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INSTITUTION fait LIBRARIES SMITHSONIAN INSTITUTION AN AN NOILNLILSNI NVINOSHLIWS NVINOSHLIWS NVINOSHLINS S31 LIBRARIES SMITHSON! LIBRARIES SMITHSON! NOILNLILSNI 1917 LIBRARIES N iT INSTITUTION NOILOLILSNI Sa1uVvuait INSTITUTION Sagruvua TION NOIMLNLILSNI NVINOSHLINS S3iu¥Vddl) SMITHSONIAN NVINOSHLIWS NVINOSHLINS SMITHSONIAN _ INSTITUTION NVINO NOILNLILSNI LIBRARIES NOILNLILSNI TION NOILNLILSNI SHLINS S3Jld¥vugil 4 INSTITUTION S3tuvusl INSTITUTION NOILNLILSNI S3iuyvudit LIBR SMITHSONIAN INSTITUTION NOILNLIL NOILNLILSNI 1 LIBR SMITHSONIAN S3ZIYVUSIT LIBRARIES Ss, y 8 ‘ oe he Xs CREE S, RS S3lyuvudl Y \_) eld A MS CrraIOy LIBF NVINOSHLIWS INSTITUTION WOIL ARIES ® SN NVINOSHLINS S3I1YVUslT SMITHSONIAN | NVINOSHLIWS LIBRARIES SMITHSONIAN” INSTITUTIO INSTITUTION NOILMLILSNI JINVUSIT LIBRARIES S NVINOSHLINS S3JiuVvad! NVINOSHLINS INSTITUTIO S SMITHSONIAN ARIES SMITHSONIAN LIBRARIE NOILNLILSNI NVINOSHLINS S318Vdad! + INSTITUTION NOILNLILSNI LIBRARIES SMITHSONIAN INSTITUTIC NVINOSHLINS S31uvusl SMITHSONIAN NOILOLILSNI NVINOSHLIWS S3iuvua . LIBRARIES SMITHSONIAN_INSTITUTIC INSTITUTION NOILLNLILSNI S31UVUSIT LIBRARIES PROCEEDINGS of the Biological Society of Washington VOLUME 100 1987 Vol. 100(1) published 25 February 1987 Vol. 100(3) published 14 October 1987 Vol. 100(2) published 19 May 1987 Vol. 100(4) published 31 December 1987 WASHINGTON PRINTED FOR THE SOCIETY EDITOR BRIAN KENSLEY ASSOCIATE EDITORS Classical Languages Invertebrates GEORGE C. STEYSKAL THOMAS E. BOWMAN Plants Vertebrates DAvip B. LELLINGER RICHARD V. VARI Insects ROBERT D. GORDON All correspondence should be addressed to the Biological Society of Washington, Smithsonian Institution Washington, D.C. 20560 ALLEN PREss INC. LAWRENCE, KANSAS 66044 OFFICERS AND COUNCIL of the BIOLOGICAL SOCIETY OF WASHINGTON FOR 1986-1987 OFFICERS President AUSTIN B. WILLIAMS Vice President KRISTIAN FAUCHALD Secretary C. W. HART, JR. Treasurer DON C. WILSON COUNCIL Elected Members STEPHEN D. CAIRNS RICHARD P. VARI MASON E. HALE STANLEY H. WEITZMAN ROBERT P. HIGGINS DONALD R. WHITEHEAD TABLE OF CONTENTS Volume 100 Alvarez-Noguera, Fernando. Pseudothelphusa mexicana, a new freshwater crab from the State of Guerrero, Mexico (Brachyura: Pseudothelphusidae) eee Alvarino, Angeles. Spadella pimukatharos, a new benthic chaetognath from Santa Cat- VINO) VISIR N ove bame Gee 1 Es oa 00 t= We luemme egaee ee ween a tNaet mee erg Oa POC aI Rea san ALT od OR ele NO Barnard, J. L. and G. S. Karaman. Revisions in classification of gammaridean Amphip- odan(Gmstacea): pant. .3) sx. see toe eS SA eas ee eee ee eee Barnard, J. L. and James Darwin Thomas. New species of Neomegamphopus from tropicaluAmenica (Crustacea Marines Aimap ini © c/a) eases ee een Barnard, J. L. and J. D. Thomas. A new species of Chevalia from the Caribbean Sea (@rustacea: Am php oda) ss ee adhe Ee a res en eee ee Bayer, Frederick M. and Jeffrey Stefani. New and previously known taxa of isidid octocorals (Coelenterata: Gorgonacea), partly from Antarctic waters. Becker, Jonathan J. Additional material of Anhinga grandis Martin and Mengel (Aves: Aminigidae) trom theplatewNeOCeme xO 1p ll Oral ayes eee acta cee an as Bishop, Gale A. Dromiopsis kimberlyae, a new Late Cretaceous crab from the Pierre Shaleof South Dakota eo sete a a a eck ee akc ee Boothe, Billy B., Jr. and Richard W. Heard. Discias vernbergi, new species, a caridean shrimp (Crustacea: Decapoda: Bresiliidae) from the northwestern Atlantic... Bowman, Thomas E. Bahalana mayana, a new troglobitic cirolanid isopod from Co- zumel Island/andithe Yucatan: Penimsulass VMiexd Goya n eerste ee enone Bowman, Thomas E. and Thomas M. Iliffe. Anopsilana lingua, a new freshwater trog- lobitic isopod from the Palau Islands (Flabellifera: Cirolamidae) .o..........eccccseeeeecsseeeeeeeeeees Brinkhurst, Ralph O. Notes on Varichaetadrilus Brinkhurst and Kathman, 1983 (Oli- eochacta: Tub ricidae) 2 oe Pe, Se cea ead eg ae eee a Cairns, Stephen D. Conopora adeta, new species (Hydrozoa: Stylasteridae) from Aus- tralias the first knowarumMattac lie dis try als tre set cl ements anna nau oe cn Child ey Allanehe Ry cnogonidabypesko lille v2 Nile let al) eee Child, C. Allan. Ammothea verenae and Sericosura venticola, two new hydrothermal vent-associated pycnogonids from the mortheast Pacifac oi. ecccccccsseeceecesssseeeecesesssnueeseeeesenenees Child, C. Allan. New and little known Pycnogonida from Antarctic and Subantarctic WALCTS ec cccsccic ce ao, SNE Me ah i i aaa I aaa ia ern ied Se RI. neo eee ie Clark, Janice and J. L. Barnard. Chono angustiarum, a new genus and species of Zo- brachoidae (Crustacea: Amphipoda) from Magellan Strait, with a revision of NGO NVA US COTS ea I Sie Se ie ee Lar LE le Ee Clarke, J. F. G. Range extension of the genus Asymphorodes Meyrick (Lepidoptera: COSTMOP ter 1A ae)) see ke a SO INE A ea eat ee Beta teem eS eee Cressey, Roger and Patricia Nutter. Reidentification of David Causey’s Caligus collec- tons'(Crustacea:sCopepod a) wee we ests cos teat miate a aot Ne We UN os Nine ore ete ase ee lees Cutler, Edward B. and Norma J. Cutler. Revision of the genus Golfingia (Sipuncula: Grol fig iae) ooo ea ne ae ela ere De AE | Re SM Ma AN Dojiri, Masahiro and Jiirgen Sieg. epee fuscina, new species (Crustacea: Amphip- oda) from the Gulf of Mexico and the Atlantic coast of North America, aud partial redescription of Lvarlantisie Mill ssel\S Oy/mmere ts eee eee ee ee Ernst, Carl H. and William P. McCord. Two new turtles from southeast Asia..................... Erséus, Christer. Records of Limnodriloides (Oligochaeta: Tubificidae) from Vene- Fauchald, Kristian. Redescription of the genus Nauphanta Kinberg, 1865 (Polychaeta: FEU CV GAC) ie eA 1 79 ae ae ag sae ee Feduccia, Alan. Two woodpeckers from the Late Pliocene of North America....................... Fernandez, Justa M. and Stanley H. Weitzman. A new species of Nannostomus (Te- leostei: Lebiasinidae) from near Puerto Ayacucho, Rio Orinoco drainage, Venezuela 1-3 125-133 856-875 147-163 ~ 532-542 937-991 358-363 35-39 506-514 659-663 347-352 915-517 141-146 552-558 892-901 902-916 75-88 596-599 600-602 735-761 494-505 624-628 272-274 375-380 462-464 164-172 Ferraris, Carl J., Jr. and Justa Fernandez. Trachelyopterichthys anduzi, a new species of auchenipterid catfish from the upper Rio Orinoco of Venezuela with notes on 7. GIB GE RULES (INCI © 1p) ere es ARN OS DNS Eran woe Phil eal, AACR emit te. tls chevy Mowe tld At Yeon, Fitzpatrick, J. P., Jr. The subgenera of the crawfish genus Orconectes (Decapoda: a carta Atel AAS) eee neem nee a ee eo iene deal veh a ee iene ed Dee fel Fitzpatrick, J. P., Jr. Fallicambarus (Creaserinus) burrisi and F. (C.) gordoni, two new burrowing crawfishes associated with pitcher plant bogs in Mississippi and Alabama Wecapod aa Garb aii Gac) pisses tials Ts ten. eee be eareen ts oe tect elas lenente ae ep ht ie Gable, Michael F. and Eric A. Lazo-Wasem. The caprellids (Amphipoda: Caprellidae) of Bermuda: a survey of specimens collected from 1876-1987, including cave inhab- itants, and the description of Deutella aspiducha, MEW SPCCIES -...iui.....-ccssnneeeeeeecesccsnseseeeeeeeeeeseeeees George, Robert Y. Storthyngura torbeni, a new species of hadal isopod from the Puerto Rico Trench and an hypothesis on its origin (Crustacea: Eurycopidae).... Goeke, Gary D. Amphipods of the family Ampeliscidae (Gammaridea). VI. Ampelisca macrodonta, a new species from the Falkland [slams occ ceeececceeeecceeeeeseeeeeeeeeees Goy, Joseph W. Microprosthema emmilltum, new species, and other records of steno- podidean shimps from the eastern Pacific (Crustacea: Decapoda) eee Graves, Gary R. and Mark B. Robbins. A new subspecies of Siptornis striaticollis (Aves: Eunniarmidae)dnom! the eastern slope of the Andes... Grygier, Mark J. Classification of the Ascothoracida (Crustacea) 0.0.00... ccccccceeeeeeeeeeeceeteeeeeee Grygier, Mark J. Antarctic records of asteroid-infesting Ascothoracida (Crustacea), in- chichnoeammewacenusrot,Ctenosculidae wise 0) cu te A ek oy is eet Harrison, Elizabeth. Type specimens of freshwater Ostracoda described by Norma C. Furtos in the collections of the National Museum of Natural History, Smithsonian Utes ois C5 1(@ 1 emanate etal hose Del ent Eva DY Yon te Pov eee AAI an eon Th a ene wll bone et 9 Harrison, Elizabeth B. and Thomas E. Bowman. Mysidium rickettsi, a new species of mysid from the Gulf of California (Crustacea: Mysidacea: Mysidae).......... Hendrickx, Michel E. and José Salgado-Barragan. A new species of stomatopod, Eu- rysquilla pumae (Crustacea: Stomatopoda: Eurysquillidae), from the Gulf of Califor- EDL, WANG SSC] eterna Se ole eR ea Re OE et ee a Hershler, Robert and Glenn Longley. Phreatoceras a new name for Hadoceras Hershler and Longley, 1986 (Gastropoda) non Strand, 1934 (Cephalopoda)... Hershler, Robert and Donald W. Sada. Springsnails (Gastropoda: Hydrobiidae) of Ash Meadows, Amargosa Basin, California—Nevada............... pratense Maybe Un inet Lata ee Aten Hobbs, Horton H., Jr. On the identity of Astacus (Cambarus) mexicanus Erichson (1846) and Cambarus aztecus Saussure (1857) (Decapoda: Cambaridae) with the description of Procambarus olmecorum, new species, from Veracruz, Me@XiCO......ceccceeecccccseeeeeessseeeeeeeone Hobbs, Horton H., Jr. and Arthur V. Brown. A new troglobitic crayfish from north- Mestemmankansas, (Decapoda: Cambaridae) 2.24.28) 28 se hd ee ee Hobbs, Horton H., Jr. and Mike Whiteman. A new, economically important crayfish (Decapoda: Cambaridae) from the Neches River Basin, Texas, with a key to the SMS CIN SMILE UCC TIVO GTUIS i: se site <2005 dace nah tees, na blve neh ss Due eestomile Lueie yA 9 Machen g Wien ie Holthuis, L. B. and Raymond B. Manning. Hypoconcha parasitica (Linnaeus, 1763), a senior synonym of Hypoconcha sabulosa (Herbst, 1799) (Crustacea: Decapoda: JES FEVE GU EOREY) cece entre test ten cast Be re eR ees BS ee Pe On eR er Be SO 9 Johnston, David W. and Daniel I. Winings. Natural history of Plummers Island, Mary- land. XX VII. The decline of forest breeding birds on Plummers Island, Maryland, MEG) WARCEODUTENY eccrine eS cane tlap sl ep OE BIS a resN On, 2 een Te RON, Ts nD. eae een Jones, Meredith L. On the status of the phylum-name, and other names, of the vesti- RETCEN IMC TeATEUUSCR VOR Gece. eon el eae iain Pe NWN aed AOE leas SL ad Kenk, Roman. Freshwater triclads (Turbellaria) of North America. XVI. More on sub- teianean species of Phagocata of the eastern United States... Kennedy, Robert S. New subspecies of Dryocopus javensis (Aves: Picidae) and Ficedula hyperythra (Aves: Muscicapidae) from the Philippimes cc cceeeeeeeecccceeceeeeceeecennneeeeeee Kennedy, Robert S. and Charles A. Ross. A new subspecies of Rallina eurizonoides (awesekallidac) from the Batan Islands, Philippines... ee Kensley, Brian. Further records of marine isopod crustaceans from the Caribbean .......... 257-261 44-74 433-446 629-639 68 1-686 4-7 717-725 121-124 452-459 700-712 371-374 674-679 529-531 402 776-843 198-215 1040-1048 403-411 1017-1021 762-768 1049-1050 664-674 40-43 459-461 559-577 Kensley, Brian. Harrieta, a new genus for Cymodoce faxoni (Richardson) (Crustacea: Isopoda: Sphaeromatidae) i. 2 anette Be el 2 eee ee eee eee Kensley, Brian and Paul Snelgrove. Records of marine isopod crustaceans associated with the coral Madracis mirabilis frorm Barta... ecescesssesscsseeeeeseceeceeeessonnnnsessseeeeeeeesenncnes Kensley, Brian and Marilyn Schotte. New records of isopod Crustacea from the Carib- beansithe Hlorida Keysyandithe Bahamase: ewe ee eee Keppner, Edwin J. Observations on three known free-living marine nematodes of the family Ironidae (Nematoda: Enoplida) and a description of Thalassironus lynnae n. SOM IMOM) NOFth west FLO ri cae eke Ml hee 1 ire ee a De Knapp, Leslie W. and Thosaporn Wongratana. Sorsogona mellanoptera, a new flathead from the northern Indian Ocean (Teleostei: Platycephalidae) oe eeeeecteeeeeeeceneeeeeeee Kornicker, Louis S. Eusarsiella thominx, a new species of myodocopid Ostracoda from the continental’ shelfiof southern @alifomia] = eee eee Kornicker, Louis S. Supplementary description of Cypridina americana (Muller, 1890), a luminescent myodocopid ostracode from the east Pacific eect Kornicker, Louis S. Ostracoda from the Skagerrak, North Sea (Myodocopina)................... Kornicker, Louis S. and R. J. Palmer. Deeveya bransoni, a new species of troglobitic halocyprid ostracode from anchialine caves on South Andros Island, Bahamas (Crus- tacea Ostracoda) sss isis Te ce ee ee Oe Ie Scan Kritsky, D. C., V. E. Thatcher, and W. A. Boeger. Neotropical Monogenea. 10. Omo- thecium, new genus (Dactylogyridae: Ancyrocephalinae) and two new species from the Piranambu, Pinirampus pirinampu (Spix), (Siluriformes), in Brazil... Kudenov, Jerry D. Four new species of Sphaerodoridae (Annelida: Polychaeta) including one new genus and three new species from Alaska cece eeeeeeeeesnnneeeeeeeeecnneeeeeeeeestnneees Kudenov, Jerry D. Five new species of Sphaerodoridae (Annelida: Polychaeta) from the Goullh Of RS KC ee ER TENE ahs eI ee EE ete ae Kyte, Michael A. Stegophiura ponderosa (Lyman), new combination, and Amphiophiura vemae and Homophiura nexila, new species (Echinodermata: Ophiuroidea) from the R/V Verma Collections i. 22 f 8e s BU) UE ae ae ae ee ie 2 aa ee Lazo-Wasem, Eric A. and Michael F. Gable. A review of recently discovered type specimens of Bermuda Amphipoda (Crustacea: Peracarida) described by B. W. Kunkel CSB 2 TOG) see aaa Ee ITN Ae nl Aa pee ROR eR Lee, Welton L. Guitarra abbotti and G. isabellae, new sponges from the eastern Pac- Lynch, John D. and Roy W. McDiarmid. Two new species of Eleutherodactylus (Am- phibia: Anura: Leptodactylidae) fromm Bolivia 20... ccccccccccccsssssseeeeceeeceesscsssnueeeesseeeeeseesetosnuessseeeeeeee Manning, Raymond B. Notes on western Atlantic Callianassidae (Crustacea: Decapoda: ‘VRalassumidea) 1c it Greasy ea eek rn Ec RA eaten 2 ee ee Re Manning, Raymond B. and Brian Morton. Pinnotherids (Crustacea: Decapoda) and leptonaceans (Mollusca: Bivalvia) associated with sipunculan worms in Hong Kong. McDermott, John J. The distribution and food habits of Nephtys bucera Ehlers, 1868, (Polychaeta: Nephtyidae) in the surf zone of a sandy beach eect Mikkelsen, Paula M. The Euphausiacea of eastern Florida (Crustacea: Malacostraca).... Milligan, Michael R. Marine Tubificidae (Oligochaeta) from Puerto Rico with descrip- tions of two new species, Tubificoides aguadillensis and Heterodrilus paucifascis .......... Modlin, Richard F. Heteromysini from Grand Bahama Island: Description of Hetero- mysis agelas, new species, first description of male H. floridensis, and notes on H. guitarti: (Crustacea: Miy sid ace a) ssi tee ace ce sae ee cle ee Modlin, Richard F. Heteromysis kensleyi and H. coralina, new species from the shallow waters off Looe Key, Florida (Mysidacea: HeteromySimi) i ..cccccccccccsccccccecceceeteetttttcceneneeee Morgan, Gary J. Two new species of Paguristes (Decapoda: Anomura: Diogenidae) from southwestern Australia .c...2.../) 7 Nees See eae eee Coe ieee A: eee Olson, Storrs L. and Evgeny N. Kurochkin. Fossil evidence of a tapaculo in the Qua- ternary of Cuba (Aves: Passeriformes: ScytalopOCidae) ...............ecceescceeeeeeeeeeeeceeeeeeeeeeeeeeteenennnee Reid, Janet W. Some cyclopoid and harpacticoid copepods from Colombia, including descriptions of three: new species 2 ee | Te eae Reid, Janet W. Attheyella (Mrazekiella) spinipes, anew harpacticoid copepod (Crustacea) from Rock Creek Regional Park, Maryland 1036-1039 186-197 216-247 1023-1035 381-385 134-140. 173-181 876-89 1 610-623 8-12 917-926 927-936 249-256 321-336 465-479 337-346 386-401 543-551 21-27 275-295 480-489 296-301 653-658 726-734 353-357 262-271 694-699 Robinson, Harold. Studies of the Lepidaploa complex (Vernonieae: Asteraceae). I. The PINUS OL CILOCED IL EIVUTIL SC IM TSP sooo eee eta a ag ae et mc el a Robinson, Harold. Studies of the Lepidaploa complex (Vernonieae: Asteraceae). II. A INNES AMUSO EGIIEI2 OGO IA 11 Cesena OL fo ea De a a ee in et Robinson, Harold. Studies in the Lepidaploa complex (Vernonieae: Asteraceae). III. inomewacenera, Cyriecymura and Mirmocepnala ne ee Ross, Charles A. and Franklin D. Ross. Identity of Crocodilus mexicanus Bocourt, 1869 AARC Po lillie @ROCOG IAS) ae oer ae aa scp catecsnsecesedecanasnastelsvesasectensceatiecononsuasberobluccreeeStdccec Matas Seiglie, George A. and Mary B. Baker. Ammoanita rosea, new genus and new species of deepwater Trochammiinidae, Forammimiferida cece ceeeeeccnneeecenseeennnneneeenenecnnnneee Shetler, Stanwyn G. and Susan K. Wiser. First flowering dates for spring-blooming plants of the Washington, D. C., area for the years 1970 to 1983 occ eeeceeeeeeeenneeeee Steadman, David W. and Marie C. Zarriello. Two new species of parrots (Aves: Psit- tacidae) from archeological sites in the Marquesas Islamds. eee Timm, Robert M. and James S. Ashe. Host and elevational specificity of parasitic beetles (Amblyopinus Solsky) (Coleoptera: Staphylinidae) in Panama eee Thomas, J. D. and J. L. Barnard. The Indo-Pacific Audulla chelifera reported from the GanibbeanvSeas(Crustacea: Amphipoda) #252. Thompson, Fred G. and Thomas M. Iliffe. Two new operculate land snails from the POTEAU, ARC DINOS! VANG) ee tees re OT IN a ee Oe Ei LS ee re Vari, Richard P. Two new species of curimatid fishes (Ostariophysi: Characiformes) POM—eRConGrande:dorsul. Brazil. ee cate Vari, Richard P. and Ann M. Williams. Headstanders of the neotropical anostomid genus Abramites (Pisces: Characiformes: AmOStommidae) oo... cece eecceecssesneeeeeeeeeccenneeeeeeenee Weitzman, Stanley H. A new species of Xenurobrycon (Teleostei: Characidae) from the RAO IV WeaATIM@ NEMO ASTM Ol ESOL V 1 ence arenes cunadeeccunecovennensteucle Weitzman, Stanley H. and Richard P. Vari. Two new species and a new genus of miniature characid fishes (Teleostei: Characiformes) from northern South America... Wetzel, Mark J. Limnodrilus tortilipenis, new North American species of freshwater mupmcdae (Annelida: Clitellata: Oligochacta) eee Wicksten, Mary K. Nicoya tuberculata, a new genus and species of spider crab from EaAceICeOStavkica (MajiGae: Pisinae) 2 Williams, Austin B. Upogebia synagelas, new species, a commensal mud shrimp from sponges in the western central Atlantic (Decapoda: Upogebiidae).... Williams, Lucy Bunkley and Ernest H. Williams, Jr. Three new species of Renocila (Crustacea: Isopoda: Cymothoidae), external parasites of coral reef fishes from the AUR TIMLS ATI GSEONO I a Palast er andl ee Ce eae ee aie ete ee SE Wolf, Paul S. Two new species of Pettiboneia (Polychaeta: Dorvilleidae) primarily from “5 SHOUT OIE 1A (GaSe tek en a ec te CHUAN HI LA eds ORNL fe OTM Sonat ene eer Wongratana, Thosaporn. Four new species of clupeoid fishes (Clupeidae and Engrau- Gao RInOTMPAUS tralian waters: ae cece Yager, Jill. Cryptocorynetes haptodiscus, new genus, new species, and Speleonectes ben- jamini, new species, of remipede crustaceans from anchialine caves in the Bahamas, WItMeneMmManks ON GIStribution and CCOlOGY cece ee cc ceecccetnenccaeesennnnncesssece Young, Martha W. and Henry Kritzler. Paraeupolymnia, a new genus of terebellid (Eomchiaetasherebellidac) from Belize 578-583 584-589 844-855 713-716 490-493 992-1016 518-528 13-20 364-370 447-451 603-609 89-103 112-120 640-652 182-185 691-693 590-595 417-432 28-34 104-111 302-320 687-690 INDEX TO NEW TAXA VOLUME 100 (New taxa are indicated in italics; new combinations designated n.c.) PROTOZOA Foraminiferida PALIT OG ATE CRE cos ak ae Ee a a fe ae Rae Se ee TODS CD i ae cc ict a in ding DOR Nek eg AMD 2 SM SNA ac SY A a8 TIT UC ACE EDS US TCC hse Sh i la al le care OM ae, LU PORIFERA GoUPAT TAXA DDOLET sic wel sac Wl a SU A AMIN 0 as Al Oca ee ESA CHC «00s. sxe Bini ats SULA es as SS ade Ne RSE So AN Se ec COELENTERATA Hydrozoa CONOPOFa QLELA scsi st LI alae OTe a OR ane ee ec nT eta ee eee Gorgonacea DAVES UH LESUS eet uN ee ede te obi ee eRe ae ERNE RA na Le cE PCa 9 Rn SCUTINCNLOSG ccc ee cre Noss el ALS TIS RS da IR RO ae SNC ce et ee? GOT UUSTS ooo ceca reo a EO DE ot Ie SEIVUION ee eS SE BAe ae ee Rede LOR et 8 C@helidonisis aurantiaca M1eCXxTCANGE ee ee ae ee hee eee oR DIDLLED DET OTISUS. cee soe eR TROL I et eae OT EIT a ee ECHUETNES1S CULO TOUTE nee es de UII as con DETSCDNONE ore Te Ie A gee CLE IED eee ol ee ae re eer ee VOTIVE oe ee Ec LE ETO eT EG MS OT BTM NGISIS PLIAGE rake eae Tena nee TS ee oe a POY EA (1 IS Io le na i ei pe Eee ce a ilar Ae co easel yee nen i ela we Ye tea ea HR CTUULUS TSI ON ae Are ees Clete Lae nace CE Oc Pe ee ORO ea ee ON SIO ee es ee PLATYHELMINTHES Trematoda Onothecivinm cs elie iu Moet SN reais, eek > ARUN ca ee tee LIC tt Loe eae Tard eee DCH SN aah RN a re bo Sy ae ia me en EUR tees 2A Uae TR a ID oh EN ete DENTE GUND 222 TEI N,N RINE 5S AS, SI SRN Pel 1 IL Re ee a Turbellaria Phapocatia Ot orchis: 2 We tve 58 Pet litt NE A eek 4 tee ae ln Rg EPS eM ee ee DY INCL is ea aR TING Sap Rae REL Mead dc RNA E ONO em MAU Pred h SPV UETU Reis MOAR ts A ROI PIRI IEA a et el EN TI ee atl a Wee ee tle Jd Zon NEMATHELMINTHES Nematoda Mhalassironus, nnGe. es wo ee eee LC re rade YOR Ue aaa Cael cal nee Fc Nas Damasio eee ANNELIDA Oligochaeta Heterodrilus paucifascis Amn OGLiNYS COP PENIS S20 NC APS AE SSL ad tM Ae the SIR Sl eared ey ade Sco oem Tubificoides aguadillensis Polychaeta ANEFSUCLET ODA DUP ec SS SPE MOSS le SS tee ET ee SIPC EO Sn 917 [cE EG DULL ICUS DOEAOE cha Sa ES NSO a SS eee, SSeS Mr eo EN etc Ee er 918 Berl @oTREU NUNN) 72 CCL CCUIT CLIN pte ata ea ee en eer a ep eee eee ae acted wd cclnnausletoces 927 TE SLi SIIY (DCI DUN CIA) wos he NN AU ne EC eee a eee tcl ane ene 929 2 ES GUID ON SAMO cca a Se RA sc a de uk Sees AG tte Eset Te Neate PLONE Rone 687 (HEN IYS ncecesne ea SN RS SS and ee JU Ce eI eee 687 TEPRETIDOST EL DULL act a Sea ge oe oe Re Sa > 32 DW OG VIP OD cxconecscest el rsh ais I ce te sad ae see Ae Sa at, Rate es cae ee eth Rn Ae ana ET Dy) PIMP AG GPITS SUAM CZCEC 111 CL Bieta eee eo tan deee cee nr tc cete acta gh Coors Ma Does to ed cnet capenaeh inna emsereescnnse Seon 930 Bee BRETCT GS TCOGECHECUIUNNTIN) 20 2CM1 cree ean Sune arte eee ete cere etre ee a een 932 IRESROCIOM ODES. KALINGA CT. (1 a eae etna O12 li ee 0 DSP e eee OUR oe OTE ESTEE SUN AO 922 PIZI TIDE RZT OSHS sce 0Ny date a PSS RDA RT Se Be EP SOA ANS PE 919 WUD IA x conocncenneseroneenege aaa bao Soe a sate 1 NA Ie ee ANS Ween hash Ra aC Cra em a Pe i ST RESPIR 933 SIPUNCULIDA Missae) PTS AR OS 71170211) ro eter ae rea ny a eaten er coe a ge a ee 740 CHAETOGNATHA ee SST Ue VL CULT ccc EN Ae IT rag hia eee OP evn ee le 125 ARTHROPODA Crustacea Sf nna VR Nel ee a a 860 Se RASS AMIE CE GOL C) 11 CL meters ete i li ee Su i UE Cote lt oe Ee a 4 2 LEEDS GQMA CTS G cert tc sks ol eT SO TI at EE ne PR SN 239 Seem BeeR RA SAN CAT (77 Si] renner ca ee fe NE | ge cee ula eA a a lel ee elie 565 [SVEN sceoccarcoacsenpocsttvetinss tert heed APA NAM See ANREs APA MERTON TSR SO Ce IOS Oa MEER RE OD ere OPS ALN 347 ASE LIS TAILOR DS IO seer area tap BT ONT et aI i re ETS (i nm eee EP 456 Eee eI MIWA AZ KIC IEA) ESI OS: mn ke 694 SOSA GUOND, a cnomenenmnatenenets ent ete VaR gia cE SERB SBN LR TEN tee ee DEE CE a PETE 869 SeamYcA UA CIMNIT C21) (01) CL rma tale ee A eR A een dal be SAL net Neal pean csp hte ele 659 EDULIS IOIIE cc cncoceent merce DIMM A tN IIe ER ONS ORDER SC TNE OE I a RN ORE MP Ie RTE SA ARETE 868 USN SLT DDS cc sccmmccmntt teats oS ee eo SD cE Te Se a EEE eo 864 SU ATMM SATA S ACU UC ATINDATUS) CULAD IIA aaa cenccccssccccccesebcbecebeslUStccccblsccccoeepepicleccvites tlle eine assess 1040 SOE CTT ITLL (2 Cm eta NS NIE RE DOE OL a ee ad shat A TIE A ON ee Nc 858 Se A ECAWENI) OAMTI7OS (LUC Cr Ruma Ian ete sist NN aed ak Sn lls alice ah nia nae ace erie ete a a aL 243 77S | | I Le NR Ras ie hey ac I init nore etna ao anlar wh he 865 ULE MMBNCARIE ITN NcAW] CAV) S Iemma anu Pose ehh merece ds ell ee nea ie een NL Nema aie gn 187 ee Yeserict AN ACE 1 7) 1] me ee i eS See ee Le ei Me 541 UG DID concn cccanoseceecccecerecencecrsincted ttt 1 TE NG HCI IE SAE AT aT A SE OREETET ETRE is fete 18 See TI 78 PLIST TAR examined Mit Net ee 1s ep oe ISP aIEE DIN REORSN re ts 20 he a Oe eS 79 NE TIDI OIL nner omneznennccenennets anette tea a ep SERS oN as ODT ROR PN redeem Cee 865 Me ORLCn CRIT ATU(Z) 11) Cm Re ee Mh ee a oe ie amas 22F BECHTIGILLCHS (| meee ora iti aeons otk ti Ade SL ne Mee Sar D 2 I a ia 236 “TITLING NODE sr crorammncin betta hy A eRe A eG ed CE oN ie OO OU Paka st ne 859 ered) CEILS 5 (am oes Lee is TE ee eee eG te ee ei aa 392 ay PTY HACIA LCL CS ene er Ns ea Noah a Wine eo aN a en ul lo A ane 310 ECE HOLLIS GUS Pena RIES So ics a eat NA 2k Es de SNe ial PIN a 310 BREN AMIS COUES (1 Imma esc ate tae Lal oe Ne ee a Oe el 611 ME RENN ASC MmEIES LID ONLNID sites, itechnenee 231 08 os els aN a tea Asal teh ease A ae 2 eA alc aye 702 emma Tact) (ASP DIRAIID) Atee rs aeertn le ee ae ee oe ee eh ot cl a | i ee ARUN ale 456 ETL) GUS WIIG NG sah cee AP AEE RI ED ure DUE ae ROE eee 630 LYLE WUD ATE tect csp nae emi Rc ean a eso eee a DAE ae aE ee PRR Oc 506 CTLAGATEAS SYOU/SI eres de de sa RE ee eRIRE, SOO eo PTR Te oe EER EOE ccc EES RI ae Oe 884 CUNEO ABALIGJBRSIG, AL OY AAA ICLP seca terre) ae ORE IR SD RIN ante al i Nala By a oe 35 TESS TISIB SGD creep etek SN Oa OSV AC ALE Ect Pe vec Lt Oe 190 Blapliordel ata dhe sachet cere cee ae 267 SUL QT OZ ec act ha ek Oe ek ee Mae eR A SR 32 Se Pee eee ee RO 263 FSU YV.GLCC CVS OFUGE 8 i ce A a acc cee UR Re 568 FE UAT SCUUME ea gp 00700 chee lel i eA a aR een Eres ra ase 529 FoUSarSie ll age Ve Ut cco a cee ce Oe TEA I eee 134 Eallicarnibanurs:(Creaseriey us) ict Sioa oe cet ate eee ee 433 ECONO (1 a SONATE Oe DE eA rn NE S RIT eR AY Se Nc ae om ee 439 Fallicambarus (Eallicannbanus) ia eve stare reer ee eer ae ee 403 CCL ET CUT ak a Sa a 2 A OR ae Ne PEN ae ce 866 (COMPLY OP SCHTO kos IIe Fea Se OSA AE a a BOON Peete OEE 705 SCL OS SUT 3: ot OA CIC sects dsce ch Sosa eee 2 eR need Den eR deen 705 FV YI TCL MR ce ala i SR eee A Le ac a Ue ver AE ale ei ae emcee Re a a Ne 1036 FAG UST ONIN SISHO LOLA 8 ee ae Sa a cae TO eR SA ere 296 COTA LCRA L I st BINS aN eg Se A le a De a a Cee A 657 [PARSE eee ee een emer ree we tele ees et Rs eee ee er 653 EL SORIA GS Mee Me ete 2 aust en AN a ee ree ete a rec ee PA 872 CTETUCL PB UTUTATNe Ch mr 8 Ta NAS a Re a a a 543 IS PDULTUGUCLCLTRCE Sh a cB SN SRI a Ua ad Po el ee 544 |G a¥ero) 51s) | CW (VRS) 797 eee eens erste Ne ASIN SHORE NO re PO ae eomeet RIO eee Mee ane tO ean ole I 495 SUV TUS ee 0h a a oe TO 864 TRAYS CE OPK. cent te ea illo ag ls BOR oe Ue acl ata eee 870 TTA GRID Ani oI 5 Sab No MRE Bie SE A Oi eT rk SG a 453 TLC DUQUE ISUCS I ec ik le at sg Re ea Ph el 866 TE VCE TAURI ekg oe th Cc ae oe Pn sen Ne cen rnd 2 Te ce eR ea Te ae Cee 216 CETUS LCE es ia cal Sat iol a i ee Oe Ew A i ec Di WVAGTLCT. OS OTION A so i i a ee FN am 860 NAC YAU ELIG D UES eI ake NS a a ar a re cael na te ee 861 IMS Seana DEV ELECT Clic sc hcl ack Ng ce Rc 559 LO OCHS TS aa Is gS SI rat eA IR Pl ae Ge Se Ne 220 MMI GEO PKOStIE NINA Grrr T2E LETITIA eo ag WV ATTT OL CGI ORI I ie hk Ss Na pt a ah EL CE oO 562 DEUS CO i i Mn a ine ee I 562 IVA y STUNT CIOL ES Ute alo ad) Lal ap 8 nes en ads et ts 674 IN GS SCV CUO os ace ct NAN ete St al lw oR IED a Re 862 ING OMS RAIA PTO PUS HRCA E nt ak aca ecto la a Oa 159 ULCER a na ta a eh RAT nl auth a Ee et eR AA Ce Oe POR 152 1571101 {1 1) Seema eae ane ee Reo) eee me NTT Pee TERE ea ce ie A ha 156 PITS UTTAR Oe ee LEME aE OME ONO O RENTS NOLIIOE WONT ie NNR FAUNAS ry ME AT ici ees 157 INTEC OY Ces oN i A I A aa RN Nl nm ea a NL BR 691 ELULD ONC ULL AL se Re Es Ssh NI aad 8 yh eR De a aN MS aL Hm Race 691 Onrcomectes CBU lec e Sy e cae cn ae al ge e 47 UB RI VENA TUES JG 1 TRS) Wen et R a2s NONSEOS SU NrNay ot BLT NGPES LP EUROS ee arty en No Roe eT ae AE. 49 COT OCI CNET UES) acta ted es are ade la a a LM a PND Its cD oD MSN FR ead 50 (GTI COTO ELIS ieee rote TIE a oh xR Pa ER Rd a en Al ees ea ree 53 (FIESPETICA IDG GUS) eee cree LINE Sa MR 28 Llc hgh = Or ea 54 (PROCETICAIAD GGUS is 2a oS ae oe eis Soe ae aN RB AD re A en ei pegee ee er 2d 57 CFR OGALES TUS) oes Bocce aA se ae AIA le LE nic ek Beat eal aL Ge eI A 58 OG eM TURVCTANG a1 070 Lac) ky einai te Dire Pie NS aa pt Tara SNe) We a elle OH INEM egy Le LE Ye 60 PapUniStes OMI SCLO SIS: coe cess ek teas ne Nace Ire eM RN sO ota rere te es PO 726 DUFDUMCQNLENITIQLUS 2 oie so Es Pee ek a SE ly AL ee ee 729 PAPACY ClOPS P71 OVCHGNULLS 0 oa cae gl acre NE geet Ss se bee 262 PAV COMM OIA CLANS, ce IN a lee ie cole ea amy Case ese Nee lato Seed a ee ae 226 Procambarus (Austrocambarus) OL CCOLUMN o.cccainccccscnecccsconweeeesovveeseeevvneseessvueeseessvneseesesvuessesssusesessouesesesouuesseesiuessesssuueseess 208 Pseudothelphusawnexican@.s ak ae ee Dn De eee ee ees i 1 RREMOCIA DOM Otte a ON eee ed a et ta ee Oe eg OD nee ce ce 427 OURO asec NS ae teh re og oe CTI Oe tote Coe en 421 VTA ZU OU Sea et Oe ok a hoes HR Oat tat ee es ae ee 418 RUM OAK OF ORG. ke ei Bae 8 ay St 5 eect i ER AL ORNS a iets ORR Eee en Le a oe 867 Baas aA CL SELICAII] CLI] Lemncs aan eran ene arom drec ines eA eee Ga a bee cae 302 DADE LCLLCUL Sperone eee ere ah tte lia Es a 869 ir aaesa HRM aT ULTRAM COTO CLI La ames te reer ee a SN ae ok aera er ee ads SO ee 682 ODE TELL AUGIED) cxcecertn eter a tte bea ce Uae ATS e Oa UTE NC ER Ee Cia oe SEEDED PE SEE ROLER OCS nee cD er rere 870 mea AU SEATIES DTI Cd Comm tie tra ace Minette tee eee eS wR ene Pe ese ak aN en Ae ta ec 456 EOE L GTR, SULCUS aaa la OR ear IO Ce Rec en 590 OSSD EEL QID OY occ cents et ee Be Sa PSE aPC nT RE EE rE 871 TE SEDI USSD smc tt as ERR PE te ORE em Ere ME RE EPO eR 863 JSD DIGG socom EE ee oS am ec PP TSO Oe fre an eee 570 RELELECTGC)| pa ee ae aE Me eee 2 eR ee ee ae SP LATE IDE OTUDID © cexecececeete nt es Ne UR A CE ROR Us SO RSI GS i CAN aR NB Oe 871 Insecta RMMIREETEINOROCC SUA ONCII mms cre set wren me ce Ve ee ae cle ee a allel 596 AEE G POU AEM OMOXESCasniC ste tee ee Oe ee 597 Pycnogonida STU SEEVS UUM, PROV LUC cect A oT I see SOE pln DD ls cla DP VS SO ER STR OL RCS 892 os AEG SRCLAS CBD OCIS aoe aU apes Mees ae Oe eT OU ees ede: Pea er CDEP 902 CHING secacescccescccurt tet eset Set ets aH Ste SE OOPS Ae Se 9 OE Se nS cn PPE Ra ce OE DEN 904 RA an Rs RTO C2 111, Cr mmr asst are are Sinaia ce NON Sele pet OC Vaal pu ee a A as 912 SPEER TECLU NSE /ITITI GI US rmmmremnte a teat sole WUE CRG Oe EN ee eae al ee a, 907 ELISVEV NS CLG ACD icc ge ae ee IE ONL IE Ms eS Sa RO eee rete 909 PESOS WLI ODUD aici SG AT A RTE i Pa eS Ra Er ETL 896 MOLLUSCA Gastropoda aan ssn CC) I ey Se FN IM lad REE ee ek SN a ol a aa ee re 447 PULP SOG CONDLS concer cenacpen is i Natt tL El a set RA coe Ted Ta ERS ER CEE ee TLE Ne TD 402 “EDPRTONG: PUOUOEIUIGL ccrinncred cen UN SUG meat em TO SO AE I I eRe EO ne nT 449 Nee ise AUNCHES SUSU Gla) SUC penne eee ee acs eth eee el a LEP Mai A ee ote ee ee oe a ae 797 Fi EEERUCIIAIGS C11 SS amen Pa Ul ect mae Be NOY NANA dy Lk Nea Dee 2 is MULE ests leila Aarau 796 ISENOTIVSS scrncwcceens tenon haskapt BC) ete BON .AOb eet ai ee sss NER ea SCID Tae Ae eT OSU MER RLS aon Oa 807 DUELS ae emerwecpceres ttt fhm Slt ESC Me EE BRUNO RD ae 20 ROUT UE ral cepts Sol EOP Tak Bl ee 802 [DUST QI ceces enn enmcseucee chose lp MAUS NWN g ONE oe hse ie SEROUS AR le Se NOE NO RROD REN RISER SRE os UIE DUNE PSI 804 TPS INUEL CHAU GID ccc SIRI Ce aD gE ED eee SN ROS Re PER IRL: ener 810 ACE GTR ie NN a Shed LEN oa rece hg IS Naame Galea ho Nee hen oN 831 GD -merecerececcccextetten tet tite, PA aa aU seems Pate Sted dhe) EP a DAP S eeell 5EM ts A eee Re 826 HONS EI DIY Tce sen est tO BOD a aS RE Ee A Sarre MN OR Ne er 817 ECHINODERMATA EOE WoL DFE Spy NINN IRY C111 Cpemrmen tatiana are aay Jie ae AN Me eM le eee has a ae D2 RaUAEL Toy PARES G1 Hammamet inh ok tab ee Mek oye wee ee ee LN SUES eat aes cee 254 Warr OSG BON TCIM [OOSELGL CLO Sel alie Cos ets sesence wee ee yO a ea an a a ee A, My CHORDATA Pisces INTIS: SU OHIOD coe niatatecal Sed A NO CLANS SNE SISCREI OTT SR DPS s OSS DIODE ALI sO ROSEN E PAS ARO oe AE 603 STTATUGSIOL exricoctr ess Nea ope OND REA AIS soe ieee A A RIL ELD ie ERE PO oe Ec PE Le 606 MENCIARALUST @ DEIN SECO LLCUL CL ier escesicne a a Nena eRe EL sear enc ke Aenea ene Wee pet yee ene eek Jat AA 105 SER ROSS CO UNILISECOTIC LIZ Crm nue Se nena) PAINS cane ION ota IIA PDR a de abel al SM Oe gh 8 i oe 165 2 NCL GIVI? sreceneecncncedicee ertth lrped att ALI OAM DOI em UID NTIMI oa, U5 Sope Fe Tet VE ea se -PREEOD Oe Oe PPC a SONS ome SRN FR 641 CE! CET TOO ALLIAGE Lib ds Ns che peau Ace See altar es aa Sle ic 2 641 J DTA UII ccm eet a NA Ss IE SEI AD ey OA ABO Da oh AE eee aL RO EE OLE LR 646 Sm RETUCAINT) CID! (7) Damm enre re teen lees re ey SNe me ee LU MN INL Me eae ua No 109 MOTI AMIITCL LOLI CN Cane menre ciara re ena tis eae ine ee a Nipe Aer Mis Dee ait tee Se ae ge, 381 Stole pH Oris GQ VOI 0 ccetecccsseccsccct ccd ccet Seca ete at ce 106 "TIRE Y SSE POI AST ccc asec ts ant a cee 108 Mirachrelty.op te rl Glnt lay S xe 11010 Cre eee eee ee eee eee ee nea ONES Cyd a ie ee 257 DEMO yCOM DOM I CISET US cesta oar lee reece ee a ee eo a eee ee 113 Amphibia Blewtherodactylus(/raudaror 222i OC Oe Cave See ee ee 337 ROK COC OS AO ec Manes oh Ba as NEE LN EAE am a Fo eg ee 340 Reptilia (CWOLALGHT SGT. AIT QI UTN cc 8h ee NO I ire es eee ee eee ae ene ee 624 Rlatystermonumegacepliallurmy:S/1ic¢t 2 5 ee ee ee ee 626 Aves Dryocopusmaviensis CeDUCNSIS 2) che to ee ees Te ee eal sia ec, Ae ea ee 40 RicedularhypemythravwQtulumiensis 22 Vesa eNO Ase ene ve aol eos ee ee 42 RallinareruizOnoides Ql vaQrezt ius 00 oe Rate CRRA ole feel ea lceetnedte Bye ote eter av ene 459 Siptonnissstmaticolllis Orton coc 2 Ss ee eee ee eee ee ee 121 VETOES 7171 OL O Tommaso en INE Pet a a Ae te I PO a 521 VELVET ane 098 MN te NER DEA A a Sean ie ec 523 PLANTES Spermaphyta Gi COGN TIT ick ON eee SSeS TREY TU aN ea ree 849 MTCC TC Meet lee a nS Se 851 102 1c a) i 1h Ce ol re ee 852 AMM PITOS AT Cos sate Nae hs TC Oe 852 TM ALLOS= SUVA IV sc 8 ote ye SIS eM RR Pk en 852 S721) 0°) 1 CA eS GPL ITTUeRS NR et ean Pn nC Tal en Oh eM CLI CLI I oe I co ert 853 SCOP IOLA SS aes Core ae URN cha uk ced En 852 TECRINOCOT VN ee ee Ra OE TC NES ae ches one Eee 586 Echinocep lala sy. Ci ee EE a RN Sele Sr pele ie ee Re a 587 HOLOSCHUCA INC ere eee IE ee ean ep 587 PUM EMS Ce ee Ee alt NS oe Se ee Ne Se ea eter cee Oe eee 587 Schiwenikidel Oliva 15 C eee SOO ae Rae ete Care ac aa cea lO an 587 SOGU CUA Cs ee ae CIS 1 URS Nek es ANT Na ea eee 588 SUnlUleatea TC ee Te Te ae teen ae ee ee pte ea 588 TE UITTOC ED IVQUG soo ere kN Ne oF atta tae eO eo tOe Oe ne Ee ee 853 Loy 2X) 0 ez bas 0 WY cia DEO la a mI Un aE LA Ua Nd ial, Pencil ded hn ees ee oo ge 854 CAIMARACHTCMSTS SEDC xe a oe Oe INE cle tee erie el GI ee nee 854 mae Bea pony il ara ee Na 2 eet Oe net ane sca ea ee an 854 Stemoce plana by Sere xc ses Sa eae laceration a ic vee he a 583 HLS Celbt evo [BN 90Ws ole commneaian inn mnelweet erat ie retanert MnumeNn Monn Ucn eel enue rn ice AU etn con ra 583 Y, f Vy MBE THE BIOLOGICAL SOCIETY OF WASHINGTON 1985-1986 Officers President: Austin B. Williams Secretary: C. W. Hart, Jr. President-elect: Kristian Fauchald Treasurer: Don E. Wilson Elected Council Stephen D. Cairns Richard P. Vari Mason E. Hale Stanley H. Weitzman Robert P. Higgins Donald R. Whitehead Custodian of Publications: David L. Pawson PROCEEDINGS Editor: Brian Kensley Associate Editors Classical Languages: George C. Steyskal Invertebrates: Thomas E. Bowman Plants: David B. Lellinger Vertebrates; Richard P. Vari Insects: Robert D. Gordon Membership in the Society is open to anyone who wishes to join. There are no prerequisites. Annual dues of $15.00 include subscription to the Proceedings of the Biological Society of Washington. Library subscriptions to the Proceedings are: $25.00 within the U.S.A., $30.00 elsewhere. The Proceedings of the Biological Society of Washington (USPS 404-750) is issued quarterly. Back issues of the Proceedings and the Bulletin of the Biological Society of Washington (issued sporadically) are available. 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WASH. 100(1), 1987, pp. 1-3 PSEUDOTHELPHUSA MEXICANA, A NEW FRESHWATER CRAB FROM THE STATE OF GUERRERO, MEXICO (BRACHYURA: PSEUDOTHELPHUSIDAE) Fernando Alvarez-Noguera Abstract.—A new species of freshwater crab, Pseudothelphusa mexicana, from La Jolla Cave, in the State of Guerrero, Mexico, is*des¢ribed. Ft doés tot -"— * exhibit any external morphological modification for a tro affinity with Pseudothelphusa granatensis is discussed. In July 1983, and January and February 1984, 11 specimens (5 2, 6 6) of a previously undescribed freshwater crab were collected in La Jolla Cave, in the State of Guerrero, Mexico. They were captured in an isolated area in complete darkness 120 m from the cave’s entrance after a 20 m vertical drop; this seems to restrict the distribution of the species to the deeper part of the cave. The presence of pseudothelphusid crabs in caves has already been reported. The genus 7)- phlopseudothelphusa, distributed from the south of Mexico to northern Guatemala, in- cludes the only three species of truly trog- lobitic American pseudothelphusids (Hobbs et al. 1977, Rodriguez 1982). Typhlopseu- dothelphusa mocinoi does not have pig- mentation on either the carapace or the ap- pendages, and the ocular peduncle lacks visual elements (Rioja 1952). Like the species described herein, Zehuana com- planata (Rodriguez 1982), reported from a cave in the State of Veracruz, exhibits no obvious adaptations to the cave environ- ment (Rodriguez and Smalley 1969, Hobbs eid l977): Pseudothelphusa mexicana, new species Fig: | Description.—Front of carapace without defined superior border, but limited by epi- gastric lobes. Inferior frontal border well marked and smooth. Slightly arched me- +4 MAR 0 9 1987 dian groove reaching- visible in frontal view. Café with smooth-surfgeesand snrall papikwereg= ularly distributed. Cervical groove deep and curved, reaching anterolateral margin of carapace. Cardiac and metabranchial re- gions with shallow grooves. Anterolateral margin bearing small denticles from cervi- cal groove to epibranchial region, number varying from 16 to 24. Margin between ocu- lar orbit and cervical groove sinuous and without denticles. Ventral surface of cara- pace smooth except on pterygostomian re- gion, which bears cylindrical setae. Ratio ischium/exopod of third maxilliped varying from 0.7 to 0.87, average 0.8. Chelipeds un- equal, right chelae larger; dactyl and propo- dus curved inward. Fingers of major chelae in male gaping. Ocular peduncle and cornea well developed, carapace with brown pig- mentation, walking legs and chelipeds sim- ilar to those of epigean species. Gonopod description.—In a caudal view the lateral process (fig. 24 in Rodriguez and Smalley 1969) changes to a caudal position at *3 of the length of the gonopod. Apex exhibiting well developed lateral lobe pos- sessing 3 acute projections laterally orient- ed. Outer margin of mesial process curved and serrate, ending in rounded tip at about same level as projections of lateral lobe. Straight inner margin projecting toward me- dian front portion of the apical cavity, with- out joining lateral lobe. Superior lobe re- z PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. duced to acute denticle as in P. granatensis. Apex of gonopod with oval cavity opening anteriorly and bearing 26 terminal pore se- tae. Internal border of cavity thicker than external one. The terminology employed for the gon- opod description is taken from Smalley (1964), and Smalley and Adkison (1984). Type.—The holotypic male is deposited in the Carcinologic Collection, Instituto de Biologia, UNAM (Catalog No. EM 3604). Type locality.—La Jolla Cave, State of Pseudothelphusa mexicana, holotype, left gonopod: A, Apical part, cephalic view, B, Detail of apex, cephalic; C, Lateral lobe; D, Mesial process. Guerrero, Mexico (18°35'N, 99°34'’W), 5 km northeast from Taxco by Highway 95, at an altitude of 1800 m. This cave was formed by chemical dissolution and always carries water in its deeper parts. Relationships.—As stated by Rodriguez (1982), few external morphological char- acteristics within the pseudothelphusids have taxonomic value. The gonopod, how- ever, provides the most important differ- entiation characters. The gonopod of P. granatensis (Rodriguez and Smalley 1969), VOLUME 100, NUMBER 1 Table 1.—Measurements of width and length of car- apace and fronto-orbital width, in mm. Males Females C.W. C.L. F.W. C.W. CJL. F.W. Sie M3351 DES DOES 18.2 17.1 ANG 12.9 NB 36.0 21.8 DA\\5)I 26.6 17.0 16.3 33.0 Wey 19.0 24.0 333 14.3 S62 21.4 AMZ 2 P24 16.3 17.0 36.0 DAY 20.5 S37 19.4 19.4 * Holotypic male. a crab which seems to be closely related to P. mexicana, has the following features: a) a well developed mesial process that ends in a rounded tip, b) a reduced lateral lobe with an upward projection that closes the apical cavity, and c) the exterior border of the cavity 1s higher than the inner one (fig. 16 in Rodriguez and Smalley 1969). In P. mexicana the mesial process is similar in shape and position but shorter; the lateral lobe emerges from the same region forming a blade which ends in three sharp tips; and the apical cavity is opened in the median front portion due to the separation between the lateral lobe and the superior lobe. Acknowledgments I wish to thank Dr. Rodriguez for his valuable help, and Dr. Soto for his com- ments on the manuscript. I also thank Jose Luis Villalobos for allowing me to examine the type specimens of P. granatensis, and Yolanda Hornelas for the micrographs. Literature Cited Hobbs, H. H., H. H. Hobbs III, and M. A. Daniel. 1977. A review of the troglobitic decapod crus- taceans of the Americas.— Smithsonian Contri- butions to Zoology 244:1-183. Rioja, E. 1952. Estudios Carcinologicos XX VIII: De- scripcion de un nuevo genero de Potamonidos cavernicolas y ciegos de la Cueva del Tio Ticho, Comitan, Chiapas.—Anales del Instituto de Biologia, Universidad Nacional Autonoma de Mexico 23(1,2):217-225. Rodriguez, G. 1982. Les crabes d’eau douce d’Amé- rique. Famille des Pseudothelphusidae. — Office de la Recherche Scientifique et Technique Outre- Mer, Faune Tropicale 22:1—223. , and A. Smalley. 1969. Los cangrejos de agua dulce de México de la Familia Pseudothelphu- sidae (Crustacea: Brachyura).—Anales del In- stituto de Biologia, Universidad Nacional Au- tonoma de Mexico, Serie Ciencias del Mar y Limnologia 40:69-112. Smalley, A. E. 1964. A terminology for the gonopods of the American river crabs.—Systematic Zo- ology 13(1):28-31. , and D. L. Adkison. 1984. Disparithelphusa pecki, anew genus and species of freshwater crab from Mexico (Brachyura, Pseudothelphusi- dae). — Journal of Crustacean Biology 4(1):127- 133. Instituto de Ciencias del Mar y Limnolo- gia, Universidad Nacional Autonoma de Mexico, Apartado Postal 70-305, Mexico 04510, D.F. Mexico. Present address: De- partment of Biology, Tulane University, New Orleans, Louisiana 70118. PROC. BIOL. SOC. WASH. 100(1), 1987, pp. 4-7 AMPHIPODS OF THE FAMILY AMPELISCIDAE (GAMMARIDEA). VI. AMPELISCA MACRODONTA, A NEW SPECIES FROM THE FALKLAND ISLANDS Gary D. Goeke Abstract.—An unusual new ampeliscid amphipod, Ampelisca macrodonta, is described from the Falkland Islands. It is characterized by a series of small spines on the posterior margin of the basis of pereopod 7 and a very large tooth on the posterior corner of epimeral plate 3. The proposed new marine amphipod was collected in 3—9 meters of water over mud and broken shell bottoms. Ampelisca macrodonta, new species Figs. 1, 2 Material examined.—HOLOTYPE (16.2 mm), USNM 216638, 23 Feb 1927, 3-9 m, Port Stanley, Falkland Islands, boat dredge over mud and broken shell bottom, coll. W. L. Schmitt; PARATYPE (9.2 mm), USNM 216639, 2 Apr 1927, Teal Inlet, East Falk- land Islands, coll. W. L. Schmitt. Diagnosis.—Moderate sized, lower front margin of head deeply excavate, head 2! body segments long, with 2 pairs of corneal lenses. Antenna I extending beyond end of antenna II peduncular segment 5; pedun- cular segments ratio 100:150:70, first seg- ment tumid, antenna I flagellum with 13 segments, setae moderate in length. Anten- na II peduncular segments 4 and 5 length ratio 100:60, flagellum 74 body length, with 20 segments, setae moderate in length. Mandible heavily sclerotized, 11 rakers, left with 4 teeth on lacinia mobilis, 6 teeth on incisor, palp segment 2 slightly curved, heavily setose, palp segment 3 %4 length of segment 2 with 5 scattered and 3 apical se- tae. Maxilla 2, upper lip, and lower lip with- out diagnostic features. Maxilliped inner plate with 2 setal spines and 2 chisel-shaped spines distally and submarginal row of plu- mose setae; outer plate with 13 chisel-shaped spines and 3 distal plumose setae; palp nor- mal. Maxilla | inner plate with 2 apical se- tae; outer plate with 11 distal spines; palp with 6 distal spines and 11 distal facial setal spines. Coxa 1 with well developed pos- teroventral notch, coxa 2 with slit, coxae 3 and 4 without slit or notch. All gills saclike. Pereopod | heavily setose with small dacty]. Pereopod 2 heavily setose, with elongate carpus. Pereopods 3 and 4 very similar but with 4 slightly more massive. Merus of pe- reopod 3 with setae of posterior margin on distal 2 only; pereopod 4 with margin en- tirely setose; dactyli of pereopods 3 and 4 longer than combined length of propodus and carpus on respective leg. Pereopod 5, anterior margin of basis rounded, posterior margin weakly biolobate; carpus with 4 an- terior spines, posterior submarginal spi- nules in clusters of 2, 3, and 4, cluster of posterodistal marginal spines; propodus with 2 posterior and 3 distal spines, dactyl with 6 accessory teeth. Pereopod 6, anterior mar- gin of basis bare proximally, 8 distal spines, posterior margin bare; anterior margin of carpus with 6 spines and with anterodistal and posterodistal clusters of spines, 4 sets of submarginal spinules in groups of 1, 2, 4, and 5, propodus with 10 antero- and 3 posteromarginal spines, dactyl with 9 ac- cessory teeth. Pereopod 7, basis inflated, posterodistal margin with small spines, ante- rior margin slightly concave, ischium short with anterodistal spine, merus with postero- distal setae, antero- and posterodistal mar- gins acuminate, carpus anterodistally acu- minate with spine, posterodistally with 3 spines, propodus inflated, dactyl acuminate VOLUME 100, NUMBER 1 5 > Fig. 1. Ampelisca macrodonta, new species: A, Head; B, Left mandible; C, Telson; D, Pereopod 4; E, Pereopod 3; F, Uropod 1; G, Uropod 2; H, Uropod 3; I, Maxilliped. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. Ampelisca macrodonta, new species: A, Pereopod 5 and detail of dactyl; B, Pereopod 7; C, Pereopod 6 and detail of dactyl; D, Lateral view of pleosome and urosome; E, Detail of slit or notch of coxa 2; F, Coxa 1. VOLUME 100, NUMBER 1 with apex curved anteriad. Epimeron 1 rounded, ventral margin slightly sinuous with plumose setae; epimeron 2 rounded with ventral plumose setae; epimeron 3 with rounded anterior margin, very strong tooth at posteroventral corner, posterior margin slightly sinuous above tooth. Uropod 1 equal to uropod 2 in length, peduncle spinose, lower margin with facial spinules, outer ra- mus without dorsal spines but with ventro- lateral spinules, inner ramus spinose. Uro- pod 2 peduncle and rami distally heavily spinose, outer ramus with long distal spine. Uropod 3 rami acuminate with plumose marginal setae. Telson cleft for more than ¥, length with few dorsal setal spines. Male. —Unknown. Variation.—Little variability is evident in the pair of specimens collected from the Falkland Islands. Differences noted in the setation and number of flagellar articles of the antennae are minor and due to differ- ences in maturity. Etymology.—The specific name macro- donta, derived from the Greek ‘“‘makros’”’ (large) and “‘odous”’ (stem odont-) (tooth), refers to the very large tooth of the 3rd epi- meral plate. Remarks. —Ampelisca macrodonta is de- scribed from an area where seven species of Ampelisca have been recognized. Four of these seven species known from the extreme southern reaches of South America and Antarctica possess characteristics of the seventh leg similar to A. macrodonta. Am- pelisca barnardi Nicholls, 1938, A. hemi- cryptops K. H. Barnard, 1930, A. richard- soni Karaman, 1975, and A. statensis K. H. Barnard, 1932, differ from A. macrodonta by the notch in the anterior margin of seg- ment 4 of pereopod 7 (absent in A. macro- donta) and the small size of the posterior process of epimeral plate 3 (very large in A. macrodonta). Ampelisca macrodonta may be separated from A. bouvieri Chevreux, 1913, A. composita Schellenberg, 1931, and A. anversensis Karaman, 1975, by the deep- ly concave lower margin of the head and broader outer ramus of uropod 3 of A. mac- rodonta. An interesting aspect of A. macrodonta is the presence of small spines on the postero- distal margin of the basis of pereopod 7. This author is aware of only two additional species which possess this feature. This pair of undescribed species has been found in the western Atlantic and the Gulf of Mexico and differs from A. macrodonta by a stron- ger superior lobe and much weaker pos- teroventral tooth on the third epimeral plate (Goeke, unpublished data). Acknowledgments The author would like to express his grat- itude to the staff of the National Museum of Natural History for their kindness and help during a recent visit to Washington. Thanks are also extended to Richard W. Heard, Jr., Gulf Coast Research Labora- tory, for his support and for a critical read- ing of the manuscript. Literature Cited Barnard, K. H. 1930. Crustacea, Part 2 Amphipo- da.—British Antarctic (“Terra Nova’’) Expedi- tion, 1910, Natural History Report, Zoology 8(4): 307-454. . 1932. Amphipoda.— Discovery Reports 5:1- 326. Chevreux, E. 1913. Amphipodes.—Deuxiéme Ex- pédition Antarctique Francaise (1908-19 10):79- 186. Karaman, G.S. 1975. Contribution to the knowledge of the Amphipoda. 68. Descriptions of two new species of the genus Ampelisca (family Ampe- liscidae), along with a redescription of A. bou- vieri Chevreux, 1913.—Beaufortia 24:37-54. Nicholls, E. L. 1938. Amphipoda Gammaridea.— Australasian Antarctic Expedition 1911-1914, Scientific Reports, series C, Zoology, and Bot- any 2(4):1-145. Schellenberg, A. 1931. Gammariden und Caprelliden des Magellangebietes, Sudgeorgiens und der Westantarktis.—Further Zoological Results of the Swedish Antarctic Expedition 1901-1903. 2(6): 1-290. U.S. Army Corps of Engineers, LMNPD- RE, P.O. Box 60267, New Orleans, Loui- siana 70160. PROC. BIOL. SOC. WASH. 100(1), 1987, pp. 8-12 NEOTROPICAL MONOGENEA. 10. OMOTHECIUM, NEW GENUS (DACTYLOGYRIDAE: ANCYROCEPHALINAE) AND TWO NEW SPECIES FROM THE PIRANAMBU, PINIRAMPUS PIRINAMPU (SPIX), (SILURIFORMES), IN BRAZIL D. C. Kritsky, V. E. Thatcher, and W. A. Boeger Abstract.—Omothecium, new genus (Dactylogyridae: Ancyrocephalinae) is proposed for two new species (O. pinirampi [type] and O. luckyi) collected from the gills of the piranambu, Pinirampus pirinampu (Spix), from Janauaca Lake near Manaus, Amazonas, Brazil. The new genus is characterized by its species having a sinistral nonsclerotized vagina opening anteriorly near the level of the copulatory complex, tandem gonads (testis postovarian), unmod- ified anchors and bars, undilated hook shanks, and a clockwise coiled cirral tube arising acutely from the cirral base. Among monogeneans collected from Bra- zil during the past decade, two species of Ancyrocephalinae were discovered on the gills of the piranambu, Pinirampus piri- nampu (Spix), Siluriformes, and were orig- inally considered members of Urocleidoides sensu Mizelle, Kritsky, and Crane (1968). The revision of Urocleidoides by Kritsky, Thatcher, and Boeger (1986) has precluded these from the restricted genus. However, common morphologic features of the two species now justify the proposal of the new genus Omothecium. The host was collected from Janauaca Lake near Manaus, Amazonas, Brazil, on 25 April 1984. Methods of host and parasite collection, preparation of helminths for study, measurement, and numbering of haptoral hook pairs are as described by Kritsky, Thatcher, and Boeger (1986). Mea- surements are in micrometers; averages are followed by ranges in parentheses. Mea- surements of the cirrus include 1) the di- ameter of the proximal ring of the coil, de- picted on the respective drawings as the interval between the solid straight lines, and 2) an approximation of total length of the cirrus obtained by using a Minerva curvi- meter on camera lucida drawings. Type specimens are deposited in the collections of the Instituto Nacional de Pesquisas da Amazonia (INPA), the U.S. National Mu- seum Helminthological Collection (USNM), and the University of Nebraska State Mu- seum (HWM.L) as indicated below. Omothecium, new genus Diagnosis. —Dactylogyridae, Ancyroce- phalinae. Body divisible into cephalic re- gion, trunk, peduncle, and haptor. Tegu- ment thin, smooth. Head organs, cephalic lobes present; cephalic glands unicellular, comprising 2 bilateral groups posterolateral to pharynx. Eyes present. Mouth subter- minal, midventral; pharynx muscular, glan- dular; esophagus present; intestinal caeca 2, confluent posterior to testis, lacking diver- ticula. Gonads tandem, intercaecal; testis postovarian. Vas deferens looping left in- testinal crus; seminal vesicle a dilation of vas deferens; prostatic reservoir not ob- served. Cirrus comprising an ovate base from which coiled tube originates at acute angle, rings clockwise (Kritsky, Boeger, and Thatcher 1985). Accessory piece not artic- ulated to cirrus, a fleshy rod serving as cirrus guide. Common genital pore midventral, at level of intestinal bifurcation. Oviduct short; VOLUME 100, NUMBER 1 uterus delicate; seminal receptacle lying near anterior end of ovary; vagina weakly scler- otized, sinistral, opening anteriorly near level of copulatory complex. Vitellaria well developed, coextensive with gut. Haptor armed with dorsal and ventral pairs of un- modified anchors, ventral and dorsal bars, 7 pairs of hooks with ancyrocephaline dis- tribution (Mizelle 1936); hooks with undi- lated shanks. Parasites of gills of siluriform fishes. Type species, host, and locality.—Omo- thecium pinirampi, n. sp. from Pinirampus pirinampu (Spix), Pimelodidae, Janauaca Lake near Manaus, Amazonas, Brazil. Other species. —Omothecium luckyi, n. sp. from Pinirampus pirinampu (Spix), Pime- lodidae, Janauaca Lake, near Manaus, Amazonas, Brazil. Etymology.— The generic name is from Greek (Omos = shoulder + théké = case) and refers to the anterior position of the vaginal opening. Remarks. — Omothecium is characterized by species possessing |) a sinistral nonscler- otized vagina opening anteriorly near the level of the copulatory complex, 2) tandem gonads (testis postovarian), 3) unmodified anchors and bars, 4) undilated hook shanks, and 5) a clockwise cirrus tube arising from an acute angle from the cirrus base. Omo- thecium resembles Cosmetocleithrum Krit- sky, Thatcher, and Boeger, 1986, by having species with tandem testes, unmodified an- chors, and a sinistral vagina. The new genus differs from Cosmetocleithrum by lacking submedian posterior projections of the dor- sal bar (present in Cosmetocleithrum), by having the vagina opening anteriorly on the left side (sinistral at level of ootype in Cos- metocleithrum), and by the nature of the copulatory complex. Omothecium pinirampi, new species Figs. 1-9 Type specimens.— Holotype, INPA PA282-1; paratypes, USNM 78798, HWML 22972, Description (based on 5 specimens).— Body fusiform, elongate with large glandu- lar area posterior to limit of vitellaria; ce- phalic margin comprising 2 terminal, 2 bi- lateral cephalic lobes; 4 head organs, well developed; each group of cephalic glands comprising relatively few cells. Usually 2 eyes, submedial at anterior pharyngeal mar- gin, frequently closely appressed giving ap- pearance of single eye; eye granules small, subspherical; accessory granules distributed in cephalic and anterior trunk regions. Phar- ynx subspherical; esophagus short. Pedun- cle moderately elongate; haptor hexagonal. Anchors similar, each with poorly devel- oped roots, large base, straight shaft, slightly curved point. Ventral bar shaped as ex- panded V; dorsal bar usually W-shaped, fre- quently fiattened. Hook distribution ancy- rocephaline, except members of hook pair 1 are situated submarginally on each side of the haptor. Hooks comprising 3 mor- phoiogic types; hook pairs 1, 3, 4, 7 with delicate point and shaft, slightly depressed thumb, elongate straight shank; pair 2 sim- ilar to previous pairs except shank length significantly reduced, erect thumb; pair 5, 6 with rapidly tapered shaft, strongly de- pressed thumb; FH loop shank length (pair 2), “% shank length (pairs 5, 6), 7% shank length (remaining pairs). Cirrus comprising enlarged fusiform base with proximal flare, tube of about one complete ring. Accessory piece a cirrus guide, spathulate. Gonads ba- cilliform. Seminal vesicle a coiled dilation of vas deferens. Measurements. —Body 952 (924-1004) long; greatest width 155 (129-189) near midlength. Greatest pharynx diameter 74 (68-78). Haptor 90 (73-112) long, 154 (122- 185) wide. Ventral anchor 44 (41-47), base width 24 (20-27); dorsal anchor 41 (37-45), base width 26 (22-30). Ventral bar 44 (41- 47); dorsal bar 36-37. Hook pair 2-9 (7- 11), all others 16 (12-20). Cirrus 155-156 long, ring diameter 44 (41-47); accessory piece 36 (30-44). Testis 191 (143-240) x 54 (41-67); ovary 68 (53-82) x 37 (31-43). Remarks. —Omothecium piniramp1 1s the OW) = = ~ ~~ N S ioe) (—— ry 8 9 Figs. 1-9. Omothecium pinirampi: 1, Ventral view of holotype; 2, Hook pair 2; 3, Hook (pairs 5, 6); 4, Hook (pairs 1, 3, 4, 7); 5, Copulatory complex; 6, Ventral bar; 7, Dorsal bar; 8, Ventral anchor; 9, Dorsal anchor. All drawings are at the same scale (30 micrometers) except Fig. 1 (200 micrometers). 10 VOLUME 100, NUMBER 1 11 Ay Wh Gy) A RN 11 13 30 15 16 Figs. 10-16. Omothecium luckyi: 10, Holotype (ventral view); 11, Hook; 12, Copulatory complex; 13, Ventral bar; 14, Dorsal bar; 15, Ventral anchor; 16, Dorsal anchor. All figures are drawn to the 30-micrometer scale except Fig. 10 (100 micrometers). 12 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON type species for the genus and is named for its host. Omothecium luckyi, new species Figs. 10-16 Type specimens.— Holotype, INPA PA283-1; paratypes, USNM 78795, HWML 22 TBs Description (based on 3 specimens).— Body fusiform; cephalic margin comprising 2 terminal, 2 bilateral poorly-developed ce- phalic lobes; head organs 4, poorly devel- oped; cephalic glands inconspicuous. Eyes 2, closely associated, frequently appearing as single submedian eye at level of anterior pharyngeal margin; eye granules small, ir- regular to subspherical; accessory granules absent. Pharynx spherical; esophagus short. Peduncle broad, elongate; haptor subhexag- onal. Anchors similar, each with poorly de- veloped roots, large base, short straight shaft, slightly curved point. Bars similar, V-shaped. Hooks similar, each with fine point and shaft, erect thumb, delicate shank; FH loop “,_ shank length. Cirral base ovate with tissue flaps at each end, tube compris- ing a coil of less than one ring; accessory piece simple, serving as cirrus guide. Go- nads subovate. Measurements.— Body 375 (334-397) long, greatest width 54 (51-55) near mid- length. Pharyngeal diameter 22 (19-27). Haptor 50 (49-52) long, 54 (50-58) wide. Ventral anchor 19-20, base width 15-16; dorsal anchor 22 (20-23), base width 14— 15. Ventral bar 29 (27-30); dorsal bar 24 (23-26). Hook (all pairs) 10-11. Cirrus 72- 73 long, ring diameter 20-21; accessory piece 22-23. Remarks.— This species differs from O. pinirampi by 1) possessing smaller sclero- tized haptoral structures, 2) the morphology of the dorsal bars (usually W-shaped in O. pinirampi; V-shaped in O. luckyi), and 3) the absence of different types of hooks in O. luckyi. Omothecium luckyi is named in honor of Dr. Z. Lucky, Faculty of Science, Brno, Czechoslovakia, who has greatly sup- ported our studies on Neotropical Mono- genea by providing specimens of his species collected from aquarium fishes in Czecho- slovakia. Literature Cited Kritsky, D. C., Boeger, W. A., and V. E. Thatcher. 1985. Neotropical Monogenea. 7. Parasites of the pirarucu, Arapaima gigas (Cuvier) with de- scriptions of two new species and redescriptions of Dawestrema cycloancistrium Price and Now- lin, 1967 (Dactylogyridae: Ancyrocephali- nae).— Proceedings of the Biological Society of Washington 98:321-331. , Thatcher, V. E., and W. A. Boeger. 1986. Neotropical Monogenea. 8. Revision of Uro- cleidoides (Dactylogyridae, Ancyrocephalinae). — Proceedings of the Helminthological Society of Washington 53:1-37. Mizelle, J.D. 1936. New species of trematodes from the gills of Illinois fishes.—American Midland Naturalist 17:785-806. —, Kritsky, D.C.,and J. W. Crane. 1968. Studies on monogenetic trematodes. XXXVIII. Ancy- rocephalinae from South America with the pro- posal of Jainus gen. n.—American Midland Naturalist 80:186-198. (DCK) Department of Allied Health Professions and Idaho Museum of Natural History, Idaho State University, Box 8002, Pocatello, Idaho 83209; (VET) Instituto Nacional de Pesquisas da Amazonia, Ma- naus, Brazil; (WAB) INPA and Department of Biological Sciences, Idaho State Univer- sity, Box 8007, Pocatello, Idaho 83209. PROC. BIOL. SOC. WASH. 100(1), 1987, pp. 13-20 HOST AND ELEVATIONAL SPECIFICITY OF PARASITIC BEETLES (AMBLYOPINUS SOLSKY) (COLEOPTERA: STAPHYLINIDAE) IN PANAMA Robert M. Timm and James S. Ashe Abstract.— The literature on staphylinid beetles of the tribe Amblyopinini, all of which are parasitic on Neotropical or Australian mammals, has provided few specifics on the natural history and host relationships of these beetles. We provide the first comprehensive data on host relationships and elevational range for a community of amblyopinines and their potential hosts. Mammals col- lected from eight separate localities along an elevational transect ranging from 900 to 1856 m were examined for parasitic beetles of the genus Amblyopinus. At these localities, mammals collected include a diverse array of marsupials, edentates, insectivores, bats, rodents, and lagomorphs. Among these potential hosts, parasitic beetles were limited to a subset of the cricetine rodents. Am- blyopinines were collected from only two species of rodents at two localities with elevations of 1425-1525 m and 1800-1825 m. Amblyopinus emarginatus Seevers was collected at both localities, where it was found on a single species of host, Oryzomys albigularis. In contrast, A. tiptoni Barrera was collected only at the higher elevation on Reithrodontomys creper. These data, as well as re- analysis of literature records, are consistent with the interpretation that these beetles are much more host restricted than has been previously recognized. Additionally, while these beetles appear to be tracking specific species of hosts, they are restricted to that subset of the hosts’ range above 1000 m. This implies that factors other than host range are of considerable importance to the dis- tribution and ecology of these beetles. Staphylinid beetles of the tribe Ambly- oOpinini have most often been collected from the fur of mammals. They are thought to be parasitic, although there is little infor- mation available on the natural history of this tribe. There are five genera in the tribe Ambly- opinini, four of which are restricted to the Neotropical region, Amblyopinus Solsky, Amblyopinodes Seevers, Edrabius Fauvel, and Megamblyopinus Seevers, and a single genus, Myotyphlus Fauvel, which occurs in Australia and Tasmania (Seevers 1955). Members of the Amblyopinini are primar- ily found at high elevations and are seldom reported on mammals collected below 1000 meters elevation. All four Neotropical gen- era are found in South America; members of only one genus, Amblyopinus, are found in Central America and Mexico. Thirty-four species of Amblyopinus currently are rec- ognized, most being known by only a few specimens with little biological data avail- able. Five species of Amblyopinus are known from Central America and Mexico (A. bo- livari Seevers and A. isabellae Barrera from Mexico, A. schmidti Seevers from Guate- mala, A. tiptoni Barrera from Costa Rica and Panama, and A. emarginatus Seevers from Panama, Colombia, and Venezuela). The most common hosts are cricetine ro- dents, although a wide range of hosts has been reported and includes marsupials; ca- viomorph, myomorph, and sciuromorph 14 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON rodents; a sloth; and bats. Of the Panama- nian amblyopinines, A. tiptoni has been col- lected most commonly on Peromyscus nu- dipes (Barrera 1966a, b), though it has also been reported on Reithrodontomys creper (Barrera 1966a, Vaughan 1982). In contrast, A. emarginatus has been reported from a wide array of hosts, though most commonly from species of Oryzomys (Barrera and Ma- chado-Allison 1968, Machado-Allison and Barrera 1972). The purpose of this paper is to report new data available for two species of Amblyopi- nus in Panama, A. emarginatus and A. tip- toni. We are now able to provide detailed information on host and elevational speci- ficity of these species. Additionally, we ex- amined specimens previously reported from Panama and provide a reevaluation of this material. Materials and Methods From 15 May through 10 July 1980, Ron- ald H. Pine and Robert J. Izor collected small mammals and their ectoparasites in the Cerro Colorado region of western Pan- ama. Mammals were collected at eight lo- calities ranging from 900 to 1856 m (see Pine and Handley, in prep.). Localities where Amblyopinus was encountered include (Fig. 1): Panama, Chiriqui-Bocas del Toro boundary, Cerro Bollo, 3.5 km E of Esco- peta, 1800-1856 m (subsequently referred to as the “Cerro Bollo”’ locality). Originally cloud forest and elfin woodland, consider- able forest remains, though some clearing has been undertaken for surveying purpos- es. Traps were set along pre-existing paths and paths cut through the forest for the trap- line. Species of small mammals taken at Cerro Bollo (numbers collected in parentheses) in- clude: Talamancan small-eared shrew, Cryptotis gracilis (1); blackish small-eared shrew, Cryptotis nigrescens (15); Cryptotis sp. (1); Tomes’ rice rat, Oryzomys albigularis (14); cloud forest pygmy rice rat, Oryzomys vegetus (3); Chiriqui harvest mouse, Rei- throdontomys creper (33); Mexican harvest mouse, Reithrodontomys mexicanus (2); and Chiriqui brown mouse, Scotinomys xe- rampelinus (9) (Pine and Handley, in prep.). Bocas del Toro, 25 km NNE of San Felix, 1425-1525 m (subsequently referred to as the “San Felix” locality). This area is cloud forest, except where disturbed. Trapping took place along a recently cut trail and a small stream, the Quebrada Alicia. Species of small mammals taken at this locality (numbers collected in parentheses) include: opossum, Didelphis marsupialis (1); South American mouse-opossum, Mar- mosa robinsoni (2); blackish small-eared shrew, Cryptotis nigrescens (5); Tomes’ rice rat, Oryzomys albigularis (50); cloud forest pygmy rice rat, Oryzomys vegetus (2); na- ked-footed deer mouse, Peromyscus nu- dipes (15); and Mexican harvest mouse, Reithrodontomys mexicanus (9) (Pine and Handley, in prep.). Other species of terrestrial small mam- mals collected at lower elevations (900, 1275-1325, and 1400-1425 m) include: water opossum, Chironectes minimus; two- toed sloth, Choloepus hoffmanni; forest rab- bit, Sylvilagus brasiliensis; Harris’ rice rat, Oryzomys aphrastus; pygmy rice rat, Ory- zomys fulvescens; yellow deer mouse, Pero- myscus flavidus; and hispid cotton rat, Sig- modon hispidus. Additionally, 19 species of bats were collected throughout the eleva- tional range (Pine and Handley, in prep.). None of these other mammals were para- sitized by Amblyopinus, nor were any Am- blyopinus found below 1425 m elevation. An attempt was made to collect every beetle encountered, although a small num- ber escaped. The actual number that es- caped is uncertain, but represents only a very small percentage of those captured. None of the beetles which escaped were from hosts or elevations which differ from those reported here (R. Izor and R. Pine, pers. comm.). Thus, these data provide an infor- mative representation of beetle distribu- VOLUME 100, NUMBER 1 tions among available hosts and elevational range at these localities. The mammals collected are deposited at the U.S. National Museum of Natural His- tory in Washington; beetles at the Field Mu- seum of Natural History. Results Staphylinids of the genus Amblyopinus were collected at two localities in Panama, the San Felix locality and the Cerro Bollo locality. Two species of Amblyopinus were collected. Amblyopinus emarginatus was obtained at both localities, although it was common at only 1425-1525 m. Amblyopi- nus tiptoni was obtained at only 1800-1856 m. These two localities are separated by a distance of less than 3 km. Amblyopinus emarginatus Seevers Amblyopinus emarginatus was taken from a single host species, Tomes’ rice rat Ory- zomys albigularis, from two elevational ranges (1425-1525 m and 1800-1856 m). It was collected from ten hosts; nine from the lower elevational range (total of 29 bee- tles) and one from the higher (1 beetle). In the lower elevational range (1425-1525 m) 50 O. albigularis were trapped, 35 males and 15 females. Of these, 9 had beetles (Ta- ble 1), 8 males and 1 female. At the higher elevation (1800-1856 m) 14 O. albigularis were trapped, 10 males and 4 females. Only 1 male had specimens of A. emarginatus (Table 1). At the San Felix locality, beetles were taken between 22 May and 11 June 1980; at the Cerro Bollo locality, the single A. emarginatus was taken on 7 July 1980. Specimens of A. emarginatus from these localities were compared with the holotype of A. emarginatus from Colombia in the collection of the Field Museum of Natural History and found to be well within the ex- pected range of variation in structural fea- tures for this species. The specimens of A. emarginatus from western Panama are 1so- lated from the other populations of this 1425 - 1525 m 1800 - 1856 m = De) wo s [o) (oe) (o} (oe) (panies oe 2) o Stel; ge 8 se) 7) 2) 3 Q2 Lo ao Qe = es o £5 a” co a = os So See (OS 1S W SS, N® N®& ES oO Bo Ss SS ~® o53 O65 Oo Xx 5 62 6> GS « ee S 8 © Q = == {S Yo o o x< eg og A. emarginatus QV 4. tiptoni Fig. 1. Numbers of individuals of mammal species captured (total bar) and corresponding numbers from which specimens of Amblyopinus emarginatus Seevers and Amblyopinus tiptoni Barrera were collected from two localities (Cerro Bollo, 3.5 km E of Escopeta, 1800- 1856 m; and Bocas del Toro, 25 km NNE San Felix, 1425-1525 m) along an elevational transect in Pana- ma. We follow Pine (pers. comm.) in recognizing Ory- zomys vegetus as distinct from O. fulvescens. species in the highlands of Columbia and Venezuela by about 900 km of the Pana- manian and northwestern Colombian low- lands. Amblyopinus tiptoni Barrera Amblyopinus tiptoni was found on only one species of host, the Chiriqui harvest mouse, Reithrodontomys creper, at only one 16 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 1.—Numbers of specimens of Amblyopinus emarginatus collected at two localities in western Pan- ama [Cerro Bollo, 3.5 km E of Escopeta (=“‘Cerro Bol- lo” locality); Bocas del Toro, 25 km NNE of San Felix (=**San Felix” locality)] from individual specimens of Oryzomys albigularis. No. beetles Host Catalog Fe- No.* Locality Males males Total 541126 San Felix — 2 2 541127 San Felix 5 5 10 541131 San Felix 1 DY, 3 541135 San Felix l 1 yD) 541136 San Felix — 1 1 541141 San Felix l y; 3 541142 San Felix 3 1 4 541354 San Felix D — y 541373 San Felix — D, 2 541356 Cerro Bollo _ al, ei Totals 13 17 30 * Host Catalog No. refers to the catalog number of mammal specimens housed at the Smithsonian Insti- tution. elevation range, 1800-1856 m. This was the only locality trapped at this high an eleva- tion, and the only locality where R. creper was obtained. Amblyopinus emarginatus was also taken here, but is represented by only a single individual. Sixteen males and 16 females of R. creper were trapped, in- cluding all age categories. A total of seven specimens of A. tiptoni were found on 5 of these (Table 2), 2 maies and 3 females. Am- blyopinus tiptoni was collected between 15 and 22 June 1980. Discussion a) Host specificity.—In this study 38 species of small mammals were collected and surveyed for parasitic arthropods over an eight week period in western Panama. This included 19 terrestrial species and 19 species of bats. Of these potential hosts, Amblyopinus emarginatus was found only on Oryzomys albigularis and was present only at the two localities (and elevations) where O. albigularis was trapped. Ambly- opinus tiptoni was found only on a single Table 2.—Numbers of specimens of Amblyopinus tiptoni collected at one locality in western Panama [Cerro Bollo, 3.5 km E of Escopeta (=“‘Cerro Bollo”’ locality)] from individual specimens of Reithrodonto- mys creper. No. beetles Host Catalog Fe- No.* Locality Males males Total 541206 Cerro Bollo 1 y 3 541212 Cerro Bollo 1 — 1 541220 Cerro Bollo — 1 1 541225 Cerro Bollo 1 — 1 541227 Cerro Bollo ale = lh Totals 4 3 i * Host Catalog No. refers to the catalog numbers of mammal specimens housed at the Smithsonian Insti- tution. species of host, Reithrodontomys creper, at the single locality and elevational range where R. creper was trapped. Several thousand specimens of mammals were collected from throughout Panama as part of the zoonoses surveys by the Gorgas Memorial Laboratory and the “Ectopara- sites of Panama” surveys during the 1950’s and 1960’s. Most of the 201 species of na- tive terrestrial mammals listed by Handley (1966) from Panama have been examined for ectoparasites. During these surveys, only three specimens of Amblyopinus emargin- atus were obtained, all from a single locality at 1525 min Bocas del Toro Province. Two species of hosts were reported, Oryzomys albigularis and Peromyscus nudipes. Elsewhere, A. emarginatus has been col- lected on a number of occasions from the highlands of Colombia and Venezuela. Throughout the range of A. emarginatus, it has been reported from a variety of mam- mals including: Oryzomys albigularis, O. caliginosus, O. alfaroi, O. concolor, O. de- vius, Akodon urichi, Peromyscus flavidus, Rhipidomys venustus, Thomasomys fusca- tus, T. laniger, and T. sp.; as well as from Didelphis marsupialis, Marmosa dryas, Bradypus infuscatus, Myotis nigricans, and Vampyrops oratus (Seevers 1955; Macha- VOLUME 100, NUMBER 1 Oryzomys albigularis Oryzomys caliginosus Other Oryzomys pb (3 species) | All other rodents (6 species) 2 Non-rodents (5 species) 3 Number of Records Fig. 2. Numbers of host individuals from which Amblyopinus emarginatus Seevers has been recorded, compiled from literature citations (Seevers 1955; Machado-Allison and Barrera 1964, 1972; Barrera 1966a, b; Barrera and Machado-Allison 1968). ! O. alfaroi, O. concolor, O. devius (O. devius is often considered conspecific with O. albigularis). 2 Akodon urichi, Peromyscus flavidus, Rhipidomys venustus, Thomasomys fuscatus, T. laniger, and Thoma- somys sp. 3 Didelphis marsupialis, Marmosa dryas, Bradypus infuscatus, Myotis nigricans, and Vampyrops oratus (these records must be considered suspect). do-Allison and Barrera 1964, 1972; Barrera 1966a; Barrera and Machado-Allison 1968). The last five of these, which include all re- ports from marsupials, the sloth, and bats, represent single records and should be con- sidered suspect. The records from the two bats were even suspected to be contami- nations in the original report by Machado- Allison and Barrera (1972). Among litera- ture records of A. emarginatus (n = 135) (Fig. 2), most reports have been from species of Oryzomys (124; 91.9%), and the large majority of these have been either from O. albigularis (n = 86; 69.4%) or O. caliginosus (n = 35; 28.2%). All other host records rep- resent single reports. This strongly suggests that species of Oryzomys represent the pri- mary hosts of A. emarginatus. Most records of A. emarginatus from Oryzomys are from O. albigularis, and in- deed it has been reported from this host throughout the known range of the beetle. However, at one locality in Colombia, A. emarginatus was commonly reported from O. caliginosus (Barrera and Machado-Alli- son 1968). Interestingly, O. albigularis was also abundant at this locality and frequently parasitized by A. emarginatus. These data, as well as results reported in this paper, provide considerable evidence that A. emarginatus is host specific on species of Oryzomys, primarily O. albigu- laris. In this study, it was found to be re- stricted to O. albigularis, though a variety of other rodent species, which could have served as hosts, were present in sympatry with O. albigularis. Thus, we conclude that A. emarginatus is much more host specific than literature reports suggest. In contrast to the large number of records for A. emarginatus, A. tiptoni has been re- ported only three times in the literature, once from Panama and twice from a single lo- cality in Costa Rica. In the original species description of specimens from Chiriqui Province in Panama, Barrera (1966a:284) reported “‘long series” of A. tiptoni from Peromyscus nudipes from several localities. However, he also noted “‘unique material” from Reithrodontomys creper. The other re- ports of A. tiptoni in the literature are from Cerro de la Muerte in Costa Rica where five 18 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON specimens were reported from Peromyscus nudipes by Barrera (1966b), and several specimens were recorded from a single in- dividual of R. creper by Vaughan (1982). In addition, we have collected A. tiptoni from several specimens of Peromyscus nudipes at Monteverde (Puntarenas Prov.) in Costa Rica (unpublished data). The results reported here suggest that A. tiptoni is limited in host range to R. creper at the localities sampled, though relatively few specimens were collected. It was col- lected at only one locality and elevation range (1800-1856 m). No specimens of P. nudipes were found at this locality; how- ever, P. nudipes was abundant at the locality where A. emarginatus was most common. Though this is the host of A. tiptoni most often reported in the literature, and the el- evation is within the range known for this beetle, no beetles were found on this host. b) Elevational zonation and host distri- bution. —In this study Oryzomys albigularis was taken at elevations above 1425 m, Reithrodontomys creper above 1800 m, and Peromyscus nudipes from 1250 to 1500 m. In a survey of mammals from Panama, Handley (1966) reported that Oryzomys al- bigularis was an uncommon, terrestrial species found in fog forest at high elevations in both extreme eastern and western Pan- ama. He found it at elevations ranging from 1220 to 2380 m. He reported Reithrodon- tomys creper to be abundant at high ele- vations in western Panama; it is primarily a terrestrial species that is found in fog for- ests and openings. He found it at elevations ranging from 2075 to 3175 m. Peromyscus nudipes was a common terrestrial species in evergreen forest. It was found at elevations ranging from 730 to 2380 m. Oryzomys albigularis and O. caliginosus are widespread species in southern Central America and northern South America and are found over a range of elevations. How- ever, O. albigularis is most common at higher elevations (above 1200 m), while O. caliginosus 18s most common at lower ele- vations (below 1000 m) to sea level (Gard- ner 1983, Handley 1966). Amblyopinus emarginatus has been collected on these hosts only at elevations above 1050 m. Herein, we found A. emarginatus common only at 1425-1525 m elevation, and rare at 1800-1856 m elevation. Several collections were made at localities ranging from 900 to 1425 m elevation. Neither O. albigularis nor A. emarginatus were found at these lower elevations, though two other species of Ory- zomys were present. It is interesting that A. emarginatus has not been found on O. caliginosus in Costa Rica or Panama where O. caliginosus is common at lower elevations. The only re- port of A. emarginatus from O. caliginosus is from a single locality in Colombia where O. albigularis and O. caliginosus were sym- patric at higher elevations (above 1600 m) (Barrera and Machado-Allison 1968). The occurrence of A. emarginatus only on O. albigularis in Panama (and perhaps else- where) may primarily be a result of the higher elevation preference of this species of Oryzomys. This implies that A. emargin- atus is restricted to higher elevations, though suitable hosts occur over a much broader elevational range. Handley (1966) reported that Peromyscus nudipes was common in Panama between 730 and 2380 m. However, A. tiptoni was collected from this species only between 1525 and 2075 m (Barrera 1966a). Barrera (1966a) reported that A. tiptoni was com- mon on P. nudipes from numerous locali- ties within the 1525 to 2075 m elevational range during the months of January and February. In addition, A. tiptoni has been collected at 3335 m from P. nudipes at Cer- ro de la Muerte in Costa Rica (Barrera 1966b) as well as at 1600 m at Monteverde (Timm, unpublished data). In comparison, Reithrodontomys creper is limited to higher elevations (2075-3350 m) in the mountain systems of northwestern Panama and central Costa Rica (Hooper 1952, Handley 1966, Hall 1981). Literature reports of A. tiptoni from R. creper have been from elevations of 3200 m in Panama VOLUME 100, NUMBER 1 (Barrera 1966a) and 3200 m in Costa Rica (Vaughan 1982). Our data indicate that A. tiptoni occurs on R. creper in Panama as low as 1800-1856 m, the lowest elevation at which this species of mammal was col- lected. It is notable that 15 specimens of P. nu- dipes were collected at 25 km NNE of San Felix (1425-1525 m). This is within the el- evational range from which A. ftiptoni has been previously reported to occur on P. nu- dipes. However, no specimens of Ambly- opinus were found on P. nudipes at this lo- cality. Conclusions Historically, the literature on staphylinid beetles of the tribe Amblyopinini, all of which are parasitic on Neotropical mam- mals, has provided few specifics on the nat- ural history and host relationships of these beetles. Our studies provide the first com- prehensive data on host relationships and elevational range for a community of these beetles and potential hosts. In Central America the mammal community includes a diverse array of marsupials, edentates, in- sectivores, bats, primates, rodents, and car- nivores. Among these potential hosts, para- sitic beetles are limited to a subset of the cricetine rodents. Within a given commu- nity of cricetines, beetles appear to be re- stricted to one or a very few species. Spe- cifically, our data are consistent with the interpretation that these beetles are much more host restricted than has been previ- ously recognized. In the communities studied, Amblyopi- nus emarginatus is host specific on Oryzo- mys albigularis. In addition, our reanalysis of distribution of A. emarginatus among hosts from throughout its known range pro- vides considerable evidence of host speci- ficity for mammals of the genus Oryzomys in general and O. albigularis in particular. The number of host records for A. emar- ginatus far exceeds those for any other am- blyopinine. Therefore, though published host data for many species are confusing 19 and inconclusive (see for examples, Seevers 1955, Machado-Allison and Barrera 1972) our analysis of data for A. emarginatus pro- vides a robust confirmation of a previously unexpected level of host specificity. Data presented herein indicate that Am- blyopinus tiptoni is restricted to Reithro- dontomys creper. However, host specificity for this beetle throughout its range is less well documented. This uncertainty is par- tially a result of lack of comprehensive col- lections. Additionally, it has been reported from two species of cricetine rodents, Pero- myscus nudipes and R. creper. In most in- stances, it has been restricted to one host locally. Considerable additional study is re- quired to clarify host relationships of this beetle. Additionally, our data show that while beetles of the genus 4mb/yopinus appear to be tracking specific hosts, they are restricted to that subset of the hosts’ range above 1000 m. Few other groups of parasites show this restricted distributional pattern in relation to host elevational range. Wenzel and Tip- ton (1966) also noted that Amblyopinus tip- toni, as well as several other species of ec- toparasites, are restricted to only a subset of their hosts’ range, and suggested that these groups were primarily temperate in distri- bution. This implies that factors other than host range are of considerable importance to the distribution and ecology of some ec- toparasites. Though factors responsible for this aspect of amblyopinine distributions are not obvious, possible candidates include temperature-humidity relationships, an- nual and seasonal climatic variation, and vegetational communities. This character- istic of amblyopinine ecology seems to be one of the most curious features of these beetles and merits further attention. Acknowledgments We wish to acknowledge gratefully a grant made by the Rice Foundation to the Field Museum of Natural History which provid- ed support for our research on this project. We also thank R. J. Izor and R. H. Pine for 20 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON conscientiously collecting the beetles and generously supplying the data to us. The original field work was supported in part by the Field Museum of Natural History and the U. S. National Museum of Natural His- tory. In Panama, J. Burt; Empresa de Cobre Cerro Colorado, S.A.; RTZ Development Interprises, Ltd.; and Texasgulf Panama Inc. provided numerous courtesies and field support. M. D. Carleton and C. O. Handley, Jr. provided identifications of specimens deposited at the Smithsonian Institution. We thank R. J. Izor, A. F. Newton, Jr., R. H. Pine, M. K. Thayer, and R. L. Wenzel for reading, and making helpful comments on the manuscript. Literature Cited Barrera, A. 1966a. New species of the genus Ambly- opinus Solsky from Panama and Mexico (Co- leoptera: Staphylinidae). Pp. 281-288 in R. L. Wenzel and V. J. Tipton, eds., Ectoparasites of Panama. Field Museum of Natural History, Chicago. . 1966b. Hallazgo de Amblyopinus tiptoni Bar- rera, 1966 en Costa Rica, A. C. (Col.: Staph.).— Acta Zoologica Mexicana 8(5):1-3. ,and C. Machado-Allison. 1968. Amblyopinus de Colombia (Coleoptera Staphilinidae). — Neo- tropica 14(45):89-98. Gardner, A. L. 1983. Oryzomys caliginosus (raton pardo, raton arrocero pardo, Costa Rican dusky rice rat). Pp. 483-485 in D. H. Janzen, ed., Costa Rican natural history. University of Chicago Press, Chicago. Hall, E.R. 1981. The mammals of North America. Volume 2. John Wiley & Sons, New York. 1181 pp. Handley, C. O., Jr. 1966. Checklist of the mammals of Panama. Pp. 753-795 in R. L. Wenzel and V.J. Tipton, eds., Ectoparasites of Panama. Field Museum of Natural History, Chicago. Hooper, E. T. 1952. A systematic review of the har- vest mice (genus Reithrodontomys) of Latin America.— Miscellaneous Publications, Mu- seum of Zoology, University of Michigan 77:1- DSc Machado-Allison, C. E., and A. Barrera. 1964. Sobre Megamblyopinus, Amblyopinus, y Amblyopi- nodes (Col., Staph.).—Revista de la Sociedad Mexicana de Historia Natural 25:173-187. , and . 1972. Venezuelan Amblyopinini (Insecta: Coleoptera; Staphylinidae).— Brigham Young University Science Bulletin, Biological Series 17(2):1-14. Pine, R. H., and C. O. Handley, Jr. In Prep. Mammals of the Cerro Colorado area, Western Panama. Seevers, C. H. 1955. A revision of the tribe Ambly- opinini: staphylinid beetles parasitic on mam- mals.—Fieldiana: Zoology 37:211-264. Vaughan, C. 1982. Parasitism of harvest mice by staphylinid beetles.—Brenesia 19/20:615. Wenzel, R. L., and V. J. Tipton. 1966. Some rela- tionships between mammal hosts and their ec- toparasites. Pp. 677-723 in R. L. Wenzel and V.J. Tipton, eds., Ectoparasites of Panama. Field Museum of Natural History, Chicago. Department of Zoology, Field Museum of Natural History, Chicago, Illinois 60605; (RMT) Present address: Museum of Natural History, University of Kansas, Lawrence, Kansas 66045. PROC. BIOL. SOC. WASH. 100(1), 1987, pp. 21-27 THE DISTRIBUTION AND FOOD HABITS OF NEPHATYS BUCERA EHLERS, 1868, (POLYCHAETA: NEPHTYIDAE) IN THE SURF ZONE OF A SANDY BEACH John J. McDermott Abstract. — Nephtys bucera was more abundant below mean low water (MLW) than above, in the surf zone of an exposed sandy beach along the southern coast of New Jersey. Food items found in the digestive tracts of 111 N. bucera, collected over several years, were identified and counted. One hundred pre- served worms ranged in length from 51 to 166 mm, wet weight from 70 to 3400 mg, and dry weight from 13 to 506 mg. Juvenile wedge clams, Donax variabilis, were the most common and abundant prey, occurring in 65.5% of worms with food and accounting for 81.3% of all food items. Clams, all young- of-the-year, were consumed whole and digested in their valves; they ranged in shell length from 1.1 to 6.4 mm (xX 2.47 + 0.82). The spionid polychaete, Scolelepis squamata, was next in dietary importance (incidence 32.3%, abun- dance 10.1%). This potentially important prey species dominates the intertidal area above MLW for all except three winter months, and thus there is usually little spatial overlap with Nephtys. The crustaceans Amphiporeia virginiana and Emerita talpoida appeared to be incidental prey. Thus N. bucera is an opportunistic carnivore (there was no evidence of deposit feeding) influencing the population dynamics of several surf-dwelling invertebrates. Nephtys bucera itself is a minor prey item for at least one species of juvenile fish inhabiting the surf zone, viz., the northern kingfish Menticirrhus saxatilis. Nephtys bucera, a relatively large sand- dwelling polychaete, is distributed from the Gulf of Saint Lawrence to South Carolina, and in the Gulf of Mexico from Florida to Mississippi (Verrill 1873; Hartman 1945, 1951; Carpenter 1956; Sanders 1958; Pet- tibone 1963; Croker 1970, 1977; Wass 1972; Day 1973'; Gardiner 1975; Kinner and Maurer 1978; Zingmark 1978; Croker and Hatfield 1980; Garlo 1980). It is found from the intertidal zone to a depth of about 200 ! Perkins (1980) examined all of the specimens from the coast of North Carolina identified as N. bucera by Day (1973), and found that 17 specimens were N. bu- cera, 43 N. simoni (a new species) and 13 may have been the young of N. picta. m (Pettibone 1963, Day 1973, Zingmark 1978), primarily in sand containing little organic matter, and has been reported in concentrations ranging from 10 to 419 worms/m~2 (Carpenter 1956, Sanders 1958, Kinner and Maurer 1978). Clark (1962) reviewed the scanty litera- ture on the genus and pointed out that, with few exceptions, all species were carnivo- rous. In apparently the only published in- formation on the food habits of N. bucera, Croker (1977) listed it as an omnivore, but gave no evidence for such a designation. The purpose of the present study was to determine the types of prey consumed by N. bucera, to relate this information to po- tential prey in its habitat, and to discuss its role in the food web of the surf zone. 22 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ee N= 100 Y >= (ae (=) = 15 LL ro) ac Lu co 55 = 0 50 100 150 BENG GASSES ss MM: Fig. 1. The length-frequency distribution of 10 mm size classes of Nephtys bucera (preserved) from the surf zone at Seven Mile Beach, Avalon, New Jersey. Materials and Methods Worms were collected in the surf of an exposed sandy beach at Avalon, New Jersey (39°04'43’N, 74°44'05”W) from 1977-1985. This was the site of an extended study (1977- 1979) dealing with interactions of the ben- thos and nekton (McDermott 1983). The spatial distribution of Nephtys was deter- mined from the analyses of 704 benthic cores (each 20 cm deep and 46 cm?) taken along transects run perpendicular to the shore from above mean high water (MHW) to 50 m seaward of mean low water (MLW). Few of the worms collected in these cores were used for food analyses because most were dam- aged. The majority of worms used for food de- terminations were collected during spring ebb tides by turning over the exposed sed- iments with a long-handled spade. Digging was usually done in a 20 m wide region (parallel to the shore) below MLW, where worms were more abundant than inshore of this mark. Worms were placed immediately into capped tubes containing ~15% sea water formalin. This isolation was to insure recovery of any food regurgitated from the gut, but no evidence of regurgitation was subsequently found. 25 N= 100 Y = © ons LU oO ~ LiJ aa) = 5 = 0 200 400 600 DRYW Gl eEASSESs MG Fig. 2. The dry weight-frequency distribution of 50 mg size classes of Nephtys bucera (preserved). The following determinations were made on each worm prior to examination: total length to the nearest mm, width of the pro- stomium (posterior to base of antennae) to the nearest 0.1 mm, and damp weight to the nearest 10 mg. Dry weight and ash-free dry weight (AFDW) to the nearest mg, were de- termined after examination. Contents of the entire digestive tract of each worm were examined with the aid of a dissecting microscope, and prey species were identified and counted. Shell lengths of one prey item, Donax variabilis, were measured to the nearest 0.1 mm with a cal- ibrated ocular micrometer. Results Eleven of the 111 N. bucera used for food analyses were slightly damaged and were not measured or weighed. The remaining worms ranged in length from 51 to 166 mm (x 110.9 mm =+ 22.7) (Fig. 1), in wet weight from 70 to 3400 mg (x 1167.1 mg + 696.3), and in dry weight from 13 to 506 mg (x 187.2 mg + 108.1) (Fig. 2). The mean AFDW/dry weight ratio for 49 worms col- lected in the fall of 1980 and 1983 was 0.832 + 0.051. Regression equations and correlation coefficients for various relation- ships follow: VOLUME 100, NUMBER 1 ee STATIONS WJ >) => >) 50 30 10 23 MLW : 3 = O >) —/ 30 10 a) fe eME ES Fig. 3. The distribution of Nephtys bucera at Seven Mile Beach, Avalon, New Jersey, in relation to the mean low water mark (MLW), based on 704 cores (each 20 cm deep, 46 cm?) taken from 1977-1979 at 250 stations above and 102 stations below MLW, plotted in 10 m intervals. The zone occupied by the dominant polychaete Scolelepis squamata is located approximately between 10 and 30 m above MLW, during all but three winter months. dry wt (Y) vs. wet wt Y = 9.3500 + (0.1525)X dry wt (Y) vs. length log Y = —3.8882 + (2.9832)X R=0.92 wet wt (Y) vs. length log Y = —3.6389 + (3.2469)X R=0.93 AFDW (Y) vs. dry wt R = 0.98 Y = 0.3726 + (0.8295)X R = 0.99 pro. w (Y) vs. length* Y = 0.1747 + (0.0127)X R = 0.91 * prostomium width of 90 worms (72 from those examined for food and 18 ju- venile worms from the benthic cores). Core samples revealed that small juvenile worms occurred in the surf zone during the summer months (June to the middle of Sep- tember), when the water temperatures ex- ceeded 15°C. These juveniles (15 of 23 were measurable) ranged in length from 5.5 to 42.0 mm (X = 19.0 mm = 11.3; 10 worms were below 20 mm), and in dry weight from 0.1 to 15.7 mg (X 2.3 mg + 4.3). They are not included in Figs. 1 and 2, and their gut contents were not analyzed. Six N. bucera were collected in 500 cores taken above MLW, and 30 were recovered from 204 cores below MLW (Fig. 3). The ratios of worms to cores were 0.012 and 0.147, respectively, approximating 2.5 worms/m~? inshore and 32 worms/m7? off- shore of MLW. Thus UN. bucera is consid- erably more abundant below the MLW mark, only slightly overlapping the narrow (20 m wide band) Scolelepis zone, which is found in the mid-intertidal region for most of the year. During the winter months (par- ticularly January through March), however, when the Scolelepis population moves to 24 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 1.—Incidence and total numbers of food items in the digestive tracts of Nephtys bucera collected in the surf zone of Seven Mile Beach, Avalon, New Jersey. Number of worms Incidence of food items 4 i Unidentified Date of Amphi- Amphipod crustacean collection Examined With food Scolelepis Donax poreia fragments Emerita fragments 13-5-78 ] 0 24-7-78 ] 1 l 14-10-78 3 3 2 l 1 28-10-78 5) 4 1 2 2 7-7-79 3 3 3 16-10-79 3 l 1 27-3-80 7 2 D, 12-10-80 7 7 i 4 22-11-80 19 16 8 13 2 1 9-9-83 5 4 y) yD) 5-11-83 24 18 13 4 l 3-6-85 33 31 30 8 1 Totals 111 90 29 59 14 4 4 2 Numbers of each item 34* 278 18 4* 5) 2® * Where no anterior ends were found, fragments were calculated as one individual. the lower intertidal and subtidal region (McDermott 1983), there may be more overlap of the populations. This assumes that Nephtys does not migrate also. Donax variabilis was the most common and most abundant prey species found in the digestive tracts of 111 worms examined from 1978 to 1985 (Table 1). These clams occurred in 65.6% of worms with food, amounted to 81.3% of all food items, and were ingested by worms from all size classes. The small spionid polychaete Scolelepis squamata was next in importance, while the two crustaceans, Amphiporeia virginiana and Emerita talpoida, appeared to be in- cidental prey. The mole crabs belonged to the 1980 year-class (carapace lengths: 3.5, 4.0, 4.5 and 5.1 mm). Most of the amphipod fragments were probably from Amphipo- relia. Eleven additional damaged worms were also examined, and in these Donax was dominant over Scolelepis and Emerita. Sand was not often found in the digestive tracts of Nephtys, but when found it was usually in small amounts in the rectum. Clams are consumed whole, i.e., the shells are not crushed by the two conical pharyn- geal jaws of the worm during ingestion. Bro- ken shells were found sporadically, but this was attributed to damage caused during dis- section. The clams were all young-of-the- year, ranging in shell length from 1.1 to 6.4 mm (xX = 2.47 mm + 0.82). The length frequency distribution of all undamaged clams recovered from Nephtys appears in Fig. 4. Clams were usually oriented with their long axes parallel to the gut of the worm, thus the longest clam (6.4 mm) re- quired a buccal opening equal only to its height (4 mm). This clam was one of two recovered from one of the longest worms in the collection (149 mm long, 441 mg dry wt). Up to 18 clams were found in a single worm (a specimen 118 mm long, 243 mg dry wt), and they ranged in length from 1.9 to 3.2 mm (X = 2.38 + 0.33). Nine of these clams were packed into a swollen part of the gut just posterior to the muscular phar- ynx. Most of the clams recovered were in the 1.5 to 2.9 mm category (Fig. 4), and prac- tically all of the larger clams were from the 22-11-80 collection (2.8 to 6.4 mm long, x 4.41 mm + 0.85, n = 30). Worms in this VOLUME 100, NUMBER 1 collection were not significantly larger than those from other collections, indicating that worms collected at other times would have been capable of ingesting larger clams had they been available. Clams are digested within their shells, the two valves usually remaining attached by the ligament even after complete digestion of the tissues. Some tightly closed clams with little or no digestion were found in the lower gut. Whether some clams escape digestion and are redeposited alive back into the sed- iments with the feces, is a question that must be left to laboratory experimentation. K. Fauchald’s (Smithsonian Institution, pers. comm.) observations on several species of nephtyids, suggest that following digestion bivalve shells of the size reported here are probably regurgitated because the anal openings of these worms are too small to allow passage in the normal manner. Discussion Nephtys bucera appears to be an oppor- tunistic carnivore, feeding on the inverte- brates most available to them in the surf. It was found previously that Scolelepis, Do- nax, Amphiporeia and Emerita are the dominant benthic species in this particular habitat along the coast of New Jersey (McDermott 1983). The virtual lack of significant amounts of sand in the digestive tracts of N. bucera, with or without macrofauna, certainly ob- viates any contention that this species is a deposit feeder. Sanders’ (1956, 1960) ob- servation that N. incisa is a non-selective detritus feeder was viewed by Clark (1962) as a condition probably peculiar to the species, and perhaps related to the unusu- ally high concentrations of the worm in the offshore waters of Long Island Sound and Buzzards Bay, Massachusetts. Sanders’ (1960) contention that it would not be fea- sible for the worm to be primarily a pre- daceous feeder because of its dominance in the soft-bottom community of Buzzards Bay, might be reevaluated in light of what 5) 100 N=291 (eo) (=>) RO jo) NUMBER OF CLAMS 0 1 2 3 4 5 6 i LENGTH CLASSES MM. Fig. 4. The length-frequency of 0.5 mm size classes of Donax variabilis removed from the digestive tracts of Nephtys bucera. appears to be a great diversity and relative abundance of potential prey species tabu- lated for the area. Suspension feeding was recently suggested for the same species by Davis (1979). Clark (1962) noted that the European species, N. cirrosa and N. hombergi, while usually not packed with food, fed mainly on a variety of polychaetes. Never were their digestive tracts filled with sand. Warwick and Price (1975) also concluded that N. hombergi was a carnivore, but their evi- dence was meager, and they were concerned that no other suitable macrofaunal animals of a lower trophic level were available in the Lynher Estuary (England) to maintain the large population of Nephtys. Ockelmann and Muus (1978) determined that Nephtys spp. from Danish waters fed on foraminif- erans, small molluscs (including the small montacutid bivalve Mysella bidentata) and smaller polychaetes. Nephtys caeca, N. hombergi and N. longosetosa all occurred in their study areas, but the food of each was not specified. Brown (1964) considered N. capensis to be an impartial feeder that ‘draws the line only at plant material,” but he gave no substantiating data. Srinivasa Rao and Rama Sarma (1978) concluded that N. oligobranchia from the east coast of In- dia feeds primarily on polychaetes. In their 26 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON recent review of food and feeding in poly- chaetes, Fauchald and Jumars (1979) con- cluded that nephtyids are predominantly carnivores, but they admitted that there is a scarcity of information on the subject. Ockelmann and Muus (1978) observed that the shells of Mysella bidentata are crushed by the pharyngeal jaws of European Nephtys spp. This did not appear to be the case with Donax ingested by N. bucera. Possible differences in feeding behavior and size of the jaws among nephtyids and the physical properties of bivalve shells, may be related to how small bivalves are handled by these predators. Although Donax was the dominant food item of N. bucera, further more intensive seasonal observations from field collec- tions, as well as laboratory experiments, are necessary in order to determine if it has a preference for this clam. It may be suggested that as the population of Scolelepis moves offshore in the winter it becomes a more important food source in the diet of N. buce- ra, but the sporadic nature of my sampling does not lend itself to a seasonal analysis. It appears likely that other species of haus- torlid amphipods, belonging to the same subfamily as Amphiporeia (Pontoporeiinae) and to the subfamily Haustoriinae, and known to exist in relatively small numbers in the Nephtys region (McDermott 1983), would be suitable prey. Nephtys’ role in the food web of the shal- low surf zone is that of a predator, influ- encing (to an undetermined extent) the pop- ulation dynamics of Donax, Scolelepis and various benthic crustaceans. All of the latter have been shown to be important prey for a variety of fishes that inhabit the surf zone (McDermott 1983). Nephtys, in turn, serves as occasional food for juvenile northern kingfish (Menticirrhus saxatilis); single specimens were found in the stomachs of four of the 159 fish examined. The kingfish feeds predominantly on Scolelepis, which makes up over 80% of its diet in the surf (McDermott 1983). Nephtys was not ob- served in the stomach contents of 236 ju- venile spot (Leiostomus xanthurus), also a member of the Sciaenidae, or in other species of fishes examined from the same environ- ment. Further, more detailed studies on the life history of N. bucera will require improved sampling techniques. The methods used in the present study are not adequate for ob- taining sufficient numbers of worms. Two devices, a suction corer and a scoop dredge, specifically developed recently for use in the surf zone by Fleischack et al. (1985) may hold promise for obtaining adequate sam- ples of Nephtys and other invertebrates liv- ing in this turbulent region. Acknowledgments I am indebted to the following for their help in collecting worms: J. M. McDermott, J. L. Sabol, and K. L. Trautmann. I appre- ciate the many kindnesses rendered to me by the staff of the Wetlands Institute of Le- high University. I thank J. L. Richardson for reviewing the manuscript. Literature Cited Brown, A. C. 1964. Food relationships on the inter- tidal sandy beaches of the Cape Peninsula. — South African Journal of Science 60:35—41. Carpenter, D. G. 1956. Distribution of polychaete annelids in the Alligator Harbor area, Franklin County, Florida.— Florida State University Studies 22:89-110. Clark, R. B. 1962. Observations on the food of Neph- tys.— Limnology and Oceanography 7:380-385. Croker, R.A., 1970. Intertidal sand macrofauna from Long Island, New York.—Chesapeake Science 11:134-137. 1977. Macro-infauna of northern New En- gland marine sand: long-term intertidal com- munity structure. Pp. 439-450 in B. C. Coull, ed., Ecology of Marine Benthos. University of South Carolina Press, Columbia. , and E. B. Hatfield. 1980. Space partitioning and interactions in an intertidal and sand-bur- rowing amphipod guild.— Marine Biology 61: 79-88. Davis, W. R. 1979. The burrowing, feeding and res- piratory activity of Nephtys incisa Malmgren, 1865 (Polychaeta: Annelida). Ph.D. Disserta- VOLUME 100, NUMBER 1 tion, University of South Carolina, Columbia, 140 pp. Day, J.H. 1973. New Polychaeta from Beaufort, with a key to all species recorded from North Car- olina.— NOAA Technical Report NMFS Circ. 375, 140 pp. Fauchald, K., and P. A. Jumars. 1979. The diet of worms: a study of polychaete feeding guilds. Pp. 193-284 in M. Barnes, ed., Annual Review of Oceanography and Marine Biology. Vol. 17. Aberdeen University Press, Scotland. Fleischack, P. C., A. J. de Freitas, and R. B. Jackson. 1985. Two apparatuses for sampling benthic fauna in surf zones. — Estuarine Coastal and Shelf Science 21:287-293. Gardiner, S. L. 1975. Errant polychaete annelids from North Carolina. — Journal of the Elisha Mitchell Scientific Society 91:77—220. Garlo, E. V. 1980. Abundance and distribution of benthic macro-invertebrates near Little Egg In- let, New Jersey, from 1972 to 1974.—Interna- tional Revue der Gesamten Hydrobiologie 65: 345-356. Hartman, O. 1945. The marine annelids of North Carolina.— Duke University Marine Station Bulletin 2:1-51. . 1951. The littoral marine annelids of the Gulf of Mexico. — Publications of the Institute of Ma- rine Science, University of Texas 2:7-124. Kinner, P., and D. Maurer. 1978. Polychaetous an- nelids of the Delaware Bay region.— U.S. Fish- ery Bulletin 76:209-224. McDermott, J. J. 1983 Food web in the surf zone of an exposed sandy beach along the mid-Atlantic coast of the United States. Pp. 529-538 in A. McLachlan and T. Erasmus, eds., Sandy Beach- es as Ecosystems. Dr. W. Junk, The Hague, Netherlands. Ockelmann, K. W., and K. Muus. 1978. The biology, ecology and behaviour of the bivalve Mysella bidentata (Montagu).— Ophelia 17:1—93. Perkins, T. H. 1980. Review of species previously referred to Ceratonereis mirabilis, and descrip- tions of new species of Ceratonereis, Nephtys, A and Goniada (Polychaeta).— Proceedings of the Biological Society of Washington 93:1-49. Pettibone, M. H. 1963. Marine polychaete worms of the New England region. 1. Aphroditidae through Trochochaetidae. — United States National Mu- seum Bulletin 227:1-356. Sanders, H. L. 1956. Oceanography of Long Island Sound, 1952-1954. X. Biology of marine bot- tom communities.—Bulletin of the Bingham Oceanographic Collection Yale University 15: 345-414. 1958. Benthic studies in Buzzards Bay. I. Animal-sediment relationships. —Limnology and Oceanography 3:245-258. 1960. Benthic studies in Buzzards Bay. III. The structure of the soft-bottom community. — Limnology and Oceanography 5:138-153. Srinivasa Rao, D.,andD. V.RamaSarma. 1978. Food and feeding habits of Nephtys oligobranchia Southern (Annelida: Polychaeta). — Indian Jour- nal of Marine Science 7:193-195. , Verrill, A. E. 1873. Report upon the invertebrate animals of Vineyard Sound and the adjacent waters, with an account of the physical char- acters of the region.—U.S. Fish Commission Report 1871-1872, pp. 295-852, 40 pls. Warwick, R. M., and R. Price. 1975. Macrofauna production in an estuarine mud-flat.— Journal of the Marine Biological Association of the United Kingdom 55:1-18. Wass, M. L. (Editor). 1972. A check list of the biota of lower Chesapeake Bay.—Special Scientific Report, Virginia Institute of Marine Science, Gloucester Point 65:1—290. Zingmark, R. G. (Editor). 1978. An annotated check- list of the biota of the coastal zone of South Carolina.— University of South Carolina Press, Columbia, 364 pp. Department of Biology, Franklin and Marshall College, Lancaster, Pennsylvania 17604. PROC. BIOL. SOC. WASH. 100(1), 1987, pp. 28-34 TWO NEW SPECIES OF PETTIBONEIA (POLYCHAETA: DORVILLEIDAE) PRIMARILY FROM THE GULF OF MEXICO Paul S. Wolf Abstract.—Two new species of the genus Pettiboneia Orensanz, 1973, are described, P. duofurca from the east and west coasts of Florida, Alabama, and Texas, and P. blakei from the east coast of Florida. The genus Pettiboneia Orensanz, 1973, is currently known for three species: P. san- matiensis Orensanz, 1973 (redescribed by Blake 1979), P. urciensis Campoy and San Martin, 1980, and P. australiensis West- heide and von Nordheim, 1985. Pettiboneia sanmatiensis is recorded from Argentina, its type locality, and from California and British Columbia (Blake 1979:1137); P. ur- ciensis is described from the Mediterranean; and P. australiensis is described from Aus- tralia. Wolf (1984) identified two poten- tially new species of Pettiboneia from the Gulf of Mexico; these are described below. The bulk of the material examined for this study was collected as part of a U.S. Bureau of Land Management (now Min- erals Management Service) Outer Conti- nental Shelf baseline study conducted dur- ing 1975-1981. MAFLA stations were those designated within the Mississippi—Ala- bama-—Florida portion of the program; SO- FLA stations were those located off south- west Florida; STOCS stations were located off the Texas coast (see Uebelacker and Johnson (1984). The remaining material was collected under the auspices of the Envi- ronmental Protection Agency (EPA) during contracts issued to Battelle, Columbus Lab- oratories (EPA/Bat stations); to Science Ap- plications International Corp. through JRB Associates, McLean, Virginia (SAI sta- tions); and under a contract issued by the U.S. Army Corps of Engineers to Barry A. Vittor & Associates, Inc. (COE station). The type material and some additional specimens are deposited in the National Museum of Natural History, Smithsonian Institution (USNM). Other specimens are in the laboratory museum of Barry A. Vittor & Associates, Inc., Mobile, Alabama. Figure Abbreviations an antenna nuO nuchal organ br branchia pa palp noto notopodium vC_ ventral cirrus Pettiboneia Orensanz, 1973 Type species.—Pettiboneia sanmatiensis Orensanz, 1973. Diagnosis. —Maxillae in 8—14 rows, each row composed of free denticles only; some maxillary rows with rasping denticles; base plates absent. Denticle rows not fused pos- teriorly. Maxillary carriers absent. Prosto- mium with well-developed, biarticulate palps; antennae digitiform, simple, shorter than palps. Notopodia present anteriorly, long, with internal acicula, without distal article; absent posteriorly. Branchiae pres- ent or absent. Supraacicular setae simple with long tapered forms and furcate setae; subacicular setae include compound falci- gers and occasionally inferior simple setae. Remarks.— Armstrong and Jumars (1978) described Protodorvillea pugettensis and P. dibranchiata, both of which probably be- long in Pettiboneia primarily because of their jaw morphology (Blake, pers. comm.); how- VOLUME 100, NUMBER 1 ever, both species are described as having maxillary carriers. Thus, their inclusion within Pettiboneia would necessitate expan- sion of the generic diagnosis to include species with maxillary carriers. Pettiboneia duofurca, new species Figs. 1, 2 Pettiboneia sp. A.—Wolf, 1984:44—47, fig. 44-1, 44-2a-j. Material examined.— FLORIDA, off Palm Beach: EPA/Bat Sta 5-2, Nov 1984, 26°46.0'N, 79°58.9'W, 118 m, medium coarse sand, 1 Paratype (USNM 98931).— Off Port Everglades: EPA/Bat Sta Dive 1, Nov 1984, 26°07.7'N, 80°05.0’W, 17 m, sand, 1 Paratype (USNM 98932).—Off Tampa Bay: MAFLA Sta 22091, Jun 1976, 27°52'30.5’N, 83°33'59.0"W, 34 m, clayey- sandy silt, 1 specimen (USNM 89597); SAI Sta A-13-1, 15 May 1982, 82°59.0'N, 27°36.5'W, 49 m, medium fine sand, | spec- imen; SAI Sta A-13-3, same date and lo- cation, 49 m, coarse sand with gravel, 1 specimen. — Northwest: MAFLA Sta 2422C, Jun 1976, 29°30’N, 84°27'W, 24 m, medi- um fine sand, 2 specimens including | ovig- erous female; MAFLA Sta 2424B, Jul 1976, 20°13'00.7”N, 85°00'01.4”W, 27 m, medi- um sand, 4 specimens (USNM 89557); MAFLA Sta 2424C, same date and loca- tion, 3 specimens; MAFLA Sta 2424E, same date and location, 1 specimen (USNM 89596); MAFLA Sta 2424H, same date and location, 1 specimen.—Off Panama City: MWAEFIEA Sta 2528H, Aug 1977, 29°54'58.6"N, 86°04'58.5”W, 37 m, coarse sand, 2 specimens (USNM 89594—5); SAI Sta 10-1, Nov 1983, 30°08'07’N, 85°45'39”W, 17.7 m, fine to medium sand with shell, 1 Paratype (USNM 98933).— ALABAMA, off Mobile Bay: COE Sta 695- 6, 1 Apr 1981, 30°01.5'N, 87°54.27'W, 22.4 m, sand, 1 specimen.—TEXAS, off Mata- gorda: SAI Sta 1-3, Nov 1983, 28°15.33'N, 96°11.91'W, 9.3 m, sand and gravel, 1 spec- imen.— Off Padre Island: STOCS Sta III/4- 2S) 1, no date, 26°58’N, 97°20'W, 15 m, sand, 1 specimen.—Off Port Isabel: STOCS Sta IV/4-1, Fall 1976, 26°10’N, 97°08'W, 15 m, sand, 2 specimens (USNM 89556, 89593). — Off Brownsville: SAI Sta 8-3, Nov 1983, 26°03.28'N, 97°04.15'W, 16.9 m, sand, Ho- lotype (USNM 98930). Description.—Length to 4.3 mm, width to 0.2 mm. Largest specimen incomplete with 45 setigers. Prostomium conical, broadly rounded anteriorly (Fig. la), with 1 pair of small eyes, when present. Antennae smooth, digitiform, about *% length of palps. Palps biarticulate. Single pair of large nu- chal organs present dorsolaterally at postec- tal corners of prostomium. Additional cil- lated areas present on prostomium, palps, and laterally on each tentacular ring (Fig. la). Notopodia present on setigers 2-8 or 9, with internal acicula, without distal article (Fig. 1b). Notopodia absent posteriorly (Fig. lc). Branchiae present on dorsal edge of neuropodia from setigers 3-5 or 8, or absent entirely. Parapodia without well-developed pre- or postsetal lobes (Fig. 1b, c). Supraacicular setae including simple ser- rate setae tapering to fine tips, and furcate setae. Furcate setae of anterior few setigers small, with tines slightly unequal in length, with spines present below short tine (Fig. 1d); thereafter tines long, slightly unequal in length, with or without spines below short tine (Fig. le, f). Furcate setae of juveniles pseudocompound, with tines unequal in length and with numerous spines below short tine (Fig. lg, h). Subacicular setae com- pound, with long to short blades having uni- dentate tips (Fig. 11, j); blade length ratio approximately 4.4—5.1:1. Far posterior parapodia with inferior simple setae (Fig. 1k). Pygidium with 4 anal cirri, 2 very long, filiform dorsal ones and pair of shorter, club- shaped ventral ones (Fig. 2a). Maxillae in 12 rows (Fig. 2b), each row composed of separate denticles, each den- ticle wider than long. Row 1 with 6-7 broad, 30 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. parapodium, posterior view; d, Furcate seta from anterior setiger of adult; e, Same, from posterior setiger; f, Same, spinous form; g, Pseudocompound furcate seta from anterior setiger of juvenile; h, Same, from posterior setiger of juvenile; i, Superior subacicular falciger; j, Inferior subacicular falciger; k, Inferior simple seta. (Figs. a-e, I-k from Wolf 1984:fig. 44-2a—h.) rounded, poorly chitinized, clear, rasp-like denticles, each denticle with numerous mi- nute teeth. Row II with about 13-15 squared, flattened denticles, each denticle with one large tooth and several smaller ones. All denticles of Row II heavily sclerotized and dark brown in color. Rows III-VI with poorly sclerotized, clear, rounded, rasp-like denticles, each denticle with numerous teeth. Mandibles each with anterior portions rounded, scalloped along inner edge. Each mandible strongly concave medially along outer edge, and with widely divergent pos- terior portions (Fig. 2c). Remarks.— Reexamination of material shows that rows I and II of Wolf (1984) were labelled and described in reverse order. It also appears that in most specimens ex- 0.03mm Pettiboneia duofurca: a, Anterior end, dorsal view; b, Anterior parapodium, posterior view; c, Posterior amined, row I is not as long as row II, but this feature is dificult to determine due to the small size and/or the condition of the worms. Indeed, row I may be analogous to maxillary carriers of other dorvilleid genera, thereby further distinguishing P. duofurca from other species of the genus. If maxillary carriers are present, this may be cause to remove P. duofurca to a new genus; how- ever, I have elected not to do this since other diagnostic characters indicate P. duofurca is best kept within Pettiboneia for the pres- ent. Among the material examined, one com- plete juvenile specimen was found that measures 2.06 mm in length for 28 setigers. It differed from the adult worms in having pseudocompound furcate setae (Fig. 1g, h) VOLUME 100, NUMBER 1 31 Fig. 2. Pettiboneia duofurca: a, Posterior end, dorsal view; b, Maxillae, dorsal view (left side not completely shown); c, Mandibles, dorsal view. (Fig. c from Wolf 1984:fig. 44-2j.) and in having dorsal cirri present on both parapodia of setiger 3 only. The mandibles and maxillae are identical to those of the adult although an entirely new set of each was developing within the juvenile speci- men. One gravid female was also found. It con- tained 1-2 eggs per setiger from setiger 11. The largest egg measured about 80 um in diameter. Pettiboneia duofurca is similar to P. blakei, described below, in having only 12 maxillary rows instead of 14 as in P. san- matiensis or eight rows as in P. australiensis and P. urciensis. Pettiboneia duofurca dif- fers from P. blakei in having denticles of all maxillary rows wider than long rather than longer than wide; in having smaller, more numerous rasp-like teeth on each denticle rather than fewer, more prominent teeth; in having narrow, widely flaring mandibles in- stead of broad, slightly divergent mandi- bles; in having two types of furcate setae instead of one; and in having branchiae in some specimens. Etymology.— The species name is taken 32 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON \ th d @ f g Fig. 3: Pettiboneia blakei: a, Anterior end, dorsal view; b, Anterior parapodium, posterior view; c, Posterior parapodium, posterior view; d, Furcate seta; e, Superior subacicular falciger; f, Inferior subacicular falciger; g, Setal shaft, edge-on view; h, Inferior simple seta; 1, Pygidium, dorsal view; j, Maxillae, dorsal view; k, Mandibles, dorsal view. (Figs. a—h, j, k, from Wolf 1984:fig. 44-4a-j.) from the Latin duo, two, and furca, fork, referring to the two types of furcate setae present along the body. Distribution.—East and west coasts of Florida to Texas, 9.3-118 m. Pettiboneia blakei, new species Fig. 3 Pettiboneia sp. B.— Wolf, 1984:44—49, fig. 44-3, 44-4a-j. Material examined.—FILORIDA, south- west: SOFLA Sta 20A, May 1981, 25°17.34'N, 82°09.73'W, 22 m, coarse sand, 1 specimen: MAFLA Sta 2211F, Nov 1977, 27°56'29.5"N, 83°52'59.5”W, 43 m, coarse sand, 2 specimens. — Off Tampa Bay: EPA/ Bat Sta 1111-III-OLD 3-10, Dec 1984, 27°29'N, 83°04.5’W, 22.6 m, sand, | Para- type (USNM 98937); EPA/Bat Sta 1111- IlI-5-1, Dec 1984, 27°28.7'N, 83°06.5’W, 24.7 m, sand, | Paratype (USNM 98935); EPA/Bat Sta 1111-III-5-3, same data, 1 Paratype (USNM 98936); EPA/Bat Sta VOLUME 100, NUMBER 1 1111-III-5-10, same data, Holotype (USNM 98934).— West coast: MAFLA Sta 2315A, Jul 1976, 28°33'59.1”N, 84°20'09.1’W, 38 m, silty-fine sand, 1 specimen; MAFLA Sta 2316C, Nov 1977, 28°42'00.3’N, 84°20'00.7” W, 35 m, silty-fine sand, 1 spec- imen.—Off Cape San Blas: MAFLA Sta 2854G, Aug 1977, 29°24'00.1’°N, 85°42'02.0”W, 42 m, medium fine sand, 1 specimen. — Off Panama City: MAFLA Sta Dose Aue 1977,- 29°54'58.6°N, 86°04'58.5”W, 37 m, coarse sand, | speci- men. Description.— Length to 8.5 mm, width to 0.58 mm. Largest specimen complete with 114 setigers. Prostomium (Fig. 3a) rounded anteriorly, expanded in ocular region, with single pair of large eyes at antennal bases. Antennae smooth, digitiform, about 2 length of palps. Palps distinctly biarticulate. Single pair of large, ciliated nuchal organs located at dorsal postectal corners of pro- stomium. Additional prominent ciliated areas present on prostomium, palps, and laterally on both peristomial rings (Fig. 3a). Notopodia present on setigers 2-12 to 24 (Fig. 3b), with internal acicula, without dis- tal article. Branchiae absent. Notopodia ab- sent posteriorly (Fig. 3c). Neuropodia with- out pre- and postsetal lobes. Supraacicular setae include long, simple, serrate setae tapering to fine tips; and furcate setae with blunt-tipped tines, slightly un- equal in length, about 4 rows of spines below short tine (Fig. 3d). Subacicular setae com- pound, unidentate, with long to short ser- rate blades (Fig. 3e, f); apical tips of setal shafts bifid when viewed edge-on (Fig. 3g). Inferior simple setae present on far poste- rior parapodia (Fig. 3h). Pygidium rounded with 2 pairs of fili- form, subterminal anal cirri, dorsal cirri about twice as long as ventral ones (Fig. 31). Maxillae arranged in 12 rows (Fig. 3)) each row composed entirely of free denticles. Each denticle longer than wide, with 1 main tooth and several smaller teeth. Maxillary carriers absent. Rows I-IV each with 6-8 denticles; 33 row V with 5-6 denticles; row VI with 2-4 denticles. Each mandible broad anteriorly, scalloped along inner edge, then tapering abruptly posteriorly and becoming slightly divergent (Fig. 3k). Remarks. —One paratype (USNM 98936) is a gravid female with numerous eggs pres- ent in each setiger from about setiger 26. The largest eggs measured 65-70 um in di- ameter. Pettiboneia blakei is most similar to P. duofurca, described above, but differs in several respects (see “REMARKS” for P. duofurca). Etymology.—The species is named in honor of Dr. James Blake, Battelle New En- gland Marine Research Laboratory, Dux- bury, Massachusetts, for his numerous con- tributions to polychaete taxonomy including those dealing with the Dorvilleidae. Distribution. — East coast of Florida, 22- 43 m. Acknowledgments I wish to thank Dr. Kristian Fauchald, Smithsonian Institution, Washington, D.C.; and Dr. Barry A. Vittor and Ms. Linda Sierke, Barry A. Vittor & Associates, Inc., for their reviews of the manuscript. Some of the material examined was ob- tained under the following contracts: En- vironmental Protection Agency (EPA) to Science Applications International Corp. through JRB Associates, McLean, Virginia, contract number 68-0106388; from EPA to Battelle, Columbus Laboratories, Colum- bus, Ohio, contract number 68-01-6986; and from the U.S. Army Corps of Engineers, Mobile District, to Barry A. Vittor & As- sociates, Inc., Mobile, Alabama, contract number DACWO1-80-C-0427. Barry A. Vittor & Associates, Inc., pro- vided monetary and material support for this study. Literature Cited Armstrong, J. W., and P. A. Jumars. 1978. Bran- chiate Dorvilleidae (Polychaeta) from the North 34 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Pacific.—Bulletin of the Southern California Academy of Science 77(3):133-138. Blake, J. A. 1979. A redescription of Pettiboneia san- matiensis Orensanz (Polychaeta: Dorvilleidae) and a revised key to the genera of the Dorvil- leidae.—Bulletin of the Southern California Academy of Science 78(2):136-140. Campoy, A., and G. San Martin. 1980. Pettiboneia uriensis sp. n.: Un nouveau Dorvilleidae (Poly- chétes: Errantes) de la Méditeranée. — Cahiers de Biologie Marine 21:201-207. Orensanz, J. M. 1973. Los anelidos poliquetos de la provincia biogeographica Argentina. III. Dor- villeidae.— Physis, Seccion A, 32(85):325-342. Uebelacker, J. M., and P. G. Johnson (Eds.). 1984. Taxonomic Guide to the polychaetes of the northern Gulf of Mexico. Final Report to the Minerals Management Service, contract 14-12- 001-29091. Barry A. Vittor & Associates, Inc., Mobile, Alabama. 7 vols. Westheide, W., and H. von Nordheim. 1985. In- sterstitial Dorvilleidae (Annelida, Polychaeta) from Europe, Australia and New Zealand.— Zoologica Scripta 14(3):183-199. Wolf, P. S. 1984. Chapter 44. Family Dorvilleidae. Pp. 44-1-44-37 in J. M. Uebelacker and P. G. Johnson, eds., Volume VI. Taxonomic Guide to the polychaetes of the Northern Gulf of Mex- ico. Final Report to the Minerals Management Service, contract 14-12-001-29091. Barry A. Vittor & Associates, Inc., Mobile, Alabama. Barry A. Vittor & Associates, Inc., 8100 Cottage Hill Rd., Mobile, Alabama 36609. PROC. BIOL. SOC. WASH. 100(1), 1987, pp. 35-39 DROMIOPSIS KIMBERLYAE, A NEW LATE CRETACEOUS CRAB FROM THE PIERRE SHALE OF SOUTH DAKOTA Gale A. Bishop Abstract.— Two specimens of an undescribed Cretaceous crab were collected from the Heart Tail Ranch in the lower Pierre Shale, Butte County, South Dakota. The specimens are assigned to Dromiopsis kimberlyae n. sp., the sec- ond North American species of Dromiopsis and the first from the Cretaceous of North America. Dromiopsis kimberlyae joins seven previously described decapod taxa as a rare faunal element in the Heart Tail Ranch Decapod As- semblage. Recollecting of the Heart Tail Ranch Decapod Assemblage, Butte County, South Dakota, on 23 June 1985, resulted in dis- covery of the left side of the carapace of an undescribed fossil crab (Fig. 2A, B) (by Kimberly Dawn Bishop). During subse- quent collecting on 31 July 1985, I found a second specimen of this crab, a complete carapace with a partly extended abdomen (Fig. 2C-I). Both specimens are preserved in apatite concretions typical of the Heart Tail Ranch Decapod Assemblage (Bishop 1985). This assemblage now consists of eight decapod species represented by about 900 specimens. The Heart Tail Ranch Decapod Assemblage is from the Gammon Ferrugi- nous Member of the Pierre Shale and is Ear- ly Campanian in age (see geologic map and stratigraphic diagrams in Bishop 1985, figs. 1 and 2). Systematic Paleontology Class Crustacea Order Decapoda Family Dynomenidae Ortmann, 1892 Dromiopsis Reuss, 1859 Diagnosis.—Carapace pentagonal, con- vex, front forming large triangular lobe, lat- eral margins tuberculate (to smooth), trans- verse grooves strong (emended after Glaessner 1969:R488). Range.— Dromiopsis is Late Cretaceous to Paleocene in age, ranging from the Cen- omanian through the Paleocene. Most taxa are from Belgium, Sweden, Denmark, and Germany (Forster 1975). One species, Dromiopsis americana Roberts, 1956, was described from the Paleocene of New Jer- sey. Dromiopsis kimberlyae, new species Figs. 1, 2 Diagnosis.— Carapace nearly circular, arched longitudinally and transversely; front downturned and triangular; anterolateral margin keeled, not tuberculate; transverse grooves strong, medial groove becoming obsolete posteriorly on intestinal region. Types.— The holotype (SDSM 10184) and paratype (SDSM 10185) are deposited in the collections of the Museum of Geology, South Dakota School of Mines, Rapid City, South Dakota 57701. Occurrence, preservation, and sample size. —Two specimens preserved in apatite concretions were collected in the SW ‘4, Sec. 16, TLIN, R2E, Butte County, South Da- kota from the Heart Tail Ranch Assemblage (Bishop 1985). This fauna is from the zone of Baculites sp. (smooth), early form and is mid-early Campanian in age. The holotype is acomplete carapace steinkern with a par- 36 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Line drawings of carapace of Dromiopsis kimberlyae showing carapace regions and grooves. Carapace grooves are: e, cervical; c, postcervical; a, brachialcardiac, and ae, position of attachment of the attractor epimeralis muscle. Carapace regions are: R, rostrum; O, orbit; Gm, mesogastric; Gp, protogastric (with epigastric boss near rostrum); U, urogastric; C, cardiac; I, intestinal; and B, the branchial regions, Be, epibranchial, Bm, mesobranchial, and Bt, metabran- chial. tially extended, but reflexed, abdomen. The paratype is the left third of a carapace steinkern. Dromiopsis kimberlyae is the eighth decapod species to be described from the Heart Tail Ranch Assemblage and the specimens represent specimens number 855 and 856 collected from the Heart Tail Ranch. Description.—Carapace pentagonal-cir- cular, slightly wider than long (19.1 mm long, 19.8 mm wide), strongly arched trans- versely, arched longitudinally. Rostrum strongly downturned, triangular with me- dial sulcus running onto tip causing up- turned rim to form a “bifid,” blunt tip. Or- bits large (59% of carapace width), divided, forming slight concavities on anterolateral margin, rims slightly raised, lower edge with suborbital spine. Anterolateral margins slightly concave around orbits then curving into a nearly circular arc, widest at crab’s midpoint, then forming posterolateral mar- gins until their junction with concave pos- terior margin. Anterolateral dorsal shield edge strongly reflexed and keeled from orbit to cervical furrow, from cervical furrow to branchiocardiac furrow, and for short dis- tance immediately behind branchiocardiac furrow, breaking up into a few parallel elon- gate tubercles. Posterior margin raised into ridge bordered by marginal furrow. Cara- pace strongly differentiated by 3 more or less transverse grooves. Cervical groove deep, relatively straight and oblique. Bran- chiocardiac groove almost as deep, nearly transverse from dorsal shield edge to medial ridge where it bifurcates to encircle cardiac region. Between cervical and branchiocar- diac furrows a shorter, third transverse groove, “post-cervical furrow,” arising just anterior of cardiac region and running sub- parallel to other 2 transverse grooves, be- coming obsolete before reaching dorsal shield edge. Cephalic arch (area anterior to cervical furrow) moderately differentiated into medial metagastric region, and proto- gastric-hepatic-epigastric region by indis- tinct groove running obliquely inward from cervical furrow then swinging forward to form triangular anterior tongue of mesogas- tric region. Posterior of mesogastric region separated into 2 low, lateral bosses by shal- low medial furrow; each boss asymmetrical, steep behind where roughened by mold of muscle insertion areas terminating in pair of small, spinelike insertion points. Epigas- tric regions raised into small, circular, epi- gastric bosses. Scapular arch (area posterior to cervical groove) differentiated into sag- ittal ridge and branchial regions by more or less continuous longitudinal groove. Sagit- tal ridge consisting of (anterior to posterior) urogastric region (“Gastrical gruben”’ and urogastric regions of Forster 1975), dia- mond-shaped cardiac region with trans- versely-paired tubercles, and poorly differ- entiated intestinal region. Branchial regions subdivided into anterior epibranchial, me- VOLUME 100, NUMBER 1 3h7/ Fig. 2. Photographs of Dromiopsis kimberlyae: A—B, Dorsal and left lateral view of paratype (SDSM 10184), x 2.0; C-I, holotype specimen (SDSM 10185) seen in: C, Dorsal view, x 1.0; D, Left oblique; E, Anterior; F, Right lateral; G, Ventral; H, Dorsal; and I, Posterior views (D-I, x 2.0). dial mesobranchial, and posterior meta- branchial regions. Major muscle insertion points lie on posteriors of mesogastric boss- es, on posterior edge of cervical furrow, and in small, paired oval areas in “‘postcervical furrow” (Fig. 2H). Both cervical and bran- chiocardiac furrow continue onto reflexed pterygostomial region separating it into an anterior part (subhepatic), medial part (an- terior subbranchial), and posterior part (posterior subbranchial). Abdomen subtile- ly grooved longitudinally, with low, later- ally placed pleural bosses paired across each somite. Appendages only preserved in cross section. Etymology.—Dromiopsis kimberlyae is named in honor of its discoverer, Kimberly Dawn Bishop. Comparison.—Dromiopsis kimberlyae can be distinguished from all congenors by its keeled, non-tuberculate anterolateral margin. Dromiopsis kimberlyae is much smoother, more circular, and has a carapace less divided by grooves than D. rugosa 38 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON (Schlotheim 1820) and its associates (D. cf. rugosa of Forster 1975). Dromiopsis kim- berlyae is not as coarsely ornamented as is D. gigas Forir, 1887, nor does it have as prominent a transverse, raised urogastric re- gion. Dromiopsis kimberlyae is more cir- cular in dorsal view, has a proportionally broader posterior margin, less rounded an- terolateral margins and much deeper cara- pace grooves than D. elegans Reuss, 1859. Dromiopsis kimberlyae is most similar to D. laevior Reuss, 1859, and the closely re- lated (Forster 1975:290) form D. depressa Segerberg, 1900, in circular carapace shape, carapace proportions, and carapace smoothness. Dromiopsis kimberlyae differs from D. depressa by being much more cir- cular, proportionally shorter, by having a much more highly grooved carapace, and by possessing a complete, incised cervical furrow. Dromiopsis kimberlyae is more cir- cular than D. /aevior, has deeper, more pro- nounced, more complete carapace grooves, retains a prominent, but incomplete, “‘post- cervical” furrow where D. /aevior does not, possesses tuberculate epigastric spines where D. laevior does not, and possesses trans- versely paired spinules on the cardiac region where D. /aevior is smooth. Dromiopsis kimberlyae differs significantly from D. americana Roberts, 1956, by being more circular, having its carapace more fully dif- ferentiated by generally deeper, almost complete grooves (except perhaps the bran- chiocardiac furrow so prominently depicted by Roberts 1956: fig. 2), by possessing the keeled, rather than tuberculate, anterolat- eral margin, and by possessing the “‘post- cervical” groove completely lacking in D. americana. Dromiopsis pulchella Secretan, 1964 (Pl. 19, fig. 7, text-figs. 99-100) bears little resemblance to congenors in Dromiop- sis and may represent a new genus-level tax- on. Remarks. — Dromiopsis kimberlyae is the second species of Dromiopsis described from North America and is the only Dromiopsis from the Cretaceous of North America. Its scarcity in the Heart Tail Ranch Assem- blage (~0.2% of the decapods) is in agree- ment with its European record (Forster 1975: 289) which consists of five species repre- sented by approximately 12 specimens, many of which are only claws associated with fragmentary carapaces. Dromiopsis kimberlyae generally fits into the phyletic scheme envisioned by Roberts (1956:8), forcing the divergence of the D. rugosa stock from the D. kimberlyae-D. elegans-D. lae- vior-D. americana stock further back in time to at least the mid-early Campanian. Acknowledgments My thanks are extended to Chance and Cindy Davis for the western hospitality they have extended to me over the last decade of collecting on the Heart Tail Ranch. The original specimen of this new species was collected by my daughter, Kimberly Dawn, whose assistance is gratefully acknowl- edged. Indirect support for this research has been provided by NSF Grant 8011570, Na- tional Geographic Society Grants 1629 and 2867-84, and Georgia Southern College through its Faculty Research Committee. The manuscript was typed by Judith Un- derwood. The paper was strengthened by critical reading by R. M. Feldmann, Rein- hard Forster, and an anonymous reviewer. Nelda Bishop proofread the typescript. Literature Cited Bishop, G. A. 1985. Fossil decapod crustaceans from the Gammon Ferruginous Member, Pierre Shale (Early Campanian), Black Hills, South Dako- ta.—Journal of Paleontology 59(3):605-624. Forir, H. 1887. Contributions a l’étude du Systéme crétacé de la Belgique.— Annales de la Socieété Géologique de Belgique 14:25-26. Forster, R. 1975. Ein Krebs aus dem oberen Campan von Misburg.—Berichte der Naturhistorischen Gesellschaft Hannover 119:285-294. Glaessner, M. F. 1969. Decapoda. Pp. R399-R651 in R. C. Moore, ed., Treatise on Invertebrate Paleontology. Part R, Arthropoda 4, Vol. 2. University of Kansas Press and Geological So- ciety of America, Lawrence, (Kansas). VOLUME 100, NUMBER 1 Mertin, H. 1941. Decapode Krebse aus dem Sub- hercynen und Braunschweiger Emscher und Untersenon, sowie Bemerkungen uber einige verwandte Formen in der Oberkreide.— Nova Acta Leopoldina, Abhandlungen der Kaiserlich Leopoldinisch-Carolinisch Deutschen Akade- mie der Naturforscher 10(68):149-264. Reuss, A. 1859. Zur Kenntnis fossiler Krabben.— Denkschriften der Kaiserlich Akademie der Wissenschaften in Wien 17:1-90. Roberts, H. B. 1956. Early Tertiary decapod crus- taceans from the Vincetown Formation in New Jersey.— Bulletin of the Wagner Free Institute of Science 31:5-12. Schlotheim, E. F. v. 1820. Die Petrefactenkunde auf ihrem jetzigen Standpunkte durch die Beschrei- bung seiner Sammlung ... erlautert. (Nach- 39 trage, etc.) 3 Pt. Gotha, Becker. 62 + 437 pp. and atlas of plates. Secretan, S. 1964. Les Crustacés Décapodes du Ju- rassique supérieur et du Crétacé de Madagas- car.—Mémoires du Muséum National d’His- toire Naturelle, série C, 14:1-226. Segerberg, K.O. 1900. De Anomura och Brachyura dekapoderna inom Skandinaviens yngre kri- ta.—Geologiska Foreningens 1 Stockholm F6r- handlingar 22(201):347-394. Institute of Arthropodology and Parasi- tology, Department of Geology and Geog- raphy, Georgia Southern College, States- boro, Georgia 30460-8149. PROC. BIOL. SOC. WASH. 100(1), 1987, pp. 40-43 NEW SUBSPECIES OF DRYOCOPUS JAVENSITS (AVES: PICIDAE) AND FICEDULA HYPERYTHRA (AVES: MUSCICAPIDAE) FROM THE PHILIPPINES Robert S. Kennedy Abstract.— Austin L. Rand prepared descriptions and named two new sub- species of Philippine birds that were never published. After examining speci- mens of the proposed subspecies and comparing them with closely allied forms, I concur with Rand’s descriptions and here formally propose recognition of the two subspecies: Dryocopus javensis cebuensis, n. subsp. (Cebu); and Ficedula hyperythra matutumensis, n. subsp. (Mt. Matutum, Mindanao). At the end of his tenure as Chief Curator of Zoology at the Field Museum of Natural History, Austin L. Rand prepared two pa- pers on Philippine birds that were never published. The first manuscript, ‘“‘The Thicket Flycatcher Muscicapa hyperythra of Mindanao,” later referred to by Rand (1970 ms) as completed in 1969, contained the description of a new subspecies of the Thicket Flycatcher, which is now usually called the Snowy-browed Flycatcher and placed in the genus Ficedula. The second manuscript, entitled ““An Annotated List of Philippine Birds’? and completed in 1970, was of book length and was the culmination of his more than two decades of work with Philippine birds. It contained the descrip- tion of a new subspecies of White-bellied Woodpecker (Dryocopus javensis). During the course of my research on Phil- ippine birds, J. T. Marshall, Jr., with the approval of G. E. Watson, kindly provided a copy of each of the Rand manuscripts which are on microfilm in the Division of Birds at the National Museum of Natural History. After discovering the unpublished descriptions, I examined specimens of the proposed subspecies, compared them to other nearby populations and concluded that Rand’s diagnoses were accurate but incom- plete and that the two subspecies are indeed valid. Here I formally describe the two subspe- cies discovered and named by Rand by pre- senting his diagnoses verbatim and by add- ing the results of my own comparative work. Dryocopus javensis: White-bellied Woodpecker Bourns and Worcester (1894), McGregor (1907, 1909), Hachisuka (1934) and Dela- cour and Mayr (1946) reported the occur- rence of the White-bellied Woodpecker on Cebu. Recently, however, Parkes (1960), duPont (1971), and Short (1982) failed to mention the Cebu population of this wood- pecker in their discussions or treatment of the Philippine subspecies. Hachisuka (1934) placed the Cebu birds in D. j. confusus of Luzon, but this was questioned by Delacour and Mayr (1946). As noted earlier, Rand (1970 ms) examined the Cebu birds and found that they repre- sent an undescribed subspecies to be known as: Dryocopus javensis cebuensis, new subspecies Holotype.—USNM 315188, adult male, 22 Jun 1892, Pandai, Cebu Island, Philip- pines, D. C. Worcester and F. S. Bourns (collectors’ no. 511 from Menage Expedi- tion). VOLUME 100, NUMBER 1 Subspecific characters. —‘‘Most like D. j. multilunatus of Mindanao in having an all black back, a moderate amount of white streaking in chin, upper throat and side of head; a scant amount of narrow pale edgings on breast feathers, and bill that is mostly black but with some yellowish or black horn in the lower mandible. “Differs from D. j. multilunatus in the shorter bill [cebuensis—(3) 46.5 + 0.64, range 46.0—47.2: multilunatus —(22) 54.4 + 2.77, range 49.0—59.1] and the shorter wing [cebuensis—(3) 195.7 + 2.08, range 194—- 198: multilunatus —(22) 209.9 + 8.30, range 197—229].”? (Rand 1970 ms). Similar to D. j. suluensis of the Sulu Ar- chipelago in most plumage characters in- cluding the presence of a concealed white patch in the lower back or rump and in being of equal size (su/uensis: bill (15) 47.6 + 1.63, range 44.9-51.7: wing (15) 193.7 + 5.15, range 186-203). Differs from su/uensis in having buffy edgings to some of the breast feathers and in having narrow buffy tips Gn two of the three specimens) to the outer primaries; some specimens of suluensis have, at most, a tiny buffy spot near the tip of one or more primaries. Dryocopus j. confusus of Luzon differs from D. j. cebuensis in being larger (confu- sus: bill (6) 198.7 + 1.21, range 197-200: bill (6) 51.9 + 2.36, range 48.0—54.8), in lacking the buffy edgings to the breast feath- ers, and in having a black bill, a mostly white throat and the black plumage char- acters a deeper black. Range. — Known only from Cebu Island, Philippines, where it has not been seen by local and visiting naturalists for several de- cades. Etymology.—Rand named this form for the island it inhabited. Remarks. —“‘It is interesting that cebuen- sis, most similar to Mindanao multilunatus [and to suluensis| and less so to Luzon con- fusus, is nearly surrounded by islands of more different forms: the white-backed phi- lippinus of Negros, Masbate, etc., and the 41 black-backed birds with heavily patterned throat and breast, pectoralis of Samar— Leyte—Bohol.”’ (Rand 1970 ms). Of the three known specimens of cebuen- sis, one (USNM 357282) is a mounted bird that was previously on display. The red of its malar mark and the top of its head are much duller and darker than the other two specimens. Its bill has been painted, and the tip of one primary, although sooty black, looks as if it may have been buffy at one time, like the outer primaries of the other two specimens. All of these plumage char- acters seem to have resulted from exposure to light and to soot while it was on display. In comparisons of cebuensis with suluen- sis, I noted that cebuensis has a concealed white patch in the lower back; thus, Rand’s mention of an all black back is incorrect. However, some specimens of multilunatus have a concealed white patch on the back as well. Parkes (1960) mentions one Basilan specimen with some white on the back and I found a concealed white patch on the lower back of at least three Basilan specimens (USNM). The differences or similarities in plumage characters noted in the description section above are valid for both sexes. Rand did not treat each sex separately presumably be- cause of the few specimens of D. j. cebuensis available for comparison. Specimens examined.—Dryocopus j. ce- buensis: 2 8, 1 2(USNM). D. j. confusus: 3 6, 3 2 (DMNB). D. j. multilunatus: Basi- lan—3 6, 2 2 (USNM); Dinagat—1 4 (DMNH); Mindanao—2 4, 2 2 (DMNH), 5 6, 6 2, 12? (USNM). D. j. suluensis: 5 6, 6 2 (DMNH); 3 4, 1 2 (USNM). Ficedula hyperythra: Snowy-browed Flycatcher As noted by Rand (1969 ms), the written history of F. hyperythra on Mindanao is short yet more forms (four subspecies in- cluding the one described here) of this species inhabit that island than of any other species (except Phylloscopus trivirgatus) with a sim- 42 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ilar range. Such divergence has come about through the isolation of populations of this species in the higher elevations (above 1000 m) of the mountains of Mindanao that have served as habitat islands for the different forms. The first subspecies discovered was F. h. montigena (Mearns 1905) from Mt. Apo, which has since been recorded on nearby Mt. McKinley and on Mt. Katanglad in cen- tral Mindanao (Ripley and Rabor 1961), and from the Piagayungan Mts. of Lanao del Sur Province (USNM 580519-580524). It is the only previously described race in the Phil- ippines in which the plumage of the male closely resembles that of the female both in the color of the upperparts and in the rufous colored tail. In 1957, D. Rabor obtained specimens of a second subspecies from Mt. Malindang in the Zamboanga Peninsula that Rand and Rabor (1957) named F. h. mal- indangensis. This form is more closely al- lied to F. h. nigrorum of Negros and shows the sexual dimorphism (including males with gray tails and females with rufous tails) typical of most races of this species. More recently, J. duPont discovered a third subspecies at Daggayan in Misamis Oriental Province in northern Mindanao, named F. h. daggayana by Meyer de Schauensee and duPont (1962). The male of daggayana has a dark tail similar to that of malindangensis even though the chest- nut-tailed montigena from Mt. Katanglad and the Piagayungan Mts. occurs between the two populations. A fourth subspecies, a chestnut-tailed form, recognized and named by Rand may be known as: Ficedula hyperythra matutumensis, new subspecies Holotype. —FMNH 275254, adult male, 23 Jan 1964, Tucay E-el, Mt. Matutum, 3300 to 3500 ft, Tupi, South Cotabato Province, Mindanao Island, Philippines, D.S. Rabor (collector’s no. 39568). Subspecific characters. —‘‘Like monti- gena in having tail and outer edges of re- miges dark red brown in both sexes but dif- fers in having the rufous of the underparts much deeper and more extensive, extending from chin to flanks and undertail coverts; male differs further in having the back more slaty with hardly a trace of olive brown pos- teriorly. Female differs further in having crown and foreback slaty gray with an olive brown wash on lower back and rump only.” (Rand 1969 ms). Similar to montigena, sex for sex, in length of tail and culmen but averages smaller than montigena, sex for sex, in wing chord (ma- tutumensis —é (8) 60.3 + 0.53, range 59.2- 61.0; 2 (7) 57.1 + 1.29, range 54.9-58.2: montigena—é (7) 62.0 + 1.31, range 60.4— 64.5; 2 (7) 58.8 + 1.30, range 56.6—-60.7) and in tarsus length (matutumensis—é (8) 17.3 + 0.24, range 17.0-17.7; 2 (5) 16.5 = 0.58, range 15.6-17.0: montigena—é (6) 18.4 + 0.26, range 18.1-18.7; 2 (7) 17.9 = 0.60, range 17.0—-18.7). “Like daggayana with underparts deeply and extensively colored from chin to un- dertail coverts. Male differing in having tail and edges of wing quills red brown as in female; side of head blacker; with little or no olive brown tinge on lower back; female differing also in having back with little or no olive brown, and in having spot in front of eye and eye ring rusty.’ (Rand 1970 ms). Measurements.—Culmen 6 (8) 13.0 + 0.26, range 12.5-13.3; 2 (7) 13.0 + 0.12, range 12.8-13.1. Tail é(8) 45.0 + 1.35, range 43.3-47.1; 2 (7) 40.8 + 1.18, range 39.8- 42.2. See above for other measurements. Weight 6 (5) 10.1 + 0.50 g, range 9.7-10.9 g; 2 (2) 9.33 + 0.11 g, range 9.25-9.40 g; 1 2 with “ripe egg” in the oviduct 12.3 g (col- lected 21 June 1966). Range.—Known only from above 1000 m on Mt. Matutum, South Cotabato Prov- ince, Mindanao Island, Philippines. Etymology. —Rand named this form for the mountain it inhabits. Remarks. — After examining more speci- mens than Rand had seen, I concluded that the best character distinguishing matutu- VOLUME 100, NUMBER 1 mensis from montigena is the absence or near absence of the olive brown tinge on the lower back of male matutumensis and the restriction of this color in females to the lower back. The more extensive and deeper rufous of the underparts is a general char- acter of matutumensis but is shared by some specimens (particularly DMNH 36738 and USNM 580521) of montigena. Differences between males of matutu- mensis and daggayana are as Rand de- scribed except for the alleged blacker sides of the head, which does not hold for all specimens of matutumensis. Females of matutumensis do have less olive brown in the back but the spot in front of the eye and the eye ring are rusty in both forms. Specimens examined. —Ficedula h. dag- gayana: | 6 (type), 1 2? (DMNH). F. h. ma- tutumensis: 3 6(AMNRH); 3 6, 3 2 (FMNH); 7 6,5 2(USNM). F. h. montigena: 1 4; 1 (AMNH); 3 6, 1 20 DMNH),; 3 6, 3 2 (FMNH); 446, 6 2 (USNM). Acknowledgments I thank the curators of the American Mu- seum of Natural History (AMNH), Dela- ware Museum of Natural History (DMNH), Field Museum of Natural History (FMNH) and the National Museum of Natural His- tory (USNM) for the loan of specimens and/ or for permission to study the specimens in their care. The Frank M. Chapman Me- morial Fund provided assistance to visit the AMNH. I thank J. T. Marshall, Jr. for pro- viding Rand’s manuscripts, K. C. Parkes for his help in examining F. hyperythra speci- mens in the FMNH, and D. Willard and F. E. Lohrer for assistance in searching for cop- ies of Rand’s notes on the two subspecies. E. C. Dickinson and K. C. Parkes kindly read an earlier draft of this manuscript. I dedicate this paper to the memory of Austin L. Rand. 43 Literature Cited Bourns, F. S., and D. C. Worcester. 1894. Prelimi- nary notes on the birds and mammals collected by the Manage Scientific Expedition to the Phil- ippine Islands.— Minnesota Academy of Natu- ral Sciences Occasional Papers 1:1-64. Delacour, J., and E. Mayr. 1946. Birds of the Phil- ippines. Macmillan, New York. xv + 309 pp. duPont, J.E. 1971. Philippine birds. — Delaware Mu- seum of Natural History Monograph 2: x + 1- 480. Hachisuka, M. 1934. The birds of the Philippine Is- lands with notes on the mammal fauna volume 2, part 3. Witherby, London. 256 pp. McGregor, R. C. 1907, Notes on birds collected in Cebu. — Philippine Journal of Science 2:298-309. . 1909. A manual of Philippine birds. Manila, Bureau of Science. No. 2. Part 1, pp. 1-412. Mearns, E. A. 1905. Descriptions of a new genus and eleven new species of Philippine birds. —Pro- ceedings of the Biological Society of Washington 18:1-8. Meyer de Schauensee, R., and J. E. duPont. 1962. Birds from the Philippine Islands. — Proceedings of the Academy of Natural Sciences of Phila- delphia 114:149-173. Parkes, K. C. 1960. Notes on Philippine races of Dryocopus javensis. — Bulletin of the British Or- nithologists’ Club 80:59-61. Rand, A.L. 1969. The Thicket Flycatcher Muscicapa hyperythra of Mindanao. Unpublished manu- script. 8 pp. 1970. An annotated list of Philippine birds. Unpublished manuscript. 511 pp. , and D. S. Rabor. 1957. New birds from the Philippines. — Fieldiana, Zoology 42:13-18. Ripley, S. D., and D. S. Rabor. 1961. The avifauna of Mt. Katanglad.—Postilla 50:1—20. Short, L. L. 1982. Woodpeckers of the world. — Del- aware Museum of Natural History Monograph 4: xvii + 1-676. Department of Zoology, Washington State University, Pullman, Washington 99164- 4220. Present address: Yale University, School of Forestry and Environmental Studies, 205 Prospect Street, New Haven, Connecticut 06511. PROC. BIOL. SOC. WASH. 100(1), 1987, pp. 44-74 THE SUBGENERA OF THE CRAWFISH GENUS ORCONECTES (DECAPODA: CAMBARIDAE) J. F. Fitzpatrick, Jr. Abstract. —The genus Orconectes, the last major cambarid genus to be reeval- uated following the major increase in species recognized in recent years, is divided into 10 subgenera. Besides the nominate subgenus, Faxonius Ortmann, 1905, is resurrected and the new names Billecambarus, Buannulifictus, Crock- erinus, Gremicambarus, Hespericambarus, Procericambarus, Rhoadesius, and Tragulicambarus proposed. Each is diagnosed and the type species illustrated; six species-groups are recognized in Crockerinus, four in Procericambarus, and © two each in Buannulifictus, Gremicambarus and Hespericambarus, with Bil- lecambarus and Tragulicambarus being monotypic. The divisions are justified with a phylogenetic discussion of morphological, geographical, and to a lesser extent, temporal considerations. In recent years all of the major genera in the Cambaridae, except Orconectes, have been reevaluated. The tremendous increase in the number of recognized species has al- most mandated that the members of the larger genera be grouped into natural as- sociations at levels lower than genus, assem- blages which have been formally recognized as subgenera and/or “Sections.” Hobbs (1969) began the current reassessments with a study of Cambarus in which he recognized several subgenera, erected the genus Falli- cambarus for one divergent group of species and recognized the validity of Hobbseus, proposed by Fitzpatrick and Payne (1968). Hobbs then turned his attention to Procam- barus and identified a number of subgeneric groupings within it (1972). Fallicambarus likewise was discovered to consist of two subgeneric-level assemblages (Hobbs 1973). Recently, Fitzpatrick proposed subgenera for the monogeneric Cambarellinae (1983). Also recent is a series of events beginning with Hobbs’ (1981) discovery of a new group, Distocambarus, which was first pro- posed as a subgenus of Procambarus, soon elevated to generic rank (Hobbs and Carl- son 1983), and then divided into two sub- genera (Hobbs 1983). Associated with these major revisions, the past two decades have seen miscellaneous reassignments of species groups, principally by the elevation of subgenera to genera (Fitzpatrick 1963, Bouchard 1972). Also a new subgenus was proposed to receive a dis- junct species, newly-discovered, and assign- able to Cambarus (Bouchard and Hobbs 1976), and the similarly erected genus Bou- chardina Hobbs, 1977, was offered. For 25 years I have been studying the members of the genus Orconectes, second only to Procambarus in the number of de- scribed species assigned to it. The species have been assembled into “Groups” and the latter into “Sections,” but they are essen- tially the same divisions proposed by Ort- mann (1905) and modified by Creaser (1934). The number of species assigned to this genus has nearly doubled since then, but there has been no comprehensive re- view of the interspecific relationships or an attempt to reorganize the species into groupings which reflect this added knowl- edge. There are still several taxa which are VOLUME 100, NUMBER 1 known to exist but await formal description; likewise uncertain are the precise limits of variation in many species (most recently noted by Cooper and Hobbs 1980: 1-2). Only two small groups of the genus have been studied in detail (Fitzpatrick 1967, Hobbs and Barr 1972). Nonetheless, certain clearly related groups seem obvious, and in com- paring them some phylogenetic trends are suggested. The nomenclatorial history of Orconectes is relatively straightforward. First proposed for troglobitic animals and established upon characteristics which are essentially adap- tations to the spelean environment, the ge- nus was inadequately defined (Cope 1872). Most of the early writers followed Faxon’s (1885) lead in rejecting the genus. In 1905, Ortmann offered a scheme of classification for North American (=Cambaridae) craw- fishes in which he proposed several subgen- era, one of which was Faxonius with As- tacus limosus Rafinesque, 1817, the second species known from North America, the type species. Faxon (1914) vigorously rejected this system, but in the interim Fowler (1912) accepted Faxonius as a subgeneric name and designated Orconectes inermis Cope, 1872, as the type species of Orconectes Cope, 1872. In 1933, Creaser accepted Faxontius, ele- vated it to generic level, and proposed a new subgenus, Faxonella, to receive the quite divergent Cambarus clypeatus Hay, 1899. In a major reorganization, Hobbs (1942) argued cogently and persuasively for the recognition of Ortmann’s (1905) subgenera, but as genera, with the exceptions that Fax- onius, aS a subjective junior synonym, be replaced by Orconectes. Since then Orco- nectes has been accepted according to his definition with one notable exception. Creaser (1962) rejected many of Hobbs’ ar- guments and proposed different generic as- signments. Of interest here is only that Or- conectes (sensu Hobbs) was divided into three equivalent “‘generic patterns”: Fax- onius, Faxonella, and Orconectes (s. s.). To 45 the latter he assigned Cambarus lancifer Hagen, 1870, Orconectes inermis Cope, 1872, and Astacus pellucidus Tellkampf, 1844 “(with subspecies—if they are subspe- cies)” (1962:3); Faxonella contained Cam- barus clypeatus and, presumably, Orco- nectes (Faxonella) beyeri Penn, 1950. In general his system was rejected in favor of Hobbs’ with the exception that Fitzpatrick’s (1963) formal elevation of Faxonella to ge- neric status is commonly accepted. No oth- er major changes in nomenclature have been suggested. The synonymies at most levels in the present paper have been kept to a minimum for brevity’s sake. The reader is referred to Hobbs’ extensive synonymies (1974b) if more detailed information is required. Taxonomic Characters in Orconectes As is the case with most crawfishes, cer- tain of the obvious external morphological features, useful for the recognition of species, are readily modified to adapt to the envi- ronmental conditions in which the animal lives. Such features are difficult to use in establishing intergroup relationships except in the broadest sense. On the other hand, structures associated with amplexus are less susceptible to environmental modification. In Orconectes, however, even these cannot be easily analyzed. For example, in the Pro- pinquus Group, studied in detail by Fitz- patrick (1967), a number of interspecific “hybrids” are reported in the literature (Crocker 1957, Crocker and Barr 1968). But Fitzpatrick (1967) strongly implied that nat- ural hybridization was a rare occurrence. Recently, Capelli and Capelli (1980) and Smith (1981b) reported “hybrids” between species that I propose should be assigned to separate subgenera (!) below. I have no reason to question the veracity of the reports of any of the workers men- tioned above. Yet I remain firmly con- vinced that no extensive interspecific hy- bridization occurs in Orconectes or any other 46 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON crawfish genus. It is very difficult to con- ceive of a species retaining its identity with- out reproductive isolation. Too many data exist not to believe that there are many, many species of crawfishes in North Amer- ica. Smith himself (1981b), as he offers an explanation of events leading to the possible origins of his specimens, acknowledges that the occurrence of his apparent hybrids is probably a transient one and related to the quite artificial situation of an introduction of members of an alien population of a pre- viously allopatric species. When such events occur, I would expect responses not unlike the well documented one in Bufo (e.g., Blair 1941, Cory and Manion 1955). Possibly the most revealing facts related to hybridization are in the experimental data gathered by Tierney and Dunham (1984) which seem to indicate that inability to recognize a con- specific mate exists principally in species which, because of allopatry, lack a stimulus to develop isolating chemical cues; natu- rally sympatric populations are probably more accurate in mate selection. Berrill (1985) assembled data from laboratory-in- duced interspecific matings between O. pro- pinquus and O. rusticus, two species in competition as the latter is expanding its range, which demonstrated that such cross- ings reduced significantly the reproductive success in both species (or more specifically, from a natural standpoint, in individuals unable to identify their own species), similar indeed to the Bufo situation. Equally, statistics fail to provide clear in- dication of relationships. Structures asso- ciated with amplexus in crawfishes are very variable and extensive overlaps occur in meristic and morphometric data (Fitzpat- rick 1967; Chambers et al. 1979, 1980; Tier- ney 1982). Sometimes sophisticated statis- tical methodology is overtly misleading. For example, when a local population of O. pro- pinquus (Girard) was subjected to discrim- inant analysis of morphology, it seemed to consist of two distinct morphological groups; pleopod morphology was among the highest discriminant functions (Fitzpatrick and Pickett 1980). Shortly thereafter, Smith (198 1a) extended the examination to a larg- er—geographically and numerically—data base and determined that the proposed sep- aration of forms was not justified. Despite these difficulties, it is still possi- ble to discern certain morphological asso- ciations which enable the recognition of species groups. In first form males, the first pleopods are straight or curved with respect to the long axis of the appendage. One may determine the proportions of the terminal elements with respect to each other and to the appendage as a whole; the elements can range from subsetiform to stout and blade- like. In females, the degree of sculpture of the annulus ventralis can be described in generalities that lead to groupings compat- ible with those based on pleopod morphol- ogy. Although no longer are groupings rec- ognized that are based upon such characters as the areola, the spinose ornamentation of the cephalothorax and the structure of the cheliped, they, too, often exhibit a similarity compatible with groupings based on struc- tures associated with amplexus. Using these, a natural system of classification can be es- tablished. Notes on the Orconectes Annulus The seminal receptacle of the Cambari- dae is a potentially very useful tool for the taxonomist. Despite this, it has received lit- tle attention. Without the artistic skills of Hobbs, our knowledge would be little ad- vanced beyond the level of the early part of the century (Andrews 1906a, b). Perhaps the most comprehensive collation of annulus morphology is in Hobbs’ checklist (1974b), but all of his species descriptions and many of his reviews contain realistic, detailed rep- resentations of the structure. Only one study of variation in any species has been made: for O. propinquus (Tierney 1982); and very few descriptive terms have been established for features of the annulus. Hobbs (1981: 10, fig. 4b) provided a labelled figure and a VOLUME 100, NUMBER 1 brief verbal discourse on its anatomy. Fitz- patrick (1983) added some observations es- pecially pertinent to the pendulent annulus as found in the Cambarellinae. To discuss Orconectes, however, requires the addition of some terminology. Hobbs’ figure (198 1:fig. 4b) is essentially that of a procambarid, not undesirable as most consider that the family arose from a procambaridlike ancestor. But in most Or- conectes there are prominent tubercular or lobular elevations associated with the ce- phalolateral surfaces (the cephalolateral prominences). These may be united-to varying degrees—along the cephalomedian margin, or they may be separated by a depression of varying development which I propose to call the trough. I interpret the fossa, as illustrated by Hobbs (1981), to be the fundus of a pit formed by a deep ingres- sion of the sinus. In many Orconectes there iS a conspicuous transverse subovate depression, located caudal to the cephalo- lateral prominences but cephalic to the higher caudal margins and not traversing the entire width of the annulus. I suggest that this depression be called the sulcus. The relationships of the sinus and fossa to this sulcus seem to be significant. One need only to refer to Tierney’s (1982) difficulties in the application of the word “‘groove’”’ to appre- ciate the need for the introduction of such terminology. Generic and Subgeneric Diagnoses of Orconectes Family Cambaridae Hobbs, 1942 Subfamily Cambaridae Hobbs, 1942 Genus Orconectes Cope, 1872 Diagnosis.—“‘Antenna never with con- spicuous fringe on mesial border. Third maxilliped with teeth on mesial margin of ischium. Mesial margin of palm of chela usually with row of less than 12 tubercles; lateral margin of fixed finger never bearing row of spiniform tubercles; opposable mar- gin of dactyl seldom with prominent exci- sion. Areola broad to obliterated at mid- 47 length. Ischium of third, rarely third and fourth, pereiopod with hook. Coxa of fourth pereiopod of male lacking caudomesial boss except in troglobitic members. First pleo- pod of first form male almost always sym- metrical, never deeply withdrawn between bases of pereiopods nor concealed by dense setal mat extending from ventrolateral mar- gins of sternum, and contiguous basally; ter- minal elements (usually 2, occasionally 3 in troglobitic members) highly variable in length and disposition—divergent, straight, or curved caudodistically or caudally; cen- tral projection never abruptly curved cau- dally at base nor forming arc of more than 90 degrees. Female with annulus ventralis immovable or slightly movable in troglo- bitic species; first pleopod usually present. Branchial formula 17 + ep.’’ (Hobbs 1974a: 14-15). Billecambarus, new subgenus Figs. 1, 14a Diagnosis. —(Based on first form male and female.) Body and eyes pigmented, latter well developed. Rostrum with small mar- ginal spines, median carina absent. Cervical spines much reduced or absent; areola about 9.5 times longer than wide with 1 or 2 punc- tations across narrowest part, and consti- tuting about 34-35% of total length of car- apace; cephaloventral surface of carapace with small squamous tubercles; devoid of spines or tubercles in hepatic region. First pleopod of male ending in 2 terminal ele- ments, elements short (about 15% total length of pleopod), subparallel, both curved caudodistally from base so that apices di- rected at angle of nearly 90° to main axis of basal portion of pleopod; mesial process slender; distal fourth of pleopod inclined caudodistally at angle of about 30°; shoulder on cephalic surface of pleopod just proximal to aforementioned inclination not strongly developed or sharply angular. Inner margin of palm of chela about 28.5% of length of outer margin; opposable margins of im- movable finger and dactyl with prominent 48 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON VOLUME 100, NUMBER 1 subequal tubercles in basal half; tuft of setae at base of immovable finger, if present, quite weakly developed. Annulus ventralis of fe- male immovable, firmly attached to pre- ceding sclerite, widest near midlength and only slightly more than twice as wide as long; cephalolateral prominences tubercu- liform and narrow; sulcus deep and arching through 80% of cephalic half of annulus; fossa not conspicuous with sinus traversing deep depression inclined at angle of about 20° to transverse axis about midpoint of annulus, then turning sharply caudally fol- lowing shallower depression to caudal mar- gin. Type species. —Cambarus Harrisonii Faxon, 1884:130. List of species. —Monotypic, Orconectes (Billecambarus) harrisonii (Faxon, 1884: 130). Gender. — Masculine. Etymology. — Bille (German = axe) plus the generic name, Cambarus, because the first pleopod of the male reminds me of a bill-hook used in pruning plants. Buannulifictus, new subgenus Figs. 2, 13b Diagnosis. —(Based on first form male and female.) Body and eyes pigmented, latter well developed. Rostrum with marginal spines, lacking median carina. Cervical spines if present reduced; areola obliterated, linear, or with room for no more than 2 punctations in narrowest part, constituting 31-36% total length of carapace; cephal- oventral surface of carapace with small squamous tubercles; devoid of spines or tu- bercles in hepatic region. First pleopod of male ending in 2 terminal elements of sub- — 49 equal length (except in O. m. meeki), mod- erately long (36-45% total length of pleo- pod), and both curved caudodistally throughout length (except in O. m. meeki), mesial process subsetiform; cephalic mar- gin of pleopod lacking shoulder. Inner mar- gin of hand 28-31% of length of outer mar- gin; opposable margin of immovable finger with prominent (except in O. palmeri creo- lanus), subequal tubercles in basal half; op- posable margin of dactyl with prominent (except in O. p. palmeri, O. p. creolanus, and O. hobbsi), subequal tubercles; tuft of setae at base of immovable finger not well developed but present in most specimens of O. palmeri subspp. Annulus ventralis of fe- male immovable, firmly attached to pre- ceding sclerite, widest caudal to midlength and about as wide as long; cephalolateral prominences lobiform and weakly devel- oped in most specimens; sulcus when ob- vious sharply constricted laterally but obvi- ous central depression always present; trough usually present but not deep or conspicuous; sinus originating in fossa set at acute angle to longitudinal axis of annulus and winding sinuously caudad but lost before reaching caudal margin. Type species. —Cambarus Palmeri Fax- on, 1884:124. List of species. —Palmeri Group (areola linear or obliterated; central projection more than 40% of total length of pleopod): Orconectes (Buannulifictus) denae Reimer and Jester, 1975:124. O. (B.) hobbsi Penn, 1950:381. O. (B.) palmeri palmeri (Faxon, 1884:124). O. (B.) palmeri creolanus (Creaser, 1933: 16). Figs. 1-3. Type species of Orconectes subgenera (all not to same scale): 1, Orconectes (Billecambarus) har- risoni; 2, O. (Buannulifictus) palmeri palmeri; 3, O. (Crockerinus) sanbornii sanbornii. a, Dorsal view of carapace; b, Mesial view of first pleopod of first form male; c, Lateral view of first pleopod of first form male; d, Annulus ventralis of female; e, Lateral view of first pleopod of second form male; f, Dorsal view of chela of first form male. 50 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON O. (B.) palmeri longimanus (Faxon, 1898: 65/5): Meeki Group (areola with room for at least one punctation across narrowest part; central projection less than 40% of total length of pleopod): Orconectes (Buannulifictus) meeki meeki (Faxon, 1898:657). O. (B.) meeki brevis Williams, 1952:348. Gender. — Masculine. Etymology. —Bu- (L., =large, prefix) + annulus (L., =ring) + fictus (L., =form), an allusion to the large annulus ventralis char- acteristic of members of this subgenus. Crockerinus, new subgenus Figs. 3, 13a Description. —(Based on first form male and female.) Body and eyes pigmented, lat- ter well developed. Rostrum with marginal spines, median carina present or absent. Cervical spines usually present and mod- erately well developed; areola 4—10 times longer than wide with room for more than 2 punctations in narrowest part, constitut- ing 29-36% total length of carapace; ce- phaloventral surface of carapace with small squamous tubercles, devoid of spines or tu- bercles in hepatic region. First pleopod of male ending in 2 terminal elements of sub- equal length or with mesial process slightly shorter, moderately long (29-36% of total length of pleopod), subparallel (except in O. shoupi); basal half of both elements sub- parallel to main axis of pleopod, distal half continuing subparallel to axis or curved caudodistally about 45°; mesial process sub- setiform; cephalic margin of pleopod with or without shoulder. Inner margin of hand 29-36% length of outer margin; opposable margins of both fingers with or without prominent tubercles in basal half, tubercles always subequal in size; small tuft of setae at base of immovable finger present or ab- sent. Annulus ventralis of female immov- able, firmly attached to preceding sclerite, widest at or slightly cephalic to midlength and 1.6—2.1 times wider than long; ceph- alolateral prominences lobiform or tubercu- liform, moderately or well developed; sul- cus varying from shallow and obscure to well developed (not developed in O. bisec- tus, O. shoupi, and O. tricuspis); trough present and well developed to absent; fossa when present small but deep (linear in O. illinoiensis), located near midpoint of an- nulus, sinus usually moving sharply later- ally and then recurving to near midline be- fore progressing sinuously caudad but lost before reaching caudal margin (except in O. erichsonianus). Type species. —Cambarus Sanbornii Faxon, 1884:128. List of species. —Sanbornii Group (cen- tral projection 24—29% total length of pleo- pod, distal half straight— except extreme tip often arced caudally—and subparallel to mesial process; lacking distinct gap between bases of fingers; opposable margin of fixed finger usually with one tubercle larger than rest; annulus 1.4—1.7 times wider than long, moderately developed cephalolateral prom- inences oriented at distinct angle to trans- verse axis, trough obscure, and sinus orig- inating near midpoint of annulus and with distinctly laterally oriented section in an- terior half): Orconectes (Crockerinus) obscurus (Hagen, 1870:69). O. (C.) sanbornii sanbornii (Faxon, 1884: 128). O. (C.) sanbornii erismophorus Hobbs and Fitzpatrick, 1962:208. O. (C.) stannardi Page, 1985:564. O. (C.) virginiensis Hobbs, 1951:122. Marchandi Group (central projection 28— 29% total length of pleopod, distal half straight or curved caudodistally — marchan- di—and subparallel to mesial process; lack- ing distinct gap between bases of fingers; opposable margin of fixed finger usually with one tubercle larger than rest; annulus 1.5— 1.6 times wider than long, well developed VOLUME 100, NUMBER 1 cephalolateral prominences oriented at dis- tinct angle to transverse axis, trough well developed, and sinus originating near mid- point of annulus and lacking distinctly lat- erally oriented section in anterior half): Orconectes (Crockerinus) eupunctus Wil- liams, 1952:330. O. (C.) marchandi Hobbs, 1948b:140. Propinquus Group (central projection 30— 35% total length of pleopod, distal half straight—except extreme tip sometimes arced caudally—and subparallel to mesial process; lacking—except in jeffersoni—dis- tinct gap between bases of fingers; opposable margin of fixed finger without one tubercle larger than rest; annulus 1.7—1.9 times wider than long, weakly to moderately developed cephalolateral prominences oriented at dis- tinct angle to transverse axis, trough weakly or only moderately developed, and sinus originating near to or just lateral to mid- point of annulus and distinctly laterally ori- ented section in anterior half): Orconectes (Crockerinus) erichsonianus (Faxon, 1898:659). O. (C.) jeffersoni Rhoades, 1944:123. O. (C.) propinquus (Girard, 1852:88). Rafinesquei Group (central projection 21- 29% total length of pleopod, distal half straight—except extreme tip often arced caudally—and subparallel to mesial pro- cess; lacking distinct gap between bases of fingers; opposable margin of fixed finger with or without one tubercle larger than rest; an- nulus 1.6—-1.8 times wider than long, ce- phalolateral prominences lobiform or broadly tuberculiform and always well de- veloped, oriented—often as much as 90°— at distinct angle to transverse axis, trough distinct and usually moderately deep, and sinus may or may not originate in deep fossa near or slightly lateral to midpoint of an- nulus before following undulant longitudi- nal route not quite to caudal margin, sinus with—except in bisectus—distinctly later- ally oriented section in anterior half): Sill Orconectes (Crockerinus) bisectus Rhoades, 1944:129. O. (C.) illinoiensis Brown, 1956:163. O. (C.) rafinesquei Rhoades, 1944:116. O. (C.) tricuspis Rhoades, 1944:117. Shoupi Group (central projection about 21% total length of pleopod, straight, sub- parallel to mesial process in basal half but distal half of latter curved caudodistally at angle of about 45° and not subsetiform; dis- tinct gap between bases of finger about as wide as width of base of dactyl; opposable margin of fixed finger without one tubercle larger than rest; annulus about 2.4 times wider than long, well developed tuberculi- form cephalolateral prominences oriented subparallel to transverse axis and occupying much of cephalic half of annulus, trough narrow but distinct, sinus originating on midline in cephalic third of annulus, arcing caudolaterally to point caudolateral to mid- dle of annulus, recurving to midpoint before turning sharply caudadally to proceed in substraight line almost to caudal margin): Orconectes (Crockerinus) shoupi Hobbs, 1948a:14. Gender. — Masculine. Etymology. —Named in honor of Denton W. Crocker in recognition of his lifelong study of many of the species assigned to this subgenus. Subgenus Faxonius Ortmann, 1905:97 Figs. 4, 15a Faxonius (subgeneric name): Ortmann, 1905 (part).— Fowler, 1912 (part). Faxonius (generic name): Creaser, 1933 (part); 1962 (part). [For a fuller synonymy and explanation see Hobbs 1942:339, 350-352; 1974a:14; 1974b:26.] Diagnosis. —(Based on first form male and female.) Body and eyes pigmented, latter well developed. Rostrum with marginal spines, median carina absent. Cervical spines well developed (except in O. indl- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 52 Oe | a e FE e LO nary, COW ge OO eA Medtasy VOLUME 100, NUMBER 1 anensis); areola 6.0—8.0 times longer than wide with room for more than 2 punctations in narrowest part, and constituting 31-33% total length of carapace; cephaloventral sur- face of carapace with small squamous or spinose tubercles, hepatic region with (in O. limosus) or without spines. First pleopod of male ending in 2 terminal elements of sub- equal length, short (less than 20% total length of pleopod), straight but divergent at angle of about 20°, neither (except central projec- tion in O. wrighti) subparallel to main axis of pleopod; mesial process slender and ta- pering apically but not subsetiform; distal half of pleopod (except in O. wrighti) slight- ly inclined caudodistally; cephalic margin of pleopod lacking shoulder. Inner margin of hand 33—40% length of outer margin; op- posable margins of fingers lacking promi- nent tubercles in basal half, setose margins obscuring small tubercles; tuft of setae at base of immovable finger lacking. Annulus ventralis of female immovable, firmly at- tached to preceding sclerite, widest near midlength and !.9-—2.1 times wider than long; cephalolateral prominences tubercu- liform and well developed, occupying much of cephalolateral half of annulus; sulcus arising lateral to midline of annulus then moving to it before turning sharply caudally forming only very slightly undulant line disappearing before reaching caudal mar- gin. Type species.—Astacus limosus Rafines- que, 1817:42. Designated by Ortmann 1905: 97. List of species. — Orconectes (Faxonius) indianensis (Hay, 1896:494). O. (F.) limosus (Rafinesque, 1817:42). O. (F.) wrighti Hobbs, 1948c:85. — 53 Gremicambarus, new subgenus Figs. 5, 12 Diagnosis. —(Based on first form male and female.) Body and eyes pigmented, latter well developed. Rostrum with or without marginal spines, median carina absent (ex- cept in O. alabamensis). Cervical spines much reduced, absent, or only moderately well developed; areola obliterated, linear or to 3.5 times longer than wide with room for more than 2 punctations in narrowest part of wider areolae, constituting 25—40% total length of carapace; cephaloventral surface of carapace with small squamous tubercles; devoid of spines and tubercles in hepatic region. First pleopod of male ending in 2 terminal elements, elements moderately long to long (20-40% total length of pleopod), subparallel or divergent with central pro- jection subparallel to main axis of pleopod through at least 90% length or curved cau- dodistally or inclined caudodistally; ele- ments subequal in length with mesial pro- cess tapering evenly from base to tip or deflected sharply caudodistally (to 90°) in apical 15% with distal part subspatulate and cephalically excavated; cephalic surface of pleopod lacking shoulder. Inner margin of hand 25-43% length of outer margin; op- posable margin of immovable finger with or without (O. alabamensis, O. compressus, O. rhoadesi) row of prominent tubercles in bas- al half and if present one markedly larger than rest; opposable margin of dactyl also with tubercles likewise disposed except one never markedly larger than rest; tuft of setae of varying degree of development present at base of immovable finger (except in O. compressus and O. mississippiensis). An- nulus ventralis of female immovable, firmly attached to preceding sclerite, widest near Figs. 4-6. Type species of Orconectes subgenera (all not to same scale): 4, Orconectes (Faxonius) limosus; 5, O. (Gremicambarus) virilis; 6, O. (Hespericambarus) difficilis. a, Dorsal view of carapace; b, Mesial view of first pleopod of first form male; c, Lateral view of first pleopod of first form male; d, Annulus ventralis of female; e, Lateral view of first pleopod of second form male; f, Dorsal view of chela of first form male. 54 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON midlength (except caudal to midlength in O. alabamensis) and (except in O. alaba- mensis and O. immunis) length and width subequal; cephalolateral prominences well developed and occupying most of width of annulus; trough of varying width and depth but always clearly evident; sulcus deep, usu- ally partially overhung along some of its width by caudal parts of cephalolateral prominences; fossa deep although some- times obscured by overhang of cephalolat- eral prominences with sinus originating lat- eral to midline in cephalic half of annulus and moving transversely across midline where making U-shaped turn to midline and thence caudad for varying distance but nev- er so far as caudal margin of annulus. Type species.—Cambarus virilis Hagen, 1870:63. List of species.—Virilis Group (central projection greater than 35% total length of pleopod and reaching coxa of first pleopod; mesial process subsetiform or tapering base to tip; caudal margin of annulus somewhat angular): Orconectes (Gremicambarus) causeyi Jest- Be, MNO Wed lisse O. (G.) nais (Faxon, 1885:140). O. (G.) virilis (Hagen, 1870:63). Alabamensis Group (central projection less than 35% total length of pleopod and reaching no farther anteriorly than coxa of second pleopod; mesial process apically subspatulate; caudal margin of annulus gen- erally rounded): Orconectes (Gremicambarus) alabamensis (Faxon, 1884:125). O. (G.) chickasawae Cooper and Hobbs, 1980:29. O. (G.) compressus (Faxon, 1884:127). O. (G.) cooperi Cooper and Hobbs, 1980: le O. (G.) etnieri Bouchard and Bouchard, 1976b:459. O. (G.) holti Cooper and Hobbs, 1980:23. O. (G.) immunis (Hagen, 1970:71). O. (G.) mississippiensis (Faxon, 1884:123). O. (G.) rhoadesi Hobbs, 1949:19. O. (G.) validus (Faxon, 1914:382). Gender. — Masculine. Etymology.—Gremius (Latin = middle, center) in combination with the generic name, Cambarus, an allusion to the dom- inance of this subgenus in the central part of North America. Hespericambarus, new subgenus — Figs. 6, 16a Diagnosis. —(Based on first form male and female.) Body and eyes pigmented, latter well developed. Rostrum with marginal spines, median carina absent. Cervical spines well developed; areola obliterated and constituting 3 1—-33% total length of carapace; cephaloventral surface of carapace with small squamous tubercles; devoid of spines or tubercles in hepatic region (except small spines in O. blacki). First pleopod of male terminating in 2 elements, elements short (less than 20% total length of pleopod) and unequal in length (except in O. blacki and O. maletae), apices divergent with longer central projection subparallel to main axis of pleopod or deflected caudodistally to an- gle of 45°, mesial process deflected 30—90°; shoulder present on cephalic surface of pleo- pod only in O. difficilis. Inner margin of hand 24-31% length of outer margin; op- posable margins of immovable finger and dactyl with prominent subequal tubercles in basal half (O. difficilis with one, centrally located, larger than rest on both fingers); tuft of setae small but obvious at base of im- movable finger. Annulus ventralis of female immovable, firmly attached to preceding sclerite, widest at midlength or nearly so, about as wide as long; cephalolateral prom- inences lobiform or obscure with poorly de- fined trough visible only in O. difficilis and O. hathawayi; sulcus-like structure evident only in O. maletae and there caused by sin- VOLUME 100, NUMBER 1 gle cephalomedian prominence; fossa if present poorly defined and sinus describing simple undulant line in caudal half of an- nulus but not reaching caudal margin. Type species.—Cambarus difficilis Fax- on, 1898:656. List of species. —Difficilis Group (mesial process sharply recurved and approaching subsetiform; central projection slender; an- terior portion of sinus of annulus deeply incised): Orconectes (Hespericambarus) difficilis (Faxon, 1898:656). O. (H.) maletae Walls, 1972:456. Hathawayi Group (mesial process broad and tapering from base to tip, not recurved more than 45°; central projection laterally compressed; sinus of annulus shallowly in- cised): Orconectes (Hespericambarus) blacki Walls, 1972:454. O. (H.) hathawayi Penn, 1952:1. O. (—H.) perfectus Walls, 1972:451. Gender. — Masculine. Etymology. — Hesperius (Latin = west- erm) in combination with the generic name, Cambarus, referring to the generally west- ern distribution of the members of this sub- genus. Subgenus Orconectes Cope, 1872:419 Figs. 7, 8, 16b Orconectes: Cope, 1872.—Fowler, 1912.— Hobbs, 1942.—Creaser, 1962 (part). Faxonius: Ortmann, 1905 (part) (subgener- ic name).—Creaser, 1933 (part) (subge- neric name). [For a fuller synonymy and explanation see Hobbs 1942:339, 350-352; 1974a:14; 1974b:26.] Diagnosis. —(Based on first form male and female.) Albinistic; eyes reduced and with- out pigment. Rostrum with or without mar- ginal spines; median carina absent. Cervical DD spines well developed to scarcely observ- able; areola 3.5—6.5 times longer than wide with room for more than 2 punctations across narrowest part, constituting 34-46% total length of carapace; cephaloventral sur- face of carapace usually with small spinose Or squamous tubercles; hepatic region usu- ally (except in O. inermis testii) with one to many spines of varying degrees of devel- opment. First pleopod of male ending in 2 or 3 very short (less than 10% total length of pleopod) terminal elements, caudal pro- cess if present always small and often ves- tigal; mesial process tapering from base to tip, divergent distally and usually subequal in length to central projection (but in O. pellucidus mesial process at least twice as long as central projection); central projec- tion more or less subparallel to main axis of pleopod; rounded shoulder (often angular but always small in O. australis packardi) present in O. australis subspp. and O. in- comptus but absent in O. inermis subspp. and O. pellucidus. Inner margin of hand 37- 39% length of outer margin; opposable mar- gin of immovable finger with at least one small tubercle in basal half, usually more and (except in some specimens of O. pel- lucidus) with one larger than rest; opposable margin of dactyl with 3—5 small tubercles in basal half but only O. australis subspp. with one larger than rest. Annulus ventralis of female slightly movable, always separat- ed from preceding sclerite by nonsclerotized cuticle, widest at or slightly cephalic to mid- length or with measurements subequal; cephalolateral prominences in strict sense absent but prominent longitudinal ridges along midline occupying cephalic two-thirds or more of annulus; ridges usually separated by narrow shallow median groove probably representing trough; sometimes shallow transverse depression associated with cau- dal margins of aforementioned ridges per- haps representing rudimentary sinus; fossa if present poorly developed and sinus usu- ally arising anterior to it in caudal fourth of 56 HOOKS BOSS Figure 8 Figure 9 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON VOLUME 100, NUMBER 1 annulus near midline and arcing gently to or nearly to caudal margin. Type species. —Orconectes inermis Cope, 1872:449; designated by Fowler 1912:339 as type-species of the genus Orconectes. List of species. — Orconectes (Orconectes) australis australis (Rhodes, 1941:142). O. (O.) australis packardi Rhoades, 1944: 121. O. (O.) incomptus Hobbs and Barr, 1972: 32. O. (O.) inermis inermis Cope, 1872:449. O. (O.) inermis testii (Hay, 1891:148). O. (O.) pellucidus (Tellkampf, 1844:684). Procericambarus, new subgenus Figs. 9, 17 Diagnosis. —(Based on first form male and female.) Body and eyes pigmented, latter well developed. Rostrum with marginal spines or distinctly angular cephalic termini of margins so that bases of acumen clearly delimited; median carina present or absent. Cervical spines present and well developed to scarcely observable; areola 4.5—17.5 times longer than wide with room for at least 2 and usually more punctations across nar- rowest part, and constituting 29-37% total length of carapace; cephaloventral surface of carapace with small squamous tubercles; devoid of spines or tubercles in hepatic re- gion. First pleopod of male ending in 2 ter- minal elements, elements long (34-55% to- tal length of pleopod) and of subequal length or with central projection 10-20% longer than mesial process, subparallel or very slightly divergent (artifactual divergence in preserved specimens not uncommon); cen- — a tral projection subsetiform, straight and subparallel to main axis of pleopod or gently arced, apical 5—10% curved sharply cau- dodistally or distally so that apex directed nearly 90° to main axis of basal part of pleo- pod; mesial process subsetiform and apex rounded distally (except subspatulate and cephalically excavated in O. nana and O. macrus), usually subparallel to main axis of pleopod for most of length but distal 10- 20% often arced cephalodistally; cephalic surface of pleopod with or without promi- nent sharply angled shoulder just proximal to base of central projection. Inner margin of hand 24-38% length of outer margin; op- posable margin of immovable finger with prominent tubercles in basal half (except in O. forceps and O. mirus), only rarely (in O. longidigitus and O. williamsi) with one larg- er than remainder; opposable margin of dactyl with small or low scale-like tubercles (except prominent in O. longidigitus, O. ozarkae, and O. williamsi, and unequal in size), all subequal in size; tuft of setae if present at base of immovable finger poorly developed, prominent only in O. ozarkae. Annulus ventralis of female immovable, firmly attached to preceding sclerite, widest near midlength and distinctly wider than long if caudal overhang present in some species not considered; cephalolateral prominences large and lobiform (except re- duced to low ridges in O. quadruncus); trough distinct and usually deep (except in O. ozarkae, O. quadruncus and some spec- imens of O. longidigitus); sulcus deep and obvious (shallow but usually distinct in O. quadruncus), cephalic parts often obscured by overhang of cephalolateral prominences; sinus arising in distinct deep fossa (fossa sometimes obscured by overhanging ceph- Figs. 7-9. Type species of Orconectes subgenera (all not to same scale): 7 and 8, Orconectes (Orconectes) inermis inermis; 8, ventral view of posterior thorax of first form male (redrawn from Hobbs and Barr 1972); 9, O. (Procericambarus) forceps. a, Dorsal view of carapace; b, Mesial view of first pleopod of first form male; c, Lateral view of first pleopod of first form male; d, Annulus ventralis of female; e, Lateral view of first pleopod of second form male; f, Dorsal view of chela of first form male. 58 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON alolateral prominences) in cephalic part of sulcus, curving sharply laterad, executing U-turn at midline, then extending sinuously caudad to disappear before reaching caudal margin. Type species. —Cambarus forceps Faxon, 1884:133. List of species. — Forceps Group (terminal elements of pleopod of unequal length, cen- tral projection 34—40% total length of pleo- pod, both elements subsetiform; cephalic shoulder present or absent; annulus about twice as wide as long, caudal margin round- ed—except in O. neglectus chaenodacty- /us—and with overhanging cephalolateral prominences creating large sulcal cavity): Orconectes (Procericambarus) barrenensis Rhoades, 1944:125. O. (P.) forceps (Faxon, 1884:133). O. (P.) longidigitus (Faxon, 1898:653). O. (P.) mirus (Ortmann, 1931:81). O. (P.) neglectus neglectus (Faxon, 1885: 142). O. (P.) neglectus chaenodactylus Williams, 1952:344. O. (P.) placidus (Hagen, 1870:65). O. (P.) rusticus (Girard, 1852:88). Hylas Group (terminal elements of pleo- pod of markedly unequal length, central projection 48-52% total length of pleopod and subsetiform, mesial process subseti- form or apex bluntly rounded or apex ex- panded and cephalically excavate; cephalic shoulder present; annulus about as long as wide or slightly longer than wide and with caudal margin produced into tongue-like projection which overhangs following scler- ite); Orconectes (Procericambarus) acares Fitz- patrick, 1965:87. O. (P.) hylas (Faxon, 1890:632). O. (P.) leptogonopodus Hobbs, 1948b:146. O. (P.) peruncus (Creaser, 1931:7). O. (P.) punctimanus (Creaser, 1933:1). Quadruncus Group (terminal elements of pleopod subequal in length, central projec- tion about 33% total length of pleopod and tapering from base to tip, mesial process spatulate and excavated cephalically in dis- tal third with small spinose projection in distal fourth of caudal margin, both ele- ments inclined caudally in distal half; ce- phalic shoulder absent; annulus about as wide as long with caudal margin projected into tongue-like protrusion which over- hangs following sclerite, cephalolateral prominences and sulcus weakly developed): Orconectes (Procericambarus) quadruncus (Creaser, 1933:10). Spinosus Group (terminal elements of pleopod of unequal length, central projec- tion 40-48% total length of pleopod, both elements subsetiform; cephalic shoulder present—absent only in O. williamsi; an- nulus at least as long as wide, caudal margin rounded and if projected only slightly over- hanging following sclerite): Orconectes_ (Procericambarus) luteus (Creaser, 1933:7). O. (P.) macrus Williams, 1952:337. O. (P.) medius (Faxon, 1884:121). O. (P.) menae (Creaser, 1933:5). O. (P.) nana Williams, 1952:333. O. (P.) ozarkae Williams, 1952:339. O. (P.) putnami (Faxon, 1884:131). O. (P.) saxatilis Bouchard and Bouchard, 1976a:439. O. (P.) spinosus (Bundy, 1877:175). O. (P.) transfuga Fitzpatrick, 1966a. O. (P.) williamsi Fitzpatrick, 1966b. Gender. — Masculine. Etymology. — Procerus (Latin = tall, thin) combined with the generic name, Camba- rus, an allusion to the long, thin terminal elements characteristic of most members of this subgenus. Rhoadesius, new subgenus Figs. 10, 14b Diagnosis. —(Based on first form male and female.) Body and eyes pigmented, latter well developed. Rostrum with marginal VOLUME 100, NUMBER 1 59 Figure 11 Figs. 10, 11. Type species of Orconectes subgenera (all not to same scale): 10, Orconectes (Rhoadesius) sloanii; 11, O. (Tragulicambarus) lancifer. a, Dorsal view of carapace; b, Mesial view of first pleopod of first form male; c, Lateral view of first pleopod of first form male; d, Annulus ventralis of female; e, Lateral view of first pleopod of second form male; f, Dorsal view of chela of first form male. spines, median carina absent. Cervical spines well developed; areola 5.5—6.5 times longer than wide with 3—4 punctations across narrowest part, and constituting 32-34% to- tal length of carapace; cephaloventral sur- face of carapace with small squamous tu- bercles; devoid of spines or tubercles in hepatic region. First pleopod of male ending in 2 short (less than 18% total length of pleopod) terminal elements of unequal length, elements subparallel (O. sloanii) or divergent (O. kentuckiensis), mesial process stout and tapering from base to tip; distal half of pleopod inclined caudodistally at an- gle of about 30° to main axis of basal por- tion; cephalic surface of pleopod with (O. sloanii) or without (O. kentuckiensis) shoul- der. Inner margin of hand about 29% length of outer margin; opposable margin of im- movable finger and dactyl with (O. sloanii) or without (O. kentuckiensis) prominent tu- bercles, never with one more strongly de- veloped than others; tuft of setae at base of immovable finger, if present, never well de- 60 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON veloped. Annulus ventralis of female im- movable, firmly attached to preceding scler- ite; widest near midlength; cephalolateral prominences developed only in O. sloanii and trough not always clearly demonstrat- ed: sulcus usually present but often not well defined; sinus arising near midpoint of an- nulus, in well defined fossa only in O. ken- tuckiensis, and following simple undulating path to be lost before reaching caudal mar- gin. Type species. —Cambarus sloanii Bundy, 1876:24. List of species. — Orconectes (Rhoadesius) kentuckiensis Rhoades, 1944:122. O. (R.) sloanii (Bundy, 1876:24). Gender. — Masculine. Etymology. —Named in honor of the late Rendell Rhoades in recognition of his con- tributions to our knowledge of crawfishes. Tragulicambarus, new subgenus Figs. 11, 15b Orconectes: Creaser, 1962:3 (part), 6 (Fig. 15), 7 (part). Diagnosis. — Based on first form male and female.) Body and eyes pigmented, latter well developed. Rostrum lacking marginal spines but with sharply angular shoulders delimiting base of very long (about 50% of total length of rostrum) acumen, median ca- rina absent. Cervical spines well developed; areola obliterated and constituting about 31% total length of carapace; cephalolateral surface of carapace with small squamous tubercles; devoid of spines or tubercles in hepatic region. First pleopod of male ending in 2 short (about 18% of total length of pleo- pod) terminal elements of subequal length; central projection subparallel to main axis of pleopod and laterally compressed into blade-like structure; mesial process stout, tapering from base to tip and divergent from central projection throughout length; ce- phalic surface of pleopod with well defined shoulder near base of central projection. In- ner margin of hand about 45% length of outer margin; opposable margin of immov- able finger and of dactyl without prominent tubercles; margins of fingers setose but tuft of setae at base of immovable finger lacking. Annulus ventralis of female immovable, firmly attached to preceding sclerite; widest near midlength and about as long as wide; cephalolateral prominences well developed and separated by well defined trough; sulcus deep but only unilaterally developed; sinus arising in distinct fossa in sulcus and lateral to midline of annulus, moving caudome- sially in gentle arc to midline, there turning caudally to traverse slightly undulant path to intersect caudal margin. Type species.—Cambarus lancifer Ha- gen, 1870:59. List of species. —Monotypic, Orconectes (Tragulicambarus) lancifer (Hagen, 1870: 59). Gender. — Masculine. Etymology. — Tragula (Latin = a javelin) combined with the generic name, Camba- rus, a reference to the trivial name of the only species. Phylogenetic Considerations Fig. 18 As outlined above, the similarities of taxonomic characters in Orconectes make it difficult to apply cladistic techniques rigidly when considering phylogeny. Nevertheless, one can make some outgroup and ingroup comparisons to identify probable apomor- phies and plesiomorphies. The latter com- parisons are sometimes confusing, because ecological channelization has led to con- vergent emergences of certain characteris- tics. Perhaps the clearest plesiomorphy is mul- tiple terminal elements. Hobbs has con- vincingly argued for a procambarid-like ancestor for the family Cambaridae (1958, 1967, 1969, 1976, 1981; Hobbs and Barr 1960, 1972). Implicit in these discussions is an ancestral pleopod of four elements, from which certain lineages can be shown 61 VOLUME 100, NUMBER 1 ee SE — ‘ht az # Figure 19 Figure 14 Figure 13 q ) Figs. 12-14. Distribution of Orconectes subgenera: 12, Subgenus Gremicambarus (horizontal rulings = con- tribution of virilis-nais complex; vertical rulings = other species of the subgenus); 13, Subgenera Buannulifictus and Crockerinus (a, horizontal rulings = Buannulifictus; b, stippling = Crockerinus); 14, Subgenera Billecambarus and Rhoadesius (a, stippling = Billecambarus; b, horizontal rulings = Rhoadesius). 62 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON t iy p 5 < cs e Sas q Fs ~ Figure 15 y Figure 17 Figs. 15-17. Distribution of Orconectes subgenera: 15, Subgenera Faxonius and Tragulicambarus (a, vertical rulings = 7ragulicambarus; b, stippling = Faxonius); 16, Subgenera Hespericambarus and Orconectes (a, vertical rulings = Hespericambarus; b, stippling = Orconectes); 17, Subgenus Procericambarus (arrows represent probable or actual introductions where large populations have been established, probably at the expense of native species). VOLUME 100, NUMBER 1 to lose specific elements. Also, in general, these elements are fundamentally short in the plesiomorphic state. Identification of the first clade.—The re- tention of short (comparatively) terminal elements by members of the subgenus Or- conectes is the more primitive form. This conclusion is reinforced by the presence of a third element (the caudal process) in O. (O.) australis australis and in O. (O.) in- comptus and the occasional presence of a vestigial caudal process in O. (O.) australis packardi (in some specimens even a vesti- gial cephalic process can be recognized: Hobbs and Barr 1972:31, fig. 8c, d). Hooks (sometimes rudimentary) on the fourth pe- reiopods of specimens of O. (O.) australis subspp., O. (O.) inermis subspp., and O. (O.) pellucidus further serve to place the subgenus close to the stem population from which the genus descended. (Only in rare specimens of other subgenera of Orconectes are hooks found on pereiopods other than the third.) Other features attributed to the procam- barid ancestor also present in the nominate subgenus include spinose ornamentation of the hepatic region of the carapace and along the cephalic part of the cervical groove, a short broad areola, and a movable (albeit slightly) annulus ventralis in the females. Equally significant is the presence of these apparently most primitive members of the genus in a geographical area which Hobbs (most recently, 1984) considers the center of diversity for the Cambaridae; members of the subgenus are troglobites in the karst along the southeastern edge of the Cum- berland Plateau. Thus, by outgroup com- parisons one is able to establish reasonably reliable plesiomorphic character states for the genus, identify the group retaining the greatest number of plesiomorphic states, and postulate probable lineages through which the several subgenera were established. Un- fortunately, of these, only the terminal ele- ments of the male pleopod and the annulus ventralis of the female are not subject to influence by the habitat. And the most strik- 63 Buannulifictus Gremicambarus Rhoadesius Tragulicambarus Procericambarus Crockerinus Orconectes Hespericambarus Faxonius ? WIN JT | procambarid ancestor Fig. 18. Cladogram expressing proposed relation- ships of subgenera of Orconectes. ing feature of the annulus in the subgenus Orconectes, its motility, 1s lost in all other subgenera. Enough plesiomorphies do exist, how- ever, to permit evaluations of other char- acter states in the members of the nominate subgenus and to compare them with mem- bers of Procambarus which Hobbs (1984) has indicated contain many familial ple- siomorphies: the subgenera Pennides and Ortmannicus (particularly the Pictus Group). From these comparisons, one can project additional probable synapomor- phies for the genus. In those species with the greatest number of plesiomorphies (O. australis subspp. and O. incomptus) a shoulder is present at the cephalic base of the central projection. A similarly situated or somewhat proximally displaced irregularity (in varying degrees of development) can be found in some mem- bers of Pennides (P. ablusus, P. lylei, P. natchitochae, P. versutus) and some mem- bers of the Pictus Group (P. enoplosternum, 64 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON P. hirsutus, P. pictus). Fitzpatrick (1967) postulated that the shoulder was plesio- morphic for members now assigned to the subgenus Crockerinus, which members likewise exhibit few character states which I consider apomorphic. Thus, a cephalic shoulder would seem to be plesiomorphic, but not its degree of development. The annuli ventrales of members of Pen- nides and the Pictus Group of Ortmannicus, although not as simply constructed as the Orconectes in question, are nevertheless comparatively weakly sculptured as a group. Further, females of the genus Cambaroides (subfamily Cambaroidinae) have only a simple, unsculptured annular plate. It is not, therefore, excessive to consider more elab- orate sculpturing of this structure to be apo- morphic. In Orconectes one can find no consistent pattern of cheliped development which can be used to establish lineages. This feature was extremely useful to Hobbs (1969) when he examined Cambarus. Equally, the car- apaces of Orconectes species do not afford a means to evaluate apomorphic trends. In most carapace characteristics, the members of the genus are remarkably similar. If one accepts the postulates I have of- fered, then some decisions about lineages can be made. In members of the subgenus Orconectes the annulus ventralis is mov- able, nearly planar and, significantly, lack- ing any structures which can be identified unequivocally as vestiges or precursors of a trough or sulcus. The central projection and mesial process are both short; the former is often flattened in the cephalocaudal plane and the mesial process is usually robust al- though tapering from base to tip. When compared with these character states and with members of other genera of the Cam- baridae, long subsetiform elements, blade- like central projections, and spatulate, cephalically excavated mesial processes be- come the apomorphic condition. The first recognizable apomorphies seem to be a firm commitment in the gonopod to only two terminal elements (the central pro- jection and the mesial process), their elon- gation, and the introduction of orconectid (for lack of a better term) sculpturing to the annulus ventralis. Shortly after these spe- cializations become established, in group II of my cladogram (that leading to Crocker- inus, Faxonius, and Procericambarus), the main shaft of the pleopod became more cy- lindrical, principally through the reduction of the caudal expansion of that area situated caudolateral to the proximal opening of the sperm groove. At about the same time, de- velopment of both a trough and sulcus oc- curred on the annulus, and the mesial pro- cess of the male gonopod became subsetiform. In group III there was no appreciable re- duction of the caudal expansion of the gon- opod, it remained rather prominent; the mesial process tended toward attenuation but was not subsetiform. In females, the trough and sulcus were probably very weak- ly developed and variable. Trends leading to Faxonius.— Returning to group II, further developments can be recognized. The divergent apices of the ter- minal elements of the gonopod were re- tained by some, as were the plesiomorphic carapace spines (evidenced by their reten- tion in O. limosus), and a clearly recogniz- able trough can be seen; only in O. wrighti is the sulcus obscure, but in all of them the cephalolateral prominences are easily iden- tifiable, and the sinus arises in a barely rec- ognizable fossa near the midwidth of the sulcus. The populations diversified into the members of the subgenus Faxonius. Other populations of group II took another tack (leading to Crockerinus and Proceri- cambarus). The central projection also ap- proached a subsetiform state, and the straight elements moved into a subparallel relationship. A shoulder was probably pres- ent on the pleopod (Fitzpatrick 1967:167), and both trough and sulcus were clearly ev- ident on the annulus. Trends leading to Crockerinus.— Of these, VOLUME 100, NUMBER 1 those that developed into members of the subgenus Crockerinus retained a relatively low degree of relief on the surface of the annulus; the sinus arose from a moderately developed fossa which was located near the midline of the annulus in the cephalic por- tion of the sulcus. From there, the sinus usually extended laterad or caudolaterad before recurving sharply to the midline (the unique annulus of O. bisectus is an excep- tion); and from there it followed a gently sinuous path nearly to the caudal margin, only rarely (in O. erichsonianus and O. shoupi) intersecting it. A fundamentally broadly spindled shape was retained throughout. The male gonopods remained in a more conservative state. They were longer than the plesiomorphic condition found in mem- bers of the subgenus Orconectes and longer than in members of the subgenus Faxonius but somewhat shorter than the conditions found in members of the subgenus Procer- icambarus. Both the mesial process and central projection became less stout, but they remained subequal in length or nearly so. The central projection never reached a state of being subsetiform. The cephalic shoulder was lost in most populations, but individ- uals in many retained it, albeit usually in a reduced form. Trends leading to Procericambarus.—In members of Procericambarus the annulus developed large, conspicuous cephalolateral prominences and a deep, distinct trough, often overhung in its cephalic parts by the cephalolateral prominences; in general, the annuli of members of Procericambarus are the most prominently sculptured in the ge- nus. The strong tongue-like projection of the caudal margin of the annulus in members of the Hylas Group also represents an ex- treme for the genus. The terminal elements of the male gonopod became much elon- gated, especially the central projection. The tip of the elements reach the coxae of the second pereiopod when the abdomen is flexed, and in some species they reach as far 65 as the mouthparts. Undoubtedly, in pleo- pods one can say that the most divergent state of the genus is reached. The subgenus seems to represent the most modified, with respect to structures used in amplexus, sit- uation in the genus, and indeed its members may well be the most specialized members of the family in this respect. Trends leading to Tragulicambarus.— Returning to the initial dichotomy of the two major groups, an enigma, O. /ancifer, is encountered. In many ways it is unlike any other Orconectes. The acumen usually accounts for at least half of the total length of the rostrum, although I have seen many specimens in which the acumen approached more typical proportions. The hand is long and narrow, the inner margin of the palm being clearly longer than the dactyl. The central projection is laterally compressed and blade-like, a characteristic shared only with O. bisectus. But all of these, except the pleopod, probably represent specializations rather than significant apomorphies. The hand is very reminiscent of Procambarus (Capillicambarus) spp. and less so of Fax- onella spp. All of these species inhabit road- side ditches or other such semipermanent standing waters. The hand would seem to be more indicative of habitat than ancestry. Never reaching the extreme of /ancifer, the acumena of several apparently distantly re- lated species of Orconectes can be quite long: O. (O.) inermis, O. (C.) virginiensis, O. (P.) longidigitus. Likewise, outgroup represen- tatives sometimes have very long acumena: Cambarellus (Cs.) prolixus, Procambarus (Ortmannicus) youngi, P. (Pennides) ablu- sus, P. (Pe.) lylei, P. (Pe.) lagniappe. Such distribution discourages the use of the acu- men to determine relationships. If the general morphology of the pleopod of /ancifer is compared with what I have taken to be the plesiomorphic condition, one notices that the reduction of the area caudolateral to the proximal opening of the sperm groove is not reduced to the degree found in members of Crockerinus, Faxo- 66 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON nius, and Procericambarus. In this respect, the pleopod is more like that of the lineage of group III of the major dichotomy than of group II. Similarly, the length of the ceph- alocaudal axis of the annulus is nearly equal to that of the transverse axis, again more like the condition obtaining in the group III stem. Discounting the apparently environ- mentally influenced features, Tragulicam- barus seems to have arisen as a unique di- vergence from the group III stock, but its precise relationship remains unclear. Less tenuous are the lineages of what I perceive to be the principal line of descent in the group III arm of the dichotomy. The greatest diameter of the gonopod remained located just distal to the proximal opening of the sperm groove, and there it retained close to twice the diameter at the base of the terminal elements. In the terminal ele- ments, especially the mesial process, a marked tendency to have their apices ori- ented caudodistad or caudad developed. The length of the cephalocaudal axis of the an- nulus approached or slightly exceeded that of the transverse axis. The development of cephalolateral prominences encouraged the appearance of distinct troughs and sulci. Trends leading to Hespericambarus.— The species which digressed least from the postulated ancestral type of group III are assigned to the subgenus Hespericambarus. In these animals, the terminal elements re- tained a relatively stout condition, were still relatively short and had minimal modifi- cation of their basic shape. The annuli were simple, and in none was the combination of a clearly defined trough and sulcus pres- ent. Only in O. difficilis (and possibly O. maletae) is a fossa present, and the sinus, although more conspicuous than that in the subgenus Orconectes, is not elaborate. Moreover, only in O. difficilis does the cephalocaudal elongation of the annulus be- come such that the structure can be de- scribed as subrhomboid. Orconectes hath- awayi seems to be the extant species which can be described as “‘closer to the ancestral type.” It has the straightest and shortest ter- minal elements in the subgenus; but also pertinent is the fact that many individuals bear spines (although most rather small) in the hepatic region of the carapace, their presence earlier herein considered a plesio- morphic character state. As a final observation, I should comment on the nomenclature employed for mem- bers of this subgenus. Lacking contrary evi- dence, I would retain the nomenclatorial combinations proposed by Walls (1972), despite that fact that I cannot comprehend the intergradations of populations visual- ized by him. But for reasons implicit in my recognizing two “Groups” within the sub- genus, I am listing each taxon as a distinct species and deferring determination of the precise limits of variations for the several populations until a later time. Trends leading to Rhoadesius.— Proba- bly closely related to but nonetheless dis- tinct from Hespericambarus is the small group of crawfishes assigned to the subgenus Rhoadesius. These differ from members of the former principally in that the distal half of the pleopod is inclined caudally to the main axis of the appendage (as established by the basal part). The annuli are relatively simple, but in O. kentuckiensis a fossa 1s present, and in O. sloanii ridge-like ceph- alolateral prominences suggest a tendency to sulcus formation. Both species have a moderately broad areola in contrast to the obliterated one in members of Hespericam- barus, yet this character is unreliable for determining relationships. Species, in any genus, which inhabit cool, tumbling waters tend to have a short, broad areola; in con- trast, species living in sluggish, warm, standing waters and burrowers most likely will have a long and obliterated areola; species living in intermediate type habitats usually exhibit intermediate characteristics of this structure. Trends leading to Billecambarus.— Per- haps the most confusing of the members of the genus is O. (Billecambarus) harrisoni. VOLUME 100, NUMBER 1 In some respects the pleopod resembles that of members of Rhoadesius, but the short terminal elements are curved throughout their length to a degree that, coupled with caudal inclination of the distal half of the pleopod, the apices of both are directed due caudad. The annulus is unique in the genus. Only in O. (R.) kentuckiensis are the ceph- alolateral regions so undeveloped that the anterior half of the annulus is essentially a deep transverse excavation similar to the condition in O. (Bi.) harrisoni. In the latter, this transverse sulcus (?) is overhung cau- dally by a median projection of the more elevated caudal half; the sinus originates nearly on the cephalolateral margin and runs obliquely in a deep groove to the midline before turning caudally to move in a scarce- ly arched path to the caudal margin. The species seems to be a digressive, trans-Mis- sissipp1 offshoot of Rhoadesius. Trends leading to Gremicambarus.—In members of the subgenus Gremicambarus the central projection is straight in its basal part, although the distal parts of the pleo- pods are frequently inclined caudally with respect to the main axis of the basal half of the appendage. The mesial process is like- wise usually straight basally. The propor- tions of the annulus approach those of Crockerinus, but the structures differ con- spicuously from those of the latter in that well defined sulci are present and, except in O. (G.) validus, equally well defined troughs are also present. The sinus originates in a deep fossa, usually in the cephalolateral por- tion of the sulcus and extends laterally be- fore recurving sharply to the midline; this produces a prominent, tongue-like, laterally oriented ridge which descends toward the lateral parts of the sinus, which latter is fre- quently partly obscured by an overhang of the cephalolateral prominence and/or lat- eral extension thereof. These are probably the most complexly organized annuli in the genus. Further evidence of the digressive nature of the members of this subgenus can be 67 found in the nearly subcylindrical main shaft of the pleopod. Members of the Alabamen- sis Group are extreme in having a spatulate mesial process with a conspicuous groove along the cephalic face. The hands, although usually not useful for determining relation- ships, help tie the species together into a cohesive group. In all (except O. compressus and O. chickasawae) a tuft of setae, of vary- ing degrees of development, occurs at the base of the opposable margin of the fixed finger; in this respect they resemble mem- bers of Crockerinus. But, most unusual in the genus, fully half of the species have at least one tubercle on the opposable margin of the fixed finger which stands out from the rest in size. Another unusual characteristic occurs in the basal third of the opposable margin of the dactyl (absent only in O. com- pressus, O. cooperi and O. rhoadesi): a broad concavity with (except in O. immunis) more than one prominent tubercle, structured not unlike that seen in Cambarus (Lacunicam- barus) spp. Trends leading to Buannulifictus. — More divergent still are members of the subgenus Buannulifictus. Except in O. meeki meeki, the distal portion of the pleopod is inclined caudally with respect to the main axis of the proximal part of the pleopod, and the cen- tral projection is curved throughout its length, the apex (except in the same sub- species) directed more or less caudally. The subsetiform mesial process is likewise curved but also from its base takes a path divergent from that of the central projec- tion, the apices of the terminal elements being at least twice as far apart as the bases. One of the most significant changes in the annulus ventralis was an increase in the cephalocaudal axis. The development of the cephalolateral prominences was more to- ward a ridge-like oval than toward a circular hillock, a situation shared with Gremicam- barus. Not surprisingly, the annuli are su- perficially similar in the two subgenera. But in Buannulifictus the lateral development of the prominences is more intrusive on the 68 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON middle of the annulus, resulting in a sulcus which is more pit-like than ditch-like. At the same time, the mediocephalic incur- sions of the prominences render the trough poorly evidenced, at best. The sinus origi- nates in a deep fossa near the caudal margin of the cephalolateral prominences, and its peregrinations are not dissimilar to those described for Gremicambarus, although a tongue-like ridge as described for the latter subgenus is clearly defined only in O. (B.) palmeri palmeri. Except for O. (B.) palmeri longimanus, the dactyl concavity typical of Gremicam- barus is absent, but a tuft of setae at the base of the fixed finger is present, albeit poorly developed, in all but the two sub- species of O. meeki. The opposable margins of the fingers usually have well developed tubercles, but no prominently large one is characteristically present. The two subgen- era are bound by many commonalities, some of which are apparently synapomorphic. But an equal number of differences exist also. One is led to the conclusion that although their members are rather closely related, they constitute two distinct species groupings, here recognized as subgenera. Geographic and Temporal Interpretations In such speculations as have been pre- sented in preceding paragraphs one invari- ably is led to postulate temporal assign- ments for proposed events. At the moment I am more prepared to examine geographic relationships than paleontologic ones. Hobbs and Barr (1972) offered explanations of the time and place of the origin of Or- conectes. They also similarly addressed the early development and migration of the ear- ly isolates of the parental population. They accepted the pre-Miocene origin of the ge- nus, located geographically at the south- eastern rim of the Cumberland Plateau, pro- posed by Hobbs (1969), as well as his Suggestion that the expansion was essen- tially westward. Their principal addition was to offer a “Tertiary” northward migration of early populations, some of which gained access to the Atlantic drainages and per- sisted as O. (F.) limosus. They also provid- ed strong arguments that the genus descend- ed from stream dwellers. Fitzpatrick (1967) associated emergence of Crockerinus and Procericambarus with (by inference) the early Quarternary and en- visioned subsequent speciation of the for- mer as a result of conditions existing during Illinoian to post-Wisconsin times. His ex- planations were more compatible with Rhoades’ (1962) proposals for events ef- fecting the distribution of northern craw- fishes than were those of Hobbs and Barr (1972). Indeed, they rejected outright Rhoades’ mechanism to explain the distri- bution of O. limosus. If I were inclined to disagree with any of the above, I could find no concrete data to refute Hobbs’ and Barr’s hypotheses. Re- cently, however, Fitzpatrick (1983) used new geologic information to speculate that a prominent pre-Pleistocene river drained the upper Tennessee into the Florida Parishes of Louisiana and argued that this river could account for the eastern distribution patterns of the Cambarellinae. He carried this pro- posal further (Fitzpatrick 1986), giving more details, in using such a drainage to account for many peculiarities of crawfish distri- bution in the eastern part of the Gulf Coastal Plain. In this scheme, the proposed “‘ances- tral home’’ of Orconectes would be related to this river. It seems more than coinciden- tal that the expansion of Orconectes is es- sentially west of this river and that of the early digressives of Cambarus (Hobbs 1969) is to the east. If one uses this proposed river to localize the eastern boundary of the eastcentral mar- gin of the Mississippi Embayment, one can begin to speculate about the dispersal routes (corridors in the sense of Hobbs 1969, 1984). One early group moved north through east- ern Tennessee and West Virginia, possibly through the New River system, and gained VOLUME 100, NUMBER 1 access into what is now the upper Ohio drainage. From Hobbs’ and Barr’s (1972) timetable this would be definitely pre-TIlli- noian and probably Tertiary time. These populations became the members of Fax- onius. Another “‘wave’’ of invasion, out of the same Cumberland source area followed, to diversify into today’s Crockerinus. This more progressive descendent of the ances- tral stock would have obliterated any extant epigean remnant of Orconectes (subgenus) and was able to bisect the range of Faxonius to leave the probably already isolated /i- mosus precursors in the Atlantic drainages and compressing the remainder into three isolated populations near the periphery of a range that at once encompassed the area from the Appalachian divide to the eastcen- tral rim of the Mississippi embayment and south of the Teays system. By Fitzpatrick’s (1967) reckoning this would have occurred before the Illinoian glaciation. West of what is now the lower reaches of the Tennessee River another stock was es- tablishing itself. This was the precursor of my proposed “Group III” (Fig. 18) lineage (Billecambarus, Buannulifictus, Gremicam- barus, Hespericambarus, Rhoadesius, and Tragulicambarus). These crawfishes seem to have invaded the northern (lower) reach- es of this drainage as several “‘waves’’; pos- sibly these are associated with the migra- tions of the ice sheets of the Pleistocene. One group, however, seems to have become established west of the Mississippi River comparatively early. Exactly how this was accomplished is highly problematical, but surely considerable opportunities exist and existed for animals not too demanding of the stream environment to cross as mean- ders of the mid-Mississippi channel occur. Apparently the ancestors of members of Hespericambarus settled in the Tertiary or early Quarternary streams of northwest Louisiana/Arkansas/Oklahoma and invad- ed the Quarternary lands as they became available. The present-day representatives east of the river are O. (H.) perfectus and 69 populations very close to it. They, in turn, are almost indistinguishable from Louisi- ana’s O. (H.) hathawayi, and I interpret them to be relatively recent immigrants into the Tombigbee drainage system. Possibly coincident with these activities, another group was moving up (down by present directions) the streams into the Ohio basin. These diversified into the members of Rhoadesius and were more widespread than today. For whatever reason—climatic change, replacement by more efficient com- petitors, or both—the original range was bi- sected and remains today as two, essentially relict, allopatric areas. Two monotypic, enigmataic subgenera attract our attention next. Orconectes (Bil- lecambarus) harrisoni can be found only in a very restricted area just west of the Mis- sissippi River flood plain in Missouri and between the latitudes of confluence of the Missouri and Ohio rivers with the Missis- sippi. As discussed above, its morphology is very difficult to interpret, but it seems to be most closely related to Rhoadesius and could represent a very disparate western iso- late from that group of populations. Orco- nectes (Tragulicambarus) lancifer is like- wise morphologically unique and even more difficult to associate with another group of species. That an almost perfect correlation of its distribution with Quarternary-Holo- cene deposits exists is inescapable, however, and one must therefore assume a compar- atively late specialization enabling its al- most unique (for Orconectes) invasion of the habitats—essentially lentic, often stag- nant, or very sluggishly flowing and lacking firm substrates for much of their extent— associated with these regions. Becoming fully established in the lower (i.e., southern) reaches of the eastern leg (1.e., upper) of the Tennessee River were the pro- genitors of Procericambarus. This may have taken place in Early Quarternary times. This stock seems to have consisted of vigorous competitors; indeed, O. (P.) rusticus is one of the most successful displacers at the pres- 70 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ent time (Crocker and Barr 1968; Capelli 1975, 1982; Berrill 1978; Capelli and Mun- jal 1982; Tierney and Dunham 1984). They eliminated Crockerinus from its ancestral home on the Cumberland Plateau, drove O. (C.) erichsonianus southward and pushed the second, northern assemblage hard as they invaded the Ohio system. They spread ex- tensively, possibly along the Erigan system, going across the Mississippi to occupy the Missouri highlands and thence southward into the Ouachitas. They tumbled off the Highland Rim into the Nashville Basin to become firmly established in central Ten- nessee and Kentucky. How this was accom- plished will probably always remain un- known. It may have resulted from an invasion down the slopes from the Cum- berland Plateau; it may have been via a union of the two segments of the Tennessee River; or it may have been both. Until the approximate time of this fusion to form the present Tennessee River is determined, fur- ther speculation seems useless. Man has helped O. (P.) rusticus in its invasions, but much of the range represents its own vig- orous and successful expansion into areas breaking free of ice cover. The lower (southern) parts of the western (lower) leg of the Tennessee drainage saw the nearly simultaneous establishment of ancestors to Gremicambarus. Most species initially probably had difficulties expanding their range northward as they encountered the vigorous Procericambarus populations and were forced to content themselves with central Tennessee and the emerging lands which are now associated with the Tom- bigbee River drainage. To the west and southwest of them Buan- nulifictus fauna was laying claim to most of Mississippi and the southern parts of the west bank of the river. Little conjecture can be made concerning this stock, for present drainage patterns in the critical areas do not well reflect the history of the region. There are extensive “‘drowned”’ drainages in northern Mississippi (Murphey and Gris- singer 1981), and serious questions of the age and sources of deposits throughout the area have been raised (May 1981, Isphord- ing 1983). Becoming teleological, one could say that Gremicambarus “‘bided its time” and ‘“‘worked to build a better mousetrap.” By the retreat of the last ice sheet, two species of the subgenus were poised “to hold their own’ as the north was exposed for coloni- zation. Surely the subgenus dominates the crawfish fauna of the central part of North America. But an examination of the ranges of the members of the subgenus leaves no doubt that the overwhelming majority of the total range is ococupied by O. (G.) im- munis and O. (G.) virilis, the latter exceeded in range only by Procambarus (Ortmanni- cus) acutus subspp. The several populations of O. (G.) virilis, O. (G.) nais and O. (G.) causeyi are morphologically nearly indistin- guishable, although Pryor and Leone (1952) reported serological differences between O. nais and O. virilis. The latter has to be eval- uated in light of Phillips’ report of possible intergrade populations in southwestern Iowa (1980); this is one of the few studies in which a detailed examination for just such a sit- uation has been reported. Regardless, the Gremicambarus invaders attempting to move out of Missouri River drainages into southern Missouri and Kansas-Oklahoma were stymied in their southern and south- western migration by well-established Buannulifictus populations and in their westward and northern movements by in- tolerable or inaccessable habitats resulting from climatic conditions. Nevertheless, they constitute the most widespread subgenus of Orconectes and are obviously successful competitors. In summary, analysis reveals a reason- ably rational division of the genus Orco- nectes into 10 subgenera which seem to re- flect historical events in the diversification of the genus. Adequate data determining VOLUME 100, NUMBER 1 precise relationships are lacking, but gen- eralities implied by the proposed groupings are supported by the information at hand. Acknowledgments Many people have contributed to this study, but Horton H. Hobbs, Jr., deserves special recognition. 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WASH. 100(1), 1987, pp. 75-88 CHONO ANGUSTIARUM, A NEW GENUS AND SPECIES OF ZOBRACHOIDAE (CRUSTACEA: AMPHIPODA) FROM MAGELLAN STRAIT, WITH A REVISION OF UROHAUSTORIIDAE Janice Clark and J. L. Barnard Abstract. —The new genus and species, Chono angustiarum, from Magellan Strait, has characters of both Zobrachoidae and Urohaustoriidae. The Zobra- choidae are redefined to include genera with well developed epimeron 1, in contrast to Urohaustoriidae containing genera with vestigial epimeron 1. The previously described Magellanic genera Huarpe and Tonocote also represent intergrades between the two families as originally described from Australia where they are very distinctive. Chono angustiarum, a new genus and species of Zobrachoidae is described from Magellan Strait. This marine fossorial am- phipod almost perfectly intergrades between the Urohaustoriidae and Zobrachoidae by combining a well developed epimeron | with a simple gnathopod 1. Overlaps between the two families, so distinct in Australian waters, have already been described in Magellan genera such as Huarpe Barnard and Clark (1982) and Yonocote Clark and Barnard (1986). We are not prepared for final fusion of the two families because the genera re- main distinctive on the basis of epimeron 1. In our redefinition the Zobrachoidae re- tain the plesimorphic state of epimeron 1 in fully developed condition, whereas Uro- haustoriidae have epimeron | vestigial or absent. The distinction remains strong. Structure of gnathopods is thereby dis- counted. Methods of analysis follow Barnard and Drummond (1982). In the text, symbols “‘E”’ and “e”’ = large and small setae, “S” and **s” = large and small spines, “‘p’’ = peni- cillate seta, ““R”’ = right sided appendage as contrasted to normal description based on left. Discussion When originally described from Australia (Barnard and Drummond 1982), the fam- ilies Zobrachoidae (then three genera) and Urohaustoriidae (then six genera) were dis- tinguished by the apomorphic characters of Urohaustorudae as follows: simple gnatho- pod 1, strongly reduced coxa 2 in compar- ison to coxa 3, adze-shaped coxa 3 with large posteroventral projection, great re- duction of epimeron |, smallness of rostrum on head, small number of setae on inner plate of maxilla 1, more strongly extended incisor with reduced development of cusps, greatly reduced number of rakers, and gen- erally the reduction in size of article 2 of outer ramus on uropod 3; to this could be added in retrospect the development of awns on the setae of the mandibular palp and the loss of lateral setae on the peduncle of uro- pod 2. Table 1 shows the overlaps between the old family extremes occurring in Huarpe, Tonocote, and Chono. Characters of the table are stated in such a way that Zobrachoidae reflect (+) and Urohaustoridae (0). The intermediate gen- era are placed in transformational order re- 76 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 1.—Characters of Zobrachoidae and Urohaustoriidae from Australia compared to transitional Magel- lanic genera. Intermediate genera are arranged in their closest affinities. Asterisks denote strongest anomalies in this particular arrangement. Urohaus- Zobra- Character __torlidae Chono Huarpe Tonocote choidae l 0 0 sr de + Coxa 3 not adze-shaped 2, 0 0 + ae Se Coxa 2 not very small 3 0) 0 0 aF + Rakers 4+ 4 0 ap 0 +