NEW RECORDS OF BRYOCYCLOPS FROM THE CONTINENTAL U.S.A., PUERTO RICO, AND BRAZIL (COPEPODA: CYCLOPOIDA: CYCLOPIDAE) Janet W. Reid

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1 NEW RECORDS OF BRYOCYCLOPS FROM THE CONTINENTAL U.S.A., PUERTO RICO, AND BRAZIL (COPEPODA: CYCLOPOIDA: CYCLOPIDAE) Janet W. Reid A B S T R A C T Collections from Brazil, Puerto Rico, and the U.S.A. yielded new records of 3 species of Bryocyclops. Both sexes of Bryocyclops muscicola are described from a probably introduced population found in a plant nursery in Florida, U.S.A., and compared to material of B. muscicola from Java. This is the first record of the genus in North America. The known distribution of Bryocyclops campaneri in Brazil is considerably expanded, and new information provided on its morphology. Bryocyclops caroli, previously known only from Brazil, is recorded for the first time from Puerto Rico. Species of the semiterrestrial cyclopoid copepod genus Bryocyclops have seldom been collected in the New World. This report presents new records from Brazil, Puerto Rico, and the U.S.A., and provides morphological details for three species. A population ascribed to Bryocyclop.s muscicola (Menzel, 1926) was found in soil in a plant nursery in Florida, U.S.A., by Paul S. Lehman of the Bureau of Nematology, Division of Plant Industry, Florida Department of Agriculture. Bryocyclops muscicola was originally described from Java and has since been reported only from Sumatra (Kiefer, 1933). Both the female and the male of the Florida population are described, and aspects of their morphology are compared with a specimen of B. muscicola from Java held in the Friedrich Kiefer Collection, Staatliches Museum fur Naturkunde, Karlsruhe, Germany. Comparisons are also made with B. caroli Bjornberg, 1985, B. campaneri Rocha and Bjornberg, 1987, and B. anninae (Menzel, 1926). Collections mainly from bromeliads in the Luquillo Mountains of Puerto Rico made by Bassett Maguire, Jr., and provided by Harry C. Yeatman included individuals of Bryocyclops caroli. Specimens of B. campaneri from northeastern Brazil sent by Sigrid Neumann- Leitao are briefly described. MATERIALS AND METHODS Specimens were preserved in 70% ethanol. Morphological observations were made on whole specimens in glycerine or lactic acid, and on dissected specimens mounted in CMC-10 or commercial polyvinyl lactophcnol with a little chlorazol black E added. The specimens provided by Harry Yeatman were mounted whole in glycerine jelly. Whole animals were drawn with the aid of a drawing tube mounted on a Wild M30 microscope, at magnifications of 400x or 600x. Dissected parts were drawn at 1,000x using an oil immersion lens. Details were verified using a Zeiss microscope set up for phase-contrast illumination. The material was deposited in the collections of the United States National Museum of Natural History, Smithsonian Institution (USNM). Order Cyclopoida G. 0. Sars, 1886 Family Cyclopidae Burmeister, 1834 Genus Bryocyclops Kiefer, 1927 Bryocyclops muscicola (Menzel, 1926) Figs. 1, 2 Cyclops mu.ccicola Menzel, 1926: 213, 215, figs Borutsky, 1927: 63.-Kiefer, 1928a: 526, Brehm, 1927: 482. (Not Cyclops muscicolus Lastochkin, 1924.) Brvoc.yclops muscieo/a.-kiefer, 1927: 306.-Kiefer, 1929: 75, 76.-Kiefer, 1933: 525, , 619, figs Kiefer, 1937: 439.-Lindberg, 1947: 49.-Thienemann, 1950: 254.-Lindberg, 1954: 104, 110, 111. Brvocvclops muscicola Group II. Lindberg, 1955: 6.- Lindberg, 1956: 76.-Dussart and Defaye, 1985: 142. Bryocyclopa (Bryocyclops) muscicola.-monchenko, 1972: 92.-Bjornberg, 1985: 237, 239, 241. Cyclops me zzeli, new name, Kiefer, 1926: 24, 25.-new name, Borutsky, 1927: 63.-Kiefer, 1928a: 526, 531, 551.-Dussart and Defaye, 1985: 142. Bryncyclop.s menzeli, new combination, Kiefer, 1927: 306, 307.-Kiefer, 1928a: Material Examined (2 completely dissected on slides) and 1 d (dissected on slide), from culture NCC- N , established in 1996 from specimens collected from organic soils of ornamental plants (plant host unknown) in a nursery at Zellwood, central Florida, U.S.A., P. S. Lehman, collector (USNM ). 1 adult 9 and 4 copepodids, Glas 0467, Java, Indonesia, R. Menzel, collector, Kiefer Collection, Staatliches Museum fur Naturkunde, Karlsruhe, Germany. The following description mainly refers to the Florida specimens. Bryocyclops muscicola was not listed as such in the Catalogue of the Kiefer Collection (Franke, 1989). Two lots from that collection labeled "Bryocyclops sp." were examined. A slide, Mikropraparat , from Java, collected by A. Thienemann, con- tains an adult female of Bryocyclops, but is dried. Ethanol-preserved specimens in Glas

2 0467, from Java, Indonesia, collected by R. Menzel, contains an adult female and four copepodids; the adult fits the descriptions of B. muscicola. The specimen was not dissected, but, although some setae are missing and the anal operculum is slightly damaged, its visible features agree with the Florida specimens, particularly with respect to the structure and setation of the antennule, antenna, legs 1-5, and the structure and proportions of the genital double-somite and caudal rami. Female.-Length of Florida specimens tm (median 430 pm, shorter specimens extremely telescoped). Length of Java specimen m (telescoped). Habitus (Fig. la) compact, dorsoventrally compressed. Sclerotized pseudosomite (Fig. 1 B, D) present between pediger 5 and genital double-somite. Cephalosome, pedigers 2-5, pseudosomite, and genital double-somite, except for surfaces of leg 6 plates and areas of genital doublesomite adjacent to leg 6 plates and around copulatory pore, covered with tiny, strongly refractile points. Cephalosome with transverse scar indicating fusion line of former cephalon and pediger 1. Cephalosome, pedigers 2 and 4, genital double-somite, and succeeding two urosomites with shallow grooves or pits on dorsal and lateral surfaces. Pedigers 2-4 each with paired dorsal pores and irregularly toothed hyaline frills, frill of pediger 3 especially wide. Pediger 5 with 2 transverse muscle scars, but paired dorsal hair-sensilla, usually present in family, not visible. Genital double-somite (Fig. 1 A, B, D-F) expanded laterally, approximately 1.5 times broader than long; seminal receptacle (Fig. I E) with broadly trapezoidal anterior expansion and narrow posterior expansion, complexly sclerotized pore-canal area, and narrow, nearly horizontal lateral canals. Spermatophores (Fig. 1 F) roughly bean-shaped. Anal somite (Fig. 1A, D, G) with spines along posterior margin; anal operculum ovate with irregularly toothed margin, produced posteriorly to approximately three-fourths length of caudal rami. Each caudal ramus about 1.5 times longer than wide, with pronounced longitudinal dorsal keel, ventrolateral pore at proximal one-fourth, tiny spine lateral to insertion of lateral seta, and row of larger spines along distolateral margin. Dorsal seta set on small prominence at distal end of keel. Middle and lateralmost terminal caudal setae stiff, fairly straight, and finely plumose. Lengths of caudal setae: lateral 21 im, dorsal 39 pm, medialmost to lateralmost terminal 15, 167, 106, and 32 11m. Rostrum (Fig. 1H) distinct from cephalothorax, broad and blunt, with pair of round pores and pair of tube-pores; its surface covered with tiny refractile points. Antennule (Fig. lea, H) of 11 segments, reaching approximately two-thirds length of cephalosome. Segment I with short row of tiny spinules; segment XI lacking hyaline membrane. Numbers of setae (s), spines (sp), and aesthetascs (ae) (Arabic numerals) per segment (Roman numerals): I(8s), II(3s), 1II(Ss), IV(3s), V(ls + 1 sp), VI(2s), VII(3s), VIlI(2s + 1 ae), IX(2s), X(2s + I ae), and XI(7s + lae). Antenna (Fig. 1I, J) of 5 segments. Segment 2 (basipodite) short, ornamented with few tiny spinules, and bearing only 1 seta on anterodistal corner, i.e., exopodite seta and 1 of 2 setae normally present on anterodistal corner lacking. Segments 3-5 (endopodite) with 1, 5, and 7 setae, respectively. Mandible (Fig. 1 K), palp represented by 1 short seta inserted directly on mandible. Maxillule (Fig. 1 L) of normal structure for family, i.e., palp lacking spinules on surface. Maxilla (Fig. 2A), claw set with tiny denticles along middle half of inner margin. Maxilliped (Fig. 2B) of 3 segments bearing 2, 1, and 3 setae. Legs 1-4 (Fig. 2C-F) with exopodites and endopodites 1-3 each composed of 2 segments, and endopodite of leg 4 of 1 segment. Margins of couplers (intercoxal sclerites) each with 2 large acute projections. Coxopodite of leg 1 with, but coxopodites of legs 2-4 without seta on distomedial corner. Leg 1 basipodite with seta on posterolateral surface and stout spine on medial expansion. Legs 2-4 basipodites with seta only on posterolateral surfaces. In fixed specimens, exopodites directed medially over endopodites (figures showing rami separated for clarity). Legs 1-4 exopodites, segment 1 with lateral spine and no medial seta; segment 2 with 3,3,3,3 spines and 5,5,5,4 setae, respectively. Two setae medial to lateralmost seta of legs 2 and 3 exopodites stout and blunt, with long plumage; all remaining setae of legs 1-4 slender, normally plumose. Legs 1-3 endopodites, segment 1 with medial seta; segment 2 with 2, 2, and 4 setae, respectively, inserted me-

3 Fig. 1. Brvocyclops muscicnla (Menzel, 1926), female, USNM A, habitus, dorsal; B, pediger 5, pseudosomite, and genital double-somite, right lateral; C, right leg 5, enlarged; D, pediger 5 and urosome, ventral (only integumental features indicated); E, genital double-somite, ventral, showing seminal receptacle; F, genital double-somite, ventral, with paired spermatophores; G, anal somite and caudal rami, dorsal; H, rostrum and antennule; I, antenna, frontal side; J, antenna basipodite, caudal side; K, mandible; L, maxillule; M, maxillular palp. Scales = 50 11m. dial to terminal spine, and 1 seta inserted lateral to terminal spine. Legs 2 and 3, distal segment of endopodites bearing accessory spinules along lateral margins. Leg 4 endopodite with original 2 segments completely fused, fusion line indicated on medial mar- gin by seta and on lateral margin by large spiniform process; distal margin with 1 short spine and 3 longer setae. Leg 5 (Fig. IB, C) consisting of 3 stiff setae, 2 more ventral setae of these set directly on posterolateral margin of pediger 5, dor-

4 Fig. 2. Bryocyclops mu.scicola (Menzel, 1926), A-F, female; G-J, male, USNM A, maxilla; B, maxilliped; C, leg 1 and coupler; D, leg 2 and coupler; E, leg 3 and coupler; F, leg 4 and coupler; G, genital somite and succeeding urosomites, dorsal; H, pediger 5, genital somite, and succeeding urosomite, left lateral; 1, leg 3 endopodite; J, leg 4 endopodite. Scales = 50 wm. salmost seta set on small prominence. Ventralmost seta longest, middle seta shortest. Leg 6 (Fig. 1A, B) consisting of large subtrapezoidal plate bearing slender seta and 2 tiny spines on dorsal margin. No egg sacs present on specimens examined. Male.-Length 360 Jlm (specimen somewhat telescoped). Habitus generally similar to female, except for normal sexual dimorphisms. Prosome, pediger 5, and genital segment (Fig. 2G, H) covered with strongly refractile points, next 2 urosomites with few, scattered points. Anal operculum (Fig. 2G) shorter and more rounded than in female. Caudal ramus (Fig. 2G) as in female except lacking dorsal keel. Good mount of antennule not obtained; antennule geniculate, resembling that of B. campaneri Rocha and Bjornberg, 1987, especially in possessing long slender aesthetascs.

5 Legs 1 and 2 as in female. Leg 3 (Fig. 21), exopodite as in female, endopodite with setae somewhat longer than in female; terminal spine modified, with subterminal, bulbous, finely serrate expansion, and acute, hooked end; and seta medial to terminal spine directed under spine (structures confirmed on several undissected specimens). Leg 4 (Fig. 2J), exopodite as in female, endopodite composed of 2 segments, segment 1 with spiniform expansions on medial and lateral distal corners, and on segment 2, setae relatively longer than in female. Leg 5 (Fig. 2H) as in female. Leg 6 (Fig. 2G, H) consisting of large trapezoidal plate (without refractile points) bearing 3 setae, ventralmost being longest and dorsalmost shortest. Color.-Living specimens colorless; eye red. Comparisons.-Bryocyclops muscicola was first discovered in damp moss on rocks at the base of a small stream in the virgin forest of Gedeh at Tapos near Buitenzorg (now Bogor), Java, at an altitude of 1,000 m (Menzel, 1926). The only subsequent collection in Asia was from leaf axils of Pandanus, near Lake Ranau in southern Sumatra (Kiefer, 1933). Menzel (1926) and Kiefer (1933) provided only partial descriptions, and only of the female. Neither Bryocyclops muscicola nor B. menzeli (a new name proposed by Kiefer, 1926, and Borutsky, 1927, which was unnecessary) is listed in the catalogue of the Kiefer Collection (Franke, 1989). The females from Java and Florida are congruent with the descriptions of both Menzel (1926) and Kiefer (1933), particularly with respect to the spination and setation of the swimming legs, and the single-segmented leg 4 endopodite with a prominent spiniform process on its lateral margin (visible in fig. 12 of Menzel, 1926, and less clearly in fig. 127 of Kiefer, 1933). The specimens are easily identified as B. muscicola in the key of Monchenko (1972). Bryocyclops mu,scicola is clearly a member of the subgenus Bryocyclops sensu Dussart (1982), in that it possesses a seta and spine on the leg 1 coxopodite and basipodite, the spine formula 3,3,3,3, and acute processes on the leg 4 coupler. The new information on the morphology of the male confirms that it belongs to Group II of Lindberg (1955, 1956), as previously supposed by Lindberg (1955, 1956) and Dussart (1982). Bryocyclops campaneri Rocha and Bjornberg, 1987 Fig. 3A-E Material Examined.-2 2.'?, I dissected on slide, from estuary of Rio Ipojuca near Suape, State of Pernambuco, Brazil, approximately 8 24'S, 34 59'W, 1986, S. Neumann-Leit5o, collector (USNM ). Female.-Lengths 352, 360 pm. Habitus (Fig. 3A) much as B. muscicola, except dorsal transverse hyaline frills on pedigers 2-4 located slightly more anteriorly, and refractile points smaller, more difficult to see, and more widely spaced than in B. muscicola. Scar at fusion line of cephalon and pediger 1 represented only by longitudinal lateral remnants. Distribution of somitic pores as in B. muscicola. No pseudosomite visible between pediger 5 and genital double-somite on either specimen. Anal operculum (Fig. 3A, B) reaching tips of caudal rami. Antenna (Fig. 3C) with 2 setae on distal corner of basipodite, and 6 setae on segment 2 of endopodite. Exopodite segment 2 of leg 2 (Fig. 3D) and of leg 3 with 2 stout blunt setae, like those in B. muscicola except with short stiff plumage. Leg 4 (Fig. 3E), exopodite segment 1 somewhat lobate; endopodite with convex margins and large rounded protrusion lateral to seta of medial margin. Comments.-Bryocyclops campaneri, originally described from the coastal Rio Una do Prelado in the Jureia Ecological Reserve, State of Sao Paulo, has also been recorded from oligohaline waters of the coastal Rio Piauf, near the town of Estancia, State of Sergipe, northeastern Brazil (C. E. F. Rocha, personal communication, 1994). In Pernambuco, B. campaneri was collected by Neumann-Leitao (1994) from June through October 1986, at a station in the oligohaline zone of the Rio Ipojuca, where the salinity varied from 0.05%c to about 32%c depending on the tide. Thus, this species is apparently able to tolerate a wide salinity range in the tidal zones of the small, slowly flowing coastal rivers. Its geographical distribution in Sao Paulo, Sergipe, and Pernambuco spans some 2,000 km and 16 degrees of latitude. C. E. F. Rocha (personal communication, 1994) confirmed the presence of refractile points on the integument of the holotype of B. campaneri, and other details conforming

6 Fig. 3. A-E, Bryncyclops cumpuneri Rocha and Bjornberg, 1987, female from Pernambuco, Brazil, USNM A, habitus, dorsal; B, anal somite and caudal rami, dorsal; C, antenna; D, leg 2 coxa-basipodite and exopodite, caudal; E, leg 4 and coupler, frontal. F-H, Bryncyclnps unninae (Menzel, 1926), female from Guam, USNM F, habitus, dorsal; G, anal somite and caudal rami, dorsal; H, leg 4. Scales = 50.1m. to the specimens from Pernambuco. He noted that the hyaline frill on pediger 4 is not obvious in the holotype. Rocha and Bjornberg (1987) noted only one seta on the antenna basipodite and 5 setae on the antenna endopodite segment 2. The same authors clearly illustrated but did not comment on the stout, blunt setae on the exopodites of legs 2 and 3, nor did they mention or illustrate a lobe on the leg 4 endopodite. The anal operculum of the Pernambuco specimens is much more produced posteriorly than in the Sao Paulo animals, in which it reaches only the level of the lateral caudal setae.

7 Bryocyclops anninae (Menzel, 1926) Fig. 3F-H Material Examined.-I 1 9, ethanol-preserved, examined while temporarily mounted whole in lactic acid, from Mangilao, Guam, 8 March 1971, D. Belk, collector (USNM ). Female.-Habitus (Fig. 3F) much as in M. muscicola, with similar density and distribution of refractile points on prosome and anterior urosomites, but lacking integumental structure indicating fusion of cephalon and pediger 1. No somitic pores or hairs, or transverse hyaline membranes visible on prosomites. Leg 6 plate with conspicuous knob near medial margin. Anal operculum (Fig. 3F, G) irregularly acuminate, with free margin smooth. Caudal ramus with longitudinal dorsal keel little developed, no spines at base of lateral seta, and dorsal seta plumose, otherwise much as in M. muscicola. Legs 2 and 3 with blunt setae on exopodites, like those of B. muscicola; leg 4 exopodite (Fig. 3H) also with 2 blunt setae. Bryocyclop.s caroli Bjornberg, 1985 Material Examined.-4,'t? and 1 J mounted whole on 1 slide, Sample B-4, from Guzmania sp., west of EI Verde experimental station, Puerto Rico, 18 May 1965 (USNM ); B. Maguire, Jr., collector, H. C. Yeatman, donor. Comments.-As far as can be seen from the whole specimens, the specimens agree in every detail with the description of B. caroli by Bjornberg (1985). Refractile points are visible on the prosome, pediger 5, and genital double-somite, but not on the remaining urosomites. These and specimens of Fimbricyclops jimhensoni Reid, 1993, as well as a previously unknown species of Fimbricyclops, were present in the series of slides from Puerto Rico provided by H. C. Yeatman. These individuals were originally reported as Bryocyclops chappuisi Kiefer, 1928b, and B. anninae by Maguire (1970), following identifications by H. C. Yeatman. Bryocyclops chappuisi and B. anninae, both Asian species, have not been recorded elsewhere in the New World. DISCUSSION The confused systematics of the genus Bryocyclops have been discussed by Lindberg (1956), Monchenko (1972), Dussart (1982), Rocha and Bjornberg (1987), and Rocha et al. (1998). Rocha et al. (1998) proposed that the subgenus Haplocyclops Kiefer, 1952, be raised to genus rank. The members of Group II were defined by Lindberg (1954) as having: the leg 3 endopodite segment 2 terminal spine different in male and female; the leg 4 coupler with acute or rounded protrusions; spine formula 3,3,3,3 and seta formula 5,5,5,4; and the leg 4 endopodite composed of 1 segment, bearing 1 spine and 4 setae. Group II includes B. muscicola; B. bogoriensis (Menzel, 1926), known from Java, Sumatra, and Bali; and B. caroli, from Brazil and Puerto Rico. Bryocyclops fidjiensis Lindberg, 1955, from Fiji, was assigned by its author to Group II. However, according to observations by Yeatman (1983) of many specimens of B. fidjiensi.s from the type locality, the female of this species actually has the leg 4 exopodite composed of only 1 segment (resulting in the spine formula 3,3,3,4) and the leg 4 endopodite always of 2 segments. The other species known from Brazil, B. campaneri, closely resembles the species of Group II except in having the seta formula 4,5,5,4. Bryocyclops bogoriensis differs from B. muscicola, according to the redescription by Kiefer (1928b), in lacking a seta on the medial corner of the coxa in all the swimming legs; in having rounded rather than acute projections on the couplers; in the female, having a small spiniform process on leg 4 endopodite; and in the male, lacking a spiniform process on the distomedial comer of the leg 4 endopodite 1. Bryocyclops caroli is now shown to be extremely similar to B. muscicola, separable only by its obvious dorsal hyaline frill on pediger 3, the lack of a large spiniform process on the leg 4 endopodite of the female, and by the thick blunt tip of the modified spine of leg 3 endopodite of the male (illustrated both by Bjornberg, 1985, and by Rocha and Bjornberg, 1987). Both B. caroli and B. campaneri share with B. muscicola the thick blunt setae of the exopodites of legs 2 and 3, as described herein, and illustrated for the type populations of the former two species by their respective authors. In B. muscicola, the seta medial to the modified spine on the leg 3 endopodite of the male is as long as the spine and curves rather gently beneath it, as opposed to the situation in the other species where this seta is straight and stiff, less than half the length of the spine, and passes beneath it at about a 45 angle.

8 The relative length and shape of the anal operculum have been treated as species discriminant characters by Monchenko (1972) and others. The length of the anal operculum is apparently a somewhat variable character, at least in different populations of B. campaneri. The degree of development of the anal operculum should be employed as a species discriminant character only with due caution. The geographical distribution of the members of Bryncyclop.s Group II in Indonesia and Brazil is rather peculiar. Moreover, species of Bryocyclops in general occur predominantly in the Old World and on islands in the Pacific Ocean (reviewed by Bjornberg, 1985). The only New World species now known are B. campaneri in Brazil, B. caroli in Brazil and Puerto Rico, and the probably introduced population of B. muscicola in Florida. There are few descriptions of ornamentation on the somites of species of Bryocyclops. Lindberg (1947: 47) noted that on the ventral surface of the genital double-somite of the female of B. constrictus Lindberg, 1947, there was "une ornamentation de la cuticule d'epinules disposees en lignes sinueuses." Defaye and Heymer (1996) observed a transverse scar or thickening on the cephalothorax of B. phyllopus Kiefer, 1939, similar to the structure at the same location on B. muscicola, and thicker than the slight remnant of such a scar on the cephalothorax of B. campaneri. Such a structure has not been observed in other members of the subfamily Cyclopinae. Transverse dorsal hyaline frills, such as those on pediger 3 in B. caroli, on pedigers 2, 3, and sometimes 4 in B. campaneri, and on pedigers 2-4 in B. muscicola have been described elsewhere only for B. phyllopus by Defaye and Heymer (1996). Por (1981), in his description of B. absalomi, observed that the cuticle was "granulated in transparence (not superficially)", and no surface pores or granulations were apparent in the SEM photographs that he provided. Defaye and Heymer (1996) described the integument of "tout le corps" of both sexes of B. phyllopus as "tres finement pointille." These descriptions apparently refer to the same structure as the cuticular re- fractile points of B. muscicola, B. campaneri, and B. anninae. The presence of similar refractile points may account for the "shiny" appearance of live specimens of B. fidjiensis, as reported by Yeatman (1983). As Monchenko (1972, and personal com- munication, 1997) noted, the genus diagnosis of Bryocyclop.s has suffered continuous erosion with the discovery of morphological features, once thought to be defining, in other related genera. It is possible that cuticular refractile points may be a general feature in species of Bryocyclops, because they occur in all species examined for the present report, and apparently in several other species. This is not to suggest that the possession of cuticular refractile points is a unique apomorphy in Bryocyclops, since apparently similar structures have been described for Halicyclops maculatus by Rocha and Hakenkamp (1993). However, this cuticular structure may be an apomorphy separating Bryocyclops from related genera that lack a similar modification, such as Fimbricyclops, Stolonicyclops Reid and Spooner, 1998, and Haplocyclop.s (cf. Rocha et al., 1998). The modified blunt setae of some swimming leg exopodites, which were also observed in all species examined, are found in very few cyclopoid species other than members of Bryocyclop.s. Improved knowledge of the distribution of heretofore nearly ignored details of the integument, mouthparts, and swimming leg setae within Bryocyclops and related genera cannot but result in improvements in the concept of the genus. The existence of a pseudosomite between pediger 5 and the genital double-somite was first discovered in Diacyclops biceri by Boxshall et al. (1993), who suggested that it might increase body flexibility. A similar pseudosomite occurs in Stolonicyclops heggiensi.s Reid and Spooner, A pseudosomite may be a common feature in soildwelling and interstitial cyclopoids. ACKNOWLEDGEMENTS I am grateful to Drs. Denton Belk, Paul S. Lehman, Sigrid Neumann-Leitao, Carlos Eduardo F. Rocha, and Harry C. Yeatman for their gifts of specimens to the collections of the National Museum of Natural History, and to Drs. Oliver Coleman, Hans-Uwe Dahms, and Hans- Walter Mittmann for their good offices in obtaining material from the Staatliches Museum fur Naturkunde, Karlsruhe, Germany. Drs. Wolfgang Janetzky, Vladislav I. Monchenko, Carlos Rocha, and Hiroshi Ueda provided valuable information and commentary. The Department of Invertebrate Zoology, NMNH, afforded me research facilities, not least the use of the C. B. Wilson Copepod Library. LITERATURE CITED Bjornberg, M. H. G. C Bryocyclops caroli sp. n. (Crustacea, Copepoda, Cyclopoida), the first representative of the genus in South America.â Hydrobiologia 124:

9 Borutsky, E. V Cyclops muscicola Menzel and Cyclops muscicolus Lastotschkin.-Zoologischer Anzeiger 71: 63. Boxshall, G. A., T. D. Evstigneeva, and P. F. Clark A new interstitial cyclopoid copepod from a sandy beach on the western shore of Lake Baikal, Siberia.â Hydrobiologia 268: Brehm, V Copepoda.â In: W. Kükenthal and T. Krumbach, eds., Handbuch der Zoologie 3: Burmeister, H Beitrà ge zur Naturgeschichte der Rankenfüsser (Cirripedia).-G. Reimer, Berlin. Pp Defaye, D., and A. Heymer Crustacà s Copà podes de litiere de la foret ombrophile du Kivu (Zaà re).â Bulletin du Museum national d'histoire naturelle, Paris, 4e Sà rie, 18 (Section A, 1-2): Dussart, B. H Crustaces copepodes des eaux intã rieures.â Faune de Madagascar 58: , and D. Defaye Repertoire mondial des Crustaces Copepodes. II. Cyclopoà des.â à ditions du Centre National de la Recherche Scientifique, Bordeaux. Pp Franke, U Katalog zur Sammlung limnischer Copepoden von Prof. Dr. Friedrich Kiefer.â Carolinea 5: Kiefer, F Beitrà ge zur Copepodenkunde. (IV). 9. Neue Cyclops-Arten.â Zoologischer Anzeiger 69: Versuch eines Systems der Cyclopiden.â Zoologischer Anzeiger 73: â â â. 1928a. Uber Morphologie und Systematik der Süà wasser-cyclopiden.â Zoologische Jahrbücher, Abteilung fur Systematik 54: b. Beitrage zur Copepodenkunde. (IX). 19. à ber drei Bryocyclops-Arten aus Java.-Zoologischer Anzeiger 76: Crustacea Copepoda. 2. Cyclopoida Gnathostoma.-Tierreich 53: Die freilebenden Copepoden der Binnengewasser von Insulinde.â Archiv fã¼r Hydrobiologie, Supplementband 12, Tropische Binnengewà sser, 4: â â â à ber Systematik und geographische Verbreitung einiger Gruppen stark verkã¼mmerten Cyclopiden (Crustacea, Copepoda).-Zoologische Jahrbucher, Abteilung fur Systematik 70: Crustacea. IV. Copepoda: Diaptomidae, Cyclopidae.-Mission Scientifique de l'omo, V (Zoologie). Memoires du Museum National d'histoire Naturelle, Paris, Nouvelle Sà rie, 9: Haplocyclops gudrunae n. g. et n. sp., ein neuer RuderfuBkrebs (Crustacea Copepoda) aus Madagascar.â Zoologischer Anzeiger 149: Lastochkin, D. A Novye i redkie Copepoda and Oligochaeta v faune Ivanovo-Voznesenskoi gubernii. [New and rare Copepoda and Oligochaeta in the fauna of Ivan-Voznesensky Province.]-Izvestiya Rossiiskoi Gidrologicheskogo Instituta 9: [In Russian; summary in English.] Lindberg, K Cyclopoà des (Crustaces copepodes) nouveaux de l'inde. I. Description des premiers Bryocyclops decouverts dans l'inde.â Records of the Indian Museum 45: Un Cyclopide (Crustace copepode) troglobie de Madagascar. Avec remarques sur un groupe de Cyclopides tres evolues, cavernicoles et muscicoles.-hydrobiologia 6: â â â Cyclopides (Crustacà s copã podes) d'iles du Pacifique Sud (Mà lanã sie et Micronà sie) et de Bornà o.â Kunglige Fysiografiska Sallskapets i Lund Forhandlingar 24: â â â Les Cyclopides (Crustaces Copepodes) tres evolues en tant qu'habitants des eaux souterraines. Revue des travaux recents concernant les Bryocyclops Kiefer et Speocyclops Kiefer.-Actes du Premier Congres International de Speleologie, Paris, 1953, 3: Maguire, Jr., B Aquatic communities in bromeliad leaf axils and the influence of radiation.â In: H. T. Odum and R. F. Pigeon, eds., A tropical rain forest: a study of irradiation and ecology at El Verde, Puerto Rico. United States Atomic Energy Commission: E.95-E.101. Menzel, R Cyclopides muscicoles et bromã licoles de Java (Indes Nà erlandaises).â Annales de Biologie Lacustre 14: Monchenko, V. I Tsiklopy (Copepoda, Cyclopidae) gruntovykh vod pustyni Kyzylkum. [Subterranean water cyclops (Copepoda, Cyclopidae) from Kisilkum.]â Trudy Zoologicheskogo Instituta 51: [In Russian; summary in English.] Neumann-Leitao, S Impactos antropicos na comunidade zooplanctã nica estuarina, Porto de Suapeâ PEâ Brasil.â Tese de Doutorado, Escola de Engenharia de Sao Carlos, Universidade de Sao Paulo, Sao Carlos, Brazil. Pp Por, F. D A new species of Bryocyclops (Copepoda: Cyclopoida) and of Parastenocaris (Copepoda: Harpacticoida) from a cave in Israel and some comments on the origin of the cavernicolous copepods.â Israel Journal of Zoology 30: Reid, J. W Fimbricyclops jimhensoni, new genus, new species (Copepoda: Cyclopoida: Cyclopidae), from bromeliads in Puerto Rico.â Journal of Crustacean Bi- ology 13: â â â. and J. D. Spooner Stolonicyclops heggiensis, new genus, new species, from Georgia, U.S.A. -Journal of Crustacean Biology 18: 405â 411. Rocha, C. E. F., and M. H. G. C. Bjornberg Copepods of the Jureia Ecological Reserve, State of Sao Paulo, Brazil. II. The genera Hesperocyclops, Muscocyclops, and Bryocyclops (Cyclopoida, Cyclopidae).â Hydrobiologia 153: â â â, and C. C. Hakenkamp New species of Halicyclops (Copepoda Cyclopidae) from the United States of America.-Hydrobiologia 259: â â â, I. C. Torres, and P. M. Maia-Barbosa Huplocyclops torresi n. sp. and Potamocaris estevesi Reid, 1991 from Brazil, with a proposal for revalidation of the genus Haplocyclops Kiefer, 1952 (Copepoda).-Beaufortia 48: Sars, G. O Crustacea, II.â Norwegian North Atlantic Expedition , Zoology 6: Thienemann, A Verbreitungsgeschichte der Süà - wassertierwelt Europas.â Binnengewà sser 18: Yeatman, H. C Copepods from microhabitats in Fiji, Western Samoa, and Tonga.-Micronesica 19: RECEIVED: 19 July ACCEPTED: 13 March Address: Department of Invertebrate Zoology/MRC- 163, National Museum of Natural History, Smithsonian Institution, Washington, D.C , U.S.A. ( reid.janet@nmnh.si.edu)