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Pawson Department of Invertebrate Zoology, Smithsonian Institution Washington, DXJ. 20560 In a recent article on the echinoderms of British Honduras ( Belize ) , Devaney ( 1974 ) described a fauna ot 34 species, of which only one, the synaptid Euapta lappa (MuUer), was a holothurian. During the Smithsonian Institutions In- vestigations of Marine Shallow Water Ecosystems (I.M.S.W.E.) Project, based largely at Carrie Bow Cay, Behze (16°48'N, 88°05'W), participating investigators have made general collections of invertebrates, and these include 12 species of holothurians, of which 3 species of the genus Leptosynapta are new. In this paper the new species are described, other species are briefly characterized and some illustrated, and a key to all species is pro\'ided. A brief de- scription of Carrie Bow Cay habitats can be found in Dahl (1973). All specimens wc>re collected from Carrie Bow Cay, unless otherwise indicated in the species accounts. They were sampled by wading and skin diving in less than 2 m water depth. Kier (1975) described the echinoids of Carrie Bow Cay, and added a further 13 species to the 7 listed by De\'aney. Thus, the total number of echinoderms now reported from Belize is 58. As is to be expected, the fauna is typically Caribbean in character. For the holothurians some interest- ing new range extensions are reported. No dendrochirotid ' Contribution \o. 30 of the Investigations of Marine Shallow Water Ecosystems (I.M.S.W.E.) Project, Smithsonian Institution. 31— Proc. Biol. Soc. Wash., Vol. 89, 1976 (369) 370 Proceedings of the Biological Society of Washington ( "cucumaria"-type ) species have been collected at Belize. In several other Caribbean areas the dendrochirotids may comprise up to 30% of the holothurian fauna. This hiatus is inexplicable, but may be related in some way to the reduced vagility of the dendrochirotids (which lack a pelagic larval stage), or to the fact that more collecting needs to be done in areas with hard substrates. I wish to thank the coordinator of the I.M.S.W.E. Program, Dr. K. Ructzler, for making the material available for me, and Mr. K. Sandved for photographs. I am grateful to the several individuals named in the text who collected the holo- thurians described here. The I.M.S.W.E. Program has re- ceived support from the Smithsonian Institution and the Exxon Corporation. Key to Holothurians Known from Belize 1 ( 12 ) Body wall usually thick. Tentacles 20. Tube feet present. 2 ( 3 ) Anus surrounded by 5 calcified "teeth" Actinopijga agassizii ( Selenka ) 3 ( 2 ) No such teeth present. 4 (5) Burrowing form, body fusiform, with inconspicuous feet. Light brown with dark brown blotches dorsally Holothiiria arenicola Semper 5 (4) Not burrowing fomis, feet more or less conspicuous. 6 (11) Tube feet placed upon conspicuous warts, at least dorsally. 7 ( 8 ) Skin thin, rough to touch. Grey mottled with brown Holothuiia impatiens (Forskaal) 8 (7) Skin thick, more or less smooth. Light to dark brown, mottled. Ossicles in body wall tables and C-shaped bodies. 9 (10) C-shaped bodies (when present) approximately as long as tables are high Isostichopus badionotus (Selenka) C-shaped bodies approximately twice as long as tables are high Isostichopus macroparentheses ( Clark ) Warts absent. Skin very thick, smooth, rigid. Dark brown dorsally, yellowish laterally and reddish to pink ven- trally Holothuiia mexicana Ludwig Body wall thin, body worm-like, tentacles less than 20. Tube feet absent. Skin sticky to touch, due to presence of projecting ossicles in form of anchors supported by anchor plates. No wheels in body wall. 14 (17) Arms of anchors smooth; vertex widi minute knobs. Anchor 10 (9) 11 (6) 12 (1) 13 (22 Seo cucumbers from Belize 371 plates with well developed bridge for support of anchors. Active, conspiciunis fomis; generally non-burrowing. 15 (16) Grey to brown, often longitudinally striped. Body wall often with characteristic hemispherical "bumps." Stock of anchor branched Eiiapta lappa (Miiller) 16 (15) Green and white mottled or brown and white mottled. No "bumps" on body wall. Stock of anchor unbranched ____ Synaptula hijdnformis (Lesueur) 17 ( 14 ) Arms of anchors with teeth; no knobs on vertex. Anchor plates lack true bridge. Inconspicuous burrowing forms. 18 ( 19 ) Anchors and anchor plates very large, anchors usually ex- ceeding 600 /iui in length, plates usually exceeding 400 ^m Leptosijnapta imswe Pawson 19 (18) Anchors and anchor plates do not exceed 200 /tm in length. 20 (21) Anchors and anchor plates very small, considerably shorter than 130 ^m. Miliary granules resemble rosettes Leptosynapta nannoplax Pawson 21 (20) Anchors and anchor plates larger, usually exceeding 140 jxm in length. Miliary granules more or less C-shaped, with enlarged ends, not resembling rosettes Leptosynapta roseogradia Pawson 22 ( 13 ) Skin more or less smooth, but with numerous scattered papillae containing aggregations of wheels. Brick red to pink Cliiridota rotifera ( Pourtales ) Actinopyga agassizii (Selenka) Figures IC, E Actinopyga agassizii. — Deichmann, 1930, 78, pi. 5, figs. 21-29. Material examined: Lagoon, west of island, coral sand and turtle grass, 19 April 1972, collected R. J. Larson, 1 specimen; lagocMi, April 1974, 1 specimen; Coral berm, 3 May 1974, collected by B. Spracklin, 1 specimen; back reef, 1-2 m, April 1975, collected by M. Carpenter, 2 specimens. Remarks: Grows to about 20 cm. Five conspicuous calcareous "teeth" surround the anus. Skin thick, leathery. Numerous tube feet dorsally and ventrally. Color variable, but usually with mottled brown and yellow predominating. Tentacles yellow. Common on sand in grassy areas in shallow water around islands of the West Indies, from Barjjados to Florida. One record from Ber- muda. The record from Belize is a new range extension into the Western Caribbean. Holothuria impatiens ( Forskaal ) Holotliuria impatiens. — Deichmann, 1930:64, pi. 3, figs. 17, 18. Material examined: Just inside reef crest, 7 May 1974, collected by F. H. C. Hotchkiss and K. Sand\ed, 2 specimens; lagoon, west side 372 Proceed iiiiis of the Bio]o<^ical Society of Washington Fic. 1. A, Holothuria mexicana, ventral; B, Ilolothuiia arenicola, dorsal; C, Actinopyga agassizii, dorsal; D, Isostidwpus niacioparen- tlieses, anterior dorsal; E, juvenile Actinopyga agassizii, ventral. of island, coral sand, turtle grass, under conch shells, 19 April 1972, collected by R. J. Larson, 1 specimen. Remarks: Grows to about 20 cm. Tube feet few, scattered, placed on distinct warts, at least dorsally. Skin thin, rough to touch. Color grey with brownish patches. A "tropicopolitan" species, often imcommon or rare where it occurs. Usually found on sand or in grassy areas. Holotluiria arciiicola Semper Figure IB Uolotluiiia au'iiicola. — Deichmann, 1930:66, pi. 4, figs. 1-9. Material examined: Under conch shells, foot of pier, 29 April 1974, collected !)>■ V. II. C. Hotchkiss and K. Sandved, 1 specimen; lagoon, Sea cucumbers from Belize 373 west side of island, near shore, 1 May 1974, collected by F. H. C. Hotchkiss and K. Sandved, 1 specimen; lagoon along west shore of island, low tide, 30 April 1972, collected by R. J. Larson, 1 specimen; back reef, 1-2 m, 1975, collected by K. Sandved, 1 specimen; same locality, 22 April 1975, collected by M. Carpenter, 2 specimens; same locality, 27 April 1975, collected by M. Carpenter, 1 specimen. Rc'Diarks: Grows to about 30 cm. Body slender, fusiform, with veiy small tentacles. Adapted to a burrowing habit, and often found con- cealed under rocks and shells. Skin relatively thin and smooth to the touch. Groimd color usually light brown, with dark brown patches in 2 series on dorsal smface. Color can vary considerably, probably de- pending upon chemical properties of the habitat; rust-colored to al- most black specimens occur in some areas of the Caribbean. A "tropicopolitan" species, commonly encountered in suitable habi- tats. Ilolotlitiria mexicana Ludwig Figmes lA, 2A Holotliuiia )>icxicana. — Deichniann, 1930:74, pi. 5, figs. 15-20. Material examined: Lagoon, south end of island, 1 specimen; May 1974, collected by F. H. C. Hotchkiss, 1 fragment; back reef, im- mediately behind crest, 1-2 m, April 1975, collected by M. Carpenter, 2 specimens. Remarks: Grows to about 50 cm. Skin very thick, smooth, extremely hard when contracted. In life, body dark brown to blackisli dorsally, with flanks yellowish brown, and ventral surface frequently reddish or pink. The reddish to pink ventral coloration disappears in alcohol. The specimen from station 1 above is unusual in being very light brown dorsally. The smaller specimen from the back reef is 110 mm in total length. The ground color is greyish white; ventral feet are light brown with dark brown endplates; dorsal surface with light brown feet and a double row of 4 large dark brown blotches. Deich- mann (1930:74) noted that young specimens of this species have similar coloration, but she did not mention the presence of large dark blotches dorsally. Ranges from Cuba to Curacao in shallow water (to 20 m) in grass\' areas or on muddy sand. This species has been frequently confused with the closely related H. floridana Pourtales. Isosticliopus Jnuliouotus (Selenka) StieIioj)tis Ijadionotus. — Deichniann, 1930:80, pi. 5, figs. 30-36. Material examined: Lagoon, west side of South Water Cay, mixed sand and Thalassia, 1 m depth, 4 May 1974, collected by F. H. C. Hotchkiss and K. Sandved, 1 specimen; Coco-Plum Cay, at edge of mangrove swamp, 2 May 1972, collected by R. J- Larson, 1 specimen; 374 Proceedings of the Biological Society of Washington lagodn, west side of island, coral sand, turtle grass, under conch shells, 19 April 1972, collected by R. ]. Larson, 1 specimen. Remarks: Grows to about 20 cm. Skin thick, with low warts dorsally and 3 crowded rows of tube feet ventrally. Color highly variable, from light brown to black, usually with numerous large spots or blotches of dark brown on a lighter background. Common on nuidd\' sand or in grassy areas, in shallow water, from Bermuda to Panama. I.sosficJwpus macroparentheses (Clark) Figure ID Sficliopus iiuiciopiiiciitlicscs II. L. Clark, 1922:61, pi. 4, figs. 1-7; Deichmann, 1930:82, pi. 5, figs. 37-43; H. L. Clark, 1933:110 [macrapaicntJicscs I . Iso.sticliopiis IxidiDiiotus. — Deichmami, 19fi3: 106. Material cxaniinecl: Back reef, 1-2 m, April 1975, collected by M. Carpenter, 1 specimen. Reniarks: Grows to about 12 cm. In tlie field, \irtually indistinguish- able from 7. badionotus, and can be positively identified only after examination of the spicules. Deichmann ( 1963 ) was inclined to re- gard I. macroparentheses as a juvenile stage of 7. hadionotus which has exceptionally large C-shaped ossicles. The difference between these C-shaped ossicles in tlie two forms is dramatic, particularly when one compares jmeniles of 7. hadionotus with 7. macroparentheses. I can- not agree with Deichmann's contention, and thus prefer to retain 7. macroparentheses as a separate species. In life, "blight brown, with very dark rings around the bases of the papillae which ha\e yellow tips" (Clark, 1933:110). The present specimen is 30 nun long. Color photograph shows light brown body with dark brown dorsal feet and light to dark brown ventral feet; tentacles are more or less colorless. The species i.^ also known from Antigua, Jamaica and the Tor- tugas, in .shallow water. Euapta lappa (Mullcr) Figure 2B Euapta lappa.— U. L. Clark, 1924:464, pi. 1, figs. 5-7; 1933:118. Material examined: Lagoon, 2 specimens; just inside reef crest, 6 May 1974, collected by F. H. C. Hotchkiss and K. Sandved, 1 speci- men; reef flat, south end, east side of island "Penicillus" rock zone, 1 specimen; lagoon, west side of island, coral sand, turtle grass, under conch shells, 19 April 1972, collected by R. J. Larson, 2 specimens. Remarks: Grows to about 100 cm. Color grey to brown, often longitudinally striped. Body wall prickly to touch, owing to presence Sea cucumbers from Belize 375 Fig. 2. A, young HoIotJuiria inexicaua, lateral; B, Ettapta lappa, anterior dorsal; C, Synaptiila liijdiiformis, dorsal. of projecting spicules ( anchors ) . Tentacles phnne-like. Body often displays rows of characteristic conspicuous hemispherical protuberances. An active reef-dwelling form often associated with Kuiips of dead coral. Can swim to a limited extent. Ranges the entire Caribbean in shallow water. Si/iuipfnia h[idriformis Figure 2C Lcsuem- Synaptiila liijdiifotmis.—H. L. Clark, 1924:473, pi. 3, figs. 5, 6, pi. 4, fig. 4; 1933:119. Material examined: Foot of boat pier, bulk sample no. 2, 29 April 1974, collected by F. H. C. Hotchkiss and K. Sandved, 8 specimens; just inside reef crest, 6 May 1974, collected by F. II. C. Hotchkiss and K. Sandved, 1 specimen; back reef, 1-2 m, April 1975. col- lected by K. Sandved, 1 specimen. Remarks: Grows to about 10 cm. Prickly to touch due to projecting spicules ( anchors ) . Two color phases, mottled green and white and mottled brown and white, can occur. Some correlation exists between habitat type (red or brown or green algae frequently) and color of body wall, but exceptions occur; for example, in Bermuda the bidwn fomi is conspicuous among clumps of the green Penicillus. Green form only known so far from Carrie Bow Cay. Viviparous; apparently can breed all year round in Bermuda. Ranges from Bermuda to Brazil. Usually found in weed in shallow water. 376 Proceedings of the Biological Society of Washington Leptosynapta Verrill, 1867 The Carrie Bow Cay collections contain a total of 11 complete specimens and several fragments of synaptids which can be referred to the genus Leptosynapta. Suiprisingly, the specimens represent 3 new and distinctive species. While I am reluctant to describe 3 fur- ther species in a genus which is already cjuite large (approximately 25 species) and requires revision, it is clear that under currently accepted taxonomic criteria for the family Synaptidae, the present species must be regarded as new. It seems likely that further collecting in sandy areas of the Caribbean will reveal a large and diverse fauna of burrowing synaptids; such habitats have received very little at- tention from collectors in the past. The 3 species described below do not appear to be closely related to one another, but each shares some important features with otlier western Atlantic congeners. Leptosynapta imswe, new species Figure 3 Diagnosis: Anchors and anchor plates of one kind, very large, an- chors usually exceeding 600 /mi in length, plates exceeding 400 fim in length. Miliary granules numerous, in fonu of C-shaped rods with enlarged ends. Material examined: Holotype (USNM E 15854, specimen 57 mm long), lagoon, sand, north end, west side of Carrie Bow Cay, Belize, 27 April 1974, collected by F. H. C. Hotchkiss and K. Sandved. Para- types 3 complete specimens and 11 fragments (USNM E 15855) from same locality as holotype. Etymology: The species is named for the Smithsonian Institution I.M.S.W.E. program (Investigations of Marine Shallow Water Eco- systems ) . Description: Total length ranges from 7 to approximately 90 mm; probably species exceeds 120 mm in life. Specimens unifonnly whitish, body wall translucent when expanded; color in life pink to light brown. Conspicuous anchors project through body wall rendering specimens very prickly to touch. Tentacles 12, each with 5-6 pairs of digits and a terminal digit; digits increase in length distally, and terminal digit is longest (Fig. 3F). Inner (oral) surfaces of tentacles with double row of well developed sensory cups. Body wall deposits comprise large anchors and anchor plates of one kind, and numerous miliary granules. Anchors and plates at an- terior, middle and posterior of body wall essentially similar, although developmental stages of these ossicles more numerous posteriorly. An- chors average 630 /xm in length (standard deviation 2.49; standard error 0.83), 374 fim in widtli, arm-tip to arm-tip (standard deviation 2.01; standard error 0.67), 126 /zm in width of stock (standard de- Sea ciicunil}ers from Belize 377 Fig. 3. Leptosynapta iniswe n. sp. A, Anchor; B, Detail of anchor arm; C, Detail of anchor stock; D, Miliary granules from body wall; E, Ossicles from tentacle stem; F, Outline of tentacle; G, Anchor plates; H, Ossicles from tentacle digits. viation 1.26; standard error 0.42). Arms carry up to 10 conspicuous sharp teeth. Stock unbranched, but eciuipped with numerous small sharp projections (Fig. 3A-C). Anchor plates elongate, approximately oval, with numerous toothed perforations (Fig. 3G); central perforations tend to be larger than others. No true bridge for support of anchors, but in area of support, anchor plate strengthened by having an ir- regular "pseudo-bridge" in form of a double layer of calcite. Anchor plates average 447 /xva in length (standard deviation 3.09; standard error 1.03), and 246 ^m in greatest width (standard deviation 0.84; standard error 0.28). Miliary granules numerous exerywhere in body wall, highly \ariable in shape, but generally tending to have enlarged, recurved ends. Granules up to approximately 30 /im in length ( Fig. 3D). Stems of tentacles with ossicles similar to miliary granules of body wall but tending to be slightly smaller ( Fig. 3E ) . In tentacle digits 378 Proceedings of the Biological Society of Wa.sliington ossicles tend to be more elongate, length up to 45 /tni, and some have perforated ends (Fig. 3H). Remarks: This is one of the few species of Leptosynapta sciisu lato which has \er>- large anchors and anchor plates. L. acantliia H. L. Clark, known on]\' from Bermuda, has anchors and plates which are similar in size to those in L. imswe, but the plates have fewer per- forations, and further, L. acanthia has numerous small anchors 140- 210 /mi in length and plates of approximately the same size (Clark, 1924:478); such deposits are apparently lacking from L. imswe. L. imiltipora II. L. ('lark also has large anchors and plates but lacks the numerous miliary granules. Leplosynapta roseogradia, new species Figure 4A-D Diagnosis: Anchors of one type, usually less than 200 /un long, anchor plates of one type, usually less than 160 /mi long. Miliary granules C- or bracket-shaped, numerous. Radial pieces of calcareous ring perforated for radial nerve. Material examined: Holotype (USNM E15856, specimen 36 mm long), lagoon, sand, north end, west side of Carrie Bow Cay, Belize, 27 April 1974, collected by F. 11. C. Hotchkiss and K. Sandved. Para- types 3 specimens (USNM E15857) from same locality as holotype. Etymology: The species is so named to indicate its superficial simi- larity to Epitomapta roseola. Description: Total length 16, 26, 36, and 40 mm. Specimens white to yellowish, body wall thin, translucent; color in life pink to light brown. Tentacles 12, each with 5-6 pairs of digits and a terminal digit, which is longest. Sensoiy cups present in small numbers on oral surface of tentacles. Radial pieces of calcareous ring perforated for passage of radial nerve. CiHated funnels all of one type, small, approximately 100 /im in length. Deposits in body wall anchors, anchor plates and numerous miliary granules. Anchors and plates at anterior, middle and posterior of body essentially similar, but differing in dimensions: nterior anchors length 167 /nil S.D. 1.36 S.E. 0.43 plates length 154 /nil S.D. 1.29 S.E. 0.42 id-body anchors length 192 /mi S.D. 2.6 S.E. 0.93 plates length 182 fim S.D. 3.6 S.E. 1.29 plates width 142 /an S.D. 2.46 S.E. 0.89 3Sterior anchors length 148 /tm S.D. 1.32 S.E. 0.44 plates length 144 iim S.D. 1.42 S.E. 0.45 Anchors (Fig. 4A ) have 1-.3 serrations on arms; stock with mmierous small teeth. Anchor plates (Fig. 4D) approximately oval, with 7 Sea cucumbers from Belize 379 50;jm 63 ^^63^ 30 Mm ^{&(f 50)jm 100 um 40 pm 40/jm G 40jjm SO^im Fig. 4. Leptosyiuii)ta wseogiatlia n. sp. A, Anchor; B, Miliary gran- ules from body wall; C, Ossicles from tentacle stems and digits; D, Anchor plates. Leptosynapta nannoplax n. sp. E, Anchor plates; F, Anchor; G, Miliary granules from body wall; H, Ossicles from stems and digits of tentacles. major perforations, always toothed, also with varying numbers of smaller perforations. Plate distinctly narrower at articular end, with several small perforations. No true supporting bridge for anchor. Body wall with minute miliary granules ( Fig. 4B ) more or less C- or bracket- shaped, with enlarged ends. Size variable, largest granules approxi- mately 25 jim in length. Tentacle stems and digits contain granules similar to those in body 380 Proceedings of the Biological Society of Washington wall, and dibits in particular have more elongate rods (Fig. 4C) up to 60 ^m in length. Remarks: Tins is another distinctive species of Leptosynapta. The anchors and anchor plates strongly resemble those of Epitomapta roseola (Verrill) from Woods Hole, Bermuda, and Jamaica, but E. roseola has imperforate radial pieces in the calcareous ring, ciliated funnels of two types, and anchor plates which are considerably smaller (110 /nn, Heding, 1928:237) than those of L. roseogradia. Within the genus Leptosynapta, L. tenuis (Ayers) from the north- eastern United States shares some features with this new species, but differs in having larger anterior and posterior anchors, smaller an- terior anchor plates, and differently shaped miliary granules and tentacle rods. Further, in L. tenuis, the anterior and posterior anchors are distinctly different in appearance, a feature not observed in L. roseogradia. Leptosynapta nannoplax, new species Figure 4E-H Diaf^nosis: Anchors and anchor plates very small, anchors usually less than li?5 /un in length, anchor plates usually less than 90 /um in length. Miliary granules resemble rosettes. Material examined: Holotype (USNM E15858, specimen 31 nun long), lagoon, sand, north end, west side of Carrie Bow Cay, Belize, 27 April 1974, collected by F. H. C. Hotchkiss and K. Sandved. Paratypes 2 specimens (USNM E 15859) from same locality as holotype. Etymology: The specific name is derived from Greek nanuos small, and plax, a plate, in reference to the diminutive ossicles in the body wall. Deseriptio)i: Total length of 3 specimens 26, 27 and 31 mm. Species may exceed 50 mm in life. Specimens white, body wall translucent; color in life pink to light brown. Twelve pinnate tentacles, with 3-5 pairs of digits and a terminal digit which is the longest. Sensory cups present on oral surface of tentacles. Body wall ossicles comprise very small anchors and anchor plates of one kind, and minute miliary granules. Anchors and plates at an- terior, nnddle, and posterior of body sinu'lar, but differing slightly in dimensions : Anterior anchors length 86 /an S.D. 0.79 S.E. 0.25 plates length 83 iim S.D. 0.51 S.E. 0.16 Mid-body anchors length 107 in-n S.D. 4.36 S.E. 1.53 plates length 90 /un S.D. 1.91 S.E. 0.64 plates width 61 /xm S.D. 1.62 S.E. 0.51 Posterior anchors length 124 /mi S.D. 0.70 S.E. 0.22 plates length 88 /un S.D. 0.71 S.E. 0.24 Sea cucumbers from Belize 381 Anchors cany vip to 4 or 5 distinct serrations on arms; stock is toothed but not strongly so (Fig. 4F). Anchor plates elongate oval, usually with 7 large dentate perforations, and several smaller holes at anchor support area ( Fig. 4E ) . No true bridge for anchor support, but usually a double layer of calcite is more or less well developed. Miliary granules scattered, usually less than 40 ^m in length (Fig. 4G ) . They bear close resemblance to rosettes found in other groups of apodous holothurians. Tentacle stems and digits contain numerous deposits similar to miliary granides of body wall; in addition, digits in particular contain elongate rods up to appro.ximately 70 ^m in length ( Fig. 4H ) . Remarks: Very few species of Leptosijnapta have anchors and plates of such small size, and apparently none have miliary granules which resemble rosettes in combination with small anchors and plates. L. mlcropatina Heding from Tobago has anchors and plates of the same order of size as L. nannoplax, but does not possess the rosette-like miliary granules. Further, the anchor plates of L. micropatina usually contain smaller perforations at the wider ends of the plates in additioir to the larger dentate perforations; these are lacking from L. nannopiax. In \'ie\\' of the relatively small size of L. nannopiax, it is conceivable that this is the young form of a known species; if this is the case, then it is to be expected that the rosette-like miliary granules disappear or are supplanted by granules of a different type as the animal grows. CJiiiidota rotifcra (Pomtales) Chiiiclota lot if era. —Heding, 1928:293, figs. 59, 60; H. L. Clark, 1933: 122. Material examined: Foot of pier, bulk sample no. 2, 29 April 1974, collected by F. H. C. Hotchkiss and K. Sandved, 5 specimens; subtidal, bulk sample, 28 April 1974, collected by F. H. C. Hotchkiss and K. Sandved, 1 specimen; back reef, in conch shell, 1 m depth, 27 April 1975, collected by K. Sandved, 1 specimen. Remarks: Grows to about 10 cm. Body smooth to touch, with more or less conspicuous aggregations of calcareous spicules ( "wheel pa- pillae" ) scattered in rows along the interradial areas, especially dor- sally. Ground color light brick-red to pink. Viviparous. Common on sand under rock in shallow water; also occurs in sandy beaches. Ranges from Brazil to Bermuda. Literature Cited Clark, H. L. 1922. The holothurians of the genus Stichopus. Bull. Mus. Comp. Zool. Harvard 45(3):40-74, 2 pis. . 1924. The holothurians of the Museum of Comparative Zoology, The Synaptinae. Ibid. 45( 13) :459-501, 12 pis. . 1933. A handbook of the littoral echinoderms of Porto 382 Proceedings of the Biological Society of Washington Rico and the other West Indian islands. Sci. Surv. Porto Rico Virgin Islands, N. Y. 16(l):l-60, 7 pis. Dahl, A. L. 1973. Surface area in ecological analysis: quantification of benthic coral-reef algae. Marine Biology 23:239-249, 8 figs. Deichmaxx, E. 1930. The holothurians of the western part of the Atlantic Ocean. Bull. Mus. Comp. Zool. Harvard 71(3):43- 226, 24 pis. . 1963. Shallow water holothurians known from the Carib- bean waters. Stud. Fauna Curacao 14:100-118. Devaxey, D. M. 1974. Shallow-water echinoderms from British Hon- duras with a description of a new species of Ophiocorna (Ophiuroidea). Bull. Mar. Sci. 24( 1 ) :122-164, 16 figs. Heding, S. G. 1928. Synaptidae: Papers from Dr. Th. Mortensen's Pacific Expedition 1914-1916, no. 66. Vidensk Medd. dansk naturh. Foren. Kjobenhavn 85:105-323, pis. 2, 3. KiER, P. M. 1975. The echinoids of Carrie Bow Cay, Belize. Smith- sonian Contrib. Zool. 206:1-45, 8 figs., 12 pis. QH 1 B4X HH vJo. 32, pp. 383-392 12 October 1976 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON A NEW TROGLOBITIC CRAYFISH (DECAPODA, CAMBARIDAE ) FROM PENINSULAR FLORIDA By Horton H. Hobbs, Jr., and David S. Lee Department of Invertebrate Zoology, Smitlisonian Institution, Washington, D.C. 20560, and North Carolina State Museum of Natural History, Raleigh, N.C. 27611 Several previously undescribed species of troglobitic cray- fishes have been reported recently from Florida. Hobbs (1971) described Procambarus (Leconticambarus) milleri, the first new form recorded since 1942 (when he reviewed the crayfishes of the state). At this time (1971), he sum- marized the known ranges, provided a key, and illustrated the diagnostic features of all of the troglobitic species known to occur in Florida. Since then, three additional crayfishes have been named: Procambarus (O.) orcinus Hobbs and Means (1972:394), P. (O.) horsti Hobbs and Means (1972: 401), and P. (O.) erythrops Relyea and Sutton (1975:8). Accompanying the description of the latter was a revised key to the ti'oglobitic crayfishes of the state prepared by Hobbs. The species described here brings the total number of known Floridian troglobitic crayfishes to 11, all except two of which are members of the genus Procambarus. We take pleasure in naming this crayfish in honor of Richard Franz, a colleague, friend, and outstanding student of cave ecosystems. His continued interest and support of this study are greatly appreciated. We wish to thank Sylvia Scudder, Richard Bradley, Bar- bara Lee, and Richard Franz for their assistance in collect- ing the specimens on which the following description is based. We also wish to acknowledge the assistance of John E. Cooper who provided useful information leading to a^ ZT- 32— Proc. Biol. Soc. Wash., Vol. 89, 1976 :383) 384 Proceedings of the Biological Society of Washington Franz's initial location of Orange Lake Cave, the source of our specimens. For their critical reading of the manuscript, we are grateful to Thomas E. Bowman, Martha R. Cooper, and Margaret A. Daniel. Procambarus (Ortmannicus) franzi, new species Diagnosis: Albinistic, eyes without pigment or faceted cornea. Rostrum with, or more often without, marginal spines; median carina absent. Carapace with cervical spine cephalo ventral to row of spines or tubercles flanking caudal margin of cervical groove. Areola 12.8 to 17.2 times as long as broad and constituting 38.7 to 41.0% of total length of carapace (47.1 to 49.5% of postorbital carapace length). Suborbital angle absent. Postorbital ridge with cephalic spine. Hepatic area with many small tubercles, some spiniform. Antennal scale ap- proximately twice as long as wide, broadest slightly distal to midlength. Ischia of third and fourth pereiopods of first form male with simple hooks, that on third overreaching basioischial articulation and that on fourth highly arched, almost reaching basioischial articulation, and op- posed by low eminence on basis; coxa of fourth pereiopod with promi- nent oblique boss. First pleopod of first form male reaching coxa of third pereiopod, asymmetrical, provided with subapical setae; distal extremity bearing subspiculiform mesial process directed caudally at approximately right angle to shaft of appendage and curved laterally; cephalic process rather short, acute, somewhat hooding central pro- jection cephalically, and directed caudodistally; caudal element con- sisting of inconspicuous, caudolaterally situated caudal knob, and prominent, corneous adventitious process caudomesially, latter distally rounded, convex mesially, and somewhat concave laterally; and corneous beaklike central projection, most conspicuous of terminal elements, directed caudodistally subparallel to cephalic process. Annulus ven- tralis freely movable, subspindle-shaped, slightly more than twice as broad as long, and completely exposed, not (partly) hidden by pro- jections from sternum immediately cephalic to it; cephalic area with convex elevated area bearing submedian J-shaped furrow; sinus origi- nating in furrow and, following sinuous curve caudally, terminating to side of median line slightly caudal to midlength of annulus. Postannular plate slightly more than half as wide and half as long as annulus with cephalic region somewhat inflated. First pleopod in female moderately well-developed. Holotypic Male, Form I: Cephalothorax (Fig. lo, I) subcylindrical. Abdomen narrower than thorax (10.1 and 13.1 mm). Greatest width of carapace greater than height at caudodorsal margin of cervical groove. Areola 14.7 times as long as wide with 1 or 2 punctations across narrowest part. Cephalic section of carapace approximately 1.5 times as long as areola, length of latter 40.7% of entire length of New cave crayfish from Florida 385 Fig. 1. Pwcambanis (Oiimannicus) franzi, new species (all illustra- tions are of holotype except c and e of morphotype, and d of allo- type): a. Lateral view of carapace; b, c, Mesial view of first pleopod; d, Annulus ventralis; e, f. Lateral view of first pleopod; g, Basal podo- meres of third, fourth, and fifth pereiopods; h, Epistome; i, Antennal scale; /, Caudosinistral view of first pleopods; k, m. Mesial view of terminal part of first pleopod showing subterminal setae; /, Dorsal view of carapace; n, Dorsal view of distal podomeres of cheliped. 386 Proceedings of the Biological Society of Washington carapace (47.9% of postorbital carapace length). Rostrum with gently convergent margins, lacking marginal spines or tubercles but with broad-based acumen clearly delimited by mesially curved cephalic ends of rostral margins. Acumen reaching midlength of penultimate segment of antennular peduncle; dorsal surface shallowly excavate and sparsely punctate. Subrostral ridge rather weak and evident in dorsal aspect along caudal three-fifths of rostrum. Postorbital ridges well- developed, grooved dorsolaterally, and terminating cephalically in small spine, more posterior spines lacking. Caudodorsal margin of cervical groove with row of spiniform tubercles, ventralmost member of row (cervical spine), larger than others. Suborbital angle virtually obsolete. Brachiostegal spine moderately strong. Entire dorsolateral and lateral surfaces of carapace studded with tubercles, some of them spiniform. Abdomen only slightly longer than carapace (25.9 and 25.3 mm). Pleura of third through fifth abdominal segments truncate ventrally, those of fourth and fifth segments subangular caudoventrally. Cephalic section of telson with single fixed spine in caudodextral corner and 2 in caudosinistral corner. Cephalic lobe of epistome (Fig. Ih) sub- rhomboidal but asymmetrical with cephalomedian projection, 3 acute prominences on cephalosinistral margin, and 2 on cephalodextral; main body of epistome with shallow cephalomedian fovea continuous with caudally disposed submedian sulcus abutting epistomal zygoma. Ventral surface of proximal podomere of antennular peduncle with prominent submedian spine near midlength. Antenna with moderately large spine on basis and small spinifonu tubercle on ischium; flagellum extending caudally beyond telson by at least length of latter. Antennal scale (Fig It) almost twice as long as broad, widest distal to mid- length, and lamellar (part) about twice as wide as thickened lateral part. Third maxilliped extending anteriorly to, or slightly beyond, level of tip of rostrum; ischium with distolateral extremity produced in acute prominence, and lateral half of ventral surface with scattered short setiferous punctations; exopod reaching distal end of carpus. Right chela (Fig. In) subovate in cross section, not strongly de- pressed. Mesial surface of palm with several irregular rows of 10 or 11 strongly elevated tubercles; remainder of palm also tuberculate, more mesial tubercles stronger than more lateral ones on both dorsal and ventral surfaces. Both fingers with well defined submedian longi- tudinal ridge flanked proximally by tubercles and more distally by setiferous punctations. Opposable margin of fixed finger with row of 8 tubercles along proximal half of finger, third from base largest, and with single, large, more ventrally situated one slightly proximal to midlength; band of minute denticles extending almost from proximal end of finger (between and ventral to row of tubercles) to base of corneous tip, band distinctly broader distal to level of ventral tubercle. Lateral surface of fixed finger with tubercles, decreasing in size dis- Neiv cave crayfish from Florida 387 Table 1. Measurements (mm) of Procambarus (Ortmannicus) franzi. Holotype Allotype Morphotype Carapace Height 10.2 11.1 10.0 Width 13.1 12.9 12.2 Total length 25.3 26.9 25.2 Postorbital length 21.5 22.1 20.9 Areola Width 0.7 0.7 0.8 Length 10.3 10.4 10.2 Rostrum Width 3.6 3.6 3.5 Length 5.2 5.9 5.6 Chela Length of mesial margin of palm 9.0 7.0 7.0 Width of palm 5.6 4.5 4.6 Length of lateral margin 25.1 21.8 20.2 Length of dactyl 14.7 13.4 11.7 Abdomen Width 10.1 10.2 9.8 Length 25.9 26.7 25.3 tally, along proximal half followed by row of setiferous punctations, latter reaching base of corneous tip of finger. Opposable margin of dactyl with row of 9 tubercles, fifth from base largest, along proximal third of finger; minute denticles between and dorsal to row of tubercles and forming broad band immediately distal to last tubercle of row, band continuing to base of corneous tip of finger. Mesial surface of dactyl similar to lateral surface of fixed finger with tubercles diminish- ing in size distally. Carpus of cheliped longer than broad, tuberculate on all surfaces, conspicuously so mesially where 1 tubercle more prominent than others; additional strong tubercle present on dorsomesial distal angle. Shallow oblique sulcus on dorsal surface flanked by small subsquamous tubercles. Ventrodistal margin of podomere with 3 tubercles: weak lateral one, moderately strong one on articular knob, and prominent acute one mesially; row of 3 tubercles extending proximally from latter. Merus of cheliped strongly tuberculate except for proximal parts of mesial and lateral surfaces. Dorsal surface with linear series of tubercles basally continuous with broadening band of them distally; tubercles on ventral surface abundant, not limited to usual 2 rows; rows poorly 388 Proceedings of the Biological Society of Washington defined but each consisting of 20 to 23 tubercles. Ischium with row of 6 tubercles ventromesially, distalmost slightly larger than proximal 5. Hooks on ischia of third and fourth pereiopods (Fig. Ig) as de- scribed in "Diagnosis." Coxa of fourth pereiopod with prominent oblique (almost vertical), somewhat inflated caudomesial boss; that of fifth pereiopod with very weak prominence corresponding to caudo- mesial boss on fourth pereiopod. Sternum between third and fourth pereiopods rather deep with conspicuous mat of plumose setae extending mesially from ventrolateral margins. First pleopods (Fig. lb, f, j, k, m) as described in diagnosis; mesial process of left member not bent nearly so strongly laterally as that on right pleopod. Uropod with both lobes of basal podomere bearing spines; distomedian spine on mesial ramus far removed from distal margin of ramus. Allotypic Female: Differing from holotype in following respects: rostrum with marginal spines and reaching proximal end of ultimate podomere of antennular peduncle; subrostral ridges evident in dorsal aspect only along basal fourth of rostrum; tubercles of spiniform row flanking caudal margin of cervical groove much less conspicuous and cervical spine smaller; cephalic section of telson with 2 spines in each caudolateral corner; cephalic lobe of epistome symmetrical with weak prominences at angles of rhomboid; opposable margin of fixed finger of chela with row of 9 tubercles, fourth from base largest, that of dactyl with row of 13 tubercles, fifth from base largest; ventromesial margin of ischium of cheliped with row of 7 tubercles. Annulus ventralis (Fig. Id) only moderately deeply situated in sternum (see "Diagnosis" for details). Sternum immediately cephalic to annulus lacking ornamentations. First pleopod reaching slightly cephalic to caudal margin of annulus when abdomen flexed (see "Mea- surements"). Morphotypic Male, Form II: Differing from holotype in following respects: rostrum with small marginal tubercle dextrally (sinistral margin as in holotype), and reaching almost midlengdi of ultimate podomere of antennular peduncle; subrostral ridges evident in dorsal aspect from caudal margin of orbit to acumen; postorbital ridge with well defined posterior tubercles, sinistral one almost serrate; cervical spine very small, hardly larger than adjacent tubercles; epistome simi- lar to that of allotype but anterolateral angles less sharp, and weak prominences present closely flanking cephalomedian prominence; ce- phalic section of telson with single fixed spine in each caudolateral comer; lateral ramus of sinistral uropod deformed, possessing bilobed distal segment; opposable margin of fixed finger of chela with row of 6 tubercles, that of dactyl with 11; ventromesial surface of ischium of cheliped with row of 8 tubercles and few others nearby; hooks on ischia of third and fourth pereiopods much reduced, that on third not overreaching basioischial articulation; boss on coxa of fourth pereiopod also much reduced. First pleopod (Fig. Ic, e) without corneous ter- ISlew cave crayfish from Florida 389 minal elements; mesial and cephalic processes much shorter and heavier, caudal element much less sharply defined, and central projection shorter, and decidedly more tumescent (see "Measurements"). Type-locality. Orange Lake Cave, 0.4 mi S of junction of U.S. Hwy. 441 and State Route 318 off Hwy. 411 (T.12S, R.21E, Sec. 33/34), Marion County, Florida. Orange Lake Cave is a small, horizontal cavern located on the north side of a quarry of several acres. Tlie status of the entrance prior to quanying operations is unknown. The two present openings (2 m X 1.5 m; 1.5 m X 3 m) lead into a small room. The main passage (approximately 1 m wide) leads north and northeast from about 20 m and terminates in a large, irregular room ( 15 m long, 5 m wide, and 3-7 m high) where the water level fluctuates markedly. On 8 Sep- tember 1974, water was encountered within 5 m of the entrance, and the floor of the entire passage and that of the room in the back of the cave were inundated. On 18 May 1975 and 19 November 1975, the water was about a meter lower and was confined to the rear of the big room. On other visits (5 January 1975, 28 September 1974, etc.) water levels were intermediate between these extremes. Water tempera- ture remained at 21-22° C throughout the collecting period. In the spring, as many as 3700 bats (Myotis austroriparius) use the back room of this cave as a maternity site. Throughout the remainder of the year, the bats are not usually present, but Richard Franz re- ported 6000 individuals having been found there on 19 November 1975. The bats are probably responsible for the major source of energy for the Orange Lake Cave crayfish population. Nearly all of the crayfish were observed in the pool directly under the bat roosts. The total number of crayfish counted on 18 May 1975 and 19 No- vember 1975, when they were confined to one pool, was 32 and 23, respectively. A large percentage of the population consisted of juveniles. On two occasions (October 1974 and 5 January 1975), several small, white crayfish, assumed to be a part of this same population, were seen in a small solution cavity in the floor of the quarry. The water level was about 1.5 m below the bottom of the quarry. Other openings in the sides and bottom of this same quarry apparently do not contain water. Disposition of Types: The holotype, allotype, and morphotype are deposited in the National Museum of Natural History (Smithsonian Institution), numbers 146992, 146993 and 146994, respectively, as are the paratypes consisting of 1 5 II, 5 9, and partly decayed remains of IS I. Size: The largest of the specimens is a female having a carapace length of 29.0 mm (postorbital carapace length 23.5 mm). Perhaps the (partly) decayed form I male was a little larger. The only other first form male is the holotype possessing corresponding lengths of 25.3 and 21.5 mm. Range and Specimens Examined: This crayfish is known only from 390 Proceedings of the Biological Society of Washington the type-locality, and the only specimens available are those included in the type-series: 1 5 1, 8 September 1974, R. Franz and D. S. Lee, coll.; UII, 29, 1 October 1974, R. F. and D. S. L., coll.; 3 9, 19 May 1975, B. Lee and D. S. L.; 151, 1?, 19 November 1975, R. F., R. Bradley, and S. Scudder, coll.; 1 5 H, 8 December 1975, R. F. coll. Variations: The most conspicuous variation observed is in the ros- trum. Except in a few specimens in which there are marginal spines present and in the morphotype in which such a spine is present on one side, all of the other specimens lack marginal spines. Instead the rostral margins curve mesially at the base of the acumen, merging with the flattened dorsal surface of the latter. The subrostral ridges which are continuous with the margin of the acumen may be visible along the length of the rostrum from the level of the caudal margin of the orbit to the tip of the acumen, or the rostral margins may obscure, in dorsal view, part of them between the orbit and the base of the acumen. The other slight variations noted are in the number, dis- position, and sizes of the tubercles, both on the carapace and on the chelipeds, but only those differences occurring on the postorbital ridges seem worthy of mention. Even in a single individual, the tubercles on the paired ridges are not identical; one ridge may have an arrange- ment of tubercles that might be described as almost serrate while the tubercles on the other are so reduced as to be easily overlooked. The only consistency is the presence of a small apical tubercle. Relationships: Procambariis (Ortmannicus) franzi is the seventh troglobitic member of the subgenus to be described, all of them from the subterranean waters of Florida, and all at least as closely related to one another as to any epigean crayfish. Three of the previously described species, P. (O.) pallidus (Hobbs, 1940:394), P. (O.) horsti and P. (O. ) orcinus share in common a somewhat laterally displaced cephalic process on the constricted distal part of the first pleopod of the male, and the sternum immediately cephalic to the annulus ven- tralis in the female is produced into caudally projecting prominences that are not present in the remaining ones : P. ( O. ) hicifugus lucifugus (Hobbs, 1940:398), P. (O.) /. alachua (Hobbs, 1940:402), P. (O.) erythrops, and the species described here. The range of P. (O. ) franzi to the south and east of P. (O. ) I. alachua, with which it seems to share more features than with the other species, tends to support an assumption of close kinship. Speci- mens that have been identified as intergrades between the two sub- species of P. ( O. ) lucifugus were obtained from several caves approxi- mately 20 mi S of Orange Lake Cave (see Warren, 1961:7). The characters cited above distinguish it from P. ( O. ) pallidus, P. (O. ) horsti, and P. (O. ) orcinus, and the absence of pigment in the eye serves readily to separate it from P. (O. ) I. alachua and P. (O. ) erythrops. The tapering rostrum distinguishes it from P. (O.) I. luci- fugus, and the terminal part of the first pleopod of the male is unique. Common Name: Because this crayfish appears to have such a limited Islew cave crayfish from Florida 391 range, the possibility of its ultimately being placed on the list of en- dangered species is very real. Inasmuch as a common name will be- come mandatory when its status of probable safety is considered, we propose that it be known as the Orange Lake Cave Crayfish. Literature Cited HoBBS, HoRTON H., Jr. 1940. Seven new crayfishes of the genus Camharus from Florida, with notes on other species. Proc. U.S. Nat. Mus. 89 (3097): 387-423, figs. 14-22. . 1942. The crayfishes of Florida. Univ. Fla. Publ., Biol. Sci. Ser., 3(2):1-179, 24 pis. . 1971. A new troglobitic crayfish from Florida. Quart. Jour. Fla. Acad. Sci. 34(2):114-124, 19 figs. , AND D. Bruce Means. 1972. Two new troglobitic cray- fishes (Decapoda, Astacidae) from Florida. Proc. Biol. Soc. Wash. 84 (46). -393-409, 2 figs. Relyea, ICenneth, and Bruce Sutton. 1975. A new troglobitic crayfish of tlie genus Procambarus from Florida (Decapoda, Astacidae). Tulane Stud. Zool. Bot. 19(l-2):8-16, 4 figs. Warren, Richard D. 1961. The obligative cavernicoles of Florida. Fla. Speleol. Soc, Spec. Pap. 1:1-10, 2 figs. 392 Proceedings of the Biological Society of Washington ^"^ QH i B4X No. 33, pp. 393-404 12 October 1976 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON NEW ENTOCYTHERID OSTRACODS FROM KENTUCKY AND TENNESSEE By Horton H. Hobbs, Jr., and Margaret Walton Department of Invertebrate Zoology, Smithsonian Institution, Washington, D.C. 20560, and Mountain Lake Biological Station Pembroke, Virginia 24136 O cr: Four new ostracods are described from crayfishes collected in the Cumberland and Tennessee River basins in Kentucky and Tennessee. One, a member of the genus Ascetocythere, is believed to be restricted to a burrowing crayfish, whereas the other three, assigned to the genus Dactylocy there, infest sti'eam dwelling members of the genus Cambarus and per- haps also two species of the genus Orconectes. The most recent keys and summary of the genera Ascetocythere and Dactylocythere are those of Hart and Hart (1974). Except for four lots of specimens, from three localities, which were collected by Perry C. Holt, (1 lot), Joseph F. Fitzpatrick, Jr., and H. H. Hobbs, Jr., (2), and Daniel J. Peters, Jean E. Pugh, and H. H. Hobbs, Jr. (1), all were donated to us by Raymond W. and Judith Way Bouchard who also supplied us with identifications of the hosts collected by tiiem. We gratefully acknowledge the gift from the Bouchards and Perry C. Holt, and the assistance given to us by the other collectors. We also wish to thank Margaret A. Daniel, C. W. Hart, Jr., and Raymond B. Manning for their critical reading of the manuscript. AscetocyShere riopeli, new species (Figure \a-d) Male: Eye pigmented and located between one-fourth and one-fifth shell lengtli from anterior margin. Shell (Fig. la) ovate with greatest 33— Proc. Biol. Soc. Wash., Vol. 89, 1976 (393) 394 Proceedings of the Biological Society of Washington Fig. la-d, Ascetoctjthere riopeli, new species; le-h, Dactylocythere apheles, new species, a, e. Shell of male; b, g, Copulatory complex of male; c, h. Clasping apparatus of male; d, f. Shell of female. Scales height one-third shell length from posterior margin where about 1.4 times height at level of eye; margin entire. Submarginal setae present except dorsally, closer together anteriorly and posteriorly than ven- trally. Copulatory complex (Fig. lb, c) with bulbous ventral portion of peniferum bearing following processes: that borne on cephaloventral margin, and serving as conduit guiding penis, swollen distally and New entocytherid ostracods 395 flared with margin produced in several small prominences; adjacent ventrally directed process slender, subsinuous with cephaloventral ex- tremity slightly broadened and turned laterally; vestigial caudal process situated near caudal base of sinuous process, its subacute apex di- rected cephalically. Penis very prominent, L-shaped, with spermatic and prostatic elements gaping for some distance at and on both sides of midlength of penis, two converging in basal part of cephaloventral process of peniferum and emerging on mesial surface of swollen distal region. Clasping apparatus only slightly curved, tapering from base, its postaxial border entire; preaxial border with 3 teeth on distal third followed by 2 apical ones. Dorsal finger moderately heavy, its setiform tip overreaching midlength of uniformly curved ventral finger, tip of which directed anteroventrally. Triunguis Female: Eye pigmented and situated approximately one- fourth shell length from anterior margin. Shell (Fig. Id) elongate ovate, highest about one-third shell length from posterior margin where 1.3 times height at level of eye. Submarginal setae disposed as in male but closer together anteroventrally than elsewhere. Genital complex consisting of small, weakly sclerotized prominence with concave apical margin and embedded in heterogeneous mass projecting posteroven- trally from posterodorsal part of body. Measurements (in mm): 4 males and 1 female. Holotype Males Allotype Length (range) 0.39 0.38-0.41 0.41 Average 0.39 Height (range) 0.22 0.22-0.24 0.22 Average 0.22 Type-locality : Poor Fork of the Cumberland River, at Upper Cumber- land School, 0.6 mi from Virginia state line, Letcher County, Ken- tucky. The specimens were retrieved from a collection of crayfishes containing Cambarus {Jiigicambarus) diibius Faxon, C. (/. ) distans Rhoades, C. (Pimcticambarus) robustus Girard, and C (P.) buntingi Bouchard. Disposition of Types: The holotypic male and allotypic female are deposited in the National Museum of Natural History (Smithsonian Institution), numbers 155317 and 155318, respectively, as are the paratypes. Range and Specimens Examined: Six specimens from two localities in Kentucky: Type-locality, 30 March 1974, R. W. and J. W. Bouchard, coll. and Marrowbone Creek at State Route 195 and Elkhorn Creek at State Route 197 (combined collection), Pike County, Kentucky, 29 March 1974, R.W.B. and J.W.B., coll. Hosis: Although five species of crayfishes are represented from the collections containing this ostracod {Cambanis {Jiigicambarus) dubius. 396 Proceedings of the Biological Society of Washington C. (J.) distans, C. (Ptincticambarus) robustus, C. (P.) buntingi, ana Orconectes juvenilis ( Hagen ) ) , it is highly probable that this ostracod is confined to C. {].) dubius. Entocytherid Associates: In the type-locality, Ascetocythere riopeli was found in a collection containing Dactylocythere spinata Hobbs and Walton (1970:860), Dactylocythere sp.?, and Donnaldsoncythere donnaldsonensis (Klie, 1931:334). In the Pike County locality, there were no associates. Relationships: Ascetocythere riopeli is a member of the Asceta Group and is perhaps more closely allied to As. sclera Hobbs and Hart (1966: 42) than to any other member of the Group. The absence of a cephalic process on the peniferum of the male and the presence of a slender sinuous sclerotized process projecting ventrally from the bulbous area provide a unique combination of characters separating this species from its congeners. Etymology: This ostracod is named in honor of our mutual friend, James L. Riopel, Director of the Mountain Lake Biological Station of the University of Virginia, who, for a number of years, has encouraged us in our studies of tlie entocytherids. Dactylocythere apheles, new species (Figure le-h) Male: Eye pigmented and located between one-fifth and one-sixth shell lengtli from anterior margin. Shell (Fig. le) subovate; greatest height slightly posterior to midlength where almost 1.4 times that at level of eye; ventral margin weakly convex. Submarginal setae some- what closer together anteriorly and posteriorly than ventrally, and absent dorsally. Sternal spine lacking. Copulatory complex (Fig. Ig, h) with slender, distally tapering finger guard provided with anteriorly eccentric tip; peniferum moder- ately slender with ventral part curved anteroventrally and tapering rather rapidly to acute anteroventral angle; ventral part of tip corneous. Peniferal groove opening anteroventrally, almost closed in apical region, its least diameter approximately one-tenth that of corresponding di- ameter of vertical ramus of clasping apparatus. Penis L-shaped with arms subequal in length. Accessory groove represented by short tri- angular clear area posterodorsal to base of penis, its apex barely or not reaching level of spermatic loop. Clasping apparatus C-shaped with short proximal area subparallel to longer distal segment (horizontal ramus); apparatus of almost uniform diameter throughout and with margins entire; distal part with distally radiating grooves extending toward 5 or 6 apical denticles. Dorsal and ventral fingers comparatively slender, latter about twice length of former and gently curved with distal half directed anteriorly. Triunguis Female: Eye pigmented, situated as in male. Shell (Fig. 1/) strongly arched dorsally and with almost straight ventral margin; New entocytherid ostracods 397 greatest height slightly posterior to midlength where about 1.5 times that at level of eye. Submarginal setae disposed as in male. Posterior margin of shell entire, smoothly rounded. Genital complex consisting of weakly sclerotized, slender, tubular papilla directed posteroventrally; usual J-shaped rod and amiculum lacking. Measurements (in mm) : 3 males and 4 females. Holotype Males Allotype Females Length (range) 0.41 0.40-0.41 0.39 0.36-0.39 Average 0.40 0.38 Height (range) 0.22 0.21-0.22 0.21 0.21-0.22 Average 0.22 0.21 Type-locality: Spring seep flowing across old road to Chimneys at Indian Gap (Little Pigeon River drainage), Sevier County, Tennessee. There the type-series was collected with Uncinocythere zancla Hobbs and Walton (1963:456) on 13 November 1971, by R. W. Bouchard and J. D. Way. Disposition of Types: The holotypic male and allotypic female are deposited in the National Museum of Natural History (Smithsonian Institution), numbers 155319 and 155320, respectively. Paratypes are in the collection of H. H. Hobbs III and the Smithsonian Institution. Range and Specimens Examined: Seven specimens collected at the type-locality ( Little Pigeon River basin ) . Host: Cambarus (J.) carolinus (Erichson). Entocytherid Associate: Uncinocythere zancla. Relationships: Dactylocythere apheles has its closest affinities with Dactylocythere leptophylax (Crawford, 1961:238). The male of both species possesses a peniferum with a reduced accessory groove, at least occasionally hardly discernible; the margins of the clasping ap- paratus are entire, and the distal extremity bears at least 4 denticles set off proximally by distally diverging striae. The females of both species lack the usual J-shaped rod and amiculum. Members of the new species differ from those of Dt. leptophylax in possessing a simple as opposed to a distally bifid or trifid finger guard; tlie ventral extremity of the peniferum is smooth, lacking tuberculiform prominences, and the distal part of the clasping apparatus is not expanded. Etymology: Apheles, G. = smooth; referring to the absence of teeth or prominences on the preaxial border of the clasping apparatus and on the ventral margin of the peniferum of the copulatory complex of the male. Dactylocythere brachydactylus, new species (Figure 2a-d) Male: Eye pigmented and situated about one-fifth shell length from anterior margin. Shell (Fig. 2c) subovate with greatest height about 398 Proceedings of the Biological Society of Washington one-third shell length from posterior margin where approximately 1.3 times height at level of eye; ventral margin straight to weakly convex. Submarginal setae present around entire shell, slightly closer to- getlier anteriorly and posteriorly than dorsally or ventrally; other setae widely scattered over surface of shell. Sternal spine lacking. Copulatory complex (Fig. 2b, d) with short, robust finger guard produced in bifurcate extension from cephaloventral side and with posterior angle at level of base of extension; peniferum comparatively heavy with ventral portion subtruncate and apical part directed antero- ventrally. Peniferal groove conspicuous, its least diameter from one- to three-fourths as wide as least diameter of vertical ramus of clasping apparatus. Penis L-shaped with rami subequal in lengdi. Accessory groove slender and long, reaching dorsally distinctly beyond level of dorsal extremity of spermatic loop. Clasping apparatus more L- than C-shaped, rami indistinctly delimited; postaxial margin of vertical ramus with slight angle near midlength, and preaxial margin of hori- zontal ramus with 1 or 2 minute teeth near 3 terminal denticles and marked by oblique striae extending proximally from teeth; remaining borders of apparatus entire. Dorsal and ventral fingers slender, former about one-half length of gently and evenly curved ventral finger, latter directed anteriorly. Triunguis Female: Eye pigmented, situated between one-fifth and one-sixth shell length from anterior margin. Shell (Fig. 2a) much more highly vaulted than in male, highest about one-third shell length from posterior margin where approximately 1.5 times height at level of eye; ventral margin slightly concave near midlength. Disposition of setae on shell similar to that on male. Posterior margin of shell entire and rounded. Genital complex consisting of slender, non- corneous tubular prominence, directed posteroventrally, flanked poste- rodorsally by similarly directed inflated sclerotized projection capped by hyalin coat and with heterogeneous material adhering to antero- ventral margin; usual J-shaped rod and amiculum lacking. Measurements {in mm): 10 males and 10 females. Holotype Males Allotype Females Length (range) 0.47 0.44-0.47 0.49 0.47-0.49 Average 0.46 0.48 Height (range) 0.27 0.25-0.27 0.29 0.27-0.29 Average 0.25 0.29 Type-locality: Goose Creek at State Route 66 southwest of Dand- ridge, Jefferson County, Tennessee. There the host was Cambarus (C) bartonii (Fabricius); specimens of Orconectes virilis (Hagen) were also collected in this locality and may have been infested with New entocytherid ostracods 399 this ostracod. No other ostracod was infesting the host. Collections were made there on 16 March 1969 by R. W. Bouchard. Disposition of Types: The holotypic male and allotypic female are deposited in the National Museum of Natural History (Smithsonian Institution), numbers 155321 and 155322, respectively. Paratypes are in the collections of H. H. Hobbs III, D. J. Peters, and the Smithsonian Institution. Range and Localities: Little Tennessee, French Broad, Nolichucky, and Tennessee river basins in Cocke, Jefferson, and Roane counties, Tennessee. Tennessee: Cocke County — (1) Jake Best Creek on road S from Citico Creek, SE of Acorn (Host, Cambarus (C. ) bartonii (Fabricius)). Jefferson County — (2) Type-locality; (3) Long Creek at State Rte. 32 and U.S. Hwy. 25 E, N of White Pine (Host, Cambarus (Hiaticam- barus) longirosiris Faxon); (4) Spring Creek on U.S. Hwy. 70 W of junction with State Rte. 113 near Oak Grove (Host, Cambarus (C ) bartonii); (5) Embayment of Spring Creek off U.S. Hwy. 70 and State Rte. 9, near Douglas Lake (Hosts, Cambarus (C. ) bartonii, Orconectes virilis ( Hagen ) , Orconectes forceps ( Faxon ) ) . Roane County — (6) Caney Creek at Buttermilk Road, off 1-40 SE of Brad- bury (Hosts, C. (C.) bartonii, C. (H.) longirostris). Collections from localities 1-4 were made by R. W. Bouchard, that from 5 by D. A. Etnier, and that from 6 by Bryant. Hosts: Although this ostracod was found in collections of crayfishes containing Cambarus (C. ) bartonii, C. (H.) longirostris, Orconectes forceps, and O. virilis, it is probable that it is actually associated with only the former two. Entocytherid Associates: In the Cocke County locality it was asso- ciated with Donnaldsoncythere donnaldsonensis (Klie, 1931:334); in the Long Creek locality in Jefferson County, it was found with Un- cinocythere simondsi (Hobbs and Walton, 1960:17); and in the Roane County locality, with Dactylocythere falcata (Hobbs and Walton, 1961:379). Relationships: Dactylocythere brachydactylus has its closest affinity with Dt. chelomata (Crawford, 1961:242). The similarities in the genitalia of the males are striking; however, members of the former possess a much longer accessory groove in the peniferum, one ap- proaching the length of that in Dt. mecoscapha (Hobbs and Walton, 1960:19) and Dt. macroholca Hobbs and Hobbs (1970:6). In addi- tion, the denticles along the distal preaxial border of the horizontal ramus of the clasping apparatus are not nearly so well-developed. The females of the two species may be distinguished by the absence of a J-shaped rod and amiculum in the genital complex of Dt. brachydacty- lus. Etymology: Brachys G. = short + Dactylus, G. = finger; refer- ring to the comparatively short finger guard on the copulatory complex of the male. 400 Proceedings of the Biological Society of Washington Fig. 2a-d, Dactylocythere hrachydactylus, new species; 2e-h, Dacty- locythere demissa, new species, a, e. Shell of female; h, h, Clasping apparatus of male; c, f, Shell of male; d, g, Copulatory complex of male. Scales in mm. Dactylocythere demissa, new species (Figure 2e-h) Male: Eye pigmented and situated about one-fourth shell length from anterior margin of shell. Shell (Fig. 2/) elongate ovate (about 1.8 times as long as high) with greatest height about one-third shell length from posterior margin where about 1.3 times height at level New entocytherid ostracods 401 of eye; ventral margin almost straight. Submarginal setae present ex- cept dorsally, closer together anteriorly and posteriorly than ventrally. Sternal spine absent. Copulatory complex (Fig. 2g, h) with robust finger guard pro- duced in bifurcate extension from cephaloventral side and with poste- rior angle at level of base of extension; penifermn comparatively stout with ventral portion truncate, anteroventral angle acute and directed anteriorly. Peniferal groove distinct, rather uniformly narrow with least diameter about one-fifth that of least diameter of vertical ramus of clasping apparatus. Penis somewhat L-shaped with distal ramus hardly more than half length of proximal ramus. Accessory groove short, not nearly reaching ventral extremity of spermatic loop, with end adjacent to penis forming inverted (ventrally directed) loop — unique in the genus in this respect. Clasping apparatus L-shaped with postaxial border distinctly angular at junction of horizontal and vertical rami; vertical ramus with postaxial border concave but otherwise entire, its preaxial border almost straight to broadly but shallowly concave; horizontal ramus with postaxial border entire, its preaxial border with 2 small teeth on distal half, and apex of ramus with 3 anterodorsally directed denticles. Dorsal finger comparatively heavier than very slender ventral finger and almost one-half as long; latter gently curved throughout most of its length and with distal portion directed anteriorly. Triungiiis Female: Eye as in male. Shell (Fig. 2e) distinctly more highly vaulted than in male (about 1.5 times as long as high) with greatest height approximately one-third shell length from posterior margin where about 1.3 times height at level of eye; ventral margin slightly concave anterior to midlength. Disposition of submarginal setae on shell similar to that on male but also with widely spaced ones dorsally. Posterior margin of shell often with eccentric prominence in vicinity of amiculum. Genital complex consisting of J-shaped rod with long vertical arm inclined subparallel to posterodorsal margin of shell and with conspicuous but short amiculum borne on curved portion of rod; amiculum with short supportive hyaline arcs and often slightly protruding from between caudal margin of valves of shell. Measurements ( in mm ) : 10 males and 10 females. Holotype Males Allotype Females Length (range) 0.42 Average Height (range) 0.22 Average Type-locality: Pokepatch Creek at County Road 4385, southwest of Pleasant Hill, Cumberland County, Tennessee (Caney Fork River drainage). Specimens were obtained from a collection of crayfishes (containing Cambarus (Veticambarus) pristinus Hobbs, Cambarus iJugicambarus) parvoculus Hobbs and Shoup, and Cambarus (De- 0.40-0.43 0.42 0.40-0.43 0.41 0.42 0.21-0.24 0.25 0.23-0.26 0.23 0.25 402 Proceedings of the Biological Society of Washington pressicambams) sphenoides Hobbs) made by R. W. Bouchard on 7 August 1969. Disposition of Types: The holotypic male and allotypic female are deposited in the National Museum of Natural History (Smithsonian Institution) number 155323. Paratypes are in the collection of H. H. Hobbs in, J. D. Peters, and the Smithsonian Institution. Range and Localities: Caney Fork and Obey (Cumberland River), Emory, and Tennessee river basins on the Cumberland Plateau. Tennessee: Bledsoe County — (1) McGill Creek at County Rd. 5881 S of Brayton (Hosts, C. (D. ) sphenoides Hobbs, C. (/. ) parvoculus Hobbs and Shoup); (2) Roaring Creek off County Rd. 5881, SW of New Harmony (Hosts, C. (D.) sphenoides, C. (J.) parvoculus); (3) Henderson Creek at County Rd. 5881 NE of Summer City (Hosts, C. (D.) sphenoides, C. (/. ) parvoculus); (4) Moccasin Creek at County Rd. 5881 SW of Milo (Host, C. (/.) parvoculus); (5) Glade Creek at State Rte. 30, NW of Pikeville (Host, C. (/.) parvoctdus). Cumber- land County — (6)Type-locality; (7) Caney Fork River at U.S. Hwy. 70 (Hosts, C. (D. ) sphenoides, C. (V.) pristinus Hobbs); (8) Clear Creek at County Rd. 4794, 3 mi W of junction with U.S. Hwy. 127 (Host, C (/. ) crinipes Bouchard); (9) No Business Creek at U.S. Hwy. 127, N of Isoline (Hosts, C. (D.) sphenoides, C. (J.) distans Rhoades); (10) Little Obed River at U.S. Hwy. 127, N city limits of Crossville (Hosts, C. (D. ) sphenoides, C. {}.) distans, C. (/. ) par- voculus); (11) Daddy's Creek at U.S. Hwy. 127 near Big Lick (Host, C. (/.) distans); (12) White Oak Creek, 3.9 mi E of White County line and 0.1 mi S of U.S. Hwy. 70 (Hosts, C. (D. ) sphenoides, C. (V.) pristinus); (13) Fox Creek off County Rd. 4252 (Catoosa Wild- life Management Area ) ( Hosts, C. ( /. ) distans Rhoades, C. ( /. ) parvoctdus); (14) South Fork of Elmore Creek at County Rd. 4252 (Hosts, C. (D. ) sphenoides, C. (J.) distans, C. (J.) parvoculus); (15) Caney Fork River at County Rd. between U.S. Hwy. 70 and 70N, NE of Pleasant Hill (Hosts, C. (D. ) sphenoides, C. {}.) parvoculus, C. {V.) pristinus). Grundy County — (16) Firescald Creek at County Rd. 4398, in Altamont (Hosts, C. (D.) sphenoides); (17) Piney Creek at State Rte. 108, at Altamont (Host, C. (D. ) sphenoides). Putnam Cotinty — (18) Dripping Springs Creek at State Rte. 62, SE of Monterey (Hosts, C. (D. ) sphenoides, C. (J.) obeyensis). Rhea County— (19) Wliites Creek at State Rte. 68, NW of Grand View (Hosts, C. (D. ) sphenoides, C. (J.) parvoculus). White County —(20) Pole Bridge Branch at County Rd. 4385, S of DeRossett (Hosts, C. (D. ) sphenoides, C. (J.) parvoctdus). All of the specimens from the above localities, except stations 8, 12, and part of those from 6 were collected by R. W. Bouchard or R.W.B. and J. D. Way. Hosts: As may be noted above, in most of the localities this os- tracod was associated with Camharus (D.) sphenoides, frequently with C. (/. ) parvoculus, also with C. (V.) pristinus in the Caney Fork drainage, and occasionally with C ( /. ) distans. 'New entocytherid ostracods 403 Entocytherid Associates: The entocytherids most frequently sharing the hosts with Dactylocythere demissa are Donnaldsoncythere don- naldsonensis, and Dactylocythere brachystrix Hobbs and Walton (1966:2). Occasionally accompanying it are Dt. pachysphyrata Hobbs and Walton (1966:3), Dt. speira Hart and Hart (1971:113), and Entocythere sp. In one locality each, it was found in association with Dt. arcuata (Hart and Hobbs, 1961:173) and Dt. spinata Hobbs and Walton (1970:853). Relationships: One of the most distinctive features of this ostracod is the very short accessory groove of the peniferum, which, in its length, is similar to that in Dactylocythere coloholca Hobbs and Hobbs (1970:7), Dt. exoura Hart and Hart (1966:5), and Dt. speira. In other respects, it also has as much in common with these species as with any of its congeners. In none of the three, however, is the finger guard of the copulatory complex produced in a bifid tip. While Dt. demissa appears to be more closely allied to Dt. speira than to the other two in possessing a similar clasping apparatus and a rather broad accessory groove, the peniferum of the former is truncate ven- trally, the accessory groove is disposed in a distinct ventral loop, and the J-shaped rod of the genital complex of the female has an almost straight shaft with a gently curved ventral part rather than being strongly curved throughout, almost forming a spiral. In addition, Dt. demissa possesses a distinct amiculum that is lacking in Dt. speira. Etymology: Demissus, L. = hanging down; so named because of the long, almost straight J-shaped rod in the genital complex of the female; also the looped ventral part of the accessory groove in the peniferum of the male suggests a collapse of the groove. Literature Cited Crawford, E. A. 1961. Three new species of the genus Entocythere (Ostracoda, Cytheridae) from North and South Carolina. Amer. Midi. Nat. 65(1) :236-245, 21 figs. Hart, C. W., Jr., and Dabney G. Hart. 1966. Four new entocy- therid ostracods from Kentucky, with notes on the troglobitic Sagittocythere harri. Notulae Naturae, Acad. Nat. Sci., Philad. 388, 10 p., 13 figs. , AND HoRTON H. HoBBS, Jr. 1961. Eight new troglobitic ostracods of the genus Entocythere (Crustacea, Ostracoda) from the eastern United States. Proc. Acad. Nat. Sci., Philad. 113(8) :173-185, 32 figs. Hart, Dabney C, and C. W. Hart, Jr. 1971. New entocytherid ostracods of the genera Ankylocythere, Dactylocythere, En- tocythere, Geocythere, and Uncinocythere — with a new di- agnosis of the genus Entocythere. Proc. Acad. Nat. Sci., Philad. 123(5) :105-125, 13 figs. , and . 1974. The Ostracod Family Entocytheridae. Acad. Nat. Sci., Philad., Monograph 18: ix -f- 238 p., 62 pis. 404 Proceedings of the Biological Society of Washington HoBBS, HoRTON H., Jr., AND C. W. Hart, Jr, 1966. On the entocy- therid ostracod genera Ascetocy there, Plectocy there, Phymocy- there (gen. nov. ), and Cymocy there, with descriptions of new species. Proc. Acad. Nat. Sci., Philad. 118(2):35-61, 37 figs. , AND H. H. HoBBS in. 1970. New entocytherid ostracods with a key to the genera of the subfamily Entocytherinae. Smithsonian Contrib. Zool. 47:19 p., 9 figs. , AND Margaret Walton. 1960. Three new ostracods of the genus Entocythere from the Hiwassee drainage system in Georgia and Tennessee. Journ. Tenn. Acad. Sci. 35(1):17- 23, 20 figs. , AND . 1961. Additional new ostracods from the Hi- wassee drainage system in Georgia, North Carolina, and Tennessee. Trans. Amer. Micros. Soc. 80(4) :379-384, 8 figs. , AND . 1963. Three new ostracods (Ostracoda, Entocy- theridae) from the Duck River drainage in Tennessee. Amer. Midi. Nat. 69(2):456-461, 10 figs. , AND . 1966. A new genus and six new species of entocytherid ostracods (Ostracoda, Entocytheridae ) . Proc. U.S. Nat. Mus. 119 (3542): 1-12, 2 figs. , AND . 1970. New entocytherid ostracods from Ten- nessee and Virginia. Proc. Biol. Soc. Wash. 82(68) :851- 864, 3 figs. Klie, W. 1931. Campagne speologique de C. Bolivar et R. Jeannel dans I'Amerique du Nord (1928). 3. Crustaces Ostracodes. Biospeol.: Archiv. Zool. Exp. et Gen. 71(3):333-344, 20 figs. \\ OH i 34, pp. 405-410 12 October 1976 NH // c:5 PROCEEDINGS |/C -i OF THE ' ^ 2 — blOLOGICAL SOCIETY OF WASHINGTON ^ 5! \ ^ ^i) TWO NEW SEA CUCUMBERS (ECHINODERMATA: y HOLOTHUROIDEA) FROM THE EASTERN UNITED STATES By David L. Pawson Department of Invertebrate Zoology, Smithsonian Institution, Washington, D.C. 20560 During the examination of holothurians collected by the Northeast Center, National Marine Fisheries Service, Woods Hole, Massachusetts, two new species were found; they are described here. Both species will be discussed further in a report on distribution patterns of holothurians off the eastern United States (Wigley and Pawson, in prep.). I am grateful to Dr. Roland L. Wigley, National Marine Fisheries Service, for giving me access to the collections in his care, and for his help in many other ways. Type-material is deposited in the National Museum of Natural History, Smithsonian Institution. Order APODIDA Chuudotidae Chiridota Eschscholtz, 1829 Chiridota wigleyi, new species Figure IF-H Diagnosis: Wheel papillae very scarce, apparently restricted to dorsal interradii. Radial and interradial areas of body wall with numerous curved rods with bifurcated ends; rods average 63 /xm in length. Ten- tacles with curved rods with branching ends; tentacle rods average 85 ^m in length. Material examined: Holotype (USNM E15904, 40 mm total length) Delaware cruise 62-7, station 24, 15 June 1962, 40°20'N, 70°15'W, 90 m, silty sand, bottom temperature 7.8° C. Paratypes 20 specimens (USNM E 15905, 25-114 mm total length) same locality data as holotype. 34— Proc. Biol. Soc. Wash., Vol. 89, 1976 (405) 406 Proceedings of the Biological Society of Washington The species was collected at 20 stations off the eastern United States, in an area bordered by latitudes 39°47'N and 40°30'N and longitudes 69°31'W and 71°46'W; bathymetric range 70-301 m; bot- tom temperature 6.6-11.6° C; bottom type sand to sandy silt to sand- silt-clay. Etymology: The species is named for Dr. Roland L. Wigley, in recognition of his many contributions to our knowledge of the marine fauna of the northwestern Atlantic. Description: Total length 25-114 mm. All specimens contracted to varying degrees, cylindrical, dark reddish-brown in alcohol. Tentacles 12 with 5-7 pairs of digits. Ossicles in body wall include wheels and curved rods. Wheels aggregated loosely into poorly defined papillae which are sparsely scattered in dorsal interradii. In one specimen of 100 mm total length only 4 wheel papillae present, containing 28, 40, 18 and 17 typical chiridotid wheels averaging 86 fim in diameter. Curved rods with bifurcated ends (Fig. IH) scattered in radial and interradial areas of body wall. Average length of rods 63 /xm (range 50-80 jum; standard deviation 3.73). Radial longitudinal muscles con- tain numerous elongate miliary granules which vary greatly in length up to a maximum of approximately 100 ^m. Granules approximately cylindrical, often slightly thickened near center (Fig. IF). Tentacle stems and digits contain rods (Fig. IG) resembling those of body wall, but with more complex terminal branches and greater variation in size. Average length of tentacle rods 85 /tm (range 55-115 fim; standard deviation 8.7). Remarks: In possessing numerous rods in the body wall, this new species is immediately distinguished from other temperate North At- lantic and Arctic species Chiridota laevis (Fabricius), C. pellucida (Vahl), C. spiroiirna Heding, C. groenlandica Heding and C. ahyssi- cola von Marenzeller, all of which lack such rods ( Heding 1928, 1935 ) . The only other known North Atlantic species, C rotifera (Pourtales), has smaller rods in the body wall (length averaging 30 /tm), and also has very numerous wheel papillae scattered in all radii. C rotifera is tropical, occurring in shallow water in Bermuda and the West Indies (Clark, 1933). Order DENDROCHIROTIDA CUCUMARHDAE Ocnus Forbes, 1841 Ocnus diomedeae, new species Figure lA-E Diagnosis: Ossicles in body wall rudimentary cups comprising a pri- mary cross with projecting knobs and averaging 41 nm in length, over- lying knobbed plates of one type which are highly variable in shape and size; average length of plates 72 fim. Tube feet contain elongate smooth or knobbed plates. New eastern U.S. sea cucumbers 407 Fig. lA-E. Ocnus diomedeae n. sp. A, Plates from tentacles; B, Os- sicles from tube feet; C, Cups from body wall; D, One radial piece and two interradials from calcareous ring; E, Plates from body wall. F-H, Chiridota wigleyi n. sp. F, Miliary granules from radial muscles; G, Rods from tentacles; H, Rods from body wall. Material examined: Holotype (USNM E 15906, 15 mm total length) Albatross iv cruise 66-9, station 1004, 15 July 1966, 42°11'N, 65°50'W, 247 m, gravel. Paratypes: 19 specimens (USNM E15907, 8-17 mm total length) same locality data as holotype; 5 specimens (USNM E 15908, 9-19 mm total length) Albatross iv cruise 66-9, station 1005, 16 July 1966, 42°13'N, 65°42'W, 229 m, gravel; 12 specimens (USNM E 15909, 10-17 mm total length) Albatross iv cruise 68-12, station 72, 15 August 1968, 42°22'N, 65°55'W, 192 m, gravel. Etymology: Species named for collecting vessel. Diomedea is generic name for the albatross. 408 Proceedings of the Biological Society of Washington Description: Body 2-3 times as long as broad, mouth and anus ter- minal. Total length 8-19 mm. Body wall thin, stiff, packed with ossicles. Tube feet restricted to radii, in 5 more or less double rows. One or 2 feet may be present in mid-dorsal interradius. Tube feet apparently only partly retractile. Tentacles 10, richly branched; ventral pair of tentacles smaller than others. Body and tube feet light orange to white in alcohol; tentacles light yellow. Calcareous ring simple (Fig. ID) with undulating posterior margin. Retractor muscles attach to radial muscles at about middle of body. Polian vesicle single. Respiratory trees well developed, extending to anterior of body cavity. Gonad a tuft of several caeca at about mid- body. In females caeca contain yolky eggs approximately 700 fira in diameter. Ossicles in body wall knobbed plates and rudimentary cups. Plates ( Fig. IE ) highly variable in shape and size, averaging 72 fim in length (range 120-300 /im; standard deviation 14.64). Basic plate apparently 4-holed, but very few 4-holed examples seen. Cups (Fig. IC) minute, averaging 41 fim in length (range 38-44 fim; standard deviation 1.81), comprising central rod with terminal bifurcation (primary cross pat- tern), each furca carrying small knobs. Tube feet with rudimentary end plates; walls of feet contain nu- merous elongate smooth or knobbed plates (Fig. IB) which are highly variable in shape and size. Tentacles packed mth curved perforated rods and small perforated plates (Fig. lA); plates and rods usually smooth, or with few small knobs. Remarks: Rowe (1970) erected the new genera Aslia and Pawsonia to accommodate some species which had originally been referred by Panning (1949) to Ludwigia (see Pawson, 1963). Rowe concluded also tliat some of the species which had originally been assembled under the preoccupied genus-name Ludwigia should provisionally be re- ferred to Ocnus. These species included planci (Brandt), lactea (Forbes), glacialis (Ljungman), hedingi (Panning) and some others, and it is with this group of species, and therefore with the genus Ocnus, that O. diomedeae appears to have its strongest relationships. In possessing rudimentary cups, O. diomedeae is immediately distin- guished from most Ocnt/s-species. O. hedingi (Panning), O. syracusanus (Grube) and O. glacialis (Ljungman) have similarly reduced cups, but in these species the knobbed plates in the body wall are either more complex or are of a different type, and the ossicles in the tube feet are distinctly different. Literature Cited Clark, H. L. 1933. A handbook of the littoral echinoderms of Porto Rico and the other West Indian islands. Sci. Surv. Porto Rico Virgin Islands N. Y. 16( 1 ) : 1-60. Heding, S. G. 1928. Synaptidae. Vidensk. Medd. dansk naturh. Foren. Kbh. 85:105-323. New eastern U.S. sea cucumbers 409 . 1935. Holothurioidea. Part I. Apoda, Molpadioidea, Gephy- rothurioidea. Danish Ingolf-Exped. 4(9): 1-84. Panning, A. 1949. Versuch einer Neuordnung der Familie Cucumari- idae (Holothurioidea, Dendrochirota ) . Zool. Jb. 78(4) :404- 470. Pawson, D. L. 1963. The holothurian fauna of Cook Strait, New Zealand. Zool. Pubis. Victoria Univ. Coll. 36:1-38. RowE, F. W. E. 1970. A note on the British species of cucumarians, involving the erection of two new nominal genera. Jour. Mar. Biol. Ass. U.K. 50:683-687. 410 Proceedings of the Biological Society of Washington QH 1 5, pp. 411-420 12 October 1976 B4X ^^^ PROCEEDINGS OF THE u.wlOGICAL society of WASHINGTON A RECLASSIFICATION OF IRIS SPECIES BEARING ' o c/ ARILLATE SEEDS - 3 "^ I ^ ^^> -: By John J. Taylor 't o ^'' University of Montana, Missoula, Montana 59801 | -i: g. °^> .<''■' ~^': ■-'' In his revision ot the genus Iris L., Rodionenko (1961) placed in the subgenus Iris only those species with aggrega- / tions of multicellular hairs (beards) on the outer perianth ' segments. Beardless rhizomatous and nonrhizomatous species were placed in other subgenera, or were transferred to genera other than Iris. In his system, subgenus Iris consisted of sec- tion Hexapogon emended to include all species with arillate seeds, and section Iris containing the remaining non-arillate species. Hexapogon consisted of the subsections Regelia, Pseudoregelia and Oncocyclus. Regelia, however, had the same circumscription as subsection Hexapogon (Bunge) Bentham emend. Lawrence (1953), and Rodionenko's use of the name Regelia was superfluous (see also under section Hexapogon below ) . Data pubhshed since the Rodionenko revision of the genus make even more obvious the heterogeneous nature of sub- section Hexapogon (Bunge) Bentham emend. Lawrence and its synonym Regelia ( Dykes ) Rodionenko. The following new system for the arillate species in Hexapogon is proposed. Iris subgenus Iris Section Iris (This type section of the subgenus lacks aril- late species.) Section Hexapogon (Bunge) Baker Section Regelia Lynch Section Oncocyclus ( Siemssen ) Baker Section Psammiris (Spach) J. Taylor Section Pseudoregelia Dykes 35— Proc. Biol. Soc. Wash., Vol. 89, 1976 (411) 412 Proceedings of the Biological Society of Washington The essential features of this reclassification are (1) the segregation of species previously included in section Hexa- pogon (Bunge) Baker emend. Rodionenko (1961) into five distinct taxa, and (2) the designation of a type of the section Regelia Lynch. A compendium of literature relevant to sec- tion Iris is also included. Iris subgenus Iris Iris sect. Pogiris Tausch in Hort. Canalius, 1 ( 1823 ) ; idem in Sehultes Additamentum Mantissum, 2:369 (1824), quoad typus. — Iris subgen. Euiris Alefeld in Bot. Zeit., 21:296 (1863), p.p.; non Boissier Fl. Orient., 5:118 (1884); non Klatt in Linnaea, 33:591-604 (1866).— 7m sect. Euiris Bentham in Bentham and Hooker, Gen. PI., 3:687 (1882). — Iris sect. Pogiris Tausch emend. Lawrence in Gent. Herb., 8(4):353 (1953). — Iris sect. Iris emend. Lawrence in Gard. Irises, 142 (1959). Type of the subgenus : 7. germanica L. Species composing subgenus Iris are characterized by distinctly rhi- zomatous stems, and flowers with beards of multicellular hairs on the outer, and occasionally the inner, perianth segments. Key to the Sections of Subgenus Iris 1. Seed without an aril; capsule usually dehiscent very near apex, but if dehiscent considerably below apex, then some internal placen- tation ruptured at maturity Section 7m 1. Seed with conspicuous creamy- white aril; capsule dehiscent sub- apically or laterally; mature placentation intact 2 2. Both inner and outer perianth segments with conspicuous more or less linear beards 3 3. Spathe valves (bracts) 3 or 4; rhizome usually compact, slowly creeping; seed with small aril; chromosome number of counted species: n = 9 Section Hexapogon 3. Spathe valves 2; rhizome stoloniferous, readily spreading; seed with large aril; chromosome number of counted species n = 1 1 Section Regelia 2. Only outer perianth segments bearded, or if all segments bearded, then beards on outer segments scattered 4 4. Beard on outer segments scattered; spathe 1-flowered; seed with large aril Section Oncocyclus 4. Beard on outer segments linear; spathe usually 2-flowered; seed with aril much smaller than seed 5 5. Rliizome creeping or spreading by stolons; flowers essentially monochromic though sometimes lightly veined Section Psammiris Reclassification of arillate Iris 413 5. Rhizome compact, gnarled; flowers usually dichromic, spotted Section Pseudoregelia Iris section 7m Iris subgen. Pogiris Tausch, I.e.; idem in Schultes, I.e. — Iris subgen. Pogoniris Spach in Ann. Sci. Nat. set. 3, 5:103 (1846); idem in Hist. Nat. Veg., 13:48-68 (1846), p.p.; Klatt in Bot. Zeit., 30:515 (1872), p.p.; Baker in Joum. Linn. Soc. Lon. Bot., 16:143 (1877), p.p.; Randolph in Bull. Amer. Iris Soc, 109:4 (1948), p.p. — Iris subgen. Euiris Alefeld in Bot. Zeit., 21:296 (1863), p.p.; non Boissier, I.e.; non Klatt, I.e. — Iris subsect. Pogoniris (Spach) Bentham in Bentham and Hooker, I.e.; Pax in Engler and Prantl, Nat. Pflanzenfam., 2(5):145 (1888), p.p.; Lawrence, I.e. — 7m sect. Pogoniris Baker, I.e., p.p. — 7ns subgen. Eupogoniris Randolph, I.e. Type of the section: 7. germaniea L. In his reclassification of Iris, Lawrence ( 1953, 1959 ) divided sub- section Pogoniris into the series Pumilae and Elatae. To the former were assigned those species which are less than 3 dm tall, and either acaulescent or if caulescent, then not branched. Series Elatae com- prised the taller species which are distinctly caulescent and branched. Although he cited these taxa, Rodionenko did not include them in his system. Recent collections of bearded irises have included forms which could justify emending series Pumilae to include the low-growing species which are nevertheless caulescent and branched, e.g., 7. fureata MB, 7. timofejewii Woron., and perhaps others. Anyone systematically treat- ing these non-arillate species should include such emendation if the taxon is retained. Because of the numerous morphological, karyological, genetic and distributional differences between the arillate and the non-arillate spe- cies, Rodionenko ( 1961 ) removed arillate species from section 7m, based on 7. germaniea L., and gave them sectional rank collectively in Hexapogon. His circumscription of section Iris is retained here. Iris section Hexapogon (Bunge) Baker in Gard. Chron. ser. 3, 5:787- 788 (1876); Boissier, Fl. Orient, 5:119 (1884). 7m subgen. Hexapogon Bunge ex Alefeld in Bot. Zeit., 21:296 (1863), nom. illegit.; Baker in Journ. Linn. Soc. Lon. Bot., 16:147 (1877), nom. illegit. — 7m subsect. Hexapogon (Bunge) Bentham in Bentham and Hooker, Gen. PL, 3:687 (1882); Pax in Engler and Prantl, Nat. Pflanzenfam., 2 ( 5 ) : 145 ( 1888 ) . Type of the section: 7. faleifolia Bunge. Species included within the section: 7. faleifolia Bunge 1847 7. longiscapa Ledebour 1853 414 Proceedings of the Biological Society of Washington The name Hexapogon, first used by Bunge ( 1847 ) in describing the new species /. falcifolia and 7. filifolia Bunge (an invalid synonym of I. longiscapa Ledeb. ) , was taken up by Alef eld ( 1863 ) for a sub- genus circumscribing the two Bunge species and I. susiana L. Because I. susiana was the type for Spach's earlier subgenus Susiana (1846), Alefeld's use of Hexapagon was illegitimate (Article 63, International Code of Botanical Nomenclature (ICBN), 1969). Baker (1876) trans- ferred I. susiana to section Oncocyclus and first validly published Hexapogon as a section comprising 7. falcifolia and 7. filifolia Bunge only. ( The following year ( 1877 ) he raised Hexapogon to the rank of subgenus, but his use of the name at that rank was illegitimate (Article 64, ICBN).) In 1892, Baker transferred the Hexapogon species to subgenus Pogoniris, and introduced die name Regelia for a subgenus containing three species described by Eduard Kegel. Although it is apparent that Baker intended to segregate Regelia from Hexapogon and Pogon- iris, he did not adequately characterize Regelia, and it remained a nomen nudum until validly published as a section by Lynch in 1904, A type was not designated. Dykes considered 7. falcifolia and 7. filifolia Bunge to be synonymous, and transferred them (1913) from Pogoniris to section Regelia. Be- cause one or the other of these Hexapogon species must be considered the type for the earlier section Hexapogon (Bunge) Baker, Dykes' use of the name Regelia was illegitimate (Article 63, ICBN). Rodion- enko's use of Regelia for a subsection ( 1961 ) was illegitimate for the same reason. Lawrence did not acknowledge section Regelia Lynch, and combined Regelia Dykes, including Hexapogon, with Psammiris species in subsection Hexapogon (1953, 1959). 7. falcifolia and 7. longiscapa are xeritic species of the Turkmenian and Uzbekian deserts in southcentral U.S.S.R., and of a few similar but restricted locahties in Iran and Afghanistan. They are characterized by weakly or non-stoloniferous rhizomes, very narrow leaves, thin and leafless stems, and spathes of 3 or 4 bracts enclosing 2 to 5 small flowers with all perianth segments bearded. 7m longiscapa has been examined karyologically (Randolph and Mitra, 1961), and is the only bearded iris species yet counted with the diploid chromosome number 18. 7m section Regelia Lynch, Bk. Iris, 56 ( 1904 ) ; Fedtschenko in Ko- marov, Fl. USSR, 4:539 (1935); nee Dykes, Gen. Iris, 123 (1913), p.p., nom. illegit. Iris subgen. Regelia M. Foster ex Baker, Handbk. Irideae, 1 (1892), p.p., nom. nudum. — Iris subgen. Pogoniris Randolph in Bull. Amer. Iris Soc, 109:4 (1948), p.p. affin. sect. Regelia Lynch; non Spach (1846), I.e.; non Baker (1876), I.e. — Iris subsect. Regelia (Dykes) Rodionenko, Gen. Iris L., 198-199 (1961), p.p., nom. illegit. Reclassification of arillate Iris 415 Type of the section: 7. korolkowii Regel. Species included within the sectiton: I. afghanica Wendelbo 1972 7. darwasica Regel 1884 7. hewed Grey- Wilson & Mathew 1974 7. hoogiana Dykes 1916 7. korolkowii Regel 1873 7. kuschkensis Grey- Wilson & Mathew, ? edit. I. lineata M. Foster 1887 7. stolonifera Maximowicz 1880 To my present knowledge, a type of section Regelia as circumscribed by Lynch has not previously been designated. After a study of that author's protologue, and in an attempt to preserve both the originally intended and the current usage of Regelia (Article 7 B, ICBN), I have selected and here designated 7. korolkowii Regel as the type of this section. The Regelia species are essentially montane species characterized by more or less stoloniferous rhizomes bearing unbranched scapes each with a 2-bracted spathe containing 2 (rarely 3) flowers with all perianth segments bearded. They differ from all other arillate sections except Oncocyclus, with which they show the greatest affinity, in general distribution, plant habit, spathe and/or floral morphology, and, except for chromosome number in Psammiris and Pseudoregelia, in karyotype. Intersectional hybrids between Regelia and the remain- ing arillate sections (except Oncocyclus) have been difficult or im- possible to obtain. The rare hybrids produced have been sterile. In contrast, there is marked fertility in intersectional diploid hybrids between Regelia and Oncocyclus, and, excepting the large metacentric chromosome of Regelia species, a striking similarity of haploid karyo- types in the two sections. The taxa are differentiated, however, by the weakly or non-stoloniferous rhizomes, 1-flowered spathes, and widely scattered beards on the outer perianth segments of Oncocyclus species. 7m section Oncocyclus (Siemssen) Baker in Gard. Chron. ser. 3, 5:788 (1876). Oncocyclus Siemssen in Bot. Zeit., 4:706-707 (1846). — 7m subgen. Susiana Spach in Ann. Sci. Nat. ser. 3, 5:110 (1846); idem in Hist. Nat. Veg., 13:70-71 (1846). — 7m subgen. Oncocyclus (Siemssen) Alefeld in Bot. Zeit., 21:296 (1863); Baker in Joum. Linn. Soc. Lon. Bot., 16:142 (1877). — 7m subsect. Oncocyclus (Siemssen) Bentham (1882), I.e.; Pax (1888), I.e. — 7m subgen. Pogoniris Randolph in Bull. Amer. Iris Soc, 109:4 (1948), p.p. affin. gen. Oncocyclus Siemssen; non Spach (1846), I.e.; non Baker (1876), I.e. Type of the section: 7. paradoxa Steven. 416 Proceedings of the Biological Society of Washington Species included within the section: /. acutiloha C. A. Meyer 1831 I. antilibanotica Dinsmore 1933 7. atrofusca Baker 1894 I. atropurpurea Baker 1889 7. auranitica Dinsmore 1933 7. barnumiae M. Foster & Baker 1888 7. biggeri Dinsmore 1933 7. bismarckiana (Dammann) Kegel 1890 7. bostrensis Mouterde 1954 7. camillae Grossheim 1950 7. calcaria Dinsmore inedit. I. cedretii Dinsmore ex Chaudhaiy 1972 7. damascena Mouterde 1967 7. demawendica Bornmueller 1902 7. ewbankiana M. Foster 1901 7. gatesii M. Foster 1890 7. grossheimii Woronov 1928, ? hyb. nat. I. hauranensis Dinsmore 1933 7. haynei (Baker) Mallet 1904 7. hetjlandiana Boissier & Reuter 1877 7. hermona Dinsmore 1933 7. iberica Hoffmann 1808 7. kirkwoodii Chaudhary 1972 7. lineolata ( Trautvetter ) Grossheim 1950 7. lortetii Barbey 1881 7. lupina M. Foster 1887 7. lycotis Woronov 1915 7. maculata Baker 1876 7. manissadjanii Freyn 1896 7. meda Stapf 1885 7. nigricans Dinsmore 1933 7. paradoxa Steven 1844 7. petrana Dinsmore 1933 7. polakii Stapf 1885, p.p. I. sari Schott 1876 7. schelkownikowii Fomin 1907, ? hyb. nat. I. schischkinii Grossheim 1950 7. sofarana M. Foster 1899 7. sprengeri Siehe 1904 7. susiana Linnaeus 1753 7. tirmiensis Hoog 1900 7. yebrudii Dinsmore ex Chaudhary 1972 Of the arillate irises, the section Oncocyclus is the most refractory to systematic treatment. The situation has resulted from the nimierous intraspecific phenotypic variations among populations and collections Reclassification of arillate Iris 417 of apparently valid species, the frequency of natural hybridization among sympatric species and the use of specific epithets for hybrids and their nothomorphs, and variations in interpretation and use of differentiating criteria among taxonomists. It is possible, therefore, that the list immediately above lacks the names of valid Oncocyclus species, and includes names which may be synonymous with others. It should be considered a tentative listing in lieu of more thorough examinations of putative species and hybrids, and of systematic anal- yses based on characteristics perhaps somewhat more instructive of this section than gross morphology and habitat alone. The Oncocyclus species are found in dry desert and montane habitats from tlie eastern Mediterranean coastal region east and northeast into Iran, Afghanistan and Turkmenian U.S.S.R. They are characterized by weakly or non-stoloniferous rhizomes, more or less falcate radical leaves, and a scape bearing a 2-bracted spathe containing a single flower. Both inner and outer perianth segments vary remarkably in outline among the species, from the much reduced sepals of /. para- doxa, the type for the section, to the narrow and acutely pointed seg- ments of the I. meda-I. acutiloba complex and the wide rounded perianth of /. susiana and similar cultivated species. All, however, possess a beard of multicellular hairs scattered widely and often densely along the haft and onto tlie blade of the sepal. Iris section Psammiris ( Spach ) Taylor comb. nov. Iris subgen. Psammiris Spach in Ann. Sci. Nat. ser. 3, 5:110 (1846); idem in Hist. Nat. Veg., 8:69-70 (1846). Tj^e of the section: 7. humilis Georgi. Species included within the section: 7. bloudowii Bunge ex Ledebour 1833 7. humilis Georgi 1775 ? 7. mandschurica Maximowicz 1880 7. potaninii Maxim.owicz 1880 Because there has been some reluctance to reject 7. flavissima Pallas as a later synonym for 7. humilis Georgi, a brief nomenclatural history of this species is included here. Messerschmidt found a low-growing yellow iris near the Trans- baikalian town of Ulan-Ude. Its description ("Iris humilis angusti- folia " ) was first published by Ammann ( 1739 ) after number 133, page 101. Gmelin (1747) described after his number 31 "iris foliis ensiformi- bus, caule bifloro. Tab. V. Fig. 2" and included as a synonym "Iris humilis angustif olia Mess. Amm. ( Stirp. rar. ) Ruth, post 133." Gmelin stated that his number 31 (as Ammanni) had also been ob- served by Messerschmidt in the hills near Ulan-Ude. It is obvious that Gmelin considered his number 31 to be the same as Messerschmidt- Ammann number 133. 418 Proceedings of the Biological Society of Washington Georgi ( 1775 ) again observed the iris in the southern Baikal region, and published the following description of it under the name 7. humilis: "Iris humilis, angustifolia, lutea. Messerm. Amman. (Stirp. rar. ) p. 101. Radix flavo alba fibrosa. Caulis 3. ad 6. poUicum, saepius biflorus, foliis gramineis, dupio longioribus." It is apparent that Georgi con- sidered his /. humilis to be the same as Messerschmidt-Ammann number 133 (the only "iris humilis angustifolia" on page 101), and his diag- nosis is compatible with Gmelin's figure 2, plate V. In 1776, Pallas published the name I. flavissima, and included as a synonym: "Iris foliis ensiformibus Gmel. Flor. Sibr. I. p. 31. tab. V. fig. 2. cum synon. Ammanni." Thus I. flavissima is the same as Gmelin number 31 and therefore Messerschmidt-Ammann number 133, and I. humilis Georgi, also conspecific with the latter, is the first validly published name for this species. Spach ( 1846 ) first published Psammiris as a monospecific subgenus based on 7. arenaria Waldst. & Kit., a later synonym of 7. humilis Georgi. Baker ( 1877 ) later combined psammirises with Fogoniris, where they remained until Lawrence ( 1953 ) transferred them to subsection Hexa- pogon. Although most forms of 7. humilis have more or less stolon- iferous rhizomes and radical leaves which are tinged with anthocyanin pigments at vernal emergence (characteristics sometimes assumed to show affinity with common garden forms of some Regelia species), the psammirises differ from other arillate species in karyotype, general distribution range, plant habit and/or floral and rhizome morphology. The psammiris karyotype (Simonet, 1934; Gustafsson and Wendelbo, 1975) is quite distinct among arillate irises, unlike even those of Regelia and Pseudoregelia species with similar chromosome numbers (Randolph and Mitra, 1961; Zakhariyeva and Makushenko, 1969). The few intersectional hybrids produced experimerttally between Psammiris and other arillate species are sterile. Psammirises are found in open meadows and on hillsides from south- central Europe east into transcaucasian U.S.S.R., and on exposed mountain slopes and in grasslands and shaded, dry mountain valleys from the western Altai region north and east into Mongolia, Manchuria and transbaikalian U.S.S.R. They are the most widely distributed of the arillate species, and the only arillate irises native to central Europe. Although there are some differences in flower stalk length and rhizome morphology among Psammiris species and geographical forms of species, all produce narrow radical leaves which frequently dry to leave fibrous vestiges near the rhizome, and thin scapes bearing 1 or 2 2-bracted spathes each with 1 or 2 short-lived flowers with more or less elongated perianth tubes. The withering flowers become charac- teristically helically twisted (Spach, 1846; Ugrinsky, 1922). 7m section Pseudoregelia Dykes, Gen. Iris, 129 (1913). 7m subgen. Pseudoevansia Baker, Hndbk. Irideae, 2 (1892), nom. nudum. — 7m sect. Pseudoevansia Lynch, Bk. Iris, 55 (1904), nom. Reclassification of arillate Iris 419 nudum. — Iris subsect. Pseudoregelia (Dykes) Lawrence in Gent. Herb., 8(4):356 (1953); Rodionenko, Gen. Iris L., 199 (1961). Type of the section: 7, kamaonensis Wallich ex D. Don. Species included within the section: I. goniocarpa Baker 1876 7. hookerana M. Foster 1887 7. kamaonensis Wallich ex D. Don 1841 ?7. leptophylla Lingelsheim 1922 7. sikkimensis Dykes 1908, ? hyb. not. I. tigridia Bunge ex Ledebour 1829 Baker ( 1892 ) created the subgenus Pseudoevansia to include several small central Asian species which he believed were related to the crested Evansia species. His differential diagnosis was based exclu- sively on the rudimentary crests terminating the beard of pseudoevansias, structures now known to have little taxonomic significance. Lynch ( 1904 ) retained Pseudoevansia as a section, but failed to describe the taxon further. The name remained a nomen nudum. Dykes (1913) examined Pseudoevansia, transferred some included species to other taxa, and fully characterized and published the name Pseudoregelia as a section circumscribing the remaining species. Both Lawrence ( 1953, 1959 ) and Rodionenko ( 1961 ) retained the name for a sub- section with the same circumscription. The pseudoregelias are essentially montane often alpine species found from the Indian Himalayas north and east into Nepal, Tibet, Mongolia and eastern Siberia, and west into the Altai region. They superficially resemble Psammiris species in dwarf plant habit and floral mor- phology, but differ significantly in rhizome morphology and in karyotype (Simonet, 1952). The known Pseudoregelia species exhibit no close affinities with other arillate irises. Fertile intersectional hybrids involving pseudoregelias have not been produced. LiTERATxmE Cited Alefeld, F. G. C. 1863. Ueber die Gattung Iris L. Bot. Zeit. 21: 296-298. Ammann, J. 1739. Stirpium rariorum in Imperio Rutheno sponte proventientium icones et descriptiones collectae ab loanne Ammano. Acad. Sci. St. Petersburg, 210 pp. Baker, (E.) J. G. 1876. A synopsis of the known species of Iris. Gard. Chron. ser. 3, 5:787-788. . 1877. Systema Iridacearum. Joum. Linn. Soc. Lon. Bot. 16:142-147. . 1892. Handbook of the Irideae. George Bell & Sons, Lon- don, 247 pp. Bunge, A. von. 1847. Beitrag zur Kenntniss der Flor Russlands und 420 Proceedings of the Biological Society of Washington der Steppen Central- Asiens. Mem. Sav. Etr. Petersb. 7:177- 536. Dykes, W. R. 1913. The genus Iris. University Press, Cambridge, 245 pp. Georgi, J. G. 1775. Bemerkungen einer Reise in Russichen Reiche im Jahre 1772. 1:196. Gmelin, J. G. 1747. Flora Sibirica sive historia plantarum Sibiriae. 1:31. GusTAFSsoN, M., AND P. Wendelbo. 1975. Karotype analysis and taxonomic comments on irises from SW and C Asia. Bot. Notiser. 128:208-226. Lawrence, G. H. M. 1953. A reclassification of the genus Iris. Gent. Herb. 8(4):34e-371. . 1959. Garden irises. The American Iris Society, St. Louis, 575 pp. Lynch, R. I. 1904. The book of the iris. J. Lane, London and New York, 214 pp. Pallas, P. S. 1776. Reise durch verschiedene Provinzen des rus- sichen Reichs. 3:716. Randolph, L. F., and J. Mitra. 1961. Karyotypes of Iris species indigenous to the USSR. Amer. Joum. Bot. 48:862-870. Rodionenko, G. 1961. The genus Iris L. Acad. Sci. USSR, Moscow, 215 pp. (in Russian). Regel, E. 1873. Iris korolkowii Rgl. Gartenfl. Deutsch. Russ. und der Schweiz 8:225-226, Tab. 766. Simonet, M. 1934. Nouvelles recherches cytologiques et genetique chez les Iris. Ann. Sci. Nat. Bot. 10*^ serie 16:229-383. . 1952. Nouveaux denombrements chromosomiques chez les Iris. Compt. Rend. Acad. Sci. (Paris) 235:1244-1246. Spach, E. 1846. Revisio generis Iris. Ann. Sci. Nat. ser. 3, 5:89- 111. Ugrinsky, K. a. 1922. Die gesamtart Iris flavissima Pall. Eine monographische studie. Fedde's Repert. Beiheft. 14:14. Zakhariyeva, O. I., AND L. M. Makushenko. 1969. Chromosome nimibers of monocotyledons belonging to the families Lili- aceae, Iridaceae, Amaryllidaceae and Araceae. Bot. Zhur. 54: 1213-1227 (in Russian). Qi-i 1 6, pp. 421-432 12 October 1976 B4K ^^ PROCEEDINGS OF THE / divjlOGICAL society OF WASHINGTON 7 AFFINITIES OF PARANIPHARGUS LELOUPARUM : MONOD, A BLIND ANCHIALINE AMPHIPOD (CRUSTACEA) FROM THE GALAPAGOS ISLANDS I By J. Laurens Barnard \\ Department of Invertebrate Zoology, Smithsonian Institution Washington, D.C. 20560 Paraniphargus lelouparum Monod, 1970, is the first blind hypogean amphipod known from the Galapagos Islands. A new genus is described for this species to recognize several distinctions it has from Paraniphargus. The possibility that the species has sternal gills is refuted, thereby clarifying its relationships. Terms: "Gammaridan" refers to Gammaridea in the hypo- thetical group level of Section. "Mark" ("M." refers to a position a stated distance from the proximal end on a scale of to 100. "Melitid gnathopod 1" refers to a small, mitten- form gnathopod with transverse palm, elongate wrist and pubescence on one or more of articles 4-6. "Hadziid gnatho- pod 2" refers to an enfeebled female gnathopod with elongate wrist, the palm and posterior margin of the hand confluent and both armed with sparse groups of stiff, apically bent, elongate setae. These setae are also found in melitids, such as Psammoniphargus Ruffo, but occur on the posterior margin of the hand outside the palm. Uropod 3 is described in the following terms : Dispariramus, outer and inner rami dissimilar; Aequiramus, outer and inner rami similar in length, shape and patterns of armament; Magniramus, inner ramus extending as far as outer ramus; Variramus, inner ramus not as long as outer ramus but medial margin with armaments; 36— Proc. Biol. Soc. Wash., Vol. 89, 1976 (421) ■j^ c^ 422 Proceedings of the Biological Society of Washington Parviramus, inner ramus much shorter than outer ramus and lacking medial armaments, inner ramus usually very short and scalelike. The presence of a conspicuous article 2 on the outer ramus results in a classification of dispariramus; such uropod 3 can be magniramus, variramus or parviramus; an aequiramus uropod 3 is always magniramus but a magniramus uropod 3 can be either dispariramus or aequiramus. Galapsiellus, new genus Type-species: Paianiphargus lelouparum Monod, 1970 (here desig- nated ) . Etymology : Contrived. Masculine. Diagnosis: Coxal gills 2-6, ovate, weakly pedunculate, not 2-articu- late. Sternal gills absent. Males and females almost identical except for penial processes of males and weak sausage-shaped oostegites 2-5 on female. Body subvermiform, all coxae very short, of similar length. Uropod 3 parviramus, outer ramus 1-articulate, peduncle greatly elon- gate, about as long as longest ramus on uropods 1-2 and nearly as long as outer ramus of uropod 3. Telson fully cleft, lobes apically turgid, each bearing apicomedial spine. Gnathopods of both sexes enfeebled, gnathopod 1 of melitid form, wrist elongate, anteriorly pu- bescent, hand weakly trapezoidal, palm scarcely oblique, short, article 4 swollen and pubescent. Gnathopod 2 broader and longer than gnatho- pod 1, wrist similarly elongate, not pubescent, article 4 similar, not pubescent, article 6 almost twice as long as article 6 of gnathopod 1, about 1.2 times broader, palm oblique. Palms of gnathopods sparsely setose, lacking spines except at defining corners. Wrists of gnathopods unlobate. Mandibular palp article 3 linear, bearing only E setae (apical). Lower lip with weak inner lobes. Medial setae on maxillae absent or sparse. Pleopods biramous. Urosomites free, naked, or with at most one dorsolateral setule on each side. Galapsiellus lelouparum ( Monod ) Paraniphargtis lelouparum. Monod, 1970:13-25, figs. 6—45. Description: Blind. Head almost truncate anteriorly, with weak but broad and truncate anterior lobe (less accentuated than shown by Monod). Article 2 of pereopods 5-7 unexpanded, elongate, weakly pyriform; posteroventral corners right angular (or weakly sharpened in Monod's specimens). Basofacial spine of peduncle on uropod 1 situated at Mark 45, apex of peduncle extended and proboscoid. Dactyl of maxilliped with strong apical nail (not shown by Monod). New material: Five specimens from JLB GAL 103, Isla Santa Cruz, Galapagos Islands, Academy Bay, mangrove tidepool 300 m from sea near lower bodega of Charles Darwin Research Station, tidepool of Affinities of blind Galapagos amphipod 423 anchialine variety, not connected to sea at surface but fluctuating with tidal level, 23 January 1964, collected by J. L. Barnard. Associated fauna, Ampithoe sp., Cheiriphotis megacheles (both amphipods) and palaeomonid shrimps. Observations: One specimen is a definite male, bearing small penial processes on sternite 7 of the thorax. No sternal gills are present. Pre- sumably, therefore, the sausage-shaped appendages noted by Monod on pereonites 2-5 represented brood lamellae of a female, probably attached to the coxae but appearing to Monod to be attached to the sterna. One of these had a seta, also suggesting their identity as oostegites. Monod's depiction of this species is excellent. New illustrations are therefore not required. Distribution: Isla Santa Cruz, Galapagos Islands, phreatic and an- chialine. Classification of Gammaridans Monod ( 1970 ) noted the potential affinities of Paraniphargus lelouparum as belonging to the broadly conceived groups proposed by Stephensen ( 1933 ) which included such diverse genera as Niphargus, Neoniphargus, Paraniphargus, Metaniphargus {= Hadzia), Uroctena, Austroniphargus, Melita and Crangonyx. He noted also the remarks of Schellenberg (1931) showing how close Paraniphargus lies to Melita and how Stephensen ( 1933 ) placed Paraniphargus into a subgroup containing Niphargus, Neoniphargus, Niphargopsis, and Metaniphargus (= Hadzia). These genera are now better divided into the following groups, some of which are briefly characterized (see in part, Bousfield, 1973). I. Crangonychoids ( Crangonychidae, Bousfield, 1973). A primitive superfamilial group characterized by either true sternal gills, paddle- shaped calceoli or the presence of densely packed bifid-trifid spines on the palm of male gnathopod 2. Including, for example, most fresh- water genera of Australia (Neoniphargus Stebbing, Uroctena Nicholls); South Africa {Paramelita Schellenberg); Falklafidella Schellenberg and Phreatogammarus Stebbing from Falkland and New Zealand, respec- tively; Pseudocrangonyx Akatsuka and Procrangonyx Schellenberg ( = Eocrangonyx Schellenberg) from east Asia; plus the Holarctic cran- gonyxes. Numerous other genera. II. Gammaroids (Gammaridae, Bousfield, 1973). Sternal gills ab- sent. Palm of male gnathopod 2 not densely lined with bifid or trifid spines. Coxal gill 7 present or occasionally absent in apomorphic forms otherwise derivable from gammaroids; or marked by plesiomorphic characters such as tympanic calceoli in males. Eighty-five genera, nu- merous groupings, examples, Gammarus J. C. Fabricius, Chaetogam- marus Martynov, Acanthogammarus Stebbing, Amathillina Sars, Saro- throgammarus Martynov, Pontogammarus Sowinsky, Micruropus Stebbing. 424 Proceedings of the Biological Society of Washington A. Anisogammarids. Palms of male gnathopods with peg-spines. Anisogammarus Birstein, Spinulogammarus Tzvetkova, Eogammarus Birstein, Bathyceradocus Pirlot, PMetaceradocoides Birstein and Vino- gradova (and see below). B. Mesogammarid. Apomorphic form. Mesogammarus Tzvetkova. C. Micruropus alaskensis Bousfield and Hubbard. Apomorphic (new genus, Bousfield, in prep.). D. Eoniphargus Ueno. Apomorphic form with tympanic calceoli. III. Gammarelloid Groups. One or both gnathopods scarcely or not prehensile, but coxal gill 7 usually retained. Divisible into groups characterized by Gammarellus Herbst, Cheirocratus Norman, Horn- ellia Walker, Megaluropus Hoek, Argissa Boeck, Melphidippa Boeck and Macrohectopus Stebbing. IV. Hadzioids ( Hadziidae, Karaman, 1943; Melitidae, Bousfield, 1973). Sternal gills absent. Coxal gill 7 absent. Gnathopods subchelate except for special form of female hadziid gnathopod 2. Calceoli absent. Loss of any other marker attributes showing immediate descent from Crangonychoids and Gammaroids. Probably polyphyletic and subject to further elaboration. A. Melitids, to contain Melita Leach, Eriopisa Stebbing, Dulichiella Stout, Melitoides Gurjanova, Psammoniphargus Ruffo, and several new genera. Uropod 3 dispariramus. Lower lip with inner lobes. Female gnathopod 2 with distinct palm lacking groups of bent setae. 1. Subgroup to contain Psammogammarus S. Karaman, characterized by loss of sexual dimorphism on gnathopod 2, possibly Eriopisa longimmus Stock and Nijssen to be distinguished generically by variramus uropod 3. Divergent from ancestors of Eriopisa. 2. Subgroup to contain Paraniphargus Tattersall characterized by loss of medial setation on maxillae and possibly by loss of sexual dimorphism in gnathopod 2; retaining enlarged coxae unlike sub- group 1. Derivative from Melita. 3. Subgroup to contain GaJapsiellus, new genus, characterized by re- duction of medial setation on maxillae, reduction of anterior coxae, partially mittenform gnathopod 2 lacking sexual dimor- phism. Derivative from Eriopisa. B. Hadziids, to contain Hadzia Karaman, Dulzura J. L. Barnard and new genera (in part, see Zimmerman and Barnard, in press). Uropod 3 dispariramus. Lower lip lacking inner lobes. Female gnatho- pod 2 lacking palm but dactyl closing against margin furnished with sparse groups of stiff, elongate, apically bent setae. Other genera re- moved to weckeliids and ceradocids. 1. Metacrangonyx Chevreux. Group characterized by entire telson and miniaturized uropod 3. C. Eriopisellids, to contain Eriopisella Chevreux, Netamelita J. L. Barnard, Indoniphargus Straskraba, and Microniphargus Schellenberg. Both sexes with fully mittenform gnathopods. Affinities of blind Galapagos amphipod 425 1. Bathyonyx subgroup to contain Bathyonyx Vejdovsky. Charac- terized by especially reduced maxillary spination. 2. Niphargus pulchellus Sayce. Australia. D. Salentinellids, to contain Salentinella Ruffo, Pamsalentinella Bou. Gnathopods mittenform but article 2 of pereopods 5-7 expanded and lobate. E. Austroniphargids, to contain Austroniphargus Monod and a new genus (Austroniphargus starmuhlneri Ruffo). Characterized by ham- mer-like gnathopods bearing lobular wrists. Urosomites coalesced. Pos- sibly of direct crangonychoid descent. F. Pseudoniphargids, to contain Pseudoniphargus Chevreux. Gnatho- pod 1 of melitid form, hammer-like. Gnathopod 2 enlarged, palm oblique, wrist short, scarcely lobate. Telson almost entire. G. Niphargids, to contain Niphargus Schiodte, Pontoniphargus Dan- cau, Haploginglymus Mateus and Mateus, Niphargopsis Chevreux, Niphargellus Schellenberg and Carinurella Sket. Characterized by hammer-like gnathopods resembling mittens but enlarged, or apo- morphically derived from enlarged hammer-like gnathopods lacking palmar spines, wrists unlobate. H. Ceradocopsids, to contain Ceradocopsis Shellenberg ( = Maera- cunha Stephensen), PMetaceradocoides Birstein and Vinogradov. Uro- pod 3 miniaturized, retaining conspicuous article 2 on outer ramus. I. Nuuanuids (see McKinney and Barnard, in prep.), to contain Cottesloe J. L. Barnard, Gammarella Bate, Nuuanu J. L. Barnard, and new genus. Uropod 3 miniaturized as in category 9 but article 2 of pereopod 7 broadly expanded, hatchet-shaped. J. Weckeliids, formerly hadziids, to contain Weckelia Shoemaker, AUoweckelia Holsinger and Peck, Mexiweckelia Holsinger and Minckley, Mexiweckelia particeps Holsinger and Minckley (new genus, Holsinger, in prep.). Uropod 3 aequiramus. Female gnathopod 2 enfeebled, palm distinct, lined evenly with weakly bifid spines, posterior bent setae not fully developed on palm. Inner lobes of lower lip weak. Possibly derivative from ceradocids (to follow) or directly from crangonychoids. K. Ceradocids, to contain all other fully marine gammaridan genera lacking coxal gill 7, bearing gills 2-6, bearing fully subchelate gnatho- pods, nonvermiform body, normal oostegites and pleopods, uropod 3 basically magniramus and aequiramus except in apomorphie genera. Including Paraweckelia Shoemaker, a former hadziid. 1. Ceradocins. Infra group a. Ceradocins, Ceradocus Costa, Ceradomaera Ledoyer, Paraweckelia Shoemaker, Ceradocoides Nicholls. Group IVJ weckeliids, probably descend at this point. Infra group b. Paraceradocins. Antenna 2 with articles 4-5 elongate, slightly thickened. Paraceradocus Stebbing, Quadrivisio Steb- bing. 426 Proceedings of the Biological Society of Washington Infra group c. Anelasmopus Oliveira, Elasmopoides Stebbing, Maeropsis Chevreux, Maera Leach. Infra group d. Elasmopus Costa. Infra group e. Parelasmopus Stebbing, Mallacoota J. L. Barnard, Ifalukia ]. L. Barnard, Beaudettia J. L. Barnard. 2. Paraphenisa group. Uropod 1 and uropod 3 aberrant. Para- pherusa Stebbing. 3. Maerellins. Peduncle of uropod 3 elongate. Maerella Chevreux, Jerbarnia Croker. L. Kergueleniolids, to include Kergueleniola Ruffo ( =: Kerguelenella Ruffo). Like ceradoccpsids but body vermiform. V. Bogidiellids, to include Bogidiella Hertzog, Bollegidia Ruffo. Body vermiform, coxal gills reduced to 4 pairs, brood plates with dense setae confined terminally, some brood plates geniculate. VI. Pseudingolfiella Noodt. Problems of Convergence The higher classification of gammaridean Amphipoda is fraught with difficulties, among them convergence. The problem of the third uropod in hadziids is just one of many for which morphologists alone may never have answers. For example, crangonychoids (Bousfield, 1973) are marked either by sternal gills or bifid spines densely lining one or more of the gnathopodal palms. Because loss of structure is the trend in gammaridean evolution, the loss of these gills and spines could result in a host of descendents unrecognized in other super- families. Characters frequently do not occur universally in a higher taxon of amphipod, though closely related or inclusive taxa can often be recognized by marker attributes. Some crangonychoids lose sternal gills, some lose coxal gill 7 and others lose bifid palmar spines, but if the first and last are present, or if an apomorphic species can otherwise be adjoined in an obvious evolutionary sequence, then one may recog- nize a crangonychoid. Eoniphargus Ueno, for example, a Japanese hypogean gammaroid, might have crangonychoid ancestry except that the male calceoli are tympanic, rather than paddle-shaped. By that character, Eoniphargus is also not a member of the greater hadzioids and probably should be derived from a gammaroid (greater Gam- maridae) ancestry, even though coxal gill 7 is absent. A geographic companion, Awacaris Ueno, is clearly analogous to Eoniphargus in scores of attributes, though uropod 3 is aequiramus, whereas uropod 3 of Eoniphargus is dispariramus and parviramus. Awacaris therefore appears related to the marine ceradocins and the bogidiellids with aequiramus uropod 3 but its gnathopods are far more apomorphic than those of Eoniphargus, which has the apomorphic uropod 3. However, it also shows plesiomoiphy in the almost imperceptible remnant of article 2 on the outer ramus. Uropod 3 of Phreatogammarus (crangony- choid) also is aequiramus in contrast to all of its congeners. The dis- pariramus uropod 3 greatly resembles that of Notogean syncarids and Affinities of blind Galapagos amphipod 427 could be conceived as a plesiomorphic attribute, but the aequiramus uropod 3 would have to be derived from that plesiomorphic condition by loss of article 2 on the outer ramus without concomitant reduction of the inner ramus. Most of the ceradocid marine gammaroids possess the aequiramus uropod 3 but it is also present in the weckeliids and in the bogidiellids, kergueleniolids, awacarids and phreatogammarids. The aequiramus uropod 3 also has the rami more or less equal in thickness and armaments. The greater proportion of gammaridans carry the dispariramus uropod 3, or its derivatives, with the outer ramus incipiently or fully biarticulate and the inner ramus of diverse shapes and lengths. Ancestry in taxa with severe reduction in any component of uropod 3 cannot necessarily be traced. The aequiramus uropod 3 can be shown to be ancestral to the fully parviramus stage (as seen in the marine Beaudettia J. L. Barnard) as much as can the dis- pariramus uropod 3. The probability is high that a consistent evolutionary trend in uropod 3 dominates the amphipods but much elucidation is required before this trend can be perceived. The question is very basic to the origins of Gammaridea because either the gammarid-like amphipods or the photidcorophiid amphipods such as Gammaropsis, characterized by fleshy telson and generally by aequiramus uropod 3, are presumed to be the most primitive living gammarideans. The ultimate question is whether or not the dispariramus uropod 3, associated generally with freshwater amphipods, is the more primitive and therefore signals a freshwater, Notogean, syncarid ancestry. In this evolutionary sequence one would assume that sternal gills mark the primitive state and that Phreatogammarus, isolated in New Zealand, would be the most plesio- moiphic of the aequiramus and sternobranchiate forms, perhaps antic- ipatory to invasion of the sea. On the other hand, the fleshy telson of corophioids, coupled with the typical aequiramus uropod 3 of marine gammarideans may mark the ancestral stock, in which case the dis- pariramus uropod 3 is an apomoiphic development. Affinities of Galapsiellus Galapsiellus differs from Paraniphargus in the elongate peduncle of uropod 3, the size reduction of gnathopod 2 in both sexes, and the distad position of the basofacial spine on uropod 1. Gnathopod 2 ap- proaches the mittenform-shape found in Eriopisella and its allies but is significantly larger than gnathopod 1 and the hand is weakly ex- panded apically in contrast to the eriopisellid genera. Reduction of gnathopod 2 in male gammaridan amphipods is a common generic character. It typifies the eriopisellids, another group of genera known as weckeliids and other scattered genera of the gammaridan group. This characteristic is especially prevalent in anchialine or phreatic or anoculate groups. Monod was undoubtedly correct in assuming a relationship of G. lelouparum to Paraniphargus. That genus, with two species, occurs 428 Proceedings of the Biological Society of Washington on western Pacific and Indian Ocean islands in freshwaters and pre- sumably the species are basically hypogean although the type-species, P. annandalei Tattersall, 1925, was found among matted rootlets of a jungle stream at 152-244 m altitude in the South Andaman Islands, whereas the second known species, P. ruttneri Schellenberg, 1931, was found in a well in east Java. Faraniphargus has the primitive gnathopod 2 exhibited by con- tiguous marine amphipods such as Melita, an enlarged appendage with short and weakly lobate wrist, and has the primitive uropod 3 with shortened peduncle. Paraniphargus can be derived from a widely distributed tropical and temperate marine genus, Melita, with more than 50 species, often found in estuaries or in anchialine situations. Paraniphargus differs from Melita in the complete loss of medial setation on the maxillae. Paraniphargus is not well-known but one would suspect that it is also characterized by a loss of sexual dimorphism in gnathopod 2, which in Paraniphargus is a blend between male and female conditions of Melita. The loss of article 2 on the outer ramus of uropod 3 and the loss of all but E setae on mandibular palp article 3 are but ex- tensions of conditions almost fully expressed in several marine species of Melita. Paraniphargus maintains the fleshy inner lobes typical of Indo-Pacific species of Melita. Galapsiellus carries the trends of reduc- tion and sexual stabilization in gnathopod 2 to an extreme almost typical of eriopisellids. This group of genera, containing Eriopisella Chevreux, Netamelita J. L. Barnard, Indoniphargus Straskraba and Microniphargus Schellenberg, is characterized by fully mittenform gnathopods. Gnathopod 1 retains the melitid (mittenform) form but gnathopod 2 is reduced to the same size and has some of the same characteristics, such as medial or posterior pubescence and elongate wrist. In Microniphargus and Indoniphargus the gnathopods are axially reversed in contrast to Eriopisella as marked by the presence of a pubescent posterior lobe on the wrist of gnathopod 1, which in Eriopis- ella occurs on gnathopod 2. Eriopisellids also tend to be blind and anchialine and some of them, Microniphargus and Indoniphargus, have penetrated fully into phreatic waters of Belgium and India. Galapsiellus is not on eriopisellid because its gnathopods are some- what more primitive, though, like Paraniphargus, it could lie ancestral to eriopisellids. This group and the eriopisellids could also lie ancestral to niphargids. Galapsiellus bears a remarkable resemblance to the enigmatic Bathy- onyx Vejdovsky, 1905, from Lough Mask, a lake of Ireland. The body of Galapsiellus is more vermiform but the odd shape of the anterior coxae, weakly sinuous posteroventrally, is similar. In Galapsiellus much of this sinuosity can actually be eliminated by pressing the coxae very flat. The large, almost truncate head of Bathyonyx is very similar to that of Galapsiellus. Bathyonyx also has mittenform gnathopods with elongate wrists but they are closer to the eriopisellid kind than to Affinities of blind Galapagos amphipod 429 the galapsiellin kind because they are identical in size or gnathopod 1 is very slightly tlie larger. Bathyonyx is characterized by the retention of medial setae on maxilla 2 (maxilla 1 inner plate is unknown), the outer plate of maxilla 1 has the spines reduced to 4, uropod 3 is parviramus but the peduncle is short and the outer ramus is 2-ar- ticulate, the mandibular palp article 3 is more tumid and setose, and the telson is elongate. Geographically and temporally Galapsiellus bears no relationship to crangonychoids, though apomorphic crangonychoids would be dif- ficult to detect once the sternal gills, coxal gill 7, urosomal setation and bifid palmar spines were lost. Crangonychoid distribution appears to have been a product of Pangaea in the early Mesozoic. Crangony- choids are strictly of freshwater provenance and probably were widely distributed over the coalesced continental masses. They have survived primarily in Notogea, South Africa, and Nearctica with an outpost in the Falkland Islands. In Palearctica they were largely replaced by modern gammaroid genera but never reached Neotropica. There is no nearby source of crangonychoids to postulate an origin for Galapsiellus and the Galapagos Islands are too youthful to retain relicts of Pangaea. Galapsiellus can also be derived from the widespread marine genus Eriopisa. The similarity between these genera is even greater than between Galapsiellus and Paraniphargus, because Eriopisa possesses the shortened anterior coxae not typical of Paraniphargus. Many species of Eriopisa have female gnathopod 2, and occasionally male gnathopod 2, more strongly reduced than in Paraniphargus, although none of them has gnathopod 2 as enfeebled as in Galapsiellus. Eriopisa is more plesiomorphic than either Paraniphargus or Galapsiellus in maxil- lae and uropod 3 and therefore could be ancestral to Galapsiellus but not to Paraniphargus. Uropod 3 of Eriopisa bears a moderately to well-developed article 2 and the medial margins of one or both maxillae bear setae. Eriopisa may not be directly descendent from ancestors like Melita because uropod 3 of the most primitive species of Eriopisa, E. longiramus Stock and Nijssen, is magniramus, a con- dition plesiomorphic to the parviramus uropod 3 of Melita. Eriopisa longiramus and E. caeca (S. Karaman) should be reestablished in Psammogammarus S. Karaman and differentiated from Eriopisa by loss of sexual dimorphism in gnathopod 2 and the evenness of spination on the palm of gnathopod 2 in the female. Eriopisa longiramus may fur- ther be distinguished generically as uropod 3 is almost magniramus, like Pontoniphargus Dancau. Galapsiellus bears a resemblance to the Maerella subgroup of the Ceradocus group because of the elongate peduncle on uropod 3 and the shape of the telson. Male gnathopod 2 of Jerbarnia in that group has undergone an elongation reminiscent of Galapsiellus but otherwise the number of evolutionary steps between Jerbarnia (Micronesian marine) and Galapsiellus is far greater than between Eriopisa and Galapsiellus. 430 Proceedings of the Biological Society of Washington The occurrence of a phreatic amphipod in the Galapagos is striking because of the isolation of the archipelago from the mainstream of gammaridean dispersal and evolution. Nearby South America has only Ingolfiella Hansen (suborder Ingolfiellidea ) and Pseudingolfiella Noodt (suborder Gammaridea) occurring in its phreatic waters. Galapsiellus is very remote from those taxa and clearly has a marine origin. In the Galapagos Islands, Galapsiellus has been collected (Monod and herein) in brackish anchialine vi'aters presumed to be a mixture of intruding seawater and phreatic freshwaters percolating downslope to the sea from the highlands of Santa Cruz Island. Brackish sink- holes and emergent aquifers in mangrove swamps are often sufficiently fresh to be potable (5 ppt) and are so used by inhabitants of the island (pers. observ. ). The presence of Ampithoe and Cheiriphotis in my sample suggests that Galapsiellus lived in far saltier water than 5 ppt as those genera are strictly marine. The mangrove pond I sampled may actually be a stratified pool of differing salinities so that my broadly cast sample may have covered several salinity regimes. If Galapsiellus is an emergent phreatic genus, the specimens of Gala- psiellus may actually have been dying of exposure to high salinities as I caught them. On the other hand, the genus may be euryhalinic and this may help to explain its immigrational adaptability from the Conclusion Galapsiellus is considered to be an apomorphic melitid with the same kind of phreatiform adaptations found in the eriopisellids and niphargids. The best ancestral fit lies near Eriopisa although close morphological similarity occurs with Paraniphargus. Acknowledgments I am indebted to J. H. Stock, E. L. Bousfield, J. R. Holsinger, and T. E. Bowman for help with but not necessarily full agreement on the classificatory system outlined herein. Literature Cited Bousfield, E. L. 1973. Shallow-water Gammaridean Amphipoda of New England. Comstock Publishing Associates, Cornell Uni- versity Press, Ithaca, 312 p. Karaman, S. 1943. Die unterirdischen Amphipoden Sudserbiens. Srpska Kral'evska Akademiia, Posebna Izdan'a, Prirodn'achki i Mathematichki Spici, Okhridski Zbornik 34:161-313, figures 1-215. Monod, Th. 1970. V. Sur Quelques Crustaces Malacostraces des lies Galapagos .... Mission Zoologique Beige lies Galapagos et en Ecuador (N. et J. Leleup, 1964-1965) 2:11-53, figures 1-104. Affinities of blind Galapagos amphipod 431 ScHELLENBERG, A. 1931. Amphipoden der Sunda-Expeditionen Thiene- mann und Rensch. Archiv Hydrobiologie, Supplement 8:493- 511, figures 1-3. Stephensen, K. 1933. Fresh- and Brackish-water Amphipoda from Bonaire, Curagao and Aruba. Zoologische Jahrbiicher, Sys- tematik 64:414-436, figures 1-8. Tattersall, W. M. 1925. Fresh water Amphipoda from the Anda- man Isles. Records Indian Museum 27:241-247, figures 1-13. Vejdovsky, Fr. 1905. Ueber einige Siisswasser-Amphipoden. III. Sit- zungsberichte Konigeliga Bohmischen Gesellschaft der Wis- senschaften, Mathematisch- Naturwissenschaftliche Classe, 1905 (28): 1-40, plates 1-2. Zimmerman, R. J., and J. L. Barnard, in press. A New Genus of Marine Hadziid (Amphipoda, Crustacea) from Bimini and Puerto Rico. Proceedings Biological Society, Washington. 432 Proceedings of the Biological Society of Washington 3 OH i-z-v '.7, pp. 433-438 12 October 1976 HH PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON tf- A NEW SPECIES OF ARICIDEA (POLYCHAETA:f/p^ ^ > PARAONIDAE) FROM FLORIDA |iS ^^ ^; By Betsy Brown \ ^ go College of Marine Studies, University of Delaware, U Lewes, Delaware 19958 \\ v. Vv A new species of the genus Aricidea (Paraonidae) from ^-^ the Indian River lagoon of the east coast of Florida is de- scribed below. Specimens were collected by Dr. David K. Young and his staff of the Smithsonian Institution's Fort Pierce Bureau (FPB). Additional specimens from the west coast of Florida, collected by Dr. John L. Taylor and Mr. Stuart L. Santos and deposited in the National Museum of Natural History (USNM), were examined and referred to the new species. I am grateful to Dr. Young for the loan of specimens from the Indian River, to Ms. Ruth Swanson, Dr. Donald Maurer, and Dr. Les Watling for useful criticism during preparation of the manuscript, and to Sandy Steele for careful typing of the final draft. I particularly wish to thank Dr. Marian H. Pettibone of the Smithsonian Institution for the loan of specimens, for considerable assistance with some of the litera- ture and systematic problems, and for critically reviewing the manuscript. Types are deposited in the Smithsonian Insti- tution (USNM). This is contribution 109, University of Dela- ware, College of Marine Studies. Paraonidae Cerruti Aricidea Webster Aricidea philbinae, new species Material examined: Holotype (USNM 53172) and one paratype (USNM 53173): Indian River lagoon adjacent to Hutchinson Island, Ft. Pierce, Florida (20°20.9'N, 80°15.7'W), on Halodule wrightii grass- flats, 1 m, in muddy sand, collected 12 April 1973. Four paratypes (USNM 53174): Indian River lagoon just N of St. Lucie Inlet, St. Lucie, 37— Proc. Biol. Soc. Wash., Vol. 89, 1976 (433) ^air-S 434 Proceedings of the Biological Society of Washington Florida (27°10.9'N, 80°10.3'W) on H. wrightii grassflats, intertidal to 1 m, in muddy sand, collected 16 November 1974. Other specimens: about 50 animals in Indian River lagoon ranging from Titusville, Florida (just N of Haulover Canal) (28°44.1'N, 80°45.5'W) to just N of St. Lucie Inlet, St. Lucie, Florida, intertidal to 1 m, collected 1974-1975, on H. wrightii grassflats, several bottom stations, D. K. Young and staff, collectors. Tampa Bay, Florida (27°47'N, 82°40'W) (USNM 48936) 26 specimens, collected September 1971, J. L. Taylor, collector. Tampa Bay, Florida (USNM 48936) 3 specimens, collected 27 October 1969, S. L. Santos, collector. Lassing Park, St. Petersburg, Florida (27°45.2'N, 82°37.7'W) (USNM 50280/1) 9 specimens, col- lected 18 April and 20 July 1970, S. L. Santos, collector. Description: Length up to 12 mm, width up to 0.5 mm, up to 77 segments, several ovigerous. The 4 ovigerous specimens ranged from 9.8-10.6 mm long and 0.36-0.44 mm wide. Body cylindrical, widest in branchial region, tapering and flattened dorsoventrally more pos- teriorly. Prostomium subtriangular (Fig. lb, c), rounded anteriorly, fused with achaetous buccal segment, forming lateral lips of ventral mouth. Posterior lip of mouth formed by anterior edge of first setiger. Median antenna clavate, short, extending to setiger 2, unequally bifid distally, with smaller subterminal process (Fig. Ic) or rarely with asymmetrical terminal process (Fig. lb). Cilia present on distal tips of bifid antenna. Nuchal slits posterolateral to median antenna. Pig- ment spots sometimes present anterior to nuchal slits, as well as on anterior region of body, particularly on dorsal surface. Branchiae IS- IS pairs, beginning on setiger 4. Branchiae widest proximally, taper- ing to rounded tips and overlapping slightly middorsally (Fig. la, c). Anterior 7-10 branchial pairs usually longer and more robust. Single small orbicular papilla posterior to and hidden by branchia on branchial segments (Fig. la). Notopodial postsetal lobes small, spherical on first 2 setigers, more elongate on third, subulate in branchial region (Fig. la, c) becoming more slender and cirriform in postbranchial region (Fig. le). Dense bundles of basally thickened and evenly tapered capillary setae in prebranchial and branchial regions, number- ing about 10 in notopodia and about 15 in neuropodia. Anterior post- branchial setigers with finer capillary setae, numbering about 5 in notopodia and 7 in neuropodia. Posterior notopodia with about 3 (2-5) slender capillaries. Modified neuropodial hooks beginning on about setiger 22 ( 19-24 ) . Neuropodia with about 5 ( 3-7 ) very slender capillaries and 5 (4-8) modified neurosetae (Fig. le). Modified neurosetae hooked or curved distally, with terminal aristae (apparently fragile, sometimes absent), with small subterminal spine on concave side (perhaps part of subterminal hood) (Fig. Id, e). Pygidium vdth pair of small anal cirri. Etymology: It gives me great pleasure to name this species for May- belline Philbin, who was an unparalleled source of inspiration prior to the preparation of this manuscript. ISIew polychaete from Florida 435 Fig. 1. Aricidea philbinae, new species: a, Parapodium from bran- chial region, posterior view; b, c. Anterior ends of 2 different speci- mens, dorsal view; d. Modified neurosetae from posterior parapodium; e, Posterior parapodium, posterior view. Distribution: East and west coasts of Florida. Intertidal to 1 m. Remarks: Aricidea philbinae is close to Aricidea Jeffrey sii (Mc- intosh) which was described by Pettibone (1965:134) for paraonids from Virginia. Cerruti first used Aricidea jeffreysii (Mcintosh) when he established the family Paraonidae. Since then, it has been used widely as A. jeffreysii sensu Cerruti by numerous authors. Strel'tsov ( 1973 ) presented a thorough revision of the family Paraonidae in which he reexamined many specimens including type material. After reviewing the holotype of Scolecolepis (?) jeffreysii Mcintosh, 1879, from the Davis Strait, he concluded (1973:106, 159) that it should 436 Proceedings of the Biological Society of Washi^igton be an indeterminable Aricidea sp. He did so because the type-specimen was a fragment of 26 anterior segments lacking the modified posterior neurosetae necessary for diagnostic species description. Therefore, Mc- intosh's name should not be used and all the subsequent records of A. jeffreysii have needed to be referred to other species. Many of them were referred by Laubier (1967:102) and Strel'tsov (1973:105) to Aricidea (Acesta) cenutii Laubier, 1966. Strel'tsov (1973:91, 159) examined 6 paratypes of Aricidea [Acesta) catherinae Laubier, 1967, from the Mediterranean ( Banyuls-sur-Mer, France). After examining specimens of each of the following species, he referred tliem to A. (Acesta) catherinae: the records of A. jeffreysii by Pettibone (1963, 1965); the records by Hartman of Aricidea lopezi (1963:38, as A. lopezi, not Berkeley and Berkeley); and Aricidea zelen- zovi Strel'tsov, 1968. From material on loan from USNM, I examined 3 paratypes of A. catherinae from France (USNM 35914) and numerous specimens identified as A. jeffreysii by Pettibone from Maine (USNM 28940), Massachusetts (USNM 28935, 31496/7) and Virginia (USNM 31498-31500). These specimens agreed for the most part with the de- scriptions of Aricidea catherinae by Laubier (1967:112) and Strel'tsov (1973:91). None of the specimens showed a bifid antenna as found on A. philhinae. The revised distribution of A. catherinae (Strel'tsov, 1973) therefore, is: Atlantic coast of North America (from Gulf of St. Lawrence to Chesapeake Bay), coast of Uruguay, Mediterranean, Barents Sea, region of Kurile Island, off Southern California, in 2 to 1929 m. Aricidea catherinae and A. philhinae agree in the following charac- ters: Median antenna is relatively short; branchiae begin on setiger 4; notopodial postsetal lobes are small, bulbous on first 2 setigers, subu- late and cirriform more posteriorly; absence of modified setae in noto- podia; neuropodial lobes lacking; modified setae found in posterior parapodia, possessing teixninal aristae and short, subterminal spines ( "hoods" ) . Median Antenna: The median antenna of A. catherinae is enlarged in the midregion by a varying amount and is thinner distally. This agrees with Laubier (1967:114, Fig. 4a, b, c) and Strel'tsov (1973: 91). Strel'tsov's (1973) illustration of the dorsal view of the median antenna might lead one to believe that it supports a secondary process. This is not tlie case, as he states in the text and as I have observed in examination of specimens. On the A. catherinae identified as A. jef- freysii by Pettibone (USNM 28935, 28940, 31496-31500) and ex- amined by me, the swelling of the midregion of the antenna was so reduced as to appear subulate. Branchiae: Strel'tsov (1973:91) found that the number of branchial pairs of A. catherinae varied with the size of the specimen, smaller specimens (0.15 mm width) with 8-12 pairs and larger ones (0.7- 0.9 mm) with up to 25 pairs. A. philhinae has 13-15 pairs of branchiae and a maximum width of 0.5 mm. New polychaete from Florida 437 Setae: The modified neurosetae of A. catherinae and A. philbinae are similar possessing terminal aristae and a short, subterminal spine (or "hood"). The "hoods," visible on the concave side, are very dif- ficult to observe. In examination of the paratypes of A. catherinae, Strel'tsov ( 1973 ) observed a short, spinelike structure below the tip of the seta on the concave side, probably corresponding to a part of the "hood." My observations of specimens borrowed from the USNM correspond with this. Distinct hoods on the modified setae were de- picted by Laubier (1967: Fig. 5a-d). The appearance on the modified neurosetae as having hoods or short, subterminal spines seems de- pendent upon the optical equipment of the observer. Strel'tsov (1972:150) documented the setal morphology of the fam- ily Paraonidae. He described 3 main groupings concerning the de- velopment of specialized setae within the family. The first group, found on the dorsal parapodial branches of the branchial and post- branchial segments, includes lyrate setae and aciculate setae bearing spines. Setae of the second and third groups are distributed on the ventral branches of the postbranchial parapodia. Group 2 contains "pseudocompound" setae, hooked setae with a subterminal spine and "hooded" setae. According to Strel'tsov's illustration (1972:90, Fig. 1) group 2 setae may be distally curved. Group 3 includes thickened setae, setae with a terminal spine, and various hook-shaped setae which lack spines. These may be either smooth or possess thin hairs on their distal ends. The classification of the setae of A. philbinae in this scheme is uncertain. They possess the character of group 3 by having a terminal spine and by being hook-shaped. There is no indication of pubescence on the setae nor do they appear thickened. However, they also possess a hood or a short subterminal spine (as discussed earlier) giving them tlie character of group 2. In Florida, A. philbinae has been collected in association with Ari- cidea fragilis Webster, 1879 and Aricidea taylori Pettibone, 1965 (Pettibone, 1965:129, 131). These 3 species differ in the following characters : A. fragilis A. taylori A. philbinae Median Antenna Subulate Clubbed Clavate, bifid Branchiae 50-60 pairs 26-29 pairs 13-15 pairs Posterior Stouter basally, Bidentate, Unidentate, Modified tapering abruptly aristae arise curved dis- Neuropodial in midregion to terminally be- tally, with short Setae capillary tips. tween teeth subterminal spine sometimes frac- (hood), terminal tured at mid- aristae point 438 Proceedings of the Biological Society of Washington Pettibone (1965:131) described the median antenna of A. taylori as "short, extending at most to first setiger, clubbed (one paratype with distal tip bifid, evidently an anomaly. . .)•" The holotype and paratype (USNM 31494/5) were examined and the terminally bifid setae and median antenna were as described. The anomalous paratype could not be located (according to Pettibone in correspondence, one paratype was sent to Dr. Laubier in exchange, the other paratype ap- parently has been lost). The relationship of its setae and antenna to those of A. philbinae could not be determined. LrrERATURE Cited Hartman, Olga. 1963. Submarine Canyons of Southern California, Part 3. Systematics: Polychaetes. Allan Hancock Pac. Ex- ped. 27(3):l-93. Laubier, Lucien. 1967. Sur quelques Aricidea (Polychetes, Para- onidae) de Banyuls-sur-Mer. Vie Milieu 18( lA) :99-132. Pettibone, Marian H. 1963. Marine Polychaete Worms of the New England Region. 1. Aphroditidae through Trochochaetidae. Bull. U.S. Nat. Mus. 227:1-356. . 1965. Two new species of Aricidea (Polychaeta, Paraonidae) from Virginia and Florida, and redescription of Aricidea fragilis Webster. Proc. Biol. Soc. Wash. 78:127-140. Strel'tsov, V. E. 1972. [On the morphology of the setae of the polychaete family Paraonidae Cerruti, 1909 (Polychaeta, Sed- entaria).] Doklady Akad. Nauk. SSSR, 202(5) : 1219-1222. [In Russian]. English translation, Proc. Acad. Sci. USSR, Biol. Sci. Sec, 202:148-150. . 1973. [Polychaete worms of the family Paraonidae Cerruti, 1909 (Polychaeta, Sedentaria).] Akad. Nauk. SSSR. Mur- manskii Morsk Biol. Inst., 1-170 [In Russian]. QH g^.^, fo. 38, pp. 439-450 12 October 1976 NH PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON OCCURRENCE OF THE CARIBBEAN STOMATOPOD, BATHYSQUILLA MICROPS, OFF HAWAII, WITH o ADDITIONAL RECORDS FOR B. MICROPS AND 1-1 B. CRASSISPINOSA HC By Raymond B. Manning and Paul Struhsaker IE: Smithsonian Institution, Washington, D.C. 20560, ,., C-' and \ ^ National Marine Fisheries Service, Honolulu, Hawaii 96812 \\^ During the sampling program of the National Marine Fish- eries Service (NMFS) aboard the NOAA research vessel Townsend Cromwell in 1972, one specimen of an unusual stomatopod was taken by trawl off Maui, Hawaiian Islands, at a depth of 731-786 m. It proved to be an adult female of the rare Bathysquilla microps (Manning, 1961), then known from only three specimens taken in 732-952 m off southeastern Florida and the Bahamas (Manning, 1969a). Bathysquilla microps was the second species of bathy- squillid to be recognized, the first being B. crassispinosa (Fukuda, 1909), originally described from Japan and subse- quently recorded from additional localities in the western Indian Ocean. It occurs in depths between 230 and 310 m. A third bathysquillid representing another genus, Indosquilla manihinei Ingle and Merrett, 1971, was described from a unique specimen taken off Cosmoledo Atoll, Indian Ocean, in a depth of 420 m. Apparently bathysquillids are restricted to moderate depths of the outer shelf and upper slope ( Fig. 2 ) . The bathysquillids live in far greater depths than the major- ity of known stomatopods. The deepest record given by Kemp (1913) in his monograph of the Indo-West-Pacific stomato- pods was 370-419 f m ( 677-767 m ) for Squilloides leptosquilla from the "Investigator" collections. Chopra (1939) described 38— Proc. Biol. Soc. Wash., Vol. 89, 1976 (439) 440 Proceedings of the Biological Society of Washington Eurysquilla sewelli from 695 m in the Gulf of Aden and re- corded two species from 1295 m in the same area. However, these two species, Gonodactylus chiragra (Fabricius) and Oratosquilla investigatoris ( Lloyd ) , are known to live in much shallower water ( especially G. chiragra, an intertidal species ) , and their occurrence at this depth is questionable. In a review of the western Atlantic stomatopods. Manning (1969a) re- corded 14 species from depths greater than 300 m, and only two of these, B. microps (732-952 m) and Squilla intermedia Bigelow ( 291-615 m ) , occurred in depths greater than 450 m. In reporting this unusual extension of range, we present here additional northwestern Atlantic records for B. microps, based on collections made since 1969 by the National Marine Fisheries Service, additional records for B. crassispinosa, and comparative diagnoses and sketches for both of these species. Terms and indices used in the accounts below have been discussed in Manning (1969a). All measurements are in milli- meters; total length (TL) is measured on the midline from the anterior margin of the rostral plate to a Hne between the apices of the submedian teeth of the telson. Station data for the Oregon II collections are on file in the Department of Invertebrate Zoology (Crustacea), National Museum of Natural History, Smithsonian Institution. All specimens are in the Smithsonian. The illustrations were prepared by Lilly King Manning. Bathysquilla crassispinosa (Fukuda, 1909) Figures la-c, 2 Lysiosquilla crassispinosa Fukuda, 1909:61, pi. 5, fig. 4. Bathysquilla crassispinosa. — Holthuis, 1967:27 [synonymy to 1967]. — Manning 1969a:95, 98.— Holthuis and Manning, 1969:fig. 350-2.— Ingle and Merrett, 1971:197. Material: Shikoku Island, Tosa Bay, Japan: 1$ TL (total length) 240 mm.— Madagascar; 18°54'S, 43°55'E; 280-310 m; A. Crosnier, leg.; 24 November 1973: 15 TL 215 mm. Diagnosis: Cornea (Fig. la) subglobular, fully pigmented, set obliquely on stalk. Rostral plate (Fig. la) with distinct median longi- tudinal groove. Antennal protopod with 2 papillae. Dactylus of claw witli 9-10 teeth. Carpus of claw with 2 dorsal spines (Fig. lb). Inter- mediate carinae prominent on thoracic and anterior 2 abdominal somites, unarmed. Third to fifth abdominal somites lacking spinules on posterior Caribbean stomatopod off Hawaii 441 Fig. 1. Front, carpus of claw, and basal prolongation of uropod of: a-c, Bathysquilla crassispinosa (Fukuda), female from Madagascar; d-f, Bathysquilla microps (Manning), female from Hawaii. 442 Proceedings of the Biological Society of Washington margin. Telson with submedian swellings anteriorly, lacking distinct dorsal submedian carinae. Proximal segment of uropodal exopod with distinct distal spine dorsally, with 10-13 movable spines laterally. Basal segment of uropod (Fig. Ic) with spine ventrally at articulation of exo- pod, endopod with inner basal spine. Remarks: Although Manning (1969a:95) suggested that the Japanese and South African populations of this species might be distinct, based on discrepancies between published figures. Ingle and Merrett (1971) compared material from the two areas and could find no differences. We observed no differences between our specimens from Japan and Mada- gascar. On our specimens of B. crassispinosa, the intennediate carinae of the eighth thoracic somite may have a minute, sharp tubercle, but the carinae are not obviously armed posteriorly as in B. microps. The posterior seg- ments of the abdomen are more heavily granulated dorsally than in B. microps. The merus and propodus of the claw are strongly inflated in large males (TL 285 mm from Tosa Bay, Japan; reported by Ingle and Merrett, 1971), and the strongly curved dactylus has an angular pro- jection as in large males of Harpiosquilla ( Manning, 1969b ) , but the antepenultimate tooth of the claw is not reduced as in B. microps. The eyes of B. crassispinosa, unlike those of B. microps and 1. mani- hinei, are relatively large and the cornea is fully pigmented. This must be a reflection of its occurrence in shallower waters than either of the other two bathysquillids. Koniai (1938:270) noted that "the ground color is of the bright orange-red very commonly found in deep-sea crustaceans. The color is especially bright on the exposed thoracic and abdominal somites partic- ularly on the anterior and posterior margins of each somite. The cara- pace, raptorial limbs and telson are relatively light in color. On the ventral side the pleopods are crimson-red." Bathysquilla crassispinosa is a large stomatopod, attaining a total length of almost 300 mm; only two other stomatopods, Lysiosquilla maculata (Fabricius) and Harpiosquilla raphidea (Fabricius), are known to grow to this size. Females of B. crassispinosa 200-297 mm long have been recorded in the literature; males appear to be only slightly smaller, with total lengths ranging from 245 to 280 mm. Our specimens have 2-3 movable and 10-12 fixed propodal spines. The propodal indices are similar to those of B. microps 076-078 in two large males examined (TL 250-285 mm) and 070 in a female 215 mm long. A feature which apparently has escaped notice in the past is the presence, in botli species of Bathysquilla, of a minute movable tooth at the inferodistal angle of the outer surface of the carpus of the claw (Figs, lb, e). So far as we are aware such spines are found in no other stomatopods. Distribution (Fig. 2): Indo-West-Pacific region, from Japan, Caribbean stomatopod off Hawaii 443 the western Indian Ocean, and off South Africa, in depths ranging from 230 to 310 m. Records include JAPAN: Sagami Sea (Fukuda, 1909, 1910); deep part of Sagami Sea (Komai, 1938); off Owase, ca. 300 m (Komai, 1927); Shikoku Island, Tosa Bay (Ingle and Merrett, 1971); Shikoku Island, Tosa Bay, 230-295 m.— MADAGASCAR: 18°54'S, 43°55' E, 280-310 m.— MOZAMBIQUE: 25n2'S, 34°04'E; 230-295 m (Ingle and Merrett, 1971).— SOUTH AFRICA: Off Natal, north of Durban, 150 fm (275 m) (Caiman, 1923; Gordon, 1929; Ingle and Merrett, 1971); off Durban, 29°42'S, 31°29'E, 132 fm (242 m) (von Bonde, 1932; Barnard, 1950; Ingle and Merrett, 1971). Bathysquilla microps (Manning, 1961) Figures Id-f, 2 Lysiosquilla microps Manning, 1961:693, fig. 5, pis. 10-11. Bathysquilla microps. — Holthuis 1967:28 [references]. — Manning 1969a: 95, Figs. 26-28. Material: Gulf of Mexico, Bay of Campeche, Mexico; 700 fm (1281 m); Oregon II Station 10957: 1 c? TL 112.5 mm.— Gulf of Mexico, south of Panama City, Florida; 395 fm (723 m); Oregon II Station 10635: 1$ TL 175 mm.— North Atlantic, off Surinam; 681-769 m; Oregon II Stations 10604, 10620, 10796: 1$ TL 187 mm, 2 5 TL 173- 220 mm.— Off French Guiana; 604-769 m; Oregon II Stations 10614, 10616, 10803, 10816, 10822: 4^ TL 176-255 mm, 2 5 TL 183-195 mm. — Pacific Ocean, near Maui, Hawaiian Islands; 21°04'N, 156°09'W; 731-786 m (400-430 fm); 12.5 m shrimp trawl; Townsend Cromwell Station 61-66; 26 October 1972: 1 5 TL 201 mm. Diagnosis: Cornea (Fig. Id) indistinctly bilobed, set very obliquely on stalk, pigmented part reduced to small, transverse bar. Rostral plate (Fig. Id) not grooved longitudinally. Antennal protopod with 2 papil- lae. Dactylus of claw with 13-15 teeth (14 in Maui specimen). Carpus of claw (Fig. le) witli 1 dorsal spine. Intermediate carinae of body prominent, armed posteriorly on eighth thoracic and usually on second to sixth abdominal somites, also on first somite in Hawaiian specimen. Third to fifth abdominal somites variously ornamented with spinules on posterior margin. Telson with distinct dorsal submedian carinae. Proximal segment of uropodal exopod unarmed dorsally, with 6-8 (6 only in Maui specimen) movable spines laterally. Basal segment of uro- pod (Fig. 1/) unaraied ventrally at articulation of exopod, endopod unarmed on inner margin. Remarks: The Hawaiian specimen of B. microps agrees in almost all respects with our material from the western Atlantic. The only major difference that we observed was that the Hawaiian specimen has armed intermediate carinae on the first abdominal somite, whereas the inter- mediate carinae of this somite are xmarmed in all of our Atlantic spec- imens. This single difference in one specimen is not, in our opinion, enough to warrant recognizing a separate Hawaiian subspecies. 444 Proceedings of the Biological Society of Washington 3 •43 3 a Caribbean stomatopod off Hawaii 445 The propodal index for five female B. microps (TL 173-220 m) ranges from 074-078; it is 074 in the female from Hawaii. Propodal indices of males of B. microps from the Atlantic (TL 68-255 mm) range from 074 to 092, the higher indices being found in the smaller specimens. The female from Hawaii has 2-3 proximal movable spines and 12-13 fixed spines on the opposable margin of the propodus of the claw. In Atlantic specimens, there are 2 movable proximal spines and 11-15 fixed spines; of 20 claws examined, 16 were armed with 13-14 teeth. The dactylus of the claw is anned with 13-15 teeth in Atlantic speci- mens, 14 in the female from Hawaii. There are 7-8 movable spines on the proximal segment of the uropod in Atlantic specimens, 6 in the female from Hawaii. Bathysquilla microps may be a slightly smaller species than B. crassispinosa which attains a total length of almost 300 mm. Males of B. microps have total lengths of 68-255 mm, and females ranging from 44.5 to 220 mm are known. The color of the female specimen from Hawaii is similar to that of Atlantic specimens, based on photographs by J. E. Randall, Bemice P. Bishop Museum, D. Opresko, University of Miami, and B. Rohr, NMFS. On the Hawaiian specimen the anterior part of the body, from about the level of the antennae and including the eyes, is white; the rostral plate, like the carapace, is pink. The proximal portion of the propodus of the claw is white, the distal portion is pink, and the dactylus is orange. The proximal half of the walking legs is whitish, the distal part is pink. The remainder of the body is pink with orange carinae on the posterior portion of the body. Atlantic specimens show a similar pattern, but specimens have been taken in two color phases, red and orange. Of the more than 300 shrimp trawl stations effected in the Hawaiian Islands by NMFS during the period 1967-72, the Bathysquilla capture occurred at the deepest successful station. Of the total of 300 stations, about 55 were in the 300-800 m depth range. Thus, it would appear that the minimum depth inhabited by Bathysquilla in the Hawaiian Island is about 750 m. At this depth the bottom temperature is approx- imately 5° C. An account of the recent NMFS surveys and the general ichthyological results are given by Struhsaker ( 1973 ) . Taxa of fishes associated with the Hawaiian capture of Bathysquilla were: sharks {Apristurus, Etmopterus); the eel families Congridae and Nettastomatidae; Halosauridae; Neoscopelidae; Ogcocephalidae; Ophid- ioidei; Macrouridae {Hymenocephalus, Malacocephalus, Matacocephalus, Nezumia, Trachonunis, Ventrifossa) ; Cynoglossidae. The ophidioid fishes represent a species previously recorded only from the western Pacific and the Indian Ocean ( Cohen, 1974 ) . Also taken in the haul were three or four species of pandalid shrimps and 25 kg of discarded military ordnance. During the 1902 expedition to the Hawaiian Islands of the U.S. Fish Commission steamer Albatross, no specimens of Bathysquilla were 446 Proceedings of the Biological Society of Washington taken. Of the 257 beam trawl stations occupied during that expedition, 88 were in the 700-2800 m depth range. Bigelow (1931) reported on the stomatopods collected during that expedition and recorded only three species which occurred on the upper slope: Echinosquilla guerinii ( White ) in 55-335 m, Odontodactyltis brevirostris ( Miers ) in 55-439 m, and O. hawaiiensis Manning (as O. japonicus) in 110-278 m. Townsley (1953) recorded E. guerinii from depths of 92-220 m; all of the other stomatopods reported by Townsley from Hawaii were taken in compara- tively shallower water. Distribution ( Fig. 2 ) : Western Atlantic, from localities neai Florida, in the Gulf of Mexico, and off northern South America, in depths be- tween 604 and 1281 m, and off Maui, Hawaiian Islands in 731-786 m. Records include: BAHAMAS: Santaren Channel (24°24'N, 80°00'W), 732-860 m (BuUis and Thompson, 1965; Manning, 1969a).— FLOR- IDA: Off Cape Canaveral (28°03'N, 78°44'W), 915-952 m, and south- east of Tortugas (24°irN, 83°21.5'W), 732 m (Manning, 1961, 1969a; BuUis and Thompson, 1965); south of Panama City (28°12'N, 86°09'W), 723 m.— MEXICO: Bahia de Campeche (2P31'N, 96°46'W), 1281 m.— SURINAM: 07°49'N, 54°22'W, 732 m; 07°47'N, 54°05'W, 769 m; 07°53'N, 54°04'W, 681-732 m.— FRENCH GUIANA: 07°37'N, 53°32' W, 723 m; 07°35'N, 53°32'W, 604-769 m; 07°32'N, 53°22'W, 641-659 m; 07°18'N, 52°59'W, 668-705 m; 07°06'N, 52°44'W, 668 m.— HA- WAII: Off Maui (21°04'N, 156°09'W), 731-786 m. General Discussion The discovery of a population in Hawaii of a species previously knoviTi from the western Atlantic raises many interesting questions, none of which can be answered with our current state of knov/ledge. Since bathysquillids represent an old stomatopod stock (see below), it seems likely that the Hawaiian and American populations are relicts of a more widely distributed species, but that the Atlantic populations colonized the Pacific or that the reverse occurred cannot be ruled out. It seems likely that two distinct populations now occur in the two areas, for gene flow between deepwater benthic populations in the western Atlantic and the central Pacific must be minimal, but if genetic differences exist they are not strongly reflected in morphological features. Too, geo- graphically intermediate populations may well exist; the faima of the upper slope is not that well known in most areas of the world. Several species or species groups of slope fishes exhibit distributions similar to tliat of B. microps. The macrourid Hymenocephalus aterrimus is known only from the tropical western Atlantic and Hawaii in depths of 450-900 m. Marshall and Iwamoto ( 1973 ) state that there are virtually no differences between the populations in the two oceans except for a tendency for a slightly greater interorbital width in the Hawaiian spec- imens. They did not feel that subspecific or other recognition of the populations were required. Iwamoto ( 1974 ) recently described a new Caribbean stomatopod off Hawaii 447 subgenus and species of macrourid presently known only from the Gulf of Mexico, the Caribbean (732-1062 m), and the Hawaiian Islands ( 623-705 m ) . He apparently did not find any morphological differences in the two populations of Nezumia { Kuronezumia) bubonis. This sub- genus also comprises a second species which occurs in the South China Sea (Iwamoto, 1974). The plectognath fish genus Hollardia is represented in the tropical western Atlantic by H. meadi (Bahamas, Cuba, and Barbados: 135- 450 m) and H. hollardi (Bermuda, Florida Keys, Gulf of Mexico, Carib- bean: 225-915 m). Tyler (1968), in describing H. goslinei from Hawaiian waters (365 m), states that it and H. hollardi are closely related and have differentiated along the same lines in several important respects, while H. meadi differentiated less from the ancestral type, re- maining more generalized than the other two species. He also discussed the peculiar distribution of the genus and speculated that the migration of the hoUardiins into what is now the tropical western Atlantic must have beeen eastward through the Pacific with goslinei being the only presently known Pacific remnant with the extinction of eastern Pacific populations after the reemergence of Central America. Finally, the Hawaiian endemic bothid flatfish Chascanopsetta prorigera was recently reported from the central western Atlantic by Gutherz ( 1967 ) on the basis of a still unpublished manuscript. Preliminary comparison of material from the two regions indicates to us that the western Atlantic population deserves recognition as a distinct species. However, these two forms are very close morphologically, and together constitute a distinctive group within the genus. Such broad distributional patterns are generally rare within the stomatopods. Of approximately 300 species now recognized, only one, Heterosquilla mccullochae (Schmitt), can be considered to be pantrop- ical, occurring in the eastern Pacific, the western Atlantic, and the Indo- West-Pacific regions. Five other species occur in both the Atlantic and the Indo-West-Pacific region but not in the eastern Pacific. Broad horizontal distribution patterns also appear to be rare, in general, in inhabitants of the upper slope (Ekman, 1953; Briggs, 1974). Briggs (1974:435) has pointed out that "many ancient, phylogenetic relicts have accumulated in the slope habitat," and docvmiented this with material recently reported in the literature on a variety of animal groups. The bathysquillids also represent one of these groups, an old stomatopod stock that occurs on the upper slope. The stomatopods are not well represented in the fossil record ( Holthuis and Manning, 1969 ) , and there is little evidence to indicate what are old or primitive charac- ters in the group. However, the primary distinction between the single fossil family, Sculdidae, and the four recent families, is the presence of an unsegmented exopod in the sculdids. As Ingle and Merrett ( 1971 ) noted, one of the characteristics of Indosquilla manihinei is its uropodal exopod, with the distal segment indistinctly sutured rather than distinctly 448 Proceedings of the Biological Society of Washington articulated as in all other recent stomatopods. It seems likely that the bathysquillids do represent an ancient stomatopod stock. Literature Cited Barnard, K. H. 1950. Descriptive list of South African stomatopod Crustacea (mantis shrimps). Ann. S. African Mus. 38:838-864, figs. 1-4. BiGELOW, R. P. 1931. Stomatopoda of the southern and eastern Pacific Ocean and the Hawaiian Islands. Bull. Mus. Comp. Zool., Harvard 72:105-191, figs. 1-10, pis. 1-2. Von Bonde, C. 1932. Report No. 9 for the year ending 1931. Rep. Fish. Mar. Biol. Surv. S. Africa 9:1-128. Briggs, J. C. 1974. Marine zoogeography. McGraw-Hill. 475 pp. Bullis, H. R., Jr., and J. R. Thompson. 1965. Collections by the exploratory fishing vessels Oregon, Silver Bay, Combat, and Pelican made during 1956 to 1960 in the southwestern North Atlantic. U.S. Fish Wildl. Serv., Spec. Sci. Rep.: Fish. 510: iii + 130 pp. Calman, W. T. 1923. Preliminary report on Crustacea procured by the S.S. "Pickle." VI. S. Africa Fish. Mar. Biol. Surv., Res. Rep. 3 (1922):1. Chopra, B. 1939. Stomatopoda. John Murray Exped., Sci. Rep. 6: 137-181, figs. 1-13. Cohen, D. M. 1974. The ophidioid fish genus Luciobrotula in the Hawaiian Islands. Pacific Sci. 28:109-110. Ekman, S. 1953. Zoogeography of the sea. Sidgwick & Jackson, London. 417 pp. FuKXTDA, T. 1909. The Stomatopoda of Japan [in Japanese]. Dobut- sugaku Zasshi 21:54-62, 4 pis. . 1910. Report on Japanese Stomatopoda, with descriptions of two new species. Annot. Zool. Japonenses 7:139-152, pi. 4. Gordon, I. 1929. Two rare stomatopods of the genus Lysiosquilla Dana. Ann. Mag. Nat. Hist. (10)4:460-462, figs. 1-2. Gutherz, E. J. 1967. Field guide to the flat fishes of the family Botliidae in the western North Atlantic. U.S. Fish Wildl. Serv., Circ. 263:iv-f47 pp. HoLTHUis, L. B. 1967. Stomatopoda I. Fam. Lysiosquillidae et Bathy- squillidae. Pages 1-28 in H.-E. Grimer, and L. B. Holthuis, eds. Crustaceorum catalogus. Pars 1. Den Haag, Dr. W. Junk N. V. , AND R. B. Manning. 1969. Stomatopoda. Pages 535-552, figs. 343-363, in R. C. Moore, ed. Treatise on invertebrate paleontology. Part R, Arthropoda 4. Ingle, R. W., and N. R. Merrett. 1971. A stomatopod crustacean from the Indian Ocean, Indiosquilla manihinei gen. et sp. nov. (Family Bathysquillidae ) with remarks on Bathysquilla crassi- Caribbean stomatopod off Hawaii 449 spinosa (Fukuda, 1910). Cmstaceana 20(2):192-198, figs. 1-9, pi. 1. IWAMOTO, T. 1974. Nezumia ( Kuronezumia ) bubonis, a new subgenus and species of grenadier ( Macrouridae : Pisces) from Hawaii and the western North Atlantic. Proc. Calif. Acad. Sci. 39(22): 507-516. 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. 4:1-217, pis. 1-10, 10 text-figs. KoMAi, T. 1927. Stomatopoda of Japan and adjacent localities. Mem. Coll. Sci. Univ. Kyoto, Ser. B 3(3):307-354, figs. 1-2, pis. 13- 14. . 1938. Stomatopoda occurring in the vicinity of Kii Peninsula. Annot. Zool. Japonenses 17:264-275, figs. 1-3. Manning, R. B. 1961. A new deep-water species of Lysiosquilla (Crustacea, Stomatopoda) from the Gulf of Mexico. Ann. Mag. Nat. Hist. (13)3:693-697, fig. 5, pis. 10-11. . 1969a. Stomatopod Crustacea of the western Atlantic. Stud. Trop. Oceanogr. 8:viii -f 380, figs. 1-91. . 1969b. A review of the genus Harpiosquilla (Crustacea, Stomatopoda), with descriptions of three new species. Smith- sonian Contrib. Zool. 36:1-41, figs. 1-43. Marshall, N. B., and T. Iwamoto. 1973. Family Macrouridae. Fishes of the western North Atlantic. Sears Found. Mar. Res. No. 1, Part 6. 698 pp. Struhsaker, p. 1973. A contribution to the systematics and ecology of Hawaiian bathyal fishes. Ph.D. thesis. University of Hawaii, Honolulu. XV 4- 482 pp. TowNSLEY, S. J. 1953. Adult and larval stomatopod crustaceans occur- ring in Hawaiian waters. Pacific Sci. 7(4) :399-437, figs. 1-28. Tyler, J. C. 1968. A monograph on plectognath fishes of the super- family Triacanthoidea. Monograph 16, Acad. Nat. Sci. Philadel- phia. 364 pp. 450 Proceedings of the Biological Society of Washington QH 1 B4X NH 3. 39, pp. 451-466 12 October 1976 PROCEEDINGS OF THE M.OLOGICAL SOCIETY OF WASHINGTON AN ANALYSIS OF PEROMYSCUS DIFFICILIS FROM THE MEXICAN-UNITED STATES BOUNDARY AREA By Victor E. Diersing \ ■ Museum of Natural History, University of Illinois at ^ U rhana-Champaign, Urbana, Illinois 61801 In April of 1964 and 1969 Woodrow W. Goodpaster, collect- ing for the Museum of Natural History, University of Illinois, obtained 15 specimens of Peromyscns of uncertain taxonomic status from the Franklin Mountains, El Paso County, Texas. Numerous mammalogists examined these 15 specimens, of- fered suggestions for possible affinities, but never were cer- tain enough to assign specific identification. Subsequent ex- amination of these and additional specimens, some of which were from the Franklin Mountains, proves that they are Peromyscus difficilis, that this species occurs in mountainous areas near the Rio Grande River, and that they are referable to a distinct subspecies. This subspecies is called Peromyscus difficilis penicillatus Mearns ( Peromyscus ])oylii penicillatus of Mearns). To establish a means of species recognition, comparative studies using 20 skin and skull features were made of samples of P. polius, P. hoylii rowleyi, P. pectoralis laceianus, P. truei triiei, P. truei comanche, P. attwateri, and P. difficilis. In addition cranial and external morphology, along with cyto- logical evidence was used in studying intrapopulation and interpopulation variation in five subspecies of P. difficilis: P. d. nasutus, P. d. griseus, P. d. penicillatus, P. d. difficilis, and P. d. petricola. Methods Over 1000 specimens representing six species of the genus Peromyscus were studied. External measurements were taken 39— Proc. Biol. Soc. Wash., Vol. 89, 1976 (451) 452 Proceedings of the Biologic^il Society of Washin , 1 -r a "1 ^ y -Tl Clj > H -D V n fl tj o c U _!_, ,. * * * * in OT t^ i^ -v 00 CO Ci CO ro 00 ^ 00 in 1—1 Cl in CO m Cl kO CO l^ i^ CO -V in rH CO i-^ 00 CO q q CO CO C5 CO 00 ■^ d d d d ci ci ci rH CO 1—1 d "' 1—! r-W d 1— H ^ ci d d 00 1^ ^H in CI CO m 1- Cl CD -t in 1—1 Cl c-i -f< CO I- t^ > CO CO CO CO CO 00 C7> a> ^H CO i- Cl q CO q i-H CD u Xt^ t-^ ci ci CO ■"^ ^^ '"' ^^ ^H ci co ci ci Tfi ci CO ■"^ ci Cl Oi oi CO CI CO o 1—1 m -f 00 CO 00 CO ^ CD G> t- Q OS Ol CO CO 1-; m -V Cl "—1 1—1 Cl r— ( 1—1 CO ^ 1—1 Cl 1—1 Cvl c/i oi CO ci d ^ d d d d d d d d d ^-^ ^ d d d d in O m o O CO 'Nf* I— 1 00 01 CO QO in Cl l^ CD i-H -f< 1—1 l>< q in in 00 O in t- 00 05 CO Cl '^ ^ CO CO CO CD 1—1 1— 1 lO f— J CO CO d t-^ d ci 1— < CO d CO -* d -^ in d d 1^ d o T— ( Gi CI cl CI CI 1—1 f— ( i-H ^^ I— 1 o^ 1—1 ^ o o O o O a> C75 o 00 Cj5 00 PI 01 Ol CI CI Cl 1—1 Cl Cl cl Cl Cl 1—1 Cl 1—1 in CO CO CO CD \~- t- CO Cl CO "^ t- Cl 00 Cl 00 ^ m CO cy> I> ■* Ol CD 00 T— 1 o CO 00 L- !>: cq t- -t ^H Oi 00 i^ 00 Cl U CO ^ Tt^ CO CO ci ci •"^ 1—1 ci ci CO ci ci -^ CO ci ci ci ci CO o t- T— I in OD o -t" ^ 00 ,_^ CO Cl CD 00 CO t- -^1 CO d ■— < 00 CO c^ CD in in Cl Cl CO 1—1 ■-H 1—1 Cl I— 1 Cl Cl Cl ■—1 d c/^ t-^ -t -r d d d d d d d d d d d d d d d d d ro l^ 1-- o t- c^ -t ^ CO G^ cl -t^ ^ 00 i> CO CO re i^ CO l^ 01 -r CO m ^ -H CZi ^H i^ CO in Tf< -t CO i^ q CO I— 1 IX CO Ol -t" CO d i>^ 1—! ci ci CO d CO Tt^ d Tfi in t~^ d ^ d o 1— { c:i CI 03 CI Cl 1— < i-H 1—1 1— 1 T— 1 Ol z lO »n in in -f ID in in m in in in in m m in in in in in ■—1 T— 1 1—1 1—1 1— 1 1—1 1— ( c -5 bC 'Xj a W) ^ bC C ^ -^ ^ ^ ^ ■^3 -^ ^. ^ tib — "cS a b£) C 1 c T3 C Is 'cS T3 0) 03 Cj _o Is bJD _cS c Cj s "cS 5c -2 c -5:1 CS C a T3 o "c3 O K W O c5 m m ^ cS 0* fil Q oi S CO Peromyscus — Mexican boiindanj area 457 ill! DA ilA ^5*5 "'■ -' |.^ (IR Jjil >u aa r)B AQ on ftO 0/"j oa a» no nn nn ft A nn «*» Fig. 3. Karyogram of P. difficilis griseus, male, UI 48397, from the Canizozo kua beds, Lineoln County, New Mexico. and females of a sample were combined in analyzing geo- graphic variation. Geographic Variation Chromosomal evidence: Species ot die genus Peromyscus arc consistent in having karyotypes ot 48 chromosomes. The autosomal complement usually consists of several pairs of large biarmed chromosomes, several pairs of small biarmed chromosomes, and a number of acrocentric chromosomes. Hsu and Arrighi ( 1968 ) described the karyotype of P. difficilis as being polymorphic. Karyotypes of P. d. nasutus from Fort Collins, Colorado, contained four pairs of large biarmed autosomes and specimens of P. d. saxicola from Jacala, Hidalgo, Mexico, contained three pairs of large bi- armed autosomes. Lee, et al. (1972) also described karyo- types of P. d. nasutus from Colorado and northwestern New Mexico with four pairs of large biarmed autosomes. 45S Froceedings of fJie Biological Society of Washington Fig. 4. Collecting localities of P. difficilis are represented by circles. Adjacent localities were usually grouped to increase sample size. Sample locality 1, Chirieahua Mountains, Arizona; 2, Springerville, Arizona; 3, Capitan, Jicarilla, and Sacramento mountains, New Mexico; 4, Carrizozo lava beds, Lincoln County, New Mexico; 5, Franklin Mountains, El Paso County, Texas; 6, Sierra del Carmen Mountains, Coahuila, Mexico; 7, vicinity Creel, Chihuahua, Mexico; 8, vicinity Sierra Guadalupe, Coahuila, and Cerra Potosi, Nuevo Leon, Mexico; 9, \'icinity Bella Union, Carneros, Sierra Encarnacion, Coahuila; and Concepcion del Ore, Zacatecas. Peromyscus — Mexican boundary area 459 30.0 298 29 6 29.4 29 2 29 288 28 6 - 28 4 28.2 - 28.0 27.8 276 - Pcf petricola Pd. difficilis Pd. nasutus Pd griseus Pd penicillatus 108 112 116 Tail Length (mm; 124 Fig. 5. Scattergram plotting mean tail length versus mean skull length in samples of P. difficilis. Numbers within the figure correspond to the locahty nmnbers of Fig. 4 and Table 2. Specimens representing two subspecies of P. difficilis were collected and karyotyped: P. d. penicillatus (Fig. 2) from the Franklin Movmtains, El Paso County, Texas, and P. d. iS^riseus ( Fig. 3 ) from the lava beds near Carrizozo, Lincoln County, New Mexico. Karyotypes of both subspecies did not differ from karyotypes of P. d. nasutus in having four pairs of large biarmed autosomes. A series of karytoypes from both subspecies may establish slight morphological differ- ences in the X chromosomes. Morphological evidence: Eleven characters were selected and geographic variation was assessed to ascertain the af- finities of each population to the Franklin Mountains' popu- lation of P. difficilis. In plotting collecting localities (Fig. 4) some localities of close proximity were grouped to in- crease sample size. 460 Proccediuii^s of the BioIo