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ES SMITHSONIAN _ INSTITUTION NOILALILSNI_ NVINOSHLINS Satay: 5 ; oe = oc a a _ a = P) = tia PROCEEDINGS of the Biological Society of Washington VOLUME 83 1970 WASHINGTON PRINTED FOR THE SOCIETY -~a\ | HSONT™ Nig JUN 3.01971 LIBRARIES EDITOR Rocrer F. Cressey 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 1970-1971 OFFICERS President MEREDITH L. JONES Vice President DANIEL M. COHEN Secretary ISABEL P. F. CANET Treasurer HARALD A. REHDER COUNCIL Elected Members K. C. EMERSON JAMES A. PETERS CLAYTON E. RAY STANWYN G. SHETLER STANLEY H. WEITZMAN tii fii / ro “OTtTHUTEON JUN 3 * TABLE OF CONTENTS Volume 83 Banse, Karl. The small species of Euchone Malmgren (Sabel- lidae;. Polychaeta) 22.06. SN ee Biffar, Thomas A. Three new species of callianassid shrimp (Decapoda, Thalassinidea) from the western Atlantic Bohart, R. M. New species of Gorytini from western North America (Hymenoptera: Sphecidae)) Bourdon, Roland, and Thomas E. Bowman. Western Atlantic species of the parasitic genus Leidya (Epicaridea: Bopyr- TAA) ee! SE we ek eh Ie Seer i ade a ee Cartwright, Oscar L. A review of the aphodiine beetles of the Galapagos Islands (Coleoptera: Scarabaeidae) _...._-__ Chace, Fenner A., Jr., and Samuel L. H. Fuller. A new shrimp of the genus Gnathophyllum (Decapoda, Caridea) from Puerto TRico | se) ee Chernoff, Neil, and Robert Pope. Two new mites of the genus Klinckowstroemia from Mexico (Acarina: Klinckowstroemi- idae))) sees oe Ue a Child, C. Allan. Pycnogonida of the Smithsonian-Bredin Pacific expedition, 1957 2 ee Cohen, Daniel M., and C. Richard Robins. A new ophidioid fish (genus Lucifuga) from a limestone sink, New Providence Tsland;, Bahamas {200 0 eee Cole, Gerald A. The epimera of North American fresh-water species of Gammarus (Crustacea: Amphipoda) Damkaer, David M. Parastephos occatum, a new species of hyperbenthic copepod (Calanoida: Stephidae) from the in- land marine waters of Washington State == Dawson, C. E. A new wormfish (Gobioidea: Microdesmidae ) from the northem Red Séa 2.0 eee Dawson, C. E. The Caribbean Atlantic blenny Lupinoblennius dispar (tribe: Blenniini), with observations on a Pacific pop- Wlation.- iui re ee Downey, Maureen E. Drachmaster bullisi new genus and spe- cies of Ophidiasteridae (Echinodermata: Asteroidea), with a key to the Caribbean species of the family Downey, Maureen E. Marsipaster acicula, new species (As- teroidea: Echinodermata), from the Caribbean and Gulf of IMie@xXG G0 2a ao ae Drugg, Warren S. Two new neogene species of Tuberculodinium and one of Xenicodinium (Pyrrhophyta) —-___________ Flint, Oliver S., Jr. Studies of Neotropical caddis flies, XI: The genus Rhyacopsyche in Central America (Hydroptilidae) __ Gates, G. E. On an interesting Philippine species of the earth- AD) 493-504 145-154 287-308 505-514 267-272 273-286 77-82 309-312 115-122 515-526 worm genus Pheretima Kinberg, 1866 (Oligochaeta, Annel- EOIe) Lape stutter Cel ets Bay BAe on Mt Au ahah eke RA Halpern, Jerald A. Biological investigations of the deep sea. 53. New species and genera of goniasterid sea stars Handley, Charles O., Jr., and Jerry R. Choate. The correct name for the least short-tailed shrew (Cryptotis parva) of Guate- malian (Mammalassinsectivora)i i. ee Hart, C. W., Jr., and Dabney G. Hart. A new ostracod (En- tocytheridae, Notocytherinae) commensal on New Zealand VENA,» cet ae hc) Ni alee Sac AR Es ae eR OR Ce SO Hobbs, H. H., III. New entocytherid ostracods of the genus Ornithocythere and the description of a new genus Hobbs, Horton H., Jr. A new crayfish from the Nashville Basin, "UN @SRLONE RISES” 8a EEE UE ae aks a OU SR ete RES TD Hobbs, Horton H., Jr. New crayfishes of the genus Cambarus from Tennessee and Georgia (Decapoda, Astacidae) Hobbs, Horton H., Jr. A new crayfish of the genus Procambarus from Mississippi (Decapoda: Astacidae) Hobson, Katharine D. Novaquesta trifurcata, a new genus and species of the family Questidae (Annelida, Polychaeta) from ape. Codibay. Massachusetts 2.2 8 ee ee Hobson, Katharine D. Some polychaetes of the superfamily Eunicea from the North Pacific and North Atlantic Oceans _ Hoffman, Richard L. Studies on spirostreptoid millipeds. IX. The status of the Indian genus Leiotelus Chamberlin Holsinger, John R., and W. L. Minckley. A new genus and two new species of subterranean amphipod crustaceans (Gam- Mande) trom northern Mexico 20.22 Hope, W. Duane, and D. G. Murphy. A redescription of Enop- lus groenlandicus Ditlevsen, 1926 (Nematoda: Enoplidae) __ Hotchkiss, Frederick H. C. North American Ordovician Ophiu- roidea the genus Taeniaster Billings, 1858 (Protasteridae) Hubbard, John P., and Richard C. Banks. The types and taxa Braco Glade anle ys AU" We arte Ue eS ane Illg, Paul L., and Arthur G. Humes. Henicoxiphium redactum, a new cyclopoid copepod associated with an ascidian in Flor- Re amTicle NOTE @ ano lnm deca ee eet enlmene ones el. LEN es sen Kenk, Roman. Freshwater triclads (Turbellaria) of North Amer- ica. II. New or little known species of Phagocata __..__. ba Kenk, Roman. Freshwater triclads (Turbellaria) of North America. III. Sphalloplana weingartneri new species, from CAV EMIT MII AT Wee tele ow Aik ik weet en eh ae eee yD be Loeblich, Alfred R., Jr., and Helen Tappan. Thysanoprobolus, a new acritarch genus from the Early Devonian (Late Gedin- nian) Haragan Formation of Oklahoma, U.S.A. _.. Lynch, John D. A new eleutherodactyline frog from Amazo- MU PATA MYO LT AC ON teste eg ses Sis ae ee ei es ae 155-160 1-12 195-202 579-584 171-182 161-170 241-260 459-468 191-194 527-544 183-190 425-444 227-240 59-76 321-332 569-578 13-34 313-320 261-266 221-226 Manning, Raymond B. Nine new American stomatopod crusta- (cre V st ae Maltese er DION IAG ROMS PC nnTNeAC A EN EN Morrison, J. P. E. Western Atlantic Donax ____- Parin, N. V., and V. E. Becker. Materials for a revision of the trichiuroid fishes of the genus Benthodesmus, with the de- scription of four new species and one new subspecies _______- Parkes, Kenneth C. A revision of the red-rumped cacique, Cacicus haemorrhous (Aves: Icteridae) _.....___.___-__ Pettibone, Marian H. Two new genera of Sigalionidae (Poly- Chaetaly si okie A NI Th Uae ta Le eee Rehder, Harald A. A molluscan faunule from 200 meters off Valparaiso, Chile, with descriptions of four new species _____ Schultz, George A. Disposition of species of terrestrial isopod crustaceans of the genera Synuropus, Spherarmadillo, Sphae- roniscus and Scleropactes (Oniscoidea, Sphaeroniscidae) _ Shear, William A. The identity of Ergethus perditus Chamber- lin (Diplopoda, Polydesmida, Paradoxosomatidae) _________ Springer, Stewart, and Victor Sadowsky. Subspecies of the western Atlantic cat shark, Scyliorhinus retifer Springer, Victor G., and William F. Smith-Vaniz. Blennius antholops, new deep-water fish, from the Gulf of Guinea, with comments on the bathymetric distribution of the family Blenmid ae w/t rte thes Be ae Plies Ee ne Taylor, William R., Robert E. Jenkins, and Ernest A. Lachner. Rediscovery and description of the ictalurid catfish, Noturus flavipinnisy (280 ey ene 8 i Dh ee Tuck, Robert G., Jr. Rediscovery and redescription of the Khu- zistan dwarf gecko, Microgecko helenae Nikolsky (Sauria: Gekkonidae’)) 22.20 its ue ie ieee Wilson, Mildred Stratton. North American harpacticoid cope- pods 9. A new Mesochra (Canthocamptidae) from the Gulf of Mexico with notes on a related Mediterranean form ___ Work, Robert C. Further comments on the range of Murex florifer Reeve: 2o.)2 ee ee ee vt 99-114 545-568 351-364 203-214 365-386 585-596 123-132 455-458 83-98 215-220 469-476 477-482 483-492 349-350 BIOLOGICAL SOCIETY OF WASHINGTON PROCEEDINGS 1042nd Meeting—16 February 1970 Ninety-First Annual Meeting The 1042nd meeting and 91st Annual Meeting of the Society was called to order by President Joseph Rosewater at 2:35 p.m. in room 43 of the National Museum of Natural History. Twenty-three members were present. The minutes of the previous annual meeting were read by the Secre- tary, and approved as read. Treasurer Harald Rehder presented the financial report. Richard H. Manville reported that the auditing committee had found the Treasurer’s books to be in order. Reports of the Treasurer and Auditing Committee were accepted. The Secretary reported on membership changes in the past year, read- ing the names of five members whose deaths had been reported, and summarized actions of the Council. Editor Roger Cressey reported that Volume 82 of the Proceedings consisted of 870 pages in 69 numbers, and included the symposium on Natural History Collections, amounting to 204 pages. President Rosewater explained that the purpose of the proposed amend- ment to the bylaws, which had been distributed to members with the announcement of the meeting, was to avoid difficulties with the Internal Revenue Service. Robert Gibbs moved adoption of the amendment; the motion was seconded. After brief discussion to ensure that the proposed wording would serve the intended purpose, the motion was passed unanimously. The Secretary reported on the results of the election of officers. All candidates were elected. President-elect Meredith Jones reported on plans for a symposium, to be held in the fall of 1970, probably October. The symposium would concentrate on an examination of the state of knowledge of the region of Panama in which a sea-level canal is proposed, and on our ability to predict the biological consequences of such a canal. More than 20 potential speakers have been contacted. Victor Springer expressed the opinion that the yield from the in- vested funds of the Society was not as great as might be expected, and moved that the investment policy of the Society be reviewed. Motion was seconded and passed. Richard Manville asked whether action would follow such review, and was assured by President-elect Jones that a Finance Committee would be formed. There being no further business, the meeting was adjourned at 3:05 p.m. Richard C. Banks Secretary aa Be ; et uy. 5 ‘ e A I OE ae DC TE lt Rt de RNA SR Ae? hai 4 Ay ee as Ee ORD. Ra ee { i ® wre rea LAK, SiS bai } ww pee B74 0673 Vol. 83, No. 1, pp. 1-12 27 May 1970 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON BIOLOGICAL INVESTIGATIONS OF THE DEEP SEA. 53. NEW SPECIES AND GENERA OF CONUS TUBE ID) a SEA STARS! SMI ONT. By Jeratp A. HALPERN JUN J Institute of Marine Sciences, University of Viena Up ees Two new species of goniasterid sea stars were collected in the tropical Atlantic by the R/V Pillsbury and R/V Gerda of the Institute of Marine Sciences, University of Miami. They are the type-species of new genera. The R/V Pillsbury also collected the second known specimen of Plinthaster productus A. H. Clark. This species does not belong in the genus Plinthaster and is made the type-species of a new genus. The type-species of Aphroditaster, A. gracilis Sladen, belongs in the genus Pseudarchaster, making Aphroditaster invalid. A new genus is erected to accomodate Aphroditaster micro- ceramus Fisher. This research was supported by National Science Foundation grant GB-4936. The operations of the R/V Gerda and R/V Pillsbury were supported by National Science Foundation grant GB-7082. The field work was part of the National Geographic Society-University of Miami Deep-Sea Biology Program. The author is grateful to these agencies. Key TO THE GENERA DISCUSSED IN THIS PAPER 1. Unpaired median spine at apex of each mouth plate pair 2 Pair of median spines at apex of each mouth plate pair 3 2. Adambulacral furrow margin straight —_ Fisheraster new genus Adambulacral furrow margin angular — __ Pseudarchaster 3. Abactinal plates completely covered by granules — 4 Abactinal plates not completely covered by granules — 6 1 Contribution No. 1181 from the Institute of Marine Sciences, University of Miami. 1—Proc. Biot. Soc. Wasu., Vou. 83, 1970 (1) bo Proceedings of the Biological Society of Washington 4, Adambulacral granulation very crowded; plate not visible _ Peltaster Adambulacral granulation spaced; plates visible between sub- ambulacral oranules. =... 5.0 ee 5 5. Irregular secondary abactinal plates present; abactinals covered only by spherical granules — Pillsburiaster new genus No secondary abactinal plates; peripheral granules of radial abactinalstangullar 2242.05 we Ceramaster 6. All abactinal plates surrounded by single row of granules; naked mien, losemIinNy Glligcy [OSS Plinthaster Some abactinal plates bearing more than single row of granules; naked, areas’ ‘smooth 2... =. eee 7 7. Radial plates mainly naked. General form stellate; more than six superomarginals in contact medially Diplasiaster new genus Radial plates having only small naked area. General form pentagonal; less than three superomarginals in contact medially GEE Bese ah Daal SU ane el ieee eR Apollonaster new genus Fisheraster new genus Diagnosis: Unpaired (but not recurved) median spine at apex of each mouth plate pair. No spines or spinules on actinal and inferomarginal plates. Adambulacral furrow margin straight. Remarks: This genus belongs in the subfamily Pseudarchasterinae but it is closer to the Goniasterinae than any of the known genera of this subfamily. Type-species: Aphroditaster microceramus Fisher. Etymology: This genus is named after Walter K. Fisher, who described the type-species and was an excellent asteroid systematist. Fisheraster microceramus (Fisher, 1913) Aphroditaster microceramus Fisher, 1913, pp. 626-627; 1919, pp. 225- OAS) Folk, BS), water, IL, yoll, le sakes, IU Remarks: This species is well described and illustrated by Fisher (1919). Plate 70, figure 5 is not a photograph of Fisheraster micro- ceramus, but of the type of Paragonaster stenostichus. Type: U.S. National Museum cat. no. 30535. Type-locality: Buton Strait, Celebes, 05°35’S, 122°20’E, 1023 m, Albatross sta. 5648. Distribution: This species is known only from the type-locality. Pillsburiaster new genus Diagnosis: Abactinal plates tabulate, completely covered by spherical granules; loosely arranged into primary and secondary plates in radial area and center of disk. Papulae irregularly distributed between primary and secondary abactinal plates. Subambulacral spines very small, not crowded. New Goniasterid 3 Remarks: The abactinal surface in this genus is similar to that of the species of Peltaster, but the secondary plates are not as regularly arranged and are not clearly delineated from the primary plates. The peripheral granules of the radial plates are not different from the central granules, as they are in Ceramaster. The actinal surface is similar to that of Plinthaster and Ceramaster. This genus belongs in the subfamily Goniasterinae. It appears to be intermediate to Ceramaster and Peltaster. Pentagonaster ernesti Ludwig, 1905, from the tropical eastern Pacific, belongs in this genus. Type-species: Pillsburiaster geographicus new genus, new species. Etymology: Pillsbury—after the R/V John Elliot Pillsbury of the Institute of Marine Sciences, University of Miami. Many species of sea stars new to science have been collected on this ship’s expeditions; aster, Greek—a common ending for genera of sea stars. Pillsburiaster geographicus new genus, new species Figures 1, 2 (top) Material studied: Holotype: R=75 mm, r=30 mm, R/r=2.8; 04°36’N, 09°46’W, 1464-1556 m, R/V Pillsbury sta. 76, 4 June 1964, USNM E11258.—Paratypes: same location as holotype, 2 spec., USNM E11259.—03°50’N, 02°33’W, 1949-1986 m, R/V Pillsbury sta. 34, 29 May 1964, 1 spec., UMML 40.240. Diagnosis: Abactinal granules coarse, spherical. Five or six adambu- lacral furrow spines. Ten or eleven mouth furrow spines. Description: Five arms. R= 75 mm, r = 30 mm, R/r = 2.8. General form stellate. Abactinal plates loosely arranged into primary and secondary plates in long, narrow radial areas and small center of disk. Primary plates only two to four times as large as secondary plates; secondary plates loosely arranged between primary plates. Abactinal plates tabulate, irregularly round; completely covered by very closely crowded, coarse, spherical granules. Many abactinal plates bearing two or three valved, spatulate sugar-tong pedicellariae with short, wide, thick jaws. Papulae in center of disk and radial areas; papular pores irregularly distributed, each containing single papula. Superomarginal and inferomarginal plates corresponding; thirty-two plates in each series. Lateral angle of marginals depressed; plates mainly in horizontal plane. Marginals more angular on distal third of arm. Superomarginals slightly wider than long. In interbrachial arc, lower half of each plate completely covered by granules similar to those on abactinals. Upper half bordered by two to three rows of similar granules; center bare or with one to three scattered granules. Naked area becoming larger distally, so that last six plates surrounded by single row of granules. Last three or four pairs of superomarginals contiguous medially. Terminal plate moderately large; in shape of truncated cone 4 Proceedings of the Biological Society of Washington Fic. 1. Pillsburiaster geographicus new genus, new species, holotype: top, abactinal view, 0.8.—bottom, abactinal view, 4.8x. New Goniasterid 5 with apical depression bearing one large, short, wide, conical spine. Inferomarginal plates slightly wider than long; completely covered by granules similar to those of superomarginals. Actinal plates moderately large, rhombic; extending about half way down arms. Each plate completely covered by coarse, rounded granules similar to those on inferomarginals. Adambulacral plates approximately square with straight furrow margin bearing five or six short, subequal, compressed furrow spines. Three to four irregular rows of four to six very short, blunt subambulacral spinules, slightly taller than actinal granules. Many adambulacral plates bearing two or three valved, spatulate sugar-tong pedicellariae, similar to abactinal pedicellariae but not as thick. Each mouth plate bearing ten or eleven furrow spines; first nine or ten spines subequal, slightly taller and thicker than adambulacral furrow spines. Median spine more strongly compressed and slightly taller than other mouth furrow spines. Rest of each mouth plate completely covered by 22-24 large, rounded granules, about twice as large as actinal granules. Madreporite irregularly round, about five times as large as adjacent abactinal plates; located approximately one-third distance from center of disk to middle of interbrachial arc. Type: United States National Museum, cat. no. E11258. Type-locality: Gulf of Guinea, off Cape Palmas, 04°36’N, 09°46’W, 1464-1556 m, R/V Pillsbury sta. 76. Distribution: This species is known from only two localities in the northern Gulf of Guinea; off Cape Three Points, 1949-1986 m and off Cape Palmas, 1464-1556 m. Discussion: Various characters of the paratypes are as follows: Pills- supero- adamb. mouth bury R i Mar- marginals furrow furrow station (mm) (mm) R/r ginals incontact spines spines 34 66 27 2.4 26 0-1 5-6 11-12 76 80 32 2.9 34 2-4 5-6 10 76 65 28 2.3 26 0-1 6 MI In the specimen measuring R = 80 mm, some of the superomarginals in the middle of the interbrachial arc are completely covered by granules. Remarks: This species is very similar to Pillsburiaster ernesti and differs from it by having actinal pedicellariae and less stout sub- ambulacral granules. Etymology: geographicus—in reference to the National Geographic Society whose grant to the Institute of Marine Sciences, University of Miami for investigations of the deep-sea fauna helped make possible the discovery of many sea stars new to science. 6 Proceedings of the Biological Society of Washington Fic. 2. Top, Pillsburiaster geographicus new genus, new species, holotype, actinal view, 6.8x.—bottom, Diplasiaster productus (A. H. Clark), specimen from R/V Pillsbury sta. 707, abactinal view, 2.9x. New Goniasterid i Diplasiaster new genus Diagnosis; Abactinal plates low-tabulate; surrounded by double row of granules in radial areas, single row elsewhere; no bosses. Supero- marginals contiguous medially throughout length of arm. General form stellate. Etymology: Diplasios, Greek—double; aster—a common ending for genera of sea stars; in reference to the double row of granules on the radial abactinal plates. Remarks: This genus belongs in the subfamily Goniasterinae. It appears to be most closely related to Apollonaster. It differs from Plinthaster in the shape and granulation of its abactinal plates and in the large number of superomarginal plates in contact. Type-species: Plinthaster productus A. H. Clark. Diplasiaster productus (A. H. Clark, 1917) Figures 2 (bottom) and 3 Plinthaster productus A. H. Clark, 1917. pp. 67-69; 1954, p. 375. Material studied: Holotype: R=27 mm, r=12 mm, R/r=2.3; 93°10'N, 82°23’W, 567 m, Albatross sta. 2154, 30 April 1884, USNM 36930.—11°22’N, 62°22’W, 78 m, B/V Pillsbury sta. 707, 19 July 1968, UMML 40.235. Diagnosis: Four or five adambulacral furrow spines. Each mouth plate bearing eight furrow spines. Description: Five arms. R = 29 mm, r=13 mm; R/r = 2.2. General form stellate, with broad, flat disk and narrow, strongly tapered arms. Interbrachial arcs wide, rounded. Abactinal plates low-tabulate, restricted to disk. Plates smooth, naked in center; surrounded by single row of flattened granules except in radial areas, where some plates bearing second row of rounded granules inside peripheral row. Twenty superomarginal plates naked except for one peripheral row of granules similar to those surrounding abactinals and U-shaped cluster of rounded granules adjacent to inferomarginals. Seven or eight pairs of superomarginals in contact medially. Terminal plate small, oval, surmounted by short spine. Twenty inferomarginal plates corresponding with superomarginals. Inferomarginals surrounded by two rows of rounded granules. Rest of each plate naked except for small area adjacent to superomarginals covered by closely crowded, rounded granules. Actinal plates large, rhombic, completely covered by moderately large, rounded granules. Adambulacral plates about half as big as adjacent actinals. Adambu- lacrals rectangular (about twice as wide as long) with straight furrow margin bearing four or five short, subequal, cylindrical furrow spines with rounded tips. First subambulacral row of two or three large, conical spines. Three to six rounded granules similar to those of actinals, in one or two irregular rows, covering rest of plate. 8 Proceedings of the Biological Society of Washington Fic. 3. Diplasiaster productus (A. H. Clark), specimen from R/V Pillsbury sta. 707: top, abactinal view, 8.5x.—bottom, actinal view, UB New Goniasterid 9 Mouth plates relatively small. Each plate bearing eight furrow spines. First seven spines subequal, similar to adambulacral furrow spines; median spine slightly taller and stouter. Eight to ten scattered spines, grading actinally into granules similar to those of actinals, covering rest of plate. Anus subcentral, inconspicuous. Madreporite irregularly round, slightly smaller than adpacent abactinals; located approximately one-third dis- tance from center of disk to middle of interbrachial arc. Type: United States National Museum, cat. no. 36930. Type-locality: 23°10'N, 82°23’W, 567 m, Albatross sta. 2154. Distribution: This species is known from only two localities: off Havana, Cuba, 567 m and off the Peninsula de Paria, Venezuela, 78 m. Apollonaster new genus Diagnosis: Abactinal plates of radial areas covered by peripheral row of large, flattened granules and one to three additional rows of rounded granules, leaving small, naked central area. Abactinal plates of inter- radial areas, center of disk and arms covered by peripheral row of small, flattened granules and zero to two additional rows of rounded granules, leaving large, naked central area. Central naked areas smooth. Remarks: This genus belongs in the subfamily Goniasterinae. It appears to be intermediate between Ceramaster and Diplasiaster. Type-species: Apollonaster yucatanensis new genus, new species. Etymology: Apollo—in honor of the flight of Apollo 11, which landed the first men on the moon; aster, Greek—a common ending for genera of sea stars. Apollonaster yucatanensis new genus, new species Figures 4, 5 Material studied: Holotype: R=36 mm, r= 24 mm, R/r=1.5; 20°57'N, 86°34’W, 40-165 m, R/V Gerda sta. 899, 10 September 1967, USNM E11285. Diagnosis: Six or seven subequal, strongly compressed adambulacral furrow spines. Thirteen mouth furrow spines. Description: Five arms. R = 36 mm, r = 24 mm, R/r = 1.5. General form arcuate pentagonal. Abactinal plates low-tabulate. Plates of radial areas covered by peripheral row of large, flattened granules and one to three additional rows of rounded granules, leaving small, naked area in center of plate. Plates of interradial areas, center of disk and very short arms covered by peripheral row of small, flattened granules and zero to two additional rows of rounded granules, leaving large naked area in center of plates. Central naked areas smooth. Some plates bearing small, excavate sugar-tong pedicellariae. Fourteen superomarginal and sixteen inferomarginal plates correspond- ing, except at tip of arm, where two inferomarginals correspond with 10 Proceedings of the Biological Society of Washington Fic. 4. Apollonaster yucatanensis new genus, new species, holotype, abactinal view; Top, actual size—Botton, 6.9. New Goniasterid il ot 3° @e @ vee, * Fic. 5. Apollonaster yucatanensis new genus, new species, holotype, actinal view, 6.9x. one superomarginal. Superomarginals slightly wider than long; sur- rounded by two rows of small, rounded granules, with several similar granules scattered about center. Some plates bearing small, excavate sugar-tong pedicellariae like those on abactinals. Terminal plate very small, oval. Inferomarginals slightly wider than long; surrounded by three to seven rows of small, rounded granules like those of supero- marginals; several similar granules scattered about center of plate. Some plates bearing small, excavate sugar-tong pedicellariae. Actinal plates large, rhombic, completely covered by moderately large, rounded granules. Some plates bearing excavate sugar-tong pedicellariae slightly larger than those on abactinals. Adambulacral plates rectangular (about twice as wide as long) with straight furrow margin bearing six or seven subequal, strongly com- pressed furrow spines. First subambulacral row of three or four large, prismatic granules. Three or four more irregular subambulacral rows of three to five granules similar to those on actinals. Some _ plates bearing excavate sugar-tong pedicellariae. Mouth plates long, narrow. Each plate bearing thirteen subequal furrow spines, similar to those on adambulacrals; median spine slightly enlarged. Rest of plate covered by 20-25 prismatic granules grading actinally into granules similar to those of actinals. Madreporite irregularly round, about same size as adjacent abactinals; 12 Proceedings of the Biological Society of Washington located approximately one-quarter distance from center of disk to middle of interbrachial arc. Type: United States National Museum cat. no. E11285. Type-locality: Arrowsmith Bank, Yucatan, 20°57'N, 86°34’W, R/V Gerda sta. 899. Remarks: The naked areas of the abactinal plates are not due to the granules being rubbed off. The granules are set in small, hemisperical pits in the plate which are readily apparent when the granules are rubbed off. LITERATURE CITED Ciark, A. H. 1917. Four new echinoderms from the West Indies. Proc. Biol. Soc. Wash., 30: 63-70. FisHer, W. K. 1913. Four new genera and fifty-eight new species of starfishes from the Philippine Islands, Celebes, and the Moluccas. Proc. U.S. Nat. Mus., 43: 599-648. 1919. Starfishes of the Philippine Islands and adjacent waters. Bull. U.S. Nat. Mus., 100(3): 1-547, pls. 1-155. Lupwic, H. 1905. Reports on an exploration off the west coasts of Mexico, Central and South America, and off the Galapagos Islands, . . . by . . . “Albatross” during 1891 ... 32. Asteroidea. Mem. Mus. comp. Zool. Harvard, 32: 1-292, pls. 1-36. 57s ad a ae Vol. 83, No. 2, pp. 13-34 27 May 1970 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON FRESHWATER TRICLADS (TURBELLARIA) OF NORTE NI HSo, Vareanet AMERICA. II. NEW OR LITTLE KNOWN NG SPECIES OF PHAGOCATA.! We By RoMAN KENK? BRARIES The George Washington University and Smithsonian rere SS During work on a revision of the North American fauna of freshwater triclads I studied planarian specimens from several states either collected on field trips or sent in by various collaborators. I wish to express my grateful appreciation to all who assisted me in the field or provided materials for study. Thanks are due particularly to Drs. Thomas E. Powell, Jr., R. O. Flagg, and Mr. Richard Whitten of the Carolina Biologi- cal Supply Company; and to Messrs. Timothy F. Hicks, Vernon M. Hawthorne, Phillip H. Baker, William F. Hart, Jerry H. Carpenter, and James F. Leser who in one way or another were instrumental in facilitating my access to the materials discussed in this paper. Phagocata was established by Leidy (1847: 248) as a sub- genus of Planaria O. F. Miller (1776) for the species Planaria gracilis described by Haldeman (1840). The chief charac- teristic distinguishing Phagocata from Planaria s. str., in Leidy’s opinion, was the presence of multiple pharynges which at that time had not been observed in other species of fresh- water triclads. Much later it was discovered that polypharyngy may occur in several genera of planarians (Crenobia, Sphal- loplana) and that it has no generic taxonomic significance. The present definition of the genus Phagocata (synonym, 1Supported by National Science Foundation Grant GB-6016 to the George Washington University. 2 Authors address: Department of Invertebrate Zoology, Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560. 2—Proc. Bron. Soc. Wasu., Vou. 83, 1970 (11:3) 14 Proceedings of the Biological Society of Washington 1mm i) Fic. 1-2. 1. Outline sketches of species of Phagocata, from life. A, P. bulbosa; B, P. crenophila; C, P. oregonensis; c. copulatory complex; i, area occupied by intestinal branches; ph, pharynx. 2. Phagocata bulbosa, semidiagrammatic view of copulatory apparatus in sagittal sec- tion. am, male atrium; b, copulatory bursa; bd, bursa duct or stalk; de, ejaculatory duct; gp, genital aperture; m, mouth; odc, common oviduct; vd, vas deferens; vs, seminal vesicle. North American Triclads 15 Fonticola Komarek) is as follows (Kenk, 1935: 103, and Hy- man, 1937: 302). Planariidae whose oviducts—without embracing the stalk of the bursa copulatrix—unite in a common oviduct which opens into the genital atrium. Male atrium without radial muscle plates. Adenodactyls absent. Eyes usually two. Type of the genus, Planaria gracilis Haldeman (1840: 3). Phagocata bulbosa new species® Type material: Holotype, series of sagittal sections of posterior half of the body, on 2 slides, U.S. National Museum No. 40967. Paratypes in the author’s collection. External features (Fig. 1A): Mature animals are up to 12 mm long and 1.7 mm wide. The anterior end is truncate, with straight or centrally bulging frontal margin and rounded auricular edges. Behind the head there is no pronounced narrowing or neck, at most a very slight constric- tion behind the auricular lobes. The lateral margins diverge very gradually up to the middle of the body where the greatest width is attained. Behind the pharyngeal region the body tapers to a rather pointed posterior end. There are usually two eyes, their distance from each other one-third or less of the width of the head at the level of the eyes. Each eye is farther removed from the frontal margin than from the lateral margin. The pharynx is inserted at about the middle of the body and measures in length approximately one-fifth of the body length. The copulatory organs occupy the anterior half of the post-pharyngeal region. The animal is usually uniformly pigmented, the dorsal surface being a variable shade of gray, often presenting a mottled appearance. The ventral surface is somewhat lighter. The ocular specks on the dorsal side and a small area around the mouth opening ventrally are free of pigment, white. As is seen from this description, the species resembles very closely a number of other American planariids from which it connot be dis- tinguished in life by external characters: Phagocata velata (Stringer), P. vernalis Kenk, P. crenophila Carpenter, Planaria dactyligera Kenk, Hymanella retenuova Castle, and probably Phagocata monopharyngea Hyman (which has not been studied in life). I have not been able to separate the new species, with any degree of confidence, in mixed collec- tions or cultures where it was associated with Planaria or Hymanella. Only the analysis of serial sections made it possible to identify the in- dividual species. Anatomy of the anterior end: There is no adhesive organ developed. The submarginal zone of adhesive gland openings, which parallels the 8 bulbosus, Latin, bulbous, referring to the size of the penis bulb. 16 Proceedings of the Biological Society of Washington body margins, continues into the head region along the frontal margin without interruption and without increase in width. There are no muscular differentiations such as are generally observed in adhesive organs. On either side of the head, along the auricular edges, the surface epithelium is modified, the cells bearing conspicuous cilia and containing very short rhabdites. It is highly probable that these two strips represent the auricular sense organs. On the ventral side of the head, immediately behind the adhesive gland zone, there is a small patch of ciliated epithelium approached by numerous gland ducts with a fine-grained, slightly eosinophilic secretion (as opposed to the deeply staining secretion of the adhesive glands). The cells of this epithelium generally lack rhabdite inclusions. The function of this organ is not clear. Reproductive system: The numerous testes are predominantly ventral, occupying on either side a broad zone extending from a short distance behind the head to approximately the level of the mouth opening. Some testicular follicles may, however, have a more dorsal position or, at full maturity, individual large follicles may occupy the entire dorso- ventral depth of the body. The two ovaries, accompanied laterally by large parovarial cell masses, lie, as is typical for freshwater triclads, on the medial side of each ventral nerve cord, generally behind the first lateral branch of the anterior intestinal ramus. The yolk glands or vitellaria occupy all available mesenchymal spaces both dorsally and ventrally in a zone extending from the ovaries to the posterior end of the body. The copulatory apparatus (Fig. 2) conforms in every way with the definition of the genus Phagocata as proposed by Hyman (1937: 302). The genital aperture or gonopore (gp) lies at or a little behind the middle of the postpharyngeal region. There is no common genital atrium developed as the male atrium (am) and the bursa duct (bd) meet at the gonopore. The male atrium is conical, tapering posteriorly, and duplicating the shape of the penis papilla. It is lined with a flattened or cuboidal epithelium below which there are two muscle layers, a circular and a longitudinal one. The penis consists of a large, spherical bulb and a finger- or cone- shaped papilla. The bulb is formed by strong muscle fibers running in various directions in a more or less concentric arrangement. It is traversed by many gland ducts with a faintly eosinophilic, fine-grained secretion, which enter it from the surrounding mesenchyme and open into its lumen. The papilla of the penis is covered with a flattened epithelium. Underlying this epithelium is a layer of very dense fine fibers which at the base of the papilla is continuous with the circular muscle layer of the male atrium; this fibrous layer may be of an elastic rather than muscular nature. In comparing the penis papilla in nine series of sagittal sections it was seen that the shape of this organ appeared quite uniform, straight, not showing any distortions or con- tractions such as are generally observed in soft muscular organs in North American Triclads 17 planarians. Apparently the relative stiffness of the papilla is due to the fibrous layer. The cavity of the penis bulb, the seminal vesicle (vs), is rather large, with an irregular outline. Its wall consists of a columnar epithelium of a glandular nature. Posteriorly the cavity narrows and continues into the papilla as a straight narrow canal, the ejaculatory duct (de), which opens at the point of the papilla. The seminal vesicle, particularly in its caudal portion, receives the gland ducts which have penetrated the tissue of the penis bulb. The two vasa deferentia (vd), which at the level of the pharynx have assumed the shape of expanded, tortuous ducts (the false seminal vesicles or spermiductal vesicles), enter the penis bulb from either side antero-ventrally. They retain their expanded shape, form a few con- volutions within the bulb, and finally empty separately through short narrow passages into the antero-lateral part of the seminal vesicle. The two oviducts or ovovitelline ducts, which run from the ovaries posteriorly along the ventral nerve cords, ascend dorsally at the level of the copulatory complex and unite above the male atrium to form a common oviduct (odc). This then proceeds posteriorly along the atrial wall and connects with the atrial cavity quite close to the gonopore. The end parts of the paired oviducts and the common oviduct receive the highly eosinophilic “shell glands” as is the general rule in the genus Phagocata. The copulatory bursa (b) is a large sac situated between the wall of the pharyngeal pouch and the penis bulb. It is highly lobate, with many diverticula extending mainly in a lateral direction. Its outlet, the bursal canal or duct (bd) runs from the bursa caudally, somewhat to the left of the penis bulb, then curves ventrally to connect with the gonopore. The duct shows no histologically distinct divisions. Some of the cells of its epitelial lining project villus-like into the lumen of the duct. The duct is surrounded by two muscle layers, an inner circular and an outer longitudinal layer. Distribution and ecology: Phagocata bulbosa was collected on 13 June 1968 on the grounds of the Warren Laboratories of the Carolina Biological Supply Company (situated on highway U.S. 158, 3 miles east of Warrenton), Warren County, North Carolina. It was taken on liver bait in two localities, one (the type locality) on the side entrance road to the Laboratories in a seepage area with highly colored water, ac- companied by Hymanella retenuova Castle; in the other locality, also on the side entrance road, it occurred together with Planaria dactyligera musculosa Kenk. Some of the specimens collected were sexually mature, others matured in laboratory cultures. I am indebted to Dr. Thomas E. Powell, Jr., and Dr. R. O. Flagg of the Carolina Biological Supply Company for kindly extending to me the hospitality of the Laboratories and aiding me in the collection of the specimens. Laboratory cultures: Phagocata bulbosa was kept in the laboratory 18 Proceedings of the Biological Society of Washington in spring water at about 14° C. and was fed beef liver and Tubifex. Young specimens became sexually mature after several weeks and de- posited egg capsules (cocoons). As the cultures were mixed, containing also the other species present at the collecting sites (which in life could not be separated from P. bulbosa), only a few measurements of the egg capsules could be made for the individual species. All cocoons were unstalked, ellipsoidal, and attached to the substrate by a jellylike transparent substance. From one egg capsule, isolated from a mixed culture, three young emerged after 38 days. They matured, or at least developed parts of the reproductive system, in about four months and could then be assigned to P. bulbosa on the basis of their anatomy. Two specimens from a mixed culture (which were later identified as P. bulbosa on the basis of serial sections) laid two cocoons measuring 0.7 x 0.9 mm and 0.8 x 1.2 mm, respectively. These cocoons hatched 36 and 50 days after their deposition. No asexual reproduction (by fissioning or fragmentation) was observed in over one year of culturing. Taxonomic position: The genus Phagocata in its broader sense (in- cluding also the forms frequently separated as Fonticola, Atrioplanaria, and Penecurva) comprises at present close to 50 species and subspecies widely distributed in the northern hemisphere (Eurasia and North America). The outstanding specific characters of Phagocata bulbosa are its pigmentation, the presence of a singie pharynx, and the anatomy of its reproductive system: large, spherical penis bulb containing a seminal vesicle, conical penis papilla with narrow ejaculatory duct opening at its tip, far caudal opening of the common oviduct, and absence of a sphincter on the duct of the copulatory bursa. With regard to the anatomy of the penis it shows certain similarities with some widely scattered exotic species of the genus (in which, however, the zone of the testes reaches to the posterior end): three Japanese species, P. kawakatsui Okugawa (1956), which has a muscular sphincter in the posterior section of the bursal duct and a different course of the vasa deferentia entering the penis; P. iwamai Ichikawa and Kawakatsu (1962a), which has a wider ejaculatory duct; P. teshirogii Ichikawa and Kawakatsu (1962b), with a more flexable penis papilla; a species from Manchuria, P. wenoi Okugawa (1939), which has on each side a group of 4-8 eyes; and P. altaica Livanov and Zabusova (1940) from Lake Teletskoye in the Russian part of the Altai Mountains in Siberia, in which the penis papilla appears to be soft and easily distorted, coming back upon itself in the published illustrations. As far as the American planarians are concerned, the superficial similarity of the external features of P. bulbosa with those of other species and even genera has already been mentioned. The principal differences between our species and other pigmented American species of Phagocata are as follows: P. gracilis (Haldeman) and P. woodworthi Hyman are polypharyngeal; P. velata (Stringer) has predominantly dorsal testes and a diverticulum on the ejaculatory duct; P. vernalis Kenk is distinguished by a U-shaped North American Triclads 19 Fic. 3. Phagocata crenophila, photographs from life. A, specimen from Logan, Utah; B, specimen from Tahoe City, California. <4. copulatory bursa and lacks a well-developed penis bulb; in P. crenophila Carpenter the ejaculatory duct opens on the ventral side of the penis papilla; P. tahoena Kawakatsu, a small species with a different external habit, has the testicular zone extending to the posterior end; and P. monopharyngea (Hyman), originally described as a monopharyngeal subspecies of P. gracilis, seems to have a very weak penis bulb and a wide lumen of the ejaculatory duct. Parasites: The majority of the specimens of Phagocata bulbosa ex- amined on sections contained the holotrichous ciliate parasite, Sieboldiel- lina planariarum (Siebold) in the intestinal cavity and frequently in the lumen of the pharyngeal pouch. Several of the animals were infested also with a peritrichous epizoite, probably Urceolaria mitra (Siebold). Phagocata crenophila Carpenter, 1969 The species was recently described by J. H. Carpenter (1969) who collected and studied it in Utah and deposited type material in the Smithsonian Institution (No. 38052-38054). The present discussion furnishes additional data on its morphology and distribution. External features: (Fig. 1B and 3): Phagocata crenophila is another of the group of pigmented planarians which cannot be positively identi- fied in life (see the remarks under the preceding species, P. bulbosa). 20 Proceedings of the Biological Society of Washington i] e | ; | 7 eq a b c d e Fic. 4. Phagocata crenophila, outline drawings of anterior end. According to Carpenter (1969: 276), mature specimens measure 12-22 mm in length and 1.5-2 mm in width. My specimens, when mature, were mostly about 17 mm long and 2 mm wide. The head is truncate but in detail may present various shapes when the animal is in motion. Generally the frontal margin bulges slightly in the center and the edges protrude very little laterally. In one population (from Tahoe City, California), however, the auricular edges protrude considerably so that a distinct narrowing or neck is formed behind them. Figure 4 shows the outlines of the head of specimens of various provenances in gliding locomotion. The dorsal surface is gray to black, the ventral side somewhat lighter. The distance between the two eyes amounts to about one-third the width of the head at the level of the eyes, and each eye is farther removed from the frontal margin than from the lateral margin. Reproductive system: A general description of the gonads, gonoducts, and the copulatory organs has been given by Carpenter (1969). The zone of the testes, which are predominantly ventral, extends from the level of the ovaries to almost the posterior end of the body. I wish to supplement Carpenter's account of the copulatory complex, with particu- lar regard to its muscular and glandular structures (Fig. 5). The genital aperture (gp) leads into two cavities: anteriorly the male atrium (am) and dorsally, somewhat to the left of the midline, the duct (bd) of the copulatory bursa (b). There is no common genital atrium discernible. The penis consists of a feebly muscular bulb and a well-developed plug- shaped papilla which bears at its distal end a characteristic lobelike appendage (pp). The two vasa deferentia (vd) enter the bulb antero- laterally, acquire a distinct coat of circular muscle fibers, and proceed transversally toward the midline where they unite. The common vas deferens thus formed runs posteriorly or postero-dorsally (depending on the state of contraction of the penis) for a short distance, then widens to a small cavity (vs) which we may consider to be the equivalent of a seminal vesicle. This cavity, which never assumes a conspicuous volume, is indicated by Carpenter (1969, fig. 12, ed) as the beginning of the ejaculatory duct. Actually the cavity continues into a straight canal, the ejaculatory duct (de) proper, running posteroventrally and opening into the male atrium on the ventral side of the penis papilla, its orifice generally being nearer to the base than to the lobular tip of the papilla. North American Triclads 21 5 b 0.5mm \ | ! ph m vd vs de am ode pp- gp bd t Fic. 5. Phagocata crenophila, semidiagrammatic view of copulatory complex in sagittal section. am, male atrium; b, copulatory bursa; bd, bursa duct; de, ejaculatory duct; gp, gonopore; m, mouth; odc, common oviduct; ph, pharynx; pp, lobe of penis papilla; t, testis; ud, vas deferens; vs, seminal vesicle. The seminal vesicle and ejaculatory duct are lined with columnar epithelia, with little histological difference between them. The entire male tract from the common vas deferens to the mouth of the ejaculatory canal receives ducts of mesenchymal glands with a granular, faintly eosinophilic secretion. The gland ducts are particularly numerous at the level of the common vas deferens and the anterior part of the seminal vesicle. A coat of circular muscle fibers surrounds all canals and is very noticeable on the ejaculatory duct. The outer surface of the penis papilla is covered with a cuboidal epithelium which gradually diminishes in thickness toward the lobe of the papilla tip where it becomes thin and flattened. Below the epithelium are the usual layers of muscle fibers, a circular layer adjoining the epithelium and a deeper layer of longitudinal muscles. Only the lobe at the tip of the papilla is provided with dorsoventral muscle fibers. The male atrium (am), which in general repeats the shape of the penis papilla, has a lining of cuboidal to columnar cells, with two muscle layers (circular and longitudinal) underlying it. The atrium opens to the outside at the genital pore (gp). On its dorsal side, a certain distance from the gonopore, the atrium receives the opening of the common oviduct (odc). The two oviducts, which in the anterior part of the body accompany the ventral nerve cords, ascend at the level of the copulatory complex toward the dorsal side and unite in the space between the male atrium and the bursal canal, forming the common oviduct (odc). The end 22 Proceedings of the Biological Society of Washington parts of the paired oviducts and the common oviduct connect with many ducts of mesenchymal glands with highly eosinophilic secretion, the so-called “shell glands.” The copulatory bursa (b) is a large sac lying between the pharyngeal pouch and the penis bulb, of variable size and shape and easily deformed by the pharynx in preserved specimens. Its outlet, the bursal canal or duct (bd) proceeds posteriorly above the atrium and behind it arches ventrally toward the gonopore. The diameter of its lumen gradually increases from the bursa posteriorly, then constricts again when ap- proaching the gonopore. The cellular lining of the bursal duct consists of tall columnar cells, apparently of a glandular (apocrine) nature. There is no conspicuous histological difference between the narrow anterior and the wider posterior parts of the duct. Both parts are coated with two muscle layers, one with circular, the other with longitudinal fibers. Distribution and ecology: Carpenter (1969: 280) collected Phagocata crenophila in five localities in Logan Canyon, Cache County, Utah, in cold mountain springs and streams. The species occurred in these localities together with Polycelis coronata (Girard). I observed sexually mature samples of P. crenophila in the following localities: Visitors Center of Eldorado National Forest, El Dorado County, California, located near the south shore of Lake Tahoe. Seepage springs on the grounds of the Center, with clear water, 7.8° C, pH 6.5. Current moderate, bottom mud with stones. Under stones and pieces of wood about 20 specimens, about half of them mature or semimature, taken 21 and 22 June 1967 and 11 July 1968. Tahoe City, Placer County, California (near northwest shore of Lake Tahoe). Overflow of spring furnishing water to the local California Fish and Game facility. Water clear, current fast, bottom with stones and algal growth. Collected 6 specimens on 23 September 1967, one of them sexually mature (water temperature, 6.7° C, pH 6.8); many animals taken 10 July 1968, both mature nad immature ones (tempera- ture 6.2° C, pH 6.6, oxygen 10 ppm). The population of this locality has the flared anterior end shown in figures 3B and 4e. Besides this species, Polycelis sp. occurred, in smaller numbers. Both Phagocata and Polycelis were also attracted to liver bait left in the stream over night. At my visit to Logan, Utah, on 27 September 1967, Mr. Jerry H. Carpenter kindly took me to two of his collecting places where I could observe P. crenophila and Polycelis coronata: Ricks Spring, Cache County, Utah, on U.S. Highway 89, 16.6 road miles east of Utah State University in Logan; and Spring Hollow Stream, Cache County, off U.S. 89, about 5 miles east of Logan. Immature planarians resembling P. crenophila were collected also in other localities under similar environmental conditions. Although no final identification could be made, they probably belonged to that species: North American Triclads 23 Sagehen Creek Biological Station of the University of California, Nevada County, California (north of Truckee, off State Road 89, close to the Sierra County line). Spring furnishing water to the Station, with clear, fast water, temperature 6.5° C, pH 7.5, oxygen 10 ppm. Under stones several specimens with wide heads, none mature, collected 10 July 1968. Besides these, more numerous Polycelis sp. Charleston Park, Toiyabe National Forest, Clark County, Nevada (west-northwest of Las Vegas). Below waterfall at end of Little Falls Trail (off Kyle Canyon Road). Water clear, 8.2° C, pH 8.8. Under stones many immature specimens collected by Timothy F. Hicks and James F. Leser, 6 July 1967. Several animals were placed in a culture kept at 14° C but did not mature and gradually died off without any having fissioned or encysted. The last specimen disintegrated 31 October 1968 (after almost 16 months). Phagocata crenophila seems to be widely distributed in the western mountains, both the Rocky Mountains and the Sierra Nevada, which have so far been little studied with regard to planarians. The species appears to be confined to cold springs and streams at high altitudes, often in the company of a species of Polycelis. Phagocata tahoena Kawakatsu Synonym: Phagocata nivea tahoena Kawakatsu, 1968: 6. Phagocata tahoena was described by M. Kawakatsu (1968) from preserved specimens collected during 1962 and 1963 in the course of the Lake Tahoe Fisheries Study carried out by the California and Nevada Fish and Game Departments. The animals had been taken in bottom samples brought up by an Ekman dredge from depths of 15 to 1632 feet. Concerning the appearance of the species in life we have only the brief description given by one of the collectors (Mr. Ted C. Frantz), quoted by Kawakatsu (1968: 6, footnote), indicating that the color of the two species occurring in Lake Tahoe varied from dark brown to translucent pink. I visited Lake Tahoe in 1967 and attempted to collect the species with a trawling dredge at a depth of about 40 m off the south shore of the lake near Al Tahoe (El Dorado County, Cali- fornia) in the company of Mr. Phillip H. Baker. Two specimens were obtained, one sexually mature, the other immature. In 1968, with the able assistance of the California Fish and Game Warden William F. Hart, I tried baiting specimens with beef liver. The method proved to be very successful, as I was able to collect numerous specimens. Baiting is often the most efficient method for the qualitative collecting of most species of freshwater triclads in both standing and running waters. The layout of the bait used in Lake Tahoe is illustrated in Figure 6. A small piece of beef liver was placed in a glass jar (7) which had a metal screw top with a number of perforations and a string attached to it. At a distance of several feet, the string was tied to a 24 Proceedings of the Biological Society of Washington ee Imm Fic. 6-7. 6. Baiting of planarians in deep water: b, canvas bag filled with stones to serve as anchor; f, float (plastic bottle); 7, jar with per- forated metal lid, containing bait. 7. Phagocata tahoena, from life. canvas bag (b) filled with stones which served as an anchor. A nylon line connected the bag to a float (f), in this case an inverted plastic bottle. With this arrangement, baits were exposed in three places, at depths of 40, 69, and 89 m. When the jars were raised to the surface after 24 hours, two of them contained many animals which had remained in them after feeding on the liver. The use of a perforated bait container for the collection of planarians had been recommended already by Sell (1916) but has hardly been in widespread practice. The effectiveness of this method was first demon- strated to me by the Powell Laboratories in Gladstone, Oregon. Liver, meat, pieces of fish, crushed snails, chopped-up earthworms, or other suitable food items may be used as bait. The perforations on the con- tainer keep larger predators or scavengers (e.g., crayfish) from reaching the bait. There are, however, planarian species that are not attracted by chemical food stimuli, such as Procotyla fluviatilis Leidy and some sub- terranean species. For these species the common collecting methods must be employed, i.e., individual searching in hiding places or the collection of debris, leaf litter, etc. in a large jar with water in which the animals will rise to the surface. External features (Fig. 7): The general aspect of Phagocata tahoena North American Triclads 25 in life has been fairly well preserved in some of the specimens studied by Kawakatsu (1968) and illustrated on his plate 1, figures A-C. Living specimens are generally 6 mm long.and about 1 mm wide, although some of Kawakatsu’s samples were considerably larger, measuring up to 12 mm in length in the preserved state. The animals are plump, with a large dorsoventral diameter. The head in life is rounded rather than truncate, lacking any lateral (auricular) protuberances. In the center of the head margin a small bluntly pointed projection may be seen when the animal is in gliding locomotion. There is no constriction or neck behind the head, as the lateral margins of the body run almost parallel up to the post-pharyngeal region where they begin to converge toward the more or less pointed posterior end. Eyes are two, separated from each other by a distance which amounts to about one-third, or slightly less, of the width of the head at the level of the eyes. The distance of each eye from the lateral margin is somewhat smaller than that from the frontal margin. The pharynx is inserted at the middle of the body and its length measures one-eighth to one-sixth the body length. The color of the dorsal side is a light hue of grayish brown, diminishing in darkness toward the margins of the body. The two small areas adjoining the eyes and a crescent-shaped rim along the frontal border are free of pigment. The ventral surface is unpigmented, white. In quiet gliding movement, the central part of the frontal margin is held somewhat elevated. Anatomy: The distinctive anatomical characteristics of Phagocata tahoena have been described by Kawakatsu (1968). I wish to add here only some details concerning glandular and sensory structures of the species. The submarginal adhesive zone (Fig. 8, az) is remarkably feebly developed, consisting of a very narrow strip of epithelial cells paralleling the lateral margins of the body, infranucleate,* and lacking rhabdite inclusions. These cells are perforated by few gland ducts con- taining an eosinophilic secretion. In the head region, along the frontal margin, the adhesive zone widens moderately, without forming a distinct adhesive organ. The auricular sense organs (Fig. 8, aw) consist of a pair of bands of modified epithelium, one on each side of the head, extending along the very edge of the frontal margin and encroaching to some extent on the dorsal surface. The epithelium is ciliated, infranucleate, but contains rhabdites similar in size to those of the normal surface epithelium. Each organ is separated from the submarginal adhesive zone by a narrow strip of normal nucleated epithelium. There is a wide gap between the two 4This term (“infranucléé”) was first introduced by Codreanu (1943: 138) for epithelia, rather commonly observed in the various classes of the Platyhelminthes, in which the nuclei with some cytoplasm have descended into a deeper layer. Such epithelia are designated as “‘eingesenkt’”” by German authors, an expression which is frequently translated as “‘insunk.’ “Infranucleate’” is a better descriptive term for this condition. 26 Proceedings of the Biological Society of Washington Fic. 8-9. Phagocata tahoena, sagittal sections through anterior end. 8. Auricular sense organ. 9. Subterminal organ. au. auricular sense organ (ciliated infranucleate epithelium, rhabdites present); az, submarginal adhesive zone; cg, cerebral ganglion; ne, normal surface epithelium; so, subterminal organ (infranucleate epithelium lacking rhabdites). 300. North American Triclads oF auricular organs in the center region of the frontal margin. Another small field of possibly sensory epithelium is seen in the very midline of the head on the ventral side immediately behind the adhesive zone (Fig. 9, so). This field, which we may call the subterminal organ, has a well-ciliated, infranucleate epithelium lacking rhabdite inclusions. It may be homologous with a similar organ described above in Phagocata bulbosa. Regarding the anatomy of the reproductive system reference is made to Kawakatsu’s (1968: 7-10) description and illustrations. The features distinguishing P. tahoena from other members of the genus Phagocata are as follows: Testes numerous, extending posteriorly almost to the tail end, predominantly ventral but with some testes ascending dorsally in the mesenchymatic spaces between the branches of the intestine; penis with well-developed bulb and conical or finger-shaped papilla; lumen of penis wide, not clearly differentiated into a seminal vesicle and an ejaculatory duct, receiving many glandular canals which have penetrated the penial tissues from the surrounding mesenchyme; penis lumen opening into the male atrium at the tip of the penis papilla; vasa deferentia opening into penial cavity separately from the antero-lateral sides; ovovitelline ducts ascending dorsally at the level of the copulatory complex, the left one passing through the space between the wall of the male atrium and the duct of the copulatory bursa, uniting with the right one above the atrium, the common oviduct thus formed opening into the posterior part of the male atrium (with regard to this situation, Kawakatsu’s figure 2 is misleading); copulatory bursa and its duct both lined with a very thick epithelium, the bursal duct displaced to the left of the midline, showing no histological division into an anterior canal and a posterior “vagina.” Distribution: Kawakatsu’s material was obtained from 40 bottom samples collected by an Ekman dredge at depths of 15 to 1632 feet in various sections of Lake Tahoe (Placer and El Dorado Counties, Cali- fornia; and Washoe and Douglas Counties, Nevada). My own collections were all made in the southern part of the lake: Off south shore of Lake Tahoe, near Al Tahoe, El] Dorado County, California. Two specimens collected by trawling dredge at about 40 m depth, one of them sexually mature, 23 September 1967. Emerald Bay of Lake Tahoe, E] Dorado County, California, on liver bait exposed for 24 hours at depth of about 40 m. About half a dozen specimens collected 12 July 1968. Off Pope Beach, El Dorado County, California, on liver bait left 24 hours at a depth of 89 m, very many (certainly over 100) specimens, both sexually mature and young, 12 July 1968. So far Lake Tahoe is the only known locality of Phagocata tahoena. However, the species may be more widely distributed in lakes of the Sierra Nevada mountain range which is still very little explored with regard to turbellarians. 28 Proceedings of the Biological Society of Washington Fic. 10. Phagocata tahoena, part of transversal section. g, gregarine parasites in mesenchyme; i, intestine; t, testis. 114. Taxonomic position: Phagocata tahoena has been described by Kawakatsu as a subspecies of P. nivea Kenk (1953). There are, how- ever, sufficient morphological differences between these two forms to justify giving tahoena the rank of a separate species. Of external char- acters, the plumpness of the body is quite remarkable, the rounded anterior end (truncate in P. nivea), the position of the eyes, and the presence of body pigment differ from the corresponding features observed in P. nivea. Among the anatomical characters, the structure of the penis, in particular the opening of the penis lumen at the tip of the penis papilla, separates P. tahoena from P. nivea (in which the penis lumen opens on the ventral side of the papilla). There is hardly any close taxonomic relationship between the two species. From an evolutionary standpoint it would also seem to be somewhat paradoxical to have an unpigmented surface species (P. nivea) closely related to a pigmented subspecies inhabiting the bottom of a deep lake. P. tahoena is, there- fore, here considered to be a species separate from P. nivea. Parasites: Some of the specimens of Phagocata tahoena examined in serial sections were infested with gregarines either encysted in the mesenchyme or more rarely free in the intestinal cavity. The cysts located in the mesenchyme were more or less spherical or ovoidal masses, each containing a pair of gametocytes (Fig. 10), similar to those de- scribed and illustrated by Holmquist (1967, fig. 7f-g) from Dendro- North American Triclads 29 11 A B e Fic. 11. Phagocata oregonensis, anterior end. A, normal two-eyed specimen; B and C, specimens with accessory eyes. _ 0.5mm bd ph m b pl vd am odc gp Fic. 12. P. oregonensis, copulatory apparatus in sagittal section. am, male atrium; b, copulatory bursa; bd, bursa duct; gp, gonopore; m, mouth; odc, common oviduct; ph, pharynx; pl, penis lumen; vd, vas deferens. coelopsis piriformis. No stages of gamete or spore formation were ob- served. Phagocata oregonensis Hyman, 1963 A brief description and rather sketchy illustrations of this species were given by Hyman (1963: 2-4), based mainly on the study of preserved specimens. No further mention has been made of the species in the literature, except for its inclusion in an analytical key to the American representatives of the genus Phagocata by Carpenter (1969: 281) and the presentation of an outline drawing of the anterior end in a recent paper (Kenk, 1969: 550). 30 Proceedings of the Biological Society of Washington External features (Fig. 1C and 11): This is an unpigmented, white, species. Mature specimens are up to 8 mm long and 1.5 mm wide. The head is truncate, with a somewhat waved frontal margin with slightly protuberant central and lateral portions (this condition is exaggerated in Hyman’s figure, 1963: 3, fig. 1). The lateral or auricular edges of the head are rounded, protruding very gently laterally so that an in- distinct necklike constriction appears behind them when the animal is in gliding motion. Behind the neck, the lateral margins of the body gradually widen up to the level of the pharynx, then converge again to meet at the more or less pointed posterior end. The root of the pharynx lies somewhat anterior to the middle of the body, and its length amounts to one-fifth the body length. The copulatory apparatus occupies less than half the length of the post-pharyngeal region. The two principal eyes are situated close together, their distance from each other amounting to one-fifth to one-fourth the width of the head and the distance of each eye from the lateral margin being about equal that from the frontal margin. Supernumerary eyes are frequently observed, particularly in “old” individuals that have been kept in a culture for several months. Individual eyes may appear lateral or posterior to the principal eyes, either on one or on both sides (Fig. 11). It is well known that planarians with one pair of principal eyes fre- quently develop accessory eye spots and that the occurrence and number of these increases with the advancing age of the animals (Balazs, 1962). The intestinal system begins a short distance behind the level of the eyes; there is no diverticulum or extension of the anterior intestinal ramus reaching forward between the eyes to a preocular level, such as is seen in some other white planarians with truncate head (e.g. Planaria occulta, see fig la in Kenk, 1969: 550). The anterior ramus bears 5-6 lateral branches, each posterior ramus 13-17 branches. Reproductive system (Fig. 12): The zone of the ventrally located testes extends on either side from the level of the second intestinal side branch to the posterior end (it does not end in the region of the copulatory complex as Hyman indicates). The ovaries, ovovitelline ducts, and yolk glands present no peculiarities. The copulatory apparatus (Fig. 12) conforms with the design typical of the genus Phagocata. The genital aperture or gonopore (gp) leads dorsally into the duct of the copulatory bursa (bd) and anteriorly into the male atrium (am); there is no common atrium developed. The penis has a bulb of moderate size with generally concentrically arranged muscle fibers and a conical, pointed papilla. The lumen of the penis (pl), situated in the posterior part of the bulb and traversing the length of the papilla, is a uniform cavity not differentiated into a seminal vesicle and an ejaculatory duct. It gradually narrows posteriorly and opens at the tip of the papilla. This cavity is lined with an epithelium of tall cells of glandular nature and is at least partially filled with the secretion produced by these cells. The outer cover of the papilla and the lining of the atrial cavity consist North American Triclads 31 of flattened to cuboidal cells, below which there is a layer of circular muscle fibers and a deeper layer of longitudinal muscles. The penial cavity also has a thin coating of circular and a more pronounced layer of longitudinal fibers. The two sperm ducts or vasa deferentia (vd), which at the sides of the pharynx expand to form the tortuous spermiductal vesicles, proceed posteriorly to the level of the male atrium, then turn forward again, enter the penis bulb laterally, and open into the penis lumen separately but close together. Hyman’s material on which she based her description of the anatomy of P. oregonensis was apparently defective since she states that she could not trace the sperm ducts into the penis bulb and assumed that they unite at the anterior margin of the bulb to form a common sperm duct which she interprets as an ejaculatory duct. My slides show clearly that each sperm duct, after forming the backward loop on the side of the atrium, enters the bulb and opens into the wide penis lumen independently. The lumen itself apparently corresponds to a combination of an anterior seminal vesicle and a distal ejaculatory duct without showing either a morphological or a histological differentia- tion into these two portions. The two ovovitelline ducts unite in the usual way in the space be- tween the dorsal wall of the atrium and the duct of the copulatory bursa. The common oviduct (odc) opens into the male atrium. Both the end parts of the paired oviducts and the common oviduct are pro- vided with eosinophilic gland ducts which penetrate their epithelia from the surrounding mesenchyme. The copulatory bursa (b) and its canal (bd) show no special differ- entiations. The canal or stalk gradually widens as it bends ventrally toward the gonopore, slightly left of the midline. The entire canal has two muscular coast, a circular and a longitudinal one. In the widened posterior part of the bursa stalk the circular layer gradually increases in thickness producing a terminal “vagina” without a sharp demarcation. Distribution: Hyman’s type material of Phagocata oregonensis was “from the vicinity of Crystal Springs in the Eastmoreland District of Portland,” Oregon. I visited this area on 12 September 1967 in the company of Mr. Richard Whitten of the Powell Laboratories of Glad- stone, Oregon, and collected numerous specimens on liver bait in Wood- bury Spring (Eastmoreland District of Portland, Multnomah County), a small limnocrene with almost stagnant, clear water. The water tem- perature was 11.2° C, the pH 6.3. The species was accompanied by two other planarian species, Dendrocoelopsis vaginata Hyman and Polycelis sp. Laboratory cultures: P. oregonensis was kept in an aquarium with spring water at 14° C for 13 months, fed alternately beef liver and live Tubifex worms (both were freely taken). Some specimens were sexually mature but did not deposit any egg capsules, nor did fission occur during the entire period. 32 Proceedings of the Biological Society of Washington LITERATURE CITED BatAzs, A. 1962. Uber die Rolle des Lebensalters in der Morphogenese der akzessorischen Augen des Strudelwurms Dugesia lugubris. Zeitschr. fiir Alternsforschung, 16: 53-41. CARPENTER, J. H. 1969. A new planarian from Utah, Phagocata crenophila n. sp. (Turbellaria, Tricladida). Trans. Amer. Microscop. Soc. 88: 274-281. CopreANnu, R. 1943. Dendrocoelum (sous-genre Dendrocoelides) clu- janum n. sp., nouveau Triclade souterrain de Transylvanie. Analele Acad. Romane, Mem. Sect. Stiint., Ser. III, 18 (7). 24 p., 4 pl. HatpEeMaAn, S. S. 1840. Supplement to number one of “A Monograph of the Limniades, or Freshwater Univalve Shells of North America.” 3 p. Hortmeuist, C. 1967. Dendrocoelopsis piriformis (Turbellaria Tri- cladida) and its parasites from Northern Alaska. Arch. fiir Hydrobiol., 62: 453-466, 6 pl., 1 folding table. Hyman, L. H. 1937. Studies on the morphology, taxonomy, and dis- tribution of North American triclad Turbellaria. VII. The two species confused under the name Phagocata gracilis, the validity of the generic name Phagocata Leidy 1847, and its priority over Fonticola Komarek 1926. Trans. Amer. Microscop. Soc., 56: 298-310. 1963. North American triclad Turbellaria. 16. Fresh-water planarians from the vicinity of Portland, Oregon. Amer. Mus. Novitates, no. 2123. 5 p. IcuHikAWA, A., AND M. KAwakatsu. 1962a. Phagocata iwamai, a new freshwater planarian, from Hokkaido. Annot. Zool. Japonenses, 35: 38-46. 1962b. Phagocata teshirogii, a new freshwater planarian, from the Tohoku region in Honshi. Annot. Zool. Japonenses, 35: 112-120. Kawakatsu, M. 1968. North American triclad Turbellaria. 17. Fresh- water planarians from Lake Tahoe. Proc. U.S. National Mus., 124 (3638). 21 p., 2 pl. Kenxk, R. 1935. Studies on Virginian triclads. Jour. Elisha Mitchell Scient. Soc., 51: 79-133, pl. 45. 1953. The freshwater triclads (Turbellaria) of Alaska. Proc. U.S. National Mus., 103: 163-186, pl. 6-8. 1969. Freshwater triclads (Turbellaria) of North America. I. The genus Planaria. Proc. Biol. Soc. Washington, 82: 539-558. Lemy, J. 1847. Description and anatomy of a new and curious sub- genus of Planaria. Proc. Acad. Nat. Sci. Philadelphia, 3: 248-251. Livanoy, N. A., AND Z. I. ZAsusova. 1940. Planarii bassema Telets- North American Triclads 33 kogo ozera i novye dannye o nekotorykh drugikh sibirskikh vidakh (Paludicoles du lac Teletzkoé et nouvelles données sur quelques formes sibériennes). Trudy Obshch. Estestvoisp. pri Kazan. Univ., 56: 83-159. MU.ier, Orno Frmericus. 1776. Zoologiae danicae prodromus, seu OkUGAWA, SELL, H. Animalium Daniae et Norvegiae indigenarum characteres, nomina, et synonyma imprimis popularium. xxxii + 282 p. Havniae. K. 1939. Probursalia (Tricladida-Paludicola) of Man- choukuo. Annot. Zool. Japonenses, 18: 155-165. 1956. A new fresh-water planaria Phagocata kawakatsui sp. nov. from Kyoto Prefecture. Bull. Kyoto Gakugei Univ., (B) 8: 19-22, 1 pl. 1916. Zur Biologie der Planarien. I. Blatter fiir Aquarien- und Terrarienkunde, 27: 315-316. 34 Proceedings of the Biological Society of Washington 74.06'7 3 Vol. 83, No. 3, pp. 35-50 27 May 1970 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON THREE NEW SPECIES OF CALLIANASSID SHRIMP (DECAPODA, THALASSINIDEA) FROM THE << coy WESTERN ATLANTIC! Ja laa f JUN 1 19/70 By Tuomas A. BirFraR School of Marine and Atmospheric Sciences, LIBRARIES University of Miami ~*~ Three new species of Callianassa were discovered during a recent study of the callianassid fauna of South Florida. Since the comprehensive publication will not appear for some time, it was deemed best to publish the descriptions of the new species separately. Specimens of the new species described below were first collected by the author in South Florida in 1967, but these specimens were later destroyed. Fortunately, additional ma- terial was available from other sources. I am most grateful to Mr. William Lyons of the Florida Board of Conservation Marine Laboratory, St. Petersburg (FBCML), for the loan of specimens of Callianassa trilobata dredged from Tampa Bay. I am indebted to Dr. Raymond B. Manning, Smithsonian In- stitution (USNM), for his assistance on a number of occasions and for the loans of specimens and to Dr. Herbert W. Levi of the Museum of Comparative Zoology (MCZ) for the loan of material containing specimens of all three new species. The material at the National Museum of Natural History was examined during a trip sponsored by the Office of Academic Affairs of the Smithsonian Institution. During a 1 Contribution No. 1184 from the University of Miami, Rosenstiel School of Marine and Atmospheric Sciences. This work was supported in part by grant GB-7075X from the National Science Foundation. This paper is part of a thesis submitted in partial fulfillment of the requirements for the Degree of Master of Science, University of Miami, Coral Gables, Florida. 3—Proc. Biot. Soc. WasH., Vou. 83, 1970 (35) 36 Proceedings of the Biological Society of Washington portion of the study, the author was supported by a National Science Foundation traineeship. Measurements given for specimens refer to total length. The term P3 is an abbreviation for third pereiopod. Other legs are referred to similarly. Callianassa trilobata new species Figure 1 Holotype: 18, 76 mm; Florida, Tampa Bay, off Pinellas Point; 18 January 1968; R. Stokes. EJ-68-2 (FBCML). Paratypes: 19, 66 mm; Florida, Miami, Virginia Key, Bear Cut beach; 12 April 1959; R. Manning. USNM. 1 juvenile, 35.5 mm; Florida, Biscayne Bay; March—April 1937; H. L. Clark. MCZ 9697. 22 2, 77-80.5 mm; Florida, Tampa Bay, SW of Bayboro Harbor; 2—4 m; 16-17 January 1968; R. Stokes. EJ-68-1 (FBCML). 1 damaged @; Florida, Tampa Bay, off Bayboro Harbor; 2.5—3.5 m; 22, February 1968; Stokes and Lyons. EJ-68-7 (FBCML). 1g, 60 mm, 1 damaged ¢, 82 2, 39.5-77.5 mm; Florida, Tampa Bay, off Pinellas Point; 2-3 m; 18 January 1968; R. Stokes. EJ-68-2 (FBCML). Diagnosis: Front with three low, subtriangular projections. Telson trilobed laterally, much wider than long. Large cheliped with triangular ventral keel on merus. Eye of fresh specimen with white spot in center of pigmented area. Description: Rostrum and lateral projections short rounded triangles. Rostrum 0.2 times length of eyestalks. Lateral projections situated be- tween eyestalks and base of antennal peduncle. Eyestalks flattened, not reaching tip of first antennular segment, length 1.45-2.0 times width; lateral margins straight, directed slightly inward; distal margins straight, more obliquely directed, lacking distal projections; pigmented area central, slightly convex, elongate longitudinally, covering about 0.5 exposed dorsal surface of eyestalk; cornea small, subterminal, covering about 0.2 pigmented area. Dorsal carapace posterior to cervical groove with median prominence set off by oblique ridges. Third segment of antennule 1-1.4 times length of second segment. Third antennular segment 0.6—0.9 times length of fourth antennal seg- ment. Fourth antennal segment 1.25—1.85 times length of fifth segment. Antennal flagellum 2.5-4.3 times length of antennular. Second segment of antennule reaching 0.6 length of fourth antennal segment. Third antennular segment to 0.25 fifth antennal segment. Second and third segments of antennule with dense rows of elongate setae on their lateral surfaces. Third segment of antenna with numerous elongate setae laterally. Incisor process of mandible with 16 acute teeth of varying sizes Three New Decapods 37 Fic. 1. Callianassa trilobata: a, mandible; b, first maxilla; c, second maxilla; d, first maxilliped; e, second maxilliped; f, third maxilliped; g, female large cheliped; h, male large cheliped; i, small cheliped; j, dorsum; k, P3; 1, tail fan; m, male first pleopod; n, female first pleopod; 0, male second pleopod; p, female second pleopod. 38 Proceedings of the Biological Society of Washington extending along entire margin, molar process bifurcate ventrally, tips acute, remainder of process entire. First maxilla with wide basal endite, slightly expanded distally, margin rounded; coxal endite with narrow, rounded distal lobe. Epipod of first maxilliped triangular, broadly rounded laterally; basal endite with straight lateral margins, rounded tip; exopod not distinctly bilobed, slight indentation on inner margin. Second maxilliped with exopod longer (1.5) than first segment of endopod. Merus and ischium of third maxilliped wide, length of both segments 1.5—2.0 times greatest width, ischium lacking spinous ridge on internal surface except for 3-4 proximally-directed free spines at proximal end; propodus expanded, width 0.73 times width of merus, with small rounded lobe distoventrally; width of dactylus 0.25 times width of propodus. Ischium of large cheliped of male slightly expanded distoventrally, ventral margin with series of 23 rounded denticles, smallest proximally. Length of merus 1.6-1.8 times greatest width, proximal end of dorsal margin raised, with inconspicuous rounded denticles; ventral margin expanded, forming triangular keel, concave and entire proximally, re- mainder of margin with about 17 rounded serrations. Length of carpus 1-1.1 times length of merus, carpus width 0.75—-0.95 times length, wider medially due to convex proximal margin; ventral margin of carpus in- conspicuously serrate on inner surface, serrations separated by short fringes of setae. Length of palm 1.2—1.4 times length of carpus, palm length 1.35-1.6 times width, ventral margin weakly serrate on inner surface. Propodal finger blunt, curving slightly inward, cutting edge with three acute, triangular teeth proximally, two moderately strong rounded teeth medially, six low rounded teeth distally; series of blisters running along internal surface of finger. Length of dactylus 0.5 times length of palm, cutting edge with two irregular, truncate teeth proximally, series of 10 rounded teeth distally. Internal surface of dactylus with series of blisters interspersed with tufts of hair, dorsal margin similar, blisters more acute. Large cheliped sexually dimorphic. Ventral margin of ischium of cheliped of female with several inconspicuous denticles. Length of merus 1.5—2.0 times width, dorsal margin convex, ventral keel present but not as strong as in male, fewer denticles on margin distally. Length of carpus 0.9-1.1 times length of merus, length of carpus 0.9-1.3 times width, ventral margin entire. Palm longer (1.05-1.55) than carpus, longer than wide (1.02-1.4), ventral margin entire. Propodal finger blistered on internal surface below cutting edge, cutting edge with proximal series of small triangular teeth, strong triangular tooth proximal to midpoint of edge, distal series of approximately 10 small rounded teeth decreasing in size toward tip. Small multilobed projection at base of notch between fingers. Dactylus narrow, curving inward, length 0.65-0.95 times length of palm, low serrate ridge running transversely across cutting edge proximally, remainder of edge straight, inconspicuously Three New Decapods 39 Taste 1. Branchial Formula of Callianassa trilobata. Maxillipeds Walking legs Gill type 1] 2 3 1 2) 3 4 5 Pleurobranchs = se = ee a Eas bs Zs Arthrobranchs = 2 2, 2 2 2 2, - Podobranchs = = = a Eb she 2 mt * A single rudimentary gill may be present. serrate; internal surface with scattered blisters, dorsal margin lacking blisters found in male. Ischium of small cheliped lacking ventral serrations. Merus with simple convex dorsal margin, ventral margin lacking keel or denticles. Carpus narrow, length 2.5 times width. Palm shorter than carpus, narrowing distally. Propodal finger long and thin, cutting edge entire. Dactylus straight, narrow, cutting edge entire. Propodus of third pereiopod (P3) bilobed, posterior lobe wide, rounded, ventral margin concave. P4 subchelate, P5 chelate. Branchial formula shown in Table 1, second maxilliped with rudimentary arthro- branch. Abdomen elongate, well developed. First abdominal somite divided by transverse dorsal ridge, anterior 0.2 of somite narrow, rounded an- teriorly, posterior area widening toward posterior margin. Well de- veloped fringe of hair on posterolateral margins of third and fourth somites, length of fringes 0.25 times somite width. Fifth somite with similar fringe midlaterally. Lateral margin of sixth somite distinctly bi- lobed by strong transverse groove separating anterior three quarters from posterior quarter; lateral margin of anterior lobe inconspicuously bilobed with indentation occurring one-third length of lobe from anterior margin; posterior margin of somite indented. Telson distinctly wider (1.14—1.6) than long, lateral margins trilobed; posterior margin concave centrally. First pleopod of male uniramous, two-segmented, distal segment with acute tip, rounded prominence distolaterally. Second pleopod of male biramous, exopod bladelike, acute at tip, 0.78 times length of first seg- ment of endopod. First segment of endopod bladelike, wider than exopod; second segment triangular with acute distal tip; third segment small, elongate, articulating laterally at junction of first and second segments. First pleopod of female uniramous, two-segmented, distal segment thick at base, bladelike distally. Second pleopod biramous, exopod bladelike, shorter than first segment of endopod. Endopod two- segmented, first segment bladelike, projecting slightly at distal tip and forming subchela with small, elongate second segment. Appendix interna of third through fifth pleopods subtriangular in outline, hooks located along lateral surface. 40 Proceedings of the Biological Society of Washington Endopod of uropod elongate oval, length 1.5-1.95 times width, narrowing distally, fringe of fine setae along distal half of posterior margin, denser fringe at distal tip. Exopod of uropod slightly longer than endopod, lower exopodal plate longest posteriorly, with dense fringe of setae. Single basal spine present. Suture distinct. Upper exopodal plate 0.6 times length of lower plate. Color; Entire animal white. Eye of fresh specimen with white spot in center of pigmented area. Dimensions of Holotype: EJ-68-2 (FBCML), ¢. Total length 76 mm, carapace length 17 mm. Telson width 5.0 mm, length 4.4 mm. Width of endopod of uropod 3.5 mm, length 5.9 mm. Large cheliped: width of merus 5.9 mm, length 10.6 mm; width of carpus 9.0 mm, length 11.9 mm; width of propodus 8.9 mm, length 14.2 mm; length of dactylus 7.25 mm. Known Range: Miami and Tampa Bay, Florida. Discussion: There are similarities between the cheliped of this species and that of C. audax de Man, but the species can be distinguished by the form of the telson, shape of the eyestalks and endopod of the uropod, and the keel on the merus of the large cheliped in C. trilobata. Remarks: Two specimens are known which, because they exhibit characters usually associated with the male (form of the large cheliped and the pleopods), have been termed males and described as such. However, both specimens have male and female coxal apertures. The reason for this phenomenon is not yet apparent. Etymology: The name is derived from the lateral contours of the telson. Callianassa quadracuta new species Figure 2 Holotype: 16, 68 mm; Venezuela, Cumana; 1859; J. P. Couthouy. MCZ 760. Paratypes: 16, 75 mm, 22 9, 27-39 mm, | ovigerous 2, 62 mm; Venezuela, Cumana; 1859; J. P. Couthouy. MCZ 760. 16, 69 mm, 19, 74.5 mm, 2 damaged 2 9; Venezuela, Cumana; 1859; J. P. Couthouy. MCZ 732. 1 ovigerous 2, 70 mm; Venezuela, Cumana; 1859; L. Agassiz. MCZ 12872. | Diagnosis: Chelipeds of first walking legs subequal, carpus with two acute distodorsal and two acute distoventral projections, distodorsal corner of palm an acute projection. Endopod of uropod triangular, extending well beyond distal margin of telson. Telson distinctly wider than long. Dactylus of third maxilliped expanded distally, width 0.75—0.9 times width of propodus. Description: Carapace with two weak ridges dorsally, one on either side of midline running parallel to body axis, terminating before reaching front. Short median ridge narrowing anteriorly, continuous with rostrum. Three New Decapods 4] Fic. 2. Callianassa quadracuta: a, mandible; b, first maxilla; c, second maxilla; d, first maxilliped; e, second maxilliped; f, third maxilliped; g, P3; h, small cheliped; i, large cheliped; j, dorsum; k, tail fan; 1, male first pleopod; m, male second pleopod; n, female first pleopod; 0, female second pleopod. 42. Proceedings of the Biological Society of Washington Convex area as wide as eyestalk located over base of eye, narrowing posteriorly. Posterior area of carapace with ridge along dorsal midline. Rostrum a low acute triangle extending 0.1-0.2 times length of eye- stalks, slightly depressed. Lateral projections low rounded triangles at external margin of eyestalks. Eyestalks flattened, extending beyond distal margin of first antennular segment, length 1.8—2.8 times width; lateral margins indistinctly concave; distal margin straight, median projection denticulate at tip; cornea lateral, subterminal, covering about 0.1 exposed dorsal surface of eyestalks. Length of third antennular segment 0.75-0.95 times length of second segment. Third antennular segment 0.6—0.8 times length of fourth antennal segment. Fourth antennal segment 1.06—-1.5 times length of fifth segment. Antennal flagellum 2.54.35 times length of antennular. Second antennular segment extending to 0.9 length of fourth antennal segment. Third antennular segment to 0.75 fifth antennal segment. Second and third antennular segments with dense rows of elongate setae laterally. Basis of mandible wide, first segment of palp shorter than second segment, incisor process with 10 truncate teeth, some inconspicuously serrate, molar process entire, bifurcate ventrally. Basal endite of first maxilla slightly expanded distally, distal margin straight; coxal endite with indication of distal lobe. Epipod of first maxilliped triangular, narrowly rounded distally; basal endite rounded; internal margin of exopod indented. Exopod of second maxilliped longer (1.3) than first segment of endopod. Merus and ischium of third maxilliped wide, length of both segments 1.55—-1.85 times greatest width of either segment, spinous ridge on ischium with seven free spines distally, distal-most largest, remainder of ridge with about eight denticles; propodus not expanded, width 0.6-0.7 times width of ischium; width of dactylus measured proximally 0.35 times width of propodus, 0.75-0.9 times if width of dactylus measured distally. Chelipeds of first pereiopods subequal. Ischium of left cheliped short, with 18 subacute denticles ventrally, longer spine near distal end. Merus with two inconspicuous denticles proximally on ventral margin, five denticles medially, length of merus 1.53-1.74 times width. Carpus shorter (0.8—0.9) than merus, length 0.65—0.85 times width, dorsal margin cristate proximally, divided on external surface distally forming two acute projections, one above the other, at distodorsal corner; ventral margin entire, secondary ridge just above margin on external surface, terminating distally as acute projection, second acute projection just above ventral-most articulation with palm. Palm longer (1—1.32) than wide, longer (1.6-1.96) than carpus; ventral margin of palm entire, rounded, dorsal margin cristate on proximal half, distal half with minute denticles, terminating distally as acute projection; distal margin of palm inconspicuously serrate along articulation with dactylus, slightly blistered behind notch between fingers. Series of raised tufts of hair on external Three New Decapods 43 surface of palm near proximoventral articulation. Propodal finger strong, blunt at tip, much shorter than dactylus, series of 15 rounded denticles proximally on cutting edge ending at median triangular tooth, remainder of cutting edge inconspicuously serrate. Weak ridge running along outer surface below cutting edge and for short distance on palm. Dactylus strong, length 0.6-0.75 times length of palm, blunt and curving inward distally; cutting edge entire on proximal third, serrate on remainder. Ischium of right cheliped as in left, ventral margin with 16 irregular denticles and single spine near distal end. Merus as in left, length 1.45— 1.8 times width. Carpus as in left with four acute projections distally, length 0.76 times width, length 0.85-0.95 times length of merus. Length of palm 1.17-1.52 times length of carpus, notably shorter and more oblique than palm of left cheliped, dorsal margin lacking distal serra- tions, ridge on propodal finger and palm more distinct. Propodal finger subacute distally, curving slightly inward, cutting edge with 10 irregular serrations. Dactylus slender, subacute distally, length 1.05-1.2 times length of palm, cutting edge with series of rounded denticles decreasing in size distally. For practical purposes, this cheliped termed small cheliped. Propodus of P3 with narrow rounded lobe posteriorly, anterior margin slightly convex, segment about as long as broad. P4 simple, P5 chelate. Branchial formula as in C. trilobata, all gills well developed, gill of second maxilliped with lamellae. First abdominal somite divided by transverse ridge; anterior quarter subrectangular, widening posteriorly; margins of posterior area diverging and continuing laterally almost to level of pleopod articulation before reaching posterior margin of somite. Third and fourth somites with well developed fringe of hair posterolaterally on dorsal surface. Fifth somite with well developed midlateral fringe. Lateral margin of sixth somite bilobed, anterior lobe distinctly wider than posterior, lateral margins convex; posterior lobe separated from anterior by lateral indentation, lacking well defined groove; posterior margin of somite with hint of indentation at midline, stronger indentation just beyond tufts of hair. Telson distinctly wider (1.46-1.76) than long; lateral margins with slight indentation in anterior third, remainder convex; posterior corners smoothly rounded, posterior margin with median convex projection. Dorsal surface of telson convex, distinct transverse ridge on proximal half interrupted medially by tuft of hair. First pleopod of male uniramous, two-segmented, distal segment elongate, rounded distally, slight indentation on internal margin. Second pleopod of male biramous, exopod bladelike, subacute at tip. Endopod two-segmented, proximal segment with pliable flap running along distal half of lateral margin, distal segment bladelike, rounded at tip. First pleopod of female uniramous, two-segmented, proximal segment extending horizontally beyond level of articulation of second seg- 44 Proceedings of the Biological Society of Washington ment; distal segment bladelike, thicker proximally, rounded at tip. Second pleopod of female biramous, peduncle extending horizontally beyond level of articulation of endopod; exopod bladelike, curving inward, longer than endopod; endopod bladelike, thicker proximally, with only one segment. Appendix interna of third through fifth pleopods subtriangular with hooks along lateral surface. Endopod of uropod triangular, length 1.2-1.7 times width, distal tip extending 2.1 times length of telson beyond posterior margin of telson; margins slightly irregular due to widely spaced setae, dense fringe of setae at distal tip. No median ridge. Exopod of uropod longer than endopod, upper exopodal plate with dense fringe of setae, posterodistal margin with two rows of approximately 14 strong setae. Suture distinct. Lower plate fringed with both hair and setae distally, strong short setae forming dorsal row; posterior corner with row of 12 strong spine- like setae. Ridge lacking, one strong basal spine present. Color: Thorax and chelipeds white, remainder unknown in fresh ma- terial, entire animal beige when preserved in alcohol. Dimensions of Holotype: MCZ 760, 6. Total length 68 mm, carapace length 17 mm. Telson width 5.5 mm, length 3.75 mm. Width of endopod of uropod 2.75 mm, length 4.5 mm. Left cheliped: width of merus 4.25 mm, length 6.85 mm; width of carpus 8.4 mm, length 5.9 mm; width of propodus 9.5 mm, length 10.9 mm; length of dactylus 6.6 mm. Right cheliped: width of merus 4.0 mm, length 6.4 mm; width of carpus 8.0 mm, length 6.1 mm; width of propodus 8.4 mm, length 8.5 mm; length of dactylus 10.0 mm. Known Range: SE Florida, Venezuela. Discussion: This species appears to be closely related to C. aequimana Baker from Australia and to C. novaebritanniae Borradaile and its variety which was described by de Man (1928). De Man separated the two species and the variety based on the telson morphology, size, morphology, of the ischium of the large cheliped, and several less distinctive characters. As Sakai (1966) noted, for practical purposes, the ridge on the dorsal surface of the telson is most useful. Callianassa quadracuta, like C. aequimana, has a notched ridge, while C. novae- britanniae has a continuous weak ridge, and the variety lacks any ridge. In addition, the acute distal projections on the carpus of the cheliped and the convex posterior border of the telson separate C. quadracuta from all these Pacific forms. Only one other species, Callianassa laticauda Otto, has been described as having a relatively well-developed gill on the second maxilliped (Gurey, 1944). The significance of the character is unknown since the branchial formula has been described for only a small number of species of Callianassa. No other characters relating C. laticauda and C. quadracuta are immediately apparent. Etymology: The name of the species is derived from the acute projections on the carpus of the chelipeds. Three New Decapods 45 Callianassa fragilis new species Figure 3 Holotype: 14, 27 mm; Puerto Rico, Punta Arenas, sand flat; 12 June 1946; H. G. Hewatt. USNM 82367. Paratypes: 1 damaged ¢; data as in holotype. 12, 16.5 mm; Antigua, Falmouth Bay, Black’s Point; 30 April 1959; Smithsonian Bredin Expedition 112-59. USNM. 26 6, 39-40 mm; Venezuela, Cumana; 1859; Capt. Couthouy. MCZ 760. Additional specimens have been collected around Miami and have been deposited in the collection at the University of Miami Marine Institute. Diagnosis: Total length less than 45 mm. Merus of large cheliped with proximal hook directed distally. Third segment of antennule ex- tending beyond distal end of fifth antennal segment. Telson shield- shaped, with posteromedian concavity. Endopod of uropod wide oval, extending beyond posterior margin of telson. Description: Rostrum and. lateral projections rounded triangles. Rostrum extending only 0.07—0.23 times length of eyestalks. Lateral projections just beyond margin of eyestalks, not extending as far an- teriorly as rostrum. Eyestalks elongate, extending almost to end of first antennular segment, length 2.2-2.7 times width; lateral margin more or less straight, directed at slight oblique angle inward; distal margin straight, more inwardly directed, with internal triangular projec- tion; pigmented area central, subterminal, covering about 0.05 exposed dorsal surface of eyestalks, irregular in shape. Length of third antennular segment 2.2-3.7 times length of second antennular segment. Third antennular segment 1.25-1.7 times length of fourth antennal segment. Fourth antennal segment 1.15-1.55 times length of fifth antennal segment. Antennal flagellum 2.75—4.2 times length of antennular. Second antennular segment extending to 0.55 length of fourth antennal segment. Third antennular segment to just beyond distal end of fifth antennal segment. Second and third antennular segments moderately fringed with setae on lateral surfaces. Mandible with 12 teeth in incisor process, teeth progressively larger toward ventral end, molar process with two strong ventral teeth and four smaller teeth dorsally. Basal endite of first maxilla expanded distally, rounded; coxal endite narrow, with small rounded distal lobe. Basal endite of first maxilliped convex distally; exopod squared distally, with indentation on internal margin; epipod lacks distal acute projection, narrowly rounded laterally. Exopod of second maxilliped only 0.75 times length of first segment of endopod. Merus and ischium of third maxilliped wide, length of segments 1.4—1.6 times greatest width, internal surface of ischium with 16 weak denticles; merus subrectangular, disto- ventral area a rounded corner; propodus only slightly expanded proxi- 46 Proceedings of the Biological Society of Washington Fic. 3. Callianassa fragilis: a, mandible; b, first maxilla; c, second maxilla; d, first maxilliped; e, second maxilliped; f, third maxilliped; g, P3; h, female large cheliped; i, male large cheliped; j, small cheliped; k, dorsum; 1, tail fan; m, female second pleopod; n, female first pleopod; o, male first pleopod. Three New Decapods AT mally, width 0.43 times width of merus; width of dactylus 0.4—0.67 times width of propodus. Ischium of large cheliped of male with five broad, inconspicuous serrations ventrally. Merus more than twice (2-2.2) as long as wide, proximoventral margin produced into distally directed hook, hook with 3-4 strong serrations ventrally, acute tip; remainder of ventral margin with 11 denticles separated by tufts of hair. Length of carpus 0.68—0.9 times length of merus, longer medially due to convex proximal margin; width of carpus 1.1—-1.4 times dorsal length, ventral margin entire. Palm longer (1.3-1.8) than carpus, length greater (1.15-1.3) than width, ventral margin entire. Propodal finger shorter than dactylus, cutting edge entire. Notch between fingers rounded, extending proxi- mally for a short distance. Dactylus 0.6—-0.82 times length of palm, truncate teeth proximally and distally with three or four rounded serra- tions on cutting edge of distal tooth, proximal tooth entire or serrate, triangular tooth medially. Tip of dactylus strongly curved, not hooked. Ischium and merus of female cheliped like those of male. Carpus longer than wide (1.14—1.22), length equalling length of merus, almost (0.9-0.95) as long as palm. Palm narrowing distally, slightly narrower than carpus. Propodal finger serrate along cutting edge, notch between fingers not extending proximally. Dactylus slightly curved, lacking strong dentition, cutting edge serrate. Ischium and merus of small cheliped lacking ventral denticles. Ventral margin of merus inconspicuously concave proximally, lacking hook. Carpus elongate, length 5-6 times width. Palm narrow, length 0.4 times length of carpus, notch not extending proximally. Propodal finger almost as long as dactylus, minute serrations on cutting edge. Dactylus straight, cutting edge entire. Propodus of P3 bilobed, dactylus with acute tip. P4 simple, P5 chelate. Branchial formula as in C. trilobata, second maxilliped lacking rudimentary gill. First abdominal somite short, lacking any transverse ridge. Third somite with tuft of hair in posterolateral corner. Fourth somite with midlateral tuft. Fifth somite with tuft of hair in anterior third of lateral margin. Lateral margin of sixth somite weakly bilobed, margins of anterior 0.75 slightly convex, fringed with fine hair, separated from posterior rounded lobes by indistinct groove; posterior margin indented medially. Telson shield-shaped, width 1.16 times length, median concavity in posterior margin; lateral margins convex, slight notch 0.25 total length from proximal end, remainder of margin smoothly rounded; tuft of elongate setae on dorsal surface set back from each posterior corner. First pleopod of female uniramous, two-segmented, distal segment bladelike, rounded at tip. Second pleopod biramous, exopod bladelike, rounded at tip, shorter than endopod (0.88); endopod similar, with transverse articulation 0.65 total length from proximal articulation. First 48 Proceedings of the Biological Society of Washington TaBLE 2. Morphological Characteristics of Callianassa atlantica and C. fragilis. Characteristic C. atlantica C. fragilis Telson posterior border acute median median indentation projection Eyestalk beyond lst antennu- shorter than Ist an- lar segment tennular segment Eyestalk projection slender, curving short, rounded 3rd antennular segment 3.6—5.3 2.2-3.7 2nd antennular segment large cheliped fingers about equal dactylus longer small cheliped carpus 3.6—4.0 4.5-5.1 length/width dactylus of 3rd slender wide maxilliped pleopod of male uniramous, two-segmented, distal segment elongate, subacute at tip. No indication of second pleopod. (Holotype lacking pleopods on first and second abdominal somites.) Appendix interna of third through fifth pleopods subtriangular, hooks only on small area of lateral surface near apex. Endopod of uropod wide oval, length 1.6 times width, extending 0.33 times length of telson beyond posterior margin of telson. Exopod of uropod extending just beyond endopod. Upper exopodal plate 0.8 times length of lower plate. Suture indistinct. Lower exopodal plate with dense distal fringe of setae, posterior corner with eight strong setae, anterior margin fringed with hair. No basal spine. Color: Transparent except for subcutaneous yellow-orange on dorsal abdominal surface and white chelipeds. Dimensions of Holotype: USNM 82367, ¢. Total length 27 mm, carapace length 6 mm. Telson width 2.25 mm, length 1.95 mm. Width of endopod of uropod 1.55 mm, length 2.5 mm. Large cheliped: width of merus 2.0 mm, length 4.25 mm; width of carpus 3.9 mm, length 2.9 mm; width of propodus 4.0 mm, length 5.25 mm; length of dactylus 4.3 mm. Known Range: SE Florida, Puerto Rico, Antigua, Venezuela. Discussion: Callianassa atlantica Rathbun is the only other western Atlantic species known which has a hook on the merus of the large cheliped. The two species appear to be closely related; both have an elongate third antennular segment, similar telson shapes, similar female pleopods, and both lack spinous frontal projections and male second pleopods. Table 2 shows several characters which can be used to separate the species. Three New Decapods 49 Remarks: This species is found in shallow burrows in sand, mud, or clay, where it lives in close proximity to burrows of other species of Callianassa or Upogebia. When excavating around burrows of other species, it is not uncommon to find specimens of C. fragilis floating in water that has seeped into the hole. The animal apparently is affected by surface tension and can escape only after a prolonged struggle. Etymology: The name reflects both the great ease with which the large cheliped becomes separated from the body and the small size of the species. LITERATURE CITED Gurney, R. 1944. The systematics of the crustacean genus Callianassa. Proc. zool. Soc. London, 114(5): 82-90, text figs. 1-19. Man, J. G. bE. 1928. A contribution to the knowledge of twenty-two species and three varieties of the genus Callianassa Leach. Capita zool., 2(6): 1-56, pls. 1-12. Saxal, K. 1966. On Callianassa (Callichirus) novaebritanniae Bor- radaile (Thalassinidea, Crustacea) from Japan. J. Fac. Agric. Kyushu Univ., 14(1): 161-171, 4 text figs. 50 Proceedings of the Biological Society of Washington b 14.0673 Vol. 83, No. 4, pp. 51-58 27 May 1970 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON A REVIEW OF THE APHODIINE BEETLES OF THE GALAPAGOS ISLANDS =< HSON] (COLEOPTERA: SCARABAEIDAE)! /~ Vy IN 70) By Oscar L. CarTwRIGHT ( FUN 1 97 Smithsonian Institution, Washington, D.C. » C/BRARIED The discovery of the genus Diastictus on Isla Wolf, confusion of names due to misidentification, and synonomy in the re- ported species of Ataenius, initiated the following brief review of the Aphodiinae found on the Galapagos Islands. The study was made possible by the generous loan of all of the many specimens in the California Academy of Sciences collection, by my good fortune in seeing many other specimens in American collections, and in Paris and London museums. Paratypes of all previously described species except Ataenius atramentarius (Erichson) have been examined. Of the latter I have seen a good series in the British Museum in London. As might be surmised, since the Galapagos Islands are nearest the South American mainland, at least part of the coleopterous fauna has been derived from this continent. Linell, 1898, and Van Dyke, 1953, as well as other authors listed in the very comprehensive bibliography covering the insects of the Galapagos Islands by Linsley and Usinger, 1966, have pointed to South America as the original source of much of the fauna. Van Dyke believed, “The coleopterous fauna is definitely related to that of the barren grounds of western South America especially Ecuador and Peru, and no doubt derived from them at some period long ago.” Hurd, 1958, in discussing carpenter bees expressed his belief that, “when the faunal relationships of the Galapagos Islands are considered 1 Financial assistance was provided by National Science Foundation Grant No. GB-1434. 4—Proc. Biot. Soc. Wasu., Vou. 83, 1970 (51) 52 Proceedings of the Biological Society of Washington in the light of those of continental areas, the faunal affinities are much closer with those of Central America. There is some evidence which suggests that the Galapagos Islands were at one time connected by a series of islands to Central America.” Linsley and Chemsak, 1966, in two papers on the Cerambycidae of the Galapagos Islands, and of Cocos Island, also found affinities with mainland South America, Central America and the West Indies. Two Cocos Island species are found in the West Indies. One of them was described from Cuba. Based on my limited knowledge of the Aphodiinae of South and Central America, and the West Indies, I believe two of the known species probably were derived from species origi- nating in South America, one from Central America, one pos- sibly from the West Indies and one from an endemic on an- other Galapagos Island. Such an assumption must be pure conjecture of course since it is impossible to be sure from which species and exact area the progenitors came. The only species of Diastictus is closely allied to a similar species from Colombia. Ataenius atramentarius (Erichson) was described from Peruvian specimens. Ataenius arrowi Hinton and cavagnaroi n. sp. are close to nugator Harold, 1880, which occurrs from southern United States, through Central America (Costa Rica) and into Argentina and Chile. Ataenius aequa- torialis Petrovitz is similar to the West Indian species A. vincentiae Arrow. Ataenius cavagnaroi, thus far found only on Isla Fernandina, may have been derived from the more widely dispersed, very closely related endemic species arrowi which is known from Isla Santiago, Isla Santa Cruz, Isla Pinta, and Isla Isabella. So far as we know at present all known Aphodiinae found on the Galapagos Islands are endemic except Ataenius atramentarius. Key To GENERA AND SPECIES 1. Length less than 3 mm.; clypeus verrucose; first segment of hind tarsus equal to the length of the following segments combined __________. Diastictus darwini n. sp. Galapagos Beetles 53 Length usually more than 3 mm.; clypeus simply punc- tate, never verrucose; first segment of hind tarsus much longer than following two combined _.. Ataenius spp. 2 2. Length 5.2 to 5.4 mm.; elytral intervals weakly convex over anterior disc, over apical third intervals carinate, laterally eroded and alutaceous _..____-__ rose yl alld Se dll eae A. atramentarius (Erichson) Length less than 5.0 mm.; elytral intervals weakly to moderately convex or carinate over entire length _... 3 3. Length 4.4 to 4.6 mm.; clypeus rounded each side of median emargination; elytral intervals weakly evenly convex; sutural interval impunctate; male anterior spur hooked; tip of tibia and first tarsal segment densely Glethedy waithy long! hair ios s ees ee 4 Length 2.9 to 3.5 mm.; clypeus finely dentate each side; elytral intervals carinately convex entire length, sutural interval strongly punctate ________ aequatorialis Petrovitz 4, Dull surface lustre; elytral striae fine, not noticeably crenate, punctures fine; intervals noticeably alutaceous, Vety weakly convex ..0 2 cavagnaroi n. sp. Shining surface lustre; elytral striae moderately crenate- punctate; intervals shining, moderately convex ______. uments ul nee AO nd es 2 arrowi Hinton Diastictus darwini Cartwright new species Holotype: Length 2.3 mm., width 1.0 mm. Shining, dark castaneous, convex, oblong-oval, widest at apical third. Antennae testaceous. Head strongly convex; clypeal margin evenly rounded each side of moderately deep median emargination, edge narrowly reflexed, surface moderately verrucose, genae sharply rounded, nearly right-angled, a sharply defined anteocular groove extending upward from gena around the eye; occiput with mixed fine and minute punctures. Pronotum convex, one-fourth wider than long, anterior angles obtuse, posterior angles broadly rounded, sides nearly straight but due to convexity not visible from directly above except over anterior third; postapical groove deep in anterior angles causing the angles to appear explanate, the groove continued upward to disc in a series of closely placed moderate punctures leaving a smooth, minutely punctate anterior edge, a similar smooth area laterally outside fovea into the posterior angles, disc with moderately coarse scattered punctures generally separated by one or more times their diameters, longitudinal, vaguely 54 Proceedings of the Biological Society of Washington impressed median line indicated by row of similar punctures; lateral and basal marginal line well defined. Elytra one-third longer than wide, sides slightly divergent to apical third; striae strong, deep, crenately punctate; intervals weakly convex, smooth but with a row of very minute punctures and indistinct very minute alutaceous sculpture near striae, the latter more noticeable on the sutural interval. Metastermum shining, disc smooth but with scattered minute punc- tures, median line weakly impressed, very finely alutaceous at sides, triangular area in front of posterior coxae merely flattened and in- distinctly alutaceous. Abdominal segments shining very minutely alu- taceous especially at sides, crenate along anterior margins, crenations of terminal segment much longer and deeper, at middle about one-third total length of segment. Pygidial transverse eroded area medially inter- rupted at base, eight pygidial setae. Middle and posterior femora smooth, shining, posterior marginal line nearly half the length of femur inward from knee. Posterior tarsus more than half the length of the tibia. First segment of posterior tarsus shorter than long spur, apically as wide as length of second segment, as long as two following segments combined. Holotype, California Academy of Sciences, and six paratypes collected on Isla Wolf, 1 February 1964, by D. Q. Cavagnaro; two paratypes from Isla Darwin, 29 January 1964, by D. Q. Cavagnaro. Paratypes in the California Academy of Sciences and Smithsonian Institution. Paratypes vary in length from 2.2 to 2.7 mm., in color from dark castaenous to piceous. The coarse pronotal punctures are slightly more numerous and the median longitudinal line slightly deeper in some specimens. Pygidial setae number 6 or 8. Diastictus darwini is nearest D. gregalis (Cartwright), new combina- tion, from Colombia, South America. It differs in being noticeably broader, with less convex elytral intervals, a shallower less noticeable metasternal midline and in being uniform in color, not darker laterally than on disc of elytra. Ataenius atramentarius (Erichson), 1847-110 Euparia atramentarius Erichson, 1847-110. Ataenius atramentarius: Harold, 1869-1066. Ataenius dampieri Petrovitz, 1961-356. (New synonymy ) Ataenius scutellaris: Van Dyke, 1953-123. (not Harold 1867-82) Erichson placed atramentarius in Euparia, noting the species had been collected in Peru. I have examined nine specimens of atramentarius in the British Museum and compared paratypes of dampieri Petrovitz with one of them. I can find no differences. In the original description of dampieri, Petrovitz states his species is distinguished by the short hairs of the elytra and the shortened posterior marginal line of the hind femur. Since these characters are exactly duplicated in the Peruvian Galapagos Beetles 55 specimen of atramentarius and I can find no differences otherwise, it is unfortunately necessary to place dampieri in synonymy. Van Dyke misidentified this species as At. scutellaris Harold. Among the specimens he listed was a single specimen from Isla Isabella (Albemarle Island) not now in the Academy Collection. I have seen A. atramentarius (Erichson) from Isla Floreana (Charles Island), Isla Pinta (Abingdon Island), and Isla San Cristobal (Chatham Island). Ataenius aequatorialis Petrovitz, 1961-355 Ataenius aequinoctialis Petrovitz, 1961-355. (Lapsus calami) This species was described from Santa Cruz in 1961. It is the smallest of the Galapagos Ataenius and of the known species it is the only one with dentate clypeus. Except for the more evident short fine elytral pubescence and subcarinately convex elytral intervals, aequa- torialis resembles exiguus Brown or a small abditus (Haldeman), as well as vincentiae Arrow as pointed out by Petrovitz. At. vincentiae differs in having the entire cariniform convex ridge of the elytral intervals broken by a median row of punctures and by having closer, shallower pronotal punctures. All specimens examined were collected on Isla Santa Cruz (Indefati- gable Island). Ataeénius arrowi Hinton, 1936—414 Fig. 1 Dr. Hinton described arrowi from James Island in 1936. The series before me were collected on the Horneman Ranch, Isla Santa Cruz, February and March 1964 and on Isla Darwin in January 1964. The males of arrowi are much more shining than the females, the pronotal punctures are less dense and finer, the anterior tibial spur is bent inward at the tip, and the end of the posterior tibia and tarsus are quite hairy. The posterior tibial fringe of arrowi is usually composed of a group of five short close setae. The short accessory spine is close to the spurs without an intervening seta. I have seen specimens of arrowi Hinton from Isla Santiago (James Island), Isla Santa Cruz (Indefatigable Island), Isla Pinta (Abingdon Island), and Isla Isabella (Albemarle Island). The specimens from Isla Pinta frequently show slight differences such as having more uniform head punctures and the 10th elytral intervals slightly flatter than 9th but I can find no other differences. Ataenius cavagnaroi Cartwright new species Fig. 2 Holotype male—Length 3.9 mm., width 1.8 mm. Piceous, finely alutaceous, weakly shining, elongate, moderately convex. Head moder- ately convex; genae nearly right angled; clypeus rounded each side of moderate median emargination, edge finely reflexed, surface very finely 56 Proceedings of the Biological Society of Washington iS. Fics. 1, 2.—Dorso-caudal view of the lateral lobes of the male genitalia: 1—Ataenius arrowi Hinton, 2—Ataenius cavagnaroi n. sp. punctate, the punctures generally separated by almost twice their diam- eters, band of frontal punctures coarse, about two to three times the diameter of the clypeal puncture and separated by less than half their diameter except in basal or occipital area where they are less dense. Parts of the clypeus have been broken away on each side of the median emargination but remainders of the sides are practically straight to angles of the genae. Pronotum 1.6 mm. wide, 1.1 mm. long, very moderately convex; lateral and basal marginal line fine, edge barely broken by extremely short, fine setae, widely spaced laterally, more closely posteriorly around posterior angles, anterior angles obtusely rounded, posterior angles distinct but rather broadly rounded; surface densely punctate with mixed coarse and fine punctures throughout, the finer punctures usually half or more the diameter of the larger, usually separated by the diameter of the smaller or less, laterally, especially in the anterior angles the punctures are shallower and alutaceous within, surface between punc- tures very finely alutaceous. Elytra 2.6 mm. long, 1.8 mm. wide, oblong oval, humeri finely dentate; striae fine, very weakly crenate with very fine punctures; intervals, all alike, weakly convex, the first three almost flat, surface alutaceous with scattered minute punctures separated by four or more times their diameters. Galapagos Beetles 57 Metasternum concave lengthwise, shining and smooth over middle disc, a moderately strong and deep midline ending anteriorly in wider deeper pore, outer edges of disc very finely and closely punctate, with slightly larger punctures outward to sides, more strongly alutaceous at extreme sides, triangular depressed area in front of hind coxae deep and finely alutaceous. Abdominal segments alutaceous, with scattered mod- erately fine rather shallow punctures throughout, a trifle larger toward sides, separated by one or more times their diameters, segments finely crenate along anterior margin. Eroded area of pygidium strongly alutaceous and with very indistinct shallow punctures, shining margin noticeably widened apically. Spur of anterior tibia bent inward at tip, tibial margin serrate above apical teeth, femur with perimarginal groove, surface closely, roughly, coarsely, shallowly punctate. Middle and hind femora with moderately fine punctures rather irregularly but closely spaced except along an- terior margin and near knee; posterior marginal line half the length of the femur inward from knee. Posterior tibia with four closely spaced fimbriae, short accessory spine close to spurs without intervening seta, first tarsal segment a trifle longer than long spur. Remaining segments of hind tarsi missing. Unique holotype, California Academy of Sciences, collected on Isla Fernandina, W. side, 1,100 ft., 5 February 1964, D. Q. Cavagnaro collector. The species is named after its collector. This species is closely related to Ataenius arrowi Hinton but the pronotal punctures are much coarser and denser than in males of that species and they show less disparity in size than in females of arrowi. The strongly alutaceous elytral intervals are much flatter, the striae and their punctures are fine, and the crenations much less noticeable than in arrowi. The distinctly serrate anterior tibiae are noticeable. The para- meres of the male aedeagus show a distinct lateral tooth, see fig. 2, which is lacking in arrowi. LITERATURE CITED Arrow, GitBeRT JoHN. 1903. On the laporostict lamellicorn Cole- optera of Grenada and St. Vincent (W. Indies). Transactions Entomological Society London, pp. 509-520. Brown, WILLIAMSON JAMES. 1932. New species of Coleoptera III. Canadian Entomologist, vol. 64, pp. 3-12. Cartwricut, Oscar Linc. 1948. The American species of Pleuro- phorus (Coleoptera: Scarabaeidae). Transactions of the American Entomological Society, vol. 74, pp. 131-145. ERICHSON, WILHELM FERDINAND. 1847. Conspectus insectorum cole- opterorum quae in Republica Peruana observata sunt. Archiv Naturgeschicte, vol. 13, pp. 67-185. HALDEMAN, SAMUEL STEHMAN. 1848. Descriptions of North American Coleoptera, chiefly in the cabinet of J. L. LeConte, M.D., 58 Proceedings of the Biological Society of Washington with references to described species. Journal Academy Nat- ural Sciences Philadelphia, vol. I, pp. 95-110. von Haroup, Epcar. 1867. V. Diagnosen neurer Coprophagen, vom Herausgeber. Coleopterologische Hefte. I., pp. 76-119. . 1869. In Catalogus Coleopterorum hucusque descriptorum synonymicus et systematicus autoribus Dr. Max Gemminger et B. de Harold. Vol. 4, Familia 33, Scarabaeidae, pp. 979-1346. Monachii. 1880. Verzeichniss der von E. Steinheil in Neu-Granada gesammelten coprophagen Lamellicornien. Entomologische Zeitung., vol. 41, pp. 13-46. Hinton, Howarp Everest. 1936. New species of Ataenius allied to A. cribrithorax Bates (Coleoptera: Scarabaeidae). Annals and Magazine of Natural History, Ser. 10, vol. 17, pp. 413-428. Hurp, Paut D., Jr. 1958. The carpenter bees of the eastern pacific oceanic islands. Journal of the Kansas Entomological Society, vol. 31, pp. 249-255. LinsLEy, EARLE GORTON, AND JOHN A. CHEMsAK. 1966. Cerambycidae of the Galapagos Islands. Proceedings of the California Academy of Sciences, vol. 33, No. 8, pp. 197-236. 1966. Cerambycidae of Cocos Island, vol. 33, No. 9, pp. 237-248. LinsLEY, EARLE GorRTON, AND Rospert L. Usincrer. 1966. Insects of the Galapagos Islands. Proceedings of the California Academy of Sciences, series 4, vol. 33, No. 7, pp. 113-196. (See this paper for complete bibliography of papers on insects of Galapagos Islands. ) PEetrovitz, RupotF. 1961. Neue und verkannte Aphodiinae aus allen Erdteilen. Entomologischen Arbeiten aus dem Museum G. Frey, Tutzing, vol. 12, No. 2, pp. 344-356. Van Dyke, Epwin Coorer. 1953. The Coleoptera of the Galapagos Islands. Occasional Papers of the California Academy of Sciences, No. 22, pp. 1-181. ery 7. 0672 Vol. 83, No. 5, pp. 59-76 27 May 1970 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON NORTH AMERICAN ORDOVICIAN OPHIUROIDEA_