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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
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PROC. BIOL. SOC. 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.
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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-
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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
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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
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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-
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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,
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Ockelmann, K. W., and K. Muus. 1978. The biology,
ecology and behaviour of the bivalve Mysella
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tions of new species of Ceratonereis, Nephtys,
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and Goniada (Polychaeta).— Proceedings of the
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Pettibone, M. H. 1963. Marine polychaete worms of
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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. He has been generous
with his time and knowledge since he first
directed my attention to Orconectes. Sub-
sequently, he has made every effort to ease
my path toward an understanding of these
creatures and crawfishes in general. Ray-
mond W. Bouchard, Horton H. Hobbs III,
David Stansbery and their students have
given many helpful insights during numer-
ous extended conversations. And not least,
personnel of the U.S. National Museum of
Natural History consistently worked to
make my visits there profitable. To each of
these I offer sincere thanks.
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1890. Notes on North American crayfishes,
family Astacidae.— Proceedings of the United
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1898. Observations on the Astacidae in the
United States National Museum and in the Mu-
seum of Comparative Zoology, with descrip-
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1914. Notes on the crayfishes in the United
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Fitzpatrick, J. F., Jr. 1963. Geographic variation in
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1965. A new subspecies of the crawfish Or-
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. 1966a. Anewcrawfish of the subfamily Cam-
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1966b. A new crawfish of the genus Orco-
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. 1967. The Propinquus Group of the crawfish
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, and J. F. Payne. 1968. A new genus and
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Girard, C. 1852. A revision of the North American
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Hagen, H. 1870. A monograph of the North Amer-
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Hay, W. P. 1891. The Crustacea of Indiana.—Pro-
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Hobbs, H. H., Jr. 1942. A generic revision of the
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1948b. Two new crayfishes of the genus Or-
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American Midland Naturalist 39:139-150.
. 1948c. Anewcrayfish of the genus Orconectes
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VOLUME 100, NUMBER 1
. 1949. A newcrayfish of the genus Orconectes
from the Nashville Basin in Tennessee, with
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1958. The evolutionary history of the Pictus
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1967. A new crayfish from Alabama caves
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73
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74 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
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Department of Biological Sciences, Uni-
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PROC. BIOL. SOC. 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 +
Le
\\
\
, ——— = _— 5
RK in |
Say — te
eae
——<
oS
aNS
AEN LE
AE |
= a} jE2 a
i Were ees
Mi ar
\ a |
H
e
Fig. 3. Cypridina americana, adult male, USNM 159080: a, Left 1st antenna, medial view; b, Endopodite,
distal part of protopodite, and proximal part of Ist joint of exopodite of right 2nd antenna, medial view; c,
Right 6th limb, medial view; d, 7th limb; e, Upper lip, anterior to right (2 posterior tusks of left side not shown).
178 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
|
\
/
i ef \
/ =
ee BS : =. I :
2
yy nN
5>6>7.
Cephalon with very low rounded rostral
point; eyes small, pigmented, of about 10
ommatidia. Pleonites short, free, 1-4 sub-
equal, 5 slightly longer, 6 with posterior
margin broadly bilobed. Telson lacking
statocysts, lanceolate, widest at midlength,
lateral margins in posterior half with 6 ser-
rations; few setae on rounded apex.
Basal antennular peduncular article long-
est and broadest; flagellum of 3 articles, ar-
ticle 2 longest, with single aesthetasc, article
3 with 2 aesthetascs. Antennal peduncle with
article 2 longest and widest; articles 3 and
187
59 30
59 40
13 20
Six Men’s (14 m) ,
SSS OO
Greensleeves (11m) ,
North Bellairs (10 m) .
Bank Reef (15 m) @ 13 10
Payne’s Bay (5m) ef
° )
Brighton (9 m) Bridgetown
5 nautical miles
Fig. 1. Map of Barbados, West Indies, with sam-
pling sites and approximate depths.
4 together slightly shorter than article 5; fla-
gellum of 7 setose articles. Mandibular palp
of 3 articles; terminal article bearing 3 spines;
lamina dentata of 5 serrations; molar short,
truncate; incisor of 2 or 3 sclerotized cusps.
Maxilla with 6 distal teeth. Maxilliped with
palp of 5 articles, article 3 longest; broad
endite almost reaching base of palp article
4, distally acute.
Pereopod 1, propodus slightly expanded,
palm straight, bearing few simple setae; un-
guis 4 length of remainder of dactylus, with
short auxiliary spine at base. Pereopods 2
and 3 similar, propodi less expanded than
in pereopod 1, with 2 sensory spines on
palm; short strong auxiliary spine at base of
dactylar unguis. Pereopod 7, carpus with
short anterior margin, shorter than poste-
rior margin, small sensory spine at postero-
distal angle; propodus with posterodistal
sensory spine and 2 elongate distally fringed
anterodistal spines. Pleopod 1, exopod
operculiform, with 9 marginal plumose se-
tae; endopod about % length and % greatest
width of exopod, with 2 distal plumose se-
tae. Uropodal sympod with mesioventral
188 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
F
\
H
A
Y
|
Fig. 2. Chalixanthura lewisi, female: A, Adult in dorsal view; B, Telson; C, Antennule; D, Antenna; E,
Maxilla; F, Maxilliped; G, Mandible; H, Pleopod 1; I, Pleopod 2.
row of plumose setae; exopod ovate, api-
cally acute, with 2-3 distal serrations on
outer margin and several elongate setae; en-
dopod equal in length to basis, distally
broadly rounded, distolateral margin ser-
rate, bearing elongate simple setae.
Male: Body about 13 x longer than wide.
Proportions:C > 1=2=3<4>5=6>
7. Cephalon with low rostral point. Eyes
enlarged, of about 30 ommatidia, almost
contiguous middorsally, extending ventral-
ly and leaving narrow median strip in which
VOLUME 100, NUMBER 1
189
Fig. 3. Chalixanthura lewisi, female: A, Uropodal sympod and endopod; B, Pereopod 1; C, Pereopod 2; D,
Uropodal exopod; E, Pereopod 3; F, Pereopod 7.
mandibular palp and maxilliped accom-
modated. Pleonites short, free, 1-4 sub-
equal, 5 slightly longer, 6 with posterior
margin middorsally bilobed. Telson lanceo-
late, posterior half of lateral margins with
8—9 serrations, several setae on rounded
apex.
Antennular flagellum of 11 articles, prox-
190
imal articles bearing numerous aesthetascs,
number of aesthetascs reduced distally. An-
tenna similar to female, flagellum of 6 or 7
setose articles. Mandibular palp of 3 arti-
cles, terminal article with 4 spines; body of
mandible reduced to narrow columnar
structure. Maxilliped segmentation ob-
scure, at least 3 distal articles, endite lack-
ing.
Pereopod 1, propodus with slightly con-
cave palm bearing 2 basally broadened
spines, mesial surface near palm with 7
spines; unguis 73 length of remainder of dac-
tylus, with auxiliary spine more slender than
in female. Pereopods 2 and 3 similar; pro-
podus more elongate and slender than in
female, with 2 (pereopod 2) or 3 (pereopod
3) sensory spines on palm. Pereopod 6, car-
pus with anterior margin shorter than pos-
terior, with posterodistal spine; propodus
with 3 serrate sensory spines on posterior
margin. Pereopod 7, carpus as in pereopod
6; propodus with strong serrate sensory spine
at posterodistal angle, and with 2 elongate
distally fringed spines anterodistally. Pleo-
pod 1, rami subequal in length, endopod
half width of exopod. Pleopod 2, copulatory
stylet elongate, slender, reaching by more
than half its length beyond distal margin of
ramus, articulating in proximal half of en-
dopod; latter with 5 distal plumose setae;
exopod broader and slightly longer than en-
dopod, with transverse suture at about mid-
length, 10 plumose setae on distal margin.
Pleopod 3, exopod longer and wider than
endopod with transverse suture. Uropodal
endopod longer than sympod, with serrate
distolateral margin, several elongate simple
setae distally; exopod narrowly lanceolate,
apically acute, outer margin with 5 serra-
tions and few elongate setae.
Remarks. —The following features place
this species in the group of genera appar-
ently related to Panathura: Short free ple-
onites, pleonite | exopod operculiform, tel-
son lacking statocysts, large apically acute
maxillipedal endite, maxillipedal palp of 5
articles, pleopods 2—5 witk biarticulate ex-
opods, pereopod 1 propodus only slightly
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
more inflated than in pereopods 2 and 3,
posterior pereopods with carpus having an-
terior margin shorter than posterior margin.
Of these genera, the present species most
closely resembles the monotypic Chali-
xanthura Kensley, 1984, in overall body
shape and especially in the modifications of
the male. The major differences lie in the
antennal and antennular flagella which have
more articles in the present species, and in
the maxillipedal endite of the female, which
is much shorter in C. scopulosa. The present
species also does not have the strongly in-
cised uropodal exopod seen in the female
of C. scopulosa. The maxillipedal endite
more closely resembles that of Expanathura
Wagele, 1981, another member of the
‘*“Panathura’”’ group of genera.
Etymology. —The species is named for Dr.
John B. Lewis, professor of Oceanography
at McGill University, who has contributed
more than anyone to the knowledge of the
marine fauna of Barbados.
Ejisothistos teri, new species
Figs. 6, 7
Material.—HOLOTYPE, NMNS NMC-
C-1986-1134, non-ovig. 2, 3.2 mm, Six
Men’s, | Jun 1985. PARATYPES, NMNH,
2 6, 2.0 mm, Brighton, 20 Aug 1985; 1 4,
damaged, Six Men’s, | Jun 1985; 1 6, 1.8
mm, Greensleeves, 26 Jun 1985; 3 4, 2.0
mm, Bank Reef, 27 Jul 1985; 2 non-ovig 8,
1.4 mm, Greensleeves, 26 Jun 1985; 2 non-
ovig. 2, 1.8 mm. | damaged, Six Men’s 1
Jun 1985; 2 non-ovig. 2, 1.3 mm, 2.4 mm,
Six Men’s, | Jun 1985; 1 non-ovig. 2, 2.4
mm, Brighton, 20 Aug 1985. PARA-
TYPES, USNM 211430, 3 6, 2.0 mm, Bank
Reef, 11 Jun 1985; PARATYPES, USNM
211431, 2 non-ovig. 2, 1.3 mm, 1.8 mm,
Greensleeves, 13 Aug 1985.
Description.—Female: Body about 7 xX
longer than wide. Only tailfan markedly in-
durate. Prominent anterolateral eye with 5
or 6 ommatidia. Pleonites short, free, 1-5
subequal, 6 longer than preceding pleonites,
with broad posterolateral lobes separated by
wide middorsal notch. Telson widening
VOLUME 100, NUMBER 1 191
Fig. 4. Chalixanthura lewisi, male: A, Adult in dorsal view; B, Mandible; C, Maxilliped; D, Uropodal exopod;
E, Pleopod 3; F, Pleopod 1; G, Pleopod 2; H, Antennule; I, Antenna; J, Telson; K, Uropodal endopod and
sympod.
192
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig. 5;
Pereopod 7.
posteriorly, broadly rounded posterior mar-
gin deeply incised into 13-14 acute to nar-
rowly rounded teeth; middorsally with row
of 8 or 9 well separated erect teeth, becom-
ing progressively larger and somewhat re-
curved posteriorly.
Antennule with basal article broad, bear-
ing mediodistal flattened, narrowly trian-
gular, apically narrowly-rounded process
bearing single seta; peduncular articles 2-3
narrow, subequal; flagellum of 7 articles, ar-
ticle 1 short, article 2 longest; single aesthet-
Chalixanthura lewisi, male: A, Pereopod 1; B, Pereopod 2; C. Pereopod 3; D, Pereopod 6; E,
asc on penultimate and antepenultimate ar-
ticles each. Antenna with peduncular article
5 longest; flagellum of 6 setose articles.
Mandible lacking palp; incisor with single
broad strongly sclerotized cusp; subtermin-
al lamina dentata of 2 teeth. Maxilla having
2 broad and 4 narrow distal teeth. Maxil-
liped elongate, slender, proximal segmen-
tation obscure; short distal article with 4
setae.
Pereopod 1, carpus triangular, with very
short free anterior margin; propodus elon-
VOLUME 100, NUMBER 1
gate, distally tapering, palm unarmed except
for few distal setae; unguis about *4 length
of remainder of dactylus. Pereopod 2, is-
chium having 11 sclerotized tubercles near
posterior margin, merus with 2 sclerotized
tubercles; carpus triangular; propodus bear-
ing row of about 20 fringed scales (appearing
as spines in lateral view) plus strong pos-
terodistal fringed spine; unguis about 2
length of remainder of dactylus. Pereopod
7, carpus roughly rectangular with row of
about 9 fringed scales plus strong postero-
distal fringed spine; propodus elongate-rect-
angular, with row of about 14 fringed scales
on posterior margin plus strong postero-
distal fringed spine; unguis slightly less than
Y length of remainder of dactylus. Pleopod
1, rami fused for *%4 length, distal endopod
and exopod with 2—4 plumose setae each.
Uropodal sympod with mesial margin ser-
rate, 4 spines near lateral margin; endopod
with mesial margin longer than lateral mar-
gin, extending well beyond telsonic apex,
margins bearing acute to rounded teeth; ex-
opod proximally broad, margins serrate,
with distal spine-like, apically acute exten-
sion bearing strong recurved tooth on dorsal
surface (2 on right, 1 on left in holotype).
Male: Body about 15 x longer than wide.
Eyes relatively larger than in female, each
with 9-10 ommatidia, extending onto ven-
trolateral surface; mouthparts obsolete. Ple-
onites free, 1-3 subequal, longer than 4 and
5. Telson widening slightly posteriorly, dor-
sally unarmed, posterior margin with 12
acute to narrowly rounded teeth.
Antennule with basal article slightly long-
er than article 2; articles 2 and 3 subequal;
flagellum of 8 articles, article 1 inserted
obliquely on peduncle, bearing 9 elongate
aesthetascs; article 2 with 2 aesthetascs; ar-
ticles 3-8 each with single aesthetasc. An-
tennal peduncle with 3 proximal peduncu-
lar articles short, 2 distal articles elongate,
article 5 just less than twice length of article
4; flagellum of 6 setose articles.
Pereopod 1, carpus triangular, with short
free anterior margin; propodus elongate-
rectangular, with 19-20 fringed spines near
193
posterior margin plus posterodorsal un-
fringed spine; unguis just more than 2 length
of remainder of dactylus. Pereopod 2, prop-
odus elongate-rectangular, with few pos-
terodistal fringed scales and strong fringed
posterodistal spine; unguis '2 length of re-
mainder of dactylus. Pereopod 7, carpus
rectangular, with 5 fringed scales and strong
fringed posterodistal spine; propodus with
about 14 fringed scales on posterior margin
and 2 strong fringed spines. Pleopod 1, basis
about *%3 length of endopod; rami separate,
each with 3-4 elongate plumose setae on
distal margin. Pleopod 2, basis % length of
rami; endopod with copulatory stylet at-
tached proximally; stylet distally rounded,
not reaching beyond rami.
Remarks. —Of the 12 species of Eisothis-
tos described, only four have middorsal spi-
nation of the telson, similar to that seen in
the present species. These are the two Med-
iterranean species E. macrurus Wagele, and
E. pumilis Wagele, E. antarcticus Vanhof-
fen from the Antarctic, and E. crateris
Kensley from St. Paul and Amsterdam Is-
lands. None of these species show the ar-
rangement of large slightly recurved teeth
increasing in length posteriorly on the telson
of the female, and none possess the heavy
recurved tooth on the “‘spike”’ of the uro-
podal exopod. One striking feature, the lam-
inate process of the basal article of the fe-
male antennule, immediately separates E.
teri from all other described species. A sim-
ilar but more elaborate structure is seen in
Licranthura amyle Kensley and Schotte,
1987. In the latter, however, the process
arises from the third peduncular article of
the antennule.
Etymology.—The species is named for
Teri Snelgrove, who has the kindness to feign
interest in marine biology.
Mesanthura paucidens
Menzies and Glynn
Mesanthura paucidens Menzies and Glynn,
1968:27, fig. 9a-g.--Kensley, 1982:335,
i, WSOs MS Ik.
194 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
=>.
C
F
Fig. 6. Eisothistos teri, female: A, Antennule; B, Antenna; C, Maxilliped; D, Left uropod and telson; E,
Pleopod 1; F, Pereopod 1; G, Pereopod 7; H, Mandible; I, Maxilla; J, Pereopod 2.
Material. — 1 specimen from Paynes Bay, Mesanthura pulchra Barnard
ea Och 8. Mesanthura pulchra Barnard, 1925:145S, fig.
Previous records.—Puerto Rico; Carrie OASIS, WRT Fete Un LOZ, LS.
Bow Cay, Belize; shallow water. Material. —3 specimens, Six Men’s, | Jun
VOLUME 100, NUMBER 1 NOS)
Fig. 7. Eisothistos teri, male: A, Antennule; B, Antenna; C, Left uropod and telson; D, pleopod 2; E, Pleopod
1; F, Pereopod 1; G, Pereopod 2; H, Pereopod 7.
1985; 1 specimen Bank Reef, 19 Jul 1985; Minyanthura corallicola Kensley
3 specimens, Paynes Bay, 13 Jul 1985; 1
specimen, Bank Reef, 8 Sep 1985; 1 spec- Minyanthura corallicola Kensley, 1982: 343,
imen, North Bellairs, 19 Jun 1985. figs. 157, 158.
196 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Material.—1 non-ovig. 2, 2 manca, Six
Men’s, 1 Jun 1985. Numerous other spec-
imens from Six Men’s (1 Jun; 19 Jul; 16 Sep
1985), Greensleeves (26 Jun; 13 Aug; 13 Oct
1985), Bank Reef (11 Jun; 27 Jul; 8 Sep
1985). Present but rare at North Bellairs (19
Jun; 2 Aug 1985), Paynes Bay (22 Oct 1985),
and Brighton (20 Aug; 10 Oct 1985). Over-
all, the most abundant anthuridean species.
Previous records. —Carrie Bow Cay, Be-
lize, 6—24 m.
Family Cirolanidae
Cirolana minuta Hansen
Cirolana minuta Hansen, 1890:347, pl. 3
fig. 5, pl. 4 fig. 1.
Material. —2 specimens, Paynes Bay, 13
Jul 1985. Other specimens from Six Men’s
(19 Jul; 16 Sep 1985), Greensleeves (26 Jun;
13 Aug; 13 Oct 1985), Bank Reef (8 Sep
1985), Paynes Bay (13 Jul 1985), Brighton
(5 Jul; 20 Aug; 10 Oct 1985). Rare at all
sites and absent from North Bellairs.
Previous records. —St. Thomas, West In-
dies.
Family Gnathiidae
Gnathia rathi Kensley
Gnathia rathi Kensley, 1984:43, fig. 27a-.
Material. —5 specimens, Six Men’s, | Jun
1985; 12 praniza, Six Men’s, 1 Jun 1985.
Numerous other specimens from Six Men’s
(1 Jun; 19 Jul; 16 Sep 1985), Greensleeves
(26 Jun 1985; 13 Aug; 13 Oct 1985), Bank
Reef (11 Jun; 27 Jul; 8 Sep 1985), North
Bellairs (19 Jun; 2 Aug; 24 Sep 1985). Pres-
ent but rare at Paynes Bay (13 Jul; 27 Aug;
22 Oct 1985) and Brighton (5 Jul; 20 Aug;
10 Oct 1985).
Previous records.—Carrie Bow Cay, Be-
lize, 0.5—36 m.
Family Stenetriidae
Stenetrium patulipalma Kensley
Stenetrium patulipalma Kensley, 1984:52,
figs. 33, 34.
Material. —4 °, Six Men’s, 1 Jun 1985.
Numerous other specimens from Six Men’s
(1 Jun; 19 Jul; 16 Sep 1985), Greensleeves
(26 Jun; 13 Aug; 13 Oct 1985), Bank Reef
(11 Jun; 27 Jul; 8 Sep 1985), North Bellairs
(19 Jun; 2 Aug; 24 Sep 1985), Brighton (5
Jul; 20 Aug; 10 Oct 1985). Present but rare
at Paynes Bay (13 Jul; 27 Aug 1985).
Previous records.—Carrie Bow Cay, Be-
lize, 9.1-15.2 m.
Stenetrium spathulicarpus Kensley
Stenetrium spathulicarpus Kensley, 1984:
55, figs. 35, 36, 37d.
Material.—1 6, 2 2, 3 juvs., Paynes Bay,
27 Aug 1985; 1 6, 3 juvs., Six Men’s, 1 Jun
1985. Numerous other specimens from
Paynes Bay (13 Jul; 27 Aug; 22 Oct 1985).
Present but rare at Six Men’s (1 June; 19
Jul; 16 Sep 1985), Greensleeves (26 Jun; 13
Aug; 13 Oct 1985), Bank Reef (11 Jun; 27
Jul; 8 Sep 1985), North Bellairs (19 Jun; 2
Aug 1985), and Brighton (5 Jul; 20 Aug; 10
Oct 1985).
Previous records. —Carrie Bow Cay, Be-
lize, intertidal—20 m.
Family Janiridae
Carpias minutus (Richardson)
Bagatus minutus (Richardson), Pires, 1982:
231, figs. 1-16.
Carpias minutus: Bowman and Morris,
1979:650, figs. 1-3, 4a-d.
Material. —1 6, 1 ovig. 2, 1 2, Six Men’s,
1 Jun 1985. Numerous other specimens
from Six Men’s (1 Jun; 19 Jul; 16 Sep 1985),
Greensleeves (26 Jun; 13 Aug; 13 Oct 1985),
Bank Reef (11 Jun; 27 Jul; 8 Sep 1985),
North Bellairs (19 Jun; 2 Aug; 24 Sep 1985),
Brighton (5 Jul; 20 Aug; 10 Oct 1985). Pres-
ent but rare at Paynes Bay (13 Jul; 27 Aug
1985).
Previous records. —Bermuda; Georgia,
U.S.A. to Brazil; Azores, intertidal to shal-
low intratidal.
VOLUME 100, NUMBER 1
Family Joeropsidae
Joeropsis personata Kensley
Joeropsis personatus Kensley, 1984:70, fig.
44.
Material. —3 6, Bank Reef, 27 Jul 1985.
Other specimens present but rare at Bank
Reef (11 Jun; 27 Jul; 8 Sep 1985). Not pres-
ent at any other site.
Previous records.—Carrie Bow Cay, Be-
lize, 1-20 m.
Acknowledgments
Financial support for sample collection
was supplied by a Natural Sciences and En-
gineering Research Council of Canada
(NSERC) grant to Dr. J. B. Lewis and an
NSERC Postgraduate Fellowship to the ju-
nior author. We wish to thank Dr. Lewis
for advice and assistance with this project,
and Dr. Wayne Hunte of Bellairs Research
Institute for his hospitality and advice while
in residence at Bellairs. Fahmida Rafi of
NMNS in Ottawa assisted in identification
of some specimens and arranged deposition
of material in NMNS. Barbara Conlin was
an enthusiastic dive partner in sample col-
lection, and Kim Silkauskas endured the te-
dious task of assisting in sorting. We are
grateful to Dr. Thomas E. Bowman of the
Smithsonian Institution for reading and
commenting on a draft of this paper.
Literature Cited
Barnard, K. H. 1925. A revision of the family An-
thuridae (Crustacea, Isopoda), with remarks on
certain morphological peculiarities. — Journal of
the Linnean Society of London, Zoology 36:109-
160.
Bowman, T. E., and B. F. Morris. 1979. Carpias
Richardson 1902, a senior synonym of Bagatus
Nobili 1906, and the validity of Carpias mi-
nutus (Richardson, 1902) (Isopoda: Asellota:
Janiridae).— Proceedings of the Biological So-
ciety of Washington 92(3):650-657.
Brander, K. M., A. A. Mcleod, and W. F. Humphreys.
1971. Comparison of species diversity and
ecology of reef-living invertebrates on Aldabra
iy
Atoll and at Watamu, Kenya.—Symposium,
Zoological Society of London 28:397-—431.
Hansen, H. J. 1890. Cirolanidae et familiae nonullae
propinquae Musei Hauniensis.—K. Danske Vi-
denskabernes Selskab Skrifter 6(3):239-426.
Kensley, B. 1982. Anthuridea (Crustacea: Isopoda)
on Carrie Bow Cay, Belize. Jn K. Ritzler and
I. Macintyre, eds., The Atlantic Barrier Reef
ecosystem at Carrie Bow Cay, Belize, 1: Struc-
ture and communities.—Smithsonian Contri-
butions to Marine Sciences 12:321-353.
1984. The Atlantic Barrier Reef Ecosystem
at Carrie Bow Cay, Belize, III: New Marine Iso-
pods.—Smithsonian Contributions to Marine
Sciences 24:1-81.
, and M. Schotte. 1987. New records of isopod
Crustacea from the Caribbean. — Proceedings of
the Biological Society of Washington 100:216-—
247.
Menzies, R. J.,and P. W. Glynn. 1968. The common
marine isopod Crustacea of Puerto Rico.—
Studies of the Fauna of Curacao and other Ca-
ribbean Islands 27:1-133.
Lewis, J. B. 1960. The coral reefs and coral com-
munities of Barbados, West Indies. — Canadian
Journal of Zoology 38:1133-1145.
, and R. Bray. 1983. Community structure of
ophiuroids (Echinodermata) from three differ-
ent habitats on a coral reef in Barbados, West
Indies.— Marine Biology 73:171-176.
1975. Community patterns on a submerged
barrier reef at Barbados, West Indies.—Inter-
nationale Revue der gesamten Hydrobiologie 60:
719-736.
Pires, A. M. 1982. Taxonomic revision of Bagatus
(Isopoda, Asellota) with a discussion of onto-
genetic polymorphism in males.—Journal of
Natural History 16:227-259.
Tomascik, T., and F. Sander. 1986. Effects of eutro-
phication on reef building corals. Part III. Struc-
ture of the scleractinian coral communities on
the fringing reefs of Barbados, W.I.— Marine Bi-
ology [In press].
Wagele, J. W. 1981. Zur Phylogie der Anthuridea
(Crustacea, Isopoda) mit Beitragen zur Leben-
sweise, Morphologie, Anatomie, und Taxon-
omie.— Zoologica 132:1-127.
Ott, B.
(BK) Department of Invertebrate Zool-
ogy, National Museum of Natural History,
Smithsonian Institution, Washington, D.C.
20560; (PS) Institute of Oceanography,
McGill University, 3620 University Street,
Montreal, P.Q. H3A 2B2, Canada.
PROC. BIOL. SOC. WASH.
100(1), 1987, pp. 198-215
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, MEXICO
Horton H. Hobbs, Jr.
Abstract. —The Mexican crayfish Procambarus (Austrocambarus) mexicanus
(Erichson, 1846), from the state of Veracruz, is redescribed and declared a
subjective senior synonym of Cambarus aztecus Saussure, 1857. The name
Procambarus (Austrocambarus) olmecorum is proposed as a substitute name
for the sympatric Procambarus aztecus Villalobos (1954) which is also rede-
scribed and illustrated. As complete a list of references as the author has been
able to assemble is offered for both species.
Uncertainty seems to have existed con-
cerning the identity of both Astacus (Cam-
barus) Wiegmanni and Astacus (Cambarus)
Mexicanus since they were described by Er-
ichson in 1846, and the quandary was com-
pounded with the description of Cambarus
aztecus by Saussure in 1857. Of those who
attempted to clarify the identity of these
crayfishes and other of their close relatives,
the most recent was Villalobos (1954, re-
peated in 1955 and 1983) who presented a
historical summary of all recorded pertinent
facts and opinions, added additional infor-
mation, and offered proposals that he be-
lieved would clarify the confusion that was
clearly evident in, and had persisted since
the publication of, Hagen’s (1870) mono-
graph of the North American crayfishes.
Only facts that seem immediately pertinent
to establishing the identity of Astacus
(Cambarus) mexicanus and de Saussure’s
(1857) Cambarus aztecus are repeated here,
although as complete synonymies for the
species treated as I have been able to ferret
are included.
The two crayfishes described by Erichson
were the first reported from Mexico, but,
unfortunately, in his rather brief accounts
of them, he did not record the locality from
which his specimens had been collected.
Apparently these crayfishes were not avail-
able to subsequent students of crayfishes.
Whether or not Saussure sought Erichson’s
material is not known, but in describing the
third and fourth species (Cambarus Mon-
tezumae and Cambarus aztecus) from Mex-
ico, he did not mention having seen Erich-
son’s “‘types.’? Almost certainly his
knowledge of these species was derived sole-
ly from Erichson’s descriptions, and only
two specific differences were noted between
his C. aztecus and C. mexicanus: “... les
mains sont comprimées, non cylindriques
comme chez l’espéce citée, puisque les bras
sont épineux, etc.’’ (Saussure 1858:461).
Hagen was unsuccessful in his search for
Erichson’s types in the Berlin Museum in
September 1870 (Faxon 1885:38) as was von
Martens (1872:131), but Hagen (1870) ex-
pressed the opinion that Saussure’s C. az-
tecus was identical to Erichson’s C. Wieg-
manni and that C. Montezumae was the
young of C. mexicanus. These opinions are
clearly without merit, for males of C. Mon-
tezumae, as described, possess hooks on the
second and third pairs of pereiopods, C.
VOLUME 100, NUMBER 1
mexicanus, on the third, and C. wiegmanni,
on the third and fourth. Hagen suggested
that the different positions of the hooks were
based upon Erichson’s counting the che-
lipeds as the first pair of legs and Saussure’s
numbering the leg immediately posterior to
the chelipeds as the first. This suggestion
cannot be taken seriously since indeed the
male of Saussure’s C. Montezumae does ex-
hibit hooks on the second and third pairs
of pereiopods. Faxon (1885), while unable
to distinguish between Cambarus mexicanus
and C. aztecus, recognized the distinctness
of C. wiegmanni and C. montezumae on
the basis of the characters just cited.
Because of the lack of material from Mex-
ico, not until 1954 was an apparently re-
warding effort made to clarify the status of
Cambarus mexicanus, C. aztecus, and C.
weigmanni. In resolving the synonymy of
C. aztecus with C. mexicanus proposed by
Faxon (1885, 1914) it was necessary to as-
sociate the latter name with specimens from
a known locality, and with good reason Vil-
lalobos (1954: 305) selected a specimen in
the Academy of Natural Sciences of Phila-
delphia from El Mirador de Zacuapan, 8 km
northeast of Huatusco, Veracruz, as the
neotype of Erichson’s A. (Cambarus) mex-
icanus. Apparently unaware that syntypes
of C. aztecus were still extant, he described
and illustrated specimens that he had col-
lected in the presumed type locality, To-
matlan, 14 km south-southeast of Huatus-
co, Veracruz, as members of Saussure’s
species.
Had the syntypes of Saussure’s C. aztecus
not been extant, the problem of the identity
of the two species would have been solved.
That the specimens described by Villalobos
as Procambarus aztecus were not conspe-
cific with a syntype of Saussure’s species in
the collections of the Smithsonian Institu-
tion came to light when, in preparing illus-
trations for a checklist of the North and
Middle American crayfishes (Hobbs 1974),
I discovered that my drawing of the cara-
pace of the Smithsonian syntype did not
199
agree with Villalobos’ illustration (Villalo-
bos 1954:pl. III, fig. 1). I pointed this out
to Dr. Villalobos, and, prior to his untimely
death in October 1983, we had agreed to
collaborate in attempting to rectify previ-
ously erroneous concepts of the two species.
The most conspicuous difference between
the specimens illustrated by Villalobos and
the syntype is in the width of the areola.
The illustration of C. aztecus provided by
Saussure strangely lacks lines representing
the branchiocardiac grooves so that in it the
expanse of the areola cannot be determined.
This could well be interpreted as the ani-
mal’s possessing an obliterated one! But in
the syntypes in the Muséum d’Histoire Na-
turelle, Geneva, and in that in the Smith-
sonian Institution currently available to me
the areola is distinctly “open”? throughout
its length, about 10 times as long as broad —
not obliterated along part of its length as
occurs in Villalobos’ P. aztecus.
On the basis of Villalobos’ (1954:312)
designation of a neotype of Astacus (C.)
mexicanus, the identity of this species be-
came established, and the locality from
which the neotype was collected is known!
Unfortunately, this specimen is in poor con-
dition, and instead of preparing an inade-
quate description based upon it, I have cho-
sen neotopotypic specimens which have
been compared with the neotype, to illus-
trate the species. In view of the existence of
syntypes, the identity of Cambarus aztecus
Saussure seems clear, and now their source
is reasonably assured. Since presumed to-
potypes proved to be members of another
species, and there are at least five ““com-
munities” in Mexico bearing the name To-
matlan, the question had to be asked as to
whether or not the locality given by Saus-
sure, “Pris a Tomatlan, dans les Terres-
Chaudes”’ is the same as that from which
the specimens described by Villalobos,
‘‘Tomatlan, 14 km S.S.W. of Huatusco, Ve-
racruz,’ came? Not until specimens that
were conspecific with the syntypes became
available from or nearby one of the To-
200
matlans could reasonable certainty of the
location of that referred to by Saussure be
assumed.
Of the five Mexican ““ccommunities”’ bear-
ing the name Tomatlan listed in the gazet-
teer available to me, only that 14km S.S.W.
of Huatusco, Veracruz (19°02'N, 97°00'W),
lies within the known range of the subgenus
Austrocambarus to which Procambarus (A.)
aztecus belongs. Moreover, specimens that
differ only in minor respects from the syn-
typic male at hand were obtained from lo-
calities both north and south of the town.
Thus, there is every reason to believe that
it lies within the range of the species and is
likely the same Tomatlan from which Saus-
sure’s specimens came.
The neotype and neotopotypes of P. (A.)
mexicanus have been compared with the
syntype of C. aztecus and with other spec-
imens assignable to it from a number of
localities in the vicinity of the two type lo-
calities (which are no more than 40 km
apart), and I am convinced that the few dif-
ferences noted between specimens assigned
to the two, represent nothing more than in-
dividual variations. Among those features
noted, none even seems to be restricted to
a limited part of the range of the species. In
view of these observations, I am proposing
that Cambarus aztecus Saussure, 1857, be
placed in the synonymy of Cambarus mex-
icanus (Erichson, 1846). Neither the char-
acters pointed out by Saussure nor differ-
ences noted between the neotypes of the
latter and syntypes of C. aztecus will serve
consistently to separate populations of one
from the other. I have examined, but do not
have before me, the specimens that Villa-
lobos described and figured as members of
Cambarus aztecus. Available, however, are
series from nearby localities that appear to
be unquestionably conspecific with his ma-
terial from Tomatlan and from near Cos-
comatepec, both localities in Veracruz.
These specimens exhibiting an areola that
is obliterated along a part of its length must
be accorded a new name, and a description
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
and illustrations of this crayfish are offered
herein. Because of the confusion surround-
ing the identities of Procambarus (A.) mex-
icanus and P. (A.) aztecus, a description and
illustrations of the syntypic male, form I, of
the latter are included.
Procambarus (Austrocambarus)
mexicanus (Erichson)
Fig. |
Astacus (Cambarus) Mexicanus Erichson,
1846:99-100 [Type locality: Mexico, re-
stricted by Villalobos (1954:305) to El
Mirador de Zacuapan, 8 km NE of Hua-
tusco, Veracruz; neotype: male, form I,
Academy of Natural Sciences of Phila-
delphia, no. 4176].—Villalobos, 1953:
352, 353; 1954:299, 300, 301, 302; 1955:
15; 1983:19.
Astacus Mexicanus. —Erichson, 1846:87,
88.—Hagen, 1870:7.— Martens, 1872:
131.
Astacus mexicanus. —Dana, 1852:522.—
Hobbs, 1972a:1.
Cambarus aztecus Saussure, 1857:503—504,
fig. 23 [Type locality: ““Pris 4 Tomatlan,
dans les Terres-Chaudes,’’ Veracruz,
Mexico. Syntypes: Muséum d’Histoire
Naturelle, Geneva, uncatalogued, 13 dry
specimens; USNM, 20682, 1 male, form
I.]; 1858:460-—461, pl. III: fig. 23.—Ha-
gen, 1870: 11, 12, 55.—Martens, 1872:
131.—Schmeltz, 1874:79.— Faxon, 1884:
141, 142; 1885:5, 9, 10, 38, 51 m, 123,
172, 174; 1914:410.—Torralbas, 1917:
596 (53), figs. 61, 62.— Villalobos, 1953:
353-363; 1954:300, 302, 314; 1955: 15;
1982:19.—Hobbs, 1972a:2.
Cambarus Mexicanus. — Saussure, 1858:460
(44).— Hagen, 1870:11, 12, 75, 84-85, 98,
102 (part). — Faxon, 1884:138, 139, 141-
142 (part); 1885:5, 8, 9, 38-39, 47-48,
50-51, 53, 76, 158, 172-174, 177 (part). —
Villalobos, 1950:381; 1955:140; 1983:
136.—Hobbs, 1972a:2.
Cambarus mexicanus. —Saussure, 1858:461
(45).—Ortmann, 1892:12 (part); 1902:
VOLUME 100, NUMBER 1
277, 283, 284, 341; 1905a:99, 103; 1905b:
435, 436.—Faxon, 1898:649; 1914:363,
410 (part).—Hay, 1899:964.—Harris,
1903:58, 108, 151, 155 (part).—Ellis,
1919:254, 264.—Goodnight, 1940:63.—
van Straelen, 1942:5.— Villalobos, 1946:
216; 1950:381, 382; 1955:140; 1983:
136.—Hart, 1961:78, 79.—Hobbs and
Villalobos, 1964:321, 324.—Holt, 1968:
LS:
Cambarus Aztecus. —Hagen, 1870:11.
Astacus Aztecus. —Martens, 1872:131.
Cambarus (Cambarus) mexicanus. —Ort-
mann, 1905a:101, 103.— Villalobos,
1946:216.
Cambarus (Procambarus) mexicanus. —
Ortmann, 1905b:437—438, 441 [by im-
plication, in part]; 1906:11, 15, 21, 23;
1913:416, 417.—Hobbs, 1942a:57, 60,
61, pl. 2:fig. 12.—Rioja, 1940:249.—Vil-
lalobos, 1953:354; 1954:302; 1955:17;
1983:20.—Hobbs and Villalobos, 1964:
SD.
Astacus Astecus.— Valdés Ragués, 1909:180
[erroneous spelling].
Procambarus aztecus.— Hobbs, 1942b:342
[by implication].— Villalobos, 1953:346,
363; 1954:300, 302, 304, 314 (in part);
1955:239; 1982:227.—Hart and Hart,
1974:124, 142.—Spitzy, 1976:445.
Procambarus mexicanus.—Hobbs, 1942b:
342 [by implication], 355, pl. III, figs. 11,
13; 1942c:38; 1943:203, 205, 206; 1966:
70, 71; 1967:8; 1969:118; 1971:3, 12, 30,
31, 41; 1972b:151, figs. 5h, 30f, 35b,
36a.— Villalobos, 1946:218; 1948:182;
1950:382, 385, pl. 4: figs. 2, 4; 1953:346,
365; 1954:299, 302-307, 310, 312-314,
317, 318, 321, 323, 328, 335, 346, 364,
375, pls. 1, 2; 1955:15, 22, 139, 140, 141,
INOS 172,173, 176, 178, 183, 201,
239, 240, pls. 28 (figs. 2, 4), 34, 35; 1983:
17, 25, 134, 136, 154-163, 166-168, 170,
iWialpmly Geel92227, 228, pls. 28 (figs. 2,
4), 34, 35.—Rioja, 1949:316, 321 (part),
322, 327.—Hobbs and Villalobos, 1964:
313, 314, 346.—Hobbs III, 1969:21.—
Straskraba, 1969:25.— Holt, 1973:4, 5,17,
201
20, 25, 26, 32.—Hart and Hart, 1974:22,
23, 86.—Spitzy, 1976:443, 444.—Fitz-
patrick, 1983:165, fig. 171.
Procambarus mexiacnus. — Villalobos,
1954:304 [erroneous spelling].
Procambarus Mexicanus. —Creaser, 1962:
7, fig. 8.
Procambarus (Austrocambarus) mexican-
us. —Hobbs, 1972a:6; 1972b (1976):43;
1974:45, fig. 173.—Feldmann et al., 1981:
794, text-fig. 3.— Fitzpatrick, 1983:202.—
Villalobos, 1982:219.
Procambarus (Austrocambarus) aztecus. —
Hobbs, 1972a:6; 1972b (1976):43, 150,
figs. 30e, 34c, 36b; 1974:44, 123, fig.
172.— Fitzpatrick, 1983:202.
Diagnosis. —Body pigmented, eyes well
developed. Rostrum of adults without mar-
ginal spine and median carina. Carapace
with or without small cervical spine or tu-
bercle. Areola 6.5 to 17 times as long as
wide and constituting 30.2 to 37.4 (average
32.8) percent of total length of carapace (37.1
to 46.0, average 40.3, percent of postorbital
carapace length). Suborbital angle weak and
obtuse; infraorbital spines lacking. Postor-
bital ridge moderately strong and anterior
extremity with or without spine or tubercle.
If present, branchiostegal spine small. An-
tennal scale about twice as long as broad,
widest slightly distal to midlength. Cheliped
studded with subsquamous tubercles from
midlength of merus to about midlength of
fingers. Ischium of third pereiopod of first
form male with simple, strong, acute hook
usually overreaching basioischial articula-
tion; hook not opposed by tubercle on cor-
responding basis. First pleopods of first form
male reaching coxae of third pereiopods,
symmetrical, usually contiguous basally,
lacking proximomesial spur; simple angular
or produced shoulder present on cephalic
surface at about base of distal eighth; lacking
subterminal setae; terminal elements con-
sisting of short, acute distolaterally directed
mesial process extending beyond short, cor-
neous subacute, cephalodistally directed
202 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig. 1. Procambarus (A.) mexicanus—All except 1 (which from km 57 on Fortin-Huatusco-Conejos Road)
from El Mirador de Zacuapan, 8 km NE of Huatusco, Veracruz, Mexico (a-c, e, g, h, j-l, n from first form male;
d, f, from second form male; i, m, from female): a, Lateral view of carapace; b, d, Mesial view of first pleopod;
c, Cephalomesial view of distal part of first pleopod; e, Epistome; f, h, Lateral view of first pleopod; g, Caudal
view of distal part of first pleopod; i, Annulus ventralis and adjacent sternal elements; j, Antennal scale; k,
Dorsal view of carapace; 1, Proximal podomeres of third, fourth, and fifth pereiopods; m, n, Distal podomeres
of cheliped.
VOLUME 100, NUMBER 1
central projection. Female with hinged an-
nulus ventralis slightly longer than prean-
nular plate, almost twice as wide as long,
tapering posteriorly, and bearing short, sin-
uous sinus on midposterior surface; postan-
nular sclerite as wide as, or slightly wider
than, annulus but shorter; first pleopod re-
duced.
Topotypic male, form I: Cephalothorax
(Fig. la, k) subcylindrical; maximum width
of carapace slightly less than height at level
of caudodorsal margin of cervical groove
(11.8 and 12.0 mm). Abdomen only little
narrower than thorax (11.2 and 11.8 mm).
Areola 10.6 times as long as wide with 2 or
3 punctations across narrowest part. Ce-
phalic section of carapace 1.5 times as long
as areola, latter constituting 34.1 percent of
entire length of carapace (40.9 percent of
postorbital carapace length). Surface of car-
apace densely punctate dorsally, granulate
laterally. Rostrum moderately broad with
weakly convergent, slightly thickened mar-
gins, latter tapering rapidly from base of
acumen which gently upturned apically and
reaching slightly beyond distal end of pen-
ultimate podomere of antennular peduncle;
upper surface distinctly concave, strongly so
anteriorly. Subrostral ridge weak and evi-
dent in dorsal view only in posterodorsal
part of orbit. Postorbital ridge moderately
well developed and terminating cephalically
in small acute spine not attaining level of
orbit. Suborbital angle obtuse and not
prominent. Branchiostegal spine clearly de-
fined. Cervical spine represented by very
small tubercle.
Abdomen longer than carapace (26.6 and
24.9 mm). Pleura of third through sixth seg-
ments subtruncate ventrally and rounded
caudoventrally. Cephalic section of telson
with 3 spines in right and 4 in left postero-
lateral corners, that adjacent to lateralmost
on both sides movable. Cephalic lobe of
epistome (Fig. le) subtriangular, lacking an-
teromedian excavation; main body with
distinct fovea; epistomal zygoma arched.
Ventral surface of proximal podomere of
antennule with strong ventromesial spine
203
short distance distal to midlength. Antennal
peduncle with short distolateral spine on
basis; ischium with small tubercle; flagel-
lum broken but probably reflexed only
slightly posterior to first abdominal seg-
ment. Antennal scale (Fig. 1j) about twice
as long as wide, greatest width distal to mid-
length; lamellar area approximately twice
width of thickened lateral part. Third max-
illiped slightly overreaching penultimate
podomere of antennal peduncle; mesial sec-
tor of ventral surface of ischium with
crowded clusters of both stiff and plumose
setae, lateral sector studded with mat of plu-
mose setae, ischium not produced distolat-
erally; merus with setae similarly dispersed.
Right chela (Fig. In) ovate in cross-sec-
tion, weakly depressed; palm about 1.3 times
as long as broad; its length little less than
half length of chela; except for ridges and
distal part of fingers, almost completely
studded with squamous tubercles. Mesial
surface of palm with tubercles somewhat
staggered but roughly forming 3 rows of 9
to 11. Both fingers with low submedian lon-
gitudinal ridges dorsally and ventrally; ridges
flanked proximally by tubercles and distally
by punctations. Opposable margin of fixed
finger with upper row of 11 tubercles, sec-
ond from base largest, extending from base
of finger almost to terminal corneous tip,
and lower row of 6, proximalmost largest,
along distal half of finger (between level of
third and ninth tubercles of upper row); band
of minute denticles extending along entire
length of finger, separating upper and lower
rows of tubercles; lateral surface of finger
with row of tubercles along proximal half
followed by row of punctations. Opposable
margin of dactyl with dorsal row of 14 tu-
bercles (third from base largest, some too
small to be included in illustration) and low-
er row of 6 lying at level between second
and thirteenth tubercles of upper row; me-
sial margin of finger with basal cluster of
tubercles narrowing distally to single row
where approaching corneous terminal part
of finger.
Carpus of cheliped, except for proximal
204
Table 1.—Measurements (mm) of Procambarus (A.)
mexicanus. (Left chela of syntype of Cambarus aztecus;
right of others.)
Neo- Topotypes
holoty Pe’ Se
él 31 Q Sit) eri
Carapace:
Entire length MD ANS) WSS HAO DA)
Postorbital
length IO) BOS) Bile Biles A783
Width NOS ts WA) UO Mies
Height LCG 2201269 RO 9a
Areola:
Width 0.7 OFSi OLN OLS Ol
Length 7.9 SN SL 824 EZ
Rostrum:
Width 4.0 42 46 4.3 3.6
Length 4.6 46) S88 Sal) ALS
Chela:
Length, palm
mesial
margin VS Sin) 656 Telia nO
Palm width 6.0 6:65.59) 54 45
Length, lateral
margin 19.0 20.0 17.1 18.6 14.8
Dactyl length soll NOLO, 84h D5» . Oa
Abdomen:
Width _ Wo WAS O02) *©.7/
Length — 26.6 28.4 268 —
* Syntype of Cambarus aztecus.
dorsal and ventral areas and distolateral
band, almost entirely tuberculate; only few
tubercles mesially and 2 distoventral ones
noticeably larger and less squamous than
others. Merus also strongly tuberculate ex-
cept for much of mesial and lateral surfaces;
no spiniform ones dorsodistally; ventrolat-
eral row consisting of 9 tubercles and ven-
tromesial one of 15; both rows flanked by
additional tubercles. Ventromesial surface
of ischium with row of 3 prominent tuber-
cles subtended by few scattered ones.
Hook on ischium of third pereiopod (Fig.
11) simple, acute, and overreaching basiois-
chial articulation, not opposed by tubercle
on basis. Coxa of fourth pereiopod without
caudomesial boss, that of fifth with small
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
rounded boss compressed in longitudinal
axis of body.
Sternum between third, fourth, and fifth
pereiopods shallow, margined with plu-
mose setae.
First pleopods (Fig. 1b, c, g, h) as de-
scribed in “Diagnosis.”
Uropods with both lobes of proximal
podomere bearing acute spines; mesial ra-
mus with distomedian spine terminating
premarginally and with well developed dis-
tolateral spine; lateral ramus with fixed dis-
tolateral spine.
Topotypic female: Differing, other than
in secondary sexual characteristics, from
male, form I, in following respects: cervical
spine obsolete; cephalic section of telson
with 3 spines in each caudolateral corner;
tubercle on ischium of antennal peduncle
vestigial; mat of plumose setae on ventro-
lateral sector of ischium of third maxilliped
less conspicuous; opposable margin of fixed
finger of right chela (Fig. 1m) with upper
row of 5 tubercles (proximalmost largest) in
proximal three-fifths and lower row of 4
along penultimate fourth; opposable margin
of dactyl with double row of 9 (10 on left)
tubercles (proximalmost largest) in proxi-
mal three-fifths; minute denticles arranged
in single row on dactyl but forming narrow
band on propodus; ventral surface of merus
with lateral row of 11 tubercles and mesial
one of 15 (10 and 14, respectively on left
member); carpus with row of 4 tubercles.
(See Table | for measurements.)
Annulus ventralis and associated sternal
sclerites (Fig. 11) as illustrated.
Topotypic male, form II: Differing from
first form male in following respects: ros-
trum not so concave, especially anteriorly;
subrostral ridges evident in dorsal aspect to
base of acumen; cephalic section of telson
with 3 spines in each caudolateral corner;
ischium of left antennal peduncle with well
developed spiniform tubercle; opposable
margin of fixed finger of cheliped with upper
row of 12 tubercles (proximalmost largest)
VOLUME 100, NUMBER 1
and ventral row of 6 (10 and 5 on left chela);
opposable margin of dactyl of cheliped with
upper row of 9 tubercles (proximalmost
largest) and lower row of 4 (5 on left); mi-
nute denticles forming much narrower band
on both fingers than in male, form I; single
spiniform tubercle present dorsodistally on
merus of cheliped, ventral surface with lat-
eral row of 9 and mesial row of 15 tubercles,
rows joined distally by row of 4 tubercles;
hook on ischium of third pereiopod not
reaching basioischial articulation. First
pleopods (Fig. 1d, f) not contiguous basally,
strongly resembling those of first form male
except both terminal elements more robust,
and shoulder on cephalic surface slightly
more strongly produced; oblique juvenile
suture also clearly defined.
Size. —The largest specimen examined is
a second form male having a carapace length
of 31.0 (postorbital carapace length, 26.0)
mm. The smallest first form male has cor-
responding lengths of 18.8 and 14.5 mm.
Neither ovigerous females nor ones carrying
young have been observed.
Type locality. —El Mirador de Zacuapan,
8 km northeast of Huatusco, Veracruz. As
pointed out above, Erichson did not cite a
specific locality in Mexico when he de-
scribed this crayfish. In his treatment of the
mexicanus group, however, Villalobos
(1954:312) chose a specimen from the lo-
cality cited here as the neoholotype, thereby
designating El Mirador de Zacuapan as the
type locality of Erichson’s species.
Types. —Neoholotype (6 I), Academy of
Natural Sciences of Philadelphia (ANSP
421); neoallotype (2) and neomorphotype (6
II), Instituto de Biologia de la Universidad
de Mexico (all designated by Villalobos
1954).
Range and specimens examined. —Spec-
imens of this crayfish have been reported
from a number of localities in Mexico south
of the Cordillera Volcanica Transversal. All
of those of which I am aware are listed here,
and those from which material has been ex-
205
amined by me are followed by an enumer-
ation of the specimens seen; at least some,
and perhaps most, of the other localities
need to be confirmed. Locations of the col-
lections cited are recorded as follows: ANSP
(Academy of Natural Sciences of Philadel-
phia), FMNH (Field Museum of Natural
History, Chicago), IBM (Instituto de Biol-
ogia, Universidad Nacional Autonoma de
Mexico), MCZ (Museum of Comparative
Zoology, Harvard University), USNM (Na-
tional Museum of Natural History, Smith-
sonian Institution).
VERACRUZ; (1) El Mirador (Faxon
1898:649) (=El Mirador de Zacuapan, 8 km
NE of Huatusco—restricted type locality;
see Villalobos 1954:312) 1 6I (ANSP), neo-
type, 1894, Dr. Sartorius, coll.; 1 61, 2 4 II,
5 2 (USNM), Nelson & Goldman, coll. (2)
Texola (near Xico), 1 6 I, 2 6 II (ANSP), 21
Mar 1899, S. N. Rhoades (Ortmann 1906:
11). (3) Jalapa (FMNH) (Faxon 1914:363).
(4) Hacienda de Tenejapa (IBM?) (Rioja
1949:316). (5) Hacienda de Jalapilla, a 5 km
al SE de Orizaba (IBM?) (Rioja 1949:322).
(6) Rincon de La Doncella cercanias de Ciu-
dad Mendoza (IBM?) (Villalobos 1953:365).
(7) Tres Puentes, Col. Emiliano Zapala, Ja-
lapa (IBM?) (Hobbs 1971:41). (8) Rio Be-
jyucos, 11.2 km S of Jalapa, 13 ¢ I, 30 4H,
44 9, 24j 6, 34) 2 (USNM), 18 May 1930,
E. P. Creaser & Gordon. (9) Rio Bejucos,
4.8 km S of Jalapa, 18 May 1930, 1 6 Il
(USNM), E.P.C. et al. (10) Trib. to Rio Be-
jucos at Coatepec, 1 61, 5 6 II, 10 2, 7j 4,
73 2(USNM), 25 Jul 1971, A. Villalobos F.
(11) Arroyo Bejucos between Jalapa and
Coatepec, among Eichornia, 1 61(USNM),
6 Sep 1971, N. Vidal. (12) Teocelo, at 300
m, 2 6I, 1 2, 1j 2 (USNM), 30 Dec 1940,
W. W. Dalquest. (13) Km 35 carretera Cor-
doba-Huatusco, 3 61, 2 6 II, 5 2, 3j 4, 3) 2
(USNM), 26 Apr 1980, A.V.F.; 6 61, 15 4
II, 26 9(USNM), 26 Apr 1980, A.V.F. (14)
Huatusco, km 57 on Fortin-Huatusco-Co-
nejos Road, | 6 I, 1 2, 4j 6, 2} 2 (USNM), 8
Jul 1962, P. C. Holt. (15) Rio Jamapa, 6 to
206
7 km NE of Coscomatepec (Holt 1973:17),
1 2, 9j 6, 23 2 (USNM), 9 Jul 1962, P.C.H.
(16) Spring at Tlilipan (Hobbs 1971:30),
141,2 611, 2 2, 2} ¢(USNM), 3 Aug 1967,
J. R. Reddell. (17) Ojo de Agua, Thilapan
(Hobbs 1971:30), 3 61, 3 2(USNM). 4 Aug
1967, J.R.R. (18) Pond 19.2 km E of Jalapa,
1 2, 4j 46, 1j 2 (USNM), 6 Mar 1941, N. E.
Hartweg. (19) Santa Maria, 1 2(USNM), 12
Feb 1894, USDA Biol. Expedition (Faxon
1898:649). (20) 3.5 km W of Huatusco on
Hwy 143, 300 m alt, 1 6 II, 1 2(USNM), 7
Jan 1978, J. R. Dixon. (21) 9.6 km SE of
Coatepec on road to Teocelo, 1180 m alt,
1 2(USNM) 8 Jan 1978, J.R.D. (22) Stream
at water house about halfway up Los
Cumbres, SW of Orizaba, 2 6 I, 3 2, lj 3
(USNM), 27 Dec 1940, H.H.H., Jr. (23) Na-
cimiento de Manzanilla, 11 km N Potrero,
5 6 Il, 4 9, 1j 6, 53 2 (USNM), 6 Jan 1977,
J. R. R. & A. Grubbs.
In addition, specimens with areolae rang-
ing from 12 to 17 times as long as broad,
from two additional localities in the State
of Veracruz (Fortin de Las Flores, 1 ¢ I, 7
2, 1) 2(USNM), 1 Nov 1969, A. Argano &
V. Sbordoni; Copes Estate, Orizaba, 2j 2
(USNM), date?, F. Sumichrast) are tenta-
tively assigned to P. (A.) mexicanus. These
specimens exhibit an array of variation suf-
ficient to point out the difficulty in identi-
fying members of the mexicanus Group of
the subgenus Austrocambarus. (See Villa-
lobos 1954:306.) Three of them possess ros-
tra with produced, angular marginal prom-
inences. One has strong cervical spines,
another exhibits a strong spine on the left
and arather weak one on the right, four have
weak ones, and, in three, there is hardly a
trace of these spines; moreover, there is no
correlation between the degree of their de-
velopment and the size of the animal. The
areola is narrower than that of most speci-
mens here assigned to P. (A.) mexicanus,
and the first form male is unique among all
Austrocambarus that I have examined in
that hooks are lacking from the ischia of the
third pereiopods. The first pleopods are sit-
uated quite far apart; the mesial process is
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
short, and the distal margin of the shoulder
on the cephalic surface lies perpendicular to
the axis of the pleopod, not produced ceph-
alodistally.
Villalobos (1953:365-—366) listed 18 lo-
calities from which he had specimens be-
longing to the ““groupo mexicanus” (=Sub-
genus Austrocambarus). In his review of the
group (1954, 1955, 1983), however, I have
found references to only three of them: “‘S.
Cerro Hueco ..., Chis.,” “10. Presidio,
Ver.,” and “11. El Castillo, .. . ,Oax.” These
were designated the type localities of Pro-
cambarus mirandai, P. veracruzanus, and
P. acanthophorus, respectively, in his 1954
publication and cited as such in 1955 and
1983. Perhaps specimens from some of the
other localities not subsequently mentioned
by Villalobos are referable to P. mexicanus.
If they are still extant they will likely be
reported by Sr. Miguel A. Morales Mora
who is currently studying the Mexican rep-
resentatives of the subgenus Austrocamba-
rus.
*““Cambarus aztecus Saussure”’
Fig. 2
Syntypic male, form I (dry): Cephalotho-
rax (Fig. 2b, e) subovoate; maximum width
of carapace greater than height at level of
caudodorsal margin of cervical groove (11.1
and 9.1 mm). Abdomen narrower than tho-
rax (9.7 and 11.1 mm). Cephalic section of
carapace about twice as long as areola; latter
10.3 times longer than broad with 1 or 2
punctations across narrowest part. Surface
of carapace densely punctate dorsally, be-
coming finely granulate ventrolaterally.
Rostrum comparatively broad with mar-
gins not thickened and gently converging
anteriorly to base of acumen where armed
with minute corneous tubercles, otherwise
not produced; upturned, corneous tip of
acumen almost reaching distal end of ulti-
mate podomere of antennular peduncle;
dorsal surface of rostrum subplane basally,
gradually becoming weakly excavate ante-
riorly, and bearing setiferous punctations
VOLUME 100, NUMBER 1
207
Fig. 2. Cambarus aztecus, syntypic male, form I: a, Dorsal view of distal podomeres of cheliped; b, Dorsal
view of carapace; c, Mesial view of first pleopod; d, Proximal podomeres of third, fourth, and fifth pereiopods;
e, Lateral view of carapace; f, Caudal view of first pleopod; g, Lateral view of first pleopod; h, Epistome.
which deeper and more abundant poste-
riorly than anteriorly. Subrostral ridge weak
and evident dorsally only along about basal
sixth of rostrum. Postorbital ridge well de-
fined, moderately swollen caudally, and ter-
minating anteriorly in small spine not
reaching caudal margin of orbit. Suborbital
angle weak and obtuse; branchiostegal spine
small but clearly defined.
Abdomen longer than carapace but can-
not be measured accurately because bent.
Pleura of third through fifth segments very
weakly arched and rounded caudoventrally.
Cephalic section of telson with 3 spines in
each caudolateral corner (middle one of each
group movable). Cephalic lobe of epistome
(Fig. 2h) subtriangular with anterolateral
margins slightly thickened and very weakly
arched; main body with clearly defined fo-
vea; epistomal zygoma broadly arched.
Ventral surface of proximal podomere of
antennular peduncle with strong spine near
midlength. Antennal peduncle with disto-
lateral spine on basis; ischium with small
subacute tubercle; (flagellum lost). Antennal
scale about twice as long as broad, widest
slightly distal to midlength, greatest width
of lamellar area about twice that of thick-
ened lateral part. Third maxilliped slightly
overreaching antennal peduncle; mesial sec-
tor of ventral surface of ischium bearing
dense clusters of stiff, simple setae, and more
delicate plumose ones; lateral sector with
shorter and fewer stiff setae; ischium not
produced distolaterally.
Left chela (Fig. 2a), right regenerated,
subovate in cross section, weakly depressed;
palm 1.7 times as long as broad, its length
almost half maximum length of chela; palm
and proximal part of fingers studded with
crowded small tubercles (median longitu-
dinal ridges polished, and distal two-thirds
of fingers with setiferous punctations). Me-
sial surface of palm with tubercles arranged
irregularly, but 2 or 3 ill-defined rows, con-
sisting of 10 or 11 in each, discernible be-
208
tween proximal and distal margins. Dorsal
and ventral surfaces of both fingers with low
median longitudinal ridges flanked along
distal two-thirds by setiferous punctations.
Opposable margin of fixed finger with row
of 5 tubercles (proximalmost largest) ex-
tending along slightly more than proximal
half of finger; broad longitudinal band of
minute denticles extending from base to
corneous tip of finger, and single massive
tubercle located below band at distal end of
proximal third of finger; lateral surface with
2 tubercles basally followed by row of con-
spicuous setiferous punctations. Opposable
margin of dactyl with row of 7 tubercles
(second from base largest; most too small
to be included in illustration) reaching level
of row on fixed finger; broad band of minute
denticles and single large tubercle situated
below band just proximal to midlength of
finger; corresponding margin of right chela
with only 2 distinct tubercles: one above
denticular band at end of proximal sixth of
margin, and that below, slightly more dis-
tally.
Carpus of cheliped longer than broad with
very shallow elongate depression; surface
almost entirely tuberculate except proxi-
moventrally, only those tubercles on mesial
surface larger than tubercles elsewhere, and
distal members there subspiniform; ven-
trodistal margin with prominent tubercle at
articular knob and smaller one mesial to it.
Merus with all surfaces tuberculate except
proximal half to fourth of mesial and lateral
surfaces; ventral surface with mesial row of
13 (14 on right) tubercles and lateral row of
8. Ischium with ventromesial row of 7 (5
on right) small tubercles.
Hook on ischium of third pereiopod (Fig.
2d) simple, acute, and reaching to, but not
overreaching, basioischial articulation, and
not opposed by tubercle on basis. Coxa of
fourth pereiopod without caudomesial boss,
but that of fifth with compressed (in lon-
gitudinal plane of body) boss at caudomesial
angle.
Sternum between third, fourth, and fifth
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
pereiopods comparatively shallow, mar-
gined with plumose setae.
First pleopod (Fig. 2c, f, g) reaching coxa
of third pereiopod; cephalic process want-
ing; mesial process and central projection
unremarkable, shoulder on cephalic surface
with cephalodistal extremity subtruncate,
only slightly produced distally.
Uropods with both proximal lobes bear-
ing acute spines; mesial ramus with disto-
median spine situated premarginally and
with well developed distolateral spine; lat-
eral ramus with fixed distolateral spine
flanked mesially by strong movable spine.
See Table 1 for measurements.
Procambarus (Austrocambarus)
olmecorum, new species
Fig. 3
Procambarus aztecus.— Rioja, 1949:321.—
Villalobos, 1954:306, 312, 314 Gn part)—
321, 323, 328, pls. 3, 4; 1955:160, 161,
169 (in part)—-176, 178, 183, pls. 36, 37;
1983:154, 155, 164-170, 171, 176, 227,
pls. 36, 37.— Hobbs and Villalobos, 1964:
313.—Hobbs, 1966:71; 1971:3, 12, 22,
27, 30, 31, 41.—Holt, 1973:4, 24, 25.—
Hart and Hart, 1974:22, 23, 86.
Procambarus mexicanus.— Rioja, 1949:321
[part: Costomatepec, Ver.]
Procambarus (Austrocambarus) aztecus. —
Villalobos, 1982:219.
All of the above synonyms are traceable
to the misidentification of crayfish from To-
matlan, first cited as hosts of the entocy-
therid ostracod Ankylocythere bidentata by
Rioja (1949:321) and later fully described
and illustrated by Villalobos (1954). Hobbs
(1966), in summarizing our current knowl-
edge of the entocytherids of Mexico, ac-
cepted the previously reported identifica-
tion of the host as did Hart and Hart (1974).
In Holt’s (1973) account of the Mexican
branchiobdellid worms, he employed the
identifications furnished by Villalobos.
After having compared Villalobos’ de-
scription and illustrations with syntypes of
VOLUME 100, NUMBER 1 209
Fig. 3. Procambarus (A.) olmecorum (a, b, f-1, n from holotype; c, e from morphotype; d, m from allotype):
a, Lateral view of carapace; b, c, Mesial view of first pleopod; d, Annulus ventralis and adjacent sternal elements;
e, f, Lateral view of first pleopod; g, Epistome; h, Caudal view of first pleopods; i, Antennal scale; j, Cephalomesial
view of first pleopod; k, Dorsal view of carapace; 1, Proximal podomeres of third, fourth, and fifth pereiopods;
m, n, Dorsal view of distal podomeres of cheliped.
210
Cambarus aztecus Saussure, Hobbs (1972b:
44) pointed out that “‘specimens described
as Procambarus aztecus by Villalobos (1954:
44) are not members of this [Saussure’s]
species and must receive a new name.”’ The
name o/mecorum, honoring the artistic Ol-
mec nation which in earlier times contrib-
uted so much to Mexican culture, is offered
as a substitute name for Villalobos’ Pro-
cambarus aztecus.
Diagnosis. —Body pigmented, eyes well
developed. Rostrum of adults without mar-
ginal spine and median carina. Carapace
with small cervical spine. Areola linear or
obliterated along part of its length and con-
stituting 30.5 to 35.7 (average 33.8) percent
of total length of carapace (38.9 to 45.5,
average 41.2 percent of postorbital carapace
length). Suborbital angle weak and obtuse;
infraorbital spines lacking. Postorbital ridge
moderately strong and anterior extremity
with or without spine or tubercle. Bran-
chiostegal spine small. Antennal scale about
twice as long as broad, widest slightly distal
to midlength. Cheliped studded with squa-
mous tubercles from midlength of merus to
midlength of fingers. Ischium of third pe-
reiopod of first form male with simple,
strong, acute hook overreaching basioischi-
al articulation; hook not opposed by tuber-
cle on corresponding basis. First pleopods
of first form male reaching coxae of third
pereiopods, symmetrical, contiguous basal-
ly, lacking proximomesial spur; subangular
shoulder present on cephalic surface at about
base of distal eighth; lacking subterminal
setae; terminal elements consisting of short,
acute, distolaterally directed mesial process
extending beyond short, corneous, acute,
cephalodistally directed central projection.
Female with hinged annulus ventralis only
slightly longer than well developed prean-
nular plate, about twice as wide as long,
broadly arched posteriorly, and bearing short
sinuous sinus on midposterior surface; post-
annular sclerite as wide as annulus and only
slightly shorter; first pleopod greatly re-
duced.
Holotypic male, form I: Cephalothorax
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
(Fig. 3a, k) subovate, weakly compressed
laterally; maximum width of carapace
slightly greater than height at level of cau-
dodorsal margin of cervical groove (12.9 and
12.0 mm). Abdomen narrower than thorax
(11.0 and 12.9 mm). Areola linear. Cephalic
section of carapace about 1.9 times as long
as areola, latter constituting 34 percent of
total length of carapace (42.2 percent of
postorbital carapace length). Surface of car-
apace densely punctate dorsally and finely
tuberculate laterally, only on anterior half
of rostrum sparsely punctate. Rostrum
comparatively broad with weakly conver-
gent margins, tapering rather strongly an-
teriorly but base of acumen not clearly de-
fined, apex slightly upturned and reaching
midlength of ultimate podomere of anten-
nular peduncle; margins not thickened; up-
per surface very weakly concave, almost flat.
Subrostral ridge weak and not evident in
dorsal aspect except along caudal margin of
orbit. Postorbital ridge moderately strong,
only faintly swollen caudally, and termi-
nating anteriorly in small spine not attain-
ing level of orbit. Suborbital angle weak and
obtuse; branchiostegal spine very small but
acute. Cervical spine small but distinct.
Abdomen longer than carapace (25.8 and
21.8 mm). Pleura of third through fifth seg-
ments weakly arched and rounded caudo-
ventrally. Cephalic section of telson with 3
spines (middle one of group movable) in
each caudolateral corner. Cephalic lobe of
epistome (Fig. 3g) subtriangular with slight-
ly excavate anterior angle, margins not
thickened, main body with distinct fovea;
epistomal zygoma arched. Ventral surface
of proximal podomere of antennular pe-
duncle with spine near midlength. Antennal
peduncle with short distolateral spine on
basis; ischium without spine or tubercle; fla-
gellum broken but reaching second abdom-
inal segment. Antennal scale (Fig. 31) about
twice as long as broad, widest slightly distal
to midlength, greatest width of lamellar area
about twice that of thickened lateral part.
Third maxilliped overreaching antennal pe-
duncle by length of ultimate podomere; me-
VOLUME 100, NUMBER 1
sial sector of ventral surface of ischium with
crowded clusters of both simple stiff and
more flexible plumose setae, lateral sector
studded with mat of plumose setae, ischium
not produced distolaterally.
Right chela (Fig. 3n) subovate in cross-
section, weakly depressed; palm almost 1.2
times as long as broad; its mesial length
almost half maximum length of chela; ex-
cept for ridges and apices of fingers, almost
entire surface studded with closely set squa-
mous tubercles. Mesial surface of palm with
3 irregular rows of about 8 tubercles, others
interspersed between rows. Both fingers with
low median longitudinal ridges dorsally and
ventrally; ridges flanked by tubercles except
distally where replaced by setiferous punc-
tations. Opposable margin of fixed finger
with row of 13 (left with 14) tubercles (more
distal ones too small to be included in il-
lustration), second from base largest, ex-
tending along proximal four-fifths of finger,
and row of 3 (2 on left) tubercles, proxi-
malmost largest, on lower level along mid-
dle fifth of finger; minute denticles present
between rows of tubercles and reaching cor-
neous tip of finger. Opposable margin of
dactyl bearing row of 12 (left with 13) tu-
bercles, third from base largest and proxi-
mal 2 (0 on left) fused; narrow band of mi-
nute denticles interspersed between tubercles
and extending to base of corneous tip of
finger, opposing those on fixed finger.
Carpus of cheliped longer than broad with
shallow furrow dorsally; furrow flanked by
squamous tubercles; entire podomere stud-
ded with similar tubercles, one or 2 on me-
sial surface and another on ventrodistal
margin slightly more elevated than others
but none conspicuously larger than others.
Merus with all surfaces tuberculate except
proximal fourth of mesial and lateral sur-
faces; dorsal surface with tubercles increas-
ing in size distally; ventral surface with me-
sial row of 12 tubercles and irregular lateral
row of 14 (left with 13). Ischium with row
of 3 (left with 5) tubercles ventromesially.
Hook on ischium of third pereiopod (Fig.
31) simple, acute, and overreaching basiois-
214
chial articulation, not opposed by tubercle
on basis. Coxa of fourth pereiopod without
caudomesial boss, but that of fifth with small
tuberculiform one at caudomesial angle.
Sternum between third, fourth, and fifth
pereiopods shallow, margined with plu-
mose setae.
First pleopods (Fig. 3b, f, h, }) as described
in “‘Diagnosis.”
Uropods with both lobes of proximal
podomere bearing acute spines; mesial ra-
mus with distomedian spine situated dis-
tinctly premarginally and with well devel-
oped distolateral spine; lateral ramus with
fixed distolateral spine flanked mesially by
very strong movable spine.
Allotypic female: Differing from holo-
type, other than in secondary sexual fea-
tures, in following respects: maximum width
of carapace distinctly greater than height;
base of acumen clearly defined by suddenly
contracted rostral margins; tip of rostrum
more strongly upturned and reaching base
of distal third of ultimate podomere of an-
tennular peduncle; postorbital spine reach-
ing level of posterior margin of orbit; ab-
domen only slightly shorter than carapace;
pleuron of fifth abdominal segment with
ventral margin straight, not arched; anterior
margin of epistome rather evenly and
strongly arched, and zygoma broadly so; is-
chium of antennal peduncle with very small,
acute, tuberculiform spine ventrally; third
maxilliped almost reaching distal end of an-
tennal peduncle.
In addition, chela (Fig. 3m) much smaller
than that of holotype with width of palm
slightly greater than length of mesial mar-
gin, latter much less than half length of che-
la; opposable margin of fixed finger with row
of 6 tubercles (most too small to be evident
in illustration), basal one largest, 3 in prox-
imal group separated from those in distal
group by considerable gap in which | tu-
bercle situated on lower level at about mid-
length of finger; opposable margin of dactyl
with row of 9 tubercles (small and hidden
by setae), proximal 3 subequal in size and
larger than more distal ones; carpus with 2
ZZ
Table 2.— Measurements (mm) of Procambarus (A.)
olmecorum.
Morpho-
Holotype Allotype type
Carapace:
Entire length 27.0 DISD 20.4
Postorbital length 21.8 20.3 16.3
Width 12.9 12.1 10.8
Height 12.0 12.1 10.0
Areola:
Width linear linear linear
Length 2 8.8 7.0
Rostrum:
Width 4.4 4.0 3:5
Length 6.0 5.6 4.2
Right chela:
Length, palm me-
sial margin 7.8 4.7 4.1
Palm width 6.7 5.0 4.2
Length, lateral
margin 17.9 12.0 10.5
Dactyl length 9.8 7.0 60)
Abdomen:
Width 11.0 10.8 8.5
Length 25.8 25.0 20.6
distal tubercles on mesial surface and | on
distal ventrolateral articular condyle dis-
tinctly spiniform and slightly larger than
neighboring tubercles; ventral surface of
merus with row of 11 tubercles mesially and
5 or 7 (left) laterally.
Sternum between third, fourth, and fifth
pereiopods shallow. Annulus ventralis and
associated sternites as figured (Fig. 3d) and
described in “‘Diagnosis.”’
Morphotypic male, form II: Differing
from holotype in following respects: base of
acumen more sharply defined than in ho-
lotype although acumen broken, probably
reaching anteriorly at least as far as mid-
length of ultimate podomere of antennular
peduncle; postorbital ridge lacking spine at
anterior extremity and not attaining level of
orbit; suborbital angle more distinctly an-
gular than in holotype; branchiostegal spine
obsolete; cephalic lobe of epistome and zy-
goma as in allotype; third maxilliped just
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
reaching utimate podomere of antennular
peduncle; tubercles and spines on chelipeds
only slightly different from those of holo-
type; hook of ischium of third pereiopod
much reduced, and coxa of fifth pereiopod
bearing only slightly reduced boss.
First pleopods (Fig. 3c, e) differing only
slightly from that of first form male, differ-
ing chiefly in the absence of cornified tip on
central projection and both terminal ele-
ments less acute.
Type locality.— Arroyo to Rio Metlac near
‘“‘edge’’ of Fortin de Las Flores, Veracruz,
Mexico.
Disposition of types. —The holotypic male,
form I, allotype, and morphotype are de-
posited in the National Museum of Natural
History (Smithsonian Institution) nos.
217626, 217627, and 217628, respectively,
as are the paratypes consisting of 1 ¢I, 1 36
II, 5 2, and 1j 2°. These specimens constitute
the type series.
Size. —The largest specimen available is
a female from Cuitahuac having a carapace
length of 36.4 mm (postorbital carapace
length 30.0 mm); the holotype (see Table 2)
is the smallest of the three first form males.
Females carrying eggs or young are un-
known.
Range and specimens examined. —Pro-
cambarus (A.) olmecorum 1s known to occur
in only six localities, all in the state of Ve-
racruz, Mexico: (1) type locality, 3 61, 2 6
II, 6 2, 1) 2 (USNM), 13 Mar 1974, R. R.
Miller et al. (2) Cuitahuac, 2 9 (USNM), 10
Aug 1964, P. J. Spangler. (3) Nacimiento
de Manzanilla, 17.6 km N of Potrero, 1 6 I
(USNM), 6 Jan 1977, J. R. Reddell & A.
G. Grubbs. (4) Tomatlan, 14 km SSW of
Huatusco (IBM?) (Villalobos, 1954:321;
1955:176; 1983:168). (5) 3 km S of Cos-
comatepec (IBM?) (Villalobos 1954:321;
1955:176; 1983:168). (6) Sinkhole on south
side of highway between Jalapa and Vera-
cruz (Hobbs 1971:30), 3 61, 1601, 3 2, lj
6, 3} 2 (USNM), 19 Dec 1966, R. R. M. &
W. L. Minckley. These lie at altitudes be-
tween 800 and 1600 m in the drainage ba-
sins of Rio Atoyac and Rio Jamapa, which
VOLUME 100, NUMBER 1
unite before entering the Gulf of Mexico a
few km south of the city of Veracruz.
Variations. —The rostrum reaches ante-
riorly from the base to the end of the ulti-
mate podomere of the antennular peduncle,
and the margins vary from being distinctly
subangular at the base of the acumen to
tapering with little interruption from their
bases; the postorbital ridges terminate an-
teriorly either in a short spine or tubercle;
the areola is either obliterated along part of
its length or is linear; and the cervical spine
although never conspicuous may be rather
well developed or reduced to a tubercle
scarcely larger than others nearby. The tel-
son is also variable, sometimes subrectan-
gular, but often with the posterior section
somewhat tapering. As in all members of
Austrocambarus, the chelipeds are studded
with crowded tubercles, but the numbers
and disposition exhibit conspicuous indi-
vidual differences.
The most distinctive of the populations
assigned to this species is that from the sixth
locality cited above. In the specimens from
there, the rostral margins converge little an-
teriorly to the base of the acumen where
they turn sharply mesially forming distinct
angles before tapering to the apex of the
acumen which just reaches, or only slightly
overreaches, the base of the distal podomere
of the antennular peduncle. The chelipeds
in one of the males are conspicuously slen-
der. Too, the shoulder on the cephalic sur-
face of the first pleopod of two of the first
form males is more strongly produced dis-
tally and more acute than that of the ho-
lotype, but in the other it is only little more
prominent.
Relationships. —Procambarus (A.) olme-
corum has its closest affinities with Procam-
barus (A.) mexicanus, P. (A.) veracruzanus,
and P. (A.) ruthveni zapoapensis. As in
mexicanus, the rostrum lacks marginal
spines, but the areola is distinctly much nar-
rower, similar to that of veracruzanus and
ruthveni zapoapensis, but in them the ros-
tral margins bear spines or spiniform tu-
bercles at the base of the acumen. The de-
213
velopment of the shoulder on the cephalic
margin of the pleopod of the first form male
is variable, ranging from being almost trun-
cate to being produced cephalodistally in
acute prominences but never sloping proxi-
mally as in ruthveni zapoapensis. Among
the members of the subgenus Austrocam-
barus, the unique combination of characters
exhibited by P. (A.) olmecorum 1s: rostrum
without produced anterolateral margins; ar-
eola linear or obliterated along part of its
length; and first pleopod of first form male
with subangular shoulder subtruncate or
produced distally but never sloping proxi-
mally.
Acknowledgments
I wish to acknowledge the cooperation
and help given me by the late Dr. Alejandro
Villalobos-Figueroa and to extend my ap-
preciation to those (see the paragraphs de-
voted to “Range and Specimens exam-
ined’’?) who donated specimens to the
Smithsonian. I am also grateful to T. E.
Bowman, a Smithsonian colleague, to Jo-
seph F. Fitzpatrick, Jr., of the University of
South Alabama, and to H. H. Hobbs III of
Wittenberg University, for their criticisms
of the manuscript.
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van Straelen, Victor. 1942. A propos de la distribu-
tion des écrevisses, des homards et des crabes
d’eau douce.— Bulletin du Musée Royal d’His-
toire Naturelle de Belgique 18(56):1-11.
Villalobos Figueroa, Alejandro. 1946. Estudios de los
cambarinos mexicanos, IV. Consideraciones
acerca de la posicion sistematica y redescripcion
de Procambarus digueti (Ortmann). — Anales del
Instituto de Biologia, Universidad Nacional
Autonoma de México 17(1, 2):215—230, 2 plates.
. 1948. Estudios de los cambarinos mexicanos,
VII. Descripcion de una nueva especie del gé-
nero Procambarus, Procambarus acanthopho-
rus, n. sp.—Anales del Instituto de Biologia,
Universidad Nacional Autonoma de México
19(1):175-182, 2 plates.
1950. Contribucion al estudio de los cam-
barinos mexicanos, IX. Estudio taxonomico de
un grupo de especies del género Procambarus. —
Anales del Instituto de Biologia, Universidad
Nacional Autonoma de México 21(2):367—413,
11 plates.
1953. Distribucion geografica y notas eco-
logicas de los cambarinos mexicanos. Memoria
del Congreso Cientifico Mexicano, VII. Ciencias
Biologicas, IV Centenario de la Universidad de
México 7:343-374, 5 figs.
. 1954. Estudios de los cambarinos mexicanos,
XII, Parte 1. Revision de las especies afines a
Procambarus mexicanus (Erichson), con de-
scripcion de nuevas formas.—Anales del Insti-
tuto de Biologia, Universidad Nacional Auto-
noma de México 25(1, 2): 299-379, 19 plates.
1955. Cambarinos de la fauna mexicana.
(Crustacea Decapoda). xvi + 290 pages, 62
plates. Tesis, Facultad de Ciencias. Universidad
Nacional Autonoma de México.
1982. Decapoda. Pp. 215-239 in Stuart H.
Hurlbert and Alejandro Villalobos-Figueroa, eds.
Aquatic Biota of Mexico, Central America and
the West Indies. San Diego, California: Aquatic
Biota SDSU Foundation.
1983. Crayfishes of Mexico (Crustacea: De-
capoda). xii + 276 pages, 62 plates. Washington:
Smithsonian Institution Libraries and the Na-
tional Science Foundation. [Translation of Vil-
lalobos 1955 by Horton H. Hobbs, Jr.]
Department of Invertebrate Zoology, Na-
tional Museum of Natural History, Smith-
sonian Institution, Washington, D.C. 20560.
PROC. BIOL. SOC. WASH.
100(1), 1987, pp. 216-247
NEW RECORDS OF ISOPOD CRUSTACEA FROM THE
CARIBBEAN, THE FLORIDA KEYS,
AND THE BAHAMAS
Brian Kensley and Marilyn Schotte
Abstract. —Fourteen species of marine isopods are recorded from Belize, the
Bahamas, and the Florida Keys. These include a new genus and species of
anthuridean, Licranthura amyle, resembling the genera Eisothistos and Stel-
lanthura, and characterized mainly by the presence of antler-like processes on
the antennae, and the following new species: Mesanthura looensis (Anthuridae),
Phycolimnoria clarkae (Limnoriidae) (apparently the first record of the genus
found in decaying wood), Cirolana albidoida, Cirolana crenulitelson (Cirolan-
idae), Ancinus belizensis, and Cassidinidea mosaica (Sphaeromatidae). Six
species of Limnoria are recorded as co-occurring: L. multipunctata, L. pfefferi,
L. platycauda, L. tuberculata, L. indica, L. unicornis. Of these, the latter two
are recorded from the Caribbean for the first time. Cirolana minuta Hansen
is redescribed, this being the second record of the species.
Since the appearance of the work by Men-
zies and Glynn (1968) on the isopod fauna
of Puerto Rico (and which summarized
knowledge of the Caribbean isopodan fau-
na), several papers have appeared dealing
with aspects of the marine isopod fauna of
the area. Careful, small-scale collecting aid-
ed by SCUBA have added considerably to
the Caribbean faunal list (e.g., Kensley 1982,
1984), and will no doubt continue to do so,
as witness this present paper. The surprising
and unsuspected richness of the isopod fau-
na is well illustrated by the growing number
of new species and records coming from a
relatively small area around the Smithson-
ian Institution’s field laboratory on Carrie
Bow Cay, Belize.
Unless otherwise indicated, all material
dealt with in this paper was collected by the
authors.
Family Anthuridae
Licranthura, new genus
Diagnosis. —Pleonites 1-5 short, free.
Telson lacking statocysts. Tailfan cup-like,
expanded, margins serrate, uropodal exo-
pod with spike-like lobe. Antennular fla-
gellum of 3 articles, terminal article with 2
aesthetascs. Antennal peduncle article 3 with
serrate triangular process, flagellum of 6 ar-
ticles. Mandible lacking molar process.
Maxilliped palp of 5 articles. Pleopod 1, rami
separate, together forming operculum. Pe-
reopod 1, propodus somewhat enlarged,
larger than pereopods 2 and 3.
Remarks. —The lack of statocysts and a
mandibular molar, the spike-like lobe on
the uropodal exopod, the serrate tailfan
margins, the free anterior five pleonites, the
non-subchelate pereopod 1, the poorly de-
veloped opercular pleopod | with both rami
contributing, all point to a similarity to two
genera, Eisothistos Haswell, 1884, and Stel-
lanthura Wagele, 1979. Licranthura differs
from both in several aspects. Eisothistos
possesses fused rami of pleopod 1, lacks a
-mandibular palp, and has pereopod 1 of
about the same size as pereopods 2 and 3.
Stellanthura possesses a mandibular palp,
a reduced molar, and four articles in the
maxillipedal palp. Neither of these genera
possess the unusual serrate lobe of the third
VOLUME 100, NUMBER 1
peduncular article of the antenna. The func-
tion of these structures can only be specu-
lated on: possibly it is related to some feed-
ing behavior. The splayed and serrate tailfan
would suggest that this species also preys on
some tubiculous organism as does Eiso-
thistos.
Etymology.—The generic name is de-
rived from the Greek J/ikros, an antler (re-
ferring to the structures on antennae), plus
the commonly-used sufhx “‘anthura.”’
Licranthura amyle, new species
Figs. 1, 2 :
Material. HOLOTYPE, USNM 211408,
2 tl 3.2 mm; sta H-15, Carrie Bow Cay,
Belize, from Agaricia coral in forereef crest,
15 m, 20 Apr 1981, coll. G. Hendler.
PARATYPES, USNM 211409, ¢ tl 3.8
mm; sta H-1, Twin Cays, Belize, Caulerpa
alga around red mangrove roots, 0-2 m, 17
Apr 1981, coll. G. Hendler.—USNM
211410, manca tl 1.9 mm; sta H-7, off
Glover’s Reef, Belize, 15—21 m, 24 Mar
1980, coll. G. Hendler.—USNM 211411, 2
tl 2.8 mm; sta H-8, Carrie Bow Cay, Belize,
from Madracis coral on forereef crest, 15.2
m, 19 Apr 1981, coll. G. Hendler. —USNM
211412, 3 2 tl 3.1, 3.5, 3.6 mm; sta H-11,
Carrie Bow Cay, Belize, from Madracis cor-
al on forereef crest, 15.2 m, 19 Apr 1981,
coll. G. Hendler.—USNM 211412, 2 tl 2.8
mm; sta H-16, Carrie Bow Cay, Belize, from
Agaricia coral on forereef crest, 15.2 m, 20
Apr 1981, coll. G. Hendler.—USNM
211414, 2 tl 3.0 mm; sta H-41, Carrie Bow
Cay, Belize, from Agaricia on reef platform,
1—2 m, 25 Apr 1981, coll. G. Hendler. —
USNM 211415, manca tl 1.9 mm; sta H-49,
Carrie Bow Cay, Belize, from Porites coral
on reef flat, 1-2 m, 26 Apr 1981, coll. G.
Hendler.—USNM 211416, 2 2 tl 2.5, 2.8
mm; sta H-85, Carrie Bow Cay, Belize, from
coral rubble and algae on forereef dropoff,
24.4-27.4 m, 21 Apr 1981, coll. G. Hend-
ler.—USNM 211417, 22 t1 2.8, 3.0 mm; sta
K-101, Carrie Bow Cay, Belize, coral rubble
De)
near drop-off, 25 m, 14 Apr 1981.—USNM
211418, 2 tl 3.8 mm; sta K-142, Carrie Bow
Cay, Belize, lower spur and groove zone, 11
m, 5 Dec 1982.—USNM 211419, @ tl 3.9
mm; sta K-162, between Carrie Bow Cay
and South Water Cay, Belize, coral rubble
from patch reef area, 6-8 m, 27 Nov 1985.
Description. —Female: Body slender,
about 12 times longer than wide, widest at
pereonite 1. Faint diffuse brown pigment
most noticeable on dorsolateral margins of
somites; cephalon with faint dorsal reticu-
lation. Body proportions: C < 1 > 2=3 <
4=5> 6 > 7. Pleonites short, similar in
length. Telson thin, dorsally faintly con-
cave, basally narrow, widening posteriorly
to truncate or faintly bilobed serrate pos-
terior margin.
Cephalon with small well pigmented eyes. |
Antennular peduncle of 3 articles, basal ar-
ticle longest and widest; flagellum of 3 ar-
ticles, article 2 about 5 times length of ter-
minal article, latter bearing 2 aesthetascs.
Antennal peduncle with article 3 produced
mediodistally into triangular process bear-
ing 6 serrations on medial margin, process
reaching distally beyond article 4, latter with
medial margin flexed, resulting in article 5
and flagellum being borne at right angle to
axis of body; flagellum of 6 setose articles.
Mandibular palp of 3 articles, article 2 three
times length of article 3, latter bearing 2
fringed spines distally; incisor of 3 blunt
cusps directed anteriorly; lamina dentata of
5 marginal serrations; molar absent. Max-
illa bearing 7 distal spines. Maxilliped with
short endite tipped with 2 setules; palp of 5
articles, basal article very short, terminal
article small, bearing 4 elongate setae.
Pereopod 1 larger than pereopods 2 and
3; dactylus articulating distally on propo-
dus, with strong sclerotized unguis having
smaller squat spine, thin-walled elongate
spine, and 4 slender setae at its base; pos-
terior margin having finely fringed ridges;
propodus basally slightly broader than dis-
tally, having 2 fringed spines anterodistally
and few elongate setae on posterior margin;
218 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
L
Ce ea
=
G
Fig. 1. Licranthura amyle, non-ovigerous ?: A, Antennule and antenna; B, Antennal process enlarged; C,
Maxilla; D, Mandible; E, Maxilliped; F, Uropod and telson; G, Pleopod 1.
VOLUME 100, NUMBER 1
¢
fal a3
ly
eee
yo
KL
219
Fig. 2. Licranthura amyle, non-ovigerous 2: A, Pereopod 1; B, Pereopod 7; C, Pereopod 2.
carpus short, with posterior margin longer
than anterior; merus, ischium, and basis
broad squat articles; basis having 4 or 5
broad serrations posterodistally. Pereopod
2, unguis about half length of remainder of
dactylus, with small accessory spine at base;
hook-like tooth present on posterior margin
near articulation with propodus; latter
roughly rectangular, with squat dentate spine
posterodistally. Pereopods 5-7, dactylus
with 2 basal serrations; propodus rectan-
gular, with strong dentate posterodistal
spine, 2 anterodistal fringed spines; carpus
with posterior margin longer than anterior
220
margin, bearing strong dentate posterodistal
spine.
Pleopods essentially similar; pleopod 1
only slightly larger than following pleopods,
rami both contributing to opercular func-
tion, of equal length, endopod about 7 width
of exopod. Uropodal exopod widening dis-
tally, outer (free) margin serrate, ending in
outer narrowly triangular acute lobe and
more broadly rounded inner lobe; endopod
ovate, with serrate margin.
Etymology.—The specific name is de-
rived from the Greek myle, a grinder, and
refers to the lack of a molar in the mandible
of this species.
Mesanthura looensis, new species
Fig. 3
Material.—HOLOTYPE, USNM 211406,
non-ovig. 2, tl 10.0 mm; sta FLK-22, Looe
Key, Florida, sandy bottom in gorgonacean
bed with scattered sponges and corals, | m,
26 Jan 1983.
PARATYPES, USNM 211407, 2 non-
ovig. 2, tl 7.0, 7.0 mm. Same data as ho-
lotype.
Description. —Female: Body proportions,
C<1=22>3 < 4=5=62>7= PI.
Pleonite 6 fused with telson ventrally; mid-
dorsally, posterior margin having tiny
emargination. Telson thin, dorsally slightly
concave, outline evenly elongate-oval.
Antennular flagellum of 4 articles, 2 distal
articles each with 2 aesthetascs. Antennal
flagellum of 5 setose articles. Mandibular
palp of 3 articles, article 2 bearing fringed
scales; article 3 with 8 fringed spines; lamina
dentata of 4 serrations; molar thin-walled,
truncate. Maxilliped lacking endite; palp of
3 articles, terminal article semicircular in
outline, with 2 fringed setae and 3 simple
setae on medial margin.
Pereopod | subchelate, unguis slightly less
than half length of remainder of dactyl;
propodus expanded, palm with proximal
rounded scale-bearing lobe armed with 6
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
setae; carpus triangular, distally having
rounded scale-bearing lobe carrying 5 sim-
ple setae. Pereopod 2, propodus rectangu-
lar, not expanded or subchelate, with serrate
sensory spine at posterodistal angle; carpus
short, triangular. Posterior pereopods,
propodus somewhat curved, with postero-
distal series of short setae and strong serrate
sensory spine at posterodistal angle; carpus
with posterior margin longer than anterior,
sensory spine at posterodistal angle.
Pleopod 1, exopod operculiform, slightly
longer and 3 times wider than endopod;
protopod bearing 5 retinacula. Uropodal
exopod ovate, with outer distal margin sin-
uate; endopod broadly ovate.
Color pattern: dorsum of cephalon, pe-
reon, pleon, telson, and uropods with strong
red-brown pigment; cephalon with broad
patch between eyes, and extending to pos-
terior margin; on pereonites 1-3 pigment
anteriorly broad, posteriorly constricted;
pereonites 4-6 covering most of dorsum,
with oval middorsal unpigmented patch;
pigment more constricted on peronite 7;
covering most of pleon with middorsal open
patch; telson and uropodal rami with patch
in central area.
Remarks.—The present species differs
from the seven previously described species
of Mesanthura mainly in its distinctive dor-
sal pigment pattern. The spination of the
mandibular palp article 3 is the other most
useful character for species separation in
Mesanthura. The series of eight spines in
M. looensis differs from all the other seven
species (M. fasciata Kensley, 1982—4, M.
hopkinsi Hooker, 1985—4, M. paucidens
Menzies & Glynn, 1968—6, M. reticulata
Kensley, 1982—6, M. punctillata Kensley,
1982—7, M. pulchra Barnard, 1925—10,
M. floridensis Menzies & Kruczynski,
1983— 12-13).
Etymology.—The specific name is de-
rived from the type locality, Looe Key, Flor-
ida.
VOLUME 100, NUMBER 1 A13)|
K
Fig. 3. Mesanthura looensis, non-ovigerous 2: A, Whole animal in dorsal view; B, Antennule; C, Antenna;
D, Mandible; E, Maxilla; F, Maxilliped; G, Pereopod 1; H, Pereopod 7; I, Pereopod 2; J, Uropodal exopod; K,
Uropodal endopod and basis; L, Pleopod 1.
222
Family Limnoriidae
Menzies and Glynn (1968), and Menzies
and Kruczynski (1983) together list six
species of limnoriids recorded from the Gulf
of Mexico and the Caribbean. These are
Limnoria platycauda Menzies, 1957, L.
pfefferi Stebbing, 1904, L. simulata Men-
zies, 1957, L. saseboensis Menzies, 1957,
L. tripunctata Menzies, 1951 (=L. tuber-
culata Sowinsky, 1884), and Paralimnoria
andrewsi (Calman, 1910). Kuhne (1975)
records L. tripunctata, L. saseboensis, L.
pfefferi, and L. multipunctata Menzies,
1957, from the Caribbean. From the Twin
Cays and Man o’War Cay areas of Belize,
six species of Limnoria have now been re-
corded, including all the abovementioned
except L. sasehboensis, plus two new records,
viz. L. indica Becker and Kampf, 1959, and
L. unicornis Menzies, 1957. Most of the
material comes from decaying red man-
grove (Rhizophora mangle) wood, with all
except P. multipunctata co-occurring in a
single station on Man o’War Cay.
A single undescribed species of Phyco-
limnoria is recorded from the Bahamas,
rather surprisingly also from decaying man-
grove wood.
The records of L. simulata are treated
with caution, as female L. indica can easily
be confused with the former species. The
question of sexual dimorphism in Limnoria
generally requires closer scrutiny, in light of
such possible confusion.
The dark-green tests of folliculinid ciliate
protozoans were found externally on several
of the Limnoria species, (see Mohr and
LeVeque 1948) and were especially com-
mon on the dorsal pleotelson and ventral
coxae of L. platycauda.
Limnoria (Limnoria) indica
Becker and Kampf, 1959
Fig. 4
Limnoria indica Becker and Kampf, 1958:
3, figs. 2-4.—Kuhne, 1975:546, figs. 3, 4.
Material. -USNM 211422, 56 speci-
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
mens, sta CBC K166, Man o’War Cay, Be-
lize, from submerged red mangrove roots,
0.5 m, 29 Nov 1985.
Remarks.—This species was described
from the east coast of the Indian Peninsula,
while Kushne (1975) records the species from
Hong Kong and Manila, as well as Madras,
India. Becker and Kampf (1959) detail the
sexual dimorphism reflected in the pleotel-
sonic structure; this dimorphism may well
be the source of confusion in the identifi-
cation of such species as L. simulata and L.
quadripunctata from the Caribbean.
Distribution. —Mandapam Camp and
Madras harbor, India; Hong Kong; Manila,
Philippines.
Limnoria (Limnoria) multipunctata
Menzies, 1957
Fig. 5
Material.—USNM 211421, 4 4, 6 ovig.
2, 3 non-ovig. 2, 7 immature, Twin Cays,
Belize, from dead red mangrove wood, coll.
J. Kohlmeyer, 28 Nov 1985.
Remarks. —Kuhne (1975) points out the
variability found in this species, with ma-
terial from Japan having the two pairs of
submedian pleotelsonic tubercles as in the
type material from the Kai Islands, while
material from Puerto Rico and Jamaica lacks
these tubercles. The present material from
Belize also laeks these anterior tubercles,
but does possess a strong middorsal longi-
tudinal pleotelsonic ridge with rounded tu-
bercles in the posterior half.
Distribution. — Puerto Rico; Jamaica; Ko-
niya, Japan; Kai Islands (South Pacific).
Limnoria (Limnoria) pfefferi
Stebbing, 1904
Limnoria (Limnoria) pfefferi: Menzies,
1957:135, fig. 15.—Kuhne, 1975: 547,
fig. 7.
Material. -USNM 221621, 12 speci-
mens, Twin Cays, Belize, under red man-
groves, 24 Oct 1979.—USNM 221623, 7
specimens, sta H-6(80), Glover’s Reef, Be-
VOLUME 100, NUMBER 1 223
3 3 mM
Rae Mies
ee baie
i ae
BAL FRE Tae,
rere
Fig. 4. Limnoria indica: A, Pleotelson 6; B, Pleotelson 2; C, Last pleonite and anterior pleotelson ¢; D,
Pleotelson 4, in lateral view; E, Pleotelsonic surface enlarged; F, Anterior pleotelson 64, enlarged.
lize, O-3 m, 24 Mar 1980, coll. G. Hend- A.Cohen.—USNM 221625, 10 specimens,
ler.—USNM 221624, 4 specimens, sta AC- sta CBC-K166, Man o’War Cay, Belize,
CBC-610B, Carrie Bow Cay, coarse Hali- submerged red mangrove wood and roots,
meda sediment, 1.5 m, 14 Jun 1981, coll. 0.5 m, 29 Nov 1985.—USNM 221626, 2
224
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig. 5. Limnoria multipunctata: A, Pleotelson; B, Pleotelson in oblique-lateral view; C, Pleotelsonic margin;
D, Middorsal tubercles of posterior pleotelson.
specimens, sta H-18(80), Twin Cays, Belize,
under mangroves, 1—2 m, 28 Mar 1980, coll.
G. Hendler.— USNM 221627, 5 specimens,
sta CBC-K45, Twin Cays, Belize, algal mat
under red mangrove roots, 0.1 m, 9 Apr
1979.
Distribution. —Indian Ocean; Florida;
Puerto Rico; Panama; Philippines; New
Guinea.
Limnoria (Limnoria) platycauda
Menzies, 1957
Limnoria (Limnoria) platycauda Menzies,
1957:139, fig. 17.—Ortiz, 1983:7.
Material. -USNM 221622, 23 speci-
mens, Twin Cays, Belize, from red man-
grove wood, 3 m, 28 Nov 1985, coll. J.
Kohlmeyer.—USNM 221628, 38 speci-
mens, sta CBC-K 166, Man o’ War Cay, Be-
lize, submerged red mangrove wood and
roots, 0.5 m, 29 Nov 1985.
Distribution. —Cuba; Puerto Rico to Cu-
racao.
Limnoria (Limnoria) tuberculata
Sowinsky, 1884
Limnoria (Limnoria) tripunctata Menzies,
1951:86, pl. 30; 1957:137, fig. 16.
Limnoria tuberculata Sowinsky, Kussakin,
1979:322, figs. 187-190.
Material. —USNM 221630, 4 specimens,
sta CBC-K166, Man o’War Cay, Belize,
submerged red mangrove wood and roots,
0.5 m, 29 Nov 1985. :
Distribution. — Yellow Sea; Hong Kong;
Rhode Island to Venezuela; Gulf of Mexico;
VOLUME 100, NUMBER 1
Jas)
Fig. 6. Limnoria unicornis: A, Pleotelson; B, Last pleonite and anterior pleotelson; C, Pleotelson in oblique-
lateral view; D, Pleotelsonic tooth enlarged.
Caribbean; Uruguay; California; Hawaii;
Australia; Mediterranean; Black Sea; India;
Ghana.
Limnoria unicornis Menzies, 1957
Fig. 6
Limnoria unicornis Menzies, 1957:173, fig.
32.
Material.—USNM 221631, 6 6, 8 ovig.
2, 3 non-ovig. 2, Pigeon Creek, San Salva-
dor, Bahamas, 22 May 1986, coll. J. Clark. —
USNM 211423, 2 6, sta CBC-K166, Man
o’ War Cay, Belize, submerged red man-
grove wood and roots, 0.5 m, 29 Nov
1985.—USNM 211424, 2 6, 1 non-ovig. 8,
Twin Cays, Belize, from green algal turf un-
der Rhizophora mangle, 0.5 m.—USNM
211425, 9 4, 3 ovig. 2, 6 non-ovig. 2, 7 juvs.,
Ngeruktabel Is., Palau, from anchialine ce-
note, 9 Feb 1985, coll. T. Iliffe and D. Wil-
liams.—USNM 102745, 1 4,12, 1 juv., Baie
de Maroe, Huahine Is., Society Islands, 1
m, 30 Apr 1957, coll. T. Bowman.
Remarks.—This species has been men-
tioned once in the literature, 1.e., in the orig-
226
inal description. For this reason, all the
Smithsonian Institution (USNM) holdings,
both from the Pacific and the Caribbean,
are included here.
Distribution. —Caroline Islands; Society
Islands; Palau; Bahamas; Belize.
Phycolimnoria clarkae, new species
Figs. 7-9
Material.—HOLOTYPE, USNM 211426,
6 4.3 mm, PARATYPES, USNM 211427,
44 8, 18 ovig. 2, 3.3-4.4 mm, 10 non-ovig.
2, 14 juvs., Pigeon Creek, San Salvador, Ba-
hamas, from dead red mangrove wood, coll.
J. Clark, 22 May 1986.
Description.—Dorsal integument, espe-
cially of pleotelson, bearing very fine, im-
bricate, minutely setulose ridges, giving ap-
pearance of fine, regular foveolation.
Pleonite 5 with broad raised middorsal re-
gion, having irregular bumps. Pleotelson
wider than long, with two rounded sub-
median ridges basally, becoming obsolete
posteriorly.
Antennular peduncle of 3 articles, basal
article longest and widest; flagellum con-
sisting of basal article, wider than long,
bearing 4 aesthetascs, and tiny terminal ar-
ticle bearing single aesthetasc and several
setae. Antennal flagellum of 5 setose arti-
cles, basal article equal in length to 4 distal
articles. Mandibular palp of 3 articles, two
basal articles subequal in length; article 2
bearing 3 distal fringed spines; terminal ar-
ticle two-thirds length of article 2, bearing
6 distal fringed spines; spine row of left
mandible of 2 laciniate spines, flanked by 2
short rounded lobes; spine row of right man-
dible of about 9 laciniate spines increasing
in length proximally, plus bilobed distal
process; incisor consisting of roughly tri-
angular, strongly sclerotized, unornament-
ed cusp. Maxilla 1 and 2 as figured. Max-
illipedal endite with single coupling hook,
8 spines on distal margin, all save one bear-
ing fine setules; palpal article 2 longest and
widest; epipod about 3'2 times longer than
basal width, distally narrowly rounded,
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
reaching to base of palp. Pereopod 1, ac-
cessory spine at base of dactylar unguis short,
bidentate; propodus with 2 fringed poster-
odistal spines; propodus, carpus, and merus
each having row of 4 or 5 rounded tuber-
cules on posterior surface. Pereopod 2 with
5 distal articles each having few tubercles
on or near posterior surface; accessory spine
at base of dactylar unguis short, bidentate;
carpus with stout dentate spine at postero-
distal angle; merus with single stout dentate
spine at anterodistal corner. Pereopod 7, ac-
cessory spine of dactyl faintly bidentate;
carpus with several fringed spines of varying
lengths on distal margin; merus with 4
fringed spines on anterodistal margin. Paired
penes on ventrum of pereonite 7. Pleopod
2, copulatory stylet sabre-shaped, articu-
lating slightly proximal to midlength of
median margin of endopod, just reaching
beyond ramus. Uropodal endopod elon-
gate-ovate, about twice longer than wide;
endopod less than half length of exopod,
triangular, tipped with short squat non-
curved spine (claw); basis with row of fringed
setae along outer margin.
Remarks. —This is the first record of the
genus Phycolimnoria from the Caribbean,
and is unusual in that the material was found,
not boring into an alga as is usually the case,
but in decaying red mangrove wood.
The absence of a “rasp” and “‘file”’ struc-
ture on the mandibles, along with the very
unequal size of the uropodal rami, place this
species in the genus Phycolimnoria Men-
zZ1es.
Two features easily separate this species
from the other 11 species of Phycolimnoria
already described. None of these species
possess a uropodal exopod with a short
straight terminal claw. Only P. zinovae Kus-
sakin, 1963, from the Sea of Japan, has a
uropodal exopod in which the terminal claw
is almost straight. This latter species, how-
ever, has a distinctive Y-shaped ridge on
pleonite 5. Phycolimnoria clarkae, with its
raised and rounded central area of pleonite
5, and the two longitudinal submedian
rounded ridges of the pleotelson with no
VOLUME 100, NUMBER 1 227
Fig. 7. Phycolimnoria clarkae: A, Cephalon in ventral view; B, Pleotelson; C, Pleotelson seen from posterior
margin; D, Pleotelsonic integumental surface enlarged; E, F, Uropod.
other ornamentation would also seem to be : ;
Family Cirolanidae
unique in the genus. SY: '
ane oo: Cirolana albidoida, new species
Etymology.—The species is named for Fics. 10-12
Janice Clark of the Department of Inver- a
tebrate Zoology, Smithsonian Institution, Material.—HOLOTYPE, USNM 211419,
who collected the type material. é tl 7.8 mm; sta GB-7, off Lucaya, Grand
228 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig. 8. Phycolimnoria clarkae: A, Left mandible; B, Incisor and spine row of right mandible; C, Antennule;
D, Antenna; E, Maxilla 1; F, Maxilla 2; G, Maxilliped.
Bahama, 180-220 m, from trap baited with imens, 4.6—9.0 mm; same data as holotype.
fish, set for 8 days, 24 May 1981, coll. D. All specimens lack internal organs and mus-
Camp. culature, suggesting that they may have been
PARATYPES, USNM 211420, 55 spec- retained dead in the fish trap for some days.
VOLUME 100, NUMBER 1
229
Fig. 9. Phycolimnoria clarkae: A, Uropod; B, Pereopod 1; C, Pereopod 2; D, Pereopod 7; E, Pleopod 2 3;
F, Penes.
Description. —Male: Body about 3'2 times
longer than wide, widest at pereonite 1. In-
tegument sparsely pitted. Cephalon width
about '3 longer than medial length, with
small rostral point between antennal bases.
Frontal lamina an equilateral pentagon.
Pereonite 1 about 2'2 times length of pe-
reonite 2; pereonites 3—6 subequal in mid-
dorsal length, pereonite 7 slightly shorter.
Coxae of pereonites 2 and 3 rounded, of
230 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig. 10. Cirolana albidoida: A, Holotype in dorsal view; B, Whole animal in lateral view; C, Antennule; D,
Antenna; E, Right mandible; F, Maxilla 1; G, Maxilla 2; H, Maxilliped; I, Pereopod 1; J, Pereopod 2.
pereonite 4 just less than right-angled, of S— 1-2 acutely produced, of pleonite 4 broader,
7 somewhat produced, acute, with oblique subacute; pleonite 5 lacking free lateral mar-
upcurving furrow. Pleonite 1 laterally over- gin.
lapped by pereonite 7; epimera of pleonites Antennule reaching to posterior margin
VOLUME 100, NUMBER 1
231
Fig. 11.
E, Pleotelsonic apex.
of cephalon; two basal peduncular articles
fused, line of fusion faintly visible, article
3% longer, but narrower than basal article;
flagellum of 10 articles, aesthetascs present
on distal 8 articles. Antennal peduncle with
three basal articles short, articles 4 and 5
more elongate, 4 slightly shorter than 5; fla-
gellum of 19 articles. Mandibular palp of 3
articles, article 2 with 5 elongate and about
11 shorter distal spines, article 3 with 18
marginal spines becoming distally progres-
Cirolana albidoida: A, Pereopod 7; B, Uropod; C, Pleotelsonic apex. Cirolana albida: D, Uropod;
sively longer; incisor of 3 cusps; molar bear-
ing 22 teeth. Maxilla 1, inner ramus with 3
stout setae; outer ramus with about 11 spines
on mediodistal margin. Maxilla 2, inner ra-
mus with 3 fringed proximal setae and about
11 simple distal setae; inner lobe of outer
ramus with 9 distal setae, outer lobe with 5
setae. Maxillipedal endite reaching to end
of basal palp article, with 2 coupling hooks
and 5 fringed setae; palp article 3 longest
and widest; articles 2—5 with dense setae on
232 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig. 12. Cirolana albidoida 6: A, Pleopod 1; B, Pleopod 2; C, Pleopod 3; D, Pleopod 4; E, Pleopod 5.
mediodistal margins, sparser setae on outer
margins.
Pereopod 1 robust, dactyl slightly more
than half anterior length of propodus; prop-
odus with stout rounded posterodistal spine
and 2 smaller sensory spines on posterior
margin; carpus triangular, with single sen-
sory spine; merus with 5 stubby rounded
VOLUME 100, NUMBER 1
spines on posterior margin; ischium with 2
acute posterodistal spines. Pereopods 2 and
3 similar, dactylus about 7% length of prop-
odus; latter with rounded posterodistal spine
and 2 smaller sensory spines on posterior
Margin; carpus with 2 posterodistal spines;
merus with 5 large sensory spines on pos-
terior margin, several anterodistal spines of
varying length; ischium with 1 elongate and
2 smaller posterodistal spines, 1 elongate
and 2 small anterodistal spines. Pereopods
4-7 similar, becoming progressively more
elongate; dactylus about '4 length of prop-
odus; latter elongate-cylindrical, with 3 small
spines on posterior margin; carpus about 73
length of propodus, with 2 spines at mid-
length of posterior margin, clump of 4 pos-
terodistal spines, clump of ‘6 anterodistal
spine of which 2 fringed; merus about %
length of carpus, with 2 spines in proximal
half of posterior margin, clump of 6 pos-
terodistal spines, clump of 9 or 10 antero-
distal spines, several of which fringed; is-
chium with several small spines on posterior
margin and clump of 5 anterodistal spines,
2 of which fringed. Pleopods, all rami fringed
with plumose setae; pleopod 1 peduncle
roughly rectangular, with 5 coupling hooks
on median margin, endopod slightly shorter
and narrower than exopod, latter elongate-
ovate; pleopod 2, peduncle roughly rect-
angular, with 4 coupling hooks on medial
margin, copulatory stylet articulating at base
of endopod, slender, tapering reaching by
half its length beyond rami; pleopod 3, pe-
duncle with 4 coupling hooks, exopod long-
er and broader than endopod, with indis-
tinct transverse articulation at midlength;
pleopod 4 peduncle with 4 coupling hooks,
exopod broadly ovate, with distinct artic-
ulation at midlength; pleopod 5 exopod be-
coming almost circular in outline, with dis-
tinct articulation at midlength. Uropodal
peduncle bearing 3 spines at laterodistal an-
gle, mesiodistal angle produced into acute
triangular setose lobe; exopod subequal in
length to, but half basal width of, endopod,
with 5 spines on lateral margin, 3 on inner
233
margin, apically bifid; endopod with 2 or 3
spines on lateral margin, 5 on inner margin,
apically bifid; apices of both rami bearing
elongate simple setae.
Remarks. —Of the Caribbean species of
Cirolana, the present species closely resem-
bles C. albida Richardson, 1901, known only
from the type locality, Sugarloaf Key, Flor-
ida. This resemblance is seen in the overall
size, the shape of the frontal lamina, the
integumental pitting, and the overall simi-
larity of the appendages. The major differ-
ences are seen in the proportions of the uro-
pods and telson. In C. albidoida, the
uropodal rami are more slender, the endo-
pod especially being almost straight-sided;
the posterior telson is also straight-sided,
apically somewhat narrowed, with four
marginal serrations anterior to the apical
spination. Cirolana albida has the outer
margin of the uropodal endopod distinctly
convex, with both endopod and exopod
proportionally broader; the telson is pos-
teriorly more rounded than in the new
species, and lacks the proximal serrations.
The copulatory stylet of pleopod 2 in the
male provides another differences, extend-
ing by almost halfits length beyond the rami
in the new species, and just barely beyond
the rami in C. albida. The number of an-
tennal flagellar articles in C. albida (20-32)
would seem to be higher than in C. albi-
doida (18-20).
Etymology. —The specific epithet, mean-
ing “like albida,” refers to the high degree
of similarity of the two species.
Cirolana minuta Hansen, 1890
Fig. 13, 14
Cirolana minuta Hansen, 1890:347, pl. 3
fig. 5, pl. 4 fig. 1.— Richardson, 1901:512;
1905:83, 92, fig. 74. — Menzies and Glynn,
1968:11.—Schultz, 1969:180, fig. 279.—
Bruce, 1981:961.
?Cirolana minuta: Stebbing, 1900:634
[Loyalty Islands].
Material. -SYNTYPES, Copenhagen
234 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
view; B, Adult in lateral view, C, Antennal, antennular bases
lla 2; G, Maxilla 1; H, Maxilliped; I, Mandible.
Fig. 13. Cirolana minuta: A, Adult 2 in dorsal
and frontal lamina; D, Antennule; E, Antenna; F, Maxi
VOLUME 100, NUMBER 1 235
Fig. 14. Cirolana minuta: A, Pereopod 1; B, Pereopod 2; C, Pereopod 7; D, Uropod; E, Pleopod 1; F, Pleopod
2; G, Pleopod 3; H, Pleopod 4; I, Pleopod 5.
236
Museum, 3 specimens, 5.0, 4.6, 4.1 mm,
probably St. Thomas, U.S. Virgin Islands.
USNM 221618, 4 4, 7.0-8.9 mm, | 2, 4.6
mm, sta GB-7, off Lucaya, Grand Bahama,
180—220 m, coll. D. Camp.
Remarks. —Hansen (1890:348) stated
under the heading ‘Occurrence’ for the de-
scription of C. minuta: 3 specimens, with-
out locality, found in a vial with 1 specimen
of C. parva, which leads to the probable
conclusion that they originate from the West
Indies, and most likely from St. Thomas.
(Translation from J. Just, in litt.). These
three syntypes were compared with the six
USNM specimens from the Bahamas; no
differences other than size could be found,
leading us to suspect that the syntypes are
immature specimens.
Stebbing’s record (1900) from the Loyalty
Islands and his suggestion that C. minuta
is Synonymous with C. /atistylis Dana, 1853,
from the Indo-Pacific, are not dealt with
here. Even though Nordenstam (1946) also
suggested this synonymy, it seems unlikely
that the Indo-Pacific species is the same as
the present Caribbean species.
Cirolana crenulitelson, new species
Figs. 15, 16
Material.—HOLOTYPE, USNM 221619,
ovig. 2, tl 6.8 mm, sta H41-80, PARA-
TYPES, USNM 221620, 2, 7.0 mm, ovig.
2, 6.2 mm, 6, 7.0 mm, sta H41-80, Carrie
Bow Cay, Belize, 36 m, 5 Apr 1980, coll.
G. Hendler.
Description. —Male: Body 2! times long-
er than wide, widest at pereonite 5. Ceph-
alon with well developed eyes; frontal lam-
ina pentagonal, longer than wide. Pereonite
2*%3 length of pereonite 1; pereonites 2-6
subequal in middorsal length, pereonite 7
slightly shorter; coxa of pereonites 2 and 3
posteriorly rounded, of pereonites 4—7 acute,
becoming more elongate posteriorly. Pleo-
nite 1 overlapped laterally by pereonite 7;
epimera of pleonites 1-3 elongate-acute, of
pleonite 4 rounded; pleonite 5 lacking free
lateral margin. Telson basally wider than
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
long, tapering gently to posterior truncate
margin; latter with small but distinct cren-
ulations.
Antennule barely reaching to end of an-
tennal peduncle; articles 1 and 2 fused, to-
gether almost 1'2 times length of article 3;
flagellum of 10 articles, aesthetascs present
on 8 distal articles. Antennal peduncle with
3 short proximal articles, 2 distal articles
more elongate, subequal; flagellum of 15 ar-
ticles. Mandibular palp of 3 articles, article
2 with about 18 spines on outer face; article
3 with row of 22 serrate spines, becoming
distally more elongate; incisor of 3 cusps;
molar bearing 10 teeth. Maxilla 1, inner ra-
mus with 3 stout setae; outer ramus with
about 11 spines, some with few lateral teeth,
on mediodistal margin. Maxilla 2, inner ra-
mus broadly truncate, bearing about 15 se-
tae, 2 proximal setae fringed; inner lobe of
outer ramus with 9 distal setae, outer lobe
with 4 elongate setae. Maxillipedal endite
reaching to distal margin of palp article 1,
bearing single coupling hook and 5 fringed
setae; palp article 3 longest and broadest,
articles 3-5 with setae on inner margins,
outer margins with setae more widely
spaced.
Pereopod 1 robust, dactyl about half
length of anterior margin of propodus; latter
with strong rounded posterodistal spines and
2 smaller acute spines on posterior margin;
carpus short with 2 small spines on poste-
rior margin; merus with 5 stout rounded
spines and 5 small acute spines on posterior
margin; ischium with few scattered small
spines posterodistally. Pereopods 2 and 3
similar, with propodus bearing strong pos-
terodistal spine and 2 smaller spines on pos-
terior margin; carpus more elongate than in
pereopod 1, roughly rectangular, with group
of 3 posterodistal spines; merus with group
of 6 anterodistal spines, posterior margin
bearing about 9 spines of varying lengths;
ischium roughly triangular, with two an-
terodistal spines and 4 posterodistal spines.
Pereopods 4—7 becoming more elongate,
similar; dactyl slightly less than half length
VOLUME 100, NUMBER 1 237
SS
E
Fig. 15. Cirolana crenulitelson: A, Adult in dorsal view; B, Adult in lateral view; C, Antennule; D, Mandible;
E, Maxilla 1; F, Maxilliped; G, Maxilla 2; H, Antenna.
238 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
F
Fig. 16. Cirolana crenulitelson: A, Uropod; B, Pereopod 1; C, Pereopod 2; D, Pereopod 7; E, Pleopod 1; F,
Pleopod 2; G, Pleopod 3; H, Pleopod 4; I, Pleopod 5.
VOLUME 100, NUMBER 1
of propodus; latter elongate-rectangular,
with 3 pairs of small spines on posterior
margin; carpus with entire distal margin
bearing spines of varying length, some elon-
gate and distally fringed, with group of 3
spines on posterior margin; merus with
group of spines at antero- and posterodistal
corners, former more elongate, group of 3
spines on posterior margin; ischium with
group of anterodistal spines, group of 4 short
posterodistal spines and several spines scat-
tered on posterior margin.
Pleopods with all rami bearing marginal
plumose setae. Pleopod 1 peduncle roughly
rectangular, with 5 coupling hooks; endo-
pod parallel-sided in proximal half; exopod
ovate. Pleopod 2, peduncle with 4 coupling
hooks; endopod with basally articulating
copulatory stylet reaching by 4 its length
beyond rami, apically acute. Pleopods 3-5,
exopods with indistinct transverse articu-
lation at about midlength.
Uropodal exopod tapering, elongate-tri-
angular, with 5 sensory spines on outer mar-
gin, 3 spines on medial margin, apically bi-
fid, with short rounded tooth and longer
acute tooth; endopod broad, somewhat ta-
pering, with 3 spines on medial margin, 2
spines on outer margin, both margins with
irregular serrations between spines, apically
bifid, with 2 subequal triangular teeth.
Remarks. —The present species bears a
close superficial similarity to Cirolana ob-
truncata Richardson, 1901, recorded from
Jamaica, and Puerto Rico. Examination of
the holotype of this species, as well as ma-
terial from Puerto Rico and the Gulf of
Mexico reveal several consistent differ-
ences. These include the posterior margin
of the telson, which in C. crenulitelson is
clearly crenulate, but entire in the earlier
species; Richardson (1905:108, fig. 87b) fig-
ures spines on the posterior margin of the
telson; these are not present in C. crenuli-
telson. The uropodal rami are distally
broadly rounded in C. obtruncata, but ta-
pering, marginally serrate, and distally acute-
easy)
bifid in C. crenulitelson. The overall size of
the two species also differs: C. crenulitelson
ovig. 2 6.2-6.8 mm, C. obtruncata 2 up to
11.2 mm. The antennular flagellum has two
fewer articles, the antennal flagellum six
fewer articles in C. crenulitelson.
Etymology. —The specific name refers to
the faint crenulations on the posterior mar-
gin of the telson.
Family Sphaeromatidae
Ancinus belizensis, new species
Figs. 17A, B, 18, 19
Material.—HOLOTYPE, USNM 211401,
2 tl 2.8 mm; sta CBC-K-153, near Colson
Point, north of Dangriga, Belize, sweepings
in seagrass beds, 0.5 m, 20 Nov 1985.
PARATYPES, USNM 211402, 2 6 tl 4.1,
4.1 mm, 2 juvs. tl 2.4, 2.0 mm; sta AC-
CBC-600, between Colson Point and Salt
Creek, north of Dangriga, Belize, sweepings
in segrass beds, 0.1—0.2 m, 7 Jun 1981, coll.
A. Cohen.
Description. —Male: Dorsal integument,
especially of cephalon and pleon strongly
pitted. Cephalon narrower than pereonites,
line of fusion with pereonite 1 marked dor-
sally by impressed line; rostrum anteriorly
rounded, about two-thirds middorsal length
of rest of cephalon. Pereonites all of equal
width; coxae in dorsal view becoming more
elongate posteriorly, except for that of pe-
reonite 7, latter shorter than preceding coxa.
Pleon somewhat inflated, lateral margins
convex, curled ventrally; posterior margin
(apex of triangular pleon) narrowly truncate.
Antennule with basal peduncular article
flexed at right angle, article 3 twice length
of, but narrower than, article 2; flagellum of
12 articles, single aesthetasc on 7 distal ar-
ticles. Antennal peduncular articles increas-
ing in length distally, flagellum of 10 arti-
cles, numerous brush setae on anterior
(medial) surfaces of peduncular and 4 prox-
imal flagellar articles. Mandibular palp of 3
articles, article 2 about twice length of ar-
ticle 3, bearing 8 fringed spines; article 3
240 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig. 17. Ancinus belizensis: A, Pereopod 1, tubercles of proximal cutting edge; B, Pereopod 1, spination of
propodal margin. Casidinidea mosaica: C, Pleotelson in dorsal view; D, Fused setal fringe of uropod.
curved, bearing 5 fringed spines in distal
half; left mandibular incisor of 3 sclerotized
and 2 non-sclerotized cusps; lacinia mobilis
non-sclerotized, with 2 distal cusps; spine
row represented by single stout serrate spine.
Right mandibular incisor as in left, lacinia
mobilis with 2 sclerotized cusps; spine row
with 2 serrate spines. Maxilla 1, outer ramus
bearing 4 stout serrate spines and 8 smaller
and more slender simple spines distally.
VOLUME 100, NUMBER 1
“
241
eT Y hy
"ING
{ Ong
Fig. 18. Ancinus belizensis: A, Adult in dorsal view; B, Antenna; C, Antennule; D, Distal part of right
mandible; E, Left mandible; F, Maxilliped; G, Maxilla 1; H, Maxilla 2.
Maxilla 2, inner ramus with 5 elongate
fringed distal setae; outer ramus with 4 dis-
tal fringed setae, plus single setae on rudi-
mentary outer lobe. Maxillipedal endite dis-
tally bluntly rounded to subtruncate; single
retinaculum on medial margin; basal palp
article short, 2! times wider than long; ar-
ticles 2-5 bearing elongate simple setae on
anterior (medial) lobed margins; terminal
article slender, 3! times longer than broad.
242 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig. 19. Ancinus belizensis 6: A, Pereopod 1; B, Pereopod 2; C, Pereopod 7; D, Pleopod 1; E, Pleopod 2; F,
Pleopod 3.
Pereopod 1, unguis about half length of
rest of dactyl, with elongate accessory spine
at base; proximal half of dactyl bearing row
of rounded scales on posterior margin;
propodus proximally inflated; palm bearing
series of densely-packed long and short dis-
tally widened setae; proximal digitiform
process curved, reaching beyond carpus.
Pereopod 2, dactyl elongate, curved, tip
reaching proximal lobe of propodus, latter
VOLUME 100, NUMBER 1
bearing 3 stubby spines; propodus strongly
curved, palm strongly concave. Pereopod 7,
dactyl % length of propodus; latter parallel-
sided, with about 8 elongate setae on distal
margin; carpus, merus, and ischium each
bearing elongate setae on antero- and pos-
terodistal margins.
Pleopod 2, exopod triangular, less than
halflength of distally tapered endopod; cop-
ulatory stylet elongate-slender, just falling
short of apex of endopod. Pleopods 3-5 typ-
ical of genus. Uropodal ramus reaching to
pleotelsonic apex, tapering, following lateral
margin of pleotelson closely, with faint dis-
tal flexure.
Remarks.—The present species differs
from the five species discussed by Glynn
and Glynn (1974) in several features:
In A. panamensis Glynn & Glynn, 1974,
from Pacific Panama and Colombia, the
dactyl of pereopod 2 of the male is short
and closes on the propodus; in 4. belizensis
the dactyl closes (i.e., the apex reaches) the
carpus. The antennular flagellum has 13 ar-
ticles, the antennal flagellum nine (A. beli-
zensis: 12 and 10 articles, resp.). The prox-
imal digitiform process of pereopod | in the
male is short (relatively elongate in A. be-
lizensis). The two distal articles of the man-
dibular palp bear 9 and 10 setae as against
8 and 5 in 4. belizensis. The coarse and
obvious integumental pitting of the latter
species is distinctive; the pitting in 4. pan-
amensis is finer and less obvious (see Brusca
and Iverson 1985).
In A. brasiliensis Lemos de Castro, 1959,
the pleotelson is as long as its basal width,
and apically narrowly truncate; in A. beli-
zensis the pleotelson is wider than long, more
broadly truncate, and the dactyl of pereopod
2 of the male is relatively longer, reaching
to the midlength of the carpus; the anten-
nular flagellum has 14-16 articles, the an-
tennal flagellum 8-10 articles.
Ancinus granulatus Holmes & Gay, 1909,
of California (with its synonym S. seticor-
nus Frask, 1970), has a densely granulate
integument.
243
Ancinus depressus (Say, 1818) of the east
coast of the U.S.A. and Gulf of Mexico is
a large species (at least three times longer
than A. belizensis), having the uropodal ra-
mus distinctly shorter than the pleotelson.
Etymology. —The specific name refers to
the country of Belize, where the present
species, as well as several others in this pa-
per, was collected.
Cassidinidea mosaica, new species
Figs. 17C, D, 20, 21
Material.—HOLOTYPE, USNM 211403,
6 tl 1.8 mm; sta CBC-K-158, Carrie Bow
Cay, Belize, rubble and coarse sediments at
base of spur and groove buttress, 8-10 m,
24 Nov 1985.
PARATYPES, USNM 211404, 10 spec-
imens (incl. | ovig. 2 tl 1.6 mm); same data
as holotype.—USNM 211405, 15 speci-
mens (incl. 3 ovig. 2 tl 1.5 mm); sta CBC-
AC-610-B, Carrie Bow Cay, Belize, silty
sand from patch reefs, 1.5 m, 14 Jun 1981,
coll. A. Cohen.
Description. — Body oval in outline, about
twice longer than wide. Dorsal integument
finely tuberculate. Lateral margins fringed
with transparent setose flange. Cephalon
embedded in pereonite 1; eyes dorsal, well
pigmented. Pereonites 2—7 subequal in
length and width. Pleonite | free, middor-
sally somewhat inflated, with free lateral
margin. Pleotelson triangular, basally
slightly inflated, posteriorly apex narrowly
rounded. Frontal lamina visible between
antennal bases.
Basal antennular peduncle article flexed,
rest of appendage directed laterally; 2 distal
peduncular articles becoming progressively
shorter; flagellum of 3 articles, distal article
bearing single aesthetasc. Antennal pedun-
cular articles becoming progressively longer
distally; flagellum of 7 setose articles. Man-
dible with strong dentate molar, 4 spines in
spine-row, 3 cusps in incisor; palp of 3 ar-
ticles, article 2 with 2 distal fringed spines,
article 3 with 5 distal fringed spines. Maxilla
1, inner ramus with 4 fringed setae, outer
244
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig. 20. Cassidinidea mosaica: A, Adult in dorsal view; B, Antennule; C, Antenna; D, Mandible; E, Maxilla
1; F, Maxilliped; G, Maxilla 2; H, Pereopod 1; I, Pereopod 2; J, Pereopod 7.
ramus with 3 broad and 3 slender distal
spines. Maxilla 2, inner ramus with 5 distal
fringed spines; inner lobe of outer ramus
with 3 dentate spines and | simple spine,
outer lobe with 3 dentate spines. Maxilli-
pedal endite reaching to article 4 of palp,
distally rounded, with single retinaculum on
median margin, several short fringed spines
distally; palp articles 2-5 with several sim-
ple setae on low inner distal lobe.
Pereopods becoming progressively longer
posteriorly. Pereopod 1 with unguis equal
VOLUME 100, NUMBER 1
245
Fig. 21. Cassidinidea mosaica 6: A, Pleopod 3; B, Pleopod 2; C, Pleopod 1; D, Pleopod 4; E, Pleopod 5; F,
Uropod; G, Penis.
in length to rest of squat dactyl; propodus
with 2 posterodistal spines; carpus trian-
gular, with single posterodistal spine; merus
with 2 anterodistal spines. Pereopod 2, dac-
tyl less squat than that of pereopod 1, prop-
odus and carpus lacking spines. Pereopod
7, carpus rectangular with 2 posterodistal,
and 3 anterodistal stout fringed spines.
246
Penile rami basally fused, distally with 2
slender elongate rami. Pleopod 1 in male,
basis broad, with 2 retinacula; endopod ba-
sally broad, distally tapering, longer than
ovate and narrower exopod. Pleopod 2 in
male, both rami ovate; copulatory stylet at-
tached basally to endopod, basally broad,
tapering distally, extending well beyond
rami. Pleopod 3, basis produced mesially
into lobe bearing 2 retinacula; endopod
elongate-ovate, half length and width of ex-
opod. Pleopod 4, both rami well developed,
pleated. Pleopod 5, exopod *% length of, and
narrower than, exopod. Uropodal basis and
endopod fused, almost reaching pleotelson-
ic apex, distally rounded; exopod short,
ovate.
Remarks. —Menzies & Frankenberg
(1966) regarded Dies and Cassidinidea as
synonymous, but noted the single penis of
the former and the double structure of the
latter. Carvacho (1977) disagreed with
Menzies & Frankenberg, maintaining that
the genital structure required separation of
the two genera. He further characterized Dies
as being estuarine, Cassidinidea as truly ma-
rine. Heard (1982), however, recorded C.
ovalis from the northeastern Gulf of Mex-
ico, from salinities of < 1%0-20%o, 1.e., truly
estuarine. Loyola e Silva (1960) also char-
acterized Dies as having a single penis.
Cassidinidea mosaica differs from C.
ovalis (Say, 1818) (=C. lunifrons (Richard-
son, 1905), see Schultz 1978, Heard 1982)
known from New Jersey to Florida, in hav-
ing a rounded posterior pleotelsonic mar-
gin, in being a smaller species (ovig. 2 1.5—
1.6 mm), and in having a finely tuberculate
dorsal integument.
Cassidinidea tuberculata Richardson,
1912, from Mexico, Brazil, Argentina (see
Pires 1982), has a relatively larger uropodal
exopod, a less ovate body outline, and a
larger body-size (2 tl 5.1 mm).
Etymology. —The specific name refers to
the dorsal integument especially of the pleo-
telson, which resembles the closely-set tes-
serae of a mosaic (see Fig. 17).
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Acknowledgments
Material covered in this paper came from
several sources, in addition to the authors’
collections. We thank Mrs. Anne Cohen,
Dr. Gordon Hendler, and Dr. Jan Kohl-
meyer for such material. We are grateful to
Dr. Jean Just of the Zoological Muesum,
University of Copenhagen, for the loan of
material, and for a translation of part of an
earlier Danish work. Drs. T. E. Bowman
and Richard W. Heard read the manuscript
of this paper; we thank them for their valu-
able comments and criticisms. Ms. Heidi
Wolf and Mrs. Suzanne Braden assisted in
the preparation of the scanning electron mi-
crographs.
This paper is Contribution No. 194 of the
Smithsonian Institution’s Caribbean Coral
Reef Ecosystems program.
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Hansen, H. J. 1890. Cirolanidae et familiae nonnul-
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Heard, R. W. 1982. Guide to common tidal marsh
invertebrates of the northeastern Gulf of Mex-
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Mississippi, 82 pp.
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of Carrie Bow Cay, Belize. Jn K. Ritzler and I.
Macintyre, eds. The Atlantic Barrier Reef Eco-
system at Carrie Bow Cay, Belize, I: Structure
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. 1984. The Atlantic barrier reef ecosystem at
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Kihne, H. von. 1975. Zur geographischen Verbrei-
tung holzzerstorender Crustaceen und Syste-
matik der Untergattung Limnoria s. str. Men-
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553%.
Kussakin, O. G. 1979. Marine and brackish water
isopod crustaceans (Isopoda) of cold and tem-
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order Flabellifera.—Opredeliteli Fauna SSSR
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Universidade do Parana, Zoologia 4:1-182.
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(Crustacea, Isopoda).— Bulletin of Marine Sci-
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villi + 93 pp.
———,,and P. W. Glynn. 1968. The common marine
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, and W. L. Kruczynski. 1983. Isopod Crus-
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Mohr, J. L., and J. A. LeVeque. 1948. Folliculinids
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Nordenstam, A. 1946. Marine Isopoda from Profes-
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1918.—Arkiv for Zoologi 37A(7):1-31.
Ortiz, M. 1983. Guia para la identificacioin de los
isopodos y tanaidaceos (Crustacea: Peracarida),
asociadoss a los pilotes de las aguas Cubanas. —
Revista de Investigaciones Marinas, Universi-
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Pires, A. M.S. 1982. Sphaeromatidae (Isopoda: Fla-
bellifera) da zona entre-mares e fundos rasos dos
estados de Sao Paulo e Rio de Janeiro. — Boletim
do Instituto Oceanografico da Universidade de
Sao Paulo 31(2):43-55.
Richardson, H. 1901. Key to the isopods of the At-
lantic coast of North America with descriptions
of new and little known species. — Proceedings
of the United States National Museum 23:493-
579.
. 1905. A monograph on the isopods of North
America.—Bulletin of the United States Na-
tional Museum 54:i-hii1, 1-727.
Schultz, G. A. 1969. How to know the marine isopod
crustaceans. Wm. C. Brown Co., Dubuque, Iowa.
i-vii + 359 pp.
1978. Four marine isopod crustaceans from
St. Catherines Island with a list of other species
from Georgia.— Georgia Journal of Science 36:
1-12.
Stebbing, T. R. R. 1900. On Crustacea brought by
Dr. Willey from the South Seas.— Willey’s Zoo-
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Department of Invertebrate Zoology, Na-
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mule,
racwiaaye
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mi
A
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CONTENTS
Pseudothelphusa mexicana, a new freshwater crab from the State of Guenere, Mexico ( rac
ura: Pseudothelphusidae) Fernando Alv:
species from the Falkland Islands
Neotropical Monogenea. 10. Omothecium, new genus (Dactylogyndae: Ancyrocephi
two new species from the Piranambu, Pinirampus pirinampu (Spix), (Siluriforme
D. C. Kritsky, V. E. Thatcher, and W.
Dromiopsis kimberlyae, a new Late Cretaceous crab from the Pierre Shale of South Dak ta
Gale A. Bishop
New subspecies of Dryocopus javensis (Aves: Picidae) and Ficedula hyperythra (Aves: Mus-
cicapidae) from the Philippines Robert S. Sue
The subgenera of the crawfish genus Orconectes (Decapoda: Cambaridae)
Ld tps ie oy
C. Hono angustiarum, a new genus and species of Zobrachoidae (Crustacea: Amphipoda) from ive
Magellan Strait, with a revision of Urohaustoriidae Janice Clark and J. L. Barnard ~
Headstanders of the neotropical anostomid genus Abramites (Pisces: Characiformes: Anostom-
idae) Richard P. Vari and Ann M. Williams _
Four new species of clupeoid fishes (Clupeidae and Engraulidae) from Australian waters —
Thosaporn Wongratana _
A new species of Yenurobrycon (Teleostei: Characidae) from the Rio Mamoré basin of Bolivia _
Stanley H. Weitzman |
A new subspecies of Siptornis striaticollis (Aves: Furnariidae) from the eastern slope of the
Andes Gary R. Graves and Mark B. Robbins
Spadella pimukatharos, a new benthic chaetognath from Santa Catalina Island, California |
Angeles Alvarino
Eusarsiella thominx, a new species of myodocopid Ostracoda from the continental shelf of
southern California Louis S. Kornicker ~
Conopora adeta, new species (Hydrozoa: Stylasteridae) from Australia, the first known unat- —
tached stylasterid Stephen D. Cairns
New species of Neomegamphopus from tropical America (Crustacea: Marine Amphipoda)
J. L. Barnard and James Darwin Thomas
A new species of Nannostomus (Teleostei: Lebiasinidae) from near Puerto Ayacucho, Rio —
Orinoco drainage, Venezuela Justa M. Fernandez and Stanley H. Weitzman
Supplementary description of Cypridina americana (Miiller, 1890), a luminescent myodocopid
ostracode from the east Pacific Louis S. Kornicker —
Limnodrilus tortilipenis, new North American species of freshwater Tubificidae (Annelida:
Clitellata: Oligochaeta) Mark J. Wetzel
Records of marine isopod crustaceans associated with the coral Madracis mirabilis from Bar-
bados Brian Kensley and Paul Snelgrove —
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, Mexico Horton H. Hobbs, Jr.
New records of isopod Crustacea from the Caribbean, the Florida Keys, and the Bahamas
Brian Kensley and Marilyn Schotte
THE BIOLOGICAL SOCIETY OF WASHINGTON
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PROC. BIOL. SOC. WASH.
100(2), 1987, pp. 249-256
STEGOPHIURA PONDEROSA (LYMAN), NEW COMBINATION,
AND AMPHIOPHIURA VEMAE AND HOMOPHIURA NEXILA,
NEW SPECIES (ECHINODERMATA: OPHIUROIDEA)
FROM THE R/V VEMA COLLECTIONS
Michael A. Kyte
Abstract.—Amphiophiura ponderosa Matsumoto is placed in the ophiurid
genus Stegophiura and is redescribed. The geographic distribution ofthisspeties—
is extended to the Southern Hemisphere. Amphiophiurg
phiura nexila collected from the R/V Vema off the
South America are described as new species.
Between November 1958 and March
1961, benthic samples were collected from
the R/V Vema of the Lamont-Dougherty
Geological Observatory of Columbia Uni-
versity along the west coast of Central and
South America between the latitudes of 13°N
and 47°S. This sampling was a part ofa long-
term benthic exploration using the Vema.
The collected echinoderm specimens were
deposited in the American Museum of Nat-
ural History.
Among the specimens that were previ-
ously unidentified, two were new species.
Also within these collections were three
specimens of Amphiophiura ponderosa (Ly-
man), which after comparison with the de-
scriptions by Lyman (1882) and Clark
(1911), is assigned to the genus Stegophiura.
Order Ophiurida Muller & Troschel, 1840
Ophiuridae Lyman, 1865
Stegophiura ponderosa (Lyman),
new combination
Fig. 1
Ophioglypha ponderosa Lyman, 1878:93,
pl. 2, figs. 52-54.
Ophiura ponderosa.— Meissner,
925.—H. L. Clark, 1911:77.
Amphiophiura ponderosa. — Matsumoto,
1917:261.
1901:
Material examined.—R/V Vema sam-
la I-V-
a EaNeBbeONUANTS and
ples: V- ie { ae Penas, Chile,
47°02'S, 75°36’ Ww. BD we 24,Marel96t, 3
individeedarerCNCHAET deposited as para-
type in American Museum of Natural His-
tory (AMNH 2758).
Description. — Disc diameter 32 mm, arms
incomplete, length estimated at 100 mm.
Disc about 10 mm high, covered by about
130 irregular, angular, thick plates. Primary
plates conspicuous, separating proximal
ends of radial shields, high, prominent,
forming low projections. Radial shields
large, nearly as wide as long, contiguous dis-
tally along straight inner margin, widely
separated proximally by primary plate. Ra-
dial shields and primary plates locked to-
gether with “dove-tail’’ notch arrangement
at distal apices of primary plates.
Ventral interbrachial spaces covered by 9
to 13 plates. Plates immediately distal to
oral shields irregularly elongate, remaining
plates irregular, angular, similar to aboral
disc plates.
Oral plates narrow, conspicuously swol-
len at proximal end forming a marked ele-
vation on face of each jaw, tapering to points
distally adjacent to adoral plates. Apical pa-
pillae 2, bluntly pointed. Oral papillae 4 to
7 on each side of jaw, varying from small,
low, and truncate to large and elongate, dis-
tinct but continuous with oral tentacle pore
papillae. Oral tentacle pores large, opening
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Stegophiura ponderosa. Dorsal view (top), lateral view (center), and ventral view (bottom).
VOLUME 100, NUMBER 2
partially into mouth slit, protected by 6 large
oral papillae proximally and by 6 smaller
papillae borne by peristomal plate distally.
Oral shields small, arrow-shaped, with
distal rounded semicircular distal lobe be-
tween the genital slits. Adoral shields rela-
tively large, rectangular, with projections at
outer distal ends separating oral shields from
first lateral arm plates. Each adoral shield
bearing 3 to 4 papillae on distal outer cor-
ners similar to opposing genital papillae.
Adoral shields unequal in length, one usu-
ally slightly longer, overriding other.
Genital slits long and narrow, continuing
nearly halfway to aboral side of arms. Each
slit bordered by 3 to 4 plates; small round
plate adjoins oral shield, followed by long,
rectangular plate continuing to aboral arm
base where it fragments into 2 to 4 increas-
ingly smaller angular plates meeting at arm
aboral apex distal to radial shields. Each
plate bearing series of low, rectangular, wid-
er proximally, narrower distally, marginal
papillae continuing aborally to arm apex
forming arm comb.
Arms higher than wide, triangular in cross-
section. Dorsal arm plates much wider than
long, rectangular, becoming pentagonal and
nearly equal, or slightly longer than wide,
near arm tip. Dorsal arm plates appearing
folded over apex of arm, broadly in contact
throughout. First ventral arm plate pentag-
onal, wider than long, adjoining next plate.
Succeeding arm plates trapezoidal to pen-
tagonal to diamond-shaped, wider than long
in distal parts of arm.
Lateral arm plates high, narrow, pointed
at both ends and interlocking with oral and
aboral plates, distal sides convex, proximal
sides concave. First lateral arm plate bear-
ing 16 arm spines, succeeding plates car-
rying approximately 10 spines. Arm spines
dimorphic, 3 on proximal plate, 2 on suc-
ceeding plates, rounded, bead-like, becom-
ing bluntly pointed distally. Second type low,
truncate, first immediately above tentacle
scales which are similar but disappear dis-
tally. Lowest rounded spine separated from
Ar)
tentacle scales by 1 truncate spine, those
above separated by 4 to 5 spines.
Tentacle pores large, protected by as many
as 8 tentacle scales on each side with outer
set overlapping partially, or entirely hiding
inner set. After third or fourth arm joint no
distal tentacle scales, after sixth or seventh
joint only 2 or 3 tentacle scales persisting
with pores to arm tip.
Color (dried from alcohol): whitish tinged
with light brown.
Distribution. — Western Pacific off Japan,
Okhotsk Sea, eastern Pacific off Alaska, cen-
tral California, and Chile; outer continental
shelf to middle slope, 137 to 923 m. Pre-
vious to the collection from the R/V Vema,
Stegophiura was known only from the
Northern Hemisphere (D’yakonov 1954,
Downey 1969). This collection extends the
range of both genus and species to the
Southern Hemisphere from the boreal North
Pacific. The bathymetric distribution of the
species 1S maintained.
Discussion.— This species has not been
described in detail since Lyman (1882), who
had only one specimen with which to work.
Also, Lyman’s description and figures were
detailed and accurate in regards to disc fea-
tures, but were not correct in depicting the
arm spines, as was noted by H. L. Clark
(1911). Because of this situation and the
suggested transference of the species to
another genus, the above detailed descrip-
tion is presented of the most complete and
typical of the three specimens collected.
The genera Amphiophiura and Stego-
phiura were erected by Matsumoto (1915,
1917) while clarifying species relations
within the genus Ophiura. The two genera
were distinguished by Matsumoto on the
basis of only the dimorphism of the arm
spines and the relation between arm height
and width. All other characters of the two
genera were similar according to Matsu-
moto’s discussion. However, the two genera
differ also by the apparent degree of stout-
ness and inflation of the aboral disc scales
with Stegophiura being the stouter.
252
Matsumoto (1917) listed Ophioglypha
ponderosa as an Amphiophiura in his Group
III containing species with the oral inter-
radial disc covered with small plates, with
quadrangular oral arm plates, and uniform
arm spines. According to Lyman’s (1878,
1882) description and figures, this was a cor-
rect generic placement even though Lyman
described the oral arm plates as “broadly
hexgonal.’’” However, the Vema specimens
and two individuals from A/batross station
5023 in the Okhotsk Sea were definitely Ste-
gophiura while also fitting Lyman’s descrip-
tion of O. ponderosa reasonably well. All
specimens had a high and stout disc, higher
than wide arms, and dimorphic arm spines.
It is because of these characters that O. pon-
derosa is transferred here to Stegophiura.
Matsumoto (1917:262) indicated that Group
III, which contained O. ponderosa, ap-
proaches Stegophiura.
Amphiophiura vemae, new species
Fig. 2
Material examined.—R/V Vema sam-
ples: V-17-1, southeast of Punta Aguja, Peru,
7°10’S, 85°50'’W, 4124 m, 26 Feb 1961, 1
individual; V-17-5, off Punta Morquilla,
Chile, 38°15’S, 76°00'W, 3739 m, 15 Mar
1961, 7 individuals, holotype (AMNH
2759), 6 paratypes (AMNH 2760).
Etymology.— The specific name was cho-
sen in honor of the R/V Vema.
Description. — Disc diameter 9 mm; arm
length approximately 12 to 14 mm.
Disc stout, slightly convex, nearly 4 mm
high. Dorsal side of disc covered by about
84 plates. Primaries and radial shield con-
spicuous, slightly swollen. Radial shields
slightly longer than wide, with rounded out-
er edges and straight inner edges for distal
two-thirds of their length; in contact for dis-
tal one-third, separated proximally by small,
wedge-shaped plates and larger, rounded,
slightly swollen plates.
Ventral interbrachial spaces covered by
large oral shields and 14 to 16 small, irreg-
ular marginal and submarginal plates.
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Oral papillae stoutly triangular, well sep-
arated, 3 to 5 on each side of jaw, 2 to 3
longer apical papillae.
Oral tentacle pores large, opening entirely
outside mouth slit, protected by 1 long, low,
flat scale on each side of pore.
Oral shields nearly oval with narrowed
proximal end, lateral sides partially cover-
ing genital slits, separated from first oral
arm plates by adoral shields. Adoral shields
short, rectangular, broadly in contact.
Genital slits long, partially obscured dis-
tally by arm comb papillae. Each slit bor-
dered by 2 plates, most ventral small, in-
conspicuous, carrying 2 or 3 wide, flat,
adjoining truncate papillae. Second plate
large, conspicuous from lateral view, car-
rying marginal series of 7 to 9 wide, flat
papillae with rounded tips.
Arms stout, slightly rounded, bluntly tri-
angular in cross-section proximally. First
dorsal arm plate broadly triangular with
convex distal edges. Subsequent dorsal arm
plates tetragonal with convex distal margins
and flaring sides, becoming pentagonal near
middle of arms, triangular near tips, broadly
in contact proximally but widely separated
near arm tips.
First ventral arm plate hexagonal, wider
than long, distal margin longer than prox-
imal. Succeeding plates tetragonal, as long
as wide with convex ends and concave sides,
becoming more rounded and smaller dis-
tally with most distal nearly circular, broad-
ly in contact until seventh to ninth joint,
after which plates rapidly separate.
Lateral arm plates approximately as wide
as high, rounded, meeting above and below
only in distal part of arm. Each plate car-
rying 2 to 3 equally spaced, small, terete
spines.
Tentacle pores large and protected by 2
long, low, flat scales. Distal scales often in-
conspicuous; tentacle pores and scales per-
sist to arm tip. Color (dried from alcohol):
ivory white.
Distribution. — Southeast Pacific upper
abyssal regions off Peru and Chile, 3739 to
4124 m.
VOLUME 100, NUMBER 2
Fig. 2. Amphiophiura vemae. Ventral view (top) and dorsal view (bottom).
254
Discussion. — While these specimens def-
initely belong to the genus Amphiophiura,
they are distinct from all other species pres-
ently in the genus. Distinctive features in-
clude the relation of the radial shields, the
shape and number of the oral and genital
papillae, and the number and appearance of
the tentacle scales. Litvinova (1971) listed
two other Amphiophiura species in the gen-
eral region from which A. vemae was col-
lected. These two, A. undata (Lyman) and
A. convexa (Lyman), as described by Lyman
(1878, 1882) differ from A. vemae in having
more tentacle scales on each pore, and the
arm comb papillae are pointed rather than
flat and truncate. Also, the oral papillae are
rectangular and truncate in A. undata rather
than triangular and pointed as in A. vemae.
The oral shield of A. undata is relatively
much smaller than in A. vemae.
Litvinova (1971) also described a new
subspecies from the Northwest Pacific, 4m-
phiophiura bullata (Thompson) pacifica
Litvinova, that was collected within the
same abyssal depth range as A. vemae.
However, in addition to the substantial geo-
graphical separation, the species differ in
that A. bullata pacifica has six to eight ten-
tacle scales where A. vemae has only one to
two. Also, specimens of A. bullata pacifica
collected from the same depths as 4. vemae
possessed relatively more dorsal disc scales
than A. vemae between the primary plates.
The relatively large number of specimens
and the variety of sizes present in the two
samples from abyssal depths suggest that
this zone is the normal habitat for the
species.
Homophiura nexila, new species
Fig. 3
Material examined.—R/V Vema sam-
ples: V-15-37, Gulf of Panama, 7°25’N,
79°23'W, 1749 m, 14 Nov 1958, 3 individ-
uals, holotype (AMNH 2761), 2 paratypes
(AMNH 2762); V-15-42, off Costa Rica,
7T°55'N, 86°00'W, 3087 m, 19 Nov 1958, 1
individual; V-15-58, off Costa Rica,
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
12°11'N, 89°34'W, 5690 m, 27 Nov 1958,
2 individuals.
Etymology. — From nexila (Latin), mean-
ing tied together, referring to the tapering,
jointed appearance of the arms.
Description. — Disc diameter 7 mm, arm
length about 12 mm.
Disc stout, flat; dorsal side covered by
approximately 90 plates. Primary plates
conspicuous, convex. Radial shields con-
spicuous, flat, twice as long as wide, nearly
oval, touching at a point one quarter of
length from distal ends, diverging from this
point. Radial shields separated distally by
small, triangular plate in contact with first
dorsal arm plates; proximally separated by
2 plates, more distal longer than wide and
acutely triangular, second nearly tetragonal.
Each radial shield bearing 2 minute, terete,
Opaque spines on most lateral interradial
margins.
Oral plates of jaw nearly trapezoidal with
extended proximal corners, each plate bear-
ing 4 low, fused truncate oral papillae. Sin-
gle, bluntly pointed apical papilla.
Oral shields small, with proximal angle,
rounded distally, well separated from first
ventral arm plates by adoral shields. Adoral
shields large, narrower at proximal end,
broadly in contact. Each plate bearing 1
large, opercular papilla protecting second
oral tentacle pore.
Genital slits short, inconspicuous, guard-
ed by | to 3 small, bead-like papillae. Each
slit bordered by 2 plates, bar-shaped with
rounded ends, distal plate approximately
twice as long as proximal. Proximal plates
bearing genital papillae, distal plates bare.
No arm comb.
Arms rounded in cross-section, rapidly
tapering, slender. First dorsal arm plate
broad, wider than long, semicircular with
straight proximal border adjoining disc plate
separating radial shields distally. Dorsal arm
plates after first small, inconspicuous, te-
tragonal, wider proximally, longer than wide,
becoming equilateral distally, widely sepa-
rated throughout.
First ventral arm plate rounded, approx-
VOLUME 100, NUMBER 2
Homophiura nexila. Ventral view (top) and dorsal view (bottom).
256
imately hexagonal, bearing 2 low, flat pa-
pillae pointed at inner ends and projecting
into mouth slit giving plate notched ap-
pearance. Succeeding ventral arm plates tri-
angular with concave sides, rapidly dimin-
ishing after fifth joint, absent after ninth,
widely separated throughout.
Lateral arm plates wider than high,
broadly in contact after first joint; each plate
carrying 3 to 4 small, evenly spaced terete,
translucent spines.
Oral tentacle pores small, inconspicuous,
opening partly into mouth space, obscured
by adoral papillae. Arm tentacle pores larg-
er, protected by 2 low rounded scales on
first pores only. After second joint only larg-
er outer scales remain. Tentacle pores pres-
ent only to fourth joint.
Color (dried from alcohol): ivory white.
Distribution.— Lower continental slope
and abyssal regions of the Gulf of Panama
and the eastern Pacific off Central America,
1749 to 5690 m.
Discussion. —Hubert Lyman Clark (1915)
established this genus with Ophioglypha in-
ornata Lyman as the type and included 19
species. Paterson (1985) revised Homalo-
phiura and confirmed that the generic name
is invalid. Paterson proceeded to place the
species formerly assigned to Homalophiura
into four groups; one of which was the new
genus Homophiura. The above described
Specimen corresponds well with Clark’s ge-
neric description and with generic charac-
ters in Lyman’s (1878, 1882) description of
O. inornata. The distinguishing specific
characters including the restriction of the
tentacle pores to the proximal four arm
joints, the long genital slits, and the poorly
developed arm combs place this species in
Paterson’s Group D or the new genus Ho-
mophiura. The small spines on some disc
scales are unusual and differentiate H. nex-
ila from other Homophiura species.
Acknowledgments
I am grateful to John H. Dearborn of the
University of Maine who directed this work
as a portion of my Master of Science thesis.
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Ernst Kirsteuer of the American Museum
of Natural History generously provided the
material for study. This research was sup-
ported in part by National Science Foun-
dation grant GV-24157 to John H. Dear-
born.
Literature Cited
Clark, H. L. 1911. North Pacific ophiurans in the
collection of the United States National Mu-
seum.— Bulletin of the U.S. National Museum
75:1-295.
1915. Catalog of recent ophiurans.—Mem-
oirs of the Museum of Comparative Zoology at
Harvard College 25:162-340.
Downey, M. 1969. Catalog of recent ophiuroid type
specimens in major collections in the United
States. — Bulletin of the U.S. National Museum
293:1-239.
D’yakonov, A. M. 1954. Ophiuroids of the U.S.S.R.
seas.— Keys to the fauna of the U.S.S.R., Zoo-
logical Institute of the Academy of Sciences of
the U.S.S.R. 55:1-123. [Translated from Rus-
sian by the Israel Program for Scientific Trans-
lations, Jerusalem, 1967.]
Litvinova, N. M. 1971. Ophiuroids of the genus 4m-
phiophiura in the Pacific and Indian Oceans. —
Trudy Instituta Okeanologii Akademiya NAUK
SSSR 92:298-316.
Lyman, T. 1878. Ophiuridae and Astrophytidae of
the Challenger Expedition, I.—Bulletin of the
Museum of Comparative Zoology at Harvard
College 5(7):65-168.
1882. Report on the Ophiuroidea dredged
by H.M.S. Challenger during the years 1873-
76.— Challenger Reports, Zoology 5(14):1-386.
Matsumoto, H. 1915. A new classification of the
Ophiuroidea: With descriptions of new genera
and species.— Proceedings of the Academy of
Natural Sciences, Philadelphia 67:43-92.
1917. A monograph of Japanese Ophiuro-
idea, arranged according to a new classifica-
tion.— Journal of the College of Science, Tokyo
38(2):1—408.
Meissner, M. 1901. Systematik (der Schlangen-
sterne).—Bronn’s Thierreichs 2(3):902—-966.
Paterson, G. L. J. 1985: The deep-sea Ophiuroidea
of the North Atlantic Ocean.— Bulletin of the
British Museum of Natural History (Zoology)
49(1):1-162.
Department of Zoology, University of
Maine, Orono, Maine 04469. Present ad-
dress: 11025 44th St., S.E., Snohomish,
Washington 98290.
PROC. BIOL. SOC. WASH.
100(2), 1987, pp. 257-261
TRACHELYOPTERICHTHYS ANDUZEIT, A NEW SPECIES
OF AUCHENIPTERID CATFISH FROM THE
UPPER RIO ORINOCO OF VENEZUELA
WITH NOTES ON 7. TAENIATUS (KNER)
Carl J. Ferraris, Jr. and Justa Fernandez
Abstract.— A new catfish species of the Neotropical siluriform family Au-
chenipteridae is given the name Trachelyopterichthys anduzei. This species can
be distinguished from all other auchenipterids by the irregularly distributed
brown spots on the dorsal and lateral surfaces of the body and by a unique
combination of vertebral and fin ray meristics. Sexual dimorphism of the anal
fin and dorsal fin spine is documented in Trachelyopterichthys taeniatus.
The auchenipterid catfish genus TJ7va-
chelyopterichthys was created by Bleeker
(1862) to accommodate Kner’s (1858)
Trachelyopterus taeniatus, a species then
known only from its holotype. Trachelyop-
terichthys taeniatus remains a poorly known
fish, being represented by only a handful of
specimens in museums. Little has been
written about this species due to the dearth
of material available for anatomical studies.
Recent collecting in the Rio Orinoco sys-
tem of Venezuela has uncovered another
species of this genus, again from a unique
specimen. Studies associated with the de-
scription of this new species and the recent
importation of specimens of 7. taeniatus
into the U.S. aquarium fish trade led to the
discovery of previously unknown sexually
dimorphic characters in Trachelyopter-
ichthys.
Materials and methods. — Vertebral counts
include all rib-bearing centra but do not in-
clude any of the anterior, complex-centrum
elements without ribs.
The following institutional abbreviations
are used below: AMNH, American Mu-
seum of Natural History; NMW, Naturhis-
torisches Museum, Vienna; MCZ, Museum
of Comparative Zoology, Harvard Univer-
sity; MZUSP, Museu de Zoologia da Uni-
versidade de Sao Paulo, Brazil; MBUCV,
Museo de Biologia, Instituto de Zoologia
Tropical, Universidad Central de Venezue-
la, Caracas.
Trachelyopterichthys anduzei, new species
Fig. |
Holotype. —MBUCV V-14627, 139.5 mm
standard length, male; Venezuela: Territo-
rio Federal Amazonas; Rio Orinoco, La-
guna de Carida, at the mouth of Cano Cari-
da, Justa Fernandez and Edgar Armas, 28
Apr 1981.
Diagnosis.—A species of Trachelyopter-
ichthys most readily distinguishable from
the only other species in the genus, 7. taeni-
atus, by a pigmentation pattern consisting
of an irregular series of dark spots scattered
over the body and caudal fin. Additional
characters which distinguish these two
species are noted under “‘Remarks.”’
Description. — Meristic values of the ho-
lotype are presented in Table 1. Head de-
pressed, broad; depth at pectoral-fin base
half of its width; fontanelle ovoid, com-
pletely contained within frontals. Nostrils
widely separated, anterior nostril tubular,
posterior with an opercular flap on anterior
margin, above eye. Eye obscure, completely
covered with skin, less than snout length.
Barbels in three pairs, maxillaries extending
to dorsal-fin origin, inner mentals short,
reaching only to more posteriorly placed
258
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig. 1.
outer mentals, outer mentals reach poste-
riorly to pectoral-fin base. Teeth in both
jaws conical, in bands of seven to eight ir-
regular rows. No teeth on vomer or pala-
tines. Lower jaw protruding slightly.
Dorsal fin with narrow base, base one-
third of head length; fin spine pungent, with
single row of feeble denticles along anterior
midline; posterior and lateral margins
smooth, except for fine grooves running par-
allel to long axis. First branched dorsal ray
extending beyond spine, fin margin round-
ed; tip of appressed spine just reaches to
vertical line through pelvic-fin origin. Adi-
pose dorsal fin absent.
Pectoral fin with pungent spine, spine
length equals distance from snout tip to pec-
toral-spine origin, spine with approximately
equal numbers of serrae on both margins.
Anterior serrae antrorse, progressively larg-
er distally; posterior serrae retrorse, all of
approximately equal length. Dorsal and
ventral surfaces of spines with fine parallel
grooves, without granulations, first branched
ray slightly longer than spine, rays becoming
progressively shorter posteriorly.
Pelvic fins broadly rounded, branched rays
3 to 5 longest; appressed fin rays reaching
past anal-fin origin.
Anal fin with branched rays of approxi-
mately equal length, posterior three shorter.
Posterior margin rounded; appressed fin rays
not reaching to caudal fin base.
Trachelyopterichthys anduzei, new species, holotype MBUCV V-14627.
Caudal fin emarginate, middle rays only
a little shorter than lateral principal rays;
lobes broadly rounded, dorsal lobe slightly
longer than ventral.
Cleithral spine long, acutely pointed;
reaching to beneath dorsal-fin origin; di-
rected posterodorsally; lateral surface cov-
ered with rows of fine rounded nodules of
uniform size.
Lateral line canal undulating posteriorly
from beneath dorsal fin, becoming less wavy
posteriorly; cephalic lateral line canal ossi-
fications of the preopercular and infraor-
bital series wholly contained beneath skin
and lack granulations.
Pigmentation in ethanol.—Head brown
dorsally, white on underside, except for
brown lower jaw margin. Body dark brown
dorsally, progressively lighter ventrally, ab-
domen white. Fins brown except for dorsal
and posterior margins of dorsal fin, poste-
rior half of pectoral fins, unbranched pelvic
ray and anterior and distal margins of anal
fin white; brown areas of caudal fin, ventral
and anal fin bases, and dorsal and lateral
surfaces of body, with scattered dark brown
spots 3 mm or less in diameter; spots largest
and most concentrated on dorsal half of lat-
eral body surface.
Etymology. — The patronymic species ep-
ithet is for Dr. Pablo Anduze, former gov-
ernor of the Territorio Federal Amazonas,
Venezuela, for his continued interest in the
VOLUME 100, NUMBER 2
natural history of the Amazonian region of
Venezuela.
Remarks. — Trachelyopterichthys anduzei
differs from its congener in quite a number
of characters. The pigment pattern of T.
taeniatus consists primarily of a series of
alternating black and white stripes running
the length of the body. All fin ray counts
except the caudal differ between the two
species (Table 1). The dorsal fin origin is
proportionally further posteriorly in 7. an-
duzei than in T. taeniatus (29 vs. 23% of
standard length (SL), respectively).
The dorsal spine of 7. anduzei has but a
single anterior row of feeble serrae, whereas
in 7. taeniatus several rows of stout, blunt
tubercles cover the anterior and lateral sur-
faces. The pectoral spine of 7. taeniatus is
also more heavily ornamented, with an ir-
regular patch of small rounded serrae proxi-
mally on its ventral surface, and an acces-
sory row of pointed serrae both dorsal and
ventral to the anterior edge of the spine. In
larger individuals, serrae are more promi-
nent and more widely distributed on the
dorsal and ventral surfaces.
The cleithral process of Trachelyopter-
ichthys anduzei has shorter, more uniform-
ly-sized tubercles and is more acutely point-
ed than in 7. taeniatus. The enlarged series
of tubercles along the dorsal surface of the
cleithral process of 7. taeniatus is unlike
that of any other auchenipterid species.
Relationships.—The genus Trachelyop-
terichthys belongs to a subgroup of the fam-
ily Auchenipteridae defined by several de-
rived characters, among which are an
expanded lateral ethmoid and an accessory
basibranchial cartilage (Ferraris, pers. obs.).
Within this group, 7. anduzei shares three
additional derived characters with 7. tae-
niatus: an elongate anal fin with greater than
38 rays, a high vertebral count, and the ab-
sence of an adipose fin. The first two char-
acters are unique within the aforementioned
subgroup of the Auchenipteridae. The ab-
sence of an adipose fin occurs in a number
of auchenipterid species, including all known
Ss)
Table 1.—Meristics of Trachelyopterichthys anduzei
holotype and 7. taeniatus. Number of specimens ex-
amined is listed in parentheses.
T. anduzei
holotype T. taeniatus
Dorsal-fin rays II,5 11,4 (6)
Pectoral-fin rays 1,8 1,8 (6)
Ventral-fin rays 1,10 1,12-14 (6)
Anal-fin rays 111,36 111,49-55 (5)
Caudal-fin rays 1,7,8,1 1,7,8,1 (5)
Vertebrae
(rib bearing/total) 11/46 7-9/47—50 (5)
species of Trachelyichthys, Epapterus, and
Trachelyopterus. Indeed, this character was
the basis for the inclusion of 7. taeniatus in
Trachelyopterus by Kner (1858). The hy-
pothesis that an adipose dorsal fin is prim-
itively present in catfishes and lost several
times within this group has been supported
recently by Vari and Ortega (1986). Pending
the outcome of a broader study of the re-
lationships among auchenipterid catfishes
(Ferraris, in prep.), it is assumed that the
absence of an adipose fin in 7rachelyopter-
ichthys is not a shared derived character of
a larger group but is, instead, a synapo-
morphy for these two species.
Sexual dimorphism in Trachelyopter-
ichthys taeniatus.—Britski (1972) noted
sexual dimorphism in the anal fin and the
placement of the urogenital pore in Tva-
chelyopterichthys. Anterior anal-fin rays of
females are of the same length and thickness
as those following. In males, the last un-
branched ray and the first two branched rays
are both longer and slightly thicker than
those immediately posterior. There is, how-
ever, no evidence of modification of the rays
or their supporting elements as was found
in Epapterus blomhi (Vari et al. 1984), En-
tomacorus gameroi (Mago-Leccia 1984), and
a number of other auchenipterid species. In
most adult male auchenipterid species, in-
cluding 7. taeniatus, the urogenital pore is
located at the distal tip of the anterior mar-
gin of the anal fin in contrast to the enlarged
pore anterior to the anal fin base in females.
260
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig. 2. Geographic distribution of Trachelyopterichthys anduzei (®) and T. taeniatus: holotype (@), examined
specimens (¥), and literature records (Britski 1972) (A).
In Trachelyopterichthys taeniatus there is,
in addition, a clear difference in the length
and serration pattern of the dorsal spine.
The spine in females and immature males
is between 35% and 43% of the predorsal
length and it lacks serrations on the poste-
rior margin. Substantially longer (60-69%)
spines with a single series of strong, re-
curved serrae on the distal half of the pos-
terior margin were observed in two males.
Distribution. — Trachelyopterichthys an-
duzei was collected in the Rio Orinoco, and
is therefore, the first record of this genus
outside of the Rio Amazonas basin. Tra-
chelyopterichthys taeniatus was described
from a specimen from the Rio Guaporé
(Kner 1858). Additional material collected
from the Thayer expedition (Eigenmann and
Eigenmann 1890), and specimens housed at
MZUSP (Britski 1972) all originate from
the Rio Amazonas and Rio SolimGes (Fig.
2).
Relevant material examined. — Trachely-
opterichthys taeniatus: NMW 43346, ho-
lotype, Rio Guaporé; MCZ 36189, 1 male,
Lake Hyanuary; MCZ 8132, 1 female, Teffe;
MCZ 7731, 1 female, Teffe; AMNH uncat.,
3 aquarium specimens.
Acknowledgments
Research associated with this study was
supported in part by funding from the IESP
Neotropical Lowland Research Program of
the Smithsonian Institution. Dr. Karel Liem
and Mr. Karsten Hartel (MCZ), Dr. Barbara
Herzig and Mr. Harald Ahnelt (NMW) and
Drs. Antonio Machado-Allison and Fran-
cisco Mago-Leccia (MBUCV) allowed us to
examine specimens in their care. Lee Finley
and Ginny Eckstein donated the aquarium
specimens to the American Museum of
Natural History. The assistance of these
people and organizations is greatly appre-
ciated.
VOLUME 100, NUMBER 2
Literature Cited
Bleeker, P. 1862. Notice sur les genres Trachelyop-
terichthys, Hemicetopsis et Pseudocetopsis.—
Comptes-rendus de l‘Academie Royale des Sci-
ences, Section Sciences exactes 14:400-—403.
Britski, H. A. 1972. Sistematica e evolugao dos Au-
chenipteridae e Ageneiosidae (Teleostei, Silu-
riformes). Unpublished doctoral dissertation,
Universidade de Sao Paulo, Brazil 1-146, 60
fig., 6 maps.
Eigenmann, C. H., and R. S. Eigenmann. 1890. A
revision of the South American Nematognathi
or catfishes.— Occasional Papers of the Califor-
nia Academy of Sciences 1:1-508.
Kner, R. 1858. Ichthyologische Beitrage. II. Abthei-
lung.—Sitzungberichte der Kaiserlichen Aka-
demie der Wissenschaften, Mathematisch-Na-
turwissenschaftiche Klasse, Wien 26:373-448,
9 pl.
Mago-Leccia, F. 1984 (1983). Entomocorus gameroi
261
una nueva especie de bagre auquenipterido (Te-
leostei, Siluriformes) de Venezuela, incluyendo
la descripcion de su dimorfismo sexual secun-
dario.—Acta Biologica Venezuelica 11(4):215-—
236)
Vari, R. P., S. L. Jewett, D. C. Taphorn, and C. R.
Gilbert. 1984. A new catfish of the genus
Epapterus (Siluriformes: Auchenipteridae) from
the Orinoco River Basin.— Proceedings of the
Biological Society of Washington 97(2):462—472.
, and H. Ortega. 1986. The catfishes of the
Neotropical family Helogenidae (Ostariophysi:
Siluroidei). —Smithsonian Contributions to Zo-
ology 442:1-20.
(CJF) Department of Ichthyology, Amer-
ican Museum of Natural History, New York,
New York 10031; (JF) Estacion Experi-
mental Amazonas, FONA, IAP, Puerto Ay-
acucho, T.F.A., Venezuela.
PROC. BIOL. SOC. WASH.
100(2), 1987, pp. 262-271
SOME CYCLOPOID AND HARPACTICOID COPEPODS
FROM COLOMBIA, INCLUDING DESCRIPTIONS OF
THREE NEW SPECIES
Janet W. Reid
Abstract.—The cyclopoid copepods Paracyclops andinus Kiefer, P. nove-
narius, new species, Eucyclops serrulatus (Fischer), Mesocyclops aspericornis
(Daday), and the harpacticoid copepods Elaphoidella suarezi, new species, and
E. radkei, new species, are recorded from outdoor artificial cultures of Aedes
aegypti in Colombia. Descriptions of P. novenarius, E. suarezi and E. radkei
are furnished.
Copepods collected from artificial con-
tainers in the municipalities of Anapoima
and Agua de Dios (Cundinamarca) and
Buenaventura (Valle), Colombia, in 1983
and 1985 were sent to me for determination
by Dr. Marco F. Suarez of the Servicio Na-
cional de Erradicacion de la Malaria, Bo-
gota. These outdoor artificial containers,
used to rear larvae of Aedes aegypti, in-
cluded asbestos-cement tanks (capacity 250
1), metal drums (200 1) and tires (4 1). Water
was supplied to the containers from larger
rainwater cisterns. Several species of cyclo-
poid and harpacticoid copepods were pres-
ent in the containers. The 1983 finding of
Mesocyclops aspericornis was reported by
Suarez et al. (1984); all records are sum-
marized below and the new species are de-
scribed.
Locality and date Species recorded
Anapoima, Cundinamarca:
8 Nov. 1983 Mesocyclops aspericornis
(Daday), 18 2
Agua de Dios, Cundinamarca:
20 Aug 1985 Eucyclops serrulatus
(Fischer), 15 2, 2
M. aspericornis, | @
24 Sep 1985 E. serrulatus, 2 9
M. aspericornis, 2 2, 1 6
Buenaventura, Valle:
29 Aug 1985 Paracyclops andinus Kie-
fer, 1 copepodite V 4
Elaphoidella suarezi,
new species, 16 2
5 Sep 1985 Paracyclops novenarius,
new species, 12 8,
10 6, 7 copepodites
18 Sep 1985 Paracyclops novenarius,
1 2, 4 copepodites
Elaphoidella radkei,
new species, | 6
Paracyclops novenarius, new species
Figs. 1-20
Material.—1 2°, holotype, National Mu-
seum of Natural History, USNM 231096,
5 Sep 1985. Paratypes: 1 92, dissected on 1
slide, USNM 231097, 5 Sep 1985; 1 6, dis-
sected on 1 slide, USNM 231098, 5 Sep
1985; 10 2, 9 6, 7 copepodites, USNM
231099, 5 Sep 1985; 1 2, 4 copepodites,
USNM 231100, 18 Sep 1985; all from
Buenaventura, Valle, Colombia. All undis-
sected specimens alcohol-preserved.
Description.— Female: Length of holo-
type excluding caudal setae 0.67 mm; range
of lengths of 11 paratypes 0.57-0.88 mm
(median = 0.63 mm). Prosome (Fig. 1) de-
pressed; posterior margins of 2 anterior pro-
VOLUME 100, NUMBER 2
somal somites smooth, posterolateral mar-
gins of 2 posterior prosomal somites with
hairs. Dorsolateral margin of first urosomal
somite with spines, remaining urosomal so-
mites with toothed hyaline membranes on
posterior margins and few rows of fine spi-
nules; posterior margin of anal somite with
spinules. Genital segment (Figs. 2, 3) slight-
ly expanded anteriorly and tapering poste-
riorly, broader than long; shape of seminal
receptacle normal for genus. Caudal rami
(Fig. 2) about 4 x longer than broad, slightly
divergent, separated at anal somite by a dis-
tance slightly less than breadth of ramus.
Length of lateral seta about equal to width
of ramus; length of dorsal seta about *4 length
of ramus. Ratios of lengths of inner to outer
apical setae 1:6.8:3.5:0.7. Medial apical se-
tae set with fine setules proximally, grading
to coarser setules distally.
Antennule (Figs. 4—7) of 9 articles in most
specimens; article 4 with partial suture on
anterodorsal surface. Article 6 with 2 setae
and 1 narrow esthetasc. In one female, ar-
ticles 3 and 4 of the right antennule are fused
posteriorly and divided anteriorly (Fig. 6);
while in the left antennule (Fig. 7) these ar-
ticles are distinct posteriorly, article 3 -ap-
pearing telescoped under article 2 anterior-
ly. Antenna, labrum and mouthparts as in
Figs. 8-12. Swimming legs 1-4 (Figs. 13-
16) each with rami of 3 articles and spine
formula 3,4,4,3. Terminal article of endo-
pod of leg 4, 1.5 x longer than broad; inner
apical spine 2.1 x longer than outer. Basal
lamellae of legs 1—3 each with crescentic row
of fine spinules on each side of anterior sur-
face and long hairs on margins; lamella of
leg 4 with short hairs near margin.
Medial spine of leg 5 (Fig. 3) reaching
midlength of genital segment when de-
pressed. Ratios of lengths of medial to lat-
eral spine and setae of leg 5, 1:1.5:1.2. Leg
6 (Fig. 2) consisting of 1 seta and 1 spinule
inserted somewhat dorsally.
Male: Lengths of 10 specimens 0.54—0.64
mm (median = 0.60 mm). Ornamentation
263
of somites (Figs. 17, 18) similar to female.
Caudal rami (Fig. 19) about 2.7 longer
than broad; setae similar to those of female.
Antennules geniculate (Fig. 20). Lateral seta
of leg 5 (Fig. 18) long, slender; medial spine
of leg 6 not reaching posterior margin of
next somite; middle seta spiniform, short
and stout.
Etymology.— From Latin “consisting of
nine,” to describe the number of articles in
the antennule.
Remarks.—Species of the genus Paracy-
clops, particularly P. fimbriatus are noto-
riously variable in such features as the pro-
portions of the caudal rami and of the
terminal article of the endopod of leg 4
(Gurney 1933, Lindberg 1958). However,
to my knowledge no population with an-
tennules of 9 articles has been found; all
other species in South America possess an-
tennules of 8 articles (Lindberg 1958, Reid
1985). Paracyclops novenarius otherwise
keys to P. fimbriatus chiltoni in the keys of
Lindberg (1958) and Reid (1985), but differs
in that the inner apical seta on the caudal
ramus is relatively longer than that of P. f.
chiltoni; the basal lamella of leg 4 lacks spi-
nules on its anterior surface; and in the male,
the medial spine of leg 6 does not reach the
posterior margin of the succeeding somite.
Elaphoidella suarezi, new species
Figs. 21-33
Material.—1 2, holotype, USNM 231093,
partly dissected on | slide. Paratypes: 15 8,
USNM 231094, alcohol-preserved; all from
Buenaventura, Valle, Colombia, 29 Aug
1985.
Description.— Female: Length of holo-
type 0.37 mm; range of lengths of 10 para-
types 0.35-0.41 mm (median = 0.38 mm).
Prosomal and urosomal somites with lateral
rows of tiny hairs and toothed posterior
margins. Genital segment (Figs. 21, 22) with
posterolateral row of 8 large spinules on each
side and discontinuous posterodorsal and
Wy yw WANVYNY VAAN ANMUEE Nyy ve nver
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WV Vovyue
‘| \\ ‘ v Re .
yvevvy WW VOUY VVUV WW TD Y yy
pyro
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vegteey wvevy OVVV Vo yf
yv Veyy vy uu yaw WAVES WV VY VY Yu \
Ny \\ if
\ fi;
dvvey \
Vivuvy
WN Ct Ue id
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vy VU VV AVY \
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o-7
8=|2 -——— 5 5
Figs. 1-12. Paracyclops novenarius, female: 1, Habitus; 2, Urosome, dorsal; 3, Genital segment, ventral; 4,
Right antennule, ventral; 5, Left antennule articles 1-5, dorsal; 6, Right antennule articles 1—4, dorsal, of a
second female; 7, Left antennule articles 1-5, ventral, of same female; 8, Antenna and labrum; 9, Mandible; 10,
Maxillula; 11, Maxilla; 12, Maxilliped. Scales = 100 um.
VOLUME 100, NUMBER 2 265
—
wai,
MNS _ Corr
igdsss
SSS:
| i
\,
N ; !
\ \/J
\!
| S/ C
nN fs =
SSAA
Yoh 3 YY
Ie ™ =
ANNAN
yV
+ |4—|6, 18-20
13
Figs. 13-16. Paracyclops novenarius, female: 13, Leg 1; 14, Leg 2; 15, Leg 3; 16, Leg 4.
Figs. 17—20. P. novenarius, male: 17, Habitus; 18, Legs 5 and 6; 19, Caudal rami, dorsal; 20, Antennule.
Scales = 100 um.
266 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Wy,
VW
"Ne eecey, ; es
|
COOOL ay Ay wave
W VAY
VyvWVV Ti voy
aa
py/ VyVPryyy
TAVIVAVAVACAUIVAVAUAVAVAVAVATAUALACA UA A M.A 10 AD Wivy "wy
Sos 22
WIN |
more
a Zz
TAMA Thi tern
roll TAAL LAL LALA BD ee L
Wy
Ke
Ro
ea kT Tee
ge a es ee ee EC OmOO
Figs. 21-30. Elaphoidella suarezi, female: 21, Urosome, lateral; 22, Genital segment, ventral; 23, Anal somite
and caudal ramus, ventral; 24, Anal somite and caudal ramus; dorsal; 25, Antennule and rostrum; 26, Antenna
(setation omitted) and exopod; 27, Mandible; 28, Maxilliped; 29, Leg 1; 30, Leg 2. Scales = 100 um.
VOLUME 100, NUMBER 2
posteroventral rows of smaller spinules.
Genital field extending to midlength of seg-
ment. Urosomal somites 2—3 each with row
of large spinules ventrally and laterally and
smaller spinules dorsally along posterior
margins. Anal somite with group of 3 long
spinules on each side and 2 long spinules
extending posteroventrally over each caudal
ramus (Figs. 21, 23). Operculum smooth,
slightly convex (Figs. 21, 24). Caudal rami
about 1.5 Ti cristata _
T. monacantha xX
T. obtusifrons —
T. orientalis xX
*T. pectinata —
T. tricuspida —
Euphausia americana —
E. brevis =
E. gibboides —
E.. hemigibba ~—
E. mutica =
E. pseudogibba —
E. tenera —
Nematobrachion boopis —
Nb. flexipes =
*Nb. sexspinosus _
Nematoscelis atlantica —
Ns. megalops —
Ns. microps _
Ns. tenella —
Stylocheiron abbreviatum —
S. carinatum —
S. elongatum —
S. maximum _
*S. robustum —
S. suhmii —
x |
x |
Xx
x |
x |
x |
xX
|
|
x |
|
aK KK XK |
xKK KK XK |
x1 Kx |
x |
x | XK |
1 x x |
lL xx xX |
KKK KKK |
|
KKK KKK KK KKK KKK MK XM
|
x |
KKK KK KK XM |
KKK KK |
KKK mK XK |
KKK KK XK |
KKK KKK KX |
x |
x |
x
|
specimens (including 13 with less than 10), IRCZM—Indian River Coastal Zone Mu-
and were suspected of being unrepresenta- seum, Harbor Branch Founda-
tive of the original collection. For this rea- tion, Inc., Fort Pierce, Florida.
son, samples with 100 or more total indi- UMML-—Invertebrate Museum, Rosen-
viduals (hereafter referred to as “100+” stiel School of Marine and At-
stations) were treated separately for analy- mospheric Science, University of
ses of species frequency and relative abun- Miami, Florida.
dance. No further attempts were made to USNM-—Division of Crustacea, National
treat these collections quantitatively. Museum of Natural History,
To confirm adult identifications, speci- Smithsonian Institution, Wash-
mens were compared to type or other iden- ington, D.C.
tified material obtained from several mu-
seums. All such material is listed in the The structure of the male copulatory
“Material examined” sections of the species structure, the petasma, is of taxonomic val-
accounts. Cited repositories are as follows: ue in euphauseacean, and was used in final
VOLUME 100, NUMBER 2
verification of species determinations. Fol-
lowing clearing of the isolated appendage,
using the method of Mikkelsen (1981), the
species-specific arrangement of processes on
the petasma could be observed without ob-
struction or further manipulation. The fe-
male copulatory structure, the thelycum, is
believed equal to the petasma in diagnostic
value (Einarsson 1942, Costanzo and Gug-
lielmo 1976a). However, because thelyca are
more difficult to prepare and observe, they
were examined only when somatic charac-
ters of females were inconclusive.
Larvae were encountered but are not con-
sidered in this study (all are deposited in the
IRCZM).
Synonymies are restricted to the original
description, previous names, petasma and
thelycum descriptions, and one or two use-
ful recent descriptions. All body lengths are
total lengths in millimeters, measured along
the dorsal midline from the tip of the ros-
trum (or frontal plate) to the tip of the tel-
son.
Taxonomic Section
The following artificial key is designed for
identifying adult specimens, and relies
heavily on structures most easily recognized
in the often imperfect specimens commonly
found in plankton tows. Previous keys (e.g.,
Boden et al. 1955, Lomakina 1978) depend
on fragile structures, such as the thoracic
appendages, which frequently suffer damage
or loss during collection. Acronyms in
brackets refer to those used in Fig. 2.
1. Eyes with few ommatidia, sur-
rounded by homogeneous organic
matrix
Family BENTHEUPHAUSIIDAE,
ROS 03s, Bentheuphausia amblyops
— Eyes with numerous, densely-
packedvomumatidia ..). 002.005.
.... Family EUPHAUSIIDAE ... 2
2. Eyes round, or nearly so [eyr],
without division into two lobes; no
29g)
GEORGIA
200m
30°N
te}
FLORIDA 28°N
BAHAMA
BANK
Key West | x?
e
24°Nn
82°w 78°w
Fig. 1. Map of eastern Florida, showing stations
utilized in this study. Open circles = previous records;
solid circles = R/V Gerda stations; solid squares = R/V
Gosnold and R/V Johnson stations (both HBF).
80°w
thoracic endopods greatly elongat-
ed ..(Euphausia, Thysanopoda) .. 3
— Eyes oblong and divided into two
lobes [eyb]; second or third pair of
thoracic endopods greatly elongat-
edufene|ii eee or (Nematobrachion,
Nematoscelis, Stylocheiron) ...... 16
3. Rostrum with secondary dorsal
spine [rss] of varying length ..... 4
— Rostrum simple, with no second-
AINASPINes oe Ae 5
4. Dorsal process of first segment of
antennular peduncle expanded as
a nhoodn See Thysanopoda cristata
— Dorsal process spine-like
Thysanopoda tricuspida
278 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
pep
Fig. 2. Diagrammatic euphausiid to accompany artificial key. Segments (1, 2, 3) of antennular (=first antennal)
peduncle and those (1 through 6) of abdomen are numbered for use with text. abf = antennular bifid process
(first antennular segment); abk = abdominal keel (fifth segment); abp = abdominal photophores (that of second
segment enlarged); abt = abdominal tooth (third segment); adp = antennular dorsal prominence (first segment);
ank = antennular keel (tooth, third segment); ans = antennular spine (second segment); cck = crested carapace
keel; ckh = carapace with humped keel; chf = false chela; cht = true chela; cvd = carapace with ventral denticle;
ene = elongated thoracic endopod (third); eyb = bilobed eye (arrow indicates crystalline cones); eyr = round
eye; mcj = mero-carpal joint (fourth thoracic endopod); obr = oblique ridge (process of first antennular segment);
peh = petasma of Euphausia hemigibba (arrow indicates median lobe); pep = petasma of Euphausia pseudogibba
(arrow indicates median lobe); rab = absent rostrum; rsp = spiniform rostrum; rss = rostral secondary spine.
5. Dorsal prominence (“hump’’) [adp]
on first segment of antennular pe-
duncle, directly anterior to eye ..
Ree (Thysanopoda, in part) .....
— Prominence absent ............
Bah ah aaad i hue naan Ns (Euphausia) .. 10
6. Middorsal tooth [abt] on third ab-
domunalysesmenteaeea et ee
ree eee Thysanopoda monacantha
—* “Foothvabsent200 m
T. obtusifrons
Nb. sexspinosus
S. elongatum
S. robustum
STA ES SS
T. orientalis
T. pectinata
Nb. boopis
S. maximum
~ZZEz
first time off eastern Florida in the present
study, all have been previously collected in
the Gulf of Mexico and so do not reflect
dramatic range extensions for circumglobal,
pelagic species such as these.
In an analysis of species abundance and
relative frequencies (Table 2), the six top-
ranking species comprised over 70% of the
entire collection. Species ranking in this cat-
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
egory was not significantly altered when only
“100+ stations’? were analyzed, unlike
ranking by relative frequency which showed
noticeable change. The figures generated by
analysis of “100+ stations” thus probably
represent a more accurate description of the
eastern Florida euphausiacean community
than those from analysis of the entire col-
lection.
Because all collections analyzed here were
made with open nets, few positive state-
ments may be made about the vertical dis-
tribution or migration of the various species.
However, some information may be ob-
tained by examining species lists from shal-
low (<50 m), medium (<200 m), and deep
(> 200 m) hauls. Table 4 shows that 14 (82%)
of the 17 species in the top 50 m at night
were not found in day hauls to the same
depth, suggesting vertical migration on the
part of those species. The list also shows
three shallow-water species (Thysanopoda
monacantha, Euphausia tenera, Stylochei-
ron carinatum) and four deep-water species
(T. orientalis, T. pectinata, Nematobrachion
boopis, S. maximum) which did not change
depth category, suggesting the absence of
marked vertical migration. Most of these
observations are consistent with the current
categorizations of these species as migratory
or nonmigratory (Table 4).
Gopalakrishnan (1975) discussed the oc-
currence of patterns of enlarged abdominal
photophores in the genus Nematoscelis. Six-
ty-nine stations analyzed here contained one
or more Nematoscelis species, with 58 sta-
tions containing males. Of these 58, 35 sta-
tions contained at least one male with en-
larged photophores (=MEP). Fifteen stations
yielded only one species of Nematoscelis; of
these, two stations contained MEP’s. Sev-
enteen stations yielded two species of Ne-
matoscelis, with 10 stations containing one
MEP species, and | station containing both
species with MEP’s. The remaining 37 sta-
tions contained all three species of Nema-
toscelis; 14 stations contained no MEP’s, 11
contained one MEP species, 11 contained
VOLUME 100, NUMBER 2
two MEP species, and one station contained
all three species with MEP’s. These data
indicate that MEP’s are more frequently en-
countered in populations containing more
than one species of Nematoscelis, lending
support to Gopalakrishnan’s (1975) sugges-
tion that enlarged photophores may play a
role in species recognition. The observation
of more fully enlarged photophores in larger
males of N. atlantica and N. microps further
supports this hypothesis.
Species recognition may also apply as ex-
planation for the species-, sex-, and size-
dependent patterns of abdominal photo-
phore reduction shown here for the three
species of Nematobrachion.
Acknowledgments
Thanks are due to Dr. Robert H. Gore
(Department of Natural Resources Man-
agement, Collier County, Florida) for his
continual interest and support and for much-
appreciated critique of the manuscript. Uni-
versity of Miami collections were made
available through the generosity of Dr. Gil-
bert L. Voss, and much additional assis-
tance was provided by R. B. Toll and R.
Lemaitre. Other specimens were loaned by
T. E. Bowman (USNM), T. Wolff (Zoolo-
gisk Museum, University of Copenhagen),
S. Pinkster [Instituut voor Taxonomische
Zoologie (Zodlogisch Museum), Amster-
dam], A. R. Gurney [British Museum (Nat-
ural History), London], and J. Forest (Mu-
seum National d’Histoire Naturelle, Paris).
A. Cohen (Los Angeles, California), K.
Metzger [Harbor Branch Foundation, Inc.
(HBF)], and C. Carter (Library of Congress,
Washington, D.C.) assisted in procurement
of essential references. C. L. Van Dover
(Marine Biological Laboratory, Woods Hole,
Massachusetts) supplied a valuable English
translation of Lomakina’s (1978) Russian
monograph. C. Baker-Lounibos (Fort Pierce,
Florida) offered criticism and heipful com-
ments on the plates. Finally, I thank the
following, all of whom provided encour-
293
agement, advice, and critical reading of var-
10us parts of the manuscript: T. E. Bowman
(USNM), L. B. Holthuis (Rijksmuseum van
Natuurlike Historie, Leiden, The Nether-
lands), R. B. Manning (USNM), P. S. Mik-
kelsen (HBF), J. E. Miller (HBF), L. E. Scot-
to (Florida Department of Natural
Resources, Fort Pierce), H. J. McDonald
(Fort Pierce), C. L. Van Dover, and L. P.
Lounibos (Florida Medical Entomology
Laboratory, Vero Beach).
This is Part II of the series ‘“‘Studies on
Euphausiacean Crustaceans from the Indi-
an River Region of Florida,’’ and Contri-
bution no. 567 of Harbor Branch Oceano-
graphic Institution.
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ania 7:1—43.
. 1885. Report on the Schizopoda collected by
VOLUME 100, NUMBER 2
H.M.S. “‘Challenger” during the years 1873-
1876.—Challenger Reports, Zoology 13(37):1-
228, 38 pls.
Springer, S., and H. R. Bullis, Jr. 1956. Collections
by the Oregon in the Gulf of Mexico. List of
crustaceans, mollusks, and fishes identified from
collections made by the exploratory fishing ves-
sel Oregon in the Gulf of Mexico and adjacent
seas 1950 through 1955.—U.S. Department of
the Interior, Bureau of Commercial Fisheries,
Special Scientific Report, Fisheries No. 196.
Stepien, J.C. 1980. The occurrence of chaetognaths,
pteropods and euphausiids in relation to deep
295
flow reversals in the Straits of Florida.— Deep-
Sea Research 27A:987-1011.
Tattersall, W. M. 1926. Crustacea of the orders Eu-
phausiacea and Mysidacea from the western At-
lantic.— Proceedings of the United States Na-
tional Museum 69(8):1-—31.
Indian River Coastal Zone Museum,
Harbor Branch Oceanographic Institution,
5600 Old Dixie Highway, Fort Pierce, Flor-
ida 33450-9719.
PROC. BIOL. SOC. WASH.
100(2), 1987, pp. 296-301
HETEROMYSINI FROM GRAND BAHAMA ISLAND:
DESCRIPTION OF HETEROM YSIS AGELAS,
NEW SPECIES, FIRST DESCRIPTION OF
MALE H. FLORIDENSTS, AND NOTES ON
H. GUITARTI (CRUSTACEA: MYSIDACEA)
Richard F. Modlin
Abstract. —Heteromysis agelas is described from waters near Freeport, Grand
Bahama Island. The first description of the male of H. floridensis, known
previously from a single female specimen, is presented. Additional ecological
information on H. guitarti is provided.
Species in the tribe Heteromysini, be-
cause of their cryptic mode of existence, are
rarely obtained when usual collecting tech-
niques are employed. The species described
and discussed herein were obtained by
SCUBA divers when they examined or col-
lected substrates (hosts) on the sea bottom
that contained heteromysids. Although
about 22 species in this tribe have been re-
ported from the western Atlantic Ocean,
only Heteromysoides dennisi Bowman
(1985) has previously been reported from
waters off Grand Bahama Island. Two other
heteromysids are known from the Bahama
Islands. Clarke (1955) described Hetero-
mysis actiniae from waters off Bimini and
discussed its commensal relationship with
the sea anemone Bartholomea annulata.
This species has since been reported from
Lyford Cay and Paradise Island (Brattegard
1970). Brattegard (1970) obtained two spec-
imens of H. guitarti from a sponge collected
off Andros Island. Herein is described a new
species of Heteromysis and the male of H.
floridensis, and some additional ecological
information is presented for H. guitarti.
Type specimens of the new species and
the collection that contained H. floridensis
have been deposited in the U.S. Museum
of Natural History (USNM). The remaining
specimens are in the museum of the Marine
Environmental Sciences Consortium at the
Dauphin Island Sea Lab, Dauphin Island,
Alabama (MESC).
Heteromysis agelas, new species
Fig. 1
Material examined. — Male, 4.7 mm, ho-
lotype (USNM 228745); 5.2 mm male, 5.0
mm female, paratypes (USNM 228746); and
3 males (4.0-5.2 mm), 4 females (4.0—5.1
mm), 3 ovigerous females (4.6-5.5 mm),
and 6 juveniles (2.4—3.0 mm) taken from 8
different specimens of the sponge Agelas
dispar collected by J. Uebelacker near the
November 1975 Hydro-Lab habitat loca-
tion off Freeport, Grand Bahama Island, at
depths from 15-17 m.
Description. — Body robust. Carapace with
anterior margin produced into triangular
rostrum, posterior margin deeply emargin-
ate, partly exposing thoracic segment 7 and
all of 8, anterolateral lobes rounded. Eyes
large, oval, distinctly stalked and directed
laterally; cornea large, oval, medial margin
slightly scalloped with strong ocular tooth
on anterosuperior edge.
Antennular peduncle 3-segmented; seg-
ment 1 about as long as segment 3; segment
2 compressed, with small spine on medial
surface; segment 3 with plumose seta on
medial surface, robust blade-like spine with
subterminal flagellum, 2 plumose setae and
VOLUME 100, NUMBER 2 MSI)
fay 8
GsWs Ins log ler 4 Qa2
@, [IK On|
Bae OOS
Fig. 1. Heteromysis agelas: A, Carapace, dorsal; B, Carapace, lateral; C, Antennular peduncle; D, Antennal
peduncle and scale; E, Mandible, right; F, Mandible, left; G, Mandibular palp; H, Thoracic endopod 3; I,
Thoracic endopod 3, carpopropodus and dactylus; J, Pleopod 3; K, Pleopod 4; L, Uropod; M, Telson. A-B =
male, 5.2 mm, USNM 228746; C-I, L-M, = female, 5.1 mm; J—K = male, 5.0 mm.
298 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
1 naked seta distomedially, 2 small plumose
setae mid-dorsally, male lobe ventrally with
few long hair-like setae.
Antennal scale blade-like, about 0.8 times
length of peduncle, medial margin strongly
convex, lateral margin slightly convex, se-
tose all around, apical segment about 0.06
times length of scale. Antennal peduncle
3-segmented, segment | inconspicuous; seg-
ment 2 1.2 times longer than 3, short robust
plumose seta near distolateral edge, 4 naked
setae distomedially; segment 3 with 4 naked
setae distomedially.
Mandibles and palp: Right and left man-
dibles with blade-like incisors, right incisor
with large anterior cusp, left with anterior
and posterior cusp; right lacinia mobilis with
5 cusps, left with 4 cusps; each mandible
with 3 strong setose accessory blades in spine
row; left molar region more strongly devel-
oped than right. Mandibular palp 3-seg-
mented; segment 1 small, inconspicuous;
segment 2 expanded, medial margin with
16 naked setae, lateral margin with 10 na-
ked setae; segment 3 medial margin sin-
uous, proximal half with 2 long naked setae
and 3 robust spined setae, distal half with
about 5 short spines and 1 long plumose
seta, apex with 2 long robust spine setae, 4
long simple setae on lateral surface.
Labrum, maxillule, maxillae, and parag-
naths typical of genus.
Thoracic endopods | and 2 typical of ge-
nus. Thoracic endopod 3, ischium about 1.4
times longer than carpopropodus, medial
margin with 3 long and 4 short simple setae,
small protuberance distomedially, lateral
margin with 1 simple seta proximally and
1 simple seta distolaterally; medial margin
of carpopropodus with 5 robust flagellated
spines arranged as 2 pairs of equal-length
spines distal to single short spine, 3 long
simple setae submarginal to spinal groups,
2 short simple setae proximally, 1 simple
seta distally; dactylus inconspicuous with 2
simple setae distolaterally, terminating in
long robust claw with 3 simple setae at mid-
length. Carpopropodus 3-segmented in tho-
racic endopod 4; 6-segmented in thoracic
endopods 5-8. Thoracic exopod | with 9
segments; exopods 2-8 each with 10 seg-
ments.
Pleopods unsegmented; male pleopods 1,
2, and 5 not differentiated. Pleopod 3 with
8 robust plumose setae along anterior sur-
face, 4 flagellated spines along distal margin
and 1 naked long robust seta distolaterally,
2 short robust plumose setae proximally on
posterior surface, distal margin of pseudo-
branchial lobe with 4 long and 1 short plu-
mose setae. Pleopod 4 with 8 robust plu-
mose setae along anterior surface, 5
flagellated spines on distal margin and 1
simple long robust seta distolaterally, 2 short
robust plumose setae proximally on poste-
rior surface, distal margin of pseudobran-
chial lobe with 4 long and 1 short plumose
setae. Female pleopods rudimentary.
Uropods: Exopod about 1.1 times longer
than endopod, lateral margin straight, me-
dial margin slightly convex, setose all
around. Endopod linguiform with 2 small
spines on medial margin in region of stato-
cyst, setose all around.
Telson 0.8 times length of exopod of uro-
pod, lateral margins slightly concave, com-
pletely spined with 15-16 spines per margin
(apical spines included), marginal spines in
distal half slightly longer than proximal
group, outer apical spine 2.5 times longer
than inner, cleft completely spined with 12
spines, 0.2 times length of telson.
Remarks. —Heteromysis agelas appears
to be intermediate between H. bredini Brat-
tegard, 1970, and H. guitarti Bacescu, 1968;
(Brattegard 1970). It differs from H. bredini
by having the antennal scale shorter than
the antennal peduncle, 5 rather than 10 flag-
ellated spines on the carpopropodus of tho-
racic endopod 3, 2 short rather than 5 long
spines in the region of the statocyst, and a
telson with 15-16 instead of 19-20 spines
on each lateral margin and 12 rather than
31 spines in the telsonal cleft. Similarily, 7.
guitarti has 7 flagellated spines on the me-
dial margin of the carpopropodus of tho-
VOLUME 100, NUMBER 2
racic endopod 3, 4 long spines on the en-
dopod of the uropod in the vicinity of the
statocyst, 9-10 spines on each lateral mar-
gin of the telson and 16-21 spines in the
cleft of the telson. Males of H. guitarti and
H. agelas have pleopods 3 and 4 modified.
Pleopods 3 and 4 of the latter have, respec-
tively, 4 and 5 flagellated spines on the distal
margins rather than 2 and 9 as does the
former. Males of H. bredini have not been
described.
Ecological notes. —Eight of the 19 speci-
mens of Agelas dispar examined contained
H. agelas. The average number of individ-
uals taken from a sponge was two, range
one to five. The three ovigerous females each
carried two eggs. Modlin (1984) obtained
specimens of H. beetoni from A. dispar col-
lected on the Florida Middle Grounds in
the Gulf of Mexico.
Etymology. —Named for the sponge, Age-
las dispar, with which this mysid was found
associated.
Heteromysis floridensis Brattegard, 1969
niga
Heteromysis floridensis was described
from a single female specimen collected in
Biscayne Bay, Florida (Brattegard 1969). No
other specimens of this species have since
been reported. This report describes the male
of the species for the first time and extends
the range of H. floridensis to include the
northern end of the Bahama Islands, spe-
cifically Little Bahama Bank.
Only those morphological features of H.
floridensis that have not been described and
that differ from Brattegard’s (1969) original
description are discussed.
Material examined. —Male (3.7 mm), 25
Oct 1982, voucher (USNM 228747); 2 fe-
males (3.2, 3.5 mm), 2 ovigerous females
(4.5 mm), 11 Nov 1983, voucher (USNM
228748); all collected near entrance to Cem-
etery Cave, Grand Bahama Island, an
oceanic blue hole about 100 m off the south
coast of the island, depth 3—4 m, collected
BY)
by Jill Yager. The actual substrate from
which the specimens were taken is un-
known.
Description. —Antennular peduncle of
male like that of female, but with prominent
male lobe on ventral surface of segment 3
that contains many long hair-like setae.
Mandibles and palp: Right and left man-
dible incisor blade-like with single cusp, right
lacinia mobilis with 2 prominent cusps, left
lacinia mobilis with 2 slightly curved cusps,
both mandibles with 2 robust serrate ac-
cessory blades, molar surface well devel-
oped on right mandible, weakly developed
on left mandible. Mandibular palp segment
2 expanded, medial margin straight, armed
with 12—13 simple setae, lateral margin con-
vex, armed with 4 simple setae; distal seg-
ment medial margin sinuous, armed proxi-
mally to distally with 5 robust spined setae,
about 5 small serrate setae, 2 long spined
setae and terminal claw-like spine, lateral
margin convex armed with 2 long and 2
short submarginal setae distally.
Thoracic endopod 3, merus without
barbed spines on lateral surface described
by Brattegard (1969); carpopropodus with
6 rather than 5 strong flagellate spines on
medial margin, distal 4 arranged in 2 paired
sets, proximal 2 slightly longer and arranged
singly.
Pleopod 4 of male modified, lateral mar-
gin armed with 5 long submarginal plumose
setae proximally, 26 closely-spaced flagel-
late denticles along distal margin, slightly
longer simple seta distomedially, medial
margin with 2 plumose setae proximally;
pseudobranchial lobe with 4 long and | short
plumose setae.
Uropod endopod with 14, rather than 19,
subequal spines along medial margin.
Telson with 8 spines on distal half of each
lateral margin (apical spines included) in-
creasing in length distally, cleft with 10-11
spinule in apical half rather than 9.
Ecological note. —The two 4.5 mm ovig-
erous females carried three and five stage
IV larvae, respectively.
Fig. 2. Heteromysis floridensis: A, Mandible, right; B, Mandible, left; C, Mandibular palp; D, Thoracic
endopod 3; E, Pleopod 4; F, Uropod; G, Telson. A-G = male, 3.2 mm.
VOLUME 100, NUMBER 2
Comments. —The characters of the male
H. floridensis strengthen the phylogenetic
relationship suggested by Brattegard (1969)
between H. bermudensis, H. b. cesari, H.
actiniae, and H. floridensis. Among other
similarities, all four species have about the
same number (6—12) of spines on the lateral
margins of the telson and in the telsonal cleft
(9-20), and all males have pleopod 4 mod-
ified. Assuming that a reduction in spines
suggests evolutionary advancement, H. ber-
mudensis would be the plesiomorph be-
cause it has the most spines on key mor-
phological features. For example, its male
pleopod 4 has 35 flagellated spines on the
distal margin (Bowman 1981) and its telson
has 18-20 spinules in its cleft. Distal mar-
gins of the fourth pleopod of H. b. cesari,
H. actiniae, and H. floridensis have, re-
spectively, 26, 17, and 26 flagellate spines,
while the clefts of their telsons contain 14,
9-13, and 9-11 spines (Brattegard 1969,
Bowman 1981).
Heteromysis guitarti Bacescu, 1968
Material examined.—8 males (3.3—3.7
mm), 6 females (2.9-3.7 mm), 2 ovigerous
(4.0, 4.1 mm), 3 juveniles (2.2—2.4 mm) tak-
en with 12 specimens of the sponge Agelas
dispar collected during November 1975 by
J. Uebelacker near the Hydro-Lab habitat
location off Freeport, Grand Bahama Island
at depths from 15-17 m.
Geographic distribution. — Originally col-
lected in waters off northern Cuba (Bacescu
1968), also reported from waters off Andros
Island, Bahama Islands (Brattegard 1970),
Lesser Antilles (Brattegard 1975), and Gulf
of Mexico (Modlin 1984).
Ecological notes.—Heteromysis guitarti
appears to be a true spongicolous species.
It was found in 12 of the 19 specimens of
Agelas dispar examined. An average of about
two individuals were taken in each sponge,
range one to four. It occurred together with
H. agelas in three sponges. Modlin (1984)
obtained specimens from the sponge /rici-
nia campana. Likewise, Bascescu (1968)
301
found specimens inhabiting J. fasciculata.
Specimens collected by Brattegard (1970,
1975) were taken from unidentified sponges.
The two ovigerous females collected each
carried two stage I larvae.
Acknowledgments
The Uebelacker and Yager collections
were kindly provided for inspection by Mr.
Michael Dardeau (MESC) and Dr. Thomas
E. Bowman (USNM), respectively. This is
Contribution 109 from the Marine Envi-
ronmental Sciences Consortium, Dauphin
Island, Alabama 36528.
Literature Cited
Bacescu, M. 1968. Heteromysini nouvaux des eaux
cubaines: Trois especes nouvelles de Hetero-
mysis et Heteromysoides spongicolan. g.n. sp.—
Revue Roumaine de Biologie, Zoologie 15:11-
16.
Bowman, T. E. 1981. First description of the male
opossum shrimp, Heteromysis bermudensis
bermudensis (Crustacea: Mysidacea).—Pro-
ceeding of the Biological Society of Washington
94(2):458-461.
1985. Heteromysoides dennisi, a new mysi-
dacean crustacean from Cemetery Cave, Grand
Bahama Island.— Proceedings of the Biological
Society of Washington 98(4):945-948.
Brattegard, T. 1969. Marine biological investigations
in the Bahamas. 10. Mysidacea from shallow
water in the Bahamas and southern Florida. Part
1.—Sarsia 39:17-106.
1970. Mysidacea from shallow water in the
Caribbean Sea.—Sarsia 43:111-154.
1975. Shallow-water Mysidacea from the
Lesser Antilles and other Caribbean regions. —
Studies on the Fauna of Curacao and Other Ca-
ribbean Islands 157:102-115.
Clarke, W. D. 1955. A new species of the genus Het-
eromysis (Crustacea, Mysidacea) from the Ba-
hama Islands, commensal with a sea-anemo-
ne.—American Museum Novitates 1716:1-13.
Modlin, R. F. 1984. Mysidacea from the Florida
Middle Ground, northeast Gulf of Mexico, with
descriptions of three new species of Heteromysis
and a key to the Heteromysini of the Western
Atlantic.—Journal of Crustacean Biology 4(2):
278-297.
Department of Biological Sciences, The
University of Alabama in Huntsville,
Huntsville, Alabama 35899.
PROC. BIOL. SOC. WASH.
100(2), 1987, pp. 302-320
CRYPTOCORYNETES HAPTODISCUS, NEW GENUS,
NEW SPECIES, AND SPELEONECTES BENJAMINI,
NEW SPECIES, OF REMIPEDE CRUSTACEANS
FROM ANCHIALINE CAVES IN THE BAHAMAS,
WITH REMARKS ON DISTRIBUTION AND ECOLOGY
Jill Yager
Abstract. —Two additional members of the crustacean class Remipedia, Spe-
leonectes benjamini n. sp., and Cryptocorynetes haptodiscus n. gen., n. sp., are
described from anchialine caves on the Little Bahama Bank in the northern
part of the Bahamas. The Bahamian anchialine caves in which remipedes are
found represent a low oxygen environment.
Since the discovery in Lucayan Cavern,
Grand Bahama Island, of the first represen-
tative of the crustacean class Remipedia,
Speleonectes lucayensis Yager (1981), ad-
ditional members of the class have been
found in the low oxygen waters of anchialine
caves throughout the Bahamian archipela-
go. Outside of the Bahamas, undescribed
remipedes were collected in 1986 from an
anchialine cenote on the Yucatan Penin-
sula, and in addition, Speleonectes ondinae
(Garcia-Valdecasas), the second species of
the genus Speleonectes, was discovered in a
submerged lava tube in the Canary Islands
(see Garcia-Valdecasas 1984, Iliffe et al.
1984, Schram et al. 1986). Remipedes ap-
pear to be common components of low oxy-
gen anchialine cave communities in the
north Atlantic Ocean, especially in the West
Indies region (Table 1).
In the summer of 1984, a joint British-
USA-Bahamian cave diving expedition sur-
veyed many anchialine caves on and near
Sweeting’s Cay at the east end of Grand
Bahama Island. Two of the caves explored,
Sagittarius and Asgard, were inhabited by
Speleonectes benjamini, described below, the
third species of the genus Speleonectes to be
described. Also in 1984 the biological sur-
vey of several previously unexplored caves
on Grand Bahama and Abaco Islands re-
sulted in the collection of an unusual new
genus and species of remipede, Crypto-
corynetes haptodiscus, described below.
Speleonectes benjamini, new species
Figs. 1-6
Material examined. — BAHAMAS: Grand
Bahama Island, Sweeting’s Cay, Asgard
Cave, holotype, adult, 16.8 mm, USNM
228199, 27 Jul 1984, R. Palmer, S. Cun-
liffe. —Sagittarius Cave, | adult, 5 Jul 1984,
R. Palmer; | adult, 25 Jul 1984, D. Wil-
liaams.— Abaco Island, Dan’s Cave, 2 adults,
26 Jul 1985, D. Williams, J. Yager. Non-
type material retained in the collection of
the author.
Diagnosis.—Antenna 1 very long, ex-
tending from one-half to two-thirds length
of body. First maxilla with long, slender dis-
tal fang; segment | with narrow endite bear-
ing 1 moderately long apical spine and at
least 10 accessory spines. Trunk sternites
developed as cuticular plates with postero-
lateral projections. Sternal bars of trunk seg-
ments 1-13 with concave posterior mar-
gins, 14th triangular, remaining bars concave
to triangular until anal segment. Trunk ap-
pendages with many small, serrate, spine-
like comb setae along distal margins of pre-
distal segments.
VOLUME 100, NUMBER 2
303
Table 1.—List of known remipedes from anchialine caves of the West Indies region.
Grand Bahama Island:
Speleonectes lucayensis
Speleonectes benjamini
Cryptocorynetes haptodiscus
Undescribed “‘juvenile-like”’
Abaco Island:
Speleonectes cf. lucayensis
Speleonectes benjamini
Cryptocorynetes haptodiscus
Godzillius cf. robustus
Undescribed godzilliid
Undescribed “‘juvenile-like”’ species
Andros Island:
Speleonectes cf. lucayensis
Description. —Body elongate, slender,
with tiny setae dispersed over surface, with-
out pigment or eyes; maximum length of
specimens examined 16.8 mm. Cephalic
shield small, tapered slightly at anterior end.
Trunk segment numbers varying with age,
maximum number in material examined 27.
First trunk segment reduced and covered by
cephalic shield; pleura from second segment
posteriad projecting laterally. Sternites (Fig.
3C) developed as cuticular plates with tri-
angular posterolateral projections. Trans-
verse cuticular sternal bars posterior to each
plate, with concave posterior margins on
trunk segments 1-13 (Fig. 3C), triangular
on segment 14, concave on several segments
posterior to 14th segment, then triangular
in shape until anal segment.
Frontal filaments long, slender, cylindri-
cal, with short, tapered medial process. An-
tenna | (Fig. 1A) biramous, very long, slen-
der, extending well beyond cephalon, about
two-thirds length of body. Peduncle 2-seg-
mented; proximal segment enlarged, bear-
ing several rows of densely packed, fine es-
thetascs (Fig. 5A, B) draping over second
antenna toward mouth; distal segment bi-
furcate. Dorsal ramus long, with at least 15
slender, elongate segments. Ventral ramus
short, less than 2 length of dorsal ramus,
with 10 to 14 segments; segments near base
Cat Island:
Speleonectes cf. lucayensis
Godzillius cf. robustus
Providenciales Island:
Lasionectes entrichoma
North Caicos Island:
Lasionectes entrichoma
Undescribed species with discoid organs
Godzillius robustus
Yucatan Peninsula:
Undescribed speleonectid
of ventral ramus indistinctly divided by
partial sutures. Segments of both rami with
short, simple setae along ventral margins,
clusters of forked esthetasc-like setae (Fig.
3B) on distoventral margins, and at least 4
terminal setae on apical segments. Antenna
2 (Fig. 1B) biramous, smaller than antenna
1, not extending beyond cephalon. Proto-
pod 2-segmented, with moderately long se-
tae on medial margins. Exopod a single,
large, oval article, extending laterally from
distal segment of protopod, bearing about
54 long, finely plumose setae along entire
margin. Endopod 3-segmented, curving lat-
erally; first segment with moderately long
setae on anterior margin; second segment
with row of moderate to long anterior setae
becoming a double row distally, and several
short setae on posterior margin; third seg-
ment with row of about 30 setae, becoming
a double row along anterior and distal mar-
gins; all setae plumose.
Labrum a prominent, fleshy lobe, with
distinct transverse groove; anterior half tri-
angular and slightly raised at apex; posterior
half with dense array of fine setae along
margin of fossa. Mandibles well devel-
oped, asymmetrical. Right mandible with
3-cusped incisor process and 3-cusped la-
cinia mobilis. Left mandible (Fig. 1C) with
4-cusped incisor process and crescent-
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
304
A |
\ ne ceonieasanaee
B
Speleonectes benjamini, new species. A, Antenna 1; B, Antenna 2; C, Left mandible; D, 14th trunk
appendage with gonopore; E, Anal segment with caudal rami and one terminal trunk appendage.
shaped lacinia mobilis. Molar process of
both broad, bearing long, thick, multitipped
setae. Paragnath (Fig. 4A) a round, flattened
lobe with dense covering of fine, hair-like
setae along margin.
First maxilla (Fig. 2A) 7-segmented, uni-
ramous, prehensile, robust. Segment | (Figs.
2B, 4A) with long, narrow endite terminat-
ing in | long spine and up to 10 shorter,
stout subterminal spines arranged in op-
posing pairs. Segment 2 (Fig. 4A) with en-
dite as broad, plate-like flap lying over par-
agnath and bearing at least 8 short spines
on terminal margin, flanked by parallel row
of many short, simple setae; a row of long
setae on anterolateral margin, and a few
VOLUME 100, NUMBER 2
ven
NN
AWE
305
Fig. 2. Speleonectes benjamini, new species. A, Maxilla 1; B, Maxilla 1, endite of first segment; C, Maxilla
2; D, Maxilliped; E, Claw complex of maxilliped.
short to long setae near base of endite. Seg-
ment 3 with conical medial endite (see Fig.
4A) terminating in 2 robust, conical setae
with fine serrations on distal halves, flanked
posteriorly by cluster of several short to
moderate setae. Segment 4 longer than seg-
ment 3, medial margin rounded, with 1 ro-
bust, serrate proximal seta and 2 rows of
setae; setae of anterior row long, serrate,
setae of posterior row less robust than those
of anterior row, short to long, simple. Prin-
ciple flexure of appendage between seg-
ments 4 and 5. Segment 5 about as long and
wide as segment 4, with cluster of setae on
anterior and posterior distomedial margins.
Segment 6 very short, with cluster of long
306
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig. 3.
Speleonectes benjamini, new species. A, Head appendages (29 x); B, Antenna 1, branching, esthetasc-
like setae (1330 x); C, Sternal plates with cuticular bars (56 x).
simple setae on anterior and posterior dis-
tomedial margins and cluster of moderate
to long setae on anterior and posterior dis-
tolateral margins. Segment 7 a single long,
slender fang (Fig. 3A) about 3 times as long
as segment 6, with terminal pore and tuft
of long, fine, simple setae at base medially.
Second maxilla (Fig. 2C) 7-segmented,
uniramous, prehensile, longer than maxilla
1. Segment 1 with 2 small, weakly devel-
oped lobes posteriorly, bearing a few short
setae each; 3 anteriorly-directed digitiform
endites, progressively larger distally; each
endite with single, curved terminal spine-
like seta, several small subterminal spine-
like setae, and several moderately long setae
anterolaterally. Segment 2 shorter than seg-
ment 1, with small, rounded endite bearing
VOLUME 100, NUMBER 2
307
Fig. 4. Speleonectes benjamini, new species. A, Mouth area (117 x), P—paragnath, El —maxilla 1, endite of
first segment, E2—maxilla 1, endite of second segment, E3—maxilla 1, endite of third segment; B, Maxilla 2,
claw complex of terminal segment (650 x); C, Maxilliped, claw complex of terminal segment (527 x).
1 short, stout, apical spine-like seta and 2
rows of setae, setae of anterior row long,
serrate, setae of posterior row short to mod-
erately long. Segment 3 long, with proxi-
momedial rounded bulge bearing 2 rows of
setae; anterior row with many moderate to
long serrate setae, posterior with fewer short
to moderate setae. Principle flexure of ap-
pendage between segments 3 and 4. Seg-
ment 4 about as long as segment 3, with row
of many moderate to long setae along dis-
tomedial margin, cluster of moderately long
setae on anterior and posterior distomedial
margins, and several small setae on disto-
lateral margin. Segment 5 shorter than seg-
ment 4, with row of many moderate to long
308 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Figs:
Speleonectes benjamini, new species. A, Antenna 1, esthetascs (85 x); B, Antenna 1, closeup of base
of esthetascs (2200 x); C, Gonopore (136 x); D, Porous plate adjacent to gonopore (1710).
setae along entire medial margin; cluster of
several small to moderate setae on anterior
and posterior distomedial margins, several
small setae on distolateral margin. Segment
6 about as long as segment 5, with row of
moderate to long simple setae along medial
margin; cluster of moderate to long setae on
anterior and posterior distomedial margins,
and several moderate to long setae on an-
terior and posterior distolateral margins.
Segment 7 (Fig. 4B) with claw complex con-
sisting of 1 long, stout, anterior claw, flanked
subterminally by smaller, stout claw, and
posterior horseshoe-shaped arrangement of
smaller spines; pores on surface of claw
complex. Thumb-like pad bearing many
long, simple, esthetasc-like setae opposing
claw complex.
Maxilliped (Fig. 2D) similar in form but
longer than maxilla 2, having 1 more seg-
ment beyond point of flexure; 8-segmented,
uniramous, prehensile. Segment 1 indis-
tinctly subdivided with several weakly de-
veloped lobes bearing a few short medial
setae. Segment 2 with medial lobe bearing
anterior row of long, serrate setae and pos-
terior row of short to moderate simple setae.
Segment 3 moderately rounded, with 2 rows
of setae, anterior setae longer and more nu-
merous than those of posterior row. Prin-
ciple flexure between segments 3 and 4. Seg-
ments 4, 5, and 6 with medial row of short
to moderate setae,-clusters of several long
anterior and posterior distomedial setae, and
several setae on distolateral margins. Seg-
ment 8 (Figs. 2E, 4C) with claw complex
and opposable pad similar to maxilla 2.
Trunk appendages (Fig. 1D) biramous,
paddle-like swimming appendages becom-
ing smaller and less setose near anal seg-
VOLUME 100, NUMBER 2
f “ae, 3 y <
—_— 2 » SE
CSS TE, ~ at
al & G@= Sa Gn DD aa ES ED, 4 BP,
pai i as — oo = SD Gil <<
a en OS 6 Gs oS pa aa
Se eS SST ~s
= IAn> Poa sofresh or Ve
a ey Kee brackish 9k
eens a, pe
—— ae 2eces oF ee a weet, A
> veeeet KS
Saal DENG YA
2 ED ay =D => =a eo GD a
Hr Sr p+ Ft INTERFACE C—
Oe BEY a ee ey
ane Ch) Sa Ge. Oe ae i a
S S52 Ga GS es, ee SS a : T,
See (2 ay aap Gee os
Tana Wh VV
habitat for
remipedes and other fauna Sed mater
Fig. 12. Diagram of typical anchialine cave of the West Indies.
tain waters with dissolved oxygen values
higher than | ppm (T. M. Iliffe, pers. comm.).
The low oxygen anchialine ecosystem can
be considered a refugium because it repre-
sents a “‘stable”’ habitat. Although the caves
accessible to divers today are geologically
young, the Bahama Banks contain numer-
ous caves at greater depths, providing a
crevicular habitat that has been available
for colonization throughout the 150 million
year history of the Bahamas Platform. The
low oxygen anchialine fauna may be de-
scendants of taxa that have inhabited this
environment for millions of years relatively
unaffected by surface climatic changes, and
the low oxygen ecosystem may have served
as a refugium for species that were able to
survive Oceanic anoxic events (Degens and
Stoffers 1976, Arthur and Schlanger 1979).
Acknowledgments
I am grateful for the support of the Cave
Research Foundation which awarded me a
Karst Research Fellowship Grant for the
summer of 1985, and my parents, Jane and
Luther Yager. Dennis Williams has contin-
ually supported this research by helping in
the field as a cave diving partner, by de-
signing and maintaining diving equipment,
by organizing expeditions, and by sharing
his wisdom about the caves. In Abaco we
had the enthusiastic and generous support
of Ian Lothian, Donna and Barry Albury,
Maria and Lesley Albury. Keith Williams
helped move diving equipment and assisted
with field work. I appreciate the help of the
following members of the East End Expe-
dition to Sweeting’s Cay: Howard Cosgrove,
Sarah Cunliffe, David Knowles, Rob Palm-
er, Pernell Poitier, Rob Parker, and Dennis
Williams. I would like to thank Keith A.
Carson for the use of the electron micros-
copy facilities at Old Dominion University
and Carol Weidman who helped take the
SEM’s and prepare the photographic plates.
Fredrick R. Schram made available two
specimens of Cryptocorynetes previously
collected under National Science Founda-
tion Grant BSR 82-12335. John R. Holsin-
ger and Thomas E. Bowman reviewed the
manuscript and gave many helpful sugges-
320
tions. Thomas E. Bowman, Jerry H. Car-
penter, John R. Holsinger, Louis S. Kor-
nicker, and Ray S. Birdsong identified some
of the associated fauna. Katherine D. Payne
and Dirk E. Peterson inked some of the
drawings; Yvonne Smith helped prepare
some of the figures. All cave diving equip-
ment and techniques used for this research
met the standards of the U.S. National Spe-
leological Society Cave Diving Section.
Literature Cited
Arthur, Michael A., and Seymour O. Schlanger. 1979.
Cretaceous “‘oceanic anoxic events’ as causal
factors in development of reef-reservoired giant
oil fields. — American Association of Petroleum
Geologists Bulletin 63(6):870-885.
Cunliffe, Sarah. 1985. The flora and fauna of Sagit-
tarius, an anchialine cave and lake in Grand
Bahama.—Cave Science 12(3):103-109.
Degens, Egon T., and Peter Stoffers. 1976. Stratified
waters as a key to the past.— Nature 263:22-27.
Garcia-Valdecasas, Antonio. 1984. Morlockiidae new
family of Remipedia (Crustacea) from Lanza-
rote (Canary Islands).—Eos 60:329-333.
Holsinger, John R., and Jill Yager. 1985. Anewgenus
and two new species of subterranean amphipod
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
crustaceans (Hadziidae) from the Bahamas and
Turks and Caicos Islands.—Bidragen tot de
Dierkunde 55(2):283-294.
Iliffe, Thomas M., Horst Wilkens, Jakob Parzefall, and
Dennis Williams. 1984. Marine lava cave fau-
na: composition, biogeography, and origins. Sci-
ence 225:309-311.
Schlager, W., and R. N. Ginsburg. 1981. Bahama
carbonate platforms: The deep and the past.—
Marine Geology 44:1-24.
Schram, Frederick R., Jill Yager, and Michael J. Emer-
son. 1986. Remipedia. Part 1. Systematics. —
San Diego Society of Natural History Memoir
15:1-60.
Sket, Boris, and Thomas M. Iliffe. 1980. Cave fauna
of Bermuda.—International Revue der ge-
samten Hydrobiologie 85(6):87 1-882.
Yager, Jill. 1981. Remipedia, a new class of Crustacea
from a marine cave in the Bahamas.—Journal
of Crustacean Biology 1(3):328-333.
, and Frederick R. Schram. 1986. Lasionectes
entrichoma, new genus, new species, (Crustacea:
Remipedia) from anchialine caves in the Turks
and Caicos, British West Indies.— Proceedings
of the Biological Society of Washington 99(1):
65-70.
Department of Biological Sciences, Old
Dominion University, Norfolk, Virginia
23508.
PROC. BIOL. SOC. WASH.
100(2), 1987, pp. 321-336
A REVIEW OF RECENTLY DISCOVERED TYPE SPECIMENS
OF BERMUDA AMPHIPODA (CRUSTACEA: PERACARIDA)
DESCRIBED BY B. W. KUNKEL (1882-1969)
Eric A. Lazo-Wasem and Michael F. Gable
Abstract. — Type specimens of six of the seventeen species of Bermuda am-
phipods described by Kunkel (1910) are reviewed: Maera tinkerensis, Melita
planaterga, Parhyalella batesoni, Eurystheus [=Gammaropsis] lina, Isaea lon-
gipalpus, and Caprella bermudia. The first four listed species are redescribed
and refigured. The lectotype and recently collected specimens of Melita plan-
aterga are compared with Karaman’s (1981) redescription of the species. The
authors conclude that the specimen used by Karaman represents a new species.
Caprella bermudia is synonymized with C. equilibra.
In 1898 and 1901 A. E. Verrill conducted
scientific expeditions to the Bermuda is-
lands and amassed large collections of ter-
restrial and marine invertebrates. Verrill
himself reviewed some of the crustaceans,
mostly decapods, in several papers. To one
of his students, Beverly Waugh Kunkel,
Verrill gave the task of elucidating the status
of the Bermuda amphipod fauna. Verrill’s
collections, along with those of G. Brown
Goode and others, formed the basis of Kun-
kel’s (1910) monograph on the Bermuda
amphipods. Prior to this publication, the
amphipods of Bermuda were virtually un-
known and, to date, Kunkel’s paper remains
the only significant study of its kind. John-
son’s (1986) eclectic summary of the Ber-
muda amphipods provides little additional
information and leaves most of the long-
standing taxonomic questions untouched.
Kunkel described 17 new species, 16
gammaridean and | caprellidean, and erect-
ed three new genera in his monograph. Un-
fortunately, he did not clearly designate type
specimens, a fact that made their recovery
from the general collections difficult and at
times serendipitous. E. L. Mills (1964) re-
viewed the status of six of Kunkel’s type
specimens. The present paper reviews six
additional species for which the type spec-
imens have been found. Because the first
author has culled all the amphipods from
YPM’s crustacean collection, including all
unidentified specimens, virtually no possi-
bility exists for finding type material of the
remaining species described by Kunkel. This
situation is unfortunate as one species, /n-
sula antennuella, may have been incorrectly
figured (Barnard 1969) and was the basis for
a new genus.
In general, the descriptions of Kunkel (es-
pecially the figures) were not executed with
sufficient detail to facilitate contemporary
comparison with other known species. Re-
vised descriptions of the recently discov-
ered type specimens are desirable, therefore,
before the description of new taxa compli-
cates the already confused state of the Ber-
muda amphipod fauna.
Systematics
The family categories used herein follow
Bowman and Abele (1982). Legend: Body
parts marked by abbreviation beginning with
uppercase letters, and lowercase letters sep-
arated from body part indications read as
follows: A, antenna; Gn, gnathopod; Hd,
head; LL, lower lip; Md, mandible; Mx,
maxilia; Mxpd, maxilliped; T, telson; U,
322
uropod; dm, damaged; plp, palp; s, setae
missing.
Melitidae Bousfield, 1973
Maera tinkerensis Kunkel
Figs. 1-2
Maera tinkerensis Kunkel, 1910:49—51, fig.
18.—Barnard, 1962:100.
Description. —Male: Dorsal surface of
body finely setose, pleonal epimera 1-3 each
with small ventroposterior tooth, epimeron
1 with 1 ventral spine, epimera 2-3 with 3-
4 ventral spines.
Head subequal to combined length of pe-
reonites | and 2, lateral lobe broadly round-
ed, eye weakly pigmented in alcohol-pre-
served specimens, ocelli separated.
Antenna | longer than antenna 2, nearly
50% body length, peduncular article 1
somewhat shorter than article 2, posterior
margin with a few spines, article 3 20% ar-
ticle 2, primary flagellum composed of at
least 10-13 articles (terminal article(s?)
missing), accessory flagellum 6-articulate,
terminus minute. Antenna 2, peduncle ex-
tending nearly to length of peduncle of an-
tenna 1, flagellum short, composed of 6 ar-
ticles.
Left mandible with cuspate lacinia, prox-
imal spine row with | large cuspate spine
followed by 8 smaller, plumose spines; palp
slender, article 1 with distal tooth, 33%
length article 2, posterior margin article 2
setose, longer than article 3. Maxilla 1, inner
plate distally with 1 short spine and 2 long
plumose setae, outer plate with 8 stout, re-
curved, bifid spines; palp 2-articulate. Max-
illa 2, inner plate less broad than outer plate.
Gnathopod 1, ventroanterior corner of
coxal plate acutely produced, posterior mar-
gin of basis with long setae, article 3 with
plumose setae on posterior margin, article
5 triangular, longer than article 6, posterior
margin and inner facial surface densely se-
tose, article 6 oval, palm oblique, margin
lined with small spines, dactyl slender, tip
bearing small spatulate process. Gnathopod
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
2 much larger than gnathopod 1, coxa
square, corners rounded, posterior margin
of basis with 2 spines and a few setae, article
5 30% length article 6, posterior lobe dense-
ly setose, article 6 oval, much longer than
broad, palm oblique, elongate, sinusoidal
and distally serrate, posteriorly defined by
sharp tooth bearing spine at base, posterior
margin lined with submarginal spines and
a few setae, inner facial surface with long,
stout spine, dactyl stout, with 4 setae on
anterior margin, tip bearing small spatulate
process, dactyl closing between palmar tooth
and inner facial spine.
Pereopods 3—4 similar, normal, coxa 4
not posteriorly excavate; pereopod 5 shorter
than pereopods 6—7; pereopods 6—7 similar,
subequal in length, pereopod 7, proximo-
posterior corner of basis produced into
rounded lobe; all dactyls singly annulate.
Uropod | extending slightly beyond uro-
pod 2, dorsal surface of peduncle with row
of short and row of long spines, distal spines
of both rows longest and very stout, ventral
surface with large proximal spine, rami
shorter than peduncle, spinose, outer ramus
shorter than inner. Uropod 2, peduncle
stout, shorter than rami, with 2 rows of
spines, rami spinose, outer ramus shorter
than inner. Uropod 3 missing from speci-
men.
Telson longer than broad, deeply exca-
vate, lobe apices pointed, outer margins
notched, stout spines submarginally at base
of each notch.
Remarks. —Kunkel’s original description
agrees fairly well with the type specimens
but lacks considerable detail, most notably
in uropod spination, the details of the
gnathopods, and in precise length of spec-
imens. The uropods are much more spinose
than as figured by Kunkel, and the lengths
of the terminal spines on uropod 2 were
exaggerated. Kunkel drew the palm of
gnathopod 2 crenulate along its entire length,
whereas in the type material the crenulation
extends only 4 the length beginning at the
distal end.
VOLUME 100, NUMBER 2
323
Fig. 1. Maera tinkerensis Kunkel: YPM 8220, male syntype, 8.4 mm.
Material examined.—YPM 8220, male,
6.7 mm, syntype, Harrington Sound, Ber-
muda, collector and date unknown. — YPM
8221, male, 8.4 mm, syntype, Harrington
Sound, Bermuda, collector and date un-
known.— YPM 8222, male, 6.3 mm, syn-
type, Harrington Sound, Bermuda, collector
and date unknown.
Melita planaterga Kunkel
Figs. 3—4
Melita planaterga Kunkel, 1910:34—37, fig.
12.—Barnard, 1962:107-108.
Melita planaterga (?): Karaman, 1981:29-
50.
Description. —Male: Head, lateral lobes
Fig. 2. Maera tinkerensis Kunkel: YPM 8220, male syntype, 8.4 mm: Al, A2, Gn2, Md, T, U1 and U2.
YPM 8221, male syntype, 6.7 mm: Mxl, Mx2.
VOLUME 100, NUMBER 2
rounded, eyes not distinct in alcohol-pre-
served specimens. Pleon dorsally smooth,
epimeron 3 produced behind to form large
tooth, urosome segment | with short medial
mucronation on posterior margin, urosome
segment 2 with lateroposterior spine.
Antenna | greater than 50% body length,
peduncular article 1 stout, with 2 proximal
and 1 distal spines on posterior margin,
article 2 120% length article 1, article 3 33%
length article 2; flagellum longer than pe-
duncle, accessory flagellum 3-articulate, ter-
minus minute. Antenna 2 slightly shorter
than antenna 1.
Mandible with 4-cuspate lacinia, molar
moderately strong; mandibular palp article
1 60% article 2, article 2 30% longer than
article 3. Maxilla 1, inner plate truncate,
with distal setae; outer plate with 7 stout,
recurved, bifid spines; palp 2-articulate, dis-
tally moderately serrate, with stout spines
and a few setae. Maxilla 2, inner plate as
broad as outer plate, inner marginal setae
extending proximally * the distance of me-
dial margin, outer plate distally setose, a few
setae plumose.
Gnathopod 1, basis with 2 long setae on
anterior margin near base, distal half an-
terior margin with dense groups of long, ex-
ceptionally fine setae, article 4, distal por-
tion extended into distinct, slightly excavate
margin with setae, article 5 much longer
than broad, anterior margin lined with
groups of setae, posterior margin with 8 or
9 fascicles of long setae, inner facial margin
with 4 groups of setae, article 6 shorter than
article 5, distoposterior corner produced into
rounded lobe with setae, palm incised near
base of article 7, base of dactyl inflated, an-
terior margin with small distal spine.
Gnathopod 2, anterior margin of basis with
groups of long setae, articles 3 and 4 sub-
rectangular, article 4, distoposterior corner
produced into small tooth, article 5, ante-
rior and posterior margins densely lined with
fascicles of setae, setae on posterior margin
serrate, inner facial margin with several
groups of long setae, article 6, length 170%
width, anterior and posterior margins
325
densely lined with long setae, palm oblique,
excavated margin result of damage, lined
with plumose setae, inner facial surface ad-
jacent to palm forming depression, article
7 longer than palm, closing against facial
depression of article 6.
Pereopods 5-7, bases of anterior and pos-
terior margins finely serrate, serrations de-
fined by small spines.
Uropod | extending slightly beyond uro-
pod 2, peduncle spinose, slightly longer than
rami, ventral margin with | stout spine, rami
spinose, rami equal, uropod 2, peduncle and
rami spinous, rami equal to each other and
to peduncle, uropod 3 missing from speci-
men.
Telson split nearly to base, each lobe
bearing 1—3 terminal spines and 0-1 spines
along inner margin.
Remarks. — Kunkel described the palm of
gnathopod 2 as slightly concave. The dactyl
in his figure, however, obscures the margin
of the palm. Our redescribed gnathopod 2
of the lectotype shows a concavity (Fig. 3).
Examination of recent material from Ber-
muda, however, suggests that the normal
condition of the palmar margin is entire,
not concave, and of the palmar setae, long
and plumose, not short.
Karaman (1981) redescribed this species
from a single male specimen collected at
Castle Harbour, Bermuda, noting many dif-
ferences between his specimen and Kun-
kel’s. The type material of Kunkel, how-
ever, confirms many points of his original
description and brings to light more differ-
ences from the specimen described by Kar-
aman.
The dorsal mucronation on urosome seg-
ment 1, as figured by Karaman, is much
more prominent than on the type specimen;
its mucronation extends approximately %
the length of urosome segment 2 whereas
Karaman’s figure shows it extending nearly
% the length of urosome segment 2. More-
over, on Kunkel’s specimen we find no notch
at the insertion of the dorsolateral spine on
the posterior margin of urosome segment 2.
Karaman figured a strongly dentate palp
326 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig. 3.
article 2 on maxilla 1. In the lectotype the
distal margin of the maxilla 1 palp is only
moderately dentate; the teeth are acutely
rounded and not pointed as in Karaman’s
specimen.
Kunkel stated that the accessory flagel-
Melita planaterga Kunkel: YPM 8227, male lectotype, 5.4 mm.
lum of antenna | is 2-segmented. Careful
examination of the type revealed that the
accessory flagellum is actually 3-segmented;
the terminus is miniscule and was certainly
overlooked by Kunkel. Karaman, however,
clearly figured a 4-segmented accessory fla-
VOLUME 100, NUMBER 2 327
Fig. 4. Melita planaterga Kunkel: a: YPM 8227, male lectotype, 5.4 mm; b: YPM 8229, male, 4.5 mm.
328 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
gellum with a large terminal segment. In
specimens examined from material recently
collected at several localities in Bermuda,
all had an accessory flagellum as we have
figured; none had 4 segments as stated by
Karaman.
Examination of these recently collected
specimens supports the differences between
Kunkel’s and Karaman’s specimens in oth-
er aspects. The posterior spination of the
first peduncular segment of antenna | is in-
variably as Kunkel illustrated as is the inner
marginal spination of the telson. Further-
more, the spines on the peduncle and rami
of uropods 1 and 2, although variable in
number, are always significantly less than
the number shown by Karaman and again
agree with Kunkel’s figure. Kunkel did not
figure uropod 3 and his description 1s vague,
but the third uropods of recent specimens
(Fig. 4b) differ from those of Karaman’s
specimen; the peduncle in his specimen. is
less spinose and the rami more spinose than
in the recent material. Finally, the serrate
margin of the basis on pereopods 5—7 men-
tioned by Kunkel but not seen by Karaman
is always present and distinct in all speci-
mens we examined.
Karaman’s specimen, which differs in so
many characteristics, undoubtedly repre-
sents a new species; without seeing his spec-
imen, however, we feel it prudent not to
erect a new taxon.
Material examined.—YPM 8227, Flatts
Village, Bermuda, collector and date un-
known, | lectotype male (5.4 mm).— YPM
8229. Whalebone Bay, St. George’s, Ber-
muda, M. F. Gable and class, 22 May 1985,
intertidal, among rocks and vegetation, 1
male [uropod 3 figured].—YPM 8230,
Whalebone Bay, St. George’s, Bermuda,
M.F. Gable and class, 22 May 1985S, inter-
tidal, among rocks and vegetation, 2 males,
2 females (ovigerous).—YPM 8231, Ferry
Reach, Bermuda, north of Long Bird Bridge,
M.F. Gable, 29 May 1985, intertidal to sub-
tidal, 1 male, 1 female, 3 juveniles.— YPM
8232, Devonshire Bay, Bermuda, M.F. Ga-
ble, 29 May 1985, under rocks, shallow sub-
tidal, 2 males, 2 females, 2 ?? [damaged
specimens].—YPM 8233, Gravelly Bay,
Bermuda, M.F. Gable, 31 May 1985, under
rocks in subtidal honeycombed substrate, 2
females (1 ovigerous).—USNM Acc. No.
346847, Hungry Bay, Bermuda, M.L. Jones,
5 Sep 1981, from transect collection along
length of bay, West Seawall, associated with
rocks, 1 male, 1 female (ovigerous).
Hyalellidae Bulycheva, 1957
Parhyalella batesoni Kunkel
Figs. 5-6
Parhyalella batesoni Kunkel, 1910:74—-76,
fig. 28.
Description. —Male: Body _ dorsally
smooth, coxa deep, anterior margins round-
ed. Pleonal epimera subquadrate, posterior
corners not produced.
Head, lateral lobes rounded; eyes mod-
erately large, oblong. Antenna 1 25% of body
length, slightly longer than antenna 2, fla-
gellum 110% length of peduncle, consisting
of 11 to 15 articles; peduncular segments
subequal in length, accessory flagellum lack-
ing. Antenna 2 nearly 25% of body length,
peduncular segments much broader than
those of antenna 1, gland cone hidden, seg-
ment 3 short and stout, as long as broad,
segment 4 as broad as 3, length of segment
4 to 5, 1:0.5, segment 5 narrower than 3 and
4; flagellum short, 25% length of peduncle,
3-articulate, basal article conjointed.
Maxilla 1, inner plate narrow, with short
medial setules and 2 stout plumose terminal
spines; outer plate with 8 stout, serrate
spines, lateral margin distinctly excavate,
pre-ecdysial serrate spines visible through
facial surface; palp absent.
Gnathopod 1 subchelate, much smaller
than gnathopod 2, posterior margin of ar-
ticle 2 produced, broadest point with 3 large
and | small setae; article 5 nearly as broad
and as long as article 6, posterior lobe with
ridge of small spines, article 6 rectangular,
posterior corner of transverse palm with
small to medium spines and a few setae,
dactyl short, stout, congruent with palm.
VOLUME 100, NUMBER 2
Llp’ oor “ang
ee MD ry A] oa
“STEER Ut
329
My
Fig. 5. Parhyalella batesoni Kunkel: YPM 8188, male holotype, 6.8 mm.
Gnathopod 2, powerful, subchelate, article
2 with anterodistal lobe; articles 3 and 4
normal, 3 with mammilliform anterior lobe;
article 5 less than 20% length article 6, pos-
terior lobe strong, distally produced be-
tween articles 4 and 6, lined with row of
spines, article 6 equal to width of article 5,
Oval, palm oblique, margin incised, sub-
marginally lined with stout spines, larger
spine at proximoposterior corner, dactyl
congruent with palm.
Pereopods 3 and 4 normal, dactyls 1-an-
nulate. Pereopods 5-7 similar, length in-
creasing consecutively; article 2 expanded
in all.
Uropod 1 extending beyond uropod 2,
peduncle spinose, rami subequal, each with
spines along dorsal margin and four ter-
330
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig. 6. Parhyalella batesoni Kunkel: YPM 8188, male holotype, 6.8 mm.
VOLUME 100, NUMBER 2
minal spines. Uropod 2 75% length uropod
1, peduncle with row of spines; outer ramus
shorter than inner, both with marginal and
terminal spines. Uropod 3, peduncle with
1 prominent spine at inner posterior corner,
inner ramus absent, outer ramus stout, 40%
length of peduncle, with 1 terminal spine.
Remarks. —The genus Parhyalella is di-
agnosed in part by the absence of a palp on
maxilla 1. The type specimen lacked nearly
all the mouthparts but fortunately still re-
tained a single maxilla 1 so its form could
be confirmed. Several species assigned to
this genus, however, have been clearly de-
scribed as having a palp, albeit vestigial, on
maxilla 1. It is apparent that the species of
Parhyalella need to be re-examined and the
generic diagnosis re-evaluated.
Recent shallow water collections made in
Bermuda by one of us (MFG) failed to turn
up any additional specimens of P. batesoni.
As the type specimen had no collection data,
One can only speculate upon the habitat of
this species. Several individuals of Parhy-
alella were recently collected at 20 m from
Taiwan (in prep.); it is possible that P. bate-
soni also inhabits deeper water and may only
rarely be collected in shallow water.
Material examined.—YPM 8188, Ber-
muda, collector and date unknown, 1 ho-
lotype male (6.8 mm).
Corophiidae Dana, 1849
Gammaropsis lina (Kunkel)
Figs. 7—9
Eurystheus lina Kunkel, 1910:81-83, fig. 31.
Eurystheus = Gammaropsis [in part], Bar-
nard, 1969:271.
Description. —Male: Coxae moderately
deep, overlapping. Pleon segments 1-3 each
with one pair dorsolateral setae on posterior
Margin; posterior margin pleonal epimera
broadly rounded, hind corner produced into
small tooth. Urosome segment | with pair
of 1 long and 1 short dorsolateral setae; uro-
some segment 2 with pair of 1 long and 2
short, dorsolateral setae.
Eye oval, situated in head lobe; head lobe
331
acute. Antenna 1 50% body length, pedun-
cular segment length ratio 1:1.4:1; pedun-
cular segment | twice as broad as segments
2 and 3, posterior margin of segments 2 and
3 densely setose; flagellum composed of 12
articles, 30% length of peduncle; accessory
flagellum 5 articulate, terminus minute. An-
tenna 2 slightly shorter than antenna |; pe-
duncular segment 4 shorter than 5, posterior
margin of both articles strongly setose; fla-
gellum 40% length of peduncle, flattened,
Ist article elongate.
Right mandible with serrate lacinia, 9
spines proximal to incisor, molar strong,
ridged, with plumose accessory seta; palp
3-articulate, Ist article 25% length article 2,
article 2 stout, subequal in length to article
3, posterior margin slightly excavate and
setose, anterior margin article 3 with 3 long
setae, distal margin armed with long, plu-
mose setae. Lower lip with inner lobes well
developed, outer lobes with pointed lateral
projections. Maxilla 1, inner plate with plu-
mose marginal setae, outer plate with about
9 stout, bifid spines; palp 2-articulate, distal
margin of second article with short, stout
spines. Maxilla 2, outer plate a little broader
than inner, inner plate with row of facial
setae. Maxilliped normal, palp 4-articulate,
terminus bearing long distal setae.
Gnathopod 1 subchelate, smaller than
gnathopod 2, article 4 distally rounded with
long posterior setae, article 5 slightly longer
than article 6, posterior margin and medial
face with fascicles of long setae, article 6
ovate, fascicles of setae on posterior and
anterior margins, palm oblique, with stout
spines at proximoposterior corner. Gnatho-
pod 2, article 5 33% article 6, posterior lobe
prominent, article 6 elongate, 2.5 times
longer than wide, rectangular, posterior
margin densely lined with curly setae, palm
transverse, convex, distoposterior corner
produced into strong thumb, article 7 stout,
with posterior tubercle hidden by long setae
of article 6.
Pereopod 4 (pereopod 3 missing), coxal
plate with long seta at distoanterior corner,
basis with long seta on posterior margin;
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
332
\
\
\
)_ AA
\
©,
Oi Yl
a Yoo UippZ
ty pe Bij
Via a Mi
rt aA \\I
Ui
ial {
SN
rSQv
————
—
SX
es
_&
LE
Z>
——_
Fig. 7. Gammaropsis lina (Kunkel): YPM 8189, male syntype, 6.1 mm: Al, Gnl, Gn2; A2 from unattributed
specimen. YPM8224, female syntype, 5.1 mm: Gn2.
VOLUME 100, NUMBER 2
= Mxt
Woes
333
WANK
ai { | ‘
t pe
a hy
v f ) ;
K\ \y
Fig. 8. Gammaropsis lina (Kunkel): YPM 8189, male syntype, 6.1 mm.
pereopod 5, basis somewhat expanded, ar-
ticles 6 and 7 appearing antero-facing be-
cause of limb splaying; pereopod 6, coxa
with spine at distoposterior corner, dactyl
bifid; pereopod 7, proximoposterior corner
of basis produced acutely, articles 6 and 7
appearing antero-facing because of limb
splaying.
Uropods extended behind subequally;
uropod |, interramal spine extending nearly
334 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig. 9.
’, length outer ramus, rami longer than pe-
duncle, outer ramus 80% inner ramus, both
spinose; uropod 2 spinose, outer ramus 80%
inner ramus; uropod 3, peduncle and rami
subequal, rami with long distal spines.
Telson emarginate, apices separated, with
1—2 short distal spines.
P7 iw
Pane
Gammaropsis lina (Kunkel): YPM 8189, male syntype, 6.1 mm.
Female: Syntypic female very similar to
male except in form of gnathopods.
Gnathopod | smaller than gnathopod 2, an-
terior margin moderately setose, posterior
margin with fascicles of setae, palm oblique,
lined with setae, distoposterior corner with
stout spine, posterior margin article 7 ser-
VOLUME 100, NUMBER 2
Fig. 10. Caprella bermudia Kunkel: YPM 8223, male lectotype, 6.3 mm: outer view (dactyl removed) and
inner view.
rate. Gnathopod 2, article 6 subrectangular,
anterior margin densely lined with setae,
posterior margin lined with fascicles of curly
setae; palm oblique, moderately crenulate,
distoposterior corner produced into weak
thumb, and armed with stout spine, article
7 serrate posteriorly.
Remarks. —The type material agrees
closely with Kunkel’s description in major
diagnostic features; Kunkel’s figures, how-
ever, lacked detail, particularly those of the
mouthparts and the gnathopods. He also
did not figure any of the pereopods or uro-
pods. In the type material there are three
specimens and not two as he stated; none
of the specimens retained an intact antenna
2, and, therefore, one antenna for figuring
was selected from several found loose inside
the original vial.
Material examined.—YPM 8189, Ber-
muda, Dr. J. L. Cole, 1903, 1 syntype male,
6.1 mm.— YPM 8224, Bermuda, J. L. Cole,
1903, 1 syntype female, 5.1 mm.— YPM
8225, Bermuda, J. L. Cole, 1903, 1 syntype
male [damaged, lacking head].
Tsaea longipalpus Kunkel
Isaea longipalpus Kunkel, 1910:85-87, fig.
Bo:
Remarks. —The type material consists of
only a few rehydrated fragments (YPM
8226) from Bailey Bay, Bermuda; their poor
condition is not satisfactory for redescrip-
tion so Kunkel’s description must stand un-
til new material can be evaluated.
Caprellidae White, 1847
Caprella bermudia Kunkel
Fig. 10
Caprella bermudia Kunkel, 1910:108—110,
fig. 42.—McCain, 1968:22; McCain and
Steinberg, 1970:13.
Remarks. — Although authenticated type
material of C. bermudia was originally not
found, one lot of C. equilibra Say, obviously
examined by Kunkel, contained an individ-
ual matching his description and figures for
C. bermudia precisely. This specimen con-
sisted of a dissected head and first pereon
segment and a matching carcass complete
with gnathopod 2. The gnathopod 2 de-
scribed and figured by Kunkel for C. ber-
mudia is identical to gnathopod 2 of this
specimen (Fig. 10). Furthermore, the dis-
section cleavage, directly posterior to the
suture between the head and pereon seg-
ment | is identical with Kunkel’s figure.
It is not readily apparent what characters
Kunkel used to diagnose C. bermudia. He
averred that the diagnostic character of C.
336
equilibra is the strong spine projecting ven-
trally between the gnathopods on pereonite
2. The specimen mentioned above has only
a very small ventral spine between the
gnathopods on pereonite 2 whereas the
specimens called C. equilibra by Kunkel
possess a very large, prominent ventral
spine. On both a specimen recently collect-
ed from Bermuda and a specimen found in
the USNM collections there was also a very
large ventral spine on pereonite 2 even
though the specimens were extremely small.
It is conceivable, therefore, that Kunkel sep-
arated the two species by the appearance of
the ventral spine, as no other distinctions
are apparent. McCain (1968) mentioned that
a variant of C. equilibra from North Car-
olina lacks the prominent ventral spine on
pereonite 2; obviously, Kunkel was not
aware of variants.
Kunkel apparently figured a female C.
equilibra and a male caprellid, calling it C.
bermudia. McCain (1968) concluded there
is no way to separate C. bermudia from C.
equilibra. Kunkel’s figured specimen of C.
bermudia is clearly indistinguishable from
C. equilibra (C. equilibra in McCain 1968,
Laubitz 1970). We believe, therefore, that
the synonymy of the two is warranted.
Material examined.—YPM 8223, Ber-
muda, Dr. J. L. Cole, 15 Jul 1903, lectotype
male (6.3 mm).— YPM 8208, Caprella equi-
libra (Say), Flatts Village, Bermuda, collec-
tor unknown, 4 Jul 1898, 2 males, 1 fe-
male.— YPM 8265S, C. equilibra, Shelly Bay,
Bermuda, M. F. Gable, 3 Jun 1985, among
hydroids, 1 male.—USNM collection, C.
equilibra, Bermuda, G. Brown Goode, 1876—
1877, 1 ovigerous female.
Acknowledgments
The authors wish to thank Dr. J. L. Bar-
nard (USNM) for the loan of material. This
study has been supported in part by a Con-
necticut State University Research Grant to
the second author. This paper is Contri-
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
bution No. 1118 from the Bermuda Biolog-
ical Station for Research.
Literature Cited
Barnard, J. L. 1962. Benthic marine Amphipoda of
southern California: Families Tironidae to
Gammaridae.— Pacific Naturalist 3:73-115.
1969. The families and genera of marine
Gammaridean Amphipoda.—Bulletin of the
United States National Museum 271:vi, 1-535.
Bowman, T. E., and L.G. Abele. 1982. Classification
of the Recent Crustacea. Pp. 1-24 in D. E. Bliss,
ed., The biology of Crustacea, vol. 1. L. G. Abele,
Systematics, the fossil record, and biogeogra-
phy. Academic Press, New York, xx + 319 pp.
Johnson, S. E. 1986. Order Amphipoda. Pp. 372-
381 in W. Sterrer, ed., Marine fauna and flora
of Bermuda. John Wiley & Sons, New York.
Karaman, G.S. 1981. Redescription of Melita plan-
aterga Kunkel 1910 from Bermuda islands with
revision of genera Melita Leach and Abludo-
melita n. gen.—Poljoprivreda 1 Sumarstvo 27:
29-50.
Kunkel, B. W. 1910. The Amphipoda of Bermuda. —
Transactions of the Connecticut Academy of Arts
and Sciences 16:1-116.
Laubitz, D. 1970. Studies on the Caprellidae (Crus-
tacea, Amphipoda) of the American North Pa-
cific. — National Museum of Natural Sciences,
Ottawa, Publications in Biological Oceanogra-
phy 1:i—vu, 1-89.
McCain, J.C. 1968. The Caprellidae (Crustacea: Am-
phipoda) of the western North Atlantic. —Bul-
letin of the United States National Museum 278:
i-vi, 1-147.
,and J. E. Steinberg. 1970. Caprellidea I. Fam-
ily Caprellidae. Pp. 1-78 in H.-E. Gruner and
L. B. Holthuis, eds., Crustaceorum Catalogus,
Amphipoda I, part 2. Dr. W. Junk Publishers,
The Hague, The Netherlands.
Mills, E. L. 1964. Noteworthy Amphipoda (Crusta-
cea) in the collection of the Yale Peabody Mu-
seum.—Postilla 79:1-41.
(EAL-W) Division of Invertebrate Zool-
ogy, Peabody Museum of Natural History,
Yale University, 170 Whitney Avenue, P.O.
Box 6666, New Haven, Connecticut 0651 1-
8161; (MFG) Department of Biology, East-
ern Connecticut State University, Willi-
mantic, Connecticut 06226-2295.
PROC. BIOL. SOC. WASH.
100(2), 1987, pp. 337-346
TWO NEW SPECIES OF ELEUTHERODACTYLUS
(AMPHIBIA: ANURA: LEPTODACTYLIDAE)
FROM BOLIVIA
John D. Lynch and Roy W. McDiarmid
Abstract. —Eleutherodactylus fraudator, new species, E. mercedesae, new
species, and E. rhabdolaemus are reported from cloud forests in Departamento
Cochabamba, Bolivia. Eleutherodactylus pharangobates Duellman is placed as
a synonym of E. rhabdolaemus Duellman.
Although the genus Eleutherodactylus is
extraordinarily speciose in many areas of
northern South America and especially on
the Andean slopes, it is poorly known in
southern South America. Presently, the
southern terminus of the genus is in north-
ern Argentina (Lynch 1971, Cei 1980) where
only a single species, Eleutherodactylus dis-
coidalis, is recognized. Three species have
been reported from Andean Bolivia in con-
trast to ten species recorded from adjacent
areas in Peru. While such a gradient is con-
sistent with the geographic pattern of di-
versity characteristic of many groups of
tropical organisms, with frogs (especially
Eleutherodactylus) this gradient is partially
a reflection of collecting efforts. Thus, a se-
ries of small collections made by Mercedes
S. Foster in 1979 as she travelled down the
eastern versant of the Bolivian Andes along
the road from Cochambamba to Villa Tu-
nari (Cannatella [1980] presented a useful
map of the area) are especially noteworthy
for the frogs they contain. Her collections,
when added to material collected in 1959
from the same area, provide us with some
important additions to the Bolivian her-
petofauna particularly within the genus
Eleutherodactylus.
The three species previously reported
from Bolivia are E. cruralis (Boulenger), E.
fenestratus (Steindachner), and EL. pharan-
gobates Duellman. The cloud forests of ad-
jacent Peru (Departamentos Ayacucho,
Cuzco, and Huanuco) harbor ten species:
E. cosnipatae Duellman, E. danae Duell-
man, E. granulosus (Boulenger), E. lindae
Duellman, &. mendax Duellman, E. phar-
angobates Duellman, E. platydactylus
(Boulenger), E. rhabdolaemus Duellman, E.
salaputium Duellman, and E. scitulus
Duellman.
In addition to the two new species of
Eleutherodactylus (described below) and E.
rhabdolaemus, Foster obtained one speci-
men of EL. fenestratus (USNM 257849, Mi-
guelito, ca. 99 km from Cochabamba on
Cochabamba to Villa Tunari road, Provin-
cia Chapare, Departamento Cochabamba,
1770 m). This record is from the southern
edge of the distribution of EF. fenestratus, a
wide-ranging species recorded from several
localities in the southern Amazon Basin
(Lynch 1980).
Our use of abbreviations and terminology
follows Lynch (1980) and Lynch and Myers
(1983).
Eleutherodactylus fraudator, new species
Figs. 1, 2A
Holotype. —USNM 257847, adult male,
taken 73.5 km from Cochabamba on Co-
chabamba to Villa Tunari road, Provincia
Chapare, Departamento Cochabamba, Bo-
livia, ca. 2690 m, on 23 Sep 1979, Mercedes
S. Foster collector.
Paratopotype. —USNM 257846, adult
male, taken with holotype.
338
Diagnosis. —A species of Eleutherodac-
tylus distinguished from others by the fol-
lowing combination of characteristics: (1)
skin of dorsum shagreened, that of venter
smooth; low dorsolateral folds present; (2)
tympanum prominent, round, *% eye length;
(3) snout outline from dorsal view rounded
to obtuse, round in lateral profile; canthus
rostralis sharp in cross section; (4) upper
eyelid broader than IOD; no cranial crests;
(5) vomerine odontophores oval; (6) males
with vocal slits; males with non-spinous
nuptial pads; (7) first finger slightly longer
than second; tips of two outer fingers trun-
cate with expanded pads, those of inner fin-
gers more rounded; (8) lateral fringes on fin-
gers weakly defined or absent; (9) no ulnar
tubercles; (10) no tubercles or folds on heel
or tarsus; (11) inner metatarsal tubercle oval
and elevated, outer low and indistinct; no
supernumerary plantar tubercles; (12) toes
with weak lateral keels; toe tips rounded to
truncate; (13) brown with darker brown
stripes; dark brown canthal-supratympanic
and labial stripes; venter cream stippled with
brown, heaviest on throat; (14) two adult
males 28.1—28.3 mm SVL.
Eleutherodactylus fraudator tentatively is
assigned to the conspicillatus group. How-
ever, the plesiomorphic nature of that group
(Lynch 1986) prevents final determination
of relationships at this time. Eleutherodac-
tylus fraudator is readily distinguished from
all others species in the group by having a
continuous brown labial stripe instead of
labial bars or spots. Additionally, the oval
vomerine odontophores are distinctive.
Description. —Statements expressed in full
in diagnosis are not repeated here. Head as
wide as body, wider than long; HW 40.3-
40.6% SVL; nostrils protuberant, directed
laterally; snout obtuse, sloping abruptly an-
terior of nostrils, rounded below tip; can-
thus rostralis sharp, slightly concave to
straight; loreal region concave, sloping
abruptly to lips; lips not flared; E-N 75.0-
82.0% eye length; low tubercles on upper
eyelid; upper eyelid width 112.0-119.2%
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
IOD; relatively large frontoparietal fonta-
nelle present, possibly indicating that these
are young, albeit adult, males; supratym-
panic fold low, barely touching tympanum;
tympanum low on head, length 40.0—46.2%
of eye length, separated from eye by dis-
tance equal to its length; choanae ovoid,
moderate-sized, about equal to tip of first
finger, well separated from palatal shelf of
maxillary arch; vomerine odontophores oval
in outline, medial and posterior to choanae,
separated by distance about equal to width
of odontophore, bearing two to four teeth;
tongue longer than wide, oval, posterior edge
with shallow notch, posterior 3 free from
floor of mouth; vocal slits long, postero-
lateral to tongue; vocal sac subgular.
Skin of dorsum shagreened but with scat-
tered low tubercles; dorsolateral folds ex-
tend from above tympanum to above groin;
skin on flanks not areolate but with scat-
tered tubercles; distinct glandular area in
groin; no ventral discoidal folds; skin pos-
teroventral to vent areolate; no anal sheath;
palmar tubercle much larger than oval the-
nar, both low; subarticular tubercles low,
longer than wide; fringes indistinct (pres-
ence at some points along margins of some
fingers may be artifact of preservation); tips
of fingers III-IV (Fig. 2) expanded, truncate,
with enlarged pads, tips of II-IV with cir-
cumferential grooves; finger tip of III less
than 4 size of tympanum; finger lengths,
shortest to longest, 2, 1, 4, 3; males with
large (covers more than half length of digit),
white and glandular nuptial pad on top of
thumb; outer metatarsal tubercle low, '2 to
3 size of inner; subarticular tubercles longer
than wide, sub-conical; toes lack fringes and
webbing; lateral keels weakly developed; all
toes bear pads on: ventral surfaces; heels
broadly overlap when flexed hind legs held
at right angles to sagittal plane; shank 57.9-
60.8% SVL.
The paratype, dissected to determine the
condition of the m. adductor mandibulae,
has the ‘“‘S”’ condition (Lynch 1986).
Coloration in preservative. — Brown above
VOLUME 100, NUMBER 2
339
Fig. 1.
with darker brown interorbital triangle con-
tinuing posteriorly as a pair of stripes vague-
ly recalling scapular and sacral chevrons (Fig.
1); lateral margin of dorsolateral fold very
dark brown delineating a brown flank band
bordered ventrally by paler coloration; ir-
regular dark brown marks forming broken
stripe beginning above forelimbs and ex-
tending ventrally on anterior '4 of flank;
canthal stripe broad, dark brown, extending
posteriorly above and below eye to join su-
pratympanic stripe; supratympanic stripe
expanded ventrally to form blotch, edged
ventrally by cream line; tympanum dark,
continuous with supratympanic stripe, an-
nulus only slightly lighter; brown stripe on
margin of upper lip; limbs irregularly barred
(bars wider or narrower than interspaces,
darkest marginally); top of thigh barred;
posterior surface of thigh essentially dark
brown with some traces of lighter mottling;
groin pigmentless except in glandular area
that is ochre in one specimen; ventral sur-
faces cream dusted with brown, heaviest on
throat and chest.
Coloration in life. —Dorsally and laterally
Eleutherodactylus fraudator, left: holotype, USNM 257847; right: paratype, USNM 257846.
mottled with tan, light and dark brown, pat-
tern edged in black in some places; eye, lip,
and lateral stripes dark brown to black; ven-
trally creamy gold, more or less iridescent,
intermixed with brown; iris gold (from field
notes of M. S. Foster).
Measurements of holotype (and paratype)
inmm.—SVL 28.3 (28.1), shank 16.3 (17.1),
HW 11.4 (11.4), head length 10.3 (10.1),
chord of head length 11.4 (11.1), upper eye-
lid width 3.1 (2.8), IOD 2.6 (2.5), tympan-
um 1.8 (1.6), eye 3.9 (4.0), E-N 3.2 (3.0),
hind leg (vent to tip of fourth toe) 53.8 (53.8).
Etymology. — Latin, meaning cheat or de-
ceiver; used because these frogs look very
much like Gastrotheca marsupiata.
Referred specimen.—In addition to the
types, we are aware of one additional spec-
imen of E. fraudator. USNM 146568, a ju-
venile female 14.3 mm SVL, was taken at
Incachaca, Provincia Chapare, Departa-
mento Cochabamba, 2100 m, on 27 Oct
1959, R. B. Cumming. We prefer not to
include it in the type series because of its
juvenile nature.
Remarks. — These frogs were calling from
340
a road-cut in the late afternoon from under
rocks or in heavy “‘moss.”’ The road-cut, a-
sheer face, was very rocky and covered with
vegetation 3 to 5 cm tall interspersed with
many bare areas. Males, completely hidden,
gave single note, high pitched calls.
We considered the possibility that this
species might be identical with that named
Leptodactylus (Plectromantis) andicola by
Boettger (1891) from Sorata, Bolivia, a lo-
cality east of Lago Titicaca at about 2650
m on the eastern Andean slopes of Depar-
tamento La Paz. Heyer (1978:33) reported
that the type had been destroyed but placed
the species in the genus Eleutherodactylus.
Our comparison of the published descrip-
tion also indicates that several features are
in agreement with Eleutherodactylus, and
E. fraudator. However, the description is
somewhat general and the following points
are at variance (states for fraudator in pa-
rentheses): snout almost 1'4 times as long
as eye (about equal); IOD as wide as upper
eyelid (narrower); first finger as long as sec-
ond (longer); narrow fringes on fingers and
toes (indistinct or absent); fold on distal '4
of tarsus (no tarsal fold); upper lip blotched
(brown labial stripe); limbs indistinctly
banded (bands prominent). Additionally,
Boettger (1891) made no mention of dor-
solateral folds; given that such folds are im-
portant in the taxonomy of Leptodactylus,
the omission suggests that the type of an-
dicola lacked such folds. We conclude that
if andicola is an Eleutherodactylus, it is dis-
tinct from E. fraudator.
In reading the description of L. andicola
we were struck by its agreement with that
of Gastrotheca marsupiata. Although Heyer
(1978:33) placed andicola in Eleutherodac-
tylus, we think it equally likely that it might
be a Gastrotheca marsupiata. Although the
type is reported to be 48 mm SVL, some-
what large for a G. marsupiata (Duellman
and Fritts 1972:17), Sorata is within the
range of G. marsupiata. Also many other
traits, as described by Boettger (1891) for
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
andicola, are similar to those found in Gas-
trotheca.
Eleutherodactylus mercedesae, new species
Figs. 2B, C, 3
Holotype. —USNM 257848, adult male,
ca. 3.3 km N of Cochabamba to Villa Tuna-
ri road on road to San Onofre, at a point
97.5 km from Cochabamba on Cochabam-
ba to Villa Tunari road, Provincia Chapare,
Departamento Cochabamba, Bolivia, ca.
1690 m, taken 29 Oct 1979, Mercedes S.
Foster collector.
Paratype. —USNM 165753, Limbo, Pro-
vincia Chapare, Departamento Cochabam-
ba, Bolivia, 1950 m, probably collected 25
Oct 1959, Robert B. Cumming.
Diagnosis.—A species of Eleutherodac-
tylus distinguished from others by the fol-
lowing combination of characters: (1) skin
of dorsum finely granular, that of venter
areolate; irregular, indistinct dorsolateral
folds; (2) tympanum prominent, superficial,
3 eye length; (3) snout subacuminate in dor-
sal view, round in lateral profile; canthus
rostralis sharp; (4) [OD about equal to width
of upper eyelid; no cranial crests; (5) vo-
merine dentigerous processes large, nearly
triangular, just medial and posterior to
choanae; (6) males with vocal slits; males
lack nuptial pads; (7) first finger slightly
shorter than second; fingers long, tips trun-
cate, pads large; (8) lateral fringes on fingers;
(9) no ulnar tubercles; (10) fold along distal
’; of inner edge of tarsus; no heel or tarsal
tubercles; (11) two metatarsal tubercles, in-
ner large, oval, about four times size of low
outer; three small supernumerary plantar
tubercles; (12) toes with narrow lateral
fringes, basal webbing; toe tips truncate with
broad pads; (13) dorsum brown with lich-
enose pattern of cream (green in life); throat
streaked and mottled with cream and brown;
posterior surfaces of thighs brown with many
cream flecks and mottling; (14) adults large,
two males 40.1-—49.5 mm SVL.
By virtue of its color pattern, E. merce-
VOLUME 100, NUMBER 2
341
Fig. 2. (A) Hand of Eleutherodactylus fraudator (USNM 257847), (B) palate and (C) hand of E. mercedesae
(USNM 257848). (A) is a composite of the left and right hands of the holotype to remove distortions caused
by encysted parasites. Scales equal 2 mm.
desae is distinguished from every other
species in the genus. Although it has char-
acteristics of Lynch’s (1976) wnistrigatus
group, we are unable to suggest any relatives
at this time.
Description. —Statements expressed in full
in diagnosis not repeated here. Head equal
to or wider than body, wider than long; HW
36.4-39.7% SVL; snout deep and lacking
tubercles at tip; nostrils moderately protu-
berant, directed laterally, much nearer tip
of snout than to eye; canthus rostralis straight
or weakly sinuous; E-N 90.4—93.5% eye
length; loreal region concave, especially an-
teriorly, sloping abruptly to lip; small tu-
bercle in center of loreal region; lips not
flared; upper eyelids with few small tuber-
cles; upper eyelid width 118.2—120.0% IOD;
supratympanic fold distinct, contacting up-
per edge of tympanum, extending postero-
ventrally to above arm insertion; tympa-
num round, its length 36.5—37.1% eye length,
separated from eye by distance equal to or
slightly more than its diameter; two, low to
moderate postrictal tubercles posteroven-
tral to tympanum; scattered tubercles on
head (three or four on upper eyelid, one
between eyes, two preinterocular, one on
loreal surface, two behind eyes on a line
between tympana), but none on margin of
lower jaw; choanae relatively large, about
equal to subarticular tubercle width, ovoid,
not concealed by palatal shelf of maxillary
arch; vomerine odontophores between and
slightly posterior to choanae (Fig. 2B), nar-
rowly separated, slightly larger than choana,
342
approximately triangular in outline, much
elevated, bearing slanted row of six or seven
teeth; tongue longer than wide, somewhat
cordate, posterior border with shallow notch,
posterior '4 free from floor of mouth; mod-
erately long vocal slits lateral to posterior
half of tongue, slit covered posteriorly by
distinct labial flap.
Skin of dorsum finely granular, bearing
some enlarged warts (more distinct in para-
type); a series in the area where dorsolateral
folds might occur, a pair in scapular region,
a series forming a fold high on flanks; skin
of flanks coarsely granular with few enlarged
warts; throat smooth; ventral discoidal folds
prominent; no anal sheath. In USNM
165753, several strigiid parasites have en-
cysted beneath the skin just above the anus
and three more below and to the right. The
resulting structures (not on the holotype)
superficially resemble specialized anal warts.
A single strigiid cyst occurs on the underside
of the left shank near the heel, and another
on the left heel.
Skin of arm smooth except for some ru-
gosity on outer surface of forearm; palmar
tubercle distinctly bifid (Fig. 2C), larger than
thenar; thenar tubercle oval, distinctly ele-
vated; supernumerary palmar tubercles
present, subconical; subarticular tubercles
raised, round to oval, basal on III subcon-
ical, others non-conical; fringes along edges
of fingers, least developed along outside of
IV, palm, and I; fingers very long and slen-
der (Fig. 2C); toe tip on I scarcely expanded,
on II expanded but smaller than tympanum,
on III and IV expanded, larger than tym-
panum, weakly emarginate; circumferential
grooves present on all fingers (least evident
on thumb).
Upper surface of thigh smooth (lower and
posterior proximal surfaces areolate); outer
surface of shanks feebly granular; one or two
minute outer tarsal tubercles; inner meta-
tarsal tubercle 2.5 times as long as wide;
supernumerary plantar tubercles low, at
bases of toes II-IV; subarticular tubercles
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
moderate-sized, mostly subconical, longer
than wide; toe tips truncate, smaller than
those of outer fingers, weakly emarginate;
heels of flexed hind legs broadly overlap ('4
length of leg) when limbs held at right angles
to sagittal plane; heel of adpressed hind leg
reaching well beyond snout; shank 69.3-—
70.1% SVL.
The paratype was dissected to record the
condition of the m. adductor mandibulae
(“‘S’’). A moderate-sized frontoparietal fon-
tanelle, slightly larger than that illustrated
by Lynch (1971:147) for E. palmeri (=E.
nyctophylax), is found in adult males of E.
mercedesae.
Coloration in preservative. —Brown above
with cream blotches in complex lichenose
pattern; light areas include a | -shaped mark
on snout (bordered posteriorally by dark in-
terocular bar that forms anterior edge of
brown interocular triangle, irregular in
shape), two irregular paired blotches on an-
terior half of trunk, more diffuse cream
blotches posteriorally (Fig. 3); canthal stripe
dark brown; two or three dark brown sub-
ocular spots; dark brown supratympanic
fold, separated from eye; tympanum dark
brown, annulus cream except dorsally where
dark color of tympanum is continuous with
that of supratympanic fold; pair of dark
brown spots on each side in scapular region;
lips incompletely barred (three bars con-
sisting of dark half-moons); flanks pale
brown, reticulated with cream; limbs cream
with brown bars, bars about equal to inter-
spaces, transverse on shank; lower lip with
two or three irregular shaped, light brown
marks; center of throat with cream,
¢-shaped mark edged in brown, posterior
sides longer than anterior ones (Fig. 3);
throat darker than belly, belly cream with
brown reticulation; undersides of limbs
cream with brown spots; posterior surfaces
of thighs dark brown with many irregular,
small cream spots and lines; anal triangle
diffuse.
Coloration in life. —Medium brown dor-
VOLUME 100, NUMBER 2
Fig. 3. Eleutherodactylus mercedesae, holotype, USNM 257848.
sally with irregular, light green splotches
(spots look like moss) and few black marks;
black eyestripe (canthal plus supratympanic
stripe) and lip marks; limbs iridescent cop-
pery with green marks; thighs and sides of
body speckled cream, green, and brownish
black; ventrally pearly yellow, mottled
brown; iris gold (from field notes of M. S.
Foster).
Measurements of holotype (and paratype)
inmm.—SVL 40.1 (49.5), shank 28.1 (34.3),
HW 15.9 (18.0), head length 13.5 (17.1),
chord of head length 16.5 (19.7), upper eye-
lid width 3.9 (4.8), IOD 3.3 (4.0), tympa-
num 1.9 (2.3), eye 5.2 (6.2), E-N 4.7 (5.8).
Etymology. —This strikingly beautiful frog
is named for Mercedes S. Foster whose her-
petological collecting efforts in Latin Amer-
ica have added significantly to our knowl-
edge of the fauna of that region.
Remarks. —The type locality is in forest
adjacent to the road to San Onofre. The
holotype was found active on the forest floor
during the day.
The throat pattern in E. mercedesae is
somewhat similar to that in E. rhabdolae-
mus (see below and Fig. 4). However, we
do not believe that this feature is a synap-
omorphy. Several species of Eleutherodac-
tylus in the Andes of Colombia and Ecuador
have similar throat markings.
Eleutherodactylus rhabdolaemus Duellman
Eleutherodactylus rhabdolaemus Duell-
man, 1978a:65 (holotype, LSUMZ 26150,
Huanhuachayocc, on trail from Tambo
to Valle del Apurimac, Departamento
Ayacucho, Peru, 1650 m).
Eleutherodactylus pharangobates Duell-
man, 1978b:426 (Holotype, KU 173236,
Buenos Aires, Departamento Cuzco, Peru,
2400 m). NEW SYNONYM Y.
Although Duellman (1978a) compared
rhabdolaemus with pharangobates (as “E.
sp. [Buenos Aires]”’), he (1978b) did not
compare pharangobates with rhabdolaemus
(but did compare it with E. cosnipatae, E.
danae, and E. granulosus, three other species
from the vicinity of the type locality). The
two species are purported to differ in several
respects (Table 1).
One of us (JDL) has examined all known
specimens of the two taxa. The following
344
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Table 1.—Purported differences between two nominal species of Eleutherodactylus (from Duellman 1978a,
b). Cited differences marked with asterisk.
rhabdolaemus
Scapular tubercles Present
Snout shape
Vomerine odontophores
Digital pads
Heel, tarsal tubercles
Throat coloration
Labial stripe
Rounded
Small
Absent
point-by-point analyses are based on direct
comparisons of the type series (KU 173236-
54) of E. pharangobates with the KU para-
types of rhabdolaemus (138877, 175082-
84), the USNM Bolivian series, and notes
on the holotype (LSUMZ 26150). The skin
of the dorsum is finely shagreened in all
specimens of each species; all specimens we
examined have a pair of scapular warts. Al-
though not mentioned in the description of
E. pharangobates, scapular warts are ob-
vious in the photograph (Duellman 1978b,
Fig. 2e) of the paratype (KU 173237). These
warts (indistinct in some specimens), often
bordered with darker pigment to form scap-
ular marks, are more prominent in paler
individuals because the dark scapular marks
show up better against the ground color.
Duellman (1978a, b) described the snout
shapes in the diagnoses of the two species
as distinct but direct comparison belies these
differences (which were not cited in the
comparison of E. rhabdolaemus to phar-
angobates by Duellman [1978a] as distin-
guishing characteristics). Both species have
long snouts with sharp canthi rostrali and
subelliptical to somewhat pointed snouts (in
contrast to all other forms except E. scitulus
in the cloud forests of adjacent Bolivia and
Peru). Duellman (1978b) reported that male
E. pharangobates lack vomerine odonto-
phores. In both sexes of both nominate
species, including type material, low vo-
merine odontophores are evident. We find
no difference in position, size, shape, or tooth
number. Duellman (1978a) distinguished
Narrowly rounded
Present, both sexes
Longitudinal stripes
pharangobates
Not mentioned
Acuminate
Absent in males*
Truncate*
Absent*
Brown with cream flecks*
Present*
pharangobates from rhabdolaemus in hav-
ing truncate digital discs. Inspection of spec-
imens under magnification fails to confirm
a shape difference. Casual inspection (with-
out magnification or direct comparison) does
give the impression of truncate finger tips
in the type series of E. pharangobates. How-
ever, in those specimens having had the fin-
gers laid out in a fixing pan, one gets the
impression of truncateness whereas in those
preserved by Richard Thomas (collector of
most of Duellman’s [1978a] specimens of
rhabdolaemus), the hands were not laid flat
in a fixing tray and the tips appear less trun-
cate (more rounded). We consider the tips
of the outer fingers of all specimens to be
weakly truncate owing to slight emargina-
tion of the ungual flap. The two species were
reported (Duellman 1978a) to differ in tu-
berculation of the heel and tarsus. Both have
a low inner tarsal fold which usually has a
tubercle on it. Both have small, nonconical
tubercles on the heels and each has smaller
tubercles along the outer edge of the tarsus.
Duellman (1978a) contrasted rhabdolae-
mus and pharangobates in two aspects of
coloration: throat pattern and labial stripe.
The type series of E. pharangobates in-
cludes a single female and 12 males. The
males have semi-inflated vocal sacs and the
throat pattern is obscure. In the female, the
throat pattern is identical to that of para-
types of E. rhabdolaemus (Fig. 4) except
that it is paler. The pattern of a dark central
streak with narrower lateral streaks is evi-
dent in all specimens from the type series
VOLUME 100, NUMBER 2
-¥
a
Fig. 4. Venters of Eleutherodactylus rhabdolaemus. Intensity of throat patterns declines to right. Paratype
of E. rhabdolaemus, KU 175083; female paratype of E. pharangobates, KU 173246; and male paratype of E.
pharangobates, KU 173250.
examined. The same general throat patterns
are repeated in several of the Bolivian spec-
imens except that a streaked throat is ob-
vious in several males as well.
Darker specimens have dark faces. The
labial stripe recorded by Duellman (1978a,
b) for pharangobates is produced by a paler
face edged by the dark canthal stripe and
the partial labial bars; the clear area evident
between these dark markings was termed a
labial stripe (see Duellman 1978b, fig. 2e).
The pattern evident in dark specimens is
identical to that seen in pale specimens. The
difference in intensity is consistent in the
type series of pharangobates but is bridged
within the series of paratypes of rhabdolae-
mus. This difference appears to be the result
of metachrosis. This interpretation is sup-
ported by Foster’s observation (unpub-
lished field notes) that a specimen of E.
rhabdolaemus (USNM 257854) varied from
“very dark to light brown—changes color.”
In the absence of sustained differences,
we combine the two taxa. Eleutherodactylus
pharangobates was described on 29 August
1978, slightly more than two months after
the description of EF. rhabdolaemus ap-
peared (27 June 1978). Duellman (1978a,
b) reported specimens from the Peruvian
Departamentos Ayacucho, Cuzco, and
Huanuco from localities between 1020 and
2650 m. He also reported one specimen
(USNM 146587, as pharangobates) from
Departamento Cochabamba, Bolivia. We
have seen the following material of E. rhab-
dolaemus from Bolivia (all Provincia Cha-
pare, Departamento Cochabamba): Limbo,
on mule trail to ““Mino Porro,” 1950 m,
USNM 146592-93; 10 km N Limbo, 1800
m, USNM 146587-91; Miguelito, ca. 99 km
from Cochambamba on Cochabamba to
Villa Tunari road, 1770 m, USNM 257850-
51; Paracti, 83 km from Cochabamba on
Cochabamba to Villa Tunari road, 2040 m,
USNM 257856-—57; road to San Onofre, ca.
3.3 km N of Cochabamba to Villa Tunari
road at point 97 km from Cochabamba,
1695 m, USNM 257855.
Acknowledgments
We would like to thank the curators of
the following museum collections for loan
of pertinent material: KU—Museum of
Natural History, University of Kansas;
LSUMZ-— Louisiana State University, Mu-
seum of Zoology; USNM—National Mu-
seum of Natural History. In addition, JDL
346
thanks W. E. Duellman for providing space.
R. I. Crombie and R. P. Reynolds read a
draft of the manuscript and made several
worthwhile comments.
Literature Cited
Boettger, O. 1891. Reptilien und Batrachier aus Bo-
livia.— Zoologischer Anzeiger 14:343-347.
Cannatella, D.C. 1980. Two new species of Centro-
lenella from Bolivia (Anura: Centrolenidae).—
Proceedings of the Biological Society of Wash-
ington 93:714-724.
Cei, J. M. 1980. Amphibians of Argentina.— Moni-
tore Zoologico Italiano (N.S.) Monografia
2:xi1 + 1-609.
Duellman, W. E. 1978a. Two new species of Eleu-
therodactylus (Anura: Leptodactylidae) from the
Peruvian Andes.—Transactions of the Kansas
Academy of Science 81:65-71.
1978b. New species of leptodactylid frogs of
the genus Eleutherodactylus from the Cosnipata
Valley, Peru.— Proceedings of the Biological So-
ciety of Washington 91:418-430.
, and T. H. Fritts. 1972. A taxonomic review
of the southern Andean marsupial frogs (Hyli-
dae: Gastrotheca).—Occasional Papers of the
Museum of Natural History, University of Kan-
sas 9:1-37.
Heyer, W. R. 1978. Systematics of the fuscus group
of the frog genus Leptodactylus (Amphibia, Lep-
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
todactylidae).— Natural History Museum, Los
Angeles County, Science Bulletin 29:1-85.
Lynch, J. D. 1971. Evolutionary relationships, os-
teology, and zoogeography of leptodactyloid
frogs.— University of Kansas, Museum of Nat-
ural History. Miscellaneous Publication 57:1-
238.
. 1976. The species groups of South American
frogs of the genus E/eutherodactylus.—Occa-
sional Papers of the Museum of Natural History,
University of Kansas 61:1-24.
1980. A taxonomic and distributional syn-
opsis of the Amazonian frogs of the genus
Eleutherodactylus.— American Museum Novi-
tates 2696:1-24.
. 1986. The definition of the Middle American
clade of Eleutherodactylus based on jaw mus-
culature (Amphibia: Leptodactylidae).— Her-
petologica 42:248-258.
,andC. W. Myers. 1983. Frogs of the fitzingeri
group of Eleutherodactylus in eastern Panama
and Chocoan South America (Leptodactyli-
dae).—Bulletin of the American Museum of
Natural History 175:481-572.
(JDL) School of Biological Sciences, Uni-
versity of Nebraska, Lincoln, Nebraska
68588; (RWM) U.S. Fish & Wildlife Ser-
vice, National Museum of Natural History,
Washington, D.C. 20560.
PROC. BIOL. SOC. WASH.
100(2), 1987, pp. 347-352
ANOPSILANA LINGUA, A NEW FRESHWATER
TROGLOBITIC ISOPOD FROM THE PALAU ISLANDS
(FLABELLIFERA: CIROLANIDAE)
Thomas E. Bowman and Thomas M. Iliffe
Abstract. — Anopsilana lingua, a blind, unpigmented cirolanid isopod, is de-
scribed from a natural well on Peleliu Island. It is distinguished from the nine
known species of Anopsilana by the shape of the telson, which is very broadly
rounded, in contrast to the triangular, pointed or angular telsons of the other
species. Anopsilana lingua is the first troglobitic cirolanid reported from a
Pacific Ocean locality.
Explorations of caves on Atlantic Ocean
islands in recent years have revealed the
presence of rich and diverse faunas of aquat-
ic troglobites (Sket and Iliffe 1980; Iliffe et
al. 1983, 1984). A summary of the species
in these faunas and those of other troglobitic
aquatic faunas of the world is given in the
recent volume edited by Botosaneanu
(1986). In contrast, the number of known
troglobites from caves of Pacific Ocean is-
lands is small. Collecting effort in Pacific
caves has been much less than in Atlantic
caves, and to remedy this situation, the sec-
ond author led an expedition to explore and
collect in Pacific island caves in 1985. One
of the discoveries of this expedition was the
isopod described below.
Anopsilana lingua, new species
Figs. 1-3
Material. —Palau, Peleliu Island, Airport
Well, 26 Feb 1985, leg. Thomas M. Iliffe,
Jeff Bozanic, and Dennis Williams, holo-
type (USNM 232000) and 16 paratypes
(USNM 232001); 2 Apr 1985, leg. Dennis
Williams and Jeff Bozanic, 25 paratypes
(USNM 232002).
Etymology.—From the Latin “lingua”
(tongue), referring to the shape of the pleo-
telson.
Description. — Blind, unpigmented.
Length of largest specimen 11 mm. Body
length/width 2.48—3.25, tending to be great-
er in longer specimens (Fig. 1). Head sub-
triangular, about 2 roe a yi} aS
Paps et NET EE
, re gm Ty , a
;
a4 dade
aa
\ Aligeet
ee 3 gti,
Figs. 34, 35.
423
-
=.
.
i>? oy
. PD :
34, (top): Female (20.7 mm) Renocila kohnoi on 73 mm SL namerayakko, Centropyge vrolicki
(Bleeker), laboratory photograph an Amitori; 35, (bottom): Male (9.6 mm) on side opposite female in Fig. 34.
on the posterior tips of the dorsal and anal
fins in some hosts. On one host, three males
were atranged in a tight triangle about mid-
body along the center of the lateral side,
anterior of the female. Two females were
attached slightly posterior to the body mid-
line, and one male was attached posterior
of the caudal peduncle. Number of isopods
per host varied from two to seven and av-
eraged 2.9.
Description.—Body ovate. Antennae 1
7-merous, antennae 2 7—8-merous. Anten-
nae | distinctly longer and wider than an-
tennae 2, distal 5 articles with tufts of setae
on distal ventral corners. Anterior margin
of head ventrally flexed, produced into
broadened lobe between bases of antennae
|. First segment of mandibular palp slightly
expanded, 3 simple setae on tip of distal
article. Incisor process of mandible with
pointed tip. Maxilla 1 with | slender and 3
more robust, recurved spines. Distal lobes
of maxilla 2 with | spine on inner lobe and
2 spines on outer lobe, spines short and al-
most obscured by long distal portions of
semilunar pectinate scales which are longer
on this appendage than on others. Distal
segment of maxillipedal palp with 2 re-
curved spines. Posteroventral angles of per-
eonite | moderately produced, those of 4—
7 produced, and that of 7 extending to lat-
eral portion of pleonite 5 or to posterior
WASHINGTON
VOLUME 100, NUMBER 3
extent of basis of uropods. Pleotelson semi-
circular, 1.43—1.75 (1.56) times wider than
long with median ridge rising from posterior
to anterior. Exopod of uropods only %4 to
longer than endopod. Exopod and not en-
dopod reaching beyond posterior border of
pleotelson but in 2 cases neither ramus at-
taining posterior border of pleotelson. Pe-
reopods 1—4 with swelling of dactyls. Pereo-
pods 1-3 with lobe at posterodistal corner
of basis. Dactyls subequal in length, none
lengthened. Pereopods 1-6 subequal in
length or increasing in length slightly from
1-6. Pereopod 7 abruptly longer than pereo-
pod 6. Color uniform slate gray.
Juvenile (attached to host) (n = 5): Total
length 3.7—4.0 (3.9), maximum width 1.0-
1.2 (1.1). Possessing 6 pairs of pereopods,
no appendix masculina and no penes lobes.
Three juveniles possessing setae on pleo-
telson and terminal portions of uropods;
pereopods 1-3 with comb-like teeth on in-
ner margin of dactyls and row of large hooks
on inner margin of propodus; large chro-
matophores on snout along line between an-
terior extreme of eyes; large darkly pig-
mented lateral stripe from eyes along pereon,
becoming more diffuse on sides of pleon,
reappearing as dark stripe on outer edge of
exopod of uropods; pigment spots outlining
edges of pleopods | and 2, spots on distal
outer margins of pleopod 3 and sometimes
4; pigment spots on midventral surface of
pereopods; pigment spots dispersed along
posterior borders and middle of pereonites
and pleonites, forming elongated oval of
pigment spots along mid-line of isopod.
These 3 juveniles similar in size and mor-
_
425
phology to juveniles found in brood pouch-
es of 2 female R. kohnoi. Remaining 2 at-
tached juveniles (3.7 and 4.0 long, 1.1 and
1.2 wide) resembling first 3 except for pig-
ment spots uniformly dispersed over dorsal
surface; comb-like teeth on inner margins
of dactyls and hooks present on inner mar-
gin of propodus of pleopods 1-3 reduced in
length. This second morphological type may
represent first molt after settling on host.
These 5 juveniles all attached to fishes also
parasitized by female R. kohnoi with repro-
ductive products other than juveniles.
Juvenile-male transitional (n = 2): Total
length 5.6 and 7.8, width 2.2 and 2.0. Both
isopods possessing 7 pairs of pereopods, ap-
pendix masculina as long as or longer than
endopod or pleopod 2, and no penes lobes
apparent. Shorter (and wider) isopod col-
ored uniformly slate gray (male coloration)
and possessing no comb-like teeth on inner
margins of dactyls or hooks on inner mar-
gins of propodus of any pereopods. Longer
isopod similar in color and pereopod mor-
phology to group of 2 attached juveniles
described above.
Male (n = 11): Total length 5.6—-11.3 (8.6),
width 2.0-6.0 (4.1), appendix masculina
linear with unmodified tip, slightly shorter
to slightly longer than endopod of pleopod
2. Penes lobes small, bilobed, conical pro-
cess. Possessing no remnants of comb-like
teeth on inner margins of dactyls or hooks
on inner margins of propodus. Color uni-
formly slate gray.
Male-female transitionals (n = 3): Total
length 9.2—-13.0 (10.5), width 4.4—7.0 (5.4)
with appendix masculina shorter than inner
Figs. 36-59. Renocila kohnoi, new species (36, 37, 39 = 15 mm female holotype with oostegites; 38 = 11.1
mm male allotype; 40-54 = 18.6 mm female paratype lacking oostegites; 55-59 = 11.3 mm male paratype; all
from Centropyge heraldi) scale bars in mm; scale of 36-38 equal; 40-41, and 51-59 equal; 43, 45, 48, and 50
equal; 42, 44, 46, 47, 49 enlargements from associated mouthpart figures). 36, Dorsal view; 37, Lateral view;
38, Dorsal view; 39, Head, ventral view; 40, Antennae 1; 41, Antennae 2; 42, Distal lobes of maxilla 2 with
semilunar pectinate scales; 43, Maxilla 2; 44, Apex of maxilla 1; 45, Maxilla 1; 46, Apex of distal segment of
mandibular palp; 47, Incisor process of mandible; 48, Mandible and palp; 49, Apex of maxillipedal palp; 50,
Maxilliped; 51, Pereopod 1; 52, Pereopod 4; 53, Pereopod 7; 54, Uropod; 55, Uropod; 56, Pereopod 6; 57,
Pereopod 7; 58, Pereopod 1; 59, Pleopod 2.
426
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Table 2.— Description of development in the brood pouch of 4 females of Renocila kohnoi (measurements in
mm).
Number Description
111 Subspherical embryos
219 Hunched embryos with internal segmentation, broad ce-
phalic end and darkly pigmented eyes
109,109 Juveniles with racing stripes (as described in text)
blade of pleopod 2 and penes lobes not ap-
parent. Color uniformly slate gray.
Female (n = 9): Total length 15.6—21.2
(18.8), width 9.7-13.2 (11.6), no remnants
of appendix masculina or penes lobes pres-
ent. All but 2 females (17.1 and 18.6 long
and 11.2 and 11.3 wide), possessing oos-
tegites.
Brood pouch reproduction (n =
each): See Table 2.
Etymology. —The specific name 1s in hon-
or of Mr. Hiroyoshi Kohno, who told us
about this isopod, and helped us to collect
specimens.
Japanese standard common name. —
Kohno-yadori-mushi (shin-sho) = Kohno’s
isopod.
Remarks.— Renocila kohnoi resembles R.
periophthalmi in general body shape. It dif-
fers from all known species of the genus by
having swelling of the dactyl of pereopod 4
in addition to those on pereopods 1-3.
The 30 Renocila kohnoi were collected
from eight Centropyge heraldi, 55—80 (68.1)
in standard length, and two C. vrolicki, 69
and 73 in standard length. Parasitized C.
heraldi were observed as deep as 27.4 m
(collected at 24.3 m at Iriomote Island) and
collected as shallow as 9.1 m at Ishigaki
Island. Parasitized C. vrolicki were collected
at 7.6 m at Iriomote Island and 6.1 m at
Ishigaki Island and observed as shallow as
4.6 m at Ishigaki Island. Of the nine species
of Centropyge reported from the Yaeyama
Islands (Masuda et al. 1984), we observed
seven. Centropyge bispinosus (Gunther), C.
tibicen (Cuvier), C. bicolor (Bloch), C. nox
(Bleeker), and C. ferrugatus Randall and
Burgess were not observed to be parasitized
20 of
Length (mean) Width (mean)
1.32-1.56 (1.44)
1.80-1.98 (1.86)
1.20-1.26 (1.22)
0.95-1.10 (1.04)
3.95-4.11 (4.03) 1.09-1.15 (1.12)
by external isopods. Twenty—thirty-three
percent (22%) (five observations for a total
of 98 fish) of C. heraldi and 20—40% (33.3%)
(two observations for a total of 15 fish) of
C. vrolicki were observed parasitized by R.
kohnoi. Only one C. heraldi was observed
with a single (male?) isopod (not collected);
all other infections observed were by more
than one isopod.
Renocila kohnoi, a dark-colored isopod,
was very visible in the water attached to the
yellow C. heraldi (Figs. 32-33). A similar
situation exists with Anilocra holacanthi
Williams and Williams, where the dark-col-
ored isopod attaches under the eye of Ho-
lacanthus tricolor (Bloch) whose face is yel-
low (Williams and Williams 1981). Renocila
kohnoi is almost invisible on C. vrolicki
which is light brown on the anterior body
and dark brown on the posterior one-third
of the body and caudal peduncle where the
isopod attaches (Figs. 34-35). The corre-
spondence of the dark isopod to the limited
dark areas of this host may be more than
coincidence. Possibly R. kohnoi was first a
parasite of C. vrolicki and later parasitized
both hosts.
Four R. kohnoi were partially molted with
the posterior three pereonites and the pleon
in a softened condition, the head and pe-
reonites 1—4 not molted.
Three of the five juveniles of R. kohnoi
attached to the hosts were identical in size,
shape, and coloration to the juveniles of the
brood pouch (Table 2). These juveniles ap-
parently attached to the host soon after leav-
ing the brood pouch, and, therefore, may
not have any intermediate planktonic pe-
riod or intermediate host.
VOLUME 100, NUMBER 3
Renocila bollandi, new species
Figs. 60-69
Type specimen. —Holotype (female),
USNM 228659.
Type locality. —“‘Horseshoe Cliffs,” 1 km
WNW Onna Village, Okinawa, Japan,
26°30.0'N, 127°50.9’E (2 Jan 1985, 38.1 m,
0800 hrs, temperature at depth 24.8°C).
Type host. —Onikasago, Scorpaenopsis
cirrhosa (Thunberg), Scorpaeniformes:
Scorpaenidae) (Fig. 60).
Additional host and locality. —Madara-
fusakasago, Scorpaena bynoensis Richard-
son (Fig. 61), Onna Point, Okinawa, Japan
(20 Sep 1985, 12.2 m, 2200 hrs) (2 under-
water photographs).
Site of infection. —Female attaches on the
427
4
Figs. 60,61. 60, (top): Female 13.9 mm holotype (USNM 228659) of Renocila bollandi on the 44 mm type-
host specimen of onikasago, Scorpaenopsis cirrhosa (Thunberg), aquarium photograph by Dr. Robert F. Bolland;
61, (bottom): Female (uncollected) of Renocila bollandi on madara-fusakasago, Scorpaena bynoensis Richardson,
from Onna Point, Okinawa, underwater photograph by Mr. Gary Hagland.
anterior, lateral portion of the host, poste-
rior of and dorsal of the eye (Figs. 60-61).
Description. —Body oval. Head im-
mersed in pereonite 1. Eyes small. Antennae
1 6-merous, not reaching posterior border
of head; antennae 2 3-merous, very short,
reaching to third or fourth article of anten-
nae |. Anterior margin of head rounded, not
ventrally flexed. Coxae of pereonites pro-
duced laterally, those of pereonites 2-3
reaching posterior extent of segments. Pos-
terolateral angles of pereonite 3 moderately
produced, those of pereonites 4—7 produced
posteriorly and laterally, that of pereonite
7 extending posteriorly to level of lateral
portion of pleonite 4—5. Pereonites and ple-
onites with projection at middle of posterior
428
border of segment, that of pleonite 5 larger
than others, raised dorsally and extending
over pleotelson. Medial ridge rises from an-
terior pereon to posterior pleon. In dorsal
view, pleotelson deeply incised anteriorly
by medial posterior projection of pleonite
5. Pleotelson 1.36 times wider than long,
rounded posteriorly, slightly pointed at me-
dial posterior border. Uropods extending to
posterior border of pleotelson, exopod lon-
ger than endopod. Pereopods 1-3 without
swelling of dactyls or lobate posterodistal
corners of propodus. Dactyls and pereopods
increasing in length from 1 to 3 or 4 (which
are sub-equal) then decreasing gradually in
length to pereopod 7. Pereopod | abruptly
shorter than pereopod 2. Color, yellowish-
brown with posterior borders of all seg-
ments and anterior border of head and pe-
reonite | outlined by darkly pigmented line.
Female (n = 1): Total length 13.9, max-
imum width 8.8. No remnants of appendix
masculina or penes lobes, with oostegites.
Etymology. — The specific name is in hon-
or of Dr. Robert F. Bolland who collected,
photographed, and provided additional in-
formation about this isopod.
Japanese standard common name. —Ze-
bra-yadori-mushi (shin-sho) = zebra iso-
pod.
Remarks.— Antennae, mouthparts, and
uropods were not detached from the holo-
type to avoid damaging the single specimen.
Renocila bollandi differs from the diag-
nosis of the genus (Bowman and Mariscal
1968) by having an immersed head, pro-
duced posteroventral angle of pereonite 4,
medial posterior projections of pereonites
and pleonites, and reduced antennae 2.
The specimen of Scorpaenopsis cirrhosa
was 44 in standard length and the observed
and photographed Scorpaena bynoensis ap-
proximately 49 in standard length. Both
hosts are young specimens. This isopod has
been observed in depths from 12.2 to
38.1 m.
This isopod is known from a 500 m sec-
tion of the west coast of central Okinawa.
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
We did not see this isopod during approx-
imately 80 hrs of day and night scuba ob-
servations around Sesoko Island, 14—20 km
north of Onna Point (May 1985—Mar 1986).
We only observed one other common
species of scorpaenid, satsumakasago, Scor-
paenopsis diabolus (Cuvier), in the Ryukyu
Islands, but it was never observed parsitized
by this isopod.
Discussion
Cymothoids are traditionally associated
in intimate, stable, long-term, female-male
pairs. This process seems to be stabilized
by neuro—endocrine control of the male by
the female (as summarized by Brusca 1981).
The female (or male) somehow excludes the
attachment and growth of additional con-
specific isopods. The constancy of this tra-
ditional pattern among almost all of the cy-
mothoid isopods is probably based on the
parasite-carrying capacity of hosts. No more
than one female isopod and one male can
be accommodated without threatening the
survival of the host. The female isopod,
through a series of non-growth reproductive
cycles interspersed with vegetative growth
periods (Williams and Williams 1982),
probably grows to an optimum size for the
size of the host. This is supported by the
direct relationship between host length and
female isopod length found in many cy-
mothoids (Trilles 1964; Williams and Wil-
liams 1982). The obvious advantage of
maximum female isopod size (as large as
can be supported by the host) is that larger
isopods (of the same species) produce more
individual offspring. The number of eggs
varies directly with body length within a
species (Brusca 1981).
We have observed two major modifica-
tions of this traditional pattern. The first
occurs in Carribean Anilocra. The juvenile
serves aS a mobile male (micromale) and
the host-carrying capacity, normally occu-
pied by the male, can be taken by a second
female. Thus the reproductive potential is
VOLUME 100, NUMBER 3
429
Figs. 62-69. Renocila bollandi, new species, 13.9 mm female holotype. (Scale bars in mm; scale of 62-63
equal, scale of 64—69 equal.) 62, Dorsal view; 63, Lateral view; 64, Uropods; 65, Pereopod 1; 66, Pereopod 2;
67, Pereopod 4; 68, Pereopod 7; 69, Head, ventral view.
Figs. 70-73. Renocila yamazatoi, new species, 15.3 mm female paratype. (Scale bars in mm; scale of 70-
77 equal.) 70, Pleopod 2; 71, Pleopod 3; 72, Pleopod 4; 73, Pleopod 5.
Figs. 74-77. Renocila kohnoi, new species, 18.6 mm female paratype. 74, Pleopod 2; 75, Pleopod 3; 76,
Pleopod 4; 77, Pleopod 5.
essentially doubled (Williams 1984). The
second modification involves a physical
separation of the female-male pair on the
host. This duplex arrangement was de-
scribed for Mothocya bohlkeorum, where a
single male and female occupy different gill
chambers on a single host (Williams and
Williams 1982). Lironeca circularis Pillai
and Lironeca sp. also follow this pattern in
33 hosts (Amblygaster sirm (Walbaum)) we
examined from Thailand and the Ryukyu
Islands.
430
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Table 3.—A developmental listing of 31 Renocila kohnoi arranged by hosts, and ranked in female-male pattern
categories.
1 2
Developing 3
toward
duplex female-
Status male
Development stage
Female l 1
Male-female transitional l
Male l
Juvenile-male transitional l l
Juvenile l
All species of the genus Renocila for which
the attachment positions are known, except
R. kohnoi, display the traditional female-
male arrangement on the host. In contrast,
R. kohnoi is strongly duplex in its female-
male pattern by having the largest two iso-
pods in the same position on opposite sides
of the host. In addition, a number of smaller
isopods also attach to the host and compli-
cate this duplex pattern in five of the ten
hosts examined. This combination of du-
plex arrangement and additional small 1so-
pods may indicate that this isopod-host re-
lationship is a fairly recent phenomenon. By
examining the specific relationships of these
isopods on each host, we may gain some
understanding of how the three basic ar-
rangements came about.
Table 3 ranks the isopods occurring on
each host by development stage, and also
divides the assortment found on each host
in terms of the three female-male patterns
(Traditional female-male, Duplex female-
male, and Double female-micromale). Our
interpretation is that isopods on hosts 1 and
2 (Table 3) are developing toward a stable
duplex female-male arrangement. The male-
female transitional in host 1 will become a
female and the juvenile-male transitionals
in hosts | and 2 will become males. The
isopods on hosts 3—5 are stable duplex fe-
male-male pairs. The attachment of the ju-
veniles in host 5 is interesting because they
could develop into males as in hosts 6-8,
Host number
9 10
4 5 6 7 8
Developing
Stable duplex Developing toward toward
female-male traditional female-male female pair
but our field and aquarium experiments
(Williams 1984) have demonstrated that
mere attachment of juveniles does not nec-
essarily guarantee survival and growth.
Hosts 6-8 have additional males. Two of
these are attached in front of the female,
very similar to some traditional female-male
pairings (Renocila colini Williams and Wil-
liams, for example). This suggests a method
by which a traditional female-male arrange-
ment could be developed from a duplex ar-
rangement, but, in the case of R. kohnoi,
some mechanism (for example: limited
space available for attachment of adult iso-
pods to the caudal peduncle, or more swim-
ming stability of hosts with large isopods
on both sides) seems to be maintaining the
duplex arrangement. Hosts 9-10 (Table 3)
have the unusual relationship of female and
male-female transitional. This suggests lack
of female control of the male over the dis-
tance of opposite sides of the host. If the
male-female transitional also became a fe-
male on this host and if a functional male
was small and mobile (as in Caribbean Ani-
locra), then a double female-micromale ar-
rangement could develop.
Thus, the arrangement of individuals of
R. kohnoi on a host displays the duplex ar-
rangement as well as possible methods of
changing to either a traditional or a double
female arrangement. It is possible that the
duplex arrangement is a precursor to the
more frequently observed female-male re-
VOLUME 100, NUMBER 3
lationships, particularly when an isopod is
parasitizing a new host.
Williams et al. (1983) reported the only
known case of more than one species of
Renocila (R. bowmani Williams and Wil-
liams and R. colini) occurring at the same
geographic location (Isla Saona, Dominican
Republic). We observed a specimen of R.
kohnoi on a Centropyge heraldi among
Chromis ovatiformis parasitized by R. ya-
mazatoi south of Saba Saki in Amitori Bay.
The extremely limited geographic ranges,
high levels of infection and strong host spec-
ificity suggested for the species of Renocila
by Williams and Williams (1980) are not
contradicted by these three new species from
the Ryukyu Islands.
Key to Renocila Species
1. Dactyls of pereopods 1-3 with
swelling on outer margin ....... 2
— Dactyls of pereopods 1-3 without
SG ae 5
2. Antennae | shorter than antennae
2 Oe R. dubia (Nierstrasz)
— Antennae | longer than antennae
MP 8a ess. CAN ead 3
3. Pleotelson longer than wide
a R. periophthalma Stebbing
— Pleotelson wider than long ..... 4
4. Dactyl of pereopod 4 with swelling
on outer margin ....R. kohnoi, n. sp.
— Dactyl of pereopod 4 without
swelling
.... R. indica Schioedte and Meinert
5. Antennae | and 2 subequal ..... 6
— Antennae | longer than 2
6. Posteroventral angle of pereonite
7 produced laterally, not covering
lateral margins of anterior pleo-
nites in dorsal view
— Posteroventral angle of pereonite
7 not produced laterally, covering
lateral margins of anterior pleo-
nites in dorsal view
7. Posteroventral angle of pereonite
5 produced, antennae 2 extending
GC 0 0 O40 0 Os OO Oo OF 0™O OO) 0-0) OO OG
431
beyond posterior border of head
.... R. colini Williams and Williams
— Posteroventral angle of pereonite
5 not produced, antennae 2 not ex-
tending to posterior border of head
....R. thresherorum Williams and
Williams
8. Brown in color, antennae 2 8-mer-
ous, antennae | slightly shorter
thane 2 erie he are ee aie R. waldneri
Williams and Williams
— Black in color, antennae 2 7-mer-
ous, antennae | slightly longer than
DETR te ea ee eee ae R. bowmani
Williams and Williams
9. Lobe of head between bases of an-
tennae 1, exopod of uropod more
than twice as long as endopod ... 10
— No lobe between bases of antennae
1, exopod of uropod only slightly
longer than endopod .......... 11
10. Posteroventral angle of pereonite
5 produced, dactyls of pereopods
2-6 lengthened . R. yamazatoi, n. sp.
— Posteroventral angle of pereonite
5 not produced, dactyls of pereo-
pods subequal ......... R. heterozota
Bowman and Mariscal
11. Pereonites and pleonites with me-
dial posterior projections, head
immersed R. bollandi, n. sp.
— Pereonites and pleonites lack me-
dial posterior projections, head not
HMIMEeTSeG! Pas R. ovata Miers
Acknowledgments
Mr. Hiroyuki Yokochi and Mr. Hiro-
yoshi Kohno, Iriomote Marine Research
Station, Tokai University, Iriomote Island;
Dr. Akinosuke Tomori, Mr. Nobuhiro
Ohshiro, and Mr. Kazuyuki Shimoike, Yae-
yama Branch Fisheries Laboratory, Oki-
nawa Prefectural Fisheries Laboratory, Ish-
igaki Island; Mr. Yonezo Ikeda, Iriomote
Island; Dr. Robert F. Bolland, University
of Maryland, Okinawa Island; Mr. Gary
Hagland, Kadena Air Force Base, Okinawa
432
Island helped collect fishes or provided ob-
servations or equipment and facilities. Dr.
Kiyoshi Yamazato, Sesoko Marine Science
Center, and the Ministry of Education, Sci-
ence and Culture supported our research for
one year (1985-1986). Additional support
was provided by the Department of Marine
Sciences, University of Puerto Rico, Ma-
yaguez, Puerto Rico. This is a contribution
of the Sesoko Marine Science Center.
Literature Cited
Bowman, T. E., and R. N. Mariscal. 1968. Renocila
heterozota, anew cymothoid isopod, with notes
on its host, the anemone fish, Amphiprion okal-
lopisos, in the Seychelles.—Crustaceana 14:97-
103.
Brusca, R. C. 1981. A monograph on the Isopoda
Cymothoidae (Crustacea) of the eastern Pacif-
ic.— Zoological Journal of the Linnean Society
73(2):117-199.
Masuda, H., K. Amaoka, C. Araga, T. Uyeno, and T.
Yoshino (Eds.). (1984) The fishes of the Jap-
anese Archipelago. Tokai University Press, 437
pp. ‘
Trilles, J. P. 1964. A propos d’un fait particulier
d’éthologie parasitaire chez les Isopodes Cy-
mothoidae: La relation de taille entre parasites
et poissons. Note préliminaire.— Vie et Milieu
15:365-369.
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Williams E. H., Jr., and L. B. Williams. 1980. Four
new species of Renocila (Isopoda: Cymothoi-
dae) the first reported from the new world.—
Proceedings of the Biological Society of Wash-
ington 93:573-592.
—., and . 1982. Mothocya bohlkeorum new
species (Isopoda: Cymothoidae) from West In-
dian cardinal-fishes (Apogonidae).— Journal of
Crustacean Biology 2:570--577.
, I. Clavijo, J. J. Kimmel, P. L. Colin, C. Diaz-
Carela, A. T. Bardales, R. A. Armstrong, L. B.
Williams, R.H. Boulon, and J.R. Garcia. 1983.
A checklist of marine plants and animals of the
south coast of the Dominican Republic.—Ca-
ribbean Journal of Science 19:39-53.
Williams, L. B. 1984. Geographic distribution and
early life history of Anilocra (Isopoda: Cymo-
thoidae) parasites of Caribbean coral reef fishes.
Auburn University, Ph.D. Dissertation, 312 pp.
,and E. H. Williams, Jr. 1981. Nine new species
of Anilocra (Crustacea: Isopoda: Cymothoidae)
external parasites of West Indian coral reef fish-
es.— Proceedings of the Biological Society of
Washington 94:1005-1047.
Foreign Visiting Researchers, Sesoko
Marine Science Center, University of the
Ryukyus, Motobu-cho, Okinawa 902-05
Japan. Permanent address: Department of
Marine Sciences, University of Puerto Rico,
Mayaguez, Puerto Rico 00708.
PROC. BIOL. SOC. WASH.
100(3), 1987, pp. 433-446
FALLICAMBARUS (CREASERINUS) BURRISI AND
F. (C. GORDONI, TWO NEW BURROWING
CRAWFISHES ASSOCIATED WITH PITCHER PLANT
BOGS IN MISSISSIPPI AND ALABAMA
(DECAPODA: CAMBARIDAE)
J. P. Fitzpatrick, Jr.
Abstract. —Two new burrowing crawfishes, closely related to Fallicambarus
(Creaserinus) byersi (Hobbs) are described from southern Mississippi and Al-
abama. They share with F. byersi and F. caesius Hobbs the characteristic, not
otherwise known in the genus, of having a row of long stiff setae along the
ventrolateral margin of the chela. They differ from them in having a more
slender central projection on the gonopod of the first form male and from each
other by the degree to which the apex of the mesial process of that member is
curved caudoproximally. Each species has a unique annulus ventralis, and other
morphological features serve to indicate the several species.
Several years ago, when the Mississippi
Academy of Science was initiating studies
to identify the state’s Rare and Endangered
species, I called attention to the unique re-
lationship between Fallicambarus (Crea-
serinus) byersi (Hobbs 1941) and pitcher
plant (Sarracenia spp.) bogs. My discovery
of a second, closely related, similarly en-
vironmentally restricted species elicited the
interest of the Mississippi Natural Heritage
Program. As a result, they have paid special
attention to the collection of burrowers, and
the results of some of their labors are re-
ported here.
Fallicambarus (Creaserinus) burrisi,
new species
Figs. 1, 2a—d
Diagnosis. — Antennal scale much re-
duced, lacking usual spiniform distal ter-
minus of lateral thickened part (Fig. le);
coxa of fourth pereiopod with large, cau-
domesially directed boss; telson undivided,
entire. Ventrolateral surface of propodus of
chela with row of long stiff setae near lateral
margin. First pleopod of first form male in-
clined caudally in distal third, apices of ter-
minal elements inclined at about 130° to
main axis of pleopod shaft. Annulus ven-
tralis of female large, about 1.3 times longer
than wide; with lateral and caudal margins
conspicuously raised; prominent inverted
U-shaped sinus arising on caudal margin,
well lateral to midline, passing cephalically
20% of length of annulus, and recurving and
disappearing beneath cephalic overhang of
caudal elevation on same side of midline as
origin; overhang of caudal part of annulus
completely obscuring postannular sclerite.
Holotype male, Form I.—Cephalothorax
subovate, compressed (Fig. 1b, h); eyes re-
duced. Abdomen much narrower than tho-
rax (6.2 and 9.6 mm); greatest width of car-
apace less than depth at caudodorsal margin
of cervical groove (9.6 and 10.8 mm). Are-
ola obliterated along most of length; length
39.6% of entire length of carapace (42.8%
of postorbital carapace length). Rostrum de-
pressed, with mildly convergent, slightly
thickened margins constricting near mid-
length to tip of indistinctly delimited acu-
men, tip slightly upturned, reaching base of
ultimate podomere of antennular peduncle;
434 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
j
#8,
Fig. 1. Fallicambarus (Creaserinus) burrisi, holotype: a, Mesial view of first pleopod; b, Dorsal view of
carapace; c, Lateral view of chela; d, Lateral view of first pleopod; e, Antennal scale; f, Ventral view of thoracic
VOLUME 100, NUMBER 3
dorsal surface deeply concave cephalically
and subplanar caudally. Subrostral ridges
poorly evident in dorsal aspect. Postorbital
ridges prominent, grooved laterally, and
terminating cephalically in weak tubercles.
Suborbital angle and branchiostegal spine
absent. Carapace sparsely punctate dorsally
with slightly granulate branchiostegites.
Abdomen shorter than carapace (17.3 and
21.7 mm); pleura quite short and broadly
rounded; cephalic lobe of pleuron of second
segment overlapping reduced pleuron of first
(Fig. 2e). Telson not clearly divided into
cephalic and caudal portions (Fig. 1j); dex-
tral margin provided with short, acute, fixed
spine posterior to midlength, spine absent
from corresponding position on sinistral
margin. Proximal podomere of uropod with
broadly rounded tubercle on medial lobe;
both rami rounded distally; lateral ramus
with very tiny fixed lateral spine and small,
acute, premarginal medial spine at end of
weak median ridge; mesial ramus with small,
fixed lateral spine and median ridge.
Cephalomedian lobe of epistome (Fig. 1g)
subovoid with slightly raised margins and
prominent cephalomedian tubercle; main
body with longitudinal median fovea and
transverse grooves along cephalolateral
margins of arched epistomal zygoma. Ven-
tral surface of proximal podomere of an-
tennule with obtuse tubercle near mid-
length. Antennal peduncle lacking spines;
flagellum reaching not quite to caudal mar-
gin of carapace. Antennal scale (Fig. le)
short, stout, lacking usual broad lamellar
part mesially and terminal distolateral spine
scarcely developed and obscured by dense
setal row; about 2.5 times longer than wide;
distal margin provided with dense row of
long, stiff setae; scale reaching about mid-
length of penultimate podomere of anten-
nular peduncle. Ventral surface of ischium
of third maxilliped (Fig. 2b) with sparse band
—
435
of long, stiff setae just mesial to midline and
irregular submedian row of much shorter
stiff setae, often clustered.
Chela (Fig. 11) about 1.7 times longer than
broad, strongly depressed; mesial margin of
palm with row of 7 tubercles, second row
of 5 just medial to it; dorsal surface with
scattered, sometimes setiferous puncta-
tions, but numerous only on fingers; ventral
surface with arched sublateral row of punc-
tations bearing tufts of long setae (Fig. Ic);
distal ridge, opposite base of dactyl, with
subacute tubercle and low rounded one me-
dial to it; middle third of lateral (outer) mar-
gin of chela with row of 4 low tubercles.
Opposable margin of fixed finger with row
of 5 tubercles in proximal two-thirds, mid-
dle one markedly largest; single row of mi-
nute denticles in distal half; small tuft of
setae obscuring margin in most proximal
part; dorsal and ventral surfaces with prom-
inent submedian longitudinal ridge flanked
by punctations; lateral margin strongly cos-
tate. Opposable margin of dactyl with row
of 4 tubercles in proximal half, proximal-
most largest; conspicuous excision in mar-
gin delimited distally by penultimate, next
largest, tubercle; single row of crowded mi-
nute denticles beginning at penultimate tu-
bercle and extending distally to horny tip.
Mesial margin with cluster of 3 tubercles,
one set ventrally to others, in proximal
fourth and row of punctations with long stiff
setae extending distally; dorsal and ventral
surfaces with prominent submedian longi-
tudinal ridge flanked by punctations.
Carpus (rotated about 20° counterclock-
wise in Fig. 11) with prominent longitudinal
furrow dorsally, flanked mesially by irreg-
ular row of 4 tubercles; mesial surface with
twin spikelike spines in distal fourth and 3
much smaller spines proximal to them; ven-
tral surface smooth with 2 small subacute
tubercles in mesio- and laterodistal corners;
region; g, Epistome; h, Lateral view of carapace; 1, Dorsal view of distal podomeres of cheliped; j, Dorsal view
of telson and uropods.
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
436
h
Fig. 2. Types of new species of Fallicambarus (a—d, F. burrisi; e-i, F. gordoni): a, Mesial view of first pleopod
of morphotype; b, Ventral view of ischium of third maxilliped of holotype; c, Lateral view of first pleopod of
VOLUME 100, NUMBER 3
lateral surface punctate with prominent
ventrolateral articulate knob. Dorsodistal
surface of merus entire with small puncta-
tions; ventromesial margin with row of 7
small tubercles, distalmost and antepenul-
timate ones subacute; ventrolateral margin
with row of 6 subacute tubercles intermixed
with row of long stiff setae. Sufflamen well
developed.
Chela of second pereiopod with row of
long setae on both margins of palm and on
dorsal and ventral margins of carpus; distal
half of ventral margin of merus with similar
row of setae.
Ischium of third pereiopod with simple
curved hook not reaching basioischial ar-
ticulation (Fig. 1f) and not opposed by tu-
bercle on basis. Coxa of fourth pereiopod
with large, caudomesially directed boss.
Coxa of fifth pereiopod lacking caudomesial
boss but with setiferous ventral membrane.
First pleopod (Fig. la, d, f) not quite
reaching coxa of third pereiopod, situated
deep within sternum and obscured, partic-
ularly distally, by setae extending caudally
and mesially from ventral margins of ster-
num and proximal margins of coxae of third
and fourth pereiopods; proximomesial spur
lacking; distal half of shaft inclined caudal-
ly; terminal elements consisting of broad,
corneous central projection, scarcely
notched subterminally; and mesial process
attenuate distally but apical third somewhat
flattened in cephalocaudal aspect, entire
process deflected sharply laterad, and tip
not extending beyond tip of central projec-
tion; cephalic process absent.
Allotypic female.—Excluding secondary
sexual characters, differing from holotype in
following respects: small spine on sinistral
margin of telson and none on dextral; mesial
margin of palm with only one tubercle dor-
somedial to mesialmost row; mesial margin
ee
437
of dactyl with row of 3 tubercles in proximal
third; distal ridges of ventral surface of palm
with only one tubercle opposite base of dac-
tyl. Carpus of cheliped with single spikelike
tubercle on distomesial surface; tubercles
associated with ventrodistal margin more
acute. Tubercles associated with ventrome-
sial margin of merus of cheliped all acute.
Annulus ventralis (Fig. 2d) deeply exca-
vate except for lateral and caudal elevated
ridges; as described in Diagnosis. Caudal
overhang of annulus completely obscuring
first abdominal sclerite. First pleopods much
reduced and biramous.
Morphotypic male, Form II.—Differing
from holotype in following respects: mesial
margin of palm with row of 8 tubercles,
second more medial row of 3 near mid-
length, and single tubercle just medial to
central tubercle of latter row; dactyl with
row of 5 tubercles on mesial margin and
with tubercle just dorsal to second from base;
distal ridge of ventral surface of palm with
2 acute tubercles opposite base of dactyl.
Carpus with single spikelike tubercle on me-
sial margin. Ventromesial margin of merus
of cheliped with row of 9 subacute tubercles,
ventrolateral margin with row of 8. Hook
on ischium of third pereiopod as well de-
veloped as in holotype, but caudomesial boss
of coxa of fourth pereiopod less well de-
veloped. First pleopod (Fig. 2a, c) with dis-
tal half not so clearly inclined caudally; both
terminal elements non-corneous, blunter;
central projection lacking subapical notch;
mesial process contiguous to central pro-
jection along basal half, tip extending no
further caudad than central projection.
Type locality. —Burrows in saturated
sandy soil of hillside Sarracenia bog; T4N,
RSW, E/2 Sec. 21, Greene County, Missis-
sIppl.
Disposition of types. —The holotypic male,
morphotype; d, Annulus ventralis of allotype; e, Lateral view of cephalic segments of abdomen of holotype; f,
Annulus ventralis of allotype; g, Dorsal view of telson and left uropod of holotype; h, Ventral view of ischium
of third maxilliped of holotype; i, Antennal scale of holotype.
438
Form I, the allotypic female, and the mor-
photypic male, Form II (USNM 206881,
206883, and 206882, respectively), are lo-
cated in the National Museum of Natural
History, Smithsonian Institution, as well as
a small paratypic series (3 éI, 1 SII, 4 2, 2 4
imm., | 2 imm.). Another series of para-
types (2 461, 9 2, 2 6imm., 3 2 imm.) is at
the Mississippi Museum of Natural Science,
plus one dubiously identified collection,
probably assignable to this species and not
designated paratypic (1 4II, 1 6 imm.).
Variations and color.—Most of the vari-
ations encountered are represented among
the primary types, but some points need to
be noted. In a male, Form II, from Wash-
ington County, Alabama, the setae are ab-
sent from the ventrolateral margin of the
palm, and in the same specimen, the is-
chium of the third maxilliped is not as hir-
sute as usual. Strikingly, in second form
males, the caudomesial boss of the coxa of
the fourth pereiopod is nearly as well de-
veloped as in the first form male. Equally
striking is the remarkable congruence of tu-
bercular ornamentation of the opposable
margins of the fingers; not only are the num-
bers constant, but the relative sizes seem to
be independent of sex and are the same. I
have no specific color notes on this species,
but my field notes in one instance (Wash-
ington Co., Alabama) indicate that the colors
do not differ significantly from those of Fal-
licambarus (Creaserinus) byersi.
Size. —The largest specimen collected is
the morphotype; the smallest first form male
is the holotype, and the largest has a 26.4
mm carapace length; the largest female, 27.4
mm. For measurements see Table 1.
Range and specimens examined.—All
specimens were collected from pitcher plant
bogs in the Chickasawhay and Escatawpa
drainages of Alabama and Mississippi.
ALABAMA. Washington County (all Es-
catawpa drainage): (1) bog, 1.5 mi (2.4 km)
SE of County Road 18 (Vinegar Bend Rd.)
on U.S. Hwy. 45, T3N, R3W, | 61,3 2,24
imm., | 2imm., 6 Apr 1974, J. F. Fitzpat-
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
rick, Jr., and B. A. Laning, colls.; (2) bog,
2.1 mi (3.4 km) NW of Mobile County line
on U.S. Hwy. 45, T2N, R3W, | éII, 25 May
1974, J.F.F., Jr., and B.A.L., colls.; (3) small
swamp stream 9.4 mi(15.1 km) N of Mobile
County line on St. Rte. 17, T3N, R3W, 1
2, 3 2imm., 22 Apr 1970, E. Black and H.
H. Hobbs, Jr., colls.; (4) 6.2 mi (10.0 km)
W of McIntosh on St. Rte. 35, hillside seep-
age, T4N, R2W, 1 61, 12, 21 Apr 1970, E. B.
and H.H.H., colls.; (5) roadside ditch, 5 mi
(8.0 km) N of Citronelle on U.S. Hwy. 45,
T2N, R3W, 1 @II, 1 29, 2 6imm., 2 2? imm.,
21 Apr 1970, E. B. and H.H.H., colls.
MISSISSIPPI. Greene County (all Chick-
asawhay drainage): (6) about 9.5 mi (15.3
km) S of town of State Line, 1 ¢I, 1 II
(morphotype), 1 2, 10 May 1979, John W.
Burris, coll.; (7) about | mi W of Yellow
Pine, Alabama, TSN, RSW, NE/4 Sec. 14,
1 61, 2 Aug 1979, J.W.B., coll; (8) type lo-
cality, 1 I, 1 2 (holo- and allotype), 25 Jun
1981, J.W.B., coll.; (9) Kurtz State Forest,
seepage area in headwaters of Brannon
Creek, T4N, R5W, SE/4 Sec. 21, 3 2, 1 9
imm., 29 Mar 1983, S. E. Mott and M. Ste-
gall, colls.; Jackson County (all Escatawpa
drainage): (10) 0.75 roadmi (1.2 km) W of
Escatawpa River on St. Rte. 614, T5S, RSW,
SE/4 SW/4 SE/4 Sec. 2, 2 2, 2 6imm., 2 2
imm., 1 unsexed carapace fragment, 19 Apr
1983, R. L. Jones and S.E.M., colls.; (11)
4.8 airmi (7.7 km) SW of George County
line at Alabama state line, T4S, RSW, NE/4
NE/4 SE/4 Sec. 16, 3 2, 15 Aug 1984, R.L.J.,
coll.; (12) (probably assignable to this
species) T4S, RSW, SE/4 SW/4 NW/4 Sec.
16, 1 61, 1 ¢imm., 12 Dec 1984, R.L.J., J.
Wiseman, and R. Lohoefnar, colls.
Environmental notes. —The species seems
always to be associated with pitcher plant
bogs, always burrowing, and usually into a
sandy clay substrate, exhibiting complex
branching patterns in the burrow design;
often a trickle of water runs through the
habitat—except perhaps in midsummer. My
best notes are from locality no. 2, above.
There Sarracenia leucophylla was the dom-
VOLUME 100, NUMBER 3
inant plant, by far, but S. alata, S. psitta-
cina, Dichomena latifolia, Drosera rotun-
diflora and Stockesia laevis were also
conspicuous. At that time, Cambarus (La-
cunicambarus) diogenes ludovicianus Fax-
on, 1884, was also dug from burrows in the
area. The site had been visited earlier (on 6
Apr) when the dominant identifiable plants
were S. alata and D. rotundiflora, the for-
mer in bloom. Also identifiable were S’.. psit-
tacina and S. leucophylla. In flower were D.
latifolia, S. laevis, Drosera filiformes and
Lachnocaulon anseps. Fallicambarus bur-
rows were provided with freshly worked and
thoroughly plugged chimneys 6-8 in (15.2-
20.3 cm) tall. A solid plug filled the burrow
to a depth of 8-10 in (20.3—25.4 cm), and
beneath this, the burrow was filled with a
slurry of about 50% water and 50% sand
and clay. Only rarely was a clear chamber
encountered, and no crawfishes were col-
lected although several burrows were ex-
cavated as far as could be detected. About
12 in (30.5 cm) below the surface, burrow
temperature fell sharply to about 65°F from
about 85°F air temperature.
Relationships. — Fallicambarus (Creaser-
inus) burrisi has its closest affinities with F.
(C.) byersi, F. (C.) gordoni, n. sp., below,
and F. (C.) caesius Hobbs, 1975, and more
remotely to F. (C.) danielae Hobbs, 1975,
and F. (C.) oryktes (Penn and Marlow 1959).
Among the common features shared by the
first four species are a caudally inclined gon-
opod of the first form male, a conspicuous
row of long setae on the ventrolateral sur-
face of the palm of the chela, and the ab-
domen of the male being markedly narrow-
er than the carapace. The annulus of the F.
(C.) burrisi female, however, is unique; in
no other species of the genus is it so scoop-
like and does it project so far caudad. The
conspicuous boss on the coxa of the male
fourth pereiopod is not matched by any close
relative. The attenuate tip of the mesial pro-
cess of the first form male extends no further
than the apex of the central projection, a
characteristic shared with F. (C.) byersi and
439
F. (C.) oryktes. It can be separated from F.
(C.) oryktes by the row of setae on the ven-
trolateral margin of the palm and from F.
(C.) byersi by the lack of an inflated part of
the mesial process which overlaps the cau-
dal margin of the central projection. The
relatively slender terminal elements of the
first pleopod and overall configuration of
the palm of the cheliped are more like F.
(C.) oryktes than the other mentioned
species, but burrisi shares more features with
F. (C.) byersi, F. (C.) caesius, and F. (C.)
danielae than with oryktes.
Fallicambarus (Creaserinus) gordoni,
new species
Figs. 2e-i, 3
Diagnosis. — Antennal scale reduced, usu-
al lamellar development of mesial part re-
duced. Ventrolateral surface of propodus of
chela with row of long stiff setae near lateral
margin; coxa of fourth pereiopod of first
form male with small, but conspicuous,
transversely-oriented caudomesial boss.
First pleopods of first form male inclined
caudally in distal one-fourth; apex of central
projection directed at angle of about 120°
to main axis of pleopod, but mesial process
at nearly right angle. Annulus ventralis of
female about as long as wide, deeply exca-
vate cephalically and sigmoid sinus so de-
veloped that opposing halves of raised cau-
dal part resembling interlocking fingers;
annulus projected caudad only slightly,
overhang scarcely, if at all, obscuring pos-
tannular sclerite.
Holotypic male, Form I.—Cephalothorax
subovate, compressed (Fig. 3b, f); eyes re-
duced. Abdomen much narrower than tho-
rax (6.0 and 9.1 mm); greatest width of car-
apace less than depth at caudodorsal margin
of cervical groove (9.1 and 10.5). Areola
obliterated along middle half of length;
length 37.9% of entire length of carapace
(43.0% of postorbital carapace length). Ros-
trum only slightly depressed, with moder-
ately converging, slightly thickened mar-
440 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig. 3. Fallicambarus (Creaserinus) gordoni, holotype: a, Mesial view of first pleopod; b, Dorsal view of
carapace; c, Lateral view of chela; d, Lateral view of first pleopod; e, Ventral view of thoracic region; f, Lateral
VOLUME 100, NUMBER 3
gins, latter constricting from near midlength
to obtuse acumen; tip not upturned and
reaching not quite to base of ultimate pod-
omere of antennular peduncle; dorsal sur-
face concave cephalically becoming sub-
planar caudally. Subrostral ridges poorly
evident in dorsal aspect. Postorbital ridges
prominent, grooved laterally and terminat-
ing cephalically in weak tubercles. Subor-
bital angle virtually obsolete; branchiostegal
spine absent. Carapace sparsely punctate
dorsally with slightly granulate branchio-
stegites.
Abdomen shorter than carapace (16.5 and
21.1 mm); pleura quite short and very
broadly rounded; cephalic lobe of pleuron
of second segment not prominent but over-
lapping reduced pleuron of first (Fig. 3h).
Telson undivided (Fig. 2g), lacking lateral
spines. Proximal podomere of uropod with
subacute tubercle on mesial lobe; both rami
rounded distally; cephalic portion of lateral
ramus with small fixed spine in distolateral
corner and row of small acute spines along
entire distal margin, median one only very
slightly larger than rest; mesial ramus with
weakly developed median ridge, but lacking
spines.
Cephalomedian lobe of epistome (Fig. 3g)
broadly subtriangular with slightly raised
margins and weak cephalomedian tubercle;
main body with small median fovea near
cephalic end and transverse grooves along
cephalolateral margins of arched epistomal
zygoma. Ventral surface of proximal podo-
mere of antennule with small tubercle near
midlength. Antennal peduncle lacking
spines; flagellum reaching about midlength
of areola. Antennal scale (Fig. 21) short,
stout, usual broad lamellar mesial part much
reduced; terminal distal spine of expanded
lateral portion small but stout and subacute;
about 2.1 times longer than wide; mesial
and mesiodistal margins with dense row of
long, stiff setae, such setae sparse on lateral
—_—
441
margin; scale reaching about midlength of
penultimate podomere of antennal pedun-
cle. Ventral surface of ischium of third max-
illiped (Fig. 2h) with row of long, stiff setae
just mesial to midline and widely scattered,
sometimes clustered submedian row.
Chela (Fig. 31) about 1.8 times longer than
broad; strongly depressed; mesial margin of
palm with single row of 6 tubercles; dorsal
surface with scattered setiferous puncta-
tions, most numerous near base of immov-
able finger; ventrolateral surface with arched
row of punctations, each with one or more
long, stiff setae (Fig. 3c); distal ridge, op-
posite base of dactyl, with low tubercle;
proximal third of lateral (outer) margin of
palm with row of 4 low tubercles. Oppos-
able margin of fixed finger with row of 4
tubercles in proximal half, second from base
markedly largest, distal subequal in size and
smaller; single row of minute denticles on
distal half, beginning just distal to penulti-
mate tubercle; tuft of setae in most proximal
part almost negligible; dorsal and ventral
surfaces with prominent longitudinal ridges
flanked by punctations; lateral margin
strongly costate. Opposable margin of dac-
tyl with 4 tubercles in proximal half, 2 in
conspicuous excision largest, and penulti-
mate one delimiting distal extent of exci-
sion; single row of minute denticles begin-
ning just distal to ultimate tubercle and
extending nearly to corneous tip; mesial
margin with complex ridge pattern in basal
third but lacking tubercles; long setae prom-
inent only in distal half; dorsal and ventral
surfaces with prominent longitudinal ridge
flanked by punctations.
Carpus with prominent longitudinal fur-
row dorsally, flanked mesially by few punc-
tations; mesial surface with spikelike tuber-
cle in distal fourth and irregular row of 4
subacute tubercles proximal to it; ventral
surface smooth with low subacute tubercle
in each distal corner; lateral surfaces sparse-
view of carapace; g, Epistome; h, Lateral view of cephalic segments of abdomen; i, Dorsal view of distal podomeres
of cheliped.
442
ly punctate with few long setae and prom-
inent ventrolateral articular knob. Dorso-
distal surface of merus entire with few small
punctations; ventromesial margins with row
of 10 rounded to slightly subacute tubercles;
ventrolateral margin with row of 7, ante-
penultimate and penultimate bifid, low tu-
bercles flanked medially by row of long stiff
setae. Sufamen well developed.
Chela of second pereiopod with row of
long setae on both margins of palm, and on
dorsal and ventral margins of carpus; distal
half of ventral surface of merus with similar
row.
Ischium of third pereiopod with simple,
slightly curved hook not overreaching ba-
sioischial margin (Fig. 3e) and not opposed
by tubercle on basis. Coxa of fourth pereio-
pod with small but prominent, transversely
oriented caudomesial boss. Coxa of fifth pe-
reiopod lacking boss but with setiferous
ventral membrane.
First pelopod reaching just beyond caudal
margin of coxa of third pereiopod, situated
deeply within sternum, and somewhat ob-
scured, particularly distally, by setae ex-
tending caudally and mesially mostly from
ventrolateral margins of sternum; proxi-
momesial spur lacking; distal fourth of shaft
inclined caudally (Fig. 3a, d, e); terminal
elements consisting of moderately broad,
corneous central projection, obtusely
notched subapically; mesial process sub-
spatulate in distal half, expanded and some-
what compressed laterally just proximal to
midlength but not contiguous with adjacent
central projection; cephalic process absent.
Allotypic female. —Excluding secondary
sexual characters differing from holotype in
following respects: merus of cheliped with
8 (none bifid) spinose tubercles on ventro-
lateral margin; ventromesial row of 9; inner
margin of palm with small squamous tu-
bercle just dorsolateral to third from prox-
imal end of row.
Annulus ventralis (Fig. 2f) about as broad
as long, deeply excavate cephalically; caudal
margin elevated, sinistral half tilted ce-
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
phalically overhanging more cephalic part
of caudal half of annulus; sinus originating
beneath overhang near midline and extend-
ing cephalically short distance before re-
curving caudally, forming inverted-U, and
disappearing before reaching caudal mar-
gin; dextral elevated portion flanking ce-
phalic and lateral sides of sinus disappear-
ing beneath sinistral overhang, sculpture
providing overall appearance of interlock-
ing fingers when viewed ventrally. Postan-
ular sclerite reduced and subconical; pro-
vided with setae. First pleopod reduced and
biramous.
Male, Form IIT.—No morphotype ts des-
ignated because in all of the second form
males I have seen there is so much vari-
ability, especially in the first pleopod, that
I cannot select one as “‘typical”’ of the species.
There is no consistency in the relative lengths
of the terminal elements; they are subequal
in 26.7%, the mesial process longer in 60.0%,
and the central projection longer in 13.3%.
The elements are bent (in relation to the
principal axis of the basal part of the pleo-
pod) about 90° in 33.3%, about 135° in
20.0%, between 90 and 135° in 33.3%, and
up to 180° in 13.4%. The two elements are
contiguous through at least 90% of their
lengths in 37.5%, but are not so in 62.5%.
The central projection is always blunt or
bluntly rounded apically, but the mesial
process, although always tapering from base
to tip, can be described as apically con-
stricted only in 20% of the specimens. None
of these characteristics seems to be corre-
lated with each other; neither does size seem
to be a factor in the nature of their expres-
sion. Two other characters seem to be more
developed as the animal gets larger: the hook
on the ischium of the third pereiopod and
the caudomesial boss of the coxa of the
fourth. Nevertheless, even this relationship
is not absolute. In five specimens the hook
was aS prominent as in the holotype (and
other first form males). Their carapace
lengths ranged from 24.1 to 30.4 mm, but
the hook was much less developed in three
VOLUME 100, NUMBER 3
other animals of the same size range (24.8,
24.8 and 25.3). Generally, the boss size was
correlated with hook size; in one second
form male, however (carapace length 28.0),
it was scarcely identifiable. In one animal,
the boss was rounded, although not as much
so as in F. (C.) burrisi, and the associated
females were clearly assignable to F. (C.)
gordoni. Other types of variation are dis-
cussed below.
Type locality.—DeSoto National Forest,
Camp Shelby Military Reservation, T2N,
R10W, SE/4 NE/4 SE/4 Sec. 5, Perry Coun-
ty, Mississippi. Here the animals were dug
from multiple-branched burrows in satu-
rated sandy soil at the edge of a pitcher plant
savannah. Nearby (within | km) burrows
yielded specimens of Cambarus (Lacuni-
cambarus) diogenes ludovicianus and Pro-
cambarus (Ortmannicus) planirostris Penn,
1953.
Disposition of types.—The holotype and
allotype (USNM 206877 and 206878, re-
spectively) are deposited in the National
Museum of Natural History, Smithsonian
Institution, together with a paratypic series
(1 411, 22, 1 2imm., | 2ov.); The Mississippi
Museum of Natural Science has a larger
paratypic series (4 41, 19 6H, 19 2, 3 $imm.,
4 2imm.).
Variations. —Some of the variability en-
countered is encompassed within the de-
scriptions of the types and discussion of the
second form male, above, but other differ-
ences which were encountered merit dis-
cussion here. In all but two of the females
(1 adult, 1 juvenile) the annulus was a mir-
ror image of the allotype. In two mature
females the sinus 1s more broad than that
of the allotype, and, independent of matu-
rity, in three the cephalic overhang of the
caudal elevation 1s more pronounced. In
about half the specimens, the tip of the ros-
trum is slightly upturned, but in a nearly
equal number, the apical development is so
unpronounced that one could not term the
slight compression there an acumen. In
nearly two-thirds of the specimens the ce-
443
phalic extremity of the postorbital ridge
merges imperceptibly into the cephalotho-
rax, a Situation somewhat more prevalent
in larger than smaller specimens. The ceph-
alolateral margins of the epistomal zygoma
in many specimens have deeper pits asso-
ciated with their cephalolateral bases than
do the type specimens.
A remarkably early establishment of the
pattern of tubercles associated with the op-
posable margins of the fingers of the chela
apparently occurs. The smallest animals I
have seen (15.4-19.0 mm cephalothorax
length) all have a hand which in size and
ornamentation allows them to be unques-
tionably assigned to this species. In some
adults, however, there are differences in the
arrangement and number of tubercles that
are not clearly associated with regeneration
of the member. Two females and a second
form male from locality (8) below have the
more basal tubercle in the excision of the
movable finger larger than the next most
distal, and the tubercle which marks the dis-
talmost limit of the excision 1s, by far, the
largest of the three; just distal to this last-
mentioned tubercle are two small rounded
ones. The proximolateral margin of the dac-
tyl has a subtriangular cluster of three low
tubercles. On the opposable margin of the
immovable finger opposite the dactyl ex-
cision is a small, but still prominent, tu-
bercle proximal to the usual two. Several
animals from several localities also have
some kind of tubercular development in the
proximomesial region of the dactyl.
Setal development is likewise variable. In
about half the specimens, the ischium of the
third maxilliped has more than one, often
arranged in a row, medial punctation in its
basal half from which a tuft of long stiff setae
emerges; in two animals from different lo-
calities, their maxillipeds were asymmetri-
cal with respect to this character. Although
always present, the row of setiferous punc-
tations along the ventrolateral surface bears
setae of different lengths and number; in
only one specimen were the setae as long as
444
Table 1.—Measurements (in mm) of types of Fal-
licambarus (Creaserinus) burrisi n. sp. and F. (C.) gor-
doni n. sp.
Morpho-
Holotype Allotype type
F. (C.) burrisi
Carapace
Total length De 2308 2926
Postorbital length 20.1 D2 26:5
Width 9.6 10.6 Ned
Height 10.8 10.9 14.5
Areola
Length 8.6 9.0 12.0
Rostrum
Length 3.5 3.7 5.0
Width 4.0 3.8 5.0
Chela
Length, mesial margin
palm 4.1 3.7 Do
Width, palm 6.9 6.1 9.5
Length, lateral margin
propodus 11.9 11.8 16.9
Length, dactyl 7.0 7.4 10.5
Abdomen
Length 17.3 ee me 73)32)
Width 6.2 6.6 9.0
F. (C.) gordoni
Carapace
Total length Paibed| 21.0 —
Postorbital length 18.6 18.6 —
Width 9.1 8.5 —
Height 10.5 9.4 —
Areola
Length 8.0 8.1 —
Rostrum
Length 3.6 4.3 —
Width 4.0 4.4 —
Chela
Length, mesial margin
palm 3.7 3.0 —
Width, palm 6.1 SED —
Length, lateral margin
propodus 11.2 11.0 —
Length, dactyl 7.0 7.3 —
Abdomen
Length 16.5 10.2 —
Width 6.0 5.6 _
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
found in F. (C.) byersi and F. (C.) burrisi.
The setae along the lateral margin of the
dactyl also vary in number and length, but
in only one specimen was there no trace of
such.
The apex of the central projection of the
first form male is blunt, but only in the ho-
lotype and one other is there any suggestion
of a notch, and such indication is most pro-
nounced in the holotype. In the first form
male from locality (4) below the hook on
the ischium of the third pereiopod is so
acutely arched that the tip clearly over-
reaches the basioischial margin.
Size. — The smallest first form male is 21.0
mm in cephalothorax length, and the larg-
est, 25.5. The largest second form male is
30.4 mm. The largest female is 32.5 mm
and is the largest animal collected; the ovig-
erous female is 28.1 mm in cephalothorax
length. For measurements see Table 1.
Range and specimens examined. —This
crawfish is known from eight locations, all
in the DeSoto National Forest (Pascagoula
River drainage) in Perry County, MISSIS-
SIPPI: (1) type locality, 1 3I, 1 2 (holo- and
allotype, respectively), 25 Oct 1982, Robert
L. Jones, coll.; (2) Camp Shelby Military
Reservation, T2N, Rl1OW, NW/4 SW/4
SW/4 Sec. 4, 1 él, 1 2, 1 2 imm., 5 Nov
1982, R.L.J., K. L. Gordon, S. Mott, colls.;
(3) Camp Shelby Military Reservation, T2N,
R1OW, SE/4 SW/4 NW/4 Sec. 15, 1 2, 1 2
ov., 9 Nov 1982, R.L.J., S.M., and K.L.G.,
colls.; (4) TZN, RIOW, NE/4 NW/4 NE/4
Sec. 8, 3.3 airmi (5.3 km) SSE of New Au-
gusta, | 6I, 3 2°, 29 Mar 1985, R.L.J., coll.;
(5) compartment 71, TZN, RIOW, NW/4
NE/4 SW/4 Sec. 20, bog on S side of Forest
Service Rd. 304, about 0.45 airmi (0.7 km)
SSW of jct. F. S. Rd. 304 and St. Rte. 29,
3 61, 7 All, 3 2, 18 Mar 1986, R.L.J. and
K.L.G., colls.; (6) compartment 62, T2N,
R1IOW, SE/4 NE/4 NW/4 Sec. 21, about 0.7
airmi (1.1 km) NE of crossing of St. Rte. 29
over Cypress Creek, 4 II, 5 2, 18 Mar 1986,
R.L.J. and K.L.G., colls.; (7) TZN, R1OW,
NW/4 SW/4 SW/4 Sec. 4, 1 ¢II, 1 2, 1 ¢
VOLUME 100, NUMBER 3
imm., 1 2imm., 29 Mar 1985, R.L.J., coll.;
(8) compartment 61, T2N, RIOW, SE/4
SE/4 SW/4 Sec. 8, small bog area just S of
active oil well, about 4.1 airmi (6.6 km) SSE
of jct. U.S. Hwy. 98 and St. Rte. 29 in New
Augusta, 6 4I1I, 6 2, 1 ¢6imm., 2 2imm., 18
Mar 1986, R.L.J. and K.L.G., colls.
Life history and environmental notes. —
First form males were taken on 18 and 29
March and 29 October; the ovigerous fe-
male was captured on 9 November, at which
time she was carrying about 36 fertile eggs
and was discovered within the chimney,
above the water table. All were taken from
saturated sandy soil associated with pitcher
plant savannahs. Collections 1-3, above,
were dug from the complex burrows, but
the subsequent ones were collected using a
modification ofa trap designed by Norrocky
(1984) for use in Ohio. These traps have
yielded remarkable results for the Missis-
sippi Museum personnel in their studies of
south Mississippi habitats. The greatly in-
creased number of specimens per unit effort
afforded by this technique has permitted
them to accumulate an excellent collection
of burrowing crawfishes from the Jackson
Prairie, which is, except for Hobbs’ efforts
in Georgia and South Carolina, spread over
many more years, possibly the most thor-
ough representation of such animals from a
particular region available anywhere.
I have not collected the animals myself,
so I have no details of associated plants or
color notes. In one formalin-preserved
specimen, however, a lavender ground col-
or, not unlike F. (C.) byersi, was evident
when I first examined the specimen. I also
was able to examine methanol-preserved
specimens from localities 5, 6, and 8, above,
within one month of their collection. In all
a distinct lavender color, similar to that seen
in byersi, was evident on the hand or dactyl
or both. In two of the first form males, the
rostral margins and postorbital ridges were
deep purple.
Relationships. — Fallicambarus (Creaser-
inus) gordoni is most closely related to F.
445
(C.) danielae but it is easily distinguished
by a broader rostrum and the presence of
two, rather than one, prominent tubercles
in the excision on the opposable margin of
the dactyl. In first form males the mesial
process extends scarcely beyond the tip of
the central projection, and the notch at the
apex of the latter, if present, is never so
obvious as in danielae. In females, the sinus
begins in the far caudal half of the annulus,
beneath an overhanging elevation of the
caudal margin. In F. (C.) danielae the tip
of the mesial process extends well beyond
the apex of the central projection, which
projection is also clearly notched subapi-
cally; the annulus of the female has no over-
hanging, conspicuously elevated caudal
margin, and the sinus originates near the
middle of the annulus. The central projec-
tion and mesial process of F. (C.) danielae
are directed so that their apices are more or
less 90° to the principal axis of the basal
part of the pleopod; in F. (C.) gordoni the
central projection apex is directed at least
150° and the mesial process as much as 145°.
See also the comments under the “Rela-
tionships’ section for F. (C.) burrisi, above.
Acknowledgments
I am indebted to all those people iden-
tified in the lists of collectors, and especially
to John W. Burris and Kenneth L. Gordon,
successive coordinators of the Mississippi
Natural Heritage Program, who made dil-
igent searches for these beasts; equally tire-
less in such efforts were Robert L. Jones, of
the Program, and Paul D. Hartfield of the
Mississippi Museum of Natural Science.
Horton H. Hobbs, Jr., was his usual gen-
erous self in offering many helpful sugges-
tions during this work. My friend and col-
league, Michel G. Lelong, helped with the
identification of plant associates.
Literature Cited
Faxon, Walter. 1884. Descriptions of new species of
Cambarus to which is added a synonymical list
446
of the known species of Cambarus and Asta-
cus. — Proceedings of the American Academy of
Arts and Sciences 20:107-158.
Hobbs, Horton H., Jr. 1941. Three new Florida cray-
fishes of the subgenus Cambarus (Decapoda,
Astacidae).— American Midland Naturalist 26:
110-121.
1975. New crayfishes (Decapoda: Cambari-
dae) from the southern United States and Mex-
ico.—Smithsonian Contributions to Zoology
201:111 + 34 pp.
Norrocky, M. James. 1984. Burrowing crayfish trap.—
Ohio Journal of Science 84:65-66.
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Penn, George Henry, Jr. 1953. A new burrowing
crawfish of the genus Procambarus from Loui-
siana and Mississippi (Decapoda, Astacidae). —
Tulane Studies in Zoology 1:71-76.
, and Guy Marlow. 1959. The genus Cam-
barus in Louisiana.—American Midland Nat-
uralist 61:191—203.
Department of Biological Sciences, Uni-
versity of South Alabama, Mobile, Ala-
bama 36688.
PROC. BIOL. SOC. WASH.
100(3), 1987, pp. 447-451
TWO NEW OPERCULATE LAND SNAILS FROM THE
PALAU ARCHIPELAGO
Fred G. Thompson and Thomas M. Iliffe
Abstract. —Georissa zea Thompson, n. sp. (Gastropoda, Prosobranchia, Hy-
drocenidae) is described from Machachar Island. It differs from other species
by its large size and regularly-spaced granular sculpture. It is the first hydrocenid
recorded from the Palau Archipelago. A second species, G. rufula, also occurs
there. Pupina nitidula Thompson, n. sp. (Gastropoda, Prosobranchia, Pupin-
idae) is described from Ngeruktabel Island. It is differentiated by its small size
and slender form.
The land snail fauna of the Palau Islands
is better known than that of most other
western Pacific archipelagos. Four of the six
numerically dominant groups of snails in-
habiting the islands have been mono-
graphed in recent years, based primarily
upon collections made by the Bernice P.
Bishop Museum Micronesian Expedition in
1936 (Baker 1938-1941; Clench 1949;
Cooke and Kondo 1960; Solem 1976, 1983).
Notwithstanding these revisionary studies,
the fauna remains only partially known, even
among those families that are mono-
graphed. Species have been reported from
only six of the nearly 200 islands that com-
prise the archipelago, and only two (Koror
and Peleliu) have been sampled at more than
a few stations.
Recent field work in Palau by the authors
has added important collections from sev-
enteen other islands and islets not reported
upon in earlier studies. The new snails de-
scribed below are from two of these islands.
Georissa zea Thompson, new species
Figs. 1-4
Diagnosis. — A moderately large species
characterized by its light yellow color, its
Ovate-turbinate shape, and teleoconch
sculpture consisting of strong spiral cords
bearing nodes that are synchronized along
growth lines.
Description. —Shell light yellow in color.
Moderately thick, opaque. About 6.5 mm
long at maturity. Ovate-turbinate in shape
(Fig. 1); about 1.4 times as wide as high in
adult shells. Spire moderately high, about
1.4 times height of aperture. Whorls 4.2.
Protoconch strongly protruding; sculptured
with dense mesh of minute longitudinal
granules (Fig. 3); initial half whorl horizon-
tal, 0.30 mm wide by 0.40 mm long. Whorls
of teleoconch uniformly rounded with deep-
ly impressed suture; sculptured with strong
spiral cords adorned with regularly-spaced
rounded nodes synchronized along growth
lines; cords and nodes nearly uniform in size
over surface of shell; 19 cords on body whorl,
10 on penultimate whorl and 8 on antipen-
ultimate whorl. Umbilicus closed but deep-
ly indented with narrow rimate impression
at maturity (Fig. 2). Columellar lip rounded,
not forming broad, flattened plate. Imma-
ture shells imperforate; with broad plate-
like expansion of columella, typical of Geo-
rissa (Fig. 4). Aperture broadly ovate-tri-
angular in shape; 0.9 times as high as wide.
Peristome continuous across parietal wall;
thickened along posterior corner; basal lip
and columellar lip nearly straight and form-
ing pronounced angle; baso-columellar cor-
ner sharp and weakly projecting forward.
Inner surface of holotype operculum with
thick calcereous peg projecting to right and
attached to plate to margin. (The peg was
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Figs. 1-4. Georissa zea Thompson, new species: 1-3, Holotype (UF 90523); 4, Paratype (UF 90524). En-
largements: 1, 2, 4 x 26; 3 x 175.
broken at this point in the only specimen
available.) Outer face paucispiral; nucleus
located at left-basal margin. Outer surface
covered with thin calcareous layer. Basal
plate surrounded by a chitinous fringe. Bas-
al plate 0.86 mm wide and 1.15 mm high.
Measurements in mm of two specimens
follow.
length width aper. h. aper. w. whorls
G4) Gell DS 2.54 4.2
3.41 Meld 1.67 1.80 3h)
Type locality. —Republic of Belau (Palau
Islands), northwest point of Machachar Is-
land, 134°22'20’E, 07°08'45”’N. HOLO-
TYPE: UF 90523; collected 22 Oct 1985 by
Fred G. Thompson. PARATYPE: UF
Specimen
Holotype
Paratype
VOLUME 100, NUMBER 3
90524: same data as holotype. The holotype
and paratype were gold-plated for SEM
study.
The area at the type locality consists of a
strongly karsted limestone terrain in a pri-
mary rainforest with a sparse understory of
woody shrubs and vines. The ground was
covered with a thin layer of dead leaves.
The holotype, the only live specimen found,
was on the underside of a rotting piece of
wood. The single paratype was found in leaf
jitter
Remarks. — This species is unique among
Georissa because of its sculpture. The um-
bilical indentation of the holotype is also
unique. All other Georissa are imperfo-
rate and have a broad plate-like colu-
mellar lip as is depicted in Fig. 4. The um-
bilical pit of the holotype may be a gerontic
teratology. It appears to be caused by rolling
of the columella. Not withstanding the
structure of the umbilical area this species
is readily identified by its sculpture and its
size. It is the largest Georissa known from
the Pacific or the Philippine Islands.
This is the first species of Hydrocenidae
recorded from the Palau Islands. A second
species that was originally described from
Panope, Georissa rufula Moellendorff, 1900,
is widely distributed in the Palau Archipel-
ago. It is distinguished from G. zea by its
minute size, being about 2.5 mm long, and
its smooth reddish shell that is sculptured
with very fine spiral striations. Usually, the
striations are apparent only on fresh shells.
Zilch (1973, pl. 13, fig. 22) illustrates the
lectotype of G. rufula. Both species were
found together on Manchachar Island. Geo-
rissa rufula also inhabits leaf-litter. Three
other species, G. elegans Quadras and
Moellendorff, 1894, G. biangulata Quadras
and Moellendorff, 1894, and G. laevigata
Quadras and Moellendorff, 1894, are de-
scribed from Guam. Numerous other species
are known from the Philippine Islands,
southeast Asia, Indonesia, Australia and
various south Pacific islands. None of these
has sculpture that is similar to that of G.
zed.
449
YES ae
ees
5
eas
6
Figs. 5-8. 5-7, Pupina nitidula Thompson, new
species, holotype (UF 79186). 8, Pupina difficilis Sem-
per (UF 84348), Peleliu Island, Kloulklubed. Scale
equals 2 mm.
Etymology.—The species name zea is
taken from the generic name for maize, Zea,
and alludes to the noded sculpture arranged
in rows much like kernels on a corncob.
Pupina nitidula Thompson, new species
Figs. 5—7
Diagnosis.—A species of Pupina s.s. that
is not closely related to other known species
of the genus. It is characterized by its small
size, slender shape, and notches in the peri-
stome. The small, slender, transparent shell
is flexed weakly to the left at the third whorl.
The palatal lip inserts on the front of the
shell (Fig. 5), not on the side as in other
members of Pupina s.s. such as P. difficilis
Semper. The posterior corner of the aper-
450
ture has a nearly tubular channel formed by
the parietal tooth and a tubercular projec-
tion on the palatal lip. The columellar pore
lies deep behind the columella and is con-
nected to the peristome by a narrow oblique
slit.
Description. — Shell small, about 4.5 mm
long; ovate-cylindrical in shape with round-
ed apex. Shell weakly flexed to left at third
whorl (Fig. 5). Slender, 0.46—0.51 times as
wide as high. Surface smooth and glossy;
transparent; grayish white. Whorls 4.8—4.9;
weakly arched between sutures. Penulti-
mate whorl flattened above front of aper-
ture. Suture weakly impressed with distinct
hyaline subsutural zone. Aperture circular,
0.49-0.51 times width of shell. Peristome
nearly continuous across parietal wall as low
rounded callus merging into parietal lamella
at base. Peristome lying in plane with an-
terior slope of penultimate whorl (Fig. 6).
Peristome incised by parietal sinus and col-
umellar pore (Fig. 7). Parietal sinus formed
by oblique parietal lamella extending into
aperture for about ’s whorl forming narrow
channel along posterior corner of aperture;
upper end of palatal lip with tubercular pro-
jection partially overlapping outer edge of
parietal lamella. Parietal lamella not ex-
tending beyond edge of aperture. Columel-
lar pore located at base of columella, con-
sisting of small elliptical pore connected to
margin of peristome by deep, narrow,
oblique slit (Fig. 6). Pore nearly vertical and
opening internally behind flattened colu-
mellar wall.
Measurements in mm of three specimens
are as follows. The aperture is measured
internally.
length width aper. h. aper. w. whorls
Holotype! 4.46 2.29 1.05 Nel 4.9
Paratype? 4.40 SI 0.99 1.05 4.9
Paratype? 4.46 DAVS 1.05 1.05 4.8
' UF 79186; ? UF 79187; ? UF 79188.
Type locality. —Republic of Belau (Palau
Islands), southeast end of Ngeruktabel Is-
land, 134°26'50”E, 07°15'30”N; 150 m al-
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
titude. The type locality is 1 km north of
the ruins of a World War II Japanese artil-
lery installation, and is just a few meters
below the crest of the island on the east
slope. The area is on limestone substrate
covered by rain forest with very little under-
story or ground vegetation. Specimens were
found deep in leaf-litter in association with
Pupina difficilis Semper. HOLOTYPE: UF
79186; collected 23 Oct 1985 by Fred G.
Thompson. PARATYPES: UF 79187, UF
79188; same data as holotype.
Pupina nitidula is known only from the
type locality. Ngeruktabel is a long narrow
crescent-shaped island about 30 km long
and 1-2 km wide. The island is uninhabited.
It is accessible only at a few points along
the shore, because it is surrounded by nearly
continuous vertical cliffs. Six field collec-
tions were made by the authors and Jeffry
Bozanic from the east end of Ngeruktabel
Over a linear distance of less than five km.
The central region is higher and broader. It
has not yet been explored for land snails.
Remarks. —Pupina nitidula is readily
identified by its small size, its slender shape
with a rounded apex, and its aperture notch-
es. It is the smallest known Pupina s.s. It is
not closely related to other known species
because of the structure of the aperture. In
other species the embayment between the
parietal lamella and the palatal lip is broad-
er, the palatal lip inserts in a more lateral
position, and the columellar pore 1s broader
with a more rounded perforation connected
to the edge of the peristome by a horizontal
transverse slit. These characters ae typically
depicted in Pupina difficilis Semper. Clench
(1949) illustrates all of the known Pacific
Pupina, including the type species, Pupina
keradreni Vignard. None approaches Pup-
ina nitidula in the characters of the aperture.
Pupina nitidula is most similar to Pupina
difficilis because of its bluntly rounded apex.
Pupina difficilis is widely distributed
throughout the Palau Islands, and is the only
other Pupina from there. It was found with
Pupina nitidula at the type locality.
VOLUME 100, NUMBER 3
Etymology. —The species name nitidula
is from the Latin nitidus, diminutive, and
refers to the elegant, shiny aspect of this
small snail.
Acknowledgments
Field work by Thompson was supported
in part by a grant from the National Science
Foundation, International Programs Divi-
sion. Field work by Iliffe was supported by
National Science Foundation grant BSR-
8417494. We are grateful to Jeffry Bozanic
and Dennis Williams, Miami, Florida, who
assisted Iliffe in the field, and who proved
to be exceptionally capable collectors. We
also wish to thank Toshiro Paulis, Director
of the Micronesian Mariculture Demon-
stration Center, Harno Adilbai, Depart-
ment of Entomology, and Herman Francis-
co, Chief, Division of Agriculture, Koror,
Republic of Belau, for the many courtesies
extended to us during our visits to the is-
lands.
Literature Cited
Baker, H. B. 1938-1941. Zonitid snails from the Pa-
cific Islands. — Bulletin of the Bernice P. Bishop
Museum (158, 165, 166): 1-370. [continuous se-
quence]
451
Clench, Wm. J. 1949. Cyclophoridae and Pupinidae
of Caroline, Fijian and Samoan islands. —Bul-
letin of the Bernice P. Bishop Museum (196): 1—
S2,
Cooke, C. M., Jr., and Y. Kondo. 1960. Revision of
the Tornatellinidae and Achatinellidae
(Gastropoda, Pulmonata).— Bulletin of the
Bernice P. Bishop Museum (221):1-303.
Moellendorff, O. F. von. 1900. The land shells of the
Caroline Islands.— Journal of Malacology 5:101—
124.
Quadras, J. F., and O. F. von Moellendorff. 1894.
Diagnoses specierum novarum a J. F. Quadras
in insulis Mariannis collectarum scripserunt. —
Deutsche Malakozoologische Gesellschaft 1894:
13-42.
Solem, A. 1976. Endodontoid land snails from Pa-
cific Islands. Part I: Endodontidae. Field Mu-
seum of Natural History, Monograph, pp. 1—
508.
1983. Endodontoid land snails from Pacific
Islands. Part II. Families Punctidae and Char-
opidae, zoogeography. Field Museum of Natu-
ral History, Monograph, pp. 1-336.
Zilch, A. 1973. Die Typen und Typoide des Natur-
museums Senckenberg, 52. Mollusca: Hydro-
cenidae.— Archiv fiir Molluskenkunde 103:263-
DAD
(FGT) Florida State Museum, University
of Florida, Gainesville, Florida 32611;
(TMI) Bermuda Biological Station for Re-
search, Inc., Ferry Reach 1-15, Bermuda.
PROC. BIOL. SOC. WASH.
100(3), 1987, pp. 452-458
CLASSIFICATION OF THE ASCOTHORACIDA
(CRUSTACEA)
Mark J. Grygier
Abstract.—The order- and family-level classification of the maxillopodan
crustacean superorder Ascothoracida is revised in light of recent advances in
taxonomical and morphological knowledge of this group. It is divided into two
orders and six families, two of which have two subfamilies. New taxa proposed
are the orders Laurida and Dendrogastrida (which supersede the suborders
Lauroidida Wagin and Synagogoidida Wagin), the family Ascothoracidae, and |
the subfamilies Introcorniinae and Ulophysematinae.
The Ascothoracida consist of about 70
described species of parasites of echino-
derms and anthozoans. Their taxonomic
rank and relationship to other maxillopo-
dan crustaceans, especially to the Cirripe-
dia, have been interpreted differently (e.g.
Grygier 1983b, Boxshall 1983, Boxshall and
Lincoln 1983). Here they are considered a
superorder coordinate with the Cirripedia
and the Facetotecta (““Hansen’s y-larvae’’)
within the maxillopodan subclass Theco-
straca (classification of Grygier 1985a).
Wagin (1976) divided the Ascothoracida,
considered by him an order of the Ento-
mostraca, into two suborders: Lauroidida
for most of the anthozoan parasites; and
Synagogoidida primarily for the echino-
derm parasites, but Synagoga mira Nor-
man, the type of its genus, infests an anti-
patharian coral. Wagin’s arrangement of
Gruvel’s (1905) four families and their con-
tained genera follows:
Lauroidida—Lauridae: Laura, Bacca-
laureus, Gorgonolaureus
— Petrarcidae: Petrarca
Synagogoidida — Synagogidae: Synagoga,
Ascothorax, Parascotho-
rax
— Dendrogasteridae (sic):
Ulophysema, Dendrogas-
ter
Numerous new species and genera have
been described since 1980, mostly by the
present author, and the family Ctenosculli-
dae, previously thought to be molluscan,
has been transferred to the Ascothoracida
(Warén 1981, Grygier 1983d). After the
separation of the crinoid-infesting Wagi-
nella from Synagoga (Grygier 1983a), Wag-
in’s suborders could no longer logically be
used if the basic, apparently sound division
by host phylum were to be preserved. Syn-
agoga, now limited to anthozoan parasites,
may reasonably be transferred to the other
suborder, but both subordinal names would
then come to apply to the old Lauroidida,
and the remaining portion of the former
Synagogoidida would be left nameless. The
recent addition of several genera of gorgo-
nian parasites to the Synagogidae (Grygier
1981, 1984a; Moyse 1983; Lowry 1985)
makes the need for an ordinal revision more
pressing, and it has also exacerbated the
paraphyletic nature of the Synagogidae,
which was already evident in Wagin (1976)
(defined by possession of a generalized body
plan).
The present paper revises the ordinal and
familial classification of the Ascothoracida
in a manner consistent with current mor-
phological knowledge of the animals (Gry-
gier 1984b) and seeks, as far as possible, to
employ monophyletic taxa.
VOLUME 100, NUMBER 3
Table | is a character state matrix for the
genera of Ascothoracida described through
early 1986. A very high level of convergence
is evident, probably because most of the
apomorphic states are easily duplicated re-
ductions associated with parasitism. There-
fore, one of the preconditions for a mean-
ingful cladistic treatment, the assumption
of relatively rare convergence, is not met.
Despite this limitation, several apparently
monophyletic taxa have been identified.
However, the family Synagogidae, as de-
fined below, and thus also the order Laur-
ida, may still be paraphyletic due to: 1) the
near absence of synapomorphies between
Waginella and other genera, and 2) the pos-
sibility that one or more of the advanced
synagogid genera, such as Thalassomem-
bracis, form the sister group of the Lauridae.
Despite these faults, which may eventually
be correctable, I believe the following clas-
sification of the Ascothoracida to be an 1m-
provement over its predecessor, and, aside
from possible further subdivision of the
Synagogidae, it will serve to place numerous
undescribed species and genera now in
preparation.
Class Maxillopoda Dahl, 1956
Subclass Thecostraca Gruvel, 1905, sensu
Grygier (1985a)
Superorder Ascothoracida
Lacaze-Duthiers, 1880
Diagnosis. — Bivalved crustaceans (valves
often fused in females), diverticula of mid-
gut and gonads in carapace. Primitively with
11 free trunk segments, first 6 with bira-
mous thoracopods, seventh with biramous
or uniramous penis in both sexes, last with
movable furcal rami. Eyes usually absent.
Frontal filaments often present, sometimes
combined with uniramous antennae(?) into
sensory organs, better developed and plu-
mose in males. Antennules primitively
6-segmented, prehensile, with claw guard
and movable claw on sixth segment, setae
on fourth, fifth, and sixth. Antennae usually
453
absent. Labrum conical or pear-shaped,
sheathing other mouthparts to form oral
cone; maxillae usually with movable sub-
terminal hooks. Female gonopores and often
filamentary appendages at base of first legs.
Thoracopods primitively paddle-like and
setose, usually abutting medially, with
2-segmented exopods, 3-segmented endo-
pods (2-segmented in legs | and 6). Seminal
receptacles usually in coxae of legs 2—4 or
2-5. Furcal rami blade-like, unsegmented,
with various arrangements of terminal and
medial setae, primitively a setal fan. Com-
monly a pair of posteroventral telsonic
spines. Apparently gonochorists (Grygier
1987) or (Petrarcidae) simultaneous her-
maphrodites. Eggs and larvae usually
brooded under carapace; larvae include
nauplii and bivalved, “‘ascothoracid lar-
vae.”’ Parasites of echinoderms and antho-
zoans.
Remarks. — Features, especially of the ap-
pendages, are modified or reduced in var-
ious ways in different ascothoracidans as
indicated in the following diagnoses.
Order Laurida, new order
Diagnosis. — Parasites of Anthozoa (ex-
cept Waginella, ectoparasitic on crinoids).
Adult morphology highly variable.
Family Synagogidae Gruvel, 1905
Diagnosis. — Parasites of antipatharians,
octocorals, and stalked crinoids. Main body
and appendages in general plesiomorphic
(see above). Carapace bivalved or dorsally
fused and expanded into stiff-walled brood
chamber (latter correlated with vertical ce-
phalic attachment zone and unpaired dorsal
humps or horns on thorax). Fifth antennular
segment with several to many setae (but O-—
2 in Thalassomembracis); sixth with prox-
imal sensory process, usually with aesthe-
tasc and 3 setae (less well developed in
Thalassomembracis and Waginella). An-
tenna(?)-frontal filament complexes pres-
ent, best developed in males. Mandibles
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
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VOLUME 100, NUMBER 3
lanceolate or broad-based but attenuate,
with complex but variable medial arma-
ment (little armament in Waginella). Max-
illules blunt with one or more longitudinal
comb-rows of setae (styliform and unarmed
in Waginella). Maxillar tips variable. Hy-
popharynx produced into long languette (not
in Waginella). Filamentary appendages
present or not. Large epaulets on thoraco-
mere 6 (not in Synagoga). Details of penis,
telsonic spines, and furca variable.
List of genera. —Synagoga Norman, 1888;
Waginella Grygier, 1983a; Gorgonolaureus
Utinomi, 1962; Jsidascus Moyse, 1983;
Thalassomembracis Grygier, 1984a; Car-
domanica Lowry, 1985.
Remarks. — Waginella is included in this
family despite its echinoderm hosts (when
known) and its somewhat exceptional
mouthparts because it and Synagoga, the
two most generalized ascothoracidan gen-
era, are in most respects extremely similar
and until recently were classified in the same
genus (Grygier 1983a).
Family Lauridae Gruvel, 1905
Diagnosis. —Endoparasites or mesopara-
sites of zoanthids. Female carapace uni-
valved, greatly enlarged relative to body,
often with lateral pouches or coils, cephalic
attachment zone inverted; males bivalved,
much smaller than females. Antennules re-
duced in females, claw vestigial or absent;
male antennules generalized. Oral cone nor-
mal. Mandibles usually with medial hairs,
maxillules blunt, unarmed, maxillae gen-
eralized. Four to 6 pairs of uniramous, seg-
mented legs with short, spine-like setae in
females; 6 pairs in males, some biramous.
Female filamentary appendages oval, plate-
like (absent in Laura). Abdomen 4-seg-
mented, last segment sometimes showing
partial division. Penis uniramous (often very
long in males). Telsonic spines small or ab-
sent. Furcal rami with up to 4 terminal se-
tae, usually no medial setae.
List of genera. — Laura Lacaze-Duthiers,
1865; Baccalaureus Broch, 1929; Zoan-
455
thoecus Grygier, 1985b; Polymarsypus Gry-
gier, 1985b.
Family Petrarcidae Gruvel, 1905
Diagnosis. —Endoparasites of scleractin-
ian corals. Carapace of two thick valves
armed with spines and/or papillae, lacking
brood chamber. Antennules 5-segmented,
not subchelate, only distal segment well
armed. Antennae absent. Oral cone vari-
able; mandibles and maxillules with short,
medial cutting edge, rarely unarmed; distal
parts of maxillae reduced. Thoracopods
uniramous, with few or no setae, first pair
reduced or absent, no filamentary append-
ages. First abdominal segment much larger
than others. Penis and terminal abdominal
segment variable. Simultaneous hermaph-
rodites.
List of subfamilies. —Introcorniinae, new
subfamily; Petrarcinae Gruvel, 1905.
Subfamily Introcorniinae, new subfamily
Diagnosis. —Carapace spherical with pa-
pillae; cephalic attachment zone vertical.
Antennular cuticle of even thickness, distal
segment generalized. Labrum normal; max-
illae with hairy, vestigially bifid tips. Thorax
clearly segmented. Thoracopod 1 absent;
other 5 pairs segmented, weakly setose, some
with seminal receptacles. Abdomen 5-seg-
mented, penis in form of 2 blunt lobes. Fur-
cal rami well developed.
List of genera. —Introcornia Grygier,
1983d (type genus).
Subfamily Petrarcinae Gruvel, 1905
Diagnosis. —Carapace ovoid or dome-
shaped, with spines and/or papillae; ce-
phalic attachment zone dorsal, horizontal.
Basal antennular segments with distinct
sclerites; conspicuous armament of fifth
segment reduced to claw and claw guard
bearing a large seta. Labrum blunt, rear edges
widely separated; maxillae forming a short,
massive labium. Thoracic segmentation ob-
scure. Five or 6 pairs of unsegmented, un-
456
armed, usually lobular thoracopods, first
pair narrow or setiform when present. Sem-
inal receptacles in thorax. Extremely long
and robust penis with short rami or none.
No more than 4 abdominal segments, all
but first very reduced. Furcal rami vestigial
or absent.
List of genera. —Petrarca Fowler, 1889;
Zibrowia Grygier, 1985c.
Order Dendrogastrida, new order
Diagnosis. — Parasites of echinoderms.
Carapace valves in females at least partly
fused, soft-walled with delicate cuticle, often
greatly enlarged. Trunk variable, general-
ized to extremely reduced. Antennules 4- or
5-segmented and subchelate at some stage
of development, or vestigial; proximal sen-
sory process of terminal segment reduced
to isolated aesthetasc and seta, aesthetasc
ribbon-like in larvae (antennular details to
be confirmed in larval Ctenosculidae). An-
tennae absent. Mandibles and maxillules
unarmed, at least medially, when present.
Thoracopods leaf-like, uniramous, or ab-
sent.
Family Ascothoracidae, new family
Diagnosis.—Bursal parasites of ophiu-
roids. Carapace in females roughly spherical
or ovoid, valves partly fused with pair of
thin-walled, dorsal brood chambers; ce-
phalic attachment zone vertical; males bi-
valved, much smaller than females. Anten-
nules 5-segmented, subchelate, fourth
segment usually with toothed process. La-
brum normal. Mandibles setiform with dis-
tal hairs. Anterior thoracomeres usually
greatly swollen with bilateral protrusions.
First thoracopods short, uniramous; fila-
mentary appendages often present. Next 4
pairs of legs leaf-like, with short, rounded
rami (legs narrow in males, endopod re-
duced or absent); sixth legs short. Seminal
receptacles small, tubular, often only in legs
2-4. Abdomen 5-segmented, sharply bent
at segment 4; penis a short lobe, even in
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
males. Furcal rami elongate, usually with
short ventral setae in females, a few distal
setae in males.
List of genera.—Ascothorax Djakonov,
1914 (type genus); Parascothorax Wagin,
1964.
Family Ctenosculidae Thiele, 1925
Diagnosis. —Mesoparasites of starfish.
Ovoid carapace with short, posteroventral
or ventral aperture. Cephalic attachment
zone inverted. Antennules minute or ab-
sent. Front side of labrum short. Mandibles
apparently absent; maxillae bifid, but not
hooked. Thorax enlarged, with humps or
long dorsal horns. Thoracopods typically
leaf-like, sometimes uniramous; filamen-
tary appendages sometimes present. Ab-
domen 4-segmented, penis vestigial or ab-
sent. Furcal rami large, variable. Adult males
unknown.
List of genera. —Ctenosculum Heath,
1910; Endaster Grygier, 1985d.
Family Dendrogastridae Gruvel, 1905
(name corrected)
Diagnosis. — Female carapace (mantle)
produced into large, soft-walled lobes or
branches. Antennules 4-segmented, subche-
late at some stage of development. Man-
dibles and maxillules vestigial or absent.
Thoracopods short and uniramous, or ab-
sent; first pair always absent. No filamen-
tary appendages or seminal receptacles. Ab-
domen at most 3-segmented, often absent,
furcal rami present as unarmed lobes or ab-
sent.
List of subfamilies. —Ulophysematinae,
new subfamily; Dendrogastrinae Gruvel,
1905.
Subfamily Ulophysematinae,
new subfamily
Diagnosis. —Endoparasites of irregular
echinoids. Mantle with anterior and pos-
terior lobe and ventral aperture, cephalic
VOLUME 100, NUMBER 3
attachment zone inverted. Antennules in
adult poorly segmented, with small distal
segment and claw. Mouthparts absent ex-
cept for small labrum. Dorsal horns on first
4 thoracomeres. Four or 5 pairs of short,
uniramous limbs with short setae. Abdo-
men 2- to 3-segmented, with or without fixed
furcal lobes. Adult males unknown.
List of genera. — Ulophysema Brattstrom,
1936 (type genus).
Subfamily Dendrogastrinae Gruvel, 1905
Diagnosis. —Endoparasites of starfish.
Female mantle developed into pair of pos-
terior lobes or bilateral system of branches
with very small aperture. Cephalic attach-
ment zone anterodorsal. Males bivalved,
each valve with posterior protrusion; living
in female brood chamber. Antennules and
maxillae well developed in adults; third an-
tennular segment lacking process, but usu-
ally with 1 or 2 spine-like setae opposing
claw. Thorax and abdomen obsolete, sac-
like, limbs generally absent, no furca.
List of genera.—Dendrogaster Knipo-
vich, 1890 (subjective synonyms: Myriocla-
dus Okada, 1925 and probably Laocoon
Nierstrasz and Entz, 1922); Bifurgaster
Stone and Moyse, 1985.
Acknowledgments
This paper is based on work performed
as a graduate student of W. A. Newman at
Scripps Institution of Oceanography and as
an exchange fellow (American-Scandina-
vian Foundation) at the Zoological Central
Institute, University of Copenhagen. I thank
Dr. J. T. Hgeg (University of Copenhagen)
for his comments on the manuscript.
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Note added in proof
The new genus Paremedius Stone, 1987 (Journal of
Natural History 21:219-—224) belongs to the Dendro-
gastrinae. I regard it as a synonym of Bifurgaster, and
in all described features it agrees with that genus in
Table 1.
PROC. BIOL. SOC. WASH.
100(3), 1987, pp. 459-461
A NEW SUBSPECIES OF RALLINA EURIZONOIDES
(AVES: RALLIDAE) FROM THE BATAN
ISLANDS, PHILIPPINES
Robert S. Kennedy and Charles A. Ross
Abstract.—Specimens of the Slaty-legged Crake (Rallina eurizonoides) col-
lected on Batan Island, Philippines, in 1981 and 1985 are the first records of
the species for the Batan Islands and are here recognized as R. e. alvarezi, new
subspecies. The first three records of R. e. eurizonoides for Fuga Island in the
Babuyan Islands are also reported.
The Slaty-legged Crake (Rallina eurizo-
noides) ranges from India through South-
east Asia to the Greater Sundas, and from
the Ryukyu Islands of Japan south through
the Philippines to Celebes. Ripley (1977)
recognized six subspecies with nominate R.
e. eurizonoides restricted to the Philippines,
where duPont (1971) listed it from the is-
lands of Basilan, Bohol, Cagayan, Cebu,
Leyte, Luzon, Mindanao, Mindoro, Negros,
Panay, and Jolo. Recently, duPont and Ra-
bor (1973) have reported it from Sanga San-
ga in the Tawitawi Group, and Dickinson,
Kennedy, and Rozendaal (in prep.) have lo-
cated specimens from Guimaras, Marin-
duque, Samar, and Siquijor taken by the
1887-1888 Steere Expedition to the Phil-
ippines. It is also known from three previ-
ously unpublished specimen records from
Fuga Island in the Babuyan Group collected
for the Delaware Museum of Natural His-
tory on 16 Mar 1980 (DMNH 70273, 70275)
and 17 Mar 1980 (DMNH 70274).
During a visit to Batan Island in the Batan
Islands north of Luzon in the Philippines
on 31 Oct 1981, Kennedy collected a spec-
imen of R. eurizonoides (LSUMZ 105082,
female) that is the first record for the island.
Recent collections we made in the Batan
Islands in collaboration with the Philippine
National Museum and Silliman University
from 28 May to 11 Jun 1985 yielded ad-
ditional specimens of R. eurizonoides (4
skins: 2 adult males, | half-grown male and
1 adult female; and 1 complete and 1 partial
skeleton).
After comparing the Batan birds with
specimens of R. e. eurizonoides from other
islands in the Philippines and samples of
the other subspecies, particularly R. e. for-
mosana from Taiwan and R. e. minahasa
from Celebes, we have concluded that the
specimens from Batan represent a new sub-
species, to be known as:
Rallina eurizonoides alvarezi,
new subspecies
Holotype.—PNM 16301, adult male in
breeding condition (largest testis—7 x 14
mm), 31 May 1985, western slope of Mt.
Iraya, 180 m elevation, Sitio Nacamaya, 3
km NE of Basco, Batan Island, Batanes
Province, Philippines, R. S. Kennedy and
party (collector’s no. 1048).
Paratypes. —LSUMZ 105082 and USNM
5828 10—582812, Batan Island, 1.5 to 4 km
N or NE of Basco, R. S. Kennedy and party,
31 Oct 1981 (LSUMZ 105082) and 31 May
to 8 Jun 1985 (USNM 582810-5828 12).
Subspecies characters. — Differs from R. e.
eurizonoides and R. e. formosana by having:
the upperparts, and outer edges of wing and
tail feathers darker olive brown; the top and
sides of head, hind neck, lower throat and
breast darker chestnut; the white barring of
460
underparts narrower and less pronounced;
the throat, in males, chestnut, not pale ru-
fous as in R. e. eurizonoides or white as in
R. e. formosana; and the inner webs of wing
and tail feathers dusky brown almost black.
Differs further from R. e. formosana in hav-
ing a longer tail (R. e. alvarezi (4) 66.9 +
1.31, range 65.5-68.0; R. e. formosana (2)
60.5 + 4.95, range 57.0—64.0), shorter cul-
men (from feathers on side of culmen: R. e.
alvarezi (4) 20.6 + 0.46, range 20.4—21.2;
R. e. formosana (2) 24.4 + 1.13, range 23.6—
25.2), and a shorter tarsus (R. e. alvarezi (5)
43.3 + 0.85, range 42.2-44.0; R. e. for-
mosana (2) 44.5 + 0.85, range 43.9-45.1).
Like R. e. minahasa in being of similar
size, in having dark olive brown upperparts
and in having narrow and less pronounced
white barring on the underparts. Differs from
R. e. minahasa in having the top and sides
of head, hind neck, lower throat and breast
darker chestnut and the throat, in males,
chestnut, not pale rufous.
Rallina e. alvarezi differs from the other
subspecies by size and color of the head and
neck (R. e. sepiaria from the Ryukyu Is-
lands), and by the narrow white barring of
the underparts and throat color of the male
(R. e. amauroptera from India; and R. e.
telmatophila from mainland South-east
Asia). Females are like males except that
they have pale rufous throats like R. e. eu-
rizonoides and R. e. minahasa.
Soft parts. —Iris dark orange in males and
bright red with a faint inner ring of orange
in females; eye ring orange; upper mandible
dark gray, almost black, with blue-green to
green base; lower mandible with blue-green
to green proximally turning to gray distally;
legs and feet gray.
Description of juvenile. -USNM 582812,
about 3—4 weeks old; forehead, superciliary
line and malar region with black down
feathers; crown, neck, upper back and wings
blackish gray with a dark olive brown wash;
lower back and rump blackish neutral gray;
throat dark neutral gray; underparts sooty
gray with a faint olive brown wash and with
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
some white flecks on the breast and belly.
Iris dark olive brown; bill and legs black.
Measurements. — Wing chord (4) 131.4 +
6.61, range 123.2—137.7; see above for other
measurements. Weight (1 6) 128 g, (1 2 with
fully developed shelled egg in the oviduct)
180 g.
Range. —Batan Island. Kennedy and A.
Fidel heard this species calling on Sabtang
Island in the Batan Islands. The species was
not encountered on nearby Ivojos Island and
is unknown to the inhabitants.
Status. — Fairly common but extremely
secretive in forest and second growth. All
specimens were located and flushed from
the forest floor by a dog. Known in the Iva-
tan dialect as Adongong.
Etymology. —Named in honor of Jesus B.
Alvarez, Jr., for his untiring efforts to con-
serve Philippine fauna and flora.
Remarks.—Of the three known speci-
mens from Fuga, two clearly resemble R. e.
eurizonoides. The third (DMNH 70274) has
the white barring on the belly as in R. e.
eurizonoides but the olive brown of the up-
perparts is nearly as dark as in R. e. alvarezi
and darker than in any of the specimens of
R. e. eurizonoides we examined. Neverthe-
less, the Fuga birds, although tending to-
ward R. e. alvarezi, appear to be closer to
R. e. eurizonoides.
The relationships of R. e. alvarezi are not
clear. It is most similar to R. e. minahasa
of Celebes and least similar to R. e. for-
mosana and the other races of R. eurizo-
noides owing to its more richly colored up-
perparts and narrower abdominal barring.
Specimens examined.—R. e. alvarezi: |
(LSUMZ), 1 (PNM), 3 (USNM). R. e.
amauroptera: 4 (USNM). R. e. eurizo-
noides: Bohol, 1 (FMNH); Cebu, 3 (DMNH);
Fuga, 3 (DMNH); Leyte, 2 (USNM); Luzon,
57 (DMNH), 3 (FMNH); Mindanao, 2
(USNM); Mindoro, 1 (FMNH); Negros, 1
(FMNH); Panay, 1 (USNM); Sanga Sanga,
Tawitawi, 3 (DMNH); Siquijor, 2 (USNM).
R. e. formosana: 3 (USNM). R. e. mina-
hasa: 3 (AMNH), | (USNM). R. e. sepiaria:
VOLUME 100, NUMBER 3
2 (AMNH), 1 (type) (USNM). R. e. telma-
tophila: 2 (USNM).
Acknowledgments
We thank the curators of the Delaware
Museum of Natural History (DMNH), Field
Museum of Natural History (FMNH), Lou-
isiana State University Museum of Zoology
(LSUMZ), National Museum of Natural
History (USNM), and Philippine National
Museum (PNM) for lending specimens and/
or for permission to study specimens in their
care. We are grateful to P. Gonzales, L.
Gonzales, and J. Alvarez, Jr., for their help
and cooperation in granting collecting per-
mits and to A. Alcala, A. Fidel, G. Fidel,
B. Gargar, and V. Samarita for help col-
lecting in Batanes. Without the assistance
of Philippine Airlines, field work in the Ba-
tan Islands would have been impossible.
Partial support for our field work was sup-
plied by the Smithsonian Institution. E.
Dickinson and K. Parkes kindly reviewed
drafts of this manuscript.
Literature Cited
Dickinson, E. C., R. S. Kennedy, and F. G. Rozendaal.
[In prep.] Notes on the birds collected in the
Philippines by the Steere Expedition of 1887/
1888.
duPont, J.E. 1971. Philippine birds.— Delaware Mu-
seum of Natural History Monograph 2:x + 1-
480.
, and D. S. Rabor. 1973. South Sulu Archi-
pelago birds: An expedition report.— Nemouria
9:1-63.
Ripley, S.D. 1977. Rails of the world. A monograph
of the family Rallidae. Boston, David R. Go-
dine, Publisher. 406 pp.
(RSK) Department of Zoology, Washing-
ton State University, Pullman, Washington
99164-4220 (Present address: Cincinnati
Museum of Natural History, 1720 Gilbert
Avenue, Cincinnati, Ohio 45202). (CAR)
Department of Vertebrate Zoology, Nation-
al Museum of Natural History, Smithson-
ian Institution, Washington, D.C. 20560.
PROC. BIOL. SOC. WASH.
100(3), 1987, pp. 462-464
TWO WOODPECKERS FROM THE LATE PLIOCENE
OF NORTH AMERICA
Alan Feduccia
Abstract.— Three bones from the late Pliocene Rexroad Formation, Meade
County, Kansas, represent two species of woodpeckers, a large species com-
parable in size to the species of the ivory-billed group (genus Campephilus),
but not of that genus, and a flicker, similar in size to modern Colaptes auratus.
Because of the extreme homogeneity in the postcranial bones of the Picidae
these bones are not assigned to a new species.
Although woodpecker fossils are known
during the Pleistocene, especially in North
America (Brodkorb 1971a), the pre-Qua-
ternary fossil record of the Picidae is ex-
tremely sparse, and is currently represented
by only three valid species. Pliopicus brod-
korbi (Feduccia and Wilson 1967) and Pa-
laeonerpes shorti (Cracraft and Morony
1969), both known from the Ogallala Group
in Kansas and Nebraska, respectively, are
moderate-sized woodpeckers, possibly re-
lated to the melanerpine assemblage. Cam-
pephilus dalquesti (Brodkorb 1971b) is from
the late Pliocene (Blancan) of Texas, and
represents a large woodpecker, comparable
in size to modern species of the ivory-billed
group, genus Campephilus. Because of the
extreme morphological homogeneity of the
postcranial skeletons of the Picidae these
generic allocations may be questioned; how-
ever, there seems at least little doubt that
the abovementioned forms represent three
species of Pliocene woodpeckers, one of large
size. In addition to the abovementioned
forms, Wetmore (1931) reported a fossil
woodpecker, represented by a nearly com-
plete right ulna, of flicker-like affinities in
the Lower Phocene Devil’s Gulch Beds
(Blancan), near Ainsworth, Nebraska. Beck-
er (1986), likewise, reported a species of
flicker (Colaptes), larger than modern C. au-
ratus from the Oreana local fauna (Blancan),
of Owyhee County, Idaho, which correlates
in age with the well-known Hagerman local
fauna, thought to be slightly younger than
the fossils reported herein from Rexroad
Formation (see Feduccia 1975).
In their 1969 paper describing Palae-
onerpes shorti, Cracraft and Morony discuss
its relationships through a laborious four
page section (pp. 4-7), giving detailed ge-
neric analyses as well as comparisons with
fossils that were not actually examined, but
seen only in published figures. I can now
confirm few, if any, of their assertions con-
cerning the generic distinctiveness of Pa-
laeonerpes, and it seems possible that P.
shorti belongs in the genus Melanerpes.
Likewise, the generic distinctiveness of
Pliopicus brodkorbi could also be ques-
tioned. Feduccia and Wilson (1967:4) stated
that ‘““Pliopicus seems to be most closely al-
lied to the genus Melanerpes, ...”’ How-
ever, I recommend leaving the above genera
until a complete revision of the family is
undertaken.
In addition to Palaeonerpes shorti and
Pliopicus brodkorbi, Olson (1985) reported
having examined fossils of a medium-sized
woodpecker from the middle Miocene (late
Barstovian) of New Mexico, which would
represent the earliest known occurrence of
the family, and Ballmann (1976) assigned
the proximal end of an ulna from the late
Miocene of Italy to the Picidae.
Two of the fossils reported here (Fig. 1)
VOLUME 100, NUMBER 3
are a left ulna (University of Michigan Mu-
seum of Paleontology no. 31711), and a left
carpometacarpus (UMMP 24754), both
from the Wendell Fox Pasture locality (UM-
K3-53) (see Feduccia 1975), and represent
a large woodpecker similar in size to species
of the genus Campephilus. Although the ulna
has no distinctive characters to ally it with
any specific genus of large woodpecker, it is
nevertheless not assignable to Campephilus,
as it lacks the strong inward curve of the
proximal end of the ulna found in the genus.
In size the fossil most closely approximates
Campephilus melanoleucus of all the forms
examined. The fossil ulna measures 52.8
mm in total length; C. melanoleucus, n =
6, was 49.8-54.2 (mean = 52.1). The car-
pometacarpus measures 25.0 in total length;
C. melanoleucus, n = 6, is 24.0-—27.2
(mean = 25.4).
A third fossil element is the distal end of
a right tarsometatarsus (UMMP 31881,
from the Fox Canyon locality, UM-K1-47),
that agrees very closely with the genus Co-
laptes. The tarsometatarsus of Colaptes is
distinguished by lacking any substantial ex-
cavation of the trochlea for digit III when
compared with that of other woodpeckers
having a robust trochlea metatarsi tertii.
This character presumably reflects the ter-
restrial habits of the genus. The greatest
width at the level of the distal foramen is
2.9, for Colaptes auratus, n = 6, the same
measurement is 2.5—3.2 (mean = 2.8). The
specimen of Colaptes described by Becker
(1986) is slightly younger than this fossil
and slightly larger than living C. auratus.
However, there 1s little to preclude the pos-
sibility that the two represent the same
species or that both represent a Blancan form
of modern C. auratus.
Two woodpeckers, one certainly different
from those currently known in the fossil rec-
ord, are represented by the fossils described
herein. These fossils, together with those
previously described, may possibly indicate
a greater diversity of woodpeckers during
the Pliocene of North America. Pending a
463
A, Left ulna, 52.8 mm, anconal view, B,
Fig. 1.
Left carpometacarpus, 25.0 mm, external view, of a
large Pliocene woodpecker. C, Distal end of right tar-
sometatarsus, 13.6 mm, anterior view, of Pliocene Co-
laptes sp. Photo by Susan Whitfield.
complete review of the osteology of the Pi-
cidae, it seems most prudent not to assign
new names to these fossils at present. They
point, however, to the need for a thorough
study of the postcranial osteology of the
woodpeckers, and suggest the possibility that
the living genera are oversplit.
Acknowledgments
I am grateful to Storrs L. Olson, Depart-
ment of Vertebrate Zoology, National Mu-
seum of Natural History, for advice and for
providing working facilities in the Division
of Birds. The late Claude W. Hibbard of the
Museum of Paleontology, University of
464
Michigan, placed these fossils at my dis-
posal.
Literature Cited
Ballmann, P. 1976. Fossil Vogel aus dem Neogen der
Halbinsel Gargano (Italien). Zweiter Teil.—
Scripta Geologica 38:1—59.
Becker, J. J. 1986. Fossil birds of the Orcana local
fauna (Blancan), Owyhee County, Idaho. — Great
Basin Naturalist 46(4):763-768.
Brodkorb, P. 1971la. Catalogue of fossil birds. Part 4
(Columbiformes through Piciformes).— Bulle-
tin of the Florida State Museum, Biological Sci-
ences 15:163-—266.
1971b. The paleospecies of woodpeckers. —
Quarterly Journal of the Florida Academy of
Sciences 33:132-136.
Cracraft, J., and J. J. Morony, Jr. 1969. A new Plio-
cene woodpecker, with comments on the fossil
Picidae.— American Museum Novitates 2400:
1-8.
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Feduccia, A. 1975. Professor Hibbard’s fossil birds.
Studies on Cenozoic paleontology and stratig-
raphy in honor of Claude W. Hibbard, Vol. 3.—
Museum of Paleontology, University of Mich-
igan, Papers on Paleontology 12:67—70.
, and R. L. Wilson. 1967. Avian fossils from
the Lower Pliocene of Kansas.— Occasional Pa-
pers of the Museum of Zoology, University of
Michigan 655:1-6.
Olson, S. L. 1985. The fossil record of birds. Pp. 79-
238 in D.S. Farner, J. R. King, and K. C. Parkes,
eds., Avian biology, volume 8. Academic Press,
New York and London.
Wetmore, A. 1931. Record of an unknown wood-
pecker from the Lower Pliocene.—Condor 33:
255-256.
Department of Biology, University of
North Carolina, Chapel Hill, North Caro-
lina 27514.
PROC. BIOL. SOC. WASH.
100(3), 1987, pp. 465-479
GUITARRA ABBOTTI AND G. ISABELLAE, NEW
SPONGES FROM THE EASTERN PACIFIC
Welton L. Lee
Abstract.—Two new species of the genus Guitarra are reported from the
eastern Pacific. One of these, G. abbotti, was collected from a submerged sea-
mount, the Cordell Bank, off the central California coast at 35-46 m. The
second, G. isabellae, was collected off Albemarle Island in the Galapagos Islands
at 82.3 m. A new spicule type, the biplacochela, is described from both species
and the nature of the microscleres has been shown to differ significantly from
those described from other Guitarra species.
Recent collections obtained through the
auspices of the Cordell Bank Expeditions
and deposited at the California Academy of
Sciences have brought to light several new
and unusual sponge species as well as range
extensions. Among those sponges new to
science is the first Guitarra species to be
reported from the eastern Pacific Ocean. In
the process of looking for comparative ma-
terial a second new species of this genus was
uncovered in the collections of the Allan
Hancock Foundation, University of South-
ern California. Dr. G. J. Bakus kindly pro-
vided for description the single specimen
that was found in collections made in the
Galapagos Islands by the Foundation in
1934. These two new species are described
below.
Guitarra Carter, 1874
Guitarra abbotti, new species
Figs. 1-13
Holotype and type locality. —CASIZ
060483 (Lee No. 34), Cordell Bank (Sta No.
821009), Marin County, California.
Paratypes. —BMNH 1986:5:7:2, BMNH
1986:5:7:2a (Lee No. 64), BMNH 1965:5:
7:3 (Lee No. 0-2), USNM 34058 (Lee No.
66), USNM 34059 (Lee No. 69), CASIZ
057844 (Lee No. 20), CASIZ 057843 (Lee
No. 0-1), CASIZ 057841 (Lee No. 2), and
CASIZ 057852 (Lee No. 29).
Distribution. — Pacific Coast of North
America, Cordell Bank, Marin County, Cal-
ifornia. Depth 35—46 m.
Local occurrence. No. 34, No. 20-Cor-
dell Bank, Marin County, 38°01.8'N,
123°25.1’W), 9 Oct 1982, 34.7—42.0 m; No.
2-Cordell Bank, Marin County, 37°59.1'N,
123°25.5’'W), 14 Sep 1980, 42—45.7 m; No.
0-1, No. 0O—2-Cordell Bank, Marin County,
38°01.8'N, 123°25.1'W, 10 Oct 1981, 34.7-
40.2 m; No. 69-Cordell Bank, Marin Coun-
ty, 38°01.8’N, 123°25.7'W, 15 Dec 1981,
40.2—43.9 m; No. 29-Cordell Bank, Marin
County, 38°01.8’N, 123°25.1'W, 10 Oct
1982, 34.7-42.1 m; No. 64, No. 66-Cor-
dell Bank, Marin County, 38°01.8'N,
123°25.1'W, 23 Oct 1982, 34.7—42.1 m.
Description. —Guitarra abbotti is an en-
crusting to massive, subtidal species. Form
ovoid or pulvinate when small to massive,
digitate and irregular in larger specimens
(Fig. 1). Larger specimens frequently lumpy
in appearance due to infolding of the surface
to form deep furrows which delineate rel-
atively smooth and gently rounded lobes,
the latter often broadly flattened as a result
of the invasion of the grooves beneath. These
lobes are closely appressed to the colony so
as not to disrupt its overall shape. The
smallest specimen (CASIZ 057854) mea-
sures 2.8 < 3.8 x 4.7 cm and the largest
(CASIZ 057843) 8.0 < 3.5 X 6.5 cm.
Color in life dark chocolate to reddish
466
a
Fig. 1. Guitarra abbotti, specimen No. WL-34 (ho-
lotype). Scale: life-size.
brown. Specimens in alcohol beige, often
mottled with light to dark gray. In a few
specimens the gray may dominate, espe-
cially on the extremities of the lobes.
Body surface superficially smooth with the
exception of deep grooves which isolate the
round or oblong lobes described above.
These lobes are found more frequently on
the larger specimens. Surface may vary
greatly (Fig. 2a, b). It can be smooth to mi-
nutely granular, the granular appearance due
to closely spaced and slightly raised spicule
brushes. Commonly smooth, minutely
raised ridges may be seen here as well. Else-
where the spicule brushes may be distinctly
elevated and more hispid with very shallow
and somewhat wide grooves isolating nar-
row plateaus or ridges containing rows of
these raised spicule brushes. Finally the
spicule brushes may form distinct opaque
conules, often coalescing to form ridges di-
vided by shallow to deep furrows. While all
conditions may be found in a single speci-
men, the smooth, granular appearance is
more frequently seen on smaller specimens,
whereas conules and deep grooves are more
characteristic of larger specimens. The
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
smoother surface areas possibly represent
portions of the sponge where the thin sur-
face membrane is still intact. However, the
surface is always minutely hispid, this
showing best when the specimen has been
dried. The spicules seen are almost always
derived from the spicule brushes on the con-
ules or ridges. The surface feels smooth to
the touch and the consistency is firm and
somewhat elastic.
Oscula (Fig. 3) often difficult to locate,
tending to be found in clusters, never evenly
spaced. Oval to oblong or diamond-shaped
with their opening commonly measuring
from 1.0-3.5 mm. Larger oscula may reach
6.0 mm. The openings frequently surround-
ed by a collar 0.25-—1.0 mm wide, with a
noticeably smoother surface than the rest of
the body, lighter in color and often very
slightly raised to 1.0 mm. Collar sometimes
flared. Ostia not seen.
Dermal membrane 4.0—6.0 um thick and
difficult if not impossible to see in all but a
few areas. It is most readily seen on the
smooth rim of the oscula where it is some-
what protected and not penetrated by spic-
ules. In most other areas it is either obscured
by dense spicules or eroded away.
Dermal membrane penetrated by styles
in the form of spicule brushes made up of
columns of 20-30 spicules (Fig. 4). Usually
7-20 spicules actually penetrate the surface,
these extending 120-200 um above the sur-
face. Surface showing a more or less regular
pattern of these columns, spaced 180-242
um apart. Penetrating spicules erect in the
center of the bundle and splayed to as much
as 45° at the edge. The result is to form a
spicule network at the surface that traps
considerable debris and foreign spicules.
The columns act as structural support for
the surface ridges and from above can be
seen to be connected by more or less loose
spicule bundles of megascleres, 3—6 per bun-
—
Fig. 2. Surface features of Guitarra abbotti, specimen No. WL-34 (holotype): A, Smooth surface area at edge
of surface furrow; B, Rough surface area. Scale: 10x.
VOLUME 100, NUMBER 3 467
468
a OE: A tes me
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig. 3. Osculum of Guitarra abbotti, specimen No. WL-34 (holotype). Scale: 5 x.
dle. These bundles as well as individual
scattered megascleres form a loose surface
network which is best seen in tangential sec-
tion (Fig. 5). Cross sections usually do not
show this network well because spicules close
to the surface other than those in the col-
umns tend to be disrupted by sectioning.
As noted above, each column is formed
by 20-30 or more spicules, these being dif-
ficult to count because they are so closely
packed. These spicules are derived in part
Table 1.—Spicule measurements (yu).
from 2 or more secondary bundles of 8-14
or more spicules which intersect from below
at about a 45° angle, these deriving from
the choanosome. The distance from the sur-
face to the point at which these bundles in-
terdigitate to form the ectosomal columns
is 400-600 um. Biplacochelae, placochelae
and “bipocillae’’ are abundant in the in-
terstices.
Choanosome (Fig. 4) consisting ofa fairly
regular isotropic reticulation of bundles of
, Megasclera Placochela
Species
(type) WL No. Length Range Width Large Range
G. abbotti
Holotype WL-34 341.05 + 38.51 259-379 7.50 + 1.80 86.65 + 7.15 74.1-97.6
Paratype WL-O1 330.12 + 38.19 279-382 Toa 22 NOS Soo (8) ae S55) 7/ 78.2-91.5
Paratype WL-02 338.88 + 36.54 278-396 Yoox 26 S573 81.61 + 7.76 64.2-97.2
Paratype WL-69 317.37 + 31.80 268-349 U3. 22 57 SMS) BE Sotho 69.1-96.2
Paratype WL-64 324.75 + 19.49 250-365 7.90 + 1.46 82.35 + 4.66 75.9-91.4
Paratype WL-66 338.48 + 20.13 254-374 Ul se AM 83.36 + 5.92 70.7--91.6
Paratype WL-20 323.96 + 21.65 249-362 UPD) ae Nye 83.18 + 2.43 80.4-88.1
Paratype WL- 2 321.44 + 16.06 272-374 7.48 + 1.65 80.55 + 9.39 65.7-93.3
Paratype WL-29 339.07 + 19.48 277-377 7.62 + 1.80 84.09 + 6.63 74.6-100.1
G. isabella
Holotype WL-AHF 354.45 + 46.58 224-406 VASA ae NWS 106 + 10.66 72.3-122.9
VOLUME 100, NUMBER 3
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Pr Lae =
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=—
RS
Pals Dy
* eA
Fig. 4. Skeletal architecture of Guitarra abbotti as
seen in cross-section.
megascleres, 2—10 spicules thick, which in-
tersect at intervals of 200-300 um. In cross-
section these areas appear as distinct nodes.
Added to this reticulate network are nu-
Table 1.—Continued.
469
: y
| Nok
b aX! d \
EA KS
aN)
ae
=
ABR
SS Sis
——_>
CLL A)
PAY ANY ES
~
SSS
‘
i
M/
Laat
1U1A
(4) Wty ‘Tunsae 1
Z G © : (A) Hy NAT OIA
I SitmSre 6C VV-CT? I I I I SUSUTIDIdS WIOpPO|]
6C oe OC ev cal OC 9°61 Oe? SNIADINOS 1U1A
I I I I I I I I BO BATH “7 ON Joqoys enyojeury
VE tv OFF VS Il GO OCC 9ST 1O1AIpiA 1U14
C ¢ el [L 56 6 9 v
6€ 8 S-0'S 60-TH SOs Y) vC-O'C ONEAS GC ¢°6C-0 LZ Vac Cm cele ByNH Pf) “eg oueH
6€ es CO+9P v9 OU aE GG 9 187¢ VCE 10jOUIS 1Uu14
SI[E19IE| sijeIpou sijeIpow YIpIM yeys Jo yeys Jo Sd JO dd dd
sn[Apuoo sn[Apuoo snjApuoo [e1siq yidap UIpIM 01 4O JO qd AO JO Jd
jo yidaq jo yidaq JO 14319 \svoT \svoT 7y13uaT :y3uaT
‘UMOUYUN X9s = (-) ‘O[VW = JY ‘o[eUIDJ = J ‘snourpuajeidng suog = Sq ‘pug [ewIxolg = qd ‘pug [eisiq = Aq ‘suenqiy e1suD = 4O ‘o10W IO
OI JO safdures 10J papraoid st uONeIASp plepuLys ‘ozis a[dures pue ‘osues ‘uUvoW YIM “(WI UT) sjoLIed pojoaTaS Jo sNSIeJOIQN dy} JO SJUDWIOINSeII|— "7 IQR
526
Discussion
Systematics and osteology. —The large size
of Vini vidivici and especially of V. sinotoi
suggested at first that these forms may be-
long in Cyanoramphus, the only other genus
of psittacid recorded from eastern Polyne-
sia. For this reason, more detailed osteo-
logical comparisons were made between Vini
and Cyanoramphus, as well as the geograph-
ically next closest genus of parrot, Prosopeia
of Fiji and Tonga (see “Systematic Paleon-
tology’). These comparisons show that the
fossils clearly pertain to Viniand are distinct
from Cyanoramphus or Propsopeia. Char-
mosyna, a polytypic genus that occurs from
Fiji west through the New Guinean region,
is also distinct from Vini, at least in the
single species of Charmosyna available (see
‘““Generic Diagnosis’’). Our comparisons for
Cyanoramphus are based upon C. novae-
zelandiae of New Zealand, as the only known
skeleton of C. zealandicus of Tahiti (re-
ported by Schauenberg 1970) was unavail-
able. Similarly, the comparisons for Char-
mosyna are based upon C. placentis of the
Moluccas rather than C. amabilis of Fiji, for
which no material is available.
The monotypic genus Phigys accommo-
dates the species solitarius of the Fijian Ar-
chipelago. We found no characters in the
tibiotarsus or tarsometatarsus to distinguish
Phigys from any species of Vini. The plum-
age of Phigys also resembles that of Vini in
both color and pattern (see duPont 1976:
plate 18, or Forshaw 1977:72-73). There-
fore we recommend synonymizing the ge-
nus Phigys G. R. Gray, 1870, with Vini Les-
son, 1831.
Although Vini is distinct osteologically
from other genera of parrots, the species
within Vini are rather uniform osteologi-
cally, with differences in size and occasion-
ally in proportions being the only diagnostic
features at the species level. We had no skel-
eton of V. stepheni, a species known only
from Henderson Island. The tarsal mea-
surements of skins of V. stepheni (16-18
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
mm, n = 8) resemble those of V. kuhlii (1 7—
18.5 mm, n = 25; Holyoak and Thibault
1984:123-124), so we are confident that V.
stepheni is much smaller than either V. sin-
otoi or V. vidivicl.
The parrot bones from the archeological
sites on Ua Huka, Hiva Oa, and Tahuata
permit an assessment of species-level rela-
tionships, although they are not adequate
for analysis of intraspecific variation. The
limited sample suggests that there may be
interisland differences in size, with both V.
sinotoi and V. vidivici being larger on Hiva
Oa and Tahuata than on Ua Huka (Table
1).
Extinction and biogeography. —The
stratigraphic association of bones of extinct
parrots with cultural materials shows that
these species survived until after the arrival
of Polynesians. A brief review of the chro-
nology and stratigraphy of the archeological
sites permits an assessment of the time of
extinction. A discussion of the causes of ex-
tinction of parrots and other birds in the
Marquesas will be presented in a future pub-
lication.
The Hane Site on Ua Huka, known also
as the Hane Dune Site, has provided much
of the basis for the four proposed cultural
phases of the Marquesas (Sinoto 1966, 1970,
1979). The Hane Site records the earliest
known aspects (Phase 1, Initial Contact) of
Polynesian culture in the Marquesas (begin-
ning about A.D. 300 according to Sinoto,
or about 200 B.C. or earlier according to
Kirch 1986). In his analysis of the faunal
material from Hane and other sites in the
northern Marquesas, Kirch (1973: see es-
pecially pp. 27-37) reported an abundance
of bird bones (not further identified) in the
strata of Phase 1, with very few bird bones
in the strata that represent Phase 2, (A.D.
600-1300), and none in the strata of Phase
3 (A.D. 1300-1600) or Phase 4 (A.D. 1600-
1800). Phase I of Marquesan culture is rep-
resented by Layers III—V of Area A and Lay-
er VI of Area B (Kirch 1973:27). The bones
of extinct parrots, as well as those of other
VOLUME 100, NUMBER 3
extinct birds yet to be described, occur only
in stratigraphic Layers II through V of Area
A and Layers V and VI of Area B. Thus
nearly all of the parrot bones are from Phase
I Layers, although a few bones of each of
the three species occur in strata assigned to
Layer II of Area A and Layer V of Area B,
both of which represent Phase 2 of Mar-
quesan culture. Unfortunately, precise
stratigraphic data were lost for some (less
than 5%) of the Hane bird bones sometime
between 1964 and 1984. Nevertheless, the
data at hand indicate that the extinction of
most landbirds on Ua Huka, including the
parrots, occurred within the first millenium
of human settlement.
The parrot bones from the Hanatekua
Shelter No. 2 Site, Hiva Oa, are from pre-
occupational levels (Y. H. Sinoto, pers.
comm.). There is no radiocarbon chronol-
ogy for this site (Bellwood 1972).
From the Hanamiai Site, Tahuata, the
specimens of Vini sinotoi and V. vidivici are
from Layers G/H and H, whereas those of
V. ultramarina are from Layers D, F, and
I. “I is the lowermost Layer in this well
stratified site. Radiocarbon ages (in years
B.P.) on charcoal for the Hanamiai Site are
130 + 100 (Layer B), 620 + 90 (Layer F),
and 850 + 60 and 1250 + 100 (Layer G)
(B. Rolett, pers. comm.). Additional radio-
carbon ages are pending. The stratigraphic
distribution of parrots at Hanamiai suggests
that V. ultramarina survived longer on Ta-
huata than its larger, extinct congeners, al-
though all three co-existed at one time. The
possible longer survival of V. ultramarina
on Tahuata is in accord with the fact that
this small species survived into historic
times on two islands in the Marquesas (Nuku
Hiva, Ua Pou), whereas the two larger
species of Vini apparently became extinct
before European contact.
The historical distribution of species of
Vini outside of the Marquesas suggests that
certain of these species as well have not sur-
vived on all of the islands that once made
up their natural ranges. For example, a fossil
All
sternum from a cave on Mangaia, southern
Cook Islands, has been referred to Vini cf.
V. kuhlii (Steadman 1985), a species pre-
viously known only from Rimatara and
Tubuai, 480 km east-southeast of Mangaia.
In light of the historic or prehistoric extir-
pations from certain islands of V. kuhlii, V.
ultramarina, V. australis, and V. peruviana,
and the complete losses of V. sinotoi and V.
vidivici, we might expect additional unde-
scribed species of Vini to show up in future
archeological or paleontological excava-
tions in Oceania. At a minimum, we can
expect new island records for the known
species. One or more species of Vini prob-
ably once existed throughout the many is-
lands of Polynesia that are not inhabited by
parrots today.
Acknowledgments
For access to collections and numerous
other courtesies, we thank Yosihiko Sinoto,
Toni Han, Elaine Jourdane, Patrick McCoy,
and Aki Sinoto of the Department of An-
thropology, BPBM. Carla Kishinami, Gail
Wine, and Allen Allison of the Department
of Vertebrate Zoology, BPBM, provided a
system for cataloguing the bones. Barry Ro-
lett allowed access to his recently collected
bones from Tahuata. For use of compara-
tive skeletons, we thank the Division of
Birds, USNM VJ. Phillip Angle, Storrs Ol-
son, Richard Zusi), the Department of Or-
nithology, AMNH (Mary Currie, Wesley
Lanyon), and the Ornithology Collection,
UCMVZ (Anne Jacobberger, Ned John-
son). William Birnbryer and Karen LaPorta
helped to sort a portion of the bones from
Hane. Partial financial support for the 1985
trip to BPBM was provided by the Smith-
sonian Associates Travel Program and by
funds generously donated to the Division
of Birds, Smithsonian Institution, by Mrs.
Alexander Wetmore. Douglas Pendleton
provided housing during the 1986 trip to
BPBM. We thank Storrs Olson for the idea
to name one of the species in honor of a
528
famous Roman quotation. Leslie Overstreet
kindly provided bibliographic assistance.
For comments on the manuscript, we are
indebted to Thomas Dye, Norton Miller,
Storrs Olson, Barry Rolett, and Yosihiko
Sinoto. Thaddeus Beblowski made the pho-
tographs. This research was supported in
part by National Science Foundation Grant
BSR-8607535. Contribution number 509 of
the New York State Museum and Science
Service.
Literature Cited
Baumel, J. J., A. S. King, A. M. Lukas, J. E. Breazile,
and H. E. Evans (eds.). 1979. Nomina Ana-
tomica Avium. Academic Press, London, 664
pp.
Bellwood, P. 1972. Asettlement pattern survey, Han-
atekua Valley, Hiva Oa, Marquesas Islands.—
Pacific Anthropological Records 17:1—50.
duPont, J. E. 1976. South Pacific birds.— Delaware
Museum of Natural History Monograph Series
3:1-218.
Evans, B. 1968. Dictionary of quotations. Delacourte
Press, New York, 2029 pp.
Forshaw, J. M. 1977. Parrots of the world. T.F.H.
Publications, Neptune City, New Jersey, 584 pp.
Holyoak, D. T., and J.-C. Thibault. 1984. Contri-
bution a l’étude des oiseaux de Polynesie ori-
entale.—Memoires du Muséum National
D’Histoire Naturelle, Serie A, Zoologie, 127:1-
209.
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Kirch, P. V. 1973. Prehistoric subsistence patterns
in the northern Marquesas Islands, French Pol-
ynesia.— Archaeology and Physical Anthropol-
ogy in Oceania 8:24—40.
Kirch, P. V. 1986. Rethinking East Polynesian pre-
history.—Journal of the Polynesian Society 95:
9-40.
Peters, J. L. 1937. Check-list of birds of the world,
volume 3. Harvard University Press, Cam-
bridge, Massachusetts, 311 pp.
Schauenberg, P. 1970. Note sur une perruche eteinte
de Tahiti Cyanoramphus zealandicus (Latham
1781) conservée au Muséum de Genéve.— Ar-
chives des Sciences (Genéve) 22:645-649.
Sinoto, Y. S. 1966. A tentative prehistoric cultural
sequence in the northern Marquesas Islands,
French Polynesia.—Journal of the Polynesian
Society 75:286-303.
Sinoto, Y. S. 1970. An archaeologically based as-
sessment of the Marquesas Islands as a dispersal
center in East Polynesia. Pp. 105-130 in R. C.
Green and M. Kelly, eds.— Pacific anthropo-
logical records 11.
Sinoto, Y. S. 1979. The Marquesas. Pp. 110-134 in
J. D. Jennings, ed. The prehistory of Polynesia.
Harvard University Press, Cambridge, Massa-
chusetts.
Steadman, D. W. 1985. Fossil birds from Mangaia,
southern Cook Islands.— Bulletin of the British
Ornithologists Club 105(2):58—66.
Biological Survey, New York State Mu-
seum, The State Education Department, Al-
bany, New York 12230.
PROC. BIOL. SOC. WASH.
100(3), 1987, pp. 529-531
A NEW SPECIES OF STOMATOPOD, EURYSQUILLA
PUMAE (CRUSTACEA: STOMATOPODA: EURYSQUILLIDAB),
FROM THE GULF OF CALIFORNIA, MEXICO
Michel E. Hendrickx and Jose Salgado-Barragan
Abstract. — A new species of the stomatopod genus Eurysquilla Manning, E.
pumae, is described from the Gulf of California, Mexico.
Two species of the genus Eurysquilla
Manning, EF. veleronis (Schmitt) and E. so-
lari Manning, are known from the Eastern
Pacific. Up to now, only the former has been
collected along the coast of Mexico, where
it is commonly found on sandy substrates
between 29 and 118 m (Hendrickx and Sal-
gado-Barragan 1987).
During sampling activities of the B/O E/
Puma in the Gulf of California, Mexico
(CORTES Project), in Mar 1985, a male
specimen of Eurysquilla sp. was collected
by trawl. The specimen, which appears to
be distinct from £. veleronis, was latter
compared with type material of E. solari at
the USNM and identified as an undescribed
species.
Eurysquilla pumae, new species
Fig. 1
Diagnosis. —Cornea subglobular, not bi-
lobed, set obliquely on eyestalk; eyestalk al-
most cylindrical, little inflated near external
base. Ocular scales broad, slightly sloping
backward (rising forward), with rounded
anterolateral corners. Antennal processes
acute, directed forward, tip reaching ante-
rior margin of ocular scales. Rostrum tri-
angular, without terminal spine, longer than
wide. Raptorial claw with 7 teeth on dac-
tylus. Five epipodites. Sixth and seventh
thoracic segments with posterolateral angle
slightly produced backward, rounded; 8th
thoracic segment laterally rounded. Pos-
terolateral angles of abdominal somites 1,
2, and 3 rounded; 4th with acute angle but
no spine. Posterior spine present on mar-
ginal carina of 5th segment (smaller sub-
marginal spine also present on right side),
and on marginal, intermediate, and sub-
median carinae of 6th segment. Telson
broader than long, with strong median ca-
rina ending in spine and one pair of strong,
continuous carinae, parallel to median,
slightly converging towards telson anterior
margin but falling short of it; in between
these carinae, and parallel to them, 2-3
strong blunt tubercles (2 on left side, 3 on
right side of median carina); strong conical
tooth present close to inner distal extremity
of each of paired intermediate carinae; par-
allel to intermediate continuous carina, on
outer side of telson, 2 tubercles, well sepa-
rated from each other, anterior one round
and strong, close to telson anterior margin,
posterior tubercle elongate, reaching ap-
proximately to distal end of intermediate
carina. Marginal carina of telson well
marked. Short carina on basis of lateral
tooth, its posterior end bent outwards; strong
tubercle on intermediate tooth; a little ele-
vated, obliquely set carina on submedian
tooth. Inner intermediate denticle elongat-
ed, with very short inner sinus; outer inter-
mediate denticle much shorter, thickened
on posterior margin and slightly upturned.
Inner lateral denticle inconspicuous, outer
one a rounded lobe. Proximal segment of
exopod of uropod with 7 movable spines
(one broken off). Basal prolongation of uro-
pod relatively thick, with 3 spines, median
terminal spine strongest, inner proximal
spine shortest; inner margin smooth.
VOLUME 100, NUMBER 3
Color. — Preserved specimen shows car-
apace with large patch of black pigment close
to posterolateral corners; posterolateral cor-
ners of lst abdominal segment with black
patch of pigment, subtriangular in shape.
Conspicuous black mark on posterolateral
corner of 6th segment, suboval, with light
linear longitudinal band about '4 away from
outer margin of mark, and circular spot close
to inner border. Median portion of all tho-
racic and abdominal somites with median
patch of black pigment. Base of lateral teeth
of telson, posterior center of telson and last
segment of both exopod and endopod of
uropod black.
Measurements. —Only one specimen col-
lected, a male of 34 mm (total length).
Holotype. —Reference Collection of Es-
tacion Mazatlan, UNAM, EMU-2415.
Distribution. —Known only from the
type locality, off Estero Tastiota, Sonora
(28°17’N, 111°37’W), Gulf of California,
Mexico. Collected at 34-37 m, on sandy
bottom.
Etymology. —The species is named for the
Oceanographic Vessel E/ Puma, of the Uni-
versidad Nacional Autonoma de Mexico.
Discussion. —Eurysquilla pumae can be
distinguished from E. veleronis, the other
species present in the area, by its much
smaller rounded, not elongated cornea which
is not as strongly bilobed as in E. veleronis.
Eurysquilla solari, a species with a smaller
eye than E. veleronis (but bilobed and larger
than that of E. pumae) has a spiny rostrum
and long, acute antennal processes that
clearly overreach the ocular scales.
The telsons of the three species of Eury-
squilla known from the East Pacific differ
in the complexity of their dorsal sculpture.
Eurysquilla solari has the most elaborate
pattern with, in addition to the median ca-
rina, a series of three lateral carinae that are
strong and tuberculate; also present is a patch
of ridges and tubercles on the base of both
—
Figs I.
531
the intermediate and the lateral teeth (Man-
ning 1970). In E. veleronis, the sculpture are
rather reduced when compared to EL. solari,
but three pairs (two well defined, a third one
less so) of lateral carinae can still be distin-
guished; the tubercles are also less conspic-
uous (Schmitt 1940). In E. pumae, a further
reduction of the carinae complex can be ob-
served, and there is only one clearly marked,
very strong dorsal pair of carinae parallel to
the median one (not tuberculate), the rest
being. reduced to a few tubercles. Eury-
squilla solari is also the only species that has
a posterior spine on the marginal carina of
the 4th abdominal segment and a well de-
fined intermediate spine on the 5th (Man-
ning 1970). The small marginal inner spine
on the basal prolongation of the uropod,
present in both E. veleronis and E. pumae,
is absent in E. solari.
Acknowledgments
This study was partly supported by
CONACYyT, México (ICECXNA-021996).
A stay at the USNM by one of us (MEH)
was supported by a grant from AID, United
States Embassy, Mexico. We thank R. B.
Manning for the help provided.
Literature Cited
Hendrickx, M. E., and J. Salgado-Barragan. 1987. Los
estomatopodos (Crustacea: Hoplocarida) del
Pacifico mexicano.—Instituto de Ciencias del
Mar y Limnologia, Universidad Nacional Au-
tonoma de México, Publicacion Especial. [in
press]
Manning, R. B. 1970. Nine new American stomato-
pod crustaceans. — Proceedings of the Biological
Society of Washington 83(8):99-1 14.
Schmitt, W. L. 1940. The stomatopods of the west
coast of America.— Allan Hancock Pacific Ex-
peditions 5(4):129-225.
Estacion Mazatlan, Instituto de Ciencias
del Mar y Limnologia, UNAM. P.O. Box
811, Mazatlan, Sinaloa, 82000, Mexico.
Eurysquilla pumae, holotype: A, Fifth and sixth abdominal somites and telson; B, Anterior part of
the body; C, Uropod in ventral view. Scale bar = 1.0 mm. (Setae omitted.)
PROC. BIOL. SOC. WASH.
100(3), 1987, pp. 532-542
A NEW SPECIES OF CHEVALIA FROM THE
CARIBBEAN SEA (CRUSTACEA: AMPHIPODA)
J. L. Barnard and J. D. Thomas
Abstract. —A new Caribbean species, Chevalia carpenteri is described; here-
tofore only one species of the genus was known, but we have examined material
that suggests morphs may represent distinct species or subspecies. We report
C. mexicana and extend its range from the northern Gulf of Mexico to Belize.
Our new species differs from the world complex of C. aviculae morphs in the
ovate article 2 of pereopods 5-7 and the posteriorly flat epimera with large
notches.
Chevalia has heretofore been monotypic,
the type species being C. aviculae Walker
(1904) from Ceylon. Two other species,
Chevalia mexicana Pearse (1913) from the
Gulf of Mexico and Neophotis inaequalis
Stout (1913) from California were synony-
mized with C. aviculae by Shoemaker (1942)
and J. L. Barnard (1962, 1970). Chevalia
mexicana is redescribed from specimens
collected at Belize and reexamination of type
material in Smithsonian Institution. A
strongly distinct new species from the Flor-
ida Keys and Belize is described and some
previous materials reported as C. aviculae
from California, Hawaii, and the Galapagos
Islands, are reviewed. This new study sug-
gests that C. aviculae has minutely distinct
populations in those places but the type ma-
terial from Ceylon remains obscure and an
elaboration of possible species of subspecies
around the tropics is not possible. A new
diagnosis of the genus and a key to these
taxa 1s presented below.
Methods
Two diagnoses each are given for Che-
valia mexicana and C. carpenteri to com-
pare each with the superspecies complex of
C. aviculae and with the (presumably spe-
cific) divisions of C. aviculae not yet pos-
sible to implement.
Legend
Capital letters as follows refer to parts;
lower case letters to the left of capital letters
refer to specimens noted in legends; lower
case letters to right of capitals refer to ad-
jective modifications in list: B, body; C, coxa;
D, dactyl; F, accessory flagellum; G, gnatho-
pod; I, inner plate or ramus; L, labium; M,
mandible; O, outer plate or ramus; P, pe-
reopod; Q, pleopod; R, uropod; S, maxil-
liped; T, telson; U, labrum; W, urosome; X,
maxilla; Y, gill; Z, oostegite; 1, left; r, right.
Chevalia Walker
Chevalia Walker, 1904:288 (Chevalia avi-
culae Walker, 1904, monotypy).—J. L.
Barnard, 1969:271.
Neophotis Stout, 1913:652 (Neophotis in-
aequalis Stout, 1913, monotypy).
Diagnosis. —Corophiidean with fleshy
telson; article 2 of antenna | longer than
article 1, article 3 shorter than 1, accessory
flagellum 2-3 articulate; antenna 2 slightly
shorter than antenna 1; labrum weakly ex-
cavate below; incisors and laciniae mobiles
serrate, rakers about 4, palp article 3 slightly
shorter than 2, thinly clavate, with ABDE
setae; inner lobes of labium large and fleshy;
inner plate of maxilla 1 present but small
and poorly armed, outer plate with 10-11
VOLUME 100, NUMBER 3
spines, palp 2-articulate, article 1 short; in-
ner plate of maxilla 2 with oblique facial
row of setae; plates of maxilliped thin, outer
with sparse but large medial spines, palp
thin, dactyl stubby, multi-armed; coxae
small, disjunct, coxae 1-2 larger than 3-7;
gnathopods of sexes alike, gnathopod | fee-
ble, almost simple, carpus elongate; gnatho-
pod 2 large, carpus short and lobate, pro-
podus large, subrectangular, palm weakly
oblique or transverse, defined by tooth, dac-
tyl thick; locking spines of pereopods 6-7
comprised of one short thick spine and one
elongate thin seta, often on pereopod 5 both
thin and short, on pereopods 3—4 both long
and seta-like; dactyls of pereopods 3—4 sim-
ple, of pereopods 5—7 strongly bent and with
large outer tooth; pereopods 5-7 short but
increasingly elongate, feeble; oostegites ab-
sent on coxa 2 (and occasionally coxa 3),
thin, strap-shaped, sparsely setose; bran-
chiae thin, on coxae 2—6; epimera simple;
urosomites 1-2 fused, urosomite 3 free; out-
er rami of uropods 1-2 shortened, armed
apically, inner rami styliform and unarmed
apically; uropod 3 short, rami subequal, thin,
setose only apically; telson short, as broad
as long, squared, well setose dorsally.
Key to the Taxa of Chevalia
1. Article 2 of pereopods 5-7 ovate,
not produced posteroventrally, epi-
mera 1-3 straight behind, with dis-
tinctly strong posteroventral notch,
coxal gill 2 shortened, oostegites 2
pairs carpenteri
— Article 2 of pereopod 7 subquad-
rate, produced or squared postero-
ventrally, epimera 1-3 convex be-
hind, with weak notch or none, coxal
gill 2 not shortened, oostegites 3
o 6 © © © © 8 6 8 ee ee lel lel
pairs ..... aviculae complex .... 2
2. Palm of gnathopod 2 transverse .. 3
Palm of gnathopod 2 oblique .... 4
3. Palm of gnathopod 2 with deep si-
nus near dactylar hinge, article 2 of
pereopod 7 obtusely lobate, anten-
ae)
nae and mandibular palp ordinary
ee eee ee yh eee eee ee HAWAII
— Palm of gnathopod 2 lacking sinus,
article 2 of pereopod 7 quadrate, an-
tennae and mandibular palp im-
mense ............ (MADAGASCAR,
WEST ATLANTIC) mexicana
4. Coxa | with sharply attenuate an-
teroventral apex ................
(CEYLON, type) aviculae
— Coxa 1 blunt and not attenuate ap-
icoventrally
5. Article 2 of pereopod 7 with pro-
tuberant posteroventral lobe, hinge
notch of palm deep .............
GALAPAGOS + FIJI
— Article 2 of pereopod 7 quadrate,
hinge notch of palm weak .......
ne ere (CALIFORNIA) inaequalis
° © © © © © © © © © ©
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Chevalia aviculae Walker, Superspecies
Diagnosis. — Article 3 of antenna | about
Q.70-0.75 times as long as article 1; acces-
sory flagellum 2-3 articulate, article | short
or long (2-articulate and article | long in
California, Galapagos, Hawaii, but 3-artic-
ulate with article | short in western Atlantic
=mexicana). Coxal gill 2 almost as long as
other gills. Three pairs of oostegites present
(on coxae 3-5). Article 2 of pereopods 5—7
pyriform, with extended or produced pos-
teroventral corner (especially on pereopod
7); article 5 of pereopods 5-7 with | (rarely
2) main emergent spine(s); article 4 of pe-
reopod 7 slender (type and Galapagos) or
broad in California-Hawau. Epimer 1-3
with convex posterior margin, posteroven-
tral notch tiny or absent. Apex of outer ra-
mus on uropod 2 with 5—6 subsidiary spines,
some of these directly basal to 3 main distal
spines.
Chevalia aviculae Walker
Chevalia aviculae Walker, 1904:288—290,
pl. 7, fig. 50, pl. 8, fig. 50; 1909:341.—K.
H. Barnard, 1916:252; 1937:169, fig. 15.
534
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Figen 1h
Chevalia sp., Galapagos, male “‘p.”’
Diagnosis. — Accessory flagellum 2-artic-
ulate, article 1 elongate (Walker probably
overlooked tiny article 2); anteroventral
corner of coxa | attenuate and sharp; coxa
2 [?weakly excavate ventrally]; palm of
gnathopod 2 oblique, notch near hinge ab-
sent; article 2 of pereopod 7 with weakly
protuberant posteroventral corner, article 4
slender. [Unknown are gills, oostegites and
minute details on outer ramus of uropod 2.]
Distribution. —Indian Ocean.
Chevalia inaequalis (Stout), California
Neophotis inaequalis Stout, 1913:653.
Chevalia aviculae: Shoemaker, 1942:39.—
J. L. Barnard, 1962:17, fig. 5.—Conlan,
1983:61, fig. 32.
Diagnosis. — Accessory flagellum 2-artic-
ulate, article 1 shortened; anteroventral cor-
ner of coxa | blunt; coxa 2 weakly excavate
or flat ventrally; palm of gnathopod 2
oblique, notch near hinge moderate; article
2 of pereopod 7 not protuberant at postero-
ventral corner, article 4 stout; outer ramus
of uropod 2 with 2 marginal setae.
Material.— Allan Hancock Foundation
Velero IV 5164 (reexamination of material
reported by J. L. Barnard 1962).
Distribution. — British Columbia to outer
Baja California.
Chevalia sp., Galapagos
Fig. 1
Chevalia aviculae: J. L. Barnard, 1979:24.
Diagnosis. — Accessory flagellum 2-artic-
ulate, article 1 elongate; anteroventral cor-
ner of coxa | blunt; coxa 2 weakly excavate
or convex ventrally; palm of gnathopod 2
oblique, notch near hinge deep; article 2 of
pereopod 7 protuberant at posteroventral
corner, article 4 stout; outer ramus of uro-
pod 2 with 2 marginal setae.
Material.—JLB Galapagos 40 (reexami-
nation of material reported by J. L. Barnard
1979).
Distribution. —Galapagos Islands.
Chevalia sp., Fiji
Chevalia aviculae: Myers, 1985:76-80, fig.
59.
Diagnosis. — Accessory flagellum 2-artic-
ulate, article 1 elongate; anteroventral cor-
ner of coxa 1 blunt; coxa 2 almost straight
ventrally; palm of gnathopod 2 oblique,
VOLUME 100, NUMBER 3
notch near hinge moderate to strong; article
2 of pereopod 7 protuberant at postero-
ventral corner, article 4 slender; outer ra-
mus of uropod 2 with 1 marginal seta.
Distribution. — Fiji.
Chevalia sp., Hawaii
Chevalia aviculae: J. L. Barnard, 1970:166—
167, fig. 105.
Diagnosis. — Accessory flagellum 2-artic-
ulate, article 1 elongate; anteroventral cor-
ner of coxa | blunt; coxa 2 weakly excavate
or convex ventrally; palm of gnathopod 2
transverse, notch near hinge deep; article 2
of pereopod 7 strongly protuberant at pos-
teroventral corner, article 4 slender; outer
ramus of uropod 2 with 2 marginal setea.
Material. —JLB Hawaii 6 (reexamination
of original material, see J. L. Barnard 1970).
Chevalia mexicana Pearse
Figs. 2-3
Chevalia mexicana Pearse, 1913:374—376,
fig. 5.
?Chevalia aviculae: Shoemaker, 1921:101.—
?Ledoyer, 1982:198, fig. 68.
Diagnosis (comparison to C. aviculae su-
perspecies and C. carpenteri).— Article 3 of
antenna | about 0.70 times as long as article
1; accessory flagellum 3-articulate, article |
short. Coxal gill 2 almost as long as other
gills. Three pairs of oostegites present (on
coxae 3-5). Article 2 of pereopods 5—6 ovate,
of pereopod 7 rectolinear, lacking extended
or produced posteroventral corner on pe-
reopod 5, but with squared posteroventral
corner on pereopods 6-7, article 5 of pereo-
pods 5-7 with 2 main emergent spines; ar-
ticle 4 of pereopod 7 slender. Epimera 1-3
with weakly convex posterior margin, pos-
teroventral notch obsolescent. Apex of out-
er ramus on uropod 2 with 5+ subsidiary
spines directly basal to 3 main distal spines.
Diagnosis (comparison to species in C.
aviculae complex).—Accessory flagellum
335
3-articulate, article 1 shortened; anteroven-
tral corner of coxa | blunt; coxa 2 weakly
excavate ventrally; palm of gnathopod 2 al-
most transverse, notch near hinge weak or
absent; article 2 of pereopod 7 not protu-
berant at posteroventral corner, article 4
slender; outer ramus of uropod 2 with 1
marginal seta.
Antenna | unlike other species, about 1.5
times as long as head plus first 4 pereonites
together, in C. mexicana antenna | about
2.3 times as long as ““comparative,”’ antenna
2 concomitant and mandibular palp simi-
larly elongate (mandibular palp cleans an-
tennae); inner basal margin of dactyl on
gnathopod 2 with hump.
Description of female ‘“‘w”’ 4.77 mm.—As
in illustrations; primary flagellum of anten-
na | with one aesthetasc on each article ex-
cept 1, 8, and 10, and one tiny one on article
11; gnathopod 2 with smooth transverse
palm, dactyl as long as palm, medial face
of article 5 lacking tiny prickles; 3 pairs of
oostegites very thin, sparsely setose; ratio of
peduncle to outer ramus to inner ramus on
pleopods = 30:45:39, outer rami with 10
articles, inner with 8-9, peduncle with 2
coupling hooks, one apical seta (remote from
coupling hooks); outer ramus of uropod |
with lateral and medial fine serrations dis-
tally, these absent on outer ramus of uropod
2, this ramus with | marginal seta; inner
ramus of uropod 2 with large lateral serra-
tions, each small tooth subserrate.
Male “vy” 3.75 mm.—Like female but
body smaller and armaments therefore few-
er.
Distinctions from C. carpenteri. —Palp of
mandible elongate; palp of maxilla | with
3 outer facial setae, inner plate of maxilla 2
shortened; outer rami of uropods 1-2 of C.
aviculae superspecies-form in presence of
5+ subsidiary spine-teeth.
I/[lustrations. —Following parts like C.
carpenteri and therefore neither described
nor illustrated: posterior body, urosome,
pleon, upper and lower lips, maxilla |, max-
illiped (but inner plate with 6 medial setae),
OR2
\\
ZZ
SED cae > eae
Fig. 3. Chevalia mexicana, female “w.”
YW.
esr ea,
LSE
SCALE
7 /
WF
\ AS | fg y [L .
< QZ Za
) ;
ZZ ‘
in,
t ¥ Ma ‘
ue SS TOU
Se
female “‘h.”
Fig. 4. Chevalia carpenteri, female “a” holotype; h
IR1
Fig. 5. Chevalia carpenteri, female “a” holotype; h = female “‘h.”
Fig. 6. Chevalia carpenteri, female “a” holotype, h = female “h.”
VOLUME 100, NUMBER 3
pereopods 3—4 (but article 6 with 4 posterior
setae besides locking setae), uropods 1-2 pe-
duncular setae shorter, outer rami with 2
and | setae respectively), telson (but with 2
rows of 5 dorsal setae).
Type locality.—South of Panama City,
Florida, 29°16'30"’N, 85°32'00”W, Alba-
tross sta 2369, 25 to 27 fms.
Material.—USNM 195160, JDT Belize
82B, Carrie Bow Cay, reef crest, outer edge,
depth 60 feet, Halimeda in coral rubble,
coll. J. D. Thomas, male “‘v”’ 3.75 mm and
female ““w” 4.77 mm.
Relationship. —Having characters of the
C. aviculae form (multitoothed outer rami
of uropods 1-2) as well as of C. carpenteri
(presence of 2 spines on posterodistal article
5 of pereopods 5-7). See key.
Distribution. —Northern Gulf of Mexico
to the Lesser Antilles, Barbados, and Belize,
0-50 m; ?Madagascar.
Chevalia carpenteri, new species
Figs. 4—6
Diagnosis. — Article 3 of antenna | about
0.60 times as long as article 1; accessory
flagellum 3-articulate, article 1 short. Coxal
gill 2 half as long as other gills. Two pairs
of oostegites present (on coxae 4—5). Article
2 of pereopods 5-7 ovate, lacking extended
or produced posteroventral corner; article
5 of pereopods (5) 6—7 with 2 main emergent
spines; article 4 of pereopod 7 slender. Epi-
mera 1-3 with straight posterior margin,
posteroventral notch strong. Apex of outer
ramus of uropod 2 with 2-3 subsidiary
spines, none of these directly basal to 3 main
distal spines.
Description of holotype, female “‘a”’ 4.64
mm. — As in illustrations; primary flagellum
of antenna | with one aesthetasc on each
article; gnathopod 2 with weakly serrate
oblique palm, dactyl shorter than palm, me-
dial face of article 5 with tiny prickles; right
gnathopod 2 abnormally stunted, regener-
ant; 2 pairs of oostegites very thin, sparsely
541
setose; ratio of peduncle to outer ramus to
inner ramus on pleopods = 23:24:29, outer
rami with 7 articles, inner with 6, peduncle
with 2 coupling hooks, one apical seta (not
near coupling hooks); outer ramus of uro-
pod 1 with lateral and medial fine serra-
tions, these absent on outer ramus of uro-
pod 2, this ramus with | marginal seta; inner
ramus of uropod | with large lateral serra-
tions, each small tooth subserrate.
Holotype.—USNM 195157, female ‘‘a”’
4.64 mm.
Type locality. —JDT Belize 105, Carrie
Bow Cay, Belize, sand bores near Wee Wee
Cay, 1 m, formalin wash of algai/inverte-
brate covered coral rubble, 21 Jul 1984, coll.
J. D. Thomas.
Material. —The type locality, female ‘“‘b”’
2.96 mm, female “‘c”’ 2.54 mm, female “‘d”’
2.79 mm, female “‘e’’ 3.04 mm, female “‘f”’
2.51 mm, female “‘g’”’ 1.93 mm, female “‘h”’
3.16 mm; Florida Keys, Looe Key Reef,
forereef, formalin wash of coral rubble, 5
m, 8 Oct 1983, coll. J. D. Thomas, female
velo 7) mim:
Etymology. —Named for Michael R. Car-
penter, Smithsonian Institution, whose as-
sistance in field endeavors has been invalu-
able.
Relationship. — Differing from the com-
plex of morphs in C. aviculae by the ovate
article 2 of pereopods 5-7 and the poste-
riorly straight epimera with large notches.
As far as we can determine (except for C.
mexicana) in the morphs of C. aviculae we
have observed, our new species differs in
the presence of two tiny spines on the pos-
terodistal apex of article 5 on pereopods 6—
7 in contrast to one spine in the type species
of the genus. Also, the new species has only
two to three subsidiary spines or teeth on
the outer ramus of uropod 2 in contrast to
five to six in C. aviculae, has only two pairs
(versus three) of oostegites, and gill 2 is
shortened. Coxa 2 appears to be more
sharply attenuate anteroventrally then in any
of the morphs of C. aviculae.
542
Acknowledgments
We thank Dr. Klaus Rutzler and Mike
Carpenter of Smithsonian Institution for
field support on Carrie Bow Cay, Belize. We
thank Linda B. Lutz of Vicksburg for inking
our plates. The second author was support-
ed by NSF Grant BSR-8515186. This is
contribution 193 from the Smithsonian Reef
and Mangrove Project, Carrie Bow Cay, Be-
lize.
Literature Cited
Barnard, J. L. 1962. Benthic marine Amphipoda of
Southern California. — Pacific Naturalist 3:1-72.
1969. The families and genera of marine
gammaridean Amphipoda.— United States Na-
tional Museum Bulletin 271:1-535.
. 1970. Sublittoral Gammaridea (Amphipoda)
of the Hawaiian Islands. —Smithsonian Contri-
butions to Zoology 34:1-286.
. 1979. Littoral gammaridean Amphipoda from
the Gulf of California and the Galapagos Is-
lands. —Smithsonian Contributions to Zoology
271:1-149.
Barnard, K.H. 1916. Contributions to the crustacean
fauna of South Africa.—Annals of the South
African Museum 15:105-298.
1937. Amphipoda.—John Murray Expedi-
tion 1933-34, Scientific Report, British Mu-
seum (Natural History) 4(6):131-—201.
Conlan, K.E. 1983. Family Isaeidae: Systematics and
distributional ecology.— National Museums of
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Canada, Publications in Natural Sciences 4:i-—
viii, 1-75.
Myers, A. A. 1985. Shallow-water coral reef and
mangrove Amphipoda (Gammaridea) of Fiji. —
Records of the Australian Museum, Supplement
5:1-144.
Pearse, A.S. 1913. Notes on certain amphipods from
the Gulf of Mexico, with descriptions of new
genera and new species.—Proceedings of the
United States National Museum 43:369-379.
Shoemaker, C. R. 1921. Report on the amphipods
collected by the Barbados-Antigua Expedition
from the University of Iowa in 1918.—Univer-
sity of Iowa Studies in Natural History 9:99-
102.
. 1942. Amphipod crustaceans collected on the
Presidental Cruise of 1938.—Smithsonian Mis-
cellaneous Collections 101(11):1—52.
Stout, V.R. 1913. Studies in Laguna Amphipoda.—
Zoologisches Jahrbucher fiir Systematik 34:633-—
659.
Walker, A.O. 1904. Report on the Amphipoda col-
lected by Professor Herdman, at Ceylon, in
1902.—Ceylon Pearl Oyster Fisheries, Supple-
mentary Report 17:229-300.
. 1909. Amphipoda Gammaridea from the In-
dian Ocean, British East Africa, and the Red
Sea.— Transactions of the Linnean Society of
London (2) Zoology, 12:323-344.
(JLB) Division of Crustacea, NHB-163,
Smithsonian Institution, Washington, D.C.
20560; (JDT) Reef Foundation, P.O. Box
569, Big Pine Key, Florida 33043.
PROC. BIOL. SOC. WASH.
100(3), 1987, pp. 543-551
PINNOTHERIDS (CRUSTACEA: DECAPODA) AND
LEPTONACEANS (MOLLUSCA: BIVALVIA)
ASSOCIATED WITH SIPUNCULAN WORMS
IN HONG KONG
Raymond B. Manning and Brian Morton
Abstract.—The intertidal soft-bottom sipunculans Sipunculus nudus Lin-
naeus and Siphonosoma cumanense (Keferstein) in Hong Kong each have a
pair of commensal associates, one leptonacean bivalve and one pinnotherid
crab, inhabiting their burrows. With the former are the bivalve Fronsella oh-
shimai Habe and members of a new genus and species of pinnotherid crab,
Indopinnixa sipunculana, and with the latter are the bivalve Nipponomysella
subtruncata (Yokoyama) and the crab Mortensenella forceps Rathbun. The
bivalves occur in mean numbers of 5 (range 1-9) and 3.3 (range 1-—7)/host,
respectively, and the crabs either singly or in pairs. The pairs of associates seem
mutually exclusive of each other, with regard to both inter- and intra-host
specificity.
Two sipunculans, Sipunculus nudus Lin-
naeus and Siphonosoma cumanense (Ke-
ferstein), build deep burrows on Hong
Kong’s soft, muddy shores. Over the course
of a number of field trips by one of us (B-M.)
to such a sipunculan habitat at Tai Tam Bay
on the south coast of Hong Kong Island
[22°15'’N, 114°11’E], a number of speci-
mens of each of these species has been col-
lected, and two species of commensals have
been found in association with each of them.
Morton and Morton (1983:204) listed the
leptonid bivalve Fronsella ohshimai Habe
(Montacutidae) and the pinnotherid crab
Asthenognathus inaequipes Stimpson as as-
sociates of Sipunculus nudus; we describe
the pinnotherid here as a new genus and
species. Morton and Morton (1983:204) also
listed the leptonid bivalve Nipponomysella
subtruncata (Yokayama) (Montacutidae)
and the pinnotherid crab Pinnixa rathbuni
Sakai as associates of Siphonosoma cuma-
nense; here we correct the identity of the
crab to Mortensenella forceps Rathbun. This
note provides further information on these
commensal associations.
Materials and Methods
Sipunculan burrows were identified by
their surface apertures, and sometimes dig-
ging would expose their deep, fast-burrow-
ing inhabitants. When a sipunculan was ob-
tained, the burrow wall was examined for
possible associates. No records were kept in
cases where the associate could not be linked
positively to the burrow owner. In this way,
definite associations have been established.
As it was impossible, however, to trace ac-
curately every burrow and collect every as-
sociate, the data presented here should be
regarded as provisional.
The crab specimens reported below have
been deposited in the National Museum of
Natural History, Smithsonian Institution,
Washington (USNM).
Results
The Pinnotherid Crabs
Indopinnixa, new genus
Definition. —Carapace much wider than
long; integument firm; front narrow, with
igeale
Fourth pereopod; E, Abdomen; F, Gonopod; G, Apex of gonopod. Female, Hong Kong: H, Second to fifth
pereopods; I, Abdomen.
median groove. Orbit broadly ovate, with
wide inner hiatus partly occupied by basal
antennal article. Antennules folded into wide
fossettes, joined medially under front. Eye-
stalks very short. Ischium of third maxil-
liped short, merus much longer, both slen-
der; palp articulated to summit of merus,
distal article arising on inner side of second
article. Chelipeds moderately large, hand
large, compressed. Third walking leg (P4)
stout, largest of all walking legs; first and
second walking legs (P2, P3) very slender,
fourth and fifth legs (P4, P5) stout. Abdo-
men of male 6-segmented, fifth and sixth
segments fused; seventh segment short,
broader than long. Female abdomen 7-seg-
mented. Gonopod simple, tapering distally.
Type species. —Indopinnixa sipunculana,
new species.
Etymology.—The generic name is de-
rived from the Greek combining form,
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
B
C
Indopinnixa sipunculana. Male, Hong Kong: A, Carapace; B, Third maxilliped; C, Cheliped; D,
““indo-,” and the generic name Pinnixa. The
gender is feminine.
Remarks. — Indopinnixa, which now con-
tains only the type species described below,
differs from Pinnixa sensu stricto in the form
of the male abdomen, which is composed
of seven free segments in Pinnixa, six seg-
ments in /ndopinnixa, because the fifth and
sixth segments are fused.
Indopinnixa sipunculana, new species
Fig. 1
Pinnixa rathbuni.—Morton and Morton,
1983:78, 204. [Not P. rathbuni Sakai,
1934.]
Material.—Tai Tam, Hong Kong, Jun
1977, in burrows of Sipunculus nudus, leg.
B. Morton, | male, holotype, USNM
221697.—Same data except date, 20 Jun
1974, | ovigerous female, paratype, USNM
MALS OO),
VOLUME 100, NUMBER 3
Description.—Carapace 1.83 (male) to
2.15 (female) times broader than long,
punctate dorsally, lacking transverse ridge
across posterior part. Branchial regions each
crossed by low, tuberculate ridge, falling
short of orbit. Front bilobed. Anterolateral
margin with 2 tomentose depressions visi-
ble dorsally, lateral to orbit, depressions
connected ventrally, defining smooth, tear-
shaped area lateral to orbit. Regions of
carapace poorly marked, with transverse
median depression, and, on each side, a
shallow groove parallel to posterolateral
margin.
Chelipeds of male subequal (of female un-
known), cheliped with fixed finger in same
plane as palm, not deflected ventrally, mov-
able finger (measured dorsally) slightly
shorter than palm, latter smooth dorsally,
about as long as high, with longitudinal ridge
on outer surface. Fingers gaping, movable
finger smooth dorsally, curved, tip crossing
that of fixed finger, with triangular tooth on
opposable margin. First 2 walking legs slen-
der, first short, second about as long as third.
Third walking leg largest in both sexes, pro-
podus 1.5 to 1.7 times longer than high,
ventral (opposable) margin bicarinate, to-
mentose; merus 2 times as long as high;
dactylus smooth, neither carinate nor tu-
berculate dorsally. Fourth walking leg stout,
extending about to end of merus of third.
Walking legs with some plumose setae on
dorsal and ventral margins.
Male abdomen of 6 segments, fifth and
sixth fused; telson short and broad. Female
abdomen of 7 segments, lacking transverse
line of hairs on posterior margin of second
segment. Male gonopod simple, tapering
distally, as figured.
Size. —Carapace length of male 2.3 mm,
of ovigerous female 2.0 mm; carapace width
of male 4.2 mm, of ovigerous female 4.3
mm. Ova measure 0.3 mm.
Remarks. —Initially we identified this
species with Pinnixa penultipedalis Stimp-
son (1858:108; 1907:143), a species origi-
nally described from Hong Kong. However,
545
Stimpson’s species was characterized by him
as having a smooth and glossy carapace with
a distinct transverse ridge posteriorly, a
small palm on the cheliped, a very broad
merus on the third walking leg (height %
length), and a line of setae on the posterior
margin of the second somite of the abdomen
in the female. Our species differs in nu-
merous features: it lacks a distinct ridge pos-
teriorly on the carapace, the surface of the
carapace 1s punctate, not smooth, there is a
distinct tuberculate ridge on each branchial
region, the palm of the cheliped is large, and
the posterior margin of the second abdom-
inal somite in the female is not setose.
Indopinnixa sipunculana also resembles
the species identified by Shen (1937:298)
with Pinnixa penultipedalis Stimpson, es-
pecially in the shape of the carapace and the
general proportions of the walking legs.
Shen’s species, which probably is unde-
scribed, differs from P. penultipedalis and
resembles J. sipunculana in lacking a dis-
tinct ridge across the posterior part of the
carapace. Our species differs from Shen’s in
having a tuberculate ridge across each bran-
chial region as well as a strong tooth on the
cutting edge of the dactylus of the cheliped.
Our species further differs in lacking dorsal
tubercles on the palm and movable finger
of the cheliped and on the upper margin of
the dactyls of the walking legs, as well as the
line of setae on the posterior margin of the
second abdominal segment of the female.
Further, the eyes of our species are shorter
and the merus of the third walking leg is not
nearly so high as in Shen’s species.
Indopinnixa sipunculana also is similar
to the species from Inhambane, Mozam-
bique, identified by Barnard (1955:19) with
Pinnixa penultipedalis, to our knowledge the
only other record of a pinnotherid crab liv-
ing with a sipunculan. Barnard’s species,
which, like Shen’s, is probably undescribed,
has a sharp ridge across the posterior part
of the carapace and lacks the tuberculate
branchial ridges as well as the toothed dac-
tylus on the cheliped. Barnard’s species fur-
546
Fig. 2.
Re WH ONO
Ss
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
B H
Mortensenella forceps Rathbun, 1909. Male, Hong Kong: A, Dorsal view; B, Third maxilliped; C,
Cheliped; D, Second pereopod; E, Third pereopod; F, Fourth pereopod; G, Fifth pereopod; H, Abdomen.
ther differs in having three segments on the
male abdomen fused, rather than two as in
I. sipunculana.
Excluding Pinnixa brevipes H. Milne Ed-
wards, 1853, from Madagascar, which may
not be a Pinnixa (see comments in Seréne
(1964:277) and Schmitt, McCain, and Da-
vidson (1973:103), for references), and rec-
ords of the American Pinnixa faba (Dana,
1851) from Australia and Tasmania, which
may be erroneous (see Rathbun 1918:142),
five species of Pinnixa are known from the
Indo-west Pacific. All five have been re-
ported from Japan and were treated by Sak-
ai (1976:583). Three of the five species, P.
balanoglossana Sakai, 1934, P. haemato-
sticta Sakai, 1934, and P. penultipedalis
Stimpson, 1858, have a distinct ridge across
the posterior part of the carapace and can
be distinguished from J. sipunculana on this
feature alone.
The other two species, P. rathbuni Sakai,
1934, and P. tumida Stimpson, 1858, lack
distinct branchial ridges, and can be distin-
guished from J. sipunculana on this feature.
So far as we can tell, the male abdomens of
both of those species are made up of seven
free segments; see Yokoya (1928: fig. 6 (P.
rathbuni) and fig. 7 (P. tumida)).
Morton and Morton (1983:78, 204) re-
ported that the entoproct Loxosomella oc-
curs on Indopinnixa sipunculana.
Etymology.—The specific name alludes
to the association of this species with si-
punculan worms.
VOLUME 100, NUMBER 3
Mortensenella forceps Rathbun, 1909
Fig. 2
Mortensenella forceps Rathbun, 1909:111;
1910:337, fig. 21, pl. 1, fig. 18. — Dai, Feng,
Song, and Chen, 1980:137, fig. 9.—Mor-
ton and Morton, 1983:177, 204, fig. 9.14.
Asthenognathus inaequipes.—Morton and
Morton, 1983:204. [Not A. Inaequipes
Stimpson, 1858.]
Material. —Tai Tam, Hong Kong; in bur-
rows of Siphonosoma cumanense, 19 Dec
1974, leg. B. Morton, 1 male, | female,
USNM 222501.
Size. —Carapace length of male 5.0 mm,
of female 4.6 mm; carapace width of male
8.0 mm, of female 7.5 mm.
Remarks.— These specimens agree well
with the accounts of this species given by
Rathbun (1909, 1910) and Dai et al. (1980)
except that figures given in each of those
accounts show the third maxilliped to have
an indistinct suture between the ischium and
merus; no suture is visible in either of our
specimens.
Rathbun’s material, from Koh Chang,
Thailand, was taken from very shallow
water, “outside mangroves.’ Morton and
Morton (1983) recorded this species from
Hong Kong as an associate of the echiuroid
worm Ochetostoma _ erythrogrammon
Leuckart and Ruppel, 1828. The material
reported by Dai et al. (1980) was taken off
Hainan Island, China; no host was recorded
by those authors. These are the only records
of this species known to us.
The Sipunculans and Their Associates
Sipunculus nudus Linnaeus
Two associates were found with this
species: the leptonacean bivalve Fronsella
ohshimai Habe, 1958, and an undescribed
genus and species of pinotherid crab (Table
1, Fig. 3). The bivalve is an unusual com-
mensal in that it often attaches directly to
the host. Up to nine individuals have been
recorded attached to a single host by byssal
threads, apparently on the posterior valve
547
Table 1.—Records of the occurrences of burrow as-
sociates of Sipunculus nudus and Siphonosoma cu-
manense from Tai Tam Bay, Hong Kong (M = male,
F = female, ovig. = ovigerous).
S. cumanense
S. nudus
Nippono-
mysella
subtruncata
Fronsella
ohshimai
Mortensenella
forceps
Indopinnixa
Date sipunculana
20/6/74 IM
20/6/74 1F ovig.
20/6/74 1F ovig.
20/6/74 1F ovig.
20/6/74 3
20/6/74 y)
19/12/74 3
19/12/74
2/6/77
2/6/77
2/6/77 1
2/6/77
2/6/77
2/6/77
23/4/79
23/4/79
23/4/79 IM
26/8/80 9
26/8/80 7
26/8/80
26/8/80 1
21/6/82 8 1F ovig.
30/1/84 2
30/1/84 5
30/1/84 D
30/1/84 2
3
Mean 5 l
1M, 1F
1M
IM
1M
Re BW
1M
margins. The brachyuran Indopinnixa si-
punculana also has been recorded regularly
in the burrows, usually as solitary individ-
uals. Ovigerous females of the crab were
taken in June of 1974 and 1982. In only one
instance (June 1982) were bivalve and crab
simultaneously recorded in the same bur-
row. Mean numbers of bivalves and crabs
per host were 5 (range 1-9) and 1, respec-
tively.
Siphonosoma cumanense (Keferstein)
Two associates were recorded with this
species: the leptonacean bivalve Nippono-
mysella subtruncata (Yokoyama) and the
pinnotherid crab Mortensenella forceps
548
Sh MES ae er AeR OTe ora See
ce ae
CEOS
SRO
S ZS Hf ip
iB
Coe
ris
Icm
Fig. 3.
Imm
Posterior <<
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
By
» Posterior
: -
ves: Anterior
Pow a
A, Sipunculus nudus with 3 specimens of Fronsella ohshimai attached. B, Fronsella ohshimai: B1,
External view of left shell valve; B2, Interior view of hinge plate of right shell valve; B3, Interior view of hinge
plate of left shell valve. C, Indopinnixa sipunculana.
Rathbun (Table 1, Fig. 4). The former lies
laterally adpressed, but not attached bys-
sally, against the burrow wall. Up to seven
individuals were recorded from a single bur-
row. The crab Mortensenella forceps was re-
corded from the burrows on two occasions.
On 26 August 1980 a single male was col-
lected, and on 19 December 1974 a pair of
crabs was taken. Mean numbers of bivalves
and crabs per host were 3.3 (range 1-7) and
1.5, respectively.
Discussion
Sipunculans are well known as hosts of
leptonacean bivalves. Boss (1965) and Ste-
VOLUME 100, NUMBER 3
Hig:
1]
me R REG re
Wee \ y :
SR Wa RASH ‘
LOS thy 1 5
S Vg) q D Vey 1
i Wea, ;
\ NS bE aT yf By
WA PTS LO Bh iy Ty
ve ea /, :
Ny CESS }
) x
sh,
Anterior
Posterior
549
Anterior
Fig. 4. A, Siphonosoma cumanense. B, Nipponomysella subtruncata: B1, External view of left shell valve;
B2, Interior view of hinge plate of right shell valve; B3, Interior view of hinge plate of left shell valve. C,
Mortensenella forceps.
phen and Edmonds (1972) have reviewed
much of the literature detailing such asso-
ciations. The most detailed study of such
an association, 1.e., Mysella cuneata (Verrill
and Bush) with Phascolion strombi (Mon-
tagu), is that of Gage (1968). Both of the
leptonaceans found with sipunculans in
Hong Kong have been recorded from the
same hosts elsewhere. Fronsella ohshimai is
a known associate of Sipunculus nudus from
Japan (Habe 1958, 1961, 1964, 1977; Koy-
ama et al. 1981); as in Hong Kong, it at-
taches to the worm by byssal threads. Sim-
ilarly, Nipponomysella subtruncata is known
to be a commensal of Siphonosoma cu-
manense elsewhere, e.g., Utinomi (1972) as
Mysella subtrucata [sic]. Koyama et al.
(1981) reported it, possible erroneously, to
be associated with Sipunculus nudus.
In Hong Kong, these two bivalves occur
550
in groups of, on the average, 5 for F. oh-
shimai and 3.3 for N. subtruncata. They al-
most certainly capitalize on the flow of ox-
ygenated water through the burrow, and,
concomitantly, on suspended food parti-
cles.
Herein we have corrected identifications
by Morton and Morton (1983) of the pin-
notherid crabs also inhabiting the burrows
of the two sipunculans.
Of interest is that each sipunculan has a
pair of associates that are different from,
and mutually exclusive of, the other pair.
Furthermore, each of the pairs of associates
seems mutually exclusive, so that each si-
punculan possesses either a bivalve, in vary-
ing numbers, or a crab, either singly (more
commonly) or as a pair. What determines
selective burrow occupation is unknown, but
it seems possible that the more active crabs
may inhibit establishment of the more sed-
entary bivalves. There is no other infor-
mation on this aspect of such relationships,
Stephen and Edmonds (1972), for example,
not recording crabs as sipunculan associates
and giving few examples of multiple com-
mensalism, e.g., the annelid, bivalve and
entoproct associates of Phascolion strombi
(Montagu), but not mutual exclusivity. The
Hong Kong sipunculans and their associates
may have much to offer in gaining an un-
derstanding of these poorly understood taxa
and neglected phenomena.
Acknowledgments
We thank M. E. Rice, Smithsonian In-
stitution, for her comments on an early draft
of this manuscript. Figures 1 and 2 were
prepared by Lilly King Manning. This is
contribution number 195 from the Smith-
sonian Marine Station at Link Port, Fort
Pierce, Florida.
Literature Cited
Barnard, K. H. 1955. Additions to the fauna-list of
South African Crustacea and Pycnogonida.—
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Annals of the South African Museum 43(1):1-
107.
Boss, K. K. 1965. Symbiotic erycinacean bivalves. —
Malacologia 3:185-195.
Dai, A.-Y., Z.-Q. Feng, Y.-Z. Song, and G.-X. Chen.
1980. New species and new records of Family
Pinnotheridae from Hainan Island.— Acta Zoo-
taxonomica Sinica 5(2):129-143. [In Chinese.]
Gage, J. 1968. The mode of life of Mysella cuneata,
a bivalve ““commensal”’ with Phascolion strombi
(Sipunculoidea).— Canadian Journal of Zoology
46:919-934.
Habe, T. 1958. Descriptions of five new bivalves
from Japan.— Venus 20:174—180.
1961. Coloured illustrations of the shells of
Japan, vol. 2. Hoikusha Publishing Co., Ltd.,
Japan, 183 pp.
. 1964. Two commensal bivalves from the west
coast of Kyushu, Japan.— Venus 23:137-139.
1977. Systematics of Mollusca in Japan: Bi-
valvia and Scaphopoda. Hoikusha Publishing
Co., Ltd., Japan, 372 pp.
Koyama, K., T. Yamamoto, Y. Toki, and H. Minato.
(Eds) 1981. A catalogue of molluscs of Waka-
yama Prefecture, the Province of Kii. I. Bival-
via, Scaphopoda and Cephalopoda. — Publica-
tions of the Seto Marine Biological Laboratory,
Special Publication Series 7(1):i-xx, 1-301.
Morton, B., and J. Morton. 1983. The sea shore ecol-
ogy of Hong Kong. Hong Kong University Press,
Hong Kong, 350 pp.
Rathbun, M. J. 1909. New crabs from the Gulf of
Siam.— Proceedings of the Biological Society of
Washington 22:107-114.
1910. Brachyura. The Danish expedition to
Siam 1899-1900, V.—Kongelige Danske Vi-
denskabernes Selskabs Skrifter, 7. Raekke, Na-
turvidenskabelig og Mathematisk Afdeling 5, 4:
301-367, pls. 1, 2, map.
1918. The grapsoid crabs of America.—
United States National Museum Bulletin 97:1-
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Sakai, T. 1976. Crabs of Japan and the adjacent seas.
[In 3 volumes: (1) English text, xx1x + 773 pages;
(2) plates, pages 1-16, plates 1-251; (3) Japanese
text, pages 1-461.! Kodansha Ltd., Tokyo.
Schmitt, W. L., J. C. McCain, and E. S. Davidson.
1973. Decapoda I: Brachyura I: Fam. Pinno-
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eds., Crustaceorum Catalogus 3:1—-160.
Seréne, R. 1964. Goneplacidae et Pinnotheridae. Pa-
pers from Dr. Th. Mortensen’s Pacific Expedi-
tion 1914-1916, 80.— Videnskabelige Medde-
lelser fra Dansk Naturhistorisk Forening 1
Kjobenhavn 126:181-282, pls. 16-24.
Shen, C.-J. 1937. Second addition to the fauna of
brachyuran Crustacea of North China, with a
VOLUME 100, NUMBER 3
check list of the species recorded in this partic-
ular region.—Contributions from the Institute
of Zoology, National Academy of Peiping 3(6):
277-312.
Stephen, A. C., and S.J. Edmonds. 1972. The phyla
Sipuncula and Echiura. The Trustees of the Brit-
ish Museum (Natural History), London, 528 pp.
Stimpson, W. 1858. Crustacea Ocypodoidea. Pro-
dromus descriptionis animalium evertebrato-
rum, quae in Expeditione ad Oceanum Pacifi-
cum Septentrionalem, a Republica Federata
missa, Cadwaladaro Ringgold et Johanne Rod-
gers Ducibus, observavit et descripsit, Pars V.—
Proceedings of the Academy of Natural Sciences
of Philadelphia 1858:93-110.
1907. Report on the Crustacea (Brachyura
and Anomura) collected by the North Pacific
Exploring Expedition, 1853—1856.—Smithson-
ian Miscellaneous Collections 49:1—240.
551
Utinomi, H. 1972. Coloured illustrations of seashore
animals of Japan. Hoikusha Publishing Co., Ltd.,
Japan, 166 pp. [In Japanese. ]
Yokoya, Y. 1928. Brachyura and crab-shaped Ano-
mura. Report of the Biological Survey of Mutsu
Bay, 10.—Science Reports of the Tohoku Im-
perial University, Sendai (4th series, Biology)
3(4, fascicle 2):757-784.
(RBM) Department of Invertebrate Zo-
ology, National Museum of Natural His-
tory, Smithsonian Institution, Washington,
D.C. 20560; (BM) Department of Zoology,
The University of Hong Kong, Hong Kong.
PROC. BIOL. SOC. WASH.
100(3), 1987, pp. 552-558
THE PYCNOGONIDA TYPES OF H. V. M. HALL
C. Allan Child
Abstract.—The five recently rediscovered pycnogonid types described by
Harry V. M. Hall (1912, 1913) are reexamined in terms of current literature
and synonymies. One species remains valid as originally named (Anoplodac-
tylus californicus), two remain valid under different but currently accepted
genera (Nymphopsis spinosissima, Callipallene californiensis), and two are re-
duced to synonymy under previously named species (Ammothea nudiuscula as
Achelia alaskensis, and Ammothella biunguiculata var. californica as A. biun- —
guiculata). The name Anoplodactylus californicus Hall, 1912, takes precedence
over the frequently encountered A. portus Calman, 1927, which is reduced to
a synonym.
Harry V. M. Hall published two short pa-
pers (1912, 1913) describing four new pyc-
nogonid species and one new variety, all
from California. He named these five An-
oplodactylus californicus, Ammothella
biunguiculata var. californica, A. spinosis-
simus, Ammothea nudiuscula, and Pallene
californiensis. These species have been
largely ignored or sometimes reported in the
literature, but have never had the benefit of
reexamination in light of current literature,
nor have Hall’s specimens ever been avail-
able for comparison. Hall did not deposit
the species in the Museum of the University
of California (Berkeley) as he stated (1913:
127), and they were therefore regarded as
lost. Hedgpeth (1939:461-463, fig. 2m-—r)
even named a neotype and redescribed one
of the species and listed several other of
Hall’s species as part of the California fauna
while republishing Hall’s inadequate figures
for three of them. Hedgpeth, in California,
could not find Hall’s types either.
The estate of Leon J. Cole donated to the
National Museum of Natural History in
1939 Cole’s pycnogonid literature, files,
types on slides and in alcohol, and other
miscellaneous specimens. A bottle of Cole’s
types in alcohol was subsequently mis-
placed and lost, and has only recently come
to light. This most important bottle of the
Cole collections contains all of Cole’s types
and other specimens not mounted on slides
and also contains most of the material pub-
lished by Hall in 1913 (127-142). The Hall
labels are presumably in his handwriting and
not Cole’s, many examples of which are
available for comparison in Cole’s files and
with his specimens. Hall (1912:91, 1913:
128) acknowledged the assistance of Cole in
his work on the pycnogonids of California
and he presumably sent Cole his later ma-
terial for examination. Cole evidently kept
this material with his own specimens and it
subsequently came to the National Museum
upon his death. There are specimens of all
five of the above species, identified by Hall,
along with several others listed in his 1913
paper, including the types of the two species
he described therein. The discovery of this
long-lost material finally permits clarifica-
tion of the status of Hall’s species, some 75
years after they were originally described.
Achelia alaskensis (Cole)
Ammothea alaskensis Cole, 1904:266-268,
pl. 12, fig. 4, pl. 17, figs. 4-12.— Losina-
Losinsky, 1933:59—-60, fig. 10.—Schmitt,
1934:70.—Exline, 1936:421.—Okuda,
VOLUME 100, NUMBER 3
1940:73-86, figs. 1—-10.—Losina-Losin-
sky, 1955:160, pl. 42, fig. 6.
Ammothea (Achelia) alaskensis: Schimke-
witsch, 1929:151-156, figs. 42,45.—
Ohshima, 1933:144—146, fig. 1; 1936:866.
Achelia alaskensis: Hedgpeth, 1949:289
[text].—Utinomi, 1954:14, figs. 6-7.—
Losina-Losinsky, 1961:91.—Utinom1,
1971:329.—Kim and Hong, 1986:44—46,
fig. 7.
Ammothea nudiuscula Hall, 1913:135—137,
pl. 3, figs. 1-8.—Hilton, 1939a:32.—
Hedgpeth, 1941:256 [key], pl. 10.—Hil-
ton, 1942¢:93.
Achelia nudiuscula: Hedgpeth, 1964:208
[key], fig. 94b.
Achelia gurjanovii Losina-Losinsky, 1961:
93-95, fig. 18.
Material examined. —Holotype (Ammo-
thea alaskensis Cole, 1904), male, Orca,
Alaska; Allotype, female, Orca; Paratypes,
2 females, Popof Island, Alaska; Holotype
(Ammothea nudiuscula Hall, 1913), female;
San Francisco, California.
Remarks.—1 can find no reference in
Cole’s papers or notes in which he identified
or recognized Hall’s species as his previ-
ously described Alaskan species. The slight
differences between Hall’s specimen and that
figured by Cole are evident in the variation
among the three females in Cole’s type lot.
Almost all of Cole’s later work on pycno-
gonids treated Atlantic species, as he was
based for field work on the Atlantic coast
of America or Bermuda and did not publish
again on California species.
This species 1s conspicuous among the
bewildering numbers of Achelia species
found in northwestern North America by
its lack of heavily setose or tuberculate lat-
eral processes. The lateral processes, aside
from bearing one or two tiny anterolateral
setae, are usually without other adornment.
The four terminal segments of the palp are
always short, only about as long as their
widths, and the first coxae each bear a single
dorsodistal tubercle which 1s broad but only
553
slightly longer than its width. The legs are
quite setose in the male but much less so in
the female, and the abdomen is most often
carried horizontally. The ocular tubercle is
shorter than its basal width. The distal tu-
bercle on each chelifore first segment 1s var-
iously pointed, rounded, or a low swelling
in the several types examined, but is always
more acutely pointed in the male.
I tentatively place Achelia gurjanovii Los-
ina-Losinsky in this synonymy even though
the type figures are rather diagramatic and
I have no specimens bearing this name. The
proboscis of Losina-Losinsky’s specimen
(1961:94, fig. 18A) is narrower than that of
the material in hand; there are apparently
randomly-placed small lateral process tu-
bercles, and a full segmentation line exists
between the first and second trunk segments
which is not evident or only faintly present
in Cole’s material. There is a great similarity
between Losina-Losinsky’s figures of the
appendages and those of Cole (1904, pl. 17).
The ovigers and legs are almost exactly the
same, but the palp segments of A. gurjanovii
are slightiy longer and more slender. I con-
sider these differences to be those of geo-
graphically different populations, particu-
larly in a genus in which some species display
enormous variation (e.g., 4. assimilis (Has-
well)). This question must be settled by col-
lection and comparison of fresh material of
A. gurjanovii and A. alaskensis from both
sides of the northern Pacific.
The distribution of this species, as pro-
posed above, 1s now known to be from the
mid-California coast through Alaska and the
Russian and Korean coasts to the middle
of Japan, in littoral habitats to as deep as
180 meters.
Ammothella biunguiculata (Dohrn)
Ammothea bi-unguiculata Dohrn, 188 1:26,
105, 158-160, pl. VIII, figs. 1-3.
Ammothella biunguiculata: Stock, 1974:12-
13 [literature], fig. 1.
Ammothella biunguiculata var. californica
554
Hall, 1912:93—95, figs. 50, 52A, E, G, K;
1913:130, 132 [key].— Hilton, 1915a:68;
1915b:204; 1920:93.
Material examined. —Topotypes, 81
specimens, Laguna Beach, California, NW
of pier under stones, coll. H. V. M. Hall,
Jul 1912.
Remarks. —Hilton carried Hall’s trino-
mial through several of his superficial pa-
pers, but it was Hedgpeth (1941:259) who
first proposed that Hall’s species was a syn-
onym of Dohrn’s species without the var-
ietal designation. Stock (1974:95) combined
all three varieties under Dohrn’s species, to
which I agree. If the varieties of this species
were to be maintained, then the genus Ache-
lia would, by the same reasoning, be crowd-
ed with varieties or subspecies unnecessar-
ily encumbering the many variable species
of this genus.
This species has been found in many shal-
low localities of the oceans of the world and
may eventually become known as a pan-
temperate species. It is known to be dis-
tributed from the littoral to 135 meters.
Nymphopsis spinosissima (Hall)
Ammothella spinosissima Hall, 1912:95-99,
figs. 51, 52B, D, H; 1913:130, 132 [key].—
Schmitt, 1934:69.—Hiulton, 1915a:68;
1915b:204; 1920:93; 1939a:32.
Nymphopsis spinosissimus: Hedgpeth, 1939:
461-463, fig. 2m-—r [Neotype].— Hilton,
1939b:72, fig. 3; Hedgpeth, 1941:256
[key], pl. 11.
Nymphopsis spinosissima: Hilton, 1942b:
278, 300-302, fig. 44; 1943:98.—Hedg-
peth, 1951:110.—Stock, 1955:253.—
Hedgpeth, 1964:206, fig. 94f.
Material examined. —Topotypes (Am-
mothella spinosissima Hall, 1912), 2 fe-
males, Laguna Beach, California, NW of pier
in tide pool, coll. Metz, Jul, 1912; Neotype
(Nymphopsis spinosissimus: Hedgpeth,
1939), male, Corona Del Mar, California,
coll. G. E. MacGinitie, 27 Mar 1938, in 22-
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
31 meters; also other specimens reported on
by Hilton and Hedgpeth.
Remarks. — The type specimen described
by Hall in 1912 was not among the Cole
and Hall types and presumably has been
lost. Hedgpeth’s male neotype therefore re-
mains valid for this easily recognized species.
Hall’s topotypes can not be regarded as type
material as they were collected subsequent
to the original description. Hedgpeth’s
(1939) figures of this species are adequate
to allow its differentiation from the other
known California species, N. duodorsospi-
nosa Hilton which has two prominent dor-
sal trunk tubercles instead of the three found
on this species.
The distribution of this shallow-water (O-
31 m) species now extends from the state
of Washington to southern California, but
it has not been found yet along the west
coast of Mexico where its place is apparently
taken by N. duodorsospinosa.
Anoplodactylus californicus Hall
?Nymphon dubium Nicolet in Gay, 1849:
307; 1854: fig. 10.
Anoplodactylus californicus Hall, 1912:91-
93, figs. 49, 52D, F, I, J; 1913:129-130,
pl. 4, figs. 14-16.—Hilton, 1915a:69;
1915b:201, 205; 1916:27 [text]; 1920:93;
1939a:29.—Marcus, 1940:40 [key].—
Hilton, 1942b:277 [list], 286-288, pl. 38;
1942d:72.
Anoplodactylus californiensis [sic]: Hedg-
peth, 1941:257 [key], pl. 11.
Anoplodactylus portus Calman, 1927:405-—
408, fig. 103.—Sawaya, 1950:70 [key].—
Stock, 1954:128; 1955:238—-239; 1958a:
4: 1958b:140-141; 1962:218.—Lipkin
and Safriel, 1971:9.—Stock, 1975:1052-
1053, fig. 4lb-e.—Child, 1975:201.—
Birkeland et al., 1976:158.—Child, 1978:
133-144, figs. 1-4; 1979:58-59.—Stock,
1979:15.—Child, 1982:373.
Anoplodactylus robustus Hilton, 1939a:28-
29 [non A. robustus (Dohrn, 1881)];
VOLUME 100, NUMBER 3
1942a:288-291, pl. 39; 1942d:72.—
Hedgpeth, 1941:257 [key].
Anoplodactylus carvalhoi Marcus, 1940:50—
54, fig. 3a—k.— Hedgpeth, 1943:46; 1948:
230-232, fig. 30e-g.—Sawaya, 1950:70
[key].—Bourdillon, 1955:592-593, pl. 1,
figs. 9-10.
Anoplodactylus projectus Hilton, 1942c:45-—
47, fig. 3; 1942d:72.
[non Anoplodactylus portus Calman var.
chilensis Hedgpeth, 1961:5-7, fig. 2].
Material examined.—Topotypes (Ano-
plodactylus californicus Hall, 1912), 2 males
with eggs, 1 male, 5 females, Laguna Beach,
California, NW of pier, coll. Hall and Metz,
16 Jul 1912. Also many specimens identi-
fied as A. portus from California.
Remarks.—I\t is unfortunate that Hall
carried his wide use of geographical epithets
to this species because it is now known to
be pantemperate-pantropical in distribu-
tion, predominantly in shallow water. It is
not surprising that this species turns out to
be widely known, first, because of Hall’s
poor figures of his type which render it al-
most impossible to compare with other
species, and second, because Cole was pre-
sumably the only other pycnogonid spe-
clalist to see the specimens, if he indeed
examined them at all. Cole never published
on pycnogonids of California after Hall’s
1912 description of the species and he was
no longer working on pycnogonids by the
time Calman (1927) published this species
as A. portus.
The species is easily recognized by its ro-
bust appearance, very hirsute oviger stri-
gilis, tiny distally pointing cement gland
tube, long sex pore tubercles on the second
coxae in male specimens, and the peculiar
alar processes placed proximoventrally on
the female proboscis. While none of these
characters 1s unique to this species, no oth-
ers, at least along the California coast, share
this set of characters.
The description and figures of A. portus
Calman var. chilensis Hedgpeth (1961:5—7,
22)
fig. 2) do not fit well with this species. The
major differences occur in the oviger, a rath-
er stable character in this genus as it is in
most Pycnogonida. In Hedgpeth’s variety,
the oviger segment lengths are different from
typical specimens and the terminal segment
is not heavily setose. The female lacks the
typical palp buds but has the proboscis alar
processes, both of which are quite promi-
nent in this species. The propodal heel has
one major spine while both Hall’s and Cal-
man’s type specimens have two heel spines.
These differences are not in themselves con-
clusive proof that the Chilean material is a
different species, particularly without ex-
amining the Chilean specimens first hand,
but it would not be surprising if, with more
material, this variety were eventually to be
called A. chilensis.
Callipallene californiensis (Hall)
Pallene californiensis Hall, 1913:131, 133-
135, pl. 4, figs. 9-13.— Hilton, 1915a:67;
1916b:465, fig. 6; 1920:93.
Palene [sic] californiensis: Hilton, 1915b:
204; 1939a:29.
Callipallene californiensis: Hedgpeth, 1941:
257 [key], pl. 11.—Hilton, 1942b:277,
281-282, pl. 36; 1942c:38.—Correa,
1948:6 [key].—Hedgpeth, 1964:204 [key].
Callipallene solicitatus Child, 1979:44—46,
ne WS
Material examined.—Holotype (Pallene
californiensis Hall, 1913), male, Laguna
Beach, California, Mussel Point, Cowards
Cove, under stones in channel, coll. Hall,
15 Aug 1912; Holotype (Callipallene soli-
citatus Child, 1979), male, Isla del Espiritu
Santo, Bahia Ballenas, Baja California (Gulf
side), Mexico, coll. J. L. Barnard, 28 Nov
1971; also, the Mexican and Panamanian
paratypic series, and twenty specimens col-
lected by W. A. Hilton between 1914 and
1934 in southern California.
Remarks. — This species has been obscure
in the literature with most authors subse-
556
quent to Hall accepting his description and
figures (and sometimes reproducing the fig-
ures) without adding further to the descrip-
tion or information. It has not been com-
monly found in California and until the
species was redescribed (as C. solicitatus
Child, 1979), no adequate figures of all di-
agnostic characters had been published.
The species has a relatively short neck, a
very setose and spinose propodus bearing
four or five heel spines, a large setose apoph-
ysis on the male fifth oviger segment, and
many small chela finger teeth.
In its more southern range (Panama), as
far as its range is known, it apparently pre-
fers subtidal depths and has been found on
settling plates in 8 to 10 meters and on sub-
tidal Balanus colonies. In California and
Mexico, most of the captures appear to be
intertidal with some listed as “‘on rocks.”’
Acknowledgments
I wish to thank personnel of the Smith-
sonian Oceanographic Sorting Center for
finding this misplaced bottle of specimens
and for bringing it to my attention. The
specimens are now redeposited in the Na-
tional Museum of Natural History under
the old catalog numbers of the United States
National Museum. I am grateful for the re-
view of this paper by Dr. T. E. Bowman and
for his comments towards its improvement.
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VOLUME 100, NUMBER 3
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the summer of 1914, at Laguna Beach.—Po-
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67-70.
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1916a. The life history of Anoplodactylus
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. 1920. Notes on Pacific Coast pycnogonids. —
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from the shores of California.— Pomona Journal
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. 1939b. A collection of pycnognids [sic] from
Santa Cruz Island.—Pomona Journal of Ento-
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. 1942a. Pycnogonids from the Allan Hancock
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1942b. Pantopoda (continued). II. Family
Callipallenidae.— Pomona Journal of Entomol-
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1942c. Pycnogonids from Hawaii.—Occa-
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Honolulu, Hawaii 17(3):43-55, 10 figs.
. 1942d. Pycnogonids from the Pacific. Family
Phoxichilidiidae Sars 1891.—Pomona Journal
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[publ. 1854].
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and of the Mediterranean coasts of Israel. Con-
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Entomologische Berichten, Amsterdam 22(1 1):
218-219.
558 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
. 1975. Pycnogonida from the continental shelf, shallow waters of Japan.— Publications of the
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—. 1979. Pycnogonida from the mediolittoral P BY
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Utinomi, H. 1971. Records of Pycnogonida from
PROC. BIOL. SOC. WASH.
100(3), 1987, pp. 559-577
FURTHER RECORDS OF MARINE ISOPOD
CRUSTACEANS FROM THE CARIBBEAN
Brian Kensley
Abstract. — Five species of isopods are described from Belize. The anthurid
Mesanthura bivittata, new species, is characterized primarily by its double-
barred dorsal color pattern. The idoteid Miratidotea bruscai, new genus and
species, is characterized chiefly by possessing a pleon consisting of two complete
and two incomplete pleonites plus pleotelson. The cirolanid Anopsilana jonesi,
new species, can be distinguished by its non-projecting pentagonal frontal lam-
ina and a color pattern having a dark central area on the pereonites. Eurydice
personata (family Cirolanidae), often confused with E. convexa and E. littoralis,
is characterized by the possession of a slender, lanceolate frontal lamina. The
new cirolanid genus and species Xylolana radicicola, taken from dead mangrove
roots, features a fused rostrum and frontal lamina, a strongly projecting clypeus,
an overlapping fourth pleonite, distally articulating copulatory stylet, and a
produced uropodal sympod. The sphaeromatid Paraleptosphaeroma glynni,
described from Pacific Panama, is recorded for the first time from the Carib-
bean.
Anyone following the literature on Carib-
bean marine isopods will have noted a series
of my papers, appearing at irregular inter-
vals, and often covering material from Be-
lize. No doubt the question has been raised:
why this dribble of short papers, instead of
a single comprehensive work? In compiling
a guide to the marine isopods of the Carib-
bean, I am attempting to deal with as many
species as possible, and as new material be-
comes available, it has been necessary to
publish new species before including them
in the abovementioned guide. A further fac-
tor has contributed to this plethora of titles,
viz. continued sampling in the area around
Carrie Bow Cay, Belize, over the past nine
years. With such concentrated sampling, it
is inevitable that rarer species eventually are
collected, and that as more and more spe-
cialized habitats are investigated, new
species will be revealed. As an example of
the latter case, see the description in this
paper, of a new genus and species, collected
less than one hundred meters from the field
laboratory on Carrie Bow Cay. This was the
first time, however, that the root-mat of the
seagrass Syringodium had been sampled. No
doubt similar concentrated collecting in
other areas of the Caribbean will reveal yet
more undescribed forms, and this trickle of
taxonomic papers will perforce continue.
Family Anthuridae
Mesanthura bivittata, new species
Figs, 1.2
Material. —Holotype, USNM 221718,
ovig. 2 TL 7.8 mm. Paratypes, USNM
221719, 1 non-ovig. 2 TL 7.8 mm, 2 6 5.2
mm, 6 immature specimens 3.0—4.8 mm.
Twin Cays, Belize, in red mangroves, 1-2
m, coll. G. Hendler 28 Mar 1980.
Description. —Non-ovigerous female:
Proportions C <1=2>3<4=5=6>
7. Cephalon with low rounded rostrum; large
well pigmented dorsolateral eyes. Articular
hollow between pereonites 1 and 2, and 2
and 3. Pleonites 1—5 fused, subequal to per-
560
Fig. 1.
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Mesanthura bivittata, ovigerous female: A, Entire animal in dorsal view; B, Antenna; C, Antennule;
D, Mandible; E, Maxilliped; F, Pereopod 1; G, Pereopod 2; H, Pereopod 7.
eonite 7 in length; pleonite 6 dorsally free,
with small middorsal notch in posterior
margin. Telson widest at midlength, pos-
terior margin broadly rounded and bearing
numerous elongate simple setae.
Antennular peduncle of 3 articles, basal
article longest and broadest; flagellum of 3
articles, terminal article bearing 3 aesthe-
tascs. Antennal peduncle of 5 articles; fla-
gellum of 4 short setose articles. Mandibular
palp of 3 articles, terminal article with 8 or
9 spines; incisor of 3 sclerotized cusps; lam-
ina dentata with 4 serrations. Maxilliped
lacking endite; palp of 3 articles, terminal
article narrower than preceding article,
bearing 3 stout fringed setae on medial mar-
gin. Pereopod 1, carpus distally rounded;
propodus expanded, palm with rounded lobe
in proximal half bearing few marginal setae;
unguis subequal to rest of dactylus in length.
Pereopod 2, carpus short, triangular, lacking
free anterior margin; propodus not expand-
VOLUME 100, NUMBER 3
\
Fig. 2. Mesanthura bivittata, ovigerous female: A, Uropodal endopod and sympod; B, Uropodal exopod; C,
Pleopod 1. Male: D, Mandible; E, Antennule, aesthetascs indicated by insertions only; F, Pereopod 1; G, Pleo-
pod 2.
ed, bearing single short posterodistal spine.
Pereopods 4—7, carpus with anterior margin
shorter than posterior, latter with single sen-
sory spine; propodus with row of short spi-
nules in distal half of posterior margin plus
single sensory spine. Pleopod 1, exopod
operculiform, subequal in length but 3 times
wider than endopod; latter with 10 distal
plumose setae. Uropodal exopod with notch
in distal margin; most of margin bearing
elongate setae; endopodal distal margin
broadly rounded, bearing numerous elon-
gate setae.
Male: Antennular flagellum of 7 articles,
first 6 bearing distal band of aesthetascs.
Mandible consisting of palp attached to short
featureless basal structure. Pleopod 2, en-
dopod with copulatory stylet articulating in
proximal half, slender, cylindrical, distally
rounded, reaching well beyond apex of ra-
mus.
Color pattern. —Similar in male and fe-
male. Large dark-brown chromatophores
arranged in broad patch on cephalon
stretching from eyes to posterior margin,
with narrow elongate gap along midline;
pereonites 1—6 with 2 broad elongate bands
separated by fairly wide gap, bands touching
anteriorly and/or posteriorly; pereonite 7
and fused pleonites 1—6 each with 2 widely
562
separated bands. Pleonite 6 with 2 small
patches. Telson with elliptical patch becom-
ing posteriorly obsolete. Uropods with broad
patch on each.
Remarks.— Mesanthura bivittata is the
eighth species of the genus to be recorded
from the Caribbean and/or the Florida Keys.
Mesanthura looensis Kensley and Schotte,
1987, like the present species, possesses eight
spines on the third article of the mandibular
palp. Ovigerous at 10 mm, the Floridean
species does not have the distinctive dou-
ble-bar pattern seen in M. bivittata. Mes-
anthura pulchra Barnard, which has 10
spines on the mandibular palp, while having
pigment patches with a central gap, does not
have the well defined double-bar. The re-
maining five species have radically different
color patterns and different numbers of
mandibular spines.
Etymology.—The specific name is de-
rived from the Latin, meaning ‘two rib-
bons,’ and refers to the double band of pig-
ment that characterizes this species.
Suborder Valvifera
Family Idoteidae
Miratidotea, new genus
Diagnosis. — Antennal flagellum of single
clavate article. Maxillipedal palp of 4 arti-
cles. Pleon consisting of 2 complete pleo-
nites, 2 incomplete pleonites, plus pleotel-
son. Uropods uniramous.
Type species. — Miratidotea bruscai, new
species. Gender: masculine.
Remarks.— Brusca (1984) placed the
subfamily Idoteinae (to which the present
genus and species clearly belong) on a sound
taxonomic footing, with a phylogenetic
analysis of the group. Using in particular
the form of the pleon, this author presented
a schematic plan (fig. 3) of the possible phy-
logenetic pathways that gave rise to the ap-
proximately 21 known genera of the
subfamily. In this plan Brusca postulated
several stages for which no forms are yet
known. The present species clearly fits one
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
of these postulated forms in the Lineage A,
with its pleon consisting of 2 complete and
2 incomplete pleonites plus pleotelson. This
pleonal formula places the present species
phylogenetically close to Cleantioides Kens-
ley and Kaufman, and indeed the similar-
ities between these two genera, in antennal,
mouthpart, pereopodal, and uropodal
structure, are marked. Apart from the
pleonal structure and minor proportional
differences in the appendages, the main dif-
ference between Miratidotea and Clean-
tioides occidentalis (Richardson) lies in the
pleotelson. In the latter, the dorsal margin
of the planiform posterior pleotelson con-
sists of two broadly rounded lobes (in dorsal
view), barely projecting when seen in lateral
view. In Miratidotea, these lobes are sub-
acute, and project markedly in lateral view.
Etymology. —The generic names derives
from the Latin ‘miratio,’ a surprise, plus
‘idotea,’ the frequently-used sufhix taken
from the family name Idoteidae.
Miratidotea bruscai, new species
Figs. 3, 4
Material. —Holotype USNM 221720,
ovig. 2 TL 13.0 mm; paratypes USNM
MI WAN A Owns QING MOLY aro, Uil.5 mans
Carrie Bow Cay, Belize, from root-mat of
seagrass Syringodium filiforme in 1.5 m
depth, coll. B. Kensley, 11 Dec 1986.
Description. —Ovigerous female: Body
elongate-cylindrical, about 6 times longer
than wide. Anterior margin of cephalon with
tiny midline notch in dorsal view. Eyes dor-
solateral, reniform. Sparsely scattered setae
over most of dorsal integument, but dense
on posterior margin of pereonite 7 and coxa
7, and anterolateral pleon. Pereonite 1 and
coxa fused; coxae 2—4 about half lateral
length of pereonite, elongate-oval in shape;
coxae 5—7 produced posteriorly into trian-
gular subacute lobe. Pleonites 1 and 2 com-
plete, subequal, ventrally tapering to nar-
rowly-rounded free margin; pleonite 3
incomplete, with narrowly-rounded free
VOLUME 100, NUMBER 3
563
Fig. 3. Miratidotea bruscai, ovigerous female: A, Holotype in dorsal view; B, Whole animal in lateral view;
C, Antennule; D, Antenna; E, Left and right mandibles; F, Maxilla 1; G,Maxilla 2; H, Maxilliped, with palp
shown separately; I, Uropod.
ventral margin; pleonite 4 incomplete, lack-
ing free ventral margin. Posterior planiform
area of pleotelson with dorsal margin armed
with 2 triangular submedian subacute pos-
teriorly-directed lobes; ventral margin
broadly rounded and forming posterior
margin of pleotelson.
Antennule with 3-articled peduncle, basal
article broadest and longest; article 2 tri-
questrous, distally hollowed for articulation
of article 3; flagellum of single article about
half length of peduncle article 3, bearing 3
distal aesthetascs. Antenna with peduncle
of 5 articles, article 2 produced ventrally
into broad lobe; flagellum of single clavate
article bearing numerous ventrodistal setae.
Mandible lacking palp; incisor of 4 cusps;
lacinia of 3—4 cusps; 8 spines in spine-row;
molar stout, distally with flattened circular
surface. Maxilla 1, inner ramus with 3 distal
fringed setae; outer ramus with about 10
distal spines, some of which serrate. Maxilla
564 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
TAN
Fig. 4. Miratidotea bruscai, ovigerous female: A, Pereopod 1; B, Pereopod 2; C, Pereopod 3; D, Pereopod
4; E, Pereopod 5; F, Pereopod 6; G, Pereopod 7.
2, both lobes of outer ramus bearing nu-
merous fringed spines; inner ramus with
simple and sparsely fringed spines along
mesial margin. Maxillipedal endite bearing
3 coupling hooks, several distal fringed
spines; palp of 4 articles, article 2 longest
and broadest, terminal article short. Pereo-
pod 1, merus with 3 posterodistal spines;
carpus triangular, with 4 posterodistal
spines; propodus somewhat expanded, with
7 spines on posterior margin and numerous
smaller spines on mesial surface; unguis
about half length of rest of dactylus. Pereo-
pods 2 and 3 similar, longer than pereopod
1; merus with 2 anterodistal fringed spines;
carpus roughly rectangular, with 5 sensory
spines and one fringed spine on posterior
margin; propodus elongate-rectangular, with
6 or 7 spines on posterior margin. Pereopod
4 short, equal in length to ischium and basis
of pereopod 3; ischium with 3 distal spines;
merus with 9 spines on posterodistal sur-
VOLUME 100, NUMBER 3
face; carpus with 8 spines on posterodistal
surface; propodus with 7 spines on poster-
odistal surface; dactylus reduced to length
of a spine but distally corneous. Pereopod
5, basis, ischium, and merus with elongate
setules; carpus with 3 spines on posterior
margin; propodus with 2 posterior spines;
dactylus strongly hooked, approaching
biunguiculate state. Pereopod 6 with elon-
gate setules on basis, ischium, and merus;
merus with 2 posterodistal spines; carpus
with 4 posterodistal spines; propodus with
5 posterodistal spines; dactylus hooked,
biunguiculate. Pereopod 7, elongate setules
on basis, ischium, and merus; posterior
margin of carpus with 2 separate, and clump
of 4 spines on posterior margin; propodus
with 3 clumps of spines on posterior margin;
dactylus hooked, biunguiculate. Uropod
with single ramus slightly less than half
length of sympod, latter with elongate
fringed seta at outer distal angle.
Color pattern. — Dorsal integument over-
all red-brown, with 6 darker longitudinal
stripes running from pereonite | onto pleon.
Cephalon with faint reticulation of color.
Posteroventral area of pleotelson with 4 faint
darker rays of brown.
Etymology. —The species is named for Dr.
Richard C. Brusca, as a small recognition
of his valuable contributions to isopod re-
search.
Family Cirolanidae
Anopsilana jonesi, new species
Figs. 5, 6
Material. —Holotype, USNM 221722, 6
TL 6.2 mm, paratypes, USNM 205679, 3
6 5.6-6.7 mm, 5 non-ovig. 2, 5.0—6.5 mm,
6 juvs. 2.2—4.5 mm, Sapodilla Lagoon, Sit-
tee River, Belize, amongst red mangrove
roots, coll. K. Fauchald, 9 Dec 1986.—
Paratypes, USNM 205680, 2 6 6.0-—7.5 mm,
4 ovig. 2 5.4-6.0 mm, 17 juvs. 2.0—5.2 mm,
Salt Creek, north of Dangriga, Belize, be-
tween red mangroves in 1 m, 31%0o, 32°C,
coll. M. Jones, 16 May 1977.—Paratypes,
565
USNM 205681, 4 4, 5.1-7.4 mm, 6 2, 5.0-
7.0 mm, Anderson Lagoon, Sittee River,
Belize, from amongst barnacles and mussels
on red mangrove roots, coll. K. Fauchald,
11 Dec 1986.
Description. —Male: Body 2.6 times long-
er than wide, widest at pereonites 5 and 6.
Cephalon somewhat sunken into pereonite
1, with small rostral point between anten-
nular bases; eyes large, dorsolateral; three
low dorsal tubercles near posterior margin;
frontal lamina narrow-pentagonal, distal
margin not projecting. Pereonite 1 with 4—
6 low tubercles near posterior margin; pere-
onites 2—7 with several low submedian
ridges. Coxae 2 and 3 narrow, posteriorly
rounded; coxae 4—7 broader, posteroven-
trally somewhat produced to subacute apex.
Pleonites 1-3 with free ventral margins
rounded; pleonite 4 lacking free ventrolat-
eral margins, with 2 low submedian tuber-
cles on posterior margin. Pleotelson broadly
triangular, posterior margin rounded, bear-
ing 8—10 apical spines and numerous setae.
Antennular peduncle of 2 articles, basal
article broader but slightly shorter than dis-
tal article; flagellum of 9-12 articles, distal
7-9 articles each bearing 1-3 aesthetascs.
Antennal peduncle of 5 articles, 3 basal ar-
ticles short, together equal in length to ar-
ticle 4, articles 4 and 5 subequal in length;
flagellum of about 30 articles, reaching pos-
teriorly to anterior of pereonite 5. Mandib-
ular palp directed anteriorly, of 3 articles,
article 2 longest, bearing about 12 fringed
spines, article 3 with comb of about 16 spines
on outer margin; incisor of 3 strong scler-
otized cusps; lacinia having 5 cusps; 4 spines
in spine-row; molar narrowly triangular,
with row of short marginal spines. Maxilla
1, inner ramus with three stout setae; outer
ramus with about 10 sclerotized spines,
some of which spinulose, plus | central seta.
Maxilla 2, 2 lobes of outer ramus short, out-
er bearing 4 spines, inner with 7—11 spines;
inner ramus with 6 distal simple spines and
6-8 fringed proximal spines. Maxilliped,
endite short, barely reaching beyond basal
566
| De
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
i,
SS
INN
XN
>
* . SSieace
x
\
Z
Anopsilana jonesi, male: A, Entire animal in dorsal view; B, Entire animal in lateral view; C, Frontal
lamina and clypeus; D, Antennule; E, Maxilla 1; F, Maxilla 2; G, Maxilliped; H, Mandible; I, Uropod; J,
Antenna.
palp article, with 2-3 distal fringed setae
and | or 2 coupling hooks; palp of 5 articles,
article 3 longest and broadest; articles 3-5
each with group of mediodistal spines. Pe-
reopods increasing in length posteriorly. Pe-
reopod |, merus with row of 6 blunt spines;
carpus short, with almost no free anterior
margin, bearing single sensory spine pos-
terodistally; propodus with spine at mid-
length and single posterodistal spine. Pereo-
pod 2 and 3 similar, ischium with 3
posterodistal spines and single large antero-
VOLUME 100, NUMBER 3
567
Fig. 6. Anopsilana jonesi, male: A, Pereopod 7; B, Pereopod 1; C, Pereopod 2; D, Pleopod 1; E, Pleopod 2;
F, Pleopod 3; G, Pleopod 4; H, Pleopod 5.
distal spine; merus with row of 8 blunt spines
on posterior margin and 3 anterodistal sen-
sory spines; carpus about half length of pro-
podus, with 3 posterodistal spines; propo-
dus with single spine at midlength and
another distally on posterior margin; unguis
about half length of rest of dactylus. Pereo-
pods 4—7 similar; ischium with 4 pairs of
spines on posterior margin, 3 spines antero-
distally; merus with 2 groups of spines on
posterior margin, group of sensory and ser-
rate spines anterodistally; carpus rectangu-
lar, with pair of spines on posterior margin,
plus dense cluster of sensory and serrate
spines along distal margin; propodus elon-
gate-rectangular, with few spines along pos-
terior margin. Pleopod 1, endopod paral-
lel-sided, narrower than ovate exopod.
Pleopod 2, copulatory stylet apically acute,
reaching slightly beyond, and articulating at
base of, endopod; exopod ovate. Pleopods
3—5, endopods narrower and shorter than
568
exopods, lacking marginal setae; exopods
broadly ovate, biarticulate, with marginal
setae. Uropodal sympod produced along
medial margin of endopod, bearing few dis-
tal setae; endopod widening distally, reach-
ing beyond pelotelsonic apex, with about 12
marginal spines plus numerous setae; exo-
pod narrow, lanceolate, apically acute, with
10-12 marginal spines.
Female: Essentially similar to male, but
lacking tubercles and ridges on cephalon,
pereonites, and pleonite 4. Setae on anten-
nal flagellum shorter than in male.
Color pattern.—Similar in both sexes.
Dorsal integument of cephalon and pereo-
nite 1 with scattered brown chromato-
phores; pereonites 2—7 with solid central
area, laterally with scattered chromato-
phores; pleonites 1—3 with middorsal patch
of pigment; pleonite 4 and basal pleotelson
with 2 patches, plus large irregular central
blotch on pleotelson; uropodal endopod with
irregular central patch.
Remarks. —The approximately nine
known species of Anopsilana (see Bruce
1986) can be grouped into those occurring
in freshwater caves (generally unpigmented
and eyeless), and those pigmented forms
such as A. browni (Van Name), A. luciae
(Barnard), and A. oaxaca Carvacho and
Haasmann, which occur in estuaries. The
present species belongs to the latter group,
and indeed, was caught along with single
specimens of A. browni from Anderson La-
goon in the Sittee River and from Salt Creek.
The two species may be separated by three
easily seen features. 1. The color pattern,
that of A. browni lacking the solid middorsal
area on the pereon. 2. The frontal lamina,
which in A. browniis distally broadly round-
ed and strongly projecting. 3. The strong
double tubercles on the cephalon and rows
of rounded tubercles on the pereonites and
pleonites of male A. browni, contrasted with
the three low tubercles on the cephalon and
the low ridges on the pereonites of male A.
jonesi.
Etymology. —The species is named for Dr.
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Meredith Jones, of the Department of In-
vertebrate Zoology, Smithsonian Institu-
tion, who collected the first specimens of
this species along with numerous other iso-
pods from localities in the Caribbean.
Eurydice personata, new species
Figs. 7, 8
Material.— Holotype, USNM 211436, 1
6, 6.0 mm, paratypes, USNM 128314, 19
6, 5.0-6.0 mm, 4 ovig. 2, 5.1-6.4 mm, 6
non-ovig. 2, Mona Island, Puerto Rico, 40-—
50 ft, coll. R. Menzies and P. Glynn, 10
May 1966.—USNM 211435, 63 6, 3.5—4.5
mm, 1 ovig. 2, 5.0 mm, 95 non-ovig. 8,
Bahamas north of Bimini Is., 1-2 m, coll.
M. Jones, 22 Aug 1962.—USNM 111388,
2 ovig. 2, 4.1 mm, Bahamas, surface plank-
ton tow at night, coll. J. McCain, 14 Jul
1964.—USNM 60689, 3 4, 3.5—4.5 mm, 2
non-ovig. 2, Bermuda, 27 Sep 1933.—
USNM 65871, 1 non-ovig. 2, Bigie Bay,
Haiti, 23 Apr 1930.—USNM 221547, 38 6,
4.5—4.8 mm, 6 ovig. 2, 4.9-5.8 mm, Grande
Cay, Cuba, 19 Apr 1937.—USNM 86369,
7 juvs., off Miami, Florida, Jun 1942.—
USNM 225445, 1 ovig. 2, 5.6 mm, off Geor-
gia, 27 m, 14 May 1981.—USNM 225440,
1 non-ovig. 2, off Georgia, 18 m, 28 Jul
1981.—USNM 225448, 1 juv., off South
Carolina, 34 m, 27 Jul 1981.—USNM
225450, 1 6, 1 non-ovig. 2, 1 juv., off Geor-
gia, 26 m, 12 Aug 1981.
Description. —Male: Body about 4 times
longer than greatest width. Coxae of pereo-
nites 2—7 ending in acute denticle. Pleonites
2-5 posteroventrally acute. Pleotelson wid-
er than middorsal length, with anterior hol-
low; posterior margin between notches
slightly convex, with 2 pairs of spines, inner
pair longer than outer, between 5 and 6 times
longer than wide. Cephalon lacking ros-
trum; frontal lamina narrow, lanceolate,
distally acute; clypeus broadly triangular,
distally acute and projecting anteroventral-
ly; eyes large, lateral, reaching to ventral
surface.
VOLUME 100, NUMBER 3 569
M
Fig. 7. Eurydice personata: A, Adult in lateral view; B, Antennule; C, Antennal peduncle; D, Antennal
flagellar article enlarged; E, Pleotelson; F, Mandible; G, Maxilla 1; H, Maxilla 2; I, Maxilliped; J, Pereopod 1;
K, Pereopod 7; L, Male pleopod 1; M, Male pleopod 2.
570
Antennular peduncle with article 2 at right
angle to article 1; flagellum of 6 articles,
article 2 longest, bearing series of aesthe-
tascs. Antennal peduncle of 4 articles, ar-
ticle 3 with 4—6 slender spines distally; fla-
gellum reaching posteriorly to level of
pereonite 7, articles with plicate organ about
half length of article. Mandibular palp of 3
articles, article 2 longest, article 3 with comb
of 10 distal setae. Maxilla 1, outer ramus
with 12 distal spines, some being spinulose.
Maxilla 2, inner ramus short, truncate, with
4 proximal fringed setae and 5 distal simple
setae; inner and outer lobes of outer ramus
with 3 and 5 fringed setae respectively.
Maxillipedal endite reaching to middle of
palp article 2, with distal fringed setae
reaching to palp article 4; palp with article
3 widest and longest. Pereopod 1-3 similar,
prehensile; ischium with anterodistal exten-
sion bearing single spine, 2 acute and 2 blunt
spines posterodistally; merus with single
short anterodistal spine, | acute and 5 blunt
spines on posterior margin; carpus lacking
free anterior margin, with 2 posterodistal
spines; propodus with 3 spines on posterior
margin plus stout posterodistal specialized
spine. Pereopods 4—7 similar, becoming
more elongate posteriorly, ischium, merus,
carpus, and propodus each bearing clumps
of sensory spines on anterior and posterior
margins. Pleopod 2, endopod with copula-
tory stylet clavate, distally blunt, reaching
by one-fifth of its length beyond ramus.
Uropodal exopod ovate, five-sixths length
of endopod; latter with broadly truncate dis-
tal margin.
Female: Essentially similar to male, but
body proportionally slightly broader.
Color pattern. —Entire dorsum except
posterior three-fourths with dense brown
pigmentation; sternites, pereopods (except
propodi and dactyli), and pleopodal sym-
pods pigmented (based on Georgia and
South Carolina material).
Etymology.—The specific name is de-
rived from the Latin personatus, meaning
wearing a mask, and refers to the fact that
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
this species has been masquerading as either
of two other species of Eurydice.
Remarks. —Examination of the material
of three western Atlantic species of Eurydice
in the Smithsonian’s collections, and pe-
rusal of the literature, let to some confusion
in my attempts to separate these species.
Closer observation revealed that an unde-
scribed species had previously been mis-
identified either as E. convexa or E. littor-
alis. Eurydice personata, the new species,
while superficially very similar to E. con-
vexa Richardson, 1900 (=E. littoralis
Moore, 1901), can most easily be distin-
guished by its slender lanceolate frontal
lamina. The following table provides fur-
ther means for separating the three species
occurring in the Florida Keys and Carib-
bean.
E. convexa |E. piperata | E. perso-
Richard- Menzies &| anata,
son, Franken- new
1900 berg, species
1966
Frontal truncate, truncate, lanceolate
lamina faintly faintly acute
bilobed bilobed
Pleotelson
apex truncate faintly
to convex
faintly
convex
spines 4, mod- 4, very 4, elon-
erate short gate
length
6 anten- ¥, length ¥, length ¥%, length
nal of of of
plicate article article article
organ
Xylolana, new genus
Diagnosis. — Frontal lamina and rostrum
fused, broad, separating antennular bases.
Clypeus conical, projecting. Antennular pe-
duncle of 3 articles; antennal peduncle of 5
(24) articles, articles 3—5 subequal in length.
Mandibular palp directed anteriorly. Max-
VOLUME 100, NUMBER 3
8
Fig. 8. Eurydice personata: A, Frontal lamina; B, Apex of pleotelson.
illipedal endite reduced, lacking coupling
hooks. Pereonite | twice length of pereonite
2. Pereopods 1—3 with meri not anterodis-
tally produced. Pereopods 4—7, articles not
flattened, lacking natatory setae. Short penes
SII
present on sternite of pereonite 7. Pleopod
2 in male with copulatory stylet articulating
in distal half of mesial margin of endopod.
Pleopods 3-5, exopod biarticulate; endopod
lacking marginal setae. Pleonite 5 lacking
1/2
free lateral margin, overlapped laterally by
pleonite 4. Uropodal sympod produced
along mesial margin of endopod.
Type species.—Xylolana radicicola, new
species.
Etymology.—The generic name is de-
rived from the Greek word xylon for wood,
referring to the woody habitat of the animal,
plus the commonly-used sufhx ‘lana,’ de-
rived from Ciro/ana, originally an anagram
of Carolina.
Remarks.— Discovery of an apparently
specialized and highly adapted species such
as the one under discussion, immediately
exposes a gap in our knowledge of the tax-
onomy of the cirolanids. (With few excep-
tions, this gap is present in most of the ma-
jor isopod groups.) With no phylogenetic
analysis at the generic level, there is no way
of knowing which characters are apomor-
phic and which plesiomorphic. Separation
of genera, while probably reflecting the phy-
logenetic relationships fairly well, is thus a
shaky and somewhat subjective process.
Well defined and long-understood genera
such as Eurydice will present little problem,
but with unusual and adapted forms such
as the present species, generic placement be-
comes very difficult. Has a projecting clyp-
eus evolved more than once? Is the condi-
tion with pleonite 4 overlapping pleonite 5
apomorphic? These and other similar ques-
tions arise in trying to place the present ma-
terial.
The projecting clypeus would indicate af-
finity with the Eurydicinae, while the lateral
overlapping of pleonite 5 by pleonite 4 would
indicate the Cirolaninae. Features such as
the fusion of the rostrum and frontal lam-
ina, the medially-articulating copulatory
stylet, and the lack of marginal setae on the
endopods of pleonites 3—5, however, all in-
dicate a stronger affinity with the Eurydi-
cinae. Within the latter subfamily, the pres-
ent species does not agree with the
definitions of any of the genera. Using the
available keys to the Cirolanidae also proves
unsatisfactory. For example, using Bruce’s
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
1986 key, the present species runs down to
Eurylana, from which it differs in several
features. There seems to be no choice but
to describe a new genus for this material, in
which small size and choice of habitat might
well have dictated several of the specialized
features.
Xylolana radicicola, new species
Material. — Holotype, USNM 211437, 2
2.6 mm, paratypes, USNM 211438, 1 ovig.
? (cephalon plus anterior 4 pereonites only),
1 non-ovig. 2 (cephalon missing), | non-
ovig. 2, 1.9 mm (SEM specimen), from dead
in-situ red mangrove prop roots, 1 m, Twin
Cays, Belize, coll. K. Rutzler, Feb 1987.—
Paratype, USNM 211439, non-ovig. 2, 3.3
mm, from dead in-situ red mangrove prop
roots, 1 m, Twin Cays, Belize, coll. B. Kens-
ley, 12 Dec 1986.
Description. —Male: Body about 4 times
longer than greatest width (at pereonites 4
and 5). Cephalon sunken into pereonite 1,
with large well pigmented eyes; area be-
tween posterior margin and line joining pos-
terior margins of eyes somewhat inflated;
antennular bases separated by broad, flat-
tened, fused rostrum and frontal lamina.
Clypeus narrowly conical, projecting dis-
tally. Pereonite 1 about twice length of pe-
reonite 2. Coxae of pereonites 2 and 3 pos-
teriorly rounded; of pereonites 4—7 becom-
ing progressively more produced and
elongate posterodistally. Pleonites 1-3 short;
pleonite 4 laterally broad, overlapping
pleonite 5 and base of pleotelson, with
oblique row of plumose setae; pleonite 5
short, lacking free lateral margin. Pleotelson
basally broad with faint rounded middorsal
ridge, becoming abruptly narrowed from
base of uropodal sympod, tapering slightly
to broadly rounded posterior margin.
Antennular peduncle of 3 articles, article
3 longest; flagellum of 6 articles, articles 3—
5 each bearing 2 aesthetascs, terminal article
with single aesthetasc. Antennal peduncle
of 4 articles, article 4 only slightly longer
VOLUME 100, NUMBER 3 S713)
F 200PM
Fig. 9. Xylolana radicicola, paratype: A, Cephalon in dorsal view; B, Cephalon in lateral view; C, Cephalon
in oblique-anterior view; D, Cephalon in ventral view; E, Pleon in lateral view; F, Pleotelson and uropods in
dorsal view.
574 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig. 10. Xylolana radicicola, male paratype: A, Adult in dorsal view; B, Adult in lateral view, pereopods
and pleopods not shown; C, Antennule; D, Antenna; E, Maxilla 1; F, Mandible; G, Maxilliped; H, Pleopod 1;
I, Pleopod 2; J, Pleopod 3; K, Pleopod 4; L, Pleopod 5.
than 2 preceding articles; flagellum of 7 se- ginal spines. Maxilla 1, outer ramus with 9
tose articles. Mandibular palp of 3 articles, distal spines; inner ramus with single distal
article 2 longest, with 3 fringed spines dis- seta. Maxillipedal endite short, reaching to
tally, article 3 with 7 fringed spines becom- middle of palp article 1, with single distal
ing more elongate distally; body of man- seta; palp of 5 articles, article 3 longest and
dible somewhat elongate, incisor of 2 widest. Pereopods 1-3 similar, becoming
rounded cusps; lacinia and spine row re- progressively shorter posteriorly. Pereopod
duced; molar triangular, with row of mar- 1, merus with | acute and 4 rounded spines
VOLUME 100, NUMBER 3
DIS)
Fig. 11.
E, Pereopod 5; F, Pereopod 6; G, Pereopod 7.
on posterior margin; carpus with very short
free anterior and posterior margins, with
single spine on posterior margin; propodus
slightly inflated, with 2 spines on posterior
margin; dactylus with strong secondary un-
guis. Pereopods 2 and 3, merus with antero-
distal spine. Pereopods 4—7 similar, becom-
Xylolana radicicola, male paratype: A, Pereopod 1; B, Pereopod 2; C, Pereopod 3; D, Pereopod 4;
ing more elongate posteriorly; basis with 2
posterodistal spines; ischium, merus, car-
pus, and propodus with groups of antero-
distal and posterodistal spines plus few scat-
tered spines on posterior surfaces; unguis
about equal in length to rest of dactylus,
with small secondary unguis. Penes on ster-
576
nite of pereonite 7 short, about twice longer
than wide, separate. Pleopod 1, endopod
half width and subequal in length to exopod.
Pleopod 2, endopod half width but about
one-fourth shorter than exopod, with cla-
vate copulatory stylet articulating in distal
half of mesial margin. Pleopods 3—S similar,
exopod broad, biarticulate, with marginal
plumose setae; endopod triangular, shorter
than exopod, lacking marginal setae. Uro-
podal sympod produced along mesial mar-
gin of endopod; latter ovate, distally broadly
rounded, wider and slightly longer than dis-
tally subtruncate exopod, latter with single
short mesiodistal spines.
Female: Very similar to male. Antennular
flagellum of 4 articles.
Color pattern. —Dorsum with strong pur-
ple-brown pigmentation. Cephalon almost
solidly pigmented; pereonite 1 with scat-
tered patches; pigment becoming denser
posteriorly; pleonites, pleotelson, and uro-
pods fairly densely pigmented; coxae of pe-
reonites 4—7 with only posterior half pig-
mented.
Habitat.—The five specimens of this
species were collected on two occasions,
from the same locality. The specimens came
from the washings of dead but in situ red
mangrove prop roots. These roots, while still
submerged, have lost most of the epiphytes
and epizooites found on live roots. The dead
roots were broken up in a bucket of sea-
water, well rinsed, and the washings
screened. In addition to the new cirolanid
genus, the washings contained numerous
polychaete worms, pycnogonidans, harpac-
ticoid copepods, amphipods, tanaidaceans,
Nebalia sp., plus gnathiid, sphaeromatid,
limnoriid, anthurid, and corallanid isopods.
All these organisms were living either on or
under the decaying and flaking outer layers
of the roots, or in the hollowed and tunneled
inner tissues.
Etymology.—The specific name, mean-
ing ‘dwelling in roots.’ refers to the habitat
of the species.
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Family Sphaeromatidae
Paraleptosphaeroma glynii Buss and
Iverson
Paraleptosphaeroma glynni Buss and Iver-
son, 1981:2, figs. 1-11.
Material. —USNM 205682, 6 specimens
(incl. 2 ovig. 2), Smithsonian-Bredin Ex-
pedition sta 75-59, Portsmouth, Dominica,
amongst boulders, rocks, and dead coral in
0.5 m, 19 Apr 1959.
Previous records. — Punta Paitilla, Pacific
Panama, intertidal.
Remarks. —This is the first Atlantic re-
cord of this monotypic genus and is there-
fore one of the few species of isopods known
to occur on both sides of the Isthmus of
Panama.
Acknowledgments
I am grateful to Drs. K. Fauchald, G.
Hendler, M. Jones, and K. Rutzler for col-
lecting some of the material described in
this paper. I thank Marilyn Schotte for as-
sistance with the sorting of samples and dis-
secting and drawing some of the specimens,
and Dr. T. E. Bowman for reading and com-
menting on the manuscript. This paper is
contribution number 219 of the Smithson-
ian Institution’s Caribbean Coral Reef Eco-
systems program. This program provided
funding for the author’s visit to Belize in
December, 1986.
Literature Cited
Bruce, N. L. 1986. Cirolanidae (Crustacea: Isopoda)
of Australia.—Records of the Australian Mu-
seum, supplement 6:1-239.
Brusca, R. C. 1984. Phylogeny, evolution and bio-
geography of the marine isopod subfamily Ido-
teinae (Crustacea: Isopoda: Idoteidae). — Trans-
actions of the San Diego SOU of Natural
History 20(7):99-134.
Buss, L. W., and E. W. Iverson. 1981. A new genus
and ee of Sphaeromatidae (Crustacea: Isop-
oda) with experiments and observations on its
reproductive biology, interspecific interactions
and color polymorphisms. — Postilla 184:1-23.
VOLUME 100, NUMBER 3
Kensley, B., and M. Schotte. 1987. New records of
isopod Crustacea from the Caribbean, the Flor-
ida Keys, and the Bahamas. — Proceedings of the
Biological Society of Washington 100(1):216-
DAT.
Menzies, R. J.,and D. Frankenberg. 1966. Handbook
on the common marine isopod Crustacea of
Georgia. University of Georgia Press, Athens,
Georgia, 93 pp.
Moore, H. F. 1901. Report on Porto Rican Isopo-
da.—U.S. Fish Commission Bulletin for 1900
2:161-176.
SVT
Richardson, H. 1900. Synopses of North-American
invertebrates. VIII. The Isopoda. Part 1. Che-
lifera, Flabellifera, Valvifera.—The American
Naturalist 34:207-309.
Department of Invertebrate Zoology, Na-
tional Museum of Natural History, Smith-
sonian Institution, Washington, D.C. 20560.
PROC. BIOL. SOC. WASH.
100(3), 1987, pp. 578-583
STUDIES OF THE LEPIDAPLOA COMPLEX
(VERNONIEAE: ASTERACEAE). I.
THE GENUS STENOCEPHALUM SCH. BIP.
Harold Robinson
Abstract.—The genus Stenocephalum Schultz-Bip. is resurrected for a group
of five tropical American species related to the genus Lepidaploa. The two
genera differ from Vernonia and are like each other in the form of their lophate
pollen with “rhizomatous” columellae under the crests. The resurrected genus |
is distinguished structurally and functionally from Lepidaploa by fewer flowers
in the heads. The surface of the pollen also has more numerous areolae, and
the bases of the styles usually lack a node.
The establishment of a series of reliable
and useful phyletic generic concepts in the
tribe Vernonieae has already progressed
considerably from the traditional core con-
cept of Bentham and Hooker (1873) and
Hoffmann (1890-94) that had numerous
segregates based primarily on pappus vari-
ations. A major advance was that of Jones
(1977) in recognizing basic differences in
chemistry and chromosome number be-
tween New and Old World groups. Within
the broad hemispheric groups are many
genera, some of which have already been
recognized in the traditional treatments,
while others still reside in the synonymy of
Vernonia. The present series of studies 1s
aimed at the restoration or naming as new
of many of the genera of the Lepidaploa
complex that have been placed in the genus
Vernonia.
The paleotropical elements of the tribe
are phyletically remote from typical Ver-
nonia in eastern North America, and are
thus more easily justified at the generic level
on the basis of the Jones’ (1977) results.
Some such genera recently resurrected in-
clude Distephanus Cass. and Gymnanthe-
mum Cass. (Robinson and Kahn 1986), with
the former not even fitting the broad con-
cept of Vernonia. The neotropical elements
fall into the same broad chemical and cy-
tological group as typical Vernonia, but they
also require segregation to a degree exceed-
ing that found in the traditional tribal con-
cepts. Many of the necessary additional seg-
regates such as Critoniopsis Sch. Bip.
(Robinson 1980) are at least as phyletically
distinct as traditional genera like Piptocarpa
R. Br. and Pollalesta H.B.K. Many small
segregates remain to be recognized, but the
largest number of neotropical species in-
volved belong to the Lepidaploa group. Six
genera, Lepidaploa (Cass.) Cass., Stenoce-
phalum Sch. Bip., and four as yet unnamed,
are to be treated in a series of studies of
which the present resurrection of Stenoce-
phalum is the first.
Significant Characters
Number of flowers in the head.—In the
series of five genera, Stenocephalum Schultz-
Bip. is notable for the comparative accuracy
of delimitation at the time it was first pub-
lished (Schultz-Bipontinus 1863). The pa-
per in which the genus was described dealt
mostly with genera with reduced numbers
of flowers in the heads such as Lychnophora
Mart. and Eremanthus Less. The reduced
number of flowers appears to have been the
VOLUME 100, NUMBER 3
primary reason Schultz (1863) elevated
Stenocephalum to generic level while leav-
ing related forms in Vernonia. No phyletic
considerations were evident in the work of
Schultz, and the reduced number of flowers
seems to have been regarded as a violation
of the limits of Vernonia. Stenocephalum
was relegated to the level of a section in
Vernonia by Baker (1873) and has remained
under the latter genus until the present. It
was made a subsection by Jones (1979).
The present concept of Stenocephalum is
more refined, and the relationship to other
groups 1s now more evident. Even though
the genus has no close relationship to the
Lychnophorinae, the number of flowers in
the head remains one of the most important
characters in the delimitation of the genus.
The heads contain usually 4—7 flowers, and
rarely as many as 10. The characteristic
number is lower than that in any of the other
members of the Lepidaploa complex and
moreover is reduced in proportion to the
total of 15—22 involucral bracts in the heads.
The reduction results in a lower proportion
of flowers in four of the five species of Steno-
cephalum than the *% to '4 ratio seen in most
of Lepidaploa. The other exceptions in the
group all have much larger numbers of both
flowers and bracts. The ratio clearly distin-
guishes Stenocephalum from typical Ver-
nonia which has more flowers and nearly
equal numbers of bracts and flowers in the
heads.
The reduced number of flowers in the head
prevents some of the types of floral displays
most common in the Vernonieae. In many
members of the tribe the peripheral flowers
spread in a manner resembling rays, a fea-
ture particularly notable in typical Vernonia
and Stokesia. In Centratherum and some
species like Vernonia santosii H. Robins.
the zone of peripheral flowers is differen-
tiated by maturing while the inner flowers
remain unopened. The inner flowers open
on another day with a distinctly unraylike
appearance. These strategies are difficult if
not impossible in the smaller heads of
579
Stenocephalum, and in many cases the flow-
ering heads are very unimpressive. Only S.
tragiaefolium seems to furnish a notable flo-
ral display, caused by the massed effect of
the whole inflorescence. One assumes that
the limited floral presentation would have
some effect on attraction of pollinators, but
field observations remain to be made.
Disposition of heads.—The genus Steno-
cephalum consists entirely of species that
have sessile heads at the nodes. The cymose
structure is basically like that of Lepidaploa
with none of the pedicellate modifications
of the heads seen in either of the related
unnamed genera. The heads may be isolated
at separate, rather remote nodes as in the
type species S. apiculatum or S. jucundum,
clustered on short lateral branches as in S.
megapotamicum, or more closely massed in
a terminal inflorescence as in S. tragiaefo-
lium. The variation of disposition of heads
in the genus seems to be a significant factor
in speciation. The two species, S. apicula-
tum and S. jucundum, that show similar
displays have nearly complete geographical
isolation.
Style base. — As noted in the study of Di-
stephanus (Robinson and Kahn 1986), sty-
lar nodes are widely distributed in the neo-
tropical Vernonieae. Stylar nodes are
particularly characteristic of Lepidaploa,
often being quite marked as disc-like exten-
sions at the base of the style just above the
nectary. The node is lacking, however, in
one related group that has previously been
placed in Vernonia series Buddleiifoliae, and
it is also poorly developed or lacking in the
genus Stenocephalum. While most species
have no node, a hint of basal differentiation
is seen in some S. megapolitanum and a
small but distinct basal rim is seen in S.
tragiaefolium.
Achene structure.—The carpopodium of
Stenocephalum was mentioned by Schultz
(1863) as being large. The size is not unusual
in the Vernonieae but the structure is uni-
form for all five species. The shape is short-
cylindrical except for the rounding of the
580
basal margins. The upper edge of the dif-
ferentiated structure is only slightly irregu-
lar at or below the level of the lowest setulae.
In contrast to the carpopodium, the
achene wall structure in Stenocephalum
varies extensively in setulosity and form of
surface cells. The setulae seem character-
istically somewhat contorted and uneven at
the tips. One of the cells may project far
beyond the other, or some setulae may be
uniseriate from at or near the base. The ribs
seem most prominent in S. jucundum, S.
megapotanicum and S. hystrix where they
stand out from the surface and separate the
setulose areas into distinct bands. The ribs
are weaker and the setulae more densely
cover the surface in S. apiculatum and S.
tragiaefolium. The ribs are particularly weak
in the latter species and occasionally there
are less than ten. In the type species, S. apic-
ulatum, the cells of the intercostal region
seem lax with weak walls that are scarcely
observable under the microscope. The cells
are most readily noticeable by the manner
that they peel away with their imbedded
setular bases from the thicker-walled elon-
gate cells below. There are no other differ-
entiated cells evident in this surface. The
Central American and northern South
American S. jucundum has surface cells that
are similar to those of the type but they are
much firmer and more easily seen. The three
remaining species are seen to have thicker-
walled elongate cells in immediate connec-
tion with the setular bases. All three species
show islands of differentiated idioblasts with
cells occurring singly or in a series of two
to three. The latter pattern is one also seen
in many species of Lepidaploa.
Pollen. — All members of the genus Steno-
cephalum show uniformity in their pollen
and its general features differ only in details
of areolation. The grains are of the type re-
defined here as Lepidaploa-type with the
ridges of the lophate grains subtended by a
horizontal “rhizome.” The whole structure
seems to easily peel away from the foot-
layer (Fig. 4). The grains of the type species
shown (Figs. 1-4) differ from other Lepi-
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
daploa type pollens in the height of the crests
and the exposure of the basal columellae,
but the differences are ones of degree.
In form of areolation all the grains fall
generally in the C-type which has areoles at
the poles. The structure is unlike the C-type
of Lepidaploa, however, in the extra num-
ber of areolae in the intercolpar region (Figs.
1-3). Observations under the light micro-
scope indicate that none of the other species
are as extreme in the number of areolae as
the type species that 1s shown, but all have
at least some extra, there often being three
across the intercolpar region. The higher
number of areolae and the presence of polar
areolae is a combination unknown in Lep-
idaploa.
Relationship of Stenocephalum
The form of the pollen in Stenocephalum
clearly indicates a phyletic position near
Lepidaploa. This is advanced over the more
primitive pollen type seen in such genera as
Stokesia where the basal columellae each
reach the footlayer individually. The pres-
ence of a trace of stylar node in S. tragiae-
folium and the presence of idioblasts on the
surface of the achene in three species of
Stenocephalum also indicate a position close
to Lepidaploa. The positions of the heads
can also be interpreted as an seriately in-
novating cyme as in Lepidaploa. Neverthe-
less, the species potentially closest to the
outgroup, S. tragiaefolium already shows
some increase in the number of areolae on
the pollen grains and already has the num-
ber of flowers in the head reduced to the
point where the whole inflorescence is need-
ed for floral display. Within Lepidaploa there
iS no species that seems particularly close
to Stenocephalum. A species of the general
Lepidaploa relationship that was initially
thought to be related to Stenocephalum,
Vernonia regis H. Robins., has enough flow-
ers to fall completely within the Lepidaploa
flower/bract ratio, a carpopodium on the
achene with a more conical shape, much
denser and straighter setulae on the achene
VOLUME 100, NUMBER 3 581
ec. Hum 1./76um
Figs. 1-4. Pollen of Stenocephalum apiculatum (Martius in DC.) Schultz-Bip. 1-3, dotted lines = 12 um; 4,
dotted line = 1.76 wm. 1, Polar view showing polar areole; 2, Oblique view showing half of intercolpus with
complete median transverse row of three areoles; 3, Colpar view; 4, Broken section of crest showing details of
“rhizomatous” columellae and attachment of perforated tectum.
582
with more equal tips, a well-developed node
at the base of the style, and type-B pollen.
Stenocephalum Schultz-Bip.
Stenocephalum Schultz-Bip., Pollichia 20/
21:385. 1863, type Vernonia apiculata
Mart. ex DC.
Vernonia sect. Stenocephalum (Schultz-Bip.)
Baker in Mart., Fl. Bras. 6(2):25. 1873.
Vernonia subsect. Stenocephalum (Schultz-
Bip.) S. B. Jones, Rhodora 81:437. 1979.
Plants herbaceous, perennial, erect to 0.3-
1.0 m tall, with little or no vegetative
branching above the base. Leaves linear to
broadly oblong, ovate or obovate, discolo-
rous, dark green and densely to evanes-
cently pilose above, pale tomentose below.
Inflorescence cymose, heads single or
grouped at primary nodes or densely clus-
tered on lateral branches. Heads cylindrical;
involucral bracts ca. 15—22 in 3—4 graduated
series, with pungent usually narrowly acu-
minate tips (short cuspidate in some S. me-
gapotamicum), slightly to distinctly re-
curved; flowers 4—7(—10) in a head; corollas
lavender, with or without hairs at tips of
lobes; anther thecae having shields of me-
dian endothecial cells variously radial or
looped with multiple nodes, anther appen-
dage glabrous; style base without or with
only slight node. Achenes with ten ribs
weakly to strongly developed, intercostal
surface with surface of lax cells or with firm-
er cells intermixed with some idioblast clus-
ters; setulae appearing evenly distributed in
species with weak ribs, otherwise restricted
between ribs, with often unequal and con-
torted tips, rarely uniseriate from base; car-
popodium short-cylindrical, rounded only
at basal margin; pappus of short outer squa-
mellae and numerous inner capillary bris-
tles. Pollen grains (40—)45—48(—S0) um in
fluid, distinctly lophate with “‘rhizomatous”’
columellar structure under crests, exine eas-
ily stripping away from footlayer, lophate
pattern of general C-type with polar areoles
but differing by extra intercolpar areoles (up
to 9).
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Key to the Species of Stenocephalum
1. Cauline and primary branch leaves
broadly ovate to obovate, distinctly
narrowed at base
2. Heads laxly disposed along stems
and branches, usually solitary in or
near axils; leaves ovate (Central
America, N. South America) .....
S. Jjucundum
2. Heads congested near apex of plant;
leaves mostly obovate (Brazil)
Ce rte ere eMC A Gee ee S. tragiaefolium
1. Cauline leaves narrowly oblong with
broadened bases or linear through-
out, broader bladed leaves restricted
to basal rosette
3. Cauline leaves linear throughout,
narrow at base (Paraguay) ..S. hystrix
3. Cauline leaves linear to oblong with
oblong or broadened bases ....... A
4. Inflorescence laxly branched with
heads mostly solitary in successive
axils; involucral bracts ca. 20, with
narrowly acuminate tips; flowers ca.
4 in a head (Brazil, Venezuela)
re eee Mee a SUNS S. apiculatum
4. Inflorescence often profusely
branched with heads distinctly clus-
tered in axils or on short branchlets;
involucral bracts ca. 15, with shortly
acuminate or cuspidate tips; flowers
ca. 6—7 in a head (Argentina, Brazil,
Paraguay, Uruguay)
ee © © © © © © © © © © © © © © © © © © © 6
The species of Stenocephalum recognized
in this study are as follows:
Stenocephalum apiculatum
(Martius in DC.) Schultz-Bip.
Figs. 1-4
Vernonia apiculata Martius in DC., Prodr.
5:51. 1836. Vernonia monticola Martius
in DC., Prodr. 5:18. 1836. Stenocephal-
um monticola (Martius in DC.) Schultz-
Bip., Pollichia 20/21:386. 1863. Steno-
cephalum apiculatum (Martius in DC.)
Schultz-Bip., Pollichia 20/21:387. 1863.
VOLUME 100, NUMBER 3
The species is almost completely restrict-
ed to Brazil, but one specimen seen in this
study from Venezuela seems to be this
species [Bolivar: Rio Villacda (Rio Auyac-
da). Jan 6, 1956. Wurdack & Monachino
41144, US].
Stenocephalum hystrix
(Chodat) H. Robinson, comb. nov.
Vernonia hystrix Chodat, Bull. Herb. Boiss.
Sem li2:298. 1902.
Stenocephalum jucundum (Gleason)
H. Robinson, comb. nov.
Vernonia jucunda Gleason, Bull Torrey Bot.
Club 46:248. 1919. Vernonia spinulosa
Gleason, Bull. Torrey Bot. Club 52:188.
1925. Vernonia llanorum Badillo, Bol.
Soc. Venez. Cienc. Nat. 10:218. 1946.
Gleason (1925) distinguished his Vene-
zuelan species from the Central American
entity by the narrower branch leaves, but
his description indicated that the type had
lost most of its leaves. The slight tendency
that has been seen for the distal leaves to
be narrower does not seem to justify sepa-
rate specific rank. The picture has been
complicated by the presence in southeastern
Venezuela of the specimen mentioned above
that has been determined here as the Bra-
Zilian S. apiculatum.
Stenocephalum megapotamicum
(Spreng.) Schultz-Bip.
Vernonia megapotamica Spreng., Syst. Veg.
3:437. 1826. Vernonia megapotamica var.
brevifolia DC., Prodr. 5:51. 1836. Ver-
nonia megapotamica var. melanotrichia
DC., Prodr. 5:51. 1836. Stenocephalum
brevifolium (DC.) Schultz-Bip., Pollichia
20/21:387. 1863. Stenocephalum mega-
potamicum (Spreng.) Schultz-Bip., Polli-
chia 20/21:388.1863. Stenocephalum
melanotrichium (DC.) Schultz-Bip., Pol-
lichia 20/21:388. 1863. Stenocephalum
penicillatum Schultz-Bip., Pollichia 20/
21:389. 1863. Stenocephalum hexan-
thum Schultz-Bip., Pollichia 20/21:390.
583
1863. Vernonia hexantha (Schultz-Bip.)
Baker in Mart., Fl. Bras. 6(2):27. 1873.
Stenocephalum tragiaefolium
(DC.) Schultz-Bip.
Vernonia tragiaefolia DC., Prodr. 5:60.
1836. Stenocephalum tragiaefolium (DC.)
Schultz-Bip., Pollichia 20/21:389. 1863.
Vernonia interjecta Baker in Mart.?, FI.
Bras. 6(2):28. 1973.
Acknowledgments
The pollen specimens were prepared by
Mary Sangrey using the facilities of the Bot-
any Department Palynological Laboratory.
The photographs were prepared by Suzanne
Braden of the Smithsonian Museum of Nat-
ural History SEM Laboratory using a Hi-
tachi 570 scanning electron microscope.
Literature Cited
Baker, J. G. 1873. Compositae I. Vernoniaceae. Jn
Martius, C. F. P., Flora Brasiliensis 6(2):2-179.
Bentham, G., and J. D. Hooker. 1873. Genera Plan-
tarum 2(1):1-544.
Gleason, H. A. 1925. Studies on the flora of northern
South America. V.— Bulletin of the Torrey Bo-
tanical Club 52:181-196.
Hoffmann, O. 1890-94. Compositae. Jn A. Engler
and K. Prantl, eds., Die Natiirlichen Pflanzen-
familien 4(5):87—387. Leipzig.
Jones, S. 1977. Vernonieae: Systematic Review. Jn
V.H. Heywood, J. B. Harborne, and B. L. Turn-
er, eds., The Biology and Chemistry of the Com-
positae, chapter 17:503-521.
. 1979. Synoptic classification and pollen mor-
phology of Vernonia (Compositae: Vernonieae)
in the New World.—Rhodora 81:425-447.
Robinson, H. 1980. Re-establishment of the genus
Critoniopsis (Vernonieae: Asteraceae). — Phy-
tologia 46:437-442.
,and B. Kahn. 1986. Trinervate leaves, yellow
flowers, tailed anthers, and pollen variation in
Distephanus Cassini (Vernonieae: Astera-
ceae). — Proceedings of the Biological Society of
Washington 99:493-501.
Schultz-Bipontinus, C. H. 1863. Geschichte der Gat-
tung Lychnophora. —Pollichia 20/21:329-439.
Department of Botany, National Mu-
seum of Natural History, Smithsonian In-
stitution, Washington, D. C. 20560.
PROC. BIOL. SOC. WASH.
100(3), 1987, pp. 584-589
STUDIES IN THE LEPIDAPLOA COMPLEX
(VERNONIEAE: ASTERACEAE). II.
A NEW GENUS, ECHINOCORYNE
Harold Robinson
Abstract.—The genus Echinocoryne is established for a group of Brazilian
species related to the genus Lepidaploa in the “rhizomatous” form of the
columellae in its lophate pollen. The new genus is characterized by its large
number of slender involucral bracts and its pedunculate heads. The genus is —
also notable for its densely sericeous pubescence and poorly differentiated style
base.
The present paper is one of a series of five
devoted to the particular problem of refin-
ing generic limits in the neotropical Lepi-
daploa relationship of the Vernonieae.
Echinocoryne is one of three genera in the
complex showing a specialized pollen that
is unique to the group. The first of the genera
treated, Stenocephalum, a small genus of
Central of South America south to Argen-
tina, is distinguished by heads with few
flowers and by pollen with some extra ar-
eoles (Robinson, 1987). The present genus,
Echinocoryne is also small, but is restricted
to Brazil. Another paper will deal with the
small V. brachiata and V. scorpioides groups.
The two remaining papers in the series will
deal with the two larger elements of the
complex, one being on Lepidaploa itself with
a fuller discussion of the basic pollen form
of the complex.
The species of Echinocoryne have been
placed together in some previous treat-
ments. Baker (1873) placed them in his Ver-
nonia catagory Lepidaploae Paniculatae I.
Oxylepidae, while Jones (1979) placed them
in Vernonia as a new series Subulatae under
his subsection Nudiflorae. The Baker treat-
ment included two additional species in the
group. The first, V. foliosa Gardn. with its
strikingly different Lychnophorine habit and
Proteopsis-like heads, seems from photo-
graphs to have no close relationship. The
second, V. virgulata, was also retained by
Jones in his series Subulatae. The latter
species has some resemblance to Echino-
coryne in its sericeous indument and vari-
ably short-pedunculate heads, but differs in
the aspect of the short-tipped multiseriate
involucral bracts. The number of bracts is
approximately 40 in head compared to ca.
10 flowers, a ratio approaching but not
equalling that in Echinocoryne. In addition,
the stems are distinctively grooved, recep-
tacles setiferous, corollas have more hairs
on their lobe tips, carpopodium has a dif-
ferent shape, while the upper achene and
corolla have larger short-stalked capitate
glands of a form not seen in Echinocoryne.
The pollen differences are cited below. The
overall impression is one of a remote rela-
tionship elsewhere in the Lepidaploa com-
plex.
The only species included here that was
not covered in the previously mentioned
works is V. echinocephala H. Robinson
which has been described since those works
(Robinson 1980).
Significant Characters
Pubescence. —The vegetative parts of
Echinocoryne are notable for their sericeous
pubescence. Such hairs occur to the exclu-
sion of evident glandular punctations. The
VOLUME 100, NUMBER 3
density of the hairs varies from the sparse
form seen in the more herbaceous EF. sub-
ulata to the usually dense, sometimes sil-
very sericeousness of parts of the other
species. The pubescence is most dense on
the undersides of the leaves, but is often
nearly as thick on the upper side without
obscuring the darker color of the leaf sur-
face. The hairs are shorter but often very
dense on the involucral bracts. In contrast,
the corollas have less hairs than those of
many other species in the Vernonieae. The
corollas are nearly glabrous, except at the
tips of the lobes, in all the species. In E.
subulata even the tips of the lobes have no
hairs, just the few characteristic minute
glands found in all the species.
Pedunculate heads.—The heads of Echi-
nocoryne, with rare exceptions, are all pe-
dunculate; however, the length of the pe-
duncles varies. The peduncles are longest in
the herbaceous E. subu/ata and subshrub E.
echinocephala, but are short in typical E.
holosericea and FE. schwenckiaefolia. The
length seems consistent in the first two
species, but in the latter two seems to be
variable. The longest peduncles in the genus
seem to be generally associated with larger
heads, but the correlation breaks down in
forms with shorter peduncles.
Involucre.—The graduated bracts of the
involucre in Echinocoryne are all similar in
their general form and show no differentia-
tion of the type seen in many Lepidaploa.
They have minimal differentiation of the
largest inner bracts. All the bracts are linear-
lanceolate with narrowly pungent almost se-
tiform tips. The appearance of the involucre
is characteristically echinate.
The most significant aspect of the invo-
lucre compared to Lepidaploa is the ratio
of the bracts to the flowers. The example
with the largest numbers, EF’. echinocephala,
has 400-500 bracts with ca. 50 flowers in a
head. The number of bracts may be as great
as any in the tribe, and the ratio of bracts
to flowers 1s larger than any in the tribe
except those with three or less flowers in a
585
head. The stability of the usually 2:1 or 3:
I ratio in the large genus Lepidaploa seems
significant, and the striking exception to that
ratio in Echinocoryne is therefore also con-
sidered significant.
Style base. —In all of the species the base
of the style has a few rows of sclerified cells
on the part that is immersed in the top of
the nectary, but no enlargement of the type
seen in Lepidaploa has been seen in any of
the species of Echinocoryne.
Achene.—The achene is densely and se-
riceously setuliferous without any interven-
ing glands or uniseriate hairs. The five weak
ribs of the surface are completely obscured
by the setulae. The carpopodium is some-
what unusual in its turbinate form which is
constricted above. The incurved upper sur-
face is further distinct for the number of
setulae borne as low as the broadest part of
the carpopodium, well below the level of
the uppermost margin of the structure. Al-
though the differentiated carpopodial cells
do not occur directly above such setulae,
but only between them, the impression 1s
nevertheless that the upper incurved third
of the carpopodium is setuliferous.
Pollen. —Echinocoryne has lophate pol-
len grains with a “rhizomatous”’ columellar
structure under the crests (Figs. 1-6). The
pollen in the group was classified as type B
by Jones (1979), which it is in the broad
sense of having colpi continuous to the poles.
That classification has proven much too
broad, however, because it includes phylet-
ically significant variations. The type B of
Jones 1s approximately the same as that
called the Vernonia argyrophylla type by Stix
(1960). The type, as represented by the latter
species, was erroneously regarded by Stix
under her general Vernonia category which
she showed in a drawing with the structure
here called “rhizomatous.” The type B based
on V. argyrophylla is not rhizomatous, and
it tends to be larger in size with three areoles
equatorially across the intercolpar region.
The Echinocoryne pollen has only two in-
tercolpar rows of areoles (Figs. 1-3) and has
586
rhizomatous structure (Figs. 5, 6). It is a
type more often seen in the genus Lepidap-
loa. The grains seem distinct from similar
forms in Lepidaploa in only one possible
way, the manner in which the rhizomatous
columellae peel away from the foot-layer
leaving slightly raised ridges but no clear
scars (Fig. 4).
Vernonia virgulata which was placed with
species of Echinocoryne by both Baker
(1873) and Jones (1979) actually does not
have the type B pollen designated for it by
Jones. The type is technically C with polar
areoles, but the grains are distinct from any
other type C grains by the non-rhizomatous
columellar structure of the crests and the
frequent extra areole in the middle of the
intercolpar region. The pollen is regarded
here as closest phyletically to the type B as
seen in V. argyrophylla although it has a
different areolation. The pollen structure
tends to reinforce the impression gained
from other characters that V. virgulata is
only remotely related to Echinocoryne.
Relationships
A phyletic position of Echinocoryne can
be proposed if one accepts certain conclu-
sions regarding the characters analyzed
above. The specialized rhizomatous crests
of the pollen place the genus in the imme-
diate relationship of Lepidaploa in which
such complex pollen is unique. Since the
bract/flower ratio of Lepidaploa extends
more widely than the rhizomatous pollen
character, then the departure from that ratio
in the small genus Echinocoryne seems de-
rived in the more immediate ancestry of
that genus. The pedunculate condition of
the heads is not found in Lepidaploa but is
found in what appears to be its nearest out-
group. Consequently, a position for Echino-
coryne from near the basal stock of Lepi-
daploa might be possible. It is also possible
that the pedunculate condition is a rever-
sion and that the genus is a more recent
derivative of Lepidaploa. However, no
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
species in the latter genus seems a likely
candidate for close relationship.
As indicated, the genus dealt with here is
well-marked in its habit, and the characters
violate the character limits by which all the
most closely related genera are most effec-
tively defined. Furthermore, a phyletic po-
sition outside of the immediate Lepidaploa
generic clade seems possible. The name
Echinocoryne, meaning hedgehog or prickly
club, is particularly suitable for the pedun-
culate heads of the plants with their many
spiny involucral bracts.
Echinocoryne H. Robinson, gen. nov.
Plantae herbaceae perennes erectae ad 2
m altae vegetative laxe ramosae, in caulibus
foliis et bracteis involucri dense albo-seri-
ceae. Folia alterna sessilia vel subsessilia li-
nearia vel oblonga margine integra anguste
reflexa apice acuta et minute apiculata sub-
tus pallidioria non glandulifera. Inflorscen-
tiae diffusae in pedunculis leniter vel arcte
elongatae. Capitula late campanulata; brac-
teae involucri super-abundentes dense sub-
imbricatae graduatae ca. 110-500 et 6—9 se-
rlatae patentes vel erecto-patentes lineares
apice longe punguntes. Flores 15-60 in ca-
pitulo; corollae lavandulae in partibus ma-
joribus glabrae in lobis subapice pauce spic-
uliferae et minute glanduliferae; cellulae
endotheciales in scutis scleroideis radiatae
vel curvatae, nodis multifidis; appendices
antherarum non glanduliferae; basi stylo-
rum noduliferi1. Achaenia longe setulifera,
costis 5 indistinctis in setulis dense obsitis,
setulis perdensis strictis sericeis, cellulis in-
tercostalis raphides elongates continenti-
bus; carpopodia turbinata in partibus su-
perioribus setulifera; setae pappi interiores
capillares persistentes ca. 30, squamae pap-
pi exteriores lineares. Grana pollinis in dia-
metro ca. 50 um valde lophata Lepidaploa-
forma (subtypus E).
Type species. — Vernonia holosericea Mart.
ex DC.
A number of names have been proposed
VOLUME 100, NUMBER 3
for various entities in the genus Echinocor-
yne, but the name V. subulata has repre-
sented the only obviously distinct species
besides the type. The other members of the
genus have seemed comparatively alike, and
the name V. holosericea has been applied
widely to them by myself and others. Con-
trary to the superficial appearances, the
present study has shown that all the avail-
able names in the genus correlate with dis-
tinguishable species having distinctive geo-
graphical distributions.
Key to the Recognized Species of
Echinocoryne
1. Plants laxly herbaceous with pale
green membranaceous leaves and
yellowish-green stems, leaf blades
broad, plane or with scarcely re-
curved margins, with sparse flex-
uose hairs; peduncles long and flex-
WOUISMEM Ee E. subulata
— Plants more stiffy herbaceous to suf-
frutescent with darker leaves and
stems, leaf blades narrow or broad
with distinctly recurved margins,
with dense hairs on at least under
surface; peduncles strict and often
short
2. Heads few or solitary on peduncles
7 cm long or longer; leaves lanceo-
late with densely whitish sericeous
WINGSESUIACC a.) 4.4. E. echinocephala
— Heads usually numerous in complex
inflorescences; peduncles usually 4
cm or less long; leaves linear to ob-
long with sordid or grayish pubes-
cense on undersurfaces
3. Leaves mostly broadly elliptical to
oblong 4 to '2 as wide as long, often
velvety pubescent beneath
50 coro ere E. schwenkiaefolia
— Leaves narrowly lanceolate to lin-
ear, less than 4 as wide as long, with
pubescence beneath variable but not
WEINIGIN ac ete alee ele enemas 4
4. Heads with 30-60 flowers, 12 mm
oOo 0'6 0 0 0
587
2. © © © © © ©
Pearce Ps MIN eae St ee E. holosericea
— Heads with 15-21 flowers, 8-10 mm
high: atimatuciiyeeg | see ee eee 5
5. Involucres pale yellowish; upper
leaves abruptly terminating in nar-
rowly obtuse apices ........ EFS SETLCTA
— Involucres tinged with red; upper
leaves tapering to narrowly acute
APICES Mi 0 hee ee E. pungens
The species recognized in the genus are
as follows:
Echinocoryne echinocephala
(H. Robinson) H. Robinson, comb. nov.
Vernonia echinocephala H. Robinson, Phy-
tologia 45:173. 1980. Southern Goias.
Echinocoryne holosericea
(Mart. in DC.) H. Robinson, comb. nov.
Figs. 1-6
Vernonia holosericea Mart. in DC., Prodr.
5:43. 1836. Bahia, southern Maranhao,
northern Minas Gerais.
Echinocoryne pungens
(Gardn.) H. Robinson, comb. nov.
Vernonia pungens Gardn., London J. Bot.
6:418. 1847. Southern Goias, western Ba-
hia, western Minas Gerais, Distrito Fed-
eral. Examination of an isotype (US)
shows that neither the odd statement of
4—5 flowers by Gardner (1847) nor the
30-40 flowers in the head claimed by
Baker (1873) is correct. The number seems
to be ca. 20, essentially the same as that
in E. stricta which has heads of about the
same size. The involucral bracts are gen-
erally less longly attentuate than in the
other species.
Echinocoryne schwenkiaefolia
(Mart. in DC.) H. Robinson, comb. nov.
Vernonia schwenkiaefolia Mart. in DC.,
Prodr. 5:44. 1836. Minas Gerais, south-
588 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Figs. 1-6. Pollen of Echinocoryne holosericea (Martius in DC.) H. Robinson, dotted lines of 1. = 15 um, 2,
3. = 13.6 um, 4. = 7.5 wm, 5. = 2 um, 6. = 2.7 wm. 1, Polar view showing convergence of colpi; 2, View of
colpus (transverse); 3, View of intercolpar region showing characteristic two rows of areoles; 4-6. Broken grains.
4, View showing nearly unscarred foot-layer with crest removed; 5, Crest in section showing “rhizome” and
structure of perforated tectum; 6, Lateral view of crest showing weak basal attachment of “‘rhizome.”’
ern Goias, southern Bahia. The species is
rather consistent in its broad short leaves
but is more variable in its heads. The lat-
ter range from nearly as pale as E. stricta
to as dark as E. holosericea, but the ma-
ture heads are mostly in the size range of
the latter species with 25—45 flowers.
Echinocoryne stricta
(Gardn.) H. Robinson, comb. nov.
Vernonia stricta Gardn., London J. Bot. 5:
219. 1846. Southern Minas Gerais, Sao
Paulo. The small pale involucres are very
noticeable, especially in contrast to the
rufus pappus and the reddish corollas.
Echinocoryne subulata
(Baker) H. Robinson, comb. nov.
Vernonia subulata Baker, FI. Bras. 6(2):108.
1973. Southern Goias, central Minas
Gerais.
Acknowledgments
The pollen specimens were prepared by
Mary Sangrey using facilities of the Botany
Department Palynological Laboratory. The
photographs were prepared by Suzanne Bra-
den of the Smithsonian Museum of Natural
History SEM Laboratory using a Hitachi
570 scanning electrom microscope.
VOLUME 100, NUMBER 3
Literature Cited
Baker, J. G. 1873. Compositae I. Vernoniaceae. /n
Martius, C. F. P., Flora Brasiliensis 6(2):2-179.
Gardner, G. 1847. Contributions towards a flora of
Brazil, being the characters of several new species
of Compositae, belonging to the tribes Vernon-
iaceae and Eupatoriaceae, from the Province of
Goyaz.— London Journal of Botany 6:417-449.
Jones, S. 1979. Synopsis and pollen morphology of
Vernonia (Compositae: Vernonieae) in the New
World.—Rhodora 81:425-447.
Robinson, H. 1980. New species of Vernonieae (As-
589
teraceae). V. Additions to Vernonia from Bra-
sil.— Phytologia 45:166-208.
1987. Studies of the Lepidaploa complex
(Vernonieae: Asteraceae). I. The genus Steno-
cephalum Sch. Bip.—Proceedings of the Bio-
logical Society of Washington 100:578-583.
Stix, A. 1960. Pollenmorphologische Untersuchungen
an Compositen.—Grana Palynologica 2:41-104.
Department of Botany, National Mu-
seum of Natural History, Smithsonian In-
stitution, Washington, D.C. 20560.
PROC. BIOL. SOC. WASH.
100(3), 1987, pp. 590-595
UPOGEBIA SYNAGELAS, NEW SPECIES, A COMMENSAL
MUD SHRIMP FROM SPONGES IN THE WESTERN
CENTRAL ATLANTIC (DECAPODA: UPOGEBIIDAE)
Austin B. Williams
Abstract. — Upogebia synagelas, a new species of mud shrimp from the west-
ern central Atlantic, lives commensally as adult male-female pairs in species
of the agelasid sponge, Age/as, in the eastern Gulf of Mexico and West Indian
region. Both adults and zoea I are described. The species is similar to U.
ramphula of the eastern Pacific.
Specimens of an undescribed species of
Upogebia were brought to my attention re-
cently by Drs. Paula J. B. Scott of McMaster
University, Hamilton, Ontario, Canada, and
Elizabeth Sides, Dublin, Ireland. Both of
these investigators have found the species
living commensally in the agelasid sponge,
Agelas sceptrum (Lamarck), and both Scott,
Reiswig, and Marcotte (in press) and E. Sides
have observed that the adults live as male-
female pairs in the sponge whereas juveniles
may occur singly. Michael Dardeau and as-
sociates of the Marine Environmental Sci-
ences Consortium, Dauphin Island, Ala-
bama, have found the shrimp in 4. dispar
Duchassaing and Michelotti on the Florida
Middle Grounds in the Eastern Gulf of
Mexico and the Bahama Islands. The pur-
pose of this paper is to describe the adult
shrimp as well as selected features of the
zoea stage I, with comparative remarks on
similar species.
Type specimens and larval material have
been placed in the crustacean collection of
the United States National Museum of Nat-
ural History (USNM) and in the research
collection of the Marine Environmental Sci-
ences Consortium (MESC), Dauphin Island
Sea Lab, Dauphin Island, Alabama 36528.
Upogebia synagelas, new species
Figs. 1-3
Material examined.—Jamaica: USNM
233572, 6 Holotype; USNM 233573, 2 Al-
lotype; USNM 233574, 1 6, 1 2, Paratypes;
Discovery Bay, 18°28'N, 77°24'W, spur and
groove reefs on Fore Reef, Long Term Sur-
vey site, 27.4-35 m (90-115 ft.), P. J. B.
Scott, SCUBA, 3 Oct 1985, host sponge
Agelas sceptrum. —USNM 233575, 2 6, 2 2
(ovig., | with eyed eggs, 1 with variably
hatched zoeae) Paratypes; Pear Tree Bot-
tom, 22 m, J. Woodley, 25 Nov 1985, oc-
curring as pairs in A. sceptrum.
Barbados: USNM 233576, 2 36, 1 8;
233577, 1 36, 3 2 ovig., Paratypes; ““Nurse’s
Jetty, on Bank Reef,’ 1 km W Holetown,
St. James, 13°10.7’N, 59°38.9'W, 18.3 m
(60 ft.), H. M. Reiswig collection No. 76-8-
9.1, SCUBA, 9 Aug 1976, host sponge A.
sceptrum (specimens in collection of Red-
path Museum, McGill University Cat. No.
RMI 2727).
MESC, Paratypes.
Florida Middle Grounds, coral reef, Gulf
of Mexico: Sta 151, 28°32’20’N, 84°
18’36”W: MESC 6183-4563, 1 juv., diver
collected, 27.4 m, 5 Oct 1978; MESC 6183-
VOLUME 100, NUMBER 3
Fig. 1.
4558, 1 juv., diver collected, 31.3-33 m, 17
Jan 1979; MESC 6183-4574, 5 juv., diver
collected, 27.4 m, 18 Jan 1979.—Sta 481,
28°30'52”N, 84°18'59"W: MESC 6183-
4589. 1 4, diver collected, 29 m, 6 Oct 1978;
MESC 6183-4556, 2 juv., diver collected 29
m, 8 Oct 1978; MESC 6183-4580, 2 6 Guv.),
diver collected, 29 m, 8 Oct 1978; MESC
6183-4587, 1 juv., diver collected, 29 m, 8
Oct 1978; MESC 6183-4552, 1 juv. (tiny),
submersible, 37.5 m, 8 Nov 1978.—Sta 491,
28°27'18"N, 84°17'02”W: MESC 6183-
Upogebia synagelas, 6 holotype: a, Cephalic region, lateral; b, Anterior carapace, dorsal; c, Mandible;
d, e, Maxilla 1, 2; f, g, h, Maxilliped 1, 2, 3. Scales all 1 mm: 1 = a—b; 2 = c; 3 = d-h.
4575, 1 8, diver collected, 29 m, 16 Oct
1978.—Sta 482, 28°31'06"N, 84°18'55’W:
MESC 6183-4578, 1 2 Guv.), diver collect-
ed, 33.5 m, 8 Jul 1979.—Sta 2315, 28°34’
O5’N, 84°14'11”"W: MESC 6183-10523, 1
juv., Capetown dredge, 38 m, 30 Aug 1977.
Bahamas: Freeport, Grand Bahama Is-
land: MESC 6183-10524, 1 juv., diver col-
lected, 21-23 m, Nov 1975; MESC 6183-
lOS2Z5,eleG ae oGlisovmic?))\. lmiras.vdiver
collected, 21—23 m, Nov 1975.
Diagnosis. — Projections to either side of
592
aN Sy SESS
E—SS TS oa Sv
Vv MyYy YY
ViViMay
1
Fign2
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Upogebia synagelas, 6 holotype: a, Cheliped, lateral; b, Cheliped, mesial; c, d, e, f, Legs 2-5; g, Parts
of abdominal segment 6, telson and uropods; h, Parts of lateral margin, segments 5—6. Scales all 1 mm: 1 = a—
(B= In,
rostrum ending in acute spine. Postocular
spine absent. First and second abdominal
segments lacking ventral spines; sixth seg-
ment bearing anterolateral process; telson
longer than wide, slightly narrowed poste-
riorly. Chelipeds with fully developed fixed
finger equal to dactyl in length, merus with
ventral row of low spines; remaining legs
Spineless.
Description.—Rostrum truncate, trian-
gular in dorsal view, slightly broader than
long, downturned tip exceeding eyestalks;
4—6 strong acute spines along each lateral
margin; dorsal surface bearing anterior mat
of long silky setae followed by field of scat-
tered spines angling toward sides posterior-
ly, gastric region smooth; ornamented part
separated from, and flanked on each side
by, slightly sinuous ridge bearing crest of
about 11 spines grading from slender an-
teriorly on lateral rostral process to sharp
tubercles posteriorly. Postorbital spine ab-
sent. Shoulder lateral to cervical groove
prominent; thalassinidean line faint but ex-
tending to posterior margin of carapace.
Abdomen weakly sclerotized; first 2 seg-
VOLUME 100, NUMBER 3
1
£——_ 3
Fig. 3.
593
4
Upogebia synagelas, Zoea I, just hatched or in process of hatching but still partly encapsulated in egg
membrane. a, Lateral view, semidiagrammatic; b, Dorsal view, cephalothorax and proximal part of abdomen;
c, Antennule; d, Antenna; e, Telson, composite of 2 specimens. Scales all 0.5 mm: 1 = a—b; 2 = c; 3 = d; 4 =e.
ments lacking spines or spinules on sterna,
pleura and bases of pleopods; pleura of seg-
ment 1 very poorly developed, of 2—5 rather
well developed and bearing plumose setae
on margins from middle of 2 to middle of
5, pilose tract extending dorsally to some
extent near anterior and posterior margin
of tergum on segment 3, scattered setae else-
where dorsally; segment 6 broader than long,
bearing slightly hooked process on antero-
lateral margin and behind it an irregularly
rounded lobe, marked dorsally on each side
with an obsolescent, irregularly lunate fur-
row.
Telson as long as uropods, slightly wider
than long, somewhat narrowed and rounded
posteriorly; median longitudinal furrow and
broader submarginal furrow at each side;
small acute spines irregularly placed along
lateral margins as well as submarginally and
irregularly on raised tracts, some spines
clustered in twos and threes.
Eyestalks rather stout, clearly exceeded
by rostrum; cornea large but narrower than
594
base of eyestalk in lateral view and directed
anterolaterally.
Antennular and antennal peduncles with
articles unspined; scale on latter minute.
Mouthparts as figured; epipod of maxil-
liped 1 minute, maxilliped 3 lacking epipod.
Extension of epistome spineless in lateral
view.
Chelipeds subequal, moderately robust,
depth of palm less than '2 length; fingers
elongate, gently curved toward each other
and tapering to tip, opposed edges entire;
carpus often with small distolateral, ventral
spine; merus bearing row of about 4—9 spines
on ventral margin. All walking legs with
smooth, spineless articles; they and che-
lipeds bearing usual tracts of long, silky se-
tae.
Two arthrobranchs arranged in biserial
rows of undivided (entire) lamellae on max-
illiped 3 and legs 1-4.
Pleopods densely setose, exopod far long-
er and broader than endopod.
Uropodal exopod with convex distal
margin, that of endopod less so, distal spine
on protopod lateral to articulation of en-
dopod; comparable acute spine proximally
on prominent rib of exopod; central rib of
endopod bearing irregular row of about 4—
12 or more small spines in distal half and
sometimes scattered spines elsewhere on
blade, exopod usually bearing sparse ob-
solescent spines on distolateral margin.
Measurements (mm).— Holotype ¢, length
anterior carapace 3.84, length carapace 6.34,
length right chela including fixed finger 4.8,
height 1.66. Allotype ovigerous ¢, same,
4.22, 6.49; 4.67, 1.34; 4.48, 1.34; eggs of
one individual approximately 0.9 x 1.04 in
diameter.
Known range. —Confined to the material
studied.
Color.—Chelipeds with distal parts of
palm colored shades of orange, fingers sim-
ilar but light at tips.
Remarks.— There is considerable varia-
tion in the amount of spination on the tail
fan and in ornamentation elsewhere; for ex-
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
ample, the cheliped merus of the holotype
lacks a dorsal subdistal tubercle whereas
such a tubercle is present on this member
in some individuals.
Chelae of selected juveniles are notice-
ably stouter than those of adults. For ex-
ample, in one juvenile with a carapace length
of 2.56 mm the right chela has a palm length
of 1.79 and a mid-palm height of 0.77; mid-
palm height of the right chela in the holo-
type is 0.35 the palm length, whereas in the
abovementioned juvenile the relationship is
0.43. In still another juvenile the relation-
ship is 0.53. The rostrum and eyes show
obvious ontogenetic changes from juveniles
of this size to the full adult phase; the eyes
are relatively large in juveniles, overshad-
owing the short narrow rostrum, though its
marginal spines are evident.
Upobegia synagelas most closely resem-
bles U. ramphula Williams (1986) in struc-
ture of the rostrum, abdomen, eyes, mouth-
parts, and walking legs. There are differences
in shape of the fingers (toothless in U. syn-
agelas) and in shape of elements of the tail-
fan which are spineless in U. ramphula.
Nevertheless, the similarity is striking, as in
many species pairs found on both sides of
Central America, and, though the exact hab-
itat of U. ramphula is presently unknown,
it may prove to be some species of sponge.
Most members of the genus Upogebia are
burrowers in subtidal substrates (Williams
1986). Some are highly specialized for bur-
rowing in stony corals (Kleemann 1984),
and still others are adapted for living in the
cavities of sponges (Barnard 1950).
Etymology. —A noun in apposition from
the Greek syn, with, and Agelas, the generic
name of the host sponge.
Notes on Zoea I
Fig. 3
Jamaican females studied (see above) are
Ovigerous, and one of them bears both
hatching and recently hatched zoeae I among
VOLUME 100, NUMBER 3
setae on the pleopods. Interesting compar-
isons can be made between these zoeae and
the zoea I of U. affinis (Say, 1818) described
and figured by Sandifer (1973). The larvae
figured here agree in general structure with
the latter but differ from them in lack of a
rostrum, in possession of developed pleo-
pods, in lack of incipient uropods which are
indicated in Sandifer’s Fig. 1A but not in
1B, and in pattern of spination on the tail
fan. Some of these differences may result
from a premature hatch due to handling.
Zoea I of U. affinis has 5 terminal spines on
each side of the telson and 2 or more smaller
lateral setae which may represent the distal
edge of the developing uropods. In contrast,
the telson of U. synagelas has 6 terminal
spines to either side of the midline. Differ-
ences in the admittedly variable abovemen-
tioned features may be additional evidence
of the clustered relationships among upo-
gebian species in the western hemisphere
(Williams 1986).
Acknowledgments
I am indebted to both P. J. B. Scott and
E. Sides for bringing this interesting species
to my attention, to P.J.B.S. for giving spec-
imens to the USNM, to M. R. Dardeau,
Dauphin Island Sea Lab, Dauphin Island,
Alabama, for loan of study specimens from
the MESC research collection, and to
95)
P.J.B.S., M.R.D., and B. B. Collette for
critical reading of the manuscript. Klaus
Rutzler gave information on the host. Keiko
Hiratsuka Moore carefully drew the adults
and inked drawings of the larvae.
Literature Cited
Barnard, K. H. 1950. Descriptive catalogue of South
African decapod Crustacea (crabs and
shrimps).—Annals of the South African Mu-
seum 38:1-837.
Kleemann, K. 1984. Lebensspuren von Upogebia
operculata (Crustacea, Decapoda) in karibisch-
en Steinkorallen (Madreporaria, Anthozoa).—
Beitrage zur Palaontologie von Osterreich, In-
stitut fur Palaontologie der Universitat Wien 1 1:
35-49.
Sandifer, P. A. 1973. Larvae of the burrowing shrimp,
Upogebia affinis, (Crustacea, Decapoda, Upo-
gebiidae) from Virginia plankton. — Chesapeake
Science 14(2):98-104.
Scott, P. J. B., H. M. Reiswig, and B. M. Marcotte. [In
press]. Ecology, functional morphology, behav-
iour and feeding in coral- and sponge-boring
species of Upogebia (Crustacea: Decapoda: An-
omura).— Canadian Journal of Zoology.
Williams, A. B. 1986. Mud shrimps Upogebia, from
the eastern Pacific (Thalassinoidea: Upogebi-
idae).—San Diego Society of Natural History,
Memoir 14:1-60.
National Marine Fisheries Service Sys-
tematics Laboratory, U.S. National Mu-
seum of Natural History, Washington, D.C.
20560.
PROC. BIOL. SOC. WASH.
100(3), 1987, pp. 596-599
RANGE EXTENSION OF THE GENUS
ASYMPHORODES MEYRICK
(LEPIDOPTERA: COSMOPTERIGIDAE)
J. F. G. Clarke
Abstract.— Asymphorodes aporema is described from Guam. The new species
is distantly removed from the Marquesas Islands, the center of distribution of
the genus Asymphorodes. Asymphorodes aporema is closely related to A. po-
liopterus but is distinguished from it by the strongly spined posterolateral mar-
gin of the sixth sternum and the annulated antenna. The lectotype of Stathopoda
monoxesta (Meyrick), new combination, is designated.
The genus Asymphorodes (Meyrick, 1929:
498-501) originally contained 13 species
from the Marquesas Islands, one from the
Paumotus (Tuamotus), one from Tahiti
(monoxesta) and one from Rapa (perfuga)
which Meyrick transferred from the genus
Ulochora.
Currently, the genus contains 75 species
from the Marquesas Islands (Clarke 1986:
180-283), two from Hawaii (Zimmerman
1978:1055-1067), one from Easter Island
(Clarke 1986:161), one from the Solomon
Islands, (Bradley 1957:100), and one from
the Tuamotus (Meyrick 1929:500).
Asymphorodes monoxesta Meyrick from
Tahiti is misplaced and is treated elsewhere
in this paper. Asymphorodes perfuga (Mey-
rick) from Rapa, was transferred to Stath-
mopoda (Clarke 1971:176).
Recently, while sorting species of Cos-
mopterigidae from Micronesia, an obvious
species of Asymphorodes from Guam was
encountered. Guam is ca. 3750 miles (6039
km.) from Hawaii and ca. 5250 miles (8454
km.) from the Marquesas Islands. These are
the nearest areas from which any species of
Asymphorodes has been reported.
The specimen was collected on an air-
plane in 1939, the “Philippine Clipper,”’ and
it is not clear whether the moth was a
stowaway or whether it boarded the air-
plane in Guam. The only place where the
Pan American Clipper Ships stopped on
their way to or from the Orient, and where
any species of Asymphorodes are known to
occur, 1s Hawaii. The two species of this
genus that occur in Hawaii are not closesly
related to the Guam species, so it is not
likely that the Guam specimen originated
there. The microlepidopterous fauna of
Guam is very poorly known and it is quite
possible that extensive collecting on that is-
land will reveal the presence of other species
of Asymphorodes, as well as additional spec-
imens of A. aporema.
In the meantime we must consider this
apparent great extension of range of the ge-
nus with some scepticism.
Asymphorodes aporema, new species
Figs. 1, 2
Description. — Alar expanse 6 mm. Labial
palpus shining white; second segment shad-
ed with fuscous on outer side; third segment
fuscous anteriorly and on outer side. An-
tenna, scape white; flagellum white annu-
lated fuscous. Head white. Thorax white;
thornlike processes of metascutum well de-
veloped. Forewing ground color white; from
base to apex a broad median, longitudinal
fuscous streak, edges of which not clearly
defined; cilia mixed white and grayish.
Hindwing sordid white; cilia grayish. Fore-
VOLUME 100, NUMBER 3
Se)
ig. 1.
leg white; femur, tibia and tarsal segments
fuscous on outer side; midleg white; tarsal
segments slightly infuscated; hindleg white;
tibia slightly infuscated dorsally; tarsal seg-
ments slightly infuscated. Abdomen fus-
cous above, whitish ventrally; segments 6,
7 and 8 strongly modified (see figures).
Male genitalia slide USNM 26411. Harpe
broad basally; neck moderately narrow; cu-
cullus moderately dilated distally. Brachia
unequal, right brachium longer than left,
distally hooked. Tegumen shorter than
broad. Aedeagus slightly S-shaped, slender.
Manica thickened distally.
Holotype.—USNM. Unique male holo-
type, 15 Mar 1939, Guam No. 1912, R. G.
Oakley.
Type locality.—Guam (no specific local-
ity).
Distribution.— Guam.
Food plant.— Unknown.
Remarks.—The nearest relative to A.
aporemais A. poliopterus Clarke (1986:241)
from the Marquesas Islands. In the male of
poliopterus the antenna is cream white but
the antenna of aporema is white, annulated
with fuscous; also, the fuscous marking of
Asymphorodes aporema, new species, right wings.
the forewing of aporema is confined to a
median longitudinal streak but in poliop-
terus the fuscous marking covers all of the
wing except the dorsal white edge. The mod-
ifications of the abdominal segments are
similar, but distinct. Compare figures of
aporema with those of poliopterus (Clarke
1986: fig. 168 c-e).
Stathmopoda monoxesta (Meyrick),
new combination
Asymphorodes monoxesta Meyrick, 1929:
501. “42. 11-13 mm.... Society Is. Ta-
hiti, SOO—2500 feet, March to May (Miss
Cheesman, Collenette); 9 ex.”
Lectotype.—é, 12mm. “‘Fautaua, Tahiti,
at light 2500 ft. 13.3.25. St. George Expedn.
Ge Collenette. slide JEGCe Now Wisi9:
Lectotype hereby designated. This speci-
men is marked “Type 2” in the British Mu-
seum. A small white label bears the number
srlzo ON
Type.— British Museum (Natural Histo-
ry).
Type locality.—Tahiti, Fautaua.
Distribution. — Tahiti.
598
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig. 2. Asymphorodes aporema, new species: a, Lateral aspect of male genitalia with aedeagus removed; b,
Aedeagus; c, 6th tergum; d, 7th tergum and sternum; e, 8th sternum.
Remarks.—The fact that the lectotype was
marked “Type 9”’ presents no problem be-
cause the gender of many “types”’ from the
Meyrick collection was mislabelled.
Acknowledgments
I wish to thank Miss Ann Whyte, Man-
ager, Pan American World Airways, New
York, for providing information on the
routes of the “Clipper Ships” during the
years 1939-40. I am indebted to Mrs. Nan-
cy McIntyre for calculating the distances in-
dicated. Also, I wish to express my appre-
ciation to Victor Krantz for the photograph
of the moth wings, to Elaine R. S. Hodges
for the drawings and to Silver West for typ-
ing the manuscript, all members of the
Smithsonian staff.
Literature Cited
Bradley, J. D. 1957. Microlepidoptera from Rennell
and Bellona Islands. Pages 87-112, plates 1-12
in The Natural history of Rennell Island, British
Solomon Islands, volume 2(19).
Clarke, J. F.G. 1971. The Lepidoptera of Rapa Is-
land.—Smithsonian Contributions to Zoology
56:1—282,figs. 1-175, pls. 1-29.
VOLUME 100, NUMBER 3 599
. 1986. Pyralildae and Microlepidoptera of the pl. 1-7; (2):883-1903, fig. 608-1355. University
Marquesas Archipelago.—Smithsonian Contri- of Hawaii Press. Honolulu.
butions to Zoology 416:1—485, figs. 1-319.
Meyrick, E. 1929. The Micro-Lepidoptera of the “‘St.
George” Expedition.— Transactions of the En- Department of Entomology, National
tomological Society of London 76:489-521. : : :
Zimmerman, E. C. 1978. Microlepidoptera. Jn In- Museum of Natural History, Smithsonian
sects of Hawaii 9(1):i-xviii, 1-882, fig. 1-607, Institution, Washington, D.C. 20560.
PROC. BIOL. SOC. WASH.
100(3), 1987, pp. 600-602
REIDENTIFICATION OF DAVID CAUSEY’S CALIGUS
COLLECTIONS (CRUSTACEA: COPEPODA)
Roger Cressey and Patricia Nutter
Abstract.—The David Causey collections of Caligus from the Gulf of Mexico
and Caribbean Sea housed at the University of Arkansas are reidentified and
names are changed for about half of the samples.
During Causey’s tenure at the University
of Arkansas he authored four papers re-
porting on parasitic copepods from the Gulf
of Mexico, Caribbean Sea and the west coast
of Mexico (Causey 1953a, b, 1955, 1960).
The reidentifications reported here are only
of those from the Gulf and Caribbean. The
identifications of the west coast of Mexico
material will follow a study by the first au-
thor of all species of Ca/igus from the east-
ern Pacific Ocean.
Our report on the collection is arranged
chronologically by Causey publication date.
There are six previously unreported collec-
tions included herein. Also, some collec-
tions reported by Causey are not present in
the collection.
We thank Dr. Nancy Glover McCartney,
Curator of Zoology at the University of Ar-
kansas Museum for bringing this collection
to our attention and for loaning us the ma-
terial for study. Other parasitic copepod
material of Causey’s is also present in
U.A.M. collections and available for study.
All University of Arkansas Museum
numbers in the table should be prefaced by
78-93-.
Causey new
UAM no. identification identification
78-93-183 C. bennetti Scianophilus tenuis
184 C. bennetti Scianophilus tenuis
194 C. haemulonis Lepeophthirus sp.
Doe C. pelamydis Scianophilus tenuis
235 C. productus Caligus bonito
236 C. productus Caligus bonito
239 C. productus Caligus bonito
Additional records not in the collection
Parasitic Copepoda of Texas coastal fishes
Causey new
UAM no. identification identification
179 C. aliuncus Tuxophorus caligoides
195 C. haemulonis Caligus haemulonis
196 C. haemulonis Lepeophthirus sp.
196 Caligus haemulonis
214 C. pelamydis caligid, not Caligus
DNS) C. pelamydis Scianophilus tenuis
216 C. pelamydis Tuxophorus collettei
ANY C. pelamydis Tuxophorus collettei
Parasitic Copepoda from Grande Isle, Louisiana
host
Lobotes surinamensis
Lobotes surinamensis
Scianops ocellatus
Pogonias cromis
Coryphaena hippurus
Coryphaena hippurus
Coryphaena hippurus
host
Rachycentron canadum
Galeichthys felis
Galeichthys felis
Pogonias cromis
Pogonias cromis
Sarda sarda
Scomberomorus cavalla
VOLUME 100, NUMBER 3
218 C. pelamydis
219 C. pelamydis
229 C. praetextus
254 C. rapax
DS C. robustus
258 C. robustus
Tuxophorus collettei
Scianophilus tenuis
Caligus praetextus
Caligus n. sp.
Caligus robustus
Caligus robustus
Additional records not in the collection
601
Scomberomorus cavalla
Pogonias cromis
Galeichthys felis
Scomberomorus maculatus
Caranx hippos
Caranx hippos
Parasitic Copepoda from Gulf of Mexico fish
Causey new
UAM no. identification identification host
185 C. bennetti Scianophilus tenuis Lobotes surinamensis
197 C. haemulonis Caligus haemulonis Galeichthys felis
198 C. haemulonis Caligus haemulonis Galeichthys felis
199 C. haemulonis Caligus haemulonis Galeichthys felis
200 C. haemulonis Caligus haemulonis Bagre marinus
203 C. haemulonis Caligus haemulonis Pogonias cromis
207 C. mutabilis Caligus mutabilis Trachinotus carolinensis
220 C. pelamydis Scianophilus tenuis Pogonias cromis
VAI C. pelamydis Scianophilus tenuis Pogonias cromis
224 C. pelamydis Caligus mutabilis Scomberomorus maculatus
D5) C. pelamydis Scianophilus tenuis Pogonias cromis
226 C. pelamydis Caligus mutabilis Scomberomorus maculatus
228 C. pelamydis Scianophilus tenuis Pogonias cromis
230 C. praetextus caligid—not Caligus Bagre marinus
231 C. praetextus Caligus sp. male Bagre marinus
232 C. praetextus caligid—not Caligus Bagre marinus
233 C. praetextus caligid—not Caligus Bagre marinus
240 C. productus Caligus bonito Coryphaena hippurus
241 C. productus Scianophilus tenuis Pogonias cromis
262 C. setosus Caligus haemulonis Galeichthys felis
Additional records not in collection
Parasitic Copepoda from Mexican coastal fishes
Causey new
UAM no. identification identification host
192 C. bennetti Scianophilus tenuis Kyphosus sectatrix
204 C. haemulonis Caligus haemulonis ‘““‘bandara-a cat”’
All other material in the collection reported in this paper is from the
Pacific coast of Mexico.
Unreported collections of interest
Causey new
UAM no. identification identification host
181 C. balistae Caligus balistae Canthidermis sabaco
186 C. bonito Caligus chelifer Brevoortia tyrannus
206 C. latifrons Caligus praetextus Scianops ocellatus
234 C. praetextus Caligus atromaculatus Lutianus griseus
237 C. productus Caligus bonito “cutlass fish”’
261 C. setosus Caligus haemulonis Bagre marinus
Collection 181 is from Grande Isle, Lou-
isiana. Collection 234 is from Alligator Har-
bor, Florida. Collections 186, 206, 237, and
261 are from Port Aransas-Laguna Madre,
Texas.
Literature Cited
Causey, D. 1953a. Parasitic Copepoda from Grande
Isle, Louisiana.—Occasional Papers of the Ma-
rine Laboratory of Louisiana State University
7:1-18.
602
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
1953b. Parasitic Copepoda of Texas coastal
fishes. — Publication of the Institute of Marine
Sciences, University of Texas 3:6—-16.
. 1955. Parasitic Copepoda from Gulf of Mex-
ico fish.— Occasional Papers of the Marine Lab-
oratory of Louisiana State University 9:1-19.
1960. Parasitic Copepoda from Mexican
coastal fishes.— Bulletin of Marine Sciences of
the Gulf and Caribbean 10:323-337.
Department of Invertebrate Zoology, Na-
tional Museum of Natural History, Smith-
sonian Institution, Washington, D.C. 20560.
PROC. BIOL. SOC. WASH.
100(3), 1987, pp. 603-609
TWO NEW SPECIES OF CURIMATID FISHES
(OSTARIOPHYSI: CHARACIFORMES) FROM
RIO GRANDE DO SUL, BRAZIL
Richard P. Vari
Abstract.—Curimata spilota is described from two localities in the Rio Uru-
guay basin in southeastern Brazil. The species is unique in the family in its
higher number of branched dorsal-fin rays (10 or 11). Curimata stigmosa is
described from the coastal rivers of southeastern Brazil and Uruguay and from
one locality in the Rio Uruguay basin, and is the first member of its phyletic
lineage known from that region. The possession of lobulate fleshy processes on
the roof of the oral cavity, the distinctive pigmentation pattern, and meristic
values distinguish the species within the family.
Recent collecting efforts by Carlos A. S.
Lucena, Luiz R. Malabarba, and Roberto
E. Reis of the Museu de Ciéncias of the
Pontificia Universidade Catolica do Rio
Grande do Sul, Rio Grande do Sul, Brazil
have produced a number of interesting
species. Included in those collections are two
undescribed species belonging to separate
lineages within the characiform family Cu-
rimatidae. The family is the subject of on-
going phylogenetic and revisionary studies
(Vari 1982, 1983, 1984), but the species in
question are members of genera that will
not be published on for some time. In order
to make the names of these species available
for ongoing ecological and faunal studies,
those taxa are described separately in this
paper.
Counts and measurements in the species
descriptions follow the methods outlined in
Vari (1982, 1984). Ranges of counts and
measurements include values of all speci-
mens. Values in square brackets are those
of the holotype. Specimens examined for
this study are deposited in the following in-
stitutions: Museu de Ciéncias da Univer-
sidade Catolica do Rio Grande do Sul, Por-
to Alegre, Rio Grande do Sul, Brazil, MCP;
Museu de Zoologia da Universidade do Sao
Paulo, Sao Paulo, Brazil, MZUSP; and Na-
tional Museum of Natural History, Smith-
sonian Institution, Washington, D.C.,
USNM.
A variety of very different classifications
have been utilized by authors publishing on
the Curimatidae. Under these classifica-
tions, the number of genera recognized in
the family has ranged from seven (Eigen-
mann 1910) to 29 (Fernandez-Yepez 1948).
Research completed (Vari 1982, 1984) or
in progress, attempts to redefine subunits of
the family along natural lines, and has or
will result in the synonymization of nu-
merous genera and the redefinition of those
taxa that will be recognized. Pending com-
pletion of those studies, the species de-
scribed herein are retained in the broadly
encompassing genus Curimata which has
included the bulk of the species of the family
in the more widely used classificatory
schemes.
Curimata spilota, new species
Figs. 1, 2
Holotype.—Brazil, Rio Grande do Sul,
Rio Santa Maria, at bridge on highway Br
293, between Dom Pedrito and Livramento
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
igeyale
Curimata spilota, holotype, MZUSP 37133, 56.9 mm SL; Brazil, Rio Grande do Sul, Rio Santa
Maria, at bridge on highway Br 293, between Dom Pedrito and Livramento.
(listed as Santana do Livramento on some
maps), C. Lucena and L. Malabarba, 22 Jan
1982, MZUSP 37133, 56.3 mm.
Paratypes.— Brazil, Rio Grande do Sul,
taken with holotype, MZUSP 37134, 7,
34.7—43.4 mm; USNM 285194, 12, 36.9-
59.7 mm (2 specimens cleared and coun-
terstained for cartilage and bone).—Brazil,
Rio Grande do Sul, headwaters of Rio Ne-
gro, at Bage, C. Lucena and L. Malabarba,
25 Oct 1983, MCP 9613, 4, 43.4-62.8 mm.
Diagnosis.—The possession of 10 or 11
(typically 10) branched dorsal-fin rays dis-
tinguishes Curimata spilota from all other
members of the family Curimatidae which
have 8 or 9, rarely 10, branched dorsal-fin
rays. The few species of curimatids reported
as having 10 branched dorsal-fin rays in oc-
casional individuals are members of the ge-
nus Potamorhina (Vari, 1984), all of which
have 75 or more lateral line scales, contrary
to the 30 to 33 scales in that series in Cu-
rimata spilota. The low lateral line count
(30 to 33) of C. spilota also distinguishes
the species from the vast majority of curi-
matids. The Curimatopsis species with such
low lateral line counts (evelynae, macrole-
pis, crypticus, myersi) all have an anteriorly
convex ventral portion of the maxilla that
is lacking in Curimata spilota, along with
pronounced sexual dimorphism that is also
absent in the latter species (see Vari 1983:
4—11).
Within the LaPlata drainage system, three
species are phenetically similar to C. spilota
and have comparable scale counts. The first
of these, C. gillii, described by Eigenmann
and Kennedy (1903:510) from the Rio Par-
aguay in Paraguay, differs in having fewer
branched dorsal-fin rays (8 or 9) than C.
spilota (10 or 11). The second similar species
in that basin is Curimata saladensis, de-
scribed by Meinken (1933) in Curimatopsis.
That species, which is not closely related
phylogenetically to Curimatopsis (see Vari
1983:10-12), can be readily distinguished
from C. stigmosa in having 8 or 9 branched
dorsal-fin rays, and having the pores of the
laterosensory canal system of the body de-
veloped only on the anterior 7 to 9 scales,
contrary to the completely developed series
of such pores in C. spilota. Curimata Stig-
mosa, a partially sympatric species de-
scribed in this paper, has very similar lateral
line counts to C. spilota, but is distinguished
by the presence of fleshy lobulate processes
on the roof of the mouth.
Description.— Body moderately elongate,
somewhat compressed, more so in juve-
niles. Dorsal profile of head convex ante-
riorly, straight from above nostrils to rear
of head. Dorsal profile of body slightly con-
VOLUME 100, NUMBER 3
vex from rear of head to origin of rayed
dorsal fin; straight and slightly posteroven-
trally slanted at base of dorsal fin, gently
convex from base of last dorsal-fin ray to
caudal peduncle. Dorsal surface of body
transversely rounded anteriorly, with indis-
tinct median keel immediately anterior to
rayed dorsal fin, smoothly rounded trans-
versely posterior to fin. Ventral profile of
body gently curved from tip of lower jaw to
caudal peduncle. Prepelvic region obtusely
flattened, with median series of scales prox-
imate to pelvic fin origin. An obtuse median
keel posterior to pelvic fin insertion.
Greatest body depth at origin of rayed
dorsal fin, depth 0.33-—0.41 [0.39]; snout tip
to origin of rayed dorsal fin 0.50—0.53 [0.52];
snout tip to origin of anal fin 0.82—0.86
[0.84]; snout tip to insertion of pelvic fin
0.55-0.58 [0.58]; snout tip to anus 0.79-
0.82 [0.81]; origin of rayed dorsal fin to hy-
pural joint 0.53-0.59 [0.53]. Rayed dorsal-
fin margin rounded; anteriormost rays three
to three and one-half times length of ulti-
mate ray. Pectoral fin pointed; length of pec-
toral fin 0.18—0.21 [0.21], extending two-
thirds to three-quarters distance to vertical
through insertion of pelvic fin. Pelvic fin
pointed, length of pelvic fin 0.22-0.26 [0.25],
reaching approximately two-thirds distance
to origin of anal fin. Caudal fin deeply forked.
Adipose dorsal fin well developed. Anal fin
emarginate, anteriormost branched rays
about two and one-half times length of ul-
timate ray. Caudal peduncle depth 0.14—
Ort Sa(O. 15):
Head obtusely pointed, head length 0.28-—
0.32 [0.31]; upper jaw slighty longer, mouth
subterminal; snout length 0.26—-0.31 [0.31];
nostrils very close, anterior circular, pos-
terior crescent shaped with aperture closed
by thin flap of skin separating nares; orbital
diameter 0.30—0.36 [0.30]; adipose eyelid
present, more developed anteriorly, with
broad vertically ovoid opening over center
of eye; length of postorbital portion of head
0.40-0.45 [0.45]; gape width 0.20-0.25
[0.22]; interorbital width 0.40-0.46 [0.42].
ATLANTIC
OCEAN
Fig. 2. Map of major drainages in southeastern
Brazil, Uruguay, and proximate portions of Argentina
showing collecting localities of specimens reported on
in this paper (all localities in Brazil, state of Rio Grande
do Sul unless otherwise noted): 1, Rio Santa Maria
(holotype and paratypes of Curimata spilota); 2, upper
Rio Negro, at Bagé (paratypes of C. spilota); 3, Rio
Jacui (holotype and paratypes of Curimata stigmosa);
4, Rio Forqueta (paratypes of C. stigmosa); 5, Arroio
Sarandi and Arroio Chasqueiro (paratypes of C. stig-
mosa); 6, Brazil, state of Santa Catarina, Rio Uruguai
near Concordia (C. stigmosa); and 7, Uruguay, Florida,
Arroio Chamizo (C. stigmosa).
Pored lateral line scales from supracleith-
rum to hypural joint 30 to 33 [31]; all scales
of lateral line pored, canals in scales straight;
2 to 4 series of scales extending beyond hy-
pural joint onto caudal fin base; 6'2 [614]
scales in transverse series from origin of
rayed dorsal fin to lateral line; 42 or 5 (5
rare) [4'4] scales in transverse series from
lateral line to origin of anal fin.
Dorsal-fin rays 11,10 or 11, or 111,10 or 11
(1,11 and 11,11 less common; when three
unbranched rays present, first ray very short)
[111,10]; anal-fin rays 11,7 or 11,7 (when three
unbranched rays present, first ray very
short)[111,7]; pectoral-fin rays 13 to 15 [14];
pelvic fin rays 1,8 [1,8].
Total vertebrae 30 (4), 31 (20), 32 (3).
Color in alcohol.—Specimens retain some
guanine on scales; silvery overall, purplish
dorsally and whitish ventrally. Head darker
606
Fig. 3.
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Curimata stigmosa, holotype, MZUSP 37135, 73.4 mm SL; Brazil, Rio Grande do Sul, Rio Jacui,
at bridge on the road between Santa Maria and Veracruz.
dorsally. Iris, opercle and ventral portions
of head silvery. Obscure midlateral dusky
band extending from supracleithrum to cau-
dal peduncle. Horizontally elongate mid-
lateral spot on caudal peduncle. Spot some-
what diffuse, more apparent in individuals
in which guanine is largely lacking. Median
fins with small chromatophores scattered
over membranes; dusky overall. Adipose
dorsal fin dusky. Paired fins hyaline.
Distribution.—Rio Uruguay basin (Fig.
2). The two localities from which this species
is known (localities | and 2, Fig. 2) although
relatively close, lie within the basins of two
different tributaries of the Rio Uruguay
whose mouths are distant from each other.
This occurrence of the species in the head-
waters of those two rivers may indicate that
the species is widely distributed through the
Rio Uruguay basin.
Etymology.—The specific name, spilota,
from the Greek for spotted or stained, refers
to the midlateral spot on the caudal pedun-
cle.
Curimata stigmosa, new species
Figs. 2, 3
Holotype.— Brazil, Rio Grande do Sul,
Rio Jacui, at bridge on the road between
Santa Maria and Veracruz (approx. 29°41'S,
53°19'W), C. Lucena, L. Malabarba, and R.
Reis, 16 Sep 1983, MZUSP 37135, 73.4
mm SL.
Paratypes.—(All specimens collected in
Brazil, Rio Grande do Sul, unless noted oth-
erwise), taken with holotype, USNM
285193, 3, 65.5-73.6 mm (one specimen
cleared and counterstained for cartilage and
bone); MCP 9224, 3, 58.1-70.4 mm;
MZUSP 37136, 3, 52.5-74.3 mm.—Rio
Forqueta, and Marqués de Souza, Munici-
pio de Lajeado, MZUSP/USNM expedi-
tion, 7 Nov 1979, USNM 285191, 1, 53.7
mm; MZUSP 21721, 1, 53.4 mm.—Arroio
Sarandi, along highway (Br 116) between
Pelotas and Jaguarao, MZUSP/USNM ex-
pedition, 14 Dec 1978, MZUSP 21728, 1,
43.0 mm.—Arroio Chasqueiro (empties into
Lagoa Mirim north of Arroio Grande), along
highway (Br 116) between Pelotas and Ja-
guarao, MZUSP/USNM expedition, 14 Dec
1978, USNM 285190, 2, 43.5-56.5 mm.
The following non-typic specimens were
also examined: Brazil, Santa Catarina, pools
along Rio Uruguai, near Concordia, MZUSP
28252, 4, 71.5-72.3 mm.— Uruguay, Flor-
ida, Arroio Chamizo, USNM 285192, 2,
64.2-70.4 mm. |
Diagnosis.—The presence of a series of
lobulate fleshy processes in the roof of the
mouth of Curimata stigmosa distinguishes
the species from the majority of curimatids
with the exception of those species phylet-
VOLUME 100, NUMBER 3
ically aligned with Curimata elegans Stein-
dachner (1874). The wide deep-lying stripe
of dark pigmentation is distinctive for C.
stigmosa in that assemblage. Only three oth-
er species with lobulate processes on the
roof of the oral cavity inhabit river systems
proximate to the known distribution of C.
stigmosa. These are C. elegans Steindachner
of the coastal drainages of Brazil north of
Sao Paulo state, C. insculpta (Fernandez-
Yepez, 1948) an inhabitant of the Rio Pa-
rana above Sete Quedas rapids, and C. ni-
tens Holmberg (1891) of the Paraguay, low-
er Parana and Uruguay rivers. Curimata
insculpta is readily distinguished from C.
stigmosa by differences in lateral line scale
counts (37 to 42 versus 30 to 33). Curimata
nitens, which is sympatric with C. stigmosa
in the Uruguay basin, differs from the latter
species in having an intense black stripe
along the entire extent of the lateral line,
and in lacking the wide deep-lying band of
pigmentation on the caudal peduncle char-
acteristic of the latter species. Curimata ele-
gans similarly lacks the pronounced deep-
lying wide band on the caudal peduncle.
Description.— Body moderately deep,
more so in larger specimens, somewhat
compressed. Dorsal profile of head convex
anteriorly, straight from over nostrils to rear
of head. Dorsal profile of body gently con-
vex from rear of head to origin of rayed
dorsal fin; straight and slightly posteroven-
trally slanted at base of dorsal fin, straight
or gently convex from base of last dorsal-
fin ray to caudal peduncle. Dorsal surface
of body transversely rounded anteriorly,
with indistinct median keel immediately
anterior to rayed dorsal fin, smoothly
rounded transversely posterior to fin. Ven-
tral profile of body gently curved from tip
of lower jaw to caudal peduncle. Prepelvic
region obtusely flattened proximate to pel-
vic fin origin. Obtuse median keel posterior
to pelvic fin insertion. Secondary obtuse keel
on each side of postventral portion of body
one scale dorsal of ventral midline.
Greatest body depth at origin of rayed
607
dorsal fin, depth 0.35—0.40 [0.40]; snout tip
to origin of rayed dorsal fin 0.48-0.53 [0.49];
snout tip to origin of anal fin 0.80-0.84
[0.82]; snout tip to insertion of pelvic fin
0.53-0.57 [0.56]; snout tip to anus 0.76—-
0.79 [0.78]; origin of rayed dorsal fin to hy-
pural joint 0.52-0.57 [0.55]. Rayed dorsal-
fin margin rounded; anteriormost rays three
to three and one-half times length of ulti-
mate ray. Pectoral fin pointed; length of pec-
toral fin 0.17-0.21 [0.19], extending about
two-thirds distance to vertical through in-
sertion of pelvic fin. Pelvic fin pointed,
length of pelvic fin 0.19-0.22 [0.22], reach-
ing about two-thirds distance to origin of
anal fin. Caudal fin forked. Adipose dorsal
fin well developed. Anal fin emarginate, an-
teriormost branched rays about two and one-
third to two and two-thirds times length of
ultimate ray. Caudal peduncle depth 0.12-
0.14 [0.12].
Head obtusely pointed, head length 0.25-
0.29 [0.26]; upper jaw slightly longer, mouth
subterminal; snout length 0.29-0.33 [0.30];
nostrils very close, anterior circular, pos-
terior crescent shaped with aperture closed
by thin flap of skin separating nares; orbital
diameter 0.30-0.36 [0.31]; adipose eyelid
present, more developed anteriorly, with
vertically ovoid opening over center of eye;
length of postorbital portion of head 0.39-
0.44 [0.39]; gape width 0.25-0.29 [0.28];
interorbital width 0.44—0.50 [0.44].
Pored lateral line scales from supracleith-
rum to hypural joint 31 to 34 [32]; all scales
of lateral line pored, canals in scales straight;
2 to 4 series of scales extending beyond hy-
pural joint onto caudal fin base; 512 to 6
[5'4] scales in transverse series from origin
of rayed dorsal fin to lateral line; 412 [414]
scales in transverse series from lateral line
to origin of anal fin.
Dorsal-fin rays 11,9 or 111,9 (41,9 rare) [11,9];
anal-fin rays 11,7 [11,7]; pectoral-fin rays 13
to 15 [14]; pelvic-fin rays 1,8 or 9 [1,8].
Total vertebrae 33 (21), 34 (1).
Color in alcohol.—Overall ground col-
oration of specimens lacking guanine on
608
scales tan to tannish-brown, darker on dor-
sal portions of head and body. Obscure mid-
dorsal band from rear of head to origin of
rayed dorsal fin, and between rayed and adi-
pose dorsal fins. Scales of dorsal portion of
body with field of dark chromatophores on
exposed surface; field more extensive on
dorsal scales. Lateral line scales with pores
surrounded by patches of dark pigmenta-
tion. Deep-lying dark band extends from
vertical through origin of dorsal fin poste-
riorly to somewhat beyond hypural joint;
both deep-lying stripe and surface chro-
matophores forming dark, anteriorly point-
ed slightly elongate spot on lateral surface
of caudal peduncle. Field of small chro-
matophores continuing posteriorly on body
surface to base of middle caudal-fin rays.
Median fins somewhat dusky; other fins
hyaline.
Distribution.— Atlantic coastal drainages
of Rio Grande do Sul in Brazil, Uruguay,
and Rio Uruguai in Santa Catarina, Brazil
(Fig. 2).
Etymology.—The specific name, stig-
mosa, from the Latin for full of marks, re-
fers to the series of dark spots surrounding
the pores of the lateral line scales.
Relationships.— Curimata stigmosa is the
first member of the Curimata elegans Stein-
dachner (1874) lineage that has been col-
lected in the coastal drainages of far south-
eastern Brazil and Uruguay. The lineage,
characterized by the presence of fleshy lob-
ulate processes on the roof of the oral cavity,
is widely distributed through the freshwa-
ters of South America. Three members of
the lineage occur in regions close to that
inhabited by C. stigmosa (see ‘“‘Diagnosis”’
for that species); however, the exact phy-
logenetic relationships of C. stigmosa to
these and other members of the lineage are
unknown.
Resumo
Duas novas espécies de Characiformes da
familia Curimatidae (Pisces: Ostariophysi)
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
do Estado do Rio Grande do Sul, no sudeste
do Brasil, sao descritas. Curimata spilota é
descrita dos rios Santa Maria e Negro, tri-
butarios do rio Uruguai. A espécie é unica
na familia pelo numero mais elevado de
raios ramificados na nadadeira dorsal (10
ou 11). Curimata stigmosa é descrita dos
rios da regiao costeira atlantica do Uruguai
e sudeste do Brasil; e de uma localidade na
bacia do rio Uruguai. A espécie é€ caracter-
izada por processos lobulares carnosos no
teto da cavidade oral e um padrao de pig-
menta¢ao distinto. Estas caracteristicas e
valores meristicos distinguem a espécie
dentro da familia. A posse dos processos na
cavidade oral € caracteristica unica do com-
plexo Curimata elegans. Nenhum membro
deste grupo foi anteriormente assinalado
para a area habitada por C. stigmosa.
Acknowledgments
The specimens that served as the basis
for this paper were made available by Carlos
A. S. Lucena, Luiz R. Malabarba, and Rob-
erto E. Reis of MCP, and Heraldo A. Britski
and Naércio A. Menezes of MZUSP. I also
thank the latter two individuals and Jose
Lima de Figueirido of the same institution
for their hospitality during my visits. Tech-
nical assistance at the Smithsonian was pro-
vided by Andrew G. Gerberich and Jeffrey
Clayton. Figures 1 and 3 were prepared by
Theophilus Britt Griswald. The map is a
modification of a base map prepared by
Marilyn J. Weitzman. This paper benefitted
from the comments and suggestions of Stan-
ley H. Weitzman, Wayne C. Starnes, Ri-
cardo M. C. Castro, and Ann M. Williams.
Prof. Castro also provided the Portuguese
translation of the ““Resumo.” I thank all of
the above for their assistance and interest.
Research associated with this study and
travel expenses for museum studies in Bra-
zil were partially supported by the Neo-
tropical Lowland Research Program of the
International Environmental Sciences Pro-
gram of the Smithsonian Institution.
VOLUME 100, NUMBER 3
Literature Cited
Eigenmann, C.H. 1910. Catalogue of the fresh-water
fishes of tropical and south temperate Ameri-
ca.— Reports of the PrincetonUniversity Expe-
ditions to Patagonia, 1896-1899, 3 (Zoology, 4):
SIS= Ne
, and C. H. Kennedy. 1903. On a collection
of fishes from Paraguay, with a synopsis of the
American genera of cichlids.— Proceedings of
the Academy of Natural Sciences of Philadel-
phia 55(2):497-537.
Fernandez-Yépez, A. 1948. Los Curimatidos (peces
fluviales do Sur América). Catalogo descriptivo
con nuevas adiciones genéricas y especificas. —
Boletin Taxonomico del Laboratorio de Pes-
queria de Caiquire 1:1-86.
Holmberg, E. L. 1891. Sobre algunos peces nuevos
0 poco conocidos de la Republica Argentina. —
Revista Argentina de Historia Natural 1:180-
193.
Steindachner, F.
1874. Die Siisswasserfishe der Su-
609
dostlichen Brasiliens.—Sitzungsberichte der
Akademie der Wissenschaften, Wien 70:499—
538.
Vari, R. P. 1982. Systematics of the Neotropical char-
acoid genus Curimatopsis (Pisces: Characoid-
e1). —Smithsonian Contributions to Zoology 373:
1-28.
1983. Phylogenetic relationships of the fam-
ilies Curimatidae, Prochildontidae, Anostomi-
dae, and Chilodontidae (Pisces: Characi-
formes).—Smithsonian Contributions to
Zoology 378:1-—60.
1984. Systematics of the Neotropical char-
aciform genus Potamorhina (Pisces: Characi-
formes).—Smithsonian Contributions to Zool-
ogy 400:1-36.
Department of Vertebrate Zoology (Fish-
es), National Museum of Natural History,
Smithsonian Institution, Washington, D.C.
20560.
PROC. BIOL. SOC. WASH.
100(3), 1987, pp. 610-623
DEEVEYA BRANSONI, A NEW SPECIES OF
TROGLOBITIC HALOCYPRID OSTRACODE
FROM ANCHIALINE CAVES ON SOUTH
ANDROS ISLAND, BAHAMAS
(CRUSTACEA: OSTRACODA)
Louis S. Kornicker and R. J. Palmer
Abstract. —Deeveya bransoni, a new species of troglobitic halocyprid ostra-
code in the subfamily Deeveyinae, from anchialine caves on South Andros -
Island, Bahamas, is described and illustrated. A supplementary description is
presented of the type species of the genus, Deeveya spiralis Kornicker and Iliffe,
1985, based on type specimens from the Turks and Caicos Islands.
The new species described herein increas-
es to five the known number of trogolobitic
ostracodes in the Halocypridoidea: 1, Dan-
ielopolina orghidani (Danielopol, 1972)
from Cuba; 2, a species of Spelaeoecia from
Bermuda described by Angel and Iliffe
(1987) and mentioned by Angel (1983:531);
3, Deeveya spiralis Kornicker and Iliffe,
1985, from the Turks and Caicos Islands;
4, Danielopolina wilkensi Hartmann, 1985,
from the Canary Islands; and 5, Deeveya
bransoni, new species, from the Bahamas.
A supplementary description is presented
of Deeveya spiralis Kornicker and Iliffe,
1985.
Superfamily Halocypridoidea
Dana, 1853
Composition. —The superfamily com-
prises the families Halocyprididae Dana,
1853, and Thaumatocyprididae Muller,
1906.
Family Halocyprididae Dana, 1853
Composition. —The family comprises the
subfamilies Halocypridinae Dana, 1852,
Conchoecinae Claus, 1891, Archiconchoe-
cinae Poulsen, 1969, Euconchoecinae Poul-
sen, 1969, and Deeveyinae Kornicker and
Iliffe, 1985.
Subfamily Deeveyinae
Kornicker and Iliffe, 1985
Composition. —The subfamily comprises
the genera Deeveya Kornicker and Iliffe,
1985, and Spelaeoecia Angel and Iliffe,
1987.
Deeveya Kornicker and Iliffe, 1985
Type species. — Deeveya spiralis Kornick-
er and Iliffe, 1985.
Composition. —The genus comprises two
species from marine caves: D. spiralis from
the Turks and Caicos Islands, and D. bran-
soni, a new species from South Andros Is-
land, Bahamas.
Terminology. —The distribution of bris-
tles on joints 7 and 8 of the Ist antenna of
Deeveya appears quite similar to those of
members of the Myodocopina; therefore, the
lettering system proposed by Skogsberg
(1920:188) for bristles of the Cypridini-
formes is used herein for Deeveya (Fig. 6e):
“The one situated anteriorly = “‘the a-bris-
tle’; the one placed distally-medially and
somewhat anteriorly = “‘the b-bristle’’; the
posterior-distal one = “‘the c-bristle.”” The
four bristles on the original 8th joint: The
anterior of the two simple sensory filaments
which are situated laterally, close to each
other, is called “‘the d-bristle,”’ the posterior
VOLUME 100, NUMBER 3
one of them is called “‘the e-bristle”’; of the
two distal-medial bristles, the anterior one
is called “‘the f-bristle,”’ the posterior one =
“the g-bristle.”” On Deeveya the e-bristle is
longer and stouter than other bristles of the
8th joint and is generally named the “‘prin-
cipal bristle’ in the Halocyprididae. The
terminology used for Deeveya herein differs
from the terminology proposed by Skogs-
berg (1920:583) for the bristles of the end
joints of the Ist antenna of members of the
genus Halocypris, but coincidentally the
e-bristle is the principal bristle in both ter-
minologies.
The terminology proposed by Skogsberg
(1920:575) for bristles of the endopodite of
the 2nd antenna of the Halocyprididae is
applied herein to Deeveya (Figs. le, f; 4d;
6f, g): a- and b-bristles refer to the 2 dorsal
bristles of the Ist joint; f-bristle (lateral) and
g-bristle (medial) refer to the 2 long distal
bristles of the 2nd joint; and h-, -1, and
j-bristles refer to the 3 long terminal bristles
of the 3rd joint. The c-, d-, and e-bristles of
the 2nd joint, which are not present on all
species of the Halocyprididae, and are
mostly found only on males, were not pres-
ent on the juveniles of known species of
Deeveya, but a small proximal peg on the
2nd joint of the A-1 females of D. spiralis
(Fig. 6g) and D. bransoni (Fig. 1f) may rep-
resent one of those bristles. An additional
short unlettered bristle is present on the 3rd
joint of D. spiralis (Fig. 6g).
Deeveya bransoni, new species
Figs. 1-5
Etymology.—The species is named for
Mr. Richard Branson of Virgin Ltd, one of
the main sponsors of the field work in which
the junior author participated.
Material.—Bahama Islands, The Bluff,
South Andros Island (map reference: Ba-
hamas 1: 25000 OS. series): Evelyn Green’s
Blue Hole (type locality) (grid reference TB
410648); 4 Jul 1986; collected from just be-
yond limit of daylight at depth of 22 m in
611
saline water; holotype, USNM 193301,
A-1 female on slide and in alcohol (Palmer
specimen SA/3/86).—Stargate Blue Hole
(grid reference TB 403681); 3 Jul 1986; col-
lected in saline water at depth of 30 m, 80
m from cave entrance; paratype, USNM
193302, juvenile female (A-2) on slide and
in alcohol (Palmer specimen SA/2/86).
Distribution. —Known only from Evelyn
Green’s Blue Hole and Stargate Blue Hole,
South Andros, Bahamas.
Habitat.—Specimens from both caves
were from clear water with salinities in ex-
cess of 18%o and a stable temperature of
25°C. They were collected just below the
mixing zone between brackish and saline
waters. The mixing zones in both caves have
an associated growth of colonial bacterial
plate, which may be providing a reducing
environment and provides a primary food
source (Cunliffe 1985). The two caves lie on
a major fracture line which has its southern
end extending offshore, and its northern end
over | kilometer onshore. The fracture par-
allels the tongue of the Ocean, a 2000 m
deep oceanic trough to the east of Andros.
Current flow has been monitored along the
line of the fracture (Smart, Whittaker, and
Palmer, in prep.), and the fracture can be
regarded as a subterranean tidal creek, which
partially drains the fresh water lenses of
South Andros. In Stargate Cave the A-2 fe-
male was free-swimming in clear water and
appeared to be from a good population.
Description of A-1 female (Figs. 1—3).—
Carapace oval in lateral view except for
straight dorsal margin and slightly concave
anterior margin (Fig. la). Right valve with
small tubercle on dorsal margin near pos-
terior end.
Ornamentation (Fig. 1c): Carapace when
viewed with transmitted light appearing re-
ticulate (reticulations appearing bright); mi-
nute rounded processes present on outer
surface of walls of reticulations (processes
generally appearing darker than walls and
may extend above surface of valve).
Bristles (Fig. la): Valve margins with few
612 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
ui;
2 ef>2 © SS <2 er Dae f
[IS ASCs 5 seitineals ANG
GS : = ( Sot
Geeta)!
ee
d * Se i} /
ai ee
he SM |
" ee eee |
| Sk Tai ae ee
a Ni ne 2 Vay
SS \ Mie C of ~~
Fig. 1. Deeveya bransoni, A-\ female, holotype: a, Complete specimen from left side (dashed oval represents
location of central adductor muscle attachment scars), length 1.68 mm; b, Detail of adductor muscle attachments
(striate) and some reticulations as viewed through shell; c, Reticulations and bosses (small filled-in circles) viewed
with transmitted light (stippled areas appear dark); d, Anterior of body showing divided Bellonci organ, left
hepatic organ (upper dashed ovoid), left 1st antenna, and upper lip; e, Endopodite of left 2nd antenna, medial
view; f, Fused joints 2 and 3 of left endopodite, lateral view; g, Left 7th limb, lateral view; h, Anterior of body
from anterior showing upper lip (arcs at top represent sockets of lst antennae); i, Anterior view of lower lip.
VOLUME 100, NUMBER 3
widely spaced bristles. Setal bristle at tip of
posterodorsal tubercle of right valve just
posterior to glandular opening.
Glands: Glandular opening on tip of pos-
terodorsal tubercle of right valve.
Central adductor muscle attachments (Fig.
1b): Indistinct attachment scars forming
cluster near valve midlength.
Shell size: USNM 193301, length 1.68
mm, height 1.27 mm.
First antenna (Fig. 1d): Elongate with 8
joints. 1st joint with distal lateral spines be-
coming longer near ventral margin; distal
end of Ist joint overlapping proximal end
of 2nd joint, especially near ventral margin.
2nd joint with distal medial spines and dor-
sal midbristle with short marginal spines;
distal end of 2nd joint overlapping proximal
end of 3rd joint. 3rd joint elongate, with
spinous ventral bristle distal to midlength,
few minute spines along dorsal margin, and
longer medial spines near ventral margin.
4th joint short with small, slender, dorsal,
terminal bristle reaching past 6th joint. 5th
joint shorter than 4th, with long, terminal,
filament-like ventral bristle bearing few,
short, widely spaced, spines. 6th joint slight-
ly shorter than 5th, bare. 7th joint about
same length as 4th joint, with | short, distal,
lateral a-bristle, and 2 long b- and c-bristles
on small terminal pedistal (both bristles
longer than bristle of 5th joint and with
widely separated marginal spines and ter-
minal papilla; medial b-bristle filament-like;
lateral c-bristle with well-defined proximal
rings and about '4 longer than b-bristle). 8th
joint small, with very long principal e-bris-
tle and 3 shorter filament-like d-, f-, and
g-bristles (all bristles with widely spaced
marginal spines and terminal papilla; prin-
cipal bristle about same length as c-bristle
of 7th joint; g-bristle slightly longer than
b-bristle of 7th joint, and longer than d- and
f-bristles).
Second antenna (Fig. le, f): Protopodite
with lateral spines forming cluster distal to
middle. Endopodite 3-jointed. Ist joint with
2 distal dorsal a- and b-bristles on small
613
protuberance (proximal a-bristle about half
length of distal b-bristle); 2nd joint forming
right angle with Ist joint, with filament-like
f-bristle slightly longer than half length of
g-bristle (with widely separated marginal
spines and terminal papilla) and medial
g-bristle about twice length of protopodite,
stouter than f-bristle and with fairly strong
rings in proximal half (both f- and g-bristles
longer than bristle of 5th joint and with
widely separated marginal spines and ter-
minal papilla); minute proximal lateral peg
present near 3rd joint (Fig. 1f). 3rd joint
fused to second, with filament-like h-, i-,
and j-bristles, each less than half length of
g-bristle (with widely spaced marginal spines
and terminal papilla). Exopodite 9-jointed:
lst joint weakly divided into long proximal
and short distal parts; proximal part with
minute faint spines near dorsal (concave)
margin; distal part with short ringed bristle
reaching middle of 5th joint; 2nd joint with
long bristle with small spines along middle
part and distal natatory hairs; joints 3-8
each with long bristle with natatory hairs;
9th joint with 4 bristles (2 short and 1 me-
dium length, all with distal ventral spines
and no natatory hairs; | long with distal
natatory hairs and ventral spines).
Mandible (Fig. 2): Coxale endite with teeth
forming 3 rows (proximal (=dorsal), mid-
dle, distal (=ventral)) (Fig. 2b—d, f). Proxi-
mal row consisting of 4 broad teeth (Fig. 2b,
f); densely packed spines between teeth and
at each end; medial and lateral spines and
hairs proximal to teeth; single bifurcate
pointed or rounded tooth present between
proximal and middle rows (about midway
between posterior tooth of proximal row and
anterior tooth of middle row), and adjacent
to 2 stout spinous pointed bristles; 2 spinous
bristles (posterior stouter) with bases just
proximal and dorsal to bases of teeth form-
ing middle row. Middle row with 5 teeth
(posterior tooth longer than others) (Fig. 2c).
Distal row forming ventral edge of endite
with 6—7 teeth (middle tooth larger than
others) (Fig. 2d). Basale with 3 proximal
614 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
i
Fig. 2. Deeveya bransoni, A-| female, holotype, mandible: a, Proximal part of basale and endopodite of left
limb, lateral view; b, Coxale endite of left limb, anterior view; c, d, Detail from b showing middle and distal
teeth; e, Basale of left limb, lateral view; f, Coxale endite of right limb, posterior view; g, 3 terminal bristles of
3rd endopodial joint of left limb, lateral view.
VOLUME 100, NUMBER 3
bristles (2 stout and plumose, | slender with
short marginal hairs) (Fig. 2a, e). Basale en-
dite (Fig. 2e): anterior margin with single
ringed bristle; posterior margin with prox-
imal hairs, | short proximal bristle and 1
short distal tube-formed bristle; lateral side
with few long hairs, 6 slender distal bristles
(2 longer than others and entwined), and 1
short, stout, tooth just proximal to distal
edge of endite; medial side with few long
hairs; ventral edge with 6 terminal cusps (5
anterior serrate proximally, 1 posterior cusp
bare, smaller than others and separated from
them by space). Endopodite 3-jointed (Fig.
2a): 1st joint with 1 spinous, terminal, dor-
sal bristle, 1 spinous, distal, ventral bristle,
and 4 spinous, distal, medial bristles; 2nd
joint with 3 spinous, terminal, dorsal bris-
tles, and | spinous, terminal, ventral bristle;
3rd joint hirsute medially and along dorsal
margin, with 4 medial bristles forming row,
and 3 stout, spinous, terminal bristles (ven-
tral of these with long spines at midlength
and shorter thinner spines distally; middle
bristle longest, with smooth sharp tip, closely
spaced posterior spines forming medial and
lateral row and more widely spaced anterior
spines) (Fig. 2g).
Maxilla (Fig. 3a): 3 well developed en-
dites (bristles of endites not shown in illus-
tration): endite I with 2 proximal bristles
with long proximal hairs and about 8 ter-
minal bristles; endite IJ with 2 proximal
bristles with short marginal spines and about
7 terminal bristles; endite III with 1 prox-
imal bristle with short marginal spines and
about 5 terminal bristles. Coxale with 1
stout, hirsute, terminal, dorsal bristle. Ba-
sale with 1 slender ventral bristle with short
marginal spines and | slender terminal bris-
tle at midwidth. Endopodite: Ist joint with
4—5 spinous bristles on or near anterior mar-
gin, 5—6 bristles at distal posterior corner,
and sparse surface hairs; 2nd joint with 2
stout claws, 5 slender bristles, and long hairs
on anterior surface.
Fifth limb (Fig. 3b): Epipodite with hir-
sute bristles forming 3 groups: dorsal group
615
with | short and 4 long bristles; middle group
with 6 long bristles; ventral group with 4
long and | short bristles. Protopodite and
endopodite with 27 bristles including 2 pec-
tinate claw-like bristles at ventral corner.
Exopodite 3-jointed: lst joint with total of
12 bristles: 2 distal dorsal bristles (longest
bare, other with long marginal hairs), 2 lat-
eral bristles (distal of these plumose), 1 me-
dial bristle near middle, 3 proximal ventral
bristles, and 4 bristles on distal ventral cor-
ner. 2nd joint elongate with 4 bristles (1
dorsal, 3 ventral). 3rd joint short with 2 long
claw-like bristles and 2 slender ringed bris-
tles (smallest bristle medial).
Sixth limb (Fig. 3c): Epipodite with hir-
sute bristles forming 3 groups: dorsal group
with | short and 6 long bristles; middle group
with 6 long bristles; ventral group with 5
long bristles. Protopodite with 2 joints:
proximal joint with 4 bristles on or near
ventral margin (2 plumose, 2 with short
marginal spines); distal joint with 4 bristles
on or near ventral margin (2 plumose, 2
with short marginal spines). Exopodite
4-jointed: Ist joint with plumose lateral
bristle, S—6 plumose bristles on or near ven-
tral margin, and process with 4 bristles (1
long bristle bare, others plumose) on distal
dorsal corner; 2nd joint with 4 bristles (bare
or with short marginal spines) on or near
ventral margin; 3rd joint separated from 2nd
joint by suture, elongate with 3 bristles (2
ventral, 1 dorsal, either bare or with short
marginal spines); 4th joint short, with 2 stout
claw-like pectinate bristles and 2 slender
bristles (1 long, ventral, 1 short, medial).
Seventh limb (Fig. 1g): Limb unjointed,
with 3 terminal bristles (1 long, 2 shorter)
with indistinct marginal hairs.
Furca (Fig. 3d): Each lamella with total
of 7 claws followed by unpaired dorsal bris-
tle; claws decreasing in width posteriorly
along lamella; claw 4 slightly shorter than
claw 5, and claw 5 about same length as
claw 6; claws 1—4 with spines along poste-
rior margin; claw 5 with longer spines along
anterior margin than along posterior mar-
616 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig. 3. Deeveya bransoni, A-1 female, holotype: a, Maxilla, endite bristles not shown; b, Left 5th limb, lateral
view; c, Left 6th limb, lateral view; d, Left lamella of furca and apron; e, Ventral view of posterior end of hinged
valves showing attached protistans.
VOLUME 100, NUMBER 3
gin; claw 6 with spines along anterior mar-
gin, claw 7 with few spines along anterior
margin; stout triangular protuberance on
each lamella following last claw, and small
process laterally between claws | and 2. Un-
paired dorsal bristle with marginal spines.
Bellonci organ (Fig. 1d): Well developed,
bifurcate distally, with branches tapering to
pointed tip.
Lips (Fig. 1d, h, 1): Anterior face of upper
lip with 14 small tooth-like processes form-
ing upper horizontal row, and 6 larger pro-
cesses forming lower horizontal row (Fig.
lh); crescent-like low process at each end
of upper row (Fig. Id, h); posteroventral
edge with slight concavity at midwidth
(when viewed anteriorly (Fig. lh) or pos-
teriorly) having stout spine at each end of
concavity (Fig. 1h); hair-like spines forming
rows near concavity (Fig. 1h). Lower lip with
triangular process at each side of mouth (Fig.
11).
Posterior of body: Evenly rounded, un-
segmented.
Apron (Fig. 3d): Curving shield anterior
to both anus and proximal leading edge of
furca.
Gut: Narrow in vicinity of mouth (Fig.
1d), broadening dorsal to central adductor
muscle, then becoming narrower in short
segment proximal to a second broad area
(not as broad as anterior broad area), then
narrowing at anus. Anterior and posterior
broad areas containing brown unrecogniz-
able organic particles.
Epizoa (Fig. 3e): Holotype with attached
protistans on posterodorsal edges of valves.
Description of A-2 female (Figs. 4, 5).—
Carapace differs from that of A-1 female in
having slightly convex dorsal margin (Fig.
4a).
Shell size: USNM 193302, length 1.21
mm, height 0.94 mm.
First antenna (Fig. 1b, c): Similar to that
of A-1 female.
Second antenna (Fig. 4d): Longest of 4
bristles of 9th exopodial joint without na-
tatory hairs or spines. Limb otherwise sim-
ilar to that of A-1 female.
617
Mandible (Fig. 5a—g): Coxale similar to
that of A-1 female (Fig. 5b, d, g). Basale
with 4 proximal bristles (3 stout and plu-
mose, | slender, bare) (Fig. 5e). Basale en-
dite (Fig. 5b—e) similar to that of adult fe-
male except entwined bristles cross in 2 or
3 places rather than 4 or 5 (Fig. Se). En-
dopodite 3-jointed (Fig. 5a): lst joint with
1 spinous terminal dorsal bristle, 1 spinous
distal ventral bristle, and 2 spinous distal
medial bristles; 2nd joint with 3 terminal
dorsal bristles, and | terminal ventral bris-
tle; 3rd joint hirsute medially and along dor-
sal margin, with 4 spinous medial bristles
forming row, and 3 stout spinous terminal
bristles similar to those of A-1 female (Fig.
Sa, f).
Maxilla: 3 well developed endites: endite
I with 2 proximal bristles (1 with long prox-
imal hairs) and 9 terminal bristles; endite
II with 2 proximal bristles and about 7 ter-
minal bristles; endite III with | proximal
bristle and about 6 terminal bristles. Coxale
with | stout, hirsute, terminal, dorsal bris-
tle. Basale with 1 slender ventral bristle and
1 shorter, slender terminal bristle at mid-
width. Endopodite: Ist joint with 3 bristles
on or near anterior margin, 3—4 bristles at
distal posterior corner, and sparse surface
hairs; 2nd joint with 2 stout claws, 4 slender
bristles, and long hairs on anterior surface.
Fifth limb (Fig. 4e, f): Epipodite similar
to that of A-1 female. Protopodite and en-
dopodite with 24 bristles including 2 claw-
like bristles and 1 small spinous medial bris-
tle (Fig. 4f). Exopodite 3-jointed: Ist joint
with total of 10 bristles: 2 distal dorsal bris-
tles longest bare, other with long marginal
hairs); 2 lateral bristles (distal of these plu-
mose), | medial bristle near middle, 2 prox-
imal ventral bristles, and 3 bristles on distal
ventral corner. 2nd joint elongate with 3
bristles (1 dorsal, 2 ventral). 3rd joint short
with 2 long claw-like bristles and 2 slender
ringed bristles (smallest bristle medial).
Sixth limb (Fig. 5h): Epipodite similar to
that of A-1 female. Protopodite with 2 joints:
proximal joint with 3 bristles on or near
ventral margin (1 plumose, 2 with short
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
618
Ms J
VG
oe
VOLUME 100, NUMBER 3
marginal spines); distal joint with 3 bristles
on or near ventral margin (1 plumose, 2
with short marginal spines). Exopodite
4-jointed: Ist joint with plumose lateral
bristle, 4 plumose bristles on or near ventral
margin, and process with 4 bristles (1 long
bristle bare, others plumose); 2nd joint with
2 ventral bristles; 3rd joint separated from
4th by weakly defined suture, elongate with
2 bristles (1 ventral, 1 dorsal); 4th joint short,
with 2 claw-like pectinate bristles and 2
slender bristles (shortest of these medial).
Seventh limb (Fig. 41): Limb unjointed,
with 3 bare bristles (1 long, 2 short equi-
length).
Furca (Fig. 4g): Each lamella with total
of 6 claws followed by unpaired dorsal bris-
tle; claws decreasing in width posteriorly
along lamella, claw 4 slightly shorter than
claw 5 and claw 6 slightly shorter than claw
5; claws 1-4 with spines along posterior
margin; claw 5 with few distal hair-like
spines along anterior margin; stout trian-
gular protuberance on each lamella follow-
ing last claw, and small process laterally be-
tween claws 1 and 2.
Bellonci organ (Fig. 4b): Well developed,
bifurcate distally with branches tapering to
pointed tip. Marginal hairs visible at high
magnification (x 100 objective).
Lips (Figs. 4b, 5i, j), posterior of body
(Fig. 4g), apron (Fig. 4g), gut: Similar to
those of A-1 female.
Comparisons. —The carapace of D. bran-
soni has bosses on walls of reticulations larg-
er than those of D. spiralis (compare Figs.
lc and 6b). The A-1 female bransoni has a
much smaller shell than that of the A-1 fe-
male of spiralis: 1.68 mm compared to 2.67-
2.87 mm. The end joints of the exopodites
of the 5th and 6th limbs of the A-1 female
—
619
bransoni bear four bristles whereas those of
spiralis bear five. The ventral bristle on the
end joint of the mandible of the A-1 female
bransoni bears near midlength long stout
marginal spines (Fig. 2g); these are less well
developed and closer to the tip on spiralis
(Fig. 61).
Ontogenetic development.—The collec-
tion comprised an A-1| and A-2 female. Their
Carapaces are similar in shape and orna-
mentation except the dorsal margin of the
Shell is slightly convex on the A-2 instar and
straight on the A-1. The Ist and 2nd anten-
nae have the same number of bristles in
both stages. The coxale and basale of the
mandible are similarly developed, except the
A-2 instar has four proximal bristles on the
basale and only three on the A-1 (possibly
a bristle broke off during dissection of the
A-1 female), and the two entwined bristles
of the basale cross each other two to three
times on the A-2 instar and four to five
times on the A-1. The endopodites of the
mandible and maxilla and the exopodites
of the 5th and 6th limbs of the A-2 instar
have in total a smaller number of bristles,
but fewer bristles are added on distal joints
(Table 1). The 7th imb and Bellonci organ
are similar for both stages. The furca of the
A-2 instar has six claws, compared to seven
for the A-1.
Deeveya spiralis
Kornicker and Iliffe, 1985
Fig. 6
Material. —Holotype, USNM 193117;
paratype, USNM 193118.
Distribution. —The Hole, a marine cave
on Providenciales Island, Caicos Islands,
Turks and Caicos Islands.
Fig. 4. Deeveya bransoni, A-2 female, paratype: a, Complete specimen from right side, length 1.21 mm; b,
Anterior of body showing divided Bellonci organ, right Ist antenna, and upper lip; c, Medial view of joints 1-
4 of right 1st antenna showing spinosity; d, Endopodite of left 2nd antenna, lateral view; e, Right 5th limb,
lateral view; f, Claw-like bristles and short stout medial bristle of endite of left Sth limb, medial view; g, Right
lamella of furca (arrow indicates anus); h, Ventral or dorsal view of Bellonci organ under cover slip; 1, Right
7th limb, lateral view.
620 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
—
aS an ga a Cote |
2900900
=
Fig. 5. Deeveya bransoni, A-2 female, paratype: a-g, Mandible: a, Posterior view of proximal parts of coxale
and basale, and lateral view of endopodite of right limb; b, Medial view of coxale endite of right limb, not under
VOLUME 100, NUMBER 3
Remarks. — Because both the holotype and
paratype of D. spiralis had unextruded eggs
in their ovaries, Kornicker and Iliffe (1985)
interpreted the specimens to be adult fe-
males; however, they could be A-1 females,
and they are so interpreted herein. A reason
for this is that a species from a cave in Ber-
muda referred to Spelaeoecia by Angel and
Iliffe (1987) has almost identical append-
ages to those of Deeveya, and the adult fe-
male of that species has eight claws on the
furca, one more than on the known speci-
mens of D. spiralis. It seems probable that
the adult Deeveya will also have eight furcal
claws, and the A-1 instar only seven.
Supplementary description of A-1 female
(Fig. 6).—Kornicker and Iliffe (1985:fig. 2)
illustrated a fragment of shell having convex
pillow structures forming surfaces of areas
within reticulate walls; further examination
suggests that convex structures probably oc-
cur only on a decalcified shell, and that the
outer surface of calcified specimens are es-
sentially flat (Fig. 6c). Reticulations visible
in transmitted light are structures within the
shell wall and not surface structures. Minute
bosses on the shell surface occur lateral to
the walls forming reticulations, generally
where walls intersect (Fig. 6b, c). Small
round structures appearing lighter (when
viewed with transmitted light) than sur-
rounding area occur on the surface of some
areas bounded by reticulate walls, but
whether these are raised or depressed could
not be ascertained (Fig. 6b), but they are
shown raised in Fig. 6c.
First antenna (Fig. 6d, e): Sth joint with
filament-like bristle with widely spaced mi-
nute marginal spines and terminal papilla.
6th joint bare. 7th joint with short ringed
spinous lateral a-bristle near distal dorsal
corner and 2 bristles on terminal ventral
pedistal (medial b-bristle about 7% length of
—
621
Table 1.—Number of bristles on mandible, maxilla,
and 5th and 6th limbs of A-2 and A-1 females of Deev-
eya bransoni.
Growth stages
A-2 A-1
Mandible, endopodite
Ist joint 4 6
2nd joint 4 4
3rd joint 7 7
Maxilla, endopodite
Ist joint 6-7 9-11
2nd joint 6 7
5th limb, exopodite
Ist joint 10 12
2nd joint 3 4
3rd joint 4 4
6th limb, exopodite
Ist joint 9 10-11
2nd joint 2 4
3rd joint 2 3
4th joint 4 4
lateral, filament-like, with minute widely
spaced marginal spines and terminal pa-
pilla; lateral c-bristle about 1'2 times length
of stem, ringed in proximal *%3 and with
widely spaced marginal spines and terminal
papilla (spines stouter than those on bristle
of 5th joint)). 8th joint with 4 terminal bris-
tles: d-bristle filament-like, short, with small
widely spaced marginal spines and terminal
papilla (tip of e-bristle of illustrated right
limb aberrant; dashed line indicates length
of e-bristle of left limb); e-bristle (principal
bristle) about twice length of stem and dis-
tinctly ringed in proximal half, with small
widely spaced marginal spines and terminal
papilla; f-bristle about 4 length of principal
bristle; g-bristle more than half length of
principal bristle, both f- and g-bristles fila-
ment-like, with small widely spaced mar-
ginal spines and small terminal papilla.
cover slip; c, Medial view of distal teeth of basale of right limb, not under cover slip; d, Anterior view of coxale
endite of right limb, under cover slip; e, Lateral view of basale of right limb; f, Ventral terminal bristle of 3rd
endopodial joint of right limb, lateral view; g, Posterior view of coxale endite of left limb, under cover slip; h,
Right 6th limb, lateral view; i, Outline of upper lip from right side, anterior to right; j}, Lower lip, ventral view.
622 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
ho ae ee Ne aa ee Nee
on
Fig. 6. Deeveya spiralis, A-1 female, paratype: a, Complete specimen from left side, length 2.67 mm; b,
Lateral view using transmitted light of part of right valve just ventral to central adductor muscle attachments
showing shell structure (stippled areas appear darker when viewed through microscope); c, Sketch of theoretical
cross-section of shell shown in b (outside of shell at top). Holotype: d, Right 1st antenna, lateral view; e, Detail
of tip of limb in d; f, Protopodite and endopodite of right 2nd antenna, lateral view; g, Detail of tip of endopodite
in f; h, Distal part of basale of right mandible, medial view; 1, Ventral bristle of end joint endopodite of right
mandible, medial view; j, Left lamella of furca showing gland between claws | and 2, lateral view.
VOLUME 100, NUMBER 3
Second antenna: Endopodite (Fig. 6f, g):
Ist joint: dorsal margin with stout promi-
nence bearing spinous a-bristle less than half
length of spinous b-bristle, both bristles di-
rected anteriorly. 2nd joint forming right
angle with Ist joint, with 2 distal bristles:
f-bristle filament-like, slightly more than half
length of g-bristle, with widely spaced mi-
nute spines (smaller than marginal spines of
g-bristle), and terminal papilla; g-bristle
more than twice length of protopodite, with
distinct rings in proximal half, with widely
spaced minute marginal spines, and ter-
minal papilla; minute, unringed, pointed peg
present proximally on dorsal margin of joint.
3rd joint with 3 similar filament-like h-, i-,
and j-bristles, each less than half length of
g-bristle, with widely spaced minute mar-
ginal spines (spines about same size as those
of f-bristle), and with terminal papilla; small,
unringed, medial, terminal bristle with mar-
ginal spines along distal quarter present near
dorsal corner of joint (length of bristle about
half width of joint).
Mandible: Distal end of basale with stout
lateral tooth (Fig. 6h); ventral (=posterior)
bristle of 3rd endopodial joint with long hairs
(Fig. 61). (Figure 10 in Kornicker and Iliffe
(1985:488) was incorrectly labelled and
should be as follows: “*b” is an illustration
of the coxa, and “‘c’’ the basis and endop-
odite.)
Maxilla: Hirsute dorsal bristle on coxale
(not on basale as stated by Kornicker and
Iliffe 1985:490).
Furca (Fig. 6j): Small lateral process be-
tween claws | and 2 of each lamella rep-
resenting glandular outlet (this process was
overlooked by Kornicker and Iliffe 1985:
491).
623
Acknowledgments
The field work was supported in part by
Virgin Diver’s World and Shell Explora-
tion. Our thanks to Dr. Thomas E. Bowman
for reviewing the manuscript.
Literature Cited
Angel, M. V. 1983. A review of the progress of re-
search on halocyprid and other oceanic plank-
tonic ostracodes 1972-1982. Pages 529-543 in
R. F. Maddocks, ed. Applications of Ostracoda.
Houston, Department of Geosciences, Univer-
sity of Houston. [Proceedings of the Eighth In-
ternational Symposium on Ostracoda July 26-
29, 1982.]
, and T. M. Iliffe. 1987. Spelaeoecia bermu-
densis, new genus, new species, a halocyprid os-
tracod from marine caves in Bermuda. —Jour-
nal of Crustacean Biology 7(3):541-553.
Cunliffe, S. 1985. The flora and fauna of Sagittarius,
an anchialine cave and lake in Grand Baha-
ma.—Cave Science 12(3):103-109.
Danielopol, D. L. 1972. Sur la Présence de Thau-
matocypris orghidani n. sp. (Ostracoda, My-
odocopida) dans une Grotto de Cuba. — Comptes
Rendus hebdomadaires des Séances de |’Aca-
démie des Sciences, Paris 247:1390-1393.
Hartmann, G. 1985. Danielopolina wilkensi n. sp.
(Halocyprida, Thaumatocyprididae), ein neuer
Ostracode aus einem marinen Lava-Tunnel auf
Lanzarote (Kanarische Inseln).— Mitteilung aus
dem Hamburgischen Zoologischer Museum und
Institute 82:255-261.
Kornicker, L. S., and T. M. Iliffe. 1985. Deeveyinae,
a new subfamily of Ostracoda (Halocyprididae)
from a marine cave on the Turks and Caicos
Islands.— Proceedings of the Biological Society
of Washington 98(2):476-493.
(LSK) Department of Invertebrate Zo-
ology, National Museum of Natural His-
tory, Smithsonian Institution, Washington,
D.C. 20560, USA. (RJP) Department of
Geography, University of Bristol BS8 1SS,
England.
PROC. BIOL. SOC. WASH.
100(3), 1987, pp. 624-628
TWO NEW TURTLES FROM SOUTHEAST ASIA
Carl H. Ernst and William P. McCord
Abstract.—Cuora chriskarannarum (Emydidae: Batagurinae), new species,
from Yunnan Province, China, and Platysternon megacephalum shiui (Platy-
sternidae), new subspecies, from northern Vietnam are described and compared
with other members of their genera.
The recent relaxation of regulations re-
garding scientific exploration and the col-
lection and export of reptiles from China
and Vietnam have introduced many un-
common and unusual animals into the pet
trade. Dr. William P. McCord, a turtle fan-
cier, obtained a number of turtles and rec-
ognized that some of these animals repre-
sented new taxa. Two of these are described
below.
Methods and Materials
Straight-line measurements of each spec-
imen were taken with dial calipers accurate
to 0.1 mm, of the greatest carapace length,
carapace width and depth at the level of the
seam between vertebrals 2 and 3, marginal
width (the difference between the carapacial
width and the width across the pleurals tak-
en between the points of juncture of the
marginals and pleurals at the level of the
seam between vertebrals 2 and 3), greatest
plastron length, greatest width and length of
both plastral lobes, greatest bridge length,
greatest width and length of vertebrals 1 and
2 and pleural 2, and the medial seam length
and greatest width of all plastral scutes.
Notes and drawings were made of head,
neck, limb, carapacial, plastral, and bridge
patterns. Colors were recorded from living
turtles and color transparencies. Shell pro-
portions are expressed as ratios of one mea-
surement to another. Sixteen ratios proved
useful (abbreviations used in the text are
given in parentheses): width/length of cer-
vical scute (W/L CS), width/length of first
vertebral (W/L Ist V), width/length of sec-
ond vertebral (W/L 2nd V), width/length of
second pleural (W/L 2nd PI), marginal
width/carapacial width (MW/CW), margin-
al width/carapacial length (MW/CL), cara-
pacial width/carapacial length (CW/CL),
carapacial depth/carapacial length (D/CL),
carapacial depth/carapacial width (D/CW),
plastral length/carapacial length (PL/CL),
bridge length/plastral length (B/PL), bridge
length/carapacial length (B/CL), length of
anterior plastral lobe/plastral length (APL/
PL), width of anterior plastral lobe/plastral
length (APW/PL), length of posterior plas-
tral lobe/plastral length (PPL/PL), and width
of posterior plasteral lobe/plastral length
(PPW/PL). The number of rows of large
scales at the lateral edge of the antibrachium
between the claw of digit V and the first
horizontal skin fold proximal to the elbow
(presented in text as foreleg scale rows) was
recorded.
Specimens from the following collections
were examined (abbreviations used in the
text are given in parentheses): William P.
McCord personal collection of living turtles
(WPM), National Museum of Natural His-
tory, Smithsonian Institution (USNM).
Cuora chriskarannarum, new species
Fig. 1
Holotype. —USNM 266162, adult male;
Ta Lau Shan, Yunnan Province, China
(23°30'N, 102°25’E); Oscar Shiu, Sep 1986.
Paratype. -USNM 266163, adult fe-
male; Chinsha (=Chin ° Ping), Yunnan
VOLUME 100, NUMBER 3
Province, China (22°46'N, 103°15’E); Oscar
Shiu, Sep 1986.
Diagnosis. — Flattened, green species of
Cuora with yellowish-green, black-bor-
dered postorbital stripe, oblique yellowish-
green stripe extending from upper jaw to
below tympanum to neck, medial carapacial
keel most pronounced on vertebrals 1-3,
vertebral 1 usually extending laterally to
seam separating marginals 1-2 or beyond,
broad black seam-following plastral pat-
tern, plastron with wide anal notch, and in-
teranal seam complete.
Description (from all specimens exam-
ined).—Carapace length to 160 mm (males
113, females 160), elliptical, flattened (D/
CL 0.29-0.42, x = 0.346; D/CW 0.40-0.60,
X = 0.498; CW/CL 0.67-0.73, x = 0.694);
widest at marginals 8, highest at posterior
of vertebral 2. Carapace sides straight, pos-
terior rim slightly serrated and with a very
small medial notch. All marginals flared,
those over bridge slightly downturned (MW
11.8-17.3 mm, x = 14.1; MW/CW 0.13-
0.18, x = 0.157; MW/CL 0.09-0.14, x =
0.109). Marginal 1 widest, marginals 3-6
smallest. Scute texture rough with growth
annuli. Cervical rectangular, longer than
wide (W/L CS 0.70-0.72, x = 0.713). Ver-
tebrals wider than long; vertebral 1 largest
and very flared anteriorly, reaching seam
separating marginals 1—2 or marginal 2 in
15 (88%) specimens; vertebral 5 posteriorly
flared. Low medial keel most pronounced
on vertebrals 3—5. Olive-brown, rim yellow,
seams outlined with dark brown or black,
keel black. Undersides of marginais yellow
with black, posteriorly-directed wedge at
each seam, some with narrow, radiating
black lines.
Plastron length to 159 mm (males 111,
females 159), longer than carapace in 7 (41%)
specimens (PL/CL 0.96-1.02, x = 0.995),
slightly upturned anteriorly, movable hinge
between pectoral and abdominal scutes.
Posterior lobe longer and wider than ante-
rior lobe (APL/PL 0.39-0.44, x = 0.414;
PPL/PL 0.55-0.60, x = 0.578; APW/PL
Fig. 1. Cuora chriskarannarum, new species.
0.45-0.50, xX = 0.474; PPW/PL 0.51-0.57,
xX = 0.536; APW/CW 0.62-0.70; x = 0.682:
PPW/CW 0.74-0.81, x = 0.769). Anterior
lobe truncated, posterior lobe with shallow
anal notch. Anal scutes laterally tapering
toward midline. Bridge moderate (B/CL
0.22-0.32, x = 0.273; B/PL 0.23-0.31, * =
0.274); O-1 axillaries; 1 inguinal. Average
plastral formula An. > Abd. > Pect. >
Gul. > Fem. > Hum.; 10 (59%) had this
formula, but 5 other formulae occurred; An.,
Abd., Pect. always longer than Gul., Fem..,
Hum. Interanal seam present and complete.
Yellow with wide black seam-following pat-
tern becoming more extensive with age
(size); bridge with black bar.
Head narrow; snout slightly projecting;
upper jaw slightly hooked. Olive (lighter
dorsally, darker laterally) with yellowish-
green postorbital stripe and second yellow-
ish-green stripe extending obliquely down-
ward from upper jaw to below tympanum;
626
both stripes with narrow black borders. Faint
thin black line may circle tympanum. Iris
green; unmarked yellow jaws and chin. Neck
olive dorsally and laterally, yellowish-green
ventrally, several faint, narrow yellow lat-
eral stripes present.
Digits webbed. Forelimbs with large scales
(10-15 rows, x = 14.0); outer surface olive,
inner surface and sockets cream to whitish-
green. Hindlimbs with smaller scales, col-
ored as forelimbs. Holotype with amputat-
ed right hindlimb. Tail olive dorsally with
two dark-bordered longitudinal stripes,
venter yellow with olive tip.
Smaller, flatter males having concave
plastra, longer thicker tails with anal vent
beyond carapacial rim, and pointed snouts;
larger, more vaulted females having flat
plastra, shorter tails with anal vent beneath
carapacial rim, and rounded snouts.
Other material.—WPM 1-15 (live; 2
males, 13 females); from localities of type
specimens.
Etymology. —The name chriskaranna-
rum is in honor of Dr. McCord’s daughters
Christine and Karen, and his wife Anne.
Remarks. —The new species is most
closely related to Cuora pani Song, 1984,
C. trifasciatus (Bell, 1825), and C. yunna-
nensis (Boulenger, 1906), and at least one
additional undescribed species. These taxa
form a distinct subgroup within Cuora char-
acterized by relatively flattened carapaces,
plastrons with wide anal notches, and com-
plete interanal seams. Cuora pani is from
Shaanxi Province, China. It has a lemon-
yellow head with a broad yellow longitu-
dinal stripe, a brown carapace with reddish
vertebrals and a keel extending over ver-
tebrals 1-5, a yellow plastron with a pattern
of black wedges that are often separated
along the seams, and a broad vertebral 1
that may extend to marginals 2. Cuora tri-
fasciatus ranges from Kwangsi and Kwang-
tung provinces and Hainan in China through
northern Vietnam and possibly to Burma.
Its head is olive dorsally, brown to black
laterally, and with a lateral longitudinal
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
stripe; an olive to brown tricarinate cara-
pace bears three longitudinal black stripes;
the black plastron has a yellow border; ver-
tebral 1 usually does not extend to the seam
separating marginals 1-2. Cuora yunnanen-
sis 18 known only from the highlands of
Yunnan Province, China. It has a brown
head with a narrow yellow stripe extending
from the nostril through the eye to the neck,
a unique mottled chin pattern, a brown car-
apace with dark seams and a keel extending
over all five vertebrals, a yellow to light
brown plastron with narrow dark seams, and
a vertebral 1 that does not extend to the
seam separating marginals 1-2.
Platysternon megacephalum shiui,
new subspecies
Fig. 2
Holotype. —USNM 266160, adult male;
vicinity of Langson, Langson Province,
Vietnam (26°50’N, 106°45’E); Oscar Shiu,
Aug 1986.
Paratype. —USNM 266161, adult male;
same collection data as holotype.
Diagnosis. —Head, shell, limbs, sockets,
ventral surface of tail heavily speckled with
yellow, orange or pink spots; cephalic shield
moderately developed, not entering orbit;
upper jaw strongly hooked; carapacial sur-
face smooth, posterior rim unserrated; no
postorbital stripe, dark plastral figure, or
small additional scales at medial juncture
of gular and humeral scutes.
Description.—(From all specimens ex-
amined.) Carapace length to 151 mm (males
151, females 107), flat (D/CL 0.26-0.31, x =
0.280; D/CW 0.38-0.47, x = 0.438; CW/
CL 0.60-0.68, x =.0.640), widest at mar-
ginals 9, highest at vertebral 1. Carapace
sides straight, or slightly indented at bridge;
anterior rim medially indented; posterior
rim smooth, but with small medial notch.
Anterior and posterior marginals flared, es-
pecially over hindlimbs, those at bridge
downturned (MW 3.6—7.9 mm, x = 5.53;
MW/CW 0.42-0.81, x = 0.65; MW/CL
VOLUME 100, NUMBER 3
0.025-0.045, x = 0.041); marginals 1 wid-
est, those over bridge smallest. Scute sur-
faces smooth. Cervical very small, wider
than long (absent in two specimens). Ver-
tebrals wider than long, 5th largest. Medial
keel low, blunt; most prominent on verte-
brals 3-5. Brown with mottled pattern of
small yellow, orange, or orangish-red spots
throughout, but most prominent on pleurals
and dorsal surface of marginals. Ventral sur-
face of marginals yellowish-brown or olive
with yellow, orange or pink speckles.
Plastral length to 112 mm (males 112,
females 94); PL/CL 0.75-0.80, x = 0.776.
Anterior lobe slightly longer than posterior
lobe, which wider (APL/PL 0.36-0.47, x =
0.433; PPL/PL 0.40-0.44, x = 0.423; APW/
PL 0.36-0.48, x = 0.443; PPW/PL 0.46-
0.52, x = 0.493; APW/CW 0.46-0.61, x =
0.537; PPW/CW 0.55-0.67, x = 0.598). An-
terior lobe truncated; posterior lobe with
anal notch. Femorals and anals laterally ta-
pering toward midline. Bridge narrow; B/CL
O.15-0.20; x = 0.173; B/PL 0.19-0.24, x =
0.223; 1-2 axillaries, 1—2 inguinals. Plas-
tron connected to carapace by ligaments at
bridge. Average plastral formula An. >
Hum. > Fem. > Pect. > Abd. > Gul.; 8
formulae present in 1 1 specimens examined
(3 with average formula); variation occur-
ring in Hum., Pect., Fem., and An.; Abd.
and Gul. consistently shortest. Yellowish-
brown or olive with many small yellow to
orangish-red spots throughout, and yellow
seams.
Head large, cannot be withdrawn into
shell. Snout pointed, projecting; upper jaw
strongly hooked. Dorsal cephalic shield only
moderately developed, not covering pos-
terior portion of orbit. Upper jaw sheath not
as well developed as in other subspecies.
Brown with numerous orange to red spots
on dorsal and lateral surfaces. Eyes bulging;
iris brown. Jaws, chin, throat brown with
yellow, orange or red mottling.
Digits webbed. Forelimb with large scales
(7-10 rows, x = 8). Thighs with tubercle-
like scales; skin of outer surface of limbs
627
Fig. 2. Platysternon megacephalum shiui, new sub-
species.
and dorsal surfaces of neck and tail brown;
limb sockets and ventral surface of neck yel-
lowish-brown. Limbs, sockets, underside of
tail with yellow, orange or red spots. Tail
long, covered dorsally with large annular
scales.
Males having concave plastra and anal
vent beyond carapacial marginals; females
flat plastra and vent beneath carapace.
Other material. — WPM 1-9 (live, 7 males,
2 females) from type locality.
Etymology. —The name shiui is in honor
of the collector Oscar Shiu.
Remarks. —Four other subspecies of
Platysternon megacephalum have been de-
scribed. Platysternon megacephalum mega-
cephalum Gray, 1831, which occurs in
southern China, has an unpatterned yellow
plastron, a slightly keeled carapace with
poorly developed growth annuli, and a
slightly serrated posterior rim, the margin-
als above the bridge flared, a well developed
628
cephalic shield that often covers the rear of
the orbit, a strongly hooked upper jaw, yel-
low mottling on the jaws, and a pattern of
radiating narrow lines on the dorsal surface
of the head. Platysternon megacephalum
peguense Gray, 1870 is found from western
Vietnam west to southern Burma. It has a
dark seam-following plastral pattern, the
carapacial keel pronounced and sometimes
indications of lateral keels, well developed
growth annuli, the posterior carapacial rim
serrated, unpatterned yellow jaws, a strong-
ly hooked upper jaw, and a black-bordered
postorbital stripe. Platysternon megaceph-
alum vogeli Wermuth, 1969 occurs in
northwestern Thailand. It is similar to P.
m. peguense in having a dark plastral figure,
but differs in having a short, narrow, less
hooked upper jaw, and a smooth, unserrat-
ed carapace. Platysternon megacephalum
tristernalis Schleich and Gruber, 1984 is
from Yunnan Province, China, and similar
to P. m. megacephalum except it has three
additional small scales at the medial junc-
tion of the gular and humeral scutes. It is
likely that further study will prove P. m.
vogeli and P. m. tristernalis invalid. Ernst
is currently studying the variation within
Platysternon.
Acknowledgments
George R. Zug critically commented on
the manuscript, and Evelyn M. Ernst typed
it.
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Literature Cited
Bell, T. 1825. A monograph of the tortoises having
a movable sternum, with remarks on their ar-
rangement and afhnities.— Zoological Journal,
London 2:299-310.
Boulenger, G. A. 1906. Descriptions of new reptiles
from Yunnan.— Annals and Magazine of Nat-
ural History (7) 17:567—568.
Gray, J. E. 1931. Characters of a new genus of fresh-
water tortoise from China. — Proceedings of the
Zoological Society of London 1831:106-108.
1870. Supplement to the catalogue of shield
reptiles in the collection of the British Museum.
Part I. Testudinata (tortoises), with figures of
the skulls of 36 genera. Trustees of the British
Museum, London, 120 pp.
Schleich, H.-H., and U. Gruber. 1984. Eine neue
Grosskopfschildkr6éte, Platysternon megaceph-
alum tristernalis nov. ssp., aus Yunnan, China
(Reptilia, Testudines, Platysternidae).—Spix-
iana 7:67-73.
Song, M.-T. 1984. A new species of the turtle genus
Cuora (Testudoformes: Testudinidae).—Acta
Zootaxonomica Sinica 9:330-332.
Wermuth, H. 1969. Eine neue Groskopfschildkrote,
Platysternon megacephalum vogeli, n. spp.—Die
Aquarien-und Terrarien-Zeitschrift (Datz) 22:
372-374.
(CHE) Department of Biology, George
Mason University, Fairfax, Virginia 22030;
(WPM) East Fishkill Animal Hospital,
Route 82, RD #8, Box 268, Hopewell Junc-
tion, New York 12533.
PROC. BIOL. SOC. WASH.
100(3), 1987, pp. 629-639
THE CAPRELLIDS (AMPHIPODA: CAPRELLIDA)
OF BERMUDA: A SURVEY OF SPECIMENS
COLLECTED FROM 1876-1987, INCLUDING
CAVE INHABITANTS, AND THE DESCRIPTION
OF DEUTELLA ASPIDUCHA,
NEW SPECIES
Michael F. Gable and Eric A. Lazo-Wasem
Abstract. — An examination of all available specimens of Bermuda caprellids,
including recently collected cave caprellids, permits a contemporary survey of
the group for Bermuda waters. A new species, Deutella aspiducha, is described.
Its transitional characters suggest the reabsorption of Luconacia incerta into
the genus Deutella. Four other Bermuda caprellids (Caprella danilevskii, C.
equilibra, Hemiaegina minuta, and Fallotritella biscaynensis) are reviewed, and
keys are provided for all six species. Zoogeographical and evolutionary theories
using cave and shallow-water specimens are not possible because of the paucity
of material from the latter habitat.
Kunkel’s 1910 monograph on the am-
phipods of Bermuda included four species
of caprellids. In 1968 McCain mentioned
two more species for Bermuda and sum-
marized the small amount of information
known for the six Bermuda species on hab-
itat preferences, morphological variation,
etc. Several recent factors predicate an up-
dated treatment of the Bermuda caprellids:
the discovery at the Yale Peabody Museum
(YPM) of many of Kunkel’s specimens
(Lazo-Wasem and Gable 1987), a collection
of Bermuda amphipods made in 1985, and
the examination of all Bermuda caprellids
housed at the United States National Mu-
seum (USNM), including specimens col-
lected by G. Brown Goode in 1876-1877,
and the cave specimens collected this de-
cade by T. M. Iliffe of the Bermuda Biolog-
ical Station. In addition, the discovery of a
possibly endemic species, Deutella aspidu-
cha, adds to the desirability of a reconsi-
deration of the Bermuda caprellids.
Systematics
Full synonymies for the Bermuda ca-
prellids are in McCain (1968); taxonomic
changes since his publication are noted in
this paper. Our synonymies apply only to
publications that mention a species as it oc-
curs in Bermuda. The classification schemes
of McCain (1970), Bowman and Abele
(1982), and Schram (1986) are used
throughout this paper. Legend: major body
parts are marked by abbreviations begin-
ning with uppercase letters. Abbreviations
are as follows: A, antenna; Ab, abdomen;
C, cephalon; Gn, gnathopod; Md, mandi-
ble; Mx, maxilla; Mxpd, maxilliped; P, pe-
reopod; Pn, pereonite.
Family Aeginellidae Vassilenko, 1968
Deutella Mayer, 1890, emend.
Diagnosis. —Flagellum of antenna 2 bi-
articulate (rarely triarticulate?), swimming
setae absent; mandibular palp 3-segmented,
630
setal formula for terminal article 1-x-1, 1,
or 2, knob on terminal article present or
absent, molar present; outer lobe of max-
illiped larger than inner lobe; gills on pereo-
nites 3 and 4; pereopods 3 and 4, 2-seg-
mented, pereopod 5, 6-segmented; abdomen
of male with pair of appendages and pair of
setose lobes, female with or without a pair
of lobes.
Deutella aspiducha, new species
Figs. 1-3
Diagnosis.—Cephalon and pereonite 2
each with 1| dorsal, anteriorly directed spine.
Males with anterior spatulate processes on
pereonite 2; pereonites 3 and 4 with out-
wardly projecting, crenulate side plates. In-
sertion of pereopod 5 in both sexes, 4 from
posterior edge of pereonite 5.
Etymology of specific modifier. — From the
Greek aspiduchos meaning “‘shield-bearer,”’
in reference to the distinct and crenulate
side plates on pereonites 3 and 4 of the male.
Description.—Male holotype, 4.8 mm.
Cephalon with | dorsal, anteriorly directed
spine, followed posteriorly by 2 smooth,
rounded dorsal humps; posterolateral
margins with small, anteriorly projecting
triangular processes. Pereonite 2 with 1
middorsal, anteriorly directed spine; an-
terolateral margins with distinct, large, an-
teriorly projecting spatulate processes. Per-
eonites 3-7 dorsally smooth, except
pereonites 3 and 4 with middorsal hump;
side plates of pereonites 3 and 4 projecting
ventrolaterally with margins distinctly cren-
ulate. Eyes present, round.
Antenna 1, 46% of body length. Pedun-
cular segment | broader than segment 2;
peduncular ratios 1:1.3:0.7. Peduncular seg-
ments heavily circumsetose with fine setae.
Flagellum 0.6 length of peduncle, of 6 or 7
articles, first article conjointed. Antenna 2,
16% of body length, swimming setae absent,
flagellum 2-articulate.
Mandible with 3-segmented palp, setal
formula for terminal article 1-x-1, terminal
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
article with knob, penultimate segment with
single distal seta. Left mandible with
6-toothed incisor, serrate lacinia, spine row
of 3 broad and short spines, molar strong.
Right mandible identical. Palp of maxilla 1
with 3 apical spines, 1 medial spine and 1
facial seta; outer lobe with 4 apical spines
and | shorter subapical spine. Outer lobe of
maxilla 2 with 4 apical setae; inner lobe with
3. Outer lobe of maxilliped narrow, long,
reaching '4 length of palp segment 2, with
2 apical setae, 2 medial setae, and | facial
seta; inner lobe oval, small, % length of
outer, with 2 apical setae; terminal article
of palp with 1 distal seta, penultimate article
with no triangular projection.
Article 6 of gnathopod | triangular, with
grasping spine, palm with few setae, a small
distal notch, and a facial spine row on inner
surface; dactyl extending length of palm,
with straight inner margin. Gnathopod 2
strongly inserted in expanded anteroventral
corner of pereonite 2; article 2 with small,
proximal, medially projecting triangular
process; article 6, proximal end of palm de-
marcated by a grasping spine, palm with a
strong poison tooth proximal to a deep notch
followed distally by a small notch, and more
distally armed with small teeth; article 7,
curved and smooth.
Pereopods 3 and 4 inserted at base of gills,
2-segmented with tiny, distal, setose seg-
ments. Pereopod 5 of 6 segments, much
smaller than pereopods 6 and 7, inserted 4
from posterior margin of pereonite 5, prop-
odus without palm, terminal article small
and setose. Pereopods 6 and 7 normal, ar-
ticles 6 with grasping spines.
Abdomen with medium-sized penes, a
pair of appendages with recurved spines,
and a pair of setose lobes.
Female allotype, 4.2 mm. Ovigerous. All
features same as those for male except as
noted. Posterolateral margins of pereonite
1 without small, anteriorly projecting pro-
cesses. Anteriorly projecting processes on
anterolateral margins of pereonite 2, small,
tooth-like. Side plates of pereonites 3 and
4 not developed.
VOLUME 100, NUMBER 3
Fig. 1.
Antenna 1, 35% of body length, flagellum
of 4 articles, ratio to peduncle, 0.3:1. Right
antenna 2, flagellum of 2 articles, first con-
jointed; left flagellum of 3 articles.
Maxilliped, terminal article with 2 setae,
1 short and 1 curved.
Gnathopod 2, article 6 subquadrate, pal-
Mar margin with 2 small teeth on distal half,
unnotched. Pereopods 3 and 4 inserted next
to gills, both larger than those of males, both
segments setose. Pereopod 5, anterior mar-
Deutella aspiducha, male holotype. USNM 195178.
gin with tubercles. Abdomen with 2 lobes,
weakly setose.
Remarks. —Even the smallest individuals
of D. aspiducha show the beginnings of the
dorsal spines on the cephalon and pereonite
2. The spatulate processes of pereonite 2
and the middorsal humps and side plate
extensions of pereonites 3 and 4 in males
definitely exhibit allometry, being barely
visible in juveniles. The crenulations on the
margins of the side plates of pereonites 3
632
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig. 2. Deutella aspiducha, male paratypes. USNM 195179: Mx1, Mx2, Md, Ab. YPM 8261: Mxpd.
VOLUME 100, NUMBER 3 633
Fig. 3. Deutella aspiducha, female allotype. USNM 195177.
and 4 in males also become more distinct pereonite processes, their shapes when pres-
with increasing size. Females do not possess__ ent, and the 6th article of gnathopod 2 make
the small, projecting process on pereonite 2 this species notably sexually dimorphic.
until ovigerous. The possession or lack of The bodies of many of the animals, when
634
preserved, are bent into an inverted “U”’ at
the junctions of pereonites 2 and 3 and pe-
reonites 4 and 5; in some males, an addi-
tional flexion at the junction of pereonites
3 and 4 forms the animals into virtual
squares. All ovigerous females exhibit a
tight, inverted U-shape. Many individuals
are covered thickly with flocculent detritus
and morphological details can be obscured
unless ultrasonically cleaned.
The female allotype, with one antenna 2
triarticulate and the other biarticulate (but
with its first article conjointed) is an enigma.
Most caprellid genera, other than some in
the Phtisicidae, have a biarticulate flagel-
lum on antenna 2 (McCain 1968). Of 44
other specimens of D. aspiducha examined,
a few do appear to have a conjointed first
article on antenna 2 and one male does ap-
pear to have a triarticulate flagellum. Per-
haps triarticulation and conjointed articles
are mere exceptions to a generally conser-
vative feature or perhaps the occasional oc-
currence of these features has been over-
looked in other genera.
The number of small setae in the formula
for the terminal mandibular palp article
varies from 4 to 5. The distal setation of the
terminal maxillipedal palp varies from 1
short, to 1 short and 1 long, to 1 long. Distal
setation of the palp of the maxilliped is nor-
mally characteristic of only one genus
(Paracaprella) other than Deutella (McCain
1968).
Deutella aspiducha differs from all other
Deutella species (sensu McCain 1968) by
the lack of a triangular projection on the
penultimate article of the palp of the max-
illiped, by the shape of the last two articles
of pereopod 5, and by the point of insertion
of pereopod 5. The genus Deutella, how-
ever, exhibits a rather wide range of vari-
ation. Deutella schieckei Cavedini 1981
(possibly the Deutella sp. of Mayer (1890)),
and D. mayeri Stebbing, 1895, are both dor-
sally smooth; D. californica Mayer, 1890
and D. venenosa Mayer, 1890, the only oth-
er two species in the genus, possess dorsal
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
spines, as does D. aspiducha. Body spina-
tion is indeed often a variable character but
even conservative characters such as the
mouthparts, which McCain (1968) and
Laubitz (1970) believe offer some of the best
taxonomic characters, show significant
variability. The same latter three species,
for example, have a terminal article man-
dibular palp formula of 1-x-1; D. schieckei
(Cavedini, 1981) and D. mayeri (McCain,
1968), however, have only one or two setae
in toto on the palp. Pereopod 5 offers another
example of extreme variability. Pereopod 5
of D. venenosa is described by Mayer (1890)
as being “‘shortly rendered”’ and pereopod
5 of D. aspiducha is distinctly shorter than
pereopods 6 and 7; the size of pereopod 5
in the other three species does not differ
distinctly from that of pereopods 6 and 7.
Based on the above examples, a case for
a Strict generic diagnosis could be made in
such a way as to eliminate any one of the
Deutella species and allow the others to re-
main united. We believe that the structure
and proportions of the antennae of D. as-
piducha, most of its mouthpart morpholo-
gy, the various pereonite projections, the
delineation of the palm of gnathopod 2 in
males, the strong anterior insertion of
gnathopod 2 on pereonite 2, the shape,
length, and insertion patterns of pereopods
3 and 4, and the structure of the male ab-
domen all link this new species to the genus
Deutella.
Deutella aspiducha was first discovered at
the YPM in a vial of specimens examined
by Kunkel and labelled Protellopsis stebbin-
gii Pearse, Bermuda, with no precise habitat
description. That the species is indeed an
inshore species was ascertained by the col-
lection of a small male from hydroids at a
1 meter depth in 1985. Subsequent exam-
ination of Iliffe’s material from Little River
Cave, collected in 1982, revealed many D.
aspiducha of both sexes at all growth stages.
Because the inshore amphipod fauna of Ber-
muda has been so poorly surveyed and stud-
ied, however, at this time no suggestion of
VOLUME 100, NUMBER 3
a stygophilic nature should be ascribed to
this species. At present, its occurrence in
caves and in shallow inshore waters can be
said only to support the statement of several
authors (e.g., Stock 1986, Maddocks and
Iliffe 1986) that species found in the marine
waters of certain types of island sea caves
often also occur in shallow inshore waters.
Material examined. —Male holotype,
USNM 195178, Little River Cave, Ber-
muda, T. M. Iliffe, 8 May 1982, 4.8 mm.—
Female allotype, USNM 195177, Little
River Cave, Bermuda, T. M. Iliffe, 8 May
1982, 4.2 mm.—19 paratypes, USNM
195179, Little River Cave, Bermuda, T. M.
Iliffe, 8 May 1982.—USNM Acc. No.
359182, Little River Cave, Bermuda, T. M.
Iliffe, 8 May 1982, 152 specimens.— 1 male
paratype, YPM 8261, Bermuda, Dr. J.
L. Cole, 15 Jul 1903.—3 male paratypes,
YPM 8263, Bermuda, Dr. J. L. Cole, 15 Jul
1903.—1 male paratype, YPM 8262, Shelly
Bay, Hamilton Parish, Bermuda, off pro-
montory NNW of Bay, M. F. Gable, 3 Jun
1985, among hydroids, shallow subtidal.
Deutella incerta (Mayer) 1903
Fig. 4
Protellopsis stebbingii: Kunkel, 1910:111-
113, fig. 43.
Deutella incerta: Steinberg and Dougherty,
1957:281, 285-286.
Luconacia incerta: McCain, 1968:53-—54,
68-72, figs. 33-35.—McCain and Stein-
berg, 1970:53.—Johnson, 1986:381, fig.
125.
Remarks. —Steinberg and Dougherty
(1957) synonymized the monotypic Luco-
nacia of Mayer (1903) with Deutella. Their
argument for combining the genera rested
on differences between the genera that they
deemed trivial, the most important of which
is the form of the distal segments of pereo-
pod 5. Steinberg and Dougherty (1957) also
claimed that swimming setae were men-
tioned in the diagnosis of Luconacia (Deu-
635
tella lacks them), but that illustrations in
various papers never showed them.
Interestingly, McCain (1968) himself
questioned the use of pereopods 3, 4, and
5 as generic characters, because of their var-
1ous degrees of reduction. Nonetheless, he
re-established Luconacia and claimed sev-
eral important differences between the gen-
era (yet he eliminated the presence of swim-
ming setae from his diagnosis of Luconacia):
apical setae are present on the terminal ar-
ticle of the maxillipedal palp and a projec-
tion is present on the penultimate article in
Deutella and both are lacking in Luconacia;
the terminal article of the mandibular palp
in Deutella has no knobs and a setal formula
of 1-x-1 or “1 or 2,” and in Luconacia there
is a knob with a formula of 1-x-1; in Deu-
tella the tip of the male abdominal appen-
dage is without papillae and fringed or not
fringed, and in Luconacia both papillae and
fringes are present; the female abdomen in
Deutella is lobed and in Luconacia it is not;
in Deutella, pereopod 5 is inserted poste-
riorly and in Luconacia, midlength; finally,
the right lacinia is 5-toothed or serrate in
Deutella and serrate in Luconacia.
The discovery of D. aspiducha provides
us with a transitional form that makes a case
for uniting Deutella and Luconacia once
again, increasing the number of known Deu-
tella species to six. D. aspiducha possesses
apical setae on the terminal article of the
maxillipedal palp yet lacks the projection
on the penultimate article. The terminal ar-
ticle of the mandibular palp bears a knob
as in Luconacia (sensu McCain). The tip of
the male abdominal appendage of D. aspi-
ducha is without papillae and is not fringed;
the female abdomen is lobed. The insertion
of pereopod 5 is neither midlength nor pos-
terior in D. aspiducha but almost precisely
halfway in between. In addition, the prop-
odus of pereopod 5 lacks a palmar margin
as does that of McCain’s Luconacia. The
right lacinia of D. aspiducha 1s serrate.
Finally, McCain’s (1968) suggestion that
the removal of D. mayeri from the genus
636
Fig. 4. Deutella incerta. Cephalon of female. USNM
Acc. No. 359182.
would make it an exclusive Pacific Ocean
genus and Luconacia, then, an exclusive At-
lantic Ocean genus, loses a foundation for
further consideration with Cavedini’s (1981)
discovery of the Mediterranean D. schieckei.
Until these rather obscure, geographically
widely separated and poorly studied species
are much better known, we believe that the
overlapping characters among them, punc-
tuated in particular by the definitely Deu-
tella-like (and Luconacia-similar) D. aspi-
ducha, argue strongly for absorbing
Luconacia into Deutella.
Many of the specimens of D. incerta from
Bermuda of moderate to large size, both
male and female, have gnathopod 2 palmar
margins much more heavily setose than
those figured by McCain (1968). In addi-
tion, these same specimens possess two large
spines on the cephalon, one pre-ocular and
one sub-ocular (Fig. 4), not figured by
McCain (1968). Body spination at the
species level is often of questionable taxo-
nomic value (McCain 1968), but the invari-
able possession of these cephalic spines in
moderate to large sized Bermuda specimens
is a distinctive population character worth
noting.
Deutella incerta seems to be one of the
most commonly encountered caprellids in
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Bermuda. It has been found associated with
hydroids in shallow inshore waters and in
several caves. Johnson (1968) stated that it
is also common on mangrove roots, Sar-
gassum, Thalassia, sponges, and ascidians.
The many specimens at Kunkel’s disposal
were presumably from non-cave waters.
Material examined.—YPM 8212, Ber-
muda, Dr. J. L. Cole, 15 Jul 1903, 186 spec-
imens.— YPM 8264, Shelly Bay, Hamilton
Parish, Bermuda, off promontory NNW of
Bay, M. F. Gable, 3 Jun 1985, among hy-
droids, shallow subtidal, 1 specimen.—
USNM Acc. No. 359182, Palm Cave, Ber-
muda, T. M. Iliffe, 13 Mar 1982, 87 spec-
imens; Palm Cave, Bermuda, T. M. Iliffe,
13 Mar 1982, from hydroids, 4 specimens;
Little River Cave, Bermuda, T. M. Iliffe, 8
May 1982, 37 specimens; Cripplegate Cave,
Bermuda, T. M. Iliffe, 22 Jul 1982, 1 spec-
imen.
Family Caprellidae White, 1847
Caprella danilevskii
Czerniavski, 1868
Caprella danilevskii: Kunkel, 1910:110-
111.—McCain, 1968:22-25, fig. 10.—
McCain and Steinberg, 1970:16-17.—
Johnson, 1986:381, fig. 125.
Remarks. — According to McCain (1968),
this species is easily distinguished from all
other species of Caprella in the western
North Atlantic by its elongate gills whose
axes lie parallel to the body, by its distinc-
tive abdomen, and by the short dactyl on
male gnathopod 2. Kunkel saw no speci-
mens of this species but relied on accounts
of it for Bermuda in the Challenger Report
(Stebbing 1888). Johnson (1986) claimed it
is common in turtle grass in Bermuda. De-
spite such reports in the literature, no spec-
imens from Bermuda were found in the
available museum collections, nor were any
collected recently. Because of its distinctive
appearance, one can only assume that re-
ports of its occurrence in Bermuda are ac-
curate.
VOLUME 100, NUMBER 3
Caprella equilibra Say, 1818
Caprella equilibra: Kunkel, 1910:106—108,
fig. 41.—McCain, 1968:25-30, figs. 12-
13.—McCain and Steinberg, 1970:19-
21.—Johnson, 1986:381, fig. 125.—Lazo-
Wasem and Gable, 1987:335-336, fig. 10.
Caprella bermudia: Kunkel, 1910:108—110,
fig. 42.—Lazo-Wasem and Gable, 1987:
335-336, fig. 10.
Remarks.—Every specimen examined,
even the smallest, possesses the ventral tooth
between the insertions of the second gnatho-
pods. This species in Bermuda has a wide
size range. Caprella equilibra in Bermuda is
found, as is D. incerta, in near-shore waters
and in caves. It has been collected from hy-
droids, and Johnson (1986) stated it is also
common on bryozoans, ascidians, 7halas-
sia, and algae.
Material examined.—YPM 8223, Ber-
muda, Dr. J. L. Cole, 15 Jul 1903, lectotype
male of C. bermudia Kunkel (6.3 mm).—
YPM 8208, Flatts Village, Bermuda, col-
lector unknown, 4 Jul 1898, 2 males, | fe-
male.—YPM 8265, Shelly Bay, Hamilton
Parish, Bermuda, off promontory NNW of
Bay, M. F. Gable, 3 Jun 1985, among hy-
droids, shallow subtidal, 1 male. —USNM
collection, Bermuda, G. Brown Goode,
1876-1877, 1 ovigerous female.—USNM
Acc. No. 359182, Cripplegate Cave, Ber-
muda, T. M. Iliffe, 22 Jul 1982, 9 speci-
mens.
Hemiaegina minuta Mayer, 1890
Hemiaegina minuta: McCain, 1968:61-64,
figs. 29-30.—McCain and Steinberg,
1970:51.
Remarks. —McCain (1968) stated that /7.
minuta was found “‘in several localities near
Bermuda,” and in 1970 McCain and Stein-
berg gave its distribution as “‘off Bermuda.”
They also mentioned that the species had
been collected in plankton tows and from
Sargassum. Two specimens, presumably
from near-shore waters, were discovered in
637
the vial of Protellopsis stebbingii used by
Kunkel. Two other specimens were taken
intertidally from hydroids under rocks in
May 1985, a time when no Sargassum was
in evidence. Very likely, therefore, H. mi-
nuta 18 a permanent inshore inhabitant of
Bermuda waters, even if new populations
arrive with each wash of Sargassum.
Specimens collected from such an inshore
Sargassum wash in January 1987 by T. M.
Iliffe greatly exceeded the maximum sizes
given by McCain (1968). Iliffe collected a
male of 4.8 mm length (McCain’s largest
male = 4.0 mm) and an ovigerous female
of 5.4 mm length (McCain’s largest fe-
male = 3.2 mm). Personal observation of
thousands of specimens of H. minuta from
New England waters always attested to the
specific epithet chosen by Mayer; perhaps
if he had seen some of the Bermuda spec-
imens the epithet chosen would have been
different. |
Material examined.—YPM 8240, Ber-
muda, Dr. J. L. Cole, 15 Jul 1903, 2 spec-
imens.—YPM 8266, Whalebone Bay, St.
George’s, Bermuda, A. J. Baldinger, 22 May
1985, hydroids under rock, 2 specimens.
Non-catalogued: St. George’s, Bermuda, T.
M. Iliffe, 10 Jan 1987, Sargassum, 8 spec-
imens.
Family Dodecadidae Vassilenko, 1968
Fallotritella biscaynensis
McCain, 1968
Fallotritella biscaynensis McCain, 1968:57-
61, figs. 27—28.—McCain and Steinberg,
LOTTO ile
Remarks. —McCain (1968) stated that this
species is difficult to detect because of its
small size and its usual covering of detritus.
The five specimens we examined were in-
deed all detritus-covered; special care by one
not familiar with caprellids would be nec-
essary in dealing with this species so as not
to confuse it with D. aspiducha. The only
habitat records for the species are from red
algae (McCain 1968); one of our specimens
638 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
was taken from Avrainvillea, a green alga,
and two other specimens were collected from
Thalassia.
Material examined. —YPM 8267, Ferry
Reach, St. George’s, Bermuda, cove W of
BBS, M. F. Gable, 28 May 1985, from
Avrainvillea, 1 female. —YPM 8268, Shelly
Bay, Hamilton Parish, Bermuda, off pro-
montory NNW of Bay, M. F. Gable, 3 Jun
1985, from mixed subtidal algae, 1 fe-
male.-—USNM Acc. No. 346847, Ferry
Reach, Bermuda, adjacent to W side of dock
at BBS, M. L. Jones, 2 Sep 1983, Cohen
net through Thalassia, 1 juvenile female. —
USNM Acc. No. 359182, Castle Harbour,
Bermuda, T. M. Iliffe, 24 Aug 1984, 1 male.
Non-catalogued: Ely’s Harbour, Bermuda,
T. M. Iliffe, 21 Nov 1986, from Thalassia,
1 male.
Key to the Caprellids of Bermuda
(Pereopods 5, 6, and 7 present)
1. Pereopod 5 much smaller than 6 and
7, not inserted posteriorly on pereo-
nite, terminal segment not claw-like
— Pereopod 5 similar to 6 and 7 in size
or, if shorter and thinner, with a
claw-like terminal segment ...... 3
2. Insertion of pereopod 5 at mid-seg-
ment; peduncular segments of an-
tennay | withitew, Setdcuens sae
aT eee a Rae ies Deutella incerta
— Insertion of pereopod 5 75% back
from front edge of segment; pedun-
cular segments of antenna | heavily
SElLOSC eae: Deutella aspiducha n. sp.
3. Pereopods 3 and 4 present, of only
1 segment, animal seldom longer
tha Oem a gey ts leeees coy ea aera 4
— Pereopods 3 and 4 absent, no man-
dibular palp, animal can be much
larger, thang. miinniiine aes eee 5
4. Dorsal surface of body with spines,
atleast onyheads Moy see
Animal usually with all appendages
splayed out in same plane as body,
dorsal body with no spines; 2 teeth
between insertions of 2nd gnatho-
POdS #6 GC Ca ecat Hemiaegina minuta
. Gills with long axis parallel to body,
dactyl (last segment) of male 2nd
gnathopod less than half as long as
AIbIClE\Gr dan yee Caprella danilevskii
Gills normal, long axis perpendic-
ular to body; ventral tooth promi- —
nent between insertions of 2nd
enathopods ee owas es C. equilibra
Key to the Caprellids of Bermuda
(Pereopods 5, 6, and 7 missing)
. Pereopods 3 and 4 present, small,
OH Cally | Or 2 eSuonesnws 2.6060. yD)
Pereopods 3 and 4 absent ....... 2)
. Pereopods 3 and 4 of 1 segment .. 3
Pereopods 3 and 4 of 2 segments,
mandibular palp present ......... 4
. Dorsal surface of body with spines,
at least.oni head) 4.2) ieee
a ag cane Fallotritella biscaynensis
Animal usually with all appendages
splayed out in same plane as body,
no dorsal body spines; 2 teeth be-
tween insertions of 2nd gnathopods
FORME Ok BRON ts, 5 Hemiaegina minuta
. Insertion of pereopod 5 (even if
missing) 75% back from front edge
of segment; peduncular segments of
antenna | heavily setose .........
Po ren Rea Ae Deutella aspiducha n. sp.
Insertion of pereopod 5 (even if
missing) at mid-segment; pedun-
cular segments of antenna 1 with few
SCAG. ite aeo Geen rae Deutella incerta
. Gills with long axis parallel to body,
dactyl (last segment) of male 2nd
gnathopod less than half as long as
ATTICS Oa ies oy wee Caprella danilevskii
Gills normal, long axis perpendic-
ular to body; ventral tooth promi-
nent between insertions of 2nd
HONathOPOd nial qwouee Sc C. equilibra
VOLUME 100, NUMBER 3
Summary
The caprellid fauna of Bermuda has been
poorly and only incidentally collected and
studied. Although not rich in numbers of
species, some of the species so far discov-
ered raise interesting taxonomic questions,
as indicated in this paper. The Bermuda
caprellids also raise several interesting eco-
logical questions. As the collections of T.
M. Iliffe demonstrate, for example, signifi-
cant populations exist in many of the island
caves, yet no evolutionary conclusions can
be drawn because the inshore fauna has been
virtually neglected.
Acknowledgments
The authors thank Dr. J. L. Barnard
(USNM) for the loan of material, Dr. T. M.
Iliffe (BBS) for the collection of additional
specimens, Dr. M. A. Levin (ECSU) for
technical assistance, and Pedro Rivas-Diaz
(ECSU) for translation of Cavedini’s paper.
This study has been supported in part by a
Connecticut State University Research
Grant to the first author. Contribution No.
1131 from the Bermuda Biological Station
for Research.
Literature Cited
Bowman, T. E.,and L.G. Abele. 1982. Classification
of the Recent Crustacea.—Pp. 1-24 in D. E.
Bliss, ed., The Biology of Crustacea, vol. 1, L.
G. Abele, ed., Systematics, the fossil record, and
biogeography. Academic Press, New York.
Cavedini, P. 1981. Contributo alla conoscenza dei
caprellidi del Mediterraneo (Crustacea, Am-
phipoda).—Bolletino del Museo Civico di Sto-
ria Naturale di Verona 8:493-531.
Johnson, S. E. 1986. Order Amphipoda. Pp. 372-
381 in W. Sterrer, ed., Marine fauna and flora
of Bermuda. John Wiley & Sons, New York.
Kunkel, B. W. 1910. The Amphipoda of Bermuda. —
Transactions of the Connecticut Academy of Arts
and Sciences 16:1-116.
Laubitz, D. R. 1970. Studies on the Caprellidae
(Crustacea, Amphipoda) of the American North
Pacific.—National Museum of Natural Sci-
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ences. Publications in Biological Oceanography
No. lu-—vii + 1-89.
Lazo-Wasem, E. A.,and M. F. Gable. 1987. A review
of recently discovered type specimens of Ber-
muda Amphipoda (Crustacea: Peracarida) de-
scribed by B. W. Kunkel (1882-1969).—Pro-
ceedings of the Biological Society of Washington
100:32 1-336.
Maddocks, R. F., and T. M. Iliffe. 1986. Podocopid
Ostracoda of Bermudian caves.—Stygologia 2:
26-76.
Mayer, P. 1890. Die Caprelliden des Golfes von Nea-
pel und der angrenzenden Meeres-Abschnitte.
Eine Monographie. Nachtrag zur Monographie
derselben.—Fauna und Flora des Golfes von
Neapel 17:vii + 157 pp.
McCain, J.C. 1968. The Caprellidae (Crustacea: Am-
phipoda) of the western North Atlantic. —Bul-
letin of the United States National Museum 278:
i-vi + 1-147.
1970. Familial taxa within the Caprellidea
(Crustacea: Amphipoda).— Proceedings of the
Biological Society of Washington 82:337-342.
,and J. E. Steinberg. 1970. Caprellidae I. Fam-
ily Caprellidae. Pp. 1-78 in H.-E. Gruner and
L. B. Holthius, eds., Crustaceorum Catalogus.
Amphipoda I, part 2. The Hague, The Neth-
erlands: Dr. W. Junk Publishers.
Schram, F. R. 1986. Crustacea. Oxford University
Press, New York. 606 pp.
Stebbing, T. R. R. 1888. Report on the Amphipoda
collected by H.M.S. Challenger during the years
1873-76.—Report on the Scientific Results of
the Voyage of H.M.S. Challenger during the years
1873-76, Zoology 29:xxiv + 1737 pp., 210 pls.
Steinberg, J. E.,and E. C. Dougherty. 1957. The skel-
eton shrimps (Crustacea: Caprellidae) of the Gulf
of Mexico.— Tulane Studies in Zoology 5:267-
288.
Stock, J. H. 1986. Two new amphipod crustaceans
of the genus Bahadzia from ‘blue holes’ in the
Bahamas and some remarks on the origin of the
insular stygofaunas of the Atlantic.— Journal of
Natural History 20:921-933.
(MFG) Department of Biology, Eastern
Connecticut State University, Willimantic,
Connecticut 06226-2295; (EAL-W) Divi-
sion of Invertebrate Zoology, Peabody Mu-
seum of Natural History, Yale University,
170 Whitney Avenue, P.O. Box 6666, New
Haven, Connecticut 06511-8161.
PROC. BIOL. SOC. WASH.
100(3), 1987, pp. 640-652
TWO NEW SPECIES AND A NEW GENUS OF
MINIATURE CHARACID FISHES (TELEOSTEI:
CHARACIFORMES) FROM NORTHERN
SOUTH AMERICA
Stanley H. Weitzman and Richard P. Vari
Abstract. —The first known miniature characid fishes apparently aligned with
the Characinae and Cynopotaminae are described as new from the Rio Negro
and Rio Orinoco drainages of Venezuela and tributaries of the Rio Amazonas
in Colombia. Although the suggested relationships of the new genus (Priocha-
rax) and species (P. ariel and P. pygmaeus) to those subfamilies appear rea-
sonable, their exact phylogenetic relationships within the Characinae and Cy-
nopotaminae remain obscure. The new species are distinguished from others
in these subfamilies primarily by a higher number of jaw teeth, a lower number
of pelvic- and anal-fin rays, retention of larval pectoral fins in adults, and a
minute adult body size of a maximum of about 17 mm in standard length.
The early explorer-naturalists who sam-
pled the South American freshwater fish
fauna focused nearly exclusively on species
of moderate to large body size, evidently
under the mistaken belief that all smaller
fishes were juveniles, or if distinct the species
were unimportant. Agassiz, during the
Thayer Expedition to Brazil in 1865, was
the first collector who fully endeavored to
collect even the smallest fishes, recognizing
that such specimens often represented in-
teresting species of small adult size. During
the twelve decades that have passed since
that trip numerous species of relatively small
adult body sizes have been described from
the freshwaters of South America. Recent
collecting efforts in Venezuela have yielded
miniature species of the family Lebiasinidae
(Fernandez and Weitzman 1987) and the
subfamily Characidiinae of the Characidae
(Weitzman 1986). Those collections also re-
vealed the existence of a miniature species
evidently aligned phyletically with the char-
acid subfamilies Characiinae and Cynopo-
taminae. A second, very similar species,
originally collected in the Colombian Am-
azon, was subsequently found in the collec-
tion of the Naturhistoriska Riksmuseet,
Stockholm. These two species, assigned
herein to a new genus, are described as new
and the possible phylogenetic relationships
of these taxa are discussed.
Methods and Materials
The counts and measurements are those
described by Fink and Weitzman (1974: 1-
2). All measurements other than standard
length (SL) are expressed as a percentage of
SL except subunits of the head which are
expressed as a percentage of head length or
as otherwise noted.
Specimens examined for this study are
deposited in the American Museum of Nat-
ural History, New York (AMNH); Acade-
my of Natural Sciences of Philadelphia
(ANSP); British Museum (Natural History),
London (BMNH); California Academy of
Sciences, San Francisco (CAS); Field Mu-
seum of Natural History, Chicago (FMNH);
Museo de Biologia, Universidad Central de
Venezuela, Caracas (MBUCV); Museu de
Ciéncias, Pontificia Universidade Catolica
do Rio Grande do Sul, Porto Alegre (MCP);
VOLUME 100, NUMBER 3
Museum of Comparative Zoology, Harvard
University, Cambridge (MCZ); Museum
d’Histoire Naturelle, Genéve (MHNG);
Museum National d’Histoire Naturelle,
Paris (MNHN); Museu de Zoologia da Uni-
versidade de Sao Paulo (MZUSP); Natur-
historiska Riksmuseet, Stockholm (NRM);
Naturhistorisches Museum Wien, Vienna
(NMW); University of Michigan, Museum
of Zoology, Ann Arbor (UMMZ); National
Museum of Natural History, Smithsonian
Institution, Washington, D.C. (USNM); and
Institute of Taxonomic Zoology (Zodlo-
gisch Museum), Amsterdam (ZMA).
Priocharax, new genus
Diagnosis. —New World characid fish
group distinguished from all other characids
by retaining larval rayless pectoral fin in
adults. Distinguished from all other New
World characids except some members of
Characinae and Cynopotaminae by having
lower jaw (dentary bone) with approxi-
mately 30 to 50 conical teeth in somewhat
irregular single row, and upper jaw (pre-
maxilla and maxilla) with approximately 45
to 90 conical teeth in somewhat irregular
single row. Adult body size ranges from
about 11 to 17 mm standard length; such
diminutive adult body size otherwise un-
known in Characinae and Cynopotaminae.
Within the tribe Characini of the Cha-
racinae (sensu Géry 1977:302-—306) and the
Cynopotaminae (Menezes 1976) 16 to 22
branched anal-fin rays of Priocharax further
distinguish its species from Acanthocharax
Eigenmann, Acestrocephalus Eigenmann,
Asiphonichthys Cope, Charax Scopoh, Cy-
nopotamus Valenciennes, Eucynopotamus
Fowler, Galeocharax Fowler, Gilbertolus
Eigenmann, Gnathocharax Fowler, Hetero-
charax Eigenmann, Hoplocharax Géry,
Lonchogenys Myers, Moralesia Fowler,
Roestes Ginther, and Roeboides Gunther
all with 26 or more branched anal-fin rays.
Priocharax posesses 1,5 pelvic-fin rays, dis-
criminating it from all other members of
641
Characinae and Cynopotaminae, all with 1,7
pelvic-fin rays.
Type species. —Priocharax ariel, new
species.
Etymology. —Prio from the Greek for saw,
and charax from the Greek characo mean-
ing pointed stake, here used in reference to
the characid genus Charax, hence a characid
fish. Priocharax means a saw-bearing char-
acid fish in reference to its numerous small
jaw teeth.
Key to the Species of Priocharax
1. Caudal peduncle depth about 32 to
46 percent of caudal peduncle length;
dentary teeth about 38 to 55; lower
limb gill-rakers 11 to 13°.........
Priocharax ariel, new species
— Caudal peduncle depth about 52 to
65 percent of caudal peduncle length;
dentary teeth about 28 to 36; lower
limb gill-rakers 8 to 10
....Priocharax pygmaeus, new species
Priocharax ariel, new species
Figs. 1-5, Table 1
Holotype. —MBUCV V-15340, male, SL
14.5 mm, Venezuela, Territorio Federal
Amazonas, Departamento Rio Negro, Cano
Manu, tributary of Rio Casiquiare approx-
imately 250 m upstream from Solano,
02°00'N, 66°57'W;, R. P. Vari, C. J. Ferraris,
Jr., O. Castillo, and J. M. Fernandez, 7 Dec
1984.
Paratypes.— Following 16 lots collected
with holotype and deposited in cited insti-
tutions; all museums received 25 specimens
unless otherwise noted; extremes of SL are
given only for USNM specimens, lengths of
other series fall within that range: USNM
272619, SL 8.8-17.1 mm, 1229 specimens,
41 cleared and counterstained for cartilage
and bone; AMNH 57007; ANSP 158006;
BMNH 1986.2.4:1-25; CAS 57944; FMNH
96689: MBUCV V-15341, 100 specimens;
MCP 9953; MCZ 63031; MHNG 2239.48;
MNHN 1986-303; MZUSP 3647, 50 spec-
642
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Table 1.—Morphometrics of Priocharax ariel and P. pygmaeus. Males of P. ariel all with anal-fin hooks.
Specimens of P. ariel and P. pygmaeus of undetermined sex lack anal-fin hooks. Standard length is expressed
in mm. First group of measurements are presented as percentages of standard length; second group as percentages
of head length.
Priocharax ariel males
Holo-
type n Range BX
Standard length 14.5 27 11.8-15.1 14.0
Depth at dorsal-fin origin 24.8 27 21.8-24.8 23.4
Snout to dorsal-fin origin 55.9 27 53.2-55.9 54.6
Snout to pelvic-fin origin 42.4 27 39.5-42.5 40.9
Snout to anal-fin origin 538 27 S3O=D00 S47
Peduncle depth VS 2 P33 7.8
Peduncle length 21.4 27 19.2-23.2 20.8
Pelvic-fin length WO 27 JOA SM3 0) Wes
Dorsal-fin height Davee DS 22 0=264= 23.7
Anal-fin anterior lobe
length AO D> WILDS. AN
Bony head length 23.4 27 23.0-26.4 24.4
Horizontal eye diameter 31.8 27 27.7-33.8 31.5
Snout length IBS) BY ZOO 22S
Interorbital width SSM le 132 la swe 34.2.
Upper jaw length 55.9 27 49.9-60.0 55.8
Caudal peduncle depth as
percent of caudal
peduncle length 35.5 26 33.3-42.4 36.8
imens; NRM A86/1984495.3603, 50 spec-
imens; NMW 81788; UMMZ 213500, 35
specimens, 10 cleared and counterstained
for cartilage and bone; and ZMA 119.456.
All following paratypes collected in Vene-
zuela, Territorio Federal Amazonas, R. P.
Vari and party unless otherwise noted:
USNM 272613, SL 13.9 mm, 1, Departa-
mento Rio Negro, lagoon northeast of air-
port at San Carlos de Rio Negro, 01°55’N,
67°02'W, 4 Dec 1984. USNM 272614, SL
14.0 mm, 1, Departamento Rio Negro, Cano
Chola, where crossed by road from San Car-
los de Rio Negro to Solano, 01°58’N,
67°00’W, 5 Dec 1984. USNM 272615, SL
11.4-13.6 mm, 10, Departamento Rio Ne-
gro, small cano off Cano Urami, just upriver
of Santa Lucia, 01°17'N, 66°51'W, 6 Dec
1984. AMNH 57008, SL 13.1 mm, 1, De-
partamento Rio Negro, Rio Negro at Santa
Lucia, 01°17’N, 66°52’W, 4 Feb 1984, C. J.
Ferraris and party. USNM 272618, SL 13.7—
Priocharax pygmaeus
Priocharax ariel sex undetermined
sex undetermined
Holo-
n Range x type n Range x
27) MACS=17 1 ARDS G4 a OSS =NGx4 eel)
27 220-2524, 23.9) 2454 37 232-2 46 Bn
LY SESS a8) Ses) SUY=9O.3 S327
27 39.1-42.4 40.7 42.1 7 42.1-44.6 43.6
DT 32-8562)" 54535525) 71 58.6 —S Ol Om SAO
2 ALO) 8.2 Hee} ccd) 7.9-8.4 8.2
27. 18sl=2307) 22.2, 922), De See lin
Sel Nell WO. 7/ 9.1-11.6 10.4
26 21.1-25.2 23.7 24.4 7 24.1-25.7 24.9
259° 2050=22799 2k Oe eee? OR eee ORO
27, 2258=2573 224 SO ee LOG eee
27 29.7-34.4 32.3 30:0" 7 *30,0=35233"1
27 A9,7=24°4 22565) 2.4 a9 42 eel
26.32.1389) 6352354) 39.0 eo Bo Ol
27 50.0-60.0 56.1 48.3 7 43.3-50.9 47.2
27 32.1=46:0) = 392 6520) 7 5226-65 Om Onl
14.9 mm, 3, Departamento Rio Negro, Cano
Loro, where crossed by road from San Car-
los de Rio Negro to Solano, 01°59'N,
66°58’W, 7 Dec 1984. USNM 272616, SL
11.9 mm, 1, Departamento Ature, small
cano crossed by road from Puerto Ayacucho
to Samariapo, 2 km south of Mirabel,
05°25'N, 67°46’W, 12 Dec 1984. USNM
272617, SL 12.2-12.9 mm, 2, Departa-
mento Ature, Rio Platanillal, where crossed
by road from Puerto Ayacucho to Samar-
iapo, 05°37'N, 67°35'W, 2 Dec 1984.
Diagnosis.—Two species of Priocharax
diagnosable on basis of complete separation
in one morphometric and two meristic
characters. In both sexes of Priocharax ariel
caudal peduncle depth about 32 to 46 per-
cent of caudal depth, contrasted to about 52
to 65 percent in P. pygmaeus. Priocharax
ariel with greater number of lower limb gill
rakers on anterior gill-arch, 11 to 13, than
P. pygmaeus, 8 to 10. Large individuals of
VOLUME 100, NUMBER 3
643
Bie le
A, Priocharax ariel, new species, holotype, MBUCV V-15340, male SL 14.5; Venezuela, Departa-
mento Rio Negro, Cano Manu; B, Priocharax ariel, new species, paratype, USNM 272619, female, SL 15.3
mm; same locality as holotype.
both species with greater number of dentary
teeth, but counts do not overlap between
species. Specimens of P. ariel with 38 to 55
dentary teeth (x = 44, n = 41), those of P.
pygmaeus with 28 to 36 teeth (X = 32.5, n =
10). Other meristic characters probably will
show significant differences, but specimens
of P. pygmaeus too poorly preserved to al-
low definitive analysis without clearing and
staining many specimens. Some overlap oc-
curs in all following characters, but covari-
ance analysis of larger numbers of P. pyg-
maeus probably would show distinct
differences between these species. Branched
anal-fin rays 16 to 21 in P. ariel (x = 18.5,
n = 96) and 20 to 22 in P. pygmaeus (X =
21.0, n = 16); upper limb rakers 3 to 5 in
P. ariel (X = 3.9, n = 41) and 2 or 3 in P.
pygmaeus (X = 2.2, n = 10); premaxillary
teeth 22 to 34 in P. ariel (X = 27.7, n = 41)
and 19 to 24 in P. pygmaeus (X = 21.7, n =
10); maxillary teeth 38 to 58 in P. ariel
(x = 47.7, n = 41) and 27 to 41 in P. pyg-
maeus (X = 31.4, n = 10).
Description.— Table 1 presents morpho-
metrics of holotype and measured para-
types. See Fig. 1A, B for body form. Body
slender, sides compressed. Greatest body
depth at dorsal-fin origin. Dorsal-fin origin
somewhat closer to caudal-fin base (at pos-
terior of hypural fan and anterior of caudal-
fin rays) than to tip of snout. Dorsal-fin or-
igin approximately intersected by vertical
line through anal-fin origin. Pelvic-fin ori-
644
Fig. 2. Anterolateral view of jaws of Priocharax
ariel, new species, paratype, UMMZ 213500, male, SL
16.7 mm, same locality as holotype.
gin nearly midway between anal-fin origin
and vertical line through most posterior
point of opercle. Snout bluntly rounded.
Dorsal profile of head and body gently con-
vex from snout tip to dorsal-fin origin. Dor-
sal profile of body along dorsal-fin base
nearly straight, gently sloping posteroven-
trally. Dorsal body and caudal peduncle
profile very slightly concave from dorsal-fin
insertion to base of procurrent rays. Caudal
peduncle slender, elongate, with sides com-
pressed. Ventral body profile gently convex
from symphysis of lower jaw to ventral to
pectoral-fin origin. Belly profile from that
point to vent and anal-fin origin may be
slightly convex, more often straight, or
sometimes slightly concave. Body profile
along anal-fin base and caudal peduncle to
anterior ventral procurrent rays gently con-
cave. Base of anal fin posterodorsally ori-
ented.
Head length approximately one-quarter
of standard length. Snout rounded in profile.
Jaws about equal, mouth terminal or lower
jaw slightly included. Mouth elongate,
somewhat posteroventrally inclined. Max-
illa elongate; upper jaw long, posterior bor-
der reaching to or posterior to vertical line
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
through posterior border of pupil of eye,
length half to nearly two-thirds of head
length. Eye about one-third of head length.
Fleshy interorbital width often over one-
third of bony head length, gently convex
transversely.
Maxilla with 38 to 58 teeth (X = 47.4, n =
41; teeth of holotype not counted); teeth
conical, small and slender, in single series
along ventral border of bone. Number of
teeth increasing with body length. Acces-
sory cusps absent. Premaxilla with 22 to 34
teeth (X = 27.7, n = 41) in a single series.
Premaxilla teeth similar in form to those of
maxilla. Dentary with 38 to 55 teeth (* =
44.6, n = 41). Dentary teeth in a single row
posteriorly; larger specimens with irregular
row of closely placed teeth anteriorly, but
two distinct anterior rows never present.
Anterior dentary teeth larger than those
posterior. All jaw teeth lingually curved to
a moderate extent (see Fig. 2).
Infraorbital series incomplete, with ant-
orbital bone only element ossified, apparent
only in cleared and stained specimens over
13.5 mm SL. Fronto-parietal fontanel large,
extending from ethmoid to supraoccipital,
completely separating parietals; frontals only
in contact at strong epiphyseal bar. Frontal
about of same width along entire length;
similar in form to that of small juvenile
characids. Supracccipital spine poorly de-
veloped (Fig. 3).
Dorsal-fin rays 11,9 in holotype; other ex-
amined specimens with 2 unbranched rays
followed by 8 to 10 mostly divided rays (x =
8.9, n = 96); usually with posterior ray or
more rarely posterior two rays not divided
to their base. Dorsal-fin height moderate,
20 to 25% of SL. Pectoral fin larval in form
(Fig. 4). Radials of two incompletely sepa-
rated hyaline cartilaginous flat plates artic-
ulating anteriorly with vertically elongate
cartilaginous coracoscapular plate and pos-
teriorly with striated actinotrich tissue
around which rays form in developing lar-
vae of other characids. In specimens about
14 to 15 mm SL coracoscapular plate par-
VOLUME 100, NUMBER 3
Fig. 3.
Lateral view of posterior portion of neu-
rocranium and Weberian apparatus of Priocharax ar-
iel, new species, paratype, UMMZ 213500, male, SL
16.7 mm, same locality as holotype.
tially ossified dorsally. Cleithrum ossified,
slender, attached dorsally to slender supra-
cleithrum by relatively loose ligamentous
tissue. No notch or spine on cleithrum. Cor-
acoid and other pectoral girdle elements
often not ossified, but slight ossification
present along ventral border in example
drawn. Posttemporal weakly ossified, not
shown in Fig. 4. Pelvic-fin rays 1,5 in all
counted specimens, n = 97. Posterior tip of
pelvic fin extending about to region of vent.
No hooks present on pelvic, dorsal or pec-
toral-fin rays. Anal fin 111,20 in holotype;
other examined specimens with three un-
branched rays followed by 16 to 21 branched
rays (X = 18.5, n = 96), posterior ray divided
to its base. Anal-fin hooks present in 35
specimens from Cano Manu, all other spec-
imens (1319) from that locality without
hooks. Hooks occur on posterior margin of
three anterior undivided rays and anterior
4 or 5 divided rays (Fig. 5). Usually 6 to 8
hooks per ray, with one short hook per ray
segment. Anal-fin margin concave, with an-
terior elongate lobe and posterior section of
short rays. Fin shape similar in specimens
with and without hooks. Caudal fin forked,
of moderate length; principal rays 10/9 in
91 specimens. Adipose fin absent.
645
\\\\
Fig. 4. Lateral view of pectoral girdle of Priocharax
ariel, new species, paratype, UMMZ 213500, male, SL
16.7 mm, same locality as holotype. Densely stippled
structures are cartilage.
Squamation almost completely absent in
most specimens, apparently lost in handling
during capture; following counts thus ten-
tative. Scales in lateral series on body about
31 or 32; with no indication of perforated
lateral line scales, but scales typically lack-
ing on anterior of body just posterior to
cleithrum and supracleithrum. Scale rows
between dorsal-fin origin and pelvic-fin or-
igin apparently 7. About 12 or 13 scale rows
around narrowest portion of caudal pedun-
cle. Predorsal scales perhaps 11 or 12; scales
often present immediately anterior to dorsal
fin but always absent or lost just posterior
to supraoccipital spine.
Vertebrae 32 to 34 (x = 33.1, n = 41).
Fig. 5.
of Priocharax ariel, new species, paratype, UMMZ
213500, male, SL 16.7 mm, same locality as holotype.
Lateral view of anterior nine anal-fin rays
646
Upper limb gill-rakers 3 to 5 (X = 3.9, n =
41) and lower limb gill-rakers 11 to 13 (¥ =
12.5, n = 41). Branchiostegal rays 4 (n =
31), 3 rays on anterior and | ray on posterior
ceratohyal.
Weberian apparatus (Fig. 3) well devel-
oped, all components except claustrum well
ossified. Os suspensorium exceptionally
large and projecting forward to anterior end
of third centrum. Neural spine of fourth ver-
tebra moderately developed. Neural pre-
zygopophyses of fifth centrum undevel-
oped. Lack of buttressing flanges on base of
fifth pleural rib, short neural spine of fourth
vertebra and little developed posterior crest
of enlarged supraneural all characteristic of
juvenile Weberian apparatus in characids.
Color in life.—Translucent without any
bright colors; some guanine pigmentation
overlying gasbladder; dark chromatophore
pattern distributed as discussed below for
color in alcohol.
Color in alcohol.—Body-color of holo-
type pale brown to white (Fig. 1). Dark chro-
matophores limited, distributed as shown
in Figs. 1A, B. Proportionally large chro-
matophores on head covering brain tissue.
Few scattered dark chromatophores on
opercle and along base of dorsal, anal and
pelvic fins. Dorsal and lateral region of cau-
dal peduncle with few scattered dark chro-
matophores, and base of caudal fin with dark
spot usually consisting of about 10 to 20
contracted chromatophores. Sometimes one
or two chromatophores at region of trian-
gular muscular hiatus in body wall posterior
to dorsal end of free opercular border. Dor-
sal, anal and caudal fins with scattered small
elongate dark chromatophores along bor-
ders of fin rays. A few dark chromatophores
present in region of vent.
Sexual dimorphism. — Males with anal-fin
hooks described above for anal fin. Anal-
fin hooks absent in females and juveniles.
Examined specimens apparently not fully
sexually mature. Females apparently with
developing eggs but no mature eggs found.
Males, even those with anal-fin hooks, with
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
only developing testes. Using anal-fin hooks
as a criterion, large population sample from
type locality with only 35 identifiable males
among 1740 specimens; remainder imma-
ture or females. Members of this population
perhaps approaching sexual maturity at time
of capture and many males not matured suf-
ficiently to develop anal-fin hooks.
Etymology. — Ariel from the Greek for an
airy spirit, in reference to the tiny and trans-
lucent nature of this fish in its natural hab-
itat.
Habitat. — The species is an inhabitant of
black acidic waters of the upper Rio Negro
and Rio Orinoco drainage systems. The ma-
jority of specimens were captured in still
waters of shaded rain forest streams. Indi-
viduals from the extensive series captured
at the type locality were most common in
emergent vegetation, along the shore line
and around terrestrial plants that were
hanging into the water.
Priocharax pygmaeus, new species
Fig. 6, Table 1
Holotype.—NRM THO/1976303.1445,
6, SL 16.4 mm, Colombia, Departamento
Amazonas, in Quebrada Pajarito, tributary
of Quebrada Bacada, tributary of Quebrada
Matamata, a tributary of Rio Amazonas,
northwest of Leticia, about 04°41’S,
69°57'W; T. Hongslo, 21 Jul 1976.
Paratypes. —Collected with holotype, 87
specimens: NRM THO/1976303.1446, SL
8.0-11.4 mm, 41 specimens, 4 cleared and
counterstained for cartilage and bone;
USNM 278479, SL 8.1-12.2 mm, 36 spec-
imens, 6 cleared and counterstained for car-
tilage and bone; MBUCV V-15340, SL 10.0-
10.9 mm, 5 specimens; MZUSP 36498,
10.2-10.7 mm, 5 specimens.
Diagnosis. —See diagnosis of P. ariel.
Description.—Table | presents morpho-
metrics of holotype and measured para-
types. See Fig. 6A, B for body form. Body
slender, sides compressed. Greatest body
depth at dorsal-fin origin. Dorsal-fin origin
VOLUME 100, NUMBER 3
647
Fig. 6. A, Priocharax pygmaeus, new species, holotype, NRM THO/1976303.1445, SL 16.4 mm, Quebrada
Pajarito, Rio Amazonas system, northeast of Leticia, Colombia; B, Priocharax pygmaeus, new species, paratype,
USNM 278479, SL 10.8 mm, same locality as holotype.
somewhat closer to caudal-fin base (hypural
joint) than to tip of snout. Dorsal-fin origin
approximately intersected by vertical line
through anal-fin origin, or slightly in ad-
vance of that line. Pelvic-fin origin nearly
midway between anal-fin origin and vertical
through most posterior point of opercle.
Snout bluntly rounded. Dorsal profile of
head and body gently convex from snout tip
to dorsal-fin origin. Dorsal profile of body
along dorsal-fin base nearly straight, gently
sloping posteroventrally. Dorsal body and
caudal peduncle profile very slightly con-
cave from dorsal-fin insertion to base of
procurrent rays. Caudal peduncle slender,
elongate, with sides compressed. Ventral
body profile gently convex from symphysis
of lower jaw to ventral to pectoral-fin origin.
Belly profile from that point to vent and
anal-fin origin may be slightly convex, more
often straight, or sometimes slightly con-
cave. Body profile along anal-fin base and
caudal peduncle to anterior ventral procur-
rent rays gently concave. Base of anal fin
posterodorsally oriented.
Head length moderate, approximately
one-quarter of standard length. Snout
rounded in profile, of moderate length. Jaws
about equal, mouth terminal or lower jaw
slightly included. Mouth elongate, some-
what posteroventrally inclined. Maxilla
elongate; upper jaw long, posterior border
648
reaching approximately to or posterior to
vertical line through center of pupil of eye,
length half or somewhat less than half of
head length. Eye large, about one-third of
head length. Fleshy interorbital width wide,
often somewhat over one-third of bony head
length, gently convex transversely.
Maxilla with 27 to 41 teeth (x = 31.6,
n = 10; holotype with 41); teeth conical,
small and slender, in single series along ven-
tral border of bone. Larger specimens with
greater number of teeth. Accessory cusps
absent. Premaxilla with 19 to 24 teeth (x =
22.7, n = 10, holotype with 24) in a single
series. Premaxilla teeth similar in form to
those of maxilla. Dentary with 28 to 36 teeth
(x = 32.5, n = 10, holotype with 36). Den-
tary teeth in single row posteriorly; larger
specimens with irregular row of closely
placed teeth anteriorly, but two distinct an-
terior rows never present. Anterior dentary
teeth larger than posterior. All jaw teeth lin-
gually curved to a moderate extent.
Infraorbital series incomplete, with ant-
orbital bone only element ossified, apparent
only in cleared and stained specimen of 12.2
mm SL. Fronto-parietal fontanel large, ex-
tending from ethmoid to supraoccipital,
completely separating parietals; frontals only
in contact at strong epiphyseal bar. Frontal
about of same width along entire length;
similar in form to that of small juvenile
characids. Supraoccipital spine poorly de-
veloped.
Dorsal-fin rays 11,9 in 15 counted speci-
mens including holotype. Usually with pos-
terior ray or more rarely posterior two rays
not divided to their base. Dorsal-fin height
moderate, about 25% of SL. Pectoral fin lar-
val in form. Radials as described above for
P. ariel. Cleithrum ossified, slender, at-
tached dorsally to slender supracleithrum
by relatively loose ligamentous tissue. Cor-
acoid and other pectoral girdle elements ap-
parently not ossified or chondrified. Post-
temporal weakly ossified. Pelvic-fin rays 1,5
in all counted specimens, n = 15. Fin of
moderate length, posterior tip extending
about to region of vent. No hooks present
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
on pelvic, dorsal or pectoral-fin rays. Anal
fin v,21 in holotype; all examined speci-
mens with four or five unbranched rays (x =
4.1, n = 15) followed by 20 to 22 branched
rays (x = 21.7,n = 15), posterior ray divided
to its base. Anal-fin hooks not present in
any specimens. Anal-fin margin concave,
with anterior elongate lobe and posterior
section of short rays. Caudal fin forked, of
moderate length; principal rays 10/9 in 91
specimens. Adipose fin absent.
Squamation completely absent in all
specimens. Although some scale pockets
visible, their distribution not consistent
enough for counts.
Vertebrae 32 or 33 (X = 32.9, n = 9, ho-
lotype with 33). Upper limb gill-rakers 2 or
3 (X¥ = 2.2 n = 10, holotype with 3) and
lower limb gill-rakers 9 in all specimens.
Branchiostegal rays 4 (n = 15), 3 rays on
anterior and | ray on posterior ceratohyal.
Weberian apparatus well developed, sim-
ilar to that of P. ariel.
Color in life. —Report as “transparent,
faint pink” by T. Hongslo.
Color in alcohol.—Holotype pigmenta-
tion very similar to that of P. ariel (Fig. 6).
Fewer dark chromatophores present in
specimens of P. pygmaeus; however, those
specimens may have faded more in preser-
vative since they were collected eight years
earlier.
Etymology. —Pygmaeus from the Greek
for dwarf, in reference to the tiny size of this
fish.
Habitat. — The species was collected from
a shaded or partially shaded rainforest
stream about two meters wide in water to
about one meter deep. The bottom was
“clay” with much leaf litter and no sub-
merged plants. The water temperature was
25.2°C, the pH about 5.5 and the water was
somewhat turbid. Water type was not noted.
Relationships
Priocharax is an example of the problems
inherent in studies of the phylogenetic re-
lationships of miniature, paedomorphic
VOLUME 100, NUMBER 3
species. The species of Priocharax have or
appear to have plesiomorphic character
states relative to the conditions described
below in various characines and cynopot-
amines. Plesiomorphic features can be dis-
tinguished from paedomorphic characters
only when phylogenetic hypotheses and de-
velopmental information for characters in
ingroup and outgroup taxa are available.
Unfortunately the state of our understand-
ing of relationships among likely close rel-
atives of Priocharax is unsatisfactory (see
discussion below). As a consequence of that
situation it is beyond the scope of this study
to advance a detailed corroborated hypoth-
esis of phylogenetic relationships based on
shared derived characters. This lack of a
corroborated hypothesis of phylogenetic
relationships also means that we cannot
presently discriminate the pattern of plesio-
morphy versus paedomorphosis in Prio-
charax and its putative relatives. Polarity
statements and hypotheses of the paedo-
morphic nature of some features are instead
advanced within the context of broader out-
group comparisons. Fink (1982), Weitzman
and Fink (1983:345-346, 390), and Weitz-
man and Fink (1985:9-10) discussed the
issues raised by paedomorphic, possibly
progenetic, characters, especially where
outgroup information may be inadequate.
The phylogenetic relationships of Prio-
charax within the Characidae are difficult
to resolve satisfactorily. The apparent de-
rived and non-paedomorphic characters of
numerous conic teeth and elongate maxillae
in the genus suggest that its relationships lie
in the putative subfamily Characinae of Gery
(1977:302), in particular with the genera
Acanthocharax, Acestrocephalus, Charax,
Cynopotamus, Galeocharax, Gnathocha-
rax, Heterocharax, Lonchogenys, and Roe-
boides. A hypothesis of a phyletic associa-
tion of Priocharax to and within the
Characinae is complicated by the lack of any
proposed derived characters that support the
concept of the monophyly of the subfamily.
Géry (1977:295) attempted to delimit the
Characinae in his admittedly artificial key
649
to the subfamilies of his Characidae. Al-
though his key has some limited use in a
typological sense, no evidence was pre-
sented that his Characinae was monophy-
letie:
According to Géry the subfamily con-
sisted of three tribes: the Characini, the Bra-
mocharacini, and the Acestrorhynchini.
Weitzman and Fink (1983:342, 344), based
on Rosen (1972:12), questioned the recog-
nition of a separate tribe for Bramocharax
Gill and Bransford, and the assignment of
that genus to the Characinae. They agreed
with Rosen (1972) that Bramocharax is most
likely a derivative of Astyanax Baird and
Girard, a member of the large characid
subfamily Tetragonopterinae. Menezes and
Géry (1983:587, 588) recognized the Aces-
trorhynchini as a subfamily rather than as
a tribe and noted that at present “‘there is
not enough information available to study
the relationships of Oligosarcus Gunther and
Acestrorhynchus Eigenmann [the genera of
the Acestrorhynchinae] with [other] charac-
iform genera.’ Furthermore those authors
also noted that the ““Acestrorhynchinae may
not be a monophyletic group.” The re-
maining tribe, the Characini, is also of
questionable monophyly. Menezes (1976)
considered the genera Cynopotamus, Gal-
eocharax and Acestrocephalus to be a
separate monophyletic subfamily, the Cy-
nopotaminae, but now Menezes (pers. com-
mun.) informs us that this separation may
no longer be tenable in light of a more de-
tailed phylogenetic analysis.
On the basis of preliminary observations,
Menezes (in Sazima 1983:88) suggested that
some of the remaining Characinae and
Characini of Géry are polyphyletic, with
Exodon Miller and Troschel being more
closely related to members of the Tetrago-
nopterinae than to the remaining genera of
the Characini. Sazima (1983:88) noted that
his behavioral evidence appeared to con-
firm Menezes’ concepts. Vari (1986:332,
333) discussed the tentative relationships of
his new genus and species, Serrabrycon ma-
goi Vari. He noted that although he placed
650 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Table 2.— Number of teeth on jaw bones and number of branched anal-fin rays in Priocharax ariel, P. pygmaeus,
and genera of the Characinae discussed in text. Dentary tooth information for Lonchogenys and dentary and
premaxillary tooth counts for Acanthocharax presented as range of outer tooth row counts followed by range of
inner tooth row counts. Tooth data for Lonchogenys based on two cleared and stained specimens, 40.1—45.2
mm SL (USNM 270232) and anal-fin ray counts on 12 specimens (USNM 270230, 270231, 270232). Tooth
data and anal-fin ray counts for Heterocharax based on 7 cleared and stained specimens, 21.7-34.0 mm SL
(USNM 278994) and for Acanthocharax on | paratype (USNM 66109).
Priocharax
ariel pygmaeus Lonchogenys Heterocharax Acanthocharax
dheeth
Premaxillary 22-34 19-24 12-16 8-11 6/10
Maxillary 38-58 27-41 42-50 20-24 60
Dentary 38-55 28-36 16-18/11-14 33-38 4/33
Branched anal-fin rays 16-22 20-22 34-38 31-35 28
it in the Tetragonopterinae, its possession
of a mosaic of characters left open the pos-
sibility that the phylogenetic relationships
of Serrabrycon Vari could be with the Cha-
racinae. Serrabrycon magoi, if a characine,
is one with a short based anal-fin (15 or 16
rays posterior to the anterior undivided rays)
similar to that of Priocharax species. The
number of teeth on the maxilla, premaxilla
and dentaries of Serrabrycon are less than
those in Priocharax and the genera also dif-
fer in tooth forms. These differences do not,
of course, refute a hypothesis of a close phy-
logenetic relationship between the taxa. The
various uncertainties associated with taxa
such as Exodon and Serrabrycon together
with the lack of proposed synapomorphies
for possible members of Géry’s Characinae
complicate a consideration of the relation-
ships of Priocharax among Charax and pu-
tatively associated genera.
The numerous conic jaw teeth and elon-
gate maxilla of Priocharax suggest a possible
relationship with genera of the Characinae
and Cynopotaminae. Among the genera
listed at the beginning of this section, Acan-
thocharax, Acestrocephalus, Cynopotamus,
Galeocharax, Lonchogenys and Priocharax,
particularly P. ariel, have very high max-
illary and total dentary tooth counts (Tables
2, 3). Heterocharax and those genera with
the exception of Acestrocephalus have high
total dentary tooth counts (above 28) com-
pared to other Characinae. These high tooth
counts might be synapomorphies relating
these genera, but we prefer not to advance
a hypothesis of relationships based solely
on that one system.
One possible synapomorphy common to
at least Priocharax, Roeboides and Cyno-
potamus 1s the retention of a larval pectoral
fin anatomy at relatively large body sizes.
Many characids convert from the larval
pectoral fin anatomy to an essentially adult
form at about 6 to 9 mm SL (pers. obs.).
Priocharax, in contrast, retains the larval
fin to at least 17 mm SL and at least one
undetermined species of Roeboides (USNM
279251) retains that fin form up to 26 mm
SL. Cynopotamus atratoensis (Eigenmann)
hasa larval pectoral fin in specimens as large
as 41 mm SL(CAS-IUM 15033, paratypes).
The distribution of this feature in most gen-
era of the Characinae and Cynopotaminae
is unknown. Thus it would be inappropriate
to use it aS a synapomorphy for putative
subgroups in those subfamilies until its phy-
letic distribution has been thoroughly ana-
lyzed.
The possession of a larval pectoral fin has
also been reported within the Characidae
by Durbin (1909:55) for the tetragonopterid
Dermatocheir catablepta Durbin. That
species, since moved to Hyphessobrycon by
VOLUME 100, NUMBER 3 651
Table 3.—Number of teeth on jaw bones and number of branched anal-fin rays in Priocharax ariel, P. pygmaeus,
and genera of the Cynopotaminae discussed in text. Dentary tooth information for genera of the Cynopotaminae
presented as range of outer tooth row counts followed by range of inner tooth row counts.
Priocharax
ariel pygmaeus Acestrocephalus Cynopotamus Galeocharax
Teeth
Premaxilla 22-34 19-24 9-13 8-12 8-12
Maxilla 38-58 27-41 30-44 42-60 36-54
Dentary 38-55 28-36 3/9-13 2—4/22-35 3-4/7-11
Branched anal-fin rays 16-22 20-22 29-36 36-53 36-45
Geéry (1977:462), is known only from an 18
mm holotype described as having the “‘Pec-
toral short and paddle-shaped with a fringe
of soft rays” (Durbin 1909:56). Although
sharing an evidently very similar form of
pectoral fin with the species of Priocharax,
H. catableptus differs from Priocharax and
the species of the Characinae and Cyno-
potaminae in the form and distribution of
the oral teeth. Although Gery’s alignment
of H. catableptus with other Hyphessobry-
con species appears reasonable, the question
of the phylogenetic relationships of the
species, and the significance of the retention
of the larval form of pectoral fin, requires
further study once additional material of the
species becomes available.
A variety of apparently derived charac-
ters listed in “keys” by Géry (1977:302-
330) such as a notch and spine along the
ventral margin of the cleithrum, a sharp
ventral preopercular angle or spine, en-
larged coracoid bones forming a keel along
the ventral border of the cleithrum, ctenoid
body scales, outwardly projecting mam-
milliform teeth (discussed by Sazima and
Machado 1982), and certain body shape
configurations (e.g., a dorsal hump in the
body profile anterior to the dorsal-fin origin)
occur in various subunits of the Characinae
or Cynopotaminae, or both. Although these
characters have been used in part to diag-
nose genera and species groups, the current
lack of cladistic phylogenetic analyses of
these subfamilies prevents an evaluation of
the usefulness of these features in diagnos-
ing monophyletic groups at this time.
Acknowledgments
The specimens that served as the basis
for the description of Priocharax ariel were
collected during a joint MBUCV-USNM
expedition made possible by funding from
the Scholarly Studies Program of the Smith-
sonian Institution. Sven Kullander and Eric
Ahlander (NRM) made available the type
series of Priocharax pygmaeus. Research as-
sociated with this project was supported in
part by the I.E.S.P. Neotropical Lowland
Research Project of the Smithsonian Insti-
tution. Antonio Machado-Allison, Carl J.
Ferraris, Jr., Otto Castillo and Justa M. Fer-
nandez assisted in collecting activities in
Venezuela. Assistance on and/or informa-
tion for this study was provided by Marilyn
J. Weitzman, Lynn Norrod, Ann M. Wil-
liams, Ricardo M. C. Castro and Andrew
G. Gerberich (USNM) and especially Naér-
cio Menezes (MZUSP). This paper was 1m-
proved by the comments and suggestions of
Ricardo M. C. Castro, Carl J. Ferraris, Jr.,
Sara V. Fink, William L. Fink, Naércio A.
Menezes, Marilyn Weitzman, and two
anonymous reviewers. Drawings were pre-
pared by Sara V. Fink.
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de Zoologia de Sao Paulo 7:35-56.
Vari, R. P. 1986. Serrabrycon magoi, a new genus
and species of scale-eating characid (Pisces:
Characiformes) from the upper Rio Negro.—
Proceedings of the Biological Society of Wash-
ington 99(2):328-334.
Weitzman, S. H. 1986. A new species of Elacho-
charax (Teleostei: Characidae) from the Rio Ne-
gro region of Venezuela and Brazil.—Proceed-
ings of the Biological Society of Washington
99(4):737-745.
, and S. V. Fink. 1985. Xenurobryconin phy-
logeny and putative pheromone pumps in glan-
dulocaudine fishes (Teleostei: Characidae).—
Smithsonian Contributions to Zoology 421:1-
IA
, and W. L. Fink. 1983. Relationships of
the neon tetras, a group of South American
freshwater fishes (Teleostei, Characidae), with
comments on the phylogeny of New World
Characiforms.— Bulletin of the Museum of
Comparative Zoology 150(6):339-395.
Department of Vertebrate Zoology (Fish-
es), National Museum of Natural History,
Smithsonian Institution, Washington, D.C.
20560.
PROC. BIOL. SOC. WASH.
100(3), 1987, pp. 653-658
HETEROM YSIS KENSLEYI AND H. CORALINA,
NEW SPECIES FROM THE SHALLOW WATERS
OFF LOOE KEY, FLORIDA
(MYSIDACEA: HETEROMYSINI)
Richard F. Modlin
Abstract. — Descriptions of Heteromysis kensleyi and H. coralina, new species
from coral reef habitats off Looe Key, Florida, are presented.
Species in the genus Heteromysis appear
to show considerable adaptive radiation in
shallow waters of the Caribbean Sea and
Gulf of Mexico (Modlin 1983). About 22
of these species are known to inhabit these
waters. Eight occur in the waters off Florida.
Brattegard (1969, 1970) reported collecting
specimens of H. formosa in the vicinity of
Ft. Pierce, H. floridensis in Biscayne Bay,
and H. nouveli and H. dispar in the Florida
Keys. Heteromysis beetoni, H. guitarti, H.
filitelsona, and H. hopkinsi inhabit the Flor-
ida Middle Ground coral reef system north-
west of Tampa, Florida, in the Gulf of Mex-
ico (Modlin 1984). This paper describes two
new species of Heteromysis from the waters
off Looe Key, Florida.
Heteromysis kensleyi, new species
Fig. 1A-K
Material examined. —Male, 8.1 mm, ho-
lotype (USNM 229527); 2 males (6.0, 7.9
mm), 2 females (6.5, 7.7 mm), 3 ovigerous
(6.4—-8.5 mm), paratypes (USNM 229528)
collected 27 Jan 1983, from upper spur and
groove, vertical buttress wall with hard and
soft corals, depth 6.1 m, sta FLK-24.— Male,
6.1 mm, collected 27 Jan 1983, from coral
rubble at base of upper spur and groove
buttress, depth 6.1 m, sta FLK-25.—Ovig-
erous female, collected 28 Sep 1982, from
buttress wall region of spur and groove,
depth 6.1 m, sta FLK-12, specimen dis-
sected. Specimens were hand-collected by
B. Kensley and M. Schotte at collecting sites
previously poisoned.
Description. —Body large, robust. Cara-
pace with anterior margin produced into tri-
angular rostrum, posterior margin emargin-
ate, exposing thoracic segments 7 and 8,
postero- and anterolateral lobes rounded.
Eyes large, oval, stalked; cornea rounded,
dorsal margin scalloped, with prominent
ocular tooth on anterosuperior edge.
Antennular peduncle 3-segmented; seg-
ment | about 1.1 times longer than segment
3; segment 2 compressed, with short spine
and plumose seta distomedially; segment 3
with 3 short simple setae on medial edge,
robust spine, inconspicuously flagellated,
about 0.7 times length of medial margin of
segment 3 (flagellum visible only with aid
of very high magnification) and three small
setae distomedially, midventral male lobe
prominent with less than 10 hair-like setae.
Antennal peduncle 3-segmented, about
1.7 times longer than scale; segment | in-
conspicuous; segment 2 about 1.6 times
longer than segment 3, one long and one
short naked seta, and one long plumose seta
distomedially; segment 3 with 3 naked setae
distomedially. Antennal scale blade-like,
medial margin strongly convex, lateral mar-
gin slightly convex, setose all around, apical
tip about 0.06 times scale length.
Right and left mandibles with blade-like
incisors, right lacinia mobilis saddle-like
with 4 tall cusps, left pedestal-like with 4
low cusps; each mandible with 3 robust se-
654 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
A 0.33 mm
8-C,G,J,K 0.10 mm
[25 3 fi, | 0,05 mm
Fig. 1. Heteromysis kensleyi, new species: A, Carapace; B, Antennular peduncle; C, Antennal peduncle and
scale; D, Mandibular palp; E, Left mandible; F, Right mandible; G, Thoracic endopod 3; H, Pleopod 3; I,
Pleopod 4; J, Uropod; K, Telson. A = male, 8.1 mm; B, H, I = male, 7.9 mm; C-—G, J, K = female, 6.5 mm.
VOLUME 100, NUMBER 3
tose accessory blades; right molar surface
rugose distally, left plate-like along entire
length. Mandibular palp 3-segmented; seg-
ment | inconspicuous; segment 2 expanded,
medial margin with 7 setae proximally, lat-
eral margin with 17-19 setae along entire
length; segment 3 about 0.4 times length of
segment 2, medial margin slightly sinuous,
proximal half with 7 naked setae, lateral
margin with 8 short spined setae and 6-7
tubercles distally, distal tip with long plu-
mose seta and robust spined claw.
Labium, maxillule, maxilla, and parag-
naths typical of genus.
Thoracic endopod 1 and 2 typical of ge-
nus. Thoracic endopod 3-merus, about 1.2
times longer than carpopropodus, medial
margin with 2 short simple setae proxi-
mally, one long and 3 short simple setae in
group distally; medial margin of carpopro-
podus with 8 robust, subequal flagellated
spines arranged in pairs, anterior edges of
center 2 pairs serrate, one long simple seta
submarginal to each anterior pair of spines,
3 short simple setae anterior to spinal group,
lateral margin with 2 simple setae in prox-
imal half, 2 groups of 3 short simple setae
distally; dactylus conspicuous with 2 simple
setae proximally. Carpopropodus of tho-
racic endopod 4 with 3 segments; carpo-
propodus of thoracic endopods 5-8 with 8
segments. Thoracic exopods 2-8 with 10
segments.
Pleopods unsegmented; male pleopods 1,
2, and 5 not differentiated. Pleopod 3 with
13 long naked setae on anterior surface, 7
flagellated spines on distal margin, one long
plumose seta distolaterally, 2 short plumose
setae on lateral margin proximally, 2 short
naked setae on medial margin, pseudobran-
chial lobe with one short and 4 long plumose
setae. Pleopod 4 with 11-13 long naked se-
tae on anterior surface, 7—8 flagellated spines
on distal margin, one long plumose seta dis-
tomedially, 2 short plumose setae on lateral
margin proximally, 1—2 short simple setae
on medial margin, pseudobranchial lobe
655
with one short and 4 long plumose setae.
Female pleopods rudimentary.
Uropod exopod about 1.1 times longer
than endopod, lateral margin straight, me-
dial margin slightly convex, setose all
around. Endopod linguiform with 5 robust
spines on medial margin in region of stato-
cyst, entire margin setose.
Telson about 0.8 times as long as exopod
of uropod, lateral margins slightly concave,
completely spined with 19-20 marginal
spines (apical spines included), outer apical
spine about 1.7 times longer than inner, cleft
0.15 times the length of the telson, com-
pletely spined with 11 spines.
Remarks. —Characteristics of H. kensleyi
place it near H. bredini (Brattegard, 1970)
and H. tuberculospina (Modlin, 1987). It
differs from both by having a very long in-
conspicuously flagellated spine on the dis-
tomedial edge of segment 3 of the antennule
peduncle. On H. bredini and H. tuberculo-
spina this spine is less than 0.2 times the
length of the medial margin of antennule
peduncle segment 3 rather than 0.7 times.
Carpopropodus of thoracic endopod 3 of H.
kensleyi has eight flagellated spines arranged
in four pairs, while that of H. bredini and
H. tuberculospina has, respectively, ten and
seven. Spines in the telsonal cleft of H. kens-
leyi, H. tuberculospina, and H. bredini num-
ber, respectively, 11,20 and 31. Mandibular
structure of H. kensleyi varies distinctly from
that of H. tuberculospina. Distal margins of
male H. kensleyi pleopods 3 and 4 each have
seven distal spines, while those of H. tub-
erculospina have ten each. Mandibles and
male pleopods of H. bredini have not been
described.
Ecological Notes. —Ovigerous females
carried 2—4 eggs or larvae. Specimens of /.
dispar Brattegard and H. coralina, new
species, were collected with H. kensleyi.
Etymology. —Named for Dr. Brian Kens-
ley, U.S. National Museum of Natural His-
tory, who kindly provided me with his Looe
Key mysid collection and who has done
656 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
SRY
AAKRAK
©
wy
Fig. 2. Heteromysis coralina, new species: A, Carapace; B, Antennular peduncle; C, Antennal peduncle and
scale; D, Mandibular palp, E, Left mandible; F, Right mandible; G, Thoracic endopod 3; H, Pleopod 3; I,
Pleopod 4; J, Uropod; K, Telson. A-K = male, 4.9 mm.
VOLUME 100, NUMBER 3
much to forward systematical and ecologi-
cal knowledge of the Caribbean Sea Isop-
oda.
Heteromysis coralina, new species
Fig. 2A-—K
Material examined.—Female, 5.2 mm,
holotype (USNM 229529); 4.9 mm male
(dissected on 6 microscope slides), 4.3 mm
female, 3.1 mm juvenile, paratypes (USNM
229530); and 5.0 mm male (badly damaged)
collected 27 Jan 1983 from upper spur and
groove, vertical buttress wall with hard and
soft corals, depth 6.1 m, hand collected at
station FLK-24 in area previously poi-
soned.
Description. —Body small, robust. Cara-
pace with anterior margin produced into tri-
angular rostrum, posterior margin emargin-
ate exposing thoracic segments 7 and 8,
anterolateral lobes rounded, posterolateral
lobes angular. Eyes large, stalked, oval with
posteromedial region produced and covered
with regularly arranged minute denticles;
cornea small, oval, prominent ocular tooth
on anteromedial edge. .
Antennular peduncle 3-segmented; seg-
ment | as large as segment 3 with distinct
lateral process having 4 plumose setae on
apex, simple seta distomedially; segment 2
compressed; segment 3 with 2 plumose se-
tae on medial margin; anterior margin with
2 long plumose setae and a strong blade-
like spine medially with inconspicuous fla-
gellum (flagellum visible only with aid of
very high magnification), male lobe with few
hair-like setae ventrally.
Antennal scale blade-like, apical segment
about 0.05 times length of scale, setose all
around. Antennal peduncle 3-segmented, 1.1
times length of scale, segment 1 small, in-
conspicuous; segment 2 with one long and
2 small simple setae, and one robust spined
seta distomedially, one minute plumose seta
distolaterally; segment 3 about 0.7 times
length of segment 2, one long and 2 short
657
simple setae distomedially, 3 minute plu-
mose setae on lateral margin distally.
Mandibles and palp: right and left man-
dibles with blade-like incisors, right incisor
with prominent anterior cusp, left incisor
with 2 anterior cusps; right lacinia mobilis,
small, with serrate edges and 2 prominent
spike-like cusps, left with 4 cusps; each
mandible with 3 strong setose accessory
blades; right molar surface strongly rugose
posteriorly with setose clumps at anterior
and posterior edges; left molar surface, an-
terior half plate-like, posterior half with mi-
nute rugae, setose clump at posterior edge.
Mandibular palp 3-segmented; segment 1
small, inconspicuous; segment 2 expanded,
anterior margin with 8 simple setae, lateral
margin with 1 1—12 simple setae and 2 distal
spined setae; segment 3 with 3 long simple
setae near anterior margin, posterior margin
sinuous, proximal half with one spined seta,
distal half with 12-14 spined setae and 2
long spined claws on apex.
Labium, maxillule, maxilla and parag-
naths typical of genus.
Thoracic endopods | and 2 typical of ge-
nus. Thoracic endopod 3 lateral margin of
merus with 2 small simple setae proximally
and | simple seta distally, medial margin
with 5 long simple setae interspersed with
1-2 small simple setae; medial margin of
carpopropodus with 9 robust flagellated
spines, 8 arranged in pairs with single spine
proximally, 3 small and 2 long setae distally,
lateral margin with 3 short simple setae
proximally and 3 longer simple setae dis-
tally; dactylus with 4 minute setae distally
and prominent claw. Thoracic endopod 4
with 3-segmented carpopropodus. Thoracic
endopods 5-8 with 6-segmented carpopro-
podus. Exopod 1 and 2 with 8 segments;
exopods 3-8 with 9 segments.
Pleopods unsegmented. Male pleopods 1,
2 and 5 not differentiated. Pleopod 3 with
10 long plumose setae on anterior surface,
distal margin with 8-11 flagellated spines
and one long plumose seta distolaterally,
658
lateral margin with 2 spined setae proxi-
mally and one minute flagellated seta dis-
tally, medial margin with one minute seta
proximally, pseudobranchial lobe with 3
long and 2 short plumose setae. Pleopod 4
with 10 long plumose setae on anterior sur-
face, distal margin with 14-15 flagellated
spines and one long plumose seta distolat-
erally, lateral margin with 2 spined setae
proximally and one minute plumose seta
distally, lateral margin with one spined seta,
pseudobranchial lobe with 3 long and 2 short
plumose setae. Female pleopods rudimen-
tary.
Uropods: exopod about 1.2 times longer
than endopod, lateral margin straight, me-
dial margin convex, setose all around; en-
dopod, linguiform, with 5 prominent spines
medially in region of statocyst, setose all
around.
Telson about 0.8 times length of exopod
of uropod, distal *4 of lateral margins each
with 10 spines (apical spines included) that
increase in length distally, outer apical spine
1.5 times longer than inner; telsonal cleft
with 19-20 small spines concentrated in
proximal half, 0.3 times length of telson.
Remarks. —Heteromysis coralina ap-
pears closely related to H. beetoni, but it
differs by having five rather than nine spines
in the region of the statocyst on the endopod
of the uropod; 19-20 spinules in the prox-
imal half of the telsonal cleft rather than
15-17; nine flagellated spines on the car-
popropodus of the thoracic endopod 3 rath-
er than eight; 9-11 and 14-15 flagellated
spines, respectively, on the distal margins
of male pleopods 3 and 4. Male pleopods 3
and 4 of H. beetoni have seven and 20, re-
spectively. Additionally, the design of the
mandibular surfaces of the two species differ
radically.
Ecological notes. —Specimens of H. dis-
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
par Brattegard and H. kensleyi occurred in
the same sample as H. coralina.
Etymology. —Named for the habitat from
which this species was collected.
Acknowledgments
I would like to extend my appreciation to
Dr. Brian Kensley for providing the Looe
Key mysid collection and to Dr. Thomas E.
Bowman for assisting in the publication of
this manuscript. Financial support for a trip
to the U.S. National Museum of Natural
History was provided in part by the Marine
Environmental Sciences Consortium, Dau-
phin Island, Alabama. This is MESC con-
tribution No. 134.
Literature Cited
Brattegard, T. 1969. Marine biological investigations
in the Bahamas 10. Mysidacea from shallow
water in the Bahamas and southern Florida. Part
1.—Sarsia 39:17-106.
. 1970. Mysidacea from shallow water in the
Caribbean Sea.—Sarsia 43:111-154.
Modlin, R. F. 1983. Zoogeography of the Hetero-
mysis (Mysidacea) in the Caribbean Sea and Gulf
of Mexico. Symposium on the origins and dis-
tribution of crustaceans in the Caribbean Sea
and Gulf of Mexico.—ASB Bulletin 30(2):72-
Bs
1984. Mysidacea from the Florida Middle
Ground, northeast Gulf of Mexico, with de-
scriptions of three new species of Heteromysis
and a key to the Heteromysini of the western
Atlantic. —Journal of Crustacean Biology 42(2):
278-297.
1987. Mysidacea from the shallow waters in
the vicinity of Carrie Bow Cay, Belize, Central
America, with descriptions of two new species. —
Journal of Crustacean Biology 7(1):106-121.
Department of Biological Sciences, The
University of Alabama in Huntsville,
Huntsville, Alabama 35899.
PROC. BIOL. SOC. WASH.
100(3), 1987, pp. 659-663
BAHALANA MAYANA, A NEW TROGLOBITIC
CIROLANID ISOPOD FROM COZUMEL ISLAND
AND THE YUCATAN PENINSULA, MEXICO
Thomas E. Bowman
Abstract. — Bahalana mayana is described from two anchialine caves, Cueva
Quebrada on the island of Cozumel, and Temple of Doom Cave on the adjacent
Yucatan Peninsula near Tulum. It differs from the two known species, both
from the Bahamas, B. geracei Carpenter from San Salvador Island, and B.
cardiopus Notenboom from Mayaguana Island, in the ventrally projecting clyp-
eus, pereopods 1-3 lacking the long processes on the merus and carpus, the
armament of pleopods 3-5, and the narrow exopod of the uropod.
The genus Bahalana was proposed by
Carpenter (1981) for an unusual troglobitic
cirolanid isopod, Bahalana geracei, from the
anchialine Lighthouse Cave on San Salva-
dor Island, Bahamas. About 50 pages later
in the same journal, a second very similar
species, B. cardiopus, was described by Not-
enboom (1981) from Mount Misery Cave,
Mayaguana Island, Bahamas. San Salvador
and Mayaguana, both in the Bahama ar-
chipelago, are separated by about 250 km.
Some 1300-1400 km to the southwest, a
third species of Bahalana, described below,
has now been discovered in anchialine caves
on the island of Cozumel and the adjacent
mainland of the Yucatan Peninsula near
Tulum, Quintana Roo, Mexico.
Bahalana mayana, new species
Figs. 1-2
Material. — Mexico, Quintana Roo, Co-
zumel Island, Cueva Quebrada, leg. Dennis
Williams and Jeffrey Bozanic, 19 Sep 1985,
127.9 mm, USNM 233292; 1 6 4.5 mm,
USNM 233300.—Leg. Dennis Williams, 13
Jun 1986, 3 35.9, 8.0, 8.2 mm, 1 28.5 mm,
1 juv. 3.7 mm; 14 Jun 1986, 1 6 10.0 mm
(holotype), USNM 233298; 16 Jun 1986, 1
6 ca. 9.4 mm (head detached), USNM
233293; 17 Jun 1986, 1 6 9.6 mm, 4 2 8.2,
8.3, 8.4, not measured (dissected, larger than
others), USNM 233295.—Tulum area,
Temple of Doom Cave, leg. Dennis Wil-
hams, 20 Jun 1986, | 6 6.6 mm, USNM
233296.—Leg. Thomas M. Iliffe, 10 Nov
1986, 136.4 mm, USNM 233299. All spec-
imens except the holotype are paratypes.
Etymology.—Named for the Maya, na-
tive people of the Yucatan region.
Diagnosis. — Bahalana mayana is distin-
guished from its two congeners by the acute-
ly produced clypeus, the long segment 2 of
antenna 1, the fewer flagellar segments in
antenna | and 2, the long subapical seta on
the mandibular palp segment 2, the short
segment 4 of the maxillipedal palp, the mor-
phology of pereopods 1-3, and the narrow
exopod of the uropod. The principal differ-
ences between the three species are set forth
in Table 1.
Habitats. — For the following information
on the caves inhabited by Bahalana ma-
yana I thank Dr. Thomas M. Iliffe and Ms.
Jill Yager.
Both Temple of Doom Cave and Cueva
Quebrada, as well as numerous other caves
in the region, were formed by the mixing of
fresh ground water with subterranean Ca-
ribbean seawater in a highly reactive geo-
chemical zone that produced enhanced car-
bonate dissolution (Back et al. 1986). Thus
=
cr
SS eo
wit ep ttt
geet [ aff
Fig. 1.
mandible; I, Lacinia of left mandible; J, Incisor of left mandible; K, 3rd segment of mandibular palp; L, Maxilla
merus and carpus, medial.
1, apex of exopod; M, Maxilla 1, endopod; N, Maxilla 2; O, Maxilliped; P, Pereopod 1, lateral; Q, Pereopod 1,
Bahalana mayana: A, Habitus, dorsal; B, Head, dorsal; C, Frontal lamina, clypeus, and labrum; D,
Clypeus, lateral; E, Posterior margin of telson; F, Antenna 1; G, Antenna 2, proximal segments; H, Right
Fig. 2. Bahalana mayana: A, Habitus, lateral; B, Pereopod 2; C, Pereopod 3; D, Pereopod 5; E, Pereopod
6; F, Pereopod 6, distal end of carpus; G, Pereopod 7; H—L, Pleopods 1-5, 2; M, Endopod of pleopod 2, 3, setae
omitted; N, Penes; O, Uropod, dorsal.
662
Table 1.—Characteristics of the species of Bahalana.
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Character P. geracei P. cardiopus P. mayana
Greatest length 15 mm 10 mm 10 mm
Clypeus Not produced Not produced Acutely produced
A-1 peduncle, longest Segment 3 Segment 3 Segment 2
segment
A-1 flagellum segments m3} 19-22 1
A-2 flagellum segments 40-45 CA, 32 ca. 22
Md palp segment 2
Mx-1 exopod apex
Mx-1 endopod apex
Mx-2 exopod apex
Mxp retinacula
Mxp palp, length of
segment 4/segment 3
P-1 merus
P-2-3 merus
Without long
subapical seta
11 spine + | seta
4 spines + | setule
8 setae
D
0.7
With long postero-
distal process
With long antero-
distal process
Without long
subapical seta
10 spines + 1 seta
4 spines + I setule
8 setae
l
0.8
With long postero-
distal process
With long antero-
distal process
With long subapical
seta
11 spines + 1 seta
2 spines + 2 setules
6 setae
1
0.4
With rudimentary
posterodistal process
With rudimentary
anterodistal process
P-4-7 unguis Pectinate
P1-3-5, distal margins With a few setae
of endopods
Convex
’, width of endopod
Without setae
Up endopod distal margin
Up exopod width
Up rami lateral margins
most caves in the region have their primary
development at depths corresponding to the
halocline or mixing zone. The depth of the
halocline, and correspondingly the depth of
the caves, tends to increase with increasing
distance inland as the freshwater lens thick-
ens.
Cueva Quebrada, Chankanaab National
Park, Isla Cozumel, Quintana Roo, Mexico:
This cave is located on the east coast of the
island of Cozumel. It has five surface open-
ings and 2759 m of surveyed passage pos-
sibly making it Mexico’s longest completely
submerged cave. The deepest point in the
cave is only —12 m. Brackish water is dis-
charged in the form of reversing tidal cur-
rents from entrances along the coast. The
cirolanid was collected about 800 meters
from the opening, at a depth of about 5
meters. Salinity was 21%o; dissolved oxygen
3.0 ppm. Associated fauna include the am-
phipod Bahadzia sp., undetermined ther-
mosbaenaceans, and Procaris sp.
Pectinate Entire, with 2 basal spines
Unarmed With setae in P1-3-4,
unarmed in P1-5
Convex Slightly concave
’, width of endopod
Without setae
3 width of endopod
With setae
As reported by Bozanic (1984:14), ““Ma-
rine life was fairly abundant ... observed
were encrusting sponges and hydroids, lob-
sters, grunts, parrotfish, big eye, tarpon,
shrimp, and many species of small (less than
Ys, inch long) animals. As in the marine caves
of Grand Bahama Island, the sponges ex-
tended from the opening as far as I pene-
trated, the fish were seen primarily near the
entrances, and the smaller shrimp and crea-
tures in the naturally lightless zones of the
caves.”
Temple of Doom Cave, Tulum, Quintana
Roo, Mexico: This cenote is located about
15 km inland from the Caribbean Sea on
the east coast of the Yucatan Peninsula. The
entrance consists of an undercut circular hole
about 6 m in diameter with a 3 m drop to
the water. Underwater the cave radiates
down and outward on all sides from a cen-
tral collapse mound. From the base of the
mound, a white-walled passage extends to
a huge submerged chamber approximately
VOLUME 100, NUMBER 3
90 m long by 40 m wide and 10 m high.
Maximum water depth was 18 m, with pri-
mary passage development occurring in the
region of the halocline at —15 m. Salinities
at the surface and at —18 m were 3 and
14%o, respectively. Water temperature was
25°C in November 1986.
Most animals were observed just above
the halocline in oligohaline waters. Trog-
lobitic species were only found in more re-
mote areas of the cave, away from the nu-
merous and voracious small fish present near
the entrance. Collecting was done with a
plankton net and suction bottle from the
water column in 12 to 8 m depths using
scuba. In addition to Bahalana mayana,
specimens of the cirolanid Creaseriella anops
(Creaser), the mysid Antromysis (Antro-
mysis) cenotensis Creaser, amphipods, and
shrimp were collected.
Acknowledgments
Once again I am indebted to skilful and
resolute cave divers for the collection of re-
markable isopods. For their successful ef-
forts my sincere thanks go to Jeffrey Boz-
anic, Thomas M. Iliffe, and Dennis
Williams. Collecting by Dr. Iliffe was sup-
663
ported by National Science Foundation
grant BSR-8417494, and he was assisted by
Dinah Drago, Juan Jose Fucat, Michael
Madden, and James Coke; logistical and
collecting aid was provided by Dr. John
Markham. This is Contribution No. 1119
of the Bermuda Biological Station for Re-
search.
Literature Cited
Back, W., B. B. Hanshaw, J. S. Herman, and J. N. Van
Driel. 1986. Differential dissolution ofa Pleis-
tocene reef in the ground-water mixing zone of
coastal Yucatan, Mexico.— Geology 14:137-140.
Bozanic, J. 1984. Caves of Cozumel.— Underwater
Speleology 11(6B):12-14.
Carpenter, J. H. 1981. Bahalana geracei n. gen., n.
sp., a troglobitic marine cirolanid isopod from
Lighthouse Cave, San Salvador Island, Baha-
mas.—Bijdragen tot de Dierkunde 51(2):259-
AGT.
Notenboom, J. 1981. Some new hypogean cirolanid
isopod crustaceans from Haiti and Mayaguana
(Bahamas).— Bijdragen tot de Dierkunde 51(2):
313-331.
Department of Invertebrate Zoology, Na-
tional Museum of Natural History, Smith-
sonian Institution, Washington, D.C.,
20560.
PROC. BIOL. SOC. WASH.
100(3), 1987, pp. 664-673
FRESHWATER TRICLADS (TURBELLARIA) OF
NORTH AMERICA. XVI.
MORE ON SUBTERRANEAN SPECIES OF
PHAGOCATA OF THE EASTERN UNITED STATES
Roman Kenk
Abstract.— Three new species of Phagocata are described, P. pygmaea and
P. spuria from caves in North Carolina, and P. notorchis from a seep in eastern
Tennessee. Additional distributional records are given for P. carolinensis and
P. procera.
The species discussed in this paper were
collected by several investigators, mainly
speleologists, in the course of several years.
In order to obtain an accurate picture of the
shape of living planarians, which can hardly
be maintained by any known method of
preservation, I had asked my collaborators
to send me live specimens. Another good
reason to insist on living specimens is the
fact that many flatworms are collected in
the immature state, lacking their reproduc-
tive systems which form the basis of their
taxonomic standing. They may be either ju-
veniles that have not yet achieved sexual
maturity, or specimens that have been sub-
jected to prolonged starvation for the lack
of proper food. It is known that planarians
may sustain starvation for several months
without detriment. They gradually reduce
some parts of their anatomy, decrease in
size, and cannot be distinguished from ju-
veniles. It is highly probable that many of
the immature planarians collected in the
field, particularly those inhabiting subter-
ranean waters, are not true juveniles but
specimens that have been kept from full de-
velopment or have reduced in size and
structure on account of deficient nutritive
conditions. These immature specimens may
be raised to maturity in the laboratory by
being kept in aquaria under proper temper-
ature conditions and being fed once or twice
a week. Species of Phagocata accept beef
liver as food and mature after several months
in the culture.
Methods
Mature specimens were preserved by kill-
ing well extended worms with a hot aqueous
solution of mercuric chloride (HgCl,), acid-
ulated with acetic acid after the killing. Se-
rial sections of 6-8 mm thickness were
stained with Ehrlich’s acid hematoxylin and
counterstained with Eosin-Phloxine B.
The type specimens of the new species
have been deposited in the National Mu-
seum of Natural History (formerly United
States National Museum, USNM), Smith-
sonian Institution, Washington, D.C.
Phagocata notorchis, new species
Figs. 1A, B, 2, 3
Type material. — Holotype: posterior part
of worm, sagittal sections on 4 slides (USNM
102760). Paratypes: sagittal sections of en-
tire animal, 7 slides (USNM 102761); sag-
ittal sections of posterior part, 4 slides
(USNM 102762); transverse sections of
posterior part, 7 slides (USNM 102763).
External features. (Fig. 1A, B).—This
white, blind, very slender species reaches a
considerable size, up to a length of 27 mm
and a width of 2 mm when quietly gliding.
The anterior end is truncate, with a slightly
convex frontal margin and a pair of slender,
VOLUME 100, NUMBER 3
A | aan Dicecual
Fig. 1. Drawings and photographs of living speci-
mens. A and B, Phagocata notorchis; C and D, P. pyg-
maea; E, P. morgan.
665
pointed auricles projecting anterolaterally
and held elevated when in gliding motion.
Behind the auricles the body margins widen,
then run parallel for the greater part of the
body length, to narrow again near the blunt-
ly pointed posterior end. The pharynx is
situated in the posterior third of the body.
Thus the species externally resembles two
other white Phagocata species of the Ap-
palachian region that have prominent au-
ricles, P. bursaperforata Darlington and P.
procera Kenk, from which it is clearly sep-
arated by its anatomical characters.
Anatomy.—The branching of the intes-
tine could not be analyzed in the living spec-
imens. There are very many, certainly over
20, lateral branches on either side of the
anterior intestinal trunk, and perhaps an
equal number on each posterior trunk.
in 0.7 mm-~
Fig. 2. _Phagocata notorchis, sagittal section through anterior region. in, intestine; fe, testis, ve, vas efferens.
666
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
ro) vd vs
Fig. 3.
gp ac va
am odc
Phagocata notorchis, semidiagrammatic view of copulatory apparatus in sagittal section. ac, common
atrium; am, male atrium; b, copulatory bursa; gp, gonopore; odc, common oviduct; p/, penis lumen; va, vagina;
vd, vas deferens; vs, seminal vesicle.
In the reproductive system, the ovaries
are located behind the third to fifth pair of
intestinal branches. The testes (Fig. 2) are
predominantly subdorsal, situated above
and between the intestinal branches. The
follicles are not directly attached to the thin
anterior vas deferens, as they are in many
other species of the genus, but connect with
it by individual efferent ductules (ve). The
testicular zone on either side begins at a
considerable distance posterior to the ova-
ries and extends to about the level of the
pharynx root.
In the copulatory apparatus (Fig. 3), the
gonopore leads into a small common atrium
that connects anteriorly with the male atrium
and posteriorly with the outlet of the cop-
ulatory bursa. The penis consists of a small
bulb and a plug-shaped papilla. Its lumen
is divided into an anterior seminal vesicle
(vs) situated in the bulb and a large cavity
(p/) that would correspond to an ejaculatory
duct. The vesicle is lined by a thick epithe-
lium of tall, apparently secretory, cells. In
sagittal sections it appears as a dorsoven-
trally oriented duct, but extends also later-
ally in the bulb, so as to represent an antero-
posteriorly compressed cavity. Ventrally it
opens into the large cavity in the penis pa-
pilla, which is lined by a cuboidal epithe-
lium and surrounded by a strong muscular
layer. The walls of this cavity are variously
folded and the cavity opens into the male
atrium in a place dependent on the state of
contraction of the muscular systems of the
cavity and the external wall of the papilla.
The two vasa deferentia, after expanding
into the usual spermiductal vesicles, enter
the penis bulb laterally and open separately
into the dorsal portion of the seminal ves-
icle.
The two oviducts or ovovitelloducts unite
in the space above the atrium and the short
common oviduct (odc), equipped with eo-
sinophilic shell glands, opens into the com-
mon atrium (ac) from the dorsal side. The
copulatory bursa (b), a cavity with irregular
outline, is located a short distance posterior
to the pharyngeal pouch. Its outlet, the bur-
sal duct, runs posteriorly somewhat to the
left of the midline, and is divided into two
different section. The anterior section is a
narrow canal proceeding along the male
atrium, surrounded by a moderate muscle
layer. The posterior section or vagina (va),
however, is greatly expanded and curves
ventrally to open into the common atrium.
It has a very thick muscular layer composed
of intermingled fibers. There is no com-
munication between the bursa and the in-
testine.
Distribution and ecology.—The species
was collected by J. J. Lewis and T. Everitt
on 7 May 1977 in Great Smoky Mountains
National Park, in a seep near the trail to
Alum Cave Bluffs, Sevier County, Tennes-
VOLUME 100, NUMBER 3
see, at an altitude of about 4900 feet (circa
1500 meters). About 20 specimens were sent
to me alive, the majority of them immature.
From its blindness and occurrence in a seep
we may consider the species to be a ground-
water inhabitant.
Taxonomic position.—The species be-
longs to a group of Phagocata widely dis-
tributed in the Appalachian region and
apparently related to the subgenus Afrio-
planaria. Its most outstanding characteris-
tic is the presence of a large vagina with an
extraordinarily developed muscle coat. A
large vagina, but without excessive mus-
culature, is seen also in P. procera. The dor-
sal position of the testes is also a good char-
acter, that is approached also in P.
bursaperforata.
Etymology.—The species name, notor-
chis (Greek noton, dorsum; orchis, testis)
refers to the subdorsal position of the tes-
ticular follicles.
Phagocata procera Kenk
Phagocata procera was described by Kenk
(1984) from Cat Den Cave in Jackson
County, North Carolina, but is more widely
distributed in caves and springs in that state.
The characteristic features of the species,
apart from the presence of elongated point-
ed auricles and ventral prepharyngeal tes-
ticles, are in the differentiation of the cop-
ulatory apparatus. The penis lumen consists
of two sections with histologically different
linings, an anterior “‘seminal vesicle”’ with
a tall, glandular epithelium and a posterior
“ejaculatory duct’ lined by a nonglandular
cuboidal epithelium. Both sections may vary
considerably in their appearance in pre-
served specimens, due to muscular con-
tractions and distortions. The anterior sec-
tion receives many faintly cyanophilic gland
ducts and may appear as a cavity or a con-
voluted duct. The posterior section may
have the aspect of a canal or of a wide cavity
of irregular outline, opening on the dorsal
side or at the tip of the penis papilla. The
667
bursal duct widens gradually as it proceeds
posteriorly and acquires a strong muscle coat
to form a vagina, without altering the his-
tological appearance of its epithelial lining.
Additional distributional records, all in
North Carolina:
Burke County: Flatworm Fissure, located in
Linville Gorge Wilderness Area, 13 May
1984, six immature specimens collected
in two small pools by Cato O. Holler, Jr.
and Christopher Holler and shipped to
me alive; they matured in a laboratory
culture.—Fifty/Fifty Fissure, in Linville
Gorge Wilderness Area, near Linville
Falls, 23 Sep 1984, 5 specimens collected
by Cato O. Holler and his family and Lee
James in a small seep; shipped to me alive.
McDowell County: Lake Tahoma Cave, lo-
cated on the road up Little Buck Creek
near Lake Tahoma, just outside of Mar-
ion, 4 Jan 1985, 4 specimens collected by
Cato O. Holler, Jr. and Christopher Hol-
ler in a small seep and shipped alive.
Mitchell County: Buckshot Cave, 9 Sep
1979, in a seep in the cave 5 specimens
collected by Cato O. Holler, Jr. and
shipped to me alive.
Yancey County: Seep on State Road 128,
1.5 miles north of Blue Ridge Parkway,
at about 6000 feet (circa 1830 m) altitude,
5 Jun 1972, 13 specimens collected by
Leslie Hubricht and shipped preserved. —
Seep on State Road 128, 10 Nov 1979, 3
specimens collected by Cato O. Holler,
Jr. and Cathy Topping and sent alive.—
Mt. Mitchell, Sep 1975, many specimens
collected by Nicole Gourbault and Ian R.
Ball and brought to me alive.
Phagocata pygmaea, new species
Rigs: (CD45
Type material. —Holotype: set of sagittal
sections on 4 slides (USNM 102764). Para-
type: sagittal sections on 3 slides (USNM
102765).
External features (Fig. 1C, D).—This is a
small and slender white species, externally
668
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig. 4. Phagocata pygmaea, sagittal section through anterior region. fe, testis; vd, vas deferens.
resembling several other species of the ge-
nus in the same general area. Mature spec-
imens are about 10.5 mm long and 0.8 mm
wide. The anterior end is truncate, with a
slightly convex central part of the frontal
margin and rounded lateral edges somewhat
protruding laterally, lacking pointed auri-
cles. There is an insignificant neck-like con-
striction, posterior to which the body wid-
ens again, the lateral margins remaining
parallel for the greater part of the body
length, to narrow again behind the level of
the copulatory complex and to meet at the
rather rounded posterior end. There are two
small eyes (diameter of the pigment cups
about 23 wm), situated very close together
at a level rather far removed from the fron-
tal margin of the head. The pharynx is in-
serted behind the middle of the body and
measures about 4% the body length. The
number of branches on the intestinal trunks
could not be counted, but is certainly very
large.
Anatomy.—The ovaries, equipped with
parovaria, are situated behind the second
or third lateral branches of the intestinal
trunk. The numerous testes (Fig. 4) are pre-
pharyngeal and are attached directly to the
ventral sperm ducts or vasa deferentia. They
must be considered to be essentially ventral,
although many of the large follicles may ex-
tend between the intestinal branches far
dorsally, occupying almost the entire dor-
soventral diameter of the body.
In the copulatory apparatus (Fig. 5), that
is located some distance behind the pha-
ryngeal pouch, the gonopore leads into a
small common atrium that connects ante-
riorly with the male atrium (am) and dor-
sally with the outlet of the bursal duct. The
penis consists of a small, but highly mus-
cular, bulb and a finger-shaped papilla. The
penis lumen (p/) is-a uniform, rather wide
cavity, not divided into a seminal vesicle
and an ejaculatory duct. It is confined main-
ly to the penis papilla and opens at the tip
of the papilla. Its wall forms a number of
villus-like projections. The vasa deferentia,
that in their posterior parts form the en-
larged and convoluted spermiductal vesi-
cles filled with sperm, ascend dorsally at the
level of the penis bulb, enter the bulb lat-
VOLUME 100, NUMBER 3
b vd
igs)
0.1 mm
669
| | \
Pl am odc gp ac bd
Phagocata pygmaea, copulatory apparatus in sagittal section. ac, common atrium; am, male atrium;
b, copulatory bursa; bd, bursal duct; gp, gonopore; odc, common oviduct; p/, penis lumen; vd, vas deferens.
erally, then curve posteriorly to open sep-
arately into the anterior end of the penis
lumen. The two oviducts unite in the space
between the male atrium and the bursal duct.
The common oviduct thus formed opens
from the dorsal side into the atrial cavity at
the border between the male atrium (am)
and the common atrium. The copulatory
bursa is a rounded cavity without any spe-
cial modifications. Its outlet, the bursal duct,
is divided histologically into two sections.
The anterior section is a straight narrow duct
lined with a ciliated cuboidal epithelium.
The posterior section is widened and some-
what convoluted and bends ventrally to
reach the common atrium. Its lining epi-
thelium consists of apparently secretory
(apocrine) cells without cilia. Both sections
are surrounded by moderate layers of mus-
cular fibers.
Distribution and ecology. —Phagocata
pygmaea was collected in Turtle Shell Cave,
Stokes County, North Carolina, located at
the base of Moore’s Wall near Hanging Rock
State Park, between Danbury and Moore’s
Springs. Seven immature specimens were
taken by Cato O. Holler and Christopher
Holler on 20 Nov 1983 and shipped to me
alive. Some matured in the laboratory in a
culture kept at 14°C and fed beef liver at
weekly intervals.
Taxonomic position. —The species 1s
closely related to the other subterranean
species of Phagocata of the Appalachian
area. Its outstanding characteristics are in
the anatomy of the copulatory apparatus,
mainly the structure of the penis and the
differentiation of the bursal duct.
Etymology. —The name pygmaea (Latin,
dwarfish) refers to the small size of the
species.
Phagocata spuria, new species
Figs. 6, 7
Type material. —Holotype: set of sagittal
sections on 4 slides (USNM 102766). Para-
types: sagittal sections of two specimens on
10 slides (USNM 102767, 102768).
External features. — This is a small, white,
two-eyed species resembling the common
sympatric Phagocata morgani morgani
(Stevens and Boring) to such an extent that
it was at first considered to be that species.
No sketches or photos were prepared of liv-
ing specimens (see Fig. 1E of P. morgani).
The anterior end is truncate, with a more
or less straight frontal margin and rounded
670
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig. 6. Phagocata spuria, sagittal section through anterior part of body. in, intestine; ov, ovary; fe, testis; vi,
vitellarium or yolk gland.
lateral edges. Eyes are two, of regular size,
placed close together and removed from the
frontal margin. The specimens were im-
mature when collected and were placed in
a laboratory culture where they matured af-
ter several months. The length of the pre-
served mature worms is 6 mm, which would
correspond to about 8-10 mm in life.
Anatomy. — Apart from the reproductive
system, the general anatomy shows no pe-
culiarities. The eyes are of normal size (di-
ameter of the pigment cup 57 um). The ova-
ries (Fig. 6) are hyperplastic, of enormous
size, and consist of amply lobed and
branched masses of darkly staining (cyano-
philic) cells. Histologically, not all these cells
are oocytes, but some, particularly in the
peripheral portions, are comparable to those
of parovaria. The yolk glands or vitellaria
are very abundant and fully developed, oc-
cupying all spaces of the mesenchyme, be-
ginning somewhat anterior to the ovaries
and extending to the posterior end. The nu-
merous testicular follicles are small, round,
and located strictly ventrally, below the in-
testinal branches. The testicular zone on
either side begins a short distance behind
the ovary and reaches posteriorly to about
the level of the mouth. In all three speci-
mens examined, the follicles contain no ripe
sperm cells, only immature spermatogonia.
The copulatory apparatus (Fig. 7) is rath-
er small and not very clearly differentiated.
The atrial cavity is undivided. The penis
has a small weakly muscular bulb and a
finger-shaped papilla. Its lumen consists of
a bulbar cavity (seminal vesicle) lined with
a tall glandular epithelium, and a narrow
duct (ejaculatory duct) with cuboidal or flat-
tened cellular lining, that runs through the
center of the papilla and opens slightly dor-
sally to its tip. The vasa deferentia, which
in other planarians appear widened, con-
voluted, and filled with sperm before en-
tering the penis, forming the spermiductal
for false seminal vesicles, are here empty
and only slightly widened. They enter the
penis bulb laterally and open separately into
the seminal vesicle. The outlet of the round-
ed copulatory bursa takes an arch-shaped
course, gradually widening in its posterior
section, without forming a histologically dif-
VOLUME 100, NUMBER 3
671
a
; r ) |
0.1mm 5b vd
Fig. 7.
|
l | Peerysanah
vs de gP odc bd
Phagocata spuria, copulatory apparatus in sagittal section. b, copulatory bursa; bd, bursal duct; de,
ejaculatory duct; gp, gonopore; odc, common oviduct; vd, vas deferens; vs, seminal vesicle.
ferent vagina. It opens into the atrial cavity
near the gonopore.
Distribution and ecology. —The species is
known only from Bennett’s Mill Cave in
McDowell County, North Carolina, located
just outside the town of Marion. Cato O.
Holler, Jr. collected 3 immature specimens
on 12 Jun 1977 and sent them to me alive.
They matured in the laboratory. No signs
of reproduction, either by fission or by co-
coons, were observed. Two specimens were
preserved in Dec 1977 and the third in Sep
1978.
Taxonomic position. —While the general
anatomical plan of the species places it
clearly in the genus Phagocata, the speci-
mens exhibit some characters rarely ob-
served in planarians. The full development
of the ovaries and yolk glands, together with
the incomplete appearance of the testes, may
seem to indicate that the female gonads ma-
ture at a time different from that of the male
gonads. This would not, however, explain
the remarkable hyperplasia of the ovaries
which are generally rather small rounded
organs adjoining the ventral nerve cords.
This hyperplastic condition 1s occasionally
observed in fissiparous races, such as has
been reported for Phagocata morgani in
Canada (Benazzi and Ball 1972) and for the
European Dugesia subtentaculata (Drapar-
naud) and some related species (Benazzi
1968, 1974; Gremigni and Banchetti 1972;
Benazzi and Deri 1980; de Vries 1986).
Nonetheless, the anatomy of the penis and
of the bursal duct separates P. spuria from
other species of the genus.
Etymology.—The name spuria (Latin,
spurious, false) alludes to the striking ex-
ternal similarity of the species to P. mor-
gan.
~Phagocata carolinensis Kenk
Fig. 8
This species was established and de-
scribed by Kenk (1979) from One Bat Cave,
Burke County, North Carolina. It is char-
acterized by having a truncate anterior end
with some lateral widening, but lacking
prominent pointed auricles; it also has a pair
of very small eyes. The most conspicuous
anatomical feature is the presence of an ex-
ceptionally large posterior section of the
672
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig. 8. Phagocata carolinensis, sagittal section through copulatory apparatus. am, male atrium; gp, gonopore;
odc, common oviduct; va, vagina.
bursal duct, or vagina (Fig. 8, va), which
may extend far posteriorly and is lined by
a glandular epithelium differing from that
of the anterior section of the duct. The va-
gina lacks any excessive muscular coating
and may appear variously folded in the pre-
served specimens. The penis, which in its
retracted state has a serpentine-like lumen
shown in the original description of the
species, may extend and assume an irreg-
ularly convoluted shape of the penis papilla
(Fig. 8).
Additional material of the species was ob-
tained from Wind Cave, McDowell County,
North Carolina, located near Marion. It was
collected by Cato O. Holler, Jr. on two visits
to the cave; 4 specimens, some mature, tak-
en from a small seep, 15 Apr 1984; and 5
specimens, all immature, collected beneath
rocks lying on moist clay in an abandoned
water course, 15 Feb 1985 (they were raised
to maturity in the laboratory).
Acknowledgments
Thanks are due to all collaborators who
furnished the materials discussed in the pa-
per. First of all, to Dr. Cato O. Holler, Jr.,
member of the North Carolina Cave Sur-
vey, an indefatigable collector of cave ani-
mals. Individual species were given to me
by Dr. Nicole Gourbault (Muséum Nation-
al d’Histoire Naturelle, Paris), Dr. Ian R.
Ball (now at Memorial University of New-
foundland), Leslie Hubricht (Meridian,
Mississippi), J. J. Lewis, and all their co-
workers mentioned under the individual
species. Photomicrographs were prepared
in Dr. John C. Harshbarger’s laboratory and
in part processed by Abbie J. Yorkoff
(Smithsonian Institution). Dr. Marian H.
Pettibone kindly reviewed the paper for sty-
listic and topical errors.
Literature Cited
Benazzi, M. 1968. Ulteriori ricerche sul controllo
genetico della scissiparita nelle planarie.— Bol-
lettino di Zoologia 35:424.
. 1974. Fissioning in planarians from a genetic
standpoint. Pp. 476-492 in N. W. Reiser and
M. P. Morse, eds., Biology of the Turbellaria.
New York, McGraw-Hill Book Company.
, and I. R. Ball. 1972. The reproductive ap-
paratus of sexual specimens from fissiparous
populations of Fonticola morgani (Tricladida,
Paludicola).— Canadian Journal of Zoology 50:
703-704, 2 pls.
, and P. Deri. 1980. Histo-cytological study of
VOLUME 100, NUMBER 3
ex-fissiparous planarian testicles (Tricladida
Paludicola).— Monitore Zoologico Italiano, new
series 14:151-163.
Gremigni, V., and R. Banchetti. 1972. Submicro-
scopic morphology of hyperplasic ovaries in ex-
fissiparous individuals in Dugesia gonocephala
s.l.—Atti della Accademia Nazionale dei Lince1,
Serie VIII, Rendiconti della Classe di Scienze
Fisiche Matematiche e Naturali 52:539-543, 3
pls.
Kenk, R. 1979. Freshwater triclads (Turbellaria) of
North America. XII. Another new cave pla-
narian from North Carolina.— Brimleyana 2:91-
96.
—. 1984. Freshwater triclads (Turbellaria) of
673
North America. XV. Two new subterranean
species from the Appalachian region. — Proceed-
ings of the Biological Society of Washington 97:
209-216.
Vries, E. J. de. 1986. On the taxonomic status of
Dugesia gonocephala and Dugesia subtentacu-
lata (Turbellaria, Tricladida, Paludicola).—
Journal of Zoology (London), Series A, 209:43-
59.
Department of Invertebrate Zoology, Na-
tional Museum of Natural History, Smith-
sonian Institution, Washington, D.C. 20560.
PROC. BIOL. SOC. WASH.
100(3), 1987, pp. 674-679
MYSIDIUM RICKETTST, A NEW SPECIES OF
MYSID FROM THE GULF OF CALIFORNIA
(CRUSTACEA: MYSIDACEA: MYSIDAE)
Elizabeth B. Harrison and Thomas E. Bowman
Abstract.—Mysidium rickettsi, the sixth species of the genus and the first
from the Pacific Ocean, is described from the Gulf of California. It is charac-
terized by the armament and proportions of the telson, the subdivision of the
sixth segment of the endopod of the pereopods, and the length of the peduncle
of antenna 2 in relation to the scale.
The genus Mysidium Dana includes M.
gracile (Dana, 1852) from Brazil, Florida,
Bermuda, Jamaica, Puerto Rico, Curag¢ao,
Venezuela, and the Virgin Islands; M. co-
lombiae (Zimmer, 1915) from Colombia
(Atlantic side), the Bahamas, the Florida
Keys, Jamaica, the Virgin Islands, Cuba,
Mexico, Belize, and Venezuela; M. inte-
grum W. Tattersall, 1951, from Florida, the
Virgin Islands, the Bahamas, the West In-
dies, Puerto Rico, Belize, and the Gulf of
Mexico; and M. rubroculatum and M. cu-
banensis Bacescu and Ortiz, 1984 from
Cuba. A 6th species, the first from the Pa-
cific is described below.
Mysidium rickettsi, new species
Figs. 1-3
Material.—Male holotype (USNM
233310) female allotype (USNM 233311),
and 50+ paratypes (USNM 81113), from
Pt. Marcial, Gulf of California 25°31’N,
111°01'W, surface, collected with night light
by E. R. Ricketts and J. Steinbeck, 24 Mar
1940; additional material (USNM 81114)
was collected with night light by E. R. Rick-
etts and J. Steinbeck in Bahia de Ohuira
(=San Carlos Bay), 25°38'N, 108°58'W, So-
nora, Mexico, 4 Apr 1940. Specimens were
also found in the stomachs of two species
of manta rays collected from four localities
in the Gulf of California by Giuseppe Me-
tabartolo di Sciara: From Mobula thur-
stoni: southern Gulf of California, penin-
sular side, near La Paz, in water 10 m deep
or less; near the eastern shore of Isla San
Francisco, Punta Arena de la Ventana,
24°04'N,109°52'W, and Bahia de la Ven-
tana, 24°05’N,109°55'W. From Mobula
munkiana: Estero on southwest side of Isla
San José, 24°55'N,110°38'W; and Bahia de
la Ventana.
Etymology. —Named for the late Edward
F. Ricketts, whose pioneering work on the
fauna of the Gulf of California is recorded
in Steinbeck and Ricketts (1941).
Description. —Length up to 10 mm. Ros-
trum round-triangular; posterior margin of
carapace emarginate, exposing last pereo-
nite. Eyestalk subquadrate, cornea covering
approximately half. Telson entire, rounded
apically; about twice as long as wide, ta-
pering posteriorly; posterior half with closely
spaced short, blunt, marginal spines (30-37
on each side).
Antenna 1: Segment | of peduncle about
one-third longer than wide, slightly longer
than segment 3; distolateral corner pro-
duced into short rounded process. Segment
2 short, trapezoidal, length of longer (me-
dial) margin less than half that of segment
3. Segment 3 nearly as long as wide, with
small lobe on distodorsal margin at base of
each flagellum, and with pair of long setae
at distomedial corner in 2. Male lobe lan-
Fig. 1. Mysidium rickettsi: A, Anterior end, dorsal; B, Appendix masculina, medial; C, Antenna | peduncle,
dorsal; D, Antenna 2, dorsal; E, Right mandible, with detail of incisor; F, Left mandible; G, Mandibular palp;
H,I, Inner and outer lobes of maxilla 1; J, Maxilla 2; K, Maxilliped; L, Labrum; M, Penis.
675
676 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig. 2. Mysidium rickettsi: A, Pereopod 1; B—G, Dactyl, “tarsal” segments, and distal part of merus of
Pereopods 2-7, setae of merus not shown; H—L, Pleopods 1-5.
VOLUME 100, NUMBER 3
Fig. 3. Mysidium rickettsi: A, Uropod; B, Telson.
ceolate, 8x as long as wide, proximal 7%
expanded ventrally, entire ventral surface
with long fine setae.
Antenna 2, scale lanceolate, slightly more
than 4x longer than greatest width, with
suture separating distal segment; lateral
margin straight, medial margin slightly con-
vex. Peduncle with 5 segments, 2 broad
proximal segments and 3 narrower distal
segments; distolateral corner of segment 2
produced into acute tooth. Peduncle and
scale subequal in length.
Right mandible: Incisor 3-cuspate; lacin-
ia subcylindrical, apex produced into 2
compound cusps, spine-row with 5-6 spines,
anterior spine robust, compound, others
denticulate; molar grinding surface striate,
medial margin denticulate, tuft of hairs on
posterior margin.
Left mandible: Incisor 6-cuspate, lacinia
sub-cylindrical, 4-cuspate, spine row with 4
spines, all denticulate. Molar blunt, poste-
rior tuft of hairs shorter than in right man-
dible. Palp with very short Ist segment, 2nd
segment 1.8 x length of 3rd; 2nd and 3rd
segments setose on medial margin, 3rd seg-
ment tapering in distal 3rd with 2 stout api-
cal spines.
Labium: With short, apically rounded
paragnaths with fringe of short setae on me-
dial margins.
Maxilla 1: Outer lobe with | surface seta
and about 9 apical spines, inner lobe with
4 long apical setae and about 7 shorter sub-
apical setae.
Maxilla 2: Endites well developed, heavi-
677
ly setose; distal segment of endopod about
1.8x as long as proximal segment, finger-
like in shape. Exopod not reaching distal
margin of proximal segment of endopod,
with ca. 13 plumose marginal setae.
Maxilliped (endopod of thoracopod 1):
Endite of basis well developed, nearly
reaching distal margin of ischium; medial
margins of segments densely setose.
Pereopod 1 (endopod of thoracopod 2):
Merus with 7 setae on anterior margin,
propus with 2 long and 2 shorter setae at
anterodistal corner, 4 setae at posterodistal
corner and row of setae on posterior margin;
dactyl with 7-8 hooked plumose setae and
3 longer, more slender setae. Pereopods 2-—
5 with 3 tarsal segments, 6 and 7 with 2.
Penis: Oval, with diagonal row of setae
near apex.
Pleopods: Pleopods 1-4 successively
longer. Pleopod 5 shorter than 2. Pleopods
1-3 and 5, viewed laterally, with straight
ventral margin with 5 or more long setae
arranged stepwise along length and one at
apex; dorsal margin straight along proximal
part, then curving inward to posteroventral
corner; pseudobranchial lobe with cluster of
short, fine, setae with inflated bases; other
single or grouped setae of same type on lat-
eral face. Pleopod 4 extending beyond distal
end of pleon, with seta of 3rd segment
reaching to end of telson. Endopod repre-
sented by short simple lobe bearing 7 setae,
distal seta 3 x as long as others; exopod very
long and slender, 4-segmented, relative
lengths (%) of segments 1—4 = 58:14:14:14.
678
Uropod endopod 1.5 as long as exopod,
both tapering distally; exopod 5x as long as
wide, endopod 7.6 x as long as wide.
Comparisons. —Mysidium rickettsi is
much longer (10 mm) than its Atlantic con-
geners, whose maximum recorded lengths
in mm are M. columbiae 7.3, M. cubanense
5.2, M. gracile 6.5, M. integrum 7.0, M.
rubroculatum 4.5. The telson of M. rickettsi
is relatively longer than in most other
species, having a length/width ratio of
slightly more than 2.0. For the other species
this ratio is M. columbiae 1.5, M. cubanense
2.0, M. gracile 1.7, M. integrum. 1.7, M.
rubroculatum 1.9. The maximum number
of marginal spines on the telson of M. rick-
ettsi, 74, 1s the highest in the genus (M. co-
lumbiae 52, M. cubanense 35, M. gracile
50, M. integrum 70, M. rubroculatum 37.
The apex of the telson is transversely round-
ed in ™. rickettsi, M. cubanense, M. inte-
grum, and M. rubroculatum, in contrast to
M. gracile, where it is shallowly emarginate,
and M. columbiae, in which it is distinctly
excavated.
The relative lengths of the tarsal segments
of pereopods 2-6 differs in species of My-
sidium. The first segment is much longer
than the following two in M. colombiae, M.
cubanense, M. gracile, and M. rickettsi. In
M. integrum and M. rubroculatum the three
tarsal segments are subequal. In M. colum-
biae there are three segments in the tarsus
of pereopod 2, but only two segments in
pereopods 3-7; in pereopod 2 the first seg-
ment is distinctly longer than the second or
third. :
The relative lengths of the segments of
the exopod of the 6 pleopod 4 is a useful
taxonomic character in Mysidium (Bratte-
gard 1969: Table 6). In M. rickettsi and M.
gracile segments 2—4 are subequal in length.
In M. rubroculatum and M. integrum seg-
ment 2 is distinctly longer than 3 or 4. In
M. cubanense and M. colombiae the endo-
pod has only three segments.
Ecology. —Steinbeck and Ricketts (1941:
152) reported that the mysids here de-
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
scribed as M. rickettsi occurred in swarms,
perhaps resulting from the light used in night
collecting. Small fish were noticed feeding
around the edges of these swarms. The pres-
ence of large numbers of M. rickettsi in ray
stomachs suggests that swarming does occur
under natural conditions, for seeking indi-
vidual mysids for food would be an un-
profitable expenditure of energy.
Brattegard (1969: 82,86,88) mentions that
shoaling in M. gracile, M. integrum, and M.
colombiae has been reported from Bermuda
and the Florida keys. Bacescu and Ortiz
(1984:21) reported shoaling in M. rubro-
culatum. Steven (1961), studying the shoal-
ing behavior of VM. colombiae in Jamaica,
reported that this species shoals when the
number of specimens 1s above a critical fig-
ure. Hahn and Itzkowitz (1986) studied site
preference and homing behavior in M. grac-
ile.
Acknowledgments
’ We thank Giuseppe Metabartolo di Sciara,
then of Scripps Institution of Oceanogra-
phy, for providing us with the specimens of
M. rickettsi from manta ray stomachs.
Literature Cited
Bacescu, M., and M. Ortiz. 1984. Contribution to the
knowledge of the Mysidacea (Crustacea) of the
Cuban insular shelf waters.—Travaux du Mu-
scum d’Histoire Naturelle Grigore Antipa, Bu-
charest 26:15-23. |
Brattegard, T. 1969. Mysidacea from shallow water
in the Bahamas and southern Florida. Part 1.—
Sarsia 39:17-106.
Dana, J. D. 1852. U.S. Exploring Expedition during
the years 1838, 1839, 1840, 1841, 1842, under
the command of Charles Wilkes, U.S.N., 13,
Crustacea, pt. I, 658 pp. Philadelphia: C. Sher-
man.
Hahn, P., and M. Itzkowitz. 1986. Site preference
and homing behavior in the mysid shrimp Mys-
idium gracile (Dana).—Crustaceana 51(2):215-
DMO),
Steinbeck, J., and E. F. Ricketts. 1941. Sea of Cortez.
A leisurely journal of travel and research. Viking
Press, New York, 598 pp.
VOLUME 100, NUMBER 3
Steven, D. M. 1961. Shoaling behaviour in a my-
sid.— Nature (London) 192(4799):280-281.
Tattersall, W. M. 1951. A review of the Mysidacea
of the United States National Museum. — Uniied
States National Museum Bulletin 201:1-x + 1-
292.
Zimmer, C. 1915. Schizopoden des Hamburger Na-
_turhistorischen (Zoologischen) Museums. —
Mitteilungen aus dem Naturhistorischen (Zool-
ogischen) Museum, Hamburg 32:159-182.
679
. 1918. Neue und wenig bekannte Mysidaceen
des Berliner Zoologischen Museums. — Mittei-
lungen aus dem Zoologischen Museum, Berlin
9:15-26.
Department of Invertebrate Zoology
(Crustacea), NHB Stop 163, National Mu-
seum of Natural History, Smithsonian In-
stitution, Washington, D.C. 20560.
PROC. BIOL. SOC. WASH.
100(3), 1987, p. 680
ERRATUM
In the article entitled ““Stegophiura ponderosa (Lyman), new combination, and Am-
phiophiura vemae and Homophiura nexila new species (Echinodermata: Ophiuroidea)
from the R/V Vema collections,” by Michael A. Kyte, published in Proceedings of the
Biological Society of Washington, volume 100, part 2, pages 249-256, an incorrect Figure
3 was printed. The correct figure and caption are given below.
Fig. 3. Homophiura nexila. Dorsal view (top) and ventral view (bottom).
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CONTENTS
Three new species of Renocila (Crustacea: Isopoda: Cymothoidae), external parasites of coral ©
reef fishes from the Ryukyu Islands of Japan
Ents ee ee TE ee
PU eth mn Nodt Uap Caterer ies
Lucy Bunkley Williams and Ernest H. Williams, Jr. Dae
Fallicambarus (Creaserinus) burrisi and F.,(C.) gordoni, two new burrowing crawfishes asso-
ciated with pitcher plant bogs in ie eipel and Alabama (Decapoda: Cambaridae)
J. P. Fitzpatrick, Jr.
Two new operculate land snails from the Palau Archipelago :
Fred G. Thompson and Thomas M. Iliffe
Classification of the Ascothoracida (Crustacea) Mark J. Grygier
A new subspecies of Rallina eurizonoides (Aves: Rallidae) from the Batan Islands, Philippines
Robert S. Kennedy and Charles A. Ross —
Two woodpeckers from the Late Pliocene of North America Alan Feduccia
Guitarra abbotti and G. isabellae, new sponges from the eastern Pacific Welton L. Lee
Marine Tubificidae (Oligochaeta) from Ruerto Rico with descriptions of two new species, —
Tubificoides aguadillensis and Heterodrilus paucifascis Michael R. Milligan ~
Ammoanita rosea, new genus and new species of deep water Trochamminidae, Foraminiferida —
George A. Seiglie and Mary B. Baker —
Ingolfiella fuscina, new species (Crustacea: Amphipoda) from the Gulf of Mexico and the
Atlantic coast of North America, and partial redescription of I. atlantisi Mills, 1967
Masahiro Dojiri and Jiirgen Sieg
Discias vernbergi, new species, a caridean shrimp (Crustacea: Decapoda: Bresiliidae) from the
northwestern Atlantic Billy B. Boothe, Jr. and Richard W. Heard
Notes on Varichaetadrilus Brinkhurst and Kathman, 1983 (Oligochaeta: Tubificidae)
Ralph O. Brinkhurst
Two new species of parrots (Aves: Psittacidae) from archeological sites in the Marquesas Islands
David W. Steadman and Marie C. Zarriello
A new species of stomatopod, Eurysquilla pumae (Crustacea: Stomatopoda: Eurysquillidae),
from the Gulf of California, Mexico Michel E. Hendrickx and José Salgado-Barragan
A new sre of Chevalia from the Caribbean Sea (Crustacea: Amphipoda)
J. L. Barnard and J. D. Thomas
Pinnotherids (Crustacea: Decapoda) and leptonaceans (Mollusca: Bivalvia) associated with
sipunculan worms in Hong Kong Raymond B. Manning and Brian Morton
The Pycnogonida types of H. V. M. Hall C. Allan Child
Further records of marine isopod crustaceans from the Caribbean Brian Kensley
Studies of the Lepidaploa complex (Vernonieae: Asteraceae). I. The genus Stenocephalum Sch.
Bip. Harold Robinson
Studies of the Lepidaploa complex (Vernonieae: Asteraceae). II. A new genus, Echinocoryne
: Harold Robinson
Upogebia synagelas, new species, a commensal mud shrimp from sponges in the western central
Atlantic (Decapoda: Upogebiidae) Austin B. Williams
Range extension of the genus Asymphorodes Meyrick (Lepidoptera: Cosmopterigidae)
J. F. G. Clarke
Reidentification of David Causey’s Caligus collections (Crustacea: Copepoda)
Roger Cressey and Patricia Nutter
Two new species of curimatid fishes (Ostariophysi: Characiformes) from Rio Grande do Sul,
Brazil Richard P. Vari
Deeveya bransoni, a new species of troglobitic halocyprid ostracode from anchialine caves on
South Andros Island, Bahamas (Crustacea: Ostracoda) |
Louis S. Kornicker and R. J. Palmer
Two new turtles from southeast Asia Carl H. Emst and William P. McCord
The caprellids (Amphipoda: Caprellida) of Bermuda: a survey of specimens collected from
1876-1987, including cave inhabitants, and the description of Deutella aspiducha, new
species Michael F. Gable and Eric A. Lazo-Wasem
Two new species and a new genus of miniature characid fishes (Teleoste1: Characiformes) from
northern South America Stanley H. Weitzman and Richard P. Vari
Heteromysis kensleyi and H. coralina, new species from the shallow waters off Looe Key,
Florida (Mysidacea: Heteromysini) Richard F. Modlin
Bahalana mayana, a new troglobitic cirolanid isopod from Cozumel Island and the Yucatan
Peninsula, Mexico _. Thomas E. Bowman
Freshwater triclads (Turbellaria) of North America. XVI. More on subterranean species of
Phagocata of the eastern United States Roman Kenk
Mysidium rickettsi, a new species of mysid from the Gulf of California (Crustacea: Mysidacea:
Mysidae) Elizabeth B. Harrison and Thomas E. Bowman
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