Heo ae Bley ‘ i PEND a eR tan hae WyEhaS yo toed ues Mareen tas AUPE whe yn ih etn May tects Tate tah tas eT aes tty ren Bi la ot Nhat Wala vee me hese ea hes musta Rahal sy 4 rahe “A Rates hetgrn sh UN eka negteedatuga ein Sys) Riis . at Ss veo eat aw Sats ae ee PTH Re ean per UAC St ome A doings) ity fie i ra be Dia ony Mave Cala alae Pio eyeeivceee Dice yn wer EBA E ry The Pi UN wees fad Darirant ster aw { ies a Pd taira 4 4) Sane shatate Wid geen Pies eemuivunt eatin ene? Ferm yuemtl ve pean sr eres Pen en 4 Tah, elt te Ey Web vaiehepeay teeny Swen ayaa See ho Dye bat a naa ina anette ops ta Sais abies oh NAb Fearing aquretee st Witetes at ve Bae shi Movi nce ha ths it is as pale retatelt Be fe OSs mine sone as ne Ba rs uses Pee arris re CR Pan Stas Salas helce ys Youth yah A Werk aS Orta pay “iis PE a seein tea fusems Metis Pan Gnesi tients tS aghenli HCE SEEN « teh bt : ' rt pa sti Hs SVAN a Pe N ML a gaat Nise be aw tae ee paicad fateh ate Ua aisig aves Abiastts Fa HU oe NUP GSES co Soak yh Riateat Se Ee ET a he signet rottae Ratar cays ahd hes busi 8147 RaLeiont ott ‘ Penney reson ict Heautule ee) fondue roe ayant PEMA el orale. ue Spt SOLE TEN be Neha cadamnraias Taree th ora reeoe ’ y GN OVE coal asl tS gaat Tribe, ¥ ser telser kat aay rear egy Ay STR UI NeS athe SRA Che DPRESTON a i HI ieshhl vouy ase Haetay Joana sida eae Sy yee ne 1 aia PD Sa aS Uk a MA Lg Ui eaeWens | cee POPC nn aL Ein sdyitndaneee }iidoavey Mute pel he teas PERG ie Dd [Mia tyes dene dae pestyia Mek ter yeek Shasiline $e 4. gird Tarver pezie Yaar gts ¢ denna CRY fice ein’ aire on 6 Maw Ae A evr ee i Nhe cate Monte wise He ca bite tesead ore enone Penoreg sat IE Gaa8 ni TVe yy en ae Hg ine teak ac ltubere (hana bese (toy ris ise Idi surge eae! Fr ety ty aay Gatton baie iid Wiis feat err its Meret arttee se 1 Prey hil ys pon aumdt by sachet 4 sage ih mt se 2 tn Doane t cade ee fai eyes ' SIX, Wage LAE Peng oe Carp eeur etaten ae ssh faeteas i feared vit seek at Co OA DLC aa " Bh Hioaiee Pa riees 4 hy tpg Rar} athe. pales PeSCPRy TEC DE Lari Maa te we beetje Aa Peed aii UIE He 8 ba “ 47 acer WN peeve tne I ad is nt ene ff y . Myre nS a ile einie cy Penn ie tea vate pie ane vat WADIA ASH Gif th Sealy tilt rat RSS ie titel van fala ue 6 Ca J 4 Pp a Hen Ma ie ee aa wei Jah An sf) ti err dy a) Haut ae iis Watealet Bai itt myeane tae Meeanel cl i atheais m8 ii aean Ht if ai He He sti im Bewdley ‘ poate ye mate if HAM tingle AQ s " Bae ead aye BY rang san lieng ly & Jats tgosos ail ge Gar Wags he tntolrgsdann Smet aR ES san arnt ela ptPaancaety TA tau eb aang i aot gS wileteags set ittancees Hobatirt! Create ie svoune ‘ a ‘ Citi Wehbe reba ye ats ARTE Ga8e ei Ay cata: ane Wath Maer PN oavee te inte! CHRP vanes re Vnceeae ete VE aisiVstan we ¥ : ‘ ee He “atest Weta apa W aE! aOKt ALM Cah NPL neyo Has etedpe tes, Metres pianenteseres tinpagentdiona Pay oe es way even Aura ea ae uate i eres aus anna terag yeu’ ett atl gt tah. Mherdaaty i! Hare tia seb tao a svtdstt Peay NIEe Haleagouaiecigee: te bites Pract eaghies sit eet ea Doren wire : Coen HL i Pata Bitten ten spt onaed platens gana ws ata ene MM EGR A ES etm SGT 8a Shudnaredianieienr yt Meatd wash ipetares tara fisetiensttetiineate rae iiorvinaseus ies eee at (aa bie F8v a8) Db Us RA 4 ests "M Ce ee Wat) ieee ae aie esa 9 ON NMS Be Areat Tater filet rie eal o ea i ee i Api twin tdsdem saya sO an rere ait e f ie aM jas nin de Prey itailys aH ur pbirie Ce Nib oag Bea hs vin fe i Hh a 9 aia a nN a} Hi jake Tua ‘1 Neat ei itt th i i) al ah See sic ona te a in ay eke aes fiers ; eet i te i ine K 14 ,) it Vier i Bf pies ‘ chs man ii uy te i ee Ratti He pi z Sauvudiy Ol NO LIE LIBRARIES SMITHSONIAN INSTITUTION SJluyvugl INSTITUTION Ss saluvugiq Lit iSONIAN NOILNLILSNI z n z < = << z = te : z a an ~f a 3c o %2 =f = 2 YY = 5 ie kes a ISHLINS SJIMYVYUdIT LIBRARIES SMITHSONIAN eke W) > Ww = XY bee o ENN pean fC ‘ < = mis ly Zi SS oc (oe ti XY a 5 2 2 xX Zz <3 z ISONIAN INSTITUTION NOILALILSNI_NVINOSHLINS — WY _— (oe) = = (e) My, je0] — (a0) — LG 3 5 ad 5 Vy ff 7 - - J: = 2 = m = m 2 (02) = wm = OSHLINS SJIYVYUEIT LIBRARIES SMITHSONIAN = Wp, 5S Woe : é 6 LO 5 2 SS . S ? FA a G8 is eel AT 2 = a sue : ISONIAN INSTITUTION | NOILALILSNI_NVINOSHLIWS 2 uw = us Y y (dp) Vip, 4 = = = i fem < 2 < Vig 7 = ac + [oad ) es fe) = z af Ze = S3IN¥VHEIT LIBRARIES SMITHSONIAN z= im z ~ 2) wo = Me) E a) 5 | >) = > -: > — wa - P°] kK = — 2 a Ab ain ISONIAN INSTITUTION NOILALILSN! NVINOSHLIWS w = : ‘ = ' thf 6 GY = Vy = Ps Saiuvuait S S 2 : = z= = = 77) 72) OSHLIWS LIBRARIES SMITHSONIAN Ke ” 2 ” z SAN (9p) ai Tet eg (Ae NS. o =. ae Lif Lf = AE : SGiy = YY [aa em, aS te = “Gp 5 : = — 2) oe) 2 za cs HSONIAN_INSTITUTION NOILALILSNI NVINOSHLIWS e 2 eS See z Y Wy joe) a ow : — Vg jf > es P \ ARE Oe = 2 We oy Fo) eos SW NVINOSHLIW NVINOSHLIWS — WAS S Nanos = INSTITUTION NOILALILSN!I NVINOSH: z S = KE x : + - ee Z s Salyvyai yp MITHSO! AGS = z 4 ro) = (7) w xr O iH 3 = = on : INSTITUTION NOILNLILSNI NVINOSH. 2 z = w « a = a S = = ro) aj 2 S3lyuvug tt BRARI ES_ SMITHSO = (e) ge : > @ > = ; 2 - Gx m w n = INSTITUTION NOILALILSNI _ NVINOSH . = Wy 2 wie Gye 4 2. GY loo 3 >’ = 3 a LIBRARIES SMITHSOI! 2 a 2 = _ a = < + ce 3 =" re ay INSTITUTION NOILNLILSNI NVINOSH! z a °o = Ks : : & i bale i= 2 Z : S3aluYvyugi yp MITHSO! = = se (oz) ro) = > Tas = 2s wiz ©) « w 6 rE a INSTITUTION NOILALILSNI_ NVINOSH. lw 2 WR é < Be a \ oO = = re) ei Zz Saluvudid Ll BRARI Est SMITRSS a “ o w —_— 2 5 > Ee wid ret ey wm hd na &, aay | rae —} 2 JYEIT LIBRARI BS PaniTiso Man MN SmnOven NOILNLILSNI NVINOSHLINS S3JI1uVvuE = = rc z — fo) = (e) ae (2) ow — jes] = o = Ra 5. 2 5 Fa iS > — > = >" ~ a i * 2 i = = m ” m 7) m w w = n = 7,) . = UTION NOILANLILSN! NVINOSHLINS S31eYvualy SMITHSONIAN w za (22) 4 w 2 i 2 << = << = < 4 = 4 Zz | Zz BS 2 g 2 g 2 g = 2 = 2 = = a s a = | LIBRARIES SMITHSONIAN INSTITUTION NOILNLILSNI NVINOSHLINS SJIYVUE — Ww = ow = (7p) a ul 1a Ww a # = ce = oc = NS a = o Ss a = Pe ° 2 - ro) = ra) tees 2 - 2 = 2 ‘et UTION NOILALILSNI _NVINOSHLINS 3 luyvug J LIBRARI ES_SMITHSO IAN _INSTITUTI fe) ; pes > fe) eee fe) a yn — = 2 = 2 5 ¢Zy,> E > fo > rast U7 > = 2 = 2 = GY ki" o m Z m g° nm z = w = wo judg mee BRARI BS a con ANN STO TION NOUGEIESN! NVINOSHLINS (S3 IyVUE Yy, a = <7 ae a YW: Wi = | tae +4 + YY Bea NN G é ig wf, (od = S . z = = = ee . > —-, =r a x 2 2 a > NOILALILSNI_ NVINOSHLIWS LIBRARIES SMITHSONIAN INSTITUTI LIBRARIES SMITHSON NOILNLILSNI NOILNLILSNI LIBRARIES LIBRARIES NOILNLILSNI /YG11 LIBRARIES SMITHSONIAN INSTITUTION NOILNLILSNI NVINOSHLINS S3IYVUE @ = sO = o ° 20 5. a 5 rt = > Low » > fj Pe = a = as - 2 = a Zee i Z i We ‘UTION NOILOLILSN! NVINOSHLINS S3IYVYEIT LIBRARIES SMITHSONIAN w za 22) = xxv w Zz, 2 < = -= ~.W< = = 5 Zz = Zz NS = = bao (e) a DO Nee & <= fe) w ; a ny 4 AW 3) 2 2 = 2 Ee ae E 5 a = a 3 a LIBRARIES SMITHSONIAN INSTITUTION NOILALILSNI NVINOSHLINS S3AIYVUE = “ = o an ean ies uJ uJ 2 uJ = we th fy eZ z g KG = — Beer? Vp =— —_ SN NY a c SO as UTION NOILALILSNI_NVINOSHLINS S3IYVNGIT LIBRARIES SMITHSONIAN INSTITUT! z ae z cS S y = ays 2 ow = Yj, a E 2 We 5 : 6 Yy,> e = WWE > = be ll = 3a WN E 5) = LAZO R ae a “o ' ry i} j ny i , , ee na I i i \ 4 hey Ot! L ; re i 4 ‘oa 7 - LA 4 ; q A rig’ 2 Ree POE Ff + aN ae a) . { “yi ‘ ey " i : e aa) ee ¥ + " 4 Wi % my r ; f 4 ee Me 5 a ane Man gare et ] ; : or ty THE 7 J - 1 5 } F i) 4 aay 4 J es WY eat a ey ‘ " Py iy | P a ee... e Pd a | i i" ak i A, , 4 , PROCEEDINGS of the Biological Society of Washington VOLUME 107 1994 Vol. 107(1) published 20 April 1994 Vol. 107(3) published 4 October 1994 Vol. 107(2) published 21 July 1994 Vol. 107(4) published 30 December 1994 WASHINGTON PRINTED FOR THE SOCIETY EDITOR C. BRIAN ROBBINS ASSOCIATE EDITORS Classical Languages Invertebrates GEORGE C. STEYSKAL Jon L. NORENBURG FRANK D. FERRARI RAFAEL LEMAITRE Plants Vertebrates Davip B. LELLINGER THOMAS A. MUNROE Insects WAYNE N. MATHIS All correspondence should be addressed to the Biological Society of Washington, Smithsonian Institution Washington, D.C. 20560 ALLEN Press INc. LAWRENCE, KANSAS 66044 OFFICERS AND COUNCIL of the BIOLOGICAL SOCIETY OF WASHINGTON FOR 1994-1995 OFFICERS President JANET W. REID President-Elect STEPHEN D. CAIRNS Secretary CAROLE C. BALDWIN Treasurer T. CHAD WALTER COUNCIL Elected Members ROBERT J. EMRY SUSAN L. JEWETT RICHARD C. FROESCHNER LYNNE R. PARENTI ALFRED L. GARDNER F. CHRISTIAN THOMPSON TABLE OF CONTENTS Volume 107 Alvarez, Fernando and José Luis Villalobos. Two new species and one new combination of freshwater crabs from Mexico (Crustacea: Brachyura: Pseudothelphusidae) ........... Arredondo, Oscar and Storrs L. Olson. A new species of owl of the genus Bubo from the Pleistocene of Cuba (Aves: StrigifOrrmes) 2.2... ccs:ccccccccsecsesssccsseeeesstencsseeseneeeestnesneseeceeeneetennnnssesene Baldinger, Adam J. and Michael F. Gable. Two new species of Podocerus Leach (Crus- tacea: Amphipoda: Podoceridae) from Bermuda 2... scccccecccceccessssenteeseseeseeeeeeeeeeeeeeeeeeennennensseees Banta, William C. and Mae M. Crosby. Electra venturaensis, a new species (Bryozoa: Cheilostomata: Membraniporidae) from southern California 2.2... cecceeeceeeceeeeeeeeeeeee eee Bockman, Flavio A. Description of Mastiglanis asopos, a new pimelodid catfish from northern Brazil, with comments on phylogenetic relationships inside the subfamily Rhamdiinae (Siluriformes: Pimrmelodidae) cece cceceecceeccceeceneneeeeeeeeeeeeeetteettttttnnnnnnnnnnnnenneneeeeeee Browning, M. Ralph. A taxonomic review of Dendroica petechia (Yellow Warbler) (Aves? Partalimae)) x20 ie oo NR ee a a eevee nen Ss ean i no Budd, Ann F. and Hector M. Guzman. Siderastrea glynni, a new species of scleractinian coral (Cnidaria: Anthozoa) from the easterm Pacific cece ccceceeeeeeeececceceeeeeseeseeeeeeeeessseeeenseenes Campos, Emesto. Parapinnixa cubana, a new pea crab from Cuba (Crustacea: Brachy- ura: Pinmotherd ae) pact: ee ee ee A ene eer Campos, Néstor H. and Rafael Lemaitre. A new Calcinus (Decapoda: Anomura: Di- ogenidae) from the tropical western Atlantic, and a comparison with other species of the @emus: frormy thre sre ge O ete ee ee Casanova, Jean-Paul. Three new rare Heterokrohnia species (Chaetognatha) from deep Benthic samples in the northeast Atlanta 2... ccccccccccccessccccccccceeeessscecececeecessndunsececeeeceesennneeceeeeceensunneeee Couri, Marcia Souto and Carlos José Einicker Lamas. A new species of Hyperalonia Rondani, 1863 (Insecta: Diptera: Bombyliidae: ExoprosOpimae) eee eeeeeeeeeeeeeeeeeeeeeeeee Cumberlidge, Neil. Louisea, a new genus of freshwater crab (Brachyura: Potamoidea: Potamonautidae) for Globonautes macropus edeaensis Bott, 1969 from Cameroon ... Donovan, Stephen K. and Brian Jones. Pleistocene echinoids (Echinodermata) from Bermudacand Barba osx 5c 2 see ae ee eee ee Eckelbarger, Kevin J. Diversity of Metazoan ovaries and vitellogenic mechanisms: Implications} forlifeshistOnys te iy eee ee Erséus, Christer and Michael R. Milligan. Peosidriloides, a new genus, and new records of Peosidrilus (Oligochaeta: Tubificidae) from the United States, with the description of a new species from the Gulf Of MeXiCO cece ccccceeceeececcccceeceeeeeeeeeeseeetttttenennnnnnncenceneeneeceeeeecneneneneeneee Evans, Wayne A. Morphological variability in warm-temperate and subtropical pop- ulations of Macrodasys (Gastrotricha: Macrodasyida: Macrodasyidae) with the de- SCTIPLIONTOMSEVENUNEWASPECIE See eee ee Felder, Darryl L. and Raymond B. Manning. Description of the ghost shrimp Eucalliax mcilhennyi, new species, from south Florida, with reexamination of its known con- geners (Crustacea: Decapoda: Calliamassid ae) ecccccccccccsccsesssseessscesesceeeeececceceeeccecennnnstsnetseeeeeeeeee Fernandez-Leborans, Gregorio and Apolonia Novillo. Three new species of ciliate in the genera Pseudocohnilembus, Pleuronema, and Urotricha (Ciliophora) -.....:..cseeeee---- Flint, Oliver S., Jr., Steven C. Harris, and L. Botosaneanu. Studies of Neotropical caddisflies, L: The description of Cerasmatrichia, new genus, a relative of Alisotrichia, with descriptions of new and old species and the larva (Trichoptera: Hydroptilidae) Formas, J. Ramon and Lila Brieva. Advertisement calls and relationships of Chilean frogs Eupsophus contulmoensis and E. insularis (Amphibia: Anura: Leptodactylidae) Gill, Anthony C. and John E. Randall. Xenisthmus balius, a new species of fish from the Persian Gulf (Gobioidei: Xemisthmidae) .......-cssssesssecccscscececeseeeceeseeeesssssscssssessssseeeeeeeeeseeseeseeeeeeneeees Glasby, Christopher J. A new genus and species of polychaete, Bollandia antipathicola (Nereidoidea: Syllidae), from black COAL... ..cnsssesssescescsccescceccssescesscecscnssnnsnsssssssssssesseceseeeeeeeeceeceeceeeeeeeeeeees 729-737 436-444 707-720 544-547 760-777 27-51 591-599 308-311 137-150 743-750 119-121 122-131 109-113 193-217 634-640 239-255 340-353 221-238 360-382 391-397 445-450 615-621 Goodman, Steven M. Description of a new species of subfossil eagle from Madagascar: Stephanoaetus (Aves: Falconiformes) from the deposits of Ampasambazimba ............. Gruia, Magdalena, Vasilica Iavorschi, and Serban M. Sarbu. Armadillidium tabacarui (Isopoda: Oniscidea: Armadillidiidae), a new troglobitic species from a sulfurous cave TboVel RCO} VONOVENE oe eee RU Mee Oe LE ER Ce Grygier, Mark J. Summary and significance of overlooked Japanese literature on My- ZAIN OY 900 I te Ee Guerrero, Ricardo. Sciurodendrium gardneri new species (Nematoda: Trichostrongy- loidea: Heligmonellidae), a parasite of Sciurus carolinensis Gmelin, 1788 (Mammalia: Sciuridae), with comments on the biogeography of Sciurodendrium Durette-Desset, NSA ae ee ee Ws ere ae eee ANA SD Le oe A ele EE eat a Haridas, P., M. Madhupratap, and S. Ohtsuka. Pseudocyclops lakshmi, a new species (Pseudocyclopidae: Calanoida: Copepoda) from the Laccadives, India ... 2... Harvey, Michael B. A new species of montane pitviper (Serpentes: Viperidae: Bothrops) fromeCochabamibasBoliviiay = 22. ee ee ee ee ee ee Healy, Brenda. New species of Marionina (Annelida: Oligochaeta: Enchytraeidae) from Spartina salt marshes on Sapelo Island, Georgia, U.S.A. oi... ceeeceeeceeeceeeeeeeeceneeeeeeeeeeeee Hiruta, Shinichi. Two new marine interstitial Ostracoda (Crustacea: Pussellidae) from RSL eters sean fa Neer rie cect. meee pate We Sante shies Oth June Uiecewid coeds the Hsieh, Hwey-Lian. Amphitrite lobocephala, a new species (Polychaeta: Terebellidae) fROM BAT WAN gee ee ele Cen nn ei En nts Se as aN Coote sar Ishida, Teruo. A-new species of Elaphoidella (Crustacea: Harpacticoida) closely related to E. bidens (Schmeil) and the genus Attheyella from Nepal ieee ceece cece eect Jara, Carlos G. Aegla pewenchae, a new species of central Chilean freshwater decapod (GirustaceavAnomurayAeelidac) = Kensley, Brian. Redescription of Jais elongata Sivertsen & Holthuis, 1980, from the South Atlantic Ocean (Crustacea: Isopoda: Asellota) 2... cccccccceeeeceeceecsececeeeseseeneeeeeeeeeseneeeeeneneeeeee Kensley, Brian and Marilyn Schotte. A new genus and species of cirolanid isopod from the western Indian Ocean (Crustacea: PeracariGa) o..c.cccccccsscsssssssssscssccssssssssscsssesssessssensssessessessesesssessee Kensley, Brian and Marilyn Schotte. Marine isopods from the Lesser Antilles and @olombial(Grustacea wena cara cl a) eee eae eae ann Du King, Gary M., Cem Giray, and Irv Kornfield. A new hemichordate, Saccoglossus bromophenolosus (Hemichordata: Enteropneusta: Harrimaniidae), from North PANTING TN Cedi teen ee eR AAP A ET A ARN RRS OE OM IAG Aa ral AD Ane ea TE Komai, Tomoyuki, and Ichiro Takeuchi. G/yphocrangon fimbriata, a new species of caridean shrimp (Crustacea: Decapoda: Glyphocrangonidae) from Sio Guyot, Mid- PAcifice Noun tain Sipe ae Be te ete ees i 29 renee ee Sd ete byes Koretsky, Irina A. and Clayton E. Ray. Cryptophoca, new genus for Phoca maeotica (Mammalia: Pinnipedia: Phocinae) from Upper Miocene deposits in the northern BlacksSeal rep On) Sete im rk based tet et Le kaneis ote Sem coe eee ee Kropp, Roy K. and Janet Haig. Petrolisthes extremus, a new porcelain crab (Decapoda: Anomura: Porcellanidae) from the Indo-west Pacific 0... ccc eeeeceeececeeeeeeeeeceeeeeeteeeeeeeeeeene Licher, Frank. Resurrection of Glyphohesione Friedrich, 1950, with redescription of G. klatti Friedrich, 1950 and description of G. /ongocirrata (Polychaeta: Hesionidae) .... Lips, Karen R. and Jay M. Savage. A new fossorial snake of the genus Geophis (Reptilia: Serpentes: Colubridae) from the Cordillera de Talamanca of Costa Rica —... Macpherson, Enrique. Occurrence of two lithodid crabs (Crustacea: Decapoda: Lith- odidae) in the cold seep zone of the South Barbados accretionary prism .. Marshall, Harold G. Chesapeake Bay phytoplankton: I. Composition Martin, Joel W. Crabs of the family Hololodromiidae (Crustacea: Decapoda: Brachy- ura), V. Dicranodromia spinosa, a new species from the western Atlantic 0... Martin, Joel W. and Deborah L. Zmarzly. Pinnixa scamit, a new species of pinnotherid crab (Decapoda: Brachyura) from the continental slope off California 0. McCord, William P. and John B. Iverson. A new species of Ocadia (Testudines: Ba- tagurinae) from southwestern Chima ees ceeeeeeeeeeeceecceeeseeeeeeeennsseeeeeeeeesnsnnenseseeeeeenennnuneseeeeee McLaughlin, Patsy A. A new genus and two new species of deep-water hermit crabs (Decapoda: Anomura: Paguridae) from the Southerm Ocean 2...c.cccccccccecccecsesececceseeeseeeseeeeeceeeeee 421-428 699-706 174-178 179-184 151-163 60-66 164-173 657-665 517-523 256-261 325-339 274-282 283-290 482-510 383-390 458-464 17-26 312-317 600-608 410-416 465-468 573-585 451-457 354-359 52-59 469-481 v1 McLelland, Jerry A. and Gary R. Gaston. Two new species of Cirrophorus (Polychaeta: Paraonidae) from the northern Gulf Of MEex1CO eee eeeceeeceeeeeeeeenteennnnnennnnnnnnenennees Mendelson, Joseph R., III and Jonathan A. Campbell. Two new species of the Hyla sumichrasti group (Amphibia: Anura: Hylidae) from Mexico 2... ccc eceeee eee ete Miura, Tomoyuki. Two new scale-worms (Polynoidae: Polychaeta) from the Lau Back- Arc and North Fiji Basins, South Pacific Oceana eee ee cece eeeeeeeeeneeenneneeeeeeeeneneee Mooi, Richard and Antony S. Harold. Anatomical observations of the sand dollar Mellita quinquiesperforata (Leske, 1778) (Echinodermata: Echinoidea) and the des- ignation of a neotype Moon, Seung Y. and Won Kim. New species of Echiniscus (Heterotardigrada: Echin- ISCOIGEABECHINISCIAAE) HORM OTS ae ae Moon, Seung Y., Won Kim, and Roberto Bertolani. Doryphoribius koreanus, a new Species Ofmandi grad api mw Te cee eee ee nneee nee James, Samuel W. New species of Diplocardia and Argilophilus (Annelida: Oligochaeta: Megascolecidae) from southern California eee ccccceceeeeeeeeeeeeettetnttnecnnnnnnenenneee Olson, Storrs L. A giant Presbyornis (Aves: Anseriformes) and other birds from the Paleocene Aquia Formation of Maryland and Virginia Opresko, Dennis M. and Stephen D. Cairns. Description of the new genus Allopathes (Cnidaria: Antipatharia) and its type species Cirripathes AesDOnni cece Owens, Joan Murrell. Letepsammia franki, a new species of deep-sea coral (Coelen- teratatScleractiniass Micralbya Circ 2) eee teeeaaan ania nannennnnnn nen uO EEE EEE Pettibone, Marian H. Additional records of polynoid polychaetes from the Jaun de Ua RDS cee Sh a coe Rr eg RL ee eee ee Price, W. Wayne, Richard W. Heard, and Lidia Stuck. Observations on the genus Mysidopsis Sars, 1864 with the designation of a new genus, Americamysis, and the descriptions of Americamysis alleni and A. stucki (Peracarida: Mysidacea: Mysidae), fromthe Gulfiof MEXICO), 22a es ee Reed, Edward B. Arctodiaptomus novosibiricus Kiefer, 1971 in Alaska and Northwest Territories, with notes on A. arapahoensis (Dodds, 1915) and a key to New World species of Arctodiaptomus (Copepoda: Calamoida) iii eceeeecccecccecceeeeeceeeeeeeeeeeeeeeeeeneeeeeneeeeeeeeeeneee Riser, Nathan W. The morphology and generic relationships of some fissiparous het- EFONEMENCIMES 1.02 OS a ee a ee 2 a ee ee Robinson, Harold. Cololobus, Pseudopiptocarpha, and Trepadonia, three new genera from South America (Vernonieae: ASteraCeae) ence eee ceeeeeeeeeeeeeeeteen cence Rodriguez, Gilberto. A revision of the type material of some species of Hypolobocera and Ptychophallus (Crustacea: Decapoda: Pseudothelphusidae) in the National Mu- seum of Natural History, Washington, D.C., with descriptions of a new species and @UMEWSUDSPE CLES ieka tees Eek ida a ee ln td Se ee Rodriguez, Gilberto and Héctor Suarez. Fredius stenolobus, a new species of freshwater crab (Decapoda: Brachyura: Pseudothelphusidae) from the Venezuelan guiana __.... Ruedas, Luis A., John R. Demboski, and Rogelio V. Sison. Biological and ecological variation in Otopteropus cartilagonodus Kock, 1969 (Mammalia: Chiroptera: Ptero- POGidAe) MrOMMIEUZ OMAP ALT PT Se cen Schotte, Marilyn. Annina mannai, a new isopod from the Ganges River, West Bengal (Crustacea: IsopodasCirolanidae)\j2 2 ee ee eee Springer, Victor G. and Marie-Louise Bauchot. Identification of the taxa Xenocephal- idae, Xenocephalus, and X. armatus (Osteichthyes: Uranoscopidae) —...cccc-ceesseeeeee- Sternberg, Richard von. Systematic implications of color pattern polymorphism in Goniopsis pulchra (Decapoda: Brachyura: Grapsidae) from Ecuador _ Strenth, Ned E. A new species of Palaemonetes (Crustacea: Decapoda: Palaemonidae) from’ northeastern MexiGol 26 s2 ts ees aurea Bae ie at ttn ee a eas ee Suarez-Morales, E. Monstrilla elongata, a new monstrilloid copepod (Crustacea: Co- pepoda: Monstrilloida) from a reef lagoon of the Caribbean coast of Mexico ................. Suzuki, Hiroshi and Eiji Tsuda. A new freshwater crab of the genus Geothelphusa (Crustacea: Decapoda: Brachyura: Potamidae) from Kagoshima Prefecture, southern Kyushu; Japamien see cee) Ae! tas dye Bek etnies Sata Rea Maire ae eRe ene en oe Tan, Cheryl G. S. and Peter K. L. Ng. A new species of anemone-carrying crab from New Caledonia (Decapoda: Brachyura: Xanthidae: Polydectimae) eee 524-531 398-409 532-543 751-759 511-513 514-516 622-633 429-435 185-192 586-590 609-614 680-698 666-679 548-556 557-568 296-307 132-136 1-16 268-273 79-89 721-728 291-295 262-267 318-324 738-742 Thiéry, Alain and Michael Fugate. A new American fairy shrimp, Linderiella santarosae (Crustacea: Anostraca: Linderiellidae), from vernal pools of California, U.S.A. ............ Timm, Robert M. and Roger D. Price. A new species of Felicola (Phthiraptera: Tri- chodectidae) from a Costa Rican Jaguar, Panthera onca (Carnivora: Felidae) ............... Tyler, James C. and Alexandre F. Bannikov. A new genus of fossil pufferfish (Tetrao- dontidae: Tetraodontiformes) based on a new species from the Oligocene of Russia and a referred species from the Miocene of Ukraime 2... cccccceeeeeeeececeeeeeeceeeeeeeeeeeeteeeeeteeee Vari, Richard P., Antony S. Harold, and Donald C. Taphorn. Creagrutus melasma, a new species of characid fish (Teleostei: Characiformes) from upland streams of north- ETAT VE TZU) eee a RE UO Ae ere ea ue St a Wilson, Larry David, James R. McCranie, and Gustavo A. Cruz. A new species of Plectrohyla (Anura: Hylidae) from a premontane rainforest in northern Honduras .... 641-656 114-118 97-108 90-96 67-78 Vii INDEX TO NEW TAXA VOLUME 107 (New taxa are indicated in italics; new combinations designated n.c.) CILIOPHORA BEA KADU CO) 8X =p 001M) 2112 ae an eer 230 Pseudocohnilembus:fluviatiliss 30 eS ae ee ee ne SOEs ER ae ee eee 226 WUrotnch@d EEE 236 CNIDARIA Anthozoa PALL ODI CS ssa A Rh 3 ES ca NM ee Aa a 185 GES Bo rn 10s Cae Ta ee OS RE 186 Dee te po Searya rn eh sf ta ACE osetia a ee Sc eee ee 586 SIGGTaAStHC AOL 12700 esos Re Ee ae 593 RHYNCHOCOELA Nemertea WAV OISOPH GCOS sack nc MN IE Ee aoe eR ee CT 548 JEN oR) E VS oe esate tena Re te eer 553 PSEUCO]ACESUS TCs ae a eae ae ae Ln ee ee 552 SEMEN UND SUS TN ese eee ee cts RO eT 549 GASTROTRICHA Macrodasys:G@chradocytalis'. ) was calculated by hand using the method of Sokal and Rohlf (1981:174—-175). Species diversity was measured using the Shannon diversity index, H’ (Shannon 1948). Differences between diversity indi- ces of the two sites were evaluated using the t test approach of Hutcheson (1970) for the Shannon formula, as described by Zar (1984). Community similarity was exam- ined using the Horn index of community overlap (Horn 1966, Brower & Zar 1984). VOLUME 107, NUMBER 1 Evenness of abundance of species was cal- culated as the ratio of the Shannon index, H', to the maximum possible diversity, 1 ax (Pielou 1969). These measures of di- versity and community similarity (hetero- geneity indices sensu Peet 1974) are pre- ferred herein over species abundance models because’ they are distribution independent (Magurran 1988, Peet 1974; but see Gra- ham 1983 for a contrasting opinion). Specimens Examined Otopteropus cartilagonodus. —Philip- pines: Luzon Island; Camarines Sur Prov- ince; 4 km N,-:18 km E Naga, Mt. Isarog, 13°40'N, 123°20’E, 475 m; external and cra- nial characters of 4 males (U.S. National Museum of Natural History [USNM] 573439, 573444-573446) and 6 females (USNM 573440-573443, 573447, 573448); external characters of 1 male (USNM 573684) and 2 females (USNM 573680, 573682). Philippines, Luzon Island; Ca- marines Sur Province; 4 km N, 21 km E Naga, Mt. Isarog, 13°40’N, 123°22’E, 1350 m; external and cranial characters of 1 fe- male (USNM 570503), external characters of 1 female (USNM 573713) and 2 males (USNM 573712, 573715). Philippines: Lu- zon Island; Camarines Sur Province; 5 km N, 20 km E Naga, Mt. Isarog, 13°40'N, 123°21'E, 900 m; external characters of 3 males (USNM 573694, 573696, 573704) and 20 females (USNM 573685-573693, 573695, 573697-573703, 573705—-573707). Philippines: Luzon Island; Camarines Sur Province; 4.5 km N, 20.5 km E Naga, Mt. Isarog, 13°40’'N, 123°22’E, 1125 m; external characters of 4 females (USNM 573708- 573711). Philippines: Luzon Island; Ca- marines Sur Province; 4 km N, 21.5 km E Naga, Mt. Isarog, 13°40’N, 123°22’E, 1550 m; external characters of 1 male (USNM 573715). Philippines: Luzon Island; Ca- marines Sur Province; 4 km N, 22 km E Naga, Mt. Isarog, 13°40’N, 123°22’E, 1750 m; external characters of 1 male (USNM 573716). Philippines: Luzon Island; Isabela Province; 3 km W mouth of Blos River, 17°30'N, 122°10’E, elev. 50 m (precise lo- cality illustrated in Fig. 1, Mudar & Allen 1986:220; external and cranial characters of 1 female (University of Michigan Museum of Zoology [UMMZ] 156972). Philippines: Luzon Island; Zambales Province; Zam- bales Mountains, 15°35’N, 120°09’E, 1140 m; external and cranial characters of 7 fe- males (National Museum of the Philip- pines/Cincinnati Museum of Natural His- tory field numbers [NMP/CMNH] 22, 39, 75, 139, 140, 148, 149); external characters of 3 females (NMP/CMNH 45, 106, 141). Philippines: Luzon Island; Zambales Prov- ince; Zambales Mountains, 15°30’'N, 120°08’E, 1500 m; external and cranial characters of 3 females (NMP/CMNH 354, 388, 407) and 9 males (NMP/CMNH 304, 312, 356, 362, 364-366, 390, 391); external characters of 4 females (NMP/CMNH 321, 332, 389, 408) and 12 males (NMP/CMNH 302, 307, 314, 318, 322, 335, 336, 355, 357, 363, 378, 409). These specimens will be as- signed permanent museum numbers as soon as the division of specimens between CMNH and NMP takes place. Specimens from the National Museum of the Philippines were examined by RVS, but the measurements are not included to main- tain consistency of measurer and instru- ment. Specimens are identified by collection date because of lack of museum number, as follows: Luzon Isl.; Laguna Prov.; Balian; 1 female (col. 16 Jul 1964). Luzon Isl.; Nueva Viscaya Prov.; Dalton; 1 male (col. 20 Apr 1966), 4 females (col. 20 and 21 Apr 1966; 3 Jun 1970; 29 Oct 1970). Luzon Isl.; Que- zon Prov.; Real, National Botanic Garden (University of the Philippines Land Grant); 2 males (3 and 8 Jun 1974). Three speci- mens (two males, one female) were collected at this last site by Andres L. Dans and Pedro L. Alviola, III, in May 1983, but were lost during a fire at the University of the Phil- ippines at Los Banos on 10 May 1990 (A. L. Dans, in litt.). One additional male was 4 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON collected during May 1992 by two students of A. L. Dans in Luzon Isl.; Cagayan Prov.; Mount Cetaceo, Sierra Madre, elev. 1500 m (A. L. Dans, in litt.) The exact where- abouts of this specimen are unknown. The type series was not examined, but includes the holotype (Senckenberg Muse- um, Frankfurt [SMF] No. 28462, male) and three paratypes (SMF 28852-28854, 2 males, 1 female) from Philippines: Luzon Island; Mountain Province; Sitio Pactil; paratype, SMF 35750, female, Luzon Is- land: Abra Province; Massisiat Resthouse; and paratype, SMF 35749, female, “Phil- ippines,”’ no specific locality. No elevations, latitude, or longitude are noted in the de- scription. Study Sites Specimens from Mount Isarog were col- lected by L. R. Heaney and his coworkers. These sites are described and illustrated by Goodman & Gonzales (1990) and Rickart et al. (1991); one O. cartilagonodus from that locality is illustrated in Heaney & Rick- art (1990). The collection locality of the sin- gle specimen from Isabela Province (UMMZ 156972), also collected by L. R. Heaney, is described by Mudar & Allen (1986). The Zambales Mountains specimens were col- lected by LAR and JRD and coworkers. The Zambales Mountains (Fig. 1) are an isolated mountain range encompassing ap- proximately 6960 km? in west central Lu- zon. Including the volcanoes of the Bataan Peninsula, the Zambales extend approxi- mately 200 km in length, running about 20° west of due north; the southern end (Bataan Peninsula) begins approximately 55 km west of the city of Manila, across Manila Bay. The greatest width of the range is about 60 km. To the east, the Zambales Mountains are isolated from the Cordillera Central (the nearest mountain range) by the extensive plains of Tarlac, also known as the Great Valley of Luzon, a wide expanse of alluvial, fluviatile, lacustrine and other sedimentary deposits (including beach and coralline), now consisting primarily of rice fields. To the west and north, the mountains end in the South China Sea; to the south, they separate Subic Bay from Manila Bay. Rainfall patterns in the area display clear- ly demarcated dry and wet seasons. Data collected between 1951 and 1970 in Iba, a coastal town in the Zambales Province near the study area, indicate rainfall maxima in late July or August (ca. 1025 mm monthly average) and minima in late January or Feb- ruary (ca. 20 mm monthly average; data from Philippine Council, 1977). Site 1.—Zambales Mountains, 15°35’'N, 120°09'E, 1140 m. Sampled 17 to 26 Feb 1992. The altitudinal transect at this site was Situated along a steep South facing ridge of Mount Apoy between 1050 m and 1265 m. During our stay at this camp, nighttime low temperatures averaged 10°C, while day- time highs rarely exceeded 23°C. This site was characteristic of the tropical moist de- ciduous forest type of Whitmore (1984), specifically, tropical lower montane rain forest; toward the top end of the transect were found elements transitional to a mossy forest type. Relatively untouched forest be- gins in the area only above approximately 1030 m (LAR, pers. obs.); below this ele- vation, extensive thickets of bamboo and other secondary growth predominate. The forest had two stories, but moving from the lower to higher elevations, the canopy grad- ually decreased in height above ground from 15-16 m to 11-13 m, and the subcanopy became increasingly broken and less con- spicuous. Due to the high number of tree falls and boulders (the latter often 4-5 m wide) at either end of the transect, the can- opy at the extremes of the transect was bro- ken and uneven, while the canopy between the extremes was closed and continuous. Vegetation varied depending on exactly where the nets were set. Nets set along a ridge near the camp (ca. 1170 m) were in a moderately to dry habitat, with volcanic ash from the explosion of Mount Pinatubo in VOLUME 107, NUMBER 1 5 120° E LINGAYEN GULF 16° N— SOUTH CHINA SEA 15N Fig. 1. Map of the geological assemblage constituting the Zambales Mountains (stippled area); the solid circle marks the study area. Inset map shows the Zambales area (box) in relation to the remainder of Luzon Island and the Philippines, with the arrow pointing to Mount Isarog, where the sample of Otopteropus cartilagonodus from the National Museum of Natural History originated. 6 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 1991 covering much of the ground and veg- etation (1—2 cm thick layer). The driest por- tion of the site was covered almost entirely by grasses (Poaceae); several charred up- right tree trunks showed the area once had other vegetation, but had since been burned to grassland. Toward the east, the area was less dry, and trees of the families Myrtaceae, Lauraceae, and Tiliaceae predominated; these had a maximum height of approxi- mately 9 m, with a diameter at breast height (dbh) no greater than SO cm, and usually less than 30 cm. Proceeding downhill from the camp (S) trees increased slightly in stat- ure. There were some, though not many, epiphytic orchids, particularly above 1250 m; some climbing bamboos, and shrubby, herbaceous undergrowth in the wooded ar- eas northeast of the camp. Below 1100 m there was abundant rattan, a climbing spiny palm (Calamus) used in manufacturing fur- niture, and the gathering of which is the livelihood for many people in the area; the area cannot be said, therefore, to be wholly undisturbed. Sampling effort was 240 net nights. Site 2.—Zambales Mountains, 15°30'N, 120°08’'E, 1500 m. Sampled 11 to 18 March 1992. Nighttime lows at this site averaged 12.6°C (range: 10.3°-14.0°), while daytime highs averaged 22.5°C (range: 19.8°-24.0°). As with Site 1, this site is in the broad cat- egory of tropical moist deciduous forest type of Whitmore (1984); however, at this ele- vation, the vegetation was more character- istic of tropical upper montane rain forest. This site was very dry, especially as it was visited near the height of the dry season in the Zambales Mountains. Ashfall from the 1991 Mount Pinatubo explosion covered the ground to a depth of 2—4 cm and coated much of the vegetation. Canopy height var- ied from less than 5 m to approximately 11 m; no trees exceeded 70 cm dbh: in fact, few were greater than 50 cm dbh. Predom- inant trees were of the families Fagaceae, Myrtaceae, Magnoliaceae, and Pinaceae, al- though other families also were represented to a lesser extent. Many epiphytic orchids were present at this site, as well as epiphytic ferns. Not much bamboo or other secondary vegetation was found; in addition, there were many dead branches, and a thick cover of leaf litter on the ground. This site was sam- pled for 200 net nights. Results Morphological variation. —Results of morphometric analyses of the Zambales Mountains population are summarized in Table | for cranial mensural characters that did not differ significantly between the two sexes. Those cranial characters that did dif- fer are summarized in Table 2. Using a probability level of a = 0.05, the expected number of sexually dimorphic characters would have been between one and two for 24 characters examined; five were found. All of these are directly or indirectly in- volved in the feeding apparatus. Length of rostrum, length of M3, and length of man- dible from condyle to canine, all affect the mechanical force applied during mastica- tion. Posterior pterygoid breadth influences the maximum size of food particles able to be ingested. An additional note with respect to sexual dimorphism in this population is that all cranial characters are larger in male O. cartilagonodus; the remaining sexually dimorphic character, height of braincase, is larger in females of the species. This last character may offset partially the size ad- vantage of males over females in mastica- tion related measurements by influencing the mechanics of the masseter muscle. Wing bone measurements are summa- rized in Table 3; no sexual dimorphism was found in any of these mensural characters. External measurements are summarized in Table 4. Of these, only total length dis- played significant sexual dimorphism, with females being somewhat longer than males. Mensural characters of the Mount Isarog population are summarized in Tables 5-7. Only two cranial measurements differed be- — VOLUME 107, NUMBER 1 Table 1.—Standard univariate statistics for cranial characters (in millimeters; abbreviations defined in text) of Otopteropus cartilagonodus from the Zambales Mountains. Characters displaying sexual dimorphism are summarized in Table 2. (Other abbreviations are: Y mean; SE, standard error; W, results of the Shapiro-Wilk test for normality [N, normal; *, 0.05 > P > 0.01; **, 0.01 > P > 0.001; ***, P < 0.001); g,, skewness statistic [n.s., not significant]; g,, kurtosis statistic.) Character x SE Range Ww 2, ty g t.> GLS 23.02 + 0.107 22.2-23.8 N 0.257 n.s —0.652 n.s PALL 11.35 + 0.072 10.9-11.9 N —0.046 n.s —1.308 n.s MAXTL 6.56 + 0.042 6.2-6.8 N —0.493 N.S —0.672 N.S UPMOLL 5.33 + 0.039 5.0-5.6 N —0.655 N.S —0.440 N.S ROSW 5.52 + 0.956 5.0-6.1 N 0.323 n.s —0.716 n.s IOB 4.20 + 0.051 3.7-4.6 N —0.224 n.s —0.100 n.s ZB 14.49 + 0.079 13.9-15.3 N 0.692 N.S 0.504 N.S BB 10.65 + 0.058 10.2-11.2 N 0.435 n.s —0.040 N.S. INTCB 2.09 + 0.034 1.8-2.4 N 0.123 n.s —0.633 N.S. INTMB 4.84 + 0.054 4.4-5.2 N —0.253 N.S —0.930 n.s APTB 3.90 + 0.036 3.7-4.2 N 0.157 N.S. — 1.030 n.s HUPCAN 1.85 + 0.071 1.2-2.3 N —0.335 n.s —0.370 n.s M3W 0.90 + 0.018 0.7-1.0 * —1.472 By 4.325 ee DENTL 16.43 + 0.088 15.7-17.6 EP 1.385 1 3.855 EE CONM1 13.26 + 0.093 12.7-14.1 N 0.406 n.s —0.722 n.s MNDTOOTH 6.64 + 0.043 6.3-6.9 N —0.523 n.s —0.991 n.s MAMASTR 7.18 + 0.079 6.5-7.8 N —0.545 n.s —0.075 n.s DENTTHK 1.93 + 0.029 1.62.2 N —0.128 n.s 0.797 N.S HTLWRCAN 1.66 + 0.046 1.2-2.1 = —0.001 n.s 3.034 =e tween males and females in this population: zygomatic breadth (P = 0.0085) and pos- terior pterygoid breadth (P = 0.0090). With respect to the wing bones, the only one found to differ between the sexes was the first pha- lanx of digit 5 (P = 0.0053). Ear length was the only external character found to differ between the sexes (P = 0.0117), despite con- siderable overlap in the ranges. Mensural characters of the single speci- men from Isabela Province (UMMZ 156972) are reported in Table 8. Because this sample consists of a single specimen, statistical comparisons between this and the other two populations were not carried out. Geographic variation. —Considerable dif- ferences in morphology exist between the two populations of O. cartilagonodus ex- amined herein: 21 out of the 37 characters examined differed significantly between Zambales and Isarog populations. With re- spect to non-sexually dimorphic characters of the skull, significantly different characters include interorbital breadth (P = 0.0000), Table 2.—Standard univariate statistics for sexually dimorphic mensural cranial characters of Otopteropus cartilagonodus from the Zambales Mountains. (Character abbreviations as in text; other abbreviations as in Table 1.) Males Females Character x SE Range X SE Range ROSL 10.67 + 0.116 10.0-11.1 10.12 + 0.118 9.7-10.7 HB 8.93 + 0.098 8.89.8 9.34 + 0.102 8.89.8 PPTB 3.48 + 0.048 3.3-3.7 3.27 + 0.058 3.0-3.6 M3L 1.12 + 0.022 1.1-1.2 1.04 + 0.020 0.9-1.1 CONCANL 15.82 + 0.105 15.4-16.3 15.50 + 0.096 15.1-16.0 8 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 3.— Wing bone measurements, in millimeters, for Otopteropus cartilagonodus from the Zambales Moun- tains. (Character abbreviations as in text; other abbreviations as in Table 1.) Character xX SE Range D2M 22.41 + 0.138 20.1—24.1 D2P1 5.23 + 0.055 4.6-5.8 D3M 32.88 + 0.173 30.0-35.2 D3P1 24.78 + 0.161 22.1-26.9 D4M 30.58 + 0.183 27.2-33.5 D4P1 17.82 + 0.145 15.0-19.9 D5M 32.25 + 0.181 28.3-34.0 DSP1 15.44 + 0.106 14.0-16.4 height of upper canine (P = 0.0000), length of dentary (P = 0.0052), length of condyle to M1 (P = 0.0001), length of mandibular toothrow (P = 0.0000), and height of lower canine (P = 0.0047). In sexually dimorphic characters, females differ between the two populations in length of rostrum (P = 0.0005), zygomatic breadth (P = 0.0003), posterior pterygoid breadth (P = 0.0370), and condylocanine length (P = 0.0201), while males differ in posterior pterygoid breadth only (P = 0.0397). Every wing bone measurement differs significantly between the two populations (FA, D2M, D3M, D4M, D4P1, DSM, P= 0.0000; D3P1, P= 0.0093; DSP1, females, P = 0.0000, males, P = 0.0005). Finally, with respect to external characters, the two populations differ in length of hind foot (P = 0.0001), mass (P = 0.0001), and total length (females, P = 0.0001, males, P = 0.0091). The specimen from Isabela Province (Table 8) is closest w 2, t 2 t, N —0.433 fs 0.864 n.s fe —0.012 n.s —1.274 n.s N —0.467 n.s 0.500 n.s N —0.361 n.s 0.919 n.s N —0.076 n.s 2.046 pa N —0.453 n.s 1.985 ba a —1.103 be 3.769 oe N —0.576 N.S. —0.388 n.s to Mount Isarog specimens rather than to Zambales specimens in all characters ex- amined, albeit without statistical confir- mation. Ecological notes. —Thirty-eight speci- mens of O. cartilagonodus were collected in mist nets set near Sites 1 and 2. Ten indi- viduals, all females, were collected at Site 1 in seven nets set between 1130 and 1235 m. Four were caught in two nets, set at 1140 m and 1180 m; the remaining six were caught singly. All individuals were flying between 125 and 270 cm above the ground when captured. Of these ten females, nine were pregnant, with embryos averaging 9.22 mm in crown rump length (mode, 12 mm; range 5-13 mm; SE, + 0.969). At Site 2, 15 males and 9 females were collected. Two nets, set at 1579 m, and 1613 m, each caught two; one net set at 1594 m resulted in three; remaining bats were caught singly in nets at elevations between 1365 m Table 4.—Standard external measurements for Otopteropus cartilagonodus from the Zambales Mountains. Total length was the only external measurement found to be sexually dimorphic. (Mass expressed in grams; all other measurements in millimeters; abbreviations as in Table 1.) Character XG SE Range Ww £, tay g ts Hind foot 10.66 + 0.102 9-12 AS —0.294 n.s 0.189 n.s Ear length 13.80 + 0.150 11-16 A —0.646 n.s 1.887 * Forearm 44.55 + 0.213 41-47 N —0.835 ne 0.864 n.s Mass 13.55 + 0.099 12-15 = —0.025 N.S. 0.192 N.S. Total length Females 69.47 + 0.515 65-74 N 0.088 n.s —0.344 N.S. Males 68.10 + 0.350 66-70 - —0.092 N.s. —1.422 N.s. VOLUME 107, NUMBER 1 9 Table 5.—Standard univariate statistics for mensural cranial characters of Otopteropus cartilagonodus from Mt. Isarog (specimens at the U.S. National Museum of Natural History, U.S.N.M.), in millimeters. (Abbrevi- ations are as in Table 1.) Character X SE Range Ww £ te 2 fa GLS 23.18 + 0.123 22.4-23.9 N —0.374 N.S 0.723 N.S ROSL 10.79 + 0.078 10.4-11.1 N —0.454 n.s —1.625 n.s PALL 11.89 + 0.077 11.4-12.3 N —0.364 N.S —0.134 n.s MAXTL 6.76 + 0.112 6.2-7.5 N 0.518 n.s 0.976 n.s UPMOLL 5.45 + 0.068 5.0-5.7 N —1.271 n.s 0.921 n.s ROSW 5.72 + 0.140 5.1-6.5 N 0.232 n.s —0.878 n.s IOB 3.73 + 0.071 3.44.0 N 0.006 n.s —1.606 n.s BB 10.85 + 0.094 10.2-11.2 N —1.144 N.S. 0.897 n.s HB 9.52 + 0.091 9.0-10.2 N 0.844 n.s 1.485 n.s INTCB 2.20 + 0.045 1.9-2.4 N —0.802 N.S. 0.361 N.S INTMB 4.77 + 0.064 4.4-5.1 N —0.081 n.s —0.779 n.s APTB 3.81 + 0.027 3.7-4.0 N 0.944 N.S 2.110 N.S. HUPCAN 2.53 + 0.077 2.0-2.8 N —0.759 N.S. 0.466 N.S. M3L 1.09 + 0.016 1.0-1.1 N —0.503 N.S —1.168 N.S M3W 0.91 + 0.021 0.8-1.0 N 0.675 n.s 0.309 N.S. DENTL 16.89 + 0.124 16.2-17.7 N 0.172 N.S 0.706 n.s CONCANL 16.10 + 0.141 15.0-16.6 N —1.128 N.S 1.947 n.s CONM1 13.92 + 0.114 13.1-14.5 N —0.828 N.S. 1.247 n.s MNDTOOTH 7.03 + 0.068 6.7-7.5 N 0.421 N.S 0.768 n.s MAMASTR 7.23 + 0.079 6.6-7.6 N —1.112 N.S 1.552 N.S DENTTHK 1.91 + 0.045 1.72.2 N 0.300 N.S —0.372 N.S. HTLWRCAN 1.90 + 0.053 1.6—2.3 N 0.612 N.S. 0.516 N.S ZB (females) 15.10 + 0.086 14.8-15.5 N 0.066 n.s 0.385 N.S ZB (males) 14.51 + 0.180 14.2-15.0 N 1.635 N.S 2.999 N.S. PPTB (females) 3.09 + 0.043 2.9-3.3 N 0.360 n.s —0.701 n.s 3.30 + 0.035 3.2-3.4 N —1.414 N.S 1.500 N.S PPTB (males) Table 6.—Wing bone measurements, in millimeters, for the Otopteropus cartilagonodus from Mt. Isarog (specimens at the U.S.N.M.); 33 females and 8 males comprise the sample. The only measurement differing at a = 0.05 was that of the first phalanx of digit 5 (D5P1; P = 0.0053); however, D3M, D3P1, and D4P1 all had P values between 0.05 and 0.10. (Character abbreviations as in text; other abbreviations as in Table 1.) Character XG SE Range Ww 21 iby g. t D2M (8) 23.94 + 0.183 21.4-25.8 N —0.038 N.S. —0.170 N.S D2M (6) 23.44 + 0.529 21.1-25.5 N —0.346 N.S. 0.382 N.S D3M (8) 34.56 + 0.191 32.3-36.7 N —0.123 n.s —0.739 n.s D3M (8) 33.78 + 0.333 32.6-35.1 N 0.016 N.S. —1.702 N.S D3P1 (9) 25.45 + 0.143 23.8-27.2 N —0.045 N.S. —0.673 N.S. D3P1 (6) 24.82 + 0.277 23.5-25.8 N —0.688 n.s —0.412 n.s D4M (8) 32.98 + 0.181 30.0-—35.2 N —0.402 N.S. 1.059 N.S. D4M (6) 32.56 + 0.469 30.8-34.7 N 0.351 N.S. —0.673 n.s D4P1 (9) 19.28 + 0.188 17.7-23.9 ba! 2.509 +4 10.050 +45 D4P1 () 18.55 + 0.245 17.5-19.4 N —0.279 N.S. —1.210 n.s DSM (8) 33.48 + 0.214 31.0-35.3 N —0.206 N.S. —0.803 N.S. DSM (6) 33.09 + 0.389 31.6-35.0 N 0.502 N.S. —0.072 n.s DS5P1 (8) 17.12 + 0.120 15.8-18.8 N 0.149 n.s 0.498 N.S. DS5P1 (6) 16.36 + 0.177 15.6-17.2 N 0.281 N.S 0.593 n.s 10 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 7.—Standard external measurements for Otopteropus cartilagonodus from Mt. Isarog (specimens at the U.S.N.M.); 40 females and 14 males comprise the sample. Ear length was the only external measurement found to be sexually dimorphic (P = 0.0117). (Mass expressed in grams; all other measurements in millimeters; abbreviations as in Table 1.) Character Xx SE Range Ww g t, g ts Total length Females 74.67 + 0.989 65-87 N 0.196 N.S. —0.861 n.s. Males 72.28 + 1.348 65-79 N —0.175 n.s. —1.422 N.S. Hind foot Females 13.78 + 0.494 11-20 CEES 1.058 ES —0.658 N.s. Males 12.88 + 0.670 11-19 cE 1.931 ae 2.892 z Ear length Females 14.00 + 0.095 12-15 a —0.753 z 2.789 BAAS Males 13.21 + 0.260 12-15 N 0.089 N.S. — 1.026 N.S. Forearm Females 48.75 + 0.256 44-52 N —0.190 N.S. 0.650 N.s. Males 47.86 + 0.404 45-51 N 0.124 N.S. 0.494 N.S. Mass Females 15.98 + 0.245 13-19 N 0.109 N.S. —0.706 N.S. Males 16.36 + 0.360 15-20 te 1.615 aS 3.369 ma and 1643 m. At this site bats were flying 14mm, the third female was fluid preserved between 30 and 210 cm above the ground intact and determined to be pregnant by when caught. Only three of the nine females external inspection (fide JRD). All males were pregnant; two embryos measured 8 and captured exhibited abdominal testes; habit- Table 8.— Measurements of the single specimen of Otopteropus cartilagonodus (UMMZ 156972) from Isabela Province, Luzon Island, Philippines. (Mass expressed in grams, all other measurements in millimeters.) Cranial measurements Greatest length of skull 23.3 Anterior pterygoid breadth 3.7 Length of rostrum 10.7 Posterior pterygoid breadth 3.0 Length of palate 12.2 Height of upper canine 2.6 Length of maxillary toothrow U2 Length of M3 1.0 Length of upper molariform row 6.1 Width of M3 0.9 Rostral breadth 4.8 Length of dentary 7 Interorbital breadth 3.5 Condylocanine length 15.7 Zygomatic breadth 15.4 Condyle to m1 14.3 Breadth of braincase 10.9 Length of mandibular toothrow 6.7 Height of braincase 9.4 Moment arm of masseter 7.8 Breadth between upper canines DP Thickness of dentary 2.0 Breadth between M3 4.8 Height of lower canine 2.0 Wing bone measurements D3M 32.6 D4M 31.3 DSM 32.3 D3P1 26.0 D4P1 18.7 D5P1 16.3 FA 47.6 External measurements (from specimen label) Total length 70 Length of hind foot 11 Length of ear 14 Mass 17 VOLUME 107, NUMBER 1 ually, this would indicate the organism is not breeding, however, many bats are able to move their testes in and out of their ab- dominal cavity, apparently at will. For 14 males where such data were taken, testes measurements averaged 3.3 by 2.4 mm. A single net set for one night just below Mt. High Peak (15°29’N, 120°07’E) at ca. 1900 m caught four male O. cartilagonodus with abdominal testes, averaging 3.2 by 2.5 mm. The nighttime low at this camp reached at least 7°C. Geographic variation. —The altitudinal segregation of sexes found in the Zambales sample was not present in the sample from Mount Isarog, as no statistically significant differences were found between capture el- evations in males versus females. The trend, however, was for a greater number of fe- males than males at the lower elevations: three males and twenty females were cap- tured at 900 m, only females at 1125 m, one female and two males were captured at 1350 m, but only males of the species were cap- tured at 1550 and 1750 m (one at each el- evation). The two populations also appear to differ in reproductive timing. Heideman et al. (1993) found that the Mount Isarog sample displayed synchrony of embryolog- ical development, with small uterine swell- ings in March, and 15-20 mm embryos in females captured in May; they extrapolated birth dates in late May or June. In contrast, the sample from the Zambales Mountains (collected mid-February to early March) contains a mixture of pregnant and non- pregnant females with embryos from 5 to 14 mm in crown rump length. Other chiropteran species. — An addition- al 12 species of bats were collected in the same two localities where O. cartilagonodus was found to be present (Table 9). The Shannon diversity index for the chiropteran fauna of Site 1 was H', = 0.648, while that for Site 2 was H', = 0.585. The ¢ test for the Shannon formula (Hutcheson 1970), in- dicated there was no significant difference between species diversities at the two sites 11 Table 9.—List of bats caught at the two localities where Otopteropus cartilagonodus also were captured'. Site 1, Site 2, Site: Zambales, Zambales, 1100m 1500m Species (n) (n) Rhinolophidae Rhinolophus arcuatus 59 39 Rhinolophus subrufus 36 2 Rhinolophus philippinensis 1 3 Hipposideros bicolor 0 1 Vespertilionidae Myotis muricola 2 20 Murina cyclotis 1 1 Pteropodidae Cynopterus brachyotis 1 0 Eonycteris spelaea 1 0 Haplonycteris fischeri 1 0 Macroglossus minimus 1 0 Otopteropus cartilagonodus 10 24 Ptenochirus jagori 5 0 Rousettus amplexicaudatus 19 0) ' Four O. cartilagonodus and two R. arcuatus that were caught in a net set at 1900 m are not included in the totals. (P > 0.05). Because diversity indices usu- ally are not calculated for particular sub- sections of a fauna (Chiroptera in this in- stance), we calculated the relative diversity, or evenness, J’, as a proportion of H’, to the theoretical maximum value for H’, or FD pmax, thereby yielding a value constrained between 0 and 1. For Site 1, H’, ax = 1.079, while for Site 2, H’> ,,., = 0.845. Thus, for Site 1, J’; = 0.600, while for Site 2, J’, = 0.692. The Horn index of community sim- larity (“community overlap’), Ro, also constrained between 0.0 (when the two communities under consideration have no species in common) and 1.0 (when species compositions and relative abundances are identical between the two sites) calculated for Sites 1 and 2 was Ry = 0.637; the overlap between the chiropteran faunas of the two sites was thus on the order of 64%. What may not readily be apparent from these indices is that there is at least one major difference between the two sites, that being in the abundances of three species: 12 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Myotis muricola, Rhinolophus subrufus, and Rousettus amplexicaudatus. This last spe- cies is particularly noteworthy because it previously has been associated with dis- turbed areas, either natural or anthropogen- ic (Heideman & Heaney 1989). Its presence at Site 1 reinforces our perception of this site as disturbed, either by rattan gathering, or by the Mount Pinatubo ashfall, or a com- bination of both factors. The few known specimens of R. subrufus have been col- lected from caves (Heaney et al. 1987); their presence at Site 1 but not Site 2 may be more indicative of the proximity of a cave, rather than of any specific habitat prefer- ence. Discussion With respect to morphological variation, we found that 21 out of the 37 mensural characters examined in the Zambales and Isarog populations differed significantly be- tween the two populations. They are most readily separated by forearm length (Isarog, 45-52 mm; Zambales, 41-46 mm), and length of upper and lower canines. The dif- ferences in dental characters may be indic- ative of differences in diet. There also are marked differences in reproductive timing and embryonic development between the Zambales and Isarog samples. Although the two populations constitute readily identifi- able clusters of individual organisms (sensu Cracraft 1983, McKitrick & Zink 1988), we prefer at this time not to make any hard and fast taxonomic decision with respect to Otopteropus, since this might obscure the fact that we still know very little about the biology of these bats, which still are known from very small samples. We were unable to examine the type series, although from measurements provided in Kock (1969), the holotype and adult paratypes most closely resemble the specimens from Mount Isarog than the geographically closer Zambales specimens. The specimen from Isabela Province also is most similar to the Isarog and type series. The specimens from the Cordillera Central and Sierra Madre Moun- tains are closer in multivariate morpholog- ical space to the Mount Isarog population (based on the available material) than they are to the Zambales population. In our Zambales Mountain study popu- lation of O. cartilagonodus we found a great- er number of sexually dimorphic mensural characters in the skull than expected by chance alone. All differences found were in characters that affect the feeding apparatus, and thereby potentially trophic behavior as well; these may also impact male compe- tition in this probably harem-polygynous species. Although there is sexual dimor- phism in characters associated with feeding and mastication, there is no difference in mass between the sexes, and only a slight (but significant) difference in total length. Four of the five dimorphic characters of the skull were larger in males than females; however, males are significantly smaller than females in overall length. Because there is no difference in mass between the sexes, and only the difference in overall length is very small, we hypothesize that the intersexual differences in masticatory characters affect (or are affected by) food preferences, rather than resulting from an overall size com- ponent. Latitudinal species succession, or succes- sion of species with temperature variation is a long known phenomenon (Fleming 1973, McCoy & Connor 1980, Rapoport 1982, Pianka 1983). Recently, however, more attention has been paid to an expected corollary of this phenomenon, that is, to elevational zonation of species. The under- lying processes controlling this phenome- non remain unclear because of the extensive variation in geographic, taxonomic, and trophic parameters (Terborg 1971, Heaney et al. 1989, Patterson et al. 1989, Rickart et al. 1991). One reason for the uncertainty underlying the patterns and processes of al- titudinal distribution may be that most re- search has focused on faunal subsets, such VOLUME 107, NUMBER 1 as mammals (Heaney et al. 1989, Heaney & Rickart 1990), small mammals (Patter- son et al. 1989, Rickart et al. 1991), bats (Graham 1983), or birds (Goodman & Gon- zales 1990; Terborgh 1971, 1977), rather than on individual species. It is possible that the different faunas, and indeed, different species within a faunal assemblage, may be prescribed by a specific paradigm governing different factors, or characteristics, of their elevational distribution and, consequently, altitudinal patterns of replacement. Fur- thermore, in habitats considered less “‘het- erogeneous”’ (where there may be fewer re- sources available to species), there may be mechanisms for. reducing intraspecific com- petition (or intersexual competition); this sort of intraspecific zonation may be a con- founding variable when it comes to exam- ining the elevational distribution of taxa and faunas. We found such an apparent segregation of sexes by habitat along an elevational gra- dient. At lower elevations (ca. 1100 m), only females, most (90%) of them pregnant, were found. At elevations surrounding the 1500 m site, a ratio of 3 females to 8 males existed and only 30% of the females were pregnant. Within the confines of a limited sampling effort, only males were found at 1900 m. A major supposition here is that capture site is correlated with day roost site, which not always is the case in bats mist netted in the mountains. However, since the nets were set in recognizably discrete habitat types at each elevation (lower and upper montane forest), the hypothesis that intraspecific sex- ual segragation takes place may confidently be stated. It has been remarked that in contrast to rodents, bats decrease in relative abundance and species richness with elevation (Gra- ham 1983, Heaney et al. 1989, Heaney & Rickart 1990). Possibly, the metabolic de- mands imposed by flight on bats (Burton et al. 1989, McNab 1989), in contrast to other mammals, together with reduced availabil- ity of trophic resources for insectivorous and i133 frugivorous organisms at higher elevations, may increase the competition for resources at these elevations. Traditional definitions of competition di- vided this population phenomenon into ex- ploitative (indirect, i.e., resource depletion) and interference competition (direct, i.e., fighting or predation [e.g., Park 1962]). An alternative taxonomy of competition was proposed by Schoener (1983). Here, “con- sumptive competition designates consump- tion of resources that deprives other indi- viduals of those resources, whereas preemptive competition occurs when a unit of space is passively occupied by an indi- vidual, thereby causing other individuals not to occupy that space before the original oc- cupant disappears” (Schoener 1988:256). We hypothesize that O. cartilagonodus from the Zambales Mountains may reduce competition between sexes by two means: consumptive competition is reduced by ad- justing to differences in food preferences (based on cranial morphometric data), and preemptive competition is reduced by ad- justing habitat preferences between the sex- es (based on capture data). A test of these hypotheses will necessitate further study in- volving examination of stomach contents from freshly captured individuals, and more extensive sampling along an elevational gra- dient. The data from Mount Isarog, cur- rently under study by L. R. Heaney and coworkers may serve as an additional test of these hypotheses. Clearly, the variation exhibited by this species is not a case of ordinary, clinal geo- graphic variation. The data therefore lend themselves to a number of alternative hy- . potheses to be tested with respect to the relationships among the various popula- tions. Given the exhibited pattern of mor- phological variation, it may well be that the Zambales population represents a long iso- lated population. Additional specimens will be needed to distinguish among competing hypotheses, including additional sampling from the Laguna and Batangas Provinces’ 14 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON montane areas, from which only a few spec- imens are known (and none closely scruti- nized), as well as a greater number of spec- imens from the type locality (in Cordillera Central) and from the Sierra Madre Moun- tains. Until such a time as the regional pop- ulational differences are resolved, the struc- tured, geographically and morphologically cohesive populations of O. cartilagonodus appear to represent an ideal study case for elucidation of zoogeographic relationships among different regions of the island of Lu- zon. Acknowledgments We thank the Protected Areas and Wild- life Bureau of the Philippine Department of the Environment and Natural Resources for facilitating the permits that allowed us to carry out this work. Within these organi- zations, we thank especially A. Alcala, A. Ballesfin, J. Caleda, C. Catibog-Sinha, C. Custodio, D. Ganapin, and S. Penafiel. We thank A.-M. and R. Tate, and the Miranda family for extending welcome hospitality during our long stays in Manila. Our stay in Coto was much helped by assistance from Benguet Mining Company (Manila), es- pecially D. R. Belmonte and E. M. Sacris; accommodations were provided in Coto by Benguet Corporation’s Masinloc Chromite Operation, especially D. B. Delo and D. V. Galang. The intercession of the Jaime Ongping Foundation on our behalf, partic- ularly C. Balaoing, V. Nano, and P. Salva- dor, was much appreciated. Field work was greatly facilitated by the help, advice and experience of R. S. Kennedy and P. C. Gon- zales; much help was obtained in the field from the rest of the CMNH and NMP staff: R. Aquino, R. M. Brown, J. Barcelona, D. T. Busemeyer, R. E. Fernandez, J. W. Fer- ner, D. Gonzales, A. Ippolito, D. D. Keller, H. C. Miranda, Jr., V. Samarita, F. T. Wac- disen, and D. Wechsler. Specimens exam- ined in this work kindly were provided by P. Myers and J. S. Arnold, University of Michigan Museum of Zoology; and M. D. Carleton and L. K. Gordon, Division of Mammals, National Museum of Natural History; L. K. Gordon was particularly helpful. Discussions that greatly enhanced the quality of the manuscript were held with J. W. Bickham, A. L. Gardner, R. L. Honey- cutt, and L. F. Marcus. We thank M. S. Simon, P. D. Heideman, K. F. Koopman, and two anonymous reviewers for review- ing and commenting on preliminary drafts of this manuscript. LAR particularly thanks L. R. Heaney for his help, advice, and ed- itorial assistance throughout the entire phase of this project. This project was funded by a grant from the John T. and Catherine D. MacArthur Foundation to P. C. Gonzales (NMP) and R. S. Kennedy (CMNH). This work constitutes contribution No. 2 of the joint National Museum of the Philippines/ Cincinnati Museum of Natural History Bio- diversity Inventory Project. Literature Cited Brower, J. E., & J.H. Zar. 1984. Field and laboratory methods for general ecology, 2nd. ed. Wm. C. Brown Publishers, Dubuque, Iowa, xi + 226 pp. Bureau of Mines, Philippines, in coordination with the Board of Technical Surveys and Maps. 1963. Geological map of the Philippines, edition 1. Single sheet map. Burton, D. W., J. W. Bickham, & H. H. Genoways. 1989. Flow-cytometric analyses of nuclear DNA content in four families of neotropical bats.— Evolution 43(4):756-765. Cracraft, J. 1983. Species concepts and speciation analysis. Pp. 159-187 in R. F. Johnston, ed., Current ornithology, Volume 1. Plenum Press, New York, xvi + 425 pp. Dickerson, R. E. 1924. Tertiary paleogeography of the Philippines. — Philippine Journal of Science 25:11-50, pls. 1-4. Fleming, T. H. 1973. Thenumber of mammal species in several North and Central American forest communities.— Ecology 54:555-563. Freeman, P. W. 1981. A multivariate study of the family Molossidae (Mammalia: Chiroptera): morphology, ecology, evolution. — Fieldiana, Zoology, New Series, No. 7, vu + 173 pp. Goodman, S. M., & P. C. Gonzales. 1990. The birds of Mt. Isarog National Park, Southern Luzon, Philippines, with particular reference to altitu- VOLUME 107, NUMBER 1 dinal distribution.—Fieldiana, Zoology, New Series, No. 60, iv + 39 pp. Graham, G. L. 1983. Changes in bat species diversity along an elevational gradient up the Peruvian Andes.—Journal of Mammalogy 64:559-571. Heaney, L. R. 1991. An analysis of patterns of dis- tribution and species richness among Philippine fruit bats (Pteropodidae).— Bulletin of the American Museum of Natural History, 206:145- 167. —,, & E.A. Rickart. 1990. Correlations of clades and clines: geographic, elevational, and phylo- genetic distribution patterns among Philippine mammals. Pp. 321—332 inG. Peters and R. Hut- terer, eds., Vertebrates in the tropics. Zoolo- gisches Forschungsinstitut und Museum Alex- ander Koenig, Bonn, 424 pp. , P. C. Gonzales, & A. C. Alcala. 1987. An annotated checklist of the taxonomic and con- servation status of land mammals in the Phil- ippines. —Silliman Journal 34:32-66. , P. D. Heideman, E. A. Rickart, R. B. Utzur- rum, & J.S. H. Klompen. 1989. Elevational zonation of mammals in the central Philip- pines. — Journal of Tropical Ecology 5:259-280. Heideman, P. D., & L.R. Heaney. 1989. Population biology and estimates of abundance of fruit bats (Pteropodidae) in Philippine submontane rain- forest.—Journal of Zoology, London 218:556- 586. , J. A. Cummings, & L. R. Heaney. 1993. Re- productive timing and early development in an Old World Fruit Bat, Otopteropus cartilagono- dus (Megachiroptera). — Journal of Mammalogy 74:62 1-630. H. S. 1966. Measurement of “overlap” in comparative ecological studies.— American Naturalist 100:419-424. Hutcheson, K. 1970. A test for comparing diversities based on the Shannon formula.—Journal of Theoretical Biology 29:151-154. Ingle, N. R., & L. R. Heaney. 1992. A key to the bats of the Philippines.—Fieldiana, Zoology, New Series, No. 69, iv + 44 pp. Johnson, D. H. 1962. Two new murine rodents. — Proceedings of the Biological Society of Wash- ington 75:317-319. Kock, D. 1969. Eine bemerkenswerte neue gattung und art flughunde von Luzon, Philippinen.— Senckenbergiana Biologica 50:329-338. Magurran, A. E. 1988. Ecological diversity and its measurement. Princeton University Press, Princeton, New Jersey, x + 179 pp. Margalef, R. 1972. Homage to Evelyn Hutchinson, or why is there an upper limit to diversity. — Transactions of the Connecticut Academy of Arts and Sciences 44:211-235. Horn, 15 McCoy, E. D., & E. F. Connor. 1980. Latitudinal gradients in the species diversity of North American mammals.—Evolution 34:193-203. McKitrick, M. C., & R. M. Zink. 1988. Species con- cepts in ornithology.—The Condor 90:1-14. McNab, B. K. 1989. Temperature regulation and rate of metabolism in three Bornean bats.— Journal of Mammalogy 70:153-161. Mudar, K. M., & M.S. Allen. 1986. A list of bats from northeastern Luzon, Philippines.—Mam- malia 50:219-225. Park, T. 1962. Beetles, competition and popula- tions.—Science 138:1369-1375. Patterson, B. D., P. L. Meserve, & B. K. Lang. 1989. Distribution and abundance of small mammals along an elevational transect in temperate rain- forests of Chile.—Journal of Mammalogy 70: 67-78. Peet, R. K. 1974. The measurement of species di- versity.—Annual Review of Ecology and Sys- tematics 5:285-307. Philippine Council for Agriculture and Resources Re- search, Soil and Water Resources Research Di- vision. 1977. Map of monthly rainfall and cy- clones in the Philippines, 2nd ed. Pielou, E.C. 1969. An introduction to mathematical ecology. Wiley Interscience, New York, 286 pp. Pianka, E.R. 1983. Evolutionary ecology. Third edi- tion. Harper & Row Publishers, Cambridge, Massachusetts, x11 + 416 pp. Rapoport, E. 1982. Areography: geographical strat- egies of species. Pergamon Press, New York, xvi + 269 pp. Rickart, E. A., L. R. Heaney, & R. C. B. Utzurrum. 1991. Distribution and ecology of small mam- mals along an elevational transect in South- eastern Luzon, Philippines.—Journal of Mam- malogy 72:458—469. Ripley, S. D., & J. T. Marshall, Jr. 1967. A new subspecies of flycatcher from Luzon, Philippine Islands (Aves, Muscicapinae).— Proceedings of the Biological Society of Washington 80:243- 244. Ruedas, L. A., & J. W. Bickham. 1992. Morpholog- ical differentiation between Rhogeessa minutilla and R. tumida (Mammalia: Chiroptera: Ves- pertilionidae).— Proceedings of the Biological Society of Washington 105:403-409. SAS Institute, Inc. 1988a. SAS/STAT user’s guide, release 6.03 edition. Cary, North Carolina, SAS Institute, Inc., 1028 pp. 1988b. SAS procedures guide, release 6.03 edition. Cary, North Carolina, SAS Institute, Inc., 441 pp. Schoener, T. W. 1983. Field experiments on inter- specific competition.—American Naturalist 122: 240-285. 16 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 1988. Ecological interactions. Pp. 255-297 in A. A. Myers and P. S. Giller, eds., Analytical biogeography; an integrated approach to the study of animal and plant populations. Chap- man and Hall, New York, xiii + 578 pp. Shannon, C.E. 1948. A mathematical theory of com- munication.— Bell System Technical Journal 27: 379-423. Simpson, E. H. 1949. Measurement of diversity. — Nature 163:688. Sokal, R. R., & F. J. Rohlf. 1981. Biometry, second edition. W. H. Freeman and Co., New York, Xvill + 859 pp. Terborgh, J. 1971. Distribution on environmental gradients: theory and a preliminary interpreta- tion of distributional patterns in the avifauna of the Cordillera Vilcabamba, Peru.—Ecology 52: 23-40. 1977. Bird species diversity on an Andean elevational gradient.— Ecology 58:1007-1019. UNESCO/ECAFE. 1971. Geologic map of southeast Asia. United Nations Publication No. 69-30632. Whitmore, T. C. 1984. Tropical rain forests of the Far East. Clarendon Press, Oxford, England, xvi + 352 pp. Zar, J.H. 1984. Biostatistical analysis, 2nd. ed. Pren- tice-Hall, Inc., Englewood Cliffs, New Jersey, xiv + 718 pp. (LAR, JRD) Department of Vertebrate Zoology, Cincinnati Museum of Natural History, 1720 Gilbert Avenue, Cincinnati, Ohio 45202-1401, U.S.A.; (SRD) Present address: University of Alaska Museum, Mammal Collections, University of Alaska, Fairbanks, Alaska 99775-1200, U.S.A..: (RVS) Division of Zoology, National Mu- seum of the Philippines, Executive House, P. Burgos Street, Rizal Park, Manila, Phil- ippines. PROC. BIOL. SOC. WASH. 107(1), 1994, pp. 17-26 CRYPTOPHOCA, NEW GENUS FOR PHOCA MAEOTICA (MAMMALIA: PINNIPEDIA: PHOCINAE), FROM UPPER MIOCENE DEPOSITS IN THE NORTHERN BLACK SEA REGION Irina A. Koretsky and Clayton E. Ray Abstract. —Review of relevant fossil and Recent material indicates that the fossil seal species Phoca maeotica Nordmann (1860) from Kishinev, Moldavia, earlier assigned to the Monachinae, actually belongs to the subfamily Phocinae. This species differs from all known seals and is placed in the separate genus Cryptophoca. Cryptophoca maeotica differs from other seals in its straight sym- physeal region of mandible, position of mental protuberance between p* and p*, rectangular greater trochanter of femur, and length of crest of greater tubercle of humerus extending 4 length of humerus. Presence of these characters not only justifies the specific independence of the Moldavian seal, but also requires its placement in a separate genus, Cryptophoca. Nordmann (1860:3 13-326) described the species Phoca maeotica on the basis of dis- sociated postcranial bones extracted from material originally assigned by Eichwald (1850) to his Phoca pontica. Nordmann sug- gested the close affinity of this large seal with monk seals. Later, Trouessart (1898-1899, 1904), never questioning this relationship, transferred the species to the genus Mon- atherium, belonging to the subfamily Mon- achinae. However, the taxonomic position of this species has since been questioned. For instance, some systematists (Grigorescu 1977, Kellogg 1922, McLaren 1960, de Muizon 1992) classified (on the basis of the size of the distal epiphysis of femur) Phoca maeotica with the monachine seals, whereas others (Alekseev 1924, 1926; Bogachev 1927; Kirpichnikov 1961; Ray 1977; Sav- age & Russell 1983) considered it to be a phocine seal, or did not mention the place- ment of this species (de Muizon 1982). In recent years, discoveries of cranial and postcranial remains of Phoca maeotica in the northern Black Sea region have greatly augmented the classic collection of von Nordmann. Although several taxa of pho- cids occur in the same deposits with the nominal species Phoca maeotica, we feel confident in our assignment here of the mandible and humeri on the basis of sim- ilarity in size and morphology and analogy with modern phocids. Making use of all available material, we conclude that the K1- shinev seal belongs to the subfamily Pho- cinae on the basis of length of the symphy- seal part of the mandible to the anterior border of the alveolus of p2, presence of the mental protuberance; trochlear crest of the humerus not separated from coronoid fossa by distinct lip; different size of the femoral condyles, small difference in the width of proximal and distal epiphyses, and absence or poor development of intertrochanteric crest. Furthermore, our study of the osteologi- cal remains of Phoca maeotica not only cor- roborates its specific distinctness, but also requires distinguishing it as a new genus, Cryptophoca, described here. Morphometric analysis of the humeri, femora, and mandibles was carried out us- ing the methods of Sergienko (1967), Burns & Fay (1970), Piérard (1971) and one of us 18 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON (Koretsky 1986). Although the question of sexual dimorphism must be kept in mind in all studies of fossil pinnipeds (Koretsky 1987, Van Bree & Erdbrink 1987), it could not be addressed meaningfully here because of the very limited material. We are well aware of the recent intensive interest in pinniped phylogeny and classi- fication, resulting in a currently unstable sit- uation (e.g., Wozencraft 1989; Wyss 1988, 1989). Studies in progress by us and others will add much new information in the near future. Meanwhile, we feel that it would be premature and unproductive to address these broad questions in the present paper of very limited scope and objectives. Family Phocidae Gray, 1825 Subfamily Phocinae Gill, 1866 Cryptophoca, new genus Type species.—Phoca maeotica Nord- mann, 1860:321, pl. 23, figs. 8, 9. Diagnosis. —Lower canine and pl very large (Fig. la, b), pl single-rooted; symphy- seal part straight, its inner part thickened from anterior alveolus of p2 to canine; men- tal protuberance located between p3 and p4. Deltoid crest up to 4 of humeral length, not reaching radial fossa and proximal border 1s widest part (Figs. 2a, b; 4a, b); lesser tu- bercle of humerus located on same level as proximal border of deltoid crest; head round. Femur with almost rectangular greater tro- chanter; trochanteric fossa deep and open; head of femur big (Figs. 3a, b; 4c, d), situ- ated on relatively narrow, short neck; small- est width of diaphysis shifted toward prox- imal epiphysis; greatest breadth across condyles 20.0%-21.0% of bone length; proximal epiphysis narrower than distal by 2.0%—8 .0%. Comparison. —The genus Cryptophoca differs from other known seals by: straight shape of symphyseal part of mandible (ex- cept for Pagophoca), smaller height of body of mandible under p2 (except for Phoca, Halichoerus, and Pagophoca); lower posi- tion of lesser tubercle of humerus relative to head and its location on same level with proximal border of deltoid crest (except for Erignathus, Praepusa, and Monachopsis): less developed spiral groove (except for Erignathus, Pagophoca, and Monachopsis); large rectangular greater trochanter (except for Praepusa); relatively smaller intercon- dylar width of femur (except for Erignathus and Praepusa); narrower neck of femur (ex- cept for Phoca, Halichoerus, Pagophoca, and Praepusa); peculiar shape of both humerus and femur (Table 1). Available material al- lows us to hypothesize that, based on sim- ilarities in the size and character of man- dible and limb bones, this genus is closely related to Pagophoca. In addition this genus differs distinctly from other genera as follows: From Phoca by flattened body of man- dible; mental protuberance not labially bent; greater length of p4 alveolus relative to m1, alveolus; larger diastemata between teeth: single-rooted pl. Relatively greater length of deltoid crest of humerus. Bigger size of intertrochanteric crest; relatively smaller condyles of femur. From Pusa by larger dimensions; greater depth of body of mandible under p2; for- ward shifted mental protuberance. Absence of intertubercular grooves of humerus; large head; greater length of deltoid crest. Large swelling of intertrochanteric crest; deeper and wider trochanteric fossa, relatively larg- er head of femur. From Erignathus by smaller dimensions; mental protuberance of mandible slightly pronounced, shifted forward and not labi- ally bent. Relatively larger length of deltoid crest of humerus and latter’s widening on proximal border; round shape of head. Large height of greater trochanter relative to fem- oral neck; smaller size of neck; relatively narrower proximal epiphysis. From Halichoerus by greater mandible flattening; pronounced mental protuber- ance; double-rooted p2-m1. Lateral posi- tion of deltoid crest of humerus; absence of 19 VOLUME 107, NUMBER 1 = =a = re = = = + UIPIM Ie[ApUodIO}UT [Jews APOAL[OY ae = = = = = - + INULYIOI 10}e913 Ie[NdUeLIOOI O31eT InwdJ st = = = = = = + BIPIOUOIOD BSSOJ S9YIVII VOPIOI[IP BISLID = = = = = tL = ae punol jndep & tL = = = = = + BOPIO}f[op BISLIO Jo pied [eullxo1d SP [OAO[ OWLS UO Po}edO] IO}UPYOI] IossaT sniouny + + + + — —+ —+ + 9ZIS Ul Ie]IUWIs |W pue pd jo TosaTy a = te = + = dt + piemioy ¢d wor 19y91U1 Apog = 4 = a = + + + jussold vBUIOISeIG = Su = = = + = + Apoq uey} JomOLeU jIed Ie[OSATY =a = = a = + = ae yied jeosAyduids jo adeys ae = = = = — ab + QousUIWIOId UIYD JO UONeI0T s[qipueyy psndavigd snyjousisq SNAIOYINOH psng DIOYd po0ydosng p20ydo1jsipy po0ydojdtuy “O[QuLIVA JajovIvYyO = — + “ILIIUIISSIP 10 JUaSqe JoJORIeYO = — “IL[TUIIS 10 jUSSoId 19j0vIeYO = + :oeUTDOYg A[IWIeJQns dy) JO SI9}DvIvYO ONsOUseIP sATIeIedUIOD—"| 21qGe 1, 20 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON (0) 5 cM Fig. 1. Cryptophoca maeotica. Cast of the left mandibular ramus, without number. Tiraspol State Pedagogical Institute, Moldavia. a = labial aspect, b = occlusal aspect. intertubercular groove; round head. Pres- of proximal border. Deeper and wider tro- ence of intertrochanteric crest of femur. chanteric fossa of femur. From Pagophoca by forward-shifted From Praepusa by far greater dimensions; mental protuberance. Lateral position of larger height of body of mandible under p2 deltoid crest of humerus and its widening to canine. Equal position of lesser tubercle DCM Fig. 2. Cryptophoca maeotica. Left humerus 64-530 from Kishinev. Institute of Zoology of Academy of Science of Ukraine, Kiev. a = lateral aspect, b = cranial aspect. VOLUME 107, NUMBER 1 21 Table 2.—Means (+.SE) and range for measurements (mm) of number in sample (n) of humeri from Cryp- tophoca maeotica. Measurements Absolute length Length of deltoid crest Height of head Height of trochlea Width of head Width of deltoid crest Width of distal epiphysis Width of proximal epiphysis Width of trochlea distally Width of trochlea, frontal view Transverse width of diaphysis Thickness of proximal epiphysis Thickness of medial condyle Thickness of lateral condyle Diameter of diaphysis with deltoid crest of humerus and proximal border of deltoid crest; slightly bigger index of head width. Smallest width of femoral diaphysis shifted toward proximal epiphysis; relatively smaller width of proximal epiphysis, large but relatively more narrowly arranged con- dyles. From Monachopsis by far bigger dimen- sions; double-rooted p2-m1 with large diaste- mata. Round head of humerus; short deltoid crest; higher and wider medial epicondyle. Deeper and more elongated trochanteric fossa of femur; relatively more narrow proximal epiphysis. Distribution. —Middle Sarmatian (Upper Miocene) of the northern Black Sea region of the Ukraine and Moldavia. Etymology. —From the Latin “‘crypti- cus,”’ 1.e., hidden, secretive; ““~phoca’’—seal. Cryptophoca maeotica (Nordmann, 1860) Figs. 1-4, Tables 2-3 Phoca pontica Eichwald, 1850 (in part):210- 218.—Eichwald, 1853 (in part):391—400. Phoca Nordmann, 1858: pl. 23, figs. 1, 2, 8 95 IO, Phoca maeotica Nordmann, 1860 (in part): 320-321, 356-357.— Van Beneden, 1877: 26.—Toula, 1898:50.—Alekseev, 1924: n X + SE Range 5 107.1 + 1.5 99.0-123.5 5 75.9 + 0.7 73.0-80.0 4 Dreilton 0) eZ) 24.0-28.0 4 20.1 + 0.7 19.0-21.5 4 25.0 + 0.9 23.0-28.0 4 28.6 + 0.2 28.0-30.0 5 37.0 + 1.8 30.0—45.0 7 34.2 + 2.0 29.0-38.5 4 19.1 + 0.6 18.0-20.0 4 23.1 + 0.8 22.0-25.0 5 14.5 + 0.9 12.0-17.0 7 40.5 + 1.8 33.8—-46.0 4 19.6 + 0.6 18.5-20.5 4 17.4 + 0.4 16.4-18.5 6 33.5 + 1.6 29.0-38.0 202.— Alekseev, 1926:138-143.—Savage & Russell, 1983:187, 292-294. Monatherium maeoticum —Trouessart, 1897:380.—Trouessart, 1904:283.— King, 1964:131.—King, 1983:132.— Friant, 1947:50 (non pl. 1, fig. 2a—c). Monotherium maeoticum— Kellogg, 1922: 114.—Kretzoi, 1941:353.—McLaren, 1960:50-52, 56—57, fig. le.—Grigorescu, 1976:407, 413-415, 417, fig. Sb.—de Muizon, 1982:202—205. Monotherium maeotica—Grigorescu, 1976: 407. “Monotherium”™’ maesticum (sic)—de Mui- zon, 1992:37. Lectotype. —Left femur N1815, Museum of Zoology, Helsinki, Finland; illustrated by Nordmann (1858: pl. 23, figs. 8, 9) as Phoca and described and named as Phoca maeo- tica in 1860. (Fig. 4c, d). Type locality. —Moldavia (Kishinev), northern Black Sea region. Geological age.—Upper Miocene, Mid- dle Sarmatian (Bessarabian Formation). Material examined.—Moldavia, Kishi- nev: Institute of Zoology of Academy of Sci- ences of Ukraine, Kiev (von Nordmann col- lection), collection 64—eight femora, a part of the material described by Nordmann 22 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON a 90M Fig. 3. Cryptophoca maeotica. Left femur 1713/23 from Kishinev. Paleontological Institute of Academy of Science of Russia, Moscow. a = caudal aspect, b = cranial aspect. (1860)—three humeri, without numbers; Paleontological Institute of Academy of Sci- ences of Russia, Moscow, collection 1713— nine femora, 1713/1329 and 1713/1330— two incomplete rami of the mandible; Pa- leontological Museum of Odessa State Uni- versity — five femora, the material described by Alekseev (1926); Tiraspol State Peda- gogical Institute, Tiraspol—incomplete ra- mus of the left mandible, without number; Paleontological Department of Museum of Zoology, Helsinki, Finland (von Nordmann collection)—right humerus 1812 (illustrat- ed by Nordmann, 1858: pl. 23, figs. 1, 2), proximal part of left femur N1816 (illus- trated by Nordmann by 1858: pl. 23, fig. 10), left femur 1815 (illustrated by Nord- mann, 1858: pl. 23, figs. 8, 9), six femora without numbers (not illustrated); cast of left femur National Museum of Natural History, Smithsonian Institution, Washing- ton, D.C., U.S., 214979, Geological Mu- seum of University of Bucarest, Romania 259/II,5c (Simionescu collection). Ukraine, Crimea, Kerchensky peninsula (Kamysh- Burun): Institute of Zoology Academy of Sciences Ukraine, collection 64 —three fem- ora. Remarks.—McLaren (1960:56—57), working from literature, designated left fe- mur N1815 as lectotype. Unfortunately, at some time subsequent to Nordmann’s work the specimen was broken and the proximal half misplaced or lost. The breakage must have occurred prior to 1972, when the spec- imen was cataloged, as its number is afhxed to the broken proximal surface. All recent efforts by the late BjOrn Kurtén and by Mi- kael Fortelius to locate the missing part have been unsuccessful. However, there are sev- VOLUME 107, NUMBER 1 a b Fig. 4. 23 a, b. Cryptophoca maeotica. Right humerus 1812 from Kishinev (from Nordmann, 1858: pl. 23, figs. 1, 2). Paleontological Department of Museum of Zoology, Helsinki. a = caudal aspect, b = lateral aspect; c, d. Cryptophoca maeotica. Lectotype, left femur 1815 from Kishinev (from Nordmann, 1858: pl. 23, figs. 8, 9). Paleontological Department of Museum of Zoology, Helsinki. c = cranial aspect, d = caudal aspect. eral good illustrations of the intact femur and the distal half of the lectotype femur survives. The Russian version of Eichwald’s mono- graph, which we cite herein, was published in 1850, 3 years earlier than the German edition. For nomenclature it is very impor- tant, because western researchers seem to have been unaware of the Russian text. A few other elements (innominates, vertebrae and a radius) are not described here, but possibly pertain to this taxon. Diagnosis.— As for the genus. Description. —True seal close in size to the contemporary genus Pagophoca. Mandible (Fig. 1). Not high, flat from lin- gual side. From labial side, body of man- dible thickened in middle from level of an- terior p2 alveolus to beginning of ascending ramus. All teeth arranged in alignment with the tooth row axis. Alveolar length of m1 smaller than that of p4; retromolar space shortened. Mental protuberance located be- tween anterior alveolus of p3 and anterior alveolus of p4. Maximal depth of mandible between alveoli of p2 and p4. Symphyseal part straight and thick, 1.e., the lower border of the mandible not elevated with respect to alveolus of canine. Alveolus of pl, sim- ilar to canine alveolus, is very large. Mea- surements of mandible: depth under ml — 20.5—26 mm; depth under p2— 18-19 mm; depth between p3-p4—20-—24 mm; depth behind m!— 18.5-18.5 mm; alveolar length of row p1-p4—35.5—40 mm; alveolar length of row pl—m1—59 mm; alveolar length of p4—8-9 mm; alveolar length of ml —6—7 mm; length of diastema p4-m1—6.5—9 mm; thickness of mandible under m1 —9-11 mm. Humerus (Fig. 2, Table 2). Lesser tubercle of humerus slightly elongated and departing only slightly from basic axis (which is prob- ably characteristic of all extinct true seals). Intertubercular groove only slightly dis- cernible. Deltoid crest widest proximally. Deltoid tuberosity located along middle of 24 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 3.—Means (+SE) and range for measurements (mm) of number in sample (n) of femora from Cryp- tophoca maeotica. Measurements Absolute length Medial length Lateral length Length of medial condyle Length of lateral condyle Length of greater trochanter Intertrochanter length Height of head Height of articular area of patella surface Width of proximal epiphysis Width of distal epiphysis Width of condyles Width of greater trochanter Width of head Width of diaphysis Anteroposterior thickness of diaphysis Thickness of medial condyle Thickness of lateral condyle Distance between condyles Diameter of neck diaphysis. Radial fossa narrow and shallow. Epicondyles well developed. Medial epi- condyle reaching distal part of deltoid crest; lateral epicondyle spreading from lower part of entepicondyloideum and ending below medial epicondyle. Spiral groove not pro- nounced. Femur (Fig. 3, Table 3). Greater trochan- ter much higher than the head, approaching rectangular shape. Trochanteric fossa wide, deep, and open. Intertrochanteric crest lo- cated along middle part of femur, below tro- chanteric fossa. Smallest width of diaphysis shifted to proximal part of femur. Distribution.—Upper Miocene, Middle Sarmatian (Bessarabian Formation) of the northern Black Sea region of Ukraine and Moldavia. Acknowledgments We wish to express our gratitude to Dr. Yury A. Semenov, Institute of Zoology, Ukrainian Academy of Sciences, Kiev, for support of this study, reading an earlier ver- sion of this manuscript, and for strength- n X + SE Range 23 106.0 + 2.4 93.0-138.0 13 96.0 + 3.2 87.0-110.4 12 92.9 + 2.8 82.0-110.4 19 18.8 + 0.4 17.0-21.0 22 22.4 + 0.5 18.0-26.5 23 26.9 + 0.8 25.0-33.5 9 Bil se il? 24.0-34.5 17 20.0 + 0.5 18.0—23.0 17 Dds) 3= (0).7/ 18.5-25.0 23 51.8 + 1.2 44.3-64.2 25 53.4 + 0.6 47.0-62.3 23 43.3 + 0.7 38.0-49.0 24 18.7 + 0.4 16.0—22.0 Dy} DN z= 0.5) 18.5—25.0 30 27.6 + 0.4 23.0-33.0 12 12.4 + 0.6 12.0-17.0 12 23.7 + 0.7 21.0-27.5 15 26.1 + 0.7 23.5-29.0 18 WoW se It3} 8.0-12.0 ay) 16.2 + 0.4 14.0-19.5 ening it with his suggestions. Dr. Arseni A. Antoniuk, Military Medical Academy, Saint-Petersburg (Leningrad), assisted Ko- retsky in the field and introduced her to the Nordmann collection in the Institute of Zo- ology, Russian Academy of Sciences, Saint- Petersburg. We thank the late Dr. Bjorn Kurtén, and Drs. Ann Forstén and Mikael Fortelius for loan of all fossil seals from the Nordmann collection in the Museum of Zo- ology, Helsinki. We also thank Prof. Léo- nard Ginsburg of Muséum National d’His- toire Naturelle, Paris, for providing casts. We thank Victor E. Krantz and Jennifer Jett, both of the National Museum of Natural History, Smithsonian Institution, for pho- tographs, and for assistance with the tables, respectively. Literature Cited Alekseev, A. K. 1924. Seals in the Sarmatian deposits of southern Russia.—Journal Nauchno-Issle- dovatelskich kafedr v Odesse 1(10-11):201-—205. (Russian text). . 1926. Seals in the Sarmatian deflection of the VOLUME 107, NUMBER 1 south of the U.S.S.R.—Journal Nauchno-Issle- dovatelskich kafedr v Odesse 11(4):138-143. (Russian text). Bogachev, V. V. 1927. The seal in the Miocene of the Caspian Basin.—Izvestia Azerbaijan Go- sudarstvennogo Institute of Yestestvoznaniye 1 medicine 6:135—148. (Russian text). Burns, J. J., & F. H. Fay. 1970. Comparative mor- phology of the skull of the Ribbon seal, His- triophoca fasciata, with remarks on systematics of Phocidae.—Journal of Zoology (London) 161(3):363-394. Eichwald, E. 1850. Lethaea Rossica. Saint-Peters- burg, 520 pp. Atlas, 13 pls. (@n Russian). 1853. Lethaea Rossica, ou Paleontologie de la Russie 3:520 pp. 37 figs., 13 pls., Stuttgart, Schweizerbart. Friant, M. 1947. Recherches sur le femur des Pho- cidae. — Bulletin,du Musée royal d’Histoire na- turelle de Belgique 23(2):1-51. Gill, T. N. 1866. Prodrome of a monograph of the pinnipeds. — Proceedings Essex Institute, Salem, Communications 5:3-13. Gray, J. E. 1825. An outline of an attempt at the disposition of Mammalia into tribes and fam- ilies, with a list of the genera apparently apper- taining to each tribe.—Annals of Philosophy, N.S. 10 (vol. 26 of the whole series):337—344. London. Grigorescu, D. 1977. Paratethyan seals.—Systematic Zoology 25(4) (for December 1976):407-419. Kellogg, R. 1922. Pinnipeds from Miocene and Pleis- tocene deposits of California.—University of California Publications. Bulletin of the Depart- ment of Geological Sciences 13(4):23-132. King, J. 1964. Seals of the world. British Museum (Natural History), London, 154 pp. 1983. Seals of the world. British Museum (Natural History), London, and Cornell Uni- versity Press, Ithaca, 2nd Edition, 240 pp. Kirpichnikov, A. A. 1961. Essay of the History of the studying of marine mammals of Sarmatian Deposit and adjoining countries.—Akademia Nauk S.S.S.R., Ikhtiologicheskaya Komissia, Trudy soveshchania, Leningrad 12:25-39. (Russian text). Koretskaya, I. A., (Koretsky). 1986. Modern status of systematics of Miocene seals of the Family Phocidae of the Northern Black Sea Province. — Abstract of [Xx All-Union Conference on Marine Mammals, Archangelsk:206—207. (in Russian). . 1987. Sexual dimorphism in the structure of the humerus and femur of Monachopsis pontica (Pinnipedia: Phocinae).— Vestnik Zoologii, Kiev 4:77-82. (in Russian). Kretzoi, M. 1941. Seehund-Reste aus dem Sarmat von Erd bei Budapest. —Féldtani Kézloény 71(7- 12):350-356. 25 McLaren, I. A. 1960. On the origin of the Caspian and Baikal seals and the paleoclimatological im- plication.—American Journal of Science 258: 47-65. Muizon, C. de. 1982. Phocid phylogeny and dis- persal.—Annals of the South African Museum 89(2):175-213, 9 figs. 1992. Paldontologie. Pp. 34-40 in R. Duguy & D. Robineau, eds., Band 6: Meeressduger. Teil II: Robben—Pinnipedia.—Jochen Niethammer & Franz Krapp, eds. Handbuch der Saugetiere Europas. AULA-Verlag, Wiesbaden: 1—309. Nordmann, A. D. von. 1860. [1858]. Palaeontologie Suedrusslands. H.C.Fric. Helsingfors, 360 pp. (Atlas of 28 pls.) Piérard, J. 1971. Osteology and myology of the wed- dell seal Leptonychotes weddelli (Lesson, 1826). Pp. 53-108 in W. H. Burt, ed., Antarctic Pin- nipedia. National Academy of Sciences—Na- tional Research Council. Antarctic Research Se- ries 18:1-226. Ray, C. E. 1977. Geography of phocid evolution.— Systematic Zoology 25(4) (for December 1976): 391-406. Savage, D. E., & D. E. Russell. 1983. Mammalian paleofaunas of the world. Don Mills, London, Amsterdam; Addison-Wesley Publishing Co., Inc., Sydney, Tokyo, 432 pp. Sergienko, N.I. 1967. On the taxonomic significance of some parts of the postcranial skeleton of the seals from genus Pusa (in Russian). — Trudy Po- lyarnogo Institute Morskogo Rybnogo Kho- Ziaistva e Okeanography (PINRO), Murmansk 21:185-193. Toula, F. 1898. Phoca vindobonensis n. sp. von Nuss- dorf in Wien. Beitrage zur Palaontologie und Geologie Oesterreich-Ungarns und des Orients, Mittheilungen des Paladontologischen Institutes der Universitat Wien 11(2):47-70. Trouessart, E.-L. 1897. Carnivora, Pinnipedia, Ro- dentia I.—Fascicle II, Catalogus mammalium tam viventium quam fossilium. Nova editio, Berlin, 644 pp. 1898-1899. Tillodontia, Ungulata. Fascicle IV (665-1469 pp.)—Catalogus mammalium tam viventium quam fossilium. Nova editio, Berlin. 1904. Primates, Prosimiae, Chiroptera, In- sectivora, Carnivora, Pinnipedia. Fascicle I (iv + 288 pp.), Catalogus mammalium tam viven- tium quam fossilium. Quinquennale supple- mentum, anno 1904, Berlin. Van Beneden, P.-J. 1877. Description des Ossements fossiles des environs d’Anvers, premiére partie. Pinnipédes ou Amphithériens.— Musee Royal d’Histoire Naturelle de Belgique, Annales 1:1— 88, 18 pls., 17 figs., 9 maps. Van Bree, P. J. H., & D. P. B. Erdbrink. 1987. Fossil Phocidae in some Dutch Collections (Mam- 26 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON malia, Carnivora).—Bleaufortia, Institute of Taxonomic Zoology (Zoological Museum) Uni- versity of Amsterdam 37(3):43-66. Wozencraft, W. C. 1989. The phylogeny of the Re- cent Carnivora. Pp. 495-593 in J. L. Gittleman, ed., Carnivore behavior, ecology, and evolution. Chapter 18. Cornell University Press, Ithaca, New York, 593 pp. Wyss, A. R. 1988. On “Retrogression” in the evo- lution of the Phocinae and phylogenetic affini- ties of the Monk seals.—American Museum of Natural History Novitates 2924:1-38. 1989. Flippers and pinniped phylogeny: has the problem of convergence been overrated?— Marine Mammal Science 5(4):343-360. Department of Paleobiology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560, U.S.A. PROC. BIOL. SOC. WASH. 107(1), 1994, pp. 27-51 A TAXONOMIC REVIEW OF DENDROICA PETECHIA (YELLOW WARBLER) (AVES: PARULINAE) M. Ralph Browning Abstract.—The taxonomy and nomenclature of Dendroica petechia (Yellow Warbler), last discussed from the entire range of the species about 57 years ago, is reviewed. There are 43 recognizable subspecies based on geographic variation of plumage color and pattern of about 2500 specimens examined. Two new subspecies are named from populations of the migratory aestiva subspecies group of North America. New subspecies are also named that rep- resent populations of the resident subspecies groups erithachorides of Middle and South American and petechia of the West Indies. The Yellow Warbler, Dendroica petechia, comprises three groups of subspecies: the aestiva group 1s migratory and breeds in the Nearctic; the petechia group is resident in the West Indies; and the erithachorides group is resident on both coasts of Middle Amer- ica and northern South America (A.O.U. [American Ornithologists’ Union] 1983). The three groups were treated as separate species until Hellmayr (1935) combined the petechia and erithachorides groups, and Al- drich (1942) combined the petechia and aes- tiva groups. The A.O.U. (1944, 1945) com- bined all three groups as D. petechia. Lowery & Monroe (1968) recognized 34 subspecies in the combined groups of D. petechia. Ol- son (1980) named three additional subspe- cies from the erithachorides group. Relatively recent discussions of geo- graphic variation and taxonomy of Den- droica petechia have been limited to re- gional studies in North America (e.g., Phillips et al. 1964, Oberholser 1974, Rav- eling & Warner 1978, Godfrey 1986), Mex- ico (e.g., Parkes & Dickerman 1967), the Pacific coast of Middle America (Olson 1980), Panama (Wetmore et al. 1984), and islands off northern South America (e.g., Voous 1957, Phelps & Phelps 1950). Re- views of the entire species that discussed subspecific characters include Ridgway (1902) and Hellmayr (1935). These studies usually characterized subspecies of D. pe- techia on the basis of variation in plumage color and pattern; measurements were used secondarily in characterizing only a few sub- species. Peter’s (1927) review of the sub- species of the petechia group included stan- dard measurements, and Hellmayr (1935) provided measurements for some subspe- cies. Hellmayr usually characterized size in terms of such as “smaller,” “‘averages slight- ly larger,” and “‘slenderer.” Raveling & Warner (1978) reported statistical differ- ences in some measurements of specimens from North American populations, but, as with Hellmayr’s measurements, means dif- fered only 2-3 mm and ranges overlapped considerably. Broad patterns of geographic variation in size within the subspecies groups (Wiedenfeld 1991) suggest that measure- ments are of little use in identifying most subspecies. The purpose of this study of D. petechia is to evaluate geographic variation and to determine the morphological limits of the subspecies. Color and pattern of breeding plumages of adult males and females form the basis for subspecific identifications. Oth- er plumages, when known, are described, but these are less useful in characterizing popu- lations than breeding plumages and are poor- 28 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ly represented in collections. Additional col- lecting is essential (see Winker et al. 1991) to document migration and winter distri- bution of subspecies of D. petechia. Methods About 2500 specimens from the breeding grounds of D. petechia were examined for variation in color. Measurements reported in the accounts include standard measure- ments of flight feathers, tarsi, and bill length (anterior edge of the nostril to the tip). Mea- surements are discussed where size is a use- ful taxonomic character. Evaluation of col- ors was by comparison of specimens in daylight conditions. Seasonally similar plumages were compared when possible. The color of the crown (excluding the chest- nut or rusty color), back, and rump is de- scribed as darker (= more black), greener (less yellow), or the opposite, more yellow = nearest Spectrum Yellow of Smithe [1975]). The yellow of the underparts is de- scribed according to observed intensity (brightness) and purity; the yellow of some subspecies is relatively dull (more white) while in some others it exhibits a greenish cast. The chestnut markings on the head and the ventral streaks on the underparts are characterized as dark or pale. The extent of chestnut markings was characterized on the basis of the relative amount on the head, throat, and breast, the relative width (wide vs. narrow) and density of ventral streaks. The character “chestnut areas”’ refers to all chestnut parts of the plumage. Color and pattern were determined subjectively and were characterized in terms relative to the populations being compared. This method served to verify many previously published taxonomic conclusions on D. petechia. Each subspecies account includes its au- thor, year and type locality. Locations of holotypes, when known, are indicated by abbreviation of museums (see Acknowledg- ments); abbreviations in parentheses indi- cate holotypes I examined. Other locations are indicated by a reference. The section ““Subspecific characters” includes breeding plumages of adult male and female, with comparisons to geographically adjacent or morphologically similar subspecies. Non- breeding plumages, when known, are men- tioned for migratory species. “Distribu- tion” includes breeding and winter ranges and localities of intergradation between ad- jacent subspecies. Winter ranges are includ- ed when they differ from those in Lowery & Monroe (1968). Maps show the breeding distributions of the subspecies in the aestiva group (Fig. 1) and in the petechia and eritha- chorides groups (Fig. 2). Use of subspecies groups in this paper is not meant to reflect historical relationships. ““‘Specimens ex- amined” are included as an abbreviated list by general locality and number examined. Few specimens were needed to characterize subspecies represented by populations with insular or linear ranges (e.g., petechia and erithachorides groups). The ““Remarks”’ sec- tion includes discussions on nomenclature, synonymies, and geographic variation. The sequence of subspecies is only slightly mod- ified from that by Lowery & Monroe (1968). A synopsis of the pattern of variation is presented at the end of each of the three subspecies groups. Dendroica petechia aestiva subspecies group Subspecies in the aestiva group differ from members of the petechia and erithachorides groups in generally lacking chestnut on the head except on the crown, where, if present, it is diffuse or concentrated on the feather shafts. Individuals in the aestiva group have more pointed wings than birds of the other groups. Dendroica petechia rubiginosa (Pallas) Motacilla rubiginosa Pallas, 1811:496 (“in insula Kodiak” = Kodiak Island, Alaska). Subspecific characters. —Males nearest Yukon subspecies (named below) but darker and greener above. Compared with Mac- VOLUME 107, NUMBER 1 : >» a brewsteri Fig. 1. kenzie subspecies (see account below), males paler and more yellowish above; bill longer (see below). Compared with all subspecies in the aestiva subspecies group, males usu- ally with greenish forehead. Females with greener (less yellow) crown and rump than Yukon subspecies; slightly greener above than Mackenzie subspecies. Distribution. — Breeds from Unimak Is- land to Alaska Peninsula, Kodiak Island, and coastal British Columbia. Intergrades with Yukon subspecies along coastal Alaska (see below); intergrades with morcomi in British Columbia (Bella Coola, Vancouver Island and adjacent mainland). Migrates rarely east of the Rocky Mountains; indi- viduals identified from Pennsylvania, parkesi Approximate breeding ranges of subspecies in the Dendroica petechia aestiva subspecies group. Washington, D.C., Mississippi, and Florida (Parkes 1968). Specimens examined.—Coastal Alaska (31); British Columbia (29). Remarks. —Some migrants that were identified in the literature as rubiginosa may represent intergrades between rubiginosa and the next subspecies, or birds from Mac- kenzie District, Northwest Territories (named below). Further collecting and study of specimens in nonbreeding plumages from these northern populations are needed. Dendroica petechia banksi, new subspecies Holotype. —USNM 468183, adult male, Old Crow Village, Yukon Territory, Cana- 30 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ) 10 ES meee , or Neen, ~ x yi, oe = ee, EA ee, Se D a Z 04 enc _ we yet “caewnanensn 4 Fig. 2. Breeding ranges of subspecies of Dendroica petechia in the petechia and erithachorides subspecies groups: 1, castaneiceps; 2, rhizophorae; 3, phillipsi; 4, xanthotera; 5, aithocorys; 6, iguanae; 7, aequatorialis; 8, jubaris; 9, peruviana; 10, aureola; 11, oraria; 12, rufivertex; 13, bryanti; 14, erithachorides; 15, gundlachi; 16, flaviceps; 17, eoa; 18, solaris; 19, chlora; 20, albicollis, 21, barthomelica, 22, melanoptera; 23, ruficapilla; 24, babad; 25, petechia; 26, alsiosa; 27, rufopileata; 28, aurifrons; 29, obscura; 30, cienagae; 31, paraguanae; 32, chrysendeta; 33, flavida; 34, armouri. da, collected 7 June 1957 by L. Irving and L. Peyton (original number 291). Subspecific characters. —Males more yel- low above, especially rump and forehead, than rubiginosa. Males paler and more yel- low above than Mackenzie District subspe- cies (named below). Males similar to am- nicola but more yellow above; slightly greener above than morcomi; chestnut streaks average darker and more prominent than rubiginosa, amnicola and morcomi. Females most similar to amnicola from cen- tral Canada but average slightly greener above. Males in fall plumage more yellow above than rubiginosa. Compared with ru- biginosa and Mackenzie subspecies, females more yellow (less green) above. Distribution. — Breeds (or probably breeds) from Old Crow to Selkirk, Yukon Territory, and in Alaska, along Yukon River (Charley River, Circle, Galena), north of the Arctic Circle (Kuguruok [= Canning] River, Um- iat, Anaktuvuk, Sheenjek and Alatna rivers, Siruk Creek, Bettles), along Tanana River (Minto Lakes, Fairbanks, Tetlin), Kusko- kwim River (Napaskiak, Bethel), interior Nushagak Peninsula, Lake Iliamna, and Anchorage. Intergrades with rubiginosa at Napaskiak (lower Kuskokwim River), Nushagak, New Iliamna, and near Anchor age. Migrant in southwestern Oregon (see Remarks). Etymology. —For Richard C. Banks, col- league and friend, in recognition of his con- tributions to ornithology. Specimens examined. — Alaska (21); Yu- kon Territory (34). Remarks. — Oberholser (1897) character- ized a specimen from Nushagak and one from Yukon River as slightly more yellow VOLUME 107, NUMBER 1 above than specimens from Kodiak Island, and Raveling & Warner (1978) remarked that the population from Arctic interior Alaska may represent an unnamed subspe- cies. Dorsal color of breeding males ranges from brighter and more yellow in specimens from the Porcupine and upper Yukon rivers to duller and greener in birds from localities west and south of Fairbanks, Alaska. The duller and greener birds are still more yellow above than amnicola and neighboring sub- species. Males from north of the Brooks Range are still duller. Males from the type locality and Lake Iliamna have darker and more prominent ventral streaks than from elsewhere in the range of banksi. Specimens from Lake Iliamna that I identified as in- tergrades between banksi and rubiginosa were identified as intergrades between am- nicola and rubiginosa by Williamson (see Williamson & Peyton 1962). A specimen (UAM) from Tvativak Bay, Nushagak Peninsula, Alaska, collected 16 June, if actually breeding, is at the south- western limit of the breeding range of bank- si. The northern limit of banksi is based on specimens from north of the Brooks Range at Umiat (West & White 1966), Anaktuvuk (Irving 1960), the upper Sheenjek River (Kessel & Schaller 1960), and middle No- atak River (Kessel & Gibson 1978). Spec- imens from those localities are duller than most examples of banksi collected south of the Brooks Range. Additional specimens are needed to better determine the breeding range of banksi. A migrant male (USNM 592840) from Brownsboro, Oregon, collected 13 May 1920, resembles breeding specimens of banksi in dorsal and ventral color. Dendroica petechia parkesi, new subspecies Holotype. —CM 129401, adult male, Richards Island, Mackenzie Delta, North- west Territories, Canada, collected 4 June 1942 by Arthur C. Twomey (original num- ber 9864). 31 Subspecific characters. —Males darker green above than all other subspecies in the aestiva group. Compared with banksi, males much darker and greener above; chestnut streaks less prominent. Males near rubigi- nosa but greener above; forehead averaging more yellow. Compared to amnicola, males greener above; average duller yellow below; chestnut streaks average darker. Females greener than banksi and amnicola; average slightly paler yellow below in fall plumage. Bill, in males shorter (6.9-7.9, mean 7.55, n= 24) than rubiginosa (7.6-8.6, mean 8.29, n= 10), and similar to banksi (7.5—8.3, mean 7.84, n = 11). Distribution. — Breeds in Northwest Ter- ritories from Mackenzie River Delta to Arc- tic Red River, Ft. Simpson, Ft. Providence, Ft. McPherson, Ft. Norman, Rae, Reliance, Ft. Resolution, Hay River, and provision- ally, northeastern Manitoba (Ft. York). In- tergrades (?) with amnicola at Ft. Provi- dence and Hay River south of Great Slave Lake. Parkes (1968) identified a specimen from Massachusetts with the Mackenzie Delta population (= parkesi). Specimens examined.—Northwest Ter- ritories (21); Manitoba (6). Etymology. —For Kenneth C. Parkes, who first discussed the distinction of the new subspecies. Remarks. —Parkes (1968) commented that birds from the interior of Alaska to the west coast of Hudson Bay represented an unnamed subspecies. Ramos & Warner (1980) believed that the range given by Parkes (1968) included two unnamed sub- species, one from Alaska (= banksi) and the other from Northwest Territories and prai- rie provinces. There is some geographic variation in parkesi, with a reduction of green color from northwest to southeast. Specimens from the Mackenzie River Delta are the darkest above, with about one-half having greenish foreheads. Males from Ft. Simpson are slightly more yellow above, and females are paler above than specimens from the Mac- kenzie River Delta. Two of five specimens 32 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON from Ft. Providence resemble amnicola in dorsal color; specimens from Hay River, south of Ft. Providence, are, however, greener above. Males from the west coast of Hudson Bay are slightly more yellow above but are definitely greener above than amnicola. Specimens from the northern prairie provinces are clearly amnicola. Dendroica petechia amnicola Batchelder Dendroica aestiva amnicola Batchelder, 1918:82 (Curslet, Newfoundland). MCZ Subspecific characters. —Males near park- esi but more yellow and paler above; usually brighter yellow below. Compared to banksi, males darker (greener) above. Compared with aestiva, males darker and greener above, usually with greener foreheads. Fe- males more yellow above than parkesi and greener above than aestiva. Males in fall plumage slightly paler above than those of rubiginosa and parkesi. Distribution. —Breeds from northeastern British Columbia to central prairie prov- inces, Manitoba (Norway House), central Ontario, southeast to southern Quebec, New Brunswick, Newfoundland, and central Labrador. Intergrades (?; see above) with parkesi in southern District of Mackenzie (Ft. Providence). Intergrades with aestiva from central-northern Alberta to southern Manitoba, southern Quebec, and Nova Sco- tia. Specimens examined.—Northwest Ter- ritories (7); British Columbia (5); Alberta (30); Saskatchewan (11); Manitoba (8); On- tario (43); Quebec (21); and Newfoundland (30). Remarks. —Originally proposed for pop- ulations in Newfoundland, the name am- nicola was generally considered a synonym of aestiva until Oberholser (1938), followed by the A.O.U. (1944), recognized amnicola for the subspecies occuring in most of Can- ada. Contrary to Oberholser (1974), the original description of aestiva was based on the pale southeastern subspecies, not the dark northeastern population now in am- nicola (Browning 1990b). Oberholser (1974) also believed that carolinensis Latham, a name synonymized with aestiva by Hell- mayr (1935), was a synonym of the northern subspecies, but carolinensis is indetermi- nate; it is doubtfully an example of D. pe- techia. Geographic variation in back color of amnicola ranges from the brightest and most yellowish specimens from Newfoundland to greener birds from northeastern British Co- lumbia, southern Mackenzie, and northern Alberta. Differences between the colors of the forehead and back of males vary geo- graphically; both characters are greenish in 13% of the specimens from Newfoundland (n= 15), 17% of the birds from the southern Hudson Bay region, and about 25% of males from the prairie provinces. Raveling & Warner (1978) concluded that 90% of the males from the southern parts of the prairie provinces were “readily distinguishable” in back and head color from all other samples, and referred the prairie birds to aestiva. I concur. Dendroica petechia aestiva (Gmelin) Motacilla aestiva Gmelin, 1789:996 (in Gu- jana, aestate in Canada = Québec, Qué- bec). Sylvia flava Viellot, 1809:31, pl. 87, (on migration in the United States = New York, New York; fide Oberholser 1974: 1000) Sylvia childrenii Audubon, 1831:180, (near Jackson, Mississippi). ?Sylvia rathbonia Audubon, 1831:333, (Gulf states, Mississippi, Louisiana, or Tennessee). Dendroica aestiva ineditus J. C. Phillips, 1911:85 (Matamoros [Tamaulipas, Mexico)]) fide A. R. Phillips (an litt.) and Ramos & Warner (1980). MCZ Subspecific characters. — Both sexes, in all plumages, more intensely bright yellow be- low than other subspecies in the aestiva group. Compared with amnicola, both sexes paler (less greenish, more yellowish) above. VOLUME 107, NUMBER 1 Compared to morcomi, both sexes more yellow above and below; males brighter yel- low below. Compared with sonorana, both sexes greener above and brighter (not pale) yellow below; chestnut streaks more prom- inent. Paler males in fall plumage brighter with more yellowish rumps and crowns than other northern subspecies and morcomi. Distribution. —Breeds from south-central Alberta to central Saskatchewan and south- ern Manitoba, southern Quebec, Prince Ed- ward Island, Nova Scotia; east of Rocky Mountains from Montana to Colorado east to Kansas, central Oklahoma, all but west- ern Texas, northern Arkansas, central Al- abama, central Georgia, and central South Carolina. Intergrades with amnicola in cen- tral Alberta, southern Manitoba, southern Quebec, and Nova Scotia. Specimens examined. — Alberta (7); Sas- katchewan (6); Manitoba (21); Nova Scotia (31); Montana and Wyoming (16); Colora- do (10); North and South Dakota (18); Ne- braska (49); Oklahoma (8); Minnesota (41); New England (42); New York (54); New Jer- sey (11); Pennsylvania (11); Maryland and Virginia (25); Georgia and Carolinas (34); others (34). Remarks. —Oberholser (1974) believed the name aestiva was based on the darker northern subspecies (amnicola) and used the name flava Vieillot for the paler southern subspecies. However, the basis of the earlier name aestiva clearly applies to the pale pop- ulation (Browning 1990b). Based on Au- dubon’s plates and descriptions (the types are missing), the name childrenii, and most probably rathbonia, are synonyms of aes- tiva. The name inedita was based on spec- imens collected in August and September. Griscom & Crosby (1926) and Hellmayr (1935) recognized ineditus, but Miller et al. (1957) and Lowery & Monroe (1968) syn- onymized the name with morcomi. A. R. Phillips (in litt.), who has examined the type of ineditus, concluded that the holotype is a migrant example of aestiva, and that the species does not breed west of Matamoros (contra Griscom & Crosby 1926). Paratypes 33 (MCZ) from the original series of ineditus resemble specimens of aestiva. Dorsally dark and greenish birds breed in southeastern Canada to Pennsylvania. There is a gradual north to south cline in reduction of dorsal green color and increased amount of yellow on the foreheads and backs of birds from Virginia to Georgia. Specimens from Nebraska, Kansas, and western Oklahoma are slightly brighter and less greenish above than specimens from the northeastern part of the breeding range and represent the end of a cline in coloration. A similar cline is represented by specimens from the north- east west to the prairie provinces and states. Specimens from Georgia resemble birds from Nebraska in reduction of green in their upper parts. Birds from Nebraska, collec- tively, are generally more intensely bright yellow above and have more yellowish rumps than do specimens of morcomi, so- norana, and aestiva from northeastern pop- ulations (Browning in Wetmore et al. 1984). Such specimens resemble the “sonorana”’ type characterized by Sutton (1967). A grad- ual west to east cline is also apparent from a large series of specimens collected at sev- eral localities across Nebraska, with speci- mens from northeastern Nebraska being less green above then birds from northwestern Nebraska. The Nebraska series, collective- ly, is slightly more yellowish above, with more yellow than green on the head, and resemble specimens from the southern prai- rie provinces in Canada and western Min- nesota. Specimens from Nebraska resemble birds from Georgia in dorsal color. Geo- graphic variation in aestiva, though consid- erable, is clinal and thus not diagnostic in recognizing additional subspecies in its range. Johnston (1964) identified specimens from western Kansas as morcomi but spec- imens from western Kansas and eastern Colorado represent the end ofa cline toward morcomi, with males being more greenish above and less yellow on the forehead than birds from other localities of the prairie states. Specimens of aestiva from central 34 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Montana and Wyoming are also slightly paler and less greenish above than are those from the northeastern United States. Dendroica petechia morcomi Coale Dendroica aestiva morcomi Coale, 1887:82 (Fort Bridger, Utah [= Fort Bridger (Camp Scott), Uinta Co., Wyoming) fide Deig- nan 1961]). (USNM) Subspecific characters. — Males near west- erm populations of aestiva but greener above; duller yellow below. Compared to brewsteri, males darker and greener on back and rump; chestnut streaks average wider. Compared to sonorana, males greener above; ventral yellow with a greenish cast. Chestnut streaks average narrower than parkesi, yukonensis, and amnicola. Ventral yellow in males usu- ally paler than aestiva and with a greenish cast compared to sonorana. Females dor- sally paler yellow than aestiva, darker (less yellow) than sonorana, and less green than aestiva; ventrally brighter yellow than sono- rana and rubiginosa. Specimens in fall plumage dorsally intermediate between am- nicola and aestiva but nearer the latter; greener above than sonorana. Distribution. —Breeds from interior Brit- ish Columbia to eastern Washington, west- ern Montana south to eastern California, Nevada (except extreme southern), north- ern Arizona, central New Mexico, and northwestern Texas. Intergrades with rubi- ginosa at Bella Coola, Vancouver Island, and adjacent mainland, western British Co- lumbia; intergrades with sonorana in the Southwest (see beyond). Specimens examined. — British Columbia (59); Washington (9); Oregon (30); Califor- nia (7); Idaho/Wyoming (15); Nevada (5); Utah (18); northern Arizona and central New Mexico (20). Remarks. —Specimens of morcomi from the northern Rocky Mountains, Utah, and the interior of British Columbia average slightly brighter yellow above than morcomi from elsewhere, but are greener above than brewsteri. Males from Vancouver Island and adjacent mainland (e.g., city of Vancouver) of British Columbia average slightly duller and greener on the back and forehead than other specimens of morcomi. The average dorsal color of males from Bella Coola, Brit- ish Columbia, is intermediate in color be- tween specimens from the Vancouver Is- land region and morcomi from central British Columbia. Females from most of British Columbia resemble others from the range of morcomi. I conclude that the pop- ulations from the Vancouver Island are in- termediate in dorsal color between morcomi and rubiginosa but are closer to the latter, while birds from Bella Coola are likewise intermediate but are closer to morcomi. Males from the northern Great Basin (e.g., Utah) are slightly more yellow (less green- ish) above than most specimens of morcomi from the same latitudes or from higher el- evations (e.g., east slopes of Sierra Nevada Mountains). Geographic variation is gen- erally clinal from north to south, with darker and greener birds breeding to the north (Phillips et al. 1964). The zone of intergra- dation between morcomi and sonorana 1s wide and identifying specimens to either subspecies or as intergrades between them have been interpreted in different ways (see Behle 1948, Oberholser 1974, Behle 1985). Dendroica aestiva brewsteri Grinnell Dendroica aestiva brewsteri Grinnell, 1903: 72 (Palo Alto, California). (MVZ) Subspecific characters. —Males near mor- comi but definitely more yellow on the back; rump and forehead average more yellow; edges of secondaries and tertials more yel- low (less greenish); chestnut streaks aver- aging thinner and sparser. Compared with western populations of aestiva, males greener above; chestnut streaks usually thinner and sparser. Many females resemble morcomi, but brightest yellow individuals more yel- low than bright yellow examples of mor- comi. Adult males in fall plumage, com- VOLUME 107, NUMBER 1 pared to morcomi, usually with ventral streaks absent or narrower; some fall males average more yellow above. Distribution. —Breeds from western Washington to western Oregon, California west of the Cascade and Sierra Nevada rang- es, and northwestern Baja California. Win- ters from Baja California to Nicaragua. Specimens examined.— Washington (4); Oregon (10); California (34). Remarks. —The subspecies brewsteri was recognized for many years while the name morcomi remained a synonym of aestiva. About the time morcomi became recog- nized as a subspecies distinct from aestiva, the name brewsteri became a synonym of morcomi. Because brewsteri is recognizable, and because the history of the names brew- steri and morcomi are closely related, a re- view of the usage of the two names follows. When Grinnell (1903) described brew- steri, he also synonymized morcomi with aestiva, stating that the holotype of mor- comi was an extreme example of aestiva. He characterized brewsteri as smaller in size, less brightly yellow, and with narrower chestnut streaks than in aestiva, and gave the breeding range of brewsteri as west of the Cascade and Sierra Nevada mountains from Oregon to southern California. Grin- nell’s comparative material of “‘aestiva” consisted of specimens from the Rocky Mountains and birds from the northeastern United States. The A.O.U. (1910, 1931) recognized brewsteri and included the pop- ulations from the Rocky Mountains (= mor- comi) in the range of aestiva. Van Rossem (1931) and Dickey & van Rossem (1938) recognized both brewsteri and morcomi, characterized brewsteri of the Pacific coast as larger (contra Grinnell 1903) and having narrower breast streaks than morcomi, and gave the range of morcomi as from the east- ern slopes of the Cascade and Sierra moun- tains to the Rocky Mountains. Gabrielson & Jewett (1940), without providing details, synonymized both brewsteri and morcomi with aestiva. Twomey (1942) reported that wing chord of brewsteri is not a useful sub- 35 specific character, but, on the basis of color, recognized brewsteri as the subspecies breeding in the Uinta Basin, Utah. Grinnell & Miller (1944) reported that specimens from east of the Sierra Nevada north of Mono Lake are intermediate between brewsteri and morcomi. The A.O.U. (1944) recognized morcomi, following Dickey & van Rossem (1938). However, Behle (1948: 77-78) reported that specimens from Cali- fornia and Utah are similar in size, dorsal color, and widths of chestnut streaks. He concluded that the topotypes of brewsteri and morcomi “are similar in their charac- ters” and that “‘brewsteri appears to be [em- phasis mine] a synonym of morcomi.”’ Al- drich (in Jewett et al. 1953) and the A.O.U. (1954), both of whom cited Behle (1948), synonymized brewsteri with morcomi. The synonymy of brewsteri was thus based on comparisons of brewsteri with more yel- lowish specimens of morcomi typical of Utah and southwestern Wyoming. The tax- onomic conclusion by A.O.U. (1954) did not account for geographic variation in morcomi, and the fact that the breeding range of brewsteri is isolated from the more yel- lowish examples of morcomi by darker and less yellowish populations of morcomi from the eastern slopes of the Cascade and Sierra Nevada mountains. Grinnell & Miller (1944) included north- eastern California as part of the breeding range of brewsteri. I did not examine spec- imens from that region, but specimens from adjacent Oregon belong to morcomi. Dendroica petechia sonorana Brewster Dendroica aestiva sonorana Brewster, 1888: 137 (Opusura [= Moctezuma], Sonora, Mexico). MCZ Dendroica petechia hypochlora Oberhol- ser, 1974:737 (3 mi north of Fort Whipple, near Prescott, Arizona). (USNM) Subspecific characters. —Compared with other subspecies in the aestiva group, both 36 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON sexes paler yellow above and below; males with dorsal chestnut streaks usually prom- inent; males with ventral chestnut streaks narrow, paler, and less prominent. Com- pared with dugesi, males with ventral chest- nut streaks averaging more prominent; shorter wings (61.0-65.7, mean 63.1 (n = 14) in sonorana and 64.5-73.1, mean 68.6 (n = 10) in dugesi. Fall plumage greenish above but more yellowish above and more pure yellow below than morcomi. Distribution. —Breeds from southeastern California, extreme southern Nevada, cen- tral Arizona, southern New Mexico, and western Texas south to northeastern Baja California, interior Nayarit, and Zacatecas. Intergrades with morcomi in northern and northeastern Arizona, Utah, New Mexico, and western Colorado; intergrades with morcomi and aestiva in southwestern Okla- homa. Specimens examined. —California (6); Arizona (132); New Mexico (58); Colorado (7); Texas (24). Remarks. — Oberholser (1974) named the populations from the area of intergradation with morcomi (above) hypochlora. The ho- lotype of hypochlora most resembles spec- imens of sonorana (Browning 1978, 1990a). Dendroica petechia dugesi Coale Dendroica dugesi Coale, 1887:83 (Moro Leon [= Moroleon], Guanajuato, Mexi- co). (USNM) Subspecific characters. —Males nearest sonorana but more greenish above; wings longer (see under sonorana). Females paler and grayer above than sonorana. Distribution. —Breeds in Central Plateau of Mexico from southern San Luis Potosi to Hidalgo, Guanajuato, Michoacan, Guer- rero, Morelos, and Puebla. Winters from Morelos to Puebla and Tlaxcala; reported in southern Veracruz (Ramos & Warner 1980). Specimens examined.—Michoacan (6); Puebla (5); Durango (1); Guanajuato (1); Tlaxcala (1); Morelos (3); México (3). Remarks. —The southern breeding range of sonorana and northern breeding range of dugesi are poorly known. Present informa- tion suggests the two ranges are disjunct, but sonorana and dugesi may come in contact somewhere in Zacatecas or San Luis Potosi. Summary of D. p. aestiva group Generally, males in the aestiva group from coastal Alaska and northern interior pop- ulations are greener (less yellow) above than birds from elsewhere. Males from the south- western range of the aestiva group are paler yellow and much less green above than oth- er populations. Ventral color is generally brighter and more purely yellow in south- eastern populations, paler in southwestern birds, and slightly tinged with green in northern populations. More specifically, the northern popula- tions east of the Rocky Mountains from Canada (parkesi and amnicola) are darker and greener (less yellow) above than adja- cent populations; parkesi is greener above than amnicola. Variation in dorsal color of birds from the eastern United States (aes- tiva) is clinal, with darker birds from the northeast and brighter and more yellowish birds from the southeast and prairie region. Except for the dark populations of coastal Alaska and British Columbia (rubiginosa), birds from west of the Rocky Mountains are generally darker and greener above in northern populations and paler and more yellow in southern populations. Birds from interior Alaska and Yukon Territory (bank- si) are more yellow above than other north- erm populations and are darker and more greenish than birds of the northern inter- mountain United States and interior British Columbia (morcomi). Populations of west- ern Washington to California (brewsteri) are more yellow above than morcomi. South- western populations (sonorana) are paler and more pure yellow (less greenish) above than VOLUME 107, NUMBER 1 the northern populations, and are smaller and slightly less greenish above than inte- rior Mexican populations (duges!). Dendroica p. petechia group Populations of the petechia group differ from birds of the aestiva group in having a generally well-defined chestnut crown and rounder wings. Birds of the petechia group differ from those of the erithachorides group in having the chestnut restricted to the crown, with the exception of the chestnut hooded birds from Martinique. Birds from Martinique are more similar in size to birds of the petechia group than to the erithachori- des group. Birds of petechia group are usu- ally found in mangroves. Dendroica petechia rufivertex Ridgway Dendroica petechia rufivertex Ridgway, 1885:21 (Cozumel Island, Yucatan [= Quintana Roo]). (USNM) Subspecific characters. —Males near gundlachi above but chestnut streaks more extensive and darker. Females less heavily streaked below than gundlachi. Distribution. —Resident on Cozumel Is- land, Quintana Roo, Mexico. Specimens examined.—Cozumel Island (18). Remarks. — The whitish plumages typical of most of the northern subspecies of the petechia group are apparently absent in this subspecies. Dendroica petechia armouri Greenway Dendroica petechia armouri Greenway, 1933:63 [= 68] (Old Providence Island). MCZ Subspecific characters. —Males nearest flavida but forehead and throat yellow (not chestnut); ventral chestnut streaks more ex- tensive. Compared with rufivertex, both sexes darker and less yellow above. 37 Distribution. —Isla Providencia, western Caribbean. Specimens examined. —I\sla Providencia (5). Remarks.—The only female of armouri compared is similar to females of rufivertex and flavida. Bond (1950) reported armouri as the rarest resident on the island. Birds were not found there by Russell et al. (1979) and Tye & Tye (1991). Dendroica petechia flavida Cory Dendroica flavida Cory, 1887:179 (St. An- drew Island, Caribbean). FM Subspecific characters. —Males nearest rufivertex but chestnut on crown reduced and ventral streaks wider. Compared to ar- mouri, forehead and throat chestnut (not yellow) and rump greener. Females darker green than males of flavida, and some heavi- ly marked with ventral chestnut streaks. Distribution. —Isla Andrés, western Ca- ribbean. Specimens examined. —Isla Andrés (17). Dendroica petechia eoa Gosse Sylvicola eoa Gosse, 1847:158 (Crab Pond, Jamaica). BM Dendroeca petechia e. jamaicensis Sun- devall, 1870:608 (near Spanishtown, Jamaica). (USNM) Dendroica auricapilla Ridgway, 1888:572 (Grand Cayman). (USNM) Subspecific characters. —Males near gundlachi but slightly brighter (more yel- low), especially above in fall plumage, with chestnut crown patch more extensive. Com- pared with albicollis, males slightly greener and darker above. Females greener above than flaviceps. Females in fall plumage dull- er above and below than gundlachi. Distribution. —Jamaica and Cayman Is- lands. Specimens examined. —Jamaica (31), Little Cayman (3), Grand Cayman Island Ga): 38 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Remarks.—Until Peters’ (1927) study, birds from Jamaica were considered to rep- resent nominate petechia. I follow Hellmayr (1935) and others in synonymizing aurica- pilla with eoa. I disagree with Buden (1979) who synonymized eoa with albicollis. Dendroica petechia gundlachi Baird Dendroica gundlachi Baird, 1865:197 (Cuba). (USNM) Subspecific characters. —Both sexes near flaviceps but duller and greener above; rump greenish (not yellowish); less brilliant yellow (more greenish) below; chestnut crown usu- ally more extensive. Compared with eoa, both sexes duller and greener above; duller yellow below; males with chestnut crown usually less extensive. Compared with so- laris and albicollis, males greener above. Distribution.—Cuba, Isle of Pines, and mangroves of southern Florida from Flor- ida Bay Island, Virginia Key near Miami, and Florida Keys (Stevenson & Anderson 1993). Specimens examined. —Cuba (99); Isle of Pines (21); Florida (2). Remarks. —This subspecies was first re- ported as breeding in southern Florida by Greene (1942) who collected two specimens (USNM) near Key West in mid-July. The two specimens have tarsi of 20.3 and 21.8 mm, and are more similar to gundlachi than to flaviceps in dorsal color. Dendroica petechia flaviceps Chapman Dendroica petechia flaviceps Chapman, 1892: 310 (Rum Cay, Bahamas). (AMNH) Subspecific characters. —Both sexes near gundlachi but more yellow above; rump yel- lowish; more yellowish (less greenish) be- low. Topotypical flaviceps are brighter (lacking a greenish cast) below than gundla- chi; specimens from elsewhere in the Ba- hamas are more similar to gundlachi. Com- pared with solaris and albicollis, males duller yellow below. Fall and spring plumages def- initely more yellowish (less greenish) above than gundlachi; most yellowish females from Cuba are still greener above than flaviceps. Distribution. —Bahama Islands. Specimens examined. — Various Bahama Islands (103). Remarks.—Peters (1927) characterized flaviceps as similar to gundlachi, but with the chestnut of the crown less extensive and the tarsi longer. Hellmayr (1935) com- mented that specimens of flaviceps are gen- erally brighter in color and the wings are more rounded than in gundlachi. Bond (1942) synonymized flaviceps with gundla- chi stating that the characters “such as more rounded wing, longer tarsus and brighter coloration did not hold.” Others (e.g., Low- ery & Monroe 1968, Buden 1979) followed Bond (1942). Wiedenfeld (1991) stated that birds with the longest tarsi are from the northern Ba- hamas. Measurements of tarsi of flaviceps (n = 17) in this study range from 20.2 to 23.6, with a mean of 21.6, and of gundlachi from Cuba (n = 14) range from 19.5 to 21.1, with a mean of 20.3. Individual measure- ments of flaviceps overlap 30% of those of gundlachi; individuals of gundlachi overlap 50% of those of flaviceps. Dendroica petechia albicollis (Gmelin) Motacilla albicollis Gmelin, 1789:983 (in insula S. Dominici [= Santo Domingo)). Subspecific characters. —Both sexes with more yellowish (less dull and greenish) backs and more yellow crowns than eoa and duller and more greenish (less yellowish) above than solaris. Males brighter yellow below than flaviceps and more yellowish above than gundlachi. Females with some pale chestnut on foreheads and grayer above than eoa and solaris. Distribution. —Hispaniola, Vache, and Tortue islands. Specimens. — Hispaniola (27). Remarks.—Buden (1979) synonymized eoa with albicollis. VOLUME 107, NUMBER 1 Dendroica petechia chlora, new subspecies Holotype. —USNM 280265, adult male, Tororu Island, Siete Hermanos islands, Domican Republic, collected 31 January 1929 by A. J. Poole and W. M. Perrygo (original no. 165). Subspecific characters. —Males nearest al- bicollis but back and crown distinctly darker green; edges of secondaries and primaries greener (less yellow); chestnut on crown darker. Compared with solaris, males much darker above and less yellow below than solaris. Compared with gundlachi, males slightly greener above and usually paler yel- low below. One of two females darker and less grayish above than albicollis (the other specimen cannot be distinguished from al- bicollis). Distribution. —Siete Hermanos islands, off Hispaniola. Specimens examined. —Siete Hermanos islands (8). Etymology. —Greek, chloros, for the greener color of the plumage. Remarks.—A male from Fort Liberty, mainland Haiti, a locality about 10 miles SSW of the type locality of ch/ora, resembles albicollis in color. Dendroica petechia solaris Wetmore Dendroica petechia solaris Wetmore, 1929:1 (Z’Etriotes, Ile de la Gonave [in Gulf of Gonaives], Republic of Haiti, Hispanio- la). (USNM) Subspecific characters. — Males nearest al- bicollis but brighter and more yellowish above, and on edges of primaries and sec- ondaries. Compared with chlora, males pal- er (less dark green) than those of chlora. Females average paler and more yellowish above than albicollis. Distribution. —Resident on Gonave and Petite Gonave islands off Haiti. Specimens examined. —Gonave (27); Pe- tite Gonave (4). 39 Remarks. — Although solaris was listed as a distinct subspecies by Bond (1930) and by Hellmayr (1935), Lowery & Monroe (1968) synonymized the name with albicollis. Dendroica petechia bartholemica Sundevall Dendroeca petechia bartholemica Sunde- vall, 1870:607 (St. Bartholemew). Royal Natural History Museum, Stockholm (fide Gydlenstolpe 1926) Dendroeca petechia cruciana Sundevall, 1870:608 (St. Croix). Cambridge Univ. (fide Hellmayr 1935) Subspecific characters.—Near albicollis, but both sexes average brighter yellow be- low; males with chestnut on the head and breast more extensive. Compared with me- lanoptera, males with crown less distinctly capped with chestnut. Distribution. —Puerto Rico, islands east of Puerto Rico, Virgin Islands, northern Lesser Antilles from Anguilla to St. Martin, St. Bartholomew, Barbuda, St. Eustatius, Nevis, St. Kitts, Antigua, and Montserrat. Specimens examined. — Puerto Rico (97); Virgin Islands (17); St. Martin (2); Barbuda (4); St. Kitts and Nevis (8); Antigua (17); Montserrat (2). Remarks. — According to Peters (1927), specimens that he identified as cruciana from St. Croix were “‘quite distinctive” from those he identified as bartholemica from St. Thomas. However, Wetmore (1927:105) concluded that the range of cruciana ex- tends east to Antigua, and that cruciana and bartholemica “are closely similar” in color. I found that overlap in color between pop- ulations is extensive, with some specimens from the northern Lesser Antilles (““bartho- lemica’’) being brighter below and having more chestnut on the crown than specimens from Puerto Rico (“‘cruciana’’). The only female from Montserrat examined has a yel- lowish rump. Use of the name bartholemica for this subspecies follows Ridgway (1903) who acted as the first revisor. 40 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Dendroica petechia melanoptera Lawrence Dendroeca petechia melanoptera Lawrence, 1879:453 (Guadeloupe). (USNM) Subspecific characters. —Males nearest bartholemica but with chestnut crown more distinct. Females average slightly more yel- lowish on rump than bartholemica. Distribution. —Central Lesser Antilles: Guadeloupe, Désirade, Marie Galante, Iles des Saintes, and Dominica. Specimens examined. —Guadeloupe (9); Dominica (7); Marie Galante (2). Dendroica petechia ruficapilla (Gmelin) Motacilla ruficapilla Gmelin, 1789:490 (in Martinica = Martinique). Dendroica rufigula Baird, 1865:204. (Martinique). ANSP Dendroeca granadensis Sharpe, 1885:284 (Colombia = Martinique). (BM) Subspecific characters. — Differs from An- tillean subspecies by having a chestnut hood, thus resembling subspecies in the erithacho- rides group. Birds from Martinique are sim- ilar in size to subspecies in the petechia group (Wiedenfeld 1991). Distribution. —Martinique, Lesser Antil- les. Specimens examined. — Martinique (10). Remarks. — According to Peters (1927), the name ruficapilla was based on females from Guadeloupe that were originally char- acterized as having a yellow throat, whereas birds from Martinique (= “‘rufigula”’ in Pe- ters, footnote) have throats that are “gen- erally clouded with rufous.” Hellmayr (1935) concluded that there are no valid grounds for Peters’ treatment and most subsequent authors have followed Hellmayr. Sharpe (1885) referred to three specimens listed by Sclater (1862:32) in his indeter- minable description of granadensis. Hell- mayr (1935), who examined one of the spec- imens, believed that the name granadensis was probably a synonym of ruficapilla. Ap- parently on the recommendation of A. Wet- more (notes on file, USNM), Warren & Har- rison (1971) listed one of the three specimens as a syntype of granadensis. Based on mea- surements and description of plumage (P. R. Colston, in litt.), the specimen listed by Warren & Harrison (1971) is identifiable as ruficapilla. The characters of a second spec- imen (BM), examined by Browning and S. L. Olson, agree with Colston’s characteriza- tions of the specimen listed by Warren & Harrison (1971). A third specimen (BM) is similar in size and color to erithachorides. Because the three syntypes of granadensis represent more than one subspecies, I des- ignate BM 1884.5.5.15.356, the same spec- imen listed by Warren & Harrison (1971), as the lectotype of Dendroeca granadensis Sharpe. Dendroica petechia babad Bond Dendroica petechia babad Bond, 1927:571 (St. Lucia, British West Indies). (ANSP) Subspecific characters. —Males nearest melanoptera but slightly more yellowish (less greenish) below; slightly darker chestnut cap. Chestnut areas more extensive than a/siosa and much paler than nominate petechia. Distribution. —St. Lucia, Lesser Antilles. Specimens examined. —St. Lucia (4). Remarks. —\ agree with Bond (1936) who stated that babad is “barely distinguish- able” from birds from Dominica (me/anop- tera). The purported differences in crown color (Bond 1927; Hellmayr 1935) between the adjacent subspecies alsiosa and mela- noptera are exaggerated. Dendroica petechia petechia (Linnaeus) Motacilla petechia Linnaeus, 1766:234 (in America septentrionali = Barbados, Less- er Antilles). Dendroica capitalis Lawrence, 1868:359 (Barbados, Lesser Antilles). (AMNH) Dendroeca petechia c. barbadensis Sun- devall:608 (Barbados). (USNM) VOLUME 107, NUMBER 1 Subspecific characters. —Males nearest rufopileata but greener (less yellowish) be- low; chestnut cap slightly darker; darker above than either rufopileata or obscura. Chestnut areas much darker than alsiosa. Females paler (less blackish) below than ru- fopileata; more yellowish (less greenish) above than babad, and greener above than alsiosa. Distribution. —Barbados, Lesser Antilles. Specimens examined. — Barbados (12). Remarks. —The name petechia was used for the Jamaican populations until Peters (1927) determined that Edwards’ plate is of a bird from the Barbados population. Dendroica petechia alsiosa Peters Dendroica petechia alsiosa Peters, 1926:41 (Prune Island, east of Union Island, Grenadines). (MCZ) Subspecific characters. —Males near me- lanoptera, but more yellow (less greenish) above; yellow (not greenish) immediately above the bill and cheeks. Compared with babad and nominate petechia, males with slightly paler ventral chestnut streaks. Crown color paler than in nominate petechia. Fe- males more yellowish (less greenish) above than babad. Distribution. —Grenadine islands, Lesser Antilles. Specimens examined. —Prune Island (5); Union and Mayero islands (2). Remarks. — Males from Prune Island have slightly darker crowns than the two males from Union and Mayero islands. Dendroica petechia rufopileata Ridgway Dendroica rufopileata Ridgway, 1884:173 (Curacao). (USNM) Subspecific characters.—Males near ob- scura but yellowish-green (not dark green) above; chestnut areas paler, especially ven- tral streaks. Chestnut crown darker than all previously listed subspecies of petechia group, except nominate petechia. Com- 41 pared with aurifrons, males darker above; chestnut streaks slightly darker and wider. Distribution. — Aruba, Curacao, and Bon- aire; Blanquilla, Margarita, and Islas Los Testigos, off northern Venezuela. Specimens examined. — Aruba (13); Cu- racao (31); Bonaire (13); Blanquilla (16); Is- las Los Testigos (11). Remarks. — Birds in the eastern and west- ern parts of the range of rufopileata are sep- arated by a darker subspecies, obscura (see below), and the taxonomy of the complex has varied historically. When Ridgway (1884) named rufopileata, birds from the islands off Venezuela were known only from Curacao. Hartert (1893) discovered similar birds on Aruba and Bonaire, and Lowe (1907) extended the range of rufopileata to include Isla Blanquilla. Cory (1909) named obscura from Isla Los Roques and, although he noted differences between the now dis- junct eastern and western parts of the range of rufopileata, he considered these too mi- nor for recognition of a third subspecies. Peters (1927) essentially followed Cory (1909) and included specimens from Isla Las Aves in obscura. Although Hellmayr (1935) synonymized obscura with rufopilea- ta, he characterized birds from the range of obscura as dark, and birds from Blanquilla, Testigos, and Tortuga as brighter yellow above, with wider yellow margins on the wings, than in birds from western islands. Phelps & Phelps (1950) followed Hellmayr (1935), but they (Phelps & Phelps 1951) lat- er recognized obscura, adding that speci- mens from Tortuga are intermediate be- tween rufopileata and aurifrons from islands to the south (see below) but closer to the latter. Voous (1957), who recognized both rufopileata and obscura, reported that 2 of 8 specimens from Aruba have faint streaks on the throat as in cienagae (see below). Voous (1957) also, followed by Phelps & Phelps (1959) and Lowery & Monroe (1968), included specimens from Isla La Orchila (formerly in rufopileata) in obscura. The two populations of rufopileata are 42 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON generally similar, but males from the west- ern islands of Isla Blanquilla and Isla Los Testigos have slightly darker ventral chest- nut streaks than 66% of the specimens from Bonaire, Curacao, and Aruba. The chestnut on the crown of the western population is darker than about 50% of the eastern birds, and about 25% of the eastern birds are more yellow on the rump than in western birds. I do not view these differences as sufficient for taxonomic recognition. Dendroica petechia obscura Cory Dendroica ruficapilla obscura Cory, 1909: 217 (isla Los Roques, Leeward Islands). (FM) Subspecific characters. —Near rufopileata but dark greenish (less yellowish) above, with chestnut area darker, especially ventral streaks. Distribution. —Islas Los Roques, Islas Las Aves, and Isla La Orchila, offnorthern Ven- ezuela. Specimens examined. —Isla Las Aves (7); Islas Los Roques (19); Isla La Orchila (1). Remarks. —See under rufopileata. Dendroica petechia aurifrons Phelps & Phelps, Jr. Dendroica petechia aurifrons Phelps & Phelps, Jr., 1950:21 (Puerto de La Cruz, Anzoategui, Venezuela). AMNH Subspecific characters. —Males more yel- lowish (less greenish) above and more yel- low (not chestnut) on the crown than ciena- gae and paraguanae; slightly paler below than cienagae. Compared with rufopileata, males slightly paler above; chestnut streaks narrower. Distribution. —Coast of Anzoategui and extreme western Sucre (at Cumana), north- central Venezuela, and islands immediately offshore, including Isla La Tortuga, Islas Las Tortuguillas, and Isla de Piritu. Specimens examined.—Isla La Tortuga (3); Venezuela mainland (2). Summary of D. p. petechia group Generally, the ventral chestnut streaks are more prominent in males from the western Caribbean islands than those from other populations. The chestnut cap ranges from pale in males from the Greater Antilles to dark in birds from the southern Lesser An- tilles and parts of islands off Venezuela. More specifically, the ventral chestnut streaks of birds from Isla Providencia (ar- mouri) are darker than those from Cozumel (rufivertex), and are more extensive than birds from Isla Andrés (flavida). Popula- tions from the northeastern Caribbean is- lands are generally greener above, with less extensive chestnut caps than birds from the southern islands. Birds from Cuba and southern Florida (gundlachi) are duller and greener above than those from the Bahama Islands (flaviceps) and are still greener above than birds from the main island of Hispan- iola (albicollis). Populations of the islands off Hispaniola from Gonave and Petite Gonave islands (solaris) are more yellow above than a/bicollis and birds from the Siete Hermanos islands (ch/ora) are darker above than albicollis and gundlachi. Birds from Puerto Rico and the northern Lesser Antil- les (bartholemica) are brighter yellow above and below, with more prominent chestnut caps, than albicollis. The chestnut cap is still more prominent in males from the central Lesser Antilles (melanoptera). The popu- lations of Martinique (ruficapilla) have a chestnut hood similar to that of the eri- thachorides group. Birds from St. Lucia (ba- bad) are more yellow above, with darker chestnut caps, than melanoptera. Males from Barbados (petechia) are still more yellow above, with still darker caps. The chestnut streaks and caps of birds from the Grena- dine islands (a/siosa) are paler than pete- chia. The caps of birds from the islands off northern Venezuela (rufopileata and ob- scura) are darker than petechia. Back color of rufopileata is paler than obscura. Males of the populations from western coastal Venezuela and adjacent islands (aurifrons) VOLUME 107, NUMBER 1 are paler above, with narrower chestnut streaks than rufopileata. Dendroica p. erithachorides group Subspecies in the erithachorides group dif- fer from those in the aestiva and petechia groups by having chestnut heads. Birds also differ from the aestiva group by having rounded wings. Members of the erithachori- des group are usually found in mangroves. Dendroica petechia oraria Parkes & Dickerman Dendroica petechia oraria Parkes & Dick- erman, 1967:87 (two miles south of Bue- na Vista (= about nine miles north of Tla- cotalpan), Veracruz, Mexico). (CM) Subspecific characters. —Males nearest bryanti but more greenish (less yellowish) above; averaging paler yellow below; aver- aging less heavily streaked with chestnut be- low, especially flanks. The only female ex- amined, from Veracruz, is paler and more yellow above and below than females of bryant. Distribution. —Coastal southern Tamau- lipas to Tabasco, Mexico. Intergrades with bryanti in eastern Tabasco and western Campeche. Specimens examined.—Tamaulipas (5); Veracruz (19); Tabasco (4); Campeche (4). Dendroica petechia bryanti Ridgway Dendroica vieillotii var. Bryanti Ridgway, 1873:605 (Belize, British Honduras = Be- lize, Belize) fide Parkes & Dickerman (1967). (USNM) Subspecific characters.—Males near or- aria but more yellowish (less greenish) above; brighter yellow below; chestnut streaks averaging narrower. Compared with erithachorides, both sexes greener above; males with narrower less prominent chest- nut streaks below. Distribution. —Caribbean coast from Yu- catan Peninsula, Mexico to Campeche and 43 Nicaragua. Possibly breeds on Isla Mujeres, Quintana Roo (Parkes & Dickerman 1967). Intergrades with oraria in eastern Tabasco and western Campeche. Specimens examined. —Quintana Roo (8); Yucatan (19); Belize (7); Honduras (3); Nic- aragua (9). Dendroica petechia erithachorides Baird Dendroica erihtachorides [sic] Baird, in Baird, Cassin, & Lawrence, 1858:283 (South America = Cartegena, Colombia, fide Hellmayr [1935]). (USNM) Dendroica Vieilloti Cassin, 1860:192 (Cartagena, Colombia). (USNM) [Dendroeca petechia| 1) panamensis ? Sundevall, 1870:609 (Cartagena, De- partment of Bolivar, Colombia). (USNM) Subspecific characters. —Males near bryanti but darker above; darker chestnut head; more conspicuous chestnut streaks. Compared with chrysendeta, males less yel- lowish below; chestnut bib more defined; less heavily streaked below. Females greener on back and rump (less yellow) than bryanti and chrysendeta. Distribution. —Caribbean coast from Costa Rica, locally, to Panama (Bocas del Toro and islands), and Caribbean coast of Colombia. Specimens examined.—Costa Rica (5); Panama (29); Colombia (7). Remarks. —Cherrie (1891:524) remarked that specimens from Limon, Costa Rica, have broader chestnut streaks than do northern examples of bryanti, and that the throat of the Limon birds is closer to that in specimens from Panama. I found that males from elsewhere along the coast of Costa Rica also have broader chestnut streaks and are slightly darker above than northern examples of bryanti. I agree with Stiles & Skutch (1989) in extending the northern range of erithachorides to Costa Rica. The populations from Limon, Costa Rica, and Bocas del Toro, Panama, are ap- 44 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON parently disjunct; there are no mangroves between the two localities. Specimens of both sexes from Escudo de Varaguas, Bocas del Toro, Panama, are duller above than specimens of erithachorides from the main- land and islands off Almirante, Panama, but I provisionally assign them to this subspe- cies. Dendroica petechia chrysendeta Wetmore Dendroica petechia chrysendeta Wetmore, 1946:52 (Laguna de Tucacas, Puerto Lopez, Guajira, Colombia). (USNM) Subspecific characters.—Males near eri- thachorides but yellow areas brighter; chest- nut of head and upper breast darker, more extensive on the head. Compared with par- aguanae, back more yellowish (less green); chestnut less extensive on the breast. Distribution. —Coastal Guajira Peninsu- la, northeastern Colombia and Bahia Por- tete, Puerto Lopez, Castilletes, and Para- guipa in western Venezuela. Specimens examined.—Guajira Penin- sula (6). Remarks.—Specimens from Bahia Por- tete are intermediate in color between eri- thachorides and chrysendeta (Wetmore 1946b). Variation in the dorsal yellow of chrysendeta and some populations of par- aguanae are similar (N. K. Klein, pers. comm.). Dendroica petechia paraguanae Phelps & Gilliard Dendroica petechia paraguanae Phelps & Gilliard, 1941:10 (La Boca, Adicora, Par- aguana Peninsula, Falcon, Venezuela). Subspecific characters. —Both sexes darker and greener above than chrysendeta. Distribution. — Paraguana Peninsula, Fal- con, northwestern Venezuela. Specimens examined.—Paraguana Pen- insula (8). Dendroica petechia cienagae Zimmer & Phelps Dendroica petechia cienagae Zimmer & Phelps, 1944:14 (La Cienaga, sea level, between Ocumare de la Costa and Turia- mo, Aragua, Venezuela). (AMNH) Subspecific characters. —Compared with aurifrons, males brighter yellow below; throat and forehead more chestnut; chest- nut streaks wider. Compared with chrysen- deta, males darker and greener above; chest- nut regions paler. Distribution.—Coast of Carabobo and Aragua, north-central Venezuela, and small islands off Falcon. Specimens examined. — Aragua (2). Dendroica petechia castaneiceps Ridgway Dendroica bryanti castaneiceps Ridgway, 1885:350 (La Paz, Lower California). (USNM) Dendroica erithachorides hueyi van Ros- sem, 1947:50 (San Ignacio Lagoon, Pa- cific coast of Baja California). SDMNH Subspecific characters. — Both sexes near- est rhizophorae but average slightly greener above; males with chestnut streaks less dense and narrower. Males with slightly longer tails (51.9-58.1, mean 55.9, n = 13) than rhi- zophorae (48.7—54.3, mean 51.0, n = 9). Distribution. —Both coasts of Baja Cali- fornia from San Ignacio and Pond lagoons (Pacific coast) south to lat. 27°N. Specimens examined.—Baja California (63). Remarks. —Specimens from San Ignacio Lagoon (“‘hueyi’’) are fairly consistently dull and green above and pale below, but are within the range of variation of castaneiceps from elsewhere. Dendroica petechia rhizophorae van Rossem Dendroica erithachorides rhizophorae van VOLUME 107, NUMBER 1 Rossem, 1935:67 (Tobari Bay, Sonora, Mexico). (SDNHM) Subspecific characters. — Both sexes near- est castaneiceps but average slightly more yellow below (less greenish); males with chestnut streaks usually wider; males with slightly shorter tails (see above) than cas- taneiceps. Both sexes more yellow on the back and rump than the subspecies from Sinaloa to Honduras (below). Compared with xanthotera, males with slightly nar- rower chestnut streaks below; chestnut bib more sharpely defined. Distribution. —Coastal Sonora from Te- popa Bay to Mazatlan, Sinaloa, Mexico. In- tergrades with the next subspecies appar- ently at Mazatlan, Sinaloa. Specimens examined. —Sonora (8). Remarks.—Van Rossem (1935) stated that the “‘tail has more yellow than in xan- thotera; more than in castaneiceps.”’ I found that the amount of yellow on the tail is too variable individually for identifying casta- neiceps from rhizophorae. There is nearly complete overlap in measurements of ex- posed culmen of rhizophorae and castanei- ceps (contra van Rossem 1935). Dendroica petechia phillipsi, new subspecies Holotype. —DMNH 36348, adult male, La Pinita de Jaltemba, El Islote, south- western Nayarit, Mexico, 16 April 1955, collected by A. R. Phillips (original number 3903). Subspecific characters. —Compared with rhizophorae and xanthotera, both sexes with greener back and rump; yellow below with slight greenish cast; edges of terials, wing coverts, and outer edges of rectrices greener. Chestnut streaks resemble rhizophorae. Distribution. —Pacific coast from Sinaloa to Honduras. Intergrades with rhizophorae apparently at Mazatlan, Sinaloa; inter- grades with xanthotera at San Lorenzo, Honduras. 45 Specimens examined. —Mexico: Sinaloa (7); Nayarit (16); Oaxaca (1); Chiapas (2); Honduras (4). Etymology. —For Allan R. Phillips in rec- ognition of his contributions to the taxon- omy of birds. Remarks. —Lowery & Monroe (1968) did not list breeding localities of D. petechia from the Pacific Coast of Mexico south of Naya- rit. The species probably breeds along the shore of Colima (Schaldach 1963), is pos- sibly a permanent resident of coastal Oaxaca (Binford 1989), and is a resident of coastal Chiapas (Alvarez 1964). The breeding sta- tus of the two males from Chiapas is un- known. One of seven old specimens from Mazatlan has a yellowish rump similar to rhizophorae. A male from San Lorenzo, Honduras, is green above but otherwise it is similar to specimens of xanthotera. Dendroica petechia xanthotera Todd Dendroica bryanti xanthotera Todd, 1924: 123 (Puntarenas, Costa Rica). (CM) Subspecific characters.—Compared with phillipsi, both sexes more yellow on the back and rump; males with chestnut more exten- sive on head; ventral chestnut streaks wider; chestnut bib not sharpely defined. Com- pared with aithocorys, males with chestnut hood restricted and darker; ventral chestnut streaks narrower. Females with less chest- nut below than aithocorys. Distribution. — Pacific coast of Nicaragua and Costa Rica. Intergrades with phillipsi at San Lorenzo, Honduras. Specimens examined. —Costa Rica (25). Dendroica petechia aithocorys Olson Dendroica petechia aithocorys Olson, 1980: 474 (5 miles east of La Honda, near Los Santos, Los Santos Providence, Panama). (USNM) Subspecific characters. —Males nearest xanthotera but slightly more yellow above; 46 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON chestnut hood paler and extends into upper breast; chestnut streaks wider and less sparse. Compared with iguanae, males greener above; chestnut hood paler. Compared with aequatorialis, males with darker chestnut hood; upper breast not streaked with yellow. Females more yellow (less greenish) below and more heavily streaked with chestnut than aequatorialis and xanthotera. Distribution. —Pacific coast of Panama from Chiriqui to Coclé including Isla Coiba and Azuero Peninsula. Intergrades with ae- quatorialis at Puerto Aguadulce. Specimens examined. —(77); same spec- imens compared by Olson (1980). Remarks. —Wetmore (1957) regarded birds from the above range as “intermedi- ates between aequatorialis and xanthotera,” and he and Lowery & Monroe (1968) re- ferred them to the former name. Specimens from Isla Coiba are slightly less streaked below than other specimens of aithocorys. More specimens from Isla Coiba have dor- sal chestnut streaks than do other specimens of aithocorys but these differences are minor and inconsistent. Three adult males, three adult females, and one subadult male (all USNM) from Isla Brincanco are slightly duller green above and paler yellow below, and the females are slightly less marked with chestnut than other specimens of aithocorys from elsewhere. Dendroica petechia iguanae Olson Dendroica petechia iguanae Olson, 1980:475 (Isla Iguana, Los Santos Province, Pan- ama). (USNM) Subspecific characters. —Males nearest aithocorys but darker, greener (less yellow) above and below; darker chestnut hood; chestnut streaks more dense. Chestnut hood darker than aequatorialis. Females near aithocorys but more greenish (less yellow) below; average more heavily streaked be- low. Distribution.—Isla Iguana, Los Santos Province, Panama. Specimens examined. —(13); same spec- imens compared by Olson (1980). Dendroica petechia aequatorialis Sundevall Dendroeca petechia aequatorialis Sunde- vall, 1870:609 (Guayaquil, Ecuador [= Panama City, Panama]). Royal Natural History Museum, Stockholm (fide Gyl- denstolpe 1926) Subspecific characters. —Males nearest aithocorys but chestnut of the head region paler; upper breast streaked with yellow. Compared with jubaris, males duller yellow below; chestnut region of head forms a def- inite but suffused hood vs. a cap. Females nearest jubaris but less yellow below; more heavily streaked with chestnut below than aithocorys. Distribution. — Pacific coast of the Prov- ince of Panama and the Pearl Islands. In- tergrades with jubaris at Rio Maje. Specimens examined. —(95); same spec- imens compared by Olson (1980); 20 others from Pearl Islands. Remarks.—Thayer & Bangs (1905) and Wetmore (1946a) reported that many of the specimens from Isla San Jose, Pearl Islands, were golden-orange where the yellow plum- age normally occurs. I found individual specimens of other subspecies, including ex- amples in the aestiva group, with traces of orange feathers. The large number of en- tirely golden-orange birds from Isla San Jose cannot be explained. Olson (1980) reported that males collect- ed at the mouth of Rio Majé show traces of yellow in the cheeks and that this may in- dicate intergradation between aequatorialis and jubaris. More noticeable in the Rio Majé males is that the crown and cheeks are in- termediate between these two subspecies in the hue of chestnut. Ventrally, the Majé specimens are more similar to jubaris than to aequatorialis in the hue and amount of chestnut streaks. The amount of yellow in the throats of the Majé series is also similar VOLUME 107, NUMBER 1 to that of jubaris but the yellow of the lower ventral regions is duller and thus similar to that of aequatorialis. Dendroica petechia jubaris Olson Dendroica petechia jubaris Olson, 1980:478 (Nuqui, [lat.] 5°40’N, Dept. Choco, Co- lombia). (USNM) Subspecific characters. —Compared to ae- quatorialis, males with dark chestnut crowns with remaining chestnut areas paler (tawny) and more suffused with yellow; chestnut hood absent; chestnut streaks wide and suf- fused on bright yellow ventral surface; dis- tinct chestnut cap; throat yellowish. Fe- males nearest aequatorialis but more heavily streaked with chestnut below; chestnut on head paler than other Pacific coast subspe- cies from Panama; more heavily streaked with chestnut below than peruviana. Distribution. —Darieén, Panama, south along the Pacific coast of Choco, at least to Buenaventura, Valle del Cauca, Colombia. Specimens examined. —(26); same spec- imens compared by Olson (1980). Dendroica petechia peruviana Sundevall Dendroeca petechia peruviana Sundevall, 1870:609 (Callao, Peru, and Guayaquil, Ecuador; type from Callao [?], fide Gyl- denstolpe [1926]). RNHM (fide Gylden- stolpe 1926). Subspecific characters. —Males nearest Jubaris but crown less suffused with pale chestnut; cheeks, lores, and chin more yel- low. Females with little to no ventral chest- nut streaking. Distribution. —Narino, extreme south- western coastal Colombia to northern coast- al Peru, and possibly as far south as Lima. Specimens examined. —Colombia: Nari- no (4); Esmeraldas (2); El Oro (8); Ecuador (6). Remarks.—The cheeks of a specimen from Narifio have some pale chestnut feath- 47 ers; chestnut on the checks of two specimens from Esmeraldas is even paler. Dendroica petechia aureola (Gould) Sylvicola aureola Gould, in Darwin, 1839: 86 (Galapagos Islands) Subspecific characters. —Nearest peru- viana but males less heavily streaked and both sexes usually less bright yellow below. Distribution. —Cocos Island and Gala- pagos Archipelago. Specimens examined. —Cocos Islands (27); Galapagos Islands (90). Remarks. —Plumage patterns in aureola resemble subspecies in the petechia group. However, specimens of aureola are more similar in size to birds of the erithachorides group (Wiedenfeld 1991). Summary of D. p. erithachorides group Generally, the chestnut streaks are nar- rower in northern males from the Pacific and Caribbean populations and are wider in most other populations. The chestnut on the hood and neck of males ranges from dark and forming (more or less) a hood in northern birds to paler and no distinct hood in southern populations from the Pacific coast and islands off western South Amer- ica. More specifically, northern populations of the Caribbean coast from Tamaulipas and Tabasco, Mexico (oraria), Yucatan to Nic- aragua (bryanti), and from the Pacific coast from Baja California (castaneiceps), Sonora (rhizophorae), Sinaloa to Honduras (phillip- si), and Nicaragua to Costa Rica (xantho- tera) have relatively narrower ventral chest- nut streaks than other populations of the erithachorides group. Back color of the northernmost populations of Pacific coast (castaneiceps) is greener than rhizophorae, but it is more yellow than in phillipsi, the next subspecies to the south. Males from along the Pacific coast of western Panama (aithocorys) have more extensive chestnut 48 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON hoods than xanthotera, the chestnut is darker in birds from Isla Iguana (7guanae), the chestnut is paler, with the upper breast be- ing more yellow, in birds from the Provice of Panama (aequatorialis), and the chestnut is even paler, with more yellow in the upper breast, in birds from eastern Pamama to Buenaventura, Colombia (jubaris). Males from coastal southwestern Colombia to northern Peru (peruviana) have less chest- nut and more yellow on the head than the other mainland populations of the Pacific coast. The populations of the Cocos and Galapagos islands (aureola) have still less chestnut on the head and neck, thus aureola is superficially similar to subspecies in the petechia group. Males from the Caribbean coasts of Costa Rica, Panama, and western Colombia (erithachorides) have darker chestnut heads than bryanti, but they are paler than birds from the Guajira Peninsula, Colombia (chrysendeta). Males from the Parajuana Peninsula, Venezuela (paragua- nae), are darker and greener above than chrysendeta. Acknowledgments I thank the staffs of the following muse- ums for the use of specimens: Academy of Natural Sciences of Philadelphia (ANSP), American Museum of Natural History (AMNH), British Museum (Natural His- tory) (BM), Carnegie Museum of Natural History (CM), Cornell University, Cowan Vertebrate Museum, Delaware Museum of Natural History (DMNH), Denver Muse- um of Natural History, Field Museum of Natural History (FM), James Ford Bell Mu- seum of Natural History, Museum of Com- parative Zoology (MCZ), Museum of Na- ture (Canada), Museum of Vertebrate Zoology (MVZ), Royal British Columbia Museum, San Diego Natural History Mu- seum (SDNHM), University of Alaska Mu- seum. Specimens from the U.S. National Museum of Natural History (USNM) were also compared. I thank K. Ackoff for pre- paring the range maps. I also thank R. W. Dickerman, B. L. Monroe, S. L. Olson, and K. C. Parkes for their critical reading of an early draft. I thank N. K. Klein, and es- pecially R. C. Banks for their useful com- ments on the present version. Literature Cited Aldrich, J. W. 1942. Specific relationships of the Golden and Yellow Warblers.—Auk 59:447— 449. Alvarez del Toro, M. 1964. Lista de las aves de Chia- pas. Instituto de Ciencias y Artes de Chiapas, Tuxtla Gutierrez, Chiapas, 82 pp. American Ornithologists’ Union. 1910. Check-list of North American birds. Third edition. American Ornithologists’ Union, Lancaster, Pennsylva- nia, 526 pp. 1931. Check-list of North American birds. Fourth edition. American Ornithologists’ Union, Lancaster, Pennsylvania, 372 pp. 1944. Nineteenth supplement to the Amer- ican Ornithologists’ Union Check-list of North American birds.— Auk 61:441-—464. . 1945. Twentieth supplement to the American Ornithologists’ Union Check-list of North American birds.— Auk 62:436-449. 1954. Twenty-ninth supplement to the American Ornithologists’ Union Check-list of North American birds.— Auk 71:310-312. 1957. Check-list of North American birds. Fifth edition. American Ornithologists’ Union, Baltimore, Maryland, 691 pp. 1983. Check-list of North American birds. Sixth edition. American Ornithologists’ Union, Washington, D.C., 877 pp. Audubon, J. J. 1831. Ornithological biography. Vol. 1. Adam Black, Edinburgh, 512 pp. Baird, S. F. 1865. Review of American birds in the museum of Smithsonian Institution.—Smith- sonian Institution Collections 181, 478 pp. ——., J. Cassin, & G. N. Lawrence. 1858. Birds. Vol. 9, Reports of explorations and surveys. . . for a railroad from the Mississippi River to the Pacific Ocean. Washington, D.C., 1005 pp. Batchelder, C. F. 1918. Two undescribed Newfound- land birds.—Proceedings of the New England Zoological Club 6:8 1-82. Behle, W. H. 1948. Systematic comments on some geographically variable birds occurring in Utah. Condor.—50:71-80. . 1985. Utah birds: geographic distribution and systematics. Utah Museum of Natural History, University of Utah, Occasional Publication No. 5, 147 pp. Binford, L. C. 1989. A distributional survey of the VOLUME 107, NUMBER 1 birds of the Mexican state of Oaxaca.—Orni- thological Monographs No. 43, American Or- nithologists’ Union, Washington, D.C., 418 pp. Bond, J. 1927. A new golden warbler from the island of St. Lucia, B. W. I. Auk 44:571-572. 1930. The resident West Indian warblers of the genus Dendroica. — Proceedings of the Acad- emy of Natural Sciences of Philadelphia 82:329- 337. 1936. Resident birds of the Bay Islands of Spanish Honduras. — Proceedings of the Acad- emy of Natural Sciences of Philadelphia 88:353- 364. 1942. Additional notes on West Indian birds.— Proceedings of the Academy of Natural Sciences of Philadelphia 94:89-106. 1950. Results of the Catherwood-Chaplin West Indies Expedition, 1948. Part 2. Birds of Cayo Largo (Cuba), and San Andrés and Provi- dencia.— Proceedings of the National Academy of Sciences of Philadelphia 102:43-68. Brewster, W. 1888. On three apparently new sub- species of Mexican birds.— Auk 5:136-139. Browning, M. R. 1978. An evaluation of the new species and subspecies proposed in Oberholser’s Bird Life of Texas. —Proceedings of the Biolog- ical Society of Washington 91:85-122. 1990a. Taxa of North American birds de- scribed from 1957 to 1987.— Proceedings of the Biological Society of Washington 103:432-451. 1990b. The nomenclatural status and type locality of the Yellow Warbler subspecies Den- droica petechia aestiva (Gmelin) (Aves: Paruli- nae). — Canadian Field-Naturalist 103:597-599. Buden, D. W. 1979. Ornithogeography of the south- ern Bahamas. Unpublished Ph.D. Dissertation, University Microfilms International, London, 277 pp. Cassin, J. 1860. Catalogue of birds collected during a survey for a ship canal across, [sic] the Isthmus of Darien, by order of the government of the United States, made by Lieut. N. Michler, of the U.S. Topographical Engineers, with notes and descriptions of species. — Proceedings of the National Academy of Natural Sciences of Phil- adelphia. Pp. 132-144; 188-197. Chapman, F.M. 1892. Notes on birds and mammals observed near Trinidad, Cuba, with remarks on the origin of West-Indian bird-life.— American Museum of Natural History Bulletin 4:279-330. Cherrie, G. K. 1891. Notes on Costa Rican birds. — Proceedings of the United States National Mu- seum 14:517-537. Coale, H. K. 1887. Description of a new species and sub-species of the genus Dendroica. —Ridgway Ornithological Club Bulletin 2:82-83. Cory, C. B. 1887. Descriptions of six supposed new species of birds from the islands of Old Provi- 49 dence and St. Andrews, Caribbean Sea.— Auk 4:177-180. 1909. The birds of the Leeward Islands, Ca- ribbean Sea.—Field Museum of Natural His- tory, Publication Number 137, Ornithological Series 1(5):193-255. Deignan, H.G. 1961. Type specimens of birds in the United States National Museum. — United States National Museum Bulletin 221, 718 pp. Dickey, D. R., & A. J. van Rossem. 1938. The birds of El Salvador. — Field Museum of Natural His- tory, Zoological Series 23, 609 pp. Gabrielson, I. N., & S. G. Jewett. 1940. Birds of Oregon. — Oregon State Monographs, Studies in Zoology No. 2, 650 pp. Gmelin, J. F. 1789. Caroli Linné. Systema naturae. Vol. | (part 1). Lugduni, 1032 pp. Godfrey, W. E. 1986. The birds of Canada. Revised edition. National Museum of Natural Science (Canada), Ottawa, Ontario, 595 pp. Gosse, P. H. 1847. The birds of Jamaica. John Van Voorst, London, 447 pp. Gould, J. 1839. Birds. In C. R. Darwin, Narrative of the surveying voyages of H.M.S. Beagle. Part 3. Smith, Elder & Company, Cornhill, 156 pp. Greene, E.R. 1942. Golden Warbler nesting in Lower Florida Keys.—Auk 59:114. Greenway, J., Jr. 1933. A name for the Golden War- bler of Old Providence Island. — Proceedings of the New England ZoGlogical Club 13:63-64. Grinnell, J. 1903. The California Yellow Warbler. — Condor 5:71-73. ,& A.H. Miller. 1944. The distribution of the birds of California.— Pacific Coast Avifauna No. 27, 608 pp. Griscom, L., & M. S. Crosby. 1926. Birds of the Brownsville region, southern Texas.—Auk 43: 18-36. Gyldenstolpe, N. 1926. Types of birds in the Royal Natural History Museum in Stockholm. — Arkiv for Zoologi 19:1-116. Hartert, E. 1893. On the birds of Aruba, Curacao, and Bonaire.—Ibis 5:299-338. Hellmayr, C.E. 1935. Catalogue of birds of the Amer- icas.— Field Museum of Natural History, Zoo- logical Series 13, part 8:1-541. Irving, L. 1960. Birds of Anaktuvuk Pass, Kobuk, and Old Crow.— United States National Mu- seum Bulletin 217, 409 pp. Jewett, S. G., W. P. Taylor, W. T. Shaw, & J. W. Aldrich. 1953. Birds of Washington state. University of Washington Press, Seattle, Wash- ington, 767 pp. Johnston, R. F. 1964. The breeding birds of Kan- sas.— University of Kansas Publications, Mu- seum of Natural History 12:575-655. Kessel, B., & D. D. Gibson. 1978. Status and distri- 50 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON bution of Alaska birds.—Studies in Avian Bi- ology No. 1, 100 pp. ——, & G. B. Schaller. 1960. Birds of the Upper Sheenjek Valley, northeastern Alaska. — Biolog- ical Papers of the University of Alaska No. 4, 59 pp. Lawrence, G. N. 1879. Catalogue of a collection of birds obtained in Guadeloupe for the Smith- sonian Institution, by Mr. Fred A. Ober. —Pro- ceedings of the U.S. National Museum No. 57: 449-462. Linnaeus, C. 1766. Systema Naturae. Edition 12. Holmiae, 532 pp. Lowe, P. R. 1907. On the birds of Blanquilla Island, Venezuela.—Ibis series 9, 1:111-122. Lowery, G. H., & B. L. Monroe, Jr. 1968. Family Parulidae. Pp. 3-93 in R. A. Paynter, ed. Check- list of birds of the world. Vol. 14. Museum of Comparative Zoology, Cambridge, Massachu- setts. Miller, A. H., H. Friedmann, L. Griscom, & R. T. Moore. 1957. Distributional check-list of the birds of Mexico. Part 2.— Pacific Coast Avifau- na No. 33, 436 pp. Oberholser, H. C. 1897. Description of a new sub- species of Dendroica. Auk 14:76-79. . 1938. The bird life of Louisiana. Department of Conservation, State of Louisiana, Bulletin 28, 834 pp. . 1974. The bird life of Texas. Volume 2. Uni- versity of Texas Press, Austin, pp. 531-1069. Olson, S.L. 1980. Geographic variation in the Yellow Warbler (Dendroica petechia: Parulidae) of the Pacific coast of Middle and South America. — Proceedings of the Biological Society of Wash- ington 93:473-480. Pallas, P. S. 1811. Zoographia Rosso-Asiatica. Vol. 1. Imperii Rossici, Petropoli. Pp. 297-568. Parkes, K. C. 1968. Some bird records from western Pennsylvania.— Wilson Bulletin 80:100-102. ——, & R. W. Dickerman. 1967. A new subspecies of Mangrove Warbler (Dendroica petechia) from Mexico.— Annals of the Carnegie Museum 39: 85-89. Peters, J.L. 1926. A new race of the Golden Warbler from the West Indies.—New England Zodlogi- cal Club 9:41. 1927. A revision of the Golden Warblers, Dendroica petechia (Linné). — Proceedings of the Biological Society of Washington 40:3 1-42. Phelps, W. H., & W. H. Phelps, Jr. 1950. Lista de las aves de Venezuela con su distribucion. Part 2, Passeriformes.—Boletin de la Sociedad Ve- nezolana de Ciencias Naturales 12:1—427. , & 1951. Las aves de las Islas Los Roques y Las Aves y descripcion de un nuevo canario de mangle. —Boletin de la Sociedad Ve- nezolana de Ciencias Naturales 13:7—30. ————>, &% 1959. Las aves de las Isla La Or- chila.—Boletin de la Sociedad Venezolana de Ciencias Naturales No. 93:252—266. , & E. T. Gilliard. 1941. Seventeen new birds from Venezuela.—American Museum of Nat- ural History Novitates 1153, 17 pp. Phillips, A., J. T. Marshall, & G. Monson. 1964. The birds of Arizona. University of Arizona Press, Tucson, Arizona, 220 pp. Phillips, J. C. 1911. A years’ collecting in the state of Tamaulipas, Mexico.— Auk 28:67-89. Ramos, M. A., & D. W. Warner. 1980. Analysis of North American subspecies of migrant birds wintering in Los Tuxtlas, southern Veracruz, Mexico. Pp. 173-180 in A. Keast & E. S. Mor- ton, eds. Migrant birds in the Neotropics: ecol- ogy, behavior, distribution, and conservation. Smithsonian Institution Press, Washington, D.C. Raveling, D. G., & D. W. Warner. 1978. Geographic variation of Yellow Warblers killed at a TV tow- er.— Auk 95:73-79. Ridgway, R. 1873. On some new forms of American birds.—American Naturalist 7:602-619. . 1884. Onacollection of birds made by Mes- sers. J. E. Benedict and W. Nye, of the United States Fish Commission steamer ‘“‘Alba- tross.” — Proceedings of the United States Na- tional Museum 7:172-210. . 1885. Some emended names of North Amer- ican birds.— Proceedings of the U.S. National Museum 8:354—-356. . 1887. Catalogue ofa collection of birds made by Mr. Chas. H. Townsend, on islands in the Caribbean Sea and in Honduras.— Proceedings of the U.S. National Museum 10:572-597. . 1902. The birds of North and Middle Amer- ica.— United States National Museum Bulletin 50, part 2, 834 pp. Russell, S. M., J. C. Barlow, & D. W. Lamm. 1979. Status of some birds on Isla San Andres and Isla Providencia, Colombia.— Condor 81:98-100. Schaldach, W. J., Jr. 1963. The avifauna of Colima and adjacent Jalisco, Mexico.—Proceedings of the Western Foundation of Vertebrate Zoology 1: 100 pp. Sclater, P.L. 1862. Catalogue ofa collection of Amer- ican birds. N. Trubner and Company, London, 368 pp. Sharpe, R. B. 1885. Catalogue of the birds in the British Museum. Volume 10. British Museum (Natural History), London, 682 pp. Slud, P. 1964. The birds of Costa Rica.—Bulletin of the American Museum of Natural History 128, 430 pp. VOLUME 107, NUMBER 1 Smithe, F. B. 1975. Naturalist’s color guide. Amer- ican Museum of Natural History, New York. Stevenson, H. M., & B. H. Anderson. 1993. The birdlife of Florida. University of Florida Press- es, Gainesville (in press). Stiles, G., & A. F. Skutch. 1989. A guide to the birds of Costa Rica. Comstock Publishing Associates, Cornell University Press, Ithaca, New York, 511 pp. Sundevall, C. F. 1870. Ofversigt af fogelslagtet Den- droica. Ofversigt af Kongl. Vetenskaps-Akade- miens Férhandhuger, 1869, no. 26:605-618. Sutton, G. M. 1967. Oklahoma birds. University of Oklahoma Press, Norman, 674 pp. Thayer J. E., & O. Bangs. 1905. The mammals and birds of the Pearl Islands, Bay of Panama.— Bulletin of the Museum of Comparative Zool- ogy 46:137-160. Todd, W.E. C. 1924: Descriptions of eight new neo- tropical birds.—Proceedings of the Biological Society of Washington 37:121-124. Twomey, A. C. 1942. The birds of the Uinta Basin, Utah. — Annals of the Carnegie Museum 28:341-— 490. Tye, A., & H. Tye. 1991. Bird species on St. Andrew and Old Providence islands, west Caribbean. — Wilson Bulletin 103:493-497. van Rossem, A. J. 1931. Report of a collection of land birds from Sonora, Mexico. — Transactions of the San Diego Society of Natural History 6:237-304. . 1935. The Mangrove Warbler of northwest- ern Mexico.— Transactions of the San Diego So- ciety of Natural History 8:67-68. . 1947. An undescribed race of the Mangrove Warbler from Baja, California, Mexico.— Transactions of the San Diego Society of Nat- ural History 11:49-52. Vieillot, J. P. 1808-1809. Historie naturelle des oiseaux de |’Amerique septentrionale. Vol 2. Chez Desfray, Libraire, De l’imprimerie, Paris, 74 pp., pl. 57bis + 124. Voous, K. H. 1957. The birds of Aruba, Curacao, and Bonaire. Pp. 1-260 in P. W. Hummelinck, ed. Studies on the fauna of Curacao and other Caribbean islands. Vol. 7. Martinus Nijhoff, The Hague. 51 Warren, R. L. M., & C. J. O. Harrison. 1971. Type- specimens of birds in the British Museum (Nat- ural History). Volume 2. Passerines. British Mu- seum (Natural History), London, 628 pp. West, G. C., & C. M. White. 1966. Range extensions and additional notes on the birds of Alaska’s Arctic Slope.—Condor 68:302-304. Wetmore, A. 1927. The birds of Porto Rico and the Virgin Islands—Psittaciformes to Passeri- formes. — New York Academy of Sciences 9:409— 598. . 1929. Descriptions of four new forms of birds from Hispaniola.—Smithsonian Miscellaneous Collections 81, 4 pp. . 1946a. The birds of San José and Pedro Gon- zalez islands, Republic of Panama.—Smithson- ian Miscellaneous Collections 106, 105 pp. . 1946b. New forms of birds from Panama and Colombia.— Proceedings of the Biological So- ciety of Washington 59:49-54. 1957. The birds of Isla Coiba, Panama.— Smithsonian Miscellaneous Collections 134, 105 pp. , R. Pasquier, & S. L. Olson. 1984. The birds of the Republic of Panama. — Smithsonian Mis- cellaneous Collections 150, part 4, 670 pp. Wiedenfeld, D. A. 1991. Geographical morphology of male Yellow Warblers.— Condor 93:712-723. Williamson, F. S. L., & L. J. Peyton. 1962. Faunal relationships of birds in the Iliamna Lake area, Alaska. — Biological Papers of the University of Alaska No. 5, 73 pp. Winker, K., B. A. Fall, J. T. Klicka, D. F. Parmelee, & H. B. Tordoff. 1991. The importance of avian collecting and the need for continued col- lecting.— Loon 63:238-246. Zimmer, T. T., & W. H. Phelps. 1944. New species and subspecies of birds from Venezuela. 1.— American Museum of Natural History Novi- tates 1270, 16 pp. National Biological Survey, National Museum of Natural History, Washington, D.C. 20560, U.S.A. PROC. BIOL. SOC. WASH. 107(1), 1994, pp. 52-59 A NEW SPECIES OF OCADIA (TESTUDINES: BATAGURINAE) FROM SOUTHWESTERN CHINA William P. McCord and John B. Iverson Abstract. —A new species of batagurine turtle, Ocadia glyphistoma, purport- edly from southwestern Guangxi Province, China, differs from Ocadia sinensis by having fewer broad neck stripes, bold dark markings on the ventral surfaces of the hind limbs, a distinctive medial notch in the tomium of the upper jaw, a broader carapace and plastron, a longer plastral forelobe, relatively shorter interpectoral, interabdominal, and interanal seams, and a relatively longer interhumeral seam. It differs from Ocadia philippeni by having bold dark mark- ings on ventral surfaces of the hind limbs, no obvious wash of ventral pink to orange pigment, a distinctive medial notch in the tomium of the upper jaw, a broader shell, a relatively shorter plastron and bridge, and relatively shorter interpectoral, interabdominal, and interanal seams. McCord & Iverson (1992) recently de- scribed a distinctive species of batagurine turtle (Ocadia philippeni) from Hainan Is- land, China, and compared that form with its only recognized congener, Ocadia sinen- sis. But during 1990 and 1991, Mr. Oscar Shiu of Hong Kong sent McCord a series of another distinct stripe-necked batagurine turtle from southwestern China with clear affinities to both O. sinensis and O. philip- peni (Fig. 1). Univariate analysis of variance (Table 1) and discriminant function analy- sis (Fig. 2) demonstrated that these new tur- tles were similar, but morphometrically dis- tinct from the latter two species, and so they are described herein as the third species of the genus Ocadia even though skeletal ma- terial is not yet available for definitive ge- neric placement (e.g., McDowell 1964, Hi- rayama 1984, Gaffney & Meylan 1988). A study of the mitochondrial genome of all Asian batagurines is underway by J. W. Bickham, Iverson, and McCord to test this allocation. Materials and Methods Preserved material was borrowed from the American Museum of Natural History (AMNHB), the British Museum of Natural History (BMNH), the California Academy of Sciences (CAS), the Field Museum of Natural History (FMNH), the Museum of Comparative Zoology at Harvard (MCZ), the Museum of Vertebrate Zoology at Berkeley (MVZ), the Florida Museum of Natural History at the University of Florida (UF), and the United States National Mu- seum (USNM), and living material was available in McCord’s private collection (WPM). Methods of measurement and analysis follow McCord and Iverson (1991). Character abbreviations are in Table 1. All measurements are in mm. Specimens examined included: Ocadia philippeni: China, Hainan Island (UF 80765-66 [holotye and paratype]; WPM 1- 7, alive). Ocadia sinensis: “Laos” (UF 80817-19 [3 specs]; WPM 1, alive), Viet- VOLUME 107, NUMBER 1 nam (BMNH 1903.7.2.1; MCZ 21051), Taiwan (FMNH 121230-32, 127172-73, 127175-78, 127180, 195492, 199750-51), China, Hainan Island (AMNH 30173, 30176-78, 30183-84, 30186-91, 30193, 30195—-96; FMNH 6613 [formerly AMNH 30194]; MCZ 20687; MVZ 23940; UF 80816 [1 skeleton]), China, mainland (BMNH 1947.3.5.26 [holotype]; MVZ 23943: WPM 1-2, alive), and No Data (BMNH 1947.3.4.24, cotype of Emys Ben- nettii). Ocadia glyphistoma: China (UF 84818 [holotype]; WPM 1-9, alive). Results and Discussion Ocadia elyphistoma, new species Guangxi stripe-necked turtle Fig. 1 Holotype. —UF #84818, an adult male, preserved in alcohol; reported to have been collected near the Vietnam border south- west of Nanning, Guangxi Province, China, but purchased from local people near Nan- ning by Mr. Oscar Shiu, in the spring of 1991. Diagnosis. —A medium-sized species of Ocadia (Table 2) most similar to O. philip- peni, with a wide, basically tricarinate car- apace having a prominent middorsal keel and weak lateral keels; an unhinged plas- tron; a medial notch in the tomium of the upper jaw (unnotched in other Ocadia); usu- ally four yellow, black-bordered lateral head and neck stripes separated by narrow brown stripes (at least eight black-bordered, nar- row, cream to yellow stripes in O. sinensis); the ventral surfaces of shell and skin not washed with pink, orange, or salmon (so colored in O. philippeni); the ventral sur- faces of thighs boldly marked with dark pig- ment (no such dark markings in O. sinensis or O. philippeni); a relatively long plastral forelobe (maximum length averages 41% of carapace length [CL] in female and 37% in male O. glyphistoma, 42% and 39% in O. Dhilippeni, and 38% and 36% in O. sinensis); a relatively long interhumeral seam (seam 53 length averages 8.8% of CL in female and 8.5% in male O. glyphistoma, 8.6% and 6.2% in O. philippeni, and 5.6% and 5.3% in O. sinensis); a relatively short interpectoral seam (length averages 18% of CL in female and 17.5% in male O. glyphistoma, 21% and 20% in O. philippeni, and 21% and 20% in O. sinensis); a relatively short interabdom- inal seam (length averages 21% of CL in female and 22% in male O. glyphistoma, 25% and 23% in O. philippeni, and 26% and 23.5% in O. sinensis); and a relatively short interanal seam (length averages 9.6% of CL in female and 8.5% in male O. glyphistoma, 11.2% and 12.4% in O. philippeni, and 10.6% and 10.1% in O. sinensis) [see also Table 2 and Fig. 3]. Description (based on two adult females, five adult males, one subadult female and two subadult males, including the holo- type). — Carapace length to at least 199 mm in males and at least 180 mm in females, elliptical, moderately tricarinate with a prominent medial keel and weak lateral keels, moderately domed (maximum shell height/CL = 0.338 to 0.435; mean = 0.41 for females, 0.38 for males), widest at mar- ginal M7 or M8 (maximum carapace width/ CL = 0.74 to 0.80 for females and 0.68 to 0.75 for males; means = 0.77 and 0.72, re- spectively), with a slightly serrated posterior margin, and with moderately obvious growth annuli (least obvious in old individ- uals). M1, 7, 8, and 9 largest (along carapace Margin), approximately coequal in length; M11 smallest; M9 tallest; M9-11 distinctly flared. Cervical scute small, usually longer than wide, wider posteriorly than anteri- orly, and indented medially along the pos- terior margin. Vertebrals V2—5 wider than long; V1 usually wider than long, but not contacting seam between M1 and M2; V5 usually not even close to contacting M10. Prominent medial keel most pronounced on V3 and V4; lateral keels weak (usually) to absent, but if present, most pronounced on costal C3. Carapace dark brown to nearly black, with seams more darkly marked; ca- “WU OST SI 1YsLI UO ofeUO; Jo Jey) PUL ‘WILT CRT SI Ifo] JOMO] Ul B[eU Jo yey) “WU gy] SI yo] Joddn ut seu Jo yI3ua] soedeIeD (7661 “UWOsIsAyT pue pIOHdl Ul tuaddijiyd ‘CE pur sisuauis ‘CE JO sainsy 10[O9 yyIM sIeduUI0d) DOJsIYdA]s DIpDIC JO suio\jed jese[d pue peofy “(10]00) “| “314 Zz ‘e) = ©) Z oe N < S ae fe) - 2 = Oo © 7) — x e Q © — e faa) aa) se = fy e) N ©) Z la jaa jaa} Q e) 4 A VOLUME 107, NUMBER 1 55 Table 1.—Results of univariate analysis of variance of residuals of 16 characters versus carapace length for three species of Ocadia (sexes analyzed separately). Differences are coded by first letter of species name; 1.e., “‘s- g” indicates a significant difference between sinensis and glyphistoma for that character for the indicated sex. Methods of character measurement are in McCord and Iverson (1991). Males Females Character PF Differences F Differences Maximum carapace width (CW) 31/2 S-g; D-S 2.64 — Maximum carapace height (CH) 1.46 — 0.01 — Maximum plastron length (PL) 0.90 — 2.95 = Maximum forelobe length (FL) 4.82* p-g; p-s 9.03** p-s Maximum hindlobe length (HL) 0.82 — 4.01* S-g; p-s Plastral width (PW1) 2.30 — 1.62 — Plastral width (PW3) 9.38*** p-g; D-s 4.24* p-s Plastral width (PW4) 10.34*** D-g; p-s 2.03 — Bridge length (BL) 1.90 — 3.30* p-g Gular width (GW) 0.69 — 0.22 — Gular length (GL) 4.44* p-g; D-s 0.60 — Interhumeral seam length (IH) 8.19** P-g; S-g 4.95* p-s Interpectoral seam length (IP) 5.67** D-g; S-g 2.77 S-g Interabdominal seam length (IAB) 1.49 — 8.41** p-g; S-g Interfemoral seam length (IF) ESS p-g; p-s 1.16 — Interanal seam length (IAN) 17.08*** all 1.44 — * 0.01 < P < 0.05. ** 0.001 < P < 0.01. *** P < 0.001. rinae usually not distinctly colored. A black notch on ventral posterolateral portion of each marginal, sometimes covering more than one-half of some marginals. Maximum plastron length shorter than carapace length (PL/CL = 0.95-0.98 in fe- males; 0.87 to 0.95 in males). Plastron slightly upturned anteriorly, with no hinge present. Plastral forelobe width (PW1) at level of junction of humeropectoral seam and lateral plastral margin relatively wide (PW 1/CL = 0.38 to 0.44 in females and 0.36 to 0.42 in males; means = 0.42 and 0.38, respectively). Anterior width of plastral hindlobe (PW3: at lateral junction of ab- dominofemoral seam) relatively wide (PW3/ CL = 0.45 to 0.47 in females and 0.40 to 0.43 in males; means = 0.46 and 0.42, re- spectively). Plastral hindlobe with relatively deep anal notch. Bridge moderately long (BL/CL = 0.37 to 0.39 in females and 0.33 to 0.37 in males; means = 0.38 and 0.35, respectively); single large axillary and in- guinal scutes on each bridge. Average plas- tral formula (see also Table 2 and Fig. 1 for diagnostic ratios): interabdominal seam (IAB) > interpectoral seam (IF) = interfem- oral seam (IP) > gular length (GL) > in- teranal seam (IH) = interhumeral seam (IAN). Plastron dark yellow-cream (Fig. 1), with a large (covering up to half of scute), black blotch on each scute (primarily on older portion of scute). A smaller black blotch also occurring on bridge area of pec- toral and abdominal scutes, and on axillary (usually) and inguinal scutes. Head narrow; upper jaw unhooked, but with medial notch; triturating surfaces of medium width. Very small tubercles evi- dent between angle of jaw and tympanum. Dorsum of head uniform dark olive-brown. Four (or sometimes five) narrow longitu- dinal black-bordered yellow stripes on side of head, separated by narrow brown or olive stripes; all four originating at orbit, upper two passing above tympanum, third from 56 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON @ philippeni O Hainan 4 Taiwan @ Laos *# China oO new e “15-125 -10 -75 -5 -25 0 25 5 7.5 DF1 Fig. 2. Plot of first two canonical axes for specimens of Ocadia based on discriminant function analysis of the residuals of 16 characters (listed in Table 1; method as in McCord and Iverson, 1991) for males (top) and females. First and second axes account for 42.7% and 30.5% of variation, respectively, for males, and 66.0% and 20.2% for females. Geographic locations represent populations of O. sinensis. top passing through tympanum and some- times broken anteriorly, and lower stripe passing below tympanum. All stripes ex- tending posteriorly to base of neck. Anterior continuations of stripes from orbit to nares variably obvious. Chin yellow, with vari- able vague black mottling (Sometimes form- ing circles), but with seven black-bordered yellow longitudinal stripes (often discontin- uous) extending from level of tympanum to base of neck. Tomia yellow with variable thin black markings. Black horizontal line across eye (through pupil); iris yellow-green. Anterior surface of antebrachium cov- ered with large, imbricate scales, the largest of which are sickle to spade-shaped; largest scales on hindlimb at heel, but much small- er than largest forelimb scales. Upper parts of limbs and tail covered with fine scales. Exposed parts of forelimbs dark olive-gray to nearly black, with faded yellow or orange @ philippeni O Hainan 4 Taiwan @ Laos * China oO new Fig. 3. Bivariate plot of relationships among males (top) and females of species of the genus Ocadia based on the characters IH/IP (interhumeral seam length/ interpectoral seam length) and IAN/FL (interanal seam length/maximum plastral forelobe length). Geographic locations represent populations of O. sinensis. or cream stripes (often discontinuous) ex- tending outward from base of limbs variable distances onto limb; stripes barely visible on dorsal surface of antebrachium, but very obvious on ventral aspect of limbs. Stripes on posterolateral margins of each limb al- ways obvious and extending at least to heel. Recessed areas of axillary region and be- tween neck and forelimbs boldly marked with alternating dark gray and yellow to fad- ed orange stripes. Recessed areas of inguinal region mostly yellow, but with some vague faded, dark gray blotching. Tail moderately long, gray-black dorsally, with a pair of vague longitudinal dorsolateral dark brown stripes extending along full length of tail; gray-black ventrally, but flecked with yellow or cream, and with a pair of longitudinal ventrolateral cream to light brown stripes extending the full length of the tail. Males with a slightly concave plastron; females with a flat plastron. Males with a longer tail than females; vent at level of pos- 57 VOLUME 107, NUMBER 1 SS (vO'T-E8'1) SET 1u(S8°7-E0'°7Z) 9E7Z ALZ 0-170) 70 (8S°0-6€'0) 150 (O8I-IZI) 6° ZSI € dj 06 I-ZS'*1) TL'T a(S7 7-68 1) 117 p87 0-172 0) €7'0 ae(LS 0-0) 60 (Sb7-ZOI) 6691 L W eulyD puojsiydajs ‘CQ (€9' 1-67 1) Sv'T (O1'7-L9'1) v8'T AS€0-S7'0) 870 A(€S O-E1°0) 870 (IL7-V11) 8161 1Z A 1(S6 I-O€ 1) vS'T a(L6 1-991) 781 p(ZE 0-770) 870 a(Iv'0-91'0) L770 (00Z-LOT) SOFT LI W ]B10.L (€S I-LED Srl (18 I-L9°1) vLI (67 0-L7 0) 870 (07 0-€1°0) L10 (8S$7Z-S61) I €7Z € dj (S61-ES 1) vLI (88 I-bL 1) 181 (1€°0-S7'0) 87:0 (8€ 0-L7'0) €£°0 (LvyI-9€1) vIvl G W «SOPT,, (LS I-O€ 1) Ev'T (16 I-€L 1) 781 (O€ 0-970) 87°0 (L€0-81°0) 870 (ILZ-1S7) 0'197 Z J (6S I-ZS' 1) 9ST (16 I-+L 1) 78'1 (ZE0-0E 0) TE0 (Ov 0-170) 0£ 0 (OSI-LI1) S'€€I (4 W eulyD (6S I-LE' 1D) Srl (Ol-Z-9L' 1) 88°1 (O€ 0-SZ'0) L770 (€S 0-170) €£°0 (SEZ-9LI) O' 107 6 d (OL I-€v' 1) €S'T (v6 I-bL 1) 98°1 (670-970) L770 (LEO-LI'0) €7'0 (OO7-SS1) L°LLI v W UeMIe |, (€9' 1-67 1) Sv'T (v0'7-89' 1) 78'T (S€0-67'0) T€°0 (9€'0-07 0) 970 (O@Z—-V11) 8°9¢1 L A (S9'I-O€ 1) IST (L6'1-99'1) 18°T (Z€°0-77'0) L770 (1v'0-91'0) £70 (SSI-LOI) €°SZI 6 W uvuley SISUIUIS “EC (T8°I-SS'1) 99°t u(66'1-66'1) 6671 (1€°0-€7'0) L770 (9¢'0-SE'0) Iv'0 (617-17) L917 Z dj (6 I-6€'1) ELT A9EC-LL 1) 761 p(LEO-77'0) TE -(09°0-SI'0) T€°0 (66I-€11) 0691 L W ueuley 1uaddyiyd ‘CQ a ae ee ee eS ee ee ee eee eee ee eee ee aVI/14 dI/1i TA/NVI dI/HI (wuz) yI3uUa] covdeieD u XS ajdures ee JS] DOUSIOYIP JULDYTUBIS Sv] (po199101d) s.Joysty Aq (S00 > q) WolayIp Apuroyrusis ore 1eY} 1OJOeILYO Jey} JOJ SUB Soloads o}¥OIPUT sodUeI BUIMOT[OJ S}dLiosiadns onoqeydje uouIWIOD ‘sosoyuored ul odueI Aq PoMoy][oy suvoul aie sonjeA ‘(74) yISUS] Sqojai0} [eNseId wWnuxeu pue “(gqy]) YI3UI] Wess eUTWIOpqeIO}UT ‘(NVI) Y13U9] Weos [eUeID}UI “(q]) YISUZ] WIvES [e10}09dI19} UT “(HI) W1ZUZ] WeaS [eIOUINYIOIUT Ie SUOTIRIAZIQGR JO}OBIeYO ‘VIPVIC snuds oY} JO sordeds BuOUIe BUTJLUTUILIOSIP UT [NJasN sisjoRIEYO OLN AWIOYdIOW—"Z 919eL 58 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON terior carapace margin in males; anterior to it in females. Etymology. —From the Greek glyphis (carved or notched) and stoma (mouth), in reference to the species’ distinctively notched upper jaw. Other material. —Two adult females, four adult males, one subadult female, and two subadult males (all from type locality, but not designated as paratypes); specimens all alive in the collection of William P. McCord (WPM 1-9), and to be deposited on their death in the UF collection. Distribution. —Known only from the re- gion of the type locality. The precise locality could not be determined since the turtles were collected by local people. Remarks. —Our studies of the genus Oca- dia have clarified the provenance of the type material of O. sinensis and O. bennettii, as well as their taxonomic relationship. Based on the original description and illustrations of Emys bennettii Gray (1844, 1855) and on Iverson’s examination of one of the co- types (BMNH 1947.3.4.24), we concur with Gunther (1864:27) that Emys bennettii is synonymous with Ocadia sinensis. In ad- dition, discriminant function analysis per- formed with that cotype as an unknown sug- gested that it originated on Taiwan and not on Hainan Island or the Chinese mainland (see range map in Iverson 1992). Measure- ments from the holotype of O. sinensis (BMNH 1947.3.5.26) were also analyzed and the specimen was confirmed to have originated on the Chinese mainland. We can also clarify somewhat the distri- bution of the genus Ocadia in Vietnam and adjacent China. Based on the descriptions of Siebenrock (1903), Bourret (1941; with illustrations), and Petzold (1963), O. sinen- sis is known from at least as far west as the Red River basin in Vietnam. Although we have not examined the specimens on which those descriptions were apparently based (e.g., Rijksmuseum van Natuurlijke His- toire of Leiden 4750, and Zoologisches In- stitut und Museum of Hamburg R00414 and RO00416), the illustrations in Bourret (1941) are clearly of O. sinensis. In addition, MCZ 21051 from Phuc Son, Vietnam, and BMNH 1903.7.2.1 from ‘““Annam” are also O. sinensis. Furthermore, given that Bour- ret (1941) called this species the most com- mon emydid in the Tonkin (Red) delta re- gion, and that Felix (1965) also found it to be common in the area west of Hanoi, its occurrence in at least the Red River basin in Vietnam seems unquestionable. How- ever, this suggests that the range of O. glyph- istoma in southwestern Guangxi Province, China, lies wholly within the range of O. sinensis, and that the two species may be broadly sympatric. Indeed, O. sinensis has been reported to occur only 40 km north of Nanning, Guangxi (at Wuming) by Lin (1984, in Buskirk, 1989). Unfortunately, until more museum material for this genus from southwestern China and adjacent Vi- etnam is available, the precise distributions of the individual species will remain uncer- tain. Acknowledgments This species was collected through the ef- forts of Oscar Shiu of Hong Kong. Curators and collection managers of the institutions listed in the text (especially D. L. Auth of UF) facilitated specimen loans. L. Carbone conscientiously typed early drafts of the manuscript. Support for Iverson was pro- vided by Earlham College, the Joseph Moore Museum of Natural History, The National Science Foundation (RFO 9021844 and DUE 9351508), and his family. Literature Cited Bourret, R. 1941. Les tortues de l’Indochine.—In- stitute Océanographique de I’Indochine 38:1- 235. Buskirk, J. R. 1989. New locality records for Chinese non-marine chelonians.—Chinese Herpetolog- ical Research 2(2):65-68. Felix, J. 1965. Turtles of Vietnam [in Czech].—Ziva 13:227-229. Gaffney, E. S., & P. A. Meylan. 1988. A phylogeny of turtles. Pp. 157-219 in M. J. Benton, ed., The phylogeny and classification of the tetrapods. VOLUME 107, NUMBER 1 Vol. 1. Amphibians, reptiles, birds. Clarendon Press, Oxford, England. Gray, J. E. 1844. Catalogue of the tortoises, croco- diles, and amphisbaenians in the collection of the British Museum. British Museum of Natural History, London, 80 pp. 1855. Catalogue of the shield reptiles in the collection of the British Museum. Part I. Tes- tudinata (tortoises). Taylor and Francis, Lon- don, 79 pp. Giinther, A. 1864. Reptiles of British India. Robert Hardwicke, London, 452 pp. Hirayama, R. 1984. Cladistic analysis of batagurine turtles (Batagurinae: Emydidae: Testudinoidea); a preliminary result.—Studias Geologicas Sal- manticensia Volumen Especial 1. Studias Pa- leonchelonologicas I 1984:141-157. Iverson, J. B. 1992. A revised checklist with distri- bution maps ofthe turtles of the world. Iverson Publishing, Richmond, Indiana, 363 pp. Lin, L. 1984. A new record of Testudinata of Gu- angxi—Ocadia sinensis (Gray).—Acta Herpeto- logica Sinica 1984-3(1):14. McCord, W. P., & J. B. Iverson. 1991. A new box turtle of the genus Cuora (Testudines: Emydi- 59 dae) with taxonomic notes and a key to the spe- cies.— Herpetologica 47:407—420. ——., & 1992. A new species of Ocadia (Testudines; Bataguridae) from Hainan Island, China.— Proceedings of the Biological Society of Washington 105:13-18. McDowell, S. B. 1964. Partition of the genus Clem- mys and related problems in the taxonomy of the aquatic Testudinidae.— Proceedings of the Zoological Society of London 143:239-279. Petzold, H. G. 1963. Uber einige Schildkréten aus Nord-Vietnam im Tierpark Berlin.—Sencken- bergiana Biologica 44:1-20. Siebenrock, F. 1903. Schildkroten des 6stlichen Hin- terindien.—Sitzungsberichte der Akademie der Wissenschaften in Wien. Mathematisch-Natur- wissenschaftliche Klasse 112(1):333-353. (WPM) East Fishkill Animal Hospital, Hopewell Junction, New York 12533, U.S.A.; (JBI) Department of Biology, Earl- ham College, Richmond, Indiana 47374, U.S.A. PROC. BIOL. SOC. WASH. 107(1), 1994, pp. 60-66 A NEW SPECIES OF MONTANE PITVIPER (SERPENTES: VIPERIDAE: BOTHROPS) FROM COCHABAMBA, BOLIVIA Michael B. Harvey Abstract. — Bothrops jonathani is a new species described from the Bolivian altiplano and adjacent xeric mountain sides, an area previously unknown to be inhabited by any species of pitviper. The new taxon is distantly allopatric from its most phenotypically similar congener B. alternatus. The new species is distinguished from other Bothrops by higher scale counts, relatively short hemipenial spines, a unique color pattern, and distinct prelacunal and second supralabial. Many of the forty-one (Campbell & La- mar 1989, 1992) currently recognized spe- cies of South American crotalines are rarely collected and remain poorly known despite recent advances in their study. South Amer- ican species formerly referred to Bothrops (sensu lato) were placed in five genera (Bur- ger 1971, Pérez-Higareda et al. 1985, Campbell & Lamar 1989). Evidence that three of these genera are monophyletic has recently come from biochemical and ana- tomical characters (Werman 1992), while the same analysis showed that Bothrops (sensu Burger 1971) is polyphyletic if Both- riopsis is recognized. Within Bothrops (sen- su stricto), evidence in support of two monophyletic lineages referred to loosely as the “neuwiedi’ and “‘atrox’ groups was provided (Werman 1992). However, these groups have yet to be formally defined or diagnosed. Although several crotalines occur at high elevations in the Andes, most species in- habit cloud forest or wet, upper montane forest. Only two species, Bothrops lojanus and B. ammodytoides occur in relatively xeric habitats above 2000 m and no pitvi- pers are known from the altiplano of Peru, Bolivia, and Argentina. Incidental to re- search (Harvey & Smith 1993, 1994) in the cis-Andean cloud forests of Santa Cruz and Cochabamba, Bolivia, a small herpetolog- ical collection was made in the altiplano and adjacent intermontane valleys of Cocha- bamba. Among material collected were two pitvipers herein described as a new species. Methods A string and meter stick were used to measure snout-vent length (SVL), tail length (TL), and tail circumference (TC) at the lev- el of the sixth subcaudal. With a dial caliper, distances were measured to the nearest 0.1 mm from the antero-ventral corner of the skin surrounding the eye to the caudal bor- der of the pit (EP), the antero-dorsal border of the skin surrounding the eye to the center of the nostril (EN), the caudo-dorsal to an- tero-ventral edges of the skin surrounding the eye (ED), and from the tip of the snout to the skin covering the caudalmost tip of the articular (HL). Nomenclature for the hemipenis is that of Dowling & Savage (1960). Scale counts of the new taxon were compared with ranges of other species re- ported by Campbell & Lamar (1989) and specimens examined in this study (Appen- dix). VOLUME 107, NUMBER | 61 Fig. 1. Bothrops jonathani, new species Figs. 1-2 Holotype.—Museo de Historia Natural “Noel Kempff Mercado” (MNK) R-1000, adult male collected on Highway 4 on 17 Jan 1992 by M. B. Harvey, approximately 35 km N (by road) of El Empalme, Provin- cia Carrasco, Departamento de Cochabam- ba, approximately 2800 m (17°45’S, 65°00’W). Paratype.—The University of Texas at Arlington (UTA) R-34564, adult female collected on 30 Dec 1991 by M. B. Harvey and E. N. Smith on Highway 4, 97 km S (by road) of Cochabamba, 3220 m. Diagnosis.— Bothrops jonathani is distin- guished from all other species of Bothrops by the following combination of character- istics: (1) prelacunal and second supralabial distinct; (2) high numbers of supralabials, intersupraoculars, and mid-body scale rows; (3) hemipenial spines two-thirds length of adjacent subcaudals; (4) anterior suprala- Bothrops jonathani, female paratype. UTA R-34564, SVL 540 mm. Photo by Eric N. Smith. bials with distinctive pattern; (5) paraven- tral spots diffuse; (6) gular stripes short. Description of holotype.—Rostral sub- triangular, about as wide as tall, about as wide as mental; nasal distinctly divided both above and below the naris; loreal single, bound dorsally by canthal; prefoveals 6/6; subfoveals in single row, increasing poste- riorly to four rows of interoculabials; post- foveals 2/2; lacunal not contacting suprala- bials; preoculars 2/2; upper preocular elongate and contributing anteriorly to can- thus; lower preocular squarish; suboculars 2/2, the first teardrop-shaped, the second elongate and crescent-shaped; postoculars 2/2; supralabials 11/12; infralabials 14 (right side incomplete, see remarks), first pair con- tacting medially; mental much broader than long; chin shields elongate, contacting first three infralabials; gulars in five rows be- tween chin shields and first ventral; seven rows of gulars separating first ventral from infralabials; two internasals; canthals 1/1, 62 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. separated posteriorly by seven intercan- thals; supraoculars about twice as long as wide, separated caudally by 10 intersupra- oculars; dorsals 3 1—33—23; ventrals 166; anal entire; subcaudals 39, all divided; supra-an- als seven; tail spine as long as adjacent five subcaudals; dorsals covering anterior 25% of tail spine. Dorsal scales three times as long as wide anteriorly becoming wider posteriorly, only about 1.5 times as long as wide on tail; dor- sal scales strongly keeled; paraventral row of scales smooth to very weakly keeled pos- teriorly, noticeably keeled on tail posterior to subcaudal 16; paraventrals about twice as wide as adjacent dorsal scales; most dor- sal head and temporal scales strongly keeled; internasals, canthals, and supraoculars smooth; scale row dorsal to supralabials smooth. Hemipenis (left): subcylindrical and bi- lobed; bifurcation of sulcus spermaticus at level of third subcaudal; bifurcation of lobes at sixth subcaudal; distribution of large to small keratinized spines asymmetrical on sulcate aspect of hemipenis, extending from level of first subcaudal along lateral surface, from level of third subcaudal on medial sur- face; largest spines on medial and lateral surfaces 2 mm or 7% length of adjacent sub- Facial pattern of female paratype of Bothrops jonathani. UTA R-34564, Head Length 30.7 mm. caudals; spines grading to finely papillate calyces at level of eighth subcaudal; papillae on calyces present only where ridges join; ridges of calyces extending medially to be- come lips of sulcus; lips of sulcus spinulate and papillate; asulcate surface of hemipenis covered in small and inconspicuous spines below bifurcation of lobes. Color in preservative (ethanol after buf- fered formalin): Facial color pattern com- plex; broad, dark brown postocular band edged in black and extending from ventral and caudal borders of eye to enclose pos- terior border of supralabial 9, most of su- pralabials 10-12, and two scale rows caudal to the rictus, extending across the adjacent infralabials and four rows of gulars; white stripe anterior to postocular band and ex- tending from ventral edge of supralabial 10 to subfoveal region and including part or all of supralabials 4-11; dorsocaudal corners of supralabials 1-6 white to grey; remainder of these scales dark tan; ventral border of prenasal white, edged in black; preocular, foveal, and nasal regions dark tan; ventral surface of head white with black and smoke- grey markings; smoke-grey stripes on gulars between chin shields and infralabials ex- tending from level of infralabial 9 to include first 3 infralabials and mental; infralabials VOLUME 107, NUMBER 1 63 Table 1.—Comparison of selected diagnostic characteristics of certain Bolivian and Argentinian Bothrops. Characteristic B. jonathani B. alternatus B. neuwiedi B. ammodytoides Supralabials 9-12 8-10 7-10 8-11 Intersupraoculars 10-12 8-13 6-9 7-11 Mid-body scale rows 30-33 24-37 21-29 23-25 Subfoveal row of scales present present absent absent Rostral normal normal normal elongate Lengths of hemipenial spines 74 subcaudals 7 subcaudals >1 subcaudal not examined Gular stripes short long short or absent short Anterior supralabials patterned without pattern without pattern without pattern Paraventral spots diffuse, not defined, over- diffuse to some- very diffuse, on ventrals lapping ventrals what defined, over- overlapping lapping ventrals ventrals 10-12 grey; other infralabials white and edged in black; most white gulars edged in black posteriorly, rarely immaculate. Ground coloration of ventrals white; a smoke-grey ventral pattern beginning an- teriorly as a medial stripe on first 20-30 ventrals but becoming many, staggered bands posteriorly; ventrals mostly smoke- grey caudally; subcaudals mostly smoke-grey becoming uniformly smoke-grey by subcau- dal 27; tail spine smoke-grey. Dorsal ground coloration dark tan; 34 pairs of large, dark brown blotches edged in cream meeting or staggered mid-dorsally; mid-dorsal blotches mostly rectangular, but C-shaped caudally; 49 small, dark brown blotches on flanks and sides of tail; most anterior blotches occupying dorsals 6—9 and 12-22; about 80 diffuse black blotches cov- ering paraventrals and dorsals 1-2, but paraventrals and dorsals 1—2 never com- pletely black; large medial blotch occupying most of frontorostrals; parallel dark stripes beginning on posterior 3 of supraoculars and extending caudally onto neck. Variations. —The female paratype is sim- ilar to the holotype with some noteworthy differences. The nasals are not completely separated dorsal to the nares. A large preoc- ular is divided into two scales, and the smaller preocular is fused to the suprala- cunal. There are 5/5 prefoveals, no post- foveals, 5/3 suboculars owing to fragmen- tation of the crescent-shaped subocular, 3/3 postoculars, 9/11 supralabials, 13/14 in- fralabials, 8 intercanthals, 12 intersupra- oculars, 28-30-21 dorsals, 175 ventrals, 37 subcaudals, and 26 dentary teeth. (Palatine and pterygoid teeth were not counted.) The tail spine is laterally compressed but bluntly rounded rather than pointed and 1.5 times as wide as the tail spine of the holotype. Dorsal and ventral colors of the paratype are similar to those of the holotype although some differences in pattern exist. A broad postocular stripe reaches the rictus, but does not extend onto the infralabials and gulars as in the holotype. Parallel stripes on the top of the head are broken into four blotch- es, a pair covering the caudal one-third of the supraoculars and 6/5 intersupraoculars, two pairs in the parietal and occipital regions, and two crescent shaped blotches on the neck. There are 30 mid-dorsal blotches and 49 lateral blotches, with the first blotches occupying dorsals 5—9 and 1 1- Dihe Measurements: (Measurements of holo- type followed by those of paratype in pa- rentheses) SVL 540 (540), TL 80 (60), TC 41 (28), HL 32.7 (30.7), EP 2.5-1.8 (1.8- 1.9), EN 6.6-6.6 (6.3-6.4), ED 3.5—4.0 (3.8- 4.0). Remarks.—The holotype had been run over by an automobile damaging the head so that measurements from the eye to the pit and the eye diameter are approximate. The caudalmost supralabials have been lost 64 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 3. presumed closest relative, B. alternatus. (Distribution of latter species after Campbell & Lamar 1989.) Distribution of Bothrops jonathani and its on the right side; only twelve remain. Some teeth may be missing, but 4 palatine, 12 pterygoid, and 16 dentary teeth remain. Distribution.— Bothrops jonathani is known from two localities at 2800 and 3220 m in Cochabamba (Fig. 3). Low, xeric- adapted shrubs cover the rocky hillsides where the holotype was found dead on the road. The paratype came from an area typ- ical of the Bolivian altiplano: a dry, rocky grassland, largely devoid of bushes except around dry stream beds. Both specimens were found on sunny days in the afternoon. Air temperature was 27°C when the para- type was collected. Etymology. —The specific epithet is a pa- tronym for Jonathan A. Campbell in rec- ognition of his considerable contributions to the biology of neotropical pitvipers. Comparisons.— Bothrops jonathani is most similar to B. alternatus. In addition to having fewer supralabials, B. alternatus ex- hibits a distinctive color pattern that readily distinguishes it from B. jonathani. In B. al- ternatus, stripes extending from the mental reach the level of the ventrals or, more of- ten, to the angle of the jaw. These stripes extend only about two-thirds as far in B. jJonathani. The first six supralabials of the new species are dark brown, but edged dor- socaudally in white to pale grey (Fig. 2); whereas, the same scales in B. alternatus are either uniformly pale or diffusely pigmented but always lack a definite pattern. In B. al- ternatus, the postocular stripe is edged dor- sally in cream. However, the postocular stripe grades abruptly to the dark tan ground coloration of the dorsum in B. jonathani. Finally, B. alternatus has a row of well de- fined spots on the lateral edges of the ven- trals, the paraventrals, and one to two rows of dorsals above the paraventrals. In B. jon- athani, these ventrolateral spots are diffuse and do not extend onto the ventrals. Most scale counts are higher in B. jona- thani than in either B. neuwiedi or B. am- modytoides (Table 1). Additionally, B. am- modytoides, and B. neuwiedi lack a row of subfoveals. In B. ammodytoides the rostral is vertically elongated so that the snout is upturned; a normal rostral occurs in the oth- er species. Finally, in B. neuwiedi the largest hemipenial spines are two to three times as long as those of B. jonathani. Five additional species of Bothrops occur in Bolivia, but are unlikely to be confused with Bothrops jonathani. The wet forest in- habitant B. microphthalmus differs from the former by having transverse cross-bands and by usually lacking a dorsal head pattern. Numbers of intersupraoculars, supralabials, infralabials, and mid-body scale rows are lower in B. microphthalmus than in B. jon- athani. Bothrops atrox, B. brazili, B. jarara- cussu, and B. sanctaecrucis possess lacun- alabial scales, whereas B. jonathani posses a distinct lacunal and second supralabial. As for B. microphthalmus, these species generally posses fewer numbers of intersu- praoculars, supralabials, infralabials, and mid-body scale rows. Within Bolivia, Bothrops jonathani is likely only to be confused with B. neuwiedi. However, the considerably more similar VOLUME 107, NUMBER | species B. alternatus reaches northern Ar- gentina and may eventually be found in Bo- livia (Fig. 3). Neither B. alternatus nor B. neuwiedi have been reported above 700 m and are almost certainly not sympatric with B. jonathani. Bothrops neuwiedi often oc- curs in dry areas, similar in this respect to the rain-shadow valleys and dry altiplano where B. jonathani occurs; but B. alternatus is found in deciduous forests and often as- sociated with swampy areas, riparian situ- ations, and generally more mesic habitats (Campbell & Lamar 1989) than the dry grasslands and desert scrub where B. jon- athani was collected. Finally, a third simi- larly patterned ‘species could eventually prove to be sympatric with B. jonathani. Bothrops ammodytoides also occurs in areas of montane desert scrub and reaches ele- vations of 2000 m in the Andes of Argentina (Campbell & Lamar 1989). Acknowledgments The manuscript benefited from many helpful comments provided by R. L. Gut- berlet, Jr. I thank A. Resetar (FMNH), T. Fritts (USNM), and J. Rosado (MCZ) for loaning specimens examined in this study. I thank D. Frost for providing me with working space at the American Museum of Natural History. I greatly appreciate the lo- gistical support provided by H. Justiniano and A. Castillo of the Fundacion Amigos de la Naturaleza. I thank the Bolivian Con- sejo Nacional, as well as I. Pinaya, Depar- tamento Vida Silvestre, MACA; and M. Avalos, Departamento Vida Silvestre, UTD-CDF SC, for granting permission for export of the specimens and for their pa- tience and kind understanding throughout the permit process. I acknowledge the kind assistance during the permit process afford- ed me by P. Bettella, T. Centurion, and N. Vacas of the Museo Noel Kempff Mercado, as well as E. Forno of the Fondo Nacional para El Medio Ambiente, and P. Ergeta of the Coleccion Boliviana de Fauna. Finally, I express my sincerest thanks to my good 65 friends B. and I. Phillips of the El Refugio project for their encouragement and logis- tical support. Literature Cited Burger, W. L. 1971. Genera of pitvipers (Serpentes: Crotalidae). Unpublished PhD. Dissertation, University of Kansas, Lawrence, Kansas, 186 pp. Campbell, J. A., & W. W. Lamar. 1989. The ven- omous reptiles of Latin America. Cornell, Ith- aca, 425 pp. ——, & W. W. Lamar. 1992. Taxonomic status of miscellaneous neotropical viperids, with the de- scription of a new genus.—Occasional Papers the Museum Texas Tech University 153:1-31. Crother, B. I., J. A. Campbell, & D. M. Hillis. 1992. Phylogeny and historical biogeography of the palm-pitvipers, genus Bothriechis: biochemical and morphological evidence. Pp. 1-19 in J. A. Campbell and E. D. Brodie, Jr., eds., Biology of the pitvipers. Selva, Tyler, Texas. Dowling, H. G., & J. M. Savage. 1960. A guide to the snake hemipenis: a survey of basic structure and systematic characteristics.— Zoologica 45: 17-28. Harvey, M. B., & E. N. Smith. 1993. A new species of aquatic Bufo (Anura: Bufonidae) from cloud forests in the Serrania de Siberia, Bolivia. — Pro- ceedings of the Biological Society of Washington 106:442-449. ——,& 1994. A new species of Bufo (An- ura: Bufonidae) from cloud forests in Bolivia. — Herpetologica (in press). Pérez-Higareda, G., H. M. Smith, & J. Julia-Zertuche. 1985. Anew jumping viper, Porthidium olmec, from southern Veracruz, México (Serpentes: Vi- peridae).— Bulletin of the Maryland Herpeto- logical Society 21:97-106. Werman, S. D. 1992. Phylogenetic relationships of Central and South American pitvipers of the genus Bothrops (sensu lato): cladistic analyses of biochemical and anatomical characters. Pp. 21— 40 inJ. A. Campbell and E. D. Brodie, Jr., eds., Biology of the pitvipers. Selva, Tyler, Texas. Department of Biology, UTA Box 19498, The University of Texas at Arlington, Ar- lington, Texas 76019-0498, U.S.A. Appendix Specimens Examined Specimens examined are followed by collection lo- cality in parentheses. Abbreviations refer to American Museum of Natural History (AMNH), Chicago Field Museum of Natural History (FMNH), Harvard Mu- 66 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON seum of Comparative Zoology (MCZ), National Mu- seum of Natural History (USNM), the Museo de His- toria Natural ‘““Noel Kempff Mercado,” Santa Cruz, Bolivia (MNK), the University of Texas at Arlington Collection of Vertebrates (UTA). Bothrops alternatus (25): BRAZIL: Rio Grande do Sul; UTA R-32427. Sao Paulo; FMNH R-2620, 171265, 171272, 171281-82, 171289, 171298. SE Bra- zil; MCZ R-17734, 17748-51. PARAGUAY: Central; UTA R-2848, 5602, 7484-85, 7573, 9721. URU- GUAY: Cerro Largo; FMNH R-12344. Treinta y Tres; FMNH R-10595. UNKNOWN (reportedly from Ar- gentina): UTA R-4999, 6306, 6789. UNKNOWN: UTA R-32420. Bothrops ammodytoides (10): ARGENTINA: Bue- nos Aires; FMNH 10830, 10832. ARGENTINA: Chu- but; MCZ 150292. La Roja; USNM 73421. ARGEN- TINA: Mendoza; FMNH 9994, MCZ 58104—07. San Luis; UTA R-16334. Bothrops microphthalmus (1): ECUADOR: Zamor- ra; UTA R-23530. Bothrops neuwiedi (19): ARGENTINA: Tucuman; FMNH 229950. BOLIVIA: Santa Cruz; AMNH 36008- 09, MNK 124, 168, 178, 189, 197, 475. BRAZIL: Goias; UTA 28232. Minas Gerais; FMNH 171255. Parana; MCZ 112528. Sao Paulo; FMNH 171277, MCZ 112526. PARAGUAY: UTA R-2849, 5603, 9834-35. URUGUAY: Lavelleja; 7601. PROC. BIOL. SOC. WASH. 107(1), 1994, pp. 67-78 A NEW SPECIES OF PLECTROHYLA (ANURA: HYLIDAE) FROM A PREMONTANE RAINFOREST IN NORTHERN HONDURAS Larry David Wilson, James R. McCranie, and Gustavo A. Cruz Abstract.—A new species of Plectrohyla, P. chrysopleura, from moderate elevations in the Cordillera Nombre de Dios of northern Honduras is described and illustrated, as is its tadpole. Its combination of prominent golden yellow flashmarks, blunt prepollex, moderate size, vocal slits present in males, spat- ulate teeth, and weakly to moderately tuberculate dorsal surfaces easily distin- guishes it from the other species of Plectrohyla. Several problems with a recent phylogenetic analysis of the genus Plectrohyla are noted and a phylogenetic reanalysis indicates that the new species is the one closest to the ancestral stock of the genus. Field work in the Quebrada de Oro region of northern Honduras continues to uncover herpetological novelties. The first trip was in June, 1980, and since that time, we have described a new genus and five new species of anurans from this area in the central por- tion of the department of Atlantida (Mc- Cranie & Wilson 1986; McCranie, Savage, & Wilson 1989; McCranie, Wilson, & Wil- liams 1989; Savage et al. 1988). Recently, a sixth new anuran described below was col- lected. Quebrada de Oro is a tributary of the Rio Viejo, in turn a tributary of the Rio Cangrejal, which flows into the Caribbean at La Ceiba, Honduras (see McCranie, Wil- son, & Williams 1989, for a description of the area). In May 1988, we collected specimens of a distinctive new species of the montane genus Plectrohyla at the Quebrada de Oro locality. Adults were found at elevations ranging from 930 to 990 m. McCranie and Eric Hedl, then a member of the United States Peace Corps stationed in Trujillo, Honduras, returned in 1989 and collected one adult at 990 m and two tadpoles at 1010 m. Although this species probably occurs in suitable localities at higher elevations in the Rio Viejo drainage, the known elevational range is near the lower limit for this genus. Duellman & Campbell (1992) indicated that the known elevational range for the other fifteen species of Plectrohyla is 615 to 3500 meters. Most species of Plectrohyla occur at intermediate elevations (1500-2700 m; sensu Stuart 1963), but a few, including the one described herein, range downward into moderate elevations (600-1500 m; sensu Stuart 1963). Plectrohyla teuchestes ranges downward to ca. 1000 m, P. guatemalensis and P. hartwegi to slightly less than 1000 m (we have collected P. guatemalensis as low as 990 m in the Sierra de Agalta above Ca- tacamas in Honduras), and P. matudai and P. quecchi occur as low as 615-700 m. The region of the “golden stream” (Que- brada de Oro) has been surprisingly pro- ductive of new taxa (others remain to be described). It is a figurative “gold mine” and once again has produced something “‘gold- en,” the “golden-sided tree frog.” Methods All measurements are in millimeters, made to the nearest tenth with dial calipers with the aid of a dissecting microscope. The 68 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Se ee Fig. 1. dorsal ground colors were compared to the color swatches in Smithe (1975). Color numbers used below refer to that publica- tion. Webbing formulas follow Savage & Heyer (1967) as modified by Myers & Duellman (1982). Subjective evaluations of tadpole jaw sheath configurations are pat- terned after Altig & Johnston (1986). The phylogenetic analysis was conducted using Swofford’s (1991) software package PAUP (Phylogenetic Analysis Using Parsimony, version 3.0s) on a Macintosh Plus. Plectrohyla chrysopleura, new species Fig. 1 Holotype. —National Museum of Natural History (USNM) 316547, an adult male from along the Quebrada de Oro (15°38'N, Plectrohyla chrysopleura, new species, male holotype (USNM 316547), SVL 63.8 mm. 86°47'W), 970 m elev., tributary of the Rio Viejo, south slope of Cerro Bufalo, Cordille- ra Nombre de Dios, Departamento de At- lantida, Honduras, collected 23 May 1988 by James R. McCranie and Larry David Wilson. Original number LDW 8907. Paratypes. —USNM 316548, adult male, same data as for holotype, except elevation 930 m; USNM 316549-S0, both adult males, same data as for holotype, except elevations unrecorded and collected 3 May, 1988 by Gustavo A. Cruz; USNM 316551, adult male, same data as for holotype, ex- cept elevation 990 mand collected 1 August 1989 by Eric Hedl and James R. McCranie. Diagnosis. —A species of Plectrohyla dis- tinguished from its congeners by the follow- ing combination of characteristics: moder- VOLUME 107, NUMBER 1 ate size (56.6—65.6 mm snout-vent length); dorsum weakly to moderately tuberculate; vocal slits present; maxillary-premaxillary teeth spatulate; no vertical rostral keel; pre- pollex flat, elongate, terminally blunt; dor- sum Smoke Gray (color 45) with a bronze sheen or Cinnamon (color 123A); promi- nent golden yellow flashmarks present on front and hind limbs, side of body, axilla, and groin (all flashmarks hidden when frog is at rest). Description of holotype. — Adult male with snout-vent length (SVL) of 63.8; tibia length 37.7, 59.1 percent of SVL; foot length 31.0, 48.6 percent of SVL; head length 21.4, 33.5 percent of SVL; head width 23.0, 36.1 per- cent of SVL. Snout of moderate length, dis- tance from anterior edge of orbit to tip of snout 8.6, 134.3 percent diameter of eye; snout truncate in dorsal aspect and in pro- file, lacking a vertical rostral keel; canthal ridge slightly thickened; loreal region slight- ly concave; lips moderately thickened, slightly flared. Nostrils protuberant, direct- ed dorsolaterally, situated near tip of snout; internarial distance 4.9; internarial area slightly depressed near point of convergence of canthal ridges; top of head flat; interor- bital distance 6.3, 27.4 percent of head width; diameter of eye 6.4; width of eyelid 4.9, 21.3 percent of head width. Moderately heavy dermal fold extending posteriorly from posterior edge of orbit, merging with body contour above point of insertion of arm, barely covering upper edge of tym- panum, remainder of tympanum distinct; diameter of tympanum 3.3 mm, 51.6 per- cent of eye diameter. Arms moderately robust, forearm slightly heavier than upper arm; distinct transverse fold on wrist. No axillary membrane. Fin- gers long, slender; length of fingers from shortest to longest, 1-2-4-3, fourth toe near- ly as long as second; disc on third finger subequal in size to tympanum; webbing ves- tigial between first and second fingers, web- bing formula IT 2-3 III 2 1/2-2* IV; subar- ticular tubercles large, subconical; distal 69 tubercle on fourth finger normal; supernu- merary tubercles in single rows on proximal segments of fingers; pollex flat, elongate, ter- minally blunt, spine not protruding through skin, lacking nuptial excrescences. Heels slightly overlapping when hindlimbs ex- tended to right angles of body; no transverse dermal fold on heel; inner tarsal fold ex- tending full length of tarsus; no outer tarsal fold; inner metatarsal tubercle ovoid, barely visible from above; no outer metatarsal tu- bercle. Toes long, slender; length of toes from shortest to longest, 1-2-5-3-4, fifth toe near- ly as long as third; discs moderately large; subarticular tubercles moderately large, subconical; supernumerary tubercles small, low, in single row on proximal segment of each digit; toes about three-fourths webbed, webbing formula I 1*-1 1/2 11 1*-2 III 1*-2 IV 2-1* V. Vent opening directed posteroventrally at level of mid-thigh; anal sheath short, broad. Skin on dorsal surface, throat, and chest weakly tuberculate; skin on ventral surfaces of forearms moderately tuberculate; skin on belly, ventral surfaces of thighs, and below vent granular; skin on ventral surface of shanks smooth. Tongue nearly round; upper jaw shallowly notched medially; maxillary- premaxillary teeth spatulate; vomerine teeth 6-5, situated on small elliptical elevations between ovoid choanae; vocal slits present, extending from midlateral edge of tongue to angle of jaw; vocal sac single, median, subgular. Color in life: dorsum of head, body, and limbs Smoke Gray (color 45) with a slight bronze sheen; tympanum pale coppery bronze; lips slightly paler than remainder of head; iris gold with black reticulations; prominent golden yellow flashmarks at ax- illa, side of chest, and underside of arm (all contiguous), similar but slightly darker flashmarks at groin, on anterior surface of thigh, underside of shank, upper surface of tarsus, and upper surface of foot (all flash- marks hidden when frog at rest); chin and chest gray with a golden sheen; belly gray 70 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 1.— Variation in measurements (in millime- ters) and proportions (percentages) in five male Plec- trohyla chrysopleura. Character abbreviations are in brackets. Means are in parentheses following ranges. Measurements Character and proportions Snout—vent length [SVL] 56.6-65.6 (62.1) Tibia length [TL] 29.3-37.7 (34.1) (TL/SVL) 46.6-59.2 (54.9) Foot length [FL] 26.5-31.6 (29.3) (FL/SVL) 45.5-48.6 (47.1) Head length [HL] 20.0-21.7 (20.7) (HL/SVL) 32.4-35.5 (35.1) Head width [HW] 20.8-23.0 (21.8) (HW/SVL) 33.5-37.1 (35.1) Snout length [SL] 7.6-9.2 (8.3) (SL/ED) 121.2-146.0 (133.1) Eye diameter [ED] 5.8-6.6 (6.3) Internarial distance 4.3-5.2 (4.8) Interorbital distance [[OD] 5.9-7.2 (6.3) (IOD/HW) 27.4-31.4 (28.8) Eyelid width [EW] 4.9-5.4 (5.2) (EW/HW) 21.3-25.7 (23.8) Tympanum diameter [TD] 3.0-3.8 (3.4) (TD/ED) 47.0-58.7 (53.1) with yellowish cast, as is undersurface of thigh; posterior thigh surface golden yellow with dense olive green smudging; palms pale gray; soles gray. Color in alcohol: dorsal surfaces grayish- brown; ventral surfaces pale gray; flash- marks dirty white. Variation. Measurements and propor- tions of all specimens are given in Table 1. Color and pattern of the paratypes are in essential agreement with that of the holo- type, except that the dorsal surfaces of USNM 316551 were Cinnamon (color 123A) in life. Most paratypes are somewhat more tuberculate on the dorsal surfaces than is the holotype. Description of tadpole.—One lot of two Plectrohyla tadpoles (USNM 316552) pre- sumed to be of this species (no other Plec- trohyla is known from the type locality nor is expected to occur there) is available. A tadpole (Fig. 2A) in stage 36 (Gosner 1960) may be described as follows: body length 16.8; tail length 27.8; total length 44.6; body slightly depressed, a little wider than high; snout semicircular in dorsal aspect, rounded in profile; eyes moderately small, widely separated, directed laterally; nostrils situ- ated at a point slightly closer to eyes than tip of snout, directed anterolaterally; spi- racle sinistral, directed posterodorsally, sit- uated near midline, at a point about two- thirds distance from tip of snout to posterior end of body; vent tube moderately long, dextral; caudal musculature robust, extend- ing nearly to tip of rounded tail; height of caudal musculature at midlength of tail greater than height of either dorsal or ven- tral fins; dorsal fin extending very narrowly onto posterior end of body. Oral disc (Fig. 2B) large, ventral, com- pletely bordered by two rows of moderately large marginal papillae (ca. 10/mm); single row of submarginal papillae surrounding A-1 and P-3 tooth rows, larger (ca. 7/mm) than marginal papillae; submarginal papil- lary row expanding to two-three rows lateral to jaw sheaths; oral disc not emarginated; keratinized jaw sheaths medium-sized, bearing short, pointed serrations; upper jaw sheath widely arched, with well-developed lateral processes; lower jaw sheath widely V-shaped; labial tooth rows 74 with second anterior row narrowly interrupted medially; anterior tooth rows subequal, long, extend- ing to lateral portion of oral disc; posterior tooth rows subequal, noticeably shorter than anterior rows. The second tadpole is in stage 26 and has a body length of 13.4, a tail length of 20.9, and a total length of 34.3. This specimen is very similar in morphological features to the larger tadpole described above. Color in life of the larger tadpole was as follows: body brown; caudal musculature creamy-tan, boldly spotted with brown; caudal fins clear with brown spots. The smaller tadpole was similar, except that it lacked the bold markings on the caudal musculature and tail fins. Natural history notes.—All adults, for VOLUME 107, NUMBER 1 71 : al ml! wea Sl! WT MLL) Css. NN) i ING 1mm Fig. 2. Tadpole of Plectrohyla chrysopleura (USNM 316552) in Gosner stage 36: (A) lateral view; (B) oral disc. which information is available, were col- lected at night on boulders in splash zones near waterfalls in the Quebrada de Oro. The tadpoles were collected in a plunge pool in a small tributary of the Quebrada de Oro with tadpoles of Ptychohyla spinipollex and Rana maculata. The vegetation of the Quebrada de Oro area lies in the Premontane Wet Forest for- mation of Holdridge (1967), and was de- scribed by McCranie, Wilson, & Williams (1989). The section of Quebrada de Oro in which we have worked since 1980 was dec- imated by a huge landslide in November of 1988, precipitated by deforestation on the steep hills above the stream. The impact on the populations of the numerous species of amphibians resident in the environs of the stream is unknown, but is thought to be substantial. 72 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Etymology.—The name chrysopleura is derived from the Greek words chrysos and pleura, meaning “‘gold”’ and “‘side,”’ respec- tively, in reference to the yellow flash mark- ings on the frog. Discussion With the description of Plectrohyla chry- sopleura, the genus now comprises sixteen described species. The recent reapprasial of the genus by Duellman & Campbell (1992) perforce leaves a number of questions un- answered. Duellman & Campbell (1992) described two new species of Plectrohyla (P. acan- thodes and P. teuchestes), segregating them from the formerly composite P. guatema- lensis. In addition, they conducted “two levels of phylogenetic analysis” of the fifteen known species. The first analysis involved an attempt to identify suitable out-groups and to establish the monophyly of Plectro- hyla. The second was an effort to create a cladogram for the in-group, Plectrohyla. As Duellman & Campbell (1992:25) ac- knowledged, “... the morphological data set that [was used] is not sufficent to resolve the phylogenetic relationships among the species of Plectrohyla.”’ Questions also exist concerning their out-group analysis. We dis- cuss our concern with the latter first. Duellman & Campbell (1992) identified four synapomorphies uniting Plectrohyla and the Hyla bistincta group as sister-taxa. Unfortunately, they do not unequivocally indicate their concept of the latter group. They make reference to the work of Duell- man (1970), who recognized the Hyla bis- tincta group as comprising nine species. They (1992:21) further stated that two of the species included by Duellman (1970) in the group, H. charadricola and H. chryses, “... lack the thick, glandular dorsal skin characteristic of the other, larger species in the group.”’ As a consequence, they did not include these species in the first out-group. No mention was made, however, of the spe- cies described and placed subsequently in the bistincta group or related groups by Ad- ler & Dennis (1972) and Caldwell (1974). The latter work is especially important, in- asmuch as Caldwell reorganized the mem- bers of Duellman’s (1970) bistincta group (plus the species described subsequently, in- cluding her own, as well as H. arborescan- dens) into four species groups (bistincta, ar- borescandens, charadricola, and crassa), leaving the bistincta group with only two members (bistincta and pentheter). Perhaps, Duellman & Campbell (1992:21) had Cald- well’s study in mind when they penned the quizzical statement, ““The monophyly of the Hyla bistincta group seems to be assured, but the limits of the group remain to be ascertained.”” This statement appears to constitute a reverse non sequitur, i.e., the second clause of the sentence is supposed to follow from the first. Group limits (i.e., content) have to be established before a case for monophyly can be made. If this had been done, making a case for the bistincta group as the first out-group and sister-taxon of Plectrohyla would rest upon a secure base instead of being equivocal. The choice of the second out-group, the species Hyla miotympanum, by Duellman & Campbell (1992:21), was acknowledged to be “... frought [sic] with uncertainty.” They appear to have selected this taxon by a process of elimination, excluding from consideration various groups of stream- breeding hylines with highly-specialized tadpoles. They doubtless had little choice, inasmuch as their decision was hampered by “. . . the absence of phylogenetic analyses of the diverse groups of hylids” (p. 21). In light of the questions concerning the choice of out-groups, we wonder what weight can be given to the determination of features as apomorphies or plesiomorphies. Pursuing the work to clarify such matters is outside of the scope of the present work and thus we tentatively accept Duellman & Camp- bell’s (1992) out-group analysis in attempt- ing to determine the phylogenetic position of Plectrohyla chrysopleura. We also accept the hypothesis that the genus Plectrohyla is VOLUME 107, NUMBER 1 73 Table 2.—Morphological characters and coded character states used in phylogenetic analysis of Plectrohyla. Number in parentheses following a character refers to the respective character number in Duellman & Campbell (1992). State 0 is the primitive condition. Character transformation is 0 — | in all characters except character 3, which is 0 > 1 > 2 —>3. Character . Squamosal (11) . Humerus (12) . Prepollex (13) 2 i=) Character states : otic ramus of squamosal articulating with the crista parotica : otic ramus of squamosal not articulating with the crista par- otica : humerus round in section : humerus having well-developed flanges : prepollex slightly enlarged, cartilaginous or ossified, rounded or elliptical : prepollex enlarged, elongated, ossified, flat, terminally blunt prepollex enlarged, elongated, ossified, terminally curved with a single spine prepollex enlarged, elongated, ossified, with two curved spines 4. Rostrum (14) 5. Linea masculinea (15) 6. Vocal slits (16) 7. Expansion of oral disc (18) 8. Serrations on upper jaw sheath (19) 9. Length of posterior tooth rows (21) 10. Lateral processes on upper jaw sheath (20) 11. Maxillary teeth Fee ie ey Sh TTS ST A. Re monophyletic, although we feel that such an Opinion would stand on much firmer foot- ing were the phylogenetic relationships within the presumed sister-taxon, the Hyla bistincta group, and those of Middle Amer- ican stream-breeding hylids in general, bet- ter understood. The in-group analysis of Plectrohyla car- ried out by Duellman & Campbell (1992), however, is particularly frustrating, in part due to the uncooperative nature of the members of the genus. Sufficient-sized se- ries of some of the species of Plectrohyla are notoriously difficult to assemble. Nonethe- less, rendering a strict consensus tree pos- sessing a polytomy of 11 clades for 15 spe- rostrum plain rostrum with vertical keel linea masculinea absent linea masculinea present vocal slits present vocal slits absent oral disc not expanded and suctorial oral disc expanded and suctorial serrations subequal in size two or more serrations enlarged, fanglike posterior tooth rows shorter than anterior rows posterior tooth rows equal in length to anterior rows lateral processes well-developed lateral processes weak or absent maxillary teeth spatulate maxillary teeth pointed cies is close to having no in-group analysis at all. For Duellman and Campbell to imply in the title that their paper somehow re- solves the “phylogenetic relationships” of the species of Plectrohyla is certainly mis- leading. The characters and states used by Duell- man & Campbell (1992) in the in-group analysis contain several mistakes and/or potential ambiguities. We thought that a more effective analysis would modify their hypothesis about the group relationships. As a result, we undertook a phylogenetic reanalysis of the genus Plectrohyla; discus- sion of our treatment of characters follows. A major oversight of Duellman & Camp- 74 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON bell was the fashion in which the states as- sociated with the prepollex were coded in the out-group (their character 8) and in- group (their character 13) analyses. The en- larged, ossified nature of the prepollex in species of Plectrohyla was cited as evidence for the monophyly of the genus. In the in- group analysis, however, three states were identified for this character, viz., (1) blunt, (2) pointed, simple, and (3) pointed, bifid. The polarization and transformation seri- alization was identified as 0 > 1 — 2 in the same order. The “0” condition, however, was stated by Duellman & Campbell (1992: 23) to be “The primitive condition (that found in both the first and second out- groups) ....” Thus, a blunt, enlarged, and ossified prepollex is indicated as occurring in Hyla miotympanum and members of the H. bistincta group. Such is patently not the case, so the character states and their po- larity were recoded (Table 2). A second concern is with Duellman & Campbell’s character 17 (rows of accessory papillae). Two states were given for this character as follows (0 = 1): one row of accessory papillae on each labium. one row of accessory papillae on an- terior labium and two rows on pos- terior labium. Five species of Plectrohyla (hartwegi, ixil, matudai, pokomchi, and teuchestes) were said to have the derived state. However, Duellman & Campbell (1984:396) stated that there is a “single row of larger papillae medial to fringing papillae” in pokomchi. Their illustration (fig. 6) of the pokomchi tadpole also shows one row of large acces- sory papillae (= submarginal papillae) on the posterior labium in this species. Duell- man & Campbell (1992:16) also stated that there are “four or five large submarginal papillae between posterior labial tooth row and posterior margin of lip” in teuchestes and (p. 10) that there are “‘6—8 large papillae medially between third posterior tooth row and posterior edge of disc” in hartwegi. Thus, the situation in hartwegi and teuchestes is radically different from the condition where there are two rows of submarginal papillae on the posterior labium as found in ixi/ and matudai tadpoles (compare illustrations of latter two species in Duellman (1970), with those of hartwegi and teuchestes in Duell- man & Campbell 1992). We have deleted this character from our analysis because of the erroneous scoring of this character by Duellman & Campbell and the fact that an examination of the tadpoles of each species of Plectrohyla would be necessary before we could confidently recode this feature. Such an analysis is outside the limits of our study, but we would urge future workers studying the phylogeny of Plectrohyla to investigate the potential utility of this character. A third problem is Duellman & Camp- bell’s character 20 (lateral processes on up- per jaw sheath). Three states were given for this character as follows (0 — 1 — 2): 0 = lateral processes long. 1 = lateral processes short. 2 = lateral processes weak or absent. Two species (acanthodes and guatemalen- sis) were said to have the state 0, two (avia and pokomchi) state 1, and the remaining species (where known) state 2. Campbell & Kubin (1990:table 1) defined the lateral pro- cesses of acanthodes (as guatemalensis: Campbell & Kubin’s source of information for this taxon was Duellman (1970), who actually described and illustrated the acan- thodes tadpole under the name guatemalen- sis), avia, pokomchi, and sagorum as “short, narrow.” These four species, all with “short” lateral processes, are coded three different ways in Duellman & Campbell’s data ma- trix. Also, numerous contradicting state- ments can be found in Duellman & Camp- bell (1992) by comparing the species diagnosis for acanthodes, dasypus, glandu- losa, guatemalensis, pokomchi, quecchi, sa- gorum, and tecunumani with their data ma- trix (table 8). Additionally, Honduran P. glandulosa tadpoles (glandulosa coded state VOLUME 107, NUMBER 1 75 Table 3.—Character matrix for the species of Plectrohyla in the phylogenetic analysis. Missing characters coded 9. Character numbers refer to Table 2. Character nN w oS Taxon Ancestor acanthodes avia chrysopleura dasypus glandulosa A glandulosa B guatemalensis hartwegi ixil lacertosa matudai pokomchi pycnochila quecchi sagorum tecunumani teuchestes oo ow ow ow G8 3S) FS OS BS 3S BS AS DS FS Ge) BS Ge) SOSOSeOSo=S=-Se000000O0 Were NN KK WN KH NY WW HK = = NY WO Ser K ODD CDCOCOCOCOCOCOCOCCSO 2 by Duellman & Campbell) have lateral processes similar in length to those of P. guatemalensis (guatemalensis coded state O by Duellman & Campbell). Because of the contradictory evidence, we have recoded this character as either well-developed, or weak or absent (Table 2). This action necessitates distinguishing Guatemalan P. glandulosa, with weak or absent lateral processes, from Honduran P. glandulosa. These taxa are la- beled glandulosa A and glandulosa B, re- spectively, in our data matrix (Table 3). The nine species coded in Duellman & Camp- bell’s (1992) data matrix as having ““weak or absent” lateral processes were similarily coded in our data matrix, whereas the re- Maining species (where the tadpoles are known) were coded as well-developed (Ta- ble 3). A final matter relates to the lack of use of a character discussed by Duellman (1970), viz., the nature of the maxillary teeth (spat- ulate vs. pointed). Maxillary teeth of the H. bistincta group (sensu Duellman 1970) are spatulate, as are those of most species of Middle American hylids (Duellman 1970). Thus, we hypothesize that spatulate teeth 5 6 7 8 9 10 11 0 0) 0 0 0 0) 0 0 1 0 0 0 0 0 0 1 0 0 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 1 0 0 0 1 1 0 1 0 0 0 0 1 0 1 0 0 0 0 0 0 1 1 0 1 1 0 1 0 0 1 0 1 0 0 1 9 9 9 9 1 1 0 0 1 0 1 0 0 0 0 0 0 1 0 0 1 9 9 9 9 0 0 0 0 0 0 1 0 0 0 0 0 0 1 1 0 1 0 0) 0) 1 1 0 1 1 0 1 1 0 represent the ancestral state and pointed teeth the derived one. If such were the case (again, what exactly are the limits of the bistincta group?), then the character would be polarized and is added to our analysis (Tables 2, 3). Duellman & Campbell (1992) did not indicate condition of the maxillary teeth in P. acanthodes, but as specimens of this species were included within the com- posite P. guatemalensis by Duellman (1970), which has spatulate teeth, then those of acanthodes probably are spatulate and we have so coded them. The characteristics for all species in the analysis are given in Table 3. We confined our searches for the most parsimonious phylogenetic hypothesis as closely as possible to the methods used by Duellman & Campbell (1992) in their PAUP analysis. However, our PAUP program (3.0) is a later version than that (2.4) used by Duellman & Campbell. The global branch- swapping option used by Duellman & Campbell is not available on version 3.0; instead a subtree pruning-regrafting (SPR) option is available that is “... approxi- mately, but not exactly, equivalent to the 76 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON glandulosa B glandulosa A tecunumani lacertosa pycnochila & chrysopleura ancestor Fig. 3. acanthodes guatemalensis hartwegi teuchestes Strict and Adams consensus cladogram for the members of the genus Plectrohyla. Character numbers (to the left of the circles) and changes in character states (to the right of the circles) refer to numbers in Table 2. ‘global’ procedure used in earlier versions of PAUP” (Swofford, 1991:31 in user’s manual). Also available is a new procedure, the tree bisection-reconnection (TBR) op- tion. Also new to PAUP 3.0 is the ability to yield polytomies by collapsing branches having a maximum length of zero. Using the data matrix in Table 3, we made search- es using both the SPR and TBR branch- swapping options. Each branch-swapping option was executed first with the zero- length branches collapsed, then executed with the zero-length branches not collapsed. Each search was conducted with the MUL- PARS option in effect, the stepwise addition option closest, and the tree rooted using the outgroup method. Strict consensus and Ad- ams consensus trees were generated for each of the four searches. All eight consensus trees generated were identical, regardless of the search options used. When the zero-length branches were not collapsed, 300 equally parsimonious trees were retained in both the SPR and TBR searches (matrix set at 300, the exact number that can be found is unknown), whereas, when the zero-length branches were collapsed, only two equally parsimonious trees were found under both the SPR and TBR searches. Each of these two trees was identical to its counterpart under the other branch-swapping option. Tree 2 differs from tree 1 (Fig. 3) only by glandulosa B + lacertosa forming a clade by assigning a reversal to character 10, which is missing datum for /acertosa. All trees have 19 steps and a consistency index of 0.684. Thus, even though our data matrix contains an equal number of characters, plus two more taxa than Duellman & Campbell’s matrix, our trees are one step shorter and have a slightly higher consistency index (0.650 in Duellman & Campbell’s trees). More importantly, the unresolved polyto- my of 11 clades for 15 taxa in Duellman & Campbell’s strict consensus tree (their fig. 19) is almost fully resolved (Fig. 3). Whereas our cladogram can undoubtably be im- proved upon by a more thorough analysis VOLUME 107, NUMBER 1 of the genus, it is a substantial improvement over that offered by Duellman & Campbell. Plectrohyla chrysopleura, the species herein described, is the species closest to the an- cestral stock of the genus, based upon our phylogenetic analysis. A more minor concern, but nonetheless irritating, with the Duellman & Campbell (1992) revision are contradictions between some tadpole descriptions and their iden- tifying characteristics in the tadpole key or one of the tadpole illustrations. Duellman & Campbell (1992:4) stated that the pos- terior tooth rows of acanthodes tadpoles are ““_. . Slightly shorter than upper [= anterior] rows, third shortest,’ whereas, in the key we are told in couplet 7, leading to acan- thodes in couplet 9, that the posterior tooth rows are subequal in length. The other part of couplet 7, “Third lower [= posterior] la- bial tooth row shorter than others,” leads to couplet 10 and past acanthodes. Plectro- hyla teuchestes tadpoles supposedly have a “Distinct smooth, sharply raised fold be- tween fringing papillae and anterior tooth row’ (Table 3; also see couplet 2 in tadpole key), whereas Duellman & Campbell’s drawing of the teuchestes tadpole (their fig. 15) indicates that the “‘fold’’ is papillate or scalloped throughout. Also, Duellman & Campbell (1992:fig. 8) illustrate the P. gua- temalensis tadpole with the caudal muscu- lature extending dorsally onto the body nearly to the eyes, quite a remarkable fea- ture. Other statements also are confusing: 1.e., acanthodes tadpoles with “‘two rows of small labial papillae fringing disc, except only one row midventrally” (p. 4) or “Lips having a single row of small fringing papil- lae”’ (couplet 5 leading to acanthodes); and guatemalensis tadpoles reported to have ““one row of small labial papillae anteriorly and laterally, two rows midventrally” (p. 8) but “Lips having two rows of small fringing papillae” (couplet 5 leading to guatemalen- sis). Considering the extent of the Duellman & Campbell inconsistencies, we suggest that anyone trying to use their tadpole key do so 77 with caution. Finally, the specific name pyc- nochila is misspelled throughout Duellman & Campbell’s paper as pychnochila and the number in the second section of tadpole cou- plet 3 leading to couplet 5 is misprinted 6. Acknowledgments McCranie and Wilson are grateful to M. Espinal for assistance in obtaining collecting permits from the office of Recursos Natur- ales Renovables. Field assistance was pro- vided to McCranie by E. Hedl, and J. Porras Orellana provided many courtesies during our stays in Tegucigalpa; to both we are indebted. Literature Cited Adler, K., & D. M. Dennis. 1972. New tree frogs of the genus Hyla from the cloud forests of western Guerrero, México.—Occasional Papers of the Museum of Natural History, The University of Kansas 7:1-19. Altig, R., & G. F. Johnston. 1986. Major character- istics of free-living anuran tadpoles.—Smith- sonian Herpetological Information Service 67: 1-75. Caldwell, J. 1974. A re-evaluation of the Hyla bis- tincta species group, with descriptions of three new species (Anura: Hylidae).— Occasional Pa- pers of the Museum of Natural History, The University of Kansas 28:1-37. Campbell, J. A., & T. M. Kubin. 1990. A key to the larvae of Plectrohyla (Hylidae), with a descrip- tion of the tadpole presumed to be Plectrohyla avia.— The Southwestern Naturalist 35:91—94. Duellman, W. E. 1970. The hylid frogs of Middle America.— Monograph of the Museum of Nat- ural History, The University of Kansas 1:1—753. , & J. A. Campbell. 1984. Two new species of Plectrohyla from Guatemala (Anura: Hyli- dae).—Copeia 1984:390-397. , & 1992. Hylid frogs of the genus Plectrohyla: systematics and phylogenetic rela- tionships.— Miscellaneous Publications Muse- um of Zoology, University of Michigan 181:1- 32. 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. Second ed. Tropical Science Center, San José, Costa Rica, 206 pp. 78 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON McCranie, J. R., J. M. Savage, & L. D. Wilson. 1989. Description of two new species of the E/euthero- dactylus milesi group (Amphibia: Anura: Lep- todactylidae) from northern Honduras.—Pro- ceedings of the Biological Society of Washington 102:483-490. —, & L.D. Wilson. 1986. A new species of red- eyed treefrog of the Hyla uranochroa group (An- ura: Hylidae) from northern Honduras.—Pro- ceedings of the Biological Society of Washington 99:51-55. ,& K.L. Williams. 1989. Anew genus and species of toad (Anura: Bufonidae) with an extraordinary stream-adapted tadpole from northern Honduras.— Occasional Papers of the Museum of Natural History, The University of Kansas 129:1-18. Myers, C. W., & W. E. Duellman. 1982. A new spe- cies of Hyla from Cerro Colorado, and other tree frog records and geographical notes from western Panama.—American Museum Novi- tates 2752:1-32. Savage, J. M., & 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. ——., J.R. McCranie, & L. D. Wilson. 1988. New upland stream frogs of the E/eutherodactylus ru- gulosus group (Amphibia: Anura: Leptodactyl- idae) from Honduras. Bulletin of the Southern California Academy of Sciences 87:50-S6. Smithe, F. B. 1975. Naturalist’s color guide. The American Museum of Natural History, New York, 182 color swatches. Stuart, L. C. 1963. A checklist of the herpetofauna of Guatemala.— Miscellaneous Publications Museum of Zoology, University of Michigan 122:1-150. Swofford, D. L. 1991. PAUP: phylogenetic analysis using parsimony, version 3.0s. Illinois Natural History Survey, Champaign. (LDW) Department of Biology, Miam1- Dade Community College, Kendall Cam- pus, Miami, Florida 33176, U.S.A.; (JRM) 10770 SW 164th Street, Miami, Florida 33157, U.S.A.; (GAC) Departamento de Biologia, Universidad Nacional Autonoma de Honduras, Tegucigalpa, Honduras. PROC. BIOL. SOC. WASH. 107(1), 1994, pp. 79-89 IDENTIFICATION OF THE TAXA XENOCEPHALIDAE, XENOCEPHALUS, AND X. ARMATUS (OSTEICHTHYES: URANOSCOPIDAE) Victor G. Springer and Marie-Louise Bauchot Abstract. — Xenocephalinae Kaup, 1858 (currently recognized as a valid fam- ily), is a junior synonym of Uranoscopidae (dating from at least 1832; no subfamilies recognized). Xenocephalus Kaup, 1858, is a senior synonym of the currently recognized genus Gnathagnus Gill, 1861. Xenocephalus armatus Kaup, 1858, is a senior synonym of the currently recognized species Gnathagnus innotabilis Waite, 1904. A neotype is designated for Xenocephalus armatus, which was erroneously described from New Ireland, but in fact was based on a specimen from New Zealand. Kaup (1858) described a new subfamily, genus, and species of fish (Xenocephalinae, Xenocephalus armatus), which he included in the family Gadidae. Kaup’s taxa have been carried along in the systematic ichthy- ological literature ever since, but their iden- tities and affinities have remained enigmat- ic. It is the purpose of our study to clarify and fix the systematic status of Kaup’s three taxa. Taxonomic History of Kaup’s Taxa Kaup (1858) stated that Xenocephalus ar- matus was distantly related to the Macru- rinae, one of the four subfamilies he rec- ognized in the Gadidae (the others, Gadinae, Brotulinae). Subsequent mention of Kaup’s taxa followed soon after their original de- scriptions, but most authors had nothing substantive to add to his description. We include here, with minimal comment, all references we have encountered in an in- tensive search of the scientific literature for mention of Xenocephalinae (or a family- group based on it), Xenocephalus, or X. armatus. Our purpose in doing so is to dem- onstrate that Kaup’s taxa cannot be consid- ered as nomina oblita, and the genus and species, at least (and the family group, usu- ally) have always, been considered as senior synonyms, albeit of questionable affinities (authors listed chronologically by earliest publication): Bleeker (1859) essentially followed Kaup by listing Xenocephalus in a subfamily Xe- nocephaliformes of a family Gadoidei. Gunther (1862) included Xenocephalus ar- matus, with no mention of Xenocephalinae, as an “Appendix to the Anacanthini ga- doidei,”’ and (1880) stated that Xenocepha- lus was “a gadoid anacanth,” but (1909) presciently opined that it appeared to be a larval form of a fish that is unrelated to the anacanthin gadoids; Gill (1872, 1884, fam- ily listed essentially according to Gunther, 1862); Gill (1888, family ““approximated to Ophidioidea’”’; 1893, family listed under Ophidioidea); Scudder (1882, genus listed); Perrier (1903, genus in Macruridae); Jordan (1905, 1907, 1925, genus included in Zoar- cidae under “the great family Blenniidae’”’; 1919, genus listed; 1923, family included in Blenniiformes); Fowler (1928, family, ge- nus, species recognized); Berg (1940, 1947, 1955, family and genus listed in Blennioi- dei); Neave (1940, genus listed); Schultz (1948, family listed in Blennioidea); Munro (1956, family, genus, species listed; 1967, family, genus, species questionably includ- 80 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ed in Blennioidei); Golvan (1962, genus in- cluded in Macrouridae; 1965, genus listed in Macrouridae, and family and genus listed in Blennioidei); Greenwood et al. (1966, family listed in Blennioidei); Norman (1966), genus and species doubtfully re- ferred to Blennioidea; this long-delayed posthumous publication was essentially complete by 1938 and contains no refer- ences more recent than 1944; Norman in- cluded the statement, erroneous even in 1938, that Xenocephalus had not been rec- ognized since its original description); Romer (1966, family listed in Blennioide1); Gosline (1968, family listed, affinities ques- tionable); McAllister (1968, family listed in Blennioidei); Lindberg (1971, 1974, family included in Blennioidei); Wheeler (1975, 1979, remarks on family, genus, species, “There is every possibility that the only known specimen was a damaged or aberrant specimen of some other fish . . . Validity of family doubtful.”); Nelson (1976, family, genus, species, questionably included in Blennioidea; 1984, family, genus, species listed; quotes V. G. Springer’s opinion that species is possibly larval form of dactylo- pterid, chaetodontid, or scatophagid); Bond (1979, family listed in Blennioidea); Ma- tarese et al. (1984, family listed in Blen- nioidea); Kailola (1987, family, genus, spe- cies incertae sedis); Eschmeyer (1990: 425, family, genus, species, species, “family placement uncertain, based on young’’; page 484, family, genus listed under Suborder Trachinoidei); Springer (1993, family, ge- nus, species probably a dactylopterid). During the course of our literature search, we encountered the descriptions of Xeno- cephalus Wasmann (1887), based on a beetle, and Xenocephalus Leakey (1965), based on a partial skull of a fossil mammal. Both are clearly junior homonyms of Xenocephalus Kaup, and have been provided with re- placement names (see Gentry & Gentry 1978:359). Except for Wasmann (1887), Leakey (1965), Romer (1966), and Gentry & Gentry (1978), we excluded consideration of literature bearing on the junior hom- onyms of Xenocephalus Kaup. Disposition of the Holotype of Xenocephalus armatus Prior to the 1990s, there is no indication that anyone made an attempt to locate the holotype of Xenocephalus armatus or an il- lustration of it, which Kaup (1858) indicat- ed he had published elsewhere. Kaup stated that the specimen was in the Paris Museum and had been sent there by [J.-R.-C.] Quoy and [P.] Gaimard, who had obtained it dur- ing the d’Urville Expedition [= Astrolabe expedition of 1826-1829 under the com- mand of J. S.C. Dumont d’Urville]. Springer (1993) reported that, at his request, M.-L. Bauchot and M. Desoutter of the Muséum National d’ Histoire Naturelle (MNHN) had searched [during January, 1991] the MNHN collection unsuccessfully for the holotype and for information about it among the un- published plates and records of the Astro- labe expedition [not all of which were known to them in 1991]. A second search of the MNHN collection in January, 1993, also was unsuccessful, although many other specimens referred to in an unpublished Quoy manuscript have been located. As we will discuss, information on the specimen was found in the Bibliothéque Centrale, MNHN, among the unpublished descrip- tions (file MS 104), drawings (MS 840), and plates (MS 106) of the Astrolabe fishes. Even though Kaup (1858) stated that the holotype was in the Paris Museum, and he had spent three months working in the fish collection at the museum on two visits dur- ing 1855 and 1856 (Heldmann 1955), he may have actually studied the specimen in London. In J. E. Gray’s preface to Kaup (1856), it is noted that Kaup had specimens from the French, Leyden, Vienna, Frank- furt, Berlin, and Stuttgart museums sent to the British Museum, which Kaup visited several times between 1846 and 1854, so that he could compare them directly with VOLUME 107, NUMBER | British Museum specimens. If the specimen was sent to the British Museum, it appar- ently is not there now (search made of sev- eral parts of the collection by D. Siebert at our request). Under the circumstances we consider the holotype lost. A fortuitous circumstance, discussed later in our study, allows us to replace it with a neotype that conforms in many ways with Kaup’s holotype. Historical Background of Kaup’s Description Kaup (1858) wrote, in German, “This strange form, of which I give a twice-size illustration in my large work, was trans- mitted by Messrs. Quoy & Gaimard, Ex- pedition d’Urville, to the Paris Museum, where it is found under the name of Gren- adier from New Ireland.” Kaup’s memory must have deceived him. He had never published an illustration of the New Ireland grenadier. The large work, to which he referred, was undoubtedly his extensive Das Tierreich (Kaup 1835-1837). In it, he mentioned, but did not illustrate, a species of macrourid to which he gave “grenadiere”’ as part of its common name. It is also possible that Kaup was thinking about the extensive unpublished portion of the manuscript and plates of the Astrolabe expedition. Quoy & Gaimard (1834) pub- lished a study of the fishes obtained by that expedition, but their report, for reasons un- known, includes the descriptions of only 49 species, accompanied by only 12 colored plates, of the large number actually pre- pared: almost 300 species descriptions and at least 120 plates. Although the unpub- lished portions were apparently known to Kaup and other of his contemporaries, their existence has been generally unrecognized for more than 100 years, until recently, when one of us (Bauchot) located them among the archives of the MNHN central library. There is, perhaps, evidence that Kaup derived part of his description from the unpublished 81 Quoy & Gaimard manuscript. In addition to the unpublished finished plates, there are numerous preliminary colored drawings, presumably prepared in the field, upon which the finished illustrations for the plates are based. Identity of Xenocephalus armatus It is important for the determination of the identity of Xenocephalus armatus to demonstrate that Kaup’s description re- ferred to a specimen that was also described and figured earlier by Quoy & Gaimard in their unpublished manuscript. We believe we have located such a description and here present a translation of it followed by a translation of Kaup’s (1858) description. Translation of Quoy & Gaimard [MS 104 (3°™* mémoir:412—413)]; new scientific name here disclaimed and not to be considered available for permanent scientific record or taxonomic purposes [see ICZN Article 8(b)]: Spotted Grenadier Lepidolepous punctatus N. Plate 223, figures 2-3 [Both the preliminary sketches and final fig- ures for Plate 223, figures 2—3 are in color. We present herein, as our Fig. 1, black-and- white reproductions of the preliminary sketches, which we believe are more accu- rate than the final figures.] This fish has an excessively large, bony, boxlike head, quite truncate in front; the mouth is quite large, almost vertical, sit- uated but little on the ventral part of the head; the teeth large and like a card [= fine and set closely in rows]; the large eye of gold color, with a very prominent [bony] orbit dorsally, which has a notch anteriorly. The opercle consists of a mov- able triangular piece. The preopercle has a very prominent posteriorly directed spine, below the opercle, and two other smaller spines more ventrally. One sees four ridges on top of the head. The belly 82 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. of head, mouth at bottom of figure; B, lateral view (original is in color). 1S prominent, round and soft. The rest of the body ends as a point while becoming compressed. A single dorsal fin placed posteriorly, reaches the origin of the cau- dal. The anal fin has the same placement [opposite to the dorsal fin]. The pectoral fins are large and round, as [is] the caudal, which appears, however, pointed when it is collapsed [not displayed]. The lateral line is slightly curved. The ventral fins are exceptionally small, placed very anteri- orly, and almost covered by the gill mem- brane covers, which are exceptionally broad. The general color is deep blue dorsally with some spots of the same color, but slightly darker. The cheeks are a very clear blue, just as the sides. The belly is silver and the fins are whitish. Original preliminary manuscript illustrations of holotype of Xenocephalus armatus: A, dorsal view This fish, 14 lines long [= 31.6 mm; according to Grand Dictionnaire Ency- clopédique Larousse, 1984, Tome 6, p. 6295, 1 ligne = 2.2558 mm; American dictionaries indicate that a line is one- twelfth of an inch, or 2.117 mm], was collected 18 Feb 1827, near the island of Mayor in the bay of Abundance [Bay of Plenty] on the coast of New Zealand [em- phasis ours]. Translation of Kaup.—First Subfamily Xenocephalinae Kp. The abnormally large head is armed with shields and spines. First dorsal fin miss- ing. Second dorsal fin and anal fin slightly separated from caudal. One genus. I. Xenocephalus Kaup. With truncated head to which the body is joined as an VOLUME 107, NUMBER 1 appendage; head and operculum armed. Pectoral and caudal developed. Anus on the posterior half of the body. Tiny teeth in both jaws, none on vomer and pala- tines. Tongue free, thick, almost filling the entire mouth, blunt in front with short tip. Lateral line on the dorsal half of the body and slightly arched [just] following the head. 1) Xenocephalus armatus Kp. The moderately large eye golden yellow, dark spotted below the eyelid [?]. Head shields yellowish brown; the naked skin between them blackish. Body blackish brown with black spots on the dorsum. Belly gold yel- low with gloss. Fins yellowish white. 2nd ID), Ty ANS IO, 1s Zi Wo 85 (CaO, This strange form, of which I give a twice-size illustration in my larger work, was transmitted by Messrs. Quoy & Gai- mard, Exped. d’Urville to the Paris Mu- seum, where it is found under the name Grenadier from New Ireland [emphasis ours]. This subfamily is so far very poor in species and other than the one above I know of no species that belongs to this animal group. It is distantly related to the Macrurinae. [Length of specimen not pro- vided.] Aside from certain differences, which we will discuss, we believe that the similarities between the Quoy & Gaimard and Kaup descriptions, and Kaup’s indication that his specimen was received from Quoy & Gai- mard, are sufficient evidence to conclude that the descriptions were based on the same specimen. We believe that the most important dif- ference between the two descriptions is in the designation of the type localities. Kaup either made a mistake in reporting the type locality as New Ireland (one of the islands in the Bismarck Archipelago northeast of eastern New Guinea) or he was misin- formed about the locality by whoever was responsible for providing him with the in- formation. 83 Other differences between the two de- scriptions are mainly additional characters given by Kaup: the fin-ray counts, lack of vomerine and palatine dentition, shape of the tongue, and position of anus (the last is in agreement with its indication on Quoy & Gaimard’s illustration). Except for the fin- ray counts and putative lack of vomerine and palatine teeth, these characters indicate that Kaup did examine the holotype of Xen- ocephalus armatus. After concluding that the holotype of Xenocephalus armatus is the species de- scribed and figured by Quoy & Gaimard from New Zealand, we searched among the species of fishes known from New Zealand for clues to the identity of XY. armatus. Con- sidering the general gestalt of Quoy & Gai- mard’s illustration, and particularly the ar- mored head and lack ofa spinous dorsal fin, we quickly narrowed the possibilities to the Uranoscopidae. No other family of New Zealand fishes contains species that ap- proach the appearance or description of X. armatus. There are five genera and seven species of uranoscopids reported from New Zea- land (Paulin et al. 1989, Kishimoto 1990, Okamura & Kishimoto 1993): 1—Pleuro- scopus pseudodorsalis Barnard, a deep- dwelling species that also occurs around southern Australia, the southwestern Indian Ocean, and the southeastern Atlantic (Ki- shimoto et al. 1988); 2— Genyagnus mono- pterygius (Schneider in Bloch & Schneider 1801 [Forster MS]), which is endemic to New Zealand (although originally and un- doubtedly erroneously, also reported to oc- cur in Tahiti; see Fowler 1928: 428, as Ane- ma monopterygium, for comment); 3— Gnathagnus innotabilis (Waite), which also occurs in Australia (Kishimoto 1989); 4— Kathetostoma giganteum Haast, which is endemic to New Zealand; 5— Kathetostoma laeve (Schneider in Bloch & Schneider), a deep-water species that is known from the Norfolk Ridge to southern New Zealand, and southern Australia (Kishimoto 1990); 84 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. Neotype of Xenocephalus armatus, NMNZ P.30131, 26.6 mm SI: A, lateral view; B, dorsal view. 6—an unnamed deep-water species of Ka- thetostoma (Paulin et al. 1989, who report- ed two unnamed Kathetostoma species, one of which we assume is K. /aeve, which they did not report), extra New Zealand distri- bution unknown; 7—Selenoscopus turbis- quamatus Okamura & Kishimoto, which is known from off the Kii Peninsula, Pacific coast of central Japan, the Kyushu-Palau Ridge, and the Norfolk Ridge, at depths from 100-510 m (Okamura & Kishimoto 1993). A detailed descriptive account of all New Zealand uranoscopids, including at least fin- ray counts, has not appeared, but we believe that we can narrow our search among those present to the single species identifiable with Xenocephalus armatus. Among all uranoscopids, the deep-dwell- ing genus Pleuroscopus has segmented dor- sal- and anal-fin ray counts closest to those reported by Kaup for Xenocephalus arma- tus (as few as 9 and 10, respectively, in Pleu- roscopus, Kishimoto et al. 1988). Even so, Pleuroscopus can be excluded from consid- eration because it has a series of 8 to 11 tubercule-like spines preceding the seg- VOLUME 107, NUMBER 1 mented-ray portion of the dorsal fin. It is also notable that very small specimens of Pleuroscopus are unknown. Kishimoto et al. (1988) reported that specimens smaller than 242 mm SL are “‘currently unobtainable.” In view of the absence of small specimens, and the apparently restricted deep-water habitat of the species, it seems unlikely that the Astrolabe would have acquired a juve- nile 31 mm TL. Genyagnus monopterygius, although commonly available at small sizes, can be excluded from consideration for several rea- sons: it has an obvious mental barbel and lingual lure (both noticeable in specimens at least as small as 27 mm TL); the head is not noticeably enlarged nor heavily ar- mored in specimens within the size range of the holotype of Xenocephalus (no large preopercular spines as in Xenocephalus); its eyes are located on top of the head (on side of head dorsally in Xenocephalus); the color pattern dorsally on the body of small ju- veniles consists of a dark stripe-like mark- ing (no large dark spots, but numerous pale spots in adults); the pectoral-fin rays num- ber only 16-18 (21 reported by Kaup for Xenocephalus); and the dorsal fin rays, 18 or 19, appear to be too numerous even if mistakes in the counts were made by Kaup. Kathetostoma (generically), of which small specimens are often collected, can be ex- cluded from consideration because even at small sizes it has a conspicuous, elongate humeral spine and all spines on the ventral border of the preopercle are relatively fine and ventrally directed; furthermore, the col- or pattern dorsally on the body does not consist of large dark spots in any specimens we have seen. No humeral spine is indicated in Quoy & Gaimard’s figure of Xenocepha- lus armatus, and the posteriormost preoper- cular spine is greatly enlarged and posteri- orly directed. We believe that Xenocephalus armatus is conspecific with Gnathagnus innotabilis Waite (1904), described from New South 85 Wales, but currently recognized as also oc- curring in New Zealand waters (Paulin et al. 1989). We are fortunate to have an ap- proximately 31.0 mm TL specimen (small terminal portion of caudal fin now broken off) of G. innotabilis from New Zealand (Fig. 2a, b) to serve as a basis for comparison. The large and heavily armored head of the specimen, including the long posteriorly di- rected preopercular spine with two smaller spines ventrally on the preopercle, and large dark spots dorsally on the body strongly cor- roborate the conspecificity of the two spe- cies. There are differences between the descrip- tion of X. armatus and characters exhibited by G. innotabilis, particularly in the denti- tion and dorsal- and anal fin-ray counts giv- en by Kaup (1858). We believe these dif- ferences are due to the inadequacy of the optical equipment available during the 1850s. Small specimens of G. innotabilis that we have examined have a few tiny, incon- spicuous teeth on the vomer and palatines (we even overlooked these in our initial ex- amination of our 31 mm specimen). The species has 11 or 12 dorsal-fin rays and 16 anal-fin rays, 4 and 6 rays more than re- ported for Xenocephalus (additionally, the last dorsal-fin ray may be simple or split to the base in G. innotabilis; we counted either condition as one ray). We can discern 11 dorsal-fin rays and 8 anal-fin rays, however, from Quoy & Gaimard’s illustration, as op- posed to the 7 and 10, respectively, of Kaup’s description. Using a Leitz widefield stereo microscope RS (ES model has same optics), which affords the finest resolution of any dissecting scope we know, it was only with difficulty that we were able to make accurate counts of dorsal- and anal-fin rays on our 31 mm specimen. We verified these counts with counts made from radiographs of the specimen. It is because of this difficulty that we believe Kaup’s dorsal- and anal-fin ray counts were in error. All adult uranoscopids have essentially 86 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON the same number of branched caudal-fin rays (10), but branching is not evident in small specimens we examined. The number of segmented nonbranched rays appears to in- crease with growth, but probably not the total number of caudal-fin elements (also includes procurrent rays and spines). The dorsal- and ventralmost of the procurrent elements decrease considerably in size se- rially as one progresses anteriorly, and are difficult to count accurately in small speci- mens, unless, as we presume, the specimens are cleared and stained. We were unable to make an accurate count of the total number of caudal-fin elements in our 31 mm spec- imen, but a total count of 20, as given by Kaup for Xenocephalus, for all caudal-fin elements is probably slightly less than the number in G. innotabilis. In order to eliminate the possibility that the New Ireland type locality might actually be correct, we attempted to identify Xeno- cephalus armatus with a species from that area, given the additional information pro- vided by the Quoy & Gaimard manuscript. In that, we were unsuccessful in identifying Xenocephalus armatus with any fish species known from the New Guinea area, partic- ularly political Papua New Guinea, which includes New Ireland. Additionally, only two genera of uranoscopids are known from New Guinea: Uranoscopus Linnaeus and Ichthyoscopus Swainson (Kailola 1987). Uranoscopus has a conspicuous spinous dorsal fin and Ichthyoscopus lacks spinous processes on the ventral margin of the pre- opercle (Pietsch 1989), characters that ex- clude both genera as possibly being conge- neric with Xenocephalus. Designation of Neotype for Xenocephalus armatus In order to fix Kaup’s species, we here designate our 31 mm specimen (26.6 mm SL from midtip of upper lip to caudal-fin base), NMNZ P.30131 (formerly USNM 325034) as neotype of Xenocephalus ar- matus Kaup. Dorsal-fin rays 11, last ray split to base; anal-fin rays 15, last ray split to base; pectoral-fin rays (r/1) 20/21; vertebrae 27; 7 nonelement-bearing pterygiophores (predorsals) anterior to pterygiophore sup- porting first dorsal-fin ray. Tiny, widely spaced teeth present on vomer and pala- tines. The posteriorly projecting preopercle spine on each side of the head is broken, but the ends are still attached. The neotype was obtained from the col- lection of the Fisheries Laboratory in Wel- lington, New Zealand, by G. D. Johnson, who brought the specimen to our attention and informed us that there was no other data associated with the specimen. Taxonomic Consequences of the Identity of Xenocephalus armatus The identification of Xenocephalus ar- matus Kaup, 1858, has the following con- sequences: Xenocephalinae Kaup, 1858, becomes a junior synonym of Uranoscop- idae (dating at least as early as Bonaparte 1832—as Uranoscopini—and Richardson 1848:i1v—as Uranoscopidae); Xenocephalus Kaup, 1858, is a valid senior synonym for a genus of Uranoscopidae, taking priority over Gnathagnus Gill, 1861, and its junior synonyms (see Pietsch 1989:294); and X. armatus Kaup, 1858, is a valid senior syn- onym for a species of Xenocephalus, taking priority over Gnathagnus innotabilis Waite, 1904. Comparative Material (Institutional abbreviations follow those in Leviton et al., 1985.) Astroscopus y-graecum. — Louisiana: Four Bayou Pass, USNM 185647 (1 specimen: ca. 41 mm SL); Barataria Bay, USNM 187947 (2: 30.9-35.2); off coast, USNM 156863 (1: 38.4). Genyagnus monopterygius.—New Zea- land: Nukumaru Reef, Wanganui, NMNZ VOLUME 107, NUMBER | P.10463 (3 specimens: ca. 40-48 mm SL); Mangakino Channel, Pourerere, NMNZ P.26278 (2: 22.2—28.2); Hauraki Gulf, NMNZ P.21876 (2: 34.8-59.1); Castle- point, NMNZ P.17311 (1: 22.1); Manakau Harbour, NMNZ P.2457 (1: 16.6); Toko- maru Bay, NMNZ P.2038 (2: 24.3-24.7); Old Wharf, Kaikoura, NMNZ P.25700 (1: 29.6); Port Hardy, D’Urville Island, NUNZ P.5332 (1: 21.4). Ichthyoscopus lebeck?.—Indonesia: Am- bon, USNM 325474 (1 specimens: 117 mm SL). Kathetostoma giganteum. —New Zea- land: off Cape Farewell, NMNZ P.16605 (1 specimen: 54.8 mm SL); Oamaru, NUNZ P.10684 (1: 78.5); Dunedin, USNM 318371 (1: 84.9) Kathetostoma sp.— Off Caribbean Pana- ma: 9°18’N, 80°35'W, USNM 187907 (2 specimens: 41.0-64.4 mm SL). Pleuroscopus pseudodorsalis. —New Zea- land: NMNZ P.19668 (1 specimen: ca. 325 mm SL), P.20151 (2: ca. 335, 350), P.22102 (1: ca. 295), P.27963 (1: ca. 270). Uranoscopus spp.— Australia: New South Wales: off Clarence River Country, AMS 1.32120005 (1 specimen: 54.7 mm SL); Col- laroy Beach, Sydney, AMS IB.4119 (1: 17.5); off Newcastle, AMS 1.33445004 (1: 53.1); Tweeds Head Country, AMS 1I.23687009 (1: ca. 29). Queensland: N of Townsville, AMS 1I.25837002 (3:52.3-52.5). Xenocephalus armatus (as Gnathagnus innotabilis).— Australia: New South Wales, Montague Island, AMS IB.1298 (4 speci- mens: 12.8-16.0 mm SL). New Zealand: NMNZ P.23224 (1: 57.8 mm SL); Bay of Plenty, NMNZ P.11115 (1:48.2); off Taur- aga Harbor, NMNZ 16118 (1: 104). Stom- ach of bluefin tuna, AMS I.B1297 (1: 30.3). Xenocephalus elongatus. —Japan: Suruga Bay, USNM 296634 (1: 105 mm SL). Phil- ippines: Romblon Island, USNM 122528 GE75.9): Xenocephalus egregius.—Texas: Gulf of Mexico off Padre Island, USNM 268445 (1 specimen: 72.0 mm SL). 87 Acknowledgments We extend our appreciation to: W. N. Es- chmeyer and C. Ferraris (CAS), who fur- nished information on the oldest known dates for a family-group name based on Uranoscopus; C. E. Ray (Paleobiology, USNM) and A. Gentry (formerly BMNH) for information on junior homonyms of Xe- nocephalus; G. Duhanmel and M. Desout- ter (NMHN), C. D. Roberts (NMNZ), M. F. Gomon (NMV), and J. Leis and M. McGrouther (AMS), who made specimens available from their institutions; G. D. Johnson (USNM), who called our attention to the specimen we designated as a neotype; T. M. Orrell (USNM) for radiography of specimens; S. Raredon and L. Palmer (USNM) for processing incoming loans. We thank M. Ducreux, Directeur de la Biblio- théque centrale, MNHN, and F. Serres, Conservateur, who authorized the exami- nation and publication of the Quoy & Gai- mard manuscript information, and G. Du- hamel, who authorized preparation of the photographs (Fig. la, b) of the manuscript figures of the holotype of Xenocephalus ar- matus. The photographs of the neotype (Fig. 2a, b) were taken by T. B. Griswold, who also rendered the black-and-white prints of Fig. 1 from a color slide. We are especially indebted to H. Kishimoto, for his excellent review of the submitted manuscript and calling our attention to several errors of omission. Literature Cited Berg, L.S. 1940. [Classification of fishes, both recent and fossil].—Travauz de l'Institut Zoologique de l’Académie Des Sciences de ’URSS 5(2):87-— 345 [in Russian]. . 1947. Classification of fishes, both recent and fossil. J. W. Edwards, Ann Arbor, Michigan, pp. 87-517. [Berg (1940) reprinted and translated into English] 1955. [Systematics of fishes, recent and past].—Trudy Zoologicheskogo Instituta 20:1— 286 [in Russian; German translation published in 1958]. Bleeker, P. 1859. Enumeratio Specierum Piscium 88 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON hucusque in Archipelago Indico.— Verhandelin- gen der Natuurkundige Vereeniging in Neder- landsch-Indié [also as Acta Societatis regiae scientiarum Indo-Neerlandicae] 6(1):i—xxxvi, 1— 276. Bonaparte, C. L. 1832. Saggio d’una distribuzione metodica degli animali vertebrati a sangue fred- do, 86 pp. Bond, C.E. 1979. Biology of fishes. W. B. Saunders, Philadelphia, 514 pp. Eschmeyer, W. N. 1990. Catalog of the genera of Recent fishes. California Academy of Sciences, San Francisco, 697 pp. Fowler, H. W. 1928. The Fishes of Oceania.—Mem- oirs of the Bernice P. Bishop Museum 10:1—540, 49 pls. Gentry, A. W., & A. Gentry. 1978. Fossil Bovidae (Mammalia) of Olduvai Gorge, Tansnia.—Bul- letin of the British Museum (Natural History), Geology 29:289-446. Gill, T. 1861. Synopsis of the uranoscopoids.—Pro- ceedings of the Academy of Natural Sciences of Philadelphia 1861:108-117. 1872. Arrangement of the families of fish- es.—Smithsonian Miscellaneous Collections 247:xlvi + 49 pp. 1884. On the anacanthine fishes. — Proceed- ings of the Academy of Natural Sciences of Phil- adelphia 1884:167-183. . 1888. Sub-Order VIII Acanthopterygii in The Riverside Natural History, J. S. Kingsley, ed. 3:18 1-262. 1893. Families and subfamilies of fishes.— Memoirs of the National Academy of Sciences 6:127-138. Golvan, Y.-J. 1962. Repertoire systematique des noms de genres de vertébrés (suite). — Annales de Pa- rasitologie Humaine et Comparée 37(5—6):870— 997. 1965. Catalogue systematique des noms de genres de poissons actuels. Masson et Cie, Paris, 227 pp. Gosline, W. A. 1968. The suborders of perciform fishes.— Proceedings of the United States Na- tional Museum, 124(3647):1-78. Greenwood, P. H., D. E. Rosen, S. H. Weitzman, & G. S. Meyers. 1966. Phyletic studies of tele- ostean fishes, with a provisional classification of living forms.— Bulletin of the American Mu- seum of Natural History 131(4):341-455. Ginther, A. 1862. Catalogue of the fishes in the Brit- ish Museum 4:xxi + 534 pp. . 1880. The study of fishes. Adam and Charles Black, Edinburgh, xv + 720 pp. 1909. Andrew Garrett’s Fische der Stidsee, 8.— Journal des Museum Godeffroy 16:1—2, 261-— 388, pls. 141-160. Heldmann, G. 1955. Johan Jakob Kaup. Darmstadt, i-x, 1-28. [Copy of pamphlet available in Smithsonian Institution Library] Jordan, D.S. 1905. Guide to the study of fishes 2:xxii + 599 pp. 1907. Fishes. H. Holt and Company, New York, 789 pp. 1919. The genera of fishes. Part II. Leland Stanford Junior University Publications, Uni- versity Series, i-ix + 163-184 + i-xiii. . 1923. Aclassification of fishes including fam- ilies and genera as far as known. —Stanford Uni- versity Publications, University Series, Biolog- ical Sciences 3(2):77—243 + i-x. 1925. Fishes. Revised. D. Appleton and Company, New York, 773 pp. Kailola, P. J. 1987. The fishes of Papua New Guinea, a revised and annotated checklist 2.— Papua New Guinea, Department of Fisheries and Marine Resources, Research Section, Research Bulletin 41:i—xxxi, 1-194. Kaup, J. 1835-1837. Das Tierreich in Seinen Haupt- formen Systematisch Beschrieben. 3 volumes, 1275 pp. 1856. Catalogue of the lophobranchiate fish in the collection of the British Museum, Lon- don, iv + 80 pp., 4 pls. 1858. Uebersich der Familie Gadidae. —Ar- chiv fiir Naturgeschichte 24(1):85—93. Kishimoto, H. 1989. A new species and a new genus of the stargazer genus Gnathagnus from north- western Australia. — Japanese Journal of Ichthy- ology 36(3):303-314. 1990. Uranoscopidae. Pp. 297-300 in K. Amaoka, K. Matsuura, T. Inada, M. Takeda, H. Hatanaka, & K. Okada, eds., Fishes collected by the R/V Shinkai Maru around New Zealand. Japan Marine Fishery Research Resource Cen- ter, 410 pp. , P. R. Last, E. Fujii, & M. F. Gomon. 1988. Revision of a deep-sea stargazer genus Pleuro- scopus. — Japanese Journal of Ichthyology 35(2): 150-158. Leakey, L. S. B. 1965. Olduvai gorge, 1951-1961. Cambridge University Press, Volume 1:xiv + 118 pp. Leviton, A. E., R. H. Gibbs, Jr., E. Heal & C. E. Daw- son. 1985. Standards in herpetology and ich- thyology: part I. Standard symbolic codes for institutional resource collections in herpetology and ichthyology.—Copeia 1985(3):802-832. Lindberg, G. U. 1971. [Families of the fishes of the world, a check list and a key]. Akadmiya Nauk SSSR, 471 pp. [in Russian] 1974. Fishes of the world, a key to families and a checklist. John Wiley & Sons, 545 pp. [English translation of 1971 publication] VOLUME 107, NUMBER 1 Matarese, A. C., W. Watson, & E. G. Stevens. 1984. Blennioidea: development and relationships. — American Society of Icthyologists and Herpe- tologists, Special Publication 1:565-573. McAllister, D. E. 1968. Evolution of branchiostegals and classification of teleostome fishes. — Nation- al Museum of Canada, Bulletin 221:xiv + 1l- 239. Munro, I. S. R. 1956. The fishes of the New Guinea region.—Papua and New Guinea Agricultural Journal 10(4):97—369 (reissued 1958 with same pagination as Territory of Papua and New Guin- ea Fisheries Bulletin 1). . 1967. The fishes of New Guinea. Department of Agriculture, Stock and Fisheries, Port Mores- by, xxxvil + 651 pp. Neave, S. A. 1940. Nomenclator zoologicus 4:758 pp. Published for the proprietors by the Zoo- logical Society of London, London. Nelson, J. S. 1976. Fishes of the world. John Wiley & Sons, xiv + 415 pp. 1984. Fishes of the world, second edition. John Wiley & Sons, xvii + 523 pp. Norman, J. R. 1966. A draft synopsis of the orders, families and genera of Recent fishes and fish- like vertebrates. Trustees of the British Museum (Natural History), 649 pp. Okamura, O., & H. Kishimoto. 1993. Selenoscopus turbisquamatus, a new genus and species of ur- anoscopid fish from Japan and the Norfolk Ridge. — Japanese Journal of Ichthyology 39(4): 311-317. Paulin, C., A. Stewart, C. Roberts, & P. McMillan. 1989. New Zealand fish, a complete guide.— National Museum of New Zealand Miscella- neous Series 19:xiv + 279 pp. Perrier, E. 1903. Traité de Zoologie, Masson, Paris, 4:2653-2726. Pietsch, T. W. 1989. Phylogenetic relationships of trachinoid fishes of the family Uranoscopidae. Copeia 1989(2):255-303. Quoy, J. R. C., & P. Gaimard. 1834. Voyage de Deé- couvertes de “L’Astrolabe,” Exécuté par Ordre 89 du Roi, pendant les Années 1826-29, sous le Commandement de M.J. Dumont d’Urville, 3, Poissons, pp. 647-720, 20 pls. Richardson, J. 1844-1848. Ichthyology of the voyage of H.M:S. Erebus & Terror, under the command of Captain Sir James Clark Ross, R.N., F.R:S., i-viii, 1-139, 39 pls. Romer, A. S. 1966. Vertebrate paleontology. Third Edition. University of Chicago Press, ix + 460 pp. Schultz, L. P. 1948. The ways of fishes. D. Van Nos- trand Co., New York, xii + 264 pp. Scudder, S. H. 1882. Nomenclator zoologicus, 2, uni- versal index to genera in zoology.— Bulletin of the United States National Museum 19:339 pp. Springer, V.G. 1993. Definition of the suborder Blen- nioidei and its included families (Pisces: Per- ciformes).— Bulletin of Marine Science 52(1): 472-495. Waite, E.R. 1904. New records or recurrences of rare fishes from eastern Australia. No. 3.—Records of the Australian Museum 5(4):231—244, pls. 25-26. Wasmann, E. 1887. Neue brasilianische Staphylini- den, bei Eciton hanatum gesammelt von Dr. W. Miieller.—Entomologische Zeitschrift 31:403— 416. Wheeler, A.C. 1975. Fishes of the world. Macmillan Publishing Co., Inc., New York, xiv + 366 pp., 96 pls. . 1979. The world encyclopedia of fishes. Mac- donald, London, xiv + 368 pp. 96 pls. (VGS) Division of Fishes, Department of Vertebrate Zoology, National Museum of Natural History, Washington, D.C. 20560, U.S.A.; (MLB) Laboratoire d’Ichtyologie, Muséum national d’Histoire naturelle, 43 Rue Cuvier 75231, Paris Cedex 05, France. PROC. BIOL. SOC. WASH. 107(1), 1994, pp. 90-96 CREAGRUTUS MELASMA, A NEW SPECIES OF CHARACID FISH (TELEOSTEI: CHARACIFORMES) FROM UPLAND STREAMS OF NORTHERN VENEZUELA Richard P. Vari, Antony S. Harold, and Donald C. Taphorn Abstract. — Creagrutus melasma, new species, occurs in a band across the northern headwaters of the Rio Orinoco from Tachira state in the west to Monagas in the east. The species is also known from several Caribbean versant drainages, including the Rios Tuy and Neveri. This small-bodied species, which barely exceeds 40 mm SL, is distinguished from congeners by its dorsal-fin and humeral pigmentation as well as a combination of meristic characters. Resumen. —Se describe una nueva especie de Creagrutus que habita una franja en el piedmonte norte de las cuencas altas afluentes del Rio Orinoco desde el estado Tachira en el oeste hasta Monagas en el este. La especie es también conocida desde varias cuencas del Mar Caribe, incluyendo los rios Tuy y Neveri. Esta especie tiene un cuerpo pequeno que raramente excede los 40 mm de longitud estandar. Se distingue de sus congéneres por la pigmentacion en la aleta dorsal, la forma de la mancha humeral, y la combinacion de varias caracteres meristicas. The presence of Creagrutus Gunther in Lago de Valencia and adjoining drainages of northern Venezuela was first noted by Eigenmann (1920:12). Eigenmann, fol- lowed by Pearse (1920:11), identified these populations as C. beni, a species described by Eigenmann (1911:172) from the Rio Beni, northeastern Bolivia. Although Eigen- mann (1920:12) noted the darkly pigmented dorsal fin in material from Lago de Valencia and the Rio Bue, a feature absent in C. beni, he considered the pigmentation a juvenile condition. The absence of such pigmenta- tion in juveniles from elsewhere in the range of C. beni (see Eigenmann 1927:421—423) was not discussed. The Lago de Valencia Creagrutus popu- lations with dark dorsal-fin pigmentation, described herein as C. melasma, were in- dependently recognized as distinct from all congeners by two of the authors (RPV & ASH) during their revisionary study of the genus and by the third author (DCT) in the course of research on fishes of the Rio Apure basin (Taphorn 1992:173-175). The ongo- ing review of Creagrutus by the first two authors indicates that the new species has distinctive derived pigmentation patterns of the dorsal fin and humeral mark and is fur- ther separable from congeners on the basis of a combination of various meristic and morphometric characters. It is broadly dis- tributed in northern headwaters of the Rio Orinoco and several Caribbean drainage ba- sins, being particularly common in the Lago de Valencia basin and northern headwaters of the Rio Apure. It is described herein to make its name available for an ongoing phy- logenetic study (RPV & ASH) of Creagrutus and putatively related taxa. Materials and methods.—Specimens are deposited in the following collections: Academy of Natural Sciences of Philadel- phia (ANSP); California Academy of Sci- ences (CAS), including former Indiana Uni- versity collections (IU); Illinois Natural VOLUME 107, NUMBER 1 Fig. 1. Parque Nacional Guatopo, Rio Orituco, first bridge along road from Santa Teresa to Altagracia. History Survey, Champaign (INHS); Museo de Biologia, Universidad Central de Ven- ezuela, Caracas (MBUCV); Museo de Cien- cias Naturales, Universidad Nacional Ex- perimental de los Llanos Occidentales Ezequiel Zamora, Guanare, Venezuela (MCNG); Museo de Historia Natural La Salle, Caracas (MHNLS), and National Mu- seum of Natural History, Smithsonian In- stitution (USNM). Counts and measure- ments were taken following methods outlined in Harold & Vari (1994). Range of standard lengths (in mm) of specimens mea- sured for meristic and morphometric data are cited, followed by the number of spec- imens measured. Creagrutus melasma, new species Fig. 1 Creagrutus beni (not of Eigenmann 1911).— Eigenmann 1920:12 (Venezuela, Lago de Valencia and adjoining rivers; specimens from Maracay, IU 15133, and Isla del Buro (=Isla El Burro), IU 15134).— Pearse 1920:12, 24, 25, 43 (Venezuela, Lago de Valencia; food items and parasites; spec- imens served as basis for Eigenmann 1920 and 1927 citations).—Eigenmann 1927: 422 (in part; specimens from Venezuela, 91 Creagrutus melasma, new species, holotype, MBUCV V-22198, 32.4 mm; Venezuela, Estado Guarico, Maracay, IU 15133, and Isla del Buro (=Isla El Burro), ITU 15134). Holotype. —MBUCV V-22198, 32.4 mm, Venezuela, Estado Guarico, Parque Na- cional Guatopo, Rio Orituco, first bridge along road from Santa Teresa to Altagracia; collected by H. Moreno and A. Machado- Allison, 20 May 1992. Paratypes. — All collected in Venezuela: Estado Aragua: MBUCV-V21257, 56 specimens; 34.2—40.7 mm (3), CAS 79622, 10 specimens; Rio Tuy basin, mouth of Rio Cagua, ~10 km from Guayes, collected by R. Royero et al., 6 Apr 1991. Estado Carabobo: ANSP 134171, 48 specimens; 33.5—34.2 mm (3), Rio Guacara basin, Rio Vigirima, ~ 10 km NNW of Gua- cara (~10°24’N, 67°55'W), collected by N. R. Foster et al., 30 Nov 1966. INHS 60021, 10 specimens; 33.2—36.0 mm (3), Lago de Valencia basin, Vigirima, Rio Las Penitas (~10°20'N, 67°52'W), collected by D. C. Taphorn et al., 29 Nov 1990. MHNLS 503, 6 specimens; Lago de Valencia, at Guata- paro dike, W of Valencia. MCNG 15354, 56 specimens; Rio Manuare, collected by D.C. Taphorn, 19 Oct 1985. MCNG 15354, 56 specimens; Rio Manuare, about 16 km along river from Manuare (09°59'N, 92 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 67°45'10”W), collected by C. Olds et al., 19 Oct 1985. MCNG 24622, 34 specimens; Cano La Camarca, N of San Diego, col- lected by D. C. Taphorn, 29 Dec 1990. Estado Guarico: MBUCV V-24020, 2 specimens; 32.0—33.9 mm (2), collected with holotype. Estado Yaracuy: USNM 219615, 3 spec- imens; 22.8-36.3 mm (3), USNM 219616, 1 specimen; 31.8 mm, Rio Cojedas basin, Quebrada Grande, between Nirqua and Chivacoa, collected by F. Mago-Leccia, 19 May 1978. Non-type specimens examined. —(all col- lected in Venezuela): Estado Anzoategui: MBUCV V-15444, 1 specimen; Quebrada Las Minas, tributary of Rio Querecual. Estado Apure: MCNG 10302, 19 speci- mens; Cano Naporal, tributary of Rio Por- tuguesa. ANSP 165139, 1 specimen; Rio Capanaparo, ~5.0 km downstream from crossing of highway between San Fernando de Apure and Puerto Paez (7°02’N, 67°25'W). Estado Aragua: CAS 69297 (formerly IU 15133), 75 specimens; Rio Bue at Maracay. MBUCV V-3045, Lago de Valencia basin, Rio Limon, east of I.N.A. (Agricultural Re- search Institute). MCNG 14201, 3 speci- mens; Rio Pao, near La Candelaria. Estado Barinas: MCNG 5271, 402 spec- imens; MCNG 5401, 9 specimens; cano at entrance to Bocono dam. MCNG 8877, 8 specimens; Rio Bocono at dam site. MCNG 5648, 12 specimens; Rio Tucupido at Las Canoas. MCNG 6347, 2 specimens; Cano Musao. MCNG 11944, 4 specimens; Cano Las Maravillas. MCNG 6546, 2 specimens; cano at Estero Chiguira. Estado Bolivar: MHNLS 7240, 2 speci- mens; Caicara. Estado Carabobo: CAS 69294 (formerly IU 15134), 266 specimens; Lago de Valen- cia, Isla El Burro. MHNLS 5882, 1 speci- men; Lago de Valencia, Muelle Nuevo, op- posite Isla El Burro. INHS 60446, 35 specimens; 32.2—35.0 mm (5), Lago de Va- lencia basin, Cano la Cumara, 3 km N of San Diego (10°16.55'N, 67°56.21'W). MBUCYV uncat., 6 specimens; Lago de Va- lencia. MBUCV V-9919, 1 specimen; Rio Onoto, Puente Onoto, about 40 km from San Carlos. MCNG 15281, 5 specimens; Rio Chirigu, tributary of Rio Pao. MCNG 15295, 7 specimens; cano near Belen. MCNG 15342, 60 specimens; Cano Gua- mita. MCNG 24647, Rio Las Penitas at Vi- . girima. _ Estado Cojedas: MCNG 6786, 2 speci- mens; Quebrada Camoruco. MCNG 13780, 6 specimens; Rio Chorreron, 10 km from Apartaderos. MHNLS 2502, 1 specimen; Quebrada Guabinas, at highway from San Carlos to Acarigua. MHNLS 499, 6 speci- mens; Rio Portuguesa basin, Rio Manrique, 2 km upstream of Manrique. MHNLS 520, 10 specimens; Rio Portuguesa basin, Que- brada Tierra Caliente, 5 km W of Manrique. Estado Miranda: MCNG 14296, 3 spec- imens; at bridge near Araguita. Estado Monagas: MBUCV V-9753, 1 specimen; Rio Caripe, Sector Salle, on the Las Parcelas Road, 6 km from Carripito. MCNG 16977, 3 specimens; Rio Cocoyal. MAHNLS 517, 2 specimens; quebrada N of San Francisco de Maturin. MHNLS 527, 9 specimens; Rio Colorado at San Antonio de Maturin. MHNLS 8064, 1 specimen; Rio Aragua, at road from Maturin to Quiriquire, about 10 km from Maturin (63°25’W). MHNLS 8879, | specimen; Rio Aragua, 10 km from Aragua de Maturin, at road from Maturin to Quiriquire. MHNLS 9437, 16 specimens; Distrito Acosta, Rio Caripe ba- sin, Embalse El Guamo. Estado Portuguesa: MCNG 122, 1 spec- imen; Cano La Lora, tributary of Rio Tucu- pido. MCNG 2443, 9 specimens; tributary of Rio Tucupido. MCNG 8835, 3 speci- mens; Rio Tucupido at dam site. MCNG 9215, 2 specimens; Rio Tucupido, Los Hierros. MHNLS 6361, 2 specimens; Rio Tucupido, Los Hierros, 7 km N of Tucu- pido. MHNLS 2678, 6 specimens; cano N of Tucupido, 6 km along road to Los Hie- VOLUME 107, NUMBER 1 rros. MCNG 19303, 3 specimens; MCNG 19638, 1 specimen; MCNG 19798, 6 spec- imens; Embalse Tucupido. MCNG 10666, 6 specimens; cano tributary of Rio Bocono, near dam. MCNG 703, 8 specimens; trib- utary of Rio Bocono, upstream from Puerto Paez. MCNG 124, 1 specimen; cano at road from Chabasquen to Barquismeto, tributary of upper Rio Guanare. MCNG 10858, 5 specimens; Cano Buchi, between Acarigua and Guanare. MCNG 11261, 1 specimen; Rio Las Marias, bridge on Highway 5. MCNG 13314, 1 specimen; MCNG 19769, 1 specimen; Rio Las Marias. MCNG 11616, 5 specimens; Rio Are, at bridge on Highway 5 between km 227 and 228. MCNG 11842, 1 specimen; Cano Bombicito, near Apari- cion. MCNG 15380, 14 specimens; Cano Volcan. MCNG 16727, 1 specimen; Cano San Rafael, at km 247 on Highway 5S. MCNG 18737, 5 specimens; Rio Saguas. Estado Sucre: MBUCV V-15419, 2 spec- imens; Rio Neveri, at road to Turimiquire, near Cambural. MBUCV V-15423, 3 spec- imens; Rio Neveri, at road to Turimiquire, near Paraparo. MBUCV V-15428, 4 spec- imens; Rio Neveri, near Paraparo. MBUCV V-15405, 2 specimens; Rio Neveri, Que- brada Carrasposo. MBUCV V-15451, 3 specimens; Cano Cruz de Agua. MCNG 17051, 1 specimen; Cano Juan Antonio. Estado Tachira: MCNG 6484, 4 speci- mens; tributary of Rio Quinimari. MCNG 6626, 3 specimens; cano tributary to Rio Chururu. MCNG 11661, 21 specimens; Cano Toronduy, at bridge on San Cristobal road. MCNG 11790, 1 specimen; Rio San Agaton. Diagnosis. — Creagrutus melasma has the unique jaw structure and premaxillary den- tition typical of Creagrutus. Form of the humeral mark in this species is unique with- in Creagrutus (see ‘““Remarks”’ below), darkest immediately dorsal to lateral line, vertically elongate, and oriented vertically to obliquely from anteroventral to postero- dorsal. Distinctive large, dark spot on an- terior portion of dorsal fin. Other characters 93 Table 1.—Morphometric and meristic features of holotype and ranges of values for 20 paratypes of Crea- grutus melasma, new species. Standard length is ex- pressed in mm; measurements | to 14 are percentages of standard length; 15 to 17 are percentages of head length. Holo: type Paratypes Morphometrics Standard length 32.4 22.8-40.7 1. Snout to anal-fin origin 61.4 61.9-66.7 2. Snout to pelvic-fin origin 47.7 46.8-51.2 3. Snout to pectoral-fin origin 26.5 24.3-27.2 4. Snout to dorsal-fin origin 49.8 48.4-53.8 5. Dorsal-fin origin to hypural joint 56.9 53.3-57.7 6. Dorsal-fin origin to anal-fin or- igin 31.3 30.7-36.8 7. Dorsal-fin origin to pelvic-fin origin 30.2 29.0-35.2 8. Dorsal-fin origin to pectoral-fin origin 34.5 32.5-39.0 9. Caudal peduncle depth 11.9 11.5-13.1 10. Pectoral-fin length 21.3 18.7-21.5 11. Pelvic-fin length 15.9 14.8-16.7 12. Dorsal-fin length 25.7 21.4-28.5 13. Anal-fin length 21.3 18.6-21.6 14. Head length 26.4 24.5-26.9 15. Postorbital head length 43.8 42.4-47.8 16. Snout length 28.3 24.0-29.0 17. Bony orbital diameter 32.8 30.9-36.2 18. Interorbital width 31.9 29.3-34.7 Meristics Lateral line scales 35 34-36 Scale rows between dorsal-fin origin and lateral line 5 5-6 Scale rows between anal-fin origin and lateral line 4 4-5 Predorsal median scales 10 10-11 Branched dorsal-fin rays 8 7-8 Branched anal-fin rays 11 10-12 Branched pelvic-fin rays a] 7 Pectoral-fin rays 10 10-12 Vertebrae 35 34-36 which, in combination, serve to distinguish the species are small body size; short, stout gill-rakers, those of ceratobranchial and epi- branchial about equal in length; and 5 teeth in main premaxillary row, with anterior tooth slightly displaced anteriorly and me- dially. 94 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Description. —Morphometric and meris- tic data for holotype and paratypes pre- sented in Table 1. Body size relatively small, maximum observed standard length about 41 mm. Maximum body depth at pelvic-fin origin. Anterior profile of snout and dorsal profile of head meeting in rounded obtuse angle near vertical line immediately ante- rior to nares. Dorsal profile of head poste- rior to that line inclined and slightly convex. Predorsal profile of body slightly arched be- tween supraoccipital and dorsal-fin origin. Dorsal profile of body straight to slightly concave between dorsal-fin origin and adi- pose fin. Dorsal profile from adipose fin to caudal-fin base straight. Ventral profile of head and body smoothly convex from mar- gin of lower lip to pelvic-fin origin or with indistinct rounded obtuse angle delimiting anteroventral angle of dentary. Upper jaw longer than, and overhanging, lower jaw. Anterior surface of snout fleshy, as in other Creagrutus, with minute papillae over surface. Greatest concentration of pa- pillae on upper lip, margin of upper jaw, and in mouth on fleshy, plicate flaps be- tween premaxillary teeth. Lower jaw with thick, fleshy anterior region and numerous papillae on lip. Infraorbitals poorly devel- oped compared to many Creagrutus species, covering less than one half of cheek, with ventral and posterior margins of series broadly separated from preopercle. Pos- teroventral margins of infraorbitals three and four rounded, with indentation or con- cavity at their juncture. Curvature of pos- teroventral margin of third infraorbital ap- proximately concentric with margin of orbit. Premaxillary dentition with three major components: 1) undulating main row of five, rounded unicuspid to tricuspid teeth; an- terior tooth slightly displaced anteromedi- ally; 2) triangular cluster of three larger tri- cuspidate teeth, crowded together on medial portion of premaxilla; and 3) single tooth, similar in morphology to those of main pre- maxillary row, occurring lateral to third or fourth tooth of that row. Maxilla with two or three, rarely four, unicuspid to tricuspid teeth. Dentary teeth six, anterior three larg- est and tricuspid followed by three unicus- pid teeth becoming successively shorter posteriorly. Unpaired fins relatively large compared with most Creagrutus; similar to that of C. lepidus Vari et al. (1993). Dorsal-fin origin slightly posterior to vertical through pelvic- fin origin. Dorsal fin 11,8, rarely ii,7; distal margin nearly straight, with slight elonga- tion of anterior rays. Anal fin ii or 111,10- 12; distal margin slightly concave with an- terior rays more elongate. Single, paired hooks present on 3 to 6 anterior branched anal-fin rays in males; hooks restricted to posterolateral surface of main shaft and pos- terior, secondary branch ofeach ray. Pectoral fin 1,10-12; fin reaching posteriorly almost to pelvic-fin base. Pelvic fin 1,7; approaching or, especially in sexually mature males, reaching anal-fin origin; with distal portion turned medially in some individuals giving fin slightly cupped appearance. Pelvic-fin hooks, when present, on all but smallest, distal branches of all branched rays. Gill-rakers short and stout, those of cer- atobranchial and epibranchial about equal in length; 3-6+ 8-10 = 11-16 (n = 21). Color in alcohol. —Dorsal surface of head with dark, shallow and deep chromato- phores. Large, stellate deep-lying chromato- phores lining interior surface of frontal, ex- cept in region of anterior fontanel. Shallow chromatophores punctate, present over most dorsal surfaces; most concentrated on snout and on ventral of upper lip, with small cres- cent of dark pigmentation immediately an- terior of nares. Three patches of chromato- phores extending posteriorly from portion of main field immediately dorsal to anterior margin of orbit; one over each orbit and one along midline over fontanel. Band of scat- tered dark chromatophores extending from pigmentation on snout posteriorly to an- teroventral margin of orbit and then around ventral and posterior margin of orbit. Scat- tered stellate dark chromatophores overly- VOLUME 107, NUMBER | ing dorsal portions of infraorbitals and opercle. Dorsal portion of body with small dark chromatophores concentrated on and below posterior portion of scales; overall pattern reticulate. Anterior one-half of pre- dorsal surface with longitudinal concentra- tion of large stellate chromatophores. Small dark chromatophores along dorsal-fin base. Humeral mark darkest immediately dorsal to lateral line, vertically elongate with ori- entation ranging from vertical to somewhat posterodorsally oblique. Dark midlateral stripe extending from pectoral girdle to cau- dal-fin base (Fig. 1). Stripe diffuse anteri- orly; most sharply defined ventrally and posteriorly; expanded into diffuse triangle extending slightly ventral to lateral line on caudal peduncle. Region of body between midlateral stripe and anal-fin base unpig- mented or with very small dark chromato- phores delineating myosepta. Dorsally ta- pered concentrations of dark pigment located between bundles of fin-ray muscu- lature at base of anal fin. Small dark chromatophores on caudal-fin membranes; greatest concentrations along central and outer branched rays and dorsal and ventral procurrent rays. Small dark chromatophores on anal-fin membranes mainly restricted to narrow bands along an- terior and posterior margins of fin rays. Slightly larger, darker chromatophores forming diffuse longitudinal band on distal one-third of anal fin; some specimens with greatly enlarged chromatophores, giving ap- pearance of dark spot on fin anteriorly. Small dark chromatophores present across dorsal- fin membranes; large, very dark chromato- phores concentrated in central portion of anterior one-half of fin, giving appearance of large dark spot (spot well developed across observed size range; present in 14.8 mm SL juvenile, MBUCV V-21257). Pectoral fins with series of dark chromatophores asso- ciated with most rays, most numerous lat- erally. Pelvic fins unpigmented. Color in life. —Dorsal and anal fins with bright red anteriorly, and black centrally (see 95 color plate in Roman 1992:169). Dorsal lobe of caudai fin red. Dorsal surface of eye with red patch overlying reflective guanine. Distribution.—Northern Venezuela east of the Andes from Estado Tachira in the west to Sucre in the east. Occurs in many upland tributaries of the Rio Orinoco basin and the Rios Tuy and Neveri of the Carib- bean versant. Ecology.—This species inhabits small, shallow, shady streams of the Andean pied- mont where the water is usually clear and substrate ranges from sand to gravel (Ta- phorn 1992:174). The diet of adults consists of small seeds, ostracods, gastropods, and aquatic insects, especially chironomid lar- vae (Pearse 1920:24, 25; Winemiller, pers. comm. in Taphorn 1992:174). Spawning occurs throughout the wet season, with in- dividuals probably spawning more than once a season (Taphorn 1992:174). Etymology. —A noun in apposition from the Greek melasma, meaning a black spot, in reference to the distinctive pigmentation of the dorsal fin. Remarks. —Material herein referred to Creagrutus melasma was identified as C. beni by Eigenmann (1920:12; 1927:422). This is puzzling, given the distinctiveness of the new species and the fact that Eigen- mann described C. beni. We have examined the holotype of C. beni and other material from near the type locality. Creagrutus me- lasma and C. beni are readily distinguished by number of vertebrae (34 or 35, 1 speci- men out of 60 had 36, in C. melasma versus 38 in the holotype of C. beni), and relative size of the infraorbital bones (poorly de- veloped, with posteroventral margin of the series distinctly separated from the pre- opercle in C. melasma compared to well developed, with the posteroventral margin approaching or contacting the preopercle in C. beni). Creagrutus melasma appears most simi- lar to C. lepidus in body and fin form, pre- maxillary dentition and gill-raker shape. The two species differ, however, in their humeral 96 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON spots (distinctly vertically elongate in me- lasma versus incorporated into midlateral stripe in /epidus), midlateral stripe (diffuse and tapering anteriorly in melasma versus broad and well developed anteriorly in /ep- idus), and dorsal-fin pigmentation (patch of dark pigmentation present in melasma ver- sus absent in /epidus; compare Fig. | with Vari et al. 1993:fig. 1). Acknowledgments We thank the following individuals and institutions for making material available: S. A. Schaefer and W. G. Saul (ANSP), T. Iwamoto (CAS), L. M. Page, K. S. Cum- mings, M. R. Retzer (INHS), F. Mago-Lec- cia, A. Machado-Allison, and F. Proven- zano (MBUCYV), C. Lasso (MHNLS), L. F. Palmer and S. J. Raredon (USNM). We also thank S. J. Raredon for technical assistance, G. C. Steyskal for his advice on the specific name, and W. C. Starnes and S. A. Schaefer for their careful reviews of the manuscript. Figure 1 was prepared by T. B. Griswold. Museum and field studies associated with this study in Venezuela were supported by the Neotropical Lowland Research Pro- gram of the Smithsonian Institution. Dur- ing this study A. S. Harold received support in the form of a Natural Sciences and En- gineering Research Council of Canada Post- doctoral Fellowship and a Tilton Postdoc- toral Fellowship, California Academy of Sciences. Literature Cited Eigenmann, C. H. 1911. New characins in the col- lection of the Carnegie Museum. — Annals of the Carnegie Museum 8(1):164—-181. . 1920. The fishes of Lake Valencia, Caracas, and of the Rio Tuy at El Concejo, Venezuela. — Indiana University Studies 7(44):1-13. . 1927. The American Characidae.— Memoirs of the Museum of Comparative Zoology, at Har- vard College 43(4):311-428. Harold, A. S., & R. P. Vari. 1994. Systematics of the trans-Andean species of Creagrutus (Ostario- physi, Characiformes, Caracidae). Smithsonian Contributions to Zoology, 551:1—31. Pearse, A.S. 1920. The fishes of Lake Valencia, Ven- ezuela. University of Wisconsin Studies in Sci- ence 1:1—51. Roman, B. 1992. Peces ornamentales de Venezuela. Fundacion La Salle de Ciencias Naturales, Ca- racas, 223 pp. Taphorn, D. C. 1992. The characiform fishes of the Apure River drainage, Venezuela. Biollania, Edicion Especial, 537 pp. Vari, R. P., A. S. Harold, C. A. Lasso, & A. Machado- Allison. 1993. Creagrutus lepidus, a new spe- cies from the Rio Aroa basin, Yaracuy State, Venezuela (Teleostei: Characiformes: Caraci- dae). Ichthyological Exploration of Freshwaters 4(4):351-355. (RPV) Division of Fishes, Department of Vertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560, U.S.A.; (ASH) Department of Ichthyology, California Academy of Sciences, Golden Gate Park, San Francisco, California 94118, U.S.A; and (DCT) Museo de Ciencias Naturales— Guanare, UNELLEZ, Guanare, Estado Por- tuguesa, 3310 Venezuela. PROC. BIOL. SOC. WASH. 107(1), 1994, pp. 97-108 A NEW GENUS OF FOSSIL PUFFERFISH (TETRAODONTIDAE: TETRAODONTIFORMES) BASED ON A NEW SPECIES FROM THE OLIGOCENE OF RUSSIA AND A REFERRED SPECIES FROM THE MIOCENE OF UKRAINE James C. Tyler and Alexandre F. Bannikov Abstract. —A new genus of tetraodontid pufferfish, Archaeotetraodon, is pro- posed for two fossil species with uniquely specialized bifurcate scale spinules: winterbottomi, a new species based on ten specimens from the Oligocene of Russia in which nearly all of the spinules are bifurcate; and jamestyleri Ban- nikov (1990), based on two complete specimens and a fragment from the Miocene of Ukraine in which only two scales in the middle of the body have bifurcate upright spinules. An especially long rayless pterygiophore extending forward from the dorsal-fin origin in Archaeotetraodon (in winterbottomi; con- dition unknown in jamestyleri) is similar to that found in the Recent Lago- cephalus but this feature cannot be unequivocally polarized and may not in- dicate relationship between these two genera. Expeditions by the Paleontological Insti- tute of the Russian Academy of Sciences to the North Caucasus have discovered a rich Lower Oligocene marine ichthyofauna in the Pshekhsky Horizon of the Lower Maikop deposits. This was first described by Danil- chenko (1960), with further studies more recently by the second listed author and his cooperating colleagues. Among newly ex- cavated fossils of early Oligocene age are ten specimens, four in counterpart plates, that are unique among tetraodontids by having most of the upright scale spinules that cover the entire body bifurcate from the base and divergent distally. The bifur- cate scale condition is shown to be a spe- cialization because all Recent and one of the other fossil tetraodontids as well as all members of the diodontid sister group have undivided upright scale spinules. We de- scribe these specimens as a new species and type of a new genus: Archaeotetraodon win- terbottomi. Our re-examination of the scales in ma- terials of all previously described fossil tet- raodontids based on relatively entire spec- imens shows that one of them, Sphoeroides jamestyleri Bannikov (1990), from the Mio- cene of Ukraine, has most of the scales with unbranched spinules but that a limited patch in the middle of the body has bifurcate spi- nules. We transfer S. jamestyleri from Sphoeroides, into which it was originally placed mostly for the convenience of not having to create a new genus, to Archaeotet- raodon on the basis of its sharing with A. winterbottomi specialized bifurcate scales. The two species of Archaeotetraodon differ from one another not only in the coverage with bifurcate scales but also in several os- teological features. Methods Length is standard length (SL) unless oth- erwise stated. Fossil materials are from the collections of the Paleontological Institute (PIN) of the Russian Academy of Sciences in Moscow, the Museo Civico di Storia Na- turale di Verona (MCSNV), the Istituto di 98 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Geologia della Universita di Padova (IGUP), and Sammlung der Philosophie- theologie Hochschule der Eichstatt (SPHE). Comparative anatomical preparations and radiographs of Recent species of tetraodon- tiforms are those listed in Tyler (1980), sup- plemented by those of tetraodontids listed in Tyler et al. (1992). In many derived groups of tetraodonti- forms, including tetraodontids, there is a bony element in the upper midline of the body just in front of the soft dorsal fin that is thought to represent a basal pterygiophore that no longer bears dorsal-fin spines. This was called a supraneural in Tyler (1980), but is here referred to as a rayless pteryg- iophore, following the recommendation of Mabee (1988:836), who demonstrated that true supraneurals (predorsals) are not ho- mologous with such pterygiophores. For tetraodontoid phylogeny we follow the cladistic analysis of Winterbottom (1974) and the evolutionary systematics of Tyler (1980), which, respectively on the ba- sis of specialized myological and osteolog- ical features, are in agreement that: diodon- tids are the sister group of tetraodontids; molids are the sister group of the tetraodon- tid + diodontid clade; triodontids are the sister group of the tetraodontid + diodontid + molid clade; and the Eocene eoplectids are the morphologically primitive sister group of all of these other tetraodontoids. Among other tetraodontiforms, the balis- toid + ostracioid clade is the first outgroup and the triacanthoid clade the second out- group. There is no cladistic analysis available for the genera of tetraodontids, and Tyler (1980) simply placed the Recent genera into three groups of relative degrees of morphological specialization in what can be considered an unresolved trichotomy. The Eocene Eotet- raodon was presumed in that work to be the morphologically primitive sister group of these three groups of Recent genera because of its retention of such plesiomorphic fea- tures as twelve principal caudal-fin rays and pleural ribs (both of which are found in tri- odontids); however, because diodontids and molids have no pleural ribs and fewer than twelve caudal-fin rays, it is more parsimo- nious to propose that the twelve caudal-fin rays and pleural ribs of Eotetraodon are re- versals. Although it was inconsistent with his pre- sumed phylogeny of tetraodontid genera, Tyler (1980) continued the practice of rec- ognizing Canthigaster as subfamilially dis- tinct from other tetraodontids even though the genus Carinotetraodon was shown to be anatomically intermediate between Canthi- gaster and other tetraodontids in many ways, including several specialized features (e.g., skin ridge-lifting behavior, highly arched vertebral column, large haemal spines on abdominal vertebrae). This led Tyler (1980) to the conclusion that Carinotetraodon and Canthigaster had a close common ancestry. Therefore, we agree with Winterbottom (1974:99) that Canthigaster cannot reason- ably be recognized as subfamilially distinct from (and sister group to) a polyphyletic subfamily for all other tetraodontids, in- cluding Carinotetraodon. With the phylog- eny of tetraodontid genera so poorly known, we compare any unusual features of Ar- chaeotetraodon with comparable conditions in all other tetraodontid genera, including the specialized Canthigaster + Carinotetra- odon clade to which we doubt Archaeotetra- odon is closely related. Family Tetraodontidae (sensu Tyler, 1980) Archaeotetraodon, new genus Type species. —Archaeotetraodon winter- bottomi, new species, by present designa- tion; other species, Sphoeroides jamestyleri Bannikov (1990), by referral herein. Diagnosis. — Differs from all other tetra- odontids by the presence of bifurcate up- right spinules on either most of the scale plates over the entire body (in winterbot- tomi) or on many of those of the middle of the body (in jamestyleri). VOLUME 107, NUMBER | Description. — With the exception ofa few species in which scales have been second- arily lost (several species of both Sphoe- roides and Takifugu and single species of both Lagocephalus and Tetraodon; Tyler 1980:297-298) all Recent tetraodontids have specialized scales in which the basal plate, which has two or more processes ra- diating out into the skin, bears an upright spinule that protrudes through the skin as a prickle. The spinule is of varying stoutness and length, but most often is short and slen- der (in contrast to the larger and stouter projecting spines in diodontids), giving a shagreen-like quality to the skin (see illus- trations of scales in Recent species of nu- merous tetraodontid genera in Tyler 1980: 291-297). That the new species of tetraodontid from the Oligocene, A. winterbottomi, has an ex- tensive covering of bifurcate scale spinules led us to re-examine the scales in the two previously described species of fossil tetra- odontids based on relatively entire speci- mens, the Eocene Eotetraodon pygmaeus (Zigno 1887) and the Miocene Sphoeroides Jamestyleri Bannikov (1990). The holotype and four other previously unreported spec- imens of E. pygmaeus are covered with un- branched spinules like those of all Recent tetraodontids. The holotype of Sphoeroides jamestyleri does not have the scales pre- served, but the paratypic entire specimen has a complete covering of scales, most of which have unbranched spinules. However, some scales in the middle of the body (it is impossible to distinguish whether this is dorsal, ventral, or lateral) are just as dis- tinctly bifurcate distally as those in A. win- terbottomi, although the spinules are pro- portionally shorter in S. jamestyleri. Because all species, both fossil and Re- cent, of the diodontid sister group have scale plates with unbranched upright spinules or spines like those of all Recent and one of the fossil species of tetraodontids (except more massive), we propose that spinules with undivided upright shafts are primitive 99 for the tetraodontid + diodontid clade and that the deeply bifurcate spinules in the Oli- gocene A. winterbottomi and Miocene S. jamestyleri are a specialization. Therefore, the bifurcate spinule condition is a syna- pomorphy of A. winterbottomi and S. jamestyleri. The latter was originally placed in Sphoeroides because it 1s thought to be a morphologically relatively primitive genus (Tyler 1980) defined by a combination of what seem to be mostly plesiomorphic fea- tures, and no features that are known to be specialized. It was simply convenient to place S. jamestyleri in that poorly defined genus pending acquisition of better pre- served materials showing more internal fea- tures which might clarify its relationships, but such materials are not yet available. However, we can state that all of the few known internal features of similarity be- tween S. jamestyleri and the species of Sphoeroides are plesiomorphic (e.g., mod- erate interorbital and ethmoid widths; mod- erate and mostly laterally directed exten- sions of the lateral ethmoids, sphenotics, and pterotics) and that there are no known specialized features of similarity that unite S. jamestyleri with Sphoeroides. Because S. jJamestyleri does share the uniquely derived feature of bifurcate scale spinules with A. winterbottomi, we remove jamestyleri from Sphoeroides and place it in Archaeotetra- odon along with winterbottomi. We note that in molids the basal plates of the scales are rounded to rectilinear and bear a central emargination or low spinule, and in at least smaller specimens of Mola some of these spinules are branched distally (see illustration in Tyler 1980:369). In all other tetraodontoids with upright spinules on the basal plate, the spinules are un- branched (a single spinule in eoplectids, the sister group of all other tetraodontoids, and several spinules in the poorly known zig- noichthyids, that are most closely related to the tetraodontid + diodontid clade). In tri- odontids the scales bear a low spiny ridge and there are no upright spinules. There- 100 fore, on the basis of the phylogeny of tetra- odontoid families proposed by both Win- terbottom (1974) and Tyler (1980), it is most parsimonious to presume that the distally branched spinules in some molids and the deeply bifurcate branched spinules in Ar- chaeotetraodon are independent acquisi- tions. Etymology. —From the Greek: archaios, old or ancient, and tetraodon, for the four tooth plates characteristic of the family Tet- raodontidae; masculine. Similarities of Archaeotetraodon to Other Tetraodontids Rayless pterygiophore. — Archaeotetra- odon winterbottomi has an exceptionally long rayless pterygiophore, averaging 18% SL as measured from the anterior end of the el- ement to the anterior end of the base of the soft dorsal fin in the three specimens in which this region is preserved. We presume that this long slender element is a single piece of bone from its posterior end at the dorsal-fin origin to its anterior end at the level of the vertical through the centrum of the sixth to seventh abdominal vertebra be- cause we cannot see any interruptions or articulations in it, although our view of the bone is somewhat obscured by the layer of spiny scales in the overlying skin. In the holotype of A. jamestyleri the region ante- rior to the dorsal fin is poorly preserved, while in the paratypic entire specimen the skeleton is much disarticulated and it is not possible to recognize a rayless pterygio- phore among the mixture of bones. Thus, the condition of the rayless pterygiophore in A. jamestyleri is unknown and in the fol- lowing discussion of the rayless pterygio- phore the statements about Archaeotetra- odon are based on the conditions in A. winterbottomi. In some Recent tetraodontids with rela- tively long rayless pterygiophores, this ele- ment does not reach posteriorly to the dor- sal-fin origin but, rather, terminates anterior PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON to it and articulates there with an anterior process on the distal end of the first basal pterygiophore of the dorsal fin. We see no evidence of such an anterior process on the first basal pterygiophore in Archaeotetra- odon, but that pterygiophore is not well ex- posed in our material. Although we cannot be absolutely sure of it, we have no reason to believe that the rayless pterygiophore in Archaeotetraodon does not extend as a sin- gle slender bone for the full length of the distance from its anterior end to the dorsal- fin origin. Nevertheless, it is possible that some small portion of our measurement of the rayless pterygiophore posteriorly in Ar- chaeotetraodon may include part of the dis- tal head of the first basal pterygiophore of the dorsal fin. Even with that caveat, we believe that the average measurement of 18% SL is a fair estimate of the length of the rayless pterygiophore in Archaeotetra- odon. No other tetraodontid has a rayless pterygiophore as long as that in Archaeotet- raodon, but a few genera contain species with rayless pterygiophores almost as long. The rayless pterygiophore is especially long and slender in the six species of La- gocephalus examined, more so in some spe- cies than in others. For example, in L. iner- mis (Temminck & Schlegel), L. /aevigatus (Linnaeus), L. /unaris (Bloch & Schneider), and L. spadiceus (Richardson) the rayless pterygiophore length averages 8—9% SL (in 2 to 12 specimens of each species exam- ined), while it averages 12% SL in L. sclera- tus (Gmelin) (in 4 specimens) and 15% SL in L. lagocephalus (Linnaeus) (in 2 speci- mens), the latter being the longest rayless pterygiophore of which we are aware among Recent tetraodontids. In all of these species of Lagocephalus the distal end of the first dorsal-fin basal pterygiophore has a prom- inent anterior process to which the rayless pterygiophore articulates. The rayless pte- rygiophore of Lagocephalus, even though shorter than in Archaeotetraodon, extends at least as far forward as in Archaeotetra- odon, and, in L. lagocephalus, extends even VOLUME 107, NUMBER | further forward, to the level of the vertical through the region of articulation between the centra of the fourth and fifth abdominal vertebrae. The two species of Lagocephalus with the longest rayless pterygiophores, L. scleratus and L. lagocephalus, are stream- lined in form and have an offshore pelagic habitat; there may be a correlation in tetra- odontids between a long and slender rayless pterygiophore and a strong swimming, pe- lagic mode of life. The rayless pterygiophore is relatively long in the nine species of Canthigaster ex- amined, averaging between 9% and 13% SL. The rayless pterygiophore of Canthigaster differs from the slender rod as found in Ar- chaeotetraodon and Lagocephalus in being heavier, deeper, concave ventrally, and curved ventrally at its anterior end, follow- ing the contour of the arched back. In most species of Sphoeroides the rayless pterygiophore is short, averaging about 4— 5% SL, but the element is somewhat longer in such species as S. spengleri (Bloch) and S. marmoratus (Lowe), averaging 6—7% SL. In S. formosus (Gunther) (sometimes rec- ognized in the monotypic Guentheridia) the rayless pterygiophore is far longer, heavier, and deeper than in the other species of Sphoeroides, averaging 12% SL (in 5 spec- imens) but without the slender form found in Lagocephalus and Archaeotetraodon. Other than the species mentioned above, we know of no other tetraodontids with no- tably long rayless pterygiophores, 1.e., of 10% SL or greater length. The condition of the rayless pterygiophore in a variety of tetra- odontids can be assessed from the illustra- tions of representative species in Tyler (1980:figs. 195, 203, 226-244), including its absence in a few species. The similarity in the length of the long, slender rayless pterygiophore between Ar- chaeotetraodon and some species of Lago- cephalus is difficult to interpret because of the unknown phylogeny within tetraodon- tids and because diodontids, their sister group, do not have a rayless pterygiophore. 101 In the molid sister group of the tetraodontid + diodontid clade the rayless pterygiophore is either present as a short deep piece (Mola), absent or fused with the first basal pteryg- iophore of the soft dorsal fin (Masturus), or perhaps consolidated into a long complex structure that connects the first basal pte- rygiophore of the soft dorsal fin with the supraoccipital crest (Ranzania). In triodon- tids a rudimentary spiny dorsal fin, when present, of two or three spines is borne on two basal pterygiophores that are connected to the basal pterygiophores of the soft dorsal fin by two short rayless elements. These two short elements presumedly are derived from basal pterygiophores that no longer support spines at the rear of the rudimentary spiny dorsal fin. In those populations (Indian Ocean) of Triodon macropterus (Lesson) (the only Recent representative of the family) that usually entirely lack the spiny dorsal fin, all four of the elements in this series that extends anteriorly from the soft dorsal-fin origin therefore are rayless pterygiophores, the first of which is elongate and the more posterior three pieces short. In eoplectids a well-developed spiny dorsal fin is present and its basal pterygiophores connect with those of the soft dorsal fin without the in- tervention of rayless pterygiophores. Among the outgroup tetraodontiforms, the six dor- sal-fin spines in triacanthoids are borne on four or five basal pterygiophores and there are no rayless pterygiophores between the basal pterygiophores of the spiny and soft dorsal fins. In balistids the three dorsal-fin spines are borne on two basal pterygio- phores that form a complex carina sup- ported by a rayless pterygial strut that braces the carina against the first basal pterygio- phore of the soft dorsal fin, with the strut apparently being derived from the third bas- al pterygiophore of the spiny dorsal fin of triacanthoids. In monacanthids there are two dorsal-fin spines, and the less robust carina, which is formed from a single basal pteryg- iophore, is not supported posteriorly by a pterygial strut. In ostracioids the spiny dor- 102 sal fin is absent and a long and deep (ara- canids) or short (ostraciids) rayless pteryg- iophore is present anterior to the base of the soft dorsal fin. Thus, when the spiny dorsal fin is absent in tetraodontiforms such as os- tracioids, rayless pterygiophores are present and apparently represent basal pterygio- phores of the absent spiny dorsal fin, while the reduction in number of dorsal-fin spines and their supporting basal pterygiophores from posteriorly in the series in balistids in comparison to triacanthoids is accompa- nied by the apparent conversion of the third basal pterygiophore of triacanthoids into the rayless pterygial strut of balistids. Most germane, however, is the situation in triodontids, the sister group of all other Recent families of tetraodontoids, in which the distinction between basal pterygio- phores and rayless pterygiophores depends simply on whether the rudimentary dorsal- fin spines are present or not. Therefore, we propose that the presence of a rayless pte- rygiophore, representing a rudimentary support of the now absent spiny dorsal fin, 1S primitive for the tetraodontid + diodon- tid + molid clade of tetraodontoids. How- ever, it is equally parsimonious to hypoth- esize that: 1) a rayless pterygiophore was present in the ancestor of the tetraodontid + diodontid clade and that the rayless pte- rygiophore was independently lost by all diodontids and by some tetraodontids; or 2) a rayless pterygiophore was lost in the ancestor of the tetraodontid + diodontid clade and the rayless pterygiophore ac- quired by most tetraodontids as a reversal to the ancestral tetraodontoid condition. Moreover, even given that a rayless pteryg- 1ophore is primitive for the tetraodontid + diodontid + molid clade, it is not known whether that element was long or short or slender or stout. Presuming that the rayless pterygiophore of the ancestral tetraodon- toid without a spiny dorsal fin was one of the four dorsal pterygial elements as found in triodontids, it could as logically be the long, stout, anteriormost first rayless pte- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON rygiophore of those Triodon macropterus lacking dorsal-fin spines as it could be one of the three short and heavy more posterior rayless elements. We have no way of know- ing at present whether the rayless pteryg- iophore in the ancestral tetraodontoid with- out a spiny dorsal fin was long or short. Therefore, similarity in the long rayless pte- rygiophore of Archaeotetraodon and Lago- cephalus may be plesiomorphic and not in- dicative of relationship. Our surmise is that the great length and, especially, the slender form of the rayless pterygiophore is a derived feature, but since we do not know of any unequivocally de- rived features shared by Archaeotetraodon and Lagocephalus, it may be that the pu- tatively derived condition of the long, slen- der rayless pterygiophore is independently acquired by these two genera. From what little is known of its osteology, Archaeotet- raodon differs from Lagocephalus in having relatively evenly tapered neural and haemal spines on the vertebrae of the caudal pe- duncle anterior to the penultimate vertebra, whereas in Lagocephalus these are expand- ed anteroposteriorly, a derived condition (absent in other tetraodontids and in all oth- er tetraodontoids). Elongate head spines.—In at least one specimen of Archaeotetraodon winterbot- tomi the spinules on the top of the head are longer than elsewhere and many of these are not bifurcate. In only one other species of tetraodontid are the spinules on the head much longer than those on the body, this being one of the several species of Ambly- rhynchotes, A. piosae. In A. piosae the body is made exceptionally prickly by spinules that are longer than in other tetraodontids, and the spinules on the top and side of the front of the head are especially elongate, up to 8% SL. These spinules are proportionally far longer than those that are slightly elon- gate on the top of the head in A. winterbot- tomi. In both cases among tetraodontids in which speciose genera have a few species in which spiny scales are lost, these species are VOLUME 107, NUMBER | not considered to be closely related within their respective genera (Sphoeroides and Takifugu) and the loss of spines has been considered to be independent (Tyler 1980: 297). With so much homoplasy in even the presence or absence of spines within tetra- odontid genera, we place no phylogenetic significance on the fact that both one spec- imen of A. winterbottomi and one of the several species of Amblyrhynchotes have the spinules longer on the head than elsewhere, especially when the spinules in A. winter- bottomi are otherwise so different (shorter and bifurcate) than those in A. piosae, and when there are no other derived features of similarity between Archaeotetraodon and any of the species of Amblyrhynchotes. Ar- chaeotetraodon differs from Amblyrhyn- chotes not only by its short bifurcate scales but most notably also by having 18 versus 19 or 20 vertebrae (in the three species of Amblyrhynchotes examined) and a long (18% SL) versus short rayless pterygiophore (5— 6% SL in three species of Amblyrhynchotes). Generic Relationships On the basis of its few known osteological and external features we are not able to place Archaeotetraodon into one of the three mor- phological groups recognized by Tyler (1980) for Recent genera of tetraodontids. Like- wise, we do not find any special similarity between the two species of Archaeotetra- odon and other fossil species of tetraodon- tids. The Eocene Eotetraodon pygmaeus (Zigno) has neither bifurcate scales nor a long rayless pterygiophore. The Pliocene Sphoeroides hyperostosus (Tyler et al. 1992) is based on skulls and anterior vertebrae but the scales and portions of the body that might include a rayless pterygiophore are unknown; it differs from Archaeotetraodon by the extensive hyperostosis of many skull bones at sizes as small as that of the larger specimens of either of the two species of Archaeotetraodon. Several other species of fossil tetraodontids have been named on the 103 basis of pieces of jaw bones of Miocene and younger age but these cannot be usefully compared with the fossil species based on more complete specimens. We can only call attention to the unique bifurcate scales that distinguish Archaeotet- raodon from all other genera of tetraodon- tids and note that while the elongate and slender rayless pterygiophore as found in A. winterbottomi and some Lagocephalus may be a specialization for a pelagic mode of life, it seems likely to have been an independent acquisition in the few pelagic species of La- gocephalus and in A. winterbottomi, which has been found in a predominantly pelagic fossil ichthyofaunal assemblage. Archaeotetraodon winterbottomi, new species Figs. 1-4 Diagnosis. — Archaeotetraodon winterbot- tomi differs from the only other species of the genus, A. jamestyleri, by having: a ver- tebral formula of 8+10 (versus 7+11 in jamestyleri); almost complete covering of bifurcate scale spinules (versus bifurcate only on middle of body); haemal spine of pen- ultimate vertebra with a moderately long posteroventral process under the parhypu- ral region (versus no prolongation); supra- cleithrum relatively long and narrow (ver- sus shorter and thicker). Description. —Body moderately elongate (Figs. 1-2). Vertebrae 18 in four specimens in which total number can be counted, with eight abdominal and ten caudal in only specimen (holotype) in which proximal end of first anal-fin basal pterygiophore can be seen in association with a haemal spine; ver- tebral column relatively straight, only gent- ly arched in abdominal region. Caudal skel- eton relatively distinct in holotype, and having normal tetraodontid pattern of a long parhypural, a lower hypural plate fused to last centrum, an upper free hypural plate and an epural (exact shape unclear) above last centrum. Penultimate vertebra (PU>) 104 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON fee Nts 78 e 2 i Fig. 1. Photograph of holotype of Archaeotetraodon winterbottomi, PIN 3363/111, 90.0 mm SL, Lower Oligocene (Maikopian) of North Caucasus, southwest Russia. with broad neural and haemal spines, the latter prolonged posteriorly under a little more than half of length of parhypural; more anterior caudal vertebrae with more slender neural and haemal spines, except haemal spines of first three caudal vertebrae short, where proximal ends of anal-fin basal pte- rygiophores are supported. First three ab- dominal vertebrae apparently with bifid neural spines and fourth abdominal verte- bra with neural spine bifid anteriorly but undivided posteriorly, where it is prolonged posteriorly over base of neural spine of fifth vertebra. Dorsal-fin rays nine in two specimens and nine or perhaps ten in one specimen. Anal- fin rays eight in only specimen in which all rays are preserved, at least basally. Basal example of scale covering pterygiophores in dorsal fin seven and in anal fin six in single specimen in which these can be counted. Caudal-fin rays 11 in four specimens, best preserved in holotype, with uppermost ray and two lowermost rays un- branched and other eight rays branched, four above middle of hypural plate and four be- low (typical tetraodontid condition). Cau- dal-fin length 23.3-27.3% SL (25.8% aver- age) in three specimens. Pectoral fin not well enough preserved to describe. A single upright spinule arising from each basal scale plate, spinules mostly short and divergently bifurcate from base (Fig. 3), length of upright spinules along top of mid- dle of body in nine specimens 0.7—1.6% SL (1.3% average); these prickly scales present continuously over most of head and body. Fig. 2. Reconstruction of Archaeotetraodon winterbottomi, based on the holotype, data as in Fig. 1. VOLUME 107, NUMBER 1 105 Fig. 3. A. Photograph of scales along dorsal surface of paratype of Archaeotetraodon winterbottomi, PIN 3363/115, ca. 40 mm SL, longest upright bifurcate spinules 0.5 mm (1.3% SL), age and locality as in Fig. 1. B. Drawing of selected scales along same dorsal surface as in A. 106 Fig. 4. Drawing of inner surface of premaxilla of holotype of Archaeotetraodon winterbottomi, data as in Fig. 1. In one specimen (PIN 3363/120) a small group of about ten spinules on top of head slightly longer than those on body (1.4% versus 0.9% SL) and mostly unbranched, with all other spinules on head and body bifurcate; we presume that having a few slightly elongate and non-bifurcate spinules on top of head is normal for at least some specimens of this species (perhaps a sexually dimorphic feature). Rayless pterygiophore long and slender, apparently a single piece, its posterior end at origin of soft-dorsal fin and its anterior end at level of vertical through centrum of sixth or seventh abdominal vertebra; its length 16.0-—20.6% SL (18.1% average) in three specimens. Inner surface of premaxilla visible in two specimens, both of which have three tritu- ration teeth, about three times as mediolat- erally wide as anteroposteriorly deep; me- dial edge of premaxilla with articular processes increasing in size posteriorly in series (Fig. 4), interdigitating with similar processes on apposed premaxilla. Inner sur- face of dentary visible in one specimen and no trituration teeth present; medial edge of dentary with articular processes like those of premaxilla. Preopercle broad and strongly curved, with ridges in middle region. Postcleithrum long and slender. Supracleithrum long and PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON gently curved, with a low medial flange. In- terorbital width moderate, least width about 4.0-4.5% SL in two entire specimens in which it is possible to recognize the lateral edges of the upper orbit in neurocrania pre- served in dorsoventral view. No other fea- tures of skeleton clearly enough exposed or preserved to warrant description. Etymology. —winterbottomi, honoring our friend and colleague Richard Winterbot- tom, Royal Ontario Museum, in recogni- tion of the excellence of his important stud- ies on the phylogeny of tetraodontiforms and of his great help to us in our own efforts with the plectognath fishes. Type materials.—Holotype: PIN 3363/ 111 (head to left) and 11la, counterpart plates, River Pshekha, 90.0 mm SL. Para- types: PIN 3363/112 (head to right), single plate, River Belaya, 53.9 mm SL; PIN 3363/ 113 (head to left), single plate, River Belaya, 36.5 mm SL; PIN 3363/114 (head to left) and 1 14a, counterpart plates, River Belaya, 27.8 mm SL; PIN 3363/115 (dorsoventral impression), single plate, River Belaya, ca. 40 mm SL; PIN 3363/116 (head to left), single plate, River Kuban, most of head missing, length of vertebral column 32.6 mm; PIN 3363/117 (head to left), single plate, River Pshekha, fragment of most of vertebral column, whose length is ca. 33 mm; PIN 3363/118 and 118a (dorsoventral impression), counterpart plates, River Be- laya, 24.7 mm SL; PIN 3363/119 (head to left) and 119a, counterpart plates, River Be- laya, 52.1 mm SL; PIN 3363/120 (head to right), single plate, River Belaya, 64.2 mm SL. Except for the two specimens preserved entirely as dorsoventral impressions, all of the above are preserved as lateral impres- sions of the body in which, however, the neocranium is often in dorsoventral view. Type locality. —Holotype from River Pshekha (at Gorny Luch). Paratypes from Rivers Pshekha, Belaya (upstream from the settlement of Abadzekhskaya), and Kuban (near the town of Cherkessk), all of which sites are within, respectively, 32 and 150 VOLUME 107, NUMBER 1 km of one another in the Pshekhsky (Pshek- ha) Horizon, lower part of the Maikop (Mai- kopian) deposits, Lower Khadum Forma- tion, Lower Oligocene, North Caucasus of southwest Russia, about 35 million years ago. Stratigraphy and ichthyofaunal associa- tions. — About 55 other species of fishes have been collected at the localities where the type series of A. winterbottomi were found in the Maikop deposits of the Lower Oli- gocene. These fishes are predominantly pe- lagic forms (see table 3 in Danilchenko 1980), including the Caprovesposus acronu- rus presettlement stage of an acanthurid (Bannikov & Tyler 1992), numerous clu- peids of the genera Sardinella and Pomo- lobus and gadids of the genus Palaeogadus. Several strata of the Pshekhsky Horizon bear rather numerous mesopelagic photophore- bearing fishes of the genera Eomyctophum, Vinciguerria and Scopeloides. Moreover, representatives of such apparently pelagic families as Scombridae (Scombrosarda, Sarda), Trichiuridae (Lepidopus), Palaeo- rhynchidae (Palaeorhynchus, Homorhyn- chus), Nomeidae (Psenicubiceps, Rybapi- na), Stromateidae (Pinichthys), etc., were abundant in the early Eocene of the North Caucasus. Coastal and benthic fishes were much rarer, although among those few ben- thic species is the only previously known Maikopian tetraodontiform, Oligobalistes robustus Danilchenko (1960). We presume that the preponderance of pelagic fishes at the localities of the type series of A. winterbottomi is evidence that it is an offshore or pelagic species of tetra- odontid, like some of the species of Lago- cephalus. The gray, flaky marls and calcareous clays of the Oligocene Pshekhsky Horizon cover light calcareous rocks of the underlying Up- per Eocene Byeloglinsky Horizon of the North Caucasus. The only fish remains that are known from the latter horizon are iso- lated scales of a large elopiform of the genus Lyrolepis. 107 Comparative Fossil Materials Eotetraodon pygmaeus (Zigno 1887): all specimens from the Lower Eocene of Monte Bolca, Italy; IGUP 6890-91, counterpart plates, holotype, 18.2 mm SL; MCSNV T137-138, counterpart plates, 16.1 mm SL; MCSNV T1339, single plate, 14.2 mm SL; SPHE 1970/48, single plate, 15.5 mm SL; SPHE 1970/47, single plate, 90.5 mm SL. Archaeotetraodon jamestyleri (Bannikov 1990): all specimens from the Tarkhanian Horizon of the Lower Miocene at Kamysh- lak, Kerch Peninsula, Crimea, Ukraine; PIN 287-9, counterpart plates, holotype, 22.6 mm SL; PIN 3974-8, single plate, paratype, vertebral column distorted, cranium pre- served as dorsoventral impression of 16.4 mm length from anterior end of vomer to rear of occipital region, estimated 60 mm SL; PIN uncatalogued, fragment represent- ing part of caudal peduncle. Sphoeroides hyperostosus Tyler, Purdy, & Oliver (1992): both specimens from the Yorktown Formation of the Lower Pliocene of Lee Creek Mine, Beaufort County, North Carolina, USA; USNM 437601, relatively complete three dimensional skull and first four vertebrae, holotype, 72.5 mm cranium length; USNM 290643, three dimensional cranium, paratype, 37.0 mm cranium length. Acknowledgments We thank the Russian Academy of Sci- ences and Smithsonian Office of Fellowship and Grants for support of travel leading to this and other joint publications by the au- thors on fossil fishes in collections of the Paleontological Institute in Moscow. We are grateful to: Lorenzo Sorbini, Museo Civico di Storia Naturale, Verona, for help in the examination of specimens of fossil tetraodon- tids from the rich Monte Bolca collections in his museum; Victor Krantz, Smithsonian Institution, for specimen photography; Gregory Klassen, then a postdoctoral fellow at the Smithsonian Institution, for discus- 108 sions of tetraodontid character polarity based on his current work on the phylogeny of the family; Eugenia B. Bohlke and Wil- liam G. Saul, Academy of Natural Sciences of Philadelphia, for loan of specimens; and Frederick V. Grady, Department of Paleo- biology, National Museum of Natural His- tory, Smithsonian Institution, for his care- ful preparation of comparative materials of Pliocene Sphoeroides hyperostosis. The manuscript benefited from the con- structive peer review comments of Colin Patterson, British Museum (Natural His- tory), Richard Winterbottom, Royal On- tario Museum, and Thomas A. Munroe, National Marine Fisheries Service, Nation- al Museum of Natural History, Smithson- ian Institution. Literature Cited Bannikov, A. F. 1990. Pervaya nakhodka iglobryu- khikh ryb (Tetraodontidae) v miotsene SSSR [The finding of a pufferfish (Tetraodontidae) in the Miocene of the USSR].—Paleontologiches- kij Zhurnal 4:75-80. — ,, & J.C. Tyler. 1992. Caprovesposus from the Oligocene of Russia: the pelagic acronurus pre- settlement stage of a surgeonfish (Teleostei: Acanthuridae).— Proceedings of the Biological Society of Washington 105(4):810-820. Danilchenko, P. G. 1960. Kostistye ryby maikop- skikh otlozhenij Kavkaza [see Danilchenko, 1967, below, for translation].—Trudy Paleon- tologicheskogo Instituta Akademii Nauk SSSR 78:1-208. 1967. Bony fishes of the Maikop deposits of the Caucasus. Tranlated from Russian. U.S. De- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON partment of Interior and the National Science Foundation, Israel Program for Scientific Trans- lations, Jerusalem, 247 pp. . 1980. Osnovnye kompleksy ikhtiofauny kai- nozoyskikh morei Tetisa [The major ichthyo- faunal assemblages of the Cenozoic Seas of the Tethys]. Pp. 175-183 in L. I. Novitskaya, ed., Iskopayemye kostistye ryby SSSR [Fossil teleost fishes of the USSR]. Trudy Paleontologiches- kogo Instituta Akademii Nauk SSSR 178:1—211. Mabee, P.M. 1988. Supraneural and predorsal bones in fishes: development and homologies. —Copeia 1988(4):827-838. Tyler, J.C. 1980. Osteology, phylogeny, and higher classification of the fishes of the Order Plectog- nathi (Tetraodontiformes).— National Oceanic and Atmospheric Administration, Technical Report, National Marine Fisheries Service, Cir- cular 434:1-422. ——., R. W. Purdy, & K. H. Oliver. 1992. New species of Sphoeroides pufferfish (Teleostei: Tet- raodontidae) with extensive hyperostosis from the Pliocene of North Carolina.— Proceedings of the Biological Society of Washington 105: 462-482. Winterbottom, R. 1974. The familial phylogeny of the Tetraodontiformes (Acanthopterygii: Pi- sces) as evidenced by their myology.—Smith- sonian Contributions to Zoology 155:1—201. Zigno, A. 1887. Nuove aggiunte alla ittiofauna dell’epoca Eocena.— Memorie del Reale Istituto Veneto di Scienze, Lettere ed Arti (Venezia) 23: 9-33. (JCT) National Museum of Natural His- tory, Smithsonian Institution (MRC-106), Washington, D.C. 20560, U.S.A.; (AFB) Paleontological Institute, Russian Academy of Sciences, Profsoyuznaya 123, Moscow 117647, Russia. PROC. BIOL. SOC. WASH. 107(1), 1994, pp. 109-113 PLEISTOCENE ECHINOIDS (ECHINODERMATA) FROM BERMUDA AND BARBADOS Stephen K. Donovan and Brian Jones Abstract. — Pleistocene fossil echinoids have been neglected compared to more ancient members of this group. Two occurrences of extant species in the Pleis- tocene are documented herein. Echinometra lucunter (Linné) from Bermuda adds to the growing list of fossil occurrences of this species, which has a durable test, but lives in areas of poor preservation potential. Meoma ventricosa (La- marck) from Barbados is the first ‘complete’ specimen of this species from the fossil record. Pleistocene echinoids have received less attention than those from earlier in the fos- sil record for a variety of reasons. For ex- ample, they are commonly poorly preserved (Gordon & Donovan 1992) and they usually belong to extant species that have already been well-studied by zoologists. The fact that these species are still extant, however, makes Pleistocene echinoids particularly suitable for investigating and testing diverse paleo- biological problems and hypotheses. Gor- don & Donovan (1992) used disarticulated plates to determine the distribution of echi- noids on a Sangamonian (late Pleistocene) raised reef, and Donovan & Gordon (1993) tested taphonomic predictions made on the basis of living species (Greenstein 1991) against the same taxa that occurred in the Plio-Pleistocene of the Caribbean region. The present paper records the occurrence of moderately well-preserved echinoid tests from the Pleistocene deposits of Bermuda and Barbados. The Pleistocene echinoid faunas of both islands are essentially un- known and it is hoped that the present com- munication may stimulate further research. The echinoid classification used herein follows Smith (1984). Terminology of the echinoid test follows Melville & Durham (1966) and Smith (1984). Specimens de- scribed herein are deposited in the Field Museum of Natural History, Chicago (FMNH), and the University of Alberta (UA). Class Echinoidea Leske Superorder Camarodonta Jackson Order Echinoida Claus Family Echinometridae Gray Genus Echinometra Gray Echinometra lucunter (Linné, 1758) Fig. | Material, locality and horizon. —A single test, FMNH PE 309, from Bermuda. The specimen label states ““Echinometrid echi- noid (labeled Echinometra lucunter). ““Cla- docora rock’’— Quaternary [presumably Pleistocene]. Bermuda. Gift of Bermuda Bi- ological Station, 1947”. Quaternary depos- its of Bermuda have been deposited over the past 250,000 years (Harmon etal. 1983). Preservation.—The test (Fig. 1) is pre- served in a well-indurated white limestone. Fragments of Cladocora are apparent in limestone adhering to the outside of the test. The Aristotle’s lantern and part of the test (approximately coinciding with interam- bulacrum | and ambulacrum IJ) are missing, and more of the test is concealed by limestone (Fig. la). The apical region is poorly preserved and the apical system is 110 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. missing (Fig. 1a). Parts of the test are broken and crushed, commonly across plates. Al- though in other regular echinoids this might mean that death was caused by predation (Smith 1984:19), the plates of extant E. /u- cunter are so well interlocked by stereom trabeculae that this breakage, associated with an essentially ‘complete’ test, was probably post-burial. This specimen appears to fall into disintegration state 5 of Kidwell & Baumiller (1990:249). Description. —For a recent description of E. lucunter, see Donovan (1993). Echinometra lucunter (Linné, 1758), FMNH PE 309. a, apical view. b, lateral view. Scale bars in mm. Remarks. —At present, E. /ucunter ranges from Florida to Brazil and off the west coast of Africa (Kier 1992). In particular, it occurs on hard substrates in high energy environ- ments around Bermuda (D. L. Pawson, pers. comm.). Previously, E. /ucunter has been reported from the Plio-Pleistocene of Ja- maica, the Dominican Republic, Florida, and Angola (Gordon 1991, Kier 1992; note that in the former reference ‘Dominica’ in table 1 should read Dominican Republic). It is also reported from the ‘middle’ Oli- gocene (Larue 1994) San Sebastian For- VOLUME 107, NUMBER 1 111 Fig. 2. Meoma ventricosa (Lamarck, 1816), UA 9496. a, apical view, b, oral view. Scale bar in mm. mation of Puerto Rico (Gordon 1963), but this could be a misidentification of E. prisca Cotteau which is abundant at about the same level in Antigua (Poddubiuk & Rose 1985: table 1). However, this possibility was dis- cussed by Gordon (1963:632, 635), who noted that the pore pairs of the Puerto Rico specimens are mainly arranged in arcs of six as in E. lucunter, but dissimilar to E. prisca. The Bermudan specimen adds to the growing data base of occurrences of fossil E. lucunter (Donovan & Gordon 1993). Al- though this taxon has a high potential for preservation when compared with other regular echinoids (Greenstein 1991), Gor- don (1991) noted that Echinometra has a poor fossil record as complete tests. This is presumably due to the preferred life habit of this taxon, on hard substrates under high energy conditions, which is usually re- moved from environments of sedimentary deposition (Smith 1984:22). In conse- quence, fossil EZ. /ucunter is rarely found as ‘complete’ tests in Pleistocene deposits (Donovan & Gordon 1993), although Echi- nometra may be locally abundant as dis- articulated plates (Gordon & Donovan 1992). Superorder Microstomata Smith Order Spatangoida Claus Family Micrasteridae Lambert Genus Meoma Gray Meoma ventricosa (Lamarck, 1816) Fig. 2 Material, locality and horizon. —A single test, UA 9496, from the Arawak Cement Quarry, northwest Barbados. The specimen came from a large boulder lying on the quar- ry floor. This test was preserved in mod- erately lithified grainstone, about 125,000 years old (Upper Pleistocene, Sangamoni- an; last interglacial), that accumulated in pockets between large Montastrea annularis colonies that are the main component of the reef at this locality. Preservation. —This specimen is pre- served largely as an internal mold, although test calcite is retained over about half of the oral surface and in part of interambulacrum 2 apically (Fig. 2). A patchy, calcareous ve- 112 neer occurs over much of the internal mold that may be a remnant of the test, although individual plates are not discernable, per- haps suggesting it is possibly diagenetic in origin. The mold is composed of lithified carbonate sand. This specimen is particu- larly noteworthy because of its large size, about 160 mm long by 135 mm wide by 65 mm high. Although not unusual for this spe- cies, few Caribbean fossil echinoids of com- parable dimensions are known. Description. —For a comprehensive de- scription of this species, see Chesher 1970; see also Kier & Grant 1965, Chesher 1969, and Kier 1975. Remarks. —The only fossil echinoids pre- viously documented from Barbados are from the Tertiary clastic sedimentary rocks exposed in the northeast of the island (Kier 1966). Kier (1966:2) commented that “It is rather surprising, considering the large numbers of echinoids known from Eocene rocks in the Caribbean ... that .. . species from the Upper Scotland Formation are quite distinct.”’ These rocks were previously thought to be shallow-water in origin (see, for example, Barker & McFarlane 1980). However, it is now generally recognized that siliciclastic and other rocks of northeast Barbados were of deep-water origin (Speed 1988). Eocene echinoids described from the Caribbean tend to be preserved in lime- stones of shallow-water origin, hence the differences between faunas. In Barbados, only the Pleistocene limestones were truly deposited in shallow-water. Extant Meoma ventricosa is a sand-bur- rowing spatangoid found in the Caribbean, Florida, the Bahamas and the Pacific coast of Panama (Chesher 1969, 1970). It still oc- curs in sandy reef environments similar to that interpreted for the Pleistocene locality in Barbados (for example, the lagoon at east Discovery Bay, north Jamaica; SKD, per- sonal observation). However, it is poorly represented in the fossil record and only ap- pears to have been reported previously from PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON the Upper Pleistocene of Jamaica (Donovan et al. 1994), based on test fragments. Two subspecies of this taxon were rec- ognized by Chesher (1970), M. vy. ventricosa (Lamarck) and M. vy. grandis Gray. Distin- guishing between these two subspecies relies on features of the test that are not preserved in the Barbadian specimen. This test is un- usually broad and could, very tentatively, be referred to M. y. grandis, which was orig- inally described from the Pacific coast of Panama. If this provisional identification is correct, then either a contraction of range since the late Pleistocene or an ecopheno- typic, rather than genetic, explanation for the two test morphologies in this species is indicated. However, Kier & Grant (1965: pl. 9, fig. 4) illustrated a test of M. v. ven- tricosa of comparable gross morphology to the specimen from Barbados. Acknowledgments This research was undertaken during the period of a Shell Distinguished Research Fellowship in Science to SKD, which is gratefully acknowledged. Fieldwork in Bar- bados by BJ was supported by Natural Sci- ences and Engineering Research Council of Canada grant #A6090. We thank Greg Buckley and Scott Lidgard of the Field Mu- seum of Natural History, Chicago, for ar- ranging the loan of FMNH PE 309. This specimen was photographed at the Smith- sonian Institution, National Museum of Natural History, Washington, D.C., using equipment made available by David Paw- son, who also kindly commented on the ex- tant echinoids of Bermuda. Literature Cited Barker, L., & N. McFarlane. 1980. Notes on some sedimentological evidence for shallow water or- igin of parts of the Scotland Formation of Bar- bados. —Journal of the Geological Society of Ja- maica 19:46. Chesher, R. H. 1969. Contributions to the biology of VOLUME 107, NUMBER 1 Meoma ventricosa (Echinoidea: Spatangoida). — Bulletin of Marine Science 19:72-110. 1970. Evolution in the genus Meoma (Echi- noidea: Spatangoida) and a description of a new species from Panama.—Bulletin of Marine Sci- ence 20:731-761. Donovan, S. K. 1993. Jamaican Cenozoic Echinoi- dea. Jn R. M. Wright & E. Robinson, eds., Bio- stratigraphy of Jamaica.— Geological Society of America Memoir 182:371-412. ,&C.M. Gordon. 1993. Echinoid taphonomy and the fossil record: supporting evidence from the Plio-Pleistocene of the Caribbean. — Palaios 8:304—-306. , H. L. Dixon, R. K. Pickerill, & E. N. Doyle. 1994. A Pleistocene echinoid (Echinodermata) fauna from southeast Jamaica.— Journal of Pa- leontology 68: 8 pp. Gordon, C. M. 1991: The poor fossil record of Echi- nometra (Echinodermata: Echinoidea) in the Caribbean region.—Journal of the Geological Society of Jamaica 28:37-41. —, &S.K. Donovan. 1992. Disarticulated echi- noid ossicles in paleoecology and taphonomy: the last interglacial Falmouth Formation of Ja- maica.—Palaios 7:157—-166. Gordon, W. A. 1963. Middle Tertiary echinoids of Puerto Rico.— Journal of Paleontology 37:628— 642. Greenstein, B. J. 1991. An integrated study of echi- noid taphonomy: predictions for the fossil rec- ord of four echinoid families.—Palaios 6:519— 540. Harmon, R. S. et al. 1983. U-series and amino-acid racemization geochronology of Bermuda: im- plications for eustatic sea-level fluctuation over the past 250,000 years. — Palaeogeography, Pa- laeoclimatology, Palaeoecology 44:41-70. Kidwell, S. M., & T. Baumiller. 1990. Experimental disintegration of regular echinoids: roles of tem- perature, oxygen, and decay thresholds. — Paleo- biology 16:247-271. Kier, P. M. 1966. Four new Eocene echinoids from Barbados.— Smithsonian Miscellaneous Collec- tions 151(9):1-28. 1975. The echinoids of Carrie Bow Cay, Be- lize.—Smithsonian Contributions to Zoology 206:1-45. 1992. Neogene paleontology in the northern Dominican Republic. 13. The Class Echinoidea 113 (Echinodermata).— Bulletins of American Pa- leontology 102(339):13-27, 31-40. , & R. G. Grant. 1965. Echinoid distribution and habits, Key Largo Coral Reef Preserve, Florida.—Smithsonian Miscellaneous Collec- tions 149(6):1-68. Lamarck, J. B. P. A. de M. de. 1816. Histoire na- turelle des animaux sans vertébres, présentant les caractéres, généralaux et particuliers de ces animaux, leur distribution, leurs classes, leurs familles, leurs genres et la citation synonymique des principales espéces qui s’y rapportent (1st edition, volume 3). Paris, 586 pp. [Not seen.] Larue, D. K. 1994. Puerto Rico and the Virgin Is- lands. Pp. 15 inS. K. Donovan & T. A. Jackson, eds., Caribbean geology: an introduction. Uni- versity of the West Indies Publishers’ Associa- tion, Kingston (in press). Linné, C. 1758. Systema naturae per regna tria na- turae, secundum classes, ordines, genera, spe- cies, cum characteribus differentis, synonymis, locis. Holmiae, 824 pp. [Not seen.] Melville, R. V.,& J. W. Durham. 1966. Skeletal mor- phology. Pp. U220—U252 in R. C. Moore, ed., Treatise on invertebrate paleontology, Part U, Echinodermata 3(1). Geological Society of America and University of Kansas Press, New York and Lawrence. Poddubiuk, R. H., & E. P. F. Rose. 1985. Relation- ships between mid-Tertiary echinoid faunas from the central Mediterranean and eastern Carib- bean and their palaeobiogeographic signifi- cance.—Annales Géologiques des Pays Helle- niques 32:115-127. Smith, A. B. 1984. Echinoid palaeobiology. George, Allen and Unwin, London, 191 pp. Speed, R. 1988. Geologic history of Barbados: a pre- liminary synthesis. Pp. 29.1—29.11 in L. Barker, ed., Transactions of the 11th Caribbean Geo- logical Conference, Dover Beach, Barbados, July 20-26, 1986. Energy and Natural Resources Di- vision, Barbados. (SKD) Department of Geology, Univer- sity of the West Indies, Mona, Kingston 7, Jamaica; (BJ) Department of Geology, Uni- versity of Alberta, Edmonton, Alberta T6G 2E3, Canada. PROC. BIOL. SOC. WASH. 107(1), 1994, pp. 114-118 A NEW SPECIES OF FELICOLA (PHTHIRAPTERA: TRICHODECTIDAE) FROM A COSTA RICAN JAGUAR, PANTHERA ONCA (CARNIVORA: FELIDAE) Robert M. Timm and Roger D. Price Abstract.—A new species of chewing louse, Felicola (Lorisicola) oncae (Phthiraptera: Trichodectidae), is described and illustrated from a jaguar, Panthera onca (Carnivora: Felidae), taken in Costa Rica. Although this louse is based only on a single male specimen, its morphological distinctiveness and occurrence on a big cat of the genus Panthera make its discovery and description of special significance. The genus Fe/icola Ewing now contains 55 species, which can be grouped into four subgenera: Felicola, Lorisicola, Paradoxuroecus, and Suricatoecus. Resumen. —Se describe e ilustra una nueva especie de piojo, Felicola (Lo- risicola) oncae (Phthiraptera: Trichodectidae), de un jaguar, Panthera onca (Car- nivora: Felidae), capturado en Costa Rica. Aunque este piojo se basa solamente en un unico ejemplar macho, su distintividad morfologica y ocurrencia en un gran felino del género Panthera le confieren a su descubrimiento y descripcion una significancia especial. El genero Felicola Ewing contiene ahora 55 especies, las cuales pueden agruparse en cuatro subgéneros: Felicola, Lorisicola, Para- doxuroecus, y Suricatoecus. Fifty-four species of chewing lice of the genus Felicola Ewing currently are recog- nized. Of these, 48 species occur on the felid carnivores, the families Felidae, Herpesti- dae, and Viverridae, 5 occur on the Canidae (Carnivora), and | is found on the Lorisidae (Primates). Eleven of these species occur on the cats of the family Felidae. The trichodectid chewing louse of the ge- nus Felicola, subgenus Lorisicola Bedford, that we describe herein was obtained from an adult male jaguar, Panthera onca (Lin- naeus), that was shot on the night of 10 June 1988 north of Delicias de Upala, about 1 km from the Nicaraguan border in extreme northern Costa Rica. The next day, the in- tact jaguar was hauled in the back of the truck to Universidad Nacional in Heredia where it was skinned. The skin was frozen and stored in a freezer for four years. We were then able to thaw and wash the skin in an attempt to recover ectoparasites and obtained the single specimen of Felicola and a single tick. This is the first louse to be reported from a free-ranging jaguar. Al- though we do not generally believe that new species should be described on the basis of a single individual, in this case we feel it is warranted. The unlikelihood of our obtain- ing additional specimens of lice from jag- uars, coupled with the extremely fortuitous collection of the type specimen, places us in the position of documenting, based upon a single individual, that a unique species of chewing louse occurs on one of the most endangered species of Neotropical mam- mals. Because of the significant unique char- acters that this louse possesses, especially VOLUME 107, NUMBER 1 its unusual genitalia and extremely large body size, we are confident that it deserves recognition as a distinct species. Felicola (Lorisicola) oncae, new species Figs. 1, 2 Type host.—Panthera onca (Linnaeus). Male.—As in Fig. 1. Head with prean- tennal margin straight, with shallow narrow medioanterior indentation; sparse scattered dorsal setae; antennal scape enlarged. Each side of pronotum with few short lateral se- tae, single median marginal seta. Pterono- tum with row of short setae laterally. Ab- domen with two short setae on tergum I and single row of short setae on terga and sterna IJ-VIII. Pleura II—VIII with single row of short setae, with those on III somewhat lon- ger and stouter than those on V. With six pairs of large abdominal spiracles. Chae- totaxy of terminalia as shown. Genitalia (Fig. 2) with straight parallel basal apodeme lat- eral struts associated with large spinose sac; parameres apically tapered, blunt, and ba- sally fused, with flat basal margin lacking indentation; mesomeral arch with promi- nent broad blunt apical process. Female. —Unknown. Dimensions of male (in mm).—Temple width, 0.59; head length, 0.52; prothorax width, 0.46; pterothorax width, 0.56; ab- domen width at IV, 0.92; total length, 2.04; genitalia width, 0.22; genitalia length, 0.55; genitalia paramere length, 0.18. Type material.—Holotype male, ex Panthera onca, Costa Rica: Alajuela Prov- ince, Upala Canton, 2 km north of Delicias de Upala, 10 June 1988; in collection of Snow Entomology Museum, University of Kansas, Lawrence. Etymology.—This species is named for the host, Panthera onca, the jaguar. Remarks. —Felicola oncae is distin- guished from the males of all other known species of the genus by the combination of its head shape, possession of six pairs of 115 prominent abdominal spiracles, the very large dimensions, and the shape of the fused parameres and mesomeral arch of the gen- italia. Although the gross head shape and spiracle number are similar to those of the majority of the species of Lorisicola from the Felidae, no members of the other three subgenera have more than four pairs of ab- dominal spiracles and most have a quite different head shape. Within the Lorisicola found on felids, the largest species known previously is F. spen- cer! Hopkins from the two species of Hol- arctic lynx, Lynx canadensis Kerr and L. lynx (Linnaeus). Felicola spenceri is 1.51 mm long, with a temple width of 0.46 mm, head length of 0.45 mm, and abdomen width of 0.70 mm; F. oncae is considerably larger in all dimensions. Felicola zeylonicus Bed- ford, of the subgenus Felicola, is the largest previously known species in the genus, with a total length of 1.66 mm. Thus, F. oncae is by far the largest of any Felicola known to date. In addition to its extremely large size, the basally fused parameres of F. oncae are unique in shape and the mesomeral arch has a broad, blunt apical process. The api- cally separated parameres with the basal margin of the fusion area flattened and with- out an indentation were known previously only in F. americanus Emerson & Price, from the bobcat, Lynx rufus (Schreber); the two species are grossly different in dimen- sions and genitalia. Felicola americanus (subgenus Lorisicola), second in size to and considerably smaller in all dimensions than F. spenceri, is thereby much smaller than F. oncae. In the Emerson & Price (1983) key to males of the New World species of the Feli- cola felis complex, F. oncae identifies readily with F. americanus in couplet 3 on the basis of the flattened basal margin of the fused parameres. If one passes through that cou- plet, F. oncae would identify further with F. spenceriin couplet 4. As explained above, gross differences in dimensions and geni- talia separate F. oncae from both of these 116 Figs. 1, 2. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 1 Felicola oncae. \. Male. 2. Male genitalia. VOLUME 107, NUMBER | species, as well as from all other members of the subgenus and genus. Lyal (1985), in his cladistic classification of the trichodectids, treated what we rec- ognize herein as the genus Felicola as two genera, Felicola and Lorisicola, with each having two subgenera, Felicola and Suri- catoecus Bedford, and Lorisicola and Par- adoxuroecus Conci, respectively. We accept Lyal’s placement of 18, 11, 12 (+ our new species), and 13 louse species, respectively, in these four subgenera. However, we be- lieve it more appropriate that all four be recognized as subgenera of Felicola. We come to this conclusion because of the dif- ficulties encountered in the key by Lyal (1985:335-338) to genera and subgenera. The longest and most complex couplets are used for Felicola sensu lato, character states are nondiscrete and overlapping, and sep- arations are ambiguous. There is simply no clear break between these groups that war- rants generic level distinction. This action on our part is not a severe departure from Lyal’s classification, and is one that we feel is justified. Eleven of the 12 Felicola now known from felids are in the subgenus Lorisicola. The single other species of Felicola reported from felids, F. subrostratus (Burmeister), is in the subgenus Felicola. Felicola subrostratus has been reported from the domestic cat com- plex, Felix catus-lybica-silvestris. These cats have been domesticated and transported by humans for at least four millennia. Domes- tic cats generally are treated as a man-cre- ated species, F. catus, that was derived from the wild cat of northern Africa and extreme southeastern Asia, F. silvestris lybica, al- though considerable interbreeding with the wild cat of Europe, F. silvestris silvestris, has occurred. Because we have been unable to examine lice from truly wild, non-feral cats, we are unable to evaluate the relationship of Felicola subrostratus to other Felicola. The family Felidae, or cats, is nearly worldwide in distribution, being found in all zoogeographic regions except for the 117 Australian and Oceanic regions, Madagas- car, and the smaller oceanic islands. The family contains some four or five Recent genera and 37 extant species. Although there is little disagreement in the number of spe- cies recognized, there has been considerable debate on the number of genera and the relationships between species (Wilson & Reeder 1993). The number of genera of Re- cent felids recognized by various authors ranges from 4 to 19 (Ewer 1973, Nowak 1991). Four main lineages of extant cats are recognized: the cheetah (genus Acinonyx), the clouded leopard (genus Neofelis), the smaller cats (genus Felis, with as many as 14 subgenera in the single genus, or as many as 16 genera, including Lynx), and the big cats (genus Panthera). In Panthera, five spe- cies are recognized—leopard, P. pardus (Linnaeus); lion, P. /eo (Linnaeus); snow leopard, P. uncia (Schreber) (often treated as a monotypic genus Uncia); tiger, P. tigris (Linnaeus); and jaguar. Historically, jaguars were found from the southern United States, through Mexico, all of Central America, and much of tropical lowland South America, to central Argen- tina. We suspect that Felicola oncae is a host-specific parasite of jaguars and, as such, occurs on jaguars throughout their range. For well over two centuries, however, jaguar numbers have been declining; populations have been reduced by hunting pressure and habitat destruction and, in recent years, jag- uars have been extirpated from much of their former range. All previous bona fide records of Felicola from felids have been from the smaller cats of the genera Felis and Lynx. However, Ponton (1870) did describe Trichodectes ti- gris supposedly originating from a tiger. Un- fortunately, he provided no illustration and the verbal description is so general as to apply to a wide range of generic possibilities. Hopkins (1949:507) suggested that the louse was “... almost certainly from a captive and perhaps a contamination.”” Hopkins and Clay (1952:354), in referring to 7. ftigris, 118 tersely state ““Type lost, description useless. Unrecognizable.”” Lyal (1985:247) adopts this approach and relates the name to the category of incertae sedis. Our discovery of Felicola oncae on the jaguar, therefore, is the first verifiable record and recognizable description for any louse from a member of the subfamily Panther- inae and suggests that Felicola may be much more widely distributed on the cats than was recognized previously. Including F. on- cae, 12 species of Felicola have been de- scribed from 15 species of felids. Where ac- curate records are available, Felicola appears to be quite host specific. Given that chewing lice have been found on only 15 of the 37 extant species of cats, we strongly suspect that numerous new species of Felicola await discovery. Acknowledgments We are extremely grateful to several col- leagues in Costa Rica for their efforts with the jaguar that made our discovery of this louse possible. Costa Rican biologist Carlos Alberto Lopez transported the jaguar on his back, by motorcycle, and by pickup truck from the site where it was killed to the Uni- versidad Nacional. David Norman of the Programa Regional en Manejo de Vida Sil- vestre para Mesoamérica y el Caribe, Uni- versidad Nacional, skinned the jaguar and generously made the skin available to us. Miguel Rodriguez and Eduardo Lopez Pi- zarro of the Direccion de Vida Silvestre provided the permits. RMT’s field work in Costa Rica was funded in part by the Uni- versity of Kansas Office of Research, Grad- uate Studies, & Public Services; Associate Vice Chancellor Robert C. Bearse’s efforts in securing funding is most appreciated. We thank Adrian Nieto for translating our abstract into Spanish for the resumen and PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Barbara L. Clauson for her constructive comments on an earlier version of this manuscript. This study was partially sup- ported by Project No. Min-17-015, Min- nesota Agricultural Experiment Station, St. Paul, Minnesota 55108, and has been as- signed Paper No. 20,553, Scientific Journal Series. Literature Cited Emerson, K. C., & R. D. Price. 1983. A review of the Felicola felis complex (Mallophaga: Tricho- dectidae) found on New World cats (Carmivora: Felidae).— Proceedings of the Entomological Society of Washington 85:1-9. Ewer, R. F. 1973. The carnivores. Cornell University Press, Ithaca, xv + 494 pp. Hopkins, G. H.E. 1949. The host-associations of the lice of mammals.— Proceedings of the Zoolog- ical Society of London 119:387-604. —, & T. Clay. 1952. A check list of the genera & species of Mallophaga. British Museum (Nat- ural History), London, 362 pp. Lyal, C. H. C. 1985. A cladistic analysis and classi- fication of trichodectid mammal lice (Phthir- aptera: Ischnocera).—Bulletin of the British Museum (Natural History), Entomology Series 51:187-346. Nowak, R.M. 1991. Walker’s mammals of the world. Vol. II. Sth edition. Johns Hopkins University Press, Baltimore, pp. 643-1629. Ponton [misspelled Penton], T. G. 1870. On anew species of parasite from the tiger. — Monthly Mi- croscopical Journal 4:147-148. Wilson, D. E., & D. M. Reeder (eds.). 1993. Mammal species of the world: a taxonomic and geograph- ic reference. 2nd ed. Smithsonian Institution Press, Washington, xvii + 1206. (RMT) Museum of Natural History and Department of Systematics and Ecology, University of Kansas, Lawrence, Kansas 66045-2454, U.S.A.; (RDP) Department of Entomology, University of Minnesota, St. Paul, Minnesota 55108, U.S.A. (Current address) 4622 Kinkead Ave., Fort Smith, Arkansas 72903, U.S.A. (direct reprint re- quests to RMT). PROC. BIOL. SOC. WASH. 107(1), 1994, pp. 119-121 A NEW SPECIES OF HYPERALONIA RONDANYI, 1863 (INSECTA: DIPTERA: BOMBYLITDAE: EXOPROSOPINAE) Marcia Souto Couri and Carlos José Einicker Lamas Abstract.—A new species of Hyperalonia from Brazil, H. diminuta, is de- scribed and illustrated. Illustrations of wings of other Neotropical taxa are also included to aid in their identification. The Neotropical genus Hyperalonia Ron- dani, 1863, comprises four species and two subspecies (Painter et al. 1978). They were revised by Painter & Painter (1968), who described a new species, H. ater, and pre- sented a diagnosis of the genus, a key to species and subspecies and recorded their distribution. Except for H. ater, all other known species and subspecies are represented in Museu Nacional, Rio de Janeiro collection. Study- ing this material, the authors found a new species of Hyperalonia, which is herein de- scribed. The unique exemplar of H. ater ob- served is deposited at Museu de Zoologia, Universidade de Sao Paulo. To contribute further to the identification of the genus, wing illustrations of Neotrop- ical taxa are also presented. Hyperalonia diminuta, new species Figs. 1, 2 Holotype.—é. Aragarcas/Goias Brasil, 28 Jan 1953, Moacir Alvarenga. Holotipo [red label]. Hyperalonia diminuta Couri & La- mas [Lamas’ handwriting]. (MNRJ). In good condition; right third antennal article bro- ken. Genital segments in glycerin inside mi- crovial pinned with holotype. Diagnosis. — Tip of wing hyaline, outline between black and hyaline areas straight; hyaline areas as follows: area in center of wing from near base of first posterior cell to near posterior margin of discal cell; this hy- aline area is largely separated from another small one, at third posterior cell, which is round; large round hyaline area at second basal cell; a small round hyaline area crossed by R2 + 3 at its basal third (Fig. 1); hairs on costal base black; alula fringe white. Mid tibia at ventral surface with a row of about 9 short black bristles; dorsal surface with 2 bristles at basal half, one bristle at apical third and an apical bristle; hind tibia on ventral and dorsal surfaces with a complete row of short and black bristles. Description. —Male: Body length— 10 mm; wing 10 mm (Figs. 1—2). Head: orange yellow, except ocellar tubercle and a trian- gular area in front of it, which are both black, mouth opening brown posteriorly; frons protruded; sparse black hairs on occiput, frons and gena; occipital fringe yellowish white; a dense tuft of black hairs in the black spot on front of ocellar tubercle; antennae brown, with black hair on first article; first and second articles about the same length, both as long as broad; third article about 3 times the length of second; proboscis hardly exceeding mouth opening; palpi brown with black hairs. Thorax: black bluish; humeral and postalar callus brown; sparse minute black pubescence on notum; orange red col- lar of hairs on anterior part of thorax; black hairs between coxa; a tuft of orange red hairs on humeral callus and between this and wing; and another one above halter; a tuft of white hairs between base of wing and halter; halter brown and yellow; calypter dark brown; anepisternum, katepisternum and anepimeron with black hairs. Legs brown, mid tibia on ventral surface with a row of about 9 short black bristles; dorsal 120 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 1,0 cm Figs. 1-7. 1-2, Hyperalonia diminuta: (1) wing of holotype; (2) male genitalia, lateral view; 3, Hyperalonia surinamensis, wing; 4, Hyperalonia morio morio, wing; 5, Hyperalonia morio erythrocephala, wing; 6, Hyper- alonia chilensis, wing; 7, Hyperalonia ater, wing. VOLUME 107, NUMBER 1 surface with 2 bristles on basal half, one bristle at apical third and an apical bristle; hind tibia on ventral and dorsal surfaces with a complete row of short, black bristles. Wing: blackish brown; tip hyaline, outline between black and hyaline areas straight; hyaline areas as follows: area in center of wing near base of first posterior cell to near posterior margin of discal cell; this hyaline area is largely separated from another small one, at third posterior cell, which is round; big round hyaline area at second basal cell; a small round hyaline area crossed by R2 + 3 at its basal third (Fig. 1); hairs on costal base black; alula fringe white. Abdomen: Black bluish as thorax with small black hairs at dorsum and bristled laterally. Male gen- italia in lateral view with basistylus sub- triangular; dististylus with hook-shaped apex; tip of aedeagus recurved dorsally; epandrium subquadrate, with prominent ventral process. Cercus short (Fig. 2). Discussion. — The color pattern of the wing easily distinguishes H. diminuta from other species in the genus. In Painter & Painter’s (1968) key, this species is near H. surina- mensis, as in both, the line between hyaline and dark parts at tip of wing is straight. The following key separates these two species. Triangular hyaline area in the end of second basal cell; a broad hyaline area in center of wing separated by a band of black bor- dering the third posterior cell, this infe- rior part reaches margin of wing (Fig. 3) PE enn hd es, H. surinamensis Rondani Round hyaline area at second basal cell; narrow hyaline area in center of wing, separated by a large band from a small hyaline area in third posterior cell, which does not reach margin of wing (Fig. 1) H. diminuta 121 The color pattern of wings of Hyperalonia morio morio (Fabricius, 1775) (Fig. 4); H. morio erythrocephala (Fabricius, 1805) (Fig. 5); H. chilensis Rondani, 1863 (Fig. 6); H. surinamensis Rondani, 1863 (Fig. 3); and H. ater Painter & Painter, 1968 (Fig. 7) are presented, to facilitate the identification of species. Acknowledgments We are grateful to Dr. Neal Evenhuis (Bishop Museum, Honolulu) for reviewing the manuscript and to Dra. Francisca do Val (Museu de Zoologia, Universidade de Sao Paulo) for the loan of H. ater. Literature Cited Fabricius, J.C. 1775. Systema entomologiae, sistens insectorum classes, ordines, genera, species adiectis synonymis, locis, descriptionibus, ob- servationibus, 832 pp. Flensburghi et Lipsiae. 1805. Systema antliatorum, secundum or- dines, genera, species, 373 + 30 pp. Brunsvigae. Painter, R. H., & E. M. Painter. 1968. A review of the genus Hyperalonia Rondani (Bombyliidae, Diptera) from South America.—Papéis Avulsos de Zoologia, Sao Paulo 22(12):107-121. —_,, ,&J. Hall. 1978. Family Bombyliidae. Pp. 1-92. A Catalogue of Diptera of the Amer- icas South of the United States. Departamento de Zoologia, Universidade de Sao Paulo. Rondani, C. 1863. Diptera exotica revisa et annotata, 99 pp. Modena [Also published as ““Dipteroum species et genera aliqua exotica’. — Archivio per la Zoologia, l’ Anatomia e la Fisiologia, Modena (1863)3(1):1-99, 1864]. Departamento de Entomologia, Museu Nacional, Universidade Federal do Rio de Janeiro, Quinta da Boa Vista, Sao Cristo- vao, Rio de Janeiro, RJ. 20.940-040, Brazil. PROC. BIOL. SOC. WASH. 107(1), 1994, pp. 122-131 LOUISEA, A NEW GENUS OF FRESHWATER CRAB (BRACHYURA: POTAMOIDEA: POTAMONAUTIDAE) FOR GLOBONAUTES MACROPUS EDEAENSIS BOTT, 1969 FROM CAMEROON Neil Cumberlidge Abstract. — Globonautes macropus edeaensis Bott, 1969 and G. balssi Bott, 1959 from Cameroon are removed from the Gecarcinucidae Rathbun, 1904 and reassigned to the Potamonautidae Bott, 1970. Globonautes m. edeaensis is recognized as a valid species, and is established as the type species of Louisea, a monotypic new genus. Louisea is defined by a combination of characters of the mandible, third maxilliped, cheliped, and gonopods 1 and 2. Louisea edeaensis is compared to, and distinguished from, other freshwater crabs oc- curring in West Africa. Globonautes balssi is close to Louisea but is regarded here as incertae sedis. A key to the West African genera of the Potamonautidae is provided. The African family Potamonautidae Bott, 1970 currently includes four genera, Pota- monautes MacLeay, 1838, Sudanonautes Bott, 1955, Liberonautes Bott, 1955, and Potamonemus Cumberlidge & Clark, 1992. Potamonautes is widely distributed throughout sub-Saharan Africa (Bott 1955), Sudanonautes is found from Cote d’Ivoire to Central Africa (Bott 1955; Monod 1977, 1980; Cumberlidge 1989, 1993a), and Libe- ronautes is found in West Africa west of Ghana (Cumberlidge & Sachs 1989a, 1989b). The fourth genus, Potamonemus, 1s known only from southwest Cameroon (Cumberlidge & Clark 1992, Cumberlidge 1993b). The present work reappraises the taxon- omy of two poorly known species from Cameroon, G. macropus edeaensis Bott, 1969, and Globonautes balssi Bott, 1959. Both species are presently included in the Gecarcinucidae Rathbun, 1904 (Bott 1959, 1969, 1970) but there is some doubt as to the validity of these assignments. For ex- ample, the form of the mandible of G. m. edeaensis indicates that this species belongs in the Potamonautidae, while Cumberlidge (1987) examined the mandible of G. balssi and concluded that this taxon also belongs in the Potamonautidae. The exact classification of G. m. edeaensis and G. balssi within the Potamonautidae is more difficult. The two species most closely resemble members of Potamonemus, which are also from the same part of Cameroon (Cumberlidge & Clark 1992, Cumberlidge 1993b). Potamonemus is characterized by a potamonautid-type mandibular palp (2- segmented, and ending in a single lobe) to- gether with a third maxilliped which com- pletely lacks a flagellum on the exopod. Two male specimens of G. m. edeaensis from Cameroon, recently discovered in the Mu- seum fur Naturkunde der Humboldt-Uni- versitat, Berlin, Germany (ZMB), allow a complete re-assessment of this taxon. G. m. edeaensis has a potamonautid-type man- dibular palp, together with a third maxilli- ped which completely lacks a flagellum on the exopod. However, differences in the form of gonopod 2 of G. m. edeaensis argue against its inclusion in Potamonemus, or in any of the other three potamonautid freshwater crab genera known from Africa. The new VOLUME 107, NUMBER | genus Louisea is therefore proposed to ac- commodate G. m. edeaensis, which is des- ignated the type species, and a key to the West African genera of the Potamonautidae is provided. Unfortunately, the position of G. balssi cannot be properly assessed because the male type specimen is in poor condition; there- fore this taxon is regarded here as incertae sedis. The following abbreviations are used: CW = carapace width at widest point; CL = car- apace length, measured along median line; CH = carapace height, maximum depth of cephalothorax; FW = front width, width of front measured along anterior margin; M = male, F = female, ad = adult, juv = juvenile, Ovig = ovigerous; SMF = Natur-Museum und Forschungs-Institut Senckenberg, Frankfurt am Main, Germany; USNM = National Museum of Natural History, Smithsonian Institution, Washington, D.C.; ZIM = Zoological Institute and Museum, Hamburg, Germany; ZSBS = Zoologische Sammlung des Bayerischen Staates, Miin- chen, Germany. Methods The holotype of L. edeaensis was loaned from the ZSBS; two other specimens dis- covered in the ZMB were subsequently loaned. The male holotype of L. edeaenis is in poor condition, and the entire left man- dible and the palp of the right mandible are missing. Fortunately the other specimens of L. edeaensis are in good condition, and were collected in Cameroon less than 90 km from the type locality. The male holotype of G. macropus (Rathbun, 1898) was examined in the USNM. The male holotype and the second female paratype of G. balssi were examined in the SMF where they were on temporary loan. The male holotype of G. balssi is a juvenile, and both second gonopods are missing. The female paratype of G. balssi was examined in the ZIM. 123 Four dimensions of the carapace, cara- pace length, carapace width, carapace height, and front width—were recorded from each specimen using digital calipers (Table 1). The relative proportions of the latter three measurements (adjusted for body size, CL) of both species were calculated (Table 1). The left mandible and the left gonopod 1 and gonopod 2 were illustrated following removal from the specimens in order to de- scribe these structures from different angles and under magnification. Characters of the gonopods, carapace, and third maxillipeds of these specimens correspond closely with those of the holotype. The carapace, man- dible and other characters of the largest of these specimens were illustrated. The length of the propodus of the right and left che- lipeds of all specimens was measured along the ventral margin (Table 2). Louisea, new genus Globonautes Bott, 1959:995, pl. 1, figs. 1- 6; 1969:359; 1970:23. Type species. —Globonautes macropus edeaensis Bott, 1969:360; 1970:24, pl. 1, figs. 3-5, pl. 26, fig. 8. Diagnosis. —Mandibular palp 2-segment- ed, terminal segment single (Fig. 3d, e). Third maxilliped lacking flagellum on ex- opod (Fig. lc). Terminal segment of gono- pod 1 directed outward, broad at base, nar- rowing sharply, final 74 almost S-shaped, tube-like, end blunt, rounded; completely lacking longitudinal groove; subterminal segment of gonopod 1 very wide at base (Fig. 1g, h). Terminal segment of gonopod 2 flagellum-like, almost as long as subter- minal segment (Fig. 1i). Dactylus of right cheliped of adult male slim, propodus with large 3-peaked proximal tooth (Fig. le). Propodus very long, as long as carapace width. Anterior dorsal margin of merus of chelipeds with 1 large jagged tooth close to distal end, rows of small pointed teeth along rest of margin (Fig. la). Carapace distinctly convex, half carapace length (mean CH/CL 124 = 0.46), carapace, anterolateral margin, lower margin of orbit, postfrontal crest, smooth (Fig. la, b). Exo-orbital, epibran- chial teeth small, low, distinct intermediate tooth present, vertical flank suture meeting anterolateral margin at epibranchial tooth (Fig. 1b). Small species, mature at CW = 22.0 mm. Distribution. —Edea (3°48'N, 10°12’E) and Yabassi (4°32'N, 9°58’E) are 90 km apart, in the rain forest zone of the Littoral Prov- ince of southwest Cameroon. Edea lies on the Sanaga River, while Yabassi lies on the Wouri River. Remarks.—This new genus of the Pota- monautidae is established to accommodate Louisea edeaensis previously considered as a subspecies of Globonautes macropus in the family Gecarcinucidae. Key to the Genera of the West African Potamonautidae 1. Terminal segment gonopod 2 very short, stump-like, '/,. length of sub- terminal segment — Terminal segment gonopod 2 very long, flagellum-like, equal in length to subterminal segment 2. Third maxilliped with long flagel- lum onexopod ........ Sudanonautes — Third maxilliped lacking flagellum Onvexopod iene Potamonemus 3. Third maxilliped with long flagel- lum on exopod; terminal segment gonopod 1 curved, tapering to pointed tip — Third maxilliped lacking flagellum on exopod; terminal segment gon- opod | weakly S-shaped, tube-like, with rounded end ........... Louisea 4. Terminal segment of gonopod 1 curving inward (toward medial line); intermediate tooth on the antero- lateral margin between the exo-or- bital tooth and the epibranchial LOOthie entrees, Aa ey Liberonautes — Terminal segment of gonopod 1 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON curving outward (away from medial line); no intermediate tooth on the anterolateral margin between the exo-orbital tooth and the epibran- chialétoothweee eee Potamonautes Louisea edeaensis (Bott, 1969) Figs. 1-3, Tables 1, 2 Globonautes macropus edeaensis Bott, 1969: 360; Bott, 1970:24, pl. 1, figs. 3-5, pl. 26, fig. 8; Cumberlidge, 1987:table 1. Material examined. —Holotype of G. m. edeaensis, adult male, CW 22.52 mm, from Edea, Cameroon, Jan 1910, ZSBS 1118/1. Two males, CWs 18.1, 13.7 mm, from Ya- bassi, Cameroon, coll. Riggenbach, ZMB 21575. Holotype of G. balssi, juvenile male, CW 12.5 mm, ZIM K-3506; 2 female para- types, CW 22.0 mm, 13.5 mm, largest fe- male ovigerous, ZIM K-3506. All G. balssi from Eosung, Bakossi highlands, Johann- Albrecht-Hohe, 1060 m, Cameroon, coll. Carl Rathke, 10 Sep 1909. Type locality. —Edea, Cameroon. Description of holotype. —Carapace (Fig. la, b): Cephalothorax ovoid, widest in an- terior third (CW/CL = 1.27), relatively high, with maximum depth in anterior region (CH/CL = 0.51). Anterior margin of front straight, curving under, front relatively nar- row, about 3 carapace width (FW/CW = 0.35). Surface of carapace smooth with no deep sutures. Postfrontal crest smooth, postorbital portions present, mid-groove broad, shallow, epigastric crests poorly de- fined, ending before meeting anterolateral margins. Exo-orbital tooth small, low, epi- branchial tooth present but almost unde- tectable. Intermediate tooth on anterolat- eral margin between exo-orbital and epibranchial teeth. Anterolateral margin of carapace smooth. Posterolateral margin curving inward, continuous with anterolat- eral margin. Posterior margin about 7 as wide as carapace width. Each flank with 2 sutures, | longitudinal, 1 vertical, dividing flank into 3 parts (Fig. VOLUME 107, NUMBER | 125 Fig. 1. Louisea edeaensis, holotype, adult male from Edea, Cameroon (CW 22.5 mm), ZSBS 1118/1. (a), whole animal, dorsal aspect; (b), carapace, frontal aspect; (c), left third maxilliped; (d), abdomen; (e), nght cheliped, frontal view; (f), left cheliped, frontal view; (g), left gonopod 1, caudal view; (h), left gonopod 1, cephalic view, (i), left gonopod 2, caudal view. Scale bar equals 10 mm (a, b, d—f), 5 mm (c), and 2 mm (g-i). 126 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON —! aie Fig. 2. Louisea edeaensis, from Yabassi, Cameroon male (CW 18.1 mm), ZMB 21575, (a), carapace, dorsal aspect; (b), carapace, frontal aspect. Scale bar equals 5 mm. 1b). Longitudinal suture dividing suborbital and subhepatic regions from pterygostomial region, beginning at respiratory opening and curving backward across flank. Short ver- tical suture dividing suborbital region from subhepatic region (Fig. 1b); suture begin- ning at epibranchial tooth, curving forward under intermediate tooth, then curving sharply down meeting longitudinal suture, continuing round to medical corner of lower orbital margin, marked by row of granules. Groove between sternal segments 2 and 3 complete; groove between sternal segments 3 and 4 consisting of 2 small notches at sides of sternum. Third maxillipeds filling entire oral field, except for transversely oval effer- ent respiratory openings in superior lateral corners. Exopod of third maxilliped lacking flagellum (Fig. 1c). Ischium of third max- illiped smooth, with faint vertical groove. Mandibular palp 2-segmented, terminal segment single, undivided, rudimentary curved flange at junction between segments, supporting fringe of long, soft hairs (Fig. 3d). Abdomen triangular, sides not indent- ed, terminal segment rounded at distal mar- gin (Fig. 1d). Chelipeds (Fig. la, e, f) greatly unequal, right longer (22.6 mm), higher (10.2 mm) than left (14.9 mm, 5.5 mm respectively). Dactylus of right cheliped narrow, teeth small, enclosing long narrow space when closed; propodus with large 3-peaked prox- imal tooth, smaller teeth distally, palm swollen. Propodus very long, as long as car- apace width. Anterior dorsal margin of me- rus of right and left chelipeds with 1 large jagged tooth close to distal end, rows of small round teeth along rest of margin, outer mar- gin with | row of fainter granules. Carpus of cheliped with 2 large pointed teeth, first smaller than second. Left cheliped showing VOLUME 107, NUMBER 1 less enlargement, dactylus also narrow, en- closing narrow space, teeth smaller than those of right cheliped. Walking legs (pe- reiopods 2—S) slender, P4 longest, P5 short- est. Dactyli P2—5 tapering to point, each bearing rows of downward-pointing sharp bristles, dactylus of P5 shortest of the 4 legs. Terminal segment of gonopod | (Fig. 1g— i) directed outward, broad at base, final narrow, weakly S-shaped, tube-like, end blunt, rounded; completely lacking longi- tudinal groove and bristles; subterminal segment gonopod 1 very wide at base. Cau- dal face of subterminal segment forming raised triangular flap extending halfway across segment, flap tapering diagonally to point at junction with terminal segment, forming roof of chamber for gonopod 2; ce- phalic face of subterminal segment narrow, forming lower floor of chamber for gonopod 2, extending outward forming wide plat- form. Terminal segment of gonopod 2 (Fig. li) flagellum-like, extremely long. Subter- minal segment gonopod 2 wide at base, oth- erwise a long, thin, slightly tapering, upright process supporting long terminal segment. Variation. —Male from Yabassi (Fig. 2a, b) with large tooth close to distal end of anterior dorsal margin of merus of che- lipeds, but this tooth not as large as in ho- lotype. First of two large pointed teeth on carpus of cheliped larger than second in male from Yabassi. Postfrontal crest well defined, meeting anterolateral margins in Yabassi specimens, in contrast to that of holotype. Distribution. —Between the Wouri and Sanaga rivers in the rain forest zone of southwest Cameroon. Size. — Measurements given in Table 1. Etymology. —The genus Louisea is named for my wife, Dr. Louise M. Bourgault, a Professor of Mass Communications at Northern Michigan University, Marquette, Michigan, U.S.A., in recognition of her ded- icated support of African freshwater crab biology, an effort which she has maintained for more than twelve years. Remarks. —Globonautes macropus edea- 127 ensis was briefly described by Bott (1969), without illustrations, and assigned to the Gecarcinucidae. A more detailed descrip- tion, including photographs of the carapace and gonopod 1, appeared in a later work (Bott 1970). There have been no further re- ports of Bott’s taxon since that time, other than brief comparison with species of Glo- bonautes by Cumberlidge (1987). Bott’s tax- on is here elevated to specific rank, and placed in the new genus Louisea. It is interesting to speculate on why L. edeaensis and G. balssi were originally as- signed to the Gecarcinucidae by earlier workers. One possibility could be that the small hard flange at the junction between segments of the mandibular palp of these taxa (which is partly obscured by a fringe of long hairs) was counted as a second lobe. For example, Bott (1955, 1965) considered this feature to warrant subfamilial recog- nition for Madagascan crabs of the genus Hydrothelphusa A. Milne-Edwards, 1872, and erected the Hydrothelphusinae Bott, 1955, a subfamily of the Gecarcinucidae, to accommodate this genus. However, similar small flanges between the segments of the mandibular palp are also found in Suda- nonautes orthostylis (Bott 1955) and S. flow- eri (De Man 1901) and this feature has not been judged to warrant familial recognition for these species which are all in the Pota- monautidae (Rathbun 1921, Cumberlidge 1993a). It should be noted that the Gecar- cinucidae and Pseudothelphusidae possess a mandibular palp with a large, distinct sec- ond lobe which is a separate, hardened pro- cess resting on the anterior face of the man- dible. Generic Comparisons Differences in the length of the terminal segment of gonopod 2, the form of the ter- minal segment of gonopod 1, and the form of the third maxilliped are diagnostic in dis- tinguishing between Sudanonautes, Libero- nautes, Potamonautes, and Potamonemus 128 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 3. Louisea edeaensis, from Yabassi, Cameroon male (CW 18.1 mm), ZMB 21575, (a), right cheliped, frontal view; (b), left cheliped, frontal view; (c), left third maxilliped; (d), left mandible anterior view; (e), left mandible posterior view; (f), merus and carpus of right cheliped superior view; (g), left gonopod 1, caudal view; (h), left gonopod 1, cephalic view; (i), left gonopod 2, caudal view. Globonautes balssi from Johann-Albrecht- Hohn, Cameroon, paratype, adult female (CW 22.0 mm), ZIM K-3506; (j), right third maxilliped; (k), left mandible anterior view. Globonautes balssi from Johann-Albrecht-Hohn, Cameroon, male holotype (CW 12.5 VOLUME 107, NUMBER | 129 Table 1.—Carapace measurements (mm) and proportions relative to body size (CL) for all known specimens of Louisea edeaensis and Globonautes balssi from Cameroon, West Africa. Sex CW CL Louisea edeaensis Holotype, ZSBS 1118/1 1.M WD,S 17.8 ZMB, 21575 2. M 18.1 13.1 3. M 13.7 10.1 Globonautes balssi Holotype, ZIM K3506 1. M Guv) 13.5 10.5 Paratypes, ZIM K3506 2. F (ad, ovig) 22.0 15.5 3. F Guv) 12.5 9.5 CH FW CW/CL CH/CL FW/CL QI 7.93 1.27 0.51 0.45 7.4 6.3 1.38 0.41 0.48 4.8 4.8 1.36 0.48 0.48 5.5 5.0 1.29 0.52 0.48 10.0 7.0 1.42 0.65 0.45 5.0 4.5 1.32 0.53 0.47 (Bott 1955, Cumberlidge & Clark 1992). Of these four genera, Louisea most closely re- sembles Potamonautes and Liberonautes, since gonopod 2 in species of these genera has a long, flagellate, terminal segment, and the mandibular palp has a single-lobed ter- minal segment. However, characters of the third maxilliped and gonopod | clearly dis- tinguish Louisea from these two genera. In both Potamonautes and Liberonautes the exopod of the third maxilliped possesses a flagellum, and the terminal segment of gon- opod | curves evenly and tapers to a pointed tip (Bott 1955, Cumberlidge & Sachs 1989a, 1989b). In Louisea the exopod of the third makxilliped lacks a flagellum, and the ter- minal segment of gonopod 1 is S-shaped, tube-like, and has a blunt, rounded end. Li- beronautes can further be eliminated on bio- geographic grounds since members of this genus are not found east of Ghana (Cum- berlidge & Sachs 1989a, 1989b). The other two potamonautid genera oc- curring in Cameroon (Sudanonautes and Potamonemus) possess a second gonopod with a short terminal segment, and are —_— therefore not closely related to Louisea. Members of Sudanonautes can be distin- guished from Louisea using the same char- acters of the third maxilliped and gonopod 1 outlined above for Potamonautes and Li- beronautes. While Potamonemus and Lou- isea both lack a flagellum on the exopod of the third maxilliped, they can be clearly dis- tinguished by the length of the terminal seg- ment of gonopod 2: it is extremely short in Table 2.— Measurements (mm) of right and left che- liped length (RCL and LCL), and height (RCH and LCH) for all known specimens of Louisea edeaensis and Globonautes balssi from Cameroon, West Africa. Sex CW RCL LCL RCH LCH Louisea edeaensis Holotype, ZSBS 1118/1 1. M 22.5 22.6 14.9 10.2 5.5 ZMB, 21575 2. M 18.1 15.4 7.3 So 3. M 1S IO Sie ah 25, Globonautes balssi Paratype, ZIM K3506 1. F(ad, ovig) 22.0 14.0 140 5.5 5.0 mm), ZIM K-3506; (1), right gonopod 1, caudal view, based on Bott (1959, Fig. 7). Scale bar equals 5 mm (d-f, k), 2 mm (a-c, g-i), and 2 mm (j). 130 Potamonemus and extremely long in Lou- isea. For the above reasons, L. edeaensis has been removed from its former designation as a subspecies of Globonautes macropus in the family Gecarcinucidae, and placed in the new monotypic genus Louisea in the family Potamonautidae. The single-lobed terminal segment of the mandibular palp of G. balssi led Cumber- lidge (1987) to doubt its assignment to the Gecarcinucidae, and therefore, to Globo- nautes. Indeed, the form of the mandibular palp, and the lack of a flagellum on the ex- opodite of the third maxilliped of G. balssi, position this taxon close to either Louisea or to Potamonemus. The best way to dis- tinguish between these two genera is by an examination of gonopod 2: a short terminal segment would place G. balssi in Potamo- nemus, while a long, flagellate terminal seg- ment would place G. balssi in Louisea. However, gonopod 2 of G. balssi is missing on both sides in the holotype, the only male specimen. For this reason G. balssi is re- garded as incertae sedis until more material is available. Despite these differences the following important characters are shared by both G. balssi and L. edeaensis: (1) the mandibular palp is 2-segmented and the terminal seg- ment is a single lobe (Fig. 3d, e, k); (2) the exopod of the third maxilliped lacks a fla- gellum (Figs. lc, 3j); (3) the terminal seg- ment of gonopod | is weakly S-shaped, tu- bular, and is not grooved (Figs. 1g, h, 31); (4) there is a distinct intermediate tooth on the anterolateral margin between the exo- orbital and epibranchial teeth (Figs. la, b, 2a, b); (5) the vertical flank suture on the flank begins at the epibranchial tooth, curves forward under the intermediate tooth, and finally curves sharply down to meet the lon- gitudinal suture on the flank (Figs. 1b, 2b); (6) the postfrontal crest is smooth, poorly defined, and ends before meeting the an- terolateral margins (Fig. la, b); (7) the groove between sternal segments 2 and 3 is com- plete, while the groove between sternal seg- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ments 3 and 4 consists of 2 small notches at the sides of the sternum; (8) the exo-or- bital tooth is small and low, the epibranchial tooth is present but is almost undetectable (Figs. la, b, 2a, b); (9) the carapace is rel- atively high (CH/CL = 0.51 L. edeaensis, 0.65 G. balssi); and (10) both species are small, reaching maturity at CW 22.0 mm. The following are reasons why G. balssi is not included here in Potamonemus: (1) the terminal segment of gonopod 1 of Po- tamonemus is evenly curved, with a clear longitudinal groove (Cumberlidge & Clark 1992, Cumberlidge 1993b), whereas that of L. edeaensis and G. balssi is S-shaped, tu- bular, and lacks a groove (Figs. lg, h, 31); (2) the anterolateral margin of Potamone- mus lacks an intermediate tooth on the an- terolateral margin, whereas both L. edeaen- sis and G. balssi possess a small but distinct intermediate tooth (Figs. la, b, 2a, b); (3) the carapace of Potamonemus is relatively flatter (CH/CL = 0.47-0.49, Cumberlidge 1993b) than that of both L. edeaensis and G. balssi (CH/CL = 0.51 and 0.65 respec- tively). Finally, the most recent specimens of L. edeaensis were collected in 1910, and the species has not been encountered since. This is undoubtedly due in part to difficulties in the identification of freshwater crabs from Cameroon, but it may also indicate that Louisea is a rare, and possibly endangered, rain forest species. Acknowledgments Iam very grateful to Dr. L. Tiefenbacher, of the Zoologische Sammlung des Bayer- ischen Staates, Munchen, Germany for loaning the holotype of Globonautes m. edeaensis. | acknowledge the kind hospital- ity and helpfulness of Dr. Hartmann and Dr. G. Andre of the Zoological Institute and Museum, Hamburg, Germany. I also thank Prof. Dr. H. E. Gruner of the Zoologische Museum of the Humboldt-Universitat, Berlin for loaning specimens of Louwisea VOLUME 107, NUMBER | edeaensis, and for his helpful cooperation during a visit. I especially thank artists Anne C. Martin and Jon C. Bedick of Northern Michigan University, U.S.A., for their skill and patience in producing the illustrations used in this paper. Part of this work was supported by a Faculty Grant from North- ern Michigan University, Marquette, Mich- igan, U.S.A. Literature Cited Bott, R. 1955. Die Siisswasserkrabben von Afrika (Crust., Decap.) und ihre Stammesgeschichte. — Annales du Musée Royal du Congo Belge, (Ter- vuren, Belgique) C. Zoologie 1(3, 3):209-352. 1959. Potamoniden aus West-Afrika.—Bul- letin de I’Institut Fondamental D’Afrique Noire, Série A 21 (3):994—1008. 1965. Die Sisswasserkrabben von Madagas- kar.—Bulletin de Muséum national d’Histoire naturelle, Paris, 2, 27 (2):335-350. 1969. Die Flusskrabben aus Asien und ihre Klassificaten (Crustacea, Decapoda).—Senck- enbergiana Biologisches, Frankfurt am Main 50: 359-366. 1970. Betrachtungen tber die Entwicklungs- geschichte und Verbreitung der Siisswasser- Krabben nach der Sammlung des Naturhisto- rischen Museums in Genf/Schweiz.— Revue Suisse de Zoologie, 77(2), 24:327-344. Cumberlidge, N. 1987. Notes on the taxonomy of West African gecarcinucids of the genus G/o- bonautes Bott, 1959 (Decapoda, Brachyura).— Canadian Journal of Zoology 65(9):2210-2215. 1989. Redescription of Sudanonautes or- thostylis (Bott, 1955), a freshwater crab from Nigeria, Cameroon and Ghana (Decapoda, Po- tamonautidae), with notes on its ecology.— Crustaceana 56(3):230-245. . 1993a. Redescription of Sudanonautes gran- ulatus (Balss, 1929) (Potamoidea: Potamonau- tidae) from West Africa.—Journal of Crusta- cean Biology (in press). 1993b. Two new species of Potamonemus Cumberlidge & Clark, 1992 (Brachyura: Pota- moidea: Potamonautidae) from the rain forests of West Africa.—Journal of Crustacean Biology 13(3):283-307. ——,, & P. Clark. 1992. A new genus and species of freshwater crab from Cameroon, West Africa (Crustacea: Brachyura: Potamoidea: Potamo- nautidae).— Bulletin of the British Museum of Natural History (Zoology), London 58(2):149- 156. 131 —., & R. Sachs. 1989a. A key to the crabs of Liberian freshwaters.— Zeitschrift fir Ange- wandte Zoologie 76:221-229. 1989b. Three new subspecies of the West African freshwater crab Liberonautes latidac- tylus (DeMan, 1903) from Liberia, with notes on their ecology.—Zeitschnft fur Angewandte Zoologie 76:425-439. De Man, J.G. 1901. Description of a new freshwater Crustacea from the Soudan; followed by some remarks on an allied Species.— Proceedings of the Zoological Society of London 94—104 (pages). MacLeay, W.S. 1838. Illustrations of the Zoology of South Africa; being a Portion of the Objects of Natural History Chiefly Collected During an Ex- pedition into the Interior of South Africa, under the Direction of Dr. Andrew Smith, in the Years 1834, 1835, and 1836; Fitted Out by “The Cape of Good Hope Association for Exploring Cen- tral Africa.” In A. Smith, Illustrations of the Zoology of South Africa; Consisting Chiefly of Figures and Descriptions of the Objects of Nat- ural History Collected During an Expedition into the Interior of South Africa, in the Years 1834, 1835, and 1836; Fitted Out by “The Cape of Good Hope Association for Exploring Central Africa.” (Invertebrates). 75 pp., 4 pls. London. Milne-Edwards, A. 1872. Note sur les crabes d’eau douce de Madagascar.—Bibliographie Ecole Hautes Etudes (Séction Sciences naturelles), Paris 5(8):1-3. Monod, T. 1977. Sur quelques crustacés Décapodes africains (Sahel, Soudan). — Bulletin de Muséum national d’ Histoire naturelle, Paris 3, 500:1201- 1236. 1980. Décapodes. Jn J-R. Durand and C. Léveque, ed., Flore et Fauna Aquatiques del’Af- rique Sahelo-Soudanienne. ORSTOM, Paris, I.D.T. 44(1):369-389. Rathbun, M. 1898. Descriptions of three species of freshwater crabs of the genus Potamon.—Pro- ceedings of the Biological Society of Washington 12:27-30. 1904. Les crabes d’Eau Douce (Potamoni- dae).— Nouvelles Archives du Muséum d’His- toire naturelle, Paris (4)6:255-312. 1921. The brachyuran crabs collected by the American Museum Congo expedition 1909- 1915.—Bulletin of the American Museum of Natural History 43:379-474. Department of Biology, Northern Mich- igan University, Marquette, Michigan 49855, U.S.A. PROC. BIOL. SOC. WASH. 107(1), 1994, pp. 132-136 FREDIUS STENOLOBUS, A NEW SPECIES OF FRESHWATER CRAB (DECAPODA: BRACHYURA: PSEUDOTHELPHUSIDAE) FROM THE VENEZUELAN GUIANA Gilberto Rodriguez and Héctor Suarez Abstract. —Fredius stenolobus, a new species of pseudothelphusid crab, is described from the Caura River basin in the Venezuelan Guiana. The species closely resembles Fredius beccarii (Coifmann, 1939), but can be easily distin- guished by the characteristic cephalic process of the first male gonopod and by the apical spinulation of the second male gonopod. SEM microphotographs of these appendages are provided for both species. In a recent revision of the genus Fredius, Rodriguez & Pereira (1992) discussed the systematics and distribution of this genus in northern South America, and advanced an hypothesis relating to the origin and ra- diation of its species. Recent collections in the Caura River and some of its tributaries revealed another new species of Fredius. The material is deposited at the Museo de Historia Natural La Salle, Caracas (MHNLS) and the Museo de Biologia, Universidad Central de Venezuela, Caracas (MBUCV). Other abbreviations used are cb for carapace breadth, and cl for carapace length. Family Pseudothelphusidae Ortmann, 1893 Tribe Kingsleyini Bott, 1970 Genus Fredius Pretzmann, 1967 Fredius stenolobus, new species Fig. 1A—D; 2A-—C; 3A, B Material. —Rio Mojagua, affluent of Rio Erebato, Estado Bolivar; E. Leon; 7 Mar 1992, 1 male holotype, cb 81.3 mm, cl 51.5 mm, 2 male paratypes, cl 18.8 and 25.0 mm, cb 62.5 and 38.7 mm, 3 female para- types, cl 37.8, 35.3 and 33.5 respectively, cb 59.0, 54.4 and 51.8 mm respectively (MHNLS 1267).—Rio Caura, Entrerios, 5°57'15"N, 64°25'30”"W, 350 m alt., Estado Bolivar; 25 May 1989; H. Castellanos; 1 male, cl 42.7 mm, cb 64.4 mm, | female, cl 54.8 mm, 86.8 mm (MBUCV XI-2923).— El Raudal, Cano Cambur, 15 km SW from Maripa, Estado Bolivar; 21 Mar 1985; J. Medina; | immature female, cl 35.7 mm, cb 54.5 mm (MBUCV XI-901). Diagnosis. —Cephalic lobe of first gono- pod auriculariform, narrow in cephalic view, projected distally into triangular spine; me- sial border of cephalic lobe sinuous, ending proximally in rounded projection covered with spinules; lateral border of cephalic lobe ending distally in oblong rounded lobe. Me- sial lobe well developed, wide, rounded, thumb-like, with bifid apex and large tri- angular cephalic spine on distal margin. Description. —(based on holotype and 5 paratypes) Cervical groove deep, narrow, slightly sinuous, ending away from lateral margin. Anterolateral margin with depres- sion behind anteroexternal angle, followed by 3-4 papillae; rest of margin behind cer- vical groove with approximately 15 blunt teeth, decreasing in size posteriorly. Post- frontal lobes low, wide, continued laterally as faint ridges; median groove wide and shallow between postfrontal lobes, obsoles- cent or absent near frontal margin. Surface of carapace in front of postfrontal lobes slightly excavated in frontal view and in- clined anteriorly. Upper border of front straight or slightly bilobed in dorsal view, VOLUME 107, NUMBER | 133 Fig. 1. a2mm Fredius stenolobus, new species, holotype: A, Dorsal view of carapace and pereiopods: B, Chela of largest cheliped, external view; C, Third maxilliped, left; D, Aperture of efferent channel, left. carinated, marked with row of indistinct pa- pillae, median notch absent or inconspic- uous. Lower margin thin and strongly sin- uous in frontal view. Surface of front between upper and lower borders high, ex- cavated, retracted backwards. Surface of carapace with numerous large papillae vis- ible to the naked eye on hepatic and bran- chial regions. Similar papillae present on dorsal surface of pereiopods. Palm of larger chela elongated, not con- spicuously inflated, fingers strongly arched inwards, gaping at base. Exopod of third maxilliped 0.30 length of ischium of en- dognath. First gonopod robust at base, strongly ta- pering to subapical bulge, with marginal, cephalic, and mesial lobes well developed. Marginal lobe truncate, not extending over field of apical spines. Cephalic lobe auricu- lariform in cephalic view, projected distally into triangular spine (Fig. 2C, ds), mesial border sinuous, continued proximally as rounded projection with spinules; lateral PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON A ASAS1 2EOkKY oS Sa Fig. 2. Apical portion of left first gonopod: A-C, Fredius stenolobus, new species, holotype; D, Fredius beccarii (Coifmann 1939), MBUCV XI-2928. A, D, caudal view; B, meso-caudal view; C, cephalic view; al, accessory lobe; cs, cephalic spine; ds, distal spine; ml, mesial lobe. VOLUME 107, NUMBER 1 Le Fig. 3. 135 Left second gonopod: A, B, Fredius stenolobus, new species, holotype; C, D, Fredius beccarii (Coifmann 1939), MBUCV XI-2928. A, C, Terminal portion; B, D, Detail of apex. Scale in D same as scale in B. border with rounded accessory lobe (Fig. 2A, B, al); field of apical spines wide, di- rected to cephalic side. Mesial lobe wide, rounded, thumb-like, with bifid apex, and large triangular cephalic spine on distal margin (Fig. 2A, cs). Marginal setae ar- ranged in dense row over proximal half of gonopod; lateral surface with numerous long plumose setae. Second gonopod with nu- merous spinules on distal portion; tip cup- shaped, with relatively strong spines di- rected distally. Color.—Holotype preserved in alcohol dark brown on the dorsal surface of the car- apace and walking legs; anterior portion of the carapace a darker shade. The chelae are dark brown on the upper portions, brown- orange on the inner surface; the distal por- tions of the dactyli are black. The ventral surface of the carapace and appendages is light brown. Etymology. —The specific name is from the Greek “‘stenos,”’ narrow, and “lobos,” lobe, and refers to the narrow cephalic lobe. Fredius beccarii (Coifmann, 1939) Figs. 2D; 3C, D Pseudothelphusa beccarii Coifmann, 1939, p. 98, figs. 2, 4a, pl. 3, 1, 2. Guinotia (Neopseudothelphusa) beccarii, Pretzmann, 1967, p. 24. Eudaniela (Aspockia) beccarii beccarii, Pretzmann, 1971, p. 16. Eudaniela (Aspoeckia) beccarii beccarii, Pretzmann, 1972, p. 18, figs. 103-104. Pseudothelphusa contorta Rodriguez, 1966, p. 263, fig. 3, pl. 2. Eudaniela (Aspockia) beccarii contorta, Pretzmann, 1971, p. 16. Eudaniela (Aspoeckia) beccarii contorta, Pretzmann, 1972, p. 19, figs. 25-28, 73, 74. Guinotia (Neopseudothelphusa) beccarii cu- yunis Pretzmann, 1967, p. 23. Fredius beccarii, Rodriguez & Pereira, 1992, p. 304, fig. 4M, N. Material. —Rio Cuyuni, Estado Bolivar; 1987; A. Machado (MBUCV XI-2928). 1 136 male (cl 39.7 mm, cb 63.0 mm; Venezuela, without other data; (MBUCV XI-0838). [Other literature records for this species are given by Rodriguez (1982)]. Remarks. —The first gonopod of Fredius stenolobus closely resembles that of Fredius beccarii, but in this latter species the ce- phalic lobe is wider in cephalic view, the accessory lobe is absent, and the proximal spiny projection of its mesial margin is more developed. The cephalic spine of the first gonopod of F. beccarii is stronger and di- rected caudally, and the lateral lobe is larger than in F. stenolobus. The morphology of the second male gonopod has been rarely used in pseudothelphusid systematics. As our SEM photographs show, there are con- spicuous differences between the second gonopods of both species. The cup-shaped tip differs in the arrangement of the spines and the conspicuous spinulation on the dis- tal portion observed in F. stenolobus is ab- sent in F. beccarii. The key to the species of Fredius in Rod- riguez & Pereira (1992) should be expanded after couplet 2 to allow for the new species as follows: Cephalic lobe narrow, accessory lobe prominent Fredius stenolobus, new species Cephalic lobe very wide, accessory lobe absent Literature Cited Bott, R. 1970. Betrachtungen tiber die Entwicklungs- geschichte und Verbreitung der Stsswasser- Krabben nach der Sammlung des Naturhisto- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON rischen Museums in Genf/Schweiz.— Revue Suisse de Zoologie 77 (fascicule 2, 24):327—244. Coifmann, I. 1939. Potamonidi della Guicana Inglese raccolti dal Prof. Nello Beccari.—Archivio Zoologico Italino 27:93-117, pl. 3. Ortmann, A. 1893. Die Dekapoden-Krebse des Strassbourg Museums, mit besonderer béruck- sichtingung der von Herr Déderlein bei Japan und bei den Liu-Kiu-Inseln gesammelten und zur Zeit in Strassburger Museum aufbewarten Formen. VII. Theil. Abteilung: Brachyura (Brachyura genuina Boas) II. Unterabteilung: Cancroidea, 2 Section: Cancrinea, 1. Gruppe: Cyclometopa. —Zoologische Jaarbiicher, Abtei- lung fiir Systematik, Geographie und Biologie der Tiere 7:41 1-495, pl. 17. Pretzmann, G. 1967. Uber einige siidamerikanische SiiBwasserkrabben (Pseudothelphusidae). Vor- laufige Mitteilung.—Entomologische Nachri- chtenblatt, Wien 14(2):23-26. 1971. Fortschritte in der Klassifizierung der Pseudothelphusidae.—Sitzungsberichten der Osterreich Akademie der Wissenschaften, Mathematisch-Naturwissenschaftliche Klasse (1) 179(1-4):14—20. 1972. The Pseudothelphusidae (Crustacea Brachyura).— Zoologica 42(120) pt. 1:1-182, textfig. 1-31, 732 figs. A.A. Rodriguez, G. 1966. Three new species of Pseudothel- phusa from Venezuela.—Zoologische Medede- lingen 41:259-267. 1982. Les Crabes d’eau douce d’Ameérique. Famille des Pseudothelphusidae.— Faune Tro- picale 22 ORSTOM: 1-223. —, & G. Pereira. 1992. New species, cladistic relationships and biogeography of the genus Fredius (Crustacea: Decapoda: Pseudothothel- phusidae) from South America.—Journal of Crustacean Biology 12:298-311. Centro de Ecologia, Instituto Venezolano de Investigaciones Cientificas, Apdo. 21827, Caracas 1010A, Venezuela. PROC. BIOL. SOC. WASH. 107(1), 1994, pp. 137-150 A NEW CALCINUS (DECAPODA: ANOMURA: DIOGENIDAE) FROM THE TROPICAL WESTERN ATLANTIC, AND A COMPARISON WITH OTHER SPECIES OF THE GENUS FROM THE REGION Néstor H. Campos and Rafael Lemaitre Abstract.—A new species of a diogenid hermit crab, Calcinus urabaensis, is described from the Gulf of Uraba, on the Caribbean coast of Colombia. The new species is the third in the genus described from the western Atlantic, and can be distinguished from the other two known species of the genus Calcinus in the region, C. tibicen (Herbst) and C. verrilli (Rathbun), by differences in coloration.and armature of the dactyl of the left cheliped, third pereopod, and telson. A comparison of the three species is included. Resumen. —Se describe una nueva especie de cangrejo ermitano pertenecienta a la familia Diogenidae, Calcinus urabaensis, colectada en el Golfo de Uraba, Caribe sur. La nueva especie es la tercera conocida de este género en el Atlantico occidental, y se diferencia de las otras dos especies del género Calcinus de la region, C. tibicen (Herbst) y C. verrilli (Rathbun), en la coloracion y espinas del dactilo de la quela izquierda, tercer pereOpodo, y telson. Se presenta una comparacion de las tres especies. Compared to other tropical regions of the world oceans, the western Atlantic contains very few species of the diogenid genus Cal- cinus Dana, 1852. Recent studies of Cal- cinus species in the Pacific, for example, have shown that nine species occur on the Ha- waiian Islands (Haig & McLaughlin 1984), 11 species on the Mariana Islands (Wooster 1984), and 17 species on the Australian coast (Morgan 1991). In contrast, only two spe- cies have been described from the western Atlantic, C. tibicen (Herbst, 1791), broadly distributed from Florida to Brazil, including Bermuda; and C. verrilli (Rathbun, 1901), considered endemic to Bermuda (Verrill 1908, Provenzano 1960, Markhan 1977, Chace et al. 1986). Two other taxa also de- scribed from the western Atlantic, Calcinus sulcatus (H. Milne Edwards, 1836) and C. formosus Neumann, 1878, are considered junior synonyms of C. tibicen (see Proven- zano 1959; McLaughlin, pers. comm.). In 1985, during an expedition to the Gulf of Uraba, on the Caribbean coast of Colom- bia (Campos & Manyarrés 1988), the senior author collected a male hermit crab be- lieved to represent an undescribed species of Calcinus. The coloration of the Gulf of Uraba specimen was most similar to that of C. tibicen, but morphologically the spec- imen was closest to C. verrilli. Because com- parative material was not easily available to the senior author, the specimen was sent to the junior author who compared it with material of C. verrilli deposited in various museums and institutions in the United States. He too concluded that the specimen represented an undescribed species, the third in the genus Calcinus from the western At- lantic. Given that only one specimen exist- 138 ed, the preparation of a manuscript was de- layed awaiting the possibility of collecting additional material. However, various col- lecting efforts since 1985 failed to produce any additional material, and we now de- scribe this new species based on the male from the Gulf of Uraba. As pointed out by Haig & McLaughlin (1984), species of Calcinus are morpholog- ically very similar, and difficult to identify, particularly if color patterns have faded away. For this reason, it is appropriate to present along with the description of the new species, a comparison of the now three western Atlantic species of Calcinus. For this purpose, representative material of C. tibicen and C. verrilli were also examined. Illustrations of selected structures with di- agnostic importance are included for all three species. The material used remains deposited in the collections of the Indian River Coastal Zone Museum, Harbor Branch Oceano- graphic Foundation, Fort Pierce, Florida (IRCZM); Rosenstiel School of Marine and Atmospheric Sciences, University of Miami (UMML), and the National Museum of Natural History, Smithsonian Institution, Washington, D.C. (USNM). The following abbreviations are used: SL, shield length (to the nearest 0.1 mm), measured from the tip of the rostrum to the midpoint of the pos- terior margin; juv: juvenile (s); ovig(s): ovig- erous; RHG, Robert H. Gore; AJP, An- thony J. Provenzano, Jr.; WLS, Waldo L. Schmitt; sta, station. Calcinus urabaensis, new species Figs. la, 2a, 3a, 4a, 5a, b, 6a, 7a, d, g Calcinus sp.—Campos & Manjarrés, 1988: 19. Material examined. —Holotype: 6 SL 5.6 mm, inlet of Pinorroa, Gulf of Uraba, Co- lombia, on rocks, ~3 m, USNM 251886. Description of holotype. —Shield (Fig. 1a) slightly longer than wide, with scattered short setae; anterior margin weakly con- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON cave; lateral projections obtusely triangular, with small terminal spine; anterolateral margins sloping. Rostrum produced, sub- triangular, slightly in advance of lateral pro- jections. Anterodorsal plate of branchioste- gite (Fig. 2a) with acute anteroventral angle. Ocular peduncles (Fig. la) long, slender (about 8.6 times as long as wide), and sub- equal to shield length; peduncles naked ex- cept for few short setae proximally on dorsal face, slightly curved outward (viewed lat- erally). Acicles terminating in strong spine; separated basally by '2 basal width of 1 aci- cle. Antennular peduncles (Fig. 2a) reaching almost to base of corneae, naked except for tuft of setae proximally on penultimate seg- ment. Ultimate segment 1.3 times as long as penultimate. Basal segment with mesial face unarmed; lateral face with distal margin armed with 4 (left) or 3 (right) small spines and | small submarginal spine. Antennal peduncles (Fig. 1a, 2a) reaching to about distal 74 of ocular peduncles, seg- ments with scattered setae. Supernumerary segment present. Fifth segment slightly curved outward. Fourth segment with dor- sodistal spine. Third segment with strong ventrodistal spine. Second segment with dorsolateral distal angle produced, termi- nating in strong bifid spine, lateral margin unarmed; dorsomesial distal angle with strong spine. First segment with 4 small spines on ventrodistal margin, lateral face unarmed. Acicles slightly exceeding distal margin of fourth antennal segment, termi- nating in strong spine, dorsomesial margin with 2 (left) or 3 (right) spines, and 2 spines on dorsolateral margin. Flagellum almost reaching to distal end of extended left che- liped, minutely setose. Third maxilliped with crista dentata formed of row of 25 small corneous teeth. Left cheliped (Fig. 3a) with outer face of merus, carpus, and chela lacking setae. Fin- gers not leaving gap when closed, with spoon-shaped tips; cutting edges with irreg- ularly-sized calcareous teeth and tufts of se- VOLUME 107, NUMBER 1 139 Fig. 1. tae. Dactyl with row of small spines on up- per margin; outer face with irregular rows of low, closely-set tubercles. Fixed finger with row of corneous-tipped spines on lower margin; lower outer face with low tubercles similar to those on dactyl but smaller (spines and tubercles continued on lower outer face of palm). Palm 1.7 times as long as wide; upper margin with row of small, low pro- tuberances; outer and inner faces smooth except for scattered tufts of short setae on inner face. Carpus with upper margin armed with 3 small spines on distal half and strong distal spine; outer distal margin with row of small tubercles on upper 1%; outer face with prominent tubercle proximally on up- per half; inner face smooth. Merus subtrian- Shield and cephalic appendages. a, Calcinus urabaensis, new species (antennular peduncles hidden under eyestalks); b, Calcinus verrilli (Rathbun, 1901); c, Calcinus tibicen (Herbst, 1791). Scales equal | mm. gular in cross-section, upper margin with scattered short setae; inner and outer lower margins with row of 2 or 3 small spines distally; outer and inner faces smooth. Right cheliped (Fig. 4a) slightly over- reaching proximal margin of dactyl of left cheliped. Fingers with spoon-shaped tips, with scattered tufts of setae, and leaving no gap when closed; cutting edges each with 3 strong calcareous teeth. Dactyl with row of corneous-tipped spines directed anteriorly on upper margin. Fixed finger with 4 small spines on outer face proximally. Palm with upper margin raised in form of crest, and armed with 5 strong, corneous-tipped spines; outer face with scattered setae, and 2 short oblique rows of corneous-tipped spines dis- 140 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. Right antennular and antennal peduncle, and anterodorsal portion of branchiostegite, lateral view. a, Calcinus urabaensis, new species; b, Calcinus verrilli (Rathbun, 1901); c, Calcinus tibicen (Herbst, 1791). Scales equal 1 mm. (1-4: antennal segments; ac: antennal acicle; ap: anterodorsal plate of branchiostegite; b: basal antennular segment; s: supernumerary segment.) tally; inner face smooth. Carpus armed on upper margin with 4 corneous-tipped spines, increasing in size distally; outer face with scattered minute tubercles; inner face smooth. Merus subtriangular, upper margin sparsely setose; inner lower margin with 6 spines; outer and mesial faces smooth. Ambulatory legs slightly asymmetrical, left shorter and with propodi broader (viewed laterally) than right. Second pereo- pod (Fig. 5a) exceeding extended left che- liped by about % length of dactyl. Dactyl subequal in length to propodus, terminating in sharp corneous claw, and with tufts of simple setae on mesial, dorsal, and lateral faces; ventral margin armed with 7 short corneous spines. Propodus with dorsodistal, corneous-tipped spine, and ventrodistal spine; outer and ventral faces with tufts of simple setae. Carpus with strong dorsodis- tal, corneous-tipped spine; ventral margin with small spine at about midlength of mar- VOLUME 107, NUMBER | 141 Fig. 3. Left cheliped, lateral view. a, Calcinus urabaensis, new species; b, Calcinus verrilli (Rathbun, 1901); c, Calcinus tibicen (Herbst, 1791). Scales equal 1 mm. 142 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 4. Right cheliped, lateral view. a, Calcinus urabaensis, new species; b, Calcinus verrilli (Rathbun, 1901); c, Calcinus tibicen (Herbst, 1791). Scales equal 1 mm. VOLUME 107, NUMBER 1 143 Fig. 5. Ambulatory legs, lateral view. a, b, Calcinus urabaensis, new species: a, right second pereopod; b, right third pereopod. Calcinus verrilli (Rathbun, 1901): c, left third pereopod. Calcinus tibicen (Herbst, 1791): d, left third pereopod. Scales equal 1 mm. (Heavily stippled areas represent color pattern, except for furrow indicated by arrow.) 144 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 6. Dactyl of right third pereopod, lateral view. a, Calcinus urabaensis, new species; b, Calcinus verrilli (Rathbun, 1901); c, Calcinus tibicen (Herbst, represent color pattern.) gin, and long setae. Merus with spine dis- tally on ventrolateral margin; ventral mar- gin with row of small spines, and several tufts of long simple setae. Third pereopod (Fig. 5b, 6a) slightly shorter, and with segments broader (viewed laterally) than second pereopod. Dactyl sub- equal in length to propodus. Dactyl and 1791). Scale equals 1 mm (a, c), 0.5 mm (b). (Stippled areas propodus with tufts of long plumose setae on ventral margin. Propodus with dorso- distal, corneous-tipped spine, and ventro- distal spine; outer face lacking longitudinal furrow; outer and ventral faces with tufts of simple setae; ventral margin with row of small corneous spines. Carpus with strong dorsodistal, corneous-tipped spine; ventral VOLUME 107, NUMBER 1 MIN nt A Fig. 7. 145 Right fourth pereopod (a-c), lateral view, anterior lobe of sternite of third pereopod (d-f), ventral view, and denuded telson (g-1), dorsal view. a, d, g, Calcinus urabaensis, new species; b, e, h, Calcinus verrilli (Rathbun, 1901); c, f, 1, Calcinus tibicen (Herbst, 1791). Scales equal 1 mm (a, ¢, i, g), 0.5 mm (b, d, f, h), and 0.25 mm (e). margin unarmed, with long setae. Merus with spine distally on ventrolateral margin; ventral margin unarmed, with row of tufts of long simple setae. Fourth pereopod subchelate (Fig. 7a). Dactyl with 13 small corneous spines ven- trolaterally. Propodus unarmed dorsally. Carpus with strong dorsodistal spine. Fifth pereopod chelate. Sternite of third pereopods (Fig. 7d) with anterior lobe broad, subrectangular, devel- oped as pair of subequal, rounded, setose projections divided by cleft, each projection with row of small tubercles distally. Telson (Fig. 7g) with posterior lobes asymmetrical, left larger than right; sepa- rated by narrow, distinct median cleft; ter- minal margins with 5 (left) or 3 (right) cor- neous-tipped, subterminal spines, and fringe of long bristles (not illustrated in Fig. 7g). Coloration in life (based on field notes). — Carapace red with tinge of purple. Ocular peduncles purple basally, fading distally, and with cream-colored band near base of cor- nea. Chelipeds purple, fingers white. Am- bulatory legs orange; dactyl cream-colored, except for dark red band proximally (Figs. 5a, b, 6a). Etymology. —The specific name is given for the Gulf of Uraba, where the only spec- imen of this species was collected. Distribution and habitat.—Known only 146 from the inlet of Pinorroa, Gulf of Uraba, Colombia; on rocks exposed to strong waves, in about 3 m of depth. Calcinus verrilli (Rathbun, 1901) Figs. 1b, 2b, 3b, 4b, 5c, 6b, 7b, e, f Clibanarius verrilli Rathbun, 1901:238.— Alcock, 1905:161.—Verrill, 1908:449, pl. 27, fig. 5, pl. 28, fig. 6.—Gordan, 1956: 310. Calcinus verrilli.—Provenzano, 1960:120, fig. 1.—Hazlett & Provenzano, 1965: 617.—Markham, 1977:131, pl. 1.—Chace et al., 1986:336, fig. 111, color pl. 10.2.— Morgan, 1991:910. Type material.— Holotype 6 (SL 7.8 mm), paratype 2? (SL 4.2 mm), Bermuda, coll. F. V. Hamlin, USNM 24818. Additional material. — Bermuda: 2 2 (SL 2.3—2.6 mm), coll. G. Brown Goode, 1876-— 1877, USNM 109433.—4 6 (SL 1.77-3.5 mm), Three Hill Shoal, 14 Nov 1958, coll. AJP, USNM 103734.—6 6(SL 3.2—5.2 mm), 2 2 (SL 3.2-4.1 mm), 3 2 ovigs (SL 3.1-4.2 mm), ~3 km north of North Rock Reef, on coral reef, 42 m, 5 Sep 1979, coll. G. Wener, USNM 265218.—2 2 (SL 4.4—4.8 mm), 1 6 (SL 4.3 mm), North Rocks Reef, 14 Nov 1958, coll. AJP, UMML 32:1643.—1 2 (SL 2.8 mm), 1 6 (SL 3.0 mm), 1 juv (SL 1.9 mm), Western Ledge Reef, 13 Nov 1958, coll. AJP, UMML 32:1644.—5 @ (SL 4.3- 4.7 mm), 3 6(SL 4.4—5.2 mm), North Rocks Reef, 14 Nov 1958, coll. AJP, UMML 32: 1645.—1 6 (SL 2.5 mm), Pt. at Shelly Bay, 8 Nov 1958, coll. AJP, UMML 32:1955.— 1 2 (SL 5.2 mm), 15 6 (SL 3.6—4.3 mm), North Rocks Reef, 14 Nov 1958, coll. AJP, UMML 32:2216.—3 2 (SL 1.7—2.3 mm), 3 3 (SL 1.9-2.0 mm), N.E. Breakers, Sep 1966, coll. Schone, UMML 32:5172. Diagnosis.—Ocular peduncles (Fig. 1b) varying in relative length with specimen size, from moderately stout, about *4 length of shield in smaller specimens (SL = 4.0 mm), to slender and subequal to length of shield PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON in larger specimens; acicles terminating in strong spine. Anterodorsal plate of bran- chiostegite (Fig. 2b) with blunt anteroven- tral angle. Left cheliped (Fig. 3b) with dactyl having upper margin and outer upper face with row of corneous-tipped spines; upper margin of palm with row of strong corne- ous-tipped spines, and outer upper face with several small spines on distal half; outer face of carpus with prominent tubercle on upper half. Fingers of right cheliped (Fig. 4b) leav- ing wide gap when closed; dactyl with strong, usually upwardly curved corneous-tipped spine proximally, and distal row of anteri- orly directed spines; palm with row of cor- neous-tipped spines on upper margin; car- pus with 2 or 3 corneous-tipped spines on dorsal margin. Ambulatory legs (Figs. Sc, 6b) with dactyl subequal in length to prop- odus and 5 to 7 long, slender corneous spines on ventral margin of dactyl; ventral margin of dactyl and propodus with several tufts of long simple setae. Third pereopod lacking longitudinal furrow on outer face of prop- odus. Fourth pereopod (Fig. 7b) with small dorsodistal spine on propodus. Anterior lobe of sternite of third pereopods (Fig. 6e) sub- rectangular, setose, often developed as pair of low, rounded projections, with or without small tubercles distally. Telson (Fig. 7h) with posterior lobes subequal, each with 7 spines on terminal margin, outermost spines on left lobe curved ventrally; terminal margin with fringe of long setae (not illustrated in Fig. 7h). Coloration in life (after Provenzano 1960: 120, and Chace et al. 1986: color pl. 10.2).— General color of body purple with red spots; eyestalks purple near base but with increas- ing red distally; area proximal to cornea white, cornea black or black spotted with white; chelipeds purple with red patches on upper distal face of carpus and merus; an irregular row of red pigment along upper margin of hand extends to dactyl; ambu- latory legs with similar rows of red pigment on faces of segments (see Figs. 5c, 6b). Distribution and habitat.—Endemic to VOLUME 107, NUMBER | Bermuda; subtidal to 110 m, in attached vermetid gastropod shells of Spiroglyphus irregularis and S. annulatus, or unattached gastropod shells (Markham 1977). Remarks. —Some specimens of this spe- cies (UMML material) exhibit sexual di- morphism in the armature of the chelipeds. The outer face of the right palm in some females is armed on the distal half with three or four small spines, whereas in some males the outer face is armed with only one distal spine. Males tend to have more spines on the right cheliped than females. The number and strength of the spines of the upper mar- gin of the left palm and dactyl varies ac- cording to sex of the individual, being fewer and stronger in large males (SL > 4 mm) than in females of similar size. Three male specimens were found to have both male and female gonopores. One male (SL 5.2 mm, UMML 32:1645) had female gonopores on the right and left side, and two males (SL 5.1 mm, 5.6 mm, UMML 32:2216) had only one female gonopore on either the left or right side. McLaughlin & Lemaitre (1993) have reported specimens with both male and female gonopores in the tube-dwelling hermit crab Paguritta kroppi McLaughlin & Lemaitre. As in P. kroppi, it is unclear whether the condition observed in the specimens of C. verrilli represents an aberration or a reproductive adaptation, such as protandry or protogyny, to a re- stricted mode of life. Calcinus tibicen (Herbst, 1791) nigse leon Scr4ce dy 6s 7 Gate Cancer tibicen Herbst, 1791:25, pl. 23, fig. 7. Pagurus sulcatus H. Milne Edwards, 1836: 279. Calcinus formosus Neumann, 1878:31.— Alcock, 1905:164.—Gordan, 1956:304.— Morgan, 1991:907. Calcinus sulcatus. — Benedict, 1901:141, pl. 5, figs. 3, 3a.—Alcock, 1905:164. Calcinus tibicen.—Provenzano, 1959:363, fig. 4.— Forest & De Saint Laurent, 1967: 147 106.—Sanchez & Campos, 1978:22, fig. 5.—Chace et al., 1986:335, fig. 111. Material examined. — Florida: 1 29 (SL 5.3 mm), Fort Pierce, Dynamite Point North, beach in rocky tide pool, 18 Mar 1972, coll. R. G. Gilmore, IRCZM 89:050.—1 6 (SL 4.2 mm), 1 2 (SL 3.3 mm), | 2 ovig (SL 4.4 mm), Vero Beach, 100 yards (91 m) off Sand Point, 10 ft (3 m), on coquina shelf, 28 Jun 1972, coll. RHG, IRCZM 89:219.—1 6 (SL 6.3 mm), St. Lucie County, Walton Rocks, in worm reef during night survey, intertidal, 27 Jan 1975, coll. RHG, IRCZM 89:2625.— 1 6 (SL 4.6 mm), Stuart, north of St. Lucie Inlet, in worm reef by “Fish Bow!” channel, intertidal, 12 Apr 1972, coll. RHG, IRCZM 89:101.—1 2 ovig (SL 5.2 mm), St. Lucie County, Walton Rocks, Hutchinson Island, across from Florida Power and Light Com- pany plant, intertidal, 9 Jul 1975, coll. J. Dudley, IRCZM 89:2302.—1 6(SL6.7 mm), Indian River, Martin County, near St. Lucie Inlet, on rocks, 4 May 1988, coll. R. S. Rox, IRCZM 89:06472.—1 2 (SL 3.3 mm), Tor- tugas, 21 Jun 1932, coll. WLS, USNM 102696.—146(SL4.5 mm), Tortugas, 19 Jul 1930, coll. WLS, USNM 102697.—2 6 (SL 3.0-6.0 mm), 1 2 (SL 4.9 mm), Tortugas, sta 34-32, 5 Jul 1932, coll. WLS, USNM 102698. Virgin Islands: 1 6(SL 3.3 mm), St. Croix, Christiansted, East Little Princess, intertid- al under rocks, 17 Aug 1971, coll. W. E. Rainey, USNM 154541. Jamaica: 1 6 (SL 4.3 mm), Port Antonio, coral reef off Navy Island, 1932, USNM 77398. Barbados: 2 6 (SL 5.8-6.3 mm), | 2 (SL 4.9 mm), Hasting’s Reef, opposite St. Mat- thias Church, 20 Jul 1959, coll. A. G. Humes & R. U. Gooding, USNM 104256. Antigua: 13 6 (SL 3.3-8.7 mm), 4 2 (SL 2.2-3.5 mm), 10 2 ovigs (SL 3.3—5.3 mm), Smithsonian-Bredin Caribbean Expedition 1956, sta 73-56, Charlotte Point (=Nut- ting’s ““Rocky Point’), English Harbor, 2 Apr 1956, USNM 265157. 148 Belize: 1 6 (SL 4.8 mm), | 2 ovig (SL 5.5 mm), 15 Jul 1930, coll. P. W. Shufeldt, USNM 102700.—1 6 (SL 6.4 mm), Carrie Bow Cay, lagoon, 2-3 m, 25 Apr 1975, USNM 184516. Isla Providencia (Colombia): 1 ¢ (SL 6.3 mm), Bahia Maracaibo, sandy-rocky beach, 28 Mar 1991, coll. G. E. Ramos, USNM 259399. Panama: 2 6 (SL 3.3-6.7 mm), Colon, Coral reef, 2 May 1911, coll. Meek & Hil- debrand, USNM 44191. Curacao: 1 6 (SL 4.0 mm), 1 2 (SL 4.8 mm), Caracas Bay, 1920, coll. C. J. van der Horst, USNM 57513. Diagnosis.—Ocular peduncles (Fig. 1c) slightly shorter than length of shield; acicles terminating in strong spine (occasionally bi- fid or trifid) distally. Anterodorsal plate of branchiostegite (Fig. 2c) with blunt antero- ventral angle. Antennular peduncle (Fig. 2c) with lateral face of basal segment armed with 4 to 6 small spines on distal margin. Che- lipeds lacking setae, surfaces minutely gran- ulose. Left cheliped (Fig. 3c) with chela un- armed; outer face of carpus with prominent subrectangular lobe proximally on upper half. Right cheliped (Fig. 4c) unarmed ex- pect for small tubercle proximally on outer face of carpus; fingers not leaving gap when closed. Ambulatory legs (Figs. 5d, 6c) each with dactyl shorter than propodus (about 0.6 times length of propodi); ventral margin of dactyl with row of 4 to 7 short corneous spines, and several tufts of short simple se- tae; lateral face of propodus of third per- eopods each with distinct longitudinal fur- row on upper half of lateral face. Fourth pereopod (Fig. 7c) with dorsal margin of propodus unarmed. Anterior lobe of ster- nite of third pereopods (Fig. 7f) subrectan- gular, setose, often developed as pair of low, rounded, unarmed projections. Telson (Fig. 71) with posterior lobes asymmetrical, each usually with 1 submarginal spine; terminal margin with fringe of long setae (not illus- trated in Fig. 71). Coloration in life (after Provenzano 1959: PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 363, and Chace et al. 1986:335).—Che- lipeds red-brown to maroon, often tinged with purple, and with white fingertips. Dac- tyls of the ambulatory legs white or yellow, with a red band (Figs. 5d, 6c). Eyestalks orange-red fading to white near cornea. Car- apace usually red, often tinged with purple (rarely green), and with white punctae. Distribution and habitat. — Florida, Gulf of Mexico, Caribbean to Brazil, including Bermuda; lower intertidal and subtidal to 18 fms (32.9 m); commonly encountered, in hard substrates such as rocky bottoms, worm reef platforms, and coral reefs. Remarks. —Several carcinologists have indicated that Calcinus tibicen is broadly distributed in the West Indian region from Florida to Brazil, including Bermuda [e.g., Alcock 1905 (as C. sulcatus), Verrill 1908 (as C. sulcatus) Schmitt 1935, Provenzano 1959, Abele & Kim 1986, Morgan 1991]. Morphological Comparison of Calcinus Species from the Western Atlantic Calcinus urabaensis is similar to C. ver- rilli but differs from it in a number of at- tributes, and specially coloration. In the ab- sence of coloration, the most reliable difference between Calcinus urabaensis and C. verrilli can be found on the chelipeds, the dactyls of the third pereopods, and the tel- son. The dactyl of the left cheliped in C. urabaensis has a row of small spines on the upper margin, and irregular rows of low, closely-set tubercles on the outer face; the dactyl in C. verrilli has two rows of corne- Ous-tipped spines, one row on the upper margin and another on the upper outer face. The left palm in C. urabaensis is armed on the upper margin with small, low protuber- ances, whereas this margin is armed with a row of strong, corneous-tipped spines in C. verrilli. On the right cheliped of C. ura- baensis the dactyl is armed with a row of spines directed anteriorly, whereas in C. verrilli the dactyl has a strong, upwardly VOLUME 107, NUMBER 1 curved spine proximally, followed distally by smaller anteriorly directed spines. The upper margin of the right palm in C. ura- baensis is crest-like and armed with a row of five strong spines; the upper margin of the palm in C. verrilli also has spines but is not raised in the form of a crest. Calcinus urabaensis is unique among the western Atlantic species of Calcinus in that dactyls of the third pereopod have several tufts of long plumose setae on the ventral margin. A similar condition is found in oth- er Calcinus species, all distributed in the Indo-Pacific region: C. vachoni Forest, C. sirius Morgan, C. spicatus Forest, and C. latens (Randall) (see Morgan 1991). The telson in C. urabaensis has the pos- terior lobes distinctly asymmetrical, left the largest, and the terminal margins of the lobes are armed with 3—5 subterminal spines. In contrast, the telson in C. verrilli has the pos- terior lobes weakly asymmetrical, and the terminal margins of the lobes are armed with 7 terminal spines. Calcinus tibicen can immediately be sep- arated from the other two western Atlantic species of the genus by the smooth, un- armed chelipeds, the longitudinal furrow on the lateral face of the propodus of the third left pereopod, the stout dactyls of the am- bulatory legs, and the distinctive color pat- tern that often persists for many years in alcohol-preserved specimens. Although the coloration of C. tibicen generally resembles that of C. urabaensis, the two species oth- erwise differ markedly. The two can readily be differentiated by the armature of the che- lipeds (armed with spines in C. urabaensis, unarmed in C. tibicen), and the setation of the dactyl of the third pereopod (plumose in C. urabaensis, simple in C. tibicen). Acknowledgments We thank the Instituto de Investigaciones Marinas de Punta de Betin, Santa Marta, for funding the expedition to the Gulf of Uraba, and its staff for their cooperation. 149 This study was made possible, in part, by grants received by one of us (NHC) from COLCIENCIAS and the Smithsonian In- stitution’s Office of Fellowships and Grants Short-Term Visitor program, which per- mitted travel to Washington, D.C. Thanks are given to Paula Mikkelsen for facilitating work with collections at the IRCZM. This study is also supported by the Smithsonian Marine Station at Link Port, Fort Pierce, Florida, and is contribution number 346 from that station. Literature Cited Abele, L. G., & W. Kim. 1986. An illustrated guide to the decapod crustaceans of Florida. Parts 1 & 2. Florida State University, Tallahassee, 760 pp. Alcock, A. 1905. Catalogue of the Indian decapod Crustacea in the collection of the Indian Mu- seum. Part II. Anomura. Fascicle I. Pagurides. Indian Museum, Calcutta, 197 pp., pls. 1-15. Benedict, J.E. 1901. The anomuran collections made by the Fish Hawk Expedition to Porto Rico.— United States Fish Commission Bulletin for 1900, 2(2):129-148, pls. 3-6. Campos, N. H., & G. Manjarrés. 1988. Decapodos Brachyura de la region del Golfo de Uraba (Ca- tribe Colombiano).—Anales del Instituto de In- vestigaciones Marinas, Punta Betin 18:17—23. Chace, F. A., Jr., J. J. McDermott, P. A. McLaughlin, & R. B. Manning. 1986. Order Decapoda (Shrimps, lobsters and crabs). Pp. 312-358 in W. Sterrer, ed., Marine fauna and flora of Ber- muda. John Wiley & Sons, Inc., New York, 742 pp. Dana, J. 1852. Crustacea, Part 1, in United States Exploring Expedition during the years 1838, 1839, 1840, 1841, 1842, under the command of Charles Wilkes, U.S.N. 13(1):1-685. Atlas (1855):1—27, pls. 1-96. Philadelphia. Forest, J., & M. de Saint Laurent. 1967. Résultats scientifiques des campagnes de la “Calypso”’, fascicule 8, Campagne au large des cétes Atlan- tiques de l’Amérique du sud (1961-1962). I. No. 6. Crustacés Décapodes: Pagurides.— Annales de I’ Institut Océanographique, Monaco, new se- ries 45(2):47-172, pl. 1. Gordan, J. 1956. A bibliography of pagurid crabs, exclusive of Alcock, 1905.—Bulletin of the American Museum of Natural History 108(3): 257-352. Haig, J.. & P. A. McLaughlin. 1984. New Calcinus species (Decapoda: Anomura: Diogenidae) from 150 Hawaii, with a key to the local species. — Mi- cronesica 19 (1—2):107-121. [1983]. Hazlett, B. A., & A. J. Provenzano, Jr. 1965. De- velopment of behavior in laboratory reared her- mit crabs.—Bulletin of Marine Science 15(3): 616-633. Herbst, J. F. W. 1791-1796. Versuch einer Natur- geschichte der Krabben und Krebse, nebst einer systematischen Beschreibung ihrer verschiede- nen Arten, 2:i—vili, 1-226, pls. 22-46. McLaughlin, P. A., & R. Lemaitre. 1993. A review of the hermit crab genus Paguritta (Decapoda: Anomura: Paguridae) with description of three new species. — Raffles Bulletin of Zoology 41(1): 1-29. Markham, J.C. 1977. Preliminary note on the ecol- ogy of Calcinus verrilli an endemic Bermuda hermit crab occupying attached vermetid shells.—Journal of Zoology 181:131-136. Milne Edwards, H. 1836. Observations zoologiques sur les Pagures et description d’un nouveau gen- re de la tribu des Paguriens.— Annales des Sci- ences naturelles, Paris, Zoologie (2)6:257-288, pls. 13, 14. Morgan, G. J. 1991. A review of the hermit crab genus Calcinus Dana (Crustacea: Decapoda: Di- ogenidae) from Australia, with descriptions of two new species.—Invertebrate Taxonomy 5:869-9 13. Neumann, R. 1878. Systematische Uebersicht der Gattungen der Oxyrhynchen. Catalog der po- dophthalmen Crustaceen des Heidelberger Mu- seums. Beschreibung einiger neuer Arten, Druck Von J. B. Hirschfeld, Leipzig, 39 pp. Provenzano, A. J., Jr. 1959. The shallow-water her- mit crabs of Florida.— Bulletin of Marine Sci- ence of the Gulf and Caribbean 9(4):349—420. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON . 1960. Notes on Bermuda hermit crabs (Crus- tacea; Anomura).— Bulletin of Marine Science of the Gulf and Caribbean 10(1):117-124. Rathbun, M. J. 1901. Untitled footnote, in A. E. Verrill, ed., Recent papers relating to the fauna of the Bermudas, with some corrections.— American Journal of Science Series (4)11(64): 326-330. Sanchez, H., & N. H. Campos. 1978. Los cangrejos ermitanos (Crustacea, Anomura, Paguridae) de la costa norte colombiana. Parte I.—Anales del Instituto de Investigaciones Marinas, Punta de Betin 10:438-453. Schmitt, W. L. 1935. Crustacea Macrura and Ano- mura of Porto Rico and the Virgin Islands. Sci- entific survey of Porto Rico and the Virgin Is- lands.—New York Academy of Sciences 15(2): 125-262. Verrill, A. E. 1908. Decapod Crustacea of Bermuda. I. Brachyura and Anomura. Their distribution, variations, and habits.—Transactions of the Connecticut Academy of Arts & Sciences 13(6): 299-474, pls. 9-28. Wooster, D.S. 1984. The genus Calcinus (Paguridea, Diogenidae) from the Mariana Islands including three new species.—Micronesica 18(2):121—162. [1982]. (NHC) Instituto de Ciencias Naturales, Universidad Nacional de Colombia, % IN- VEMAR, A. A. 1016, Santa Marta, Colom- bia; (RL) Department of Invertebrate Zo- ology, National Museum of Natural His- tory, Smithsonian Institution, Washington, D.C. 20560, U.S.A. PROC. BIOL. SOC. WASH. 107(1), 1994, pp. 151-163 PSEUDOCYCLOPS LAKSHMI, A NEW SPECIES (PSEUDOCY CLOPIDAE: CALANOIDA: COPEPODA) FROM THE LACCADIVES, INDIA P. Haridas, M. Madhupratap, and S. Ohtsuka Abstract. —Pseudocyclops lakshmi, new species, is described from the Kadmat and Agatti atolls of the Laccadives, India, along with its dimorphic males. Characters separating the new species from the rest of its congeners are dis- cussed. This species is epibenthic and emerges in large numbers into the water column at night. The hyperbenthic calanoid copepod ge- nus Pseudocyclops Giesbrecht, 1893 at pres- ent consists of 29 species described from shallow waters of the Atlantic and Indo- Pacific regions (Brady 1872; Brady & Rob- ertson 1873; Giesbrecht 1893; Sars 1903, 1919; Esterly 1911; Gurney 1927; Sewell 1932; Nicholls 1944a, 1944b; Noodt 1958; Bowman & Gonzalez 1961; Vervoort 1964; Tanaka 1966; Wells 1967; Fosshagen 1968; Por 1968; Dawson 1977; Andronov 1986; Othman & Greenwood 1989; Barr & Ohtsu- ka 1989). Nine species of the genus have hitherto been recorded/described from the Indian Ocean including the Red Sea and Suez Canal, viz. P. obtusatus Brady & Rob- ertson, 1873 from the Arabian Sea and Sri Lanka (Thompson & Scott 1903); P. um- bricatus Giesbrecht, 1893 and P. /atens Gurney, 1927 from the Suez Canal; P. /a- tisetosus Sewell, 1932, and P. simplex Sew- ell, 1932 (type localities of these two species are not given in Sewell 1932); P. reductus Nicholls, 1944b, P. gohari Noodt, 1958 and P. steinitzi Por, 1968 from the Red Sea, and P. xiphophorus Wells, 1967 from Mozam- bique. Their occurrences so far are known only from type localities except P. obtusatus which is recorded broadly from the North Atlantic Ocean, the Mediterranean Sea and the Indian Ocean (see Thompson & Scott 1903, Rose 1933, Vervoort 1964, Othman & Greenwood 1989). Pseudocyclops obtu- satus from the Indian Ocean was reported but not illustrated by Thompson & Scott (1903); there remains the possibility that not all of its records are correct. Similarly, P. australis Nicholls, 1944a, originally de- scribed from South Australia has been re- corded from South Japan (Tanaka 1966) al- beit with differences in the structure of exopods of antenna and mandibular palp. During investigations on zooplankton of the atolls of the Laccadive Sea (Madhupra- tap et al. 1991a, 1991b), an undescribed species of Pseudocyclops was found to be a dominant calanoid copepod in the night- time net tows in the shallow lagoons of Kad- mat and Agatti atolls. Although Pseudocy- clops species are usually distributed near or on the seabottom, they are often found in the water column due to their vertical mi- gration (e.g., Esterly 1911, Gurney 1927, Vervoort 1964, Fosshagen 1968, Dawson 1977, Othman & Greenwood 1989, Oht- suka unpublished data). The present paper describes both sexes of the new species of Pseudocyclops including the apparently di- morphic males. Pseudocyclops lakshmi, new species Figs. 1-7 Material.—Specimens were collected in surface hauls at night using a square net (mesh width 200 um) from Agatti on 10- 152 WLW EEE S O-I1mm PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Pre er c 1 ery reer TNE Pony yet ” : Pep, ae rv Pre renee nny rv npe veneTTERE RITE wren ey yore Yer HUNK WG i LUNI VOLUME 107, NUMBER 1 153 am Sater ¢ 2 : e 5 Fig.2. Pseudocyclops lakshmi, Female: SEM photomicrographs. A. Urosome dorsal view: B. Urosome ventral view; C. Distal margin of genital double-somite dorsal view; D. Genital double-somite ventral view. All scale bars = 10 wm. — Fig. 1. dorsal; E. Habitus lateral; F. Urosome. Pseudocyclops lakshmi, Female: A. Habitus, dorsal; B. Habitus lateral; C. Urosome. Male: D. Habitus, 154 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 3. 11 Dec 1988 and from Kadmat between 1 4— 16 Dec 1988. The morphological terminol- ogy follows Huys & Boxshall (1991). The urosomes of females of the new species were examined with a Scanning Electron Micro- scope (SEM-JEOL JSM T-20). All types are deposited at the Indian Ocean Biological Centre (IOBC), Cochin. Types. — Holotype: Female, total length 0.90 mm, IOBC 0405-08-46-1992. Allo- type: Male, morph “A”—IOBC 0406-08- 46-1992. Paratypes: 10 females & 11 males (6 morph “A” and 5 morph “B’’), IOBC 0407-08-46-1992 plus two females and four males dissected and mounted on glass slides IOBC-0408-08-46-1992. Description. —Female. Body (Fig. 1A, B) 0.86—0.95 mm in total length (average 0.90 mm, 7 = 13). Prosome oval in dorsal view, cephalosome separate from first pedigerous somite, fourth and fifth pedigerous somites completely separate, posterior corner of fifth pedigerous somite smoothly rounded, reaching to one-third length of genital dou- ble-somite. Rostrum pointed, triangular, Pseudocyclops lakshmi, Female: A. Antennule. Male: B. Right antennule. with a pair of minute sensilla. Urosome (Fig. 1C) 4-segmented, distal margins of first 2 somites lamellar. Genital double-somite largest, as long as following 2 somites com- bined and covered with numerous minute prominences (Fig. 2A, C). Gonopores and copulatory pores paired, closed off by oper- culum-like leg 6 (Fig. 2B, D). Second and third urosomal somites with subterminal transverse row of minute spinules; third urosomal somite produced posterodorsally into 2 triangular processes reaching beyond midlength of caudal ramus (Fig. 2A); anal somite small and telescoped into third uro- some somite. Caudal ramus with serrate posterior margin dorsomedially and 1 bluntly pointed process ventromedially and with 1 dorsal, 4 terminal and 1 outer sub- terminal setae. Antennule (Fig. 3A) 21-segmented, not quite reaching to posterior end of cephalo- some; first segment with 3 large aesthetascs and 11 setae; fourth and fifth and eighteenth and nineteenth segments partly fused; ter- minal segment with one aesthetasc. Anten- VOLUME 107, NUMBER | 155 0:05mm —EEE—EE Fig. 4. Pseudocyclops lakshmi, Female: A. Antenna; B. Mandible; C. Maxillule; D. Maxilliped. na (Fig. 4A) basis with seta on inner distal corner; endopod 3-segmented, first segment with 2 setae at midlength of inner margin and subterminal patch of minute spinules, second segment with 5 inner marginal and 4 terminal setae, third segment bearing 7 setae terminally and 4 rows of spinules; ex- opod 7-segmented, third to fifth segments incompletely fused, first 6 segments each having | seta, seventh segment with 1 me- dial and 4 terminal setae. Mandible (Fig. 4B) gnathobase with 8 blunt teeth; basis with 2 inner setae and patch of minute promi- nences subterminally; endopod 2-segment- ed, first segment small, having 4 inner ter- minal setae, second segment tapering proximally, with 10 terminal setae; exopod 4-segmented, first 3 segments each bearing 156 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 0-05mm 0:05mm Fig. 5. Pseudocyclops lakshmi, Male: A. Morph B maxilla; B. Morph A maxilla. inner seta, terminal segments carrying 3 se- tae. Maxillule (Fig. 4C) well developed; praecoxal arthrite with 5 setae on posterior surface and 1 weak and 8 stout spine-like setae along inner margin; coxal and 2 basal endites having 3, 4 and 5 setae respectively; coxal epipodite furnished with 9 setae of unequal lengths; basal exite having short VOLUME 107, NUMBER | single seta; endopod 2-segmented, first seg- ment bearing 5 middle and 5 terminal setae along inner margin, second segment with 6 terminal setae; exopod 1-segmented, with 11 setae. Praecoxal and coxal endites of maxilla (Fig. 5B, female maxilla similar to morph A of male) having 6, 3, 3 and 3 setae respectively; basis completely fused with first endopod segment to form allobasis, fur- nished with 7 setae; endopod 3-segmented, first to third segments having 2, 2 and 3 setae respectively. Maxilliped (Fig. 4D) syn- coxal endites with 1, 2, 3 and 3 setae re- spectively; allobasis with 3 medial and 2 terminal setae along inner margin; second to fifth endopod segments each with 4 setae. Legs 1-4 (Fig. 6A—-D) each with 3-seg- mented rami, all bearing numerous minute spinules on both anterior and posterior sur- faces. Exopod segments 1| and 2 with | seta, exopod segment 3 with 5 setae except in leg 1 (4 setae), endopod segments 1 and 2 with 1 and 2 setae respectively, endopod segment 3 with 8 setae except in legs | (6 setae) and 4 (7 setae). Outer distal corners of second endopod segments of legs 2-4 each pro- duced into bifid process; distal corners of third endopod segments of legs 14 acutely produced. Second exopod segment of leg 1 with bulbous process on outer distal comer; terminal spines of third exopod segments of legs 1-4 elongate but not flanged. Leg 5 (Fig. 6E). Basis with outer subter- minal seta on posterior surface; endopod 3-segmented, first segment having a small outer spinule, first and second segment with inner terminal seta, third segment having 1 inner, | outer 2 terminal setae; exopod 3-segmented, third segment with 4 flanged spines and 4 inner setae. Male: Dimorphism was observed only in males. The differences between the two morphs are found in body length and max- illa. In particular, the two morphs differ in the size of maxilla (see Fig. 5A, B). In this paper, males with small and large maxillae are referred to as morphs A and B respec- tively. Body length of morph A 0.81-0.85 157 mm (average 0.83 mm, 7 = 10) and that of morph B 0.85-0.95 mm (average 0.90 mm, n = 10), prosome of both morphs similar to female (Fig. 1D, E). Urosome (Fig. 1F) 5-segmented, covered with minute prominences as in the female; first 3 somites with finely striated posterior margin as in female; 2 triangular dorsal pro- cesses of fourth somite reaching to distal end of caudal ramus; anal somite telescoped into fourth somite. Caudal ramus with ser- rate dorsomedial margin and ventromedial blunt process. Right antennule (Fig. 3B) 18-segmented, geniculate between 14th and 15th segments; first segment with 3 large aesthetascs and 3 rows of minute spinules; seventh and eighth segments fused or separate; 14th segment with sinuous process along whole length of anterior margin; 16th segment produced distally into triangular process reaching midlength of 17th segment; terminal seg- ment with one aesthetasc. Left antennule 21-segmented; fourth and fifth, and eigh- teenth and nineteenth segments incom- pletely fused as in female antennule. Antenna, mandible, maxillule, maxilli- ped and legs 1-4 of both morphs similar to those of the female. Maxilla of morph B (Fig. 5A) about twice as long as that of morph A (Fig. 5B); 2 ser- rate setae on basis (indicated by small ar- rows) much stouter in morph B than in morph A; seta on second endopod segment (indicated by large arrow) longer and stouter in morph B than in morph A. Legs 5 of both morphs (Fig. 7A—I) similar to each other except being slightly smaller in morph A. Right leg: coxa and intercoxal sclerite fused; basis separate from coxa, with 1 seta on posterior surface; endopod (Fig. 7A, D, E, F) 1-segmented with small ter- minal spinules and minute surface spinules, usually without terminal seta; exopod 2-segmented, first segment with stout, flanged terminal spine, second segment with 3 processes, outer long, thick, and medially curved, middle small at base of outer pro- Yc Le = VOLUME 107, NUMBER 1 159 0:05mm Fig. 7. Pseudocyclops lakshmi, Male leg 5: A. Posterior view (left exopod segment 2 omitted); B. Left exopod segment 2; C. Left endopod; D. Anterior view; E-F. Right endopod variability; G-I. Pointed process on basis of left leg showing variability in shape. 160 cess, inner slender and recurved appearing as chela. Left leg: coxa incompletely fused with basis on both surfaces; basis covered with minute surface prominences, with elongate, pointed process originating from anterior surface (Fig. 7A, D, G, H, I) and 1 seta on posterior surface; endopod (Fig. 7A, C) l-segmented, with numerous surface prominences and 5 plumose terminal setae; exopod 2-segmented; first segment with outer terminal flanged spine; second seg- ment (Fig. 7B) spoon-shaped, complex and membranous, proximal lobe oblong with 2 hemispherical hyaline processes originating from medial margin and 2 terminal pro- cesses, 2 outer processes attached on to out- er margin; proximal process rod-shaped with bulbous projection and short seta at base and distal process more or less napiform with subterminal cleft. Variation. —Segment fusion pattern is variable in right antennule of morph A male—seventh and eighth segments com- pletely or incompletely fused. Right endo- pod of leg 5 of morph B male has none or 1 terminal seta. The process originating from left basis of leg 5 of morph A male is vari- able in shape, smoothly tapering distally or having a knob medially (Fig. 7G—I). Etymology. —This new species is named after goddess ““Lakshmi”’ (Hindu mytholo- gy) who symbolizes abundance and wealth. As mentioned earlier, this species is the most abundant calanoid copepod in the two la- goons. Discussion. —The new species lakshmi is one of the most primitive species of the ge- nus Pseudocyclops, having 5 separate pe- digerous somites, 21-segmented female an- tennule and 7-segmented antenna exopod; it has an outer basal spine on leg 3, 3-segmented endopod in leg 5 of female and 5 plumose setae on the left endopod of male leg 5. Such primitive characters as 5 sepa- rate pedigerous somites, 3-segmented en- dopod of female leg 5 and 5 setae on left endopod of male leg 5 are found also in P. australis, P. gohari, P. kulai Othman & Greenwood, 1989, P. lerneri Fosshagen, PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 1968, P. mathewsoni Fosshagen, 1968, P. reductus, P. rubrocinctus Bowman & Gon- zalez, 1961 and P. Steinitze although their antennules and antennae are more ad- vanced. The present new species, however, has several advanced morphological characters present on the urosome, leg 1 and female leg 5. The urosome is covered with minute prominences, the second exopod segment of leg 1 has an outer bulbous process, and there are 4 setae on the third endopod seg- ment of the female leg 5 (6 setae in P. kulai, P. lepidotus Barr & Ohtsuka, 1989, P. ma- thewsoni, P. rubrocinctus, and P. steinitzi). The urosome with numerous minute prom- inences appears to be unique to the new species although P. /epidotus has foliaceous scales on the urosome. The bulbous process on the second exopod segment of leg | is known in P. australis and P. gohari; some congeners have a well developed process distal to the outer spine on the second ex- opod segment of leg 1 different in shape from that of P. lakshmi. Pseudocyclops lakshmi seems to be most closely related to P. australis from South Australia and Japan and P. gohari from the Red Sea in having 3-segmented endopod of the female leg 5 with 4 setae on the terminal segment and a bulbous process on the second exopod seg- ment of leg 1. In particular, the structures of leg 5 of both sexes of P. Jakshmi resemble those of P. gohari. The leg 5 of female P. reductus from the Red Sea also shows sim- ilarity to that of female P. /Jakshmi, but Ni- choll’s (1944b) description is too incom- plete to compare the two in detail. Pseudocyclops lakshmi is distinguishable from P. australis, P. gohari, and P. reductus in metasome, urosome, and legs | and 5 in addition to antennule and antenna. Cepha- losome and the first pedigerous somite are separate in P. lakshmi and P. australis, but fused in P. gohari. Male P. gohari has 8, 8 and 6 well-developed posterodorsal pro- cesses on the second, third and fourth uro- somal somites respectively; the terminal se- tae on the caudal ramus are extremely swollen. VOLUME 107, NUMBER 1 The numbers of the antennular segments of the females of P. lakshmi, P. australis and P. gohari are 21, 18 and 18 respectively. The antenna exopod is 7-segmented in P. lakshmi while 4-segmented in P. australis (shown as 5-segmented by Tanaka 1966) and P. go- hari. The mandibular exopod is 2-segmented (Nicholls 1944a) or 5-segmented (Tanaka 1966) in P. australis and 3-segmented in P. gohari while it is 4-segmented in P. lakshmi. The inner distal corner of the basis of leg 1 is prominent in P. australis but is not pro- duced in P. lakshmi or P. gohari. The outer distal corners of the first and second en- dopod segments of the female leg 5 are strongly produced in P. australis, P. gohari and P. reductus, but are rather rounded in P. lakshmi; there are 4 setae along the inner margin of the third exopod segment of the female leg 5 except for P. reductus which has only | seta. The right endopod of the male leg 5 has 5 terminal prominences in P. lakshmi and 6 to 7 in P. gohari, in ad- dition the 2 hemispherical hyaline processes originating along the second exopod medial margin of the left leg 5 in P. /akshmi are absent in P. gohari. The structure of the male leg 5 of P. australis is quite different from that of P. lakshmi. Do et al. (1984) reviewed dimorphism in copepod males. According to them, dimor- phism of copepod males is so far known in Calanoida, Cyclopoida, Harpacticoida and Poecilostomatoida. In Calanoida dimor- phism in male is reported in families Pon- tellidae and Pseudodiaptomidae (Johnson 1964, Fleminger 1967, Shen & Mizuno 1984, Walter 1989) and differences occur in characters such as body size and morphol- ogy of antennule and leg 5. Dimorphic asymmetry is found in the calanoid genus Pleuromamma mostly in the reproductive system (Ferrari 1984). Males of the calanoid copepod Euchaeta antarctica produce two types of spermatophores and the same male may produce both kinds of spermatophores (Ferrari & Dojiri 1987). Variations in size ranges of both females and males without morphological differences have been ob- 161 served in the poecilostomatoid copepod Oncaea spp. (Ferrari 1975). The pelagic har- pacticoid Euterpina acutifrons has two forms of males, namely large and small forms, which have differences in antennule, anten- na, developmental rate, sexual behavior, etc. (Haq 1965, 1972). The two forms have varying metabolic rates at different temper- atures (Moreira & Vernberg 1968). Varia- tions in the two forms of males of the poe- cilostomatoid Pseudomyicola spinosus parasitic on the blue mussel Mytilus edulis galloprovincialis are discernible in body length and slenderness, antennule, swim- ming legs and caudal ramus. Do etal. (1984) suggested that these differences are indica- tors of different swimming activity. Simi- larly, the parasitic copepod Pachypygus gib- ber has a smaller atypical male which is an active swimmer unlike the typical larger male and the former is more efficient re- productively (Hipeau-Jacquotte 1978). Although there remains a possibility that the two morphs of male Pseudocyclops lakshmi may be two distinct species, the absence of females with large-sized maxilla and the co-occurrence of both morphs of males led us to the conclusion that these morphs belong to the same species. Since species of the family Pseudocylopidae usu- ally have well-developed sexual characters in urosome, antennule and leg 5, the invar- iable sexual characters in both morphs of male P. lakshmi support this. It is possible that the feeding behaviors of these two morphs of male P. Jakshmi may be different because maxilla plays an important role in feeding. Acknowledgments We wish to thank three anonymous re- viewers for helpful suggestions. Literature Cited Andronoy, V. N. 1986. Bottom Copepoda in the area of Cape Blanc (Islamic Republic of Mauritania). 3. The family Pseudocyclopidae. — Zoologiche- ski Zhurnal 65:456—462. 162 Barr, D. J., & S. Ohtsuka. 1989. Pseudocyclops lep- idotus, a new species of demersal copepod (Cal- anoida: Pseudocyclopidae) from the northwest- ern Pacific.—Proceedings of the Biological Society of Washington 102:331-338. Bowman, T. E., & J. G. Gonzalez. 1961. Four new species of Pseudocyclops (Copepoda: Calanoida) from Puerto Rico.—Proceedings of the United States Museum 113:37-59. Brady, G. S. 1872. Contribution to the study of En- tamostraca, VII. A list of non-parasitic Cope- poda of the northeast coast of England.—Nat- ural History Transactions of Northumberland and Durham 4:432-445. , & D. Robertson. 1873. Contributions to the study of Entomostraca. 8. On marine Copepoda taken in the west of Ireland.—The Annals and Magazine of Natural History 12:126-142. Dawson, J. K. 1977. A new species of Pseudocyclops (Copepoda: Calanoida) from the southern Cal- ifornia coast.—Transactions of the American Microscopical Society 96:247-253. Do, T. T., T. Kajihara, & J. S. Ho. 1984. The life history of Pseudomyicola spinosus (Raffaele & Monticelli, 1885) from the blue mussel, Mytilus edulis galloprovincialis in Tokyo Bay, Japan, with notes on the production of atypical male. — Bul- letin of Ocean Research Institute 17:1-65. Esterly, C. O. 1911. Calanoid Copepoda from the Bermuda Islands.— Proceedings of the Ameri- can Academy of Arts and Science 47:219-226. Ferrari, F. D. 1975. Taxonomic status of the genus Oncaea (Copepoda: Cyclopoida) from the Gulf of Mexico and Northern Caribbean Sea.—Pro- ceedings of the Biological Society of Washington 88:217-232. 1984. Pleiotropy and Pleuromamma, the looking glass copepods (Calanoida).—Crusta- ceana Supplement 7:166-181. —, & M. Dojiri. 1987. The calanoid copepod Euchaeta antarctica from Southern Ocean At- lantic sector midwater trawls, with observations on spermatophore dimorphism.—Journal of Crustacean Biology 73:458—480. Fleminger, A. 1967. Taxonomy, distribution, and polymorphism in the Labidocera jollae group with remarks on evolution within the group (Co- pepoda: Calanoida).— Proceedings of the Unit- ed States National Museum 120:1-61. Fosshagen, A. 1968. Marine biological investigations in the Bahamas. 4. Pseudocyclopidae (Cope- poda, Calanoida) from the Bahamas. —Sarsia 32: 39-62. Giesbrecht, W. 1893. Mitteilungen tiiber Copepoden 1-6.— Mitteilungen aus der zoologischen Sta- tion zu Neapel, Berlin 11:56-104. Gurney, R. 1927. Zoological results of the Cambridge PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Expedition to the Suez Canal, 1924. Report on the Crustacea: Copepoda (littoral and semi-par- asitic).— Transactions of Zoological Society of London 22:451-577. Haq, S. M. 1965. Development of the copepod Eu- terpina acutifrons with special reference to di- morphism in the male.—Proceedings of Zoo- logical Society of London 144:175-201. 1972. Breeding of Euterpina acutifrons, a harpacticoid copepod, with special reference to dimorphic males.— Marine Biology 15:221—235. Hipeau-Jacquotte,R. 1978. Existence de deux formes sexuelles males chez le Copépode ascidicole No- todelphyidae Pachypygus gibber (Thorell, 1859).—Competes Rendus Hebdomadaires des Seances de |’Academie des Sciences 287D:253- 256. Huys, R., & G. A. Boxshall. 1991. Copepod evolu- tion. The Ray Society, London, 468 pp. Johnson, M. W. 1964. Ona new species of Pseudo- diaptomus from the west coast of Mexico, Costa Rica and Ecuador. —Crustaceana 7:33-41. Madhupratap, M., C. T. Achuthankutty, & S. R. S. Nair. 199la. Zooplankton of the lagoons of the Laccadives: diel patterns and emergence. — Journal of Plankton Research 13:947-958. , & 1991b. Estimates of high absolute densities and emergence rates of de- mersal zooplankton from the Agatti atoll, Lac- cadives.—Limnology and Oceanography 36: 585-588. Nicholls, A.G. 1944a. Littoral Copepoda from South Australia. II. Calanoida, Cyclopoida, Notodel- phyoida, Monstrilloida and Caligoida.—Rec- ords of South Australia Museum 8:1-62. . 1944b. Littoral Copepoda from the Red Sea. — The Annals and Magazine of Natural History 11:487-503. Noodt, W. 1958. Pseudocyclops goharin. sp. aus dem Eulitoral des Roten Meers (Copepoda Calan- oida).— Zoologischer Anzeiger 161:150-157. Othman, B. H. R., & J. G. Greenwood. 1989. Two new species of copepods from the family Pseu- docyclopidae (Copepoda, Calanoida).—Crus- taceana 56:63-77. Por, F. D. 1968. Copepods of some land-locked ba- sins in the islands of Entedebir and Nocra (Dah- lak Archipelago, Red Sea).—Bulletin of Sea Fisheries Research Station, Israel 49:32-50. Rose, M. 1933. Copépodes pélagiques.—Fauna de France 26:1—374. Sars, G. O. 1903. Copepoda. Calanoida. An account of the Crustacea of Norway, 4. Bergen Museum, Bergen, 171 pp. . 1919. Copepoda. Supplement. An account of the Crustacea of Norway. 7. Bergen Museum, Bergen, 115 pp. VOLUME 107, NUMBER 1 Sewell, R. B. S. 1932. The Copepoda of the Indian Seas. Calanoida.— Memoirs of the Indian Mu- seum 10:223-407. Shen, C., & T. Mizuno. 1984. Freshwater Copepoda from China and Japan. Tatarashobou, Yonago, 650 pp. (in Japanese). Tanaka, O. 1966. Neritic Copepoda from the north- west coast of Kyushu.—Symposium on Crus- tacea, Part 1. Marine Biological Association of India, 38-50. Thompson, I. C., & A. Scott. 1903. Report on the Copepoda collected by Professor Herdman at Ceylon in 1902.—Report of Government of Ceylon Pearl Oyster Fisheries. 1 (Supplement), 227-307. Vervoort, W. 1964. Free-living Copepoda from Ifa- luk Atoll in the Caroline Islands with notes on related species.— Bulletin of the United States National Museum 236:1—431. 163 Walter, T.C. 1989. Review of the new world species of Pseudodiaptomus (Copepoda: Calanoida) with a key to the species. — Bulletin of Marine Science 45:590-628. Wells, J. B. J. 1967. The littoral Copepoda (Crusta- cea) from Inhaca Island, Mozambique. — Trans- action of the Royal Society of Edinburgh 67: 189-358. (PH) Regional Centre of National Insti- tute of Oceanography, P.B. 1913, Cochin 682 018, India; (MM) National Institute of Oceanography, Dona Paula, Goa 403 004, India; (SO) Fisheries Laboratory, Hiroshi- ma University, Takehara, Hiroshima 725, Japan. PROC. BIOL. SOC. WASH. 107(1), 1994, pp. 164-173 NEW SPECIES OF MARIONINA (ANNELIDA: OLIGOCHAETA: ENCHYTRAEIDAE) FROM SPARTINA SALT MARSHES ON SAPELO ISLAND, GEORGIA, U.S.A. Brenda Healy Abstract. — Three new species of Marionina are described from Spartina salt marshes where they were associated with live and dead Spartina stems. All three species appear to be confined to this habitat. Marionina spartinae is distinguished by paired sigmoid setae, an anteclitellar origin of the dorsal vessel and elongated sperm bundles in the spermatheca. Marionina waltersi has 2-3 sigmoid setae per bundle, a long, narrow sperm funnel and a poorly developed seminal vesicle. Marionina paludis resembles M. appendiculata in having fan- like bundles of 3-6 sigmoid setae but differs in its larger sperm funnel and well developed seminal vesicle and the presence of a compact penial bulb. These are the first records of Enchytraeidae from the marine littoral zone of Georgia. There have been few studies of marine littoral Enchytraeidae in the east of North America. Published records concern only the northern United States from Maine to South Carolina (Moore 1905, Welch 1917, Lasserre 1971) or Bermuda (Moore 1902, Lasserre & Erséus 1976, Giere 1979). With the exception of a single record of Stephen- soniella marina (Moore, 1902) (Coates 1983) and some preliminary data from West Flor- ida (Healy 1989), there is no information from more southern states. The new species described in this paper are thus the first rec- ords of Marionina from the southern At- lantic coast. Sparina salt marshes, with S. alterniflora Loisel, 1907 (smooth cordgrass) the domi- nant species, cover extensive areas of the intertidal zone from Canada to mid-Florida (where they are replaced by mangroves) and on the Gulf coast as far as Texas (Reimold 1977). On the coast of Georgia, they form a belt approximately 7 km wide and the plants here grow luxuriantly, reaching 2-3 m in height with a basal diameter of 3 cm on the banks of creeks (Wiegert et al. 1981). There is extensive literature on many as- pects of the Sapelo Island marshes, includ- ing their invertebrate fauna, but while oli- gochaetes have been shown to be an important component of the fauna of the marsh surface (Teal 1962), they have not, so far, been identified, even to family. Preliminary investigations of salt marsh oligochaetes on the Sapelo marshes showed that Tubificidae and Enchytraeidae were present on the marsh surface, mainly in plant debris, but that both were far more abun- dant in the lower regions of Spartina stems. The three new species described in this pa- per, together with another enchytraeid, Marionina appendiculata Nielsen & Chris- tensen, 1959 and a tubificid, Monopylepho- rus parvus Ditlevsen, 1904, were among the most abundant faunal species living in the aerenchyma of Spartina leaf sheaths, a mi- crohabitat that supplies worms with food and oxygen and protection from predators (Healy & Walters 1993). The presence of aquatic Oligochaeta in plant aerenchyma has not previously been reported. The three new species can all be referred to Marionina as defined by Nielsen & Chris- tensen (1959). The genus was erected by Michaelsen (1989) to contain a group of spe- cies removed from Pachydrilus, but his di- VOLUME 107, NUMBER 1 agnosis was inadequate to describe the new taxon. Cernosvitov, in his review of the En- chytraeidae (1937), considered the system- atic position of Marionina to be unclear but retained the taxon as a sub-genus of Pachy- drilus in which he included 37 species. Niel- sen & Christensen (1959) subsequently re- moved several species to other genera but admitted that Marionina remained hetero- geneous. Their diagnosis is concerned en- tirely with negative or variable characters and there is, at present, no derived character that distinguishes the genus as a whole. Marionina species are recognized by their small size and the absence of characters that define other genera such as peptonephridia, intestinal diverticula, an anterior origin of the dorsal vessel, two kinds of coelomo- cytes, lobed seminal vesicles and setae with enlarged ental hooks. The genus has thus become a deposit for any small species that do not conform to other existing generic diagnoses. The 70 or so species currently included in Marionina constitute a hetero- geneous assemblage comprising at least two lineages (Coates 1987, 1989) and some spe- cies of doubtful affinity (Nielsen & Chris- tensen 1959; Coates 1980, 1987). Most of the marine species, however, form a ho- mogeneous monophyly, characterized by an anterior bifurcation of the dorsal vessel in III or IV, 1.e., posterior to the brain, instead of anterior to the brain as in most enchy- traeids (Coates 1987, 1989, 1990). The three new species described in this paper fall into this group. They are distinguished from oth- er marine Marionina by setal shape and number, the point of origin of the dorsal blood vessel, the form of the sperm funnel and development of the seminal vesicle and by characters of the spermatheca, including the arrangement of sperm in the ampulla. Materials and Methods Most of the material was collected in May, 1991 from the Kenan Field salt marsh on Sapelo Island (31°23’N, 81°17'W) where 165 samples were taken from high and low marsh and creekside as part of an investigation into the microdistribution of Oligochaeta (Healy & Walters 1993). The substrate was silt and the salinity 20—30%o. A few samples containing one or more of the new species were also taken from Spartina marshes in other parts of Sapelo Island, and from marshes at two sites in N. Florida. The latter collections were part ofa survey of intertidal and supratidal habitats in Georgia and east Florida in 1990 and 1991 (unpublished) which has provided information on the eco- logical limits of the species described in this paper. Samples were taken from surface mud, roots and leaf debris and the lower stems of live and standing dead Spartina plants. Worms were extracted from mud by sieving and from plant material by a modified ver- sion of the wet-funnel method, using 100 W light bulbs to raise the temperature in the funnels over 40°C in 1-2 hours (Healy & Rota 1992). Leaf sheaths were shredded lengthwise before extraction. Specimens were fixed in 70% ethanol, stained in para- carmine and whole-mounted in Canada Balsam. Type specimens and other material are deposited in the United States National Museum of Natural History, Washington, D.C. (USNM). Other whole mounts are in the collection of the University of Georgia Marine Institute, Sapelo Island (UGMI), or in the author’s collection. All material was collected by the author. Marionina spartinae, new species Fig. 1 Material examined. —Holotype: USNM 163805, stained whole mounted specimen, Kenan Field salt marsh, Sapelo Island, May 1991. Paratypes: USNM 163806-163807, stained whole mounted specimens from the type locality, May 1991. Other material from the type locality: USNM 163808-163812; UGMI two whole mounted specimens; 98 166 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON wrlost Fig. 1. on several fixed specimens; B, setal bundle; C, spermatheca of live specimen; D, spermatheca of fixed specimen; E, penial bulb; F, sperm funnel of fixed specimen; G, sperm funnel of live specimen; H, coelomocytes; I, clitellar glands; J, testes from two fixed, mounted specimens; K, nephridium at 8/9, live worm. a, ampulla; b, brain; c, collar of sperm funnel; dbv, dorsal blood vessel; ec, ectal duct; en, ental duct, i, intestine; mp, male pore; oe, esophagus with chloragocytes; pb, penial bulb; phb, pharyngeal bulb; pg, pharyngeal gland; ph, pharnyx; pm, pharyngeal muscles; s, setal bundle; sb, sperm bundle; spt, spermatheca. whole mounted specimens in the author’s collection. Approximately 25 live speci- mens examined. Etymology. —The specific name refers to the salt marsh plant Spartina with which the species is almost exclusively associated. Description. —Length of live worms 5-8 mm, width 0.27 mm at VII, 0.30 mm at the clitellum. Fixed, mounted specimens 4—5 mm long and 0.16—0.22 mm wide. Seg- ments (22)26—32(42) (n = 130). Setae two per bundle in all segments and positions, usually absent in XII, sigmoid without ental hooks, 38-48 um in the preclitellar region, 100 tim Marionina spartinae, new species. A, schematic view of anterior region from combined observations 48-56 um in posterior segments, of roughly equal length within a bundle (Fig. 1B). Cu- taneous gland cells inconspicuous in live specimens but about three double rows may be seen in each segment in stained mounts. Clitellum extending over XII—'2XIII, slight- ly raised, gland cells either in transverse rows, poorly developed, or irregular when well developed, absent mid-ventrally be- tween the penial bulbs (Fig. 11). Head pore in the middle of the prostomium. Three pairs of pharyngeal glands, the first two united dorsally without ventral lobes, the third pair separate with elongated ven- VOLUME 107, NUMBER 1 tral lobes and small dorsal lobes (Fig. 1A). A pair of bulbs present on the posterior bor- der of the pharynx (Fig. 1A). Esophageal diverticula absent. Esophagus merging gradually with the intestine from 6/7. Chloragocytes forming a dense layer from V, 5-8 cells across the intestine, containing small, sparse droplets. Coelomocytes nucle- ate, round or oval, sometimes with small, blunt points, filled with refringent granules, the cells appearing gray or light brown by transmitted light in live worms, length 19- 23 um (Fig. 1H). Blood colorless. Dorsal vessel originating in the preclitellar region, usually at 9/10, occasionally at 10/11, an- terior bifurcation in III (Fig. 1A). Brain about twice as long as its maximum width, 115-120 wm long, indented posteriorly (Fig. 1A). Nephridia starting at 7/8, the antesep- tal part ovoid with coils of the canal sur- rounding the nephrostome, postseptale more or less cylindrical, the efferent duct termi- nal, short and stout (Fig. 1K). Testes bulky, somewhat lobed (Fig. 1J), seminal vesicle unpaired, confined to XI or extending forward asymmetrically to X or 9/10. Sperm funnel half to two-thirds the diameter of the worm, about twice as long as wide in live worms (Fig. 1G), about 1.5 times as long in fixed specimens (Fig. 1F), with a tall, somewhat flared, asymmetrical collar to which abundant, dark-staining sperm are attached. The funnel has a gran- ular appearance and in living worms has an irregular outline. Sperm duct stout, 7-12 um in diameter, of medium length, opening at a small, compact penial bulb, maximum diameter 25-37 um. One to three mature eggs present at a time. Spermathecal am- pulla ovoid, about twice as long as wide, 60-80 um xX 40-55 um, thick-walled in live specimens, the wall not easily distinguish- able in stained mounts, containing elongat- ed sperm bundles (Figs. 1C, D). In live worms, the bundles are transversely striat- ed, but striations are hard to see in stained mounts. Ectal duct short, only about 1.5 167 times the thickness of the body wall, with- out separated, projecting gland cells. Ental duct broad and short, thick-walled, uniting with the esophagus at about the level of the setae in V (Fig. 1A). Remarks. —The distinctive characters of M. spartinae are its paired, sigmoid setae, the preclitellar origin of the dorsal vessel and the unusual arrangement of sperm in the spermathecal ampulla. Only a few spe- cies of Marionina are described as having distinctly sigmoid setae and none of these have paired setae in all segments. An an- teclitellar origin of the dorsal vessel at 8/9 or in IX is unusual in Marionina and has only been reported for three terrestrial spe- cies from S. America: M. ecuadoriensis Righi, 1981, which has free spermathecae, 1.e., not attached to the esophagus; M. cana Marcus, 1965, which has straight setae; and M. nea Marcus, 1965, which has 2-S slight- ly sigmoid setae. In other species the origin is intraclitellar or postclitellar. The arrange- ment of sperm in elongated bundles in the spermatheca is unique in the Enchytraeidae, although in Grania and in several species of Marionina there are spherical sperm bun- dles. The bundles differ from similar shaped bundles found in some Tubificidae (Baker & Brinkhurst 1981, Erséus 1982) in lacking an outer hyaline layer which, as shown by Braidotti et al. (1980), is formed by helically wound, modified spermatozoa with degen- erate nuclei and which surrounds a core of fertilizing spermatozoa. The cross striations seen in the bundles of M. spartinae suggest a different arrangement, but electron mi- croscopic studies would be needed to elu- cidate the structure. Habitat.— Spartina salt marshes, chiefly in the leaf sheaths of live and standing dead Spartina plants, less frequently in surface mud and decaying plant debris. One record from plant debris at the edge of a brackish lake, salinity 22%o. Distribution. —Common and widespread 168 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON wi ost 150 Lum Fig. 2. Marionina waltersi, new species. A, schematic view of anterior region from combined observations on several fixed specimens; B, setal bundle; C, coelomocytes; D, spermatheca of live specimen; E, spermatheca of fixed specimen; F, sperm funnel and part of duct in live specimen; G, segments XI and XII. b, brain; c, collar of sperm funnel; dbv, dorsal blood vessel; e, egg; ec, ectal duct; i, intestine; 0, ovary; oe, esophagus with chloragocytes; n, nephridium; pb, penial bulb with male pore; pd, duct of pharyngeal glands; pg, pharyngeal gland; pm, pharyngeal muscle; s, setal bundle; sf, sperm funnel; spa, spermathecal ampulla; spd, spermathecal diverticulum; sv, seminal vesicle, poorly defined; vdd, distal part of vas deferens; vdp, proximal part of vas deferens. on Sapelo Island; Guana Lake, near St. Au- gustine, Florida. Marionina waltersi, new species Fig. 2 Material examined. —Holotype: USNM 163813, stained, whole mounted specimen, Kenan Field salt marsh, Sapelo Island, May 1991. Paratypes: USNM 163814—-163815, stained, whole mounted specimens from the type locality, May 1991. Other material: UGMI, one stained, whole mounted spec- imen; 25 stained whole mounts in the au- thor’s collection. Approximately 16 live specimens examined. Etymology. —The species is named for Dr. Keith Walters, meiofaunal specialist at the Marine Institute, Sapelo Island, who first drew my attention to the presence of en- chytraeids in Spartina stems. Description. —Live worms 5-6.5 mm, width 0.22 mm in the preclitellar region, 0.27 mm at the clitellum. Fixed, mounted specimens 3-4 mm long and 0.16-0.24 mm wide. Segments (21)25—29(35) (n = 25). Se- tae slightly sigmoid without ental hooks (Fig. 2A), (2)3(4) in preclitellar segments, 2—3 be- VOLUME 107, NUMBER | hind the clitellum, 43-56 wm anteriorly, 48— 60 um in posterior segments. Within a bun- dle, the outer setae are 4—6 wm longer than those near the midline. Cutaneous gland cells not apparent. Clitellum extending over XII-— ¥4X III, gland cells more or less in transverse rows near anterior and posterior borders, arranged irregularly in the central zone, gland cells present in the mid-ventral region. Head pore near the middle of the prostomium. Three pairs of pharyngeal glands, the first two broadly united dorsally, the third unit- ed by a narrow band of tissue, all without ventral lobes (Fig. 2A). Pharyngeal and esophageal diverticula absent. Esophagus merging gradually with the intestine from 6/7. Chloragocytes present from V, forming a dense layer from VI, about 5-6 cells across the intestine, containing fine droplets. Coe- lomocytes oval, 19-25 um, one-half to one- third the length of the setae, with coarse and small granules, appearing gray or light brown by transmitted light in living worms (Fig. 2C). Blood colorless. Dorsal vessel origi- nating at 12/13, in XIII or at 13/14, anterior bifurcation in III or IV. Brain about 1.3— 1.4 times as long as broad (fixed, mounted material), slightly indented. Nephridia starting at 7/8 or 8/9, anteseptale ovoid with coils of the canal surrounding the nephro- stome, postseptale more or less cylindrical, slightly more swollen than in M. spartinae, efferent duct terminal, short and stout. Testes small, unlobed, elongate, some- times extending from 10/11 to mid XI. Seminal vesicles poorly developed (Fig. 2G) or absent. Sperm funnel long and narrow, five to six times as long as wide, roughly equal in length to the diameter of the worm (Fig. 2F). A tall, funnel-shaped collar is usu- ally bent towards the midline while the fun- nel itself is straight or slightly sinuous, ta- pering distally as it passes through 11/12 to form a thin-walled, sinuous duct with iso- lated cells on its surface, about half as long as the funnel (Fig. 2F). The sinuous duct, which represents the proximal region of the 169 vas deferens, is followed by a long, narrow, much-coiled distal region, confined to XII (Fig. 2G). Sperm attached to the funnel no- tably long and wavy (Fig. 2F). Penial bulb compact, 42-48 um in diameter. One or two mature eggs present at a time. Spermathecal ampulla surrounded by a ring of five or six, more or less spherical, thick-walled diver- ticula on short stalks (Figs. 2D, E). Sperm present in the central chamber, in the ra- diating canals and in rings in the bulbous chambers of the diverticula. In live worms, the rings of sperm often rotate. Ental duct very short, the ampulla apparently closely applied to the lateral wall of the esophagus at about the level of the setae in V (Fig. 2A). Ectal duct thick-walled, about 15 um in di- ameter, with a narrow canal, 2.5 times the length of the ampulla, swelling slightly near its opening where there may be a small nar- row chamber. Remarks. — Marionina waltersi resem- bles M. southerni (Cernosvitov, 1937 pro Enchytraeus lobatus Southern, 1909) in having a ring of sperm-containing divertic- ula around the spermathecal ampulla but in M. southerni these are sessile and more nu- merous, there is a rosette of glands sur- rounding the orifice of the ectal duct, the seminal vesicle is large, the sperm funnel thick, the coelomocytes are dense and the paired setae are straight. Marionina south- erni has been recorded in Massachusetts and North Carolina (Lasserre 1971). The only other Marionina for which a ring of sper- mathecal diverticula has been described is the poorly known M. georgiana (Michael- sen 1888) from the Antarctic, which is a larger species (around 35 segments), has up to 6 setae per bundle and a funnel only twice as long as broad. The long, narrow sperm funnel of M. waltersi, followed by a duct in two sections of different thickness has not been described for any other species of Mar- tonina. Habitat.—In Spartina salt marshes, in plant stems and occasionally in surface plant 170 150 um Fig. 3. D, spermatheca of fixed specimen; E, coelomocytes; F, sperm funnel of live specimen; G, sperm funnel of fixed specimen; H, segments XI and XII. a, spermathecal ampulla; b, brain; c, collar of the sperm funnel; dbv, dorsal blood vessel; e, egg; ec, ectal duct; en, ental duct; i, intestine; oe, esophagus with chloragocytes; pb, penial bulb; pg, pharyngeal gland; ph, pharynx; pm, pharyngeal muscle; s, setal bundle; sf, sperm funnel; sv, seminal vesicle; vd, vas deferens. debris. One record from the edge of a brack- ish lake, 22%o. Distribution. —Sapelo Island, Georgia; Guana Lake, near St. Augustine, Florida. Marionina paludis, new species Fig. 3 Material examined. — Holotype: USNM 163816, stained, whole mounted specimen, Kenan Field salt marsh, Sapelo Island, May, 1991. Paratypes: USNM 163817-163818, two stained, whole mounted specimens from the type locality, May 1991. Other material: PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Marionina paludis, new species. A, anterior region; B, setal bundle; C, spermatheca of live specimen; UGMI, three stained whole mounted spec- imens from the type locality; 18 whole mounted specimens from several marshes on Sapelo Island in the author’s collection. Approximately 20 live specimens exam- ined. Etymology.—From the Latin palus—a marsh, because the species is associated with salt marsh habitats. Description. —Live worms 4—6 mm, width 0.18-0.21 mm, 0.24—0.28 mm at the clitel- lum. Fixed, mounted specimens 3.5—4.5 mm long and 0.13-0.26 mm wide. Segments (21)24—28 (n = 25). Setae sigmoid, without VOLUME 107, NUMBER | ental hooks, usually 3-6 per bundle, (2— 3)4(5) laterally and 4—6(7) ventrally in the preclitellar region, 3-4 laterally and 4—5 ventrally behind the clitellum. Setae are ar- ranged fanwise within a bundle, those near the midline being smaller than the outer zones with a stronger curvature (Fig. 3B), size range 28-46 um in the preclitellar re- gion, 40-46 um in posterior segments. Cu- taneous gland cells variable, inconspicuous or in 3-8 rows of transversely elongated cells per segment. Sometimes the cells are fused to form a continuous, irregular line up to half the diameter of the worm which may possess small branches. Clitellum over XII-— YXTII, clitellar gland cells irregularly dis- tributed, absent ventrally in most speci- mens. Head pore at 0/1. Three pairs of pharyngeal glands, the first two pairs broadly united dorsally without ventral lobes, the third pair free with long ventral lobes and small dorsal ones (Fig. 3A). Pharyngeal and esophageal diverticula absent. Esophagus merging gradually with the intestine from 6/7. Chloragocytes form- ing a dense layer from VI, 5-6 cells across the intestine. Coelomocytes mainly oval, some with small blunt points, occasionally round, 19-24 um 1.e., about two-thirds the length of anterior setae, with small granules, appearing gray or light brown by transmit- ted light in live worms. Blood colorless. Dorsal vessel originating at 12/13 or in XIII, anterior bifurcation at around 3/4. Brain about 1.5 times as long as wide in fixed worms, slightly indented posteriorly. Ne- phridia starting at 6/7, similar to those of M. spartinae. Testes more or less globular, not or only slightly lobed (Fig. 3H). Seminal vesicle un- paired, confined to XI (Fig. 3H). Sperm fun- nel cylindrical, confined to XII, twice as long as broad, 80 < 40 um in live worms, about half the diameter of the worm (Figs. 3F, G). Sperm duct of medium length, penial bulb compact, 30 um in diameter. Usually only one mature egg present at a time. Sperma- 171 thecal ampulla cylindrical or cone-shaped, with a rather thick wall (Figs. 3C, D), united with the lateral esophageal wall in the pos- terior part of V (Fig. 3A). Ectal duct about half as long as the ampulla, covered with a layer of rounded cells (Figs. 3C, D). Remarks. — Marionina paludis closely re- sembles M. appendiculata Nielsen & Chris- tensen, 1959 in size and general anatomy, especially in having fan-like bundles of 4— 7 sigmoid setae. It differs in its much larger spermathecal ampulla with shorter ectal duct covered in a layer of large cells, in its larger sperm funnel, larger rounded testes and the presence of a well developed seminal vesicle and compact penial bulb. Marionina ap- pendiculata is characteristic of muddy sub- strates in marine and estuarine littoral zones and is present on the Sapelo marshes. It is a widespread species and, as currently rec- ognized, displays some variations in the size of the sperm funnel, in the development of the seminal vesicle and in the organs as- sociated with the male pore. Some variants may indeed prove to be distinguishable as separate species. In North American ma- terial from western Canada (Coates & Ellis 1981) and from Georgia and both Atlantic and Gulf coasts of Florida (pers. obs.) the sperm funnel is always small, usually only 0.25 the diameter of the worm, the testis is small (elongated in Sapelo specimens), a seminal vesicle is absent and there are one or two free glands at the male pore which are occasionally fused around the opening. Where the species occur together, therefore, M. paludis can be recognized by its larger sperm funnel, larger, rounded testes and well developed seminal vesicle and penial bulb, even when the spermatheca is not clearly distinguished. Immature specimens are dif- ficult to separate although M. appendiculata is generally smaller, with somewhat smaller and finer setae which may reach eight per bundle. Habitat.—Spartina marshes, chiefly in standing dead stems, less frequently in plant 172 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON debris on the marsh surface and in live stems of creekside Spartina. Distribution. —Sapelo Island, Georgia. Discussion The three new species here described were common throughout the Sapelo marshes, Marionina spartinae being dominant. They were accompanied in all microhabitats ex- cept tidal debris by M. appendiculata and on the banks of creeks by immature En- chytraeus. No other enchytraeid species were found on the marshes. Two of the new spe- cies were also present at a Spartina marsh in N. Florida. A variety of other intertidal and supralittoral habitats have been inves- tigated on the S. Georgia and N. Florida coasts, including Salicornia marshes, sea- grass and other tidal debris, and plant roots near high water mark on the open coast. Other species of Marionina were found in these habitats but the new species appear to be confined to Spartina marshes. Acknowledgment Support for my visit to Sapelo Island was provided by the University of Georgia, Sa- pelo Island Research Foundation in the Ma- rine Institute’s Visiting Scientist Program. This is contribution No. 733 of the Uni- versity of Georgia Marine Institute. Literature Cited Baker, H. R., & R. O. Brinkhurst. 1981. A revision of the genus Monopylephorus and redefinition of the subfamilies Rhyacodrilinae and Bran- chiurinae (Tubificidae: Oligochaeta).— Canadi- an Journal of Zoology 59:939-965. Braidotti, P., M. Ferraguti, & T. P. Fleming. 1980. Cell junctions between the spermatozoa flagella within the spermatozeugmata of Tubifex tubifex (Annelida: Oligochaeta).—Journal of Ultra- structure Research 73:299-309. Cernosvitov, L. 1937. System der Enchytraeiden.— Bulletin de l’Association suisse pour les Re- cherches scientifiques 4 Prague 5:263-294. Coates, K. A. 1980. New marine species of Marion- ina and Enchytraeus (Oligochaeta, Enchytraei- dae) from British Columbia.—Canadian Jour- nal of Zoology 58:1306—1317. . 1983. A contribution to the taxonomy of the Enchytraeidae (Oligochaeta). Review of Ste- phensoniella with new species records.—Pro- ceedings of the Biological Society of Washington 96:411-419. 1987. Phylogenetic analysis of some Enchy- traeidae (Annelida: Oligochaeta): parsimony analysis of structural changes. Unpublished PhD thesis, University of Victoria, Canada. 1989. Phylogeny and origins of Enchytraei- dae.— Hydrobiologia 180:35—-45. 1990. Marine Enchytraeidae (Oligochaeta, Annelida), of the Albany area, Western Austra- lia. Pp. 13-41, in F. E. Wells, D. I. Walker, H. Kirkman & R. Lethbridge, eds., Proceedings of the third international marine biological work- shop: the marine flora and fauna of Albany, Western Australia. Western Australian Muse- um, Perth. ——.,, & D. V. Ellis. 1981. Taxonomy and distri- bution of marine Enchytraeidae (Oligochaeta) in British Columbia.— Canadian Journal of Zo- ology 59:2129-2150. Erséus, C. 1982. Taxonomic revision of the marine genus Limnodriloides (Oligochaeta: Tubifici- dae).— Verhandlungen naturwissenschaftlichen Vereins in Hamburg, NF 25:207-277. Giere, O. 1974. Marionina istriae n. sp., ein mariner Enchytraeide (Oligochaeta) aus dem mediter- ranean Hygropsammal.—Helgolander wissen- schaftliche Meeresuntersuchungen 26:359-369. 1979. Studies on marine Oligochaeta from Bermuda, with emphasis on Phallodrilus species (Tubificidae).— Cahiers de Biologie marine 20: 301-314. Healy, B. 1989. Preliminary report on the Enchy- traeidae (Oligochaeta) of West Florida.—Hy- drobiologia 108:47-S6. —, & E. Rota. 1992. Methods for collecting En- chytraeidae during expeditions.—Soil Biology and Biochemistry 24:1279-1281. ——,, & K. Walters. 1993. Oligochaeta in Spartina stems: the microdistribution of Enchytraeidae and Tubificidae in a salt marsh, Sapelo Island, U.S.A.—Hydrobiologia (in press). Lasserre, P. 1971. The marine Enchytraeidae (An- nelida, Oligochaeta) of the eastern coast of North America with notes on their geographical dis- tribution and habitat.— Biological Bulletin of the Marine Biological Laboratory, Wood’s Hole 140: 440-460. —, & C. Erséus. 1976. Oligochétes marins des Bermudes. Nouvelles éspéces et remarques sur la distribution géographique de quelques Tu- bificidae et Enchytraeidae. — Cahiers de Biologie marine 17:447—-462. VOLUME 107, NUMBER 1 Marcus, E. 1965. Naidomorpha aus brasilianischem Brackwasser.— Beitrage zur neotropischen Fau- na 4:61-83. Michaelsen, W. 1988. Die Oligochaeten von Sud- Georgien nach der Ausbeute der deutschen Sta- tion von 1882-1883. Jahrbuch Hamburger wis- senschaftlichen Austatten 5:53—73. 1889. Synopsis der Enchytraeiden.—Ab- handlungen des naturwissenschaftlichen Ve- reins zu Hamburg | 1:1—60. Moore, J. P. 1902. Some Bermuda Oligochaeta with a description of a new species.— Proceedings of the Academy of Natural Sciences of Philadel- phia 54:80-84. 1905. Some marine oligochaeta of New En- gland.— Proceedings of the Academy of Natural Sciences of Philadelphia 57:373-399, Plates 22— D3. Nielsen, C. O., & B. Christensen. 1959. The Enchy- traeidae. Critical revision and taxonomy of Eu- ropean species.— Natura jutlandica 8—9:1—160. Reimold, R. J. 1977. Mangals and salt marshes of eastern United States. Pp. 157-166 in V. J. Chapman, ed., Ecosystems of the world. 1. Wet 173 coastal ecosystems. Elsevier Scientific Publish- ing Co., Amsterdam. Righi, G. 1981. Alguns Oligochaeta cavernicolas do Equador.—Papéis Avulsos de Zoologia 34:235— 249. Southern, R. 1909. Contributions towards a mono- graph of the British and Irish Oligochaeta.— Proceedings of the Royal Irish Academy 27B: 119-182. Teal, J. M. 1962. Energy flow in the salt marsh eco- system of Georgia.— Ecology 43:614-624. Wiegert, R. G., L. R. Pomeroy, & W. J. Wiebe. 1981. Ecology of salt marshes: an introduction. Pp. 3— 19 in L. R. Pomeroy and R. G. Wiegert, eds., The Ecology of a salt marsh. Springer-Verlag, New York. Welch, P.S. 1917. The Enchytraeidae (Oligochaeta) of the Wood’s Hole region, Mass.—Transac- tions of the American Microscopical Society 34: 119-138. Department of Zoology, University Col- lege, Belfield, Dublin 4, Republic of Ireland. PROC. BIOL. SOC. WASH. 107(1), 1994, pp. 174-178 SUMMARY AND SIGNIFICANCE OF OVERLOOKED JAPANESE LITERATURE ON MYZOSTOMIDA Mark J. Grygier Abstract. —The content of hitherto uncited or unreviewed Japanese scientific literature on Myzostomida is summarized with comments. The range of My- zostoma willemoesii von Graff has been extended to Japan, with an additional host record (a recent unpublished record from the Ogasawara Islands is also given). An associated myzostome has been reported from the hexactinellid sponge Farrea. Myzostoma crassum Okada, 1922 is an overlooked but available nomen dubium. Cleavage has been studied in M/. deani McClendon. Myzostoma echinus von Graff has been misidentified consistently as M/. costatum Leuckart in Japanese literature, and another species whose true identity is unclear has often been misidentified as M. ambiguum von Graff. Other possible recent misidentifications are pointed out herein. Notopharyngoides Fishelson, nomen nudum, has inadvertently been made available as Notopharyngoides Uchida, 1992, with the type species Myzostoma ijimai Hara & Okada, 1921. Most of the original descriptions of Jap- anese species of myzostomidan worms (Jap- anese name: kydk6chu or suikuchimushi), found as quasi-parasitic commensals or as endoparasites of crinoids and ophiuroids, were published by European or American zoologists (e.g., von Graff 1884, McClendon 1906, Jagersten 1937, Fedotov 1938). While several taxonomic and developmental pa- pers written in English by Japanese scien- tists are well known (Hara & Okada 1921; Okada 1922b, 1933; Kato 1952), a consid- erable amount of related literature, mostly written in Japanese, seems not to have come to the attention of Western zoologists. A few such papers have been cited in bibliogra- phies by Hartman (1951), Kato (1952), and Prenant (1959), but the findings were either not reviewed or misquoted. The contents of these largely ignored works are summarized here, with remarks on their significance. It should be noted in advance that the cited faunal encyclopedias and field guides were written and edited by eminent zoologists and are or once were in day-to-day use as reference books in Japanese laboratories; thus they are regarded here as an intrinsic part of the zoological literature. Although Myzostoma, not Myzostomum, is the cor- rect original spelling of the main genus in question (Grygier 1992), names are spelled here following the cited Japanese authors. Historical Review and Comments Hara (1895) reviewed a controversy sur- rounding the supposed excretory organs (1.e., metanephridia) of Myzostoma. On p. 247 the nomen nudum M. Jjimei appeared in the context of histological information about an undescribed species. This was evidently a reference to the species later published as M. timai Hara & Okada, 1921. Hara (1896) reported that 20 specimens of Antedon inaequalis Carpenter collected at Misaki bore galls caused by Myzostomum willemoesii von Graff. The supposed host now belongs to the genus G/yptometra, but the range of G. inaequalis does not extend to Japan (Clark 1950); two other nominal species of G/yptometra have been recorded from Misaki, G/yptometra lata (A. H. Clark) VOLUME 107, NUMBER 1 and G. septentrionalis (A. H. Clark), and presumably one of these was the host. Gry- gier (1990) was unaware of Hara’s (1896) record and neither included M. willemoesii in a count of Japanese myzostome species nor took the otherwise unrecorded host into account in a discussion of host specificity. This myzostome has been known to infest eight other hosts, including four species of Glyptometra, and Hara’s report extends its range northward from the nearest record in the central Philippines (Grygier 1990). The Philippine record, based on speci- mens in the National Museum of Natural History, Smithsonian Institution (USNM 167184), is fully documented here: two galls on an arm of one Glyptometra tuberosa (Carpenter) (USNM 35664), “Albatross” sta. 5537, 19 August 1909, between Negros and Siquijor, 9°11'00”N, 123°23’00’E, 465 m. There is another lot of M. willemoesii in the Osaka Museum of Natural History (Cat- alogue no. Iv 1293) from an intermediate locality near the Japanese Ogasawara (Bo- nin) Islands: four galls formed from en- larged, twisted pinnules on one G. tuberosa (no. 700(2)), “Sdéy6-Maru” Cruise 5 (dl- d10), 4 August 1976, 27°05.9-23.0'N, 142°03.8-05.8'E, 165-345 m. Komai (1919) briefly introduced the My- zostomida as a whole, summarized Mc- Clendon’s (1906) taxonomic paper, and re- produced figures from it. Okada (1920) reported four specimens of Myzostoma collected from the hexactinellid sponge Farrea sp. He assigned the speci- mens to Myzostoma antennatum von Graft based on morphological resemblance, al- though they were of a different color. This is the only report of myzostomes collected from a sponge, although it might be sus- pected that the worms had been lost acci- dentally from a crinoid or ophiuroid during collection. Okada (1922a) published a key to the My- zostomida of Japan preceded by a long, lit- erature-based, general introduction of the group. The key included 13 nominal species 175 of Myzostoma, one of which, My. (sic) cras- sum seems never to have been fully de- scribed. In the accompanying table, this species was referred to as M. crassum H. (for Hara), but I have been unable to locate any published description by Hara. Inas- much as this name appeared in a key, it was accompanied by a diagnosis and is available with Okada as its author; however, it is an unrecognizable nomen dubium. According to the relevant couplet, in Mg. (sic) anten- natum the position of the female (i.e., male; lapsus or misprint of a Japanese kanji char- acter) genital opening is directly outside the third parapodium while in M. crassum the male opening lies between the third para- podium and (which?) sucker (i.e., lateral or- gan). This is a very subtle distinction. The host of M. crassum is Antedon macrodiscus Hara (i.e., Tropiometra afra macrodiscus) from near-coastal waters off Misaki, Kana- gawa Prefecture. My own extensive survey (Grygier 1990) turned up only three species of myzostomes on this host, M. bocki (Ja- gersten), /. nasonovi Fedotov, and M. iji- mai, all of which can be distinguished from M. antennatum by gross morphology and are thus unlikely to be synonymous with M. crassum. Okada (1930) briefly described the early cleavage of embryos of Myzostoma deani McClendon, misspelled as M. cleani when reviewed by Kato (1952), from Enoshima, Tateyama Bay, Chiba Prefecture. This is the only report about developmental biology in this species. Much more recently, Utinomi & Kogo (1965) recorded: 1) Myzostomum sp. from Comanthus (Cenolia) japonica (Miller) (i.e., Oxycomanthus japonicus), 2) Myzostomum sp. and M. costatum Leuckart from Coman- thus (Comanthus) parvicirra (Miller), and 3) Myzostomum sp. from Comanthina schlegeli (Carpenter). All were from the southwestern part of the Kii Peninsula (es- pecially Tanabe Bay), Wakayama Prefec- ture. The worms from C. schlegeli were found in sub-epidermal cysts on the arms 176 or discs, with openings to the ambulacral grooves. The collections of the Seto Marine Bio- logical Laboratory, where Kogo did this work, include four catalogued vials labelled Myzostomum costatum, but all are mis- identified. Poly 138 was isolated from Co- manthus parvicirra but now the vial con- tains only a dry crinoid arm fragment. Poly 139, host unstated, contains four M. fissum von Graff and seven small specimens that may also belong to that species or to an undescribed, closely related species. Poly 140, host unstated, contains two individu- als of a probably undescribed species in the M. crosslandi species-group sensu Grygier (1990). Poly 137, host unstated, contains one specimen which is similar to the Jap- anese myzostome, host unknown, which Ja- gersten (1940) identified as M. cf. insigne Atkins and which was reexamined by the author in the Zoological Museum in Co- penhagen in 1986. Being unpigmented, both specimens more closely match Atkins’s (1927) original description of M. pottsi. Grygier (1990) considered M. insigne and M. pottsi to be junior synonyms of M. echi- nus von Graff. Photographs of so-called M. costatum in some handbooks (Utinomi 1964; Imajima 1975a, 1983a) actually show M. echinus with the color pattern described by Atkins (1927) for M. insigne. Uchida (1992) mentioned that M. costatum was found on Lamprometra palmata (Miller) and Comanthina schlegeli in Japan, but did not include an illustration by which to judge his identification. The species referred to as M. ambiguum in all Japanese primary and secondary lit- erature is also misidentified. Treatment of itas M. ambiguum dates to Jagersten (1937), who showed that it was distinct from M. antennatum, and whose opinion was fol- lowed by Fedotov (1938), Kato (1952), and the compilers of several field guides and fau- nal encyclopedias (Kato 1960; Okada & Kato 1965, 1979: Nishimura & Suzuki 1971; Imajima 1975b, 1983b). Grygier (1990) PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON considered the Japanese “M. ambiguum” to be a different species, but whether some of the earlier records actually correspond to M. longimanum (Jagersten), M. vastum von Graff (sensu von Graff 1884), or an unde- scribed species is still uncertain. True M. ambiguum has a wide tropical distribution and can easily be recognized on the basis of von Graff's (1887) illustration, particularly with regard to the rather narrow and trun- cated rear end of the body and the barely submarginal location of the proboscis pock- et opening. In Japan, true M. ambiguum does occur in Okinawa, based on specimens identified by the author in the Department of Marine Sciences, University of the Ryu- kyus. A recent guide to marine invertebrates of Okinawa Prefecture, southern Ryukyus Is- lands (Iwase et al. 1990:91) included pho- tographs of living Myzostomum elegans van Graff (sic) and Myzostomum limai Hare et Okada (sic), the former on an arm of Co- manthus parvicirra, the latter removed from the mouth of Tropiometra afra macrodis- cus. The bull’s-eye color pattern of the for- mer corresponds to that of Myzostoma polycyclus Atkins (cf. Atkins 1927), which was (implicitly) treated as separate from M. elegans by Grygier (1990). Uchida (1992) labelled other photographs of perhaps the same individuals as Myzostomum polycy- clus. The supposed M. ijimai, the same in- dividual of which appears in a photograph in Uchida (1992), has a series of discrete submarginal lappets like those of M. platy- pus von Graff and M. aruense Remscheid, unlike the continuous, wavy-edged, sub- marginal brim of M. ijimai (cf. Hara & Oka- da 1921, Jagersten 1937, Kato 1960, Okada & Kato 1965; the species is called Myzos- tomum ijimae in the last two references). The specific identification of the Ryukyuan specimen is thus questionable despite its oc- currence on the proper host. Noting that the proboscis protrudes from the dorsal surface in M. ijimai, Uchida (1992) remarked that the genus Notophar- VOLUME 107, NUMBER 1 yngoides had been proposed for that reason. For lack of a diagnosis, Notopharyngoides was a nomen nudum when introduced by Fishelson (1974, 1976), but Uchida’s (1992) remarks constituted a diagnosis for it, and its use in connection with a single species name constituted a type species designation. Therefore, Uchida (1992) inadvertently validated this former nomen nudum and must be recognized as the author of Noto- pharyngoides, with the type species Myzos- toma ijimai. Notopharyngoides may be con- sidered equivalent to the M. platypus species-group sensu Grygier (1990). Uchida (1992) presented a photograph of Myzostomum bocki on its host, Tropiome- tra afra macrodiscus, but the banded color pattern of the worm strongly suggests that it is really Myzostoma nasonovi. Grygier (1990) considered these two species to be synonymous, but after examining speci- mens of both found that they can be distin- guished unambiguously by the banding and the parapodial hook apparatus (Grygier 1992). Acknowledgments I conducted the literature search at the Seto Marine Biological Laboratory while holding a Postdoctoral Fellowship from the Japan Society for the Promotion of Science under the auspices of the National Science Foundation Japan Program. I thank my host, Dr. E. Harada, and, for translation assis- tance, Dr. S. Yamato. Dr. R. Yamanishi kindly allowed me to isolate myzostomes from crinoids in the Osaka Museum of Nat- ural History. The manuscript was prepared during a term as a Visiting Foreign Re- searcher at the Sesoko Marine Science Cen- ter, supported by the Japanese Ministry of Education, Science and Culture, and I thank my host, Dr. K. Yamazato. Literature Cited Atkins, D. 1927. Report on the Myzostomida col- lected by Mr. F.A. Potts in Torres Strait, to- gether with a description of a species obtained 177 by Professor J. Stanley Gardiner from the Mal- dives. — Proceedings of the Zoological Society of London 1927:339-357. Clark, A. H. 1950. A monograph of the existing cri- noids. Volume 1. The comatulids. Part 4c.— Superfamily Tropiometridae (the families Tha- lassometridae and Charitometridae).— United States National Museum Bulletin 82(1,2):xxv + 1-795, pls. 1-57. Fedotov, D. M. 1938. Spezialisation und Degrada- tion im K6rperbau der Myzostomiden in Ab- hangigkeit von der Lebensweise.— Acta Zoolo- gica, Stockholm 19:353—385. Fishelson, L. 1974. Ecology of northern Red Sea cri- noids and their epi- and endozoic fauna. — Ma- rine Biology 26:183—192. —. [1973] 1976. Ecology of the crinoids of the northern Red Sea with emphasis on epi- and endozoic fauna associated with them.— Journal of the Marine Biological Society of India 15: 461-473. Grygier, M. J. 1990. Distribution of Indo-Pacific My- zostoma and host specificity of comatulid-as- sociated Myzostomida.— Bulletin of Marine Science 47:182-191. 1992. Hong Kong Myzostomida and their Indo-Pacific distributions. Pp. 131-147 in B. Morton, ed., The marine flora and fauna of Hong Kong and southern China III. Proceedings of the Fourth International Marine Biological Workshop: the marine flora and fauna of Hong Kong and Southern China, Hong Kong, 11-29 April 1989. Hong Kong University Press, Hong Kong. Hara, J. 1895. [On the excretory organ of Myzostom- ida (from an unfinished manuscript).]—Dobu- tsugaku Zasshi (Zoological Magazine, Tokyo) 7:244-249 (in Japanese). 1896. [A crinoid collected at Misaki] (Ante- don inaequalis, P.H.C.).—Dobutsugaku Zasshi (Zoological Magazine, Tokyo) 8:330-333 (in Japanese). ——, & Y. Okada. 1921. Two new species of My- zostoma.—Annotationes Zoologicae Japonen- ses 10:33-39. Imajima, M. 1975a. Tatesuikuchimushi Myzosto- mum costatum Leuckart. P. 264 in H. Utinomi, ed., Gakken chukosei zukan. Suisei dobutsu. (Junior and senior high school student illus- trated encyclopedia. Aquatic animals). Gakken Co., Ltd., Tokyo (in Japanese). 1975b. Tsunosuikuchimushi Myzostomum ambiguum Graff. P. 270 in H. Utinomi, ed., Gakken chukosei zukan. Suisei dobutsu. (Junior and senior high school student illustrated en- cyclopedia. Aquatic animals). Gakken Co., Ltd., Tokyo (in Japanese). 1983a. Tatesuikuchimushi Myzostomum 178 costatum Leuckart. P. 264 in H. Utinomi, ed., Gakken illustrated nature encyclopedia. The aquatic lower animals of Japan. Gakken Co., Ltd., Tokyo (in Japanese). 1983b. Tsunosuikuchimushi Myzostomum ambiguum Graff. P. 270 in H. Utinomi, ed., Gakken illustrated nature encyclopedia. The aquatic lower animals of Japan. Gakken Co., Ltd., Tokyo (in Japanese). Iwase, F., K. Nomura, T. Fukuda, H. Omae, & H. Uchida. 1990. Okinawa kaichu seibutsu zu- kan. (Okinawa marine life illustrated encyclo- pedia). Lesser animals.—Shinsei Zukan Series, Marine Parks Center, Tokyo 11:1—272. (in Jap- anese). Jagersten,G. 1937. Myzostomiden von Prof. Dr. Six- ten Bocks Expedition nach Japan und den Bo- nin-Inseln 1914.—Arkiv for Zoologi 29A(17): 1-35, Taf. 1-2. . 1940. Neue und alte Myzostomum-Arten aus dem Zoologischen Museum Kopenhagen.— Vi- denskabelige Meddelelser fra Dansk Naturhis- torisk Forening 1 Kgbenhavn 104:103-125, Taf. 1. Kato, K. 1952. Onthe development of myzostome. — Science Reports of Saitama University, Series B (Biology and Earth Sciences) 1:1—16, pls. I- III. 1960. 6. Tsunosuikuchimushi Myzostomum ambiguum Graff. 7. lijimasuikuchimushi My- zostomum ijimae Hara & Okada. In Y. K. Oka- da, & T. Uchida, eds., Encyclopaedia zoologica illustrated in colours. Hokuryu-kan Publishing Co., Ltd., Tokyo, 4:49 (in Japanese). Komai, T. 1919. [Myzostomida (strange and rare an- imals, part 8).] Dobutsugaku Zasshi (Zoological Magazine, Tokyo).—31:364—366 (in Japanese). McClendon, J. F. 1906. The myzostomes of the ‘Al- batross’ Expedition to Japan.— Bulletin of the American Museum of Natural History 23:119- 130, pls. XV—XVII. Nishimura, S., & K. Suzuki (supervised by H. Uti- nomi). 1971. Common seashore animals of Japan in color. Hyojun genshoku zukan zenshu, Hoikusha Publishing Co., Ltd., Osaka, 16:1— 196, pls. 1-64 (in Japanese). Okada, Y. 1920. [One species of Myzostomida at- tached to a hexactinellid sponge.].—Dobutsu- gaku Zasshi (Zoological Magazine, Tokyo) 32: 324-325 (in Japanese). 1922a. [Key to the Myzostomida of Ja- pan.].—Dobutsugaku Zasshi (Zoological Mag- azine, Tokyo) 34:365-373 (in Japanese). 1922b. Ona new Protomyzostomum (P. sa- gamiense, sp. nov.) from the ovary of Gorgono- cephalus.—Annotationes Zoologicae Japonen- ses 10:165-169. 1933. Mesomyzostoma katoi, n. sp., an in- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON teresting myzostome found in the gonad of Comanthus japonicus.—Annotationes Zoolo- gicae Japonenses 14:185-189. 1930. [Early development of Myzostomida parasitizing comatulids.].—Dobutsugaku Zas- shi (Zoological Magazine, Tokyo) 42:363-364 (in Japanese). ——,, & K. Kato. 1965. 305. Sagamisuikuchimushi Protomyzostomum sagamiense Okada. 306. Tsunosuikuchimushi Myzostomum ambiguum Graff. 307. lijimasuikuchimushi Myzostomum Yimae (Hara et Okada). 308. Butosuikuchi- mushi Myzostomum robustum Hara et Okada. In Y. K. Okada, S. Uchida, & T. Uchida, eds., New illustrated encyclopedia of the fauna of Ja- pan. Hokuryu-kan Publishing Co., Ltd., Tokyo, 1:575 (in Japanese). ——,, & 1979. 702. Sagamisuikuchimushi Protomyzostomum sagamiense Okada. 703. Tsunosuikuchimushi Myzostomum ambiguum Graff. P. 206 in T. Uchida, ed., Illustrated en- cyclopedia of the fauna of Japan newly com- piled. Hokuryu-kan Publishing Co., Ltd., Tokyo (in Japanese). Uchida, H. 1992. Annelida. Pp. 310-313 in S. Nishi- mura, ed., Guide to seashore animals of Japan with color pictures and keys, Vol. I. Hoikusha Publishing Co., Ltd., Osaka (in Japanese). Utinomi, H. 1964. Coloured illustrations of seashore animals of Japan.— Hoikusha no genshoku zu- kan, Hoikusha, Osaka 8:1—168, pls. 1-64 + I- 12 (in Japanese). ——,, & I. Kogo. 1965. On some comatulids from the coastal sea of Kii Peninsula.— Publications of the Seto Marine Biological Laboratory 13: 263-286. von Graff, L. 1884. Report on the Myzostomida col- lected during the voyage of H.M.S. Challenger during the years 1873—76.—Report of the Sci- entific Results of the Voyage of H.M.S. Chal- lenger during the years 1873-76, Zoology 10:1— 82, pls. I-XVI. 1887. Report on the Myzostomida collected during the voyage of H.M.S. Challenger during the years 1873-76. Supplement.— Report of the Scientific Results of the Voyage of H.M.S. Chal- lenger during the years 1873-76, Zoology 20:1— 16, pls. I-IV. Seto Marine Biological Laboratory, Kyo- to University, Shirahama, Nishimuro, Wa- kayama 649-22, Japan and Sesoko Marine Science Center, University of the Ryukyus, Sesoko 3422, Motobu-cho, Okinawa 905- 02, Japan. Current address: 14804 Notley Road, Silver Spring, Maryland 20905, U.S.A. \ PROC. BIOL. SOC. WASH. 107(1), 1994, pp. 179-184 SCIURODENDRIUM GARDNERI, NEW SPECIES (NEMATODA: TRICHOSTRONGYLOIDEA: HELIGMONELLIDAB), A PARASITE OF SCIURUS CAROLINENSIS GMELIN, 1788 (MAMMALIA: SCIURIDAE), WITH COMMENTS ON THE BIOGEOGRAPHY OF SCIURODENDRIUM DURETTE-DESSET, 1971 Ricardo Guerrero Abstract. —Sciurodendrium gardneri, new species, is described from an east- ern gray squirrel, Sciurus carolinensis, collected in Virginia, U.S.A. A nearctic origin for the genus Sciurodendrium is proposed on the basis of the paleobioge- ography of the host. Sciurodendrium Durette-Desset, 1971, is a genus found only in New World squirrels (Sciuridae). There are five known species (Durette-Desset & Justine 1992): one nearc- tic, Sciurodendrium hassalli (Price 1929); and four neotropical, S. oliverai (Lent & Freitas 1938), S. hepaticum (Lent & Freitas 1938), S. aripense (Baylis 1947), and S. lan- dauae (Durette-Desset 1970). Sciuroden- drium hassalli is the name that has been used for heligmonellid parasites of squirrels in the United States (Harkema 1936, Chan- dler 1942) on the assumption that there was only one species of this genus in the North- ern Hemisphere. However, a routine ex- amination for parasites in an eastern gray squirrel from Virginia revealed an unde- scribed species. The parasites were collected in an isoton- ic solution of NaCl, preserved in ethanol (70%), and clarified in lactophenol. All mea- surements are in microns and given as the mean followed by the range in parentheses. Types are deposited in the US National Par- asite Collection, USDA, ARS Beltsville, Maryland (USNP), and the Coleccion de Parasitologia, Museo de Biologia, Univer- sidad Central de Venezuela, Caracas (CP- MBUCYV). Sciurodendrium gardneri, new species Description. —Heligmonellidae, Pudici- nae; small slender worms, reddish in color when fresh. Cuticle of the anterior end di- lated and coarsely striated. The mouth opening is triangular and bears an internal ring of six small papillae and an external ring consisting of the amphids and four well- developed papillae (Fig. 1A). Synlophe with a well-developed carene consisting of two continuous ridges (Fig. 1C). In addition, the synlophe includes nine dorsal discontinu- ous ridges arranged in nonalternating inter- rupted longitudinal lines and eight ventral discontinuous ridges in alternating lines (Fig. 1D) resulting in the doubled number of ven- tral ridge “‘tips”’ seen in cross-sectional view (Fig. 1C). Male (10 specimens): Body 3720 (3179- 4493) long by a maximum width of 82 (74— 94) near mid body. Cephalic vesicle is 52 (47-57) long and 32 (28-38) wide. Esoph- agus 1s simple, 250 (208-288) long. Nerve ring and excretory pore located 146 (132- 165) and 169 (154-184), respectively, from the anterior end. Caudal bursa is type 2-2-1 and the dorsal ray is deeply cleft to the level of the origin of 8th ray; ray 9 shorter and PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 180 ‘1 ¢Z sjenbs seq yor “(e0vjIns [eI}USA = A “[esIOP = P) OpIs 143 Wo sospL [euIpNyTsuo] Jo mata “s ‘q ‘Apo JO Yjdua]-piu Ivou sydojuAs Zurmoys UONO9S-SSOID “ “D ‘O[OISAA OT[eYdeo Jo aseq JO JOAg] 1¥ OYdoyUAS BuIMOYs UOT}das-ssolo “s ‘gq ‘MOIA [eoIde “ ‘VY ‘soloads Mou ‘UaUpsDs wntpuapouning *| BL VOLUME 107, NUMBER | hs ne Ge Fig. 2. Sciurodendrium gardneri, new species: 6, posterior end showing characteristics of bursa and rays. Bar equals 25 w. thicker than ray 10; ray 4 gross and with papilliform end; ray 6 arises from basal third of ray 5 (Fig. 2). Spicules are subequal, fi- liform, alate, fused at the tips, and measure 370 (334420) long and 4—5 wide. Telamon is elongated, weakly cutinized, and located close to cloacal papillae. A gubernaculum is absent. Female (10 specimens measured): Body 5360 (4801-5881) long with a maximum width of 98 (92-104). Cephalic vesicle is 54 (48-60) long and 33 (30-37) wide. Esoph- agus is simple, 272 (246-298) long. Nerve ring and excretory pore situated 187 (156— 210) and 221 (200-254), respectively, from the anterior end (Fig. 3). Vulva located 173 (156-188) from the posterior end of the body, and followed by a vagina vera 31 (2440) in length, a ves- tibule 87 (76-100) long, a sphincter 33 (28— 40) long, and an infundibulum 140 (94-210) long (Fig. 4). Uterus measures 316 (220- 490) in length and contains 6 (4-7) thin- shelled eggs, each measuring 73 (68-84) by 38 (36-42). Tail is 84 (59-108) in length. Host: Sciurus carolinensis Gmelin, 1788 (Rodentia: Sciuridae) Location: Small intestine. Locality: Oakton, Fairfax Co., Virginia, U.S.A. Type specimens: Holotype: CP-MBUCV No. 3942 (male). Allotype: CP-MBUCV No. 3943 (fe- male). Paratypes: CP-MBUCV No. 3109 (19 males and 11 females); USNM Helm. Col- lection No. 82704 (2 males and 2 females). Etymology. —Patronym, in honor of Al- fred L. Gardner, of U.S. National Biological Survey, due to his very important contri- butions to Neotropical mammalogy. Remarks. —Sciurodendrium gardneri is 182 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON \ Fig. 3. Sciurodendrium gardneri, new species: 2°, anterior end of body showing positions of nerve ring and excretory pore. Bar equals 25 uy. similar to S. hassallii and S. oliverai and differs from other known congeners on the basis of small body size and deeply cleft dorsal ray of caudal bursa (cleft to level of origin of ray 8). It differs from S. hassalli and S. oliverai in that rays 4 and 9 are con- spicuously gross and broader than the other rays. All rays in S. hassalli are thin as is usual in other members of the group. Also, S. gardneri is smaller than S. hassalli in size and differs from S. oliverai in having a tel- amon (absent in the latter). Discussion. —Durette-Desset (1985:290) stated ““Trichostrongyloids of Neotropical Sciuridae belong to a single genus, Sciuro- dendrium, a Pudicinae derived from forms in Caviomorph rodents.” “It is therefore likely that the Sciuridae lost their brevis- triatine parasites during their migration into North America [from the Old World] and were reinfected after contacting South American pudicines.” Durette-Desset (1971) introduced this hypothesis in her revision of heligmosomes, and has repeated it in subsequent papers (Durette-Desset 1982, 1985; Durette-Des- set & Chabaud 1977, 1981; Durette-Desset & Justine 1991). However, my finding an- other species of Sciurodendrium in a North American squirrel and a review of holarctic heligmonellids does not support Durette- Desset’s (1985) hypothesis on the origin of pudicine parasites of squirrels. The Sciuridae are known in Europe from the Lower Oligocene (+35 million years ago [mya]; Hartenberg 1985), in North America from the Hemingfordian (+16 mya; Webb 1985), but only recently in South America from the Lujanensian (<1 mya; Reig 1981). Two major intercontinental migrations, first across Beringia into North America and then across the Isthmus of Panama (Moore 1961), occurred before squirrels became es- tablished and radiated in South America. Durette-Desset (1985:290) wrote “We be- lieve that sciurids were devoid of tricho- strongyloids when they moved into South America. Trichostrongyloids are rare in holarctic Sciuridae and those that exist be- long to a recent family, the Heligmosomi- dae.’ According to the cladistic analysis of Durette-Desset & Justine (1992), Sciuro- VOLUME 107, NUMBER 1 dendrium must have appeared only a few hundred thousand years ago and evolved from parasites of the caviomorph families Dasyproctidae, Echimyidae, and especially the Capromyidae. An alternative explana- tion is that sciurids entering South America carried brevistriatine heligmonellids, and that pudicines subsequently acquired from caviomorph rodents quickly displaced brevistriatines in all New World squirrels. My research suggests, however, that nei- ther of these hypotheses is supported by the evidence. Other Recent mammals that en- tered South America from North America retained their trichostrongyloid parasites. For example, Longistriata, a genus of par- asites of holarctic Soricidae also is present in South American shrews (Guerrero 1982). The same is true for Vexillata, a parasite (Guerrero 1984) of the Heteromyidae, a New World rodent family that has its greatest diversity in North America. It seems un- likely that the Sciuridae were without trich- ostrongyloids throughout their 16-million- year history in North America before they were able to disperse across the Panamanian isthmus. If derived from caviomorph ro- dents, how did Sciurodendrium reach tem- perate North America? There is no evidence of a “reverse” migration of sciurids from South America northward. Durette-Desset (1971) commented that Brevistriata (Brevistriatinae), a parasite of Oriental Sciuridae, and Sciurodendrium, a parasite of New World sciurids, are quite similar. This she attributed to convergence, pointing out that in the former the cuticular ridges are discontinuous and in alternating lines, and ray 4 is equal to or longer than the 5th. The characteristics I describe above for S. gardneri show that discontinuous al- ternating cuticular ridges are not diagnostic only of Brevistriata. Relative lengths of rays 4 and 5 may be the only remaining major diagnostic character distinguishing the two subfamilies. I suggest that Sciurodendrium is nearctic in Origin and closely related to some Brev- 183 Fig. 4. Sciurodendrium gardneri, new species: 2, posterior end of body. Bar equals 25 pu. 184 istriatinae such as, for example, Calypso- strongylus Schmidt, Myers & Kuntz, 1967, sensu Durette-Desset (1976). Relationships between Old World Brevistriatinae and Western Hemisphere Pudicinae need to be reexamined. Acknowledgments I wish to express thanks to Drs. Alfred L. Gardner and Juhani Ojasti for providing useful suggestions on the original manu- script. Literature Cited Baylis, H. A. 1947. Some roundworms and flatworms from the West Indies and Surinam. I. Nema- todes and Acanthocephala. — Journal of the Lin- nean Society, Zoology 41:394—405. Chandler, A.C. 1942. Helminths of tree squirrels in southeast Texas.—Journal of Parasitology 28: 135-140. Durette-Desset, M. C. 1970. Nematodes héligmo- somes d’Ameérique du Sud. VIII. Description de six nouvelles espéces parasites de Cricétidés. — Bulletin du Muséum National d’Histoire Na- turelle, Paris, Série 2, 42:730-744. 1971. Essai de classification de Nematodes Héligmosomes. Corrélation avec la paléobi- ogéographie des hétes.— Mémoires du Muséum National d’Histoire Naturelle, Paris, Série A, Zoologie 49:1—-126. 1976. Brevistriatinae (Nematoda: Heligmo- somidae). IV. Conclusions phylétiques et sys- tematiques.— Bulletin du Muséum National d@ Histoire Naturelle, Paris, Série 3, Zoologie 270: 698-720. 1985. Trichostrongyloid nematodes and their vertebrate hosts: reconstruction of the phylog- eny of a parasitic group.— Advances in Parasi- tology 24:239-306. ——, & A. G. Chabaud. 1977. Essai de classifi- cation des Nématodes Trichostrongyloidea.— Annales de Parasitologie Humaine et Comparée 52:539-558. , & 1981. Nouvel essai de classifica- tion des Nématodes Trichostrongyloidea. — An- nales de Parasitologie Humaine et Comparée 56:297-312. , & J.-L. Justine. 1992. A cladistic analysis of the genera in the subfamily Pudicinae (Nema- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON toda, Trichostrongyloidea, Heligmonellidae). — Systematic Parasitology (in press). Guerrero, R. 1982. Trichostrongyloidea (Nematoda) parasitos de mamiferos silvestres de Venezuela. I. Los géneros Bradypostrongylus Price, 1928, Longistriata Schulz, 1926 y Durettestrongylus n. gen.—Acta Biologica Venezuelica 11:11—131. 1984. Nematoda: Trichostrongyloidea par- asites of Venezuelan wild mammals. III. The genus Vexillata Travassos, 1937.—Annales de Parasitologie Humaine et Comparée 59:253- 261. Harkema, R. 1936. The parasites of some North Car- olina rodents.— Ecological Monographs 6:151— 232. Hartenberger, J.-L. 1985. The order Rodentia: major questions on their evolutionary origin, relation- ships and suprafamilial systematics. Pp. 1—33 in W. P. Luckett & J.-L. Hartenberger, eds., Evolutionary relationships among rodents. A multidisciplinary analysis. NATO ASI Series, Series A: Life Sciences Volume 92. Plenum Press, New York. Lent, H., & J. F. T. Freitas. 1938. Pesquisas helmin- thologicas realisadas no Estado do Para. IV. Tri- chostrongyloideos de mamiferos.—Memorias do Instituto Oswaldo Cruz 33:363-—380. Moore, J. C. 1961. The spread of existing diurnal squirrels across the Bering and Panamanian land bridges.—American Museum Novitates 2044: 1-26. Price, E. W. 1929. Two new nematode worms from rodents. — Proceedings of the U.S. National Mu- seum 74:1—5 + 2 pls. Reig, O. A. 1981. Teoria del origen y desarrollo de la fauna de mamiferos de America del Sur.— Monographiae Naturae del Museo Municipal de Ciencias Naturales Lorenzo Scaglia 1:1—162. Schmidt, G. D., B. J. Myers, & R. E. Kuntz. 1967. Nematode parasites of Oceanica. I. Brevistriata sundasciuri sp. n. and Calypsostrongylus ogdeni gen. et sp. n. (Heligmosomatidae: Longistriati- nae) from squirrels of Palawan and Taiwan.— Journal of Parasitology 53:613-617. Webb, S. D. 1985. Late Cenozoic mammal dispersals between the Americas. Pp. 357—386 in F. G. Stehli & S. D. Webb, eds., The great American biotic interchange. Plenum Press, New York. Instituto de Zoologia Tropical, Facultad de Ciencias, Universidad Central de Vene- zuela, P.O. Box 47051, Caracas 1041A, Venezuela. PROC. BIOL. SOC. WASH. 107(1), 1994, pp. 185-192 DESCRIPTION OF THE NEW GENUS ALLOPATHES (CNIDARIA: ANTIPATHARIA) AND ITS TYPE SPECIES CIRRIPATHES DESBONNI Dennis M. Opresko and Stephen D. Cairns Abstract.—The rarely collected species originally described as Cirripathes desbonni is redescribed based on two specimens. A neotype is designated be- cause the type is presumed to be lost. A new genus, A//opathes, is proposed for this species based on its unusual colony form: numerous elongate, mostly unbranched stems originating from a short, trunk-like base. Antipathes robil- lardi Bell is also placed in this genus, which gives the genus a range of western Atlantic and Mauritius at 129-161 m. In September, 1989 the second author participated in a cruise of the Johnson-Sea- Link I research submersible (stations 2582— 2595) off the southeastern coast of Louisi- ana. Eight species of Antipatharia were ob- served, photographed, and collected (Cairns et al. 1994), one of which was Allopathes desbonni. This species is composed of nu- merous long, slender, mostly unbranched stems arising from a stout, trunk-like base. We consider the distinctive nature of the corallum of this species and one other, A. robillardi, to justify establishing a new ge- nus, and include a redescription of the spe- cies. Allopathes, new genus Diagnosis.—Corallum consisting of nu- merous elongate stems arising from a short trunk-like base. Stems generally straight but may be slightly coiled at their distal end; mostly unbranched, but occasionally with first and second-order branches arising from the lowermost parts of the stems. Spines typically arranged in verticils around cir- cumference of axis. Polyps in a single row along length of axis. Discussion.—As was noted by Duchas- saing & Michelotti (1864) the generic affin- ites of Cirripathes (=Allopathes) desbonni are not clearly defined. The individual stems resemble those of Cirrhipathes and Stichop- athes, and the arrangement of the polyps in a single row would tend to associate the species more with Stichopathes. The latter was treated as a subgenus of Cirrhipathes by some (e.g., Van Pesch 1914). However, the morphology of the spines of Allopathes, and its tendency to develop long branches, would suggest an affinity with Antipathes verticillata (Brook 1889). Placing Antipathes desbonni in Stichopathes or Cirrhipathes would substantially alter the major diag- nostic character of those genera, i.e., the un- branched growth form of the corallum, which we do not believe to be justified. Likewise, placing A. desbonni in Antipathes would diminish the significance of the unique morphology of the corallum. Al- though the genus Antipathes currently is a heterogeneous assemblage of species, it is our view that natural groupings of species tend to center around distinct types of skel- etal morphology which, when evaluated in association with characteristics of the spines and polyps, are likely to define generic or subgeneric taxa. Therefore, we consider it appropriate to establish a new genus for des- bonni and also include in it Antipathes robil- lardi Bell, 1891. Etymology. —From Greek allos, other + 186 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. stems, x 0.75; b, Section of stem with polyps, <6. pathes, second component of Antipathes, in allusion to its taxonomic relationship. Gen- der: feminine. Type species. — Cirripathes desbonni, here designated. Cirripathes is an incorrect spell- ing of Cirrhipathes. Allopathes desbonni (Duchassaing & Michelotti, 1864), new combination Figs. 1-4 Cirripathes Desbonni Duchassaing & Mi- chelotti, 1864:142. Not Antipathes (Cirrhipathes) Desbonnii. — Pourtalés, 1874:46.—1878:209 (=Stich- opathes pourtalesi Brook). Antipathes (Cirrhipathes) Desbonni. — Pourtalés, 1880:114, pl. iii, figs. 6-7. Stichopathes? desbonni. — Brook, 1889:92. Material examined. —Gulf of Mexico, off southeastern Louisiana, 27°44.62'N, 91°07.9'W, 129-144 m, Johnson-Sea-Link T Stn 2585, USNM 88327, neotype.—Off Allopathes desbonni, neotype (USNM 88327). a, Basal section of corallum showing the origin of the Montserrat, Lesser Antilles, Blake Stn 155, 88 fm (=161 m), one specimen in the Mu- seum of Comparative Zoology, Harvard University. Diagnosis. —Corallum large, about 1 m in height, with numerous mostly unbranched stems arising from a short trunk-like base (Fig. la); stems generally straight and stiff, but flexible and somewhat coiled near apex; occasionally branched near base. Spines typically conical, acute, and usually 0.10- 0.14 mm (but up to 0.20 mm) from mid- point of base to apex; with cone-shaped tu- bercles on upper two-thirds or more of sur- face. Spines arranged in verticils of varying regularity and in longitudinal rows with 3 or 4 spines per millimeter in each row. Pol- yps arranged in one row along stems and branches; 1.0—1.2 mm in transverse diam- eter (from proximal side of proximal lateral tentacles to distal side of distal lateral ten- tacles); interpolypar space about 0.4—0.5 mm; from 5 to 7 polyps per centimeter (Fig. 1b). VOLUME 107, NUMBER 1 i 4 Ml i ef x 1 Fig. 2. Allopathes desbonni, neotype (USNM 88327). a, b, Section of stem 0.5 mm in diameter, showing arrangement of spines, stereo pair, x 59; c, Section of stem 1.5 mm in diameter, x 25; d, Section of stem 0.85 mm in diameter with bifid spines, x 46. 188 Description.—The neotype consists of a cluster of about 40 stems attached to a 4cm long “‘trunk,” as well as several dozen de- tached pieces. This specimen was only part of a much larger colony, the remainder of which was not collected. Individual stems reach a maximum length of about 90 cm and are 0.5—2.3 mm in diameter at their base (average 1.47 mm, n = 24). Stems 46, 53 and 75 cm long have basal diameters of 1.1, 2.3, and 1.5 mm, respectively. All stems arise directly from the base (Fig. la) and most are unbranched; however, a few have a single branch that originates not more than 1 centimeter above the basal end of the stem. The branches are unbranched and can be as long as the stem from which they arise. The axial diameter of a stem or branch decreases regularly from base to apex, e.g., for one 52- cm long stem, the diameter is 2.3 mm at the base, 1.7 mm at a height of 10 cm, 1.1 mm at 30 cm, 0.7 mm at 40 cm, and 0.3 mm at its distal end. The stems and branches of the corallum radiate upward from the base. They are rel- atively straight and stiff for most of their length; however, near the apex (1.e., upper 10 cm or more) they curve and even form a loose coil. This occurs regardless of the overall length of the stem or branch, and is associated with an increased flexibility of the axis due to a very thin sclerenchymal layer and a relatively wide central axial ca- nal (0.20—0.36 mm). Because of these fac- tors the tips of the stems and branches col- lapse when dried. Axial spines (Figs. 2, 3, 4a—c) are arranged in longitudinal rows and also in verticils, although the regularity of the verticils varies considerably from point to point. Six or sev- en longitudinal rows of spines can be seen in viewing one side of the axis and 13 rows were counted around the entire circumfer- ence of one segment of axis having a di- ameter of 1.6 mm. The distance between the spines in each row varies on different parts of the axis and ranges from 0.24 to 0.40 mm; however, in general, there are PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON usually 3 or 4 spines per millimeter in each row. Spines are generally uniform in size at any given point on the corallum, though they vary in size and appearance along the length of the axis, as well as around the circum- ference. Spines on the polyp-side of the axis are generally larger than those on the ab- polypar side. The spines also increase in size from the distal end of a stem or branch to the basal end; however, the rate of increase is not uniform from one stem or branch to another. About 1 cm from the distal end where the axial diameter is about 0.2 mm, the spines are quite small, 0.04—0.06 mm from the tip to the center of the base, some- what compressed laterally, triangular in shape, and have only a few small conical protuberances on their surface. With in- creasing axial diameter the spines become larger; i.e., 0.07-0.09 mm (axial diameter about 0.3 mm), 0.10-0.12 mm (axial di- ameter 0.4-0.7 mm, Fig. 3a), 0.10—0.16 mm (axial diameter 0.7-1.0 mm, Fig. 3b), and 0.12—0.20 mm (axial diameter over 1.0 mm). Overall, the typical polypar spine is 0.10- 0.14 mm tall, conical in shape, with an acute to slightly rounded apex, and with cone- shaped tubercles scattered over the surface. The corresponding abpolypar spines are generally 0.02 to 0.06 mm smaller and usu- ally have fewer surface tubercles. For ex- ample, at the midpoint of one 30-cm long stem (axial diameter about 0.9 mm) the pol- ypar spines measure 0.10—0.12 mm and the abpolypar spines 0.08 mm. The spines on the lowermost portion of each stem (usually within 5 to 10 cm of the basal end and re- gardless of total length of the stem) differ from the typical spines in being almost smooth-surfaced (Figs. 2c, 3c). These spines are large (usually 0.16-0.20 mm), have a sharp apex and tend to be directed distally. In addition, on some stems, and particularly where the axial diameter is 1.0 mm or more, the polypar spines can become rather wide and blunt with numerous surface tubercles (Figs. 4b, c). The location and abundance VOLUME 107, NUMBER 1 189 es fe Se See ae OS a Fig. 3. Allopathes desbonni, neotype (USNM 88327). a, Spine from section of stem about 0.5 mm diameter, x 365; b, Spine from section of stem about 0.8 mm in diameter, VA Vy / = / W/, I 7 RECRUITMENT / RE / Zz vy, CRUITMENT Li = ie a is — si ~ N YEAR 1 YEAR 2 YEAR 3 B. ITEROPAROUS BREEDER A BREED & BREED & BREED & 3 s3 Vi 3 Vi VA Vi, 5 fe YA Ue s Ji VA 7, 2 YA WA Le ha eal N YEAR 1 YEAR 2 YEAR 3 C. SEMI-CONTINUOUS BREEDER () CBREED = Se ee eee SEXUAL MATURITY YEAR 1 Fig. 4. Metazoan breeding patterns. A) semelparous breeder; B) iteroparous breeder; C) semi-continuous breeder. VOLUME 107, NUMBER 1 ing oocytes is often positively correlated with the rate of oocyte growth. Semelparous and annual iteroparous species generally show low levels of endocytotic activity during vi- tellogenesis because there is no premium on rapid yolk precursor incorporation and yolk production. For example, endocytotic ac- tivity is minimal in most iteroparous echi- noderms based on ultrastructural studies of asteroids, echinoids, crinoids, ophiuroids, and holothuroids (see review of Eckelbarger & Young 1992). However, the lack of sig- nificant endocytotic activity in growing 0o- cytes does not suggest that exogenous pre- cursors are not being incorporated but rather that they are being incorporated at a slow rate. The oocytes of the asteroid Asterias rubens are known to incorporate high mo- lecular weight exogenous molecules despite the absence of ultrastructural documenta- tion of endocytotic activity (Beijnick et al. 1984). Low endocytotic activity in growing oocytes may merely indicate a rate of pre- cursor incorporation that is not easily de- tectable by ultrastructural methods. Oogen- esis extends for a period of more than a year in the semelparous nereid polychaete Nereis virens (Brafield & Chapman 1967), and 7— 8 mo in UN. grubei (Schroeder 1968). The oocytes of both species show minimal en- docytotic activity even though a female- specific yolk protein is known to be pro- duced by coelomocytes and incorporated slowly over the long growth period (Fischer & Dhainaut 1985). Likewise, other semel- parous nereids such as Nereis diversicolor, N. pelagica, Perinereis cultrifera, and Platy- nereis dumerilii have long periods of vitel- logenesis and have little or no detectable endocytotic activity in the oocytes (re- viewed in Fischer & Rabien 1985). Other polychaetes with slowly growing oocytes show a similar pattern of low endocytotic activity during vitellogenesis (Eckelbarger 1983, 1986). In contrast, oogenesis occurs in less than three months in the nereid Neanthes arenaceodentata (Davis 1969) and ultrastructural studies of their faster devel- 205 oping oocytes have shown high levels of endocytosis during vitellogenesis. Other polychaetes such as Phragmatopoma lapi- dosa, Streblospio benedicti, and Polydora ligni undergo vitellogensis in only a few days and show high levels of endocytotic activity during this period (Eckelbarger 1979, 1980, 1983, 1986). Similarly, oocytes of the semaeostome jellyfish, Aurelia aurita, an iteroparous neritic species that undergoes rapid egg production, show intense endo- cytotic activity during vitellogenesis (Eckel- barger & Larson 1988). In insects, including most of the primitive Apterygota, vitello- genesis is a heterosynthetic process in which the fat body produces yolk protein precur- sors which are secreted into the hemo- lymph, and rapidly sequestered by the developing oocytes through a receptor-me- diated process (Hagedorn & Kunkel 1979, Raikhel & Dhadialla 1992). Ovarian nurse cells are frequently asso- ciated with rapidly developing oocytes, no- tably in the insects. Vitellogenesis occurs in only 25 h in the mosquito (Roth & Porter 1964) and about 18 h in the fruit fly, Dro- sophila (Brownes 1982), species having the more complex meroistic ovary containing nurse cell complexes. In contrast, in the more primitive panoistic insect ovary lacking nurse cells, vitellogenesis is relatively long, for instance, 3—6 mo in the cricket (in Da- vidson 1986). Ovarian nurse cells are also responsible for rapid egg growth in cteno- phores, rotifers, annelids, and crustaceans (Davidson 1986). In the Ctenophora, the majority of biosynthetic activity during oo- genesis appears to be performed by the large population of nurse cells that establish a syncytium with the oocyte (Pianka 1974). Yolk bodies are formed in the ooplasm by the fusion of pre-yolk bodies originating from the nurse cells. Vitellogenesis in cteno- phores is believed to occur in only about two days in Beroe gracilis and Pleurobrachia pileus (Greve 1970). In the rotifer As- planchna there is a 100-fold increase in 00- cyte volume within 4-6 h resulting from the 206 abrupt transfer of cytoplasmic materials to the oocyte from the syncytial nurse cells of the vitellarium (Bentfield 1971). Acceler- ated egg production occurs in the small- bodied polychaete, Ophryotrocha puerilis with the assistance ofa single polyploid nurse cell (Ruthmann 1964). Among hydrozoans, nurse cells-assisted vitellogenesis reduces oogenesis to about four days in Hydra car- nea (Honegger 1981) and less than eight days in Tubularia crocea (Mackie 1966). Follicle cells are perhaps the most com- mon accessory cell found in invertebrate Ovaries and there are some examples in which their function(s) is directly correlated with rapid egg production. The opportunis- tic polychaete Capitella jonesi (Eckelbarger & Grassle 1982) undergoes frequent and rapid egg production with the aid of hyper- trophic follicle cells that contribute to the production of yolk precursors during vitel- logenesis. Follicle cells also have been shown to be the source of yolk precursors in the rapidly growing eggs of some bryozoans (Reed 1988, 1991). In chaetognaths, egg production can occur at daily intervals and may be facilitated by the endocytotic in- corporation of precursors produced by fol- licle-like accessory cells (Shinn 1992). The adaptive significance of semelparity and iteroparity has been debated extensive- ly (see Grahame & Branch 1985); theory suggests that these respective life history patterns have evolved to attune the life style of the organism to its environment (Stearns 1976). Often overlooked is the fact that the duration of vitellogenesis is strikingly dif- ferent between semelparous and iteropa- rous species. As reviewed above, different mechanisms of yolk synthesis have evolved in these organisms, including those that have endowed some species with the ability to rapidly accumulate yolk reserves. Rather than categorizing species as semelparous or itereroparous, it is more appropriate to view them on a continuum ranging from rela- tively fast egg producers characterized by heterosynthetic mechanisms designed to ac- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON celerate yolk production, to relatively slow egg producers characterized by slower “mixed”? mechanisms of yolk synthesis. As the period of vitellogenesis shortens (and the potential rate of egg production increas- es), the adoption of heterosynthetic mech- anisms assisting the egg in yolk production increases. In the examples presented above, these mechanisms range from the direct transport of vitellogenin to the ovary through the blood, hemolymph or coelomic fluid, to trophic support from the biosynthetic ac- tivity of follicle cells and/or nurse cells (Fig. 1). The rapid incorporation of yolk precur- sors by receptor-mediated endocytosis is a common mechanism observed in species having rapidly growing oocytes. Fast egg producers and accompanying heterosyn- thetic mechanisms of yolk production are generally not found among annual spawners because selective pressures do not favor ac- celerated egg growth. In these species, “mixed” vitellogenic mechanisms have been adopted in which endocytotic incorporation of yolk precursors plays a lesser role during oocyte growth. Long-lived species have adopted mechanisms for slow egg produc- tion consistent with seasonal, continuous, or predictable food supplies and relatively stable environments that characterize shal- low water populations at temperate lati- tudes. Unstable environments with unpre- dictable food supplies, on the other hand, favor fast egg producers (e.g., opportunistic species) and the accompanying evolution of accelerated mechanisms of yolk synthesis. Energy Allocation and Oogenesis Among Slow vs. Fast Egg Producers The manner in which organisms allocate energy among growth, maintenance and re- production during each reproductive period is critical to their life history and will differ between species depending upon the vitel- logenic mechanism employed. Reproduc- tion and somatic growth are generally con- sidered antagonistic and relatively long-lived VOLUME 107, NUMBER 1 semelparous species (e.g., Nereis virens) of- ten allocate some energy to both, whereas many species that breed semi-continuously (e.g., Capitella and Streblospio) appear to convert a large proportion of their energy directly into reproduction. In crustaceans, some species can sustain simultaneous go- nadal maturation and somatic growth (in- cluding molting) while in others, growth is sacrificed at the expense of reproduction (Harrison 1990). Storage of energy reserves in the somatic tissues for later use during oogenesis is common in semelparous spe- cies. Many polychaetes, for example, store glycogen and lipid in the gut, peritoneum and coelomic cells for later transfer to the oocytes (reviewed in Eckelbarger 1983). Lipids are commonly stored by copepods from higher latitudes for later use in repro- duction (Sargent & Henderson 1986). In nereid polychaetes and cephalopods, ma- ternal muscle tissue is catabolized during vitellogenesis and is gradually incorporated from the coelomic fluid and the circulatory system, respectively, by the oocytes. Some species show an immediate reproductive re- sponse to environmental stress or an inter- ruption of food supply. For example, in some fast-breeding polychaetes, the ovarian ac- cessory cells are involved in the resorption of unspawned eggs presumably as a means of energy recycling and they also abruptly resorb oocytes in response to food depri- vation or other environmental stresses which may create an energy deficit (Eckelbarger 1986). Regeneration also may impose en- ergetic demands on the female by altering nutrient allocation during sexual reproduc- tion. In some opportunistic polychaetes (e.g., Capitella), regenerating females show a de- lay in maturation and a marked decline in fecundity as compared to non-regenerating worms (Hill & Grassle 1981, Hill etal. 1982). In semelparous nereid polychaetes, regen- eration is progressively inhibited as females approach reproductive maturity, indicating a shift in allocation of nutrient resources toward oogenesis (Hofmann 1976). Energy 207 allocation, therefore, differs significantly be- tween fast and slow egg-producing species due to different physiological priorities (Fig. 5). Reproductive Responses to Food Levels Nutrition, vitellogenic mechanisms, and egg production are inextricably linked but species show significantly different re- sponses to spatial and temporal variations in food quality and quantity in the marine environment. For example, some species show abrupt reproductive responses to food levels while others do not. These differences may be related to the type of vitellogenic mechanism employed during oogenesis. Reproductive responses to variations in food levels have been extensively documented in pelagic invertebrates (Checkley 1980) yet are poorly understood in benthic species (Levin & Creed 1986). Pelagic species show vari- able rates of food conversion into egg pro- duction and reproductive effort has been observed to be very sensitive to food level (Tester & Turner 1990, Razouls et al. 1991). The semaeostome scyphozoan, Aurelia au- rita, maintains a high rate of egg production when fed but quickly resorbs its oocytes when deprived of food for as little as two days (Eckelbarger & Larson 1988 and un- publ.). Oogenesis is very short in this species and oocytes show high levels of endocytotic activity during vitellogenesis. In copepods, egg production for some species follows food ingestion within hours whereas, for others, the response may take days (Tester & Tur- ner 1990). Vitellogenesis in Calanus pacific- us 1s very rapid after feeding (Runge 1984) with labeled food appearing in oocytes in less than 8 h after food ingestion (Marshall & Orr 1956). Lower food levels slow the rate of yolk synthesis so that spawning in- tervals are longer and egg production rates are lower (Runge 1984). This species is adapted for rapidly converting nutrients to yolk and will have a short term advantage Over species utilizing slower pathways for 208 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON rctiaion J» [respon] —> | [2h - —_> vitellogenesis => SLOW EGG PRODUCERS somatic storage growth regeneration cell aeesion] —> [transport] > => | vilellogenesis| => Food FAST EGG PRODUCERS Source SS TIME Fig. 5. Energy flow pathways from food source to spawned egg in slow and fast egg producers. nutrient incorporation. Although the ultra- structural features of oogenesis have not been described in C. pacificus, they have been reported in Labidocera aestiva, anoth- er species that produces eggs rapidly. The oocytes of this species rapidly incorporate exogenous precursors endocytotically from the surrounding hemolymph (Blades-Eck- elbarger & Youngbluth 1984). In contrast, the oceanic subantarctic copepod, Neocala- nus tonsa, exhibits seasonal changes in feed- ing behavior, egg production response, and organic composition. Winter females dwell- ing in mesopelagic depths can release eggs in the absence of particulate food due to lipid reserves that allow recruitment to be decoupled from or out of phase with pri- mary production in surface waters. Spring females, on the other hand, have limited lipid reserves and must obtain exogenous sources of nutrition for egg production. They are, therefore, directly coupled to primary production for recruitment (Ohman 1987). There are many examples of species that respond to feeding or starvation by abruptly switching egg production on or off. For ex- ample, while most sea anemones are annual breeders, store food reserves, have extended gametogenic periods, and show little im- mediate response to food levels (Jennison 1979), the small, estuarine anemone WNe- matostella vectensis, shows dramatic re- sponses to food levels by producing eggs at 8-day intervals as long as feeding continues (Hand & Uhlinger 1992). The remarkable ability of some benthic species to translate increased food supply into accelerated egg production may underlie opportunistic dy- namics in shallow water macrobenthos (Levin 1986). Populations of Capitella sp. I, for example, are extremely sensitive to changes in food supply and can adjust their VOLUME 107, NUMBER 1 reproductive rates accordingly, an impor- tant determinant of opportunistic popula- tion dynamics (Marsh et al. 1989). The cap- itellid ovary is capable of rapidly producing yolk precursors in response to food intake with the aid of proteosynthetic follicle cells (Eckelbarger & Grassle 1982). However, even in capitellids, significant variation is observed in the way food is utilized in egg production (Eckelbarger 1986). Sibling spe- cies of Capitella occupy similar habitats and appear to have similar methods of vitello- genesis, but they show wide variation in egg volume, brood size, and the types of yolk materials deposited in the eggs (Eckelbarger & Grassle 1983): Despite their close genetic relationship and similar habitats, food is utilized differently by the ovary. In organic enrichment studies of Streblospio benedicti, a polychaete exhibiting both planktotrophic and lecithotrophic modes of larval devel- opment, females exhibiting lecithotrophy showed an increase in egg production while those with the planktotrophic mode showed no increase (Levin & Creed 1986). The eggs of S. benedicti contain two kinds of yolk bodies produced by separate autosynthetic and heterosynthetic pathways (Eckelbarger 1980) but the lecithotrophic egg contains a higher proportion of heterosynthetically- derived yolk bodies (Eckelbarger 1986). The circulatory system of S. benedicti is capable of rapidly transporting yolk precursors to the ovary following feeding (Eckelbarger 1980). It has been suggested that this in- crease in egg production by females exhib- iting lecithotrophic development could be due to a greater availability of heterosyn- thetic yolk precursors derived from the en- hanced food supply (Levin & Creed 1986). The fact that reproductive responses to food varies (e.g., the existence of opportu- nistic and non-opportunistic species) sug- gests that there are complex interrelation- ships between trophic dynamics and reproductive biology that are poorly under- stood. However, recent comparative studies of gut morphology and kinematics of di- 209 gestive processing indicate that species op- erate under different digestive constraints related to their respective feeding ecologies (Penry & Jumars 1990). For example in polychaetes, the guts of carnivorous species differ morphologically and functionally from those of deposit feeding species (Penry & Jumars 1990) with carnivorous species havy- ing significantly less gut volume than de- posit-feeding species. The diet of carnivo- rous species also contains greater proportions of high quality foods (higher protein, lower in ratios of carbon to nitro- gen). Food quality rather than quantity is probably limiting for deposit feeders and the diets of deep-sea deposit feeders are be- lieved to be, on average, of lower quality than nearshore and shelf deposit feeders (Penry & Jumars 1990). Food quality also influences digestion time. For example, di- atoms and other labile foods are digested rapidly and absorbed efficiently while “‘sea- weed detritus” requires long residence times. “Lignin” and other refractory organic mat- ter is never appreciably absorbed (Lopez & Levington 1987). Many deposit feeding, op- portunistic species such as those in Capi- tella, Armandia, Tharyx, Cirratulus, Chae- tozone, Paraprionospio, Levinsenia, and Streblospio have simple guts that may be relatively inexpensive to construct and maintain but, due to short throughput times, may limit them to inhabiting areas of or- ganic enrichment in order to exploit higher quality foods to support their high rate of reproduction (Penry & Jumars 1990). Species apparently differ with respect to their trophic transfer efficiency (net pro- duction/food supplied) (Tenore 1983), so it cannot be assumed that similar foods will be energetically processed to support egg production in the same way in different spe- cies. In crustaceans, for example, optimal dietary balance and the efficiency of utili- zation of dietary carbohydrates, lipids, and proteins vary widely among species (Har- rison 1990). In polychaetes, food quality and particularly nitrogen content appear to reg- 210 ulate growth and reproduction (Gremare et al. 1988). Juvenile growth has been shown to be especially sensitive to food quality with different growth responses being observed in relation to spring vs. summer detritus (Marsh et al. 1989). When considering pop- ulation dynamics, interspecific dietary re- quirements and digestive constraints must be factored because they ultimately influ- ence reproduction and growth. If species show different reproductive responses to nutrients, then nutrient levels in different habitats will present strong selective pres- sures favoring one species over another. For example, organic enrichment can have a dramatic effect on community structure and population biology of shallow-water organ- isms by decreasing diversity and increasing the resulting dominance by rapidly-repro- ducing, opportunistic species (Levin et al. 1993). Individual responses to elevated food quantity and/or quality may include in- creases in fecundity, early age at first repro- duction, and modified egg composition (Levin & Creed 1986, Marsh et al. 1989, Qian & Chia 1991). Notable respondents to these enrichments include Capitella spp.., Polydora ligni and Streblospio benedicti, which brood their young, have very high reproductive rates (Levin & Huggett 1990) and utilize heterosynthetic mechanisms of yolk acquisition (Eckelbarger 1983). Vitellogenic Mechanisms and Habitat Instability Some species have adopted mechanisms for rapid egg production that appear to be less related to food levels than to a response to inherent habitat instability. For example, life history studies of barnacles have dem- onstrated that species behave opportunis- tically to exploit changing conditions in rig- orous, unpredictable environments by rapidly increasing yolk accumulation in re- sponse to food levels (Hines 1979). The in- tertidal, reef-building polychaete Phrag- matopoma lapidosa is an opportinistic species with early age of first reproduction, PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON rapid and continuous egg production (Eck- elbarger 1976) and unpredictable mortality due to storms and heavy wave action. In- dividuals are sexually mature soon after set- tling and are capable of spawning continu- ously. Oogenesis occurs in only two days and involves the transport of yolk precur- sors directly to the ovary via the circulatory system (Eckelbarger 1979) in a manner sim- ilar to that reported in the polychaete Streb- lospio (Eckelbarger 1980). Hydrothermal vent communities also represent ephemeral habitats whose age may be as short as 1-10 years (see Scheltema 1994). The fauna as- sociated with these vents is largely unique and must mature and reproduce within a relatively short period of time so new active regions can be colonized. Recent studies of ovarian morphology and oogenesis in the vestimintiferan, Riftia sp. indicate that oo- cytes develop rapidly in close association with blood vessels (Gardiner et al. 1992) in a manner similar to some opportunistic polychaetes (e.g., Streblospio). Studies of vent-associated molluscs have shown that many species demonstrate life history traits that appear to be adaptive in this severe environment. Notable among these traits are rapid growth and continuous reproduc- tion, which are probably related to a nutri- ent source that is relatively constant (Gus- tafson & Lutz 1994). Iteroparous molluscs that live longer and reproduce either inter- mittently or continuously throughout their reproductive life, tend to be found in more stable deep-sea environments (Scheltema 1994). Based on comparative studies of vi- tellogenesis, one would predict that unstable or unpredictable environments would select for species that mature early and reproduce rapidly using heterosynthetic mechanisms of egg production. Vitellogenic Mechanisms, Food and Habitat Selection If species convert food into egg produc- tion at different rates, one would expect that community structure would be influenced VOLUME 107, NUMBER 1 by temporal variation in food levels. Com- munity level responses to variable organic input are much better known for near-shore populations than for the deep sea (Jumars & Wheatfcroft 1989). There has been a re- surgent interest in benthic responses to tem- poral input of organic matter in the deep sea because of the growing realization that deep-sea organisms show a much higher fre- quency of seasonal reproduction than pre- viously imagined, perhaps in response to environmental cues (Tyler et al. 1994). The relatively recent discovery of episodic in- puts of significant organic food falls, plant material, and seasonal deposition of fresh phytodetritus on the deep-sea floor (see Scheltema 1994, for review) has stimulated great interest in determining whether ben- thic organisms show a reproductive re- sponse to these cues (Tyler et al. 1994). The deep-sea, wood-boring bivalve, Xylophaga sp., for example, shows an opportunistic life history by undergoing early sexual maturity, rapid egg reproduction, high population densities, and the ability to utilize a highly transient habitat (wood) similar to shallow water opportunistic species (Turner 1973). However, most deep-sea molluscs show no obvious coupling between seasonal peri- odicity of reproduction and primary pro- ductivity occurring at the ocean surface (Scheltema 1994). Scheltema (1994) sug- gests that seasonal differences in organic particulate flux may be largely irrelevant to many deep-sea molluscs except for filter feeders (e.g., Pectinidae), and possibly de- posit-feeders that feed on the surface (e.g., protobranch bivalves). Omnivores, scav- engers, and predators (1.e., majority of abys- sal gastropods) may show little response to organic fluxes. Scheltema (1994) further suggests that other molluscs would only be secondarily affected. They include Calyp- togena sp., a hydrothermal vent species har- boring chemosynthetic bacteria, predatory or parasitic species affected directly by the relative abundance of their prey or host, and species that can derive much of their nu- trition from dissolved organic materials in 211 the pore-water of deep-sea sediments (Southward & Southward 1982). In a study of species pairs of deep-sea organisms, one being a quasi-continuous breeder and the other a seasonal breeder, Tyler and col- leagues (1994) studied the effects of seasonal phytodetrital pulses on reproduction. Their results indicated that all the quasi-contin- uous breeders conformed to the pattern pre- dicted by showing no seasonal variation in their diet while in seasonal breeders, the pattern of reproduction did not vary with the supply of phytodetritus from the sur- face. Although seasonal reproduction has been widely reported among deep-sea in- vertebrates, control of these cycles is not necessarily related to these surface-derived fluxes (Tyler et al. 1994). Do species show different reproductive responses to differing temporal patterns of organic input? Some regions of the deep sea receive a steady rain of organic material while others receive seasonal pulses result- ing from surface plankton blooms (reviewed in Gage & Tyler 1991). In a recent study of energy metabolism by deep-sea benthic fo- raminifera, it was determined that at least two survival strategies have evolved in this group that reflect individual physiological responses to this variable input of food. Based on turnover times of ATP, some sus- pension feeding species respond to condi- tions in which they receive a steady rain of organic particles by showing a reduced ATP turnover rate, while some scavenging spe- cies respond rapidly to sudden, seasonal nu- trient inputs with large seasonal ATP turn- over rates (Linke 1992). This study supports the notion that adaptation to nutritional conditions rather than mass properties or physical conditions are the sole controls over foraminiferal distribution. The author sug- gested that organisms will evolve different survival strategies depending upon whether they live in environments receiving season- al food input, such as epibenthic species liv- ing in temperate and Arctic latitudes, or if they are species exposed to little seasonality in food supply, such as in oligotrophic 212 regions of the ocean (Linke 1992). The downward vertical flux of surface phyto- detritus has been viewed as a potential con- trolling factor in gametogenic cycles and re- cruitment of seasonally breeding species (Tyler et al. 1994). It remains an enigma that two or more sympatric species can have different repro- ductive patterns under the same environ- mental conditions (Tyler et al. 1994). How- ever, it is relatively common to find congeners in similar habitats with com- pletely different life histories even though the causes are often difficult to determine (Spight 1979). This situation implies that reproductive patterns are attuning the or- ganism to something other than the habitat itself. When viewed from the perspective of vitellogenesis, these differences could be ex- plained by interspecific differences in vitel- logenic mechanisms and the manner of nu- trient cycling during oogenesis. Since species have very different capacities to respond to organic enrichment it should not be sur- prising that even closely related species will show different responses to the same food levels. Animals of similar feeding types might exploit different organic components in different ways and at varying rates within the same habitat. Thus organic pulses of surface phytodetritus may be utilized in dif- ferent ways by individual deposit feeders (or other animals) resulting in different life his- tory responses. Studies of deep sea com- munities suggest that the strategy of se- questering labile food material by rapid assimilation and population growth is most effective for small organisms with rapid growth rates. This strategy is documented only for bacteria, microfaunal, and meio- faunal populations whereas the evidence for effectively sequestering pulses of raining or- ganic matter for common deposit feeders of intermediate size (between echiurans and meiofauna or the macro-infauna) is very circumstantial (Jumars et al. 1990). Since organic windfalls to the deep sea bottom are PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON often unpredictable, it might explain why so many deep-sea populations contain in- dividuals with only a few mature eggs at any given time (Rokop 1974, Gage & Tyler 1991) and the observation of a few juveniles pres- ent on average at any time (Grassle & Sand- ers 1973). Assuming that juvenile survival is highly variable, it would be useful for the adult to have many reproductive episodes with little energy and offspring invested in each (Jumars et al. 1990). As Jumars and his colleagues point out: “The relevant and unresolved issue for macrofaunal surface are dominant, less important but still sig- nificant, or unimportant sources of the mat- ter and energy shunted into production of this group of animals” (Jumars et al. 1990). Echinoderms are among the most abun- dant macrofaunal organisms in the deep sea and their reproductive biology is better doc- umented than for most invertebrate groups (Gage & Tyler 1991). Continuous repro- duction in deep sea echinoderms is the most common pattern observed in the group but it involves the production of relatively few large eggs (Gage & Tyler 1991). Rapid and frequent egg production is rare in echino- derms, with the notable exception of the shallow water echinoid Diadema (Levitan 1988). Specialized heterosynthetic mecha- nisms for rapid yolk production have not been documented in the phylum and, thus far, vitellogenic mechanisms are highly con- served and of the ““mixed” variety (Eckel- barger & Young 1992). Further, most echi- noderms do not demonstrate abrupt reproductive responses to food levels. One would predict, therefore, that echinoderms would be well adapted to food conditions in the deep sea due to the prevalence of slow mechanisms of yolk synthesis characteriz- ing the phylum. Seasonal cuing of organic pulses to reproduction are not likely to be strong because most species lack heterosyn- thetic pathways that would tightly couple nutrient input to egg production. VOLUME 107, NUMBER 1 213 Table 1.—Some life history correlates of slow vs. fast egg producing species. Fast egg producers Intermediate Slow egg producers Heterosynthetic yolk production Relatively “r-selected”’ Small body size High food input Low food reserve storage Variable/unpredictable environments Short-lived Low fecundity Early sexual maturity High brood frequency “Mixed” (auto/heterosynthetic yolk production) Relatively “K-selected” Large body size Low or seasonal food input High food reserve storage Stable predictable environments Long-lived High fecundity Late sexual maturity Low brood frequency Conclusion Efforts to comprehend the evolutionary forces that have shaped metazoan life his- tory patterns must consider the role played by the ovary and the diverse mechanisms of yolk synthesis that have arisen through selection. Species have different capacities for converting food into egg production re- sulting in wide variation in the interval be- tween reproductive episodes. These varying capacities are strongly influenced by the rate at which developing oocytes can synthesize yolk during oogenesis. Some species living in environments that are inherently unsta- ble or experience sudden, large scale food inputs have evolved specialized mecha- nisms of yolk synthesis that enable them to respond to sudden nutrient input through rapid egg production. Other species, partic- ularly long-lived iteroparous species, utilize slower methods of egg production and are adapted to stable environments that have seasonal or steady (predictable) inputs of nutrients. Therefore, vitellogenic mecha- nisms should play a central role in deter- mining optimal species success in any given habitat based on differential reproductive responses to nutrients. In order to avoid simplistic life history models, the complex coupling between habitat, food, feeding strategies, digestive constraints, and vitel- logenic mechanisms must be better under- stood in order to appreciate observed dif- ferences in marine community structure. Table 1 lists some life history features that correlate with slow and fast egg-producing species. Acknowledgments This paper is based on the eighth annual Riser Lecture preseuted in July 1992 at Northeastern University’s Marine Science Center. It is dedicated to Professor Nathan W. “Doc” Riser who taught me to appre- ciate the beauty of the whole organism while engrossed in a study ofits parts. The author thanks Edward Ruppert and William Jaeck- le for reviewing and commenting on an ear- lier version of the manuscript. Pamela Blades-Eckelbarger assisted with some fig- ure preparation. The author is particularly grateful to Les Watling who made numerous constructive suggestions for improving the final manuscript and who patiently assisted me in making the paper (relatively) intelli- gible to ecologists. However, any errors con- tained herein are solely the responsibility of the author. This represents contribution no. 271 of the Darling Marine Center. Literature Cited Armant, D. R., D. D. Carson, G. L. Decker, J. K. Welply, & W. J. Lennarz. 1986. Characteriza- tion of yolk platelets isolated from developing embryos of Arbacia punctulata. —Developmen- tal Biology 113:342-355. 214 Beijnink, F. B., C. W. Walker, & P. A. Voogt. 1984. An ultrastructural study of relationships be- tween the ovarian haemal system, follicle cells, and primary oocytes in the sea star, Asterias rubens. Implications for oocyte nutrition. — Cell & Tissue Research 238:339-347. Bentfield, M. E. 1971. Studies of oogenesis in the rotifer Asplanchna. IJ. Oocyte growth and de- velopment. —Zeitschnift fiir Zellforschung und mikroskopische Anatomie 115:184—195. Blades-Eckelbarger, P. I., & N. Marcus. 1992. The origin of cortical vesicles and their role in egg envelope formation in the “spiny” eggs of a cal- anoid copepod, Centropages velificatus. — Bio- logical Bulletin 182:41—53. — .,,&M.J. Youngbluth. 1984. The ultrastructure of oogenesis and yolk formation in Labidocera aestiva (Copepoda: Calanoida).—Journal of Morphology 179:33-46. Bownes, M. 1982. Hormonal and genetic regulation of vitellogenesis in Drosophila.—The Quarterly Review of Biology 57:247-274. Brafield, A. E.,& G. Chapman. 1967. Gametogenesis and breeding in a natural population of Nereis virens.—Journal of the Marine Biological As- sociation United Kingdom 47:619-627. Checkley, D. M. 1980. Food limitation of egg pro- duction by a marine planktonic copepod in the sea off southern California.—Limnology and Oceanography 25:991-998. Clark, R. B., & P. J. W. Olive. 1973. Recent advances in polychaete endocrinology and reproductive biology.—Oceanography and Marine Biology Annual Review 11:176-223. Cole, L. C. 1954. The population consequences of life history phenomena.—The Quarterly Re- view of Biology 29:103-137. Davidson, E. H. 1986. Gene activity in early devel- opment. Academic Press, New York, 368 pp. Davis, W.R. 1969. Oogenesis and its relationship to dissolved oxygen suppression in Neanthes ar- enaceodentata (Polychaeta: Annelida). Unpub- lished M.A. thesis, California State University, Long Beach, 62 pp. Eckelbarger, K. J. 1979. Ultrastructural evidence for both autosynthetic and heterosynthetic yolk for- mation in the oocytes of an annelid (Phrag- matopoma lapidosa: Polychaeta). — Tissue & Cell 11:425-443. 1976. Larval development and population aspects of the reef-building polychaete Phrag- matopoma lapidosa from the east coast of Flor- ida.— Bulletin of Marine Science 26:117—132. 1980. An ultrastructural study of oogenesis in Streblospio benedicti (Spionidae), with re- marks on diversity of vitellogenic mechanisms in Polychaeta.— Zoomorphology 94:241-263. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 1983. Evolutionary radiation in polychaete ovaries and vitellogenic mechanisms: their pos- sible role in life history patterns.—Canadian Journal of Zoology 61:487-504. 1986. Vitellogenic mechanisms and alloca- tion of energy to offspring in polychaetes. — Bul- letin of Marine Science 39:426-443. . 1994. Oocytic nutrition in the lower metazoa: the Scyphozoa. Jn H. W. Wilson, S. A. Stricker, & G. L. Shinn, eds., Reproduction and devel- opment of marine invertebrates. Johns Hopkins University Press (in press). , & J. P. Grassle. 1982. Ultrastructure of the ovary and oogenesis in the polychaete Capitella jJonesi (Hartman, 1959).—Journal of Morphol- ogy 171:305-320. ——,, & J. P. Grassle. 1983. Ultrastructural differ- ences in the eggs and ovarian follicle cells of Capitella (Polychaeta) sibling species. — Biolog- ical Bulletin 165:379-393. ——,, & R. L. Larson. 1988. Ovarian morphology and oogenesis in Aurelia aurita (Scyphozoa: Se- maeostomae): ultrastructural evidence of het- erosynthetic yolk formation in a primitive meta- zoan.— Marine Biology 100:103-115. <= 1992. Ultrastructure of the ovary and oogenesis in the jellyfish Linuche ungui- culata and Stomolophus meleagris, with a re- view of ovarian structure in the Scyphozoa.— Marine Biology 114:633-643. ,& C.M. Young. 1992. Ovarian ultrastructure and vitellogenesis in ten species of shallow-wa- ter and bathyal sea cucumbers (Echinodermata: Holothuroidea).—Journal of the Marine Bio- logical Association of the United Kingdom 72: 759-781. Emlet, R. B., L. R. McEdward, & R. R. Strathmann. 1987. Echinoderm larval ecology viewed from the egg. Pp. 55-136 in M. Jangoux & J. M. Law- rence, eds., Echinoderm studies. A. A. Balkema, Rotterdam. Fautin, D. G., & R. N. Mariscal. 1991. Cnidaria: Anthozoa. Pp. 267-358 in F. W. Harrison & J. A. Westfall, eds., Microscopic anatomy of in- vertebrates. Wiley-Liss, New York, 436 pp. Fischer, A., & A. Dhainaut. 1985. The origin of yolk in the oocytes of Nereis virens (Annelida, Poly- chaeta).— Cell & Tissue Research 240:67-76. , & H. Rabien. 1986. Molecules and cellular functions driving oocyte growth in nereid an- nelids. Pp. 195-205 in M. Porchet, J.-C. An- dries, & A. Dhainaut, eds., Advances in inver- tebrate reproduction, Vol. 4. Elsevier Science Publishers, New York. Gage, J. D., & P. A. Tyler. 1991. Deep-sea biology: a natural history of organisms at the deep-sea floor. Cambridge University Press, 504 pp. VOLUME 107, NUMBER | Gardiner, S. L., S. E. Shrader, & M. L. Jones. 1992. Preliminary observations on oogenesis in the tube worm Riftia pachyptila Jones (Vestimen- tifera).— American Zoologist 32:124A. George, S. B. 1990. Population and seasonal differ- ences in egg quality of Arbacia lixula (Echino- dermata: Echinoidea).—Invertebrate Repro- duction and Development 17:111-121. Ghiselin, M. T. 1978. The economy of the body.— American Economic Review 68:233-237. Gould, S.J. 1977. Ontogeny and phylogeny. Harvard University Press, Cambridge, 501 pp. Grahame, J., & G. M. Branch. 1985. Reproductive patterns of marine invertebrates. — Oceanogra- phy and Marine Biology Annual Review 23:373- 398. Grassle, J. F., & H. S. Sanders. 1973. Life histories and the role of disturbance.—Deep-Sea Re- search 20:643-659. ——., & J. P. Grassle. 1974. Opportunistic life his- tories and genetic systems in marine benthic polychaetes.—Journal of Marine Research 32: 253-284. Gremare, A., A. G. Marsh, & K. R. Tenore. 1989. Fecundity and energy partitioning in Capitella capitata type I (Annelida: Polychaeta). — Marine Biology 100:365-371. Gremigni, V., & A. Falleni. 1991. Ultrastructural fea- tures of cocoon-shell globules in the vitelline cells of neoophoran platyhelminths.—Hydro- biologia 227:105—-111. Greve, W. 1970. Cultivation experiments on North Sea ctenophores.—Helgolander wissenschaftli- che Meeresuntersuchungen 20:304-317. Gustafson, R. G., & R. A. Lutz. 1994. Molluscan life history traits at deep-sea hydrothermal vents and cold methane/sulfide seeps. Jn C. M. Young & K. J. Eckelbarger, eds., Reproduction, larval biology and recruitment of the deep-sea ben- thos. Columbia University Press, New York (in press). Hagedorn, H. H., & J.G. Kunkel. 1979. Vitellogenin and vitellin in insects.—Annual Review of En- tomology 24:475-S0S. Hand, C.,& K.R. Uhlinger. 1992. Theculture, sexual and asexual reproduction, and growth of the sea anemone Nematostella vectensis. — Biological Bulletin 182:169-176. Harrison, K. E. 1990. The role of nutrition in mat- uration, reproduction and embryonic develop- ment of decapod crustaceans: a review. —Jour- nal of Shellfish Research 9:1—28. Hill, S.D., & J. P. Grassle. 1981. Caudal regeneration as a measure of senescence in Capitella sp. I (Polychaeta). — Biological Bulletin 161:327. , & S. W. Mills. 1992. Regeneration and maturation in two sympatric Capitella ANS) (Polychaeta) sibling species.— Biological Bulle- tin 163:366. Hines, A. H. 1979. The comparative reproductive ecology of three species of intertidal barnacles. Pp. 213-234 inS. E. Stancyk, ed., Reproductive ecology of marine invertebrates. University of South Carolina Press, Columbia, 283 pp. Hofmann, D. K. 1976. Regeneration and endocri- nology in the polychaete Platynereis dumeri- lii.— Archives Entwicklungsuchungen. Org. 180: 47-71. Honegger, T. G. 1981. Light and scanning electron microscopic investigations of sexual reproduc- tion in Hydra carnea. —International Journal of Invertebrate Reproduction 3:245-255. Huebner, E., & E. Anderson. 1976. Comparative spiralian oogenesis— structural aspects: an over- view.— American Zoologist 16:315-343. Hutchinson, G.E. 1967. A treatise on limnology, Vol. 2, Introduction to lake biology and the limno- plankton. Wiley, New York. Kimble, J., & W. J. Sharrock. 1983. Tissue-specific synthesis of yolk proteins in Caenorhabditis ele- gans. —Developmental Biology 96:189-196. Jamieson, B.G. M. 1991. Oligochaeta. Pp. 217-322 in F. W. Harrison & S. L. Gardiner, eds., Mi- croscopic anatomy of invertebrates, Volume 7, Annelida. Wiley-Liss, New York, 418 pp. Jennison, B. L. 1979. Gametogenesis and reproduc- tive cycles in the sea anemone Anthopleura ele- gantissima (Brandt, 1835).—Canadian Journal of Zoology 57:403-411. Jumars, P. A., & R. A. Wheatcroft. 1989. Responses of Benthos to changing food quality and quan- tity, with a focus on deposit feeding and bio- turbation. Pp. 235-253 in W. H. Berger, V. S. Smetacek, & G. Weler, eds., Productivity of the ocean: present and past. John Wiley & Sons, New York. , L. M. Mayer, J. W. Deming, J. A. Baross, & R. A. Wheatcroft. 1990. Deep-sea deposit- feeding strategies suggested by environmental and feeding constraints.— Philosophical Trans- actions of the Royal Society of London 331:85- 101. Lawrence, J. M., J. B. McClintock, & A. Guille. 1984. Organic level and caloric content of eggs of brooding asteroids and an echinoid (Echinoder- mata) from Kerguelen (South Indian Ocean).— International Journal of Invertebrate Repro- duction and Development 7:249-257. Levin, L.A. 1984. Life history and dispersal patterns in a dense infaunal polychaete assemblage: com- munity structure and response to disturbance. — Ecology 65:1185-1200. 1986. Effects of enrichment on reproduction in the opportunistic polychaete Streblospio ben- 216 edicti (Webster): a mesocosm study. — Biological Bulletin 171:143-160. , & E. L. Creed. 1986. Effect of temperature and food availability on reproductive responses of Streblospio benedicti (Polychaeta: Spionidae) with planktotrophic or lecithotrophic develop- ment.— Marine Biology 92:103-113. ——, & D. V. Huggett. 1990. Implications of al- ternative reproductive modes for seasonality and demography in an estuarine polychaete. —Ecol- ogy 71:2191-2208. ——, G. R. Plaia, & C. L. Huggett. 1994. The influence of natural organic enhancement on life histories and community structure of bathyal polychaetes. Jn C. M. Young & K. J. Eckelbar- ger, eds., Reproduction, larval biology and re- cruitment of the deep-sea benthos. Columbia University Press, New York (in press). Levitan, D. R. 1988. Asynchronous spawning and aggregative behavior in the sea urchin Diadema antillarum Philippi. Pp. 181-186 in R. Burke et al., eds., Proceedings of the sixth international echinoderm conference. A. A. Balkema, Rot- terdam, The Netherlands. Linke, P. 1992. Metabolic adaptations of deep-sea benthic foraminifera to seasonally varying food input.— Marine Ecology Progress Series 81:51- 63. Lopez, G. R., & J. S. Levinton. 1987. Ecology of deposit-feeding animals in marine sediments. — The Quarterly Review of Biology 62:235-269. Mackie, G. O. 1966. Growth of the hydroid Tubu- laria in culture. Pp. 397-410 in The Cnidaria and their evolution. Academic Press, New York, 449 pp. MacArthur, R. H. 1960. On the relative abundance of species. — American Naturalist 94:25-36. ——, & E. O. Wilson. 1967. The theory of island biogeography. Princeton University Press, Princeton, New Jersey. Manahan, D. T., S. H. Wright, G. C. Stephens, & M. A. Rice. 1982. Transport of dissolved amino acids by the mussel, Mytilus edulis: demonstra- tion of net uptake from natural seawater.—Sci- ence 215:1253-1255. Marsh, A. G., A. Gremare, & K. R. Tenore. 1989. Effect of food type and ration on growth of ju- venile Capitella sp. 1 (Annelida: Polychaeta): macro- and micronutrients.—Marine Biology 102:519-527. Marshall, S. M., & A. P. Orr. 1956. Experimental feeding of the copepod Calanus fimarchicus (Gunnerus) on phytoplankton cultures labelled with radioactive carbon. — Papers on Marine Bi- ology, Oceanography, and Deep Sea Research 3:110-114. McClintock, J. B., & J. S. Pearse. 1986. Organic and energetic content of eggs and juveniles of Ant- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON arctic echinoids and asteroids with lecithotroph- ic development.—Comparative Biochemistry and Physiology 85A:341-345. McEdward, L. R., & S. F. Carson. 1987. Variation in egg organic content and its relationship with egg size in the starfish So/aster stimpsoni. —Ma- rine Ecology Progress Series 37:159-169. ——,, & F-S. Chia. 1991. Size and energy content of eggs from echinoderms with pelagic lecitho- trophic development.—Journal of Experimen- tal Marine Biology and Ecology 147:95-102. ——, & L. K. Coulter. 1987. Egg volume and en- ergetic content are not correlated among sibling offspring of starfish: implications for life history theory.—Evolution 41:914—917. Ohman, M.D. 1987. Energy sources for recruitment of the subantarctic copepod Neocalanus ton- sus. —Limnology and Oceanography 32:1317— 1330. Pearse, J.. & R. A. Cameron. 1991. Echinodermata: Echinoidea. Pp. 513-662 in A. C. Giese, J. S. Pearse, & V. B. Pearse, eds., Reproduction of marine invertebrates, Vol. VI, echinoderms and lophophorates. The Boxwood Press, Pacific Grove, California, 808 pp. Penry, D.L., & P. A. Jumars. 1990. Gutarchitecture, digestive constraints and feeding ecology of de- posit-feeding and carnivorous polychaetes. — Oecologia 82:1-11. Pianka, E.R. 1970. On r- and K-selection.—Amer- ican Naturalist 104:592-597. Pianka, H. D. 1974. Ctenophora. Pp. 201-265 in A. C. Giese & J. S. Pearse, eds., Reproduction of marine invertebrates, Vol. 1, Acoelomate and pseudocoelomate metazoans. Academic Press, New York, 546 pp. Qian, P-Y., & F-S. Chia. 1991. Fecundity and egg size are mediated by food quality in the poly- chaete worm Capitella sp.—Journal of Experi- mental Marine Biology & Ecology 148:11—25. Raff, R. A., & T. C. Faufman. 1983. Embryos, genes and evolution. Macmillan, New York. Raikhel, A. S., & T. S. Dhaialla. 1992. Accumulation of yolk proteins in insect oocytes. — Annual Re- view of Entomology 37:217—251. Razouls, S., C. Razouls, & M. Huntley. 1991. De- velopment and expression of sexual maturity in female Calanus pacificus (Copepoda: Calanoi- da) in relation to food quality.— Marine Biology 110:65-74. Reed, C. G. 1988. The reproductive biology of the gymnolaemate bryozoan Bowerbankia gracilis (Ctenostomata: Vesiculariidae).— Ophelia 29:1— 23) 1991. Bryozoa. Pp. 85-245 in A. C. Giese, J. S. Pearse, & V. B. Pearse, eds., Reproduction of marine invertebrates, Vol. VI. The Boxwood Press, Pacific Grove, California, 808 pp. VOLUME 107, NUMBER 1 Rieger, G. E., & R. M. Rieger. 1980. Fine structure and formation of eggshells in marine gastrotri- cha.—Zoomorphologie 96:215-229. Rivest, B. R. 1983. Development and the influence of nurse cell allotment on hatching size in Sear- lesia dira (Reeve, 1846) (Prosobranchia: Buc- cinidae).—Journal of Experimental Marine Bi- ology and Ecology 69:217—242. Rokop, F.J. 1974. Reproductive patterns in the deep- sea benthos.—Science 186:743-745. Roth, T. F., & K. R. Porter. 1964. Yolk protein up- take in the oocyte of the mosquito Aedes ae- gypti.—The Journal of Cell Biology 20:313-332. Runge, J. A. 1984. Egg production of the marine planktonic copepod Calanus pacificus Brodsky: laboratory observations.—Journal of Experi- mental Marine Biology and Ecology 74:53-66. Ruthmann, A. 1964. Zellwachstum und RNS- Synthese im Ei-Nahrzellverband von Ophryo- trocha puerilis.—Zeitschnift fur Zellforschung und Mikroskopische Anatomie 63:816—829. Sargent, J. R., & R. J. Henderson. 1986. Lipids. Pp. 59-109 in E. D. S. Corner & S. C. M. O’Hara, eds., The biological chemistry of marine cope- pods. Clarendon Press, Oxford. Scheltema, R.S. 1994. Adaptations for reproduction among deep-sea benthic molluscs: an appraisal of the existing evidence. Jn C. M. Young & K. J. Eckelbarger, eds., Reproduction, larval biol- ogy and recruitment of the deep-sea benthos. Columbia University Press, New York (in press). Schroeder, P. C. 1968. On the life history of Nereis grubei (Kinberg), a polychaete annelid from Cal- ifornia.— Pacific Science 22:476—-48 1. Scott, L. B., & W. J. Lennarz. 1989. Structure of a major yolk glycoprotein and its processing path- way by limited proteolysis are conserved in echinoids.— Developmental Biology 132:91- 102. ——.,, W. J. Lennarz, R. A. Raff, & G. A. Wray. 1990. The “lecithotrophic” sea urchin Helio- cidaris erythrogramma lacks typical yolk plate- lets and yolk glycoproteins.— Developmental Biology 138:188-193. Selman, K., R. A. Wallace, & V. Barr. 1988. Oogen- esis in Fundulus heteroclitus. V. The relation- ship of yolk vesicles and cortical alveoli.—The Journal of Experimental Zoology 246:42-56. Schechtman, A.M. 1955. Ontogeny of the blood and related antigens and their significance for the theory of differentiation. Pp. 3-31 in E. G. But- ler, ed., Biological specificity and growth. Princeton University Press. Shinn, G. L. 1992. Ultrastructure of somatic tissues in the ovaries of a chaetognath (Ferosagitta his- pida).—Journal of Morphology 211:221-241. Southward, A. J., & E. C. Southward. 1982. The role of dissolved organic matter in the nutrition of ANT deep sea benthos.— American Zoologist 22:647— 658. Spight, T. M. 1979. Environment and life history: the case of two marine snails. Pp. 135-143 in S. E. Stancyk, ed., Reproductive ecology of ma- rine invertebrates. University of South Carolina Press, Columbia, 283 pp. Stark, J. 1984. Synthesis of oogenesis specific proteins in Caenorhabditis elegans: an approach to the study of vitellogenesis in a nematode.—Inter- national Journal of Invertebrate Reproduction and Development 7:149-160. Stearns, S.C. 1976. Life-history tactics: a review of the ideas.— Quarterly Review of Biology 51:3- 47. Strathmann, R. R., & K. Vedder. 1977. Size and organic content of eggs of echinoderms and oth- er invertebrates as related to developmental strategies and egg eating.— Marine Biology 39: 305-309. Stricker, S. A. 1986. An ultrastructural study of oo- genesis, fertilization, and egg laying in a ne- mertean ectosymbiont of crabs, Carcinone- mertes epialti (Nemertea, Hoplonemertea).— Canadian Journal of Zoology 64:1256-1269. Tester, P. A., & J. P. Turner. 1990. How long does it take copepods to make eggs?— Journal of Ex- perimental Marine Biology and Ecology 141: 169-182. Telfer, W. H. 1975. Development and physiology of the oocyte-nurse cell syncytium.— Advances in Insect Physiology 11:223-319. Tenore, K. R. 1983. Organic nitrogen and caloric content of detritus III. Effects on growth of a deposit-feeding polychaete, Capitella capita- ta.—Estuarine, Coastal and Shelf Science 17: 733-742. Turner, R. D. 1973. Wood-boring bivalves, oppor- tunistic species in the deep sea.—Science 180: 1377-1379. Turner, R. L., & J. M. Lawrence. 1979. Volume and composition of echinoderm eggs: implications for the use of egg size in life history models. Pp. 25-40 in S. E. Stancyk, ed., Reproductive ecol- ogy of marine invertebrates. University of South Carolina Press, Columbia, 283 pp. Tyler, P. A., L. S. Campos-Creasey, & L. A. Giles. 1994. Environmental control of quasi-contin- uous and seasonal reproduction in deep-sea ben- thic invertebrates; preliminary observations. Jn C. M. Young & K. J. Eckelbarger, eds., Repro- duction, larval biology and recruitment of the deep-sea benthos. Columbia University Press (in press). Wallace, R.A. 1985. Vitellogenesis and oocyte growth in nonmammalian vertebrates. Pp. 127-177 in L. W. Browder, ed., Developmental biology. Plenum Publishing Corporation, New York. 218 Williams, J. 1967. Yolk utilization. Pp. 341-382 in R. Weber, ed., The biochemistry of animal de- velopment. Academic Press, New York. Wourms, J. P. 1987. Oogenesis. Pp. 50-157 in A. C. Giese, J. S. Pearse & V. B. Pearse, eds., Repro- duction of marine invertebrates, Vol. IX, Gen- eral aspects: seeking unity in diversity. Black- well Scientific Publications and the Boxwood Press, Pacific Grove, California, 712 pp. Wild, A. E. 1980. Coated vesicles: a morphologically distinct subclass of endocytotic vesicles. Pp. 1- 24 in C. D. Ockelford & A. Whyte, eds., Coated vesicles. Cambridge University Press. Wynne-Edwards, V. C. 1962. Animal dispersion in relation to social behavior. Oliver and Boyd, Edinburgh. REQUEST FOR SCIENTIFIC LITERATURE Do you have scientific books or journal series that you no longer use and would like to donate for distribution to Cuba? The Association of Systematics Collections (ASC) is developing a program to exchange systematics and bio- diversity information between North American institutions and Cuban insti- tutions. Please contact the ASC office with the titles of books, journals, or reports in systematics/biodiversity (especially related to Caribbean biota) that you are willing to donate, and we will arrange to ship the materials to an appropriate institution in Cuba. For more information and to contribute con- tact: Elizabeth Hathway, ASC, 730 11th Street, NW, Second Floor, Wash- ington, D.C. 20001-4521, (209) 347-2850, fax: (202) 347-0072. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Yamashita, O., & L. S. Indrasith. 1988. Metabolic fates of yolk proteins during embryogenesis in arthropods.— Development, Growth & Differ- entiation 30:337-346. Young, C. M. 1990. Larval ecology of marine inver- tebrates: a sesquicentennial history.—Ophelia 32:1-48. Darling Marine Center, University of Maine, Walpole, Maine 04573 and De- partment of Animal, Veterinary & Aquatic Sciences, University of Maine, Hitchner Hall, Orono, Maine 04469, U.S.A. The 121st Annual Meeting of the Biological Society of Washington will be held on Wednesday, 11 May 1994, at 11:00 a.m. in the Waldo Schmitt Room, National Museum of Natural History, Washington, D.C. PROC. BIOL. SOC. WASH. 107(1), 1994, pp. 219-220 INTERNATIONAL COMMISSION ON ZOOLOGICAL NOMENCLATURE % The Natural History Museum Cromwell Road London, SW7 5BD, U.K. Tel. 071-938 9387 Applications published in the Bulletin of Zoological Nomenclature The following Applications were published on 30 September 1993 in Vol. 50, Part 3 of the Bulletin of Zoological Nomenclature. Comment or advice on these Applications is invited for publication in the Bulletin and should be sent to the Executive Secretary, I.C.Z.N., % The Natural History Museum, Cromwell Road, London SW7 5BD, U.K. Case No. 2854 Robulina nodosa Reuss, 1863 (currently Lenticulina nodosa; Foraminiferida): proposed retention of neotype despite rediscovery of syntypes. 2855 Cristellaria humilis Reuss, 1863 (currently Astacolus humilis; Foraminifer- ida): proposed replacement of neotype by rediscovered lectotype, and Rotalia schloenbachi (currently Notoplanulina? schloenbachi; Foraminiferida): proposed placement on the Official List. 2871 Helix nitidula Draparnaud, 1805 and H. nitens Michaud, 1831 (currently Aegopinella nitidula and A. nitens; Mollusca, Gastropoda): proposed conservation of the specific names and designation of a neotype for H. nitidula. 2860 Pleurotoma meneghinii Mayer, 1868 (currently Asthenotoma meneghinii; Mollusca, Gastropoda): proposed replacement of neotype by redis- covered lectotype. 2841 Platynectes Régimbart, 1879 (Insecta, Coleoptera): proposed conservation. 2880 Polycentrus schomburgkii Miller & Troschel, 1848 (Osteichthyes, Perci- formes): proposed conservation of the specific name. 2868 Hydromantes Gistel, 1848 (Amphibia, Caudata): proposed designation of Salamandra genei Temminck & Schlegel, 1838 as the type species. 2873 Emys Dumeéril, 1806 (Reptilia, Testudines): proposed conservation. 220 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Opinions published in the Bulletin of Zoological Nomenclature The following Opinions were published on 30 September 1993 in Vol. 50, Part 3 of the Bulletin of Zoological Nomenclature. Copies of these Opinions can be obtained free of charge from the Executive Secretary, I.C.Z.N., % The Natural History Museum, Cromwell Road, London SW7 5BD, U.K. Opinion No. 1738. Mopsea Lamouroux, 1816 (Cnidaria, Anthozoa): Isis encrinula Lamarck, 1815 designated as the type species. 1739. Strombiformis albus Da Costa, 1778 (currently Melanella (Balcis) alba; Mol- lusca, Gastropoda): specific name conserved. 1740. Amicytheridea Bate, 1975 (Crustacea, Ostracoda): A. triangulata Bate, 1975 designated as the type species. 1741. Gerris paludum Fabricius, 1794 (currently Aquarius paludum; Insecta, Het- eroptera): specific name conserved. 1742. Lincus Stal, 1867 (Insecta, Heteroptera): conserved; L. croupius Rolston, 1983: specific name not conserved. 1743. TACHINIDAE Fleming, 1821 (Insecta, Coleoptera): spelling emended to TA- CHINUSIDAE to remove homonymy with TACHINIDAE Robineau-Des- voidy, 1830 (Insecta, Diptera), and TACHYPORIDAE MacLeay, 1825 (Insecta, Coleoptera): given precedence over TACHINUSIDAE Fleming, 1821. Cheilosia Meigen, 1822 and Pyrophaena Schiner, 1860 (Insecta, Diptera): conserved. Copromyza limosa Fallén, 1820 (currently Leptocera (Rachispoda) limosa; Insecta, Diptera): lectotype replaced, so conserving the usage of the specific name and also that of Leptocera (Rachispoda) lutosa (Sten- hammar, 1855). Drosophila putrida Sturtevant, 1916 (Insecta, Diptera): holotype replaced by a neotype. Eristalis Latreille, 1804, Helophilus Fabricius, 1805, Xylota Meigen, 1822 and Eumerus Meigen, 1822 (Insecta; Diptera): conserved. EPHYDRIDAE Zetterstedt, 1837 (Insecta, Diptera): given precedence over GYMNOMYZIDAE Latreille, 1829. Epicrium Wagler, 1828 and ICHTHYOPHIIDAE Taylor, 1968 (Amphibia, Gym- nophiona): conserved, and EPICRIIDAE Berlese, 1885 (Arachnida, Acari): conserved by the emendation of EPICRIIDAE Fitzinger, 1843 (Amphibia, Gymnophiona) to EPICRIUMIDAE. INFORMATION FOR CONTRIBUTORS Content.—The Proceedings of the Biological Society of Washington contains papers bearing on systematics in the biological sciences (botany, zoology, and paleontology), and notices of business transacted at meetings of the Society. Except at the direction of the Council, only manuscripts by Society members will be accepted. Papers are published in English (except for Latin diagnoses/descriptions of plant taxa), with an Abstract in an alternate language when appropriate. Submission of manuscripts.—Submit three copies of each manuscript in the style of the Proceedings to the Editor, complete with tables, figure captions, and figures (plus originals of the illustrations). Mail directly to: Editor, Proceedings of the Biological Society of Washington, National Museum of Natural History NHB-108, Smithsonian Institution, Washington, D.C. 20560. (Do not submit manuscripts to an associate editor). 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 and diagnoses. The style for the Proceedings is described in “GUIDELINES FOR MANUSCRIPTS for Publications of the BIOLOGICAL SOCIETY OF WASHINGTON” a supplement to Volume 103, number 1, March 1990. Authors are encour- aged to consult this article before manuscript preparation. Copies of the article are available from the editor or any associate editor. The establishment of new taxa must conform with the requirements of appropriate inter- national codes of nomenclature. Decisions of the editor about style also are guided by the General Recommendations (Apendix E) of the International Code of Zoological Nomenclature. When appropriate, accounts of new taxa must cite a type specimen deposited in an institutional collection. Review. —One of the Society’s aims is to give its members an opportunity for prompt pub- lication of their shorter contributions. Manuscripts are reviewed by a board of Associate Editors and appropriate referees. Proofs. — Authors will receive first proofs and original manuscript for correction and approval. Both must be returned within 48 hours to the Editor. Reprint orders are taken with returned proofs. Publication charges.— Authors are required to pay full costs of figures, tables, changes in proofs ($3.00 per change or revision), and reprints. Authors are also asked to assume costs of page-charges. The Society, on request, will subsidize a limited number of contributions per volume. If subsidized manuscripts result in more than 12 printed pages, the additional pages must be financed by the author(s). Multiple authorship will not alter the 12 page limit (each author will be viewed as having used his/her 12 subsidized pages). Payment of full costs will facilitate speedy publication. Costs. —Printed pages @ $60.00, figures @ $10.00, tabular material @ $3.00 per printed inch per column. One ms. page = approximately 0.4 printed page. CONTENTS Morphological and ecological variation in Otopteropus cartilagonodus Kock, 1969 (Mammalia: Chiroptera: Pteropodidae) from Luzon, Philippines Luis A. Ruedas, John R. Demboski, and Rogelio V. Sison Cryptophoca, new genus for Phoca maeotica (Mammalia: Pinnipedia: Phocinae), from Upper Miocene deposits in the northern Black Sea region Irina A. Koretsky and Clayton E. Ray A taxonomic review of Dendroica petechia (Yellow Warbler) (Aves: Parulinae) M. Ralph Browning A new species of Ocadia (Testudines: Batagurinae) from southwestern China William P. McCord and John B. Iverson A new species of montane pitviper (Serpentes: Viperidae: Bothrops) from Cochabamba, Bolivia : Michael B. Harvey A new species of Plectrohyla (Anura: Hylidae) from a premontane rainforest in northern Honduras Larry David Wilson, James R. McCranie, and Gustavo A. Cruz Identification of the taxa Xenocephalidae, Yenocephalus, and X. armatus (Osteichthyes: Urano- scopidae) Victor G. Springer and Marie-Louise Bauchot Creagrutus melasma, a new species of characid fish (Teleostei: Characiformes) from upland streams of northern Venezuela Richard P. Vari, Antony S. Harold, and Donald C. Taphorm A new genus of fossil pufferfish (Tetraodontidae: Tetraodontiformes) based on a new species from the Oligocene of Russia and a referred species from the Miocene of Ukraine James C. Tyler and Alexandre F. Bannikov Pleistocene echinoids (Echinodermata) from Bermuda and Barbados Stephen K. Donovan and Brian Jones A new species of Felicola (Phthiraptera: Trichodectidae) from a Costa Rican jaguar, Panthera onca (Carnivora: Felidae) Robert M. Timm and Roger D. Price A new species of Hyperalonia Rondani, 1863 (Insecta: Diptera: Bombyliidae: Exoprosopinae) Marcia Souto Couri and Carlos José Einicker Lamas Louisea, a new genus of freshwater crab (Brachyura: Potamoidea: Potamonautidae) for Glo- bonautes macropus edeaensis Bott, 1969 from Cameroon Neil Cumberlidge Fredius stenolobus, a new species of freshwater crab (Decapoda: Brachyura: Pseudothelphu- sidae) from the Venezuelan Guiana Gilberto Rodriguez and Héctor Suarez A new Calcinus (Decapoda: Anomura: Diogenidae) from the tropical western Atlantic, and a comparison with other species of the genus from the region Néstor H. Campos and Rafael Lemaitre Pseudocyclops lakshmi, a new species (Pseudocyclopidae: Calanoida: Copepoda) from the Lac- cadives, India P. Haridas, M. Madhupratap, and S. Ohtsuka New species of Marionina (Annelida: Oligochaeta: Enchytraeidae) from Spartina salt marshes on Sapelo Island, Georgia, U.S.A. Brenda Healy Summary and significance of overlooked Japanese literature on Myzostomida Mark J. Grygier Sciurodendrium gardneri, new species (Nematoda: Trichostrongyloidea: Heligmonellidae), a parasite of Sciurus carolinensis Gmelin, 1788 (Mammalia: Sciuridae), with comments on the biogeography of Sciurodendrium Durette-Desset, 1971 Ricardo Guerrero Description of the new genus A//opathes (Cnidaria: Antipatharia) and its type species Cirripathes desbonni Dennis M. Opresko and Stephen D. Cairns Diversity of Metazoan ovaries and vitellogenic mechanisms: implications for life history theory Kevin J. Eckelbarger Announcement International Commission on Zoological Nomenclature 79 90 97 109 114 119 122 132 137 151 164 174 179 185 193 218 219 Via BIOLOGICAL SOCIETY THE BIOLOGICAL SOCIETY OF WASHINGTON 1994-1995 Officers President: Janet W. Reid Secretary. Carole C. Baldwin President-elect: Stephen D. Cairns Treasurer: YT. Chad Walter Elected Council Robert J. Emry Susan L. Jewett Richard C. Froeschner Lynne R. Parenti Alfred L. Gardner F. Christian Thompson Custodian of Publications: Austin B. Williams PROCEEDINGS Editor: C. Brian Robbins Associate Editors Classical Languages: George C. Steyskal Invertebrates: Jon L. Norenburg - Frank D. Ferrari Plants: David B. Lellinger Rafael Lemaitre Insects: Wayne N. Mathis Vertebrates: Thomas A. Munroe Membership in the Society is open to anyone who wishes to join. There are no prerequisites. Annual dues of $25.00 (for USA and non-USA addresses) include subscription to the Pro- ceedings of the Biological Society of Washington. Library subscriptions to the Proceedings are: $40.00 for USA and non-USA addresses. Non-USA members or subscribers may pay an additional $25.00 to receive the Proceedings by Air Mail. The Proceedings of the Biological Society of Washington (USPS 404-750) is issued quarterly. Back issues of the Proceedings and the Bulletin of the Biological Society of Washington (issued sporadically) are available. Correspondence dealing with membership and subscriptions should be sent to the Biological Society of Washington, P.O. Box 1897, Lawrence, Kansas 66044, U.S.A. Payment for membership is accepted in US dollars (cash or postal money order), checks on US banks, or MASTERCARD or VISA credit cards. Manuscripts, corrected proofs, editorial questions should be sent to the Editor, Biological Society of Washington, National Museum of Natural History, Smithsonian Institution, Wash- ington, D.C. 20560. Known office of publication: National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560. Printed for the Society by Allen Press, Inc., Lawrence, Kansas 66044 Second class postage paid at Washington, D.C., and additional mailing office. POSTMASTER: Send address changes to PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON, P.O. Box 1897, Lawrence, Kansas 66044. This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper). PROC. BIOL. SOC. WASH. 107(2), 1994, pp. 221-238 THREE NEW SPECIES OF CILIATE IN THE GENERA PSEUDOCOHNILEMBUS, PLEURONEMA, AND UROTRICHA (CILIOPHORA) Gregorio Fernandez-Leborans and Apolonia Novillo Abstract.—The morphological and biometric characteristics are given for three new species of freshwater ciliates: two of them belonging to the order Scuticociliatida (Pseudocohnilembus fluviatilis and Pleuronema ovata) and one to the order Prorodontida (Urotricha rotunda). P. fluviatilis is especially dis- tinguished by the presence of an inner oral membrane consisting of six clearly differentiated kinetosomal segments. P. ovata shows a reduced oral infracilia- ture with an anterior membrane and two posterior membranoid segments, as well as a paroral membrane. In this species, there is an area in “V,” located near the posterior pole, where several dorsal kineties converge. U. rotunda is characterized by the number and composition of the caudal kinetosomic groups and of the brush kineties. The taxonomic placement of these species is dis- cussed. A number of species in the genera Pseu- docohnilembus and Pleuronema have been described. Apart from the outstanding work of Evans & Thompson (1964) and Thomp- son (1966a, 1966b) on Pseudocohnilembus further, more recent, descriptions have been made, including those of Foissner & Wilbert (1981), Fernandez-Leborans & Castro de Zaldumbide (1984) and Foissner (1985). Morphogenesis within this genus has been dealt with by Evans & Corliss (1964) and Fernandez-Leborans & Castro de Zaldum- bide (1986a). Dragesco (1960, 1968) pre- sented some early descriptions of silver- stained Pleuronema material, followed more recently by Dragesco & Dragesco-Kernéis (1986), Agamaliev (1983), Groliére & Detcheva (1974), and Small & Antipa (Pleu- rocoptes, 1978). Dragesco (1960), Foissner (1979, 1983, 1984), Alekperov (1983), Patsch (1974), Martin-Gonzalez et al. (1985) and Munoz et al. (1987, 1989) provided de- scriptions of various species in the genus Urotricha. Throughout these studies the morpholog- ical and morphogenetic features of various species in these genera have been described, a noticeable evolution having been ob- served with respect to the significance of certain structures, such as the infraciliature of Urotricha (Munoz et al. 1989). On the other hand, the number of new species de- scribed has been slowly increasing, due to the contributions of Martin-Gonzalez et al. (1985) (U. vitrea), Munoz et al. (1987) (U. nais), Munoz et al. (1989) (U. ondina) and Song & Wilbert (1989) (U. corlissiana and U. valida), and it would be necessary to car- ry out a comparative analysis of the known species and then determine which charac- teristics and which order of biometric vari- ability, serve to differentiate the species. In addition, further biometric and statistical studies of each species are necessary, their scarcity contrasting greatly with the large number of morphological descriptions. These aspects, among others, are considered here. Methods The samples containing the ciliates stud- ied were collected from three areas from the 222 Fig. 1. Pseudocohnilembus fluviatilis. General view showing the cilia and the nuclear components. outskirts of Madrid (Spain). Pseudocohni- lembus species were found in samples from the Navacerrada reservoir (48 km, NW of Madrid, 4°00'W, 40°45’N); Pleuronema species were found in the Guadarrama river at the village of Villalba (4°00’W, 40°35'N, 39 km of Madrid); Urotricha species were collected in the reservoir at La Jarosa (60 km of Madrid, 4°10’W, 40°12’N). The cil- lates were fixed with 2% OsO4 to preserve them for general biometric measurements. Some of each samples were stained with the silver carbonate impregnation technique (Fernandez-Leborans & Castro de Zald- umbide 1986b) in order to obtain perma- nent slides and photomicrographs. The sta- tistical treatment of the biometric data was carried out using the Statgraphics program. The terminology used in the different de- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON scriptions corresponds to that defined by Lynn (1988). Results Pseudocohnilembus species General morphology.—The ciliates are oval in shape, 33-36.2 um in length and 24— 27 wm in width. They have a spherical or slightly oval macronucleus with a length of 12.6-14.4 wm and a width of 10.8-13.7 um and a single spherical micronucleus, 3-3.8 um in diameter (Fig. 1, Table 1). Somatic infraciliature. —There are 8 ven- tral kinetics, each one possesses 15-17 pairs of kinetosomes. On the dorsal side there are 8-9 kineties, each with 15-16 pairs of ki- netosomes. Each pair of kinetosomes of the somatic kineties show three types of deriv- atives: 1) an anterior one extends from the right kinetosome of each pair towards the anterior right area of the ciliate (kinetodes- mal fibril) and is 1.51—1.61 um in length; 2) a posterior one extends from the right ki- netosome of each pair towards the posterior right area of the ciliate (postciliary micro- tubules) and is 0.66—0.72 um in length; and 3) a derivative extends from the left kineto- some of each pair to the anterior left area of the ciliate (transverse microtubules) and is 0.48—0.51 wm in length (Figs. 2, 3 and 4; Table 1). Oral infraciliature.—The oral area ex- tends from near the anterior pole of the cil- late to 74 of the body length. The undulating membrane or paroral formation (PF, Fig. 2) is 16.8—18.5 um in length; its posterior end is 11.88—13.2 wm from the posterior pole and 24.6—25.8 wm from the anterior pole of the ciliate. It is made up of 54-56 dikinetids that curve round in the posterior area near the cytostome. On the left side of the oral area is located the inner membrane (IM) consisting of six kinetidal structures that are described in or- der, from the anterior pole of the ciliate. 1) al is a group of 8-9 kinetosomes, 1.2—-1.4 um long and about 1.6 wm wide. 2) a2 isa 223 VOLUME 107, NUMBER 2 08 CLO 99°0 v8 0 vETe 00°0 c0'0 L9°0 sofnqny O11 Areiio}sod oeuros ay} JO yIZUT 08 1¢'0 870 80 80°C 00°0 10°0 8r'0 SO[NQNIOIOIU ISIOASULI} OIVBUIOS BY} JO YISUIT 08 19'T Ic £90 £90 00°0 100 8c Jaqy IWsapojJOULY 9eWIOS 9Y] JO YIsUT 08 OT cl L90 61 € 600 6r'0 EE SI SONIUTY [VSIOP 9} JO SIled 9UIOSOJOUTY JO IoquINNy 08 LI SI €L0 LOS 80°0 6L°0 8osl SOTOUTY [BIVUIA sy} JO Siled sUOSOJOUTY JO JoquINN 08 91 vl SL0 00°9 60°0 88°0 99 FI ge JO SQUIOSOJOUTY JO JoquINN 08 Cl Ol £90 COL 800 6L0 OS OI GB JO SOUIOSOJOUTY JO JOQUINN] 08 Ol 8 68°0 €S Ol O10 £60 £88 pe JO SOULOSOJOUTY JO IOGuINNY 08 9 iS cv 0 9EL 00 8£0 91'S eB JO SoufosojouTy Jo ToquinNy 08 8e 9E L9'0 LOC O10 860 99°9€ CB JO SOUTOSOJOUTY JO TOGUINNY 08 6 8 cv 0 So” 00 8e0 918 [® JO sourosojoury JO ToquINN] 08 9¢ VAS £90 vy 800 640 Os rs [e1oied oY} JO SoOUIOSOJOUTY JO JoquINN| 08 9ST L Yl 690 09'T c0'0 ve 0 96 FI g[0d 10L19}sod—ce sourisiq 08 8° CC 102 A Os 60°€ LO0'0 $90 OO TC god JoLiaj}uR—ce souRIsSIG 08 Cel 88 II 810 ece 00 tv 0 8ICI 90d 10L10}sod-[e101ed Jo pus 1oLIa\sod sour\siq 08 8c XG C10 61 £00 0c 0 €0°S7 [od 1oL19}ue-[e101ed Jo pus 10LI9\sod souvIsIq 08 c60 8L0 870 L8V 00°0 v00 (4:50) ge JO UIPIM 08 C8 P SIP Lv'0 LOE 100 L10 8C (QueiquioU JouuT) ge Jo yIdUST 08 Ic 1 ell cv 0 OL T 00°0 c0'0 LVI ce JO WIPIM 08 cv vIV 00 VIC 10°0 600 Oc rv (Queiquisul Jouul) Ce Jo yIdUST 08 ell LOT €£0 c6 0 00°0 100 801 pe JO WIPIM 08 ve OEE 69: 0— Iv0 00°0 100 RS (eueiquioU JouUT) pe Jo BUST 08 80 90 Sc 0 £9 00°0 r00 IZ 0 ce JO UIPIM 08 cl 801 990 OL 00°0 £0°0 ITT (ueIquioul JouUT) ¢v JO yIsUST 08 TOI 96 67 0 cS 100 S10 86 (Queiquioul Jouut) Ze Jo yIsUST 08 col 9c 1 Ie 0 £90 00°0 10°0 8c l Te JO UPIM 08 cel cl IL0 Oc Ee 00°0 00 col (euvIquIoUI JouUT) Te Jo yIsUST 08 CsI 891 T€ 0 €6°C S00 1S 0 9E LI wuOneULIOJ [e1oIEd oy} JO yIsUIT 08 6 8 L9'0 88'S 00 6¢'0 €€8 SotouTy [eSIOp JO IOGUINNY 08 8 8 00°0 00°0 00°0 00°0 00°8 Sotoury [BUSA JO IOGUINN] 08 BLE € py 0— 879 (400) cc 0 6E€ SNgfONUOISTUL OY} JO IoJoUIeI| 08 Let 801 6€ 0—- 9°°9 80°0 8L0 68 TT SNIJONUOISVU OY} JO YIPIAA 08 vri 9TCI L9°0 Ory 90°0 690 OV el sngfonuoIoew oY} JO YISUIT 08 LC VC £90 SCE 60°0 £80 eo SC YIPIM 08 C9E tt cc 0- VOT 80°0 8L0 O8 PE yisuoT suonea WNWIxey UWINUITUTY JUSIOYJS09 UOTIELIVA JO JOLIO uoneiAsp ueoul -13SqQO uosivog JUSTOWJI0D, piepuris piepuris snowy “SIIDIAN]L SnquiajluYyooopnasg JO SONStI9\IeIeYS OLNSWOIg—"] IGeL Fig. 2. Pseudocohnilembus fluviatilis. Ventral side. PF: paroral formation or outer membrane. al, a2, a3, a4, a5 and a6: the different elements of the inner mem- brane. double row of kinetosomes made up of a linear portion of 30 kinetosomes accom- panied in its anterior part by a group of 6-— 8 kinetosomes. a2 is about 10 um long. 3) Slightly separated from a2, a3 isa triangular group of 5-6 kinetosomes, about 1.1 um xX 0.7 um in size. Behind a3, there are two polykinetids running more or less transver- sally to the anterio-posterior axis of the cil- iate. The more anterior, a4, has a length of about 3.4 wm and a width of about 1.1 wm and is made up of 8-10 kinetosomes in two rows. The more posterior, @5, is just ante- rior to the cytosome and is slightly larger than a4: about 4.2 um in length x about 1.2 wm in width. It has 10-12 kinetosomes in two rows. Posterior to the cytostome and parallel to the posterior portion of the pa- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON roral, there is a group of 14-16 kinetosomes (a6) in two rows, which has a length of 4.2— 4.8 um and a width of about 0.8 um. Ina number of specimens, a pair of kinetosomes were observed posterior to a6, and corre- spond to the scutica (Figs. 2 and 3; Table 1). Taxonomic position.—These ciliates be- long to the class Oligohymenophorea de Puytorac et al. 1974, order Scuticociliatida Small 1967, Family Pseudocohnilembidae Evans & Thompson 1964, genus Pseudo- cohnilembus Evans & Thompson 1964 (Small & Lynn 1985, Corliss 1979, Small 1967, Evans & Thompson 1964). There are 10 species that are most similar to the cil- iates studied: Pseudocohnilembus persali- nus Evans & Thompson 1964; P. hargisi Evans & Thompson 1964; P. longisetus Evans & Thompson 1964; P. cantabricus Fernandez-Leborans & Castro de Zaldum- bide 1984; P. antoniensis Fernandez-Le- borans & Castro de Zaldumbide 1986; P. portuensis Fernandez-Leborans & Castro de Zaldumbide 1986; P. marinus Thompson 1966 (Foissner & Wilbert 1981); P. putrinus Foissner & Wilbert 1981; P. pusillus Fois- sner & Wilbert 1981; P. caeci Foissner 1985. These species have been compared with the ciliates studied in the following character- istics: 1, body length; 2, body width; 3, size of macronucleus; 4, number of somatic ki- neties; 5, number of kinetosomes in each somatic kinety; 6, arrangement of the so- matic kinetosomes; 7, derivatives of so- matic kinetosomes; 8, size of the oral area; 9, kinetosomal structure of the paroral for- mation; 10; kinetosomal structure of the in- ner membrane; and 11, habitat (Table 2). Finding that our specimens (not taking into account the habitat), differ respect P. per- salinus in 7; P. hargisi in 9; P. longisetus in 8; P. cantabricus in 4; P. antoniensis in 7; P. portuensis in 7; P. marinus in 5; P. pu- trinus in 8; P. pusillus in 6; and P. caeci in 6 characteristics of the 10 analyzed (Table 2). The size of the body is found 1n the range of P. hargisi, P. cantabricus and P. marinus. VOLUME 107, NUMBER 2 225 Figs. 3-4. 3, Pseudocohnilembus fluviatilis. Ventral side, the cilia, nuclear components and infraciliature can be seen (x 1640). 4, Dorsal side (x 1640). The macronucleus is similar in size to the P. cantabricus, P. portuensis and the P. cae- ci. The kinetosomes of the somatic kineties are grouped in pairs as in P. antoniensis and P. portuensis, while in P. marinus, P. pu- trinus, P. pusillus, the somatic kinetosomes are only in pairs in a part of the total length of the somatic kinety. The oral area takes up *%3 of the body length of our specimens as in P. cantabricus, while in the other spe- cies it only takes up 2 or '4 of the body length. Regarding the oral infraciliature there are two diplostichomonads, which are only present in P. antoniensis, but while this spe- cies only shows a small group of 6-8 kine- tosomes near the posterior end of the short- est diplostichomonad (IM), in our specimens there are five groups of kinetosomes, two anterior and two posterior of the IM (inner membrane: kinetosomic structures of the left side of the oral area), and a short double row of kinetosomes near the posterior end of the paroral formation. Taking into ac- count these data, and especially, the number of differences from the other species, the ciliates observed could correspond to a new species, which we have named Pseudocohn- ilembus fluviatilis. On the other hand, and taking into account the variability margins and the principal morphological character- istics, various species described could be put into one group. This is the case of P. per- salinus, P. marinus, P. pusillus, P. longisetus and P. putrinus which have no fundamental differences and could be classified together as P. persalinus, the first one to be described. They all have a similar size, number of so- matic kineties and structure of oral infra- ciliature (Table 2). Note. —Foissner (1985) points out that P. cantabricus is a synonym for P. marinus redescribed by Foissner & Wilbert (1981). However, we differ in this opinion, above all when we analyze in detail both works (Foissner & Wilbert 1981, Fernandez-Le- borans & Castro de Zaldumbide 1984): the redescription by Foissner & Wilbert (1981) is very brief and contains very little bio- metric data or explanations about the so- matic and oral infraciliature of P. marinus. But, above all, there are two fundamental features that differentiate the two species. First, the somatic kineties of P. cantabricus are each made up of a single row of 20 monokinetids, each one of these has a clear- ly visible kinetodesmal fibril. In contrast, 226 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 2.—Comparison between the species of Pseudocohnilembus. (P, pairs; s, single; st, stichomonad; sd, stichodyad; dt, diplostichomonad; Th, Thompson 1966; gk, group of kinetosomes; kd, kinetodesmic fiber; mt, transverse microtubules; mp, postciliar microtubules; m, marine; f, freshwater; s, saline; sl, soil; ec, ectoparasite; bl, body length.) P. persalinus Length (um) 30 Width (um) 14 Size of the macronucleus (um) 4.5 Number of somatic kineties 8-9 Kinetosomes in each somatic kinety 20 Arrangement of somatic kinetosomes st Derivatives of somatic kinetosomes — Size of oral area relative to body % bl length Structure of paroral formation (PF) sd Structure of the inner membrane sd (*% PF) Habitat s the kineties of P. marinus (Foissner & Wil- bert 1981) are each made up of dikinetids for the majority of their length. The number of dikinetids in the dorsal kineties, 17-19, makes the total number of kinetosomes in each kinety much greater, 31-35, than in P. cantabricus. Second, the oral infraciliature of P. cantabricus is composed of one single paroral membrane with two different seg- ments, the anterior one being greater in length, made 39-40 dikinetids, and a pos- terior segment of 12 dikinetids in zig-zag formation. In P. marinus (Foissner & Wil- bert 1981), the oral infraciliature is com- posed of two membranes that show the ki- netosomes in zig-zag formation, the shorter (inner membrane) with a posterior kineto- somal group. It is evident taking these dif- ferences into account, that we are not deal- ing with the same species, as Foissner (1985) indicates. Pseudocohnilembus fluviatilis, new species Diagnosis. —Rounded, oval in shape, of 33-36 wm in length and 24—27 um in width. A spherical or oval macronucleus of 12.6- 14.4 wm X 10.8-13.7 um with an adjacent P. hargisi P. longisetus P. cantabricus 44 26.6 34.8-40.8 18 11.5 22.8-25.8 4.2-6.7 3.7 13.2-20.4 x 12-17.4 14 11 10 (12) 27 16 20 st st st — — kd /, bl /, bl %, di 21.8 12.8 21-25.2 x 3.6-4.2 sd sd dt + sd sd (=PF) sd (=PF) — Ss m m micronucleus of 3—3.8 wm of diameter. Eight ventral kineties and 8—9 somatic dorsal ki- neties, each with 15-17 dikinetids. The oral area with a paroral formation, which is 16.8— 18.5 um in length with a short polykinetid of two rows near its posterior end and, on the left side, a linear polykinetid (inner membrane) of 14-14.9 um in length with two small polykinetids anterior and two posterior. Freshwater. Pleuronema species General morphology. —Ciliates, oval in appearance, of 70.8—82.8 um in length and 53.4-60.6 wm in width. A rounded mac- ronucleus is usually located in the anterior half of the body, 15.6-—21.2 um long and 15.9-20.1 um wide. There are two spherical micronuclei 2.3—2.5 um in diameter located beside the macronucleus. (Fig. 5; Table 3). Somatic infraciliature. —There are 29-31 somatic kineties, of which 15—16 are ventral and 14-15 are dorsal. The majority of the kineties are bipolar, except for 6 dorsal ki- neties and 5—6 ventral kineties. The shortest ventral kineties are found anterior and left of the posterior of the oral area. They have a length of 42-46.8 um and are made up VOLUME 107, NUMBER 2 227 Table 2— Extended. P. antoniensis _ P. portuensis P. marinus P. putrinus P. pusillus P. caeci P. fluviatilis 12.9-16.5 18.6-24.9 32-36 17-27 25-42 59-105 33-36.2 10-13.5 11.8-17.7 20-22 6.6-14.6 12-26 22-42 24-27 5.8-8.4 x 7.2-12.6 10.5-11 x 9.3-10 48 x 46.6 5-8 x 5-8.1 10-14 x 8-14 12.6-14.4 x 2.7-4.9 10.8-13.7 10 10 8-9 (10 Th) 10 10-11 10-14 16-17 12-18 p 18-20 p 29-33 p + 5-8 s 14-17 p+s 15-23p+s 3346p+s 15-17 p sd sd sd + st (st Th) sd + st sd + st sd + st sd kd kd, mt,mp — — = = kd, mt, mp Y, bl Y bl Y bl Y bl Y% bl Y%. bl *%, bl 9.15-11.4 dt st + sd sd + gk (st Th) sd sd sd + st dt + dt di+6-8k — sd (st Th) sd sd sd + st 2 gk + dt + 3 gk m m f (m Th) sl f m (ec) f of 22—26 dikinetids. The shortest dorsal ki- neties are found between the anterior pole and an area in the left posterior region where various kineties converge in a ““V” shaped suture. In the center of this area, there are two parallel kineties (4 and 5) (somatic ki- nety | is situated on the right of the oral infraciliature) of 53.1-54 wm in length and 38—40 dikinetids each. Lateral to these two kineties another two are found (3 and 6) that converge beneath the posterior end of the previous two; they have 42—44 dikinetids each. Lateral to these last two kineties (3 and 6) are another two (7 and 2) that also converge posteriorly and have 46-48 diki- netids each. The remaining somatic kineties have 50-54 dikinetids each. In each pair of somatic kinetosomes, the one on the right has a thick derivative that runs from the kinetosome to the right anterior area of the ciliate (kinetodesmal fiber) and is 1.8—3 um long. There is a fibrillar net that circles and accompanies the pairs of somatic kineto- somes (Figs. 6-9; Table 3). The oral infraciliature.—The oral area takes up a large part (51—57 um) of the total length of the individual and is composed of three kinetosomal structures: 1) the paroral formation (PF); 2) membrane 1 (M1) and 3) the pericytostomal structures (oral for- mation, OF). The paroral formation (PF) has a length of 39.6—45 um and is longitudinally located on the left side of the oral area. The anterior end of this structure is 21.3—23.2 um from he anterior pole and is 60.6-62.8 wm from the posterior pole. The posterior end of the paroral is found near the anterior area of the oral formation (OF). This structure is made up of 140-144 dikinetids. Accom- panying the paroral formation there is a fi- ber that runs parallel to this structure for its whole length and extends posteriorly, hav- ing a length of 45—5S0.4 um (subparoral fiber, SPF). Paroral dikinetids connect by means of fine prolongations (a) with the subparoral fiber. M1 is found near the anterior end of the paroral formation with its posterior end slightly separated from this structure. It is 5.94-8.4 um and is made up of 24-38 ki- netosomes grouped in pairs (12-19 diki- netids). The oral formation (OF) is in the poste- rior oral area, encircling the cytostome. This structure is divided into two parts: one lon- ger one made up of a single row of 60-70 kinetosomes (stichomonad), M2, and an- 228 he | bY E < if AR is Hyer W =) Ge Nay Aad ANN meee, SE] 7207 SSN =) 99997, Nee [ee RAR nin 117 SRN RS We Oe Dae Ay Fig. 5. General view of Pleuronema ovata showing the cilia and nuclear components. other shorter one separated of the anterior for a zone without kinetosomes, found on top of the corresponding cytostome area, and made up of a single row (stichomonad) of 12-16 kinetosomes (M3). Parallel to the oral formation is a thick fibrous structure called the suboral fiber (SOF), which forms fine connections (b) with each of the kineto- somes of the oral formation. The suboral fiber is 45—48.6 wm long and connects up with the subparoral fiber. The oral formation circles a fibrous group (ribbed field, Small, 1967) that is constitut- ed of various structures. First, a closed fi- brillar structure, more or less circular, im- mediately defines the entrance to the cytostome, and is called oral inner fiber (OIF). It is 3.4-5.2 um in length. Second, an open fibrillar structure, the external oral fiber (OEF), runs parallel to and accompa- nies the suboral fiber along part of its length. It is 36.8-38.5 um long. Third, between the oral inner fiber and the external oral fiber PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 6. Pleuronema ovata. Ventral side. SPF: sub- paroral fiber; a: fibrillar connections between the pa- roral formation (PF) and the SPF; IOF: intermediate oral fibers; OEF: oral external fiber; b: fibrillar con- nections between the OEF and the M2; SOF: suboral fiber. there is a group of fibers called intermediate oral fibers (IOF). Some of these fibers (4-6) are connected to an oral inner fiber by one end while the other end remains free in the oral cavity; these fibers measure 1.8—3.6 um. The remaining fibers (10-14) have as much connection with the oral internal fiber as with the external oral fiber, and are 9-12 um long (Figs. 6 and 8; Table 3). Taxonomic position. —The specimens studied belong to class Oligohymenophorea De Puytorac et al. 1974, order Scuticocilia- tida Small 1967, suborder Pleuronematina Fauré-Fremiet in Corliss 1956, family Pleu- ronematidae Kent 1881, genus Pleuronema Dujardin 1836 (Small & Lynn 1985; Corliss 1979: Small 1967; Groliére & Detcheva 1974; Small & Antipa 1978; Dragesco 1960, 1968; Dragesco & Dragesco-Kernéis 1986; Agamaliev 1983). The structural simplicity of the oral infraciliature of these individuals 229 VOLUME 107, NUMBER 2 = 08 £81 VST 790 pss O10 £60 8L 91 90d 10119}sod—uoneui0y [210 JO pua 10119}sod goue}sIq 08 IS 8 9P (GGR0) lec C10 ell 68 8P god 1oLIo]Ue—uONeUIIO; [B10 JO pUud JOLIO}UR dOURISIG 08 ICE 9°87 LV0O- SOE O10 £60 br Of god JoLoyue—Ioqgy [eUID}UT [210 JO pus JO1I9}sod souRISIq 08 CCS OLY S10 OL 7 C10 yaaa 8 6P 90d 1oLIo}ue—Ioqy [eUID}UT [eIO JO pus JOLIO}Ue doULISIG 08 8°79 9°09 €£0-— ZO'I L0'0 €9'0 6S 19 aJod 10L10}sod—uonew.i10y [e1o1ed JO pud 1OLIg}Ue 90UR\SIG 08 CEC C1? £00 97 c0'0 €S°0 CV CT god 1OLIo}Ue—uONeULIOJ [e101ed JO pud JOLID}Ue J0ULISIG 08 vs OS 8c 0 61°C C10 SIT €ECS SONOSUTY [BSIOP JO 1So1 OY) JO Sited sUIOSOJOUTY 08 8YV OV €c70- IST L400 IL 0 €8°9V L pue Z sonoury [esiop Jo sired owosojouny 08 bY (G4 00°0 69'T 80°0 €L0 eV 9 pure ¢ soljoury [esiop Jo sired awosojouny 08 vs Tes L8°0 LS‘0 £00 1€0 LOES ¢ pur p Soljoury [esiop Jo ysuST 08 OV SE €10- 69'T L400 99:0 16°8€ ¢ pur p Soljoury [esIop Jo sired suIOsojaUrYy 08 8S cS 60°0 90'€ 810 69° ESS Ajoury [2IUSA Yea JO sited UIOSO}JOUTY JO JoquInNy 08 98 cv ¢8°0 SEC C10 am 96 91 Joqy [e1oqns Jo yy3uaT 08 T€ 67 0 (6 90°0 09°0 Of SOTOUTY IVULOS JO IOUINNY 08 pOs oY v0'0 vLT 1721 0) 6c 1 90°LY Joqy [e1oredqns jo yy3ueT 08 C8 OL ¢0'0- S07? 810 co T 16°82 UOTCULIOJ [VIO SY} JO SOUIOSOJIUTY JO JoquINNy 08 bri orl 00°0 460 vO bel GA UONLUIOJ [e101ed JO SoWIOSOJOUTY JO JoquINN] 08 SE VC $80 60°ST 0s 0 ISp L8°6C IW JO sowosojoury Jo Joquinn 08 CBE 89¢ 17 0- IST 90°0 LS0 SOLE Joqy [BUI9}xe [vIO dy} JO YIZUST 08 £9 12 COT 06 CI 60°0 (G:00) 61S 9uoz 1oqy [BUIOJUT [BIO 94} JO YIPIM 08 (ES VE Iv 0- STE! 90°0 09°0 coP SUZ JoqY [PUIOIUT [PIO oY} Jo YIsUST 08 V8 v6'S 0S 0- 09°8 90°0 19°0 60°L TW 2y) Jo yausT 08 oY 9°6£ 110 v6C vIO Lot SO 2 UOTIEUIO} [e1OIed Jo yysUIT 08 CSC VET 0c 0 LOT 00°0 ¢0'0 Ipc TQPONUOIOIUL JY} JO JoJQUIvIG 08 9102 6ST L9'0 (GM) C10 SIT 8I8I SnogpINUuoIDVUr 9Y} JO YIPIM 08 PCI 9ST 6c 0 cv 8 L10 9S I or 81 snoponuoJoeU! 94} JO YISUIT 08 9°09 pes 690 eve 1c 0 £61 9096 WIpIM 08 8°C8 8 OL c70- v6E ve 0 SOE ELL yysusT SUOT]BA WNUWIxey] UINWEIUTAY JUSTOWJO09 UOTIELIVA JO JOLNIS uOol}eIASp ueouwl -138SqO uosivog JUSIOWJI0D, Piepuris piepuris onewy Ly ‘DIDAO DULAUOANA/_ JO eyep IIIWOIG—"¢ IIqe_L 230 Fig. 7. Pleuronema ovata. Dorsal side showing the nuclear arrangement and the “V” area near the pos- terior end of the ciliate. Figs. 8-9. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON makes it unnecessary to compare, from bi- ometric point of view, this Pleuronema spe- cies with those previously described, which undoubtedly have a greater kinetosomal complexity, particularly in the posterior zone of the oral area. Taking into account the published data (indicated above), these cil- iates can be placed in the Pleuronema sim- plex species (Dragesco 1960), which is the one that shows a greater reduction of the oral infraciliature. However, our specimens lack the posterior segment of M2 (Dragesco & Dragesco-Kernéis 1986), although it is somewhat reduced in Pleuronema simplex. Furthermore, this species is from a marine habitat, while the Spanish population is from freshwater. Thus, a new species is proposed, Pleuronema ovata. Pleuronema ovata, new species Diagnosis. —Oval-shaped ciliates, 70.8— 82.8 wm long and 53.4-60.6 wm wide. Pleuronema ovata. 8, Ventral side of a stained specimen (x 1510). 9, Dorsal side (x 1360). VOLUME 107, NUMBER 2 Spherical macronucleus of 15.6—21.2 um x 15.9-20.1 um in size, with two adjacent mi- cronuclei of 2.3—2.5 um in diameter. The oral area is of 51-57 wm in length with a infraciliature reduced to a paroral, an an- terior membrane M1, and two membranoid segments M2 and M3 (stichomonads). Fif- teen—16 ventral kineties, 6 of which con- verge in a “V” zone located anterior to the posterior pole. Freshwater. Although the oral region of Pleuronema is situated laterally, we call the zone that includes the oral region “‘ventral’’ and the opposite “dorsal” to make this description correspond to those of other scuticociliates. The pericytostomal structures have tradi- tionally been included in the paroral for- mation, but their kinetosomic composition is different from that of the latter, and they include two zones, M2 and M3 (sticho- monads), separated by an area without k1- netosomes. The fibrillar components of the pericytostomal structures are also different from those of the paroral formation. Urotricha species General morphology. —Ciliates, rounded oval in appearance, 48-55.2 um in length, and 45-48 um in width. They have an oral opening located in the anterior pole of the individual, which is 2.9-3.6 um long and 2.2—3.2 wm wide. The oval macronucleus is 19.2—22.2 wm long and 13.2—15.4 um wide. The micronucleus, is located beside the macronucleus, and is spherical with a di- ameter of 4.8-6 um. The contractile vacuole pore is located half-way along the body be- tween the kineties 12 and 13, and is 31.2- 32.4 um from the anterior pole and 21.6— 22.4 wm from the posterior pole of the cil- late (Fig. 10; Table 4). Somatic infraciliature. —There are 45-48 somatic kineties that run between the area near the oral opening and a posterior zone without kinetosomes. Three of these kine- ties are shorter than the rest as they abut on the brush. The posterior end, where the so- Fig. 10. General view of Urotricha rotunda show- ing the cilia and the nuclear components. matic kineties are broken off, is 40.8—42 um from the anterior pole and 14.6-15.2 wm from the posterior pole. Each somatic ki- nety has 30-36 pairs of kinetosomes. The number of pairs of kinetosomes is slightly less on the dorsal side than on the ventral: 30-32 vs. 33-36. The 3—4 most anterior pairs of each kinety are more closely grouped forming a border 1.8-2.4 um wide. Each pair of somatic kinetosomes has three de- rivatives: the two associated with the right kinetosome are kinetodesmal fibril and postciliary microtubules, and those associ- ated with the left kinetosome are transverse microtubules. In the posterior area of the ciliate there are 6-8 groups of 2—4 kinetosomes each. The kinetosomes of each group appear con- nected to each other by means of a fibrous structures, in such a way that each group as a whole, has a circular appearance. These caudal kinetosomal groups (CKG) give use to the caudal cilia (Figs. 11-13; Table 4). Oral infraciliature. —This is made up of two structures: the perioral formation (PF) and the adoral organellar complexes (brush). The perioral formation (PFO) consists of PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 232 SS SS 08 Ov CC 09 17 1&0) 660 c0'0 (G0) (4K6 g]0d 1oL19}sod—sjondeA a[1}9e.11UOd Jo a10d gour\sIq 08 OV CE Oc TE crv 0 L8°0 £00 870 76 TE gJOd 1oLI9}Ue—sJONdeA 9[119e1]U09 Jo 910d JDue1sSIG 08 eel OTT 80°0 £30 000 100 Oc I dno1d eusosojoury [epnes yovs Jo YIPI A 08 891 9c 1 990 L831 000 £0°0 09'T dnosd [ewrosojoury [epneo yove jo y}sus] 08 06° LY 03°97 00°0 690 £00 ££ 0 OC LY gjod 101L19}sod—¢g s0urisiq 08 OVLI 00ST 6L°0 8f 7 L0°0 69°0 SL SI gjod JoLiajue—-¢g s0uRISIG 08 006+ 00 Lr £10- eel 90°0 650 C6 LY gjod 10110}sod—7g s0uK\sIq 08 Or vl OO! LS0 LOT 400 0r'0 tv el gjod JoLiajue-—7g s0uRIsSIq 08 08°0S Ol 8r 0) 6c I L0'0 790 Of 6h gjod 10110}sod—| g s0ur\siq 08 06 TT 08 OI 1S0 09°C £00 670 STITT gjod 1oLiajue—] gq s0uRIsSIq 08 O07 SI 09 FI (0) 0) 8c (40K0) 610 (6: [0d 1o110}sod-sonoury seUlOs Jo pus JoL9\sod souR\sIq 08 00°Cr 08 OF £0°0 0L0 £00 6c 0 IC Iv god 1o119]Ue—soroULy O1eBWOs JO Pua JOLIA}sod souLIsIq 08 O00'CE 00°0¢ vl O- IS 80°0 6L 0 Iv le AjQuTy [eSIOp Yyove ur sired 9UIOSOJOULY JO Joquinyy 08 00°9¢ OO EE 80°0 06°C ITO 660 80° VE AJQUIY [BUSA Yoo ul sited aUIOSOJoUTY JO JaquinNy 08 OVC 08'T 8c 0 0s'8 100 L10 00°C SONOULY NUIOS 9} JO JUSWUBEs JOLIO}Ue 9} JO YIZUIT 08 00°F 00'€ cv0 CO'CI 00 8£°0 OTE SONOUL] IVWOS JO JUSWIZaS IOLIOJUL Jo sited JUIOSOJIUPY “ON 08 00°€7 00°CZ cc0 (406 c0'0 Sv 0 CC CC SoTjouLy [eSIOp JO JoquinNy 08 00°SZ 00°€7 660 8re 60°0 £80 €8 EC Sonoury [eUdA Jo Joquinyy 08 00°F 00°C 00°70 00°07 90°0 09°0 00'€ dnoig [eWIOsoj}oUTy [epned Yora JO SOWOSOJOUTy “ON 08 00°8 00°9 €10- cS6 L0°0 99°0 169 sdno1d [ewosojoury [epned Jo Joquiny| 08 00°77 00°72 00°0 09°C 90°0 090 00'€C UONeUIIOS [e1OLIed Jo so[pung Ie][LQYy jo aquinNy 08 00'r 00'€ 08°0 C6 rl ¢0'0 1g) Ive €_ JO Sowlosojoury fo Joquinyy 08 00° 00°9 cs 0 OC L S00 20) S79 Ca JO SOULOSOJOUTY JO IoquINN 08 00°01 00°24 9€ 0 SII ITO COT LE8 Tq JO sowulosojoury JO Joaquin] 08 061 0c I L70 0601 70'0 810 ool €a@ JO ysusT 08 977 891 ¢0':0- b0°6 (40x0) 810 66 1 Ca JO ysusT 08 Of € v9T 97 0- (GO) 700 610 SOE 1a Jo yisusT 08 00°9 08 P 1¢0 6S°9 £0°0 ce 0 Tes SNIPINUOAITUL OY} JO JoJoUTeIG 08 Ov'sI Oc EI 9°°0 CE V 90°0 790 Se vl SngfonuoioeU 24} JO YIPIAA 08 07 C7 07 61 650 SOV 60°0 £80 6 07 sngponuoIDeU 9Y} JO YIBUIT 08 Os T 97 T cL0 L19 00°0 80°0 9E | UONeUIIOJ [eIOLIod oy} JO sa_pung IeT[LUQY sy} Jo yIUST 08 O8'L 09°9 60°0 Ory £00 1€0 Of L QUOZ UONLUIIOJ [e1OLIAd 9y} JO YIPIA 08 Or IT 08 01 ££ 0 cel 100 c10 SO TT 9UOZ WON eUIIO} [e1OLIed ay} Jo yIsUaT 08 OC E 917 cv 0 68°8 (40K) veo0 IL 7 Buluado [e1O yi JO YIPIM 08 OS v6 9£ 0 169 700 (Gav) 8I€ sutuado [e10 ay) Jo yI3UusT 08 00°8r 00°SP Sr0 pL 60°0 780 L89v YIpIM 08 OT SS 00°8r 8r'0 6f € 610 OL I pS is yisueT SUOT]EA WINUIIXey] wnuulyy JUSIOLJ309 UONBLIBA JO JO.LI9 uoneiAsp uvoul -138qQ uosieag JUSTIOYJ3OD piepuris piepuris onowYyILy ‘DPUNIOA DYIIAJOAQ JO Rep MUjJOWOIg —“p IIGeL VOLUME 107, NUMBER 2 a crown of 22-24 pairs of kinetosomes that circle the oral opening. From each of these pairs of kinetosomes and towards the buccal opening, there is a fibrous bundle of 1.3-1.5 um long. The perioral formation as a whole is 10.8-11.4 wm long and 6.6—7.8 wm wide. There are three adoral organellar com- plexes (brush kineties) (B1, B2 and B3) run- ning more or less meridionally from the an- terior part of the ciliate to its equatorial zone. B1 is the most anterior, 10.8—11.9 um from the anterior pole and 48.1-50.8 um from the posterior pole of the ciliate; it is 2.6—3.3 um long and is made up of two rows that have a total of 7-10 kinetosomes. B2 is 13-— 14.4 wm from the anterior pole, and 47-49 um from the posterior pole of the individ- ual; it is 1.7—2.3 um long and is made up of 6-7 kinetosomes grouped in two rows. B3, the most posterior, is 15—17.1 um from the anterior pole and 46.8-47.9 um from the posterior pole of the ciliate; it is 1.2-1.9 um long and has 3-4 kinetosomes (Figs. 11 and 12; Table 4). . 7 o es i Fig. 12-13. Dorsal surface. The caudal kinetosomic groups can be observed (x 1800). 233 Fig. 11. Urotricha rotunda. PFO: perioral forma- tion; B1, B2 and B3: brush kineties; CKG: caudal ki- netosomic groups. Taxonomic position. —The specimens studied belong to the class Prostomatea Schewiakoff 1896, order Prorodontida Cor- liss 1974, Family Urotrichidae Small & 13 12, Urotricha rotunda. The ventral side of a stained specimen (x 1800). 13, Urotricha rotunda. (ysniq) Z le) al (©) g so < =) € os € € (Ux 9)E S t € t t sop[ouersi0 [elope JO JoquUINN S Soljouly 6 = COE! = d ¢1 zt (a9 € s G S fi=S onBWOs TOYS JO JEquINN] SS un d o€ UI 4 O€-LZ uOT}eULIO} [e1OLIod fe = I = = dol d Q7-SI po dce-0e dg9z 417-97 d S7-€7 34} Ul S9UIOSO}SUTY JO JOQUINN oO (dnoi3 yore ul souios eo) -oJaury Jo oquinu) dnos3 a L-S I 1 C (II (DI = (1)81 (II = I dTWOSOJUTY [BpNed JO IOQUINN, O = 91-6 = = I ua I Ol 81 I s L-S eITID epnes JO IoquinN So) Ajyoury oneulos yors © = 97-£7 ~—s oI 9TVC aa O0t—-0¢ oe = = = 8I-91 Jo sourosojoury JO JoquinNy © Ux L9-09 ma = Vat = = = 7-61 Iv-S€ OS-SP 09 19-6S 1S-8P OS-SP sotjoury INeUIOS JO JIOQUINN = souOsO}oUTY JO Joquinu = = 9SS/G/= =A = --/(C-S1\ —/€/t=8°C a = a = CO YVG- /SMOI JO Jaquinu/y}sus| -¢ a 6 sauiosojaury Jo Joquinu n = OOK = = = SPHRE-G' AHS at = al = Owe /SMOI JO Joquinu/YyIsue| °7 A g souIOsSOJoUTY JO Joquinu ra) oe = =a = S$NE-G\' AEC =i = = — 8gI-O1/C/— /SMOI JO Jaquinu/yisug] :1 a ea gjod 10110}3sod—au0z Iejjoues1o0 ©) — — = — = 9€-SZ — — — _ — [e10pe Jo pus 1oLe}sod soue3sIq g gjod IOLIo}uUev—sUOZ Ie][aues10 pu - — — _ — SI-8 — — — — — je1ope JO pus JoLIasod s0urisIq Is IS 6 IS — = II-S°L _ — — — — Sna[ONUOIOeU 9Y} JO UIPIM Is UM 12 LI sngjonu Is IS [I-L Is AO BT AO CI-Ol 12 IS6I-IT -o10eUI ay} Jo adeys pue yIZUST — ce-91 = = 8t—-07 CEVT OS-8E 09-SS 1S-9P = ¢9-6E YIPIM Ov-S7 Cr-lE 08 Or-61 0c-0l Or-0€ UN Sb-St UA SS-LY ¢9-0S cL CS-8P 09-0S L9--V yysueT (,)DIDUUD puipuo S113D “1 DUuojs pjvAo “1 DIDULID “() poispjad p21uo paiuanyds ‘() 1ovs0}And “/) D1DISDI “1 n n -OLJDU ‘() -4aysdD “() *(QUOIOD BIQNOP “pd ‘gZuts ‘s ‘SE61 eM UI ‘sired ‘d ‘oureyds “Is = 7 = ce—-Cl vy—-9T cv UIPIM €9-cS €€-ST Cc SS-8P 08-S9 - OV-SE ct-61 99 pe-91 LY-LT 07 yisusT SSS Eee Dpijoa DUDISSIJ109 ppunjos D91]]0q jaanvf Djo40f x1jDUaA sipu ‘:) Dasqia pjiydosjos a 0) a a a n a n a nd a ee 2 Ee a 8 De 2 er eee eee ‘ponunuoy—"s 21921 VOLUME 107, NUMBER 2 236 Lynn 1985, and genus Urotricha Claparéde & Lachmann 1895. The best-known species of this genus are: Urotricha castalia Munoz etal. 1987; U. puytoraci Dragesco et al. 1974; U. sphaerica Groliére 1977; U. apsheronica Alekperov 1983; U. pelagica Wilbert 1986; U. armata Kahl 1927 (Foissner 1984); U. ovata Kahl 1926 (Foissner 1979); U. ma- crostoma Foissner 1983; U. agilis Stokes 1886 (Foissner 1979); U. ondina Munoz et al. 1989; U. armata Kahl 1927 (Dragesco 1960); U. satrophila Kahl 1935 (Patsch 1974); U. vitrea Martin-Gonzalez et al. 1985; U. nais Munoz et al. 1987; U. venatrix Kahl 1935; U. farcta Dragesco et al. 1974; U. faurei Dragesco et al. 1974; U. baltica Cza- pik & Jordan 1976; U. corlissiana Song & Wilbert 1989, and U. valida Song & Wilbert 1989. The species that permit a more de- tailed comparison, due to the fact that there is more biometric data available, are A, U. castalia, F, U. armata, G, U. ovata, J, U. ondina, L, U. satrophila, M, U. vitrea, N, U. nais. With respect to these 7 species, our specimens differ from U. castalia in 10; U. armata in 15; U. ovata in 10; U. ondina in 11; U. satrophila in 10; U. vitrea in 10 and from U. nais in 13 of the 16 characteristics it has been possible to consider: 1, body length; 2, body width; 3, length of the mac- ronucleus; 4, width of the macronucleus; 5, distance between the posterior end of the adoral organelles (brush) and the anterior pole; 6, distance between the posterior end of the adoral organelles (brush) and the pos- terior pole; 7, B1 length/number of kineties/ number of kinetosomes; 8, B2 length/num- ber of kineties/number of kinetosomes; 9, B3 length/number of kineties/number of ki- netosomes; 10, number of somatic kineties; 11, number of kinetosomes of the somatic kineties; 12, number of caudal cilia; 13, number of caudal kinetosomal groups (CKG); 14, number of kinetosomes of the perioral formation (circumoral corone); 15, number of short somatic kineties; and 16, number of adoral organelles (brush). With PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON regard to the ciliates studied, the length of the body is greater in U. apsheronica, U. armata (Dragesco 1960), U. venatrix and U. baltica. The body is shorter in U. ovata, U. macrostoma, U. agilis, U. ondina, U. satro- phila, U. vitrea, U. nais, U. farcta, U. faurei and U. corlissiana, being similar in the rest of the species. The number of somatic kineties is higher in U. sphaerica, U. ap- sheronica, U. venatrix, U. faurei and U. val- ida, and lower in U. armata, U. ovata, U. macrostoma, U. agilis, U. ondina, U. satro- phila, U. vitrea, U. nais, U. farcta and U. baltica. The number of caudal cilia is sim- ilar in U. apsheronica, U. venatrix, U. faurei and U. valida, being lower in the rest of the species. The number of kinetosomes in the perioral formation is higher in U. puytoraci, U. sphaerica, U. apsheronica and U. valida, and lower in U. armata, U. ovata, U. ma- crostoma, U. ondina, U. vitrea, U. nais, U. farcta, U. faurei and U. corlissiana. Due to these various differences, we conclude that our specimens represent a new species, Uro- tricha rotunda (Table 5). Urotricha rotunda, new species Diagnosis. — Ciliates, round or oval in ap- pearance, 48-55.2 um long and 45-48 wm wide. Oval macronucleus (19.2—22.2 um x 13.2-15.4 wm in size) with an adjacent spherical micronucleus 4.8-—6 wm in diam- eter. 45-48 somatic kineties broken off in the posterior zone of the body, with 30-36 pairs of kinetosomes each. 6-8 caudal ki- netosomal groups. Perioral formation of 22— 24 pairs of kinetosomes. Type specimens: permanent slides stained with silver carbonate technique, deposited in the Laboratorio de Biologia General, De- partamento de Biologia Animal I (Zoolo- gia), Facultad de Biologia, Universidad Complutense, ref. n. 2314 a—f (Pseudocohn- ilembus fluviatilis), ref. no. 3126 a—g (Pleu- ronema ovata), ref. no. 2788 a-l (Urotricha rotunda). VOLUME 107, NUMBER 2 Acknowledgments Special thanks are due to professor D. Lynn for the revision and the critical com- ments of this manuscript. This work was supported by the C.I.C.Y.T. grant NAT90- 0059. Literature Cited Agamaliev, F.G. 1983. Ciliates of the Caspian Sea. Systematics, Ecology, Zoogeography. Nauka, Leningrad, 220 pp. Alekperov, I. Kh. 1983. New species of ciliates (Gymnostomata) from water bodies of Azer- baijan.— Zoologicheskii Zhurnal 63:1417—1420. Claparéde, E., & J. Lachmann. 1895. Etudes sur les infusoires et les rhizopodes. — Mémoires Institut Naturel Genévois 5:1-260. Corliss, J. O. 1974. The changing world of ciliate systematics: historical analysis of past efforts and a newly proposed phylogenetic scheme of clas- sification for the protistan phylum Ciliopho- ta.—Systematic Zoology 23:91-138. . 1979. The Ciliated Protozoa. Pergamon Press, Oxford, 455 pp. Czapik, A., & A. Jordan. 1976. Les ciliés psammo- philes de la mer Baltique aux environs de Gdansk.— Acta Protozoologica 15:423-446. De Puytorac, P., et al. 1974. Proposition d’une clas- sification du phylum Ciliophora Doflein 1901.— Comptes Rendus Academic Science Paris 278: 2799-2802. Dragesco, J. 1960. Ciliés mésopsammiques litto- raux.— Travaux Station Biologie Roscoff 12:356 pp. . 1968. Les genres Pleuronema Dujardin, Schi- zocalyptra nov. gen. et Histiobalantium Stokes (ciliés holotriches hyménostomes).—Protisto- logica 4:85-106. —., & A. Dragesco-Kérneis. 1986. Ciliés libres de l’Afrique intertropicale. Ed. Institut Fran¢ais de Recherche Scientifique pour la Développe- ment en coopération. Collection Faune Tropi- cale n° 26, 559 pp. , F. Iftode, & G. Fryd-Versavel. 1974. Con- tribution a la connaisance de quelques ciliés Holotriches Rhabdophores: I. Prostomiens.— Protistologica 10:59-75. Dujardin, F. 1836. Sur les organismes inferieurs. — Annales de Sciences naturelles (Zoologie) 5:193- 205. Evans, F. R., & J. C. Thompson, Jr. 1964. Pseudo- cohnilembus n. fam., a Hymenostome ciliate family containing one genus, Pseudocohnilem- 237 bus n. g., with three new species.—Journal of Protozoology 11:344-352. ——,, & J. O. Corliss. 1964. Morphogenesis in the hymenostome ciliate Pseudocohnilembus per- salinus and its taxonomic and phylogenetic im- plications.—Journal of Protozoology 11:353- 370. Fernandez-Leborans, G., & M. Castro de Zaldumbide. 1984. Morphology of three species of free-liv- ing marine ciliata: Pseudocohnilembus canta- bricus n. sp., Paralembus asturianus n. sp., Uro- nema castellonensis n. sp. (Ciliophora, Scuticociliatida).—Archiv fiir Protistenkunde 128:159-168. ——,, & 1986a. Two new marine scutico- ciliates: Pseudocohnilembus antoniensis and Pseudoconhnilembus portuensis (Ciliophora). — Microbios 47:7—22. , & 1986b. The morphology of Ano- phrys arenicola sp. nov. (Ciliophora, Scutico- ciliatida).—Journal of Natural History 20:713- 721. Foissner, W. 1979. Okologische und Systematische Studien tiber das Neuston alpiner Kleingewas- ser, mit besonderer Berticksichtigung der Cil- iaten.—International Revue der gesammten Hydrobiologie 64:99-140. 1983. Taxonomische Studien tiber die Cil- iaten des GroBglocknergebietes (Hohe Tauern, Osterreich).— Annales der Naturhistorische Mu- setim Wien 84:49-85. 1984. Morphologie und Infraciliatur einiger limnischer Ciliaten (Protozoa: Ciliophora).— Schweizerische Zeitschrift fir Hydrobiologie 46: 210-223. 1985. Die Morphologie und die Infraciliatur einiger limnischer Ciliaten (Protozoa: Cilioph- ora) aus dem Darm der Seeigel (Euechinoidea) Paracentrotus lividus und Arbacia lixula. — Ar- chiv flir Protistenkunde 130:355-366. ——,, & N. Wilbert. 1981. A comparative study of the infraciliature and silverline system of the freshwater scuticociliates Pseudocohnilembus putrinus (Kahl, 1928) nov. comb., P. pusillus (Quennerstedt, 1869) nov. comb., and the ma- rine form P. marinus Thompson, 1966.—Jour- nal of Protozoology 28:291-297. Groliére,C. A. 1977. Contribution a l’étude des ciliés des sphaignes et des étendues d’eau acides. I. Description de quelques espéces de gymno- stomes, hypostomes hymenostomes et hetero- triches.—Annales Station Biologie Besse-en- Chandesse 10:265-297. , & R. Detcheva. 1974. Description et sto- matogénese de Pleuronema puytoracin. sp. (Cil- iata, Holotricha).—Protistologica 10:91-99. 238 Kahl, A. 1926. Neue und wenig bekannte Formen der holotrichen und heterotrichen Ciliaten.— Archiv fiir Protistenkunde 55:197—438. 1927. Neue und erganzende Beobachtungen holotricher Ciliaten. I.—Archiv fiir Protisten- kunden 60:34-129. . 1935. Urtiere order Protozoa. I. Wimpertiere order Ciliata (Infusoria). In F. Dahl, Die Tier- welt Deutschlands, G. Fischer, Jena. Kent, W. S. 1881. A manual of the infusoria. D. Bogue, London, 913 pp. Lynn, D. H. 1988. Cytoterminology of cortical com- ponents of ciliates: somatic and oral kinetids. — BioSystems 21:299-307. Martin-Gonzalez, A., S. Serrano, & D. Fernandez-Ga- liano. 1985. Urotricha vitrea n. sp. (Cilioph- ora, Prorodontina): general morphology and cy- tological events during the conjugation process. — Canadian Journal of Zoology 63:1885-1891. Munoz, A., C. Tellez, & D. Fernandez-Galiano. 1987. Morphology and infraciliature in Urotricha nais sp. nov. Urotricha castalia sp. n. (Ciliophora, Prorodontina).—Acta Protozoologica 26:197-— 204. ,& 1989. Description of the infraciliature and morphogenesis in the ciliate Urotricha ordina n. sp. (Prorodontida, Urotri- chidae).—Journal of Protozoology 36:104-109. Patsch, B. 1974. Die Aufwuchsciliaten des Natur- lehrparks haus Wildenrath.— Dissertation Mathematical- Natural Fakul., University of Bonn. Schewiakoff, W. 1896. Beitrage zur Kenntniss der holotrichen Ciliaten.—Bibliographie Zoolo- gische 5:1-77. Small E. B. 1967. The Scuticociliatida, a new order of the class Ciliatea (Phylum protozoa, Subphy- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON lum Ciliophora).—Transactions of the Ameri- can Microscopical Society 86: 345-370. —,&G.A.Antipa. 1978. Cortical differentiation in Pleurocoptes furgasoni n. sp., thigmotactic pleuronematine scuticociliate epizoic on the bat starfish.— Transactions of the American Micro- scopical Society 97:540-548. ——, & D.H.Lynn. 1985. Phylum Ciliophora. Pp. 393-575 in J. J. Lee, S. H. Hutner, & E. C. Bovee, eds., Illustrated guide to the protozoa. Society of Protozoologists, Allen Press, Law- rence, Kansas. Song, W., & N. Wilbert. 1989. Taxonomische Un- tersuchungen an Aufwuchsciliaten (Protozoa, Ciliophora) im Poppelsdorfer Weiher, Bonn.— Lauterbornia H. 3:2-221. Stokes, A. C. 1986. A preliminary contribution to- wards a history of the freshwater infusoria of the United States.— Journal of the Trenton Nat- ural History Society 1:71-344. Thompson, J.C.,Jr. 1966a. Pseudocohnilembus mar- inus n. sp., a hymenostome ciliate from the Vir- ginia coast.—Journal of Protozoology 13:463— 465. 1966b. Potomacus pottsi n. g., n. sp., a hy- menostome ciliate from the Potomac river.— Journal of Protozoology 13:459-462. Wilbert, N. 1986. Beitrag zur Morphologie und 6ko- logie einiger planktischer Ciliaten aus dem Pop- pelsdorfer Weiher in Bonn (BRD).—Archiv fir Protistenkunde 131:59-69. Laboratorio Biologia General, Departa- mento de Biologia Animal I (Zoologia), Fa- cultad de Biologia, Universidad Complu- tense, 28040 Madrid, Spain. PROC. BIOL. SOC. WASH. 107(2), 1994, pp. 239-255 MORPHOLOGICAL VARIABILITY IN WARM-TEMPERATE AND SUBTROPICAL POPULATIONS OF MACRODASYS (GASTROTRICHA: MACRODASYIDA: MACRODASYIDAE) WITH THE DESCRIPTION OF SEVEN NEW SPECIES Wayne A. Evans Abstract.—Seven new species of Macrodasys, M. ancocytalis, M. achrado- cytalis, M. deltocytalis, M. meristocytalis, M. dolichocytalis, M. blysocytalis and M. stenocytalis (Gastrotricha: Macrodasyida: Macrodasyidae) are described from warm temperate and subtropical marine sediments along the Atlantic and Gulf coasts of Florida, U.S.A. Multivariate analyses of morphometric char- acters reveal that the shape and size of the reproductive organs are the most useful characters for discriminating among species. The numerically abundant species M. achradocytalis and M. meristocytalis displayed the widest geographic range. High morphological variability within Macrodasys populations at certain locations can be attributed to the presence of two or more species. During previous investigations of the ma- rine gastrotrich fauna of warm temperate and subtropical sandy sediments along the Atlantic and Gulf coasts of Florida, U.S.A (Evans 1992), several populations belong- ing to the genus Macrodasys were observed that exhibited differences in both internal and external morphology. Subsequently, I sampled nine locations in southern Florida (Fig. 1): Honeymoon Island (three loca- tions), Crandon Park on Key Biscayne (two locations), Sombrero Beach on Vaca Key (one location), Bahia Honda Key (two lo- cations), and Key West (one location). Vari- ability of morphology of the forms within Macrodasys populations at each location was quantified and overall variability within populations was used to discriminate among forms, to identify critical taxonomic char- acters, and to assess the taxonomic status of each form. Particular attention was paid to the size and shape of the reproductive organs. Members of the genus Macrodasys are si- multaneous hermaphrodites with a repro- ductive system that consists of paired testes with vasa deferentia, a single ovary, a fron- tal organ which receives and stores allo- sperm, and a caudal organ which gathers autosperm and passes them to the partner during copulation. The frontal organ con- sists of an anterior seminal receptacle and a posterior spermatheca. The caudal organ comprises an anterior glandulomuscular structure and a posterior glandular sac (“‘an- trum feminum” of Remane 1924). Ruppert (1978) provides a histological account of the functional anatomy of the reproductive sys- tem in two undescribed species of Macro- dasys. Materials and Methods Littoral and sublittoral sediments were collected at each location in January, 1992 with a hand-held piston corer. The gastro- trichs were extracted by means of serial de- cantation with isosmotic MgCl, (see Evans & Hummon 1991). Specimens were located at 30x under a stereomicroscope, mounted on glass slides, and observed under No- marski differential interference contrast op- 240 ot e*° 0.6). Character loadings based on standardized- canonical-variate scores are interpreted as follows: CV 1—shape of the middle portion of the glandulomuscular structure of the caudal organ, CV2—relative lengths of the anterior and middle portions of the glan- dulomuscular structure and the presence of an auxiliary chamber on the seminal recep- tacle of the frontal organ. A 3rd canonical variate (not plotted) explained an additional 3.9% of the variation and is interpreted as the size and shape of the seminal receptacle. The seven forms are given species status and formally described below. Taxonomy Order Macrodasyida Rao & Clausen, 1970 Family Macrodasyidae Remane, 1924 Macrodasys Remane, 1924 Macrodasys ancocytalis, new species Fig. 4 Holotype.— Adult specimen 650 um in length, mounted on glass slide. Honeymoon Island, Florida, U.S.A., sand spit facing the 241 VOLUME 107, NUMBER 2 “soqny DAISOUPR JOLIOIUE JO ¢—] SP[et{—Cd ‘ZA ‘TA {UesIO [eUOY Jo aJouldooo1 [euIUIES— YS ‘UeBIO [epNed Jo oINjonI}s Je|NosNULO[NpUL]s—SWH ‘suoneIAsiggy ‘odueI oY} SB SIQIOVILYO ONSLIOUI ‘UONVIASP piepue]s F ULOUT 9Y} Se UOATS ore SIOJOBIY OINI] ‘SAsMpOIIDP JO SO1dIdS UAAAS JO SIO}OBILYS SIVISHOUI Pue INIA, “7 “SIY 0 0 0 0 0 (4 0 SMOY 9GN] SATSOUPY resi09 | 0 c 0 4 @ 0 (4 SMOY OGM] PATSOYPY Te1{USA 0 0 0 0 0 @= i 0 C4 Sqn], SATSeYsy TOT 19} UY 0 S = G= 1 0 0 a= 0 2d SOQN] SATSOYPY FOTI9,UY v1 - CL Ciaaee lL = 9 6-79 8-9 8-G = Td Sqn] SATSOYpY 1OT 197 UY CL +9 L2 89 + G LZ Oe + 8°61 Ld + 2° £2 Ge + 9T¢ Lette p°0 + 9°08 YIPTM UOTI10d STPPTN SHO 8 IL + G°8S Cpl + 8°69 £8 + 069 6°6 + 0°85 97 +915 6°¢CL + £86 9°0 + 1'eS Yybuey uot}10d STPPTW SHO Ge +6 FL QC FE OL 81 + 7 LT Ve #0 01 91 + 6 II 1?@ + O81 L°0 + 161 YIPTM UOTII0g FOTI9UY SHO pL t7 GS GLI + Te Lb + 8°19 DIL + € Sh 6°G + 7'8E $8 + 9°06 y'€ FG°€8 | YybueT uoT{I0g JOTIeqUY SND 0 pe + 2°82 0 rs = LO 0 0 0 ygbuey requeyg Arosseooy YS Lt + 9°02 0°01 + 9°7E 1 + C1 v0 + 8°12 L°? + £81 6€ + G92 6°0 + 2c? UIPIM eToe}deoey TeUTUAS 18 + 7°79 9°6 + 2 OL Cv +679 Ll + 9°85 Ona aec po + 6 UE 87 + £29 ygbuey atoe}desey TeuTuas CLEE TL LL 0S 09 + 0SH COL + 0°85 99 + L°6% C Il # GEL C6 + 19 S}Td UOJSTd J YIPTM Peo T1S6 +9699 cG&l + & T&L 9°Pl + 2069 9°GL + 1 1E9 16S + 6699 1°98 + 6°88 Cle + § 129 dnpy Jo yybuey Te,OL (¢ = u) (¢ = u) (¢ = u) (oh = u) (¢ = u) (¢ = u) STTeyAooueys STpTeyAOOSATG STTeyooyOTTop Stpeyfooystreu StpeyAo0jTep stpeyAoopeyoe StTTe Adu jUuaWeINseay STSEPOTIR 242 Canonical Variate 1 -15 -10 -5 0 5 Canonical Variate 2 Fig. 3. Canonical variate (CV) plots of Macrodasys species, based on 14 characters. CV1 and CV2 are interpreted as the size and shape of the glandulomus- cular structure of the caudal organ and the presence of an accessory chamber on the seminal receptacle of the frontal organ. Solid circles—M. ancocytalis, open cir- cles— M. achradocytalis, solid squares— M. deltocytal- is, open squares— MM. meristocytalis, solid triangles— M. dolichocytalis, open triangles— M. blysocytalis, and X—WM. stenocytalis. Gulf of Mexico [28°05'N, 82°50’W]. USNM 168056. Etymology. —anco (L.) meaning curved, after the shape of the seminal receptacle; cyto (L.) meaning chamber or receptacle; alis meaning possession. Diagnosis. —Macrodasys with trunk much longer than pharyngeal region. Lateral and ventral adhesive tubes present on trunk; dorsal tubes lacking. Anterior adhesive tubes ventral, a single row of from five to seven tubes on each side of body, near anterior margin of head. Frontal organ a small sper- matheca and elongate seminal receptacle with small nozzle; accessory chamber ab- sent. Glandulomuscular structure of caudal organ with anterior glandular portion longer than middle portion bearing circular mus- culature; posterior portion a small, curved “neck” with terminal pore; copulatory tube : without branches. Glandular sac of caudal organ a rounded, triangular shape. Description. —Elongate, strap-shaped body; adults 602-685 um long, 49-65 um wide. Caudum tapers into long, narrow PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON “tail” (Fig. 4a). Dorsal and lateral body sur- faces covered with long sensory bristles. Pis- ton pits (“stempelgrube”’ of Remane 1924) on each side of head. Ventral surface en- tirely covered with locomotor cilia except small, bare area around female pore (Fig. 4b). Dorsal ciliary band present on head. Anterior adhesive tubes a ventral, single row, an arc of from five to seven tubes on each side of body, adjacent to mouth (Fig. 4c); most medial tube shortest, most lateral lon- gest. About 16 lateral adhesive tubes on each side of trunk; an additional 10-12 tubes on each side of tail. Two ventral rows of ad- hesive tubes near lateral margins of trunk, about 16 per row. Adhesive tubes on trunk begin just anterior to pharyngeal-intestinal junction; lateral and ventral rows merge where trunk narrows into tail. Mouth leads into buccal cavity, which opens into pharynx. Pharyngeal-intestinal junction at U40 (UO, anterior-most tip; U100, posterior-most tip; in the terminol- ogy of Schoepfer-Sterrer 1969); pharyngeal pores (U24) small. Intestine narrows con- tinuously from pharyngeal-intestinal junc- tion to terminus; anus ventral. Small, paired lateral testes at pharyngeal- intestinal junction taper into vasa deferen- tia; male pores separate and ventral, adja- cent to frontal organ (Fig. 4a). Frontal organ an anterior seminal receptacle and posterior spermatheca (Fig. 4d). Seminal receptacle with weak circular musculature and small, lightly cuticularized anterior pore (nozzle); length about three times width. Spermathe- ca hollow with few secretory droplets. Ova- ry adjacent to spermatheca; ova increase in size anteriorly. Ovum adjacent to nozzle of spermatheca receives sperm. Large caudal organ a spindle-shaped glandulomuscular structure and a glandular sac with opening to ventral surface (Fig. 4e); anterior tip of glandulomuscular structure adjacent to spermatheca. Anterior portion of glandulomuscular structure longer than middle portion and sheathed in longitudinal muscles. Middle VOLUME 107, NUMBER 2 243 TbA Spe Fig. 4. MM. ancocytalis, new species. Single row of anterior adhesive tubes. Ventral tubes present. Curved seminal receptacle of frontal organ lacks an auxiliary chamber. Anterior portion of glandulomuscular structure of caudal organ longer than middle portion. a) internal organs, b) ventral surface, c) ventral and dorsal head, d) frontal organ, and e) caudal organ. c-e not to scale. Abbreviations: BrS—sensory bristle, CiD—dorsal ciliary band, CiV—ventral locomotor cilia, Co—caudal organ, Eg—egg, Fo—frontal organ, GrR—refractile granules, Gs—glandular sac, MuC—circular muscles, MuL—longitudinal muscles, Nz—nozzle, Ov—ovum, Ph—pharynx, PoF—female reproductive pore, POM—male reproductive pore, Pp—pharyngeal pore, PtP—piston pit, Sdp— secretory droplet, Smr—seminal receptacle, Spm—spermatozoa, SpT—spermatheca, TbA—anterior adhesive tube, TbD—dorsal adhesive tube, TbL—lateral adhesive tube, TbV—ventral adhesive tube, Ts—testis. portion with longitudinal muscles on ven- tral side only, but entirely sheathed in ro- bust circular muscles. Posterior portion of glandulomuscular structure an angled neck that extends into center of glandular sac. Glandulomuscular structure filled with re- fractile granules of various diameters; un- branched copulatory tube in middle portion of glandulomuscular structure leads to opening in neck. Glandular sac of caudal organ a rounded, triangular shape. Distribution and habitat.—Uncommon species, found in littoral and sublittoral zones in coarse, poorly-sorted sediments with both siliceous and carbonate fractions. Honeymoon Island (two locations). Macrodasys achradocytalis, new species JENS D) Holotype.— Adult specimen 825 um in length, mounted on glass slide. Bahia Hon- da Key, Florida, U.S.A., sandy beach facing the Atlantic Ocean [24°38'N, 81°35’W]. USNM 168055. Etymology. —achrado (L.) meaning wild pear, after the shape of the seminal recep- tacle; cyto (L.) meaning chamber or recep- tacle; alis (L.) meaning possession. 244 TbL Fig. 5. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON M. achradocytalis, new species. Dorsal adhesive tubes in two rows of 10 tubes each. Anterior adhesive tubes in three fields. Seminal receptacle and spermatheca of frontal organ of similar size. Anterior portion of glandulomuscular structure of caudal organ much shorter than middle portion; neck long and narrow. Abbre- viations and a-< as in Fig. 4; c-e not to scale. Diagnosis. —Macrodasys with trunk much longer than pharyngeal region. Lateral and dorsal adhesive tubes present on trunk; ven- tral tubes lacking. Anterior adhesive tubes ventral in three fields on each side of body, near anterior margin of head. Anterior field a row of from five to eight tubes arranged in two distinct groups at 95° angle to each other; middle field a row with from two to four tubes; posterior-most field with one or two tubes. Frontal organ a rounded sper- matheca and pear-shaped seminal recepta- cle; nearly equal in size. Seminal receptacle sheathed in circular muscles; nozzle large and cuticularized; accessory chamber ab- sent. Anterior, glandular portion of glan- dulomuscular structure of caudal organ short and narrow, about one-half length of middle portion; neck long and slightly curved; cop- ulatory tube without branches. Glandular sac of caudal organ oval. Description. —Very long strap-shaped body; adults 661-1033 wm long and 54-97 um wide. Caudum narrows abruptly into short tail (Fig. 5a). Dorsal and lateral body surfaces covered with long sensory bristles. Ventral surface entirely covered with loco- motor cilia except large, bare area around VOLUME 107, NUMBER 2 female pore (Fig. 5b). Sparse dorsal ciliary band on head. Anterior adhesive tubes ven- tral, in three fields on each side of body just posterior to mouth (Fig. 5c). Anterior field with row of seven tubes in two distinct groups at about 95° angle to each other; tubes increase in length from medial to lateral. Middle field a transverse row with from two to four tubes. Posterior-most field with one or two tubes. About 40 lateral adhesive tubes on each side of trunk; tubes longest where trunk narrows into tail. An additional five or six tubes on each side of tail. Two dorsal rows of about 10 small tubes each (Fig. 5a). Ventral adhesive tubes lacking. Adhesive tubes on trunk begin just anterior to pha- ryngeal-intestinal junction. Lateral tubes continue onto tail; dorsal tube rows end just anterior to tail. Mouth leads into large buccal cavity, which opens into pharynx. Pharyngeal-in- testinal junction at U35; pharyngeal pores at U23. Intestine narrows toward caudal end of trunk; anus ventral. Paired lateral testes at pharyngeal-intes- tinal junction taper into vasa deferentia; male pores separate and ventral, adjacent to seminal receptacle. Spermatheca of fron- tal organ rounded with internal cavity and large secretory droplets. Seminal receptacle of frontal organ pear-shaped, anterior end with large, heavily cuticularized D-shaped nozzle; accessory chamber lacking. Seminal receptacle sheathed in circular muscles (Fig. 5d). Small tube leads from posterior ventral portion of seminal receptacle to ventral fe- male pore. Spermatheca and seminal recep- tacle approximately the same size; secretory droplets often large. Ovary lies to right of frontal organ; ova increase in size anteri- orly. Glandulomuscular structure of caudal organ with anterior portion short (one-third the length of middle portion) and narrow. Long, narrow neck of glandulomuscular structure extends to posterior end of glan- dular sac; copulatory tube unbranched. Re- fractile granules densest and smallest in pos- 245 terior end of glandulomuscular structure (Fig. Se). Glandular sac of frontal organ oval. Distribution and habitat. Common spe- cies, found in littoral and sublittoral zones in both coarse, poorly-sorted carbonate sed- iments and medium-fine siliceous sedi- ments. Abundant where found. Crandon Park (two locations), Bahia Honda (two lo- cations), and Sombrero Beach. Macrodasys deltocytalis, new species Fig. 6 Holotype.— Adult specimen 670 um in length, mounted on glass slide. Crandon Park, Florida, U.S.A., small, sandy beach facing harbor on Biscayne Bay [25°44'N, 80°10'W]. USNM 168058. Etymology. —delto (L.) meaning in the shape of a triangle, after the shape of the seminal receptacle; cyto (L.) meaning cham- ber or receptacle; a/is (L.) meaning posses- sion. Diagnosis. —Macrodasys with pharynx and trunk of about equal length. Lateral and ventral adhesive tubes present on trunk; dorsal tubes lacking. Anterior adhesive tubes ventral, in one row of from six to eight tubes on each side of body, near anterior margin of head. Tubes arranged in arc with shortest tubes medially and longest tubes laterally. Spermatheca of frontal organ simple, round; seminal receptacle triangular-shaped, twice as long as diameter of spermatheca and without visible circular musculature; acces- sory chamber absent. Anterior portion of glandulomuscular structure of caudal organ wide and about two-thirds length of middle portion; neck strongly curved; copulatory tube without branches. Glandular sac of caudal organ small. Description. —Strap-shaped body; adults 602-685 um long, 49-65 um wide. Caudum tapers quickly into medium-length tail (Fig. 6a). Dorsal and lateral body surfaces sparse- ly covered with long sensory bristles. Ven- tral surface entirely covered with locomotor cilia except small, bare area around female PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 246 Mo ThA gee il i th CiV i i : TbL AI ih 4 ‘i TbV A PoF fe} PoM — il i) Bi BS ral Fig. 6. M. deltocytalis, new species. Pharyngeal region and trunk equal in length. Anterior adhesive tubes in a single row of eight tubes. Ventral adhesive tubes present. Seminal receptacle of frontal organ triangular in shape. Abbreviations and a- as in Fig. 4; c-e not to scale. pore (Fig. 6b). Sparse dorsal ciliary band on head near anterior margin. Anterior adhe- sive tubes a single ventral row of from six to eight tubes on each side of body near anterior margin of head and arranged in an arc with tubes increasing in length from me- dial to lateral (Fig. 6c). About 10 lateral adhesive tubes on each side of trunk; ad- ditional seven or eight tubes on each side of tail. Lateral tubes longest in mid-trunk region. Two ventral rows of about seven tubes each. Adhesive tubes on trunk begin just anterior to pharyngeal-intestinal junc- tion. Lateral and ventral rows converge where trunk narrows into tail. Mouth leads into shallow buccal cavity, which opens into long pharynx. Pharyngeal- intestinal junction at USO; pharyngeal pores at U30. Intestine narrows at caudal end of trunk; anus ventral. Small, paired lateral testes at pharyngeal- intestinal junction taper into vasa deferen- tia; male pores separate and ventral, adja- cent to posterior edge of spermatheca. Sper- matheca of frontal organ rounded, with large internal cavity; secretory droplets not ob- served. Seminal receptacle of frontal organ triangular-shaped with small nozzle at an- terior apex; thick-walled tubular chamber leads to ventral female pore from lateral apex. Ventral pore surround by epidermal sculpturing. Right wall of seminal recepta- VOLUME 107, NUMBER 2 CiV 100um Fig. 7. 247 Spm = Sane M. meristocytalis, new species. Ventral adhesive tubes present. Seminal receptacle of frontal organ with a small accessory chamber containing female pore. Testes large. Anterior portion of glandulomuscular structure of caudal organ about one-half as long as middle portion. Abbreviations and a-« as in Fig. 4; c-e not to scale. cle thickened: circular muscles not evident. Accessory chamber lacking. Seminal recep- tacle approximately 2.5 times as long as di- ameter of spermatheca. Ovary lies posterior and right of frontal organ; ova increase in size anteriorly. Glandulomuscular structure of caudal organ with anterior portion broad and about two-thirds as long as middle por- tion (Fig. 6e). Short, severely curved neck of glandulomuscular structure extends to left margin of glandular sac; copulatory tube un- branched. Refractile granules present from anterior end of copulatory tube to anterior tip of glandulomuscular structure. Glan- dular sac of caudal organ small. Distribution and habitat.—Uncommon species, found only in littoral zone of small beach facing harbor on Biscayne Bay. Well- sorted, medium-fine siliceous sediments. Crandon Park (one location). Macrodasys meristocytalis, new species Fig. 7 Holotype. — Adult specimen 670 um in length, mounted on glass slide. Key West, Florida, U.S.A., small sandy beach facing Atlantic Ocean [24°35’N, 81°50’W]. USNM 168060. Etymology. — meristo (L.) meaning divid- ed, after the division of the seminal recep- tacle into two chambers; cyto (L.) meaning chamber or receptacle; a/is (L.) meaning possession. Diagnosis. — Macrodasys with trunk slightly longer than pharyngeal region. Lateral and 248 ventral adhesive tubes present on trunk; dorsal tubes lacking. Anterior adhesive tubes ventral, a single row of from four to nine tubes on each side of body, near anterior margin of head. Testes very large. Frontal organ a simple, round spermatheca and elongate seminal receptacle. Seminal recep- tacle 2.5 times as long as diameter of sper- matheca and without visible circular mus- culature; accessory chamber present. Anterior portion of glandulomuscular structure of caudal organ broad and short, about two-thirds length of middle portion; neck very short; copulatory tube un- branched. Glandular sac of caudal organ oval. Description. —Strap-shaped body; adults 495-797 um long, 38-89 wm wide. Caudum tapers gradually into narrow tail (Fig. 6a). Dorsal and lateral body surfaces covered with numerous sensory bristles. Ventral surface entirely covered with locomotor cil- la except a V-shaped, bare area posterior to anterior series of adhesive tubes and a large, bare area surrounding female pore (Fig. 7b). Sparse dorsal ciliary band on head at level of sensory pits. Anterior adhesive tubes ventral, a single row of from four to nine tubes on each side of body near anterior margin of head, arranged in an arc with tubes increasing in length from medial to lateral (Fig. 7c). About 20 lateral adhesive tubes on each side of trunk; an additional eight or nine tubes on each side of tail. Lateral tubes longest where trunk narrows into tail. Two ventral rows of about 13 small tubes each. Adhesive tubes on trunk begin just anterior to pharyngeal-intestinal junction. Lateral and ventral rows converge where trunk narrows into tail. Mouth leads into shallow buccal cavity, which opens into long pharynx. Pharyngeal- intestinal junction at U41; well-developed pharyngeal pores at U26. Intestine narrows at caudal end of trunk; anus ventral. Very large, paired lateral testes at pha- ryngeal-intestinal junction taper into vasa deferentia; male pores separate and ventral, PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON adjacent to seminal receptacle of frontal or- gan. Spermatheca of frontal organ rounded with dual internal cavities; secretory drop- lets present. Seminal receptacle of frontal organ a rounded cone, with bulbous acces- sory chamber on right side; small nozzle present at anterior apex (Fig. 7d). Thin- walled, cylindrical tube leads from acces- sory chamber to simple ventral female pore. Seminal receptacle approximately 2.5 as long as the diameter of spermatheca;: circular muscles not visible. Ovary lies posterior to and left of frontal organ; ova increase in size anteriorly with largest ovum in front of sem- inal receptacle. Glandulomuscular structure of caudal organ with broad anterior portion that tapers to rounded point anteriorly, about three-quarters as long as middle por- tion. Stubby, slightly-curved neck of glan- dulomuscular structure extends to center of glandular sac; refractile granules sparse; copulatory tube unbranched. Glandular sac of caudal organ oval. Distribution and habitat. —Common spe- cies, found in littoral and sublittoral zones in coarse, poorly-sorted carbonate sedi- ments and in medium-fine, siliceous sedi- ments. Abundant where found. Honey- moon Island (one location), Crandon Park (two locations), Bahia Honda (one loca- tion), and Key West. Macrodasys dolichocytalis, new species Fig. 8 Holotype.— Adult specimen 690 um in length, mounted on glass slide. Honeymoon Island, Florida, U.S.A., beach facing the Gulf of Mexico near causeway [28°05'N, 82°50'W]. USNM 168059. Etymology. —dolicho (L.) meaning elon- gate, after the shape of the seminal recep- tacle; cyto (L.) meaning chamber or recep- tacle; alis (L.) meaning possession. Diagnosis. —Macrodasys with trunk lon- ger than pharyngeal region. Lateral adhesive tubes present on trunk; ventral and dorsal tubes lacking. Anterior adhesive tubes ven- VOLUME 107, NUMBER 2 100um 249 Fig. 8. MM. dolichocytalis, new species. Lateral adhesive tubes abundant, especially in tail region. Anterior adhesive tubes in two fields. Seminal receptacle of frontal organ about six times as long as wide. Sperm with very long tails. Anterior portion of glandulomuscular structure of caudal organ slightly longer than middle portion. Abbreviations and a-« as in Fig. 4; c-e not to scale. tral, in two fields on each side of body, near anterior margin of head. Anterior field a row of six or seven tubes of equal size in two distinct groups of three or four tubes each, at about 130° angle to each other. Second field with one or two tubes. Seminal recep- tacle of frontal organ long and narrow, about six times as long as wide, with strong cir- cular musculature; accessory chamber lack- ing. Spermatheca of frontal organ small. Anterior portion of glandulomuscular structure of caudal organ nearly as wide as, and slightly longer than, middle portion; neck wide; copulatory tube without branch- es. Glandular sac of caudal organ ovoid. Description. —Strap-shaped body; adults 680-701 wm long, 41-49 um wide. Caudum tapers gradually into wide tail. Dorsal and lateral body surfaces densely covered with long sensory bristles (Fig. 8a).Ventral sur- face entirely covered with locomotor cilia except small, bare area posterior to anterior series of adhesive tubes and bare, diamond- shaped area surrounding female pore (Fig. 8b). Dorsal ciliary band on head at level of sensory pits. Anterior adhesive tubes ven- tral in two fields, on each side of body, near anterior margin of head (Fig. 8c). Anterior field a row of six or seven tubes of equal size in two distinct groups of three or four each, at about 130° angle to each other. Sec- ond field with one or two tubes. No anterior 250 tubes reach level of sensory pits. About 26 lateral adhesive tubes on each side of trunk; an additional 10 tubes on each side of tail. Lateral tubes slightly longer near caudum. Adhesive tubes on trunk begin just anterior to pharyngeal-intestinal junction. Mouth leads into small buccal cavity, which opens into pharynx. Pharyngeal-in- testinal junction at U35; pharyngeal pores at U23. Intestine narrows in mid-trunk re- gion; anus ventral. Round, paired lateral testes at pharyn- geal-intestinal junction taper into vasa de- ferentia; male pores separate and ventral, adjacent to seminal receptacle of frontal or- gan. Spermatheca of frontal organ small and irregular in shape, without secretory drop- lets. Seminal receptacle long and narrow, six times longer than wide with rounded apex; very small nozzle present at apex (Fig. 8d). Thin-walled tube leads from posterior end of seminal receptacle to simple ventral fe- male pore. Seminal receptacle approxi- mately five times as long as the diameter of spermatheca; circular muscle bands pres- ent. Ovary not observed, large ovum in front of seminal receptacle. Glandulomuscular structure of caudal organ with anterior por- tion long and broad with broadly rounded apex, about same length as middle portion; refractile granules sparse; copulatory tube unbranched. Stubby, curved neck of glan- dulomuscular structure extends to left side of glandular sac of caudal organ. Glandular sac ovoid. Distribution and habitat.—Uncommon species, found only in sublittoral zone of small beach facing Gulf of Mexico. Coarse, poorly-sorted siliceous/carbonate mixed sediments. Honeymoon Island (one loca- tion). Macrodasys blysocytalis, new species Fig. 9 Holotype.—Adult specimen 780 um in length, mounted on glass slide. Honeymoon Island, Florida, U.S.A., inside of sand spit PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON beach facing St. Joseph Sound [28°05’N, 82°50’'W]. USNM 168057. Etymology. —blyso (L.) meaning bubble, after the bubble-like accessory chamber on the seminal receptacle; cyto (L.) meaning chamber or receptacle; a/is (L.) meaning possession. Diagnosis. — Macrodasys with trunk much longer than pharyngeal region. Lateral and ventral adhesive tubes present; dorsal tubes lacking. Anterior adhesive tubes ventral, in two fields on each side of body, near anterior margin of head. Anterior field a row of seven or eight tubes of equal size in two distinct groups of three or four tubes each, at about 100° angle to each other. Second field a row of two or three tubes. Additional, single lat- eral adhesive tube on each side of head sep- arate from anterior series. Frontal organ a large, cone-shaped seminal receptacle with large, ovoid dorsal accessory chamber and a thick-walled spermatheca, about same size as seminal receptacle. Anterior portion of glandulomuscular structure of caudal organ broad with rounded tip, about half as long as middle portion; neck short; copulatory tube T-shaped. Glandular sac of caudal or- gan small. Description. —Strap-shaped body; adults 678-784 wm long and 53-56 um wide. Caudum tapers gradually into indistinct tail (Fig. 9a). Dorsal and lateral body surfaces covered with sensory bristles. Ventral sur- face entirely covered with locomotor cilia except small, bare area surrounding female pore (Fig. 9b). Sparse dorsal ciliary band on head at level of sensory pits. Anterior ad- hesive tubes ventral, in two fields on each side of body, near anterior margin of head (Fig. 9c). Anterior field a row of seven or eight tubes in two distinct groups of three or four tube each, at about 100° angle to each other; tubes in medial group one-half as long as tubes in lateral group. Posterior field a row of two or three tubes about 1.5 times as long as tubes in anterior field. Ad- ditional, single ventrolateral adhesive tube on each side of head separate from usual VOLUME 107, NUMBER 2 at Me S) Pp 251 Fig. 9. M. blysocytalis, new species. Caudum tapers gradually into indistinct tail. Anterior adhesive tubes in three fields; additional pair of tubes laterally in head region. Seminal receptacle of frontal organ with accessory chamber in the form of a large, ventral blister. Copulatory tube of glandulomuscular structure of caudal organ branched. Abbreviations and a-< as in Fig. 4; c-e not to scale. anterior series. About 15 lateral adhesive tubes on each side of trunk; additional six or seven tubes on each side of tail. Aside from single pair of tubes near anterior series, lateral tubes begin nearly at level of pha- ryngeal pores and are of uniform size. Two ventral rows of adhesive tubes begin pos- terior to second pair of lateral tubes and are much smaller than lateral tubes; about 10 ventral tubes per row. Lateral and ventral rows begin to converge on posterior third of trunk; ventral rows stop where tail begins. Anterior mouth leads into buccal cavity, which opens into pharynx. Pharyngeal-in- testinal junction at U30; pharyngeal pores at U21. Intestine narrows abruptly in mid- trunk region; anus ventral. Elongate, paired lateral testes at pharyn- geal-intestinal junction taper gradually into vasa deferentia; male pores separate and ventral, adjacent to anterior end of seminal receptacle of frontal organ. Spermatheca of frontal organ small and irregular in shape, with thick wall and large secretory droplets. Seminal receptacle cone-shaped with large, ovoid dorsal accessory chamber, about same size aS spermatheca (Fig. 9d). Small nozzle present in anterior end of seminal recepta- cle; circular muscles not evident. Female pore exits from anterior end of accessory 252 chamber of seminal receptacle. Ovary not observed; large ovum lies in front of sem- inal receptacle. Anterior portion of glan- dulomuscular structure of caudal organ wide with rounded apex, about half as long as middle portion. Copulatory tube T-shaped, branching as it enters middle portion of glandulomuscular structure (Fig. 9e); re- fractile granules larger anteriorly. Stubby, curved neck of glandulomuscular structure extends to left side of glandular sac of caudal organ. Distribution and habitat.—Uncommon species, found only in littoral zone on inside of sand spit facing St. Joseph Sound. Me- dium-fine, well-sorted siliceous sediments. Honeymoon Island (one location). Macrodasys stenocytalis, new species Fig. 10 Holotype. —Adult specimen 670 um in length, mounted on glass slide. Sombrero Beach, Vaca Key, Florida, U.S.A., small beach facing Atlantic Ocean [24°41'N, 81°0S5’W]. USNM 168061. Etymology. —steno (L.) meaning narrow or constricted, after the constriction of the seminal receptacle; cyto (L.) meaning cham- ber or receptacle; a/lis (L.) meaning posses- sion. Diagnosis. — Macrodasys with trunk about same length as pharyngeal region. Lateral adhesive tubes present; ventral and dorsal tubes lacking. Anterior adhesive tubes ven- tral, in single row of from 12 to 14 tubes on each side of body, near anterior margin of head. Tubes arranged in arc with shortest medial and longest lateral. Frontal organ a seminal receptacle with muscularized bulb anterior to narrow constriction and an ovoid spermatheca about one-half as long as sem- inal receptacle. Anterior portion of glan- dulomuscular structure of caudal organ broad, both with narrow extension to tip, slightly more than one-half as long as mid- dle portion; copulatory tube without PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON branches; refractile granules sparse. Glan- dular sac of caudal organ small. Description. —Broad, strap-shaped body; adults 661-680 um long, 67-74 um wide. Caudum tapers abruptly into short tail (Fig. 10a). Dorsal and lateral body surfaces cov- ered with long sensory bristles. Ventral sur- face entirely covered with locomotor cilia except small, bare area surrounding female pore (Fig 10b). Sparse dorsal ciliary band on head at level of sensory pits. Anterior adhesive tubes ventral, in single row of from 12 to 14 tubes on each side of body, near anterior margin of head (Fig. 10c). Medial tubes shortest, lateral tubes longest. About 24 lateral adhesive tubes on each side of trunk; an additional six to eight tubes on each side of tail. Lateral tubes begin just anterior to pharyngeal-intestinal junction and are longest and most numerous where trunk constricts to form tail. Mouth leads into buccal cavity, which opens into the pharynx. Pharyngeal-intes- tinal junction at U45; pharyngeal pores at U30. Intestine gradually narrows toward tail; anus ventral. Paired lateral testes at pharyngeal-intes- tinal junction taper into vasa deferentia; male pores separate and ventral, adjacent to seminal receptacle of frontal organ. Sper- matheca of frontal organ large and ovoid, with various-size secretory droplets. Semi- nal receptacle with ovoid posterior section which narrows before forming a bulbous, muscularized, anterior portion; small noz- zle present (Fig. 10d). Seminal receptacle about twice as long as spermatheca. Female pore exits from posterior portion of seminal receptacle. Ovary not observed; large ovum lies in front of seminal receptacle. Anterior portion of glandulomuscular structure of caudal organ broad, but with narrow tip; slightly more than one-half as long as mid- dle portion (Fig. 10e). Middle portion broad at midsection, narrow at ends; copulatory tube straight; refractile granules present. Short, curved neck of glandulomuscular structure extends to left side of glandular VOLUME 107, NUMBER 2 ve Wy —7 mee ae ie) ond EAE FF Ho 100um 253 Fig. 10. M. stenocytalis, new species. Numerous small anterior adhesive tubes in single row. Seminal recep- tacle of frontal organ constricted in the middle; portion anterior to constriction sheathed in circular muscles. Glandular sac of caudal organ very small. Abbreviations and a-< as in Fig. 4; c—e not to scale. sac of caudal organ. Glandular sac very small. Distribution and habitat.—Uncommon species, found only in sublittoral zone on small beach facing the Atlantic Ocean. Coarse, poorly-sorted carbonate sediments. Sombrero Beach. Discussion Comparing the species described above to previously described species on the basis of reproductive morphology is difficult be- cause descriptions of the reproductive sys- tem have been either omitted or described in insufficient detail by previous workers (e.g., Boaden 1963, Ganapati & Rao 1967, Roszczak 1939, Thane-Fenchel 1970). Where details of the reproductive system are given (e.g., Remane 1936, Schmidt 1974, Valbonesi & Luporini 1984, Wieser 1957) they do not match any of the species de- scribed herein; however, M. meristocytalis does closely resemble the Macrodasys sp. II of Ruppert 1978, fig. 1c, p. 210 from Flor- ida. The reproductive morphologies of the seven Florida species described here are unique and provide an excellent basis for discriminating among species. Schoepfer- Sterrer (1974) found this to be the case with the only other genus, Urodasys, in the fam- ily Macrodasyidae. It is also possible to make correct species assignments, at least in south Florida pop- ulations of Macrodasys, on the basis of ex- ternal features such as length, the number of rows of lateral adhesive tubes, and the arrangement of the anterior tubes, when these features are used in combination. However, adhesive tubes, especially of the anterior, dorsal, and ventral series can be difficult to see without differential interfer- 254 ence contrast optics. The number of tubes in each series is hard to determine and var- ies with the age of the individual, making them an unreliable character (Luporini et al. 1973). Conversely, the reproductive or- gans, once formed, are stable in size and shape, large, and easily observed in adult specimens, even with transmitted-light mi- croscopy. They are often visible even when the specimen otherwise is in poor condition and would be the preferred characters for identifying species in this genus. What might appear to be a considerable amount of morphological variability in Ma- crodasys populations at particular geograph- ic locations is, in the Florida case, largely caused by species that are superficially sim- ilar and whose ecological distributions overlap. For example, the Crandon Park in- side-location had three species occurring to- gether in the littoral zone that contributed to the overall variability of the Macrodasys population as a whole. Morphometric anal- yses based on reproductive characters can help to resolve such variability into species- specific components. The two most abundant species (M. ach- radocytalis and M. meristocytalis) had much wider geographic distributions than the rar- er species. Each of the two occurred at five locations, in both littoral and sublittoral zones, and in a wide variety of sediment types.When found in the same beach, these two species exhibited non-overlapping hor- izontal distributions in steep, tidal beaches, but were mixed in narrow, atidal beaches. Less abundant species were found only at one, or at most two, sampling locations and were always restricted to either the littoral or sublittoral zone. Vertical overlap of spe- cies within beaches and seasonal changes in geographic and ecological distributions were not investigated. The large number of species found in a limited (on a global scale) geographical area in this and similar studies (Schmidt 1974, Valbonesi & Luporini 1984), suggests that Macrodasys is a very speciose genus. This PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON requires that claims of global distributions for members of this genus, particularly the frequently reported M. caudatus, be sup- ported by detailed morphological analyses on local and regional bases. Acknowledgments This work was completed while a Post- doctoral Fellow at Ohio University under National Science Foundation Grant BSR- 9006798 to William D. Hummon, and was supported, in part, by Ohio University Re- search Committee Grant 2335 to the author and by the Ohio University Meiofaunal Re- search Support Fund. W. D. Hummon and M. A. Todaro helped collect material for preliminary analyses and critically reviewed the manuscript. Literature Cited Boaden, P. 1963. Marine Gastrotricha from the in- terstitial fauna of some North Wales beaches. — Proceedings of the Zoological Society of London 140:485-502. Evans, W. A. 1992. Five new species of marine Gas- trotricha from the Atlantic Coast of Florida.— Bulletin of Marine Science 51:315-328. —., & W. D. Hummon. 1991. A new genus and species of Gastrotricha from the Atlantic Coast of Florida.— Transactions of the American Mi- croscopical Society 110:321-—327. Ganapati, P., & G. Rao. 1967. On some marine in- terstitial gastrotrichs from the beach sands of Waltair coast.— Proceedings of the Indian Acad- emy of Science 66B:214—225. Luporini, P., G. Magagnini, & P. Tongiorgi. 1973. Gastrotrichi macrodasioidei delle coste della Toscana.—Pubblicazioni della Stazione Zoolo- gica di Napoli 38:267—288. Rao, G. C., & C. Clausen. 1970. Planodasys margin- alis gen. et sp. nov. and Planodasyidae fam. nov. (Gastrotricha: Macrodasyoidea).—Sarsia 42:73— 82. Remane, A. 1924. Neue aberrante Gastrotrichen. I: Macrodasys buddenbrocki nov. gen. nov. spec.— Zoologischer Anzeiger 61:289-297. . 1936. Gastrotricha. Pp. 1-242 in H. G. Bronn, ed., Klassen und Ordnungen des Tierreichs Band 4. Abteilung II. Buch 1. Teil 2. Akademische Verlags gesellschaft, Leipzig. Roszezak, R. 1939. Die Psammitgastrotricha des pol- VOLUME 107, NUMBER 2 nischen Ostsee-strandes.— Zoologica Poloniae 4:1-24. Ruppert, E.E. 1978. The reproductive system of gas- trotrichs II. Insemination in Macrodasys: a unique mode of sperm transfer in metazoa.— Zoomorphologie 89:207—228. Schmidt, P. 1974. Interstitielle fauna von Galapagos IV. Gastrotricha.— Mikrofauna des Meeresbod- ens 26:1-76. Schoepfer-Sterrer, C. 1974. Five new species of Uro- dasys and remarks on the terminology of the genital organs in Macrodasyidae (Gastrotri- cha).— Cahiers de Biologie Marine 15:229-254. Thane-Fenchel, A. 1970. Interstitial gastrotrichs in 255 some south Florida beaches.— Ophelia 7:113- 138. Valbonesi, A., & P. Luporini. 1984. Researches on the coast of Somalia: Gastrotricha Macrodasy- oidea.— Monitore Zoologico Italiano, Suppl. 19: 1-34. Wieser, W. 1957. Gastrotricha Macrodasyoidea from the intertidal of Puget Sound.— Transactions of the American Microscopical Society 76:372-381. Department of Biological Sciences, Ohio University, Athens, Ohio 45701-2979, U:S.A. PROC. BIOL. SOC. WASH. . 107(2), 1994, pp. 256-261 A NEW SPECIES OF ELAPHOIDELLA (CRUSTACEA: HARPACTICOIDA) CLOSELY RELATED TO E. BIDENS (SCHMEIL) AND THE GENUS ATTHEYELLA FROM NEPAL Teruo Ishida Abstract. —Elaphoidella nepalensis, n. sp., from a small stream in Kathman- du, Nepal, is described. The new species is closely related to Elaphoidella bidens, and the genus A/theyella since the species has two setae on the basoendopodite of the male fifth leg. In a sediment sample from a small stream in the Botanical Garden of Kathmandu, the herein described new species, Elaphoidella nepalensis was found. Dr. Tomiko Ito (Hokkaido Fish Hatchery, Japan) collected the sample by scraping the bottom with a fine mesh hand net. E. nepalensis, n. sp. is in many aspects very similar with Elaphoidella bidens (Schmeil) but differs from the latter by its caudal rami and by the peculiar ornamen- tation of the male fifth leg, bearing two el- ements. Specimens were mounted in gum-chloral medium; drawings and measurements were made from the mounted specimens. Spec- imens were deposited in the U.S. National Museum of Natural History, Smithsonian Institution (USNM). Order Harpacticoida Sars, 1903 Family Canthocamptidae Brady, 1880 Genus Elaphoidella Chappuis, 1929 Elaphoidella nepalensis, new species Figs. 1-19 Material.—Holotype: female, dissected on | slide (USNM 259575). Allotype: male, dissected on 5 slides (USNM 259576). Para- types: 14, (leg 4 and abdomen only), dis- sected on 2 slides (USNM 259577): 14, dis- sected on | slide (USNM 259578): 599, habitus, on | slide (USNM 259579); 20¢9, in 70% ethanol (USNM 259580). All from a small stream in the Botanical Garden (alt. ca 1400 m), Godavari, Kathmandu, | 1 May 1983. Female.—Length of holotype excluding caudal setae 0.52 mm: range of lengths of 5 paratypes 0.52-0.67 mm. Cephalothorax (Fig. 1) with elongate planarian-shaped nu- chal organ. All somites except anal somite with posterior margins serrated. All somites except cephalothorax with transverse rows of minute spinules (Fig. 13). Genital double somite with remnant of division beneath integument; genital field as in Fig. 2, reach- ing midlength of double somite. Two uro- somites posterior to genital double somite (Figs. 1, 2) each with one row of long spines on ventral and lateral margin; 3rd uroso- mite also with grouped fine spines in middle of ventral surface. Anal somite (Fig. 2) with- out spines near the posteroventral margin; anal operculum (Fig. 3) with 18 spines, slightly convex. Caudal ramus (Figs. 1, 2, 4) about 1.7 times longer than broad, sub- rectangular, with dorsal, terminally hooked longitudinal keel extending over *4 of the length of ramus. Ramus with basally biar- ticulate dorsal seta inserted near end of keel, two lateral setae, a transverse row of three spines ventrolaterally, group of slender spines distal to medial lobe, and three ter- minal setae. Median terminal seta inserted a little above outer and inner terminal setae, VOLUME 107, NUMBER 2 lacking proximal breaking plane, basally ex- panded, and about 1.3 times longer than urosome. Outer distal seta (Fig. 2) slender, with bulbous base. Inner terminal seta na- ked, slightly longer than half the outer one. Rostrum very small, with two sensilla. Antennule (Fig. 5) of eight articles, article 4 with long esthetasc reaching past end of antennule, article 8 with short and slender esthetasc. Antenna (Fig. 6) biarticulate. Ex- opodite uniarticulate bearing four setae. Palp of mandible (Fig. 7) biarticulate. Proximal article with a single seta. Distal article with four terminal setae and one lateral seta. Legs 1-4 (Figs. 8-11) with triarticulate ex- opodites; endopodite of leg 1 triarticulate, longer than exopodite; endopodites of legs 2-4 biarticulate. Setal formula as follows: Leg 1 basis 1-1 exp 0-1; 1-1; 0,2,2 enp 1-0; 1-0; 0,2,1 Leg 2 basis O-1 exp 0-1; 1-1; 1,2,2 enp 1-0; 2,2,1 Leg 3 basis O-1 exp 0-1; 1-1; 2,2,2 enp 1-0; 3,2,1 Leg 4_ basis O-1 exp 0-1; 1-1; 2,2,2 enp 1-0; 2,1,1 Couplers of legs 1 and 2 with row of spines on each side; those of legs 3 and 4 with smooth surfaces. Leg 5 (Fig. 12), inner expansion of ba- soendopodite reaching '3 length of exopod- ite, with four setae; edge of inner expansion produced into denticles between first and second and second and third setae (com- mencing at medial margin). Exopodite with five setae of which outer lateral two setae are short and next to innermost seta longest. Leg 6 (Fig. 2) consisting of protrusion bear- ing two plumose setae. Male.—Length of allotype 0.47mm, of paratypes 0.42 and 0.43 mm. Body form similar to female. Second to 4th urosomal somites (Fig. 13) with one row of long spines on ventral and lateral margin. Anal somite (Fig. 13) with one or two spines near pos- teroventral margin above each caudal ra- mus. Caudal ramus (Fig. 13) narrower pos- 257 teriorly. Lateralmost terminal seta with slightly bulbous base. Legs 1, 2, and leg 4 exopodites similar to those of female. Leg 3 (Fig. 16) exopodite, major lateral spines of article 1, 2, thick; endopodite triarticulate, modified, spini- form process of article 2 slender to tip, seeming to lack hook, reaching only mid- length of exopodite article 3, article 3 with two long apical plumose setae. Leg 4 en- dopodite (Fig. 17) article 2 shorter than that of female. Leg 5 (Fig. 18) basoendopodite reduced, with two setae, inner one longer than outer one. Exopodite slightly longer than broad and bearing five setae. Leg 6 (Fig. 19) ru- dimentary forming posterior margin of so- mite, only one seta on each side, setae of different lengths. No variation was observed among spec- imens of either sex. Etymology. —Specific name refers to its distribution in Nepal. Affinities. — Elaphoidella nepalensis, n. sp. is most closely related to E. bidens. The morphological characteristics of the female specimens are the same as those of E. bidens s. s. and E. bidens coronata (Sars), except the leg 4 endopodite and the shape of the caudal rami (Chappuis 1931, Gurney 1932, Coker 1934, Lang 1948, Carter & Bradford 1972, Tai & Song 1979, Hamond 1987), and similar to E. bidens decorata (Daday), including the leg 4 and caudal rami (Chap- puis 1931, Tai & Song 1979). Differences between the two species exist in the male. Important diagnostic features of the male of the new species, in contrast to the male of E. bidens s. |. are the arma- ment of the biarticulate endopodite of leg 4 which bears an inner seta on the proximal article and four setae or spines on the distal article (versus uniarticulate in E. b. coronata or biarticulate in E. bidens s. s. and E. b. decorata with two setae on the distal article, except E. bidens s. s. with four setae or spines on the distal article). Also, leg 5 basoen- dopodite bears two setae and the exopodite 258 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Figs. 1-5. Elaphoidella nepalensis, n. sp., female, holotype: 1, Habitus, dorsal; 2, Abdomen, ventral; 3, Anal operculum; 4, Anal somite and caudal ramus, lateral; 5, Antennule. Scales = 100 um. VOLUME 107, NUMBER 2 SOO Le. SPD SZ2RS 1 Wet Treen We mt AAR LLUTTT eae Hany g inn “ny ! 1 w\, TOE QUUEE TEL UU ee ey Ee nt ee Figs. 13-19. Elaphoidella nepalensis, n. sp., male, allotype: 13, Abdomen except Ist urosomite, ventral; 14, Antennule; 15, Leg 2 endopodite; 16, Leg 3 and coupler; 17, Leg 4 endopodite; 18, Leg 5; 19, Leg 6. Scale = 100 um. VOLUME 107, NUMBER 2 bears five setae (versus naked basoendo- podite and exopodite bearing four setae in Elaphoidella bidens s. |.) (Chappuis 1931, Dussart 1967, Carter & Bradford 1972, Tai & Song 1979, Reid & Ishida 1992). Among the already known Elaphoidella species, as far as I know, only Elaphoidella caeca Miura iS equipped with two or three setae on the basoendopodite of leg 5 of the male (Miura 1964). However, this species is quite differ- ent from E. nepalensis in the formula for major armament of legs 2-4 and the rudi- mentary leg 5. The genus Elaphoidella is closely related to Attheyella, and in particular E. bidens is close to Attheyella crassa (Sars) (Lowndes 1950). Leg 5 of the male of E. nepalensis is closer to that of A. crassa than to E. bidens. It can be said that the new species occupies a transitional position between the E/a- phoidella and the Attheyella. The discovery of this species further confuses the generic distinctions within the Canthocamptidae, providing support for Hamond’s (1987) statement that the family is in need of re- vision. Hamond returned several generic and subgeneric taxa including Elaphoidella to the synonymy of the genus Canthocamp- tus Westwood, 1836 s. |. pending eventual revision of the family Canthocamptidae. The revision must be comprehensive, and until then, Iemploy the more familiar genus name. Acknowledgments I wish to pay most cordial thanks to Dr. T. Ito, Hokkaido Fish Hatchery, for her stimulating sampling in Nepal. Sincere thanks are also due to Dr. J. W. Reid for reading the manuscript. Several anonymous reviewers made valuable suggestions for improvement of the manuscript. Literature Cited Brady, G.S. 1880. A monograph ofthe free and semi- parasitic Copepoda of the British Islands, 2:1- 182, pls. 34-82. Ray Society, London. 261 Carter, M. E., & J. M. Bradford. 1972. Postembry- onic development of three species of freshwater harpacticoid Copepoda.— Smithsonian Contri- butions to Zoology 119:1—26. Chappuis, P. A. 1929. Révision du genre Cantho- camptus Westwood (Note préliminaire).—Bu- letinul Societatii de Stiinte din Cluj 4:41—50. . 1931. Copepoda Harpacticoida der Deutsch- en Limnologischen Sunda-Expedition. — Archiv fur Hydrobiologie, Supplement 8:512-584. Coker, R. E. 1934. Contribution to knowledge of North American freshwater harpacticoid cope- pod Crustacea.—Journal of the Elisha Mitchell Scientific Society 50:75-141. Dussart, B. 1967. Les copepodes des eaux continen- tales d’Europe occidentale. Tome I: Calanoides et Harpacticoides. N. Boubee & Cie, Paris, 500 pp. Gumey, R. 1932. British freshwater copepoda. Vol. 2 (Harpacticoida). Ray Society, London, 336 pp. Hamond, R. 1987. Non-marine harpacticoid cope- pods of Australia. I. Canthocamptidae of the genus Canthocamptus Westwood s. lat. and Fi- bulacamptus, gen. nov., and including the de- scription of a related new species of Cantho- camptus from New Caledonia.—Invertebrate Taxonomy 1:1023-1247. Lang, K. 1948. Monographie der Harpacticiden. Vols. I, II. Nordiska Bokhandeln, Stockholm, 1648 Lowndes, A. G. 1950. The males of Canthocamptus bidens Schmeil.— Proceedings of Zoological So- ciety of London 120:395-403. Miura, Y. 1964. Subterranean harpacticoid copepods from a driven well in Japan.—Japanese Journal of Zoology 14:133-141. Reid, J. W., & T. Ishida. 1993. New species and new records of the genus Elaphoidella (Crustacea: Copepoda: Harpacticoida) from the United States. — Proceedings of the Biological Society of Washington 106:137-146. Sars, G.O. 1903-1911. An account of the Crustacea of Norway. V. Copepoda Harpacticoida. Bergen Museum, Bergen, 449 pp., 284 pls. Tai, A., & Y. Song. 1979. Harpacticoida Sars, 1903. Pp. 164—300 in Fauna Editorial Committee, Ac- ademia Sinica, ed., Fauna Sinica Crustacea Freshwater Copepods. Science Press, Peking, 450 pp. 372 Irifunecho, Yoichimachi, Hokkaido, 046 Japan. PROC. BIOL. SOC. WASH. 107(2), 1994, pp. 262-267 MONSTRILLA ELONGATA, A NEW MONSTRILLOID COPEPOD (CRUSTACEA: COPEPODA: MONSTRILLOIDA) FROM A REEF LAGOON OF THE CARIBBEAN COAST OF MEXICO E. Suarez-Morales Abstract. — A new species of a monstrilloid, Monstrilla elongata, is described from plankton samples collected in a reef lagoon along the northern portion of the eastern coast of the Yucatan Peninsula. The new species is described from a single female and can be distinguished from the other species of Mon- strilla by the combination of the body proportions, a single-lobed fifth leg bearing only two setae, furcal rami with five setae, and the unusual annulated structure of the ovigerous spines. Monstrilloid copepods are occasional el- ements in plankton samples, since only the reproductive adult stage is free-living (Da- vis 1984). Naupliar and juvenile stages of these copepods are parasites of polychaetes and gastropod molluscs (Hartmann 1961, Huys & Boxshall 1991). This group is one of the least known within the Copepoda. The number of known species is relatively small (around 90 nominal species) and sev- eral have been described from a single spec- imen (Davis 1947, 1949: Suarez-Morales & Gasca-Serrano 1992). Valuable, but not comprehensive revisions of the group have been made by Davis (1949) and by Isaac (1975). Only three genera are now recog- nized, Monstrilla, Monstrillopsis and Thau- maleus (Huys & Boxshall 1991). Some previous records of Monstrilla in the western tropical Atlantic include: M. floridana Davis, 1947, M. rugosa Davis, 1947, M. reticulata Davis, 1949, M. hel- golandica Giesbrecht, 1892 and M. grandis Giesbrecht, 1891 (Davis 1947, 1949; Isaac 1975, Fish 1962, Reid 1990). Additional records of monstrillids have been stated from material obtained on the northern and central portions of the eastern coast of the Yucatan Peninsula. In these areas, several new species of Monstrilla (M. barbata Sua- rez-Moralez & Gasca-Serrano, 1992: M. reidae Suarez-Morales, 1993a; M. rebis Sua- rez-Morales, 1993b; M. mariaeugeniae Suarez-Morales & Islas-Landeros, 1993) as well as the new species Monstrillopsis cigroi Suarez-Morales, 1993b and Thaumaleus boxshalli Suarez-Morales, 1993c have been reported. During plankton surveys carried out by CIQRO in a reef lagoon located off Puerto Morelos along the northern portion of the Yucatan Peninsula’s eastern coast (Suarez & Gasca 1990), one undescribed species of monstrilloid copepod belonging to the ge- nus Monstrilla was collected. This species was previously misidentified by Suarez & Gasca (1990) as Monstrilla leucopis Sars, 1921. Monstrilla elongata, new species Type locality.—Reef lagoon off Puerto Morelos, northern portion of the eastern coast of the Yucatan Peninsula (20°51.40'N; 86°54.15'W). Date of collection 1988 Jan 16. Water column. Over Thalassia testu- dinum beds. Material examined. —Holotype; female, undissected, deposited in the U. S. National Museum of Natural History, Smithsonian Institution, under number USNM-259488. Paratype; female, undissected, deposited in VOLUME 107, NUMBER 2 263 Fig. 1. A. Monstrilla elongata n. sp. habitus, dorsal. B. habitus, lateral. C. nght antennule, dorsal view. D. left antennule, dorsal view. E. head and first antennular segments. F. urosome, ventral view. G. fifth legs, ventral view. 264 the same institution under number USNM- 259665. Specimens preserved in 70% eth- anol. Paratype; female, dissected, author’s collection at CIQRO. Female.—Length 4.2 mm. Cephalic seg- ment long and slender, almost 0.65 of total body length. Oral papilla located 0.43 of way back along cephalic segment. Eyes ab- sent (Figs. 1A, B). Antennulae 5-segmented, with segments 2-5 partially fused. Antennulae armed with three spines on first segment, and ten spines and six setae on remaining four segments. Three of these setae ripped away, sockets remaining. Large aesthetasc at midlength. Ratio of length of first segment and re- mainder being: 14:86 = 100 (Figs. 1C, D). Antennulae 0.24 of total body length. First segment with small lateral protuberance on basis, visible in dorsal view (Fig. 1E). Incorporated first thoracic somite and succeeding three thoracic segments bearing well developed, biramous swimming legs with triarticulated rami (Figs. 2J, K, L). Swimming legs equal in length and armed as follows: basis endopodite exopodite leg 1 I-0 QO-1; 0-1; O-1, 1,3 I-0; 0-1; 1, 1, 3 leg 2 O-O £O-1; 0-1; 0-1,1,3 I-0; 0-1; I, 1,4 leg 3 O-O0 £O-1; 0-1; 0-1, 1,3 I-0; 0-0; I, 1, 3 leg 4 0-0 O-1; 0-1; 0-1,1,3 I-0; 0-1; I, 1,3 Fifth leg 1-segmented with broad single lobe bearing two setae (Fig. 1G); outer seta slightly longer than inner but both reaching beyond distal end of furcal rami (Fig. 1F). Urosome consisting of fifth pedigerous somite, genital double and two free abdom- inal somites (Fig. 2H), length ratio of these 4 segments being: 39.6:33.3:14.6:12.5 = 100. Genital complex with two thick, annulated Ovigerous structures, as shown in Fig. II. Distal ends of ovigerous structures reaching slightly beyond distal end of furcal rami. Furcal rami 2.3 times longer than wide, bearing five setae, four of them strongly de- veloped, remaining one being thinner and ¥% as long as others. One of large setae borne PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON on proximal outer margin, small seta on distal outer margin, remaining three setae terminal. Male. —unknown. Etymology.—The specific name makes reference to the unusual proportional length of the cephalic segment. Discussion The new species has been assigned to the genus Monstrilla on the basis of the presence of two free abdominal somites posterior to the genital double somite, the absence of eyes, and the location of the oral papilla more than 0.25 of the way back along the cephalic segment (Isaac 1975). Monstrilla elongata differs from all other species of the genus Monstrilla in some rel- evant features. The presence of a single lobe with two setae on the fifth leg is a feature shared with M. conjunctiva Giesbrecht, 1902, M. helgolandica Claus, 1863, M. lon- gipes A. Scott, 1909 and M. ghardagensis Al-Kholy, 1963. The structure of the lobe is different in each case; in M. helgolandica, it is narrow and bent in the middle (Park 1967, Isaac 1975), but in M. conjunctiva, the same structure is broad at base and nar- rows abruptly (Isaac 1975). Monstrilla lon- gipes exhibits a very long and slender fifth leg lobe (Davis 1949, Scott 1909), and in M. ghardagensis is short and slender (AI- Kholy 1963). In M. elongata, this lobe is broad both at the base and at distal portion, with a slight medial constriction, as shown in Fig. 1G. The relative length of the antennulae dif- fers in the five species; in M. conjunctiva, the antennulae constitute 0.35 of the total body length, this proportion is 0.28 in M. helgolandica, 0.19 in M. ghardagensis, 0.22 in M. longipes and 0.24 in M. elongata. Moreover, neither of these species have fused antennular segments, a condition clearly present in M. elongata. This feature, however, is not uncommon throughout the genus; in M. longiremis Giesbrecht, 1892, VOLUME 107, NUMBER 2 th l aM) ri 265 Fig. 2. HH. urosome, lateral view. I. genital segment with ovigerous structures, and furcal rami. J. second leg. K. third leg. L. first leg. only the proximal segment is clearly de- fined, and in M. grandis Giesbrecht, 1891, only the first two segments are separated. The presence of a spine on the medial side of the basis of the first legs is another dis- tinctive feature of M. elongata when com- paring it with M. helgolandica, M. longipes and M. ghardagensis. It is only present in a few other monstrilloids (Grygier, pers. comm.). Furthermore, M. elongata differs 266 from M. helgolandica, M. longipes and M. ghardagensis in the number of furcal setae, six in M. helgolandica and M. longipes, and four in M. ghardagensis, but only five in the new species. Monstrilla conjunctiva also has five furcal seta, with the same arrangement found in M. elongata (Sewell 1949). In both M. helgolandica and M. conjunc- tiva, the genital double somite is at least 1.5 times longer than the free abdomen (Sewell 1949, Isaac 1975). It is shorter in M. /on- gipes. In M. elongata and in M. ghardagen- sis, the genital somite is almost the same length as the free abdomen. The structure of the genital complex is also different in these species. In M. helgolandica, M. ghar- dagensis and in M. conjunctiva, the oviger- Ous spines are long and slender, reaching beyond the distal end of the furcal rami (Sewell 1949). The structure of the genital complex or of the ovigerous spines are not described in the original description of M. longipes (Scott 1909). In M. elongata, the Ovigerous structure is broader and shorter, and is not slender, but exhibits a thick, an- nulated aspect. The same type of structure has also been found for M. mariaeugeniae from the same locality (Suarez-Morales & Islas-Landeros 1993). These kind of appar- ently undeveloped genital structures can not be related to copepodids or other immature stages since the development of monstril- loids takes place up to the fully mature adult within the host, and the adult burrows out of the host as a planktic form (Davis 1984). Finally, measuring 4.2 mm, the new spe- cies is clearly larger than M. conjunctiva (3.3— 3.8 mm), M. longipes (1.83 mm), M. ghar- dagensis (1.35 mm) and M. helgolandica (1.4-2.3 mm). It is also is one of the largest species of the genus, after M. mariaeugeniae (4.4 mm) and M. clavata (4.5 mm). Acknowledgments Thanks to Dr. Thomas E. Bowman for his comments on the drawings. This work was carried out with the logistic and finan- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON cial support of the Centro de Investiga- ciones de Quintana Roo (CIQRO) (Proj. 01- 02-009) and of CONACYT (Proj. 1118- N9023). Additional material of this species was sorted by Ma. Eugenia Islas-Landeros. Ms. Janice Clark, of the National Museum of Natural History, Smithsonian Institu- tion, kindly made the deposition and ca- talogation of the type specimens. Mark J. Grygier granted relevant literature. Literature Cited Al-Kholy, A. A. 1963. Some semi-parasitic Cope- poda from the Red Sea.—Publications of the Marine Biological Station, al-Ghardagqa (Red Sea) 12:127-136. Davis, C.C. 1947. Two monstrilloids from Biscayne Bay, Florida.—Transactions of the American Microscopical Society 66:390-395. 1949. A preliminary revision of the Mon- strilloida, with descriptions of two new spe- cies.— Transactions of the American Micro- scopical Society 68:245-255. 1984. Planktonic Copepoda (including Mon- strilloida). Pp. 67-91 in K. A. Steidinger & L. M. Walker, eds., Marine plankton life cycles strategies. CRC Press, Florida. Fish, A.G. 1962. Pelagic copepods from Barbados. — Bulletin of Marine Science of the Gulf and Ca- ribbean 12:1-38. Giesbrecht, W. 1892. Systematic und faunistik der pelagischen Copepoden des Golfes von Neapel und der angrenzenden Meeres-Abschnitte. — Fauna und Flora Golfes Neapel 19:1-831. Hartmann, O. 1961. A new monstrillid copepod par- asitic in capitellid polychaetes in southern Cal- ifornia.— Zoologische Anzeigen 167:325-334. Huys, R., & G. Boxshall. 1991. Copepod evolution. The Ray Society, London, 468 pp. Isaac, M. J. 1975. Copepoda. Sub-Order: Monstril- loida. Conseil International pour L’exploration de la Mer, Fiche Identification Zooplankton 144/ 145, pp. 1-10. Sewell, R. B.S. 1949. The littoral and semi-parasitic Cyclopoida, the Monstrilloida and Notodel- phyoida.—Scientific Reports of the John Mur- ray Expedition 1933-34. 9:17-199. Reid, J. W. 1990. Continental and coastal free-living Copepoda (Crustacea) of Mexico, Central America and the Caribbean region. Pp. 175-214 in D. Navarro L. & J. G. Robinson, eds., Div- ersidad Biologica en la Reserva de la Biosfera de Sian Ka’an Quintana Roo, Mexico. CIQRO/ Univ. of Florida. Chetumal, México. VOLUME 107, NUMBER 2 Scott, A. 1909. The Copepoda of the Siboga Expe- dition. Part I. Free-swimming, littoral and semi- parasitic Copepoda.—Siboga Expeditie 29:323 pp. + 69 pls. Suarez, E., & R. Gasca. 1990. Variacion dial del zoo- plancton asociado a praderas de Thalassia tes- tudinum en una laguna arrecifal del Caribe Mex- icano.— Universidad y Ciencia 7:57—64. Suarez-Morales, E. 1993a. Monstrilla reidae, a new species of monstrilloid copepod from the Ca- ribbean Sea off Mexico.—Bulletin of Marine Science 52:717-720. 1993b. Two new monstrilloids (Copepoda: Monstrilloida) from the eastern coast of the Yu- catan Peninsula.—Journal of Crustacean Biol- ogy 13:349-356. 1993c. A new species of Thaumaleus (Co- 267 pepoda: Monstrilloida) from the Eastern coast of the Yucatan Peninsula. —Crustaceana 64:85-— 89. —., & R. Gasca-Serrano. 1992. A new species of Monstrilla (Copepoda: Monstrilloida) from mexican coasts of the Caribbean Sea.—Crus- taceana 63:301-305. , & M. E. Islas-Landeros. 1993. A new species of Monstrilla (Copepoda: Monstrilloida) from a reef lagoon off the Mexican coast of the Carib- bean Sea.— Hydrobiologia (in press). Centro de Investigaciones de Quintana Roo, A.P. 424, Chetumal, Quintana Roo 77000, Mexico. PROC. BIOL. SOC. WASH. 107(2), 1994, pp. 268-273 ANNINA MANNAT, A NEW ISOPOD FROM THE GANGES RIVER, WEST BENGAL (CRUSTACEA: ISOPODA: CIROLANIDAE) Marilyn Schotte Abstract.—Annina mannai, n. sp., the fifth known species of Annina, is described from fresh water of the Ganges River in West Bengal, India. It is distinguished from its congeners by a single, crescent-shaped dorsal projection in the male on pereonite 2. The genus Annina was erected in 1908 by Budde-Lund to accommodate A. J/acustris, taken from a salt-water pool in Zanzibar off the East Africa coast. Three others have since been described from mangrove swamps, es- tuaries and streams (A. kumari [Bowman 1971] from Malaysia; A. fustis Bowman & Iliffe 1991 from Thailand; and A. mesopo- tamica [Ahmed 1971] from Iraq). Jones (1983) provided a revised diagnosis of An- nina, declaring it distinct from the closely- related Excirolana due to the presence of dorsal projections on the male cephalon and/ or pereonites 1 and/or 2, a transverse non- facetted band in the eye, and other char- acters. Herein is described a fifth species, col- lected by Dr. A. K. Manna in fresh water of the Ganges River along with atyid shrimp. All known species of Annina have been de- scribed from near the northern perimeter of the Indian Ocean region (see Fig. 4). Non- type localities include Singapore (A. /fustis) and Kenya and Comoros Islands, where A. lacustris has been found (Bowman & Iliffe 1991). Genus Annina Budde-Lund, 1908 Annina mannai, new species Figs. 1-3 Material.—USNM 252750 Holotype 4, TL 7.0 mm; USNM 252751 Allotype 2°, TL 8.9 mm; USNM 252752 Paratypes, 1é, 18 2, 11 juvs., shore of Ganges River in District Murshidabad, West Bengal, India, coll. A. K. Manna, 1992. Description. — Length up to 8.9 mm. Body widest at pereonite 6; pleon narrower than pereon. Marked sexual dimorphism; adult male with dense patch of setae between eyes and crescent-shaped middorsal process on pereonite 2; blunt, raised “shoulders” at an- terolateral parts of pereonite 2; pereonite 1 with middorsal depression. Female cepha- lon without setae or processes; no processes on pereon. Cephalon produced anteriorly into rounded rostrum projecting between anten- nal bases. Lateral incision posterior to each eye, reaching medially 4 width of cephalon. Clypeus triangular, pointed, directed an- tero-ventrally. Eyes large, with transverse, unfacetted gap dividing dorsal and ventral halves. Coxae of pereonites 1—3 rounded posteriorly; coxae 4—7 progressively more sharply pointed. Lateral margins of pleo- nites 1—2 straight; those of 3—5 increasingly more extended. Telson triangular, posterior 73 finely scalloped, bearing plumose setae, without spines. Uropodal endopod similar- ly scalloped on both margins, bearing plu- mose setae, endopod barely extending be- yond apex of telson. Uropodal exopod lanceolate, faintly notched on medial mar- gin, acuminate at apex, reaching distal *4 length of endopod. Pigment densest near posterior margins of cephalon, pereonites, VOLUME 107, NUMBER 2 269 Fig. 1. Annina mannai, new species. A, 6 habitus. B, 4, lateral view of anterior. C, lateral view of °. D, antennae | and 2. E, frontal lamina and clypeus. F, maxilla 1. G, maxilla 2. H, maxilliped. I, right mandible. J, spine row and lacinia mobilis, detail. K, telson and uropods. 270 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON S\ Fig. 2. Annina mannai, new species. A, pereopod 7. B, pereopod 1. C, cleft spine on pereopod |. D, pereopod 2. E, penes. F, pleopod 2, 6. G, pleopod 1. H, pleopod 3. I, pleopod 4. J, pleopod 5. VOLUME 107, NUMBER 2 Fig. 3. Annina mannai, new species. A, lateral view of anterior, 6. B, cephalon and pereonites 1, 2 of 6. C, cephalon and pereonites 1, 2 of 6. D, eye. E, interocular area of cephalon, closeup. F, detail of setae on scales of cephalon. G, cephalon, ventral view. H, cephalon, dorsal view, 2. I, ventrolateral view of cephalon. pleonites and anterior part of telson; pig- ment absent from medial area of pleon, forming inverted triangular shape. Antenna | reaching pereonite 4; flagellum with 15-16 articles. Antenna 2 flagellum with 9 articles in female, unknown in male; peduncle more robust and longer in male. Mandible tricuspidate; spine row composed of 9 spines, several fine setae also present. Maxilla 1, exopod with 1 seta and 12 spines PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON SO) a A. lacustris IC WA el ; Mm TOF za Sailors | Mseuseeee ie) Fig.4. Distribution of known Annina species. @ Type localities. HM Additional record for A. fustis. ® Additional records for A. /acustris. (most with accessory spinules) as figured; endopod typical of genus. Maxilla 2 bearing 8 setae on both palp and exopod; endopod with about 11 setae, simple and plumose. Maxilliped typical of genus. Pereopods ambulatory. Posterior margin of pereopod 1 with dentate, cleft spines. Pe- reopods 2 and 7 with many spines and setae, as figured. Pleopod 1 with 4 coupling hooks and 3 plumose setae. Pleopod 2 with 3 coupling hooks and 4 plumose setae, appendix mas- culina not reaching apex of endopod. Pleo- pod 3 and 4 each with 3 coupling hooks and 3 setae; pleopod 5 without hooks or setae. Exopods and endopods of pleopods 1—2 with plumose marginal setae, endopods of ple- opods 3-5 lacking setae. Rami of all pleo- pods undivided. Penes short, apically rounded. Etymology. —The species is named for its collector, Dr. A. K. Manna, Professor of Zoology at Sripat Singh College, West Ben- gal, India. Remarks. —Dr. Manna (pers. comm.) re- ported that the species, present year-round, is abundant in its Ganges River habitat dur- ing the rainy season and was found to be feeding on dead shrimp (Caridina sp., Fam- ily Atyidae), suggesting that it is a scavenger. An isopod, Tachaea sp., was also found in association with these shrimp. Annina mannai is the only known mem- ber of the genus with a single, salient mid- dorsal process on pereonite 2 in the male. In all others the processes are paired and sublaterally placed. Below is presented a key to the five species of Annina, modified from that of Jones (1983) and based on characters of the adult male. VOLUME 107, NUMBER 2 Key to Annina species 1. Posterior margin of telson truncate peers ey ad ere eR A. lacustris Budde-Lund cate 2. Male with 1 large middorsal projec- tion of pereonite 2 .. A. mannai n. sp. — Male with 2 dorsolateral projections or horns on pereonite 2 ......... 3 3. Projections club-shaped in lateral view, longer than half-length of pe- reonite 1 .... A. fustis Bowman & Iliffe — Projections short (less than one-half length of pereonite 1), either blunt or acute 4. Paired and pointed horns on cepha- lon and pereonites | and 2 Er duictemnins ee ise A. kumari (Bowman) — Paired horns, short and blunt, on pereonite 2 only ot Sah ah eta A. mesopotamica (Ahmed) Acknowledgments I thank Dr. Brian Kensley, Department of Zoology, NMNH, for reviewing the 273 Manuscript, and anonymous reviewers for further suggestions. Dr. A. K. Manna kindly donated his collected material to the NUNH collection. Literature Cited Ahmed, M. M. 1971. New Isopoda (Flabellifera) from Iraq and Arabian Gulf.— Mitteilungen aus dem Zoologischen Museum in Berlin 43(1):77-83. Bowman, T. 1971. Excirolana kumari, a new tubi- colous isopod from Malaysia. —Crustaceana 20: 70-76. Bowman, T., & T. Iliffe. 1991. Annina fustis, a new isopod from Phang Nga, Thailand (Crustacea: Isopoda: Cirolanidae).— Proceedings of the Bi- ological Society of Washington 104:247-252. Budde-Lund, G. 1908. Isopoda von Madagaskar und Ostafrika mit Diagnosen verwandter Arten, in A. Voeltskow, Reise in Ostafrika in dem Jahren 1903-1905.—Wissenschaftliche Ergebnisse 2:265-308, pl. 12-18. Jones, D. A. 1983. On the status of the cirolanid isopod genera Annina Budde-Lund, 1908 and Excirolana Richardson, 1912.—Crustaceana 45(3):309-3 12. Department of Invertebrate Zoology, NHB Stop 163, National Museum of Nat- ural History, Smithsonian Institution, Washington, D.C. 20560, U.S.A. PROC. BIOL. SOC. WASH. 107(2), 1994, pp. 274-282 REDESCRIPTION OF JAIS ELONGATA SIVERTSEN & HOLTHUIS, 1980, FROM THE SOUTH ATLANTIC OCEAN (CRUSTACEA: ISOPODA: ASELLOTA) Brian Kensley Abstract. —lais elongata is redescribed from a freshwater stream on Inac- cessible Island, Tristan da Cunha archipelago. The presence of several males, each in amplexus with a stage 1 manca, demonstrates that precopula or mate guarding, which also provides some measure of offspring protection in a swift- flowing stream, occurs in this species. About 130 specimens of a tiny asellote isopod were collected from a freshwater stream on Inaccessible Island (37°02’S, 12°12’W) in the Tristan da Cunha archi- pelago of the South Atlantic, by Mrs. Helen James of the Albany Museum, Grahams- town, South Africa. The collection was made in the course of a survey of the invertebrate fauna of the island, and submitted to the Smithsonian Institution for identification. The bulk of the collection is housed in the Albany Museum; a representative sample has been deposited in the National Museum of Natural History, Smithsonian Institu- tion. Although the isopods were determined to be a recently described species, it was de- cided to provide a fuller redescription with illustrations, especially as some interesting details of the biology were revealed. Family Janiridae Tais elongata Sivertsen & Holthuis, 1980 Figs. 1—5 Tais elongata Sivertsen & Holthuis, 1980: 104, Fig. 34. [Described from 2 6 and 1 9]. Material examined.—Trondheim Uni- versity Museum, Holotype, 6 TL 1.2 mm, paratypes | 6, 1 2, (3 slide preparations), sta 136, pool of brackish water in cave 2 m above beach, North Point, Inaccessible Is- land.—Stream flowing to Blenden Hall beach, Imnaccessible Island, 37°02’S, 12°12'W: Sample TDC 2A, 2 ovig. 2, 21 2, 16 6 (6 6 with manca), 13 Oct 1989.—Sam- ple TDC 11], 4 ovig. 2, 20 2, 10 6 (1 6 with manca), 23 Oct 1989.—Sample TDC II, 5 Ovig. 2, 26 2, 30 6 (3 with manca), 23/24 Oct 1989 (3 ovig. 2, 10 2, 10 6 in NMNH collection). Description. — Body (Fig. 1A) about 3.5— 3.8 times as long as wide. Integument bear- ing short scattered setae, some stouter than others. Cephalon about one-third wider than midlength, slightly narrower than pereonite 1; anterior margin with broadly rounded rostrum reaching to about midlength of first antennular article. Eyes well pigmented, of 2 ommatidia each. Coxae visible in dorsal view on all pereonites. Pereonite | slightly shorter than pereonite 2; pereonites 2—4 subequal in length and width, each with se- tose anterolateral rounded lobe; pereonite 5 shortest and narrowest; pereonites 6 and 7 with setose posterolateral tergal lobes. Pleon consisting of single short anterior segment plus pleotelson; latter subequal in length to pereonites 6 and 7, lateral margins weakly convex with about 15 short setae, posterior margin weakly convex between uropodal insertions. Antennule (Fig. 1C) of 6 articles, almost VOLUME 107, NUMBER 2 275 CO a aa | Dl Fe Fig. 1. Jais elongata: A, Adult in dorsal view, scale = 0.2 mm; B, Antenna; C, Antennule; D, Upper lip; E, Lower lip; F, Distal part of left mandible; G, Mandibular palp; H, Right mandible; I, Ventral view of last pereonite and pleon of male; J, maxilla 1; K, Maxilla 2: L, Maxilliped. 276 reaching distal margin of antennal article 5, article 1 widest and longest, twice length of article 2; terminal article bearing single aesthetasc. Antenna (Fig. 1B) equal in length to cephalon plus pereonites 1—5 and half of pereonite 6; peduncle articles 1—4 shorter than wide; article 3 with minute distolateral scale bearing 2 short setae; articles 5 and 6 longer than wide, article 5 about three- fourths length of article 6; flagellum of about 19 setose articles in male, 16 in female. Up- per lip (Fig. 1D) about 1.8 times broader basally than midlength, distally broadly rounded, setose. Mandibular palp (Fig. 1G) of 3 articles, article 1 about two-thirds length of article 2, with single strong distolateral seta; article 2 bearing 2 strong bilaterally pectinate setae laterally in distal half; article 3 with mesial margin convex, lateral margin straight with 5 spines increasing in length distally; mandibular molar cylindrical, grinding surface sclerotized, truncate; inci- sor of 4 sclerotized cusps; spine row of 3 short and 2 elongate spines (left, Fig. 1F), 4-toothed lacinia mobilis, 2 short and 2 elongate spines (right, Fig. 1H). Lower lip (Fig. 1E) deeply cleft, lobes laterally broadly convex, mesiodistally strongly setose. Max- illa 1 (Fig. 1J) with mesial lobe bearing 1 distomesial and 6 distal simple setae; lateral lobe bearing 11 stout toothed spines distal- ly. Maxilla 2 (Fig. 1K), mesial lobe bearing about 13 mostly simple setae mesiodistally; 2 lateral lobes each with 4 elongate unilat- erally pectinate setae. Maxillipedal palp (Fig. 1L) of 5 articles, article | broader than long, article 2 twice length and slightly wider than article 1, ar- ticle 3 three-fourths length and two-thirds width of article 2, articles 4 and 5 slender, article 4 2.7 times length of article 5; endite reaching to midlength of palp article 4, with 2 coupling hooks on mesial margin, distal margin with submarginal row of 6 stout pec- tinate spines, marginal row of 7 slender se- tae; epipod ovate, distally broadly rounded, reaching palp article 3. Pereopods (Figs. 2, 3) all ambulatory, pereopod | only slightly PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON shorter than pereopod 2, with biunguiculate dactylus; pereopods 2—3 and 5-7 in male and pereopods 2-7 in female essentially similar, becoming more elongate posteri- orly, with basis moderately broad, bearing strong seta on anteroproximal margin, is- chium with 2 or 3 setae on anterior margin, merus with group of 3 anterodistal setae, carpus subequal in length to merus and is- chium together, with short posterodistal spine, propodus about half width and sub- equal length of carpus, with short postero- distal spine, dactylus with 3 hooked claws. Pereopod 4 in male about two-thirds length of pereopod 3 or 5, propodus with stout posterodistal spine, dactylus bearing ter- minal hooked claw, and reflexed subter- minal claw. Female pleopod 2 forming broad oper- culum (Fig. 4A), midlength subequal to greatest width, distally broadly rounded with 2 submesial setae. Male with short conical penes (Fig. 4B) on posterior sternal margin of pereonite 7, just reaching base of pleopod 1. Male pleopods | and 2 together forming operculum; pleopod | elongate (Fig. 4B), rami with fused portion 3.5 times longer than free, distal rami rounded and bearing about 7 simple setae, distolateral projection short, narrowly rounded. Pleopod 2 (Fig. 4C) peduncle roughly triangular, lateral Margin convex; exopod with 2 articles sub- equal, article 2 distally rounded; endopod with article | about half length of article 2, latter slender, curved, tapering to narrow apex reaching distal end of peduncle, with narrow open furrow. Pleopod 3 (Fig. 4D) with exopod of 2 articles subequal in length, article 1 wider than 2, lateral margin con- vex, setose; article 2 tapering distally lateral and mesial margins setose, with single sim- ple setae apically, endopod roughly ovate, with irregular margin, reaching distal half of exopod article 2. Pleopod 4 (Fig. 4E) ex- opod of single slender article bearing single elongate distal seta; endopod broad, roughly ovate, with irregular margin. Pleopod 5 (Fig. 4F) of single irregular roughly ovate ramus. VOLUME 107, NUMBER 2 277 0.1 Fig. 2. Jais elongata, female, pereopods 1-7. Uropod (Fig. 4G) about half length of pleo- telson, peduncle with single strong dis- tomesial seta; endopod 2.8 times length of exopod, with several terminal simple setae; exopod with 3 terminal setae. Habitat.—The specimens were collected from a narrow, fairly swiftly flowing stream (about 60 cm wide, 50 cm deep), arising from a spring and running less than one kilometer before dropping to a boulder beach. The stream water was clear, pH neu- tral; the grass Spartina arundinacea was growing closely along the banks. The spec- imens were found under stones in the stream, usually several animals per stone. Gut contents. — Determined from 2 cleared specimens: several kinds of diatoms, spore- like structures, and fine filamentous algal- like structures in addition to unidentifiable fragments. The female paratype, cleared and mounted on a microscope slide, has the en- tire gut packed with diatoms. Size.— Males total length (tl) 1.08-1.30 mm, possibly in 2 size-groups; males with 278 1 3 ee Fig. 3. Jais elongata, male pereopods 1-7. manca tl 1.13—1.30 mm, manca tl 0.43 mm; females tl 0.75—1.45 mm, possibly in 2 size- groups; ovigerous females tl 1.20—1.48 mm. Ovigerous females (average for 11 spec- imens tl 1.32 mm) somewhat larger than mature males (average for 9 specimens tl 1.22 mm). Brood size. —Contents of brood pouch in ovigerous females: | egg—1, 2 eggs—2, 3 eggs—2, 2 manca—4, 3 manca—1. Mate guarding. —Ten of the 56 males col- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON lected were each found to be carrying a sin- gle large manca, tucked between the pereo- pod bases (Fig. 5). All the mancas (manca I stage, 0.43 mm total length) were female; all were oriented in the same way, i.e., dor- sum of manca pressed to ventrum of male, head directed posteriorly with respect to the male. This is clearly a case of mate guarding, with the juvenile partner held in precopula. The fourth pereopods of the male, which hold the manca in position, are two-thirds VOLUME 107, NUMBER 2 279 0.1 0.05 Fig. 4. Jais elongata: A, Female pleopod 2 operculum; B, Male pleopod 1 and penes; C, Male pleopod 2; D, Pleopod 3; E, Pleopod 4; F, Pleopod 5; G, Uropod. the length of the third or fifth pereopods, and have one of the dactylar claws reflexed. This precopulatory mate guarding strat- egy (see Dunham & Hurshman 1991, Franke 1993) has been observed in a number of crustaceans (Ridley 1983) including iso- pods. Precopula ensures that the male is present when a female is receptive during the brief biphasic molt, either by being pas- sively attached to the female, or by actively carrying her around. Copulation in isopods takes place during the biphasic molt to the adult brooding phase. The posterior half molts first, fol- lowed by insemination, then the anterior half molts, along with deployment of oos- tegites. Eggs are released into the brood- pouch once molting is complete (Veuille 1978). Most isopods have internal insemi- nation (Ridley 1983). Several asellote gen- era have been noted to resort to mate guard- ing; species of Jaera and Munna carry 280 Fig. 5. of adult male holding female manca. Tais elongata, composite SEM photograph around potential mates (Veuille 1980, Hes- sler & Stromberg 1989, Franke 1993). As in the present species, in Jaera the fourth pereopod of the male is shortened and bears specialized spines for clasping the female PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON during precopulation and copulation (Veuille 1980). The same leg is used in am- plexus in the primitive asellotes Caecidotea (Lewis & Bowman 1981) and Asel/lus (Gru- ner 1965). At a maximum total length of 1.48 mm, this species is within meiofaunal size-range. Many meiofaunal organisms exhibit regres- sive evolution of body size (Swedmark 1964) accompanied by reduction in number of eggs and offspring per brood. In this case, four eggs was the maximum number observed within a brood-pouch while two manca per brood pouch was the commonest number (n = 4) of manca observed. Given this very low number of offspring per brood, contin- ued care of offspring after release from the brood-pouch would greatly improve the survival rate, especially given the habitat of a fairly swiftly flowing stream. This post- marsupial care is afforded by the male’s use of precopulatory mate guarding of the fe- male manca. The male is already adapted for amplexus, and this adaptation becomes a form of offspring care. This double use of an adaptation is more parsimonious than if the female, which lacks any clasping mod- ifications, were to provide some form of post-marsupial care. The use of the fourth pereopod of the male for hanging onto the female is probably a primitive character within the Asellota, being found in the asel- lids, while the loss of one of the three dac- tylar ungui, and the flexing of one of the remaining ungui along with marked short- ening of the leg seen in the janirids, repre- sent a more advanced state than that seen in the asellids. Remarks. —The genus Jais Bovallius, 1886, at present contains five species (see Nierstrasz 1941:53; Menzies & Barnard 1951:138; Coineau 1977:436). Menzies & Barnard, 1951, expressed the opinion that true species of Jais are always found in as- sociation with sphaeromatid isopods, and that the free-living species may well belong to a different genus. There are at least three records of free-living Jais: I. pubescens of Barnard, 1965 from Gough Island, J. aquilei VOLUME 107, NUMBER 2 Coineau, 1977 from St. Helena Island, and I. elongata Sivertsen & Holthuis, 1980 from Inaccessible Island in the Tristan archipel- ago. All three of these South Atlantic island records are from freshwater habitats at varying (but never great) distances from the sea. Possibly we are dealing with a complex of island species of a genus other than Jais. Material from Tristan da Cunha, taken from the fully marine sphaeromatid Jsocladus tristanensis and identified as I. pubescens by Barnard (South African Museum, A2286, A2452) appears to be closely similar to J. pubescens, widespread through the southern ocean. Examination of commensal material from Auckland, New Zealand, taken from Exosphaeroma gigas, from southern Peru taken from Sphaeroma peruvianum, along with the abovementioned Tristan material reveals considerable variation in body and appendage proportions, suggesting a com- plex of species rather than a single circum- polar species, masquerading under the name I. pubescens. Tais elongata is very similar to I. aquilei from St. Helena Island, even in the total length of adult males and females. Coi- neau’s thorough description and figures (1977), however, do reveal some subtle dif- ferences: the basal article of the antennule is relatively broader in J. aquilei; the third article of the mandibular palp has 3 distal spines (5 in J. elongata); the maxillipedal endite has 8 distal broad spines (6 in I. elon- gata), maxillipedal palp article 4 is rela- tively longer in J. elongata; there are fewer distal setae on pleopod 1 6 in J. elongata; the endopod of pleopod 3 is relatively smaller compared with the exopod in J. elongata; the exopod of pleopod 4 and es- pecially the single distal seta are relatively larger compared with the endopod in J. elongata; the uropodal exopod is relatively more elongate compared to the endopod in I. aquilei. Acknowledgments I am grateful to Mrs. Helen James of the Albany Museum, Grahamstown, South Af- 281 rica, for making the material of 7. elongata available for study. The expedition to In- accessible Island that yielded this material was organized by the Percy Fitzpatrick In- stitute for African Ornithology at the Uni- versity of Cape Town. I thank Mrs. Michelle van der Merwe of the South African Mu- seum, Cape Town, and Dr. Tor Stromgren of the Trondheim University Museum, Norway, for the loan of comparative ma- terial. Dr. George Wilson of the Australian Museum generously shared his thoughts on asellote evolution, in correspondence over this paper. Literature Cited Barnard, K. H. 1965. Isopoda and Amphipod col- lected by the Gough Island Scientific Survey. — Annals of the South African Museum 48(9):195-— 210. Bovallius, C. 1886. Notes on the Family Asellidae. — Bihang till Kongliga Svenska Vetenskaps-Aka- demiens Handlingar, Stockholm 11(15):3-54. Coineau, N. 1977. La faune terrestre de l’ile de Sainte- Héléne. 4. Isopodes aquatiques.—Annales du Musée Royal de I|’Afrique Centrale, Tervuren, Belgique, ser. 8°, Sciences Zoologiques, 220:427— 443. Dunham, P. J.,.& A.M. Hurshman. 1991. Precopula- tory mate guarding in aquatic Crustacea: Gam- marus lawrencianus as a model system. Pp. 50- 66, in R. T. Bauer & J. W. Martin, eds., Crus- tacean Sexual Biology. Columbia University Press, New York. Franke, H.-D. 1993. Mating system of the commen- sal marine isopod Jaera hopeana (Crustacea). — Marine Biology 115:65-73. Gruner, H.-E. 1965. Krebstiere oder Crustacea V. Isopoda.— Die Tierwelt Deutschlands und der angrenzenden Meeresteile, nach ihre Merkma- len und nach ihrer Lebensweise 51:1-149. Hessler, R. R., & J.-O. Stromberg. 1989. Behaviour of Janiroidean isopods (Asellota), with special reference to deep-sea genera.—Sarsia 74:145-— 159. Lewis, J. J.,& T.E. Bowman. 1981. Thesubterranean asellids (Caecidotea) of Illinois (Crustacea: Isop- oda: Asellidae). —Smithsonian Contributions to Zoology 335:1-66. Menzies, R. J., & J. L. Barnard. 1951. The isopodan genus /ais (Crustacea).— Bulletin of the Southern California Academy of Sciences 50(3):136-151. Nierstrasz, H. F. 1941. Die Isopoden der Siboga- Expedition. IV. Isopoda Genuina III. Gnathi- 282 idea, Anthuridea, Valvifera, Asellota, Phreato- coidea.—Siboga-Expeditie monographie 32d: 235-306. Ridley, M. 1983. The explanation of organic diver- sity: the comparative method and adaptations for mating. Clarendon Press, Oxford, 272 pp. Sivertsen, E., & L. B. Holthuis. 1980. The marine isopod Crustacea of the Tristan da Cunha Ar- chipelago.—Gunneria 35:1—128. Swedmark, B. 1964. The interstitial fauna of marine sand.— Biological Reviews 39:1—42. Veuille, M. 1978. Biologie de la reproduction chez Jaera (Isopode Asellote). 2. Evolution des or- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ganes reproducteurs femelles. — Cahiers de Biol- ogie Marine 19:385-395. . 1980. Sexual behaviour and evolution of sex- ual dimorphism in body size in Jaera (Isopoda Asellota).— Biological Journal of the Linnean Society 13:89-100. Department of Invertebrate Zoology, Na- tional Museum of Natural History, Smith- sonian Institution, Washington, D.C. 20560, U.S.A. PROC. BIOL. SOC. WASH. 107(2), 1994, pp. 283-290 A NEW GENUS AND SPECIES OF CIROLANID ISOPOD FROM THE WESTERN INDIAN OCEAN (CRUSTACEA: PERACARIDA) Brian Kensley and Marilyn Schotte Abstract. —Seychellana expansa new genus, new species, is diagnosed and described from shallow water off the Seychelles Islands, western Indian Ocean. The genus is characterized by the possession of a slender, blade-like frontal lamina, a somewhat reduced pereopod 7, a strongly bipartite epimeron on pleonite 2, a broadly expanded epimeron on pleonite 3, and a ventrally inserted uropod. Records of the cirolanid isopod fauna of the western Indian Ocean are scattered through a number of publications (see Bruce & Jones 1978, Bruce & Javed 1987, Javed & Yasmeen 1990); the steady stream of new taxa being described, though, would indi- cate that many new forms await discovery. To date, the majority of records are from coastal intertidal and very shallow waters. Very little is known of the peracaridan fauna of the tropical oceanic islands of the Indian Ocean, as is demonstrated by the present record of a new genus from what is relatively shallow water in the Seychelles, an area that has received less attention than some. The material was collected in the course of an environmental survey carried out for the government of the Republic of Sey- chelles, and was submitted to the Smith- sonian Institution for identification. Suborder Flabellifera Family Cirolanidae Seychellana, new genus Diagnosis.—Cephalon lacking rostrum. Frontal lamina slender, blade-like, poste- riorly well separated from flattened, non- projecting clypeus. Mandibular molar not reduced; incisor broad. Maxillipedal endite having 2 coupling hooks. Pereopods 1-3 with merus anterodistally somewhat pro- duced. Pereopods 4—7 ambulatory, articles not markedly flattened, bases lacking na- tatory setae. Pereopod 7 noticeably shorter and more slender than pereopod 6. Epi- meron of pleonite 2 bipartite, consisting of strong acute dorsal and ventral parts. Epi- meron of pleonite 3 enormously enlarged, overlapping pleonites 4 and 5 laterally. Pleonite 5 with free lateral margin over- lapped by epimeron 3. Endopods of pleo- pods 3 and 4 with marginal setae, of pleopod 5 lacking setae; all exopods with marginal setae. Uropod inserted ventrally; peduncle produced along mesial margin of endopod; latter fairly slender, lateral margin excavate. Type species.—By present designation, Seychellana expansa, new species. Discussion. — The present material essen- tially resembles a species of Eurydice, over- lain with a number of specializations. The similarities with Eurydice can be seen in the structure of the frontal lamina, clypeus, an- tennule, pleon, and in the ventral insertion of the uropods. The specializations include the reduced pereopod 7 (one-third shorter and less robust than pereopod 6); the lack of a complete transverse suture on any of the pleopodal rami; the strongly bipartite epimeron of pleonite 2 embracing that of pleonite 3; the enormously enlarged epi- meron of pleonite 3 which completely over- laps those of pleonites 4 and 5. While some 284 of these features appear individually in some other cirolanid genera, e.g., the slender fron- tal lamina of Pseudolana, Eurydice, and Na- tatolana, the plate-like epimera of pleonites 2 and 3 in Booralana, the strongly expanded epimeron of pleonite 3 as in Aatolana schioedtei (Miers 1884) (see Bruce 1986, fig. 135c; 1993:5, 9), none of the presently rec- ognized genera exhibit all of these special- izations, along with a considerably reduced pereopod 7 (compared to pereopod 6), and ventrally inserted uropods. Etymology. — The generic name is a com- bination of Seychelles, the type locality, and ‘lana’, the suffix frequently used in cirolanid taxonomy. Seychellana expansa, new species Figs. 1-4 Material. —Holotype, USNM 252889, ¢ tl 6.6 mm, Paratypes, USNM 252890, ¢ tl 6.3 mm, 2 tl 4.2 mm, sta C2R1, 6/1/92; 6 tl 6.0 mm, SEM whole mount, sta C2R2; 6 tl 6.4 mm, sta C2R3, 6/1/92 (dissected); 4°41'N, 55°33’E, 30 m, Van Veen grab on sandy bottom, off Anse aux Pins, Mahé, Seychelles, coll. J. Elliott, S. Elliott, & P. Harper, | Jun 1992. Description. —Male: Body length about 3 times greatest width at pereonite 3. Red- brown chromatophores scattered over most of dorsum of cephalon, pereonites and pleonites, pleotelson dorsally almost unpig- mented. Cephalon with anterior margin mesially broadly rounded but lacking dis- tinct rostrum; bases of antennules separat- ing cephalon from frontal lamina; latter an- teriorly rounded in ventral view, posteriorly elongate, narrowed, well separated from broadly rounded clypeus. Eyes large, well pigmented. Coxal plates of pereonites 2 and 3 posteriorly truncate, of pereonites 4-6 be- coming larger and slightly produced, of pe- reonite 7 posteriorly acute, much smaller than and almost hidden by coxa of pereonite 6. Posterior margins of pereonites and pleo- nites finely denticulate. Pleonites 1—5 free; PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON pleonite 1 short, with very short free epi- meron; pleonites 2—4 subequal in length, epimeron of pleonite 2 bipartite, with strong acute dorsal and ventral portions embracing enlarged epimeron of pleonite 3; latter broadly ovate, reaching posteriorly beyond pleonites to protopod of uropod, completely covering short epimera of pleonites 4 and 5; pleonite 5 longer than 4, with posterior margin mesially convex. Pleotelson as long as wide, strongly arched, with finely cren- ulate posterior margin broadly rounded, bearing 12 short plumose setae; lateral and posterior margins directed ventrally. Antennular peduncle with articles 2 and 3 each bearing 2 spines on inner (ventral) surface; flagellum consisting of 9 articles, row of 3 or 4 aesthetascs on articles 2-7, penultimate article with single aesthetasc. Antennal peduncle articles | and 2 short, articles 3—5 increasing in length distally, ar- ticle 4 with 2 stout distal spines, article 5 with 5 distal spines; flagellum of 21 articles, reaching posteriorly to level of pereonite 5. Mandibular incisor of 3 cusps on each side, mesial cusp somewhat elongate and acute, 2 lateral cusps rounded; 7 or 8 spines in spine row; molar having row of 20 spines on upper surface; palp of 3 articles, article 1 subequal in length to article 3; article 2 2.5 times length of article 1, bearing 10 setae on distal half of lateral margin; article 3 bearing 13 fringed spines increasing in length distally. Maxilla 1, endopod having 3 stout mesiodistal circumplumose setae; exopod having 8 distal entire, comb, and denticu- late setae. Maxilla 2, palp bearing 2 distal elongate setae; exopod with 7 mesiodistal setae; endopod broadly rounded, with sev- eral simple and circumplumose setae on mesial margin. Maxillipedal endite reaching base of article 2 of palp, bearing 4 sparsely plumose setae distally, 2 coupling hooks on mesial margin of both left and right ap- pendage; palp articles 2—5 bearing setae on mesial and lateral margins. Pereopod 1, merus somewhat anterodis- tally produced, bearing strong terminal VOLUME 107, NUMBER 2 285 Fig. 1. Seychellana expansa. A, Holotype in dorsal view, scale = 1 mm; B, Holotype in lateral view; C, Lateral pleon enlarged, with epimera 1, 2, and 3 indicated. D, Antennule; E, Antenna; F, Left and right mandibles; G, Maxilla 1; H, Maxilla 2; I, Maxilliped; J, Mandibular palp; K, Uropod. 286 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. Seychellana expansa. Pereopods 1-7. VOLUME 107, NUMBER 2 Fig. 3. Seychellana expansa. Pleopods 1-5. spine, posterior margin having single elon- gate and several short spines; carpus short, triangular, lacking free anterior margin; propodus with 5 short spines on posterior margin; dactylar unguis lacking accessory spine. Pereopod 2, basis with 2 elongate se- tae on anterior margin; ischium with single strong anteromesial and posteromesial spine; merus with short anteromesial lobe bearing several spines; carpus having free anterior and posterior margin, with 2 short posterodistal spines; propodus with 2 pos- terodistal spines. Pereopod 3, basis with row of palmate setae on anterior margin; ischi- um with few small spines at posterodistal angle, single strong spine at anterodistal an- gle; merus with several strong anterodistal spines, posterior margin sinuous, bearing 288 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 4. Seychellana expansa. A, Cephalon, antennular bases, and apex of frontal lamina in dorsal view; B, Anterior cephalon in ventral view; C, Dactylar unguis of pereopod |; D, Dactylar unguis of pereopod 5; E, Pleon in lateral view; F, Pleon in ventral view, showing ventral insertion of uropods (protopods indicated by arrows). VOLUME 107, NUMBER 2 about 7 short and long spines; carpus, prop- odus, and dactylus as in pereopod 2. Pereo- pod 4, basis with row of elongate setae on lateral surface; ischium with anterodistal clump of spines, clump of spines at mid- length and at posterodistal angle of poste- rior margin; merus and carpus each with anterodistal and posterodistal clump of spines; propodus slender, elongate, with sin- gle spine at about midlength of posterior margin, 2 small posterodistal spines. Pereo- pods 5 and 6 similar, basis with double row of setae; ischium with 3 clumps of spines on posterior margin, clump of anterodistal spines; merus and carpus with clump of spines at midlength of posterior margin, plus anterodistal and posterodistal clump; prop- odus with clump of spines at midlength of posterior margin, 2 small posterodistal spines. Pereopod 7 about one-third shorter and less robust than pereopods 5 and 6, basis with double row of setae; ischium with slender spines and setae on anterior and posterior margins; merus with clumps of slender spines at midlength of posterior Margin, anterodistal and posterodistal an- gles; carpus with numerous slender distal spines; propodus with single spine at mid- length of posterior margin, 2 spines at pos- terodistal and anterodistal angles. Penes on sternite 7 well separated low rounded struc- tures. Pleopod 1, protopod with 4 retinaculae; endopod slightly longer and narrower than exopod, with few distal plumose marginal setae; exopod ovate, with distal third of me- sial margin and all of lateral margin bearing plumose setae. Pleopod 2, protopod with 3 retinaculae; copulatory stylet articulating at base of endopod, slender, distally curved and barely surpassing apex of rami; ovate exopod with plumose marginal setae on dis- tal third of mesial margin and all of lateral margin. Pleopods 3 and 4 similar, protopod with 3 retinaculae; endopod with mesial margin straight, few plumose marginal setae distally; exopod ovate, with plumose mar- ginal setae on distal third of mesial margin 289 and all of lateral margin, latter having in- complete suture forming small notch at about midlength. Pleopod 5, protopod lack- ing retinaculae; endopod ovate, having rounded proximomesial lobe; exopod ovate, bearing plumose marginal setae on distal third of mesial margin and all of lateral mar- gin, with incomplete suture forming small notch just short of midlength of both mesial and lateral margins. Uropodal protopod in- serted ventrally and completely hidden by pleotelson in dorsal view, produced dis- tomesially for about one-third length of en- dopod; latter with lateral margin excavate, ramus appearing flexed, with numerous se- tae on distomesial margin, few spines and setae on lateral margin; exopod slender, lan- ceolate, with 4 short spines and few setae along lateral margin, several setae along me- sial margin, apex having 2 short spines. Etymology.—The specific epithet, from the Latin expansus, expanded or spread out, refers to the expanded character of the epi- meron of pleonite 3. Acknowledgments We thank Jim Elliott, Susan Elliott, and Patricia Harper of Ogden Environmental and Energy Services of San Diego, Califor- nia, for collecting the material and making it available for study. The material was col- lected under a subcontract to Black and Veatch International. Funding was provid- ed by The African Development Bank through a contract let and administered by the Seychelles Public Utilities Corporation. We thank Mrs. Susann Braden for assis- tance with production of the SEM photo- graphs. We are grateful to Dr. Richard Brus- ca, The University of Charleston, South Carolina, and to two anonymous reviewers for their valuable comments on this paper. Literature Cited Bruce, N. L. 1986. Cirolanidae (Crustacea: Isopoda) of Australia.—Records of the Australian Mu- seum, supplement 6:1—240. 290 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON . 1993. Two new genera of marine isopod crus- taceans (Ciro lanidae) from Madang, Papua New Guinea.— Memoirs of the Queensland Museum 31(1):1-15. ——, & W. Javed. 1987. A new genus and two new species of cirolanid isopod Crustacea from the northern Indian Ocean.—Journal of Natural History 21:1451-1460. —,&D.A.Jones. 1978. Thesystematics of some Red Sea Isopoda (Family Cirolanidae) with de- scriptions of two new species.—Journal of Zo- ology, London 185:395-413. Javed, W., & R. Yasmeen. 1990. A new species of cirolanid isopod of the genus Neocirolana from Pakistan with a review of the genus. —Crusta- ceana 58(1):67-73. Miers, E. J. 1884. Crustacea. Isopoda. Pp. 299-311, Report of the zoological collections made in the Indo-Pacific Ocean during the voyage of H.M.S. “Alert,” 1881-1882. Trustees of the British Mu- seum (Natural History), London. Department of Invertebrate Zoology, Na- tional Museum of Natural History, Smith- sonian Institution, Washington, D.C. 20560, U.S.A. PROC. BIOL. SOC. WASH. 107(2), 1994, pp. 291-295 A NEW SPECIES OF PALAEMONETES (CRUSTACEA: DECAPODA: PALAEMONIDAE) FROM NORTHEASTERN MEXICO Ned E. Strenth Abstract. —Palaemonetes hobbsi is described from the headwaters of the Rio Mante near Ciudad Mante in the state of Tamaulipas, México. While this new species is similar to Palaemonetes mexicanus Strenth from the nearby state of San Luis Potosi, it differs with respect to rostral dentition, first maxilliped morphology, and spination of the appendix masculina. It also exhibits a disjunct nonoverlapping range. Standard starch gel electrophoretic comparisons also confirm that this new taxon exhibits biochemical differences that are distinct from P. mexicanus. Restimen. —Se describe una nueva especie de Palaemonetes de agua dulce de Mexico: Esta especie de camaron ocurre en las aguas de cabecera del Rio Mante y Rio Frio cerca de Ciudad Mante en el estado de Tamaulipas. Es similar a Palaemonetes mexicanus Strenth conocida del estado de San Luis Potosi. Se distingue esta especie nueva de la especie P. mexicanus por las diferencias en la denticion del rostro, en la morfologia del primer maxilipedo, en la disposicion de las espinas del appendix masculina, y el analisis de bioquimica. Following a review (Strenth 1976) of the North American species of freshwater Pa- laemonetes Heller, continued field work in northeastern Mexico revealed the presence of several populations of shrimp from two large springs near Ciudad Mante in the state of Tamaulipas. It was apparent from col- lections made during 1980 and 1983 that these populations differed morphologically only slightly from specimens of P. mexi- canus Strenth from San Luis Potosi. Field work was continued in Mexico during the middle and late 1980’s to establish the range of P. mexicanus as well as the Ciudad Mante populations. It was during this time that Hobbs & Hobbs (1989) reported the pres- ence of these shrimp as a range extension of P. mexicanus. Subsequent field work now appears to support the conclusion that these populations are in fact quite distinct and separate in their distribution from that of P. mexicanus. This undescribed taxon is restricted to the springfed headwaters of the Rio Mante and Rio Frio. More importantly, this aquatic decapod inhabits the headwaters of a sep- arate and independent drainage system from that of Palaemonetes kadiakensis Rathbun or P. mexicanus which are also known from northern Mexico (Strenth 1976). The Rio Mante and Rio Frio flow into the Rio Gua- yalejo which is a part of the Rio Tamesi system. Palaemonetes kadiakensis is not known to occur south of the Rio Bravo del Norte (Rio Grande) drainage system and the distribution of P. mexicanus 1s restricted to the headwaters of a small stream which flows into the Rio Tampaon which is a part of the Rio Panuco drainage system to the south in the states of San Luis Potosi and Veracruz. Extensive downstream collec- tions from the type locality of P. mexicanus have failed to produce specimens from ei- ther the Rio Tamuin or the Rio Panuco. 292 Similar downstream collections from the Rio Mante and Rio Frio conducted in the Rio Guayalejo and Rio Tamesi have like- wise failed to yield specimens of freshwater Palaemonetes. Recent laboratory analyses confirm that the Rio Mante population exhibits bio- chemical differences that are distinct from P. mexicanus. It is now evident that this population represents a currently unde- scribed taxon. This conclusion is based upon morphological differences, restricted distri- bution, and the absence of overlapping ranges of this form with any other currently known species of Palaemonetes. Materials and Methods Specimens selected for biochemical anal- ysis were immediately frozen in liquid ni- trogen in the field and returned to the lab- oratory. Abdominal tissue samples of 20 specimens collected from the headwaters of the Rio Mante were biochemically com- pared with similar tissue samples of 20 spec- imens of P. mexicanus from the type- locality west of Ciudad Valles in San Luis Potosi. These samples were subjected to electrophoretic analysis using standard hor- izontal starch gel techniques (Selander et al. 1971, Murphy et al. 1990). Four of 11 iso- zyme systems examined were observed to exhibit significant variation between the two populations. Specimens which were pre- served in alcohol upon collection are de- posited in the National Museum of Natural History, Smithsonian Institution, Washing- ton, D.C. (USNM) and the Instituto de Bio- logia de la Universidad Nacional Auton- oma de México, Mexico City (EM). Palaemonetes hobbsi, new species Fig. | Palaemonetes mexicanus Strenth, 1976.— Hobbs & Hobbs, 1989:222. Type material. —Nacimiento del Rio Mante, 8 km southwest of Ciudad Mante, PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Tamaulipas, México, 20 May 1980, coll. N. E. Strenth. Male holotype, USNM 264749; one male, one gravid female, paratypes, USNM 264748; one male, one gravid fe- male, paratypes, EM 11972. Description. — Rostrum (Fig. 1a) high; up- turned at end; extending to near anterior margin of scaphocerite; dorsal margin with 5 to 6 teeth, one of which is placed behind the orbit; ventral margin with 2 teeth. An- tennal spine sharp, distinct; overreaching anterior margin of carapace. Branchiostegal spine sharp and distinct; situated on ante- rior margin of carapace just below bran- chiostegal groove. Abdomen normal; pleura of third somite rounded; pleurae of fourth and fifth somites angular; sixth somite 1.5 times as long as fifth. Telson (Fig. 1b) equal in length to sixth somite; anterior pair of dorsal spines locat- ed 4 of telson length from posterior margin; posterior pair on or near posterior margin. Posterior margin of telson with sharp me- dian point flanked by 2 pair of spines and 1 pair of plumose setae; lateral pair of spines short, failing to reach median point of tel- son; mesial pair of spines elongate extending 3 to 4 times length of lateral spines. Eyes (Fig. la) well developed; cornea globular and well pigmented. Stylocerite (Fig. le) sharp and extending % length of basal segment of antennular peduncle; an- terolateral spine sharp, extending to ante- rior margin of segment; lateral antennular flagellum with rami fused for 13 to 22 ar- ticles; free portion of shorter ramus con- sisting of 3 to 7 articles. Scaphocerite (Fig. 1d) almost 3 times as long as wide, lateral margin slightly concave, blade extending well past lateral tooth. Mouthparts (Figs. 1 f-k) typical for the ge- nus. First maxilliped (Fig. 11) with endopod; epipod nonbifurcate. First pereopod (Fig. 11) extending to distal margin of third seg- ment of antennular peduncle; dactyl and propodus without teeth, bearing numerous setae distally; carpus twice as long as chela, 1.2 times as long as merus. Second pereopod VOLUME 107, NUMBER 2 (Fig. 1m) extending to anterior margin of scaphocerite; dactyl and propodus without teeth, bearing few setae distally; carpus 1.4 times as long as chela; merus and chela equal in length. Third pereopod (Fig. 1n) extend- ing to base of third segment of antennular peduncle; propodus 1.8 times as long as car- pus; merus 1.9 times as long as carpus. Fourth pereopod extending to anterior mar- gin of second segment of antennular pedun- cle; fifth pereopod extending to near ante- rior margin of scaphocerite. Appendix masculina (Fig. 1c) of male with 4 apical spines which extend to distal mar- gin of endopod; appendix interna '2 length of appendix masculina. Eggs of gravid fe- males 1.2—-1.5 mm in length. Lateral ramus of uropod with or without movable spine between fixed distolateral tooth and margin of blade. Size. — Males with carapace lengths to 6 mm (including rostrum, to 11 mm); fe- males, to 7 mm (including rostrum, to 12 mm). Variation.—The movable exopod spine on the lateral ramus of the uropod is quite variable; both movable spines may be pres- ent, both may be absent, or only a left or right one may be present. No specimens were observed to exhibit a subapical tooth on the ventral margin of the rostrum. Color.—Living specimens are transpar- ent. The eggs of gravid females are dark green in coloration. Range. —This species is currently known only from the headwaters of the Rio Mante and Rio Frio near Ciudad Mante in the state of Tamaulipas, México. Etymology. —This species is named in honor of Dr. Horton H. Hobbs, Jr. of the Smithsonian Institution in Washington D.C., and Dr. Horton H. Hobbs III of Wit- tenberg University, Springfield, Ohio. Remarks.— Palaemonetes hobbsi shares characteristics with P. mexicanus as well as with Palaemonetes texanus Strenth from the southwestern United States. All three spe- cies are similar in the variation of movable 293 exopod spination of the uropods. Palae- monetes hobbsi is similar to P. mexicanus with respect to the number of fused articles of the lateral antennular flagellum. Palae- monetes hobbsi ranges from 13-22 fused ar- ticles while P. mexicanus ranges from 14- 24. Palaemonetes texanus differs somewhat in exhibiting 19-29 fused articles. Palae- monetes hobbsi is similar to P. texanus in that both species exhibit angular pleurae of the fourth and fifth somites. The first max- illiped of P. hobbsi is similar to P. texanus in exhibiting an endopod; this same struc- ture is also similar to P. mexicanus in pos- sessing a non-bifurcated epipod. This in- termediacy of characters is possibly related to the fact that the distribution of P. hobbsi lies between the distributions of both P. mexicanus and P. texanus. All three species appear to be closely related. Palaemonetes hobbsi differs, however, from both P. mexicanus and P. texanus with respect to rostral dentition and spination of the appendices masculinae. Palaemonetes hobbsi exhibits only five or six dorsal rostral teeth while P. mexicanus exhibits six or sev- en and P. texanus ranges from five to eight. Palaemonetes hobbsi exhibits four apical se- tae on the appendix masculina while P. mexicanus and P. texanus exhibit five and six apical spines respectively. In addition, there is virtually no significant difference in the size of adult male and female specimens of P. hobbsi. Female specimens exhibit car- apace lengths that are less than 10% longer than male specimens. Female specimens of both P. mexicanus and P. texanus are con- siderably larger and exhibit carapace lengths that are 28 to 45% longer than their male counterparts. In addition to the above anatomical dif- ferences, preliminary electrophoretic anal- yses of both P. mexicanus and P. hobbsi reveal differences in the isozyme production of malate dehydrogenase 1, malate dehy- drogenase 2, phosphoglucomutase and a-glycerophosphate dehydrogenase. These differences are considered significant in light PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ee sai & Se J k m n Palaemonetes hobbsi, new species, holotype, male: a, anterior region; b, telson and uropods; c, appendix masculina; d, first antenna; e, second antenna; f, mandible; g, first maxilla; h, second maxilla; i, first maxilliped; j, second maxilliped; k, third maxilliped; 1, first pereopod; m, second pereopod; n, third pereopod. (Scales = 1.0 mm). Fig. 1. VOLUME 107, NUMBER 2 of the fact that palaemonids are known to exhibit low levels of allozyme variation (Boulton & Knott 1984). While the exact nature of the above biochemical variation is under current investigation, it is inter- preted here only as additional supportive evidence of the premise that P. hobbsi rep- resents a taxon which is morphologically distinct and geographically separate from P. mexicanus. Acknowledgments This study was supported by a Research Enhancement Grant from Angelo State University. Special thanks go to T. G. Lit- tleton and J. T. Collins for the electropho- retic analyses and interpretation. Appreci- ation is extended to M. Onofre-Madrid for assistance with the Spanish translation of the abstract. Literature Cited Boulton, A. J., & B. Knott. 1984. Morphological and electrophoretic studies of the Palaemonidae 295 (Crustacea) of the Perth Region, Western Aus- tralia.—Australian Journal of Marine and Freshwater Research 35:769-783. Hobbs, H. H., Jr., & H. H. Hobbs III. 1989. New locality records for two poorly known Mexican freshwater shrimps (Decapoda, Palaemoni- dae). —Crustaceana 57:220-—222. Murphy, R. W., J. W. Sites, Jr., D. G. Buth, & C. H. Haufler. 1990. Proteins I: Isozyme Electro- phoresis. Pp. 45-126 in D. M. Hillis & C. Mo- ritz, eds., Molecular Systematics. Sinauer As- sociates, Inc., Sunderland, Massachusetts, 588 pp. Selander, R. K., M. H. Smith, & S. Y. Yang. 1971. Biochemical polymorphisms and systematics in the genus Peromyscus. 1. Variation in the old- field mouse (Peromyscus polionotus).—Univer- sity of Texas Studies in Genetics 6:49-90. Strenth, N. E. 1976. A review of the systematics and zoogeography of the freshwater species of Pa- laemonetes Heller of North America.—Smith- sonian Contributions to Zoology 228:1-28. Department of Biology, Angelo State University, San Angelo, Texas 76909, U.S.A. PROC. BIOL. SOC. WASH. 107(2), 1994, pp. 296-307 A REVISION OF THE TYPE MATERIAL OF SOME SPECIES OF HYPOLOBOCERA AND PTYCHOPHALLUS (CRUSTACEA: DECAPODA: PSEUDOTHELPHUSIDAE) IN THE NATIONAL MUSEUM OF NATURAL HISTORY, WASHINGTON, D.C., WITH DESCRIPTIONS OF A NEW SPECIES AND A NEW SUBSPECIES Gilberto Rodriguez Abstract. —The descriptions of a number of species of freshwater crabs from Panama and Colombia held in the USNM are revised. These are Hypolobocera andagoensis (Pretzmann 1965), H. beieri Pretzmann, 1968, H. canaensis Pretz- mann, 1968, H. martelathami (Pretzmann 1965), and H. smalleyi Pretzmann, 1968, and three species of Ptychophallus from Panama, P. cocleensis Pretz- mann, 1965, P. goldmanni Pretzmann, 1965, and P. lavallensis Pretzmann, 1978. The present work provides illustrations of the first gonopods of the above species and corrects several inaccuracies in their descriptions and those of H. chocoensis Rodriguez, 1980, and P. colombianus (Rathbun 1893). The collec- tions included a new species, Ptychophallus micracanthus, and a new subspecies, Hypolobocera bouvieri rotundilobata, which also are described. Due to the scarcity of reliable taxonomic characters in the carapace and appendages of the Pseudothelphusidae, the systematics of this family of neotropical freshwater crabs is based almost exclusively on the mor- phology of the male first gonopods. For this reason, the taxonomic status of species de- scribed without an adequate illustration of these appendages is uncertain (Rodriguez 1982). This is the case with three lots of crabs obtained by E. A. Goldman near the Panama Canal in 1912, and by Marte La- tham in Colombia in 1957 and 1962, from which Pretzmann (1965, 1968) described five new species of Hypolobocera. Unfor- tunately illustrations of the first gonopods were not included with the original descrip- tions, and a latter work (Pretzmann 1972) provided only photographs in which mor- phological details are not discernible. De- scriptions of the first gonopods were not in- cluded for the species of Ptychophallus from Panama obtained by several collectors, and described by Pretzmann in 1965, 1978, and 1980. The present work provides illustra- tions of the gonopods of the holotypes and other materials of these species in the col- lections of the National Museum of Natural History, Smithsonian Institution, Washing- ton, D.C. (USNM), and corrects several in- accuracies in the descriptions of these taxa. A new species of Ptychophallus and a new subspecies of Hypolobocera bouvieri were discovered among the materials in these collections. Abbreviations used are cb. for carapace breadth, and cl. for carapace length. Pseudothelphusidae Ortmann, 1893 Genus Hypolobocera Ortmann, 1897 Hypolobocera andagoensis (Pretzmann, 1965) Fig. 1A-—C Strengeria (Strengeria) andagoensis Pretz- mann, 1965:6. Hypolobocera (Hypolobocera) andagoen- VOLUME 107, NUMBER 2 sis. —Pretzmann, 1971:17; 1972:51, figs. 170-172. Hypolobocera andagoensis. — Rodriguez, 1982:67, figs. 21c, e.—von Prahl, 1988: 183. Material. —Colombia: Andagoya, Choco, May 1957, leg. Marte Latham, No. 996, 1 male holotype, cl. 19.2 mm, cb. 31.6 mm (USNM 106405).— Colombia: no data, leg. Marte Latham, 1957, 15 males, largest cl. 21.3 mm, cb. 35.8 mm, smallest cl. 11.9 mm, cb. 18.2 mm, 2 females, cl. 17.6 and 21.0 mm, cb. 29.0 and 35.0 mm (USNM 106407).—Colombia: no data, leg. Marte Latham, 23 males, 28 females (USNM 106409). Diagnosis.— First gonopod with caudal ridge long, straight; lateral lobe prominent, square, faintly crenulated; apex small, rounded, strangled. Endognath of first max- illiped strongly reduced, approximately 0.25 length of ischium of endognath. Remarks.—In his original description Pretzmann (1965) designated the male (cb. = 31.6 mm) specimen in lot USNM 106405 as holotype, but later (Pretzmann 1972) stated the number of the holotype to be USNM 106407. In the same work 14 males and two females in lot “106405” (presum- ably 106407) were designated as paratypes and that author incorrectly stated that the two lots were from the same locality. Pretzmann (1972) illustrated the whole animal (figs. 170, 171; the negatives were inverted so that the animal appears left handed), and a first gonopod which had been removed from the animal (figs. 311, 312). This appendage, however, does not belong to the holotype (USNM 106405) since this specimen still had the first gonopods at- tached until the present author removed them. The first gonopod of the original ho- lotype of H. andagoensis (USNM 106405) is illustrated in figs. 1A—C. Hypolobocera beieri Pretzmann, 1968 Fig. 2D 297 Hypolobocera (Hypolobocera) bouvieri beieri Pretzmann, 1968:9; 1971:17; 1972:46, figs. 176-181, 308, 309. Hypolobocera beieri. —Rodriguez, 1982:46, figs. 19a, 1, 20b, g, 24 a-—d.—von Prahl, 1988:172, fig 2. Hypolobocera (Hypolobocera) monticola steindachneri. —Pretzmann, 1972:46 (part). Material.—Colombia: Bitaco, Valle del Cauca, Andes Occidentales, May 1957, leg. Marte Latham, | male holotype, cl. 22.8 mm, cb. 40.6 mm (USNM 106410).—Same, 2 male paratypes, cl. 18.9 and 17.0 mm, cb. 33.5 and 27.2 mm (USNM 128123 ex 106410).—Mountainous area of upper San Juan River, Choco jungle, Department of Choco, nearest village Playa de Oro, 28 Mar 1962, leg. Marte Latham, 3 males, cl. 46.2, 32.2 and 31.5 mm, cb. 68.4, 49.9 and 46.9 mm (USNM 240101). Diagnosis. — First gonopod with caudal ridge straight, well defined; lateral lobe small, rounded, placed relatively far from apex, with conspicuous depression below; apex small, rounded, moderately strangled, with two rounded and flat papillae on distal sur- face. Endognath of first maxilliped strongly reduced, approximately 0.3 length of ischi- um of endognath. Remarks. —Pretzmann (1972) reassigned the paratypes of this species to Hypolobo- cera monticola steindachneri, but characters of the gonopods and carapace clearly place these paratypes in H. beieri. Hypolobocera bouvieri rotundilobata, new subspecies Fig. 2A—C Material. —Colombia: Mountains of up- per San Juan River, Choco jungle, Depart- ment of Choco, nearest village Playa de Oro, 28 Mar 1962, leg. Marte Latham, 1 male holotype, cl. 46.2 mm, cb. 68.4 mm (USNM 240103).—Same data, 5 male paratypes, largest cl. 32.2 and 31.5 mm, cb. 49.9 and 46.9 mm (USNM 240104). 298 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON O Imm AYN sages SS wee Fig. 1. A-C, Hypolobocera andagoensis (Pretzmann), holotype, USNM 106405, left gonopod; D-F, Hypo- lobocera martelathami (Pretzmann), holotype, USNM 106408, right gonopod; G-I, Hypolobocera smalleyi Pretzmann, holotype, USNM 54042, left gonopod; J-N, Hypolobocera canaensis Pretzmann, holotype, USNM 54039, left gonopod. A, D, G, J, caudal view; B, E, H, K, lateral view; C, F, I, L, apex, distal view. Diagnosis. —Caudal ridge of first gono- pod strong, with transverse wrinkles; lateral lobe small, semicircular, covered by minute pores and scattered short setae, without crenulations over distal margin; apex fun- nel-form, subtriangular in distal view. En- dognath of first maxilliped reduced, ap- proximately 0.4 length of ischium of VOLUME 107, NUMBER 2 endognath. Fixed finger of chelipeds with- out conspicuous tubercle at its base. Upper margin of front advanced, angled, marked by conspicuous row of coalescent papillae. Description (based on holotype and 5 male paratypes). — Carapace relatively narrow (cb/ cl = 1.48—-1.55). Cervical grooves sinuous, deep, narrow, attaining margins of cara- pace. Postfrontal lobes small, rounded, well marked; median grooves not deeply im- pressed between lobes, but making deep in- cision on upper margin of front; this margin advanced, marked by conspicuous row of coalescent papillae and strongly bilobed in dorsal view; lower orbital margin strongly bent upward in holotype (probably a mal- formation). Exognath of third maxilliped 0.38-0.40 length of ischium of endognath. Cheliped without a large tubercle over base of fixed finger, but with scattered papillae over palm and merus, particularly on inner surface; fingers tinged reddish brown. First gonopod slender, with small semicircular lateral lobe, covered by minute pores and scattered short setae; strong caudal rib with transverse wrinkles; apex funnel-form, sub- triangular in distal view. Remarks. —Hypolobocera bouvieri is widely distributed over the Cordilleras of northern South America, with four subspe- cies: H. bouvieri angulata in the Sierra of Santa Marta, Sierra of Perija and the Ven- ezuelan Andes; H. bouvieri bouvieri and H. bouvieri stenolobata in the Eastern Cordil- lera of Colombia; and H. bouvieri monticola on the slopes of the Western Cordillera which descend to the Cauca valley. H. bou- viert rotundilobata is the only subspecies so far recorded from the Pacific slopes of the Andes. The type locality, close to the town of Playa del Oro (5°20'N, 76°23'W), is near the headwaters of the San Juan River, where the areas of distribution of H. andagoensis, H. chocoensis and H. malaguena von Prahl, 1988 overlap. Although the San Juan River runs parallel, and next to, the Cauca River, where H. bouvieri monticola occurs, these two river basins are separated by the water 299 divide of the Western Cordillera of Colom- bia. Hypolobocera bouvieri rotundilobata can be clearly distinguished from the other sub- species by the semicircular lateral lobe of its first gonopod. In addition, the new sub- species differs from H. b. bouvieri and H. b. monticola in the absence of a large tubercle at the base of the fixed fingers of chelipeds; and from H. b. bouvieri, H. b. angulata, and H. b. stenolobata in the absence of crenu- lations over the distal margin of the lateral lobe. The tuberculation of the upper border of the front in the new subspecies somewhat resembles that of H. b. angulata and H. b. stenolobata, but it is clearly different from that of H. b. bouvieri and H. b. monticola. Etymology. —The subspecific name is from the Latin “rotundus,” rounded, and “lobatus,”” lobed, a reference to the shape of the lateral lobe of gonopod. Hypolobocera canaensis Pretzmann, 1968 Fig. 1J—L Hypolobocera (Hypolobocera) canaensis Pretzmann, 1968:3; 1971:17; 1972:47, figs. 211-213, 313-315. Material.—Panama: Cana, altitude 850 m, 24 May 1912, leg. E. A. Goldman (U.S. Biological Survey donation), 1 male holo- type, cl. 32.6 mm, cb. 51.2 mm (USNM 54039).—Cana, altitude 760 m, 21 May 1912, leg. E. A. Goldman (U.S. Biological Survey donation), | immature female para- type (fragmented) (USNM 54036).—Cana, Setiganti River, altitude 610 m, 24 Mar 1912, leg. E. A. Goldman (U.S. Biological Survey donation), | immature female para- type, cl. 45.7 mm, cb. 72.8 mm (USNM 54037).—Cana, altitude 760 m, 1 Jun 1912, leg. E. A. Goldman (U.S. Biological Survey donation), | male with broken carapace, cl. 34.2 mm, cb. 53.8 mm (USNM 54038). Diagnosis. —First gonopod slender, slightly arched; caudal ridge straight, long, narrow; lateral lobe long, narrow, more ex- panded distally; apex funnel-form, elliptic 300 in distal view, with 1 flat rounded papilla on distal surface. Endognath of first max- illiped strongly reduced, approximately 0.3 length of ischium of endognath. Remarks. —It is doubtful whether the two paratype specimens listed by Pretzmann (1972) belong in this species since both are immature females collected at lower alti- tudes on different days. There are only frag- ments of the carapace and some pereopods of one of them (USNM 54036). For details of the type locality Cana see below under Ptychophallus goldmanni. Hypolobocera chocoensis Rodriguez, 1980 Hypolobocera (Hypolobocera) dubia. — Pretzmann, 1972:48, figs. 224-226, 230- 232, 236, 237 (not Pseudothelphusa dubia Colosi, 1920). Hypolobocera chocoensis Rodriguez, 1980: 891; 1982:59, figs. 19f, q, 21b, d, 31 a-d. Material. —Colombia: Mountains of up- per San Juan River, Choco jungle, Depart- ment of Choco, nearest village Playa de Oro, 28 Mar 1962, leg. Marte Latham, 10 males, the largest cl. 22.8 mm, cb. 36.7 mm, 26 females, the largest cl. 23.8 mm, cb. 39.7 mm (USNM 240102). Diagnosis. — First gonopod strongly con- stricted below lateral lobe, with caudal ridge long, irregular; lateral lobe wide proximally, narrow distally; apex funnel shaped, ex- panded, rounded in distal view, with two flat, wide papillae on distal surface and small subtriangular spine on mesial side. Endog- nath of first maxilliped strongly reduced, approximately 0.2 length of ischium of en- dognath. Remarks. —The following characters should be added to the description of the species: The regions of the carapace (Rodri- guez 1982) are strongly marked; the post- frontal area is excavated. The front is well defined by a tuberculated ridge; the surface of the front is conspicuously excavated. The postorbital notch is very deep; the antero- lateral border is not continuous with the PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON margin of this notch, but runs dorsally to it, forming a rounded lobe. The anterior portion of the carapace is covered by rough papillae, barely visible to the naked eye. The male chelipeds are strongly unequal; when fully developed the fingers of the largest che- la are strongly gapping, the dactylus is re- curved; the distal half of the dactylus, the tip of the fixed finger and the teeth of both fingers are brown black. Hypolobocera chocoensis closely resem- bles Hypolobocera malaguena von Prahl, 1988. The lateral lobe of the first gonopod of H. malaguena is smaller and almost rounded; the apex in lateral view is not strangled or funnel-shaped like in H. cho- coensis. In distal view the apex is clearly triangular, not rounded, and possesses a well developed transverse mesial lobe. Other features which separate H. malaguena from H. chocoensis, include the smooth carapace with weakly marked regions and unexca- vated frontal and postorbital regions, the upper border of the front which is weak to- ward the sides, and the anterolateral margin of the carapace which is continuous with the margin of the postorbital notch. Hypolobocera martelathami (Pretzmann, 1965) Fig. 1D-F Strengeria (Strengeria) martelathani (sic) Pretzmann, 1965:6. Hypolobocera (Hypolobocera) martela- thami 1971:17; 1972:50, figs. 159-161, 242-244. Hypolobocera martelathami.— Rodriguez, 1982:52.—Campos & Rodriguez, 1984: 538, figs. 4c, f. Hypolobocera (Hypolobocera) plana orien- talis Pretzmann, 1968:2; 1971:17; 1972: 60, figs. 162-164, 214-221. Hypolobocera orientalis. —Rodriguez, 1982: 52, figs. 19d, 20c, h; 26a-c. Material. —Colombia: No other data, 1957, leg. Marte Latham, | male holotype, VOLUME 107, NUMBER 2 \ ‘\ AST SS \ [((t 301 Fig. 2. A-—C, Hypolobocera bouvieri rotundilobata, new subspecies, holotype, USNM 240103: A, left gonopod caudal view; B, same lateral view; C, same caudal view; D, Hypolobocera beieri Pretzmann, holotype, USNM 106410, caudal view. cl. 16.0, cb. 25.7 mm (USNM 106408).— Colombia: No other data, 1957, leg. Marte Latham, 1 male paratype, cl. 13.0 mm, cb. 20.0 mm, 7 males, cl. 12.7, 12.7, 12.1, 12.0, 10.6, 10.2, 9.8 mm, cb. 19.8, 19.8, 18.7, 15.6, 15.6, 15.2, 14.0 mm, 2 females, cl. 16.7, 15.2 mm, cb. 26.8, 23.8 mm, | imm. female, cl. 12.0 mm, cb. 18.4 mm, 5 juve- niles (USNM 122602). Diagnosis. — First gonopod with distal half wide in caudal and mesial views, caudal ridge straight; lateral lobe large, reaching middle of first gonopod, narrow proximally, wide distally, covered with minute wrin- kles, pores and scattered short setae; apex not funnel-shaped or conspicuously ex- panded, oblong or rectangular in distal view, 2 flat rudimentary papillae on distal surface. Endognath of first maxilliped moderately reduced, approximately 0.55 length of is- chium of endognath. Remarks. —The first gonopod of H. mar- telathami is identical to that of Hypolobo- cera orientalis Pretzmann, 1968. According to Pretzmann (1968) this last species has a broader carapace, and the exognath of third maxilliped is longer, and does not possess an exorbital tooth. Actually the mean cb/cl ratio is 1.54 in both H. martelathami (n = 12) and H. orientalis (n = 10, Rodriguez 1982); the mean ratio of the exognath to ischium of the endognath is 0.65 in H. ori- entalis (Rodriguez 1982) and 0.55 (n = 9; spread 0.46-0.66) in H. martelathami (USNM 106408, USNM 122606); the post- orbital area is similar in both species. Con- sequently, Hypolobocera orientalis should be considered as a junior synonym of Hy- 302 polobocera martelathami, as has already been suggested by Rodriguez (1982). Hy- polobocera plana (Smith 1870) is an incer- tae sedis species (Rodriguez 1982). Hypolobocera smalleyi Pretzmann, 1968 Fig. 1G-I Hypolobocera (Hypolobocera) smalleyi Pretzmann, 1968:4; 1972:50, figs. 233- D355 2X8, 23D, Hypolobocera smalleyi. —Rodriguez, 1982: 60. Material. —Panama: Cana, Canal Zone, altitude 760 m, 1912, leg. E. A. Goldman (U.S. Biological Survey don.), 1 male ho- lotype, carapace broken, cl. 39.5 mm, cb. 24.7 mm (USNM 54042). Diagnosis. —First gonopod with caudal ridge long, straight; lateral lobe prominent, rounded, more expanded distally, with con- spicuous coalescent papillae on cephalic side; apex funnel shaped, expanded, subtrian- gular in distal view, with | acute papilla on distal surface. Endognath of first maxilliped strongly reduced, approximately 0.3 length of ischium of endognath. Genus Ptychophallus Smalley, 1964 Ptychophallus cocleensis Pretzmann, 1965 Fig. 3D, E Ptychophallus (Ptychophallus) montanus cocleensis Pretzmann, 1965:5. Ptychophallus (Ptychophallus) cocleensis. — Pretzmann, 1971:21; 1972:88, figs. SO5— 507, 534, 535. Diagnosis.—First gonopod with lateral projection divided in 2 lobes by deep me- dian notch, distal lobe small, rounded close to apex, proximal lobe finger-like, directed distally; distal caudal ridge short; 2 mesial apical processes large, triangular. Material. —Panama: Rio Coclé del Norte, 1951, leg. M. W. Stirling, | male holotype (USNM 119869).—Barro Colorado Island, Wheeler trail, 9 Jul 1969, leg. R. Foster, 1 male (USNM 230097). PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Ptychophallus colombianus (Rathbun, 1893) Fig. 3A—C Pseudothelphusa colombiana Rathbun, 1893:653, pl. 74, fig. 10, pl. 75, fig. 1; 1898:533.— Young, 1900:219.—Rath- bun, 1905:302.—Coifmann, 1939:107.— Smalley, 1964:10. Ptychophallus (Semiptychophallus) colum- bianus (sic).—Pretzmann, 1965:5. Ptychophallus (Semiptychophallus) colum- bianus (sic) columbianus (sic).—Pretz- mann, 1971:21; 1972:88, figs. 514, 515, 525, 526. Material. —Panama: Chiriqui, David River, about lat. 8°28’N, long. 82°24'W, el- evation 1220 m above sea level, very rapid stream, descending from Mount Chiriqui, Jul 1883, leg. J. A. McNeil, 2 female types (USNM 5512).— Volcan, Rio Chiriqui Vie- jo, | Mar 1924, leg. Foster, 1 male (soft shell) cl. 14.8 mm, cb. 26.8 mm (USNM 58182).—Chiriqui, Hato del Volcan, Pacific drainage, c. 3000 m altitude, 18 Jun 1960, leg. Robinson et al., 5 specimens (USNM 230098). Diagnosis.— First gonopod with lateral projection narrow, with slight depression at middle; distal caudal ridge short, narrow; mesial apical process small, triangular. Remarks.—Rathbun (1893) based her description on two female syntypes. Pretz- mann (1972) assigned to this species the male specimen from Volcan, Panama, men- tioned above (that he stated to be a female, USNM 58182), and two specimens (male and female) from Chiriqui, Panama, in the Zoologisches Museum, Berlin. Examina- tion of the USNM material revealed that the illustration of the carapace in Pretz- mann’s (1972, figs. 514, 515) corresponds to one of the female paratypes (USNM 5512), and that the first gonopod illustrated in figs. 525 and 526 was taken from the specimen from Volcan (USNM 58182). Morphological details of the first gonopods are not clearly discernible from these figures and has been redrawn in the present work. VOLUME 107, NUMBER 2 303 Fig. 3. A-C, Ptychophallus colombianus (Rathbun), USNM 58182: A, left gonopod, caudal view; B, same, meso-caudal view; C, same, apex, distal view; D, E, Ptychophallus cocleensis Pretzmann, holotype, USNM 119869: D, left gonopod, caudal view; E, same, apex, cephalic view; F—H, Ptychophallus goldmanni Pretzmann, holotype, USNM 54044: F, left gonopod, caudal view; G, same, cephalic view; H, same, apex, distal view; I, J, Ptychophallus lavallensis Pretzmann, USNM 240100: I, left gonopod, caudal view; J, same, apex, cephalic view. 304 Ptychophallus goldmanni Pretzmann, 1965 Fig. 3F—H Ptychophallus (Microptychophallus) gold- manni Pretzmann, 1965:5; 1971:21; 1972:90, figs. 527-529, 544-546. Ptychophallus goldmanni. — Rodriguez, 1982:86. Material. —Panama: Cana, 850 m, 24 May 1912, leg. E. A. Goldman, U.S. Biological Survey, | male holotype (USNM 54044). Diagnosis.— First gonopod with lateral projection almost absent, except for small lobe located distally; caudal ridge forming distally large triangular process which ex- tends slightly beyond apex; strong longitu- dinal ridge on cephalic surface ending dis- tally in flat round lobe; mesial apical process rounded. Remarks.—The name “Cana” refers to Mount Cana, altitude 1615 m, 7°48’N, 77°32'W, in Darien Province, Panama. The locality is situated in the northern outskirts of the Serrania del Baudo. The area is drained by the Balsas River which empties into a southern extension of the Gulf of San Miguel. In this mountain, E. A. Goldman also collected Hypolobocera canaensis and H. smalleyi (see above) at altitudes between 610 and 850 m. Ptychophallus lavallensis Pretzmann, 1978 Fig. 3I-J Ptychophallus (Ptychophallus) exillipes la- vallensis Pretzmann, 1978:1; 1980:651— 666. Material. — Panama: Coclé, headwaters of Rio Indio, N of La Mision, above El Valle, where 3 springs begin stream, c. 700 m al- titude, 14 Sep 1962, leg. Loftein and Kosan, 3 males, cl. 21.7, 13.7 and 12.3 mm, cb. 37.0, 21.9 and 19.2 mm (USNM 240100). — El Aguacate, 22 Feb 1973, leg. A. Smalley, 1 male, cl. 23.4 mm, cb. 39.8 mm (USNM 184338).—Same data, 1 male, cl. 23.5, cb. 29.9 mm (USNM 184339). PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Diagnosis.— First gonopod with lateral projection very wide, simple, its proximal margin transverse and slightly concave, its distal margin convex, both with minute se- tae; distal caudal ridge short, weak; apex strongly bent toward cephalic side, making contact with cephalic surface, field of spines directed laterally. Description. —Cervical grooves recurved backward, narrow and shallow; do not reach margins of carapace; anterolateral margins with shallow postfrontal notch, rest of bor- der entire. Postfrontal lobes wide, delimited anteriorly by transverse depression; median groove narrow, deep, making deep incision on upper margin of front. Surface of cara- pace between postfrontal lobes and front flat, horizontal, only slightly inclined down- ward. Upper margin of front slightly convex in dorsal view, thin, well marked, with few small tubercles; lower margin thin, mod- erately sinuous; front between upper and lower margin high, vertical. Exognath of third maxilliped 0.68 length of ischium of endognath. Palm of largest cheliped moderately swollen, lower and up- per margins convex; fingers with rows of black points on external surfaces. Lateral lobe of first gonopod very wide, simple; proximal margin transverse and slightly concave, distal margin convex; apex strong- ly bent toward cephalic side, with field of spines directed laterally; distal caudal ridge short, weak; margin of lateral lobe only have minute setae. Remarks. —This species was only briefly diagnosed by Pretzmann (1978, 1980) and no illustration of the first gonopod was pub- lished. The description given above, which supplement Pretzmann’s short description, is based on the specimens from Rio Indio and El Aguacate, Panama, recorded above. The Rio Indio runs parallel to, and approx- imately 30 km from, the valley of Rio Cocle, where the type locality of the species is found. The first gonopod of P. /avallensis has a wide undivided lobe, like P. exi/lipes (Rath- VOLUME 107, NUMBER 2 b 305 O 2mm Fig. 4. Ptychophallus micracanthus, new species, holotype, USNM 240106: A, left gonopod, caudal view; B, same, apex, cephalic view; C, opening of left efferent channel; D, third maxilliped; s, cephalic spine. bun 1898), but it has a different shape and lacks the long setae; the apex is relatively smaller, more strongly bent toward the ce- phalic side, making contact with the ce- phalic surface of the appendage; the distal caudal ridge is weaker, and relatively small- er. Ptychophallus micracanthus, new species Fig. 4A-D Material. —Panama: Pacific drainage, 1971, leg. L. G. Abele, 1 male holotype, cl. 24.6 mm, cb. 42.4 mm (USNM 240106). — Same data, 1 female paratype, 36 juveniles (USNM 240107).—Same data, 4 males, 2 females, 22 juveniles (USNM 240108).— San Blas, tributary to Rio Carti Grande at trail NW from Nuragandi, 5 Mar 1985, leg. R. W. Bouchard, 9 juveniles (USNM 240105).—San Blas, Quebrada Pingandi at Llando-Carti road, 4 Mar 1985, leg. R. W. Bouchard, 3 males (USNM 240110).—San Blas, Nuragandi off Llando-Carti road, 1 Mar 1985, leg. R. W. Bouchard, | juvenile male (USNM 240111).—Tributary to Pa- cora River, creek by road about 8 miles N of Cerro Azul, 27 Aug 1962, leg. Loftein and Kosan, 3 males, 1 female, 3 juveniles (USNM 240109). Diagnosis. — First gonopod with large lat- eral projection divided in 2 subequal round- 306 ed lobes by median notch, margins with minute setae; distal caudal ridge short, strong; mesial apical process small, trian- gular, with conspicuous spine on cephalic surface; apex strongly bent toward cephalic side, with field of spines directed toward latero-cephalic side. Description of holotype. —Cervical grooves recurved backward, narrow and shallow, not reaching margins of carapace; anterolateral margins with shallow postorbital notch, rest of borders entire. Postfrontal lobes wide, delimited anteriorly by transverse depres- sion; median groove narrow, deep, making incision on upper margin of front. Surface of carapace between postfrontal lobes and front flat, horizontal, only slightly inclined downward. Upper margin of front slightly convex in dorsal view, thin, well marked, with few small tubercles; lower margin thin, moderately sinuous; front between upper and lower margin high, vertical. Exognath of third maxilliped 0.54 length of ischium of endognath. Palm of largest cheliped moderately swollen, with lower and upper margins convex; fingers slightly gap- ing, with rows of small black-brown points on external surface. First gonopod with large lateral projection divided in 2 subequal rounded lobes by median notch, margins with minute setae; distal caudal ridge short, strong; mesial apical process small, trian- gular, with conspicuous spine on cephalic surface; apex strongly bent toward cephalic side, with field of spines directed toward latero-cephalic side. Remarks. —The first gonopod of P. mi- cracanthus resembles that of P. tumimanus (Rathbun, 1898) in the shape of the lateral lobe, but in P. micracanthus it is less ex- panded laterally and the middle notch is deeper. This species differs from all others in the genus by the presence of a small but clearly visible spine on the mesial projec- tion of the first gonopod. The species is well represented in the col- lections of the National Museum of Natural History, but the only fully mature male is PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON the holotype specimen (USNM 240106). However, immature crabs are easily iden- tifiable by means of the spine located on the mesial apical process of the gonopod which is present even in specimens with a carapace length of 10.8 and 12.5 mm. Etymology.—The specific name is from the Greek “‘mikros,”’ small, and “‘acanthus,”’ spine, in reference to the spine on the mesial process of the gonopod. Acknowledgments I thank Dr. Raymond B. Manning for the facilities provided during my stay at the Na- tional Museum of Natural History, Smith- sonian Institution. The manuscript was greatly improved by the comments provid- ed by two anonymous reviewers. Literature Cited Campos, M. R., & G. Rodriguez. 1984. New species of Freshwater crabs (Crustacea: Decapoda:Pseu- dothelphusidae) from Colombia.— Proceedings of the Biological Society of Washington 97:538— 543. Coifmann, I. 1939. Potamonidi della Guiana Inglese raccolti dal Prof. Nello Beccari.— Archivio Zoologico Italiano 27:93-116, pl. 3. Colosi, G. 1920. I Potamonidi dell R. Museo Zool- ogico di Torino.—Bolletino dei Musei di Zoo- loogia ed Anatomia Comparata della R. Univer- sita di Torino, 35(734):1-39. Ortmann, A. 1893. Die Dekapoden-Krebse des Strassburger Museums, mit besonderer Bertick- sichtigung der von Herr Doderlein bei Japan und bei den Liu-Kiu-Inseln gesammelten und zur Zeit in Strassburger Museum aufbewarten Formen. VII. Theil. Abtheilung: Brachyura (Brachyura genuina Boas) IJ. Unterabtheilung: Cancroidea, 2 Section: Cancrinea, 1. Gruppe: Cyclometopa.— Zoologische Jarbiicher, Abthei- lung fur Systematik, Geographie und Biologie der Thiere 7:41 1-495, pl. 17. 1897. Carcinologische Studien.—Zoolo- gische Jarbiicher, Abtheilung fur Systematik, Geographie und Biologie der Thiere 10:258-372, pl. 17. Prahl, H. von. 1988. Fresh-water crabs (Crustacea: Decapoda: Pseudothelphusidae) of the Pacific drainage of Colombia.—Zoologische Jahrbtch- er, Systematik, Okologie und Geographie der Tiere. 115:171-186. VOLUME 107, NUMBER 2 Pretzmann, G. 1965. Vorlaufiger Bericht uber die Familie Pseudothelphusidae.—Anzeiger der Mathematisch Naturwissenschaftliche Klasse der Osterreichischen Akademie der Wissenschaften (1)1:1-10. 1968. Neue Siidamerikanische Stisswasser- krabben der Gattung Pseudothelphusa. —Ento- mologisches Nachrichtenblatt, Wien 15(1):1-15. 1971. Fortschritte in der Klassifizierung der Pseudothelphusidae.—Sitzungsberichten der Osterreichischen Akademie der Wissenschaf- ten, Mathematisch Naturwissenschaftliche Klasse (1)179(1-4):15-24. 1972. Die Pseudothelphusidae (Crustacea Brachyura).— Zoologica 42(120) pt. 1:1-182. . 1978. Neue Potamocarcinini, Poglayen-Neu- wall leg. 1975 (Vorlaufige Mitteilung).—Sit- zungsberichten der Osterreichischen Akademie der Wissenschaften, Mathematisch-Naturwis- senschaftliche Klasse (1)1978(2):5 1-54. 1980. Von Dr. Ivo Poglayen-Neuwall 1975 in Mittelamerika gesammelte Krabben.—An- nalen des Naturhistorische Museum, Wien 83: 651-666. Rathbun, M. J. 1893. Descriptions of new species of American freshwater crabs. — Proceedings of the United States National Museum 16(959):649-— 661, pls. 73-77. 1898. A contribution to a knowledge of the 307 freshwater crabs of America. The Pseudothel- phusidae.— Proceedings of the United States National Museum 21(1158):507—537. 1905. Les crabes d’eau douce (Potamoni- dae).— Nouvelles Archives du Muséum d’His- toire Naturelle, Paris 7:159-321. Rodriguez, G. 1980. Descriptions préliminaire de quelques espéces et genres nouveaux de crabes d’eaux douce de l’Amérique Tropicale.—Bul- letin du Muséum national d’Histoire naturelle (Paris) (4) 2, Section A (3):885-894. 1982. Les crabes d’eau douce d’Amerique. Famille des Pseudothelphusidae.— Faune Trop- icale 22:1-223. Smalley, A. E. 1964. The river crabs of Costa Rica, and the subfamilies of the Pseudothelphusi- dae.— Tulane Studies in Zoology 12:5-13. Smith, S. I. 1870. Notes on American Crustacea. I. Ocypodoidea.—Transactions of the Connecti- cut Academy of Arts and Sciences 2:113-176, pls. 2-5. Young, C. G. 1900. The stalk-eyed Crustacea of the British Guiana, West Indies and Bermuda. Lon- don, 514 p., 4 pl. col. Centro de Ecologia, Instituto Venezolano de Investigaciones Cientificas, Apartado 21827, Caracas 1020-A, Venezuela. PROC. BIOL. SOC. WASH. 107(2), 1994, pp. 308-311 PARAPINNIXA CUBANA, A NEW PEA CRAB FROM CUBA (CRUSTACEA: BRACHYURA: PINNOTHERIDAE) Ernesto Campos Abstract.—One male specimen of Parapinnixa cubana, new species, was collected on Diego Pérez Reef, Cuba. Parapinnixa cubana most closely resem- bles P. magdalenensis Werding & Miller, 1990, and can be distinguished from this and all other species of the genus by the shape and proportions of the carapace and chela. Resumen.—Un especimen macho de Parapinnixa cubana, nueva especie, fue colectado en el Arrecife Diego Pérez, Cuba. Parapinnixa cubana se asemeja mayormente a P. magdalenensis Werding & Miiller, 1990, y se puede distinguir de ésta y todas las otras especies del género por la forma y proporciones del caparazon y la quela. During the study of the marine crustacean fauna of Cuba by J. C. Martinez-Iglesias and colleagues, Instituto de Oceanologia, Acad- emia de Ciencias de Cuba (ACC), one male pinnotherid specimen was collected on Die- go Peréz Reef. The specimen was sent to me for study and proved to be a new species of Parapinnixa. The holotype of the new spe- cies has been deposited in the Crustacean Collection, ACC. Abbreviations used in this paper include: third maxilliped as MXP3, carapace length as CL, carapace width as CW, and pereopods as P1 (the cheliped) to PS. Measurements are in millimeters. Parapinnixa cubana, new species Fie, Ll, 2 Material.—Diego Pérez Reef, Golfo de Batabano, Cuba, 20 m depth, Jul 1991, male holotype (CW = 3.0, CL = 1.8), coll. J. C. Martinez-Iglesias, ACC 1880. Description of holotype.—Carapace ellip- tical, width 1.6 times length, dorsally and laterally with short setae (Fig. 1A); front deflexed, triangular, with shallow medial groove, covered with minute setae; poste- rior margin almost straight. Eyes large, fill- ing orbits, extending far beyond anterior margin of carapace; orbital hiatus occupied by basal antennal article (Fig. 1B). Anten- nulae plicate in wide fossettes, fronto-or- bital distance subequal to carapace length. Buccal area triangular, epistome linear. MXP3 (Fig. 2A—B) with merus widely tri- angular, outer and distal margin almost straight, outer margin with plumose setae; carpus rounded, with long setae, and longer than combined length of elongated propo- dus and minute, inconspicuous dactylus; dactylus distally with 2 long tufts of setae (Fig. 2C); exopod ovate, without flagellum (Fig. 2D). P1 stout (Fig. 1C—D), as long as P2, merus dorsally subtriangular, with setae (Fig. 1C), lateroexternal surface trapezoidal and flat- tened (Fig. 1D); carpus rounded, setae shorter than those on merus. Chelae sym- metrical, tomentose; length of palm sub- equal to height, longer than fingers. External surface of palm somewhat convex proxi- mally, slightly concave distally, with fingers ornamented with several teeth; internal sur- face slightly swollen and with tuft of setae at summit, dorsally convex with tubercles, ventral margin somewhat sinuous. Fingers VOLUME 107, NUMBER 2 r,s BS 4 US " Sz, y uy EDGY LEG AN AeQeee ys hy Fig. 1. cheliped, inner and outer view respectively. Scales equal 0.5 mm. Plumose setae omitted on dactylus and ventral margin of carpus and propodus of P3 and P4; subacute teeth not visible on inner surface of dactylus in C. triangular, curving at tip where they cross; cutting surface of both fingers with row of subacute teeth. Dorsal surface of dactylus with crest of 7 subacute and acute teeth. Walking legs (Fig. 1A) decreasing in length and width from P2 to PS, meri flattened, dorsally and ventrally pubescent, PS much the shortest. Carpi dorsally subelliptical and convex, laterally subtriangular, with short setae. Propodi tapering distally. Fringe of extremely long plumose setae on outer sur- face of carpus, propodus and dactylus of P3 and P4, others placed on ventral margin of carpus and propodus of same legs. Dactyli triangular, naked at long corneus tip, those fH AN GD 309 Parapinnixa cubana, new species, male holotype, A, Dorsal view; B, frontal view; C and D, left of P2 to P4 somewhat falcate, that of P5 straighter. Abdomen (Fig. 2E) with 7 free somites, third widest and with convex lateral mar- gins, fourth through seventh gradually ta- pering, seventh longest, subtriangular, its length/width ratio 1.66. Gonopods (Fig. 2F) almost straight to- ward proximal third, folding mesially and tapering distally, ending in pore of sper- matic channel. Comparison with other species of Para- pinnixa. —Parapinnixa cubana mainly dif- fers from its congeners as follows: P. nitida (Lockington 1876), P. glasselli Garth, 1939, 310 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. Parapinnixa cubana, new species, male holotype. A—D, third maxilliped; A, outer view; B, palp, inner view; C, dactylus; D, exopod; E, abdomen; F, gonopod. Scales equal 0.125 mm (A-B, D, F), 0.05 mm (CQ), 0.5 mm (E). and P. hendersoni Rathbun, 1918 have a CW which is more than twice the CL, in- stead of 1.6 as in P. cubana. In addition, the three former species have two rows of hairs on the outer surface of the smooth palm, which contrast with the absence of such hairs and the presence of sub-acute and acute tubercles on the surface of the palm in P. cubana (see Garth 1939: plate 9, fig. 4; Williams 1984: fig. 358). Parapinnixa af- finis Holmes, 1900 has the dactylus of the chela hooked and a row of hairs on the sur- face of the palm which are lacking in P. cubana (see Glassell 1933: fig. 1-2). Para- pinnixa bouvieri Rathbun, 1918, and sim- ilarly P. affinis, have small eyes (see Wil- liams 1984: fig. 357) and the fronto-orbital width about one third the CW, whereas P. cubana has large eyes and the fronto-orbital width larger than one third the CW. In ad- VOLUME 107, NUMBER 2 dition P3 and P4 are smaller in P. bouvieri than in P. cubana. Parapinnixa beauforten- sis Rathbun, 1918 has a tuft of hair on either side of the dorsal surface of the carapace near the lateral margin, which are absent in P. cubana. A male specimen identified with hesitation as P. beaufortensis by Werding & Muller (1990: fig. la and le) has, in addition to the tufts noted above, a less elongated abdomen and more robust gonopod than does P. cubana. Parapinnixa cubana most closely resem- bles P. magdalenensis Werding & Miller (1990). The two species are easily distin- guished by the articles of the palp on MXP, and the pattern of setation on the carapace, chelipeds and walking legs. Furthermore the antero-lateral margin of the carapace is crenulate in P. magdalenensis but not in P. cubana; the merus of the cheliped is more elongated and possesses tubercles in P. magdalenensis than in P. cubana. Finally, the posterior margin of the caparace is wider than the fronto-orbital margin, and the gon- opod is more robust in P. magdalenensis (see Werding and Muller 1990: fig. 2a—e) than in P. cubana. Acknowledgments I am indebted to Juan Carlos Martinez- Iglesias for allowing me to study and de- scribe Parapinnixa cubana. I gratefully ac- knowledge the help of my wife Alma Rosa who illustrated and prepared the figures for publication. My deep gratitude is due to Raymond B. Manning and Rafael Lemaitre (National Museum of Natural History, Smithsonian Institution), and Deborah Zmarzly (Marine Biology Laboratory, San 311 Diego) for reviewing my manuscript with great care. This work has been supported by project ““Crustaceos simbiontes de Baja California” of the Facultad de Ciencias, Universidad Autonoma de Baja California, and the 1989 Scientific Collaboration Agreement between México (Consejo Na- cional de Ciencia y Tecnologia) and Cuba (ACC). Literature Cited Garth, J. S. 1939. New brachyuran crabs from the Galapagos Islands.— Allan Hancock Pacific Ex- pedition 5(2):9-29. Glassell, S. A. 1933. Notes on Parapinnixa affinis Holmes and its allies. — Transactions of the San Diego Society of Natural History 7(27):319-330. Holmes, S. J. 1900. Synopsis of California stalk-eyed Crustacea. — Occasional papers of the California Academy of Science 7:260 pp. Lockington, W. N. 1876. Remarks on the Crustacea of the West coast of North America, with a cat- alogue of the species in the museum of the Cal- ifornia Academy of Science.— Proceedings of the California Academy of Science 7:145-156. Rathbun, M. J. 1918. The grapsoid crabs of Amer- ica.— United States National Museum Bulletin 97:1461. Werding, B., & H. G. Miller. 1990. Die Gattung Parapinnixa Holmes 1894 an der Nordkiste Kolumbiens, mit Beschreibung von Parapin- nixa magdalenensis n. sp. (Crustacea: Decapo- da: Pinnotheridae).—Senckenbergiana Biologi- ca 70(1/3):221—227. Williams A. B. 1984. Shrimps, lobsters and crabs of the Atlantic coast of the eastern United States, Maine to Florida. Smithsonian Institution Press, Washington, xvii + 550. Facultad de Ciencias, Universidad Au- tonoma de Baja California, Apartado Postal 2300, Ensenada, Baja California, Mexico. PROC. BIOL. SOC. WASH. 107(2), 1994, pp. 312-317 PETROLISTHES EXTREMUS, A NEW PORCELAIN CRAB (DECAPODA: ANOMURA: PORCELLANIDAE) FROM THE INDO-WEST PACIFIC Roy K. Kropp and Janet Haig Abstract. — Petrolisthes extremus, new species, is described from several lo- cations in the Indo-west Pacific. The new species is most similar to P. coccineus and P. carinipes, with which it shares a transversely rugose carapace, sinuously triangular front, and one epibranchial spine. Petrolisthes extremus is distin- guished from the P. coccineus by having the lateral margins of the carapace not converging strongly between its widest point and the epibranchial spines and by having the carpus of the cheliped relatively shorter and wider. Petrolis- thes extremus is distinguished from P. carinipes by having a narrower front of the carapace that has sharply oblique lateral lobes. The new species is known from Cocos Keeling, the Kermadec Islands, Lord Howe Island, Easter Island, the Mariana Islands, and Taiwan. Among several collections of porcellanid crabs from the Indo-west Pacific were spec- imens belonging to an undescribed species of Petrolisthes. Study of the material re- vealed that several previous literature rec- ords also could be referred to the new spe- cies. The material used in this study is located in the National Museum of Natural History, Smithsonian Institution, Washington, D.C. (USNM), the Natural History Museum of Los Angeles County (the collections for- merly were associated with the Allan Han- cock Foundation), Los Angeles, California [LACM (AHF)], the Bernice P. Bishop Mu- seum, Honolulu, Hawaii (BPBM), the Aus- tralian Museum, Sydney, Australia (AM), the National Museum of New Zealand, Wellington, New Zealand (NMNZ), and the Institute of Zoology, Academia Sinica, Tai- pei, Taiwan (IZAS). Abbreviations used are: cl, carapace length; cw, carapace width; coll., collector; ft, feet; m, meters; mm, millimeters; and Ovig., Ovigerous. Petrolisthes extremus, new species Bigesl Petrolisthes, sp.—Whitelegge, 1889:36 [list, Lord Howe Island]. Petrolisthes lamarcki. —Gillett & McNeill, 1959:158, pl. 152, figure at bottom of page [Lord Howe Island]. [Not Petrolisthes la- marckii (Leach, 1820)]. Petrolisthes coccineus. —Gillett & McNeill, 1962:158, pl. 152, figure at bottom of page; Gillett and McNeill, 1967:158, pl. 152, figure at bottom of page [Lord Howe Island]. [Not Petrolisthes coccineus (Owen, 1839)]. Petrolisthes n. sp. 2 [Haig, ms].—Kropp et al., 1981:39 [Guam]; Kropp & Eldredge, 1982:125 [Cabras Island, Guam]. Type material.—Holotype: Mariana Is- lands: Guam; Asan Point; shore at low tide; 13 Jun 1980; coll. R. K. Kropp; 1 ¢é LACM 80-151.1 (AHF 8014). Paratypes: Cocos Keeling Island: West Island (ocean side); cove on N end (12°8'22"S, 96°49'0”E); dead coral; <1 m; 22 Feb 1974; coll. Smith- VOLUME 107, NUMBER 2 Vaniz etal., 1 2(ovig.) USNM 190765. Ker- madec Islands: Sunday Island; 1909-1910; coll. R. S. Bell & W. R. B. Oliver; 9 4, 7 2 (3 ovig.) NMNZ. Taiwan: Hsan Tiao Chiao; 10 May 1969; coll. Ting; 1 ¢IZAS. Yeh Liu Pi; poison station; 2 m; 28 Jun 1978; coll. L. G. Eldredge; 2 6, 1 2 (ovig.) LACM 78- 237.1 (AHF); Kuei-An; 47 km S of Hualien; subtidal under rock; 2 Jul 1978; coll. L. G. Eldredge; 1 2 (ovig.) USNM 210587. Mar- iana Islands: Guam; Asan Point; shore at low tide; 13 Jun 1980; coll. R. K. Kropp; 1 2 (ovig.) LACM 80-151.2 (AHF); Piti Bay; outer reef flat W of Camel Rock; down 1-— 2 m in consolidated rubble; 11 Jun 1986; coll. B. D. Smith & H. Conley; 2 2 (ovig.) BPBM. Easter Island: from starfish stom- ach; Jan 1965; coll.t. Efford & J. Mathias; 1 6 LACM 65-325.2 (AHF 654); Anakena; 20-25 ft [7-8 m]; 15 Jan 1965; coll. I. Efford & J. Mathias; 1 6 (damaged) LACM 65- 325.1 (AHF); Motu Iti; rock; 8 Aug 1972; coll. H. I. Moyano; | 2 LACM 72-358.1 (AHF). Other material examined. —Taiwan: Kuei-An; 47 km S of Hualien; on Pocillo- pora danae Verrill and Stylophora pistillata (Esper); 1-3 m; 23 Jul 1979; coll. R. K. Kropp; 3 2, 2 6 USNM 210585, 210586; San Hsien Tai; on Pocillopora danae; 3 m; 25 Jul 1979; coll. R. K. Kropp; specimen parasitized by rhizocephalan, USNM 210584. Mariana Islands: Anatahan; ““Ob- servation Spot’’; intertidal under rocks; 19 Jul 1981; coll. L. G. Eldredge; 3 2 USNM 210588; subtidal under rock; 5—7 m; 19 Jul 1981; 1 2 USNM. Pagan; Katsu; subtidal under rock; 2 m; 7 Mar 1981; 2 2 USNM; “Palapala Bay’; under rock; 1 m; 15 Jul 1981; 1 2 USNM. Guam; Piti Bay; outer reef flat; intertidal; 13, 15 Jun 1980; 3 Jun 1981; 3 6, 2 2 (1 ovig.) USNM; outer reef flat; down 1-2 m in consolidated coral rub- ble; 18 May, 11 Jun 1986; coll. B. D. Smith & H. Conley; 2 6, 2 2 (ovig.) USNM; Lu- minao; reef front under rock; 6 m; 8 Sep 1980; coll. V. Tyndzik; 1 6 USNM. Pago Bay; outer reef flat; intertidal; 3 May 1980; 313 1 juv.; reef front under rock; 2—3 m; 24, 31 May 1986; 1 6, 1 2 (ovig.) USNM. Lord Howe Island: collected before 1900; 1 2 AM G.2420, 2 6, 2 2 AM G.2512; collected be- fore 1909; coll. A. R. McCulloch; 2 2 AM P.1131; 1921; coll. A. R. McCulloch; 2 ¢ AM P.5248, 1 6,5 2 AM P. 5429: collected before 1924; coll. G. P. Whitely; 4 6 AM P.6883; Ned’s Beach; Jul 1959; coll. E. Pope; 1 2 AM P.15168; Ned’s Beach; from coral; Oct 1962; coll. J. Booth; 1 é AM P.15169; 6 Mar 1963; coll. J. Booth; 1 6, 2 2 AM P.15170. Measurements. — Holotype: cl, 6.6 mm, cw, 6.5 mm. Paratypes: largest male: cl, 11.3 mm, cw, 10.5 mm; largest female: cl. 9.8 mm, cw, 9.7 mm; smallest ovigerous fe- male: cl, 5.0 mm, cw, 4.6 mm. Diagnosis. —Carapace with distinct transverse rugae each with anterior row of setae, 1 epibranchial spine present; front sinuously triangular, margin serrated, su- praocular spine present. Chelipeds slightly unequal in size; carpus with squamate me- dian longitudinal ridge, anterior margin with 3-4 spine-tipped teeth, posterior margin with 3 spines; dorsal surface of manus di- vided by longitudinal squamous ridge, outer portion with granules, outer margin serrated and with 6 or more spines on proximal half; inner edges of fingers serrated, gape with very short setae, projecting only slightly above surfaces of fingers. Merus of walking legs with transverse striations, anterior mar- gin with 3-8 spines and fringed with plu- mose setae, posterodistal spines 2, 2, 0; car- pus of first walking leg with or without anterodistal spine. Description. —Carapace slightly longer than broad, broadest at posterior branchial region, lateral margins strongly converging anteriorly, cristate. Front sinuously trian- gular, lateral lobe oblique, less produced than median lobe; margin crenulate; protogastric lobes fringed anteriorly with plumose setae, divided by median groove. Supraocular lobe well-developed, spine usually present, oc- casionally obsolescent. Orbits shallow, very 314 oblique; outer orbital angle not produced, or produced into very small tooth. Epi- branchial spine well-developed. Gastric re- gion with strong transverse striae, some- times interrupted at midline; hepatic, anterior branchial, outer margin of poste- rior branchial with shorter striae; frontal re- gion crossed with fine transverse lines; car- diac, posterior branchials except near margins finely punctate, appearing smooth. Striae of carapace lined anteriorly with very fine, short setae, extending less than half way between crests of adjacent striae. Basal segments of antennules with several irregularly-spaced denticles of varying size on anterior margin. First movable segment of antennae with strong, spinnule-tipped lobe on anterior margin; second rugose ‘or strongly granulate along anterior margin, tubercle sometimes developed anteroprox- imally; third smooth. Dorsal extension of ocular peduncle onto cornea triangular, lined with simple setae, single larger seta distally (often missing in preserved specimens); cornea round in lat- eral view. Merus of chelipeds transversely rugose on dorsal and ventral surfaces; inner margin with strong rugose tooth or lobe, its edges crenulate; distal margin with one or two spines, lined distally with plumose setae; median or subproximal spine usually pres- ent on outer portion of dorsal surface. Car- pus (excluding inner marginal teeth) about twice as long as wide; inner margin with 3— 4 strong teeth, these serrate or crenulate along edges, often with smaller denticles be- tween them; series of 5—6 spines along distal half of outer margin, including 1 at outer distal angle; dorsal surface with median lon- gitudinal row of broad, flat squamae, inner portion (including marginal teeth) covered with smaller flattened granules; ventral sur- face with transverse rugae. Palm with outer margin thin, strongly curved, serrated and proximally spinulate in smaller specimens, crenulate or with blunt teeth in larger ones; dorsal surface with strong longitudinal crest PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON extending from base of dactyl nearly to ar- ticulation with carpus; surface inside this crest with oblique rugae or flattened squa- mae; surface to outside covered with flat- tened or somewhat upstanding granules; ventral surface obliquely rugose. Dactyl with longitudinal row of prominent, flattened squamae; surface of fixed finger slightly con- cave, usually granulate; cutting edges meet- ing for entire length; in some individuals of both sexes, fingers of 1 cheliped gaping and cutting edge of dactyl with strong conical tooth proximally; ventral surfaces of fingers with flattened granules, inner side with short, sparse pubescence or none. All segments of cheliped with very short, fine setae arising from distal side of granules and rugae, but these scarcely visible except at margins; in smaller specimens a fringe along outer mar- gin of chela; in larger adults this fringe ab- sent or confined to proximal portion of mar- gin; usually fringe of plumose setae along distal side of each inner carpal tooth. Walking legs rugose dorsally; dorsal sur- face of all segments with simple setae of varying length; merus with thick fringe of plumose setae along anterior margin. Leg 1: merus with 6—9 spines on anterior margin, 2 well-developed posterodistal spines (the smaller, more anterior spine occasionally obsolescent); carpus with anterodistal spine; propodus with 2—3 movable spinules along midline of posterior margin. Leg 2: merus with 4-8 spines on anterior margin, 2 well- developed posterodistal spines (the smaller, more anterior spine occasionally obsoles- cent); carpus without anterodistal spine; propodus with 2—3 movable spinules along midline of posterior margin. Leg 3: merus with 3—5 spines on anterior margin, pos- terodistal margin unarmed; carpus without anterodistal spine; propodus with 2 mov- able spinules along midline of posterior margin. Dactyls of all legs with 3 movable spinules on posterior margin. Coloration. —Overall color of carapace, chelipeds, and walking legs mottled light green, blue, and bright white with scattered VOLUME 107, NUMBER 2 315 Fig. 1. burgundy spots; red setae line rugae of car- apace. Propodus and dactylus of walking legs with transverse burgundy and white bands. Remarks.—In the Indo-west Pacific, P. extremus is most similar to P. coccineus (Owen) and P. carinipes (Heller), with which it shares a transversely rugose carapace, sin- Petrolisthes extremus, new species, paratype 6 (Kermadec, NMNZ): a, b, chelipeds (dorsal view); c, carapace; d, left cheliped (ventral view); e-g, right walking legs 1, 2, 3; h, right eye (dorsal, lateral views); i, basal segment of right antennule. Scale: 2.2 mm (a, b, d); 1.5 mm (¢, e-g); 1 mm (h-1). uously triangular frontal region, a supra- ocular spine, and a single epibranchial spine. The ranges of the three species overlap in the western Pacific. Petrolisthes extremus ranges from Taiwan and the Mariana Is- lands south to Lord Howe Island and east to Easter Island. Petrolisthes carinipes oc- curs from the Red Sea and western Indian 316 Ocean to the Ryukyu, Ogasawara, Mariana, and Chesterfield Islands in the western Pa- cific (Haig 1983, 1987). Petrolisthes cocci- neus ranges throughout the Indian Ocean, the western Pacific from Indonesia to Japan, and extends eastward from the Ogasawara and Mariana Islands to the Hawaiian Is- lands and the Tuamotu Archipelago (Haig 1983). The coloration of the three species differs sharply. In contrast to the mottled light green, blue, and bright white with scattered burgundy spots marking the carapace and chelipeds of P. extremus, those of P. cari- nipes are dark red-brown, appearing black to the unaided eye (RKK, personal obser- vation of material from Guam). The cara- pace of P. carinipes also is marked with white spots at the tip of the rostrum, at the su- praocular spines, and along the lateral and medial regions. White spots also occur on the chelipeds of P. carinipes. In P. coccineus, the carapace is pale blue-green with the gas- tric region marked with pale yellow-orange and a dark yellow-orange gastric ridge (RKK, personal observation of material from Guam). The manus of P. coccineus is blue- green and marked with a distinctive orange longitudinal crest and yellow-orange along the outer margin. Petrolisthes carinipes and P. extremus are very similar in the striation pattern of the carapace and in the form and armature of the chelipeds and walking legs. However, in P. carinipes the lateral carapace margins are evenly convex between the epibranchial and posterolateral angles, the lateral lobes of the front are nearly transverse, and the orbits are regularly concave between the supra- ocular spine and the outer orbital angle. In P. extremus the carapace is strongly diver- gent posteriorly, the lateral lobes of the front are sharply oblique, and the orbits oblique between the supraocular spine and the outer orbital angle. Petrolisthes extremus may be most easily distinguished from P. coccineus by the rel- ative proportions of the carpus of the che- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON liped and the spination on the posterior margin of the merus of the third walking leg. The carpus of the cheliped is about twice as long as wide in P. extremus, but much more than twice as long as wide in P. coc- cineus. The posterodistal margin of the third walking leg is unarmed in P. extremus, but armed with one or two spines in P. cocci- neus. Also, in P. coccineus the front is nar- rowly triangular with the lateral lobes even more oblique and obscure than in P. extre- mus; the strong transverse striae on the gas- tric region are interrupted in the midline by a short but distinct interspace; and the num- ber of spines on the anterior margin of the merus of the walking legs averages a little higher (5 to 9, 7 to 9, and 5 to 8 on legs 1, 2, and 3, respectively). Etymology. —From the Latin, extremus, meaning outermost or farthest away. When study of this taxon began by one of us (JH), the only material available had been col- lected from Lord Howe, Kermadec, and Easter Islands, the “‘outposts”’ of the trop- ical or subtropical Indo-west Pacific. Al- though later collections from Taiwan and the Marianas counter the idea that the spe- cies 1s an “‘outpost taxon,” the initial name was retained, at least in part, to refer to the interesting distribution of the taxon in the southern hemisphere portion of its range. Habitat. —In the Mariana Islands, found under rocks, generally subtidal to a depth of 7 m; occasionally in the intertidal zone. In Taiwan, also on base of pocilloporid cor- als. Distribution. —FPetrolisthes extremus ranges from Taiwan and the Mariana Is- lands in the western Pacific south to Lord Howe Island and eastward to the Kermadec Islands and Easter Island. Acknowledgments We thank I. E. Efford for donating ma- terial to AHF from the Medical Expedition to Easter Island (METEI), 1964-1965, and J. C. Yaldwyn and the late F. A. McNeill VOLUME 107, NUMBER 2 for the loan of specimens to AHF. RKK thanks the University of Guam Marine Laboratory, the University of Maryland, and the USNM for support during field work in the Marianas. Funding for field work has been supported in part by a grant from the Biological Oceanography Section of the Na- tional Science Foundation to G. J. Vermeij. Field work in Taiwan was part of a joint NSF/Republic of China-sponsored study conducted by the University of Guam. This is University of Guam Marine Laboratory Contribution Number 349. Literature Cited Gillett, K., & F. McNeill. 1959. The Great Barrier Reef and adjacent isles. Coral Press Pty. Ltd., Sydney, xiii, 194 pp. ——.,, & 1962. The Great Barrier Reef and adjacent isles. Revised edition. Coral Press Pty. Ltd., Sydney, xiii, 209 pp. ——, & . 1967. The Great Barrier Reef and adjacent isles. 2nd revised edition. Coral Press Pty. Ltd., Sydney, xiii, 194 pp. Haig, J. 1983. Porcellanidae (Decapoda, Anomura) from the Seychelles, western Indian Ocean.— Crustaceana 45:279-289. . 1987. Porcellanid crabs from the Coral Sea. — The Beagle, Records of the Northern Territory Museum of Arts and Sciences 4:1 1-14. Kropp, R. K., & L. G. Eldredge. 1982. Macroinver- tebrates. Pp. 107-131 in R. H. Randall & L. G. 317 Eldredge, eds., Assessment of the shoalwater en- vironments in the vicinity of the proposed OTEC development at Cabras Island, Guam.—Uni- versity of Guam Marine Laboratory Technical Report 79:1-208. ——, D. S. Wooster, & L. G. Eldredge. 1981. Pre- liminary checklist of anomuran crustaceans from Guam. Pp. 39-41 in A working list of marine organisms from Guam.—University of Guam Marine Laboratory Technical Report 70:1-88. Leach, W. E. 1820. Galatéadées.— Dictionnaire des Sciences Naturelles 18:49-56. Owen, R. 1839. Pp. 77-92 in The zoology of Captain Beechey’s voyage; compiled from the collections and notes made by Captain Beechey, the officers and naturalist of the expedition, during a voyage to the Pacific and Behring’s Straits performed in His Majesty’s ship Blossom, under the com- mand of Captain F. W. Beechey, R.N., F.R.S., in the years 1825, 26, 27, and 28. Whitelegge, T. 1889. Pp. 35-36 in R. Etheridge junr., The general zoology of Lord Howe Island; con- taining also an account of the collections made by the Australian Museum collecting party, Aug.— Sept. 1887.—Memoirs of the Australian Mu- seum 2:1-42. Addresses: (RKK) Battelle Ocean Sci- ences, 397 Washington Street, Duxbury, Massachusetts 02332, U.S.A.; (JH) Allan Hancock Foundation, University of Southern California, University Park, Los Angeles, California 90089-0371, U.S.A. PROC. BIOL. SOC. WASH. 107(2), 1994, pp. 318-324 A NEW FRESHWATER CRAB OF THE GENUS GEOTHELPHUSA (CRUSTACEA: DECAPODA: BRACHYURA: POTAMIDAE) FROM KAGOSHIMA PREFECTURE, SOUTHERN KYUSHU, JAPAN Hiroshi Suzuki and Esji Tsuda Abstract.—A new freshwater crab, Geothelphusa exigua, is described from Kagoshima Prefecture, southern Kyushu, Japan. It is differentiated from G. dehaani (White 1847) and G. candidiensis Bott, 1967, in possessing distally narrowed eyes, presence of granules both on the lower edge of the epistome and on the groove between the subhepatic and pterygostomian regions, and laterally curved male first gonopod with a subterminal genital opening. This is the second species known from the Japanese mainland. Among the eight species of the genus Geo- thelphusa known from Japan, G. dehaani (White 1847) is the only species known to occur on the Japanese mainland (north of Honshu southward to Nakano-shima of the Tokara Islands, south of Kyushu) (de Haan 1835; White 1847; Stimpson 1858; Rath- bun 1898, 1904, 1905; Miyake & Chiu 1965; Miyake & Minei 1965; Bott 1967, 1970; Minei 1973, 1974b). The other seven spe- cies, G. obtusipes Stimpson, 1858, G. saka- motoana (Rathbun, 1905), G. aramotoi Mi- nei, 1973, G. tenuimana (Miyake & Minei 1965), G. levicervix (Rathbun 1898), G. can- didiensis Bott, 1970, and G. miyazakii (Mi- yake & Chiu 1965), are restricted to the Ryukyu Islands, including Amamiohshima. During our current study of the geograph- ical distribution of G. dehaani in southern Kyushu, unusual specimens of a freshwater crab of the genus Geothelphusa were found on the Ohsumi Peninsula of Kagoshima Prefecture. The unusual eyes and male first gonopods of these crabs indicate that they represent a new species which is herein de- scribed and illustrated. The holotype is deposited in the National Science Museum, Tokyo (NSMT), and the paratypes are in the Marine Biological Lab- oratory, Faculty of Fisheries, Kagoshima University, Kagoshima (KUMB) and the Kitakyushu Museum of Natural History, Kitakyushu (KMNH). Measurements shown in parentheses under “‘Material examined” indicate the maximum carapace width in millimeters. Abbreviations used include: Cr and cr, crustacea; IvR, Invertebrate Recent. Family Potamidae Ortmann, 1896 Genus Geothelphusa Stimpson, 1858 Geothelphusa exigua, new species Figs. 1-3 Material examined. —Kimotsuki River: Futamata-gawa, 400 m alt., 11 Jul 1991; 7 6 (13.1-21.8), 3 2 (12.2—13.0), holotype, ¢ (21.0), NSMT-Cr 11314, KUMBcer 1039, KUMBcr 1041: Iwaya-gawa, 220 m alt., 25 Sep 1991; 1 6(20.5), KUMBer 1042: Nana- tsudani, Kushira-gawa, 400 m alt., 11 Jul 1991; 1 6 (18.8), 2 2 (13.2, 13.4), KUMBer 1041: Takakuma valley, Kushira-gawa, 180 m alt., 22 Oct 1991; 1 6 (16.8), KUMBcr 1045: Naganomaki, Aira-gawa, 180 m alt., 11 Jul 1991; 6 3(13.3-22.6), 42 (15.9-24.4), KMNH-IvR 900001-900004, KUMBcr 1041. Honjo River: Sarugajo valley, 150 m alt., 22 Oct 1991; 16(19.6), KUMBcr 1045. Ohsumi Kamino River: Hangaishi, 300 m alt., 25 Sep 1991; 1 6(20.8), KUMBcr 1042. VOLUME 107, NUMBER 2 O River: Onigauto, Fumoto-gawa, 300 m alt., 9 Aug 1989; 3 6 (11.2-14.8), 2 2 (16.2, 17.8), KUMBcr 1038, KUMBcr 1040: Uchinomaki, Shibatate-gawa, 380 m alt., 9 Aug 1989; 2 6 (12.6, 14.3), 4 29 (13.0-28.7), KUMBcr 1040: Shinden, 330 m alt., 25 Sep 1991; 1 6 (18.7), KUMBcr 1042: Ohfuji- gawa, 520 m alt., 9 Aug 1989; 2 2 (12.7, 26.7), KUMBcr 1040: Ohtakeno, Akase- gawa, 600 m alt., 9 Aug 1989; 1 young (6.9), 2 6 (8.7, 17.8), 3 2 (13.9-15.6), KUMBcr 1037, KUMBcr 1040. Hirose River: Ma- gome, 450 m alt., 2 Oct 1991; 1 2 (24.7), KUMBcr 1043. Kubota River: Himekado, 160 malt., 25 Sep 1991; 1 6(20.4), KUMBcr 1042. Funama River: Gorogamoto, 380 m alt., 2 Oct 1991; 16(9.9), 129(19.0), KUMBcr 1043. Hitotsutani River: Uchinoura, 390 m alt., 2 Oct 1991; 1 6 (16.7), KUMBcr 1043. Ohura River: Uchinoura, 300 m alt., 8 Oct 1991; 1 6 (14.4), KUMBcr 1044. Hetsuka River: Sarutubo, 490 m alt., 8 Oct 1991; 1 6 (25.0), 1 2 (27.5), KUMBcr 1044: Doh- gabaru, 180 m alt., 8 Oct 1991; 1 6 (16.4), KUMBcr 1044. All specimens were collected by H. Su- zuki and E. Tsuda. Diagnosis.—Male first gonopod saber- like, penultimate segment slightly curved laterally, ultimate segment strongly curved laterally, tapering, genital opening subter- minal. Ocular peduncle swollen proximally, cornea small. Description. —Carapace much broader than long, smooth and devoid of hair (Fig. la), postfrontal and postorbital regions in- distinctly rugose, faint oblique striae on epibranchial and posterolateral regions, epi- and uro-gastric regions distinct, former di- vided into two parts by median depression, cervical groove obsolete on epibranchial re- gion. Anterolateral margin of carapace cris- tate, lined with fine rounded granules, epi- branchial notch rudimentary. Frontal margin 0.35 (0.33-0.40 in male, 0.31-0.38 in female) times as broad as carapace. Pos- terior margin of epistome divided into three parts by 2 deeper notches (Fig. 1b), granules 319 present on lower edge of epistome, absent medially. Lower orbital margin and groove between subhepatic and pterygostomian regions lined with small rounded granules. Eyestalk short, proximallly swollen, dis- tally slim (Fig. la, b). Cornea small, slightly wider than distal portion of ocular pedun- cle Merus of third maxilliped broad, square, with deep punctum (Fig. lc). Three-seg- mented palp connected on inner distal angle of merus, tip of palp not below distal margin of ischium. Exopod slender, longer than is- chium, with small flagellum (Fig. Ic, d). Chelipeds asymmetrical in males over 13.0 mm carapace width (right larger than left in 26 out of 28 males, left larger than right in remainder), symmetrical in all fe- males and in males less than 13.0 mm; large chela 2.09 (1.82—2.27) times as long as high (Fig. le), palm smooth and surfaces rela- tively rounded, fingers with 2 longitudinal ridges on outer lateral surface (Fig. le, f). Carpus of large cheliped slightly smooth, with stout inner tooth below which is a low swelling (Fig. 2a). Carpus of small cheliped without any swelling below stout inner tooth. Palp of mandible 3-segmented (Fig. 2b), distal segment uniramous and sickle-shaped, median segment longer than wide, distal half expanded, proximal segment short, stout. Adult male first gonopod saber-like (Fig. 2c-h), penultimate segment slightly curved laterally, synovial membrane short, about 3 times as long as broad (Fig. 2c), ultimate segment strongly curved laterally (Fig. 2c, d), tapering, with genital opening subter- minal in position (Fig. 2e). Tip of first gon- opod papilla-like in specimen 8.7 mm car- apace width (Fig. 2f), and tapered in specimens 11.2 and 14.6 mm carapace width (Fig. 2g, h). Male second gonopod slender, flat, weakly convex, with small lamella on distal one-third (Fig. 21, j). Sizes (carapace width). —Males, 8.7—25.0 mm; females, 12.2—28.7 mm. Color in life. —Chocolate brown or dark brown with scattered black speckles on car- 320 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON e qe b,c, d, e&, f; 10mm Fig. 1. Geothelphusa exigua, new species, male holotype (NSMT-Cr 11314): a, dorsal view; b, frontal view; c, third maxilliped, frontal view; d, exopod of third maxilliped, frontal view; e, right cheliped, lateral view; f, fingers, ventrolateral view. VOLUME 107, NUMBER 2 321 Fig. 2. Geothelphusa exigua, new species, male holotype (NSMT-Cr 11314): a, carpus of right cheliped, dorsal view; b, right mandibular palp, ventral view; c, left first gonopod, dorsal view; d, same, ventral view; e, distal part of same, distal view; i, left second gonopod, dorsal view; j, same, ventral view. Male paratypes (KUMBer 1037-1039): f, left first gonopod (8.7 mm carapace width), ventral view; g, same (11.2 mm carapace width), ventral view; h, same (14.6 mm carapace width), ventral view. 322 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON O ie) 3I°40N iG) é O O O 8 Tarumizu ; City ; Baas aad Takakuma Mt. * 4, Shibushi Bay Kagoshima OO —~ 31°20'N o Kunimi Mt. oa Q ; 2 3I°10'N an) Inao,-Peak I30°45'E I31°O0'E 31°00'N Fig. 3. Map showing the distribution and abundance of G. exigua, new species (black part) and G. dehaani (white part) in Ohsumi Peninsula, Kagoshima. Arabic numerals corresponding to size of circles in the square at bottom right indicate the numbers of crabs captured per ten minutes by one person. Broken line shows 150 m contour line. VOLUME 107, NUMBER 2 apace and pereopods. Palm of chelae choc- olate brown, fingers milk white. No color variation between sexes and sizes. Etymology.—The specific name is de- rived from the Latin “‘exiguus”’ (small), al- luding to the small cornea, characteristic of the new species. Remarks. — The saber-like male first gon- opod and the 3-segmented mandibular palp with an uniramous distal segment displayed by the new species are characteristics of the genus Geothelphusa (see Bott 1970). The medium-sized carapace, smooth palm, and broad frontal region ally the species with G. dehaani and G. candidiensis, from which it is distinguished by several features. The most definitive differences may be seen on the male first gonopods. The ulti- mate segment in G. dehaani and G. candi- diensis is straight or slightly curved mesially (Bott 1967, 1970; Minei 1973, 1974a), end- ing in a papilla-like tip with a terminal gen- ital opening. In the new species, however, this segment is strongly curved laterally and tapering, having a subterminal genital open- ing. The eyestalks in both related species are constricted at the middle, and the cornea and the proximal part of the ocular peduncle are swollen. In the new species, only the proximal part of the ocular peduncle is swollen, the cornea and the distal part of the peduncle are proportionately narrower. The postorbital and epibranchial regions of the carapace bear finely crenulate striae in G. candidiensis (see Minei 1973), instead of faint striae as in G. exigua. Examination of specimens of G. candidiensis reported by Minei (1973), and now in the collection of the Kitakyushu Museum of Natural His- tory, Kitakyushu (5 6, 4 °, ZLKUm 1016, Maezato, Ishigaki-jima, 28 Oct 1962, leg. S. Kudaka; 19 6, 18 ¢, ZLKUm 1019, Pen- san-gara, Ishigaki-jima, 19 May 1963, leg. S. Kudaka), shows that there are distinct granules on the median part of the lower edge of the epistome which are barely dis- cernible in the new species. The presence of distinct granules on the groove between the 323 subhepatic and pterygostomian regions also differentiates G. exigua from G. dehaani. The carapace and pereopods of G. dehaani show color variation, for example red, or- ange, blue, brown, purple, or yellowish white (Chokki 1976, 1980; Suzuki & Tsuda 1991). The palm and fingers of G. dehaani, how- ever, are usually yellowish white, regardless of carapace color. In G. exigua, the carapace and pereopods are chocolate brown or dark brown, and only the fingers are milk white. In addition, allelic substitution was ob- served at General protein-1 and -2, Lactate dehydrogenase, and Isocitrate dehydro- genase-2 loci between G. exigua and G. de- haani (Suzuki & Tsuda, pers. comm.). Distribution.—The specimens of G. exi- gua examined have been obtained only in the area above 150 m altitude on Takakuma Mountain, Inao Peak, and Kunimi Moun- tain Ranges, in Ohsumi Peninsula, Kago- shima Prefecture (Fig. 3), where the Mio- cene granitic rock and quartz porphyry are exposed. Geothelphusa exigua and G. de- haani are sympatric, having been taken to- gether at many locations. Acknowledgments We thank K. Baba of Kumamoto Uni- versity, C. L. McLay of the University of Canterbury, and M. Tirkay of the Natur- museum und Forschungsinstitut Sencken- berg, for their critical readings of the manu- script. Thanks are also extended to P. K. L. Ng and R. Lemaitre for their valuable com- ments on the manuscript. We are also in- debted to M. Takeda of National Science Museum, for his comments and to M. Ota and Y. Yabumoto of the Kitakyushu Mu- seum of Natural History, for allowing us to examine the specimens under their care and the use of laboratory facilities. Literature Cited Bott, R. 1967. Potamiden aus Ost-Asien (Parapot- amon De Man, Sinopotamon n. gen., Candidi- 324 opotamon n. gen., Geothelphusa Stimpson) (Crustacea, Decapoda).—Senckenbergiana bio- logica 48(3):203-220. 1970. Die SiBwasserkrabben von Europa, Asien, Australien und ihre Stammesgeschichte. Eine Revision der Potamoidea und der Para- thelphusoidea. (Crustacea, Decapoda). — Ab- handlungen der Senckenbergishen Naturfor- schenden Gesellschaft 526:1—338 + pls. 1-58. Chokki, H. 1976. Preliminary report of the colour- ation of freshwater crab, Geothelphusa dehaani (White), with special reference to its distribu- tion.— Researches on Crustacea 7:177-182. (in Japanese with English summary). . 1980. Notes on the colouration of freshwater crab, Geothelphusa dehaani (White), in northern districts of Japan. — Researches on Crustacea 10: 57-60. (in Japanese with English summary). Haan, W. de 1833-1850. Crustacea.—Jn P. F. von Siebold, ed., Fauna Japonica sive descriptio an- imalium, quae in itinere per Japoniam, jussu et auspiciis superiorum, qui summum in India Ba- tava imperium tenent, suspecto, annis 1823- 1830 collegit. Notis, observationibus et adum- brationibus illustravit, XVII + XXXI + IX—- XVI + 243 pp., pls. A-J, L-Q + 1-55; Lugduni- Batavorum. [Leiden]. Minei, H. 1973. Potamoid crabs of the Ryukyu Is- lands, with descriptions of five new species (Crustacea, Decapoda, Potamoidea).—Journal of the Faculty of Agriculture, Kyushu Univer- sity 17:203-226. 1974a. Potamoid crabs of Taiwan, with de- scription of one new species (Crustacea, Decap- oda).—Journal of the Faculty of Agriculture, Kyushu University 18:239-251. 1974b. Studies on the freshwater crabs of Japan I. Genus Geothelphusa Stimpson.—The Nature and Animals 4(3/4):8—12. (in Japanese). Miyake, S., & J. K. Chiu. 1965. A new potamonid crab, Potamon (Geothelphusa) miyazakii sp. nov., aS an intermediate host of the lung-fluke from Formosa.—Journal of the Faculty of Ag- riculture, Kyushu University 13:595-600. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON —,, & H. Minei. 1965. A new fresh-water crab, Potamon (Geothelphusa) tenuimanus sp. nov., from Okinawa-jima, the Ryukyu Islands. —Sci- ence Bulletin of the Faculty of Agriculture, Kyu- shu University 21:377—382. (in Japanese with English summary). Ortmann, A. E. 1896. Das System der Decapoden- Krebse.—Zoologische Jahrbiicher. Abteilung fiir Systematik, Geographie und Biologie der Tiere 9:409-453. Rathbun, M. J. 1898. Descriptions of three new spe- cies of fresh-water crabs of the genus Pota- mon. —Proceedings of the Biological Society of Washington 12:27-30. 1904. Les crabes d’eau douce (Potamoni- dae).— Nouvelles Archives du Muséum d’His- toire Naturelle 6:225-312. 1905. Les crabes d’eau douce (Potamoni- dae).—Nouvelles Archives du Muséum d’His- toire Naturelle 7:159-321. Stimpson, W. 1858. Prodromus descriptionis ani- malium evertebratorum, quae in Expeditione ad Oceanum Pacificum Septentrionalem, a Repub- lica Federata missa, Cadwaladaro Ringgold et Johanne Rodgers Ducibus, observavit et des- cripsit, pars 5, Crustacea Ocypodidea.—Pro- ceedings of the Academy of Natural Sciences of Philadelphia 10:93-111. Suzuki, H., & E. Tsuda. 1991. Study on the color variation and distribution of a freshwater crab, Geothelphusa dehaani (White), in Kagoshima Prefecture.—Benthos Research 41:37-46. (in Japanese with English summary). White, A. 1847. List of the specimens of Crustacea in the collection of the British Museum. viii + 143 pp. British Museum, London. Marine Biological Laboratory, Faculty of Fisheries, Kagoshima University, 4-50-20 Shimoarata, Kagoshima 890, Japan. PROC. BIOL. SOC. WASH. 107(2), 1994, pp. 325-339 AEGLA PEWENCHAE, A NEW SPECIES OF CENTRAL CHILEAN FRESHWATER DECAPOD (CRUSTACEA: ANOMURA: AEGLIDAE) Carlos G. Jara Abstract.—The description of Aegla pewenchae, a new species of freshwater anomuran crab from the Rapel, Maule, Itata, Bio Bio, Imperial, and Tolten river basins is given. Its diagnostic characters are: 1) rostrum long and narrow, styliform, scarcely troughed both sides of rostral carina; 2) anterolateral angles of carapace spiniform, slightly divergent; 3) marginal scales of rostrum and hepatic lobes minute; 4) orbital spine clearly defined, with a smaller second one some distance below; 5) anterolateral angle of first hepatic lobe spiniform; 6) branchial borders smooth; 7) palmar crest wide and thin, its border deeply serrate or microdenticulate; 8) chelipeds with dorsal surface of propodus densely covered by minute lens-like scales; 9) anterolateral angle of second abdominal epimeron spiniform. Aegla pewenchae resembles, to a certain extent, A. rostrata Jara, 1977, and A. abtao Schmitt, 1942a. The zone between Angostura de Paine and Chillan, in central Chile, is one of the most ancient and densely populated zones be- cause of the fertility of its soils, mild climate and abundance of streams and rivers that provide water for agricultural irrigation. Ae- gla has seldom been registered in these flu- vial bodies. However, this seems to be re- lated more to the difficulties in identifying the specimens collected than with low col- lection efforts (N. Bahamonde, pers. comm.). Bahamonde & Lopez (1963) reported the presence of A. laevis talcahuano Schmitt, 1942b, from seven localities between the Zamorano River (tributary of the Cacha- poal-Rapel River system) and the Nuble River in Chillan, as well as A. concepci- onensis Schmitt, 1942a, from the Tronco River (Colchagua Province), and A. mau- lensis Bahamonde & Lopez, 1963, from La- guna del Maule. This paper describes a new species of Ae- gla, which was found almost continuously between the Colchagua Province in the north and the Cautin Province in the south. The specimens were collected during an eight- year period, while sampling the Andean or upper river stretches and the Coastal or low- er river stretches, on both sides of the Chil- ean main longitudinal highway (Carretera 5) which crosses most of the river systems along continental Chile. Aegla pewenchae, new species Figs. 1, 2, 3 Type material.—Holotype: Instituto de Zoologia, Universidad Austral de Chile, IZUA C-338, adult male collected in the Bio-Bio River, 16 km S of Los Angeles, un- der bridge of Carretera 5 (37°35'45’S, 72°16'30”W), Province of Bio-Bio, VIII Re- gion, Chile, 21 Feb 1983, by C. G. Jara. Allotype: IZUA C-338, adult female. Paratypes: IZUA C-338, 5 adult 66 (P1 to P5) and 3 adult 22 (P6 to P8). Same locality and date as holotype. Diagnosis. —Carapace longer than wide; rostrum elongate, styliform; anterolateral angles of carapace acute, slightly divergent; scales on rostral and hepatic borders very small; orbital spine well developed, dorsal 326 to a second smaller one: anterolateral angle of first hepatic lobe spiniform: branchial borders smooth, noticeably arcuate: palmar crest laminar, expanded, its border micro- denticulate to dentate: dorsum of prop- odus of chelae densely covered by minute lens-like scales: anterolateral angle of sec- ond abdominal epimeron sharply acute, spi- niform. Description of holotype.—Rostro-frontal end narrow, bound by slightly inflated an- terolateral lobes scarcely distinct from pro- togastnic prominences. Anterolateral lobe prolonged in conical acute spine, well sep- arated from orbital spine, its apex reaching posterior border of cornea. Orbital spine small, slender, acute, recurved towards an- terolateral angle: its length ¥ to % length of anterolateral spine. Frontal width about half precervical width. Orbits wide, compara- tively shallow. their depth about half their maximum width. Extraorbital sinus well defined, wide, slightly asymmetrical. Orbit- al margins with four or five minute well spaced scales. Rostrum narrow, styliform, its width at level of postenor margin of orbits *% its length. Cross-section of proximal half thombic; distal half subcircular. Apex end- ing in acute conical scale. Rostral margins defined only on proximal half of rostrum. Rostral carina low. narrow. reaching mid- point of rostrum flanked by shallow troughs: its proximal end marked by pair of tiny pits between protogastric lobes: its dorsum with two rows of minute scales which merge into one, distally. Carina is replaced by irregular row of well spaced scales which increase in size distally. on distal half of rostrum. Dorsum of precervical area uniformly convex; nO marginal plateau on dorsum of hepatic lobes. Epigastric prominences scarcely distinguishable, except by 1 or 2 nodules bearing 5 to 8 minute apical scales. Protogastric lobes less prominent, marked by 7 (left) and 5 (mght) apical scales in ar- quate row. Dorsum of carapace smooth, polished. slightly punctate. Margins of car- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON apace, between anterolateral lobe and cer- vical groove, almost straight. Hepatic lobes well delimited by shallow furrows: all of them with one acute apical scale. Remain- ing Margin with irregular row of small, un- equal scales. Dorsum of postcervical carapace mark- edly and uniformly convex. Sutures (lineae) fine, shallow. Cardiac area and areola wide: areola slightly inflated, its dorsum leveled with gastric area. Border of branchial areas smooth, narrowly marginated and slightly recurved, with irregular row of fine acu- minate scales mingled with short stiff setae. Dorsum of abdominal tergae slightly punctate: small tufts of fine short setae pro- trude from punctae: tufts thicker and more numerous on flanks of epimera. Anterolat- eral angle of second epimeron prolonged in short, stout, conical spine, its apex Over- reaching adjacent branchial border. Pleural angle of third and fourth epimera sharply acute. Telson cordiform, medially articu- late. Ventral surface of fourth thoracic ster- num fiat, slightly convex: its frontal border Straight at center and slightly concave near anterolateral angles abutting in short blunt cones. Chelipeds robust, left largest. Chelae stout, ovoidal. Left propodus markedly convex. inflated over its proximal 7s; nght subtrian- gular. Dorsum of propodus with oblique. blunt, low mdge between carpus-propodus and propodus-dactylus joints, and parallel to base of palmar crest. Palmar crest sub- rectangular, slightly excavate, its border clearly denticulate, merging anteriorly into predactylar lobe. Right crest with 7 acute denticles, left with 6, becoming progres- sively more recurved towards proximal end of crest; nght crest ends proximally in ro- bust denticle partially separated from pre- ceding ones by deep, wide notch. Postcrestal sinus deep. wide. Dorsum of chelae covered by tiny. blunt. conical scales that become larger toward distal end of both propodus and dactylus where they intermingle with bundles of short stiff setae. Dactylar lobe as VOLUME 107, NUMBER 2 Fig. 1. low blunt tubercle with apical scale 2 or 3 times larger than those on surface of prop- odus. Ventral surface of chelae slightly punctate, convex, polished, without scales or setae except at propodus-dactylus joint. Dorsum of carpus globular, with field of minute conical scales over lateroexternal half. Internal border with 3 robust acute conical spines that decrease in size proxi- mally; apex of second right spine bifid. Car- pal lobe spiniform, separated from adjacent articular nodule by shallow furrow; sepa- rated from distalmost spine of carpal crest by wide sinus. Ridge along dorsum of car- pus well marked, made up by 9 coalescent, little prominent, tubercles, each with 2 to 4 apical scales in oblique row. Lateroventral nodule of carpus-propodus joint with mi- nute flattened conical scale and short setae. Ventral face of carpus gently convex, with- out spine. Distodorsal vertex of merus of chelipeds as spiniform tubercle crowned by 2 acute scales and some stiff short setae. Dorsal border of merus sharp, with row of spiniform cones decreasing in size proxi- mally; distal cone twice size of subdistal; 10 Aegla pewenchae, new species. A, male holotype, dorsal view. B, female allotype. (right) and 9 (left) cones; the five distalmost procumbent. Ventral borders smooth, end- ing in acute conical spine; inner one slightly curved; external border, next to merus-car- pus articular node, with distally directed small acute cone. Ventral border of ischium slightly concave, with small tubercle crowned by conical scale and few stiff short setae at both ends. Distodorsal angle of me- rus of second and third pereiopods fringed by stiff short setae mingled with conical scales; 1 or 2 are central and prominent. Dorsal border of merus of second pereiopod fringed by long plumose setae; fringe absent on third pereiopod. Distal third of ventral median line of dactylus of second to fourth pereiopods with row of 3 to 7 acicular scales decreasing in size proximally. Description of allotype.— Aside from the relatively larger abdomen and smaller che- lae (secondary sexual characters), the allo- type differs from the holotype in the follow- ing aspects: scales on dorsum of chelae larger, particularly over distal half of propodus and dactylus; tubercles on dorsal midline of car- pus prominent, some spiniform; ventral 328 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. Aegla pewenchae, new species, male holotype. A, precervical carapace in dorsal view; B, same in lateral view; C, anterolateral angle of second abdominal epimeron in lateral view; D, telson plate; E, fourth thoracic sternum. border of merus of chelipeds with spine at limit between median and distal thirds of article; 2 similar spines at distal third of external lateroventral border of merus of right cheliped; predactylar lobe well de- fined, specially when seen from ventral face of chelae; midventral line of propodus of chelae with 1| (left) and 3 (right) scales form- ing row; anterolateral angles of fourth tho- racic sternum flattened and little scalloped; external flank of carpus-propodus articular node with spiniform tubercle; punctae on dorsum of carapace coarse, deep; extraor- bital sinus comparatively narrow. VOLUME 107, NUMBER 2 329 Fig. 3. Aegla pewenchae, new species, male holotype (continuation). F, left cheliped in dorsal view; G, left chela seen from dactylus top; H, right cheliped in dorsal view; I, right chela seen from dactylus top; K, same in ventral view; L, ischium and merus of left cheliped in ventral view. Color. —In life, dorsum of carapace uni- form in color varying among specimens from light olive green to dark green; color more intense in furrows and depressions of the carapace, over the frontorostral area and proximodorsal area of chelae. Distal zone of chelae and dactyls of pereiopods yellow- ish orange; intensity varying greatly among individuals. Scales amber-like, translucent, mounted on top of tubercles and spines ivo- ry white to light yellow changing progres- sively to general background color of car- 330 apace toward their bases. Ventral surface white in recently molted individuals, and smokey tan in animals in pre-ecdysis. In alcohol-preserved specimens, carapace creamy white varying from almost trans- lucent in just molted individuals to yellow- ish brown with dark brown spots in indi- viduals in pre-ecdysis. Superimposed on general background color, blueish tones mingle with other colors in varying intensity and extension. Dorsum of chelae, gastric, and cardiac areas, and dorsum of second to fourth abdominal segments of P7 and P8 light grayish blue changing to ivory white over branchial areas; center of posterobran- chial areas dark rose; same hue, but more intense, stains dactylus of pereiopods of P7. Etymology. —The name pewenchae is the latinized genitive feminine singular form of ““pewenche,” the aboriginal amerindians in- habiting the upland plateau at the origin of the Bio Bio River. Distribution. — Table 1 contains the basic data on records of Aegla deemed to be con- specific with A. pewenchae. Noteworthy is the distribution of A. pewenchae which ap- pears closely related to the Andean and pre- Andean stretches of the drainage systems of Rapel, Maule, Itata, Bio Bio, and Imperial rivers. Figure 4 shows the geographic range of A. pewenchae. The species is found from the Chimbarongo River, in the drainage ba- sin of the Rapel River, to the Donguil River, tributary of the Toltén River basin. The spe- cies is distributed along 480 km of the Chil- ean territory. In the Bio Bio River basin A. pewenchae is found along most of the 380 km of mainstream, including lakes Galletué and Icalma, sources of the Bio Bio River. The absolute upper limit of the species al- titude range (1150 m) is found in these lakes, while the lower limit (ca. 100 m) is found at several points along the Chilean Central Valley. In the drainage system of the Toltén River, A. pewenchae is only found in a small basin that drains the north-central area of the extra-Andean Valdivian territory. Remarks. —Aegla pewenchae presents PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON such a diverse combination of characters that it is difficult to advance a hypothesis about its phyletic relationships. Aegla pewenchae resembles A. rostrata, from lakes in the Toltén and Valdivia river basins, in having: rostrum narrow, elongate, scarcely troughed; orbits ample; orbital spine well defined, accompanied by second one in subordinate position; palmar crest wide, lightly built, its border dentate to micro- denticulate; dorsum of chelae covered by fine scales; and, dorsum of carapace dark green contrasting with marble-white ventral surface. In the Imperial River basin, both species coexist, being difficult to differen- tiate. A. pewenchae differs from A. rostrata in lacking denticles on the branchial border of the carapace, and in having the dorsal surface of anterior branchial area convex. Aegla pewenchae resembles A. abtao Schmitt, 1942a, distributed between the Toltén River basin and the Chiloé Island, in having: comparatively short, triangular, scarcely elongate rostrum; male chelae markedly unequal in size, fingers short and robust; dorsum of carapace coarsely punc- tate; palmar crest thick and narrow, its bor- der nodulate to dentate; spines on inner bor- der of carpus and dorsal ridge of merus of chelae short, thick, and stout. Aegla pew- enchae differs from A. abtao in lacking a dense row of conical scales along rostral bor- ders and thick protuberant scales on the sur- face of carapace. For comparison with the holotype of A. pewenchae, Fig. 5 shows the frontal end and the left chela ofa full grown male of A. abtao. Aegla pewenchae is a phenotypically well defined species, associated with the Andean piedmont zone of several Central Chilean river systems. Towards the West its distri- bution extends to the Central Valley, in gen- eral coinciding with the fluvial zone where the mean current velocity allows for the de- position of gravel and sand. These gradually replace the boulders and coarse gravel that predominate in the upper part of the basins where A. pewenchae is commonly found. VOLUME 107, NUMBER 2 331 Table 1.—Records of A. pewenchae, new species, in addition to the type series. All samples are deposited in the Collection of the Instituto de Zoologia of the Universidad Austral de Chile (IZUA-C). R. stands for river, L. for lake, and J for juveniles (specimens in which the gonopores are not visible). F Specimens Collection Latitude/ Date of number Locality longitude collection é 2 J 503 R. Chimbarongo 34°46’S, 71°08’W 12 Dec. 91 11 15 49 344-A R. Claro 35°11'S, 71°24’W 14 Feb. 83 14 13 — 393-A R. Claro 35°25'S, 71°41'W O01 Aug. 87 3 2 — 492-B R. Lircay 35°25’S, 71°34’W 13 Dec. 91 19 21 2 347-A R. Maule 35°28’S, 71°S7’'W 15 Feb. 83 23 25 24 342 R. Longavi 35°37'S, 71°46'W 17 Feb. 83 2 — — 493-A R. Putagan 35°46’S, 71°40’'W 13 Dec. 91 15 11 32 490 R. Ancoa 35°54’S, 71°30'W 13 Dec. 91 29 22 6 395-A R. Liguay 35°57'S, 71°41'W 10 Nov. 85 — 4 — 394-A R. Longavi 36°00’S, 71°43’W 10 Nov. 85 2 _ =— 489 R. Longavi 36°14'S, 71°30’W 13 Dec. 91 6 4 33 491-A R. Cato 36°15'S, 71°41'W 13 Dec. 91 6 7 4 396 R. Cato 36°17'S, 71°40'W 03 Mar. 73 1 2 _ 482 R. Nuble 36°29'S, 71°45'W 07 Dec. 91 14 40 83 480 R. Bustamante 36°34'S, 71°45’W 07 Dec. 91 D, 4 — 479 R. Chillan 36°41'S, 71°54’W 07 Dec. 91 8 16 6 478 R. Diguillin 36°54’S, 72°05’W 07 Dec. 91 4 9 22 477 R. Danicalqui 37°02'S, 72°01'W 07 Dec. 91 10 12 6 481 R. Cholgiian 37°11'S, 71°59'W 06 Dec. 91 16 6 7 470 R. Huepil 37°13'S, 71°57'W 06 Dec. 91 16 14 9 336 R. Bio Bio 37°17'S, 72°43'W 19 Feb. 83 _ 8 =— 474 R. Laja 37°18'S, 71°58'W 06 Dec. 91 20 15 43 520-B R. Cholguahue 37°29'S, 72°13'W 10 Oct. 92 10 16 2 472-A R. Quilleco 37°30'S, 71°59’W 06 Dec. 91 18 22 13 425 R. Bio Bio 37°33'S, 72°35'W 26 Jun. 85 2 7 — 475 R. Duqueco 37°35'S, 72°09'W 06 Dec. 91 19 23 28 427 R. Huequecura 37°41'S, 71°46'W 09 Apr. 87 3 8 1 340 R. Mulchén 37°43'S, 72°15'W 21 Feb. 83 13 10 1 222-A R. Malleco 37°47'S, 72°41'W 06 Jun. 81 _ 4 _ 335 R. Queuco 37°51'S, 71°38’W 21 Feb. 83 1 14 — 339 R. Renaico 37°51'S, 72°23’W 21 Feb. 83 3 3 = 334 R. Malleco 37°58’S, 72°26'W 22 Feb. 83 6 16 7 318 R. Traiguén 38°14’S, 72°19'W 23 Dec. 82 9 18 =— 323 R. Quino 38°18'S, 72°25'’W 23 Dec. 82 41 25 10 317-A R. Colpi 38°19'S, 72°47'W 22 Dec. 82 5 4 - 324-D R. Quillén 38°24'S, 72°47'W 22 Dec. 82 7 4 — 326-B R. Quillén 38°25'S, 72°56'W 22 Dec. 82 3 10 7 266 R. Bio Bio 38°38'S, 71°06’W 04 Feb. 69 6 5 12 186 L. Galletué 38°40'S, 71°19'W 16 Feb. 77 6 11 _ 319 R. Bio Bio 38°43'S, 71°09'W 05 Mar. 83 45 31 22 426 R. Bio Bio 38°46’S, 71°14’W 08 Apr. 87 — 16 2 187 L. Icalma 38°48’S, 71°17'W 17 Feb. 77 11 10 1 320-B R. Quepe 38°51'S, 72°37'W 21 Dec. 82 6 3 = 327-C R. Donguil 39°06'S, 72°41'W 21 Dec. 82 48 47 66 327-D R. Donguil 39°06'S, 72°41'W 21 Dec. 82 63 54 46 332 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON pr ool ersncacua, ae, —_— SAN FERNANDO / 0 ~ de) ee | f J \ — ( Maulgg § _ ies s 4 ya €+- i 7 ub on La lo b> ( Ue ANGELE 24 \ de Io iajad \ an \ Zo Vice R. A ( SP - 7} ° SCoHCOR ae SRE a8 cS (fi. \ Villarrica eaburgig\ <-ememe= INTERNATIONAL BOUNDARY f J —..—— REGIONAL BOUNDARY @.oncocHE [_ Se rH4 uo Fig. 4. Geographical range of A. pewenchae, new species. Stars indicate sampling localities; black dots, geographical localities. 333 VOLUME 107, NUMBER 2 1ew. iped, dorsal vi B, left chel lew; 1 carapace, dorsal vi , precervica Aegla abtao Schmitt, adult male. A Fig. 5. 334 Aegla abtao, which replaces A. pewenchae to the south of the Toltén River basin, also prefers this biotope (see Jara 1980:93-96; as A. riolimayana?). Both species share characters seemingly associated with living in moderate to fast running water environ- ments, 1.e., the smooth longer-than-wide oval carapace, the relatively short but nar- row acute rostrum, and the marked heter- ochely of adult males. However, this mor- phological similarity is not necessarily indicative of common ancestry between the two, and may well be the result of conver- gence. Aegla pewenchae, throughout its ample geographical range, shows a relatively wide exophenotypical variation which mostly af- fects the rostral and precervical morphology and the maximum size of the specimens. However, this species does not show mor- phological variations related to the lacus- trine environment. In A. rostrata, A. abtao (Jara 1986a), and A. denticulata (Jara 1986a, 1989), the lacustrine environment does seem to induce an overspinulation or spination of the carapace edges. The latitudinal limits of A. pewenchae seem to coincide, more or less, with well defined zoogeographical boundaries. In fact, its present northern limit lies at the Cacha- poal-Rapel River system, just to the south of the Maipo River system where two other freshwater decapod crustacean species reach their southern limit, namely, A. papudo Schmitt, 1942b (unpublished data), and Cryphiops caementarius (Molina 1782) (Ba- hamonde & Lopez 1963). A third species, the burrowing crayfish Parastacus pugnax (Poeppig 1835), reaches its northern limit at the Aconcagua River (Bahamonde & Lopez 1963), just to the north of the Maipo River system. At the Toltén River basin the situation is less clearly defined. Here A. ab- tao reaches its northern limit, while P. pug- nax, and the trichomycterid fish Bullockia maldonadoi (Eigenmann 1927), reach their southern limit (unpublished data). PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Morphological Variations The expression of the taxonomic char- acters varies among type specimens. In P1 and P5 the rostral apex overreaches corneae ca. two times their length, but in P6 and P7 it overreaches corneae by less than once their length. The orbital spine is comparatively small in the holotype and in P2, P7 and P8, its length being one fourth the length of an- terolateral spine of carapace, but in Pl equaling one half the length of anterolateral spine. The first left hepatic lobe of the ho- lotype, Pl, P7, and P8 ends in one acute scale, but the right in two; the opposite oc- curs in P6. The curvature of branchial bor- ders varies in relation to the relative width of carapace; it is minimum in the holotype and maximum in P1. Indentation of the pal- mar crest is notorious in the holotype, but insignificant in P1, P2, P3 and PS. The pre- dactylar lobe merges completely into the palmar crest of the left chela of Pl, P2 and P3, but in PS it does in both chelae. The ventral face of carpus of chelipeds is smooth in all specimens except in P1 and P3 (left chela), and PS (right chela), which have a spiniform tubercle. The inner ventral bor- der of merus of chelipeds is smooth, with one distal spine in all specimens except in P1, P4, and PS, which have two, and in P8, which has three. Table 2 contains the morphometric data of the type series. Measurements were taken with a digital caliper to the nearest 0.1 mm. The morphometric parameters here consid- ered were defined by Jara & Lopez (1981). The morphological variations most com- monly found among the specimens exam- ined, additionally to the type series (Table 1), are related with the rostrum and the fron- tal area. Figure 6 shows the variation ex- tremes in 29 specimens from Malleco Riv- er. A negative correlation between length and width of rostrum, and also between ros- tral length and frontal width appears among them. Figure 7 records the extreme varia- VOLUME 107, NUMBER 2 tions of the rostro-frontal area in 58 spec- imens from River Putagan; a negative cor- relation seems to exist between rostrum length and the upwards inclination of its apex. Figure 5-Al shows that the external border of the anterolateral lobes may be ar- quate instead of straightlined as in the type series. Maximum size variation along the Bio Bio River basin was found to conform to a cline (Fig. 8), between Icalma Lake and Na- cimiento. The most obvious environmental factors to which this phenomenon could be related are the thermic regime and the or- ganic productivity along the river. Lake Icalma is situated at 1150 m altitude while Nacimiento is at 130 m. However, no con- sistent field data are available to support this hypothesis. The graph suggests that the most favorable environmental conditions for A. pewenchae are those found in the mesorithral facies of the river (sector B). Natural History Field notes characterize A. pewenchae as a rheophilic species, consistently associated with rithral or fast running water environ- ments. Aegla pewenchae specimens were mostly found in places where water flowed over hard substrates (boulders and stone blocks) with little to no deposition of fine sediments. In the monomictic ultraoligo- trophic lakes Galletué and Icalma, A. pew- enchae was also found on boulders and hard substrate along the shoreline. In these pris- tine environments, the specimens typically presented clean carapaces. In contrast, when A. pewenchae was found in eutrophic bio- topes with high fine sediment deposition rates and high benthic primary productiv- ity, 1.e., the rivers associated with intensive fruticulture, between Chimbarongo and Talca, the specimens presented dirty cara- paces, covered by muddy deposits and epi- bionts. The thermic conditions in which 4. pew- enchae was found also present an interesting 3mm Fig. 6. Morphological variation in A. pewenchae, new species. Rostrum and rostral end of three males from Malleco River (IZUA C-334). contrast. In lakes Icalma and Galletué the thermic regime ranges between 5.5°C for Icalma Lake and 7.5°C for Galletue Lake, in winter, to 19°C and 17.5°C in summer, respectively (Parra et al. 1993). In the rivers of the fruticultural zone, water temperatures rise to approximately 25°C in summer (Jara, field notes). At least one instance of migratory behav- 336 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 7. Morphological variation in 4. pewenchae, new species. Rostrum and frontal end of two extreme variant males from Putagan River (IZUA C-493). Al & B1, rostrum and frontal end in dorsal view; A2 & B2, rostrum in dorsal view; A3 & B3, rostrum in lateral view. VOLUME 107, NUMBER 2 34 32 30 28 26 24 22 20 Male maximum size (CL mm) A B 337 Cc D Arbitrary river sectors Fig. 8. Variation of male maximum size (CL, carapace length) in A. pewenchae, new species, along Bio Bio River mainstream. n = number of populations sampled per river sector. In each sample only the size of the largest male was recorded. Sample size ranged between 7 and 98 specimens (X = 22.5; SD = 27.2). River sectors are roughly defined as follows: A, upland river in a high mountain steppe-like environment, 0 to 20 km downstream from sources (localities: Icalma, Galletué, Marimenuco and Liucura); B, torrential river in a deep piedmont valley, 115 to 135 km from sources (localities: Queuco and Huequecura); C, lowland moderately fast running river, 185 to 210 km from sources (localities: bridge under highway #5 and Coihue); D, lowland slow running river on sandy substrate, 245 km from sources (locality: La Laja). ior was observed for this species. On 21 February 1983, a great number of individ- uals, mostly adult males, were seen migrat- ing upstream along the banks of the Queuco River (Bio Bio River basin). Animals moved at arate of 600 to 1200 individuals per hour, 24 hours a day; the cause of this migration could not be established. In relation to the molting period, in the Queuco River, females are known to molt in October. According to field notes, most of the individuals sampled between Los An- geles to the south and Chillan to the north during December 1991 were in advanced pre-ecdysis or in recent post-ecdysis. No data are available about reproductive biology or trophic niche of A. pewenchae. The benthic community in which A. pew- enchae is found varies in the number of species and in the abundance of organisms from one river to another. In general, it is composed by rheophilic “‘clean water”’ spe- cies such as leptophlebiid and baetid ephemeropterans, hydropsychid, rhyaco- philid, and sericostomatid trichopterans, gripopterygid and diamphipnoid plecopter- ans, elmid and psephenid coleopterans, and trichomycterid fishes. Other species of Aeg/a are occasionally part of these communities. In the rivers 338 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 2.—Morphometrics of A. pewenchae, new species, type series. All measurements in mm. M = male; F = female; Holo = holotype; Allo = allotype; P1 to P8 = paratypes. CL, carapace length, distance between rostral apex and posterior margin of cephalothorax; RL, rostral length, distance between rostral tip and a transverse line tangent to deepest point of orbits; PCL, precervical length, distance between rostral tip and midpoint of cervical groove; FW, frontal width, distance between tips of anterolateral angles of carapace; PCW, maximum precervical width, distance across third hepatic lobes; CW, maximum carapace width; LCL, left cheliped length; RCL, right cheliped length; L2PL, length of second pereiopod; L2DL, dactylar length of second left pereiopod; L4DL, dactylar length of fourth left pereiopod; TL, telson length. * = rostrum broken; e = estimated. Holo Allo Pl P2 P3 P4 PS P6 P7 P8* Sex M F M M M M M F F F CL 25.8 23.8 29.7 26.9 26.8 26.0 ADI 23.9 23.8 24.7e RL Sol! 4.8 6.1 4.9 5.6 6.2 4.8 4.7 4.0 4.9e PCL 17.0 15.6 19.5 17.4 17.6 16.9 16.6 15.6 IS.3 16.5e FW WS 6.8 8.9 8.5 8.1 Voi) 7.3 7.1 6.9 7.8 PCW 13.5 13.0 16.2 14.6 14.6 13.2 14.3 13.0 13.6 13.5 CW 19.9 18.9 24.5 21.4 21.5 19.6 20.5 19.2 19.8 20.4 LCL 35.4 27.9 43.4 40.4 38.2 32.8 36.9 28) 28.6 29.1 RCL 32.3 27.9 40.4 37.0 35.4 30.6 35.5 27.6 PPS 28.9 L2PL 36.5 32.9 40.7 39.1 39.4 36.7 38.4 34.2 33.4 34.8 L2DL 8.4 7.5 8.8 8.7 8.9 8.0 8.4 7.4 7.6 7.8 L4DL 8.7 7.9 9.7 8.9 — 8.7 8.3 8.2 8.0 8.4 TL 5.3 5.9 5.1 5.3 5.0 4.7 4.8 5.8 5.9 D2) Claro, Lircay, Maule, Putagan, Liguay, and Cato (Maule River system) and Cholguahue and Quilleco (Bio Bio River system) A. pew- enchae was collected together with A. aff. laevis talcahuano Schmitt, 1942b. In rivers Malleco (Bio Bio River system) and Don- guil (Toltén River system) it was collected together with A. denticulata Nicolet, 1849. In rivers Colpi, Quillén, and Quepe (Im- perial River system) and Donguil (Toltén River system) it was collected together with A. rostrata Jara, 1977 (fluvial form), and in the river Cholchol (Imperial River system) it was collected with A. spectabilis Jara, 1986b. Acknowledgments The author thanks Mr. Raul Arriagada for his valuable collaboration during field work in December 1991, and Messrs. René Navarro and David Manriquez for their col- laboration during the field work in Decem- ber 1982. Thanks are extended to Miss M. C. Vasquez for the curation of IZUA’s Aegla collection, and to P. Stegmaier for the art work. Improvement of the English version is due to Mrs. P. Araya, and two reviewers. This paper was financed by grants FON- DECYT 91-0900 and DID S-91-4 of Di- reccion de Investigacion y Desarrollo of the Universidad Austral de Chile. Literature Cited Bahamonde, N., & M. T. Lopez. 1963. Decapodos de aguas continentales en Chile.—Investiga- ciones Zoologicas Chilenas 10:123-149. Eigenmann, C. H. 1927. The fresh-water fishes of Chile.—Memoirs National Academy of Sci- ences 22(2):1-63. 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). Unpublished M.S. thesis, Universidad Austral de Chile, Valdivia, 126 pp. 1986a. El “efecto lago” sobre el exofenotipo de macrocrustaceos dulceacuicolas del Sur de Chile. Pp. 239 in Programa Cientifico y Resu- menes de Simposios y Comunicaciones Libres. X Congreso Latinoamericano de Zoologia. Vina del Mar, Chile. 1986b. Aegla spectabilis, a new species of VOLUME 107, NUMBER 2 freshwater crab from the eastern slope of the Nahuelbuta Coastal Cordillera, Chile.—Pro- ceedings of the Biological Society of Washington 99:34—-41. 1989. Aegla denticulata lacustris, new sub- species, from Lake Rupanco, Chile (Crustacea: Decapoda: Anomura: Aeglidae).— Proceedings of the Biological Society of Washington 102: 385-393. , & M. T. Lopez. 1981. Anew species of fresh- water crab (Crustacea: Anomura: Aeglidae) from insular South Chile.— Proceedings of the Bio- logical Society of Washington 94:88-93. Molina, G. I. 1782. Saggio sulla storia naturale del Chili. Bologna, pp. 1-367. Nicolet, H. 1849. Crustaceos. Jn C. Gay, ed., Historia Fisica y Politica de Chile, Zoologia 3:115-318. Atlas Zoologico, Crustaceos, Paris. 339 Parra, O., H. Campos, W. Steffen, G. Agiiero, S. Ba- sualto, & M. Vighi. 1993. Estudios Limnolé- gicos en los Lagos Icalma y Galletué.— Mono- grafias Cientificas EULA (in press). Poeppig, E. 1835. Reise in Chile, Peru und auf dem Amazonenstrome wahrend der Jahre 1827— 1832. Vol. 1, Leipzig, pp. 1-466. Schmitt, W.L. 1942a. Two new species of Aegla from Chile. — Revista Chilena de Historia Natural, vol. 44(1940):25-31, pl. 5. . 1942b. The species of Aegla, endemic South American freshwater crustaceans. — Proceed- ings of the United States National Museum 91: 431-520, pl. 25-28. Instituto de Zoologia, Universidad Aus- tral de Chile, Casilla 567, Valdivia, Chile. PROC. BIOL. SOC. WASH. 107(2), 1994, pp. 340-353 DESCRIPTION OF THE GHOST SHRIMP EUCALLIAX MCILHENNYI, NEW SPECIES, FROM SOUTH FLORIDA, WITH REEXAMINATION OF ITS KNOWN CONGENERS (CRUSTACEA: DECAPODA: CALLIANASSIDAE) Darryl L. Felder and Raymond B. Manning Abstract. — Eucalliax mcilhennyi, new species, is described from an intertidal sandflat bordering Fort Pierce Inlet on the Atlantic coast of Florida. The species is distinguished from known congenerics of the Eucalliinae, all of which are restricted to the western Atlantic. Detailed comparisons are made to E. jonesi (Heard 1989), from Bimini Harbor, Bahamas. Specimens from Florida were first thought to represent E. jonesi because of superificial resemblance, simi- larities in habitats, and proximity of collection localities. The two species differ from each other and their congenerics in a number of morphological characters, including relative development of the front and rostrum of the carapace, spi- nation of chelipeds, shape of gonopods, and ventral plating of abdominal so- mites. Abbreviated larval development and limited capacity for dispersal are inferred by the large eggs found on ovigerous females of species in this genus, and may serve to maintain isolation of regional populations. This would be consistent with morphological evidence that Eucalliax has extensively endem- ized within the tropical western Atlantic, even in the absence of evidence for major historical disjunctures in appropriate habitat. Over the last decade, we have used yabby pumps (see Hailstone & Stephenson 1961, Manning 1975) to collect extensively from intertidal substrates in the vicinity of Fort Pierce, Florida. Our efforts there have re- vealed a number of previously unknown in- faunal decapods, some of which we have described in previous papers (Felder & Manning 1986; Manning & Felder 1989, 1992). Materials from this region have also provided a basis for systematic revisions and new distribution records, especially for members of the Callianassidae (Manning 1987, 1993; Manning & Felder 1986, 1991; Manning & Heard 1986; Manning & Le- maitre 1994). One of our collecting sites within the southern Indian River lagoon, a small in- tertidal sandflat just inside Fort Pierce Inlet (see Felder & Manning 1986), has produced a particularly rich assemblage of fossorial stomatopod, thalassinid, and alpheid crus- taceans. The thalassinids taken from this small area have included representatives of Upogebia, Callichirus, Neocallichirus, Bif- farius, and a new genus (Manning & Le- maitre 1994). Our collections there have also included infrequent occurrence of two spe- cies of ghost shrimp that we assigned to a new genus, Eucalliax Manning & Felder, 1991. One of these species, perhaps the same as that previously reported from south Flor- ida as ““Eucalliax quadracuta” (Biffar 1971; specimen destroyed by fire), we have ten- tatively grouped with the ‘Eucalliax quad- racuta complex’ until such time as we can complete further comparative studies from throughout the range of that group. The oth- VOLUME 107, NUMBER 2 er, allied to E. jonesi (Heard 1989) from the nearby northern Bahamas and E. cearaensis from Brazil (Rodrigues & Manning 1992), is herein recognized as a new species. Material examined is listed by location followed by date, collector, number of spec- imens per sex and condition (imm = im- mature, mutl = mutilated, ov = ovigerous), and, if applicable, museum number. Size is expressed as postorbital carapace length (CL) measured in millimeters (mm), except where compared as total length under ““Remarks’”’ section. The holotype and some paratypes of Eucalliax mcilhennyi have been depos- ited in the National Museum of Natural History, Smithsonian Institution, Washing- ton, D.C. (USNM). Paratypes have been de- posited in the University of Southwestern Louisiana Zoological Collections, Lafa- yette, Louisiana (USLZ). In addition to type materials of E. cearaensis and E. jonesi available at the Smithsonian Institution, the paratype of Eucalliax jonesi was obtained on loan from the Gulf Coast Research Lab- oratory (GCRL) in Ocean Springs, Missis- sippi, and the types of Eucalliax quadracuta (Biffar, 1970) were obtained on loan from the Museum of Comparative Zoology, Har- vard University, in Cambridge, Massachu- setts. Eucalliax Manning & Felder, 1991 Eucalliax mcilhennyi, new species Figs. 1-6 Type material. —Station RBM FP-89-4, sandflat with sparse seagrass, 27°27.7'N, 80°18.7'W, south side of Fort Pierce Inlet, north side of US Highway A1A causeway between Fort Pierce and South Hutchinson Island, Indian River lagoon, St. Lucie County, Florida, 11 Aug 1989, coll. R. B. Manning, R. Brown, and W. Lee, é holotype (CL 10.1 mm), USNM 267112; (same site as holotype, except where otherwise indi- cated) station RBM FP-85-4, voucher for color photo, 23 Jul 1985, coll. R. B. Man- ning and D. L. Felder, 1 6 (CL 9.0 mm), USLZ 3538; station RBM FP-86-1, 11 Aug 341 1986, coll. R. B. Manning, D. L. Felder and W. D. Lee, | ov 2 (CL 10.3 mm), USNM 267113; station RBM FP-86-2, voucher for color photo, 11 Aug 1986, coll. R. B. Man- ning, D. L. Felder and W. D. Lee, 2 2 (1 ov, 1 mutl; CL 11.6 mm, 10.5 mm), USNM 267114; station RBM FP-86-3, 12 Aug 1986, coll. R. B. Manning and D. L. Felder, 1 2(CL 10.9 mm), USNM 267115; station RBM FP-86-4, 12 Aug 1986, coll. R. B. Manning, D. L. Felder and W. D. Lee, | 2 (CL 9.9 mm), USNM 267116; station RBM FP-86-6, 14 Aug 1986, coll. R. B. Manning, D. L. Felder and W. D. Lee, 3 2 (1 ov, 1 mutl; CL 11.2, 11.2, 11.0 mm), USNM 267117; no station number, 4 Jun 1993, coll. D. L. Felder, 1 ov 2 (CL 9.7 mm), USLZ 3537; station RBM FP-88-3, hard packed sand along shore, 27°28.2'N, 80°18.2'W, south side of Coon Island, north shoreline of Fort Pierce Inlet, Indian River lagoon, St. Lucie County, Florida, “with Pinnixa,” 21 Apr 1988, coll. R. B. Man- ning, W. D. Lee, M. Schotte, and C. King, 1 6(CL7.5 mm), USNM 267118. Diagnosis. —Rostrum broad, weakly pro- duced. Carapace dorsally lacking strong, longitudinal ridges. Antennal peduncles overreaching antennular peduncles. Che- lipeds with hooked marginal spinules on is- chium, lacking acute teeth or spines at distal corners of carpus. Male first pleopod originating from distinct ovoid ventral plate on abdomen, terminally bifurcate, with sin- gle, short subapical process. Description. —Dorsally, carapace much less than (about 73) combined lengths of ab- dominal segments 1 and 2 (Fig. 1a). Frontal margin of carapace with broad, triangular rostrum; rostrum acute terminally and flanked by weakly excavate shoulders (Fig. 2a) forming anteriorly produced promi- nences just lateral to margins of eyestalks; rostrum extending less than '2 visible length of eyestalks in dorsal view, ventrally bearing tuft of setae, longest of which extend ante- riorly between eyestalks to cornea. Carapace lacking distinct dorsal oval and cardiac 342 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. mm) USNM 267114: a, Lateral view; b, External surface of right (major) chela. Scale lines indicate 5 mm. prominence; lacking rostral carina except for slightly raised postrostral area between pairs of postrostral punctae. Cervical groove evident as suture, disjunct at dorsal midline, extending anteroventrally to complex net- work of sutures in posterior 7% of carapace; one branch from this point continued an- teriorly as weakly carinate, sinuous, longi- tudinal suture, of which longest tract ter- minates anteriorly in antennal notch of carapace margin. Strong, raised hepatic boss in anterior '4 of carapace just dorsal to cer- vical suture. Linea thalassinica strong, par- allel to midline of carapace over most of length, diverging slightly posterior of car- diac suture. Cardiac suture well defined, in- complete across dorsal midline of carapace. Eyestalks dorsally flattened, length equal to or just greater than 2 times width, in dorsal view reaching beyond basal antennal article; mesial surfaces broadly triangular, flattened so eyestalks fit closely together at midline; anterolateral margin of eyestalk ar- cuate, joining mesial margin anteriorly in narrow, upturned tip; pigmented region in distal 2 of dorsal surface, area of dark pig- mentation variable, sometimes exceeding the weakly evident corneal surface; some- times with | or more setose punctae dorsally near midlength of eyestalk. Antennular pe- Eucalliax mcilhennyi, new species, from Fort Pierce Inlet, Florida, paratype ovigerous 2 (CL 11.6 duncle shorter than and not so heavy as antennal peduncle; basal article laterally and ventrally inflated to accommodate stato- cyst, opening to which is occluded by closely set fan of anteromesially directed setae overlain by eyestalk; second article slightly longer than basal article, third article about */; length of second; second and third articles with ventrolateral row of long, ventrally di- rected setae, continued onto ventral ramus of flagellum; rami of flagellum about equal in length, near 5 times length of third article of peduncle; ventral ramus ventrally setose, line of long setation ventrolaterally and line of slightly shorter setation ventromesially; dorsal ramus primarily with sparse short setae, subterminal articles of dorsal ramus heavier than those of ventral ramus, and endowed with thick line of ventral aesthet- ascs. Antennal peduncle more than 1.5 times length of antennular peduncle; basal article with dorsolateral carina bearing regular line of fine setae above laterally produced ex- cretory pore; second article with deep, di- agonal ventrolateral furrow, distally with field of long setae below ventrolateral suture and another on dorsolateral surface, broad, articulated dorsal scale at joint with third article; third article elongate, slightly longer than fourth or combined lengths of first two, VOLUME 107, NUMBER 2 b.d c.g a 343 Fig. 2. Eucalliax mcilhennyi, new species, from Fort Pierce Inlet, Florida; a—e, g, holotype 6 (CL 10.1 mm) USNM 267112: f, paratype 2 (CL 11.6 mm) USNM 267114: a, Anterior carapace, eyestalks, and antennae, dorsal view; b, Right (major) cheliped, external surface; c, Right (major) cheliped internal surface; d, Left (minor) cheliped, external surface; e, Ischium of right (major) cheliped of 6, internal surface; f, Ischial dentition, nght (major) cheliped of 2, internal surface; g, Posterior abdomen, telson, uropods, dorsal surface. Scale lines indicate 2 mm. proximolaterally with unfused condylar process articulated to distolateral extreme of second article; fourth article narrower than third; flagellum sparsely setose and more than 4 times length of antennular flagella. Mandibles set below the produced, rounded, median lobe of epistome; man- dible (Fig. 3a) with large, terminally setose, 3-segmented palp, elongated third article of palp terminally rounded; incisor process with well-defined teeth on cutting margin, field of 3-4 large distal teeth separated from large proximal tooth by line of subpectinate lower teeth, internal surface with lip giving rise to molar process proximal to incisor teeth; paragnath (not figured) uncalcified, set against proximal surface of molar process, distolateral corner slightly produced and opposing teeth of molar process. First max- illa (Fig. 3b) with endopodal palp long, nar- row, terminal article deflected proximally at articulation; proximal endite densely se- tose on concave margin, terminally with dense field of complex setae; distal endite elongate, terminally truncate and armed with stiff bristles; exopodite low, rounded. Sec- ond maxilla (Fig. 3c) with endopod nar- rowed abruptly at distal end, terminus di- rected mesially, first and second endites each longitudinally subdivided, exopod forming large, broad, scaphognathite. First maxilli- ped (Fig. 3d) with proximal endite trian- gular, marginal setation including stronger, curved setae at distal corner; distal endite elongate, ovoid, mesial half of external sur- face and all margins heavily setose, internal surface concave; exopod triangular, divided by transverse suture; distal part broader and 344 with longer marginal setation at its mesial end, proximal part with field of mesially directed setae near mesial end; epipod large, broad, weakly subdivided by transverse su- ture, its anterior end tapered, angular. Sec- ond maxilliped (Fig. 3e) with long, narrow endopod; endopodal merus arcuate, slightly heavier in proximal half than in distal, flex- or margin with dense fringe of long, close- set setae; carpus short; propodus heavy, weakly arcuate, length equal to or less than 2 times width, equal to or less than 2 length of merus; dactylus short, about 12 length of propodus, extensor margin arcuate; exopod about as long as endopodal merus, margin- ally fringed by long setae, subdivided by weak transverse suture at '4 length; epipod small, with short, rounded proximal lobe and narrow distal lobe. Third maxilliped (Fig. 3f) without exopod; endopod with long, dense setation on mesial margin; endopodal ischium subtriangular, slightly longer than broad, proximomesial lip forming produced lobe or subacute corner, internal surface with low medial, longitudinally oriented eleva- tion bearing well-defined curved row of about 9-11 sharp teeth, usually with 2-3 smaller supplementary teeth trailing ventral to proximal end of primary row; merus subquadrate, slightly broader than long; car- pus strongly flexed in proximal third, with setose lobe on flexor margin, internal sur- face faceted, superior facet glaborous except for marginal setae, and inferior facets setose; propodus large, subtriangular, about as broad as long, proximal %4 of inferior margin forming large, rounded, densely setose lobe; dactylus broad terminally, slightly longer than broad, fringed with very dense field of close-set, stiff setae on broad terminal mar- gin. Branchial formula includes exopods and epipods as described for first and second maxillipeds above; branchiae limited to sin- gle rudimentary arthrobranch on second maxilliped, pair of arthrobranchs on third maxilliped, and pair of arthrobranchs on each of the first through fourth pereopods. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON First pereopods with major and minor cheliped strongly developed (Fig. 1a), near equal in size but dissimilar in shape of prop- odus and dentition of fingers, especially in males; major cheliped located on either right or left side of body. Major cheliped of ma- ture male (Fig. 2b, c) massive and strongly calcified; ischium slender, superior margin sinuous, inferior (flexor) margin with row of small, distinctly hooked denticles; merus unarmed, about 1.8 times longer than broad; carpus broad, increasing in breadth distally, inferior margin arcuate, superior and infe- rior margins keeled, terminated distally in blunt corners; propodus heavy, length (in- cluding fixed finger) about 1.7 times height, inner surface of palm smooth, without swol- len proximal boss; weak unarmed furrow extending posteriorly from just below gape of fingers on outer face of palm; superior and inferior propodal margins keeled, keel of inferior becoming ill-defined beyond midlength and absent on fixed finger; fixed finger thick, heavily calcified, prehensile margin armed with 2 small well-separated triangular teeth in proximal 4 and broad, microserrated tooth just proximal to mid- length, with distal half of margin unarmed and terminated at subacute upturned tip; dactylus with subacute, hooked tip, external shoulder of superior margin with setose punctum abutted against low tubercle in proximal % of length in addition to line of about 6 setose punctae on internal side of superior margin, inferior (prehensile) mar- gin with low, sinuously margined tooth en- compassing distal 2 and separated from tooth on proximal 2 by rounded gap, prox- imal tooth with weakly bicarinate margin bearing scant small tubercles or microser- ration, proximal tooth separated from prox- imal end of inferior margin by rounded gap. Major cheliped of female also massive (Fig. la, b) but less heavily calcified and slightly different in sculpture than that of typical mature males; teeth of dactylus usually of slightly lower profile than in males, those of fixed finger usually centered more proxi- VOLUME 107, NUMBER 2 345 Fig. 3. Eucalliax mcilhennyi, holotype 6 (CL 10.1 mm), from Fort Pierce Inlet, Florida, USNM 267112, right appendages; a—e, external surface; f, internal surface: a, Mandible, excluding paragnath; b, First maxilla; c, Second maxilla, setae not shown; d, First maxilliped, setae not shown; e, Second maxilliped, setae not shown; f, Third maxilliped. Scale lines indicate 1 mm. mally than in males, both fingers relatively less massive, more narrow, sometimes more acutely tipped, than in males; propodus not as long relative to height, and with margins more arched, than in males. Minor cheliped (Fig. 2d) slightly lighter, less armed than major, inferior (flexor) mar- gin of ischium with line of distinct hooked spinules; merus unarmed; carpus with blunt distal corners; propodus with distinct un- armed furrow extending posteriorly from just below gape of fingers on outer surface, fixed finger tapered to very narrow acute or subacute tip, prehensile margin proximally serrate; propodus less elongate, relative to height, in females (Fig. la) and juvenile 346 males than in mature males; dactylus nar- row, with subacute tip, unarmed on pre- hensile margin. Second pereopod (Fig. 4a) chelate, most of flexor margins of ischium and merus lined with evenly spaced long setae, similar setae patchy and restricted primarily to distal patches on flexor margin in carpus, inferior margin of propodus with similar setal patches which are long proximally, pro- gressively more reduced in length and stiff- ened distally, subterminally becoming dense patch of short, stiff bristles; prehensile mar- gins of both fingers corneous, finely and uni- formly microserrate along straight edge over most of length, microserration terminating proximally at small corneous tooth and ter- minating distally in thickened corneous tips of fingers; superior margin of dactylus slightly sinuous, with patches of stiff, arched bristles becoming increasingly reduced in length, close-set and more arched distally. Third pereopod (Fig. 4b) merus length about 2 times width, flexor margin weakly sinu- ous, typically with 2 small prominences bearing tufts of setae; carpus broadly flared distally to produce strong inferior lobe, width at this point about *%4 length, inferior lobe terminally with field of long arched setae, diminishing in length toward articulation with propodus; propodus with strong prox- imally directed lobe on inferior margin, lobe terminally with field of long arched setae diminished distally along margin to close- set shorter bristles that become slightly lon- ger at distal extreme, superior margin with tufts of long arched setae, patterned tufts of lighter setae on outer face of article; dactylus tear-shaped, length about 1.4 times width, terminating in narrow corneous tip hooked toward external side, inferior margin sinu- ous, Outer (external) face crossed by fields of short, slightly hooked setae in which lon- gest setae are near superior margin, with separate, dense field of slightly heavier short weakly hooked setae along lower extreme of external face and inferior margin. Fourth pereopod (Figs. la, 4c) not subchelate, in- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ferodistal corner of propodus rounded with- out evidence of fixed finger; dense setation on outer surface of both propodus and tear- shaped dactylus divided into upper and lower fields, setae slightly stronger in lower fields of both, densest on dactylus, especial- ly on and near inferior margin; internal sur- face of propodus distally with single large very long seta originating from near supe- rior margin and reaching distally well be- yond tip of dactyl; dactyl terminated in nar- row corneous tip hooked toward external side. Fifth pereopod (Figs. 1a, 4d) minutely chelate, opposable surfaces of propodus and minute dactylus excavate, spooned, termi- nally rounded, forming beak-like chela ob- scured by dense fields of setation on distal % of propodus and superior surface of dac- tylus; corneous prehensile lip of propodus finely divided into arched row of close-set denticles. Texture of abdominal somites (Figs. la, 5a, b) smooth dorsally, glabrous, typically with setae limited to isolated pairs of setose punctae on first tergite, strongest of which are in posterior half; second tergite with pos- terolateral crescentic line of small, lightly setose granules, anterior to which is short oblique line and posterior of which are sev- eral small fields of similar granules and punctae, posterolateral-most of which bears long setae; third to fifth tergites each with a distinct, lateral, transverse field of long soft setae, posterior to each of which lies a small field of long stiff setae on the lateral margin; sixth tergite (Figs. 1a, 2g) with lateral, lon- gitudinal finely setose lines of small gran- ules, primary line turning to transverse and directed toward midline in posterior half on lateral lobe of tergite, posteriorly with strong tuft of long stiff setae at each posterolateral corner, similar tuft on posterior margin overlying each anterolateral corner of tel- son. Shape of first abdominal tergite nar- rowed anteriorly, anterior 4 offset by lateral notch and subquadrate (most striking in mature males); second tergite elongate, at least 1.6 times median length of third; third VOLUME 107, NUMBER 2 347 a,b,c,d e Fig. 4. Eucalliax mcilhennyi, new species, from Fort Pierce Inlet, Florida; a—c, holotype ¢ (CL 10.1 mm) USNM 267112; d-e, paratype 2 (CL 11.6 mm), USNM 267114: a, Right second pereopod, external surface; b, Right third pereopod, external surface; c, Right fourth pereopod, external surface; d, Right fifth pereopod, posteromesial surface; e, Enlarged terminus of fifth pereopod. Scale lines indicate 2 mm. tergite with deep, elongate anterolateral sul- cus extending across anterior 3 of each side. Anterior 2 of first abdominal somite wrapped ventrally by girdle of thickened, leathery integument, girdle rounded later- ally and transversely bisected by a furrowed suture, posterior half of somite ventrally with pair of conspicuous ovoid plates comprised of similar thickened integument, each of which (in males and females) articulates to first pleopod at its posterior extreme; medial posterior margin of first abdominal somite marked a triangular or rounded, anteriorly extended plate of thickened, leathery integ- ument continuous with massive continuous covering of leathery integument that forms articular membrane between first and sec- ond abdominal somites and that covers en- tire ventral surface of second abdominal so- mite; similar leathery integument largely covering ventral surfaces of remaining ab- dominal somites. First pleopod of male and female unira- mous, composed of 2 articles; in male (Fig. 6a, b), total length about % that of second pleopod, distal article about equal in length 348 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON a,b,c,d Fig. 5. Eucalliax mcilhennyi, new species, holotype 6 (CL 10.1 mm) from Fort Pierce Inlet, Florida, USNM 267112: a, First and second abdominal segments, ventral surface, setae not shown; b, First and second abdominal segments, right lateral surface, setae not shown. Eucalliax jonesi (Heard 1989) from Bimini Harbor, Bahamas; c, d, paratype 6 (CL 9.6 mm) GCRL 1136; e, holotype 6 (CL 9.8 mm) USNM 221861: c, First and second abdominal segments, ventral surface, setae not shown; d, First and second abdominal segments, right lateral surface, setae not shown; e, Left (major) chela, internal surface, setae not shown. Scale lines indicate 5 mm. to proximal and bifurcate at about *4length, article, terminal article with longest setae with acute tip of spooned terminal end di- on broad shoulder just beyond midlength. rected laterally; in female (Fig. 6c) total Second pleopod of male and female bira- length subequal to that of second pleopod, mous, with appendix interna on endopod; proximal article about % length of terminal in male (Fig. 6d), dense setation largely re- VOLUME 107, NUMBER 2 stricted to distal extreme of exopod, distal lobe of endopod and appendix masculina, appendix masculina markedly overreaching distal lobe of endopod and with small ap- pendix interna at its base; in female (Fig. 3e), both rami with long setae, appendix interna small and constricted distally. Third to fifth pleopod pairs (Figs. la, 6f) forming large, posteriorly cupped fans when cross- linked by hooked setae of appendices inter- nae on opposed margins of endopods; endopod of each subtriangular, appendices internae finger-like, movably articulated to mesial margin of endopod. Telson (Fig. 2g) broader than long, subrectangular, broadest at lateral lobes in posterior half, posterior margin weakly bilobate; dorsal surface with medial tuft of large setae separating two halves of strong, transverse carina; lateral margins sinuous, without setae; posterior margin with tuft of setae on each of the weak lateral lobes. Uropod (Fig. 2g) with heavy, blunt, posterolaterally directed tooth on protopod, tooth over-reaching anterolateral margin of endopod; endopod broad, subov- al, slightly longer than broad, dorsal surface with tuft of long setae on posterior 1, setae of posterior margin longest posterolaterally; exopod with anterodorsal plate falling well short of distal endopod margin, distal edge of plate lined with short, thick spiniform setae grading to thinner longer setae of ex- opod margin and long stiff, spiniform setae at posterodistal corner of plate, distal mar- gin of exopod with dense fringe of setation grading to large spiniform setae of postero- distal margin. Size.—Among the materials examined, the largest male is the holotype (CL 10.1 mm) and the largest female is an ovigerous paratype (CL 11.6 mm). Egg size (max. di- ameter) on this ovigerous specimen ranged from 0.76—-0.96 mm, prior to preservation. Color (from notes and color photographs of live specimens). — Overall whitish opaque to very faint rosy pink; may lack color pat- tern or sometimes have very faint dorsal patterning of pink on carapace, abdominal 349 segments, and uropods; when present, pat- tern usually strongest on posteriormost ab- dominal segments and telson; carapace sometimes with small median square of red- dish pigment just posterior to cervical groove. Chelipeds usually opaque white; sometimes with slight evidence of pale yel- low at articulations of chelipeds and on uro- pods. Known range and habitat.— Known from intertidal burrows in the immediate vicinity of the type locality on the Atlantic coast of Florida, U.S.A. The type locality is a tidally exposed sandflat sparsely vegetated with sea grass, located on the south margin of Fort Pierce Inlet, Florida. A single specimen was taken from a second site a few hundred me- ters distant, on the lower intertidal reaches ofa well-packed sand beach on Coon Island, which forms a northern margin on this same inlet. All of these fossorial specimens were extracted from their burrows with yabby pumps. The burrow of at least one specimen appeared to also harbor a small commensal crab of the genus Pinnixa. Etymology.—This species is named for Mr. John S. MclIlhenny of Avery Island and Baton Rouge, Louisiana. True to tradition in his family, Mr. McIlhenny has melded his passion for Tabasco® pepper sauce with an equal enthusiasm for nature and studies in natural history. The financial support that he has contributed to a number of research biologists through his Coypu Foundation, has furthered research in many subdisci- plines, including crustacean biology. Remarks. — Of the four species of the ge- nus, Eucalliax mcilhennyi, new species, E. Jonesi (Heard 1989) from the Bahamas, and E. cearaensis Rodrigues & Manning, 1992 from Brazil, all lack the acutely projecting armature that characterizes distal corners of the carpus in the chelipeds of E. quadracuta (Biffar 1970) from Venezuela. While each of these corners has typically developed a doublet of spines in E. quadracuta, they are instead weakly produced to a single sub- acute or rounded corner in the other afore- 350 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ) so a \ ae: ip s We e c,d,f a,b Fig. 6. Eucalliax mcilhennyi, new species, from Fort Pierce Inlet, Florida; a—b, d, f, holotype ¢ (CL 10.1 mm) USNM 267112; c, e, paratype 2? (CL 11.6 mm) USNM 267114: a, Right first pleopod of 4, external surface; b, Right first pleopod of 6, internal surface; c, Right first pleopod of 2, external surface; d, Right second pleopod of 4, posterior surface; e, Right second pleopod of 2, posterior surface; f, Right third pleopod, posterior surface. Scale lines indicate 1 mm. VOLUME 107, NUMBER 2 mentioned speices. Materials from Florida that were assigned to E. quadracuta by Bif- far (1970, 1971) and subsequently de- stroyed, as well as some materials that we have collected from Fort Pierce Inlet, also have characteristic “‘quadracuta”’ armature of the chelipeds, but we prefer to recognize them only as members of the “E. quadra- cuta complex” until such time as we can complete a careful comparison with the Venezuelan species. Such caution seems warranted given what appears to be a ten- dency for restricted distribution in species of this genus. As in E. mcilhennyi, the ‘E. quadracuta-complex’ materials from Flor- ida include ovigerous females with large eggs (1.1-1.3 mm), a characteristic that may re- flect an overall tendency toward abbrevi- ated development and reduced larval dis- persal in the genus. This also appears to favor regional endemization in other calli- anassid populations with such eggs (see Felder & Rodrigues 1993). Owing to this possibility, and in the absence of voucher materials, we must reserve judgment on a report of Eucalliax jonesi from the British Virgin Islands (Murphy & Kremer 1992), at least until such time as materials from that locality can be carefully compared to existing types. In addition to the marked difference in cheliped armature, the strong dorsal ridges present on the carapace in the E. quadracuta complex will also serve to readily separate this group from E. mcilhennyi and other members of the genus which lack them. Likewise, in mature individuals, size will readily distinguish E. mcilhennyi and other members of the genus from the E. quad- racuta complex. Biffar (1970) reported total lengths of 68-75 mm for the type materials of E. quadracuta from Venezuela, with a carapace length of 17 mm in the smallest individual which was the mature holotype male. Our materials of this complex from Florida are even larger with total lengths of 85-91 mm and carapace lengths of 21-22 mm. By contrast, the other known members 351 of the genus are much smaller, with cara- pace lengths of mature specimens ranging from 10.1-11.6 mm in E. mcilhennyi, 9.6— 9.8 mm in E. jonesi, and 6.0—7.2 mm in E. cearaensis. Among the many characters that can serve to separate E. mcilhennyi from both E. jo- nesi and E. cearaensis are shape of the ros- trum, armature of the cheliped ischium, rel- ative lengths of the antennular and antennal peduncles, shape of the male first pleopod, and ventral plating of the first and second abdominal somites. Eucalliax cearaensis differs from both the other species in that its antennular and antennal peduncles are subequal in length, and in that the male first pleopod is not bifurcate and instead ter- minates in a single hook. In both E. mcil- hennyi and E. jonesi the antennal peduncles markedly overreach the antennular pedun- cles, and the male first pleopods are bifur- cate along their length, producing a short subapical process. Eucalliax mcilhennyi can in turn be separated from E. jonesi by sev- eral characters that are not usually subject to striking sexual dimorphism in this group, and should therefore apply equally well in males and females (despite the fact that fe- male specimens of E. jonesi have yet to be described). In comparison to E. jonesi, E. mcilhennyi has (1) a much broader and less produced rostrum, (ii) hooked rather than straight spinules lining the ischium of the cheliped, and (i11) a unique pattern of leath- ery plates (Fig. 5a, b, c, d) on ventral sur- faces of the first two abdominal somites. The unique pattern in the ventral integu- ment of the first and second abdominal so- mites in E. mcilhennyi consists not only of the distinct ovoid plates from which the first pleopods originate (Fig. 5a), but also a unique shape in the leathery integument at the posterior margin of the first abdominal somite and the anterior margin of the sec- ond abdominal somite. In E. jonesi, the me- dian posterior margin of the first abdominal somite is marked ventrally by a short, subquadrate extension of the thick leathery 352 covering on the intersegmental joint and second abdominal somite (Fig. 5c); imme- diately posterior to this, the leathery integ- ument is deflected ventrally (Fig. Sd). In E. mceilhennyi, the median extension on the posterior margin of the first abdominal so- mite is anteriorly rounded (Fig. 5a) or sub- triangular in shape, and the leathery integ- ument posterior to this structure is not markedly deflected to the ventral side (Fig. 5b). In addition, mature males of E. mcil- hennyi have first pleopods with an entire rather than bifid subapical lobe and with a more spatulate and laterally directed apical lobe. Our reexamination of the E. jonesi holotype also revealed a prominent proxi- mal boss on the internal surface of the major palm in this male (Fig. Se), and this feature is lacking in both sexes of E. mcilhennyi. In the course of examining morphology in E. mcilhennyi, several features were not- ed which may be of particular significance in phylogenetic placement of the genus. The male second pleopod, previously thought to lack an appendix masculina (see Manning & Felder 1989), does appear to have a large terminal article that may be homologous to structures termed the appendix masculina in various ctenochelid genera. If so inter- preted, the presence or absence of an ap- pendix masculina must be dropped as a character to distinguish members of the Callianassidae from members of the Cten- ochelidae. Also, a uniquely enlarged and elongate, singular seta was found to occur on the internal superodistal corner of the propodus on the fourth pereopod. This unique seta, which is directed distally and overreaches the tip of the dactyl, was also evident in the type materials of E. jonesi (GCRL 1136, USNM 221861) and E. cea- raensis (USNM 252546) that we have sub- sequently examined, and was manifest as a distinct elongate pair of setae in materials of the E. quadracuta complex from Vene- zuela (MCZ 760) and Florida. Given their location, these setae appear to serve a spe- cialized cleaning function. We also note that PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON such a pair of setae occurs in the Mediter- ranean species, Calliax punica (Saint Lau- rent & Manning 1982) (USNM 172356), which represents a genus closely allied to Eucalliax within the Eucalliinae Manning & Felder, 1991. As a character, they may be unique to the Eucalliinae or perhaps may define some larger taxocene within the Cal- lianassidae. Acknowledgments We sincerely thank R. Lemaitre, Smith- sonian Institution, and R. W. Heard, Gulf Coast Research Laboratory (GCRL), who facilitated our access to comparative spec- imen materials. We thank Ardis B. John- ston of the Museum of Comparative Zo- ology, Harvard University (MCZ) for the loan of material. Among many individuals who assisted with or facilitated our field col- lections and observations, we especially thank R. Brown, J. M. Felder, C. King, W. Lee, M. E. Rice, M. Schotte, and J. L. Sta- ton. This study was supported in part under funding to D. L. Felder through Louisiana NSF/EPSCOoR Program grant R11-8820219, Louisiana Education Quality Support Fund grants 86-LUM(2)-084-09 and (1987-88)- ENH-BS-10, US Minerals Management Service Cooperative Agreement 14-35- 0001-30470, and a Coypu Foundation grant. Direct support was also provided through an ongoing program of Smithsonian Marine Station Link Port project grants to R. B. Manning and D. L. Felder. This is contri- bution No. 42 of the USL Center for Crus- tacean Research and contribution No. 350 for the Smithsonian Marine Station. Literature Cited Biffar, T. A. 1970. Three new species of callianassid shrimp (Decapoda, Thalassinidea) from the western Atlantic. — Proceedings of the Biological Society of Washington 83:35—-49. . 1971. The genus Callianassa (Crustacea, De- capoda, Thalassinidea) in South Florida, with VOLUME 107, NUMBER 2 keys to the western Atlantic species. — Bulletin of Marine Science 21(3):637-675. Felder, D. L., & R. B. Manning. 1986. A new genus and two new species of alpheid shrimps (De- capoda: Caridea) from south Florida. — Journal of Crustacean Biology 6:497-508. ——,, & S. de A. Rodrigues. 1993. Reexamination of the ghost shrimp Lepidophthalmus louisi- anensis (Schmitt, 1935) from the northern Gulf of Mexico and comparison to L. siriboia, new species, from Brazil (Decapoda: Thalassinidea: Callianassidae).— Journal of Crustacean Biolo- gy 13:357-376. ——,, J. L. Staton, & S. de A. Rodrigues. 1991. Patterns of endemism in the ghost shrimp genus Lepidophthalmus (Crustacea, Decapoda, Calli- anassidae): evidence from morphology, ecology and allozymes.—American Zoologist 31(5): 101A. Hailstone, T.S., & W. Stephenson. 1961. The biology of Callianassa (Trypaea) australiensis Dana 1852 (Crustacea, Thalassinidea).— University of Queensland Papers, Department of Zoology 1(12):259-285. Heard, R. W. 1989. Calliax jonesi, n. sp. (Decapoda: Thalassinidea: Callianassidae) from the north- western Bahamas.— Gulf Research Reports 8:129-136. Manning, R. B. 1975. Two methods for collecting crustaceans in shallow water.—Crustaceana 29: 317-319. 1987. Notes on western Atlantic Callianas- sidae (Crustacea: Decapoda: Thalassinidea).— Proceedings of the Biological Society of Wash- ington 100:386-401. . 1993. Two new species of Neocallichirus from the Caribbean Sea (Crustacea: Decapoda: Cal- lianassidae). — Proceedings of the Biological So- ciety of Washington 106:106-114. ——, & D. L. Felder. 1986. The status of the cal- lianassid genus Callichirus Stimpson, 1866 (Crustacea: Decapoda: Thalassinidea).—Pro- ceedings of the Biological Society of Washington 99:437-443. —,& 1989. The Pinnixa cristata com- plex in the western Atlantic, with descriptions 353 of two new species (Crustacea, Decapoda, Pin- notheridae).—Smithsonian Contributions to Zoology 474:i-111, 1-26. ,& 1991. Revision of the American Callianassidae (Crustacea: Decapoda: Thalas- sinidea). — Proceedings of the Biological Society of Washington 104:764—-792. ——, & 1992. Gilvossius, a new genus of callianassid shrimp from the eastern United States (Crustacea: Decapoda: Thalassinidea). — Bulletin of Marine Science 49(1-2)[1991]:558- 561. ——, & R. W. Heard. 1986. Additional records for Callianassa rathbunae from Florida and the Ba- hamas (Crustacea: Decapoda: Callianassi- dae). — Proceedings of the Biological Society of Washington 99:347-349. —, & R. Lemaitre. 1994. Sergio, a new genus of ghost shrimp from the Americas (Crustacea: De- capoda: Callianassidae).—Nauplius (Brazil) 1: 39-44. Murphy, R. C., & J. N. Kremer. 1992. Benthic com- munity metabolism and the role of deposit-feed- ing callianassid shrimp. — Journal of Marine Re- search 50:321-340. Rodrigues, S. de A., & R. B. Manning. 1992. Two new cCallianassid shrimps from Brazil (Crusta- cea: Decapoda: Thalassinidea).— Proceedings of the Biological Society of Washington 105:324— 330. Saint Laurent, M. de, & R.B. Manning. 1982. Calliax punica, espéce nouvelle de Callianassidae (Crus- tacea, Decapoda) des eaux méditerranéennes. — Quaderni di Laboratorio di Tecnologia della Pesca, Ancona 3(2-5):211-—224. (DLF) Department of Biology and Center for Crustacean Research, University of Southwestern Louisiana, Lafayette, Loui- siana 70504, U.S.A.; (RBM) Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560, U.S.A. PROC. BIOL. SOC. WASH. 107(2), 1994, pp. 354-359 PINNIXA SCAMIT, A NEW SPECIES OF PINNOTHERID CRAB (DECAPODA: BRACHYURA) FROM THE CONTINENTAL SLOPE OFF CALIFORNIA Joel W. Martin and Deborah L. Zmarzly Abstract.—A new species of the pinnotherid crab genus Pinnixa, P. scamit, is described. Collected off the coast of Pt. Arguello, California, U.S.A., at 311 m, the new species belongs to the Pinnixa occidentalis complex, species of which are characterized by a highly sculptured carapace, well developed cardiac ridge, and deflexed fixed fingers on the chelae. Pinnixa scamit differs from P. occidentalis in having a well developed but granular (rather than acute) cardiac ridge, and larger, more acute, slightly curved teeth along the anterolateral mar- gin of the carapace. The most salient difference between the two species is the length of the propodus of the third walking leg (pereiopod 4), which is at least 2.5 times its width in the new species. In a recent review of crabs in the genus Pinnixa White, 1846, known from Califor- nia, Zmarzly (1992) redescribed 11 previ- ously recognized species and described two new species, P. forficulimanus and P. mi- nuscula, bringing the number of species of this genus known from California waters to 13. Several of the species treated by Zmarzly were described as exhibiting significant morphological variation. In particular, Zmarzly (1992) reiterated Hart’s (1982) comment that the species Pinnixa occiden- talis Rathbun, 1893, in part because of its variability and in part because of its wide geographic and bathymetric distribution, may represent a species complex rather than a single morphologically variable species. While examining several decapod crus- tacean specimens collected as part of a fau- nal survey of the Santa Maria Basin and western Santa Barbara Channel, conducted by the Minerals Management Service (MMS) of the U.S. Department of the In- terior, we noticed two specimens that shared certain features with Pinnixa occidentalis Rathbun, 1893, but that did not entirely agree with the original description or Zmarzly’s (1992) redescription of this spe- cies. Subsequent examination and compar- ison with true P. occidentalis has convinced us that the differences are sufficient to war- rant the erection ofa new species of Pinnixa, which is described below. The holotype and sole paratype are deposited in the National Museum of Natural History, Smithsonian Institution, Washington, D.C. (USNM). Pinnixa scamit, new species Figs. 1; 2 Material examined.—Holotype female, USNM 267500, carapace width 7.4 mm, carapace length 3.7 mm; 29 Nov 1983; MMS station data 071-BSS-01-TX, 34°29.040’N, 120°44.013'W, western Santa Barbara Channel, just seaward of, and SSW of, Pt. Arguello, California; single core replicate; 1020 feet (311 m); 1 mm screen. Paratype juvenile, USNM 267501 (sex indetermi- nate), carapace width 3.4 mm, carapace length 2.1 mm, same collecting data as for holotype. Diagnosis. —Carapace highly sculptured, with anterolateral ridge bearing pronounced and slightly anteriorly-curved teeth; frontal margin with deep median cleft; cardiac ridge VOLUME 107, NUMBER 2 Fig. 1. Pinnixa scamit, female holotype, USNM 267500, and juvenile paratype, USNM 267501 (c only), western Santa Barbara Channel, California. a, female holotype, dorsal view of carapace and right side appendages, pereiopods 1-3 drawn in situ, pereiopods 4 and 5 removed and figured separately (to assure correct proportions); b, same specimen, frontal view showing sculpturing of carapace and subhepatic tooth (arrow); c, dorsal view of immature paratype; note that subhepatic tooth (arrow) and acute serrations along anterolateral margin are already present at this size; d, third maxilliped of holotype, left side, outer view; e, abdomen of holotype, ventral view illustrated in situ (first two segments not visible, and segment 3 shortened due to curvature of abdomen). Scale bar = 2.0 mm for a—, e; 1.0 mm for d. 355 356 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. Pinnixa scamit, female holotype, USNM 267500, chelipeds. a, right (minor) cheliped, outer view; b, left (major) cheliped, outer view, with fingers enlarged below (arrow) to show minute teeth along cutting border; length of merus and basi-ischium distorted (overly shortened) in both a and b because of orientation of illustration; c, merus of left cheliped, dorsal view, with carpus toward top of figure. Scale bar = 1.0 mm for all figures except for enlargement of b (which is not to scale). present, granular, not sharp. Fixed finger or major cheliped slightly deflexed; both fin- gers of major cheliped with single stout tooth located almost at midlength on inner mar- gin. All ambulatory pereiopods with mark- edly serrate and setose dorsal and especially ventral margins. Propodus of pereiopod 4 at least 2.5 times longer than wide. Description of holotype. —Carapace (Fig. la, b) two times wider than long, surface with numerous granules and scattered, short setae. Carapace highly sculptured, with are- olations distinct and defined by deep grooves. Frontal margin with deep median cleft. Anterolateral margins broadly arcu- ate, with pronounced ridge bearing well de- veloped acute teeth, each tooth curved slightly to anterior. Subhepatic region of carapace just lateral to orbits bearing small but obvious tooth (arrow, Fig. 1a; more pro- nounced in juvenile paratype, Fig. 1c). Gas- tric depression deep. Cardiac ridge well de- veloped but granular, obtuse, not acute, slightly bilobed. Chelipeds (Figs. 1a, 2) slightly dimorphic, left larger. Fixed finger slightly deflexed, nearly 2 length of palm. Opposing borders of dactylus and fixed finger of each cheliped with pronounced tooth at approximately midlength and with row of small transpar- ent teeth merging distally into thin sharp ridge along cutting edges (Fig. 2b). Palm with scattered short setae and granules on outer surface and serrate ridge on dorsal border. Carpus and merus (Fig. 2c) with well de- veloped acute teeth on dorsolateral and dor- sodistal surfaces. Ambulatory pereiopods (Fig. la, where ambulatory pereiopods are denoted by P2- PS) long and relatively slender compared to those in most other species of genus. Each leg with well developed row of teeth along VOLUME 107, NUMBER 2 dorsal and ventral borders of merus, carpus, and propodus, less obvious on carpus. Dac- tylus of each ambulatory leg more or less straight, slightly bent to flexor side on pe- reiopods 2 through 4, to extensor side on pereiopod 5. All articles with combination of scattered short plumose and simple setae. Pereiopod 4 (third walking leg) longest, with propodus 2.6 times longer than wide. Prop- odus of pereiopod 4 with ventral margin bicarinate, each carina serrate. Dactylus of pereiopod 4 slightly longer than propodus. Tip of dactylus on each walking leg not strongly curved. Third maxilliped as figured (Fig. 1d), typ- ical of genus. Endopod with short distally plumose setae on merus and carpus and with long smooth or sparsely setulose setae on distal half of dactylus and propodus; distal border of carpus with thick, brush-like tuft of setae. Proximal article of exopod with obvious lateral protrusion at approximately midlength and plumose setae along lateral border. Abdomen (Fig. le) broadly rounded, well developed, setose, consisting of 7 free seg- ments, only distalmost 5 visible in ventral view (Fig. le). Pleopods mature, well de- veloped, and setose. Male unknown. Juvenile paratype (Fig. 1c).—Similar to holotype in the serrate anterolateral border and possession of subhepatic tooth. Am- bulatory pereiopods also similar to those of holotype. Paratype differs from holotype in having a slightly reduced carapace width: length ratio (a difference that we attribute to normal ontogenetic changes), a slightly more produced front, and a slightly more pronounced subhepatic tooth relative to carapace size (arrow, Fig. Ic). Type locality. —Western Santa Barbara Channel, SSW of Pt. Arguello, California, 34°29.040'N, 120°44.013’W, 311 m, soft bottom. Distribution. —Known only from the type locality. Etymology. —The species name origi- 357 nates from an acronym being used as a noun in apposition. We are pleased to take the name of this species from SCAMIT, the ac- ronym for the Southern California Associ- ation of Marine Invertebrate Taxonomists, a largely unheralded organization of pro- fessional biologists that has done much to further our knowledge of marine inverte- brates in southern California. Remarks.—Most California species of Pinnixa White, 1846, have a ridge, which may be granular to tuberculate, along the anterolateral margin of the carapace. Ex- ceptions include P. faba (Dana 1851), P. longipes (Lockington 1876), P. tubicola Holmes, 1894, and P. weymouthi Rathbun, 1918, where an anterolateral ridge is either absent or at best very weakly developed; and P. forficulimanus Zmarzly, 1992, P. minuscula Zmarzly, 1992, and P. littoralis Holmes, 1894, where it is absent (Zmarzly 1992). In the remaining California species of Pinnixa, this ridge is obvious in dorsal view, and is sometimes as pronounced as in P. scamit. However, in none of the adults of these species are the individual antero- lateral teeth as sharp or as anteriorly curved as in P. scamit. Juvenile stages of two spe- cies, P. occidentalis Rathbun, 1893, and P. franciscana Rathbun, 1918, sometimes ex- hibit acute teeth on the anterolateral ridge; however, this feature changes dramatically between the juvenile and adult stages. In addition, P. scamit has relatively slender legs with acute marginal serrations as an adult, which is also characteristic only of juvenile stages in P. occidentalis and P. fran- ciscana. Thus P. scamit appears to have re- tained several ‘“‘juvenile’”’ characters in the adult stage. All other California species are easily dis- tinguished from P. scamit by the length: width ratio of the propodus of pereiopod 4. In true P. occidentalis Rathbun, the spe- cies to which P. scamit appears to be most closely allied, the propodus of P4 is dis- tinctly broader, nearly as broad as long. In contrast, in P. scamit the propodus of pe- 358 reiopod 4 is approximately 2.6 times longer than wide. The geographic range is of little help in determining species affinities; the collecting locality of P. scamit is within the range of nearly all of the 13 California species of Pinnixa listed by Zmarzly (1992, fig. 1), in- cluding the wide ranging Pinnixa occiden- talis. It is unfortunate that we do not have a male specimen, as gonopod morphology would undoubtedly help clarify the rela- tionship between P. scamit and other Cal- ifornia species. The species also shares cer- tain morphological similarities with Pinnixa affinis Rathbun, 1898, known only from 26 fm (47.5 m) in Panama Bay (see Rathbun 1918: 168), but can be distinguished by the more deflexed major chela in P. affinis (see Rathbun 1918: fig. 106). The extent to which some systematically important morphological characters vary in California species of Pinnixa is not known. A detailed knowledge of such characters as the morphology of the orbits, eyes, epi- stome, antennae, and mouthparts, appar- ently useful in resolving placement within pinnotherid species complexes in other geo- graphic areas, is at present lacking. In-depth study of these features may resolve some of the questions of relationships within the P. occidentalis complex. If the key to California species of Pinnixa given by Zmarzly (1992: 678) is employed, then the new species would key to P. occi- dentalis. For separation of the species, we suggest insertion of the following couplet, to replace couplet 8 in Zmarzly’s key. 8. Fixed finger of chela straight or curving upward, not deflexed ... 9 (as in original key) — Fixed finger of chela deflexed, an- gled obliquely downward relative to line delineated by ventral mar- gin of palm (deflection stronger in males than in females) ......... 8A. Length of propodus of pereiopod 8A PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 4 at least 2.5 times width ...... LE aoa terry tee ate P. scamit, new species — Length of propodus of pereiopod 4 approximately 1.5—2.0 times width P. occidentalis Rathbun, 1893 Acknowledgments This manuscript resulted from a 1993 meeting of the Southern California Asso- ciation of Marine Invertebrate Taxono- mists (SCAMIT). We are grateful to the members of that organization for their as- sistance and for their consistent dedication to marine invertebrate systematics. We also thank Dr. George C. Steyskal for his in- valuable help and advice concerning our choice of a specific epithet. Literature Cited Dana, J.D. 1851. Conspectus crustaceorum quae in orbis terrarum circumnavigatione, Carolo Wilkes e classe Reipublicae Faederate duce, lexit et des- cripsit.— Proceedings of the Academy of Natu- ral Sciences, Philadelphia 5:247—254. Hart, J. F.L. 1982. Crabsand their relatives of British Columbia.— British Columbia Provincial Mu- seum, Victoria, British Columbia, Canada. Pp. 1-267. Holmes, S. J. 1894. Notes on West American Crus- tacea.— Proceedings of the California Academy of Sciences, series 2, 4:563-588. Lockington, W. N. 1876. Description of a new genus and species of decapod crustacean.—Proceed- ings of the California Academy of Sciences, 1877, series 1, 7:55—57. Rathbun, M. J. 1893. Scientific results of explora- tions by the U.S. Fish Commission steamer A/- batross. XXIV. Descriptions of new genera and species of crabs from the west coast of North America and the Sandwich Islands.— Proceed- ings of the United States National Museum 16: 223-260. 1898. The Brachyura collected by the U.S. Fish Commission steamer A/batross on the voy- age from Norfolk, Virginia, to San Francisco, California, 1887—1888.—Proceedings of the United States National Museum 21:567-616. . 1918. The grapsoid crabs of America. — Unit- ed States National Museum Bulletin 97:1—445. White, A. 1846. Notes on four new genera of Crus- tacea.—Annals and Magazine of Natural His- tory, 18(118):176-178, pl. 2. VOLUME 107, NUMBER 2 Zmarzly, D. L. 1992. Taxonomic review of pea crabs in the genus Pinnixa (Decapoda: Brachyura: Pinnotheridae) occurring on the California shelf, with descriptions of two new species. — Journal of Crustacean Biology 12:677-713. (JWM) Natural History Museum of Los Angeles County, 900 Exposition Boulevard, 359 Los Angeles, California 90007, U.S.A.; (DLZ) Stephen Birch Aquarium-Museum, Scripps Institution of Oceanography, Uni- versity of California, San Diego 0207, La Jolla, California 92093, U.S.A. PROC. BIOL. SOC. WASH. 107(2), 1994, pp. 360-382 STUDIES OF NEOTROPICAL CADDISFLIES, L: THE DESCRIPTION OF CERASMATRICHIA, NEW GENUS, A RELATIVE OF ALISOTRICHIA, WITH DESCRIPTIONS OF NEW AND OLD SPECIES AND THE LARVA (TRICHOPTERA: HYDROPTILIDAE) Oliver S. Flint, Jr., Steven C. Harris, and L. Botosaneanu Abstract. —Cerasmatrichia, new genus (type species C. trinitatis), is estab- lished and defined in the adult and larval stage. Alisotrichia dominicensis Flint, A. wirthi Flint and A. adunca Flint are transferred thereto, and their males and females redescribed. Cerasmatrichia spinosa (Venezuela), C. argylensis (To- bago, Trinidad), C. trinitatis (Trinidad), are newly described in the male and female sexes; the larva of the first is firmly established by metamorphotypes, and therefore used as the basis for the generic level description of the larva (modified if necessary to include the larvae of species A—-E). Larvae and females of an additional six species serve to extend the range of the genus to Costa Rica, south to Peru, east through Venezuela and north in the Lesser Antilles to Guadeloupe. Since the genus Alisotrichia was described in 1964 for a Puerto Rican species, the genus has grown to 32 species and become very heterogeneous. As part of studies by the first two authors on the definition and limits of the tribe Stactobiini in the New World, we have been attempting to define natural groups in this melange, and when warrant- ed, establish new genera for them (e.g., Har- ris & Holzenthal 1993, Harris & Bueno 1994). One of the most obvious groups cen- ters around A. dominicensis Flint, which 1s the focus of this study. The recent discovery of several undescribed species by the third author, and the firm association of the larval stage has confirmed the distinctiveness of this assemblage of species. We are here es- tablishing the new genus Cerasmatrichia, redescribing three species transferred there- to, describing three newly discovered spe- cies, and the larval stage. Material is deposited in several museums as indicated by the following acronyms: IRSNB, Institut Royal des Sciences Natu- relles de Belgique, Brussels; NMNH, Na- tional Museum of Natural History, Wash- ington D.C.; IZAM, Instituto de Zoologia Agricola, Maracay, Venezuela; UWI, De- partment of Zoology, University of the West Indies, St. Augustine, Trinidad; ZMUA, Zoologisch Museum, Universiteit van Am- sterdam, Amsterdam. Genus Cerasmatrichia, New Genus Type Species. — Cerasmatrichia trinitatis new species. Adult (Figs. 1-14).—Of moderate size, forewing length 2-3 mm; color generally fuscous, sometimes with indistinct whitish maculae. Head with 3 ocelli. Antennae sim- ple, scape elongate, with 25—35 segments in male, 22—26 in female. Maxillary palpus with 5 segments: 2 basal segments very small, globular, 3 remaining segments of equal length, each 3—4 times as long as basal VOLUME 107, NUMBER 2 361 SPE me ext A 4 ie 0 % YS Kk US SC © “ Q Figs. 1-7. 1-4, Cerasmatrichia trinitatis, adult. 1, head and thorax, dorsal; 2, fore femur and tibia; 3, mid femur and tibia; 4, hind femur and tibia. 5-7, C. wirthi, adult. 5, metanotum, dorsal; 6, maxillary palpus; 7, fore femur and tibia. 362 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Figs. 8-14. 8-10, Cerasmatrichia trinitatis, wings. 8, forewing venation, male; 9, hindwing venation, male; 10, forewing venation, female. 11, C. wirthi, male forewing schematic showing modified areas. 12, same of C. dominicensis. 13, same of C. argylensis. 14, same of C. trinitatis. VOLUME 107, NUMBER 2 2; labial palpus 3-segmented. Tentorial bridge divided mesally. Wings with reduced venation; forewing with small, pointed bas- al lobe, CuP (?) with pale, fracture-like line bordering its anterior side in both male and female; forewing in male with leathery, modified posterobasal and often costal ar- eas bearing specialized setae; hindwing nar- row, attenuate. Mesoscutellum with trans- verse suture; metascutellum with anterior margin strongly convex, truncate laterally. Spur on foreleg very short and lightly scler- otized, smaller in female than male (often unobservable except under highest magni- fication); 3 spurs on midleg; 4 on hindleg. Seventh sternum in most males with short, fat lobe; females rarely with small point on sixth sternum. Males have eighth sternum large, projecting posteroventrally, with small spines posteroventrally; tergum smaller, roughly quadrate. Ninth segment produced anterolaterally, thus mostly open dorsally and ventrally, sometimes with ventrolateral lobate process. Tenth tergum variously formed, with paired lateral sclerites, spines, or single mesal process. Inferior appendages usually divided into two arms: one more slender and arising and curving ventrolat- erally, other usually broader and directed dorsomesally. Subgenital plate strong, pro- jecting posteriad. Phallus with narrow basal section and broader, tubular apical portion; with rather simple, internal, tubular ejacu- latory duct. Female genitalia with simple, conical seventh sternum; tergum produced into pair of posterolateral, darkened, hirsute lobes. Eighth segment simple, tubular, rare- ly somewhat ornamented, or with postero- lateral lobes dorsally, with long, slender an- terolateral apodemes and ring of stout setae along posterior margin. Ninth segment slen- der, elongate, sclerotized laterally, with long, slender, anterolateral apodemes. Tenth seg- ment small, lightly sclerotized, narrowed apically with pair of slender apicolateral pa- pillae. Vaginal sclerites complex, with small, spherical sclerite pierced by central opening, sometimes with strong anterior plate with 363 projecting teeth, often with only membra- nous sacs variously sclerotized in certain species. Larva (Figs. 15—24).— Length, 3.5—5 mm. Head pale brown, unpigmented around eye and ventrally, slightly elongate, coronal su- ture weak, other sutures lacking. Head with several setae quite prominent, especially an- teriorly; antenna elongate. Labrum about twice as wide as long, with anterolateral membranous lobe, densely hairy, 2 pair of major setae clearly visible dorsally. Man- dibles short, broad, slightly concave api- comesally with 2 or 3 weak cusps; each with median brush of several long setae. Pro- notum divided longitudinally, meso- and metanota entire; all with many small, dark points; bearing numerous, enlarged setae arising from large pale cuticular spots. Meso- and metanota each with small, secondary sclerites along posterior margin and region of rugosities along anterior margin. Meso- and metapleura each with 2 of these large setae arising from short protuberance; pro- pleuron with only posterior seta, trochantin small, pale, conical with small seta. All legs short, stout, virtually identical; middorsal seta of each femur enlarged, feathered; dor- sal surfaces of femora and tibiae roughened (in some species each point is seen to bear small seta); claw without evident seta. First to ninth abdominal terga with large scler- ites, those on one to eight each with trans- verse, central, dark area and several, large, dark setae. Row of small sclerites between tergites and row of rugosities along posterior margin. First to eighth segments each with conical lobe laterally capped by small scler- ite bearing pair of large setae, one dark, oth- er smaller and pale; ninth tergum with large shield-like sclerite bearing number of setae. Anal prolegs short, each with several slen- der setae and large apicomesal, pale blade; claw hooked sharply ventrad. Distribution. —Examples are known from about 10°N in Costa Rica south along the mountains to 13°S in Peru, and east across northern South America to south-central 364 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Figs. 15-16. 15, Cerasmatrichia species B, larva, dorsal. 16, C. spinosa, pupa and case, ventral. VOLUME 107, NUMBER 2 365 Figs. 17-24. Cerasmatrichia spinosa, larva. 17, foreleg; 18, midleg; 19, hindleg; 20, head; 21, mandibles; 22, labrum; 23, eighth and ninth terga; 24, anal proleg. 366 Venezuela (Puerto Ayacucho), and north along the Lesser Antilles to Guadeloupe. Biology.—All species are rheophilous, with larvae found either near the waterline on boulders in fast flowing rivers and small streams or on the rocks in madicolous hab- itats. Adults have rarely been taken at ul- traviolet light at night in large numbers, usually they are taken in rather small num- bers at light or by net in the day. The larvae are free-living until near pu- pation time. The pupae and their shelters of the two available collections are nearly identical. There is a thin, dorsal, sheetlike covering, beneath which (next to the sub- strate) is an oval, firmly spun cocoon within which is found the pupa (Fig. 16). Strangely, the dorsal covering is pierced both anteri- orly and posteriorly by a large circular open- ing; this dorsal sheet is thus like the covering of the final instar larvae of the Leucotri- chiini. The inner cocoon is attached along the middorsal line to the dorsal sheet, ex- cept at the head end where it is broadly attached. When the pharate adult emerges it uses its mandibles to cut a circular open- ing dorsally, the silken flap is left attached by a small strap left uncut on the posterior side. Remarks. —A preliminary cladistic anal- ysis of the species originally placed in A/i- sotrichia is presented by Harris & Holzen- thal (1993). In it the dominicensis group was clearly defined and separated from the other recognizable groups in Alisotrichia s.1. The retention in the group of the plesiomorphic state of tibial spur present on foreleg serves to distinguish the group from the sister lin- eage in which this spur is lost. As shown herein, this spur is greatly reduced and well on the way to total loss even in Cerasma- trichia. However, they did propose 3 syn- apomorphies to define the group which still seem valid: female seventh tergum (mistak- enly called VIII tergum) with paired setal patches, male eight sternum with patch of heavy spine posteromesally, and ejacula- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON tory duct of phallus protruding ventrally. Probably many of the larval structures men- tioned above are synapomorphic, but it would be premature to infer phylogeny based on them while evidence is still lacking from larvae of many other groups. However, con- sidering what is known of this stage in other species, it is suggested that some (or all) of the following are likely larval synapomor- phies: elongate, fusiform shape; the pres- ence of enlarged, swordlike setae on the tho- rax and abdomen; a large, dorsal, fimbriate seta on each femur; and the large, conical lateral protuberances on the abdomen which are capped by a sclerite bearing two large setae. In addition, the presence of 3 ocelli, unmodified antennae, spur count of 1 (but virtually lost), 3, 4, and modified forewing structure in the males will help differentiate the species from various other groups in Alisotrichia s.1. Warning should be given concerning some of the differences apparent in the illustra- tions of the female genitalia. The seventh tergum appears to show small differences in shape between the various species. The comparative widths and lengths are greatly dependant on the degree of flattening, ro- tation and orientation of the specimen. It is presently deemed unwise to use this seg- ment alone to identify the species. Likewise the apparent differences in the transverse, rake-like bar of C. wirthi and C. adunca are due to orientation. This plate can be arti- ficially rotated in either species to produce the aspect shown in the figure of the other. Key to Described Species 1. Males with process from seventh sternum, hooked in lateral aspect (Fig. 25). Female vaginal apparatus consisting of a spherical sclerite and possibly other sclerites more pos- terior (Fig. 40) (dominicensis group) Males lacking process from seventh NO VOLUME 107, NUMBER 2 367 Figs. 25-29. Cerasmatrichia spinosa, male genitalia. 25, lateral; 26, ventral; 27, dorsal; 28, phallus, lateral: 29, phallus, dorsal. 368 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON sternum (Fig. 57). Female vaginal apparatus consisting of spherical sclerite and a more anterior, trans- verse, rake-like sclerite (Fig. 40) (wirthi group) 2. Male tenth tergal area without spines, with a long, central strap-like sclerite (Fig. 32). Female vagina with only a spherical sclerite (Fig. 37). . ens A PRE Cate ra he dominicensis Male tenth tergal area bearing either a pair of divergent pointed processes (Fig. 43) or clusters of spines (Fig. 27). Female vagina with a spherical sclerite (Fig. 56), and often a more posterior, complex sclerite (Fig. 40) 3 3. Male tenth tergum consisting of a pair of divergent, pointed processes (Fig. 43). Female vagina with only a spherical sclerite (Fig. 56) argylensis Male tenth tergum with many spine- like processes (Fig. 27). Female va- gina with both spherical sclerite and a more posterior sclerite (Fig. 40) 4 4. Male tenth tergum basolaterally with a cluster of short spines (Fig. 27); eighth sternum laterally with many, enlarged setae (Figs. 25, 26, 27). Fe- male eighth sternum lightly sclero- tized and longitudinally striate (Fig. SiS) RRR esc? ae alive ae oR Pea Ye spinosa Male tenth tergum consisting of 6 pairs of long, slender spines (Fig. 48). Female eighth sternum with ven- tral, lateral and dorsal sclerotized areas distinct and not striate (Fig. iD aml etcabay eee! ele tas trinitatis 5. Male tenth tergum consisting of pair of diverging, tapering sclerites whose apices end in sharp point (Fig. 57). Female indistinguishable from fol- lowing species wirthi Male tenth tergum consisting of pair of broadly oval sclerites whose api- ces are narrowed and compressed (Fig. 63). Female similar to preced- NEC NSNES gonasscvacnepocevca adunca Cerasmatrichia spinosa, new species Figs. 16-29, 38-40 Alisotrichia wirthi Flint.—Flint, 1981:26 [misidentification, in part]. This is a readily identifiable species in both the male and female. The male of no other species bears a cluster of short, fat setae basolaterally on the tenth tergum, which is then continued as a pair of lateral, rodlike plates, nor does any other species have the cluster of enlarged setae dorsolat- erally from the eighth sternum. The female genitalia show a lightly striate surface to the eighth segment, and the posterior vaginal apparatus is distinctively sclerotized. Adult. —Length of forewing 2.3 mm. Col- oration in alcohol, mottled brown (probably mottled fuscous in life). Male with apparent sexual modifications: 3rd segment of max- illary palpus bearing modified setae mesal- ly; fore coxa with patch of modified setae basolaterally, femur slightly inflated basal- ly, otherwise unmodified; forewing with large basal pocket and marginal cell of scale- like hairs (as in Fig. 8). Male seventh ster- num with strong process; female sixth ster- num bearing small point. Male genitalia. Eighth sternum with dorsolateral corner produced as large angulate lobe and bearing cluster of long, enlarged spines, mesally pro- duced with cluster of small spines. Ninth segment widely open dorsally and ventrally, strongly produced anterolaterally as round- ed lobe. Tenth tergum developed on each side as rounded, convoluted basolateral lobe bearing cluster of short, very broad setae from which cluster arises elongate rodlike sclerite with slightly twisted tip. Inferior ap- pendage with large lateral, lightly sclero- tized plate (or possibly this is an extension of lateral margin of ninth segment); elongate and broad mesal plate with apex curved slightly ventrad, with small basal, membra- nous lobe. Subgenital plate long, arising ventrolaterally from basal area of tenth ter- gum, tip hooked ventrad, appearing as dark- VOLUME 107, NUMBER 2 ened ovoid in ventral aspect. Phallus with tubular basal portion, and enlarged, more membranous, apical portion separated by sharp constriction; apical portion with cen- tral ejaculatory and some basal, internal sclerites. Female genitalia. Eighth segment lightly sclerotized, surface striate; posterior row of stout setae. Vagina with anterior spherical sclerite; posterior area with elon- gate lightly sclerotized region bearing more heavily sclerotized lateral pockets. Larva. —Length, 4 mm. Head pale brown, unpigmented around eye and ventrally. Pronotum with enlarged setae: anterior margin bearing row of 5-6 each side, behind margin submesal pair, at midlength sub- mesal pair and 3 laterally, with scattered, small, dark points. Meso- and metanota with enlarged setae: anterior margin with 3 each side, midlength with 2 each side, with scat- tered small, dark points and point free areas. Dorsal surfaces of femora and tibiae rough- ened without evident seta. First to ninth abdominal terga with large sclerites, those on one to eight each with transverse, central, dark area; first tergum with 3, large, dark setae each side, second through eighth terga with 4 such setae (one extra seta in lateral cluster) each side. Row of small sclerites between tergites and row of rugosities along posterior margin of all segments. Ninth ter- gum with large shield-like sclerite bearing 3, enlarged, dark setae and 3 slender, point- ed setae, each side. Material examined. —Venezuela, Edo. Aragua, Rio El Limon, fish hatchery, Ma- racay, 16 Feb 1976, C. M. & O. S. Flint, Jr., é holotype, 2 allotype, 13 6, 1 2 paratypes (all metamorphotypes), 51 prepupae, pupae and empty cases, 78 larvae (NMNH); same data, except 19-20 May 1975, F. H. Wei- bezahn, | 6 paratype (IZAM). Cerasmatrichia dominicensis (Flint), new combination Figs. 12, 30-37 369 Alisotrichia dominicensis Flint, 1968:44; 1970:29.—Botosaneanu, 1989:97: 1990: 44 [Martinique]. Ochrotrichia (O.) species. — Flint & Sykora, 1993:58 [misidentification, Guadeloupe]. This is a readily identifiable species in the male. The male of no other species bears a single dorsomesal, sclerotized strap on the tenth tergum, which appears to be con- nected laterally into the subgenital plate which is directed straight down. The female, however, offers no clear-cut differences from that of C. argylensis. Adult. —Length of forewing, 2 mm. Color fuscous; antennae and fore and mid tarsi cream colored, head anteriorly and meso- notum with white hair, forewings with white maculae basally and at midlength. Male with minor sexual modifications: 3rd segment of maxillary palpus seemingly unmodified; fore femur slightly inflated basally, otherwise unmodified; forewing with small midbasal area modified and bulging, costal area from base to midlength with membrane thick- ened. Male seventh sternum with strong process; female sixth sternum with small point. Male genitalia. Eighth sternum with posterolateral margin produced as large rounded lobe, posteromesally with cluster of small spines. Ninth segment widely open dorsally and ventrally, strongly produced anterolaterally as narrowly angled lobe. Tenth tergum developed as broad, lightly sclerotized region basally, abruptly narrow- ing into long, parallel-sided, heavily scler- otized, middorsal band with apex slightly hooked dorsad. Inferior appendage with lat- eral arm short, arising basolaterally from elongate mesal plate which is angulate api- cally; mesal arm twice as long as lateral arm, tapered to blunt apex in lateral aspect, slightly capitate and lobed in ventral aspect. Subgenital plate long, slender, arising ven- trolaterally from basal area of tenth tergum, apex directed ventrad, tip appearing as darkened ovoid in ventral aspect. Phallus 370 with tubular basal portion, and enlarged, more membranous apical portion separated by sharp constriction; apical portion with central ejaculatory and some basal, internal sclerites. Female genitalia. Eighth segment lightly sclerotized ventrally, unornamented; posterior margin with row of stout setae. Vagina with anterior spherical sclerite with small central opening; posterior area com- pletely membranous. Material examined.—Dominica: 2.2 mi E Pont Casse, 7 May 1964, O. S. Flint, Jr., 1 éholotype (NMNH). 2.5 mi E Pont Casse, 16 Jan 1965, W. W. Wirth, 3 2 allotype and paratypes (NMNH). Fond Figues, 25 Jan 1965, W. W. Wirth, | 6 paratype (NMNH). Morne Nicholls, 9 Nov 1964, P. J. Spangler, 1 2 paratype (NMNH). Martinique: Riviére Coco at Morne Vert, 14 Feb 1986, L. Bo- tosaneanu, 1 6 (ZMUA). Ravine 1’Abbé, Morne Vert, 3 Mar 1989, L. Botosaneanu, light, 1 ¢(ZMUA). Guadeloupe: Basse Terre, Cascade aux Ecrevilles, 9 Apr 1992, L. Bo- tosaneanu, | 6 (ZMUA). Riviére Laurant, near Belleville, 8 Apr 1979, StarmuhlIner & Therezien, 1 2 (NMNH). Cerasmatrichia argylensis, new species Figs. 13, 41-45, 54-56 Hydroptilid genus, sp. 2, Botosaneanu & Sakal, 1992:201. The species is readily recognized in the male sex by the tenth tergum developed into a pair of strong, divergent hooks. The fe- male, however, can not be distinguished with absolute certainty from that of C. domini- censis. Adult. —Length of forewing, 2 mm. Color of female fuscous; antennae and fore and mid tarsi cream colored, head with white hair anteriorly, mesonotum and forewings with indistinct, silvery-white maculae. Male with sexual modifications: 3rd segment of maxillary palpus slightly cupped, with con- cave face modified and bearing specialized setae; fore femur inflated basally, integu- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ment modified and bearing specialized se- tae; forewing with midbasal area modified and strongly bulging, costal area from base to midlength with membrane thickened. Male seventh sternum with strong process; female sixth sternum bearing small point. Male genitalia. Eighth sternum with pos- terolateral margin produced as large round- ed lobe, posteromesally with cluster of small spines. Ninth segment widely open dorsally and ventrally, strongly produced anterolat- erally as narrowly rounded lobe. Tenth ter- gum developed as pair of heavily sclerotized hooks with apices directed laterad. Inferior appendage with lateral arm terete, directed laterad; mesal arm bilobed and mitten-like in ventral aspect; setate lobe ventrally at junction of two arms. Subgenital plate long, slender, tip enlarged, decurved in lateral as- pect, bilobed in ventral aspect. Phallus with tubular basal portion, and enlarged, more membranous apical portion separated by sharp constriction; apical portion with cen- tral ejaculatory duct extending through more sclerotized cavity. Female genitalia. Eighth segment lightly sclerotized ventrally, unor- namented; posterior row of stout setae. Va- gina with anterior spherical sclerite appear- ing transversely divided at equator; posterior area completely membranous. Material examined.—Tobago: [St. Paul Parish], Argyle River at Argyle Waterfall, 22 Apr 1991, L. Botosaneanu, Mary Alkins- Koo, M. Koo, at light: 6 holotype, 2 allotype, 1 6 paratype (ZMUA), 1 ¢ paratype (NMNB). [St. John Parish], 6.5 km N Rox- borough [on Parlatuvier Rd.], B1/5 (11°17'N, 60°35’W), 15-16 Jun 1993, O.S. Flint, Jr., UV light, 1 6 paratype (NMNH); same, but 14 Jun 1993, O. S. Flint & W.N. Mathis, by net, 1 2 paratype (NMNH). 3 km S Charlotteville, WASA intake stream (11°19'N, 60°33’W), 125 m, O. S. Flint & N. E. Adams, UV light, 1 2 paratype (NMNH). Trinidad: [St. George County], Northern Range, below Maracas Waterfall (10°44'N, 61°24'W), 250 m, 18 Jun 1993, N. E. Adams & W.N. Mathis, 1 2(NMNBH). VOLUME 107, NUMBER 2 371 31 Figs. 30-34. Cerasmatrichia dominicensis, male genitalia. 30, lateral; 31, ventral; 32, dorsal; 33, phallus, lateral; 34, phallus, dorsal. 372 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON P) 5 ¢ PED Pee rare ae aero ters Figs. 35-40. 35-37, Cerasmatrichia dominicensis, female genitalia. 35, ventral; 36, seventh tergum, dorsal; 37, vaginal sclerites, ventral. 38-40, C. spinosa, female genitalia. 38, ventral; 39, seventh tergum, dorsal; 40, vaginal sclerites, ventral. VOLUME 107, NUMBER 2 373 Figs. 41-45. Cerasmatrichia argylensis, male genitalia. 41, lateral; 42, ventral; 43, dorsal; 44, phallus, lateral; 45, phallus, dorsal. 374 Cerasmatrichia trinitatis, new species Figs. 1-4, 8-10, 14, 46-53 Hydroptilid genus, sp. 1, Botosaneanu & Sakal, 1992:201. The species is readily recognized in the male sex by the very spinose tenth tergum, perhaps C. argylensis with its single pair of spines in this area is its closest relative. The female is also readily recognized as no other species has such a highly ornamented eighth segment, nor such a strongly sclerotized and complex posterior portion of the vaginal re- gion. Adult. — Length of forewing, 2.5 mm. Col- or fuscous; antennae and fore and mid tarsi cream colored, head with white hair ante- riorly, mesonotum and forewings with in- distinct, silvery-white maculae; female pal- er overall. Male with sexual modifications: 3rd segment of maxillary palpus slightly cupped, with concave face modified and bearing specialized setae; fore femur inflat- ed basally, integument modified and bear- ing specialized setae; forewing with mid- basal area modified and strongly bulging, costal area with membrane thickened. Male seventh sternum bearing strong process; fe- male sixth bearing small point. Male geni- talia. Eighth sternum with posterolateral margin produced as large rounded lobe, posteromesally with cluster of small spines. Ninth segment widely open dorsally and ventrally, strongly produced anterolaterally as narrowly rounded lobe. Tenth tergum greatly modified, bearing cluster of heavily sclerotized spines basally: 3 pairs mesally, of which ventralmost much the longest, lat- eral spine long, twisted laterally, then ven- trally and mesally, around a shorter spine curved laterad; thin, lightly sclerotized plate beneath these spines and arched dorsome- sally. Inferior appendage with lateral arm terete, directed laterad; mesal arm broad in ventral aspect with distinct apicomesal point, small setate lobe ventrally at junction of two arms. Subgenital plate long, slender, tip distinctly narrowed. Phallus with tubu- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON lar basal portion, and enlarged, more mem- branous apical portion separated by sharp constriction; apical portion with central ejaculatory duct and few basal sclerites. Fe- male genitalia. Eighth segment with large, bilobed, rugose ventral sclerotized area, di- vided anteromesally, paired dorsolateral, sclerotized pockets, and pair of oval, dorsal plates; posterior row of stout setae with dor- solateral portion arising from narrow, dark- ened sclerite. Vagina with anterior spherical sclerite and strongly sclerotized posterior complex, in ventral aspect, with sclerotized lateral surface, sclerotized, projecting, mid- ventral ridge, and small, transverse anterior plate. Material examined. —Trinidad: [St. George County], Northern Range, Maracas Waterfall (10°44’N, 61°24’W), 270 m, 12 Apr 1991, L. Botosaneanu & D. Sakal, at light: 6 holotype, 2 allotype (ZMUA), 25 4, 27 2 paratypes, (ZMUA,NMNH, UWI, IRSNB); same, but 18 Jun 1993, O. S. Flint, Jr., 44 6, 112 2 paratypes (ZMUA, NMNH, UWI); same, but below falls, 250 m, N. E. Adams and W.N. Mathis, 1 6, 3 2 paratypes (NMNH). Northern Range, streamlet at “La Laja” (10°43’'N, 61°17’W), Guanapo River catchment, 520 m, N. E. Adams & W. N. Mathis, UV light, 2 2 paratypes (NMNH); same, but Ist order stream, 14 Apr 1991, L. Botosaneanu & D. Sakal, at light, 4 4, 1 ? paratypes (ZMUA); same, but 2nd order stream, 13 Apr 1991, 3 6, 1 2 paratypes (ZMUA). North Coast Road, hygropetric habitat, west from Maracas Bay, 30 Apr 1991, L. Botosaneanu & D. Sakal, at light, 1 6, 5 2 paratypes (ZMUA). Blue Basin Wa- terfall, 17 Apr 1991, L. Botosaneanu & D. Sakal, at light, 23 2 (ZMUA, NMNH). Cerasmatrichia wirthi (Flint), new combination Figs. 5-7, 11, 57-62, 68-70 Alisotrichia wirthi Flint, 1968:46; 1970:29: 1981:26 [in part]. VOLUME 107, NUMBER 2 375 Figs. 46-50. Cerasmatrichia trinitatis, male genitalia. 46, lateral; 47, ventral; 48, dorsal; 49, phallus, lateral; 50, phallus, dorsal. 376 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON \ Mi i u W, i Figs. 51-56. 51-53, Cerasmatrichia trinitatis, female genitalia. 51, ventral; 52, seventh tergum, dorsal; 53, vaginal sclerites, ventral. 54-56, C. argylensis, female genitalia. 54, ventral; 55, seventh tergum, dorsal; 56, vaginal sclerites, ventral. VOLUME 107, NUMBER 2 This species is quite similar to its sister, C. adunca, in the male and not apparently separable in the female. The two are most easily distinguished in the male by the form of the tenth tergites. In C. wirthi they are rather slender and divergent in dorsal aspect with a sharp, upturned tip, but in C. adunca they are broad, tapering apicad to a thin, vertical plate that appears as a dark line. Adult.—Length of forewing, 2-3 mm. Color fuscous; antennae pale brown, fore tarsi and tibia, and mid tarsi cream colored; forewings almost uniformly fuscous. Male with sexual modifications: 3rd segment of maxillary palpus with integument very dark and bearing specialized setae; fore femur unmodified; forewing with bulging, modi- fied midbasal area, costal area for halflength of wing with membrane thickened. Male seventh sternum without process, with nar- row, transverse, projecting carina; female sixth sternum lacking point. Male genitalia. Eighth tergum and sternum almost com- pletely fused into a cylinder; posteroventral margin almost truncate, produced into pair of small, submesal lobes, with cluster of small spines. Ninth segment widely open dorsally and ventrally, produced ventrolat- erally as small lobe over base of its inferior appendage. Tenth tergum developed as pair of elongate, divergent sclerites with pointed, upturned tip. Inferior appendage simple, elongate structure, curved ventrad and en- larged apically with single large seta dorsally at midlength; with elongate, oblique setose lobe basally. Subgenital plate as long as in- ferior appendages, tip hooked ventrad, ap- pearing as darkened ovoid in ventral aspect. Phallus with tubular basal portion, and en- larged, apical portion separated by sharp constriction; apical portion with outer sur- face strongly sclerotized (and frequently de- taching from phallus if phallus is removed), centrally with ejaculatory duct and some basal, internal sclerites. Female genitalia. Eighth segment lightly sclerotized laterally; posterior row of stout setae. Vagina with spherical sclerite, anteriad to which is 377 strongly sclerotized, transverse bar bearing many long teeth, comblike. Material examined. —Dominica: Fond Figues River, 13 Mar 1965, W. W. Wirth, 6 holotype; same data, but 6 Apr 1964, O. S. Flint, Jr., 1 6 paratype (NMNH). Pont Casse, 1.5 mi N, 12 Feb 1965, W. W. Wirth, 1 2(NMNH). Venezuela: Edo. Aragua, Dos Riitos, 6 km N Rancho Grande, 4 Feb 1976, C. M. & O. S. Flint, Jr., 1 ¢ (NMNH). Es- tacion Experimental Cataurito, ca. 32 km E Villa de Cura, 1100 m, 1 Feb 1983, O. S. Flint, Jr., 4 6, 1 2 (NMNH). Edo. Miranda, Santa Cruz de Rio Grande, Parque Gua- topo, 7 Feb 1976, C. M. & O. S. Flint, Jr., 1 6 (NMNB). Cerasmatrichia adunca (Flint), new combination Figs. 63-67, 71-73 Alisotrichia adunca Flint, 1991:44. As mentioned under C. wirthi, these two species are quite similar and perhaps not separable in the female. The form of the male tenth tergites are the most distinctive. In C. wirthi they are rather slender and di- vergent in dorsal aspect with a sharp, up- turned tip, but in C. adunca they are broad, tapering apicad to a thin, vertical plate that appears as a dark line. As in C. wirthi, the phallus of the male has a large shield-like sclerite surrounding the apical portion. This shield mimics the ““penis-sheath”’ of the Leucotrichiini and has been a source of confusion. It was clearly shown in the original description of this spe- cies (Flint 1991, figs. 106-108) as a large, apically rounded plate in lateral aspect. Careful examination shows it to be distinct from what is here called the subgenital plate, although they are easily confounded as in his figs. 107, 108. This plate often detaches from the phallus if one tries to pull the phal- lus back through the abdomen (as happened in Fig. 63 herein, where it is shown as the large, rectanguloid structure within seg- ments 8 and 9). 378 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 62 Figs. 57-62. Cerasmatrichia wirthi, male genitalia. 57, lateral; 58, ventral; 59, dorsal; 60, seventh and eighth sterna, ventral; 61, phallus, lateral; 62, phallus, dorsal. Adult. — Length of forewing, 2.5mm.Col- tions: 3rd segment of maxillary palpus and or fuscous; frontal hairs of head, antennae, fore femur apparently unmodified; forewing fore tarsi and tibia, and mid tarsi cream with membrane of midbasal area thickened, colored; forewings fuscous with many small, slightly bulging, but apparently not other- pale maculae. Male with sexual modifica- wise modified, costal area for half length of VOLUME 107, NUMBER 2 379 Figs. 63-67. Cerasmatrichia adunca, male genitalia. 63, lateral; 64, ventral; 65, dorsal; 66, phallus, lateral; 67, phallus, dorsal. 380 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON wing with membrane thickened. Male sev- enth sternum without process, with long, narrow, transverse, projecting carina; fe- male sixth sternum lacking point. Male gen- italia. Eighth tergum and sternum distinct; posteroventral margin almost truncate, pro- duced into pair of small, submesal lobes, with cluster of small spines. Ninth segment widely open dorsally and ventrally, sharply produced anterolaterally. Tenth tergum de- veloped as pair of broad, ovoid, lateral sclerites each with thin, vertical apex ap- pearing as dark mark in dorsal aspect. In- ferior appendage single, elongate structure, curved ventrad and slightly enlarged api- cally with large seta dorsally at midlength; with elongate, oblique setose lobe basally. Subgenital plate slightly shorter than infe- rior appendages, tip hooked ventrad, ap- pearing as darkened point in lateral aspect. Phallus with tubular basal portion, and en- larged, apical portion separated by sharp constriction; apical portion with outer sur- face strongly sclerotized (detached from phallus and shown internally in Figs. 63 and 65, herein), centrally with ejaculatory duct and some basal, internal sclerites. Female genitalia. Eighth segment lightly sclerotized laterally; posterior marginal row of stout se- tae. Vagina with spherical sclerite, anteriad to which is strongly sclerotized transverse bar bearing many long teeth, comblike. Material examined.—Colombia: Dpto. Antioquia, 10 km E Medellin, road to Guarne, 7 Feb 1983, O. S. Flint, Jr., ¢ ho- lotype (NMNH). Quebrada El Pozo, 8 km W EI Penol, 9 Feb 1983, O. S. Flint, Jr., 1 6, 1 2 paratypes (NMNH). Cerasmatrichia species We here record a series of collections, lar- vae or females, that can not be definitively placed to species. They serve to expand the known range of the genus and give some idea of specific differences to be found in the larvae. Species A.—This collection of pupae and prepupae contains several nearly mature metamorphotypes, but not in good enough condition to permit description. They do represent an undescribed species of the C. dominicensis group. The larva bears an ex- tra row of enlarged setae along the anterior margins of the pro-, meso- and metanota, the points dorsally on the femora and tibiae are very large and clearly setate, and the membranous surface of the abdomen dor- sally is covered with dark spicules. Material: Venezuela, Edo. Merida, Mucujun Valley, 11 km NE Merida, 21 Feb 1976, C. M. & O. S. Flint, Jr., 1 6, 4 2 metamorphotypes, 13 pupae, 2 prepupae, 2 empty cases (NMNB). Species B.—These are the larvae shown in Fig. 15. They probably are those of C. dominicensis, whose adults were taken close to larger rivers such as those from which these larvae came. Material: Dominica, Laurent River [at Bells], 21 Jul 1963, O. S. Flint, Jr., 1 larva (NMNH). Geneva Estate, 9 Dec 1964, Paul J. Spangler, 1 larva (NMNH). Fond Figues River, 9 Feb 1965, W. W. Wirth, rocks in river, 1 larva (NMNH). Martinique, Riviere Blanche, near Alma, 19 Apr 1979, Starmihlner, 2 larvae (ZMUA). Species C.—This single larva is damaged in the prothoracic region and is somewhat shriveled and thus not fully distended, therefore telescoping the segments. It ap- pears to be virtually identical to the larva of Species B although the enlarged setae seem exceptionally long and dark. Material: Cos- ta Rica, Cartago Prov., Reserva Tapanti, Rio Dos Amigos, ca. 6 km (road) NW tun- nel, 9.704°N, 83.783°W, elev. 1500 m, 9- 10 Jun 1988, C. M. & O. S. Flint, Jr., Hol- zenthal, 1 larva (NMNH). Species D.—This larva agrees, again, with the basic structure, but its entire dorsal sur- face (sclerites and membrane) is covered with dark points, the points on the tibiae are produced into a comblike row of clear projections, and the abdominal terga lack the posterior band of rugosities. Material: VOLUME 107, NUMBER 2 381 73 Figs. 68-73. 68-70, Cerasmatrichia wirthi, female genitalia. 68, ventral; 69, seventh tergum, dorsal; 70, vaginal sclerites, ventral. 71-73, C. adunca, female genitalia. 71, ventral; 72, seventh tergum, dorsal; 73, vaginal sclerites, ventral. Peru, Dept. Cuzco, San Pedro (at km 152 & 2 km east), 13°09’S, 71°26’W, 1430 m, 31 Aug 1989, R. A. Faitoute (colln. 11), 1 larva (NMNB),. Species E.—This is another larva that is very similar to those of C. spinosa. It differs in lacking the posterior bands of rugosities on the abdominal terga, but has the mem- brane darkened and with dark points later- ad, and the femora and tibiae seem smooth dorsally. Material: Venezuela, T.F. Ama- zonas, 40 km S Puerto Ayacucho at To- 382 bogan, 24 Feb 1986, leaf packs, P. J. Span- gler (colln. 12), 1 larva (NMNBH). Species F.—A single female, unassociated with any male has been found in the col- lections. The genitalia are identical to the species of the C. wirthi group. Material: Ec- uador, Tungurahua Prov., 13 km E Banos, 15 Sep 1990, O. S. Flint, Jr., 1 2 (NMNH). Acknowledgments We thank Dr. Mary Alkins-Koo for help in making local arrangements, suggesting collecting sites on Trinidad and Tobago, and providing access to the collections at the University of the West Indies. The Research Opportunities Fund of the Smithsonian In- stitution provided funding for O. Flint to collect on Trinidad and Tobago in 1993. The trips of L. Botosaneanu to Martinique (1986, 1989) were made possible by grants from the Netherlands Royal Academy of Sciences (K.N.A.W.) and Treub-Maat- schappij, whereas his trip to Trinidad and Tobago (1991) was made with financial help from the Netherlands Foundation for the Advancement of Tropical Research (W.O.T.R.O.). Fig. 16 was prepared by Young T. Sohn and Fig. 15 by the late André D. Pizzini, to both of whom we are most grateful. Literature Cited Botosaneanu, L. 1989. Seconde contribution al’étude des Trichoptéres de la Martinique. — Annales de la Société entomologique de France (Nouvelle Série), 25:95-104. . 1990. Results ofa trichopterological (Insecta: Trichoptera) travel to the Lesser Antilles in 1989.—Bulletin de l’Institut Royal des Sciences Naturelles de Belgique, Entomologie 60:39—48. —., & M. Alkins-Koo. 1993. The caddis flies (In- secta: Trichoptera) of Trinidad and Tobago, West Indies.—Bulletin de l'Institut Royal des Sci- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ences Naturelles de Belgique, Entomologie 63: 5-45. ——,, & D. Sakal. 1992. Ecological observations on the caddisflies (Insecta: Trichoptera) from Trin- idad and Tobago (W. Indies). — Revue d’Hydro- biologie Tropicale 25:197—207. Flint, O. S., Jr. 1968. Bredin-Archbold-Smithsonian biological survey of Dominica, 9. The Trichop- tera (caddisflies) of the Lesser Antilles.—Pro- ceedings of the United States National Museum 125(3665):1-86. 1970. Studies of neotropical caddisflies, X: Leucotrichia and related genera from North and Central America (Trichoptera; Hydroptili- dae).—Smithsonian Contributions to Zoology 60:1-64. 1981. Studies of neotropical caddisflies, XXVIII: The Trichoptera of the Rio Limon Ba- sin, Venezuela.— Smithsonian Contributions to Zoology 330:1-61. . 1991. Studies of neotropical caddisflies, XLV: The taxonomy, phenology, and faunistics of the Trichoptera of Antioquia, Colombia.—Smith- sonian Contributions to Zoology 520:1-113. ——., & J. L. Sykora. 1993. New species and re- cords of caddisflies (Insecta: Trichoptera) from the Lesser Antilles, with special reference to Grenada.— Annals of Carnegie Museum 62:47— 62. Harris, S. C., & J. Bueno-Soria. 1994. Scelobotrichia, a new genus of microcaddisflies from Mexico (Trichoptera: Hydroptilidae).—Folia Entomo- logica Mexicana (in press). , & R. W. Holzenthal. 1993. Phylogeny of the species groups of Alisotrichia, Sensu Lato, with the description of a new species from Costa Rica (Trichoptera: Hydroptilidae). Pp. 155—160 in C. Otto, ed., Proceedings of the Seventh Interna- tional Symposium on Trichoptera. Backhuys Publishers, Leiden, 312 pp. (OSF) Department of Entomology MRC 105, National Museum of Natural History, Washington, D.C. 20560, U.S.A.; (SCH) Department of Biology, Clarion University, Clarion, Pennsylvania 16214, U.S.A.; (LB) Instituut voor Taxonomische Zodlogie, Universiteit van Amsterdam, Plantage Middenlaan 64, 1018 DH Amsterdam, The Netherlands. PROC. BIOL. SOC. WASH. 107(2), 1994, pp. 383-390 A NEW HEMICHORDATE, SACCOGLOSSUS BROMOPHENOLOSUS (HEMICHORDATA: ENTEROPNEUSTA: HARRIMANIIDAE), FROM NORTH AMERICA Gary M. King, Cem Giray, and Irv Kornfield Abstract. — A new species, S. bromophenolosus, is distinguished from its con- gener, S. kowalevskii (Agassiz 1873), on the basis of the following morpholog- ical, biochemical, and genetic criteria: placement of gill pores; prominence of the dorsal ridge; structure of the proboscis skeleton; presence of bromophenols or bromopyrroles; relative electrophoretic mobility of allozymes (e.g., super- oxide dismutase); molecular weights of fragments from restriction endonuclease digestion of mitochondrial DNA. S. bromophenolosus occurs from southern Maine, U.S.A., to Nova Scotia, Canada, with an additional record from Willapa Bay, Washington, U.S.A. Three families of enteropneusts (Hem1- chordata: Enteropneusta: Harrimaniidae, Ptychoderidae, and Spengeliidae) occur along the coasts of North America (Milne & Milne 1973, Bullock 1975, Ruppert & Fox 1988). The best known species belong to the genus Saccoglossus (Harrimaniidae), which occurs on both the Atlantic and Pa- cific coasts (Milne & Milne 1973, Bullock 1975). Of the several saccoglossids, S. ko- waleyskii (Agassiz 1873) has been described in greatest detail. Various aspects of its dis- tribution, biology and ecology have been reported subsequent to Agassiz’s (1873) ini- tial description (e.g., Bateson 1886, Bullock 1940, Tweedell 1961, Colwin & Colwin 1962, Barrington 1965, King 1986, Woodin et al. 1987, Balser & Ruppert 1990, Carey & Mayer 1990). S. kowalevskii has been considered the only member of its genus on the Atlantic coast of North America, and has been noted in intertidal collections from Nova Scotia to Georgia (Dorjes 1972, Bromley 1979). Although there are certain characteristics common to all saccoglossids along this range, a comparison of biochem- ical attributes reported for populations from Maine and South Carolina has raised ques- tions about the taxonomic status of the spe- cies (King 1986, Woodin et al. 1987). Spe- cifically, animals collected in Maine and northward typically accumulate high con- centrations of two secondary products, 2,4- dibromophenol and a dibromoindole (King 1986, unpubl. data), while animals from other locales accumulate 2,3,4-tribromo- pyrrole (King, unpubl. data, Woodin et al. 1987). The differential accumulation of these haloaromatics does not appear to correlate with any major environmental variables or gradients. Further, the presence of a given haloaromatic phenotype appears to be a fixed trait associated with populations from specific regions. We now report that the genus Saccoglos- sus consists of at least two distinct species along the Atlantic coast of North America. These two species are readily differentiated by gross external morphological characters, the accumulation of halogenated aromatic compounds, morphology of the proboscis skeleton, electrophoretic mobilities of sev- eral enzymes, and the sequence composi- tion of the mitochondrial DNAs (mtDNA). We propose the name S. bromophenolosus for the new species occurring from southern 384 Maine northward and possessing features as detailed below. S. kowalevskii occurs from southern Maine southward, as noted by Verrill (1873) and Gosner (1979). Materials and Methods Saccoglossids were collected from the in- tertidal zones of sites ranging from Halifax, Nova Scotia, to Georgetown, South Caro- lina, by excavating sediments to a depth of about 10 cm at low tide. The proboscis, collar and anterior portions of the trunk were obtained readily by removing the animals directly from the substrate, but complete specimens were difficult to collect due to the fragility of the posterior region of the trunk. Specimens were examined in the field for external morphological characteristics (see below) and then placed individually in small vials (20 ml) containing local seawater; the animals were subsequently transported to Walpole, Maine, for further processing. MtDNA was extracted from live animals using procedures modified from Lansman et al. (1981) within 24—48 h of return to the laboratory. A minimum of 10 live animals from Lowes Cove and York, Maine (69°34’N, 43°56'W; 43°09'N, 70°39’W) and from Portsmouth, New MHampshire (43°06'N, 70°42'W) were also homogenized for electrophoretic analyses of enzymes at the following loci using standard methods (Murphy et al. 1990): glucose phosphate isomerase (GPI; E.C. 5.3.1.9), malate de- hydrogenase (MDH; E.C. 1.1.1.37), phos- phoglucomutase (PGM; E.C. 2.7.5.1), and superoxide dismutase (SOD; E.C. 1.15.1.1). Analyses of haloorganic contents utilized extracts of animals that were extracted with hexane immediately after collection and re- moval of adhering sediment. In addition, animals used for mtDNA and allozyme analyses were “‘halotyped”’ by collecting 1— 2-ul samples of the seawater immediately adjacent to the proboscis with a 10-l gas chromatography syringe (Hamilton Inc., Reno, Nevada). The epidermis of the pro- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON boscides was irritated with the syringe nee- dle prior to sample collection in order to stimulate mucus and haloorganic excretion. The samples were analyzed by direct injec- tion into a gas chromatograph. Details of the extraction and analytical procedures have been reported elsewhere (King 1986, 1988). The proboscideal skeletons of 5 in- dividuals from Lowes Cove and York, Maine, were examined after dissection of live animals. Saccoglossus bromophenolosus, new species Figs. 1-4, Table 1 Saccoglossus sp. Bullock 1975, p. 619; Koz- loff 1987, p. 478. Saccoglossus kowalevskii.— Milne & Milne 1973, (p.p.): 230; Bromley 1979, p. 533. Saccoglossus kowalewskyi.—Brinkhurst et al., 1976, p. 156. Saccoglossus kowalevskyi.— King, 1986, p. QDS. Saccoglossus kowalewskii. — Linkletter et al. 1977, (p.p.): 42, Gosner 1979, p. 265; Meinkoth, 1981, (p.p.): 726; Carey & Mayer, 1990, p. 79. Diagnosis. —Saccoglossus bromopheno- losus attains a length up to 20 cm. It has an elongate proboscis that extends 1.5—2 cm in narcotized specimens. The proboscis has a shallow dorsal groove and a single pore at the base of the proboscis. Concretions fill the primary shaft of the proboscis skeleton. Hexane extracts of the proboscis contain 2,4- dibromophenol and a dibromoindole; these compounds account for the characteristic ‘““‘bromoform” odor of live specimens. The collar is differentiated into 4-5 zones, and is generally rectangular from above with an aspect ratio of about 3:5 (width : length). The trunk is differentiated into distinct bran- chial and hepatic-genital regions. Dorsolat- erally placed gill pores appear ellipsoid in relaxed and ventilating specimens, with the major axis oriented vertically; the gill pores do not occur in folds and vary from about VOLUME 107, NUMBER 2 60-110 pair per individual. Esophageal pores, which occur slightly posterior to the gill pores vary from about 4-8 pair in num- ber. Gonads begin at the terminus of the branchial region in both sexes, and are dor- solateral in placement. Material examined. —Holotype (USNM 168049) and 3 paratypes (USNM 168050- 168052) from Lowes Cove, Maine (69°34’N, 43°56’W), collected by G.M. King 20 Oc- tober, 1993. Description. —The holotype is a sexually immature female with the following char- acteristics observed while the animal was relaxed in a solution of 7% MgCl,. The in- complete specimen was 82 mm in length; an additional 60-70 mm of the posterior- most region of the trunk was lost unavoid- ably during collection. The posterior-most section of the trunk was extremely thin and fragile; in color and diameter, it resembled the fecal coils present at the sediment sur- face. The relaxed and extended proboscis was creamy white in color with a shallow dorsal groove running from the base to the proboscis tip. Striations perpendicular to the major axis of the proboscis were observed at low power on a dissecting microscope. The proboscis was 16 mm in length and 3 mm at the base. The basal-most region was a rust orange in color. The orange collar was 5 mm in length and 2.5 and 3 mm in di- ameter at the anterior and posterior ends, respectively. Both ends of the collar were distinctly thickened, with a prominent light- er-colored ridge circling the posterior of the collar. The branchial region of the trunk was about 2.5 mm in diameter where it joined the posterior of the collar. A prominent, raised dorsal ridge began immediately pos- terior to the collar and ran the length of the branchial region, terminating just anterior to the esophageal pores. Near the collar, the dorsal ridge was about 1 mm in width with a central groove about 0.5 mm in width. This region of the trunk was pinkish-orange to orange and about 14 mm in length. A total of 61 pairs of gill pores were located 385 dorsolaterally, beginning at the junction of collar and trunk. Slightly posterior to the terminal gill pores, 4 pairs of dorsal esoph- ageal pores were observed; they formed an angle of about 30° opening toward the trunk. The hepatic-genital region of the trunk ex- tended about 23 mm, measured from the termination of the gill pores to a point where genital structures were not observable. Na- scent grayish egg masses were visible dor- solaterally just beneath the epidermis. The hepatic-genital region was grayish-brown dorsally and yellowish-brown laterally. The posterior-most region of the trunk was 34 mm in length, yellowish-brown in color and characterized by pairs of carmine-colored spots located dorsally. General dimensions for 3 male paratypes with maturing gonads were similar to those of the holotype. However, gill pore numbers ranged between 68-101 and esophageal pores numbered 4—6. The sperm sacs were dorsolateral and pink in color. Remarks. — Although the range of body size, coloration, location and appearance of the gonads, and external features of the col- lar and proboscis of S. bromophenolosus and S. kowaleyskii are very similar, gill pore placement and the morphology of the dorsal ridge allow discrimination of the species by visual inspection in the laboratory or field (Fig. 1, 2). In contrast to S. bromopheno- losus, the dorsal ridge of S. kowalevskii is relatively broad and flattened immediately posterior to the collar, covering >50% of the dorsal area. The dorsolateral gill pores of S. kowalevskii occur within lateral grooves that can be closed, thereby obscuring them from view (Fig. 2). This state is typical of specimens preserved without prior relaxa- tion. In addition to external morphology, S. bromophenolosus and S. kowalevskii can be distinguished by other diagnostic charac- teristics requiring dissection or various lab- oratory analyses (Fig. 3, Table 1). Collec- tions to date indicate that S. kowalevskii and S. bromophenolosus exist sympatrically only within a narrow range around the mouth of PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. A. Dorsolateral view of relaxed, sexually immature female of S. bromophenolosus, new species illustrating superficial features of the proboscis (P), collar (C, limits indicated by arrows) and trunk (T); gill pores (gp, indicated by arrow) are evident immediately posterior to the collar. B. Detailed view of the posterior proboscis, collar, and anterior of the trunk, illustrating the prominent, raised dorsal ridge (DR) and gill pores. VOLUME 107, NUMBER 2 387 Fig. 2. Anterior portion of S. kowalevskii showing the broad, flattened dorsal ridge (DR), collar (C) and proboscis (P); gill pores are obscured by lateral folds (F, indicated by arrow). The “sole” (S) occurs ventrally. the Piscataqua River, New Hampshire (43°06’N, 70°42'W): S. kowalevskii has not been observed north of York, Maine, while S. bromophenolosus has not been found south of Portsmouth, New Hampshire. Allozyme analyses indicated substantial divergence between S. bromophenolosus and S. kowalevskii since GPI, MDH, and PGM were fixed for alternative, diagnostic alleles within each taxon; SOD was represented at 2 monomorphic loci in S. bromophenolo- sus, and by a single, different monomorphic locus in S. kowalevskii. MtDNA analyses also revealed substantial divergence (Fig. 4). Table 1.—Diagnostic characters that distinguish among 3 species of North Atlantic saccoglossids. S. kowalevskii S. ruber SS. bromophenolosus Characteristic Dorsal ridge raised, narrow Gill pore placement central Coloration! (proboscis-collar-trunk) W/P-O/R-O Halotype? DBP/DBI Proboscis skeleton? narrow curve-co broad, flattened raised, narrow lateral central W/P-O/R-O P/R/R-O TBPy DBP/TBP broad curve-no co narrow curve-co ' Color code: W/R, ranges from white to pale pink; O/R, ranges from orange to red; O, orange; P/R, pale pink to pale red. ? Compound code: DBP = 2,4-dibromophenol; DBI = dibromoindole (positions of bromine atoms uncertain); TBP = 2,4,6-tribromophenol; TBPy = 2,3,4-tribromopyrrole. 3 Co refers to concretions within the proboscis skeleton. 388 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ' if i Fig. 4. Autoradiogram of fragments from the di- gestion of pooled mtDNA from specimens of S. bromo- phenolosus, new species (lanes 1, 3) or S. kowalevskii (lanes 2, 4) with the restriction endonucleases Hind III (lanes 1, 2) or Sty I (lanes 3, 4). Fragments were sep- arated after digestion by agarose gel electrophoresis and radiolabelled with *°S using a nick translation proce- dure. Lane 5 contains a molecular weight standard (1 kb ladder, BRL); “‘a” and “b” indicate 5090 and 1018 base pair fragments, respectively. — Fig. 3. A. Diagram of the proboscideal skeleton of S. bromophenolosus, new species; dark central region consists of concretions as described for S. ruber (Bur- don-Jones & Patil 1960). B. Diagram of the proboscis skeleton of S. Kowalevskii; note absence of central con- cretions and more deeply curved skeletal arms. VOLUME 107, NUMBER 2 No common DNA fragments were found in comparisons of restriction endonuclease di- gests of the mtDNA of the two taxa based on Hind Ill, XbaI, NdeI, Sty 1, Stu 1, Dra I or Ava I. In contrast, mtDNA haplotypes for populations of each taxon were homog- enous. For example, an Xba I digest yielded the following approximate fragment molec- ular weights: S. bromophenolosus—8750, 1075, 925, and 625, S. kowalevskii— 5000, 3225, 2275, 1875, and 1000. Etymology.—The species name is de- rived from its characteristic haloorganic content, 2,4-dibromophenol, and the Latin suffix,-osus; thus S. bromophenolosus, Sac- coglossus “‘with bromophenol.” Distribution. —On the east coast of North America, the range of S. bromophenolosus extends north from the mouth of the Pis- cataqua River separating Maine and New Hampshire to at least Halifax, Nova Scotia. S. bromophenolosus is found in silty sands throughout the intertidal zone in this range. It is often distributed in patches, with den- sities from about 10—> 100 individuals m ~?. In addition, a sub-tidal form has been re- corded from the Damariscotta River (69°34'N, 43°56’W) at a depth of 10-20 m. Specimens obtained from Willapa Bay, Washington (46°37’N, 124°00’W) have very similar mtDNA haplotypes and 16S ribo- somal DNA sequences (pers. observations); in addition, the external morphologies and haloorganic contents of these organisms are indistinguishable from S. bromophenolosus (K. Woodwick, in litt.). Since these north- western Pacific saccoglossids represent a previously unnamed species (Kozloff 1987), we incorporate them as trans-Arctic repre- sentatives of S. bromophenolosus. The range of the western North American populations is uncertain, but appears to include Oregon and Washington (Bullock 1975, Kozloff 1987). Acknowledgments GMkK dedicates this manuscript to the memory of H. W. King. We thank Dr. J. 389 Grant for assistance with collections from Nova Scotia; Drs. E. Ruppert and D. Hub- bard for information about and assistance with collections from Georgetown, South Carolina; A. Laursen and Drs. C. Plante and G. Kozloff for information about and col- lections of specimens from Willapa Bay, Washington. This work was partially sup- ported by funds from the National Science Foundation, OCE-9203342 and EHR91- 08766. Literature Cited Agassiz, A. 1873. The history of balanoglossus and tornaria.— Memoirs of the American Academy of Arts Science 1X:421—429. Balser, E. J., & E. E. Ruppert. 1990. Structure, ul- trastructure, and function of the preoral heart- kidney in Saccoglossus kowalevskii (Hemichor- data, Enteropneusta) including new data on the stomochord.—Acta Zoologica 71:235-249. Barrington, E. J. W. 1965. The biology of hemi- chordata and protochordata. W. J. Freeman Co., San Francisco. 176 pp. Bateson, W. 1886. Continued account of the later stages in the development of Balanoglossus ko- walevskii, and of the morphology of the Enter- opneusta.—Quarterly Journal of Microscopical Science 26:511-533. Brinkhurst, R. O., L. E. Linkletter, E. I. Lord, G. A. Conners, & M. J. Dadswell. 1976. A prelim- inary guide to the littoral and subtidal marine invertebrates of Passamaquoddy Bay. Hunts- man Marine Laboratory. St. Andrews, N.B. 166 pp. Bromley, J. E. C. 1979. A preliminary checklist of marine fauna of Minas Basic and Minas Chan- nel.— Proceedings of the Nova Scotia Institute of Science 29:517-541. Bullock, T. H. 1940. The functional organization of the nervous system of Enteropneusta. — Biolog- ical Bulletin 79:91-113. 1975. Phylum Hemichordata. Pp. 618-619 in R. I. Smith & J. T. Carlton, eds., Light’s manual: intertidal invertebrates of the Central California Coast, 3rd ed. University of Califor- nia Press, Berkeley, 716 pp. Burdon-Jones, C. 1951. Observations on the spawn- ing behaviour of Saccoglossus horsti Brambell and Goodhart, and of other Enteropneusta. — Journal of the Marine Biological Association of the United Kingdom 29:625-638. ——, & A.M. Patil. 1960. A revision of the genus Saccoglossus (Enteropneusta) in British wa- 390 ters. — Proceedings of the Zoological Society of London 134:635-645. Carey, D. A., & L. M. Mayer. 1990. Nutrient uptake by a deposit-feeding enteropneust: nitrogenous sources. — Marine Ecology Progress Series 63: 79-84. Colwin, L. H., & A. H. Colwin. 1962. Induction of spawning in Saccoglossus kowaleyskii (Enter- opneusta) at Woods Hole.— Biological Bulletin 123:493. Dorjes, J. 1972. Georgia coastal region, Sapelo Is- land, U.S.A.: sedimentology and biology. VII. distribution and zonation of macrobenthic an- imals.—Senckenbergiana maritima 4:169-182. Gosner, K. L. 1979. A field guide to the Atlantic seashore. Houghton Mifflin Co., Boston, 329 pp. King, G. M. 1986. Inhibition of microbial activity in marine sediments by a bromophenol from a hemichordate. — Nature 323:257-259. 1988. Dehalogenation in marine sediments containing natural sources of halophenols.— Applied and Environmental Microbiology 54: 3079-3085. Kozloff, E. N. 1987. Marine invertebrates of the Pa- cific Northwest. University of Washington Press, Seattle, Washington, 511 pp. Lansman, R. A., R. O. Shade, J. F. Shapiro, & J. C. Avise. 1981. The use of restriction endonu- cleases to measure mitochondrial DNA se- quence relatedness in natural populations. III. Techniques and potential applications. —Jour- nal of Molecular Evolution 17:214—226. Linkletter, L. E., E. I. Lord, & M. J. Dadswell. 1977. A checklist of marine fauna and flora of the Bay of Fundy. Huntsman Marine Laboratory. St. Stevens, New Brunswick, 68 pp. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Meinkoth, N. A. 1981. The Audubon Society field guide to North American seashore creatures. A.A. Knopf, New York, 813 pp. Milne, L. & M. Milne. 1973. Invertebrates of North America. Doubleday, Inc., New York, 249 pp. Murphy, R. W., J. W. Sites, Jr., D. G. Buth, & C. H. Hanfler. 1990. Proteins I: isozyme electro- phoresis. Pp. 45-126 in D. M. Hillis & C. Mo- ritz, eds., Molecular Systematics. Sinauer As- sociates, Inc., Sunderland, Massachusetts, 588 pp. Ruppert, E. E., & R. S. Fox. 1988. Seashore animals of the southeast. University of South Carolina Press, Columbia, 429 pp. Tweedell, K. S. 1961. Regeneration of the enterop- neust, Saccoglossus kowalevskii. — Biological Bulletin 120:118-127. Woodin, S. A., M. D. Walla, & D. E. Lincoln. 1987. Occurrence of brominated compounds in soft- bottom benthic organisms.—Journal of Exper- imental Marine Biology and Ecology 107:209- 217. Verrill, A. E. 1873. Explorations of Casco Bay by the U.S. Fish. Comm. B. Natural History. Proceed- ings of the American Association for the Ad- vancement of Science pp. 340-395. (GMK and CG) Darling Marine Center and Center for Marine Studies, University of Maine, Walpole, Maine 04573, U.S.A.; (IK) Department of Zoology and Center for Marine Studies, University of Maine, Orono, Maine 04469, U.S.A. Contribution 268 from the Darling Marine Center. PROC. BIOL. SOC. WASH. 107(2), 1994, pp. 391-397 ADVERTISEMENT CALLS AND RELATIONSHIPS OF CHILEAN FROGS EUPSOPHUS CONTULMOENSIS AND E. INSULARIS (AMPHIBIA: ANURA: LEPTODACTYLIDAE) J. Ramon Formas and Lila Brieva Abstract. — Advertisement calls of E. contulmoensis and E. insularis are de- scribed. The calls of both species consist ofa single short note. Call data, together with chromosomal information, indicate that E. contulmoensis, and E. insu- laris, E. roseus, E. calcaratus and E. migueli comprise a related group distinct from E. vertebralis and E. emiliopugini. Calls of frogs can be useful in revealing evolutionary relationships at the species level. In general, closely related species have some acoustic characteristics in common (Blair 1958, Barrio 1965, Mecham 1971, Martin 1972, Schiotz 1973, Kuramoto 1974, Ryan 1988, Straughan & Heyer 1971). In order to establish evolutionary trends re- lated to the diversification of the genus Eup- sophus, we analyzed the advertisement calls of E. contulmoensis and E. insularis. Both calls are compared with those other mem- bers of the genus that were previously de- scribed by Formas & Vera (1980), Formas (1985, 1989), and Penna & Veloso (1990). Eupsophus is the most speciose taxon of the rather reduced anuran fauna of the Nothofagus forests of southern Chile and Argentina. Seven species have been de- scribed: E. roseus, E. calcaratus, E. migueli, E. contulmoensis, E. insularis, E. vertebralis and E. emiliopugini. E. contulmoensis has been collected in the type locality of Contulmo (37°52’S, 73°12'W), Nahuelbuta Range (Ortiz et al. 1989), and E. insularis is endemic to Mocha Island (38°22'S, 73°56'S; 38 km W of the coast of Arauco Province) (Formas & Vera 1982). Methods and Materials Field recordings were made at 19 cm/sec on an Uher 4000 Report-IC portable tape- recorder and an Uher 517 microphone. Call durations, number of notes per call, and note duration were analyzed with a storage os- cilloscope Tektronix 5113. Audiospectro- grams were made with a Kay Elemetric au- diospectrograph model 675, using 85—8000 Hz frequency scale and narrow (45 Hz) band filter. Call repetition rates were measured in the field by counting consecutive calls over one minute. Water temperature, location and behavior of the individuals used in the analysis were taken at the time of recording. Specimens and tapes were deposited in the collection of Amphibians of the Institute of Zoology, Universidad Austral de Chile (IZUA), Valdivia. Eupsophus contulmoensis Ortiz, Ibarra-Vidal, & Formas, 1989 The advertisement call of E. contulmoen- sis was recorded at the type locality (Natural Monument of Contulmo, Malleco Prov- ince). Description is based on 25 calls from seven specimens recorded at 11.8°C (water temperature) during November 1990. Males 392 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 1.—Call characteristic (mean and range) of Eupsophus species (S, short call; L, long call) Call Notes Call length Dominant Species type per call sec Repetition rate frequency (Hz) E. contulmoensis 1 0.18 (0.15—0.20) 23.3 (15-34) (1100-2000) E. insularis 1 0.16 (0.14—0.18) 7.8 (4-12) (1500-2100) E. roseus S) 1 0.20 (0.19-0.21) 64.0 (60-72) (1600-2900) vid S 1 0.10 (0.07-0.16) 25.1 (11.1-60) (1250-1350) aes L 32.1 (8-47) 2.73 (0.65—40.0) 10.2 (93-112) (1220-1470) E. migueli +} S) 1 0.24 (0.20-0.35) 6.0 (3-8) (1500-2500) ks S 1 0.20 (0.16-0.26) 4.2 (2.4-6.6) (1170-1820) tT L 24.0 (19-33) 3.40 (3.30-4.40) 6.0 (5-8) (900-1500) 4 iL 12.3 (4-23) 1.07 (0.30-2.16) 5.4 (2.4-8.4) (1210-2000) E. calcaratus ++ 1 0.19 (0.15-0.21) 19.0 (16-25) (1100-2700) E. vertebralis + 5.0 (4-6) 0.60 (0.40-0.80) 4.0 (2-10) (1100-2500) i 5.6 (3.8) 0.64 (0.40-0.88) 27.8 (18.6—36.6) (700-1110) E. emiliopugini * 2) 0.50 = (729-1320) + From Formas & Vera (1980); +} Formas (1985); * Formas (1989); ** Penna & Veloso (1990). were observed and collected while they were calling from cavities in the ground at the border of a stream close to the forest. In- dividuals were organized into duets and tri- os spaced apart by 40-180 cm. Generally males called in alternation. The advertise- ment call of E. contulmoensis is composed of only one note (Fig. la, d; Table 1) that lasts 0.15-0.18 seconds. The dominant fre- quency is spread between 1100-2000 Hz and the repetition rate is 15—34 calls/min- ute. All calls showed modulation and de- fined harmonics are present at about 500 Hz intervals. Eupsophus insularis (Philippi) Description is based on 18 calls recorded from three individuals at Isla Mocha, dur- ing December 1989. Calls were recorded at a water temperature of 12°C. Males were observed and captured while they were call- ing from cavities in the ground close to a stream at the border of the forest. The in- dividuals call isolated and neither duets nor trios were detected. Calls consist of a single note (Fig. 1b, e; Table 1) lasting 0.14—0.18 seconds. The call is modulated and defined harmonics are present at about 500 Hz in- tervals. Maximum energy is spread over a frequency range of 1500—2100 Hz. E. roseus (Dumeril & Bibron), E. migueli Formas, E. calcaratus (Gunther), E. vertebralis Grandison, and E. emiliopugini Formas Call characteristics of the above cited spe- cies were described previously (Formas & Vera 1980; Formas 1985, 1989; Penna & Veloso 1990) and are depicted in Table 1. Discussion It is noteworthy that all Eupsophus spe- cies call during the spring-time from cavi- ties along margins of small streams in the forest (Formas & Vera 1980; Formas 1985, 1989; Penna & Veloso 1990). Among the anuran fauna of the temperate Nothofagus forest of South America, males of Eupso- phus species are unique in calling from flooded underground cavities near streams. Based only on temporal parameters (call length and notes per call) of advertisement calls, Formas (1985) established two clus- ters of species within the genus Eupsophus. The first group contains EF. roseus, E. mi- gueli and e. calcaratus, whose calls consist of only one note lasting 0.10 to 0.20 seconds (Table 1). Calls of E. contulmoensis and E. insularis (a single note lasting 0.14 to 0.18 seconds) provide good evidence for group- VOLUME 107, NUMBER 2 kHz Yo wo F-F HT Awa yQ ow 393 Fig. 1. 1 sec, Audiospectrograms of the advertisement call of Eupsophus contulmoensis (a), E. insularis (b), and E. roseus (c), band filter 45 Hz. Oscillograms of the advertisement call of E. contulmoensis (d) and E. insularis (e). ing both of these species within this species group. The second group includes E. ver- tebralis, whose call contains 4—6 notes last- ing 0.09 seconds. The advertisement call of E. emiliopugini shows that this species emits two notes lasting 0.20 seconds. Based on this information, we consider that E. emi- liopugini should be grouped with E. verte- bralis, rather than the roseus group. Both species groups (i.e., roseus and ver- tebralis) are also supported when spectral parameters of the calls are considered. Members of the vertebralis group show low- er mean values of dominant frequencies (1024-1350 Hz) than do members of the roseus group (1550-1900 Hz) (Table 1). On the other hand, members of the vertebralis group exhibit marked pulsed calls (11-34 394 Sula 3.6 r=0.996 > 2 35 y = 6.33—1.90 X 5 oO 2 Se 84 e = = 3.3 E °o mo) Coe fo) =! 3.1 EvO \Of&é 3.0 14 1.5 1.6 1.7 1.8 Log snout-vent length Fig. 2. Relation between mean snout—vent length and mean dominant frequency of the calls of Eupso- phus species. Snout—vent length measurements corre- spond to samples of males from localities where re- cordings were made. Abbreviations (sample size in parentheses): E.r., Eupsophus roseus (12), E.m., Eup- sophus migueli (7), E.ca., E. calcaratus (6), E.i., E. in- sularis (5), E.co., E. contulmoensis (7), E.v., E. verte- bralis (14), E.e., E. emiliopugini (17). pulses per note), while the roseus group spe- cies show calls poor in pulsations. As indicated earlier, different authors have suggested that calls of frogs are useful in revealing systematic and evolutionary re- lationships. Temporal and spectral param- eters of the advertisement calls of Eupso- phus species may also reveal information concerning the evolutionary history of these species. Spectral components of the call (dominant frequencies) show wide overlap within Eupsophus (Table 1). The frequency range (700-2900 Hz) at which these species call is also shared by other frogs of the tem- perate Nothofagus forest (Penna & Veloso 1990), and thus cannot be related to partic- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ular physical features of the call sites of these frogs. When the mean dominant frequen- cies of Eupsophus species are compared, E. emiliopugini and E. vertebralis show lower values (1024-1350 Hz) than those present- ed by E. contulmoensis, E. insularis, E. ro- seus, E. calcaratus and E. migueli (1550- 1900 Hz) (Table 1). The relationship be- tween the mean dominant frequency of the call varies inversely with the logarithm of animal size (r = —0.996) (Fig. 2). The ad- vertisement calls of FE. emiliopugini and E. vertebralis have been described as conspic- uously pulsed (Formas 1989). The former species emits 25.45 (17-34) pulses per note and the latter 15.90 (1 1-23) pulses per note. In contrast, calls of E. roseus, E. migueli, E. contulmoensis, E. insularis and E. cal- caratus are poor in pulsation; however the “B” call (or long call) of E. migueli is mark- edly pulsed; 6-9 pulses per note (Formas 1985). Eupsophus species exhibit considerable variation in calls, especially in the number of notes and their durations. Extremes are the single note (short call) of E. roseus, E. migueli, E. calcaratus, E. contulmoensis, and E. insularis (0.10—0.20 seconds), and the complex trilled calls (long call) with more than 40 notes of E. roseus and E. migueli (3.3-4.4 seconds). This latter vocalization has been interpreted as a territorial call (Formas 1985, Penna & Veloso 1990). Be- tween these extremes are the advertisement calls of E. emiliopugini (2 notes) and E. ver- tebralis (4-6 notes lasting 0.40 to 0.80 sec- onds). If the length of each individual note is considered, the individual notes of the call of E. emiliopugini lasts 0.20 (0.19-0.23) seconds in duration, a value similar to the single notes of the advertisement calls of E. roseus, E. calcaratus, E. migueli, E. con- tolmoensis, and E. insularis (X = 0.18; range 0.14—0.26 seconds). On the other hand, in- dividual notes of the advertisement call of E. vertebralis have a duration (¥ = 0.089; range 0.062—0.187) similar to the individual notes of the long call of E. migueli (X = VOLUME 107, NUMBER 2 0.083; range 0.051—-0.092). In spite of this similarity, both calls are different in the number of pulses per note (X = 15.90 in E. vertebralis; X = 7.5 in E. migueli), number of notes per call (4-6 in E. vertebralis; more than 40 notes in EF. migueli), and duration of the call (¥ = 0.6; range 0.4-0.8 seconds in E. vertebralis and X = 3.4; range 3.3—4.4 seconds in E. miguel). Although members of the roseus group exhibit some similar patterns in advertise- ment calls, differences can be found when intervals among harmonics are examined. Calls of E. contulmoensis and E. insularis have harmonics at about 500 Hz intervals, while E. calcaratus, E. migueliand E. roseus show harmonics at about 1000 Hz intervals. A comparison among the calls of E. roseus, E. contulmoensis and E. insularis is shown in Fig. 1. In the case of the vertebralis species group, both members (E. vertebralis and E. emiliopugini) have harmonics at about 250 Hz intervals (Formas 1989). Recognizing two species groups within the genus Eupsophus based on some temporal (call length and notes per call) and spectral (mean dominant frequencies and pulses) characteristics of advertisement calls agrees with the same proposal based on chromo- somal data (Formas 1980). Members of the vertebralis group (E. vertebralis and E. emi- liopugini) share a 28-chromosome karyo- type (Formas 1991), while members of the roseus group have a 30-chromosome karyo- type (Formas 1980, Iturra & Veloso 1981, 1989). Cuevas (personal communication) found that E. insularis and E. contulmoensis have a 30-chromosome karyotype, as oc- curs in E. roseus, E. migueli, and E. cal- caratus. Members of the Eupsophus roseus species group (E. roseus, E. calcaratus, E. insularis, E. contulmoensis, and E. migueli) are al- lopatric species; however, E. roseus and E. migueli are sympatric but not syntopic at the locality of Mehuin (39°26'S, 73°10’W) (Iturra & Veloso 1981). Members of the E. vertebralis species group (E. vertebralis and 395 E. emiliopugini) are also allopatric. In spite of the fact that species of each group (roseus and vertebralis) are allopatric, it is possible to find members of the two different species groups in sympatry. For example, at the lo- cality of Mehuin, EF. vertebralis and E. mi- gueli were observed while calling from cav- ities separated by two or three meters (Brieva, personal observation). A similar case of sympatry between members of the two different species group was observed at Cuesta de Soto (7 km by road from Valdivia city) between E. roseus and E. vertebralis. In three localities from southern Chile (La Picada, 42°07'S, 73°49’'W; Rio Rollizo, 41°27'S, 72°27'W; and Puntra, 42°07'S, 73°49'W) E. emiliopugini and E. calcaratus were observed calling from cavities sepa- rated by 25-50 cm (Formas 1989). Finally, E. vertebralis and E. contulmoensis were ob- served while calling from cavities separated by two to four meters at the locality of Con- tulmo (Natural Monument of Contulmo) (Formas, personal observation). The four pairs of Eupsophus species re- corded to occur in sympatry (EZ. migueli and E. vertebralis; E. roseus and E. vertebralis; E. calcaratus and E. emiliopugini; E. ver- tebralis and E. contulmoensis) are members of the two different species groups. These pairs of sympatric species exhibit a strong divergence among their advertisement calls. Passmore (1981) found that sympatric spe- cies of the African frog genus Ptychadena show remarkable differences in temporal and spectral components of their calls. In order to explain the origin of this phenomenon he used the notion of the Specific Mate Rec- ognition System (SMRS) (Paterson 1978, 1982). According to this view, interspecific differences in signals do not agree with the traditional hypothesis that call divergence among closely related species develops due to selection for increasing reproductive iso- lation upon re-encounter of previously iso- lated populations (Dobzhansky 1970). Passmore’s (1981) explanation for Ptycha- dena species is that the “pattern of vocal 396 divergence probably resulted from indepen- dent evolutionary histories, and concom- itant with the development of their respec- tive acoustical recognition mechanism, rather than from selective pressure for in- creasing reproductive isolation” (Penna & Veloso 1990). In hight of the SMRS concept we propose a hypothesis addressing the divergence of vocal patterns and the evolutionary history of the genus Eupsophus. As shown above (Table 1) the vertebralis group (E. vertebralis and E. emiliopugini) exhibits two different advertisement calls. Both members of this species group occur allopatrically (Formas 1989) and their karyotypes show differences in relation to chromosome pair No. 14 (telo- centric in E. vertebralis and metacentric in E. emiliopugini,; Formas 1991). From an evolutionary point of view, the genus Eup- sophus has a long history, with early generic differentiation documented by fossil rec- ords dating back to the Oligocene (Schaeffer 1949, Baez & Gasparini 1979). Conditions for further acoustical and chromosomal di- versification within these species existed during Pleistocene glaciations, during which reduced areas free of ice provided scattered refugia for the survival of flora and fauna (Vuilleumier 1968, Heusser 1974, Ash- worth et al. 1991). Since the present geo- graphical ranges of E. vertebralis and E. em- iliopugini (especially the southermost part) are in an area that was affected by glacia- tions, we hypothesize that ice-free areas were the appropriate scenarios where acoustical and chromosomal differentiation took place. Frogs of the roseus group (E. roseus, E. mig- ueli, E. calcaratus, E. contulmoensis, and E. insularis) are allopatric species sharing a similar one note advertisement call; how- ever, E. contulmoensis and E. insularis have harmonics at about 500 Hz intervals while E. roseus, E. migueli and E. calcaratus have harmonics at about 1000 Hz intervals. Vo- cal differences among these allopatric spe- cies suggest that these species represent a recent stage of speciation. All members of PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON the roseus group have the same chromo- somal formula (2N = 30), although differ- ences have been found among the karyo- types of some species. For example, the karyotypes of E. roseus and E. migueli (Itur- ra & Veloso 1989) exhibit marked similar- ities, but sex chromosomes differ. In these two species, the Y chromosome is a small metacentric one without paracentromeric heterochromatin, and corresponds to pair No. 14. In E. roseus, the sex chromosomes have the same metacentric morphology, but in E. migueli the X chromosome is telo- centric. This example of chromosomal di- vergence between two members of the ro- seus group (which share the same temporal and spectral characteristic of the advertise- ment calls) suggests that chromosomal dif- ferentiation has played a more important role in the diversification of some members of the roseus species group than has the ad- vertisement call. Finally, based on chro- mosome data, we suggest that vocal differ- ences noted for Eupsophus species that occur in sympatry results from the presumed in- dependent evolutionary histories of the taxa involved. Acknowledgments This study was supported by Fondo Na- cional de Investigacion Cientifica y Tec- nologica (Proyecto 0050-89) and Direccion de Investigacion y Desarrollo, Universidad Austral de Chile (Proyecto RS 89-06). We are grateful to Corporacion Nacional Fores- tal (CONAF) for its hospitality and field facilities at Parque Nacional Contulmo and Isla Mocha. The audiospectrograms were done by Alfredo Brain, Instituto de Acus- tica, Universidad Austral de Chile. Mario Penna assisted us with the osilloscopic anal- yses. We wish to thank Carlos Jara for his useful comments on the manuscript. Raul Arriagada aided with collections and tape recordings in the field. Our thanks to Ron- ald Heyer and two anonymous reviewers for their suggestions. VOLUME 107, NUMBER 2 Literature Cited Ashworth, A., V. Markgraf, & C. Villagran. 1991. Late Quaternary climate of the Chilean Channels based on fossil pollen and beetle analyses with an analysis of the modern vegetation and pollen rain. — Journal of Quaternary Science 6:279-291. Baez, A. M., & Z. B. Gasparini. 1979. The South American Herpetofauna: an evaluation of the fossil record. In W. E. Duellman, ed., The South American Herpetofauna: its origin, evolution and dispersal.— Museum of Natural History of Kan- sas Monographs 7:29-54. Barrio, A. 1965. El género Physalaemus (Anura, Lep- todactylidae) en la Argentina. —Physis 25:421- 448. Blair, W. F. 1958. Call structure and species group in USS. tree frogs (Hy/a).—Southwestern Nat- uralist 3:77-89. Dobzhansky, T. 1970. Genetics of evolutionary pro- cess. Columbia Press, New York. Formas, J.R. 1980. The chromosomes of E. calcara- tus and the karyological evolution of the genus Eupsophus (Anura: Leptodactylidae).—Exper- lentia 36:1163-1164. 1985. The voices and relationships of the Chilean frogs Eupsophus migueli and Eupso- phus calcaratus (Amphibia: Anura: Leptodac- tylidae).— Proceedings of the Biological Society of Washington 98:411-415. 1989. A new species of Eupsophus (Am- phibia: Anura: Leptodactylidae).— Proceedings of the Biological Society of Washington 102: 568-576. 1991. The karyotypes of Eupsophus emilio- pugini and E. vertebralis (Amphibia: Anura: Leptodactylidae).— Proceedings of the Biologi- cal Society of Washington 104:7-11. ——, & M.A. Vera. 1980. Reproductive patterns of Eupsophus roseus and E. vittatus.—Journal of Herpetology 14:11-14. ——, &M.I. Vera. 1982. The status of two Chilean frogs of the genus Eupsophus.—Proceedings of the Biological Society of Washington 95:594— 601. Heusser, C. J. 1974. Vegetation and climate of the Southern Chilean Lake District during and since the last glaciation. — Quaternary Sciences (N.Y.) 4:290-315. Iturra, P., & A. Veloso. 1981. Evidence for hetero- morphic sex chromosomes in male amphibians (Anura: Leptodactylidae).— Cytogenetic and Cell Genetics 31:108-110. ——, & 1989. Further evidence for early 397 sex chromosome differentiation of anuran spe- cies. — Genetica 78:25-31. Kuramoto, M. 1974. Mating calls of Japanese tree frogs (Rhacophoridae).— Bulletin of the Fuku- oka University, Faculty of Education 26:67-77. Martin, W. F. 1972. Evolution of vocalization in the genus Bufo. Pp. 279-309 in W. F. Blair, ed., Evolution in the genus Bufo. University of Tex- as Press, Austin and London. Mecham, T.S. 1971. Vocalization of the leopard frog, Rana pipiens and the related Mexican species. — Copeia 71:506-516. Ortiz, J. C., H. Ibarra-Vidal, & J. R. Formas. 1989. A new species of Eupsophus (Anura: Leptodac- tylidae) from Contulmo, Nahuelbuta Range. — Proceedings of the Biological Study of Wash- ington 102:1031-1035. Passmore, N. I. 1981. The relevance of the specific mate recognition concept to anuran reproduc- tive biology.—Monitore Zoologico Italiano (N.S.) Supplement XV:93-108. Paterson, H.E.M. 1978. More evidence against spe- ciation by reinforcement.—South African Jour- nal of Science 74:369-371. . 1982. Perspective on speciation by reinforce- ment.— African Journal of Science 78:53-57. Penna, M., & A. Veloso. 1990. Vocal diversity in frogs of the South American temperate forest. — Journal of Herpetology 24:23-33. Ryan, M. J. 1988. Constraints and patterns in the evolution of anuran acoustic communication. Pp. 637-677 in B. Fritzsch, M. Ryan, W. Wil- ezynski, T. Hetherington, & W. Walkowiak, eds., The evolution of the Amphibian Auditory Sys- tem. John Wiley, New York. Schaeffer, B. 1949. Anurans from the early Tertiary of Patagonia.—Bulletin of American Museum of Natural History 93:41-68. Schigtz, A. 1973. Evolution of anuran mating calls, ecological aspects. Pp. 311-319 in J. L. Vial, ed., Evolutionary biology of the Anurans. Uni- versity of Missouri Press, Columbia. Straughan, I. R., & R. Heyer. 1976. A functional analysis of the mating calls of the neotropical frog genera of the Leptodactylus complex (Am- phibia, Leptodactylidae).—Papéis Avulsos de Zoologia 29:221—245. Vuilleumier, F. 1968. Origin of frogs of Patagonian forests.— Nature 219:87-89. Instituto de Zoologia, Universidad Aus- tral de Chile, Casilla 567, Valdivia, Chile. PROC. BIOL. SOC. WASH. 107(2), 1994, pp. 398-409 TWO NEW SPECIES OF THE HYLA SUMICHRASTI GROUP (AMPHIBIA: ANURA: HYLIDAE) FROM MEXICO Joseph R. Mendelson III and Jonathan A. Campbell Abstract. —Two new species belonging to the Hyla sumichrasti group of Mex- ico are described; one is from the cloud forests of the southeastern Oaxacan highlands and the other from the extremely xeric Zapotitlan Valley of Puebla. The new species from Oaxaca can be distinguished from other members of the group based on body proportions and external morphology. The new species from Puebla resembles 7. sumichrasti, but differs in hand size and morphology, snout shape, and presence of vomerine teeth. A canonical analysis of morpho- metrics of the four species in the group reveals that H. sumichrasti and the new species from Puebla are similar, but distinguishable, whereas H. smar- agdina and the new species from Oaxaca are distinct from the other members of the group. Hyla sumichrasti is reported for the first time from the state of Guerrero. Resumen. —Se describen dos especies nuevas que pertencen al grupo Hyla sumichrasti de México, una del bosque de neblina de las montanas del sureste de Oaxaca y la otra del valle arido de Zapotitlan en Puebla. La especie nueva de Oaxaca se puede distinguir de los otros miembros del grupo sobre la base de sus proporciones corporales y su morfologia externa. La especie nueva de Puebla es similar a H. sumichrasti, pero tiene manos de forma y tamano diferentes, hocico de forma diferente y dientes vomerinos. Un analisis canonico de la morfometria de las cuatro especies del grupo revela que Hyla sumichrasti y la especie nueva de Puebla son similares, pero se pueden distinguir mientras que Hyla smaragdina y \a especie nueva de Oaxaca son distintas de los otros miembros del grupo. Se reporta por primera vez una poblacion de Hyla su- michrasti para el estado de Guerrero. Nearly 20 years have passed since the de- scription of any new members of the spe- ciose genus Hyla from the vast and geo- graphically complex regions of Mexico (see Caldwell 1974). We examined two enig- matic series of specimens belonging to the Hyla sumichrasti group (Duellman 1970), which is endemic to Mexico, and conclude that each represents an undescribed species. Members of the Hyla sumichrasti group (H. sumichrasti and H. smaragdina, from southern and western Mexico, respectively) have larvae with immense mouths contain- ing three upper and seven lower tooth rows and the adults have distinctively broad, flat heads with a unique combination of cranial characters— viz. massive nasals that are in broad contact medially, a short and broad sphenethmoid, a large frontoparietal fon- tanelle, and absence of quadratojugals (Duellman 1970). By virtue of these char- acters, the group is remarkably distinct from other species groups in Middle America (Duellman 1970). Herein we describe two VOLUME 107, NUMBER 2 new species which we place in the H. sumi- chrasti group based on the cranial and ex- ternal characters of the adults. Materials and Methods Terminology and measurements follow those described by Duellman (1970) and the descriptions are formatted similarly for ease of comparison. Webbing formulae for hands and feet follow Myers & Duellman (1982). All measurements were taken by the same person (JRM) using digital calipers. Obser- vations were made under a dissecting mi- croscope. The condition of the nasal bones was evaluated by lifting the skin of the snout posteriorly until the bones were exposed. Comparative material examined 1s listed in Appendix 1; specimens examined are from the collections at The University of Kansas (KU), The University of Texas at Arlington (UTA), and the California Academy of Sci- ences (CAS). All localities were presented in metric measurements. Localities of type specimens are presented here as they appear in the original catalogues. Small sample sizes limited comparisons and morphometric analysis to male specimens only. Stepwise discriminant function analysis (DFA) was performed using the BMDP computer program on the following mor- phometric measurements (all log-trans- formed): (1) snout—vent length (SVL), (2) tibia length, (3) foot length, (4) head length, (5) head width, (6) anterior margin of tym- panum—posterior margin of eye distance, (7) eye—nostril distance, (8) diameter of tympanum, (9) length of first finger, (10) length of third finger, and (11) diameter of disc on third finger. Systematics Hyla chimalapa, new species Figs. 1; 2A; 3A Holotype. —UTA A-13365 (original number JAC 9324), a male from Colonia Rodulfo Figueroa, 19 km NW Rizo de Oro 399 (Chiapas), 1542 m, Oaxaca, Mexico; ob- tained on 4 Aug 1983 by J. A. Campbell, D. M. Hillis, and W. W. Lamar. Paratopotype.—UTA A-13366, a male, other data as for holotype. Paratypes.—CAS 170121, 170122, 170124, 170125, 170127, 170130, adult males, and CAS 170119, 170120, 170123, 170126, 170128, 170129, adult females, from 5-7 mi NW Rizo de Oro along road to Cerro Baul, Chiapas, Mexico, 3300 feet elev., collected by D. E. Breedlove on 19 Apr 1972; CAS 170114, an adult female from 12 mi W of Rizo de Oro (Chiapas) along ridge south of Cerro Baul, Oaxaca, Mexico, 5000 feet elev., obtained by D. E. Breedlove on 28 Apr 1972. Referred specimens. —KU 179072, an adult male from 6.5 km (by road) NE Mil- tepec (=Niltepec), Oaxaca, Mexico, ob- tained by D. R. Frost and S. Aird on 16 Jun 1977. CAS 163309, an adult female from 1.6 km W Rizo de Oro, crest of ridge be- tween Tapanatepec and Cintalapa Valley, 853 m, Chiapas, Mexico, obtained by D. E. Breedlove on 12 Jun 1965. Diagnosis. —Hyla chimalapa is referred to the H. sumichrasti group because of the broad, flat head and large nasals that are in broad contact medially. This species may be distinguished from other members of this group by the following combination of char- acters: (1) tympanum distinct; (2) snout acutely pointed in dorsal view and protrud- ing, pointed in lateral view; (3) vomerine teeth absent; (4) tongue round; (5) tarsal fold distinct; (6) tibiotarsal articulation extend- ing to snout; (7) fingers long, slender, about half webbed; (8) transverse dermal fold on heel absent; (9) ventral coloration extending from axilla onto dorsomedial surface of up- per arm; and (10) row of ulnar tubercles present. Hyla chimalapa differs from H. sumi- chrasti by having relatively longer legs; a relatively wider head; a distinct tympanum; round tongue; snout protruding and pointed in lateral view (Fig. 3a, c); fingers longer, 400 Fig. 1. more slender; distinct tarsal fold present; transverse dermal fold on heel absent; dor- somedial surface of upper arm light; row of ulnar tubercles present; and vomerine teeth absent. Hyla chimalapa differs from H. smaragdina by being larger, and by having relatively longer legs; a relatively wider head; flecking on dorsum less extensive; dorso- medial surface of upper arm light; snout more pointed in dorsal and lateral view (Fig. 3a, d); tongue round; row of ulnar tubercles present; fingers relatively longer, more slen- der, about half webbed; transverse dermal fold on heel absent; and vomerine teeth ab- sent. Hyla chimalapa differs from Hyla xera (described below) by being smaller, and by having relatively longer legs; snout more pointed in profile; fingers longer, more slen- der, about half webbed; distinct tarsal fold present; row of ulnar tubercles present; pal- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Hyla chimalapa, holotype, UTA A-13365, photographed in life. mar tubercle present; transverse dermal fold on heel absent; and dorsomedial surface of upper arm light. Description of holotype.—Body robust; head as wide as body, slightly wider than long; head width 34.5 percent SVL; head length 33.7 percent SVL; snout acutely pointed in dorsal view, protruding and pointed in profile; distance from eye to nos- tril equal to diameter of eye; nostril four- fifths distance from eye to tip of snout; top of head flat, smooth; canthus rostralis dis- tinct, rounded; loreal region flat; lips thin, barely flared; interorbital distance about 50 percent greater than width of eyelid; tym- panum distinct with raised annulus, diam- eter about equal to distance from eye to tympanum; supratympanic fold thin, ob- scuring posterodorsal margin of tympanum, extending posteriorly from orbit, postero- VOLUME 107, NUMBER 2 401 Fig. 2. Ventral aspect of the hand of (A) Hyla chimalapa, holotype, UTA A-13365, and (B) Hyla xera, holotype, UTA A-13387. Scale bar represents 2 mm. ventrally from point above tympanum, be- coming indistinct at point over the insertion of the arm. Axillary membrane extending about one- fourth length of upper arm; ulnar tubercles present, weakly defined; dermal fold on wrist distinct; fingers long, slender, bearing ex- panded, slightly ovoid terminal discs; di- ameter of disc on third finger slightly larger than diameter of tympanum; relative length of fingers 1 < 4 < 2 < 3; fingers about half webbed with slight lateral fringe on all fin- gers except first, and medial surface of sec- ond; webbing formula I2* —2+*II2+ — 2+1112+ —2-IV; subarticular tubercles in- distinct except for round, elevated tubercles on first finger; supernumerary tubercles ab- sent; palmar tubercle small, low, bifid; pre- pollex barely evident, lacking horny excres- cence. Hind limb long; shank robust; tibia length 56.6 percent SVL; foot length 49.4 percent SVL; heels of adpressed limbs over- lapping by one third length of shank; tibio- tarsal articulation extending to snout; tarsal fold distinct, extending full length of tarsus; transverse dermal fold on heel absent; inner metatarsal tubercle small, oval, low; outer metatarsal tubercle absent; subarticular tu- bercles round, low, flat; supernumerary tu- bercles absent; toes long, slender, bearing round discs slightly smaller than those on fingers; toes about three-fourths webbed with lateral fringe except on outer edges of first and fifth toes; webbing formula I1 —2-II1 — 2-WIl—2-IV2-—1V. Skin on throat and belly granular, other surfaces smooth; cloacal opening directed posteroventrally at midlevel of thighs; clo- acal sheath short. Vomerine teeth absent. Choanae small, subcircular, widely spaced. Tongue round, barely free posteriorly. Vo- cal slits extending from midlateral base of tongue nearly to angle of jaw; vocal sac sin- gle, median, subgular. Measurements (mm): Measurements of the holotype followed by those of the para- topotype in parentheses. SVL 24.9 (23.3), tibia length 14.1 (13.6), foot length 12.3 (11.4), head length 8.4 (7.9), head width 8.6 (8.0), interorbital distance 3.7 (3.2), eyelid 402 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Ze ie SS a D Fig. 3. Dorsal (upper) and lateral (lower) profiles of the snouts of (A) Hyla chimalapa, holotype, UTA A-13365, (B) Hyla xera, holotype, UTA A-13387, (C) Hyla sumichrasti, KU 100941, and (D), Hyla smaragdina, KU 75314. Scale bar represents 2 mm. VOLUME 107, NUMBER 2 width 2.4 (2.5), eye—nostril distance 2.9 (2.7), eye diameter 3.2 (2.7), tympanum diameter 1.4 (1.4). Color in preservative (70 percent ethanol after formalin): Dorsum of head, forearms, fourth and fifth fingers dark brown; dorsum of body, and hind limbs pale brown with few, scattered, dark brown flecks; dorsum of first and second fingers cream with fine brown stippling; axillary area and dorso- medial surface of arms cream; flanks, all ventral surfaces cream, except posterior sur- face of thighs, periphery of lower jaw and ventral surface of hands and feet cream with fine brown stippling; tarsal fold sharply sep- arating dorsal and ventral colors on foot. Color in life (from UTA Slide No. 133): Dorsal surfaces pale brown with scattered small dark brown flecks; head and tympanic area darker than other surfaces; tympanum pale brown with cream flecks; posterior sur- face of thigh and ventral surface of tibia red- orange; first finger pale yellow; flanks cream with brown stippling; iris dull gold with black reticulations. Variation.—The measurements of the paratypes and the referred specimens are as follows (mm; range followed by mean in parentheses for males and females, respec- tively): SVL 20.3-23.6 (22.1), 18.4-26.6 (23.7); tibia length 10.9-12.3 (11.5), 10.8- 14.2 (12.7); foot length 9.2-10.2 (9.6), 9.2— 12.9 (11.2); head length 6.7—7.4 (7.2), 6.6— 9.0 (7.8); head width 6.7—8.0 (7.3), 6.4—8.7 (8.0); eye—nostril distance 2.2—2.5 (2.4), 2.1- 3.1 (2.6); tympanum diameter 1.0-1.2 (1.1), 1.1-1.6 (1.4). The proportions of the female specimens are given here rather than in Ta- ble 1 because we did not examine compa- rable numbers of females of the other spe- cies in the group: tibia/SVL 0.50-0.56 (0.54); foot/SVL 0.45-0.50 (0.47); head length/SVL 0.29-0.36 (0.33); head width/SVL 0.32-0.37 (0.34). The paratypes and referred specimens are similar to the holotype in proportion and are similar in color in preservative. The ul- nar tubercles are more distinct in these spec- 403 imens than in either the holotype or the paratopotype and the profile of the snout is more rounded; the protuberance illustrated in Fig. 3A is less pronounced in the para- types. Distribution and ecology.—Hyla chima- lapa is known only from the southeastern Oaxacan Highlands, which includes the Si- erra Atravesada (sensu Campbell 1984). This area is known locally as the Chimalapa region. All specimens have been collected between the town of Rizo de Oro and Cerro Baul—the highest peak in the region, except one (KU 179072) from the Sierra Atrave- sada. In August 1992, the Cerro Baul area still supported some disturbed cloud forest. This region is characterized by a complex mosaic of mesic cloud forest and relatively xeric pine-oak forests. The type locality is in cloud forest but there are no habitat notes associated with the remaining specimens and they may have been collected in the more xeric areas. The other species in the H. su- michrasti group inhabit moderately to ex- tremely xeric habitats and it 1s possible that H. chimalapa occurs in the dry pine-oak forests of the region. The holotype and para- topotype were discovered at night along a wide, shallow stream at the edge of Colonia Rodulfo Figueroa. This stream flows swiftly over a sand and gravel substrate, forming numerous sandbars; in August 1992, these sandbars were covered by dense stands of young willows (Salix sp.) that were not pres- ent when the types of H. chimalapa were collected. Other species of anurans from this area include Bufo marinus, B. valliceps, Rana maculata, Eleutherodactylus lineatus, E. leprus, Ptychohyla euthysanota, and Plec- trohyla matudai. Life history.—The advertisement call, eggs, and tadpole of H. chimalapa are un- known. Adult females (CAS 170128, 170129) collected on 19 Apr 1972 contain partially developed ova, and another adult female (CAS 170123) from this date con- tains no apparent ova. An adult female (CAS 170114) obtained on 28 Apr 1972 contains 404 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 1.—Comparison of selected measurements (mm), proportions, and morphological features of adult males of the H. sumichrasti group. Character abbreviations SVL, snout—vent length; TL, tibia length; FT, foot length; HL, head length; HW, head width. Range of values followed by means in parentheses. Character SVL TL/SVL FT/SVL HL/SVL HW/SVL Snout (dorsal) Snout (lateral) Tarsal fold Heel fold Tibiotarsal articula- tion Hand webbing Vomerine teeth Palmar tubercle Ulnar tubercles H. sumichrasti 22.5-27.1 (24.5) 0.43-0.53 (0.48) 0.39-0.48 (0.42) 0.29-0.33 (0.31) 0.30-0.34 (0.32) Bluntly rounded Flattened Variable Present Orbit 12+—2', I12-—2'4 I11234——2+1V Variable Present Absent H. smaragdina 22.7—-25.5 (24.0) 0.43-0.49 (0.46) 0.40-0.48 (0.44) 0.30-0.33 (0.32) 0.27-0.30 (0.28) Rounded Rounded Strong Present Orbit 12+——2%4 112-—3+ II3-—2tIV Present Present Absent H. chimalapa 20.1-24.9 (22.1) 0.50-0.58 (0.54) 0.43-0.49 (0.45) 0.31-0.34 (0.33) 0.32-0.36 (0.34) Acutely pointed Protruding Strong Absent Snout 12+——2+ 112*——2* 112+——2-IV Absent Present Present H. xera 25.8-27.9 (26.8) 0.46-0.50 (0.48) 0.42-0.45 (0.43) 0.29-0.32 (0.31) 0.30-0.32 (0.31) Acutely rounded Bluntly rounded Weak Present Orbit I2',—2% I112-—3- 112'2—2+ IV Absent Absent Absent a large number of mature ova, as does an adult female (CAS 163309) obtained on 12 Jun 1965. Only one male (KU 179072; 16 Jun 1972) appears to be in breeding con- dition. This specimen bears nuptial excres- cences and a distended vocal sac. Etymology. —The specific name is treated as indeclinable and refers to the spectacular and still incompletely surveyed Chimalapa region of eastern Oaxaca, from which this species 1s known. Remarks.—The type locality may be reached by way of a Jong, tortuous drive along an unpaved road bearing NW from the small town of Rizo de Oro, Chiapas (NE of Tapanatepec on MX Hwy 190). Whether the type locality is in Oaxaca or Chiapas is a matter of local dispute; however, most maps show Cerro Baul to be in Oaxaca. This area is near the continental divide but Co- lonia Rodulfo Figueroa lies near the head- waters of the Rio Mono Blanco, a tributary of the Rio Negro (an Atlantic drainage). The southeastern Oaxacan highlands have complex biogeographic relationships with the proximal highland areas of southern Mexico (Campbell 1984). This area shares approximately the same number of am- phibian species with the distant Sierra de los Tuxtlas of Veracruz as it does with the more proximal Sierra Madre de Chiapas (Campbell 1984), but no member of the H. sumichrasti group is known from these two regions. The Sierra Madre de Chiapas shares several disjunct, but poorly differentiated, species of cloud forest anurans with the southeastern Oaxacan highlands (Campbell 1984). It is possible that H. chimalapa will be discovered in this range, especially if it does inhabit the more widespread pine-oak forests (Breedlove 1973) of the region. Hyla sumichrasti is known to the east of the southeastern Oaxacan highlands, in the xeric areas of the Sierra Madre del Sur and the Isthmus of Tehuantepec, and to the northwest from the more humid areas in the Central Depression and the cloud forests near Pueblo Nuevo Solistahuacan, Chiapas (Duellman 1970); this species has not been collected in the southeastern Oaxacan high- lands. Hyla sumichrasti is more variable in its external morphology than the other members of the group. Variation exists both within and among populations in such char- VOLUME 107, NUMBER 2 405 Fig. 4. Hyla xera, holotype (preserved), UTA A-13387. acters as the presence of vomerine teeth, presence and degree of development of the tarsal fold, overall body proportions (Table 1) and details of color pattern evident in preserved specimens. For this reason we ex- amined populations from both east and west of the southeastern Oaxacan highlands (Ap- pendix 1) in preparing the diagnosis for H. chimalapa and the morphometric analysis. Despite the variation evident in H. sumi- chrasti, H. chimalapa is diagnosable on the basis of traditional external characters and proportions (Table 1, Fig. 5). Duellman (1970) commented that H. sumichrasti and H. smaragdina may be con- specific, and cited the existence of a large distributional hiatus between the two spe- cies along the west coast of Mexico, despite the existence of suitable habitat in Guer- rero. We here take the opportunity to report a series of specimens referable to H. sumi- chrasti collected by T. Pappenfuss in 1976 in Guerrero (Appendix 1). The discovery of this population does not support the hy- pothesis that H. sumichrasti and H. sma- ragdina are conspecific. Hyla xera, new species Figs. 2B; 3B; 4 Holotype. —UTA A-13387 (original number JAC 6577), an adult male from 5.6 km SSW Zapotitlan Salinas, 1490 m, Pueb- la, Mexico; obtained on | July 1981 by J. A. Campbell. Paratopotypes. —UTA A-13381-83, 13385, 13386, 13388 adult males, and UTA A-13384, an adult female, all collected with the holotype by J. A. Campbell. 406 CAN II CAN I Fig. 5. Plot of canonical discriminant scores for all species in the Hyla sumichrasti group: H. chimalapa (triangles), H. smaragdina (circles), H. sumichrasti (squares), and H. xera (diamonds). Closed symbols represent individual scores, open symbols indicate group mean scores. Diagnosis.— Hyla xera is referred to the H. sumichrasti group because it has a broad, flat head and large nasals that are in broad contact medially. This species can be dis- tinguished from all other members of the group by the following combination of char- acters: (1) large SVL in adult males (¥ = 26.8); (2) a uniformly dull gray dorsum, with darker loreal and tympanic areas (in pre- servative); (3) tympanum distinct; (4) su- pratympanic fold thick, darkly colored; (5) snout acutely rounded in dorsal view, blunt- ly rounded in lateral view, sometimes with weak rostral keel; (6) dermal folds on wrist and heel present; (7) tarsal fold weakly de- veloped; (8) fingers long, slender, about one- third webbed; (9) palmar tubercle absent; (10) tibiotarsal articulation extending to an- terior margin of orbit; (11) vomerine teeth absent; (12) tongue cordiform; (13) row of ulnar tubercles absent; and (14) transverse dermal fold on heel present. Hyla xera differs from H. sumichrasti by having a distinct tympanum with a thick, darkly colored supratympanic fold; snout PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON more rounded profile (Fig. 3B, C); palmar tubercle absent; fingers relatively longer; vomerine teeth absent; and SVL of adult males probably larger (Table 1; but see Duellman 1970). Hyla xera differs from H. smaragdina by being larger; snout less rounded in dorsal and lateral view (Fig. 3B, D); supratympanic fold thick, darkly col- ored; flecking on dorsum less extensive; tar- sal fold weakly developed; palmar tubercle absent; fingers longer, more slender; and vo- merine teeth absent. Hyla xera differs from H. chimalapa by being larger and having legs relatively shorter; snout more rounded in dorsal and lateral view (Fig. 3A, B); tarsal fold weakly developed; ulnar tubercles ab- sent; palmar tubercle absent; fingers about one-third webbed; transverse dermal fold on heel present; and supratympanic fold thick, darkly colored. Description of holotype.—Body robust; head as wide as body, slightly wider than long; head width 31.7 percent SVL; head length 30.0 percent SVL; snout acutely rounded in dorsal view with weak rostral keel, bluntly rounded in profile; distance from eye to nostril equal to diameter of eye; nostril four-fifths distance from eye to tip of snout; top of head flat, smooth; canthus rostralis distinct, rounded; loreal region slightly concave; lips thin; interorbital dis- tance about 70 percent greater than width of eyelid; tympanum distinct without raised annulus, diameter about equal to distance from eye to tympanum; supratympanic fold thick, obscuring dorsal and posterodorsal margins of tympanum, extending posteri- orly from orbit, posteroventrally from point above tympanum, becoming indistinct an- terior to arm. Axillary membrane extending about one- third length of upper arm; ulnar tubercles absent; dermal fold on wrist distinct; fingers long, slender, bearing expanded, round, ter- minal discs; diameter of disc on third finger slightly smaller than diameter of tympa- num; relative length of fingers 1 < 2 < 4 < 3; fingers about one-third webbed, with- VOLUME 107, NUMBER 2 out lateral fringe; webbing formula I2!2— 234112 —3~- II2'4%2—2+* IV; subarticular tu- bercles indistinct except for round, elevated tubercles on first finger; supernumerary tu- bercles absent except on prepollex; palmar tubercle absent; prepollex enlarged, bearing horny excrescence barely visible ventrally, covering posterior and dorsal portions of prepollex. Hind limbs long; shank robust; tibia length 48.7 percent SVL; foot length 42.2 percent SVL; heels of adpressed limbs overlapping by one-third length of shank; tibiotarsal articulation extending to anterior margin of orbit; tarsal fold absent; trans- verse dermal fold on heel distinct; inner metatarsal tubercle small, oval, low; outer metatarsal tubercle absent; subarticular tu- bercles small, round, slightly elevated on first and second toes, low, round, flat on fourth and fifth toes; supernumerary tuber- cles absent except on base of first toe; toes long, slender, bearing round discs about the same size as those on fingers; toes about three-fourths webbed, lacking lateral fringe except on fourth toe; webbing formula I1 — 2*H1 —2'”lll1 —2t+1V2—1V. Skin on throat and belly granular, other surfaces smooth; cloacal opening directed posteroventrally at midlevel of thighs; clo- acal sheath short. Vomerine teeth absent. Choanae small, slightly ovoid, widely spaced. Tongue cordiform, barely free pos- teriorly. Vocal slits extending from midlat- eral base of tongue to nearly to angle of jaw; vocal sac single, median, subgular. Measurements of the holotype (mm): SVL 26.7; tibia length 13.0; foot length 11.3; head length 8.0; head width 8.5; interorbital dis- tance 3.0; eyelid width 2.0; eye—nostril dis- tance 2.7; tympanum diameter 1.6. Color in preservative (70 percent ethanol after formalin): All dorsal surfaces dull gray except feet and thighs pale brown; loreal and tympanic areas darker than dorsum of head, dark coloring following and clearly demar- cating supratympanic fold; supratympanic areas and dorsal surfaces of shanks with dull silver marbling; posterior surface of thigh 407 dull yellowish brown; flanks and all ventral surfaces dull cream; palmar and plantar sur- faces cream with very fine gray stippling. Variation. — The range of variation (mm; with means in parentheses) of the male paratopotypes are followed by those of the female paratopotype. SVL 25.6—27.9(26.9), 31.2; tibia length 12.5-13.3(12.8), 15.4; foot length 11.0-12.4(11.8), 13.1; head length 8.1-8.9(8.3), 9.1; head width 8.1-8.8(8.4), 10.3; interorbital distance 2.9—3.1(3.0), 3.5; eyelid width 1.9-2.4(2.1), 2.2; eye—nostril distance 2.7—2.8(2.8), 3.1; tympanum di- ameter 1.3-1.8(1.5), 1.6. Variation in body proportions are summarized in Table | and, along canonical axes, in Fig. 5. The color- ation (in preservative) of all paratopotypes is very Similar to that of the holotype, except UTA A-13382 (male) and UTA A-13384 (female), which have distinctive dull silver and dull brown marbling on the flanks and posterior surfaces of the shanks and tarsi. The transverse dermal fold and tarsal fold are weakly developed in this species and appear to be absent in some individuals in the type series. Distribution and ecology.—Hyla xera is known only from the type locality. This area supports arid tropical scrub (sensu Leopold 1950) and is characterized by scattered mes- quite trees (Prosopis), other leguminous trees, and many species of cactus. Patches of thorn forest and tropical deciduous forest occur locally, especially in the draws. No permanent water occurs naturally in the Za- potitlan Valley, even the largest stream draining the valley is seasonal. Most spec- imens (UTA A-13381-—87) were taken by day from beneath rocks along a small stream; one male (UTA A-13388) was calling from a rock in this stream at night. Other species of anurans taken from this region are Bufo occidentalis, Eleutherodactylus nitidus, and Scaphiopus multiplicatus. Life history.—Little is known about the life history of H. xera. The advertisement call, eggs, and tadpole remain undescribed. All male specimens reported herein have 408 well developed nuptial excrescences and may be in breeding condition; however, their vo- cal sacs are not distended. The oviducts of the female paratopotype contain many well- developed ova. Etymology.—The specific name is de- rived from the Greek xeros meaning dry, in allusion to the desert habitat of this species. Morphometric analysis. —We performed a stepwise DFA of 11 morphometric vari- ables (see Materials and Methods) on adult males of H. sumichrasti (Chiapas, n = 16; Oaxaca: Portillo Nejapa, m = 13; Oaxaca: Mitla, n = 14—populations combined for analysis), H. smaragdina (n = 18), H. xera (n = 7), and H. chimalapa (n = 8). Mor- phometric variables 1, 2, 3, 5, and 10 varied significantly among groups (F < 0.001) in the stepwise analysis and were included in the canonical discriminant analysis. Group means were different (F < 0.001) at each step in the five-step model and 100 percent of the variation was displayed on three ca- nonical axes; the first two axes displayed 95 percent of the variation (CAN I, CAN II; Fig. 5). The standardized (pooled within group variances) coefficients for the canon- ical variables were: CAN I (CAN ID) = 1: =,62 COD), 22 O95 (0.78), 33 =O77/ G 0102) 5126370260) 03 O17 (GOF36): All individuals of H. xera, H. chimalapa, and H. smaragdina were correctly classified in the jackknifed classification matrix of the five-step model. Three individuals of H. sumichrasti were misclassified as H. xera and one as H. chimalapa. Hyla smaragdina is well separated from the other species along CAN I and H. chi- malapa is well separated from the other spe- cies along both axes. Hyla sumichrasti and H. xera are significantly different in this analysis, albeit weakly separated from one another along the first two canonical axes; although these two species are similar in overall proportions (Fig. 5), they are clearly distinguishable on the basis of other exter- nal characters. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Acknowledgments This paper benefitted from the comments of W. E. Duellman, L. Trueb, D. Kizirian, G. Ten Eyck, A. Lathrop, and K. Toal. A. Nieto assisted with the translation of the resumen. J. Simmons assisted in the prep- aration of photographs; Fig. 1 was repro- duced from a photograph by W. W. Lamar. JRM received funds to work at the CAS from the Stearns student travel grant pro- gram. We thank J. Gauthier (CAS) for the loan of specimens from that collection. Literature Cited Breedlove, D.E. 1973. The phytogeography and veg- etation of Chiapas (Mexico). Pp. 149-165 in A. Graham, ed., Vegetation and vegetational his- tory of Northern Latin America. Elsevier Sci- entific Publishing Co., Amsterdam. Caldwell, J. 1974. A re-evaluation of the Hyla bis- tincta species group, with descriptions of three new species (Anura: Hylidae).— Occasional Pa- pers of the Museum of Natural History, Uni- versity of Kansas 28:1-37. Campbell, J. A. 1984. A new species of Abronia (Sau- ria: Anguidae) with comments on the herpeto- geography of the highlands of southern Mexi- co.— Herpetologica 40(4):373-381. Duellman, W. E. 1970. The hylid frogs of Middle America.— Monograph of the Museum of Nat- ural History, University of Kansas 1:1—-753. Leopold, A. S. 1950. Vegetation zones of Mexico.— Ecology 31:507-518. Myers, C. W., & W. E. Duellman. 1982. A new spe- cies of Hyla from Cerro Colorado, and other tree frog records and geographical notes from western Panama.—American Museum Novi- tates 2752:1-25. (JRM) Museum of Natural History and Department of Systematics and Ecology, The University of Kansas, Lawrence, Kan- sas 66045-2454, U.S.A.; (JAC) Department of Biology, The University of Texas at Ar- lington, Arlington, Texas 76019, U.S.A. Appendix | Comparative material examined (all from Mexico): Hyla smaragdina: Sinaloa: 1.6 km E Santa Lucia, 1280 m (KU 68719); Santa Lucia, 1097 m (KU 75295- VOLUME 107, NUMBER 2 333); 2.2 km NE Santa Lucia, Hwy 40, 1157 m (KU 78380-82). Hyla sumichrasti: Chiapas: Linda Vista, ca 2 km NW Pueblo Nuevo Solistahuacan, 1675 m (KU 57857- 85). Guerrero: Sierra Madre del Sur, 3.1 km SW Rio Santiago on road to Atoyac, 762 m (CAS 143116- 409 143149). Oaxaca: 14 km E El Camaron, 1240 m (KU 10093446); 6.0 km N Mitla (UTA A-2827, 2881-95); 8.8 km N Mitla (UTA A-2916-19); 6.4 km E Mitla (UTA A-2920-29, 2931-33, 3255-56, 3284): 8.8 km E Mitla (UTA A-4673-77); 6.0 km ENE Mitla (UTA A-5889-90). PROC. BIOL. SOC. WASH. 107(2), 1994, pp. 410-416 A NEW FOSSORIAL SNAKE OF THE GENUS GEOPHIS (REPTILIA: SERPENTES: COLUBRIDAE) FROM THE CORDILLERA DE TALAMANCA OF COSTA RICA Karen R. Lips and Jay M. Savage Abstract. —Geophis talamancae, a new species of colubrid snake from south- central Costa Rica is described. The new form belongs to the sieboldi group, which includes five Mexican species, G. nasalis (Guatemala), G. hoffmanni (Honduras to western Panama), G. ze/edoni (central Costa Rica), G. nigroalbus (eastern Panama to Colombia), and G. brachycephalus (northern Costa Rica, Panama and Colombia). We reallocate Geophis betaniensis of Colombia, pre- viously referred to the championi group, to the sieboldi group; present scale counts of G. godmani from populations geographically intermediate to those previously known; comment on distinguishing between Geophis and Atractus on the basis of chin shields and temporal scales; and present a key to the Geophis species of Lower Central America and Colombia. Resumen.—En este trabajo se describe a Geophis talamancae, una nueva especie de serpiente colubrida del centro-sur de Costa Rica. Esta nueva forma pertenece al grupo sieboldi, el cual incluye a 5 especies mexicanas y a G. nasalis (Guatemala), G. hoffmanni (Honduras a Panama occidental), G. zeledoni (Costa Rica central), G. nigroalbus (Panama oriental hasta Colombia) y G. brachy- cephalus (norte de Costa Rica, Panama, y Colombia). Se reasigna a Geophis betaniensis de Colombia, previamente en el grupo championi, al grupo sieboldi; se proporcionan numeros de escamas de G. godmani de poblaciones inter- medias geograficamente a las conocidas anteriormente y se comenta sobre la distincion entre Geophis y Atractus con base en las escamas geneiales y tem- porales. Finalmente se presente una clave para las especies de Geophis del sur de Centroamérica y Colombia. A single example of a rather nondescript snake of the genus Geophis was collected during a survey of the herpetofauna of the Zona Protectora Las Tablas of the Reserva de la Biosfera la Amistad of Costa Rica, near the Costa Rica—Panama border by the se- nior author in 1992. A comparison with other Geophis confirms that the unique type appears to represent a population that may be called: Geophis talamancae, new species Fig. | Holotype.—CRE 5343, an adult female from Costa Rica: Puntarenas Province: Canton Coto Brus: Zona Protectora Las Tablas, Finca Jaguar, 1800 m elevation; taken 1 Sep 1992 by Karen R. Lips. Etymology.—The species name is de- rived from the Cordillera de Talamanca, the mountain range from which the specimen was collected. Definition.—The features listed below characterize the species and follow the for- mat used by Downs (1967) in his revision of the genus: 1) dorsal scale rows in 15-15- 15 rows, strongly keeled on posterior half of body; 2) no anterior temporal; 3) one supraocular and one postocular scale; 4) snout blunt and shovel-shaped, rostral bare- VOLUME 107, NUMBER 2 ly projecting posteriorly between interna- sals; 5) dorsal surfaces of body and of head uniform iridescent black; 6) venter white with black bands on posterior edges of scutes. Diagnosis. —The new form is a member of the sieboldi group (Downs 1967), previ- ously represented by 12 species ranging from Mexico to Colombia and including four that occur in Costa Rica and Panama. Geophis talamancae may be distinguished from G. brachycephalus by having the dorsal scales on the anterior half of the body smooth and a uniform dorsal coloration (in brachyceph- alus the dorsal scales are keeled except on the neck and there is often a distinct dorsal pattern of light, usually red, bars, spots and/ or stripes). The new form differs from both Geophis zeledoni and G. hoffmanni in the keeling of the dorsal scales since these two forms have keels only on dorsal scales above the vent. Unlike G. nigroalbus, which has the postocular and supraocular separated by an anterior projection of the parietal, the supraocular is in broad contact with the postocular in G. talamancae. The new spe- cies is distinguished from G. betaniensis of west-central Colombia (features for that form in parentheses) by having keeled pos- terior dorsal scales (smooth dorsal scales) and one (two) postocular. Finally, it differs most obviously from the several species of the G. championi group, all of which are endemic to Costa Rica and/or Panama, in the shape of the head and associated fea- tures of scutellation. Geophis talamancae has an elongate snout that is rounded in dorsal outline, a rostral that barely extends posteriorly between the internasals, and a short postnasal that is higher than wide (Fig. 1). In the four members of the championi group (G. championi, G. downsi, G. god- mani, and G. ruthveni), the elongate snout is pointed, the rostral separates the inter- nasals for much of their length and the post- nasal is broad, the width being at least 75% of its height. General characteristics. Head not dis- tinct from neck; snout elongate, rounded in dorsal profile; rostral not extending poste- 411 riorly between internasals; its length from above about ¥% its distance from frontal; internasals large, rounded anteriorly, slight- ly shorter than suture with prefrontal; pre- frontals short, their median suture about *4 length of frontal; frontal slightly wider than long, quadrangular, in contact with prefron- tals, supraoculars, and parietals, distinctly angulate anteriorly; parietals moderately long, broad, their median suture almost equal to length of frontal; parietal does not contact prefrontal above middle of orbit, but meets the supraocular and postocular scales. There is one postocular and one su- praocular scale on each side of the head (Fig. 1). Nasal divided, postnasal about the same size as prenasal, their combined length about 70% length of loreal; loreal relatively long, slightly more than 2 length of snout, slightly more than 2 times eye diameter; eye small, contained about 4'4 times in snout length (tip of snout to anterior border of eye), its vertical diameter about equal to distance from supralabials; supralabials 6-6, 3 and 4 in contact with orbit on both sides, 5th in contact with parietal; anterior temporal di- rectly above 6th supralabial, not fused with nuchals along parietal margin. Mental rounded anteriorly, definitely broader than long, separated from chin shields by first pair of infralabials; infrala- bials 6-6, first 3 in contact with anterior chin shields; anterior chin shields slightly longer than broad, longer than posterior chin shields; posterior chin shields short, in con- tact anteriorly, diverging posteriorly; 3 gu- lars separate chin shields from first ventral. Dorsal scale rows 15-15-15, keeled on posterior half of body and tail; posterior dorsal scales without discernible apical pits. Ventrals 138; anal entire; subcaudals 33. Ventrals + subcaudals 171. Standard length (snout-to-vent) 185 mm, tail length 33 mm; tail length 16 percent of total length. Coloration. — Dorsal surfaces of head and body uniform dark charcoal grey to black. Head color extends ventrally to supralabi- als, infralabials, mental, and chin shields. VOLUME 107, NUMBER 2 Gular area immaculate creamy white, be- coming flecked with black on first ventral and becoming progressively more flecked along anterior edge of scutes, until only a very thin white posterior border remains on scutes near vent. Subcaudals completely black, without white markings. Distribution. —Known only from the type locality in the Lower Montane Rainforest (Holdridge 1967, Tosi 1969) in the Cordil- lera de Talamanca of Costa Rica near the Panama border at 1800 m elevation. Remarks. —The type specimen was col- lected from beneath a series of saplings whose roots were enclosed in plastic bags that had recently (within 4 months) been transported from the region of the central Cordillera de Talamanca along the Carre- tera Interamericana, about 100 km north- west of the type locality. There is a slight possibility that this snake might have been brought to the Las Tablas area in this ship- ment. However, a second specimen prob- ably of the same species (white belly with distinct black bands on ventral scutes, uni- form black dorsum) was collected from the Las Tablas area earlier in the year but es- caped before an accurate identification could be made. Four specimens of Geophis godmani were also collected from the Las Tablas area, rep- resenting an intermediate population be- tween the two previously known but dis- junct localities: a population from the Cordillera Central and extreme northern edge of the Cordillera de Talamanca in Cos- ta Rica, and a population 250 km ESE of Las Tablas on Volcan Baru in Chiriqui, Panama. The Panamanian specimens are represented only by heads and necks (Downs 1967) so variation in ventral and subcaudal counts from the southern part of the range was unknown. Because G. brachycephalus and G. hoffmanni have distinctly lower seg- mental counts on Volcan Baru than in Costa Rica, Downs (1967) predicted that Pana- manian G. godmani would vary similarly. All four specimens have 15-15-15 scale 413 rows; no postocular, no supraocular, and no temporal scales. The number of ventrals for the three males (CRE 5344, 5319, 5072) ranges from 139-141; subcaudals 34-38; ventrals + subcaudals 173-179. The single female specimen (CRE 5337) has 140 ven- trals, 27 subcaudals, and 167 ventral + sub- caudals. These values do not differ substan- tially from those of specimens from the north in Costa Rica contra Downs’ (1967) pre- diction. Geographical variation in G. god- mani exists, however, in the amount of ven- tral pigmentation between those specimens from the area of the Barva-Poas volcanoes (venter almost uniformly bright yellow in color) and those from Volcan Turrialba south through the Cordillera de Talamanca to Las Tablas (bright yellow color of venter limited to anterior edge of scutes, posterior edge black). It is clear from the differences in ventral color that the escaped snake men- tioned above was not G. godmani. Relationships. —The new species is re- ferred to the Geophis sieboldi group (Downs 1967) on the following basis: snout long, projecting well beyond lower jaw, rounded in dorsal outline; rostral not produced pos- teriorly between internasals; internasals short, their greatest length 33-62% of suture between prefrontals; postnasal short, width about 50% of height; prefrontals and loreals elongate; no anterior temporal; rounded mental; maxillary extends forward to suture between second and third supralabials, with 14 subequal teeth, tip of maxillary toothless; posterior end of maxillary tapering to a blunt point. The sieboldi group as understood by Downs (1967) in his generic revision in- cluded four Mexican species (Geophis pe- tersi, G. russatus, G. sallei, and G. sieboldi), one Guatemalan form (G. nasalis), one Nic- araguan endemic (G. dunni) and G. brachy- cephalus (Costa Rica to Colombia), G. hoff- manni (Honduras to western Panama), G. nigroalbus (eastern Panama to Colombia) and G. zeledoni (Costa Rica). Downs pro- vided detailed descriptions of most of these 414 forms but gave belated recognition to G. nigroalbus and G. russatus only in footnotes (p. 146 and p. 138, respectively). Campbell and Murphy (1977) added another Mexican species (G. pyburni) to the group. Within the genus, G. talamancae most closely re- sembles G. nigroalbus but differs most ob- viously in the relation of the parietal to the supraocular and postocular (bordering them posteriorly in talamancae, separating them in nigroalbus). Restrepo & Wright (1987) on the occa- sion of describing Geophis betaniensis from Colombia expressed difficulty in accepting Downs’ (1967) definitions of species groups and referred their new form to the cham- pioni group. This decision was reached in part because G. betaniensis keyed out to G. championi in Downs’ key (couplet 29) to the genus. When they used Savage’s (1981) key to Costa Rican and Panamanian Geo- phis, they reached a couplet (number 4) that distinguished between G. hoffmanni and G. zeledoni (sieboldi group species) and this seems to be the basis for their puzzlement Over species group definitions. Both keys (Downs 1967, Savage 1981) are artificial ones based upon the most obvious characters of the included species and are designed for field identifications. No at- tempt was made by either author to design a key in which presumably related form (i.e., members of the same species group) were placed together. Downs (1967), contrary to Restrepo & Wright (1987), provided unambiguous def- initions of the seven species groups that he recognized within Geophis. Members of the championi group differ from G. betaniensis (features for that species in parentheses) in the following characteristics: snout elon- gate, pointed (rounded); rostral produced posteriorly between internasals (barely pro- jecting between internasals); internasals elongate, greatest length 67—-100% of pre- frontal suture (50%); postnasal long, width at least 75% of height (postnasal short, width about 50% of height); mental acuminose PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON (rounded). These differences clearly pre- clude inclusion of G. betaniensis in the championi group. Comparison of the features of head shape and scutellation of Geophis betaniensis with those of the other species groups of Geophis (Downs 1967) show a complete concor- dance between the Colombian species and members of the sieboldi group (see list of characteristics at the beginning of this sec- tion). In terms of skeletal and dentitional features the championi and sieboldi groups are very similar except that in the latter there 1s no tooth on the tip of the maxillary (pres- ent in the former) and the hemipenes of the championi group differ in lacking capita- tion. Unfortunately, features of the hemi- penes were not included by Restrepo & Wright (1987) in their original description because the only specimens referred to G. betaniensis are both females. Nevertheless we believe that evidence from physiognomy (general shape of the head) and scutellation support the inclusion of G. betaniensis in the sieboldi group. Because Colombian species of Geophis are sympatric with members of the superficially similar genus Atractus, another fossorial group, Restrepo & Wright (1987) discussed features of external morphology that may be utilized to distinguish between them. Af- ter some discussion with Frances J. Irish, who is undertaking a systematic revision of Atractus, they concluded that the presence (in Atractus) and absence (in Geophis) of an anterior temporal scale and the number of chin shields (two pairs in Geophis and one pair in Atractus) are diagnostic for those species found in Central and South Amer- ica. Unfortunately, the situation is more com- plicated than these authors suggest. As pointed out by Savage (1960) and Downs (1967), presence or absence of the anterior temporal exhibits both interspecific and some intraspecific variability in both gen- era; those Geophis characteristically having an anterior temporal are confined to Mex- VOLUME 107, NUMBER 2 415 Fig. 2. ico, and most examples of Atractus consis- tently have one. Savage (1960) had previously noted that Geophis and Atractus from the area of geo- graphic overlap in Panama and Colombia could be distinguished on the basis of the chin shield feature. Downs (1967) however, questioned the utility of this character since it is difficult in some cases to distinguish the posterior chin shield from the adjacent gular scales in Geophis, which approaches the condition found in Atractus. Nevertheless, although the posterior pair of chin shields are often small and usually separated by a median gular scale, there is no ambiguity in using this feature to separate lower Central and South American Geophis from Atractus (Fig. 2). Key to Species of Geophis from Lower Central America and Colombia la. Supraocular shields present .... 2 1b. No supraocular scales ......... 10 yae AND): 3a. 3b. 4a. Ab. Sa. Sb. 6a. Diagrammatic differences in chin shields between A) Geophis and B) Atractus. Two postoculars ............. mb caret atl p G. betaniensis (Colombia) Onespostoculanme se seen. 3 Uppermost dorsal scales keeled at least on posterior half of body andatamlwewe a, ei. cose ns oe ae 4 Dorsal and caudal scales smooth or smooth except for some faintly keeled scales above vent ...... 8 Dorsal scales on body (exclusive of neck) and tail distinctly keeled 5 Dorsal scales on anterior half of body smooth Dorsal scales in 15 rows; dorsum dark, often with light lateral blotches, crossbands, or stripes . ...G. brachycephalus (Costa Rica to Panama) Dorsal scales in 17 rows; dorsum light with dark blotches or sad- Giese hee G. dunni (Nicaragua) Postocular and supraocular in contact, excluding parietal from Margin or orbit 416 6b. Postocular and supraocular sep- arated by extension of parietal thatenmeetsronbitae es eee De ea) oe ee G. nigroalbus (Colombia) 7a. Snout pointed; rostral markedly produced posteriorly between in- ternasals; mental pointed ...... a PONE ee G. ruthveni (Costa Rica) 7b. Snout rounded; rostral barely produced posteriorly between in- ternasals; mental rounded ..... oad ely er G. talamancae (Costa Rica) 8a. Five or fewer supralabials ..... .G. hoffmani (Honduras to Panama) 8b. Six or more supralabials 9a. Snout pointed; rostral markedly produced posteriorly between in- ternasals; mental pointed ante- riorly; ventrals plus subcaudals 156-158 .. G. championi (Panama) 9b. Snout rounded; rostral barely produced posteriorly between in- ternasals; mental rounded; ven- trals plus subcaudals 180-191 Sy Set 2 pe G. zeledoni (Costa Rica) Uppermost dorsal scales keeled at least on posterior third of body and on tail; ventrals 122-133; subcaudals 41-46 ............ A RAES AO Pr G. downsi (Costa Rica) Dorsal scales smooth; ventrals 132-145; subcaudals 26-36 G. godmani (Costa Rica to Panama) 10a. 10b. Acknowledgments We would like to thank the family of Mi- guel Sandi C. for their assistance to KRL. We would also like to thank Juan Diego Alfaro of the Reserva Biosfera Amistad and Lic. Miguel Rodriguez R. of the Servicio de Parques Nacionales del Ministerio de Re- cursos Naturales, Energia, y Minas de Costa Rica for issuing permits for work within the Zona Protectora Las Tablas of the Amistad PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Biosphere Reserve, and to Marybel Soto Ramirez of the Organization for Tropical Studies for expediting permits. David Auth (Florida Museum of Natural History) pro- vided specimens for comparison. We are grateful to M. Donnelly for comments on the manuscript and to V. Sosa for providing a Spanish translation. KRL’s field work was supported by the American Society of Ich- thyologists and Herpetologists’ Gaige Fund, Society for the Study of Amphibians and Reptiles Grants-in-Herpetology, Organiza- tion for Tropical Studies Pew/Mellon post- course grant, University of Miami Tropical Biology Fellowship, American Museum of Natural History Roosevelt Fund, Explorer’s Club, and the National Geographic Society. Literature Cited Campbell, J. A., & J.B. Murphy. 1977. Anew species of Geophis (Reptilia, Serpentes, Colubridae) from the Sierra de Coalcoman, Michoacan, Meéxi- co.—Journal of Herpetology 11:397—403. Downs, F. L. 1967. Intrageneric relationships among colubrid snakes of the genus Geophis Wagler.— Miscellaneous Publications of the Museum of Zoology of the University of Michigan 131:1- 193. Holdridge, L. R. 1967. Life zone ecology. 2nd ed. Tropical Science Center, San Jose, Costa Rica. Restrepo, J. H., & J. W. Wright. 1987. A new species of the colubrid snake genus Geophis from Co- lombia.—Journal of Herpetology 21:191-196. Savage, J. M. 1960. A revision of the Ecuadorian snakes of the colubrid genus Atractus.—Mis- cellaneous Publications of the Museum of Zo- ology, University of Michigan 112:1—-86. 1981. New species of the secretive colubrid snake genus Geophis from Costa Rica.—Copeia 1981:549-553. Tosi, J. A., Jr. 1969. Mapa Ecologico. Tropical Sci- ence Center, San Jose, Costa Rica. Department of Biology, University of Mi- ami, P.O. Box 249118, Coral Gables, Flor- ida 33124, U.S.A. PROC. BIOL. SOC. WASH. 107(2), 1994, pp. 417-418 INTERNATIONAL COMMISSION ON ZOOLOGICAL NOMENCLATURE % The Natural History Museum Cromwell Road London, SW7 SBD, U.K. Tel. 071-938 9387 Applications published in the Bulletin of Zoological Nomenclature The following Applications were published on 30 March 1994 in Vol. 51, Part 1 of the Bulletin of Zoological Nomenclature. Comment or advice on these Applica- tions is invited for publication in the Bulletin and should be sent to the Executive Secretary, I.C.Z.N., % The Natural History Museum, Cromwell Road, London SW7 SBD, U.K. Case No. 2886 Doris grandiflora Rapp, 1827 (currently Dendrodoris grandiflora; Mollusca, Gastropoda): proposed conservation of the specific name. 2859 Johnstonia Quatrefages, 1866 (Annelida, Polychaeta): proposed conserva- tion. 2889 Mastotermes darwiniensis Froggatt, 1897 and Termes meridionalis Froggatt, 1898 (currently Amitermes meridionalis) (Insecta, Isoptera): pro- posed retention of neotypes following rediscovery of syntypes. 2713 COLYDIIDAE Erichson, 1842 (Insecta, Coleoptera): proposed precedence over CERYLONIDAE Billberg, 1820 and ORTHOCERINI Blan- chard, 1845 (1820); and Cery/on Latreille, 1802: proposed conser- vation of Lyctus histeroides Fabricius, 1792 as the type species. 2783 Cryptophagus Herbst, 1792, Dorcatoma Herbst, 1792, Rhizophagus Herbst, 1793 and Colon Herbst, 1797 (Insecta: Coleoptera): proposed con- servation as the correct spellings, and proposed conservation of Lyctus bipustulatus Fabricius, 1792 as the type species of Rhizo- phagus. 2861 ELMIDAE Curtis, 1830 and E/mis Latreille, 1802 (Insecta, Coleoptera): proposed conservation as correct spelling and of feminine gender respectively. 2858 Hydrophoria Robineau-Desvoidy, 1830 (Insecta, Diptera): proposed desig- nation of Musca lancifer Harris, [1780] as the type species. 2881 Sicus Scopoli, 1763 and Myopa Fabricius, 1775 (Insecta, Diptera): proposed conservation by the designation of Conops buccata Linnaeus, 1758 as the type species of Myopa. 2835 Allestes Miller & Troschel, 1844 (Osteichthyes, Characiformes): conserva- tion proposed. 418 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Opinions published in the Bulletin of Zoological Nomenclature The following Opinions were published on 30 March 1994 in Vol. 51, Part 1 of the Bulletin of Zoological Nomenclature. Copies of these Opinions can be obtained free of charge from the Executive Secretary, I.C.Z.N., % The Natural History Mu- seum, Cromwell Road, London SW7 5BD, U.K. Opinion No. 1752 Zanclea costata Gegenbaur, 1856 (Cnidaria, Hydrozoa): generic and specific names conserved. 1753. Gebia major capensis Krauss, 1843 (currently Upogebia capensis, Crustacea, Decapoda): neotype replaced, so conserving the usage of G. capensis and also that of G. africana Ortmann, 1894 (currently Upogebia africana). 1754 Histoire abrégée des insectes qui se trouvent aux environs de Paris (Geoffroy, 1762): some generic names conserved (Crustacea, Insecta). 1755 Podisus Herrich-Schaeffer, 1851 (Insecta, Heteroptera): P. vittipennis Her- rich-Schaeffer, 1851 designated as the type species. 1756 ANTHRIBIDAE Billberg, 1820 (Insecta, Coleoptera): given precedence over CHORAGIDAE Kirby, 1819. 1757 Cryptus Fabricius, 1804 and CRYPTINAE Kirby, 1837 (Insecta, Hymenoptera): conserved. 1758 Véipio Latreille, 1804 (Insecta, Hymenoptera): Agathis longicauda Boheman, 1853 designated as the type species. 1759 Acamptopoeum Cockerell, 1905 (Insecta, Hymenoptera): Camptopoeum sub- metallicum Spinola, 1851 designated as the type species. 1760 Rhipidocystis Jackel, 1901 (Echinodermata, Eocrinoidea): R. baltica Jaekel, 1901 designated as the type species. 1761 Filimanus Myers, 1936 (Osteichthyes, Perciformes): Filimanus perplexa Feltes, 1991 designated as the type species. 1762 Cynolebias opalescens Myers, 1942 and C. splendens Myers, 1942 (Osteich- thyes, Cyprinodontiformes): specific names conserved. 1763 Megophrys montana Kuhl & van Hasselt, 1822 (Amphibia, Anura): generic and specific names placed on Official Lists, and Leptobrachium par- vum Boulenger, 1893 (currently Megophrys parva): specific name conserved. Anas arcuata Horsfield, 1824 (currently Dendrocygna arcuata; Aves, Anser- iformes): specific name conserved. PROC. BIOL. SOC. WASH. 107(2), 1994, p. 419 REVIEWERS The following people reviewed manuscripts for the Proceedings in 1993. C. C. Baldwin, R. C. Banks, F. M. Bayer, A. Berta, J. A. Blake, R. W. Bouchard, T. E. Bowman, G. A. Boxshall, J. M. Bradford, R. O. Brinkhurst, M. R. Browning, N. L. Bruce, R. C. Brusca, E. Campos, M. R. Campos, A. Carvacho, F. A. Chace, Jr., T.-Y. Chan, A. Cheetham, B. Cheroff, K. Coates, J. Corliss, M. M. Criales, G. B. Corbet, N. Cumberlidge, E. B. Cutler, N. J. Cutler, M. Dardeau, M. O. Dillon, R. Drewes, D. Dudgeon, R. J. Emry, K. Fauchald, D. L. Felder, F. D. Ferrari, C. Ferraris, F. Fiers, R. D. Fisher, K. Fitzhugh, O. S. Flint, Jr., A. Fosshagen, C. Fransen, R. Franz, R. J. Gagné, J. Garcia-Raso, A. L. Gardner, J. S. Garth, C. J. Glasby, W. Goldberg, M. Gomon, G. R. Graves, M. J. Grygier, J. Haig, G. Hall, C. W. Hart, Jr., W. Hartman, G. Hartmann, K. Hayashi, L. R. Heaney, R. H. Heard, P. Heideman, M. E. Hendrickx, R. Hershler, R. P. Higgins, B. Hilbig, D. Hillis, H. H. Hobbs, Jr., H. H. Hobbs III, E. Hoberg, D. Holdich, L. B. Holthuis, R. W. Holzenthal, K. Hulsemann, W. Hummon, P. Hutchings, H. F. James, R. F. Jezerinac, D. A. Jones, D. Kathman, B. Kensley, P. Kier, Y. Kikuchi, H. Kishimoto, N. Klein, K. F. Koopman, F. Krapp, R. K. Kropp, J. D. Kudenov, D. Laubitz, M. Ledoyer, D. B. Lellinger, R. Lemaitre, D. Lipscomb, M. Litvaitis, J. Lundberg, A. Mackie, E. Macpherson, C. Magalhaes, R. B. Manning, J. C. Markham, J. W. Martin, W. N. Mathis, R. W. McDiarmid, P. A. McLaughlin, E. Moll, J. C. Morse, H.-G. Muller, T. A. Munroe, G. G. Musser, D. Nelson, T. Newberry, P. K. L. Ng, N. Ngoc-Ho, J. N. Norenburg, S. L. Olson, L, R. Parenti, B. D. Patterson, C. Patterson, D. L. Pawson, R. Paynter, Jr., T. Peterson, » M. H. Pettibone, T. Pietsch, F. Pleijel, G. Pretzmann, J. Pruski, J. W. Reid, J. V. Remsen, Jr., R. P. Reynolds, N. Riser, G. Rodriguez, G. Rouse, K. Ruetzler, L. Sandberg, J. M. Savage, S. Schaefer, A. E. Smalley, L. Soto, W. C. Starnes, D. W. Steadman, G. S. Steyskal, G. W. Stocker, J.-O. Stromberg, J. D. Thomas, F. G. Thompson, P. Tongiorgi, M. Tiirkay, J.-W. Wagele, D. Wake, T. C. Walter, K. Warheit, S. Werman, M. J. Wetzel, D. Wiedenfeld, A. B. Williams, D. E. Wilson, G. D. Wilson, L. D. Wilson, J. Winston, R. Winterbottom, N. E. Woodley, K. Wouters, H. Yeatman, G. R. Zug. -" ol y a , ecopyrntit TH eye a7 : ees to (bagi eT mee) ot aud Nie pene ee We yin i yay (shia e net es Wy re a wae A oe r 7A he 4 wD inthdl ee ahha"? a: eet oh pera” a Cat i 1 PTD mad, iy i peg uh Wy a ave wi oe - iv 7 sire ant ii ; = 16 pa a mt ewuniinds ph Yh ’ ba Shah ile Ta \ oe ie eh ii 1 Ay * ch ced Fac LO Eagan dant es i) om bp yet ee, nhs a ete : ite =a tre 1? causing a y Lah, mt A ve nae kd wit + 5 ; Ane vt if + ‘a ae “4 4 Ws, a why | su) ae} hat" of 4 Perret 7 i fia 2] ri 4 5 Mim : ar Fae ealbal Oni or INFORMATION FOR CONTRIBUTORS Content.—The Proceedings of the Biological Society of Washington contains papers bearing on systematics in the biological sciences (botany, zoology, and paleontology), and notices of business transacted at meetings of the Society. Except at the direction of the Council, only manuscripts by Society members will be accepted. Papers are published in English (except for Latin diagnoses/descriptions of plant taxa), with an Abstract in an alternate language when appropriate. : Submission of manuscripts.—Submit three copies of each manuscript in the style of the Proceedings to the Editor, complete with tables, figure captions, and figures (plus originals of the illustrations). Mail directly to: Editor, Proceedings of the Biological Society of Washington, National Museum of Natural History NHB-108, Smithsonian Institution, Washington, D.C. 20560. (Do not submit manuscripts to an associate editor). 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 and diagnoses. The style for the Proceedings is described in “GUIDELINES FOR MANUSCRIPTS for Publications of the BIOLOGICAL SOCIETY OF WASHINGTON” a supplement to Volume 103, number 1, March 1990. Authors are encour- aged to consult this article before manuscript preparation. Copies of the article are available from the editor or any associate editor. The establishment of new taxa must conform with the requirements of appropriate inter- national codes of nomenclature. Decisions of the editor about style also are guided by the General Recommendations (Apendix E) of the International Code of Zoological Nomenclature. When appropriate, accounts of new taxa must cite a type specimen deposited in an institutional collection. Review.—One of the Society’s aims is to give its members an opportunity for prompt pub- lication of their shorter contributions. Manuscripts are reviewed by a board of Associate Editors and appropriate referees. Proofs. — Authors will receive first proofs and original manuscript for correction and approval. Both must be returned within 48 hours to the Editor. Reprint orders are taken with returned proofs. Publication charges.— Authors are required to pay full costs of figures, tahles, changes in proofs ($3.00 per change or revision), and reprints. Authors are also asked to assume costs of page-charges. The Society, on request, will subsidize a limited number of contributions per volume. If subsidized manuscripts result in more than 12 printed pages, the additional pages must be financed by the author(s). Multiple authorship will not alter the 12 page limit (each author will be viewed as having used his/her 12 subsidized pages). Payment of full costs will facilitate speedy publication. Costs. — Printed pages @ $60.00, figures @ $10.00, tabular material @ $3.00 per printed inch per column. One ms. page = approximately 0.4 printed page. CONTENTS Three new species of ciliate in the genera Pseudocohnilembus, Pleuronema, and Urotricha (Ciliophora) Gregorio Fernandez-Leborans and Apolonia Novillo Morphological variability in warm-temperate and subtropical populations of Macrodasys (Gas- trotricha: Macrodasyida: Macrodasyidae) with the description of seven new species Wayne A. Evans A new species of Elaphoidella (Crustacea: Harpacticoida) closely related to E. bidens (Schmeil) and the genus Attheyella from Nepal Teruo Ishida Monstrilla elongata, a new monstrilloid copepod (Crustacea: Copepoda: Monstrilloida) from a reef lagoon of the Caribbean coast of Mexico E. Suarez-Morales Annina mannai, a new isopod from the Ganges River, West Bengal (Crustacea: Isopoda: Cirolanidae) Marilyn Schotte Redescription of Jais elongata Sivertsen & Holthuis, 1980, from the South Atlantic Ocean (Crustacea: Isopoda: Asellota) Brian Kensley A new genus and species of cirolanid isopod from the western Indian Ocean (Crustacea: Peracarida) Brian Kensley and Marilyn Schotte A new species of Palaemonetes (Crustacea: Decapoda: Palaemonidae) from northeastern Mexico Ned E. Strenth A revision of the type material of some species of Hypolobocera and Ptychophallus (Crustacea: Decapoda: Pseudothelphusidae) in the National Museum of Natural History, Washington, D.C., with descriptions of a new species and a new subspecies Gilberto Rodriguez Parapinnixa cubana, a new pea crab from Cuba (Crustacea: Brachyura: Pinnotheridae) Ermesto Campos Petrolisthes extremus, a new porcelain crab (Decapoda: Anomura: Porcellanidae) fromthe — Indo-west Pacific Roy K. Kropp and Janet Haig A new freshwater crab of the genus Geothelphusa (Crustacea: Decapoda: Brachyura: Potamidae) from Kagoshima Prefecture, southern Kyushu; Japan Hiroshi Suzuki and Eiji Tsuda Aegla pewenchae, a new species of central Chilean freshwater decapod (Crustacea: Anomura: Aeglidae) Carlos G. Jara Description of the ghost shrimp Eucalliax mcilhennyi, new species, from south Florida, with reexamination of its known congeners (Crustacea: Decapoda: Callianassidae) Darryl L. Felder and Raymond B. Manning Pinnixa scamit, a new species of pinnotherid crab (Decapoda: Brachyura) from the continental slope off California Joel W. Martin and Deborah L. Zmarzly Studies of Neotropical caddisflies, L: The description of Cerasmatrichia, new genus, a relative of Alisotrichia, with descriptions of new and old species and the larva (Trichoptera: Hy- droptilidae) Oliver S. Flint, Jr., Steven C. Harris, and L. Botosaneanu A new hemichordate, Saccoglossus bromophenolosus (Hemichordata: Enteropneusta: Harmi- mantidae), from North America Gary M. King, Cem Giray, and Irv Kornfield Advertisement calls and relationships of Chilean frogs Eupsophus contulmoensis and E. in- sularis (Amphibia: Anura: Leptodactylidae) J. Ramon Formas and Lila Brieva Two new species of the Hyla sumichrasti group (Amphibia: Anura: Hylidae) from Mexico Joseph R. Mendelson III and Jonathan A. Campbell A new fossorial snake of the genus Geophis (Reptilia: Serpentes: Colubridae) from the Cordillera de Talamanca of Costa Rica Karen R. Lips and Jay M. Savage International Commission on Zoological Nomenclature Reviewers— 1993 22M 340 354 360 383 391 398 410 417 419 bh ROCEEDINGS — OF IHE BIOLOGICAL SOCIETY OF WASHINGTON THE BIOLOGICAL SOCIETY OF WASHINGTON 1994-1995 Officers President: Janet W. Reid Secretary: Carole C. Baldwin President-elect: Stephen D. Cairns Treasurer: T. Chad Walter Elected Council Robert J. Emry Susan L. Jewett Richard C. Froeschner Lynne R. Parenti Alfred L. Gardner F. Christian Thompson Custodian of Publications: Austin B. Williams PROCEEDINGS Editor: C. Brian Robbins Associate Editors Classical Languages: George C. Steyskal Invertebrates: Jon L. Norenburg Frank D. Ferrari Plants: David B. Lellinger Rafael Lemaitre Insects: Wayne N. Mathis Vertebrates: Thomas A. Munroe Membership in the Society is open to anyone who wishes to join. There are no prerequisites. Annual dues of $25.00 (for USA and non-USA addresses) include subscription to the Proceed- ings of the Biological Society of Washington. Library subscriptions to the Proceedings are: $40.00 for USA and non-USA addresses. Non-USA members or subscribers may pay an addi- tional $25.00 to receive the Proceedings by Air Mail. The Proceedings of the Biological Society of Washington (USPS 404-750) is issued quarterly. Back issues of the Proceedings and the Bulletin of the Biological Society of Washington (issued sporadically) are available. Correspondence dealing with membership and subscriptions should be sent to the Biological Society of Washington, P.O. Box 1897, Lawrence, Kansas 66044, U.S.A. Payment for membership 1s accepted in US dollars (cash or postal money order), checks on US banks, or MASTERCARD or VISA credit cards. Manuscripts, corrected proofs, editorial questions should be sent to the Editor, Biological Society of Washington, National Museum of Natural History, Smithsonian Institution, Wash- ington, D.C. 20560. Known office of publication: National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560. Printed for the Society by Allen Press, Inc., Lawrence, Kansas 66044 Second class postage paid at Washington, D.C., and additional mailing office. POSTMASTER: Send address changes to PROCEEDINGS OF THE BIOLOGICAL SOCI- ETY OF WASHINGTON, P.O. Box 1897, Lawrence, Kansas 66044. © This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper). PROC. BIOL. SOC. WASH. 107(3), 1994, pp. 421-428 DESCRIPTION OF A NEW SPECIES OF SUBFOSSIL EAGLE FROM MADAGASCAR: STEPHANOAETUS (AVES: FALCONIFORMES) FROM THE DEPOSITS OF AMPASAMBAZIMBA Steven M. Goodman Abstract. —Several bones recovered from subfossil deposits at Ampasam- bazimba, Madagascar, are described as a new species of eagle, Stephanoaetus mahery. These bones presumably date from the Holocene. S. coronatus, oc- curring on the African mainland, is the only extant member of this genus, and throughout much of its range feeds extensively on primates weighing up to about 12 kg. Bones of a remarkable assortment of lemurs have been recovered from Ampasambazimba and it is suggested that S. mahery may have preyed upon primates. Further, the legendary Rokh, a giant bird reputed to have occurred ori Madagascar, from the tales of Sinbad and Marco Polo, may have been derived from S. mahery. Résumé.—Une nouvelle espéce d’aigle, Stephanoaetus mahery, fait Pobjet d’une description réalisée 4 partir de divers ossements provenant des dép6ts subfossiles d’Ampasambazimba, Madagascar. L’Age estimé de ces ossements est de ’holocéne. Stephanoaetus coronatus, qui se rencontre sur le continent africain, est actuellement l’unique espéce connue dans ce genre. Cette espéce se nourrit communément de primates pesant environ 12 kg. De remarquables ossements de lémuriens ont été retrouvés 4a Ampasambazimba et il a est suggéré que S. mahery pourrait étre un prédateur de primates. Par ailleurs, la legende de l’oiseau rock, célébre oiseau-géant ayant été vécu 4 Madagascar dont on trouve des détails dans |’Histoire de Sinbad le marin et de Marco Polo pourait étre celle de S. mahery. Famintinana.—Taolambiby maro hita avy tao amin’ny farita misy taratsie- fan-taolambiby ao Ampasambazimba, Madagasikara, no noheverina fa kara- zamboromahery vaovao, Stephanoaetus mahery. Araka ny tombatombana mi- kasika ireo taolana ireo, dia efa hatramin’ny vanimpotoana jeolozika “Holocene” no nisian’izy ireo tao ambanin’ny tany. S. coronatus, hita any amin’ny kaoti- nanta afrikana, no hany karazana mitovy amin’io mbola velona ankehitriny izay mivelona ara-dalana amin’ny fihinanana karazana varika milanja sahabo ho 12 kg eo ho eo izay hitany ao amin’ny faritra iainany. Mbola nahitana karazan-taolambiby voafaritra ho varika ihany koa tao Ampasambazimba, noho izany azo heverina fa io S. mahery io dia nihaza sy nihinana karazam- barika. Ankoatr’izany, ilay angano mikasika an’ilay hoe ““Rokh” na ilay vo- rombe malaza niana tany Madagasikara, nalaina avy tao amin’ny Tantaran’i Sinbad sy Marco Polo dia mety tsy ho hafa fa’ity atao hoe S. mahery ity ihany. The modern bird fauna of Madagascar is course of the last few millennia the island well-known for its high level of endemism, has undergone drastic ecological change, a of the 201 extant resident species 105 (52%) portion of which clearly is human induced are endemic (Langrand 1990). During the (Perrier de la Bathie 1921, Battistini & Vér- 422 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Tarsometatarsus of Stephanoaetus mahery, new species, holotype, MAD 5491, Muséum National d@’Histoire Naturelle, Service de Paléontologie, Paris, left dorsal and right ventral views. in 1972, Burney 1987). In general, little is known about what effects these changes have had on the avifauna. The best known group of extinct birds on the island is the elephant birds (Family Aepyornithiformes) which consisted of at least seven species in two different genera (Brodkorb 1963). The exact sequence of events that led to their extinc- tion is unknown, but at least one species may have been extant until the turn of the 17th-century, about the same time Euro- peans arrived on Madagascar (Flacourt 1658, p. 165). Other smaller bird species are known to have gone extinct in the past few millennia (Milne Edwards & Grandidier 1895, Andrews 1897, Goodman & Ra- voavy 1993). One of the best known subfossil localities on Madagascar is Ampasambazimba, lo- cated about 85 km west of the capital city of Antananarivo. The site is at the edge of a former lake created by the damming of a river by a lava flow. Subsequently, the lava barrier was downcut, and the lake drained and successionally infilled to form a marsh (MacPhee et al. 1985). During excavations at Ampasambazimba a large number of bird bones were recovered, which until now re- mained largely unstudied. The majority of this material consists of waterbirds, but bones of raptorial birds were also recovered. Within this material are elements of a large eagle previously unknown to science and here it is proposed to call it: Stephanoaetus mahery, new species Figs. 1, 2 Holotype. —Left tarsometatarsus, collec- tions of the Laboratoire de Paléontologie, Museum National d’Histoire Naturelle, Paris, MAD 5491 (Figs. 1, 2). The element is complete except a portion of the hypo- tarsus is broken. Locality. — Ampasambazimba, Madagas- car, and accessioned in 1925 as part of a collection of over 750 specimens donated by “Gouvernment general de Madagascar Fig. 2. Comparison of Stephanoaetus coronatus (MNHN 1901.209) tarsometatarsus (top) and S. mahery (MNHN MAD 5491) tarsometatarsus (bottom). 424 par l’intermediaire de M. Samat” and con- sisting of ““ossements d’oiseaux sub-fossiles provenant d’Ampasambazimba, Antsirabe, et Morarano prés de Betafo (W. d’Antsir- abe).”’ The locality listed in the general cat- alog for the holotype is Ampasambazimba. Before 1925 several different excavations were conducted at this site and it is impos- sible to determine during which season the holotype was collected. Chronology. —No radiometric date is available for the Stephanoaetus mahery re- mains. They are presumed to be Quater- nary, probably Holocene. This is supported by material of Megaladapis grandidieri, an extinct lemur excavated from Ampasam- bazimba, that has yielded a radiocarbon date of 1035 + SO years B.P. (Tattersall 1973). However, since no stratigraphic informa- tion is available for the Ampasambazimba material reported on herein, it is unknown if the Megaladapis and Stephanoaetus re- mains were associated. Measurements of holotype. —Greatest length 108.0 mm; proximal breadth 26.1 mm; and distal breadth 27.9 mm. (See von den Driesch 1976, fig. 62a— oye IA] EL6I ‘AOZTAIG snjpsu191J40f ~D (4: P-€ € quasqe quosqe € o1eIA] ‘ds‘u sisuaopipsad -D +801 vI-ZI € juesqe juesqe 9-1 O}BIA] (PI61 ‘WoyInos) Dud] “D (p61 9€ I € juasqe quosqe 7 rejnoe ‘IopoIyoS-uUeUILIeH) sypuadanl “Dp LE P-E € juosqe juosqe Z Iepno1oe ‘ds-u isu1ysad snsoydossy (‘ou) (sired) (ou) 9ejasomou vuuojue 1981198 LIBS adh sa1seds S198N39¢ ovrourig siodnas remMony oreyds SS SS Terqouriqoig 9819S0j]0U poys07 (1661) 9PpeRW Pure “(¢g6T) MleIseD “(pL61) Jeporyog-uueUILIeHY “(€261) AOZTENS “(9961) S21eWIETDH Aq poystqnd eyep sopnjouy “oZIs WINUITXeUI JO SWIIOM JOJ SONSLIO}ORIeYO [eOTsO;oYdIow sAT}eIedUIOD SUIOS YIM SNAOYdOM1D snuds dy} Ut soIdads Jo 1SIT—"| 9IGeL VOLUME 107, NUMBER 3 (USNM 168092, USNM 168093, USNM 168094, USNM 168095, USNM 168096, USNM 168097, USNM 168098, USNM 168099, USNM 168100, USNM 168101, USNM 168102, GCRL 1304, GCRL 1305, GCRL 1306, GCRL 1307, GCRL 1308, FSBC I 54472, FSBC I 54473, FSBC I 54474, FSBC I 54475). Florida Keys: Pi- geon Key (24°50’N, 80°45’W), 1 specimen in seagrass kicknet sweeps, | m, coarse cor- alline sand, collected 17 Oct 1991 by J. McLelland. Description. —Holotype 2.9 mm total length, 39 setigers, maximum body width 0.12 mm. Body circular in cross section. Prostomium approximately 1.5 times lon- ger than wide, rounded anteriorly, lacking antenna (Fig. 1B). Pair of small, subdermal, reddish-orange eye spots, distinct but faint in preserved specimens. Two ciliary bands on head, anteriormost limited to dorsal sur- face anterior to eye spots, posterior band emerging laterally from nuchal slits and cir- cumscribing ventral surface. Ciliary bands present on dorsum of prebranchial and branchial segments. First segment setiger- ous. Branchiae three (rarely four) pairs, be- ginning on setiger 4, each as long as segment width, lateral margins ciliated. Dorsal po- dial lobes (postsetal) short, tuberculate on prebranchial setigers, prominent and uni- form in length on setigers 4 through 9; be- coming tuberculate and reduced in size on setigers 10-31, then progressively longer to- ward end of body beginning 3-5 setigers from end. Pygidial region (Fig. 1C, D) with nine cirri consisting of three pairs arranged dorsolaterally to ventrolaterally, apparently representing dorsal podial lobes of three re- duced segments, and three anal cirri, a single ventromedial one and a lateral pair; pos- terior cirri of nearly equal length, longer than pygidium, approximately as long as dorsal podial lobes of last setiger. Forked setae rel- atively slender, asymmetrically lyrate (Fig. 1E), one tine up to twice length of other, tines of equal thickness; occurring singly in inferior notopodial position from setiger 3 527 to at least 4th setiger from end. Remaining notosetae and all neurosetae simple capil- laries. Remarks. — Among the additional mate- rial examined, the number of setigers varied between 30 and 42; the largest specimen observed was 3.5 mm long. A few speci- mens were observed with four pairs of bran- chiae. In the posterior regions of several specimens, large eggs, nearly equal to the body width in diameter, occurred and dark granular material (sperm masses?) was ob- served in some presumed males. Similar go- nadal conditions were also observed for C. perkinsi except that the eggs appeared pro- portionally smaller. Cirrophorus perdidoensis is unique among members of the genus for its combination of small size, lyrate forked setae, and four or fewer pairs of branchiae. Among Cir- rophorus species having lyrate forked setae, C. perdidoensis is similar to C. furcatus and C. lyra by having such setae beginning on the 3rd setiger, and by having podial lobes in the branchial region of uniform length and substantially longer than those of the prebranchial and most of the postbranchial lobes (Strelzov 1973, Mackie 1991). Cir- rophorus perdidoensis differs from the for- mer species, however, by lacking a cephalic antenna, and from both species by having substantially fewer setigers and pairs of branchiae (Table 1). Etymology. —The specific name refers to the type locality, Perdido Key, Florida. Cirrophorus perkinsi, new species Fig. 2 Type material. —Northwest Florida, Per- dido Key (30°17'31”N, 87°25'12’”W). Ho- lotype (USNM_ 168103), two paratypes (USNM 168104), station A9, 12 Dec 1990, 6.1 meters depth, sediment type: 95.4% sand, 4.6% silt-clay. Additional material examined. — Perdido Key: 58 specimens from 11 collections made between 6 Oct 1989 and 3 Dec 1991, 1.6- 528 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Cirrophorus perdidoensis, new species. A. Entire animal, dorsal view. B. Anterior end, dorsal view. C. Posterior end, dorsal view. D. Posterior end, lateral view, neurosetae and cirri from right side omitted (a, podial lobes of reduced segments; b, anal cirri). E. Lyrate forked notosetae. Scales: A = 0.5 mm, B, C, and D = 0.1 mm, E = 0.01 mm. VOLUME 107, NUMBER 3 529 Fig. 2. Cirrophorus perkinsi, new species. A. Anterior end, dorsal view. B. Posterior end, dorsal view. C. Acicular forked notosetae. Scales: A, B = 0.1 mm, C = 0.01 mm. 7.0 m depth, sediment type and hydro- graphic conditions similar to type material (USNM 168105, USNM 168106, USNM 168107, USNM 168108, GCRL 1309, GCRL 1310, GCRL 1311, GCRL 1312, GCRL 1313, GCRL 1314, FSBC I 54476, FSBC I 54477). Florida east coast: six spec- imens from three collections off Hutchinson Island, St. Lucie County (27°21'N, 80°13'W), 10 May 1972 (USNM 54571), 2 Nov 1972 (FSBC I 39860) and 14 Mar 1977 (FSBC I 39861), all 10.9 m depth. Description. —Holotype 2.5 mm total length, 37 setigers, maximum body width 0.18 mm. Body circular in cross section. Prostomium approximately 1.7 times lon- ger than wide, rounded anteriorly, lacking antenna; eye spots absent (Fig. 2A). Two ciliary bands on head, anteriormost limited to dorsal surface, posterior band emerging laterally from nuchal slits and circumscrib- ing ventral surface. Ciliary bands present on dorsum of prebranchial and branchial seg- ments. First segment setigerous. Branchiae simple, three to four pairs, beginning on se- tiger 4, each longer than width of segments, lateral margins ciliated. Dorsal podial lobes absent or reduced to small, indistinct tu- bercles on all but posterior two setigers where they are well-developed. Pygidial region (Fig. 2B) with nine cirri consisting of three pairs arranged dorsolaterally to ventrolaterally, apparently representing dorsal podial lobes of three reduced segments, and three anal 530 cirri, a single ventromedial one and a lateral pair; posterior cirri of nearly equal length, longer than pygidium, approximately as long as dorsal podial lobes of last setiger. Forked setae stout, acicular, bayonet-shaped (Fig. 2C), slender tine about twice as long as stout tine; occurring singly in inferior notopodial position from setiger 2 posteriorly to last fully developed setiger. Remaining noto- setae and all neurosetae simple capillaries. Remarks. —The holotype was the largest specimen observed. Other Perdido Key specimens ranged from 1.5 to 2.5 mm long, had 23-37 setigers, and had three or four pairs of branchiae. Of the Hutchinson Is- land specimens, one (USNM 54571), al- though in poor condition, was observed with about 45 setigers and five pairs of branchiae; the others had 30-35 setigers and four pairs of branchiae. Among the species having acicular bay- onet type modified setae, Cirrophorus per- kinsi is distinguished from C. branchiatus and C. armatus by being much smaller, hav- ing fewer pairs of branchiae, by lacking ly- rate notosetae in the branchial region, and by having tuberculate or reduced dorsal po- dial lobes throughout the body except for the pre-anal region. Cirrophorus perkinsi further differs from C. branchiatus by hav- ing three prebranchial segments instead of 4 or 5. Cirrophorus juvenalis, a small species from southern Africa, is similar to C. per- kinsi in having only acicular type special- ized notosetae and undeveloped notopodial postsetal lobes on all but the last two seti- gers, but C. juvenalis differs from C. perkinsi in having the specialized notosetae begin- ning on the 4th setiger rather than the 2nd setiger and having only one pair of bran- chiae rather than three or more (Table 1). A reduction of three pre-anal segments, resulting in two remnant podial lobes per segment and a disappearance of setae, ap- parently occurs in C. perkinsi and in C. per- didoensis, giving the appearance of a pygid- ial region with nine anal cirri rather than the characteristic three. A similar reduction PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON of segments also apparently occurs in C. juvenalis, which Hartmann-Schroder (1974) described and illustrated as having seven filamentous cirri surrounding the anus. This profusion of cirri at the posterior ends of these three small species, along with the lengthened dorsal podial lobes of the last two or three setigers, could represent an ad- aptation for an interstitial coarse sand hab- itat (T. H. Perkins, in litt.). Etymology.—The species is named for Thomas H. Perkins honoring his many sig- nificant contributions to the study of poly- chaetes and for his helpful suggestions in producing this manuscript. Acknowledgments This research was supported by funds provided by the U.S. National Park Service, contracts CA-5320-9-8001 and CA-5320- 9-8002. We are grateful to T. H. Perkins, Florida Marine Research Institute, St. Pe- tersburg, for his helpful suggestions, efforts in examining and commenting on the mor- phology and sexual maturity of the speci- mens, and reviewing the manuscript. Mr. Perkins arranged the loan of material from Hutchinson Island, Florida. We thank R. W. Heard, C. R. Rakocinski, and S. E. LeCroy for their helpful critiques of the manuscript and G. H. Meyer for translating German literature. An anonymous reviewer made many contributions to this work in- cluding drawing our attention to additional literature and expanding our understanding of paraonid morphology. Literature Cited Castelli, A. 1985. Paraonidae (Annelida, Polychaeta) des fonds meubles infralittoraux des cdtes tos- canes.— Cahiers de Biologie Marine 26:267-279. . 1988. Censimento dei Policheti dei mare It- aliani: Paraonidae Cerruti, 1909.—Atti della Societa Toscana di Scienze Naturali, Memorie, Serie B 94:319-340. Cerruti, A. 1909. Contributo all’anatomia, biologia e sistematica delle Paraonidae (Levinsenidae) con particolare riguardo alle specie del golfo di VOLUME 107, NUMBER 3 Napoli.— Mitteilungen aus der Zoologischen Station zu Neapel 19:459-512. Fauvel, P. 1927. Polychétes sedéntaires. Addenda aux Errantes, Archiannélides, Myzostomaires. — Faune de France, Paris 16:1—-494. Gaston, G. R. 1984. Chapter 2, Family Paraonidae. Pp. 2-1—2-53 in J. M. Uebelacker & P. G. John- son, eds., Taxonomic guide to the polychaetes of the northern Gulf of Mexico. Barry Vittor and Associates, Inc., Mobile, Alabama, 7 vols. Glémarec, M. 1966. Paraonidae de Bretange. De- scription de Paradoneis armata nov. sp.— Vie et Milieu, Série A: Biologie Marine, 17, 2A:1045— 1052. Hartley, J. P. 1981. The family Paraonidae (Poly- chaeta) in British waters: a new species and new records with a key to species.—Journal of the Marine Biological Association of the United Kingdom 61:133-149. Hartmann-Schroder, G. 1974. Zur Polychaetenfauna von Natal (Siidafrika).— Mitteiliingen aus dem Hamburgischen Zoologischen Museum und In- stitut 71:35-73. Katzmann, W., & L. Laubier. 1975. Paraonidae (Po- lychétes sédentaires) de l’Adriatique.— Annalen des Naturhistorischen Museums, Wien 79:567— 588. Laubier, L., & J. Paris. 1962. Faune marine des Pyr- énées-Orientales 4. Annélides polychétes.— Vie et Milieu (Supplement) 13:1-79. ——, & J. Ramos. 1973. Paraonidae (Polychétes sédentaires) de Méditerranée. — Bulletin du Mu- séum National d’Histoire Naturelle, Paris, Série 3, 168 (Zoologie 113):1097-1148. Lopez-Jamar, E., B. O'Connor, & G. Gonzalez. 1987. Demography and gametogenic cycle of Para- doneis armata Glémarec (Polychaeta, Paraoni- 531 dae) in Ria de La Coruna, northwest Spain.— Ophelia 27:127-136. Mackie, A. S. Y. 1991. Paradoneis eliasoni sp. nov. (Polychaeta: Paraonidae) from northern waters, with a redescription of Paradoneis lyra (Southern, 1914).—Ophelia Supplement 5:147-155. Rakocinski, C. F., R. W. Heard, S. E. LeCroy, J. A. McLelland, & T. Simons. 1993. Seaward change and zonation of the sandy-shore mac- rofauna at Perdido Key, Florida, U.S.A.—Es- tuarine, Coastal and Shelf Science 36:81-104. Strelzov, V. E. 1968. Polychaetous annelids of the family Paraonidae (Polychaeta, Sedentaria) of the Barents Sea.—Academy of Sciences of the USSR, Kirov Kola affiliate, Murmansk Marine Biology Institute 17(21):74-95. . 1973. Polychaete worms of the family Parao- nidae Cerruti, 1909—Polychaeta Sedentaria. Academy of Sciences of the USSR, Order of Lenin, S. M. Kirov Kola affiliate, Murmansk Marine Biology Institute (English translation, Amerind Publishing Company Private, Limit- ed, New Delhi, 1979). 212 pp. Taylor, J.L. 1971. Polychaetous annelids and benthic environments in Tampa Bay, Florida. Unpub- lished Ph.D. Dissertation, University of Flori- da, Gainesville. 1332 pp. (JAM) Invertebrate Zoology Section, Gulf Coast Research Laboratory, P.O. Box 7000, Ocean Springs, Mississippi 39566, U.S.A.; (GRG) Biology Department, University of Mississippi, University, Mississippi 38677, U.S.A. PROC. BIOL. SOC. WASH. 107(3), 1994, pp. 532-543 TWO NEW SCALE-WORMS (POLYNOIDAE: POLYCHAETA) FROM THE LAU BACK-ARC AND NORTH FIJI BASINS, SOUTH PACIFIC OCEAN Tomoyuki Miura Abstract. — Thermopolynoe branchiata, a new genus and species of branchiate polynoid polychaete from the Lau Back-Arc and the North Fiji Basins, South Pacific Ocean, is described. The new species belongs to the subfamily Lepi- donotopodinae in having well-developed notopodial bracts. It, however, differs from the other species of the subfamily in having arborescent branchiae. Ther- miphione fijiensis, a new species of the subfamily Iphioninae from the North Fiji Basin, differs from 7. tufari by the first occurrence of hooked neurosetae: segment 3 instead of segment 4 of the latter. The hydrothermal community of the cen- tral part of the North Fiji Basin was found by bottom observations using a deep towed camera system during the R/V Kaiyo cruise conducted under the French-Japanese proj- ect in 1988. A series of cruises named STARMER were realized in the same proj- ect in succeeding years by the French sub- mersible Nautile and its mother ship R/V Nadir (Azuende et al. 1989). The objective of the second cruise, STARMER II, was the study of biology and ecology of organisms associated with the ac- tive vents of the North Fiji Basin. Beds of deep-sea mussels, numerous galatheid crabs, colonies of tiny anemones, and hundreds of gastropods were observed during these sur- veys (Jollivet et al. 1989). Close to the North- Fiji Basin, other hydrothermal fields occur at the spreading center of the Lau Back-Arc Basin west of the Tonga Islands (Fouquet et al. 1991). A scientific team from France, Germany, and Tonga explored the Lau Ba- sin in 1989 and numerous organisms were collected using Nautile and Nadir (the NAUTILAU group 1991). Among the an- imals collected from the above two basins, were numerous specimens of polynoid poly- chaetes. These collections were provided to me for study. Nine species were identified and some were found in both basins. In the present study two new species are described. One is thought to belong to the subfamily Lepidonotopodinae and occurs in both ba- sins. Another species from the North-Fiji Basin is attributed to the subfamily Iphioni- nae. The types are deposited in the Museum National d’Histoire Naturelle a Paris (MNHN), the National Museum of Natural History, Smithsonian Institution (USNM), and Japan Marine Science and Technology Center (JAMSTEC). Some specimens were used for SEM or histological observation at Kagoshima University (KU). Family Polynoidae Subfamily Lepidonotopodinae Pettibone, 1983, emended The subfamily is emended to include Thermopolynoe branchiata, new species. There may be parapodial arborescent bran- chiae instead of lacking branchiae. Thermopolynoe, new genus Type species. — Thermopolynoe branchia- ta, new species. Gender. — Feminine. VOLUME 107, NUMBER 3 Diagnosis. — Body flattened; 27 segments (first achaetous). Elytra 11 pairs, on large elytrophores on segments 2, 4, 5, 7, 9, 11, 13, 15, 17, 19, and 21. Dorsal cirri on non- elytrigerous segments; dorsal tubercles large. Branchiae well developed, arborescent. Prostomium bilobed; anterior lobes cylin- drical, with frontal filaments; median an- tenna with short ceratophore and subulate style, inserted in anterior notch; palps with slender tips. First or tentacular segment not distinct dorsally; tentaculophores achaetous lateral to prostomium with two pairs of ten- tacular cirri. Segment 2 with first pair of elytrophores, biramous parapodia, and buc- cal cirri. Parapodia biramous, with noto- podia shorter than neuropodia. Notopodia subconical, with large bracts. Neuropodia truncate, with fimbriated margins. Noto- setae and neurosetae numerous, spinous. With or without ventral segmental papillae on middle segments. Pygidium bulbous, wedged between posterior parapodia, with pair of anal cirri. Pharynx with seven pairs of papillae and two pairs of jaws. Etymology.—The generic name is from Greek, Thermos, hot and Polynoe, genus name of a polynoid worm. Thermopolynoe branchiata, new species Figs. 1-4 Material examined. —White Lady, North Fiji Basin, DSRV Nautile Dive 16, 11 Jul 1989, STARMER II Station 4, 16°59.5’S, 173°55.5'E, 2000 m, Holotype (MNHN UD 264), 14 paratypes (MNHN UD 262, JAM- STEC, KU). Same site, Dive 13, 8 Jul 1989, paratype (MNHN UD 265). Same site, Dive 20, 15 Jul 1989, 4 paratypes (USNM 168325). Vailili, Lau Basin, DSRV Nautile Dive 10, 22 May 1989, BIOLAU Station 2, 23°13’S, 176°38’E, 1750 m, 2 paratypes (MNHN UD 261). Description. — Holotype 36 mm long, 12 mm wide including parapodia, with 27 seg- ments including first achaetous tentacular segment. Largest paratype 53 mm long, 20 533 mm wide, with 27 segments. Body sturdy, slightly tapered anteriorly and posteriorly, flattened ventrally, slightly arched dorsally (Fig. la, b). Integument smooth. Preserved specimens brownish to tan. Elytra 11 pairs, on segments 2, 4, 5, 7, 9, 11, 13, 15, 17, 19, and 21, large, covering dorsum, oval, stiff, rough with numerous brownish papillae on white bases (Figs. la, 2d-g). Elytra on segments 2-19 oval, wider than long (Fig. 2d, f); last pair on segment 21 subreniform, longer than wide (Fig. 2e). Dorsal cirri on non-elytrigerous segments with short cylindrial cirrophores, and short styles with slender tips, extending to tips of neurosetae; dorsal tubercles nodular (Figs. 3c, f, 4a—c). Branchiae arborescent, present on segments 3-26, around bases of dorsal tubercles and elytrophores; separated into two groups on anterior and posterior sides of notopodia on segments 3 and on some less-developed posterior segments; forming single large branchial areas encircling cen- tral parts of notopodia on other fully de- veloped parapodia with about 20 tufts of branchiae; each tuft with 1-15 filaments; each parapodium maximally with about 80 branchial filaments (Figs. 3f-g, 4a, b). Prostomium bilobed. Anterior lobes prominent, cylindrical, extending anterior- ly, with small frontal filaments; median an- tenna, inserted in anterior notch, with short cylindrical ceratophore and subulate style extending to about tip of palp; palps thick, slightly wrinkled, extending beyond prosto- mium, with slender tips. Tentacular seg- ment not distinct dorsally; tentaculophores lateral to prostomium achaetous, with two pairs of tentacular cirri slightly shorter than palps (Fig. 3a, b). Segment 2 with first pair of elytrophores and biramous parapodia; ventral or buccal cirri attached basally on prominent cirro- phores lateral to mouth, with styles similar to tentacular cirri, longer than following ventral cirri (Fig. 3a, b, e). Mouth opening situated between segments 1 and 2. Mus- cular pharnyx encircled by seven pairs of 534 Fig. 1. bulbous papillae, subequal in size; dorsal and ventral pairs of jaws fused medially, each with 5-7 teeth on basal sides (obser- vation on dissected paratypes). Segment 3 with first pair of arborescent branchiae, dorsal cirri, short ventral cirri, and setal lobes similar to segment 2 (Fig. 3a, b, f). Following parapodia biramous, with short notopodia on anterodorsal sides of large neuropodia (Fig. 3g). Notopodia sub- conical, with projecting acicular lobes hid- 10 mm PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Thermopolynoe branchiata, new species. Paratype (MNHN UD 265): a, Dorsal view; b, Ventral view. den by numerous notosetae, and enclosed antero-dorsally by well-developed large flaring bracts (Figs. 3e-g, 4a—c). Neuropodia diagonally truncate, deeply notched on up- per part; distal margins fimbriated with slender filaments covered with numerous filamentous bacteria (Fig. 2a—c). Notosetae numerous, forming radiating bundles, much stouter than neurosetae, serrated on distal margins; tips bare, blunt, tapered (Fig. 4d). Neurosetae numerous, VOLUME 107, NUMBER 3 Fig. 2. 535 aimm b,g 0.3 mm c 0.03 mm d,e 10 mm f 3 mm h 2mm Thermopolynoe branchiata, new species. Paratype (MNHN UD 265): a, Left cirrigerous parapodium from segment 10, anterior view; b, Filament of neuropodium from same, with filamentous bacteria; c, Bacterial filament; d, Left (Sth) elytron from segment 9, papillae eliminated; e, Left (11th) elytron from segment 21, papillae eliminated; f, Left (1st) elytron from segment 2, surface micropapillae illustrated; g, Papillae from same; Paratype (MNHN UD 262): h, Left halves of segments 10-13, ventral view. forming fan-shaped bundles. Supraacicular neurosetae with numerous prominent spines in two rows; tips bare, tapered (Fig. 4e). Subacicular neurosetae with numerous prominent spines in single rows; tips bare, slightly hooked (Fig. 4f). Ventral segmental papillae long, attached to bases of neuropodia, and extending to bases of ventral cirri; three pairs present on segments 12, 13, and 14 (Fig. 2h), or lacking in half of large specimens (Fig. 1b). Pygid- ium visible dorsally as bulbous lobe, wedged between parapodia of posterior smaller seg- ments (26-27), with pair of long ventral anal 536 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON g-3 Fig. 3. Thermopolynoe branchiata, new species. Paratype (MNHN UD 265): a, Anterior end, dorsal view; b, Same, ventral view; c, Posterior end, dorsal view; d, Same, ventral view; e-1, Left elytrigerous parapodium from segment 2, elytron removed, anterior view; e-2, Same, dorsal view; e-3, Same, posterior view; f-1, Left cirrigerous parapodium from segment 3, anterior view; f-2, Same, dorsal view; f-3, Same, posterior view; g-1, Left elytrigerous parapodium from segment 9, elytron removed, anterior view; g-2, Same, dorsal view; g-3, Same, posterior view. VOLUME 107, NUMBER 3 537 a-c 4mm \\ ISS d-f 0.1mm GAZE | pop aes Begewon c-3 Fig. 4. Thermopolynoe branchiata, new species. Paratype (MNHN UD 265): a-1, Left cirrigerous parapodium from segment 10, anterior view; a-2, Same, dorsal view; a-3, Same, posterior view; b-1, Left cirrigerous para- podium from segment 26, anterior view; b-2, Same, dorsal view; b-3, Same, posterior view; c-1, Left cirrigerous parapodium from segment 27, anterior view; c-2, Same, dorsal view; c-3, Same, posterior view; d, Notoseta from segment 9; e, Supraacicular neuroseta from same; f, Subacicular neuroseta from same. 538 cirri, similar to posterior dorsal cirri (Fig. 3c, d). Variation in paratypes. —Among the paratypes, two small (young) specimens had less than 27 segments. They were 8.1 and 13 mm long by 4.4 and 6.1 mm wide with 24 and 25 segments, respectively. The spec- imens with 27 segments varied from 12 mm to 53 mm in length and from 6.5 to 20 mm in width. Of the 22 specimens examined, 12 had three pairs of elongate ventral pa- pillae on segments 12-14, a single specimen had two pairs on segments 12-13, and nine lacked ventral papillae. Etymology. —The name was derived from the arborescent branchiae of the species. Remarks.—The structure of prostomi- um, tentacular segment, and pharynx of the new species recalls the subfamilies Macel- licephalinae and Lepidonotopodinae (Pet- tibone 1976, 1983, 1984b, 1985b, 1988, 1989, 1990). Of the genera of these two sub- families, Thermopolynoe is the closest to Lepidonotopodium in having well-devel- oped notopodial bracts, 11 pairs of elytra, long segmental papillae, seven pairs of pha- ryngeal papillae, and fimbriated neuropo- dia. However, Thermopolynoe branchiata clearly differs from the species of Lepidono- topodium in having well-developed arbo- rescent branchiae instead of lacking them. In the family Polynoidae, the species of three subfamilies, Branchipolynoinae, Branchinotogluminae and Branchiplicati- nae, have well-developed branchiae. The first two families have arborescent type of branchiae and the Branchiplicatinae, folded type (Pettibone 1985a). Peinaleopolynoe sil- lardi Desbruyéres & Laubier, whose affili- ation on the subfamily was not assigned by the authors, also has arborescent branchiae (Desbruyéres & Laubier 1988). The species was recently examined and referred to the Branchinotogluminae by Pettibone (1993), along with a new species of Peinaleopoly- noe: P. santacatalina. The presence of these well-developed arborescent branchiae is an unusual feature in the family Polynoidae as mentioned by Pettibone (1984a). Thermo- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON polynoe branchiata is, however, unique among these branchiate polynoids in having well-developed bracts encircling the noto- podia. The new species differs from them also in the position of branchiae: branchiae are separated in anterior and posterior groups or forming single continued bran- chial areas in 7. branchiata, but divided in upper and lower groups in others. Subfamily Iphioninae Baird, 1865 Thermiphione Hartmann-Schroder, 1992 Thermiphione fijiensis, new species Figs. 5-7 Material. —White Lady, North Fiji Basin, DSRV Nautile Dive 16, 11 Jul 1989, STARMER II Station 4, 16°59.5'S, 173°55.5'W, 2000 m, Holotype (MNHN UD 266) & 8 paratypes (USNM 168326, MNHN UD 263, KU); Dive 20, 15 Jul 1989, 1 paratype (JAMSTEC); Dive 21, 16 Jul 1989, 1 paratype (JAMSTEC). Description. —Holotype 14 mm long, 7 mm wide including setae, with 31 segments including first setigerous tentacular seg- ment. Largest paratype 20 mm long, 10 mm wide, with 30 segments. Body short, ovate, flattened ventrally and slightly arched dor- sally (Fig. 5a, b). Pygidium without ap- pendages (Fig. Se). Elytra 14 pairs, large, strongly imbricat- ed, light yellow, stiff, rough with numerous papillae (Figs. 5a, 6a—e), present on seg- ments, 22745555 7. 9ealel. Sets. lee eee 23, 26, and 27. Eltyra on segment 2 oval, with fringes of short papillae, covered with filamentous bacteria (Fig. 6a, g, h). Other elytra elongate subreniform, narrower me- dially, wider laterally; lateral borders with fringes of short papillae (Fig. 6b—e). Elytral surfaces covered with hexagonal or polyg- onal areas with secondary areolae (Fig. 6f). Elytrophores bulbous, transversely elongat- ed; places of attachment with latero-poste- rior extensions (Fig. 5c, e). Dorsal tubercles on cirrigerous segments bulbous, promi- nent, transversely elongated, continuous with enlarged cirrophores of dorsal cirri VOLUME 107, NUMBER 3 539 Fig. 5. Anterior end, dorsal view; d, Same, ventral view; d, Posterior end, dorsal view; Paratype (JAMSTEC): f, Distal end of pharynx showing jaws and border papillae. (Figs. 5c, 7e). Dorsal cirri on non-elytriger- ous segments with long cylindrical cirro- phores and short styles, extending to tips of neurosetae (Figs. 5c, 7c, e). Dorsal and ven- tral cirri with short clavate papillae (Fig. 7f). Prostomium partially fused to tentacular segment, and withdrawn in anterior seg- Thermiphione fijiensis, new species. Holotype (MNHN UD 266): a, Dorsal view; b, Ventral view; c, ments (Fig. Sc, d). Prostomium bilobed, forming separate rounded lobes, with an- terolateral extensions; without antennae; palps thick, smooth, ventral to lateral pro- stomial extensions (Fig. 5c). Tentacular seg- ment with long cylindrical tentaculophores lateral to prostomium; each with single aci- 540 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 6. Thermiphione fijiensis, new species. Holotype (MNHN UD 266): a, Right first elytron from segment 2; b, Right elytron from segment 5; c, Right elytron from segment 17; d, Right elytron from segment 26; e, Right elytron from segment 27; f, Enlarged hexagonal area enclosing small areolae of right elytron from segment 2; g, Enlarged margin of right elytron from same; h, Enlarged marginal papilla of same, with filamentous bacteria. culum, few capillary setae, and dorsal and __ than 20 papillae on anterior 7 segment; ar- ventral tentacular cirri, shorter than palps ranged in two or more rows before segment (Figs. Sc, d, 7a, g). Nodular papillae arising 14, thereafter in single row, absent posterior from dorsum of anterior segments; more to segment 20 (Fig. Sc). VOLUME 107, NUMBER 3 541 i a-e 1 mm f-k 0.1 mm Fig. 7. Thermiphione fijiensis, new species. Holotype (MNHN UD 266): a, Left tentaculophore, inner ventral view, single aciculum dotted; b, Left elytrigerous parapodium from segment 2, anterior view, noto- and neu- roaciculum dotted; c, Left cirrigerous parapodium from segment 3, posterior view; d, Left elytrigerous para- podium from segment 13, anterior view; e, Left cirrigerous parapodium from segment 14, posterior view; f, Enlarged papillae on ventral buccal cirrus of segment 2; g, Capillary seta from tentaculophore; h, Notoseta; 1, Upper feathered neuroseta; j, Middle slightly hooked neuroseta; k, Lower hooked neuroseta. 542 Segment 2 with single rounded nuchal lobe, elongated elytrophores, first pair of el- ytra, biramous parapodia, and ventral buc- cal cirri attached basally on prominent cir- rophores lateral to mouth, with styles similar to tentacular cirri, longer than following ventral cirri (Figs. 5c, d, 7b). Muscular phar- ynx encircled by nine pairs of bulbous pa- pillae subequal in size; dorsal and ventral pairs of jaws fused medially, without teeth (Fig. 5f). Segment 3 not visible dorsally, with dor- sal cirri, and parapodia wedged between elytrophores of segments 2 and 4 (Figs. 5c, 7c). Biramous parapodia with short noto- podia on anterodorsal sides of large truncate neuropodia (Fig. 7d, e). Notopodia subcon- ical, with projecting acicular lobes hidden by numerous notosetae (Fig. 7d). Notosetae forming radiating bundles of dense tufts, shorter than neurosetae, feathered, with slender axes and long capillary tips (Fig. 7h). Upper neurosetae feathered, stouter than notoseta, with short capillary tips (Fig. 71); middle neurosetae stout, hooked, with long, faint spinous areas (Fig. 7j); lower neuro- setae similar but shorter, hooked, with short, faint spinous areas (Fig. 7k). Hooked neu- rosetae first present on segment 3. Variation in paratypes. — Among the type specimens, three small (young) specimens had less than 30 segments, with 10-12 pairs of elytra. The other specimens had 30 seg- ments and 14 pairs of elytra, except for the holotype with 31 segments. These larger specimens varied from 10.0 to 19.6 mm in length and from 5.5 to 10.0 mm in width. In all specimens, hooked neurosetae started from segment 3. Etymology.—The species name is de- rived from the type locality, the North Fiji Basin. Remarks. —The presence of only a rem- nant or the complete absence of a median prostomial antenna is characteristic for the subfamily Iphioninae. The Iphioninae differ from other polynoids also by the unique reticulated elytra with hexagonal or polyg- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON onal honey-comb-like areas, the tentacu- lophore of the first segment with a few cap- illary setae, and styles of the dorsal and ventral parapodial cirri. In four previously known genera of the subfamily, [phionides differs from others in having secondary lat- tice structure in each polygonal area of el- ytron, small dorsal tubercles, up to 39 seg- ments, and up to 20 pairs of elytra (Pettibone 1986). Both Iphione and Iphionella have 29 segments and 13 pairs of elytra. Thermiphi- one differs from the above three genera in having 30-31 segments and 14 pairs of el- ytra. The last two elytra occur on segments 23 and 27 in Iphione, on 23 and 26 in Iphi- onella, and on segments 26 and 27 in Ther- miphione. Thermiphione fijiensis differs from the other congeneric species T. tungari by the first occurrence of hooked neurose- tae: segment 3 instead of segment 4 of the latter (Hartmann-Schroder, 1992). Acknowledgments I wish to thank Daniel Desbruyéres of the Institut Francais de Recherche pour l’Ex- ploitation de la Mer (IFREMER) and Mich- el Segonzac and Patrick Briand of the Centre National de Tri d’Océanographie Biolo- gique for their kind communication of the materials examined in this study. The manuscript benefited from the reviews of Marian H. Pettibone and James A. Blake. This work was supported in part by grants- in-aid from the Ministry of Education, Cul- ture and Science, Japan (No. 03640630) and Kato Foundation for the Promotion of the Bioscience Researches. Literature Cited Auzende, J.-M. et al. 1989. Le cadre géologique d’un site hydrothermal actif: la campagne STAR- MER 1 du submersible Nautile dans le bassin Nord-Fijien.—Comptes Rendus Hébdoma- daires des Séances de |’Academie des Sciences, Série II 309:1787-1795. Desbruyéres, D., & L. Laubier. 1988. Exploitation d’une source de matiére organique concentrée dans l’océan profond: intervention d’une an- VOLUME 107, NUMBER 3 nélide polychéte nouvelle.—Comptes Rendus Hébdomadaires des Séances de l’Academie des Sciences, Série III 307:329-335. Fouquet, Y. et al. 1991. Hydrothermal activity and metallogenesis in the Lau back-arc basin. — Na- ture, London 349:778-781. Hartmann-Schréder, G. 1992. Zur Polychaetenfauna in rezenten hydrothermalen Komplexmassiv- sulfiderzen (“Schwarze Raucher’’) am Ostpazi- fischen Riicken bei 21°30’S.—Helgolander Meeresuntersuchungen 46:389—-403. Jollivet, D. et al. 1989. Premiéres observations de communautés animales associées a l’hydrother- malism arriérre-arc du bassin Nord-Fijien.— Comptes Rendus Hébdomadaires des Séances de l’Academie des Sciences, Série III 309:301- 308. The NAUTILAU Group. 1991. Hydrothermal ac- tivity in the Lau basin.—Eos (Transactions, American Geophysical Union) 71:678-679. Pettibone, M. H. 1976. Revision of the genus Ma- cellicephala McIntosh and the subfamily Ma- cellicephalinae Hartmann-Schréder (Polychae- ta: Polynoidae). —Smithsonian Contributions to Zoology 229:1-71. 1983. A new scale worm (Polychaeta: Poly- noidae) from the hydrothermal rift-area of west- em Mexico at 21°N.—Proceedings of the Bio- logical Society of Washington 96:392-399. 1984a. A new scale-worm commensal with deep-sea mussels on the Galapagos hydrother- mal vent (Polychaeta: Polynoidae).—Proceed- ings of the Biological Society of Washington 97: 226-239. 1984b. Two new species of Lepidonotopo- dium (Polychaeta: Polynoidae: Lepidonotopo- dinae) from hydrothermal vents off the Gala- pagos and East Pacific Rise at 21°N.— Proceedings of the Biological Society of Wash- ington 97:849-863. . 1985a. An additional new scale worm (Poly- chaeta: Polynoidae) from the hydrothermal rift area off western Mexico at 21°N.— Proceedings 543 of the Biological Society of Washington 98:150- 157. 1985b. New genera and species of deep-sea Macellicephalinae and Harmothoinae (Poly- chaeta: Polynoidae) from the hydrothermal rift areas off the Galapagos and western Mexico at 21°N and from the Santa Catalina Channel.— Proceedings of the Biological Society of Wash- ington 98:740-757. . 1986. Review of the Iphioninae (Polychaeta: Polynoidae) and revision of [phione cimex Qua- trefages, Gattyana deludens Fauvel, and Har- mothoe iphionelloides Johnson (Harmothoi- nae).—Smithsonian Contributions to Zoology 428:1-43. . 1988. New species and new records of scaled polychaetes (Polychaeta: Polynoidae) from hy- drothermal vents of the Northeast Pacific Ex- plorer and Juan de Fuca Ridges. — Proceedings of the Biological Society of Washington 101: 192-208. . 1989. New species of scale-worms (Polychae- ta: Polynoidae) from the hydrothermal rift-area of the Mariana back-arc basin in the western central Pacific.—Proceedings of the Biological Society of Washington 102:137-153. . 1990. New species and new records of scaled polychaetes (Polychaeta: Polynoidae) from the Axial Seamount Caldera of the Juan de Fuca Ridge in the northeast Pacific and the east Pa- cific Ocean off northern California.—Proceed- ings of the Biological Society of Washington 103: 825-838. 1993. Polynoid polychaetes associated with a whale skeleton in the bathyal Santa Catalina basin. — Proceedings of the Biological Society of Washington 106:678-688. Faculty of Fisheries, Kagoshima Univer- sity, 4-50-20, Shimoarata, Kagoshima 890, Japan. PROC. BIOL. SOC. WASH. 107(3), 1994, pp. 544-547 ELECTRA VENTURAENSIS, A NEW SPECIES (BRYOZOA: CHEILOSTOMATA: MEMBRANIPORIDAE) FROM SOUTHERN CALIFORNIA William C. Banta and Mae M. Crosby Abstract. — Electra venturaensis is described from shallow subtidal water near Ventura, California. It is distinguished from E. monostachys (Busk) by lacking typical lateral spines and possessing a calcified operculum, and from E. crus- tulenta (Pallas) by possessing numerous frontal heterozoids and occupying a non-estuarine habitat. Adventitious buds derived from reparative buds or ken- ozoids derived from spines may form pseudoancestrulae and subcolonies in later astogenetic stages. Methods and Materials Pelecypod shells and other substrates were collected by snorkel in shallow subtidal wa- ters off Rincon Beach, about 5 miles north of Ventura, California on 24 March 1968. Fragments of about 15 colonies were pre- served in 70% ethyl alcohol. Some colony fragments were dried and treated varying times in 5% NaOCl or cleaned with dilute detergent in an ultrasonic cleaner. Dry spec- imens were dyed with 1% aqueous congo red and coated with sublimed ammonium chloride for light photomacrography (Kier et al. 1976). Some material was dehydrated in ethanol to propylene glycol and vacuum- imbedded in Epotuf resin for thin-ground sections containing calcareous tissues in place next to soft tissues (Nye et al. 1972). Additional topotype material: material en- crusting sandstone collected 29 Jan 1965, one dry colony and one preserved in 70% alcohol; one dried colony on sandstone col- lected Dec 1963. Family Electrinidae Stach, 1937 Genus Electra Lamouroux, 1816 Electra venturaensis, new species Type material. —Holotype: National Mu- seum of Natural History bryozoan Type USNM 477681, slide of dried material (Figs. 1, 2). Paratype: USNM 15005 and 15006, slides of dried material. Topotype material originally identified and labelled Electra crustulenta arctica (Borg) was collected at the same site in Dec 1963, and 29 Jan 1965 and preserved dry. Diagnosis. —Colony pale tan, encrusting shells, unilaminar to multilaminar (1-3 lay- ers). Autozoids arranged in quincunx, com- pletely covering the substrate, without uni- serial rows. Ten tentacles. Early astogenetic stages: unilaminar, with proximal, smooth, mostly imperforate gymnocyst occupying about one third of the frontal surface; cryp- tocyst a narrow, minutely tuberculated rim completely surrounding the opesium. A me- dian proximal, pointed, cuticular spine, a quarter to a third the length of the opesium, projects obliquely forward over the ope- sium. The base of the spine becomes in- creasingly calcified with age, forming a cen- trally perforate hemispherical boss on the gymnocyst. Minute gymnocystal windows, from which frontal-marginal kenozoids will form, are present at the distolateral zoid cor- ners (Fig. 1, arrows). No ovicells, avicularia or other spines. Operculum shaped like a thumbnail, with a straight transverse prox- imal border; always evenly calcified, white, translucent. Basal wall complete, thinly cal- VOLUME 107, NUMBER 3 Figs. 1-3. heterozoids develop, are indicated by arrows. Scale, 0.50 mm; 2: Another part of the holotype colony, showing heterozoids. Scale, 0.50 mm; 3: Transverse Epotuf thin section of paratype encrusting a pelecypod shell, showing heterozoids (h) and a spine (s). Scale, 0.10 mm. cified. Later astogenetic stages: ectocystal buds, calcified basally and laterally, are pro- duced frontolaterally from windows at the distolateral zoidal corners. These buds fill zoidal margins by a rim of frontal keno- zoids, which may become partly to com- pletely occluded by annularly calcifying gymnocyst and cryptocyst. Parts of the col- ony at this stage resemble species of Cono- 1: Electra venturaensis, new species, holotype. Distolateral gymnocystal windows, from which peum or Antropora. Finally, some keno- zoids may enlarge into pseudoancestrular autozoids, which grow out over older col- ony layers to form subcolonies: superficial layers of autozoids and kenozoids which smother out older layers. Reparative buds, growing from septulae of damaged zoids, may also form new pseudocolonies. Additional description of the holotype. — 546 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 1.—Measurements of the holotype, of Electra venturaensis n = 30, measurements in mm. Autozoid Opesium Operculum Metric Length Width Length Width Length Width Mean 0.457 0.236 0.335 0.157 0.111 0.095 SD 0.034 0.021 0.028 0.017 0.014 0.011 Min 0.380 0.190 0.270 0.120 0.080 0.080 Max 0.500 0.290 0.380 0.190 0.140 0.120 Range 0.120 0.100 0.110 0.070 0.060 0.040 The holotype comprises several NaOCl- treated fragments of a single colony which encrusted a pelecypod shell. Measurements are given in Table 1. All the features de- scribed in the diagnosis are represented ex- cept parts lost in cleaning, notably opercula and distal parts of the spines. Distal and lateral septulae are multiporous pore plates, which become surrounded by thick vertical and oblique calcareous buttresses. The gym- nocyst is usually imperforate, but minute cuticular windows may occur on its distal- frontolateral face. These windows are the source of kenozoid buds, which appear in later astogeny. The asynchronous develop- ment of buds and autozoids is reflected in the structure of the interzoidal walls viewed in section (Fig. 3). Cuticle is present between autozoids and heterozoids in all planes of section except those passing through the rel- atively small pore plates. Kenozoids con- tain cells and sometimes parietal muscles, but no polypide. They communicate basally with the parent zoid; this arrangement, to- gether with their location suggests that they represent highly modified, flattened spines. The first signs of frontal kenozoids occur in distal regions after at least 10 zoidal gen- erations of simple Electra morphology. Frontal budding in this species, therefore, may represent either programmed astoge- netic change or a response to some microen- vironmental cue (Boardman et al. 1969). Generic placement. —The most widely ac- cepted generic concept of Electra is that of Ryland & Hayward (1977:64) who studied European type material. It includes simple malacostegans with relatively large gym- nocysts, small cryptocysts, and a median proximal spine, often with additional spines near the opesium. A consistently calcified operculum is rare among cheilostomes, but found in some species of Electra, notably E. crustulenta Pallas, 1766:39. The generic diagnosis is herein revised to include E. ven- turaensis, which possesses frontal keno- zoids. Etymology.—The species name refers to the city of Ventura, California, derived from archaic Spanish, meaning ‘good luck,’ “‘hap- piness,’ and ‘fair destiny.’ Related species.—Horowitz (1992) lists 38 species and subspecies referred to Elec- tra. Most closely related is E. crustulenta typica (Borg 1931:27), a European steno- haline marine species (Prenant & Bobin 1966:153). A proximal median spine is usu- ally the only spination and there is a cal- cified operculum with a straight or concave proximal border. It differs from FE. ventu- raensis in lacking regular frontal budding, in having more elongated autozoids with a greater average length, 0.72 mm long ac- cording to Borg 1931:30, compared to 0.46 for E. venturaensis). Electra crustulenta bal- tica (Borg 1931:27) is similar to E. crustu- lenta typica, an exclusively estuarine species frequently lacking median proximal spines. Electra monostachys (Busk 1854:61), a Brit- ish estuarine species, forms uniserial to oli- goserial (2-3 rows) stellate or dendritic patches. The median proximal spine may be lacking and numerous spines frequently occur around the opesium; a pair of spines VOLUME 107, NUMBER 3 is usually present lateral to the operculum and the operculum is uncalcified (Ryland & Hayward 1977:70). Electra artica (Borg 1931:27), is circumpolar, stenohaline (Pow- ell 1968:2282), sometimes pluriserial, the operculum is calcified and the gymnocyst is extensive and imperforate; frontal hetero- zoids are absent (see Powell & Crowell 1967: 339). Specimens from Alaska identified by Osburn as Electra crustulenta arctica (Borg) in the reference collections of the National Museum of Natural History are largely oli- goserial and lack frontal heterozoids. Acknowledgments We thank Alan H. Cheetham and Joanne Sanner, Smithsonian Institution, National Museum of Natural History (U.S.A.) for help with specimens and with other aspects of this research. Kelley R. Corneles and Heather Burkholder, AU undergraduates, helped prepare illustrations. Literature Cited Boardman, R.S., A. H. Cheetham, & P. L. Cook. 1969. Intracolony variation and the genus concept in Bryozoa.— Proceedings of the North American Paleontological Convention, Part C: 249-320. Borg, F. 193i. On some new species of Membrani- pora.—Arkiv for Zoologie 22A(4): 1-35. Busk, G. 1854. Catalogue of marine Polyzoa in the collection of the British Museum, Part II, Chei- lostomata (Part). Reprinted 1966 Johnson Re- print Co. New York, pp. i—viii + 55-120. Horowitz, A.S. 1992. List of Recent species of bryo- zoans. Electronic database available from Alan S. Horowitz, Department of Geology, Univer- sity of Indiana, 1005 E. 10th Street, Blooming- ton, Iowa 47461, U.S.A. Email Telenet horo- witz@gizmo.geology.india.edu. 547 Kier, P. M., R. E. Grant, & E. L. Yochelson. 1965. Whitening fossils. Pp. 453-456 in B. Krummel & D. Raup, eds., Handbook of Paleontological Techniques. W. H. Freeman, San Francisco. xiii + 852 pp. Lamouroux, J. V. F. 1816. Historie di polypiers cor- ralligenes flexibles, vulgarairment nommes zoo- phytes. Caen. pp. 1-559. Nye, O. B., Jr., Donald A. Dean, & Robert W. Hinds. 1972. Improved thin section techniques for fos- sil & Recent organisms.— Journal of Paleontol- ogy 46(2):271-275. Osburn, R. C. 1950. Bryozoa of the Pacific coast of North America. Part I, Cheilostomata-Anas- ca.—Allan Hancock Pacific Expeditions 14(1): 1-269. Pallas, P. S. 1766. Elenchus zoophytorum, sistens generum adumbrationes generaliores et specie- rum cognitarum succinctas descriptiones, cum selectis auctorum synonymis.—Hagae-Comi- tum, xxvii + 451 pp. Powell, N. A. 1968. Bryozoa (Polyzoa) of arctic Can- ada.—Journal of the Fisheries Research Board of Canada 25(11):2269—2320. ——_., & G. D. Crowell. 1967. Studies on Bryozoa (Polyzoa) of the Bay of Fundy region. I.— Bryo- zoa from the intertidal zone of Minas Bay and Bay of Fundy.—Cahiers de Biologie Marine 8:331-347. Prenant, M., & G. Bobin. 1966. Bryozoaires. Deux- ieme Partie, Chilostomes Anasca.—Fauna de France 68:1-647. Ryland, J.R., & P. J. Hayward. 1977. British anascan bryozoans. Pp. 1-188 in D. M. Kermack, ed., Synopsis of the British fauna (new series). Lin- nean Society of London. Stach, L.W. 1937. Reports of the McCoy Society for Field Investigation and Research. Lady Julia Percy Island. 13. Bryozoa.— Proceedings of the Royal Society of Victoria, n.s. 49:373—84. Department of Biology, American Uni- versity, Washington D.C. 20016, U.S.A. PROC. BIOL. SOC. WASH. 107(3), 1994, pp. 548-556 THE MORPHOLOGY AND GENERIC RELATIONSHIPS OF SOME FISSIPAROUS HETERONEMERTINES Nathan W. Riser Abstract. —No morphological difference could be found between a fissiparous heteronemertine from New Zealand and other fissiparous species with similar external appearances. All constitute a single species. Significant anatomical characters are shared with Lineus lacteus (Rathke, 1843) and L. pseudolacteus Gontcharoff, 1951. The genus Myoisophagos is erected to contain the three species with M. sanguineus (Rathke, 1799) designated as type species. Signif- icant morphological characters separate the new genus from the genus Lineus. Clumps of slender nemertines were en- countered beneath rocks on the reef just sea- ward of the rubble beach along the outer face of the Kaikoura Peninsula, New Zea- land during the early months of 1983 and 1986. The clusters were reminiscent of Lin- eus socialis (Leidy, 1855) from New England midtidal rocky shores as was the pigmen- tation and rather regularly spaced annuli be- hind the head region of the body to that of the New England species. Five specimens of comparable size were bisected, one at the first annulus and the others successively one annulus further back to determine if the an- imals could regenerate. The heads were re- tained in a single jar, and each of the pos- terior pieces was placed in a separate jar of sea water maintained on a sea water table. Two days later, the posterior pieces had fragmented; with most of the fragments bearing one or two annuli, rarely three. Fourteen days after fission, the fragments had developed white heads with two red eyes. Existing descriptions of fissiparous lin- eid species contain no anatomical data by which species can be identified. Living spec- imens of “species” to which names had been assigned were obtained and a morphologi- cal comparison was undertaken. Materials and Methods In addition to the specimens from Kai- koura, New Zealand, living specimens of Lineus lacteus (Rathke, 1843), L. pseudo- lacteus Gontcharoff, 1951, L. sanguineus (Rathke, 1799), and L. nigricans Birger, 1892 were obtained from the colonies main- tained by Prof. J. Bierne. Living specimens of L. vegetus Coe, 1931 were furnished by Prof. P. Roe from central California rocky beaches, and L. socialis (Leidy, 1855) was collected from a number of rocky shoreline outcroppings between Nahant, Massachu- setts and Georgetown, Maine. Anestheti- zation, fixation, sectioning, and staining were as described in Riser (1988). Systematic Account Lineidae sensu Gibson, 1982 Myoisophagos, new genus Type species. — Myoisophagos sanguineus (Rathke, 1799), new combination. Diagnosis. — With three apical sensory or- gans; frontal glands absent; rhynchodaeum attached ventrally and thus with a single cephalic blood lacuna arched over the rhyn- chodaeum; rhynchocoel muscle layers not interwoven nor interwoven with muscles of body wall; proboscis with two muscle lay- ers, inner longitudinal and outer circular; vascular plexus in foregut region; neuro- chord cells and caudal cirrus absent; sub- epidermal gland cells penetrate OLM, some contact medullary nerve plexus; with sub- epithelial esophageal gland cells and longi- VOLUME 107, NUMBER 3 tudinal muscles; longitudinal muscle plate between rhynchocoel and foregut absent; ra- dial muscles to either side of the rhyncho- coel pass beneath the rhynchocoel crossing to the opposite side and continue around the foregut. Etymology. —myos Gr. muscular, oisoph- agos Gr. esophagus. Radial muscles from the body wall ar- boresce through the subepithelial esopha- geal gland layer (Figs. 8, 11). They, with the subepithelial esophageal longitudinal mus- cles control expansion and contraction of the foregut. The dorsal radial muscles form a sling around the rhynchocoel (Fig. 11) crossing beneath that organ and continuing around the foregut as circular bands. These bands lie amidst the bases of the esophageal glands of the empty foregut (Fig. 8), but are pressed closer to the subepithelial longitu- dinal muscles of the inflated organ. Fried- rich (1935) referred to these radial fibers in the foregut region as dorsoventral muscles. Burger (1897-1907; table 3, fig. 18) illus- trated the penetration of the sub-epidermal glands to the ICM, as later described by Coe (1931). Myoisophagos sanguineus (Rathke, 1799), new combination Figs. 5, 8-11 Planaria sanguinea Rathke, 1799:83. Lineus sanguineus McIntosh, 1873-1874: 188-190. auctt. Lineus nigricans Birger, 1892:159. auctt. not L. nigricans Isler 1901, 1902. Nemertes socialis Leidy, 1855:143; Verrill, 1873:324, 392, 628. Lineus socialis Verrill 1892:424—425. auctt. Lineus ruber, forme 8 Oxner 1909, 1910; Nusbaum & Oxner, 1910a, 1910b. not Lineus ruber (Miller, 1771). Lineus vegetus Coe, 1931:54-60. auctt. Species diagnosis.—Gliding individuals may attain 20 cm in length and 1.2 mm diameter. Brain region red, body of small individuals (less than 15 mm long) trans- parent white or cream color, larger speci- 549 mens vary from greyish green to brownish black or red. Three to seven ocelli in dorsal wall of cephalic fissures on each side, oc- casionally the same number on both sides. Anterior end of buccal cavity close to pos- terior ends of cerebral organs. Subepidermal glands do not penetrate the inner circular muscles. Subepidermal longitudinal muscle bundles are almost as large (Fig. 11) as the bundles of the OLM. Ganglion layer of brain enclosed with connective tissue (Fig. 4). Ex- cretory system usually with a short segment in dorsal wall of right preoral blood lacuna. Sexual reproduction abortive, oocytes ab- errant, fissiparous. Contract by spiral coil- ing. Dr. Janet Moore generously recorded length versus color and ocelli number of living specimens at Kaikoura. She noted that individuals under 20 mm, “slightly extend- ed,’ were white with three ocelli on each side; specimens from 25-35 mm were red with 3-6 ocelli per side, and beyond that were “very dark’ with 5-7 eyes on each side. This corresponds with personal obser- vations of individuals in the Gulf of Maine and with regenerating fragments of the other “species.”’ Specimens collected from mussel beds on exposed cliff faces at Nahant are frequently bright red while those beneath rocks on the nearby reef vary much like the New Zealand individuals. Riser (1993) not- ed that in culture all of these “‘species” were indistinguishable by color when fed a diet of Mytillus/Buccinum gonad/digestive gland. The uroporphyrin pigment (Vernet & Bierne 1983, Vernet 1983) diminishes in regenerating fragments, and the very small and widely scattered pigment granules are not readily evident in histological prepa- rations of regenerates that have not begun feeding. Regenerating individuals usually pass through a color phase similar to figure 12 in the description of Lineus flammeus Monastero, 1930. The closeness of the buc- cal cavity to the cerebral organs is obvious in living animals. Preoral contraction may alter the shape of the mouth from elongate to round; draw 550 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Kj f : yee (Lt af os Ri “sy ay, S ay KS 4 { Sis = a Y Py bs Fey é i ' y || py Tay) ; Ay aT (eligi OAS, ‘ ie 4 md | a Py i ty ‘ci ule Dad i ‘cae SA, ee ald iene Pu | ’ LN , ; ‘ uf i a wi ; wh wend af ” hs r uy = an 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 man- uscripts by Society members will be accepted. Papers are published in English (except for Latin diagnoses/descriptions of plant taxa), with an Abstract in an alternate language when appropri- ate. Submission of manuscripts.—Submit three copies of each manuscript in the style of the Proceedings to the Editor, complete with tables, figure captions, and figures (plus originals of the illustrations). Mail directly to: Editor, Proceedings of the Biological Society of Washing- ton, National Museum of Natural History NHB-108, Smithsonian Institution, Washington, D.C. 20560. (Do not submit manuscripts to an associate editor). 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 and diagnoses. The style for the Proceedings is described in “GUIDELINES FOR MANUSCRIPTS for Publications of the BIOLOGICAL SOCIETY OF WASHINGTON” a supplement to Volume 103, number 1, March 1990. Authors are encouraged to consult this article before manuscript preparation. Copies of the article are available from the editor or any associate editor. The establishment of new taxa must conform with the requirements of appropriate interna- tional codes of nomenclature. Decisions of the editor about style also are guided by the General Recommendations (Appendix E) of the International Code of Zoological Nomenclature. When appropriate, accounts of new taxa must cite a type specimen deposited in an institutional col- lection. Review.—One of the Society’s aims is to give its members an opportunity for prompt publi- cation of their shorter contributions. Manuscripts are reviewed by a board of Associate Editors and appropriate referees. Proofs.—Authors will receive first proofs and original manuscript for correction and ap- proval. Both must be returned within 48 hours to the Editor. Reprint orders are taken with re- turned proofs. Publication charges.—Authors are required to pay full costs of figures, tables, changes in proofs ($3.00 per change or revision), and reprints. Authors are also asked to assume costs of page-charges. The Society, on request, will subsidize a limited number of contributions per vol- ume. If subsidized manuscripts result in more than 12 printed pages, the additional pages must be financed by the author(s). Multiple authorship will not alter the 12 page limit (each author will be viewed as having used his/her 12 subsidized pages). Payment of full costs will facilitate speedy publication. Costs.—Printed pages @ $60.00, figures @ $10.00, tabular material @ $3.00 per printed inch per column. One ms. page = approximately 0.4 printed page. CONTENTS Description of a new species of subfossil eagle from Madagascar: Stephanoaetus (Aves: Fal- coniformes) from the deposits of Ampasambazimba Steven M. Goodman A giant Presbyornis (Aves: Anseriformes) and other birds from the Paleocene Aquia Forma- tion of Maryland and Virginia Storrs L. Olson A new species of owl of the genus Bubo from the Pleistocene of Cuba (Aves: Strigiformes) Oscar Arredondo and Storrs L. Olson Xenisthmus balius, anew species of fish from the Persian Gulf (Gobioidei: Xenisthmidae) Anthony C. Gill and John E. Randall Crabs of the family Homolodromiidae (Crustacea: Decapoda: Brachyura), V. Dicranodromia spinosa, anew species from the western Atlantic Joel W. Martin Glyphocrangon fimbriata, a new species of caridean shrimp (Crustacea: Decapoda: Glypho- crangonidae) from Sio Guyot, Mid-Pacific Mountains Tomoyuki Komai and Ichiro Takeuchi Occurrence of two lithodid crabs (Crustacea: Decapoda: Lithodidae) in the cold seep zone of the South Barbados accretionary prism Enrique Macpherson A new genus and two new species of deep-water hermit crabs (Decapoda: Anomura: Paguri- dae) from the Southern Ocean Patsy A. McLaughlin Marine isopods from the Lesser Antilles and Colombia (Crustacea: Peracarida) Brian Kensley and Marilyn Schotte New species of Echiniscus (Heterotardigrada: Echiniscoidea: Echiniscidae) from Korea Seung Y. Moon and Won Kim Doryphoribius koreanus, anew species of Tardigrada from Korea Seung Y. Moon, Won Kim, and Roberto Bertolani Amphitrite lobocephala, a new species (Polychaeta: Terebellidae) from Taiwan Hwey-Lian Hsieh Two new species of Cirrophorus (Polychaeta: Paraonidae) from the northern Gulf of Mexico Jerry A. McLelland and Gary R. Gaston Two new scale-worms (Polynoidae: Polychaeta) from the Lau Back-Are and North Fiji Basins, South Pacific Ocean Tomoyuki Miura Electra venturaensis, anew species (Bryozoa: Cheilostomata: Membraniporidae) from south- ern California William C. Banta and Mae M. Crosby The morphology and generic relationships of some fissiparous heteronemertines Nathan W. Riser Cololobus, Pseudopiptocarpha, and Trepadonia, three new genera from South America (Ver- nonieae: Asteraceae) Harold Robinson Biological Society of Washington: 121st Annual Meeting International Commission on Zoological Nomenclature 421 429 570 ~PROCEEDINGS _ OF THE BIOLOGICAL SOCIETY OF WASHINGTON THE BIOLOGICAL SOCIETY OF WASHINGTON 1994-1995 Officers President: Janet W. Reid Secretary: Carole C. Baldwin President-elect: Stephen D. Cairns Treasurer: T. Chad Walter Elected Council Robert J. Emry Susan L. Jewett Richard C. Froeschner Lynne R. Parenti Alfred L. Gardner F. Christian Thompson Custodian of Publications: Austin B. Williams PROCEEDINGS Editor: C. Brian Robbins Associate Editors Classical Languages: George C. Steyskal Invertebrates: Jon L. Norenburg Frank D. Ferrari Plants: David B. Lellinger Rafael Lemaitre Insects: Wayne N. Mathis Vertebrates: Thomas A. Munroe Membership in the Society is open to anyone who wishes to join. There are no prerequisites. Annual dues of $25.00 (for USA and non-USA addresses) include subscription to the Proceed- ings of the Biological Society of Washington. Library subscriptions to the Proceedings are: $40.00 for USA and non-USA addresses. Non-USA members or subscribers may pay an addi- tional $25.00 to receive the Proceedings by Air Mail. The Proceedings of the Biological Society of Washington (USPS 404-750) is issued quarterly. Back issues of the Proceedings and the Bulletin of the Biological Society of Washington (issued sporadically) are available. Correspondence dealing with membership and subscriptions should be sent to the Biological Society of Washington, P.O. Box 1897, Lawrence, Kansas 66044, U.S.A. Payment for membership is accepted in US dollars (cash or postal money order), checks on US banks, or MASTERCARD or VISA credit cards. Manuscripts, corrected proofs, editorial questions should be sent to the Editor, Biological Society of Washington, National Museum of Natural History, Smithsonian Institution, Wash- ington, D.C. 20560. Known office of publication: National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560. Printed for the Society by Allen Press, Inc., Lawrence, Kansas 66044 Second class postage paid at Washington, D.C., and additional mailing office. POSTMASTER: Send address changes to PROCEEDINGS OF THE BIOLOGICAL SOCI- ETY OF WASHINGTON, P.O. Box 1897, Lawrence, Kansas 66044. This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper). PROC. BIOL. SOC. WASH. 107(4), 1994, pp. 573-585 CHESAPEAKE BAY PHYTOPLANKTON: I. COMPOSITION Harold G. Marshall Abstract. —A list of 708 phytoplankton taxa from 10 algal divisions has been compiled for the Chesapeake Bay and is based on the analysis of 2384 water samples over three decades of study. The flora represents a diverse assemblage of algae, dominated annually by a variety of diatoms and phytoflagellates. Chesapeake Bay is the largest estuary in the United States. It is a temperate estuary, located within Virginia and Maryland, with a drainage basin: of 165,760 km? (Wright & Phillips 1988). Schubel & Pritchard (1987) characterize the Bay as having a mean depth of 8.42 m, a surface area of 6.5 x 10? km?, and a north-south orientation of about 320 km. Pritchard (1952) stated that the Bay has a net surface flow seaward and a net bottom flow of higher saline waters landward. The turbidity maximum is located north of the Patapsco River, in the upper portion of the Bay (Wright & Phillips 1988). The stratifi- cation and vertical homogeneity in the Bay are influenced by monthly flow rates, tem- peratures, tides, storms and the Coreolis ef- fect (Haas 1977, Wright & Phillips 1988, Schubel & Pritchard 1987, Pritchard 1952). Highest freshwater influx follows the spring rains and occurs in March and April, with the lowest rates from August to October. Schubel & Pritchard (1987) classify Ches- apeake Bay as a partially mixed estuary. It has at different times seasonally, and in dif- ferent segments of the Bay, either different degrees of vertical homogeneity, or strati- fication, and is dissimilar to the character- istic salt wedge estuary. The presence of the pycnocline will vary seasonally, being most developed during the summer months. Sub- pycnocline transport is a mechanism in which phytoplankton species may be dis- tributed up the Bay (Tyler & Seliger 1978). The earliest study of phytoplankton com- position and abundance in Chesapeake Bay was by Wolfe et al. (1926), reporting on sev- eral seasonal collections taken along the length of the Bay between 1916 and 1922. They collected whole water samples, fol- lowed by a centrifugation procedure to ob- tain a concentrate from which a sub-sample was analyzed. Their taxon list for Chesa- peake Bay included 83 diatoms, 12 dino- flagellates and 4 chrysophytes (total = 99), with late spring and fall maxima, dominated by Skeletonema costatum and Prorocen- trum micans respectively. Cowles (1930) discussed this data set in more detail and made geographic groupings of species, not- ing that temperate neritic forms were most abundant, with the phytoplankton having frequently higher cell concentrations along the western side of the Bay. Three decades later, Griffith (1961) prepared an illustrated guide of phytoplankton genera in Chesa- peake Bay and its tributaries. These were listed under 4 taxonomic categories that in- cluded 217 diatoms, 80 dinoflagellates, 33 cyanobacteria (cyanophytes), and 16 chlo- rophytes. The first comprehensive study of phyto- plankton composition and abundance in the lower Chesapeake Bay was by Patten et al. (1963) Using both whole water and net col- lections, they compared 24 data sets taken at 5 station locations over a 12 month pe- riod. Their net collections contained rep- resentative taxa from only the diatoms (123) and dinoflagellates (12). Greater species 574 representation, over a broader size range of cells, was obtained from their set of whole water samples. In these, they noted 107 taxa, composed of 57 diatoms, 33 dinoflagellates, 2 euglenoids, 7 chrysophytes (which includ- ed 2 cryptomonad taxa), and 8 chloro- phytes, with highest cell counts along the western Bay in the vicinity of the tributaries. Their results also indicated multiple sea- sonal maxima during the year. Earlier re- ports on phytoplankton data from these same cruises were made by Mulford (1962, 1963). Other Bay surveys were conducted by Marshall (1966), using data from a single cruise, and Whaley & Taylor (1968), who conducted a one year survey in the Bay us- ing net collections. They recorded 63 dia- toms, 19 dinoflagellates, 10 chlorophytes, and one cyanobacterium, with 28 of these as generic listings only. Mulford (1972) commented on the status of phytoplankton studies in Chesapeake Bay and described 36 dominant phytoplankters. These included 1 silicoflagellate, 28 dia- toms, and 7 dinoflagellates. In a later study and using concentrates from whole water samples, Marshall (1980) reported 219 taxa in the lower Chesapeake Bay during a 14 month study. These included 129 diatoms, 42 dinoflagellates, 8 chrysophytes, 7 eugle- noids, 7 prasinophytes, 2 xanthophytes, 8 cyanophytes, 6 haptophytes, 3 chloro- phytes, and 7 cryptomonads. Additional pa- pers on the composition and abundance of Chesapeake Bay phytoplankton include those by Marshall (1982, 1989, 1991, 1992), Marshall & Lacouture (1986), and Marshall & Alden (1990a, 1993). The importance of the Bay’s nano-pico- plankton components has been emphasized by McCarthy et al. (1974), Van Valkenburg & Flemer (1974), Marshall (1980) and Af- fronti & Marshall (1993). Specific phyto- plankton relationships to ecological vari- ables in the Bay have been addressed by Fisher et al. (1988, 1992), Harding et al. (1986), Loftus et al. (1972), Magnien et al. (1992), Marshall (1992), Marshall & Alden PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON (1991), among others. Productivity reports for the Bay region have included those by McCarthy et al. (1974), Sellner (1987), Mar- shall (1992) and Marshall & Nesius (1993). The Chesapeake Bay flora is presently dominated by diatoms that are responsible for high periods of growth during late win- ter-spring, summer, and fall (Marshall 1992). Dinoflagellates, cryptomonads, and auto- trophic picoplankters (e.g., cyanobacteria) are dominant categories during the summer and early fall. Although the shelf waters en- tering the Bay provide for an intrusion of neritic species, these are mainly concen- trated at the Bay entrance, along the south- eastern side of the Bay, and in sub-pycno- cline waters throughout the lower Bay (Marshall & Alden 1990a). In this lower strata, they may be transported not only up the Bay, but may also be introduced into adjacent tributaries (Marshall & Alden 1990b). Different degrees of turbulence are found in this system, which allow these spe- cies to be mixed with waters above the pyc- nocline. Thus, these waters contain a gen- eral mixture of riverine, estuarine, and neritic species. The degree to which this mixture changes in composition throughout the year is influenced by the spring rains and subsequent flow rates in the rivers and Bay. In addition, climatic and/or local con- ditions may influence turbulence and mix- ing of the water column. Within this system there is a seasonal succession of species within the Bay, as well as spatial distribu- tion patterns that occur annually (Marshall & Alden 1990a). The purpose of this paper is to provide a comprehensive list of phytoplankton taxa that have been observed by the author in Chesapeake Bay. This information has been derived from the analysis of 2384 water samples collected between 1963 and 1993 and analyzed in the author’s phytoplankton laboratory at Old Dominion University. Samples collected prior to 1985 came main- ly from seasonal studies or single cruise col- lections by the author. Since 1985, water VOLUME 107, NUMBER 4 samples were obtained consistently every month above and below the pycnocline at stations in the lower Bay as part of the Ches- apeake Bay Plankton Monitoring Program (Marshall 1992). This list is intended to pro- vide the reader with an overall representa- tion of the phytoplankton that currently characterize the Chesapeake Bay flora. It is expected that other species will subsequent- ly be added to this list. Methods All collections that were made and ana- lyzed in this study were whole water sam- ples taken by either hydrocasts with collec- tion bottles, or by using a pump to obtain water from various depths, to subsequently obtain 0.5—1.0 liters of water for analysis. Over the years of collections, the samples were preserved in either acidic or neutral solutions of Lugol’s (Throndsen 1978), or buffered formalin. These samples were pro- cessed through a settling and siphoning pro- cedure to obtain a final concentrate that was examined with an inverted plankton micro- scope using a modified Utermohl technique (Marshall & Alden 1990a). Cell identifica- tions and counts were made at either 125 x, 350, or 500. In addition, a scanning electron microscope (SEM) was used when needed to identify species that were not clearly defined with light microscopy. Auto- trophic picoplankters were also enumerated using epifluorescence microscopy (Affronti & Marshall 1993). This report is based on the examination of 610 samples prior to 1985 and 1774 samples taken during the following 8 years. The classification system used here is pat- terned after Van Landingham (1967-1979), Parke & Dixon (1976), and Hartley (1986). Taxonomic references include Lebour (1925), Hustedt (1927-1966, 1955), Ge- meinhardt (1930), Schiller (1930, 1933- 1937), Geitler (1932), Cupp (1943), Prescott (1951), Butcher (1959), Hendey (1964), Hulburt (1965), McIntyre & Bé (1967), Fott 575 (1968), Campbell (1973), and Dodge (1982). Also used were the various volumes of Siiss- wasserflora von Mitteleuropa, Das Phyto- plankton des Susswassers, and the series in Rabenhorst’s Kryptogamen-Flora. There were also numerous journal publications (e.g., Journal of Phycology, Phycologia, Di- atom Research, etc.), on specific taxa that are too numerous to list here, but represent essential reading to the systematist and may be obtained through routine searches of the literature. The reader must also contend with the changing systematics that is associated with several of the phytoplankton taxa and realize many present listings are destined to be changed. A major revision of the cyano- bacteria (Cyanophyceae) is given in the se- ries by Anagnostidis & Komarek (1985, 1988, 1990) and Komarek & Anagnostidis (1986, 1989). The classification of the cryp- tophyceae is also under revision; suggested references within this group include those by Butcher (1967), Anton & Duthie (1981), Santore (1984, 1985), Hull & Wetherbee (1989), and Hill (1991). Other categories in need of more taxonomic work are the var- ious groups often included under the col- lective term naked microflagellates. There are also several genera that are traditionally included among the phytoplankton (e.g., Calycomonas, Dictyocha, Distephanus) whose status with the algal flora is ques- tionable. Other references on classification of marine phytoplankton include Chrétien- not-Dinet et al. (1993) and Tomas (1993). Results A total of 708 phytoplankton taxa have been identified from water samples taken from Chesapeake Bay. These include 360 Bacillariophyceae, 125 Dinophyceae, 87 Chlorophyceae, 38 Cyanophyceae (cyano- bacteria), 24 Haptophyceae, 22 Eugleno- phyceae, 22 Chrysophyceae, 17 Crypto- phyceae, 9 Prasinophyceae and 4 Xantho- phyceae. Within the Phytoplankton Moni- toring data set from 1985 through 1992, 21 576 Table 1.— Phytoplankton that occurred in more than 50% of the water samples taken above and below the pycnocline from Chesapeake Bay, 1985-1992. I. Diatoms Asterionella glacialis 51.0% Cerataulina pelagica 65.5 Chaetoceros pendulus 51.8 Cyclotella spp. 86.3 Cylindrotheca closterium 76.6 Ditylum brightwellii 60.5 Leptocylindrus minimus 62.6 Pseudonitzschia pungens 61.3 Pleurosigma angulatum 53.5 Rhizosolenia calcar-avis 66.4 Rhizosolenia delicatula 54.3 Rhizosolenia setigera 50.0 Skeletonema costatum 84.0 Thalassionema nitzschioides 89.4 II. Dinoflagellates Ceratium lineatum 52.5% Gymnodinium spp. 83.1 Gyrodinium spp. 63.4 Katodinium rotundatum 60.6 Prorocentrum micans 84.0 Prorocentrum minimum 68.9 Protoperidinium spp. 59.9 taxa were found in greater than 50% of the 1774 samples, and are considered ubiqui- tous (Table 1). These consist of 14 diatom and 7 dinoflagellate taxa that represent many of the most common phytoplankters in the Bay. Not included in this list are other ubiq- uitous forms, often collectively counted in routine sample analysis under broad cate- gories, or size groups, e.g., autotrophic pi- coplankton, cryptomonads, and unidenti- fied microflagellates. Among the most common species throughout the study were the diatoms Thalassionema nitzschioides, Skeletonema costatum, and Cyclotella spp.., plus the dinoflagellates Prorocentrum mi- cans, P. minimum, and Katodinium rotun- datum. A total of 38 taxa were recorded in 25% of the samples, and 128 species (18.1%) had only one record of occurrence in the Bay. These results indicate that the plank- ton flora in the Bay is dominated by a small PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON number of species that occur annually and that these are accompanied by a large va- riety of background species that change sea- sonally. Occasionally, several of these back- ground species produce pulses of rapid growth and abundance. Generally, these ex- pressions are of short duration, and are not consistently produced in following years. The list also includes numerous benthic spe- cies (pennate diatoms), whose presence in the water column is not common. These periodically may appear in water samples and are often associated with areas of tur- bulence, or with active current flow. Over the period of these collections, the number of taxa recorded per sample (0.5 1) ranged from 20 to 92, with a mean of 43. Several common producers of sporadic blooms in the Bay, which are usually of lim- ited scope and duration, are Ceratium furca, Cochlodinium heterolobatum, Gymnodini- um nelsoni, Noctiluca scintillans, and Pro- rocentrum minimum. Flora associated with longer lasting seasonal expressions of abun- dance include Asterionella glacialis, Cyclo- tella spp., Heterocapsa triquetra, Katodi- nium rotundatum, Leptocylindrus minimus, and Skeletonema costatum. Although 6 or 7 seasonal assemblages may occur annually within the Bay (Marshall & Alden 1990a), there tends to be an overlying composition of “cold” and “‘warm”’ water species cate- gories when these forms are most abundant. For instance, assemblages common in win- ter-spring may include Asterionella glaci- alis, Cerataulina pelagica, Heterocapsa tri- quetra, Leptocylindrus danicus, L. minimus, Paralia sulcata, Rhizosolenia fragilissima, R. setigera, Skeletonema costatum, and Pseudonitzschia pungens. A summer-early fall assemblage may consist of Chaetoceros affinis, C. compressus, C. constrictum, C. neogracillis, Cyclotella spp., Eucampia zo- diacus, Lauderia borealis, Lithodesmium undulatum, Rhizosolenia alata, and R. stol- terfothii. One of the most common diatom assem- VOLUME 107, NUMBER 4 blages in the Bay is a Cyclotella complex that is present in high concentrations (10° cells/l) throughout most of the year. This group is dominated by a small (<10 wm diam) species of Cyclotella that was previ- ously identified as C. caspia in the Chesa- peake Bay (Marshall 1980, Marshall & Al- den 1990b) and in European waters (Hasle 1962, Kiss et al. 1988). Hakansson et al. (1993) have compared several of the smaller Cyclotella species to the original type ma- terial for C. caspia Grunow. They conclud- ed that the species which is common in the Bay is distinct from C. caspia and that it should be classified as C. choctawhatchee- ana Prasad, with C. hakanssoniae Wendker as a synonym (see Prasad et al. 1990, for a description of C. choctawhatcheeana). It is C. choctawhatcheeana that dominates the higher (> 18 ppt) saline waters of the lower Bay; with this species, plus C. striata and C. meneghiniana, forming an abundant Cy- clotella complex in the lower saline regions of the Bay and in its tidal tributaries. There is considerable morphological variability among many of these small Cyclotella spe- cies, so that SEM usage is necessary for ac- curate identification. Further studies of this assemblage may reveal that additional va- rieties or species are present in the Bay. Several phytoplankton species that have been reported as potential toxin producers are included in this Bay list: 1) the dino- flagellates Cochlodinium heterolobatum, Dinophysis acuminata, D. acuta, D. fortii, Gyrodinium aureolum, Noctiluca scintil- lans, Prorocentrum micans, P. minimum, and 2) the diatoms Amphora coffeaeformis and Pseudonitzschia pseudodelicatissima. Although not observed in the current col- lections of the lower Chesapeake Bay, two other toxin-producing species have been re- ported from this system: the diatom Pseu- donitzschia pungens f. multiseries (Hasle) Hasle was noted years ago in a sample from the Bay (Hasle 1965), and the dinoflagellate Pfiesteria piscimorte was identified recently S17 from Jenkins Creek off the Choptank River (Blankenship 1993). The increased number of species reported here, in comparison to past studies in Ches- apeake Bay, is attributed to several factors. The long term monitoring of this flora has provided a larger number of samples for analysis. These have been collected over a broader areal coverage of the Bay, and spa- tially throughout the water column. The re- peated monthly collections, under a variety of changing environmental conditions, pro- vided species in the samples that frequently have more limited periods of growth and abundance and which easily could be missed in a less concentrated sampling program. For example, many of the earlier studies were made along limited cruise tracts, with fewer representative stations and more lim- ited temporal coverage. The use of nets in several of these earlier studies would have biased their results by omitting many of the smaller species. Throndsen (1978) has also indicated that the use of different fixatives will effect the preservation of different algal categories, which in turn may influence their identification. In this study, 577 of the 705 species (81.8%) were recorded more than once, with a mean count of 43 species found in each sample. This level of species rep- resentation is attributed to using whole wa- ter samples and to following an Utermohl methodology in the analysis. Additional SEM usage provided for the identification of species whose traits could not be ade- quately discernable with light microscopy. Acknowledgments Appreciation is given to the Virginia De- partment of Environmental Quality for their financial support in the Phytoplankton Monitoring Program of the lower Chesa- peake Bay and to the numerous graduate research assistants who contributed to this study from 1985 through 1992. Special thanks is given to S. Wendker for providing 578 additional identifications of benthic dia- toms from the lower Chesapeake Bay, to H. Hakansson for verification of C. choctaw- hatcheeana, and to T. West for her work with the data set. Phytoplankton Recorded from the Chesapeake Bay CHRYSOPHYTA I. Bacillariophyceae Centrales: Actinocyclus normanii (Gregory) Hustedt. Actinoptychus sp., A. senarius (Ehrenberg) Ehrenberg, A. undulatus (J. W. Bailey) Ralfs. Asteromphalus heptactis (Brébisson) Ralfs. Aulacoseira distans (Ehrenberg) Simonsen, A. granu- lata (Ehrenberg) Simonsen, A. granulata var. an- gustissima (O. Miller) Simonsen, A. islandica (O. Miiller) Simonsen, A. islandica var. helvetica (O. Miller) Simonsen. Auliscus sculptus (W. Smith) Ralfs. Bacteriastrum sp., B. delicatulum Cleve. Biddulphia sp., B. alternans (J. W. Bailey) Van Heurck, B. biddulphiana J. E. Smith, B. reticulum (Ehrenberg) Boyer. Campylosira sp., C. cymbelliformis (A. Schmidt) Gru- now. Cerataulina pelagica (Cleve) Hendey. Cerataulus radiatus (Roper) R. Ross. Chaetoceros sp., C. affinis Lauder, C. atlanticus Cleve, C. atlanticus var. neapolitana (Schroder) Hustedt, C. brevis Schutt, C. coarctatus Lauder, C. compressus Lauder, C. constrictus Gran, C. convolutus Castra- cane, C. costatus Pavillard, C. crinitus Schutt, C. danicus Cleve, C. debilis Cleve, C. decipiens Cleve, C. densus (Cleve) Cleve, C. didymus Ehrenberg, C. didymus var. protuberans (Lauder) Gran & Yendo, C. difficilis Cleve, C. laciniosus Schiitt, C. neogracilis Van Laningham, C. pelagicus Cleve, C. pendulum Karsten, C. peruvianus Brightwell, C. pseudocurvi- setus Mangin, C. rostratus Lauder, C. socialis Laud- er, C. subtilis Cleve. Climacodium frauenfeldianum Grunow. Corethron sp., C. criophilum Castracane. Coscinodiscus sp., C. asteromphalus Ehrenberg, C. cen- tralis Ehrenberg, C. concinnus Wm. Smith, C. granii Gough, C. kuetzingii A. Schmidt, C. marginatus Eh- renberg, C. nodulifer A. Schmidt, C. oculus-iridis Ehrenberg, C. perforatus var. cellulosus Grunow, C. radiatus Ehrenberg, C. subbulliens Jorgensen, C. wai- lesii Gran & Angst. Coscinosira sp., C. polychorda (Gran) Gran. Cyclostephanos sp., C. dubius (Fricke) Round. Cyclotella sp., C. atomus Hustedt, C. caspia Grunow, PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON C. cryptica Reimann, Lawin & Guillard, C. glome- rata Bachmann, C. choctawhatcheeana Prasad, C. meneghiniana Kiitzing, C. striata (Kiitzing) Gru- now. Dactyliosolen mediterraneus (H. Peragallo) H. Pera- gallo. Detonula confervacea (Cleve) Gran. Ditylum brightwellii (T. West) Grunow. Ellerbeckia arenaria (Moore) Crawford. Eucampia cornuta (Cleve) Grunow, E. zodiacus Eh- renberg. Guinardia flaccida (Castracane) H. Peragallo. Hemiaulus sp., H. hauckii Grunow, H. membranaceus Cleve, H. sinensis Greville. Lauderia borealis Gran. Leptocylindrus danicus Cleve, L. minimus Gran. Lithodesmium sp., L. undulatum Ehrenberg. Melosira dickiei (Thwaites) Kitzing, M. moniliformis (O. F. Miiller) C. Agardh, M. nummuloides Agardh, M. varians C. Agardh. Neodelpheneis pelagica Takano. Odontella sp., O. aurita (Lyngbye) Agardh, O. mobi- liensis (J. W. Bailey) Grunow, O. obtusa Kitzing, O. rhombus (Ehrenberg) Kiitzing, O. rhombus f. trigona (Cleve) R. Ross, O. sinensis (Greville) Grunow. Paralia sulcata (Ehrenberg) Cleve. Rhizosolenia sp., R. acuminata (H. Peragallo) H. & M. Peragallo, R. alata Brightwell, R. alata f. gracillima (Cleve) Grunow, R. alata f. indica (H. Peragallo) Gran, R. bergonii H. Peragallo, R. calcar-avis Schultze, R. castracanei H. Peragallo, R. delicatula Cleve, R. fragilissima Bergon, R. hebetata f. hiemalis Gran, R. hebetata f. semispina (Hensen) Gran, R. imbricata Brightwell, R. rhombus Karsten, R. ro- busta Norman, R. setigera Brightwell, R. stolterfothii H. Peragallo, R. styliformis Brightwell, R. styliformis var. /atissima Brightwell, R. styliformis var. longi- spina Hustedt. Schroederella delicatula (H. Peragallo) Pavillard. Skeletonema sp., S. costatum (Greville) Cleve, S. po- tamos (Weber) Hasle. Stephanodiscus sp., S. hantzschii Grunow. Stephanopyxis sp., S. palmeriana (Greville) Grunow, S. turris (Greville & Arnott) Ralfs. Streptotheca tamesis Shrubsole. Thalassiosira sp., T. aestivalis Gran & Angst, T. baltica (Grunow) Ostenfeld, T. decipiens (Grunow) Jorgen- sen, 7. delicatula Ostenfeld, T. eccentrica (Ehren- berg) Cleve, 7. gravida Cleve, T. lacustris (Grunow) Hasle & Fryxell T. nordenskioeldii Cleve, T. oestrupii var. venrickae Fryxell & Hasle, T. pseudonana (Hus- tedt) Hasle & Heimdal, 7. rotula Meunier, T. /ep- topus (Grunow) Fryxell & Hasle, T. subtilis (Osten- feld) Gran, T. tenera Proschkina-Laurenko T. visurgis Hustedt. Triceratium sp., T. acutum Ehrenberg, T. favus Ehren- berg. VOLUME 107, NUMBER 4 Pennales: Achnanthes sp., A. danica (Flégel) Grunow, A. deli- catula (Kiitzing) Grunow, A. lemmermannii Hus- tedt. Amphiprora sp., A. alata (Ehrenberg) Kiitzing. Amphora sp., A. angusta Gregory, A. coffeaeformis (C. Agardh) Kitzing, A. costata W. Smith, A. crassa Gregory, A. lineolata Ehrenberg, A. /uciae Cholnoky, A. obtusa Gregory, A. ovalis (Kiitzing) Kiitzing, A. proteus Gregory, A. rhombica Kitton, A. sabyii Salah. Asterionella sp., A. formosa Hassall, A. glacialis Cas- tracane, A. gracillima (Hantzsch) Heiberg. Bacillaria paxillifer (Miller) Hendey. Berkeleya rutilans (Yrentepohl) Grunow. Caloneis sp., C. lepidula (Grunow) Cleve. Campylodiscus echeneis Ehrenberg, C. limbatus Bré- bisson. Catenula adhaerens (Mereschkowsky) Mereschkow- sky. Cocconeis sp., C. costata Gregory, C. disculus (Schu- mann) Cleve, C. placentula Ehrenberg, C. scutellum Ehrenberg, C. scutellum var. ornata Grunow. Cylindrotheca closterium (Ehrenberg) Reiman & Lew- in. Cymbella sp., C. excisa Kitzing, C. tumida (Brébisson) Grunow. Delphineis surirella (Ehrenberg) G. Andrews. Denticula subtilis Grunow. Diatoma sp., D. hyemale (Roth) Heiberg, D. tenue C. Agardh. Dimeregramma minor (Gregory) Ralfs. Diploneis sp., D. bombus (Ehrenberg) Ehrenberg, D. crabro (Ehrenberg) Ehrenberg, D. elliptica (Kitzing) Cleve, D. litoralis (Donkin) Cleve, D. subcincta (A. Schmidt) Cleve. Eunotia sp. Fragilaria sp., F. crotonensis Kitton, F. oceanica Cleve, F. pinnata Ehrenberg, F. virescens Ralfs. Gomphonema sp., G. acuminatum Ehrenberg, G. ex- iguum Kitzing, G. sphaerophorum Ehrenberg. Grammatophora sp., G. marina (Lyngbye) Kiitzing. Gyrosigma sp., G. balticum (Ehrenberg) Rabenhorst, G. balticum var. similis (Grunow) Cleve, G. fasciola (Ehrenberg) Griffith & Henfrey, G. hippocampus (Eh- renberg) Hassall, G. macrum (W. Smith) Griffith & Henfrey, G. spenceri (Quekett) Griffith & Henfrey, G. wansbeckii (Donkin) Cleve. Hantzschia sp., H. amphioxys (Ehrenberg) Grunow, H. marina (Donkin) Grunow. Licmophora sp., L. abbreviata C. Agardh, L. flabellata (Greville) C. Agardh, L. paradoxa (Lyngbye) C. Agardh, L. paradoxa var. tincta (C. Agardh) Hustedt. Mastogloia sp., M. exigua Lewis. Meridion circulare (Greville) C. Agardh. Navicula sp., N. amphipleuroides Hustedt, N. apiculata Brébisson, N. arenaria Donkin, N. cancellata Don- kin, N. caterva Hohn & Hellerman, N. crucicula (W. 579 Smith) Donkin, N. cruciculoides Brockmann, N. de- lawarensis Grunow, N. digitoradiata (Gregory) Ralfs, N. directa (W. Smith) Ralfs, NV. distans (W. Smith) Ralfs, N. escorialis Simonsen, N. forcipata Greville, N. gastrum (Ehrenberg) Kiitzing, N. gracilis var. neg- lecta (Thwaites) Grunow, N. granulata J. W. Bailey, N. gregaria Donkin, N. hanseni Moller, N. humerosa Brébisson, N. /aevissima Kiitzing, N. longa (Gregory) Ralfs, N. Jundstroemii Cleve, N. maculosa Donkin, N. marina Ralfs, N. peregrina (Ehrenberg) Kiitzing, N. phyllepa Kitzing, N. placenta Ehrenberg, N. pusil- la W. Smith, N. rhynchocephala Kitzing, N. sali- narum Grunow, N. sovereignae Hustedt, N. tripunc- tata (O. F. Miller) Bory. Nitzschia sp., N. acicularis (Kiitzing) W. Smith, N. ber- gii A. Cleve-Euler, N. bilobata var. minor Grunow, N. calida Grunow, N. clausii Hantzsch, N. compressa (J. W. Bailey) Boyer, N. constricta (Kiitzing) Ralfs, N. dissipata (Kiitzing) Grunow, N. frustulum (Kit- zing) Grunow, N. gracillima Heiden & Kolbe, N. gracilis Hantzsch, N. granulata Grunow, N. hybrida Grunow, N. lanceolata W. Smith, N. liebethruthii Rabenhorst, N. linearis (C. Agardh) W. Smith, N. longissima (Brébisson) Grunow, NV. microcephala Grunow, JN. obtusa var. scalpelliformis Grunow, N. ovalis Arnott, N. palea (Kiitzing) W. Smith, N. pel- lucida Grunow, N. pusilla Grunow, N. sigma (Kiit- zing) W. Smith, N. spathulata Brébisson, N. socia- bilis Hustedt, N. valida Grunow, N. vermicularis (Kiuitzing) Hantzsch. Operhora martyi Héribaud, O. olsenii Miiller, O. schul- zili (Brockmann) Simonsen. Pinnularia sp., P. major (Kiitzing) Rabenhorst. Plagiogramma sp., Plagiogramma van-heurckii Gru- now. Pleurosigma sp., P. anculatum (Quekett) W. Smith, P. aestuarii (Brébisson) W. Smith, P. delicatulum W. Smith, P. elongatum W. Smith, P. obscurum W. Smith, P. rigidum W. Smith, P. salinarum (Grunow) Grunow, P. strigosum W. Smith. Pseudonitzschia fraudulenta (Cleve) Hasle, P. pseu- dodelicatissima (Hasle) Hasle, P. pungens f. pungens Grunow, P. salinarum (Grunow) Grunow, P. seriata (Cleve) H. Peragallo, P. subpacifica Hasle. Rhabdonema minutum Kiitzing. Rhaphoneis sp., R. amphiceros (Ehrenberg) Ehrenberg, R. nitida (Gregory) Grunow. Rhoicosphenia abbreviata (C. Agardh) Lange-Bertalot. Rhopalodia gibberula (Ehrenberg) O. Miiller, R. oper- culata (C. Agardh) Hakansson. Scoliotropis latestriata (Brébisson) Cleve. Stauroneis sp., S. membranacea (Cleve) F. W. Mills, SS. phoenicenteron (Nitzsch) Ehrenberg, S. salina W. Smith. Striatella sp., S. unipunctata (Lyngbye) C. Agardh. Surirella sp., S. fastuosa Ehrenberg, S. gemma Ehren- berg, S. ovalis Brébisson, S. robusta Ehrenberg, S. striatula Turpin. 580 Synedra sp., S. acus Kitzing, S. closterioides Grunow, S. crystallina (Agardh) Kiitzing, S. fasciculata (Agardh) Kiitzing, S. fulgens (Greville) W. Smith, S. ulna (Nitzsch) Ehrenberg, S. undulata (J. W. Bailey) W. Smith. Tabellaria sp., T. fenestrata (Lyngbye) Kitzing, T. floc- culosa var. asterionelloides Grunow. Tetracyclus sp. Thalassionema sp., T. nitzschioides (Grunow) Gru- now. Thalassiothrix sp., T. delicatula Cupp, T. frauenfeldii (Grunow) Grunow, T. mediterranea Pavillard. Tropidoneis sp., T. lepidoptera (Gregory) Cleve. II. Chrysophyceae Ochromonadales: Dinobryon sp., D. cylindricum Imhof. Mallomonas sp. Ochromonas sp., O. caroliniana Campbell, O. varia- bilis Meyer. Olisthodiscus carterae Hulburt, O. luteus N. Carter, O. magnus Hulburt. Synura sp., S. uvella Ehrenberg. Chromulinales: Apedinella spinifera (Throndsen) Throndsen. Calycomonas ovalis Wulff, C. wulffii Conrad & Kuf- ferath. Chromulina sp. Chloromonales: Vacuolaria virescens Cienkowsk1i. Dictyochales: Dictyocha fibula Ehrenberg. Distephanus speculum (Ehrenberg) Haekel. III. Xanthophyceae Mischococcales: Monodus guttula Butcher. Tribonematales: Tribonema sp., T. affine West, T. minus (Wille) Hazen. DINOPHYTA Dinophyceae Prorocentrales: Prorocentrum sp., P. aporum (J. Schiller) Dodge, P. balticum (Lohmann) Loeblich III, P. compressum (Bailey) Abé, P. dentatum Stein, P. gracile Schitt, P. lima (Ehrenberg) Dodge, P. micans Ehrenberg, P. minimum (Pavillard) Schiller, P. triestinum Schiller. Dinophysiales: Ceratocorys horrida Stein Dinophysis sp., D. acuminata Claparede & Lachmann, PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON D. acuta Ehrenberg, D. caudata Kent, D. fortii Pav- illard, D. norvegica Claparede & Lachmann, D. ovum Schitt, D. pulchella (Lebour) Balech, D. punctata Jorgensen, D. rotundata Claparade & Lachmann, D. schuettii Murray & Whitting. Ornithocercus sp., O. magnificus Stein. Phalacroma sp. Gymnodiniales: Amphidinium sp., A. bipes Herdman, A. crassum Loh- mann, A. /acustre Stein, A. longum Lohmann, 4A. operculatum Caparéde & Lachmann, A. ovoideum (Lemmermann) Lemmermann, A. sphenoides Wulff, A. steinii (Lemmermann) Kofoid & Swezy. Cochlodinium sp., C. brandtii Wulff, C. helicoides Le- bour, C. heterolobatum Sousa Silva. Gymnodinium sp., G. arcticum Wulff, G. danicans Campbell, G. galesianum Campbell, G. marinum Kent, G. nelsoni Martin, G. roseostigma Campbell, G. simplex (Lohmann) Kofoid & Swezy, G. splendens Lebour. Gyrodinium sp., G. aureolum Hulburt, G. dominans Campbell, G. estuariale Hulburt, G. fusiforme Ko- foid & Swezy, G. spirale (Bergh) Kofoid & Swezy, G. uncatenatum Campbell. Katodinium asymmetricum (Massart) Loeblich III, K. rotundatum (Lohmann) Loeblich III. Oxyrrhis marina Dujardin. Polykrikos kofoidii Chatton. Noctilucales: Noctiluca scintillans (Macartney) Ehrenberg. Pyrocystales: Dissodium asymmetricum (Mangin) Loeblich III. Peridiniales: Amphidoma sp. Ceratium sp., C. arietinum Cleve, C. extensum (Gour- ret) Cleve, C. furca (Ehrenberg) Claparede & Lach- mann, C. fusus (Ehrenberg) Dujardin, C. hirundi- nella (O. F. Miiler) Dujardin, C. horridum (Cleve) Gran, C. inflatum (Kofoid) Jorgensen, C. lineatum (Ehrenberg) Cleve, C. Jongipes (Bailey) Gran, C. macroceros (Ehrenberg) Vanhéffen, C. massiliense (Gourret) Jorgensen, C. minutum Jorgensen, C. se- taceum Jorgensen, C. teres Kofoid, C. tripos (O. F. Miller) Nitzsch. Corythodinium reticulatum (Stein) Taylor. Diplopsalis lenticula Bergh. Glenodinium gymnodinium Penard. Gonyaulax sp., G. verior Sournia, G. digitale (Pouchet) Kofoid, G. polyedra Stein, G. polygramma Stein, G. spinifera (Claparede & Lachmann) Diesing, G. tria- cantha Jorgensen. Heterocapsa triquetra (Ehrenberg) Stein. VOLUME 107, NUMBER 4 Micracanthodinium claytonii (R. W. Holmes) Balech. Oblea rotunda (Lebour) Balech. Oxytoxum sp., O. milneri Murray & Whitting, O. scep- trum (Stein) Schroder, O. scolopax Stein. Peridinium aciculiferum Lemmermann. Podolampas sp. Protoperidinium sp., P. bipes (Paulsen) Balech, P. breve (Paulsen) Balech, P. brevipes (Paulsen) Balech, P. ce- rasus (Paulson) Balech, P. claudicans (Paulsen) Bal- ech, P. conicoides (Paulsen) Balech, P. conicum (Gran) Balech, P. depressum (Bailey) Balech, P. diabolum (Cleve) Balech, P. divergens (Ehrenberg) Balech, P. excavatum (Martin) Balech, P. globulum (Stein) Bal- ech, P. granii (Ostenfeld) Balech, P. minutum (Ko- foid) Loeblich III, P. oblongum (Aurivillius) Parke & Dodge, P. oceanicum (Vanh6ffen) Balech, P. ova- tum Pouchet, P. pallidum (Ostenfeld) Balech, P. pel- lucidum Bergh, P. pentagonum (Gran) Balech, P. subinerme (Paulsen) Loeblich III, P. steinii (Jorgen- sen) Balech, P. thorianum (Paulsen) Balech. Pyrodinium bahamense Plate. Pyrophacus horologium Stein. Scrippsiella trochoidea (Stein) Loeblich III. Triadinium polyedricum (Pouchet) Dodge. Zygabikodinium lenticulatum (Paulsen) Loeblich & Loeblich III. HAPTOPHYTA Haptophyceae Isochrysidales: Emiliania huxleyi (Lohmann) Hay & Mohler. Genhyrocapsa oceanica Kamptner. Hymenomonas roseola Stein. Pleurochrysis carterae (Braarud & Fagerland) Chris- tensen. Coccosphaerales: Acanthoica quattrospina Lohmann. Algirosphaera robusta (Lohmann) Norris. Calciosolenia sp., C. granii Schiller, C. murrayi Gran. Discosphaera tubifera (Murray & Blackman) Ostenfeld. Michaelsarsia elegans Gran. Ophiaster hydroideus (Lohmann) Lohmann. Pontosphaera sp., P. syracusana Lohmann. Rhabdosphaera sp., R. claviger Murray & Blackman, R. hispida Lohmann. Scyphosphaera apsteinii Lohmann. Syracosphaera histrica Kamptner, S. pulchra Loh- mann. Prymnesiales: Chrysochromulina sp., C. minor Parke & Manton. Prymnesium parvum N. Carter. Pavlovales: Pavlova salina (N. Carter) Green. 581 CRYPTOPHYTA Cryptophyceae Cryptomonadales: Campylomonas reflexa (Skuja) Hill. Chilomonas sp. Chroomonas sp., C. amphioxeia (Conrad) Butcher, C. caroliniana Campbell, C. pusilla (Backman) Hap- pey-Wood, C. vectensis N. Carter. Cryptomonas sp., C. erosa Ehrenberg, C. ovata Ehren- berg, C. stigmatica Wislouch. Hemiselmis sp. H. virescens Droop. Hillea sp. Rhodomonas baltica Karsten, R. salina(Wislouch) Hill & Wetherbee. Teleaulax acuta (Butcher) Hill. CYANOPHYTA (CYANOBACTERIA) Cyanophyceae Chroococcales: Anacystis sp. Aphanothece sp. Aphanocapsa sp. Chroococcus dispersus (Keissler) Lemmermann, C. limneticus Lemmermann, C. limneticus var. elegans G. M. Smith. Gomphosphaeria sp., G. aponina Kitzing. Marssoniella elegans Lemmermann. Merismopedia sp., M. elegans A. Braun, M. glauca (Ehrenberg) Nageli, 7. punctata Meyen, M. tenuis- sima Lemmermann. Microcystis sp., M. aeruginosa Kitzing, M. incerta Lemmermann. Rhabdoderma sp. Rhabdogloea sp., R. fascicularis Lemmermann. Synechococcus sp. Nostocales: Anabaena sp., A. spiroides K\ebahn. Calothrix sp. Nostoc sp., N. commune Voucher. Richelia intracellularis Schmidt. Trichodesmium erythraeum (Ehrenberg) Gomont. Oscillatoriales: Lyngbya sp. Oscillatoria sp., O. limnetica Lemmermann, O. lutea C. Agardh, O. tenuis C. Agardh, O. submembranacea Ardiss & Strafford. Phormidium sp. Planktolyngbya contorta (Lemmermann) Anagnostidis & Komarek. Spirulina sp., S. subsalsa Oersted. 582 EUGLENOPHYYA Euglenophyceae Euglenales: Euglena sp., E. acus Ehrenberg, E. agiles Carter, E. ehrenbergii Klebs, E. deses Ehrenberg, E. mutabilis Schmitz, E. proxima Dangeard, E. spirogyra Ehren- berg. Eutreptia sp., E. lanowii Steuer, E. viridis Perty. Eutreptiella sp., E. marina Cunha. Phacus sp., P. lemmermannii (Swirenko) Skvortzow, P. longicauda (Ehrenberg) Dujardin, P. suecicus Lemmermann. Trachelomonas sp., T. hispida (Perty) Stein, T. hispida var. punctata Lemmermann, TJ. intermedia Dan- geard, 7. volvocina Ehrenberg. CHLOROPHYTA I. Chlorophyceae Volvocales: Chlamydomonas sp., C. vectensis Butcher. Dunaliella sp. Eudorina sp. Gonium sp. Volvox tertius A. Meyer. Zygnematales: Closterium sp., C. gracile Brébisson. Coelastrum sp., C. cambricum Archer. Cosmarium sp., C. turpinii Brébisson. Desmidium sp. Euastrum sp., E. abrurtum West & West. Staurastrum sp., S. americanum (West & West) G. M. Smith, S. paradoxum Meyen. Zygnema sp. Oedogoniales: Oedogonium sp. Chlorococcales: Actinastrum sp., A. hantzschii Lagerheim, A. hantzschii var. fluviatile Schroder. Ankistrodesmus sp., A. convolutus Corda, A. falcatus (Corda) Ralfs, A. falcatus var. acicularis (A. Braun) West, A. spiralis (Turner) Lemmermann. Arthrodesmus sp. Chlorella sp., C. vulgaris Beijerinck. Closteriopsis longissima Lemmermann. Crucigenia sp., C. apiculata (Llemmermann) Schmidle, C. crucifera (Wolle) Collins, C. fenestrata Schmidle, C. irregularis Wille, C. lauterbornii Schmidle, C. quadrata Morten, C. tetrapedia (Kirchner) West & West. Dictyosphaerium sp., D. planctonicum Tiffany & Ahls- trom, D. pulchellum Wood. Elakatothrix gelatinosa Wille. Errerella bornhemiensis Conrad. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Franceia ovalis (Francé) Lemmermann. Kirchneriella sp., K. contorta (Schmidle) Bohlin, K. /u- naris (Kirchner) Mobius. Micractinium sp., M. pusillum Fresenius. Microspora sp. Mougetia sp. Nannochloris atomus Butcher. Oocystis sp. Pediastrum biradiatum Meyen, P. duplex Meyen, P. duplex var. reticulatum Lagerheim, P. simplex (Mey- en) Lemmermann, P. simplex var. duodenarium (Bailey) Rabenhorst, P. tetras (Ehrenberg) Ralfs. Polyedriopsis spinulosa Schmidle. Quadrigula sp., Q. lacustris (Chodat) G. M. Smith. Scenedesmus sp., S. arcuatus Lemmermann, S. acu- minatus (Lagerheim) Chodat, S. armatus (Chodat) G. M. Smith, S. bijuga (Turpin) Lagerheim, S. bijuga var. alternans (Reinsch) Hansgirg, S. denticulatus Lagerheim, S. dimorphus (Turpin) Kiitzing, S. hys- trix Lagerheim, S. perforatus Lemmerman, S. quad- ricauda (Turpin) Brébisson. Selenastrum sp. Schroederia setigera (Schr6der) Lemmermann. Tetraédron sp., T. gracile (Reinsch) Hansgirg, T. mini- mum (A. Braun) Hansgirg, 7. pentaedricum West & West, T. regulare Kiitzing, T. regulare var. incus Teiling, T. trigonium (Nageli) Hansgirg. Tetradesmus smithii Prescott. Tetrastrum glabrum (Roll) Ahlstrom & Tiffany, T. staurogeniaeforme (Schréder) Lemmermann. II. Prasinophyceae Pyramimonadales: Pyramimonas sp., P. amylifera Conrad, P. grossii Parke, P. micron Conrad & Kufferath, P. obovata N. Carter, P. plurioculata Butcher. Prasinocladales: Tetraselmis sp., T. gracilis (Kylin) Butcher, T. macu- lata Butcher. OTHER CATEGORIES Unidentified microflagellates (<10 microns) Unidentified picoplankters (<2 microns) Literature Cited Affronti, L. F., & H. G. Marshall. 1993. Diel abun- dance and productivity patterns of autotrophic picoplankton in the lower Chesapeake Bay.— Journal of Plankton Research 15:1-8. Anagnostidis, K., & J. Komarek. 1985. Modern ap- proach to the classification system of cyano- phytes. 1. Introduction.—Archiv fir Hydro- biologie/Supplementband 71, Algological Studies 38/39:29 1-302. VOLUME 107, NUMBER 4 , & 1988. Modern approach to the classification system of cyanophytes. 3. Oscil- latoriales.—Archiv fur Hydrobiologie/Supple- mentband 80, Algological Studies 50—53:327— 472. , & 1990. Modern approach to the classification system of cyanophytes. 5.—Sti- gonematales.— Archiv fur Hydrobiologie, Sup- plementband 86, Algological Studies 59:1-73. Anton, A., & H. Duthie. 1981. Use of cluster analysis in the systematics of the algal genus Cryptom- onas.—Canadian Journal of Botany 59:992- 1002. Blankenship, K. 1993. Researchers discover phan- tom “fish killer’ algae in the Bay area.— Bay Journal, Alliance for the Chesapeake Bay, Bal- timore 3(2):1 & 10. Butcher, R. W. 1959. An introductory account of the smaller algae of British coastal waters. Part I. Introduction and Chlorophyceae. Fishery In- vestigations Series IV, Ministry of Agriculture, Fisheries and Food, London, 74 pp., 14 pl. . 1967. Anintroductory account of the smaller algae of British coastal waters. Part IV. Cryp- tophyceae. Fishery Investigations Series IV, Ministry of Agriculture, Fisheries and Food, London 54 pp., 20 pl. Campbell, P.H. 1973. Studies on brackish water phy- toplankton. Sea Grant Publications UNC.-SG- 73-07, University of North Carolina, Chapel Hill, 406 pp. Chrétiennot-Dinet, M.-J., A. Sournia, M. Ricard, & C. Billard. 1993. A classification of the marine phytoplankton of the world from class to ge- nus.—Phycologia 32:159-179. Cowles, R. P. 1930. A biological study of the offshore waters of Chesapeake Bay.— Bulletin U.S. Bu- reau of Fisheries 46:277-381. Cupp, E. E. 1943. Marine plankton diatoms of the west coast of North America. University of Cal- ifornia Press, Berkeley, 237 pp. Dodge, J.D. 1982. Marine dinoflagellates of the Brit- ish Isles. Her Majesty’s Stationary Office, Lon- don, 303 pp. Fisher, T. R., L. W. Harding, Jr., D. W. Stanley, & L. G. Ward. 1988. Phytoplankton, nutrients, and turbidity in the Chesapeake, Delaware and Hud- son estuaries. — Estuarine, Coastal and Shelf Sci- ence 27:61-93. , E.R. Peel, J. W. Ammerman, & L. W. Harding. 1992. Nutrient limitation of phytoplankton in Chesapeake Bay.— Marine Ecology Progressive Series 82:51-63. Fott, B. 1968. Das Phytoplankton des Siisswassers, Band XVI, 3. Teil, 2. Auflage, Cryptophy- ceae, Chloromonadophyceae, Dinophyceae. Schweizerbart’sche Verlagsbuchhandlung, Stuttgart, 322 pp. 583 Geitler, L. 1932. Cyanophyceae von Europa unter Beriksichtigung der anderen Kontinente. Pp. 1— 1196 in L. Rabenhorst, ed., Kryptogamen Flora von Deutschland, Osterreich und der Schweiz, Band XIV. Akademische Vertlagsgesellschaft, Leipzig. Gemeinhardt, K. 1930. Silicoflagellateae. Pp. 1-88 in L. Rabenhorst, ed., Kryptogamen Flora von Deutschland, Osterreich und der Schweiz, Band X, Section II. Akademische Vertlagsgesell- schaft, Leipzig. Griffith, R. E. 1961. Phytoplankton of the Chesa- peake Bay—an illustrated guide to the genera. Chesapeake Biological Laboratory Contribution 172, Maryland. Department of Research and Education, 79 pp. Haas, L.W. 1977. The effect of the spring-neap tidal cycle on the vertical salinity structure of the James, York, and Rappahannock Rivers, Vir- ginia, U.S.A.—Estuarine and Coastal Marine Science 5:486-496. Hakansson, H., S. Hajdu, P. Snoeijs, & L. Loginova. 1993. Cyclotella hakanssoniae Wendker and its relationship to C. caspia Grunow and other sim- ilar brackish water Cyclotella species.— Diatom Research 8:333-347. Harding, Jr. W. H., B. W. Meeson, & T. R. Fisher, Jr. 1986. Phytoplankton production in two east coast estuaries: Photosynthesis-light functions and patterns of carbon assimilation in Chesa- peake and Delaware Bays.—Estuarine, Coastal and Shelf Science 23:773-806. Hartley, B. 1986. A check-list of the freshwater, brackish and marine diatoms of the British Isles and adjoining coastal waters.—Journal of the Marine Biological Association of the United Kingdom 66:531-610. Hasle, G. R. 1962. Three Cyclotella species from marine localities studied in the light and elec- tron microscopes.— Nova Hedwigia 4:299-307. 1965. Nitzschia and Fragilariopsis species studied in the light and electron microscopes. II. The group Pseudonitzschia. —Skrifter utgitt av det Norske Videnskaps-Akademi, serie 18: 1-45, 17 pl. Hendey, N. I. 1964. An introductory account of the smaller algae of British coastal waters. Part V. Bacillariophyceae (Diatoms). Fishery Investi- gations Series IV, Ministry of Agriculture, Fish- eries and Food, London, 317 pp. Hill, D.R. 1991. A revised circumscription of Cryp- tomonas (Cryptophyceae) based on examina- tion of Australian strains. — Phycologia 30:170- 188. , & R. Wetherbee. 1989. A reappraisal of the genus Rhodomonas (Cryptophyceae). Phycolo- gia 28:143-158. Hulburt, E.M. 1965. Flagellates from brackish waters 584 in the vicinity of Woods Hole, Massachusetts. — Journal of Phycology 1:87-94. Hustedt, F. 1927-1966. Die Kieselalgen Deutsch- lands, Osterreichs und der Schweiz mit Bertick- sichtigung der Ubrigen Lander Europas sowie der angrenzenden Meeresgebiete. Jn L. Raben- horst, Kryptogamen-Flora von Deutschland, Osterreich und der Schweiz. Band VII, Teil 1. 925 pp. (1927-1930); Teil 2. 845 pp. (1931- 1959); Teil 3. 816 pp. (1961-1966). Akade- mische Verlagsgesellschaft, Leipzig. 1955. Marine littoral diatoms of North Car- olina.— Bulletin Duke University Marine Sta- tion 6:1-67. Kiss, K. T., M. Costa, R. Le Cohu, & M. Nausch. 1988. Cyclotella caspia (Bacillariophyceae) in some rivers and lakes in Europe (morphological observations).—Cryptogamie, Algologie 9:27- 42. Komarek, J., & K. Anagnostidis. 1986. Modern ap- proach to the classification system of cyano- phytes. 2. Chroococcales.— Archiv fir Hydro- biologie/Supplementband 73, Algological Studies 43:157—226. —_, & 1989. Modern approach to the classification system of cyanophytes. 4. Nosto- cales.— Archiv fiir Hydrobiologie/Supplement- band 82, Algological Studies 56:247-345. Lebour, M. V. 1925. The dinoflagellates of northern seas. Marine Biological Association of the Unit- ed Kingdom, Plymouth, 250 pp. Loftus, M. E., D. V. Subba Rao, & H. H. Seliger. 1972. Growth and dissipation of phytoplankton in Chesapeake Bay. I. Response to large pulse of rainfall.— Chesapeake Science 13:282-299. Magnien, R. E., R. M. Summers, & K. G. Sellner. 1992. External nutrient sources, internal nu- trient pools, and phytoplankton production in Chesapeake Bay.— Estuaries 15:497-516. Marshall, H. G. 1966. The distribution of phyto- plankton along a 140 mile transect in the Ches- apeake Bay.— Virginia Journal of Science 17: 105-119. 1980. Seasonal phytoplankton composition in the lower Chesapeake Bay and Old Plantation Creek, Cape Charles, Virginia. —Estuaries 3:207— 216. 1982. The composition of phytoplankton within the Chesapeake Bay plume and adjacent waters off the Virginia coast, U.S.A. —Estuarine, Coastal and Shelf Science 15:29-43. 1989. Phytoplankton composition at the en- trance of the Chesapeake Bay.—Journal of the Elisha Mitchell Scientific Society 104:81-89. 1991. Seasonal phytoplankton assemblages associated with the lower Chesapeake bay plume.—Journal of the Elisha Mitchell Scien- tific Society 107:105—114. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON . 1992. Phytoplankton monitoring program. — Pp. viii, & 1-108 in Virginia Chesapeake Bay Water Quality and Living Resources Monitor- ing Programs: Comprehensive Task Report, 1985-1989. Old Dominion University Applied Marine Research Laboratory, Tech. Rept. No. 848, June, Norfolk, Virginia, 365 pp. —, & R. W. Alden. 1990a. Spatial and temporal diatom assemblages and other phytoplankton within the lower Chesapeake Bay, U.S.A. Pp. 311-322 in H. Simola, ed., Proceedings of the 10th International Diatom Symposium. Koeltz Scientific Books, Koenigstein, Germany, 592 pp. ———,, 1990b. A comparison of phyto- plankton assemblages and environmental rela- tionships in three estuarine rivers of the lower Chesapeake Bay.—Estuaries 13:287—300. —_, & 1991. Phytoplankton abundance, composition and trends within the lower Ches- apeake Bay and three tributaries. Pp. 517-522 in J. Mihursky & A. Chaney, eds., New per- spectives in the Chesapeake System: a research and management partnership. Chesapeake Re- search Consortium, Solomons, Maryland. , & 1993. A comparison of phyto- plankton assemblages in the Chesapeake and Delaware estuaries (USA), with emphasis on di- atoms. —Hydrobiologia 269/270: 251-261. , & R. Lacouture. 1986. Seasonal patterns of growth and composition of phytoplankton in the lower Chesapeake Bay.— Estuarine, Coastal Shelf Science 23:115—130. ——., & K. K. Nesius. 1993. Seasonal relationships between phytoplankton composition, abun- dance, and primary productivity in three tidal rivers of the lower Chesapeake Bay.—Journal of the Elisha Mitchell Scientific Society 109:141— 151. McCarthy, J. J.. W. R. Taylor, & M. E. Loftus. 1974. Significance of nanoplankton in the Chesapeake Bay estuary and problems associated with the measurement of nanoplankton productivity. — Marine Biology 24:7-16. McIntyre, A., & A. Bé. 1967. Modern coccolitho- phoridae of the Atlantic Ocean. I. Placoliths and crytoliths.— Deep Sea Research 14:561-597. Mulford, R. 1962. Diatoms from Virginia tidal wa- ters.— Virginia Institute of Marine Sciences, Special Report, Gloucester Point, Virgina, 30: 1-33. 1963. The net phytoplankton taken in Vir- ginia tidal waters. — Virginia Institute of Marine Sciences, Special Report, Gloucester Point, Vir- ginia 43:1-22. 1972. Anannual plankton cycle on the Ches- apeake Bay in the vicinity of Calvert Cliffs, Maryland. June 1969—May 1970.— Proceedings VOLUME 107, NUMBER 4 of the Academy of Natural Sciences, Philadel- phia, 124:17—40. Parke, M., & P. Dixon. 1976. Check-list of British marine algae— Third revision. — Journal Marine Biological Association of the United Kingdom 56:527-594. Patten, B. C., R. A. Mulford, & J. E. Warinner. 1963. An annual phytoplankton cycle in the lower Chesapeake Bay. — Chesapeake Science 4:1—20. Prasad, A. K., J. A. Nienow, & R. J. Livingston. 1990. The genus Cyclotella (Bacillariophyta) in Choc- tawhatchee Bay, Florida, with special reference to C. striata and C. choctawhatcheeana sp. nov.— Phycologia 29:418-436. Prescott, G. W. 1951. Algae of the western Great Lakes area.—Cranbrook Institute of Science Bulletin 31:1—946. Pritchard, D. 1952. Salinity distribution and circu- lation in the Chesapeake Bay estuarine sys- tem.— Journal of Marine Research 11:106-123. Santore, U. 1984. Some aspects of taxonomy in the Cryptophyceae.— New Phytology 98:627-646. . 1985. A cytological survey of the genus Cryp- tomonas (Cryptophyceae) with comments on its taxonomy.—Archiv fiir Protistenkunde 130:1- 52. Schiller, J. 1930. Coccolithinae. Pp. 89-273 in L. Rabenhorst, ed., Kryptogamen Flora von Deutschland, Osterreich und der Schweiz. Band X, Section II, Akademische Verlagsgesellschaft, Leipzig. 1933-1937. Dinoflagellatae (Peridineae) in monographischer Behandlung. Jn L. Raben- horst, ed., Kryptogamen Flora von Deutsch- land, Osterreich und der Schweiz. Band X, Sec- tion III, Teil 1. 617 pp. (1933); Teil 2. 589 pp. (1937). Akademische Verlagsgesellschaft, Leip- Zig. Schubel, J. R., & D. W. Pritchard. 1987. A brief physical description of the Chesapeake Bay. Pp. 1-32 in S. Majumdar, L. Wall, & H. Austin, eds., Contaminant problems and management of living Chesapeake Bay resources. Pennsyl- vania Academy of Science, Philadelphia, 345 pp. Sellner, K. G., 1987. Phytoplankton in Chesapeake Bay: role in carbon, oxygen and nutrient dy- namics. Pp. 134-157 in S. Majumdar, L. Hall, 585 & H. Austin, eds., Contaminant problems and management of living Chesapeake Bay re- sources. Pennsylvania Academy of Science, Philadelphia, 345 pp. Throndsen, J. 1978. Preservation and storage. Pp. 69-74 in A. Sournia, ed., Phytoplankton man- ual. Monographs on Oceanographic Method- ology No. 6. United Nations Educational, Sci- entific and Cultural Organization. Page Brothers, Norwich, 337 pp. Tomas, C. R. ed. 1993. Marine phytoplankton. A guide to naked flagellates and coccolithophores. Academic Press, San Diego, 263 pp. Tyler, M. A., & J. H. Seliger. 1978. Annual subsurface transport ofa red tide dinoflagellate to its bloom area: water circulation patterns and organisms distribution in the Chesapeake Bay. —Limnol- ogy and Oceanography 23:227-246. Van Landingham, S. L. 1967-1979. Catalogue of the fossil and recent genera and species of diatoms and their synonyms. Part 1, 493 pp. (1967); Part 2, pp. 494-1086 (1968); Part 3, pp. 1087-1756 (1969); Part 4, pp. 1757-2385 (1971); Part 5, pp. 2386-2963 (1975); Part 6, pp. 2964-3605 (1978); Part 7, pp. 3606-4241 (1978); Part 8, pp. 4242-4654 (1979). Vaduz, J. Cramer. Van Valkenburg, S. D., & D. A. Flemer. 1974. The distribution and productivity of nannoplankton in a temperate estuarine area.—Estuarine and Coastal Marine Science 2:311-322. Whaley, R. C., & W. R. Taylor. 1968. A plankton survey of the Chesapeake Bay using a contin- uous underway sampling system.— Chesapeake Bay Institute, Technical Report, Johns Hopkins Univ. 36:1-93. Wolfe, J. J., B. Cunningham, F. Wilkerson, & J. T. Barnes. 1926. An investigation of the micro- plankton of Chesapeake Bay.—Journal of the Elisha Mitchell Scientific Society 42:25—54. Wright, D. A., & D. J. Phillips. 1988. Chesapeake and San Francisco Bays. A study in contrasts and parallels.— Marine Pollution Bulletin 19: 405-413. Department of Biological Sciences, Old Dominion University, Norfolk, Virginia 23529-0266, U.S.A. PROC. BIOL. SOC. WASH. 107(4), 1994, pp. 586-590 LETEPSAMMIA FRANKT, A NEW SPECIES OF DEEP-SEA CORAL (COELENTERATA: SCLERACTINIA: MICRABACIIDAE) Joan Murrell Owens Abstract.—Letepsammia franki, a new species of deep-sea, solitary corals be- longing to the family Micrabaciidae, is described and figured. This new species resembles the type species, L. formosissima (Moseley, 1876), in having highly perforated septa and wall, and well-developed deltas. It differs in the distinctly beaded appearance of its septa, deltas, and columella; its slightly biconvex corallum with prominent basal apex; its projection of costae beyond septa at the distal edge, forming a narrow marginal shelf; and its more open basal wall. It is Recent in origin and is found in the Indian Ocean off the southeast coast of Africa at depths varying from approximately 50 to 650 meters. It thus falls within both the geologic and bathymetric ranges of the type species, L. for- mosissima. Letepsammia was erected by Yabe & Eguchi in 1932, as a subgenus of Stephano- phyllia. They based this subgenus by mono- typy on the well-developed, spongy colu- mella, and highly perforated septa and wall of the subgenotype, S. formosissima Mose- ley. More recent authors, such as Squires (1965, 1967), Keller (1977), Cairns (1982), and Owens (1984a, 1984b, 1986a), how- ever, deemed these differences, along with its larger size, thinner base, and more prom- inent marginal shelf, sufficient to denote ge- neric differences, and informally acknowl- edged the subgenus as a genus in itself. Eventually, Owens (1986b) formally de- scribed and officially elevated Letepsammia to generic rank. At present, Letepsammia consists of only two species, L. formosissima and the new species described herein, L. franki. Order Scleractinia Suborder Fungiida Superfamily Fungioidea Vaughan & Wells, 1943 Family Micrabaciidae Vaughan, 1905 Genus Letepsammia Yabe & Eguchi, 1932 Letepsammia franki, new species Figs. 1, 2 Description. —Corallum large, loose, slightly biconvex, with narrow but promi- nent shelf. Wall thin, highly perforate. Ca- licular depression deep, narrow, elongate. Fusion of proximal margins of tertiary septa with inner edges of secondary septa form broad, porous, coarsely dentate deltoid structures. Diameter of specimens 10.5-31 mm, height 3-11 mm, average H:D ratio ().3)9). Costae long, thin, nearly smooth, thick- ening slightly distally. Intercostal loculi wider than costae, but interrupted by closely spaced synapticulae that connect adjacent costae with base of intervening septum. Outer margins of costae finely serrated. Cos- tae begin as six (first cycle) at apex of convex base and bifurcate immediately (second cy- cle); outer costae of each system bifurcate about one-eighth the distance from center, with inner pairs dividing soon after (third cycle); fourth and fifth cycles overlap, with outermost costae of each system undergoing VOLUME 107, NUMBER 4 587 408 Fig. 1. Letepsammia franki: A, Oral view of holotype (USNM 75638), Anton Bruun 390-S; B, Aboral view of holotype; C, Side view of a paratype (USNM 75639), Anton Bruun 390-S; D, Thin section of a paratype (USNM 75640), under reflected light, showing thin, widely spaced trabeculae, radial perforations, and coarse dentation of distal margin of septum, Anton Bruun 390-S. Scale bars = 2 mm. 588 Fig. 2. Diagrammatic representation of one system of Letepsammia franki. Heavy lines represent septa; thin lines, costae. 5x. two bifurcations before innermost again di- vide; sixth cycle incomplete, with only one outer pair and one inner pair of each system bifurcating (Fig. 2). Septa thin, highly perforate, and, except for primaries, irregularly lined with vepre- culae. Vepreculae closely and evenly spaced on primaries. Interspaces much broader than septa. Synapticulae scarce except along dis- tal base of septa. Primaries free, straight, lower than adjacent septa; finely serrated proximally but coarsely dentate with short, clubby spines near distal margin. Second- aries straight, as tall as neighboring septa, and dentate with short, broad spines along full length of margin. Adjacent septa in ter- tiary position unite proximally with sec- ondaries in conspicuous, triangular, trabec- ular masses that form broad, porous deltas. Tertiary septa do not extend to distal mar- gin, but instead give rise to a series of bi- furcations immediately beyond deltas to produce higher cycle septa, for a total of 120 septa. Columella spongy, elongate, narrow, with single row of stubby spines running full length. Trabeculae simple, wavy, grouped in three’s in loose fan system; interareas ar- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON cuate between groupings, undifferentiated within; perforations of various sizes roughly aligned horizontally and radially. Types. —Holotype: USNM 75638, Anton Bruun 390-S (29°35’'S, 31°42’E; 138 m). Paratypes: USNM 75639 (1), 75640 (36), Anton Bruun 390-S (same as holotype); 75641 (1), Anton Bruun 370-H (24°41’S, 35°28’E; 311-320 m); 75642 (16), Anton Bruun 370-G (24°40'S, 35°28’E; 635 m); 75643 (2), Anton Bruun 372-G (24°53’S, 34°56'E; 55 m); 75644 (6), Vema Cruise 14, Sat 6 (29°48'S, 31°16’E; 232 m). Occurrence. —Recent, Indian Ocean off southeast coast of Africa; 50-650 meters. Discussion. — Letepsammia franki resem- bles both Rhombopsammia niphada and L. formosissima, R niphada because of its slightly biconvex corallum, its deep, long, narrow calicular depression, and its narrow but prominent shelf; and L. formosissima because of its highly perforate septa and wall, its well-developed deltas, and the réduced, restricted trabecular pattern of its septa— similarities of generic significance. Letep- sammia franki is unique in having a papil- lose columella and coarse septal dentation that give its corallum a distinctly beaded appearance. Owens (1986a), in noting the similarities between L. formosissima and R. niphada, suggested that R. niphada may be a morphologic intermediate between Letep- sammia and Rhombopsammia. The simi- larities between L. franki and R. niphada strongly reinforce this suggestion. Cairns (1989), who had seen the speci- mens of L. franki deposited at the National Museum of Natural History, believed that specimens of L. formosissima from the western Indian Ocean referred to by Van der Horst (1927) and Boshoff (1981), and those from the Red Sea mentioned by Gar- diner & Waugh (1939) are probably L. fran- ki, the then undescribed species of Letep- sammia alluded to by both Squires (1967) and Owens (1986b). Cairns & Keller (1993) also remarked on the same undescribed spe- cies, which Cairns in a later personal com- VOLUME 107, NUMBER 4 munication considered to be this new spe- cies. Similarly, the “button coral” figured by Williams (1986), which he referred to as “senus Stephanophyllia,” is most likely L. franki. When Yabe & Eguchi (1932, 1934) erect- ed the subgenus Letepsammia, they includ- ed in their subgenus a fossil form from the Plio-Pleistocene boundary, Stephanophyl- lia (L.) japonica nov. This species was sub- sequently determined by Squires (ca. 1967) to be synonymous with S. superstes Ort- man, which he further reassigned as L. su- perstes. Thus, when Owens (1986b) for- mally described Letepsammia, she accepted Squires’ redesignation and included in the genus two species: L. formosissima and L. superstes. More recently, however, Cairns (1989) re- counted a personal communication with H. Zibrowius, in which Zibrowius stated that he had examined the holotype of S. su- perstes and believed it to be a juvenile form of L. formosissima. If these synonymies are correct, then the geologic range of L. for- mosissima must be extended to the Plio- Pleistocene boundary, and thus exceeds the geologic range of the new species, L. franki. Consequently, the genus Letepsammia presently includes only L. formosissima and L. franki. Etymology. —The species name is in hon- or of Frank A. Owens, my husband, whose photographs of specimens are an integral part of my published works. Acknowledgments I wish to thank S. D. Cairns of the Smith- sonian Institution for making available to me the specimens used in this study; D. A. Dean of the Smithsonian Institution for his assistance in thin-sectioning some of the specimens; and F. A. Owens and E. H. Hughes for photographing the specimens used in this paper. I also wish to make spe- cial acknowledgment of D. F. Squires whose unfinished work on micrabaciid corals in- 589 spired me to undertake my own study of this group. Literature Cited Boshoff, P. H. 1981. An annotated checklist of Southern African Scleractinia.—South African Association for Marine Biological Research, Oceanographic Research Institute, Investiga- tional Report 49:45 pp. Cairns, S. D. 1982. Antarctic and subantarctic Scle- ractinia.— Antarctic Research Series 5, 34(1):1- 74. . 1989. A revision of the ahermatypic Sclerac- tinia of the Phillipine Islands and adjacent wa- ters, Part I: Fungiacyathidae, Micrabaciidae, Turbinoliinae, Guyniidae, and Flabellidae.— Smithsonian Contributions to Zoology 486:15- 18. ——.,, & N. B. Keller. 1993. New taxa and distri- butional records of azooxanthellate Scleractinia (Cnidaria, Anthozoa) from the tropical south- west Indian Ocean, with comments on their zoo- geography and ecology.—Annals of the South African Museum 103(5), 1993:213-292. Gardiner, J. S., & P. Waugh. 1939. Madreporaria excluding Flabellidae and Turbinolidae.—Sci- entific Reports of the John Murray Expedition 1933-34, 6(5):225-242. 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. [In Russian.] Owens, J. M. 1984a. Microstructural changes in the Micrabaciidae and their ecologic and taxonomic implications.— Palaeontographica Americana 54:519-522. 1984b. Evolutionary trends in the Micra- baciidae: an argument in favor of preadapta- tion.—Geologos 2(1):87—93. - . 1986a. Rhombopsammia, a new genus of the family Micrabaciidae (Coelenterata: Scleracti- nia).— Proceedings of the Biological Society of Washington 99:248-256. 1986b. On the elevation of the Stephano- phyllia subgenus Letepsammia to generic rank (Coelenterata: Scleractinia: Micrabaciidae).— Proceedings of the Biological Society of Wash- ington 99:486—-488. Squires, D. F. 1965. A new record for Leptopenus, a rare deep-water coral.— Nature 207:878-879. 1967. The evolution of the deep-sea coral family Micrabaciidae.—Studies in Tropical Oceanography 5:502-510. . [ca. 1967]. Tertiary and extant Scleractinian corals of the family Micrabaciidae. Unpub- lished partial manuscript on deposit at the Na- 590 tional Museum of Natural History, Smithsonian Institution. Van der Horst, C. J. 1927. Eupsammid corals from South Africa.— Fisheries and Marine Biological Survey, Report Number 5, for the year 1925, 7 Williams, G. C. 1986. What are corals?— Sagittarius 1(2):11-15. Yabe, H., & M. Eguchi. 1932. Some Recent and fossil corals of the genus Stephanophyllia H. Michelin PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON from Japan.— Tohoku Imperial University Sci- entific Reports, series 2 (Geology), 15(2):55-63. . 1934. Probable generic identity of Stephano- phyllia Michelin and Micrabacia M. Edwards and J. Haime.—Proceedings of the Imperial Academy of Japan, 10(5):278-281. Department of Biology, Howard Univer- sity, Washington, D.C. 20059, U.S.A. PROC. BIOL. SOC. WASH. 107(4), 1994, pp. 591-599 SIDERASTREA GLYNNI, A NEW SPECIES OF SCLERACTINIAN CORAL (CNIDARIA: ANTHOZOA) FROM THE EASTERN PACIFIC Ann F. Budd and Hector M. Guzman Abstract. —A new species of Siderastrea (S. glynni) has been found in shallow (7-8.5 m deep) reef rubble north of Isla Uraba, Bay of Panama, in the eastern Pacific region. The species is extremely rare. Its colonies are unattached, sphe- roidal in shape, and approximately 7-10 cm in diameter. Siderastrea glynni is distinguished by relatively small corallites (2.5—-3.5 mm); numerous thin septa (40-48 per corallite); a porous columella; and a distinctive synapticular mesh- work. The discovery of the new species is unusual because the genus Siderastrea typically occurs today in the Atlantic and Indian Oceans and is known in the Pacific only from a rare occurrence in the Philippine Islands. A total of 19 species and seven genera of modern hermatypic scleractinian corals have been reported from the eastern Pacific coast of Panama (Holst & Guzman 1993). The 19 species consist of: two ““Cycloseris,”’ one Gardineroseris, five Pavona, five Pocillo- pora, two Porites, three Psammocora, and one Siderastrea. Six or more of the species are restricted to the extreme eastern Pacific (Wells 1983); however, the distributions of all seven genera except Siderastrea are wide- spread across the Indo-Pacific (Veron 1993). Here we describe an unusual new species of Siderastrea. It was recently discovered by one of us (HMG) while diving at depth of 7-8.5 m along an upper reef slope north of Isla Uraba in the Bay of Panama near the Pacific entrance to the Panama Canal (Fig. 1). Despite extensive search, only one pop- ulation of the species has been found. It originally consisted of five unattached col- onies, all of which were spheroidal in shape and approximately 7-10 cm in diameter (Fig. 2). The five colonies were found in a small patch (<8 m7) over coral rubble with- in a 1 m distance from one another. One of the five colonies was collected and is de- scribed below. Since field observations sug- gested that the species may be close to ex- tinction, the other four colonies were left alive at the original discovery site in an ef- fort to preserve the species. The rarity of this species is similar to Millepora boschmai Weerdt & Glynn, 1991, a recently discov- ered eastern Pacific species of Hydrozoa. The discovery of the Siderastrea glynni is particularly noteworthy because the genus occurs today mainly in the Atlantic and In- dian Oceans (Veron 1986). Although Veron (1986) alludes to possible Indo-Pacific oc- currences of S. radians, only one species, S. savignyana, 1s well-documented in the cen- tral Indo-Pacific, represented by one spec- imen from the Philippine Islands (Veron 1993). Another species of the genus (Side- rastrea mendenhalli), however, was ex- tremely abundant in south-central Califor- nia at the northernmost end of the Gulf of California during early Pliocene time (Fos- ter 1979, 1980a; Budd 1989). Comparisons among Siderastrea glynni, the two Pacific species noted above, and three modern and two fossil Atlantic species (Table 1) indicate that the new species, S. glynni, is morphologically unique. S. g/ynni differs from the modern and the fossil Pa- cific species primarily in its small, well- rounded, unattached colony shape and in 592 79 (pe Fig. 1. of 7-8.5 m on the upper reef slope north of Isla Uraba, Bay of Panama. its small corallite size and numerous septa (Fig. 3). It also has a relatively thinner wall and a shallower columellar fossa than mod- ern Indo-Pacific S. savignyana. Siderastrea glynni differs similarly from the two com- mon modern Caribbean species of Side- rastrea (S. radians, S. siderea) by having small, shallow calices and numerous thin septa (Fig. 4). Of the two modern Caribbean species, S. g/ynni is most similar to S. ra- dians, but differs in septal number and thickness, and in the development of the columella and synapticulae. Of the three Neogene Caribbean species in Table 1, Side- rastrea glynni is most similar to S. men- denhalli, the species noted above whose dis- tribution extended to the eastern Pacific. However, S. g/ynni has smaller corallites and fewer septa than S. mendenhalli. Pacific Ocean PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON |. Naos Boo |. Culebra? |. Changame ‘pl. Venado 3 |. Tortolita |. Cocori ¢ |. Tortola 8°50’ |. Melones |. Taboguilla f) . Farallon Uraba » |. Tarapa Type locality (black dot) of Siderastrea glynni. Five colonies were found in a small patch at a depth On the basis of these distributional and morphologic comparisons, we recognize S. glynni as sufficiently distinct to describe it below as a new species. Abbreviations of Repository Institu- tions. -USNM: U.S. National Museum of Natural History, Department of Inverte- brate Zoology, Smithsonian Institution, Washington, D.C. 20560. SUI: University of lowa, Department of Geology, Iowa City, IA 52242. Order Scleractinia Bourne, 1900 Suborder Fungiina Verrill, 1865 Family Siderastreidae Vaughan & Wells, 1943 Genus Siderastrea de Blainville, 1830 Type species. — Madrepora radians Pallas, 1766:322—323. The holotype is figured in VOLUME 107, NUMBER 4 Seba, 1756, pl. 122, figs. 12, 14, 18; and is currently lost. Diagnosis.— Massive, branching or en- crusting colonies. Cerioid corallites formed by extratentacular budding. Well-defined, synapticulothecate wall structure. Septa straight, generally not fusing. Distribution. —Cretaceous to Recent; Ca- ribbean, eastern and western Atlantic, Med- iterranean, Red Sea, Indian Ocean (Wells 1956, Chevalier 1961, Veron 1993). One specimen is reported in the Pacific from the Philippine Islands (Veron 1993). The new occurrence in the present report adds the eastern Pacific to the known distribution of the genus. Siderastrea glynni, new species Figs. 2, 3, 5, 6 Etymology. —Named after Peter W. Glynn for his pioneering work on eastern Pacific coral reefs. Diagnosis. —Small corallites. Well-devel- oped outer synapticular rings forming a dis- tinctive regular meshwork. Low. straight, moderately thick corallite wall. Numerous equally thin, dentate septa in four cycles, the last sometimes incomplete. Columella porous with a shallow fossa. Description. —Colonies massive, unat- tached, spheroidal; 7-10 cm in diameter; with a well-rounded, smooth outer surface. Calices hexagonal or pentagonal; relatively small in diameter (2.5—3.5 mm). Outermost synapticular rings regular and well-devel- oped, forming a distinctive meshwork on the upper calical surface. Corallite wall low, solid, straight, continuous, intermediate in thickness (~0.15 mm). Synapticulae ar- ranged in 3 or 4 rings, intermediate in thick- ness (~0.15 mm). Septa relatively thin, equal in thickness, usually continuous be- tween adjacent corallites. Septal margins strongly dentate, with 8-10 dentations per primary septum. Septal surfaces weakly or- namented. Four septal cycles, with the fourth cycle sometimes incomplete; 40-48 septa per corallite. First and second cycles free; 593 b Fig. 2. Siderastrea glynni. Holotype, USNM 93956. Recent, 7-8.5 m depth, Isla Uraba, Bay of Panama. (a) Whole colony with soft tissue soon after collection, x). (b) A 5 mm thick slab cut through the growth axis of the colony, x1. third cycle fuses with second near columel- la. Fourth cycle free, intermediate in length (~0.5 mm). Columella porous, papillose, intermediate in thickness (~0.4 mm). Ca- licular fossa shallow. Endothecal dissepi- ments thin, at 0.3-0.5 mm intervals. Holotype. —USNM 93956 (Figs. 2, 3b, 5, 6); collected 3 Sep 1992, by H. M. Guzman, at 7-8.5 m depth on the upper reef slope along the northern tip of Isla Uraba, Bay of Panama (Fig. 1). Material. —One colony: USNM 93956. Comparison. — Siderastrea glynni 1s mor- phologically most similar to S. radians which also forms spheroidal, unattached colonies with small corallites. However, it differs from S. radians by having more numerous, thin septa that are equal in thickness. The PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 594 € ‘71 6 9 ‘b ‘7 6 ‘9 ‘Pb 7 SOdT[2O U99Mj9q snonuluos Ale -nsn ejdos ‘ssul ‘deus ¢—p ‘yoru SooI[eO uoeMjJOq snonuT} -u0d ejdos ‘ssuLl ‘deuds ¢-¢ ‘ury} SOor[eO uooMjJoq snonuT} -u0d eydas ‘ssuLl ‘deus p—¢ ‘YoY? SOOdI[BD UI9A\} -9q SnonuUT}UOD Fe} -das ‘s3ut ‘deus €-Z “yory) Atoa SOdITBO U99M19qQ snonutjuoo Aye -nsn ejdos ‘ssu ‘deuAds p—¢ “uly Sol] -89 U99Mj0q 9}8U -J9}]@ eydas ‘ssuLs ‘deuds ¢—¢ ‘uly SOdI[2D U99M19qQ snonunuod Aye -nsn ejdos ‘ssul ‘deuds ¢-Z ‘4o1u} SddT[2D UI9MI19qQ snonutuoo ATV -nsn ejdos ‘ssul ‘deuds p-¢ ‘ssou -4OIY} 9}IPSULIO} UI BSSOF MOT “Teys ‘prfos “yor essoy doop ‘ssou -YOIY) 9}IPSULI9} UI BSsOoy MOTTeYS YOry} yidep BSSO} OILIPOU ~19}UT ‘pryos “yory} essoy doap AIOA ‘osoyjided ‘ury3 vssoy doop sosorided ‘ury} yidep BSSOJ 91VIPSUT -19}UT ‘pros “Yorys BSsO} MOTTeUS ‘osorided ‘ssou -YOIY} 91eIPSULIO}UT cV< v-SC (SOLIOS UT) o~ cE-ST ce-S'°C 8b—-0V 09<—8P vS-8P CE-87 OS-br Ov—-0t 8r—-0P epHoly ‘9U990}STO[q A[IeO 0} 9UDD0T[g II PpIw oqndoy uvorurwmogd ‘epLiol{ £9U990}SI9[q AjIVd 0} sUSDOTYY ATIvS BIUIO;ITeD ‘ogqndoy uvoruiu -0q ‘ouss0T[q Aj1e9 0} susD0IPy AyIvO Uv900O ULIPU] ‘vag poy “1U999R [Izelg ‘1U909Ry ROLY “MG ‘uBaggLeD ‘1U9909y 0} DUSD0TJY AjIvO BOLIY “M ‘[Izelg ‘epnuliog ‘ue -OqqUueyD ‘U909yY 0} SUSDOI[g o[Ppru oyloed UW1o1Sed “JUd09x% 616] ‘UeYysneA DIIUaI0Id ‘5 616] ‘UeYsNeA sisuadgjis -§ LI6l “ueysne,A jpyuapuau -¢ OSs] ‘omrey wy Splempy” ull, Duvdusiavs ‘¢ 8981 THEA DIDI]2IS ‘S (98LI ‘Japuelog 2 si[q) vasapis ‘5 (99LI ‘SeI[ed) suDipos ‘S so1ioods Mou ‘1UUud]s “§ saoualayal TeuonIppy Trem aiyqerop eyjeumnyjoD (uw) JojouleIp aITTeIOD OTTeV109 Jod vydas jo Joquinyy uonnqisiq sa1sedg “SEG SBUOA = 6 *€661 UOIOA = 8 ‘ESET Ill JOOYOS = L “PL6I [910QET = 9 +6961 [9F0QRT = ¢ :G086I 19}S0.J = p ‘2086 191804 = € ‘P66I ‘18 19 PPNG = Z {6861 PPNE = I :soUdIOJoY vausvsapig Jo sotoads 1YySIo SUTYSINSUN|SIP s1o}OeIEYO SIZO;OYdIOW— "| 2qQeL VOLUME 107, NUMBER 4 595 - . * | eH ’ * Pe! aoe AAA Fig. 3. Siderastrea glynni. (a) A closeup ofa healthy colony in the field. (b) Holotype, USNM 93956. Calicular surface, x8. synapticulae of S. g/ynni are more numer- ous and regular, forming a diagnostic mesh- work on the upper calical surface. Unlike most S. radians, calices are typically shal- low, and columellae are not solid and prom- inent. Distribution and ecology. —Siderastrea glynni is known only from Isla Uraba in the eastern Pacific, and it is extremely rare and possibly endangered. Colonies occurred clumped in a single patch in shallow reef rubble. 596 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 4. SEM photos of calices of two common Caribbean species of Siderastrea. (a) S. siderea. USNM 93957. Recent, 20-25 m depth, Limones, San Blas Archipelago, Panama. x 20. (b) S. radians. SUI 84539. Recent, <1 m depth, southeast Cayos Zapatilla, Bocas del Toro, Panama. x 20. VOLUME 107, NUMBER 4 597 Fig. 5. (a, b). SEM photos of calices of the holotype of Siderastrea glynni, USNM 93956. x20. 598 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 6. SEM photos of the (a) wall (x20) and (b) columella (x80) on the holotype of Siderastrea glynni, USNM 93956. Acknowledgments figures. Funds for SEM work were provided by the Graduate College of the University We thank S. D. Cairns, N. Knowlton, J. of Iowa. B. C. Jackson, and J. E. N. Veron for com- ments on the manuscript; J. Jara, G. Ja- come, and J. Mate for field assistance; and _Bjainville, H. M. de. 1830. Dictionnaire de sciences X. Guerra and R. A. Nessler for help with naturelles.—Zoophytes (Paris) 60:297-364. Literature Cited VOLUME 107, NUMBER 4 Bourne, G. C. 1990. Anthozoa. Pp. 1-84 in E. R. Lankester, ed., Treatise on Zoology, Part II, London. Budd, A. F. 1989. Biogeography of Neogene Carib- bean reef-corals and its implications for the an- cestry of eastern Pacific reef-corals. Memoirs of the Association of Australasian Palaeontol- ogists 8:219-230. , T. A. Stemann, & K. G. Johnson. 1994. Stratigraphic distributions of genera and species of Neogene to Recent Caribbean reef corals. — Journal of Paleontology 68:951—977. Chevalier, J. P. 1961. Recherches sur les Madrépo- raires et les formations récifales miocénes de la Méditerranée occidentale.— Mémoires de la So- ciété géologique de France 93:1—562 + pls. 1- 26. Ellis, J., & D. Solander. 1786. The natural history of many curious and common zoophytes. London, 208 pp., 63 pls. Foster, A. B. 1979. Environmental variation in a fossil scleractinian coral.—Lethaia 12:245-262. 1980a. Ecology and morphology of the Ca- ribbean Mio-Pliocene reef coral Siderastrea. — Acta Palaeontologica Polonica 25:439-450. 1980b. Environmental variation in skeletal morphology within the Caribbean reef corals Montastraea annularis and Siderastrea side- rea. — Bulletin of Marine Science 30:678-709. Holst, I., & H. M. Guzman. 1993. Lista de corales hermatipicos (Anthozoa: Scleractinia; Hydro- zoa: Milleporina) a ambos del istmo de Pana- ma.— Revista de Biologic Tropical 41:535-540. Laborel, J. 1969. Madréporaires et hydrocoralliaires récifaux des c6tes brésiliennes. — Annales del In- stitut Océanographique 47:171—229. . 1974. West African reef corals: an hypothesis on their origin.— Proceedings of the Second In- ternational Coral Reef Symposium 1:425-443. Milne Edwards, H., & J. Haime. 1850. Recherches sur les polypiers. Sixiéme mémoire. Monogra- phie des fongides.— Annales des sciences natu- relles, ser. 3, Zoologie 15:73-144. Pallas, P.S. 1766. Elenchus Zoophytorum. Den Haag, 451 pp. Scheer, G., & C. S. G. Pillai. 1983. Report on the stony corals from the Red Sea.— Zoologica 133: 1-198 + pls. 1-41. Seba, A. 1756. Locupletissimi rerum naturalium the- sauri accurata descriptio et iconibus artificio- sissimus expressio, per universam physices his- torium. Vol. 3. Amstelaedami. Vaughan, T. W. 1917. The reef-coral fauna of Carrizo Creek, Imperial County, California and its sig- 599 nificance.— U. S. Geological Survey Profession- al Paper 98T:355-395 + pls. 92-102. 1919. Fossil corals from Central America, Cuba, and Porto Rico, with an account of the American Tertiary, Pleistocene, and Recent cor- al reefs.—U. S. National Museum Bulletin 103: 189-524 + pls. 68-152. ——,, & J. W. Wells. 1943. Revision of the sub- orders, families, and genera of the Scleracti- nia. — Geological Society of America Special Pa- per 104:1-363. Veron, J. E. N. 1986. Corals of Australia and the Indo-Pacific. Angus and Robertson Publishers, Sydney, 644 pp. 1993. A biogeographic database of herma- typic corals.—Australian Institute of Marine Science Monograph Series 10:1—433. Vermill, A. E. 1865. Classification of polyps: (Extract condensed from a synopsis of the polypi of the North Pacific Exploring Expedition under Cap- tain Ringgold and Rodgers, U.S.N.).—Essex In- stitute Communications 4(9):145-152. 1868. Notes on the Radiata in the Museum of Yale College, with descriptions of new genera and species. No. 4. Notice of corals and echi- noderms collected by Prof. C. F. Hartt, at the Abrohlos Reefs, Province of Bahia, Brazil, 1867.—Connecticut Academy of Arts and Sci- ences Transactions 1:351-371. Weerdt, W. H. de, & P. W. Glynn. 1991. A new and presumably now extinct species of Millepora (Hydrozoa) in the eastern Pacific.—Zoologo- gische Mededelingen 65:267-276. Wells, J. W. 1956. Scleractinia. Pp. F328-444 in R. C. Moore, ed., Treatise on invertebrate pale- ontology, vol. F. Geological Society of America and University of Kansas Press, Lawrence, Kan- sas. . 1983. Annotated list of the scleractinian cor- als of the Galapagos. Pp. 212-291 in P. W. Glynn & G. M. Wellington, Corals and coral reefs of the Galapagos Islands. University of California Press, Berkeley, 330 pp. Yonge, C. M. 1935. Studies on the biology of Tor- tugas corals. II. Variation in the genus Sideras- trea. —Carnegie Institute of Washington, Papers from the Tortugas Laboratory 29:199-208. Department of Geology, The University of Iowa, Iowa City, Iowa 52242, U.S.A.; Smithsonian Tropical Research Institute, Apartado 2072, Balboa, Republic of Pan- ama. PROC. BIOL. SOC. WASH. 107(4), 1994, pp. 600-608 RESURRECTION OF GLYPHOHESIONE FRIEDRICH, 1950, WITH REDESCRIPTION OF G. KLATTI FRIEDRICH, 1950 AND DESCRIPTION OF G. LONGOCIRRATA (POLYCHAETA: HESIONIDAE) Frank Licher Abstract.—The formerly monotypic genus Glyphohesione Friedrich, 1950 is removed from synonymy with Syne/mis Chamberlin, 1919 and emended. The type species, G. klatti Friedrich, 1950, from northern European waters, is redescribed, and G. /ongocirrata, a new species from the east coast of North America, is described. G/yphohesione is transferred from the Pilargidae to the Hesionidae, where it was originally placed, as the pilargid stem-species might have evolved from a hesionid species by progenesis. Friedrich (1950) described the monotypic Glyphohesione klatti from Helgoland and recognized it as a hesionid. Eliason (1962a, 1962b), who reported the species from the Skagerrak and Oresund, considered it to be a pilargid belonging to the genus Ancistro- syllis McIntosh, 1879. Pettibone (1966) transferred it to the pilargid genus Synelmis Chamberlin, 1919. Comparison of brain morphology of different pilargid genera caused Fitzhugh & Wolf (1990) to doubt that specimens identified as Synelmis klatti in the collection of the U.S. National Mu- seum, Washington, belong to this genus, and they suggested resurrecting the older taxon Glyphohesione for them. In a phylogenetic analysis of the Pilargidae (note: the correct spelling is Pilargidae, not Pilargiidae (see: International Commission on Zoological Nomenclature (1985): International Code of Zoological Nomenclature, art. 35 d (11))), Licher & Westheide (1994) argued that the species of this family might form a mono- phyletic group within the Hesionidae. A subfamily Pilarginae beside the “‘Hesioni- nae” Hartmann-Schroder, 1971 and “Mi- crophthalminae”’ Hartmann-Schroder, 1971 should not be erected until a comprehensive revision of the Hesionidae including pilar- gids has been made of the “‘true’’ hesionid taxa and is beyond the scope of the present study. The ““Hesioninae”’ as well as the for- mer “‘Hesionidae”’ (““Hesioninae” + “‘Mi- crophthalminae’’) are very likely paraphy- letic (Licher & Westheide 1994). The ““Microphthalminae” represents a polyphy- letic group (Westheide 1977). Reinvestiga- tion of material of different species identi- fied as Synelmis klatti in the course of the analysis of Licher & Westheide 1994 in- duced them to reestablish the old generic name. The present paper emends the diagnosis of Glyphohesione, redescribes the European G. klatti and describes the eastern North American Glyphohesione longocirrata, new species. Materials and Methods For light microscopical preparations the fixed specimens (stored in 70% ethanol) were transferred into glycerine. Observations, drawings, and measurements were made by means of a LEITZ Diaplan microscope with interference-contrast optics and a camera lucida. For SEM investigations one speci- men was dehydrated and critical-point dried with carbon dioxide. After sputtering with gold, it was analyzed with a JEOL JSM 820. VOLUME 107, NUMBER 4 Material examined originates from the following museums: Gothenburg Natural History Museum (GNM); Senckenberg Mu- seum, Frankfurt (SMF); U.S. National Mu- seum of Natural History, Smithsonian In- stitution, Washington, D.C. (USNM); Zoological Museum, University of Copen- hagen (ZMUC). Glyphohesione Friedrich, 1950, emended Type species. —Glyphohesione klatti Friedrich, 1950, by monotypy and original designation. Additional material examined. — The ho- lotype of the type species of Synelmis Chamberlin, 1919, S. simplex Chamberlin, 1919 (USNM 19480, type) and some non- type specimens of S. al/bini (Langerhans, 1881) (type locality the Canary Islands, type material lost) from the Galapagos Islands (W. Westheide coll., Osnabriick) have been examined. Diagnosis. —Hesionidae with body dor- soventrally flattened; some anterior seg- ments distinctly wider, appearing some- what inflated. Integument smooth, without papillae. Prostomium bilobed anteriorly, with two palps consisting of palpophores fused totally with prostomium and elon- gated palpostyles. Three slender antennae; lateral antennae located at anterior prosto- mial margin, close to palps; median antenna positioned at posterior margin. Pharynx un- armed. Peristomium achaetous, with two pairs of slender tentacular cirri. Parapodia biramous. Notopodia each with elongated dorsal cirrus, one notoacicula, and one stout emergent spine-like notochaeta, the latter in median and posterior segments only. Neu- ropodia well developed, each with slender ventral cirrus, one neuroacicula and simple chaetae only. Pygidium with two elongated anal cirri. Remarks. —Eliason (1962a, 1962b) placed the monotypic Glyphohesione in Ancistro- syllis McIntosh, 1879, based on similarities with Synelmis albini (Langerhans, 1881) 601 (=Ancistrosyllis albini). However, Ancistro- syllis possesses a hook-shaped notochaeta, not a straight one, which is characteristic for Synelmis. Pettibone (1966) transferred both A. albini and A. klatti to Synelmis, assuming S. k/atti to be a juvenile of S. albini. Pearson (1970), Hartmann-Schroder (1971), and Katzmann et al. (1974) consid- ered S. klatti to be a member of Synelmis. Fitzhugh & Wolf (1990) investigated Amer- ican material identified as Synelmis klatti and suggested resurrecting the original ge- neric name. Glyphohesione Friedrich, 1950 clearly dif- fers from Synelmis Chamberlin, 1919 in lacking the two emergent neuropodial spines which are apomorphic for Synelmis (Fitz- hugh & Wolf 1990, Licher & Westheide 1994). In Glyphohesione, dorsal cirri are longer than ventral cirri, and the dorsal cirri of the first chaetiger are longer than those of the following ones, whereas in Synelmis dorsal and ventral cirri of all chaetigers are subequal. In addition, this taxon is known to possess nuchal organs, which were not found in Synelmis, and the brain is similar to that of Sigambra Miler, 1858 (Fitzhugh & Wolf 1990). Glyphohesione clearly differs from Sigambra in having notopodial spines and in lacking hook-shaped notochaetae. Discussion. —Licher & Westheide (1994) conclude that there are good indications that the pilargid stem-species might have evolved by progenesis from a juvenile stage of a large- bodied hesionid species. This induced them to include the family Pilargidae in the He- sionidae, a view generally adopted by earlier authors (e.g., Ehlers 1908, Fauvel 1923, Au- gener 1927, Monro 1933, Treadwell 1941). According to Licher & Westheide (1994), within the pilargids Glyphohesione is the taxon with the highest number of plesio- morphic characters shared with juvenile he- sionids, e.g., (1) possession of elongated pal- postyles, (2) lateral antennae located at the anterior prostomial margin, and (3) prosto- mial, peristomial, parapodial and pygidial appendages elongated and well developed. 602 Fig. 1. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Glyphohesione klatti Friedrich, 1950 (ZMUC POL-178). A. Anterior and median part of the body, dorsal view. B. Anterior end, dorsal view. C-F. Parapodium 21 (ZMUC POL-178-A): C. Parapodium, posterior view, half of neuropodial fascicle omitted. D. Emergent spine-like notochaetae. E. Ventralmost neurochaeta. F. Dorsalmost neurochaeta. Scales: A = 500 um; B = 250 um; C = 100 wm; D-F = 25 um. Glyphohesione is monophyletic and the adelphotaxon of the stem-species of all oth- er pilargid genera (Licher & Westheide 1994). Genus Glyphohesione Friedrich, 1950, resurrected Glyphohesione klatti Friedrich, 1950 Figs. 1-2 Glyphohesione klatti Friedrich, 1950:171- 173, figs. 1-2. Ancistrosyllis klatti.—Eliason, 1962a:241; 1962b:29-32, fig. 3. Synelmis klatti. —Pettibone, 1966:190- 191.—Pearson, 1970:74-75, fig. 2b, c.— Hartmann-Schroder, 1971:144-145, fig. 49.—Katzmann et al., 1974:27—28. [Not Synelmis klatti of Wolf 1984; not Fitz- hugh & Wolf 1990 (both = G. longocir- rata, new species)]. Material examined. —Kattegatt: Off Ska- gen, Denmark, 58°01'N, 10°52’E, 190 m, fine mud, M. E. Petersen coll., 22 Sep 1968 (ZMUC POL-178, 5 incompl. specimens; ZMUC POL-178-B, 1 incompl. SEM prep- aration). Laholmsbukten, off Laholm, Swe- den, St. 150/417, 21 m, stiff clay and some sand, “Akka,” L. A. Jagerskiold coll., 17 Jun 1933 (GNM 11347, 1 incompl. speci- men). —Skagerrrak: Saltkallefjord, off Gull- maren, Sweden, 56 m, P. Bagge coll., 24 VOLUME 107, NUMBER 4 Fig. 2. Glyphohesione klatti Friedrich, 1950 (ZMUC POL-178-B). A. Anterior end, lateral view. B. Parapodia of median region, right side, dorsal view. Scales: A-B = 100 ym. Nov 1964 (GNM 12839, 1 incompl. spec- imen).— North Sea: Raunefjord, off Bergen, Norway, 245 m, sandy loam, Fosshagen coll., 05 Oct 1962 (GNM 12949, 2 incompl. specimens). German Bight, 54°40'N, 06°00’E, 43 m, silt and clay, M. Tiirkay coll., 24 May 1987 (SMF 4432, 1 incompl. spec- imen).—English Channel: Survey Sta. M 16T, 51°24.6'N, 08°05'W, 112 m, J. P. Hartley coll., Aug 1975 (USNM 58901, 2 incompl. specimens). Type locality.—Tiefe Rinne off Helgo- land, North Sea. Type material lost (H. Friedrich, pers. comm.). Description. — Body robust, dorsoventral- ly flattened; anterior segments inflated, much wider than long, median segments about as wide as long, posterior segments longer than wide (Fig. 1A). Largest incomplete speci- men (USNM 58901) with 40 segments and a length of 9.6 mm, 1.1 mm wide with par- apodia (chaetiger 1), 0.8 mm wide without parapodia (chaetiger 5). (Up to 13 mm long, 0.5 mm wide for 71 segments according to Eliason 1962b:29). Color in ethanol brown. Living specimens with yellowish gut and transparent body with yellowish pigment in anterior region on posterior part of prosto- mium (M. E. Petersen, pers. comm.; see also fig. 3a, fin Eliason 1962b). Prostomium slightly wider than long, di- vided anteriorly by a wide furrow into two lobes, lateral margins slightly concave (Figs. 1B, 2A). Palps with palpophores fused to prostomium and free elongate palpostyles, somewhat shorter than the lateral antennae, inserted anteroventrally. Three slender an- tennae, with slightly inflated bases. Lateral antennae slender, located at anterior pro- stomial margin; median antenna very slen- der, about 1.5—2 times as long as lateral ones, positioned at posterior prostomial margin. Eyes lacking. Nuchal organs not visible. Peristomium almost as long as each of following anterior segments, dorsally fused with prostomium and encircling posterior half of latter. Two pairs subequal, slender tentacular cirri of similar shape and size as median antenna (Figs. 1B, 2A). Specimens in ethanol usually with two indistinctly de- limited, brown-pigmented bulging areas on posterior part of peristomium. Parapodia biramous, usually distinctly set apart from trunk except for some in inflated anterior region. Notopodia each with slen- der dorsal cirrus, extending slightly beyond tip of neuropodial lobe (Figs. 1C, 2B). Lon- gest dorsal cirri (ca. 200 wm) occurring in first chaetiger. Following cirri much shorter, not longer than tentacular cirri. One thin notoacicula. Parapodia of median and pos- terior segments with one stout emergent spine-like notochaeta (first present on chae- tigers 5-8) (Fig. 1C, D). 604 Neuropodia conical, each with a longer, triangular prechaetal lobe; a fan-shaped bundle of up to ca. 25 simple chaetae (Figs. 1C, 2B), decreasing in number posteriorly; and a shorter, distally truncate postchaetal lobe. Chaetae of different lengths, shorter ones stouter, all finely serrated, distally pointed with smooth, entire tips (Fig. 1E, F); serration becoming shorter distally and dificult to recognize. One neuroacicula present. Ventral cirri similar to dorsal ones but shorter, not extending beyond neuro- podial lobe. Posterior end lacking in all of the avail- able specimens, but pygidium with two ven- tral filiform anal cirri according to Eliason (1962b:fig. 33). Distribution. —Glyphohesione klatti is known only from European waters: Ore- sund and Kattegat (Eliason 1962a, this pa- per); Skagerrak (Eliason 1962b, Bagge 1969, this paper); Tiefe Rinne off Helgoland, North Sea (Friedrich 1950); German Bight, North Sea (this paper); Loch Linnhe, Scotland (Pearson 1970); English Channel (this pa- per); and the Catalonian coast of the Med- iterranean Sea (Katzmann et al. 1974). In northern Europe it has been taken at depths of 20-680 m and in the Mediterranean at depths of 10-185 m. Glyphohesione longocirrata, new species Fig. 3 Synelmis klatti. —Wolf, 1984:29-31 to 29- 35, figs. 29-29 and 29-30 (USNM 86983, 86984, 86985, 86986, 86987); Fitzhugh & Wolf, 1990:1-16 (USNM 86986). Material examined. —Gulf of Mexico: off Florida: SOFLA Sta. 20A, 25°17.20'N, 82°09.44'W, 22 m, coarse sand, Apr 1981 (USNM 86985, 1 compl. specimen, holo- type); SOFLA Sta. 20C, 25°17.20'N, 82°09.44'’W, 22 m, coarse sand, Nov 1980 (USNM 86986, 2 specimens, paratypes); SOFLA Sta. 20E, 25°17.20'N, 82°09.44'W, 22 m, Apr 1981 (USNM 86987, | speci- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON men, paratype); MAFLA Sta. V-2531, 29°47.59'N, 86°09.29'W, 45 m, coarse sand, Aug 1977 (USNM 86983, | specimen, para- type).— Northwest North Atlantic Ocean: Gulf of Maine: Massachusetts, off Cape Cod: NEEB Sta. 41, 41°37.30'N, 69°15.42’W, 164 m, 27 Feb 1977 (USNM 91310, 5 speci- mens); NEEB Sta. 41, 41°37.34'N, 69°15.46’'W, 165 m, 27 Feb 1977 (USNM 91311, 1 specimen); NEEB Sta. 41, 41°37.21'N, 69°14.54'W, 172 m, 27 Feb 1977 (USNM 91312, 1 specimen); NEEB Sta. 41, 41°37.00'N, 69°14.59'W, 165 m, 27 Feb 1977 (USNM 91313, 1 specimen); NEEB Sta. 41, 41°37.36'N, 69°15.49'W, 178 m, 17 May 1977 (USNM 91320, 1 speci- men); NEEB Sta. 41, 41°36.58'N, 69°15.35'W, 175 m, 17 May 1977 (USNM 91321, 1 specimen); NEEB Sta. 42, 41°50.26'N, 69°29.30'W, 185 m, 26 Feb 1977 (USNM 91314, 1 specimen); NEEB Sta. 42, 41°50.10'N, 69°29.10'W, 185 m, 26 Feb 1977 (USNM 91315, 1 specimen); NEEB Sta. 42, 41°50.06’N, 69°29.09’W, 185 m, 26 Feb 1977 (USNM 91316, 1 speci- men); NEEB Sta. 42, 41°49.53’N, 69°28.54'W, 191 m, 16 May 1977 (USNM 91322, 1 specimen); NEEB Sta. 42, 41°50.35'N, 69°29.28’W, 179 m, 16 May 1977 (USNM 91323, 2 specimens).— Georges Bank: Northern Slope: NEEB Sta. 35, 42°13.08'N, 67°34.20'W, 242 m, 19 May 1977 (USNM 91317, 1 specimen); NEEB Sta. 35, 42°13.05'N, 67°33.49'W, 243 m, 19 May 1977 (USNM 91318, 1 specimen); NEEB Sta. 35, 42°13.19'N, 67°34.36'W, 239 m, 19 May 1977 (USNM 91319, 1 speci- men); Southern Slope: NEEB Sta. 16, 40°42.30'N, 67°34.26'W, 87 m, 19 Feb 1977 (USNM 91309, 1 specimen); Nantucket Shoals: NEEB Sta. 3, 40°39.38'N, 69°27.23'W, 56 m, 15 Feb 1977 (USNM 91308, 1 specimen). Type locality. —Gulf of Mexico, off south- ern Florida, 22 m, coarse sand. Description.—Body dorsoventrally flat- tened, with 2—4 inflated anterior segments (Fig. 3A). Largest complete specimen ex- VOLUME 107, NUMBER 4 { A Q NZ ~~ | — —_— SSS Sree = oO 605 sf, an — Aa j x eae, ey = SS af ee ee ee Oy See ss = BEE WAP sig cv aL ee = = ~ b= = 7 Panis Spek { ———— U4 G & — = Zo ? C ~ Pe F E Fig. 3. Glyphohesione longocirrata, new species (USNM 86985, holotype). A. Anterior and median part of the body, dorsal view. B. Anterior end, dorsal view. C. Parapodium of median region, posterior view. D. Posterior end, lateral view. E-G. Posterior chaetiger: E. Emergent spine-like notochaetae. F. Ventralmost neurochaeta. G. Dorsalmost neurochaeta. Scales: A = 250 um; B = 100 wm; C—D = 50 um; E-G = 25 um. amined (USNM 86985, holotype) with 43 segments, 5.1 mm long, up to 0.5 mm wide including parapodia (chaetiger 4), 0.2-0.25 mm wide without parapodia. (Largest spec- imen examined by Wolf (1984) incomplete, with 53 segments, 12.0 mm long and 0.5 mm wide.) Color in ethanol light greyish yellow, nearly transparent. Prostomium slightly wider than long, with concave lateral margins (Fig. 3B). Palps 606 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 1.—Characters of Glyphohesione klatti Friedrich, 1950 and G. longocirrata, new species. Characters Maximum length x width, no. of chaetigers plete) Habitus; color Robust; brown No. anterior inflated seg- ca. 8 ments Tentacular and dorsal cirri of anterior segments Notochaetae: first appearance of spines Neurochaetae: structure, number per fascicle Distribution Chaetigers 5-8 681 m consisting of two filiform palpostyles only, inserted anterolaterally at ventral side. Three filiform antennae; lateral ones near anterior prostomial margin, somewhat longer than palpostyles; median antenna longer than lateral ones, positioned at posterior prosto- mial margin. Eyes lacking. Nuchal organs not visible. Peristomium shorter than following seg- ments. Anterior peristomial margin slightly overlapped by prostomium. Two pairs sub- equal, filiform tentacular cirri, of similar shape as median antenna (Fig. 3B). Some specimens with two pigmented areas on posterior part of the peristomium (see also fig. 29-30a in Wolf 1984). Parapodia biramous, usually distinctly set apart from trunk. Parapodia of second chae- tiger usually smaller than those of first one. Notopodia each with filiform dorsal cirrus, extending far beyond tip of neuropodium (Fig. 3C). Dorsal cirri of anterior segments nearly as long as tentacular cirri. One thin and transparent notoacicula. Parapodia of median and posterior segments with one long stout emergent spine-like notochaeta (first present on chaetigers 10-15), nearly as long as shortest neurochaetae, decreasing in length posteriorly, distinctly bent in pre- pygidial segment (Fig. 3D, E). Neuropodial lobe conical, with bundle of simple serrated and distally minutely bifid Glyphohesione klatti Friedrich, 1950 13 x 0.5 mm, 71 chaetiger (com- At most one-fourth of body width Finely serrated with entire, curved tips; up to 25 per fascicle Northern and southern Europe, 43- G. longocirrata, new species 12+ x 0.5 mm, 53 chaetigers (in- complete) Delicate; light greyish yellow 2-4 As long as body width Chaetigers 10-15 Finely serrated with minutely bifid tips; up to 14 per fascicle North American east coast and Gulf of Mexico, 22-243 m chaetae, 8-14 in anterior segment, 6-9 in posterior ones (Fig. 3C, F, G). One neu- roacicula present. Ventral cirri similar to dorsal ones, extending slightly beyond tip of neuropodial lobe. Pygidium rounded, with two long filiform anal cirri, about twice as long as median antenna (Fig. 3D). Distribution. — Glyphohesione longocir- rata, new species, is widely distributed off the east coast of North America: from the Gulf of Maine (off Massachusetts) to the Gulf of Mexico (off Florida and Alabama), at depths of 20—240 m, on bottoms of coarse to medium-fine sand, silty sand and clayey silt. Remarks. —This species is assigned to the genus Glyphohesione because of the pres- ence of simple chaetae, a straight spine-like notochaeta, elongated palpostyles, and the position of the lateral and median antennae. Glyphohesione longocirrata, new species, differs from G. klatti Friedrich, 1950 in pos- sessing longer, more developed tentacular and dorsal cirri (Table 1). Dorsal cirri in anterior region in G. /ongocirrata are nearly as long as the body width; in G. k/atti they are a quarter of the body width at most. In G. longocirrata the neurochaetae per fasci- cle are few and distally bifid (see Wolf 1984). Furthermore, specimens from North Amer- ica have a more delicate body shape. VOLUME 107, NUMBER 4 Etymology.—This species is named for its elongated tentacular and dorsal cirri. Acknowledgments I am indebted to Prof. Dr. K. Fauchald, Smithsonian Institution, Washington, D. C. (USNM), for access to material and for per- mission to describe it. 1am pleased to thank Dr. D. Eibye-Jacobsen and Dr. M. E. Pe- tersen, Zoological Museum, University of Copenhagen (ZMUC); Dr. D. Fiege, Senk- kenberg Museum, Frankfurt (SMF); and Prof. Dr. L. Orrhage, Gothenburg Natural History Museum (GNM) for the kind and courteous loan of specimens. Special thanks are due to Dr. D. Rode and Dipl.-Ing. H.- J. Hemschemeier, KM-kabelmetal, Osna- brick for providing use of their SEM. Fur- thermore, I thank Prof. Dr. W. Westheide and Dr. G. Purschke, Universitat Osna- briick, and especially Dr. M. E. Petersen (ZMUC) for their critical reviews of a pre- liminary draft of this paper. Literature Cited Augener, H. 1927. Polychaeten von Sidost- und Sid- Australien. — Videnskabelige Meddelelser fra Dansk naturhistorisk Forening i Kobenhayn 83: 71-275. Bagge, P. 1969. Effects of pollution on estuarine eco- systems. I. Effects of effluents from wood-pro- cessing industries on the hydrography, bottom nd fauna of Saltkallefjord (W. Sweden). II. The succession of the bottom fauna communities in polluted estuarine habitats in the Baltic-Skag- erak region.— Merentutkimuslaitoksen Julkaisu Havsforskningsinstitutets Skrift 228:3-118. Chamberlin, R. V. 1919. The Annelida Polychaeta. — Memoirs of the Museum of Comparative Zo- ology at Harvard College 49:1-514. Ehlers, E. 1908. Die bodensdssigen Anneliden aus den Sammlungen der Deutschen Tiefsee-Expe- dition. — Wissenschaftliche Ergebnisse der Deutschen Tiefsee-Expedition 1897-1899, 16(1): 1-168. Eliason, A. 1962a. Die Polychaeten der Skagerak- Expedition 1933.—Zoologiska Bidrag fran Uppsala 33:207-293. 1962b. Undersékningar 6ver Oresund, 41: Weitere Untersuchungen tiber die Polychaeten- 607 fauna des Oresunds.—Lunds Universitets Arsskrift, n.f. 58(9):1-97. Fauvel, P. 1923. Polychétes Errantes. Faune de France 571-488. Fitzhugh, K., & P.S. Wolf. 1990. Gross morphology of the brain of pilargid polychaetes: Taxonomic and systematic implication.—American Muse- um Novitates 2992:1-16. Friedrich, H. 1950. Zwei neue Bestandteile in der Fauna der Nordsee.—Neue Ergebnisse und Probleme in der Zoologie, Festschrift Klatt, Zoologischer Anzeiger (Erganzungsband) 145: 171-177. Hartmann-Schroder, G. 1971. Annelida, Borsten- wurmer, Polychaeta.—Die Tierwelt Deutsch- lands 58:1-594. International Commission on Zoological Nomencla- ture. 1985. International Code of Zoological Nomenclature adopted by the 20th General As- sembly of the International Union of Biological Sciences. International Trust for Zoological No- menclature, London, 338 pp. Katzmann, W., L. Laubier, & J. Ramos. 1974. Pi- largidae (Annélides Polychétes errantes) de Mé- diterranée.— Bulletin de l'Institut océanogra- phique 71(1428):1—40. Langerhans, P. 1881. Uber einige canarische Anne- liden.— Deutsche Akademie der Naturforscher Nova Acta 42:93-124. Licher, F., & W. Westheide. 1994. The phylogenetic position of the Pilargidae with a cladistic anal- ysis of the taxon—facts and ideas. In J. C. Dau- vin, L. Laubier, & D. J. Reish, eds., Actes de la 4éme Conférence international des Poly- chétes. —Meémoires du Muséum National d’His- toire Naturelle (A) 162:223-236. McIntosh, W. C. 1879. On the Annelida obtained during the Cruise of H.M.S. ‘Valorous’ to Davis Strait in 1875.—Transactions of the Linnean Society 2(1):499-511. Monro, C.C. A. 1933. Onanew species of Polychaeta of the genus Pilargis from Friday Harbor, Wash- ington.— Annals and Magazine of Natural His- tory 10(11):673-675. Miiller, F. 1858. Einiges tiber die Annelidenfauna der Insel Santa Catharina an der brasilianischen Kiiste. — Archiv fiir Naturgeschichte 24:21 1-220. Pearson, T.H. 1970. Litocorsa stremma, a new genus and species of pilargid (Polychaeta: Annelida) from the west coast of Scotland with notes on two other pilargid species.— Journal of Natural History 4:69-77. Pettibone, M. H. 1966. Revision of the Pilargidae (Annelida: Polychaeta), including descriptions of new species, and redescriptions of the pelagic Podarmus ploa Chamberlin (Polynoidae).— Proceedings of the U.S. National Museum 118(3525):155-208. 608 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Treadwell, A. L. 1941. Polychaetous annelids from the New England region, Porto Rico and Bra- zil.—American Museum Novitates 1138:1-4. Westheide, W. 1977. Phylogenetic systematics of the genus Microphthalmus (Hesionidae) together with a description of M. hartmanae nov. sp. Pp. 103-113 in D. J. Reish & K. Fauchald, eds., Essays on polychaetous annelids in memory of Dr. Olga Hartman.—Allan Hancock Founda- tion Special Paper, Los Angeles. Wolf, P.S. 1984. Chapter 29. Family Pilargidae Saint- Joseph, 1899. Pp. 29-1 to 29-41 in J. M. Uebe- lacker & P. G. Johnson, eds., Taxonomic guide to the polychaetes of the northern Gulf of Mex- ico, 7(4). Final report to the Minerals Manage- ment Service, contract 14-12-001-29091. Barry A. Vittor & Ass., Mobile, Alabama. Universitat Osnabriick, Fachbereich 5, Spezielle Zoologie, D-49069 Osnabriick, Germany. PROC. BIOL. SOC. WASH. 107(4), 1994, pp. 609-614 ADDITIONAL RECORDS OF POLYNOID POLYCHAETES FROM THE JUAN DE FUCA RIDGE Marian H. Pettibone Abstract. — Additional specimens of two polynoid polychaetes, Bathycatalina filamentosa (Moore) (Macellicephalinae), and Harmothoe macnabi Pettibone (Harmothoniae), were collected from a new vent site on the Juan de Fuca Ridge. Based on these materials, both the subfamily Macellicephalinae and genus Bathycatalina are emended, the former to include Bathylevensteinia bi- cornis (Levenstein) and Gesiella jameensis (Hartmann-Schroder), and the de- scriptions of B. filamentosa and H. macnabi supplemented. A grab-sample of Ridgia tube-worms was taken from an isolated black smoker during the Atlantic/Alvin cruises to the Juan de Fuca Ridge in October 1993. Polynoid poly- chaetes obtained from the sample were sent to me for identification, and found to be Bathycatalina filamentosa (Moore) (Macel- licephalinae), and Harmothoe macnabi Pet- tibone (Harmothoniae). Descriptions of both species are supplemented based on these ad- ditional materials. The genus Bathycatalina Pettibone is emended, and the Subfamily Macellicephalinae is also emended to in- clude Gesiella jameensis (Hartmann- Schroder), and Bathylevensteinia bicornis (Levenstein). Specimens are deposited in the Depart- ment of Invertebrate Zoology, National Museum of Natural History, Smithsonian Institution (USNM). Family Polynoidae Kinberg, 1856 Subfamily Macellicephalinae Hartmann- Schroder, 1971, emended Pettibone, 1976. Additions and emendations were added to the Subfamily Macellicephalinae Hart- mann-Schroder, 1971, by Pettibone (1976: 6; 1985a:129; 1985b:740; 1989:159; & 1993: 679). Two new genera and species that were placed in the Subfamily Harmothoinae Horst, 1917, by Pettibone (1976:60) are herein included in the Subfamily Macelli- cephalinae, based on previous incorrect in- terpretations of the structures on the ante- rior lobes of the prostomia, as lateral antennae, rather than as anterior rounded- filiform extensions and not distinct lateral antennae. They include Bathylevensteinia Pettibone, 1976, with B. bicornis (Leven- stein, 1962) and Gesiella Pettibone, 1976, with G. jameensis (Hartmann-Schréder, 1974). Bathylevensteinia bicornis (Levenstein, 1962) Macellicephala bicornis Levenstein, 1962: 1143, fig. 1.1), in Macellicephalinae. Bathylevensteinia bicornis. —Pettibone, 1976:62, fig. 35a—e, in Harmothoinae. Remarks. — Bathylevensteinia bicornis was incorrectly place in Harmothoinae, due to an incorrect interpretation of the so-called bifurcate frontal horns on the prostomium by Levenstein. Pettibone referred to the bi- lobed prostomium with subtriangular fron- tal horns, and, more medially, lateral an- tennae with cylindrical ceratophores with styles missing. However, the medial pro- cesses are on the same level as the lateral frontal horns, not ventral, and terminal styles are absent, thus not distinct lateral antennae. 610 Gesiella jameensis (Hartmann-Schroder, 1974) Macellicephala (Macellicephala) jameensis Hartmann-Schroder, 1974:76, figs. 1-8, in Macellicephalinae. Gesiella jameensis. — Pettibone, 1976:64, fig. 36a-j, in Harmothoinae.—Muir, 1982: 156, in Gesiellinae Muir, 1982. Remarks. —The small spherical lobes with distal filaments on the prostomium are not to be considered as distinct lateral antennae, as indicated by Pettibone (1976:64). The presence of unique filamentous sensory or- gans on the cirrophores of the dorsal cirri, the basis for Gesiellinae by Muir (1982) does not seem to merit a separate Subfamily. Genus Bathycatalina Pettibone, 1976, emended Type species.—Polynoe (?) filamentosa Moore, 1910, by original designation and monotypy. Type locality: Southern California, off Santa Catalina Island, in 61 1- 1097 m. Remarks. —Based on an additional spec- imen from Juan de Fuca Ridge, referred herein to B. filamentosa, the genus is emended as follows: Body with 24 seg- ments, last three small, with 11 pairs of ely- trophores (not 12 pairs), on segments 2, 4, 5, 7, 9, 11, 13, 15, 17, 19, 21. Bilobed pro- stomium with rounded frontal processes and terminal filaments (not distinct lateral an- tennae, damaged on holotype); palps very long (missing on holotype). Pharynx with nine pairs of papillae (damaged on holo- type). Bathycatalina filamentosa (Moore, 1910) Fig. 1 Polynoe (?) filamentosa Moore, 1910:366, pl. 31:figs. 52-56. Bathycatalina filamentosa. —Pettibone, 1976:38, fig. 23a—e. Material.—Juan de Fuca Ridge, 46°09.3'N, 129°48.4’W, 2059 m, Alvin Dive PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 2681 in 273°C Beard Chimney, 24 Oct 1993, C. van Dover, collector, from V. Tunni- cliffe, 1 specimen (USNM 169153). Description. —Length 15 mm, width with setae 8 mm, segments 24, last three very small. Elytra (all missing except very small elytra on segment 21) and bulbous elytro- phores 11 pairs, on segments 2, 4, 5, 7, 9, 11, 13, 15, 17, 19, 21. Dorsal cirri on non- elytrigerous segments; cirrophores long, cy- lindrical, with long styles; dorsal tubercles elongate, forming digitiform ciliated bra- chial-like processes; some long clavate sen- sory papillae on dorsal cirrophores (Fig. 1A, B, F). Prostomium deeply bilobed, rounded an- terior lobes, with subconical processes and terminal filaments; ceratophore of median antenna in anterior notch of prostomium, large, cylindrical, style missing; palps very long; without eyes; tentaculophores of seg- ment | lateral to prostomium, with few no- tosetae on inner sides and two pairs of ten- tacular cirri (missing) (Fig. 1A). Segment 2 with first pair of elytrophores, biramous parapodia, and long ventral buccal cirri, longer than following ventral cirri (Fig. 1A). Biramous parapodia with both notopodial and neuropodial rami subequal in size and length, with projecting acicular processes (Fig. 1C). Notosetae numerous, stout, form- ing radiating bundles, short and longer, some as long as neurosetae, with spinous rows and blunt bare tips (Fig. 1C, D). Neurosetae very numerous, forming dense brushlike bun- dles, thin, transparent, flattened distally, paddle-like, with serrated margins (Fig. 1C, E). Ventral cirri short, tapered (Fig. 1C). Posterior end with pygidium rounded, en- closed in small posterior parapodia (seg- ments 22-24); cirrophores of dorsal cirri with clavate sensory papillae (Fig. 1B). Long extended pharynx encircled with nine pairs of dorsal and ventral papillae and two pairs of inner jaws (Fig. 1A). Groups of large yolky eggs attached to ventral surfaces of neuro- podia on some posterior segments. Distribution.—Northeastern Pacific, off Southern California (Santa Catalina Island), VOLUME 107, NUMBER 4 611 py )-ODDO Fig. 1. Bathycatalina filamentosa (USNM 169153): A, Dorsal view of anterior end, with pharynx fully extended; styles of median antenna, dorsal and ventral tentacular cirri, elytra of segment 2, and left dorsal cirrus of segment 3 missing; B, Dorsal view of posterior end (segments 21-24), dorsal cirri missing, long clavate sensory papillae on cirrophores; C, Left elytrigerous parapodium of segment 5, anterior view, acicula dotted, elytron missing; D, Short and distal tip of long notosetae from same; E, Distal tip of flattened neuroseta from same; F, Right cirrigerous notopodium, anterior view, showing cirrophore of dorsal cirrus with sensory clavate papilla (style missing) and ciliated branchial process on dorsal tubercle. Scales = 0.5 mm for C, F; 0.1 mm for D, E; A, B, not to scale. 612 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. Harmothoe macnabi (USNM 169154): A, Dorsal view of anterior end, pharynx partially extended; B, Right first elytron from segment 2, with detail of microtubercles, surface and border papillae; C, Right elytrigerous parapodium, anterior view, acicula dotted; D, Right cirrigerous parapodium, posterior view; E, Short and distal end of long notosetae from same; F, Lower and middle neurosetae from same, with detail of some tips. Scales = 0.5 mm for B—D; 0.1 mm for E, F; A, not to scale. in 611-1097 m, and Juan de Fuca Ridge, rigerous segments. G. jameensis has fewer off British Columbia, in 2059 m. segments (18-19) and elytrophores (nine Remarks. —Bathycatalina filamentosa pairs) and the notopodia are much shorter shows affinities to Gesiella jameensis. Both than the neuropodia. Ciliated cirriform species have the unique clavate sensory pa- _ _ branchial structures on the dorsal tubercles pillae on the dorsal cirrophores of the cir- of B. filamentosa are also found on Bathy- VOLUME 107, NUMBER 4 fauvelia affinis (Fauvel) and Bathybahamas charleneae Pettibone. (See Key to the three species in Pettibone, 1985a;141). Dorsal tu- bercles are indistinct in G. jameensis. Subfamily Harmothoinae Horst, 1917 Genus Harmothoe Kinberg, 1856 Harmothoe macnabi Pettibone, 1985 Fig. 2 Harmothoe macnabi Pettibone, 1985b:749, figs. 6, 7. Material.—Juan de Fuca Ridge, 46°09.3’N, 129°48.4'W, 2059 m, Alvin Dive 2681 in 273° Beard Chimney, 24 Oct 1993, C. van Dover, collector, from V. Tunni- cliffe, 2 specimens (USNM 169154). Remarks. —The two specimens agree for the most part with the holotype from the Galapagos Rift in 2482 meters. Description. — Complete specimen 12 mm long, 8 mm wide with setae, and 30 seg- ments, last three very small; incomplete specimen 10+ mm long, 9 mm wide, and 15+ segments; (holotype much larger: 33 mm long, 14 mm wide with setae, and 31 segments, last three small). Body showing brownish pigmentation: on distal tips of buccal and ventral cirri, along lateral sides of ventral nerve cord, and dark pharynx showing through body wall. Elytra and bulbous elytrophores 14 pairs, on segments 2, 4, 5, 7, alternate segments to 23, 26, 29. Elytra missing, except for first right elytron on segment 2 and very small elytra on segments 19 and 23; remaining large elytron round and covered with con- ical microtubercles and scattered long sur- face papillae and short border papillae (Fig. 2B). Dorsal cirri and non-elytrigerous seg- ments, with short cylindrical cirrophores and long papillate styles extending beyond tips of setae; dorsal tubercles nodular (Fig. 2D). Bilobed prostomium wider than long, with subtriangular cephalic peaks; median an- tenna with large ceratophore in anterior notch of prostomium; lateral antennae with distinct ceratophores inserted ventrally, 613 styles papillate, about length of prostomi- um; without eyes; stout palps slightly longer than median antenna; tentaculophores (seg- ment 1) lateral to prostomium, each with three or four setae on inner side and papil- late dorsal and ventral tentacular cirri, slightly shorter than median antenna (Fig. 2A). Segment 2 with first pair of bulbous elytrophores, biramous parapodia, and long ventral papillate buccal cirri, much longer than following ventral cirri (Fig. 2A). Biramous parapodia with notopodia rounded basally, with projecting acicular processes on lower sides, about as long as neuropodia; neuropodia with conical ante- rior lobes and projecting acicular processes and rounded posterior lobes (Fig. 2C, D). Notosetae numerous, forming radiating bundles, much stouter than neurosetae, short, slightly curved to longer, straight, with spinous rows and tapered bare tips (Fig. 2E). Neurosetae numerous, slender, with faint spinous rows and bare, slightly hooked tips with small subterminal tooth; lower ones with entire curved tips (Fig. 2F). Ventral cirri short, tapered, with slender tips (Fig. 2C, D). Some eggs in parapodia medial to bases of ventral cirri. Distribution. —East Central Pacific in Ga- lapagos Rift, in 2482 m and North Pacific in Juan de Fuca Ridge, 2059 m. Acknowledgments I wish to thank Cindy van Dover of the Woods Hole Oceanographic Institution and Verena Tunnicliffe of the University of Vic- toria for their part in collecting and sorting of the material on which this study is based. The manuscript benefited from the com- ments by two anonymous reviewers. Literature Cited Hartmann-Schréder, G. 1971. Annelida, Borsten- wurmer, Polychaeta.—Die Tierwelt Deutsch- lands und der angrenzenden Meeresteile 58:1- 549. 1974. Die Unterfamilie Macellicephalinae Hartmann-Schréder, 1971 (Polynoidae, Poly- 614 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON chaeta). Mit Beschriebung einer neuer Art, Ma- cellicephala jameensis n. sp., aus einem H6h- lengewasser von Lanzarote (Kanarische Inseln).— Mitteilungen aus dem Hamburgi- schen Zoologischen Museum und Institut 71- 75-85. Horst, R. 1917. Polychaeta Errantia of the Siboga Expedition Part 2. Aphroditidae and Chryso- petalidae. Pp. 1-140 in number 24b of Siboga- Expedite. E. J. Brill, Leiden. Kinberg, J.G. H. 1856. Nye slagter och arter af An- nelider.— Ofversigt af Konglia Vetenkaps-Aka- demiens Forhandlingar, Stockholm 12:381-388. Levenstein, R. 1962. [Polychaete worms from three abyssal trenches of the Pacific Ocean].—Zoo- logicheskii Zhurnal Akademiia Nauk SSSR 41: 1142-1148. [In Russian, English summary]. Moore, J. P. 1910. The polychaetous annelids dredged by the U.S:S. “Albatross” off the coast of south- ern California in 1904: Polynoidae, Aphroditi- dae and Segalionidae. — Proceedings of the Academy of Natural Sciences of Philadelphia 62:328—402. Muir, A. I. 1982. Generic characters in the Poly- noinae (Annelida, Polychaeta), with notes on the higher classification of scale-worms (Aphroditacea).— Bulletin of the British Muse- um (Natural History), Zoology, London 43:153- 177. Pettibone, M. H. 1976. Revision of the subfamily Macellicephala McIntosh and the subfamily Macellicephalinae Hartmann-Schréder (Poly- chaeta: Polynoidae.—Smithsonian Contribu- tions to Zoology 229:1-71. . 1985a. Polychaete worms from a cave in the Bahamas and from experimental wood panels in deep water of the North Atlantic (Polynodae: Macellicephalinae, Harmothoinae).—Proceed- ings of the Biological Society of Washington 98: 127-149. 1985b. New genera and species of deep-sea Macellicephalinae and Harmothoinae (Poly- chaeta: Polynoidae) from the hydrothermal rift areas off the Galapagos and Western Mexico at 21°N and from the Santa Catalina Channel.— Proceedings of the Biological Society of Wash- ington 98:740-757. 1989. Polynoidae and Sigalionidae (Poly- chaeta) from the Guaymas Basin, with descrip- tions of two new species, and additional records from Hydrothermal Vents of the Galapagos Rift, 21°N, and Seep-Sites in the Gulf of Mexico (Florida and Louisiana).—Proceedings of the Biological Society of Washington 102:154—168. 1993. Polynoid polychaetes associated with a whale skeleton in the bathyal Santa Catalina Basin. — Proceedings of the Biological Society of Washington 106:678-688. Department of Invertebrate Zoology, Na- tional Museum of Natural History, Smith- sonian Institution, Washington, D.C. 20560, U.S.A. PROC. BIOL. SOC. WASH. 107(4), 1994, pp. 615-621 A NEW GENUS AND SPECIES OF POLYCHAETE, BOLLANDIA ANTIPATHICOLA (NEREIDOIDEA: SYLLIDABE), FROM BLACK CORAL Christopher J. Glasby Abstract.—A new genus and species of syllid polychaete, Bollandia anti- pathicola, is described. This is the first syllid species to be found in association with an antipatharian coral. It differs from other Syllidae in having one pair of peristomial cirri and two pairs of tentacular cirri (i.e., a total of three pairs of cirri on the anterior end), lacking palps and antennae, and in having few specialized stout setae. The species is hermaphroditic and possesses many paired ventral reproductive papillae that may be used in copulation or for ejecting spermatophores. Routine sorting of a collection of benthic invertebrates from 52-61 m off Okinawa, Japan, yielded an unusual polychaete that was attached to Black Coral, Antipathes sp. The species clearly belonged to the order Phyllodocida as it had a muscular, axial type of eversible pharynx, unlike the ventral type found in other polychaete orders having a pharynx such as the Eunicida and Amphi- nomida (Dales 1962, Orrhage 1973). Fur- ther, the species was suspected of belonging to the Nereidoidea as it had at least two pairs of tentacular cirri presumably result- ing from the cephalization of the parapodia of the first two segments, as for example in some species of the Hesionidae (Glasby 1993). The Nereidoidea currently contain six families: the Chrysopetalidae, Hesioni- dae, Nautiliniellidae, Nereididae, Pilargi- dae, and Syllidae (Glasby 1993; Glasby & Fauchald 1991). The species could not ini- tially be assigned to any of these based on external features; however, a histological study revealed an axial pharynx that was fully differentiated (sensu Glasby 1993), that is one having a proventriculus and muscular ventriculus, a synapomorphy of the Sylli- dae. The association between the new syllid polychaete and the Antipathes may be com- mensal, although there is insufficient infor- mation available on the biology of either organism to be certain. Commensalism has been reported previously among the Sylli- dae with species associated with poriferan hosts (Cognetti 1957, Pearse 1934), ophiu- roid hosts (e.g., Hendler & Meyer 1982) and a eunicid polychaete (Hempelmann 1931). Although there are several records of cni- darian hosts (Utinomi 1956, Hartmann- Schréder 1960, Laubier 1960, Wright & Woodwick 1977, Hartmann-Schroder 1991, 1992) there appear to be no previous rec- ords of syllids being associated with anti- patharian corals (Anthozoa: Ceriantharia). However, Pettibone (1991) recorded four polynoid species from antipatharians. Fur- ther, Pettibone (and references therein) found that the commensal annelids could induce the formation of tunnels in the coe- nenchyme of the main stems of the coral: tunnels were formed by anastomosed twigs. No such modification to the coral was ob- served in the present study. Other commensal nereidoids include An- tonbruunia viridis Hartman & Boss, recently moved to the Pilargidae (Glasby 1993) and species of the Nautiliniellidae all of which 616 are commensals or parasites of bivalve mol- luscs (Hartman & Boss 1965, Miura & Lau- bier 1989, Blake 1993). Specimens described in this paper have been deposited with the National Museum of Natural History, Smithsonian Institution (USNM), Natural History Museum, Lon- don (BMNH), and the Australian Museum (AM). Family Syllidae Grube, 1850 Bollandia, new genus Type species.—Bollandia antipathicola, new species. Diagnosis. — Antennae absent; palps ab- sent; nuchal organs not externally exposed (sensu Glasby 1993); three pairs of append- ages on anterior end, most probably rep- resenting one pair of peristomial cirri and two pairs of tentacular cirri (1.e., parapodia of segments 1 & 2 cephalized during ontog- eny; but see Remarks); pharyngeal appara- tus consisting of sinuous, unarmed pharynx, and a barrel-shaped proventriculus; noto- podia and notosetae absent; neurosetae sim- ple, occur in the sub-acicular position only; anal cirri paired. Etymology. —The genus has feminine gender, and is named in honor of R.F. Bol- land who collected the specimens. The spe- cific name refers to the genus of Black Coral, Antipathes, on which the new species was found. Bollandia antipathicola, new species (Figs. 1A—H; 2A—D) Material examined.— All material from Japan, Okinawa, Horseshoe Cliffs, 1 km WNW Onna village, 26°30.0'N, 127°50.9’E, coll. R. F. Bolland. Holotype: Stn RFB 1240, 61 m, coll. 5 Aug 1984 (USNM 169157). Paratypes: Stn RFB 1240, 61 m, coll. 5 Aug 1984, 7 specimens (USNM 169158), 2 spec- imens mounted for SEM (USNM 169159), 2 specimens sectioned (USNM 169160), 2 specimens (AM W21804), 2 specimens PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON (BMNH 1994.3202-3203); Stn RFB 1235, 57.9 m, coll. 3 Aug 1984, 2 specimens (USNM 169161); Stn 1243, 64.0 m, coll. 10 Aug 1984, 6 specimens (USNM 169162); Stn RFB 1276, 51.8 m, coll. 10 Oct 1984, 1 specimen (USNM 169163); Stn RFB 1974, 61 m, coll. 28 Aug 1988, 2 specimens (USNM 169164), 1 specimen mounted for SEM (USNM 169165). Description.—Holotype 73 setigers, 6.0 mm long, 0.4 mm wide. Paratype material ranged in size from 16 setigers, 1.3 mm long, 0.2 mm wide (USNM 169165) to 47 setig- ers, 6.2 mm long, 0.3 mm wide (USNM 169160). Body approximately uniform in width throughout, highly arched dorsally particularly over setigers 1-4 (Figs. 1A, C) which carry the pharyngeal apparatus, flat- concave ventrally with parapodia directed ventrolaterally (Figs. 1B, 2D). Preserved material with diffuse brown pigment on ventral surface of setiger 4 (absent in smaller paratypes) and from setiger 8 to near py- gidium, particularly intense around base of ventral cirri. Dorsum, parapodia and cirri covered with clumps of long cilia (Fig. 1A). Prostomium small, ill-defined, directed anteroventrally, lacking antennae or palps (Figs. 1B, 2D). Eyes very small, red, two pairs, posterior pair slightly further apart than anterior pair. Peristomium indistinct, carrying a pair of smooth peristomial cirri (C1), about twice width of prostomium, di- rected anteriorly (Fig. 2D). First two seg- ments cephalized, fused together and with peristomium, ventral surface slightly raised above level of succeeding segments (Fig. 1B); first pair of tentacular cirri (C2) smooth, lacking aciculae, slightly dorsally displaced, similar in shape though slightly longer than peristomial cirri, basally with pair of minute vestigial ventral cirri (V1; not visible in ho- lotype); second pair of tentacular cirri (C3) also smooth and lacking aciculae, similar in size and shape to C2, basally with a pair of small vestigal ventral cirri (V2; Figs. 1B, 2D). VOLUME 107, NUMBER 4 617 Fig. 1. A-H: Bollandia antipathicola, new genus, new species, Paratypes (USNM 169160): A, in situ on Antipathes sp.; B, anterior end, anteroventral view; C, anterior end, longitudinal section; D, midbody, longitudinal section; E, midbody, transverse section; F, sperm containing tubules, midbody; G, reproductive papillae, mid- body; H, posterior end, ventral view. Scale bars: 100 um (A, C, D, E), 40 um (B, H), 20 um (F), 5 um (G). 618 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. A-—D: Bollandia antipathicola, new genus, new species: A, Paratype (USNM 169158) anterior end, dorsal view; B, Holotype, parapodium from midbody, anterior view; C, Holotype, setae from parapodia of midbody; D, anterior end, ventral view, composite illustration compiled from 3 paratypes (USNM 169158) [Cl = peristomial cirri; C2, C3 = tentacular cirri; V1—3 = ventral cirri]. Scale bars: 0.10 mm (A, D), 0.05 mm (B), 0.02 mm (C). VOLUME 107, NUMBER 4 Pharyngeal apparatus housed in anterior dorsally vaulted region (Fig. 1C). Pharynx cylindrical, extending back to about second setiger, sinuous, apparently unarmed (al- though not dissected). Proventriculus bar- rel-shaped, extending from pharynx back to about setiger 4, with about 28-35 groups of radial muscle columns (Fig. 1C). Segment 3 first setigerous. Parapodia con- sisting of a single conical neuropodial lobe supported by one acicula and carrying two setae (Figs. 2B, D). Dorsal cirri atop very short cirrophore, tapered, faintly articulated distally, with very closely spaced articles; about equal in length to neuropodial lobe. Ventral cirri very slender, adpressed to par- apodia, about ' (4—'2) length of neuropo- dial lobe (Figs. 2B, D). Subsequent para- podia similar to first, except length of dorsal cirri 1-3 times length of corresponding neu- ropodial lobes, generally increasing slightly in length posteriorly, though longer ones oc- curring every 4—7 segments. Setae of two types, both only slightly less stout than cor- responding acicula: highly tapered flail- tipped setae, slightly bent near tip; and slightly stouter setae, distally bent at right- angles forming beak-like tip (Fig. 2C). Pygidium swollen, glandular, pair of con- ical ventrolateral anal cirri (Fig. 1H; absent in holotype). Reproductive papillae present on elevat- ed pads at ventral base of parapodia (Fig. 1G); resemble ventral cirri in both size and shape. Beginning on setigers 2—5 reproduc- tive papillae extend posteriorly to last se- tigers. Papillae fed by internal tubules filled with mature sperm (Figs. 1D-F). Perhaps also early stages of sperm present, but not classical spermatids and no intermediate stages between these early stages and mature stages. Oocytes spherical, unpigmented, ranged in diameter from about 10-50 um (1 specimen, USNM 169163), present in all specimens in posterior half of body from setigers 9-11 to near pygidium. Remarks. —The species is hermaphrodit- 619 ic, although it is difficult to estimate from so few specimens whether it is a sequential or a simultaneous hermaphrodite. Although resembling some types of seminal recepta- cles (spermathecae), the ‘internal tubules’ are more likely to represent seminal vesicles or testes since they appear to empty to the exterior via reproductive papillae. Repro- ductive papillae may be used for direct cop- ulation, as in Pisione remota (Stecher 1968), although no gametal pores could be iden- tified on any specimen. Alternatively they may function for ejecting spermatophores that could be fixed to (and later penetrate) the body of another individual, as in He- sionides arenaria (Westheide 1967). The precise nature of the anterior end ap- pendages is uncertain, since the distinction between anterior segments, peristomium and prostomium is unclear. However, long, slender appendages in the Nereidoidea are more likely to represent cirri, either of peri- stomial or segmental origin, than antennae or palps. Assuming that the presence of the vestigial ventral cirri (Fig. 2D) at the ventral base of the anterior end cirri indicates a segmental origin, then clearly two segments have been cephalized. The question is whether the first pair of ventral cirri (V1) is associated with the first (Cl) or the second (C2) pair of cirri. If associated with Cl then the first pair are tentacular cirri derived from segment | and the second and third pairs (C2, C3) are also tentacular cirri perhaps both derived from segment 2 (Fig. 2D). If this were the case then Bollandia would lack peristomial cirri. However, a more likely hypothesis, and the one proposed here, is that V1 is associated with C2, implying that Cl are peristomial cirri and the C2 and C3 are derived from segments 1 and 2 respec- tively. Most Syllidae have one or two pairs of anterior end cirm (Fauchald 1977, Gar- wood 1991). Strictly speaking the cirri are peristomial in origin and should therefore be referred to as peristomial cirri (Glasby 620 1993), although the term tentacular cirri has also been used in the literature. Exceptions are those genera that lack peristomial cirri altogether, including Exogonella Hartman, Fauvelia Gravier, Haplosyllides Augener, and Nudisyllis Knox & Cameron; Irmula Ehlers supposedly has six pairs of tentacular cirri. Bollandia antipathicola, new species, differs from these and other species of Syl- lidae in having one pair of peristomial cirri and two pairs of tentacular cirri (=three pairs of anterior end cirri). In addition it lacks antennae and palps and has few specialized stout setae. The lack of palps appears to have been recorded for only one other syllid genus, Haplosyllides Augener. Haplosyllides differs from Bollandia in having three an- tennae and lacking both peristomial and tentacular cirri (Augener 1922). Most Syllidae undergo some form of structural (epitokous) modification associ- ated with sexual maturity (Garwood 1991). Neither schizogamy, in which sexual indi- viduals are budded off from the adult, nor epigamy, in which the whole animal un- dergoes modification were observed in the present specimens of Bollandia. Like the new Bollandia species, palps are often absent in epigamous (=epitokous) syllids (Estapé & San Martin 1991). However, the new spe- cies can not represent an epitokous syllid since epitokes generally lack peristomial cir- ri, have enlarged eyes and many capillary (swimming) setae and their bodies are often turgid with gametes. The species resembles most closely those of the subfamily Eusyllinae, which was re- cently defined by Garwood (1991). In com- mon with the Eusyllinae, Bollandia has smooth or indistinctly articulated dorsal cirri and cirri of the anterior end, the presence of ventral cirri, and segmental ciliation that is retained in adults. It differs from the Eu- syllinae however, in having 3 pairs of an- terior end cirri (cf. two pairs, according to Garwood) and in lacking palps, although other members of the Eusyllinae may have reduced palps (Garwood 1991). PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON In my opinion the subfamilial groupings within the Syllidae remain controversial (see also Fauchald 1977) despite the findings of Garwood (1991), which support the tradi- tional four subfamilial groupings. The Syl- lidae have never been subject to any sort of rigorous phylogenetic analysis and therefore the Eusyllinae, along with other syllid sub- families, may not constitute a monophyletic group. Until monophyletic groups within the Syllidae can be identified, the phyloge- netic relationships of Bollandia must re- main obscure. Acknowledgments Research toward this paper was conduct- ed during a Postdoctoral Fellowship at the National Museum of Natural History, Smithsonian Institution. I would like to thank Barbara Littman of the Smithsonian Oceanographic Sorting Center for drawing my attention to this unusual polychaete. Also I thank Dr. Dennis Opresko for iden- tifying the Black Coral, Ms. Belinda Alvarez for her help in printing photographs for Fig. 1 and Dr. Gregory Rouse for his help with the Scanning Electron Micrography and dis- cussions on the reproductive biology of the species. The comments of an anonymous reviewer are also appreciated. Literature Cited Augener, H. 1922. Ueber litorale polychaeten von Westindien. — Gesellschaft Naturforschender freunde zu Berlin, Sitzungsberichte, 38-53. Blake, J. A. 1993. New genera and species of deep- sea polychaetes of the family Nautiliniellidae from the Gulf of Mexico and the Eastern Pa- cific.— Proceedings of the Biological Society of Washington 106:147-157. Cognetti, G. 1957. I sillidi del Golfo di Napoli.— Pubblicazione della Stazione Zoologica di Na- poli 30:1—100. Dales, R. P. 1962. The polychaete stomodeum and the inter-relationships of the families of Poly- chaeta.— Proceedings of the Zoological Society of London 139(3):389-428. Estapé, S., & G. San Martin. 1991. Descripcion de los estolones reproductores de algunas especies de la subfamilia Syllinae (Polychaeta, Sylli- VOLUME 107, NUMBER 4 dae). — Miscelanea zoologica, Barcelona 15:43- 62. Fauchald, K. 1977. The Polychaete Worms. Defini- tions and Keys to the Orders, Families and Gen- era.— Natural History Museum of Los Angeles County, Science Series 28:1—190. Garwood, P. R. 1991. Reproduction and the classi- fication of the family Syllidae (Polychaeta). Pp. 81-87 in M. E. Petersen & J. B. Kirkegaard, eds., Systematics, biology and morphology of world Polychaeta. Proceedings of the 2nd In- ternational Polychaete Conference Copenhagen 1986.—Ophelia, Supplement 5. Glasby, C. J. 1993. Family revision and cladistic analysis of the Nereidoidea (Polychaeta: Phyl- lodocida).—Invertebrate Taxonomy 7:1551- 1573. —,, & K. Fauchald. 1991. Redescription of Hel- metophorus rankini Hartman, 1978 (Polychae- ta: Helmetophoridae) and its transfer to the Fla- belligeridae. — Proceedings of the Biological Society of Washington 104:682-685. Hartman, O., & K. J. Boss. 1965. Antonbruunia viridis, a new inquiline annelid with dwarf males, inhabiting a new species of pelecypod, Lucina fosteri, in the Mozambique Channel.—The An- nals & Magazine of Natural History, Zoology, Botany, and Geology (Series 13) 87 & 88:177-— 186. Hartmann-Schroder, G. 1960. Polychaeten aus dem Roten Meer. —Kieler Meeresforschungen 16:69— 15). . 1978. Einige sylliden-arten (Polychaeta) von Hawaii und as dem Karibischen Mer. —Mittei- lungen aus dem Hamburgischen Zoologischen Museum und Institut 75:49-61. 1991. Syllis onkylochaeta sp. n., ein koral- lenfressender polychaet (Syllidae) aus dem Ko- rallenaquarium des Lobbecke-Museums. — Hel- golander Meeresuntersuchungen 45:59-63. . 1992. Drei neue polychaeten-arten der fami- lien Polynoidae und Syllidae von Neu-Kale- donien, assoziiert mit einer verkalkten Hydro- zoe.—Helgolander Meeresuntersuchungen 46: 93-101. Hempelmann, F. 1931. Erste und zweite klasse der Vermes Polymera (Annelider). Archiannelida and Polychaeta. Jn W. Kiikenthal & T. Krum- bach, eds., Handbuch der Zoologie 2(7):1—212. Hendler, G., & D. L. Meyer. 1982. An association of a polychaete, Branchiosyllis exilis with an 621 ophiuroid, Ophiocoma echinata, in Panama.— Bulletin of Marine Science 32(3):736-744. Laubier, L. 1960. Une nouvelle sous-espéce de syl- lidien: Haplosyllis depressa Augener ssp. nov. chamaeleon, ectoparasite sur l’octocoralliaire Muricea chamaeleon von Koch.— Vie et Milieu 11:75-87. Miura, T., & L. Laubier. 1989. Nautilina calypto- genicola, a new genus and species of parasitic polychaete on a vesicomyid bivalve from the Japan Trench, representative of a new family Nautilinidae. — Zoological Science 6:387-390. Orrhage, L. 1973. Two fundamental requirements for phylogenetic-scientific works as a background for an analysis of Dales’s (1962) and Webb’s (1969) theories.—Zeitschnft fiir Zoologische Systematik und Evolutionsforschung 1 1(3):161- 173. Pearse, A. S. 1934. Inhabitants of certain sponges at Dry Tortugas.— Carnegie Institute of Washing- ton, Papers of Tortugas Laboratory 28(7):117- 124. Pettibone, M. H. 1991. Polynoid polychaetes com- mensal with antipatharian corals. — Proceedings of the Biological Society of Washington 104: 714-726. Stecher, H.-J. 1968. Zur Organisation und Fortpflan- zung von Pisione remota (Southern) (Polychae- ta, Pisionidae).— Zeitschrift flir Morphologie der Tiere 61:347-410. Utinomi, H. 1956. On the so-called ““Umi-utiwa’’, a peculiar flabellate gorgonacean, with notes on a syllidean polychaete commensal. — Publications of the Seto Marine Biological Laboratory 5(2): 243-250, pl. 27. Westheide, W. 1967. Monographie der Gattungen Hesionides Friedrich und Microphthalmus Me- cznikow (Polychaeta, Hesionidae). Ein Beitrag zur Organisation und Biologie psammobionter Polychaeten. — Zeitschrift fir Morphologie der Tiere 61:1-159. Wright, J. D., & K. H. Woodwick. 1977. A new spe- cies of Autolytus (Polychaeta: Syllidae) com- mensal on a California hydrocoral.— Bulletin of the Southern California Academy of Science 76: 42-48. Australian Biological Resources Study, P.O. Box 636, Canberra ACT 2601, Aus- tralia. PROC. BIOL. SOC. WASH. 107(4), 1994, pp. 622-633 NEW SPECIES OF DIPLOCARDIA AND ARGILOPHILUS (ANNELIDA: OLIGOCHAETA: MEGASCOLECIDAE) FROM SOUTHERN CALIFORNIA Samuel W. James Abstract. —Five new species of megascolecid earthworms, Diplocardia cali- forniana, D. woodi, D. montana, Argilophilus woodi, and A. margaritae are described from material collected in various wildland habitats in the mountains neighboring the Los Angeles Basin. No other native earthworms were previ- ously known from the region. The Diplocardia species are related to D. keyesi, known from Baja California, and all occur to the south of Los Angeles. Other Argilophilus are known from sites farther north; those described here are found to the north of the greater Los Angeles area. The northern limit of Diplocardia in California appears to be the San Gorgonio Pass. This may also be the southern limit of Argilophilus in California, but further collecting will be needed for confirmation. It has been almost a century since anyone made an organized attempt to study terres- trial Oligochaeta in Southern California. The species presented here came to light as a result of the industriousness of Hulton B. ‘Hutch’ Wood during his preliminary in- vestigations of biotic factors affecting soils in the San Dimas Experimental Forest. To- gether we collected extensively in the moun- tains and wildlands surrounding the Los Angeles Basin. It may come as a surprise to learn that an area noted for a long dry period would support indigenous earthworms in unirrigated land. Though many sites were in riparian zones or adjacent to other sources of water, several were not so favored and not located at elevations where precipita- tion is abundant. A full account of the sites visited, their soils and vegetation, and the earthworm species encountered is in Wood & James (1993). Five species are new and are described here. Additional material was collected in early 1993, and further data on the new species were taken from these col- lections. There is also one more new species from the 1993 collections. All specimens were obtained by digging and handsorting, and were killed in 50% ethanol and fixed in 5% formalin. Exami- nations were conducted by dorsal dissection under a stereomicroscope with a drawing tube. Diplocardia californiana, new species Fig. 1A—C Type material. —Holotype: USNM 169803, In grass/lupine meadow near ver- nal pool on Mesa de Colorado, Santa Rosa Plateau, near Temecula, Riverside Co., Cal- ifornia, 5 Apr 1990, S. W. James, H. B. Wood, K. L. Olivier, collectors; Paratypes: USNM 169804, same locality as holotype. Additional material. —In grass pasture of Love Valley, Cleveland National Forest, 22 Apr 1991, S. W. James and H. B. Wood, collectors; In grass among pines, Upper French Valley near Mt. Palomar Observa- tory, 22 Apr 1991, S. W. James and H. B. Wood, collectors; In grass adjacent to live oak grove, Falcon Camp, Cleveland Na- tional Forest, 24 Apr 1991, S. W. James and H. B. Wood, collectors; Long Canyon, Cleveland National Forest, near California Highway 74 in grassy area among oaks, 24 Apr 1991, S. W. James and H. B. Wood, VOLUME 107, NUMBER 4 collectors; In grass/geranium mix and oak chaparral near Tenaja Creek, Cleveland Na- tional Forest, 24 Apr 1991, S. W. James and H. B. Wood, collectors. In grass/gera- nium mix and oak chaparral near Tenaja Creek, Cleveland National Forest, 6 Apr 1993, K. L. Olivier and H. B. Wood, col- lectors. Miller Mountain, in the bowl, basalt rock substrate with clay soil, grass, 15 Apr 1993, K. L. Olivier and R. Mees, collectors. Description. —External characteristics: Dimensions 58-85 mm by 2.4—3.0 mm at segment xxx, 2.8-3.3 mm at vii; body cy- lindrical throughout, segments 106-164. Setae closely paired throughout; setal for- mula AA:AB:BC:CD = 3:1:4:1.3 at x, 3.5: 1:3:1 at xxx, DD > % circumference throughout. Prostomium epilobous to near- ly tanylobous, segments with postsetal sec- ondary annulus v, pre- and postsetal annuli vi-end, in x+ each third has tertiary an- nulus. Brown pigmentation present in i—v, vi, vii, denser dorsally; sometimes also pres- ent in male field area. Nephridiopores at D, first dorsal pore 9/10 or 10/11, spermathe- cal pores at leading edges of viii, ix lateral to A. Ovipores presetal median to A in xiv; male pores at 21/22; prostatic pores and penial setae at ends of seminal grooves in AB in xxi—xxili. Clitellum xii—xxi, xxii, sad- dle-shaped; no genital markings (Fig. 1A). Internal characteristics: Septa 5/6—-11/12 muscular, greatest thickness at 8/9; 5/6 only faintly muscular. Alimentary canal with two gizzards in Vv, vi; appearing as one unit with- out substantial demarkation; esophagous with pebbly internal texture ix—x11, low lon- gitudinal folds 13/14—xvili, ventral esoph- ageal ridge xi—xvii, esophagous valvular in XVlll, xix, intestinal origin xx; typhlosole a simple fold originating over xxili—xxv, height less than one tenth lumen diameter. No cal- ciferous glands. Stomate meganephridia 2 per segment, exoic with duct entering body wall in CD, avesiculate, tubules in elongate flat coil over AD. Vascular system with ventral trunk, single dorsal trunk, these connected by lateral 623 trunks in v—ix, latero-esophageal hearts in x—xil. Extra-esophageal vessel from pharyn- geal glands, along ventral-lateral face of giz- zard, ventral face of esophagous v—xiii, in xiii branching out to body wall of xili—xx. Supra-esophageal vessel x—xill. Fan-shaped ovaries composed of long strings, with funnels in xiii; paired sper- mathecae in vili, ix, each an ovoid ampulla with sessile diverticulum composed of 3-6 internal lobes arranged in one row (Fig. 1B); ampulla under esophagous, ampulla long axis usually perpendicular to duct axis. Male sexual system holandric, testes and funnels free in x, xi; acinous equal-sized seminal vesicles in ix, xil; vasa deferentia superficial, very thin, enter body wall at 21/ 22; tubular prostates with very short slightly muscular ducts, gland in several folds with- in segments of origin (xxi, xxili); penial setal follicles just anterior to ducts. Penial setae 0.25-0.3 mm by 0.015 mm, nearly straight, gradually tapering to blunt tip (Fig. 1C); genital setae lacking. Diagnosis. —Diplocardia with male field iM XX1—XX1ll. Remarks. —Specimens from Love Valley, Tenaja Creek and Falcon Camp had abun- dant melanocytes in the linings of the body cavity, on blood vessels, nephridia, and oth- er organs in the region xi—xxv. External pig- mentation was slightly variable among sites, the darkest being those of Mesa de Colorado and the lightest coming from Falcon Camp and Upper French Valley. Oil droplets were abundant in the coeloms of worms from Long Canyon and Love Valley. Much of the material showed evidence of reduction in male functions. Seminal vesicles varied from partly filled to very small to rudimentary, even in fully clitellate individuals. Irides- cence of male funnels was uncommon, and iridescence of spermathecal diverticula was seen only in the 1993 Tenaja Creek mate- rial. Prostates were quite small for Diplo- cardia, and penial setae were little more than straightened ambulatory setae. Some of the lack of male development could be season- 624 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 1.0mm 0.01 mm wil O'S Fig. 1. Diplocardia californiana: A. ventral view, B. spermatheca, C. penial seta. VOLUME 107, NUMBER 4 al, many of the sites having been at high elevations. However, all material examined was fully clitellate except the worms from Upper French Valley, the highest elevation site. Diplocardia californiana is referred to as Diplocardia CA\ in Wood & James (1993). Diplocardia keyesi Eisen, 1900 has the male field in xx—xxul. This would place it as the nearest relative of the new Diplocardia described here. Diplocardia keyesi is known only from a single location in northern Baja California, near Ensenada. The location of the male field or male pores is usually very conservative in earthworms, but is quite variable in Diplocardia. Diplocardia woodi, new species Fig. 2A, 2B Type material. —Holotype: USNM 169806, Mesa de Burro, Santa Rosa Pla- teau, Riverside Co., California. 1 Apr 1993, K. L. Olivier, collector; Paratypes: USNM 169807, same locality as holotype. Additional material.—Mesa de Burro, Santa Rosa Plateau, Riverside Co., Califor- nia. 5 Apr 1990, S. W. James, H. B. Wood and K. L. Olivier, collectors. Description. —External characteristics: Dimensions 63-78 mm by 2.7—2.9 mm at segment xxx, 3.2 mm at 1x; body cylindrical throughout, segments 134—153. Pigmenta- tion lacking or present as slight rings of brown pigment at segmental equators of some or all of ii—v. Setae closely paired throughout; setal formula AA:AB:BC:CD = 3.5:1:3:1.3 at x, 4:1:3.5:1.3 at xxx, DD > Y% circumference throughout. Prostomium epilobous, segments with postsetal secondary annulus vi—vii, pre- and postsetal annuli viii—end, in xxii+ postsetal third has tertiary annulus. Nephridiopores not seen, first dorsal pore 9/10 or 10/11, spermathecal pores at leading edges of viii, ix lateral to A. Ovipores presetal median to 625 A in xiv; male pores at 21/22; prostatic pores and penial setae at ends of seminal grooves in AB in xxi—xxili. Clitellum saddle-shaped, interrupted at mid-ventral line, x1i-xx1; no genital markings (Fig. 2A). Internal characteristics: Septa 6/7-11/12 muscular, greatest thickness at 8/9; 13/14 only faintly muscular. Alimentary canal with two gizzards in v, vi; appearing as one unit with thin region at 5/6; esophagous with pebbly internal texture 1x—xvili, ventral esophageal ridge 12-18, valvular in xix, in- testinal origin xx; typhlosole simple fold originating over xxiil—xxv, height one fifth lumen diameter. No calciferous glands. Sto- mate meganephridia 2 per segment, exoic with duct entering body wall in CD, ave- siculate, tubules in elongate flat coil over BD. Vascular system with ventral trunk, single dorsal trunk, these connected by lateral trunks in v—1x, latero-esophageal hearts in x—-xil. Extra-esophageal vessel from pharyn- geal glands, along ventral-lateral face of giz- zard, ventral face of esophagous v—xiil, in xiii branching out to body wall of xili—xx. Supra-esophageal vessel ix—or x—xiil. Fan-shaped ovaries composed of long strings, with funnels in xiii; paired sper- mathecae in viii, 1x, each an ovoid ampulla with sessile diverticulum composed of 1-3 internal lobes (Fig. 2B). Male sexual system holandric, testes and funnels free in x, x1; seminal vesicles in 1x, xii, both quite small; vasa deferentia super- ficial, very thin, enter body wall at 21/22; tubular prostates with very short slender ducts less than one-tenth gland length, gland in several folds within segments of origin (xxi, xx1i1); penial setal follicles just anterior to ducts, penial setae not enlarged (4) or lacking (1); genital setae lacking. Diagnosis.—Diplocardia with male field in xxi—xxili, and distinguished from D. cal- iforniana by the greater height of the typh- losole, smaller nephridia, fewer lobes of the spermathecal diverticulum, lack of mus- 626 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON N ) 3 3 C D 1.0 mm 1.0mm Fig. 2. Diplocardia woodi: A. ventral view, B. spermatheca. Diplocardia montana: C. ventral view, D. spermatheca. VOLUME 107, NUMBER 4 cularity of prostatic ducts, and lack of de- velopment of penial setae. Remarks. —Iridescence of the male fun- nels was detected on one individual from the 1993 material, though coagulum was present in segments x and xi of all material examined. Spermathecal diverticula of the 1993 material were iridescent, but those of the 1990 material were not. The species is named after Hulton B. Wood. It is referred to as ““Diplocardia CA2” in Wood & James (1993). Diplocardia montana, new species Fig. 2C, 2D Type material.—Holotype USNM 169805, Miller Mountain, San Diego Co. California, top end of jeep trail, 900 m elev., clay loam soil and grass; 15 Apr 1993, K. L. Olivier and R. Mees, collectors. Description. —External characteristics: Dimensions 103 mm by 3.5 mm at segment xxx, 4 mm at 1x; body cylindrical through- out, segments 152. Pigmentation present as slight rings of brown pigment at segmental equators on ii—vuli. Setae closely paired throughout; setal formula AA:AB:BC:CD = 4:1.3:4:1 at x, 3:1:4:1.2 at xxx, DD > % circumference throughout. Prostomium prolobous, segments with postsetal second- ary annulus vi—vii, pre- and postsetal annuli vili—end, in xxii+ postsetal third has terti- ary annulus. Nephridiopores not seen, first dorsal pore 10/11, spermathecal pores on small bumps at leading edges of viii, ix in A. Ovipores presetal median to A in xiv; male pores at 21/22; prostatic pores at ends of seminal grooves in AB in xxi—xxili. Cli- tellum saddle-shaped, interrupted at mid- ventral line, xii—xxii; no genital markings (Fig. 2C). Internal characteristics: Septa 6/7—11/12 muscular, greatest thickness at 8/9; 12/13 only faintly muscular. Alimentary canal with two gizzards in v, vi; appearing as one unit 627 with thin region at 5/6; esophagous with pebbly internal texture ix—xvili, paired ven- tral esophageal ridges xiii—xviii, valvular in xix, intestinal origin xx; typhlosole simple fold originating 22/23, height one tenth lu- men diameter or less. No calciferous glands. Stomate meganephridia 2 per segment, ex- oic with duct entering body wall in CD, avesiculate, tubules in elongate flat coil over BD. Vascular system with ventral trunk, single dorsal trunk, these connected by lateral trunks in v—1x, latero-esophageal hearts in x—xil. Extra-esophageal vessel from pharyn- geal glands, along ventral-lateral face of giz- zard, ventral face of esophagous v—xili, in xill branching out to body wall of xili—xx. Ventral esophageal ridges contain blood vessels connected to extra-esophageals. Su- pra-esophageal vessel x—x1il. Fan-shaped ovaries composed of long strings, with funnels in xiii; paired sper- mathecae in vill, 1x, each an ovoid ampulla with sessile diverticulum composed of 1-3 internal lobes (Fig. 2D). Male sexual system metandric, testes and funnels free in xi; seminal vesicles in xii; vasa deferentia superficial, very thin, enter body wall at 21/22; tubular prostates with very short slender ducts so that prostates are nearly sessile, gland in several folds within segments of origin (xxi, xxill) or one adjacent segment; penial setal follicles just anterior to ducts, penial setae vestigial; gen- ital setae lacking. Diagnosis. — Diplocardia with metandric reduction of the male gonads, male field in XXI—XXI11. Remarks.—Apart from the metandric condition, D. montana is very similar to its southern California congeners. However, its one pair of male funnels and its sperma- thecal diverticula were iridescent, and there was no other evidence of reduction of male functions. It is also somewhat larger than the other species. So far it is the only known metandric Diplocardia. 628 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Argilophilus woodi, new species Fig. 3A—C Type material. —Holotype, USNM 169799. Type locality: in oaks and sage- brush, 1300 m elev., near milepost 41.50, California Hwy. 33, Ventura Co. California, 4 Apr 1990, S. W. James and H. B. Wood, collectors. Paratypes (USNM 169800) from same locality, three adults, 4 Apr 1990, S. W. James and H. B. Wood, collectors. Description. —External characteristics: Dimensions 51-58 mm by 3.5-4 mm (strongly contracted), width at segment xxx, body cylindrical throughout, segments 125- 140. Pigmentation lacking. Setae ab closely paired throughout; setal formula AA:AB: BC:CD = 3.5:1:2.5:2.5 at xxx, DD > 4 cir- cumference throughout. Prostomium epi- lobous, segments with postsetal secondary annulus vi—ix, pre- and postsetal annuli x— xul. Nephridiopores not seen, first apparent dorsal pore 18/19, but dorsal pores merely thin spots in body wall, few if any actually Open; spermathecal pores at leading edges of viii, ix at B. Ovipores presetal median to A in xiv; male pores in xviii on small pa- pillae; penial setae emerge from papillae in xviil. Clitellum annular xiii—xvili, no genital markings (Fig. 3A). Internal characteristics: Septa 6/7—13/14 muscular, greatest thickness 8/9-11/12; 13/ 14 only faintly muscular. Alimentary canal with gizzard in vi; esophagous with pebbly internal texture x—xiv, low longitudinal folds Xv; ventral esophageal ridge x—xv, valvular in XVi, intestinal origin xvii, no caecum; typhlosole simple fold originating over 20/ 21-xxv, terminates in region Ixxxiv—ci, height one-fourth lumen diameter. No cal- ciferous glands. Stomate meganephridia 2 per segment, exoic with duct entering body wall in CD, avesiculate, tubules in elongate flat coil over BC. Vascular system with ventral trunk, single dorsal trunk, these connected by lateral trunks in vi—x, esophageal hearts in xi—xiii. Lateral trunk of vi with branch to gizzard, in v trunk from dorsal vessel to gizzard. Extra-esophageal vessel present but not traceable. Supra-esophageal vessel x—x1u1. Fan-shaped ovaries composed of long strings, with funnels in xiil; paired sper- mathecae in vill, ix, each an ovoid ampulla with one or two sessile hemispherical di- verticula (Fig. 3B). Male sexual system holandric, testes and funnels free in x, xi; seminal vesicles in xi, xii, that of xii twice size of seminal vesicle in x1; vasa deferentia superficial, enter distal end of tubular prostate glands in xviii; pros- tates with slender ducts about one quarter length of glandular portion; penial setal fol- licles just anterior to ducts, penial setae bowed, 540 x 12 microns (Fig. 3C). Diagnosis. —Argilophilus with last hearts In X11, gizzard in vi, no genital markings or papillae other than those of male pores. Remarks.—Argilophilus woodi is most similar to the new species described below, and otherwise seems close to A. sierrae Mi- chaelsen, 1921 based on somatic characters. The material used to describe A. sierrae was immature, so comparisons are difficult. However, that species does have much larg- er and more ornamented penial setae, adi- verticulate spermathecae, a midventral pa- pilla in xviii, and only one pair of seminal vesicles (Michaelsen 1921). Argilophilus woodi differs from previously published species in having the diagnostic combina- tion given above. The species A. panuliris MacNab & McKey-Fender, 1959, A. mar- moratus Eisen, 1893, A. papillifer Eisen, 1893 and A. collinus Eisen, 1900 all have the last hearts in xii (except A. papillifer which also has last hearts in xiii) and gizzard in v and vi (the first) or in v only (the last three) (Eisen 1894, 1900; Gates 1941, 1962, 1977; MacNab & McKey-Fender 1959; McKey-Fender 1970). There are other dif- ferences, such as the clitellae of most of these species being saddle-shaped, and there be- ing genital markings on some part of the body in each. Argilophilus hammondi McKey-Fender, 1970 has its gizzard in v and an earlier intestinal origin than 4. woodi, a saddle-shaped clitellum, several sets of 1.0 mm Fig. 3. Argilophilus woodi: A. ventral view, B. spermatheca, C. penial seta. 630 paired genital markings, a longer typhlosole and longer penial setae (McKey-Fender 1970). Argilophilus garloughi Smith, 1937 has last hearts in xiii but also has calciferous lamellae, a later intestinal origin and the gizzard in v (Smith 1937). The species is named for Hulton B. Wood, whose interest in earthworms led to its dis- covery, and who was the first to come up with a specimen in a seemingly unlikely spot for earthworms. It is referred to as “‘Argi- lophilus I” in Wood & James (1993). Argilophilus margaritae, new species Fig. 4A—C Type material.—Holotype USNM 169801 and Paratypes USNM 169802, in canyon live oak and bay forest, Sec. 13 R12W, T2N along Forest Service Road 2N24, 2 Apr 1990, S. W. James and H. B. Wood, collectors. Additional material. —Oak-sycamore ri- parian forest, West Fork San Gabriel River, 2 Apr 1990, S. W. James and H. B. Wood, collectors. Canyon live oaks on north-facing slope of San Gabriel River valley, Forest Service Road 2N24, 2 Apr 1990, S. W James and H. B. Wood, collectors. Oak and Coul- ter Pine with grass understorey, 1670 m. elev., 5 km west of Running Springs on Cal- ifornia Hwy. 330, 23 Apr 1991, S. W. James and H. B. Wood, collectors. Ponderosa pine- oak forest with grass understorey at mile- post 33.93 on California Hwy. 138 east of Crestline, 23 Apr 1991, S. W. James and H. B. Wood, collectors. Description. —External characteristics: Dimensions 45-78 mm by 4—5 mm, width at segment xxx, generally strongly contract- ed during fixation; body cylindrical throughout, segments 124-148. Pigmenta- tion lacking. Setae ab closely paired throughout; setal formula AA:AB:BC:CD = 3.7:1:3:2.3 at xxx, DD > '2 circumference throughout. Prostomium epilobous, seg- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ments with postsetal secondary annulus vi— X, pre- and postsetal annuli x—xu. Nephri- diopores not seen, first dorsal pore 12/13- 19/20 but dorsal pores mostly merely thin spots in body wall, few actually open; sper- mathecal pores at leading edges of viii, 1x at B. Ovipores presetal median to A in xiv; male pores in xviii. Clitellum xiii—xviul, an- nular only over xili—xiv, genital marking at 9/10 broad oval intersegmental papilla in BB or genital markings lacking (Fig. 4A). Internal characteristics: Septa 6/7-11/12 muscular, greatest thickness 8/9-10/11; 12/ 13 only faintly muscular. Alimentary canal with gizzard in vi; esophagous with pebbly internal texture xi—xill, low longitudinal folds xiv-xv; sometimes a ventral esopha- geal ridge xii—xiv, valvular in xvi, intestinal origin xvii or one-half xvii, no caecum; typhlosole simple fold originating over 20/ 21-xxiv,xxv, terminates li-lvii; height one third lumen diameter or less. No calciferous glands. Stomate meganephridia 2 per seg- ment, exoic with duct entering body wall in CD, avesiculate, tubules in rounded flat coil over BC. Vascular system with ventral trunk, single dorsal trunk, these connected by lateral trunks in vi—x, esophageal hearts in xi—x1il. Lateral trunk of vi with branch to gizzard, in v trunk from dorsal vessel to gizzard. Extra-esophageal vessel from under phar- ynx, along ventral-lateral face of gizzard, ventral face of esophagous iv—xili, in xill branching out to body wall of xiii-xviu. Su- pra-esophageal vessel xi—xiii, with lateral bulges to points of attachment of hearts. Fan-shaped ovaries composed of long strings, with funnels in xiii; paired sper- mathecae in vili, ix, each an ovoid ampulla with 2-3 lobed sessile diverticulum broadly attached to spermathecal duct (Fig. 4B). Male sexual system holandric, testes and funnels free in x, xi; seminal vesicles in x1, xii, that of xii slightly larger; vasa deferentia superficial, enter distal ends of tubular pros- tate glands in xviii; prostates with slender VOLUME 107, NUMBER 4 NO ro] 3 3 1.0 mm Www 10°0 Fig. 4. Argilophilus margaritae: A. ventral view, B. spermatheca, C. penial seta. C 632 ducts about one tenth length of glandular portion or shorter; penial setae absent or setae ab of xviii not differentiated from am- bulatory setae (Fig. 4C). Diagnosis.—Argilophilus with last hearts in xill, gizzard in vi, no penial setae, one genital marking at 9/10 or none, partially annular clitellum. Remarks.—This species differs from its previously published congeners in the same ways as A. woodi, with the sometime ex- ception of the presence of a genital marking. It differs from A. woodi in having a lesser extent of annularity in the clitellum, a short- er typhlosole, no penial setae, and a pro- static duct that is shorter relative to the length of the prostate gland. The configu- ration of the spermathecal diverticula is also different. Argilophilus margaritae, under the name ‘“‘Argilophilus 2” was found in a variety of habitats and soil types, ranging from fine- to coarse-textured (Wood & James 1993). The species name is the genitive case Lat- in equivalent of the name of the author’s older daughter. In Wood & James (1993) we noted that Argilophilus species were absent from the region south of 34°N latitude, coinciding with San Gorgonio Pass. This low desert and the lowlands to the west (presently the Los Angeles metropolitan area) may have presented a barrier to the dispersal of earth- worms. Diplocardia species were found to the south of this boundary, except for the San Jacinto Peak region which appears not to harbor any native earthworms. Exotic Lum- bricidae are present there as there are in many of the sites sampled, so there is noth- ing inherently inimical to earthworms about the highlands around San Jacinto Peak. However, south of the boundary we col- lected one immature specimen of a mega- scolecid worm with a single gizzard in one segment, greatly resembling a young Argi- lophilus. Reserving judgment on that record PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON until more material can be obtained, the range of Argilophilus is now extended south to 34°N latitude. Acknowledgments This research was supported by a grant from the United States Department of Ag- riculture Forest Service to Hulton B. Wood and the author. Dr. Wood and his staff se- lected field sites and arranged cooperation with numerous Forest Service personnel and with The Nature Conservancy. Without his interest in earthworms and perseverance this research would not have been done. Jessica Malloy assisted with preliminary identifi- cation and cataloguing of collections. Literature Cited Eisen, G. 1894. On California Eudrilidae.— Memoirs of the California Academy of Science (2), 2(3): 21-62. . 1900. Researches in American Oligochaeta, with especial reference to those of the Pacific Coast and adjacent islands. — Proceedings of the California Academy of Sciences (3), 2(2):85-276. Gates, G. E. 1941. Notes on a California earthworm, Plutellus papillifer (Eisen 1893).— Proceedings of the California Academy of Sciences (4), 23(30): 443-452. 1962. On some earthworms of Eisen’s col- lection.— Proceedings of the California Acade- my of Sciences (4), 31(8):185—225. . 1977. On the correct generic name for some West Coast native earthworms, with aids for a study of the genus.— Megadrilogica 3(2):54—60. Fender, W. M., & D. McKey-Fender. 1990. Oligo- chaeta: Megascolecidae and other earthworms from western North America. Pp. 357-378 in D. L. Dindal, ed., Soil biology guide. John Wiley and Sons, New York. MacNab, J. A., & D. McKey-Fender. 1959. A new species of Plutellus from western Oregon (Oli- gochaeta: Megascolecidae).— Northwest Sci- ence 33:69-75. McKey-Fender, D. 1970. Description of endemic earthworms from eastern Oregon (Oligochaeta: Acanthodrilidae). — Northwest Science 44:235- 244. VOLUME 107, NUMBER 4 633 Michaelsen, W. 1921. Neue und wenig bekannte Oli- woodland, and riparian zones in southern Cal- gochaten aus skandinavischen Sammlungen. — ifornia. General Technical Report PSW-GTR- Arkiv for Zoologi Stockholm 13:1-25. 142. Albany, California, Pacific Southwest Re- Smith, F. 1937. New North American species of search Station, Forest Service, USDA, 20 p. earthworms of the family Megascolecidae.— Proceedings of the United States National Mu- : aati seu 84(2174):157-181. Department of Biology, Maharishi Inter- Wood, H. B., & S. W. James. 1993. Native and in- national University, 1000 N. Fourth Street, troduced earthworms from selected chaparral, Fairfield, Iowa 52557-1056, U.S.A. PROC. BIOL. SOC. WASH. 107(4), 1994, pp. 634-640 PEOSIDRILOIDES, A NEW GENUS, AND NEW RECORDS OF PEOSIDRILUS (OLIGOCHAETA: TUBIFICIDAE) FROM THE UNITED STATES, WITH THE DESCRIPTION OF A NEW SPECIES FROM THE GULF OF MEXICO Christer Erséus and Michael R. Milligan Abstract.— Peosidrilus biprostatus Baker & Erséus, 1979, is recorded from the Gulf of Mexico, P. coeloprostatus (Cook, 1969) from Maine, and both P. biprostatus and P. acochlearis (Ers¢us & Loden, 1981) are reported from bathyal depths (off North Carolina) for the first time. Peosidriloides, new genus, does not have lateral spermathecal pores, the feature regarded as an autapomorphy of Peosidrilus Baker & Erséus, 1979. Peosidriloides hornensis, new species, from the northern Gulf of Mexico, shares some apomorphic features (a short clitel- lum, long vasa deferentia) with Peosidriloides flabellifer (Erséus, 1984), new combination, but is distinguished from that species by its more numerous anterior setae, its more ventral spermathecal pores, its wider vasa deferentia, and its possession of penial organs. The marine tubificid genus Peosidrilus was established by Baker & Erséus (1979) to ac- commodate one species, P. biprostatus Ba- ker & Erséus, 1979, from the east coast of the United States (New Jersey and North Carolina). Although this species was trans- ferred to Phallodrilus Pierantoni, 1902 ina subsequent paper by Erséus (1984), Peosi- drilus was resurrected when the paraphy- letic taxon Phallodrilus was split into a number of genera (Erséus 1992). Peosidrilus was then recognized by the assumedly au- tapomorphic, lateral, position of the sper- mathecal pores, and was enlarged to include fourteen species; however, one species (P. flabellifer Erséus, 1984) with spermathecal pores located in line with the ventral setae was included in this genus too. Many species of Peosidrilus are recorded from shallow-water or continental shelf bot- toms of the northwestern part of the Atlan- tic Ocean; one, P. simplidentatus (Erséus, 1979a), is a High Arctic deep-sea form. An Atlantic origin of the genus has therefore been suggested (Erséus 1992). The present paper presents geographic as well as bathymetric range extensions of three North Atlantic species. Peosidrilus bipros- tatus and P. acochlearis (Erséus & Loden, 1981) are recorded from the continental slope off North Carolina, P. biprostatus also from the Gulf of Mexico, and P. coelopros- tatus (Cook, 1969) is reported from as far north as Maine. Moreover, a new species, described from the northern Gulf of Mex- ico, as well as P. flabellifer, are attributed to a new genus. The new material studied originates from three sources: (1) the Study of Biological Resources on the U.S. South Atlantic Con- tinental Slope and Rise (Blake et al. 1987), supported by contract number 14-12-0001- 30064 from the U.S. Department of Inte- rior, Minerals Management Service, to Bat- telle New England Research Laboratory, Woods Hole Oceanographic Institution, and Lamont-Doherty Geological Observatory (courtesy Dr. Nancy Maciolek-Blake), (2) oligochaetes collected at Perdido Key, near Pensacola, northwestern Florida, and Horn VOLUME 107, NUMBER 4 Island, off the coast of Mississippi, by Dr. J. McLelland, Gulf Coast Research Labo- ratory, Ocean Springs, Mississippi, and (3) material from a study of trophic coupling and benthos in Sheepscot River, Maine (Dr. L. Watling and Dr. R. Langton, principal investigators; courtesy also Ms. L. McCann; University of Maine, Darling Marine Cen- ter). The specimens were stained in paracar- mine and mounted whole in Canada bal- sam. Material of the species is deposited in the United States Museum of Natural His- tory (USNM), Smithsonian Institution, Washington, D.C., the Swedish Museum of Natural History (SMNH), Stockholm, and in the reference collection of Darling Marine Center. Peosidrilus Baker & Erséus, 1979 Peosidrilus Baker & Erséus, 1979:505- 506.—(In part) Erseus 1992:27. Adelodrilus Cook (in part).—Erséus & Lod- en 1981:823. Phallodrilus Pierantoni (in part); Erséus 1979a:203.—Erséus 1984:823.—Erséus 1990:54. Diagnosis (modified after Erséus 1992) (assumed autapomorphy in italics).—Ma- rine tubificids. Somatic setae bifid, with up- per tooth thinner and shorter than lower. Penial setae, when present, generally with clubbed ectal ends, each bearing an apical hook; however, in some species the ‘clubs’ are indistinct, or even absent. Penial setae small, generally straight, and numerous in bundle. Spermathecal pores lateral, or in line with dorsal setae [P. dorsospermathe- ca]. Vasa deferentia ciliated, narrow in all but one species [vasa much dilated in P. acochlearis|, entering apical ends of atria. Atria generally somewhat elongate, more or less horizontal and curved; occasionally short and almost erect. Each atrium with two prostate glands, anterior one attached at entrance of vas deferens, posterior one attached to ectal end of atrium. Atria either 635 opening directly to exterior through simple pores, small male projections, or true pen- dent penes; copulatory sacs absent or pres- ent. Spermathecae of varying shape; sper- mathecal vestibules present in a few species. Type species. — Peosidrilus biprostatus Ba- ker & Erséus, 1979. Other species. —Twelve species listed by Erséus (1992); i.e., excluding P. flabellifer, which is transferred to Peosidriloides, new genus, below. Remarks. — Peosidrilus was revised by Er- séus (1992) to include a number of species formerly regarded as members of Phallodri- lus, primarily those species that have clubbed, apically hooked penial setae, and lateral spermathecal pores, the latter feature considered as a possible autapomorphy of Peosidrilus. However, the genus is hetero- geneous. Due to the lack of distinct syna- pomorphies, other species have been in- cluded on the basis of overall (partly symplesiomorphic) similarity with the more typical members. With the removal of Peo- sidriloides flabellifer, new combination, however, Peosidrilus is now more unam- biguously diagnosed by its lateral sperma- thecal pores. The dorsal position of the pores in P. dorsospermatheca can be regarded as a secondary transformation from the lateral position. Peosidrilus may have to be revised again when new information becomes available. Other taxa with clubbed, apically hooked penial setae (Adelodrilus Cook, 1969, Ber- mudrilus Erséus, 1979b) will then also have to be considered (see Erséus 1992). Peosidrilus biprostatus Baker & Erséus, 1979 Peosidrilus biprostatus Baker & Erséus, 1979: 506-508, figs. 1-2.—Erséus & Loden 1981:819-820.—Erséus 1992:27, fig. 12F. Phallodrilus biprostatus; Erséus 1984:819- 820.—Erseus 1986:296-297.—Davis 1985:table 1. New material.—USNM 169815, 1 spec- 636 imen from off Cape Fear, North Carolina, U.S.A., 32°32.22'N, 77°15.31’'W, 605 m, sand, South Atlantic cruise 5, station 14A, box core 3 (20 Sep 1985; see Blake et al. 1987). USNM 169816, 1 specimen from 800 m off Perdido Key (Gulf of Mexico), near Pensacola, Florida, U.S.A., about 6 m, sand, station A 0920C89C (collected by J. Mc- Lelland, October 1989). Remarks.—The specimen from 605 m depth off North Carolina represents the deepest record of this species; P. biprostatus has never been found deeper than 73 m be- fore (Baker & Erseus 1979). The worm is not complete. It is 3.5 mm long, comprising anterior 21 segments only. Its penial setae are about 9 per bundle. The Perdido Key material provides the first record of P. biprostatus from the Gulf of Mexico. The single worm is the smallest (complete, and sexually mature) individual of the species reported to date; it is 4.5 mm long, with about 38 segments. The penial setae appear to be 6 on one side of worm, whereas the penial setae and male efferent duct are not developed on the other side. Distribution and habitat.—NW Florida (first record for Gulf of Mexico), eastern United States (Florida through Massachu- setts). Largely coarse sand, known from 5.5— 605 m depth. Peosidrilus acochlearis (Erséus & Loden, 1981) Adelodrilus acochlearis Erséus & Loden, 1981:821-823, figs. 1B—C, 2.—Erséus 1983:77-78. Phallodrilus acochlearis,; Erséus 1986:297-— 298, fig. 7. Peosidrilus acochlearis; Erséus 1992:27. New material. —USNM 169817-169818, 2 specimens from off Cape Fear, North Car- olina, U.S.A., 32°32.22'N, 77°15.31'W, 605 m, sand, South Atlantic cruise 5, station 14A, box core 3 (20 Sep 1985; see Blake et al. 1987). Remarks. —One of the two specimens is PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON complete: 3.6 mm long, with 36 segments. This specimen has a few modified bifid se- tae, with much prolonged lower teeth, in some dorsal bundles of the posteriormost segments (see Erseus & Loden 198 1:fig. 1C). The new material conforms to the previous descriptions in other characters too, but one individual appears to have at least 15 penial setae per bundle. Previously described ma- terial had maximally 14 such setae per bun- dle (Erséus 1986). This species had not been collected deep- er than 11 m before. The present record is from a continental slope station. Distribution and habitat.—East coast of the United States (Florida through North Carolina). Sand, known from 5.5-—605 m depth. Peosidrilus coeloprostatus (Cook, 1969) Phallodrilus coeloprostatus Cook, 1969:16— 17, fig. 5.—Erséus 1979a:189-190, fig. 4.—Erséus 1984:813-815, fig. 1. Peosidrilus coeloprostatus; Erséus 1992:27. New material.—USNM 169819-169823 (5 specimens) and Darling Marine Center reference collection (3 specimens), all from Outer Sheepscot Bay, Maine, 43°43.10’N, 69°43.40’W, Station B7, 37 m, coarse sand and gravel with a large amount of vascular plant detritus, annual salinity range 32.4— 33.2%o0 (collected by L. Watling, 8 Sep 1988). SMNH Main coll. 1414, 4 specimens from same area, kind of sediment, collector and date, but 43°42.85’N, 69°43.65’W (Station B9), 38 m. Remarks.—The new material from Sheepscot River conforms well to the pre- vious descriptions. The specimens are 3.9— 7.2 mm long, with 38-78 segments. Their penial setae are about 35-50 um long, (8)9- 15(16) per bundle. This species has been reported from as far south as Maryland/Delaware (Diaz et al. 1987), but not north of Massachusetts (Cook 1969) before. VOLUME 107, NUMBER 4 Distribution and habitat.—East coast of United States (Maryland through Maine; new record for Maine). Sand, known from 3.4-78 m depth. Peosidriloides, new genus Phallodrilus Pierantoni (in part); Erséus 1984:823. Peosidrilus Baker & Erséus (in part); Erséus 1992:27. Etymology. —Named for its resemblance (-oides Greek for ‘resembling’) to Peosidri- lus. The type species was previously clas- sified as a member of that genus. Diagnosis. —(assumed autapomorphies in italics). — Marine tubificids. Somatic setae bifid. Clitellum short, maximally extending over posterior third of segment X, whole XT, and anterior two thirds of XII. Somatic setae bifid, with upper tooth thinner and shorter than lower. Penial setae with single-pointed, curved tips (tips also somewhat clubbed in P. flabellifer). Penial setae small, numerous, densely packed, generally over 10 per bun- dle. Spermathecal pores in line with ventral setae, or even ventral to this line. Vasa de- ferentia ciliated, narrow, several times lon- ger than atria, entering apical ends of atria. Atria cylindrical or somewhat spindle- shaped, horizontal, but slightly curved to- wards male pores; latter simple (P. flabel- lifer) or as penis-like organs (P. hornensis). Each atrium with two prostate glands, an- terior one attached at entrance of vas def- erens, posterior one attached to ectal end of atrium. Spermathecae with discrete ducts and thin-walled ampullae; spermathecal vestibules not distinct. Type species. —Phallodrilus flabellifer Er- seus, 1984. Other species.—Peosidriloides hornensis, new species. Remarks. —The inclusion of Phallodrilus flabellifer in Peosidrilus was problematic as this species does not have lateral sperma- thecal pores, which is the assumed autapo- 637 morphy of Peosidrilus (see Erséus 1992; and above). With regard to the (ventral) location of the spermathecal pores, the new genus Peosidriloides is plesiomorphic. The penial setae of Peosidriloides hornen- sis and P. flabellifer have tips that are either unmodified (Erséus 1992: fig. 1D, state 0), or with indistinct ectal swellings (Erséus 1992: fig. 1D, state 3), both relatively ple- siomorphic conditions vis-a-vis the dis- tinctly clubbed penial setae (Erséus 1992: fig. 1D, state 4) found in most species of Peosidrilus, Adelodrilus and Bermudrilus. The new genus therefore may be phyloge- netically separated from these other genera. Monophyly of Peosidriloides is supported by the unusually short clitellum, and the very long vasa deferentia. Both species of Peosidriloides are from the Northwest Atlantic. Peosidriloides flabellifer (Erséus, 1984), new combination Phallodrilus flabellifer Erséus, 1984:818- 819, fig. 4. Peosidrilus flabellifer; Erseus 1992:27. Remarks. — As argued above, this species is better placed outside Peosidrilus. The dis- tinguishing features of P. flabellifer and P. hornensis are noted in Remarks for the lat- ter below. Distribution and habitat. —Georges Bank, off Massachusetts (Northwest Atlantic). Coarse sand, 78-79 m depth. Peosidriloides hornensis, new species Fig. | Holotype. -USNM mounted specimen. Type locality. —N shore of about the mid- dle of Horn Island, about 1 1 km off the coast of the state of Mississippi, northern Gulf of Mexico, swash and supratidal zones in an area severely impacted by an oil spill, Sta- tion no. 11 (collected by J. McLelland, 21 Sep 1989). 169824, whole- 638 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. [ 100 um Peosidriloides hornensis, new species. A, Free-hand drawing of anterior somatic setae; B, Free-hand drawing of postclitellar somatic seta; C, Free-hand drawing of penial seta; D, Male genitalia in segment XI; E, Spermathecae in segment X (one spermatheca slightly damaged). Abbreviations: a, atrium; pr1, anterior prostate gland; pr2, posterior prostate gland; ps, penial seta; vd, vas deferens. Paratypes. —USNM 169825 and SMNH Type coll. 4632, 2, whole-mounted speci- mens from type locality. Etymology. —Named for Horn Island. Description. —Length of two USNM specimens, 8.2 and 6.1 mm, 59 and 60 seg- ments respectively (SMNH specimen not complete). Width at clitellum (fixed, com- pressed specimens) 0.32—0.39 mm. Prosto- mium rounded, well set off from peristo- mium. Clitellum extending over 73X—XII, well developed in all specimens. Somatic setae (Fig. 1A—B) bifid, with upper tooth shorter and distinctly thinner than lower, and with subdental ligaments. These setae generally 35-50 um long, 2.5—3.5 wm thick (smaller in a few anteriormost segments), two or three (occasionally four) per bundle anteriorly, two per bundle in postclitellar segments. Penial setae (Fig. 1C; D, ps) sin- gle-pointed, 25-35 um long, 1—2 um thick, about 7-12 per bundle (exact numbers dif- ficult to establish), with curved, thin tips (details not clear); setae densely packed within bundle. Male and spermathecal pores paired, both pairs ventral to lines of ventral setae; pores thus rather close to each other within each pair (see Fig. 1E). Male pores posteriorly in segment XI, spermathecal pores in most anterior part of X. Pharyngeal glands in segments IV-V. Male genitalia (Fig. 1D) paired. Vas defer- ens variably wide (ectal part dilated, up to about 15-20 um wide), coiled, several times longer than atrium, appear to enter apical end of latter (details not clear). Atrium cy- lindrical or somewhat spindle-shaped, slightly curved, 45-60 um long, 20-22 wm wide, with very thin outer (muscle) lining, and granulated and ciliated inner epitheli- um. Atrium terminating in simple copula- tory organ; probably a penis, but difficult to see whether organ is fully pendent within a penial sac. Copulatory organ 23-28 um long, 23-28 um wide. Two small, compact pros- tate glands present; anterior one attached near junction between vas deferens and atri- um, posterior one located near penial organ. VOLUME 107, NUMBER 4 Spermathecae (Fig. 1E) totally about 135- 160 long, with slender duct-like, at middle somewhat constricted, part, and pear- shaped, thin-walled ampullae; latter 55—70 um wide, filled with non-organized sperm. Typical spermathecal vestibules not pres- ent, but outer (Somewhat swollen) parts of ducts possibly homologous to such struc- tures. Remarks. — This new species differs from Peosidriloides flabellifer in several aspects. It has up to three or four setae in some anterior segments (setae two per bundle throughout body in P. flabellifer), its sper- mathecal pores are closer together ventrally (pores in line with ventral setae in P. fla- bellifer), its vasa deferentia appear dilated (vasa narrow throughout in P. flabellifer), and it has penis-like copulatory organs (atria opening directly to the exterior through sim- ple pores in P. flabellifer). With regard to the dilation of the vasa deferentia and the possession of penial or- gans, P. hornensis is similar to Peosidrilus acochlearis (Erséus & Loden, 1981), but the latter taxon is in other respects (with nu- merous somatic setae, distinctly clubbed penial setae, clitellum and vasa deferentia of normal length, and lateral spermathecal pores) a typical member of Peosidrilus. The resemblance may therefore be due to con- vergence. Distribution and habitat.—Known only from Horn Island (northern Gulf of Mexi- co). Inter- and supratidal sand. Discussion Peosidrilus and Peosidriloides are both taxa with a more or less Northwest Atlantic distribution, at least by conclusion from the present records. Their general appearance seems to indicate phylogenetic membership in a larger group of phallodriline genera ex- tending across the North Atlantic to Europe and the Mediterranean and Black Seas; this larger group contains also Adelodrilus and Bermudrilus. The character patterns within 639 this larger group are, however, confusing, and it is probable that there is homoplasy (both convergence and reversal) in the lo- cation of spermathecal pores as well as in the morphology of penial setae (Erséus 1992). Twelve of the thirteen species of Peosi- drilus are known only from the east coast of the United States, including the Gulf of Mexico, and from the Caribbean (Erséus 1992; present paper). The genus thus ap- pears to have undergone a unique radiation in this part of the Northwest Atlantic. All previous records of Peosidrilus have been from either intertidal or continental shelf habitats. The present records of P. bi- prostatus and P. acochlearis from a station 605 m deep indicate that the genus also oc- curs at bathyal depths. Acknowledgments We are grateful to N. Maciolek-Blake (formerly at Battelle New England Marine Research Laboratory), J. McLelland (Gulf Coast Research Laboratory), L. Watling and L. McCann (Darling Marine Center), for providing the material used in this study; to Barbro Lofnertz (University of Gote- borg), and Christine Hammar (Swedish Museum of Natural History), for technical assistance; and the Swedish Natural Science Research Council, for financial support. Literature Cited Baker, H. R., & C. Erséus. 1979. Peosidrilus bipros- tatus n.g., N.sp., a Marine tubificid (Oligochaeta) from the eastern United States. — Proceedings of the Biological Society of Washington 92:505- 509. Blake, J. A. et al. 1987. Study of biological processes on the U.S. South Atlantic slope and rise. Phase 2. Report No. MMS 86-0096. U.S. Department of the Interior, Minerals Management Service, Washington, D.C., 414 pp. Cook, D. G. 1969. The Tubificidae (Annelida, Oli- gochaeta) of Cape Cod Bay with a taxonomic revision of the genera Phallodrilus Pierantoni, 1902, Limnodriloides Pierantoni, 1903 and 640 Spiridion Knollner, 1935.—Biological Bulletin 136:9-27. Davis, D. 1985. The Oligochaeta of Georges Bank (NW Atlantic).— Proceedings of the Biological Society of Washington 98:158-176. Diaz, R. J., C. Erséus, & D. F. Boesch. 1987. Dis- tribution and ecology of Middle Atlantic Bight Oligochaeta.— Hydrobiologia 155:215-225. Erséus, C. 1979a. Taxonomic revision of the marine genus Phallodrilus Pierantoni (Oligochaeta, Tu- bificidae), with descriptions of thirteen new spe- cies.— Zoologica Scripta 8:187—208. . 1979b. 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. 1983. New records of Adelodrilus (Oligo- chaeta, Tubificidae), with descriptions of two new species from the Northwest Atlantic. — Hy- drobiologia 106:73-83. 1984. 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. 1986. Marine Tubificidae (Oligochaeta) at Hutchinson Island, Florida.— Proceedings of the Biological Society of Washington 99:286-315. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON . 1990. The marine Tubificidae and Naididae (Oligochaeta) of south-western Australia. Pp. 43— 88 in F. E. Wells, D. I. Walker, H. Kirkman, & R. Lethbridge, eds., Proceedings of the Third International Marine Biological Workshop: the marine flora and fauna of Albany, Western Aus- tralia, volume 1. Western Australian Museum, Perth, 437 pp. . 1992. A generic revision of the Phallodrilinae (Oligochaeta, Tubificidae).— Zoologica Scripta 21:5-48. —, & 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 Sur- vey of Washington 94:819-825. Pierantoni, U. 1902. Due nuovi generi di Oligocheti Marini rinvenuti nel Golfo di Napoli.—Bollet- tino della Societa di Naturalistii Napoli 16:113- 117. (CE) Department of Invertebrate Zoolo- gy, Swedish Museum of Natural History, Box 50007, S-104 05 Stockholm, Sweden; (MRM) Center for Systematics and Tax- onomy, P.O. Box 37534, Sarasota, Florida 34278, U.S.A. PROC. BIOL. SOC. WASH. 107(4), 1994, pp. 641-656 A NEW AMERICAN FAIRY SHRIMP, LINDERIELLA SANTAROSAE (CRUSTACEA: ANOSTRACA: LINDERIELLIDAE), FROM VERNAL POOLS OF CALIFORNIA, U.S.A. Alain Thiéry and Michael Fugate Abstract. — A new species of fairy shrimp, Linderiella santarosae (Anostraca: Linderiellidae), is described from vernal pools on the Santa Rosa Plateau Re- serve, Riverside Co., California, U.S.A. The form of the basal outgrowth on antenna 2 of males and the spines on the resting eggs are the primary characters which distinguish L. santarosae from the other Linderiella species of western North America, northwestern Africa and southwestern Europe. Linderiella santarosae is also distinguished from L. occidentalis, the other Californian species, by fixed differences at two electrophoretic loci (PGM and PEP-D). The distribution of the five known species, each locally endemic to Mediterranean climatic regions, may be linked to a wider distribution of the genus in the past across the Laurasian continent. Brtek (1964) erected the family Linderi- ellidae and the genus Linderiella based on an enigmatic anostracan (Linderiella occi- dentalis Dodds, 1923) from a lake (Lagun- ita) on the Stanford University Campus in Palo Alto, Santa Clara Co., California, U.S.A. Dodds (1923) originally placed L. occidentalis in the genus Branchinecta, and Linder (1941), in his review of the Anos- traca, transferred it to the family Chiroce- phalidae and the genus Pristicephalus. Brtek (1964) separated L. occidentalis from the Chirocephalidae based on differences in the male reproductive structures. The genus Linderiella is currently restrict- ed to regions with a Mediterranean climate in the Northern Hemisphere. Species are found in Morocco (Linderiella africana Thiéry, 1986), France (Linderiella massa- liensis Thiéry & Champeau, 1988), Spain (Alonso 1985) and California, U.S.A. (L. occidentalis Dodds, 1923). The genus in- habits pools and ponds with low mineral- ization that fill after winter rains (Alonso 1985, Thiéry & Champeau 1988, Eng et al. 1990). During fieldwork in California from 1988 to 1991, one of us (MF) collected two spe- cies of anostracans from several vernal ponds and pools within the Nature Conser- vancy Reserve on the Santa Rosa Plateau, Riverside Co., California. One of the species was Branchinecta lynchi Eng et al., 1990, and the other was the new species in the genus Linderiella described in this paper. Methods Specimens were fixed in 10% formalin and preserved in 70% ethanol. Antennae, tho- racic appendages and penes to be illustrated were removed under a WILD M7 stereo- microscope and observed with a WILD M20 compound microscope, both equipped with camera lucida. Measurements are given to the nearest 0.1 mm for the adults and to the nearest 2 mm for the diameter of the resting eggs. Total length includes the distal setae of the cercopods. Specimens observed on the scanning electron microscope (SEM) were dehydrated through a graded ethanol series to absolute ethanol, critical-point 642 N Gir /} t 1 / / ‘ J T y Sa ew \ ‘ y ‘ a 1 1 1 a oS 1 —_— Ui eo — 7 I 1 me q i ‘ San Francisco I pe Oo x 5 PACIFIC ® \ i 2 ‘ 24 1 : Es 1 OCEA \ i ay ‘ eee Q Pas Sty 4 \ i > DS i V Los Angeles ¥ (] Fig. 1. California collection sites for populations of Linderiella used in this study. Open star indicates the type locality of Linderiella santarosae, n. sp.; solid star, the type locality of Linderiella occidentalis (Palo Alto); solid circle, Fields Rd. and Tim Bell Rd.; solid square, Prairie City OHV; open circle, sites sampled by Eng et al. (1990), open triangles, DB. 430 population and solid triangle, Marin Co. (DB. 878). dried and coated with gold-palladium for observation in a Cambridge Stereoscan 360. Specimens of related species used for mor- phological comparisons are from the per- sonal collections of A. Thiéry (L. africana: Daya Azigza, 6 Mar 1985, Middle Atlas, Morocco and L. massaliensis: pond of Saint Maximin, 7 Feb 1988, France) and D. Belk (L. occidentalis: California, DB, 430). Ad- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ditional specimens of L. occidentalis either were collected in the field or lab-reared from dry mud (see locations on Fig. 1): Lagunita, Stanford University, Palo Alto, type locality of L. occidentalis (soil sample collected Nov 1989, C. Sassaman): 4 6, length, 8.8-10.9 mm, 3 2(1 ovigerous), length, 9.5—10.0 mm; Tim Bell Road (12 Mar 1988, coll. S. Mor- ey): 5 6, length, 9.7-11.2 mm, 4 2 (2 ovi- gerous), length, 11.0-13.3 mm; Fields Road (12 Mar 1988, coll. S. Morey): 7 4, length, 9.7-10.6 mm; 7 2 (5 ovigerous), length, 9.6— 12.0 mm; Prairie City OHV. (2 Feb 1990 coll. M. Fugate): 21 4, length, 10.0—-13.2 mm, 25 2 (20 ovigerous), length, 12.6—18.0 mm, and Tehama Co. DB. 430 (28 Mar 1982): 8 4, length, 10.1-12.4 mm, 9 2 (9 ovigerous), length, 10.2-14.6 mm. Allele frequencies were determined using starch gel electrophoresis (see Fugate 1992 for detailed methods) for three populations (Fields Road, 14 individuals; Tim Bell Road, 6 individuals; Lagunita, 10 individuals) of Linderiella occidentalis, one population (Mesa de Burro, 14 individuals) of Linderi- ella santarosae and one population of (7 individuals from Connecticut Valley Bio- logical Supply) Eubranchipus vernalis (Ver- rill, 1869) at the following eight loci: malic enzyme (ME), phosphoglucomutase (PGM), glucose-6-phosphate isomerase (PGI), phenylalanyl-proline peptidase (PEP-D), leucyl-alanine peptidase (PEP-C), Glucose- 6-phosphate dehydrogenase (G6PD), iso- citrate dehydrogenase (IDH), and aspartate aminotransferase (AAT). Nei’s pairwise ge- netic distances (D) were calculated from the allele frequency data and clustered with the UPGMA method using PHYLIP 3.4 (Fel- senstein 1989). Nei’s genetic distance (D) is an estimate of the number of allelic substi- tutions per locus between two populations (Nei 1987 chap. 9). Linderiella santarosae, new species Figs. 2-10 Material examined. — Holotype 6, USNM 266798, allotype 2, USNM 266797 and 20 VOLUME 107, NUMBER 4 643 Fig. 2. Linderiella santarosae, n. sp.: habitus male, left lateral view (scale in mm). paratypes, USNM 266796, Mesa de Burro; 18 paratypes, USNM 266795, Mesa de Col- orado; 4 paratypes, Mesa de Colorado, Mu- seum National d’Histoire Naturelle, Paris (MNHN Bp. 549). Additional specimens, fixed 1 August 1989 from Mesa de Colo- rado, also were used for the description. These individuals were lab-reared from soil samples collected 3 May 1988. Type locality. — Vernal pools on the Santa Rosa Plateau Reserve, Riverside Co., Cal- ifornia (Fig. 1). The reserve, located 6.5 km southwest of Interstate 15 on Clinton Keith Road (33°32'N, 117°17'W), is bordered on the northwest by the Santa Ana Mountains and on the east by agricultural and urban development surrounding the cities of Mur- rieta and Temecula. Etymology. —The species is named after the type locality, the Santa Rosa Plateau, Riverside Co., California. Male.—Length, 10.1 mm; to telson, 9.0 mm. Habitus (Fig. 2). Lengths of males ex- amined 6.4—10.2 mm. Antenna 2 in form of claspers without frontal process. Distal segment of antenna 2 slender, slightly curved, with row of transverse ridges on inner surface (Fig. 6a, f, h). Tip of segment slightly inflated and bent medially (Fig. 6a, g). Basal portion of row slightly sinuous, contiguous to external edge on distal half of segment (Fig. 10h), not reaching tip. Basal segment of antenna 2 stout with dorsome- dially projecting basomedial outgrowth (Figs. 3A, 5F, 6a). Outgrowth with slender, laterally curving tip and spinose plateau on inner face (Figs. 3B, C, 6a, 7a, f). Spines on plateau smooth and slightly recurved (Fig. 7a, f). Maxilla 2 of common type with single stiff seta on tip and 2-3 soft setae toward base (Cannon & Leak 1933, Linder 1941). Preepipodites of thoracic appendages ser- rated (Fig. 3E). Endopodite rounded with small spines at the base of setulose setae (Fig. 3D). Penes rectilinear, parallel. Basal part non-retractile, square in cross section, reaching end of third abdominal segment, with slightly curved medially-directed spur ornamented with 3-5 curved spines on in- ner side (Figs. 3F, 8a, c). Eversible tip of penes with smoothly tapered spine (Fig. 8b). Abdomen with dorsal and ventral setae 60- 70 mm long (Fig. 8g, h). Cercopods with fine setae on surface and setulated setae along edges (Fig. 8e, f). Female. —Length 11.1 mm, 10.0 mm to telson, ovigerous. Lengths of females ex- amined 7.2-11.3 mm. Antenna 2 with straight horn-like process originating from inner side of basal segment (Fig. 4A, B). Process with thick base, denticulate on dis- tal half (Fig. 4B). Anterior surface of basal segment hirsute, bearing irregular row of se- tae at base of the inner process (Fig. 4A, B). Distal segment of antenna 2 small, tapering to point. Brood pouch globular, 1.2 times wider than long, extending to abdominal segment 3. Pore opens subdistally and ven- trally (Fig. 4C, D). Resting eggs (=cysts) spherical, with numerous short, tulip-shaped spines (Figs. 9a—i, 10a, c); diameter 230- 283 um, mean diameter, 260.7 + 13.8 um, 644 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 3. Linderiella santarosae, n. sp. male: (A) head frontal view; (B) & (C) detail of the basomedial outgrowth; (D) shape of the endopodite of the appendage 7 with an enlarged detail; (E) shape of the preepipodites of the appendage 7; (F) penes in ventral view, tip not everted, with detail of the spurs of the basal nonretractile part (scales in ym, except for (A) in mm). VOLUME 107, NUMBER 4 200 j 645 Fig. 4. Linderiella santarosae, n. sp. female: (A) & (B) second antenna dorsal view; (C) brood pouch ventral view; (D) brood pouch left lateral view (scales (A) & (B) in mm & (C) (D) in um). n = 35 from three mature females. Spines, 18-23 wm long, with flat tops, 5-16 wm wide (Fig. 9d-i). Most tops open with no pore visible at base of spines. Irregularly, spines fused at bases in twos or threes, mak- ing short walls not reaching more than half of total length of spine (Fig. 9a, b). Spinose outer layer surrounding spongy inner layer (Fig. 10d). One of largest mature females (8.2 mm) with 14 eggs in brood pouch. Females larger than males through same cohort as previously observed for other Lin- deriella species (Dodds 1923, Heath 1924, Thiery 1986a, Thiery & Champeau 1988). Living fairy shrimps white, sometimes pale green or blue, quite translucent, with dark red eyes and reddish orange cercopods. Brood pouch of mature female with white shell glands and gold yellow to dark bronze resting eggs. Remarks. —The five species in the genus Linderiella are distinguished from one an- other by only minor morphological differ- ences (Thiéry & Champeau 1988). The un- described Spanish Linderiella was initially identified as L. occidentalis (Alonso 1985) and Linderiella santarosae was as well in a recent survey of Californian anostracans (Eng et al. 1990). The form of the male an- tenna 2 and the resting egg are the two most reliable characters for distinguishing L. san- tarosae and L. occidentalis (Figs. 5-10), but minor differences are also found in the form of the medial spur on the penis (Fig. 8a—d). The row of transverse ridges on the distal segment of the male antenna 2 of L. occi- dentalis is more sinuous and the tip of the segment is neither sharply bent nor slightly inflated (Figs. 6 & 10h, i). The paired ba- somedial outgrowths on the basal segment 646 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 5. Comparison of basomedial outgrowth of basal segment of male second antenna between Linderiella santarosae, n. sp. and different populations of Linderiella occidentalis (Dodds): L. occidentalis, (a) Tim Bell Rd.; (b) Olsen Rd.; (c) Lagunita Palo Alto; (d) Prairie City OHV; (e) DB. 430; Linderiella santarosae, n. sp.; (f) Santa Rosa Plateau, type locality (scale in mm). of the male antennae 2 of L. occidentalis are short, tulip-shaped (less than 2% of the similar in form to those of L. santarosae, spines of L. occidentalis eggs have a flat top) but lack the slender, outward curving tips and the spines are longer in L. occidentalis (Figs. 5, 6a—c, & 7). The resting eggs of L. (27-33 wm) than in L. santarosae (18-23 occidentalis have acute spines instead of mm). VOLUME 107, NUMBER 4 Fig. 6. SEM-micrographs in male second antenna. (a) L. santarosae, n. sp., lateral view; (b) inner view of L. occidentalis (DB. 430); (c) inner view of L. occidentalis (Prairie City); (d) inner view of L. africana (Daya Azigza, 6 March 1985, Middle Atlas Morocco); (e) inner view of L. massaliensis (pond of St. Maximin, 7 February 1988, France); (f) L. santarosae, n. sp., detail of row of transverse ridges, medial part of second segment, lateral view; (g) L. santarosae, n. sp., inner view of second segment; (h) L. santarosae, n. sp., detail of row of transverse ridges, apical view (scales in um). 648 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 7. SEM-micrographs of basomedial outgrowth on inner side of second antennae of males (inner views). (a) L. santarosae, n. sp.; (b) L. africana; (c) L. massaliensis; (d) L. occidentalis (DB. 430); (e) L. occidentalis (Prairie City); (f) L. santarosae, n. sp. (scales in um). Arrows on (a), (d), (e), (f) indicate the straith or curved tip of the basomedial outgrowth. Linderiella africana and L. massaliensis (the Spanish Linderiella is still undescribed) also differ from L. santarosae in the form of the basomedial outgrowth on antenna 2 of the male (Fig. 7) and the spines of the resting eggs (Mura & Thiéry 1986, Thiéry & Champeau 1988). The resting eggs of the Spanish Linderiella resemble those of L. Fig. 8. SEM-micrographs of genital and abdominal segments of males. Linderiella Santarosae, n. sp. (a—c) & (e-h); (a) ventral view of genital segments of abdomen, nonretractile structures and invaginate penes; (b) idem, with devaginate penes; (c) detail of nonretractile structure, with teeth on inner side (arrow); Linderiella occidentalis (DB. 430): (d) ventral view of genital segments of abdomen, with nonretractile parts and invaginate penes (arrow indicates curved inner edge); Linderiella santarosae, n. sp.; (e) ventral view of telson and cercopods; (f) detail of cercopods with hirsute surface (arrow); (g) ventral view of abdomen with setae (arrows); (h) detail of an abdominal seta (scales in um). 650 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 9. Linderiella santarosae, n. sp. SEM-micrographs of resting eggs, (a) egg from single female, collected 1 January 1989, (b) & (c) eggs from two different females, collected 1 January 1988; (d) detail of spines, lateral view; (e) detail of spines, apical view; (f) idem, lateral view; (g) detail of a characteristic flat tip, lateral view; (h) detail of a flat tip in apical view; (i) idem, with opened spine (scales in um). VOLUME 107, NUMBER 4 651 Fig. 10. Linderiella santarosae, n. sp. (a), (c), (d), (h) and L. occidentalis (b) & (e)—Lagunita. Palo Alto, (f)— Prairie City OHV, (g)—DB. 430; (a) whole egg entangled with mucus; (b) whole egg, with some broken spines (arrows)—(a) & (b) same scale; (c) lateral view of flat top spines; (d) detail of spongy inner layer of egg; (e-g), three detail of acute spines of L. occidentalis eggs, with some pores at their bases (arrows indicate pores and some flat top spines); (h) inner view of basal part of second antenna of male (L. santarosae, n. sp.) note—slightly sinuous line of rows (arrow); (i) idem, L. occidentalis, note the sinuous line more marked (arrow) (scales in pm). 652 Table 1.—Allele frequencies for populations of Lin- deriella santarosae, new species (1 = Mesa de Burro). Linderiella occidentalis (2 = Fields Rd., 3 = Tim Bell Rd., 4 = Lagunita), and Eubranchipus vernalis (5). Population 1 2 3 4 5 Locus (14) (14) (6) (10) (7) PEP-C 0.143 0.857 0.000 1.000 0.000 1.000 0.000 0.000 0.000 0.000 0.000 1.000 0.000 0.000 0.000 1.000 0.000 0.000 0.000 0.000 1.000 0.000 0.000 1.000 0.000 0.000 1.000 0.000 1.000 0.000 1.000 0.000 0.000 0.000 1.000 0.000 0.000 0.000 1.000 0.071 0.857 0.071 0.000 0.000 0.000 0.000 0.000 0.429 0.571 0.000 1.000 0.000 0.000 1.000 0.000 1.000 0.000 0.833 0.167 0.000 0.167 0.833 0.000 0.000 0.333 0.667 0.333 0.667 0.000 0.000 0.000 0.000 0.000 0.000 0.500 0.500 0.000 1.000 0.000 0.000 1.000 0.100 0.900 0.100 0.900 0.000 0.000 0.500 0.500 0.000 0.000 0.100 0.900 0.100 0.900 0.000 0.000 0.000 0.000 0.000 0.000 0.100 0.900 0.100 0.800 0.100 0.000 1.000 0.000 1.000 0.000 0.000 1.000 0.571 0.429 0.000 0.500 0.500 0.000 0.000 0.000 0.000 0.000 0.000 0.571 0.429 0.786 0.214 0.000 0.000 1.000 0.000 0.000 0.929 0.071 PGI PGM G6PD PEP-D ME IDH AAT cm OoocoMmPAaAangHeRe OAD gH AdnGgA aAagkhk Age oP santarosae; this undescribed species also has tulip-shaped spines (Alonso & Alcaraz 1984, Thiéry & Champeau 1988). These differ- ences in resting egg morphology confirm previous studies on the taxonomic value of resting egg ornamentation as a tool for iden- tification of most anostracans (see review in Thiéry & Gasc, 1991, and Mura 1991, 1992a, 1992b). A preliminary electrophoretic survey po- tentially provides two additional characters for distinguishing between L. santarosae and L. occidentalis. Although samples sizes are small (14 L. santarosae & 30 L. occiden- talis), two loci (PEP-D & PGM) show fixed PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON allelic differences (Table 1). Additional sampling may show these allelic differences to be incompletely fixed, but its unlikely to substantially alter the large genetic distances between the two species. The average D within Linderiella occidentalis (3 popula- tions) is 0.054 and between L. occidentalis and L. santarosae is 0.378 (Fig. 11). These genetic differences are similar to those found between closely related species in the an- ostracan genus Branchinecta (Fugate 1992). For example, Branchinecta lynchi and Branchinecta sandiegonensis Fugate, 1993 have ranges similar to L. occidentalis and L. santarosae, respectively (Eng et al. 1990, Fugate 1992, 1993). The average D within B. lynchi is 0.154 (9 populations) and be- tween B. lynchi and B. sandiegonensis (3 populations) is 0.477. A limitation in the electrophoretic analysis arises due to the limited range of L. santarosae; the close proximity of the pools on the plateau (<5 km) allows water birds to freely move rest- ing eggs from pool to pool and makes the entire range of the species effectively one population. Distribution and habitat.—Linderiella santarosae is currently known only from vernal pools on the Santa Rosa Plateau Re- serve, Riverside Co., California. The Santa Rosa Plateau is approximately 300 km southeast of the nearest known populations of Linderiella in southern California (Ca- chuma Canyon, Santa Barbara Co., 34°42'N, 119°54'W, personal collection of Clyde Er- iksen, A2-25; Sulfur Mountain, Ventura Co., 34°26'N, 119°06'W, personal collection of Denton Belk DB. 693). These two popula- tions have not been studied electrophoret- ically, but morphologically appear to be Linderiella occidentalis. Another pool, one mile northwest of Olema, Marin Co. and across the road from the Visitors Center at Point Reyes National Seashore, 38°03'’N/ 122°48'W (coll. Gary M. Fellers, collection DB. 878), is reported to have individuals that “look like L. occidentalis, but have eggs with some short flat-topped spines” (D. Belk, VOLUME 107, NUMBER 4 OLS) GS) OL OB 0.5 0x4! Nei's Genetic Distance (D) Fig. 11. 653 L. santarosae L. occidentalis 2 L. occidentalis 3 L. occidentalis 4 E. vernalis UPGMA tree of Nei’s pairwise genetic distances between populations of Linderiella occidentalis and L. santarosae. Linderiella santarosae 1—Mesa de Burro, L. occidentalis 2—Fields Rd., 3—Tim Bell Rd., 4—Lagunita, Eubranchipus vernalis 5—Connecticut Valley Biological Supply. in litt.). We have not seen these individuals, but it would appear unlikely that these are individuals of L. santarosae. The Santa Rosa Plateau Reserve is an iso- lated unit of grassland and oak-woodland- chaparral in southeastern Riverside Co., California, owned and managed by the Na- ture Conservancy (Lathrop & Thorne 1968, 1978). Two large (~3 km7), flat-topped me- sas of volcanic origin are prominent features of the reserve and contain thirteen vernal pools ranging in size from 25 m? to over 100,000 m?. The mesas are capped by ol- ivine basalt lavas that have weathered to form soils of the Murrieta series. The av- erage elevation of the plateau is 610 m (La- throp & Thorne 1976a, 1976b, 1983). Collie & Lathrop (1976) report that pools on the Santa Rosa Plateau normally fill after winter rains (November to April) totaling 30-40 cm. The physico-chemical features of the 13 vernal pools are similar to those of pools containing L. occidentalis in other parts of the state (Eng et al. 1990); the con- ductivity of the water is low (147-189 wmhos cm! in February to 500-1050 in June), the PH is near 7.0 and water temperatures are usually below 20°C when L. santarosae is present (Collie & Lathrop 1976). Linderiella occidentalis is also commonly found at tem- peratures below 20°C and higher tempera- tures may be limiting to adults (Lanway 1974, Patton 1984, Eng et al. 1990). The flora of the reserve was listed by Munz 654 & Keck (1959) and later was characterized in a series of papers by Lathrop (1976), La- throp & Thorne (1968, 1976a, b, 1978, 1983) and Thorne & Lathrop (1969, 1970). Ko- pecko & Lathrop (1975) quantified the veg- etation in and around the vernal pools on the reserve and reported the following gen- era and species, also common in pools con- taining Linderiella from France, Spain and Morocco: Isoetes, Eleocharis, Callitriche, Juncus, Marsilea and Ranunculus aquatilis (Maire 1924, Barbero et al. 1982, Alonso 1985, Thiéry 1987). These genera have been labeled “‘typical” vernal pool plants in Cal- ifornia (Jain 1976). The fauna of vernal pools on the Santa Rosa Plateau is poorly characterized (Pe- quegnat 1951), but the branchiopod diver- sity is much lower than that found associ- ated with the other four Linderiella species (Alonso 1985, Thiéry 1986a, Thiery & Champeau 1988). The other four species are often found in pools containing notostra- cans and conchostracans, but neither are present in pools on the plateau. Two am- phibians are known to breed in the pools (Bufo sp. and Hyla sp.), and the following invertebrates are also present after winter rains: Branchinecta lynchi (Anostraca), Moina sp. (Cladocera), calanoid copepods, Ostracoda, Rotifera and Platyhelminthes. Discussion The modern disjunct distribution of the genus Linderiella has been the subject of some speculation (Belk 1984, Thiery 1986a, 1986b; Thiery & Champeau 1988, Bana- rescu 1990). These authors, using indirect evidence from a variety of sources, have attributed the disjunction to a vicariance event (sensu Bernardi 1986, Wiley 1988) involving the fragmentation of an ancestral taxon distributed across Laurasia. A num- ber of other animal and plant taxa with sim- ilar ecologies have either modern or fossil distributions across the Holarctic (e.g., Ra- ven 1971, DiCastri & Mooney 1973, Ban- arescu 1990). The limited fossil records of PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON branchiopods indicates that modern genera may have been formed before the Creta- ceous (Trusheim 1938, Tasch 1969, Tru- sova 1971, Jell & Duncan 1986). Also, dis- persal is passive, limited, and more likely to occur in a north-south rather than an east-west direction (Proctor 1964, Proctor & Malone 1965, Cruden 1966, Proctor et al. 1967, Boileau et al. 1992, Fugate 1992, Saunders et al. 1993). Although the vicariance hypothesis is consistent with the evidence for both an- cientness and limited dispersal, the alter- native hypothesis of long-distance dispersal cannot be fully eliminated due to the lack of a fossil record and an inadequate under- standing of phylogenetic relationships with- in the Anostraca. The lack of a fossil record is difficult to surmount, but a phylogeny could be reconstructed using morphologi- cal, biochemical or molecular characters. The phylogeny could then be employed to test the coincidence between the history of the species within the genus and the history of the geographic regions in which they are currently found (Brooks 1990, Wiley et al. 1991). Until such a test is performed, the vicariance hypothesis will remain specula- tive. Acknowledgments We thank C. Sassaman (University of California, Riverside), S. Morey (Univer- sity of California, Riverside), M. Simovich (University of San Diego, California) and G. Bell (Nature Conservancy) for shrimp or soil, D. Belk (Our Lady of the Lake Uni- versity, Texas) for loaning samples of L. occidentalis (DB. 430, DB. 693) and for comments on sample DB. 878, C. Eriksen (Claremont Colleges, California) for loaning samples A2-25, and C. Grill (University of Montpellier, France) for help with the scan- ning electron microscopy. Literature Cited Alonso, M. 1985. A survey of the Spanish Euphyl- lopoda.— Miscellana Zoologica 9:179-208. VOLUME 107, NUMBER 4 , & M. Alcaraz. 1984. Huevos resistentes de crustaceos eufilopodos no cladoceros de la pe- ninsula Ibérica: Observacion de la morfologia externa mediante técnicas de microscopia elec- tronica de barrido. —Oecologica Aquatica 7:73— 78. Banarescu, P. 1990. Zoogeography of fresh waters. Vol. 1: General distribution and dispersal of freshwater animals. AULA-Verlag, Wiesbaden, 511 pp. Barbero, M., J. Giudicelli, R. Loisel, P. Quezel, & E. Terzian. 1982. Etude des biocénoses des mares et ruisseaux temporaires 4 éphémérophytes dominants en région méditeranéenne fran- gaise.— Bulletin d’Ecologie 13(4):387—400. Belk, D. 1984. Patterns in anostracan distribution. Pp. 168-172 inS. Jain & P. Moyle, eds., Vernal pools and intermittent streams. Institute of Ecology Publication No. 28, Davis, California. Bernardi, G. 1986. La vicariance, la pseudovicari- ance et la convergence allopatrique.— Bulletin d’Ecologie 17(3):145-154. Boileau, M. G., P. D. N. Herbert, & S. S. Schwartz. 1992. Non-equilibrium gene frequency diver- gence: persistant founder effects in natural pop- ulations. —Journal of Evolutionary Biology 5:25— 39. Brooks, D.R. 1990. Parsimony analysis in historical biogeography and coevolution. Methodological and theoretical update.— Systematic Zoology 39: 14-30. Brtek, J. 1964. Eine neue Gattung und Familie der Ordnung Anostraca.— Annotationes Zoologicae et Botanicae, Bratislava 7:1-—7. Cannon, H. G., & F. M. Leak. 1933. On the mouth parts of the Branchiopoda.— Philosophical Transactions of the Royal Society of London, Series B 222:340-352. Collie, N., & E. W. Lathrop. 1976. Chemical char- acteristics of the standing water of a vernal pool on the Santa Rosa Plateau, Riverside County, California. Pp. 27-31, in S. Jain, ed., Vernal pools: their ecology and conservation. Institute of Ecology, Publication No. 9, Davis, Califor- nia. Cruden, R. W. 1966. Birds as agents of long-distance dispersal for disjunct plant groups of the tem- perate western hemisphere. —Evolution 20:517- 5312. Di Castri, F., & H. A. Mooney, eds. 1973. Mediter- ranean type ecosystems: origin and structure. Springer-Verlag, Berlin. Dodds, G. S. 1923. A new species of phyllopod.— Occasional Papers of the Museum of Zoology, University of Michigan 141:1-3. Eng, L. L., D. Belk, & C. H. Eriksen. 1990. Califor- nian Anostraca: distribution, habitat, and sta- tus.—Journal of Crustacean Biology 10(2):247— DY 655 Felsenstein, J. 1989. PHYLIP—Phylogeny inference package (Version 3.2).— Cladistics 5:164—266. Fugate,M. 1992. Speciation in the fairy shrimp genus Branchinecta (Crustacea: Anostraca) from North America. Unpublished Ph.D. Dissertation, University of California, Riverside, 270 pp. 1993. Branchinecta sandiegonensis, a new species of fairy shrimp (Crustacea: Anostraca) from western North America.— Proceedings of the Biological Society of Washington 106:296- 304. Heath, H. 1924. The external development of certain phyllopods.—Journal of Morphology 38:453- 483. Jain, S. 1976. Some biogeographic aspects of plant communities in vernal pools. Pp. 15-21 in S. Jain, ed., Vernal pools. Their ecology and con- servation. Institute of Ecology Publication No. 9, Davis California. Jell, P. A.. & P. M. Duncan. 1986. Invertebrates, mainly insects, from the freshwater, Lower Cre- taceous, Koonwarra Fossil Bed (Korumburra Group), South Gippsland, Victoria.— Associa- tion of Australasian Palaeontologists, Memoirs 3:111-205. Kopecko, K. J. P., & E. W. Lathrop. 1975. Vegetation zonation in a vernal marsh on the Santa Rosa Plateau of Riverside County, California. — Aliso 8(3):281-288. Lanway, C.S. 1974. Environmental factors affecting crustacean hatching in five temporary pools. Unpublished M.S. thesis, Department of Bio- logical Sciences, Chico State University, Chico, California, 89 pp. Lathrop, E.W. 1976. Vernal pools of the Santa Rosa Plateau, Riverside County, California. Pp. 22— 27 in S. Jain, ed., A symposium, Vernal pools, their ecology and conservation. Institute of Ecology, Publication No. 9, Davis, California. —., & R. F. Thorne. 1968. Flora of the Santa Rosa Plateau of the Santa Ana Mountains, Cal- fornia.— Aliso 6(4):17—40. ——,, & 1976a. The Vernal pools of Mesa de Burro of the Santa Rosa Plateau, Riverside County, California.— Aliso 8(4):433-445. ——, & 1976b. Vernal pools of the Santa Rosa Plateau, Riverside County, California. — Fremontia 4:9-11 (illustrations also 12-17). —_, & 1978. A flora of the Santa Ana Mountains, California.— Aliso 9(2):197-278. ——., & 1983. A flora of the vernal pools on the Santa Rosa Plateau, Riverside County, California.— Aliso 10(3):449-469. Linder, F. 1941. Contribution to the morphology and the taxonomy of the Branchiopoda Anostra- ca.—Zoologiska Bidrag fran Uppsala 20:101- 302. Maire, R. 1924. Etudes sur la végétation et la flore du Grand Atlas et du Moyen Atlas marocains. — 656 Mémoires de la Société des Sciences Naturelles du Maroc 7:5-200 + 16 pl. Munz, P. A., & D. D. Keck. 1959. A California flora. University of California Press, Berkeley, 1681 pp. Mura, G. 1991. SEM morphology of resting eggs in the species of the genus Branchinecta from North America. — Journal of Crustacean Biology 1 1(3): 432-436. 1992a. Pattern of egg shell morphology in thamnocephalids and streptocephalids of the New World (Anostraca).—Crustaceana 62(3): 300-311. 1992b. Additional remarks on cyst morpho- metrics in the anostracans and its significance. Part II: egg morphology.—Crustaceana 63(3): 225-246. —,, &A. Thiéry. 1986. Taxonomical significance of scanning electron microscopic morphology of the Euphyllopods’ resting eggs from Morocco. Part I. Anostraca.— Vie et Milieu 36:125-131. Nei, M. 1987. Molecular evolutionary genetics. Co- lumbia University Press, New York, 512 pp. Patton, S. E. 1984. The life history patterns and the distribution of two Anostraca. Linderiella oc- cidentalis and Branchinecta sp. Unpublished M.A. thesis, California State University, Chico, California, 50 pp. Pequegnat, W. E. 1951. The biota of the Santa Ana Mountains. — Journal of Entomological Zoology 42:1-84. Proctor, V. W. 1964. Viability of crustacean eggs recovered from ducks. —Ecology 45(3):656-658. —., & C.R. Malone. 1965. Further evidence of the passive dispersal of small aquatic organisms via the intestinal tract of birds.—Ecology 46: 728-729. ,& V.L. DeVlaming. 1967. Dispersal of aquatic organisms: viability of disseminules recovered from the intestinal tract of captive Kildeer.— Ecology 48(4):672-676. Raven, P.H. 1971. The relationships between ‘Med- iterranean’ floras. Pp. 119-134 in P. H. Davis, P. C. Harper, & I. C. Hedge, eds., Plant life of South-West Asia. The Botanical Society of Ed- inburgh, Edinburgh. Saunders, J. F., D. Belk, & R. Dufford. 1993. Per- sistence of Branchinecta paludosa (Anostraca) in southern Wyoming with notes on zoogeog- raphy.—Journal of Crustacean Biology 13(1): 184-189. Tasch, P. 1969. Branchiopoda. Pp. 128-191 in R. C. Moore, ed., Treatise on invertebrate paleontol- ogy, Part R—Arthroda 4, vol. 1. Geological So- ciety of America & University of Kansas Press, Lawrence, Kansas. Thiéry, A. 1986a. Linderiella africana n. sp., premier PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON représentant des Linderiellidae (Branchiopoda, Anostraca) en Afrique du Nord (Maroc). —Crus- taceana 51(2):155-162. 1986b. Les Crustacés Branchiopodes (An- ostraca, Notostraca et Conchostraca) du Maroc occidental I. Inventaire et répartition. — Bulletin de la Société d’Histoire Naturelle de Toulouse 122:145-155. 1987. Les Crustacés Branchiopodes Anos- traca, Notostraca et Conchostraca des mares temporaires (Dayas) du Maroc occidental. Tax- onomie, Biogéographie, Ecologie. Unpublished Thesis Doctorat es Science, Université d’Aix- Marseille III, 405 pp. —., & A. Champeau. 1988. Linderiella massa- liensis, new species (Anostraca: Linderiellidae), a Fairy shrimp from Southeastern France, its ecology and distribution.—Journal of Crusta- cean Biology 8(1):70-78. —, & C. Gasc. 1991. Resting eggs of Anostraca, Notostraca, and Spinicaudata (Crustacea, Bran- chiopoda) occurring in France: identification and taxonomical value.—Hydrobiologia 212:245— 259. Thorne, R. F., & E. W. Lathrop. 1969. A vernal marsh on the Santa Rosa Plateau of Riverside County, California.— Aliso 7(1):85-95. ——., & 1970. Pilularia americana on the Santa Rosa Plateau, Riverside County, Califor- nia.— Aliso 7:149-155. Trusheim, F. 1938. Triopsiden (Crustacea, Phyllo- poda) aus dem Keuper Frankens.— Palaeonto- logische Zeitschrift 19:198-216. Trusova, Y. K. 1971. O pervoy nakhodkye v ma- zozoye predstavitoley otryada Anostraca. (First discovery of members of the order Anostraca (Crustacea) in the Mesozoic.).— Paleontological Journal 4:68-73. Verill, A. E. 1869. Descriptions of some new Amer- ican phyllopod Crustacea.—American Journal of Science 48:244-254. Wiley, E. O. 1988. Vicariance biogeography. —An- nual Review of Ecology and Systematics 19:513- 542. —,, D. Siegel-Causey, D. R. Brooks, & V. A. Funk. 1991. The compleat cladist: a primer of phy- logenetic procedures.— The University of Kan- sas, Museum of Natural History, Special Pub- lication No. 19, 158 pp. (AT) Department of Animal Biology-Hy- drobiology, Faculty of Sciences, 33 rue Lou- is Pasteur, University of Avignon, 84000 Avignon, France; (MF) Department of Bi- ology, University of California, Riverside, California 92521, U.S.A. PROC. BIOL. SOC. WASH. 107(4), 1994, pp. 657-665 TWO NEW MARINE INTERSTITIAL OSTRACODA (CRUSTACEA: PUSSELLIDAE) FROM FIJI Shinichi Hiruta Abstract. — Danipussella rhamphodes and Pussella fijiensis (Podocopida, Bair- diacea) are described based upon specimens collected from coarse intertidal sand on the coral reef in Viti Levu, Fiji. These two species are discernible from congeneric species, mainly in the morphology of carapace and/or copulatory appendage. Marine interstitial ostracods are one of the useful animal groups in reconstructing historical biogeography (Danielopol & Wouters 1992). However, our knowledge of them is still poor in respect to tropical en- vironments, especially the coral reefs (Dan- ielopol & Hartmann 1986). The present pa- per deals with two new marine interstitial species of the genera Danipussella Wouters, 1988 and Pussella Danielopol, 1973 (Pus- sellidae: Bairdidacea: Podocopida) from Fiji. Several pussellid species including an as yet undescribed new one are known from the southwestern Pacific (Cabioch et al. 1986, Danielopol & Wouters 1992). The ostra- cods were collected during the survey for marine invertebrates, which was carried out by Dr. Ken-Ichi Tajika, during the Koshida Team Expedition, a part of ODMT (the Re- search and Exchange Program of Osaka University in the South Pacific Region), in 1985 (Koshida et al. 1986). Four pussellid specimens were discov- ered in samples from coarse intertidal sand on the coral reef of the Suva Barrier Reef in the vicinity of Suva in Viti Levu, Fiji. Two of them, possibly a female and a ju- venile, sustained damage to the carapace and appendages. Two other, male speci- mens belonging to the above-mentioned genera were found in good condition. The male holotypes of the two new species are deposited in the National Science Museum, Tokyo (NSMT). Superfamily Bairdiacea Sars, 1865 Family Pussellidae Danielopol, 1976 Genus Danipussella Wouters, 1988 Danipussella rhamphodes, new species (Figs. 1, 2) Specimen examined. —Holotype male: Fiji (NSMT-Cr11412). Specimen was col- lected by Dr. Ken-Ichi Tajika from the Suva Barrier Reef in the vicinity of Suva in Viti Levu, 16 Oct 1985. Etymology.—The specific name is de- rived from Greek rhamphodes meaning beaklike, in reference to the beak shape of the ventral process of the copulatory ap- pendage. Description. —Female unknown. Male (holotype). Carapace (Fig. 1-1-3; Fig. 2-1- 4) elongate, thin, 0.568 (both valves) mm long, 0.205 (right) mm high; dorsal margin evenly arched (Fig. 1-1 somewhat de- formed), connected smoothly with posterior rounded margin; ventral margin concave in the middle; posteroventral margin convex; anterior margin straight, connected with dorsal and ventral margins at almost right angles; anterior surface of valve near ante- rior margin with two large processes which are triangular in lateral view and have ter- minal stout spines: internally these pro- cesses are connected with the vestibulum. Surface smooth, with some long hairs along the ventral margin. Inner margin parallel to anterior margin; posterior inner lamella 658 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON -__________13-6, 813,15 50 2 9-12.14 3 Saar SG Fig. 1. Danipussella rhamphodes, new species. Male (holotype: NSMT-Cr11412). 1. internal view of right valve; 2. internal view of left valve; 3. internal view of anterior part of right valve; 4. right first antenna; 5. right second antenna; 6. right mandible; 7. idem, distal part; 8. right maxillula; 9. left first walking leg; 10. left second walking leg; 11. left third walking leg; 12. furca (arrow), abdominal bristle, and copulatory appendage; 13. brush- like organ; 14. lip; 15. Oesophagal chewing apparatus. VOLUME 107, NUMBER 4 Fig. 2. Danipussella rhamphodes, new species. Male (holotype: NSMT-Cr11412). 1. internal view of left valve; 2. idem, anterior part; 3. internal view of right valve; 4. idem, anterior part; 5. lip and testis; 6. legs (leg 2 broken) and testis; 7. furca (arrow) and copulatory appendage. Each bar represents 0.05 mm. broad, forming a large vestibulum. Adduc- tor muscle scar pattern not clearly visible. Normal pore cannals scattered. First antenna (Fig. 1-4) seven-segmented; first to third segments gradually decreasing in size; fourth segment quadrangular in lat- eral view; fifth to seventh segments very short, each with three, four, and three long setae respectively. Second antenna (Fig. 1-5) five-segment- ed; first segment short, with a long ven- trodistal seta; second segment somewhat shorter than fourth segment; third segment (basis) short, with a long mediolateral seta 660 and a very small exopodite which is one- segmented and has one short terminal seta; fourth segment (first endopodite segment) with a long dorsal subterminal seta, a short ventral seta arising from the distal third of the ventral margin, and a long ventrodistal seta which is 1.6 times as long as the fourth segment; terminal segment small, about one- seventh the length of fourth segment, with a short thin seta and a strong long claw, which is 3.75 times as long as the fourth segment. Mandible (Fig. 1-6, 7). Masticatory pro- cess with a short seta on the anterior margin; distal edge toothed, with nine teeth, of which some teeth are bifurcate or trifurcate. Palp four-segmented; first segment as long as sec- ond and third segments combined; second segment about one-half the length of third, with a long posterodistal seta; third segment with two setae, an anterior subterminal one and a posterodistal one; fourth segment ro- bust, having a thicker cuticle than preceding three segments, and as long as third seg- ment, with a short anterior subterminal seta, at least four distal setae of which the ante- riormost is longer and stronger than the oth- ers, and one very short posterior subter- minal seta. Maxillula (Fig. 1-8) furnished with three masticatory lobes; first and second lobes ar- ticulated with basal part, with three and four terminal setae respectively of which the ventral one is thicker than the others, and furnished with minute some terminal short teeth; third lobe connected with basal part, with seven terminal setae. Palp unseg- mented, with two setiferous ledges along dorsodistal part, each with one long seta, and two long distal setae of which the ven- tral one is thicker than the other and fur- nished with minute terminal teeth. Two long mouthward directed setae present. Respi- ratory plate with about ten setae. First walking leg (Fig. 1-9; Fig. 2-6) four- segmented; first segment with two long ven- tral setae arising from a short process on the ventral margin as illustrated; third segment PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON with a short anterodistal seta; length ratio of first to fourth segments and terminal claw 10:11.25:6:1:11.25. Second walking leg (Fig. 1-10) slender, with an anterodistal seta on second and third segments; length ratio of segments and terminal claw 35:30:25:1:45. Third walking leg (Fig. 1-11; Fig. 2-6) slen- der, with an anterodistal seta on second and third segments; length ratio of segments and terminal claw 17.5:15:13.5:1:25. Furca and abdominal bristle (sensu Wou- ters 1988) (Fig. 1-12; Fig. 2-7: arrow). Furca consisting of a pair of proximally thickened short setae. Abdominal bristle forming a short thin seta. Copulatory appendage (Fig. 1-12; Fig. 2-7). Each hemipenis with two anteriorly directed large processes, terminating in a round tip; dorsal process straight, tapered; ventral process beaklike, articulated with basal part, having a thick proximal part and a thin distal part which is slightly curved ventrally. Basal posterodorsal part with a small dorsally pointing process and a long recurved tube, whose proximal part is thick- ened, near preceding process. Testis (Fig. 2-5, 6) as shown in figures. Brush-shaped organ (Fig. 1-13) located near the base of the first walking leg, consisting of a pair of lobes, whose length is about five times as long as wide; each lobe with long filaments. Lip (Fig. 1-14; Fig. 2-5) with hairs along anterior margin and several teeth-like struc- tures, as illustrated. Oesophagal chewing apparatus (Fig. 1-15) present near the base of lip. Remarks.—Wouters (1988) established the genus Danipussella based upon the ma- terials from the Comoros as a monotypic genus within the family Pussellidae, con- taining three genera: Anchistrocheles, Pus- sella, and Danipussella. In this connection, Warne (1990) proposed the classification of the family Bythocyprididae (Bairdiacea), in which he considered Pussellinae as a sub- family of the family, and assigned Orlovi- bairdia McKenzie, 1978 and the new genus Bythopussella to the Pussellinae. Further, VOLUME 107, NUMBER 4 Danielopol and Wouters (1992) mention an as yet undescribed new pussellid genus from Papua New Guinea. Further study on the pussellid ostracods is needed. The type species Danipussella serpentina and the present new species are quite similar to each other in morphology of both hard and soft parts. Furthermore, the size of the present new species (0.568 mm) is almost the same as in D. serpentina (0.59 mm), which is much larger than Pussella-species (0.26 mm). As Wouters (1988:87) predict- ed, this character seems to have taxonomic value. While the structure of the copulatory appendage of both species is basically sim- ilar, the morphology of the ventral process is a useful character in differentiating be- tween them. D. serpentina has a long snake- like ventral process, while D. rhamphodes has a beaklike one. Since these two Danipussella-species were found at remote distance from each other (the Comoros and Fiji), other species of the genus might be distributed widely in trop- ical seas. Pussella fijiensis, new species (Figs. 3, 4) Specimen examined.—Holotype, male: Fiji (NSMT-Cr11413). The specimen was collected by Dr. Ken-Ichi Tajika from the Suva Barrier Reef in the vicinity of Suva in Viti Levu, 16 Oct 1985. Description. —Female unknown. Male (holotype). Carapace (Fig. 3-1, 2; Fig. 4-1- 6) thin, 0.452 (right), 0.445 (left) mm long, 0.181 (might), 0.177 (left) mm high. Dorsal and ventral margins arched, almost parallel to each other in anterior two-thirds of the length; posterodorsal margin strongly arched, terminating in a blunt triangular tip where ventral and dorsal margins meet; posteroventral margin with a large poste- riorly pointing process, which is connected with vestibulum, and has a wide base and a terminal spine; anterior margin truncate, weakly arched; anterior surface near ante- 661 rior margin with two large anteriorly point- ing processes; upper one smaller than lower one, both connected with vestibulum and furnished with terminal spine. Surface of carapace smooth, with scattered normal pore canals; radial pore canals with short hairs and some long hairs. Inner margin almost parallel to anterior, ventral, and postero- dorsal margins; anterior inner lammella broad. Hingement indistinct, but a small round process detected at the anterodistal edge of the right hinge. Adductor muscle scar (Fig. 4-7) consisting of four scars of different sizes as illustrated. First antenna (Fig. 3-3) seven-segmented; first to third segments gradually decreasing in size; fourth segment quadrangular in lat- eral view; fifth to seventh segments very small, with three, four, and three long setae respectively. Second antenna (Fig. 3-4) five-segment- ed; first segment with a long ventrodistal seta; second segment somewhat longer than fourth; third segment (basis) short, with a ventrodistal seta and a very small exopodite consisting of one segment, terminating in a short seta; fourth segment (first endopodite segment) with two subterminal setae, a long distal one, and a short ventral one, and a short ventrodistal claw-like seta; terminal segment small, about one-seventh of the length of fourth, with a short ventrodistal seta and a long claw which is about 3 times as long as the fourth and fifth segments com- bined. Mandible (Fig. 3-5) similar to preceding species; fourth segment of palp with four distal setae of different lengths. Maxillula (Fig. 3-6) furnished with three masticatory lobes; first and second lobes separated from base of maxillula, with two terminal setae, the ventral one being thicker than the other, and furnished with some short terminal teeth; third lobe connected with the body, with four distal setae. Palp unsegmented, with one long dorsal seta and two long distal setae, the ventral one thicker than the other, and furnished with some 662 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 3. Pussella fijiensis, new species. Male (holotype: NSMT-Cr11413). 1. internal view of right valve; 2. internal view of left valve; 3. right first antenna; 4. right second antenna and lip; 5. right mandible; 6. nght maxillula; 7. right first walking leg; 8. right second walking leg; 9. right third walking leg; 10. furca (f) and copulatory appendage (c); 11. brush-like organ; 12. oesophagal chewing apparatus. VOLUME 107, NUMBER 4 _—" a Fig. 4. Pussella fijiensis, new species. Male (holotype: NSMT-Cr11413). 1. internal view of right valve; 2. idem, anterior part; 3. idem, posterior part; 4. internal view of left valve; 5. idem, anterior part; 6. idem, posterior part; 7. adductor muscle scar (left); 8. furca and copulatory appendage; 9. testes. Each bar represents 0.05 mm. 664 short teeth. Two long mouthward directed setae present. Respiratory plate with about ten setae. First walking leg (Fig. 3-7) four-segment- ed; first segment with two long ventral setae of equal length, arising from a process on the middle of the ventral margin, and a long dorsodistal seta; third segment with a short subterminal seta on the anterior margin; distal claw strong; length ratio of first to fourth segments and distal claw 8.75:10: 8.75:1:10.75. Second walking leg (Fig. 3-8) more slender than first leg; first segment with a long ventral seta; second segment with a thin seta on the anterodistal edge; third seg- ment with a short seta on the anterodistal edge; length ratio of first to fourth segments and distal claw 12.3:10:10.7:1:19.3. Third walking leg (Fig. 3-9) similar to second walking leg; length ratio of first to fourth segments and distal claw 11.7:10:10.3:1:25. Copulatory appendage (Fig. 3-10; Fig. 4-8). Each hemipenis consisting of a round basal lobe, clasping apparatus (hook-like process with beak-like tip and a long curved claw-like process), and long copulatory tube bending acutely at proximal third and coil- ing along distal third. Testis (Fig. 4-9) coiled, as shown in figure. Furca (Fig. 3-10; Fig. 4-8) composed of two juxtaposed claw-like processes (shafts), terminating in a sharp point. A bristle is present between the two shafts. Brush-shaped organ (Fig. 3-11) consisting of a pair of lobes, whose length is about twice as long as wide; each lobe with long filaments. Lip (Fig. 3-4) with hairs along anterior margin. Oesophagal chewing apparatus (Fig. 3-12) present. Remarks.—The genus Pussella has one fossil species: P. infraturonica Pokorny, 1989 from Lower Turonian (90 million years B.P.) of Bohemia, Czechoslovakia, and three liv- ing ones including the present new species: P. botosaneanui Danielopol, 1973 from Cuba, P. danielopoli Maddocks, 1976 from Bermuda, and P. fijiensis sp. nov. Daniel- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON opol & Wouters (1992) discussed the evo- lution of the interstitial ostracods based upon these fossil and recent pussellids and other interstitial species. The present new species is discernible from P. botosaneanui and P. danielopoli in the shape of its carapace. As described above, P. fijiensis has two anterior and two posterior processes on both valves, but the latter two species have one anterior and one posterior processes and three anterior and three posterior processes. In this respect, the new species resembles P. infraturonica, but it is larger and more elongate than Turonian species (Pokorny 1989). In the male of P. botosaneanui, the third endopodite segment of the first walking leg is fused with the distal claw, while that of P. fijiensis clearly separated from it. The copulatory appendage of P. fijiensis is much different from that of P. botosaneanui in the shape of the clasping apparatus and copu- latory tube. The male of P. danielopoli is unknown. As Maddocks (1976) pointed out, further research is needed in the tropical interstitial environments, since our knowledge of the members of the genus Pussel/a is still poor. Acknowledgments I would like to express my sincere grati- tude to ODMT Koshida Team members, especially Dr. Y. Koshida, Mr. S. Horiuchi (Osaka University), Dr. K.-I. Tajika (Nihon University), Dr. U. Raj and his assistants (The University of the South Pacific, Fiji) who gave me the opportunity to study the interesting animals reported in this paper. Cordial thanks are also due to anonymous reviewers for useful information and helpful comments. Literature Cited Cabioch, G., R. Anglada, & J.-F. Babinot. 1986. Mi- crofaunes et paleoenvironnements des recifs frangeants quaternaires de Mamie et Ricaudy (Nouvelle-Caledonie).—Cahiers de Micropa- leontologie, N. S. 1(1—2):5-36. VOLUME 107, NUMBER 4 Danielopol, D. L. 1973. Preliminary report on the new family Pussellidae (Ostracoda, Podocopa). Pp. 145-149 in T. Orghidan et al., eds, Resultats des expéditions biospéléologiques cubano-rou- maines a Cuba Editura Academiei Republicii Socialiste Romania, Bucuresti. —, & G. Hartmann. 1986. Ostracoda. Pp. 265- 294 in L. Botosaneanu ed., Stygofauna Mundi, A faunistic, distributional and ecological syn- thesis of the world fauna inhabiting subterra- nean waters (including the marine interstitial). E. J. Bnll/Dr. W. Backhuys, Leiden. ——, & K. Wouters. 1992. Evolutionary (pa- leo)biology of marine interstitial Ostracoda. — Geobios 25(2):207—211. Koshida, Y., K.-I. Tajika, & S. Horiuchi. 1986. In- terim report of ODMT (Research and Exchange Program of Osaka University in the South Pa- cific region) Koshida Team Expedition in Viti Levu, Fiji, in 1985.—Science Reports. — College of General Education, Osaka University 35:13- 35. 665 Maddocks, R. F. 1976. Pussellinae are interstitial Bairdiidae.— Micropaleontology 22(2):194—214. Pokorny, V. 1989. Pussella and Saipanetta (Ostra- coda: Crustacea) in the lower Turonian of Bo- hemia, Czechoslovakia.—Casopis pro miner- alogii a geologii 34(3):225-233. Warne, M.T. 1990. Bythocyprididae (Ostracoda) from the Miocene of the Port Phillip and Western Port Basins, Victoria.— Proceedings of the Roy- al Society of Victoria 102(2):105—115. Wouters, K. 1988. Two interesting new interstitial Ostracoda (Crustacea) from the Comoros, with the description of Danipussella gen. nov.—Bul- letin de L’institut Royal des Sciences Naturelles de Belgique, Biologie 58:85-93. Biological Laboratory, Kushiro Campus, Hokkaido University of Education, Kushi- ro 085, Japan. PROC. BIOL. SOC. WASH. 107(4), 1994, pp. 666-679 ARCTODIAPTOMUS NOVOSIBIRICUS KIEFER, 1971 IN ALASKA AND NORTHWEST TERRITORIES, WITH NOTES ON A. ARAPAHOENSIS (DODDS, 1915) AND A KEY TO NEW WORLD SPECIES OF ARCTODIAPTOMUS (COPEPODA: CALANOIDA) Edward B. Reed Abstract. —Diaptomid copepods identified in North American literature as Diaptomus bacillifer Koelbel, 1885 lack the prominent process on the posterior surface of the second article of the exopodite of the male right fifth leg, which is recognized as a hallmark of 4. bacillifer and its immediate allies. Of the currently recognized species of Arctodiaptomus, the so-called bacillifer most clearly resembles 4. novosibiricus Kiefer, 1971 and may be that species. If so then A. novosibiricus is known from the New Siberian Islands, St. Paul Island, St. Matthew Island and the northern coast of Alaska; Victoria Island, Bernard Harbour and Adelaide Peninsula, N.W.T. Ecological circumstances suggest that future collecting may fill in gaps in the present discontinuous distribution. A. arapahoensis (Dodds, 1915) bears a great morphological similarity to A. acu- tilobatus (G. O. Sars, 1903). Presently A. acutilobatus is known from the Altai mountains, Caucasus and Kurdistan mountains. A. arapahoensis is known in the Rocky Mountains of Colorado, Montana, British Columbia and Alberta. Future collecting seems unlikely to erase the present discontinuous distribution; therefore it seems advisable to retain both names. The best-conceived, best-illustrated and most complete keys for the identification of North American freshwater copepods are contained in the 2nd Edition of Ward & Whipple’s Freshwater Biology (Edmondson 1959) (Reid 1990). Keys become dated not only through the discovery of “new” species but through studies of “‘old’’ species that lead to new insights and corrections in nomenclature, presumed affiliations and distribution. Not infrequently the identity of the species re- mains fairly constant, while its presumed affiliation at the genus level undergoes one or more revisions. Ideally keys would fulfill two functions: identification in a systematic manner and express phylogenetic relationships (Gurney 1931). For reasons discussed by Gurney, methodical routes to identification rarely satisfy the second function. One approach to methodical treatment is to reduce large and seemingly intractable groups to smaller, more manageable subgroups. The genus Diaptomus Westwood, 1836 in the broad sense surely qualifies as a large group; Dussart & Defaye (1983) listed 406 species plus 15 incertae sedis and several subspecies. Kiefer (1932a, 1932b) attempt- ed to find morphological groupings among the many named species of Diaptomus; in the latter paper he established a new sub- family, Diaptominae, in the family Diap- tomidae G. O. Sars, 1903. In the Diapto- minae, Kiefer place the genera Diaptomus Westwood, 1836, Hemidiaptomus G. O. Sars, 1903 and 10 new genera. In general non-North American copepodologists have VOLUME 107, NUMBER 4 accepted Kiefer’s genera. Students of North American diaptomids, especially those making comprehensive or intensive region- al investigations, have been inclined to view Kiefer’s genera as subgenera of Diaptomus Westwood, 1836; for example, Light (1939), Wilson (1959), Smith & Fernando (1978a) and Williamson (1991). However many others including Torke (1979) and Boileau & Hebert (1988) have accepted Kiefer’s gen- era. Light (1939) noted that while many spe- cies possessed definite specific characters, the range of morphological variation within the genus was not great. Wilson (1959) noted that the genera pro- posed by Kiefer lack the morphological gaps characteristic of other calanoid genera and added that more precise diagnoses, evalu- ation and interpretation of variation and distribution of North American species are required before formally defining genera. Sadly we are no nearer today to a com- prehensive taxonomic treatment of North American diaptomids than we were in 1959. Part of the difficulty may be in deciding on what constitutes a genus gap; a second less obvious difficulty may be in attempting to assess ecological niches in anthropocentric rather than copepodocentric terms. Fewer additions and corrections have oc- curred among the North American Calanoi- da since 1959 than among the Cyclopoida or Harpacticoida (Reid 1990). However there are reasons to reexamine the status of two diaptomids appearing in Wilson’s 1959 key: Diaptomus bacillifer Koelbel, 1885 and D. arapahoensis Dodds, 1915. I use Kiefer’s Arctodiaptomus genus be- cause the arctodiaptomids possess a set of morphological characters none of which alone is sufficient to constitute a gap but in concert they produce a fairly coherent mor- phological, if confusing ecological grouping. Specimens were dissected and examined in glycerin or lactic acid. Insofar as possible, examinations and drawings were made without cover slips. Details were verified with oil immersion lenses. 667 Marsh (1920) identified as Diaptomus ba- cillifer Koelbel, 1885 some copepods col- lected 6 October 1915 from a pond one foot in depth on a ridge 100 feet above sea level at Bernard Harbour, N.W.T. (68°45’N, 114°44'W). Concerning these animals Marsh wrote (1920 p. 6j): “Up to the present time no species of Diaptomus found on the American continent has been considered identical with those of Europe or Asia. Therefore a good deal of care was used to make certain that the identification of this species was correct. The determination is based on the original description of Koelbel, 1884 (sic), supplanted by the later descrip- tions, especially those of Sars and Schmeil.” In the same paper, Marsh said that he had examined specimens of D. bacillifer from Saint Paul Island, Alaska (57°07’N, 170°17'W). Marsh (1924) synonymized Diaptomus arapahoensis Dodds, 1915 with D. bacillifer Koelbel, 1885. Marsh (1929) reafhirmed this synonymy with the words “‘an examination of the description reveals that D. arapa- hoensis is D. bacillifer.” Wilson (1953) stated that she had ex- amined Marsh’s specimens of D. bacillifer and found his identification to be correct. She, however, believed D. arapahoensis not to be conspecific with D. bacillifer, and treated them as two species (Wilson 1959). Reed (1962, 1963) followed the lead of Marsh and Wilson and identified arcto- diaptomids from Alaska and N.W.T. as Diaptomus bacillifer. In 1956 while the late Mrs. Wilson was gathering specimens for a long-planned monograph on North American diaptomids (Damkaer 1988), I had the good fortune to work with her at the University of Saskatch- ewan for a short time. Wilson was com- pletely aware that some European copepo- dologists were questioning Marsh’s (1920) records of Diaptomus bacillifer. First, the press of larger, more demanding taxonomic problems, then ill health which cut short the proposed monograph (Damkaer 1988) pre- 668 vented Wilson from fully dealing with ques- tions surrounding the identity of North American D. bacillifer. Kiefer (1971) provided a complete review of taxonomy of the Arctodiaptomi; it is nec- essary here to recount only the portions which impinge directly on North American forms. In 1885 Koelbel described and named Diaptomus bacillifer from specimens col- lected in Plattensee, Hungary. Also in that year Imhoff named Diaptomus alpinus from an alpine lake in Berner Oberland (Kiefer 1971). Schmeil (1893) noted only one absolute difference between bacillifer and alpinus, the presence ofa ““Cuticularvorsprung am zwei- ten Aussenastsegments des richten mann- lichen Greiffusses” of D. bacillifer (Kiefer 1971 p. 127). Schmeil did not believe that the “Cuticularvorsprung” was sufficient to differentiate alpinus and bacillifer as species and synonymized alpinus with bacillifer, thus placing his not inconsiderable prestige behind Koelbel’s species. Schmeil’s decision had far-reaching ef- fects. For more than the next half century the works of many prominent copepodol- ogists would show the influence of Schmeil, including Giesbrecht and Schmeil (1898), Sars (1898, 1903b), Marsh in several pa- pers, Damian-Georgescu (1966) and Dus- sart (1967) among others. Kiefer (1932b) placed D. bacillifer in one of his new genera, Arctodiaptomus, and per- haps under the influence of Schmeil and others, omitted mention of D. alpinus. By the late 1960’s the nomenclature of Arcto- diaptomus had become so confused, partic- ularly in regard to bacillifer and alpinus, that Kiefer (1971) undertook a revision of the genus. Arctodiaptomus novosibiricus Kiefer, 1971 Fig. | Specimens examined.—Northwest Ter- ritories.— Victoria Island: Lady Franklin Point, 22 Aug 1957, Cambridge Bay PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON (69°03'N 104°05’W) 3 tundra ponds, 16 & 17 Sep 1957; Bernard Harbour: (68°44'N 114°27'W) little tundra pond, 20 Aug 1957; Adelaide Peninsula: shallow lake, 3 Sep 1957 (68°09'N 97°45’W). Alaska. — Near junction of Kikiakrorak and Colville rivers: (70°01'N 151°36'W), small tundra pond, 4 Aug 1955; Oliktok Point: (70°32'N 150°30'W), 3 tun- dra ponds, 24 & 25 Aug 1955; Point Bar- row: (71°33'N 156°30'W), 3 tundra ponds, several dates 1963-64; Saint Matthews Is- land: (60°30’N 172°45’W), tundra ponds. Description. —Most of the following de- scription is based on Point Barrow speci- mens and checked against specimens from other localities. Length without caudal setae: Most fe- males ranged between 1.4 and 1.6 mm, one female from Adelaide Peninsula measured 1.8 mm. Males ranged between 1.2 and 1.45 mm. Female: Body widest at mid-prosome (Fig. la). Pedigers 4 and 5 not divided dorsally, laterally a small furrow marks their joining (Fig. 1b). Posterior lateral margins pediger 5 moderately produced with lateral and dor- sal sensilla. Genital segment little widened anteriorly, nearly symmetrical, lateral pro- cesses small, left with oblique postero-lat- erally directed sensillum, that of right pro- jecting laterally (Fig. la), genital segment nearly as long as 2 succeeding segments and ramus combined. Urosome 2-segmented. Caudal rami about twice as long as wide, hairs on medial and lateral margins. Anten- nule extending to mid-genital segment (Fig. la); with 2 setae on articles 11 and 13, 1 on articles 14-19, seta on article 1 long, reach- ing beyond distal margin of article 5 (Fig. 1c). Rostrum small, pointed (ca. 25 um) (Fig. 1d). Lappet (Schmeil’s organ) present on posterior surface of second article of en- dopodite leg 2 (Fig. le). Leg 5: posteriorly basipodite 1 extending over proximal end of basipodite 2; basipodite 2 with lateral seta near distal margin (Fig. 1f); exopodite 1 about 2.5 to 3 times longer than wide; exopodite 2 seta well developed, claw with VOLUME 107, NUMBER 4 spinules on outer margin; exopodite 3, sep- arated, setae strong, inner about twice as long as outer, endopodite of 2 articles, but dividing suture not always distinct, termi- nating in truncate margin, no setae, sub- apical row of hairs (Fig. 1f). Male: Smaller, body shape similar to fe- male. Rostral points short (ca. 25 um), rounded, usual protrusion on right margin (Fig. 1g). Antennule: left similar to female in setation, except seta on article 1 short. Right antennule short spine on article 8, moderately long slender spines on articles 10 and 11, very small spine on article 12, short heavy spine on article 13 (Fig. 1h), seta on article 12 occasionally spine-like (Fig. 11). Process of antepenultimate article long, straight, tip often slightly thickened (Fig. 1J), extending at least to mid-length of ultimate article. Lappet of leg 2 similar to that of female but smaller. Leg 5 left; basipodite 1 lacks process but may overhang proximal margin of basipodite 2 (Fig. 1k); basipodite 2 inner margin with thin, flexible, cuticular process (Fig. 1k) extending from mid-seg- ment to mid-exopodite 1; exopodite ter- minal article with long, blunt fingerform outer process and curved, sharply pointed spine-like inner seta with spinules; proximal pad relatively small, hairy (Fig. 11), endo- podite 1- or 2-segmented, rounded, blunt tip, about equal to exopodite 1 in length, few small hairs. Leg 5 right: basipodite 1 with prominent process and sensillum (Fig. 1k, 1m) basipodite 2 with prominent semi- circular or elongate oval process on poste- rior surface (Fig. 1m), rounded process on proximal inner margin; flap-like cuticular membrane extending along middle % to 2 of inner margin; exopodite 1 projecting dis- tally over exopodite 2 at outer margin, small prominences on posterior distal margin; ex- opodite 2 about twice as long as wide; lateral spine straight, about 1.3 times article width, inserted at distal “4 of article, usually small cuticular process near base of spine; claw long, slender, recurved near tip. Endopodite 1- or 2-segmented, widened and rounded 669 on outer proximal margin, terminating in rounded and slanted tip bearing a sharp point on inner margin (Fig. In). The Point Barrow specimens belong to the genus Arctodiaptomus as characterized in the key of Kiefer (1978) on the basis of these morphological features: 1) configura- tion of the terminal processes of the exo- podite of male left fifth leg, 2) endopodite of female leg 5 armed only with a row of hairs, 3) two setae on article 11 of female and male left antennules and 4) spine of article 13 male right antennule not enlarged, spines of articles 10 and 11 strong, process of antepenultimate article straight. Comparison. —Inexplicably, Marsh (1920) overlooked the prominent process on the posterior surface of basipodite article 2 of the male right fifth leg exopodite. This process is clearly visible in lateral view of the intact specimen (Fig. 1m). Cover slip pressure on a dissected leg five may cause the protrusion of basipodite article | to flat- ten, obscuring the process of article 2. Marsh’s (1920) description and drawings do not mention or show a cuticular process on the second segment of the male right exopodite leg five. Notes in the Marsh Col- lection at the National Museum of Natural History, Smithsonian Institution reveal that Marsh compared descriptions and drawings of several authors in regard to this process. Perhaps the most telling is this note in re- spect to the Canadian Arctic Expedition specimens: “‘Differs from Giesbrecht’s anal- ysis only in 2nd seg rt exop having no [cu- ticular-vorsprung].”” Since Schmeil had al- ready accepted the presence or absence of the ““Vorsprung” as less than specific value, Marsh did not think it worth mentioning in his account of North American bacillifer. The Point Barrow males resemble males of A. alpinus in lacking a process on the posterior surface of the right second exo- podite article, but differ in minor respects of the right antennule. The Point Barrow females are similar to some other arctodiaptomid females in se- 670 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Arctodiaptomus novosibiricus. Female: a, habitus; b, pedigers 4 and 5, lateral view; c, antennule articles 1-6; d, rostrum; e, Schmeil’s organ; f, leg 5. Male: g, rostrum; h, right antennule; i, spines on right antennule articles 10-13; j, last three articles, right antennule; k, fifth legs, posterior view; 1, exopodite left leg 5; m, right exopodite, lateral view; n, fifth legs, anterior view. Scale: a, 400 um, all other bars 50 um. VOLUME 107, NUMBER 4 671 Table 1.—Subgeneric classification (A, H, R) number of setae () on articles 11, 13, 15 of female and male left antennule and presence (+) or absence (—) of process on posterior surface of 2nd article, right exopodite, male leg 5 of some Arctodiaptomus species. Data from Kiefer (1971). 11 (1) 11 (2) kurilensis Kiefer, 1937 byzantinus (Mann, 1940) H + A- parvispineus Kiefer, 1935 burduricus Kiefer, 1939 H? R - nepalensis Ueno, 1966 dentifer (Smirnov, 1928) H? A+ dudichi (Kiefer, 1932) A = saltillinus (Brewer, 1898) kerkyrensis (Pesta, 1935) (A) + A+ floridanus (Marsh, 1926) _pectinicornis (Wierzejski, (A) + 1887) A — dorsalis (Marsh, 1907) salinus (Daday, 1885) (A) + R - asymmetricus (Marsh, spinosus (Daday, 1891) 1907) (A) + R — steindachneri (Richard, 1897) A + stephanidesi (Pesta, 1935) A = wierzejski (Richard, 1885) A = arapahoensis (Dodds, 1915)R + acutilobatus (G. O. Sars, 1903) R + 11 (2), 13 (2) 11 (2), 13 (2), 15 (2) alpinus (Imhoff, 1885) acutilobatus (G. O.. Sars, R - 1903) R bacillifer (Koelbel, 1885) centetes (Brehm, 1938) R+ R+ laticeps (G. O. Sars, 1863) similis (Baird, 1859) A + A = niethammeri (Mann, arapahoensis (Dodds, 1940) R — 1915)R novosibericus Kiefer, 1971 R = piliger (Brehm, 1955) A — osmanus Kiefer, 1974 A ? Subgenera: A, Arctodiaptomus s. str.; H, Haplodiaptomus; R, Rhabdodiaptomus. tation of articles 11 and 13 (Table 1) and in possessing a lateral furrow between the fourth and fifth thoracic segments. The ros- trum of female bacillifer is long (40-50 um); whereas those of alpinus, laticeps and no- vosibiricus are short (20-25 um). Sars (1898) identified the New Siberian diaptomids as D. bacillifer Koelbel. Sars’ description of the New Siberian specimens is sketchy but his figures clearly show no process on the posterior surface of the sec- ond article of the male right fifth leg. Sars (1898:332) stated: “This (D. bacillifer) is one of the most characteristic Copepoda of the territory, occurring in great abundance as far north as the Expedition has reached.” Sars also stated (1898:333): “This species is also recorded by Prof. Lilljeborg from Nor- denskjolds Expedition, as occurring at In- serowa on the Siberian continent.” In the synonymy of D. bacillifer, Sars listed “‘D. retusus, Lilljeb. Ms.” Perhaps Lilljeborg’s D. retusus name was never published; nei- ther Kiefer (1971, 1978) nor Dussart and Defaye (1983) mention a D. retusus. The lack of the process on the posterior surface of article 2, male right exopodite fifth leg led Kiefer (1971) to reject Sars’ identification as D. bacillifer for the New Siberian specimens. He placed them in the A. alpinus group and named them 4A. no- vosibiricus. Apparently no specimens of no- vosibiricus from Siberia have come to hand since Sars’ (1898) collections. Borutskyi et al. (1991) in their review of freshwater cal- anoids of U.S.S.R. and discussion of A. no- 672 vosibiricus list only the island of Lachovsk1, one of the New Siberian Islands from whence Sars’ specimens came. The couplet which distinguishes male no- vosibiricus from male alpinus in Kiefer’s (1971) key reads: Seitendorn des 2 Aussenastgliedes rechts insertiert nahe der Basis der Endklaue (Abb. 41); Fortsatz des drittletzten Gliedes des Greifan- tenne sehr lang und zugespitzt (Abb. 41) novosibiricus Seitendorn sitzt etwa im 2 Drittel des Aussenrandes (Abb. 28-30); Fortsatz des drittletzten Gliedes der Greifantenne wie (Abb. 16a, 22, 25, 265 28—30537=39) i eee alpinus The females of A. alpinus and A. novo- sibiricus fall out together in Kiefer’s (1971) key. In Kiefer’s (1978) key males of A. novo- sibiricus are separated from males of A. al- pinus by the length of the process on the antepenultimate article of the right anten- nule: that of alpinus being at most as long as the following segment or usually shorter and that of novosibiricus at least as long as the following segment or much longer. Again in the key to females, a/pinus and novosibiricus fall out together along with 4. laticeps. Male A. laticeps are quite different from males of novosibiricus and alpinus in several characters. If one accepts Kiefer’s (1971) revision of the Arctodiaptomi, then the lack of a pro- cess on the posterior surface of the second article of the male fifth right exopodite is a strong argument to reject Marsh’s designa- tion of D. bacillifer for the Canadian Arctic Expedition specimens and by extension for the present animals. The morphological dif- ferences between alpinus and novosibiricus are not great and ultimately they may be shown to be conspecific. For the time being the least objectionable course is to designate the North American form as A. novosibiri- cus Kiefer, 1971, partly on morphological PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON evidence and partly on distributional and ecological considerations. Sars (1898) provided no ecological data for A. novosibiricus collected on the New Siberian Islands. In North America, A. no- vosibiricus is an inhabitant of small and shallow ponds. The Point Barrow speci- mens came from tundra ponds less than 250 m? in area, that varied in depth through the summer from 15 to 51 cm (Kalff 1967). The Adelaide Peninsula pond was about 6 ha, but less than 1 m deep. A polygon pond containing A. novosibiricus at Kikiakroarak was about 40 m? in area and 10 cm in depth. Arctodiaptomus arapahoensis (Dodds, 1915) Fig. 2 Specimens examined. —Colorado.—Lar- imer County: East (3295 m) and West Rain- bow lakes, Rawah Wilderness Area; Jack- son County: Lower Slide Lake; Hinsdale County: Emerald Lake (3112 m); San Juan County: Silverking, Ice (3752 m), Fuller (3843 m) lakes; Pitkin-Lake counties: alpine tundra ponds, Independence Pass (3689 m). Description.—Most of the following de- scription and drawings are based on East Rainbow and Silverking specimens and checked against specimens from other lo- calities. Female: Dodds (1915) gave the lengths of females that he examined as from 1.6 to 2.1 mm and lengths of males between 1.35 and 1.7 mm. Sixteen females from East Rain- bow Lake ranged between 1.46 and 1.75 mm, mean 1.53; 28 males ranged between 1.32 and 1.48, mean 1.41 mm. Wings of last prosome segment produced posteriorly past level of sensilla on genital segment (Fig. 2a). Reflexed antennules ex- tend to midlength of genital segment, which is essentially symmetrical, about 1.1 times longer than greatest width, sensilla directed postero-laterally. Urosome segments 2 & 3 combined about equal to length of genital segment. Rami haired on inner margins. Rostrum well developed, about 100 um VOLUME 107, NUMBER 4 673 Fig. 2. Arctodiaptomus arapahoensis. Female: a, pediger 5 and urosome; b, rostrum; c, antennule segments 1-8; d, Schmeil’s organ; e, leg 5. Male: f, rostrum; g, right antennule, spines on articles 8—13; h, last three articles, right antennule; i, right exopodite leg 5, lateral view; j, fifth legs, posterior view; k, fifth legs, anterior view; 1, terminal articles, left exopodite; m, right endopodite. Scale: a, 400 wm, all other bars 50 >. 674 in length (Fig. 2b). Seta of first segment of antennule about 260 um long, extending to midlength of article 7 (Fig. 2c). Antennule articles 9, 11, 13 and 15 with 2 setae each. Second article of endopodite leg 2 bearing hyaline lappet (Schmeil’s organ) (Fig. 2d). Leg 5. Basipodite 2 with small process bearing a short, moderately strong spine (Fig. 2e). Inner margin basipodite 2 much longer than outer margin. First article of exopodite long, slender, about 2 times as long as wide; claw of second article sinuous on inner mar- gin, finely denticulate on outer. Article 3 of exopodite well developed, distinct from ar- ticle 2, outer seta about twice the length of inner, both well developed and slightly spi- nulose. Endopodite of one article, end square bearing few hairs, with spine on inner cor- ner. Male: Rostrum smaller than that of fe- male, tips 40 to 50 um, right side with usual protuberance on margin (Fig. 2f). Left an- tennule articles 9 and 11 with 2 setae, one seta on articles 12-17. Right antennule, ar- ticle 8 with short spine, spines of articles 10 and 11, moderately long, slender; spine of article 12 short, spine ofarticle 13 moderate (Fig. 2g); antepenultimate article with long slender process extending nearly to distal end of last article (Fig. 2h). Schmeil’s organ smaller than that of fe- male. Right leg 5, basipodite article 1 with prominent posterior process; basipodite 2 with posteriorly directed process on poste- rior margin (Fig. 21, k), flap of hyaline mem- brane mid-inner margin; exopodite article 1 produced into point on outer margin (Fig. 21, k) exopodite article 2 with process on posterior face at midlength (Fig. 21, k) small hyaline process near base of lateral spine; lateral spine stout, inserted at distal 3 of article and about equalling or exceeding ar- ticle length. Claw long, curved, slender, fine- ly denticulate, with recurved tip. Endopo- dite long, nearly reaching midlength of exopodite 2, apex oblique, acute. Left leg (Fig. 2j, k), basipodite 2 with long, PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON hyaline process on inner margin; digital pro- cess of terminal article of exopodite elon- gate, somewhat broadened, with roughened inner border (Fig. 21); inner process a stout, spinulose spine (Fig. 21). Endopodite 1 ar- ticle, terminating in sharp tip. Curiously Dodds (1915) failed to either mention or figure the prominent process on the posterior surface of the second basipo- dite of the exopodite of the right male fifth leg. A. arapahoensis bears an unmistakably clear morphological resemblance to A. acu- tilobatus, (G. O. Sars, 1903) (Sars 1903b). In 1953 when Wilson called attention to this resemblance, the setation of the antennules of acutilobatus was not known, and she be- lieved more information was needed to de- cide conspecificity. Kiefer (1971) found the setation of acutilobatus to be exactly the same as that reported for arapahoensis, but perhaps for want of further information, took no stand on the conspecificity of ara- pahoensis and acutilobatus. Arctodiaptomus arapahoensis is known from the Rocky Mountains of Colorado (Dodds 1915, Reed & Olive 1958, Wilson 1959), Montana (Wilson 1959) and British Columbia and Alberta (Anderson 1974). Arctodiaptomus arapahoensis has been most frequently found in lakes ranging from 1.4 to 150 ha and 2.5 to 34 m deep; however the Independence Pass record shows that it is capable of inhabiting smaller and shal- lower waters. Kiefer (1971) listed A. acutilobatus from the Altai mountains, lakes and ponds in the Caucasus mountains from 1800 to 3200 m and the mountains of Kurdistan (Turkey). To these locations Kiefer (1978) added Mongolia and ““Mundungsgebiet der Sam- ara.” In spite of the morphological congruity of Colorado specimens with Kiefer’s figures and descriptions of A. acutilobatus, it is my opinion that Dodd’s name arapahoensis should be retained for the North American form on distributional considerations. VOLUME 107, NUMBER 4 Discussion The morphological similarity of A. no- vosibiricus to an arctodiaptomid occurring along the northern coast of North America and that of A. acutilobatus and A. arapa- hoensis, at least in the characters used, is unquestioned. In the understanding of pres- ent day biological species, morphological similarity or lack thereof is not a sure guide to conspecificity or species distinctness. In addition to external morphology, ecologi- cal, physiological, behavioral, and most of all, information on reproductive compati- bility must be evaluated in answering ques- tions concerning conspecificity. Breeding experiments, chromosomal studies, allo- zyme analyses, electrophoresis and other biochemical techniques have supported dis- tinctness of species initially based on minor morphological differences (Boileau & He- bert 1988). In some instances of morpho- logical differences, conspecificity has been established by chromosomal techniques (Einsle 1963). In still other instances repro- ductive isolation has been shown in species that could not be differentiated morpholog- ically (Price 1958, Boileau 1991). A decision regarding the designation of North American arctodiaptomids requires a presently unverifiable assumption regard- ing reproductive status. The data which would shed light on their standings as bio- logical species are not available, nor are they likely to be available soon. A comparative study requiring live specimens would be useful. However, for practical reasons, the probability of such a study actually hap- pening must be vanishingly small. The North American populations of A. arapahoensis and Eastern Hemisphere pop- ulations of A. acutilobatus if reproductively isolated could represent two or more sibling species; if not reproductively isolated, then one species and possibly several subspecies. We are still learning about gene flow be- tween widely separated and nearby popu- lations. A. acutilobatus and A. arapahoensis 675 seem to occur in mountainous areas that are separated from each other by broad steppes, prairies, deserts and a broad ex- panse of ocean. Future collecting will almost certainly increase the number of waters known to contain these species in their re- spective mountainous areas but seems un- likely to produce populations in the vast areas separating the mountainous regions. Allopatry on the scale shown by acutilo- batus and arapahoensis generally hints at different species rather than discontinuous distribution of one species. Hence my opin- ion that both be retained as species pending information on reproductive compatibility. It is not known if A. novosibiricus from the new Siberian Islands would in fact pro- duce viable offspring with Point Barrow or Bernard Harbour copepods. The few eco- logical data available suggest that they may occupy similar habitats. The Siberian coastal regions north of the tree line contain very numerous bodies of water which range greatly in extent and depth (Zhadin & Gerd 1961). Black & Barksdale (1949) estimated that areas on the Alaskan Coastal Plain are 20 to 90 percent covered by water. Holmquist (1975) emphasized not only the number of ponds and lakes along the North American northwestern coast, but also the diversity of ecological conditions in them. Because of the similarity of terrain along the Arctic coasts of Siberia and North America, future collecting of copepods seems likely to fill in spaces between the presently known populations, thus reducing the discontinuity of distribution, and the likelihood of reproductive isolation. Kiefer’s 1971 study clearly illustrated the overall great similarity of many Arctodiap- tomus species and simultaneously illustrat- ed variation in morphological characters and combinations of characters. His study also underlines the importance of examining both sexes before reaching conclusions about identity. Females that are virtually specifi- cally indistinguishable may belong with 676 males that are clearly separable and vice ver- Sa. In establishing subgenera of Arctodiap- tomus Kiefer relied heavily on the form of the process of the third from last article of the male right antennule and on the 14th article of the antennule, as well as characters of the 5th pair of legs. Which of the morphological characters are chosen for emphasis will greatly influence the species groupings within the Arctodiap- tomi. Kiefer’s emphasis on male right an- tennules led to grouping together forms which have quite different patterns of an- tennular setation. Grouping by antennular setation leads to mixing together species with other different morphological characters. At least four patterns of setation on the female antennules and male left antennule are known in the genus Arctodiaptomus (Table 1). The presence or absence of a process on the posterior surface of the second exopo- dite article of the male right leg 5 does not seem to bear a consistent relationship to antennular setation. Key to New World species of Arctodiaptomus la. One seta on article 11 of 4 left and both 2 antennules 1b. Two setae on article 11 2a. Two setae on article 11, one seta 13-17 .... 6A. arapahoensis (Dodds, 1915) Two setae on article 13, one or two on articles 13-17 3a. Two setae on 13, one on article 15 | A. novosibiricus Kiefer, 1971 — New Siberian Islands, Alaska, Northwest Territories . Two setae on articles 13 and 15 . .... 2 A. arapahoensis (Dodds, 1915) — Colorado, Montana, British Co- lumbia, Alberta 4a. Genital segment: 2, right side pro- duced into a large lateral lobe dis- tal to usual process bearing sensil- 4b. Sa. SID) 6a. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON lum. Leg 5 6 right exopodite 2, lateral spine thick, less than length of article ae ae A. symmetricus (Marsh, 1907) Cuba Marsh (1907) Genital segment 8, large distal lobe lacking. Leg 5 6 exopodite 2, length of lateral spine about equal to or greater than that of article Leg 5 2, endopodite as long or nearly as long as exopodite 1, 2 pediger 4, usually with single, dou- ble or triple medial dorsal process. Leg 5 4, right exopodite 2, lateral spine inserted proximal to mid- length of article, longer than arti- cle; left basipodite 2, width and length about equal SURE et A. dorsalis (Marsh, 1907) Arizona (Cole 1961), Louisiana (Marsh 1907), Mississippi (Harris 1978), Oklahoma (Robertson 1970), Cuba (Smith & Fernando 1978b), Haiti (Kiefer 1936) as D. proximus, Costa Rica (Collado et al. 1984), Mexico (Suarez 1991), Colombia (unpub. Suarez & Reid 1992), Nicaragua (Herbst 1960) as D. alter, Venezuela (Gonzalez 1968) as D. proximus. Leg 5 2, endopodite '2 to %4 length of inner margin exopodite 1; 2 ped- iger 4, with or without dorsal me- dial process. Leg 5 6, right exop- odite 2 lateral spine inserted at mid-length or distally, spine length about equal to article length; left basipodite 2 longer than wide... 6 Right metasomal wing 2 produced outward beyond lateral margin of body (dorsal view). Leg 5 4, right exopodite greatly exceeding length of inner margin exopodite 1, lat- eral spine exopodite 2 inserted at mid-length of article. Right anten- nule 6, antepenultimate article without process A Oa eh an) A. kurilensis Kiefer, 1937 VOLUME 107, NUMBER 4 — Kurile Islands (Kiefer 1937), Aleu- tian Islands (Wilson 1959) 6b. Right metasomal wing ? not pro- truding beyond lateral margin of body. Leg 5 6 right exopodite, lat- eral spine inserted near distal end of article; right endopodite little if any longer than inner margin of exopodite 1. Right antennule 4, an- tepenultimate article with curved distal process 7a. Caudal ramus 2, usually with hairs on inner margin only, pediger 4 with or without dorsal medial pro- cess. Leg 5 2, claw of exopodite 2 about as long as exopodite 1. Leg 5 6, basipodite 2 with small hyaline process on inner margin ........ A. saltillinus (Brewer, 1898) — Nebraska (Brewer 1898), Oklaho- ma (Robertson 1970), Texas (Wil- son 1953). 7b. Caudal ramus 2, Usually with hairs on inner and outer margins. 2 ped- iger 4 with or without dorsal me- dial process. Leg 5 2, claw of ex- opodite 2 much shorter than exopodite 1. Leg 5 6, basipodite 2 lacking inner hyaline process RNs tote es A. floridanus (Marsh, 1926) — Florida (Marsh 1926), Georgia (Humes 1950) as D. albuquerquen- sis det. by Wilson (1953), Okla- homa (questionable record by Keeton 1959); discussed by Rob- ertson (1970). Acknowledgments I am indebted to the following people for the use of specimens of A. novosibiricus from Alaska and Northwest Territories: J. Kalff, Alaska; J. Bond, Bernard Harbour and Vic- toria Island; E. and A. MacPherson, Ade- laide Peninsula. W. C. Nelson loaned spec- imens of A. arapahoensis from Colorado lakes. I thank Janet W. Reid for providing cop- 677 ies of literature from the Wilson Copepod Library and Marsh’s notes concerning spec- imens from the Canadian Arctic Expedi- tion. I am particularly indebted to Janet W. Reid for aid in developing the key to New World arctodiaptomids and to her and M. F. Suarez for permission to use their un- published record of A. dorsalis from Colom- bia. Thomas Bowman kindly offered many helpful comments on an early draft manu- script. Two anonymous reviewers provided helpful comments. Literature Cited Anderson, R. S. 1974. Crustacean plankton com- munities of 340 lakes and ponds in and near the National Parks of the Canadian Rocky Moun- tains. — Journal of the Fisheries Research Board of Canada 31:855-869. Black, R. F., & W. L. Barksdale. 1949. Oriented lakes of northern Alaska. — Journal of Geology 57:105— 118. Boileau, M. G. 1991. A genetic determination of cryptic species (Copepoda:Calanoida) and their postglacial biogeography in North America.— Zoological Journal of the Linnean Society 102: 375-396. ——, & P. D. N. Hebert. 1988. Electrophoretic characterization of two closely related species of Leptodiaptomus.— Biochemical Systematics and Ecology 16:329-332. Borutskyi, E. V., L. A. Stepanova, & M.S. Kos. 1991. Keys to the freshwater Calanoida of U.S.S.R. Zoological Institute, Russian Academy of Sci- ence, Saint Petersburg, 503 pp. (In Russian with English summary). Brewer, R.H. 1898. A study of the Copepoda found in the vicinity of Lincoln, Nebraska.—Journal of the Cincinnati Society of Natural History 19: 119-138. Cole, C. A. 1961. Some calanoid copepods from Ar- izona, with notes on congeneric occurrences of Diaptomus species.— Limnology and Oceanog- raphy 6:432-443. Collado, C., D. Defaye, B. H. Dussart, & C. H. Fer- nando. 1984. The freshwater Copepoda (Crus- tacea) of Costa Rica with notes on some spe- cies. —Hydrobiologia 199:89-99. Damian-Georgescu, A. 1966. Calanoida (forma de apa dulce). Jn Fauna Republicii Populare Rom- ine, Crustacea 4(8):1—130. Damkaer, D. M. 1988. Mildred Stratton Wilson, co- 678 pepodologist (1909-1973).—Journal of Crus- tacean Biology 8:131-146. Dodds, G. S. 1915. Descriptions of two new species of Entomostraca from Colorado, with notes on other species. — Proceedings of the United States National Museum 49:97-102. Dussart, B. 1967. Les Copépodes des eaux continen- tales d’Europe occidentale, Volume 1, Calanoi- des et Harpacticoides. N. Boubée et Cie, Paris. ——,, & D. Defaye. 1983. Répertoire Mondial des Crustacés Copépodes des Eaux Intérieures. I. Calanoides-Centre National de la Recherche Scientifique, Paris. Edmondson, W. T. (ed.). 1959. Freshwater biology. 2nd ed. John Wiley & Sons, Inc., New York. Einsle, U. 1963. Untersuchungen tiber die Variabi- litat in Cyclops furcifer Claus, 1957.—Crusta- ceana 5:193-—204. Giesbrecht, W., & O. Schmeil. 1898. Copepoda. I. Gymoplea. Jn Das Tierreich 6 Liefering. Berlin. XIV + 169 pp. Gonzalez, S. A. 1968. Desarrollo Larvario de Diap- tomus proximus Kiefer (Copepoda, Calanoi- da). — Hydrobiologia 32(3—4):528-544. Gurney, R. 1931. British fresh-water Copepoda. Vol- ume I, Ray Society, London. Harris, M. J. 1978. Copepoda of northern Mississippi with a description of a new subspecies. — Tulane Studies in Zoology and Botany 20(1-2):27-34. Herbst, H. V. 1960. Copepoden (Crustacea, Ento- mostraca) aus Nicaragua und Stdperu.—Ge- wasser und Abwasser 27:27-54. Holmquist, C. 1975. Lakes of Northern Alaska and nordwestern Canada and their invertebrate fau- na.— Zoologische Jahrbucher fur Systematik 102: 333-484. Humes, A. G. 1950. Experimental copepod hosts of the broad tapeworm of man, Dibothriocephalus latus (L.).— Journal of Parasitology 36:54 1-547. Imhoff, O. E. 1885. Uber die “blassen Kolben” und den vorderen Antennen der Siisswasser-Calani- den.— Zoologischer Anzeiger 8:353-356. Kalff, J. 1967. Phytoplankton abundance and pri- mary production rates in two arctic ponds.— Ecology 48:558-565. Keeton, D. 1959. Limnological effects of introducing oil field brine in farm ponds to reduce the tur- bidity.—Oklahoma Fishery Research Labora- tory, Report 72. (cited in Robertson, 1970). Kiefer, F. 1932a. Versuch eines Systems der altwelt- lichen Diaptomiden (Copepoda Calanoida).— Zoologischer Anzeiger 100(7/8):2 13-220. . 1932b. Versuch eines Systems der Diaptomi- den (Copepoda Calanoida). — Zoologische Jahr- bucher, Abteilung fiir Systematik, Okologie und Geographie der Tiere 63:451-520. 1936. Freilebende Siiss- und Salzwasserco- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON pepoden von der Insel Haiti.—Archiv fiir Hy- drobiologie 30:263-317. 1937. Stisswassercopepoden aus Ostasien.— Zoologischer Anzeiger 118(11/12):299-301. 1971. Revision der bacillifer-Gruppe der Gattung Arctodiaptomus Kiefer. (Crustacea Co- pepoda: Calanoida).—Memorie dell’Istituto It- aliano di Idrobiolgia 27:113-267. 1978. Das Zooplankton der Binnengewasser 2. Teil Freilebende Copepoda.—Die Binnen- gewasser 26:1-343. Koelbel, C. 1885. Carcinologischer.—Sitzungabe- richte der Akademie von Wissenschaften Wien 90:312-326. Light, S. F. 1939. New American subgenera of Diap- tomus Westwood (Copepoda, Calanoida).— Transactions of the American Microscopical Society 58:473-484. Marsh, C.D. 1907. A revision of the North American species of Diaptomus.—Transactions of the Wisconsin Academy of Science 15:381-516. 1920. Report of the Canadian Arctic Expe- dition 1913-18. Volume VII: Crustacea, Part J: Freshwater Copepoda, 1-25. J. de La-roquerie Tache. Ottawa. 1924. A new locality for a species of Diap- tomus.—Science 59:485—486. . 1926. Onacollection of Copepoda from Flor- ida with a description of Diaptomus floridanus, new species. — Proceedings of the United States National Museum 70, Article 10:1—4. 1929. Distribution and key of the North American copepods of the genus Diaptomus, with the description of a new species. — Proceedings of the United States National Museum 75(14): 1-27. Price, J. L. 1958. Cryptic speciation in the vernalis group of Cyclopoida.— Canadian Journal of Zo- ology 36:285-303. Reed, E. B. 1962. Freshwater plankton Crustacea of the Colville River area, northern Alaska.— Arc- tic 15:27-S0. 1963. Records of freshwater Crustacea from arctic and subarctic Canada. Jn Contributions in Zoology.— Bulletin of the National Museum of Canada 199:29-62. —, & J.R. Olive. 1958. Altitudinal distribution of some entomostraca in Colorado.—Ecology 39:66-74. Reid, J. W. 1990. Taxonomic reference list for North American free-living Copepoda. — Bulletin of the North American Benthological Society 7:227— 232. Robertson, A. 1970. Distribution of calanoid cope- pods (Calanoida, Copepoda) in Oklahoma.— Proceedings of the Oklahoma Academy of Sci- ence 50:98-103. VOLUME 107, NUMBER 4 Sars, G.O. 1898. The Cladocera, Copepoda and Os- tracoda of the Jana Expedition.— Annuaire du Museé Zoologique de I’ Academie Imperiale des Sciences de St. Petersbourg 3;324-359 + 6 plates. 1903a. Copepoda Calanoida. An account of the Crustacea of Norway, with short descrip- tions and figures of all the species, 4:1—xii 145— 171, Bergen Museum, Bergen. 1903b. On the crustacean fauna of Central Asia. Part III. Copepoda and Ostracoda.—An- nuaire de Museé Zoologique de l’Academie Im- periale des Sciences de St. Petersbourg 8:196— 232. Schmeil, O. 1893. Copepoden des Rhatikon-Ge- birges.— Abhandlungen Naturforschung Gesell- shaft Halle 19:1—40. Smith, K., & C. H. Fernando. 1978a. A guide to the freshwater calanoid and cyclopoid copepod Crustacea of Ontario. Department of Biology, University of Waterloo, Ontario, 74 pp. ——_, & 1978b. The freshwater calanoid and cyclopoid copepod Crustacea of Cuba.— Canadian Journal of Zoology 56:2015—2023. Suarez, E. 1991. Nuevo registro de Diaptomus dor- salis Marsh (Copepoda: Calanoida) en Mexico y su distribucion en Ja zona epicontinental cen- tral del Caribe Mexicano.—Caribbean Journal of Science 27:200—203. (In Spanish with English summary). 679 Torke, B. G. 1979. Crustacean zooplankton data for 190 selected Wisconsin inland lakes.— Wiscon- sin Department of Natural Resources Research Report 101:1-69. Westwood, J.O. 1836. Art. Cyclops. Pp. 227-238 in C. F. Partington, ed., The British cyclopaedia of natural history Vol. II. London. Williamson, C. E. 1991. Copepoda Pp. 787-822 in J. Thorp & A. Covich, eds., Ecology and clas- sification of North American freshwater inver- tebrates. Academic Press, Inc., San Diego. Wilson, M. S. 1953. New and inadequately known North American species of the copepod genus Diaptomus.—Smithsonian Miscellaneous Col- lections 122(2):1-30. 1959. Calanoida. Pp. 738-794 in W. T. Ed- mondson, ed., Fresh-water biology. John Wiley & Sons, Inc., New York. Zhadin, V. I., & S. V. Gerd. 1961. Fauna and flora of the rivers, lakes and reservoirs of the U.S.S.R. State Training and Pedagogical Publishing House, Moscow, 599 pp. (In Russian, English Translation by Israel Program for Scientific Translations. Jerusalem, 626 pp. 1963). 1901 Stover Street, Fort Collins, Colo- rado 80525, U.S.A. PROC. BIOL. SOC. WASH. 107(4), 1994, pp. 680-698 OBSERVATIONS ON THE GENUS MYSIDOPSIS SARS, 1864 WITH THE DESIGNATION OF A NEW GENUS, AMERICAM YSIS, AND THE DESCRIPTIONS OF AMERICAMYSIS ALLENI AND A. STUCKI (PERACARIDA: MYSIDACEA: MYSIDAE), FROM THE GULF OF MEXICO W. Wayne Price, Richard W. Heard, and Lidia Stuck Abstract.— Americamysis, a new genus, 1s proposed to receive six American species: Mysidopsis bigelowi Tattersall, 1926; M. almyra Bowman, 1964; M. bahia Molenock, 1969; M. taironana Brattegard, 1973; and two new species, Americamysis alleni and A. stucki. Mysidopsis bigelowi is designated as the type species of the new genus. Americamysis is defined by a combination of characters, most notably: the presence of two articles in the carpo-propodus of thoracic endopods 3-8; the absence of a terminal article on antennal scale; and the presence of less than eight ventrolateral spines on uropodal endopod, all of which are confined to the region of the statocyst. Americamysis, Mysidopsis, and its subgenera are distinguished from each other by a variety of characters, including the morphology of the mandibular palp, antennal scale, and telson. A complete systematic review may indicate that the genus Mysidopsis sensu lato is polyphyletic and represents three or more distinct genera. The new species, Americamysis alleni and A. stucki, which are both only known from the Gulf of Mexico, are most similar and apparently most closely related to A. bigelowi. They can be separated from each other, A. bigelowi and the other species of Americamysis by the setation of their telsons and second thoracic en- dopods. A diagnostic table and key for separating the species of Americamysis and related species of Mysidopsis is presented. The distribution of the species within Americamysis extends along the Atlantic coasts of the Americas from the northeastern United States to Colombia. The genus Mysidopsis Sars, 1864 pres- ently contains a heterogeneous group of 49 nominal species, demonstrating a wide size range and a variety of morphological char- acters (Bacescu 1968b, Brattegard 1969, Tattersall 1969, Bacescu & Gleye 1979, Mauchline 1980). Brattegard (1969) dis- cussed the similarity of Mysidopsis almyra, M. bahia, and M. bigelowi and designated these three northwestern Atlantic species as the “‘almyra group.”’ Later (1973), he de- scribed M. taironana from the Caribbean coast of Colombia and noted its affinities with this group. Tattersall (1951), Clarke (1956), Stuck et al. (1979a, 1979b), Modlin (1982), Price (1982) and Price et al. (1986) reported M. bigelowi from the coastal and shelf waters of the northern Gulf of Mexico. Reexami- nation of the material used in these studies indicates the specimens identified as “/M. bigelowi”’ actually represent two unde- scribed species within the “‘a/myra group” and that M. bigelowi sensu Tattersall (1926) does not occur in the Gulf of Mexico. Based on Brattegard’s observations and our ex- amination of specimens of M. almyra, M. bahia, M. bigelowi, and the two undescribed species from the Gulf of Mexico, we now believe that the members of the “‘a/myra VOLUME 107, NUMBER 4 group” represent a distinct genus, which is endemic to the Atlantic coastal regions of the Americas. Americamysis, new genus Diagnosis.—Thoracic endopod 1 com- posed of 4 articles, preischium and ischium fused; thoracic endopods 3-8 having carpo- propodus with 2 articles; antennal scale lacking distal article and spinose tip; telson linguiform, midapical spine setae much (4 times) longer than lateral spine setae; uro- podal endopod armed with 8 or fewer spines adjacent to statocyst or along inner margin. Type species: Americamysis bigelowi (Tattersall, 1926), new combination. Additional species: A. almyra (Bowman, 1964), new combination; A. bahia (Mole- nock, 1969), new combination; A. tairon- ana (Brattegard, 1973), new combination; A. alleni, new species; and A. stucki, new species. Remarks. — Americamysis appears to form a natural group containing six species, all of which seem to be confined to the north- western Atlantic. The known species of this genus are endemic to estuarine and shallow shelf waters along the east coasts of the Americas from New England to Colombia. The combination of a distally unarticu- lated antennal scale, a 2-articulate carpo- propodus on the endopods of thoracic limbs 3-8, distinctly fewer articles in the exopod of the fourth male pleopod, and the presence of one or more mesial pairs of apical spine- setae on the telson distinguishes the genus Americamysis from the type species of Mys- idopsis, M. didelphys (Norman, 1863). Since Sars’ (1885) generic diagnosis, the generic limits of Mysidopsis have been broadened, resulting in the inclusion of many apparently distantly related species. Among the characters given by Sars (1885) in his generic definition of Mysidopsis were (1) the presence of an apical or distal article on the antennal scale and (2) three articles in the carpo-propodus of thoracic endopods 681 3-8. The morphological limits were broad- ened by the description of Mysidopsis acuta Hansen, 1913, the first species assigned to the genus having an antennal scale termi- nating in an acute spinose process and lack- ing an apical suture. W. Tattersall (1926) further expanded the generic definition with the description of M. bigelowi, a species characterized by having two (instead of three) articles in the carpo-propodus of tho- racic endopods 3-8. Like M. acuta, M. bige- Jowi has an antennal scale lacking a distal suture. Unlike M. acuta, however, the tip of M. bigelowi’s antennal scale is blunt with distal setae. Excluding M. bigelowi and the other species being assigned to America- mysis, there are now six nominal species of Mysidopsis that lack a distal article on the antennal scales (Hansen 1913, O. Tattersall 1962, Brattegard 1973, Brattegard 1974b, Gleye 1982, and Bacescu 1984). In a synopsis of Mysidopsis, O. Tattersall (1969) erroneously stated in her rediagnosis of the genus that the carpo-propodus of tho- racic endopods 3-8 “is divided by a trans- verse suture into two subsegments ...,” which would exclude the type species and most of the other described species of the genus. She also considered the 4-articulate endopod of first thoracic appendages which was due to the fusion of the preischium and ischium, a unique character of the genus Mysidopsis. At present, however, there ap- pear to be at least two other genera, Meta- mysidopsis W. Tattersall, 1951 and Taph- romysis Banner, 1953 that have their first thoracic endopods composed of four arti- cles as in Mysidopsis, instead of the five normally found in other genera of the family Mysidae (see Mauchline 1980). Based on the illustrations of Brattegard (1969, p. 62, fig. 18E) and Gleye (1982, fig. 1f), respec- tively, the genera Brasilomysis Bacescu, 1968 and Cubanomysis Bacescu, 1968 also may have 4-articulate first thoracic endo- pods. Brattegard’s (1969) illustrations of Brasilomysis castroi Bacescu, 1968 were based on specimens from the Bahamas, 682 while those for Cubanomysis mysteriosa Gleye, 1982 accompanied the original de- scription based on specimens from Califor- nia waters. Though not described in the text, the illustration (fig. 4C) for the only other nominal species of Brasilomysis, B. (=Mys- idopsis) inermis Coifmann, 1937) indicates four articles in the first thoracic endopod. The generic status of B. inermis is still un- settled. Its original and only description was quite incomplete and the type material, which came from Pacific waters off Ecuador, appears to be no longer extant. The number of articles in the first limb of the type and remaining species of Cubanomysis, C. ji- menesi Bacescu, 1968b, has not been re- ported or illustrated. O. Tattersall (1969) treated 21 species of Mysidopsis. Since her work the number of described species has more than doubled to 49 (including the four species being placed in Americamysis). Of the 28 species added to the genus since O. Tattersall’s synopsis, the majority (21) are from the western At- lantic (Bacescu 1968b, 1984; Brattegard 1969, 1973, 1974a, 1974b; Modlin 1987, Moleneck 1969, da Silva 1979, Hoffmeyer 1993, Ortiz & Lalana 1993). One of these, M. cojimarensis Ortiz & Lalana, 1993, may be synonymous with Antromysis (Parvi- mysis) bahamensis (Brattegard 1970); how- ever, it is tentatively retained in the genus Mysidopsis pending a comparison of the types of these two species. The remaining seven species are from the coastal waters of East and West Africa, the Pacific coast of North America, and Japan (Murano 1970, Bacescu & Vasilescu 1973, Bacescu 1975, Bacescu & Gleye 1979, Gleye 1982, Woold- ridge 1988). The combination of a 2-articulate carpo- propodus in thoracic endopods 3-8 and an antennal scale lacking a distal article distin- guishes Americamysis from the heteroge- nous group of species tentatively retained within the genus Mysidopsis. Eight of these species, like the members of the genus Americamysis, have thoracic endopods 3-8 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON with a 2-articulate carpo-propodus, but their antennal scales have a terminal article. Five of these, M. californica W. Tattersall, 1932; M. intii Holmquist, 1957; M. tortonesi Ba- cescu, 1968b; M. robustispina Brattegard, 1969 and M. ankeli Brattegard, 1973, differ strikingly from both the type species (M. didelphys) and the species being referred to the genus Americamysis by the presence of numerous spine-setae or true spines (VM. an- keli, M. intii) along the inner margin of the uropodal endopod. These species, which are known only from the Atlantic and Pacific waters of the Americas, appear to be more closely related to the genus Americamysis than to the genus Mysidopsis. We have tem- porarily retained them within Mysidopsis pending further systematic revision of the genus sensu O. Tattersall (1969). Because of the spiniform tip of their antennal scales, three of these species (M/. californica, M. robustispina, M. tortonesi), along with M. acuta and M. coelhoi Bacescu, 1968b, were included in Brattegard’s (1969) “coelhoi group.” Three other species of Mysidopsis, two from South Africa (M. camelina O. Tatter- sall, 1955 and M. suedafricana O. Tatter- sall, 1969) and one from the tropical north- western Atlantic (M. bispinulata Brattegard, 1974a), also have thoracic endopods 3-8 with a 2-articulate carpo-propodus. How- ever, in other respects these species are quite distinct from each other and are not appar- ently closely related to other species with similarly articulated thoracic endopods. Based largely on the presence of highly modified mandibular palps and telson se- tation, Bacescu & Gleye (1979) created the two monotypic subgenera, Mysidopsoides and Pseudomysidopsis, to receive Mysidop- sis bispinosa O. Tattersall, 1969 and Mys- idopsis camelina O. Tattersall, 1955, re- spectively. Both of these subgenera are endemic to South Africa and may represent distinct genera. In conclusion, based on our own obser- vations and a review of the literature, we VOLUME 107, NUMBER 4 consider that the genus Mysidopsis sensu lato represents a polyphyletic melange of taxa. We believe that in addition to the “al- myra”’ group (Americamysis), the other sub- generic groupings proposed by Brattegard (1969), the subgenera described by Bacescu & Gleye (1979), and other distinctive, ‘atypical’ species or species groups (i.e., M7. angusta, M. acuta, M. ankeli-intii) may be unrelated and may represent distinct gen- era. Except for members of the “almyra group’, we have been unable to critically examine many of the other nominal species of Mysidopsis, especially those from Euro- pean and African waters, belonging to these other subgeneric groupings or “atypical” species. Systematic studies of the heterog- enous species or species complexes still ac- commodated within the genus Mysidopsis is needed. Such studies using the systematic tools of cladistics and molecular genetics will likely result in the removal of additional species from the genus. Table 1 presents a listing of the nominal species currently retained within the genus Mysidopsis with information on their gen- eral distribution and distinctive characters for some of the atypical species or species groups. Mysidopsis incisa Sars, 1885, which has only been tentatively reported once (Thompson 1894) since its original descrip- tion, is included. Its taxonomic status, how- ever, remains uncertain. Eight species orig- inally assigned to the genus, including four in this report, have been transferred to other genera. These are listed in Table 2 with their current generic designations. Americamysis alleni, new species Figs. 1-2 Mysidopsis bigelowi. — Tattersall 1951:139 (in part).— Price 1982:16 (in part), figs. 17, 20.—Price et al. 1986:49 (in part). Type material.—Holotype: adult male (USNM 253072), length 5.1 mm, Biloxi Beach, Mississippi; Mississippi Sound (30°23.3'N, 88°58.5'W): depth 1 m, sand 683 substratum, dredge net, David Hard (coll.), 27 Oct 1991.—Paratypes: 5 6, 5 2 (USNM 253073), 56, 5? (GCRL 1317), same collec- tion data as for holotype. Additional material examined. —Texas coast: 4 6 (6.3—-7.3 mm), 4 2 (7.3-7.4), La- vaca Bay, station | (28°35.3'N, 96°31.7'W), depth 2.1 m, otter trawl, 4 Mar 1971.—6 6 (6.6—7.3 mm), 2 2 (6.9-7.4 mm), Lavaca Bay, station 7 (28°33.3'N, 96°28.5’W), depth 2.7 m, otter trawl, 5 Dec 1970, 2 Feb 1971.— 6 6 (4.4—5.3), 2 2 (5.0—5.1 mm), West Bay, station WB6 (94°58.5'N, 29°15.8’W), depth 1.2 m, Oct 1973.—15 6 (5.3-6.0 mm) 15 2 (5.3-5.6 mm), West Bay, station WB2 (94°57.2'N, 29°14.5W), depth 1.5 m, Oct 1973.—15 6 (3.8-5.2), 15 2 (3.8-5.4 mm), West Bay, station WB4 (94°57.8'N, 29°15.0’'W) depth 1.5 m, June 1973. West Bay specimens collected with epibenthic sled and all Texas material in collection of WWP. Diagnosis.— Apex of antennal scale not pointed; endopod of thoracic leg 2 normally developed in both sexes, merus with 2-3 simple setae on inner proximal margin and 4 simple setae along outer margin; inner margin of carpo-propodus armed with 5- 10, occasionally 4, spiniform setae; carpo- propodus of thoracic endopods 3-8 2-segmented; exopod of pleopod 4 of male 7-segmented; endopod of uropods armed with 3—4 (rarely 2 or 5) spine-setae near statocyst; apex of telson with 3 pairs of strong spine-setae, outermost pair '2—%4 length of inner pairs, innermost pair 4 or less length of telson and shorter than, or equal in length to, adjacent pair. Description. —General body form (Fig. 1A): moderately slender, adult males to 7.3 mm and females to 7.4 mm; anterior margin of carapace produced between eyes into short, triangular rostrum; posterior dorsal margin emarginate, exposing thoracic seg- ment 8; anterolateral corners rounded. Antennular peduncle (Fig. 1B): more ro- bust in males than females; segment | about as long as segments 2 and 3 combined, dis- tomedial margin armed with 4—5 long and 684 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 1.—Species presently included in the genus Mysidopsis by region. Distinctive characters of some species groups are presented. General location Character* Eastern Atlantic M. angusta Sars, 1864 North Europe, Western Mediterranean (3) M. (Mysidopsoides) bispinosa O. Tattersall, 1969 South Africa (1, 5) M. (Pseudomysidopsis) camelina W. Tattersall, 1955 South Africa M. didelphys (Norman, 1863): Type species North Europe, Iceland, Western Mediterra- nean M. eremita W. Tattersall, 1962 South Africa M. gibbosa Sars, 1864 North Europe, Western Mediterranean M. major Zimmer, 1912 South Africa M. schultzei (Zimmer, 1912) South Africa M. similis (Zimmer, 1912) South Africa M. suedafrikana O. Tattersall, 1969 South Africa (1) Western Atlantic M. acuta Hansen, 1913 Argentina (4) M. ankeli Brattegard, 1973 Colombia (1, 2) M. arenosa Brattegard, 1974 Panama M. badius Modlin, 1987 Belize, Mexico M. bispinulata Brattegard, 1974 Colombia, Panama (1, 5) M. brattstroemi Brattegard, 1969 Bahamas, South Florida, Panama M. coelhoi Bacescu, 1968 Brazil (4) M. cojimarensis Ortiz & Lalana, 1993** Cuba M. cultrata Brattegard, 1973 Colombia M. eclipis Brattegard, 1969 South Florida M. furca Bowman, 1957 Southeastern United States M. juniae da Silva, 1979 Brazil M. mathewsoni Brattegard, 1969 Bahamas M. mauchlinei Brattegard, 1974 Panama M. mortenseni W. Tattersall, 1951 Bahamas, South Florida, Caribbean M. rionegrensis Hoffmeyer, 1993 Argentina M. robusta Brattegard, 1974 Colombia M. robustispina Brattegard, 1969 Bahamas, Colombia (1, 4) M. sankarenkuttkyi Bacescu, 1984 Brazil M. tortonesei Bacescu, 1968 Brazil, Colombia (1, 4) M. velifera Brattegard, 1973 Caribbean M. virgulata Brattegard, 1974 Colombia Eastern Pacific M. acuta Hansen, 1913 sensu Holmquist (1957) Chile (4) M. brattegardi Bacescu & Gleye, 1979 California M. californica W. Tattersall, 1932 California (1, 4) M. cathengelae Gleye, 1982 California M. intii Holmquist, 1957 Chile (1, 2) M. onofrensis Bacescu & Gleye, 1979 California Western Pacific M. japonica Ii, 1964 Japan M. surugae Murano, 1970 Japan M. incisa Sars, 1884 incertae sedis Southern Australia (3) Indian Ocean M. buffaloensis Wooldredge, 1988 South Africa M. coralicola Bacescu, 1975 Tanzania Le VOLUME 107, NUMBER 4 Table 1.—Continued. M. helvillensis Nouvel, 1964 M. indica W. Tattersall, 1922 M. kempi W. Tattersall, 1922 M. kenyana Bacescu & Vasilesco, 1973 685 General location Mozambique India India Kenya * Character codes: (1) 2-articulate carpo-propodus on thoracic endopods 3-8; (2) margin of uropodal endopod with serrate spines; (3) telson distinctly cleft; (4) antennal scale with acute, spinose tip; (5) lateral margins of telson lacking spine-setae. ** This species may be synonymous with Antromysis (Parvimysis) bahamensis (Brattegard, 1970). 2-3 shorter plumose setae, distolateral mar- gin with | short and 4 long plumose setae; medial margin of segment 2 with strong simple spine and 5 plumose setae; segment 3 with 2 plumose setae along medial margin and 5-6 plumose setae distally, distal mar- gin with dorsomedial lobe bearing 2 tooth- like processes and 2 plumose and 2-3 sim- ple setae, 1 plumose and 1 simple seta grouped proximal to dorsomedial lobe, males with densely setose lobe on outer ven- tral surface. Antenna (Fig. 1C): scale lanceolate, reaching beyond distal end of antennular peduncle, 5.5—6 times as long as maximum width, inner margin slightly convex, outer margin slightly concave, setose all around, lacking distal article; antennal peduncle ex- tending about 0.6 length of scale, article 2 about 1*4 times as long as 3, with 2-3 plu- mose setae on inner distal margin and 1 plumose seta on outer distal margin; article 3 with 2 plumose and 3 simple setae on inner distal margin and 2 plumose setae along outer margin; distolateral corner of sympod with prominent tooth. Eyes (Fig. 1A): large, cornea wider than distal end of eyestalk, lacking ocular papilla. Labrum (Fig. 1D): rounded anteriorly; posterior margin with medial emargination, middle 7% with fine setae. Mandible (Fig. 1E): cutting edges typical of genus. Palp (Fig. 1F) 3-articulate, articles 2 and 3 more robust in males than females; article 2 armed with 10-14 submarginal simple setae along outer margin and 10-15 simple setae on inner margin, distal margin with 3-5 simple setae; article 3 about 2 as long as 2, inner surface with 3-4 marginal and 3—4 submarginal simple setae, distal portion armed with 1 long simple seta, 1 long curved spine-seta with medial barbs and 9 thick blunt densely barbed spine-se- tae, outer margin with 2 simple setae. Maxillule (Fig. 1G): outer lobe with 9 stout, sparsely serrate apical spine-setae; in- Table 2.—Nominal species of Mysidopsis transferred to other genera. Present generic Oniginal designation designation Reference M. bigelowi W. Tattersall, 1927 Americamysis Present study M. almyra Bowman, 1964 Americamysis Present study M. bahia Molenock, 1969 Americamysis Present study M. taironana Brattegard, 1974 Americamysis Present study M. inermis Coifman, 1937 Brasiliomysis Bacescu (1968) M. elongata Holmes, 1900 Metamysidopsis W. Tattersall (1951) M. munda Zimmer, 1918 Metamysidopsis W. Tattersall (1951) M. pacifica Zimmer, 1918 Metamysidopsis W. Tattersall (1951) 686 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Wa Ye LQ tp Fig. 1. Americamysis alleni, new species. A, adult female, dorsal view; B, antennular peduncle; C, antennal peduncle and scale; D, labrum; E, mandibles; F, mandibular palp; G, maxillule; H, maxilla; I, endopod, thoracic limb 1; J, K, endopod, thoracic limb 2, female, male, respectively. Scales in mm. VOLUME 107, NUMBER 4 0.37 E 1.40 687 Fig. 2. Americamysis alleni, new species. A, endopod, thoracic limb 3; B, endopod, thoracic limb 8; C, male pleopod 1; D, male pleopod 4; E, uropod; F, G, telsons. Scales in mm. ner lobe with 2 large serrate apical setae subequal in length. Maxilla (Fig. 1H): typical of genus; exo- pod with 3-6 plumose setae; endopod 2-articulate, distal article armed with 2-3 submarginal and 16-24 plumose setae on apex and inner margin; lobe of sympodal article 2 with 5 plumose setae; sympodal article 3 bilobed, inner lobe with 10-15 plu- mose setae, outer lobe with 8—10 plumose setae. Endopod of thoracic limb | (Fig. 11): ro- bust typical of genus; carpo-propodus twice length of dactyl, distal part with numerous simple setae and serrate spine-setae; dactyl wider than long with numerous simple se- tae, serrate spine-setae and a single large claw, slightly longer than dacty]. 688 Endopod of thoracic limb 2: (Fig. 1J, K): large and robust, more developed in females than in males; ischium about '2 length of merus with several simple setae on inner margin; merus subequal in length with car- po-propodus, with 2-3 simple setae on in- ner proximal margin, 4 simple setae along outer margin; carpo-propodus with 5-10, occasionally 4, flared spiniform setae and numerous simple setae on distal '2 to % of inner margin; dactyl ‘45 length of carpo- propodus armed with numerous curved, serrate spine-setae, simple setae and a ser- rate distal claw. Endopod of thoracic limb 3 (Fig. 2A): ba- sis with 2 plumose setae; ischium 1.5 times as long as merus, setose along inner margin; merus setose along inner margin with ser- rate seta and simple seta on outer distal margin; 2-segmented carpo-propodus slightly shorter than merus with segments subequal in length, proximal segment with 1 small submarginal serrate seta and 1 large serrate seta on outer distal margin; dactyl small with long, slightly curved claw on dis- tal tip surrounded by several simple setae extending from distal margin of the carpo- propodus. Endopod of thoracic limb 8 (Fig. 2B): ba- sis with 2 plumose setae; ischium slightly longer than merus with 2 simple setae on distal part of inner margin; merus with many simple setae along inner margin and | ser- rate seta and | simple seta on outer distal margin; 2-articulated carpo-propodus 0.7 times length of merus with articles subequal in length, proximal article with 2 serrate setae on outer distal margin; dactyl with slender claw surrounded by several long, simple setae, extending from distal article of carpo-propodus. Female pleopods reduced to setose uni- ramus plates. Male pleopod | (Fig 2C): 1-articulated endopod with | plumose seta distally, 5 plu- mose setae proximally and pseudobranchial lobe furnished with 5 terminal plumose se- tae; 7-articulated exopod. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Male pleopods 2, 3, 5: exopods and en- dopods 7-segmented. Male pleopod 4 (Fig. 2D): 7-articulated endopod with pseudobranchial lobe similar to first pleopod; 7-articulated exopod, slightly longer than endopod, setule on out- er distal margin of article 7, with long barbed apical spine-setae as long as the 5 distal ar- ticles combined. Uropods (Fig. 2E): exopod 1.3—1.4 times as long as endopod and 1.8—1.9 times as long as telson, outer margin concave, inner margin convex; endopod 1.3-1.4 times as long as telson, inner margin almost straight, outer margin concave, armed with 3-4, oc- casionally 2 or 5 spine-setae near statocyst; exopod and endopod setose along both in- ner and outer margins. Telson (Fig. 2F, G): entire, linguiform, shorter than 6th abdominal segment, 1.4— 1.5 times as long as maximum width, lateral Margins concave and armed with 10-15 short, stout spine-setae; apex with 3 pairs of strong spine-setae, outermost pair 2-4 length of inner pairs, innermost pair %3 or less length of telson and shorter than, or equal in length to, adjacent pair. Coloration.—The following description is of preserved specimens collected in Mis- sissippi Sound, Mississippi. Pairs of black chromatophore located dorsally at base of telson and ventrally at base of mandibles and thoracic legs 3 and 7. Posterior ventral margin of abdominal segments 1-5 with a single black chromatophore. Antennular peduncles mottled brown; male lobe with brown along distal and medial margins. Brown along distal margin of sympod of antenna and on distomedial margin of scale. Females with one chromatophore at base of each posterior oostegite. Distribution. —Inshore and continental shelf waters in the northern Gulf of Mexico from Lavaca Bay, Texas to Mississippi Sound, Mississippi. Habitat.—Hypoplanktonic in meso or polyhaline waters of bays and shallow con- tinental shelf waters to a depth of 15 m. VOLUME 107, NUMBER 4 Etymology.—This species is named for Dennis Allen in recognition of his contri- butions to our knowledge of the biology of mysids along the east coast of North Amer- ica. Americamysis stucki, new species Figs. 3-4 Mysidopsis bigelowi, Brattegard 1969:53, fig. 15.—Farrell 1979:32, figs. 1,2, 1le, f, g.— Stuck et al. 1979:235, figs. 2j, 3), 4j, 5j.— Price 1982:16 (in part), figs. 18, 21. Type material.—Holotype: adult 6 (USNM 253074), length 5.2 mm, SEAMAP station 35056 (30°11.6'N, 88°11.2’W), con- tinental shelf waters off Mississippi, 16 Oct 1981.—Paratypes: 3 6, 42 (USNM 253075), 5 6, 5 2(GCRE 1316), same collection data as for holotype. Additional material examined. —Conti- nental shelf waters off Texas: 3 6 (4.4-5.2 mm), 4 2 (5.7-6.0 mm), NMFS St. 1 (29°22.5'N, 94°34.0'W), depth 8.2 m, 13 Jan 1969.—14 6 (4.3-6.4 mm) 18 2 (4.5-5.0 mm), NMFS St. 17 (28°42.4'N, 94°58.0’W), depth 19.2 m, 14 Jan 1969. West coast of Florida: 10 6 (3.8-4.6 mm), 10 2 (3.9-5.0 mm), Anclote Key St. 15 (28°11.1'N, 82°52.4'W), depth 3.2 m, 23 Jul 1982.—25 6 (3.5—5.1 mm), 25 2 (3.9-5.0 mm), Tampa Bay St. 9 (27°37.3’N, 82°35.3’W), depth 3.5 m, 15 Jun 1983.—10 6 (4.1-4.6 mm), 10 2 (4.0-5.2 mm), offshore St. 5 (27°38.8'N, 82°49.7'W), depth 7.6 m, 3 Oct 1981.—8 6 (3.6—4.8 mm), 8 2 (3.7-4.8 mm), Sarasota Bay St. 8 (27°20.2’N, 82°33.7'W), depth 3.7 m, 7 Jun 1982.—15 6 (3.7-4.1 mm) 15 2 (3.5-4.9 mm), Charlotte Harbor St. 34 (26°43.5'N, 82°15.4'W), depth 3.7 m, 8 Aug 1982.—7 8 (4.8-5.0 mm), 5 2 (4.6-5.5 mm), San Carlos Bay St. 55 (26°27.2'N, 82°0.2’W), depth 5.5 m, 11 Jul 1982.—1 6 (3.8 mm), 1 2 (4.3 mm), Bear Point, Big Marco River St. 63 (25°58.0’N, 81°42.2'’W), depth 3.7 m, 12 Jul 1982. All Texas and Florida material collected with epibenthic sled and in col- lection of WWP. 689 Diagnosis. — Apex of antennal scale not pointed; endopod of thoracic limb 2 exhib- iting sexual dimorphism, in males proximal portion of merus very slender, bearing 4—8 setae along inner margin and 4-6 setae along outer margin, proximal part of merus not as slender in females as in males, inner mar- gin bearing 4—7 setae and outer margin with 3-5 setae; inner margin of carpo-propodus armed with 5-9 spiniform setae; carpo- propodus of thoracic endopods 3-8 2-articulated; exopod of pleopod 4 of male 7-articulated; endopod of uropods armed with 4—5 (rarely 3) spine-setae near stato- cyst; apex of telson armed with 3 pairs of strong, widely spaced spine-setae, outer- most pair % to 4 length of inner pairs, in- nermost pair greater than '4 length of telson and slightly longer to subequal in length with adjacent pair. Description.—General body form (Fig. 3A): moderately slender, adult males to 6.4 mm, females to 6.0 mm; anterior margin of the carapace produced into a short, trian- gular, rostrum reaching to the bases of the eye-stalks; posterior dorsal margin broadly emarginate exposing thoracic segment 8; anterolateral corners rounded. Antennular peduncle (Fig. 3B): more ro- bust in males than females; article 1 about as long as articles 2 and 3 combined, dis- tomedial margin armed with 3-4 long sparsely plumose setae and 3—4 shorter plu- mose setae; distolateral margin with | short and 3-4 long plumose setae; medial margin of article 2 with strong simple spine-seta and 5 plumose setae; article 3 with 2 plu- mose setae along medial margin and group of 6 plumose setae distally, distal margin with dorsomedial lobe bearing 2 tooth-like processes and 4 plumose setae, lateral mar- gin with 1 long and 1 short submarginal plumose seta; males with densely setose lobe on outer ventral surface. Antenna (Fig. 3C): scale lanceolate, 5.0- 5.5 times as long as maximum width, inner margin slightly convex, outer margin slight- ly concave, all margins setose; lacking distal 690 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 2.32 BuJ,K 0.38 C,F 0.27 D,I 0.33 H 0.17 E,G 0.25 Fig. 3. Americamysis stucki, new species. A, adult female, dorsal view; B, antennular peduncle; C, antennal peduncle and scale; D, labrum; E, mandibles; F, mandibular palp; G, maxillule; H, maxilla; I, endopod, thoracic limb 1; J, K, endopod, thoracic limb 2, female, male, respectively. Scales in mm. VOLUME 107, NUMBER 4 article; antennal peduncle extending about */, length of scale, article 2 about 1.5 times as long as 3, having 3 plumose setae on inner distal margin and | plumose seta on outer distal margin, article 3 with 4 simple and 2 plumose setae on inner distal margin and 2-3 short plumose setae along outer margin; distolateral corner of sympod with promi- nent tooth. Eyes (Fig. 3A): large, cornea wider than distal end of eyestalk, lacking ocular papilla. Labrum (Fig. 3D): rounded anteriorly; posterior margin with medial emargination, middle *% with fine setae. Mandible (Fig. 3E): with cutting edges typical of the genus. Palp (Fig. 3F) 3-articulated; article 2 more robust in males than females, armed with 11-12 submar- ginal setae on mesial surface and 1 7—19 sim- ple setae on inner surface, distal margin with 4-6 strong simple setae; article 3 % as long as 2, inner surface with 5—9 marginal and 3—5 submarginal simple setae, distal portion armed with | long simple seta, 1 long curved spine barbed along its middle part, and 9 thick, blunt densely barbed spines. Maxillule (Fig. 3G): outer lobe with 9 stout serrate apical spine-setae; inner lobe with 2 serrate setae subequal in length. Maxilla (Fig. 3H): typical of genus; exo- pod with 4 to 7 plumose setae; endopod 2-articulated, distal article armed with 2-4 submarginal and 20-22 plumose setae on apex and inner margin. Lobe of sympodal segment 2 with 4 long plumose setae; sym- podal segment 3 bilobed, inner lobe with 12-14 plumose setae, outer lobe with 11 plumose setae. Endopod of thoracic limb 1 (Fig. 31): ro- bust, typical of the genus; carpo-propodus twice length of dactyl, distal part with nu- merous simple setae and strong serrate spine-setae; dactyl as long as wide with nu- merous simple setae, serrate spine-setae and single large claw, slightly longer than dactyl. Endopod of thoracic limb 2 (Fig. 3J, K): exhibiting sexual dimorphism; in males (Fig. 3K) merus subequal in length with carpo- 691 propodus, proximal portion very slender, bearing 4-8 simple setae along inner mar- gin, 4 to 6 setae along outer margin and 1 seta on outer distal margin; inner margin of carpo-propodus armed with 5—9 flared spi- niform setae gradually increasing in length distally, and a short submarginal seta at the base of each spiniform seta. In females (Fig. 3J) merus % as long as carpo-propodus, proximal part not as slender as in male, in- ner margin bearing 4—7 setae, outer proxi- mal margin with 3—5 setae and distal margin with | seta; inner margin of carpo-propodus armed with 5-9 flared spiniform setae as in male; dactyl 4 length of carpo-propodus, armed with numerous curved serrate spine- setae and serrate distal claw. Endopod of thoracic limb 3 (Fig. 4A): ba- sis with 3 plumose setae, ischum 1.5 times as long as merus, setose along inner margin; merus setose along inner margin with ser- rate seta on outer distal margin; 2-articulated carpo-propodus slightly shorter than merus with segments subequal in length, proximal segment with | large serrate seta on outer distal margin; dactyl small with long slightly curved claw on distal tip surrounded by sev- eral simple setae extending from distal mar- gin of the carpo-propodus. Endopod of thoracic limb 8 (Fig. 4B): ba- sis with 2 plumose setae; ischium slightly longer than merus with simple setae scat- tered along entire inner margin; merus with many simple setae along inner margin and 1 serrate and 1 simple seta on outer distal margin; length of 2-articulated carpo-prop- odus slightly shorter than merus, articles subequal in length, proximal article with 1 submarginal serrate seta and 2 serrate setae on outer distal margin; dactyl with slender terminal claw and minute setule surrounded by several long simple setae extending from distal margin of carpo-propodus. Female pleopods reduced to setose uni- ramous plates. Male pleopod 1 (Fig. 4C): endopod l-articulate with 1 plumose seta distally, 4— 5 plumose setae proximally and pseudo- 692 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 4. Americamysis stucki, new species. A, endopod, thoracic limb 3; B, endopod, thoracic limb 8; C, male pleopod 1; D, male pleopod 4; E, uropod; F, telson. Scales in mm. branchial lobe furnished with 1 subterminal and 4—5 terminal plumose setae; exopod 7-articulated. Male pleopods 2, 3, 5: exopods and en- dopods 7-articulated. Male pleopod 4 (Fig. 2D): endopod 7-articulated with pseudobranchial lobe similar to first pleopod; exopod 7-articulat- ed, slightly longer than endopod, setule on outer distal margin of article 7, with long barbed apical spine-seta as long as the 5 distal articles combined. Uropods (Fig. 4E): exopods 1.2—1.3 times as long as endopod and 2.0 times as long as telson, outer margin concave, inner margin convex; endopod 1.5—1.6 times as long as telson, inner margin almost straight, outer Margin concave, armed with 4—S (rarely 3) spine-setae near statocyst; exopod and en- dopod setose along both inner and outer margins. Telson (Fig. 4F): entire, linguiform, shorter than 6th abdominal segment, 1.3- 1.5 times as long as maximum width, lateral VOLUME 107, NUMBER 4 Margins concave and armed with 9-1 1 short stout spine-setae; apex armed with 3 pairs of strong widely spaced spine-setae, outer- most pair %4—'4 length of inner pairs, inner- most pair greater than 3 length of telson, slightly longer or subequal in length to ad- jacent pair. Coloration.—The following description is of a population from Tampa Bay, Florida. In living specimens, pairs of black chro- matophore located dorsally at base of tel- son, and ventrally at base of mandibles and thoracic legs 3 and 7. Posterior ventral mar- gin of abdominal segments 1—5 with a single median black chromatophore; when dis- persed, abdominal chromatophore extend laterally and dorsally giving abdomen a mottled brownish appearance; pigmenta- tion from chromatophore of adjacent seg- ments never merges. Abdomen often yel- lowish, ventral half with pink-purple cast. Eyestalks with brown pigmentation dorsal- ly; cornea black. Antennular peduncles usu- ally mottled light brown; inner flagella pig- mented on 10 or fewer proximal segments; male lobe partly brown. Brown along the distal margin of sympod of antenna and on distomedial margin of scale. Carpo-propo- dus and dactyl of thoracic leg | partially or totally brown. Proximal 4 of inner margins of merus of thoracic legs 3-6 brown. Lon- gitudinal brown line extending length of proximal segment of thoracic exopods 2-7. Females with one chromatophore at base of each posterior oostegite. Sympods of male pleopods 1-5 partially brown. Exopod of uropod with 4 or less of distomedial margin lined with brown. Endopod of uropod with Y/, of distomedial margin lined with brown. In preserved specimens the following col- oration persists. Pairs of chromatophore at bases of telson, mandibles, thoracic legs 3 and 7. Single chromatophore on abdominal segments 1—5S, posterior oostegites. Partial pigmentation on male lobe and distomedial margins of uropods. Distribution. — Continental shelf waters in 693 northern Gulf of Mexico between Texas and Alabama. Continental shelf waters to 20 m and inshore waters of west coast of Florida from Anclote Key to Marco Island. Habitat. —Bottom plankton in deeper waters of bays and bottom, mid and surface water plankton in continental shelf waters to a depth of 200 m. Etymology.—This species is named for Kenneth Stuck in recognition of his excel- lent work on malacostracan Crustacea of the Gulf of Mexico. Remarks.—Americamysis alleni, n. sp. and A. stucki, n. sp. are morphologically most similar and apparently most closely related to A. bigelowi. They can be distin- guished from A. bigelowi and each other by the setation of the second thoracic endopod and the telson. Table 3 presents characters that may be used to distinguish the six spe- cies of Americamysis. The following key separates the species of Americamysis and the seven species of Mys- idopsis (excluding M. cojimarensis) that have a 2-articulate carpo-propodus on thoracic endopods 3-8 and a normally developed mandibular palp. As mentioned earlier M. cojimarensis may not belong to Mysidopsis. Key to the Species of Americamysis and the Species of Mysidopsis having a 2-articulate Carpo-propodus on Thoracic Endopods 3-8 1. Lateral margin of telson lacking spine-setae (only terminal spine setae present) — Lateral margin of telson with spine- setae 2. Telson entire with 3 pairs of ter- minal spine-setae; uropodal en- dopod with 2 ventral spine-setae adjacent to statocyst near inner VAG OT See Mecsas WS PAN bee Ak et oe .. Mysidopsis suedafricana O. Tatter- sall, 1969 — Telson weakly cleft with single spine seta on each apical lobe; uro- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 694 i FIV 1 suisvoiour Apdniqe 01-60 Suisvorour Ajdnige TI-01T suiseoiour Apdniqe T I-01 sutsvoiour Ajdniqe CI-TT €I-l I sulseolo suiseorour Ajdniqe -ul Ayjenpei3 Iied qusoelfpe JO yIZUST/UOS]A1 JO 9e19S -ouids jeorde jsow Jouul Jo usu] 9ejas-oulds [e19}e] 0} UOTIL[OI UT 9ejaS-ouIds UOs[9} [eoIde Jo ysUST oe} iG € € € 9-€ 8-p _-as-autds uosjs} [eorde jo sited ‘ony pod € Ajensn ‘p—-Z ¢ 10 Z Ajasel ‘p—¢ ¢ Ajerer ‘C—p p-€ Ajorel ‘¢ €-z Ajjensn ‘¢-] ] -Opuo jepodoin uo ovjes-surds ‘ON p podosjd 9 L i 8-L L L g[ew Jo podoxoe ou saporyie “ON poley polepy Z podopuod o1dvs0y} Jo snpodoid € ‘wo0jruids “QO [—p ‘uLojturds ‘6-¢ wuojiuids ‘7-9 €-7 €-Z -Od1¥d JO UISIVUI JOUUI UO aeIaS “ON Z podopus 919e10Y} Jo Y €-Z 8-7 Z SI-L QI-L SMIoUl JO UISIeUI JOUUI UO Bas “ON podo podo 33 UBY} PP UI IOp podo podo podojoa -[oAop A]]eUuLIOU -JoAop A][euLoU -U9Z[S SIOUW SMIOW -[J2Aop AT[euL0U -[oAop AT[euLIou -op Ajjewi0ou 3 ‘9 JO Z podopus s1ov10y g-€ Spodopus d1dvI0Y} G iG G Z Z Z jo snpodoid-od1ie9 ut sajoryie ‘ON ayeos os 09-S'S ¢S-0'S 09 ¢°9-0'9 ¢°9-0°9 jeuusjue Jo Ones YIpIM/yIdUaT quosqe yuosqe quosqe juosqe quosqe juosqe gjeos [euus}Ue UO JjdILIv [eISIG wn} WINIjsO1 wind}sol wInIjsOo1 winI}sol po -SOI podo[sasp [Tan IeynsueLy “0Ys iepnsueL} “0Ys IvjnsueLy “10ys Ie[NZueLN ‘WoYs -punol A[peoiq govdeieo JO UISIVUI [eSIOp IOLIOWUY i ————— DUDUOLID] soiseds mou soloeds mou 10/2819 piyDpqg Dido si9joeIeYyD ‘1ua[D ‘Pons Nene een Se Se 0 OO eee ee so1seds DT EE ‘sarads sistwupoiuauMp JO sio\oeleYyo 1[Npe oyloeds pue uoWUIUIOD—"¢ 2IQRL VOLUME 107, NUMBER 4 podal endopod with 10-20 ventral spine-setae along inner margin .. ... Mysidopsis bispinulata Brattegard, 1974 . Antennal scale with distal article 4 Antennal scale lacking distal arti- CEE UH SS See Re tte 8 . Tip of antennal scale with setae, not spiniform; uropodal endopod with inner margin wavy, armed with groups of spines inserted be- tween setae Tip of antennal scale without se- tae, acute, spiniform; uropodal en- dopod inner margin not wavy, lacking armed processes along in- ner margin, 18-50 ventral spine- setae extending distally from stato- cyst near inner margin . Area adjacent to statocyst with sin- gle spine-seta; uropodal endopod with inner margin having 6—7 wavy shallow processes each armed with DAES PINES SHIsaee ee Weak. .. Mysidopsis ankeli Brattegard, 1973 Area adjacent to statocyst lacking spine-seta; uropodal endopod with inner margin with 8 wavy pro- cesses each armed with 5-7 spines ....Mysidopsis intii Holmquist, 1957 . Uropodal endopod with more than 45 spine-setae along inner ventral MALoin yee ar AA). tera. se eIM ee ... M. californica W. Tattersall, 1932 Uropodal endopod with 15-25 spine-setae along inner ventral margin . Uropodal endopod having inner ventral surface near posterior mar- gin of statocyst with stout rounded spine-setae producing pad-like ap- pearance; telson with marginal spine-setae gradually becoming larger and longer toward apex ... .. Mysidopsis tortonesi Bacescu, 1968 Uropodal endopod lacking stout rounded spine-setae on inner sur- face near statocyst; telson with api- 10. Wile 695 cal 2 pairs of spine-setae much more robust and distinctly longer than anterolateral pair . . Mysidopsis robustispina Brattegard, 1969 . Rostrum well-developed, extend- ing to distal '4 of first peduncular article of antennule; telson with in- nermost apical pair of spine-setae over 13 longer than adjacent pair Americamysis taironana (Brattegard, 1973) Rostrum not well-developed, not extending past proximal ' of first peduncle article of antennule; tel- son with innermost apical pair of spine-setae subequal, equal, or only slightly longer than adjacent pair 9 . Carpo-propodus of second thorac- ic endopod with 2-3 setae on distal part of inner margin, merus with 7-18 setae on proximal *% of inner IND AG OTD ges payer eg Br eyeesy bl slave, 10 Carpo-propodus of second thorac- ic endopod with series of 4—12 spi- niform setae on distal '2 to % of inner margin, merus with 2-8 spine-setae on inner margin .... 11 Anterior margin of carapace (ros- tral shield) broadly rounded; uro- podal endopod with 1 spine near statocyst; apex of telson of adults usually with 4-8 pairs long slender spine-setae .... Americamysis almyra (Bowman, 1964) Anterior margin of carapace pro- duced to form short triangular ros- trum; uropodal endopod with 1—S (normally 2-3) spine-setae near statocyst; apex of telson of adults usually with 3-6 pairs of long slen- der spine-setae Fao Americamysis bahia (Molenock, 1969) Telson with innermost spine or pair of spines on apex shorter than, or equal in length to adjacent pair, 696 never longer; uropodal endopod usually with 3-4, occasionally 2— 5, spine-setae near statocyst .... Americamysis alleni, new species — Telson with innermost spines slightly longer, never shorter than adjacent pair; uropodal endopod usually with 5 (occasionally 4, rare- ly 3) spine-setae near statocyst... 12 12. Endopod of second thoracic limb normally developed in both sexes, merus (article 5) with 2 setae on proximal inner margin; carpo- propodus with inner margin armed with an irregularly alternating row of short and long spiniform setae ... Americamysis bigelowi (Tattersall, 1951) — Endopod of second thoracic limb with merus in male very slender, merus in both sexes with 4-8 setae spaced along entire inner margin, carpo-propodus with inner margin armed with spiniform setae grad- ually increasing in length distally Americamysis stucki, new species The genus Americamysis is presently known only from the Northwest Atlantic. Its species occur in estuarine and shelf wa- ters along the coasts of the Americas from the northeastern United States to Colom- bia. The type species, A. bigelowi has been reported from off Massachusetts (Georges Bank) southward to Florida and westward along the Gulf coast to Aransas Bay, Texas (Tattersall 1951; Clark 1956; Bowman 1964; Brattegard 1969; Wigley & Burns 1971; Stuck et al. 1979a, 1979b; Stuck & Heard 1981, Modlin 1982, Price et al. 1986). Based on our present studies A. bigelowi appears to be restricted to the east coast of the Unit- ed States. All the Gulf of Mexico records for this species are now referable to either A. alleni or A. stucki, which both appear to be endemic to this region. Americamysis al- leni, an inshore bay species, is currently PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON known from the northwestern and northern central Gulf, whereas, A. stucki, commonly found in higher salinity, near shore waters, is known from southeastern Gulf westward to Texas. Americamysis almyra and A. bahia are closely related species with similar distri- butions, however, the former is generally found in lower salinities than the latter spe- cies. Both are reported from inshore waters along the entire coast of the Gulf of Mexico from Terminos Lagoon, Mexico to the southwestern Everglades in South Florida (Bowman 1964; Brattegard 1969, 1970; Molenock 1969; Price 1978, 1982: Stuck et al. 1979a, 1979b; Escobar-Briones and Soto 1988). The ranges of both species extend northward along the Atlantic east to Patap- sco River, Maryland (Grabe 1981) for A. almyra and to Narragansett, Rhode Island (Lussier et al. 1988) for A. bahia. Americamysis taironana, the only mem- ber of the genus not found in North Amer- ican waters, occurs along the Caribbean coasts of Colombia and Panama (Brattegard 1973, 1974a, 1974b). Acknowledgments We are indebted to the Gulf Coast Re- search Laboratory (GCRL), the United States Environmental Protection Agency (EPA), and Technical Resources, Inc. (TRI) for their support of the 1989 Mysid Tax- onomy Workshop that precipitated the in- terest and need for this study. William Walker (GCRL) and Richard Montogomery (TRI) were instrumental in coordinating the EPA Workshop and initial funding for this research. Thomas E. Bowman kindly ar- ranged for the loan of specimens from the National Museum of Natural History (Smithsonian Institution) and made helpful suggestions at various stages of this re- search. Sara E. LeCroy and Jerry McLelland made constructive comments on the drafts of the manuscript. Ken Stuck, David Burke, Dawne Hard, David Hard, J. McLelland, S. VOLUME 107, NUMBER 4 LeCroy, Chet Rakocinski, Barry Lenz, Craig Hawkinson, and David Camp helped in the collection of specimens and other phases of this work. We greatly appreciate the en- couragement and contributions of K. Stuck, who contributed much insight into the tax- onomy of the ““Mysidopsis complex” during the early phases of this study. We thank Dennis Allen for making information on mysids and specimens of Mysidopsis bige- lowi available to us. The encouragement, constructive comments, and specimens provided by EPA reserach personnel, es- pecially Steve Ward, Suzanne Lussier, Ger- aldine Cripe, and Larry Goodman, were very helpful. This research was funded in part by TRI and the EPA Environmental Monitor- ing and Assessment Program (EMAP) Con- tract No. CR-818218-01-0. The first author was supported through a University of Tampa Faculty Development Grant. Literature Cited Bacescu, M. 1968a. Afromysis guinensis n. sp. and Brasilomysis castroi n.g., n. sp. from the wa- ters of the tropical Atlantic.— Revue Roumaine de Biologie, Série de Zoologie 13(2):75—-86. 1968b. Etude des quelques Leptomysini (Crustaceae Mysidaceae) des eaux du Bresil et de Cuba; description d’un genre et de cing autres taxons nouveaux.—Estratto dagli Annali del Museo civico di Storia Naturale di Genova 77(5): 232-249. 1975. Contributions to the knowledge of the mysid (Crustacea) from Tanzanian waters.— University Science Journal, University of Dar es Salaam 1:39-61. 1984. Katerythrops n. gen., Amathimysis brasiliana n.sp. and Mysidopsis sankarankuttyi n. sp. of the N E. Brazilian littoral waters. — Revue Roumaine de Biologie, Biologie Ani- malia 29:3-7. ——, & L.G. Gleye. 1979. New Mysidacea from Californian waters.—Travaux du Museum d@ Histoire naturelle Grigore Antipa 20:131—-141. ——,, & E. Vasilescu. 1973. New benthic mysids from the littoral waters of Kenya: Mysidopsis kenyana n.sp. and Nouvelia natalensis mobasae N.g., n. sp. — Revue Roumaine de Biologie, Série de Zoologie 18:249-256. Banner, A. H. 1953. Ona new genus and species of mysid from southern Louisiana (Crustacea, 697 Malacostraca).— Tulane Studies in Zoology 1:3-8. Bowman, T. E. 1964. Mysidopsis almyra, a new es- tuarine mysid crustacean from Louisiana and Florida.—Tulane Studies in Zoology 12(1):15— 18. Brattegard, T. 1969. Marine biological investigations in the Bahamas 10. Mysidacea from shallow water in the Bahamas and southern Florida. Part 1.—Sarsia 39:17-106. 1970. Marine biological investigations in the Bahamas 11. Mysidacea from shallow water in the Bahamas and southern Florida. Part 2.— Sarsia 41:1—35. 1973. Mysidacea from shallow water on the Caribbean coast of Colombia.—Sarsia 54:1-66. 1974a. Additional Mysidacea from shallow water on the Caribbean coast of Colombia.— Sarsia 57:47-86. . 1974b. Mysidacea from shallow water on the Caribbean coast of Panama. —Sarsia 57:87-108. Clarke, W. D. 1956. Further description of Promysis atlantica Tattersall (Crustacea, Mysidacea).— American Museum Novitiates 1755:1—5. Coifmann, I. 1937. Misidacei raccolti dalla R. Cor- vetta “Vettor Pisani” nelgli anni 1882-85.— Annuario del Museo Zoologico della R. Univ- ersita di Napoli (Nuova Serie) 7(3):1-14. da Silva, V.M.P. 1979. Mysidopsis juniae nova spe- cies of Crustacea-Mysidacea.— Avulso 30:14. Escobar-Briones, E. & L. A. Soto. 1988. Mysidacea from Terminos Lagoon, southern Gulf of Mex- ico and description of a new species of Taph- romysis.—Journal of Crustacean Biology 8(4): 639-655. Gleye, L.G. 1982. Two new species of Leptomysinid mysids (Crustacea, Mysidacea) from southern California.— Proceedings of the Biological So- ciety of Washington 95:319-324. Grabe, S. A. 1981. Occurrence of Mysidopsis almyra Bowman, 1964 (Mysidacea) in the Patapsco River estuary (Upper Chesapeake Bay), Mary- land, U.S.A.— Proceedings of the Biological So- ciety of Washington 94:863-865. Hansen, H. J. 1913. Report on the Crustacea Schi- zopoda collected by the Swedish Antarctic Ex- pedition, 1901-1903. Copenhagen, 56 pp. Holmquist, C. 1957. Mysidacea of Chile. Reports of the Lund University Chile Expediton 1948-1949 (Report 28).—Lunds Universitets Arsskrift 53: 1-52. Hoffmeyer, M. S. 1993. Mysidopsis rionegrensis, a new species of Mysidacea from San Matias Gulf, Argentina. — Physis 48(114—115):15-19. Lussier, S. M., A. Kuhn, M. J. Chammas, & J. Sewall. 1988. Techniques for the laboratory culture of Mysidopsis spp. (Crustacea, Mysidacea).—En- 698 vironmental Toxicology and Chemistry 7:969— 978. Mauchline, J. 1980. The biology of mysids and eu- phausiids. — Advances in Marine Biology 18:1— 369. Modlin, R. F. 1982. Contributions to the ecology of the mysid crustaceans in the shallow waters of Dauphin Island, Alabama.— Northeast Gulf Science 5:45-49. 1987. Mysidacea from shallow waters in the vicinity of Carrie Bow Cay, Belize, Central America, with descriptions of two new spe- cies.— Journal of Crustacean Biology 7:106—121. Molenock, J. 1969. Mysidopsis bahia, a new species of mysid (Crustacea: Mysidacea) from Galves- ton Bay, Texas. — Tulane Studies in Zoology and Botany 15(3):113-116. Murano, M. 1970. A small collection of benthic Mys- idacea from coastal waters in Suruga Bay, Ja- pan.—Crustaceana 18:251-268. Norman, A. M. 1863. Report of the dredging expe- dition to the Dogger Bank and the coasts of Northumberland. Crustacea.—Transactions of the Tyneside Naturalist’s Field Club 5:263-280. Oritz, M. & R. Lalana. 1993. Mysidopsis cojimar- ensis, una nueva especie de misidaceo marino (Crustacea, Mysidacea) de Cuba.—Caribbean Journal of Science 29(1—2):50—53. Price, W. W. 1978. Occurrence of Mysidopsis almyra Bowman, M. bahia Molenock and Bowmaniella brasiliensis Bacescu (Crustacea, Mysidacea) from the eastern coast of Mexico.—Gulf Research Reports 6:173-175. 1982. Key to the shallow water Mysidacea of the Texas coast with notes on their ecology. — Hydrobiologia 93:9-21. , A. P. McAllister, R. M. Towsley, & M. Del Re. 1986. Mysidacea from continental shelf waters of the northwestern Gulf of Mexico.— Contributions in Marine Science 29:45-58. Sars, G. O. 1864. Beretning om en i Sommeren 1863 foretagen Zoologisk Reise i Christiania-stift.— Nyt Magazin for Naturvidensskaberne 13:225— 260. 1872. Monographi over de ved Norges Kys- ter forekommende Mysider. — Andet Hefte Car- cinologiske Bidrag til Norges Kyster forekom- mende, Christiana 1-34. 1884. Preliminary notices on the Schizopoda of HMS “Challenger” expedition.—Forhand- linger i Videnskabsselskabet i Kristiania 7:1—43. 1885. Report on the Schizopoda collected by HMS “Challenger” during the years 1873 to 1876.—The Voyage of H.M.S. Challenger 13: 1-228. Stuck, K. C., & R. W. Heard. 1981. Amathimysis PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON brattegardi, a new peracarid (Crustacea: Mysi- dacea) from continental shelf waters off Tampa Bay, Florida.—Journal of Crustacean Biology 1:272-278. , H. M. Perry, & R. W. Heard. 1979a. An annotated key to the Mysidacea of the North Central Gulf of Mexico.—Gulf Research Re- ports 6(3):225-238. , H. M. Perry, & R. W. Heard. 1979b. Records and range extensions of Mysidacea from coastal and shelf waters of the eastern Gulf of Mexico. — Gulf Research Reports 6(3):239-248. Tattersall, W. M. 1926. Crustaceans of the orders Euphausiacea and Mysidacea from the western Atlantic. — Proceedings of the United States Na- tional Museum 69(2634):1—31, 2 plates. 1932. Contributions to the knowledge of the Mysidacea of California, I. On a collection of Mysidae from La Jolla, California.— University of California Publications in Zoology 37:301— 314. . 1951. Areview of the Mysidacea of the Unit- ed States National Museum.—Bulletin United States National Museum 201:1-292. Tattersall, O. S. 1955. Mysidacea.—Discovery Re- ports 28:1-190. 1962. Report on a collection of Mysidacea from South Africa off-shore and coastal waters (1957-1959) and from Zanzibar. — Proceedings of the Zoological Society of London 139:221— 247. 1969. A synopsis of the genus Mysidopsis (Mysidacea, Crustacea) with a key for the iden- tificaton of its known species and descriptions of two new species from South African waters. — Journal of Zoology, London 158:63-79. Thompson, G. M. 1894. Ona freshwater schizopod from Tasmania.—Transactions of the Linnean Society of London (Series 2) 6:285-303, 3 pl. Wigley, R. L., & B. R. Burns. 1971. Distribution and biology of mysids (Crustacea, Mysidacea) from the Atlantic coast of the U.S. in the NMFS Woods Hole collection.— Fishery Bulletin 69(4): 717-746. Wooldridge, T. H. 1988. Anew species of Mysidopsis (Mysidacea) from coastal waters of southern Af- rica and a key to the known species from the subcontinent.—Annals of the South African Museum 98:93-103. (WWP) Department of Biology, Univer- sity of Tampa, Tampa, Florida 33606, U.S.A.; (RWH, LS) Gulf Coast Research Laboratory, P.O. Box 7000, Ocean Springs, Mississippi 39566, U.S.A. PROC. BIOL. SOC. WASH. 107(4), 1994, pp. 699-706 ARMADILLIDIUM TABACARUI (ISOPODA: ONISCIDEA: ARMADILLIDIIDAE), A NEW TROGLOBITIC SPECIES FROM A SULFUROUS CAVE IN ROMANIA Magdalena Gruia, Vasilica Iavorschi, and Serban M. Sarbu Abstract.—Four species of terrestrial isopods belonging to Caucasonethes, Haplophthalmus, Trachelipus and Armadillidium were collected in Movile Cave, Romania. Trachelipus troglobius was described recently and Armadillidium tabacarui, a new species, is described here. It was collected in the lower level of the cave where it is present in large numbers on the walls of the cave’s air- bells. It feeds on the rich chemoautotrophic microbiota covering the walls. A diverse troglobitic community was re- cently discovered in Movile Cave in south- ern Dobrogea, Romania (Sarbu & Popa 1992). Thirty-one previously unknown ter- restrial and aquatic species have been iden- tified so far and further discoveries are ex- pected. Most are characterized by an advanced degree of troglomorphy suggest- ing that they have been isolated from the surface for a long time. Four species of ter- restrial isopods (Oniscidea) inhabit Movile Cave: Trachelipus troglobius Tabacaru & Boghean, 1989 (Trachelipidae), Haplo- phthalmus sp. and Caucasonethes sp. (Trichoniscidae). A new species of Arma- dillidium Latreille, 1804 (Armadillidiidae) is described here. Armadillidium tabacarui, new species Fig. 1 Derivation of name. —This new species is named in honor of our colleague, Dr. Ionel Tabacaru, a well-known specialist in terres- trial isopod taxonomy. Holotype. — Adult male, “Emil Racovita”’ Speleological Institute (ERSI), Bucharest, Romania, from Movile Cave, leg. Viorel Boghean, 28 Dec 1990. Paratypes.—Three males; two females, ERSI, from Movile Cave, leg. Serban M. Sarbu and Dumitru Pegulescu, 28 Dec 1990. Distribution. — The lower level of Movile Cave, southern Dobrogea, Romania. Diagnosis.—The new species of Arma- dillidium is typified by the following char- acters: volvation of mesospheric type; smooth white tegument; pleotelson trian- gular with rounded posterior margin; ar- madillidiid type cephalon; frontal plate pro- jects slightly above vertex and is slightly prismatic; frontal fossa hidden by posterior margin of plate; male ischium of pereiopod VII elongate, thickened distally with slightly concave ventral margin with zone of pili- form scales; merus of pereiopod VII strong- ly widened distally; internal lobe of exo- podit of male pleopod 1, pointed and slightly curved laterally; dorsal valvula of stomach, rectangular shaped. Description.—Body length—6.7 mm in males, 6.45 mm in females; Body width— 3.35 mm in males, 3.05 mm in females; Color—white, white-pinkish; Eye— 16-18 small ommatidia, with slight traces of pig- ment (Fig. 1). Tegument: smooth, lacks granulation, with small hyaline semicircular scales (Fig. 2A). Triangular scale-spines with large base and rounded tip on posterior margin (Fig. 2B) and on surface of pereionites. Trian- gular scale-spines, with narrow base and pointed tip (Fig. 2C), on anterolateral sur- 700 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. face of pereionites II-VII and especially on uropods. Simple short spines on surfaces of all pereionites (Fig. 2D). Long acutely point- ed spines along interior margin of pereio- nites (Fig. 2E) and on anterolateral surface of pereionites II-VII, along with simple hairs. Long triangulate flexible hyaline scales forming bundles on anterolateral margin of pereionites IJ-VII (Fig. 2F). Lateral nodes small and inconspicuous (Fig. 2G), located posterolaterally, close to posterior margin; in reference to lateral margin, lateral nodes situated on line extending past posterior an- gle of pereionites; no major differences re- garding position of nodes on all pereionites. Glandular fields of Armadillidium type. Cephalon: rectangular in dorsal view. Scutellum convex, slightly prismatic; ante- rior margins of scutellum forming wide an- gle in dorsal view (Fig. 2H), posterior mar- gin straight. Scutellum triangulate in frontal view (Fig. 21). Subscutellar fossa slightly deepened and widened, hidden by posterior margin of scutellum, bent posteriorly (Fig. 2J). Posterior margin extending very slight- ly above vertex (Fig. 2K). Postscutellar line Armadillidium tabacarui, new species: lateral view. continuing toward eyes, bordered by 2 lat- eral prominent rectangular lobes. Antennal lobes with thick upper margin in frontal view; lobes protrude laterally in dorsal view (Fig. 2L). Frontal line short. Pereion: anterior angle of pereionite I frontally directed, slightly lifted (meso- spheric type). Acute posterior angle, pos- terior margin only slightly concave (Fig. 1). Pereionites II—VII, of different shapes: pe- reionites II-IV decrease progressively in length. Epimeron of tergite IV smallest, with rounded edge. Tergite VII with largest epi- meron, posterior angle 90° (Fig. 1). Pleon: size of epimera decreases from pleonites 3 to 5 (Figs. 1, 3A), posterior angle changing from acute (3) to obtusely rounded (5). Telson: width—length ratio 1.5:1. Telson triangular, rounded tip extending only slightly beyond posterior most part of en- veloping pereionite 5. Telson limited ven- trally by clearly visible carina (Fig. 3A). Antennule: articles of unequal length, basal article longest. Short spine present dis- tally on dorsal convex edge of article 1. Ar- VOLUME 107, NUMBER 4 701 Fig. 2. Armadillidium tabacarui, new species: holotype. A, semicircular scales; B, triangulate scale-spines on surface of pereionites; C, triangulate scale-spines on surface of uropods; D, spine on anterolateral surface of pereionites; E, spine on anterolateral surface of pereionites; F, scales on anterolateral margin of pereionites; G, lateral nodes; H, cephalon, dorsal view; I, same, frontal view; J, same, lateral view; K, scutellum, posterior view; L, scutellum, anterior view. ticle 2 shortest. Apical article bearing 3 rows of aesthetascs (10-12), tip pointed, with 1 aesthetasc (Fig. 3B). Antenna: long (Fig. 3C), reaching poste- rior edge of pereionite I. Flagellum with ar- ticle 2 1.25—1.30x longer than article 1; S— 6 aesthetascs on article 1 (Fig. 3C, b). Mandible: molar compound; 4 setae in setal row on both mandibles; seta and hair- like setae on lacinia mobilis (Fig. 3D). Maxillule: outer ramus with 10 apical teeth, outer 4 plain, inner 6 thin with bi- or tri-lobed tip (Fig. 3E). Inner ramus tipped with 2 subequal penicillate setae. Maxilliped: endite longer than wide, with 3 short spines on upper edge; especially stout spine on endite surface; palp with 2 large spines on basal article; one on edge and complex of spines (one large) on article 2; apex with scale-like spined tip with several spines (Fig. 3F). Pereiopods: robust (Fig. 4A—C), tegument uniformly covered by semicircular hyaline scales; rod-shaped spines of various sizes and shapes variously placed on pereiopods (Fig. 4B—C, a). Pereiopod I (Fig. 4A): inner surface with 2 regions covered by long, flexible, hyaline hairs (14-15 rows) on carpus, and short hairs with 4 rows of spines on propodus, both regions bordered on both sides by rows of moderately long setae (Fig. 4A, a). Sternal edge with different shaped rods (Fig. 4A, b— c), forming two rows on carpus and one row of four rods on propodus. Uropods: length—width ratio of basis 1:2; 702 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 3. Armadillidium tabacarui, new species: holotype. A, telson; B, antennule; C, antenna: a, apex ofantennal flagellum;b, aesthetasc on flagellar article 1; c, terminal spine of peduncular segment 5; D, left mandible; E, outer ramus of maxillule; F, maxilliped. exit region from basis of flattened wide ex- opod oblique; exopod with medial margin rounded and lateral corner pointed (Fig. 4D); endopod long and narrow, extending slight- ly past tip of telson (Fig. 3A). Stomach (Fig. 4E, F): dorsal lamella of stomach in A. tabacarui n.sp. rectangular. Dorsal lamella short and wide, with lateral margins about half as long as posterior and basal margins. Posterior margin sinuous, with central excavation (Fig. 4F). Dorsal la- mella covering only 4 of stomach length. Lateralium exhibiting complex three di- mensional structure: triangulate in dorsal view, and anteriorly covered by dorsal val- vula. Upper anterior margin covered by spines. Internal interior surface hyaline and covered by short thin hairs (Fig. 4E, a). Up- per plate with well developed tritulating part; anterior zone with abundant hairs (Fig. 4E, b). Trituration zone of lower plate with rows of long curved, laterally oriented hairs (Fig. 4E, c). Male sexual characters.—Pereiopod VII (Fig. 4B, C): longest; basis very narrow with few tegumentary formations; ischium elon- gate, wider distally with slightly concave ex- ternal edge and elongate area covered by long hyaline hairs next to inner surface (Fig. 4C); merus narrow proximally, very wide distally especially toward external edge, be- ing 1.5 longer than wide; last 3 segments with characteristic rods (Fig. 4C, a) on outer edge of both surfaces. VOLUME 107, NUMBER 4 703 Fig. 4. Armadillidium tabacarui, new species: holotype. A, pereiopod I, inner surface: a, short spines on surface of pereiopod I; b, rod on outer surface of pereiopod I; c, rod on rostral surface of pereiopod I; d, sensory hair on pereiopod I; B, pereiopod VII, inner surface; C, same, outer surface: a, rod on sternal edge of pereiopod VII; D, uropod; E, stomach: a, lateralium, interior hyaline surface; b, upper plate; c, trituration zone of lower plate; d, filtration zone of lower plate; F, same, dorsal valvula. Pleopod 1 (Fig. 5A): triangular exopodit with convex posterolateral margin, pseu- dotrachea obvious, tip pointed, fringing se- tae on medial and posterolateral margins of point; endopodit elongate and straight, dis- tal extremity slightly curved with 2 un- equally long, curved tips: internal tip longer, with row of foliaceous spines increasing in size distally, extending for short distance along endopod; external tip tiny, curved lat- erally and plain (Fig. 5B). Pleopod 2-5 (Fig. 5C-—F): exopodits pointed with rows of short hairs; endopodit 2 (Fig. 5C) only slightly longer than exopod. Discussion Eight species of Armadillidium have been recorded from Romania. Three are found in Dobrogea: A. jaqueti Dollfus, 1897 (banks of the Danube at Isaccea, in western Dob- rogea), A. traiani Demianowics, 1932 (abundant in Bessarabia, also in Dobro- gea— Radu 1985), and A. vulgare (Latreille) 1804, cosmopolitan (Radu 1985). Armadillidium tabacarui can be included in the vulgare-maculatum group (Strouhal 1927), but a perfect attribution of this spe- cies to any of these groups is difficult to make. Compared to A. vulgare and A. traiani which belong to the vu/gare group and are surface species present in Dobrogea, the subterranean A. tabacarui differs mainly through: the shape of the head (mesospheric type) and through the presence of a pilifer- ous area on the ischium of the pereiopod VII in males. Resemblance with other spe- cies of the vulgare group consists mainly in 704 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Sn Ne ee (WN ~ Fig. 5. Armadillidium tabacarui, new species: holotype. A, pleopod 1; B, same, distal extremity of endopod; C, pleopod 2; D, exopod of pleopod 3; E, exopod of pleopod 4; F, exopod of pleopod 5. the shape of the frontal plate of the cephalon which is slightly prominent close to the level of the vertex (Vandel 1962). The presence of the piliferous area in A. tabacarui suggests a close resemblance to A. delattini Verhoeff, 1943 which inhabits the Marmarean region. The broad telson of A. tabacarui resembles that of A. traiani and especially A. absoloni Stroughal, 1939 (present in caves in Bosnia and Herzegovina), as well as A. azerbaid- zhanum Schmalfuss, 1990 from Azerbaijan and the northern Caucasus (Schmolzer 1965, Schmalfuss 1990). The presence of several characters sug- gests that A. tabacarui may belong to the maculatum group. These characters are: the mesospheric type of the shape of the body; the triangulate shape of the telson; the shape of the ischium of the pereiopod VII in males. The later character is present in A. banati- cum Verhoeff, 1907, an endemic species for western Romania (Radu 1985). The com- parison of A. tabacarui with species of the maculatum group present in the Balkan re- gion such as A. klugi Brandt, 1883 and A. bulgaricum Frankenberger, 1941, reveals additional affinities with this group such as the shape of the pleopod I in males, as well as differences such the shape of the merus of the pereipod VII in males (Frankenberger 1941). The shape of the frontal plate is the main character that determines the imper- fect fit of A. tabacarui into the maculatum group as described by Vandel (1962). The anatomical analysis of the stomach in our new species reveals differences in the shape of the dorsal lamella and the later- alium compared to A. vulgare. In the latter the dorsal lamella has a deeper central ex- cavation on the posterior margin and the interior hyaline surface has a rounded me- dian margin as opposed to 4. tabacarui where this margin is straight. The comparative analysis of the charac- VOLUME 107, NUMBER 4 ters shows that A. tabacarui presents a mix- ture of characters which makes its precise attribution to a certain group of species dif- ficult. The additional comparison of A. taba- carui with taxa from Greece and Turkey, such as those described by Schmalfuss (1981, 1982, 1985), Strouhal (1937, 1956), and Vandel (1980), shows affinities of our new species with eastern mediterranean forms. Ecological Observations In the lower level of the cave, the surfaces of the water and the cave walls are covered by microbial mats consisting of heterotro- phic and chemoautotrophic sulfur-oxidiz- ing microbes (Sarbu & Popa 1992). All four oniscids were found exclusively in the vi- cinity of the sulfurous pools, especially in the air-bells of the lower level of the cave (Tabacaru & Boghean 1989). They feed on the microbial mats that cover the walls. Chemoautotrophically produced food (Sar- bu & Popa 1992) is in such abundance that A. tabacarui reaches densities of several hundred specimens per square meter in the air-bells. On several occasions A. tabacarui was seen being eaten by the clubionid spider Lascona cristiani and by the large centipede Cryptops anomalans. The physico-chemical parameters of the cave’s atmosphere are highly stable without marked diurnal or annual fluctuation. The air temperature is 20.9°C (+0.3) and the relative humidity 100%. The atmosphere of the air-bells is low in oxygen (7—10%) and rich in carbon dioxide (2.5—3%). It contains up to | ppm hydrogen sulfide and 1% meth- ane. Acknowledgments The authors are grateful to Viorel Bogh- ean and Dumitru Pegulescu for their help with collecting the material and to all those who made the collecting trips possible. We would like to express special thanks to Dr. G. Schultz, Dr. S. Taiti, and Dr. H. Schmal- 705 fuss for their suggestions regarding the final preparation of the manuscript. This re- search was supported by the “Emil Racov- ita”’ Speological Institute in Bucharest, Ro- mania, the Dept. Biological Sciences of the Univ. of Cincinnati, the National Geo- graphic Society (4639-91), the Cave Re- search Foundation, the Explorers Club, the National Speleological Society, and the Sig- ma Xi Scientific Research Society. Literature Cited Frankenberg, Z. 1941. Prispevek ke znalosti fauny Bulharskych Isopod. Entomologické listy. — Fo- lia Entomologica 4:1—10. Radu, G. V. 1985. Fauna RSR; Crustacea; Ordinul isopoda; Crinochaeta. —Editura Academiei RSR, Bucuresti 4(14), 154 pp. Radu, V. V. 1961. Noi contributii la studiul anato- miei comparate a stomacului la isopodele ter- estre. Cercetari la specii de Platyarthrus. —Co- municarile Academiei RSR 11:1195—1202. Sarbu, S. M., & R. Popa. 1992. A unique chemo- autotrophically based cave ecosystem. Pp. 635- 666 in A. I. Camacho, ed., The natural history of biospeleology, monografias Museo Nacional de Ciencias Naturales, Madrid, 680 pp. Schmalfuss, H. 1981. Die Landisopoden (Oniscidea) Griechenlands. 2. Beitrag: Gattung Armadilli- dium, Teil I (Armadillidiidae). —Spixiana 4:275- 289. . 1982. Die Landisopoden (Oniscidea) Griech- enlands. 3. Beitrag: Gattung Armadillidium, Teil II (Armadillidiidae).—Spixiana 5:217—230. . 1985. Die Landisopoden (Oniscidea) Griech- enlands. 6. Beitrag: Gattung Armadillidium, Teil III (Armadiliidae).—Sitzungsberichten der Os- terreichischen Akademie der Wissenschaften Mathematische- und naturwissenschaftliche Klasse Abteilung I 193:289-301. 1990. Land-Isopoden aus dem Kaukasus- Gebiet. 3. Porcellionidae, Armadillidiidae, Ar- madillidae.—Stuttgarter Beitrage zur Natur- kunde. Serie A (Zoologie) 144:1-11. Schmolzer, K. 1965. Ordnung Isopoda (Landasseln). Pp. 1-186 in Bestimmungsbicher zur Boden- fauna Europas. Akademie-Verlag, Berlin. Strouhal, H. 1937. Isopoda terrestria. II. Armadil- lidiidae, Armadilidae. Pp. 45-65 in M. Beier, ed., Zoologische Forschungsreise nach den Ion- ischen Inseln und den Peloponnes. X VIII. Teil.— Sitzungsberichten der Osterreichischen Akade- mie der Wissenschaften Mathematische- und 706 naturwissenschaftliche Klasse Abteilung I 146: 1-65. . 1956. Isopoda terrestria. II. Armadillidiidae, Armadilidae. Pp. 585-618 in M. Beier, ed., Zoologische Studien in West-Griechenland, VI. Teil.—Sitzungsberichten der Osterreichischen Akademie der Wissenschaften Mathematische- und naturwissenschaftliche Klasse Abteilung I 165:1-618. Tabacaru, I., & V. Boghean. 1989. Découverte, en Dobrogea (Roumaine) d’une espéce troglobie du genre Trachelipus (Isopoda, Onioscoidea, Trachelipidae).— Miscellanea speologica Ro- manica 1:53-75. Tomescu, N. 1974. Morfologia stomacului la citeva speci de isopode terestre. — Studia Universitatis “Babes-Bolyai”’, seria Biologie 1:77-83. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Vandel, A. 1962. Faune de France. Isopodes ter- restres (deuxieme partie).— Le chevalier 66:417— 931. Vandel, A. 1980. Les Isopodes terrestres recueillis en Turquie orientale et en Irak occidental par le Prof C. Kosswig.— Bulletin de la Societé d’His- toire Naturelle de Toulouse 116:83-119. (MG, VI) Institutul de Speologie “Emil Racovita’”’, Str. Frumoasa 11, Bucuresti 78114, Romania; (SMS) University of Cin- cinnati, Department of Biological Sciences, Cincinnati, Ohio 45221-0006, U.S.A. PROC. BIOL. SOC. WASH. 107(4), 1994, pp. 707-720 TWO NEW SPECIES OF PODOCERUS LEACH (CRUSTACEA: AMPHIPODA: PODOCERIDAE) FROM BERMUDA Adam J. Baldinger and Michael F. Gable Abstract. —Two new species, Podocerus tachyrheo and Podocerus lazowasemi, are described from Bermuda. Podocerus tachyrheo is a non-carinate, highly pigmented species lacking uropodal interramal spines; P. Jazowasemi possesses distinct uropodal interramal spines and dorsal carinations, but usually lacks pigmentation. In Kunkel’s (1910) monograph on the amphipods of Bermuda no specimens of the genus Podocerus were recorded. Johnson (1986) and Gable et al. (1988), however, both refer to the existence of a single species of Podocerus in Bermuda. Examination of collections at the Yale Peabody Museum (YPM) and at the National Museum of Nat- ural History (USNM) clearly documents the presence of two species of Podocerus in Ber- muda, both of which are newly described in this paper. In the figures, body parts are marked by abbreviations as follows: A, antenna; Gn, gnathopod; LL, lower lip; Md, mandible; Mx, maxilla; Mxpd, maxilliped; P, pereo- pod; T, telson; U, uropod; UR, urosome. Podocerus tachyrheo, new species Figs. 1-5 Podocerus sp.—Johnson, 1986:378, fig. 125.—Gable et al., 1988:148-149. Etymology. —Named for the ability of in- dividuals of this species to live in areas of extremely swift (G. tachy-) currents (G. rheo), such as those at The Flatts, Bermuda, where all tidal flow from Harrington Sound moves through one narrow channel. Diagnosis. —Pereon without dorsal cari- nations. Pereon segments 5—7 with dorsal and lateral spine groups, pleon segments 1-— 2 with dorsal spine groups only. Adult males and females with varying degrees of pig- mentation. Coxal plates 1-5 with a strong distal spine. Male antennae 1-2, gnatho- pods 1-2, maxillae 1-2, mandibular palp, and maxilliped with plumose setae. Male antenna 2, flagellum 3-articulate and or- namented with submarginal spines. Female antennae 1-2, gnathopod 1, maxillae 1-2, mandibular palp, and maxilliped with plu- mose setae. Male gnathopod 2, article 5 masked by articles 4 and 6, palm with ir- regular margin bearing a proximal conical tooth and a distal rectangular process. Ar- ticle 6 of all pereopods with strong bifurcate spines. Uropods 1-2 with bifurcate spines but lacking peduncular interramal spines. Description.—Male: Body and append- ages with pigmentation, body lacking dorsal carinae, pereon segments 5—7 and pleon seg- ments 1-2 with dorsal distoposterior spine groups, pereon segments 5-7 also with lat- eral distoposterior spine groups. Coxae re- duced with serial discontiguity, coxae 1-5 each with a large marginal spine, located anteriorly on coxae 1-4. Head less than pe- reonites 1 and 2 in length, cuboidal with ocular bulges. Eyes pigmented. Antenna 1, 33% of total body length, pe- duncular ratio 1:3:2.7, flagellum 5-articulate; accessory flagellum 1-articulate and prom- inent. Antenna 2 greater than antenna | in 708 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. length, articles 4 and 5 subequal, flagellum 3-articulate with distinct submarginal spines. Mandible with 3-segmented palp, pen- ultimate article triangular with marginal se- tae, terminal article clavate with facial and apical setae; incisor and lacinia both with 4 teeth; spine row of 3 short, broad spines; molar normal. Maxilla 1, inner plate ves- tigial; outer plate with 9 strong apical spines; palp 2-articulate, terminal article with 3 apical, medially bulging spines and | apical seta, 4 submarginal setae, and 4 facial spines. Maxilla 2, outer plate with 10 long apical setae; inner plate with 10 apical setae, inner margin with fine setae distally. Maxilliped, inner plate with marginal and submarginal setae and a single outer marginal spine; out- Podocerus tachyrheo, male, 4.0 mm, YPM No. 9239. er plate reaching 50% length of palp, with an inner marginal spine row and submar- ginal setae; palp 4-articulate, terminal ar- ticle triangular. Lower lip, normal. Gnathopod 1, coxal plate rhomboidal with a distinct distoanterior spine; article 6, palmar margin longer than hind margin; dactyl with 3 marginal setae. Gnathopod 2 heavily pigmented, robust, and much larger than gnathopod 1; article 5 hidden behind articles 4 and 6; palm of article 6 with ir- regular margin, bearing a proximal conical tooth and a distal rectangular process, mar- gin heavily setose with 2 proximal spines. Bases of pereopods 3-7 with posterior plate- like extensions; article 6 of all pereopods with distally bifurcate spines. Uropods 1 and 2 elongate, biramous, VOLUME 107, NUMBER 4 709 o — << | AN Poce LEN \ \\ Nes Sah OWihe a Hall , Cf Ai} xy Fig. 2. Podocerus tachyrheo, male, 4.0 mm, YPM No. 9239: Al, Gnl, Gn2, UR (with left U1, right U2, right and left U3, T). Male, 2.9 mm, YPM No. 9242: A2. 710 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 3. Podocerus tachyrheo, male, 4.1 mm, YPM No. 9241. VOLUME 107, NUMBER 4 Fig. 4. Podocerus tachyrheo, male, 2.9 mm, YPM No. 9242: P3—P4. Male, 4.0 mm, YPM No. 9239: P5-7. lacking interramal spines; peduncles and rami with marginal bifurcate spines, rami with distinct apical spines. Uropod 3 leaf- like with 3 apical setules. Telson dorsally produced and armed with 4 long setae and 6 short setules. Female: All features same as those for male except as noted. Body lacking dorsal spine groups. Antenna 1, 40% of total body length; flagellum 4-articulate. Flagellum of antenna 2 lacking distinct submarginal spines. Gnathopod 1, article 6, proximal corner of palm demarcated by a spine. Gnathopod 2 resembling gnathopod 1 but twice the size and less ornamented. Remarks.—The degree and patterns of pigmentation are variable among individ- uals of P. tachyrheo; juveniles, smaller males, and females may completely lack pigment. Larger males appear to have a uni- form pigment band along the musculature of the pereon segments. Pigment has also been observed in males on almost every body segment, even on the most distal an- tennal segment. In females, however, pig- mentation appears to become much denser on the ventral margin of the pereon, and may extend to the basal portions of the oos- tegites. Male/female differences in body spi- nation, in the flagella of both antennae, in both gnathopods, and in pigmentation ob- viously make P. tachyrheo sexually dimor- phic. Podocerus tachyrheo is morphologically most closely related to Podocerus multis- pinis K. H. Barnard, 1925 and Podocerus multispinis var. levis K. H. Barnard, 1925. The only major difference between P. mul- tispinis and P. multispinis var. levis is the almost complete absence of the dorsal spi- niform tubercles in the variety (Barnard 1925). Based upon the description and one illustration of gnathopod 2 by both Barnard (1925) and Griffiths (1976), the most sig- nificant differences between P. tachyrheo and P. multispinis are: the number of flagellar 712 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON aA ’ : r ¥ -— ~~ Long, slender and sinuous 1 tooth on movable finger, 2 back- ward curving teeth on movable finger Trilobed (inclusive of external or- bital angle) first epibranchial lobe large, with 2 small lobes on either side, lobes acute to sub- acute > Sakai (1961:144) described the male abdomen in L. hatagumoana as consisting of ““seven free segments. . . .” Whether the segments are freely movable or fused but with visible sutures can only be ascertained when the type specimens are examined. In all other species of Lybia, including the new species, L. tutelina, segments 3— 5 are fused. with segments 3-5 fused, sutures faint; dis- tal end of segment 6 bilobed; telson with truncated apex. G1 stout, 2.3 times longer than G2, distal portion sparsely covered with long setae and conical spines, apex bifur- cated, with petaloid terminal process; G2 with pointed distal process, short spines subterminally. Paratype female.—Female similar to male, except for margins of second and third anterolateral lobes more distinctly dentic- ulated. Etymology.—The Latin ‘“‘tutelina’”’ is a general term for any guardian deity, and is here used as a noun in apposition. Discussion There are certain difficulties in placing this new species in the existing species al- liances suggested by Guinot (1976) and Se- réne (1984). From the figures given by Sakai (1961:142, figs. 2a—d; 1965:162, pl. 80, fig. 1; 1976:pl. 180, fig. 3), L. hatagumoana ap- pears related to L. tutelina, in terms of the sub-globular carapace which is only slightly longer than broad, the fingers of the cheliped being slightly longer than the palm, and the dentition of the cheliped fingers. However, L. tutelina has a much stouter, straighter and shorter G1 relative to G2, whereas in L. hatagumoana, the G1 is longer, slender and sinuous. In this respect, L. tutelina re- sembles L. plumosa (see Guinot 1976:94, fig. 21A—-C) and L. Jeptochelis (see Guinot 1976:94, fig. 21E, F). Differences between L. tutelina and L. hatagumoana are tabu- lated in Table 1. L. hatagumoana appears to be a rare spe- cies, being originally recorded from Amai- daba, off the coast of Hayama, by Sakai (1961) and thereafter only from the Wa- kayama Prefecture, Japan, by Nagai (1990). The male holotype of L. tutelina was car- rying an anemone in each chela as is typical of crabs in this genus. The ovigerous para- type female carried only one anemone (in the left chela), the right chela was free. Acknowledgments The authors wish to thank Drs. A. Cros- nier and B. Richer de Forges for loan of specimens from the ORSTOM collections. This work was partially supported by grant RP900360 from the National University of Singapore. Drs. Peter Davie and Daniele Guinot kindly reviewed the manuscript. 742 Literature Cited Alcock, A. 1896. The Brachyura Oxystomata: ma- terials for a carcinological fauna of India, No. 2.—Journal of the Asiatic Society of Bengal 65(II: 2)134-296, pls. 6-8. Barnard, K. H. 1947. Descriptions of new species of South African decapod Crustacea, with notes on synonomy and new records. — Annals and Mag- azine of Natural History, series 11, 13:361-392. Dana, J. D. 1851. On the classification of the Can- croidea.— American Journal of Science and Arts, series 2, 12:121—131. Guinot, D. 1976. La superfamille des Bellioidae et trois sous-familles de Xanthidae (Polydectinae Dana, Trichiinae de Haan, Actaeinae Alcock): Constitution de quelques groupes naturels chez les Crustacés Décapodes Brachyoures, I.—Meé- moirs du Muséum national d’Histoire naturelle (Paris), new series A (zoology), 97:1—308, figs. 1-47, pls. 1-19. Latreille, P. A. 1812. Crustacés et Insectes. In: J. Milbert, Voyage pittoresque a I’lle-de-France, au Cap du Bonne-Espérance et a I’Ile de Téné- riffe, t. 2. Paris, Le Normant.—Crustacés: 270- 280. MacLeay, W.S. 1838. On the Brachyurous decapod Crustacea brought from the Cape by Dr Smith. Pp. 53-71, pls. 2, 3 in Illustrations of the An- nulosa of South Africa; being a portion of the objects of natural history chiefly collected dur- ing an expedition into the interior of South Af- rica, under the direction of Dr Andrew Smith, in the years 1834, 1835 and 1836; fitted out by “Cape of Good Hope Association for Exploring South Africa.”” London. Milne Edwards, H. 1834. Histoire naturelle des Crus- tacés, comprenant l’anatomie, la physiologie et la classification de ces animaux. Paris 1:468. . 1837. Histoire naturelle des Crustacés, com- prenant l’anatomie, la physiologie et la classi- fication de ces animaux. Paris 2:532. Atlas [1834, 1837, 1840]: 32 pages, pls. 1-14, 14bis, 15-25, 25bis, 26-42. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Nagai, S. 1990. Brachyuran fauna of Wakayama Pre- fecture IV.—Nankiseibutu, the Nanki Biologi- cal Society 32(2):115-119. Nobili, G. 1906. Diagnoses préliminaires de 34 es- péces et variétés nouvelles et de 2 genres nou- veaux de Décapodes de la Mer Rouge. — Bulletin du Muséum d’Histoire naturelle, Paris, 11(6), 1905 (1906):393-411. Sakai, T. 1961. New species of Japanese crabs from the collection of His Majesty the Emperor of Japan.—Crustaceana 3:131-150, 1 pl. 1965. The crabs of Sagami Bay. East-West Centre Press, Honolulu, 206 pp., 100 pls. . 1967. Notes from the carcinological fauna of Japan III.— Researches on Crustacea 3:68-83. 1976. Crabs of Japan and the adjacent seas. Kodansha Ltd, Tokyo, 725 pp., 151 pls. Seréne, R. 1968. The Brachyura of the Indo-West Pacific region. Jn Prodromus for a checklist of the nonplanctonic marine fauna of Southeast Asia.—Singapore National Academy of Sci- ences, special publication 1:33-112. 1984. Crustacés Décapodes Brachyoures de l’Océan Indien Occidental et de la Mer Rouge, Xanthoidea; Kanthidae et Trapeziidae, Faune Tropicale, XXIV, Paris, 349 pp. Takeda, M., & S. Miyake. 1970. Crabs from the East China Sea. IV. Gymnopleura, Dromiacea and Oxystomata.—Journal of the Faculty of Agri- culture, Kyushu University 16:193-—235, pl. 1. Ward, M. 1933. New genera and species of Marine Decapoda, Brachyura from the coasts of New South Wales and Queensland.— Australian Zo- ologist 7(5):377-394. Zehntner, L. 1894. Crustacés de l’Archipel malais.— Revue Suisse de Zoologie et Annales du Musée d’Histoire Naturelle de Genéve 2:135-—214, pls. 7-9. Department of Zoology, National Uni- versity of Singapore, Lower Kent Ridge Rd, Singapore 0511, Republic of Singapore. PROC. BIOL. SOC. WASH. 107(4), 1994, pp. 743-750 THREE NEW RARE HETEROKROANIA SPECIES (CHAETOGNATHA) FROM DEEP BENTHIC SAMPLES IN THE NORTHEAST ATLANTIC Jean-Paul Casanova Abstract.— Three new rare chaetognath species of the genus Heterokrohnia, each represented by a single specimen, are described from deep benthic samples, two off northwest Africa (H. angeli and H. discoveryi) and one in the Bay of Biscay (H. biscayensis). They differ by many characters from hitherto known species, most important among these the number and shape of teeth. The diversity of the benthoplanktonic chaetognath fauna is probably as high as that of the planktonic one. Chaetognath species of the family Het- erokrohniidae are known to live in the water layer just above the sea bed (Casanova 1986a). Most of them have been described from the large R. V. Discovery collections: seven from deep near-bottom planktonic samples and another (Heterokrohnia mir- abiloides Casanova & Chidgey, 1990) from benthic sledge samples. However, the gears used, trawls or sledges, are poorly adapted to catch these fragile animals, which thus are often more or less damaged. Neverthe- less, even if slightly twisted, a single speci- men may be described as a new species if its characteristics are so particular as to avoid any confusion with previous known species. This is the case for the three new species described below. Heterokrohnia angeli, new species Figs. la, 2a, 3a—d Material examined. — Discovery St. 8976, BN2-4, 5 Aug 1976, 32°54.4’N, 11°38.5’W, 3610-3646 m, holotype (Natural History Museum, London, 1994. 2095). Description. —The specimen is 6 mm in body length without tail fin. Tail constitutes 40% of this length. Body stumpy (Fig. 1a) and opaque. Head triangular with a small apical gland cell complex. Anterior teeth, 8/9, short and conical (Figs. 2a, 3a, d). Posterior teeth, 13/ 14, slightly longer (Figs. 2a, 3b, c). All teeth with apical part differentiated (Figs. 2a, 3c, d). Hooks not numerous, 9 on each side. Vestibular organs oval, thin and smooth (Figs. 2a, 3c). Eyes absent. Corona ciliata and glandular neck canals not observed. Neither collarette (very probably stripped off), nor gut diverticula present. Transverse musculature very thin and difficult to ob- serve (for this purpose the body has been cleared with lactic acid and stained with methylene blue); it extends from neck to slightly beyond the large ventral ganglion in trunk and in about the first sixth of tail. Lateral fins begin beyond the ventral gan- glion, at a distance less than half the gan- glion length. Tail fin damaged. All fins with numerous rays. Ovaries not developed. Seminal vesicles large, opening posterola- terally, in contact with both lateral and tail fins. Comparisons with other species.—The shape of teeth immediately differentiate H. angeli from all known Heterokrohnia spe- cies but one, H. bathybia Marumo & Kitou, 1966 from southern Japan. Indeed the latter has both anterior and posterior teeth de- scribed as “‘thick and short’’, but their ex- tremity is not differentiated and they are less PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 744 ~~ QQ Ly Se is ms LLL a, Heterokrohnia angeli, new species in ventral view; b, Heterokrohnia discoveryi, new species in dorsolateral view; c, Heterokrohnia biscayensis, new species in ventral view. AG = apical gland cell complex, Fig. 1. transverse musculature, VG = ventral = ovaries, SV = seminal vesicles, TM GC = glandular canals, Ov ganglion. VOLUME 107, NUMBER 4 745 b Fig. 2. Teeth and vestibular organs of the three new Heterokrohnia species: a, H. angeli in ventral view; b, H. discoveryi in ventrolateral view; c, H. biscayensis in lateral view. AT = anterior teeth, PT = posterior teeth, VO = vestibular organs. numerous at comparable size: respectively 1-2 and 0-4. Other characters distinguish the Japanese species from H. angeli, for in- stance its notched vestibular organs or its very short ventral ganglion. But it must be noted that the presence of a collarette in H. bathybia cannot be regarded as a specific difference since this fragile tissue is often lost during sampling. Etymology. —This species is named after Dr. Martin V. Angel who gave me this spec- imen. Heterokrohnia discoveryi, new species Figs. 1b, 2b, 3e-g Material examined. — Discovery St.10141 N° 1, SBN, 3 Oct 1979, 24°34.8'N, 19°40.7’'W, 3460-3470 m, holotype (Nat- ural History Museum, London, 1994. 2096). Description. —Another small species: length 7.1 mm without tail fin. Tail repre- sents 42.2% of body length. Body stumpy (Fig. 1b), with the four longitudinal muscle masses well separated. Head with a small apical gland cell com- plex. Anterior teeth, 4/5, short, regularly in- creasing in size, the two innermost being the stoutest and the outermost hardly vis- ible (Figs. 2b, 3e, g). Posterior teeth, 4 on each side, slightly longer and stout, distant from each other (Figs. 2b, 3f, g), the out- ermost being reduced. Hooks, 14 on each side, gently curved. Vestibular organs bracket-shaped (Figs. 2b, 3g), narrow and posteriorly festooned. Eyes absent. Corona ciliata abraded away. Remnants of collar- ette tissue (small vacuolar cells) on neck. Short glandular canals visible laterally on neck (Fig. 1b), embedded in collarette. Gut without diverticula. Transverse muscula- ture thick, extending from neck to end of ventral ganglion in trunk and in the first sixth of tail. Lateral fins originate posterior to the ven- tral ganglion. Tail fin sheathing deeply the 746 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 3. Heterokrohnia angeli, new species (a—d): a, Left anterior teeth in lateral view; b, Left posterior teeth in lateral view; c, Dorsal view of right vestibular organ and teeth; d, Detail of the anterior tooth shown in c (in c and d, arrowheads indicate the differentiated apical part of teeth). Heterokrohnia discoveryi, new species (e— g): e, Left anterior teeth in lateral view; f, Left posterior teeth (indicated by arrowheads) in lateral view; g, Ventral view of anterior part of head (the left vestibular organ is indicated by an arrowhead). Heterokrohnia biscayensis, VOLUME 107, NUMBER 4 tail extremity. All fins rayed throughout. Ovaries slightly developed. Seminal vesi- cles empty, elongated, laterally open, in contact with both lateral and tail fins. Comparisons with other species.—The shape and number of teeth set H. discoveryi apart from the 13 Heterokrohnia species de- scribed hitherto. But the shape of vestibular organs as well as the outermost anterior teeth very reduced in size may indicate that this species is related to H. furnestinae Casanova & Chidgey, 1987 in which the anterior teeth are all reduced or absent. The latter is easily recognizable by its acute needle-like pos- terior teeth. Further specimens are needed to confirm this claim of relationship. Etymology. —This species is named after the R. V. Discovery on board of which the specimen was caught. Heterokrohnia biscayensis, new species Figs. lc, 2c, 3h—1 Material examined.—St TS0O3 (Cruise ECOFER IV), suprabenthic sledge, 10 May 1991, 44°43.254’N, 2°19.509'W, 2410 m, holotype (Muséum national d’Histoire Na- turelle, Paris, UD 271). Description. —Body length 12.4 mm without tail fin. Tail is 40.7% of this length. Body transparent and relatively rigid. Head without apical gland cells (Fig. Ic). All teeth short and more or less conical, 3/3 anterior and 3/4 posterior (Figs. 2c, 3h, j— 1). Hooks, 12 on each side, very curved dis- tally (Fig. 31). Vestibular organs prominent and papillated on their edge (Figs. 2c, 3}, 1). Eyes absent. Corona ciliata, collarette and glandular canals on neck not observed. Gut diverticula absent. Transverse musculature thin, from neck to end of ventral ganglion — 747 in trunk and in less than the first sixth of tail. Lateral fins begin beyond the ventral gan- glion, at a distance equal to the ganglion length. Dense rays on both lateral and tail fins. Ovaries not developed. Seminal vesi- cles empty, oval, laterally open, well sepa- rated from lateral fins and slightly apart from tail fin. Comparisons with other species. —In this case also, teeth easily differentiate H. bis- cayensis from all its congener except H. bathybia of which part of the head armature has been described above. But the two spe- cies cannot be confused, the latter being rec- ognizable at first sight by its more developed transverse musculature that extends far be- yond the ventral ganglion in the trunk and almost into the anterior quarter of the tail. Etymology. —The name of this species re- calls that it was caught in the Bay of Biscay. Diversity of the deep bottom chaetognath fauna The benthoplanktonic habitat of hetero- krohniids explains why Heterokrohnia mi- rabilis Ritter-Zahony, 1911, which is the least linked with the bottom, remained for so long the sole known species of this family. It was 55 years before the second represen- tative, H. bathybia, was described. When Dawson (1968) found a few specimens of H. mirabilis in bottom trawls and not in plankton hauls above, he suspected that it lives “‘at or very close to the bottom.” I was able to confirm this view when I studied the chaetognaths sampled by the R. V. Discov- ery on St. 9541 off Mauretania, between the surface and 4000 m (i.e., # within 10 m of the sea bed): in the few metres above the new species (h-l): h, Dorsal view of anterior part of head; i, Distal part of hooks; j, Ventral view of left anterior part of head; k, Right anterior teeth in lateral view; 1, Right posterior teeth in lateral view. AT = anterior teeth, H = hooks, PT = posterior teeth, VO = vestibular organs. Magnification: x 110 (a, b, g-l) and x 270 (c-f). 748 me, “Tg “ae OS +d a ‘ PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 4. SEM photographs of head armature. Heterokrohnia sp. 1 from SW of Ireland (a-c): a, Ventral view of head (the left anterior tooth is indicated by an arrow and the place of the posterior ones by an arrowhead) (x75); b, Area of teeth (x 350); c, Detail of the left vestibular organ (x 430). Heterokrohnia sp. 2 from the Bay of Biscay (x 225); d, Anteroventral view of head. AT = anterior teeth, H = hooks, PT = posterior teeth or place of them, VO = vestibular organs. bottom (mab) were four new Heterokrohnia and one species belonging to a new genus Archeterokrohnia (Casanova 1986a, 1986b). Since then, further species have been de- scribed until today 20 species of this family are known: 16 Heterokrohnia, 3 Archetero- krohnia and the curious Xenokrohnia sorbei Casanova, 1993. All but two of these species, Heterokroh- nia involucrum Dawson, 1968 and H. mi- rabilis, which may be found around 1500 mab, never occur > 500 mab. Even the three Antarctic species H. /ongidentata and H. fragilis Kapp & Hagen, 1985 and Archetero- krohnia longicaudata (Hagen & Kapp, 1986), said to be planktonic when caught with plankton nets, are more likely bentho- planktonic since hauls started in every case between 16 and 77 mab. Heterokrohniids are said to be very deep- living organisms. This is only partly true. In fact, two species have been caught at depths less than 1000 m: 800-815 m for H. wishnerae Casanova, 1992 in the Pacific off Mexico (Casanova 1991 as Heterokrohnia sp.) and 677-738 m for Xenokrohnia sorbei in the Bay of Biscay (Casanova 1993). Thus, it appears that the specific diversity VOLUME 107, NUMBER 4 (richness) of benthoplanktonic chaetog- naths is high in deep water, probably com- parable with that of the planktonic species. In almost every near-bottom sample deeper than 700 m, new species are discovered in newly investigated areas. Even in the north- east Atlantic, at present the best studied re- gion, rare new species are wanting to be de- scribed, e.g., two incomplete specimens from benthic hauls: one (Heterokrohnia sp. 1) off southwest Ireland (Discovery st. 9779, 49°20’N, 12°49.5'W, 1404-1398 m), with small delicate bulb-like teeth and short sau- sage-shaped vestibular organs (Fig. 4a-c); another (Heterokrohnia sp. 2) from the Bay of Biscay (same station as H. biscayensis), which seems to-be related to the species of the /ongidentata group (Casanova 1992) be- cause of its arrangement and shape of teeth (Fig. 4d). They cannot be named today, since they may have relatives with the same type of head armature but differing by other fea- tures. Further research on deep living bentho- planktonic chaetognaths is needed to en- large our knowledge of these interesting spe- cies whose body organization is more variable than those strictly planktonic ones. First, we need a faunistic survey and nu- merical data. Note that in addition to the mew species described or reported above, one genus and two other species are also known from single specimens only: Bathy- spadella edentata Tokioka, 1939, Krohnit- tella tokiokai Bieri, 1974 and Archetero- krohnia longicaudata. Sometimes, this scarcity may be due to the small number of samples realized in an area. But this may be also a reality, since there are abundant and rare benthoplanktonic species as for the planktonic ones. In spite of the maladjust- ment of gears used to catch these organisms, it is obvious that the number of specimens collected is different according to the spe- cies. For instance, in the numerous Discoyv- ery samples in the northeast Atlantic that I observed (both near bottom and epibenthic samples), there were nearly two hundred Heterokrohnia heterodonta Casanova, 1986, 749 about ten H. curvichaeta Casanova, 1986 and only one H. angeli and H. discoveryi. Second we need to describe more fully the structure and function of interesting organs such as the archaic genital apparatus of the heterokrohniids and the curious ventral se- cretory gland (probably digestive) of Xe- nokrohnia sorbei. The discovery of other archaic features in other deep benthoplank- tonic chaetognaths might also be revealed and throw further light on this curious phy- lum. Acknowledgments I am pleased to thank the Curator of the R. V. Discovery collections at IOSDL, K. Chidgey, for sorting out the specimens and providing them on loan. My thanks also to Dr. M.V. Angel for allowing me to study the specimens from the Discovery collec- tions and to Dr. J.-Cl. Sorbe (Laboratoire d’Océanographie biologique, Université de Bordeaux I) for collecting the biscayan spec- imens. Lastly, my thanks to the reviewers who have improved the text. Literature Cited Bieri, R. 1974. First record of the chaetognath genus Krohnittella in the Pacific and description of a new species.— The Wasmann Journal of Biology 32(2):297-301. Casanova, J.-P. 1986a. Quatre nouveaux chaetog- nathes atlantiques abyssaux (genre Heterokroh- nia). Description, remarques éthologiques et biogéographiques.—Oceanologica Acta 9(4): 469-477. . 1986b. Archeterokrohnia rubra n. gen., n. sp., nouveau chaetognathe abyssal de l’Atlantique nord-africain: description et position systéma- tique, hypothése phylogénétique.— Bulletin du Muséum national d’Histoire naturelle, Paris, 4é sér., 8, section A(1):185-194. . 1991. Chaetognaths from the A/vin dives on the Seamount Volcano 7 (east tropical Pacif- ic).—Journal of Plankton Research 13(3):539- 548. . 1992. Chaetognaths from Alvin dives in the Santa Catalina Basin (California), with descrip- tion of two new Heterokrohnia species. —Jour- nal of Natural History 26:663-674. 750 1993. A new genus and species of deep-sea chaetognath from the Bay of Biscay with a strange ventral secretory gland.—Journal of Natural History 27:445-455. , & K. Chidgey. 1987. Une nouvelle espéce d’Heterokrohnia (chaetognathe) des campagnes du “Discovery” dans |’Atlantique nord-orien- tal.—Bulletin du Muséum national d’Histoire naturelle, Paris, 4é sér., 9, section A(4):877-883. 1990. A new benthopelagic species of Het- erokrohnia (Chaetognatha) from the North At- lantic Ocean.—Bulletin ZoGdlogisch Museum Universiteit van Amsterdam 12(8):109-116. Dawson, J. K. 1968. Chaetognaths from the Arctic Basin, including the description of a new species of Heterokrohnia.—Bulletin of the Southern California Academy of Sciences 67(2):112-124. Hagen, W., & H. Kapp. 1986. Heterokrohnia longi- caudata, new species of Chaetognatha from Ant- arctic waters. — Polar Biology 5:181-183. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Kapp, H., & W. Hagen. 1985. Two new species of Heterokrohnia (Chaetognatha) from Antarctic waters.— Polar Biology 4:53-59. Marumo, R., & M. Kitou. 1966. A new species of Heterokrohnia (Chaetognatha) from the West- em North Pacific.—La Mer (Bulletin de la So- ciété franco-japonaise d’océanographie) 4(3): 178-183. Ritter-Zahony, R. Von. 1911. Revision der Chae- tognathen.— Deutsche Stidpolar-Expedition 13(5):1-71. Tokioka, T. 1939. Three new chaetognaths from Jap- anese waters. — Memoirs of the Imperial Marine Observatory 7(1):129-139. Laboratoire de Biologie animale (Planc- ton), Université de Provence, 13331 Mar- seille cedex 3, France. PROC. BIOL. SOC. WASH. 107(4), 1994, pp. 751-759 ANATOMICAL OBSERVATIONS OF THE SAND DOLLAR MELLITA QUINQUIESPERFORATA (LESKE, 1778) (ECHINODERMATA: ECHINOIDEA) AND THE DESIGNATION OF A NEOTYPE Rich Mooi and Antony S. Harold Abstract.—A neotype for the sand dollar Mellita quinquiesperforata (Leske, 1778) is designated and described. Quantitative and qualitative data are given for the test, spines, and pedicellariae of the neotype. Figures of plate and food groove patterns are also given for non-type specimens. The genus Mellita L. Agassiz, 1841 com- prises seven living species. Four occur only on the Pacific side of the Isthmus of Pan- ama, and three on the Atlantic side (Harold & Telford 1990). Mellita quinquiesperforata (Leske, 1778), one of the Atlantic species, is the type species and most widely distrib- uted member of the genus. In spite of its widespread occurrence, and its prominence in both appropriate habitats and shell shops, M. quinquiesperforata has a taxonomic his- tory described as “strange” by Harold & Telford (1990:999). In their recent revision of the genus, Harold & Telford (1990) de- scribed this history, synonymizing Mellita lata Clark, 1940 and Mellita latiambulacra Clark, 1940 with M. quinquiesperforata, but raising M. quinquiesperforata tenuis Clark, 1940 to the status of species. For the Mellita occurring along the eastern coast of the United States, they named a new species, Mellita isometra Harold & Telford, 1990. These actions restricted the name M. quin- quiesperforata to mainland populations oc- curring west of the Mississippi delta along the Central and South American coasts to southern Brazil, and to populations from some Caribbean Islands in association with terrigenous sand substrates. The only reference to the deposition of the specimens in Klein’s (1734) figures of Mellita testudinata (as pre-Linnean, this name is unavailable), upon which Leske’s description of “Echinodiscus quinquies per- foratus”’ is based, is ““Hospitatur in museo Trieriano” (Leske 1778:198). At the time, the practice of designating type specimens was not well established, and it is even con- ceivable that Leske did not have the actual material in hand. Apparently, none of the earlier monographers of the genus (L. Agas- siz 1841, Clark 1940, Mortensen 1948) made attempts to locate and examine Klein’s specimens. Although Harold & Telford (1990:998) did make such attempts, they were “unable to ascertain whether or not these specimens still exist.”? There are cur- rently no museums with natural history col- lections in Trier, Germany. Correspon- dence with the other museums that do exist there has failed to uncover the specimens. The importance of this species to studies in systematics, biogeography, ecology, and physiology compels us to designate a neo- type. The name Mellita quinquiesperforata has been applied to a wide variety of taxa and used in many different senses. In par- ticular, the eastern U.S. fossil Mellita ampla Ravenel, 1848 was placed in synonymy with M. quinquiesperforata by Harold & Telford (1990), but our preliminary examination suggests that MM. ampla should be main- tained as a separate taxon. Therefore, it is imperative that comparisons be made with types of both M. quinquiesperforata and M. isometra. Before these analyses can be done, 752 the name M. quinquiesperforata must be stabilized by designating and describing in detail a neotype. Most aspects of the anatomy of M. quin- quiesperforata remain poorly or not at all figured. This is particularly true for external appendages such as spines and pedicellar- iae, which are notably variable within and between species of clypeasteroids (Mooi 1989). Although presented under the name Mellita quinquiesperforata, the majority of previous figures are actually of M. isometra, the species most commonly encountered in marine laboratories of the eastern seaboard (for example, see Mooi [1986] and Telford et al. [1985] for spination and podia of /. isometra). Louis Agassiz (1841) recognized two species that are together synonymous with M. quinquiesperforata. He was the only one to figure spines of any kind from ™. quinquiesperforata, in the sense of Harold & Telford (1990). Under the name of ™. testudinata, Agassiz showed a single spine from the aboral margin of one of the lunules (L. Agassiz 1841: plate 4a, fig. 7a), although he does not indicate which one. Under the name M. quinquefora, he showed a poorly drawn club-shaped spine (L. Agassiz 1841: plate 3, fig. 11a) from the aboral surface, and three views of spines (L. Agassiz 1841: plate 3, figs. 111, c, d) from the aboral mar- gins of unspecified lunules. He described these spines as resembling elongated spoons, an observation that is not consistent with our own. Clark (1914: plate 125, figs. 16- 21) figured several spines and pedicellariae from what he calls M. quinquiesperforata, but without knowing the locality of the spec- imen, or its appearance, it is impossible to know if this is indeed M. quinquiesperfor- ata, or (as we suspect) M. isometra. In ad- dition, Clark’s figures are not especially ac- curate, and the figure of the “miliary spine” (Clark 1914: plate 125, fig. 16) seems to be a geniculate spine. A single bidentate ped- icellaria from “‘Mellita 5-perforata lata H.L. Clark” was figured by Mortensen (1948), PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON but it lacks detail in the tooth structure of the valves. No figures of biphyllous pedi- cellariae exist in the literature. The plate pattern of the oral surface of M. quinquiesperforata from the “Gulf of Mexico” was figured by Durham (1955: fig. 17A). He also shows the basicoronal plate system (Durham 1955: fig. 16B). Unfortu- nately, there are errors in the positions and numbers of plate sutures associated with the anal lunule in Durham’s drawing of M. quinquiesperforata (similar errors appear in his figure of Leodia sexiesperforata [Leske, 1778]). No figures of the plate pattern from the aboral surface exist in the literature, al- though some details can be made out in L. Agassiz’s figure of /. quinquefora (L. Agas- siz 1841: plate 3, fig. 1) and M. testudinata (L. Agassiz 1841, plate 4a, fig. 10). We could locate only 2 figures that unambiguously show the internal structure of M. quinquies- perforata: those in Mortensen (1948: plate 62, fig. 3) and Harold & Telford (1990: fig. 7). Only in the latter is the Aristotle’s lan- tern visible. Alexander Agassiz (1872) noted that /. quinquiesperforata “‘has a wide geographical range, and is liable to great variations.” In the past, and in conjunction with taxonomic confusion stemming from these variations, it has been difficult to determine an un- equivocal suite of characters for M. quin- quiesperforata. For example, D. Pawson of the National Museum of Natural History, Washington, D.C. recently brought to our attention some difficulties in the use of Har- old & Telford’s (1990) key, particularly with respect to spine morphology, posterior paired lunule angle, and petaloid size. The present state of knowledge regarding the variation within the species suggests that the populations occurring in the northwestern Gulf of Mexico, including the type locality of Veracruz, Mexico, unequivocally belong to a single taxon. Harold & Telford (1990) pointed out that there was some distinc- tiveness of populations, particularly those VOLUME 107, NUMBER 4 from islands in the Caribbean and from the southern parts of the range, but they pre- ferred to recognize them only as geographic or ecophenotypic variants of M. quinquies- perforata. Some of these variants could turn out to be separate species, making it es- pecially important to associate a specimen with the name M. quinquiesperforata. It is clear that a full description, especially of the spines, pedicellariae, and features of MW. quinquiesperforata’s test architecture are necessary for adequate comparisons with types and other specimens of the rest of the species in Mellita. Methods Terminology of external appendages and test features is that of Mooi (1989), and the plate columns are labeled according to Lo- vén (1892). We provide an abbreviated syn- onymy, including all names that have been applied to M. quinquiesperforata. For ref- erences to more literature on this species, see Mortensen (1948) and Harold & Telford (1990). For a description, summary of in- traspecific variation, and diagnosis of the species as a whole, see Harold & Telford (1990). Unless indicated otherwise, mea- surements are made directly from the neo- type using dial calipers, with the value given in parentheses after each measurement rep- resenting percent test length. Additional comments are intended to cover anatomical aspects not emphasized by Harold & Tel- ford (1990). Lunule angle is the number of degrees subtended by axes drawn through the lunules in ambulacra I and V. Spines and pedicellariae were removed from the test of the neotype and placed in droplets of approximately 5% sodium hypochlorite (Clorox). Fully cleaned spines were drawn using a camera lucida mounted on a com- pound microscope. Ten each of miliary spines (from aboral interambulacrum 2), aboral club-shaped spines (from aboral in- terambulacrum 2), geniculate spines (from 753 oral ambulacrum I), locomotory spines (from oral interambulacrum 2), and anal lunule fringe spines (from the aboral edge at the mid-point of the lunule) were mea- sured to the nearest wm using a calibrated eye-piece micrometer. To avoid damaging the neotype, specimens of similar size to, and collected with the neotype were pre- pared or dissected as follows. Plate patterns were made visible by polishing the speci- men with a graded series of water-proof sand papers, and then painting the specimen with a light coat of a solution of equal parts glyc- erol and 95% ethanol. The plate patterns of both surfaces were drawn with the aid of a camera lucida mounted on a binocular dis- secting microscope. The Aristotle’s lantern was exposed through dissection of the ab- oral surface of another specimen and mea- sured. Order Clypeasteroida L. Agassiz, 1835 Family Mellitidae Stefanini, 1911 Genus Mellita L. Agassiz, 1841 Mellita quinquiesperforata (Leske, 1778) Figs. 1-3 Echinodiscus quinquies perforatus Leske, 1778:197, pl. 21C, D. Echinus pentaporus.—Gmelin, 1788:3189. Clypeaster pentaporus. —Lamarck, 1801: 349. Scutella quinquefora. —Lamarck, 1816:9. Scutella pentapora. —Blainville, 1830:223. Mellita quinquefora. —L. Agassiz, 1841:36, pl. 3 (plate is erroneously labelled “En- cope pentapora’’). Mellita testudinata.—L. Agassiz, 1841:40, pl. 4a, figs. 7-9. Mellita nummularia.—L. Agassiz & Desor, 1847:139. Mellita testudinea. —Gray, 1855:22. Mellita pentapora. —Liitken, 1864:107. Mellita quinquiesperforata.—H. L. Clark, 1911:599. Mellita quinquiesperforatus.—H. L. Clark, 1925:174. 754 Mellita lata. —H. L. Clark, 1940:437, pl. 60, fig. 1, pl. 61, fig. 1, pl. 62, figs. 1, 2. Mellita latiambulacra. —H. L. Clark, 1940: 439, pl. 62, figs. 3-6. Mellita quinquisperforata var. latiambula- cra.—Penchaszadeh & Layrisse, 1985: 393. Neotype.—California Academy of Sci- ences, Invertebrate Zoology (CASIZ) 096152, 44.2 mm test length, beach 2 mi south of Veracruz, Mexico, A. G. Smith, March 1954 (collected with CASIZ 087802, geology accession number 34684, 11 spec- imens in lot not counting neotype, ranging from 26.6 mm to 44.6 mm test length). Description of neotype. —Test dimensions (Fig. 1): Length 44.2 mm; maximum width 48.0 mm (108.6%); distance from ambitus at ambulacrum III to anterior edge of mad- reporic plate 19.4 mm (43.9%); test thick- ness at center of madreporic plate 5.6 mm (12.7%); test thickest anterior to madreporic plate, thickness at highest point 5.9 mm (13.3%); distance from ambitus at ambu- lacrum III to highest point of test 13.2 mm (29.9%); distance from ambitus at ambu- lacrum III to anterior edge of peristome 13.0 mm (29.4%); mouth diameter 2.0 mm (4.5%); distance from anterior edge of peri- stome to anterior edge of periproct 4.0 mm (9.0%); periproct indenting basicoronal, in depression leading to anterior edge of anal lunule, length 1.9 mm (4.3%). Petaloids (Fig. 1A): Only a single trailing podium in each of petaloids Ib, IIb, IIIa, IVa and Va (see also Fig. 3). Following mea- surements represent length of indicated pet- aloid pore pair column in ambulacra: Ib 11.2 mm (25.3%); IIb 10.00 mm (22.6%); IIa 11.3 mm (25.6%); [Va 11.0 (24.9%); Va 11.7 mm (26.5%). Following measurements rep- resent greatest width of petaloid in ambu- lacra: 15.4 mm (12.2%); II 5.2 mm (11.8%); III 5.0 mm (11.3%); IV 5.1 mm (11.5%); V 5.5 mm (12.4%). Food grooves: Primary grooves bifurcate PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON on, or just distal to ambulacral basicoronal plates; two main branches in each ambu- lacrum restricted to ambulacral plates; dis- tal, secondary branches extend onto inter- ambulacral plates (see also Fig. 3). Following measurements represent greatest distance (orthogonal to lunule axis) between main branches of food grooves in ambulacra: I 9.5 mm (21.5%); II 10.1 mm (22.9%); III 6.3 mm (14.3%); IV 10.0 mm (22.6%); V 9.3 mm (21.0%). Lunules (Fig. 1): Lunules closed. Follow- ing measurements represent lengths and widths respectively of lunules in ambulacra: I 8.4 mm (19.0%) and 1.7 mm (3.8%); II 8.6 mm (19.5%) and 1.7 mm (3.8%); IV 7.7 mm (17.4%) and 1.8 mm (4.1%); V 8.4 mm (19.0%) and V 1.7 mm (3.8%); lunule angle 73.5°; distance from anterior edge of mad- reporic plate to anterior edge of anal lunule 6.6 mm (14.9%); length and width of anal lunule 11.8 mm (26.7%), 2.5 mm (5.7%). Spines (Fig. 2): Measurements given here are mean spine lengths, followed by (in pa- rentheses) range and standard deviation. Aboral miliary spines (Fig. 2A) slender, bent at base so that whole spine leans ““down- slope”’ from apical system, distal end with typical sac-bearing tip architecture (see Mooi 1986), length 508.2 um (486-528, 11.8). Aboral club-shaped spines (Fig. 2B, C) fairly stout, with thick shaft bent at base so that whole spine leans ““down-slope”’ in manner similar to aboral miliaries, tip moderately expanded and club-shaped, length 642 um (605-658, 15.0). Geniculate spines (Fig. 2D) of oral surface slender and sharply bent just distal to point half way up shaft, length 572 um (546-598, 18.5). Locomotory spines (Fig. 2E) of oral surface long, slender, sharp- ly bent proximal to base, moderately point- ed, length 2397 wm (2073-2611, 153.0). Anal lunule fringe spines (Fig. 2F) from ab- oral edge of lunule paddle-shaped, but broadest and most greatly flattened at lu- nule’s mid-point, shaft proximal to base slightly bent in same plane as flattening, tip VOLUME 107, NUMBER 4 755 »; os AN Hani) Fig. 1. Mellita quinquiesperforata, neotype (CASIZ 096152, 44.2 mm test length). A. Photograph of aboral surface; B. Photograph of oral surface. 756 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. Mellita quinquiesperforata, neotype (CASIZ 096152), skeletal elements of external appendages. A. Miliary spine from interambulacrum on aboral surface, apex left; B. Club-shaped spine from interambulacrum on aboral surface; C. Club-shaped spine from interambulacrum on aboral surface, view orthogonal to that in B, apex left; D. Geniculate spine from ambulacrum on oral surface; E. Locomotory spine from interambulacrum on oral surface, details of stereom structure omitted; F. Anal lunule fringe spine from aboral edge at mid-point VOLUME 107, NUMBER 4 chisel-like and square, length 1610 um (1370-1772, 115.8). Pedicellariae (Fig. 2): Pedicellariae of two types, both with two valves forming a “jaw” attached to stem by long, flexible neck of almost same length as stem. Valves of bi- phyllous pedicellariae (Fig. 2G) with minute teeth all of similar size, valve length ap- proximately 45 wm. Stem slender, tapering distally but ending in slight swelling, stem length approximately 110 um. Valves of bi- dentate pedicellariae (Fig. 2H) with variably sized teeth, 2 or 3 long, distal ““canine”’ teeth on a valve interlocking with those of op- posing valve, valve length approximately 95 wm. Stem slender, same general shape as for biphyllous, stem length approximately 140 um. Plate pattern (Fig. 3), Aristotle’s lantern of non-types.—Aboral plating around am- bulacral lunules festooned except in am- bulacrum III, which lacks lunule. Aborally, narrowest point of interambulacrum 5 pos- terior to anal lunule. At ambitus, interam- bulacra and ambulacra approximately same width. Orally, interambulacra discontinu- ous by single ambulacral plate except for interambulacrum 5, which is continuous. During ontogeny, last post-basicoronal plates to become disjunct from their cor- responding basicoronals are those in inter- ambulacra 1b, 2a, 3b and 4a. Four inter- ambulacral plates form perimeter of anal lunule on oral surface, two circumferential sutures close to posterior end of lunule. In- terambulacral basicoronal plates longer than ambulacral basicoronals. Interambulacral basicoronal 5 deeply indented by periproct. In specimen 42.5 mm long, Aristotle’s lan- tern 8.8 mm long (20.7% test length). Remarks.—Klein (1734) described and figured specimens of this species from Ve- 757 & net Fig. 3. Mellita quinquiesperforata (CASIZ 087802, 44.6 mm test length, collected with neotype), plate and food groove patterns. Food grooves represented by dot- ted lines, details of plating in petaloids omitted. Mouth and periproct in solid black, interambulacral plates shaded. racruz and Leske (1778), the first post-Lin- naean writer to describe M. quinquiesper- forata (as Echinodiscus quinquies perfora- tus) referred specifically to Klein’s figures (Harold & Telford 1990). Accordingly, we have chosen a specimen from a lot from Veracruz as the neotype. The neotype does not differ significantly from the material ex- amined by Harold & Telford (1990). How- — of anal lunule, details of stereom structure omitted; G. Valves (upper) and stem (lower) of biphyllous pedicellaria; H. Valves (upper) and stem (lower) of bidentate pedicellaria. All scale bars 100 um long. 758 ever, as noted by Harold & Telford (1990), there is substantial variation within M. quinquiesperforata, particularly in width to length ratio, profile, and spine dimensions. In spite of previous attempts to do so (for example, see Clark 1940), Harold & Telford (1990) could find no basis upon which to partition this variation into species. We concur based on the characters examined by previous authors as well as on new ob- servations (D. Pawson, A. S. Harold, & R. Mooi, unpublished observations). The neo- type here designated not only comes from the type locality, it also exhibits the sharply inclined anterior profile, anteriorly placed site of maximum thickness, very broad test, and spatulate spines bordering the lunules listed in Harold & Telford’s (1990) diag- nosis. Although pedicellarial types can be diagnostic of certain clypeasteroid clades (Mortensen 1948, Mooi 1989), the pedicel- lariae of the neotype illustrated here (Fig. 2) are typical not only of Mellita quinquies- perforata, but of the genus as a whole, and do not offer additional taxonomic infor- mation. The neotype typifies the apomorphies shared by M. quinquiesperforata and other Mellita. Phylogenetic revision of fossil Mel- lita (R. Mooi & A. S. Harold, unpublished observations) indicates that Leodia sexies- perforata is a member of the clade encom- passing both fossil and extant Mellita spe- cies. Taking this into account, the apomorphies of Mellita include a single trailing podium at the end of each petaloid, and the periproct indenting the basicoronal plate. Harold & Telford (1990) added the loss of the lunule in ambulacrum III to this list. However, this character is actually ap- plicable only to a smaller clade of largely extant Mellita, and not to the genus as a whole (R. Mooi & A. S. Harold, unpub- lished observations). Our examination of the neotype and associated material also affords an opportunity to rectify Durham’s (1955) error in interpretation of the plate patterns around the anal lunule (compare our Fig. 3 with Durham 1955: fig. 17A). PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON A survey of spine morphometrics within the Mellitidae (D. Pawson, A. S. Harold & R. Mooi, unpublished observations) reveals unexpected divergences from the characters given in the key by Harold & Telford (1990). They use the relative lengths of anal lunule fringe spines and locomotory spines in their first couplet: in M. tenuis and M. isometra, the anal lunule fringe spines are supposed to be “substantially longer’’ than the loco- motories. In all other Mellita, the reverse is the case. The anal lunule fringe spines are shorter than the locomotories in the neotype and other examples of M/. quinquiesperfor- ata and is therefore in accord with Harold & Telford’s key. However, with the recog- nition that some populations of M. isome- tra, and perhaps of M. tenuis, have longer locomotories than anal lunule fringe spines, more comprehensive study of intraspecific variation (including all available types) is essential to assess the systematic, environ- mental, and ontogenetic significance of pre- viously unrecognized variation in relative spine lengths. Acknowledgments We would like to thank Bob Van Syoc and Liz Kools for their help in the collec- tions and with obtaining catalogue data. We are also grateful to Charlotte Fiorito for her skill in producing the photographs. The re- search was supported in part by a California Academy of Sciences Tilton Fellowship to the second author. Literature Cited Agassiz, A. 1872-1874. Revision of the Echini.— Memoirs of the Museum of Comparative Zo- ology at Harvard College 3:383-762. Agassiz, L. 1841. Echinites. Famille des clypéas- troides. Des Scutelles.—Monographies des Echinodermes Vivans et Fossiles 2:1—151. —., & P. J. E. Desor. 1847. Catalogue raisonné des familles, des genres et des éspéces de la classe échinodermes.—Annales des Sciences Natu- relles, series 3, 6—8:1-167. Blainville, H. M. D. 1830. Zoophytes.—Dictionaire des Sciences Naturelles 60:169-—245. VOLUME 107, NUMBER 4 Clark, H. L. 1911. The genera of Recent clypeaster- oids.— Annals of the Magazine of Natural His- tory, series 8, 7:593-605. . 1914. Hawaiian and other Pacific Echini. The Clypeastridae, Arachnoididae, Laganidae, Fi- bulariidae, and Scutellidae.—Memoirs of the Museum of Comparative Zoology at Harvard College 46(1):1-78. . 1925. A Catalogue of the Recent Sea-Urchins (Echinoidea) in the Collection of the British Mu- seum (Natural History). British Museum (Nat- ural History), London, 250 pp. . 1940. Revision of the keyhole urchins (Mel- lita). — Proceedings of the United States Nation- al Museum 89:435-444. Durham, J. W. 1955. Classification of clypeasteroid echinoids.— University of California Publica- tions in Geological Sciences 31(4):73-198. Gmelin, J. F. 1788. Linnaei Systema Naturae, eighth edition. Lipsiae, Leipzig, 3200 pp. Gray, J. E. 1855. Catalogue of Recent Echinida in the British Museum. British Museum (Natural History), London, 69 pp. Harold, A. S., & M. Telford. 1990. Systematics, phy- logeny and biogeography of the genus Mellita (Echinoidea: Clypeasteroida).— Journal of Nat- ural History 24:987-1026. Klein, T. 1734. Naturalis dispositio echinoderma- tum. T. J. Schreiber, Gedani, unpaged. Lamarck, J.B. 1801. Systéme des Animaux sans Ver- tébres. Paris, 432 pp. 1816. Histoire Naturelle des Animaux sans Vertébres, III. Paris, 130 pp. Leske, N.G. 1778. Additamenta ad Jacobi Theodori Klein naturalem dispositionem Echinoderma- 759 tum et lucubratiunculam de aculeis echinorum marinorum. Lipsiae, Leipzig, 278 pp. Loven, S. 1892. Echinologica.—Bihang Svenska Ve- tenskap Akademie Handlingar 18:1-73. Litken, C. F. 1864. Bidrag til Kundskab om Echini- derne. Copenhagen, 139 pp. Mooi, R. 1986. Structure and function of clypeas- teroid miliary spines (Echinodermata, Echi- noides).— Zoomorphology 106:212-223. 1989. Living and fossil genera of the Cly- peasteroida (Echinoidea: Echinodermata): an il- lustrated key and annotated checklist.—Smith- sonian Contributions to Zoology 488:1-51. Mortensen, T. 1948. A monograph of the Echinoidea, 4(2). Clypeasteroida. Copenhagen, Denmark, C.A. Reitzel, 471 pp. Penchaszdeh, P. E., & M. E. Layrisse. 1985. Ecology of the sand dollar, Mellita quinquiesperforata latiambulacra on the west-central coast of Ven- ezuela. Proceedings of the International Echi- noderm Conference, Rotterdam, 1984, p. 393. Telford, M.,R. Mooi, & O. Ellers. 1985. Anew model of podial deposit feeding in the sand dollar, Mel- lita quinquiesperforata (Leske): the sieve hy- pothesis challenged.— Biological Bulletin 169: 431-448. (RM) Department of Invertebrate Zool- ogy and Geology, California Academy of Sciences, Golden Gate Park, San Francisco, California 94118-4599, U.S.A.; (ASH) De- partment of Ichthyology and Herpetology, Royal Ontario Museum, Toronto, Ontario MS5S 2C6, Canada. PROC. BIOL. SOC. WASH. 107(4), 1994, pp. 760-777 DESCRIPTION OF MASTIGLANIS ASOPOS, A NEW PIMELODID CATFISH FROM NORTHERN BRAZIL, WITH COMMENTS ON PHYLOGENETIC RELATIONSHIPS INSIDE THE SUBFAMILY RHAMDIINAE (SILURIFORMES: PIMELODIDAE) Flavio A. Bockmann Abstract. —Mastiglanis asopos, a new genus and species of the neotropical catfish family Pimelodidae, is described from the Amazon basin, and rio Capim and its adjacencies, in northern Brazil. Its monophyly is supported by various autapomorphies in external and internal morphology, such as the first dorsal- and pectoral-fin elements filamentous distally, and the presence of a remarkable osseous process at the anterior portion of the premaxilla. Mastiglanis shares unambiguously all diagnostic features proposed for the subfamily Rhamdiinae, and also those that delimit a subset called the Nemuroglanis sub-clade. Eleven new synapomorphies are proposed to corroborate the monophyly of the latter group. Additional character evidence supports a hypothesis placing Mastigla- nis as the sister group of the remainder of the Nemuroglanis sub-clade. Resumo.— Mastiglanis asopos, um novo género e espécie da familia Pime- lodidae é descrito para a regiao norte brasileira (bacia Amaz6nica e rio Capim e arredores). Seu monofiletismo é indicado por varios caracteres apomorficos observaveis na morfologia externa e interna, dentre. os quais destacam-se: os primeiros elementos (“espinhos’’) das nadadeiras dorsal e peitoral extrema- mente alongados e a presenca de um notavel processo Osseo na regiao anterior da premaxila. Mastiglanis compartilha todos os caracteres diagnosticos da subfamilia Rhamdiinae, assim como, todos aqueles que definem um subgrupo mais restrito chamado “‘Nemuroglanis sub-clade.”’ O monofiletismo deste ul- timo é€ corroborado por onze novas sinapomorfias. Mastiglanis é provisoria- mente posicionado como grupo-irmao de todos os outros componentes do ““Nemuroglanis sub-clade.”’ The most recent account on the situation of the systematics of South American fresh- water fishes is that of Bohlke et al. (1978), who estimated that a large portion of that fish fauna, especially that from the Amazon basin, remains undescribed and even un- known. In spite of the considerable progress made since the publication of Bohlke et al.’s paper, only recently have catfishes from the Amazon basin been dealt within a phylo- genetic context (e.g., Stewart 1986a, 1986b; Ferraris & Mago-Leccia 1989; Pinna 1989). The fish herein studied is an illustrative example of the above-mentioned situation, for it remained unknown until now in spite of its distinctive external features and wide distribution throughout the Amazonian ba- sin. This paper describes this pimelodid cat- fish as a new genus and species, and dis- cusses its relationships within a recently provided synapomorphy scheme of the Rhamdiinae (Lundberg et al. 1991a). Ad- ditional new corroboration is given to a for- merly suggested rhamdiine subset (Ferraris 1988) as well as a discussion about the ar- rangement of the genera therein included. VOLUME 107, NUMBER 4 Material and Methods Morphometric values were taken with calipers and expressed to the nearest 0.1 mm. All measurements are made point-to- point on the specimens’ left side whenever possible. Morphometric data are expressed as proportions of standard length (SL), or head length (HL; all subunits of the cephalic region, excepting the barbels which were ex- pressed in proportions of SL). Each pro- portion is provided with its arithmetic mean, followed by the range and standard devia- tion. The number of specimens tallied in counts is presented enclosed in parentheses. When meristics vary, the count for the ho- lotype is indicated by an asterisk (*). Meth- odology and terminology for measurements followed Lundberg & McDade (1986), with the following measurements added: prepel- vic length, preadipose length, dorsal-fin base length (taken as the gap between the inser- tion point of the first and the last elements), maxillary-barbel length, outer- and inner- mental barbel length, head depth (taken at the vertical through the posterior margin of the fleshy opercular flap), head width (taken at its maximum width), fleshy interorbital distance (the space between the fleshy upper orbital rims), preorbital length (measured from snout tip to anterior orbital rim), an- terior internarial space, and length of first and second branched rays of the dorsal and pectoral fins. Some paratypes of Mastiglanis asopos and additional comparative material of Pime- lodidae were cleared and counterstained employing the Taylor & Van Dyke (1985) technique, which provides a differential staining of the skeletal system elements. Vertebral counts included the first five ver- tebrae transformed in the Weberian appa- ratus, and the compound caudal centrum (PU1+U1) was counted as one. Principal caudal-fin ray counts included all branched rays plus one unbranched ray in each lobe, following Hubbs & Lagler (1958). Counts for each lobe, upper first, are separated by 761 a plus sign. Pterygiophores, vertebrae, gill rakers, ribs, and branchiostegal rays were counted on cleared and stained specimens only. Anatomical illustrations were sketched using a Zeiss stereomicroscope with a cam- era lucida attachment. In the drawings, bone is represented by stipple and cartilage by open circles. Institutional abbreviations are: Califor- nia Academy of Sciences, San Francisco (CAS); Museu de Zoologia da Universidade de Sao Paulo, Sao Paulo (MZUSP); Museu Nacional do Rio de Janeiro, Rio de Janeiro (MNRJ); Universidade Federal do Rio de Janeiro, Rio de Janeiro (UFRJ); Departa- mento de Biologia Animal e Vegetal da Universidade do Estado de Rio de Janeiro, Rio de Janeiro (DBAV-UERJ), National Museum of Natural History, Smithsonian Institution, Washington, D.C. (USNM). Comparative material examined (CS- cleared and stained preparations).—Acen- tronichthys leptos, UFRJ 289 (1 ex., CS), UFRJ 505 (1 ex., CS); Brachyglanis sp., MZUSP 45895 (1 ex., CS); Brachyplatysto- ma filamentosa, MZUSP 45896 (1 ex., CS); Brachyrhamdia meesi, UFRJ 397 (2 ex., CS); Calophysus macropterus, MZUSP 26415 (1 ex., CS), MZUSP 45904 (1 ex., CS); Cetop- sorhamdia theringi, UFRJ 689 (2 ex., CS); Cetopsorhamdia picklei, USNM 121218 (2 ex., paratypes, CS); Chasmocranus longior, MZUSP 45909 (2 ex., CS); Chasmocranus truncatorostris, UFRJ 322 (1 ex., CS), UFRJ 504 (1 ex., CS); Cheirocerus goeldi, MZUSP 45905 (2 ex., CS); Gladioglanis conquis- tador, MZUSP 45906 (2 ex., CS); Goeldiellal eques, MZUSP 45907 (1 ex., CS); Heptapterus mustelinus, UFRJ 291 (1 ex., CS); Hypophthalmus edentatus, MZUSP 43304 (1 ex., CS); I[heringichthys labrosus, UFRJ 690 (1 ex., CS), DBAV-UERJ 146 (1 ex.); Imparales mariai, USNM 121251 (1 ex., holotype); Imparales panamensis, USNM 293454 (1 ex., CS); Imparfinis min- utus, MZUSP 39990 (1 ex., CS), UFRJ 320 (4 ex.); I. mirini, MZUSP 45899 (1 ex., CS); I. nemacheir, USNM 121163 (2 ex., CS); I. 762 piperatus, CAS 63636 (1 ex., holotype); I. schubarti, MZUSP 45897 (1 ex., CS); Im- parfinis sp. 1, MZUSP 45900 (1 ex., CS); Imparfinis sp. 2, MZUSP 45898 (1 ex., CS); Imparfinis sp. 3, UFRJ 393 (1 ex., CS); Lo- Dhiosilurus alexandri, UFRJ 042 (1 ex., CS); Megalonema platanus, MZUSP 45902 (1 ex., CS); Megalonemasp., DBAV-UERJ 122 (1 ex., CS); Microglanis parahybae, UFRJ 693 (2 ex., CS); Nannoglanis bifasciatus, MZUSP 45903 (2 ex., CS); Nannorhamdia stictonotus, UFRJ 292 (3 ex., CS); Nemu- roglanis sp.,. MZUSP 45908 (1 ex., CS); Par- apimelodus valenciennis, DBAV-UERJ 230 (1 ex., CS); Phenacorhamdia _boliviana, MZUSP 42296 (2 ex., CS); Phenacorham- dia sp., MZUSP 45901 (1 ex., CS); Pime- lodella lateristriga, UFRJ 503 (2 ex., CS); Pimelodella sp., UFRJ 502 (2 ex., CS); Pi- melodus maculatus, UFRJ 691 (2 ex., CS); P. ornatus, DBAV-UERJ 934 (1 ex., CS); Pseudopimelodus sp., DBAV-UERJ 118 (1 ex., CS); Rhamdia sp., UFRJ 321 (3 ex., CS); Rhamdiopsis sp., UFRJ 708 (1 ex., CS). Mastiglanis, new genus Diagnosis.—A small pimelodid catfish displaying the suite of apomorphic char- acters ascribed to the subfamily Rhamdi- inae (Lundberg, 1991a). It also exhibits all attributes that characterize a more restrict- ed monophyletic unit called ““Nemuroglanis sub-clade”’ (Ferraris 1988; see also Phylo- genetic Relationships section below). Mas- tiglanis is distinguishable from all other rhamdiines by the following putatively au- tapomorphic traits: 1 —integument pigmen- tation very reduced; 2—anteriormost ele- ment of the dorsal fin (homologous to dorsal- fin spine) prolonged as a long filament; 3— first pectoral-fin element (homologous to pectoral-fin spine) prolonged as a long fil- ament; 4—anterior internarial width greater than the posterior one; 5—frontals narrow at supraorbital portion; 6—an anterodorsal oriented shelf-like process at the symphysial region of premaxilla; 7— anterior cornua of PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON mesethmoid abruptly assuming a reversed direction at their mid-length; 8—an elon- gated metapterygoid; 9—a lamina at the an- terodorsal margin of the opercle; 10—rear portion of opercular bone tapered and curved ventrally. Other characters consid- ered plesiomorphic or of unknown polarity but still useful for identification include: eyes large; long maxillary barbels extending be- yond adipose-fin origin; mouth ventral: bones of cranial roof thin, lacking orna- mentation and connected via lap junctions (dentate sutures absent); well-developed su- praoccipital process; and triangular pectoral fins. Type species.—Mastiglanis asopos, new species. Etymology. —From the Greek mastix (mastigo, in latinized form) meaning whip, in allusion to the filamentous elements of pectoral and dorsal fins; and glanis, the name of the Greek catfish of Aristotle, a common denomination for fishes of the order Silur- iformes. The last two letters of the word mastigo were suppressed for the sake of eu- phony. Gender masculine. Mastiglanis asopos, new species Figs. 1, 2, Table 1 Diagnosis. —As for the genus. Holotype. —MNRJ 12227, SL 65.9 mm, Brazil, Para, Igarapé Saracazinho, tributary of rio Trombetas, near Porto Trombetas, coll. E. P. Caramaschi & D. F. Moraes Jr., 14 Dec 1990. Paratypes.—MNRJ 12228, 10 ex., SL 42.5-53.1 mm, taken with holotype; MNRJ 12229, 1 ex. cleared and stained, SL 48.8 mm, taken with holotype; MZUSP 7446, 2 ex., SL 44.8-51.5 mm, Brazil, Amazonas, municipio de Silves, rio Sanabani, coll. EPA, 7-8 Dec 1967; MZUSP 23299, 2 ex., SL 37.0-—39.1 mm, Brazil, Amazonas, rio Jauaperi, beach 30 km upstream from the river mouth, coll. T. Roberts, 19 Nov 1968; MZUSP 23533, 1 ex., SL 39.6 mm, Brazil, Amazonas, left margin of the rio I¢a, Cuiaua, VOLUME 107, NUMBER 4 Igarapé da Cachoeira, coll. EPA, 18 Oct 1968; MZUSP 23541, 10 ex., SL 30.8-58.2 mm, Brazil, Amazonas, left margin of the rio Iga, upstream from Cuiaua, Igarapé Boa Vista, coll. EPA, 19 Oct 1968; MZUSP 23549, 1 ex., SL 43.0 mm, Brazil, Ama- zonas, left margin of the rio Solimoes, 7 km downstream from the Santo Antonio do I¢a, Igarapé Ica, coll. EPA, 20 Oct 1968; MZUSP 23875, 1 ex., SL 35.8 mm, Brazil, Para, rio Capim, Caranandéua beach, coll. EPA, 17 Aug 1970; MZUSP 24282, 2 ex., SL 28.3- 34.7 mm, Brazil, Para, municipio de Sao Luis, rio Tapajos, coll. EPA, 5 Nov 1970; MZUSP 25635, 1 ex., SL 27.4 mm, Brazil, Para, pov. Pimental, right margin of the rio Tapajos, mouth of the Igarapé Pimental, coll. J. C. de Oliveira, 15-31 Jul 1979; MZUSP 30617, 84 ex., SL 16.5—40.0 mm, Brazil, Amazonas, rio Negro, 2 hours up- stream from Barcelos, Urumari beach, coll. M. Goulding, 6 Oct 1979, 0000 h; MZUSP 30618, 11 ex., SL 19.6—41.8 mm, Brazil, Amazonas, beach at the Tamaquaré island, coll. M. Goulding, 10 Oct 1979, 2100 h; MZUSP 30633, 2 ex., SL 32.9-33.8 mm, Brazil, Amazonas, Massarabi, rio Negro, coll. M. Goulding, 18 Oct 1979; 2000 h; MZUSP 30635, 1 ex., SL 35.5 mm, Brazil, Amazonas, beach at Parana do Jacaré, coll. M. Goulding, 7 Oct 1979, 2100 h; MZUSP 30636, 2 ex., SL 30.7-34.7 mm, Brazil, Amazonas, beach near mouth of rio Ari- rara (tributary of rio Negro), coll. M. Gould- ing, 6 Oct 1979, 2100 h; MZUSP 34953, 6 ex., SL 39.0—49.4 mm, Brazil, Amazonas, rio Daraa (tributary of rio Negro), Cachoei- ra do Aracu, coll. M. Goulding, 10 Feb 1980; MZUSP 34954, 17 ex., SL 26.4-38.1 mm, Brazil, Amazonas, near mouth of rio Mar- auia (tributary of rio Negro), coll. M. Goulding, 13 Oct 1979; MZUSP 34955, 6 ex., SL 29.6—33.2 mm, Brazil, Amazonas, confluence of rio Arirara with rio Negro, coll. M. Goulding, Oct 1979; MZUSP 44215, 1 ex. cleared and stained, SL 52.0 mm, taken with MZUSP 34953: UFRJ 381, 2 ex., SL 35.7—38.6 mm, Brazil, Para, muni- 763 Coes <= oe as rae? =a hy ee. — ss Mihi Age IER ONG rave bi ai 1702 oiAvqoadel Drea [Poel] galeonkia / th x theanisioi 48 cBigerzel Jiatey teak mi j s groreciy ihe: ad a\ cranny mAs a 2) fotewtisil AyeVeai chet Aye © hes ae 4 wer Pa Be vain -s eae NA Wa fiw veld ins yh ee, (ules rug ee nA si Ome. cdteyt alte reg ashy 2) Nighusa staat mete te *,: sine _ afthy: icy ery, weil a rreiig sft nd paia tlt SGM ayy et nutans Ploy chee eee i RSH ak ryote aie i eet es Daath nn) gs r pineapple hy : suber teen, Sr Chi cata ant tr bof Rr iy ere Mace auth ine’! Memetquasag it olor] ais Niet al wars COS 14, tab beiroah reat v7 ao (Wl } At ec tiaa a (Tee re sen rt Loh darenie ett Ne Meats osu naif’ ) a Lo te, » i slid sr ttiglaal oilva Deetarras fersrquel ky Beenie 7 ORD mainte Mio 0 jhosbantsnt Os weil bape rae re = s iyatiaercd web ios erties p ny atyyole eicer ie d) eas hee cL maa ihe x nomen yibsth oe Ahold iw aeitone ts | i a ; win Hy ws, Geom eon. . ' 0 Bak conilitalnet t) , () imate bad vee oscar ates deals ag | byaiqe . JieoS) PRL seladO aq eab yore mae silane) Wel oaaye nn ot hewiicacet SOU) ainrwtl < wienitebl) ° 2) .2ibliet vilitveyatt _— i?) Bin uuvIN 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 man- uscripts by Society members will be accepted. Papers are published in English (except for Latin diagnoses/descriptions of plant taxa), with an Abstract in an alternate language when appropri- ate. Submission of manuscripts.—Submit three copies of each manuscript in the style of the Proceedings to the Editor, complete with tables, figure captions, and figures (plus originals of the illustrations). Mail directly to: Editor, Proceedings of the Biological Society of Washing- ton, National Museum of Natural History NHB-108, Smithsonian Institution, Washington, D.C. 20560. (Do not submit manuscripts to an associate editor). 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 and diagnoses. The style for the Proceedings is described in “GUIDELINES FOR MANUSCRIPTS for Publications of the BIOLOGICAL SOCIETY OF WASHINGTON” a supplement to Volume 103, number 1, March 1990. Authors are encouraged to consult this article before manuscript preparation. Copies of the article are available from the editor or any associate editor. The establishment of new taxa must conform with the requirements of appropriate interna- tional codes of nomenclature. Decisions of the editor about style also are guided by the General Recommendations (Appendix E) of the International Code of Zoological Nomenclature. When appropriate, accounts of new taxa must cite a type specimen deposited in an institutional col- lection. Review.—One of the Society’s aims is to give its members an opportunity for prompt publi- cation of their shorter contributions. Manuscripts are reviewed by a board of Associate Editors and appropriate referees. Proofs.—Authors will receive first proofs and original manuscript for correction and ap- proval. Both must be returned within 48 hours to the Editor. Reprint orders are taken with re- turned proofs. Publication charges.—Authors are required to pay full costs of figures, tables, changes in proofs ($3.00 per change or revision), and reprints. Authors are also asked to assume costs of page-charges. The Society, on request, will subsidize a limited number of contributions per vol- ume. If subsidized manuscripts result in more than 12 printed pages, the additional pages must be financed by the author(s). Multiple authorship will not alter the 12 page limit (each author will be viewed as having used his/her 12 subsidized pages). Payment of full costs will facilitate speedy publication. Costs.—Printed pages @ $60.00, figures @ $10.00, tabular material @ $3.00 per printed inch per column. One ms. page = approximately 0.4 printed page. CONTENTS Chesapeake Bay phytoplankton: I. Composition Harold G. Marshall Letepsammia franki, a new species of deep-sea coral (Coelenterata: Scleractinia: Micrabaci- idae) Joan Murrell Owens Siderastrea glynni, a new species of scleractinian coral (Cnidaria: Anthozoa) from the eastern Pacific Ann F. Budd and Hector M. Guzman Resurrection of Glyphohesione Friedrich, 1950, with redescription of G. klatti Friedrich, 1950 and description of G. longocirrata (Polychaeta: Hesionidae) Frank Licher Additional records of polynoid polychaetes from the Juan de Fuca Ridge Marian H. Pettibone A new genus and species of polychaete, Bollandia antipathicola (Nereidoidea: Syllidae), from black coral Christopher J. Glasby New species of Diplocardia and Argilophilus (Annelida: Oligochaeta: Megascolecidae) from southern California Samuel W. James Peosidriloides, a new genus, and new records of Peosidrilus (Oligochaeta: Tubificidae) from the United States, with the description of a new species from the Gulf of Mexico Christer Erséus and Michael R. Milligan A new American fairy shrimp, Linderiella santarosae (Crustacea: Anostraca: Linderiellidae) from vernal pools of California, U.S.A. Alain Thiéry and Michael Fugate Two new marine interstitial Ostracoda (Crustacea: Pussellidae) from Fiji Shinichi Hiruta Arctodiaptomus novosibiricus Kiefer, 1971 in Alaska and Northwest Territories, with notes on A. arapahoensis (Dodds, 1915) and a key to New World species of Arctodiaptomus (Copep- oda: Calanoida) Edward B. Reed Observations on the genus Mysidopsis Sars, 1864 with the designation of a new genus, Ameri- camysis, and the descriptions of Americamysis alleni and A. stucki (Reracarida: Mysidacea: Mysidae), from the Gulf of Mexico W. Wayne Price, Richard W. Heard, and Lidia Stuck Armadillidium tabacarui (Isopoda: Oniscidea: Armadillidiidae), a new troglobitic species from a sulfurous cave in Romania Magdalena Gruta, Vasilica Iavorschi, and Serban M. Sarbu Two new species of Podocerus Leach (Crustacea: Amphipoda: Podoceridae) from Bermuda Adam J. Baldinger and Michael F. Gable Systematic implications of color pattern polymorphism in Goniopsis pulchra (Decapoda: Brachyura: Grapsidae) from Ecuador Richard von Sternberg Two new species and one new combination of freshwater crabs from Mexico (Crustacea: Brachyura: Pseudothelphusidae) Fernando Alvarez and José Luis Villalobos A new species of anemone-carrying crab from New Caledonia (Decapoda: Brachyura: Xanthi- dae: Polydectinae) Cheryl G. S. Tan and Peter K. L. Ng Three new rare Heterokrohnia species (Chaetognatha) from deep benthic samples in the north- east Atlantic Jean-Paul Casanova Anatomical observations of the sand dollar Mellita quinquiesperforata (Leske, 1778) (Echino- dermata: Echinoidea) and the designation of a neotype Rich Mooi and Antony S. Harold Description of Mastiglanis asopos, a new pimelodid catfish from northern Brazil, with com- ments on phylogenetic relationships inside the subfamily Rhamdiinae (Siluriformes: Pimelo- didae) Flavio A. Bockmann International Code of Zoological Nomenclature Table of Contents, Volume 107 Index to New Taxa, Volume 107 520 PAS, 75g 10/05/95 198115 ses 573 586 591 600 609 615 622 634 641 657 666 680 699 707 721 729 738 743 We! < = ns => =< Ne = z =i) upg ie = Zz W oO Pc s Ly (eo) a = (e) \ AS < Sle B i J Mf 2 B ZR 8 = 2 Vf r= 2 cE Se = > @ = 5 = is 7) a. PA Ww a w ae VWINOSHLINS S3IYVYSIT_ LIBRARI ES SMITHSONIAN INSTITUTION NOILNLILSNI NVINOS| #4 a z 2 = S ca n 2 an = nw Ss NY BW : é = : < 2 Qs y oo = a —_ foo) —, =a (e} oe [e) — (S) Se) at} Zz ej Dae a Zz SMITHSONIAN INSTITUTION NOILALILSNI NVINOSHLINS S3!IYVYSIT LIBRARIES SMITHS SN z far z e ae wo — wo ‘ — [ve] = = = Pe) Fo) Wo, E = F > Ng .e > : - b = Wb = be o = idyll ne o z NVINOSHLINS (S34 luVvud eet B RARI ES SMITHSONIAN INSTITUTION NOILALILSNI ee = f,5 = ate! = pe —_— Zz ~ = Y, fy, 2 YGy.% Nx: 2 \y i B32 g Lo 2 AN 2 » —/ 2 a | Z yey 2 = SMITHSONIAN INSTITUTION NOILALILSNI_ NVINOSHLINS S3IYVYSIT LIBRARIES SMITHS zZ Ww 2 Ww Zz eg - 4 Ne 4 WY: a os ” it we - rs w PS Sore w 2 i = ne. = z = Zz 5 z NS = Zz Ew re) = DO MSS = re) g : g 3 ZW, 8 Z S = 20 = ae Ss > « = - > = ‘\ > =} Zz 77) Zz 77) SS - Ns LIBRARIES SMITHSONIAN INSTITUTION NOILNLILSNI_NVINOSHLINS saluvual ma Ww z= tw a — aR Ww w A aW rx wn Qed 4 ee ay eee = a AS ow a “PUY a < = \ = a “Gy 5 = a Y ac S 0 Ys 3 S = Sa z 24 Zz = z Ss ped TUTION NOILMLILSNI NVINOSHLINS. S3INVUGIT LIBRARIES SMITHSONIAN INSTITUTIOI z it : E Zbl eyes e = wo = o = w = 2 Leg, = 0 = _ = F \ WWREE zs = = fe = WW" 2 = B° = z Cie NS D z y VUGIT LIBRARIES SMITHSONIAN INSTITUTION NOILQLILSNI NVINOSHLINS S31uvual ; ra 5 n za * w ze w I, ¥ < < =< < = Ps Vy pao \: 6 = 8 : ye Ty a 7) GD AD a) fy 7 — \ (e) xr YS) me (e) ; EE \ 2 = ZH = Z SES ‘ 5 Oe ie a 5 S 2 TUTION NOILNLILSNI NVINOSHLIWS INSTITUTIO! w z * Ww > (ép) S = 7) oO n a wn a = a = w Peat < a Ne SY y Ww — eae 5 a = W'S = Be - IVUSIT atl B RARI ES SMITHSONIAN INSTITUTION NOILNLILSNI NVINOSHLINS §S3 1yVvuydl Zz = = bY =i jo: a z Zz = £ aS = ay Ys : ” 2) WY Yk Dn n Goes 1 SS \ e) ae * O yp the Bc: Oo hf: Ee \ z2 -E Ss Ng f 7 = = : “ = > >" > 7) x 2 = = S Fs TUTION NOILQLILSNI 1IRRARIES SMITHSONIAN INSTITUTIO SMITHSONIAN INSTITUTION LIBRARIES || niin 3 9088 01205 2320 . ‘ pee iy ‘ , ' : or i ; io / Vs “ ’ i f 1 BOF ET ; APU on 4 ‘ . . oe Mt! 1 » . a iene i : ] é aH : 24 ‘ Ren WEG ebace aarp eeu nie : aay i wast ML DHT EM RTA on iv : ; ties i? ’ Liebe Aeaheseb rar tsce eps lial ‘ : ‘ Sa) Mbaryer te usy nt} , aes ' Anche nar AEE racettis E > % ¥ iu Wl H : beowtte “1 BE Uo OEE Ey POMOU, APH ae } , ; ety ' Heap DME ests ature nebe : ' } CNS ae eipbs ae tel crd { ( HNL. Veal to : Mie ; : ah hie UP uPai align su okt WT Ne it RAL \ D yak 5 rbnonioe pe ysbuinie-seyyay ly cy pat AY F i { , , Novnearupe Mis) f P / i . Me ay S, Bae teucat aM ; My Ra \ tres TON Dt page n ‘ rn . ; Pe ee { at) i Set bap eh f : \ ‘ OL open ge ag: n ‘ \ ‘ CORT et \ Ha Yeah te keys H ; no} nba h \ \ t i" > yi i 4, te » VEE ! } \ . he ay vines \ ‘ : f wee . . (ies LATA Awa f 4} ‘ \ . . \ or Whe ut ie nthe te { Pit Gane iN f Phat hia ‘1