VIVISIUN OF FISHES Ue. Se NATIONAL MUSEUM 17 MAR +7 1968 OIVISION OF FISHES Ue Se NATIONAL MUSEUs USSS 1S <2: SMITHSONIAN INSTITUTION ae UNITED STATES NATIONAL MUSEUM PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM VOLUME 125 NUMBERS 3651-3668 FINAL VOLUME OF SERIES SMITHSONIAN INSTITUTION PRESS WASHINGTON : 1968 Publications of the United States National Museum The scientific publications of the United States National Museum include two series, Proceedings of the United States National Museum and United States National Museum Bulletins. In these series are published original articles and monographs dealing with the collections and work of the Museum and setting forth newly acquired facts in the fields of anthropology, biology, geology, history, and technology. Copies of each publication are distributed to libraries and scientific organizations and to specialists and others interested in the various subjects. The Proceedings, begun in 1878, are intended for the publication, in separate form, of shorter papers. These are gathered in volumes, octavo in size, with the publication date of each paper recorded in the table of contents of the volume. In the Bulletin series, the first of which was issued in 1875, appear longer, separate publications consisting of monographs (occasionally in several parts) and volumes in which are collected works on related subjects. Bulletins are either octavo or quarto in size, depending on the needs of the presentation. Since 1902, papers relating to the botanical collections of the Museum have been published in the Bulletin series under the heading Contributions from the United States National Herbarium. FRANK A. TAYLOR Director, United States National Museum IT CONTENTS Nicuotas A. Kormitev. Notes on Aradidae in the U.S. National Museum, V (Hemiptera: Heteroptera). 16 pages, 8 figures. Published April 5, 1968 . New species: Acaricoris haitiensis, Hretmocoris eh tek) pn coris drakei, Aneurus haitiensis, Mezira pusilla, M. equatoriana, M. sanmartini, M. paraguayensis, M. crenulata, M. placida. New subspecies: Mezira placida haitiensis. Mitton A. Mituer. Isopoda and Tanaidacea from buoys in coastal waters of the continental United States, Hawaii, and the Bahamas (Crustacea). 53 pages, 4 figures. Pub- lished April 24, 1968 cL nbhitcdnet hk BeAcortaah Zredbe Roger F. Cressey. Caligoid copepods parasitic on Jsurus oryrinchus with an example of habitat shift. 26 pages, 58 figures, 4 plates. Published March 28,1968... . J. F. Gates CLarKeE. Neotropical Microlepidoptera, XVI: A new genus and two new species of Oecophoridae (Lepidop- tera). 8 pages, 3 figures, 2 plates. Published March 28, 1968 Re ak sew « Whgshne, A, soem al guess New genus: Maesara. New species: Maesara gallegoi, Gonionota insulana. HERBERT FRIEDMANN. Parallel evolution in the small species of Indicator (Aves). 10 pages, 2 re Published March 28, 1968 . ats Auan LEwis. Gohepuch toa pene on yAalieg 2s Eniwetok Atoll. 78 pages, 30 figures. Published June 7, 1968 : , New species: iegedttacteb lebewhntiil Sheudiath acca baie: Caligus pseudokalumai, C. alatht. Nicuouas A. Kormitev. North and Central American species of Aneurus Curtis (Hemiptera: Aradidae). 12 pages, 19 figures. Published May 17, 1968 itt New species: Aneurus pusillus, A. aibonilensis, A. pisoniae, A. froeschneri, A. championi. A. StanLtey RAnp and StepHen S. Humpurey. Interspecific competition in the tropical rain forest: Ecological distribu- tion among lizards at Belém, Pardé. 17 pages, 2 figures. Published July 30,1968 . . shine miiqowyh {eres Number 3651 3652 3653 3654 3655 3656 3657 3658 Im IV CONTENTS Kart Krompein. Records and descriptions of additional Scoliidae from New Guinea, Bismarck Archipelago, and Solomon Islands (Hymenoptera: eae 19 pages. Published May 17, 1968 . New species: Campsomeris (Linhaee ers Cs (GD papuana. New subspecies: Campsomeris (Laevicampsomeris) solomonis sedlaceki, C. (Radumeris) extranea shanahani, C. (Phalerimeris) loriae kolombangarensis. New status: Campsomeris (Laevicampsomeris) solomonis solomonis. J. F. Gates Criarxe. Neotropical Microlepidoptera, XVII: Notes and new species of Phaloniidae. 59 pages, 30 figures, 4 plates. Published July 30, 1968 New species: Hvysterosia iodes, H. alphitopa, H. melasma, Phalonia tornosema, P. vesta, P. charma, Cochylis caesiata, C. serena, C. constantia, C. mendora, C. laetitia, Amallectis nephelodes, A. peiiai, A. frangula, A. anazia, A. domna, Phalonidia heterophaea, Irazona icogramma, I. turbula, I. ademonia, I. melanoleuca, I. platina, La- siothyris dysmorphia, L. diamphidia, L. hemitephras, Carolella molyb- dina, C. asthenia, Phtheochroa arrhostia. Artuur G. Humes and Ju-SHry Ho. Cyclopoid copepods of the genus Lichomolgus associated with octocorals of the family Nephtheidae in Madagascar. 41 pages, 113 figures. Published June 4, 1968 2) Lalefess ener New species: Lichomolgus varirostratus, L. exilipes, L. gentilis, L. fissisetiger, L. cuneipes, L. aculeatus. J. R. Napier. Prospects in primate biology. 30 pages, 2 fig- ures, 1 plate. Published April 8, 1968 Arnotp Ross. Bredin-Archbold-Smithsonian Panlgeteel Sie vey of Dominica, 8: The intertidal balanomorph Cirripedia. 23 pages, 3 figures. Published June 25, 1968 ‘ Marvin C. Mryrer. Moore on the eemadines with erie on his oa oy 32 pages. Published November 19, 1968 : 5s 1B dwn CSI ign AMI ge alc hale laeces gs git Ouiver S. FLINT, 1 R. " Bredin-Archbold-Smithsonian Biologi- cal hee of emangen, 9: The Trichoptera (caddisflies) of the Lesser Antilles. 86 pages, 231 figures. Published December 13, 1968 . si New genus: Bredinia. New species: Protoptila dominicensis, Chimarra dominicana, Cc: antilliana, C. caribea, Xiphocentron fuscum, X. albolineatum, Cer- notina lutea, C. cadeti, Polyplectropus bredini, Smicridea caraba, S. simmonsi, S. grenadensis, Leptonema archboldi, Zumatrichia antil- liensis, Z. anomaloptera, Alisotrichia orophila, A. lobata, A. domint- censis, A. wirthi, A. septempunctata, Neotrichia corniculans, Bredinia dominicensis, Hydroptila antilliarum, H. grenadensis, Ochrotrichia 3659 3660 3661 3662 3663 3664 3665 CONTENTS brayt, O. ponta, O. campana, O. similis, O. exclamationis, Brachy- setodes insularis, Atanatolica dominicana, Phylloicus monticolus, Helicopsyche apicauda. Nasmma M. Trrmiz1and Raymonp B. Mannina. Stomatopod crustacea from West Pakistan. 48 pages, 17 figures. Pub- lished December 13, 1968 Pe eine Kart Bansse and Karuerine D. Honson. Benthic poly- chaetes from Puget Sound, Washington, with remarks on four other species. 53 pages, 8 figures. Published Decem- ber 20, 1968 . New species: Hunoe uniseriata, Hulalia ( Plerocirrus) parvoseta, Lao- nice pugettensis, Paraspio cirrifera, Chaetozone acuta, C. berkeleyorum, Tharyx secundus, T. serratisetis. New combinations: Caulleriella annulosa, Neoamphitrite edwardst. Perry C. Hour. The genus Pterodrilus (Annelida: Branchiob- dellida). 44 pages, 12 figures. Published November 19, 1968 New species: Pterodrilus missouriensis, P. choritonamus, P. cedrus, P. hobbsi, P. simondst. 3666 3667 3668 wna) Ans fe errs a Ps 03) jee tae iyi) uae = a n a ukavanet 1 us’ =~ cit] AS hoa ; t ; | s . | ! : | f t ' i ’ | wl yf : oF I q . ; a # a thane J if 120) f 4 oe 0 ' a ¥ | : ‘ + ae? Gi 4 | 1 | | if : a foreedin BS 9° G f Peruntted Stats. ‘1s 1onal Vi nse @en Th Ght hee © ANSBTIUTUTWOR © - WASHING TON Tit pte ce ee ee ee —— wu 135 0 i = ee a ae ee bles i Awa VINVAE 10 hie U. 7 Netional ‘Waay Biiiy Yo: : Reriiptera: Heterortera) ' peel Ue SViehelas Kéorailew® p wae pork valinage at tnd Vidi. 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Volume 125 1968 Number 3651 Notes on ARADIDAE in the U.S. National Museum, V (Hemiptera: Heteroptera) ' By Nicholas A. Kormilev ? For the privilege of studying the unidentified Aradidae in the collections of the U.S. National Museum, Smithsonian Institution, Washington, D.C., and Ohio State University, Columbus, Ohio, | wish to express my sincere appreciation to Dr. Richard C. Froeschner, Curator, Division of Hemiptera, and Dr. Charles A. Triplehorn, Curator of Insects, respectively. All types of new species are deposited at the U.S. National Museum, except two from the author’s collection, which he retains. In the descriptions, 25 units equal 1 mm. In ratios, the first figure represents the length; and the second, the width of measured part. All species treated in this paper belong to the Neotropical Region. Subfamily CarvVENTINAE Usinger Described below are two new apterous species, each in a different genus, from Haiti, and a third, belonging to a third genus, from the island of Dominica, B.W.I. 1 Previous parts of this same series are: I, Kormilev, 1958, Proc. U.S. Nat. Mus., vol. 109, no. 3413, pp. 209-222; II, Kormilev, 1960, Journ. New York Ent. Soe., vol. 68, pp. 36-47; III, Kormilev, 1964, Proc. U.S. Nat. Mus., vol. 115, no. 3483, pp. 245-258; IV, Kormilev, 1966, Proc. U.S. Nat. Mus., vol. 119, no. 3548, pp. 1-25. 2 365 Lincoln Place, Brooklyn, New York 11238. 2 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 125 Drake and Kormilev (1958, p. 243) suppressed the genus Eretmocoris Harris and Drake as a synonym of Acaricoris Harris and Drake. Usinger and Matsuda (1959, p. 132) separated them and indicated that the former is related more closely to Aglaocoris Drake and Maldonado. They further noted that Acaricoris is related closely to a new genus that they erected, Kolpodaptera. Actually, all four genera are related to each other and, pending a needed revision, I prefer to leave them separate. Genus Acaricoris Harris and Drake Acaricoris Harris and Drake, 1944, p. 128. Acaricoris haitiensis, new species Figure 1 FrmMALE.—Elongate ovate, rugose, and glabrous; legs with very fine, inclined hairs. Body covered with a very thin layer of whitish incrustation. Apterous. Head shorter than width across eyes (19:22). Anterior process moderately strong, parallel sided, notched anteriorly; clypeus reaching middle of antennal segment I, much higher than genae, with a small preapical tubercle. Antenniferous tubercles, strong, dentiform, diver- gent. Eyes globose, protruding, but not pedunculate. Postocular borders strongly converging backward, with a thin, longitudinal carina. Vertex raised in the shape of a trident, with median tooth much shorter than laterals. Laterad of trident with two (1+1) elongate ovate callosities. Antennae slender, one-and-a-half times as long as head width (33.5:22); relative length of antennal segments I to IV 10:7:8.5:8; segments I and II clavate, III slightly tapering toward base, IV fusiform. Labium not reaching base of head; labial groove wide, shallow, open posteriorly. Pronotum short and wide (12.5:42), separated from mesonotum by a distinct sulcus. Collar thin, straight anteriorly. Anterolateral angles rounded; lateral borders strongly converging forward, slightly sinuate, with two (1+1) small tubercles posteriorly. Posterior border angularly, deeply incised medially, twice (1+1) sinuate laterally. Disc rugose, with two (1+1) subrectangular tubercles anteriorly, separated from each other by a thin median sulcus that becomes much deeper and wider behind them. Along anterolateral borders, from collar to posterior border, extend two (1-+1) thin carinae. Mesonotum completely fused with metanotum and terga I—-II; the median portion occupied by a spear-shaped elevation with a very thin median sulcus, separated from lateral portions by depressions passing anteriorly into deep sulci. Lateral portions of mesometanotum roundly raised in the middle, discs rugose. NO. 8661 ARADIDAE V—KORMILEV 3 Abdomen almost as long as width across segment IV (64:63), flat; central dorsal plate, consisting of terga III to VI, slightly raised in the middle (area of the first dorsal scent gland scar). Connexivum wide, moderately reflexed. Exterior borders of connexiva straight from IT to V, slightly convex on posterior half of VI and VII. Tergum VII with a short, transverse ridge in the middle of posterior border. Spiracles IIT and III ventral, not visible from above; IV and V sub- lateral and slightly visible; VI to VIII lateral. Paratergites short, subtriangular, reaching middle of tricuspidate segment IX. Cotor.—Piceous, partially black; labium and tarsi yellow brown. MEASUREMENTS.—Total length 4.50 mm, width of pronotum 1.68 mm, width of abdomen 2.52 mm. Hototyer.—lemale, Morne Guimby, 22 km southeast of Fond Verrettes, 6500 feet, Haiti, B. and B. Valentine, 20.VII.1956 (USNM type 69611). Remarks.—Acaricoris haitiensis is related to A. ignotus Harris and Drake but differs by lateral borders of pronotum slightly sinuate and provided with a small tubercle posteriorly, by exterior border of con- nexivum VI forming a rounded angle and connexivum VII rectangular with rounded tip. Genus Eretmocoris Harris and Drake Eretmocoris Harris and Drake, 1944, p. 130. Eretmocoris dominicus, new species Figure 2 Maue.—Subrectangular, parallel sided; females ovate; both sexes flat, heavily punctured and with short, erect bristles on head, borders of the body, antennae, and legs; covered with grayish incrustation. Head as long as width across eyes (co, 34:34; 9, 35:35). Anterior process strong, parallel sided, slightly incised in front, reaching middle of antennal segment I. Clypeus much higher and shorter than genae. Antenniferous tubercles robust, dentiform, strongly divergent. Hyes distinctly stalked. Postocular borders strongly converging back- ward, with four (2+2) small, round tubercles. Vertex with a raised ridge, which is bifurcate anteriorly and separated laterally and in front by deep sulci; laterad of ridge are two (1+1) ovate, raised, rugose callosities. Neck with two (1+1) small tubercles laterally. Antennae short; relative length of antennal segments I to IV: 0, 17.5:9:10:9; 9, 17:10:11:10; I fusiform, robust; II and III slightly tapering toward the base; IV pyriform. Labium reaching hind border of labial groove; latter deep, wide, closed posteriorly. Pronotum short, wide (o, 15:49; 9, 15:55). Collar strong, raised, separated from disc by a deep sulcus. Anterolateral angles produced 4 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 into conical lobes, rounded apically, with a row of small, round tubercles. Anterior border with two (1+1) short ridges placed behind collar, separated from each other by a thin, median sulcus. Behind calli median sulcus becomes wider and deeper. Hind border straight laterally, deeply incised in the middle. Disc rugose, partially deeply punctured. Mesonotum fused with metanotum and terga I and II. Median portion of this plate triangularly raised, with apex inserted into incisure of hind border of pronotum. Mesometanotal plate laterally depressed between mesonotum and metanotum, and raised on metano- tum. Disc densely, deeply punctured, particularly in the middle. Lateral borders of mesonotum produced into two (1+1) lobes similar to those of pronotum; lateral borders of metanotum convex, rounded. Abdomen as long as width across segment V (07, 74:75; 9, 90:91). Terga I and II completely fused with metanotum, but separated from central dorsal plate by a deep sulcus. Central dorsal plate consists of terga III to VI completely fused together, raised in area of dorsal scent gland scars, with carinae dividing disc into a few irregular portions; these portions deeply punctured along carinae; tergum IIIT is punctured on the whole surface. Connexivum wide; segments IT and III fused; exterior border of connexivum II roundly produced anteri- orly; PE angles III to V angularly produced with blunt tip; PE VI and PE VII produced as conical lobes with rounded tip. Tergum VII raised posteromedially. Paratergites short, reaching middle of hypopy- gium; latter caudal in position, deeply inserted; disc of hypopygium with a short median ridge. In female, tergum VII conically raised posteromedially; paratergites conically produced as far as segment IX; latter truncated posteriorly. Spiracles IJ ventral; III sublateral; IV to VIII lateral and visible from above. Cotor.—Dark brown, shiny, concealed by gray incrustation. MerasuREMENTs.—Total length: &@, 5.64; 9, 6.84 mm; width of pronotum: 6, 1.96; 2, 2.20 mm; abdomen width: 0, 3.0; 9, 3.64 mm. Hototypr.— Male, Dominica, W.1I., VI.1963, J. Maldonado (USNM type 69612), from Drake collection. ALLoTYPE.—Female, collected with holotype ‘in copula’; in the same collection. Remarks.—Lretmocoris dominicus is related to EH. gigas Usinger and Matsuda from which it differs by much smaller tubercles on the neck; by absence of thin projections on the foreborder of pronotum; by conical lobes on lateral borders of pronotum and mesonotum; and by progressively produced PE angles of connexiva. NO. 3651 ARADIDAE V—-KORMILEV 5 Genus Aglaocoris Drake and Maldonado Aglaocoris Drake and Maldonado, 1955, p. 291. FIGURE 3 Aglaocoris drakei, new species Frma.Le.—Elongate ovate, shiny; covered with very short, fine bristles, sparse on body and more numerous on head, antennae, and legs. Body completely covered with a thick, whitish incrustation. Apterous. Head almost rectangular, shorter than width across eyes (27:30). Anterior process moderately strong, incised anteriorly, genae slightly longer than clypeus, reaching middle of antennal segment I. Antennif- erous tubercles strong, acute, diverging. Eyes stalked. Postocular borders forming a right angle, with a round tubercle on its tip. Vertex raised as a bifurcate ridge; laterad of which are placed two (1-+1) small, ovate callosities. Antennae slender; relative length of antennal segments I to IV: 13:9:—:— (two apical segments missing); I fusi- form, II tapering toward base. Labium short, reaching hind border of wide, deep labial groove, latter closed posteriorly. Pronotum short, very wide (14:45). Collar thick, separated from disc by a thin sulcus. Anterior borders laterad of collar sinuate; anterolateral angles slightly produced forward, bordered with a dense row of very fine granules. Hind border deeply incised medially. Disc with a transverse carina behind collar, and with a median sulcus behind carina; laterad of median sulcus two (1+1) round callosities; rest of disc irregularly rugose. Mesonotum wider than pronotum (55:45), medially fused with metanotum and in combination with terga IJ-II, forming a subtri- angular, elevated plate, with tip entering into incisure of hind border of pronotum. Mesonotum laterally separated from metanotum by transverse depressions, which mesad turn backward along tri- angular plate; lateral disc of mesonotum with round elevation; mesonotum laterally produced far beyond lateral border of pronotum, forming with the latter a rectangular incisure. Lateral borders of mesonotum straight, parallel, very finely granulate. Abdomen subrectangular, slightly shorter than maximum width across segment IV (65:67.5). Terga I and II flat, completely fused with metanotum but clearly separated from central dorsal plate, which consists of terga III to VI completely fused; disc slightly raised on median line; two (1+1) zigzag carinae separated outside rows of round, calloused spots from dise. Connexivum wide, reflexed. Exterior borders of abdomen straight and parallel from segment II to V, then converging. Connexiva II and III fused; exterior 6 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 125 border of connexivum VI with an elongate, calloused tubercle bearing spiracle; VII with similar, but larger tubercle. Tergum VII raised posteromedially. Paratergites small, conical, blunt, produced as far as segment IX, latter short, truncate posteriorly. Spiracles II to IV ventral, V to VIII lateral and visible from above. Cotor.—Uniformly yellow brown; clypeus, tips of antenniferous tubercles, antennae, and legs ochraceous. MEASUREMENTS.—Total length 5.4 mm, width of pronotum 1.8 mm, width of abdomen 2.7 mm. Houotyre.—lemale, Port-au-Prince, Haiti, 11.1956, C. J. Drake (USNM type 69618), from Drake collection. ReMArKs.—This new species is dedicated to the memory of my late friend Dr. Carl J. Drake, an eminent American hemipterologist. Aglaocoris drakei is related to A. rectangulatus Usinger and Matsuda from San Domingo but differs by genae with parallel sides; longer antennal segment II (two-thirds as long as I, 9:13); by spiracle VI also placed on a tubercle as is spiracle VII; and by smaller size. Subfamily ANEURINAE Douglas and Scott Genus Aneurus Curtis Aneurus Curtis, 1825, pl. 86. Aneurus haitiensis, new species Fiaures 4-6 Marze.—Elongate ovate; pronotum finely granulate anteriorly, with a wide, transversely striate band in the middle, and a narrow, striate band along hind border. Scutellum subtriangular, apex broadly rounded. Head as long as width across eyes (7, 17:17; 9, 17:17.5). Anterior process subparallel, rounded anteriorly, reaching to tip of antennal segment I. Genae slightly shorter than clypeus. Antenniferous tubercles short, truncate anteriorly. Eyes semiglobose, protruding. Postocular border forms a right angle, with three minuscule granules laterally. Vertex transversely striate, almost rugose. Two (1-+1) large, oblique, ovate callosities mesad of eyes. Antennae moderately slender; relative length of antennal segments I to IV: a4, 5.5:5.5:5.5:12.5; 9, 6:6:6:12.5; I obovate, II clavate, III tapering toward base, IV cylindrical. Labium short, not reaching hind border of wide, shallow, transversely rugose labial groove. Pronotum less than half as long as maximum width (07, 17:39; 9, 16.5:39). Collar very thin, sinuate in front. Anterolateral angles subangularly rounded with truncate anterior border and slightly convex lateral border. Lateral notch sharp, forming an obtuse angle. NO. 3651 ARADIDAE V—KORMILEV i Lateral borders of hind lobe subparallel, slightly convex, converging anteriorly. Hind border sinuate in the middle, convex laterally. Fore- dise with four (2+2) flat callosities; hind disc with two (1+1) ovate, transverse callosities surrounded by striated areas. Scutellum flat, concentrically rugose, without callosities. Hemelytra reaching middle (<) or hind border (9) of tergum VII. Corium short, reaching basal one-third of scutellum. Membrane very finely punctured. Abdomen ovate, longer than maximum width across segment IV (@, 71:56; 9, 72.5:54). Connexivum moderately wide, exterior borders of segments straight; PE angles slightly protruding. Para- tergites rounded posteriorly, slightly shorter than hypopygium; the latter small, slightly shorter than wide (5.5:6), not produced beyond exterior borders of connexiva VII. In female, paratergites very short, rounded; segment IX slightly convex posteriorly. Spiracles IT, VI, and VII lateral, visible from above; III to V ventral; VIII terminal. Cotor.—Red brown; disc of scutellum lighter; antennae and membrane darker; labium yellow brown. MEASUREMENTS.—Total length: o, 4.92; 9, 4.96 mm; width of pronotum: o, 1.56; 9, 1.56 mm; width of abdomen: do, 2.24; 9, 2.16 mm. Hototyrr.—Male, Morne Guimby, 22 km southeast of Fond Verrettes, Haiti, 6500’, B. and B. Valentine, 20.VII.1956 (USNM type 69614). ALLotTyPE.—Female, collected with holotype; Ohio State Univer- sity collection, Columbus, Ohio. ParaTyPES.—T'wo males, one collected with holotype, another a day earlier in the same place; in Ohio State University collection and collection of the author. REMARKS.—Aneurus haitiensis is related to the North American A. pygmaeus Kormilev from which it may be separated by antennal segment II as long as I or III, antennal segment IV more than twice as long as III, and by larger size. Subfamily MEztrinaE Oshanin Genus Mezira Amyot and Serville Mezira Amyot and Serville, 1843, p. 305. In my key (1962, p. 260) for American Mezira species, 69 of the 71 species were listed. Mezira horvathi (Bergroth) and Mezira novella Blatchley were excluded because of lack of specimens and insufficient data. Later, three species were described. Now, six new species and one new subspecies are added. 8 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 125 Mezira pusilla, new species FIGURE 7 Fremaue.—Elongate ovate, with finely setigerous granulations bearing extremely short, curled hairs. Head almost as long as width across eyes (18.5:19). Anterior process constricted at base, dilated apically; genae forming two (1+1) large, rounded lobes in front of clypeus, reaching to three- fourths of antennal segment J. Antenniferous tubercles acute, strongly divergent. Eyes semiglobose, protruding. Postocular tubercles thin, acute, slightly produced beyond outer border of eyes. Infraocular carinae moderately high, crenulate. Vertex with a U-shaped row of granules. Antennae slender, one-and-a-half times as long as width of head (29.5:19); relative length of antennal segments I to IV: 8:5:9.5:7. Labium reaching to hind border of labial groove, latter closed posteriorly. Pronotum less than half as long as maximum width (16:36); fore- lobe narrower than hind lobe (29:36). Collar thin, slightly sinuate anteriorly. Anterolateral angles expanded and rounded, produced forward almost as far as collar. Lateral notch deep, forming a slightly obtuse angle. Lateral borders of hind lobe slightly convex, converging anteriorly. Hind border almost straight. Foredise with four (2+2) granulate ridges. Hind dise roughly granulate. Scutellum shorter than basal width (14:20); all borders carinate, apex cut out; lateral borders sinuate at apical half; median ridge cross shaped; disc roughly granulate. Hemelytra reaching to hind border of tergum VI; basolateral borders of corium straight, carinate; apical angle rounded; apical border convex, slightly sinuate interiorly. Abdomen longer than maximum width across segment V (60:41). Lateral borders slightly convex. Connexivum wide, slightly raised exteriorly; PE angles slightly protruding; PE VII rounded. Para- tergites large, rounded, reaching to middle of a moderately long segment IX, latter incised posteriorly. Spiracles II to VI ventral; VII sublateral, barely visible from above; VIII lateral and visible from above. Cotor.—Testaceous, partly ferrugineous; labium and tarsi yellow; membrane fuscous, white at base. MrasurEMEnNts.—Total length 4.4 mm; width of pronotum 1.24 mm; width of abdomen 1.64 mm. Hoxrorypr.—Female, Cacao, Trece Aguas, Alta v. Paz, Guatemala, Barber and Schwarz, (USNM type 69615). Remarks.—WMezira pusilla is one of the smallest Mezira species known from the Neotropical Region, only Mezira nana (Champion) NO. 3651 ARADIDAE V—KORMILEV 9 being smaller. In my key (1962, p. 260) for American Mezira species, it runs to M. angustata (Champion) but is much smaller, genae are ex- panded and rounded, apical border of corium more convex, para- tergites and segment IX are relatively longer. Mezira equatoriana, new species Fremaue.—In general aspect similar to Mezia pacifica Usinger but differing from it by: Head longer than width across eyes; anterior process shorter, reaching three-fifths of antennal segment I. Eyes relatively large, semiglobose. Postucular tubercles smaller, blunt, adherent to eyes, not reaching outer border of latter; antennal seg- ment III moderately longer than II (15:12), much longer in M. pacifica (17.5:10). Pronotum more sinuate laterally; interior ridges of forelobe less raised. Scutellum relatively shorter, wider at base. Apical border of corium almost straight (sinuate interiorly and con- vex exteriorly in M. pacifica). Paratergites with posterior border convex exteriorly and straight interiorly, reaching as far as a short segment IX; latter truncate posteriorly. All spiracles ventral. Cotor.—Brown; membrane black; antennae and legs lighter, red brown; labium and tarsi yellow brown. MEASUREMENTS.—Head longer than width across eyes (30:27.5) ; relative length of antennal segments I to IV: 15:12:15:15. Pronotum less than half as long as wide (26:66); forelobe narrower than hind lobe (52:66). Scutellum shorter than basal width (25:41). Abdomen longer than maximum width across segment IV (92:87). Total length 7.08 mm; width of pronotum 2.64 mm; width of ab- domen 3.48 mm. Ho.otyre.—Female, Esmeralda, south of Mateo, Ecuador, J. Foerster, 5.X.1956, deposited in the collection of the author. Mezira sanmartini, new species Frma.e.—Elongate ovate, roughly but sparcely granulate; granules with distinct, yellow, curled hairs. Head distinctly shorter than width across eyes (21:26). Anterior process dilated anteriorly, rounded and incised in front, reaching three-fourths of antennal segment I. Antenniferous tubercles wide, acute, crenulate outside and slightly divaricating. Eyes large, semi- globose, protruding. Postocular tubercles clawlike, acute, produced slightly beyond outer border of eyes. Infraocular carinae high, cren- ulate. Vertex with a V-shaped, rough, setigerous granulation. An- tennae moderately strong; relative length of antennal segments I to IV: 10:6:—:— (two apical segments missing). Labium reaching to hind border of labial groove, latter closed posteriorly. 10 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 Pronotum less than half as long as its maximum width (22.5:50) ; forelobe narrower than hind lobe (43:50), separated from the latter by a deep depression. Collar moderately wide, angularly incised anteriorly. Anterolateral angles strongly expanded, rounded and crenulate, produced anteriorly as far as collar. Lateral notch deep. Lateral borders of hind lobe rounded and crenulate. Foredisc with four (2 + 2) high, granulate ridges. Hind lobe roughly granulate. Scutellum shorter than basal width (19:26). All borders carinate, tip incised; lateral borders straight, almost without sinus on apical half; median ridge cross shaped; disc roughly granulate. Hemelytra reaching three-fourths of tergum VI. Basolateral border of corilum carinate and crenulate; apical border straight; apical angle rounded; all veins with a rough, setigerous granulation. Abdomen longer than its maximum width across segment IV (77:60). Midlateral glabrous areas separated from central dorsal plate by high, crenulate carinae. Connexivum wide and raised exteriorly; disc partially covered with whitish incrustation. PE angles barely pro- truding; PE VII rounded. Paratergites subtriangular reaching three- fourths of segment IX, with a large, lateral spiracle; segment IX slightly incised posteriorly. Spiracles II to VII ventral, on a tubercle far from the border, VIII lateral. Cotor.—Ferrugineous, partially dark ferrugineous; labium and tarsi orange yellow. MrasurEeMents.—Total length 5.72 mm, width of pronotum 2.00 mm, width of abdomen 2.40 mm. Hototyer.—Female, Magdalena, Estado Bolivar, Rio Cauza, Venezuela, P. San Martin, 5.X.1957, deposited in the collection of the author. REMARKS.—It is a pleasure to dedicate this species to Mr. Pablo San Martin, an Uruguayan entomologist who collected this specimen and donated it to the author. Mezira sanmartini runs in my key (1962, p. 260) for American Mezira species to M. sangabrielensis Kormilev from Amazonas, Brazil, from which it differs by anterior process of head rounded anteriorly, not crenulate; by anterolateral angles of pronotum pro- duced forward as far as collar; by lateral notch of pronotum much deeper and angular; by paratergites (?) produced only to three-fourths of segment IX; and by setigerous granulation with distinct, yellow, curled hairs. Mezira paraguayensis, new species Mauy.—Elongate ovate, slightly widening backward; covered with a dense and fine granulation bearing extremely short setae. Head shorter than width across eyes (c7, 23:25; 9, 25:27). An- NO. 3651 ARADIDAE V——KORMILEV 11 terior process stout, with large, expanded, and rounded anteriorly genae, contiguous in front of clypeus and produced slightly beyond tip of antennal segment I. Antenniferous tubercles narrow, acute, diverging, reaching to the middle of antennal segment I. Eyes semi- globose, protruding. Postocular tubercles small, acute, produced as far as eyes, or slightly beyond. Infraocular carinae high, thin, finely granulate. Vertex with M-shaped group of granulations. Antennae slender; relative length of antennal segments I to IV: o&, 9:6:12:7.5; 9, 9:6.5:12:8.5. Labium short, reaching to hind border of labial eroove, latter closed posteriorly. Pronotum half as long as maximum width (07, 25:51; 9, 25:54); forelobe narrower than hind lobe (o%, 43:51; 9, 44:54). Collar thin, slightly sinuate in front. Anterolateral angles expanded, rounded, slightly reflexed, crenulate, produced forward as far as collar. Lateral notch distinct, forming an obtuse angle. Lateral borders of hind lobe subparallel, converging anteriorly, crenulate; hind border feebly sinuate. Foredise with four (2 + 2) high, granulate ridges; hind disc more roughly granulate. Scutellum shorter than basal width (co, 20:25; 9, 25:27). Lateral borders carinate, sinuate on apical half; apex widely rounded. Median ridge cross shaped, roughly granulate. Hemelytra reaching to hind border of tergum VI in both sexes. Corium with convex basolateral borders; apical border convex, rounded; apical angle also rounded. Veins of corium densely granulate. Abdomen ovate in both sexes; longer than maximum width across segment IV (o, 74:62.5; 9, 80:65). Connexivum wide, slightly reflexed exteriorly, exterior borders of segments straight, crenulate; PE angles not, or barely protruding; PE VII angularly rounded in both sexes. Paratergites (co) small, clavate, reaching slightly over two-thirds of hypopygium; latter cordate, shorter than wide (15:20), with a median ovate ridge not reaching tip of hypopygium. Parater- gites (2) large, rounded, reaching middle of segment IX, latter incised posteriorly. All spiracles ventral, not visible from above. Cotor.—Piceous with rusty brown granulations; genae, antennae, tips of antenniferous tubercles, lateral borders of forelobe of pro- notum, PE angles, and hind borders of connexiva, coxae, trochanters, and tarsi, yellow brown to rusty brown. Exterior borders of connexiva on forehalf, black. Round callous spots on connexiva and venter reddish brown. MeAsuREMENTS.—Total length: o, 5.80; 9, 6.12 mm; width of pronotum: o’, 2.04; 9, 2.16 mm; width of abdomen: 0, 2.50; 2, 2.60 mm. Houotypr.—Male, Horqueta, 45 m east of Paraguay River, 12 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 125 Paraguay, Alberto Schulz, 4.V1I.1933 (USNM type 69616), from Lutz collection in U.S. National Museum. AutLotyPE.—Female, collected with holotype; in the same col- lection. ParaTyPES.—T'wo males and one female collected with holotype; in the same and author’s collection. Remarks.—Mezira paraguayensis runs in my key (1962, p. 260) for American Mezira species to M. punctiventris (Stal) but is much smaller, piceous; granulation is rusty brown, connexivum tricolored, brown, black, and yellowish. Mezira crenulata, new species FIGuRE 8 Maue.—Elongate ovate, with sharp, setigerous granulation; setae short, rusty, curled; lateral borders of pronotum distinctly crenulate. Head as long as width across eyes (32:32). Anterior process stout, constricted at base, rounded anteriorly, apex incised in the middle, reaching three-fifths of antennal segment I. Antenniferous tubercles blunt, with convex, subparallel outer borders. Eyes semiglobose, protruding; placed behind the middle of lateral border. Postocular tubercles minute, not reaching outer borders of eyes. Infraocular carinae low, with minute, setigerous granulation. Vertex with \- shaped granulation. Antennae moderately stout; relative length of antennal segments I to IV: 20:12.5:15.5:13. Labium short, not reaching to hind border of labial groove, latter closed posteriorly. Pronotum half as long as maximum width (37:76); forelobe much narrower than hind lobe (60:76). Collar narrow; anterolateral angles expanded, rounded, and crenulate; lateral notch forming a slightly obtuse angle. Foredisce with four (2+2) high, granulate ridges. Lateral borders of hind lobe convex, crenulate; hind disc granulate; hind border shallowly trisinuate. Granulations bearing distinct, curled, rusty hairs. Scutellum shorter than basal width (31:37). All three borders carinate; lateral borders sinuate before apex; median carina cross shaped; disc granulate. Hemelytra reaching slightly over fore border of tergum VII; basolateral border of corium reflexed, slightly sinuate and crenulate; apical angle rounded; apical border convex outside, sinuate interiorly. Abdomen ovate, longer than maximum width across segment IV (105:98). Connexivum wide; exterior borders of connexiva barely convex; PE angles slightly protruding; PE VII forming a right angle with rounded tip, reaching to the middle of hypopygium. Paratergites thin, clavate, reaching three-fifths of hypopygium; latter cordate, NO. 3651 ARADIDAE V—KORMILEV lie? shorter than wide (20:30), depressed above, with an elevated ovate median ridge not reaching to hind border. Spiracles small, far from border on II to VII; sublateral, but not visible from above on VIII. Cotor.—Ferrugineous, partly piceous; connexivum concolorous. MeasurEMENtTS.—Total length 8.56 mm, width of pronotum 3.04 mm, width of abdomen 3.92 mm. Houoryrpr.—Male, collected in Hoboken, New Jersey, on Orchids imported from Venezuela, 1.VIII.1940 (USNM type 69617). Remarks.—Mezira crenulata, in my key (1960, p. 260) for American Mezira species runs to M. mexicana Kormilev but looks more lke M. boliviana Kormilev from which it differs by spiracles VIII sub- lateral and not visible from above (lateral and visible in M. boliviana); head as long as wide; lateral borders of pronotum distinctly crenu- late; hypopygium different: its upper surface produced as far as its lower portion (in M. boliviana the upper portion is distinctly shorter than its lower portion). Mezira placida, new species Mauzr.—Elongate ovate, shghtly widening backward, then narrow- ing again; covered with setigerous granulations; setae short and curled. Median ridge of hypopygium reaching to three-fourths its length; paratergites of the female subtriangular with rounded tip, divergent and reaching to one-fourth segment IX. Spiracles II to VII ventral, placed far from border; VIII sublateral, but not visible from above. Cotor.—Dark ferrugineous, connexivum and venter lighter. MaAsuREMENTS.—Head 22.5:25; relative length of antennal seg- ments I to IV: 10:9:11:10; pronotum 21:51; scutellum 19.5:28; abdomen 80:58 across segment V; hemelytra reaching to foreborder of tergum VII; hypopygium 20:24. Total length 6.0 mm, width of pronotum 2.04 mm, width of abdomen 2.32 mm. Hotoryrr.—Male, Hugural Rd., Puerto Rico, J. R. Johnston, 10.1V.1913 (USNM type 69618). ALLoTyPE.—Female, collected with holotype; in the same collec- tion. ParatyPes.—T'wo males and four nymphs of different instars, collected with holotype; in the same collection and collection of the author. Remarks.—This species is based on a series of specimens from Puerto Rico and one series from Haiti. The Puerto Rico series has one specimen with a label ‘‘Mezira n.sp., det. H. G. Barber,” and the Haiti series has one specimen with a label ‘“Brachyrhynchus sp.? 14 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 125 very near B. angustatus Champ. O. H.” (presumably by Otto Heide- mann). In my key (1962, p. 260) for American Mezira species both series run to M. granulata Say. Comparison of both series with a paratype of M. angustata (Champion) reveals that they are not related. Com- parison with various specimens of M. granulata Say shows that they are closely related. The general shape of these two series and M. granulata is the same; biometric measurements show the same pattern though the specimens from Puerto Rico and Haiti are slightly larger. The main differences are: postocular tubercles are shorter, not reaching to outer border of eye in specimens from Puerto Rico and Haiti, reaching it in M. granulata; lateral notch of pronotum in specimens from Puerto Rico and Haiti is angular, but in M. granu- lata sinuate, rounded, though J have seen specimens of M. granulata also with angular lateral notch; paratergites in the female are slightly shorter in specimens from Puerto Rico and Haiti reaching to one- fourth segment IX, while in M. granulata they reach to the middle of segment IX; the last but not the least difference is in pilosity: M. granulata has setigerous granulation with straight hairs, whereas specimens from both Puerto Rico and Haiti have distinct curled hairs. Mezira granulata shows a certain variability in the length of setae; specimens from Texas and Mississippi have setae distinctly longer than specimens from Georgia, or Virginia, but the setae always are straight, never curled. The differences between specimens from Puerto Rico and Haiti are even smaller. Besides color, which in Mezira usually has no specific value, the main difference is the median ridge on the hypo- pygium: in the specimens from Puerto Rico it reaches three-fourths the hypopygial length, whereas in the specimens from Haiti it reaches to, or almost to, the hind border of the hypopygium. The best solution is to consider the specimens from Puerto Rico as a new species, herein named Mezira placida, new species, and the specimens from Haiti as its geographical subspecies, M. p. haitiensis, new subspecies. The following key will separate the two new taxa from each other and from M. granulata: 1. Postocular spines produced as far as outer borders of eyes; lateral borders of pronotum mostly roundly sinuate; setigerous granulations with straight setae; paratergites in the female reaching to the middle of segment IX. Mezira granulata Say, United States Postocular tubercles not reaching to outer borders of eyes; lateral borders of pronotum with an angular notch; setigerous granulations with curled hairs; paratergites in the female reaching to one-fourth segment IX. Mezira placida, new species, West Indies 2. Median ridge extending three-fourths length of hypopygium; color dark ferruginous ............ . M. placida placida, Puerto Rico NO. 3651 ARADIDAE V—KORMILEV 15 Median ridge of hypopygium reaching almost to its tip; color testaceous. M. placida haitiensis, new subspecies, Haiti Mezira placida haitiensis, new subspecies Ma.re.—Similar to Mezira placida placida, but median ridge of hypopygium reaching almost to the tip of the latter. Cotor.—Testaceous; membrane fuscous, whitish at base; labium and tarsi yellow. MEASUREMENTS.—Head 21.5:22.5; relative length of antennal segments I to IV: 10:8:11:9; pronotum 22:46; scutellum 19:26; ab- domen 73:54 across segment IV; hemelytra reaching one-fifth tergum VII; hypopygium 17:22. Total length 5.54 mm, width of pronotum 1.84 mm, width of abdomen 2.16 mm. Ho.totrypr.—Male, Port au Prince, Haiti, E. D. Ball, 1940 (USNM type 69619). ALLoTYPE.—Female, Port au Prince, Haiti, Herbert Osborn, deposited in the collections of Ohio State University, Columbus, Ohio. PaRATYPES.—Six males, collected with holotype, in the collections of U.S. National Museum and in author’s collection; seven males, collected with allotype, in the collections of Ohio State University and in collection of the author. Literature Cited Amyot, C. J. B., and AupINET-SERVILLE, J. G. 1843. Histoire naturelle des insectes: Hemipteres, lxxvi+681 pp., 12 pls. Curtis, J. 1825. British entomology, being illustrations and descriptions of the genera of insects found in Great Britain and Ireland; containing colored figures from nature of the most rare and beautiful species, and in many instances of the plants on which they are found, vol. 7, pl. 86. Drake, C. J., and MaLtponapo CapRiLss, J. 1955. New apterous Aradidae from Puerto Rico. Journ. Washington Acad. Sei., vol. 45, pp. 289-294, 9 figs. Drake, C. J.. and Kormitev, N. A. 1958. Concerning the apterous Aradidae of the Americas (Hemiptera). Ann. Ent. Soc. America, vol. 51, pp. 241—247, 2 figs. Harris, H. M., and Draxg, C. J. 1944. New apterous Aradidae from the Western Hemisphere. Proc. Ent. Soc. Washington, vol. 46, pp. 128-132. Kormi.Lev, N. A. 1962. Notes on Aradidae in the Naturhistoriska Riksmuseum, Stockholm: Hempitera-Heteroptera. Ark. Zool., ser. 2, vol. 15, no. 14, pp. 255-273, 15 figs. Usinearr, R. L., and Marsupa, R. 1959. Classification of the Aradidae, vii + 410 pp., 102 figs. London: British Museum. 16 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 Ficures 1-8.—1, Acaricoris haitiensis, new species, 9 , head and pronotum; 2, Eretmocoris dominicus, new species, ¢’, head, pronotum, and mesonotum; 3, Aglaocoris drakei, new species, 9, head pronotum, and mesonotum; 4, Aneurus haitiensis, new species, o, head, pronotum, scutellum, and corium of hemelytra; 5, Aneurus haitiensis of, tip of abdomen, dorsal aspect; 6, Aneurus haitiensis 9, tip of abdomen, dorsal aspect; 7, Mezira pusilla, new species, 9 , head and pronotum; 8, Mezira crenulata, new species, do’, hypopygium, dorsal aspect. U.S. GOVERNMENT PRINTING OFFICE: 1968 Proceedings” of the United States National Museum SMITHSONIAN INSTITUTION + WASHINGTON, D.C. Volume 125 1968 Number 3652 Isopoda and Tanaidacea from Buoys in Coastal Waters of the Continental United States, Hawaii, and the Bahamas (Crustacea) ! By Milton A. Miller? This article is based on collections made during World War II by the author and other biologists employed by Woods Hole Oceano- graphic Institution on a survey of marine fouling. The research was conducted under supervision of Dr. Alfred C. Redfield and the late Dr. Louis M. Hutchins. It was done under contract with the Bureau of Ships, U.S. Navy, with invaluable logistic support from the U.S. Coast Guard. The author wishes to express deep appreciation to Dr. Redfield for his guidance and encouragement on this and other wartime investigations on the marine fouling problem. The author is indebted to Mrs. Lynn Rudy for illustrations and other assistance in the preparation of this paper. Thanks are due also to Dr. Thomas E. Bowman of the U.S. National Museum for a critical review of the manuscript. 1 Contribution No. 1816 from Woods Hole Oceanographic Institution, Mass., and a contribution from the Department of Zoology, University of California, Davis. 2 Department of Zoology, University of California, Davis 95616. ys PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 Fouling communities include not only various sessile organisms (barnacles, mussels, bryozoans, tube worms, tunicates, sponges, algae, etc.) that attach themselves to submerged structures, but also many free-living forms associated with them. The free-living animals may feed on the sedentary forms, find cover among them, and exhibit various degrees of symbiosis with them. Among the nonsedentary forms commonly found in the fouling association are isopods, tanaidaceans, and other Crustacea. The isopods and tanaidaceans reported in this paper were ex- tracted from samples of fouling scraped from buoys and their moorings in the coastal waters of the United States. As mentioned above, these collections were made during World War II in connection with a survey of marine fouling conducted by Woods Hole Oceanographic Institution. Biological objectives included determination of the amount and kinds of fouling, rates of accretion, regional and seasonal variation, and ecological factors affecting establishment of fouling organisms. Much valuable systematic, ecological, and distributional data may be derived from investigations of this sort. Such informa- tion is needed badly for most invertebrate groups, especially in regions in which the biota has not been investigated adequately. Besides intrinsic interest, the biological data also might have practical application in the development of more effective methods for control of fouling. Additionally, it was thought that data on buoy fouling could provide clues as to the origin and possibly the course of drifting mines and military flotsam. Cosmopolitan or widely dis- tributed forms obviously would be of little or no value as tracers. Attached forms clearly would serve the purpose better than free- living types. In any event, the distributional limits of species that might be found on buoys would have to be established definitely before any valid conclusions could be drawn. Unfortunately, our present biogeographical knowledge of most invertebrates is too in- complete to be of much help in determining origin or drift. The results of the marine fouling survey are incorporated in a treatise on marine fouling and its prevention that has been pub- lished by Woods Hole Oceanographic Institution (1952). The present paper extends the preliminary account of the Isopoda and Tan- aidacea given in that publication. Mernops.—The buoy fouling survey was initiated in 1943. Biologists were assigned to various naval districts with authorization to accom- pany buoy tenders of the U.S. Coast Guard in order to investigate fouling on buoys when they were relieved, serviced, or otherwise tended. Generally, at this time, the buoy with mooring chain and anchor were hoisted onto the deck, which gave the biologists an opportunity for observation and sampling. NO. 3652 ISOPODA AND TANAIDACEA—MILLER 3 i 1:BUCY STATIONS or Tanaid sites (numbered = fem stations ( ) cattle California Coast | oO 50 100 150 - cape Hatteras RILOMETE Ss | ve (Ls _ é Lokout a Atlantic Coast} Ir 26p° KAUAI aia is | i) meetin TERS sya | samt o a - : : * » rains, ae Bei te Stikses | Pics at Nn os @ ~~ MRein fassigtitin= v. Ceo, 1G, ne ey ee te es eee Wl ve. Cabs ae! Oe Han a ad ¥ - gut oe oe hae hi Cnvcandvin Ma = hits “1? re ie Lie ‘ort 4 an Vy Mats ek Li eerie ad: Ma heat ccc a oo | | ry Piper, > Pn app ae (ape Onan, eee s, Cima : ean ae mia | eT ee RRMA File oynt Al tharenlhls eva Yara *) meeting = yy SOD ih EAL, TET DAS {howl Geer ee yee pees, Satine | as a ee hn a tee, Se ne ey fe dei wei) eapsy 8's pgilet FAD, ta 1% aie hi @ wit ter Cease hh entoekee lneales ee “l Al Grk geale we Uinmsiern yrdivedy (UG tal os Ties vee het, fe Ghajirems aril 40” re Lard (a shui ny Ben ef ats aro ae heres Da a, hie ng ae Ball aaah Ba . ne bie Ate | a7 bee re > ee Bary a . if Fe 2) 7 < a ee iw _ af - -_ : 4 ‘a wis) pee ie Ve, hd oe, ee ee Proceedings of the United States National Museum SMITHSONIAN INSTITUTION +« WASHINGTON, D.C. Volume 125 1968 Number 3653 Caligoid Copepods Parasitic on Isurus oxyrinchus With an Example of Habitat Shift By Roger F. Cressey Associate Curator, Division of Crustacea Several recent collections of parasitic copepods from 17 specimens of the mako shark, Jsurus oryrinchus, have made possible redescrip- tions of certain species of these parasites commonly found on this shark. These, plus 15 collections from the Indian and Atlantic Oceans previously reported (Cressey, 1967a and 1967b), form the basis of this paper. Ten species of caligoid copepods are reported herein. Both sexes of Nemesis lamna Risso and the males of Dinemoura producta (Miiller) and D. latifolia (Steenstrup and Liitken) are redescribed. The last copepodid instar and three chalimus stages of Phyllothereus cornutus (Milne-Edwards) males are described for the first time. The remaining species have been redescribed recently by various workers (Shiino, 1955; Lewis, 1966; and Cressey, 1967a), and no further redescription of these species is considered necessary. All drawings have been made with the aid of a camera lucida. All specimens are preserved in 70 percent alcohol and deposited in the U.S. National Museum. 1 2 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 125 Station data for the R. V. Anton Bruun stations cited in this paper | are as follows: station nwmber date latitude longitude 552 2/17/66 33°02’8 77°02’ W 553 2/19/66 33°24/5 79°00’ W 554 2/20/66 32°58'S 81°31’W 558 2/24/66 30°28’S 89°31’/W 564 3/ 7/66 20°37'S 83°20’ W 567 3/10/66 12°50’S 83°31/W 568 3/11/66 10°33’S 83°38’ W 569 3/12/66 08°49’S 83°33’ W 571 3/14/66 08°47'S 81°20’ W 572 3/15/66 10°28’S 79°36’ W I acknowledge the support of the National Science Foundation- Southeastern Pacific Biological Oceanographic Program for supporting | a portion of the field work and the Smithsonian Research Foundation — for supporting a portion of the laboratory work. I thank Mr. John Casey, Sandy Hook Marine Laboratory, for a collection of shark copepods, a portion of which have been used in this study. Pandarus satyrus Dana Pandarus satyrus.—Lewis, 1966, p. 74. Pandarus cranchti.—Hewitt, 1967, p. 249 [not P. cranchii Leach]. For earlier synonymy, see Cressey, 1967a, p. 6. MareriaLu.—Four collections from off the west coast of South America, R. V. Anton Bruun, Cruise 14, Stations 553, 564, 569, and 571; two collections from the Indian Ocean previously reported (Cressey, 1966b); one collection from off New Jersey. This copepod has been well figured and described in recent literature, and no further description is included here except as related to the remarks below. Remarks.—In the seven collections reported above, this copepod was found in the mouth or on the gill arches of the host. Pandarus satyrus is a common parasite on the body surface of Prionace glauca. In all cases the egg-bearing females of P. satyrus from I. oxyrinchus were pigmented less noticeably than individuals of the same species when they occur on the body surface of blue sharks. The record of P. cranchii cited by Hewitt from the “jaws and skin” of J. oryrinchus appears to be P. satyrus according to his figures (figs. 141-147). Pandarus smithii Rathbun Pandarus smithit—Lewis, 1966, p. 91. For earlier synonymy, see Cressey, 1967a, p. 11. MatEriALu.—Three collections from the North Atlantic (two off New Jersey, one at 35°N-70°W): one collection from the Indian Ocean, R. V. Anton Bruun, Cruise 5, Station 282. NO. 3653 COPEPODS—CRESSEY 3 As in P. satyrus, this copepod has been redescribed recently (Cressey 1967a), and no further description will be given here. Remarks.—The copepods of this species from the Atlantic were found in the mouth of the host. The single female representing the Indian Ocean collection from this host was on the body surface. Like P. satyrus, this copepod is a common parasite on the body sur- face of sharks, primarily carcharinids. Also, as in P. satyrus, those specimens from inside the buccal area of the host are not pigmented as heavily as individuals of the same species occurring on the body surface of other sharks. Pandarus katoi, Cressey Pandarus katoi Cressey, 1967a, p. 17. Mareriau.—Four collections from off the west coast of South America, R. V. Anton Bruun, Cruise 14, Stations 567, 568, 569, and 571. Remarks.—Pandarus katoi has been reported previously from the body surface of carcharinid sharks in the eastern Pacific (Cressey, 1967a). The collections recorded above from J. oxyrinchus were removed from the mouths and gill arches of the hosts. The specimens in collections of this species from the body surface of other sharks are not pigmented as heavily as most other species of the genus Pandarus; nevertheless, the P. katoi from the mouths of makos are pigmented still less than those on the body surface of other sharks, consistant with the situation with P. satyrus and P. smithw reported above. The change in habitat for these three species is discussed at the end of this paper. Gangliopus pyriformis Gerstaecker For earlier synonymy, see Cressey, 1967a, p. 28. MarTErRIAL.—One collection off the west coast of South America, R.V. Anton Bruun, Cruise 14, Station 552 (1 9 specimen only). RemMArkKS.—The single specimen from the 32 sharks examined indicates a low incidence of this parasite. This copepod is more common on Prionace glauca and, since these two sharks often are found in the same waters, this may be a reflection of that association. Phyllothereus cornutus (Milne-Edwards) Figures 1-26 Phyllothereus cornutus—Lewis, 1966, p. 96.—Hewitt, 1967, p. 233. For earlier synonymy, see Cressey, 1967a, p. 25. Mareriau.—Four collections off the west coast of South America’ R. V. Anton Bruun, Cruise 14, Stations 553 (two collections), 554: and 572. 4 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 Many immature stages including the last copepodid stage were collected from the sharks off South America. When some of these copepodids were cleared in KOH and lactic acid, the first chalimus stage could be seen within. By carefully splitting open the copepodid, I teased out the developing chalimus. These chalimus stages were identi- cal to first chalimus stages collected free in the gill chamber of the shark. Two successive chalimus stages also were found. These im- mature stages are described below. For a description of the adults of this species, see Cressey (1967a). Last copEPopID.—Body form as in figure 1. Total length 6.77 mm (6.23-7.43) based on an average of 240 specimens, greatest width 4.07 mm (3.78—4.38) based on an average of 20 specimens (all measured specimens taken from a single shark at R. V. Anton Bruun Station 554). Dorsal surface unornamented. Two “eye spots” with more heavily pigmented area surrounding and between them in anterior third of cephalon. Posterior corners of first free thoracic segment produced, extending nearly to end of second free thoracic segment. Genital segment slightly longer than wide (1.3 x 1.1 mm in one specimen). Abdomen 1-segmented, somewhat triangular, wider than long (.3 x .4 mm in one specimen). Caudal ramus (fig. 2) 536u wide, 590 long at outer edge, 295u long at inner edge (based on one specimen) ; armed with six plumose setae, four long terminal and two short subterminal. Adhesion pad present on ventral surface immediately posterior to first antenna. First antenna (fig. 3) 2-segmented: basal segment with several short plumose spines, proximal two more heavily spinose; terminal segment with seven setae, posterior four plumose. Second antenna (fig. 4) with terminal segment in form of a claw, tip of claw twisted. Mouth tube and mandible of usual caligoid form, mandible with 10 teeth. First maxilla (fig. 5) a small lobe with three short setae and posterior projection bearing process with rugose tip. Second maxilla (fig. 6) with two terminal claws and subterminal hirsute process, claws ornamented as in figure 7. Maxilliped (fig. 8) stout, bearing bladelike claw opposed by striated area on basal segment. Legs 1-4 (figs. 9-12) biramose. Rami of legs 1-3 2-segmented. Exopod of leg 4 1-segmented, endopod 2-segmented (see spine and seta formula below). Leg 5 (fig. 13) a knob bearing two setae, one naked and one plumose, on lateral margin of genital segment. Leg 6 absent. Color in life: dark mahogany brown. CHALIMUS I, MALE.—Body form as in figure 14. Total length 6.6 mm, greatest width 3.1 mm (all measurements based on a single specimen taken from the same shark as copepodids). NO. 8658 COPEPODS—CRESSEY 5 Dorsal surface unornamented. Eye spots less obvious than in copepodid. Outer posterior corners of first free thoracic segment produced, extending nearly to end of second free thoracic segment. Genital segment as wide as long (1.4 x 1.4 mm), posterior corners produced to form short rounded lobes. Abdomen 2-segmented; each segment about same length but second segment somewhat wider than first and triangular (widest posteriorly). Caudal rami triangular; with four long terminal and two short subterminal plumose setae. Adhesion pad on ventral surface posterior to base of first antenna. First antenna (fig. 15) 2-segmented, similar to copepodid except that segments longer, bearing more setae. First maxilla (fig. 16) similar to copepodid except rugose process of copepodid is short pointed process in chalimus I. Second maxilla of chalimus I and all subsequent stages with plumose processes at base of two terminal spines (see Cressey, 1967a, fig. 129). Maxilliped (fig. 17) with pointed claw, seta on inner margin of claw. Legs 1-4 biramose. All rami 2-segmented. Leg 1 exopod with four spines equal in length on outer terminal margin of second segment, endopod of first segment with adhesion pad on inner distal corner. Leg 2 exopod with four outer spines, distalmost about twice length of proximalmost spine. Leg 3 exopod with four outer spines, proximalmost very small; other three becoming progressively longer, distalmost twice length of next longest. Leg 4 exopod with four spines, lengths as in leg 3. Leg 5 consisting of spine and two plumose setae at notch at posterior three-fourths of genital segment. Leg 6 a single spine near junction of abdomen and genital segment. Color in life: cream. CHALIMUS II, MALE.—Body form as in figure 18. Total length 8.6 mm, greatest width 4.6 mm (all measurements based on single speci- men). Clear fringe around cephalon and outer corners of first free segment (not seen in previous stage). Principle changes in body form from previous stage are proportionately wider cephalon, elongation of genital segment, and more prominent caudal rami. Adhesion pads present ventrally on cephalon, one posterior to base of first antenna and a small one on basal segment of second antenna. First antenna (fig. 19) with both segments longer and with in- creased ornamentation over previous stage. Second antenna (fig. 20) with well-developed claw; bearing two short setae, one at base of claw and one median. Maxilliped (fig. 21) with well-developed claw. Other head appendages as in previous stage. Legs 1-4 biramose (see pls. 1-4). Spine and seta formula for legs 1-4 of last three immature stages and adult male are as follows (Ro- man numerals refer to spines, arabic to setae) : ~ 6 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 copepodid chalimus I chalimus IT chalimus IIT leg seg. exo. end, ezo. end. exo. end. exo. end. 1 Ht 1:0 0:0 1:0 0:0 1:0 0:0 1:0 0:0 2 Vid 3 IV:3 3 LVi33 3 Waste 3 2 1 Meal 0:1 ial O:1 leit 0:1 Weil On 2 IV:5 7 IV:5 8 IV:6 8 IV:6 8 3 1 1:0 0:1 itaal 0:1 es Ost Hleentt 0:1 2 IIl:4 4 IV:5 5 Ves 5 6 IV:5 6 4 1 VERS O31 EO 0:1 Jia (eal Jigal 0:1 2 3 EVis5 4 LVS Is IV:5 5 Leg 5 (fig. 22) consists of single stout spine and three short plumose setae at notch in margin of genital segment (an increase of one seta over previous stage). Leg 6 a short spinelike process with plumose seta near junction of abdomen and genital segment. The lengths in mm of each of the free body segments for the last five developmental stages of the male are as follows (specimens measured along mid-dorsal, anterior-posterior axis): chalimus chalimus chalimus I II IIT copepodid adult cephalon 2.4 2. 6 SH 3. 6 5. 3 first free thoracic segment 9 .6 G a) .9 second free thoracic segment 7 .6 8 8 12 third free thoracic segment 9 .8 9 209) 1.3 genital segment 1.3 1.4 Ieee 2.3 3. 8 abdomen 3 .3 5 1.0 1.4 caudal ramus 1 .6 8 At) 2.0 total body length 6. 6 6. 6 8167) 084 15.9 CHALIMUS III, MALE.—Body form as in figure 23. Total length 10.4 mm, greatest width 4.9 mm (measurements based on single specimen). This form is ornamented in all respects like chalimus JI. There is a general increase in size and further elongation of genital segment. Cephalic appendages show elongation of terminal segments when compared with previous stage (claw of second antenna, maxilliped and terminal segment of first antenna are proportionally longer). Other than subtle changes in proportions, appendages are the same as in chalimus IJ. For comparison of legs 1-4, see plates 1-4. ApuULT MALE.—Total length 15.9 mm, greatest width 7.2 mm (measurements based on one specimen). The appendages of the adult do not differ significantly from the previous stage except for subtle changes in lengths of terminal segments of cephalic appendages, the adult appendage is somewhat longer in proportion (compare adult first maxilla, fig. 24 with fig. 16). The maxilliped of the adult male (fig. 25) bears two striated pads opposing the claw instead of just one as in previous stages. Changes in ornamentation and proportionate lengths of the segments of the first antenna of all stages of the male are as follows: NO. 3653 COPEPODS—CRESSEY 7 segment 1 segment 2 no. no. percent spines percent spines total and total and length length setae length length setae copepodid 266u 66 14 136 34 5 chalimus I 342u 68 23 165y 32 12 chalimus IT 413u 64 27 236u 36 14 chalimus III A72u 61 27 307 uy 39 14 adult 53lp 58 27 378yu 42 14 ADULT, FEMALE.—Body form as in figure 27. For a complete description of the female of this species, see Cressey (1967a). Remarks.—The description of the copepodid instar of this species is the first to be linked conclusively to a known species of pandarid copepod. Similar forms have been described as the males of the genus Nesippus (Dana, 1852; Steenstrup and Liitken, 1861; Beneden, 1892; Gnanamuthu, 1949; and Hewitt, 1967). This description substantiates my previous remarks regarding these Nesippus males (1967a, p. 59). Since no forms intermediate between chalimus III and the adult male were found, it is assumed that there are only three chalimus stages. A few immature females were collected but not enough to determine their stage of development. Description of these forms has been de- ferred until more material can be collected. The copepodids were found either on the body surface of the host or within the gill chamber, usually just inside the gill slit. The chalimus stages and adults all were found within the gill chamber but not usually on the filaments themselves. Most often these forms were found near the tips of the filaments, firmly attached to the epidermis. The females often cause the tissue to swell, almost covering the cephalon. Males were not observed to produce this host reaction. This parasite is not restricted to Zsurus but also is found commonly on other lamnid sharks and Prionace glauca. Dinemoura latifolia (Steenstrup and Liitken) FiGuRES 26-27 Dinematura latifolia—Lewis, 1966, p. 102.—Hewitt, 1967, p. 195. For earlier synonymy, see Cressey, 1967a, p. 42. Marteriau.—Three collections in the western North Atlantic (two previously reported by Cressey, 1967a) ; one collection from the Carib- bean Sea, 13°38’N, 75°50’W;; five collections from the Indian Ocean, previously reported by Cressey (1967b); eight collections from off the western coast of South America, R. V. Anton Bruun, Stations 553, 554, 564, and 567); one collection from the central Pacific, vicinity of Christmas Island. 8 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 Remarxks.—Both sexes of this copepod have been described, most recently by Lewis (1966). In my pandarid revision (1967a), I noted the lack of information regarding the male of this species. Lewis provides a good description of this sex in his 1966 paper. My material conforms to his description in every way except in regard to the first antenna (antennule in Lewis). According to Lewis, the first segment bears 14 setules and the second segment, five setae. In my material I have found each segment with more setae than described by Lewis (25 on the first segment, 12 on the second). A figure of the male (fig. 26) and its first antenna (fig. 27) is provided to supplement the description by Lewis. This copepod is a common parasite on the body surface of mako sharks. Large clusters of females often occur in the posterior half of the body and usually on the ventral surface. Males generally are fewer in number and are scattered over the body surface. Dinemoura producta (Miiller) FIGURES 28-42 Dinemoura producta.—Hewitt, 1967, p. 204. For earlier synonymy, see Cressey, 1967a. MartertAu.—A single collection previously reported from the North Atlantic (Cressey, 1967a); three collections from off Scripps Pier, La Jolla, Calif.; 12 collections off Chile and Peru, R. V. Anton Bruun, Stations 552, 553, 554, 567, 568, 569, 571, and 572. Frema.e.—A good description of the female of this species was given by Shino (1957). Additions to his description were provided by Cressey (1967). Further description of the female will not be included herein. Ma.r.—The male of this species was described by Wilson (1923) but a redescription of this sex is needed and is provided below. Body form as in figure 28. Total length 8.25 mm, greatest width 3.6 mm (measurements based on a single specimen from Station 567). Lengths in mm of each of the body segments measured along the mid- dorsal line are as follows: cephalon 2. 55 first free thoracic segment zo second ‘‘ se f . 60 third ‘ se i led genital segment 2, 18 abdomen (2 segments) . 90 caudal ramus . 90 No dorsal plates present. Abdomen 2-seemented, second segment about twice as long as first. Genital segment (fig. 29) nearly twice as NO. 3653 COPEPODS—CRESSEY 9 long as wide (2.18 x 1.30 mm). Caudal rami (fig. 30) conspicuous, longer than wide (.90 x .52 mm); armed with four prominent plumose setae and two small subterminal plumose setae, inner margins of rami with row of short hairs. First antenna (fig. 31) 2-segmented; basal segment with 26 short spines and setae, terminal segment with 12 short naked setae. Second antenna (fig. 32) stout; terminal claw with accessory process near base and two setae, one short and one long; appendage armed with number of adhesive areas as indicated in figure. First maxilla (fig. 33) with 3- segmented terminal ramus, basis bearing group of three short setae. Second maxilla (fig. 34) terminal claw with rows of fine hairs; second segment bearing short stout spine covered with fine spinules, patch of hairs on outer distal corner. Maxilliped (fig. 35) well developed; ter- minal claw opposed by two adhesion areas on basal segment. Legs 1-4 biramose. Leg 1 (fig. 36) both rami 2-segmented. Exopod first segment large, bearing spine on outer distal corner; second seg- ment with fringe of digitiform processes along outer edge and bearing three short spines and four setae. Endopod first segment unarmed, distal end produced to form broad lamella overlapping base of terminal segment; terminal segment with three stout setae and patch of hairs on outer edge. Leg 2 (fig. 37) both rami 3-segmented. Exopod first two segments of each with spine and seta on outer and inner distal corners respectively, terminal segment with two lateral spines and six setae. Endopod first segment with seta on inner distal corner, second seg- ment with two setae at inner distal corner, terminal segment with six setae. Leg 3 (fig. 38) both rami 3-segmented. Exopod with same arma- ture as leg 2. Endopod with last two segments modified as in figure 39. Leg 4 (fig. 40) rami 2-segmented. Exopod first segment with spines on outer distal corner, second segment with spine on outer and seta on inner distal corners, terminal segment with two spines and five setae. Endopod with one, two, and three setae on the inner margin of each segment respectively. Spine and seta formula of legs 1-4 is as follows: leg 1 leg 2 leg 3 leg 4 seg exp. end. exp end, exp. end. erp. end, 1 1:0 0:0 Meet 0:1 Megat 0:1 1:0 0:1 2 JIT: 4 3 i{gu 0:2 Jugal 0:2 ipgil 0:2 3 I1:6 5 LEG [:4 ibe 3 Leg 5 (fig. 41) represented by process on ventral surface of genital segment bearing one outer fringed spine and two inner plumose setae (see fig. 29); a fourth plumose seta internal to process. Leg 6 (fig. 42) a process near posterior corner of genital segment bearing one outer naked spine and one short inner plumose seta. 10 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 REMARKS.—This copepod normally is found on the fins of the host, most frequently on the trailing edges of the dorsal and pectoral fins. Echthregaleus denticulatus Smith Echthrogaleus pellucidus Shiino, 1963, p. 357. For earlier synonymy, see Cressey, 1967a, p. 56. MarTerrau.—One collection from off Scripps Pier, La Jolla, Calif.; one collection from the central Pacific in the vicinity of Christmas Island; six collections off Chile and Peru, R. V. Anton Bruun, Stations 553, 554, 568, 569, and 571; and four collections from the Indian Ocean previously reported (Cressey, 1967b). Remarks.—No description of either sex of this copepod will be given herein as that of Shiino (1963) and of Cressey (1967a) are adequate. I have placed E. pelludicus Shiino in synonomy after re- examining the type of E. denticulatus Smith deposited in the USNM. The only differences among all the material I have collected recently, the description of E. pellucidus by Shiino, and the type-specimen is that the type is larger. The type-specimen was collected in the North Atlantic, whereas all other material has come from the Indo-Pacific. Cressey (1967a, p. 27) reported a similar situation for another species of parasitic copepod, Phyllothereus cornutus, where the specimens collected from the colder waters of the Atlantic were larger than those from the relatively warmer Indian Ocean. These copepods generally are found on the body surface of the host, frequently in the anterior part near the gill slits. Although there was a high incidence of infestation on mako sharks off Chile and Peru, the number of individuals on any one shark was never more than 20. This same species was collected in the Indian Ocean from thresher sharks, Alopias vulpinus, and often occured on that host in clusters of more than 100 individuals. Anthosoma crassum (Abildgaard) For earlier synonymy, see Lewis, 1966, p. 66. Mareriau.—F our collections from off Scripps Pier, La Jolla, Calif. ; four collections from the central Pacific in the vicinity of Christmas Island; one collection from off Sandy Hook, N.J.; two collections from the Indian Ocean, previously reported (Cressey, 1967b) ; one collection from the North Atlantic taken during a cruise of the M. V. Delaware; seven collections from off Chile and Peru, R. V. Anton Bruun, Stations 552, 553, 554, 558, 564, 568, and 571. ReEeMARKS.—This species has been well described by Shino (1955) and Lewis (1966) and no further description will be given herein. On the mako sharks caught off the western coast of South America, this species was very common and was always found imbedded be- NO. 3653 COPEPODS—CRESSEY 11 tween the teeth on either or both the upper and lower jaw. It has been recorded also from other species of lamnid sharks. Nemesis lamna Risso Ficures 43-58 Nemesis lamna Risso, 1826, p. 1386.—Roux, 1828, p. 174.—Gueérin, 1837, pl. 35.—Krgyer, 1837, p. 199.—Milne-Edwards, 1840, p. 486.—Wilson, 1922, p. 59; 1932, p. 461.—Fage, 1923, p. 282.—Heegaard, 1962, p. 184.— Cressey, 1967b, p. 6. Nemesis carchariarum Roux, 1828, p. 176.—Kr¢yer, 1837, p. 199.—Milne-Edwards, 1840, p. 486 [misspelled “‘carcherium’’]. Nemesis mediterranea Heller, 1868, p. 220.—Valle, 1880, p. 66.—Richiardi, 1880, p. 151.—Stossich, 1880, p. 262.—Carus, 1885, p. 365.—Brian, 1898, p. 15; 1899, p. 201; 1902, p. 44; 1905, p. 7.—Bassett-Smith, 1899, p. 476. Nemesis lamna var. sinuata.—Valle, 1878, p. 91. Marteriau.—Four collections from the Indian Ocean, previously reported (Cressey 1967b); three collections from off Scripps Pier, La Jolla, Calif.; nine collections from off the Coast of Chile and Peru, R. V. Anton Bruun, Stations 552, 553, 554, 558, 564, and 572. FEMALE.—Body form as in figure 43. Total length 6.61 mm (5.85- 7.95), greatest width 2.0 mm (1.73-2.40). Widths of cephalon and four free thoracic segments are as follows: cephalon 1.25 mm (1.19- 1.50), first free thoracic segment 1.92 mm (1.80—2.33), second free thoracic segment 2.00 mm (1.73-2.40), third free thoracic segment 1.99 mm (1.73-2.40), fourth free thoracic segment 1.74 mm (1.50- 2.25). All measurements are based on an average of 10 ovigerous females randomly selected from the collection at Station 552. Fourth free thoracic segment (bearing leg 5) with many sclerotized papillae on dorsal surface. Genital segment (fig. 44) about as long as wide (.8 x .8 mm) and subdivided ventrally giving the appearance of two segments. Abdomen (see fig. 44) 2-segmented. Each segment with a few papillae on dorsal surface and armed with short hairs and spinules as in figure. Caudal rami short, about twice as long as wide (363u x 218), bearing rows of short spinules on ventral surface. First antenna (fig. 45) composed of 13 segments. Hach segment armed with short setae as in figure. Second antenna (fig. 46) 4- segmented, basal segment with weak triangular process on inner sur- face, antepenultimate segment with short seta on inner distal corner, penultimate segment unarmed, terminal segment in form of short claw bearing two setae on inner border. Mouth tube and mandible of usual caligoid type, mandible with eight to nine teeth. First maxilla (fig. 47) biramose; each rami with two setae, anteriormost ramus with one short and one long seta. Posterior ramus with two equal long setae. Tip of second maxilla (fig. 48) with short claw, claw covered with short papillae. Patch of spines present near base of claw 12 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 and patch of long hairs on outer corner of segment bearing terminal claw. Maxilliped (fig. 49) in form of stout claw. Penultimate segment with triangular process on inner proximal corner. Claw heavily sclerotized and bearing two short setae on inner margin. Legs 1-4 biramose, each ramus 2-segmented. All legs with con- spicuous patches of spinules as indicated in figures. Leg 1 (fig. 50) exopod first segment long and recurved, second segment small bearing three short spines at tip; endopod first segment with outer distal corner produced to form prominent claw, last segment with two weak setae at tip. Second leg (fig. 51) exopod first segment with inner and outer corners produced to form spines, last segment short, bearing six short spines of about equal length; endopod first segment similar to that of exopod, last segment with five spines. Leg 3 (fig. 52) exopod as in leg 2 except last segment bearing seven short spines; endopod first segment as in leg 2, last segment with prominent spinelike process on outer distal corner and one small outer and four small inner spines. Leg 4 (fig. 53) exopod first segment with spine on outer distal corner only, last segment with six small spines; endopod first segment as in legs 2 and 3, last segment with five spines, next to outermost larger than other four. Leg 5 represented by knob bearing three short setae on outer distal corners of genital segment. Ma.r.—Body form as in figure 54. Total length 4.7 mm, greatest width 1.6 mm measured across first free thoracic segment (measure- ments based on single specimen from Station 552). Cephalic appendages as in female. Leg 1 as in female. Leg 2 (fig. 55) exopod first segment with prominent spine on outer distal corner and small seta on inner margin, second segment with six small setae on terminus and inner margin; endopod first segment with prominent spine on outer distal corner, second segment with three short but somewhat recurved setae. Leg 3 (fig. 56) exopod first segment with outer prominent spine and small seta on inner distal corner, second segment with six setae, fourth from outer considerably longer than other five; endopod first segment with outer distal corner produced to form prominent spine, endopod with two terminal spines and three small setae on inner margin, spine on inner distal corner strong and somewhat recurved. Leg 4 (fig. 57) exopod first segment with promi- nent spine on outer distal corner, second segment with six setae, inner three longer than outer three; endopod first segment with outer corner produced to form strong spine and inner margin bearing con- spicuous seta, second segment with two terminal spines and two inner short setae, spine on inner distal corner strong and slightly recurved as in leg 3. Leg 5 (see fig. 58) a short free segment bearing three setae plus single seta near base of segment. Leg 6 (see fig. 58) on outer distal corner of genital segment consisting of prominent inner spine NO. 3653 COPEPODS—CRESSEY is and two outer setae. Spermatophores visible within genital segment. ReMArRKs.—Wilson (1932) redescribed both sexes of this species and stated that no description had been presented since Heller (1865). He apparently was unaware of the redescription of the female and descrip- tion of the male by Fage (1923). Although Fage’s description is of more value than that of Wilson, both lack certain details and, thus, a re- description of both sexes has been presented herein. This species is very common on mako sharks and, as in other members of the genus, the adults are found only on the gill filaments of the host. It has been reported from other lamnid sharks as well. Habital Shift in the Genus Pandarus All species of the genus Pandarus are found commonly on the body surface of their respective shark hosts. In this paper three species are reported from Jsurus oxyrinchus. Of the 16 collections of Pandarus from this host, only one was taken from the body surface (P. smithit in the Indian Ocean). All others were collected from the mouth and/or the gill arches. During the course of my studies with parasitic cope- pods, it has become evident that there is considerable specificity with regard to the site of infestation on the host. One can predict with a high degree of certainty sites of infestation for any species. Needless to say, any marked deviation from the usual mode of existence presents some interesting problems. Such is the case with the genus Pandarus as found usually on Jsurus oxyrinchus. While collecting Pandarus from the mouths of mako sharks, I suspected that they might represent new species due to the unusual habitat and their noticeable reduction in pigmentation. Subsequent examination, however, showed that they all could be assigned to the known species—satyrus, smithii, and katoi—differing from other members of each species only in the reduced pigmentation. An exami- nation of the collection records showed that in all cases where Pandarus was present in the mouth, either Dinemoura latifolia and/or D. producta were present at the sites where one might expect to find Pandarus on other hosts. In the one collection where Pandarus occurred on the body surface, Dinemoura was not present. Apparently, the presence of Dinemoura inhibits Pandarus from becoming established in its usual places. The fact that it does become established in a new loca- tion is interesting. I suspect that in time, if they continue to be success- ful in this habitat, they will result in new species of the genus being produced. By the usual criteria of separating the species of the genus, they cannot at present be considered as different species. If, however, their progeny result in forms that will not attach to any other host on the body surface, then they represent something new. A definitive answer to this would come only from experimental evidence and cannot 280-626—68——2 14 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 125 be ascertained at this time. I suspect that this situation represents new species in the early stages of formation. It is interesting to note that Hewitt (1967) cites two records of P. cranchii (which are actually P. satyrus) from the “buccal cavity” and “jaws and skin’ of Zsurus oxyrinchus and Galeorhinus australis. Dinemoura latifolia was collected from both of these sharks. As previously indicated, adult females of the three species in the mouth were pigmented less noticeably than adult females of the same species that occurred on the surface of other hosts. Two possible explanations come to mind. First, the pigmentation may be influenced by the amount of light available to the copepod, those in the mouth receiving less. Second, the pigmentation may be influenced by the amount of pigmentation of the shark at the point of attachment of the parasite. From my own observations of other parasitic copepods, I favor the latter explanation. Often parasitic copepods assume the color of the host tissue to which they are attached, or those parasites with little or no pigment are associated with lighter areas and the pig- mented forms are on darker areas. Many questions basic to under- standing this departure remain unanswered. Most of these deal with the life history process about which practically nothing is known. No attempt at a definitive explanation is offered here, but rather I have presented the situation in the hope that future investigations will shed more light on this interesting situation. PROC. U.S. NAT. MUS. VOL. 125 CRESSEY—PLATE 1 Phyllothereus cornutus, male: a, copepodid leg 1; 6, chalimus I leg 1; c, chalimus II leg 1; d, chalimus III leg 1. PROG] US. INAT MUSS VOLE, 125 GRESSEY—PEATES2 Phyllothereus cornutus, male: a, copepodid leg 2; b, chalimus I leg 2; c, chalimus II leg 2; d, chalimus III leg 2. PROC. U.S. NAT. MUS. VOL. 125 CRESSEY—PLATE 3 Phyllothereus cornutus, male: a, copepodid leg 3; b, chalimus I leg 3; c, chalimus II leg 33 d, chalimus III leg 3. PROC, U.S. NAT. MUS. VOL.. 125 GRESSEY—PEAGE?4. Phyllothereus cornutus, male: a, copepodid leg 4; 6, chalimus I leg 4; c, chalimus 11 leg 4; d, chalimus III leg 4. Literature Cited BassEert-SmiTH, P. W. 1899. A systematic description of parasitic Copepoda found on fishes, with an enumeration of the known species. Proc. Zool. Soc. London, pt. 2, pp. 438-507. BENEDEN, P. J. VAN 1892. Le male de certain caligides et un nouveau genre de cette famille. Bull. Belgique, year 62, ser. 6, vol. 23, no. 3, pp. 220-235. Brian, A. G. G. 1898. Catalogo di copepodi parassiti dei pesci della Liguria. Atti. Soc. Lig. Sci. Nat. Geogr., vol. 9, pp. 5-31. 1899. Crostacei parassiti dei pesci dell’Isola d’Elbe, 2: Contribuzione. Atti. Soc. Lig. Sci. Nat. Geogr., vol. 10, pp. 197-207. 1902. Note su alcuni crostacei parassiti dei pesci del Mediterraneo. Atti. Soc. Lig. Sci. Nat. Geogr., vol. 18, pp. 30-45. 1905. Sui copepodi raccolti nei golfo di Napoli da Oronzio G. ed Achille Costa. Ann. Mus. Zool. Univ. Napoli, vol. 1, no. 24, pp. 1-11. Carus, J. V. 1885. Prodromus faunae mediterraneae ... , vol. 1, xi +525 pp. Cresszy, R. F. 1967a. Revision of the family Pandaridae (Copepoda: Caligoida). Proc. U.S. Nat. Mus., vol. 121, no. 3570, pp. 1-133. 1967b. Caligoid copepods parasitic on the sharks of the Indian Ocean. Proc. U.S. Nat. Mus., vol. 121, no. 3572, pp. 1-21. Dana, J. D. 1852-53. Crustacea, pts. 1 and 2. Vol. 13 im United States exploring expedition . . . 1888-42 ... , pt. 1 (1852), viii +685 pp.; pt. 2 (1853), pp. 686-1618. Faas, L. 1923. Sur deux copépodes Dinemoura producta (Muller) et Nemesis lamna (Risso) parasites du pélerin, Cetorhinus maximus (Gunner). Bull. Soc. Zool. France, vol. 48, nos. 6 and 7, pp. 280-287. GNANAMUTHIU, C. P. 1949. Two male parasitic copepods from Madras. Ann. Mag. Nat. Hist., ser. 12, vol. 2, pp. 359-367. GuUERIN-MENEVILLE, F. E. 1837. Crustacés. Vol. 26 in Iconographie du regne animal. [1829-1843: plates; 1844: text.] HEEGAARD, P. 1962. Parasitic Copepoda from Australian waters. Rec. Australian Mus., vol. 25, no. 9, pp. 149-233. HELLEmR, C. 1868. Crustaceen. No. 8 in vol. 2 of Zoologischer Theil zn Reise der Oesterreichischen Fregatte Novara. . ., 280 pp., 25 pls. Hewitt, G. C. 1967. Some New Zealand parasitic Copepoda of the family Pandaridae. New Zealand Journ. Mar. Freshw. Res., vol. 1, no. 2, pp. 180-264. 15 16 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 Krgyer, H. 1837-38. Om synltekrebsene, isaer med Hensyn til den Danske Fauna. Naturh. Tidsskr., vol. 1, no. 2, pp. 172-208; no. 3, pp. 252-304; no. 5, pp. 476-506; no. 6, pp. 605-628; (1838), vol. 2, pp. 8-52, 131-157. Lewis, A. G. 1966. Copepod crustaceans parasitic on elasmobranch fishes of the Hawaiian Islands. Proc. U. 8S. Nat. Mus., vol. 118, no. 3524, pp. 57-154. MItNnge-Epwarps, H. 1840. Histoire naturelle des crustacés, comprenant l’anatimie, la physiologie et la classification de ces animaux, 638 pp., 42 pls. RIcHIARDI, 8S. 1880. Contribution alla fauna D’Italia: Catalogo sistematico dei corstacei che vivono sul corpo degli animali aquatici. In Catalogo degli Espositori, pp. 147-152. Risso, A. 1826. Historie naturelle des principales productions de l’ Europe méridionale et particulierement de celles des environs de Nice et des Alpes martitimes, vol. 5, 402 pp. Roux,.P. 1828. Crustacés de la Mediterranee et de son littoral, 176 pp., 45 pls. Paris. Surno, S. M. 1955. Copepods parasitic on Japanese fishes, 8: The Anthosomidae. Rep. Fac. Fish. Pref. Univ. Mie, vol. 2, no. 1, pp. 50-69. 1957. Copepods parasitic on Japanese fishes, 13: Parasitic copepods col- lected off Kesennuma, Miyagi Prefecture. Rep. Fac. Fish. Pref. Univ. Mie, vol. 2, no. 3, pp. 359-375. STeEnstTRvP, J. J. S., and Litxen, C. F. 1861. Bidrag til Kundskab om det aabne Havs Snyltekrebs og Lernaeer semt om nogle andre nye eller hidtil kun ufuldstaen digt kjendte parasitiske Copepoder, vol. 5, pp. 341-342. Srossicu, M. 1880. Prospetto della fauna del mare Adriatico, 3: Classe V, Crustacea. Bull. Soc. Adriatica Sci. Nat. Trieste, vol. 5, no. 5, pp. 178-271. VaLLe, A. D. 1878. Sopra due specie di crostacei parassiti dell’Oxyrrhina spallanzant Raf. Estratto Boll. Sci. Nat., vol. 4, no. 1, pp. 89-92. 1880. Crostacei parassiti dei pesci del mare Adriatico. Boll. Soc. Adriatic Sci. Nat., vol. 6, pp. 55-90. Witson, C. B. 1922. North American parasitic copepods belonging to the family Dichelesthiidae. Proc. U. S. Nat. Mus., vol. 60, no. 2400, pp. 1-100. 1923. Parasitic copepods in the collection of the Riksmuseum at Stockholm. Ark. Zool., vol. 15, no. 3, pp. 1-15. 1932. The copepods of the Woods Hole region, Massachusetts. U.S. Nat. Mus. Bull. 158, 635 pp. 17 COPEPODS—CRESSEY NO. 3653 Ficures 1-8.—Phyllothereus cornutus, male copepodid: 1, dorsal; 2, caudal ramus; 3, first antenna; 4, second antenna; 5, first maxilla; 6, second maxilla; 7, base of terminal claws on second maxilla; 8, maxilliped. 18 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 ( FO — So — Os Tr ‘) N : LN I, Vl & 4 Z > A J a i) Ay Ficures 9-13.—Phyllothereus cornutus, male copepodid: 9, leg 1; 10, leg 2; 11, leg 3; 12, leg 4; 13, leg 5. NO. 3653 COPEPODS—CRESSEY 19 a na i l a =— Ficures 14-19.—Phyllothereus cornutus, male chalimus I: 14, dorsal; 15, first antenna; 16, first maxilla; 17, maxilliped. Male chalimus II: 18, dorsal; 19, first antenna. 20 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 oe ee 25 Ficures 20-25.—Phyllothereus cornutus, male chalimus II: 20, second antenna; 21, maxilli- ped; 22, leg 5. Male chalimus III: 23, dorsal; 24, first maxilla; 25, maxilliped. NO. 3653 COPEPODS—CRESSEY 21 AN h\ Figures 26-31.—Dinemoura latifolia, male: 26, dorsal; 27, first antenna. D. producta, male: 28, dorsal; 29, genital segment and abdomen, ventral; 30, caudal rami; 31, first antenna. 22, PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 Ficures 32-38.—Dinemoura producia, male: 32, second antenna; 33, first maxilla; 34. second maxilla; 35, maxilliped; 36, leg 1; 37, leg 2; 38, leg 3. NO. 3653 COPEPODS—CRESSEY Zo Ficures 39-44.—Dinemoura producta, male: 39, terminal segment on endopod of leg 3; 40, leg 4; 41, leg 5; 42, leg 6. Nemesis lamna, female: 43, dorsal; 44, genital segment and abdomen, ventral. 24 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 Ficures 45-49.—Nemesis lamna, female: 45, first antenna; 46, second antenna; 47, first maxilla; 48, tip of second maxilla; 49, maxilliped. NO. 3653 COPEPODS—CRESSEY 25 vvv Vy vy vuvvyyvvY v v v v ryv vv y wv yy vv Yue ¥ CA cp ey bev Ficures 50-54.—Nemesis lamna, female: 50, leg 1; 51, leg 2; 52, leg 3; 53, leg 4. Male: 54, dorsal. 26 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 125 Ficures 55-58.—Nemesis lamna, male: 55, leg 2; 56, leg 3; 57, leg 4; 58, edge of genital segment showing legs 5 and 6. U.S. GOVERNMENT PRINTING OFFICE: 1968 Proceedings of the United States National Museum SMITHSONIAN INSTITUTION + WASHINGTON, D.C. Volume 125 1968 Number 3654 Neotropical Microlepidoptera, XVI’ A New Genus and Two New Species of Oecophoridae (Lepidoptera) By J. F. Gates Clarke Senior Entomologist, Department of Entomology The first species described herein was received from Dr. F. Luis Gallego M., Chief, Departmento de Entomologia, Facultad de Agro- nomia e Instituto Forestal, Universidad Nacional, Medellin, Colombia, who states that the larvae of the species damage apple by boring in the twigs. A search of the literature and museum collections failed to reveal any described form that approaches this species. The generic affinities are with the Australian Sphyrelata Meyrick. The second species described herein is noteworthy because it is the only species of the family Oecophoridae—a family exhibiting an enor- mous development of species in the Neotropical region—recorded from the island of Dominica, British West Indies. The specimens were collected by my colleagues, my wife, and me during the Archbold- Bredin-Smithsonian Biological Survey of Dominica. The photographs for this paper were made by Mr. Victor Krantz, staff photographer. The drawings of the genitalia of Gonzonota insulana and those of the head, palpus, wing venation, and leg of Maesara 1 See list at end of paper. Pe PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 125 gallegoi were made by Mrs. Elsie H. Froeschner; the genitalia of M/Z. gallegoi were drawn by Mr. André del Campo Pizzini. Maesara, new genus Type-species: Maesara gallegor, new species. Antenna about two-thirds the length of forewing, rather stout, not ciliated; scape with pecten. Labial palpus upturned, second segment slightly roughened beneath; third segment acute, in male much shorter than second; in female nearly as long as second. Max- illary palpus short. Tongue well developed, heavily scaled. Head roughened, sidetufts spreading; ocelli absent. Thorax with posterior crest. Posterior tibia heavily clothed with moderately long hairlike scales. Forewing smooth, termen oblique; costal and dorsal edge nearly parallel in female, in male costa strongly arched, 12 veins; 1b furcate, 2 from near angle of cell; 2, 3, and 4 about equidistant; 6 parallel to 7; 7 and 8 long stalked, 7 to apex; 9 approximate to base of stalk of 7 and 8; 11 from near basal third of cell. Hindwing with 8 veins; 2 from well before angle of cell; 3 and 4 short stalked, from angle of cell, 5, 6, and 7 about equidistant. Male genitalia: Uncus present. Gnathos present, divided. Aedeagus unarmed. Female genitalia: Signa present. Maesara is related very closely to the Australian Sphyrelata Meyrick and keys to it in Meyrick’s key to the genera of Oecophoridae (“Genera Insectorum,”’ 1922, fasc. 180). Maesara differs from Sphyrelata by the smooth, somewhat shorter antenna, the absence of ocelli, the basad origin of vein 11 of the forewing and the short-stalked veins 3 and 4 of the hindwing. Maesara gailegoi, new species Figures 1, 2; Puate 1 Alar expanse 18-36 mm. Labial palpus light ochraceous buff mottled with brown and fuscous. Antenna ochraceous buff basally shading to chestnut brown distad; extreme apex ochraceous buff; scape ochraceous buff shaded with fuscous. Head ochraceous buff somewhat infuscated; side tufts mixed with blackish fuscous. Thorax ochraceous buff, the ground color obscured by chestnut brown and blackish fuscous. Forewing ground color umber brown; costa broadly edged with chestnut brown, this band of color narrowing toward apex; veins in costal half of wing emphasized by chestnut brown; in cell, at one-third, an ochraceous buff spot mixed with a few black scales; on fold, slightly beyond one-third, a similar spot (absent in some specimens); at end of cell a more conspicuous, well-defined, ochraceous buff spot; ochraceous buff NO. 3654 MICROLEPIDOPTERA, XVI—CLARKE 3 scales irregularly and sparsely scattered over surface of wing; cilia composed of alternating ochraceous buff and fuscous groups producing a spotted appearance. Hindwing grayish fuscous, thinly scaled basad ; cilia grayish fuscous mixed subapically with ochraceous buff. Foreleg ochraceous buff; femur and tibia strongly overlaid fuscous; tarsal segments irregularly marked fuscous; midleg ochraceous buff; femur mottled fuscous on outer side; tibia almost wholly overlaid fuscous on outer side except distally; tarsal segments with ochraceous buff Figure 1.—Maesara gallegoi, new species: a, ventral view of male genitalia with left harpe and aedeagus removed; b, aedeagus; c, venation of right wings. reduced to narrow annulations. Hindleg fuscous except ochraceous buff distally on femur and tibia, and tarsal annulations. Abdomen blackish fuscous with a slightly paler lateral stripe; anal tuft ochraceous buff. Male genitalia slides JFGC nos. 10582, 11703. Harpe very broad basally, constricted beyond middle, then dilated into a suboval cucullus; costa with a long, curved, digitate projection before con- striction, and at base a thick pointed process (triangular in cross section). Gnathos divided, consisting of two long elements flattened 4 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 125 Figure 2.— Maesara gallegoi, new species: a, ventral view of female genitalia with bursa copulatrix at right side; b, lateral aspect of head of female showing labial palpus; ¢, labial palpus of male; d, hindleg. PROG, UES] NAT MUS VOL. 1125 CLARKE—PLATE b Maesara gallegoi, new species: a, male (holotype); 4, female, left wings. PROC. U.S. NAT. MUS. VOL. 125 e Gontonota insulana, new species: a, male (holotype), left wings; left wings; c, female (dark form), left wings. CLARKE—PLATE 2 b, female (light form), NO. 8654 MICROLEPIDOPTERA, XVI—CLARKE 5 and dilated distally. Uncus triangular, hood shaped, clothed with stiff setae. Vinculum produced into a point. Tegumen narrow, some- what longer than harpe. Anellus consisting of a narrow ventral band and a strongly developed, broad dorsal tube; laterally, on each edge of the tube a pyramidal process with a granular surface. Aedeagus curved, swollen at middle, unarmed. Female genitalia slides JFGC nos. 10939, 11704. Ostium transverse, oval. Antrum strongly sclerotized. Ductus bursae long, spiraled; posterior third lined with a narrow, longitudinal scobinate band. Bursa copulatrix membranous. Signa 11, in two longitudinal rows of small thorns; lamella antevaginalis and lamella postvaginalis membranous. Holotype: U.S. National Museum No. 69716. Type-locality: Colombia (Antioquia) Rionegro. Distribution: Of this species, Dr. Gallego states (in litt.): ‘In several municipalities of Oriente Antioqueno and other departments of Caldas, Boyaca and Cundinamarca (2200-2600 m.).”’ Food plant: Pyrus malus L. (manzano). Larva boring in the twigs. Described from the holotype male, 7 7 and 18 99 paratypes all from the same locality (September and October dates). Paratypes will be placed in the U.S. National Museum; in the Departmento de Entomologia, Facultad de Agronomia e Instituto Forestal, Universidad Nacional, Medellin, Colombia; and in the British Museum (Natural History). Presumably, this moth is a native Neotropical species; it is interest- ing that it should be found damaging apple, an introduced plant, while its native host is still unknown. The habits and economic significance of this species will be published by Dr. Gallego and his associates. It gives me very great pleasure to dedicate this species to Dr. F. Luis Gallego M., dean of South American entomology. Gonionota Zeller Gonionota insulana, new species Fieure 3; Pate 2 Alar expanse 17-19 mm. Labial palpus orange yellow irrorate with scarlet and brown scales; apex of third segment ocherous white. Antenna light brown suffused reddish in some examples; scape orange yellow irrorate with scarlet. Head orange yellow with scattered scarlet scales along sides and on frons. Thorax orange yellow mixed with scarlet and brown scales. Forewing ground color orange yellow; costa to three-fifths russet; extreme edge narrowly black interrupted by a white streak at two- fifths; on costa, beyond three-fifths an orange yellow spot with extreme 6 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 125 costal edge white; from basal third to costal three-fifths an outwardly oblique, russet band, confluent with dark costal shade, terminating in point, the latter confluent with an outwardly curved series of black Figure 3.—Gonionota insulana, new species: a, ventral view of male genitalia with left harpe and aedeagus removed; b, aedeagus; c, ventral view of female genitalia; d, signum, enlarged. spots; terminal area of forewing russet; discal stigmata, two, blackish, at one-third in cell and at end of cell; irregular lines and blotches scarlet; cilia yellow to tornus then fuscous; underside with scarlet and brown and much white scaling. Hindwing buff shaded with brown NO. 3654 MICROLEPIDOPTERA, XVI—CLARKE i and russet toward edges; cilia russet and brown terminally, buff toward anal angle. Foreleg white overlaid with light brown and scarlet on outer side; midleg white; femur with scattered brown and scarlet scales on outer side; tibia with scarlet dash distally on outer side; tarsus shaded with scarlet distally; hindleg white with scattered brown and scarlet scales; tarsus suffused reddish distally. Abdomen ocherous white dorsally, somewhat more yellowish toward middle; ventrally irrorate with ocherous white, scarlet, and fuscous scales. Male genitalia slide JFGC no. 11681. Harpe subrectangular; cucullus pointed dorsally; at about middle of cucullus a large cluster of heavy spines; sacculus a prominent ridge from base to about middle of harpe. Gnathos an oval, spined knob. Uncus divided into two divergent arms from a central stalk. Vinculum narrow, truncated. Tegumen as long as harpe. Anellus a very narrow plate, dilated distally. Aedeagus stout, curved, with a cluster of bristles distad. Female genitalia slides JFGC nos. 11678, 11679, 11680. Ostium small, slitlike, membranous. Genital plate membranous. Antrum narrowly and very lightly sclerotized. Inception of ductus seminalis from anterior edge of antrum. Ductus bursae membranous. Bursa copulatrix membranous with a small, slightly sclerotized signum, dentate along the edges. Holotype: U.S. National Museum No. 69717. Type-locality: Dominica, .5 mile east of Pont Casse (13 January 1965; J. F. G. and Thelma M. Clarke). Distribution: Dominica. Food plant: Unknown. Described from the holotype male and 7 99 paratypes as follows: 2 99, Springfield Est., 20-26 July 1963; 2 99, .4 mile east of Pont Casse, 23 June 1964, 21 May 1964; 9, 1.6 miles west of Pont Casse, 16 June 1964 (all collected by O. S. Flint); 9, 1 mile east of Pont Casse, 1800 feet, 29 January 1965 (J. F. G. and Thelma M. Clarke); 9, Pont Casse, 7 June 1965 (Don R. Davis). Only eight specimens of insuwlana came to hand during many nights of collecting over a two-year span. Moreover, all were collected within a radius of three miles although the island was covered, during the two-year period, by many collectors. At best the species is rare. This is an unusually interesting species for two reasons: First, it is the only representative of the family Oecophoridae and the sole representative of the genus Gonionota on the island. Both the family and the genus are highly developed in the neotropics yet only the one species has succeeded in establishing itself on Dominica. Second, it is unusually variable, a characteristic seldom found in the genus Goni- onota. In plate 2a I have illustrated the only male available that shows a clearly defined pattern and contrasting coloring. Plate 26 illustrates 8 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 126 a female with a less well-defined pattern and suffused coloring. In the female shown in plate 2c practically the whole pattern has dis- appeared, and the whole upper surface of the wing is overlaid with light brownish scales. Careful examination with a microscope reveals the same markings, though greatly subdued, that are found in the other specimens. In general, the male of insulana appears similar to that of G. comastis (Meyrick), but it is much smaller and lacks the conspicuous, white discal spot of forewing of that species. The evenly colored fe- male (pl. 2c) resembles that of G@. isastra (Meyrick) in color, but it is little more than half the size of that species. In the male genitalia, both isastra and comastis lack the large disc of coarse spines found in insulana. Since the females of isastra and comastis are unknown, no comparison of the genitalia can be made. Neotropical Microlepidoptera Series in the Proceedings (Papers are prepared with the aid of National Science Foundation Grants) paper author subject year volume number I, II Clarke Blastodacnidae and 1962 113 3457 Aegeriidae III Clarke Gonionota melobaphes and 1964 115 3480 relatives TV Duckworth A new genus of Stenomidae 1964 116 3497 VY Obraztsov The tortricid genus Proeulia 1964 116 3501 VI Clarke The genera Orsotricha and 1964 116 3502 Palinorsa VIL Obraztsov The tortricid genus 1966 118 3527 Pseudomeritastis VIII Duckworth The stenomid genus 1966 118 3531 Falculina IX Obraztsov The tortricid genus 1966 118 3535 Pseudatteria X Duekworth Taxa erroneously placed in 1966 119 3540 Stenomidae XI Obraztsov The tortricid genus 1966 119 3543 Idolatteria XII Duckworth The stenomid genus Lethata 1967 122 3585 XIII Duckworth The stenomid genus 1967 122 3590 Loxotoma XIV Clarke Chilean Microlepidoptera 1967 122 3591 described by Blanchard XV Duckworth The stenomid genus 1967 123 3620 Thioscelis U.S, GOVERNMENT PRINTING OFFICE: 1968 Proceedings of the United States National Museum SMITHSONIAN INSTITUTION - WASHINGTON, D.C. Volume 125 1968 Number 3655 Parallel Evolution in the Small Species of Indicator (Aves) By Herbert Friedmann ' é Ornithologist Emeritus, Department of Vertebrate Zoology One of the aspects of avian taxonomy that inevitably has to wait on the gradual amassing of considerable material is the appreciation, and the consequent elucidation, of sympatric sibling species. Until suf- ficient numbers of specimens become available, the investigator can only treat the minor differences that he may notice as inconsistent, but apparently individual, variations in a not completely homogeneous taxon. The genus /ndicator presents an unusual array of sympatric, closely similar species distinguished primarily by size. In the Acacia grasslands of much of eastern Africa there are three similar species (in descending order of size): minor, meliphilus, and narokensis. In the forests of western and central Africa there are conirostris (a race of minor), exilis, willcocksi, and pumilio. Not only are the species of the two groups similar in appearance, but their habits vary but slightly. Chapin (1962) found pumilio and exilis (pachyrhynchus) together at Tshibati and saw no difference in their behavior. We can infer that minor and meliphilus also must be extremely similar in their actions and vocalisms, or variances would have been recorded by the numerous observers who 1 Director, Los Angeles County Museum of Natural History, Los Angeles, California 90007. 1 2, PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 have written about them. Some collectors were not aware that there. were two species involved until they studied the dried specimens after their field experience. 2) MOIST WOODLANDS AND SAVANNAS Ficure 1.—Simplified vegetation map of Africa. The large equatorial forest area is the home of the following honeyguides discussed in this paper: maculatus, conirostris, exilis, willcocksi, and, at its eastern end, chiefly at higher altitudes, of pumilio. The woodlands and sa- vannahs are the home of vartegatus, minor, meliphilus, and narokensis. Through the cooperation of the American Museum of Natural History, the Field Museum, and the National Museums of Rhodesia, together with the now considerable holdings of the Los Angeles County Museum of Natural History, plus earlier study of the specimens in the Royal Museum of Natural History in Brussels, the British Museum (Natural History), the Musée d’Histoire Naturelle of La Chaux-de- Fonds, Switzerland, and the United States National Museum, I have NO. 3655 PARALLEL EVOLUTION—FRIEDMANN 3 been able to study in depth the morphology and distribution of the smaller Indicator species. Some of the specimens were collected under National Science Foundation Grant GB-5107. The exilis-willcocksi Relationship The small west African forest honey-guides, erilis and willcocksi, were considered conspecific prior to Chapin’s 1962 study when, for the first time, the specific identity of the latter sibling form was elucidated. Until then willcocksi had been treated as a western race (Ghana and Togoland) of ezilis; the fact that willcocksi-like indi- viduals occurred here and there throughout the range of nominate exilis was, if anything, looked upon as evidence that the latter race provided the variational trends that had become “‘solidified”’ in will- cockst. A good number of specimens of willcocksi collected in 1966 and 1967, along with those of ezilis (pachyrhynchus) in extreme western Uganda (Kibale and Impenetrable Forests), clearly upholds Chapin’s con- clusions in regard to the distinctness of the two sympatric species. When series are laid out, the differences between the species is more marked than that suggested by single examples. Chapin’s attention was brought to bear on the ezilis-willcockst situation as a result of his discovery and description of the still smaller, partly sympatric pumilio. While the validity of pumilio as a species distinct from its sympatric congeners is clear as has been universally accepted by all students of African birds, Chapin (1958, p. 47) did not pursue the question of its relation to the paler, small Indicator species of the open country of eastern Africa beyond commenting that the very small beak of pumilio caused him to think at first that it might be conspecific with meliphilus. The striking difference in coloration of the two species persuaded him to name pumilio bi- nomially, but he felt constrained to add that he was still of the opinion that “its small beak may well indicate relationship with mel- apis? The meliphilus-narokensis Relationship Just as willcocksi had been looked upon as a part of the variational limits of ezilis until sufficient series of specimens became available for study, meliphilus too has been assumed to comprise within its extremes, the species appelator Vincent (1933) and narokensis Jackson (1906). Ever since Chapin’s discovery of pumilio in the Kivu forests I have wondered whether or not meliphilus, like willcocksi, might also have a smaller, sympatric sibling form. The description and dimensions of appelator, described from Zobué on the border of Mozambique and Malawi, were too close to those of meliphilus to 4 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 cause any concern, but narokensis was described as much smaller > than the average meliphilus as pumilio was than the average ezilis | (or willcocksi). As long as narokensis, however, was known from only | one or two odd specimens, to decide if it was a taxon in its own right — or merely the smallest specimens of meliphilus was impossible. Accord- _ ingly, in three different publications (1954, 1955, 1958) I could treat | it only as a probable synonym of meliphilus, which, until 1958, was still considered to be an eastern, pale, gray-breasted race of evilis. As long ago as 1938 Grant and Mackworth-Praed (pp. 143-144) examined van Someren’s pair of narokensis from Mt. Moroto and | found that the two specimens agreed with Jackson’s type from Mt. Doinyo Narok and with Granvik’s specimen form Kacheliba. Grant and Mackworth-Praed concluded that comparison with examples of | meliphilus ‘clearly show that J. narokensis is a distinct species having a smaller bill and a smaller wing measurement ... . It is a very re- markable fact that there are two birds so very similar in coloration. . . and, although they both occur in the same general area in Kenya Colony, it may be found that they inhabit different types of country, as Granvik records J. narokensis in tall acacias in dry open country, and Moreau records I. e. meliphilus in the vestiges of coastal hOres tic wier.t I now feel certain that only lack of critical specimens in most of the large museums caused Grant’s and Mackworth-Praed’s con- clusion to be overlooked. In the absence of such material, it had been impossible to test it. At the time of their report, their conclusion seemed too remarkable to be likely, and I left narokensis in the un- verified synonymy of meliphilus. White (1965) does not even mention narokensis, either as a valid taxon or as a synonym—an indication that he was content to go along with current disbelief in it. The re- markable, and hence unlikely aspect of a distinct species, narokensis, being sympatric with meliphilus has, of course, been changed com- pletely by Chapin’s findings of three sympatric sibling Indicator species in the Congo. In the light of his results, it is not surprising that a comparable situation should exist among the small Indicator species of eastern Africa. In 1957 Mackworth-Praed and Grant (p. 745) again treated naro- kensis as a species, smaller and generally paler in color than meli- philus, and listed a number of specimen-based locality records: Mt. Moroto in eastern Uganda; Kacheliba in Turkanaland, northwestern Kenya; Mt. Doinyo Narok, Sokoke Forest, and Malindi, all in southern Kenya; and Lushoto, in northeastern Tanzania. Subse- quently, I have seen other records that have reported the species from Shimba Hills, near Kwale, and southern Guaso Njero, in southern Kenya, and from Sigor, West Pokot, western Kenya. With these No. 3655 PARALLEL EVOLUTION—FRIEDMANN 5 last three examples, it became obvious that there were indeed two very similar species living side by side in eastern equatorial Africa. This caused me to assemble as complete a series as possible in order to review the situation again and to seek new conclusions. Although very similar to each other, narokensis and meliphilus should be treated as specifically distinct. In narokensis, the wing length varies from 65 to 70 mm in four females, 67 and 69 mm in two males; in meliphilus, 18 females have wing lengths from 69 to 78.5 mm, 18 males, from 73.5 to 85.5 mm. The bill is ‘“stubbier’’ to the eye in narokensis, more so than actual measurements reveal; length from distal end of nostril to tip of culmen is 4.2 to 4.8 mm (4.7 to 5.9 mm in meliphilus), the exposed culmen is 5.7 to 7 mm (7.1 to 8.5 mm in meliphilus). Actually, the bills of narokensis are very similar to those of pumilio. This supports Chapin’s comment, cited above, that pumilio may be related not merely, as he put it, to meliphilus, but even more closely to that part of meliphilus (olim) now separated as narokensis. In my 1958 paper I explored the question of the status of the western race of meliphilus, which Monard (1934, p. 55) had des- ignated angolensis as a subspecies of minor! I still believe that there are two recognizable races of meliphilus, although they are based on average differences and not on more trenchant characters; angolensis is usually paler gray, especially on the throat, breast, and upper abdomen than nominate meliphilus. Chapin (1954, p. 633; 1962, p. 44) considered angolensis indistinguishable from meliphilus, but he also put narokensis in this category. More recently, Traylor (1963, p. 108) has supported the belief that angolensis is the south- western race of meliphilus, and I may add that the additional speci- mens I have seen since my 1958 paper also have supported this division. The removal of narokensis from meliphilus increases the average duskiness of the residual nominate meliphilus since the palest members of that taxon, as formerly treated, are all narokensis. The narokensis-pumilio Relationship The relationship of narokensis and pumilio parallels very closely that of meliphilus and exilis and that of minor and conirostris. Inas- much as meliphilus and exilis have been found to overlap sympatri- cally in the Mwinilunga district, Zambia, and in the Upemba Park, in the Congo (Verheyen, 1953, pp. 406-407), their specific distinctness has to be granted. The situation between minor and conirostris, however does not include this convenient sympatry, and they usually are still considered to be races of one species. While conirostris is a dark-colored forest-inhabiting form with a discontinuous range from western Kenya (Kakamega and Kapenguria) west to southern 282-405—68——2 6 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 Cameroon and Gabon, typical minor is the bird of the open country of southern and eastern Africa, west and south of the Congo forest to Angola. It now seems that pwmilio is the dark forest counterpart (from the eastern Congo, discontinuously to Kakamega, in western Kenya) of the pale narokensis, which is now known to occur from Turkanaland south to northeastern Tanzania and west to the Upemba Park in the Congo. @ MELIPHILUS @® NAROKENSIS A PUMILIO Ficure 2.—Specimen records of the three small species of Indicator whose relationships are reinterpreted in this paper. The subspecies conirostris is not wholly constant in its color characters; in fact, in parts of western Africa, especially in Upper Guinea, it and its geographic replacement riggenbachi frequently show some inter- gradation toward the adjacent, paler forms of minor—alezanderi, NO. 3655 PARALLEL EVOLUTION—FRIEDMANN 7 ussheri, and pallidus, some of which are such intermediates as to be doubtfully recognizable as racial entities. Similarly, pumilio has been found to be geographically variable. Its data are much less extensive, being based merely on the fact that its easternmost example (Kakamega Forest, western Kenya) is different from Kivu topotypes, a difference that was sufficient to cause Williams and Friedmann (1965) to raise the possibility of the Kakamega bird being racially separable from the eastern Congo population. We found the single Kakamega specimen to be slightly larger (wing 70 mm), to have more uniformly grayish underparts without the streaks that are present in the birds from the Kivu and the Impenetrable Forest, southwestern Uganda, and to have the top and sides of the head less greenish, more grayish. If additional material from Kakamega should agree in these color characters, which are in the direction of a more grayish and paler shade—or, to an extremely slight, almost incipient degree, in the direction of the very much paler and grayer narokensis—there would be a situation faintly resembling that ex- isting in conirostris and riggenbachi in Upper Guinea and other races of minor in the open country north of the forest belt. Kakamega, where pumilio occurs, and Sigor, West Pokot, where narokensis has been taken, are barely 75 miles apart. Enough has been said to warrant raising the question whether or not pumilio is a true species or an unusually distinct race of narokensis. To be wholly consistent, I would have to treat the two as conspecific forms by the same standards that I use when considering conirostris and minor in this matter. The real question is whether they are repre- sentative races or whether they have differentiated to the point where they are specifically distinct. Unfortunately, this question could be answered only by bringing them into contact under natural con- ditions, which cannot be done. I, therefore, favor considering the two as races of a single species merely to point out their mutually representative nature and to suggest their closer relationship in the not too distant past. That they are closely allied is intimated by anal- ogy with minor and conirostris, and while this is certainly no proof, it may yet reveal a pattern or trend in these small species of Indicator. Without pressing the point beyond its realistic implications, I add the suggestion that maculatus (with stictithorar as a race) is no more than a west African forest representative of variegatus of eastern and southern Africa. It is a comparable case. Morphological Overlap and Sympatry in Sibling Species While I conclude that narokensis is a species distinct from meliphilus, I realize that other observers may have the experience of finding specimens that are difficult to place in either species. It would be 8 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 comforting to find reliable and rigid key characters by which to separate narokensis from small meliphilus, but this is not always feasible. A similar state of affairs occurs, however, with such generally accepted species as extlis (pachyrhynchus) and minor. Small individuals of the latter often are difficult to distinguish from examples of the former, but this does not cause serious doubts as to their specific distinctness. In these birds, not only is there a serial gradation in size among sympatric species, but also the very limits of the species are at times difficult to distinguish. The resulting situation makes one wish for more observational data regarding the critical isolating mechanisms involved. In the case of pumilio and exilis (pachyrhyn- chus), I was able to find skeletal characters that corroborated their external specific distinctness, but these differences are not necessarily isolating mechanisms (1963). Benson and Irwin (1964, p. 110) write that meliphilus has a thinner, softer skin than ezilis and other local species of Indicator. They even suggest that, in this respect, meli- philus is more like Prodotiscus than like other species of Indicator. So far, however, no pertinent observations on the feeding habits of meliphilus have been brought forth that indicate whether or not it is more or less given to feeding on small insects generally than are other species of Jndicator. It is known that it does feed on bee comb like its congeners and unlike Prodotiscus. Sympatry and Parallelism in Sibling Species Considering the fact that sibling species, or even species of only approximately similar external characters, usually are related fairly closely, and considering the further fact that the origin and establish- ment of the discontinuities between such species ordinarily involve allopatry of some duration, it is noteworthy how quickly and how extensively some of these small Jndicator species once more have become sympatric. This fact argues for effective, but as yet unrecog- nized, isolating mechanisms. The number of instances of two or more of these morphologically similar honey-guides occurring together is very impressive—Kabompo District, Zambia: eatlis (pachrhynchus) and meliphilus (Benson and Irwin, 1965, p. 4); Tshibati, Kivu District, Congo: pumilio and exilis (pachyrhynchus) (Chapin, 1958, p. 48); Doinyo Narok, Kenya: meliphilus (specimen in American Museum of Natural History, collected by Jackson) and narokensis (Jackson, 1906, p. 20); Kibale Forest, west Uganda: exilis, willcocksi, conirostris; Impenetrable Forest, west Uganda: willeocksi, exilis pumilio; Sigor, West Pokot, western Kenya: narokensis, meliphilus; Mt. Moroto: minor, narokensis; Victoria, British Cameroons: minor (controstris), exilis, willcocksi (Serle, 1965, p. 77). NO. 3655 PARALLEL EVOLUTION—-FRIEDMANN 9 Here, then, is a series of remarkably parallel situations of similarly colored, sympatric species, each with a paler, grayer form in the open grasslands of eastern Africa and each with a darker, greener represent- ative in the forests of central and western Africa. In some, the repre- sentatives are clearly conspecific; in others, they have achieved specific distinction—minor and conirostris; meliphilus, willcocksi, and exilis; narokensis and pumilio; variegatus and maculatus. It is, of course, impossible to state with any certainty either that these similar species all originated as greenish forest birds that gave rise to paler, grayer races in the open country, or that all began as erayish denizens of the savannas and each evolved a darker, greenish representative form in the wooded areas. It is equally possible that some may have gone one way, and others, the opposite direction; but the end result, which is all we have for study, shows a remarkable parallelism. As Mayr (1963, p. 609) noted, true parallelism is due to the necessarily similar response of a common genetic heritage to similar selection pressures. Inasmuch as all of these species of honey-guides seem in every way to be closely related, and, hence, to have a basically similar genetic composition, their parallel evolutionary picture seems to be true parallelism rather than mere convergence, a situation wherein similar phenotypic developments have resulted as a response to similar environmental selection factors. The small species of Indicator have, at best, a very restricted phenotypic potential, much like some of the tyrant flycatchers of the genera Empidonaz and Elaenia. Literature Cited Benson, C. W., and Irwin, M. P. S. 1964. Some additions and corrections to “A check list of birds of Northern Rhodesia.’”’ Occas. Pap. Nat. Mus. Southern Rhodesia, no. 27B, pp. 106-127. 1965. Some birds from the north-western province, Zambia. Arnoldia, vol. 1, no. 29, pp. 1-11. CuaPin, J. P. 1954. The birds of the Belgian Congo, pt. 4. American Mus. Nat. Hist. Bull. 75B, 846 pp. 1958. A new honey-guide from the Kivu District, Belgian Congo. Bull. British Ornith. Club, vol. 78, pp. 46-48. 1962. Sibling species of small African honey-guides. Ibis, vol. 104, pp. 40-44. FRIEDMANN, H. 1954. A revision of the classification of the honey-guides, Indicatoridae. Ann. Mus. Congo, Tervuren, Zool., vol. 50, pp. 21-27. 1955. The honey-guides. U.S. Nat. Mus. Bull. 208, 292 pp. 1958. The status of the gray-breasted least honey-guides. Proc. Biol. Soc. Washington, vol. 71, pp. 65-68. 1963. Morphological data on two sibling species of small honey-guides, Los Angeles Co. Mus. Contr. Sci., no. 79, pp. 1-5, 10 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 Grant, C. H. B., and MackwortH-PRAgED, C. W. 1938. On the status of A. Melignothes pachyrhynchus, B. Indicator pyg- maeus, C. Melignothes meliphilus, D. Indicator narokensis, and EK. Indicator appelator. Bull. British Ornith. Club, vol. 58, pp. 141-145. JacKsoN, F. J. 1906. Description of new species of birds from equatorial Africa. Bull. British Ornith. Club, vol. 19, pp. 19-20. MackKworTu-PrRAED, C. W., and Grant, C. H. B. 1957. Birds of eastern and northeastern Africa, vol. 1, ed. 2, 845 pp. Mayr, E. 1963. Animal species and evolution, 797 pp. Monarp, A. 1934. Ornithologie de PAngola. Arq. Mus. Bocage, vol. 5, pp. 1-107. SERLE, W. 1965. A third contribution to the ornithology of the British Cameroons. Ibis, vol. 107, pp. 60-94. Traytor, M. A. 1963. Check list of Angola birds. Mus. Dundo Publ. Cult., no. 61, 250 pp. VERHEYEN, R. 1953. Oiseaux. Jn Exploration du pare national de l’?Upemba, Mission G. F. De Witte, fase. 19, 687 pp. VINCENT, J. 1933. Descriptions of new species and subspecies of birds collected in Portuguese East Africa. Bull. British Ornith. Club, vol. 53, pp. 130-131. WuitTz, C. M. N. 1965. A revised check list of African non-passerine birds, 299 pp. WiuuiaMs, J. G., and FriepMaAnn, H. 1965. The pygmy honey-guide, Indicator pumilio Chapin, in East Africa. Bull. British Ornith. Club, vol. 85, pp. 21-22. U.S. GOVERNMENT PRINTING OFFICE: 1968 Proceedings of the United States National Museum SMITHSONIAN INSTITUTION «+ WASHINGTON, D.C. Volume 125 1968 Number 3656 Copepod Crustaceans Parasitic on Fishes of Eniwetok Atoll ’ By Alan G. Lewis ? Copepod parasites of fishes of Eniwetok Atoll, one of the Marshall Islands, have not been previously studied. The only reference dealing with parasitic copepods from this general region of the Pacific is that of Heegaard (1943) concerning some caligids from the Gilbert Islands. Both Heegaard (1940, 1962) and Kabata (1964, et al.), however, have described copepods from fishes from the neighboring Australian region, some of which are also found on fishes from Eniwetok Atoll, and Hewitt (1963, et al.) and Thomson (1889) have described copepods from fishes of New Zealand. These studies, as well as those of Lewis (1964a, et al.) concerning copepods from Hawaiian fishes, cover most of the named species although some of them have previously been described from other areas, primarily the western Pacific and the Indian Ocean. The study herein reported is based on two collections made by the author during the summers of 1959 and 1961. He is indebted to the U.S. Atomic Energy Commission for the use of the facilities at the Eniwetok Marine Biological Laboratory. He is also grateful for the 1 This study was supported by a grant (GB-3932) from the National Science Foundation. 2 Associate Professor of Zoology and Oceanography, Institute of Oceanography, University of British Columbia, Vancouver, B.C., Canada. ah 2, PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 assistance given to him by James Coatsworth, John Shoup, and David Au, who were serving as research assistants at the Eniwetok Marine Biological Laboratory during the periods when the collections were made. Finally, the author appreciates the help given to him by members of the Division of Crustacea of the Smithsonian Institution during the time of final examination of the specimens in the collections. Metuops.—The external surface, gill cavities, buccal cavity, and nasal cavities of the hosts were examined for parasitic copepods. Copepods collected were killed and preserved in 95 percent ethyl alcohol. Specimens to be drawn or dissected were placed in 85 per- cent lactic acid to clear and soften them, stained with Chlorazol Black E dissolved in 85 percent lactic acid, and then placed in benzyl alcohol for final clearing and for dissection and drawing. Drawings of the entire animal were made from specimens placed in benzyl alcohol and covered with a cover slip, supported so that the shape of the organism was not distorted. A camera lucida was used in making the drawings. The appendages and processes were TaBLE 1.—Armature of hypothetical thoracic leg shown in figure 1 Inter- Protopodite Exopodite Endopodite Leg | Surface 0 Ga] | Dae eee Se eee ae ee ee ee Plate 2 1 2 3 1 2 3 Outer m,h,rh,r r,D,d,dh H fH,Z,mH, C c mp’,mp,Q dmH. Inner a,SSS,S, e,fm,H CE Cl,cl,dH Pp p,p’ 2PL@? dm,fm drawn in situ or were removed and mounted in either Hoyer’s mount- ing medium or a 1:1 mixture of Turtox’s CMC-10 and CMC-S. Measurements were made with an ocular micrometer on specimens softened in lactic acid and held loosely in place by a cover slip sup- ported by spacers. In the figures, the 2 and & signs are used separately under each drawing to indicate a difference between the appendage or body part of the female and that of the male. The symbols are used to- gether (2 o) to indicate the similarity of the appendage or body part in both sexes. In the latter case the sex of the specimen from which the drawing was made is indicated by a line under the appropriate symbol. TrerMInoLocy.—The term “cephalothorax” is used to indicate a condition in which one or more of the thoracic segments are fused with the cephalon. The maxilliped-bearing segment is considered as the first thoracic segment. The term ‘‘pedigerous segment” is used to indicate a leg-bearing thoracic segment, while the terms “free thoracic NO. 3656 COPEPODS—LEWIS 3 ays RUAN Ficure 1.—Hypothetical thoracic leg showing the various armament components (a=ad- hesion pad; C=large plumosities; c=small plumosities; Cl=large spikelike extension of segment; cl=small spikelike extension of segment; D=large denticulations; d=small denticulations; dH=large spine with denticulations; dh=small spine with denticulations; dm=denticulated membrane; dmH=large spine with denticulated membrane or mem- branes; fH=large spine with frilled membrane or membranes; fm=frilled membrane; H=large spine; h=small spine; [P=interpodal plate; m=membrane; mH=large spine with simple membrane or membranes; mp=setule with membrane on both sides; mp’= setule with membrane on one side, naked on other; n 1,2,3=endopodite segments one, two, three; P=plumose seta; P’=naked seta; PROT=protopodite; p=plumose setule; p’=naked setule; Q=seta with plumosities on one side, membrane on other; Q’ =seta with plumosities on one side, naked on other; r=numerous minute projections giving rugose appearance; rh=spinule; s=hairlike process; x 1,2,3=exopodite segments one, two, three; Z=spine with frilled membrane on one side, simple membrane on other) . 4 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 segments” and “free pedigerous segments” are used to designate those thoracic segments that are not fused with the cephalon. The term “oenital segment” is used, with some reservation, to designate the fused sixth and seventh thoracic segments (= fifth and sixth pedigerous segments) in the Caligidea (Caligoidea of Yamaguti, 1963). The term “ovigerous” is used to indicate a female with egg strings, the term “nonovigerous” to indicate a female without egg strings even though the genital segment may contain eggs. The terminology applied to the appendages and processes is the same as that in Lewis (1967). To facilitate the use of the thoracic leg tables, a hypothetical thoracic leg is shown in figure 1, giving the various thoracic leg armature elements used in the tables; an analysis of the hypothetical thoracic leg is given in table 1. Order CYCLOPOIDA Family BOMOLOCHIDAE Nothobomolochus gibber (Shiino) FIGuRES 2-4 Bomolochus gibber Shiino, 1957, p. 411, figs. 1-2. Nothobomolochus gibber (Shiino).—Vervoort, 1962, p. 70. Pseudartacolax gibber (Shiino).— Yamaguti, 1963, p. 14, pl. 8, fig. 1. Host AND DISTRIBUTION.—Tylosurus giganteus, Japan. MatTeErRIAL.—8 females and 2 males (USNM 120739) from the gill cavity of Belones platyura Bennett. MEASUREMENTS (in mm).—7 females and 2 males: female mean (range) male Greatest length, excluding caudal setae 1. 82 1. 71-2, 02 AE Sees Length of prosome 1.15 1. 03-1.30 1.01, 0.95 Width of prosome 0. 98 0.90-1.08 0.81, 0. 86 Length of cephalothorax 0. 67 0. 58-0. 74 0.50, 0. 58 Length of genital segment 0. 21 0. 18-0.25 0.16, 0. 14 Width of genital segment 0. 29 0. 22-0. 36 0.29, 0. 25 Length of egg sac (6 sacs) 1, 23 0. 99-1. 37 Description.—Cephalothorax of ovigerous female (fig. 2a) wider than long, consisting of cephalon and first 2 thoracic segments. Anterior end of cephalothorax forming 2 bilobed projections separated by rounded dorsal and bilobed ventral median surfaces. Lateral margins irregular, lateral posterior margins lobate. Dorsal cephalo- thoracic surface arched, with anterior median fissure. Width of free second pedigerous segment slightly more than two and one-half times the length, slightly more than half the width of cephalothorax. Lateral surfaces winglike, extending slightly past base of second NO. 3656 COPEPODS—LEWIS 5 thoracic leg. Third and fourth pedigerous segments separable ventrally, fused dorsally; dorsal surface topshaped although irregular, greatest width slightly more than that of second pedigerous segment; posterior surface overlapping fifth pedigerous (first urosomal) segment. b+ ++ +4 +4 pos ep 1 mm.g¢e O01 mm.g Ficure 2.—Nothobomolochus gibber (Shiino, 1957): a, female dorsal view; b, male, dorsal view; c, male, genital segment, ventral view; d, female, genital segment and part of egg string, ventral view; e, same, dorsal view; f, caudal ramus, ventral view. 6 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 Female fifth pedigerous segment slightly less than half the width of third and fourth, widest medially, in region of leg attachment. Genital segment (figs. 2d, e) broadest medially, tapered to posterior and anterior ends. Oviducal openings situated on lateral dorsal surface, region around openings ornamented with heavily sclerotized platelets. Sixth legs situated on dorsal surface, adjacent to oviducal openings, consisting of 3 lightly plumose setae originating from nodule. First segment of female abdomen subrectangular, length approxi- mately three-fourths that of genital segment. Second segment sub- rectangular, four-fifths the length of first. Third segment slightly longer than second, tapered posteriorly, posterior surface with bilobed anal indentation. Caudal ramus (fig. 2f) subconical, bearing 3 naked or lightly plumose setules on outer surface and large, lightly plumose seta as well as 2 lightly plumose setules from distal surface. Cephalothorax and free second pedigerous segment of male (fig. 26) generally similar to those of female. Third and fourth pedigerous segments distinct, not fused as in female. Both segments of about equal length, third approximately one and one-third times the width of fourth. Fifth pedigerous, genital, and abdominal segments similar to those of female. Female and male antennule (fig. 3a) 5-segmented, attached to ventral surface of inner lobe of bilobed projection at anterior end of cephalothorax. Segment lengths, in micra, 205:135:90:95:100. First segment with base at right angles to rest of segment and appendage, bearing 14 lightly plumose setae along anterior surface, 3 naked setules on posterior distal surface. Second through fifth segments rodlike; second with 6 lightly plumose and 3 naked setules, third with 3 naked setules, fourth with 2 naked setae on distal surface. Fifth segment with single, naked setule on distal posterior surface, also with 3 naked setules and one naked seta on distal surface. Female and male antenna (fig. 3b) 3-segmented, attached slightly posterior and lateral to antennule base. First segment approximately equal to combined lengths of remaining two, tapered towards distal end, with single naked seta distally. Second segment short, unarmed. Third segment with ventrally facing rows of clawlike denticulations along entire surface and extending on toelike projection of distal surface. Distal surface also with 2 denticulated spines (one not shown in figure), one clawlike spine, and 5 clawlike setae. Female and male mandible (figs. 3c,d) 1-segmented, basal region enlarged, with heavily sclerotized medial ridge; distal half rodlike. Distal end of segment bearing 1 large and 1 small, flattened spines serrated along anterior margins. Maxillule (fig. 3c) nodular, situated immediately posterior to mandible base, with 2 plumose setae and 2 plumose setules. Maxilla (figs. 3c,e) 2-segmented, proximal segment NO. 3656 COPEPODS—LEWIS 7 whe TET eit ad y at Ab hth Ficure 3.—Nothobomolochus gibber (Shiino, 1957), female, ventral views: a, left antennule; b, right antenna; c, left side of oral region, showing labrum, mandible, paragnath, maxil- lule and maxilla; d, right mandible; ¢, right maxilla; f, left maxilliped. 8 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 large, irregular; second segment short, rodlike, bearing 2 stiffly plumose terminal spines and 1 naked subterminal setule. Labrum (fig. 3c) apron-shaped, anterior surface with very fine projections giving fuzzy appearance to surface. Paragnath (fig. 3c) situated medial and poste- rior to maxillule base, bladeshaped, distal half of margins finely denticulated. Anteriorly V-shaped, posteriorly rounded padlike projec- SSN ae <= eet tt 0.2 mm. Qo . Ce O2 mim: oe Figure 4.—WNothobomolochus gibber (Shiino, 1957), thoracic legs: a, left first, ventral view; b, left second, anterior view; c, right third, anterior view; d, right fourth, anterior view; e, right fifth, ventral view. No. 3656 COPEPODS—LEWIS 9 tion present immediately posterior to maxilla, extending between maxillae bases. Female and male maxilliped (fig. 3f) 2-segmented, base situated lateral to maxilla base, body extending anteriorly to antenna base. First segment strongly developed, broad proximally, tapered distally, bearing single, long, finely plumose seta from knob on distal inner surface. Second segment short, inner surface cuplike, outer surface rounded; distal surface with large, sharply curved claw and finely plumose seta. Thoracic legs 1-4 biramous, fifth leg uniramous. Ramal count 3-3 on first 4 legs. For nature of legs and armature, see figure 4 and table 2. TaBLE 2.—Armature of thoracic legs I-V of the female and male of Nothobomolochus gibber (Shino, 1957) Inter- Exopodite Endopodite Leg | Surface poda! |Protopodite Plate | | 1 2 3 1 2 3 I} Outer ss dh dh dh Cc ¢ e,5P Cc Inner P,sss ic 3P 3P 12) c,P c II} Outer ¢,p c,fH fH 3fH,2P c c ¢,2h c Inner ss P 4P 1 C2P; ¢,3P III} Outer fH fH 2fH ,2P e c c,2h Inner 12) 12 4P 12 ¢,2P c,2P IV| Outer fH fH 2iH2P c c c,h,P Inner 12 4P 12) 12 h V | Outer* p D,r,p Inner r,2p *The designations ‘‘protopodite’”’ and ‘‘exopodite’”’ are used with some reservation. Discussion.—The Eniwetok specimens differ from Shiino’s original description (1957) in several respects: 1. The total length is only 80 percent of that given for the holotype (2.28 mm for Shiino’s holotype specimen, 1.82 mm average for the Eniwetok specimens) although Shiino (1957, p. 416) indicates that the holotype is the largest specimen in his collection. Other measure- ments also differ although it is suggested that these differences are due to both the size of the holotype and the effect of natural variation. 2. The caudal rami of the Eniwetok specimens have 3 setules on the outer surface while Shiino indicates only 2. The third setule, however, may have been overlooked as all 3 are small and almost transparent. 10 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 3. Part of the surface of the labrum has a “fuzzy” appearance due to numerous, minute projections. Shino figures the labrum without the projections. 4. Shino states (p. 415) that, in the first thoracic leg, “both the rami (are) fringed by stout long spines with multiserial barbs; . . .” This condition was not evident in the Eniwetok material. 5. The antennae of the Eniwetok specimens possess 2 denticulated subterminal setae on the third segment while Shiino indicates that there is only 1. In spite of these differences, the Eniwetok specimens agree closely with Shiino’s original description and, based on this similarity, are believed to be conspecific with N. gibber. Pseudotaeniacanthus species Ficures 5, 6 MatTeriAL.—1 adult male (USNM 120740) from the external surface of Acanthurus gahhm (Forskal). MEASUREMENTS (in mm).—1 male: Greatest length, excluding caudal setae 1. 73 Length of prosome 0. 81 Width of prosome 0. 63 Length of cephalothorax 0. 49 Length of genital segment 0. 18 Width of genital segment 0. 27 Derscription.—Body (fig. 5a) of general cyclopoid shape: cephalo- thorax of general bell shape, consisting of cephalon, maxilliped- bearing and first pedigerous segments. Anterior end of cephalothorax rounded although slightly concave medially, set off from rest of cephalothorax by lateral indentation and line of heavy sclerotization extending medially and curving posteriorly from lateral indentation. Lateral margins flatly convex, posterior margin irregularly concave. Second pedigerous segment distinct from cephalothorax, width approximately nine-tenths that of cephalothorax; lateral surfaces winglike, lateral margins flatly convex. Third pedigerous segment distinct from second, appearing slightly narrower than second although of similar shape. Fourth pedigerous segment distinct from third, approximately three-fourths its length, lateral surfaces winglike, as in preceding free pedigerous segments, although lateral margins more rounded. Fifth pedigerous segment without winglike lateral projections, forming first segment of urosome; slightly narrower anteriorly than posteriorly. Genital segment (fig. 56) barrel-shaped, slightly more than one and one-half times the length of fifth pedigerous segment; No. 3656 COPEPODS—LEWIS 11 ——s er tn eee Ficure 5.—Pseudotaeniacanthus species, male: a, dorsal view; }, left side of genital segment ventral view; c, left caudal ramus, ventral view; d, left antennule, ventral view; ¢, antenna, region showing antennule base (A-1), antennal base (A-2), and adjacent processes; f, right antenna, posterior view. 12 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 lateral posterior region lobate, lobes bearing genital openings and sixth legs. Sixth leg consisting of single finely plumose setule. Four segments present posterior to genital segment, first approximately equal to length of genital segment, second approximately nine-tenths its length, third and fourth approximately three-fourths its length. Posterior segments narrower than anterior, lateral margins of first 3 almost parallel, fourth tapered posteriorly, posterior margin with small anal indentation and distinct lateral concavities, at place of attachment and articulation of caudal rami. Caudal rami (fig. 5c) elongate, subrectangular, with setule-bearing indentation in distal half of outer surface; distal surface with 3 naked setae and 1 naked setule, additional naked setule present on dorsal surface just proximal to distal end. Antennule (fig. 5d) 6-segmented, attached to anterior-lateral ven- tral surface of cephalothorax, adjacent to small, spikeshaped projec- tion from lateral anterior surface of cephalothorax. First segment (may represent 2 fused segments) strongly flattened proximally, rodlike distally; bearing 19 naked or very lightly plumose setules. Second segment approximately one-third the length of first, with 4 naked or lightly plumose setules. Third segment approximately half the length of first, with 3 naked setules. Fourth segment slightly more than one-third the length of first, bearing 3 naked setules. Fifth segment slightly less than one-third the length of first, with 2 naked setules. Sixth segment slightly more than one-third the length of first, rounded distally, bearing single naked setule from medial posterior surface, 5 naked setules from distal surface. Antenna (fig. 5f) 4- segmented, situated posterior and medial to antennule base. First segment strongly developed, distal surface with naked setule. Second segment approximately half the length of first, subrectangular. Third segment, excluding terminal processes, slightly shorter than second with deep indentation in medial outer surface; distal surface with 2 spatulate and one clawlike projection, inner lateral surface of segment and spatulate process with denticulated membrane or individual denticulations. Fourth segment arising from indentation in third, rodlike, bearing 7 naked setae and setules distally. Mandible (fig. 6a) 1-segmented, broad proximally, irregularly tapered distally, with spineshaped projection distally; second spine- shaped projection arising from distal half of posterior surface; both projections with denticulations, distalmost with single longitudinal row on posterior surface, proximal projection with 2 rows. Maxillule (fig. 6a) nodular, situated immediately posterior to mandible base, bearing 4 naked setae. Maxilla (fig. 6@) appearing 2-segmented, first segment swollen, nodular, second segment arising from inner lateral surface of first, rodlike, bearing 2 naked setules and single spatulate NO. 3656 COPEPODS—LEWIS 13 Ficurre 6.—Pseudotaeniacanthus species, male, ventral views: a, right side of oral region, showing mandible, paragnath, maxillule, and maxilla; b, left maxilliped. ‘Thoracic legs: c, right first, posterior view (see table 3 for discussion of position of rami); d, right second, anterior view; e, right third, anterior view; f, left fourth, anterior view; g, left fifth, an- terior view. 14 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 process distally, spatulate process denticulated distally. Paragnath (fig. 6a) small, nodular, without distinctive armature. Ventral surface of cephalothorax with numerous heavily sclerotized knobs, adhesion processes and spikes. Postantennal process (fig. 5e) most prominent, extending as ridge from lateral anterior surface posteriorly to oral region, terminating as spinelike projection; ridge bearing 2 adhesion pads. Postantennal processes contiguous at base but not fused. Prominent pair of spinelike projections situated im- mediately lateral to antenna base, in position of postantennal process of many caligoids. TABLE 3.—Armature of thoracic legs I-V of the male of Pseudotaeniacanthus species Inter- | Protopodite Exopodite Endopodite Leg | Surface || podal cts ee eee ae Plate 1 | 2 1 2 3 1 | 2 | 3 I | Outer* m iP 3P c c ¢,2P Inner m,p GE 18 4P P ¢;P ¢,3P II | Outer p dm,H H 2H,Q’#,P c c c,h,2p’ Inner Ve fm c De 4P ey 2P 3 III | Outer p H H 2H QP c c c,h,p’,fm, p’, Inner 12) fm ¢ 2 4P iP c,2P ¢,2P IV | Outer p dm,H dm,H | dm,H,H,2P | fm fm fm,h,fm,P Inner c ? 4P iP iP DLE V | Outer dm,p’,dm, PD jam; H *Position of rami in figure 6c reversed due to flattening and consequent movement during mounting; endopodite situated almost behind (posterior) exopodite in appendage in situ. #Seta-like process with plumosities on one side, naked on other. ?Armature element may have been lost in handling. Maxilliped (fig. 66) 3-segmented, situated posterior and slightly medial to maxilla base. First segment irregular, base partially formed by heavily sclerotized ring; single naked seta present, on anterior inner surface. Second segment strongly developed, inner surface with large, heavily sclerotized, knob-tipped projection, projection with single row of denticles; inner distal surface with 2 contiguous oval adhesion pads. Third segment small, distinct from long, spinelike terminal process, bearing single, setalike accessory process. Thoracic legs I-IV biramous, rami 3-segmented; fifth leg uniramous, 2-segmented. For nature of legs and armature see figures 6c-g and table 3. eal NO. 3656 COPEPODS—LEWIS 5 Discussion.—The single male specimen from Eniwetok differs from the diagnosis of the genus given in Yamaguti (1963) in the following characteristics: 1. The lateral extensions of the second to fifth pedigerous segments are not as large. 2. The antenna is 4-segmented, not 3-segmented. The condition exhibited in the third and fourth segments of the antenna of the Eniwetok specimen suggests that the terminal segment of the antenna of previously described species may be composed of 2 fused, or par- tially fused segments. 3. The postantennal processes are contiguous at their base but are not fused and do not form the characteristic Y-shaped structure pres- ent on other members of the genus. Additionally, although there are 2 adhesion pads on each postantennal process, the projections on these pads do not have the characteristic brushlike appearance of those on other members of the genus. These two differences, however, suggest that the structure, in the group to which the Eniwetok speci- men belongs, is of a primitive nature. In spite of these differences, the specimen is placed in the genus Pseudotaeniacanthus not only because the characteristics, other than the above, coincide most closely with those previously described for other members of the genus but also because there is an overall simi- larity of the postantennal process of the Eniwetok specimen with that of the other members. Family PANDARIDAE Pandarus cranchii Leach Pandarus cranchit Leach, 1819, p. 535.—Lewis, 1966, p. 81, figs. 10-13.—Cressey, 1967, p. 9, figs. 27-33. For remaining synonymy, see Lewis (1966) and Cressey (1967). Hosts AND DISTRIBUTION.—See Lewis (1966). Mareriat.—4 females (USNM 120746) from the external surface of Galeocerdo cuviert (Lesueur). MEASUREMENTS (in mm).—4 females: mean range Length of body, excluding caudal setae 6. 87 6. 51-7. 14 Length of cephalothorax, including frontal region 4. 06 3. 85-4. 33 Width of cephalothorax 4. 26 3. 96-4. 48 Length of genital segment 2.61 2. 37-2. 74 Width of genital segment 2. 94 2. 74-3. 03 Length of caudal rami Its Be 1. 30-1. 70 Length of anal lamina 1. 63 1. 52-1. 74 Length of egg strings (6 strings) 4. 07 3. 55-5. 00 Descrirtion.—See Lewis (1966). 16 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 Family DissonrpAE Dissonus heronensis Kabata FIGuRES 7-9 Dissonus heronensis Kabata, 1966a, p. 221, fig. 5. Host AND DISTRIBUTION.—‘‘Unspecified fish of the family Balisti- dae” (Kabata, 1966a, p. 221), Heron Island, Queensland, Australia. MareriAL.—7 females and 1 male (USNM 120741) from the gill cavity of Balistoides viridescens (Bloch and Schneider). MEASUREMENTS (in mm).—5 females and 1 male: emale mean range male Total length, excluding caudal setae 1. 96 1. 82-2. 03 1. 51 Length of prosome Teas 1. 12-1. 19 0. 88 Width of prosome 0. 85 0. 79-0. 88 0. 67 Length of cephalothorax 0. 87 0. 83-0. 94 0. 61 Length of genital segment 0. 56 0. 49-0. 63 0. 52 Width of genital segment 0. 47 0. 43-0. 50 0. 31 Length of abdomen 0. 20 0. 11-0. 25 0.11 Length of egg strings (8 strings) 1. 43 1. 37-1. 48 Derscription.—Female cephalothorax (fig. 7b) consisting of cepha- lon and first 2 thoracic segments, strongly arched in transverse section. Frontal region (fig. 7g) membranous, narrow, forming slightly less than 10 percent of length of cephalothorax; with small, median notch in anterior margin. Lateral margin irregularly convex, with fine membrane posteriorly. Posterior lateral cephalothoracic region lobate, projecting to posterior half of free second pedigerous segment. Major dorsal cephalothoracic grooves consisting of pair of convex grooves extending anteriorly from junction of inner margin of pos- terior lateral cephalothoracic region and outer margin of free second pedigerous segment, terminating lateral and slightly posterior to pair of small, slightly projecting heavily sclerotized ridges. Ocular region indistinct, situated on median longitudinal axis of body, anterior to ridges. Female free second pedigerous segment slightly less than twice as wide as long, division between segment and cephalothorax distinct; lateral regions lobate, extending posteriorly to posterior region of third pedigerous segment; lobes with membrane along outer margin. Third pedigerous segment distinct from second, width slightly more than twice the length, segment without posterior extensions of lateral regions. Fourth pedigerous segment distinct from third, width slightly more than one and one-half times the length; broadest medially, tapered to anterior and posterior ends. Female genital segment (fig. 7d) generally ovoid although shape variable; indistinctly separable from fourth pedigerous segment. | | | | | NO. 3656 COPEPODS—LEWIS 17 Platelike formation of cuticle covering most of dorsal surface, ventral surface with 2 series of nodules, each nodule bearing bifid hairlike projections, anterior series consisting of transverse row of 4 nodules, second series usually forming longitudinal row of 3 nodules on each be 4 4 $$ O01 mm 9° b/+—++—+—_ + 005 mm. okey Ficure 7.—Dissonus heronensis Kabata, 1966, dorsal views: a, male; b, female. Ventral views: c, male genital segment; d, female genital segment; ¢, female first, second, and third pedigerous segments (P-I, II, II1); f, right caudal ramus; g, right antennule and adjacent frontal region. 283-932—68——2 18 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 side although nodules not always in row. Fifth leg (fig. 9e) situated on lateral ventral surface in posterior half of segment, consisting of palp bearing 4 plumose setules, Abdomen 1-segmented, distinctly separable from genital segment. Lateral margins, in adult, irregular, anterior surface broader than posterior. Posterior surface flat laterally, broadly bilobed medially, junction of lobes forming anal indentation. Caudal rami (fig. 7f) small, subrectangular, with 3 plumose terminal setae, 1 plumose setule on inner distal surface, and plumose seta and setule on proximal lateral surface. Male cephalothorax and free pedigerous segments similar to those of female (fig. 7a). Genital segment (fig. 7c) elongate, rounded an- tt tt a | O41 mm. 9 O1 mmc Figure 8.—Dissonus heronensis Kabata, 1966, ventral views: a, female oral region, showing antenna, mouth cone, mandible, maxillule, maxilla base (ma-—2), postantennal process and postoral process; b, male left antenna and postantennal process; ¢c, right maxilla; d, male left maxilliped; ¢, female left maxilliped. NO. 3656 COPEPODS—LEWIS 19 teriorly, tapered slightly posteriorly; process-bearing nodules present although longitudinal row extending posterolaterally, consisting of 4 nodules, not 3 as in female. Fifth leg (fig. 9/) situated on lateral ventral surface, consisting of 5 lightly plumose setules originating from slight swelling. Sixth leg (fig. 9g) lappet-shaped, situated on lateral posterior surface of segment, bearing 2 lightly plumose setules. Abdomen 1-segmented, short; narrow anteriorly, flared sharply posteriorly, posterior surface as in female. Caudal rami as in female. Female and male antennule (fig. 7g) 2-segmented, situated on lateral anterior ventral surface of cephalothorax and adjacent frontal region. Second segment approximately nine-tenths the length of first; first broader proximally than distally, lateral margins irregular; anterior ventral surface of first segment bearing approximately 28 lightly plumose setae and setules. Second segment rodlike, medial posterior surface with one naked setule, distal surface with 12 naked setules. Female antenna (fig. 8a) 3-segmented, situated posterior and medial to antennule base. First segment subrectangular, attached to heavily sclerotized padlike projection of cephalothorax. Second segment sub- rectangular, slightly longer than first segment. Third segment in- completely fused to clawlike terminal process, with setule-like acces- sory process proximally. Segments of male antenna (fig. 8b) similar to those of female although second with slight indication of adhesion surfaces (not shown in figure), terminal process of third segment with accessory claw proximally, segment with second accessory process at. indistinct junction with terminal process. Female and male mandible (fig. 8a) 4-parted, distalmost part flattened, inner margin with 10-12 denticulations. Female and male postantennal process (figs. 8a, 6) situated posterior and slightly lateral to antenna base, consisting of 3 minute nodules, each with one or more hairlike processes. Female and male postoral process consisting of long, spatulate spine, male spine with light, spinelike accessory process distally. Female and male maxillule (fig. 8a) situated immediately lateral to mouth cone base, consisting of lobate projection, forming base of postoral process, bearing nodule, with seta and 2 setules, on proximal outer surface. Projection with slight adhesion surface on distal half of outer surface. Female and male maxilla (fig. 8c) 2-seg- mented, situated lateral and slightly posterior to maxillule. Segment lengths about equal, first more strongly developed, second rodlike, with small membrane on distal inner surface and 2 saber-shaped terminal processes. Innermost terminal process approximately one and one-half times the length of outermost, both with fine membranes, distal portion of membranes forming denticulations on innermost terminal process. 20 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 Female maxilliped (fig. 8¢) 2-segmented, situated immediately posterior and medial to maxilla base. First segment irregular, strongly developed, proximal end fused with cephalothorax; large, shelflike projection present on proximal inner surface, receiving distal end of second segment terminal process when segment flexed. Second segment indistinctly separable from clawlike terminal process, with accessory rN J 2: ae
: Ci} a8 Yt LA 4, y Ficure 23.—Caligus confusus? Pillai, 1961, female, right thoracic legs, anterior views: 4, first; b, distal region of second segment of exopodite of first; ¢, second; d, third, ¢, exopodite of third; f, fourth. Left fifth leg: g, ventral view. 58 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 125 (fig. 22f) nodular, situated lateral and slightly posterior to mandible base, bearing single naked seta and 2 naked setules. Maxilla (fig. 22g) 2-segmented, situated lateral and slightly posterior to maxillule. First segment approximately four-fifths the length of second, tapered proximally and distally to articulation surface. Second segment rodlike with U-shaped membrane on distal half of inner surface; distal surface with 2 saber-shaped processes, innermost approximately one and one-half times the length of outermost, both with fine mem- branes along outer and inner margins. Postoral adhesion pads (fig. 22f) evident as V-shaped ridges of heavy sclerotization adjacent to median longitudinal axis of body, slightly posterior to maxilla bases. Maxilliped (fig. 22h) 2-segmented, situated posterior and slightly medial to maxilla base. First segment strongly developed, inner TABLE 9.—Armature of thoracic legs I-IV of the female of Caligus confusus? Pillat, 1961 Inter- Protopodite Exopodite Endopodite Leg | Surface |} podal Plate 1 2 1 2 3 1 2 3 SSS SS SS SS | SS a SSS SSS I | Outer Dp m, rh 3dH Inner D;r* C,C Bea ed II | Outer m,p | m,dmH]} dmH dmH, Z#,Q|C le ¢, 3P m Inner sp mM iDe jACe Ge c, 5P Pc, 2e wee IiI | Outer a,m,p H Cip)s Crepe Cy |e; 2 m Inner 1D; 2s,)-P.'s: Cie ce 3b iP |'4P m,s IV | Outer p fm,mH | fm, mH | 3fm,2mH, H *Roughened by minute projections giving brushlike appearance to element. #Spine with simple membrane on one side, frilled membrane on other. proximal half with heavily sclerotized transverse ridge; additional small, knoblike projection present on posterior surface (not shown on figure). Second segment and incompletely separable terminal process clawlike, segment with setule-like accessory process. Sternal furca (fig. 227) situated between and slightly posterior to maxilliped bases, tines angled outward, sinus between tines broad. Thoracic legs I-III biramous although endopodite of first leg reduced; fourth leg uniramous. For nature and armature of legs, see figures 22a-f and table 9. Discussion.—The Eniwetok specimens differ from Pillai’s figures and description of Caligus confusus (1961) in two respects: 1. Pillai figures a membrane-like structure around the major spine on the postantennal process, another around the major spine on the No. 3656 COPEPODS—LEWIS 59 postoral process, and one on the outer margin of each ramus of the sternal furca. The Eniwetok specimens do not have any of these membranes although the edge of these structures is thin and semi- transparent and unless closely examined, gives a membrane-like appearance. 2. Pillai figures a blunt secondary projection on the proximal inner portion of the postoral process that is not present on the Eniwetok specimens. The identification of the Eniwetok material as C. confusus Pillai is questioned because of the two differences mentioned above and the absence of an adequate first description of the species. Both Pillai (1961) and Kirtisinghe (1964) list Wilson’s (1937a,b) and Shiino’s (1959a) ‘“‘Calligus constrictus”’ as a synonym of C. confusus because of the very similar morphology of the figured specimens. There are some differences in the morphology that could be consid- ered to be of a minor nature (e.g., shape of the postoral process), but there is a difference in the size of the specimens that should be evalu- ated with sufficient material. Pillai’s female specimen(s) are 2.9 mm while Shiino’s are 3.75 mm and Wilson’s 5.0 mm in length. Even the female specimen(s) of Kirtisinghe (1964), from Ceylon, measure 4.5 mm in length, more than 1.5 times the length of Pillai’s specimen(s) from southern India. Because of the difference in size, as well as the other characteristics, it is felt that there is some question (as noted in the synonymy) about the inclusion of Wilson’s and Shiino’s material and perhaps even that of Kirtisinghe. Caligus pseudokalumai, new species FIGurRes 24, 25 MarTeErRIAL.—1 female (holotype, USNM 120765) from the external surface of Gymnosarea nuda Giinther. MEASUREMENTS (in mm).—1 female: Total length, excluding caudal setae 2. 89 Length of cephalothorax 2. 04 Width of cephalothorax 1. 48 Length of genital segment 0. 81 Width of genital segment 0. 89 Length of abdomen 0. 19 Length of egg strings (strings broken) Description.—Cephalothorax (fig. 24a) ovoid, consisting of ceph- alon, maxilliped-bearing and first 3 pedigerous segments. Frontal region approximately one-ninth the length of cephalothorax, anterior surface with narrow membrane. Lunules (fig. 24e) continuous with membrane along anterior margin, extending posteriorly, on ventral surface, to junction of frontal region and rest of cephalothorax. Lateral edge of 60 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 cephalothorax smooth, with slight indentation in anterior third, bearing narrow membrane and several hairlike processes. Posterior lateral region with small, cupshaped indentation. Posterior sinus (fig. 24c) distinct, anterior end sharply angled, lateral margins flatly concave, outer lateral surface bearing filmy membrane. Posterior median region of cephalothorax extending posteriorly well past lateral regions, pos- terior margin flatly convex in dorsal view; pair of minute, hairlike projections present, slightly anterior to posterior margin. Major dorsal cephalothoracic grooves forming irregular H, posterior legs flatly convex, anterior flatly concave, cross groove extending anteriorly from junction with legs of H. Ocular region distinct, in anterior fourth of cephalothorax. Free fourth pedigerous segment distinct from cephalothorax although anterior half overlapped by posterior median cephalothoracic region. Fourth pedigerous and genital segments fused dorsally and ventrally. Surfaces of genital segment (fig. 246) irregular; anterior region of segment broad, posterior bilobed, lobes extending posteriorly to caudal rami. Fifth legs (fig. 25g) situated on ventral surface, at beginning of posterior lobes, consisting of 2 nodules, anteriormost bearing single, lightly plumose setule, posterior bearing 2 plumose setules. Abdomen 1-segmented, distinct from genital segment, lateral mar- gins smooth, posterior margin V-shaped, almost covered by proximal surface of caudal rami. Caudal rami (fig. 24d) broad, lobate, width almost twice the greatest length. Inner surface plumose, distal surface bearing 3 plumose setae; inner distal surface with 1 plumose setule, distal outer surface with 2 plumose setules. Antennule (fig. 24¢) 2-segmented, attached to lateral posterior ventral surface of frontal region and adjacent surface of cephalothorax. First segment of general parallelogram shape, anterior and anterior ventral surface with 21 naked and lightly plumose setae and setules. Second segment clubshaped, length slightly less than two-thirds the length of anterior surface of first segment, with single naked seta from distal half of posterior surface, 13 naked setae from distal surface. Antenna (fig. 24f) 3-segmented, situated posterior and medial to antennule base. First segment squat, irregular, with narrow, dagger- like projection from posterior surface. Second segment slightly broader proximally than distally, distal surface irregular. Third segment and terminal process continuous, clawlike, with small, setule-like accessory process on proximal inner surface, second on medial surface. Ficure 24.—Caligus pseudokalumat, new species, female: a, dorsal view; c, posterior cephalo- thoracic sinus, same view. Ventral views: b, fourth pedigerous and genita] segments; d, left caudal ramus; ¢, left lunule and antennule; f, oral region, showing antenna, mouth cone, mandible, maxillule, maxilla base (ma-2), postantennal, and postoral processes; g, right maxilla; h, right maxilliped; 7, sternal furca. NO. 38656 COPEPODS—LEWIS 61 62 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 126 Mandible (fig. 24f) rodlike, 4-parted, distalmost part curved inwards, inner surface with 12 denticulations. Postantennal process (fig. 24/) situated posterior and lateral to antenna base, consisting of heavily sclerotized clawlike projection with 2 nodules proximally, third present just posterior to base of projection; nodules each bearing single, large, hairlike process. Postoral process (fig. 24f) dagger-like, immediately adjacent to maxilule. Maxillule (fig. 24f) situated lateral and slightly posterior to mouth cone base, consisting of node bearing 2 short and one long seta-like projections. Maxilla (fig. 24g) 2-sezmented, situated lateral and slightly posterior to maxillule. First segment slightly more than three-fourths the length of second, proximal articulation and muscle attachment surface directed at sharp angle to axis of segment. Second segment rodlike, bearing small, horsehoe-shaped membrane from medial inner surface and 2 saber-shaped terminal processes. Inner terminal process approximately twice the length of outer, both with fine membrane along inner and outer surfaces. Maxilliped (fig. 24h) small, 2-segmented, situated posterior and slightly medial to maxilla base. First segment narrow, with small, spikelike projection from proximal posterior surface. Second segment rodlike, distinct from short, clawlike terminal process, with setule-like accessory process on distal inner surface. Sternal furca (fig. 242) situated on median longitudinal axis of body posterior to maxilliped bases. Basal region of furca subconical, tines flattened, chisel-shaped distally. Thoracic legs I-III biramous although endopodite of first leg rudimentary. Fourth thoracic leg uniramous, 3-segmented. For na- ture and armature of legs, see figures 25a—f and table 10. Discusston.—Caligus pseudokalumai most closely resembles C. kalumai Lewis (1964a) recorded from Acanthurus guttatus from the Hawaiian Islands. The relatively minor differences between the two species include the following: 1. The cephalothorax is more elongate in C. pseudokalumai, and the posterior median cephalothoracic region projects posteriorly past the posterior lateral cephalothoracic regions more than twice as far as it does in C. kalumai (0.36 mm to 0.16 mm). As a percent of the length of the cephalothorax, this equals 12 and 9 respectively. 2. The postantennal and postoral processes as well as the sternal furca are bulkier and more heavily sclerotized in C. kalumai. Even with this, however, the 3 nodules associated with the postantennal process each bears a single large (in comparison with other species of the genus), hairlike process (not 2 as stated in Lewis, 1964a, p. 175) as do those in C. pseudokalumai. 3. The first segment of the endopodite of the second thoracic leg bears a tuft of plumosities in C. kalumai but does not in C. pseudo- NO. 3656 COPEPODS—LEWIS 63 Ficure 25.—Caligus pseudokalumai, new species, female, right thoracic legs, anterior views: a, first; b, distal region of second segment of exopodite of first; c, second; d, third; é, exopodite of third; f, fourth. Left fifth leg; g, ventral view. 64 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 126 kalumai. The plumosities on the outer margin of the second and third segments of the endopodite are heavier in C. pseudokalumai than in C. kalumar. 4. The first segment of the exopodite of the fourth leg bears a filamentous (or slightly denticulated) membrane at the base of the spine in C. pseudokalumai which is missing in C. kalumai. With one possible exception, these do not appear to be gross differ- ences and the apparent close relationship between the two species should be more closely examined when larger collections are available. The single difference that may be of a relatively gross nature is the tuft of plumosities (or their absence) on the first segment of the endopodite of the second thoracic leg. This is the easiest characteristic to use in separating the two species and, because of the relatively stable nature of the second thoracic leg, may indicate more divergence than is presently believed. TaBLE 10.—Armature of thoracic legs I-IV of the female of Caligus pseudokalumai, new species Inter- Protopodite Exopodite Endopodite eg:|| Surface: ||| podal i. 2 = s late 2 1 2 3 1 2 3 I | Outer sss, D th SH! Inner p c 3P II | Outer m,p m,mH | mH rh, mH, Q Cc C, 2P m Inner Spe m, s CG, (Ode 5P Pic, 2P. icv4e III | Outer* m,p s,m C.S; Dp: ce, 3p’, D c | ¢,3P m Inner 3S;-2,1n 8 lel iP c, 3P Perse IV | Outer 3s, p fm, mH | 2H * Armature tabulation does not include several hairlike processes in middle of protopodite. The species name indicates the similarity between the new species from Eniwetok and Caligus kalumai from Hawaii. Caligus alaihi, new species FIGURES 26-28 Mareriau.—1 female (holotype, USNM 120766) and 1 male (allotype, USNM 120767) from the buccal cavity of Holocentrus sammara (Forskal). NO. 3656 COPEPODS—LEWIS 65 MEASUREMENTS (in mm).—1 female and 1 male: female male Total length, excluding caudal seta 2.77 2.07 Length of cephalothorax 1.55 e271 Width of cephalothorax 1.53 i altZ Length of genital segment 0.65 0.40 Width of genital segment 0.68 0.36 Length of abdomen 0.34 0.32 Description.—Female cephalothorax (fig. 26a) ovoid, consisting of cephalon, maxilliped bearing and first 3 pedigerous segments. Frontal region approximately one-seventh the total length of cephalothorax, with fine membrane along anterior edge. Lunules (fig. 26g) distinct, extending posteriorly, on ventral surface, past junction of frontal region and remaining cephalothorax. Lateral margin of cephalothorax flatly convex although slightly irregular, with slight indentation anteriorly. Lateral ventral surface bearing 2 narrow membranes, first extending laterally, second medially, on ventral surface. Small, cup- shaped depression present in dorsal posterior lateral surface, with single, hairlike process arising from middle of cup. Posterior sinus irregularly U-shaped, with membrane along outer lateral surface. Median posterior surface extending posteriorly slightly past lateral posterior surfaces, overlapping anterior end of free fourth pedigerous segment. Major dorsal cephalothoracic grooves forming irregular H, anterior legs terminating well posterior to ocular region. Ocular region distinct, in anterior third of cephalothorax. Female free fourth pedigerous segment short, length slightly more than half the width; segment widest medially, in region of fourth leg attachment. Segment distinct from genital segment ventrally, fused dorsally. Genital segment (fig. 26c) of general apple shape from dorsal viewpoint, anterior end narrowest. Fifth legs (fig. 28h) arising from posterior lateral surface, consisting of 2 setule-bearing nodules, anteriormost with single, lightly plumose setule, posterior with 3 lightly plumose setules. Female abdomen 1-segmented, distinct from genital segment, lateral margins essentially parallel except at constricted anterior end; posterior surface angular, forming attachment surface for caudal rami, median posterior surface bilobed. Caudal rami (fig. 26f) con- stricted proximally, otherwise subrectangular, distal surface with 3 plumose setae and 1 plumose setule, on inner edge, distal outer surface with 2 plumose setules; distal half of inner surface plumose. Male cephalothorax (fig. 26) and free fourth pedigerous segment similar to that of female. Genital segment (fig. 26d) of general barrel 283-932—68——_5 66 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 shape, fifth legs (fig. 28g) situated on posterior lateral surface, con- sisting of 2 setule-bearing nodules, anteriormost with single plumose setule, posterior with 2 plumose setules; sixth legs (fig. 289) situated on posterior surface, adjacent to junction of genital segment and abdomen, consisting of nodule bearing 3 plumose setules. Abdomen Ficure 26.—Caligus alaihi, new species, dorsal views: a, female; 6, male; ¢, posterior cephalothoracic sinus. Ventral views: c, female fourth pedigerous and genital segments; d, male fourth pedigerous, genital, and first abdominal segments; f, right caudal ramus; g, left lunule and antennule. NO. 3656 COPEPODS—-LEWIS 67 2-segmented, distinct from genital segment dorsally, indistinctly separable ventrally. First segment short, slightly more than half the length of second, lateral margins flatly convex; second segment constricted anteriorly, margins parallel posterior to constriction, posterior surface angular, as in female. Caudal rami as in female. Female and male antennule (fig. 26g) 2-segmented, attached to both lateral ventral surface of frontal region and lateral anterior ventral surface of cephalothorax. First segment broad proximally, tapered to narrow distal end, anterior and ventral anterior surface with ap- proximately 20 naked and lightly plumose setae and setules. Second pp ed . Hee rt 02 mm, & 0.2 mm. 9 0.2 mm. oe Ficure 27.—Caligus alaihi, new species, ventral views: a, female oral region, showing antenna, mouth cone, mandible, maxillule, maxilla base (ma-—2), postantennal, and postoral processes; b, male right antenna and postantennal process; c, right maxilla; d, male right maxilliped; e, female right maxilliped; f, sternal furca. 68 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 segment rodlike, approximately four-fifths the length of first segment, posterior surface with single naked setule distally, distal surface with 12 naked setae and setules. Female antenna (fig. 27a) 3-segmented, situated posterior and medial to antennule base. First segment short, squat, outline subtriangular. Second segment subrectangular; third segment and incompletely fused terminal process clawlike, with sub- conical accessory process proximally, setule-like, accessory process medially, at indistinct division between segment and terminal process. Male antenna (fig. 27b) 3-seemented, situated posterior to antennule base. First segment elongate, fused proximally with cephalothorax, angled ventrally toward distal end. Second segment well developed, broader proximally than distally, with adhesion pad on distal half of inner surface. Third segment fused with short, bifurcate terminal process, tines of terminal process pointed; setule-like accessory process present, arising just proximal to terminal process. Female and male mandible (fig. 27a) rodlike, 4-parted, distalmost part curved inward, flattened, inner surface with 12 denticulations. Female postantennal process (fig. 27a) situated lateral and slightly posterior to antenna base, consisting of short, clawlike projection and 3 nodules, first 2 on base of projection, third shghtly posterior to base, all 3 with several hairlike processes. Male postantennal proc- ess (fig. 276) as in female except clawlike projection much longer. Female postoral process (fig. 27a) spinelike, situated immediately posterior to maxillule, spine projecting from middle of base, distal end rounded. Male postoral process as in female except spine project- ing from outer portion of base. Female and male maxillule (fig. 27a) nodular, situated lateral and slightly posterior to mandible base, nodule bearing 2 short and 1 long, seta-like processes. Female and male maxilla (fig. 27c) 2-segmented, located posterior and lateral to postoral process; both segments slender although first approximately two and one-half times the diameter of second, length slightly more than four-fifths that of second. Second segment rodlike, with small flapshaped membrane on distal third of outer surface, bearing 2 saber- like terminal processes, innermost longer than outer, both with fine membrane along outer and inner margins. Female maxilliped (fig. 27e) 2-segmented, situated posterior and slightly medial to maxilla base. First segment strongly developed, with slightly recurved proximal region terminating in knoblike proxi- mal end; inner medial surface with small, shelflike projection (not shown on figure) receiving distal end of terminal process of second segment when segment flexed. Second segment distinct from clawlike terminal process, bearing setule-like accessory process on distal inner surface. Male maxilliped (fig. 27d) similar to that of female except first segment more strongly developed and recurved proximal end | NO. 3656 COPEPODS—-LEWIS 69 Ficure 28.—Caligus alaihi, new species, right thoracic legs, anterior views: a, first; b, distal region of second segment of exopodite of first; c, second; d, third; ¢, exopodite of third (posterior view); f, fourth. Legs, ventral views: g, male left fifth and sixth; h, female right fifth. 70 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 reduced. Female and male sternal furca (fig. 27/) well developed, Y-shaped, distal ends of rami rounded. Thoracic legs I-III biramous although endopodite of first leg rudimentary; fourth thoracic lez unirmaous, 3-segmented. For nature and armature of thoracic legs, see figures 28a—f and table 11. Discusston.—The diagnostic characteristic of Caligus alaihi is the combination of a 2-segmented abdomen on the male, denticulations on the second segment of the endopodite of the second thoracic leg (but not on the first segment), and the presence of only 1 plumose seta on the inner surface of the second segment of the exopodite of the first thoracic leg. Of these three, the most uncommonly found feature in other members of the genus is the presence of a single seta on the inner surface of the second segment of the exopodite of TABLE 11.—Armature of thoracic legs I-IV of the female and male of Caligus alaihi, new species Inter- Protopodite Exopodite Endopodite Leg | Surface podal SS oe SECON Raa 5 Plate 1 2 1 2 3 1 2 3 I | Outer 8s, p rh 3H e Inner p c p II | Outer m,p m,mH | mH rh, mh,Q | ¢ D 3P m Inner Sie m,s Che cr B 5P 12 c,; 2P 3P ee NA Pe ls || are ee | ee ee | ed Ee eee | ee a III | Outer r,m,p ¢c, p’ Cy3p’, e c c, 3P m Inner 3s,P,m,s H CweE 3P 1? 3P IV | Outer 2s, p fm, mH | 4fm, 3mH, : | H the first thoracic leg. This feature, however, is found in at least one other species, Caligus pagrosomi Yamaguti, 1939. Caligus alaihi can be most readily distinguished from C. pagrosomi by the presence of denticulations on the second segment of the endopodite of the second thoracic leg, in contrast to the plumosities presumed to be found on the same segment of C. pagrosomi. (Yamaguti, 1939, does not provide a description of the second thoracic leg of C. pagrosomi but does indi- cate, p. 445, that the species is very similar to C. epinepheli Yamaguti, 1936, which does possess plumosities on the second segment of the endopodite of the second thoracic leg.) Caligus alaihi can also be distinguished from C. pagrosomi by the apparent length of the abdo- men of the female, that of C. pagrosomi being approximately twice the length of that of C. alaiki. This characteristic, however, should be used with some reservation as the variability (or potential variation) NO. 3656 COPEPODS—LEWIS v1 is not discernable from the single female in the collection of copepods from Eniwetok fishes. The species name is derived from the Hawaiian name, “alaihi,’’ for several of the holocentrid fishes, the family to which the host of this copepod belongs. Caligus laticaudus Shiino Figures 29, 30 Caligus laticaudus Shiino, 1960, p. 482, figs. 5, 6.—Pillai, 1961, p. 91, fig. 3— Yamaguti, 1963, p. 55, pl. 68, fig. 4. Hosts AND DISTRIBUTION.—3 host records: locality host reference Japan Pagrosomus major Shiino, 1960 South India Polynemus heptadactylus Rhabdosargus sarba Pillai, 1961 Mareriau.—1 female (USNM 120769) from the external surface of Acanthurus olivaceous Bloch and Schneider. MEASUREMENTS (in mm).—1 female: Total length, excluding caudal setae Length of cephalothorax Width of cephalothorax Length of genital segment Width of genital segment Length of abdomen Length of egg strings (not present, condition suggests specimen an immature adult) QS et RATAN BNO Description (female only).—Cephalothorax (fig. 29a) ovoid, con- sisting of cephalon, maxilliped-bearing and first 3 pedigerous segments. Frontal region approximately one-ninth the length of cephalothorax, with narrow membrane along anterior margin. Lunules (fig. 29e) distinct, extending posteriorly, on ventral surface, to posterior por- tion of frontal region, of relatively simple construction in comparison with those of Caligus coryphaenae. Division between frontal region and rest of cephalothorax distinct. Lateral margin of cephalothorax flatly convex, bearing single, narrow membrane extending laterally; posterior lateral surface lappet-shaped, with small, cupshaped in- dentation bearing 2 clusters of hairlike processes. Posterior sinus (fig. 29c) U-shaped, with membrane along outer margin. Median posterior surface of cephalothorax extending slightly past lateral posterior surfaces, posterior margin irregular on single specimen in collection. Major dorsal cephalothoracic grooves forming irregular H, anterior legs of H terminating well posterior to ocular region. Ocular region distinct, in anterior third of cephalothorax. Ficure 29.—Caligus laticaudus Shiino, 1960, female: a, dorsal view; d, right caudal ramus, same view. Ventral views: }, posterior region of genital segment and adjacent surface of abdomen, showing fifth leg and attached spermatophore; c, posterior cephalothoracic sinus; ¢, left lunule and antennule; f, oral region, showing antenna, mouth cone, mandible, maxillule, maxilla base (ma—2), postantennal, and postoral processes; g, right maxilla; A, left maxilliped; 7, sternal furca. No. 8656 COPEPODS—-LEWIS 73 Free fourth pedigerous segment distinct from cephalothorax, indistinctly separable from genital segment; segment widest medially, at junction with fourth thoracic legs, tapered anteriorly and poster- iorly. Genital segment (fig. 29b) subovoid, posterior region irregular in single available specimen. Fifth legs (fig. 30g) situated on posterior lateral surface of segment, consisting of node bearing 2 or 3 plumose setules (2 present on specimen, indication of additional broken setule) and nodule bearing single plumose setule (= sixth leg of Shiino, 1960, p. 485). Abdomen indistinctly 2-segmented, distinct from genital segment ventrally, indistinctly separable dorsally. First segment with flatly convex lateral surfaces, second tapered slightly to posterior surface. Division between segments consisting of break in cuticle of lateral regions only, muscle-segment associations not distinct enough to indicate segmentation. Caudal rami subpalmate, bearing 3 plumose setae and 1 plumose setule distally, plumose seta and setule present on distal half of outer lateral surface; distal half of inner surface plumose. Antennule (fig. 29¢) 2-segmented, attached to anterior lateral ventral surface of cephalothorax and adjacent surface of frontal region. First segment approximately twice the length of second, broad proximally, tapered distally; anterior surface sharply angled distally. First segment bearing approximately 22 lightly plumose and naked setae and setules. Second segment rodlike, bearing single naked setule from distal half of posterior surface, 11 naked setae and setules (including 1 bifurcate seta) distally. Antenna (fig. 29f) 3-segmented, situated posterior and slightly medial to antennule base. First segment squat, subtriangular from ventral viewpoint; second segment well developed, broad proximally, tapered slightly to distal end. Third segment and terminal process fused, with falciform outline; single, setule-like accessory process present on node at proximal end of fused segment and terminal process. Mandible (fig. 29f) rodlike, 4-parted, distalmost part curved inward, flattened, inner surface with 12 denticulations. Postantennal process (fig. 29f) situated lateral and slightly posterior to antenna base, con- sisting of spikelike projection and 3 nodules bearing several hairlike processes. Projection arising from small, subtriangular, heavily sclerotized plate, plate also bearing 2 of 3 nodules, third nodule immedi- ately posterior to plate. Postoral process (fig 29f) spinelike, curving out- ward slightly. Maxillule (fig. 297) nodular, situated immediately anteri- or to postoral process, nodule bearing 1 long, 2 short, naked seta-like processes. Maxilla (fig. 29g) 2-segmented, situated immediately posterior and slightly lateral to postoral process. First segment ap- proximately nine-tenths the length of second, second elongate, with 74 PROCEEDINGS OF THE NATIONAL MUSEUM VOI. 125 ET, oS, = 4% 7 “a Ld LIT, ‘rs: ZZ E75 KH Se Lil = x —— ar S yyw Ficure 30.—Caligus laticaudus Shiino, 1960, female, right thoracic legs: a, first, anterior view; b, distal region of second segment of exopodite of first, anterior view; c, second, posterior view; d, third, anterior view; e, exopodite of third, posterior view; f, fourth, posterior view. Right fifth leg: g, ventral view. NO. 3656 COPEPODS—LEWIS 75 fine membrane on distal half of outer surface, distal surface bearing 2 saber-shaped processes, innermost approximately one and one-fourth times the length of outermost, both fringed with fine membrane. Mazxilliped (fig. 20h) 2-segmented, situated posterior and slightly medial to maxilla base. First segment strongly developed, irregular in outline; widest medially, proximal end narrow, distal approximately half the greatest width; segment with socket-like shelf on medial inner surface, receiving distal end of second segment terminal process when segment flexed. Second segment distinct from clawlike terminal process, bearing setule-like accessory process on distal inner surface. Sternal furea (fig. 292) situated between and slightly posterior to maxilliped bases, bifurcations flattened anteroposteriorly, lobate in outline. TABLE 12.—Armature of thoracic legs I-IV of the female of Caligus laticaudus Shiino, 1960 | Inter- Protopodite Exopodite Endopodite Leg | Surface podal Plate 1 2 1 2 3 1 2 3 I | Outer ss, p rh h, 2H, p’ Inner p c 3P II | Outer m, p m,dmH| H Jays 1SIGKA) c (ed i m Inner GIP m, 5, Cee Oy 5P Pp Cree alte or III | Outer mee m, s Cape Cc, 3p’ c c, 3P m Inner P,s,m,s H P 4P P 3P peas eres ens | | ee Lene meee, ——— | IV | Outer p fm, mH | fm, mH | fm, 3m | Thoracic legs I-III biramous although endopodite strongly reduced in first leg; fourth thoracic leg uniramous, 4-segmented. For details of legs and armature, see figures 30a—/ and table 12. Discusssion.—The Eniwetok specimen differs from Shiino’s original description in several points: 1. There is a setule on the inner surface of the protopodite of the first thoracic leg that is not listed or figured in Shiino’s publication. 2. The sternal furca appears broader than that shown by Shiino for C. laticaudus although the angle from which the drawing was made may at least partially explain this. 3. The total length of the Eniwetok specimen is smaller than that listed by Shiino (3.51 mm) although Pillai (1961) lists 2.6 mm as the length of his female specimen(s?). The difference in length between the Eniwetok specimen and Shiino’s is presumed to be, at least in part, due to the possible immature adult condition of the Eniwetok specimen, Literature Cited CressgEy, R. F. 1967. Revision of the family Pandaridae (Copepoda: Caligoida). Proc. U.S. Nat. Mus., vol. 121, no. 3570, pp. 1-138, figs. 1-356. Dana, J. D. 1853; 1855. Crustacea. Pt. 2 of vol. 13 in United States Exploring Expedi- tion during the years 1838, 1839, 1840, 1841, 1842 under the command of Charles Wilkes, U.S.N., 1618 pp. (1853); atlas, 27 pp., 96 pls. (1855). HeEEGARD, P. 1940. 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Copepods parasitic on Japanese fishes, 1: On the species of Caligus and Lepeophtheirus. Rep. Fac. Fish. Pref. Univ. Mie, vol. 1, no. 2, pp. 79-113, figs. 1-14. 1954a. Note on a new parasitic copepod, Caligus brevis n. sp. Bull. Japanese Soe. Sci. Fish., vol. 20, no. 3, pp. 178-183, figs. 1-2. 1954b. Copepods parasitic on Japanese fishes, 3: On two new species of the genus Anuretes. Rep. Fac. Fish. Pref. Univ. Mie, vol. 1, no. 3, pp. 260-272, figs. 1-5. 1957. Copepods parasitic on Japanese fishes, 16: Bomolochidae and Taenia- canthidae. Rep. Fac. Fish. Pref. Univ. Mie, vol. 2, no. 3, pp. 411- 428, figs. 1-6. 1959a. Ostpazifische parasitierende Copepoden. Rep. Fac. Fish. Pref. Univ. Mie, vol. 3, no. 2, pp. 267-333, figs. 1-25. 78 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 125 Suto, 8. M.—Continued 1959b. Sammlung der parasitischen Copepoden in der Prafekturuniversitat von Mie. Rep. Fac. Fish. Pref. Univ. Mie, vol. 3, no. 2, pp. 334— 374, figs. 1-17. 1960. Copepods parasitic on the fishes collected on the coast of Province Shima, Japan. Rep. Fac. Fish. Pref. Univ. Mie, vol. 3, no. 3, pp. 471-500, figs. 1-10. 1964. Results of Amami Expedition, 6: Parasitic Copepoda. Rep. Face. Fish. Pref. Univ. Mie, vol. 5, no. 1, pp. 243-255, figs. 1-4. 1965a. Parasitic copepods of eastern Pacific fishes. Rep. Fac. Fish. Pref. Univ. Mie, vol. 5, no. 2, pp. 391-420, figs. 1-9. 1965b. Parasitic copepods of the eastern Pacific fishes, 8: Lepeophthetrus. Rep. Fac. Fish. Pref. Univ. Mie, vol. 5, no. 2, pp. 441-454, figs. 1-4. STEENSTRUP, J. J. S., and Liirxen, C. F. 1861. Bidrag til Kundskab om det aabne Havs Snyltekrebs og Lernaeer samt om nogle andre nye eller hidtil kun ufuldstaendigt kjendte parasitiske copepoder. Kongel. Danske Vidensk. Selsk. Skr. Naturh. Math. Afdel., 5R., vol. 5, pp. 341-432, pls. 1-15. THomson, G. M. 1889. Parasitic Copepoda of New Zealand, with descriptions of new species. Trans. Proc. New Zealand Inst., vol. 22, pp. 353-376, pls. 25-29. VERVOORT, W. 1962. A review of the genera and species of the Bomolochidae (Crustacea, Copepoda), including the description of some old and new species. Zool. Verh., no. 56, pp. 1-111, figs. 1-12. Witson, C. B. 1905a. North American parasitic copepods belonging to the family Caligidae, 1: The Caliginae. Proc. U.S. Nat. Mus., vol. 28, no. 1404, pp. 479-672, pls. 5-28, figs. 1-50. 1905b. New species of parasitic copepods from the Massachusetts coast. Proc. Biol. Soc. Washington, vol. 18, pp. 127-132. 1906. Report to the government of Ceylon on the Pearl Oyster Fisheries of the Gulf of Manaar, Supplementary report 34: On some para- sitic copepods. Jn Ceylon Pearl Oyster Rep., pt. 5, pp. 189-210, pls. 1-5. 1937a. Parasitic copepods taken during the third Hancock Expedition to the Galapagos Islands. Jn Hancock Pacific Expeditions, vol. 2, no. 4, pp. 23-30, 1 pl. 1937b. Some parasitic copepods from Panama Bay. Journ. Washington Acad. Sci., vol. 27, no. 10, pp. 423-431, figs. 1-34. YAMAGUTI, S. 1936. Caligoida, I. Pt. 2 in Parasitic copepods from fishes of Japan, 22 pp., 12 pls. [Publ. by author.] 1939. Caligoida, III. Pt. 5 in Parasitic copepods from fishes of Japan, pp. 443-487, pls. 14-33. [Vol. Jub. pro Prof. Sadao Yoshida, vol. 2.] 1954. Parasitic copepods from fishes of Celebes and Borneo. Publ. Seto Mar. Biol. Lab., vol. 3, no. 3, pp. 375-398, 6 pls. 1968. Parasitic Copepoda and Branchiura of fishes, vii + 1104 pp., 333 pls. YAMAGUTI, S., and Yamasu, T. 1959. Parasitic copepods from fishes of Japan with descriptions of 26 new species and remarks on two known species. Biol. Journ. Okayama Univ., vol. 5, nos. 3, 4, pp..89-165, pls. 1-14. U.S. GOVERNMENT PRINTING OFFICE: 1968 - —— = 7 _ 7 | EBroceedings, of the United States National Museum SMITHSONIAN INSTITUTION + WASHINGTON, D.C. Volume 125 1968 Number 3657 North and Central American Species of Aneurus Curtis (Hemiptera: Aradidae) By Nicholas A. Kormilev! Fifteen species of the genus Aneurus Curtis, 1825, have been re- corded for North and Central America. In the present paper five additional species are described and a key to the 20 species is presented. This study is based on specimens from the U.S. National Museum, Washington, D.C., and the American Museum of Natural History, New York, N.Y., loaned by Drs. Richard C. Froeschner and Peter Wygodzinsky, respectively; and information from types in the British Museum (Natural History) furnished by Dr. William J. Knight. To all these men I offer my sincere thanks. Measurements in this paper were taken with a micromillimeter eyepiece; 25 units equal 1 mm. For convenience, the length of the abdomen was taken from the tip of scutellum to the tip of hypo- pygium in the male and to segment IX in the female. There has been some confusion concerning the identity of the Florida species, Aneurus politus Say (1832, p. 31). The original description made no reference to the important shape of the hypo- pygium. On the basis of five specimens from Guatemala, Champion (1898, p. 115) gave a short redescription with two drawings (table 7; figs. 29, 29a) and remarked about a “small and transverse’ hypo- pygium. Comparison of three Florida specimens with three Central 1565 Lincoln Place, Brooklyn, New York 11238. 2, PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 American (Mexico and Guatemala) specimens found them to be superficially similar but specifically distinct. In addition to smaller differences, the shapes of the scutellum and hypopygium are quite different (cf. figs. 14 with 17, and 15 with 18); the proportions of the antennal segments do not coincide; and the production of the para- tergites of the females differ (cf. figs. 16 and 19). Thus, it is evident that Champion’s specimens probably were not politus but an un- named species that will be described below as championi. To establish this usage and avoid further confusion, a neotype for politus will be proposed and described under the specific treatment of that taxon. Key to North and Central American Species of Aneurus Clavus fully developed, reaching tip of scutellum; latter triangular with pointed BDO Ke tt obo ahd ais este yes tae wet Subgenus Aneurosoma Champion, 1898 (eantaine a single species, dissimilis (Bergroth), 1889) Clavus greatly reduced to small triangle near basolateral angle of scutellum; apex of scutellum arcuate . .... . . . . Subgenus Aneurus Curtis, 1832 SUBGENUS Aneurus 1. Spiracles II to VIT lateral, visible from above, VIII terminal. vauriei Kormilev, 1964, Jamaica, Guadeloupe At least spiracles III ventral or sublateral, not visible from above . .. 2 Spiracle III sublateral, placed close to border, though not visible from above. nasutus Kormilev, 1966, Guadeloupe Spiracle III ventral, placed far from border . . . . ir et ee 3. Spiracles IV ventral, placed near border, not visible fom abowe! 1. pusillus, new species, Guatemala, Panama Spiracles IV ventral, placed far from or close to border; when near border, iS) spiracle V sublateral but not visiblefrom above .......... 4 4. Spiracle V lateral, visible from above . ise ion Cheek Eo ee FO Spiracle V ventral or sublateral, not visible from Shore a bo pels Ree EL 5. Scutellum longer, ratio between length and width at least 2:3 6 Scutellum very short, ratio between length and width 1:2. simplex Uhler, 1871, northeastern U.S. 6. Smaller species, length less than 4.0 mm; anterior process of the head not reaching tip of antennal segment I; antennal segment II obovate. 2. aibonitensis, new species, Puerto Rico Larger species, length about 5.0 mm; anterior process of head produced beyond tip of antennal segment I; antennal segment II clavate. septentrionalis Walker, 1873, eastern Canada 7. Spiracle V sublateral, not visible from above. 3. pisoniae, new species, Puerto Rico Spiracle V ventral, placed far from border. . .......454. +545 8 8. Spiracle VI lateral, visiblefrom above ....... . - «..+ +» me. Spiracle VI ventral, not visible from above .. . ne Rone aE 9. Head longer than width across eyes (35:33); Hntcnor nrouees of the head slightly produced beyond tip of antennal segment I. barberi Kormilev 1960, St. Lucia (Lesser Antilles) Head as long or shorter than width across eyes; anterior process at most reaching as far as tip of antennal segmentI ........... #10 NO. 3657 ANEURUS—KORMILEV 3 10. Head shorter than width across eyes (11:12). pygmeus Kormilev, 1966, southern U.S. Head as long as width across eyes . . . NON Poe eG 11. Antennal segment II clavate, as long as if larger peeciest leneth over 4.0 mm ela tan cel nin 7 penne song II iouowace: iuoner then hie emailer ippeeies) length less than4.0mm .. . stl dhsa Geeels 12. Anterior process of the head reaching BS of pntennal segment I; postocular tubercle (composed of 3 minute teeth) very short, not seats to outer border of eyes . . . . . . . . haitiensis Kormilev 1968, Haiti Anterior process not peachine to tip of antennal segment I; postocular tubercles strong, acute, reaching outer border of eyes. 6. championi, new species, Guatemala, Mexico 13. Scutellum wider, ratio between length and width 11.5:19; lateral borders of abdomen finely serrate. . . . . . 4. froeschneri, new species, Panama Secutellum narrower, ratio between length and width 12:16; lateral borders of abdomen roughly serrate. minutus Bergroth, 1886, southern U. S., Central America 14. Antennal segment II obovate, III and IV cylindrical . ....... 15 Antennal segment II clavate, III tapering toward base,.1V fusiform . 17 15. Scutellum subtriangular, rounded apically; ratio between length and width 10.5:19; hypopygium small, transverse (3.5:4). tenuis Champion, 1898, Panama, Cuba? Scutellum semicircular; ratio of length to width 2:3 or 3:4... ... 16 16. Larger species, length over 5.0 mm; hypopygium long and wide (10.5:10). montanus Champion, 1898, Guatemala Smaller species, length less than 4.5 mm; hypopygium smaller (8:7). fiskei Heidemann, 1904, U.S., Cuba? 17. Scutellum short, ratio of length to width 10:16. ..... scart Snedodireed Keo) Scutellum longer, ratio of length to width 15:19 or 20:24. 7. inconstans Uhler, 1871, U.S. 18. Smaller species, length less than 4.0 mm; antennae shorter, less than twice as long as head is wide (24:12.5). 5. politus Say, 1832, southern U.S. (Florida) Larger species, length over 5.0 mm; antennae longer, more than twice as long as head is wide (38:16). leptocerus Hussey, 1957, Guatemala, Panama 1. Aneurus pusillus, new species FIGurEs 4-6 Maue.—Elongate, sides subparallel; minutely granulate. Head slightly shorter than width across eyes (co 10:11.5, 9 10.5:12). Anterior process conical, not reaching to tip of antennal segment I; jugae slightly visible from above. Antenniferous tubercles small, acute, convex exteriorly. Eyes large, semiglobose, protruding. Post- ocular tubercles conical with blunt tip, produced as far as, or a little less than, outer borders of eyes. Vertex very finely, transversely striate. Vertex mesad of eyes with two (1-+1) large, ovate callosities. Antennae thin, one and one-half times as long as length of the head 4 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 (o 14:10, 9 15:10.5); segment I barrel shaped; II obovate, much smaller and narrower; III and IV cylindrical. Relative lengths of antennal segments I to IV: &@ 3:3:2.5:5.5, 93.5:2.75:2.75:6. Labium very short, reaching middle of head. Pronotum less than half as long as maximum width (o 9:19.5, 2 8.5:20). Collar tiny, sinuate in front. Anterolateral angles evenly rounded, produced neither forward, nor sideways. Lateral borders of forelobe parallel; humeri parallel, strongly converging anteriorly. Hind border sinuate in the middle. Foredisc with two (1+1) bean- shaped callosities, and a few small callosities laterad. Hind disc extremely finely punctured, with an arcuate, slightly raised, trans- verse, longitudinally striate strip. Scutellum almost as long as basal width (co 10:11, ? 10:11), evenly arcuate apically; disc finely, concentrically striate. Hemelytra reaching to the middle of tergum VII in both sexes; corium extended along basal third of scutellum. Abdomen elongate ovate, longer than maximum width across seg- ment IV (@ 40.5:25, 9 40.5:25). Connexivum narrow; PE-angles not protruding; PE-VII rounded in the male, angularly rounded in the female. Paratergites (7) moderately large, subtruncate apically, reaching apical fourth of hypopygium; latter moderately large, as long as maximum width (5:5). Paratergites (9) triangular, very short, produced almost as far as segment IX; latter truncate posteriorly. Spiracles IT lateral, III ventral, IV ventral placed nearer to border, V to VII lateral, VIII terminal. Cotor.— Yellow brown or pale testaceous, partially testaceous. MEASUREMENTs.—Total length: @ 2.85, 9 2.85 mm; width of pro- notum: o 0.78, 2 0.80 mm; width of abdomen: o 1.00, 2 1.00 mm. Honotypr.—o’, Guatemala, Cacao, Trece Aguas, Alta v. Paz; Schwarz and Barber coll. (USNM type 69794). ALLOTYPE.—@, collected with holotype; in the same collection. PARATYPES.—2 oo’ and 4 99, Panama, C.Z., Bohio; Schwarz coll., 6-9.II.; 1 9, Porto Bello; Schwarz coll., 26.II; in the same and Kormilev collections. Remarks.—Aneurus pusillus is the smallest known American species of Aneurus. It belongs to the group with evenly rounded scutellum and relatively large hypopygium. Its relation to other species is seen from the key. 2. Aneurus aibonitensis, new species Figures 7, 8 FrmaLe.—Hlongate ovate; head, pronotum, and lateral borders of abdomen, densely and relatively roughly punctured; lateral borders of pronotum finely serrate. NO. 3657 ANEURUS—KORMILEV 5 Head shorter than width across eyes (11.5:13). Anterior process conical; jugae slightly shorter than clypeus, their tips still visible from above; tip rounded, reaching to apical fourth of antennal seg- ment I. Antenniferous tubercles short, acute, divergent. Eyes large, semiglobose, protruding. Postocular tubercles strong, dentiform, acute, with minuscule serrations on the foreborder. Vertex trans- versely rugose, with two (1+1) large, ovate callosities mesad of eyes. Antennae moderately robust, twice as long as head (21.5:11.5); antennal segment I obovate, thick; II obovate, but half as thick as I; III and IV subcylindrical, very slightly tapering toward the base. Relative lengths of antennal segments I to IV: 4:4:4.5:9. Labium short, reaching line connecting hind borders of eyes. Pronotum less than half as long as maximum width (12.5:27.5). Collar very thin, feebly sinuate in front. Anterior borders truncate; anterolateral angles angularly rounded, produced neither forward, nor sideways; lateral borders of forelobe straight, convergent; laterai borders of hind lobe slightly convex; both finely, but distinctly serrate. Hind border sinuate in the middle, convex laterally. Foredise with four (2+2) low callosities; hind disc densely punctured and trans- versely striate along posterior border laterally. Scutellum semicircular, shorter than basal width (12.5:18); disc concentrically rugose. Hemelytra reaching slightly over foreborder of tergum VII; corium very short, reaching basal one-fourth of scutellum; membrane densely and deeply punctured. Abdomen elongate ovate, longer than maximum width across segment IV. Connexivum moderately wide; connexiva III to V twice as long as wide; exterior borders slightly raised, densely granulate; mesad of granulation obliquely rugose around callosities. Tergum VIIT roughly, obliquely rugose. Paratergites very short, serrate, reaching middle of segment IX; latter subtruncate, slightly sinuate in the middle of posterior border. Spiracles II and V to VII lateral and visible from above, III and IV ventral and placed far from border, VIII terminal. Cotor.—Reddish brown; membrane darker, with pale, opaque, basolateral border. MeasurEMENTs.—Total length 3.70 mm, width of pronotum 1.10 mm, width of abdomen 1.40 mm. Ho.ioryre.— ¢, Puerto Rico, Aibonito BF, Medina coll., VI.1961 (USNM type 69795). Remarks.—Aneurus aibonitensis, new species, is related to A. fisket Heidemann, 1904, but is smaller, narrower, and with different position of spiracles. 286—-783—68——2 6 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 125 3. Aneurus pisoniae, new species Figures 9-11 Ma.ue.—Elongate ovate. Head shorter than width across eyes (co? 12.5:14, 9 13:15). An- terior process conical with rounded tip; reaching apical fifth of anten- nal segment I; jugae not visible from above. Antenniferous tubercles short, subtruncate anteriorly, blunt. Eyes moderately large, pro- truding. Postocular tubercles blunt, with minute granulation, short, not approaching outer borders of eyes. Vertex roughly, transversely rugose, with two (1+1) large, ovate callosities placed mesad of eyes. Antennae moderately thin; antennal segment I obovate, robust; II also obovate, but much narrower; III and IV subcylindrical, slightly tapering toward base. Relative lengths of antennal segments I to IV: | 4.5:4:5:10, 9 5:4.5:5.25:10.5. Labium reaching line connecting the middles of eyes. Pronotum less than half as long as maximum width (co 12.5:29, 9 14:31). Collar thin, sinuate in front. Anterior borders straight, slightly oblique; anterolateral angles rounded and slightly produced forward beyond collar; lateral borders of forelobe straight, converging; lateral borders of hindlobe subparallel, convergent, rounded anteri- orly. Hindborder sinuate in the middle, convex laterally. Foredisc with two (1+1) bean-shaped callosities and a few callous rugae laterad of them. Hind disc finely, transversely striate anteriorly; very finely, longitudinally striate posteriorly. Scutellum subtriangular; lateral borders moderately convex, apex angularly rounded. Scutellum shorter than basal width (o 12.5:19, Q 12.5:20). Disc concentrically rugose along the borders; longitu- dinally rugose in the middle. Hemelytra reaching to the middle of tergum VII in both sexes; corium reaching basal fourth of scutellum. Abdomen ovate in both sexes, longer than wide (o 53:38.5, 9 58:42.5). Connexivum moderately wide; segments IV to VI twice as long as wide. Exterior borders of connexiva slightly raised, granulate; disc around callosities sparsely rugose. Paratergites (o') short, rounded, reaching apical fourth of hypopygium; latter small, shorter than its maximum width (5:6). Paratergites (2) short, forming an obtuse angle, their interior border twice as long as exterior, produced as far as segment IX; latter truncate posteriorly. Spiracles II and VI to VII lateral and visible from above, [II to IV ventral, V placed close to border but not visible from above, VIII terminal. Cotor.—Head, pronotum, and scutellum yellow brown; connex- ivum brown; membrane black; ventral side ochraceous medially, yellow brown laterally. NO. 3657 ANEURUS—KORMILEV yf MeEaAsurEMENTS.—Total length: o&@ 3.70, ? 4.00 mm; width of pronotum: o' 1.16, 9 1.24 mm; width of abdomen: o& 1.54, 2 1.70 mm. Ho.totypr.—o’, Puerto Rico, Maricao; De Leon coll., 16.[V.1940, on Pisonia subcordata (USNM type 69796). AuuotyPE.—®, collected with holotype; in the same collection. Paratypes.—4 oo and 2 99, collected with holotype; in the same and Kormilev collections. REMARKS.—Aneurus pisoniae, new species, is related to A. haitiensis Kormilev (1968, p. 2), but is smaller, with lateral borders of forelobe of pronotum straight, hypopygium more narrowly rounded, para- tergites shorter, and color more contrasting: yellow brown with black membrane. 4, Aneurus froeschneri, new species Ficures 12, 13 FrmMALe.—Elongate ovate, finely granulate. Head slightly shorter than width across the eyes (12.5:13). Anterior process conical, rounded apically, reaching to tip of antennal segment I; jugae reaching almost to tip of clypeus, clearly visible from above. Antenniferous tubercles small, acute, convex exteriorly, and slightly divergent. Eyes moderately large, protruding. Postocular tubercles strong, dentiform, acute, reaching as far as outer margins of eyes. Vertex finely, transversely rugose, with two (1+1) large, ovate callosities mesad of eyes. Antennae slender, less than twice as long as head (20.25:12.5); antennal segment I stout, obovate; II narrowly obovate; III and IV cylindrical; relative lengths of antennal segments I to IV: 4:3.75:3.75:8.75. Labium reaching line connecting middles of eyes. Pronotum less than half as long as maximum width (11.5:27). Collar thin, sinuate in front. Anterolateral angles rounded, produced forward beyond collar; lateral borders of forelobe together with lateral notch slightly sinuate; lateral borders of hind lobe slightly convex, converging anteriorly; both finely serrate, particularly in the lateral notch. Hind border sinuate medially, angularly produced laterally. Foredisc with two (1+1) large ovate callosities, and granulate laterad of them. Hind disc very finely granulate anteriorly and medially; very finely, transversely striate along posterior border laterally. Scutellum shorter than basal width (11.5:19); lateral borders weakly convex, apex evenly rounded. Hemelytra reaching almost to hind border to tergum VII. Corium reaching basal third of scutellum; membrane densely and roughiy punctured. & PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 Abdomen elongate ovate, longer than maximum width across segment IV (53:38). Connexivum moderately wide. Paratergites small, rounded, reaching as far as segment IX; latter truncate pos- teriorly. Spiracles II and VI to VII lateral and visible from above, III to V ventral and placed far from border, VIII terminal. Cotor.—Dark yellow brown; connexivum brown; membrane dark brown. MerasurEMENTS.—Total length 3.64 mm, width of pronotum 1.08 mm, width of abdomen 1.52 mm. Houoryrr.—9, Panama, El Valle; N.L.H. Krauss coll., XI.1946 (USNM type 69797). REMARKS.—It is a pleasure to name this species for Dr. Richard C. Froeschner, by whose kind offices I have had an opportunity to study this lot as well as many other Aradidae from the collections in his charge. Aneurus froeschnert, new species, is related to A. minutus Bergroth, 1886, from which it differs as indicated in the key. 5. Aneurus politus Say Figures 14-16 Aneurus politus Say, 1832, p. 31. Matzr.—Elongate ovate; shiny. Head as long as width across eyes (o' 12.5:12.5, 2 15:15). Anterior process conical, rounded anteriorly, reaching slightly beyond the tip of antennal segment I; jugae adherent to clypeus, not visible from above. Antenniferous tubercles short, truncate anteriorly, not acute. Eyes moderately large, protruding. Postocular tubercles stout, dentiform, blunt, produced as far as lateral margins of eyes. Vertex transversely rugose, with two (1+1) mederately large, ovate callosities mesad of eyes. Antennae almost twice as long as head (co 24:12.5, 9 29.5:15); antennal segment I barrel shaped; II and III clavate; IV elongate fusiform; relative lengths of antennal seg- ments I to IV: o& 3.5:5:5.5:10, 9 4:6.5:7.5:11.5. Labium reaching line connecting hind borders of eyes. Pronotum less than half as long as maximum width (o' 10:25, Q 12.5:30). Anterior borders straight, slightly oblique; anterolateral angles rounded, slightly produced beyond collar; lateral borders of forelobe together with lateral notch moderately sinuate; lateral borders of hind lobe slightly convex, strongly converging anteriorly; hind border sinuate, medially. Foredisc with four (2+2) large, ovate, blurred callosities. Hind disc very finely, transversely striate an- teriorly and along hind border laterally; shiny in the middle. Scutellum transverse, much shorter than basal width (o& 10:16.5, Q 11:20); tip widely arcuate. Disc with an elongate callous spot NO. 3657 ANEURUS—KORMILEV 9 in the middle anteriorly; concentrically striate around callosity. Hemelytra reaching to the middle of tergum VII; corium reaching basal third of scutellum; membrane finely punctured, finely wrinkled around punctures; exterior border of membrane smooth, transparent. Abdomen longer than maximum width across segment IV (1 55:37.5, 9 60:56). Connexivum moderately wide. Paratergites () long, subtriangular, rounded apically, reaching apical fourth of hypo- pygium; latter large, elongate, rounded at base and apically (10:8). Paratergites (2) large, rounded apically, produced far beyond small and short segment IX, latter slightly sinuate posteriorly. Spiracles IT and VII lateral, visible from above; III to VI ventral, placed far from border; VIII terminal. Cotor.—Reddish brown; forelobe of pronotum, tergum, proster- num, meso- and metasternum around coxae, pale yellow brown; membrane brown. MEAsUREMENTS.—Total length: o 3.60, 2 4.20 mm; width of pronotum: o 1.00, 2 1.20 mm; width of abdomen: &@ 1.50, 2 2.24 mm. Neorypr.—o', U.S.A., Florida, Biscayne; 21.V.1887, ex Uhler collection (USNM type 69798). EXAMINED SPECIMENS.—1 Q, Florida, Biscayne; 26.V.1887, ex Ubler collection (USNM). 1 9, Florida, Belle Air; ex collections of Mrs. A. T. Slosson (AMNBH). 6. Aneurus championi, new species Figures 17-19 FremaLe.—Elongate ovate, shiny. Head almost as long as width across eyes (9 15.5:16, @ 15:14.5). Anterior process conical, reaching three-fourths of antennal segment I; tips of jugae slightly visible from above. Antenniferous tubercles acute exteriorly, slightly divergent. Eyes semiglobose, protruding. Postocular tubercles dentiform, acute, slightly produced beyond outer border of eyes (9), or reaching to latter (co). Infraocular, ovate, callous spots large. Vertex transversely rugose. Antennae slender, less than twice as long as head (Q 27.5:15.5, & 24.5:15); antennal segment I obovate; II clavate; III subcylindrical, tapering toward base; IV cylindrical; relative lengths of antennal segments I to IV: 9 5.5:5.5:5.5:11, o 5:5:5:10.5. Labium reaching line con- necting hind borders of eyes. Pronotum less than half as long as its maximum width (9 15:34, o 12.5:30). Collar sinuate. Anterolateral angles rounded, very slightly produced forward beyond collar. Lateral borders parallel at humeri, strongly convergent and sinuate anteriorly. Hind border sinuate in the middle. Foredise with two (1+1) bean-shaped calloused spots and with a few semifused callosities laterad of them; granulate 10 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 in front of callosities. Hind disc finely longitudinally striate, with exception of two (1+1) transverse, glabrous spots. Scutellum subtriangular with subtruncate or widely arcuate apex. Dise concentrically rugose around mediobasal longitudinal rugae. Hemelytra reaching middle of tergum VII (9), or hind border of tergum VI (<‘). Abdomen longer than its maximum width across segment IV (2 69:50, o& 60:46). Connexivum relatively wide; PE-angles barely protruding. Tergum VIII (2) small, narrower than head (12:16). Segment IX more than one-third width of head (6:16); paratergites rounded, reaching segment IX. In male, paratergites small, divergent, produced beyond small, transverse hypopygium; the latter much shorter than wide (3:5). Spiracles: II, VI, and VII lateral and visible from above; III and IV ventral and placed far from border; VIII terminal. Cotor.— Yellow brown, venter paler; membrane infuscate; labium and tarsi yellow. MEAsuREMENTS.—Total length: 9 4.68, o 3.90 mm; width of pronotum: @ 1.36, @ 1.20 mm; width of abdomen: ? 2.00, o* 1.84 mm. Houotyrr.—9, Guatemala, Chicacao; E. J. Hambleton coll., 7.VII.1945 (USNM type 69799). ALLoTyPE.— co’, Mexico, Vera Cruz, Pefuela; H. Dybas coll., 17.VII.1941; deposited in the Field Museum of Natural History, Chicago. ParatyPes.—2 92 and 1 4, collected with allotype; in the Field Museum of Natural History, Chicago, and Kormilev collection. REMARKS.—This species is named for the eminent British ento- mologist, G. C. Champion, who did the first comprehensive survey of Central American Aradidae. 7. Aneurus inconstans Uhler Fieurss 1-3 Aneurus inconstans Uhler, 1871, p. 105. In the collections examined, this species was sometimes confused with A. septentrionalis Walker. The key given earlier plus the three drawings will permit ready separation of the two forms. Literature Cited Bererortn, E. 1866. Ueber einige amerikanische Aradiden. Wiener Ent. Zeit., vol. 5, pp. 97-98. CHAMPION, G. C. 1898. Aradidae. Pp. 65-117 (with 3 pls.) in vol. 2 of Heteroptera in God- man and Salvin, Biologia Centrali-Americana. Curtis, J. 1825. British entomology, vol. 2, pl. 86. London. HEIDEMANN, O. 1904. Notes on North American Aradidae, with descriptions of two new species. Proc. Ent. Soc, Washington, vol. 6, pp. 161-165. Hussey, R. F. 1957. Two changes of name in Hemiptera (Aneuridae and Miridae). Flor- ida Ent., vol. 40, p. 80. KormiLey, N. A. 1960. Notes on Neotropical Aradidae, XI (Hemiptera). Journ. New York Ent. Soc., vol. 68, pp. 208-220. 1966. Notes on Aradidae in the U.S. National Museum, IV. Proc. U.S. Nat. Mus., vol. 119, no. 3548, pp. 1-25. 1968. Notes on Aradidae in the U.S. National Museum, V. Proc. U.S. Nat. Mus., vol. 125, no. 3651, pp. 1-16, 1 fig. Say, T. 1832. Descriptions of new species of heteropterous Hemiptera in North America, 39 pp. New Harmony, Indiana. [Reprinted in Le Conte, ed., Complete writings of T. Say ... , 1859, vol. 1, pp. 310— 368.] UHLER, P. R. 1871. Notices on some Heteroptera in the collection of Dr. T. W. Harris. Proc. Boston Soc. Nat. Hist., vol. 14, pp. 93-109. WaALkKER, F. 1873. Catalogue of the specimens of Hemiptera Heteroptera in the collection of the British Museum, pt. VII, pp. 1-213. 1a E 12 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 125 Ficures 1-19.—Aneurus inconsians Uhler 1871: 1, head, pronotum, and scutellum, o; 2, apex of abdomen from above, o; 3, same, 9. Aneurus pusillus, new species: 4, head pronotum, and scutellum, o’; 5, apex of abdomen from above, o’; 6, same, 9. Aneurus aibonitensis, new species: 7, head, pronotum, and scutellum, 9 ; 8, apex of abdomen from above, 9. Aneurus pisoniae, new species: 9, head, pronotum, and scutellum, o; 10, apex of abdomen from above, o*; 11, same, 9. Aneurus froeschneri, new species: 12, head, pronotum, and scutellum, 9; 13, apex of abdomen from above, 9. Aneurus politus Say 1832: 14, head, pronotum, and scutellum, co; 15, apex of abdomen from above, o';16,same, 2. Aneurus championi, new species: 17, scutellum, co; 18, apex of abdomen from above, o’; 19, same, ?. U.S. GOVERNMENT PRINTING OFFICE: 1968 Proceedings of the United States National Museum SMITHSONIAN INSTITUTION +« WASHINGTON, D.C. Volume 125 1968 Number 3658 Interspecific Competition in the Tropical Rain Forest: Ecological Distribution Among Lizards at Belém, Para By A. Stanley Rand and Stephen S. Humphrey! The ecological differences between sympatric species have proved useful foci for ecological studies, particularly those with an evolu- tionary viewpoint. The information provided by such studies is relevant to considerations of interspecific competition and to the problem of the causes of tropical species diversity. In general, it is assumed that a species’ closest competitors in an area are its nearest relatives. This is probably true within generic and specific groups, but, as this study suggests, it may not be true for higher categories. Studies on the ecology of sympatric lizards (usually restricted to sympatric congeners) have been made at several localities by various workers, among them Milstead (1957), Inger (1959), Collette (1961), Rand (1964). Little has been published about the ecology of the lizards of lowland tropical forests and almost nothing about those in the rain forests of the Amazon. The study most comparable to the present one is that of Sexton, Heatwole, and Knight (1964) in Panama. *Rand: Smithsonian Tropical Research Institute, Balboa, Canal Zone; Humphrey: 612 Louisiana St., Lawrence, Kansas. 1 2 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 As a result of the observations made in conjunction with other field studies, the authors have collected considerable information about the natural history of lizards in the vicinity of Belém, Para, Brazil. Some of this has already been published (Rand, 1965; Rand and Rand, 1966), and additional papers are planned. In this paper we present our data on microdistribution patterns among the common diurnal lizard species and its relevance to possible interspecific com- petition among lizards. We also present our data on lizard body temperature and their relation to habitat distribution. The senior author, who has made four trips to Belém, has spent a cumulative total of almost four months there between 1963 and 1966. These visits were all made during the period of May through August, a time of intermediate rainfall. The junior author visited Belém from July through August, 1964, and from June through August, 1965. The observations reported herein were made in the vicinity of the city of Belém, Par4, at lat. 1°27’S, long. 48°30’W, altitude less than 100 feet. The climate is hot tropical with abundant rain and a marked but not severe or prolonged dry season in the southern winter. The climax vegetation is forest, of which three types are locally recog- nized: “mata de terra firme” or upland forest; “varzea” or forest that is flooded daily by the freshwater tides; and “igapo” or permanently flooded swamp forest (Ducke and Black, 1953). We were able to sample the first two forests much more adequately than the latter. In addition to the above, we visited a variety of sites that had been disturbed by man and were in various stages of regeneration—from scattered bunches of grass and bare ground to old second growth. The main localities where our observations were made are as follows: Parque do Museu Goeldi — a small patch of forest in the city, the undergrowth much modified and denser than normal because of edge effect in some places, cleaned out in others, but the canopy only slightly disturbed. Bosque Municipal — a somewhat larger but still small patch of forest in the city, modified in much the same way as the Parque do Museu, but less so. Utinga and Agua Preta — localities in the preserve around the city water supply on the edge of the city, which include “mata de terra firme,” selectively cut but quite good in places, and various ages of second growth from just abandoned “rocas” through dense tangles to forest that betrays its second growth nature only in the species composition. IPEAN and Guama Forest Reserve — localities of “mata de terra firme,” “varzea” and some “igapo” forest. A variety of ages of NO. 8658 LIZARDS—RAND AND HUMPHREY 3 second growth and agricultural situations also were sampled in this area. The following species were seen frequently enough to provide some idea of habitat distribution: Gonatodes humeralis, Iguana iguana, Tropidurus torquatus, Plica wmbra, Uranoscodon superciliosa, Poly- chrus marmoratus, Anolis punctatus, Anolis fuscoauratus, Cnemido- phorus lemniscatus, Ameiva ameiva, Kentropyx calcaratus, Tupi- nambis nigropunctatus, Mabuya mabuya. All are diurnal and together these species include over 99 percent of all the lizards we saw active during the day. Two geckos, Hemidactylus mabouia and Thecadactylus rapicaudus, the first a common species, are omitted because they are nocturnal. Also seen only rarely were Anolis ortoni, Amphisbaena alba, Croco- dilurus lacertinus, and several microteiids. We must thank Dr. Dalcy Albuquerque, Director of the Museu Paraense “Emilio Goeldi,” and Dr. Robert Shope, then Director of the Belém Virus Laboratory, for their help during our visits to Belém. We are much indebted to Dr. P. Humphrey of the University of Kansas and Dr. P. Vanzolini of the Departamento de Zoologia, Sao Paulo, for their field assistance and encouragement and for arranging support for the field work of the junior and senior authors, respec- tively. We also wish to thank the directors of the Instituto de Pesquisas Experimentacio Agropecuarias do Norte (IPEAN) and the Belém Virus Laboratory for their many courtesies and assistance. SPHAERODACTYLIDAE Gonatodes humeralis Hasrrat.—Found only in the forest and in old second growth in the “mata de terra firme” and, less commonly, in the “varzea.” They were not found in the young second growth or in open areas without a closed canopy. These geckos perch above the ground on the trunks of small to very large trees, on large logs and stumps. They are almost always within one and one-half meters of the ground. Rarely were they seen on twigs or leaves or in small bushes, but most frequently on buttressed trees, trees with loose bark, or many small branches or suckers against the trunk that provide complex hiding places, but they are not restricted to such trees. They may leave a perch to hide in leaves on the ground and sometimes come to the ground to catch food. Axsunpancy.—This is the most frequently seen lizard in the forest and probably the most abundant. It was most common in the somewhat 4 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 disturbed areas (Bosque and Parque), where densities of several per 100 square meters were probably reached. In the less disturbed and more extensive forests they were much less common. Usually a single lizard was seen on a tree, occasionally two (a male and female), but seldom more. Foop.—Insects and other small invertebrates. ‘TeEMPERATURE.—Probably a nonheliotherm, it was never seen bask- ing, but no temperatures were taken. IGUANIDAE Iguana iguana Hapsrrat.—Very few of these large lizards were seen but they are reported to be most common along the edges of the rivers. One was seen on the edge of a large clearing. It is quite possible that they live in the canopy of the forest away from clearings and rivers as well as on edges, but they are very difficult to see in such situations. They did well in the forested Parque do Museu Goeldi. In most forested localities (Panama, Swanson, 1950; Costa Rica, Hirth, 1963; British Honduras, Neill and Allen, 1959), iguanas are highly arboreal lizards as adults, though one occasionally comes to the ground. The juveniles are reported as living closer to the ground. Apparently this is true around Belém for local men who hunt them say that they live on the branches of trees. Those running loose in the Museu Goeldi Parque were seen usually 10-20 meters up in the larger trees. ABUNDANCE.—We saw them rarely, even when we were examining the river margin. Due to hunting they are probably very shy. Foopv.—Primarily herbivorous and probably largely eat leaves. TEMPERATURE.—A. heliotherm (McGinnis and Brown, 1966). No temperatures were taken but iguanas in the Parque do Museu Goeldi were seen frequently basking. Tropidurus torquatus Hasrrat.—F ound only in open areas and along the edges of clear- ings, and not in dense second growth or forest. These lizards perch above the ground, usually below two meters, on tree trunks, logs, rocks, walls, stumps, etc. They seldom climb above three meters except when frightened and are very rarely seen in bushes, on twigs, leaves, etc. They frequently come to the ground to feed. AxsunpDAnce.—This is the most frequently seen lizard in the open and probably the most common. Only one adult male is seen per perch but sometimes one or two females and several juveniles may be associated with him. No. 3658 LIZARDS—RAND AND HUMPHREY 5 Foopv.—Insects and other invertebrates. Trmrrrature.—A heliotherm (fig. 2). It frequently basks and shows difference in posture related to substrate temperatures. (For more detailed data on this species, see Rand and Rand, 1966.) Plica umbra Hasrrat.—F ound only in the forest and older second growth. These lizards occur in the “mata de terra firme” and, at least occasionally, in the “varzea” and are not found in the open or on isolated trees. Individuals of Plica umbra were seen on the trunks and branches of moderate to large trees from close to the ground up to at least several meters. None were seen in bushes or on twigs. Foop.—Insects and other invertebrates. Anunpancn.—Moderately common. Seldom were two seen on the same perch, but several times a male and a female were found only a few meters apart, more frequently than we would have expected by chance alone. Perhaps some sort of pair bond is formed. Temprraturn.—A nonheliotherm (fig. 2). No basking behavior was seen. Uranoscodon superciliosa Hasrrat.—Found only along the shaded margins of streams and lakes where the forest interior comes to the edge of the water in both “mata de terra firme” and “varzea.” Individuals were seen up to two meters high on moderate to large diameter tree trunks and on logs at the edge or in the water, some of which the lizards could have reached only by swimming. They fre- quently jumped into the water when approached and sometimes dove to the bottom and hid. ABUNDANCE.—This species is moderately common in its habitat. No more than one individual was seen on a perch. Foop.—Insects and other invertebrates. Trmprerature.—A nonheliotherm (fig. 2). No basking behavior was seen. Polychrus marmoratus Hasrrat.—An animal collector for the Belém Virus Laboratory took large numbers in second growth along the edges of forest. Our few specimens were found in similar places. It is likely that this species also occurs in the crown of the forest, as Rand has seen it in such habitats in Trinidad. Polychrus can and does climb easily on small twigs and branches. ABUNDANCE.—Though we saw few individuals, many were taken in the area by the Virus Laboratory collector. 6 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 126 Foop.—Insects and other invertebrates plus an appreciable amount of plant material. TEMPERATURE.—Probably a heliotherm (fig. 2). Anolis punctatus Hasrrat.—In the forest, found in “mata de terra firme” and older second growth; not in “varzea” though this lizard probably occurs there at least occasionally. It was not found in the open, on isolated trees, or in young second growth. Individuals were seen most commonly on the trunks and branches of moderate to large trees up to four meters. They probably also climb up into the branches and lower crown. One individual was seen once coming to the ground to catch something. ABUNDANCE.—This is a moderately abundant species; usually single individuals were found, occasionally a male and female were found on the same perch or adjacent perches. Foop.— Insects and other invertebrates. TemperaTure.—A nonheliotherm (fig. 2). No basking behavior was seen. Anolis fuscoauratus Hasrrat.—In the “mata de terra firme,” in old second growth and at least edge of “varzea,” and not in the open areas on isolated trees or bushes, nor in young second growth. Individuals were seen sometimes on tree trunks but somewhat more frequently on bushes and slender twigs close to the ground; they were found usually below two meters but occasionally as high as three or four, and also they were seen on the leaf litter on the ground. AsuNnDANCE.—Moderately common. Usually isolated individuals were found, but once a male and tiny juvenile were found on the same tree. There is some suggestion that they group themselves in loose col- onies with adjacent home ranges rather than distributing themselves evenly or randomly over the available habitat. Foov.—Insects and other invertebrates. Tremperatoure.—A nonheliotherm (fig. 2). No basking behavior was seen. TENDAE Cnemidophorus lemniscatus Hasrrat.—Found in open areas where the vegetation is so sparse that there is bare, unshaded ground exposed between tussocks of grass or bushes and not in the forest or even young second growth. The juve- niles seem to occur in the most open parts of the habitat. NO. 3658 LIZARDS—-RAND AND HUMPHREY 7 The species is terrestrial though individuals occasionally climb short distances up bushes or onto logs and hide in holes in the ground. AsunpDancn.—Occurring in only a few places but abundant there. Foov.—Insects and other invertebrates. TemperAturse.—A heliotherm (fig. 2). Individuals were seen bask- ing and were active only in the heat of the day on sunny days. Ameiva ameiva Hasrrat.—Primarily along edges of clearings in forest and second growth. It ranges out into open areas but seldom far from bushes and other cover, and also into second growth and forest but seldom more than a few meters from an edge. Ameivas turn up in clearings com- pletely surrounded by forest, indicating that they do disperse through the forest. Ameivas are terrestrial, occasionally climbing on rocks and logs, hid- ing in holes in the ground and among rocks. ABUNDANCE.—Widespread and common in edge and edificarian situations. Foop.—Insects and other invertebrates. Temprraturs.—A heliotherm (fig. 2). Ameivas were seen basking; they are usually active only during the heat of the day and primarily on sunny days. Kentropyx calcaratus Hasrrat.—Primarily in the forest, “mata de terra firme,” old second growth, and high spots in the “varzea”; very occasionally individuals were seen In open areas. Though terrestrial, they not uncommonly were seen climbing upon a log, the base of a tree, or branches of a fallen treetop. They hide in holes in the ground and under logs. ABUNDANCE.—Widespread in the forest but only moderately common. Foopv.—Insects and other invertebrates. ‘TemprrsatTure.—A heliotherm (fig. 2). Individuals were seen bask- ing in sun patches on the forest floor; they were active only in the mid- dle of the day and on sunny days. Tupinambis nigropunctatus Hasrrat.—This seems to be a wide-ranging species, moving out into open areas, occurring in second growth and far inside the forest in “mata de terra firme” and “varzea.” Largely terrestrial, it hides in holes in the ground. Axsunpance.—Not commonly seen but, because it is hunted, it is very shy. 8 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 Foop.—A variety of animal prey including small vertebrates. TemMPERATURE.—Probably a heliotherm, as several were seen basking (no temperature records). ScINCIDAE Mabuya mabouia Hasrrat.—Most common along the edges of forest and of second growth, it also occurs well inside the forest and out in open grassy situ- ations where hiding places are available. With the possible exception of Tupinambis, this has the widest habitat range of all the species con- sidered in this paper. This species is partly terrestrial, but also it is seen frequently on logs, wood piles, and even short distances up tree trunks. It is one of the few species regularly seen in tall grass areas, where they are seen on tree trunks and on posts that extend above the grass. ABUNDANCE.—Moderately common, least so inside the forest. Foop.—Insects and other invertebrates. TemperAture.—A heliotherm (fig. 2). Frequently seen basking. Discussion The ecological profile given in figure 1 summarizes the microdistri- butions of the lizards under consideration in the Belém area. No species ranges throughout the whole spectrum of the area though some species certainly are more widely distributed than are others. Of the habitats studied, the forest is the richest in number of species, but most of the species there are not abundant—at least they are not commonly seen. Some of the species high in the forest canopy and others in the leaf litter are certainly more abundant than indicated by the number of times that we saw them. Restricting consideration to areas where our observations are best, among the species that live on tree trunks near the ground or are conspicuously active on the ground, most of the species in the relatively undisturbed forest seem rarer than those outside it. The localities where most of the forest spe- cies were most common—the Parque do Museu and the Bosque Munici- pal—were places where the canopy was only slightly disturbed but the understory more so and where many things, particularly certain snakes, birds, and mammals that might be predators or competitors, were probably absent. In the forest habitats not only density of individuals per species, but also total density of individuals of all lizard species taken together seemed less than in some more disturbed areas. Second growth has a lizard fauna much like that of the forest except that certain species are missing and a few like Amezva are more com- NO. 3658 LIZARDS—RAND AND HUMPHREY 9 mon. Canopy species, particularly Polychrus marmoratus, are seen here, more frequently than in older forest apparently because the canopy is closer to the ground. Clearings and edges have few species but they are sometimes ex- tremely abundant, particularly Z’ropidurus torquatus. These species are undoubtedly native in areas such as Belém, where rain forest is the climax vegetation, but they also occur in other major habitat areas such as “cerrado,” where rain forest is absent. Though the lizards typical of clearings disperse through and even live inside the rain forest (e.g., abuya), they are never abundant there. IGUANA en | ANOLIS PUNCTA POLYCHRDS MARMORATUS PLICA 7 | rheusl RATUS URANOSCODON. HUMERALIS SUPERCILIOSA jena MABOUIAI, ORQUA ax KENTROPYX CALCAI i Me U TUPINAMEIS NIGROPUNCTATUS Ficure 1.—Habitat distribution in Belém lizards: Schematic cross section of Belém habitats from upland forest, with a small forest stream at left, through second growth and grassland to bare ground at right (each species located in area of maximum observed abundance). There are a number of factors that may be involved in making the forest the environment where the largest number of species occur. Of the environments in the Belém region, the forest comprises the greatest area, it is the most stable on any one site, and it contains the widest floristic diversity and probably the most numerous species of other animals, (These factors were all certainly true before the advent of modern man.) All of these may well be involved in promoting species diversity, and this paper provides no evidence for emphasizing or excluding any of them. Figure 1 suggests that the greater structural complexity of the forest may also be important. There are lizards in the forest occupying structural situations that do not exist or exist only rarely outside the forest. Their restriction to the forest may be, in part, the direct result of the fact that the forest’s more complex structuring provides places for the lizards to live. The physical structures in an environment that a lizard uses as look- outs, hiding places, basking spots, etc., form an important aspect of the ee ee 10 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 environment from the lizard’s point of view. They have been called the lizard’s “structural niche” (Rand, 1964). When the term was pro- posed for West Indian anoles, it was found that certain measurements of the perches on which the lizards were seen provided a useful descrip- tion. Though the concept of structural niche seems useful for Belém lizards, measurements used for West Indian anoles are not sufficient and other environmental parameters must also be considered. In the case of Uranoscodon superciliosa, the proximity of water seems im- portant. No system of quantifying the various aspects of structural niches was devised, and they have been described verbally in the fore- going species accounts. Another set of parameters, in addition to structural niche, must be considered in describing the ecological distribution of lizards. These can be grouped as the lizard’s “climatic niche.” There are a variety of environmental factors that show parallel gradients here, among them insolation, light, temperature, and humidity. In general, one set of extremes for all these factors in the area is reached on or near the floor of the mature forest, and the other set is reached on the ground in clearings devoid of vegetation. Edges, second growth, and the top of the canopy show intermediate conditions. The various lizards arrange themselves differently with respect to these gradients. To understand better the relationships of the lizards to the microclimates, we took a number of cloacal temperatures of the former while they were active in periods when the sun was shining; behavioral thermoregulation was possible at this time. In West Indian anoles similar data showed that the lizards in shaded habitats had lower body temperatures than those in open habitats (Ruibal, 1961; Rand, 1964). The Belém data are a bit more complicated. Figure 2 shows the body temperature plotted against air temperatures taken in the immediate vicinity at the same time. These plots suggest that there are two very different kinds of thermal] relationships represented among the Belém lizards : those whose temperature approximates that of the surrounding air and those whose temperature is much above it. A review of observa- tions made of the various species shows that those in the first class were not seen basking while those in the second class were observed with definite basking behavior. Regarding this behavior, it seems safe to conclude that the high temperatures of the second group are due to behavioral thermoregulation and that they are heliotherms while the other group is not—at Jeast at the ambient temperatures observed. (Since their temperature approximates the air and there is no evidence that they use substrates for thermoregulation, we prefer not to use “thigmotherm” but rather the more neutral “nonheliotherm.”) NO. 3658 LIZARDS—-RAND AND HUMPHREY tia | There is a definite relationship, but not unvarying, between thermal relationship and habitat used. The nonheliotherms are all inside the forest, and this is thermally the most constant habitat. The heliotherms occur in the more extreme environments in clearing, second growth, and canopy. One species of heliotherm, however, is almost entirely restricted to the area within the forest (Hentropyx), where it basks in the numerous patches of sun that reach the forest floor. URANOSCODON PLICA UMBRA SUPERCILIOSA ANOLIS ANOLIS FUSCOAURATUS PUNCTATUS | POLYCHRUS CNEMIDOPHORUS MARMORATUS LEMNISCATUS Ficure 2.—Temperature relationships in Belém lizards: Cloacal temperature plotted against ambient air temperature at time of capture (temperatures in centigrade of just- captured lizards in natura! habitat during day). It should be noted from figure 2 that, under the conditions tested, the nonheliotherms in the forest are about as homeothermic as are the heliotherms that practice behavioral thermoregulation outside the for- est (see Rand and Rand, 1966, for details of thermoregulatory behavior in Tropidurus). It certainly seems likely that the body temperatures of the open habitat animals would fluctuate much more than they are ob- served to do if the lizards did not practice some sort of thermo- regulation. These data suggest that, if heliothermy has evolved to increase homeothermy rather than to produce high body temperature per se, then it has probably evolved in response to the fluctuating con- ditions in open environments. The data also suggest that the high body temperatures observed in heliotherms reflect the general principle | PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 that, if an animal is to be diurnally active, heating is easier to accom- plish than cooling in an area of high insulation. The foregoing indicates that Kentropy« may be a recent invader of the forest from a macroteiid ancestor that was adapted basically to living in open habitats. Though Hentropyz is the only primarily forest heliotherm, other heliotherms like Z’wpinambis and Mabuya also occur there. Low eccritic temperatures usually are associated with low temperature tolerances. Because of this and because the upper threshold of tolerance is usually sharper and more quickly lethal than the lower, it is probably easier and safer for open habitat animals to disperse through the forest than for forest animals to disperse through open habitats. The role of interspecific competition is a basic problem in animal ecology. One of the reasons for our interest in the distribution of Belém lizards was that the evolution of differences in microdistribution has been suggested as an important strategy in reducing interspecific com- petition among sympatric lizard species (Rand, 1964; Milstead, 1957; and others). The basis for this suggestion in lizards has come usually from studies of sympatric congeners, but there seems no reason that it could not occur between sympatric relatives of different genera. Kcologists( e.g., MacArthur, Recher, and Cody, 1966) who attempt to discover environmental parameters that will predict bird species di- versity seem to be assuming that a phylogenetic group as large as birds can be treated as an ecological unit for these purposes. We can make a similar assumption about the Belém lizards but, as we will show, it is probably wrong. The clearest cases wherein differences in microdistribution can be associated with interspecific interactions are those wherein two or more species occur in the same area, utilize the same structural niche, and replace one another in different habitats or different climatic niches. The only case of such replacement suggested by the distributions plotted in figure 1 is the three smaller macrotelids. There is some over- lap between these, but each species reaches its maximum abundance in a different environment: Cnemidophorus in the bare open patches, Ameiva in the edges and second growth, and Hentropyzx in the forest. All three have similar foraging habits and could well compete if they occurred syntopically. There are suggestions that size may be playing a role in reducing interaction between overlapping species. This is a strategy much seen in birds (Schoener, 1965, and others). In general, larger lizards take larger prey and this tends to reduce food competition among species of different sizes. In the Belém lizards, the very large macroteiid Tupinambis nigropunctatus widely overlaps Ameiva ameiva and Ken- NO, 3658 LIZARDS—-RAND AND HUMPHREY 13 tropyx calcaratus. Tupinambis is several times the size of the others and as an adult takes much larger food. Another suggestive case is the two forest Anolis that occupy very similar structural and climatic niches but differ sharply in size. The third case is the difference between Polychrus and Iguana, the two lizards of the tree crowns. It seems un- likely that this size difference is related to food competition since Iguana is largely and Polychrus is partly herbivorous. There is no evidence of the sort of temporal replacement during the day that Inger (1959) recorded in forest skinks in Borneo. Diurnal versus nocturnal replacement (perhaps involving the large nocturnal trunk gecko of the forest, Thecadactylus rapicaudus) is possible, but it is an unstudied problem. The only major diet difference we now know (our examination of stomach contents has been limited and casual and there is much still to be learned) lies between the two species that regularly take a consider- able amount of leaves and flowers, 7guana iguana and Polychrus mar- moratus, and the remainder, which take largely or exclusively animal food. This diet difference plus the difference in structural niche sepa- rate the two herbivorous species very sharply from the rest. To return to microdistribution as a mechanism for reducing species competition, there is considerable difference among species with respect to structural niches as there is with respect to climatic niches. Taken together, the microdistributions (fig. 1) do not show a regular pattern of replacement of one lizard by another because of the presence of overlaps between species (e.g., Plica umbra and Anolis punctatus) and of gaps between others (e.g., Uranoscodon and T'ropidurus torquatus) in distribution. Since there is a lack of regular replacement, micro- distribution does not suggest itself as an important factor in reducing interspecific competition among the lizards in general, though it may be so between certain species. As we have discussed, our observations on other strategies for reducing possible interspecific competition sug- gest that, even when all are considered together, no regular pattern of ecological separation among species appears. We are left with the im- pression, which we cannot prove rigorously, that the relationships of the lizard species to each other are not the over-riding factors in shap- ing their ecologies. There are several possible explanations. It may be that our data are so incomplete that we have been given a completely erroneous impres- sion. Further study may show that other species occur at Belém. This may fill the gaps in our picture and reveal differences among common species that would eliminate the apparent overlaps. Certainly, much more is to be learned and the latter possibility cannot be eliminated, 14 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 but, if for no other than heuristic reasons, it cannot be accepted as the only possibility. There are two other possibilities not mutually exclusive that seem to us more likely. One is that lizards do not make an ecological unit. They, as a group, do not occupy a “lizard niche” that is subdivided among them; rather, the broad niche or niches that lizards occupy are occupied also by organisms of widely differing phylogenetic rela- tionships with the result that several lizards in the same habitat may — each have as its close competitors animals that are not lizards at all but other vertebrates or even invertebrates. It may not be coincidental that the most convincing studies of interspecific competition and mecha- nisms for reducing it in lizards come from the arid areas of North America and from those areas of the West Indies where lizards are very abundant both as individuals and species and play an important role in the trophic ecology. Certainly, one has the impression in the Belém area that inside the forest the disappearance of all lizards would not greatly upset the forest ecological economy if at all. The relative rarity of lizards in the Belém forests suggests another possible explanation for the apparent lack of regularity in interspecific relationships. This is that populations of lizards are held at such low levels (perhaps by predation) that the populations never reach densities wherein interspecific competition for food or space occurs. It may be relevant that forest lizards are most common in the park areas, where many of the larger predatory birds, mammals, and per- haps snakes are rare or absent; these are also areas where the number of competitors may also be reduced. Certainly, it is striking how little agonistic behavior one sees among iguanids in the Belém forests. This is in contrast to desert iguanids and those in the West Indies, where fighting or displaying between males is a common sight. One has the impression that the problem the forest lizards are usually faced with is not that of spacing out the population to avoid overcrowding but rather that of bringing together mates at the proper time. Among Plica umbra, Anolis punctatus, and Gonatodes humilis, we have the impression that we found individuals of the opposite sex near one another more often than we would have expected on chance alone. It may be that these lizards form pairs or at least that males and females permanently establish adjacent or overlapping home ranges in preference to living solitarily. The absence of territorial defense and the large amount of potential home range that seems unoccupied suggest that population densities are so low that direct intraspecific competition for exhaustible resources is nonexistent, and it also suggests that probably there is no inter- NO. 38658 LIZARDS—-RAND AND HUMPHREY 15 specific competition between most lizard species and, consequently, no selection pressure for mechanisms that would reduce the competition. If this is true, how can one account for the observed specialization in both climatic and structural niche? The answer is, of course, intraspecific competition in the Darwinian sense. When any species can exploit the available resources (in the broadest sense) more effectively by specializing in part of them, the specialization itself will be selected for unless this selection pressure is countered by selection in other directions. Specialization, by definition, involves loss of eco- logical amplitude, and selection against specialization is largely generated by temporal fluctuations in environmental conditions that generate a selection pressure for ecological amplitude (Pianka, 1967). In stable habitats—and the inside of the tropical rain forest is one of the most stable—the selection against specialization should be at its lowest. It may be that low counterselection against specialization, rather than any unusually strong selection pressure for it, has produced some of the extreme specializations we see in tropical forest forms. We have not the data to choose between these two hypotheses, which, in fact, are not mutually exclusive. It seems most likely that lizards as a block do not occupy a monolithic trophic niche but share a niche in a complex way with many other animals. It also seems likely that the low population levels in the forest lizards result in mini- mizing the importance of selective forces that arise from interspecific competition between many lizard species. Conclusions (1) The Belém area contains several different habitats and no species of lizard studied ranges uniformly through the area. (2) The forest is richest in number of species and the evidence suggests that this is at least in part because of its greater structural complexities. (3) Two types of thermal relationships with the environment are demonstrated by the lizard fauna: the heliothermic in both open and forest habitats and the nonheliothermic inside the forest. The latter show about as much homeothermism as the former because of the relatively low ambient temperature variation inside the forest. (4) The distribution of three species of macroteiid suggests that they re- place one another ecologically. A fourth macroteiid that is syntopic with at least two of these is much larger. Two Anolis species that are syntopic and occupy very similar structural niches are different in size. (5) These replacements are probably important, but the im- pression of the distributions is one of irregular overlaps and gaps between niches of species. (6) The apparently unoccupied niches are 16 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 125 probably in part filled by animals that are phylogenetically dis- tantly related to lizards, but ecologically the former should be con- sidered with the latter. The lizards do not make an ecological unit. (7) The low population densities within the forest are such that interspecific competition may be relatively unimportant and ecological specialization may be the result of intraspecific competition for maxi- mum efficiency of resource exploitation. Selection pressure for this must always exist, but the tropical rain forest, because of its great continuous extent and stability, probably generates unusually few counterselection pressures and, consequently, a high relative selection pressure toward specialization. Literature Cited CoLuetTtTE, B. B. 1961. Correlations between ecology and morphology in anoline lizards from Havana, Cuba and southern Florida. Bull. Mus. Comp. Zool., vol. 125, no. 5, pp. 137-162. Ducks, A., and Buackx, GrorGsE A. 1953. Phytogeographical note on the Brazilian Amazon. An. Acad. Brasileira Ciene., vol. 25, no. 1, pp. 1-46. Hirtna, H. F. 1963. Some aspects of the natural history of Jguana iguana on a tropical strand. Ecology, vol. 44, no. 3, pp. 613-615. IncER, R. F. 1959. Temperature responses and ecological relations of two Bornean lizards. Ecology, vol. 40, no. 1, pp. 127-136. MacArruor, R.; Recuer, H.; and Copy, M. 1966. On the relation between habitat selection and species diversity. Amer. Nat., vol. 100, no. 913, pp. 319-332. McGinnis, SAMUEL M., and Brown, CHARLES W. 1966. Thermal behavior of the green iguana, Iguana iguana. Herpetologica, vol. 22, no. 3, pp. 189-199. Miustwap, W. W. 1957. Some aspects of competition in natural populations of whiptail lizards (genus Cnemidophorus). Texas Journ. Sci., vol. 9, no. 4, pp. 410-447. Neiuu, W. T., and ALLEN, R. 1959. Studies on the amphibians and reptiles of British Honduras. Publ. Res. Div. R. Allen’s Reptile Inst. Inc., vol. 2, no. 1, pp. 1-76. PianKA, Eric R. 1967. On lizard species diversity: North American flatland deserts. Ecology, vol. 48, no. 3, pp. 333-351. Ranp, A. S. 1964. Ecological distribution in anoline lizards of Puerto Rico. Ecology, vol. 45, no. 4, pp. 745-752. 1965. On the frequency and extent of naturally occurring foot injuries in Tropidurus torquatus (Sauria, Iguanidae). Pap. Avuls. Dept. Zool. Séo Paulo, no. 17, pp. 225-228. NO. 3658 LIZARDS—RAND AND HUMPHREY 7 Ranp, A. S., and Ranp, P. 1966. Aspects of the ecology of the iguanid lizard Tropidurus torquatus at Belém, Pard. Smithsonian Misc. Coll., vol. 151, no. 2, pp. 1-16. RviBaL, RoDoLFo 1961. Thermal relations of five species of tropical lizards. Evolution, vol. 15, pp. 98-111. ScHorNnER, THomas W. 1967. The ecological significance of sexual dimorphism in size in the lizard Anolis conspersus. Science, vol. 155, no. 3761, pp. 474-477. Sexton, O. J.; HEATwoLe, Haroup; and Knieut, DENNIS 1964. Correlation of microdistribution of some Panamanian reptiles and amphibians with structural organization of the habitat. Caribbean Journ. Sci., vol. 4, no. 1, pp. 261-295. Swanson, P. L. 1950. The iguana Iguana iguana iguana (L). MHerpetologica, vol. 6, pp. 187-193. U.S, GOVERNMENT PRINTING OFFICE: 1968 Pa: ete er ee id 7 % a a Ts sta ys ous me whee a , ny Manet ae al i an a ! ary Pi} a UORwiL ae UF a SUE E ieee sh ure un..!! _ hi of : HY qe, Gili pcre q wi Ae 7 fa i beat fi - is au ante ih vhs =} uv - a | Ch i, ; = ’ y y el ee | a 7 oF Tur Th ri awe rh ilaeh ie ee | re “00h a) 1 ' Bi Ades a ® Np rue : io aarie ie yi ie ' - f : : iv : ie ui —_ Mipol oy ne vim ie? ay a is % Ame rr thy He These pit iy ee a i ns Te ‘7 er ae a ee ya ay i. ae wiih i A inate mH ie id aay. or it i * Vale ah ma i 1 > a) Wh ei et MA ra kL { i ; = 7 aie NM 4 ne “4 aa thee ee Tye a Aa insole phan he Rr | = hy 1 pre fy pean tw ea nti Age “1 ae a a Ae 4 Boe or Dun Ah jhe hinges well ioe oa Ma vee oe Tike ro Yd AAV tit, Gui past a 4 atla i ot jugnitel: ube Hie aGU Rt ay me ip) Se a, \ mt sh epeaey ae : yo Toman + oliylit = a “ae [ ; Mi! ALG - ANDROS fh Gili wy 7 ‘ile ’ .? 4 Wu = 7 & 7 ; q! Jah: tur Ob =i " -,& : tp) eps : os” >* ; ‘ ah ee L640 Le Ree: eid #7 ti : perma 44) Abdi er 2 : ‘ei. as ae Oates. By 1, ory 5 Lon a) Wie NBO ein ee ye Ris, i pes) eh ar 2 \ bet Fg Cire a oS ‘0d rom dank) ; ieee , @ : “4 = lie Gy Tate! ‘peter & 7A ) & 7 ane ie G6 TR ( Cd De ih i 2nsss, 8 eee ee ee \ erect > Pee Olen se i's (aes 4 oo ee 4a Were *) i >rieg t* 3s > ‘i ‘rae ie 1 wales e os ioe my _ a * > eighivee Waele as Dyed sth Sate Maa § Cas | HP nage Trt) mt > a Peter oe Pata 28k eel eng tea Rew be ey ie ce. ee Kersey » , aa dint ati, wis, Saeed oP ¥ ty | (Teo ERS piety Marea. of ee 1 , Bee Neem tort = hy Peas — ek ie Weert mary Broecee Gin ss of mee United States National Museum SMITHSONIAN INSTITUTION +- WASHINGTON, D.C. Volume 125 1968 Number 3659 Records and Descriptions of Additional Scoliidae from New Guinea, Bismarck Archipelago, and Solomon Islands (Hymenoptera: Aculeata) By Karl V. Krombein Chairman, Department of Entomology Several hundred Scoliidae have been obtained in New Guinea, Bismarck Archipelago, and Solomon Islands by field parties of the Bernice P. Bishop Museum, Honolulu, since those reported upon in my revisionary study of these wasps (Krombein, 1963). I am erateful to J. L. Gressitt and C. M. Yoshimoto for supplying me the specimens for study. I have prepared the following brief paper in- asmuch as included among the specimens are several distinctive new species and subspecies of Campsomeris subgenera Laevicamp- someris and Phalerimeris, and Campsomeris, sensu lato, the previously unrecognized males of two other species belonging to the subgenus Laevicampsomeris, and the previously unknown female of Diliacos r. ribber (Betrem). Also, many specimens came from a number of new localities and provide additional data on seasonal occurrence of many species. The species are arranged in systematic order as in my revisionary paper. The nomenclature employed is also the same except for the changes necessitated by two recent papers on classification of the Scoliidae (Betrem, 1962, 1967; Bradley and Betrem, 1967). 1 2, PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 125 Campsomeris (Laevicampsomeris) bonguensis Betrem Northern District, Papua: 30’, Popondetta, 60 meters, 2 and 26 September 1963, P. Shanahan and J. Sedlacek, in Malaise trap. Central District, Papua: 3 o, Cape Rodney, 10 meters, 2-4 November 1960, J. L. Gressitt, 1 in Malaise trap. 2 #, Otomata Plantation, east of Port Moresby, 1 meter, 2 November 1960, J. L. Gressitt, in Malaise trap. 1 o, Brown River, 23 October 1960, J. L. Gressitt. West New Guinea: 12, Dawai River, south-southeast of Sumber- baba, Japen Island, 3 November 1962, N. Wilson. 1 &, Nabire, 5-50 meters, 25 August-2 September 1962, J. Sedlacek, in Malaise trap. 1 &, Kebar valley, west of Manokwari, 550 meters, 4-31 January 1962, L. W. Quate. Campsomeris (Laevicampsomeris) nigerrima (Smith) Northern District, Papua: 19 , Popondetta, 60 meters, 26 September 1963, P. Shanahan, in Malaise trap. West New Guinea: 12, Dawai River, south-southeast of Sumber- baba, Japen Island, 3 November 1962, N. Wilson. Bismarck Archipelago: 22, 1 <, Illugi, Upper Warangoi, Gazelle Peninsula, New Britain, 230 meters, 8-15 December 1962, J. Sedlacek. 1 &, Upper Warangoi, Gazelle Penin., 250-600 meters, 28-30 Novem- ber 1962, J. Sedlacek. 2 #, Gaulim, Gazelle Penin., 130-140 meters, 23-28 October and 19-20 November 1962, J. Sedlacek. 19, Mt. Sinewit, Gazelle Penin., 900 meters, 7-16 November 1962, J. Sedlacek. Males of nigerrima have not been recognized previously. These key to laglaizer Betrem, couplet 9 (Krombein, 1963, p. 553), and agree with that species in details of punctation and coloration. The genitalia of both laglaizer and nigerrima appear to be indistinguishable from those of bonguensis Betrem. The males of nigerrima recorded above are 18-23 mm long, and the forewing is 16-21 mm. It is presumed that the single bonguensis male recorded earlier from New Britain (Krom- bein, 1968, p. 568) is actually a specimen of nigerrima. Campsomeris (Laevicampsomeris) laglaizei Betrem West New Guinea: 12, Dawai River, south-southeast of Sumber- baba, Japen Island, 3 November 1962, N. Wilson. Campsomeris (Laevicampsomeris) brandti Krombein Bismarck Archipelago: 19, 1 &, Lelet Plateau, Schleinitz Mts., New Ireland, October 1959, W. W. Brandt. This pair bears exactly the same label data as the female holotype (Krombein, 1963, p. 570), the only specimen known previously. The No. 3659 SCOLIIDAE—KROMBEIN 3} male is 18 mm long with the forewing 16 mm. The genitalia are identical with those of bonguensis Betrem and other known Melanesian males of the subgenus Laevicampsomeris. The male keys to solomonis Krombein, couplet 10 (Krombein, 1963, p. 553), and cannot be separated from males of that species except by reference to label data. Campsomeris (Laevicampsomeris) solomonis solomonis, new status I am reducing this taxon to subspecific rank because of the discovery of the distinctive new subspecies described below. Solomon Islands: 1 &, Kukugai, Bougainville, 150 meters, Novem- ber 1960, W. W. Brandt. 1 o&, Ulo Crater, Vella Lavella Island, 10 meters, 16 December 1963, P. Shanahan. 1 o, Pepele, Kolombangara Island, 0-30 meters, 3 February 1964, P. Shanahan. 12, 107, Dala, Malaita Island, 50 meters, 9-14 and 21 June 1964, J. and M. Sedlacek. 2 #, Tambalia, 30 km west of Honiara, Guadalcanal, 22-25 May 1964, R. Straatman, J. and M. Sedlacek. 19, Betikama River, Guadalcanal, August 1960, W. W. Brandt. 19, Wugiroga, San Cristoval, 9 August 1960, C. W. O’Brien. 1 o, Kira Kira, San Cris- toval, 0-50 meters, 10 November 1964, R. Straatman. Campsomeris (Laevicampsomeris) solomonis sedlaceki, new subspecies At first glance, this anomalous member of the subgenus Laevicamp- someris reminds one of a female of Campsomeris (adumeris) extranea levert Krombein without the narrow yellow bands on the first three abdominal terga; however, it has such Laevicampsomeris characters as the lack of a carina between the dorsal and lateral propodeal sur- faces, the very short, impunctate, nontuberculate median area of the dorsal propodeal surface, the pronounced oblique furrow on the scapula, large impunctate areas on the front, vertex, and thoracic dorsum, and the lack of a second recurrent vein in the forewing. What makes it so distinctive among the taxa of Laevicampsomeris is the bright fulvous vestiture on the entire body and the yellowish wings, whereas the vestiture and wings are black in the more typical members of the subgenus. In my key the female runs to solomonis, couplet 8, if one disregards the character of wing color in couplet 4. It is separated at once from typical solomonis by the color of the vestiture and wings. I assume that the male from Gizo Island, which I included in the type-series of typical solomonis, is actually the opposite sex of solomonis sedlaceki. It has the black integument, vestiture, and wings of solomonis males from other of the Solomon Islands. If this assumption is correct, we have in solomonis sedlaceki a taxon of Laevicampsomeris exhibiting a most unusual form of sexual dimorphism. 4 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 Ho.otryrr.— , Gizo Island, New Georgia Group, Solomon Islands, 100 meters, 16 July 1964, J. and M. Sedlacek, Bishop Museum. Length 23 mm, forewing 18 mm. Exactly like typical solomonis in punctation and shape of longer hind tibial spur, but the vestiture fulvous, wings yellowish and lacking the green and blue reflections, and veins testaceous. Maue.—Presumably identical to typical solomonis in color of vestiture and integument. PARATYPE.—1 @ ,same data as holotype, but 19 July 1964, USNM. It is very similar to the holotype but is only 20 mm long. Campsomeris (Laevicampsomeris) quatei, new species This is known from a single female from West New Guinea. It is unique among females of this subgenus in having the entire posterior half of the vertex, scutellum, postscutellum, and median area of dorsal surface of propodeum with rather close punctures. It keys to couplet 8 and is distinguished from nigerruma (Smith) and solomonis Krombein as noted above. Ho.oryprse.—, Bokondini, 40 km north of Baliem Valley, ca. 1300 meters, West New Guinea, 16-23 November 1961, L. W. Quate, Bishop Museum. Length 25 mm, forewing 22 mm. Integument black; wings dark brown with purplish reflections, the margin of forewing darker. Vestiture black except that on clypeus and a few hairs on vertex light brown. Clypeus with only a small median area and the apical rim impunc- tate; area frontalis and spatium frontale coarsely and closely punc- tate, upper part of front with scattered punctures; a narrow space anteriorly on vertex impunctate, elsewhere closely punctate. Pronotum with moderately large, close punctures bearing stout erect setae, a narrow strip along posterior margin and anterior declivity impunctate; scutum with large contiguous punctures on anterior half and lateral third posteriorly, the median posterior space nearly smooth except for some punctures near midline; disks of scutum and scutellum with moderately sized, subcontiguous punctures except along midline; metapleuron and propodeum devoid of large punctures except median area of dorsal surface of propodeum with mostly subcontiguous, moderately large punctures. Longer spur of hind tibia spatulate at tip, about two-thirds as long as hind basitarsus. Maue.— Unknown. Campsomeris (Laevicampsomeris) papuana, new species The presence of abundant pale hair on the front, sides, and venter of head, the sides and venter of thorax (dorsum also in male), legs, No. 3659 SCOLIIDAE—-KROMBEIN 5 and on part (female) or all (male) of the abdomen distinguish this handsome species at a glance from other Melanesian members of the subgenus Laevicampsomeris. The female runs to couplet 8 of my key (Krombein, 1963, p. 558). It has spatulate hind tibial spurs and moderately dense punctures on scutellum and postscutellum. This combination of characters coupled with the distinctive vestiture separate it from nigerrima (Smith) and solomonis Krombein, the two species keyed out in that couplet. The male keys to couplet 9. It is separated from laglaizei Betrem and the species keying out in the subsequent couplet, solomonis and bonguensis Betrem, by a combination of characters such as the pale erect vestiture, lack of a propodeal tubercle, moderately dense punc- tation of the propodeal dorsum, and small yellow mark opposite the posteroventral margin of the eye. The genitalia, as is true in all Melanesian Laevicampsomeris, afford no differentiating characters. Houotyrr.—@, Karimui, south of Goroka, Eastern Highlands Dist., Terr. New Guinea, 1000 meters, 6 June 1961, J. L. and M. Gressitt, Bishop Museum. Length 29 mm, forewing 24 mm. Integument black; wings fuscous, the forewing with purplish reflections. Erect vestiture brown to black on clypeus, vertex, dorsum of thorax including scapulae, terga three to six, and sterna five and six, whitish with an ocherous cast on front, sides, and venter of head, sides and venter of thorax, legs, and first and second abdominal segments; appressed preapical fringes of sterna two to four pale, apical fringes on sterna dark except for a few pale intermixed setae on second; sides and venter of thorax with abundant, silvery, sericeous pubescence. Clypeus contiguously punctate except middle; area frontalis, spa- tium frontale, and a narrow strip on lower half of front between eyes densely and coarsely punctate; vertex and occiput densely and coarsely punctate except for a longitudinal median strip on vertex; setae arising from frontal, vertexal and occipital punctures coarse, erect and moderately long. Scapulae, and anterior and lateral areas of scutum with dense, small punctures bearing conspicuous, erect setae, the impunctate median space on posterior half of scutum about a third as wide as scutal width and with a few scattered larger punctures; scutellum, postscutellum, and propodeal dorsum with subcontiguous punctures except for a narrow median strip; propodeum not tuberculate. Posterior areas of terga and sterna shining; terga two to four mat on anterior two-thirds. Legs with spurs of hind tibia spatulate at apex as in bonguensis, the longer one two-thirds as long as basitarsus. AuLotyPpr.—o’, Karimui, 3 June 1961, J. L. and M. Gressitt, in Malaise trap, Bishop Museum. 6 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 Length 24 mm, forewing 20 mm. Integument mostly black, the dorsum of thorax and abdomen with weak blue reflections; the follow- ing creamy to yellowish: clypeus above and broadly on sides, narrow strip along inner eye margin to ocular sinus, short stripe adjacent to posteroventral eye margin, and apical half of foredistitarsus; wings fuscous, the forewing anteriorly with darker streak. Head, thorax, legs, and abdomen clothed with erect white setae except for a few intermixed dark setae on vertex and scutum, and the sixth and seventh terga with all dark setae; silvery sericeous pubescence on sides of thorax very conspicous, longer and denser than in female. Propodeum with denser, larger punctures than in bonguensis, those on dorsal surface separated from each other by about the diameter of a puncture, tubercle absent; genitalia as in bonguensis (Krombein, 1963, p. 597, fig. 18). PARATYPE.—1 <’, same data as holotype, but 5 June 1961, J. L. and M. Gressitt, in light trap, USNM. The paratype is 25 mm long and agrees with the allotype in vestiture, punctation, and coloration. Couplets 9 and 10 of my key may be expanded to include the males of these additional species of Laevicampsomeris as follows: 9. Median area of dorsal surface of propodeum with a distinct tubercle posteri- orly, most of punctures on this area large and contiguous to subcon- tiguous ... Sixes See OS Median area of dorsal Shei of pueondeumn noe foberculats most of punc- tures on this area finer and separated by more than the diameter of a puncture except in papuana and solomonis .. . ¢ hockae see 9a. Side of clypeus with narrow yellow streak; pieeedenl pUNeration denser on dorsum in ne many of punctures contiguous; New Guinea, Waigeu. «7. . -.. . . .Campsomeris laglaizei Betrem Clypeus entirely Bick: Poronodcal punctation slightly sparser on dorsum in middle, many of punctures separated from each other by about half the diameter of a puncture; New Guinea, Misool, New Britain. Campsomeris nigerrima (Smith) 10. Head, thorax, and most of abdomen with long, erect white hair; punctures on dorsum of median area of propodeum larger and denser, usually separated by no more than the diameter of a puncture; New Guinea. Campsomeris papuana, new species Erect hair almost entirely dark; propodeal ees finer and more sepa- rated except in solomonis. . . . 3 Ae peer LO) 10a. Clypeus with a narrow yellow stripe lone fateral mere prooedaal punc- tation fine and well separated; New Guinea. ee ae bition Betrem Clypeus entirely black . .... sap ee aeyelOb 10b. Foredistitarsus with a small yellowish spot Deane on median area of propodeal dorsum finer and shallower, many of them separated by more than the diameter of a puncture; New Ireland. Campsomeris brandti Krombein Foredistitarsus black; punctures on median area of propodeal dorsum coarser, many of them separated by less than the diameter of a puncture; Solomon Islands. . .... .. . . Campsomeris solomonis Krombein NO. 3659 SCOLIIDAE—-KROMBEIN a Campsomeris (Lacvicampsomeris) formosa (Guérin) Northern District, Papua: 3 9, Popondetta, 25 meters, June 1966, P. Shanahan and Lippert, in light trap. Central District, Papua: 1 9, Bisianumu Station, 40 km north- northwest of Port Moresby, 29 April 1960, C. W. O’Brien. 1 o’, Brown River, 23 October 1960, J. L. Gressitt. Morobe District, Terr. New Guinea: 15 2, 6 o&, Wau, 900-1300 meters, 25 February 1962, 23 March 1963 and 1964, 27-31 March 1964, 12 April 1963, 13 April 1964, 27 June 1961, 27 July 1964, 14, 15, 16, and 19 August 1964, 11 September 1964, 17 September 1961, 11 October 1962, 19 October 1965, 15 December 1961, J. and M. Sedlacek, C. Montieth, P. Shanahan, 3 o& in Malaise trap. 1 9, 1 o&, Mt. Kaindi, Wau, 900-1800 meters, 12-14 June 1963, P. Shanahan. 3 9, 2 o’, Mt. Misim, Wau, 900-1100 meters, 16 February 1963, 14 August and 25 September 1964, J. and M. Sedlacek. 1 o, Bulldog Road, ca. 14 km south of Edie Creek, 2405 meters, 4-10 July 1966, G. A. Samuelson, in Malaise trap. 1 9, Pindiu, Huon Peninsula, 860 meters, 22 April 1963, J. Sedlacek. 1 9, Singaua River, Lae, 30 meters, 8 April 1966, O. R. Wilkes. Eastern Highlands District, Terr. New Guinea: 9 o, Kassam, 48 km east of Kainantu, 1350 meters, 30 October 1959, T. C. Maa. West New Guinea: 1 9, 1 o, Ifar, Cyclops Mts., 300-900 meters, 23-25 June 1962, J. L. Gressitt and J. Sedlacek. 1 o&, Kampong Landbouw, 30 km northeast of airstrip, Biak Island, 16 July 1957, J. L. and M. Gressitt, in light trap. 1 o&, Sucumi Camp, near head of Ransiki River, Vogelkop, 300 meters, 6 August 1957, D. E. Hardy. Bismarck Archipelago: 1 9, Riat, Upper Warangoi, Gazelle Peninsula, New Britain, 250 meters, 1 November 1962, J. Sedlacek. 1 9, Illugi, Upper Warangoi, Gazelle Penin., 15 December 1962, J. Sedlacek. 1 o&, Gaulim, Gazelle Penin., 130 meters, 23-28 October 1962, J. Sedlacek. Solomon Islands: 1 &, Kokugai, Bougainville, 150 meters, October 1960, W. W. Brandt. 2 &, Gollifer’s Camp, Kolombangara Island, 100-700 meters, 22-23 January 1964, P. Shanahan, one in Malaise trap. 1 9, Kolosulu, Guadalcanal, 20 May 1960, C. W. O’Brien. 1 &, Betikama River, Guadalcanal, August 1960, W. W. Brandt. The male from Gaulim, New Britain, has more reduced yellow markings than in any specimens noted in my revision. The scutellar spots are reduced in size, the band on postscutellum is lacking, reduced posterolateral spots are present on the first two terga only, and the sternal spots are smaller on two and three. The Vogelkop male lacks yellow maculations on both the scutellum and postscutel- lum, but abdominal bands are present on the first three terga, only 8 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 125 those on the third being narrowly separated. The vestiture in the latter specimen is normal for formosa. Campsomeris (Radumeris) tasmaniensis (Saussure) Northern District, Papua: 1 9, 1 o&, Popondetta, 20-60 meters, May 1966 and 2 September 1963, P. Shanahan and J. Sedlacek, 3 in Malaise trap. Central District, Papua: 1 &, Boroko, Port Moresby, 6-7 Novem- ber 1960, J. L. Gressitt, in Malaise trap. 1 &@, Otomata Plantation, east of Port Moresby, 1 meter, 2 November 1960, J. L. Gressitt, in Malaise trap. 1 o, Bisianumu Station, 40 km north-northwest of Port Moresby, 29 April 1960, C. W. O’Brien. Western District, Papua: 1 9,1 o, Daru Island, 2-3 meters, 19 July 1964 and 24-25 October 1960, J. L. Gressitt and H. Clissold, 3 in Malaise trap. 1 9, Oriomo River, 6 meters, 14 February 1964, H. Clissold, in light trap. Morobe District, Terr. New Guinea: 1 9, 1 o&, Wau, 1200 meters, 12 April 1963 and 19 August 1964, J. Sedlacek. West New Guinea: 7 o, Nabire, south of Geelvink Bay, 0-50 meters, 2-9 July and 25 August-20 September 1962, J. L. Gressitt, J. Sedlacek, H. Holtman, 1 in light trap and 6 in Malaise trap. 2 @, Enarotadi, Wisselmeeren, 1800-1900 meters, 27 July 1962, J. Sedlacek. Campsomeris (Radumeris) extranea moluccensis Betrem Bismarck Archipelago: 2 o&, Kandan, New Britain, 24 December 1959, W. W. Brandt. 1 o, Talliligap, Gazelle Peninsula, New Britain, 300 meters, 17-18 December 1962, J. Sedlacek. Solomon Islands: 1 9, Fauro, northeast Solomon Islands, 12 April 1964, P. Shanahan. One of the males from New Britain has the yellow band on the fourth tergum narrowly interrupted in the middle; the other male bearing identical label data has the normal coloration for this subspecies. In my revision (Krombein, 1963, p. 585) I recorded eztranea moluccensis from localities in New Guinea, Bismarck Archipelago, and Solomon Islands. In the latter group it was present only on the northernmost island, Bougainville, and was replaced by eztranea lever. Krombein on the more southern islands, Santa Ysabel, Russell, Ulawa, and Guadalcanal. The present shipment from the Bishop Museum includes a female and two males of extranea moluccensis labeled as having been taken at Kira Kira, San Cristoval, 27-28 July 1960, by C. W. O’Brien. I am of the opinion that these specimens are mislabeled because San Cristoval lies to the southeast of Guadalcanal, NO. 3659 SCOLIIDAE—-KROMBEIN 9 and we would expect to find it occupied by eztranea levert. This record becomes even more suspect when we note that in this material there is a specimen of the eastern New Guinea race, Campsomeriella mano- kwariensis lanhami (Krombein), bearing a “Kira Kira, San Cristoval’” label as well as several specimens of Austroscolia nitida nitida (Smith), another taxon occurring in New Guinea and the Bismarcks. Both Campsomeriella manokwariensis and Austroscolia nitida are represented by other races in the Solomons. It appears probable that some of these presumably mislabeled specimens may have been collected in eastern New Guinea because of the specimen of manokwariensis lanhami, a race that occurs on the Huon Peninsula, Markham River Valley, and at Port Moresby. Campsomeris (Radumeris) extranea leveri Krombein Solomon Islands: 1 &, Tambalia, 35 km west of Honiara, Guadal- canal, 30 meters, 22-25 May 1964, J. and M. Sedlacek. Campsomeris (Radumeris) extranea shanahani, new subspecies The single female of this previously unrecognized taxon from Gizo Island in the New Georgia group differs from females of the other races of eaztranea in lacking yellow integumental markings on the abdomen and in the very reduced yellow spot adjacent to the upper inner eye margin. Disregarding the lack of yellow abdominal markings it keys without difficulty to extranea (couplet 23). The putative male, represented by a short series from Vella Lavella Island in the New Georgia group, runs to eatranea (couplet 34) if one disregards the lack of a yellow band on the postscutellum. It differs from males of the other races of eztranea in having a very tiny yellow spot adjacent to the upper inner eye margin and in lacking yellow markings on the pronotum and postscutellum. Houotyps.—®, Gizo Island, New Georgia group, Solomon Islands, 100 meters, 20 July 1964, J. and M. Sedlacek, in Malaise trap, Bishop Museum. Length 18 mm, forewing 14 mm. Black, yellow markings lacking except for a tiny spot adjacent to upper inner eye margin. Vestiture dark fulvous on head and thoracic dorsum, whitish on sides of thorax and first abdominal segment; apical fringes of all terga and of fourth and fifth sterna dark brown, of second and third sterna light brown; decumbent setae on pygidium dark brown. Wings and punctation as in extranea lever. ALLOTYPE.— ’, Pusisama, Vella Lavella Island, Solomons; Novem- ber 1963, P. Shanahan, in Malaise trap, Bishop Museum. Length 19.5 mm, forewing 14.5 mm. Black, abdomen with weaker blue reflections than in eztranea leveri, pale yellow maculations as 288-033—68——2 10 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 follows: mandible at base, sides and apex of clypeus more narrowly than in eatranea leveri, line along inner eye margin to top of antennal insertions, very tiny spot in ocellocular area, narrow streak along posterior eye margin on lower two-thirds, a pair of small transverse spots on scutellum, tiny posterolateral spot on first tergum, a some- what larger, transverse posterolateral streak on outer fourth of second tergum, a similar but shorter posterolateral streak on second sternum, small spot at apex of anterior surface of mid and hind femora, ex- terior surface of foretibia, and outer surface of forebasitarsus and distitarsus. Erect vestiture dark brown, appressed pubescence on side of thorax silvery. Wings slightly more infumated than in eztranea leverr. PARATYPES.—2 co’, same data as allotype. 167, same data as allo- type but 25 November 1963, USNM and Bishop Museum. Paratypes are 17-19 mm long. The color pattern is the same as described for the allotype except that one specimen has larger spots on the scutellum that are separated only narrowly on the midline, and the third tergum has a short posterolateral streak on the outer fourth. The genitalia are identical with those of the other subspecies of eztranea. Campsomeris (subgenus?) oresbios oresbios Tuijn Morobe District, Terr. New Guinea: 1 9, 2 o’, Wau, 1200 meters, 10 November 1965, 9 December 1961, and 17 December 1965, J. and M. Sedlacek and P. Shanahan, in Malaise trap. Bismarck Archipelago: 1 o, Gaulim, Gazelle Peninsula, New Britain, 130 meters, 23-28 October 1962, J. Sedlacek. This race has not been recorded previously from New Britain, although it has a wide distribution in New Guinea. This specimen agrees well with darker maculated and haired specimens from New Guinea except that there is slightly less yellow on the clypeus and the wings are darker. Additional material, especially of females, may demonstrate that the population occurring in the Bismarck Archipelago should be recognized as a race discrete from the New Guinea taxon. Campsomeris (subgenus?) microspatulata Tuijn Morobe District, Terr. New Guinea: 1 9, Wau, 1200 meters, 3 March 1963, J. Sedlacek. 1 o&, Finschhafen, 14 April, J. and M. Sedlacek. 1 9, ca. 8 km west-northwest of Lab Lab, Umboi Island, 300 meters, 8-19 February 1967, G. A. Samuelson, in Malaise trap. West New Guinea: 1 <, Nabire, south of Geelvink Bay, 10-40 meters, 7 October 1962, N. Holtmann, in Malaise trap in jungle. Campsomeris (Phalerimeris) loriae loriae (Mantero) Central District, Papua: 1 &, Laloki, 10 February, F. Muir. Eastern Highlands District, Terr. New Guinea: 1 o, Kassam, 48 km east of Kainantu, 1350 meters, 30 October 1959, T. C. Maa. NO. 3659 SCOLIIDAE—-KROMBEIN Tt West New Guinea: 3 9, 3 6%, Sibil Valley, Star Mts., 1250 meters, 18 October-8 November 1961, S. and L. Quate. 1 o, Ifar, Cyclops Mts., 300-500 meters, 28-30 June 1962, J. L. Gressitt and J. Sedlacek, in light trap. 5 9, 4 &, Bokondini, 40 km north of Baliem Valley, about 1300 meters, 16-23 November 1961, L. W. Quate. 19,2 o&, Moanemani, Kamo Valley, Wisselmeeren, 1500 meters, 15 August 1962, J. Sedlacek. 1 9, Itouda, Kamo Valley, Wisselmeeren, 1500-1700 meters, 18 August 1962, J. Sedlacek. 1 9, Dawai River, south-southeast of Sumberbaba, Japen Island, October 1962, N. Wilson. Campsomeris (Phalerimeris) loriae millironi Krombein Solomon Islands: 1 2, Buin, Bougainville, 2 June 1956, J. L. Gressitt. 1 9, Pusisama, Vella Lavella, 17-18 November 1963, L. and M. Gressitt. 3 9, 1 o&, Betikama River, Guadalcanal, August 1960, W. W. Brandt. 1 o, Tambalia, 35 km west of Honiara, Guadalcanal, 30 meters, 20 May 1964, J. and M. Sedlacek. Campsomeris (Phalerimeris) loriae kolombangarensis, new subspecies This distinctive melanic race of loriae, known only from males, is separated from the other races of loriae by the reduced yellow markings. Although it is now known to occur only on Kolombangara in the New Georgia group, it is quite likely that it will be found on the other islands in that group when more extensive collecting has been accom- plished. The closest congener, loriae milliront Krombein, is known from Guadalcanal, Florida, Rendova, Vella Lavella, and Bougainville Islands in the Solomons, and from New Britain in the Bismarck Archipelago. It differs from loriae millironi in having the clypeus black except sides, yellow markings very reduced or lacking on the pronotum, and the postscutellum, abdomen (almost entirely), and legs black. The genitalia are identical with those of the other subspecies. Houotypz.—<’, Iriri, Kolombangara, New Georgia group, Solomon Islands, 2 meters, 3 July 1964, J. and M. Sedlacek, Bishop Museum. Length 14 mm, forewing 12 mm. Black, abdomen with faint blue reflections, yellow as follows: mandible on basal two-thirds, spot at lower corner of clypeus, short stripe along inner eye margin to antennal insertion, a pair of tiny spots anteriorly on pronotal disk, and short posterolateral streak on second tergum; tip of mandible and lower edge of clypeus reddish. PaRrATYPES.—3 o’, same data as holotype, USNM and Bishop Museum. These are uniform in size. Two of them lack yellow spots on the pronotum. Campsomeris (Phalerimeris) ferrugineipes ferrugineipes (Mantero) West New Guinea: 3 9, Moanemani, Kamo Valley, Wisselmeeren, 1500 meters, 15 August 1962, J. Sedlacek. 12 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 125 Campsomeris (Phalerimeris) ferrugineipes pembertoni Krombein Bismarck Archipelago: 7 o, Talliligap, Gazelle Peninsula, New Britain, 300 meters, 17-18 December 1962, J. Sedlacek. Campsomeriella (Campsomeriella) manokwariensis manokwariensis (Cameron) West New Guinea: 6 o, Hollandia [Kota Baru], 25-28 June 1962, N. Wilson. 1 9, Bokondini, 40 km north of Baliem Valley, about 1300 meters, 16-23 November 1961, S. and L. Quate. 1 o, Itouda, Kamo Valley, Wisselmeeren, 1500-1700 meters, 18 August 1962, J. Sedlacek. 3 o', Enarotadi, Wissel Lakes, 1800-2000 meters, 2-11 and 27 July 1962, N. Wilson and J. Sedlacek, 2 by sweeping. 1 9, Dawai River, south-southeast of Sumberbaba, Japen Island, 3 November 1962, N. Wilson. 2 9, Biak Island, 22-30 June 1962, J. L. Gressitt and J. Sedlacek, in light trap. 1 o, southeast Biak Island, 1 July 1962, J. L. Gressitt. 9 2, 16 o&, Nabire, south of Geelvink Bay, 0-50 meters, 2-9 July, 25 August-2 September, 1-4, 7, and 17 September, 4, 7, 10, and 13 October 1962, J. Sedlacek, J. L. Gressitt, H. Holtmann, N. Wilson, 1 each in Malaise and light traps. Bismarck Archipelago: 1 9, Kerawat, New Britain, 6 May 1956, G. Dun. Campsomeriella (Campsomeriella) manokwariensis lanhami (Krombein) The Bishop Museum material included one female of this eastern New Guinea race undoubtedly mislabeled as being from Kira Kira, San Cristoval, Solomon Islands, 27 July 1960, C. W. O’Brien (see my earlier remarks on these presumably mislabeled specimens under Campsomeris extranea moluccensis). This particular female agrees with the eastern New Guinea race in having the more lightly infuscated basal two-thirds of the forewing and white tergal fringes on the first four terga in contrast to the Solomon Islands race manokwariensis manni (Krombein), which has similarly colored wings but only the first three tergal fringes white, that on the fourth being brown. Campsomeriella (Campsomeriella) manokwariensis manni (Krombein) Solomon Islands: 1 9, Pusisama, Vella Lavella Island, 29 November 1963, P. J. Shanahan. Trisciloa saussurei Gribodo Morobe District, Terr. New Guinea: 1 o, Wau, 1250 meters, 3 January 1963, J. Sedlacek. 1 o&, Lae, 26 November 1961, J. Sed- lacek. 1 o&, Zenag-Lae, 200 meters, 15 January 1965, J. Sedlacek. West New Guinea: 1 o, Ifar, Cyclops Mts., 300-500 meters, 28-30 June 1962, J. L. Gressitt and J. Sedlacek, in light trap. 19, NO. 2659 SCOLIIDAE—-KROMBEIN 13 50, Kebar Valley, west of Manokwari, 550 meters, 4-31 January 1962, L. W. Quate. 1 #, Oransbani, south of Manokwari, February 1962, L. Richards. Liacos fulgidipennis (Smith) Northern District, Papua: 1 2, Popondetta Mission, 60 meters, 18 October 1963, P. Shanahan. Morobe District, Terr. New Guinea: 1 9, 2 co’, Wau, 1050-1400 meters, 26 May 1965, 9 July 1963, and 11 September 1961, P. Shana- han, J. and M. Sedlacek, 1 & in Malaise trap. 1 9, Finschhafen, Huon Peninsula, 20-150 meters, 15 April 1963, J. Sedlacek. West New Guinea: 1 9, Kebar Valley, west of Manokwari, 550 meters, 4-31 January 1962, S. Quate. Diliacos quadriceps (Smith) In my revision (1963, pp. 613-616) I used the name Scolia (Diliacos) larradiformis Smith, 1863, for a polytypic species occurring in New Guinea and the Admiralty Islands, and I suggested the possibility that the taxon I regarded as typical larradiformis might be identical with quadriceps Smith, 1859, from Aru. Betrem has restudied these types subsequently and has found that they are indeed identical (Bradley and Betrem, 1967, p. 311). The correct names for the taxa, therefore, that I placed under larradiformis are as follows: Diliacos quadriceps grandiceps (Micha), p. 613 Diliacos quadriceps quadriceps (Smith), p. 615 Diliacos quadriceps manus (Krombein), p. 615 Diliacos quadriceps grandiceps (Micha) Morobe District, Terr. New Guinea: 4 9, 2 o‘, Wau, 1050-1200 meters, 27 January 1966, 24 May 1965, and 16 September, 22 October, 9 November 1965, and 19 November 1961, J. and M. Sedlacek, 1 9, 3 co in Malaise trap. 3 o&, Kunai Creek, Wau, 1250-1500 meters, 28-30 May and 26 August 1963, P. Shanahan and J. Sedlacek, 1 in Malaise trap. Diliacos ribbei stueberi (Krombein) Eastern Highlands District, Terr. New Guinea: 3 9, 13-20 km southeast of Okapa, 1650-2250 meters, 24 and 26 August 1964, J. and M. Sedlacek. West New Guinea: 3 2, Hollandia, 0-300 meters, 22 December 1961-21 January 1962, S. and L. Quate. 1 &, Bokondini, 40 km north of Baliem Valley, ca. 1300 meters, 16-23 November 1961, L. W. Quate. 1 o&, Oransbari, northwest of Geelvink Bay, 16 February 1963, R. Straatman. 14 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 125 Diliacos ribbei ribbei (Betrem) Bismarck Archipelago: 2 9, 1 o, Illugi, Upper Warangoi, Gazelle Peninsula, New Britain, 230 meters, 8-11 and 12-15 December 1962, J. Sedlacek. 1 9, Gaulim, Gazelle Penin., 140 meters, 21-26 October 1962, J. Sedlacek. Females of the typical subspecies of ribbet were unknown previously. These specimens from New Britain key to couplet 62 (Krombein, 1963, p. 563). Females of typical ribbei are intermediate in density of punctation between ribbei stueberi (Krombein) from northern New Guinea and ribbei hurdi (Krombein) from the Solomon Islands. As in ribbet stueberi the front between the spatium frontale and the anterior ocellus is impunctate except for a few scattered punctures in the ocular sinus and along the inner eye margin, but the ocellar triangle has one or two punctures instead of being impunctate and the posterior half of the scutum has 15-20 scattered punctures between the parapsidal furrows instead of being impunctate. In contrast, the more densely punctate ribbet hurdi has about 30 punctures on the front between the spatium frontale and anterior ocellus, the ocellar triangle has half a dozen punctures, and the posterior half of the scutum has 44-50 punc- tures between the parapsidal furrows. Each female of ribber ribber has only two discoidal cells in the forewing. They are 13-15 mm long, and the forewing is 10-12 mm long. Diliacos ribbei hurdi (Krombein) Solomon Islands: 1 9, Pepele, Kolombangara Island, 0-30 meters, 3 February 1964, P. Shanahan. 1 Go, Iriri, Kolombangara, New Georgia Group, 2 meters, 3 July 1964, J. and M. Sedlacek. 1 &%, Nalimbu River, 29 km southeast of Honiara, Guadalcanal, 5 June 1960, C. W. O’Brien. Diliacos glabrata glabrata (Micha) Northern District, Papua: 1 9, Popondetta Mission, 60 meters, 18 October 1963, P. Shanahan. 1 9, Gona Road, Popondetta District, 60 meters, 30 October 1963, H. W. Clissold. Morobe District, Terr. New Guinea: 3 9, 1 o&, Wau, 1200-1300 meters, 1 and 14-17 January 1963, 17 May 1965, and 14 November 1965, J. and M. Sedlacek and P. Shanahan, 2 9 in Malaise trap. 1 9, 24 km west of Bulolo, Upper Watut River, 760 meters, 5-6 March 1963, J. Sedlacek. 1 9, ca. 8 km west-northwest of Lab Lab, Umboi Island, 300 meters, 8-19 February 1967, G. A. and S. L. Samuelson, and P. H. Colman. West New Guinea: 1 9, Genjam, 40 km west of Hollandia, 100-200 meters, 1-10 March 1960, T. C. Maa. NO. 3659 SCOLIIDAE—-KROMBEIN Py Diliacos glabrata praslini (Bradley) Bismarck Archipelago: 1 9, Illugi, Upper Warangoi, Gazelle Penin- sula, New Britain, 230 meters, 12-15 December 1962, J. Sedlacek. Solomon Islands: 1 &, Kukugai, Bougainville, 150 meters, October 1960, W. W. Brandt. 1 9, 1 o&, Gizo Island, New Georgia Group, 50- 120 meters, 16-26 April 1964, J. Sedlacek, in Malaise trap. 6 <”, Pepele, Kolombangara Island, New Georgia Group, 30 meters, 7-9 February 1964, P. Shanahan, in Malaise trap. 2 o, Kow, Vella Lavella, 30 meters 28 November 1963, P. Shanahan, in Malaise trap. 7 o, Ulo Crater, Vella Lavella, 10 meters, 16-21 December 1963, P. Shanahan, 4 in Malaise trap. 1 &, Kolotuve, Santa Ysabel, 21 June 1960, C. W. O’Brien. 1 o&, Kolotuve-Sukapisu, Santa Ysabel, 20 June 1960, C. W. O’Brien. 1 9, Takopekope, Florida Group, 12 November 1960, C. W. O’Brien, in light trap. 1 9, Guadalcanal, 12-20, J. A. Kusche. 3 9,4 &%, Betikama River, Guadalcanal, August and September 1960, W. W. Brandt. 1 9, Kiwi Creek, Guadalcanal, 18 September 1944, H. E. Milliron. 2 o&, Napagiwae, San Cristoval, 19 August 1960, C. W. O’Brien. Microscolia maindroni (Betrem) West New Guinea: 1 o, Oransbari, northwest of Geelvink Bay, 16 February 1963, R. Straatman. Carinoscolia lorentzi (Cameron) Morobe District, Terr. New Guinea: 1 o’, Wau, 1050 meters, 18 December 1961, J. and J. H. Sedlacek. 1 o, Lae, 26 November 1961, J. Sedlacek. 1 o, Finschhafen, Huon Peninsula, 150 meters, 15 April 1963, J. Sedlacek. Carinoscolia foveifrons (Cameron) Bismarck Archipelago: 1 @, Illugi, Upper Warangoi, Gazelle Peninsula, New Britain, 230 meters, 8-11 December 1962, J. Sedlacek. 1 2, Lelet Plateau, Schleinitz Mts., New Ireland, October 1959, W. W. Brandt. Solomon Islands: 1 &, Ulo Crater, Vella Lavella Island, 10 meters, December 1963, P. Shanahan. 1 o, Dala, Malaita Island, 50 meters, 22 June 1964, J. and M. Sedlacek, in Malaise trap. 1 o, Betikama River, Guadalcanal, August 1960, W. W. Brandt. The present records extend the range of this species considerably to the south in the Solomons, the only previous record from that group being from Bougainville. 16 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 125 Austroscolia pulchripennis pulchripennis (Cameron) Western Highlands District, Terr. New Guinea: 2 o, Tomba, 2450 meters, 24 May 1963, J. Sedlacek. Bismarck Archipelago: 1 9, 5 o, [llugi, Upper Warangoi, Gazelle Peninsula, New Britain, 230 meters, 8-15 December 1962, J. Sedlacek. 2 o&', Gaulim, Gazelle Penin., 140 meters, 21-27 October 1962, J. Sedlacek. 1 o, Talliligap, New Britain, 300 meters, 17-18 December 1962, J. Sedlacek. 1 o&, Lelet Plateau, Schleinitz Mts., New Ireland, October 1959, W. W. Brandt. Solomon Islands: 21 o&, Dala, northwest Malaita, 6-22 June 1964, R. Straatman, J. and M. Sedlacek, 1 in MV light trap. In my revision (Krombein, 1963, p. 638) I questioned the occurrence of this taxon in New Guinea on the basis of an old, possibly mislabeled specimen from West New Guinea. The two males recorded above from Tomba in the western part of the Territory of New Guinea definitely establish the presence of this subspecies in New Guinea as well as in the Bismarcks and on Malaita Island alone of the Solomons. Austroscolia pulchripennis franclemonti (Krombein) Morobe District, Terr. New Guinea: 1 2, 1 km north of Awelkom, Umboi Island, 600 meters, 21-28 February 1967, 5. L. and G. A. Samuelson. Solomon Islands: 2 #, Kukugai, Bougainville, 150 meters, October and December 1960, W. W. Brandt. 1 2, 4 o&, Gizo Island, New Georgia Group, 30-120 meters, 16-26 April and 19 July 1964, J. and M. Sedlacek, 9 in Malaise trap. 1 o&, Iriri, Kolombangara Island, 2 meters, 29 June 1964, J. and M. Sedlacek. 2 o&, Kolotuve, Santa Ysabel, 21 June 1960, C. W. O’Brien. 1 o&, Betikama River, Guadal- canal, August 1960, W. W. Brandt. The female from Umboi Island is only 24 mm long. The smaller male is only 17 mm long. Both of these are smaller by a millimeter or two than the previously recorded minimal length. Austroscolia betremianus (Krombein) Bismarck Archipelago: 1 9, 3 o, Mt. Sinewit, Gazelle Peninsula, New Britain, 900-1200 meters, 5-16 November 1962, J. Sedlacek. 1 &@, Upper Warangoi, Gazelle Penin., 250-600 meters, 28-30 Novem- ber 1962, J. Sedlacek. 1 , Ilugi, Upper Warangoi, 230 meters, 12-15 December 1962, J. Sedlacek. 2 9, 2 o, Lelet Plateau, Schleinitz Mts., New Ireland, October 1959, W. W. Brandt. The present series establishes the range of length in females as 20-25 mm. NO. 3659 SCOLIIDAE—KROMBEIN ily Austroscolia nitida nitida (Smith) Northern District, Papua: 1 2, 1 o&@, Popondetta, 5-60 meters, June 1966 and 26 September 1963, P. Shanahan, Gressitt and Tawi, o in Malaise trap. Central District, Papua: 1 &, Port Moresby, 6-7 November 1960, J. L. Gressitt, in Malaise trap. Morobe District, Terr. New Guinea: 1 o, 35 km south of Lae, 120 meters, 6 August 1964, J. and M. Sedlacek. 1 &, Markham River, 60 meters, 8 August 1964, J. Sedlacek, in Malaise trap. 9 9, 36 o', Wau, 650-2400 meters, 9-12 January 1962, 15, 21-25, and 26 January 1963, 3 and 27-31 March 1964, 2 April 1966, 16-20 April 1962, 14 and 18 May 1965, 12-14 June 1963, 15 and 29 August 1961, 2, 10, 12, 17, 19, and 30 September 1961, 16-18 September 1964, 4 and 5 October 1962, 12 October 1965, 19 and 30 October 1961, 6-7 November 1961, 3 December 1962, and 18 December 1961, J., J. H., and M. Sedlacek, C. Monteith, P. Shanahan, and a native, several in Malaise trap, 3 on lucerne, 1 in light trap. 2 9, 10 @#, Mt. Misim, Wau, 950-1600 meters, 13 and 16 February 1968, 14 August and 25 September 1964, J. and M. Sedlacek, 7 @ in Malaise trap. 1 9, Kunai Creek, Wau, 1500 meters, 28-30 May 1963, P. Shanahan. 1 9, 11.2 km north of Bulolo, 600 meters, 11 March 1962, J. H. Sedlacek. 1 o&@, Adelbert Mts., 800-1000 meters, 25 October 1958, J. L. Gressitt. Eastern Highlands District, Terr. New Guinea: 1 as at ( ? - TA’ — ) id a m, mana de Ade rh a’ 7 My ; i - aan fl ie? Sy Aeensk: Pqutiaiit $4 vee ie Gra Ae : ry =i : = Jor : Se " 7 mi cay Chitin vd 79; SPOS MVCN 1 aan ar Mueqp ie ’ noah 4 an. : iS er a a re A _ > ee Aer 4 Vide E sai) 0 (wey =e a PIGS er ealy TT AS) AeA Te) I) heise i . 44 a TiAe @&'s 4g. ie y aid | @itha i) owe ut % a) rt a od he @arnades jhiieMas ui com, erred er reins Vitee hone irk Los yy 3 oie, ron dahy TR itn = Nea a aver wile Ga’ agvin(ttan, 7” Proceedings of mie. nd. ed, 5.t.a.te8 National Museum SMITHSONIAN INSTITUTION + WASHINGTON, D.C. Volume 125 1968 Number 3663 BREDIN-ARCHBOLD-SMITHSONIAN BIOLOGICAL SURVEY OF DOMINICA! 8. The Intertidal Balanomorph Cirripedia By Arnotp Ross? The present study on the intertidal balanomorph barnacles of Dominica is based on incidental collections made during April and May of 1966 by Dr. Ernst Kirsteuer of the American Museum of Natural History and by Dr. Klaus Riitzler of the Smithsonian Institution. These collections are of considerable value largely because there have been no previous studies or mention of the Cirripedia of Dominica. Represented in the collections are: Chthamalus angusti- tergum Pilsbry, 1916; Yetraclita (Tetraclita) stalactifera (Lamarck), 1818; Tetraclita (Tetraclitella) divisa Nilsson-Cantell, 1921; and Balanus (Megabalanus) stultus Darwin, 1854. That this list does not embrace all the barnacles inhabiting this region is recognized, but it probably includes the more common species. All the Dominica species have been reported previously from other regions in the Caribbean and western Atlantic with the exception 1See list at end of paper. 2 Department of Living Invertebrates, American Museum of Natural History, New York, N.Y. 10024. Present address: Department of Invertebrate Paleon- tology, Natural History Museum, San Diego, Calif. 92112. 1 De PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 of T. divisa; however, these barnacles remain poorly known because earlier workers failed to provide detailed descriptions and illustrations of their distinctive features, especially of the mouth field and cirral appendages. The addition of such information, provided herein for the Dominica fauna, should aid materially in an understanding of the morphological and biological characters of other populations of the same species as well as closely related species in the Caribbean and elsewhere. The dissected specimens, which have been figured, are deposited in the collections of the American Museum of Natural History (AMNH). Representative samples of all the species have been placed in the collections of the American Museum and the U.S. National | Museum (USNM). | The author is indebted to Drs. Ernst Kirsteuer and Klaus Riitzler, who made these collections available for study. Dr. Wiliam K. Emerson of the American Museum kindly read a manuscript draft of this paper and offered several suggestions for improvement. Dr. William A. Newman, Scripps Institution of Oceanography, first brought to the writer’s attention the fact that Balanus stultus was a megabalanid. The writer should also like to thank Dr. Newman for many informative discussions on the systematics, evolution, and phylogeny of the Cirripedia. Family CHTHAMALIDAE Darwin, 1854 Genus Chthamalus Ranzani, 1817 Chthamalus angustitergum Pilsbry Figure 1 Chthamalus stellatus angustitergum Pilsbry, 1916, p. 305, text-figs. 85, 86, pl. 71, figs. 5, 5a, 5b; 1927, p. 37, fig. 1—Nilsson-Cantell, 1933, p. 506.—Kolosvary, 1939, p. 161, figs. 6-1, 6-2; 1941, p. 68, fig. lsa.—Stephensen and Stephensen, 1950, p. 389; 1954, p. 80.—Henry, 1954, p. 444.—Wells, 1966, p. 92. Chthamalus stellatus—Smith, Williams, and Davis, 1950, p. 134.—Marshall, 1953, p. 435.—Voss and Voss, 1960, p. 102.— Werner, 1967, p. 70. Chthamalus angustitergum.—Newell, Imbrie, Purdy, and Thurber, 1959, p. 209. Mareriau.—Western side of Panto Hole Bay, east of town of Marigot, approximately 15°32’21”N, 61°17’31"W; intertidal, on Tetraclita (Tetraclita) stalactifera; May 1-10, 1966; about 100 specimens. Diaenosis.—Articular ridge of scutum straight to slightly convex, basal end of which evenly rounded and not projecting beyond the basitergal angle. Tergum narrow, about twice as high as broad, and nearly twice as thick as scutum. Mandible quadridentoid, with basal NO. 3663 CIRRIPEDIA—ROSS ay comb containing 22-28 teeth. Maxilla I with subapical notch, below which spines divided into two distinct clusters. SUPPLEMENTARY DESCRIPTION.—In the majority of the specimens examined, prominent radial ribs or their remnants ornament the parietes. The shell is pale grey as is the outer surface of the operculum. Internally, near the apices, the opercular plates are tinted pale pink. Measurements (in mm) of the five specimens dissected are as follows: shell opercular plates carino-rostraé specimen diameter lateral diameter height scutum height tergum height i 5.8 5. Pel 1s 1.6 2 6.1 6.0 745°C 1.8 1.9 3 8.0 8.0 ayal 155) 1.6 4 Oe 4.9 2.0 1.2 16.3} 5 Sy 5.4 5 JL Vhs i 1.2 The height of the scutum is about one-half its width. The articular ridge extends about three-fifths the length of the tergal margin. It is straight, and the basal margin is evenly rounded and does not project. There is no adductor ridge; the adductor muscle depression is very small and deep. Situated on the basitergal angle, which pro- jects beyond the tergal margin, is the pit for the depressor muscle. The tergum is about one-half as wide as it is high, and the plate is exceedingly thick. There is a small, narrow, and clearly delimited spur. Crests for the tergal depressor muscle number five to seven. The multidenticulate labrum has a shallow, medial, saddle-like depression, on each side of which are 11-14 teeth. Short setae are dispersed between the teeth along the crest of the labrum (fig. If). A large elongate-oval patch of downward-directed spines extends obliquely from each side of the superolateral margins toward the basimedial portion of the labrum. The superior margin of the palp is straight and free of setae (fig. Ic). The basal margin is covered with short setae, but those on the supero- distal margin are extremely long; all of the setae are clothed with spinules. The cutting edge of the mandible is armed with five teeth including the inferior angle (fig. 1a). The distance between the second and third teeth is one-half that between the first and second teeth. The fourth tooth is bifid. The inferior angle bears three to four spines, above which is the typically developed comb, containing about 22-28 spinules. Maxilla I has a somewhat trilobed cutting edge. There is a promi- nent subapical, V-shaped notch, and on some specimens there is a suprabasal notch. The spination along the edge is readily separable into three zones, the apical consisting of both long and short, stout 4 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 spines; the medial, below the subapical notch, containing long, slender spines; and the basal, which may or may not be set off from the medial by a slight notch, containing only short, slender spines (fig. 15). Ficure 1.—Chihamalus angustitergum Pilsbry, 1916, AMNH 12458, Panto Hole Bay, Dominica: a, mandible; b, maxilla I; c, palp; d, maxilla IT; ¢, cirrus I; f, labrum; g, distal end of penis; h, cirrus II; 7, comb seta from cirrus II; 7, cirrus III; &, cirrus VI. (Setae omitted on cirri I and II; on numbered segments of cirri III and VI, only one set of setae shown; a=anterior ramus; p=posterior ramus; scale in millimeters.) NO. 3663 CIRRIPEDIA—ROSS a Maxilla IT has a distinctly bilobed anterior margin. The appendage is broad, but the height is somewhat greater. The anterior ramus of cirrus I is slightly longer than the posterior ramus, and all of the articles of both rami are broader than they are high. The terminal article of the posterior ramus bears a few comb setae. The same relationships as found in cirrus I hold for cirrus II except that comb setae are found on the terminal articles of both rami. The comb setae on both cirri lack basal guards. Cirrus III is not modified. Cirri III-VI are essentially equal in length with equal rami. At each articulation along the posterior curvature of the inter- mediate articles of cirri [II-VI there are 1-3 long, slender setae and 1-3 short, slender setae. There are no bristles or teeth arrayed along the lateral faces of the posterior cirri. Pigment spots occur at each articulation along the posterior border of the cirri and along the whole length of the anterior border of the cirri. Chaetotaxis of the anterior border of the intermediate articles of cirri IV-VI is typically ctenopod, there being 5 pairs. A count of the segments of specimen number 1, illustrated in figure 1, is as follows: I I UI IV Vv VI anterior 8 6 14. We? 19 19 right ; ef a ray a oor ars posterior 6 6 15 17 19 19 anterior 7 if is 18 19 19 left = = a ais ar Bo posterior 6 if 15 XX 19 19 Data on the cirral counts of the five specimens dissected are pre- sented below. The range (R) and mean (X) values for the number of segments in the anterior (a) and posterior (p) rami of the right side are as follows: ] II III I\ Ny VI 1 fain. et ee haat Silas amped” aiid adic ae Seana heals Gn) rele SG aes aa aa 5 5 5 5 5 5 5 R 6-8 5-6 6-9 5-7 14-15 14-16 13-17 15-17 16-19 16-19 17-19 17-19 AO.600) v4 6.0) 14/5 1570° 154)” 16-0 ° ‘1747 1678) 18.0 1746 The penis is annulated throughout its length, and it is covered with long bristles. The distal extremity is densely clothed with bristles. There is no basidorsal point. Remarxs.—The nominal subspecies Chthamalus stellatus angusti- tergum is readily separable from C. stellatus stellatus (Poli, 1791) on taxonomic as well as geographic and reproductive grounds. Al- though apparently alike superficially, differences exist that clearly warrant the recognition of this Caribbean barnacle at the specific level. 6 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 Although there are notable differences in the coloration of these two species, by far the greatest number of distinguishing characters are found in the opercular plates. The scutum of the Caribbean species is about one-half as tall as it is wide, whereas in C. stellatus it is about two-thirds. In the latter species the adductor muscle pit is exceedingly large and the basal end of the articular ridge is acute, extending out to the basitergal angle; the opposite conditions exist in C. angustitergum. In C. angustitergum, furthermore, the tergum is narrow—hence, the name—the width being about one-half the height and the spur being well set off from the basiscutal angle, but in C. stellatus the spur is virtually confluent with the basiscutal angle and the height of the plate exceeds the width by one-fourth or less. Crests for the tergal depressor muscles, of which there are four in CO. stellatus, are low and feebly developed. In C. angustitergum there are five crests, which are both high and strongly developed. Insular populations of C. angustitergum in the Caribbean are obviously isolated geographically from the Asia Minor, Mediter- ranean, and eastern Atlantic populations of C. stellatus—indeed, there is no indication of recruitment of C. stellatus in the Caribbean; consequently, the populations of these two species are reproductively isolated and more than likely have been for a significantly long period. Such isolation, both reproductive and spatial, speaks in favor of eliminating the antequated subspecific label attached to C. angustitergum. There are at least three other species of Chthamalus in the Caribbean and western Atlantic: C. fragilis Darwin, 1854; C. stellatus thompsont Henry, 1958; and ©. rhizophorae de Oliveira, 1940. There should be little confusion regarding the identity and distinction among these species. Ohthamalus fragilis is confined for the most part to the Atlantic coast of the United States, although it has been reported in the Caribbean and on the west African coast (Stubbings, 1967, p- 262). Bermuda is apparently the only locality where C. stellatus thompsoni occurs (Henry, 1958, p. 220). The euraphian, C. rhizophorae, has been collected by the present writer at several localities in the Bahamas, although previously it was known only from Brazil (Oliveira, 1940b, p. 379; Stubbings, 1967, p. 257). Family TETRAcLITIDAE Nilsson-Cantell, new status TerRacLitina® Nilsson-Cantell, 1921, p. 357. At the present time, students of balanomorph systematics favor inclusion of the tetraclitids in the family Balanidae. Although Nilsson- Cantell (1921) took the first and only bold step when he segregated the two dozen or so species of this complex into a distinct subfamily, NO. 3683 CIRRIPEDIA—ROSS 6 there apparently has been no further attempt to reevaluate the systematic position of these barnacles. Assignment of the tetraclitids as a subfamily of the Balanidae seems untenable at this time because they obviously represent a different and distinct lineage that in many respects parallels closely that of the Chthamalidae. The tetraclitids are compounded of forms with a shell consisting of only four tubiferous plates, the number of tubes ranging from one row in Tesseropora to several rows in Tetraclita and Tetraclitella. In the latter group the radii are also tubiferous, and only in this group are the radi as well developed as they are in the Balanidae. The basis with few exceptions is membraneous, and where calcareous, it is neither tubiferous nor does it possess complex ridges marginally that interdigitate with the basal margin of the wall. One of the salient structures of primary importance in the classi- fication of the Balanomorpha is the state of development of the labrum. In the Chthamalidae there is no notch in the labrum, and it is effectively bullate, whereas in the Balanidae the labrum is dis- tinctly notched and not bullate. In the tetraclitids this structure is clearly not bullate, nor is it notched, but rather it holds an inter- mediate position since it is thin, as in the balanids, and the crest has a saddle-like groove, as in Chthamalus. The presence of a bullate or simple un-notched labrum indicates a primitive evolutionary state, which is readily recognizable in the chthamalids and, of the other Thoracica, in the verrucomorphs and lepadomorphs. On the basis of the foregoing evidence alone it is not likely that an un-notched labrum could have evolved from a notched labrum; hence, the tetra- clitids could not have evolved from the balanids. Modifications in tetraclitid mandibular structure closely parallel that found in the chthamalids, wherein there are groups with either a comblike or serrate inferior margin. The inferior margin is never molariform as is frequently the case in the Balanidae. The first three pairs of cirri in the Balanidae are highly modified and effectively serve as mouth appendages. In Chthamalus and related genera only the first two pairs of cirri are so modified. In the tetra- clitids, cirrus III is modified as a mouth appendage but significantly less so than in the Balanidae. In the Balanidae the third cirrus is never antenniform, but it commonly is in the Chthamalidae and rarely in tetraclitids of the Tetraclita squamosa complex. Other features that argue for exclusion of the tetraclitids from the Balanidae are the absence of a basidorsal point on the intromittant organ and the presence of comb setae on the anterior cirri. Caudal appendages, present in many of the chthamalids, are lacking in the tetraclitids as well as the balanids. 8 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 In all likelihood the tetraclitids and chthamalids shared a common ancestry, but unlike the chthamalids the tetraclitids probably evolved and deployed far more rapidly. The balanids, on the other hand, probably evolved from the chthamalid lineage, but at a much later time. Unfortunately, the fossil evidence documenting these evolu- tionary steps remains to be discovered. Inclusion of the tetraclitids in the Balanidae would clearly appear to be precluded on the basis of the foregoing. By the same token their inclusion in the Chthamalidae would serve only to weaken the definition of this group also. Because no intergrades or transitionary forms are known to occur between the Balanidae and tetraclitids, separation of these two groups at the familial level is clearly warranted. Genus Tetraclita Schumacher, 1817 Tetraclita (Tetraclita) stalactifera (Lamarck) FIGURE 2 Balanus ponderosus [Lightfoot, 1786,] p. 89.—Dillwyn, 1825, p. 23. Balanus stalactiferus Lamarck, 1818, p. 894.—-Lamy and André, 1932, p. 222. Balanus latus Lamarck, 1818, p. 397.—Lamy and André, 1932, p. 222. C[lonia] stalactifera.—Chenu, 1843, pl. 4, figs. 6-8. Tetraclita porosa var. communis Darwin, 1854, in part, p. 329, pl. 10, fig. la, li (?), 1k (?). Tetraclita porosa.—Verrill, 1901, p. 22.—Bigelow, 1901, p. 180. Tetraclita squamosa stalactifera.—FPilsbry, 1916, p. 254, pl. 59, figs. 1, la, 1b.— Oliveira, 1940a, p. 138; 1941, p. 7, pl. 1, figs. 1, 2, pl. 2 (fig. 4), pl. 10 (figs. 1, 3, 6); 1947, p. 715.—Stephensen and Stephensen, 1950, p. 388; 1952, p. 8.— Henry, 1954, p. 444; 1958, p. 224, pl. 1, fig. b, pl. 5, figs. a, b.—Newell, Imbrie, Purdy, and Thurber, 1959, p. 209.—Voss and Voss, 1960, p. 102, 106.— Ross, 1962, p. 31.—Rehder, 1967, p. 16. Tetraclita stalactifera.—Pilsbry, 1927, p. 38. Tetraclita porosa stalactifera—Nilsson-Cantell, 1933, p. 508; 1939, p. 5. Tetracliia squamosa.—Smith, Williams, and Davis, 1950, p. 184.—Marshall, 1953, p. 435.—Werner, 1967, p. 70.—Multer and Milliman, 1967, p. 260, fig. 2. MarTERIAL.—Scotts Head Bay, at southern end of Soufriere Bay, approximately 15°12’40” N, 61°22’40” W; 0.5 meters; May 17-28, 1966; about 150 specimens. Middle Bay, adjacent to town of Marigot, approximately 15°32’21” N, 61°17’28” W; intertidal, on basaltic rocks; April 15-30, 1966; 4 specimens; same locality as that cited by Kirsteuer (1967). Western end of Panto Hole Bay, approximately 15°32’21" N, 61°17/31" W; intertidal, on basaltic rocks; May 1-10, 1966; about 125 specimens. Dracnosis.—Sheath of shell tinted plumbeous black and opercular plates violet black or raisin black with whitish or paler borders and ridges. Parietes relatively thin; tubes large, commonly in 3-5 rows. Radii very poorly developed or obsolete. Labrum with slight notch NO. 3663 CIRRIPEDIA—ROSS 9 and 4 teeth on each side of notch. Mandible with 6 teeth including inferior angle; second and third teeth bifid; fifth tooth rudimentary; inferior angle strongly pectinate. Maxilla I with 2 spines above Ficure 2.—Tetraclita stalactifera (Lamarck), 1818, AMNH 12459, Scotts Head Bay, Dominica: a, mandible; b, enlarged view of fourth tooth and inferior angle of mandible shown in a; c, maxilla I; d, maxilla I; ¢, palp;f, cirrus 1; g, labrum; h, distal end of penis; 2, comb seta from cirrus II; 7, cirrus II; &, cirrus III; /, cirrus VI. (Setae omitted on cirri I-III; on numbered segments of cirrus VI, only one sct of setae shown; a=anterior ramus; Pp=posterior ramus; scale in millimeters.) 288-726—68 2) -_ 10 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 subapical notch, and 18-24 spines beneath, unequally divided into 2 clusters. Anterior ramus of cirrus [ longer than rami of cirri IT and III. Comb setae of cirri IT and III lacking basal guards. SUPPLEMENTARY DESCRIPTION.—The shell is_ typically conic, spreading, commonly eroded externally, and tinted various shades of white with deep purple. The radii are extremely narrow or obsolete, and often external demarcation of the four parietal plates is virtually impossible to recognize. The surface of the body cavity wall is smooth or slightly irregular but never ribbed or ridged. Externally, the surface of the scutum is eroded, as is the tergum. The adductor ridge of the scutum extends nearly the whole length of the valve. It is long, low, deeply undercut, and the ridge moderately thin. The articular ridge occupies about two-thirds the length of the tergal margin, and it is evenly and broadly rounded distally. Prominent ridges exist for the insertion of the lateral and rostral depressor muscles. Dentition of the occludent margin is limited to 4-6 strong, oblique teeth. The number and development of these teeth, however, is no doubt contingent upon the age and degree of corrosion of the specimen. The apex of the tergum is produced into a slight but noticeable beak. The articular ridge is high, and its thickness varies. There are 6-8 strong, well-spaced, inclined crests for the insertion of the depressor muscles. Parameters of the shell and opercular plates of the five dissected specimens (in mm) are as follows: shell opercular plates carino-rostral specimen diameler lateral diameter height seutum height tergum height J 19.5 17.2 13.4 OL 6.0 2 ieyers 1726 122 50.4 6.5 3 20.6 18.6 12.9 6.1 40 t 30.8 21.2 16.2 (8 8.6 5 19 17.6 4.7 2.9 2.8 On both sides of the shallow, saddle-like groove in the crest of the labrum there are four short teeth, although in one specimen dissected there were four teeth on one side and only three on the other. Inter- spersed among the teeth in the notch and extending laterally along the rim of the labrum are short, slender bristles (fig. 2g). Two elongate- oval patches of spines extend obliquely from the superolateral margin on each side of the crest toward the basicentral portion of the labrum. The palps are long, high, and free of setae along their basal margins; however, the basal portion of the appendage is uniformly covered with ctenoid scales (fig. 2e). The setae on the distal extremity are long and setulose whereas those on the superior margin are shorter and not covered with setules. NO. 3663 CIRRIPEDIA—ROSS if Along the cutting edge of the mandible there are six teeth including the inferior angle (fig. 2a). The distance between teeth decreases by about one-half, proceeding from the first tooth to the fifth. Both the second and third teeth have subsidiary cusps. Along the superior slope of the third and fourth teeth there are serrations. Similarly, the inferior angle is serrate, but very coarsely. Maxilla I has a deep subapical notch (fig. 2c). Above the notch there are two long, stout spines and one or two shorter, slender spines. Below the notch 11-16 long, slender spines and 8-14 shorter, slender spines are present. On larger and older specimens there may be little distinction between these two spine clusters. Maxilla IT is taller than broad and distinctly bilobed. The setae on the basal lobe are setulated. The rami of cirrus I are grossly unequal in length, the posterior ramus being about one-half the length of the anterior ramus (fig. 2/). The rami of cirrus IT are essentially equal in length while the posterior ramus of cirrus III is slightly longer than the anterior ramus. The intermediate and basal articles of the anterior three pairs of cirri are squat, not appreciably protuberant. Both rami of cirrus IT and III are clothed with long comb setae, which lack basal guards. Cirri IV-VI are essentially equal in length with equal rami. At each articu- lation along the posterior curvature of the intermediate articles of cirri [V—VI there is one short, stout seta, 2-4 long, slender setae, and 1-2 short, slender setae. On the lateral face of both rami of cirri IV-VI immediately below each articulation of the intermediate articles, there is a single row of short spines. Chaetotaxis of the inter- mediate articles of the posterior three pairs of cirri, along the anterior curvature, is ctenopod, there being three pairs of setae. Between or at the bases of the distal one or two pairs of setae there are one or two short slender setae. A count of the segments of the cirri of specimen 1 above, illustrated in figure 2, is as follows: I II Ii LV V VI anterior ramus 22 wn 13 19 21 23 right Sh OBR om posterior ramus 12 il 20 22 25 26 anterior ramus 23 11 12 18 22 2) left Sar ase wes ee par fe posterior ramus 12 10 18 21 24 21 Counts of the cirral segments are summarized below for the five specimens dissected. The range (R) and mean (x) values for the number of segments in the anterior (a) and posterior (p) rami of the cirri of the right side are as follows: 12 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 125 if II IIL IV V VI a aoe a p a Dp a p a p a Dp nD o 5 5 5 5 5 5 5 5 5 5 5 XY 15-26 8-17 9-14 8-13 8-14 11-20 16-21 16-22 18-24 18-26 19-26 19-26 x 20540 11-8310. SelOcSe We aiie2) 1S 58" 208610 2100) 230g Rae oe. The intromittant organ is distinctly annulated throughout its length. It is sparsely hirsute except for the tip where there are two or three distinct clusters of setae. There is no basidorsal point. Remarks.—In a recent study Rehder (1967) listed the valid zoological names from an anonymously authored auction catalogue of the Portland Museum collections. Of the names cited in this catalogue, which is purported to have been authored by the Reverend John Lightfoot, only one, Balanus ponderosus, is of concern here. in listing this name, Lightfoot referred to a published figure of Lister’s (1770, pl. 442, fig. 284). The species represented by that figure most closely resembles Tetraclita stalactifera of all the pres- ently known Caribbean barnacles. Although this illustration of a specimen from Barbados is poorly executed, Rehder (1967, p. 17) stated that, ‘“The form, sculpture, and locality cited under Lister’s figure seem to point to this tropical Western Atlantic barnacle but because Lightfoot’s name has never been used for this subspecies, it should be considered a nomen oblitum.” In concurring with Rehder, it need only be added that Lamarck’s species should be conserved because of its widespread and common usage for 150 years, whereas Lightfoot’s name, as noted by Rehder, has never appeared in the primary literature. Passing mention should be made of the species Balanus latus Lamarck (1818, p. 397). This name was also based on the same figure as that of B. ponderosus. Under these circumstances it is obviously not amiss to include this name in the synonymy of 7. stalactifera. Tetraclita stalactifera ranges from South Carolina, through the Caribbean to southern Brazil (Stephensen and Stephensen, 1952; Pilsbry, 1916). It also has been reported from Bermuda (Verrill, 1901; Henry, 1958). Its range in the eastern Pacific is from the Gulf of California to Acapulco, Mexico. The presence of this species in the eastern Pacific, although over a considerably restricted range, indicates it is a relatively old species, probably ranging as far back as the Miocene. It is readily separable from all other forms of Tetraclita on the basis of its geologic history, distribution, color, shell and trophic morphology, and cirral counts; NO. 3663 CIRRIPEDIA—ROSS 13 | consequently, it is regarded herein as a full species and cited accordingly. Subgenus Tetraclitella Hiro, 1939 Tetraclita (Tetraclitella) divisa Nilsson-Canteil Tetraclita divisa Nilsson-Cantell, 1921, pp. 93, 362, text-figs. 8, 83, pl. 3 (fig. 11).— | Visscher, 1927, p. 201, fig. 68.—Hiro, 1939, p. 275, fig. 15.—Zevina and Tarasov, 1963, p. 96, fig. 13.—Stubbings, 1967, p. 291, fig. 21. Tetraclita (Tetraclitella) divisa subquadrata Ross, 1961, p. 210, pl. 4, fig. 1-5. Tetraclita costata.—Pilsbry, 1928, p. 316. MarTeriAu.—Western side of Panto Hole Bay, east of town of Marigot, approximately 15°32’21” N, 61°17/31” W; intertidal, on Tetraclita stalactifera; May 1-10, 1966; 2 specimens. Draenosis.—Shell covered by persistent, hirsute, chitinous mem- brane; parietes furnished with prominent, radial ribs. Radii with horizontal summits parallel to base; external surface covered with horizontal ridges scored at regular, close-spaced intervals. Scutum broader than high; adductor ridge of variable degrees of prominence, and apically fused with articular ridge to effectively form roof. Tergum only slightly higher than wide; both terga form about one- third of bulk of operculum; spur extremely broad and separated from basiscutal angle by less than one-half its own width. Few low denticles on both sides of shallow, saddle-shaped groove in labrum. Cutting edge of mandible armed with 5 teeth including strongly serrate inferior angle, not furnished with comb. SUPPLEMENTARY DESCRIPTION.—The small size, peltate form; regu- larity of shell ornamentation and outline, and the pale purple or whitish-purple shell of this species should serve as readily recognizable features for field identification. Measurements (in mm) of the two specimens in the present col- lections are as follows: shell opercular plates carino-rostral specimen diameter lateral diameter height scutum height tergum height 1 feel Ileal 3.9 4.2 4.0 2 8.2 7.6 2.6 3.7 3.0 In view of the excellent available descriptions of this species by Nilsson-Cantell (1921), Hiro (1939), Zevina and Tarasov (1963), and Stubbings (1967), nothing more need be added at this time; however, the specimens were dissected, and cirral counts are given below for each of them: 14 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 I II III IV Vv VI i anterior 9 6 6 fal 15 15 right a = a cee as aw posterior 5 5 5 12 15 15 specimen 1 anterior a 6 5 Wil 14 15 left . oa i ire = an as posterior 5 5 4. 13 14 15 ; anterior % 6 6 12 14. 15 right me ac — aes — posterior 5 5 5 12 14 15 specimen 2 anterior 9 cf 5 11 te 15 left ; He, 7% i es ar foig posterior 5 6 4 13 14 15 In the larger of the two dissected specimens there were about 45 embryos. In the smaller specimen there were 32 cyprids. The latter have been figured by Nilsson-Cantell (1921, text-fig. 8). REMARKS.—Based on specimens collected by A. E. Verrill in the Hawaiian Islands the present writer described 7. divisa subquadrata (Ross, 1961, p. 210). After critical reexamination of the type speci- mens the author has concluded that this subspecies does not warrant recognition, and should be considered a synonym of the nominate species. Nilsson-Cantell (1921, p. 364) described 7. divisa from specimens collected in Sumatra. Subsequent workers have reported it from Formosa (Hiro, 1939, p. 275), the South China Sea (Zevina and Tarasov, 1963, p. 97), and Ghana on the west coast of Africa (Stub- bings, 1967, p. 293). The presence of this species in the Caribbean and the Hawaiian Islands extends greatly its known distribution. The fact that it has virtually a circumtropical distribution tends to confirm the long history of members of the Tetraclitidae. Family BALANIDAE Leach, 1817 Genus Balanus DaCosta, 1778 Subgenus Megabalanus Hoek, 1913 Balanus (Megabalanus) stultus Darwin FIGURE 3 Balanus stultus Darwin, 1854, p. 216, pl. 3, figs. 2a-—2d.— Weltner, 1897, p. 262.— Gruvel, 1905, p. 221, fig. 243.—Pilsbry, 1916, p. 235.—Nilsson-Cantell, 1929, p. 1, figs. 1c, 1d, 2; 1939, p. 5.—Pilsbry, 1953, p. 25, pl. 2 Gigs) J=3).—— Henry, 1954, p. 443. Tetraclita radiata.—Pilsbry, 1927, p. 38. Balanus stultus morycowae Kolosvary, 1966, p. 69, pls. 1, 2. NO. 3663 CIRRIPEDIA—ROSS 15 Marerrau.—Scotts Head Bay, at southern end of Soufriere Bay; approximately 15°12’40” N, 61°22’40” W; 0.5 meters; May 17-28, 1966; 5 living and 2 dead specimens on Millepora complanata Lamarck; locality no. 2 of Kier (1966). Diaenosis.—Basis, radii, and parieties tubiferous. Basal margin of scutum sinuous and strongly protuberant medially; growth ridges of plate straight and narrow at lateral extremities, but flexed basally in medial third. Labrum with 3 teeth on each side of medial notch. Mandible with 5 teeth including inferior angle; second tooth bifid. Cutting edge of maxilla I straignt. Living strictly on milleporine corals. SUPPLEMENTARY DESCRIPTION.—This barnacle is readily recognized by its habitat, although the shell is commonly overgrown by the milleporine. In all of the present specimens the shell is high, conical, the parietes slightly to strongly ribbed, and the basis flat. The radii are moderately broad with summits parallel to the basis. The sutural surface of the radii and opposed sutural surfaces possess septa on which both the superior and inferior margins are denticulate. Shell and opercular dimensions (in mm) of four dissected specimens are as follows: shell opercular plates specimen diameter lateral diameter height height scutum height tergum 1 23.8 18.9 14. 0 5.9 6.9 2 19.1 15.0 ie}, ag2 6.1 3 2050 18.7 Hoe A 6.2 Uo ® 4 23.4 15.9 NG 7 6.8 Cell Both the scutum and tergum are tinted apically, either pink or pale purplish blue. The articular ridge of the scutum is about two- thirds the length of the articular margin, and it terminates in an acute point. In young stages the adductor ridge is prominent, acute, and it extends nearly to the midpoint of the basal margin, but in older specimens it is less prominent and less broadly rounded. The adductor muscle depression is deep, and its outline varies from elongate-oval to oval, to nearly circular. The tergum is broad, triangular, and thinner than the scutum. The spur is separated by about its own width from the basiscutal angle. Externally, the longitudinal furrow is commonly closed but sometimes open. Crests for the attachment of the tergal depressor muscles are weakly developed, and few in number. On each side of the deeply notched crest of the labrum there are three teeth (fig. 3d). Interspersed among the teeth and extending laterally the complete length of the crest, there are short, soft setae. Extending obliquely from the superolateral margin on each side of the crest toward the basicentral portion of the labrum, there are 16 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 125 large, elongate-oval patches of spines directed downward toward the esophagus. The palps are long, high, and spatulate. Along the superior margin the setae are moderately long and densely concentrated, whereas Ficure 3.—Balanus (Megabalanus) stultus Darwin, 1854, AMNH 12460, Scotts Hea: Bay, Dominica: a, mandible; b, maxilla I; ¢c, palp; d, labrum; e, maxilla II; f, cirrus I g, cirrus II; h, cirrus I; 7, distal end of penis; j, cirrus VI. (Setae omitted on cirri I-III on numbered segments of cirrus VI, only one set of setae shown; a=anterior ramus p=posterior ramus; scale in millimeters.) NO. 3663 CIRRIPEDIA—ROSS 1 7 at the distal extremity they are significantly longer and more sparsely concentrated. A single row of long setae occurs on the outer lateral face, above and oblique to the basal margin of the appendage. Five teeth, including the inferior angle, occur along the cutting edge of the mandible (fig. 3a). The distance between the teeth de- creases by about one-half proceeding from the first tooth to the inferior angle. The inferior angle is simple, not molariform, and does not project forward beyond the fourth tooth. The second and occa- sionally the third tooth is bifid, or it bears subsidiary cusps. Maxilla I has a straight cutting edge, although on one specimen dissected there appears to be a minute notch above the center. The spines are separable into three poorly defined groups, the apical consisting of two long, stout spines, a central group of 10-14 spines of varying lengths and diameters, and a basal cluster of about 7-12 short, very slender spines. Mazxilla II is tall, slender, and the anterior edge distinctly bilobed (fig. 3e). At the juncture of the two lobes there is a large patch of short, stiff bristles. The anterior ramus of cirrus I is slightly longer than the posterior ramus, the intermediate articles of which are strongly protuberant, whereas those of the anterior ramus are less so (fig. 3f). The anterior ramus of cirri II and III are longer than the posterior. The inter- mediate articles of both rami of cirrus II are strongly protuberant, but less so in cirrus III (figs. 3g, 4). Cirri IV-VI are essentially equal in length, with rami of more or less similar lengths. At each articu- lation along the greater curvature of the intermediate articles of cirri [V-VI there are two to three short, slender bristles, one short, stout bristle, and one to two very short, fine bristles. Immediately below each articulation of the inner lateral faces of both rami of the posterior three pair of cirri there is a row of short, triangular spines. Chaetotaxis along the anterior curvature of intermediate articles of cirri [V-VI is ctenopod, there being four pairs of setae on each article. A count of the segments for specimen 1 above (fig. 3) is as follows: I IL III IV Vv VI anterior 19 16 14 25 26 29 right ¢ a we aa aaa a aes. posterior es 12 12 31 31 28 eta ee LS ERS I RD posterior 15 14 13 31 32 ay A summary of the data on the cirri of the four specimens dissected are summarized below. Range (R) and mean (x) values for the number of segments in the anterior (a) and posterior (p) rami of the cirri of the right side are as follows: 18 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 126 T Il Iil IV Vv VI a Pp a p a Pp a Dp a 7) . a p n 4 4 4 4 4 4 4 4 4 3 2 4 R 17-21 13-18 13-16 10-13 12-15 11-14 25-29 29-31 26-32 31-35 29-32 28-34 KWo Vas MAO TAN o MSY 2N 2) 22882) “3050 29n2 S256 SOs omeo Ieee The penis is annulated throughout its length, but it is sparsely hirsute except for the distal extremity (fig. 32). The basidorsal point is small, bluntly rounded, and sparsely covered with short, stiff bristles. Remarxs.—Balanus stultus is herein assigned to the subgenus Megabalanus. Darwin (1854, p. 216), however, obviously could not reconcile the morphological evidence with the sectional (=subgenus) diagnosis he proposed for other megabalanids, and, consequently, he was forced to assign B. stultus to a different section of Balanus, subsequently regarded as the subgenus Conopea. Pilsbry (1958, p. 27) also could not accept the fact that B. stultus is a megabalanid; he stated: In Florida when I first saw these barnacles I thought they were a species of Megabalanus (the B. tintinnabulum group), but the absence of pores in the radii was against this view. Later, upon identifying them as B. stultus, I was amused to find that Darwin had been similarly embarrassed by the structure of this barnacle. The failure of earlier workers to note the obvious affinities of B. stultus stems from the fact that the tubes of the radii are not readily observed unless the radius is broken away from the parietes. Particu- larly significant, in regard to facies similarity, are the form and topog- raphy of the opercular valves and the structure of the trophic as well as the cirral appendages. It should be pointed out at this time, however, that there are at least two recognizable species groups in the mega- balanids that are based in part on certain elements of the mouth field and on certain shell and opercular characters. Some semblance of these two groups may be seen in the key to the species of Megabalanus presented by Pilsbry (1916, p. 53). Of these groups, B. stultus perhaps is allied most closely to the B. tintinnabulum group. The specimens that Pilsbry (1927, p. 38) reported as Tetraclita radiata were subsequently reexamined by Nilsson-Cantell (1939, p- 5), who found them to be specimens of B. stultus. It would also appear that the specimen Pilsbry cites and illustrates in the same paper as B. tintinnabulum antillensis, growing on Millepora alcicornis, may also prove, on closer scrutiny, to be a specimen of the present species. In a recent paper, Kolosvaéry (1966) reported the occurrence of what he believed to be a new subspecies of B. studtus from Cuba, which he called morycowae. The cardinal differences between mory- cowae and the nominate subspecies is that, in the former the adductor NO. 3663 CIRRIPEDIA—ROSS 19 muscle pit of the scutum is circular in outline and the spur faciole of the tergum is extremely narrow and apparently closed. To judge from the studies of earlier workers and from specimens on hand, it appears that the degree and length of closure of the longitudinal furrow of the tergum is highly variable and of little diagnostic value in this species. In regard to the outline of the adductor muscle de- ression, this too appears to be quite variable and no doubt dependent on the age of the specimen, among other things. Consequently, the author unhesitantly considers Kolosvdry’s taxon a junior synonym f B. stultus and not worthy of recognition. Although Darwin (1854, p. 216) reported B. stultus from Singapore las well as from the West Indies, the western Pacific record obviously is an error. Distrisution.—Balanus stultus ranges from the Dry Tortugas (Nilsson-Cantell, 1929) and Florida (Pilsbry, 1953) to Cuba (Kolos- dry, 1966) and Curagao (Pilsbry, 1927; Nilsson-Cantell, 1939). Pilsbry (1927) reported it growing on Millepora aleicornis Linnaeus, and Weltner (1897) found it to be living on M. complanata. | Summary | Four species of intertidal cirripeds, reported for the first time from the island of Dominica, West Indies, are described and illustrated. Chihamalus angustitergum and Tetraclita stalactifera are elevated to the rank of species. Tetraclita divisa is shown to be of circumtropical distribution, and 7. divisa subquadrata is accordingly placed in synonymy with the nominate subspecies. Balanus stultus is removed from the subgenus Conopea and assigned to Megabalanus on the basis of facies similarity and the presence of tubes in the radi. The subfamily Tetraclitinae is elevated to the rank of family. Literature Cited BicELow, Maurice A. 1901. The Cirripedia collected near Porto Rico by the Fish Hawk Expedition in 1898-99. Bull. U.S. Fish. Comm. for 1900, vol. 2, pp. 177-180. CHENU, J. C. 1843. Illustrations conchyliologiques, pt. 12, pl. 4 [no pagination or text]. Paris. Darwin, Charles R. 1854. A monograph on the sub-class Cirripedia, with figures of all the: species: The Balanidae, (or sessile cirripedes) ; the Verrucidae, ete., | 684 pp., 30 pls. London: Ray Society. | Ditiwyn, L. W. 1823. An index to the Historia Conchyliorum, with the name of the species to which each figure belongs, and occasional remarks, pp. 1-48. Oxford. GRUVEL, ABEL 1905. Monographie des cirrhipedes ou thecostraces, 472 pp., 427 figs. Paris: Masson et Cie. Henry, Dora P. 1954. Cirripedia: The barnacles of the Gulf of Mexico. Jn Galtsoff, ed., Gulf of Mexico: Its origin, waters, and marine life. Fish. Bull. U.S. Fish Wildl. Serv., vol. 55, no. 89, pp. 443-446. 1958. Intertidal barnacles of Bermuda. Journ. Mar. Res., vol. 17, pp. 215-234, fig. 1, pls. 1-9. Hiro, Fusio 1939. Studies on the cirripedian fauna of Japan, IV: Cirripeds of Formosa (Taiwan), with some geographical and ecological remarks on the littoral forms. Mem. Coll. Sci., Kyoto Imp. Univ., ser. B, vol. 15, no. 2, art. 8, pp. 245-284, figs. 1-16. Kier, Porter 1966. Bredin-Archbold-Smithsonian biological survey of Dominica: 1, The echinoids of Dominica. Proc. U.S. Nat. Mus., vol. 121, no. 3577, pp. 1-10, figs. 1-8, pls. 1-2. KirstEvER, ERNstT 1967. Bredin-Archbold-Smithsonian biological survey of Dominica: 3, Marine archiannelids from Dominica. Proc. U.S. Nat. Mus., vol. 123, no. 3610, pp. 1-6, fig. 1. Kotosviry, G. 1939. Uber die Variabilitat der Cirripedien-Unterart Chthamalus stellatus stellatus (Poli). Zool. Anz., vol. 127, nos. 5-6, pp. 159-169, figs. 1-8. 1941. Die Formenkreise der Chthamaliden. Zool. Anz., vol. 133, nos. 3-4, pp. 67-81, figs. 1-7. 1966. Balanus (Conopea) stulius morycowae n. ssp. (Crustacea, Cirripedia). Bull. Soe. Amis Sci. Lett. Poznan, see. D, vol. 7, pp. 69-71, pls. 1-2. Lamarck, J. B. P. A. pz M. DE 1818. Histoire naturelle des animaux sans Vertebres, vol. 5, pp. 375-410. Paris: Deterville. 20 No. 3663 CIRRIPEDIA—ROSS PA ‘Lamy, Epovarp M., anv Anpré, M. Marc | 1932. Notes sur les espéces Lamarckiennes de Cirripédes. Compt. Rend. Congr. Soc. Sav. Paris Depart., Sec. Sci., no. 65, pp. 212-228. {Lignrroot, JoHN] 1786. os ete ee) eee Midtibia without preapical spur. . . . id) 6. Terminal segment of maxillary sige elongate, rN ponent with suture-like cross striae. . .. BRP Ne ith Terminal segment of pal Roan same lenin as cprcnedine scemenel without cross striae. .. . Peo OweOer Ge 7. Mesoscutum with a pair of staal etal varie: forctibia often Tate a preapical SPUR wien .. . « 6 PSycnomyiman Mesoscutum Tisha Rotal rariae fereuibinG never ouatn a preapical spur . HyYDROPSYCHIDAE 8. Antennae much longer than wings. ....... ... . . LEPTOCERIDAE Antennae about length of forewings. . .... .. . . HELICOPSYCHIDAE 9. Mesoscutellum small and rectangular. . . . . . . . . CALAMOCERATIDAE Mesoscutellum large and domelike. . ..... =... . . ODONTOCERIDAE Family GLOSSOSOMATIDAE Although the family is found over most of the world, the Pro- toptilinae, to which all the antillean species belong, is exclusively of New World distribution. The two genera known from the Greater Antilles seem to belong to a different section of the subfamily from where Protoptila belongs. The larvae of the family all construct cases of small sand grains shaped like a turtle’s shell. At pupation the ventral strap is cut away and the domelike dorsal covering is firmly anchored to the substrate, usually a rock, and a separate silken inner cocoon is spun to enclose the pupa. Genus Protoptila Banks Protoptila Banks, 1904, p. 215. [Type-species: Beraea? maculata Hagen, 1861, by original designation.] The genus Protoptila, which contains many species and is found throughout the New World, is here recorded from the Antilles for the first time. Preliminary descriptions of the larvae have been given by Ross (1944) and Flint (1963). The immature stages of the species described herein agree structurally with the other known species of the genus. NO. 3665 CADDISFLIES—FLINT 7 Protoptila dominicensis, new species FiaurReEs 1-9 This species is a member of the maculata group, c'osest to P. resolda Mosely. From this species it differs in the nondivergent apices of the ninth sternum, the shape of the apical portion of the tenth tergite, and the very different aedeagus. Avutt.—Length of forewing 3 mm. Color brown; legs paler, fore- wing with intermingled brown and gold hairs, a pale line at anasta- Figures 1-9.—Protoptila dominicensis, new species: 1, male genitalia, lateral; 2, male genitalia, dorsal; 3, female genitalia, ventral; 4, female genitalia, lateral; 5, pupal labrum, dorsal; 6, larval foreleg, posterior; 7, larval thorax, dorsal; 8, larval head, anterior; 9, pupal mandible, anterior. 8 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 125 mosis. Male genitalia: eighth sternum elongate, tapering, tip slightly bifid; ninth sternum flattened, scooplike, tip bifid; lateral arms of tenth tergum rectangular, with a sharp apicomesal tooth; lateral process of aedeagus with a curled apical spine; aedeagus with dorso- mesal lobe, basal complex, narrow neck, and greatly enlarged apex. Female genitalia: eighth sternum with rounded ventrolateral lobes; internal plate rectangular in ventral view with a circular mesal opening, apically with a smaller ventral plate, a long internal whip attached to apex. Larva.—Length to 4 mm. Sclerites brown, head yellowish around eyes. Prosternum with a pair of broad plates filling the venter. Meso- and metasterna with narrow sclerites along posterior margins. All legs very similar; mid- and hindlegs with ventroapical seta of tibia nonfimbriate. Abdominal segments 1-6 with 1 pair, segments 7-8 with 2 pairs of dorsal setae; segments 1-8 with a lateral seta; segment 1 with 2 pairs, segments 2-9 with 1 pair of ventral setae. Ninth tergite with 2 pairs of long setae. Anal claw with 3 pairs of accessory teeth. Pupa.—Length 3 mm. Mandibles and labrum as in figures 5, 9. Face with 3 pairs of setae. Abdomen with hook-plates anteriorly on seg- ments 2-8 (2 and 8 lightly sclerotized and may be lacking), posteriorly on 4. Case.—Length 4 mm, width 2 mm. Made of small sand grains in typical shape. Irregular respiratory openings dorsally at anterior and posterior ends. MateriIAL.—Holotype, pharate male: Dominica, Morne Nicholls, 9 Nov. 1964, P. J. Spangler, USNM Type 69879. Allotype, female: Fond Figues, 6 Apr. 1964, O.S. Flint, Jr. Paratypes: same data as holotype, 1 2 ; same data as allotype, 19; Clarke Hall, 11-20 Feb. 1965, W. W Wirth, 19. Other: same data as holotype, 7 larvae, 1 prepupa, 6 pupae; Laudat, 20 Nov. 1964, P. J. Spangler, 1 larva. Brotocy.—This species has been encountered only a few times on the island of Dominica. The larvae were found in clear, forested streams about 5 yards wide, where they were attached to gravel and rocks on the bottom. Family PHILopoTaMIDAE The philopotamids are nearly ubiquitous wherever there is flowing water in most of the regions of the world. Many of the genera, in- cluding Wormaldia, are more or less limited to the cooler, spring-fed streams in mountainous regions, whereas others, especially Chimarra, are more diverse in the regions of larger, warmer, but still rapidly flowing, lowland rivers. The larvae construct long, tubular, silken shelters attached to the underside of a rock or stick, where the flow of water will keep the NO. 3665 CADDISFLIES—FLINT 9g shelter distended. The pupae are enclosed in a rather loose, domed shelter of sand and silk. Key to Genera LARVAE Frontoclypeus with anterior margin evenly convex. . . ... . . Wormaldia Frontoclypeus with anterior margin asymmetrically emarginate . . . Chimarra PUPAE Mandible broad, subapical teeth close together, often arising from a single pro- PCUIOITUERR Re ences re Bet wei fo pt et ata: oak “all ete aserr ia oo ps OMIM AER Mandible narrower, teeth well separated, never arising from a single projection Wormaldia ADULTS Brostmpiawith. 1 apical spuri. .eic5 Be ae yet ee ees se SQ &. Chimarra Brenton with.2, apical spursis..o/. <.-.)! 3,2. «3 3 « » « -.. .Wormaldia Genus Wormaldia McLachlan Wormaldia McLachlan, 1865, p. 140. [Type-species: Hydropsyche occipitalis Pictet, 1834, designated by Ross, 1949.] There are species placed in the genus Wormaldia from all regions of the world save the Australian; however, the genus seems to be most diverse in the Northern Hemisphere. This is the first record of the genus from the West Indies. Larvae and pupae of the genus were described by Ross (1944), Lepneva (1964), and others. Wormaldia planae Ross and King Fiaures 10, 11 Wormaldia planae Ross and King, 1956, p. 64. This species was described from Mexico, but I have seen examples from Panama, Colombia, and Trinidad, as well as these recorded here. It is thus a circum-caribbean species, and it may also be expected in the Andean region further south. Apuut.—Length of forewing 4 mm. Color brown, legs slightly paler. Male genitalia: no sternal processes; eighth tergum with a U-shaped dorsomesal excision, flanked by a pair of knoblike pro- tuberances; tenth tergum narrow, with apex enlarged in lateral aspect, dorsally with an expansion and a blunt tooth laterally, then narrow- ing to a subapical constriction, and a sharp apicolateral tooth; cercus reaching posteriad to subapical constriction of tenth tergum; clasper with basal segment short and broad, apical segment about as long as basal segment, somewhat narrowed, with a dense apicomesal patch of black spines; aedeagus with internal sclerotizations in the form of long 10 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 lateral rods, a domelike basal hood, and an internal complex at midlength. Larva AND PuPpA.—Unknown. MateEriAu.— Grenada, 2 miles west of Lake Grand Etang, 4-8 Aug. 1963, O.S. Flint, Jr., 40. Brotocy.—The specimens were taken at an ultraviolet light placed beside a small tumbling, mountain brook. Genus Chimarra Stephens Chimarra Stephens, 1829, p. 318. [Type-species: Phryganea marginata Linnaeus, 1767, by monotypy.] This is a very large genus with species found on every region of the earth. One or more species have been found on all the Antillean islands. The larvae have been described many times. They live in silken tubes on the undersurface of rocks in fast water. Key to Species LARVAE Anterior margin of frontoclypeus with a central tooth. . . . . C. dominicana Frontoclypeus without a small central tooth... ..... . . C. antilliana PUPAE Mandibles with 2 separate subapical teeth. . .... .. . . €. dominicana Mandibles with 2 teeth arising from a common projection. . . . C. antilliana ADULTS 1. Clasper of male with a long apicomesal projection; M in hindwing with only 2) branches’. ie 6st se eg es Se we ee 2 CR omuncana Claspers without a long apicomesal projection; M with 3 branches. . . . 2 2. Clasper considerably enlarged basally; female with an elongate internal, ventral sclerite. ... Z ~ . « « « (Co caribea Clasper barely broader basally; female without an elongate ventral sclerite. C. antilliana Chimarra dominicana, new species Figures 12-17 This species is apparently related to C. puertoricensis Flint. It differs in the ornamentation of the eighth tergum, the presence of Figures 10-26.—Wormaldia planae Ross and King, male genitalia: 10, lateral; 11, dorsal. Chimarra dominicana, new species: 12, male genitalia, dorsal; 13, male genitalia, lateral; 14, aedeagus, lateral; 15, pupal mandible, anterior; 16, apex of larval frontoclypeus and mandibles, dorsal; 17, female genitalia, lateral. C. caribea, new species: 18, clasper, posteroventral; 19, male genitalia, lateral; 20, aedeagus, lateral; 21, female genitalia lateral. C. antilliana, new species: 22, apex of larval frontoclypeus and mandibles, dorsal; 23, pupal mandible anterior; 24, male genitalia, lateral; 25, clasper, posteroventral; 26, female genitalia, lateral. 11 FLINT CADDISFLIES NO. 3665 12 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 two apical teeth on the ventral half of the tenth tergum, and the long apicomesal process of the clasper. The larvae and pupae are attributed to this species on circumstantial evidence. Adults of two species of Chimarra have been taken on Dominica and the adults and larvae of C. antilliana have been defi- nitely associated by means of metamorphotypes. By elimination, therefore, the larvae herein described must be those of C. dominicana. Apu.tr.—Length of forewing 4 mm. Color fuscus, coxae and femora slightly paler. Vein M in hindwing with 2 branches. Male genitalia: eighth tergum with a broad mesal exicision, lateral angles of which are developed into elongate points; ninth sternum with a short, terete apicomesal process; tenth tergum divided mesally, each side divided again into dorsal and ventral lobes; dorsal lobe rodlike, ventral lobe widened apically with two apicolateral teeth; cercus elongate; clasper slender and elongate, with a long apicomesal process; aedeagus (partially everted) with a pair of small dark apicoventral spines, and a longer pair of spines, a basal ring and rod with a pair of lateral wings. Female genitalia: eighth segment without anterolateral proc- esses; posterior margin with a rounded lateral lobe bearing three large setae, ventral surface truncate; ninth tergum with a short anteroventral process, and a pair of dorsoapical lobes; bursa copulatrix consisting of several elongate sclerotized straps. Larva.—Length to § mm. Sclerites pale yellowish brown, posterior margin of pronotum black. Anterior margin of frontoclypeus with a slightly asymmetrical emargination that bears a short, pointed process centrally. Left mandible with a short pointed molar tooth, right mandible with a small molar area. Pupa.—Length 5 mm. Mandibles with inner margin serrulate, with 2 large teeth. Apex of abdomen with a few bristly hairs. MareriaLu.—Holotype, male: Dominica, .4 miles east Pont Casse, 6 May 1964, O. S. Flint, Jr.. USNM type 69880. Allotype, female: 2.2 miles east of Pont Casse, 11 May 1964, O. 8. Flint, Jr. Paratypes: Pont Casse, .5 miles south, 22-24 July 1963, O. S. Flint, Jr., 10749; same, but 8 Apr. 1964, 39; same, but 23 Apr. 1964, 1o°1?; same, but 17 June 1964, 19; Pont Casse, .4 miles east, 21 Apr. 1964, O. 8. Flint, Jr., 29; same, but 27 Apr. 1964, 5719; same, but 6 May 1964, 10759; same, but 23 June 1964, 29; Pont Casse, 1.3 miles east, 29 Apr. 1964, O. 8. Flint, Jr., 19; same, but 12 May 1964, 19; same, but 18 May 1964, 19; same, but 11 June 1964, 19; Pont Casse, 2.2 miles east, 1 May 1964, O. S. Flint, Jr., 1c7; same, but 14 Apr. 1964, 19; Pont Casse, 27-30 Nov. 1964, P. J. Spangler, 19. Other: D’leau Gommier, 27 Apr. 1964, O. S. Flint, Jr., 4 larvae, 2 prepupae, 1 pupa. Biotoay.—This species has been taken at elevations of around 2000 feet near Pont Casse and on the central divide. Larvae believed to be of this species were taken in a small, rapid stream on the under- side of rocks in the bottom gravel. NO. 3665 CADDISFLIES—FLINT 13 Chimarra antilliana, new species Figures 22-26 This species is clearly related to the following new species as is shown by the structure of the tenth tergum and aedeagus. It may be recognized by the more elongate, narrower clasper, which bears sub- apically a thin mesal shelf. Aputr.—Length of forewing 4-5.5 mm. Color fuscus; coxae and femora pale brown. Vein M in hindwing with 3 branches. Male geni- talia: eighth tergum unmodified; ninth sternum with a short terete apicomesal process; tenth tergum with a dorsomesal flaplike process, lateral lobe with a long straplike apicoventral process and a short apicodorsal one; cercus short, rounded; clasper elongate, straight, and barely widened basally in lateral aspect; in ventral aspect, bowed outwardly, with a narrow shelflike mesal projection sub- apically; aedeagus with 2 internal spines, a scabrous pouch with a darkened rounded lobe, and basal rod and ring. Female genitalia: eighth segment completely divided dorsomesally, with anterolateral processes, posterior margin with a few large setae; ninth tergum with long anterolateral processes; bursa copulatrix consisting of a heavily sclerotized ring basally, and apically of a nearly vertical plate capped dorsally by a hoodlike structure. LarvaE.—Length to 9 mm. Sclerites yellowish brown, posterior margin of pronotum black. Anterior margin of frontoclypeus asym- metrically emarginate. Left mandible with enlarged molar area and blunt tooth, right mandible with arcuate molar area and basal tooth. Pura.—Length 4-5-mm. Mandibles with inner margin serrulate, with a large subapical, bifid tooth. Marertau.—Holotype, male: Dominica, Mannett Gutter, near Clarke Hall, 4 April 1964, O. S. Flint, Jr. USNM Type 69881. Allotype, female: same data. Paratypes: same data, 407; same, but 10 Mar. 1965, W. W. Wirth, 1c; Clarke Hall, 11-20 Jan. 1965, W. W. Wirth, 107; same, but 18-19 Jan. 1965, J. F. G. Clarke, 22; same, but 27 Jan. 1965, 12; same, but 1-10 Feb. 1965, W. W. Wirth, 17; same, but 10 Feb. 1965, J. F. G. Clarke, 10; same, but 11-20 Feb. 1965, W. W. Wirth, 19 ; same, but 21-28 Feb. 1965, 1c; same, but 21-31 Mar. 1965, 3c° 42 ; same, but 31 Mar. 1965, D. R. Davis, 16; same, but 1 June 1964, O. S. Flint, Jr., 1; same, but 9 June 1964, 19; same, but 3-8 Oct. 1964, P. J. Spangler, 19; same, but 1-7 Dec. 1964, 1% 59; Fond Figues, 23 Jan. 1965, W. W. Wirth, 1? ; same, but 9-13 Mar. 1965, 17 1692; same, but 16-17 Mar. 1964, D. F. Bray, 2c’ 59; same, but 6 Apr. 1964, O. S. Flint, Jr., 60% 63 9 ; same, but 7 May 1964, 707, 449; same, but 1 May 1965, D. R. Davis, 1c 29; same, but 10 June 1964, O. S. Flint, Jr., 42% 2219; same, but 1 Dec. 1964, P. J. Spangler, 1 9 ; Pagua Bay, 19 Nov. 1964, P. J. Spangler, 1; Cabrit Swamp, 3 Nov. 1964, P. J. Spangler, 19; Clarke Hall, 1 mile east, 4 Apr. 1965, D. R. Davis, 136° 29; same, but 19 Apr. 1965, 5c’ 29; same, but 22 May 1965, 40°; Layou Valley (upper bridge), 22-25 July 1963, O. S. Flint, Jr., 8c"; Springfield Estate, 20-26 July 1963, O. S. Flint, Jr., 1%; Mahaut, Oct. 1966, E. L. Todd, 14 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 126 1; Syndicate Estate, 5 Mar. 1964, D. F. Bray, 19 ; Pont Casse, 12 Jan. 1965, J. F. G. Clarke, 1? ; Pont Casse, 2.5 miles north, 8 Apr. 1965, D. R. Davis, 8 9 ; Pont Casse, 3.5 miles north, 5 Dec. 1964, P. J. Spangler, 27 104? ; Pont Casse, 2.2 miles east, 1 May 1964, O. S. Flint, Jr., 5c? 19; same, but 2 May 1964, 70°29; same, but 7 May 1964, 4c7 39 ; same, but 11 May 1964, 17 49; same, but 6 June 1964, 8o° 19; same, but 19 June 1964, 29; Pont Casse, 1.3 miles east, 29 Apr. 1964, O. S. Flint, Jr., 1? ; same, but 10 May 1964, 4¢° 59 ; same, but 12 May 1964, 3c? 139; same, but 18 May 1964, 29; same, but 11 June 1964, 1c 19; Pont Casse, .4 mile east, 27 Apr. 1964, O. S. Flint, Jr., 19; Pont Casse, .5 mile south, 8 Apr. 1964, O. 8. Flint, Jr., 3o° 39; same, but 11 Apr. 1964, 27 39 ; same, but 17 June 1964, 2 9 ; same, but 22-24 July 1963, 167° 39; Pont Casse, 1.6 miles west, 28 Apr. 1964, O. S. Flint, Jr., 3o7; same, but 16 June 1964, 1c’ 49; Pont Casse, 3 miles east, 13-16 Oct. 1966, A. B. Gurney, 19; same, but 23 Oct. 1966, E. L. Todd, 1c 9 ? ; same, but 26 Oct. 1966, 1 9 ; La Ronde River, 15 Feb. 1964, H. Robinson, 19; Trafalgar, 21 May 1965, D. R. Davis, 20° 19; La Plaine, 23 Nov. 1964, P. J. Spangler, 19. St. Lucta: Vergallier River near Marquis, 31 July 1963, Flint and Cadet, 2? ; same, but 2 Aug. 1963, 19; R. Galet, south of Dennery, 1 Aug. 1963, Flint and Cadet, 217 159. Other: Dominica: Roseau River, swift water, 16 June 1911, 12 larvae; Springfield River, Springfield Estate, 20-26 July 1963, O. S. Flint, Jr., 8 larvae, 3 prepupae, 2 pupae, 1ch 19 metamorphotypes; Pont Casse, 2.2 miles east, 3 May 1964, O. S. Flint, Jr., 5 larvae; same, but 15 June 1964, 1 pupa, 1c metamorphotype; Pont Casse, .5 mile south, 22-24 July 1963, O. S. Flint, Jr., 1 larva, 1 prepupa, 3 pupae; Rosalie, 30 Nov. 1964, P. J. Spangler, 1 larva, 2 prepupae; Belfast River, .75 mile above mouth, 31 Jan. 1964, H. H. Hobbs, Jr., 11 larvae; Batali River, north of Salisbury, 21 Feb. 1964, R. Zusi and H. H. Hobbs, Jr., 1 larva; Espagnole River, cascades on Mt. Diablotin, east of Syndicate Estate, 26 Jan. 1964, H. H. Hobbs, Jr., 1 larva; Fond Figues, 1 Dec. 1964, P. J. Spangler, 2 larvae, 3 pupae, 3c’ metamorphotypes; Deux Branches, 30 Nov. 1964, P. J. Spangler, 2 larvae, 1 9 metamorphotype; Pagua Bay, 19 Nov. 1964, P. J. Spangler, 2 larvae. St. Lucia: Vergallier River, near Marquis, 31 July 1963, Flint and Cadet, 8 larvae, 1 prepupa, 1 pupa, 3c metamorphotypes; R. Galet, south of Dennery, 1 Aug. 1963, Flint and Cadet, 12 larvae, 2 pupae, 1? metamorphotype. Briotocy.—This species is one of the most frequently encountered Trichoptera on Dominica and St. Lucia. It has been taken in the larger lowland rivers and the small tumbling mountain brooks. The immature stages were taken under stones in the fast water of riffles and cascades. Chimarra caribea, new species Figures 18-21 This species appears to be the same as the one I have from the island of Trinidad and to be extremely close to C. duckworthi Flint from Costa Rica. The differences from the latter species lie in the narrower lateral lobes of the tenth tergites and the slightly more elongate clasper with a prominent thumblike apicomesal lobe in (@. caribea. Additional material from Venezuela and Colombia may show these differences to be clinal, but for the present I consider them specifically distinct. Chimarra antilliana is also related, but very distinct in the shape of both the clasper and the tenth tergum. NO. 3665 CADDISFLIES—FLINT 15 Apuut.—Length of forewing 4.5-5 mm. Color fuscus; coxae and femora yellowish brown. Vein M of hindwings 3 branched. Male genitalia: ninth sternum with an elongate, terete, apicomesal process; tenth tergum with a dorsomesal flaplike lobe, lateral lobe with a darkened slightly protruding area at midlength bearing sensillae, ventrally with a pair of straplike sclerites articulating beneath aedea- gus; clasper elongate, enlarged basoventrally, apex enlarged, curved mesally, with a thumblike, subapical projection; aedeagus with a spine- like apicoventral lip, a scabrous sac with darkened pouch, two spines, and ring and rod. Female genitalia: eighth segment divided dorsomesally, with anterolateral processes, posterior margin with a few enlarged setae; ninth tergum with very long anterolateral processes; bursa copulatrix with a basal ring, elongate ventral plates, and a dorsal hoodlike structure. LaRvA AND PupA.—Unknown. Materrau.—Holotype, male: Grenada, 2 miles west of Lake Grand Etang, 4-8 Aug. 1963, O. S. Flint, Jr., USNM Type 69882. Allotype, female: same data. Paratypes: same data, 14o° 39. Other: Trinidad, Simla, Arima Valley, 9-12 Feb. 1966, S. S. and W. D. Duckworth, 607; same, but 13-19 Feb. 1966, 1107; same, but 20-26 Feb. 1966, 6c. Brotogy.—The adults were collected on Grenada beside a small (3 feet wide by 3-6 inches deep) tumbling, mountain brook. It is assumed that the larvae were living in the same stream although none were found. Family PsycHOMYIIDAE The psychomyiids are found throughout the world. Although most species breed in flowing water, there are a number that breed in lentic situations. The genera Xiphocentron, Cernotina, and Polycentropus are found throughout the Antilles although the species of the latter genus on the Greater Antilles are not closely related to the Lesser Antillean species. The West Indian endemic genus Antillopsyche is limited to the Greater Antilles, but the widespread genus Polyplectropus has not been found on these islands. The immature stages of most genera make flimsy silken nets to trap their prey although certain others make long silken tubes affixed to the substrate. The pupae are generally sheltered in a domelike case of silk and sand. Key to Genera LARVAE 1. Tibia and tarsus fused; a broad flat process extending anteriorly from the MHESOPICULO Msi... Nay Meena, ZeNeW Suaeos ate eid. Ae .... . . . Xiphocentron Tibia and tarsus not fused; no process from mesopleuron. . ..... . 2 2. Mandibles with dorsal row of teeth overhanging ventralrow. .. . «2a iS Mandibles with dorsal row not overhanging ventralrow . . . Polycentropus 16 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 3. Head with dark muscle scars; anal claw with large ventral teeth. Polyplectropus Head uniformly pale; anal claw without ventral teeth... . . Cernotina PUPAE 1. Tip of mandible hooked; apical process with 4 setae. . . . . Xiphocentron Tip of mandible not hooked; apical process with many setae. ..... 2 2. Apical process with about 10 setae; gills absent. . . .. . . Cernotina Apical process with many more than 10 setae; gills present. . Polycentropus ADULTS 1. Foreleg with preapical sputic 3.06: 0s as 6 Foreleg without preapical spur. . . be a aol sce he 2. Hindwing with Rz and R; fused to wing Sarin . .. . . . Polyplectropus Hindwing with R, separating from R3 before wing margin . . Polycentropus 3. Anterior scutal warts of mesonotum delimited by a distinet lateral suture; general color black. .... ... . . Aiphocentron Anterior scutal warts without cree Sitpats es; eters color yellowish. Cernotina Genus Xiphocentron Brauer Xiphocentron Brauer, 1870, p. 66. [Type-species: Xiphocentron bilimeki Brauer, 1870, by monotypy.] Species of the genus are found from southwestern United States south through South America. All the Antillean islands intensively collected for Trichoptera have proven to support one or two species. The immature stages were described by Edwards (1961) and Flint (1964b). The adults of the genus are generally diurnal and rarely attracted to lights, which accounts in part for their scarcity in col- lections. Key to Species ADULTS Color totally black. .... wee « te) oes OSC rm Color black, with a line of white meine alone he Tne ior osetia of the forewings. X. albolineatum Xiphocentron fuscum, new species Figures 27, 28 The species seems to be closest to X. borinquensis Flint from Puerto Rico. From this it differs in the apically widened cerci, much shortened anterior process of the ninth sternum, and differently shaped mesal lobes of the ninth tergum. Apuut.—Length of forewing 3.4-4 mm. Color fuscus, coxae and femora pale brown, hairs on face ventrad of antennae pale brown. Apical spur of hindleg in male about 4% length of basal tarsal seg- ment. Male genitalia: ninth sternum with anterolateral process short and blunt, posterior margin truncate; ninth tergum rectangular in lateral aspect, posterior margin with a V-shaped mesal incision; tenth NO. 3665 CADDISFLIES—FLINT 17 Ficures 27-37.—Xiphocentron fuscum, new species, male genitalia: 27, lateral; 28, dorsal. X. albolineatum, new species, male genitalia: 29, lateral; 30, dorsal. Cernotina lutea, new species: 31, female genitalia, ventral; 32, male genitalia, lateral; 33, apex of clasper, ventral; 34, male tenth terga and cerci, dorsal. C. cadeti, new species, male: 35, genitalia, lateral; 36, tip of clasper, ventral; 37, tenth terga and cerci, dorsal. 313-152—68——2 18 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 tergum sclerotized laterally, tip decurved; cercus about 10 times as long as broad, widened subapicaily; clasper with apical portion narrow and sharply curved dorsomesally, mesally with many dark spicules; aedeagus very long, slightly enlarged apically. Female genitalia: eighth segment divided dorsally with long anterolateral processes; ninth segment long and slender with long anterolateral processes; apex of tenth segment with a pair of slender papillae. LARVA AND PUPA.—See Xiphocentron species. MateEr1aL.—Holotype, male: Dominica, Brantridge, 9 May 1964, O.S. Flint, Jr., USNM type 69883. Allotype, female: same data. Paratypes: same data, 507; same, but 30 April 1964, 1267; Pont Casse, .4 miles east, 7-8 May 1964, O. 8. Flint, Jr., 197; Pont Casse, 2.5 miles east, 16 Jan. 1965, W. W. Wirth, 10; Pont Casse, 1.5 miles north, 12 Feb. 1965, W. W. Wirth, 1c. Brotoagy.—Apparently this species is limited to the higher eleva- tions on the island of Dominica. I have swept them in numbers from rocks in nearly dry streambeds around Pont Casse. Xiphocentron albolineatum, new species FIGURES 29, 30 This is a species closely related to the preceding, from which it differs in possessing a line of white hairs along the posterior margin of the forewings and mesally on the head. The male genitalia differ most noticeably in the narrower clasper and the more elongate dorso- mesal lobes of the ninth tergum. Apuut.—Length of forewing 3-4 mm. Color fuscus, coxae and femora slightly paler, posterior margin of forewing, pro- and mesonota, and head mesally with a band of whitish hairs. Apical spur of hind tibia of male about 1 as long as basal tarsal segment. Male genitalia: ninth sternum with anterolateral process long and attenuate; ninth tergum rounded basally, with dorsomesal lobes slightly elongate; tenth tergum with tip sharply decurved; cercus slender, narrowing apically, about 12 times as long as broad; apex of clasper slender, sharply curved dorsomesad, mesal surface with elongate spiculate patch. Female genitalia: identical to that of preceding species. LARVA AND PUPA.—See Xiphocentron species. Mater1aL.—Holotype, male: Dominica, Pont Casse, 1.7 miles east, 12 March 1965, W. W. Wirth, USNM type 69884. Allotype, female: near Clarke Hall, 13 Feb. 1964, D. F. Bray. Paratypes: Pont Casse, 12 Oct. 1964, P. J. Spangler, 19; Mannett Gutter, 7 March 1965, W. W. Wirth, 19. Brotoay.—This species is known only from Dominica but appears to be more widespread on the island than the preceding. It undoubtedly has a slightly different habitat preference, which unfortunately is unknown. NO. 3665 CADDISFLIES—FLINT 19 Xiphocentron species I list here the immature stages of this genus that I have collected on the Lesser Antillean islands. Characters have not been found that will serve to separate the larvae of the various species; therefore, they are all listed together and described as a unit. Larva.—Length to 8 mm. Sclerites pale yellow, oral margin of head capsule darker. Structure apparently identical to other described species. Pupa.—Unknown. Casre.—A long silken tube attached to the substrate, generally a rock, often in great numbers crisscrossing the rocks both above and below the water line. Matertat.—Dominica, Pont Casse, .5 miles south, 22-24 July 1963, O. S. Flint, Jr., 4 larvae; same, but 15 June 1964, 1 larva. Springfield Estate, .5 miles east, 21 July 1963, O. S. Flint, Jr., 1 larva. Pont Casse, 2.2 miles east, 15 June 1964, O. S. Flint, Jr., 2 larvae. Fond Figues, 6 April 1964, O. S. Flint, Jr., 2 larvae. Mannett Gutter, 23 April 1964, O.S. Flint, Jr., 2 larvae. St. Lucia, Vergallier River, near Marquis, 31 July 1963, Flint and Cadet, 2 larvae. Genus Cernotina Ross Cernotina Ross, 1938, p. 136. [Type-species: Cernotina calcea Ross, 1938, by original designation.]} The genus is widely distributed in North and Central America and in the Greater Antilles. The immature stages of the genus still have not definitely been correlated with the adult. It seems probable, however, that the larvae and pupae described by Flint (1964b) from Puerto Rico as an unknown Polycentropodinae belong to a species of this genus. Key to Species ADULTS Ventrolateral lobe of cercus of male with tip bent sharply dorsomesad. C. cadeti Wensrointeral lobe of cercus spinelike . 2.050. 2 ee ee eel tek) C. lutea Cernotina lutea new species Figures 31-34 Cernotina lutea is a member of the calcea section of the genus closest to C. ohio Ross. From this species it differs in the elongate mesal lobe and in the long ventrolateral spines of the cerci. Apuutt.—Length of forewing 3.5-4 mm. Color yellowish brown, a slightly paler band of hairs mesally on head and thorax. Male genitalia: ninth segment expanded basoventrally; tenth tergites elongate, conical, 20 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 and semimembranous; cercus developed dorsally into a long slender, mesally curving process that bears on outer surface near base a short tooth and ventrolaterally a long curving spine; mesoventrally devel- oped into an elongate quadrate plate; clasper with prominent dorsal arm, tip bifid; aedeagus semimembranous, with a heavily sclerotized dorsomesal rod. Female genitalia: lateral lobes of eighth sternum elongate and slender; dorsally with a transverse sclerite between ninth and tenth segments; bursa copulatrix with an elongate ventral plate and a ringlike sclerite between dorsolateral rods. Larva AND pupA.—Unknown. MarerraAL.—Holotype, male: Dominica, Pont Casse, 1.3 miles east, 18 May 1964, O. S. Flint, Jr.. USNM Type 69885. Allotype, female: same data. Paratypes: same, but 10 May 1964, 19? ; Pont Casse, .4 miles east, 15 June 1964, O. S. Flint, Jr., 12; same, but 23 June 1964, 2 19; Brantridge, 30 April 1964, O. 8. Flint, Jr., 19 ; Cabrit Swamp, 23 Feb. 1965, W. W. Wirth, 17. Brotocy.—The adults have generally been taken near small streams at higher elevations on the island of Dominica. Presumably the larvae will be found in these streams. The one specimen from Cabrit Swamp probably represents an individual that wandered considerably from its breeding site. Cernotina cadeti, new species Figures 35-37 As to be expected, this species is clearly related to the preceding. It may be recognized by the lack of external teeth, by the more sharply angulate ventral spine of the cerci, and by the more mesally displaced dorsal plate at the tip of the clasper. Apuut.—Length of forewing 3 mm. Color in alcohol, uniformly pale brown. Male genitalia: ninth segment considerably expanded baso- ventrally; tenth tergites conical, semimembranous; cercus developed into a long curving dorsal process and a ventrolateral spine whose tip is sharply angulate dorsomesad; mesoventral process elongate, rod- like; clasper with a dorsal arm; tip bifid, with dorsal plate displaced mesally; aedeagus semimembranous, with a long dorsomesal sclerite. Larva AND pupA.—Unknown. Marertau.—Holotype, male: St. Lucia, Vergallier River, near Marquis, 31 July 1963, Flint and Cadet, USNM Type 69886. Brotogy.—The specimen was taken at a light near a slowly flowing stream about a yard wide, in which the larva probably developed. Genus Polyplectropus Ulmer Polyplectropus Ulmer, 1905, p. 103. [Type-species: Polyplectropus flavicornis Ulmer, 1905, by monotypy.] NO. 3665 CADDISFLIES—FLINT PA Ecnomodes Ulmer, 1911, p. 17. [New synonymy. ‘Type-species: Henomodes buchwaldi Ulmer, 1911, by monotypy.] Cordillopsyche Banks, 1913, p. 238. [Type-species: Cordillopsyche costalis Banks, 1913, by monotypy.] Ecnomodellina Ulmer, 1962, p.5. [Replacement name for Ecnomodes Ulmer 1911. New synonymy.] Genus C Flint, 1964a, p. 476. Species from tropical America, Africa, and the Orient have been placed in this genus; however, I expect that many of those from the Old World are not truly congeneric. On the basis of adult morphology, the genus is close to Polycentropus; indeed, most North American workers have not recognized the two as distinct. The adults of this genus may be recognized by the fusion of Re and R; in the hindwing and generally by the division of the clasper in the male into distinct dorsolateral and ventromesal lobes. The larvae have the mandibles with the dorsal row of teeth overhanging the ventral row, the tibia and tarsus of the fore- and midlegs have a row of enlarged and generally black setae on the posterior face, and the anal claw bears several long teeth ventrally. I have already synonymized the genus Cordillopsyche (Flint, 1967), and am here synonymizing Eenomodellina (=Ecnomodes) Ulmer with Polyplectropus Ulmer. The original description of HL. buchwaldi leaves little doubt that it is a typical species of Polyplectropus. The following New World species, mostly described in Polycentropus, must be transferred to Polyplectropus (all new combinations): Hcnomodes buchwaldi Ulmer, Polycentropus charlesi Ross, P. santiago Ross. P. thilus Denning, P. allent Yamamoto, P. altmani Yam., P. deltoides Yam., P. elongatus Yam., P. laminatus Yam., P. robacki Yam., P. recurvatus Yam. Polyplectropus bredini, new species Ficures 38-41, 47-49 This species seems quite unrelated to any other known species although there may be a slight relationship to P. charlest (Ross). From this species it is easily distinguished by the more distinctly bipartite clasper that bears large black spines. The species is named for Mr. J. Bruce Bredin, a cosponsor of the Dominica Survey. The relationship of the island populations is very close, but slight differences do exist. The males from St. Lucia have only three dark setae on the dorsal lobe of the clasper; the other dark seta on the Dominican specimens is pale. The Grenadan male (fig. 40) also has only three dark setae, but the ventralmost one is on an elongate process. Considering, the overall similarity however, I am considering these all one species. 46 Ficures 38-49.—Polyplectropus bredini, new species: 38, male genitalia, lateral; 39, claspers, ventral; 40, clasper of Grenadan specimen, lateral; 41, female genitalia, ventral. Poly- centropus insularis Banks: 42, male genitalia, lateral; 43, male genitalia, ventral; 44, tenth terga and cerci, dorsal; 45, female genitalia, ventral; 46, larval head and pronotum, dorsal. Polyplectropus bredini, new species, larva: 47, mandibles, dorsal; 48, anal claw, | lateral; 49, head and pronotum, dorsal. NO. 3665 CADDISFLIES—FLINT 28 Aputt.—Length of forewing 5-5.5 mm. Color light brown, with whiter hairs middorsally, anterior margin of forewing dark brown. Male genitalia: ninth sternum moderately rounded basally; tenth tergites developed as long terete processes; cercus with a rounded dorsolateral lobe, posteroventrally developed into a sharp point; clasper divided into a thin, concave upper lobe bearing 4 short, broad, black setae, generally on elongate bases; basal lobe scooplike with a distal row of short dark setae; aedeagus with apicoventral portion developed into an elongate, narrow, hoodlike process. Female geni- talia: lateral lobes of eigth sternum broad; subgenital plate produced and rounded apically; bursa copulatrix with a complex of sclerites apically, and a donut-shaped mesal structure. Larva.—Length to 8 mm. Head and pronotum pale brown with conspicuous dark muscle scars. Labrum, maxillolabium, and legs as illustrated by Flint (1964a, fig. 4). Mandibles with dorsal row of teeth overhanging ventral row. Basal segment of anal proleg with a few short setae mesally; claw with 3 long and 1 short ventral teeth, apex curved at right angles. Pupa.—Unknown. MareriAu.—Holotype, male: Dominica, Pont Casse, 1.3 miles east, 29 Apr. 1964, O. S. Flint, Jr. USNM Type 69887. Allotype, female: Syndicate Estate, 5 March 1964, D. F. Bray. Paratypes: same as holotype, but 11 June 1964, 10; same, but 12 May 1964, 10°; Fond Figues, 13 March 1965, W. W. Wirth, 10; D’leau Gommier, 24 Feb. 1965, J. F. G. Clarke, 59. Other: Grenada, 2 miles west Grand Etang, 4-8 Aug. 1963, O. S. Flint, Jr., 1¢. St. Lucia, Cul de Sae River, at mile post 9, 29 July 1963, Flint and Cadet, 1; R. Galet, south of Dennery, 1 Aug. 1963, Flint and Cadet, 1.7, 8 larvae. Brotocy.—The adults generally have been collected near small clear streams. The larvae were taken on St. Lucia in a stream only a few feet wide by several inches deep. They were found under rocks near the head or tail ends of pools in the stream. Genus Polycentropus Curtis Polycentropus Curtis, 1835, pl. 544. [Type-species: Polycentropus irroratus Curtis, 1835, by original designation.] Polycentropus, as it is presently recognized by most North American workers, is found over most of the world; however, it seems to be best developed in the Northern Hemisphere. There is at least one species on all the Antillean islands. The larvae of the genus are well known and described. They build various types of silken trap nets. 24 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 Polycentropus insularis Banks Figures 42-46 Polycentropus insularis Banks, 1938, p. 302.—Fischer, 1962, p. 83.—Flint, 1967, D0: Polycentropus insularis was described from Grenada, but the specimens collected on Dominica do not seem to differ significantly from the type. The species is rather distantly related to the other species of Antillean Polycentropus. From these it may be recognized by the lack of dorsomesal process on the clasper and the elongate ventromesal lip of the aedeagus. ApuLt.—Length of forewing 7-8 mm. Brown, body and wings flecked with spots of golden hair. Male genitalia: ninth segment rounded in lateral aspect; cercus divided into a dorsolateral ovate lobe bearing a smaller mesal lobe, and a long, angled, pointed process; clasper with a rounded dorsolateral lobe, and an elongate ventromesal section; aedeagus with a long pointed ventromesal lip. Female geni- talia: lateral lobes slightly elongate, directed apicomesally; subgenital plate rounded apically; bursa copulatrix complex, with a heavily sclerotized basal plate and lateral supports. Larva.—Length to 12 mm. Sclerites pale brownish, muscle scars conspicuously darker, head irregularly clouded with darker brown. Structure typical of other West Indian species. Pupa.—Unknown. MarertaL.—Holotype, male: Grenada, Grand Etang, Sept. 1910, Allen and Brues, collection MCZ. Other: Dominica, Pont Casse, .5 miles south, 22-24 July 1963, O. S. Flint, Jr., 1¢, 1 larva; Pont Casse, .4 miles east, 21 April 1964, O. S. Flint, Jr., 19; same, but 27 April 1964, 10°; same, but 6 May 1964, 39 ; same, but 16 May 1964, 19 ; same, but 12 June 1964, 1 larva; same, but 23 June 1964, 1c*; Pont Casse, 1 mile east, T. M. and J. F. G. Clarke, 2%; Pont Casse, 1.3 miles east, 29 April 1964, O. S. Flint, Jr., 19 ; same, but 10 May 1964, 19 ; same, but 18 May 1964, 19; same, but 26 May 1964, 1c; same, but 11 June 1964, 1c° 19; Pont Casse, 2.2 miles east, 1 May 1964, O.S. Flint, Jr., 1c; Pont Casse, 3 miles east, 15 Oct. 1966, E. L. Todd, 19 ; Sylvania Estate, 28 Jan. 1965, T. M. and J. F. G. Clarke, 1? ; Boeri Lake, 22 Feb. 1964, D. F. Bray, 27; Fresh- water Lake, 13 Oct. 1964, P. J. Spangler, 1 larva. Biotoay.—The species has been taken only at higher elevations on the islands. Larvae were found sparingly in small streams. The two adults taken at Boeri Lake and the larva from Freshwater Lake indicate that the species may breed in high elevation lakes as well. Family HypRopsYCHIDAE The Hydropsychidae are a very common and widely distributed family, breeding in flowing waters throughout the world. Represent- atives of two subfamilies, the Macronematinae and Hydropsychinae, NO. 3665 CADDISFLIES—FLINT 25 have been found on the Antilles. Leptonema and Smicridea, respec- tively, are the representatives throughout the Antilles, but Macronema and Hydropsyche are found only on the Greater Antilles. The larvae construct a complex silken trap-net to strain their food from the water. The trap-net is attached to a tubular retreat in some crevice of the substrate. At pupation time a domelike shelter is constructed of silk with included sand or organic matter. Key to Genera LARVAE Gills consisting of a long central stalk giving rise to many lateral filaments. Leptonema Gills branching into filaments near base. ......... .. . .Smicridea PUPAE Hook-plate present posteriorly on segment 3 only. .... . . . .Leptonema Hook-plates present posteriorly on segments 3 and 4... . . . .Smicridea ADULTS Antennae at least twice as long as wing; large and pale green. . . . Leptonema Antennae subequal to forewings; small and dark. .... . . . .Smicridea Genus Smicridea McLachlan Smicridea McLachlan, 1871, p. 134. [Type-species: Smicridea fasciatella McLachlan, 1871, by designation of Milne, 1936.] Smicridea is a genus of many species found from the southwestern United States to the southern tip of South America and Australia. All of the Antillean islands support at least one species. The larvae and pupae have been described on several occasions (Ross 1944, Flint 1964b). Key to Species PUPAE Hook-plate 3P equidimensional with 3 long hooks. ..... . .S. cariba Hook-plate 3P nearly twice as wide as long, with 5 hooks. . . . .S. simmonsi ADULTS 1. Forewing black with 2 transverse bands of white setae. ........2 Forewing uniformly grayish. ... . ste : . S. grenadensis 2. Tenth tergite short and broad, aedeagus boat tera plate. .S. simmonsi Tength tergite long and narrow, aedeagus with lateral plate. . .S. cariba Smicridea cariba, new species Friaures 50-56 This species, found on Dominica, is most closely related to the following species from St. Lucia. It differs strikingly in the structure of the tenth tergite and aedeagus. 26 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 125 Ficures 50-65.—Smicridea caribea, new species: 50, male genitalia, lateral; 51, aedeagus, lateral; 52, male genitalia, dorsal; 53, female genitalia, dorsal; 54, bursa copulatrix, dorsal; 55, larval mandibles, dorsal; 56, pupal hook-plates, dorsal. S. simmonsi, new species: 57, pupal hook-plates, dorsal; 58, aedeagus, lateral; 59, male genitalia, lateral; 60, male genitalia, dorsal. S. grenadensis, new species: 61, male genitalia, lateral; 62, aedeagus, lateral; 63, male genitalia, dorsal; 64, female genitalia, dorsal; 65, bursa copulatrix, dorsal. NO. 3665 CADDISFLIES—FLINT pare Avuur.—Length of forewing 4-5 mm. Color fuscus, leg bases slightly paler, forewing with a transverse band of iridescent white hair at level of anastamosis and another halfway to wing base, apical fringe white. Sixth and seventh segments of male with internal reticulate sacs. Male genitalia: ninth segment with anterior margin angulate at middle; tenth tergite elongate, rounded apically, with lateral margin heavily sclerotized; clasper widening uniformly to apex of basal segment, apical segment terete; aedeagus large and complex, apicoventrally heavily sclerotized and scooplike, a ventrally directed spine laterally at midlength, a short dorsal hood, a flat lateral plate in apical membrane that bears 2 rows of short spines and an apicodorsal angulate rod. Female genitalia: lobes of eighth sternum slightly longer than broad, evenly rounded apically; ninth segment without clasper groove or receptacle, with anterolateral angle greatly enlarged; internal plate with a quadrate central section bearing short apical arms and a pair of long basal arms; bursa copula- trix with a central ring and broad basolateral wing-like supports. Larva.—Length to 7 mm. Sclerites brownish. Each gill on basal abdominal segments with 3-4 filaments. Abdomen with many short, broad, black setae. Pupa.—Length 4-5 mm. Right mandible with 3 inner teeth, left with 4. Hook-plates anteriorly on segments 2-8, posteriorly on 3 and 4; posterior plates about equidimensional, anterior plates with 2-3 teeth, apex of posterior plates considerably elevated above body. Apical processes widely separated, with a brush of setae on apical third. MatTERIAL.—Holotype, male: Dominica, Pont Casse, 2.2 miles east, 2 May 1964, O. S. Flint, Jr., USNM Type 69888. Allotype, female: same data. Paratypes: same data, 8o' 49; same, but 14 April 1964, 1c; same, but 1 May 1964,7¢ 39; same, but 7 May 1964, 1c 5? ; same, but 11 May 1964, 16°; same, but 21 May 1964, 19 ; Pont Casse, 1.3 miles east, 10 May 1964, O.S. Flint, Jr.,2719; same, but 12 May 1964, 6¢° 19; same, but 18 May 1964, 19 ; same, but 11 June 1964, 19; Pont Casse, .4 miles east, 21 April 1964, O. S. Flint, Jr., 17; same, but 27 April 1964, 2¢°?; same, but 6 May 1964, 4o° 12; same, but 7 May 1964, 1; same, but 15 June 1964, 1<; Pont Casse, .5 miles south, 22-24 July 1963, O.S. Flint, Jr., 2° 19 ; Sylvania, 9 Feb. 1964, D. F. Bray, 1; Trafalgar Falls, 15 March 1964, D. F. Bray, 1¢° 19; Fond Figues, 23 Jan. 1965, W. W. Wirth, 19; D’leau Gommier, 15 Feb. 1965, W. W. Wirth, 1c’. Other: Pont Casse, 2.2 miles east, 3 May 1964, O. S. Flint, Jr., 8 larvae; Pont Casse, .4 miles east, 20 May 1964, O.S. Flint, Jr., 2 larvae; same, but 12 June 1964, 8 larvae; same, but 25 June 1964, 3 larvae, 1 prepupa, 1 pupa; Pont Casse, .5 miles south, 22-24 July 1963, O. 8. Flint, Jr., 36 larvae, 2 prepupae, 16 pupae, 29 metamorphotypes; same, but 15 June 1964, 6 larvae, 4 pupae, 1 ? metamorphotype; D’leau Gommier, 27 April 1964, O. S. Flint, Jr., 5 larvae, 1 pupa; Springfield Estate, 20-26 July 1963, O.S. Flint, Jr., 1 larva, 1 pupa; Roseau River, swift water, 16 June 1911, 5 larvae; Espagnole River, cascades on Mt. Diablotin, east of Syndicate Estate, 26 Jan. 1964, H. H. Hobbs, Jr., 1 larva; Boeri Lake, outlet, 10 Nov. 1964, P. J. Spangler, 4 larvae. 28 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 Biotocy.—The species is most abundant around the small, tum- bling mountain brooks; however, the collection of larvae in the Roseau River suggests that they breed sparingly in the large lowland rivers as well. Smicridea simmonsi, new species Figures 57-60 This species appears to be most closely related on the basis of male genitalia to S. grenadensis, although in coloration it is apparently the same as S. cariba. From S. grenadensis it may be separated by the more evenly rounded apices of the tenth tergites, by the straight lateral process of the aedeagus, and by the very long apical segment of the clasper, Aputt.—Length of body 4.5 mm, forewing probably about 5 mm. Wing dark with a transverse white band at region of anastamosis, apparently some white obliquely along the base of Cu. Sixth and seventh abdominal segments of male with internal reticulate sacs. Male genitalia: ninth segment with a pronounced dorsolateral angle; tenth tergite short, broad, apex rounded; basal segment of clasper short, apical segment subequal in length; aedeagus with well-developed ventral scoop, lateral spine straight, no lateral plate, dorsal hood semimembranous, membranous portion with about 6 pairs of long spines, apex with a crenulate crecentic hood above an elongate U-shaped sclerite. Larva.—Length to 7 mm. No differences found from the larva of cariba. Pupa.—Length 4.5 mm. As in S. cariba, except hook-plate 3P almost twice as wide as long and with 5 hooks. Marteriau.—Holotype, pharate male: St. Lucia, Vergallier River, near Marquis, 31 July 1963, Flint and Cadet, USNM Type 69889. Other: same data, 22 larvae, 1 pupa; Grand Riviere du Mabouya, 29 July 1963, Flint and Cadet, 1 larva; Cul de Sac River at mile post 9, 29 July 1963, Flint and Cadet, 2 larvae, 1 pre- pupa, 1 pupa; R. Galet, south of Dennery, 1 Aug. 1963, Flint and Cadet, 32 larvae, 1 prepupa, 3 pupae. Brotocy.—The larvae of this species are most abundant in the small clear streams on the island of St. Lucia, but they are also found in the larger rivers in smaller numbers. Smicridea grenadensis, new species Fiaures 61-65 This species, known only from Grenada, is most closely related to S. simmonsi of St. Lucia; however, it is nearly unicolorous, and and in the male the tenth tergite bears a distinct anteapical bulge, NO. 3665 CADDISFLIES—FLINT 29 the basal segment of the clasper is longer, and the lateral spine of the aedeagus is hooked. Apuutt.—Length of forewing 4-4.5 mm. Color uniform grayish brown. Males with two pairs of ovoid internal sacs in abdominal segments 6 and 7. Male genitalia: ninth segment with pronounced dorsolateral angle; tenth tergite short, blunt, with anteapical dorsal bump; clasper with apical segment about half as long as basal segment; aedeagus with a strong ventral scoop, lateral spine hook-shaped, membranous portion with about 6 pairs of long spines. Female genitalia: lobes of eighth sternum slightly longer than broad, pro- duced apicomesally; ninth segment without clasper groove or recep- tacle, greatly produced anterolaterally; internal plate with central part narrowly quadrangular, with long posterior and anterior arms, anterior pair arising mesally; bursa copulatrix with a central heart- shaped plate with a central hole, lateral supports U-shaped. Larva AND pupa.—Unknown. MareriaL.—Holotype, male: Grenada, 2 miles west Lake Grand Etang, 4-8 Aug. 1963, O. S. Flint, Jr.. USNM Type 69890. Allotype, female: same data. Brotocy.—The adults were taken beside a small tumbling mountain brook in which the larvae are probably to be found. Genus Leptonema Guerin Leptonema Guerin, 18438, p. 396. [Type-species: Leptonema pallidum Guerin, 1848, by monotypy.] This is a genus of many species throughout the American tropics and Africa. Species are known from Cuba and Puerto Rico on the Greater Antilles. The larvae have been described previously (Flint, 1964b). Key to Species LARVAE Coxa of foreleg with anteroapical process short and with setae on anterior OTR, shee ey Seam ciate an aan eg ee nema are ee L. archboldi Coxa of foreleg with process longer but without setae. ..... L. albovirens ADULTS Males with a very long, basally directed process from apex of aedeagus; females with a shallow groove laterally on 9th segment. ..... L. archboldi Males with a short lateroventrally directed process from apex of aedeagus; females with a deep, complex lateral groove on 9th segment . . L. albovirens Leptonema archboldi, new species FIGURES 66-70 This species is very distinct, not being closely related to any described species. It probably belongs to the stigmosum section of the 30 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 genus, from which it may be separated by the lack of the spot of dark hair on the forewing and the longer apical and shorter lateral pro- cesses of the aedeagus. I take pleasure in naming this species for Mr. John D. Archbold, one of the sponsors of this survey. 14 Ficures 66-74.—Leptonema archboldi, new species: 66, male genitalia, lateral; 67, tip of aedeagus, dorsal; 68, tip of aedeagus, lateral; 69, larval coxa, dorsal; 70, female genitalia, lateral. L. albovirens (Walker): 71, male genitalia, lateral; 72, tip of aedeagus, lateral; 73, female genitalia, lateral; 74, larval coxa, dorsal. Apvuut.—Length of forewing: male 12-13 mm, female 14-15 mm. Color pale green when alive, becoming brownish after death. Female with a yellow cellule on 1A of hindwing. Male genitalia: ninth segment narrow, slightly produced dorsomesally; tenth tergite broadly tri- angular, with a produced hirsute knob near ventral angle, and a setose patch dorsally; clasper long and slender, without basomesal process, basal segment with rows of spinelike setae apicomesally, NO. 3665 CADDISFLIES—FLINT 31 apical segment short with many spinelike setae mesally; aedeagus angled near base, with a long, basally directed, fimbriate process arising apically, a short fimbriate process subapically, and a dorso- mesal stub arising from a semicircular basal structure. Female genitalia: lobes of eighth sternum about % as broad as long; ninth segment with clasper receptacle shallow, entered from above, a narrow groove on surface ventrad to receptacle; with an indistinct angulate plate internally whose attachment to posterior margin of ninth segment is heavily sclerotized. Larva.—Length to 20 mm. Head and pronotum dark reddish brown, meso- and metanota paler, pale around eyes and on posterior of head. Anteroapical process of forecoxa short, with setae on anterior margin. Body covered densely with broad, erect, or decumbent, black setae. Pupa.— Unknown. Mareriau.—Holotype, male: Dominica, Pont Casse, .5 miles south, 22-24 July 1963, O. S. Flint. Jr.,. USNM Type 69891. Allotype, female: Pont Casse, 2.2 miles east, 2 May 1964, O. 8. Flint, Jr. Paratypes: same data as holotype, but 8 April 1964, 1°19; same, but 23 April 1964, 19 ; same data as allotype, but 14 April 1964, 1¢° 39; same, but 1 May 1964, 19 ; same, but 7 May 1964, 19; same, but 11 May 1964, 29; same, but 21 May 1964, 29; Pont Casse, 3 miles east, 26 Oct. 1966, E. L. Todd, 29; Pont Casse, 1.3 miles east, 18 May 1964, O. S. Flint, Jr. 19; Pont Casse, .4 miles east, 27 April 1964, O. S. Flint, Jr., 19; Boeri Lake, 10 May 1964, P. J. Spangler, 1 9; Pont Casse, 2.5 miles north, 8 April 1965, D. R. Davis, 3 9. Other: Pont Casse, .5 miles south, 22- 24 July 1963, O. S. Flint, Jr., 1 larva; same, but 16 Feb. 1964, H. H. Hobbs, Jr., 1 larva; Pont Casse, 2.2 miles east, 3 May 1964, O. 8. Flint, Jr., many larvae, 2 prepupae; same, but 15 June 1964, 1 larva, 1 prepupa; Espagnole River, Cas- cade on Mt. Diablotin, east of Syndicate Estate, 26 Jan. 1964, H. H. Hobbs, Jr., 3 larvae; Morne Nicholls, 9 Nov. 1964, P. J. Spangler, 2 larvae. Brotoagy.—This species seems to be restricted to the fast waters of small highland streams, and it may also breed around the rocky margins of Boeri Lake. The larvae make typical trap-nets and sand- enclosed retreats. Pupation takes place in a silk and sand cocoon attached tightly to a rock in the substrate. Leptonema albovirens (Walker) Figures 71-74 Macronema albovirens Walker, 1852, p. 76. Leptonema albovirens (Walker).—Mosely, 1933, p. 45.—Fischer, 1963, p. 166. The species is known from the extreme northeast of Mexico through- out Central America, across northern South America to Trinidad, and north in the Lesser Antilles to Grenada and St. Vincent. I give only a few more pertinent references to this species; a complete bibliography is to be found in Fischer (1963). 32 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 The species is related to L. dissimile Mosely, from which it differs in the much shorter apical process of the aedeagus. The following descriptions are based on Grenadan specimens. Apvuutr.—Length of forewing: male 11 mm, female 12-14 mm. Color pale green. Female with a yellowish cellule on vein 1A in hind- wing. Male genitalia: ninth segment narrow, posterior margin angulate above base of clasper; tenth tergite trianguloid in lateral view, with an apical setate patch, dorsoapically with 2 short processes; clasper long and slender, no mesobasal lobe; basal and apical segments with patches of short spinelike setae mesoapically; aedeagus with apex bearing 2 pairs of fimbriate processes, a ventrally directed process basad of gonopore, and a posteriorly directed fimbriate process appressed to side of aedeagus. Female genitalia: lobes of eighth sternum about 34 as broad as long; ninth segment with a shallow pouchlike dorsally directed receptacle and a lateral groove with a heavily sclerotized ventral margin leading to a rounded internal plate. Larva.—Length to 17 mm. Head and thoracic notae brown, paler around eyes and posteriorly on head. Process anteroapically on forecoxae arising from inner margin, with setal row passing anteriorly of it. Abdomen covered very densely with erect or decumbent broad, black setae. Pupa.—Unknown. MareriaL.—Grenada, Lake Grand Etang, 4-6 Aug. 1963, O. S. Flint, Jr., 29. Lake Grand Etang, 2 miles west, 4-8 Aug. 1963, O. S. Flint, Jr.. 17 29, 2 larvae. Great River, Balthazar, 8 Aug. 1963, O. S. Flint, Jr., many larvae. St. Vincent, 2c? 1? recorded by Mosely, 1933, p. 47 and verified by Kimmins (pers. comm.). Brotocy.—The larvae have been taken in both a small tumbling mountain brook and a riffle in a large lowland river. Both, however, are clear and rapid with a bottom of rocks and gravel. Family HypRopriLiDAE This family contains the smallest species of the order. Most of the adults are only a millimeter or two long with the largest reaching about five millimeters. The Lesser Antules contain at least 23 species in 8 genera, or 50 percent of the total species in the order from these islands. The larvae undergo hypermetamorphosis in which the first four instars are slender, bear long setae, are free living, and are brief in duration. In the fifth instar, the larvae construct their cases and undergo the greater part of their growth, often changing their shape radically. NO. 3665 CADDISFLIES—-FLINT an Since the cases show good generic differences—except between Leucotrichia and Zumatrichia, and between Hydroptila and Ochro- trichia—they are more useful in the placement of the pupal stage than the pupae themselves, which are only slightly different. In the majority of the genera it is impossible to key the immature stages to species. The larvae and pupae of Bredinia are unknown. Key to Genera LARVAE AND CASES 1. Larvae with abdominal terga bearing sclerites; case flattened and tightly appressed to substrate, never movable. . ... ee Larvae with a sclerite on ninth tergum only; case eros enemed! but carried by larva and attached at pupation . . . es BOS | 2. Abdominal segments 3-6 greatly cscs case Oe domeal with openings ae DOUM Ends! 2 jo. 6 ..« yaa No abdominal segment salieaned case Pacasly Snaviks, flat, ain ein openings .... Sees ss ee 6 ey Alisotrichia 3. Ninth tergum with ationt, euiarged sta ees, es a) > IMA triChia Ninth tergum with only normal hairlike setae. . . . . . . . Leucotrichia 4. Mid- and hindlegs about 3 times as long as forelegs; case silken, anterior end cylindrical, becoming enlarged and compressed posteriorly . . . Oxyethira All legs of about same length; case generally with some sand or organic matter, differently shaped .... . : A ae nei OO 5. Anal prolegs projecting freely from abdomen; case sabular made of BCA Sr es ... . . Neotrichia Anal prolegs fused to sil of apdercene case ome Dae, Geren th ie or 6. Metanotum with anterolateral angle enlarged. . . . . . . . Ochrotrichia Metanotum with anterolateral angle not enlarged. . . . . . . Hydroptila ADULTS PEOLCCUIEDTCSON Gi.” cauc EAMEy dee ct” Seti vote WEG Os )ae) ad Saucy a eee ener ee Ocelliabsent ..... . .. . . Hydroptila 2. Mesoscutellum with a GRATES autre een aera angles! (iy 2%... 3 iNMesoscutellumeentire tr is stills, PNT Aa Sod. Oe ed ao S-eeloretibia with an.apicalspur .).)elgler a ble 4% wot me bh ue are AA Foretibia without an apical spur... . Se Peo ar et a An Gee) 4. Males with basal antennal segment aalaeeedh covering face, with 2 Ocelli 2. 3 ss ... . . Zumatrichia Basal antennal peemieas enodiaed ae 3 econ et As Macao sseeee eee ey 1. 5. Forewing fuscus, with linear, bright green marks. . .. . . Leucotrichia Forewing fuscus, sometines with silvery-gray blotches Ochrotrichia (Metrichia) Game vacdiibias with a@preapical spurs. lye. sods eos da Spacer mh ence yerrd Midtibia without a preapical spur ... . ay Pe ec eey ms 7. Male genitalia not greatly modified, Snepers inte Gna. easily recognizable OS FOG Ss a6 6 6 . . . . . Ochrotrichia (O.) Male genitalia gr andy po cined dagen suell and often difficult to homolo- IZ ed Te cate el ce ah ae el cere eh aeta Ode 32? Alisotrichia 313-152—68——3 34 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 8. Male with basal antennal segment enlarged, covering face, 2 ocelli Alisotrichia Male with antennae unmodified, 3 ocelli . . . .... =... . Bredinia 9. Hindtibia with only a preapical spur ......... ... . Neotrichia Hindtibia with 2 preapical spurs .. . a sh gw oe 6 ey eRe LO) 10. Metascutellum narrow, almost Gnadrane nis ~ 4. « « . » oe Alisotrichia Metascutellum wider, distinctly triangular. . ...... . . Oxyethira Genus Zumatrichia Mosely Zumatrichia Mosely, 1937, p. 187. [Type-species: Zumatrichia filosa Mosely, 1937, by original designation.] The genus has heretofore been known only from several Mexican species. The genus is closely related to Leucotrichia, but the males differ greatly in the modified basal antennal segment, in the possession of only two ocelli, and in a different style genitalia. The immature stages of the genus are herein described for the first time, and as expected they show a close relationship to those of Leucotrichia. The larvae of Zumatrichia are distinguished by the short, broad, setae on the eighth and ninth terga. Key to Species ADULTS Color gray, mottled with grayish green; forewing of male unmodified. Z. antilliensis Color, brown; forewing of male with a basal patch of modified setae. Z. anomaloptera Zumatrichia antilliensis, new species Figures 75-79, 86, 99-103 This species has been the most commonly encountered trichopteran on the island of Dominica, and it has also been taken on Guadeloupe, St. Lucia, and Grenada. It is related to Z. galtena Mosely from Mexico but is distinguished by the dorsal process of the clasper not being digitate apically and by the long hooked ventral process of the aedeagus. Aputt.—Lenegth of forewing 3 mm. Color intermingled patches of eray and gray green. Male genitalia: eight sternum with ventrolateral angles slightly produced; ninth segment with anterolateral angles pro- duced as narrow lobes; cercus, rodlike, with a single long apical seta; clasper with a long basodorsal process, slightly sinuate apically; ventral portion with a slender apicolateral lobe; tenth tergum divided into platelike lateral lobes, rounded apically, with a midventral tooth; aedeagus with a middorsal process apically, a pair of lateral spines, and a pair of appressed, hooked, ventral processes; a complex No. 3665 CADDISFLIES—FLINT 35 Ficures 75-84.—Zumairichia antilliensis, new species, male: 75, genitalia, lateral; 76, genitalia, ventral; 77, genitalia, dorsal; 78, aedeagus, lateral; 79, female, bursa copulatrix, lateral. Z. anomaloptera, new species, male: 80, genitalia, lateral; 81, genitalia, ventral; 82, genitalia, dorsal; 83, aedeagus, lateral; 84, female, bursa copulatrix, lateral. 36 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 at midlength and a basal loop. Female genitalia: eight and ninth segments weakly sclerotized, with short anterolateral rods; bursa copulatrix with basal group attached to apical group by a slender rod, apical group with long lateral supports. Ficures 85-86.—Alisotrichia species 2: 85, larva, dorsal. Zumatrichia antilliensis, new species: 86, larva, dorsal. Larva.—Length to 3 mm. Head, thorax slender, abdominal seg- ments 5-7 greatly distended at maturity. Sclerites pale brown, marked with fuscus. Head unmodified. Legs short and broad; all airs very similar in structure. Meso- and metanota divided mesally. NO. 3665 CADDISFLIES—FLINT 37h Abdomen with tergites dorsally on segments 1-9, 1 small lateral sclerite on segments 2 and 8, 2 sclerites on segments 3-7. Anal proleg with basal segment terete, short; claw sharply decurved. Pura.—Length 3 mm. Mandible sickle shaped, broad, without serrations. Labrum membranous. Hook-plates anteriorly on seg- ments 3-7, posteriorly on 3-5; each plate with many small teeth. Apex of abdomen with a membranous lobe containing genitalia. Casze.—Length 3 mm, width 1% mm. Silken, tightly attached to substrate. Larval case with round anterior and posterior openings. Pupa enclosed in an inner silken cocoon. MatTeErRIAL.—Holotype, male: Dominica, Clarke Hall, 17 April 1964, O. S. Flint, Jr.. USNM type 69892. Allotype, female: same data. Paratypes (many thousands of specimens taken at the following localities, on many dates throughout the year, and by most collectors): Clarke Hall, Fond Figues, Layou Valley, Layou River Mouth, Grand Bay, Rosalie, Cabrit Swamp, Trafalgar, 2.5 and 3.5 miles north of Pont Casse, 1.6 miles west of Pont Casse, 1.3 and 2.2 miles east of Pont Casse. St. Lucia, Cul de Sac River at mile post 9, 29 July 1963, Flint and Cadet, 5o7; Vergallier River near Marquis, 2 Aug. 1963, Flint and Cadet, 2°. Grenada, 2 miles west of Grand Etang, 4-8 Aug. 1963, O. S. Flint, Jr., many o'c'9 9; Balthazar, 7 Aug. 1963, O. S. Flint, Jr., 407%. Guadeloupe, Petit-Bourg, Duclos, March 1966, J. Bonfils, many 7079 ¢. Other: Dominica, Fond Figues, 3 May 1964, O.S. Flint, Jr., larvae, o'9 metamorphotypes; Roseau River, 1 mile above Roseau, 23 July 1963, O. S. Flint, Jr., larvae and pupae; Layou River, 23-25 July 1963, O. S. Flint, Jr., larvae, o 9 metamorpho- types; Springfield, 20-26 July 1963, O.S. Flint, Jr., larvae, co 9 metamorphotypes; R. Laurent, Bells, 21 July 1963, O. S. Flint, Jr., larvae, o& 9 metamorphotypes; Pont Casse, 3.5 miles north, 5 Dec. 1964, P. J. Spangler, larvae and pupae. Brotogy.—This is the commonest species of Trichoptera on the island of Dominica. It is the only species that has successfully adapted to breeding in the large lowland rivers. The flat cases are attached in ereat abundance to the large boulders in the fast-flowing sections of these rivers. A few adults have also been taken near the small streams at high elevations; however, these may well have been carried up by winds from the lowlands. The species seems less abundant on St. Lucia and Grenada. On the latter, Z. antilliensis seems to be replaced in the lowland rivers by Z. anomaloptera. Zumatrichia anomaloptera, new species FiaureEs 80-84 On the basis of head, genitalia, and spurs, this species is a typical member of the genus Zumatrichia; however, it is the only species so far discovered that has the patch of modified setae present on the basal half of the forewing in the male. AputtT.—Length of forewing 3 mm. Color brown; basal half of forewing of male with a large patch of deep brown, scalelike setae. 38 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 Male genitalia: eighth sternum bearing laterally from posterior margin an enlarged setae from a large base, and ventrally a pair of caliper- like lobes; ninth segment with anterolateral angle broad; cercus rod- like with a single long apical seta; clasper with a short, rodlike baso- dorsal process, ventral lobes rounded; tenth tergum divided into flat lateral plates, rounded apically, with a subapical ventral tooth; aedeagus with usual mesal and basal structures, apically with a large number of slender spines. Female genitalia: eighth and ninth segments with lateral rods; bursa copulatrix complex, basal group supported by a slender rod from apical group that has short lateral supports. LARVA, PUPA, AND cASE.—Not different from Z. antilliensis. MaTERIAL.—Holotype, male: Grenada, Balthazar, 7 Aug. 1963, O. S. Flint, Jr.. USNM Type 69893. Allotype, female: same data. Paratypes: same data, many o'o'? ?. St. Lucia, R. Galet, south of Dennery, 1 Aug. 1963, Flint and Cadet, 1; Vergallier River, near Marquis, 21 July 1963, Flint and Cadet, many 3h? 9; same, but 2 Aug. 1964, 29° 49. Dominica, Clarke Hall, 1-10 March 1965, W.W. Wirth, 1. Other: Grenada, Great River, at Balthazar, 8 Aug. 1963, O. S. Flint, Jr., larvae, 7 o' 2? 9 metamorphotypes. Brotogy.—The immature stages were found on large rocks in a large lowland river on Grenada. It thus seems that it has identical breeding habits to the preceding species, but that it has replaced Z. antilliensis in the lowlands of Grenada and St. Lucia. Genus Leucotrichia Mosely Leucotrichia Mosely, 1934, p. 157. [Type-species: Leucotrichia melleopicta Mosely, 1934, by original designation.] This and the preceding genus are closely related, but Leucotrichia is found over most of North America as well as Central America. The male of Leucotrichia may be recognized by the unmodified antennae and presence of three ocelli. The larvae are also similar in the two genera, but those of Leuco- trichia lack the modified setae on the eighth and ninth terga. The immature stages are described in Ross (1944) and Flint (1964b), and so are not treated in detail here. Leucotrichia sarita Ross Fiaures 87-91 Leucotrichia sarita Ross, 1944, p. 274. This species, described from Texas, is common in Mexico and Central America. There are some minute differences in the genitalia between the insular and mainland populations, but these are no greater than differences found on the mainland. The description is based on Grenadan examples. NO. 3665 CADDISFLIES—FLINT 39 Apvutr.—Length of forewing 3 mm. General color fuscus, with bright green linear marks on forewings, tegulae, and head; antennae alter- nating black and white. Male genitalia: eighth sternum produced into an acute angle laterally, broadly U-shaped ventrally; ninth seg- ment oblique, open anteroventrally, with a row of stout setae along posterior margin; claspers fused mesally, elongate, with a subapical seta dorsally, dorsally with a pointed mesal structure, partially enclosed by claspers that articulate basally with a linear lateral sclerite; tenth tergite strongly sclerotized, produced into a ventro- lateral point; aedeagus with a pair of small apicolateral rods, a central dome with basolateral rods, and a basal loop. Female geni- talia: eighth and ninth segments with anterolateral rods, posterior margin of eighth segment with a row of large setae; bursa copulatrix without an apical complex of supporting sclerites, but with a large complex basal group. Larva.— Unknown (from Grenada). Pupa.—Length 2.5 mm. Mandibles sickle shaped, inner surface smooth. Hook-plates present anteriorly on segments 3-7, posteriorly on 3-5, each with many small hooks. Casr.—Length 4 mm, width 2 mm. Silken, oval, domed, attached to substrate. Pupal case with a tight inner cocoon. MatreriAu.— Grenada, Beausejour River, 2 miles west of Grand Etang, 4-8 Aug. 1963, O. S. Flint, Jr..many 7 &@ 2? 9, o& 2 metamorphotypes. Brotogy.—The pupae were taken on rocks in a cascade of a small mountain brook several feet wide by a few inches deep. The adults were attracted to a blacklight placed nearby. Genus Alisotrichia Flint Alisotrichia Flint, 1964b, p. 46. [Type-species: Alisotrichia hirudopsis Flint, 1964b, by original designation.] Known from Puerto Rico and Jamaica previously, five species are assigned to the genus from Dominica. The typical males have enlarged basal antenal segments that cover the face, only two ocelli, and a spur count of 0, 2, 4. In addition to two species belonging to this typical group, I am assigning three species to this genus that belong to a rather different group. The males of this second group have unmodified antennal segments, three ocelli, and a spur count of 0, 3, 4. The thoracic structure is very similar in the two groups, however, and both have extremely modified genitalia. The female seventh tergum in this second group bears two straplike sclerites, rather than a single tri- angular one, but both have a simple ringlike bursa copulatrix. The larvae assumed to belong to this second group are very similar to the typical ones but also show several differences. Pending a total review 40 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 tt “ ems we - we ' s ” \ ‘ 96 ‘ “ 94 Ficures 87-98.—Leucotrichia sarita Ross, male: 87, genitalia, lateral; 88, genitalia, dorsal; 89, genitalia, ventral; 90, aedeagus, lateral; 91, female, bursa copulatrix, lateral. Aliso- trichia orophila, new species, male: 92, genitalia, lateral; 93, genitalia, dorsal; 94, eighth sternum, ventral. 4. lobaia, new species: 95, male genitalia, dorsal; 96, male eighth sternum, ventral; 97, male genitalia, lateral; 98, female genitalia, dorsal. NO. 3665 CADDISFLIES—FLINT 4] of this group, I prefer to leave both groups in the genus Alisotrichia, rather than establish a second genus. Key to Species LARVAE 1. Abdominal terga with large, elongate-oval setae... .. . . A. species 2 Abdominal terga with short, truncate setae... ........e84.8-. 2 2. First abdominal tergum with a central dark mark ... .. . A. orophila First abdominal tergum without a central mark. ...... A. species 1 ADULTS ieee Madleg: without a preapical spur . . 6. . 6. 6 6 0 ele os Be oes ee Madierawithy a-preapiCAl Spur sts vwlece oe) ce) a) es ees ay) eel aah 3 2. Highth sternum of male with a bifurcate ventrolateral lobe; forewing gray, WAGHEWIMILENPALGHES 5 0s os, wt, tee lo oe gee ener A. orophila Highth sternum with a rounded dorsolateral lobe and a pointed ventrolateral ROHS MMete Ys ciat her) “al snes Cd cath Ace pie 2) a oar crdeiawo ye! Soko acne A. lobata 3. Claspers fused into a single median rodlike process; forewings fuscus with brighitvereentspOtstitms 1 yioceco sense ne eitiey ate A. septempunctata Claspers not fused: without green.spots .. . . . . « « © « «ss 6 « 4 4. With an elongate process extending ventrad from venter of tenth segment A. dominicensis Without an elongate midventral process from tenth sternum .. A. wirthi Alisotrichia orophila, new species Figures 92-94, 105 The species seems to be closely related to the following from which it differs strikingly in the shape of the eighth sternum, which possesses a long dorsolateral spine and a bifurcate apicolateral hook. Avu.tt.—Length of forewing 1.5 mm. Color grayish, forewing with alternating bands of silver and fuscus hair. Structure of head and ap- pendages conforming closely to type species. Male genitalia: eighth sternum bearing a long dorsolateral spine, apicolateral angle with a strong mesally directed, shallowly bifurcate hook; ninth segment with extremely long anterolateral rods, with a dorsomesal sclerite; tenth tergum with paired dorsal, straplike sclerites, apex with a semimem- branous process; aedeagus tubular, with conical base. Female genitalia: seventh tergum narrowly triangular; eighth and ninth segments with long anterolateral rods; apparently the same as following species. Larva.—Length 1.5 mm. Sclerites pale brown. Structurally very similar to larva of type species, but differing in the possession of a ter- gite on first abdominal segment. Pupa.—Length 1.2 mm. Mandibles with a short apical blade, lack- ing mesal serrations. Hook-plates anteriorly on segments 3-7, pos- teriorly on 3-5; each with many hooks. 42 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 Ficures 99-105.—Zumatrichia antilliensis, new species, larva: 99, head, anterior; 100, mandible, dorsal; 101, labrum, dorsal; 102, ninth, eighth, and seventh abdominal terga, dorsal; 103, thorax and first abdominal tergum, dorsal. Alisotrichia species 1: 104, larval thorax and first abdominal tergum, dorsal. 4. orophila, new species: 105, larval thorax and first abdominal tergum, dorsal. NO. 3665 CADDISFLIES—FLINT 43 Casr.—Width 1 mm, length 2 mm. Oval, silken, appressed to substrate. Consisting of an outer flange attaching inner cocoon to rock. MaTeERIAL.—Holotype, male: Dominica, D’leau Gommier, 15 Feb. 1965, W. W. Wirth, USNM type 69894. Allotype, female: Pont Casse, 2.5 miles east, 16 Jan. 1965, W. W. Wirth, stream margin. Paratype: Pont Casse, .4 miles east, 6 May 1964, O. S. Flint, Jr., 1c. Other: D’leau Gommier, 27 April 1964, O. S. Flint, Jr., 1 larva, 2 pupae, 1o7 19 metamorphotypes. Brotoay.—The immatures of this species were collected on large stones kept wet by the cascade of a small mountain stream at the place where it emerged from the forest. The habitat is very similar to that of the type-species on Puerto Rico. Alisotrichia lobata, new species FiaurEs 95-98 The males of this species are easily recognized by the large dorso- lateral lobe of the eighth sternum and the long lateral process of the ninth segment. Aputt.—Length of forewing 1.5 mm. Color grayish, forewing with alternating bands of silver and fuscus hairs. Corresponding closely to type-species in structure of head and appendages. Male genitalia: eighth tergum narrow, sternum greatly prolonged apicoventrally, with a large dorsolateral lobe, apicoventral angles produced into a sharp spine, midventrally with narrow incision; ninth segment with long, anterolateral, rodlike processes, divided dorsally on midline, with posterolateral swordlike processes; tenth tergum with paired, dorsal, straplike sclerites, with ventrolateral margin sclerotized, ventrally with a complex of sclerites; aedeagus tubular, with base slightly en- larged. Female genitalia: seventh tergum with a narrow, apicomesal, triangular sclerotization; eight and ninth segments with long antero- lateral rods; tenth segment with long apical papillae. LARVA, PUPA, AND CASE.—Not known for certain, but see following species. MateriAu.—Holotype, male: Dominica, Clarke Hall, malaise trap, 11-20 Jan. 1965, W. W. Wirth, USNM Type 69895. Allotype, female: Clarke Hall, cocoa trail, 18 Jan. 1965, W. W. Wirth. Paratypes: same data as allotype, 29 ; Fond Figues, 3 Feb. 1965, W. W. Wirth, 1c. Brotogy.—The few adults of this species have been taken in proximity to the larger lowland rivers. Alisotrichia species 1 Figure 104 There is a single larva of a species closely related to A. orophila in the collections. The larva is a bit larger and the thoracic notae 44 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 and abdominal terga posses a few more setae than those of the larvae of A. orophila. The first abdominal tergum lacks the central dark mark, and the pores on the following segments are in front of the black mark rather than surrounded by it as in A. orophila. The data with the specimen is: ‘‘Pont Casse, .5 miles south, 22-24 July 1963, O. S. Flint, Jr.”’ Because this is a small high altitude stream rather than a lowland stream, I hesitate to definitely attribute this larva to A. lobata, to which it may well belong. Alisotrichia dominicensis, new species Figures 110-114 This species is easily recognized by the long ventral process and the middorsal sclerite of the tenth tergum and the bilobed claspers. Avutt.—Length of forewing 2 mm. Color fuscus; antennae yellow- ish, face and mesonotum with white hairs, forewing with a white band at midlength and basally. Spurs, 0, 3, 4. Basal antennal segment not modified. Ocelli 3. Maxillary palpus with 2 basal segments minute, third, fourth, and fifth segments elongate, subequal. Male genitalia: seventh sternum with an apicomesal process; eighth tergum broad, posterior margin of sternum slightly produced laterally, almost squarely truncate lateroventrally; ninth segment produced antero- laterally, with margins heavily sclerotized; claspers arising ventro- laterally from ninth segment, bilobed, semimembranous; tenth tergum with a mesal straplike sclerite upturned apically on dorsum, ventrally with an elongate mesal structure extending directly ventrad; aedeagus mostly membranous, with basal and apical cylindrical portions separated by a constriction, apical portion with indistinct, short, internal spines. Female genitalia: sixth sternum with a small apicomesal point; seventh sternum with posterior margin slightly bilobed, tergum with a pair of straplike sclerotizations ending in an oval mark; eighth and ninth segments with long lateral rods, eighth with a row of setae along posterior margin; bursa copulatrix an internal sphere. LARVA, PUPA, AND cASE.—Not known for certain, but see Aliso- trichia species 2. MareriaLt.—Holotype, male: Dominica, Pont Casse, 2.2 miles east, 7 May 1964, O. S. Flint, Jr.. USNM Type 69896. Allotype, female: Pont Casse, 2.5 miles east, 16 Jan. 1965, W. W. Wirth. Paratypes: same data as holotype, but 3 May 1964, 19 ; same data as allotype, 2? ; Fond Figues, 25 Jan. 1965, W. W. Wirth, 27; Morne Nicholls, 9 Nov. 1964, P. J. Spangler, 19. Brotocy.—The adults were taken near either a small cascading mountain brook or a larger lowland river also with some cascades. NO. 8665 CADDISFLIES—FLINT 45 Ficures 106-118.—Alisotrichia sepiempunctata, new species, male: 106, genitalia, lateral; 107, genitalia, ventral; 108, genitalia, dorsal; 109, aedeagus, dorsal. 4. dominicensts, new species: 110, aedeagus, lateral; 111, male genitalia, lateral; 112, male genitalia, ventral; 113, male genitalia, dorsal; 114, female genitalia, dorsal. A, wirthi, new species, male: 115, genitalia, lateral; 116, genitalia, ventral; 117, genitalia, dorsal; 118, aedeagus, lateral. 46 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 125 Alisotrichia wirthi, new species Fiaures 115-118 This species is closely related to the preceding, from which it is easily separated by the very different claspers and tenth tergum. Aputt.—Length of forewing 2.56 mm. Color in alcohol uniformly fuscus. Basal antennal segments unmodified. Ocelli 3. Maxillary palpus with 2 basal segments very short, apical segments long, each slightly longer than preceding. Spurs 0, 3, 4. Abdominal sterna lacking processes. Male genitalia: eighth tergum broad, sternum produced ventrally, truncate in ventral aspect; ninth segment quad- rate, with a middorsal point, a small lateral flap, and a ventral rodlike support; claspers with a ventrolateral lobe, and a flattened mesal lobe bearing a stout setae from its dorsal margin; tenth tergum con- sisting of paired elongate dorsal sclerites, a triangular lateral sclerite, and flattened ventromesal sclerites lying within the mesal lobes of the claspers; aedeagus with cylindrical basal and apical portions separated by a constriction, apical portion with an indistinct internal spine, with outer surface sclerotized in basoventral region. LARVA, PUPA, AND CASE.—Not known. MateriALt.—Holotype, male: Dominica, Fond Figues, 13 March 1965, W. W. Wirth, USNM Type 69897. Paratype: same, but 6 April 1964, O. S. Flint, Jr., 1c. Brotogy.—Both specimens were taken at light near a large, clear, fast-flowing lowland river. Alisotrichia septempunctata, new species Figures 106-109 Both the coloration of this species and the unique structure of the genitalia are diagnostic. Avutt.—Length of forewing 2.5 mm. Black, each forewing with 3 bright green spots, mesonotum with a green mesal spot. Basal antennal segment unmodified. Ocelli 3. Maxillary palpus with 2 basal segments very short, third and fifth long, subequal, fourth % length of third. Male genitalia: no processes from sterna; eighth tergum broad, sternum produced ventrolaterally, posterior margin broadly V-shaped in ventral aspect; ninth segment with ventrolateral angles prolonged anteriorly, with posterior margin produced into an up- turned, swordlike process; claspers completely fused, developed as a long, slender rodlike process from apex of ninth sternum; a terete, semimembranous process arising mesad of lateral process of ninth segment; tenth tergum flattened, developed as broad lateral plates with an apicoventral point, dorsally with a pair of mesal parenthesis- like sclerites; aedeagus with apical and basal cylindrical portions NO. 3665 CADDISFLIES—FLINT 47 separated by a constriction, apical portion with several pairs of elongate internal spines. LARVA, PUPA, AND CASE.— Unknown. MarertaL.—Holotype, male: Dominica, Pont Casse, 2.2 miles east, 14 April 1964, O. S. Flint, Jr.,. USNM Type 69898. Biotocy.—The only known specimen was taken at a black light at a small tumbling brook. Alisotrichia species 2 FiaureE 85 I am here describing the larvae of a species that is clearly related to Alisotrichia although they do possess a number of unique charac- teristics. The possession of numerous small intercalary sclerites on the thorax and abdomen, and the large, elongate-oval setae are very distinctive. It seems quite probable that these are the larvae of one of the last three species. Larva.—Length 3.5 mm. Sclerites brown. Slightly flattened, abdomen gradually tapering toward posterior. Structurally much like the larva of type-species, differing as follows. Thoracic notae and abdominal tergites bearing long, broad, pointed, black setae; pro- pleuron with 1, meso- and metapleura with 2 of these setae. Meso- and metanota anteriorly and abdominal terga posteriorly with a broad border of sclerotized spots. Small intercalary sclerites present between segments from the mesonotum to the eighth segment. Abdomen with 9 tergites; that of first segment barely different from those of segments beyond. MartertaL.—Dominica, R. Laurent, 21 July 1963, O. S. Flint, Jr., 1 larva. Geneva Estate, 9 Dec. 1964, P. J. Spangler, 1 larva. Fond Figues R., 9 Feb. 1965, W. W. Wirth, 1 larva. BioLtocy.—The larvae were collected from boulders in rocky sections of swiftly flowing lowland rivers. Genus Neotrichia Morton Neotrichia Morton, 1905, p. 72. [Type-species: Neotrichia collata Morton, 1905, by monotypy.] The genus contains numerous species, all of New World distribution. The species are all very small, generally being less than two milli- meters long. The larvae were described by Flint (1964b) and Ross (1944). They construct small, cylindrical cases made of very small sand grains. 48 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 Key to Species ADULT Male genitalia with subgenital plate elongate, directed posteriad; female eighth sternum with a large heart-shaped midventral lobe ....... N. iridescens Male subgenital plate with apex slender, angled sharply ventrad; female eighth sternum without midventral lobe . (2.02 & .a!sa = ale a oe N. corniculans Neotrichia iridescens Flint Figures 119-123 Neotrichia tridescens Flint, 1964b, p. 51; 1968a, p. 37. Originally described from Puerto Rico, this species has since been recorded from Jamaica and is now recorded from Dominica and St. Lucia. The Dominican examples agree closely with the type except that the darker ventral processes of the subgenital plate are nearer its apex. The St. Lucian specimens seem to have a narrower and more twisted apex of the aedeagus and the dorsolateral process of the ninth segment has both dorsal and ventral points apically. Aputt.—Length of forewing 1.5-2 mm. Color mottled grayish brown; forewing of female with a patch of purplish iridescent scales centrally. Male genitalia: ninth segment produced basoventrally, posterior margin with a dorsolateral process whose tip is developed into a dorsal (and sometimes ventral) point; clasper elongate and rectangular in lateral view, in ventral view with apex obliquely truncate; bracteole slender, arising near base of clasper; subgenital plate elongate, slender, with dark, ventrally directed lobes at mid- length; aedeagus with a membranous process arising near midlength, and a more strongly sclerotized process near apex, central tube trough shaped apically. Female genitalia: eighth sternum with anterior margin heavily sclerotized, with a pair of short anterolateral rods, mesally with a heart-shaped lobe, and posteriorly with a rec- tangular sclerite, posterior margin with 3 pairs of setae; bursa copula- trix elongated apically, basolateral angles rodlike, with a basomesal complex. Ficures 119-135.—Neotrichia iridescens Flint: 119, male genitalia, ventral; 120, aedeagus, dorsal; 121, male genitalia, lateral; 122, female eighth sternum, ventral; 123, bursa copu- latrix, ventral. NV. corniculans new species: 124, male genitalia, lateral; 125, male geni- talia, dorsal; 126, male genitalia, ventral; 127, aedeagus, dorsal; 128, female eighth ster- num, ventral; 129, bursa copulatrix, ventral. Bredinia dominicensis, new species: 130, male genitalia, lateral; 131, male genitalia, dorsal; 132, male genitalia, ventral; 133, aedeagus, dorsal; 134, female genitalia, ventral; 135, bursa copulatrix, ventral. NO. 3665 131 313-152—68———-4 CADDISFLIES—FLINT 49 50 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 Larva.—Length to 2 mm. Nearly cylindrical in outline. Sclerites pale yellowish. Head with a spinelike process behind eyes. Apical abdominal segments with many long, dark setae; anal prolegs elongate. Casr.—Nearly cylindrical, posterior slightly flattened and tapered. Made of very small sand grains. Materiau.—Dominica, Clarke Hall, 11-20 Jan. 1965, W. W. Wirth, 107; same, but 21-29 April 1964, O. S. Flint, Jr., 1c? 19; Pont Casse, .4 miles east, 25 June 1964, O. S. Flint, Jr., 1 pupa; Pont Casse, 1.6 miles west, 24 April 1964, O. 8. Flint, Jr., 2 cases. St. Lucia, Vergallier River, near Marquis, 31 July 1963, Flint and Cadet, 6c 39, 3 larvae, 2 pupae. Brotocy.—The immature stages are found on rocks in the more slowly flowing sections of both the larger and small rivers. Neotrichia corniculans, new species Fiaures 124-129 This species appears to be related to the group of species named Dolotrichia by Mosely. From all known species it is easily recognized by the hornlike shape of the tenth tergite and the shape of the claspers. Aputt.—Length of forewing 1.5 mm. Color silvery gray. Male genitalia: ninth segment rounded anteriorly, posterior margin devel- oped into a thin lateral shelf that bears two pointed processes, dorsal one longest; tenth tergum developed as a pair of curving hornlike processes directed first laterad then posteriad; subgenital plate long, attenuate, and angled sharply ventrad; clasper elongate, slightly narrowed apically; aedeagus with apical half divided into two proc- esses, longer one twisted apically and with a bifid tip, a whiplike process arising at midlength. Female genitalia: eighth sternum with a dark lateral mark and a small raised central structure shaped like an arrowhead; bursa copulatrix elongate with a basal opening and two pairs of lateral pouchlike invaginations. LARVA, PUPA, AND CASE.— Unknown. MartertaLu.—Holotype, male: Dominica, D’leau Gommier, 16 March 1965, W.W. Wirth, USNM Type 69899. Allotype, female: same data. Paratypes: same data, 19 ; same, but 15 Feb. 1965, 307 3 9 ; Pont Casse, 1.0 mile east, 29 Jan. 1965, W.W. Wirth, 19. Brotocy.—Little is known of the biology of this species. The adults have been taken only at higher elevations on the island. Bredinia, new genus Three ocelli. Spurs 0, 2, 4. Mesoscutellum divided by a transverse suture, with a broad sclerite along posterior margin. Metascutellum NO. 3665 CADDISFLIES—FLINT 5l as wide as scutum, short and rectangular. Wings narrow, evenly acuminate. Type-species: Bredinia dominicensis, new species. The relationship of the genus is rather obscure. It would seem to be most closely related to Neotrichia as is shown by the presence of ocelli, minute size, and wing shape. The spur count is the same as is found in Mayatrichia, a genus closely related to Neotrichia; however, the structure of the thoracic notae (almost exactly as shown in Ross, 1944, fig. 438) is totally different from Neotrichia or Mayatrichia in which the mesoscutellum is entire and the metascutellum trianguloid. It may well be that the genus is related to Alisotrichia as is suggested by the structure of the thorax and dorsolateral rods of the male ninth segment. The discovery of the larval stages will be necessary to settle the exact relationship of the genus. It is with great pleasure that I name this genus for Mr. J. Bruce Bredin. Bredinia dominicensis, new species Figures 130-135 Since this is the only known species in the genus, it is not possible to point out specific characters. Apu.t.—Leneth of forewing 1.5 mm. Color silvery gray, forewing with bands of white hair apically and basally. Male genitalia: eighth sternum divided midventrally; ninth segment narrow, oblique, lateral halves divided ventrally by the claspers, anterior margin with long, slender supports dorsolaterally; tenth tergum a large membranous lobe; clasper rather quadrate, with narrow dorsal and ventral ridges, a long seta from ventral ridge; subgenital plate elongate, rectangular, slightly widened apically; aedeagus tubular, apex flattened, tridentate, with a central tubule in apical fourth. Female genitalia: seventh segment with apical margin squarely truncate; eighth segment simple, with anterolateral rods, and a row of setae along posterior margin; ninth segment elongate with anterolateral rods; tenth segment trianguloid, with apical papillae; bursa copulatrix with a long baso- mesal process and a pair of basolateral tubules, a central complex, apically with a pair of dark oval sclerites. LARVA, PUPA, AND CASE.—Unknown. MateErr1aLt.—Holotype, male: Dominica, Hodges River mouth, swamp forest, 27 Feb. 1965, W. W. Wirth, USNM Type 69900. Allotype, female: same data. Paratypes: same data, 10°; Fond Figues, 3 Feb. 1965, W. W. Wirth, 10; same, but 18 March 1965, 2°. Biotogy.—The adults have been taken only near the larger lowland rivers. 52 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 Genus Oxyethira Eaton Oxyethira Eaton, 1873, p. 143. [Type-species: Hydroptila costalis Curtis, 1834, by original designation.] Oxyethira is a genus of worldwide distribution, containing numerous species in most regions. Each of the Antillean islands support several species. The immature stages and the cases they make are very similar throughout the genus. Nielsen (1948) has given an excellent account of the morphology and biology of the immature stages of the type- species. Key to Species ADULTS 1. With aedeagus, etc.: males. ...... 2 Without aedeagus: females. ...... 5 2. Ninth sternum elongate, scooplike, apex pide 3 Ninth sternum not elongated. ....... vane 4 3. Ninth sternum deeply and widely divided apical 5 Ges £0); azteen Ninth sternum with a short, narrow apical division. . . . . .O. janella 4. Eighth segment dorsally with a sabre-like process, with a broad lateral flap. O. cirrifera Eighth segment with a pair of short, flattened dorsal lobes, and erect dorso- laterallobe. ... . os a a 2 Os tega 5. Bursa copulatrix with pated: fone? proaliees antcr ior ad posterior supports. O. janella Bursa without rodlike supports 21. . 6 % 4.5. 0s ou ee Sa ge? ene 6 6. Eighth sternum with sclerotized apicolateral invaginations . . O. cirrifera Highth sternum without invaginations. . .. . aes Pets 7. Bursa rounded basally, shield shaped, with partie apical lobes . .O. tega Bursa angulate basolaterally and mesally, apical lobes not earlike. O. azteca Oxyethira janella Denning Fiauress 136-137, 148 Oxyethira janella Denning, 1948, p. 397.—F lint, 1968a, p. 42. Oxyethira neglecta Flint, 1964b, p. 57; 1968a, p. 42. This species appears to be the most widespread Antillean caddisfly. It has been found on Jamaica, Puerto Rico, Dominica, St. Lucia, Ficures 136-149.—Oxyethira janella Denning, male genitalia: 136, lateral; 137, dorsal. O. azteca Mosely, male genitalia: 138, lateral; 139, dorsal. O. cirrifera Flint, male geni- talia: 140, lateral; 141, ventral. O. tega Flint, male genitalia: 142, lateral; 143, ventral. O. cirrifera Flint: 144, aedeagus, lateral. O. tega Flint: 145, aedeagus, dorsal; 146, female genitalia, ventral. O. cirrifera Flint: 147, female genitalia, ventral. O. janella Denning: 148, female genitalia, ventral. O. azteca Mosely: 149, female genitalia, ventral. 53 CADDISFLIES—FLINT NO. 3665 “a 54 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 | | Grenada, in Florida, and in Central America. It is closely related to — O. puertoricensts Flint and O. azteca Mosely. From both it differs in the male sex in the long narrow ninth sternum. Aputt.—Length of forewing 2-3 mm. Color various shades of brown and white. Seventh sternum with a small spine posteromesally. Male genitalia: eighth segment surrounding most of genital capsule, divided ventromesally, with dorsolateral lobe surpassing ventro- lateral lobe; ventral part of ninth segment narrow, scooplike much surpassing eighth segment, dorsal structure Y-shaped; aedeagus a single spinelike structure resting in fork of the Y. Female genitalia: eighth sternum with posterior margin bilobate; bursa copulatrix with anterior processes twice length of bursa. LarvA, PUPA, CASE.— Unknown. MatTeErtAL.—Dominica, Clarke Hall, 11-20 Jan. 1965, W. W. Wirth, 2 39; same, but 21-30 Jan. 1965, 1c’ 49 ; same, but 27 Jan. 1965, J. F. G. Clarke, 2 9 ; same, but 1-10 Feb. 1965, W. W. Wirth, 19; same, but 11-20 Feb. 1965, 2¢ 13 9 ; same, but 18 Feb. 1964, D. F. Bray, 1? ; same, but 21-28 Feb. 1965, W. W. Wirth, 29; same, but 25 Feb. 1964, D. F. Bray, 49; same, but 1-10 March 1965, W. W. Wirth, 69; same, but 11-20 March 1965, 39; same, but 21-30 March 1965, 39 ; same, but 10-20 April 1964, O. 8S. Flint, Jr., 129 ; same, but 21-29 April 1964, 7c 619 ; same, but 1-10 May 1964, 1 19; same, but 11-20 May 1964, 3o° 199 ; same, but 1-15 June 1964, 11 9 ; same, but 15-30 June 1964, 11¢ 1069; Layou Valley, 23-25 July 1963, O. S. Flint, Jr., 19; Springfield Estate, 20-26 July 1963, O. S. Flint, Jr., 59; Fond Figues, 13 March 1965, W. W. Wirth, 2° 329 ; same, but 6 April 1964, O. S. Flint, Jr., 1c; same, but 7 May 1964, 1c 29; Grand Bay, 13 March 1964, D. F. Bray, 19 ; same, but 13 April 1964, O. S. Flint, Jr., 39 ; Pont Casse, 1.6 miles west, 24 April 1964, O. S. Flint, Jr., 19 ; same, but 28 April 1964, 19 ; same, but 19 May 1964, 107 49; same, but 16 June 1964, 2o7 49? ; Pont Casse, 3.5 miles north, 5 Dec. 1964, P. J. Spangler, 39 ; Pont Casse, 1.3 miles east, 10 May 1964, O. S. Flint, Jr., 19; same, but 12 May 1964, 1? ; Pont Casse, 2.2 miles east, 1 May 1964, O. S. Flint, Jr., 49; same, but 19 June 1964, 109; Pont Casse, 3 miles east, 13-16 Oct. 1966, A. B. Gurney, 1c 39; Pont Casse, .4 mile east, 6 May 1964, O. S. Flint, Jr., 29; Sylvania, 23 Jan. 1965, W. W. Wirth, 70 149. St. Lucia, Vergallier River, near Marquis, 31 July 1963, Flint and Cadet, 19; Cul de Sac River, at mile post 9, 27 July 1963, Flint and Cadet, 59. Grenada, 2 miles west of Grand Ktang, 4-8 Aug. 1963, O. S. Flint, Jr... 7799. Brotogy.—Although the immature stages of this species have not been taken, they will probably be found in most flowing water habitats. Adults have been attracted to lights at most elevations on many of the islands. Oxyethira azteca (Mosely) FIGurEs 138-139, 149 Lozotrichia azteca Mosely, 1937, p. 165. Oxyethira azteca (Mosely).— Ross, 1944, p. 295. This species described from Mexico, where it is quite common, will undoubtedly be found to be widely distributed in Central and northern South America. It is here recorded from Grenada for the first time. NO. 3665 CADDISFLIES—FLINT 55 The species is related to O. janella but is easily recognized by the more deeply divided ninth sternum and the very different dorsal processes of the same segment. Apu.t.—Length of forewing 2 mm. Color various shades of brown and white. Male genitalia: eighth segment divided midventrally and dorsally, posterolateral margin bilobate; ninth sternum produced posteriorly, deeply divided apically, dorsally giving rise to a pair of curving rods laterad to the aedeagus, each with an apical spine, ventrad to aedeagus a mesal, pointed structure; aedeagus tubular, basally ex- panded. Female genitalia: eighth sternum with posterior margin slightly bilobate, with a pair of ovate depressions; bursa copulatrix with posteriolateral supports, with basal portion angulate, complex, with a large basal membranous sac. Larva, PupPA, cAsE.—Not known for certain. MareriAL.— Grenada, 2 miles west of Grand Etang, 4-8 Aug. 1963, O. S. Flint, Jr., 24o7 119. Balthazar, 7 Aug. 1963, O. S. Flint, Jr.,107 29. Brotocy.—A number of typical larvae and pupae of this genus were taken on rocks in the small brook two miles west of Grand Etang and may belong to either this or the preceding species. The stream is about three feet wide by six inches deep on the average, clear, and tumbling over bedrock and boulders. Oxyethira cirrifera Flint Figures 140-141, 144, 147 Oxyethira cirrifera Flint, 1964b, p. 57; 1968a, p. 42. The species was described from Puerto Rico and has since been found on Jamaica, and now Dominica. It is related to the following species, from which it differs most noticeably in the structure of the eighth segment and aedeagus. Aputt.—Length of forewing 2 mm. Color pale brown with some darker flecks. Male genitalia: eighth tergum produced laterally into a long saber-like process, sternum produced dorsolaterally in a broad flap; clasper dark, tightly attached to ninth sternum, quadrate in ventral aspect; subgenital plate C-shaped in lateral aspect, in ventral aspect produced into 2 submesal points; an elongate, seta- bearing rod dorsal of clasper; aedeagus with a flattened process arising at midlength and twisted around central tube 1% times; central tube attenuate beyond origin of process, tip slightly twisted. Female genitalia: eighth sternum produced as a semicircular flap mesally, apicolaterally with sclerotized, pouchlike invaginations; bursa copulatrix short with a pair of jawlike apical processes. Larva, PUPA, CASE.— Unknown. MarteriaLt.—Dominica, Cabrit Swamp, 23 Feb. 1965, W. W. Wirth, 129; same, but 18 June 1964, O.S. Flint, Jr.,207 29. 56 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 Brotocy.—The species has been taken on the island of Dominica only near a large lowland swamp with slowly flowing streams. It has been found, however, near fast waters on other islands. Oxyethira tega Flint Ficures 142-143, 145-146 Oxyethira tega Flint, 1968a, p. 44. This species recently described from Jamaica is here recorded from Dominica. The species is easily distinguished from the preceding by the shape of the eighth segment, claspers, and aedeagus. Apvutt.—Length of forewing 2—2.5 mm. Color pale brown, with darker flecks. Male genitalia: eighth segment with a pair of flat dorsomesal lobes, and an erect dorsolateral lobe; anterior margin of ninth segment developed into a rounded lobe; claspers forming a ribbon-like band along ventral margin of ninth segment; subgenital plate developed into a triangular plate beneath aedeagus in ventral aspect, with lateral processes surrounding aedeagus, and developed into dorsal points; two pairs of membranous rods above clasper; aedeagus with apical half divided into 2 processes, 1 of which bears an internal tubule. Female genitalia: eighth sternum slightly bilobate; ninth sternum sclerotized laterally; bursa copulatrix shield shaped with a central opening, and a pair of apicolateral earlike flaps. LARVA, PUPA, CASE.— Unknown. Marerrau.—Dominica, Freshwater Lake, 21 Jan. 1965, W. W. Wirth, 1c; same, but 22 Feb. 1964, D. F. Bray, 5oc' 79; Sylvania, 23 Jan. 1965, W. W. Wirth, 3o° 39; Clarke Hall, 21-31 Jan. 1965, W. W. Wirth, 1; same, but 1-10 Feb. 1965, 19 ; same, but 11-20 Feb. 1965, 2c° 49 ; same, but 21-28 Feb. 1965, 1c’; same, but 1-10 March 1965, 1°; same, but 21-31 March 1965,29719 ; same, but 21-29 April 1964, O. S. Flint, Jr., 19; same, but 15-30 June 1964, 49° 49; Mannett Gutter, 10 March 1965, W. W. Wirth, 19 ; Grand Bay, 13 April 1964, O. S. Flint, Jr., 1o7; Fond Figures, 13 March 1965, W. W. Wirth, 380° 29. Brotocy.—The adults have been taken at light both near the larger lowland rivers, and a highland lake. Genus Hydroptila Dalman Hydroptila Dalman, 1819, p. 125. [Type-species: Hydroptila tineoides Dalman, 1819, by monotypy.] The genus Hydroptila has been found in all the faunal realms of the world. One or two species have been encountered on each of the Antillean islands that have been collected in depth. The immature stages have been described in detail by Nielsen (1948). They inhabit in the final instar a “purse-case” made of silk, often incorporating sand and plant matter. NO. 3665 CADDISFLIES—FLINT OF Key to Species ADULTS Clasper of male in lateral aspect very narrow, developed into an apicodorsal point; eighth tergum of female slightly less produced than sternum, trilobate. H. grenadensis Clasper broader, widening regularly from base to apex; eighth tergum deeply ange rectangularly “emarpinate. 2) ss, soi jelis Oe eh els H. antilliarum 153 157 Ficures 150-158.—Hydroptila antilliarum, new species: 150, male genitalia, ventral; 151, male genitalia, lateral; 152, aedeagus, lateral; 153, female eighth sternum, ventral; 154, bursa copulatrix, ventral. H. grenadensis, new species: 155, male genitalia, ventral; 156, male genitalia, lateral; 157, aedeagus, lateral; 158, female eighth sternum, ventral. Hydroptila antilliarum, new species Fiaures 150-154 This species is related to a species from Mexico that I believe to be H. paschia Mosely. From this species it differs in the more deeply divided tenth tergum, longer claspers, and smaller membranous process of the aedeagus. Apu.t.—Length of forewing 2 mm. Color brown, mottled with patches of white hair. Male genitalia: ninth segment with anterior margin slightly expanded ventrally, posterior margin developed into a sharp point laterad of clasper; tenth tergum with declivent, sclerot- ized, lateral band, divided dorsally almost to base; clasper evenly 58 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 expanded toward apex in lateral view, with two dark lateral points, in ventral view almost parallel sided; aedeagus with a flattened, heavily sclerotized, apical portion, giving rise subbasally to a mem- branous process, twisted process wrapped around apical section 14% times, shorter than apical section. Female genitalia: eighth sternum elongate apically, dorsal margin deeply and rectangularly emarginate, midventral mark goblet shaped, with a long stem; bursa copulatrix with lyre-shaped apicolateral arms, a starlike central complex and a short basal stem. LARVA, PUPA, CASE.— Unknown. MarteriaL.—Holotype, male: Dominica, Pont Casse, 1.6 miles west, 27 June 1964, O. S. Flint, Jr.,. USNM type 69901. Allotype, female: Clarke Hall, 11-20 Feb. 1965, W. W. Wirth. Paratypes: same data as allotype, 19 ; same, but 21-31 March 1965, 1? ; same, but 11-20 May 1964, O. S. Flint, Jr., 19; same, but 15-30 June 1964, 19; Fond Figues, 7 May 1964, O. S. Flint, Jr., 19. Other: St. Lucia, Vergallier River, near Marquis, 31 July 1963, Flint and Cadet, 19. Brotogy.—The adults have been taken at light primarily in the lowlands, but the holotype was taken near a slowly flowing, small highland stream. Hydroptila grenadensis, new species Figures 155-158 This species is quite closely related to the preceding although there is considerable difference in the shape of the clasper, which is very narrow with an apicodorsal point. Aputt.—Length of forewing 2 mm. Color pale brown, forewing mottled with brown and white. Male genitalia: ninth segment rounded anteriorly, with a sharp spine from posterior margin laterad of clasper; tenth tergum sclerotized laterally, with dorsum divided mesally about a third of its length; clasper narrow, apex developed into a distinct dorsolateral point bearing a black spot; aedeagus flattened apically with a membranous process exiting from near its base, twisted process wrapped around stem 1% times, shorter than apical portion. Female genitalia: eighth segment with posteroventral margin evenly rounded, dorsal margin shallowly emarginate, trilobate, ventral surface with a goblet-shaped mark; bursa copulatrix identical to that of preceding. LARVA, PUPA, CASE.— Unknown. Marterrau.—Holotype, male: Grenada, 2 miles west of Grand Etang, 4-8 Aug. 1963, O. S. Flint, Jr., USNM Type 69902. Allotype, female: same data. Para- types: same data, 3067 4? ; Bathazar, 7 Aug. 1964, O. S. Flint, Jr., 159. Brotocy.—The adults have been taken near both a large, rapid lowland river and a small, tumbling mountain brook. NO. 3665 CADDISFLIES——FLINT 59 Genus Ochrotrichia Moesly Ochrotrichia Mosely, 1934, p. 162. [Type-species: Ochrotrichia insularis Mosely, 1934, by original designation.] The genus, which is of exclusively New World distribution, is divided into two subgenera, Metrichia and Ochrotrichia. The adults of the former are easily recognized by the presence of an apical spur on the foreleg. The mature larvae inhabit compressed, purse-shaped cases, similar in appearance to those of Hydroptila. To date no characteristics have been found that will serve to separate the larvae of the two subgenera. The larvae of Metrichia were described by Flint (1964b); Ochrotrichia, by Ross (1944). Key to Species ADULTS 1. With claspers and aedeagus: males. Without claspers and aedeagus: females . 2. Foretibia without apical spur . Foretibia with apical spur . a, 3. Clasper short with posterior margin developed ie meas spurs. O. spinosissima aw NT bb Clasper elongate, not developed into spurs. . . . Figg ee 4. Clasper elongate, paralled sided, produced into an pcodersl leben O. brayi Clasper shorter, higher, hind margin bilobate. . . ... . . . O. ponta 5. Abdominal segments 4 and 5 with hair tufts and inter sacs. O. exclamationis These segments unmodified. . ... Se Seer CoO! 6. Apicodorsal lobe of clasper narrowed ara deveined ‘invies a ue . O.campana Apicodorsal lobe pointed, but not spurlike. . ....... . .. O.similis Pemitboutapical spur omforetibia. ps. 2.) ft 0 os cs ee we With apical spur on foretibia. . . . . PRs dae ata ey A) 8. Eighth sternum with a tongue-like process o pagina. . . O. spinosissima Highth sternum with a small mesal lobe. . ....... 2... O.ponta 9. Eighth sternum with a large internal sclerite. . .... . . O. species 2 Highth sternum with a posterior collar, but no internal plate . . O. species 1 Ochrotrichia (O.) spinosissima Flint Fiaures 159-163 Ochrotrichia spinosissima Flint, 1964b, p. 58. The species was described from Puerto Rico. The shape of the clasper and aedeagus is distinctive. The female is ascribed to this species primarily on coloration, which is quite different in O. brayi, the only other species of the subgenus in which the female is unknown. Apu.t.—Length of forewing 2-2.5 mm. Color black, legs annulate, forewing with a narrow transverse white band, and small white spots 60 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 apically. Male genitalia: ninth segment developed into a large rounded flap dorsolaterally; clasper short, rather quadrate in lateral view, with a small basoventral lobe; posterior margin developed into numerous short spurs, ventral one heaviest and curving mesad; tenth tergum a simple, broad flap, apex slightly bilobed and asymmetrical; aedeagus KUN Sd 171 Ficures 159-171.—Ochrotrichia (O.) spinosissima Flint: 159, male genitalia, lateral; 160, male genitalia, dorsal; 161, aedeagus, dorsal; 162, female eighth sternum, ventral; 163, bursa copulatrix, ventral. O. (O0.) ponta, new species: 164, male genitalia, lateral; 165, aedeagus, dorsal; 166, male genitalia, dorsal; 167, female eighth sternum, ventral; 168, bursa copulatrix, ventral. O. (O.) brayi, new species: 169, male genitalia, lateral; 170, aedeagus, dorsal; 171, male genitalia, dorsal. NO. 3665 CADDISFLIES—FLINT 61 long, with 2 apical processes, 1 heavily sclerotized and gradually decurved, the other membranous and tubular. Female genitalia: eighth sternum with an elongate tongue-like apicomesal process; rods attached apicolaterally; bursa copulatrix expanded apically with a large central opening and a pair of apicolateral earlike lobes. LARVA, PUPA, CASE.— Unknown. Marertau.—Dominica, Carholm Estate, 7 Feb. 1965, W. W. Wirth, 3c’. Clarke Hall, 11-20 Jan. 1965, W. W. Wirth, 16. Pont Casse, 2.5 miles east, W. W. Wirth, 16 Jan. 1965, 19. Biotocy.—Nothing is known of the biology of the species. Ochrotrichia (O.) brayi, new species Fiaures 169-171 This new species, like so many others in the genus, is not clearly related to any other described species although the simple aedeagus suggests a relationship with O. marica Flint and 0. lobifera Flint. The rather simple tenth tergum with the heavy apical spine is unique. Apu.t.—Length of forewing 3 mm. Color brown. Male genitalia: ninth segment nearly quadrate, without a dorsolateral lobe; tenth tergum elongate, narrow, right side membranous especially sub- apically, left side heavily sclerotized, apex with a short, stout, black spine; clasper elongate with an apicodorsal lobe; black peglike setae on mesal face of apicodorsal lobe, and a few near ventral margin at midlength; aedeagus a long, slender tube with a small subapical tooth. Larva, PuPA, caseE.—Unknown. MarTERIAL.—Holotype, male: Dominica, Freshwater Lake, 22 Feb. 1964, D. F. Bray, USNM Type 69903. Biotogy.—Nothing is known of the biology of this species. Ochrotrichia (O.) ponta, new species Fiaures 164-168 The species would seem to be related to 0. insularis Mosely, from which it differs in possessing a simple tenth tergum and a divided aedeagus. Avu.t.—Length of forewing 2-3 mm. Color fuscus, antennae cream colored, forewing with a narrow, transverse white band at midlength, apically with several small white spots. Male genitalia: ninth segment with a large dorsolateral lobe; tenth tergum elongate, pointed, with a sclerotized band along the dorsolateral margins; clasper elongate with anterodorsal margin straight, hind margin shallowly bilobed, each lobe bearing a patch of dark peglike setae; aedeagus with apical half divided into 2 elongate tubules. Female 62 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 126 genitalia: eighth sternum with posterior margin bearing small mesal lobe, and a thin, projecting sclerite marked with a pair of crescentic dark marks; bursa copulatrix heavily sclerotized, streamlined, with an elongate central opening. LARVA, PUPA, cASE.— Unknown. MarteriaLt.—Holotype, male: Dominica, Pont Casse, .4 miles east, 27 April 1964, O.S. Flint, Jr.. USNM Type 69904. Allotype, female: same data. Paratypes: same data, 11o’ 19; same, but 6 May 1964, 1c; same, but 28 June 1964, 20’; Pont Casse, 1.3 miles east, 10 May 1964, O. S. Flint, Jr., 1c; same, but 18 May 1964, 1h 19; Pont Casse, 1.7 miles east, 24 Mar. 1965, W. W. Wirth, 10’; Pont Casse, 12-14 Oct. 1964, P. J. Spangler, 1o&; Freshwater Lake, 5-8 Nov. 1966, A. B. Gurney, 19 ; Mannett Gutter, 7 Mar. 1965, W. W. Wirth, 1c. Brotogy.—Adults of this species have been taken primarily near small, clear brooks, in which the larvae will probably be found. Ochrotrichia (Metrichia) campana, new species FieureEs 172-174 This and the following species are very closely related. In this species the dorsal lobe of the clasper is developed into a narrow spurlike process. Apvuut.—Length of forewing 2 mm. Color fuscus, antennae and legs paler. A pair of large pouchlike sacs, each filled with many modified setae, between sixth and seventh terga. Male genitalia: ninth sternum produced and rounded anteroventrally, with a heavily sclerotized dorsal lobe; a lightly sclerotized, elongate lobe dorsally extending posteriad between spinelike process arising from dorso- lateral angles of ninth sternum; clasper broad, with a strong dorso- lateral spur curving mesad, and a broad ventrolateral lobe; aedeagus with 2 subapical spines, a heavily sclerotized central tubule, and a slightly inflated basal half. LARVA, PUPA, CASE.—Unknown. Mareriau.—Holotype, male: Dominica, D’leau Gommier, 16 March 1965, W. W. Wirth, USNM Type 69905. Brotocy.—Nothing is known of the biology of the species. Ochrotrichia (M.) similis, new species Figures 175-177 This species is extremely close to the preceding, differing primarily in the dorsal lobe of the clasper, which is broad in O. similis and narrow in O. campana. The pouches between terga six and seven also seem different in the two species. Apuit.—Length of forewing 2.56 mm. Color in alcohol brown. A pair of small pouches between sixth and seventh terga; seventh NO. 3665 CADDISFLIES—FLINT 63 182 ie Fr i 181 171 Ficures 172-185.—Ochrotrichia (Metrichia) campana, new species, male: 172, genitalia, lateral; 173, genitalia, dorsal; 174, aedeagus, dorsal. O. (M.) similis, new species, male: genitalia, lateral; 176, genitalia, dorsal; 177, aedeagus, dorsal. O. (M.) exclamationis, new species, male: 178, genitalia, lateral; 179, fourth and fifth abdominal terga, dorsal; 180, genitalia, dorsal; 181, aedeagus, dorsal. O. (M.) species 1: 182, female eighth sternum, ventral; 183, bursa copulatrix, ventral. O. (M.) species 2: 184, female eighth sternum, ventral; 185, bursa copulatrix, ventral. 64 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 tergum with a dark, trianguloid mark mesally. Male genitalia: ninth segment produced and rounded anteroventrally, with a dumbell- shaped dorsal lobe; apex of lightly sclerotized dorsal lobe with a V-shaped mesal excision; dorsolateral spine long, and slender; clasper, with broad, dorsal and ventral lobes; aedeagus with 2 subapical spines a central tubule, and enlarged base. LARVA, PUPA, CASE.— Unknown. MateErIAL.—Holotype, male: Dominica, Boiling Lake, 19 Nov. 1964, P. J. Spangler, USNM Type 69906. Paratype: Guadeloupe, altitude 3000 ft., 30 July, A. Busck, 1c. Briotocy.—The immature stages and breeding sites are unknown. Ochrotrichia (M.) exclamationis, new species Figures 178-181 This species is not clearly related to any other in the subgenus. The long subapical spine of the aedeagus, the short clasper, and the pouches and hair tufts of the abdomen are all extremely distinctive. Apuutr.—Length of forewing 2 mm. Color fuscus, forewing irregu- larly mottled with silvery patches of hair. A pair of elongate internal tubes arising ventrally between fourth and fifth segments, extending the length of fifth segment; a pair of black, scale-filled sacs postero- ventrally on fourth segment; a pair of large, dorsal hair tufts basally on fourth segment. Male genitalia: ninth segment with anteroventral margin extended and rounded; dorsal lobe short, rounded, lightly sclerotized; dorsal lobe rounded, short, and indistinct; dorsolateral spine broad, curved ventrad; clasper short and broad, with a small mesal spine along posterior margin; aedeagus with a long ribbon-like subapical spine, a twisted, lightly sclerotized process on right, an internal tubule, and expanded base. LARVA, PUPA, CASE.— Unknown. MarrriAL.—Holotype, male: Dominica, Clarke Hall, cocoa trail, 16 Feb. 1965, W. W. Wirth, USNM Type 69907. BroLtogy.—The biology is totally unknown. Ochrotrichia (M.) species 1 Ficures 182-183 There are two females of the subgenus Metrichia in the collections. On the basis of coloration, they are probably not the female of 0. exclamationis, but there is no way to assign these to either O. campana or O. similis. Aputt.—Length of forewing 2-2.5 mm. Color black, antennae and legs paler. Female genitalia: eighth sternum with a thin apical collar NO. 3665 CADDISFLIES—FLINT 65 beyond which extends a thinner, more membranous collar that bears mesally a dark point to which are attached long, internal rods; bursa copulatrix with a short basal stem, a broad rounded apical lobe, and a complex of central structures; ninth and tenth segments extremely long and extensile. MatTERIAL.—Dominica, Freshwater Lake, 14 April 1964, O. S. Flint, Jr., 19 ; Pont Casse, .4 miles east, 16 May 1964, O. 8S. Flint, Jr., 19. Ochrotrichia (M.) species 2 Figures 184-185 On the basis of size and coloration, these are possibly the females of O. exclamationis. The structure of the eighth sternum is extremely distinctive. Aputt.—Length of forewing 1.5-2 mm. Color silver gray, forewing with several transverse bands of white hair, antenna pale basally, dark apically. Female genitalia: eighth sternum with posterior margin slightly lobate mesally; with an internal posteromesal sclerite giving rise to internal rods from its lateral angles and with its posterior mar- gin developed into a projecting mesal lobe; eighth segment dorsally with a pair of knobbed submesal processes; bursa copulatrix ovoid, with lateral margin bandlike and a central complex. MarteErtaL.—Dominica, Pont Casse, 2.5 miles east, 16 Jan. 1965, W. W. Wirth, 39. Family LEPTOCERIDAE The leptocerids are worldwide in distribution but perhaps a little more abundant and diverse in the warmer regions than in the colder. This is probably because they are one of the few families that have successfully invaded the larger, warmer lakes and slowly flowing rivers. The larvae construct cases that are generally tubular and made of sand or organic matter although a few genera hollow-out twigs or utilize discarded cases of other species. Pupation takes place within the larval case, which has been firmly attached to some object in the substrate. Key to Genera LARVAE 1. Metanotum with large sclerotized plates ......4.4... Atanatolica MGtANOLUM. WHOMY MGMpPranOUs, 2) ccc. cs tee olpice sh os) fe. ous wesipy woh es 2 2. Anal proleg ventrally with linear rows of hairs and spines . Brachysetodes Anal prolegs without hairs and spines ventrally ......... Oecetis ADULTS 1. Forewing with M unbranched from base to apex ........ Oecetis Forewing with M branched near wing margin. ........4-.2... 2 313-152—68——_5 66 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 125 Ficures 186-195.—Oecetis pratti Denning: 186, male genitalia, lateral; 187, female genitalia, ventral; 188, internal structure of female seventh segment. Brachysetodes insularis, new species: 189, male genitalia, lateral; 190, clasper, posterior; 191, male genitalia, dorsal; 192, female genitalia, ventral. Atanatolica dominicana, new species: 193, female genitalia, ventral; 194, male genitalia, lateral; 195, male genitalia, ventral. NO. 8665 CADDISFLIES—FLINT 67 2. Hindwing with anal area expanded, wing broader than forewing. Atanatolica Hindwing with anal area compressed, wing narrower than forewing. Brachysetodes Genus Oecetis McLachlan Oecetis McLachlan, 1877, p. 329. [Type-species: Leptocerus ochraceus Curtis, 1825, by subsequent designation of Ross, 1944.] A single species of this worldwide genus has been found on Dominica. The larvae of this genus generally make rather bulky cases of plant material although some make more trim cases of sand grains. The long mandibles and palpi of the larvae are apparently correlated with a trend toward predation. Oecetis pratti Denning Figures 186-188, 196-201 Oecetis pratti Denning, 1947, p. 656.— Wolcott, 1950, p. 93.—Flint, 1964b, p. 62.— Fischer, 1966, p. 140. The species has heretofore been known only from Puerto Rico although there are closely related, if not identical, species in Florida and Peru. The Dominican specimens differ slightly from the Puerto Rican in possessing a more elongate tenth tergum. Apuit.—Length of forewing 7-8 mm. Color brown, forewing with small tufts of black hair where Rs branches and just ventrad on M, where Re+3 and Cu, branch, and where the branches of Cu meet the wing margin. Male genitalia: ninth segment narrow, slightly widened laterally; cercus ovate about twice as long as broad; tenth tergum rodlike, longer than cerci, slightly expanded apically; a conical pro- cess slightly ventrad of tenth tergum; clasper broad basally, tapering to a blunt apex, inner margin near apex bearing short, stout, recurved setae, basomesal lobe with short setae; aedeagus tubular, slightly enlarged apically, internally with a pair of angular sclerites. Female genitalia: eighth sternum without strongly sclerotized marks; bursa copulatrix elongate; seventh segment internally with a hollow, more- or-less spherical structure. Larva.—Length to 5 mm. Head mottled yellow and brown; thoracic notae more uniformly brown. Foreleg short and broad, mid- and hind-legs slender, hindleg especially long, with tibia and tarsus divided at midlengths. Spacing humps well developed; gills single; lateral line lacking. Ninth tergum with 3 pairs of long, black setae. Anal claw with 3 small accessory teeth in a transverse row. Pupa.—Unknown. 68 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 125 CasE.—Length to 9 mm, width 2-3 mm. Constructed of small bits of flat organic matter; occasionally placed to form a case square in cross-section, or more frequently polygonal to round. Ficures 196-201.—Oecetis pratti Denning, larva: 196, case, lateral; 197, head and thorax, dorsal; 198, left mandible, dorsal; 199, right mandible, ventral; 200, foreleg, posterior; 201, hind leg, posterior. MaTERIAL.—Dominica, Pont Casse, 1.6 miles west, 24 April 1964, O. S. Flint, Jr., 1c, 30 larvae; same, but 28 April 1964, 1? ; same, but 29 April 1964, 6 larvae; same, but 18 May 1964, 19 ; same, but 16 June 1964, 29; Pont Casse, .4 mile east, 8 May 1964, O. S. Flint, Jr., 1 larva; same, but 15 June 1964, 29; Pont Casse, .5 mile east, 27 Jan. 1965, J. F. G. and T.M. Clarke, 1c 19 ; Pont Casse, 1.3 miles east, 18 May 1964, O.S. Flint, Jr., 19 ; Pont Casse, 12-14 Oct. 1964, P. J. Spangler, 5 9 ; same, but 23 Nov. 1964, 1? ; same, but 8-13 Oct. 1966, A. B. Gurney, 19; Freshwater Lake, 22 Feb. 1964, D. F. Bray, 1; Clarke Hall, 14 Oct. 1966, E. L. Todd, 19. NO. 3665 CADDISFLIES—FLINT 69 Biotocy.—The larvae were found in pools and other slowly flowing sections of two small mountain streams. On occasion they were found to be abundant, crawling over rocks and leaves in a favored site. Genus Brachysetodes Schmid Brachysetodes Schmid, 1955, p. 134. [Type-species: Brachysetodes trifida Schmid, 1955, by original designation. | I am placing with some hesitation the following species in the Chilean genus Brachysetodes. Although venation is the same in the Dominican species and the type-species, the plan of the male genitalia seems quite different. Perhaps the immature stages of the Chilean species when they are discovered will show that the Dominican species is not congeneric. The larvae are quite similar to those of Leptocella in many charac- teristics; however, the brushes of hair and spines on the ventral portion of the anal prolegs are unique. Brachysetodes insularis, new species Figures 189-192, 202-208 This species seems to be closest to B. bifida Schmid, from which it differs in the round cerci and in the shorter clasper with a differ- ently shaped appendage. Aputt.—Lengths of forewing 4-5 mm. Color brown, forewing golden, with interspersed brown spots roughly arranged in transverse rows. Male with 2 branches of M in forewing, female with 3 branches. Male genitalia: cercus almost spherical, with a small apicoventral point, cerci broadly united dorsomesally; tenth tergum elongate, with thin lateral expansion; clasper terete, nearly upright, apex with a cluster of setae, posterior face with a hooked process; aedeagus tubular, with an apicoventral liplike expansion. Female genitalia: cerci well developed, united ventromesally by a semicircular plate; bursa copulatrix with apical supports, and central keyhole-like opening. Larva.—Length to 6 mm. Sclerites pale yellow brown. Mandibles with 3 apical teeth; maxillary palpi short. Pronotum with many setae on anterior half, anterolateral angles set off by sutures. Metano- tum with a group of setae anterolaterally; sternum with a transverse row of about 20 setae. Spacing humps present, lateral ones poorly developed. A few single gill filaments on basal abdominal segments. Ninth tergum with 3 pairs of long setae. Mesal portion of anal prolegs with a linear brush of hairs margined laterally by spinelike setae, another row of spines mesad of claw. 70 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 125 Pupa.— Unknown. Casr.—Length to 6 mm. Evenly tapered and curved, made of small sand grains. MartTErRIAL.—Holotype, male: Dominica, Pont Casse, 1.6 miles west, 24 April 1964, O. 8. Flint, Jr., USNM Type 69908. Allotype, female: same data. Paratypes: same data, 22o' 39; same, but 27 April 1964, 1°; same, but 28 April 1964, 8¢ 59; same, but 2 May 1964, 19; same, but 9 May 1964, 15c° 59; same, but 16 June 1964, 20’; same, but 25 June 1964, 2c’; same, but 27 June 1964, 1d 19; Ponte Casse, .4 mile east, 7 May 1964, O. S. Flint, Jr., 1c’; Pont Casse, Figures 202-208.—Brachysetodes insularis, new species, larva: 202, case, lateral; 203, head and thorax, dorsal; 204, foreleg, posterior; 205, midleg, posterior; 206, hind leg, posterior; 207, foretrochantin, lateral; 208, anal prolegs, ventral. NO. 3665 CADDISFLIES—FLINT a” 1.3 miles east, 10 May 1964, O. S. Flint, Jr., 2o°; Pont Casse, 1.7 miles east, 12 March 1965, W. W. Wirth, 1c; Pont Casse, 2.2 miles east, 2 May 1964, O. S. Flint, Jr., 16%; Pont Casse, .5 mile south, 22-24 July 1963, O. S. Flint, Jr., 19 ; Pont Casse, 1.5 miles north, 12 Feb. 1965, W. W. Wirth, 30° 3 9 ; Brantridge, 30 April 1964, O. 8. Flint, Jr., 1@; D’leau Gommier, 15 March 1965, W. W. Wirth, 1o; same, but 16 March 1965, 2°; Freshwater Lake, 2 Oct. 1964, P. J. Spangler, 19; Guadeloupe, 4000 feet altitude, 30 July, August Busck, 19. Other: Pont Casse, 1.6 miles west, 29 April 1964, O. S. Flint, Jr., 4 larvae, Brotocy.—The larvae were found sparsely in a slowly flowing pool of a small mountain stream. The adults were taken mostly in the daytime when they were easily beaten from the vegetation surrounding the stream. Genus Atanatolica Mosely Atanatolica, Mosely, 1936, p. 85. [Type-species: Mystacides brasilianus Brauer, 1865, by original designation.] I place the Dominican species in this genus primarily because of the close similarity of the male genitalia to the type-species. There are a few differences in the venation in the two species: in A. domini- cana, fork 1 in the forewing is petiolate, and there is no crossvein in the hindwing between R»2,3 and R4,;. The larvae and pupae of the genus are described herein for the first time. They are especially unusual because they spend most of their time out of water crawling and pupating on moist rocks. Atanatolica dominicana, new species Fiavures 193-195, 209-221 On the basis of male genitalia, this species is closely related to A. brasiliana (Brauer), from which it differs in the apicolateral angles of the tenth tergum, which are finger-like rather than truncate; the apex of the aedeagus, which is bilobate; and the apicomesal angle of the basal part of the clasper, which is obliquely truncate. There are also differences in the venation as stated above. Avutt.—Length of forewing: male 7-9 mm, female 5-6 mm. Head, basal antennal segment with a cream-colored middorsal line, remainder of antennae black with white annulae, forewing brown (darker in female) with apex, apical fringe, and posterior margin cream colored. Male genitalia: ninth segment annular; cercus elongate, narrowing apically; tenth tergum with paired apicodorsal finger-like appendages, and apicolaterally with short digitate lobes; clasper with enlarged basal section whose apicomesal angle is obliquely truncate, a small appressed appendage posteriorly, apical portion with many spinelike setae on mesal surface; aedeagus with an elongate, bilobed, apical lip. Female genitalia: cerci rounded; ventrally to cerci a thin, ventrally 72 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 125 219 200 218 Ficures 209-221.— cal Yenc, chad ob pene bo TODA 4. Protosquilla pulchella (Miers, 1880) Fiaure 5 Gonodactylus pulchellus—Kemp, 1913, p. 177, pl. 10 (figs. 117-118).—Chopra, 1934, p. 41.—Holthuis, 1941, p. 288, fig. 9b [older references].—Ingle, 1963, p. 30, figs. 29, 49. Gonodactylus pulchellurs.—Tirmizi, 1967, p. 35, fig. 2 [erroneous spelling]. Protosquilla pulchella——Holthuis, 1967b, p. 42.—Manning, 1968b, p. 54. MATERIAL. 29, 28-35 mm; western end of Astola Island, ca. 177 miles west of Karachi; rocks, sand, scanty, scattered coral; 0-8 ft; L. P. Woods, et al.; Sta. LW-1; IIOE; 27 November 1963; USNM.—1<’, 45 mm; off Karachi; University of Karachi; USNM.—4<¢, 36-49 mm; 6 fragmented specimens; G. M. Hut, about 22 miles west of Karachi; Zoological Survey no. 1898.—1 broken o', CL 8.7 mm; 39 (2 broken), 45 mm; off Karachi; University of Karachi. Description.—Anterior margin of ocular scales sinuous, scales produced laterally into acute lobes; eyes usually extending beyond end of basal segment of antennular peduncle; cornea subglobular; rostral plate sharply trispinous, median spine slenderer and longer than laterals; anterolateral margins of lateral plates of carapace con- cave, anterolateral angles acute but rounded; mandibular palp 2- segmented; dactylus of claw (fig. 5c) lacking basal notch on outer margin; first 4 abdominal somites almost smooth, at most marked by obscure lateral grooves; fifth abdominal somite with pits arranged in 2 rows on either side of ridge separating the smooth median portion from irregular lateral portions of somite; carinae of sixth abdominal somite and telson covered with short hairs; distal margin of telson (fig. 5d) divided into 2 halves by long, narrow, median fissure, distal margin of each half with 4 teeth or lobes, submedians with movable apices; several small submedian denticles present and 1 denticle present between each of remaining teeth; dorsal surface of telson with 3 bosses (fig. 5d) median triangular, external bosses on each side rounded or oval, not extending much past midlength of telson; surface of telson appearing pitted rather than coarsely reticulate; outer spine of basal prolongation of uropod (fig. 5e) much larger than inner. 313-150—68 9 v4 18 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 Cotor.—Faded in most specimens; male has dark rectangular patches on the sixth thoracic and first and fourth abdominal somites, color most prominent on first abdominal somite; other specimens show traces of light banding over the body. Discusston.—The presence of short hairs on the dorsal surface of the last abdominal somite and telson, the smaller lateral dorsal bosses on the telson, and the four pairs of marginal teeth on the telson will immediately distinguish this species from P. lenzt. The hook process of the petasma of P. pulchella is well developed and extends beyond the tube process. The latter is ornamented with a triangular projection, the margins armed with small blunt spines. Ficure 5.—Protosquilla pulchella (Miers), female, TL 45 mm, off Karachi: a, carapace and rostral plate; b, eye; c, raptorial claw; d, last abdominal somite, telson and uropod; e, uropod, ventral view. Female (broken), off Karachi: f, outline of telson. Male, TL 45 mm G. M. Hut: g, petasma. NO. 3666 CRUSTACEA—TIRMIZI AND MANNING 19 One specimen, a female from off Karachi, has an abnormal telson, with five marginal teeth developed on the left side; it is shown in figure 5f. DisrrisuTion.—Indian Ocean, from the Red Sea and the coast of Africa to Australia. It was recorded previously from West Pakistan by Tirmizi (1967). 5. Protosquilla lenai (Holthuis, 1941) FIGURE 6 Gonodactylus glaber.—Kemp, 1913, p. 182, pl. 10 (fig. 121). Gonodactylus lenzi Holthuis, 1941, p. 288 [older references].—Tiwari and Biswas, 1952, p. 362.—Ingle, 1963, p. 31, fig. 31. Protosquilla lenzi.—Tirmizi, 1967, p. 32, fig. 1—Holthuis, 1967b, pp. 36, 42.— Manning, 1968b, p. 54. MATERIAL.—2 9? , 27-42 mm; G. M. Hut, about 22 miles west of Karachi; 13 February 1965; University of Karachi.—3 , 21-32 mm; 49, 24-35 mm; western end of Astola Island, ca. 177 miles west of Karachi; rocks, sand, scanty, scattered coral; 0-8 ft; L. P. Woods, et al.; Sta. LW-1; IIOE; 27 November 1963; USNM. Description.—Anterior margin of ocular scales rounded, scales acute but rounded laterally; eyes extending beyond end of first seg- ment of antennular peduncle; cornea subglobular; rostral plate Ficure 6.—Protosquilla lenzi (Holthuis), female, TL 42 mm, G. M. Hut: a, carapace and rostral plate; b, eyes; c, propodus and dactylus of raptorial claw; d, last abdominal somite, telson, and uropod. 20 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 125 sharply trispinous, median spine slenderer and longer than laterals; anterior margins of lateral plates of carapace concave, anterolateral angles rounded; mandibular palp 2-segmented; dactylus of claw with prominent notch on outer margin; first 4 abdominal somites smooth, with lateral groove flanked dorsally by slightly irregular area; fifth somite smooth dorsally, with 2 broad longitudinal swollen areas above lateral margin; carinae of sixth abdominal somite and telson smooth, not ornamented with setae; distal margin of telson (fig. 6d) divided into 2 halves by long, narrow, median fissure, distal margin of each half with 3 teeth or lobes, submedians with movable apices; several small submedian denticles present, at most 1 very small intermediate and lateral denticle present, either or both occasionally missing; dorsal surface of telson with 3 bosses, median rounded, smaller than more oval submedians, which extend posteriorly beyond midlength of telson (fig. 6d); outer spine of basal prolongation much broader and larger than inner. Cotor.—Body marked with diffuse bands of dark chromatophores; sixth thoracic somite with median and lateral dark patches, seventh thoracic and first abdominal somites with dark median patch; telson with small dark spot at anterior end of each submedian boss. Discussion.—These specimens agree well with accounts of this species in the literature. Protosquilla pulchella (Miers), the only other species of this genus known from West Pakistan, differs in having the dorsal surface of the sixth abdominal somite and telson covered with short hairs, in having smaller submedian bosses on the dorsal surface of the telson, and in having four pairs of marginal teeth on the telson. One of the males from Astola Island differs from the remainder of the specimens in having but two teeth on one side of the telson margin. DistriBpuTIoN.—Indo-West Pacific region, from the western Indian Ocean to the Philippines. It was recorded from West Pakistan by Tirmizi (1967). Gonodactylus Berthold, 1827 Diaanosis.—Cornea subglobular; rostral plate with apical spine, anterolateral angles usually rounded, rarely acute, not spiniform; anterolateral margins of carapace extending beyond base of rostral plate; mandibular palp present; dactylus of claw unarmed, inflated basally; sixth abdominal somite free, not fused with telson; basal segment of uropodal exopod extending beyond articulation of distal segment, marginal spines straight. Typr-species.—Gonodactylus chiragra (Fabricius, 1781). NO. 3666 CRUSTACEA—TIRMIZI AND MANNING 21 Remarks.—This genus includes the species assigned by Kemp (1913) in his monograph of the Indo-West Pacific stomatopods to Gonodactylus Group I. Nine species have been reported from the northwestern Indian Ocean, and representatives of four of these are reported herein. The other five species, not now known to occur off West Pakistan, are: (1) G. platysoma Wood-Mason, 1895, a common species characterized by its broad body and by the presence of only two pairs of marginal teeth on the telson, the laterals being absent; (2) G. falcatus (Forskal, 1775), also a common species, which has five dorsal carinze in the center of the telson; (3) G. choprat Manning, a small, possibly rare species from moderate depths that lacks the fixed distal spine on the ventral surface of the proximal segment of the uropodal exopod; (4) G. spinosus Bigelow, a small species related to G. lanchesterti Manning and G. demanit Henderson (both reported below), which has a narrow telson tapering distally, with the inter- mediate teeth poorly developed, numerous small dorsal spinules on the telson, and a normal complement of setae on the uropod; and (5) G. segregatus Lanchester, a small species inhabiting moderate depths, which resembles G. choprai but has the distal ventral spine on the proximal segment of the uropod exopod. Any of these species could occur off West Pakistan if suitable habitats were available. The four species of Gonodactylus now known from West Pakistan may be distinguished by means of the following key. Key to Gonodactylus from West Pakistan i= Sorsal'surtace of telson lacking spinules’. . . . . 0. 2.0.0. «© 2s os 2 oe Dorsal surface of telson with spinules ... . Ruse 3 2. Anterolateral angles of rostral plate rounded; carinae ‘of telson inflated) Maren ceeth DlLUNb ede se ee elie BS OP Se SS . . G. chiragra Anterolateral angles of rostral plate cones: carinae of telson slender, usually with apical spinule, marginal teeth sharp. . ...... . . G. smithii 3. Uropodal endopod completely fringed with setae. . . . . . G. lanchesteri Most of inner margin of uropodal endopod smooth, nonsetose, 10 or less proxumal setae present, .* 4 cs, 6, 3) s,s ot ai.om ese . . . G. demanii 6. Gonodactylus chiragra (Fabricius, 1781) FIGURE 7 Gonodactylus chiragra.—Kemp, 1913, p. 155, fig. 2 on p. 161, pl. 9 (fig. 107).— Holthuis, 1941, p. 277, fig. 7 [older references].—Barnard, 1950, p. 861.— Baig, 1954, p. 143.—Manning, 1966, p. 113.—Holthuis, 1967b, pp. 25, 41.— Manning, 1968b, p. 43.—Chhapgar and Sane, 1968, p. 45 [key]. MATERIAL.—2 0’, 1 soft, other 68 mm; 49, 43-78 mm; off Karachi; University of Karachi—19?, 74mm; Hyderabad; University of Karachi.—1?, 52 mm; Manora Island, Karachi; 8. M. H. Balgrammi; 10 October 1953; Zoological Sur- vey reg. no. 272.—1o¢’, 71 mm; 19, 69 mm; same; N. Tirmizi, R. B. Manning, Pipe PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 et al.; 9 March 1967; University of Karachi—1o, 77 mm; off Karachi; Mo- hammed Abdullah el Husseini; USNM.—10, 53 mm; 19, 52 mm; off Karachi; USN M.—1 9, 73 mm; Pasni, Makran coast; F. Townsend; BMNH reg. no. 1898.5. 23.2.6 7, 24-66 mm; 79, 38-53 mm; western end of Astola Island, ca. 177 miles west of Karachi; rocks, scanty, scattered coral; 0-8 ft; L. P. Woods, et al.; Sta. LW-1; IIOE; 27 November 1963; USNM.—2?, 50-56 mm; same data; Sta. RF-2; IOE; USNM. Ficure 7.—Gonodactylus chiragra (Fabricius), male, TL 68 mm, off Karachi: a, anterior portion of carapace and rostral plate; b, carpus, propodus, and dactylus of raptorial claw; d, last abdominal somite, telson, and uropod (setae omitted). Male, TL 71 mm, Manora Island: c, petasma. Diaenosis.—Anterolateral angles of rostral plate rounded (fig. 7a), anterior margins straight or with slight anterior slope; ocular scales large, truncate; telson broader than long, dorsal surface unarmed, dorsal carinae each with at most an apical tubercle; accessory median carinae short, forming anchor; 3 pairs of marginal teeth present, submedians with movable apices, intermediates broad, blunt, laterals poorly formed but distinct; carinae of marginal teeth broad, inflated; numerous small submedian denticles and 2 intermediate denticles NO. 3666 CRUSTACEA—TIRMIZI AND MANNING 23 present, intermediates recessed anteriorly; uropod with full comple- ment of setae. Cotor.—Faded in most specimens; in two fresh specimens from Manora Island the male was dark brownish green, the female a lighter ereen; on both specimens the display patches on the dorsal surface of the merus of the chelae were whitish, lined with light green, with 2 distal spots. Discussion.—Gonodactylus chiragra is the most common species of the genus in the Indo-West Pacific region, and it can be recognized without difficulty through the characters outlined above. Gonodactylus smithii Pocock, discussed below, differs in having acute anterolateral angles on the rostral plate and in other features as well. The display patch on the merus of the claw in G. smithii is blue or crimson in life, not whitish as in G. chiragra. On the male petasma, the hook process is swollen in the middle and extends as far as or a little beyond the tube process; the latter has a truncated distal margin, armed with small distolateral spines (fig. 7c). In the largest male examined (CL 18.1 mm) the median carina is so swollen that it completely obliterates the accessory medians with the result that no anchor is visible (fig. 7d). In general, the carinae of the telson in males are more swollen than in females. The rostral plate figured by Ingle (1963) for a specimen of G. chiragra is the typical shape of the rostral plate of G. smithii. The two specimens collected by us at Manora Island were found burrowing in a rocky flat exposed at low tide. DistrisutTion.—Throughout the Indo-West Pacific region, from the Red Sea and East Africa to Japan, with the exception of Hawaii. 7. Gonodactylus smithii Pocock, 1893 FIGURE 8 Gonodactylus smithii Pocock, 1893, p. 475, pl. 20B (fig. 1)—Manning, 1966, p. 112.—Holthuis, 1967b, pp. 28, 41.—Manning, 1968b, p. 44 [references]. Gonodactylus chiragra chiragra.—Ingle, 1963, p. 27, figs. 27, 47, 63. MarTErRIAL.—19, 74 mm; off Karachi; University of Karachi —1¢, 29 mm; western end of Astola Island, ca. 177 miles west of Karachi; rocks, sand, scanty, scattered coral; 0-8 ft; L. P. Woods, et al.; Sta. LW-1; IIOEH; 27 November 1963; USNM. Diaenosis.—Anterolateral angles of rostral plate acute, sharp, anterior margins sloping anteriorly; ocular scales large, truncate; sixth abdominal somite with 6 sharp carinae, most ending in spines; telson broader than long, dorsal surface unarmed, dorsal carinae each usually with an apical tubercle; dorsal carinae slender, accessory medians present, fusing with median to form anchor; 3 pairs of marginal teeth present, submedians with movable apices, interme- 24 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 diates sharp, laterals poorly formed but distinct; carinae of marginal teeth sharp; numerous small submedian and 2 sharp intermediate denticles present, intermediates recessed anteriorly. Cotor.—Fresh specimens and most specimens in preservative show traces of a crimson or blue display patch on the dorsal surface of the merus of the claw; the dactylus of the claw is pink. Discussion.—The sharp anterolateral angles of the rostral plate will immediately distinguish this species from G. chiragra as well as most other Indo-West Pacific species of the genus. The only other FicurE 8.—Gonodactylus smithii Pocock, female, TL 74 mm, off Karachi: a, carapace and rostral plate; b, last abdominal somite, telson, and uropod; ¢, uropod, ventral view. (Setae omitted.) species from the Indo-West Pacific region with a similar rostral plate is G. hendersoni Manning (see Manning, 1967b), which differs in numerous features, including the presence of dorsal spinules on the telson. In his account of this species from Madagascar, Manning (1968b) noted that G. smithit and G. acutirostris de Man were probably con- specific. Manning also placed Ingle’s (1963) record of G. chiragra chiragra from the Red Sea in the synonymy of G. smithii; the rostral plate of G. chiragra illustrated by Ingle is almost certainly that of G. smithir. NO, 3666 CRUSTACEA—TIRMIZI AND MANNING 25 The small specimen of this species from Astola Island has a faintly sinuate inner margin on the uropodal endopod, as in the specimens from Madagascar reported by Manning (1968b). DistriBuTion.—Indo-West Pacific, from Vietnam, Australia, and the western Indian Ocean. It has not been recorded previously from West Pakistan. 8. Gonodactylus lanchesteri Manning, 1967 FIGuRE 9 Gonodactylus spinosus.—Holthuis, 1967b, pp. 34, 42. Gonodactylus lanchestert Manning, 1967b, p. 11, fig. 4 [other references]; 1968b, job ile MatTEriAt.—1<’, 30 mm; 19, 24 mm (in 2 lots); off Karachi; University of Karachi. Draenosts.—Anterolateral angles of rostral plate rounded or sub- acute, anterior margins straight, ocular scales small, erect; carinae of sixth abdominal somite swollen, each usually with apical tubercle; telson broader than long, dorsal surface ornamented with numerous small spinules and tubercles; all carinae of dorsal surface inflated; anterior dorsal carinae smooth dorsally, spinulose laterally; carinae of marginal teeth very spinulose dorsally; 3 pairs of marginal teeth present, submedians with movable apices; submedian and interme- diate teeth blunt, broad, lateral teeth sharper; numerous small sub- median and 2 sharp intermediate denticles present, inner larger and set at level of apex of intermediate tooth, outer more recessed an- teriorly ; submedian teeth lacking well-marked ventral carinae; uropod with full complement of setae; outer spine of basal prolongation of uropod broader and slightly longer than inner. Cotor.—Almost completely faded; there are traces of black spots in no particular pattern on the dorsum of the male. Discussion.—Manning (1967b) noted that specimens of this species showed two different patterns of dorsal spinulation on the telson. The specimens reported herein are of the form with numer- ous small spinules approaching the condition found in G. spinosus Bigelow. The well-developed intermediate teeth of the telson will immediately distinguish this species from G. spinosus, to which it is closely related. The full complement of setae on the uropod will distinguish this species from G. demaniz (discussed below), in which most of the inner margin of the uropodal endopod and exopod is smooth and devoid of setae. The larger male specimen differs from the female in having all of the carinae of the telson more inflated and in having fewer, blunter 26 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 tubercles on the dorsal surface of the telson. In the male the median — carina is almost subglobular. It also differs from the female in having | the outer intermediate denticle set slightly posterior to the apex of © the intermediate tooth. | The petasma in the male is similar to that of G@. chiragra. DistriputTion.—Western Indian Ocean, where it has been recorded from numerous localities. It has not been recorded previously from West Pakistan. Ficure 9.—Gonodactylus lanchesteri Manning, female, TL 24 mm, off Karachi: a, anterior portion of carapace, rostral plate, and eyes; b, propodus of fourth maxilliped; ¢, last abdominal somite, telson, and uropod. 9. Gonodactylus demanii Henderson, 1893 Figure 10 Gonodactylus demanii.—Manning, 1967b, p. 8, fig. 3 [older references].—Holthuis, 1967b, pp. 32, 41.—Manning, 1968b, p. 50. Gonodactylus demani demani.—Chhapgar and Sane, 1968, p. 45 [key]. NO. 8666 CRUSTACEA—TIRMIZI AND MANNING 2G MATERIAL.—1 9, 22 mm; western end of Astola Island, ca. 177 miles west of Karachi; rocks, sand, scanty scattered coral; 0-8 ft; L. P. Woods, et al.; Sta. LW-1; IIOE; 27 November 1963; USNM 120473. Driaenosis.—Anterolateral angles of rostral plate subacute, usually rounded; ocular scales small, erect; carinae of sixth abdominal somite swollen, each usually with apical tubercle; telson broader than long, dorsal surface ornamented with numberous spinules and small tuber- cles; anterior dorsal carinae smooth dorsally, spinulose laterally; Ficure 10.—Gonodactylus demanii Henderson, female, TL 19 mm, Dahlak, Red Sea: a, last abdominal somite, telson, and uropod; b, submedian teeth of telson, ventral view; c, uropod, ventral view; d, rostral plate. (From Manning, 1967b.) carinae of marginal teeth spinulose dorsally; 3 pairs of marginal teeth present, submedians with movable apices; submedian and intermediate teeth sharper than laterals; numerous small submedian and 2 sharp intermediate denticles present, intermediates both recessed anteriorly; submedian teeth each with low carina on inner ventral surface; endopod slender; most of inner margin of both endopod and exopod of uropod smooth, devoid of setae, endopod with 9-10 small, prox- imal setae; outer spine of basal prolongation of uropod broader and longer than inner. 28 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 Cotor.—Largely faded in the present specimen; there are scattered dark spots on the sixth thoracic and first, third, and fourth abdominal somites, and an anterior pair of dark spots on the telson. Discusston.—The smooth inner margin on the uropodal endopod and exopod will serve to distinguish this species from the other species of the genus occurring off West Pakistan. The illustrations that we give herein have been taken from Manning (1967b) because the condition of our single specimen from Astola Island is not good enough to be illustrated. Distripution.—Western Indian Ocean. It has been recorded from Karachi (Kemp, 1913) and from Astola Island (Manning, 1967b). Family SQuitLipaE Latreille, 1803 DriaGnosis.—Propodi of third, fourth, and fifth thoracic appendages longer than broad, not beaded or ribbed ventrally (fig. 15c) ; telson with sharp median dorsal carina and 4 or more intermediate denticles on margin. Discussion.—Representatives of four of the genera assigned to this family by Manning (1968a) occur off West Pakistan. These genera can be distinguished by means of the key given below. Representatives of five other genera may also occur off West Pakistan; these include: Carinosquilla carinata (Seréne), known from the Red Sea (Ingle, 1963) and Madagascar (Manning, 1968b); Squilloides gilesi (Kemp), which has been reported from the Persian Gulf by Kemp (1913), the Gulf of Oman by Chopra (1939), and the Red Sea by Holthuis (1967b); Alima suppler (Wood-Mason), re- ported from Bombay by Kemp (1913) and Chhapgar and Sane (1968) ; and both Anchisquilla fasciata (de Haan) and Leptosquilla schmeltzii (A. Milne-Edwards), recorded from the Red Sea by Holthuis (1967b). Key to Genera of SQuILLIDAE from Pakistan 1. Cornea small, usually not as broad as stalk; ocular scales fused; submedian teeth of telson with movable’apices .°. .. =... 3). see Clerida Cornea small or large, always broader than stalk; ocular scales separate; sub- median teeth of telson with fixed apices |... '. .. . #7!) FSSNE 2 2. Carapace with deep posterolateral excavation; propodus of raptorial claw withsrow Of erectispinesi-150 wells) a) eee ce Dee eee ee Harpiosquilla Carapace rounded posterolaterally; propodus of raptorial claw with pectina- tions. but not erect;spines|. <.. % «2 3 5!_« «je. 4 <2 se 3 3. Lateral processes of fifth, sixth, and seventh thoracic somite single, not bi- lobed; less than 4 epipods present. . ......4+: +. -:+. se Cloridopsis Lateral processes of fifth, sixth, and seventh thoracic somites bilobed; 4 or more epipods present. 2 6.6344. 2 Sale ESE Shee Oratosquilla | NO. 8008 CRUSTACHA—TIRMIZI AND MANNING 29 Clorida Eydoux and Souleyet, 1842 DiaGnosis.—Hye small, cornea bilobed, rarely broader than dilated stalk; ocular scales fused along midline; carapace rounded postero- laterally; mandibular palp usually present; 2-4 epipods present; dactylus of raptorial claw with 4-5 teeth, upper margin of propodus pectinate; lateral processes of fifth, sixth, and seventh thoracic somites not bilobed, process of fifth somite usually a slender spine; abdomen broad, depressed, carinae usually reduced in number; tel- son with movable apices on submedian teeth; basal prolongation of uropod with spines on inner margin. Typx-species.—Clorida latreillei Eydoux and Souleyet, 1842. Discussion.—Five species of Clorida are known to occur in the northwestern Indian Ocean and one of these is reported herein from West Pakistan. Clorida fallax (Bouvier) has been reported from the Red Sea by Holthuis (1967a) and C. latreillet has been recorded from the Persian Gulf by Kemp (1913). More recently, Chhapgar and Sane (1967) described two new species from Bombay, C. denticauda and C. bombayensis. Any of these species could occur off West Pakistan. Manning (1968b) gave a key to all the species of Clorida. 10. Clorida microphthalma (H. Milne-Edwards, 1837) FIGuRE 11 Squilla microphihalma—Kemp, 1913, p. 31, pl. 1 (figs. 17-20.— Holthuis, 1941, p. 242.—Tiwari and Biswas, 1952, p. 350.—Manning, 1968b, p. 5 [key]—Chhapgar and Sane, 1968, p. 44 [key]. Material.—1 ? , 32 mm; off Karachi; University of Karachi. Duscription.—Eye small, cornea bilobed, set obliquely on stalk; stalk expanded proximally, expanded portion broader than cornea (fig. 115); eyes appressed for most of their length, extending about to end of first segment of antennular peduncle; ocular scales sub- truncate; rostral plate triangular, as long as broad, apex rounded; carapace strongly narrowed anteriorly, without carinae except for reflected marginals; anterolateral spines of carapace strong but not extending past base of rostral plate (fig. 11a); dactylus of claw with 5 teeth, outer margin sinuous; dorsal ridge of carpus undivided, terminating in blunt angle (fig. 1le); mandibular palp present; 4 epipods present; thoracic somites lacking submedian carinae; low, unarmed intermediate carinae present on last 3 somites; lateral process of fifth thoracic somite an angular lobe, directed antero- laterally; fifth somite also with 1 pair of ventrolateral tubercles; lateral processes of sixth and seventh thoracic somites rounded anterolaterally and posterolaterally; abdomen smooth, depressed, 30 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 125 lacking submedian carinae on first 5 somites; abdominal carinae spined as follows: submedian 6, intermediate 5-6, lateral 5-6, marginal 4-5; telson broad, inflated, with 3 pairs of marginal teeth, submedians with movable apices, intermediates and laterals sharp; prelateral lobes not developed; dorsal surface of telson with median carina, a submedian row of tubercles coverging under its distal apex, and several curved rows of tubercles or denticles on the lateral surface; marginal denticles spiniform, 1-2, 7-8, 1; postanal keel absent; uropodal exopod with 5-6 movable spines on outer margin of proximal Ficure 11.—Clorida microphthalma (H. Milne-Edwards), female, TL 32 mm, off Karachi: a, outline of carapace and rostral plate; b, eyes; c, raptorial claw; d, telson and uropod; e, uropod, ventral view (setae omitted). segment; basal prolongation of uropod with 5-6 fixed spines on inner margin and broad, rounded lobe on outer margin of inner spine (fig. 11e). Cotor.—hLargely faded in the present specimen, but carapace, last 3 thoracic and first 5 abdominal somites each with posterior black line; anterolateral plates of abdomen darker than body; proximal segment of uropodal exopod dark distally. Discussion.—Among the characteristic features of C. microphthalma are the elongate eyes, with a very small cornea, the small lateral processes on the fifth thoracic somite, the lack of submedian carinae | NO. 3666 CRUSTACEA—TIRMIZI AND MANNING 3l on the last three and first five thoracic somites, and the small number of spines on the abdominal carinae. DistTRIBuTION.—Indo-West Pacific, from East Africa to China and Australia. Kemp (1913) listed two specimens from Karachi. Harpiosquilla Holthuis, 1964 Diacenosis.—Eye large, T-shaped, cornea bilobed, distinctly broader than stalk (fig. 13a); ocular scales separate; carapace with deep posterolateral excavations; mandibular palp 3-segmented; 5 epipods present; dactylus of raptorial claw with teeth, upper margin of propodus with row of large, erect spines and intervening smaller spines or denticles (fig. 12e); lateral processes of next 2 somites not strongly bilobed, sinuous, sharp posterolaterally; abdomen broad, submedian carinae usually present; telson with apices of submedian teeth fixed; basal prolongation of uropod with at most tubercles on inner margin. Typr-speciEs.—Squulla harpax de Haan, 1844. Discussion.—This genus now includes four species, all of which occur in the western Indian Ocean. Two of the species, H. harpax (de Haan) and H. raphidea (Fabricius), occur off West Pakistan. A third species, H. annandalei (Kemp), has been recorded from the Gulf of Oman by Chopra (1939), and the fourth species, H. melanoura Manning, is known only from Madagascar. We have included here an account of a single specimen of H. raphidea from East Pakistan from the collection of the Zoological Survey Department. The two species known from Pakistan may be distinguished by means of the following key. Key to Species of Harpiosquilla from West Pakistan Lateral process of fifth thoracic somite spined; propodus of claw with alternate LOM ERANGRSHOFbYSPINES. fo 1S GR Bhs Se evs 20a ee seh ela es 8 H. raphidea Lateral process of fifth thoracic somite rounded; propodus of claw with more than 1 short spine between longer ones..............46-. H. harpax 11. Harpiosquilla raphidea (Fabricius, 1798) FIGuRE 12 Squilla raphidea.—Kemp, 1913, p. 88, pl. 7 (fig. 77) [part; older references].— Tiwari and Biswas, 1952, p. 356, figs. 3a, ¢, e. Harpiosquilla raphidea.—Manning, 1968b, p. 14 [key].—-Chhapgar and Sane, 1968, p. 45 [key]. MatTERIAL.—19, 261 mm: Cox’s Bazaar, Chittagong, East Pakistan: Zo- ological Survey reg. no. 888.—19 , 189 mm; Fish Harbour, Karachi; 17 April 1967; University of Karachi. 32 PROCEEDINGS OF THE NATIONAL MUSEUM VoL, 125 Draenosis.—Size large to very large, TL 300 mm or more; rostral plate triangular, apex blunt; carinae of carapace well developed; dactylus of raptorial claw with 8 teeth; upper margin of propodus of claw with widely separated alternate long and short spines; lateral process of fifth thoracic somite produced into an acute spine, ventral process on each side acute; last 3 thoracic somite with well-formed eye ee Ficure 12.—Harpiosquilla raphidea (Fabricius), female, (L 261 mm, Cox’s Bazaar: a, rostral plate; b, lateral processes of exposed thoracic somites; ¢c, raptorial claw; d, telson. Female, TL 189 mm, Karachi: ¢, telson. submedian and intermediate carinae, intermediates of sixth and seventh somites armed posteriorly on large specimens; submedian carinae of abdomen well developed; abdominal carinae spined as follows: submedian 6, intermediate 1-6, lateral 1-6, marginal 1-5; telson with broad median carina and 3 pairs of marginal teeth; prelaterals lobes absent; denticles rounded, 5-6, 7, 1; outer margin of basal NO. 3666 CRUSTACEA—TIRMIZI AND MANNING 33 segment of uropodal exopod with 8 spines, last short; lobe on outer margin of inner spine of basal prolongation rounded, margin concave. Cotor.—Faded in the specimen from East Pakistan. The specimen from Karachi was examined while fresh; one of us (N.T.) made the following color notes: ocular peduncles light pink; posterior border of carapace with a black band; antennal scale yellowish, outlined with dark pigment; claw with merus pink, also marked with a greenish- yellow patch; distal end of propodus with bright yellow streak; thorax and abdomen appearing speckled; last 3 thoracic and first 3 abdominal somites pink, last 3 abdominal somites more cream- colored; tips of spines of last 4 abdominal somites yellow; posterior margin of first 4 abdominal somites black; carinae of telson bluish, apices of teeth yellow; telson with pair of submedian yellow-brown spots; uropod lightly marked with yellow and black, inner half of distal segment of exopod grayish, outer half yellow. Discussion.—The large size of this species and the acutely pointed lateral process of the fifth thoracic somite will immediately distinguish it from H. harpax. Other differences have been noted under the account of the latter species. DistTRIBuTION.—Indian Ocean, from East Africa to Australia. Kemp (1918) recorded it from Karachi. 12. Harpiosquilla harpax (de Haan, 1844) FIGURE 13 Squilla raphidea.—Kemp, 1913, p. 88 [part].—Chopra, 1939, p. 158.—Barnard, 1950, p. 851, figs. 1c, g. Squilla harpax.—Tiwari and Biswas, 1952, p. 358, figs. 3b, d, f.—Ingle, 1963, p. 18, figs. 9, 59. Harpiosquilla harpax.—Holthuis, 1967b, pp. 14, 40.—Manning, 1968b, p. 15, fig. 4—-Chhapgar and Sane, 1968, p. 45 [key]. MATERIAL.—1<, 107 mm; Ahsan; University of Karachi—19?, 142 mm; off Karachi; Central Fisheries Department.—1 dry ?, CL 35.6 mm; near Karachi; Mohammed Abdullah el Husseini; USNM. DIaGNosis.—Size moderate to large, TL 200 mm or less; rostral plate acutely pointed, lateral margins sinuous (fig. 13a); dactylus of raptorial claw with 8 teeth; upper margin of propodus of claw with series of long, erect spines, with 1 or more shorter spine or spinule between the long ones (fig. 13c); lateral process of fifth thoracic somite rounded (fig. 136), ventral process on each side acute; last 3 thoracic somites with submedian and unarmed intermediate carinae, submedians low; submedian carinae of abdomen poorly developed but present; abdominal carinae armed as follows: submedian 6, intermediate 1-6, lateral 1-6, marginal 1-5; telson with sharp median carina and 3 pairs of sharp marginal teeth, prelateral lobes 313-150—68——3 34 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 125 absent; denticles spiniform, 5, 13, 1; outer margin of basal segment of uropodal exopod with 9-10 spines, last short; lobe on outer margin of inner spine of basal prolongation rounded, margin concave. Cotor.—Fresh specimens are brightly colored, as follows: eye- stalks bright yellow; antennal peduncle and scale with pink patches; carinae of carapace lined with dark spots; merus and carpus of rap- Ficure 13.—Harpiosquilla harpax (de Haan), male, TL 107 mm, Ahsan: a, anterior portion of carapace, rostral plate, and eyes; b, lateral processes of exposed thoracic somites; c¢, spines on upper margin of propodus of claw; d, telson; ¢, petasma. torial claw with pink patches; posterior margin of each of thoracic and abdominal somites lined with black spots; lateral portions of abdominal somites pink; sixth abdominal somite pink between sub- median carinae; spines on abdomen, telson, and uropod yellow; telson with grey pits and carinae except for red median carina; prox- imal submedian patches of telson maroon (black in preservative) ; base of uropod pink. NO. 3666 CRUSTACEA—TIRMIZI AND MANNING 35 Discussion.—Tiwari and Biswas (1952) were the first to show that H. harpax (de Haan) is distinct from the larger H. raphidea (Fabricius), which is also widely distributed in the Indo-West Pacific region. Harpiosquilla harpax differs from H. raphidea in having a shorter rostral plate, in lacking the lateral spine on the fifth thoracic somite, in having more than one spine or spinule between the major spines on the propodus of the claw, and in having the carinae of the abdomen less well developed. In H. harpaz the submedian carinae of the abdomen are present but they are poorly-defined. The petasma is illustrated in figure 13e. The tube process is well developed; the hook process has an additional hook-like projection on its inner, distal margin. DistRIBUTION.—Indo-West Pacific region, from the Red Sea and South Africa eastward to Japan. Cloridopsis Manning, 1968 Diacnosis.—Eye small, stalk dilated or margins subparallel, cornea broader than stalk; ocular scales separate; carapace rounded posterolaterally; mandibular palp present or absent; 2-3 epipods present; dactylus of raptorial claw with 5-6 teeth, upper margin of propodus pectinate; lateral processes of fifth, sixth, and seventh thoracic somites not bilobed, process of fifth somite a broad, curved spine; abdomen broad, submedian carinae present; telson with apices of submedian teeth fixed; basal prolongation of uropod with at most tubercles on inner margin. Typr-sprEcies.—Squilla scorpio Latreille, 1825. Discussion.—Two species of Cloridopsis occur in the north- western Indian Ocean, and both are reported herein from West Pakistan. They may be differentiated by means of the key given below. Key to Cloridopsis from West Pakistan Lateral process of fifth thoracic somite with a large black spot; apex of rostral PELE MALO We tls EY fos ahd foie dy satel eehomite: serawhseats aa (Mays C. scorpio Lateral process of fifth thoracic somite lacking a large black spot; apex of rostral A PRLERSROR CS ee rotten tg Bees tac GaN eal eens abn Sioa 7 C. immaculata 13. Cloridopsis scorpio (Latreille, 1825) FicuRres 14a-e Squilla scorpio.—Kemp, 1913, p. 42, pl. 2 (fig. 30).—Holthuis, 1941, p. 243 [older references].—Tiwari and Biswas, 1952, p. 353.—Baig, 1954, p. 143.— Chhapgar and Sane, 1968, p. 44 [key]. MareriaL.—2 9 , 72 mm (1 broken); off Karachi; University of Karachi.—1 , 81 mm; off Karachi; Mohammed Abdullah el Husseini; USNM. 36 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 DescripTion.—Eye small, cornea bilobed, broader than stalk and set obliquely on it; ocular scales rounded laterally, separated by median indentation; rostral plate as long as broad, narrowed anteriorly, apex truncate, with median carina on anterior half; carapace narrowed anteriorly, anterior width slightly less than one-half median length; anterolateral spines of carapace strong but not extending to base of rostral plate, each spine with a rounded ventral lobe; median carina of carapace lacking anterior bifurcation, intermediate carinae Figure 14.—Cloridopsis scorpio (Latreille), female, TL 72 mm, off Karachi: a, anterior portion of carapace and rostral plate; b, eye; c, propodus and dactylus of raptorial claw; d, lateral process of fifth thoracic somite; ¢, last abdominal somite, telson, and uropod (setae omitted). Cloridopsis immaculata (Kemp), female, CL 16.5 mm, off Karachi: f, rostral plate; g, lateral process of fifth thoracic somite. present, poorly marked, not extending to anterior margin of lateral plates; dactylus of raptorial claw with 5 teeth, outer margin of dactylus sinuous, with shallow proximal notch; dorsal ridge of carpus undivided; mandibular palp absent; 2 epipods present; lateral process of fifth to seventh thoracic somites each composed of a single lobe, that of fifth somite broad, produced into an anteriorly directed spine; lateral processes of next 2 somites triangular, rounded posterolaterally ; last 3 thoracic somites with submedian and intermediate carinae, NO. 3668 CRUSTACEA—TIRMIZI AND MANNING ou intermediates unarmed; submedian carinae present on abdomen; abdominal carinae spined as follows: submedian 6, intermediate 5-6, lateral 5-6, marginal 2-5; telson broader than long, with 8 pairs of marginal teeth, submedians fixed, prelateral lobes present, denticles large, rounded, 2, 3-4, 1; postanal keel absent; outer margin of uropodal exopod with 7 short, movable spines; lobe on outer margin of inner spine of basal prolongation of uropod large, rounded. Cotor.—Lateral process of fifth thoracic somite with large, well- marked black spot; last 3 thoracic and all abdominal somites with dark posterior line; second abdominal somite with rectangular dorsal patch of dark color; proximal segment of uropodal exopod with dark spot. Discusston.—Living or freshly caught specimens of C. scorpio can immediately be recognized by the presence of the black spot at the base of the lateral process of the fifth thoracic somite; the spots can persist for many years in preservative. Cloridopsis immaculata, which is discussed below, lacks these black spots and exhibits other differ- ences as well. One of the specimens has only three intermediate denticles on one side of the margin of the telson although there is an indication of a fourth denticle appressed to the outer margin of the submedian tooth. Kemp (1913) reported three to six intermediate denticles in his material. Distripution.—Indo-West Pacific region, throughout the Indian Ocean into the western Pacific. Both Kemp (1913) and Baig (1954) have reported the species from Karachi. 14. Cloridopsis immaculata (Kemp, 1913) Figures 14f, g Squilla scorpio var. immaculata Kemp, 1913, p. 45, pl. 2 (fig. 31). Matrriau.—1 fragmented 9, CL 16.5 mm; off Karachi; University of Karachi. Discussion.—This specimen is so fragmented that it is not possible to prepare a description, but from what we can see of the specimen it agrees well with Kemp’s account. Kemp treated this species as a “variety” of Squilla scorpio, but there seems to be no good reason not to recognize it as a distinct species. Both species are distinct throughout their range. Cloridopsis immaculata differs from C. scorpio as follows: (1) the rostral plate is longer and has a broader apex; (2) the median and lateral carinae of the carapace are more distinct; (3) the lateral processes of the sixth and seventh thoracic somites are slightly more upturned; and (4) the lateral process of the fifth thoracic somite is never marked with a conspicuous black spot. 38 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 125 DistTRIBUTION.—Northern Indian Ocean. It was recorded from Karachi by Kemp (1913) but is known from no other localities in the Arabian Sea. Oratosquilla Manning, 19638 Diaenosis.—Eye large, cornea bilobed, noticeably broader than stalk; ocular scales separate; carapace rounded posterolaterally ; mandi- bular palp usually present; 4 epipods present; dactylus of raptorial claw with 5 or more teeth (usually 6), upper margin of propodus pecti- nate; lateral processes of fifth, sixth, and seventh thoracic somites bilobed; abdomen with paired submedian, intermediate, and lateral carinae; telson with fixed apices on submedian teeth; basal prolonga- tion of uropod with at most serrations or tubercles on inner margin. TyprE-spEcies.—Squilla oratoria de Haan, 1844. Discussion.—Nine species of Oratosquilla have been recorded from the northwestern Indian Ocean; three of these are recorded herein from West Pakistan. The other species which are known from the general area are: (1) O. quinquedentata (Brooks, 1886), reported from Bombay by Kemp (1913); (2) O. investigatoris (Lloyd, 1907), reported from the Persian Gulf by Kemp (1913) and the South Arabian coast by Chopra (1939); (8) O. gonypetes (Kemp, 1911), recorded from the Persian Gulf by Kemp (1913) and the Gulf of Oman by Chopra (1939); (4) O. perpensa (Kemp, 1911), reported from the Persian Gulf by Kemp (1913); (5) O. massavensis (Kossmann, 1880) from the Red Sea (records summarized by Holthuis, 1967b); and (6) O. simulans (Holthuis, 1967) from the Red Sea. Kemp (1913) also reported 0. woodmasoni (Kemp) and O. massa- vensts from localities in the western Indian Ocean; those specimens are probably referable to O. hesperia (Manning) (see below). All three of the species discussed below are extremely abundant in the waters off West Pakistan; most of the specimens were obtained at the local fish market in Karachi. Key to Oratosquilla from West Pakistan 1. Median carina of carapace distinct throughout its length, bifurcation open posterior to dorsal pit; anterior lobe of lateral process of sixth thoracic somite with apex truncate 9.0. 2): Si. sh es O. nepa Median carina of carapace either interrupted or indistinct anteriorly; anterior bifurcation, if present, open anterior to dorsal pit; anterior lobe of lateral process of sixth thoracic somite with apex acute .......... 2 2. Anterior width of carapace less than one-half median length; lobe on outer margin of inner spine of basal prolongation of uropod rounded, margin CONVER: ha, Bert MT Ay ae US io Bebe See a on O. interrupta Anterior width of carapace more than half median length; lobe on outer margin of inner spine of basal prolongation of uropod sharp, acute, margin CONCAVE a) ei Fake Be eke fariond tg ole esoaltss eH gots ad Pa Oe O. hesperia NO. 8666 CRUSTACEA—TIRMIZI AND MANNING 39 15. Oratosquilla nepa (Latreille, 1825) FicurE 15 Squilla nepa.—Kemp, 1913, p. 60, pl. 4 (fig. 49).—Chopra, 1934, p. 23.—Holthuis, 1941, p. 245 [older references].—Barnard, 1950, p. 847, figs. lb, 2a.—Kurian, 1954, p. 85.—Holthuis, 1967b, p. 7—Manning, 1968b, p. 31, fig. 10.—Chhap- gar and Sane, 1968, p. 45 [key]. MatTErIAu.—6 6, 72-93 mm; 6 2, 538-83 mm (in 8 lots) ; off Karachi; University of Karachi.—2o, 75-92 mm; 1?, 80 mm; same; USNM.—1<’, 99 mm; off West Pakistan; A. H. Qadri; USNM. DescripTion.—Eye of moderate size, cornea bilobed, set almost transversely on stalk, cornea width less than length of stalk; ocular scales subquadrate or rounded, inclined laterally; rostral plate sub- quadrate, tapering distally, apex truncate or rounded, lateral margins upturned, obscure median tubercle occasionally present; anterior bifurcation of median carcina of carapace uninterrupted, bifurcation opening posterior to dorsal pit, secondarily closing anterior to pit in some specimens; intermediate carinae of carapace not extending to anterior margin, converging anteriorly with laterals; anterolateral spines of carapace strong, extending to or beyond base of rostral plate; dactylus of claw with 6 teeth, outer margin sinuous; dorsal ridge of carpus irregular, with 2-3 large tubercles; inferodistal angle of merus with blunt spine; mandibular palp present; 4 epipods present; last 3 thoracic somites rough, pitted, with submedian and intermediate carinae, neither armed, intermediates pitted; anterior lobe of lateral process of fifth thoracic somite a slender, anteriorly directed spine, posterior lobe shorter, slender, apex acute but rounded; lateral proc- esses of next 2 somites bilobed, anterior lobe on sixth somite large, obliquely truncate, posterior lobe larger, triangular, apex blunt, anterior lobe of lateral process of seventh somite an acute but rounded lobe, posterior lobe larger, triangular, apex rounded; abdomen rough, pitted, surface irregular; abdominal carinae spined as follows: sub- median 4-6, intermediate 3-6, lateral 2-6, marginal 1-5; telson flattened, about as long as broad, with 3 pairs of marginal teeth, sub- medians and intermediates slender, elongate; prelateral lobes present; denticles large, rounded, 2—4, 6-9, 1; dorsal surface of telson, either side of median carcina, with lines of pits, but lacking rows of tubercles or carinae except for carinae of marginal teeth; long postanal keel present; uropod with 8-9 short, movable spines on outer margin of proximal segment; lobe on outer margin of inner spine of basal prolongation of uropod low, rounded, margin concave. Cotor.—Most specimens faded in preservative; some show traces of dark, rectangular patches on the second and fifth abdominal somites; uropod with dark color on exopod at articulation of distal 40 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 126 segment, inner half of distal segment dark, endopod dark distally. Fresh specimens may be brightly colored, with the median carina of the carapace red, remainder of carinae and grooves on the carapace light green or orange; display patch on merus of claw light blue with ereen anterior border; carinae of abdomen light green, some with distal portions orange or red; carinae of telson green, apices of median carina and submedian and intermediate marginal teeth reddish; uropods marked with blue, green, yellow, and black, spines orange. Fielden, lea Ficure 15.—Oratosquilla nepa (Latreille), male, TL 77 mm, off Karachi: a, outline of anterior portion of carapace and rostral plate; b, outline of lateral processes of fifth, sixth, and seventh thoracic somites; ¢, propodus and dactylus of third maxilliped. Male, TL 87 mm: d, eye; ¢, median carina of carapace. Median carina of carapace: f, male TL 72 mn; g, female, TL 79 mm; h, male, TL93 mm. Female, TL 83 mm: 7, abnormally developed posterior portion of carapace. Male, TL 93 mm: j, petasma. NO. 3666 CRUSTACEA—TIRMIZI AND MANNING Al Not all fresh specimens exhibit dark patches on the second and fifth abdominal somites, but these are well marked in a few specimens. Discusston.—The relatively small eyes, long anterior bifurcation on the median carina of the carapace, and large, truncate anterior lobe on the lateral process of the sixth thoracic somite will immediately distinguish O. nepa from the other two species of the genus found in the waters off West Pakistan, 0. interrupta and O. hesperia. Orato- squilla nepa is the least abundant of the three species, but it still occurs in relatively large numbers. Several of the specimens show unusual variation in the configura- tion of the anterior bifurcation of the median carina of the carapace; these variations are shown in figures 15e—-h. The typical shape of the bifurcation is shown in figure 15¢; in some specimens the arms of the bifurcation converge anteriorly, beyond the dorsal pit, but do not meet. In one specimen the bifurcation is closed by a short bar (fig. 15f). Finally, in each of a series of three specimens the bifurcation closes anterior to the dorsal pit, continues anteriorly for a short distance as a single ridge, and reopens again (fig. 154). There does not seem to be any correlation of these variations with sex or size, and the specimens are typical of the species in all other respects. Some specimens exhibit a uniform dusky-gray color pattern, whereas others show distinct rectangular patches on the second and fifth abdominal somites, suggesting a possible dimorphism in color. Kemp (1913) noted similar variation in other collections. In one of the specimens the posterior margin of the carapace is deformed, and the median posterior projection is bifurcate (fig. 157). DistTRIBUTION.—Widely distributed in the Indo-West Pacific re- gion, from West Pakistan and Mogambique to Hong Kong and Aus- tralia. It recorded from Karachi by Kemp (19138). 16. Oratosquilla interrupta (Kemp, 1911) Fiaure 16 Squilla interrupta—Kemp, 1913, p. 72, pl. 5 (figs. 60-62).—Chopra, 1934, p. 25.— Holthuis, 1941, p. 253 [older references].—Baig, 1954, p. 143.—Manning, 1966, p. 97, fig. 4.—Chhapgar and Sane, 1968, p. 45 [key]. MaTEerRIAL.—1@, 71 mm; off Karachi; Central Fisheries Department.—10<, 88-113 mm; 59, 938-123 mm (in 7 lots); off Karachi; University of Karachi.— ldry ?, CL 17.2 mm; near Karachi; Mohammed Abdullah el Husseini; USNM.— 1o¢, 128 mm; West Pakistan; A. H. Qadri; USNM.—1<, 100 mm; off Karachi; University of Karachi; USNM. Description.—Hye large, cornea bilobed, set obliquely on stalk; ocular scales subtruncate, inclined laterally; rostral plate subquadrate, without carinae, lateral margins upturned, apex truncate; anterior bifurcation of median carina of carapace with basal interruption, 42 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 125 carina and arms of bifurcation distinct anterior and posterior to smooth interrupted portion; intermediate carinae of carapace not extending to anterior margin; anterolateral spines of carapace strong but not extending to base of rostral plate; dactylus of raptorial claw with 6 teeth, outer margin sinuous; dorsal ridge of carpus of claw with 2 tubercles; inferodistal angle of merus of claw with broad, blunt spine; Ficure 16.—Oratosquilla interrupta (Kemp), male, TL 95 mm, off Karachi: a, outline of anterior portion of carapace and rostral plate; b, lateral processes of fifth, sixth, and seventh thoracic somites. Male, TL 98 mm, off Karachi: c, outline of basal prolongation of uropod; d, petasma. mandibular palp present; 4 epipods present; last 3 thoracic somites with submedian and intermediate carinae, none armed; anterior lobe of lateral process of fifth thoracic somite an anteriorly directed spine, posterior lobe slender, acute, directed laterally; lateral processes of next 2 somites bilobed, anterior lobe smaller than triangular posterior lobe on both somites, anterior lobe of process of sixth somite slenderer NO. 3666 CRUSTACEA—TIRMIZI AND MANNING 43 and longer than that of seventh somite; abdominal carinae spined as follows: submedian 5-6, intermediate 4-6, lateral 3-6, marginal 1-5; telson broad, with 3 pairs of slender marginal teeth, prelateral lobes present; denticles rounded, 2-3, 7-9, 1; dorsal surface of telson lacking well-marked tubercles or ridges lateral to median carina other than short carinae of marginal teeth; postanal keel present; uropodal exopod with 8-9 movable spines on outer margin of proximal segment; lobe on outer margin of inner spine of basal prolongation of uropod rounded, margin convex. Cotor.—Usually faded in preservative. Fresh specimens have median carina and gastric grooves of carapace red; posterior margin of carapace, last 3 thoracic somites, and first 5 abdominal somites red; median carinule and submedian and intermediate carinae of body red, color most intense on submedians; carinae of marginal teeth of telson green, apices of teeth reddish; telson with a prominent, large red or green spot on the anterior portion of the median carina; spine of uropod pink. There may be considerable variation in overall pattern. Discussion.—The specimens examined by us agree well with ac- counts of the species in the literature. The best feature for recognition of O. interrupta is the convex lobe on the outer margin of the inner spine of the basal prolongation of the uropod; in no other species is this lobe so shaped. This is the second most abundant stomatopod taken by local fisherman off Karachi. DistrisutTion.—Widely distributed in the Indo-West Pacific, from the Persian Gulf to Australia and Formosa. Both Kemp (1913) and Baig (1954) recorded its occurrence off Karachi. 17. Oratosquilla hesperia (Manning, 1968) FiqureE 17 Squilla nepa.—Miers, 1880, p. 25 [part; specimen from Zanzibar]. Squilla woodmasont.—Kemp, 1913, p. 74 [part; specimens from Zanzibar, Muscat, and Aden reidentified]. Squilla massavensis.—? Kemp, 1913, p. 76 [part; specimens from the Gulf of Oman and Persian Gulf].—Ingle, 1963, p. 15 [part; specimens from Zanzibar and the Persian Gulf reidentified]. Squilla hesperia Manning, 1968b, p. 25, fig. 8. MartTERIAL.—1<, 62 mm; fish market, Fish Harbour, Karachi; 24 November 1966; University of Karachi —6<, 52-89 mm; 39, 66-101 mm (in 5 lots); off Karachi; University of Karachi.—1 ? , 89 mm; off Karachi; University of Karachi; USNM.—19, 113 mm; off Karachi; Central Fisheries Department.—1<, 89 mm; 19, 66 mm; off Karachi; Zoological Survey Department reg. no. 1877.—8 0, 60-80 mm; 89, 66-102 mm; 25°04’ N, 65°24-26’E; off West Pakistan; 26 m; hard packed mud; Anton Bruun Sta. 237A; IIOE; 22 November 1963; USNM. 44 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 DerscripTion.—LHye large, cornea bilobed, set obliquely on stalk; ocular scales subtruncate, inclined laterally; rostral plate as long as broad, or broader than long, trapezoidal, upturned lateral margins converging on truncate or rounded apex, median carina absent; median carina of carapace lacking well-marked anterior bifurcation; intermediate carinae of carapace not extending to anterior margin; anterolateral spines of carapace well developed but not extending to base of rostral plate; dactylus of raptorial claw with 6 teeth, outer margin sinuous; dorsal ridge of carpus of claw with large, irregular tubercles; inferodistal angle of merus of claw with broad, obtuse projection; mandibular palp present; 4 epipods present; last 3 thoracic somites with submedian and intermediate carinae, none armed; anterior lobe of lateral process of fifth thoracic somite a slender, anteriorly directed spine, posterior lobe short, slender, apex rounded, directed laterally; anterior lobe of lateral process of sixth thoracic somite slender, posterior lobe much larger, triangular, apex acute but not spiniform; anterior lobe of lateral process of seventh thoracic somite more obtuse than on sixth, posterior lobe similar to that of sixth somite; second to fifth abdominal somites with anterior tubercles between intermediate and lateral carinae; abdominal carinae spined as follows: submedian (8-4) 5-6, intermediate 3-6, lateral (1) 2-6, marginal 1-5; telson broad, with 3 pairs of marginal teeth, submedians and intermediates slender, sharp; prelateral lobes present; denticles rounded, 2-4, 7-9, 1; dorsal surface of telson roughened, with sub- median row of tubercles converging under posterior apex of median carina, lateral surface between lines of pits raised, irregular, almost carinate; margin of telson in males more swollen than in females; postanal keel present; uropod with 8-9 movable spines on outer margin of proximal segment of exopod; basal prolongation of uropod with small rounded lobe, margin concave, on outer margin of inner spine in adults, lobe spiniform in juveniles and subadults. Cotor.—In preservative, carapace with median oval patch and posterior dark line, body segments each with posterior dark line; second and fifth abdominal somites with broad, dark median patch; narrower on fifth somite; uropodal exopod with dark spot at articu- lation of distal segment, inner half of distal segment dark; distal half of endopod dark. Color in living specimens may be bright and variable; there is also evidence of sexual dimorphism in color of the telson. The carinae and gastric grooves of the carapace are orange, and the posterior margin is orange and yellow. The submedian carinae of the abdomen are orange, and each abdominal somite is lined posteriorly with orange; other carinae are blue or green anteriorly, more yellow posteriorly, and the intermediate and lateral carinae of the sixth NO. 3666 CRUSTACEIA—-TIRMIZI AND MANNING 45 somite are green. In females, the telson carinae and margins are primarily orange, whereas in males they are green and blue green. Discusston.—Manning (1968b) described O. hesperia from a single female taken off Madagascar. In his account, he pointed out that O. hesperia was very similar to O. massavensis in general appearance and he suggested that records of O. massavensis from localities south of the Red Sea actually might be referable to 0. hesperia. Examination Ficure 17.—Oratosquilla hesperia (Manning), female, TL 92 mm, off Karachi: a, outline of anterior portion of carapace and rostral plate; b, lateral processes of fifth, sixth, and seventh thoracic somites. Basal prolongation of uropod: c, male, TL 52 mm; d, female. TL 92 mm; e, female TL 101 mm. of specimens in the British Museum from Zanzibar (1c, 104 mm; 19, 75 mm; not registered), Muscat (1c, 34 mm; reg. no. 87-16), and Aden (1c, 42 mm; reg. no. 1894.5.16.4), all of which are O. hesperia rather than O. massavensis, bear out this suggestion. Oratasquilla hesperia differs from O. massavensis in several important features as follows: (1) the rostral plate is shorter and broader, and the lateral margins are rarely markedly concave as in 0. massavensis; (2) the submedian carinae of the abdomen are divergent on each so- 46 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 125 mite rather than subparallel; (3) there is a dark patch of chromato- phores on both the second and fifth abdominal somites; and (4) there are many less tubercles on the telson (in O. massavensis there are two rows of erect tubercles flanking the median carina, one row on each side of the convergent lines of pits; in O. hesperia, only the inner of these two rows is well developed; the tubercles on the anterior half of the lateral surface of the telson, which are present in O. massavensis, are rarely developed in O. hesperia). The two dark dorsal patches on the abdomen are not visible on the type; they are not always visible on the specimens from Karachi reported herein. Both O. hesperia and O. massavensis resemble O. woodmasoni (Kemp) in several features, but both of the former species can be distinguished from O. woodmasoni by the presence of dorsal tubercles on the telson and by the well-developed lobe on the inner spine of the basal pro- longation of the uropod. All three species have a smooth, polished carapace, lacking a well-developed anterior bifurcation on the median carina, and in all three the anterior width of the carapace is greater than half the median length of the carapace. Small specimens of O. hesperia have a sharp spine instead of a rounded lobe on the inner spine of the basal prolongation of the uropod; in adults the lobe is rounded and may be very inconspicu- ous. The lobe is illustrated in figures 17c-e for three specimens, TL 52, 92, and 101 mm. In adult males (TL 75 mm or more) the margin of the telson is noticeably more inflated than in the females. The denticles and carinae of the teeth may all be affected by the enlargement. Oratosquilla hesperia is the most abundant stomatopod in the fishing grounds off Karachi. DIstTRIBUTION.—Western Indian Ocean, from Muscat, Aden, West Pakistan, Zanzibar, and Madagascar. Literature Cited Baie, M. M. 1954. Species of Stomatopoda of Karachi coast. Proc. 6th Pakistan Sci. Conf., vol. 3, pp. 143-144. Batss, H. 1938. Stomatopoda. Jn Bronn, Klassen und Ordnungen des Tierreichs, vol. 5, sec. 1, book 6, pt. 2, pp. 1-178, figs. 1-114. BARNARD, K. H. 1950. Descriptive list of South African stomatopod Crustacea (mantis shrimps). Ann. South African Mus., vol. 38, pp. 838-864, figs. 1-4. 1962. New records of marine Crustacea from the East Africa region. Crus- taceana, vol. 3, no. 3, pp. 239-245, figs. 1-3. NO. 3666 CRUSTACEA—TIRMIZI AND MANNING 47 Cuuapaar, B. F., and Sang, S. R. 1967. Two new species of Squilla (Stomatopoda) from Bombay. Crusta- ceana, vol. 12, pt. 1, pp. 1-8, figs. 1-2. 1968. The Stomatopoda of Bombay. Journ. Biol. Sci., vol. 9, nos. 1-2 (1966), pp. 48-46. Cuopra, B. 1934. On the stomatopod Crustacea collected by the Bengal Pilot Service off the mouth of the River Hughli, together with notes on some other forms. Rec. Indian Mus., vol. 36, pp. 17-43, figs. 1-5. 1939. Stomatopoda. Jn The John Murray Expedition, Sci. Reps., vol. 6, no. 3, pp. 137-181, figs. 1-13. Ho.urauis, L. B. 1941. The Stomatopoda of the ‘‘Snellius’’ Expedition. Jn Biological Results of the “Snellius’”’ Expedition, XII. Temminckia, vol. 6, pp. 241-294, figs. 1-9. 1964. Preliminary note on two new genera of Stomatopoda. Crustaceana, vol. 7, no. 2, pp. 140-141. 1967a. Fam. Lysiosquillidae et Bathysquillidae. Stomatopoda I in Gruner and Holthuis, eds., Crustaceorum catalogus (1), ed. a, pp. 1-28. Den Haag: W. Junk. 1967b. The stomatopod Crustacea collected by the 1962 and 1965 Israel South Red Sea Expeditions. In The Second Israel South Red Sea Expedition, 1965, rep. no. 1. Israel Journ. Zool., vol. 16, pp. 1-45, figs. 1-7. Inez, R. W. 1963. Crustacea Stomatopoda from the Red Sea and Gulf of Aden. In Contributions to the knowledge of the Red Sea, no. 26. Bull. Sea Fish. Res. Sta., Haifa, no. 33, pp. 1-69, figs. 1-73. Kemp, S. 1913. An account of the Crustacea Stomatopoda of the Indo-Pacific region based on the collection in the Indian Museum. Mem. Indian Mus., vol. 4, pp. 1-217, pls. 1-10, 10 text-figs. Kurian, C. V. 1954. Contributions to the study of the crustacean fauna of Travancore. Bull. Centr. Res. Inst., Univ. Travancore, Trivandrum, ser. C, vol. 3, no. 1, pp. 69-91, 3 figs. Mannina, Raymonp B. 1963. Preliminary revision of the genera Pseudosquilla and Lysiosquilla, with descriptious of six new genera (Crustacea: Stomatopoda). Bull. Mar. Sci. Gulf and Caribbean, vol. 13, no. 2, pp. 308-328. 1966. Notes on some Australian and New Zealand stomatopod Crustacea, with an account of the species collected by the Fisheries Investiga- tion Ship Endeavour. Rec. Australian Mus., vol. 27, no. 4, pp. 79-137, figs. 1-10. 1967a. Notes on the genus Manningia with description of a new species. Proc. U.S. Nat. Mus., vol. 122, no. 3589, pp. 1-13, figs. 1-3. 1967b. Notes on the demanii section of genus Gonodactylus Berthold with descriptions of three new species. Proc. U.S. Nat. Mus., vol. 123, no. 3618, pp. 1-27, figs. 1-8. 1968a. A revision of the family Squillidae (Crustacea, Stomatopoda), with the description of eight new genera. Bull. Mar. Sci., vol. 18, no. 1, pp. 105-142, figs. 1-10. 48 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 125 1968b. Stomatopod Crustacea from Madagascar. Proc. U.S. Nat. Mus., vol. 124, no. 3641, pp. 1-61, figs. 1-16. 1969. Notes on some stomatopod Crustacea from South Africa. Smithsonian Contrib. Zool., no. 1, in press. Miers, E. J. 1880. On the Squillidae. Ann. Mag. Nat. Hist., ser. 5, vol. 5, pp. 1-30, 108-127, pls. 1-3. Nositi, G. 1906. Décapodes et stomatopodes. Jn Faune carcinologique de la Mer Rouge. Ann. Sci. Nat. Zool., ser. 9, vol. 4, pp. 1-347, figs. 1-12, pls. 1-11. Pocock, R. I. 1893. Report upon the stomatopod crustaceans obtained by P. W. Bassett- Smith, Esq., surgeon, R.N., during the cruise, in the Australian and China seas, of H.M.S. ‘“‘Penguin,’’? Commander W. U. Moore. Ann. Mag. Nat. Hist., ser. 6, vol. 11, pp. 473-479, pl. 20B. SERENE, R. 1962. Révision du genre Pseudosquilla (Stomatopoda) et définition des genres nouveaux. Bull. Inst. Océanogr. Monaco, no. 1241, pp. 1-27, figs. 1-5. Tirmizi, Nasima M. 1967. Occurrence of Protosqwilla lenzi (Crustacea: Stomatopoda) from Karachi coast with notes on the other species. Univ. Stud., Univ. Karachi, vol. 3, no. 3, pp. 32-39, fig. 1-2. Tiwari, K. K., and Biswas, 8. 1952. On two new species of the genus Squwilla Fabr., with notes on other stomatopods in the collections of the Zoological Survey of India. Rec. Indian Mus., vol. 49, nos. 3-4, pp. 349-363, figs. 1-5. U.S. GOVERNMENT PRINTING OFFICE: 1963 BEQeecdings of the United States National Museum SMITHSONIAN INSTITUTION + WASHINGTON, D.C. Volume 125 1968 Number 3667 Benthic Polychaetes from Puget Sound, Washington, with Remarks on Four Other Species ' By Karl Banse and Katharine D. Hobson? We describe here eight new polychaete species, give 25 new records, and discuss 35 other forms. Forty-one species (including one unnamed species discussed below, and three new species and four new records to be published by K. Banse and F. H. Nichols in other papers) are added to 394 benthic polychaetes previously known from waters of Washington and British Columbia. The new records probably reflect incomplete knowledge of the region rather than recent immigration into the area. There is no trend toward predominantly arctic or warm water additions, as one would expect with new immigration resulting from climatic changes. Incomplete knowledge of the polychaete fauna of this area is suggested further by the fact that the 41 additions mentioned above were among 162 named polychaete species in our collection. For comparison, there is no new record among 49 named bivalve species of the same collection (Mr. D. Kisker, pers. comm.). Polychaetes were identified from 116 samples taken in 1963 at eight subtidal stations in Puget Sound by the Department of Ocean- ography, University of Washington, under the direction of U. Lie 1 Contribution No. 470, Department of Oceanography, University of Wash- ington, Seattle, Wash. 98105. 2 Banse: Department of Oceanography, University of Washington, Seattle, Wash. 98105; Hobson: Systematics-Ecology Program, Marine Biologica Laboratory, Woods Hole, Mass. 02543. 1 2 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 and K. Banse. Samples were collected with a 0.1-m? van Veen grab and washed through a screen with square meshes of 1 mm side length. The formalin-preserved animals were separated from the residue in the laboratory under slight magnification. The nomenclature follows Hartman (1959, 1965b) except as noted otherwise. A reference to other descriptions is omitted if the species is included in Berkeley and Berkeley (1948, 1952). All colors refer to formalin-preserved material. Drawings illustrating new species are made from holotypes if not stated otherwise. Station numbers are given only for new species and new records. A list of all species in our collection with specimen numbers per station and dates of sampling is contained in Lie (in press). A set of identified species is deposited in the U.S. National Museum. The station characteristics are given below (from Lie, in press). The mean depth is calculated from seven quarterly visits (this is given on the labels of the material in the U.S. National Museum). The depth range, however, refers to the recordings of those visits when polychaetes were identified. Quarterly samples were studied for stations 2 and 7, whereas for the other stations polychaetes were identified only from winter and spring collections. Sediment was subsampled for particle size analysis from about one-third of the grab samples from the winter and spring collections. The average results are presented below (‘‘clay” refers to particles smaller than 0.0031 mm, “silt”? to particles between 0.0031 and 0.0625 mm in diameter, and “‘eravel” to particles larger than 2.0 mm). Station 1 (47°41/33’’ N, 122°24’/18’’ W). Mean depth 23 m, range 15-24 m. Mean particle size 0.139 mm (fine sand). Clay 3.0%, silt 11.1%, sand 84.2%, gravel 1.7%. Station 2 (47°42/16’’ N, 122°26’24’’ W). Mean depth 196 m, range 172-210 m. Mean particle size 0.0067 mm (silt). Clay 34.0%, silt 59.3%, sand 6.8%. Station 3 (47°44/31 N, 122°31'53’/’ W). Mean depth 22 m, range 17-22 m. Mean particle size 0.126 mm (fine sand). Clay 4.7%, silt 11.8%, sand 83.9%. Station 4 (47°44/31"’ N, 122°32'41'’ W). Mean depth 12 m, range 10-18 m. Mean particle size 0.104 mm (very fine sand). Clay 3.0%, silt 18.8%, sand 77.9%, gravel 0.4%. Station 5 (47°10'48’’" N, 122°50’00’’ W). Mean depth 22 m, range 17-37 m. Mean particle size 0.43 (medium sand). Clay 4.5%, silt 5.5%, sand 82.8%, gravel 7.2%. Station 6 (47°10'40’’ N, 122°48’48’’ W). Mean depth 35 m, range 30-40 m. Mean particle size 0.22 mm (fine sand). Clay 5.9%, silt 20.5%, sand 66.2%, gravel 7.5%. Station 7 (47°13’30’’ N, 122°49’36’’ W). Mean depth 68 m, range 47-84 m. Mean particle size 0.009 mm (silt). Clay 26.7%, silt 59.3%, sand 13.8%, gravel 02%: Station 8 (47°07/24"’ N, 122°50’06’’" W). Mean depth 15 m, range 9-21 m. Mean particle size 0.837 mm (medium sand). Clay 3.6%, silt 7.8%, sand 79.4%, gravel 9.7%. NO. 3667 POLYCHAETES—BANSE AND HOBSON 3 The annual temperature range of the shallow stations during 1964 was 8-13° C, that of station 7 was 8-12° C, and that of the deepest station (2) was 8-10.5° C. The average salinity for stations 1-4, off Seattle, was 29.8°/o0; that for stations 5-8 in the southern Sound (Case Inlet) was 29.2, with very little variation. At station 8, the shallowest of the eight stations, the seasonal range was 28.0 to 29.7°/oo S, and at station 2, the deepest one, it was 29.4 to 30.8°/o0S. The oxygen concentrations were between 58% and 87% of saturation. The data of 1964 were representative for other years (Lie, in press). Water movement near the sea bed was always noticeable at the stations during studies with a television camera. The preparation of this paper was supported by U.S. Public Health Service Grant No. GM 10817 to K. Banse and U. Lie, and National Science Foundation Grant No. GB-4902 to K. Banse and M. M. Pamatmat. We are indebted to these agencies for the support, to Mr. U. Lie for the material, to Mr. F. H. Nichols for valuable dis- cussions, and to Drs. M. L. Jones and M. H. Pettibone for critical reading of the manuscript. The Allan Hancock Foundation, Los Angeles, the Academy of Natural Sciences, Philadelphia, the Natur- historiska Riksmuseum, Stockholm, and the U.S. National Museum, Washington, D.C., kindly lent us specimens for study. Family PoLYNOIDAE Eunoe uniseriata, new species FicureE |] Eunoe sp. I Banse et al. [in press]. Typrs.—Holotype: USNM 36273, from Station 2, 47°42/16” N, 122°26'24’’ W (November 1963). Paratype: USNM 36274 (1, in frag- ments) from Station 7, 47°13’30’’ N, 122°49’36’’ W (February 1963). There are eight anterior and some posterior fragments from stations 2 and 7. The holotype has 18 setigers and is 12 mm long without cirri. It is 7 mm wide without cirri or setae and 2.5 mm without parapodia. The width of a mature female (paratype, 32 setigers) is 8 and 3 mm, respectively. DescripTion.—The prostomium (fig. 1a), slightly broader than long, is divided by a mid-dorsal furrow, is blunt anteriorly and without peaks. There are two pairs of eyes, which, after two years of preser- vation, have disappeared in some specimens. The appendages of the prostomium bear scattered short filiform papillae. The antennae arise from large cirrophores. The median antenna of the paratype is as long as the tentacular cirri. The lateral antennae insert termino- ventrally. The styles are disproportionately thin and short compared with the cirrophores. The palps are about 4.5 mm long. There are two smooth spines in the base of each dorsal tentacular cirrus. 4 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 The dorsal and ventral surfaces of the body are smooth save for the distinct segmental borders. The first segment with elytra forms a low collar behind the prostomium. Its ventral cirri are elongated, almost approaching the tentacular cirri in length. Median parapodia (fig. 16) have long dorsal cirri with numerous long papillae. Neuropodia e [Marian arartrcrr terre. 0.1 mm TTY wouneeue el” val TTT ca 0.025mm i a | (d,e,g,h) Ficure 1.—Eunoe uniseriata, new species: a, dorsal view of anterior end, without elytra; b, anterior view of seventh parapodium; c-e, notoseta, with detailed views of tip and middle section; f-h, upper neuroseta, with details of scales (g, optical section); 7, lower neuroseta; j-/, elytron, microtubercles, and macrotubercle. are longer than the notopodia and terminate in finger-like extensions above the aciculae. The ventral cirri are smooth. The presence of elytrophores on a tail end (paratype) indicates that the elytra cover most of the body. All setae have entire tips. Notosetae are stouter than neurosetae; they are yellowish and finely serrated almost to the pointed tip; the distal rows of spines are irregularly spaced (figs. 1c-e). There are two kinds of neurosetae. The upper neurosetae (figs. 1f-h) have two longi- tudinal rows of spinose scales. Similar but more delicate scales are NO. 3667 POLYCHAETES—-BANSE AND HOBSON 5 present on the lower neurosetae (fig. 12); with spines, they are 20u-25y long. It could not be made out whether there are two or three rows, but in any case, the rows of scales are restricted to one-half to three- fifths of the circumference of the setae so that, in side view, usually only the cutting edge of the seta appears as pectinated. The spines on these scales are finer than those of the upper neurosetae, and there are almost twice as many spines per scale. Only two thin, wrinkled, and colorless elytra (fig. 17) are left. Among fairly numerous conical microtubercles of 20y-30y height (fig. 1k) there are some filiform papillae, which become very numerous toward the outer margin of the elytron. Very conspicuous is a row of conical macrotubercles about 110y high (fig. 10). The color of the body is yellow brown in reflected light. The para- podia of the paratype, collected in February, are filled with exceedingly numerous, loose, and almost round eggs of about 654 diameter from setigers 7 or 8 onward. The fragment of a juvenile collected in July, with a width of almost 2 mm with parapodia but without cirri, and 0.7 mm without para- podia, shows the same characters as the adults, except that the prostomium is more pointed and the number of setae is small; all elytra had been lost. The name refers to the row of macrotubercles on the elytra. Diaenosis.—A Eunoe species with blunt prostomium and dispro- portionately slender paired antennae. Notosetae with pointed tips. Two kinds of neurosetae. Elytra with conical microtubercles and one row of conical macrotubercles, and numerous filaments on the margins. DIFFERENTIAL DIAGNOSIS.—Hunoe uniseriata is best characterized by its blunt prostomium, the slender paired antennae, the arrange- ment and form of the elytral macrotubercles, and the tips of the noto- setae. There are few species with a blunt prostomium or reduced prostomial peaks among the Hunoe species listed by Hartman (1959, 1965b). Among these, H. ewra Chamberlin, E. sentiformis Ushakov, E. shirikishinai Imajima and Hartman, F. spinicirris Annenkova, LE. subtruncata Annenkova, and possibly E. oerstedi Malmgren have notosetae with blunt tips, which is not the case in EL. wniseriata. Notosetae are pointed in /. crassa (Treadwell) and E. nodosa dybow- skyi Augener, but the conical and other macrotubercles on the elytra are arranged differently from those on EH. unisertata. Hunoe nodosa (Sars) as described by Pettibone (1954, 1963) can have a blunt prostomium; there are papillae, however, in addition to the dorsal tubercles above the base of the dorsal cirri. Further, the very con- spicuous row of macrotubercles near the external borders of the elytra are broad and rounded or jagged, not pointed as in the present species. Only one kind of neurosetae has been reported for . nodosa. 6 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 Harmothoe fragilis Moore Evarnella fragilis —Hartman, 1959, p. 69. Harmoihoe fragilis —Pettibone, 1963, p. 39. One damaged incomplete specimen with eggs was collected in February 1963. Only the supracicular neurosetae are bidentate. The single remaining elytron conforms to figure 4, table 1, by Annenkova (1937) and figure b, table 6, by Imajima and Hartman (1964) for H. impar (Johnston). Pettibone (1963) already included Annenkova’s record (as reported by Ushakov, 1955) in the synonymy of H. fragilis. We consider our specimen to belong to Moore’s species because the European representatives of H. wmpar as described by McIntosh (1900) and Fauvel (1923) have inverted-conical or droplike papillae on the margins of the elytra, which is not the case with H. fragilis. Imajima and Hartman (1964) have doubted the justification of the genus Evarnella Chamberlin, and we follow them herein. Found on station 6. New for Puget Sound. Previously recorded from southern California and the northwestern Pacific. Hesperone complanata (Johnson) Johnson (1901) stated that the paired antenna is less than half the length of the prostomium (cf. Pettibone, 19538, pl. 18: fig. 155). Johnson’s largest animal was 21 mm long. In a small, incomplete animal, as well as in an incomplete specimen of 30 mm length with 26 segments, and in another fragment of about the same size, we find that the antennae are slightly more than half as long as the prostomia. Family POLYODONTIDAE Peisidice aspera Johnson A 5-mm-long specimen collected in May contains moderately numerous polygonal eggs of up to 105u x 95y, and one-third to one- half of this in the third dimension. Four specimens of about 5 mm length, collected in daytime, have empty intestines, excepting one from station 6 that contains two sand grains of about 0.35 mm diameter. Family CHRYSOPETALIDAE Paleanotus bellis (Johnson) Paleanotus bellis—Hartman, 1961, p. 57. In California material examined by Hartman (1961), the ventral tentacular cirri of the right side were lacking, and this asymmetry was thought to be characteristic of the species. The type-locality is Monterey, Calif. In Puget Sound material, the cirri are present in NO. 8667 POLYCHAETES—BANSE AND HOBSON 7 five animals (apparently damaged in one); the cirrus is rudimentary in one. Thus, asymmetry of the front end is not the rule. Family PayLLopociDAE Eulalia (Hypoeulalia) bilineata (Johnston)? Eulalia bilineata.—Imajima and Hartman, 1964, p. 61. Although there are great similarities of characters, the first sezments in our six specimens are clearly separated from the prostomia as in Eulalia sensu stricto, rather than being fused with them as reported by Bergstrém (1914) for European material. Imajima and Hartman (1964) have found the same separation in Japanese animals, a fact that suggests the Pacific form is not identical with the European species. The ventral cirri of the second tentacular segments of our specimens are not filiform but slightly broader than the others. Eulalia (Eulalia) levicornuta Moore FIGURES 2a-c Eulalia levicornuta Moore, 1909a, p. 346. Not Eulalia levicornuta.—Berkeley and Berkeley, 1943, p. 130. We consider a well-preserved anterior fragment to represent this species although it does not completely agree with the type-specimen (USNM 17288). Some additions to the description follow, based on the Puget Sound specimen. The length (without proboscis, 125 segments) is about 20 mm. The greatest width (without parapodia) is 0.65 mm. The pros- tomium is quadrate except for the narrower anterior part (fig. 2a). The unpaired (broken) antenna inserts between the small eyes. The partially everted proboscis is densely covered with rounded, conical papillae except proximally; the smooth section is shorter than the proboscis is wide. There is a fairly distinct dorsal furrow between the prostomium and the first segment. The first segment is not reduced dorsally and is enlarged laterally. The tentacular cirri are filiform except the ventral one on the second segment, which is slightly flattened. The dorsal tentacular cirri on the second and the third segments are about one-and-a-half times the length of the normal anterior dorsal cirri. There are setae on the second segment. The parapodia (fig. 2b) have rounded lips of equal length. The dorsal cirri are thick and of a broad shape. The ventral cirri are slightly longer than the parapodia. There are approximately 15 setae per parapodium. The shaft of each seta ends in a conspicuous spine with secondary teeth on both sides (fig. 2c); the blade is long. 8 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 The color of the animal is pale brown; the dorsal cirri have a greenish tint. Comparing our specimen with the description by Moore (1909a) we find that the form of the shafts of the setae is characteristic of E. levicornuta. In our animal, the median as well as the accessory spines are slightly stronger than in the type specimen, and are exactly as in Moore’s figure (1909a, pl. 16: fig. 30). Moore stated that although 0.125 mm Figure 2.—Eulalia levicornuta: a, anterior end, from above and slightly from the left (proboscis and parapodium with ventral cirrus on third segment omitted); b, posterior view of 80th parapodium; c, terminal end of shaft of seta. Lulalia parvoseta, new species: d, dorsal view of anterior end (all dorsal cirri and part of tentacular cirri missing); e, papilla of proboscis; f, anterior view of 19th parapodium; g, seta. the first segment is usually sharply separated from the prostomium, in one of his specimens they were dorsally almost continuous; this approaches the situation in our animal. The ventral cirrus of the second segment is flattened in the Puget Sound specimen contrary to the California material, but this character may be of lesser im- portance in view of the variability reported for E. viridis (Bellan, 1964). The length of the parapodia of our specimen is about one-third the body width, not one-fourth as with Moore’s animals. The dorsal cirri in the type are rather leaflike, whereas in out animal they are fleshy. NO. 8667 POLYCHAETES—BANSE AND HOBSON 9 Found on station 2. New for Puget Sound. Previously known from California. Berkeley (1924) had recorded a specimen for the coast of Vancouver Island but did not include it in the “Canadian Pacific Fauna” (Berkeley and Berkeley, 1948). This specimen is not in the Berkeley Collection of the U.S. National Museum (Dr. M. H. Petti- bone, pers. comm.). Two specimens collected in Wakeham Bay, near Ungava (Canada), on 2 September 1927 by Johansen and identified by Berkeley and Berkeley (1943) as E. levicornuta (USNM 32672) belong to another species of Eulalia sensu stricto. Among other characters, the shafts of the setae end in long spines of equal thickness. Eulalia (Pterocirrus) parvoseta, new species FiaureEs 2d-g Eulalia (Pterocirrus) sp. I Banse et al. [in press]. Holotype, USNM 36269, from station 5, 47°10’48’’ N, 122°50’00’’ W (February 1963). There is one complete specimen (holotype), with about 50 segments, of 3.0 mm length (without proboscis) and 0.55 mm greatest width (without parapodia). An anterior fragment (fig. 2d) from station 4 with seven setigerous segments of slightly greater width was lost after the figure had been drawn. DeEscriPpTION.—The oval prostomium carries five long antennae. The unpaired antenna inserts somewhat posterior to the paired ones and reaches the third segment. The small eyes, without lenses, are near the posterior margin of the prostomium. The everted proboscis is 1.5 mm long and is densely covered with clavate papillae (fig. 2e), about 50p high. The first two segments seem to be fused ventrally. The first one is reduced dorsally. The four pairs of tentacular cirri originate from large cirrophores; that of the ventral cirrus of the second segment (V. II) is supported by a large acicula. All tentacular cirri are filiform except V. II, which is leaflike with a tip; its posterior margin is thicker than the anterior margin. The tentacular cirrus of the first segment is about as long, the cirrus of the second segment is about twice as long, and the one of the third segment is about two-and-one- half times as long as V. II (see fig. 2d). There are no setae on the second and third segments, but on the third segment there is a reduced parapodial lobe, about two-fifths the length of the following one. Its acicula is much thinner than that in the cirrophore of V. II. : 01 Ol The tentacular formula thus is 140570" The parapodia have rounded lips of about equal length. The dorsal cirri are broadly lanceolate (fig. 2f); toward the posterior end of the 10 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 125 animal their length is about twice their width. The ventral cirri are oval and slightly longer than the parapodia. In the type specimen, the number of setae per parapodium is about 10, and in the second fragment it is about 15. The ends of the shafts of the setae are 6u-7y thick and have large fangs with a few secondary teeth (fig. 29). The blades are very short (20u—25y). Anal cirri were lost. The color of the animals is pale greenish with marked black pig- ment bands dorsally on the posterior margin of each segment. The name refers to the noticeably short blades of the setae. Diacnosis.—A small Pterocirrus species with an oval prostomium, small eyes, and a long unpaired antenna. Clavate papillae on proboscis. First segment dorsally reduced. Shafts of setae with large fangs and very short blades. DIFFERENTIAL DIAGNOSIS.—The species belongs to the subgenus Pterocirrus Claparéde of Eulalia, as defined by Banse (1959; see also Day, 1960). It is distinguished from the type-species, E. (Pterocirrus) macroceros (Grube) (cf. Banse, 1959), by the oval shape of the pros- tomium, the small size of the eyes, the clavate papillae on the proboscis, the toothed tips of the shafts, and the short blades of the setae. Possibly also H. marginata Claparéde (see Rullier, 1964) belongs to Pterocirrus as defined now, although Claparéde (1868) stated that all cirrophores of the second and third segments are supported by aciculae. The new species differs from EH. marginata in this character, the small eyes, the toothed ends of the shafts, and the short blades of the setae. Phyllodoce (Anaitides) nr. multiseriata Rioja FIGURE 3a Phyllodoce (Anaitides) multiseriata Rioja, 1941, p. 684. There is one broken specimen, presumably a mature male, from station 2, August 1963. The animal is at least 25 mm long and about 0.75 mm broad (without parapodia). It is particularly distinguished by the proximal papillae of the proboscis, which form seven or eight fairly irregular rows of about nine round papillae each on both sides. The papillae leave broad mid-dorsal and midventral gaps, the dorsal one tending to be V-shaped. Because of its broad base, the shape of the dorsal gap is not nearly as regular as shown by Rioja (1941, pl. 1: fig. 2). There is a tiny nuchal papilla. Only the ventral tentacular cirri of the second segment are left, which are filiform and three- fourths as long as the body is wide. Setae start on the third tentacular segment. The parapodia have large supra-acicular lips (fig. 3a). Cirri are preserved only in median-posterior segments. There are about a dozen setae per parapodium. Their shafts end in numerous small teeth; the NO. 3667 POLYCHAETES—BANSE AND HOBSON 11 blades are long. The color of the animal is pink with dark interseg- mental furrows. The form bears some resemblance to P. multiseriata Rioja, which, however, has 12 pairs of rows of papillae with 11-15 papillae in the long rows. Also, Rioja (1941) emphasized the V-shaped form of the dorsal gap. On the other hand, the large supra-acicular lips of the parapodia (stressed by Rioja in the text rather than in the figures) support our tentative identification. Hartman (1961) has described Ficure 3.—Phyllodoce nr. multiseriata: a, posterior view of medioposterior parapodium. Gyptis brevipalpa (setae not drawn to scale): b, anterior view of median parapodium; c, thick notopodial spine; d, forked notopodial seta; e¢, tip of neuropodial compound seta; f-h, notopodial setae, including a thick capillary seta, from type of G. brevipalpa (figs. f-h by G. Hartmann-Schréder). Micropodarke dubia: i, posterior view of median parapodium; j, compound seta. Sigambra tentaculata: k, seta and acicula of a median notopodium squeezed under the cover glass (outline of basis of dorsal cirrus sketched). an Anaitides nr. multiseriata with 12 pairs of lateral papillae. Her specimen had a nuchal papilla that in Rioja’s animals was reported to be wanting.. Our form is not similar to any other known member of the subgenus Anaitides from the temperate northeastern Pacific. Phyllodoce (Anaitides) williamsi (Hartman) Anaitides williamst Hartman, 1936, p. 126. Our six specimens have about nine papillae per row on the proboscis, approximately rectangular dorsal cirri, rounded ventral cirri, and three bars across the dorsum of median segments, all of which support the identification. Adding to the description, we note that there are no 12 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 setae on the second tentacular segment, as to be expected in the sub- genus Anaitides. A female with polygonal eggs of up to 75u diameter was collected in February. Found at stations 1, 5, and 8. New for Puget Sound. Previously known from Oregon and California. Family HEsIONIDAE Gyptis brevipalpa (Hartmann-Schréder) FicurREs 3b-h Ozydromus brevipalpa Hartmann-Schrider, 1959, p. 105. Oxydromus arenicola glabra Hartman, 1961, p. 68. Many specimens are available that are referred to G. brevipalpa, originally described from El Salvador, despite minor differences in the notopodial setae. The largest, though incomplete, specimen is 17 mm long. No cilia are visible between the 10 widely spaced terminal papillae of the proboscis. Occasionally the unpaired antenna seems to arise from a small elevation rather than from the straight anterior margin of the prostomium. Also, the posterior margin of the prostomium is straight. The longest tentacular cirri reach the twelfth setiger. Tentacular as well as dorsal cirri on body segments can be annulated with the joints being slightly shorter than wide, or they can appear pseudoannulated. The dorsal cirri are alternately short and long; the long cirri surpass the short cirri by about one-half their length and are almost as long as the body (without parapodia) is wide. Anal cirri are smooth. Notosetae are absent on the first three or four setigers although aciculae are present dorsally. In posterior notopodia (fig. 36) there are three to four short, nearly smooth spines about one-half as thick as the aciculae (fig. 3c) and about twice as many forked setae with flattened blades and short spurs, distinctly serrated (fig. 3d). The greatest width of the blades is about 5y. In addition, two very long slightly curved capillaries can occur, which are finely serrated on the convex sides. The neurosetae have serrated blades that are almost smooth distally. The tips are excavated (fig. 3¢). The blades may occasionally have hoods as indicated by the broken line in the figure; also one of the notopodial forked setae was found with such a hood. The tips of the shafts of the ventral neurosetae are slightly bifid when seen from below. The proximal parts of all setae, but not of aciculae, are distinctly 2-layered, the core being transversely barred. A specimen collected in February is packed with many polygonal egos of about 60u-70y diameter. Dr. G. Hartmann-Schréder inspected the type of G. brempalpa, provided drawings, and permitted us to add the following to the description: NO. 3667 POLYCHAETES—BANSE AND HOBSON 13 Notopodial setae appear on the fifth setiger. There is an upper short, needle- like bristle, subdistally finely serrated [fig. 3f]. There are two thin, smooth cap- illaries almost twice as long [fig. 3g]. There are usually three forked bristles which appear to be smooth with 20 X 100 magnification [fig. 3h]. With the same mag- nification, a very fine serration of the blades of neuropodial setae can barely be seen. The type of O. arenicola glabra was studied at the Allan Hancock Foundation and was found to have forked setae, as in figure 3d, from the fourth and fifth setigers. Also, some neuropodial setae are hooded. Another specimen, studied in sample 4829, has notopodial setae as in figure 3c. The forms differ from G. arenicola (La Greca) in the absence of cilia on the proboscis and the presence of the forked notosetae. Apparently, the articulation of cirri is not a useful character, in view of the variability of the present material and our experience with Micropodarke dubia, wherein regenerating cirri are not clearly annu- lated (see below). The insertion of the unpaired antenna does not seem to be of significance either, Neither Dr. O. Hartman nor we wish to separate material from California and Washington from the Central American species on the basis of details in notopodial setae, Gyptis capensis (Day) seems very close to G. brempalpa and is perhaps identical. Found at stations 2, 4, 5, 7, and 8. New for Puget Sound. Previously known from California and Central America, Micropodarke dubia (Hessle) FIGURES 37, 7 Kefersteinia dubia Hessle, 1925, p. 32. Micropodarke dubia.—Imajima and Hartman, 1964, p. 83. There are many broken specimens that agree with the description by Hessle (1925) except that the everted proboscis has about 25 fairly regularly spaced terminal papillae with some additional super- numerary ones, rather than ‘about 20.” Adding to Hessle’s description, the dorsal tentacular cirri can be quite long, the longest reaching the fourteenth setigerous segment. Regenerating cirri are not clearly annulated. The ventral tentacular cirri are about as long as the body is wide. Lobes that originate slightly posteriorly to the bases of the parapodia reach two-thirds the length of the parapodia in the first three or four setigerous seg- ments; they are about four times as long as they are wide. Posteriorly they are much shorter (fig. 37). The anterior lips of the first parapodia are almost bilobed as described by Okuda (1938) for M. anemiyar Okuda (this record has already been considered a synonym of M. dubia 14 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 by Imajima and Hartman, 1964). A compound seta is shown in figure 3). Fairly numerous polygonal eggs of up to 60u diameter are filling the parapodia of a specimen collected in May at station 5. The species seems to be a selective deposit feeder: Intestines of specimens from station 5 (mean particle size 0.45 mm) are primarily filled with fine materials; in addition, there are sand grains smaller than 0.05 mm. Some sand grains and diatom frustules of more than 0.1 mm shortest dimension are also present. There is no doubt that our specimens belong to Hessle’s species; however, the genus Micropodarke Okuda, based on only one specimen of M. anemiyai, is said to have no notopodial aciculae, that is, to have uniramous parapodia. In view of the variability in the number of notopodial aciculae reported by Hessle (1-3; in the specimens checked by us, 1-2), it is feasible that Okuda’s specimen was exceptional. If this is the case, the generic diagnosis for Micropodarke should be amended. The genus would then be very close to Nereimyra, except that there are no jaws in Micropodarke. Also N. punctata (Miiller) has the mentioned lobes (fig. 37) mediad to the ventral cirri (Banse, 1956). Other close genera are Parasyllidea Pettibone and possibly Neopodarke Hartman, both with slightly different termination of the proboscis; the latter genus lacks notopodial aciculae. Since the armature of the proboscis is not a very practical character, it is doubtful that all of these mostly monospecific genera should be retained. Nereimyra would take precedence over the other names. Found at stations 1, 4, 5, and 8. New for the eastern Pacific. Pre- viously known from Japan. Family PILARGIDAE Sigambra tentaculata (Treadwell) FIGURE 3k Ancistrosyllis tentaculata—Hartman, 1947a, p. 498. Sigambra tentaculata.—Pettibone, 1966, p. 182. There are several animals. The longest one has at least 135 setigers and is 2 cm long. The deeply indented posterior margin of the prostomium is rarely visible. The proboscis has 12-13 triangular closely spaced terminal papillae of equal size and also some medium-sized papillae on the outside. There seems to be some variability of this character—14 in the type (Pettibone, 1966), 8 in other Atlantic material (Hartman, 1965a). The curved notopodial hooks are usually found from the third setigerous segments onward, rather than from the fourth as in the type material from Long Island Sound (Pettibone, 1966); occasionally they begin with the fourth setigerous segment in our material. A fine NO. 3667 POLYCHAETES—BANSE AND HOBSON 15 short capillary seta may be present (fig. 3k). In the median and posterior parapodia short, thick spines occur, which are not as strongly bent as the aciculae. Neurosetae are all spinose. In short ones, the hairs are as long as the shafts are wide, so that the setae appear comblike. In the longest thin setae, the serration becomes just visible under 400 magnification. Found at stations 6-8. New for Washington waters. Previously known in the North Pacific from southern California. Family SYLLIDAE Eusyllis blomstrandi Malmgren FIGURES 4a-c Adding to the description of the Pacific and Arctic material (Petti- bone, 1954), we note that the pharynx opening in our material is Figure 4.—E£usyllis blomstrandi: a, opening of pharynx; b and c, setae from a median parapodium. LExogone lourei (setae not drawn to scale): d, tip of upper simple seta, e, compound seta with short blade. Pionosyllis uraga: f, posterior view of median parapodium with a dorsal acicula (neurosetae not shown); g, compound seta with long blade. Syllis harti: h, partly everted pharynx from the lower right hand side (the palps are indicated); 7, blade of median seta from 41st parapodium of type. Dorvillea pseudorubrovittata (setae not drawn to scale): 7, tip of simple, flattened seta; 8, compound seta. 16 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 chitinous with the usual large dorsal tooth and small teeth on about three-fourths of the circumference (fig. 4a); it thus differs from the material of Malmgren (1867) and Imajima (1966). Two rows of soft papillae encircle the outside of the pharynx. The proventricle has 50 to 70 rows of papillae and is followed by a pair of spherical caeca. The dorsal cirri, annulated in the anterior region of the body, are smooth in the middle and posterior regions. Setae from the middle region of the body have short blades (figs. 4b, c). Setae with long, straight blades as reported by Berkeley and Berkeley (1945) for animals from Vancouver Island are absent in our animals. Swimming setae occur from setigers 17 or 18 onward. Exogone lourei Berkeley and Berkeley Figures 4d-e There are several specimens. An inspection of the type (USNM 32895) showed that the proventricle is rather long, extending from the third to the sixth setiger (fourth to seventh in our specimens). In a median parapodium of one of our specimens, the tip of the simple dorsal seta is slightly serrated (fig. 4d); compound setae with short blades have small teeth above the main fangs (fig. 4e). Pionosyllis uraga Imajima Figures 4f, g Pionosyllis uraga Imajima, 1966, p. 114. There are many anterior fragments with up to about 30 setigers, 4 mm long and 0.7-0.8 mm at greatest width (without parapodia). Commenting on the description by Imajima (1966), we note that the proventricle of our specimens has about 35 rows of papillae and extends through 12-14 setigers. The dorsal cirri alternate in length. The long ones of the anterior setigers as well as the dorsal tentacular cirri are 1.5-2 mm long. Farther posteriorly, the dorsal cirri decrease in size and are twice the body width at about the thirtieth setiger although there are cirri of 2 mm length even on the twentieth setiger. Neuropodial setae are numerous in the anterior 12-15 setigers and appear faintly yellow in direct light. Subsequently the number of setae is sharply reduced to approximately 20 or less. In this region, there are up to four neuropodial aciculae. In one animal there is a dorsal acicula but natatory setae are absent (fig. 4f). The uppermost setae with very long blades (about 100u, fig. 4g) have faintly serrated cutting edges at their bases and bidentate tips. The compound setae with blades of ordinary length (25u-40u) have strongly spinose cutting edges, except the lowermost ones with the shortest blades. The ends of the shafts of these setae are slightly serrated but do not NO. 3667 POLYCHAETES—BANSE AND HOBSON 1% have hairs as stated by Imajima (1966). There is very faint serration at the tips of the shafts of the upper compound setae as well. Found at stations 5 and 6. New for the eastern North Pacific. Previously known from Japan. Syllis (Typosyllis) harti Berkeley and Berkeley Ficurss 4h, 7 Our many specimens have been compared with the type material (USNM 32671), and we can add to the description by Berkeley and Berkeley (1938): The pharnyx extends through the tenth to twelfth setigers. There are the usual 10 soft papillae on the outside and, alternating with them, rows of four small soft papillae (fig. 4h). The proventricle has 40-60 rows of papillae and extends in the type through 12 setigers and in our material through 9-11 setigers. The ends of the shafts of median (fig. 42) and lower setae from the thirteenth and forty-first feet of the type bear minute teeth. The blades are coarsely serrated and have subbidentate tips. The blades of the upper setae are about twice as long as those of the median setae but are less coarsely serrated and have bidentate tips (fig. 47; see also original description, Berkeley and Berkeley, 1938). The species reproduces by stolons. The intestines of all our speci- mens are empty. Family NEREIDAE Platynereis bicanaliculata (Baird) Platynereis dumerilit agassizi—Berkeley and Berkeley, 1948, p. 60. Paragnath groups VII and VIII of specimens from the San Juan Archipelago collected by M. H. Pettibone (USNM 28458, 28460, 28463, and 28464) form five subgroups of pectinate paragnaths (sometimes in a double row), rather than a continuous band as stated by Hartman (1954) for P. bicanaliculata from Vancouver Island. Ehlers (1868) described six subgroups of two rows each for his P. agassizt from the Strait of Georgia and from Mendocino (see also Izuka, 1912). Family NEPHTYIDAE Nephtys assignis Hartman Nephtys assignis Hartman, 1950, p. 112. A fairly relaxed specimen with about 145 setigers is just 20 cm long. Found at stations 2 and 7. New for Puget Sound. Previously known from southern California and Guatemala. 313-605—68——2 18 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 125 Nephtys ferruginea Hartman This species has not been reported previously to possess recurved aciculae, but they are clearly recurved in our many specimens. The interramal cirri usually point downward or may appear to be involute; however, the pigmentation pattern, the appearance of the interramal cirri on the third setiger, and the shape of the posterior neuropodial acicular lobe make identification certain. Usually, the intestinal content is not conspicuous even in trans- parent posterior ends. A specimen with sediment boluses in the pos- terior intestine was dissected and was found to have numerous syllid bristles engulfed in the sediment. Females with eggs and young specimens, the latter presumably be- longing to this species, were observed from February through July. Family SPHAERODORIDAE Sphaerodoridium sphaerulifer (Moore) Spaerodorum sphaerulifer— Ushakov, 1955, p. 222. Spaerodoridium sphaerulifer —Liitzen, 1961, p. 415. Found at stations 4 and 7. New for Puget Sound. Previously known from California, the Sea of Okhotsk, and the Sea of Japan. Family GLYCERIDAE Glycera Savigny In addition to a note on G. siphonostoma (della Chiaje), we report observations on the intestinal contents of other species of this genus, occurring locally. Glycera siphonostoma (delle Chiaje) Glycera siphonostoma. —Imajima and Hartman, 1964, p. 164. Our specimens agree with the description by Imajima and Hartman (1964): Each aileron has a connecting plate, there are ridges on the proboscidial organs, the prostomium is 9-ringed, branchiae are absent, there are pointed postsetal lobes that are much shorter than the pre- setal lobes, and there are large ventral cirri. The ventral cirri of middle parapodia of our specimens reach nearly to the presetal lobes, agreeing with the figures given by McIntosh for the species (1910; pl. 76; fig. 2b). Found at stations 2, 7, and 8. New for Puget Sound; hitherto known in the Pacific from Japan. NO. 3667 POLYCHAETES—BANSE AND HOBSON 19 Intestinal Contents of Local Glycera species Intestinal contents of the midportion of animals were studied, or Squeezed specimens were inspected (as indicated). All the material had been collected in the daytime. Three specimens of G. americana Leidy (about 3.5, 5, and 10 cm long) had empty intestines. A 5-cm- long specimen of G. capitata Oersted (including G. nana Johnson; see Banse et al., in press) had an empty intestine as seems to be the rule with the species (from inspection). A 4-cm-long animal from sta- tion 7 contained a few boluses of sediment that were full of setae of Sigambra tentaculata, together with fine sediment material and sand grains up to about 0.075 mm diameter. Specimens of G. sipho- nostoma (two of 2 cm, one of 6 cm length; from inspection) had empty intestines, A 35-cm-long specimen of G. robusta had an empty middle intestine; a 25-cm-long animal contained a small amount of sediment particles together with many bristles, possibly of Aphroditidae. The observations of largely empty intestines corroborate the findings by earlier investigators (see Klawe and Dickie, 1957; Sanders et al., 1962). According to these authors, who quote earlier studies, Glyceridae may be considered deposit feeders that swallow larger animals incidentally. Our material is not large enough to investigate whether the grain size composition of the intestinal content is that of the habitat; the organization of the proboscis would suggest non- selective deposit feeding only. Family GONIADIDAE Goniada maculata Oersted Goniada maculata.—St¢p-Bowitz, 1941, p. 209.—Hartman, 1950, p. 20. Found at stations 1, 3, 4, 6, and 8. New for Puget Sound. Pre- viously known in the Pacific from Kodiak Island and the northwestern Pacific. Family LuMBRINERIDAE Lumbrineris Blainville In addition to a morphological note on L. californiensis Hartman, we make observations on the intestinal contents of several other species of this genus occurring locally. Lumbrineris californiensis Hartman Lumbrineris californiensis Hartman, 1944, p. 163. In specimens of 4-5 cm length and about 1 mm width (without parapodia), composite hooks are found only to the fifteenth or six- 20 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 teenth setiger, rather than to the twenty-fourth to thirtieth or a little beyond as in California material 8.5-13 cm long (Hartman, 1944). Found at stations 1, 3-6, and 8. New for Puget Sound. Previously known from California. Intestinal Contents of Local Lumbrineris species From inspection of entire animals, we note that the intestines of specimens of the local species of one to a few cm length are filled only to about one-fourth of their length, sometimes less. The sediment occurs in boluses at least in the middle and posterior sections of the body. Dissection of two specimens each of four species showed the following (width is without parapodia; all animals had been collected in the daytime): LUMBRINERIS BICIRRATA TREADWELL.—The intestinal contents of a posterior fragment from station 4 of 3 mm width contained sand grains up to 0.2 mm wide, inbedded in a fair amount of organic and inorganic fine material. The intestinal contents of an almost 10-cm- long specimen of 6 mm greatest width from station 2 were a large amount of fine sediment with some diatom frustules and sand of up to 0.15 mm diameter. In neither case could recognizable organic remains be seen. LUMBRINERIS CALIFORNIENSIS HartMAN.—The hind intestine of a small individual (about 1 mm wide) from station 7 contained very fine sediment of less than 0.01 mm grain size and two healthy locking nematodes. A specimen about 1.5 cm long and 1 mm wide from station 1 contained fine detrital material, with a few sand grains up to 0.15 mm in diameter, and several pieces of a terrestrial plant epidermis about the same size. LUMBRINERIS CRUZENSIS HARTMAN.—A young specimen about 1 em long in length and <1 mm wide from station 1 contained fine detrital material, with a few sand grains <0.07 mm in diameter. Many other specimens of approximately the same size from this station and station 7 had empty intestines. LUMBRINERIS LUTI BERKELEY AND BERKELEY? (for identification, see Banse et al., in press).—T wo posterior fragments 1.5 cm long and about 1 mm wide from station 4 contained largely coarse material, including unbroken Coscinodiscus species frustules 0.2 mm in diameter. Most specimens had sediment about the same grain size or particles smaller than the mean grain size of their habitat in their intestines, suggesting predominantly selective feeding. Because of the partial filling of the guts, it may be doubtful whether the species are ex- clusively selective deposit feeders as these have to cope with large amounts of sediment to obtain enough organic material. The absence ; : | | NO. 3667 POLYCHAETES—BANSE AND HOBSON 21 of animal remains (except for one specimen of ZL. californiensis), is certainly striking. Harlier studies have suggested that Lumbrineris species are not carnivorous: L. impatiens Claparéde feeds on macroscopic algae (Hempelmann, 1931); L. minima Hartman is a detrital feeder (deposit feeder) (Reish, 1959); LZ. tenuis (Verrill) and probably L. fragilis (Miller) are selective feeders as is the related Drilonereis longa Webster (Sanders et al., 1962). Family ARABELLIDAE Notocirrus californiensis Hartman Notocirrus californiensis Hartman, 1944, p. 175. Found at stations 1, 3, 4, and 6. New for Puget Sound. Previously known from California. Family DorvVILLEIDAE Dorvillea pseudorubrovittata Berkeley Fiaures 4j, k The setae were studied in the syntypes (USNM 32674). There are some superior thin capillary setae, slightly broadened in the distal third, where they are finely serrated The bidentate tips have the same outline as those of the thicker spines below (fig. 47), which are slightly broadened and coarsely serrated in the distal halves. The serration near the bifid tips is fine. Rioja (1962) has not mentioned the bifid tips. The ends of the compound setae are serrated (fig. 4k). The spurs below the secondary teeth of the blades can be as small as in our figure, or they can form a membrane on the cutting edges of the blades reaching their tips. Family PARAONIDAE Aricidea (Aricidea) ramosa Annenkova Figures 5a-d Aricidea? (Aedicira) ramosa.—Hartman, 1957, p. 312, p. 327; 1959, p. 371. Aricidea (Aedicira) ramosa.—Hartman, 1963, p. 37. Aedicira ramosa.—Hartman, 1965b, p. 50. We have 15 individuals, 2 of which are complete and have about 80 setigers. The antenna is short and palmately divided into three to five fairly equal lobes (fig. 5a); Annenkova described longer lobes. There are 13-17 pairs of gills beginning on setiger 4. Notopodial 2, PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 postsetal lobes are short in the anterior and longer in the posterior segments. The character of the neuropodial setae changes at the fortieth to fiftieth setigers, where some setae taper abruptly (fig. 50). Two or three setigers farther, modified setae, usually with an arista, are also present (fig. 5c). The pygidium of one specimen is well preserved and has three ventral cirri, a short median one and two longer ventrolateral ones (fig. 5d). Ficure 5.—Aricidea ramosa: a, dorsal view of anterior end; b, seta from about the 40th neuropodium; c, modified neuropodial seta; d, posterior view of pygidium. Paraonis lyra: e, furcate seta from posterior notopodium. Paraonis ivanovi: f, posterior neuro- podial hook. The neuropodial hooks place this species in Aricidea sensu stricto not in Aedicira as tentatively suggested by Hartman (1957). Found at stations 4, 6, and 7. New for Puget Sound. Previously known from the Sea of Japan and southern California. Paraonis (Paradoneis) lyra Southern FIGURE 5e Paraonis (Paraonides) lyra Southern, 1914, p. 94. Paradoneis lyra.—Hartman, 1965a, p. 140. Our single specimen has a prostomium longer than wide. It lacks the ciliated papilla on the anterior margin figured by Southern (1914), but this may be an artifact of preservation. There are 14 pairs of NO. 3667 POLYCHAETES—BANSE AND HOBSON 23 gills beginning on the fourth setiger. Furcate setae are present in the notopodia beginning on the last few branchial segments (fig. 5e). They differ from those figured by Southern in having spines on the inside of both arms, rather than on the inside of the long arm only. Found at station 4. New for the temperate northeastern Pacific. Previously known from the northern Atlantic and southern California. Paraonis (Paraonis) ivanovi Annenkova Figure 5f Paraonis ivanovi—Hartman, 1957, p. 333. Our specimens have conical prostomia and no eyes, five to six prebranchial setigers, and 10-16 (usually 13) pairs of gills. The modified neuropodial, acicular setae are curved, and have hoodlike structures on the convex sides. The hooks resemble the figure of Annenkova (1934, fig. la) for P. wanovi under X 400. Under X 1000, however, the hood appears to be feathered (fig. 5f), and further, it is noticeable only on the convex side of the hook, whereas Annenkova showed it on both sides of the hook. The pygidium of one specimen consists of one large median ventral lobe from which arise two lateral cirri. Ushakov (1950) regarded P. wanovi and P. gracilis Tauber as the same species; however, we find that the modified neuropodial acicular setae of P. gracilis specimens from near the type-locality (kindly sent by Dr. A. Eliason, from his station 2 in the southern Oresund; Eliason, 1920) are without hoods or any similar structures. We therefore consider the two forms to be different species, which are distinguished only by these setae, as far as we can see from the descriptions. The Swedish material did not allow a detailed compari- son of other characters. Found at stations 2-4 and 6-8. New for the eastern Pacific. Pre- viously known from the Sea of Okhotsk and possibly from Murmansk (fide Ushakov, 1950). Ushakov (1955) has recorded P. gracilis sensu lato also from the Bering Strait, and Chukchi Sea. Family APISTOBRANCHIDAE Apistobranchus ornatus Hartman Apistobranchus ornatus.—Hartman, 1965a, p. 144. Numerous specimens were found in January at station 4 but not in April. They are characterized by their palps, if present, reaching to the fourth setiger, the absence of the ventral cirrus on the first setiger, the serrated postsetal neuropodial lamella on the fifth, and the large lobes on the postsetal neuropodial lamella on the seventh setiger. Notopodia 24 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 are present at least through the twenty-second setiger, and interramal cirri are present from the first through at least the seventh setiger. The setae of the first six (sometimes seven) parapodia appear bushy, as in California animals (Hartman, 1961), but contrary to Hartman’s (1965a) description. New for Puget Sound. Previously known from California. Family SPIONIDAE Laonice Malmgren While describing a new species from Puget Sound, we found it necessary to study the types of Z. antarcticae Hartman and L. japo- nica (Moore). We add here to the original descriptions of these species. Laonice antarcticae Hartman Laonice cirrata antarcticae Hartman, 1953, p. 40; 1959, p. 378. Laonice antarcticae—Hartman, 1965a, p. 147. We consider L. cirrata antarcticae Hartman a species of its own as did Hartman (1965a), for the following reasons: In the two types of L. antarcticae (Naturhistoriska Riksmuseet, Stockholm, type no. 613), the dorsal sense organ reaches to about the thirteenth setiger. The gills start, barely visible, on the second setiger. On the fourth setiger, they are one-fourth as long as the notopodial postsetal lamella, and only on the sixth to tenth setiger do they attain the same length as the lamellae. In the segments immediately posterior to the gill-bearing region, the ridges connecting the notopodial lamellae with the dorsum occupy about one-fourth of the distance between the lamellae, as in L. pugettensis, and are otherwise inconspicuous. Hooks in the forty- sixth and fifty-seventh setigers have two teeth, side by side above the main fangs, as in Brazilian material (Hartman, 1965a). Laonice cirrata (Sars)? Our specimens differ from the European representatives as de- scribed by Séderstrém (1920) by having two small teeth rather than one, side by side above the main fangs of the ventral hooks (no exception in the four parapodia of two specimens studied). This is one of the characters by which Séderstrém distinguished his L. bahusiensis from L. cirrata (later authors have not separated these forms). According to Séderstrém, genital pouches begin with the for- mer species at the fifteenth to seventeenth setiger, with the latter at the twenty-eighth to thirty-fifth setiger. Hartmann-Schréder (1965) has reported Z. cirrata from Chile, with similar hooks as in our form. The Chilean form is clearly different from ours, as well as from Séderstrém’s form, because of its short dorsal sense organ, although the beginning of brood pouches is not known. NO. 3667 POLYCHAETES—BANSE AND HOBSON 290 In our material, the occipital tentacle inserts at the level of the first parapodium. The dorsal sense organ reaches the twenty-sixth to thirty-fourth setiger (average 29, S.D. 3.7, n=6). The first occurrence of genital pouches ranges from the twelfth to twenty-eighth setiger (average 20, S.D. 4.5, n=13). In the largest animals, which are mature females, neuropodial hooks start at the forty-eight and sixty- second setigers; hooks occur in more anterior segments (thirtieth to thirty-fifth) in younger animals. In view of this variability we include our material, for the time being, in L. cirrata in spite of differences in the shape of the hooks. BEgg-carrying females are dark reddish-brown. The largest anterior fragment, with 62 setigers, is 5.5 cm long and 3.5 mm wide, without parapodia. The diameter of eggs taken from the coelom is slightly larger than 200u, similar to Swedish material of both forms. A juvenile animal had been regenerating the prostomium and five anterior setigers. The U.S. National Museum has an uncataloged adult specimen, also regenerating the anterior end. Found at stations 1-4 and 6-8. Laonice japonica (Moore) Sptonides japonicus Moore, 1907, p. 204. Laonice cirrata—Imajima and Hartman, 1964, p. 281 partim. Not Laonice japonica.—Monro, 1933, p. 1047. We consider L. japonica to be a species separate from L. cirrata (Sars), as suggested by Monro (1933), because of the beginning of genital pouches at the fourth to fifth steiger. Also, the dorsal sense organ reaches only to the eighteenth steiger in the holotype of L. japonicus (Acad. Nat. Sci., Philadelphia, type no. 1055) rather than to the twenty-eighth to thirtieth setiger as in L. cirrata (Séderstrém, 1920). The gills on the second setiger are half as long, and on the third setiger about as long as the notopodial postsetal lamellae. The holo- type of L. zaponicus has been collected off Japan at Albatross station 3771, not 5771 as stated by Moore (1907). Monro (1933) has observed gills from the third setiger on a frag- ment from the Gulf of Panama identified as L. japonica; the dorsal sense organ extended to the twelfth setiger. Therefore, his form seems to be a different species. Laonice pugettensis, new species FIGURE 6a Laonice cirrata.—Berkeley and Berkeley, 1936, p. 27 partim. Laonice sp. I Banse et al. [in press]. Typrs.—Holotype: USNM 36263. Paratypes: USNM 36262 (4). Both from station 5, 47°10’48’” N, 122°50’00’’ W (February 1963). 26 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 There are many anterior fragments from stations 1-3, 5, 6, and 8: mature females with up to 70 setigers about 3 cm long and almost 2.5 mm wide without parapodia, some posterior fragments that pre- sumably belong to this species, and several anterior fragments of small individuals about 0.5 cm long and 0.5 mm wide. Description.—The prostomium is broader than long and anteriorly flattened. There are two large crescentic eyespots and a distinct occipital antenna. The dorsal sense organ (caruncle) reaches to about the twenty-fifth setiger. The palps are lost in all specimens. Gills start at the second setiger and number 24 pairs in a well-preserved young animal, and 27, 32, and 33 pairs in mature females. The first pair is not quite as long as the notopodial lamella; the next two or three pairs, as well as the last one or two pairs, are shorter than the rest. A fully developed gill is shown in figure 6a. Notopodial postsetal lamellae in the middle of the gill-bearing region are auricular ventrally. Their distal ends are fairly blunt in adults (fig. 6a), whereas in younger specimens the tip is tapering in the long axis of the lamella so that the lamella appears to point upward. Posterior to the gill-bearing region a ridge from the notopodial lamella on to the dorsum occupies about one-fourth the distance between the cirri on each side in the holotype but is not distinct in the other large specimens. Genital pouches occur from the second to seventh setiger onward through at least the seventieth setiger. Neuropodial hooks are found in an old and a young specimen from the thirtieth to thirty-third setiger onward. There are two secondary hooks side by side above the main fang. Dorsal hooks are absent on the anterior ends, as well as on the posterior end checked. Stout ventral neurosetae begin on the twenty-sixth and thirty-third setiger with two old individuals, at the twenty-fourth with the young animal mentioned above, and farther forward with another young specimen. The pygidium has nine slender cirri that are slightly longer than the pygidium is wide; possibly their number varies. Polygonal eggs from genital pouches are about 125y in diameter. The animals are yellowish pink. The name refers to the type-locality. Diaenosis.—A small Laonice species with genital pouches from the second to seventh setiger. Dorsal sense organ (caruncle) to about the twenty-fifth setiger. Approximately 30 pairs of gills. Dorsal ridges posterior to the gill-bearing region inconspicuous. Dorsal hooks absent. Ventral hooks with two secondary teeth side by side above the main fangs. Discussion.—We follow Séderstrém (1920) in using the occurrence of genital pouches for separating species of the genus. Sdderstrém has found a fairly close relation between the end of the atokous region NO. 3667 POLYCHAETES—BANSE AND HOBSON 2¢ and the beginning of the brood pouches in three spionid genera. We note, however, that eggs can be seen through the body wall of three spec- imens of L. pugettensis only at the twenty-fifth setiger or slightly post- erior to this segment although genital pouches start at the second to seventh setiger. Omitting Monro’s form, the known Laonice species with brood pouches starting before the tenth setiger may be distinguished as follows: 1. Dorsal sense organ (caruncle) to about the 25th setiger; dorsal ridges behind the gill-bearing region inconspicuous. . .. . L. pugettensis new species Dorsal sense organ not reaching the 20th setiger; dorsal ridges present or SERIE TAD Moat merce no kict bh aharcenla Sgt jo Mistyeretet Walia wics). wal top caked ue te inden iey Serene 2 2. Neuropodial hooks occurring before the 20th setiger. L. appeloefi Séderstré6m Neuropodial hooks starting posterior to the 20th setiger . ....... 3 3. Gills distinct from second setiger onward; conspicuous dorsal ridges behind Gill pearTing FEAIOM Ar Ss sis 3 see as) ef ve L. japonica (Moore) Gills distinct only posterior to second setiger; dorsal ridges inconspicuous. L. antarcticae Hartman Laonice pugettensis is different from the incompletely known Arici- deopsis megalops Johnson (1901) from Puget Sound, considered to be a species of Laonice (Hartman, 1959), because of hooks on the seven- teenth somite in the latter. Also, neither the occipital antenna nor the gills on the tenth setiger of old or young specimens of the new species are as large as figured by Johnson. A few records of L. cirrata from waters of British Columbia must refer also to LZ. pugettensis as evident from the description of the former species by Berkeley and Berkeley (1952). Further, an un- numbered sample in the U.S. National Museum, collected in Depar- ture Bay, B.C., between 1918 and 1925, contains 15 L. cirrata and 2 L. pugettensis (cf. Berkeley and Berkeley, 1936). Paraspio cirrifera, new species Fiaures 6b-f Paraspio sp. I Banse et al. [in press]. Typrs.—Holotype: USNM 36270. Paratypes: USNM 36271 (2) and 36272 (3). All come from station 3, 47°44’31’’ N, 122°31’53’" W (February 1963). Twenty-four specimens from stations 1, 3, 4, 6, and 8 are available. About half of our specimens come from station 3. The largest anterior fragment of a mature female has 73 setigers and a length of 21 mm. One complete animal possesses about 85 setigers. Description.—The prostomium (fig. 66) is rounded anteriorly. In most specimens there is a high bilobed protuberance on the upper side; in such a case, the eyes are not visible when the animal is seen 28 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 125 from the front (fig 6c). The posterior end of the prostomium is obtuse. Palps have been lost. There is a very small gill on the first setiger. In a well-developed parapodium (fig. 6d), the notopodial and neuro- podial postsetal lamellae are present as usual. Also, there is a fairly large neuropodial presetal lamella and an interramal papilla. The gills are partially fused with the notopodial lamellae. Near the pos- Ficure 6.—Laonice pugettensis, new species: a, median parapodium (setae omitted). Paraspio cirrifera, new species: b, dorsal view of anterior end; c, frontal view of anterior end (setae of the first setiger are shown on the animal’s right side, setae of the second setiger on the left side); d, anterior view of parapodium of 10th setiger (light broken lines indicate setae); ¢, hook from 25th parapodium; f, anal cirrus. Trochochaeta multi- setosa: g, anterior view of third setiger (light broken lines indicate notosetae). terior end of the animals, the lamellae are reduced, and the gills are long and straplike. In the tenth parapodium, a dorsal extension of the gill reaches forward, toward the origin of a most characteristic notopodial presetal cirrus, which is fairly round in cross-section. This cirrus is present at least through the twenty-fifth setiger but visible only in well-preserved material without dissection. There are notopodial and neuropodial setae in the first setiger. The number of neurosetae is about 20. Apart from the wings, the setae appear dotted rather than striated. Tridentate neuropodial hooks NO. 3667 POLYCHAETES—BANSE AND HOBSON 29 (fig. 6e) begin on the sixteenth to seventeenth setiger in young and mature specimens. There are fewer than 10 hooks per parapodium. Two ventral stout bristles start from the thirty-second or thirty-third setigers. Notopodial hooks are absent. There are four anal cirri (fig. 6f), round in cross-section and about 0.4 mm long. Two large brown pigment spots occur on the prostomium behind its anterior constriction; sometimes similar pigmentation is present also on the anterior end of the lateral wings of the prostomium. There is some irregular pigment pattern dorsally behind the prostomium, and a thick band of pigment is found laterally on the peristomium. In the first 8 to 10 setigers, some brown pigment occurs in the inter- segmental borders between the neuropodia and ventrally on both sides of the midline, separated by the pigment-free midline. Polygonal eggs of about 250u diameter are observed in one specimen (February). The animals construct a transparent fragile mucus tube. The name refers to the presetal notopodial cirrus. Diaenosis.—A Paraspio species with an anteriorly rounded prostomium. Notosetae and a small gill on the first setiger. Dorsal presetal cirri in the anterior region of the body. Tridentate neuro- podial hooks from setiger 16 or 17, ventral neuropodial acicular setae from setiger 32 or 33. Four elongated anal cirri. DiFrFERENTIAL DIAGNOSIS.—The new species may be distinguished from the members of the genus listed by Hartman (1959, 1965b; including Microspio) as follows: P. africana (Rullier), P. armata (Thulin), P. minuta Hartmann-Schréder, P. kussakini (Chlebovich), P. theeli (Séderstrém), and P. wireni (Augener) do not have tri- dentate hooks. Among the Paraspio species with tridentate hooks, P. mecznikourana (Claparéde) and P. rolasiana (Augener) lack setae in the first notopodium (as does P. minuta); P. arctica (Séder- stroém) has a pointed prostomium (as does P. theeli). The presetal dorsal cirrus has been reported only for P. africana and P. atlantica (Langerhans); the latter species has been separated from P. arctica by Hannerz (1956). Paraspio africana has unidentate hooks and peculiar liimbate notosetae; P. atlantica has a rather pointed pros- tomium, and its tridentate hooks start on the ninth segment. Prionospio pinnata Ehlers In our material, the third pair of gills is always only half as long as the first two pairs. There is no appendix on the third pair of gills as reported by Caullery (1915) for Malayan material. Ehlers (1901) has implied that the species is a selective deposit feeder. Specimens from station 3 (mean grain size diameter 0.126 mm) have numerous sand grains not exceeding 0.07 mm imbedded in fine detrital material, which confirms Ehlers’ observation. 30 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 125 Family DisoMIDAE Trochochaeta multisetosa (Oersted) FIGuRE 69 Disoma multisetosum.—Hartman, 1959, p. 394. Trochochaeta multisetosa.—Pettibone, 1963, p. 312. There is one incomplete specimen from station 2. The dorsal postsetal lip of the third setiger is strongly digitate (fig. 6g); on the undissected animal, the uppermost lobe of the lip protrudes like a cirrus, which is not so on the preceding and following parapodia. There is no acicula in the notopodium. The shape of the notopodial lamella on the third setiger is very close to that drawn by Friedrich (19388, fig. 88b) from material close to the type-locality. We therefore assign the specimen to 7. multisetosa rather than to T. franciscanus (Hartman). A detailed comparison of better preserved material from Puget Sound with the description of 7. franciscanus by Hartman (1947b) would be desirable. Trochochaeta franciscanus is considered a synonym of 7. multisetosa by Pettibone (1963). New for Puget Sound. Previously known in the Pacific from Sakhalin. Family CirRATULIDAE Caulleriella alata (Southern)? Because of the differences in the insertion of the palps and in the setation, we have some doubt that our animals are identical with the European C. alata (Southern), but we do not wish to decide on this before having studied material from other localities on the west coast of the Americas. As in C. viridis pacifica (Berkeley), the palps insert on the first setiger rather than on the preceding segment. Caulleriella viridis pacifica is considered a synonym of C. alata (Berkeley and Berkeley, 1950). In the first neuropods of our animals are 8 to 10 winged, bifid hooks and 1 to 2 capillary setae in small and large specimens (about 2-3 mm long, with 0.4-0.5 mm greatest width, and 1-1.5 mm long and 0.6-0.8 mm greatest width, respectively). Capillary setae can be absent in posterior neuropodia. Notopodial hooks are found from the forty-fifth to fifty-fifth setigers on, rather than from about the twentieth as in the European representatives (Southern, 1914). Notopodial hooks from the twentieth setiger have also been reported for C. alata from southern California (Hartman, 1961). Hartmann-Schréder (1962) has recorded hooks starting from the sixth and the seventeenth notopodia in Chilean material. Our form 1s certainly different from C. alata maculata (Annenkova). NO. 3667 POLYCHAETES—BANSE AND HOBSON ol Caulleriella annulosa (Hartman), new combination FiGurE 7a Tharyx annulosus Hartman, 1965a, p. 167; 1965b, p. 59. Among Tharyz species kindly sent by Dr. O. Hartman, there are specimens of T. annulosus from station SL 4, off New England (Hart- man, 1965a). There are acicular spines (fig. 7a) in neuropodia of the middle section of the body, and in both rami of the posterior section, in addition to capillary setae. These spines were not mentioned in the original description. We transfer the species therefore into Caulleriella as defined by Hartman (1961). Chaetozone Malmgren Because we have seen six North Pacific species of Chaetozone, a key for the temperate and subarctic North Pacific species is given. It considers primarily characteristics of the front end. Chaetozone abranchiata Hansen is not included in the genus on the basis of the original description (from Hansen, 1882; Levinsen, 1883) and that by Moore (1903), which report only capillary setae but not acicular spines. There are two North Pacific records that are not included in the key: Ushakov (1950) briefly described a Heterocirrus species with unidentate acicular spines starting in the fifteenth neuropod, which well might have been a Chaetozone species. Hartman (1963) mentioned a Chactozone species from California with neuropodial spines present from the fifteenth segment. 1. Neuropodial spines from Ist setiger; spines posteriorly arranged in cinctures. C. corona Berkeley and Berkeley Neuropodial spines absent on Ist setiger but starting anterior to 15th to to 20th setiger .... wee 2 Neuropodial spines Sart ‘onttor iB ‘15th ie 20th ie Savion to 35th setiger; spines posteriorly not arranged in cinctures. ........ 83 Neuropodial spines starting posterior to about 35th setiger. . ..... 4 2. Neuropodial spines from 7th to 10th setiger; spines posteriorly not arranged in cinctures. Serrated capillary setae after the 7th to 10th neuropodium. C. berkeleyerum, new species Neuropodial spines from about the 10th setiger; long buccal region. Smooth capillary setae only. Posteriorly, 2 to 3 spines per ramus tending to form Cinetures\. nace % ..... .C. multioculata Hartman 3. Neuropodial spines, fairly need ont about the 17th setiger. Posteriorly, single spines without capillary setae in each ramus. C. armata Hartman Neuropodial spines from about 26th (18th to 40th) setiger. Short capillary setae in the middle region of body that appear to be broken off obliquely. C. acuta, new species Neuropodial spines from about 30th setiger. Long, ordinary, almost smooth capillary, setaeyi jad, tsuki eiedes) ss ais, a0ues |. . Cs, gracilis: Moore 32 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 4. Neuropodial spines, serrated at the tip, from the 35th to 40th setiger, posteri- orly not arranged in cinctures. Capillary setae ordinary. C. gracilis sensu Hartman (1961) (see below) Neuropodial spines from the 45th to 90th setiger, posteriorly arranged in very marked cinctures. Capillary setae ordinary. . .C. setosa Malmgren Neuropodial spines from about 70th neuropod, tending to form cinctures posteriorly. Long, threadlike capillary setae from about the 13th setiger. C. spinosa Moore REEL Ujja/77**“ “thy thE ig Ficure 7.—Caulleriella annulosa: a, acicular spine. Chaetozone acuta, new species (seta from median foot): b, capillary seta; c, acicular spine. Chaetozone berkeleyorum, new species (setae from 40th neuropodium): d and g, capillary setae; f, acicular spine. Chaeto- zone gracilis (holotype, setae from about 85th setiger): g, capillary seta; 4, acicular spine. Tharyx secundus, new species: i, detail of anterior capillary seta; j, neuroseta with short tip from 43rd setiger. Tharyx serratisetis, new species: k, detail of notoseta from 10th parapodium; /, serrated neuroseta from median region of body. Chaetozone acuta, new species FIGURES 7), c Chaetozone sp. I Banse et al. [in press]. Typrs.—Holotype: USNM 36275, from station 5, 47°10’48”” N, 122°50’00’’ W (May 1963). Paratypes: USNM 36276 (2) and 36277 (2), from the same station (February and May 1968). Many complete immature specimens of up to 155 setigers, 15 mm long and 1.1 mm wide, are at hand from stations 3, 5, 6, and 8, but no complete, mature individuals are available. Description.—A cirratulid species with a conical prostomium on which one pair of eyespots is visible in freshly preserved material. There is a 2-ringed achaetous region that is about as long as it is NO. 3667 POLYCHAETES—-BANSE AND HOBSON 33 wide. Palps and the first pair of gills insert prior to the first setiger. Gills, which can be 5-7 mm long anteriorly, occur throughout the body but are mostly lost. The anterior setigers (at least 50) are crowded; posteriorly, the length of the segments becomes almost one- half their width. The setae of the posterior region originate from slight ridges which, however, are not nearly so pronounced as in C. setosa Malmeren. The anus is dorsal, with a crenulated margin dorsal and a ventral semicircular lip. Anteriorly, there are about one dozen long capillary setae in each ramus, which are only slightly limbate. When bent, their fibers tend to separate with the result that the former cutting edges of the setae appear to be feathered. In the notopodia, prior to the occurrence of acicular spines, these setae gradually are replaced by shorter setae, which resemble obliquely broken-off bristles and have a finely drawn tip (fig. 7b). In the middle region of the body they are the only cap- illary setae present in noto- and neuropodia (two to four per ramus). In young animals near the end of the body, they are replaced by or- dinary capillary setae. Acicular spines are slightly curved and have rounded tips and slight wings (fig. 7c). Ventrally, they occur from the eighteenth to thirty-eighth setigers onward (average 26; n=11). Dorsally, they begin 30-40 setigers posterior to the ventral spines. There are usually five to seven neuropodial, and two to five notopodial spines. Posteriorly, notopodial and neuropodial rami are only slightly separated laterally, but the spines do not nearly encircle the body. A posterior end of 51 setigers (10 mm long, 1.2 mm wide), apparently of a maturing female, and collected in May, carries in the last setigers dorsally four to five, ventrally five to six spines, with almost the same number of the described capillary setae (fig. 7b) between them. The setae are restricted to the sides of the body. There are numerous polygonal eggs of 50u-60u freely floating in the body cavity, including the last setigers. The species seems to be a selective deposit feeder. The largest particles in the intestines of two specimens from station 5 (mean particle size 0.43 mm) are 0.05 mm in diameter; most are below 0.02 mm in diameter. The name refers to the capillary setae of the middle region of the body. Diaenosis.—A Chaetozone species with ventral and dorsal acicular spines starting ventrally from approximately the twenty-sixth (range, eighteenth to fortieth) setiger, with slight wings. Short capillary setae, which appear to be obliquely broken-off, in the middle region of the body. DIFFERENTIAL DIAGNOSIS.—For the characterization of the new species, it seems questionable how much significance is to be attached 313-605—68——3, 34 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 to the wings of the acicular spines since they occur also in the local representatives of C. setosa Malmgren; this feature seems not to have been reported previously for C. setosa. Among the Chaetozone species listed by Hartman (1959, 1965b) to which C. berkeleyorum, new species, and C. curvata Hartmann-Schréder are to be added, the new species seems to be unique by the character of the capillary setae in the middle part of the body. For the characters distinguishing C. acuta from the other Pacific species, see page 31. Chaetozone berkeleyorum, new species Ficures 7d-f Caulleriella viridis pacifica.—Berkeley and Berkeley, 1942, p. 197.—Not Berkeley, 1929, p. 307 [fide Berkeley and Berkeley, 1950, p. 57]. Caulleriella gracilis—Berkeley and Berkeley, 1950, p. 57; 1952, p. 37 partim. [Not Chaetozone gracilis (Moore); see p. 35 of this paper]. Chaetozone sp. II Banse et al. [in press]. Typrs.—Holotype: USNM 35286. Paratypes: USNM 36251 (2). All from the littoral of Skidegate Narrows, Queen Charlotte Islands. (Collected in 1935.) There are two complete specimens and an anterior fragment from the littoral of Skidegate Narrows in the Queen Charlotte Islands, which are in the Berkeley Collection of the U.S. National Museum and which have been reported by Berkeley and Berkeley (1950), and one badly damaged anterior fragment from our station 7. The holotype is the largest specimen, has 72 setigers, and is 12 mm long, with 1 mm greatest width. Description.—The prostomium is triangular but unusually blunt by being slightly broader than long. Eyes are not visible. The palps seem to insert on the anterior margin of the first setiger. In the vial from Skidegate Narrows, detached grooved palps of 4-5 mm length are present. In the Canadian animals, the segments lengthen in the general! region of the fortieth setiger. In the Puget Sound specimen, only the approximate 10 first setigers are fairly crowded; thereafter they are about half as long as they are wide. There are very short gills above some of the notopodia, at least to about the sixtieth setiger. Even in posterior setigers, the segments are fairly cylindrical although with clear demarcations, in contrast to those of C. setosa Malmgren. The anus appears to be displaced dorsally. In the anterior 8 to 10 setigers, there are less than a dozen each of dorsal and ventral capillary setae, some of them one-third the body width. Subsequently, the notosetae shorten. From the seventh to tenth neuropodium, one to three capillary setae appear that are slightly widened in their middle portion (fig. 7d). Two-thirds of the widened portion is finely serrated when seen under 400. Scales NO. 3667 POLYCHAETES—BANSE AND HOBSON 3a (rather than teeth) are well visible under 1000 (fig. 7¢). These capillary setae occur dorsally at least from the twenty-fifth setiger together with smooth ones. Starting from the seventh to tenth neuro- podia, the widened capillary setae are accompanied by two to three smooth acicular yellow spines without wings (fig. 7f). About two to three notopodial acicular hyaline spines appear at about the forty- fifth setiger (not the twenty-fifth as stated by Berkeley and Berkeley, 1950). They are straighter than the neuropodial ones, and only as thick as the accompanying capillary setae. The spines do not encircle the posterior portion of the body. The serration of the notopodial capillary setae is weak. The specimens from Skidegate Narrows had originally been mis- identified as Caulleriella viridis pacifica, according to Berkeley and Berkeley (1950). The authors state in the same paper that C. viridis pacifica as described by Berkeley (1929) is a synonym of C. alata (Southern). The specimens on which the 1929 paper has been based are no longer available (Pettibone, 1967). The new species is named for Edith and Cyril Berkeley. Diacenosis.—A Chaetozone species with blunt prostomium and fairly smooth posterior region. Smooth neuropodial spines without wings and serrated capillary setae from the seventh to tenth setiger. Thin notopodial spines from the forty-fifth setiger. Some notopodial serrated capillary setae. DIFFERENTIAL DIAGNosIs.—Among the Chaetozone species listed by Hartman (1959; 1965b) to which C. acuta, new species, and C. curvata Hartmann-Schrider are to be added, C. abranchiata Hansen, C. atlantica McIntosh, and C. pacifica McIntosh have only capillary setae, accord- ing to the original descriptions (for C. abranchiata, see p. 31). Among the others, neuropodial spines occur before the twentieth setiger in C. andersensis (Augener) (see Hartman, 1967; spines from ninth neuropodium; posteriorly, one to two per ramus); C. armata Hartman; C. carpentert McIntosh (spines from about the tenth notopodium and neuropodium, with some very large spines among them); C. multi- oculata Hartman; and C. gayheadia Hartman (nearly straight spines from the first neuropodium; apparently no posterior cinctures). Spines can occur in C. acuta from the eighteenth neuropodium. None of the above species has serrated capillary setae. The distin- guishing features for the North Pacific species are given on p. 31. Chaetozone gracilis (Moore) Figures 7g, h Tharyz gracilis Moore, 19238, p. 187, partim. Not Caulleriella gracilis—Berkeley and Berkeley, 1950, p. 57; 1952, p. 37 [see p. 34 of this paper]— Hartman, 1960, p. 125. Not Chaetozone gracilis—Hartman, 1961, p. 111. 36 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 We include here a study of Moore’s original material made when describing Chaetozone berkeleyorum. In the holotype of C. gracilis (USNM 17398) neuropodial hooks start at about the thirtieth setiger. They are smooth and have rounded tips (fig. 72; also checked near the fiftieth setiger). The acicular spines are fairly inconspicuously arranged and do not encircle the posterior part of the body. The accompanying capillary setae (fig. 7g), which are quite similar to those in the notopodia, seem to be serrated, but this is irregular and might rather be a sign of decay of the setae. The oval eggs are up to 120u by 95u. Moore’s description fits the holotype only. The cotype (Acad. Nat. Sci., Philadelphia, no. 3097) had been dried up at some time but is clearly another species. Notopodial capillary setae, slightly longer than the body is wide, occur throughout the body. There are only capillary setae on the twenty-eighth to thirtieth setigers. The neuropodia of posterior setigers (past the hundredth) have only broad capillary setae with serration of the edges well visible under 400, which are similar to those of Tharyx secundus, new species (p. 37). All setae are hyaline. The cotype thus seems to be a Tharyz species. Dr. O. Hartman (in litt.) suggested already that the holotype and the cotype of Moore’s material may be different forms. Neither species is identical with Chaetozone gracilis of Hartman (1961). This form is characterized by serrated neuropodial spines and is a species to be named. Chaetozone setosa Malmgren In our material, neuropodial spines occur from the forty-fifth to sixty-fifth setigers onward, and even from about the thirty-fifth to fortieth in specimens under 1 cm length, whereas Hartman (1961) reported for California animals a beginning at the seventy-ninth to nintieth setigers. The spines are faintly winged, similar to those of C. acuta described above although they are more pointed than in the latter species. The species occurs at all stations, but, regardless of the sediment composition, the intestine is filled with masses of fine detritus, among which are some sand grains or diatom frustules that do not surpass 0.05-0.06 mm in diameter. The species is thus a selective deposit feeder, at least in coarse sediment. Chaetozone spinosa Moore Chaetozone spinosa.—Imajima and Hartman, 1964, p. 297. There is one anterior fragment with 36 setigers, which is about 4 mm long. The first 12 setigers are crowded; the body attains its great- No. 3667 POLYCHAETES—BANSE AND HOBSON 37 est width of 0.5 mm here. Thereafter the setigers become longer, but the width is only 0.8 mm. Rudiments of palps and gills are visible. Whereas the neuropodial bristles in the crowded section of the body are all about normal length, there are some threadlike setae in the notopodia. From about the thirteenth setiger these occur in notopodia and neuropodia, with a length of 0.5-0.6 mm. At the beginning of the exposed part, their width is about 4y, farther out only 3y; the thickness is about 1y here. These setae are “‘striated having a slight turn” as described by Moore (1903, p. 470); however, the scales ob- served on the large specimen from Japan could not be seen with 1000 magnification in our material. Acicular spines occur in the species from the seventieth neuropod onward (Hartman, 1960). Our identification is based on the threadlike setae. Found at station 6 in May. New for Puget Sound. Previously recorded from Japan and California. Tharyx multifilis Moore Contrary to the original description by Moore (1909b), the anus of one complete specimen from Puget Sound is displaced dorsally by a very pronounced triangular ventral lip. This holds also for an in- dividual from Velero IV station 5027-57 off Pt. Fermin, kindly sent by Dr. O. Hartman. Thus the separation from this species of T. pacifica Annenkova continues to be doubtful as also pointed out by Chlebovich (1961); 7. pacifica has eyes, T. multiilis does not. The caudal ends of our two specimens are very slightly thickened (inflated), whereas Moore (1909b) and Hartman (1961) reported a slender taper- ing posterior end. A large portion of the intestinal contents of two specimens from two hauls at the fine-sand station 4 (mean diameter 0.10 mm) is made up of grains of 0.07-0.10 mm diameter, with a very few larger (0.15 mm) particles present. Thus there seems to be only slight selection of particles when feeding. Tharyx secundus, new species FIGURES 72,7 Tharyx sp. II Banse et al. [in press]. Typrs.—Holotype: USNM 36278. Paratype: USNM 36279 (1) drawings from paratype). All from station 3, 47°44’31’’ N, 122°31’53’” W (February 1963). There are several anterior and posterior fragments 1-2 cm long and 0.38-0.5 mm wide. In all, there are at least 100 setigers. Description.—A pointed prostomium without eyes is followed by a 3-ringed achaetous region. The scars left by the broken-off palps at 38 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 the anterior margin of the first setiger are separated from each other by a gap that is as wide as their diameter. The gills of this segment arise between the palps and the notopod; on the following segments, they insert just above the notopod. Gills are at least 2 mm long and are found at least to the seventieth setiger (length of the largest anterior fragment, apparently a juvenile). Approximately the first 20 setigers are widened laterally, but the dorsum is low and not inflated. The external borders of the segments are dorsally well visible. In an anterior fragment of 65 setigers, 13 mm long, this anterior region is 2 mm long (of which the achaetous rings contribute 0.4mm) and 0.5mm wide. The anterior region is followed by more than 40 setigers, which are about as wide as they are long and are separated from each other by deep furrows. Sometimes they appear beadlike. Near the fortieth setiger, segments are 0.3 mm long and 0.3 mm wide. The parapodia arise on the hindmost portion of the segments. The posterior region consists of about 30 setigers, which are crowded and laterally widened (inflated). A posterior portion is 1.5 mm long. The anus is dorsal. In the anterior region, all setae are capillary. Notosetae are half as long as the body is wide. By the twentieth to twenty-fifth setiger, the setae of both rami become limbate and weakly serrated (fig. 77). The neurosetae shorten in the subsequent parapodia, become very broad, and are beset with distinct hairs on the cutting edge (fig. 77). The serration is just visible under 400, and thus the setae are not as strongly serrated as in 7. serratisetis described below (under oil im- mersion, the neurosetae of local specimens of 7. multifilis Moore are not quite smooth either but do not appear to be hairy). In the forty- fifth setiger of an apparently mature male, there are about 20 of these setae in each ramus. In the posterior region, neurosetae are narrow and limbate again, similar to figure 77. The tube is apparently ephem- eral, being made of soft and fairly transparent mucoid material. The name refers to the provisional name used by us prior to the description. Driaenosis.—A Tharyz species with beadlike setigers in the middle section of body. Posterior end inflated. Anus dorsal. Very broad neurosetae with distinct hairs on cutting edge in the middle section of body. DIFFERENTIAL DIAGNOSIS.—Among the Tharyx species in Hartman (1959, 1965b) but omitting 7. annulosus (see p. 31), and adding Tharyz sp. described by Berkeley and Berkeley (1941, from California) and YT. serratisetis, new species, the following have serrated (or ap- parently so) neurosetae: 7. dorsobranchialis (Kirkegaard), T. ser- ratisetis, new species, 7. tesselata Hartman, and the mentioned Tharyx sp. Among these, 7. tesselata has an inflated posterior end like 7. NO. 3667 POLYCHAETES—BANSE AND HOBSON 39 secundus, but the middle segments are short, whereas there is a bead- like middle section in the new species. Tharyx serratisetis, new species Fiaurgs 7k, l Tharyx sp. I Banse et al. [in press]. Types.—Holotype: USNM 36266 (possibly the posterior fragment in the vial is of a second individual), from station 3, 47°44’31’’ N, 122°31’53’’ W (February 1963). Paratypes: USNM 36267 (2) and 36268 (1), from the same station (February and April 1963). There are numerous fragments, in part of mature animals, from several stations, with up to about 125 setigers, 2 cm long and 2 mm greatest width. The total length is more likely to be 4 cm, and the number of setigers may well reach 200. Most specimens come from station 3, fewer come from stations 6 and 8, some were found at station 7 (in 2 of 35 hauls from the station), where an egg-bearing female occurred. Description.—The prostomium is conical and can be slightly longer than wide. Eyes are not visible. The achaetous region appears to consist of three indistinct rings. Palps are lost. Gills are rare on the preserved material and not longer than the body is wide; they arise just above the notopod. The anterior setigers are very crowded, and external borders of segments are not well visible. The dorsum is high. Starting from the fifth to tenth setiger, the ventrum is glandular through the sixtieth to seventieth setiger in mature specimens, and to the forty-fifth in one young, complete animal 2 cm long and 0.5 mm at greatest width. In moderately contracted animals, segments be- come about three times as long as in the anterior region from about the fiftieth to seventy-fifth setiger. From about the one hundred twenty-fifth setiger, segments are about five times as long as the anterior ones. The middle region of the body is not beadlike. The body tapers gradually to the pygidium, which carries a semicircular ventral lip with the result that the anus is dorsal. Notosetae are long thin capillaries that are one-half to three-fourths as long as the body is wide; neurosetae are one-third to one-half as long as the notosetae of this region. All these setae are slightly limbate and have cutting edges with fine hairs (fig. 7) that are just visible under 400 magnification. The hairs appear to be outcroppings of the fibers that make up the setae. After the seventy-fifth to one-hundredth setigers, these thin neurosetae are replaced by flattened serrated setae, measuring about 75u from the beginning of the serrated region to the tip (fig. 71), and by about a dozen that are twice as long owing to a 40 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 long drawn-out tip. When the preparation of a parapodium is squeezed under the cover glass, the long and short setae seem to alternate. Tubes are not known. Exceedingly numerous polygonal eggs, about 150u by 110y, are observed in a female in May. The intestinal content of two specimens from station 3 (mean particle size 0.126 mm) con- tains many sand grains 0.2 mm in diameter and even an unbroken Coscinodiscus species test 0.25 mm in diameter, in addition to smaller grains and much fine detritus. Thus, the species may be a not very selective deposit feeder. The name refers to the serrated neurosetae. Diaenosis.—A Tharyx species with serrated neurosetae after the seventy-fifth to one-hundredth setigers. Median segments not bead- like. Posterior end not inflated. Anus dorsal. DirFERENT Dragnosis.—Among the Tharyx species in Hartman (1959, 1965b), but omitting 7. annulosus Hartman (p. 31) and add- ing T. secundus, new species, and Tharyzx sp. described by Berkeley and Berkeley (1941, from California), only 7. dorsobranchialis (Kirke- gaard), JT. tesselata Hartman, and the mentioned Tharyz sp. are known to have serrated setae. Of these, 7’. dorsobranchialis has a very peculiar arrangement of gills. Tharyx secundus has a beadlike middle body region and an inflated posterior end. Tharyx tesselata has an inflated posterior end and a characteristic tube. Also, the serrated broad neurosetae start very roughly at the fiftieth setiger (from material kindly sent by Dr. O. Hartman). The Berkeleys’ species has four achaetous rings following the prostomium, and the serrated setae seem to start at a low-numbered setiger. Thus, the new form is easily distinguished from the species with serrated setae. The form seems also to be different from all those species that are known to have flattened posterior neurosetae for which serration has not been described. We note, however, that there is a very minute serration on the flattened neurcsetae of T. multifilis Moore from our samples; in fact, we cannot distinguish anterior fragments of the new species from those of T. multifilis. Family FLABELLIGERIDAE Brada sachalina Annenkova? Brada sachalina.— Ushakov, 1955, p. 310. There are about 10 specimens up to 3 cm long that are profusely covered with sand grains and usually have 23 setigers (22-25). The notosetae (three to five per bundle) are longer and thinner than the neurosetae (five per bundle). The dermal papillae are dome shaped with a short filiform tip; there are two (sometimes three) rows per segment. The nephridial papillae arise on the anterior portion of the NO. 3667 POLYCHAETES—BANSE AND HOBSON 41 fifth setiger but are barely noticeable. From Annenkova’s description (1922), B. sachalina is distinguished from B. ochotensis Annenkova mainly by the number of segments. Our specimens are intermediate in this respect. Both species have been described from the southern Sea of Okhotsk and occur also in the Bering Sea. Found only at station 2. Neither species has been found in the eastern Pacific before. Family OPHELIIDAE Travisia brevis Moore Our specimens, about 1.5 cm long, have 25 setigers followed by four achaetous segments, whereas Hartman (1961) stated that there are only two achaetous rings. Imajima (1963) has reported 24 setigers and four achaetous segments for specimens from the Okhotsk Sea, and Imajima (1964) has found 23 setigers in animals collected off Hokkaido. Family MALDANIDAE Isocirrus longiceps (Moore) Very numerous spherical (or slightly depressed) eggs about 250u in diameter were observed in November. Macroclymene? species Figures 8a-f Macroclymene? sp. I Banse et al. [in press]. Incomplete mature specimens with up to 27 setigers are about 6 em long and 1 mm thick. The longest animal is without the anterior three setigers and the rear end and must thus have had at least 30 setigers; probably it had at least 34 setigers. The cephalic plate (figs. 8a, 6) has a low rim that dorsally is almost absent; there is no dorsal incision. The palpode is obtuse. Nuchal erooves are straight and one-half to three-fifths as long as the cephalic plate. The proboscis has conspicuous papillae that are triangular in side view. There are two to three slightly bent acicular spines (fig. 8c) in each parapodium of the first three setigers. These setigers are glandularized over their entire length. There are two bundles of 14-18 limbate capillary setae each per ramus in the fifth and sixth setiger, with the fibrilles arranged as in figure 8d. The edges of the setae are not quite smooth (not shown in fig. 8d). There are six to seven short and long capillary setae each per ramus in the middle part of the body. The shorter ones are beset with hairs (fig. 8¢) up to the tip; the longer ones seem to carry hairs only in the proximal half. Thoracal unicni (14 per ramus in the fifth setiger) have one or two fewer secondary 313—605—68——4 42 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 teeth than the abdominal ones (fig. 8f; 10 per ramus), The pygidium is not known in spite of very many specimens obtained with a grab digging to about 15 cm depth. Figure 8.—Macroclymene? species: a, anterior end from above and slightly from the left side; b, left side view of anterior end; ¢, acicular spines from second setiger; d, detail of capillary seta from fifth setiger; ¢, detail of short capillary seta from middle portion of body; f, hook from middle portion of body. Pista fasciata: g, thoracic hook from third setiger; h, thoracic hook from 17th setiger. Scionella japonica, setae from median thoracic setiger: i, long capillary seta; 7, short capillary seta. Potamilla myriops: k, dorsal view of anterior segments; /, thoracic hook. The tube is made of a fairly firm, thin layer of very fine sand. Im- mature flattened eggs taken in August from an anterior end of 12 setigers of 35 mm length and 1.1 mm width measured up to 250u across, NO, 3667 POLYCHAETES—BANSE AND HOBSON 43 This is not one of the species known from this area and may well represent the genus Macroclymene, not yet recorded for the northern Pacific. Found at station 2. The drawings are of a mature specimen taken in August 1963. Praxillella affinis pacifica Berkeley Having many specimens, we add the following to the descriptions by Berkeley (1929) and Berkeley and Berkeley (1952): The dorsal rim of the cephalic plate can be split or more or less united. The pro- boscideal papillae are pointed as in P. gracilis (Sars) rather than smooth as in our specimens of P. affinis (Sars). The fourth setiger is the first fully glandular one as in P. gracilis, rather than the third setiger as in P. affinis. Feathered setae occur also in the posterior bundle of the twelfth setiger, rather than in the tenth or tenth and eleventh only. The constriction preceding the pygidium, emphasized by Berkeley (1929) for the subspecies, is much more marked in our material of P. gracilis than in the form discussed here. Numerous eggs of 240y-280u diameter are observed in animals collected in November. Young specimens occur together with adults in February. Rhodine bitorquata Moore Posterior ends of this species have apparently not been described yet. From one specimen we find that double rows of uncini occur through the fifteenth setiger and the first posterior collar occurs on the seventeenth setiger, as in R. lovent Malmgren. Arwidsson (1907) has stressed that these characters seem to be completely constant within species. The papillae of the proboscis of our specimen seem to have about the same form as in R. loveni, but details could not be made out. The margin of the posterior collar of many fragments is practically smooth with a slight dorsal notch; the hooks of our specimens also agree with the figure given by Arwidsson (1907, fig. 235 for R. loveni) so that the only gross difference of R. bitorquata from R. loveni continues to be the sharply bent nuchal groove of the former, as emphasized by Berkeley and Berkeley (1950). Individuals with moderately numerous, lens-shaped eggs of up to 140 largest diameter are observed in January. Young animals are found in February. Two individuals from station 4 (mean grain size 0.104 mm) have very coarse sediment (many particles of 0.15-0.20, up to 0.25 mm) and two specimens from station 7 (mean grain size 0.009 mm) have fine material (few grains reaching 0.05 mm) in their intestines. On the sediment studied, the species thus seems to be a nonselective deposit feeder. 44 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 Family PECTINARIIDAE Pectinaria (Cistenides) granulata (Linné) Pectinaria granulata.—Pettibone, 1954, p. 312. Pectinaria brevicoma.—Berkeley and Berkeley, 1952, p. 106. We follow Pettibone (1954) and include P. brevicoma Johnson de- scribed from Puget Sound in P. granulata because of the variability of some characters used for separation of the species. In our animals there are 10-11 pairs of brassy cephalic spines, about 20-40 papillae on the antennal membrane, and 5-7 scaphal hooks. The uncini have three to four major teeth. In small specimens, there can occur on one torus uncini with one row of major teeth, as typical for the subgenus Cistenides, and uncini with a double row, as in the subgenus Pectinaria. The large specimens all have a single series of major teeth in their uncini. Moore (1923) has stated that ‘this species unites Pectinaria and Cistenides.” Family AMPHARETIDAE Ampharete acutifrons (Grube) The tentacles of our specimens are smooth contrary to the generic diagnosis by Malmgren (1866) and Day (1964). Mature females are about 15 mm long and 1 mm thick but there are two broken specimens, otherwise very well preserved, that are at least 20 mm long and 2.5 mm thick, and apparently are not mature. Pettibone (1954) has stated a length of 55 mm and a width of 8 mm as the greatest size for the species. The species is a selective deposit feeder, as seen from the comparison of intestinal content with the sediment of the habitat. There are animals from all seasons. Many specimens collected in August and one found in May carry fairly numerous oval-lens-shaped egos, up to 240u x 200 x 50u in size. Ampharete gagarae Ushakov Ampharete arctica gagarae.— Ushakov, 1955, p. 369. In this form, which we consider to be a species of its own, the neuropodial lappets of the middle region of the abdomen are about one-fourth as long as the segments, whereas they are nearly as long as the segments in one specimen of A. arctica from Puget Sound. No anal cirri were found even in well-preserved animals. The species is a selective deposit feeder. Very few (50-100) irregular oval-lens-shaped eggs of about 220u x 1904 x 504-70» dimensions are observed in one specimen collected in January. NO. 3667 POLYCHAETES—BANSE AND HOBSON 45 Found at stations 1-4 and 6-8. New for the northeast Pacific. Previously known from the Sea of Okhotsk. Melinna elisabethae McIntosh Melinna elisabethae.— Ushakov, 1955, p. 363. There are four moderately well-preserved specimens. The gills form groups of one anterior and three posterior filaments. The thoracal setigers are arranged as in M. cristata (Sars) although the fourth bundle of the fine ventral capillary setae is visible only in transparent animals. The strong dorsal hooks have relatively short and broad bases, and the tips are bent backward by almost 180°, more so than pictured by Ushakov (1955, fig. 1834p). There is no canal in the tip. The uncini have four teeth. The species is a selective deposit feeder. Found at stations 1, 3, and 4. New for the northeastern Pacific. Previously known from the Sea of Okhotsk and the Arctic. Family TEREBELLIDAE Neoamphitrite edwardsi (Quatrefages), new combination Amphitrite edwardsii—Fauvel, 1923, p. 245.—Hartman, 1961, p. 497.—Imajima and Hartman, 1964, p. 335. There are three large specimens up to 13.5 cm long. Eyes appear to be absent. Seventeen setigers have notosetae; double rows of uncini occur from the eighth to the seventeenth setiger. Lateral folds are large on the two first segments bearing gills and are practically absent on the third one. Nephridial papillae are present on 9 consecutive segments, not on 12 as in WN. robusta (Johnson), starting on the second segment with gills. There are 11 or 12 ventral shields. We place the species into the genus Neoamphitrite because of the arborescent gills, but we have not studied the nephridia. Hessle (1917) has tentatively suggested this action. Found at station 2 and near station 7. New for Puget Sound. Previously known in the North Pacific from Japan. Pista fasciata (Grube) sensu Marenzeller FicuREs 8g, h Pista fasciata.—Imajima and Hartman, 1964, p. 343. Since there is some confusion about this species in the literature, the following may be observed: Our specimens, which are more than 10 cm long, agree well with the accounts of Marenzeller (1884) and Imajima and Hartman (1964). The lateral folds are shaped as de- scribed by the latter authors. Dorsally, the fourth segment (first setiger) is very much higher than the preceding ones. Nephridial 46 POLYCHAETES—BANSE AND HOBSON VOL. 125 pores are present on setigers 3 and 4 (segments 6 and 7), contrary to Indian material (Fauvel, 1932). Nearly rectangular ventral shields occur through the thirteenth setiger. From the first setiger onward a wide glandular band is present ventrally and laterally behind the tori (without a torus being on the first, it is in the corresponding place). In the anterior part of the thorax it is laterally as broad as the tori but is hardly visible ventrally between the ventral shields. In the middle region of the thorax, glands occupy laterally all the space between the tori and the posterior border of the segments. Between the tori and the shields, they occupy one-half to over three-fourths of the length of the setigers. Ventrally, on the last four thoracic setigers, only the anterior one-third to one-half of the segments is free of glands All thoracic uncini have narrow, long, thin shafts (figs. 8g, h). In side view, the number of teeth above the rostrum appears to be five to six in anterior segments, and four to five in posterior segments of the thorax. The protuberance (‘‘x”’ in fig. 8h) on the lower anterior curvature of the posterior thoracic hooks is often less pronounced than shown. Proclea graffi (Langerhans) Proclea grafi.— Ushakov, 1955, p. 395. There are three fairly well-preserved specimens from three hauls at station 3 in February. New for Puget Sound. Previously known in the North Pacific from the Bering Sea. Scionella japonica Moore FIGURES 82, 7 The gills of our two specimens from station 7 agree with the descrip- tion by Moore (1908) and are like those of S. winogradow (Ushakov, 1955, figs. 1483p, £), but not like those of S. japonica Berkeley and Berkeley (1952, fig. 176). Since the differences between these two species do not seem to be very clear, we give figures for the thoracal setae (figs. 8i, 7). Strong, long setae with very broad blades alternate with slightly more slender, shorter capillary bristles whose shafts are serrated above the short and narrow wings. Berkeley and Berkeley (1950) have pointed out that Moore figured only the larger setae. Family SABELLIDAE Potamilla (Pseudopotamilla) myriops (Marenzeller) Fiaures 8k, l Pseudopotamilla myriops.—Imajima and Hartman, 1964, p. 360. NO. 3667 POLYCHAETES—BANSE AND HOBSON 47 One anterior end of about 8 cm total length (about 2 cm for the tentacular crown) and 6 mm greatest width with nine thoracic and 30 abdominal setigers comes from station 7. Several other pieces in the sample appear to be from the same specimen, adding to about 21 cm total length and 195 setigers. The pygidium is missing. The species is particularly distinguished by the eyespots on the slightly more than 30 pairs of radioli. Ventrally there are about 10, dorsally up to 20, eyes per radiolus, the most distal ones leaving the last third of the rachis free. The basis of the most dorsal radiolus on each lophophore is widened on its dorsal side to give a sharp, straight edge. Ventrally, the lophophores are slightly rolled inwardly The collar (fig. 8k) is reduced dorsally; ventrally it is drawn out into two fairly long lips separated by a cleft. In a median thoracal setiger, there are 20 limbate capillary setae, about 55 spatulate bristles with fine tips, and almost 70 hooks and as many pickaxe-shaped setae. A hook is depicted (fig. 82) to show the length of the manubrium. The number and length of the hooks is about the same in the last thoracic setiger. The abdominal setae are as figured by Marenzeller (1884, pl 3: fig. 2m). The color of the trunk of the animal is red brown; the parapodia are only slightly lighter. The ventral shields are cream white. There are no colored bands in the tentacular crown in the preserved material. Augener (1914) has included this species with hesitation among the synonyms of P. oligophthalmos Grube, and Ushakov (1955) has fol- lowed him. Our specimens seems to be different from P. ehlersi (Gravier), another synonym of P. oligophthalmos, because of the dif- ferent size of the ventral lobe of the collar alone. The two records by Zachs (1933) and Annenkova (1938) of P. oligophthalmos included by Ushakov (1955) are certainly not concerned with P. myriops. Found at station 7. Also collected in March and September 1966 at 47°12’00’’ N, 122°44’45’’ W at 18 m and about 10 m depth. New for the northeastern Pacific. Previously known from Japan. Summary Hight species are newly described: Eunoe uniseriata, Eulalia (Pterocirrus) parvoseta, Laonice pugettensis, Paraspio cirrifera, Chaeto- zone acuta, C. berkeleyorum, Tharyx secundus and T. serratisetis. Information on type material is provided for Eulalia (Eulalia) lericornuta, Gyptis brevipalpa, Exogone lourei, Syllis (Typosyllis) harti, Dorvillea pseudorubrovittata, Laonice antarcticae, L. japonica, and Chaetozone gracilis. Caulleriella annulosa and Neoamphitrite edwards, are new combinations. New characters are described for Micropodarke dubia, Pionosyllis uraga, Aricidea (Aricidea) ramosa, 48 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 Paraonis (Paraonis) wanovi, Praczillella affinis pacifica, Rhodine bitorquata, Scionella japonica and Potamilla (Pseudopotamilla) myriops. Twenty-four new records for Washington and British Columbia waters are given. Intestinal contents of local species of Glycera and Iumbrineris suggest deposit feeding but it is not certain that it is the exclusive mode of nutrition. Literature Cited ANNENKOvA, N. 1922. Apergu de la famille des Chloraemidae (Annelida Polychaeta) de la collection du Musée Zoologique de l’Académie des Sciences de Russia. C.R. Acad. 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Zur Kenntnis des Sublitorals der chilenischen Ktste unter beson- derer Beriicksichtigung der Polychaeten und Ostracoden, Teil II: Die Polychaeten des Sublitorals. Mitt. Hamburgischen Zool. Mus. Inst., vol. 62 (suppl.), pp. 59-305. HEMPELMANN, F. 1931. Erste und zweite Klasse der Vermes Polymera (Annelida): Archian- nelida und Polychaeta. Jn Kiikenthal and Krumbach, Handbuch der Zoologie, vol. 2, pp. 1-212. Hesse, C. 1917. Zur Kenntnis der terebellomorphen Polychaeten. Zool. Bidr. Uppsala, vol. 5, pp. 39-258. 1925. LHiniges iiber die Hesioniden und die Stellung der Gattung Ancistro- syllis. Ark. Zool., vol. 17, pp. 1-87. Imasima, M. 1963. Polychaetous annelids collected off the west coast of Kamchatka. II. Notes on species found in the collection of 1959. Publ. Seto Mar. Biol. Lab., vol. 11, pp. 345-372. 1964. Benthie polychaetes collected by the second cruise of the Japanese Expedition of Deep Seas (JEDS-2). Bull. Nat. Sci. Mus. Tokyo, vol. 7, pp. 235-254. 1966. The Syllidae (polychaetous annelids) from Japan, III: Eusyllinae. Publ. Seto Mar. Biol. Lab., vol. 14, pp. 85-116. Imasima, M., and Hartman, O. 1964. The polychaetous annelids of Japan, pts. I and II. Allan Hancock Found. Publ., Occas. Pap. 26, pp. 1-452. Izuxka, A. 1912. The errantiate Polychaeta of Japan. Journ. Coll. Sci., Tokyo, vol. 30, pp. 1-262. JOHNSON, H. P. 1901. The Polychaeta of the Puget Sound region. Proc. Boston Soc. Nat. Hist., vol. 29, pp. 381-437. Kuawe, W. L., and Dicriz, L. M. 1957. Biology of the blood worm, Glycera dibranchiata Ehlers, and its re- lation to the blood worm fishery of the Maritime Provinces. Bull. Fish. Res. Bd. Canada, vol. 115, pp. 1-37. LEvINSEN, G. M. R. 1883. Systematik-geografisk oversigt over de nordiske Annulata, Gephyrea, Chaetognathi og Balanoglossi. Vidensk. Medd. Naturh. Foren. Kj¢b., pt. 2, pp. 92-354. Lig, U. In press. A quantitative study of benthic infauna in Puget Sound (Wash- ington) 1963-1964. FiskDir. Skr. Ser. Havunders., vol. 14. o2 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 Luzern, J. 1961. Sur une nouvelle espéce de polychéte, Sphaerodoridium commensalis, n. gen., n. spec. (Polychaeta Errantia, famille des Sphaerodoridae) vivant en commensal de Terebellides stroemi Sars. Cah. Biol. Mar., vol. 2, pp. 409-416. McInrosu, W. C. 1900. A monograph of the British Annelids, vol. 1, pt. 2, Polychaeta: Amphinomidae to Sigalionidae. Ray Soc. London, pp. 215-442. 1910. A monograph of the British Annelids, vol. 2, pt. 2, Polychaeta: Syllidae to Ariciidae. Ray Soc. London, pp. 233-524. MaumGren, A. J. 1866. Nordiske Hafs-Annulata. Ofvers. K. Vet.-Akad. Férh. 1865, vol. 22, pp. 355-410. 1867. Annulata polychaeta Spetsbergae, Grénlandiae, Islandiae et Scandi- naviae hactenus cognita. Ofvers. K. Vet.-Akad. Forh. 1867, vol. 24, pp. 127-235. MARENZELLER, E. von 1884. Siidjapanische Anneliden, II. Denkschr. Akad. Wiss. Wien, vol. 49, pp. 197-224. Monro, C. C. A. 1933. The Polychaeta Sedentaria collected by Dr. C. Crossland at Colén in the Panama region and the Galapagos Islands during the expedition of the S.Y. St. George. Proc. Zool. Soc. London 1933, pt. 2, pp. 1039-1092. Moors, J. P. 1903. Polychaeta from the coastal slope of Japan and from Kamchatka and Bering Sea. Proc. Acad. Nat. Sci. Philadelphia, vol. 55, pp. 401-490. 1907. Description of new species of spioniform annelids. Proc. Acad. Nat. Sci. Philadelphia, vol. 59, pp. 195-207. 1909a. The polychaetous annelids dredged by the U.S.S. Albatross off the coast of southern California in 1904, 1: Syllidae, Sphaerodoridae, Hesionidae and Phyllodocidae. Prod. Acad. Nat. Sci. Philadelphia, vol. 61, pp. 321-351. 1909b. Polychaetous annelids from Monterey Bay and San Diego, California. Proc. Acad. Nat. Sci. Philadelphia, vol. 61, pp. 235-295. 1923. The polychaetous annelids dredged by the U.S.S. Albatross off the coast of southern California in 1904. 4. Spionidae to Sabellaridae. Proc. Acad. Nat. Sci. Philadelphia, vol. 75, pp. 179-259. OxupaA, 8S. 1938. Polychaetous annelids from the vicinity of the Mitsui Institute of Marine Biology. Japanese Journ. Zool., vol. 8, pp. 75-105. PreTTiBons#, M. H. 1953. Some scale-bearing polychaetes of Puget Sound and adjacent waters, pp. 1-89. Seattle: Univ. Washington Press. 1954. Marine polychaete worms from Point Barrow, Alaska, with additional records from the North Atlantic and North Pacific. Proc. U.S. Nat. Mus., vol. 103, no. 3324, pp. 203-356. 1963. Marine polychaete worms of the New England region, 1: Aphroditidae through Trochochaetidae. Bull. U.S. Nat. Mus., vol. 227, pp. 1-356. NO. 3667 POLYCHAETES—BANSE AND HOBSON 53 1966. Revision of the Pilargiidae (Annelida: Polychaeta), including de- scriptions of new species, and redescription of the pelagic Podarmus ploa Chamberlin (Polynoidae). Proc. U.S. Nat. Mus., vol. 118, no. 3525, pp. 155-208. 1967. Type-specimens of polychaetes described by Edith and Cyril Berkeley (1923-1964). Proc. U.S. Nat. Mus., vol. 119, no. 3553, pp. 1-23. Retsu, D. J. 1959. An ecological study of pollution in Los Angeles-Long Beach Harbors, California. Allan Hancock Found. Publ., Occas. Pap. 22, pp. 1-119. Rosa, E. 1941. Estudios anelidolégicos, III: Datos para el conocimiento de la fauna de Poliquetos de las costas del Pacffico de México. An. Inst. Biol. Mexico, vol. 12, pp. 669-746, 1962. Estudios anelidolégicos. XXVI. Algunos anélidos poliquetos de las costas del Pacifico de México. An. Inst. Biol., vol. 33, pp. 131-229. Ruiter, F. 1964. Campagne de la ‘“‘Calypso’’: [les de Cap Vert, 5: Annélides Polychétes. Ann. Inst. Océan. Monaco, vol. 41, pp. 113-218. SanpeErs, H. L.; Goupsmira, HE. M.; Miuus, E. L.; and Hampson, G. E. 1962. A study of the intertidal fauna of Barnstable Harbor, Massachusetts. Limnol. Oceanogr., vol. 7, pp. 63-79. SoprrstrOm, A. 1920. Studien tiber die Polychaetenfamilie Spionidae, pp. 1-286. Uppsala: Almquist and Wicksells. SouTHERN, R. 1914. Clare Island Survey, no. 47. Archiannelida and Polychaeta. Proc. Roy. Irish Acad., vol. 31, pp. 1-160. Stgp-BowirTz, C. 1941. Les glycériens de Norvége. Nytt Mag. Naturvidensk., vol. 82, pp. 181-250. Usuakovy, P. VY. 1950. [Polychaeta of the Sea of Okhotsk.] Akad. Nauk SSSR, Issled. Dal’nevost. Morei SSSR, vol. 2, pp. 140-237. [In Russian.] 1955. [Polychaeta of the Far Eastern Seas of the USSR.] Akad. Nauk SSSR, Opredelitel Faune SSSR, vol. 56, pp. 1-445. [In Russian. Transl. (1965) Israel Program Scient. Transl., U.S. Dept. of Com- merce, Washington D.C., pp. 1-419.] ZaAcus, I. 1933. [The annelid fauna of the northern Sea of Japan.] Issled. Morei SSSR, no. 19, pp. 125-137. [In Russian.] U.S. GOVERNMENT PRINTING OFFICE: 1968 dedytet Var ai Oy betes TOR 11 Ply eGR sew Ty Of te ea i 7 us 4 7 4 _ a - “otto tw! Sel eae ft 7 _ 7 L. clivon gs) chapliiatt De pes. bossa . i fae wat, beds ad na i eee ae yu a fie Oe Ae or. ’ ‘ f ¥ i { U ; - i i? { eee arene. @) - : . .. y WHiAs raam/e i080 9 cua aun ot tay 67 eS ; i Sey ey Aah? lias, Up, Peleg aay oleh Rie op a Sime, Seb oti anal item ipeyibeilap! Gato 7 Vivagy cari * ites Gin OF ik cer: op eeam Proce dines’ “ot the United States National Museum SMITHSONIAN INSTITUTION + WASHINGTON, D.C. Volume 125 1968 Number 3668 The Genus Pterodrilus (Annelida: Branchiobdellida) —_——_—.. _—__——— By Perry C. Holt? Visiting Research Associate Department of Invertebrate Zoology The branchiobdellid worms, once greatly neglected, have received somewhat more attention in recent years. Occurring as epizoites on freshwater decapods and, in a few cases, on other crustaceans of the Northern Hemisphere, their distribution, evolution, and relationships with their hosts furnish several interesting problems. In spite of increased interest in them, however, the systematic account of no genus of North American branchiobdellids is at a stage adequate for a satisfactory consideration of many of these broader problems. Such is the case for the genus Pterodrilus, whose members form a distinctive part of the branchiobdellid fauna of eastern North America. In the last 20 years, however, I have gathered together a large num- ber of specimens of the previously known species of the genus plus material representing five new species. It is now possible, therefore, to present a more nearly adequate definition of the genus, redescriptions and new distributional data for the previously described species, descriptions of the new species, and a discussion of the evolutionary and geographical relationships of the genus. These are the objectives of the present paper. 1 Department of Biology, Virginia Polytechnic Institute, Blacksburg, Va. 24061. 1 Ds PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 125 ACKNOWLEDGMENTS.—Part of this paper is based upon an un- published doctoral dissertation written at the University of Virginia (Holt, 1951). Some of the material studied was collected with the aid of grants from the National Science Foundation (NSF G—4439, G-9828, GB-372). The paper was prepared during the tenure of a Visiting Research Associateship at the Smithsonian Institution. As always, I am grateful to Dr. Horton H. Hobbs, Jr., for his interest in and support of my studies in branchiobdellid systematics, for a careful and helpful reading of the manuscript of this paper, and for identifying the host crayfishes. I wish to thank a number of collectors: Dr. Hobbs; Drs. Warwick R. West, Denton W. Crocker, and Joseph F. Fitzpatrick, Jr.; Messrs. C. W. Hart, Jr., and Kenneth W. Simonds; Mrs. Virgie F. Holt. METHODS AND DISPOSITION OF MATERIAL.—The material used in this study that was collected by persons other than Mrs. Holt and myself was collected and preserved in 70% ethyl alcohol. My collect- ing methods and procedures have been described elsewhere (Holt, 1960a, p. 57). Except for some paratypes of new species and a small number of specimens retained for reference purposes in my collections at the Virginia Polytechnic Institute, the material is deposited in the collections of the United States National Museum. Where appro- priate, United States National Museum catalog numbers (USNM) and my personal catalog numbers (PCH) are listed with the locality data given for a species. In all cases, complete locality data are avail- able from my files or from the Registrar, United States National Museum. The terminology used in branchiobdellid taxonomy and evaluation of the taxonomic utility of various characters have also been discussed previously and the reader is referred to these papers (Holt, 1953, 1960a, Holt and Hoffman, 1959), but some relevant explanations and anatomical comparisons are presented, where appro- priate, in discussions of certain species. REVIEW OF THE LITERATURE.—Knowledge of the genus Pterodrilus dates from Moore’s paper entitled “Pterodrilus, a remarkable discod- rilid’’ (Moore, 18952). He separated the genus from previously known branchiobdellids (=“‘discodrilids” of authors, e.g., Vejdovsky, 1884) on the basis of the striking dorsal “appendages” of the two species (P. alcicornus and P. distichus) that he described and assigned to his new genus. His descriptions are excellent for the state of knowledge of the branchiobdellids of his time, and his species are easily recognized. Since then, Pierantoni (1912, pp. 24-25) and Stephenson (1930, p. 801) mentioned the genus in their literature survey. Ellis (1918, pp. 49-51), by means of a key, assigned his species durbini to Pterodrilus; sub- sequently (1919, pp. 254-255), he formally described and ilustrated P. durbini, described P. mexicanus, and listed new locality records for NO. 3668 PTERODRILUS—HOLT 3 P. distichus. Goodnight (1940, pp. 58-63) quoted the original de- scriptions of the four nominal species of Pterodrilus and added a new locality record for P. durbini. Later, he recorded the presence of P. aleicornus in Sinking Creek, Giles County, Va. (Goodnight, 1941b, p-. 468). A new species was recognized and previously known ones of the genus were redescribed in my unpublished dissertation (Holt, 1951, pp. 100-148). Later I reassigned P. durbini, placing it in the newly established genus Hilisodrilus (Holt, 1960b, pp. 173-176). Recently, P. aleicornus and its distribution have been discussed (Hobbs, Holt and Walton, 1967, pp. 61, 71, 73-74). Causey (1955, p. 44) recorded the presence of P. mexicanus in Arkansas. Other than passing refer- ences (e.g., Hoffman, 1963, pp. 294, 295) or mention in various keys, nothing else has been written about the genus Pterodrilus by North American authors. In Europe, however, Moszyifiski (1937, pp. 71-72; 1938, pp. 99-103) and Georgévitch (1955, pp. 200-203; 1957, p. 14) described species that they had assigned to Pterodrilus, but Pop (1965, pp. 223-225) pointed out the obvious fact that these European species were based upon material belonging to the genus Branchiobdella and synonomized them with B. parasita Henle, 1835. They are as follows: Pterodrilus karamani Moszyfiski, 1937; Pterodrilus bidens Georgévitch, 1955; Pterodrilus megas Georgévitch, 1955; Pterodrilus prion Georgé- vitch, 1955; Pterodrilus megodont Georgévitch, 1955; Pterodrilus aliata Georgévitch, 1957; Pterodrilus dantata Georgévitch, 1957. Pteredrilus Moore, 1895 Typr-species.—Pterodrilus alcicornus Moore, 1895a, pp. 449-450, by subsequent designation (Goodnight, 1940, p. 58). Diacnosis.—Small branchiobdellids (known forms less than 2.0 mm in length) of delicate appearance; cylindrical; prosomite of seg- ment VIII always with elevated dorsal ridge, those of other segments often so, dorsal ridges often bearing fan- or finger-like projections; jaws delicate, light in color or colorless, triangular in shape, dental formula 5/4; prostate present, incompletely divided from spermiducal gland; bursa ovoid to pyriform, penis sheath short, penis non-eversible; spermatheca with long ectal duct, bulb clavate or spatulate; anterior nephridia open by common dorsal pore on segment III. ArFInities.—The close relationship of the species of Pterodrilus to those of Cambarincola has been discussed earlier (Hoffman, 1963, pp. 294-295), and the exclusion of the species at present assigned to Cambarincola from the older genus Pterodrilus has elements of arbit- rariness that require discussion. Part of the argument for maintaining the generic staus of the two groups of closely related species is based upon a conservative desire 4 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 126 to preserve nomenclatural stability. The genus Cambarincola, as presently understood, is composed of, by far, the largest number of species of any genus of the order Branchiobdellida. With the exception of the Eurasian genus Branchiobdella, it has the greatest geographical range of any genus now known. In almost all localities where they occur, the species of Cambarincola are the dominant elements of the branchiobdellid fauna. To transfer all these species to the much less well-known and smaller genus Pterodrilus could result only in a period of nomenclatural confusion. This argument alone, however, cannot justify excluding the species now assigned to Cambarincola from Pterodrilus. Moore was struck with the unusual appearance of P. alcicornus and P. distichus with their ornamentation of dorsal projections, and he established a new genus for them. Although he described the male reproductive system of both species (Moore, 1895a, pp. 453, 454), the importance of this system to the systematics of the branchiobdellids was not appreciated at that time, nor, indeed, by Ellis who did, however, present a diagram of it in his paper establishing the genus Cambarincola (Ellis, 1912, p. 483). The difficulty arises from the fact that the basic plan of the repro- ductive system of species of Pterodrilus does not differ from that of the members of Cambarincola as much as it does from other genera of the order. Recent workers (Holt, 1960a, 1960b, 1967a, 1967b, Hoffman, 1963; Laing, 1963) have derived their generic concepts from the major variations in pattern of the male reproductive system, and I regard these variations as furnishing the most usable characters for marshalling groups of species into genera. Also, the jaws of species of Pterodrilus are quite similar in shape and arrangement of the teeth to those of species of Cambarincola. But the jaw patterns are shared by two or more genera in other cases, and the jaws of all species of Pterodrilus are of essentially the same form. We have, however, in Pterodrilus a group of distinctive species that obviously belong together as a specialized offshoot from the main direction of the evolution of Cambarincola. A formal diagnosis obscures by its brevity and technical language the distinctiveness of such a group. The species of Pterodrilus are smaller than those of Cambarin- cola and are characteristically delicate in appearance. The jaws are correspondingly reduced in size and pigmentation. Always there are ridges on some of the segments and usually these ridges bear projec- tions. It is true that both segmental ridges, produced by supernumer- ary muscles (Holt, 1960b, p. 172), and projections of various sorts occur in other genera and that several species of Cambarincola have such ridges. None of the latter species, however, are easily confused with those of Pterodrilus. The male reproductive systems of species of Pterodrilus vary, but the spermiducal gland is relatively short and No. 3668 PTERODRILUS—HOLT 5 thick and the prostate is less completely divided from the spermiducal gland than in any species of Cambarincola. The dorsal projections of such species as P. alcicornus, P. distichus, P. mexicanus and three of the five new species described herein readily set them apart from Cambarincola. There would be no difficulty in maintaining the generic separateness of these species except for the last two of the new species treated herein, which are closely related to the others but lack fan- or finger-like projections on the ridge of segment VIII. This is not unexpected: the species assigned to Ptero- drilus are believed to have arisen from a generalized stock of Cambar- incola as animals adapted to a niche that favored a reduction in size and the production of the ridges and their projections. The species of Péerodrilus are a distinct group that might be placed within a larger group which includes the species assigned at this time to Cambarincola. Since, however, generic status has been accorded these two groups for many years, I prefer to retain both names and assign such taxa as the new species without dorsal projec- tions to one or the other of the existing genera on the basis of judg- ments as to the closeness of affinities with species previously assigned to them. There are precedents for such decisions in many groups; for instance, among the hosts of the branchiobdellids, the genera Procambarus and Orconectes are united by intermediate species that must be assigned rather arbitrarily to either genus (Hobbs, 1967, pes): One species, P. durbini Ellis (1919, pp. 254-255), previously assigned to Pterodrilus has been removed from the genus and referred to the genus Ellisodrilus Holt (1960b, pp. 173-176). Ellisodrilus is one of a group of genera related to Cambarincola and hence to Pterodrilus. It differs from Pterodrilus in the absence of a spermatheca and the asymmetry and other unique features of the bursa. Ceratodrilus, Oedipodrilus, and Magmatodrilus are other related genera. Mag- matodrilus Holt (1967b) lacks a prostate, the bursa is proportionally quite large and there are no dorsal projections; the penial sheath of Oedipodrilus is elongated, enclosing an eversible penis, the prostate is relatively very small and dorsal projections are absent (Holt, 1967a, p. 58); Ceratodrilus Hall (1914, p. 191) is composed of larger worms in which the prostate is extremely reduced in size and the penis is eversible (Holt, 1960a, p. 57). DisTRIBUTION.—The genus Pterodrilus is confined to eastern North America including Mexico. Within this area there are three distinct centers of distribution: the Southern Appalachians with adjacent portions of the Interior Plateau east of the Mississippi River and the glaciated region north of the Ohio River to the Great Lakes and the Saint Lawrence River; the Ozarkian uplift 6 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 north of the Arkansas River in Arkansas, Missouri, and Oklahoma; the eastern slopes of the Sierra Madre Oriental in Veracruz. Obviously, this fragmented range must at some time have been continuous. A discussion of possible migration routes and the evolution of the genus follows the systematic accounts. Key to the Species of Genus Pterodrilus 1. Dorsal projections present:.34. 6 i ndes vs ool se ot 5s OM ee eee 3 Dorsal projections absent, prosomite of segment VIII raised....... 2 2(1). Dorsal ridges on segments I-VIII..... P. missouriensis, new species Dorsal ridge on segment VIII only....... P. choritonamus, new species 3(1). Dorsal projections on raised prosomite of segment VIII only......... 4 Dorsal projections on raised prosomites of other segments in addition to segmenity WI ein el Se ce Peale le ee Baar ote, Sree 6 4(8). Segments I-VIII with ridges............... P. cedrus, new species Segments other than- VIII without ridges. ...-2 .s:.23: av sei oe 5 5(4). Fanlike dorsal projection of segment VIII with 5 prongs; bursa small, ejaculatory, (duct plone mia. acest -ialcmerelene ene P. hobbsi, new species Fanlike dorsal projection of segment VIII with 4 prongs; bursa large, ejaculatory duct shorten cece aieteenenc mieten eee ee P. mexicanus Ellis 6(3). Two finger-like dorsal projections on segments II-VII, 5 on segment AV TTT i satya Caged ateestie eh ne oncmer oe’ Sach eS banat payee hes lees P. distichus Moore Fanlike dorsal projections on segments III-V, VIII................. ef 7(6). Fanlike dorsal projection lacking on segment II...P. alcicornus Moore Fanlike dorsal projection present on segment II. . . P. simondsi, new species Pterodrilus alcicornus Moore Figures 1, 10 Pterodrilus alcicornus Moore, 1895a, pp. 450-453.—Pierantoni, 1912, p. 25.— Ellis, 1919, p. 254.—Goodnight, 1940, pp. 58-50; 1941, p. 468.—Hobbs, Holt, and Walton, 1967, pp. 61-62. TYPE-SPECIMENS.—The material from Johns River, Watauga County, N.C., upon which Moore based this species, has not been found among Moore’s collections now in the U.S. National Museum nor among the collections of the Academy of Natural Sciences of Philadelphia. The species is distinctive, subsequent identifications are not disputed, and no neotype has been designated. DiaGnosis.—Dorsal ridges on segments II-VIITI; those of III-V, VIII bearing fanlike dorsal projections; bursa ovoid, less than 4 body diameter in length; ejaculatory duct of medium length; length of spermiducal gland about 3 times its diameter; prostate subequal in diameter to that of spermiducal gland and % to % its length, histo- logically differentiated from the latter; spermatheca longer than body diameter, bending dorsad to gut, ectal duet long and narrow, bulb clavate, ental process absent. NO. 3668 PTERODRILUS—HOLT rf Description.—The length of individuals of Pterodrilus alcicornus (based on 10 specimens) is about 1.3 mm. The head is slender, its length about % that of the body and its diameter about ¥% that of the greatest body diameter. The intersegmental grooves of the head, except for that setting off the peristomium, are indistinct. The trunk or body is cylindrical throughout and increases grad- ually in diameter up to the reproductive segments (V—-VII), which are all essentially the same diameter (about 0.25 mm). The sucker, formed from segment XI, is subequal to or slightly greater than the head in diameter. The dorsal appendages or projections are borne on ridges of the prosomites. In P. alcicornus, they are paired lobes that diverge some- what and extend laterally and anteriorly in the case of the anterior three to form forward-facing concavities. The projection of segment VIIT is similar, except that it faces posteriorly. The lobes (‘‘wing- like”? projections) bear conical prongs, usually three on each side, although the number varies from one to four prominent prongs, with smaller ones frequently present. The lobes of the projection of seg- ment V do not flare out quite so much as do those of the others and the prongs project more nearly upward. Dorsal ridges are present on the prosomites of segments VI and VII. The spermiducal gland is thick for its length, with a length- diameter ratio of about 3:1. In length and diameter, the prostate is about *4 of these dimensions of the spermiducal gland though it often appears in whole mounts to be subequal in diameter to that of the latter. The prostate is differentiated. The ejaculatory duci is about % the length of the bursa and slightly expanded along its midlength. The diameter of the bursa is approximately * its length. The bursal glands mentioned by Moore (1895a, p. 454) are not present. He described as glands the fold of the wall of the bursal atrium that becomes the ‘“‘rim of the cup” of the everted bursa. The ectal duct of the spermatheca is long and slender, widening gradually into the bulb, which is also long and bends over the gut dorsally; the total length of the spermatheca exceeds the body diameter. There is no ental process of the spermatheca though the ental end of the bulb may resemble such a process when incom- pletely filled with spermatozoa. Discussion.—The following account of the anatomy of P. alcicornus is a condensation of my unpublished earlier treatment (Holt, 1951, pp. 101-115). Serial sections were used as well as whole mounts and the earlier observations confirmed by more recent examination of many specimens. The abundance of this material affords an oppor- tunity to describe P. alcicornus is some detail and, thereby, present 8 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 125 a treatment of the anatomy of the species that will serve to introduce the reader more fully to features common to all species of the genus and as a basis for the shorter descriptions of the other species of Pterodrilus that follow. Ficure 1.—Pvrerodrilus alcicornus: a, lateral view of reproductive systems; b, animal from Giles County, Va. (b=bursa; ejd=ejaculatory duct; pr=prostate; sb=bulb of sper- matheca; sd=spermathecal duct; sg=spermiducal gland; vd=vas deferens.) No. 3668 PTERODRILUS—HOLT 9 The dorsal ridges, which may bear projections in P. alcicornus and other species of Pterodrilus and may occur without projections in species of other genera, are formed by the attachment of muscles (‘supernumerary muscles,” Holt, 1960b, pp. 171-172) that are shorter than the segment to the cuticle and that, by their contraction, differentially shorten the dorsal surface of the prosomites in which they occur. There has been no suggestion by anyone, nor is it indicated in their structure, as to the function of the dorsal projections. They consist of flat ‘“‘wing-like” or cylindrical ‘‘finger-like” (prongs) exten- sions of the epidermis covering the dorsal ridges. The wall of the projections is a single-cell-layer thick (as is the epidermis) with unicellular glands as a prominent feature. The interior of the pro- jections is an irregular cavity that does not appear to communicate with the coelom. The finger-like projections are usually set off by a slight constriction. Dorsal ridges that are present on segments VI and VII do not bear projections on these reproductive segments in any species of the genus. Since the jaws of all species of Pterodrilus are monotonously similar, they have not been illustrated for all the species included in this study (see, however, figs. 2, 4, 5, 7). They are small, delicate and light yellowish brown (but see p. 11, below). The upper jaw bears five sharply pointed teeth, the lower four (dental formula 5/4); and they are more nearly quadrate in shape than is usual among species of Cambarincola with the same dental formula. Moore (1895a, p. 425) believed both jaws of P. alcicornus to be quadridentate, but this was probably because of the smallness of the lateral teeth of the upper jaw, which may cause one of them to be overlooked. All branchiobdellids possess two pairs of nephridia and the anterior pair may open by either separate pores or by a common pore on the dorsum of segment III. The anterior nephridia of Pterodrilus and related genera open by a common pore, which in P. alcicornus is located at the base anteriorly of the dorsal projection. The nephridio- pore usually cannot be seen in animals mounted entire, but there is little doubt that this arrangement is consistently present in Pterodrilus. The innermost parts of the branchiobdellid male reproductive system consist of testes in segment V or in segments V and VI, a pair of sperm funnels and sperm ducts (vasa efferentia) in each testicular segment, and a vas deferens from each of these segments that is formed by the union of the sperm ducts. These elements are quite similar throughout the order and will not be described here (but see Moore, 1895b, pp. 519-521; Holt, 1949, pp. 538-541, 550-552). The spermiducal gland is formed by the union of the vasa deferentia and does vary in shape and structure. Other than its peritoneal 10 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 covering and a very thin layer of circular muscles, the gland consists of a tube composed of a single layer of columnar epithelium, the individual cells of which open into a narrow lumen. All of these gland cells are filled with granules though differences in the secretory cycle of individual cells can be detected in sectioned material. There are no apparent differences in the histological appearance of the spermi- ducal gland of P. alcicornus and the other species of Pterodrilus and that previously described for a species of Cambarincola (Holt, 1949, p. 552). The prostate is a glandular diverticulum of the spermiducal gland that lies along the anteriodorsal border of the latter, ends blindly entally and opens either into the spermiducal gland somewhere along its ectal portion or with it into the ejaculatory duct. In P. alcicornus and the other species of Pterodrilus, the prostate opens into the sper- miducal gland somewhat entally to the junction with the ejaculatory duct; that is, it is incompletely divided from the former. The prostate is of the same basic structure as the spermiducal gland from which it arises, but in P. alcicornus its glandular epithelium consists of highly vacuolated cells with different staining properties from those of the spermiducal gland. This histological difference between the two glands is readily apparent in well-prepared whole mounts and is present in all mature individuals of species of Pterodrilus with the exceptions mentioned below (pp. 16, 25). The expressions ‘differentiated’ and “undifferentiated” are used to distinguish between such prostates as those of P. aleicornus and those that are histologically lke the sper- miducal gland in both sectioned material and whole mounts. In many species of the genus Cambarincola, the ental end of the prostate consists of a thin-walled bulb, the interior of which is a cavity (Holt, 1949, p. 553; 1960a, p. 63). There is no prostatic bulb in P. alcicornus. The ejaculatory duct is a muscular tube which is found in most branchiobdellids and unites, if present, the spermiducal gland and the copulatory bursa. That of P. alcicornus is not unlike that of other species of the genus except in length. The copulatory bursa of P. alcicornus and other species of the genus is quite similar to that of members of the genus Cambarincola. The penial sheath region of the bursa in P. alcicornus is not demarcated externally from the bursal atrium and composes about half of the organ. When the atrial portion of the bursa is everted, the penis is protruded as a short and relatively slender tube surrounded by the cuplike everted bursal atrium. More detailed descriptions of the type of bursa found in the species of Pterodrilus may be found in Holt (1949, pp. 553-555) and Hoffman (1963, pp. 289-290). The ovaries and ovipores of all branchiobdellids seem to be basically similar (Moore, 1895b, pp. 524-525; Holt, 1949, pp. 545-547, 560). NO. 3668 PTERODRILUS—HOLT 11 The spermatheca, however, does vary. The length and diameter of both the ectal duct and bulb may differ among the species of Ptero- drilus; the bulb may be thin walled as it is in P. alcicornus and many other branchiobdellids (Holt, 1949, p. 560, fig. 18), and an ental process may be present. The terms used for these parts of the spermatheca are defined in Holt (1960a, p. 64). The organ systems of P. alcicornus and its congeners, with the exception of those discussed above, are not noticeably different from those of other branchiobdellids. VariaTions.—The foregoing description and discussion of P. alcicornus is based primarily upon specimens from the New River drainage in Virginia. Differences in methods of killing and preserva- tion, that is, the use of dilute solutions of alcohol, produce some distor- tion of the specimens. There is little of note in the way of intrapopula- tional variation, except for differences in the number of prongs of the dorsal projections. A count of these for 10 specimens from Gules County, Va., gave the following results: segment IIL segment IV segment V segment VIII 5 4 5 7 a 6 6 a 5 5 5 a 7 5 5 a 6 6 5 6 8 a at 7 7 5 5 6 if s 5 6 5 3 4 5 5 5 3 i) Similar results were obtained from specimens from other parts of the range of the species. The “wings” of the dorsal projections of at least some specimens from the Watauga River system in North Carolina and Tennessee and the headwaters of the New River in North Carolina are of greater extent than those of specimens from other parts of the species’ range. The latter material, however, is poorly preserved, which may account for the apparent differences. The jaws of some specimens from Alleghany County, N.C., are much darker than usual. That this may reflect something other than geographical variability is indicated by the presence of dark jaws in at least one collection from Giles County, Va., where most specimens have lightly colored jaws. The extent of variability in the species is not enough, or at least it is not well enough defined geographically, to allow one to consider the erection of subspecies. ArFinities.—Pterodrilus alcicornus, in external appearance, is 12 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 125 most like P. simondsi and P. distichus. It differs from both in the absence of dorsal projections on segment II and from P. distichus in the fanlike instead of finger-like nature of the projections. The re- productive systems of P. alcicornus differ in only minor details, mostly in length and shape of the spermatheca, from those of P. hobbst and P. distichus and in the fully differentiated prostate and thin- walled spermathecal bulb from P. simondsi. Hosts.—Pterodrilus alcicornus was found with the following cray- fishes: Cambarus sciotensis Rhoades, C. bartonii bartonit (Fabricius), C. robustus Girard, C. bartonii subspecies, C. longulus longirostris Faxon, C. longulus chasmodactylus James, Cambarus species, Or- conectes juvenilis Hagen, Cambarus parvoculus Hobbs and Shoup, C. bartonit cavatus Hay, C. veteranus Faxon, C. acuminatus Faxon, C. longulus longulus Girard, Orconectes sanborni sanborni (Faxon). The most frequent hosts are C. sciotensis and C. bartonit bartoni. DistripuTion.—Pterodrilus alcicornus is widespread in the streams of the Tennessee and New Rivers in Tennessee, North Carolina, and Virginia. In addition, it has moved—apparently recently since it is not common there—into other adjacent drainages: the Savannah River in Transylvania County, N.C., the James River drainage in Craig County, Va., the Roanoke River drainage in Franklin and Patrick Counties, Va., the Big Sandy River drainage in Buchanan and Dickenson Counties, Va., and Wyoming County, W. Va. (fig. 10). Most of my collections of P. alcicornus are from Virginia, and the greater number of known localities for the species in the New River drainage in Virginia may be, but probably is not, a peculiarity of collecting. The range as given here may not be complete for it is possible that P. alcicornus occurs in other adjoining drainages. MATERIAL EXAMINED.—Several hundred specimens from 122 collections were studied. The bulk of this materia! is deposited in the U.S. National Museum (USNM 36184-36250). Pterodrilus distichus Moore Ficurss 2, 10 Pterodrilus distichus Moore, 1895a, pp. 453-454.—Pierantoni, 1912, p. 25.—Hall, 1914, pp. 190, 193.—Ellis, 1919, p. 254.—Goodnight, 1940, pp. 60-61; 1943, p. 100. TYPE-SPECIMENS.—The material from western New York, upon which Moore based this species, has not been found. The species is distinctive, subsequent identifications are not disputed, and no neotype has been designated. Diacnosis.—Low, somewhat indistinct ridges on segments I-VIII, those of segments II-VII each with two bluntly pointed cylindrical NO. 3668 PTERODRILUS—HOLT 13 \ Ficure 2.—Pterodrilus distichus: a, lateral view of reproductive systems; b, en face view of jaws, upper jaw above; c, animal from Seneca County, N.Y. 14 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 dorsal projections, that of segment VIII with five projections; bursa subspherical, small, hardly reaching ventral border of gut; ejaculatory duct of medium length; prostate broadly joined to spermiducal gland, subequal in length and diameter to latter, differentiated; spermiducal gland small, its length about twice its diameter; spermatheca clavate, length subequal to body diameter, ectal duct gradually merging into bulb, without ental process. DescriIpTion.—In size the members of this species differ very little from those of P. alcicornus, but are perhaps slightly larger, with an average length of 1.4 mm. The dorsal ridges would hardly attract attention if they did not bear projections. The latter are relatively short and about io the greatest body diameter in length. There are two of these on segments JI-VIT and five on segment VIII. The spermiducal gland, ejaculatory duct, bursa, and penis differ from those of P. alcicornus only in their smaller size. The spermatheca is not proportionally as long as that of P. alcicornus and does not bend appreciably over the gut although its length is approximately equivalent to the body diameter; the ectal duct appears to be somewhat wider than that of P. alcicornus and is not greatly different in diameter from the bulb at the union of the two. VarriatTion.—Minor differences that may be noted in the size of specimens, the length of the dorsal projections, and the size and proportions of the reproductive systems perhaps are best correlated with differences in age or nutrition and in the methods of killing and preserving. ArFinitiges.—The differences between P. distichus and P. alcicornus have been noted (p. 12). The two species are closely related. In the number of dorsal appendages, P. distichus agrees with P. simondst, but the reproductive systems of these two species are significantly different (p. 25). Hosts.—Pierodrilus distichus has been associated with Orconectes propinguus (Girard), O. immunis (Hagen), O. obscurus (Hagen), O. juvenilis (Hagen), O. rusticus rusticus (Girard), Cambarus robustus Girard, C. bartonii barton (Fabricius) and C. longulus chasmodactylus James, of which the two most frequent hosts are O. propinguus and C. robustus. Distrripution.—Pterodrilus distichus has been taken from the states of New York, Ohio, Kentucky, Indiana, Illinois, and Michigan (fig. 10). All of these records are from regions covered by ice during the Wisconsin glaciations except those from Breathitt, Madison, Jessa- mine, and Harrison Counties, Ky. The first three of these Kentucky records are from the Kentucky River drainage, the last from the Licking River system, both streams of the Ohio drainage. The con- clusion is that the ancestors of P. distichus have moved from some- NO. 3668 PTERODRILUS—HOL' 15 where near the Kentucky River across the Ohio into the glaciated areas of the Ohio-Mississippi and Great Lakes drainage systems since the melting of the Wisconsin glacier. MarterraL Examrnep.—Approximately 200 specimens from 25 collections have been examined. Specimens from all these collections are deposited in the U.S. National Museum (USNM 17651-17653, 36160-36183). Pterodrilus mexicanus Ellis FIGURES 3, 9 Pierodrilus mexicanus Ellis, 1919, p. 254.—Goodnight, 1940, p. 63.—Causey, 1955, p. 44. TyYPE-sPECIMEN.—Holotype, USNM 17654, from Mirador, Vera- cruz, Mexico. Host: Cambarus mexicanus Erichson; Nelson and Gold- man, collectors. Draenosis.—Dorsal ridge on segment VIII, typically bearing four conical projections, remainder of segments without ridges; bursa large, elongate, length exceeding 4 body diameter; ejaculatory duct short; prostate about }; diameter of and subequal in length to spermiducal gland, undifferentiated; spermatheca shorter than body diameter, bulb thick walled. Description.—Pterodrilus mexicanus differs externally from other species of Pterodrilus in the arrangement and number of the dorsal ridges and projections. There are no ridges present, except that on segment VIII, which has four finger-like, conical projections, very similar to those of P. distichus. The total length averages 1.1 mm. The spermiducal gland is about three times its diameter in length and lies along the upper border of the gut. The prostate, subequal in length to and about half the diameter of the spermiducal gland, is histologically undifferentiated in most specimens although some specimens show a vacuolation of some cells along its ental and dorsal borders. The ejaculatory duct is very short. The bursa, however, is large, about 1% times longer than that of P. alcicornus and 3 times that of P. distichus. This great increase in size is primarily accounted for by an increase in the length of the atrial area, which is not only larger but has additional inwardly directed folds of the bursal wall. The penial sheath region and the penis itself is as in other species of Pterodrilus. Specimens with the bursa everted have not been seen; but one would expect a cup-within-a-cup structure to be produced by eversion. The spermatheca of P. mexicanus is shorter than that of most species of the genus, hardly extending above the upper border of the gut. The inner part of the ectal duct is often enveloped in an expanded ectal part of the bulb. The blind end of the spermatheca frequently 16 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 125 resembles an ental process, except that often it is distended with spermatazoa. The wall of the spermathecal bulb is thicker than in all other species of the genus except P. simondsi. VARIATION.—One or two specimens have only three dorsal pro- jections on segment VIII instead of four. The length of the prongs vary, those of the type are larger than those of most specimens. This difference appears to be of sporadic occurrence and of no systematic importance. The prostate of some specimens is partially vacuolated (cf. P. simondsi, p. 25, below). AFFINITIES.—Pterodrilus mexicanus is related to P. missouriensis, P. choritonamus, P. cedrus, and P. hobbst. In features of the repro- ductive system, P. mezxicanus is most similar to P. missouriensis. These two Ozarkian species differ in that in P. mexicanus the ejacu- latory duct is short, the bursa is larger, the prostate is partially differentiated, the spermatheca is shorter and the wall of the sper- mathecal bulb is thicker, and there are no dorsal ridges on segments J-VII. The absence of dorsal ridges except on segment VIII ally P. mexicanus with P. hobbst, a more advanced member of the same lineage (see below, p. 36), which differs from P. mezxicanus in the fully differentiated prostate, long ejaculatory duct, small bursa, and thin-walled spermathecal bulb. Pterodrilus mezxicanus shares the absence of dorsal ridges, except on segment VIII, with P. choritonamus, which, however, lacks projections on this dorsal ridge. In addition, the latter species differs from P. mexicanus in that the bursa is smaller, the ejaculatory duct is longer, the prostate is differentiated, and the spermatheca has an ental process. Pterodrilus cedrus belongs in the same lineage as P. missourvensis and differs from P. mezxicanus in the presence of dorsal ridges on segments I-VIJ, a smaller, more nearly spherical bursa, a longer ejaculatory duct, a differentiated prostate, and a longer spermatheca without an ental process. Hosts.—Pterodrilus mezicanus has been taken from 10 species of the genus Orconectes: O. punctimanus (Creaser), O. ozarkae Williams, O. mecki meeki (Faxon), O. neglectus neglectus (Faxon), O. nana nana Williams, O. nais (Faxon), O. luteus (Creaser), and O. hylas (Faxon). DistrisuTion.—Two of my collections are from the Arkansas River drainage in northwestern Arkansas and eastern Oklahoma; four are from the St. Francis River system in Missouri; the remainder are from the White River drainage in Arkansas and Missouri. All of these streams, however, drain the Ozark highlands centered in south- central Missouri. The range of P. mezicanus, thus, is compact and well delimited except for the type-locality, Veracruz, Mexico. MATERIAL EXAMINED.—Approximately 100 specimens from 22 collections from Arkansas, Missouri, and Oklahoma were studied. NO. 3668 PTERODRILUS—HOLT 17 Ficure 3.—Pterodrilus mexicanus: a, lateral view of reproductive systems 35, animal from Wayne County, Mo. 813-169—68——3 18 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 This material, for the most part, is deposited in the U.S. National Museum (USNM 36138-36159). Remarks.—The holotype of P. mezicanus Ellis (1919) from Mirador, Veracruz, Mexico, is poorly preserved, making a study of the internal structures impossible; it was separated from other bran- chiobdellids by Ellis (1919, p. 254) on the sole basis of the ‘‘simple four-horned appendage like that on the same segment of P. distichus.”’ I have collected branchiobdellids in Mexico and unsuccessfully have tried to locate Mirador. Among the 64 collections from Mexico that I have studied, there are no specimens that can be assigned to the genus Pterodrilus. I have, therefore, with considerable hesitation, referred my material from the Ozarks to P. mexicanus. The possibility remains that a future discovery of P.mezicanus at or near the type- locality will necessitate the renaming of the Ozarkian animals. Pterodrilus hobbsi, new species Fiaurss 4, 9 TypE-sPECIMENS.—Holotype, USNM 36486, and five paratypes, USNM_ 36487, from Cambarus rusticiformis Rhoades, Orconectes quvenilis (Hagen), and O. placidus (Hagen) taken from Spring Creek, 1.4 miles north of the Putnam County line on State Highway 43, Overton County, Tenn., by Perry C. and Virgie F. Holt, July 26, 1961. Draenosis.—Dorsal ridge present on segment VIII, bearing fanlike projection with five prongs, other segments without dorsal ridges and projections; bursa small, ovoid, length less than half body diam- eter; ejaculatory duct of normal length; spermiducal gland relatively large; prostate about %4 diameter of and subequal in length to sper- miducal gland; spermatheca clavate, bending dorsally over gut. Erymo.tocy.—I take great pleasure in naming this species in honor of Dr. Horton H. Hobbs, Jr., as a token of my gratitude for the many years of friendly help he has given me in my study of the branchi- obdellids. Description.—In shape and size, P. hobbst is much like other members of the genus, differing from all except P. choritonamus and P. mezxicanus in the absence of ridges on the prosomites of all segments except the eighth. The dorsal projection of this segment is fanlike and bears five tapering prongs of which the median is the longest. The length ranges from about 1.3 to 1.7 mm. The spermiducal gland is not markedly different from that of P. mexicanus; the prostate, however, is distinctly vacuolated; that is, it is histologically differentiated, but there is no distinct prostatic bulb. The two organs are broadly joined and often the spermiducal NO. 3668 PTERODRILUS—HOLT 19 gland lies so that the true extent and appearance of the prostate is obscured. The ejaculatory duct is markedly longer than that of the unusually short one of P. mezxicanus and is expanded along its mid- length. The bursa is small and ovoid, intermediate in size between those of P. alcicornus and P. distichus, and much _ smaller that of P. mexicanus. ‘ Ficure 4.—Pterodrilus hobbsi: a, lateral view of reproductive systems; b, en face view of jaws, upper jaw above; c, animal from Lee County, Va. 20 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 125 The spermatheca is quite similar to that of P. distichus, perhaps slightly broader along the midlength of the bulb. It is approximately as long as the body diameter and bends dorsad over the gut. The ental end is composed of larger cells and in many specimens there is a, small ental process. VARIATION.—The prongs of the dorsal projections vary in length, but this may be related in part to differences in the degree of con- traction. The reproductive organs show only minor differences in apparent shape and proportions—except for the prostate, which appears to be variable in length, and the extent of vacuolation. The ectal portion near its junction with the spermiducal gland is often not differentiated, but in all individuals the blind end is vacuolated to a greater extent and more consistently than in P. mezxicanus. Arrinities.—Pterodrilus hobbsi is similar to P. choritonamus and P. mexicanus in the absence of dorsal ridges on all segments except the eighth, but it differs from both of these species in the smaller bursa and from P. mezxicanus in the presence of five instead of four prongs of the dorsal projections and in the consistently differentiated prostate (see p. 16). The reproductive systems are most like those of P. distichus and P. alcicornus, species with dorsal projections on several segments. Hosts.—Pterodrilus hobbsi has been taken in association with 17 species and subspecies of Cambarus and 4 species of Orconectes: Cambarus tenebrosus Hay, C. longulus longirostris Faxon, C. parvoculus Hobbs and Shoup, C. longulus chasmodactylus James, C. robustus Girard, C. veteranus Faxon, C. friaufi Hobbs, C. extraneus Hagen, C. bartonii cavatus Hay, C. sciotensis Rhoades, C. distans Rhoades, C. bartonii bartonii (Fabricius), C. longulus longulus Girard, C. latimanus (LeConte), C. striatus Hay, Cambarus species, C. barton subspecies; Orconectes erichsonianus (Faxon), O. juvenilis (Hagen), O. rusticus forceps (Faxon), and Orconectes species. The most common hosts were Cambarus longulus longirostris, C. bartonii cavatus, and Orconectes juvenilis. DistrisutTion.—Fterodrilus hobbsi inhabits most of the upper Tennessee drainage system and is especially common in tributaries of the Nolichucky, Watauga, Holston, Powell, and Clinch Rivers. It has also invaded the New River in Bland and Carroll Counties, Va., and Alleghany County, N.C.; the Big Sandy in Dickenson County, Va.; and is at home in a wide stretch of the Cumberland River drainage in Tennessee and Kentucky (fig. 9). MATERIAL EXAMINED.—Types and over 300 specimens from 62 collections have been examined. The major part of this material is deposited in the U.S. National Museum (USNM 36488-36508). No. 3668 PTERODRILUS—HOLT 21 Remarks.— Unfortunately, much of the material on which the description of P. hobbsi is based is poorly preserved since it was col- lected by students of crayfishes whose requirements are such that the branchiobdellid material in their collections often proves unsuitable for careful study or positive identifications. There is no doubt that P. hobbsi is a distinct species occupying an extensive range; future studies based upon larger series of collections that are better preserved may reveal the presence of other and similar species among the animals presently assigned to this species (see below, p. 32). Pterodrilus cedrus, new species Figures 5, 10 TYPE-SPECIMENS.—Holotype and five paratypes, USNM 36464, from Orconectes placidus (Hagen) and Cambarus tenebrosus Hay taken in a small stream at the intersection of State Highways 52 and 53 at Celina, Clay County, Tenn., by Perry C. and Virgie F. Holt, July 25, 1961. Draenosis.—Dorsal ridges on segments I-VIII, that of VIII bearing four short conical projections; bursa subspherical, small, reaching ventral border of gut; ejaculatory duct of medium length; prostate about % diameter of and equal in length to spermiducal gland, differentiated; spermatheca frequently exceeding body diameter in length, strap shaped to clavate, ectal duct long. ErymMoLoey.—Latin, cedrus, the cedar tree, by extension as a common name, the red cedar, Juniperus virginiana, for the cedar glades that are such a conspicuous part of the landscape of middle Tennessee. Description.—Pterodrilus cedrus is a small worm, about 1.0 to 1.3 mm long; the combination of dorsal ridges on the first eight body segments and the four finger-like projections borne on the ridge of segment VIII are distinctive. These projections are short and resemble closely those of the corresponding segment of P. distichus. The spermiducal gland is small, approximately twice its diameter in length. The prostate is broadly joined to the spermiducal gland and composed of highly vacuolated cells that end abruptly at the level of the separation of the two. It extends entally to the ental end of the spermiducal gland. The ejaculatory duct is about equal in length to the bursa and therefore longer than that of P. mexicanus and perhaps somewhat shorter, relative to the size of the organs, than that of P. hobbsv. The bursa is much like that of all members of the genus, except P. mezxicanus, P. choritonamus, and P. missouriensis, that is, small and subspherical in shape. 313-169—68——4 22 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 The spermatheca is long, with a long ectal duct. The bulb is elongate oval and usually bent mesiad over the gut dorsally. There is no ental Ficure 5.—Pterodrilu cedrus: a, \ateral view of reproductive systems; }, oblique view of jaws; c, holotype. NO. 3668 PTERODRILUS—HOLT 23 process. The organ is narrower and the ectal duct is longer than is usual in the genus. VariatTion.—Observable variability is confined to the spermatheca and seems to depend upon the degree of distension of the bulb with spermatozoa. When incompletely distended, there appears to be an ental process and the ectal duct is long; when fully distended, the ental process disappears and the ectal duct is shorter; that is, the ental part of the duct becomes part of the bulb. A cursory inspection might lead to the conclusion that there are structural differences of the spermatheca among individuals of the same population. AFFINITIES.—Pterodrilus cedrus is superficially most like P. hobbsi but differs in having dorsal ridges on segments I-VIII, the shorter and finger-like dorsal projections of segment VIII instead of the fanlike projection with five prongs of the latter, the longer ectal duct of the spermatheca, and the narrower spermathecal bulb. The dorsal ridges of P. cedrus allies it, however, with the lineage culminating in P. alcicornus. Among these species (P. simondsi, P. distichus, and P. alcicornus), P. cedrus differs from P. distichus most markedly in the absence of two finger-like projections on the dorsal ridges of segments II-VII and the presence of four, instead of five projections on segment VIII. Hosts.—Pterodrilus cedrus has been taken with the following cray- fishes: Orconectes placidus (Hagen), O. rusticus subspecies, O. guvenilis (Hagen) and Cambarus tenebrosus Hay. Distripution.—Pterodrilus cedrus is known only from a small series of collections taken in the eastern Highland Rim and Nashville Basin regions of Tennessee (fig. 10). Both its anatomical features and re- stricted distribution impute to it the status of a phylogenetic relict. MATERIAL EXAMINED.—Types and 37 additional specimens. With the exception of three paratypes (PCH 1396) from the type-locality, this material is deposited in the United States National Museum (36465-36468). Pterodrilus simondsi, new species Fiaurss 6, 10 TypE-SPECIMENS.—Holotype, USNM 36477, five paratypes, USNM 36478, from Cambarus bartonii bartonii (Fabricius) taken in a tribu- tary to the Ocoee River, 12.2 miles south of Morganton, on State Highway 60, Fannin County, Ga., by Kenneth W. Simonds, Nov. 6, 1958; four paratypes, PCH 989, from Cambarus bartonii bartonii taken in a tributary to the Ocoee River, 8.8 miles south of Morganton, Fannin County, Ga., on State Highway 60, by Kenneth W. Simonds, Nov. 6, 1958. 24 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 DiaGnosis.—Dorsal ridges on segments II-VIII, those of segments IL-V, VIII bearing fanlike projections; bursa of medium size, with expanded atrial region; spermiducal gland relatively long, exceeding slightly anteroposterior dimension of segment VI in length; prostate % to subequal to spermiducal gland in diameter, subequal in length, - f Me ( Ficure 6.—Pterodrilus simondsi: a, lateral view of reproductive systems; }, holotype. NO. 3668 PTERODRILUS—HOLT 2a histologically differentiated in some specimens, not in others; sper- matheca slightly longer than body diameter, ectal duct long, bulb clavate with thick muscular wall. Erymotoey.—I am pleased to name this species in honor of its discoverer, Mr. Kenneth W. Simonds. Description.—The dorsal ridges reach a greater height than in other species and more of them bear the expanded projections with prongs that may in turn be bifurcated or bear secondary prongs. The generalized description of the dorsal projections of segments III and VIII of P. alcicornus apply to those of segments II-IV and VIII of P. simondsi. The dorsal projection of segment V of P. simondsi, however, lacks the membranous lateral expansions of those of the other segments and are similar to the projections of segment VIII of P. distichus. Pterodrilus simondsi is composed of small worms about 0.9 to 1.8 mm long. The bursa is smaller than that of P. mezicanus, but still larger, or at least longer, than is usual. The ejaculatory duct is prominent and rather noticeably expanded along its midlength. The spermiducal gland is somewhat longer than the anteroposterior dimension of the segment in which it lies and is usually oriented diagonally in the segment, extending dorsally above the gut. The diameter of the prostate ranges from % to subequal that of the spermiducal gland and extends entally to the ental end of the latter. It is more nearly separated from the spermiducal gland than in other species of the genus, the separation between the two extending almost to the junc- tion of the spermiducal gland with the ejaculatory duct. The spermatheca of P. simondsi is comparable in length and general shape to that of such species as P. alcicornus and P. hobbsi, but it differs in the heavier muscular investment of the bulb that distinctly persists even when the bulb is distended to the maximum with spermatozoa. There is no ental process. VaRIATION.—The only detectable variations in the material I have studied are those involving the prostate, which is, in some specimens, histologically identical to the spermiducal gland; in others it is com- posed of large, clear cells (vacuolated cells); and in still others some of the cells are composed of dense cytoplasm with many granules, and others are filled almost entirely with a clear material. In other words, in this species, the distinction between differentiated and un- differentiated prostates breaks down. The degree of distension of the spermathecal bulb is also variable. There can be no doubt that these are individual, intrapopulational variations. AFFINITIES.—Pterodrilus simondsi is closest in external appear- ance to P. alcicornus, from which it differs most noticeably in the presence of an additional dorsal projection on segment II, but the 26 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 reproductive systems of these two species are quite dissimilar. The prostate of P. alcicornus is always differentiated and its spermatheca lacks the muscular investment of the bulb characteristic of P. simondsi. Pterodrilus simondsi, then, is a less advanced member of a lineage derived from ancestors much like P. missouriensis and P. cedrus (see p. 21) that has also produced P. distichus and P. alci- cornus. Hosts.—The following crayfishes were found associated with P. simondsi: Cambarus bartonit bartonii (Fabricius), C. latimanus (LeConte) and Cambarus species. This is the only species of Pterodrilus for which no species of Orconectes is known to serve as a host (but see p. 32 below). Moreover, only once was it found in the absence of C. 6. bartonii, the one record outside the Ocoee River, where it is associated with an unnamed species of Cambarus. Distrisution.—Pterodrilus simondsi is known only from the collec- tions taken by Mr. Simonds from small tributaries to the Ocoee River in Fannin County, Ga., and Cherokee County, N.C., and one collec- tion from a tributary to the Nottely River in Union County, Ga. In 1958-59 Mr. Simonds took 84 collections of crayfish from the Hiwassee River drainage to which the Ocoee and Nottely Rivers are tributary. Of his 19 stations in the upper Ocoee, P. simondsi was present at 14. The streams in which these stations were located are described as “small . . . with cold cascading waters, the bottoms of which are composed almost entirely of large flat stones often with several layers superimposed .... In such streams the water is clear even after heavy rains” (Simonds, unpubl. ms.). The thorough- ness of Mr. Simonds’ collecting efforts in similar streams of the Hiwassee River system to the north (75 collections, only one of which contained P. simondsi) leads to the conclusion that P. simondsi is a highly localized species. It should be searched for in the headwaters of the Savannah River to the east, the Chattahoochee River to the southeast, and the Coosa River to the southwest, but presumably P. simondsi is a relic of an early invasion of the area by primitive rela- tives of P. alcicornus that were adapted to cold, clear mountain streams. MATERIAL EXAMINED.—Types and 53 specimens from 15 localities. The major part of this material is deposited in the U.S. National Museum (USNM 36479-36485). Pterodrilus choritonamus, new species FIGuREs 7, 9 TYPE-SPECIMENS.—Holotype, USNM 36471, and two paratypes, USNM 36472, from Cambarus tenebrosus Hay taken in a tributary to NO. 3668 PTERODRILUS—HOLT 7 Eagle Creek (Holt Spring Branch) about 4.5 miles north of Living- ston, Overton County, Tenn., by Perry C. and Virgie F. Holt, July 24, 1961; five paratypes, PCH 1393, from Cambarus tenebrosus and Orconectes placidus (Hagen) taken in Little Eagle Creek about 0.5 miles above confluence with Eagle Creek and about 6.0 miles north of Livingston, Overton County, Tenn., by Perry C. and Virgie F. Holt, July 24, 1961. Diaenosis.—Without dorsal projections, dorsal ridge present on segment VIII; bursa pyriform, small, extending at most to ventral border of gut; ejaculatory duct of medium length; prostate sub- equal to or shorter than spermiducal gland, diameter about % that of latter, differentiated; spermatheca with long ectal duct, median bulb and ental process. Ficure 7.—Pterodrilus choritonamus: a, lateral view of reproductive systems; J, lateral view of jaws; c, holotype. 28 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 126 Erymo.tocy.—From Greek, choritos, native, and namos, spring or stream, native spring, for the spring branch of my boyhood home. Description.—Pterodrilus choritonamus is a small and delicate worm about 1.1 to 1.5 mm long. In body proportions and outline it is similar to P. hobbsi and P. mexicanus, differing in the absence of projections on the dorsal ridge of segment VIII. The bursa approaches half the body diameter in length and is pyriform in shape: the penial sheath region is set off externally by a slight constriction and is less in diameter than the atrial portion of the bursa. The spermatheca has the midlength of the organ (the spermathecal bulb) normally expanded, the ental portion not, so that there is an ental process that is lined with a columnar epithelium instead of a thin layer of flattened cells as is the bulb. Although the total length of the spermatheca is subequal to the body diameter, it is not as long as that of the other species of the genus except that of P. mezi- canus, which it exceeds in length. VaRIATIONS.—The prostate varies in length, the ental end usually approaching the ental end of the spermiducal gland, but sometimes not. The spermatheca varies in the degree of the distension of the bulbular region, with the result that the extent of the ental process is reduced by a greater expansion of the bulb, but in the specimens I have seen the process is present and may, then, be a constant feature of the species. Arrinities.—Pteordrilus choritonamus is related to P. missouriensis, P. cedrus, P. mexicanus, and P. hobbsi. Its affinities with the first three of these species have been discussed (p. 16). It differs from P. hobbsi in the absence of projections on the dorsal ridge of segment VIII, in the larger bursa and in having an ental process of the spermatheca. Hosts.—The known crayfish hosts of P. choritonamus are Cambarus tenebrosus Hay, C. extraneus Hagen, Orconectes placidus (Hagen) and Orconectes species. Disrrisution.—Pterodrilus choritonamus frequents tributaries of the Cumberland River in the Eastern Highland Rim in Tennessee. MATERIAL EXAMINED.—Types and 28 specimens. The material for the most part is deposited in the United States National Museum (USNM 36473-36476). Pterodrilus missouriensis, new species Figures 8, 9 TyYPE-SPECIMEN.—Holotype, USNM 36469, two paratypes, USNM 36470, and two paratypes, PCH 1476, from Orconectes luteus (Creaser) No. 3668 PTERODRILUS—HOLT 29 taken in Whetstone Creek on U.S. Highway 60, 5 miles west of Moun- tain Grove, Wright County, Mo., by Perry C. Holt, August 23, 1961. Draenosis.—Low dorsal ridges on segments I-VII, higher one on VIII, no dorsal projections; bursa large, its length equalling or ex- Ficure 8.—Pterodrilus missouriensis: a, lateral view of reproductive systems; b, holotype. 30 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 ceeding % body diameter; spermiducal gland relatively long, its length equalling or exceeding anteroposterior dimension of seqment VI, narrowing at ectal end; prostate short, in diameter about % that of spermiducal gland, incompletely divided from latter, histologically undifferentiated, with ental bulb; spermatheca spatulate, its length subequal to body diameter, ectal duct long, ental process absent. Erymo.tocy.—The adjectival form of Missouri. Description.—The length, based on five animals, averages 1.6 mm (range 1.5-1.8 mm). The dorsal ridges of segments I-VII are poorly developed and in some extended specimens might be over- looked. That of segment VIII, however, is well developed. The anterior nephridiopore is clearly visible on the dorsum of segment III. The teeth of the jaws appear to be longer and more sharply pointed than is usual. The male reproductive system in the totality of its primitive aspects, is unlike that of any other species of Pterodrilus. The spermiducal gland is relatively long and slender, its length more than three times its diameter. The prostate arises as a diverticulum of the gland rather far from the latter’s junction with the ejaculatory duct. There is an abrupt narrowing at the point of origin of the prostate, from which point the spermiducal gland continues to decrease in diameter until it passes into the ejaculatory duct. The prostate has a diameter of about half that of the spermiducal gland and its ental end is located about % of the length of the latter from its ental end: in ail, the prostate is about 4 the length of the spermiducal gland and lies along the median third of the gland. The prostate is not histologically differentiated, but there is an ental “bulb”, a cavity of rather small extent. The ejaculatory duct is prominent and noticeably expanded along its midlength. The bursa is large, exceeding half the body diameter in length. The penis is prominent and the penial sheath region of the bursa is larger than usual. The spermatheca has a long ectal duct that expands entally before it merges into the elongated, spatulate bulbular portion. There is no ental process, but in at least some specimens the entire wall of the bulb appears to be composed of large, granular cells with the result that the wall is much thicker than usual. VARIATION.—The prostate appears to be of variable length, but this is probably because of the difficulty of estimating the compara- tive lengths of the prostate and spermiducal gland in specimens in which these organs are viewed from different directions. The ental part of the spermathecal bulb does not always appear to be filled with a glandular epithelium, but this is most likely a reflection of differences in degree of distension of the bulb with spermatozoa. NO. 3668 PTERODRILUS—HOLT 31 Arrinitins.—Pterodrilus missourvensis is a primitive pterodrilid related to P. choritonamus, P. mexicanus, and P. cedrus (p. 21). It shares with P. cedrus the dorsal ridges of segments J-VIIT but differs in the absence of projections on the dorsal ridge of segment VIII, the undifferentiated prostate, the shape of the spermiducal gland, the larger size of its bursa, and in the thicker-walled spermathecal bulb. Pterodrilus missouriensis and P. choritonamus both lack dorsal projec- tions on segment VIII and have large bursae but differ in the presence of dorsal ridges on other segments, the undifferentiated prostate, the thicker-walled spermathecal bulb in the former, and an ental process mg | GA baa Yj. CHORITONAMUS a P. MISSOURIENSIS : P. MEXICANUS P. HOBBSI Ficure 9.—Distribution of certain species of Pterodrilus. of the spermatheca in the latter. Pterodrilus missouriensis shares with P. mexicanus the primitive nature of the prostate (p. 15) (though that of the latter is often partially differentiated), the large size of the bursa, and a spermatheca with a thicker-walled bulb; it differs from the latter in its much longer ejaculatory duct, the presence of dorsal ridges on segments anterior to segment VIII, and in the absence of dorsal projections. Hosts.—The only known host is Orconectes luteus (Creaser). DistRiBuTion.—Pterodrilus missouriensis is known only from the type locality, Whetstone Creek in Wright County, Mo. The one collection was taken from shallow pools in the headwaters of the stream, where there was little or no flow at an elevation of approxi- mately 1260 feet. This locality is near the divide between the south- 32 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 125 ward-flowing White River drainage system and the northward-flowing Gasconade River and is a part of the latter system. There is no other record of a species of Pterodrilus from the Missouri River basin. MATERIAL EXAMINED.—Five type-specimens. Pterodrilus species Poorly preserved material taken from three localities in the Hiwassee River drainage in Union County, Ga., and Cherokee County, N.C. (PCH 915, 974, 979), by Mr. Kenneth W. Simonds may well represent another species of Pterodrilus. These specimens appear to differ from those of other species of Pterodrilus in that there are three or four prongs of the dorsal projection on segment VIII, there appear to be dorsal ridges without projections on the other segments, and the prostate seems to be undifferentiated with a a thick-walled “prostatic bulb.”? The latter two points cannot be confirmed in my material, which raises the question as to whether the differences in the number of prongs of the dorsal projection may not be due to intraspecific variability in P. hobbsi. If the prostate should be differentiated and there are no dorsal ridges other than that bearing the projection on segment VIII, these animals could be distinguished from P. hobbsi only by the number of prongs of the dorsal projection. Better preserved material will almost surely show that these specimens represent a new species, but I am unwilling to describe a species on the basis of such poor material. These specimens are from the following localities in the upper Hiwassee drainage: Union County, Ga., 2.6 miles east of the Fannin County line on U.S. Highway 76, hosts Cambarus latimanus (LeConte), O. bartonti bartonit (Fabricius), Nov. 5, 1958, K. W. Simonds, coll. (PCH 915); Union County, Ga., 0.5 mile north of Vogel State Park on U.S. Highway 19, hosts Cambarus longulus longirostris Faxon, C. carolinus Erichson, Cambarus species, Nov. 5, 1958, K. W. Simonds, coll. (PCH 979); Cherokee County, N.C., 1.4 miles off Joe Brown Road, in Grape Creek, hosts Cambarus bartonii bartonia (Fabricius), Cambarus species, June 6, 1959, K. W. Simonds, coll. (PCH 974). Evolutionary Considerations The genus Pterodrilus is a group of closely related species derived from a primitive stock of the genus Cambarincola that specialized in the direction of small size and presumably a relatively narrow niche on the crayfish host. It would be of considerable importance if we knew more precisely what this niche is. Brown (1961, p. 25) has shown that P. alcicornus is randomly distributed over the ventral surface of the hosts. The other species of the genus almost surely NO. 3668 PTERODRILUS—HOLT aa occupy the same microhabitat. Diatoms make up a goodly part of the food of the species of Pterodrilus and they inhabit creeks and branches in upland regions, but nothing else is known about their ecological requirements. One is forced, then, to discuss their primitive char- acteristics and their subsequent specializations as adaptations fitting them for unknown ways of life. I shall proceed by describing the ee orn Yj P, CEDRUS sp. DISTICHUS \\ / = P. SIMONDSI © P. ALCICORNUS Ficure 10.—Distribution of certain species of Péerodrilus. hypothetical primitive pro-pterodrilus as I conceive it to have been and by defending, along the way, the reasons its various character- istics must be considered primitive. From these hypothetical con- siderations a tentative phylogeny will be derived and this in turn will be tested against the distributional data. Thus, a reasonable, if not necessarily true, story of the evolution of the genus can be written. THE PRIMITIVE Preropritus—The ancestors of Pterodrilus were the smallest of the North American branchiobdellids, not 34 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 greatly, if at all, exceeding 2.0 mm in length. As branchiobdellids go, they were graceful animals, and their size was probably an adaptation that enabled them to escape competition with their larger relatives by retreating farther into the smaller crevices found on the underside of a crayfish than their relatives could and there exploiting the food found in such crannies. The dorsum of the prosomite of segment VIIT on these animals was raised into a ridge by the existence of supernumerary muscles. Such ridges are found on the prosomites of one or several segments of a number of branchiobdellids in genera that are not closely related to Pterodrilus, as well as among the species of the genera Cambarincola and Oedipodrilus Holt (1967a, p. 58). Perhaps this arrangement of the body-wall musculature is related in a mechanical sense to the hirudinoid mode of locomotion adopted by the branchiobdellids. One might conclude, then, that the absence of these ridges on all segments other than segment VIIT is a primitive condition and that the evolutionary trend in Pterodrilus has been in the direction of an increasing number of such ridges. The tendency in the genus Pterodrilus for the dorsal ridges to bear projections of unknown adaptive significance is shared with Ceratodrilus and the Asian Cirrodrilus, genera that are dissimilar to Pterodrilus in most other respects. The primitive progenitor of Ptero- drilus lacked these projections, as the species P. missouriensis and P. choritonamus attest. In spite of our ignorance of the adaptive significance of these projections, it is assumed that the species with few or none are more primitive in this respect than are those with dorsal projections on several segments. The jaws of pro-pterodrilus were generally small and delicate in appearance: the upper bore five teeth; the lower, four. Except for the reduction in size, this is the usual, and probably primitive, pattern in the genus Cambarincola and that found in all species of Pterodrilus. The cylindrical body shape, common anterior nephridiopore and 5/4 dental formula are features shared by Pterodrilus and Cambarincola and hence by the progenitor of Pterodrilus. The innermost parts of the male reproductive system are basically the same in all branchiobdellids (Holt, 1965, p. 26) and nothing needs to be said about the testes in segments V and VI, the efferent funnels and ducts, or the deferent ducts. The spermiducal gland received the deferent ducts entally without the deferent lobes (Hoffman, 1963, p. 286) that are found in some putatively primitive species of Cam- barincola. The gland had a lesser relative diameter and a proportion- ally greater length than that in all the species of today except P. missouriensis and, to a lesser extent, P. mexicanus. NO. 3668 PTERODRILUS—HOLT 35 The prostate was a small gland, about % the length of the spermi- ducal gland, that arose about \ the latter’s length from its junction with the ejaculatory duct. The prostate of the more advanced species of Cambarincola and Pteredrilus is differentiated. In pro-pterodrilus it was undifferentiated and consisted of a lobe of glandular epithelium that was histologically indistinguishable from that of the spermiducal gland. There may have been a prostatic ‘‘bulb” at the ental end that consisted of a few differentiated cells. In more advanced species of Cambarincola, the bulb is a distinctive and specialized part of the prostate. In all species of Pterodrilus the lumen of the prostate opens into that of the spermiducal gland some distance entad to the junction of the latter with the ejaculatory duct. In Cambarincola, the prostate and spermiducal gland usually open together into the ejaculatory duct. That the prostatic glands of the two genera are homologous cannot be doubted, but that of Pterodrilus is closer in this respect than is Cambarincola to Ceratodrilus Hall (Holt, 1960a, p. 57), Elliso- drilus Holt (1960b, p. 172), and Oedipodrilus Holt (1967a, p. 58). The latter genera must on this account and others be considered as primitive relatives of Cambarincola. The histological differentiation of the prostate occurs in the more advanced species of both Cam- barincola (Hoffman, 1968, pp. 287, 301, et seq.) and Pterodriius (only P. missouriensis has a completely undifferentiated prostate). The evolutionary trend in the specialization of the prostate seems to be clear. The ejaculatory duct was probably short; though this supposition is based upon the length of the ejaculatory duct of P. mezicanus, it is strengthened by the fact that in the presumably primitive genera of the branchiobdellids, the ejaculatory duct is absent or short (Holt, 1968). The bursa of pro-pterodrilus was proportionally larger than that found in Cambarincola and all the pterodrilids except P. missouriensis and P. mexicanus. The penial sheath region of the bursa may not have been unusually large, but the penis may have been partially eversible. This conjecture is based upon the opinion (Holt, 1968) that the primitive members of the lineage leading to Cambarincola and Pterodrilus possessed an eversible as opposed to a protrusible penis. The known members of this lineage (Magmatodrilus Holt, 1967b, and an unnamed Mexican genus), which lack a prostate, possess a bursa with a large penial sheath enclosing an eversible or semi-eversible penis; those (Oedipodrilus and Ceratodriius) with incompletely separated prostates likewise have large bursae with eversible penes. Arguments based on the spacial relationships of the set of tubes that is the male reproductive system of the branchiobdel- lids and the conditions in other annelids have been set forth elsewhere 36 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 (Holt, 1968) supporting the hypothesis that an eversible penis, as opposed to the protrusible one, is primitive. If these arguments be allowed, it would be expected that pro-pterodrilus may have been provided with a penis that was proportionally longer and less in diameter than the cone-shaped one of Cambarincola. Such a penis is found in P. missourtensis, P. mexicanus, and P. choritonamus. The spermatheca had an ectal duct that was heavily muscular and entally expanded at its junction with the spermathecal bulb, which in turn was provided with a muscular wall or a thick lining of tall collumnar epithelial cells. There may have been an ental process, but in any case the spermatheca consisted of more diverse elements than the simple muscular tube that is the spermathecal duct and the thin- walled expanded bulb without an ental process characteristic of the advanced members of the genus and of Cambarincola. This opinion is based upon conditions in related but more primitive genera (Holt, 1960a, 1967b, 1968) and upon a consideration of conditions in what are otherwise thought to be primitive species of Pterodrillus, i.e., those with an undifferentiated prostate. A PHYLOGENY OF THE GENUS Preropritus.—Except that it has low dorsal ridges on segments I to VII and that the spermatheca varies in ways difficult to evaluate, P. missouriensis fits remarkably well the above description of the primitive Pterodrilus. But three other species form with this one a group of primitive phylogenetic relicts in the genus: P. choritonamus, P. mexicanus, and P. cedrus. The major problem remaining in the attempt to reconstruct the history of the genus is that of convergence. If one bases a proposed phylogeny on the evolution of dorsal ridges and projections, a quite satisfactory scheme is produced except that there are two distinct lineages of which the more advanced members of each have very similar re- productive systems. Conversely, a phylogeny based on the evolution of the reproductive systems produces a phylogenetic dendrogram that is almost a straight line and places closely together such species as P. hobbsi and P. alcicornus that otherwise are unlike. The solution has been a modified compromise (fig. 11) that assumes a condiserable degree of convergence in the evolution of the reproductive systems, mostly because the alternative would suggest that at least some limeages alternately acquired and lost dorsal ridges and projections, an inherently improbable hypothesis. Two levels of structural specialization were reached in the evolution of Pterodrilus and two minor radiations occurred. Four species (P. missouriensis, P. choritonamus, P. mexicanus, and P. cedrus) compose a group, derived from the original pro-pterodrilus stock, that is characterized by primitive features of the reproductive system and dorsal projections on only one segment or none at all. From the NO. 3668 PTERODRILUS—HOLT alcicornus distichus hobbsi simondsi mexicanus cedrus choritonamus missouriensis PRO-PTERODRILUS Ficure 11.—A phylogeny of the genus Péerodrilus. 37 38 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 125 radiation that produced these species, a form similar to P. mexicanus gave rise to P. hobbsi, which evolved a more advanced type of re- productive system and stands at the second evolutionary level. Pterodrilus cedrus is the survivor of a stock with dorsal ridges and projections that gave rise to the other main lineage composed of P. distichus, P. alcicornus, and, at a more primitive stage of the development of the reproductive systems, P. stmondsi, the members of the second radiation. PLACES OF ORIGIN AND MIGRATIONS.—When the distribution of the species of Pterodrilus (figs. 9, 10) is considered along with the hy- pothesis of their phylogeny that has been sketched here, some con- clusions immediately emerge. The phylogenetically primitive species are scarce and localized. The most primitive of all, P. missouriensis, is known from a single location in the headwaters of the Gasconade River in Missouri. The more abundant but still relatively scarce P. mezicanus is essentially confined to the White River system in Missouri and Arkansas since it is otherwise known only from the nearby St. Francis River in Missouri, a tributary to the Arkansas River in Oklahoma, and Veracruz, Mexico. Pterodrilus choritonamus and P. cedrus are inhabitants of tributaries to the Cumberland River in the Eastern Highland Rim and Nashville Basin regions of Ten- nessee, P. hobbsi is a widespread and successful species of the Cumber- land and Tennessee River systems with outliers in the Big Sandy and New Rivers. Of the species of the lineage with dorsal projections on multiple segments, the most primitive, P. simondsi, is localized in the Hiwassee River drainage of the Tennessee basin; P. distichus is a species of the Kentucky River that has crossed the Ohio to invade the eastern Great Lakes and St. Lawrence drainages; P. alcicornus is found in the Tennessee and New River systems, again with outliers to the east and north in the Savannah, Roanoke, James, and Big Sandy Rivers. The ancestral home of the genus Pterodrilus most likely is in the headwaters of the Cumberland River in the Eastern Highland Rim region of Tennessee. Two of the four most primitive species, P. choritonamus and P. cedrus, still persist as phylogenetic and geographic relicts in this region. The other species are arranged radially around this center in a fashion that almost requires that their ancestors come from the Cumberland (fig. 12). The same general region was the postulated home of the ancestors of the host animals, primitive Procambarus crayfishes that gave rise to the genera Orconectes and Cambarus, with Orconectes spreading mostly to the north and west, Cambarus to the east and south, and some stocks of Procambarus southwestward into Mexico (Hobbs, 1967, p. 15). The modern host relationships of species of Pterodrilus can NO. 3668 PTERODRILUS—HOLT 39 afford little insight into the problems of evolution and migrations of either the hosts or their epizoites: it is well established (Goodnight, 1940, p. 65; Hobbs, Holt, and Walton, 1967, p. 75) that host specificity in the classical sense of a species-to-species correspondence does not occur. Yet it is worthy of note that the crayfish-branchiobdellid as- sociations as recorded under ‘‘Hosts’” for each species is consistent with the hypothesis that Pterodrilus originated in the Cumberland basin and spread from there with the ancestors of the hosts of today, mostly species of Orconectes. An attempt is made (fig. 12) to diagram more precisely the geo- P. distichus Great Lakes P baits, New River RP missouriensis P. distichus Kentucky River PR mexicanus R hobbsi Ozarks P choritonamus R alcicornus — Propterodrilid P- simondsi P cedrus Tennessee River Cumberland River Mexico Figure 12.—The evolution of the genus Pterodrilus. eraphical relationships of the species of Pterodrilus. The times at which all these migrations occurred cannot be determined on the basis of the evidence now available, but the original diversification of the early Pterodrilus stock took place well back in the Tertiary, and the movement of P. distichus into the glaciated regions of the north, of necessity, has happened since the last glaciation. Without attempting to pmpoint the events in time on the basis of the hypothesis developed, we can note that an early diversification of pro-pterodrilus stocks oc- curred in the Cumberland basin. Of this radiation, P. choritonamus and P. cedrus remain in the general area of their ancestral home as relict forms. A primitive species, represented today by P. missouriensis, moved early into the northward-flowing streams of the Missouri 40 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 125 Ozarks. Whether P. mexicanus is a descendant of this stock that moved over the Ozarkian divide into the White River and adjacent drainages cannot be determined with certainty: the postulated phylogeny sug- gests that it hkewise came into the Ozarks by the same route, but if so, it seems somewhat strange that it is unknown from the Missouri basin. Yet few collections have been taken from the northern Ozarks in Missouri, and further field work may well reveal the presence of P. mexicanus there. But P. mezicanus is obviously extinct over much of the route it or its ancestral form must have taken to reach the Ozarks, and the same or similar factors that caused this restriction of range may operate in the streams of the Missouri River system in southern Missouri. It is not surprising that an early stock of Ptero- drilus may have moved into Mexico: the crayfish hosts did so some- time before the end of the Miocene (Hobbs, 1967, p. 15). The possi- bility remains that the Ozarkian worms are not conspecific with the type of P. mexicanus (see p. 18 above), but any solution of the problem of the status of P. mezicanus will fit these ideas; for if it is recovered from Mexico and a new name assigned to my specimens from the Ozarks, the Mexican worms are, on the basis of my study of the type, very similar to the Ozarkian ones. Such a solution, however, would date the early migrations of Pterodrilus stocks in the Miocene or earlier (Hobbs, 1967, p. 15). Turning now to the north and east, we note that there are large gaps in the range of P. distichus (fig. 10) that can only be attributed to inadequate collecting. The records from the Kentucky and Licking Rivers are near the postulated place of origin of the species and may represent the Pleistocene refugium from which P. distichus has moved north and northeastward, most likely by way of the Miami and Scioto Rivers, since the Wisconsin glaciation. The gap in the range of P. distichus in the Lake Erie basin in Pennsylvania and New York surely represents inadequate collecting. Pterodrilus hobbsi has arisen from a stock that also produced P. mexicanus, but it has reached a higher level of development in the structures of the reproductive systems. Its sympatry with its prim- itive relative P. choritonamus argues for its origin in a part of the Cumberland basin, perhaps the headwaters of the Cumberland in southeast Kentucky, not inhabited by the latter and a reinvasion of the homeland. From such a region, the invasion of the Tennessee basin, where P. hobbsi is widespread and successful, of the Big Sandy, and of the New River is entirely possible. There are, however, gaps in its known range, and other histories of the species are possible. Its absence from the central part of the Cumberland Plateau in Tennessee appears to be real, but further collecting can be expected NO. 3668 PTERODRILUS—HOLT 4] to connect the parts of its range that now appear to be disjunct. If so, the upper reaches of the Cumberland in Kentucky remain the likely site of origin for P. hobbsi. The few scattered records from the Big Sandy and the New Rivers indicate that the species is still ac- tively extending its range, and much of the spreading of P. hobbsi may well have occurred quite recently. The migrations of P. distichus, P. hobbsi, and P. alcicornus may have occurred rather recently, but the movement of the stock that gave rise to P. simondsi must be older. Although it is believed that a common ancestor gave rise to both P. simondsi and P. alcicornus, the former clearly stands at a lower level of evolutionary advance as is indicated by the primitive nature of its reproductive systems. It is known only from the tributaries to the Ocoee River and one locality in the Nottely, both parts of the Hiwassee River system of the Ten- nessee River basin. Pterodrilus simondsi is found, then, at the south- eastern periphery of the range of the genus in an isolated part of the somewhat isolated Hiwassee basin. Its ancestors came from the Cumberland and its known distribution can be explained by postu- lating that the species was once widespread in the Tennessee basin but has been eliminated throughout all of its range except the small part in the Hiwassee by the more advanced, successful, and wide- spread species, P. hobbsi and P. alcicornus. In any case, though the origins of P. simondsi may not be as ancient as those of the relict species in middle Tennessee and the Ozarks, it is an older relative of P. alcicornus holding out in a relict status in a part of the Tennessee basin not yet successfully invaded by the latter. If the hodbsi-like animals mentioned above (p. 32) are conspecific with other populations of P. hobbsi, the Hiwassee drainage is being invaded by this more advanced species, but if, as seems more likely, these specimens represent a survival of the primitive stock that gave rise to P. cedrus, we have at the periphery of the present range of the genus a relict of the first radiation within Pterodrilus. Pterodrilus alcicornus is the most advanced and successful species of the genus. It is a native of the New River basin that has in recent times extended its range, probably by stream captures, into the James and Roanoke basins to the east, into the Big Sandy to the north, and, amazingly, into the Savannah in the south. The latter invasion can only have occurred by means of the streams of the Tennessee system that lie between the headwaters of the New and the Savan- nah in western North Carolina, a region that has been inadequately sampled. Still earlier, P. alcicornus had moved into the upper reaches of the Tennessee River system in southwestern Virginia and north- eastern Tennessee, where it is sympatric with P. hobbsi, often occupy- 42 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 125 ing the same streams and presumably the same hosts. A cedrus-like stock that gave rise to P. distichus to the north and a more primitive member, P. simondsi, to the south, moved by way of former con- nections with the Cumberland into the New River basin to produce P. alcicornus. The history of the genus Pterodrilus is conceived in broad outline, then, to be like this: In early Miocene or pre-Miocene times a primi- tive stock of cambarincoloid branchiobdellids were epizoites carried by the progenitors of the modern crayfish fauna of the upland regions of eastern North America. These animals lived along the slopes of the present Appalachian uplift, represented today by the Cumberland Plateau and the Highland Rim, which was drained by a stream that corresponded to the present day Cumberland. From this center, early stocks moved into the Ozarks and perhaps on into Mexico with the progenitors of the Mexicanus Section of the crayfish genus Procambarus (Hobbs, 1967, pp. 13-15) and produced the species P. missouriensis and P. mexicanus. Pterodrilus choritonamus and P. cedrus are the survivors (and representatives of the two lineages produced) of this early diversification that remained in the area of their origin and P. hobbsi is a more advanced member of the choritonamus-mexicanus lineage that has not only remained in the Cumberland basin but has successfully invaded the Tennessee system and more recently the New River drainage. Some of the early members of the missouriensis- cedrus lineage have also moved eastward, with one and possibly two (P. simondsi and the unnamed animals) remaining today in the Hiwassee basin as relicts. This lineage also gave rise to the advanced species, P. alcicornus, in the New River basin. Moving to the north, most likely by way of the Kentucky River or a nearby stream, another branch of this lineage gave rise to P. distichus, which remained in the Kentucky region throughout the Pleistocene, and in Recent times has followed its crayfish hosts (primarily species of Orconectes) into the Great Lakes and the St. Lawrence basins. These migrations have left three regions in which primitive species remain today: the original home, the Cumberland basin; the Ozarks in the Missouri and Arkansas river systems (and possibly the eastern slopes of the Sierra Madre Oriental in Veracruz); and the Hiwassee basin along the southwestern slope of the Blue Ridge. 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