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Ss x ~ = = Ee E SMITHSONIAN _INSTITUTION NOILALILSNI_ NVINOSHLIWS S3i1uvysg oe LIBRARIES SMITHSON z rf zZ Xs vt = ws ‘Ss e aA: : =. ih fy A B Cc D PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON E F G H I Fig. 1. The tibiotarsus of Megapodius in cranial (A-C), medial (D-F), and lateral (G—I) aspects. A, D, G, Holotype of Megapodius alimentum, new species, Lifuka, Tonga, BPBM 165686; B, E, H, M. freycinet freycinet, male, Halmahera, Northern Moluccas, USNM 557015; C, F, I, M. pritchardi, sex unknown, Niuafoou, Tonga, USNM 319633. Scale bars = 10 mm. al. (1979). Measurements were taken with dial calipers with increments of 0.05 mm, rounded to the nearest 0.1 mm. CU = Cul- tural Unit. Systematic Paleontology Class Aves Order Galliformes Family Megapodiidae Six specimens are referred to the Mega- podiidae rather than to Gallus gallus of the Phasianidae, the only other galliform in Oceania, because of the following charac- ters: coracoid—ventro-medial margin of hu- meral end of shaft more rounded, cotyla scapularis not extending beyond lateral margin of shaft in dorsal aspect, shaft (just sternal to cotyla scapularis) more rounded in cross-section; tibiotarsus—condylus me- dialis and condylus lateralis short and wide, epicondylus medialis large; tarsometatar- sus—distal half of facies dorsalis convex rather than concave, tuberculum intercon- dylarum small, tuberositas musculo tibialis cranialis large and protruding dorsad to fa- cies dorsalis, fossa metatarsi large and deep, extending beyond medial edge of facies dor- salis; pedal digit I, phalanx 1—large size, straight shaft; pedal digit II-IV, terminal phalanx—large size, dorso-ventrally com- pressed, medio-laterally expanded. Genus Megapodius Within the Megapodiidae, the fossils are referred to Megapodius (including Eulipoa, following Ripley 1960) rather than to Le- poa, Alectura, Aepypodius, Tallegalla, or Macrocephalon by the following combina- tion of characters: tibiotarsus—distal mar- gin of pons supratendineus nearly perpen- dicular (less diagonal) to the long axis of the shaft, ventral margin of condylus lateralis protrudes more ventrad from the shaft; tar- sometatarsus—shaft wide but dorso-ven- trally compressed, fossa metatarsi I distinct, deep, and circular in deeper portion, fora- men vasculare distale large; pedal digit II- IV, terminal phalanx—dorso-ventrally compressed, medio-laterally expanded. Megapodius alimentum, new species Figs. 1-3 Holotype. —Distal end of tibiotarsus, BPBM 165686, Pit ONOW, Layer IV (CU- III), Tongoleleka archeological site (To-Li), Lifuka, Haapai Group, Tonga. Tom Dye and field party Aug 1984. Paratypes.—All from Tongoleleka site. Proximal end of tarsometatarsus, BPBM 165689, Pit ONOW, Layer IV (CU-III). Tar- sometatarsus lacking both ends, BPBM 165670, Pit 45N1W, Layer IV (CU-IID. VOLUME 102, NUMBER 3 A B Cc D 339 s G ,, ’ G H E F Fig. 2. The tarsometatarsus of Megapodius in dorsal (A—D) and plantar (E—H) aspects. A, E, Holotype of M. molistructor, New Caledonia, MNHN 600; B, F, Paratypes of M. alimentum, new species, Lifuka, Tonga, BPBM 165689, 165670; C, G, M. freycinet freycinet, male, Halmahera, Northern Moluccas, USNM 557015; D, H, M. pritchardi, sex unknown, Niuafoou, Tonga, USNM 319633. Scale bar = 10 mm. Pedal digit I, phalanx 1, BPBM 165674, Pit ON11W, Layer IV pit (CU-III). Pedal digit IJ-IV (exact number uncertain), terminal phalanx, BPBM 165675, Pit ON11W, Layer IV pit (CU-IID). Diagnosis.—A large species of Megapo- dius, exceeded in size only by M. molis- tructor (Tables 1-3). Megapodius alimen- tum differs from M. freycinet as follows: tibiotarsus—incisura intercondylaris wider, tuberositas retinaculi musculo fibularis larger and more distinctly offset from con- dylus lateralis; tarsometatarsus— foramina vascularia proximalia more deeply inset be- low the dorsal surface of facies dorsalis, lat- eral margin of facies dorsalis more rounded at level of foramina vascularia proximalia; digit I, phalanx 1 —stouter; digit II-IV, ter- minal phalanx—broader medio-laterally. Megapodius alimentum differs from M. molistructor in the same characters of the tarsometatarsus, as well as in the smaller foramina vascularia proximalia and the less prominent crista plantare medialis. Mega- podius alimentum differs from M. pritchardi in the same characters of the tibiotarsus and phalanges, but not those of the tarsometa- tarsus. Etymology. — From the Latin alimentum, meaning “‘food.’”’ The name alimentum re- fers to the presumed eating of this species by the early Tongans who deposited the bones at Tongoleleka. Remarks.— Although modern skeletons were not available for Megapodius lape- rouse of Micronesia (Palau, Marianas), measurements of skins (Baker 1951:106- 113) indicate that M. laperouse is smaller than M. freycinet freycinet, and therefore would be significantly smaller than M. ali- mentum. In spite of their large difference in size (Tables 1-3), the qualitative similarity between the tarsometatarsi of M. alimen- tum and M. pritchardi suggests that the for- mer may be more closely related to M. prit- chardi than to M. freycinet. 540 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON A B Cc Fic. 3. The digit I, phalanx 1 in dorsal aspect (A—C) and digit I-IV (exact number uncertain), terminal phalanx in lateral (D-F) and ventral (G-I) aspects in Megapodius. A, Paratype of M. alimentum, new species, Lifuka, Tonga, BPBM 165674; D, G, Paratype of M. alimentum, new species, Lifuka, Tonga, BPBM 165675; B, E, H, M. freycinet freycinet, male, Halmahera, Northern Moluccas, USNM 557015; C, F, I, M. pritchardi, sex unknown, Niuafoou, Tonga, USNM 319633. Scale bars = 10 mm. coideus, facies articularis clavicularis, and processus coracoideus, BPBM 165682, Pit ON11W, Layer IV (CU-IID, Tongoleleka archeological site (To-Li), Lifuka, Haapai Referred material. —Humeral half of cor- Group, Tonga. Tom Dye and field party acoid, lacking most of processus acrocora- Aug 1984. Megapodius cf. molistructor Balouet & Olson Fig. 4 Table 1.—Measurements (in mm) of the tibiotarsus in Megapodius, giving mean, range, and sample size. For sample sizes larger than 10, the mean is rounded to the nearest 0.05. F = female. M = male. U = sex unknown. Distal width Length Width of Distal width through Depth of through incisura Least width Least depth through epicondylus condylus pons supra- inter- of shaft of shaft condyles medialis lateralis tendineus _condylaris Megapodius alimentum 55) 4.8 ee 12.8 10.5 340) PAE, Holotype, BPBM 165686 l 1 1 1 1 1 1 Lifuka, Tonga (U) M. pritchardi 3.9 3.4 8.2 8.3 18 1.8 2.0 Niuafo‘ou, Tonga (2U) 3.84.0 3.3-3.5 8.1-8.3 8.1-8.5 7.1-7.5 1.6-2.0 1.9-2.0 a D 2 2 2 2 2 M. freycinet freycinet 5.60 4.35 P36 eT 9.40 2.65 PAS. Halmahera, Moluccas 4.8-6.4 3.7-4.7 10.3-12.2 11.1-12.5 8.8-10.1 2.0-3.2 1.5-2.6 (12M, 10F) 2 22 yA | 75, | 20 22 19 M. freycinet pusillus 5.9 4.6 123 12.6 9.9 2.6 26 Philippines (M) 1 l 1 1 1 1 1 M. freycinet gilberti 4.6 3.8 9.4 9.6 TH) 2D 1.6 Celebes (1M, 1F) 4.3-5.0 3.7-3.8 9.1-9.6 9.6—-9.7 8.1-8.3 DD, 1.4-1.7 2 ye 2 2 2 2 2 M. freycinet abbotti 5.6 4.6 11.4 Le, pe Draih 2S Nicobar Islands (2M) 5-3=5.9 4.5--4°7 "04 3-16 Visio 1 2.5-2.9 2.2-2.4 Z Z 2 2 jz 2 VOLUME 102, NUMBER 3 541 Table 2.— Measurements (in mm) of the tarsometatarsus in Megapodius, giving mean, range, and sample size. For sample sizes larger than 10, the mean is rounded to the nearest 0.05. The values for M. molistructor are estimations extrapolated from similar measurements in Table 3 of Balouet & Olson (in press b). FVP = foramina vascularia proximalia. F = female. M = male. U = sex unknown. Width of Minimum Depth of lateral shaft at width of Depth of shaft Proximal depth side of facies proximal edge shaft through just proximal Length of to hypotarsal dorsalis at level of fossa fossa to fossa fossa canal of FVP metatarsi I metatarsi I metatarsi I metatarsi I Megapodius alimentum 7.0 3.4 6.7 6.6 5 ae | 9.5 Lifuka, Tonga (U) l ] 1 1 1 1 BPBM 165670, 165689 M. pritchardi 4.4 1.9 4.2 4.3 2.4 5.9 Niuafo‘ou, Tonga (2U) 4.3-4.4 1.8—2.0 4.2-4.3 4.24.4 2.3-2.5 5.6-6.2 2 2 2 2 2 2 M. molistructor “cr al gad | _ ca. 8.7 ca. 8.9 ca. 4.4 = New Caledonia (U) l l 1 1 M. freycinet freycinet 5.80 2.45 6.10 6.25 3.25 8.85 Halmahera, Moluccas 4.8-6.3 1.7-2.9 5.5-6.8 5.6—7.0 2.9-3.6 7.9-9.6 (12M, 10F) 19 20 22 22 22 22 M. freycinet pusillus 6:2 2.6 6.4 6.7 33 9.6 Philippines (1M) 1 1 1 1 1 1 M. freycinet gilberti 4.8 2.4 5.0 52 any 7.9 Celebes (1M, 1F) 4.8 2.0—2.7 4.9-5.0 5.1-5.3 2.6—2.8 7.8-8.0 2 Z 2 2 2 2 M. freycinet abbotti 5.8 2.4 6.4 6.5 32 9.0 Nicobar Islands (2M) 5.8-5.9 1 6.2-6.5 6.2-6.8 3.1-3.4 8.7-9.2 2 2 2 Z 2 Table 3.— Measurements (in mm) of the digit I, phalanx 1 (DI, P1) and digit II-IV, terminal phalanx (DII- IV, TP) in Megapodius, giving mean, range, and sample size. For sample sizes larger than 10, the mean is rounded to the nearest 0.05. F = female. M = male. U = sex unknown. Minimum width of Minimum depth of Maximum width of DI, Pl DI, Pl Length of DI, P1 DII-IV, TP Megapodius alimentum od see 3.0 2.9 4.0+ Lifuka, Tonga (U) 1 1 1 1 BPBM 165674, 165675 M. pritchardi 16.3 2.0 1.6 pg | Niuafo‘ou, Tonga (2U) 16.0-16.6 1.9-2.0 1.6-1.7 2.6—2.8 2 2 2 2 M. freycinet freycinet 2225 235 255 3.20 Halmahera, Moluccas 20.8—23.9 2.2-2.8 2.3-2.8 2.7-3.5 (12M, 10F) 21 22 22 22 M. freycinet pusillus Fa 2.6 2.4 3.0 Philippines (1M) l | l ] M. freycinet gilberti 18.8 2.0 2.0 2.4 Celebes (1M, 1F) 18.3-19.3 1.9—2.1 1.9-2.0 | 2 2 2 M. freycinet abbotti 21S aes 2.4 a2 Nicobar Islands (2M) 21-5 5 2.3-2.4 3.0-3.5 Z 2 2 2 542 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 4. The coracoid of Megapodius in dorsal (A, B) and ventral (C, D) aspects. A, C, M. freycinet freycinet, male, Halmahera, Northern Moluccas, USNM 557015; B, D, M. molistructor, Lifuka, Tonga, BPBM 165682. Scale bar = 10 mm. Remarks. —This specimen is referred to Megapodius molistructor on the basis of its being much larger than any other species of Megapodius (Table 4), which is the only ge- nus of Megapodiidae that occurs in Ocea- nia. Intergeneric comparisons were not made. BPBM 165682 differs further from the coracoids of M. pritchardi, M. freycinet, and M. wallacei in having a less concave facies articularis humeralis. Although ad- ditional material is needed to determine with certainty whether the coracoid from Lifuka is conspecific with that of MM. molistructor, this specimen does demonstrate that an ex- tremely large species of megapode, approx- imately the size of M. molistructor, once lived on Lifuka. The type series of Megapodius molistruc- tor, an extinct species known otherwise only from New Caledonia (Balouet & Olson, 1989), does not include a coracoid. Never- theless, BPBM 165682 is much larger than in M. freycinet and larger than would be expected for the coracoid of M. alimentum. In coracoidal measurements (Table 4), /. cf. molistructor from Lifuka is from 1.30+ to 1.45 times larger (x = 1.35+, n = 5) than the means for M. freycinet freycinet. This corresponds with ratios of the measure- ments of the scapula and ulna from the type series of M. molistructor, which are from 1.25 to 1.40 times larger (X = 1.33, n = 7) than the means for M. freycinet frey- cinet (Balouet & Olson, 1989:Table 3). No leg elements of M. cf. molistructor are avail- able from Lifuka. The tarsometatarsus in M. molistructor from New Caledonia is much more massive than in M. alimentum VOLUME 102, NUMBER 3 543 Table 4.— Measurements (in mm) of the coracoid in Megapodius, giving mean, range, and sample size. For sample sizes larger than 10, the mean is rounded to the nearest 0.05. CS = cotyla scapularis. FAC = facies articularis clavicularis. FAH = facies articularis humeralis. ILA = impressio hgamentum acrocoracoideum. F = female. M = male. U = sex unknown. Depth between Length of Minimum width Depth of FAC ILA & FAH Width of FAH FAH & CS of shaft M. pritchardi 4.1 2.4 3.8 7.8 3.0 Niuafo ou, Tonga (2U) 4.04.2 2.2-2.5 3.64.0 7.6—7.9 2.9-3.1 2 2 2 2 2 M. cf. molistructor 7.9+ 4.2 6.3+ 13.6 5.7 BPBM 165682 1 1 1 1 Lifuka, Tonga (U) M. freycinet freycinet 5.95 2.90 4.85 10.20 4.20 Halmahera, Moluccas 5.4-6.4 2.6—3.4 4.6—5.2 9.5-11.2 3.64.6 (8M, 8F) 16 16 16 16 M. freycinet pusillus 6.4 2.9 4.8 10.3 4.5 Philippines (M) 1 I I M. freycinet gilberti BH 2.6 4.3 9.2 3.8 Celebes (1M, 1F) 5.6-6.2 2.4—2.7 1 9.1-9.2 3.8-3.9 2 2 2 M. freycinet abbotti 6.2 3.0 4.8 10.1 4.2 Nicobar Islands (2M) 6.0-6.3 2.8-3.1 1 l 4.14.4 2 2 M. wallacei 5.6 ae | 4.5 9.6 3 Halmahera, Moluccas (M) 1 1 1 1 (Fig. 2). Measurements of the tibiotarsus and tarsometatarsus of M. alimentum from L1- fuka are, respectively, from 0.98 to 1.21 times larger (X = 1.10, n = 7) and 1.05 to 1.39 times larger (X = 1.16, n = 6) than the means for M. freycinet freycinet, while mea- surements of the femur and tarsometatarsus of M. molistructor from New Caledonia are, respectively, from 1.32 to 1.54 times larger (x = 1.39, n = 3) and 1.14 to 1.43 times larger (x = 1.31, n = 4) than the mean values for M. freycinet freycinet. The validity of these calculations is not likely to be com- promised by a sexual dimorphism in size, which is extremely slight or non-existent in species of Megapodius (Mayr 1938, Ama- don 1942). Order Columbiformes Family Columbidae Genus Ducula Two specimens are referred to the genus Ducula rather than other genera of pigeons from Polynesia or eastern Melanesia (Co- lumba, Ptilinopus, Caloenas, Gallicolumba, Goura, Didunculus) because of these char- acters: coracoid—medio-ventral side of hu- meral end of shaft rounded, sulcus musculo supracoracoidei smooth and shallow, facies articularis sternalis medio-laterally expand- ed but dorso-ventrally compressed, impres- sio musculo sternocoracoidei deepest in medio-sternal corner; tibiotarsus—size and placement of prominent muscle scar on me- dio-distal surface of shaft, degree of con- cavity on the distal portion of shaft. Ducula, undescribed species Fig. 5 Material. —Tibiotarsus lacking both ends, BPBM 165685, Pit ONOW, Layer IV (CU- III), Tongoleleka archeological site (To-L1i), Lifuka, Ha‘apai Group, Tonga. Tom Dye and field party Aug 1984. Remarks. — This tibiotarsus is larger than in any other species of Ducula (Table 5). 544 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON A B C D Fig. 5: E F G H The tibiotarsus of Ducula in cranial (A—D) and caudal (E-H) aspects. A, E, Undescribed species, Lifuka, Tonga, BPBM 165685; B, F, D. galeata, Hanatekua Shelter No. 2 Archeological Site, Hiva Oa, Marquesas, BPBM 166055; C, G, D. aurorae, male, captive (original stock presumably from Makatea Island, Tuamotus), USNM 344776; D, H, D. pacifica, male, Rarotonga, Cook Islands, USNM 559586. Scale bar = 10 mm. The proximo-ventral and disto-lateral por- tions of the shaft have smoother, more rounded surfaces than other species. This tibiotarsus is too fragmentary to be named, yet it represents one of the largest of all columbids, being exceeded in size only by the crowned pigeons of New Guinea (Goura Spp.). Ducula cf. david Balouet & Olson Fig. 6 Referred material. —Nearly complete coracoid, lacking processus acrocoracoid- eus and part of facies articularis clavicularis, BPBM 165692, Pit ONOW, Layer IV (CU- III), Tongoleleka archeological site (To-L1), Lifuka, Haapai Group, Tonga. Tom Dye and field party Aug 1984. Remarks.—Ducula david, recently de- scribed from Wallis Island, was character- ized mainly by being larger than any extant congeners (Balouet & Olson, 1987). Among living species of Ducula, the largest of which occurs on oceanic islands, only D. galeata of eastern Polynesia and D. goliath of New Caledonia approach the size of D. david, although even these species are slightly smaller and less robust (Table 6). Ducula david is exceeded in size only by the huge extinct species noted above. Al- though comparable elements are not avail- able for the undescribed species and D. da- vid, comparisons of measurements of these VOLUME 102, NUMBER 3 Table 5.—Measurements (in mm) of the tibiotarsus in Ducula and Goura, giving mean, range, and sample size. F = female. M = male. U = unknown. Length from distal end of Length of Least width Least depth fibular crest to distal knob fibular crest of shaft of shaft LLL Ss Ducula, undescribed sp. 57.4 23.3 5.5 4.8 Lifuka, Tonga (U) l 1 1 1 D. galeata 39.7+ _ 3.9 323 Nuku Hiva, Marquesas (M) (est. 41-42) 1 1 | D. galeata 40.5 — 4.3 3.3 Henderson Island (U) (composite) 1 1 BPBM 160464, 160267 D. galeata 41.5 — 4.2 3.6 Hiva Oa, Marquesas (U) 1 1 1 BPBM 166055 D. goliath wife 12.7 4.3 3.8 New Caledonia (1M, 2F) 26.2—28.1 11.7-13.6 4.1-4.7 3.6—4.1 3 3 3 3 D. aurorae 26.6 10.9 3.9 3.0 Captive (M) l 1 | 1 D. pacifica 30.9 11.0 3.6 2.9 Niuafo ou, Rarotonga 30.2-31.8 10.9-11.2 3.3-3.7 2.8-3.0 (1M, 1F, 1U) 3 3 3 3) D. oceanica 30.0 ED 3.0 2.5 Palau, Ponape (2U) 28.5-31.4 11.0-11.4 3.0-3.1 2.42.6 D ) 2 D D. aenea 28.4 14.2 4.1 308) Philippines (F) ] 1 1 1 D. perspicillata 28.9 13.6 3.9 Sr2 Halmahera, Moluccas 28.2—29.6 13.2-14.1 3.9 3.0-3.4 (1M, 1F) D 2 2 2 D. bicolor 38.5 13.0 37 3.0 Halmahera, Moluccas (M) 1 1 1 1 D. luctuosa 28.2 13.9 3.5 Sul Celebes (F) 1 1 l 1 D. spilorrhoa D522 14.2 Sed 2.9 Australia (U) 1 1 1 l D. badia WB) 13.1 3.4 2.8 Thailand (M) l 1 1 1 D. pinon 29.4 14.0 4.0 Sul Captive (M) 1 1 l l D. radiata 19.4 10.7 3.1 2.4 Celebes (M) 1 1 1 1 Goura victoria 68.0 24.7 6.7 5.9 Captive (M) 1 1 species with those of D. galeata and D. go-_ ticular, the tibiotarsus of the undescribed liath indicate that the undescribed species species is 1.40 times longer than that of D. is larger than D. david (Tables 5 and 6herein; galeata, whereas the holotypical tarsometa- Balouet & Olson, 1987: Table 1). In par- tarsus of D. david from Wallis Island is only 546 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON VOLUME 102, NUMBER 3 1.01 times longer than that of D. galeata and 1.12 times longer than that of D. goli- ath. These ratios correspond well with those of the coracoid in D. cf. david from Lifuka, which is 1.07 times longer than that in D. galeata and 1.12 times longer than that in D. goliath. The amount of sexual dimor- phism in size is poorly understood in ocean- ic species of Ducula owing to the worldwide scarcity of skeletons. In measurements of D. goliath, the male is consistently larger than the female (Tables 5 and 6), although these are captive individuals that may not be reliable indicators of the size of wild birds. No skeletons were available for D. /atrans of Fiji. Based upon measurements of skins (duPont 1976:83-85), D. latrans is approx- imately the same size as D. aurorae or D. pacifica, and thus would be much smaller than D. david. Ducula pacifica (Gmelin) Referred material.—Ulna lacking both ends (BPBM 165676), Pit 126NOW, Layer III (CU-IIb), Tongoleleka archeological site (To-Li), Lifuka, Haapai Group, Tonga. Tom Dye and field party Aug 1984. Remarks. — The curvature of the shaft and the prominent papillae remigiales caudales refer this ulna to the Columbidae. The spec- imen agrees in size and other features with the ulna of Ducula pacifica. Each of the oth- er three species of columbids reported here is much larger than D. pacifica, which is the only species of columbid (other than Gal- licolumba stairii and species of Ptilinopus, which are very small) surviving on Lifuka or anywhere else in Tonga. Genus Caloenas Among the bird bones from the Tongo- leleka Site is another coracoid of a large — 547 columbid. This specimen, slightly smaller than in D. cf. david (Table 6), is referred to the genus Caloenas rather than to Ducula or other pertinent genera of columbids be- cause of these characters: greater pneuma- ticity in humeral end of sulcus musculo su- pracoracoidei; facies articularis humeralis protrudes more ventrad from surface of shaft; sharp medio-ventral edge of humeral end of shaft; in medial aspect, portion of shaft between cotyla scapularis and facies articularis clavicularis faces more perpen- dicularly (less diagonally); facies articularis clavicularis deeper. Caloenas cf. canacorum Balouet & Olson Fige o/ Referred material. —Humeral end of cor- acoid, including facies articularis humeralis and cotyla scapularis, BPBM 165678, Pit ON20E, Layer II (CU-II), Tongoleleka ar- cheological site (To-Li), Lifuka, Haapai Group, Tonga. Tom Dye and field party Aug 1984. Remarks.—Caloenas canacorum is an extinct species recently described from late Holocene fossils (sternum, coracoids, scap- ula, and humerus) from New Caledonia (Balouet & Olson, 1989). Although direct comparison of the holotype coracoid of C. canacorum with BPBM 165678 was not possible except in photographs (Fig. 7), BPBM 165678 is referred to C. cf. cana- corum because of similarity in qualitative generic characters and in size, being signif- icantly larger than in C. nicobarica, the only living species in this distinctive genus (Ta- ble 6). Discussion Although the detailed implications of these findings will be reported elsewhere, a Fig. 6. The coracoid of Ducula in dorsal (A—D) and ventral (E—-H) aspects. A, E, D. cf. david, Lifuka, Tonga, BPBM 165692; B, F, D. galeata, Hanatekua Shelter No. 2 archeological site, Hiva Oa, Marquesas, BPBM 166056; C, G, D. aurorae, male, captive (original stock presumably from Makatea Island, Tuamotus), USNM 344776; D, H, D. pacifica, male, Rarotonga, Cook Islands, USNM 559586. Scale bars = 10 mm. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 548 suivjndeos RIA1O9 Soro JO YIPIM solory Jo yIBUTT] PUd [BULIO}S 0} 199RJ Jeulojs WO YIdUIT (6 6 Z Z Z iG Z Z Z (41 ‘WI) 9°8I-p'8I l't-6°€ TPB’ O'S-L'b Ct-b'r [8-Gel ¢°S-7'S CSE-b' PE 7 Le-6'SP seoonjoyy ‘eloyeuyey CsI Or O'r SP vy OL v's OSE 9°9P vyoppioidsiad °q I I I I I I I I I (4) sourddiy1yg ELI SE O'r xs O'r 9°L LY I've T Sb pauav ‘q I I I I I I (I) eAndeD aH Gay Or — = 0°8 aS 6 VE SLY DAOAIDNSIAAU “Gl G € € Z € € € € € (N¢) SI-V' El 87-17 0'€-9°7 L€-9'€ €°€-0'€ 0'S-S'P O'r-S'€ GA CAN ASIC C 9€-L'ZE odeuog ‘nejed 9°€EI 9°7 87 9'€ I'€ LY SE O'LZ 6'VE po1upa20 ‘qd 74 7 7 7 7 7 7 17 7 (AT “AT SWZ) esuojo1ey 9°SI-r' El l'€-'7Z O'€-S°7Z Ev-S€ 6 €-T'E 6'S-€°S l'p-6°€ 7 7E-b 87 O'Ip-€°9€ ‘NO, OJeNIN] “S}1IqQ]IH (anal 87 87 6'€ 9°€ cs O'r v'6C SLE poyfiaod ‘q I I I I I I I I (WW) 9AndeD EST L€ 9'€ os O'r 79 o'r IIe J It ADAOAND ‘GT € € € G G G € € € (AZ ‘WI) 1'0@-r'LI Sr-l'b L'v-0'P 9°S-0'S p'S-€'P t'L-6'°9 ¢°9-S'°S pb LeE-l've 9°6¢-6' br BIUOps[eD MON LSI vp (On ES SP Gill 19 BSE OLY yj01038 ‘q iG G G (Nn) 950991 Ndda I I I TS-6'P 9'b-0'r O'8-I'L I I I sesonbieyy “8O BATH O81 SE 6'€ (xs ev OL 9°¢ 7’ 8E 9°6b DIvavs ‘q I (A) purysy ste — — Tp eo — — _ = = = Piavp ‘d I I (1Z 389) I I I I I (€¢ “1S9) (Q) esuol “exnjrT +6°SI ES 6'P 69 +¢°¢ ~ = SIP +0°8p plapp “j9 ‘q PUd [BUIOIS JO YIPIAA yeys jo yeys Jo snoplooe1090108 STjeIoWINY SI[eIoUINY puo [elowny Ssije1oumny yidus] WINWIxes| YIpIM sea] yidop iseoT snssoo01d pue Sup[Norpie supine jo yidoq SLUIB[NOILIe So1OR] JO usemjeq 1eYs jo uidop iseoT ‘UMOUYUN XOS = () ‘JPW = PA ‘O[BUIDJ = J (6861) UOSTO % JONO[eg WoO ‘suOT}eWIT}Sso poejodenxs WIOIF IO ATJO9IIP JOYU9 ‘UdZeR] de PIUOPI[VD MON WIOIJ WNIOIDUDI ‘Dd IO} SONTeA YJ, (L861) UOSIO FW JONoOTeg JO | IQUI, Ul SJUSWIOINSeOU ILIWIIS WOT, pajye[oOde1x9 SUOTIeUITISO 218 PURIS] SIT[VAA WOI PiAvp ‘Gq JO} sonjeA dU ‘9ZIS o[dures pue ‘osues ‘URDU BUTAIZ ‘SPUIOJVD puke DINING Ul PlodesOS 9Y} JO (UI UT) SJUDUOINSBI|I — "9 BIQVL 549 VOLUME 102, NUMBER 3 8L9S91 Wddd (Q) esuoy ‘eynyry anes — — 19 oS 6'8 +¢°9 —. — WUNnNAOIDUDI ‘po =) Z Z eserae yh p'6S-0'°6S (NZ) Bluopsyed MON sont - -_ ~ = —e 78 _ ORAS WUNAOIDUDI *y ¢ ¢ ¢ ¢ ¢ ¢ ¢ ¢ ¢ (4S) 8°91-6'rl O'r-7'€ S'E-E'€ ¢'S-9'p 0'S-b'P 6'L-8'9 C'S-8'b 8'PE-£°7E C'9b-S Eb svoonjoyy “eioyeUyeH gol 9°€ 9°€ ae Lv pL I'S L'€€ 8'PP DILADGOINU SDUIO]D) I I I I I I I I I (WN) S9q9[9D pr 67 67 9°€ O'€ 09 ge €°67 6'8E DIDIPDA ‘] I I I I I I I (I) 9ANdeD = 6'€ O'P = i OL p's L'9€ 9°8p uould ‘q I I I I I I I () PuepieyL = 8'€ ce = 6'€ €°9 CP 9°7E 6 TP pipvg ‘d I I I I I I I (AQ) Byensny = ge L€ ne 0'P OL 8'P COE L'0v poysdopids ‘ I I I I I I I (4) s9qojaD = 8°€ [his = lv VL O'S OEE OED psonjon] “q I I I I I I I I (IA) seoonjoyy ‘eroyeuyey = ce 9°€ 9" O'P (ofl cP OIE SIP 40]0I19 ‘d pus [BUIO}S JO YIPIA yeys jo yeys Jo Ssnoploor10900108 SI[eJOWINY Sije1ouNny puo [e19wny Ssijesouiny yisuo] WNWIxeyyy YIPIM seo] yidop iseo] snssooo1d pue SLIBNOIE sue[norpie jo yidoq SLIB[NIAe So1oRy JO siivjndeos RIAIOO us9mI10q WeYs jo yidop isvoT SO1ORJ JO YIPIAA sSoloRy JO yIdUST PUD [PUIO}S 0} 190RF [eusojs WO YIsUIT ‘panunuoy—"9 91921 550 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 7. The coracoid of Caloenas in dorsal (A-C) and ventral (D-F) aspects. A, D, C. nicobarica, female, Halmahera, Northern Moluccas, USNM 557089; B, E, Caloenas cf. canacorum, Lifuka, Tonga, BPBM 165678; C, F, C. canacorum, paratype, New Caledonia, MNHN 300. Scale bar = 10 mm. few comments are pertinent here. The un- described species of Ducula is of unknown interspecific relationships. The other extinct birds from Lifuka are related to species found west or north of Tonga. It seems like- ly that all of these species, or closely related ones, once occurred through much of the region of New Caledonia, Vanuatu, Fiji, Tonga, and Samoa. Megapodius molistruc- tor and Caloenas canacorum are known only from New Caledonia and tentatively from Lifuka. Megapodius alimentum and the un- described species of Ducula are known only from Lifuka, although there is no reason to believe that they were confined to that is- land. Aside from Lifuka, D. david is known from Wallis (Uvea) Island, which is north of Lifuka, between Samoa and Fiji. The late Holocene extinction of two species of megapodes and three species of pigeons on Lifuka shows that there has been a significant loss of birds in western Poly- nesia since the arrival of man. Numerous extinctions have been documented by fossil records from more remote parts of Poly- nesia, such as Hawaii (Olson & James 1982a, b), Marquesas (Steadman, in press), Hen- derson Island (Steadman & Olson 1985), Cook Islands (Steadman 1985, in press), and New Zealand (Cassels 1984). More exca- vation is needed in Tonga and Samoa. Based upon the limited record available, the de- gree of avian extinction in western Poly- nesia may have been just as severe as that from elsewhere in Polynesia. The fossils from Lifuka indicate that two species of Megapodius once occurred there. Megapodius pritchardi, restricted to the iso- lated Tongan island of Niuafo ou, is the only species of megapode that survives anywhere in Polynesia, although we presently do not know to what extent the natural range of VOLUME 102, NUMBER 3 megapodes has been reduced by human im- pact. The widespread M. freycinet reaches the eastern limit of its range in Vanuatu (New Hebrides). That the absence of mega- podes in the Fijian region may be an artifact of human disturbance was noted by Olson (1980) and confirmed several years ago by our examination of bones of Megapodius (species undetermined) from an archeolog- ical site on Lakeba, Lau Group, Fiji (re- ported in Gibbons & Clunie 1986). Else- where in the western Polynesian and Me- lanesian region, extinct megapodes (species undetermined) have been reported from as yet unconfirmed historical accounts in the Kermadec Islands (Lister 1911), archeolog- ical sites on Tikopia (Kirch & Yen 1982: 282; Green 1976), an egg collected in 1847 from Samoa (island undetermined; Gray 1862), and an egg collected before 1862 from an undetermined island in the Haapai Group of Tonga (Gray 1862, 1864). Oates (1901) referred the last two specimens to M. pritchardi, a determination that should be reconfirmed. A better understanding of the systematics and natural distribution of megapodes in Oceania depends upon the reexamination of historic specimens and documents, and more fully upon the discovery and study of bones from prehistoric sites on many ad- ditional islands. It now seems likely that one to three species of megapode occurred on most or all islands of eastern Melanesia and western Polynesia before the arrival of hu- mans. Four species of megapodes still exist, for example, on the Papuan island of Misool (Ripley 1960). Columbids also have suffered much ex- tinction in Oceania. The hunting of pigeons by prehistoric Tongans was extensive and highly organized (McKern 1929:19-27). Ducula pacifica is the largest pigeon known historically from anywhere in Tonga, where it is found essentially throughout the group. Fossils from Tongoleleka represent three additional species of columbids, each ex- tinct and larger than D. pacifica. Although 551 it may seem remarkable that four large species of columbids, including three species of Ducula, once lived on Lifuka, we really do not yet know the natural (=pre-human) distribution and diversity of Pacific co- lumbids. From Mangaia in the Cook Is- lands, for example, late Holocene fossils represent five species of columbids where none exists today (Steadman 1985, 1989). Acknowledgments I thank T. Dye for his cooperation and generosity in making available the Lifukan specimens and associated information. S. L. Olson and J. C. Balouet kindly allowed ac- cess to specimens and data on extinct birds from New Caledonia, including photo- graphs of Megapodius molistructor and Ca- loenas canacorum. For access to specimens, I thank J. P. Angle, S. L. Olson, C. A. Ross, and R. L. Zusi (USNM) and A. Allison, C. H. Kishinami, and G. Wine (BPBM). D. Pahlavan, S. E. Schubel, and M. C. Zarriello assisted in other curatorial matters. D. Shi- deler, L. Woodward, and A. Ziegler sorted and preliminarily identified the bones. The photographs are by T. Beblowski and C. Supkis. The manuscript was improved by comments from T. Dye, N. G. Miller, S. L. Olson, and G. K. Pregill. The research was supported by National Science Foundation Grant BSR-8607535. This paper is contri- bution number 538 of the New York State Science Service. Literature Cited Amadon, D. 1942. Birds collected during the Whit- ney South Sea Expedition. XLIX. Notes on some non-passerine genera, 1.—American Museum Novitates 1175:1-11. Baker, R. H. 1951. The avifauna of Micronesia, its origin, evolution, and distribution. — University of Kansas Publications, Museum of Natural History 3:1-359. Balouet, J. C., & S. L. Olson. 1987. A new extinct species of giant pigeon (Columbidae: Ducula) from archeological deposits on Wallis (Uvea) 352 Island, South Pacific.— Proceedings of the Bio- logical Society of Washington 100:769-775. ,& 1989. Fossil birds from late Qua- ternary deposits in New Caledonia. —Smithson- ian Contributions to Zoology 469:1-38. Baumel, J. J., A. S. King, A. M. Lukas, J. E. Breazile, & H. E. Evans (eds.). 1979. Nomina anato- mica avium. Academic Press, London, 664 pp. Cassels, R. 1984. The role of prehistoric man in the faunal extinction of New Zealand and other Pa- cific islands. Pp. 741-767 in P. S. Martin and R. G. Klein, eds., Quaternary extinctions. Uni- versity of Arizona Press, Tucson. duPont, J. E. 1976. South Pacific birds.— Delaware Museum of Natural History Monograph Series 3:1-218. Gibbons, J. R. H., & F. G. A. U. Clunie. 1986. Sea level changes and Pacific prehistory.—Journal of Pacific History 21:58-82. Gray,G.R. 1862. List of species composing the Fam- ily Megapodiidae, with descriptions of new species, and some account of the habits of the species. — Proceedings of the Zoological Society of London for 1861:288-296. 1864. Ona new species of megapode.— Pro- ceedings of the Zoological Society of London for 1864:41-44. Green, R. C. 1976. Lapita sites in the Santa Cruz Group. Pp. 245-265 in R. C. Green and M. M. Cresswell, eds., Southeast Solomon Islands cul- tural history.—Royal Society of New Zealand Bulletin 11. Karch, P. V., & D.E. Yen... 1982. Tikopia= The: pre- history and ecology of a Polynesian outlier. — Bernice P. Bishop Museum Bulletin 238:1-396. Lister, J.J. 1911. The distribution of the avian genus Megapodius in the Pacific Islands.— Proceed- ings of the Zoological Society of London for 1911:749-759. Mayr, E. 1938. Birds collected during the Whitney PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON South Sea Expedition. XX XIX. Notes on New Guinea birds.—American Museum Novitates 1006:1-16. McKern, W. C. 1929. Archaeology of Tonga.—Ber- nice P. Bishop Museum Bulletin 60:1-123. Oates, E. W. 1901. Catalogue of the collection of birds’ eggs in the British Museum (Natural His- tory). British Museum (Natural History), Lon- don, 252 pp. Olson, S.L. 1980. The significance of the distribution of the Megapodiidae.—Emu 80:21-24. —, & H. F. James. 1982a. Fossil birds from the Hawaiian Islands: Evidence for wholesale ex- tinction by man before Western contact.—Sci- ence 217:633-635. ——., & 1982b. Prodromus of the fossil avifauna of the Hawaiian Islands.—Smithson- ian Contributions to Zoology 365:1-59. Ripley, S. D. 1960. Distribution and niche differ- entiation in species of megapodes in the Mo- luccas and western Papuan area.— Proceedings XII International Ornithological Congress 2: 631-640. Steadman, D. W. 1985. Fossil birds from Mangaia, southern Cook Islands.— Bulletin of the British Ornithologists’ Club 105:58-66. 1989. Fossil birds and biogeography in Po- lynesia.—Acta XIX Congressus Internationalis Ornithologici II:1526-1534. —, & S. L. Olson. 1985. Bird remains from an archaeological site on Henderson Island, South Pacific: Man-caused extinctions on an “unin- habited”’ island.— Proceedings of the National Academy of Sciences USA 81:4448-4451. Biological Survey, New York State Mu- seum, The State Education Department, Al- bany, New York 12230. PROC. BIOL. SOC. WASH. 102(3), 1989, pp. 553-554 GEOGRAPHIC VARIATION IN THE YELLOW-RUMPED TANAGER (AVES: THRAUPINAE) Robert W. Storer Abstract.—Females and young males of the Yellow-rumped Tanager (the icteronotus group of the Flame-rumped Tanager, Ramphocelus flammigerus) from Panama to west central Colombia are grayer and less green on the back with less black on the crown than are those from Ecuador. Birds from south- western Colombia and northeastern Ecuador are intermediate between those populations, the latter being more similar to those of the other Ecuadorean populations. The name Ramphocelus flammigerus varians Lafresnaye should be applied to the northern populations and R. f. icteronotus Bonaparte to those from Ecuador. Yellow-rumped Tanagers (the icteronotus group of the Flame-rumped Tanager, Ram- Dhocelus flammigerus) are widely distrib- uted from western Panama to southwestern Ecuador (American Ornithologists’ Union 1983). Wetmore et al. (1984:449) comment that “preliminary examination reveals that females and immatures from Ecuador are much greener, less brownish or gray, above than are most birds from Colombia and Panama. Thus it is possible that the Pana- manian birds should take the name varians Lafresnaye 1847 (type locality, Buenaven- tura Colombia).” To determine the correct name for the Panamanian birds, it was necessary to de- termine first, if there is sufficient geographic variation to recognize more than one race, and second, if so, whether the birds from near the type locality of varians are closer to birds from Panama or western Ecuador (the restricted type locality of icteronotus, Berlepsch, 1912). Should the latter situation apply, a new name would be needed for the Panamanian birds. I examined 143 specimens of females and immature males of the Yellow-rumped Tanager for analysis. (Adult males do not vary in the deep black of the back.) Wing length (chord), tail length, and length of bill from nostril to tip of females were measured with dial calipers, the first two to the nearest 0.5 mm and the last to the nearest 0.1 mm. (The sample of males was too small to treat Statistically.) The geographic range was di- vided into five areas with gaps between them to avoid comparisons between contiguous populations and to reduce overlap between samples. The selection of the areas was based primarily on the distribution of the mate- rial. These areas were: Western Panama (Bocas del Toro Province and the Canal Zone), Eastern Panama (Darien Province), Central Colombia (provinces of Cauca, To- lima, and Valle), Northern Ecuador (prov- inces of Esmeraldas, Imbabura, and Pichin- cha), and Southern Ecuador (provinces of Canar, El Oro, Guayas, and Loja). Color comparisons were made by ex- amination of specimens in north light. Bad- ly worn or faded specimens, approximately one half of those examined, were not used in the comparisons. The measurement data for females (Table 1) show a cline in increasing wing and tail length from western Panama to southern Ecuador. Bill length is greatest in birds from eastern Panama and least in those from western Panama and central Colombia, but mean differences are small. In none of the 554 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 1.—Sample size, mean, and standard deviation of measurements of female Yellow-rumped Tanagers. Measurement Western Panama Eastern Panama Central Colombia Northern Ecuador Southern Ecuador Wing length fT WW 0) 9 S29" aale9 23) OS te 1.8 10 80.4 + 2.6 12) 83:4 27189 Tail length 6 68.2, 4233.2 $68.9: 3.2 21 GOR say 2yi 1Q; 69.7 22.23 12. 138.4537) Bill length A AZ EOS 9) 13:4,-©.0:6 22. 12:8 2106 10, 32.203 12, 13,125)0e measurements is the difference between the largest and smallest mean greater than the sum of the standard deviations. Therefore, none of the differences approaches a degree of difference useful in separating subspecies. In wing and tail measurements, the birds from central Colombia are closer to those from eastern Panama than to those from northern Ecuador. The data for bill length are equivocal. In color, specimens from Panama are de- cidedly grayer (less black) on the back than those from Ecuador, the crowns are less black, the yellow edging on the back feathers is paler and contrasts less with the dark col- or of the rest of the feather, and the yellow of the underparts is paler, especially on the abdomen. The brighter yellow edgings and darker central parts of the feathers combine to produce a greener tone to the back feath- ers of the Ecuadorean birds. Five specimens from the province of Valle, central Colom- bia, (including Buenaventura, the type lo- cality of varians) are nearer birds from Pan- ama in color than those from Ecuador. In Ecuador, there is a noticeable difference in color between specimens from the northern and those from the central and southern parts of the country, the latter being bright- er. However, birds from northern Ecuador are more similar to those from the south than to those from Panama and northern Colombia. Thus, birds from Panama through central Colombia (Valle) vary little in color, the major change occurring from Cauca to Ecuador, with a smaller change occurring between the northern and central parts of Ecuador. In conclusion, differences in wing, tail, and bill lengths are not sufficient to warrant recognition of subspecies. Color differences, however, are greater than in many recog- nized subspecies. I therefore propose that the birds from Panama through the prov- ince of Valle, Colombia, be called Ram- Dhocelus flammigerus varians Lafresnaye, those from Ecuador, Ramphocelus flam- migerus icteronotus Bonaparte, and those from southwestern Colombia, intergrades. Acknowledgments I acknowledge the assistance of the cu- rators of the bird collections of the Amer- ican Museum of Natural History, the Field Museum of Natural History, and the Na- tional Museum of Natural History, Smith- sonian Institution, for the loan of speci- mens, and Richard C. Banks and Thomas R. Howell for reading the manuscript and offering helpful suggestions. Literature Cited American Ornithologists’ Union. 1983. Check-list of North American Birds, 6th Edition, 662 pp. Berlepsch, H. von. 1912. Bericht ueber den V. In- ternationalen Ornithologen-Kongress Berlin 1910, p. 1061. Wetmore, A., R. F. Pasquier, & S. L. Olson. 1984. The Birds of the Republic of Panama.—Smith- sonian Miscellaneous Collections 150(4):448- 449. Museum of Zoology and Department of Biology, University of Michigan, Ann Ar- bor, Michigan 48109. PROC. BIOL. SOC. WASH. 102(3), 1989, pp. 555-558 TWO OVERLOOKED HOLOTYPES OF THE HAWAIIAN FLYCATCHER CHASTEMPIS DESCRIBED BY LEONHARD STEJNEGER (AVES: MYIAGRINAE) Storrs L. Olson Abstract.—The holotypes of two of the three taxa of Elepaio (Chasiempis) described by Leonhard Stejneger in 1887 have been overlooked for almost a century but were located in the collections of the British Museum (Natural History) and restudied. That of Chasiempis ridgwayi Stejneger presents no problems as it is representative of the subspecies from the windward parts of the island of Hawaii currently known as Chasiempis sandwichensis ridgwayi. The holotype of Chasiempis ibidis Stejneger, 1887, however, is identified as being from the Oahu population and this name thus takes precedence over Chasiempis gayi Wilson, 1891. In the early literature on systematics and nomenclature of the Hawaiian flycatcher known as the Elepaio, Chasiempis sand- wichensis (Gmelin), there was considerable disagreement concerning the number of taxa and their distribution. Some proponents held out for a single species (e.g. Sclater 1885, Berlepsch & Leverkuhn 1890), while others recognized as many as five or six (e.g. Steyneger 1887, Wilson 1891), with the is- land of origin often being completely ig- nored, however. It was eventually deter- mined that the Elepaio occurred only on three of the Hawaiian islands, with the pop- ulation of each island coming to be recog- nized under a single name: C. sclateri Ridg- way, 1882, on Kauai; C. gayi Wilson, 1891, on Oahu; and C. sandwichensis (Gmelin, 1789) on Hawaii. These taxa are now gen- erally regarded as subspecies of C. sand- wichensis. Geographic variation within the island of Hawaii caused Henshaw (1902) to recognize two forms there, with the second taking the name C. ridgwayi Stejneger, 1887. Pratt (1979, 1980) recognized these and de- scribed a third subspecies from Hawaii, C. s. bryani. Leonhard Stejneger was in the thick of the early confusion surrounding the system- atics of Chasiempis and proposed no less than three new taxa (Stejneger 1887). The only actual specimens available to him, however, were those taken on Kauai by Val- demar Knudsen and forwarded to the Smithsonian Institution. Part of the prolif- eration of taxa resulted from the two dis- tinct plumage types found on each island, now generally regarded as adult and “im- mature.”’ Thus, Stejneger’s name Chasiem- pis dolei, the type of which (USNM 110040) is a gray-backed adult bird from Kauai, is a pure synonym of C. sclateri Ridgway, 1882, the cotypes of which (USNM 41955, 41956) are brown-backed immature birds from Kauai (see Deignan 1961:460). Stejneger’s other two names, C. ridgwayi and C. ibidis, were based on a color plate published in Jbis (hence the latter name) by Sclater (1885). Although Stejneger attempt- ed to forestall criticism by saying that if C. ibidis were not distinct from C. sclateri “‘then I can only say that the published figure of the former is worse than useless” (Stejyneger 1887:88), his contemporaries nevertheless roundly excoriated him. “It is a pity that Dr. Stejneger, with so much good material before him, should think it necessary to manufacture ‘new species’ out of other peo- 556 ple’s figures without seeing the specimens” (Sclater 1888:143). “On the genus Chasiem- pis | would offer only one remark, and that is a word of caution to those who would, on the evidence of from a couple to half-a-doz- en of specimens, or perhaps even on the evidence of a badly-coloured plate, attempt to break it up into definable ‘species’ ”’ (Newton 1892:469). With the benefit of hindsight, it is fair to note that the plate in question is actually quite accurate, and that both of Stejneger’s taxa based on it are now seen to be valid. Although both figures were stated to be based on particular specimens, these have been overlooked in the general collections of the British Museum (Natural History) for nearly a century and have long gone unrec- ognized as types (neither is mentioned in Warren & Harrison 1971). They still exist, however, and I was able to examine and compare them with the series of Chasiempis in the National Museum of Natural History, Smithsonian Institution (USNM), and with a selection of specimens from the American Museum of Natural History that were in plumage comparable to that of C. ibidis (see Material Examined). One of Stejneger’s names was applied to ‘the brown and chestnut colored bird from Hawaii, Ch. ridgwayi, figured on plate 1, Ibis, 1885” (Stejneger 1887:87). Sclater (1885: 18) had mentioned two specimens collected by the Challenger Expedition at Hilo, Ha- wail, in August, 1875, and stated that “‘the figure (Plate I fig. 1) has been taken from one of them.” Both specimens are in nearly identical plumage but one of them (BMNH 80.11.18.445, original no. 529) is in much worse condition, with most of the feathers of the rump missing and many of the rec- trices broken off, so that only one remains that has a white tip. Because the plate shows a bird with a large white rump patch and white tips on most of the rectrices, if it were drawn from a single specimen, as stated by Sclater, this would have to be BMNH PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 80.11.18.444 (original no. 528), which I here affirm to be the holotype of Chasiempis ridgwayi Stejneger (culmen, 13.1 mm; wing, 67.7; tail 54.5, tarsi not measurable). No nomenclatural problems attach to this iden- tification, as the specimen is of known provenance and is clearly the bird from windward Hawaii currently known as Cha- siempis sandwichensis ridgwayi. The specimen from which Sclater’s re- maining figure was drawn, the type of Cha- siempis ibidis, has a much more curious and enigmatic history. Sclater (1885:18) re- ceived the specimen, labelled “‘Chili,” prior to 1862 from the dealer Verreaux and mis- took it for some undetermined species of Tyrannidae. Thus he once listed it as “Cni- polegus __?” (Sclater 1862:203), an error that he later corrected to Chasiempis sand- wichensis (Sclater 1873) after comparing the specimen with material in the Berlin Mu- seum (Sclater 1885), where the only speci- mens of Chasiempis then were those col- lected on Oahu by Deppe in 1837. The same specimen was listed by Sharpe (1879) as Chasiempis sandvicensis (sic), at which time it was the only example of Elepaio in the British Museum. It is the only Elepaio from the Sclater collection ex Verreaux in the British Museum collections or elsewhere and is thus certainly identifiable as the holotype of Chasiempis ibidis Stejneger (BMNH 73.8.6.3; culmen broken; wing, 62.9 mm; tail, 59.3; tarsus, 25.1). Itisin fresh, unworn plumage and agrees perfectly with the figure in Sclater’s (1885) plate except that the ochraceous color at the posterior margin of the throat is somewhat darker than depict- ed. Circumstantial evidence alone would suggest that this specimen came from Oahu, as most commerce then, as now, was through Honolulu. That it had been labelled “Chili” is understandable as Chile was then a dis- patch point for cargo going “around the Horn” to Europe. In the Senckenberg Mu- seum, Frankfurt, are at least five specimens VOLUME 102, NUMBER 3 of Hawaiian birds labelled as received or exchanged from Chile in 1842, the notation “Erh[halten] von Chili’? having been mis- taken by Banko (1979:31, 32, 80) for the collector’s name. Of these, three are of the Oahu form of Loxops virens; the other two belong to species that show no interisland variation (Psittirostra psittacea and Ves- tiaria coccinea). Itis uncertain who was sup- plying specimens from Oahu through Chile at that time, but it was very likely the same source whence Verreaux obtained the spec- imen of Chasiempis sold to Sclater. As far as the actual characters of the ho- lotype of Chasiempis ibidis are concerned, Stejneger (1887:88) was perfectly correct in describing it as differing from C. sclateri of Kauai, which is “much deeper and richer tawny color. . . and this color extends much further on breast, flanks, and tibiae than in Ch. ibidis.” Wilson (1891) considered C. ibidis to have come from Oahu. The new form from Oahu that he called Chasiempis gayi he described as a second species from that island in ad- dition to C. ibidis. Why then has the Oahu bird come to be known as C. gayi rather than C. ibidis? This results entirely from Rothschild’s (1893:71) statement that the type of C. ibidis ‘agrees best with the young Hawaiian bird, so there is no doubt it really came from Hawaii.” This is erroneous. Rothschild’s conclusion appears to have been unduly influenced by some exceptional specimens (e.g. AMNH 607136, 607138) collected by Palmer on the Kona coast of Hawaii, the first mentioned having served as the model for the immature of C. sand- wichensis in Rothschild’s accompanying plate. These birds are much more rufescent, especially on the throat and breast, than typ- ical immature birds from Hawaii, which are dark brownish above, with a grayish crown, and white lores and underparts. The light tawny ochraceous color of the type of C. ibidis is very unlike this and is matched only by specimens from Oahu. The exceptionally 557 rufescent specimens from Hawaii are still much darker, more chestnut, above, espe- cially on the rump, than in C. ibidis. Another overlooked difference is in the shape of the bill, which in birds from Oahu and Kauai appears broader and flatter than in birds from Hawaii. Although the bill in the type of C. ibidis is damaged and lacks the tip of the upper part, its shape agrees better with birds from Oahu than with those of Hawaii. The holotype of Chasiempis ibidis is un- questionably representative of the Oahu population of Elepaio, as Wilson (1891) himself recognized. Because Chasiempis ibidis Stejneger, 1887, has priority over Chasiempis gayi Wilson, 1891, the Oahu Elepaio should now be known as Chasiem- pis ibidis or Chasiempis sandwichensis ibi- dis. Material examined.—Chasiempis s. sandwichensis:. AMNH 607118, AMNH 607125, AMNH 607136, AMNH 607138. C. s. ridgwayi: BMNH 80.11.18.444 (ho- lotype), BMNH 80.11.18.445, AMNH 193362, AMNH 193366, AMNH 193368, plus about 30 USNM specimens in imma- ture plumage. C. ibidis: BMNH 73.8.6.3 (holotype), AMNH 193354, AMNH 193355, AMNH 193357, AMNH 168638, AMNH 199353, AMNH 607160, USNM 301122. C. sclateri: AMNH 168639, AMNH 193347, AMNH 607188, AMNH 607189, AMNH 607190, AMNH 607198, USNM 41955 and 41956 (cotypes), USNM 110040 (type of C. dolei), USNM 110037, USNM 110038, USNM 116782, USNM 116783, USNM 493863, USNM 493864, USNM 591935, USNM 591936. Acknowledgments I am exceedingly grateful to Graham Cowles, Sub-Department of Ornithology, British Museum (Natural History) (BMNH), Tring, for lending the specimens that proved to be the types of C. ridgwayi and C. ibidis, and to Richard A. Sloss and Mary LeCroy, American Museum of Natural History 558 (AMNH), New York, for supplying addi- tional comparative material. I also thank D. S. Peters for access to specimens of Hawai- ian birds in the Senckenberg Museum, Frankfurt. Literature Cited Banko, W. E. 1979. History of endemic Hawaiian birds [sic] specimens in museum collections. — Cooperative National Park Resources Study Unit, University of Hawaii, Avian History Re- port 2:1-80. Berlepsch, H. von, & P. Leverkiihn. 1890. Studien iiber stidamerikanische Vogel nebst Beschrei- bungen neuer Arte.—Ornis 6:1-32. Deignan, H. G. 1961. Type specimens of birds in the United States National Museum.— United States National Museum Bulletin 221:1-718. Gmelin, J.F. 1789. Systema Naturae. Volume 1, part 2. Lipsiae, G. E. Beer, pp. 501-1032. Henshaw, H. W. 1902. The Elepaio of Hawaii.— Auk 19:225-232. Newton, A. 1892. Omnithology of the Sandwich Is- lands. — Nature 45:465-469. Pratt, H. D. 1979. A new subspecies of the Elepaio, Chasiempis sandwichensis, from the island of Hawatii.— Bulletin of the British Ornithologists’ Club 99:105-108. 1980. Intra-island variation in the ‘Elepaio on the Island of Hawai’i.— Condor 82:449-458. Ridgway, R. 1882. Description of a new fly-catcher and a supposed new petrel from the Sandwich Islands. — Proceedings of the United States Na- tional Museum 4:337-338. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Rothschild, W. 1893. The avifauna of Laysan and the neighbouring islands: With a complete his- tory to date of the birds of the Hawaiian pos- sessions. Part 2. London, R. H. Porter, pp. 59- 126. Sclater, P.L. 1862. Catalogue ofa collection of Amer- ican birds. London, N. Trubner and Co., 368 pp. 1873. [Corrections to Sclater’s Catalogue of American Birds.]— Proceedings of the Zoolog- ical Society of London 1873:554—555. 1885. On the muscicapine genus Chasiem- pis. —Ibis, series 5, 3:17—19, plate 1. . 1888. [review] Stejneger on Hawaiian birds. — Ibis, series 5, 6:143-144. Sharpe, R. B. 1879. Catalogue of the birds in the British Museum. London, British Museum, 494 pp. Stejneger, L.H. 1887. Birds of Kauai Island, Hawai- ian Archipelago, collected by Mr. Valdemar Knudsen, with descriptions of new species.— Proceedings of the United States National Mu- seum 10:75-102. Warren, R. L. M., & C. J. O. Harrison. 1971. Type- specimens of birds in the British Museum (Nat- ural History). Volume 2, passerines. London, British Museum (Natural History), 628 pp. Wilson, S. B. 1891. On the muscicapine genus Chasi- empis, with a description of a new species.— Proceedings of the Zoological Society of London 1891:164-166. Department of Vertebrate Zoology, Na- tional Museum of Natural History, Smith- sonian Institution, Washington, D.C. 20560. PROC. BIOL. SOC. WASH. 102(3), 1989, pp. 559-567 A NEW LIZARD OF THE GENUS LEPIDODACTYLUS (REPTILIA: GEKKONIDAE) FROM BATAN ISLAND, PHILIPPINES Hidetoshi Ota and Ronald I. Crombie Abstract. — A new species of Lepidodactylus, L. balioburius, is described from Batan Island in the northernmost archipelago of the Philippines. It is most closely related to the recently described L. yami from Lanyu Island, Taiwan. Morphological variation in both species is analyzed. The herpetofauna of mountainous north- ern Luzon and the island groups stretching north toward Taiwan remains poorly known, despite intensive field work in other parts of the Philippines during the past 20-30 years. The Batan Island group, the north- ernmost archipelago in the Philippines, is of considerable zoogeographic interest since it is located almost midway between the Philippines and Taiwan (220 km north of Luzon and 200 km south of Lanyu Island, see Fig. 1). A few specimens from Batan reported in the literature perished when the Bureau of Science collection in Manila was destroyed during World War II. During May and June 1985, a multidisciplinary team of biologists, coordinated by Angel C. Alcala (Silliman University) and Charles A. Ross (Smithsonian Institution), collected on sev- eral islands in the Batan Group. Their col- lections contained a small series of a dis- tinctive new species of Lepidodactylus, a genus unreported from the extreme north- ern Philippines. The new species is super- ficially similar to the recently described L. yami from Lanyu Island, Taiwan (Ota 1987). Materials and Methods Data were taken from the series of Lep- idodactylus from Batan Island (n = 14) and all other Philippine/Taiwanese species of the genus (see Specimens Examined). Addition- al information was taken from Brown & Alcala (1978). Nine meristic and 21 mor- phometric characters were used for com- parisons. The meristic characters are the number of: upper labials (UL), lower labials (LL), internasal scales (INS), interorbital scales (IOS), midbody scale rows (MSR), enlarged preanal and femoral scales (PFS), preanal and femoral pores in males (PFP), toe I scansors (TIS), toe IV scansors (TIVS). The mensural characters are: snout to vent length (SVL), head length (HL), head width (HW), head depth (HD), snout to eye length (SEL), eye diameter (ED), eye to ear length (EEL), internasal distance (IND), interor- bital distance (IOD), snout to arm length (SAL), axilla to groin length (AGL), body width (BW), body depth (BD), thigh length (THL), tibia length (TBL), toe I length (TIL), toe IV length (TIVL), toe IV width (TIVW), the length of scansor series beneath toe IV (SL), tail width (TW), and tail depth (TD). All morphometric characters were mea- sured to the nearest 0.1 mm with dial cal- ipers. Meristics were compared using Wil- coxon’s 2-sample test and morphometric characters were examined by principal com- ponent analysis, using the PRICOMP pro- cedure of SAS (1985) with correlation ma- 560 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON t.. Philippines 2" 8) Fig. 1. Map of the Philippines and Taiwan, showing the type locality of Lepidodactylus balioburius sp. nov. (Batan Island, inset), in relation to the distribution of its close relative L. yami (Lanyu Island, inset) and L. planicaudus (shaded portions in the central and the southern Philippines). trix. Skeletal characters were examined in radiographs. Museum acronyms follow Leviton et al. (1985). Lepidodactylus balioburius, new species Fis2 Holotype. — Philippine National Museum (PNM) 984 (Original number USNM-FS 121559), an adult male collected 2 km (by road) SE of Mahatao, Mahatao Municipal- ity, Batan Island, Batanes Province, Phil- ippines, on 6 Jun 1985, by Angel C. Alcala, Ven Samarita, and Braulio Gargar. Paratypes.—(n = 13, all from Batan Is- land). USNM 266559, 3 km NE of Basco, collected by Charles A. Ross & B. Gargar on 28 May 1985; OMNH 2349 (USNM-FS 121200), 1-2 km E of Basco along road to Balugdh Bay, A. C. Alcala & D. Catada, 27 May 1985; USNM 266560-61, 2.5 km ENE of Basco on W slope of Mt. Iraya, 150 m, Robert S. Kennedy & Fred G. Thompson, 30 May 1985; OMNH 2348 (USNM-FS 121372), Basco, C. A. Ross, 31 May 1985; USNM 266562, 1.5 km N of Basco, near airstrip, C. A. Ross & A. C. Alcala, 4 Jun 1985; USNM 266563, Itbud, C. A. Ross & B. Gargar, 5 Jun 1985; USNM 266564, 3 km ENE of Basco, W slope Mt. Iraya, 150 m, C. A. Ross & R. S. Kennedy, 7 Jun 1985; CAS 162489, Mahatao, A. C. Alcala, 8 Jun 1985; USNM 266565-67, CAS 162490, 2 km E of Mahatao, C. A. Ross & A. C. Alcala, VOLUME 102, NUMBER 3 561 D Fig. 2. Holotype (PNM 984) of Lepidodactylus balioburius, adult male. A) dorsal view (scale = 10 mm), B) lateral and C) ventral views of snout (scale = 1 mm), D) ventral view of the preanal and femoral region, showing pores and enlarged scales (scale = 2 mm). 6 Jun 1985. (five males and eight females, all adults). Etymology.—The specific name is de- rived from the Latin roots balius (brown) and burius (beast), an appropriate descrip- tor for this species. The name also acknowl- edges the significant contributions to Phil- ippine herpetology by Walter C. Brown and Charles A. Ross. Diagnesis.—A small (males 27.2-34.9, females 33.5—38.7 mm), bisexual, Group III (sensu Brown & Parker 1977) species of Lepidodactylus, characterized by slight but distinct digital webbing, relatively few en- larged scales and pores in the femoral/ preanal region, the rostral separated from the nostril by a scale, presence of lateral serration on the tail, and the absence of dis- tinctive pattern elements (see Figs. 2, 3). Description of holotype.—Habitus mod- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 3. & D). erately depressed, SVL 30.4 mm. Snout ta- pering, rounded at tip, length 3.9 mm. Eye diameter 2.1 mm. Internasal distance 1.6 mm. Rostral separated from nostril by a small quadrangular scale. Nostril surround- ed by two supranasals, first upper labial, one small scale anteriorly and a slightly enlarged scale posteriorly. Anterior supranasals sep- arated by two small scales that border the rostral. Eleven upper labials on the right, 12 on the left, the 9th beneath the center of the orbit; the last 2 only about twice as large as the surrounding scales. Eleven lower labials. Mental triangular, smaller than adjacent la- bials. Three to 5 rows of slightly enlarged scales on anterior part of chin. Scales on the snout larger than those on the dorsal surface of the body. Dorsal and lateral body scales very small, granular, with no enlarged tu- bercles. Forty-one interorbital scale rows at the midpoint of orbits, 131 scale rows at midbody. Ventral scales flat, cycloid, dis- tinctly larger than dorsals. Ventral view of toes and lateral tail margins of Lepidodactylus balioburius (A & C) and L. yami (B Length of extended hind limb 11.0 mm. Digits moderately dilated, distal three-fifths to three-quarters of undersurface bearing scansors as follows: fingers—I 7, II 8, III 10, IV 11 (ight) or 12 (left), V 8; toes—I 8, I 9, II 11 eft) or 12 (ight), IV 9 (ight) or 10 (left), V 8 (left) or 9 (right). Distal two to three scansors, including the terminal one, divided on all digits except the first. First digit with complete terminal and two divided subterminal scansors. All digits ex- cept the first clawed. Compressed claw- bearing phalanges arising from distal mar- gin of the dilated part and extending only a short distance beyond. Phalangeal formula of hand and foot 2-3-4-5-3. Webbing slight but evident between toes III and IV, ex- tending to about one-eighth to one-fifth length of toe IV. Twenty-three enlarged preanal and fem- oral scales bearing a continuous series of 21 pores, extending over proximal 60% of thigh. Series of pore-bearing scales followed by one VOLUME 102, NUMBER 3 row of slightly enlarged scales on thigh, and four to five rows of enlarged scales in pre- anal region. Two pairs of cloacal spurs on both sides of vent. Tail unregenerated, moderately depressed; its depth just pos- terior to the basal swollen area 73% of its width; lateral flange of skin lacking, but en- larged, spine-like scales present every five to seven marginal scales. Scales on the ven- tral surface of the tail slightly larger than those on dorsal surface. Interclavicle dag- ger-shaped, without lateral projections. Clavicles perforated. Twenty-six presacral vertebrae. Nasals fused at midline. Color in alcohol.—Dorsal ground color light grayish tan, with numerous minute dark dots; slightly darker areas on snout and be- tween orbits; a wide, indistinct, dark band from the tip of the snout, through the nos- tril, eye, along the dorsal margin of the ear, and fading out between the ear and fore- limb. Several dark spots on upper and lower labial regions. Indistinct rusty gray mark- ings forming vague dorso-lateral lines. Ven- ter creamy white, with minute blackish dots, much sparser than those on dorsum. Tail with 10 dark gray annular bands. Variation. — Variation in counts and measurements of the type series is presented in Tables 1 and 2. In the five specimens with regenerated tails the enlarged, spine- like lateral scales are absent. Coloration in the series is variable. In two specimens, the dorsal ground color is much darker and more rusty than the holotype, with a lighter middorsal region forming a broad longitudinal stripe. The dark dorso- lateral markings and annular bands on the tail are indistinct or absent in seven speci- mens. In two others, however, these mark- ings are more distinct than in the holotype. Four specimens have black spots on the lat- eral region of the original tail and on the neck. Natural history.—Specimens were col- lected in both disturbed and forested hab- itats from sea level to 150 m on Mt. Iraya. The field notes of C. A. Ross indicate that Table 1.—Comparison of nine meristic characters in L. balioburius sp. nov., L. yami, and other Philippine congeners. See the text for abbreviations. Data marked with an * taken from Brown and Alcala (1978:82-101 + table 6), but note that numbers in the text do not always agree with those in the table. In these cases, inclusive values were used. IOS MSR PFS PFP TIS TIVS INS LL UL Species 10.00 Jd 0.83 6-9 24.33 22.79 40.00 138.57 4.71 0.47 10.07 yal L. balioburius 0.68 9-11 HS Bee) Loy 19-23 a 20-25 S32 131-151 3.57 34-46 43.27 0.83 9-11 10.93 0.94 10-14 SD 4-5 4.20 0.68 range 8.40 0.74 7-9 pe) 19.63 139.60 AKO) L. yami 1.28 10-15 1.92 15-21 1.51 19-24 18-26* 20-28* 30-40* 32-40* 28-35* oo 126-151 3.81 36-49 0.88 10-12 0.80 11-14 10-13* 10-13* 11-13* 10-13* 10-13* SD 3-5 range 7-12* 7-9* 11-16* 12-17* 12-18* 7-9* 6-9* 18-34* 20-27* 26-40* 28-46* 21-32* 135-145* 165-168 34-42* 40-41 38-42 3-5* 4-5* 3-5* 3-5* 10-12* 10-12* range L. planicaudus L. christiani range 9-10* 9-11* 9-13* 120-140* 78-108* 120-140* 9-13* 9-13* 9-12* range L. aureolineatus L. herrei 24-34* 32-40* range 3-5* range L. lugubris 563 564 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 2.—Comparison of 21 morphometric characters of Lepidodactylus balioburius sp. nov. and L. yami, and the factor loadings on the first three principal components. See text for abbreviations. L. balioburius L. yami Characters x SD Range x SD Range PRIN I PRIN II PRIN III SVL 34.08 3.81 27.2-38.7 36.16 20 31.7-42.1 0.27 0.08 —0.94 HL 8.69 0.67 7.6—9.6 9.47 0.69 8.6-10.9 O27 0-01 —0.03 HW 6.36 0.47 5.6-7.1 6.76 0.95 6.0-9.8 0.25 0.15 0.10 HD 3.59 0.28 3.1-3.9 4.23 Erg 3.4-7.1 0.25 0.04 =O313 SEL 4.11 0:35 3.5-4.6 4.19 0.28 3.9-4.8 0.25 0.11 0.02 ED 20> 0.20 2.0—2.6 2.29 0.23 2.0-2.6 0.23 0.02 0.13 EEL 2:83 TOL 2.5-3.0 5.02. «uON3 2.8-3.4 0.24 —0.07 0.00 IND 163 ©.13 1.4-1.8 1.68 0.14 1.5-2.0 0.23 0.08 0.06 IOD 4.16 0.29 3.7-4.7 4.59 0.36 4.0-5.0 O:17, 023 =@.11 SAL [2259 1.30 10.6-14.8 13.44 1.05 11.5-15.1 0.24 0.10 0.15 AGL 16.19 23 12.7-19.3 17.81 1.82 14.9-21.5 0.24 0.04 —0.38 BW 8.23 1.25 6.2-9.9 7.04 1.23 5.6-9.4 0.05 0.48 =0:35 BD 4.23 0.86 2.8-5.4 5.25 Ipallig/ 4.0-8.6 0.23 0.01 —0.36 THL 4.43 0.60 3.5-5.3 4.80 0.68 3.5-5.8 0.24 0.06 —0.10 TBL 4.38 0.38 3.64.8 4.90 0.36 4.3-5.4 0.26" —0: 1 —0.09 Te 1.65 0.21 1.42.1 1.95 0.16 1.7—2.2 O22- = =—O0H9 0.12 TIVL 3305 0.37 2.4-3.7 3.72 0.34 3.0-4.2 O22 =" O71 0.29 TIVW eal Osh 1.0-1.6 1.19 0.14 0.9-1.5 0.20 0.00 0.51 SL BMD 0.36 1.6-2.9 2.10. “O21 2.2-2.9 Onl 94 —0:29 —0.00 TW 4.51 0.64 2.9-5.6 3.50 0.45 2.7-4.1 0.02 0.53 0.11 TD Zo 0.32 2.3-3.4 2.82) 10:37 2.1-3.6 0.06 0.44 0.34 Eigenvalue 12.34 3.20 E15 Difference oS 2.06 O23 Proportion 0.59 0.15 0.05 Cum. prop. 0.59 0.74 0.79 the species was commonly found under loose bark on trees during the day, in disturbed areas of fields and gardens, in coastal vege- tation, and along a forested stream. Of those found after dark, one was active on the guest house in Basco, and another was on the un- derside of a banana leaf in forest on Mt. Iraya. Two eggs (USNM 266568), four hatched eggshells (not collected), and an adult were found under bark in a ravine running through a garden area, eventually leading to forest. Other gekkonids collected with L. balioburius include Hemidactylus frenatus, Gehyra mutilata, and Gekko po- rosus. Since L. balioburius is ecologically tolerant and not restricted to forest, it is likely that it will be found on other islands in the Batan group, none of which has been adequately collected to date. Distribution.—Known only from Batan Island, Batanes Province, Philippines, but expected on other islands in the group (see above). Remarks. —Brown & Alcala (1978) con- veniently separated the Philippine species of Lepidodactylus into two species com- plexes (=Sections), based largely on digital morphology, habitus, and caudal scalation. Section A (consisting of aureolineatus, her- rei and lugubris) was characterized by high scansor counts (11-18, usually more than 12), the scansors covering most of the mod- erately to broadly dilated digits, slight dig- ital webbing, moderately depressed habitus, and a slightly to moderately flattened tail with lateral denticulation. Section B species (christiani, planicaudus) have fewer scan- sors (7-10, usually less than 10), confined to the distal half of broadly dilated and strongly webbed digits. The body and tail are strongly flattened, the latter with a broad flange of skin. VOLUME 102, NUMBER 3 Ota (1987) placed his new species, L. yami from Lanyu Island, Taiwan, in Section A, but with some reservations. He also re- stressed the importance of the nostril po- sition relative to the rostral (in contact in Section A species, separated by a scale in christiani and yami, separated or in contact in planicaudus). Brown & Alcala (1978:81) dismissed this character as having “‘little significance at the species level,’ due to the variation in planicaudus, but our data in- dicate that it may be more useful than Brown & Alcala thought. Lepidodactylus balioburius shares the na- sal-rostral separation character with L. yami and the Section B species, but has little else in common with the latter. Consequently, it requires comparison only with Section A species. Within this group, L. balioburius has lower PFS, PFP, TIS, and TIVS counts than all species except L. yami (see Table 1). Lepidodactylus aureolineatus and herrei are further distinguished by the presence of bright head stripes beginning on the snout and extending to near the ear. Lepidodac- tylus lugubris is a functionally all-female, parthenogenetic species; the few males re- ported to date have been sterile (Cuellar & Kluge 1972, Pasteur et al. 1987). In color and pattern, L. /ugubris is often very pale, almost white, with a variable pattern of dark spots, occasionally resembling those in the much darker L. balioburius. Lepidodactylus lugubris is capable of color change, how- ever. When in the dark phase, a more com- plex, ladder-like pattern becomes obvious, but this pattern is distinctly different from the plain brown, spotted balioburius. Lepidodactylus balioburius and L. yami share a number of characters unique in Sec- tion A species. Both are small, brown, un- specialized species isolated on small islands far to the north of their Philippine congeners (reports of L. /ugubris from Taiwan and as- sociated islands are possibly recent intro- ductions [Ota 1986, Cheng 1987]). Besides the similar habitus, coloration, and nostril- rostral separation, UL, MSR, PFS, and PFP 565 showed no statistically significant differ- ences between the two species (P = 0.05). Although the ranges overlapped somewhat, the means of LL, INS, IOS, TIS, and TIVS were significantly different (Table 1); INS of L. balioburius was larger than yami (P < 0.05), whereas LL (P = 0.05), IOS (0.05), TIS (0.05), and TIVS (0.001) were larger in L. yami. Principal component analysis of morphometric characters revealed approx- imately 80% of the total variation in shape as expressed in the first three components— PRINs I, II and III. Of these, PRIN I was of little use in separating L. balioburius from L. yami. This component consists wholly of positive variable loadings, and is pri- marily a size component (Table 2). PRIN II, accounting for about 15% of the total variance, tends to discriminate L. balio- burius from L. yami more strongly. Variable loadings on this component revealed sev- eral characters chiefly contributing to the shape. The greatest proportion of the vari- ance on PRIN II was expressed by differ- ences in TW. The BW and TD were also heavily loaded, and followed by SL, IOD, TIVL, TIL and HW in descending order. Outlines of scatter plots of the component scores on PRINs I and II separated L. ba- lioburius from L. yami without overlap (Fig. 4). The two species can also be distinguished by toe webbing and lateral caudal scalation, although these two characters are more sub- jective. Lepidodactylus balioburius has slightly more extensive webbing than L. yami (Fig. 3A, B) and the lateral denticu- lation of the original tail of balioburius con- sists of strongly enlarged, spine-like scales (Fig. 3C). In L. yami, slightly enlarged scales are present along the lateral tail edge, but they are neither projecting nor spinose (Fig. 3D). We originally intended to summarize and discuss the relationships of Philippine- Taiwanese Lepidodactylus in this paper. Unfortunately, the unprecedented variation in L. planicaudus requires further re-eval- uation so we defer our taxonomic summary and key to a later paper. 566 9 -8 -7 6-5 -4 3 PRIN | Fig. 4. Two-dimensional plots of scores of Lepidodactylus balioburius (closed circles) and L. yami (open circles) on principal components (PRINs) I and II. See Table 2 for the factor loadings of each component. The apparent absence of Lepidodactylus from the large island of Luzon is zoogeo- graphically puzzling. Although L. planicau- dus is found on small islands both east and southwest of Luzon (Brown & Alcala 1978, see also Fig. 1), the genus remains unre- ported from the entire large island; even the widespread human commensal L. lugubris is absent from its towns and cities. Although Luzon has been reasonably well collected (Taylor 1922; Brown & Alcala 1970, 1978), this hiatus may not be real. Many Philippine Lepidodactylus are ecologically restricted and difficult to collect. They may inhabit axils of palms or aerial ferns 10 meters or more from the ground. The montane areas of northern Luzon have never been ade- quately sampled and we would not be sur- prised if a new species allied to L. balio- burius and yami is eventually discovered in that area. Specimens examined.—L. yami: Osaka Museum of Natural History (OMNH) R2291 (holotype), 690, 691, R2855-61, California Academy of Sciences (CAS) 158254 (paratypes), USNM 267943-44; L. planicaudus: CAS 60570, 128566, 139930, 2-10 4 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 12 +3 +4 +5 +6 139931, L. christiani: CAS 128877, 128878. L. aureolineatus: CAS-SU 28411, 26127, CAS 60226, 139941. L. h. herrei: CAS-SU 24228, 26342. L. h. medianus: CAS 125239, 131856 (paratypes). L. lugubris: OMNH R1772, 2201, 2202, 2320, 232 ers 158255, 60595, 137835, Australian Mu- seum (AMS) R82602-82610, 82724-82730, 109804—109809, 110141-110146, 110238- 110242. Acknowledgments We are greatly indebted to Angel C. AI- cala, Walter C. Brown, and Charles A. Ross for their unstinting cooperation in provid- ing notes, unpublished data, and specimens for our use. We have enjoyed stimulating discussions on the Philippine fauna with the above individuals, Lawrence R. Heaney, Robert S. Kennedy, and Fred G. Thomp- son. Yasuhiko Shibata (OMNH), Jens V. Vindum (CAS), Harold Cogger and Allen E. Greer (AM) kindly loaned specimens for comparative purposes. Special thanks are due Tsutomu Hikida for computing the data, using the facilities of the Data Processing VOLUME 102, NUMBER 3 Center, Kyoto University. Radiographs were made with facilities of Biological Labora- tory, Yoshida College, Kyoto University through the courtesy of Masafumi Matsui. The Batan field work was made possible by a Smithsonian Research Opportunity Fund award to C. A. Ross. Crombie’s mu- seum work on the Pacific herpetofauna was also supported by two ROF grants, for which David Challinor has our thanks. Ota’s work was supported in part by a Grant-in-Aid for Special Project Research on Biological As- pects of Optimal Strategy and Social Struc- ture from the Japan Ministry of Education, Science and Culture. We also thank W. Ronald Heyer and George R. Zug for their critical reading of the manuscript and Linda K. Gordon for valuable assistance in producing the final manuscript copy. Literature Cited Brown, W. C., & A. Alcala. 1970. The zoogeography of the herpetofauna of the Philippine Islands, a fringing archipelago.— Proceedings of the Cali- fornia Academy of Sciences 38:105-—130. —,, & 1978. Philippine lizards of the Family Gekkonidae. Silliman University Nat- ural Science Monograph Series (1), Dumaguete City, Philippines, 146 pp. —., & F. Parker. 1977. Lizards of the genus Lep- idodactylus from the Indo-Australian archipel- ago and the islands of the Pacific, with descrip- tions of new species.— Proceedings of the California Academy of Sciences 41:253-265. 567 Cheng, H.-Y. 1987. The record of a gekkonid lizard Lepidodactylus lugubris (Dumeril and Bibron, 1836) from Taiwan.—Journal of the Taiwan Museum 40(1):85-89. Cuellar, O., & A. G. Kluge. 1972. Natural partheno- genesis in the gekkonid lizard Lepidodactylus lugubris.—Journal of Genetics 61:14—26. Leviton, A. E., R. H. Gibbs, Jr., E. Heal, & C. E. Dawson. 1985. Standards in herpetology and ichthyology: Part I. Standard symbolic codes for institutional resource collections in herpetology and ichthyology.—Copeia (3):802-832. Ota, H. 1986. The mourning gecko Lepidodactylus lugubris (Dumeril and Bibron, 1836); an addi- tion to the herpetofauna of Taiwan. —Journal of the Taiwan Museum 39(1):55—58. . 1987. Anew species of Lepidodactylus (Gek- konidae: Reptilia) from Lanyu Island, Tai- wan.—Copeia (1):164-169. Pasteur, G., J.-F. Agnese, C. P. Blanc, & N. Pasteur. 1987. Polyclony and low relative heterozygos- ity in a widespread unisexual vertebrate, Lepi- dodactylus lugubris (Sauria).—Genetica 75:71- 79. 1985. SAS user’s guide: Statistics, version 5. SAS Institute Inc., Cary, North Carolina. Taylor, E. H. 1922. The lizards of the Philippine Islands. — Philippine Bureau of Science Publ. No. 17:1-269. SAS. (HO)Department of Zoology, Faculty of Science, Kyoto University, Kitashirakawa, Sakyo, Kyoto, 606 Japan (Present address, Department of Biology, University of the Ryukyus, Nishihara-cho, Okinawa 903-01, Japan); (RIC) Department of Vertebrate Zoology (Amphibians & Reptiles), National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560. PROC. BIOL. SOC. WASH. 102(3), 1989, pp. 568-576 A NEW SPECIES OF EUPSOPHUS (AMPHIBIA: ANURA: LEPTODACTYLIDAE) FROM SOUTHERN CHILE J. Ramon Formas Abstract.—Eupsophus emiliopugini, a new species of leptodactylid frog, is described from southern Chile. This species is distinguished from its congeners by the olive-green cephalic pigmentation. The mating call is composed of two notes and the tadpole is typically an inhabitant of ground water-filled cavities. This frog occurs from eastern Osorno Province through Llanquihue, Chiloé, and Aisén provinces. The genus Eupsophus Fitzinger, 1843 is restricted to the temperate Nothofagus for- est of the southwestern border of South America. At present five species have been described (E. roseus, E. vittatus, E. calca- ratus, E. migueli, and E. insularis) (Formas & Vera 1982, Formas 1985), and for this reason Eupsophus can be considered the most diverse genus within the reduced an- uran fauna of the temperate forest system of southern Chile and Argentina. Among the genera of the sub-family Telmatobiinae, Eupsophus is remarkable by having tad- poles (E. roseus, E. vittatus, and E. calca- ratus) which live in small water-filled cav- ities in the ground. The larval mouthparts (denticles) are reduced, and the larvae feed only upon yolk reserves (Formas & Pugin 1978a, b; Formas 1989). Between September 1975 and December 1987, Carlos Varela, Lila Brieva, Gonzalo Aguilar and I collected specimens of a new species of Eupsophus in the temperate Noth- ofagus forest of southern Chile. In this paper the new taxon is described on the basis of a series of materials (adults, juveniles, mat- ing call, eggs, and tadpoles) collected from seventeen localities. Eupsophus emiliopugini, new species Fig. 1 Holotype. —IZUA (Instituto de Zoologia, Universidad Austral de Chile) 1587, adult male; Ramon Formas, Nov 28, 1975, at La Picada, 23 km NE (by road) of Ensenada (Fig. 2), Osorno Province, Andean Range, 41°04’S, 72°26’W, 480 m. Paratypes. —Seven adult males from the type locality: IZUA 1585-86, 1593, 1596, 1602, 1607-8. Diagnosis.—Eupsophus emiliopugini is a medium-sized frog which is characterized by having a distinctive olive-green band be- tween the eyes, which is absent in E. roseus, E. calcaratus, E. vittatus, E. migueli and E. insularis. Furthermore it differs from E. vit- tatus in snout—vent length (Table 1). The mating call of E. emiliopugini has two notes while that of E. vittatus possesses five (4—6) notes (Table 2). Adult description. —(Based on 25 living frogs and 29 fixed specimens.) Head wider than long. Snout pointed in dorsal view, slightly sloping in lateral profile; canthus rostralis indistinct and rounded; loreal re- gion slightly concave; nostrils dorso-lateral, closer to tip of snout than to the orbit; length of the eye greater than distance between eye and nostril; inter-orbital distance narrower than length of eye, greater than internarial distance. Tympanic membranae medium and well defined, tympanum diameter three fourths the distance between eye and nostril. Well developed supratympanic fold extend- ing posteriorly from the corner of the eye to the posterior part of the tympanum, cov- VOLUME 102, NUMBER 3 569 Figc ft. ering upper part of tympanic annulus but not reaching insertion of arm. Tongue large, ovoid, with notch at the tip. Choanae small, round dentigerous process of vomers lying slightly below choanae; each process bear- ing 3-4 slightly oblique teeth close to me- dian line. Forelimbs thin. First finger equal in length to second; third finger much longer than fourth; digital length in decreasing or- der 3-4-2-1. Palmar webbing absent; tips of fingers rounded and slightly prominent. In- ner median palmar tubercle ovoid; outer palmar tubercle horseshoe-shaped; subar- ticular tubercles globular and moderate in size; supernumerary palmar tubercles pres- ent. Hind limbs slender. Toes long, slender, and moderately fringed; tips of toes round- ed; third and fifth toes equal in length; toes in decreasing order of length 4-(3,5)-2-1. In- ner metatarsal tubercle ovoid and promi- nent, outer rudimentary. Subarticular tu- bercles ovoid; supernumerary tubercles absent. Tarsal fold absent. Rudiment of web among toes. Anal opening oriented trans- versely and directed postero-ventrally at dorsal level of thighs. Dorsal and ventral skin smooth. Two weakly developed para- Eupsophus emiliopugini, new species. Holotype (IZUA 1587). vertebral folds extending from posterior part of head, converging behind it. Post-tym- panic areas and flanks with scarce minute granules. Some specimens with slight gran- ular area around vent and posterior part of thighs. External measurements of males and females of this species shown in Table 1. Coloration in preservative. —Dorsal ground color light gray with few whitish mi- nute spots; a delicate vertebral line extend- ing from the tip of snout to the vent. A dark greenish band on the eyes. Venter whitish and gular area gray. Lips gray with two or three irregular spots reaching the tympanic region. Arms light gray and crossed by two or three irregular transverse bars of dark gray color in dorsal area; arms and legs whit- ish ventrally. Coloration in life. —Dorsal ground color grayish brown to leaden; vertebral line lem- on-yellow. Band on eyes olive-green col- ored. Some specimens with bright yellowish reticulations on the thighs. Belly whitish and the gular area of mature males bright or- ange. Distribution. —The presently known range of E. emiliopugini extends from northeast- 570 Table 1.—Comparative measurements (mm) of Eupsophus emiliopugini and E. vittatus. Means, SD, and ranges (parenthesis). PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON E. emiliopugini E. vittatus Character Females Males Females Males n 14 40 19 19 Snout—vent length SOLO == 5:01 46.71 + 24.00 59 Olne=s6. 58 55.44 + 8.68 (41.0-64.0) (42.9-50.0) (47.0-71.8) (44.5-66.6) Head length PES See yl 282 16.45 + 0.69 20.43 + 1.50 £3.36 2 23350 (13.4—20.8) (14.8-18.1) (16.7-23.9) (15.4-21.3) Head width PGES Sect Deng 18.48 + 0.33 23:66 = 2:02 DAT 2 E5238 (15.9-25.6) (16.9-19.1) (19.9-26.6) (18.1-26.0) Femur length 23:38 2247 2S6 == 06 28.45 + 1.89 26.37 = 3:85 (17.1-29.2) (16.6-24.3) (24.7-31.2) (21.7-31.6) Tibia length DAD = 2) 24.25 + 4.43 D8: 985 2.05 27.2 + 4.18 (19.9-29.6) (19.30-33.1) (25.0-32.2) (22.8-33.1) Foot length Soe ee S50 365222110 41.83 + 3.56 Al .26 225337 (29.1-44.9) (29.7-49.1) (34.4-48.2) (33.8-49.1) ern Osorno Province (Termas de Puyehue) to Aisén Province (Caleta Vidal) (Fig. 2). This area is covered by humid and cool Nothofagus forests, which are found at the Andes Cordillera (below 1000 m), the Coastal Range, the Central Valley, and on Chiloé Island. The altitudinal distribution of E. emiliopugini ranges between the sea level (Caleta Vidal) and 700 m in the Andes Cordillera (Termas de Puyehue). Over much of its range, E. emiliopugini occurs sym- patrically with E. calcaratus. It has never been found in sympatry with E. vittatus. Habitat. —La Picada (type locality) (Fig. 2) is a small subandean valley surrounded by forests (Nothofagus, Aextoxicum, and Eucryphia). During winter and summer, frogs were found there under logs, however during the reproductive period (springtime) the animals were collected at the border of a small stream shaded by ferns (Dryopteris) and mosses (Sphagnum and Hygroambly- stegium). From the ecological point of view this area 1s situated in the oceanic region of med- iterranean influence (di Castri 1968). The annual mean temperature of this region is 10.5°C the relative humidity is 84% and the rainfall ranges between 2000 to 2500 mm. Of seventeen examined localities, fifteen are included in the preceding region; however the southernmost localities (Rio Cisnes and Caleta Vidal) are situated in the oceanic cold- temperate region (di Castri 1968). The an- nual mean temperature is 8.8°C, the relative humidity is 87% and the rainfall ranges from 2500 to 3000 mm. At the type locality the following species of amphibians were also collected: Rhino- derma darwinii, Bufo variegatus, Batrachyla leptopus, B. antartandica, Alsodes monti- cola, Pleurodema thaul, Hylorina sylvatica and Eupsophus calcaratus. Breeding sites and breeding season. —Ten breeding sites were examined (Termas de Puyehue, La Picada, El Traiguén, Rio Rol- — Fig. 2. Distribution of Eupsophus emiliopugini (dots) and E. vittatus (triangles) in Southern Chile. Arrow indicates location of the type locality (La Picada) of E. emiliopugini. 1) Ramadillas, 2) Contulmo, 3) Mafil, 4) Linguento, 5) Mehuin, 6) San Martin, 7) Los Molinos, 8) Valdivia, 9) Huellelhue, 10) Tres Chiflones, 11) Cordillera Pelada, 12) Bahia Mansa, 13) Pucatrihue, 14) Puyehue, 15) Piedras Negras, 16) El Traiguén, 17) La Picada, 18) Rio Rollizo, 19) Rio Lenca, 20) Camino Maullin, 21) Ancud, 22) Lechagua, 23) Chepu, 24) Cucao, 25) Puntra, 26) Tepuhueico, 27) Yaldad, 28) Quellon, 29) Rio Cisnes, 30) Caleta Vidal. VOLUME 102, NUMBER 3 @ Frutillar Llanqu/hue Lake © Valdivia At0 : ' Ue -¥ i _ Scanning electron micrographs of Nocticanace texensis: 13, Head, lateral view; 14, Head, anterior quately outline characters to distinguish be- tween them and other congeners. The only distinguishing characters between the three species of the fexensis group that I have found are those of the male terminalia. Nocticanace texensis (Wheeler) Figs. 13-27 Canaceoides texensis Wheeler, 1952:92. Nocticanace texensis.— Wirth, 1954:62 [ge- neric combination]; 1965:734 [nearctic catalog]; 1975:3 [neotropical catalog]. Diagnosis.—Small to moderately small beach flies, length 1.60 to 2.95 mm (holo- 596 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Figs. 22-27. and different localities, lateral views: 22, Belize, Stann Creek District, Carrie Bow; 23, USA, North Carolina, Wrightsville; 24, USA, North Carolina, Wrightsville; 25, Mexico, Tabasco, Paraiso (5 km N); 26, Antigua, Dutchman Bay; 27, Dominica, David Bay. type 2.15 mm). Coloration generally brown dorsally and gray laterally. Head: Frons generally brown to oliva- ceous or slightly charcoal brown, coloration of frons uniform or ocellar triangle more grayish, and fronto-orbits charcoal gray. Ocellar setae divergent, lateroclinate; in- trafrontal setae anteroclinate, slightly con- vergent; ocellar area with three or four smaller setulae. Face, clypeus, and gena (to an extent) whitish, with faint tinges of blue or gray, gena becoming darker posteriorly, more charcoal gray. Orientation and size of genal setae as follows (anterior to posterior setae): lst seta large, anteroclinate; two large anaclinate setae with one, rarely zero or two anaclinate setulae between larger anaclinate setae; anaclinate setula about one-third length of larger setae. External male terminalia of Nocticanace texensis: Variation in shape of the surstylus from same Thorax: Mesonotum from dorsal view mostly brown to olivaceous brown, darker around anterior margins; postpronotum, anterior half of notopleuron, and to a lesser degree the posterior fourth of scutum, and scutellum gray; coloration of thorax in lat- eral view brown dorsally, becoming gray ventrally around area of notopleuron (in some specimens just above, others at or be- low notopleuron), remainder of pleural areas mostly unicolorous, gray. Notopleuron bearing only one seta, inserted posteriorly; anepisternum with three large setae and from three to five smaller setulae between larger ones, larger setae with insertions and ori- entations as follows: posterior seta, usually the largest, with posteroclinate orientation, ventral seta, usually the next largest, with declinate orientation, dorsal seta smallest VOLUME 102, NUMBER 3 (not much larger than setulae) and with an- aclinate orientation; one large katepisternal seta and one or two smaller setulae inserted ventrad or anteroventrad. Fore femur with four or five long setae along posteroventral margin and four or five shorter setae pos- terodorsally; mid tibia bearing apicoventral spinelike seta, length about equal to tibial width; hind basitarsomere lacking basoven- tral pair of larger setae; legs mostly concol- orous, gray to dark gray apically. Wing gen- erally lightly infumate, brownish; costal vein ratio averaging 0.13 (holotype 0.15); M vein index averaging 0.43 (holotype 0.49); apical section of vein CuA, slightly more than twice length of crossvein dm-cu. Abdomen: Dorsum mostly olivaceous brown, gray laterally and ventrally. Male terminalia (Figs. 22—27) as follows: poste- rior margin of surstylus densely setulose, posterior half of surstylus in lateral view roughly triangular, distinctly angulate to form a large and wide posterior projection and a tapered, more slender process ven- trally (Sometimes curved inward), anterior margin with a subapical emargination (see remarks section below for discussion of variation). Type material.—The holotype male is la- beled “‘Galveston[,] Tex[as] 9.13.50 [hand- written|/M. R. Wheeler Collectors/Holo- type Canaceoides texensis Wheeler 1952 [red; handwritten].’’ The allotype female (not examined) is reported to bear the same lo- cality label data as the holotype. The ho- lotype is double mounted (glued to a paper point), is in excellent condition (the abdo- men has been removed, dissected, and the parts are stored in an attached plastic mi- crovial), and is deposited in the California Academy of Sciences. Other specimens examined.— Antigua. Dutchman Bay, 7 Jan 1965, W. W. Wirth (4 6; USNM). Belize. Stann Creek District: Carrie Bow, 4 Mar 1984, 2 Jun 1985, 15- 27 Jan 1987, W. N. Mathis, C. Feller (18 4, 9 2; USNM). Dominica. Calibishie (sea shore), 27 Feb—22 Mar 1965, 1989, W. N. 597 Mathis, W. W. Wirth (15 6, 3 2; USNM); Coulibistri, beach, 21 Mar 1989, W. N. Mathis (23 6, 10 2; USNM); Layou River (mouth), 9 Jan 1965, W. W. Wirth (1 2; USNM); Macoucheri (sea shore), 1 Feb 1965, W. W. Wirth (1 2; USNM); Pagua Bay, 18 Feb 1965, W. W. Wirth (6 4, 5 2; USNM); Rosalie (cobble beach), 23 Mar 1989, W. N. Mathis (7 3, 4 2; USNM); Sou- friére Bay, 24 Mar 1989, W. N. Mathis (9 6, 6 2; USNM); St. David Bay (sea shore), 23 Jan 1965; W. Ws Wirth (15. 6, 10.9; USNM). Mexico. Tabasco Province: Para- iso (5 km E), 6 May 1985, W. N. Mathis, A. Freidberg (26 6, 25 2; USNM). St. Vin- cent. Buccament Bay, 25-28 Mar 1989, W. N. Mathis (7 6, 4 2; USNM); Cumberland Bay, 28 Mar 1989, W. N. Mathis (4 34; USNM); Wallilabou (beach), 27 Mar 1989, W. N. Mathis (8 6, 4 2; USNM). United States. North Carolina: Wrightsville Beach (light trap), 3-7 Sep 1953, R. H. Foote (9 6, 23 2; USNM). Distribution. —East (North Carolina south) and Gulf coasts of North America, West Indies (Antigua, Dominica, and St. Vincent), and Caribbean coast of Mexico and Belize. Natural history.—All of the specimens. from Belize were collected on Carrie Bow Cay, formerly Ellen Cay, which is a highly disturbed, vegetated sand cay that is about 18 km off the coast of Hopkins, Stann Creek District. The cay is small, 100 by 40 m, and the former woodland, mostly red man- grove, has been cleared. Twenty-two species of plants have been recorded from Carrie Bow, including a few that have been intro- duced. Stoddart et al. (1982) and a more recent paper by Rutzler and Ferraris (1982) provide more extensive information on Carrie Bow and its habitats. The specimens of N. texensis were col- lected by sweeping an aerial net with a fine- meshed bag immediately over and between rocks and exposed coral on the surrounding beach, which is mostly sand covered, es- pecially on the north, east, and south sides. 598 Remarks.—Externally, this species is very similar to N. wirthi but can be distinguished from it and other congeners by the following characters: anterior notopleural seta absent; acrostichal setae absent; apical scutellar se- tae nearly straight in lateral view, not dis- tinctly curved dorsally (anaclinate); and shape of the male terminalia, especially the unique surstylus (see figures). The shape of the surstylus is quite similar to that found in males of N. arnaudi Wirth (the galapagos group), but specimens of the latter are usually larger (3-3.5 mm), are much darker brown dorsally with the dark coloration extended ventrally to the dorsal third of the anepisternum, and the noto- pleuron usually bears an anterior and pos- terior seta, although the latter seta is usually much better developed. The shape of the surstylus also differs slightly from locality to locality (see Figs. 22-27 from different localities), and I in- terpret this to be intraspecific variation, per- haps clinal in nature. The differences found in populations from the Lesser Antilles, however, especially those from Dominica, could represent interspecific variation. The surstylus from a male collected in Antigua (Fig. 26), for example, appears to be some- what intermediate between that found on Dominica (Fig. 27) and the more typical shape, as found in males from North Car- olina, Texas, Mexico, and Belize (Figs. 22- 25). Unfortunately no specimens are avail- able from the Greater Antilles that could provide important evidence to help resolve this issue, and for the present, I prefer to recognize a single species. Nocticanace wirthi, new species Fig. 28 Nocticanace texensis [of authors, not Wheeler].— Wirth, 1954:61-62 [illustra- tion of male terminalia]; 1965:734 [nearctic catalog]. Diagnosis.—As in the preceding species except as follows: small to moderately small beach flies, length 1.75 to 2.25 mm. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Abdomen: Male terminalia as follows (Fig. 28): surstylus in lateral view with two large, ventral lobes; posteroventral lobe wide, pointed anteroventrally, and densely setulose, especially along outer margin; an- terior lobe blunt, thumblike, and setulose. Type material.— The holotype male is la- beled “‘Boyton Beach, Fl[orid]a [Palm Beach Co.]/WWWirth Collector/10 VIII 51 [10 Aug 1951] Intertidal rocks/é/Nocticanace texensis (Wheeler) det WWirth [species name handwritten; black sub-border]/HO- LOTYPE 4 Nocticanace wirthi Mathis USNM [species name handwritten; red].” The allotype female and 14 paratypes (7 4, 7 2; CAS, USNM) bear the same locality label data as the holotype. Other paratypes are as follows: Cuba. Matanzas Province: Playa Larga, 1 May 1983, W. N. Mathis (3 6, 9 2; IZAC, USNM). Bermuda. Smith’s Parish: Spittal Pond (intertidal rocks), 18 Nov 1987, N. E. Woodley (21 6, 25 2; BDAF, USNM). The holotype is double mounted (minute nadel in a cork block), is in good condition, and is in the Smithsonian Insti- tution (USNM). Distribution.—Bermuda, Cuba, southern Florida. Natural history.—Wirth (1954:62) re- ported that the type series was “... col- lected from a shelf of limestone rock about a hundred yards long on the Atlantic Ocean beach. The rock projected from the water only at low tide and was covered with a scanty growth of filamentous green algae.” The specimens collected in Cuba were swept from a large limestone outcrop that extend- ed into the sea. The limestone was deeply eroded, and the exposed surface had nu- merous sharp ridges. Etymology.—It is a pleasure to name this species after Dr. Willis W. Wirth, who has contributed substantially to our knowledge of beach flies and who first illustrated this species. Remarks. —This species is very similar to N. texensis, and the only distinguishing characters that I have found between these two species are those of the male terminalia, and VOLUME 102, NUMBER 3 599 Figs. 28-29. Lateral view of male terminalia: 28, Nocticanace wirthi; 29, Noticanace panamensis. especially the bilobed shape of the surstylus (see description and figures). Externally the specimens are virtually impossible to sep- arate from those of N. texensis. Nocticanace panamensis, new species Fig. 29 Diagnosis.—As in N. texensis except as follows: moderately small beach flies, length 2.0 mm (based on the single pinned 2 para- type). Abdomen: Male terminalia as follows (Fig. 29): surstylus in lateral view more or less rectangular, posterior margin conspic- uously setulose; inner lobe not developed as a process, bearing several setulae along me- dian surface; outer lobe a large, thumblike posteroventral process that bears several setulae. Type material.— The holotype male is la- beled ‘“[Panama] Mojinga Swampf,] Ft. Sherman, C. Z. Jan. 1953[,] F. S. Blanton ct. [collector] Nocticanace texensis (Whee- ler) 6 5 [°5” is circled][handwritten].’’ The allotype female (double mounted) and three paratypes (3 2; USNM;; slide mounted) bear the same label data as the holotype. The holotype is slide mounted in balsam (most body parts have been dissected and are sep- arated) and is in the Smithsonian Institution (USNM). Distribution.— Panama. Fort Sherman is located at the Caribbean mouth of the Pan- ama Canal. Etymology.—The specific epithet, pana- mensis, refers to the country of Panama, where the type series was collected. Remarks.—Like N. wirthi, this species dif- fers from N. texensis in characters of the male terminalia, especially the shape of the surstylus (see figures and description above). 600 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ey» NY wy, Af ACA , ~ Figs. 30-38. Scanning electron micrographs of Paracanace hoguei: 30, Head, lateral view; 31, Gena and setae, lateral view; 32, Head, dorsal view; 33, Antenna, lateral view; 34, Ocellar triangle, dorsal view; 35, Fronto- orbital setae, dorsal view; 36, Notopleuron and setae, lateral view; 37, Katepisternum and setae, lateral view; 38, Scutellum, dorsal view. Genus Paracanace Mathis & Wirth Paracanace Mathis & Wirth, 1978:524. Type species: P. hoguei Mathis & Wirth, by original designation. Diagnosis.—Small to moderately small beach flies, 1.40 to 2.60 mm; general col- oration whitish gray to brownish black. Head: Intrafrontal setae two pairs; post- ocellar setae well developed, proclinate and very slightly divergent, subequal in length to intrafrontal setae; ocelli arranged to form isosceles triangle, with greater distance be- tween posterior ocelli. Two to three large anaclinate genal setae; anteroclinate genal seta well developed, subequal in length to larger anaclinate genal setae; epistomal margin sinuate; clypeus low, width more than 4x height; palpus yellowish. Thorax: Acrostichal setulae present, in about four rows, with a distinctly larger pre- scutellar pair; scutellar disc lacking setae; apical scutellar setae not anaclinate; ante- rior notopleural seta usually present (very weak or absent in one species); proepisternal seta(e) present; anepisternum with scattered setulae; katepisternal seta present. Femora and tibiae grayish black; tarsomeres yellow- VOLUME 102, NUMBER 3 ish brown to dark brown, becoming darker apically; mid femur of male with comblike row of setae; hind tibia lacking spinelike setae apically. Wing with length of apical section of vein CuA, twice or more that of crossvein dm-cu; vein M index 0.38. Discussion.— Like Canacea, all of the de- scribed species of Paracanace occur in the Western Hemisphere, with primarily trop- ical or subtropical distributions (Mathis and Wirth 1978). I have recognized two species groups within Paracanace. This arrangement ad- heres to the cladogram for the species of this genus that Mathis and Wirth (1978:535) published. The two species groups are char- acterized in the key that follows. All known Caribbean species belong to the hoguei group (Figs. 30-38). Key to Species Groups of Paracanace 1. Fore femur of male with from three to four long setae along posteroven- tral surface, setal length subequal to 2x femoral width; mid femur of male bearing a posteroventral, comblike row of setae along entire length, setae at proximate one-fourth pale; costal vein between humeral crossvein and subcostal break usu- ally bearing a row of long spinelike setae, setal length subequal or great- er than width of Ist costal cell oy ae The hoguei Group — Fore femur of male lacking three or four setae as described above; mid femur of male bearing a postero- ventral comblike row of setae along distal one-half only; setae along an- terior margin of wing much shorter, not more than one-half width of Ist costal cell The maritima Group Key to Species of the hoguei Group 1. Three, dorsally curved, genal setae subequal in length; body strongly se- tose (Figs. 30-38) (Cocos Island. Costa Rica) cy ae eee ee ae P. hoguei Mathis & Wirth — Middle, dorsally curved, genal seta about one-half length of setae on either side; body moderately setose 2. Surstylus broader on distal half, es- pecially evident in lateral view; ven- tral, surstylar margin broadly trun- cate in lateral and posterior views; posterior margin of surstylus bear- ing distinct row of longer setae (Florida, Jamaica) 48) OF er ee. . He Bs a P. lebam Mathis & Wirth — Surstylus in lateral view swollen along anterior margin near middle, tapered ventrally to broadly round- ed, ventral margin; posterior margin of surstylus lacking distinct row of longer setae; posteroventral angle of surstylus noticeably produced api- cally (Figs. 39, 40) (Belize, Cuba, Dominica) .. P. aicen Mathis & Wirth Paracanace aicen Mathis and Wirth Figs. 39-42 Paracanace aicen Mathis & Wirth, 1978: 533. Specimens examined.—Belize. Stann Creek District: Carrie Bow Cay, 15 Jan—2 Jun 1984-1987, W. N. Mathis, C. Feller (19 6, 25 2: USNM); South Water Cay, 1 Jun 1985, W. N. Mathis (2 6, 6 2; USNM). Distribution.— United States (Florida: Lee Co., Sanibel Island), Belize, West Indies (Cuba, Dominica, and St. Vincent). This the first record of Paracanace from the United States, although earlier, Mathis (1988:330) had included this genus in a key to the beach-fly genera from the United States with the expectation that the genus was likely to be found there. The new lo- calities from the western Caribbean sub- stantially increase the known distribution of this species, which previously was known only from Dominica in the Lesser Antilles. 602 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 4] Figs. 39-42. External male terminalia of Paracanace aicen: 39, Epandrium and surstylus (holotype from Dominica), lateral view: 40, Epandrium and surstyli (holotype from Dominica), posterior view; 41, Epandrium and surstylus (Florida, Lee Co., Sanibel Island), lateral view; 42, Epandrium and surstyli (Florida, Lee Co., Sanibel Island), posterior view. VOLUME 102, NUMBER 3 The species may eventually be found to be circumcaribbean, but the paucity or absence of canacid collections from Colombia, Ven- ezuela, and most countries of Central Amer- ica precludes assessment of this possibility. Natural history.—The specimens from Florida were collected on the causeway be- tween the Gulf Coast of Florida and Sanibel Island (Lee Co.). The sides of the causeway, particularly the south side, were partially lined with broken pieces of concrete and large rocks to moderate the erosive action of waves. Much of the surface of the con- crete and rocks was covered with algae. In addition to this species, beach flies of the following two species were also found in this habitat: Procanace dianneae (very abun- dant; see treatment below) and Canacea macateei (uncommon; see treatment above). Although this species and Nocticanace texensis both occur on Carrie Bow Cay and are found in essentially the same habitats, I did not collect the two species together during a particular season. In the Lesser An- tilles (Dominica and St. Vincent), I found the two species in the same sweep of the net, although in very unequal numbers. Where one species was common, the other was not. Whether this is an artifact of sampling or is indicative of temporal partitioning needs further investigation. Indeed, to what de- gree these two species may compete for the same or similar resources merits closer scru- tiny. See comments under the treatment of N. texensis for further information concerning Carrie Bow Cay and the specific habitat where this species occurs. Remarks.—This species is closely related and similar to congeners of the Hoguei group but can be distinguished from the latter by the following combination of characters: middle anaclinate genal seta about one-half length of seta on either side; spinelike setae along costal margin variable, short, not as long as width of Ist costal cell, or long (the holotype), length equal or slightly greater than width of Ist costal cell; general ap- 603 pearance setose but less so than in P. hoguei; surstylus of male terminalia as illustrated (Figs. 39-42). After study of several males from the new material collected in Belize, Cuba, Dominica, and Florida, I have noted some variation, although slight, in the shape of the surstylus. The range in variation is as illustrated (Figs. 39-42); the illustrations also facilitate identification of this species. Based on the sampling available, the shape of the ventral surstylar margin in the male from Florida, both lateral and posterior views (Figs. 41, 42), is more characteristic of the species than that of the holotype, which is from Dominica (Figs. 39, 40). Paracanace lebam Mathis & Wirth Figs. 43-44 Paracanace lebam Mathis & Wirth, 1978: 530. Distribution.—Jamaica. Point Hender- son. Remarks.— Externally, this species and P. aicen are very similar, but it may be distin- guished from the latter by the following characters from the male terminalia: sur- stylus (Figs. 41, 42) broader on distal half, especially evident in lateral view; ventral margin of surstylus broadly truncate in lat- eral and posterior views; posterior margin of surstylus bearing distinct row of longer setae. Genus Procanace Hendel Procanace Hendel, 1913:93. Type species: Procanace grisescens Hendel, by original designation.— Mathis, 1988:329-333 [first record of genus from Western Hemi- sphere]. Diagnosis.—General coloration whitish gray, olivaceous, to blackish brown. Head: Intrafrontal setae absent, but with a few setulae inserted anteriorly; fronto-or- bital setae three; ocelli arranged to form equilateral or isosceles triangle, if isosceles, 604 / pele \ NS a dy lle 43 \ Figs. 43-44. 44, Epandrium and surstyli, posterior view. the greater distance is between posterior ocelli. Arista pubescent over entire length. Two large anaclinate genal setae; anterocli- nate genal seta moderately well developed. Palpus not bearing long setae. Epistomal margin, in lateral view, more or less hori- zontal. Thorax: Acrostichal setae, especially a prescutellar pair of large setae, usually lack- ing (setulae present in species of the wil- liamsi group); scutellar disc lacking setae (one or two pairs of scutellar disc setulae occur in P. nakazatoi Miyagi of the william- si group); two pairs of marginal scutellar setae, apical pair not anaclinate; anterior and posterior notopleural setae present, length of both subequal, anepisternum with scattered setulae. Katepisternal setae usually present (lacking in species of the grisescens PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON External male terminalia of Paracanace lebam: 43, Epandrium and surstylus, lateral view; group). Hind tibia lacking spine-like setae apically. Abdomen: Male genitalia as follows: Epandrium in posterior view wider than high; cerci reduced, poorly sclerotized; sur- stylus with an anterior and posterior lobe, the latter larger, sometimes markedly so and shape unique to species. Discussion. — Mathis (1988) first reported the occurrence of Procanace in the Western Hemisphere from specimens collected in Virginia along the tidal shores of the Po- tomac River. The species, P. dianneae, was then only known from Virginia, but recent collecting in North and South Carolina, as well as the Gulf Coast of Florida has re- sulted in discovery of this species along much of the eastern coast of the United States. VOLUME 102, NUMBER 3 605 Figs. 45-53. Scanning electron micrographs of Procanace dianneae: 45, Head, lateral view; 46, Gena and setae, lateral view; 47, Antenna, lateral view; 48, Mesonotum, dorsal view; 49, Frons and ocellar triangle, dorsal view; 50, Fronto-orbital and vertical setae, dorsal view; 51, Scutellum, dorsal view; 52, Notopleuron and setae, lateral view; 53, Katepisternum and setae, lateral view. Annotated Key to Species Groups of Palaearctic, Oceanian, Mala- Procanace Hendel gasy, Seychelles] 1. Katepisternal seta absent ........ — Katepisternal seta present ....... 2 RES inti ced hom Ls dthys the grisescens group 2. Clypeus high, width about twice the [four species; Oriental, eastern height; palpus blackish brown; 606 54 a7 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Figs. 54-56. External male terminalia of Procanace dianneae: 54, Epandrium and surstylus, lateral view; 55, Surstyli, posterior view; 56, Fourth and fifth sterna, ventral view. proepisternal seta absent be See net rad nates the nigroviridis group [seven species; Hawaiian Is- lands] — Clypeus low, width at least four times the height; palpus yellowish; proepisternal seta(e) present ..... 3 3. Acrostichal setulae present, in two irregular rows .... the williamsi group [four species; Hawaiian and Ryukyu Islands] — Acrostichal setulae absent 4. Postocellar setae either absent or much reduced y.. 3s the fulva group [nine species; Oriental and east- ern Palaearctic] — Postocellar setae present, subequal to length of ocellar seta .......... ithe WSF lap Dae aie the cressoni group [three species; Oriental, Nearc- tic] Procanace dianneae Mathis Figs. 45, 46 Procanace dianneae Mathis, 1988:330-333. Distribution.— Eastern coast of the United States from Virginia south through North and South Carolina to Florida, including the Gulf Coast. Natural history.— The type series was tak- en along the tidal shore of the Potomac Riv- er where the water is only slightly brackish. The additional distribution sites reported here are oceanic (Atlantic and Gulf of Mex- ico), with distinctly saline water. On Kure Beach (North Carolina, Brunswick Co.), the specimens were extremely abundant on the sandy shore within a protected area where boats could be launched. The exposed sand was largely covered with algae that had washed ashore. At Cherry Grove (South Carolina, Horry Co.), the specimens were found exclusively on the concrete founda- tions ofa fishing pier. The foundations near- est the shore are exposed at low tide and most high tides and were partially covered with algae. A species of Tethina (probably albula (Loew); Tethinidae) and Fucellia (Anthomyiidae) also occurred on the foun- dations. The causeway leading to Sanibel Island (Gulf Coast side of Florida, Lee Co.) had portions of the shore that were lined with large chunks of concrete and rock to brake the erosive action of waves, and the specimens P. dianneae mostly occurred VOLUME 102, NUMBER 3 where the wave action was dampened by the rocks and concrete that were also ex- tensively covered by algae. Two other beach flies, Canacea macateei and Paracanace aicen, also occurred on the causeway. Remarks.— Externally this species is very similar to those of the cressoni group. It differs from the two species of that group, P. cressoni Wirth and P. taiwanensis Del- finado, as well as other congeners by the following combination of characters: Post- ocellar setae well developed, subequal in length to ocellar setae; clypeus low, height one-fourth width; palpus yellowish. Scutum mostly bluish black, sparsely microtomen- tose, scutum densely microtomentose, brown; proepisternal seta present, pale; kat- episternal seta present; acrostichal setae ab- sent. Shape of the male genitalia unique (see figs. and description above). As noted previously, this species was only recently discovered in Virginia, and it has now been found to occur along the East Coast of the United States from Virginia to Flor- ida, including the Gulf Coast. Apparently the species either dispersed rapidly along the East Coast after its introduction, per- haps similar to Brachydeutera longipes Hendel (Mathis and Steiner 1986; Ephyd- ridae), or it has resided here for some time without being detected. Acknowledgments I am grateful for the assistance in the field from Jorge L. Fontenla (IZAC) and the oth- er collectors as noted in the text. For the loan of the holotype of Canaceoides tex- ensis, | thank Paul H. Arnaud, Jr. (CAS). For critically reviewing a draft of this paper, I thank Amnon Freidberg, Oliver S. Flint, Jr., and I. Candida Feller. The pen and ink illustrations were skillfully inked by George L. Venable, and Susann Braden provided technical support for the scanning electron micrographs. Funding for this research project was provided in part by the Amer- ican Philosophical Society (grant number 9284, Penrose Fund) and a grant from the 607 Research Opportunity Fund (Smithsonian Institution). Their support is gratefully ac- knowledged. This is contribution number 257, Carib- bean Coral Reef Ecosystems (CCRE), Smithsonian Institution, partly supported by a grant from the Exxon Corporation. Literature Cited Cresson, E. T., Jr. 1924. Descriptions of new genera and species of the dipterous family Ephydridae. Paper VI.—Entomological News 35(5):159-164. 1936. Descriptions and notes on genera and species of the dipterous family Ephydridae. IT. — Transactions of the American Entomological Society 62:257-270. Curran, C.H. 1934. The families and genera of North American Diptera. Privately published, New York, 512 pp., 235 figs., 2 pls. Hendel, F. 1913. Acalyptrate Musciden (Dipt.). II.— Supplementa Entomologica 2:77-112, 7 figs. Johnson, C. W. 1910. Order Diptera. Jn J. B. Smith, ed., The insects of New Jersey. New Jersey State Museum Annual Report 1909:703-814. 1925. 15. List of the Diptera or two-winged flies. Jn Fauna of New England. Occasional Pa- pers of the Boston Society of Natural History 7: 1-326, 1 fig. Malloch, J.R. 1924. A new species of Canacea from the United States (Diptera: Ephydridae).—Pro- ceedings of the Entomological Society of Wash- ington 26(3):52-53. 1933. Some Acalyptrate Diptera from the Marquesas Islands.— Bulletin of the Bernice P. Bishop Museum 114:3-31, 9 figs. Mathis, W. N. 1982. Studies of Canacidae (Diptera), I: Suprageneric revision of the family, with re- visions of new tribe Dynomiellini and new ge- nus Jsocanace.—Smithsonian Contributions to Zoology 347:1-29. 1988. The first record of Procanace Hendel from North America, with the description of a new species. — Proceedings of the Entomological Society of Washington 90(3):329-333. , & W.E. Steiner. 1986. An adventive species of Brachydeutera Loew in North America (Dip- tera: Ephydridae).—Journal of the New York Entomological Society 94(1):56-61. ——., & W. W. Wirth. 1978. A new genus near Canaceoides Cresson, three new species and notes on their classification (Diptera: Canaci- dae).— Proceedings of the Entomological Soci- ety of Washington 80(4):524—537, 12 figs. Ritzler, K., & J. D. Ferraris. 1982. Terrestrial en- vironment and climate, Carrie Bow Cay. Pp. 608 77-91 in K. Riitzler & I. G. MacIntyre, eds., The Atlantic barrier reef ecosystem at Carrie Bow Cay.—Smithsonian Contributions to the Marine Sciences 12:1-539. Stoddart, D., F. R. Fosberg, & D. L. Spellman. 1982. Cays of the barrier reef and lagoon.— Atoll Re- search Bulletin 256:1076. Teskey, H. J., & I. Valiela. 1977. The mature larva and puparium of Canace macateei (Diptera: Canaceidae).— The Canadian Entomologist 109: 345-347, 7 figs. Tomlinson, P. B. 1986. The botany of mangroves. Cambridge Tropical Biology Series, Cambridge University Press, Cambridge, 413 pp. Wheeler, M. R. 1952. The dipterous family Cana- ceidae in the United States.—Entomological News 63(4):89-94. Wirth, W. W. 1951. A revision of the dipterous fam- ily Canaceidae.— Occasional Papers of Bernice P. Bishop Museum 29(14):245-275, 6 figs. 1954. A new intertidal fly from California, with notes on the genus Nocticanace Malloch (Diptera: Canaceidae).—The Pan-Pacific Ento- mologist 30(1):59-62. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON . 1965. Family Canaceidae. Pp. 733-734 in A. Stone et al., eds. A catalog of the Diptera of America north of Mexico. United States De- partment of Agriculture, Agriculture Handbook 276. Washington, D.C. . 1970. The American beach flies of the Cana- ce snodgrassii group.— Proceedings of the En- tomological Society of Washington 72(3):397- 403, 4 figs. 1975. 76. Family Canaceidae. Pp. 1-5 in N. Papavero, ed., A catalogue of the Diptera of the Americas south of the United States. Museu de Sao Paulo, Universidade de Sao Paulo, Brasil. 1987. Camnacidae [chapter 102]. Pp. 1079- 1083 in J. F. McAlpine, ed., Manual of Nearctic Diptera. Volume 2. Monograph 28, Research Branch, Agriculture Canada. Hull, Quebec. Department of Entomology, NHB 169, Smithsonian Institution, Washington, D.C. 20560. PROC. BIOL. SOC. WASH. 102(3), 1989, pp. 609-612 A SMALL COLLECTION OF HETEROPTERA FROM THE GALAPAGOS ISLANDS, WITH THE DESCRIPTION OF THE NEW SPECIES NIESTHREA ASHLOCKI AND A LIST OF NIESTHREA SPECIES (RHOPALIDAE) Richard C. Froeschner Abstract.—Report on seven species of Heteroptera from the Galapagos Is- lands: Five confirm reported occurrences on Santa Cruz Island; one new island record of Harmostes disjunctus Barber from Fernandina Island; and description of one new species, Niesthrea ashlocki taken from Sida acuta Burmann [Mal- vaceae] on Santa Cruz Island, most closely allied to the Brazilian N. digna Chopra. Description of the new species accompanied by dorsal habitus drawing and sketch of male genital capsule. Included is a checklist of the species of Niesthrea Spinola. Subsequent to the appearance of my syn- opsis of the Heteroptera of the Galapagos Islands (1985) Dr. Peter D. Ashlock (Uni- versity of Kansas, Lawrence) submitted for my study a small collection of Heteroptera he made on the Galapagos Islands during the period of January to May of 1964. All specimens but one were from Santa Cruz Island and included the following: Beryti- dae: Metacanthus galapagoensis (Barber) [in abandoned garden]; Coreidae: Anasa ob- scura Dallas [from ‘“‘“Mamortlca” (probably a misspelling for the Cucurbitaceae genus Momordica) indica; in abandoned garden]; Miridae: Horcias lacteiclavus Distant [part of the population discussed by Carvalho (1968:200); Pentatomidae: Acrosternum viridans (Stal) [at light; in abandoned gar- den]; Podisus sordidus (Stal) [from Psidium sp.]; Rhopalidae: Harmostes disjunctus Bar- ber, including nymphs; Niesthrea ashlocki, new species described below [from Sida acuta Burmann]. One new island record was included based on a broken specimen of Harmostes disjunctus taken in the Miconia Belt at 1300-2100’ on the SW side of Fer- nandina Island, 4 Feb 1964, P. D. Ashlock. The New World genus Niesthrea Spinola (1837:245), in the tribe Niesthrini, was not previously reported for the Galapagos Is- lands. In my (1985) synopsis it would key to Liorhyssus in couplet 2 on page 49. Niesthrea’s pronotum lacks the subapical, calloused, impunctate, transverse ridge that is characteristic of Liorhyssus. Chopra (1973) presented a revision of the genus Niesthrea based principally on male geni- talic characters. Niesthrea ashlocki, new species Figs. 1-2 Diagnosis.— Males are easily recognized to species by the shape of the medioventral lobe on the genital capsule plus the length of the claspers (Fig. 2): Medioventral lobe broad, lateral expansion subangularly con- vex; exposed part of clasper gently curved, apically more strongly incurved to a sub- acute tip, exposed part of clasper elongate, extending by half its length beyond apex of medioventral lobe. Description (measurements in millime- ters).—Holotype male, length 5.35; general 610 2 Figs. 1-2. Niesthrea ashlocki, new species: 1, dorsal view; natural length 5.3 mm; 2, male genital capsule, ventral view, left clasper omitted. color yellowish; head clouded with fuscous on midline between eyes, with a pair of di- verging, deep-black lines between ocelli. Antenna yellow, segment I mesally and lat- erally with an oblique, fuscous line; seg- ments II and III with a blackened line ex- tending almost full length, II blackened apically. Pronotum with median fuscous area divided by pale median carina; pos- terior lobe with humeri, subbasal margin, and diagonal row of a few fuscous spots. Scutellum with subbasal pair of brown spots. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON or S f cA Vi S i i V3) Y : Hemelytral veins with a few reddish-brown dots. Dorsal disc of abdomen (viewed through hyaline hemelytra) mostly black; connexival segments apically pale, visible segments III-—V basally broadly black, each with an included pale dot, the black ex- tending onto margin of venter. Legs yellow, appearing annulate due to transverse dark marking extending more or less around fe- mur and tibia. Head.—Length 0.90, width across eyes 1.15; preocular part convex, tylus distinctly produced anteriorly; antennal tubercles short, apex transverse. Antennal, segment lengths I-IV, 0.36:0.87:0.87:0.95, segment I reaching apex of tylus. Labium reaching basal segment of abdomen, lengths of seg- ments J-IV, 0.53:0.71:0.53:0.76, segment I reaching hind margin of eye. Pronotum, length 0.90, width 1.77. Genital capsule (Fig. 2) with medioven- tral lobe broader than long, constricted ba- sally, apical margin subangularly concave, laterally subangularly convex; dorsolateral lobe (best viewed from above) distinctly in- curved, markedly surpassing apex of medio- ventral lobe. Exposed part of clasper gently curved, apex projecting mesally as an acute tip, inner margin near apex of capsule with a small tooth. Female.—Length 5.58. General appear- ance similar to male but more abundantly dotted on pronotum; ventrally with red dots on thoracic pleurae. Last abdominal tergum rounded. Last abdominal sternum not notched. Head, length 1.15, width 1.23. An- tennal segment lengths, I-IV 0.37:1.50:1.50: 1.20. Labial segment lengths, I-IV, 0.57: 0.76:0.62:0.82. Pronotum, length 1.25, width 2.06. Holotype male.—“‘Galapagos, 4 mi. [6.4 km] N Academy Bay, Santa Cruz Is., 21 Feb 1965, P. D. Ashlock, Sida acuta Burm. [Malvaceae].’’ Paratype: one female, Gala- pagos Arch.[ipelago], Santa Cruz Is., 2.4 km N Academy Bay, 25 Feb 1964, P. D. Ash- lock. Holotype and lone paratype in the Na- tional Museum of Natural History. Placement of this new species in the phy- VOLUME 102, NUMBER 3 logeny (based solely on males) proposed by Chopra (1973) in his revision of the genus Niesthrea may be made as follows. Three modifications of the male genital structures (Fig. 2) place it on the same branch of Cho- pra’s (1973:457) “‘Phylogenetic tree’? with N. digna Chopra: genital capsule with dor- solateral lobes incurved apically, medio- ventral lobe broader than long, and clasper apically incurved to form an acute angle. It differs from N. digna in having both the dorsolateral lobes and the clasper greatly surpassing the apex of the medioventral lobe, the claspers by almost half their own length and the dorsolateral lobe extending beyond them. The Galapagos Islands occurrence of N. ashlocki is geographically remote from the Brazilian homeland of N. digna. The species name is a dedication to Dr. Peter D. Ashlock, the collector whose many contributions to heteropterology have soundly advanced that science. Checklist of the Species of Niesthrea Spinola The following list is an expanded version of that given on pages 52-56 of Gollner- Scheiding’s (1983) catalog of the family Rhopalidae. agnes Chopra, 1973:455 Argentina ashlocki, new species Galapagos Islands brevicauda Chopra, 1973:455 Peru dentata Chopra, 1973:454 Brazil digna Chopra, 1973:453 Brazil fenestrata (Signoret), 1859:93 Chile flava Grillo & Alayo, 1978:43 Cuba louisianica Sailer, 1961:297 U.S.A.; Mexico parasidae Grillo & Alayo, Cuba 1978:46 pictipes (Stal), 1859:239 Argentina; Brazil; Paraguay subsp. pictipes (Stal), see species entry subsp. casinii Gdllner- Scheiding, 1984:116 Argentina; Uruguay sidae (Fabricius), 1794:169 Greater and Lesser Antilles; 611 Colombia; Mexico; United States; Venezuela Argentina; Brazil Guatemala; Mexico; United States Greater and Lesser Antilles; Argentina; Brazil; Paraguay; Venezuela similis Chopra, 1973:453 ventralis (Signoret), 1859:89 vincentii (Westwood), 1842:6 and 26 Acknowledgments My appreciation is expressed to Dr. John J. Wurdack, Smithsonian Institution De- partment of Botany, for his help in deci- phering the misspelled host genus for Anasa obscura; to Ms. Silver B. West for help in preparing the manuscript; to Ms. Elsie Froeschner for the drawings, and to Dr. Oliver S. Flint and Mr. Thomas J. Henry for helpful reviews of the manuscript. Literature Cited Carvalho, J. C. M., & W. C. Gagne. 1986. Miridae of the Galapagos Islands (Heteroptera).—Pro- ceedings of the California Academy of Sciences, Series 4 36(7):147-219. Chopra, N. P. 1973. Arevision of the genus Niesthrea Spinola (Rhopalidae: Hemiptera).—Journal of Natural History 7:441-459. Fabricius, J.C. 1794. Ryngota. Jn Entomologia sys- tematica emendata et aucta, secundum classes, ordines, genera, species, adjectus synonymis, lo- cis, observationibus, descriptionibus 4:[I-IV], 1-229. Froeschner, R. C. 1985. Synopsis of the Heteroptera or true bugs of the Galapagos Islands. —Smith- sonian Contributions to Zoology 407:1-84. Gollner-Scheiding, U. 1983. General-Katalog der Familie Rhopalidae (Heteroptera). — Mitteilun- gen aus dem Zoologischen Museum in Berlin 59:37-189. 1984. Erganzungen zu den Gattungen Lio- rhyssus Stal, 1870, Niesthrea Spinola, 1837, und Rhopalus Schilling, 1827 (Heteroptera, Rho- palidae).— Mitteilungen aus dem Zoologischen Museum in Berlin 60:115-121. 612 Grillo R., H. & P. Alayo. 1978. La Familia Rho- palidae (Heteroptera:Coreidae) en Cuba. Centro Agricola. Facultad de Ciencias Agricoles, Uni- versidad Central de Las Villa, Septiembre-Di- ciembre 1978:41-64. Sailer, R. I. 1961. The identity of Lygaeus sidae Fa- bricius, type species of the genus Niesthrea (He- miptera: Coreidae).— Proceedings of the Ento- mological Society of Washington 63:293-299. Signoret, V. 1859. Monographie du genre Corizus.— Annales de la Societe Entomologique de France. Series 3 7:75-105. Spinola, M. 1837. Essai sur les genres d’insects ap- partenants a l’ordre des Hemipteres, Lin. ou Rhynchotes, Fab., et a la section des Heterop- Note: The honoree of the new species, Dr. Peter D. Ashlock, died 26 January 1989. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON teres, Dufour. Geneva, Yves Graviers. 383 pp., 15 tabs. Stal, C. 1859. Hemiptera: Species novas descripsit. Kongliga Svenska Fregattens Eugenies Resa Omkring Jorden, III (Zoologi, Insekter). Pp. 219- 298, pls. 3-4. Westwood, J.O. 1842. A catalogue of Hemiptera in the collection of the Rev. F. W. Hope, with short Latin descriptions of the new species. 2:1—26. Department of Entomology, NHB Stop 127, National Museum of Natural History, Washington, D.C. 20560 PROC. BIOL. SOC. WASH. 102(3), 1989, pp. 613-619 CANCER JOHNGARTHI, N. SP. AND CANCER PORTERI (BELL) (CRUSTACEA, DECAPODA): COMPARISONS AND HYPOTHESIS Alberto Carvacho Abstract.— Cancer johngarthi, long mistaken for C. porteri, is described. It is known in the eastern Pacific from Isla Guadalupe, Mexico (29°N), south to Panama (7°N), on soft bottoms at depths exceeding 90 m. Differences from C. porteri are discussed with an emphasis on biological aspects: C. johngarthi shows an isometric growth of chelipeds in relation to carapace width, while in C. porteri a strong positive allometry is evident after the puberal molt. These two conditions may reflect different mating systems. The species Cancer porteri, described by Bell (1835) as C. longipes, was transferred to the genus Platycarcinus, synonym of Cancer, by H. Milne-Edwards & Lucas (1844), and given its present name by Rath- bun (1930). Nations (1975) included it in the subgenus Cancer s.s. The holotype was collected in Valparaiso, Chile. Faxon (1895) recorded it from Panama Bay as deep as 523 m (Albatross). These two eastern Pacific localities, 33°S and 7°30’N, respectively, were long considered as the geographic dis- tributional limits of the species. Garth (1957) cited a continuous distri- bution “from Callao, Peru to Valparaiso, Chile, 0-24 fms” and an extralimital record from Panama. Nevertheless, he also includ- ed in the list of examined material one male collected by the Lund University Chile Ex- pedition at Talcahuano (36°41'S), some 450 km south of Valparaiso. This latter record has been confirmed by Retamal & Yanez (1973). Garth (1961) recorded C. porteri from the coast of Sinaloa in the Gulf of California, between 108 and 128 m, and mentioned that the species ““may now be reported as a bi-temperate species that transgresses the tropics by submergence, being found in the Gulf of California, the Bay of Panama in 210 to 286 fathoms, and from Peru to Chile in the Sublittoral.”’ Information gathered during almost 150 years supported the idea that C. porteri was a eurybathic species with a wide geographic distribution. In fact, the case has been used as a paradigm of the peculiar tropical sub- mergence distribution pattern (Ekman 1953, Garth 1961). Careful study of several specimens re- cently collected off Baja California Sur and the reexamination of virtually all specimens identified with Cancer porteri from the Northern Hemisphere, along with several specimens from Chile and Peru, leads to the conclusion that they belong to two different species. The morphological differences, scarcely evident in young specimens, may express divergence in their mating systems. Cancer johngarthi, new species Figs. 1, 3A, 4B Cancer longipes, Faxon, 1895:16; Rathbun, 1930:199 (in part). Cancer porteri Rathbun, 1930:199 (in part); Garth, 1957:50 (in part); 1961:122; Par- ker, 1964:173; Chirichigno, 1970:45 (in part); Retamal & Yanez, 1973:12 (in part); 614 Nations, 1975:43 (in part); 1979:154, 156, 178 (in part); Retamal, 1981:30 (in part). Carapace granulated, widely oval, very convex and moderately areolated, with pro- tuberances on proto- and mesogastric re- gions and on borders of epi- and meso- branchial regions. Both mesobranchial regions swollen and nearly meeting in me- dian line. Frontal region convex. Front pro- jected and furnished with 3 teeth, median slightly longer and narrower than others. In- ner orbital tooth pointed and slightly short- er than frontal teeth. Anterolateral margin finely granulated, cut into 9 teeth; granu- lations enhanced posteriorly. Posterolateral margin granulated and furnished with 2 teeth, first small and second vestigial, some- times imperceptible. Pterygostomial region swollen and coarsely granulated towards outer edge. Whole carapace remarkably thin; epi- and subbranchial and pterygostomial regions may be easily flexed. Buccal cavity well delimited anteriorly by projections of pterygostomial border, with 2 strong vaults separated by a longitudinal keel. Maxillipeds granulated, with ischium and merus widened distally. Merus with outer face concave and a notch on distal half of inner margin where palp inserts. Chelipeds: fingers with tips and cutting edges dark, starting from proximal tooth. Palm granulated, with 4 longitudinal cari- nae on lower half of outer face. Propodus 2.7 times as long as wide in adult males. Carpus rough, with irregular granulated ca- rinae and anterosuperior pyramidal tooth. Merus with subtriangular section and upper distal margin granulated. Walking legs long and slender, without spines or setae on proximal articles. Prop- odus with scarce setae on distal end of lower margin. Dactylus with 4 symmetrical lon- gitudinal rows of setae and a deep groove along inner and outer faces, respectively. Abdomen in adult males with terminal segment narrowly rounded distally, lateral margins slightly concave and 1.1 times as PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON long as wide. Adult males with second pleo- pod slightly overreaching second segment of thorax; first pleopod slightly overreach- ing third segment of thorax. Holotype.—In the collection of the Allan Hancock Foundation: male (carapace 14 cm wide, 8.7 cm long); Isla Guadalupe, Mexico; 183 m; 15 Nov 1968, Velero IV sta. 12460- 68; catalog number AHF 6815. Material examined.—Paratypes: Isla Guadalupe, Mexico (29°N); 183 m, 15 Nov 1968, Velero IV sta. 12460-68; 4 males, 1 female, AHF 6816. Off Rio San Lorenzo, Sinaloa, Mexico (24°15'N), 108-128 m; May 1959; 2 males, 6 females; AHF 5929. Off Bahia Magdalena, B.C., Mexico (24°15'N), 90-125 m; Jul 1987, 1 male, 3 females, CIB, La Paz. Bahia de Panama; 384 m; Mar 1891; Albatross sta. 3389; 1 female, MCZ, Har- vard University. Distribution.—Eastern Pacific from Isla Guadalupe, Mexico to Bahia de Panama. Southern Gulf of California. Habitat.—Soft bottoms, 90-523 m. Etymology.— Named in honor of Dr. John S. Garth, Chief Curator Emeritus, Allan Hancock Foundation, University of South- ern California, Los Angeles, California. Comparison with Cancer porteri The observations listed below and also data for Figs. 3 and 4 resulted from the ex- amination of 17 specimens of C. johngarthi (carapace widths from 35 to 140 mm) and 43 specimens of C. porteri (c.w. 22.8 to 123 mm). These latter came from Valparaiso, Chile and from the following localities in Peru: Bahia Independencia, Bahia San Juan, Isla San Lorenzo, Bahia San Nicolas, Ca- llao, Isla Lobos de Afuera. 1. Chelipeds of adult males noticeably stronger in C. porteri (Fig. 2a). As shown in Fig. 3B this allometric character is better expressed after the molt of puberty. 2. Darkening in cutting edges of cheliped fingers starts proximally in C. porteri but in C. johngarthi it starts at first tooth. 3. The most remarkable difference at any VOLUME 102, NUMBER 3 615 = negra pa i ; pert haere i \ sre iyfig. LCR ose SOOT L ¢ “is GH Fig. 1. Cancer johngarthi, male: a, Carapace, dorsal; b, Abdomen; c, Left cheliped; d, Second pleopod, distal end; e, First and second pleopods; f, First pleopod, distal end; g, Third maxilliped. 616 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. Cancer porteri, male: a, Left cheliped; b, First pleopod, distal end; c, Abdomen. age is consistency of the carapace. C. john- garthi seems to be a paper shell species. 4. Terminal segment in abdomen of males with straight lateral margins in C. porteri and slightly concave margins in C. john- garthi (Figs. 1b and 2c). 5. Pereopods proportionately longer and dactyli of walking legs longer in relation to propodi in C. johngarthi. 6. Width/length relation of carapace sig- nificantly greater in C. johngarthi (Fig. 4). 7. As shown in Figs. 1f and 2b, apex of first male pleopods quite different in each species. Habitat preferences of each species are not sufficiently documented, but C. john- garthi seems to prefer deeper waters; it has not been collected shallower than 90 m while C. porteri inhabits waters from the intertidal to more than 350 m (Yanez 1974). This scheme agrees with the well known relation of Cancer and water temperature: species in this genus usually live at latitudes greater than those where the surface isotherm of 20°C is to be found (MacKay 1943, Nations 1979). At lower latitudes depth compen- sates thermal needs; such is the case of C. borealis and C. irroratus in Florida, and es- pecially C. guezei in Madagascar (Crosnier 1976). The Hypothesis The genus Cancer originated in the north- eastern Pacific (Ekman 1953, Nations 1975, 1979) and dispersed southward along the west coast of the Americas. Four species may be found in Peru and Chile; one of these is C. porteri. This species, now sepa- rated from C. johngarthi, ranges from Isla Lobos de Afuera (6°57’S) to Talcahuano (36°41’S), covering most of the Peruvian- Chilean province (sensu Briggs 1974). The morphological afiinity and geographic dis- tribution of the two species suggest the ex- istence of a common ancestor that trav- eled—perhaps in the Miocene—between North and South America (Nations 1979). Morphological divergence between C. johngarthi and C. porteri may have resulted from different mating systems. Orensanz & Gallucci (1988) explain some differences among four sympatric species of Cancer, such as dimorphic development of che- lipeds, according to the models of polygyny established by Emlen & Oring (1977). Mat- ing systems of species with precocious de- velopment of a strong cheliped may be in- terpreted as a case of resource defense polygyny: in C. oregonensis each male holds a refuge area—limited resource—which al- VOLUME 102, NUMBER 3 Cheliped height (cm) Carapace width (cm) Cheliped height (cm) 2 4 6 8 10 - 12 14 Carapace width (cm) Fig. 3. Cheliped height plotted against carapace width in males: A, Cancer johngarthi; B, Cancer por- teri. lows him to monopolize females. Preco- cious development of strong chelipeds is re- quired for an early appropriation of adequate refuges. The case of harem defense polygyny involves direct access to females; therefore, defense strategies are only needed once re- productive size is reached. Positive allo- metric growth of chelipeds, consequently, starts just after the molt of puberty. This seems to be the case for C. porteri, as in- dicated by allometric growth of male che- lipeds (Fig. 3B) and also by data in Antezana 617 y L57 l Fig. 4. Frequencies of width to length ratios in car- apaces: A, C. porteri; B, C. johngarthi. L; 54 1.56 Uh UY, 1.5I 1.53 SS 1.45 1.47 1.48 L50 160 162 W/L RATIO et al. (1965) who determined a figure of 5 females per each male after one year of monthly sampling. In the third model, male dominance polygyny, mates or critical resources are not economically monopolizable. Males aggre- gate during the breeding season and females select males from these aggregations. Sexual dimorphism in the development of che- lipeds is not expected here. Orensanz & Gal- lucci (1988) included C. magister in this category, stressing the fact that C. magister is the only species in the genus Cancer in which chelipeds are of the same size in males and females. I suggest that this is also the case for C. johngarthi, in absence of di- morphic development of chelipeds. Several additional arguments uphold this hypoth- esis. As expected, sex ratio is almost 1:1 and sexual selection nearly null; from a total of 618 27 known specimens of C. johngarthi, 15 are males and the rest females. Since breed- ing assemblages are not permanent, breed- ing season should be normally restricted to a short period of time; this seems a reason- able explanation for the lack of ovigerous females in the collected material. On the other hand, there is a relation be- tween size of cheliped and quality of sub- strate. Species inhabiting soft and homog- enous bottoms of fine sand have chelipeds proportionately weaker than those from ir- regular rocky substrates. Lawton & Elner (1985) stated that these differences account basically for the type of feeding, but they left aside an important element of analysis, the role of chelipeds in sexual selection, which is common to most decapod Crus- tacea. Evidently, differences in size and shape of chelipeds of dimorphic species are not due to differences in diet of each sex; male chelipeds fulfill other functions such as the defense of a territory or of a harem. Sudden development of cheliped dimor- phism at molt of puberty in many species is strong evidence of cheliped morphology depending primarily upon sex require- ments. Open soft-bottom environments, compared to complex rocky substrates, al- low fewer possibilities of delimitation and defense of a territory; therefore, different mating systems are involved. Following this idea, lack of dimorphism in chelipeds of C. Johngarthi may be interpreted as a conse- quence of a male dominance polygyny mat- ing system that in turn results from inhab- iting open soft bottoms. Acknowledgments This paper has been enriched with critical comments by John Garth, Janet Haig, Ray- mond Manning, José Orensanz and Rubén Rios. I also thank Dr. Mario Monteforte (Centro de Investigaciones Bioldgicas, La Paz, Baja California Sur) and Ardis B. John- ston (Department of Invertebrates at the Museum of Comparative Zoology, Harvard PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON University), who kindly loaned specimens. Illustrations were prepared by Clara Yanez and Carlos Sepulveda. Literature Cited Antezana, T., E. Fagetti, & M. T. Lopez. 1965. Ob- servaciones bioecologicas en Decapodos co- munes en Valparaiso. — Revista de Biologia Ma- rina (Chile) 12:1-60. Bell, T. 1835. Observations on the genus Cancer of Dr. Leach (Platycarcinus Latreille) with descrip- tions of three new species. — Proceedings of the Zoological Society, London 3:86-88. Briggs, J.C. 1974. Marine zoogeography. New York, McGraw-Hill, 475 pp. Chirichigno, N. F. 1970. Lista de Crustaceos del Peru (Decapoda, Stomatopoda) con datos sobre su distribucion geografica.—Informe del Instituto del Mar, Pert 35:1-115. Crosnier, A. 1976. Donnés sur les Crustacés Déca- podes capturés par M. Paul Gueze a Vile de la Réunion lors d’essais de péche en eau pro- fonde.— Travaux et Documents, ORSTOM 47: 225-256. Ekman, S. 1953. Zoogeography of the sea. London, Sidgwick & Jackson, 417 pp. Emlen, S. T., & L. W. Oring. 1977. Ecology, sexual selection and the evolution of mating systems. — Science 197:215-223. Faxon, W. 1895. The stalk-eyed Crustacea: Report on an exploration off the west coast of Mexico, Central and South America, and off the Gala- pagos Islands, in charge of Alexander Agassiz by the U.S. Fish Commission Steamer ALBA- TROSS during 1891. XV.—Memoirs of the Museum of Comparative Zoology, Harvard 18: 1-212. Garth, J. S. 1957. Reports of the Lund University Chile Expedition 1948-49, No. 29. The Crus- tacea Decapoda Brachyura of Chile.—Lund University Arsskrifter, (2)53:1-128. 1961. Distribution and affinities of the brachyuran Crustacea.—Systematic Zoology 9: 105-123. Lawton, P., & R. W. Elner. 1985. Feeding in relation to morphometrics within the genus Cancer: Evolutionary and ecological considerations. Pp. 357-379 in B. R. Melteff, ed., Proceedings of the Symposium on Dungeness Crab Biology and Management. — University of Alaska, Alaska Sea Grant Reports. MacKay, D. C. G. 1943. Temperature and world distribution of crabs of the genus Cancer.— Ecology, 24:113-115. Milne-Edwards, H., et H. Lucas. 1844. Crustacés. Pp. VOLUME 102, NUMBER 3 1-37 in A. D’Orbigny, ed., Voyage dans l’Amé- rique Meridionale. Nations, J. D. 1975. The genus Cancer (Crustacea: Brachyura): Systematics, biogeography and fos- sil record.—Scientific Bulletin of the Natural History Museum, Los Angeles County 23:1-104. 1979. The genus Cancer and its distribution in time and space.—Bulletin of the Biological Society of Washington 3:153-187. Orensanz, J. M., & V. F. Gallucci. 1988. A compar- ative study of postlarval life-history schedules in four sympatric Cancer species (Decapoda: Brachyura: Cancridae).— Journal of Crustacean Biology 8:187-—220. Parker, R. H. 1964. Zoogeography and ecology of some macroinvertebrates, particularly mol- lusks, in the Gulf of California and the Conti- nental Slope of Mexico. — Videnskabelige Med- delelser fra Dansk Naturhistorisk Forening 126: 1-178. Rathbun, M. J. 1910. The stalk-eyed Crustacea of Peru and the adjacent coast.— Proceeding of the United States National Museum 38:531-620. 619 . 1930. The cancroid crabs of America.—Bul- letin of the United States National Museum 152: 1-609. Retamal, M.A. 1981. Catalogo ilustrado de los Crus- taceos Decapodos de Chile.—Gayana (Zoolo- gia) 44:7-110. ,& L. A. Yafiez. 1973. Analisis cuali y cuan- titativo de los decapodos de los fondos subli- torales blandos de la Bahia de Concepcion, Chile. —Gayana (Zoologia) 23:1-47. Yanez, E. 1974. Distribucion y abundancia relativa estacional de los recursos disponibles a un arte de arrastre camaronero frente a la costa de Val- paraiso (Invierno y Primavera 1972).—Inves- tigaciones Marinas (Chile) 5:125-138. Centro de Investigacion Cientifica y de Educacion Superior de Ensenada, Apartado Postal 2732, Ensenada, B.C., México. Pres- ent address, Instituto Professional de Osor- no, Casilla 933, Osorno, Chile. PROC. BIOL. SOC. WASH. 102(3), 1989, pp. 620-636 STENORHYNCHUS YANGI, A NEW WESTERN ATLANTIC SPECIES OF ARROW CRAB (CRUSTACEA, BRACHYURA, MAJIDAE) AND A REDESCRIPTION OF S. SETICORNIS (HERBST, 1788) Gary D. Goeke Abstract.—The common arrow crab of the western Atlantic has a complicated taxonomic history as two species have long been confused under the name Stenorhynchus seticornis (Herbst, 1788). Through study of living specimens and preserved material in museums of North and South America, as well as Europe, the genus Stenorhynchus Lamarck, 1818, is reviewed, S. seticornis is restricted, its synonyms discussed, and a previously undescribed species, S. yangl, is recognized. Adults of these two species are described and illustrated, their morphologic variations analyzed, colors differentiated, known geographic and bathymetric ranges recorded, and their larvae compared. The species differ in rostral setation, shape of male first pleopod, and in other characters. More- over, two ecophenotypes related to substrate can be recognized within S. se- ticornis. The common shallow-water arrow crab, Stenorhynchus seticornis (Herbst, 1788), of the western North Atlantic is such an ob- vious component of the marine fauna that it has been described under several different scientific names over the past 200 years. It has become evident through the work of Yang (1967, 1976) on larval development in majids and on taxonomic problems in the genus Stenorhynchus Lamarck, 1818, that two species are confused under the name Stenorhynchus seticornis (Herbst, 1788) (Fig. 1). Historical Review The group of spider crabs currently as- signed the generic name Stenorhynchus has a wide distribution in warm and temperate waters of the Atlantic and eastern Pacific oceans. Species now grouped within this ge- nus were previously assigned to various genera until taxonomic consistency was reached with the use of the generic name Leptopodia Leach, 1814. Arrow crabs are very common and led Milne-Edwards (1875: 173) to state, ““Cette espéce est si bien con- nue, et elle a été si souvent figurée, qu’il est inutile d’en donner ici une description.” Leptopodia is now known to be a junior synonym of Inachus Weber, 1795, and not a valid generic name for this group. Cancer sagittarius Fabricius, 1793 (=Stenorhyn- chus seticornis) was transferred to the genus Leptopodia by Leach in 1815 and was con- sidered the type for the genus. However, Leptopodia was erected for Cancer pha- langium Pennant, 1777, and Leptopodia tenuirostris Leach, 1814. Because Cancer sagittarius was not mentioned in the origi- nal description of Leptopodia, it could not serve as the type species. Lamarck (1818) erected the genus Stenorhynchus for Cancer seticornis Herbst, 1788, and Cancer pha- langium. The latter species, however, is a member of the genus /nachus. Since the des- VOLUME 102, NUMBER 3 621 ir .43 . bd 5 J re - TAN Ly, AUMLMES MAU Fig. 1. Stenorhynchus seticornis, a, b, d, e, f, and g; Stenorhynchus yangi, c. 622 ignation of the name Stenorhynchus by Rathbun (1897) as an available name for the group, it has been widely accepted. Garth & Holthuis (1963) petitioned the Interna- tional Commission on Zoological Nomen- clature (ICZN) to designate Cancer seticor- nis Herbst, 1788, the type species of the genus and to officially emend the spelling of the generic name from Stenorynchus to Stenorhynchus. These recommendations were followed in opinion 763 of the ICZN. Five binomials may bear on western At- lantic species of Stenorhynchus, the earliest of which is the description of the “‘Oost- Indische Zee-Krabbe” by Slabber (1778). This “East Indies Sea Crab’ was described in very general terms which dealt primarily with the gross morphology of the carapace and legs. The specimen, a female from the description of the abdomen, was character- ized as having small setae on the sides of the rostrum. The very generalized illustra- tion accompanying Slabber’s (1778: pl. 18, fig. 2) description includes a single useful morphologic feature for specific taxonomic purposes, and even that is of limited diag- nostic value for taxa within this group. The figure indicates a rostrum twice the length of the postorbital region of the body. How- ever, the drawing of the original figure is questionable because the crab is not accu- rately depicted, as evidenced by the lack of spination on the ambulatory legs. Although no doubt exists as to the genus with which Slabber dealt, the morphological characters considered useful for specific identifications by today’s standards were not detailed by that author. The type locality of the “East Indies Sea Crab” as given by Slabber is incorrect. Hol- thuis (1959:185) noted that the genus Ste- norhynchus is not represented in the Indo- West Pacific and that material from which the description was drawn was apparently mislabeled. Holthuis (1959) restricted the type locality to Guadeloupe because mate- rial examined by Herbst, following Slab- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ber’s description, was collected from Gua- deloupe. The binomial nomenclature introduced by Linnaeus (1758) was used by Herbst (1788:229) to designate the species Cancer seticornis. His abbreviated description was based on the work and material of Slabber and even reproduced the figure used by Slabber (Herbst, 1788: tabl. XVI, fig. 91). As noted by Yang (1967:211), this figure was also reproduced by Bosc (1802). A more detailed and accurate representation of a male Stenorhynchus species was later given by Herbst (1803: tabl. LV, fig. 2) from Guadeloupe material. This latter figure shows the rostrum nearly twice the length of the postorbital carapace. The margins of the rostrum are nearly parallel, and the rel- ative length of the dactyl of the cheliped suggests that the specimen figured repre- sents S. seticornis. However, the very reg- ular placement of spines and absence of postocular spines indicate that the figure is somewhat stylized and not an accurate rep- resentation. The type material for S. seticornis has not been located and must be assumed lost or destroyed. Slabber’s original material was offered for sale to the ““Zeeuwsch Genoot- schap van Wetenschappen”’ (Society of Sci- ences of the Province of Zeeland) in Mid- delburg, the Netherlands. The price asked was not agreeable and the offer was de- clined. The collection later reputedly was sold to the Leiden Museum. No record of this acquisition exists and no material is present which can be attributed to Slabber’s “Sea Crab.’ The type specimens must be assumed lost (L. B. Holthuis, pers. comm.). Because the original type locality is in error and the original description lacks diagnostic features, a neotype is proposed for the re- description of S. seticornis. The specimen on which the redescription is based was col- lected from the Dutch West Indies, Curacao (USNM 42956). The previously designated type locality of Guadeloupe (Holthuis 1959) VOLUME 102, NUMBER 3 623 ee en A IT IIL UA UU DN a UMA AU au a aU a ee ee ee es baa HK KK KK KL KK Soe iene ae ee Poe OO... HB Fig. 2. Cancer sagittarius Fabricius, 1793: Above, ““Type” specimen from the Copenhagen Museum: Below, the specimen originally deposited in the Kiel Museum. 624 is superseded by Curacao with the selection of the male neotype. Curacao is a location from which collectors are known to have sent Slabber material for examination (Hol- thuis, pers. comm.). The second name to be considered is Cancer sagittarius (Fabricius, 1793). The original description of Cancer sagittarius 1s inconclusive. However, from an examina- tion of the syntypes, it has been confirmed the Fabrician material is conspecific with Stenoyrhynchus seticornis. A syntype was originally deposited in the Zoological Mu- seum, Copenhagen, and a second syntype, along with the remainder of the Fabrician material, has been transferred from the Kiel Museum to Copenhagen. The syntype of Cancer sagittarius transferred from the Kiel Museum has deteriorated to fragments that are very nearly unrecognizable. This spec- imen (Fig. 2) was listed by Rathbun (19235: 14) as type material. However, it was not listed by Zimsen (1964) as a syntype. Prof. Torben Wolff (Zoological Museum, Copen- hagen) knows no reason why one specimen was listed as the type by Zimsen and not both (pers. comm.). This female syntype possesses all the characters listed below as typical of S. seticornis. The eastern Atlantic species of Steno- rhynchus was considered conspecific with the western Atlantic species until Yang (1967, 1976) showed it to be distinct. Apart from the unavailable Leptopodia vittata Kingsley, two names have been applied to the eastern Atlantic form, Leptopodia lan- ceolata Brullé, 1837, and L. canariensis Brullé, 1839, both described from the Ca- nary Islands. As Yang pointed out, the cor- rect name for the eastern Atlantic taxon is Stenorhynchus lanceolatus and L. canarien- sis must be considered a junior synonym. Yang (1967) showed how S. lanceolatus dif- fers from both S. seticornis and the pro- posed new species and Manning & Holthuis (1981) presented considerable data on the biology and distribution of S. /anceolatus. Paula (1987) has most recently described PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON the first zoeal stage of S. lanceolatus and compared it with the works of Yang (1967, 1976). The next name possibly available is Lep- topodia ornata Guilding, 1825, described from St. Thomas in the Caribbean Sea. The original description in Latin is very brief. It treats a majid crab with a serrate rostrum, and includes few details as to general shape of the legs or other features. Guilding also corrected the definition of Leptopodia by detailing the arrangement of the abdominal segments. By today’s standards, however, there are no characters described for L. or- nata that are of diagnostic value. Rathbun (1925) listed the type specimen of L. ornata as not in the British Museum and perhaps not extant. Yang (1967) requested Dr. Isa- bella Gordon to make another search for the specimen, but that search was also fruit- less and the type must be assumed lost. Lep- topodia ornata Guilding, 1825, is herein designated a junior synonym of Stenorhyn- chus seticornis (Herbst, 1788). ; Another name in the literature which may have precedence for the proposed new species is Leptopodia vittata. Kingsley (1880) reported the presence of a specimen within the collections of the Museum of the Acad- emy of Natural Sciences of Philadelphia la- beled “‘Leptopodia vittata Guer., Senegal.” He stated that it might represent a manu- script name and that no published descrip- tion could be found of the species. Lepto- podia vittata Kingsley, 1880, is not available because it was published only in synonymy and was not adopted before 1961 as a name for a species. Manning & Holthuis (1981) listed this species as synonymous with the West African S. /anceolatus Brullé, 1837. Goeldi (1886) described Leptopodia lin- eata from Rio de Janeiro and Cabo Frio, Brazil. The description is somewhat vague, not detailed enough to determine which of the 2 western Atlantic species it represented, and the illustrations are not sufficiently de- tailed to accurately identify the taxon by today’s standards. An attempt to locate the VOLUME 102, NUMBER 3 type series of L. /ineata at the major zoo- logical museums of Europe and Brazil was fruitless; the type must be assumed lost. Stenorhynchus material obtained from Cabo Frio and Rio de Janeiro revealed only S. seticornis. A search by Dr. W. Zwink (Mu- seu Nacional, Rio de Janeiro) of holdings of local specimens of Stenorhynchus showed no representatives of a second species. It may be conclusively presumed that this tax- on 1s conspecific with S. seticornis and is herein designated a junior synonym. The only remaining specific name that needs to be mentioned here is Pactolus bos- ci, Leach, 1815. This species was based on a single specimen of unknown origin found in the holdings of the British Museum (Nat- ural History). As the name has been sup- pressed by the ICZN (opinion 763) it is not available for use. Systematics Due to the extremely common nature of Stenorhynchus species and their wide geo- graphic range, it is nearly impossible to de- tail all of the workers who have dealt with western Atlantic members of the genus in the past 200 years. It is not attempted here. In most cases, it is not feasible to determine accurately from the literature which of the two species was the subject of each report. A great many of the citations are species listings and not accompanied by diagnoses, ecological data, or illustrations which would help to determine the identity of the taxon reported. However, a few records are suf- ficiently detailed (e.g., Hay & Shore 1918; Williams 1965, 1984) or have illustrations which help to clear up some of the confu- sion. For these reasons, the synonymies that follow are abbreviated and by no means de- tailed accounts of references to Stenorhyn- chus species in the literature. The materials examined in the following species accounts are housed at the United States National Museum of Natural History (USNM), Florida Department of Natural 625 Resources, Marine Research Laboratory (FSBC), Museu Nacional, Rio de Janeiro, Gulf Coast Research Laboratory (GCRL), Dauphin Island Sea Lab (MESC), and the University of Southwestern Louisiana (USLZ). Stenorhynchus seticornis (Herbst, 1788), redescription Figs. la, b, d—g, 2, and 3 Oost-Indische Zee-Krabbe Slabber, 1778: 162: pli 18; fig. 2. Cancer seticornis Herbst, 1788:229, pl. 16, hie tee 1803-27, pl 55; fig.. 2: Cancer sagittarius Fabricius, 1793:442. Leptopodia ornata Guilding, 1825:335. Leptopodia lineata Goeldi, 1886:37, pl. 3, figs. 24-31. Stenorynchus sagittarius.—Rathbun, 1901: 53.—Hay & Shore, 1918: 455, pl. 37, fig. 8 (in part). Stenorynchus seticornis.—Rathbun, 1925: 14, pls. 2 and 3 (in part).—Abele, 1970: 137 p. Stenorhynchus seticornis.— Williams, 1965: 244, figs. 222 and 223K (in part).—Feld- er, 1973:48, pl. 7, fig. 1.—Yang, 1976.— Felder & Chaney, 1979:27.—Wicksten, 1980:150.— Williams, 1984:304 (in part). Material examined.—Table 1. Diagnosis.—Carapace naked, rostrum covered with short dense felt and setae, be- coming longer and thicker distally. No spines at distal end of basal antennal article, single inter-antennular spine directed posteriorly. Chelipeds hairy, palm from three to four times length of fingers in mature males, not as stout in females. Merus of third maxil- liped normally with small spine on antero- distal angle. Pereopods, abdomen, and ster- num bearing short pubescence. Description of male neotype (USNM 42956).—Carapace subtriangular, smooth, naked, regions slightly defined; intestinal and cardiac region inflated and separated by shallow furrow from posterior regions; branchial region inflated, delimited by shal- 626 Table 1.— Material examined; Stenorhynchus seticornis. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Catalog number Location Material Depth (m) USNM 42956 (Neotype of Stenorhynchus seticornis) Curacao, Suriname 16 = USNM 19941 Rio de Janeiro, Brazil 16 — USNM 76441 Gallows Bay, St. Croix, Virgin Islands 13 = USNM 46697 Aucilla, FL 19 13 USNM 43065 Montego Bay, Jamaica 26 — USNM 43066 Jamaica 1¢ — USNM 19580 Kingston Harbor, Jamaica 1¢ — USNM 31040 Grolding Cay, Bahamas 16 _ USNM 11232 33°34'N, 77°42'W iS. 9 16.5 USNM 19940 Bay of Bahia, Brazil 19 — USNM 22554 Santa Marta, Colombia 1¢ — USNM 69601 18°30'N, 66°04’W Puerto Rico 1¢ 366 USNM 101712 16°05'N, 82°05’'W 16 44 USNM 137761 Grand Island, Trinidad 36 69 USNM 137020 00°15'S, 46°45’W 14,22 28 USNM 171562 Tongue of the Ocean, Bahamas 16 — USNM 184122 Palm Beach, FL 16 22. USNM 58047 Shoal Banks, Barbados 16 55 USNM 154711 Grenada 1¢ — USNM 321390 Belize 1¢ — USNM 103268 06°51'N, 54°53’W Suriname 12 ait USNM 43060 Montego Bay, Jamaica 1¢ — USNM 105004 08°40'N, 77°10’'W Gulf of Darien 19 61 USNM 137020 00°15'S, 46°45’W Para, Brazil | row) Xe: 27, USNM 107804 off Houma, LA 1¢ _ USNM 103267 06°49'N, 55°54’W 12 47 USNM 103504 07°40'N, 57°34’'W 1¢ — USNM 17374 22°18'N, 87°04’W 1¢ 44 USNM 49082 10 miles south of Key West, FL 12 229 USNM 42955 Curacao 12 _ USNM 101713 01°S50’N, 47°93 1'W 19 85 USNM 43062 Montego Bay 14 _ USNM 7653 St. Thomas 1¢ _ USNM 17373 (in part) 35°08'30’N, 75°10’'W 1é 90 USNM 137019 04°46’N, 51°21’W 16 59 USNM 103265 06°48'N, 54°54’W 26 46 MESC 6187-0121 28°26'N, 84°21'W 3: 3.2 ~ MESC 6187-0113 28°25'N, 84°19'W 64,42 — MESC 6187-0114 30°02'30”N, 86°06'30”W 26 ul! MESC 6187-0109 28°36'N, 84°16'W To 52 _ GCRL 164:695 29°43'N, 88°26'W 14,32 — FSBC I 31042 (in part) 30°30'N, 80°15’W 22 47 FSBC I 31045 (in part) 30°20'N, 80°14’'W 13 65 FSBC I 31043 (in part) 30°31'N, 80°10'W Lig 22.9 64 FSBC I 31044 (in part) 30°20'N, 80°17'W 1¢ 46 FSBC I 31032 27°40'N, 80°06'W 1¢ 27 FSBC I 31030 27°10'N, 80°01'W 1¢ 46 FSBC I 31029 27°10'N, 80°01'W 19 45 Museu Nacional Rio de Janeiro Cabo Frio, Brazil 14,12 — Museu Nacional Rio de Janeiro Rio de Janeiro, Brazil 146,12 — Museu Nacional Rio de Janeiro Guanabara, Brazil ivGul-e — VOLUME 102, NUMBER 3 low margin; hepatic region inflated, with well-defined ventral margin, but remaining ventral margin ill defined. Simple, strong deflexed postorbital spine; supra-orbital furrow shallow; rostrum flattened basally between eyes; short setae from base of ros- trum to tip, setae dense and increasingly long distally; subhepatic region inflated pos- teriorly, margins well defined; Ist rostral spine directed laterally, slightly deflexed, followed by up to 16 large spines on lateral margins, directed forward and occasionally downward; rostrum 1.8 times length of car- apace behind transverse line connecting base of eyestalks dorsally. Basal antennal article elongate, with longitudinal ventral furrow, strong spine anterolaterally directed on ven- tral margin; septum dividing antennular si- nuses with posteriorly directed spine; an- terolateral margin of sinus defined by upturned border; lateral and anterior mar- gins of buccal cavity with raised ridge, acute small spine at anterolateral angle of mouth frame. Exopod of third maxilliped with maximum width one-third distance from base, narrowing distally; internal margin of merus straight, with strong spine at antero- mesial angle and small spine on exterior margin posterior to articulation with palp. First pereopods greatly elongate, covered with short dense pile; basis inflated, tuber- culate, with spine on interior margin; merus tuberculate, with region of few tubercles dorsolaterally, six to nine strong spines on mesial row, single spine dorsally, two spines in lateral row, ventral spine toward distal margin, three or four spines at articulation with carpus; carpus with three strong spines dorsally, three distal spines ventrally and numerous tubercles; propodus covered with strong tubercles and low pile which becomes denser and longer at base of finger, pile ex- tends laterally along propodus and onto ventral surface at base of finger, propodus three times length of dactyl; dactyl stout, with long dense setae dorsally and laterally at base, thinning distally to become sparse long setae, tuberculate dorsally. 627 Second pereopod longer than first and covered with short dense pile; merus with 4 longitudinal rows of spines, 5 or 6 spines in dorsal row, 5-7 spines in lateral row con- centrated in distal '2, 11 mesial spines dis- tributed along length of segment, 2 ventral spines in distal 12; 3 spines at articulation with carpus slightly longer than others; car- pus with pair of dorsal spines at midlength and 3 spines at articulation with propodus longer than others; length of carpus and propodus equal to dactyl but shorter than merus; propodus with 1 1-13 spines, slightly compressed laterally, 2 spines at articula- tion ventrolaterally; dactyl elongate, slightly curved, little compressed laterally, with 5 longitudinal rows of setae. Third pereopod shorter than second, cov- ered with short dense pile; merus with 5 dorsal spines along length, 8—10 spines along inner row, 3—4 spines in distal '2 on external row, single spine on ventral margin in distal Y), 3 spines at articulation with carpus; car- pus with pair of dorsolateral spines at mid- length and 3 spines at articulation with propodus; propodus with 11-13 small spines, slightly compressed laterally, with 2 spines at articulation; dactyl % length carpus and propodus combined, with 5 longitudi- nal rows of setae, curved slightly in distal i. Fourth pereopod shorter than third, with covering of short pile; merus with four to five spines on internal row and ventrolateral spine at articulation slightly enlarged; car- pus with pair of dorsal spines at midlength and three spines at articulation with prop- odus; propodus with eight or nine small spines; dactyli damaged, slightly com- pressed laterally, and bearing five longitu- dinal rows of setae. Fifth pereopod shorter than fourth, cov- ered with short dense pile; merus with 4 dorsal spines, single ventrolateral spine and 2—4 mesial spines; carpus with pair of spines dorsolaterally at midlength, 3 terminal spines at articulation; propodus with 9-10 low spines, slightly compressed; dactyl elon- 628 gate, curved distally with 5 rows of longi- tudinal setae. Abdominal segments 5 and 6 fused, with sutures indicated; segment 1 slightly longer than wide, naked along elevated midlength but with setae in depressions; segments 3 and 4 subequal, longer than 2; 3 widest an- teriorly, segments 5 and 6 fused, segment 7 is 1.75 times long as wide with hollowed protuberance for locking mechanism and a small tubercle medially in proximal one- half. Sternum with 22 large tubercles and few low setae; sternite 5 with 2 tubercles at articulation with basis of maxilliped; ster- nite 4 with 8—9 tubercles at articulation with first pereopod; plastron with arcuate ridge in anterior '/ lined with setae, lateral margin parallel. Color.—Carapace with alternating stripes of off-white and brown-maroon, white stripes on dorsum of carapace reminiscent of inverted V’s, large white bands originat- ing dorsally between fourth and fifth legs join anterior to eyes, every other white stripe smaller than preceding; white bands origi- nating at tip of dactyl of pereopods, con- tinuing dorsally along leg onto carapace, or originating at posterior margin of carapace between coxae of legs. Carapace with 4 ma- jor white longitudinal bands, one running along the pterygostomial ridge. Distal *4 of fingers on chelae blue. Inner surface of fin- gers each with 2 whitish spots, forming rough circle when fingers close. Merus of cheliped with large orange-yellow spot at base of dis- tal spine. White band bordered with ma- roon extending obliquely across merus and palp of third maxilliped. Broad yellow-white band connecting coxae of first pereopods with area under third maxilliped. Range.— Material assignable to S. seti- cornis has been examined from Cape Fear, North Carolina, through the Gulf of Mexico southward to the mouth of the Amazon River, and Cabo Frio, Brazil. Bathymetric records are from | to 366 m. Remarks.—The above technical descrip- tion is drawn from the designated neotype, PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON a mature male collected at Curacao, Dutch West Indies. This specimen is the form commonly found on the rock outcroppings or coral reefs in temperate and tropical waters of the western Atlantic, and reported by Herbst (1803) from Guadeloupe. Variation.— Stenorhynchus seticornis ex- hibits considerable morphological variation which appears to be related to the substrata from which the specimens are collected. One variant, herein designated S. seticornis form A, is most often collected on hard rocky bottoms (i.e., rock outcroppings, reefs, jet- ties) or immediately adjacent to these hard- bottom types. It is a large heavy bodied form, having an extremely long rostrum with sub- parallel margins throughout most of the length. This form was mentioned by Yang (1967) as “atypical.” Stenorhynchus seticornis form B differs from the above by characters enumerated below. Form B is the ecotype most often found in the northern Gulf of Mexico on mud bottoms and in grass beds. It was listed by Yang (1967, 1976) as Stenorhynchus se- ticornis and the complete larval develop- ment has been described. It is the smaller of the two forms. In SEM micrographs, the male pleopod of S. seticornis form A (Fig. 3) shows little variation, other than size, from that of S. seticornis form B (Fig. 3). The somewhat narrower opening of the apex and the slightly shorter apex may represent differences in the orientation of the gonopod at the point where the photographs were taken. This very minor difference, and the observed gradation between the two forms, are well explained by ecophenotypic vari- ation. Stenorhynchus seticornis form B is sepa- rated from S. seticornis form A by: 1) small- er overall body size, 2) females and subadult males with large hiatus at base of moveable fingers, and 3) setae on the dactyl and prop- odus of the first leg not forming thick mat continuous on dorsal surface. Considerable intergradation is found be- tween the two forms of S. seticornis, espe- VOLUME 102, NUMBER 3 Fig. 3. Scanning electron micrographs of the tips of male gonopods: Above, Stenorhynchus seticornis form A; Below, S. seticornis form B. 630 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 2.— Material examined; Stenorhynchus yangi. eee eee Catalog number Location Material Depth (m) a USNM 211812 (HOLOTYPE) 33°49'N, 76°43'W ihc 81 USNM 211855 (PARATYPE) 26°45'70"N, 84°00'13"W 18 89 USNM 211803 (PARATYPES) 33°48'46’N, 76°35'W 346,22 60 USNM 67800 18°30'30’N, 66°23'50”"W 16 He USNM 101715 34°17'N, 76°01'W 19 137 USNM 120135 Dominican Republic 1¢ pa USNM 103269 6°52'N, 54°53'W 14,12 288 USNM 73401 Barbados ia = USNM 73086 Grand Cayman 1é 220 USNM Uncatalogued Oregon St. 4391 12°33) NE 09 Ww. rote li i: USNM Uncatalogued Oregon St. 3605 12° T6IN 82°53 W 16 1D USNM 69601 18°30'30’N, 66°04'05”W 16 365 USNM 92646 2T°-1S NSO 2 TOW 16 64 USNM 211815 33°48'42”N, 76°34'12"W 1. dyke 102 USNM 24418 Mayaguez Harbor, Puerto Rico 16 137 USNM 11379 Key West, FL 12 108 USNM 11219 Martha’s Vineyard 16 62 USNM 211807 33°47'36’N, 76°34'24"W 22 116 USNM 9496 Key West jewel) S _ USNM 103504 70°40’N, 57°34'W 12 — USNM 6934 9°30'15”N, 76°20'30”W Sid oe — USNM 9862 33°18'30’N, 77°07'W 16 174 USNM Uncatalogued Oregon St. 4391 12°33'N, 71°09'W [Boe LS 73 USNM Uncatalogued Oregon St. 3605 12°16'N, 82°53'W 16 7h: USNM 17373 (in part) 35°08'30’N, 75°10’'W 22 90 GCRL 160:165 25°35'N, 83°42'’W 5:62 110 FSBC I 31031 27°40'N, 79°59'W ye toe |e: 64 FSBC I 31049 30°00'N, 80°15’W 16 91 FSBC I 31042 (in part) 30°30'N, 80°15'W 12 47 FSBC I 31048 30°11'N, 80°15’W 16 64 FSBC I 31033 27°50'N, 79°58’'W seks 92 FSBC I 31045 (in part) 30°20'N, 80°14’W 104,42 65 FSBC I 31041 30°40'N, 80°06’W PL 91 FSBC I 31036 28°40'N, 80°06’W 192 64 FSBC I 31043 (in part) 30°21'N, 80°10’W leds Ee 64 FSBC I 31046 30°20’N, 80°12’W 16 92 FSBC I 31040 30°40'N, 80°07’W 16 64 FSB@ RSt037 28°50'N, 80°09’W 16 64 FSBC I 31034 28°30'N, 80°01'W 192 91 FSBC I 31047 30°10'N, 80°14’W 12 91 FSBC I 31038 29°00'N, 80°10’W 1¢@ 64 FSBC I 31044 (in part) 30°20'N, 80°17’W 192 46 cially in areas of rock outcroppings adjacent to muddy bottoms. In this situation, spec- imens are larger than those found on muddy bottoms, but often possess the hiatus at the base of the fingers and the greatly elongated rostrum of the form common to reefs. Sim- ilarly, specimens from grass beds adjacent to coral heads exhibit characteristics of both forms A and B. Specimens of S. seticornis exhibit great variation in a number of characters, includ- ing length of the rostrum and the fingers. To document this variation, I measured the total postorbital carapace lengths and total carapace lengths on specimens for which the rostrum was unbroken on specimens which represented the entire geographic range. The postorbital measurement was determined VOLUME 102, NUMBER 3 from the posterior margin of the carapace to a point midway between the base of the eyestalks. Additional measurements of the moveable finger and palm length were tak- en. The range of the ratio of postorbital length to total carapace length is slightly dif- ferent in males (n = 20:0.27-0.40) and fe- males (n = 5:0.33-0.42), but no real di- morphism in carapace dimensions is evident which is attributable to the sex of the spec- imens. These differences bear no relation- ship to either the size of the individual or the location from which it was collected. However, differences in the range of ratios of the finger to palm lengths do seem at- tributable to sexual dimorphism. Although there is some overlap in the range, males (n = 22) tend to have shorter fingers, relative to the palm length (0.24—0.39) than do fe- males (n = 7:0.37—0.45). This trend is also evident in S. yangi. A detailed examination of the variation found in S. seticornis has not produced suf- ficient data to warrant the establishment of separate subspecies. Until the larval devel- opment of S. seticornis form A has been documented, I prefer the designation “‘form”’ to a questionable subspecies status. Stenorhynchus yangi, new species Fig. lc Stenorynchus sagittarius.—Hay & Shore, 1918:455, pl. 37, fig. 8 (in part). Stenorynchus seticornis.—Rathbun, 1925: 14, pls. 2 and 3 (in part). Stenorhynchus seticornis.— Williams, 1965: 244, figs. 222 and 223K (in part).— 1984: 304 (in part). Stenorhynchus sp. A.— Yang, 1967; 459 p.— 1976:158. Material examined.—Table 2. Diagnosis.—Carapace naked, rostrum devoid of setae or felt; no spines at distal end of basal antennal article; interanten- nular septum without posteriorly directed spinous process; chelipeds hairy, palm only twice length of moveable fingers in males, 1.5 to 2 times length of dactyl in females. 631 Merus of third maxilliped with vestigial spine at anteromedial angle. Ambulatory legs, abdomen and sternum without pubes- cence. Regions of carapace well defined and inflated. Description of holotypic male (USNM 211812).—Carapace subtriangular, smooth, naked, regions well defined; branchial re- gions inflated; intestinal and cardiac regions elevated in midline and separated from branchial and cardiac regions by sulcus; gas- tric region inflated; hepatic region moder- ately inflated and defined by furrows on all sides. Strong postorbital tooth bifid, de- flexed and directed slightly forward. Shal- low supraorbital furrow; rostrum flattened between eyestalks, devoid of setae; subhe- patic region inflated posteriorly with well defined margins; first rostral spine directed slightly forward; rostrum about equal to length of postorbital carapace, naked, broad basally and tapering to acute apex, bearing 17 to 19 large lateral spines; basal antennal article with shallow longitudinal furrow and strong anteriorly directed spine on ventral margin; septum dividing antennular sinuses well developed, with rounded ventrum; an- terolateral margin of sinus simple. Lateral and anterior margins of buccal frame with raised margin and weak spine at anterolat- eral angle; exopod of third maxilliped with maximum width at one-half length from base, narrowing distally, internal margin of merus concave, occasionally with small spine at antero-internal angle, small spine posterior to articulation with palp. All pereopods with few scattered setae. First pereopod greatly elongate; ischium slightly inflated, smooth; merus with nu- merous spines in longitudinal rows; two spines in distal one-half of lateral row; three spines in dorsal row evenly spaced, enlarged terminal spine at articulation with carpus; six spines on interior row; carpus with two dorsal spines at midlength and pair of ter- minal spines ventrally; propodus with small scattered tubercles. Felt and dense setae lat- erally at base of finger in small patch with longer sparse setae on immoveable finger; 632 dactyl with small patch of setae at base, with few long scattered setae on surface, small hiatus at base. Second pereopod elongate, longer than first; ischium very short, tuberculate dor- sally; merus greatly elongate, with about 20 spines arranged in 4 longitudinal rows, 3 larger terminal spines; carpus short, about . length of merus, with 3 dorsal and 3 ter- minal spines; propodus elongate, *4 length of merus, with numerous spinules and tu- bercles, | lateral terminal spine; dactyl elon- gate, 44 length of merus, slightly compressed laterally, lined with spinules and with cor- neous tip. Third pereopod shorter than second; is- chium very short, tuberculate; merus elon- gate, with approximately 15 spines arranged primarily in 2 dorsal rows, 3 terminal spines; carpus short, %4 length of merus, 3 dorsal and 3 terminal spines; propodus elongate, *3; length of merus, with approximately 10 dorsal and 2 terminal spines; dactyl elon- gate, ¥, length of merus, with numerous spi- nules, slightly compressed laterally with corneous tip. Fourth pereopod shorter than third, is- chium tuberculate, very short; merus elon- gate, roughly 20 spines arranged in 2 pri- mary dorsal rows, 3 terminal spines; carpus short, 4 length of merus with 3 dorsal and 3 terminal spines; propodus elongate, * length of merus with about 10 spines dor- sally and 3 terminal spines; dactyl elongate, 47, length of merus with numerous spinules and corneous tip, slightly compressed lat- erally. Fifth pereopod shorter than fourth, is- chium very short, slightly tuberculate; me- rus elongate, with 10 spines in 2 dorsal rows, and 3 terminal spines; carpus short, 14 length of merus, with 3 dorsal and 3 terminal spines; propodus elongate, equal in length to merus, with spinules mainly in single dor- sal row, slightly compressed laterally, nu- merous spinules along length; dactyl with corneous tip. First abdominal segment little longer than PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON wide, elevated along midline, segments 2 and 3 very short but broad; segments 4 and 5 somewhat longer and constricting to nar- rowest part of abdomen, segments 6 and 7 fused, no external indication of fusion vis- ible. Tubercles scattered on sternum with very few setae apparent; anterior extension of sternum with ridge below mouth frame bearing setal row and two spines at center of process. | Color.—Carapace with four triangular white stripes, yellow and orange-reddish- brown areas surrounding white stripes. Lat- eral white stripe passes through orbital area and is continued onto lateral edge of ros- trum. Lateral teeth of rostrum white; tip of rostrum dark brown. Fingers of cheliped purple for distal two-thirds to three-fourths of length, teeth white, remainder of fingers tan; carpus tan with brown stripe proxi- mally; merus dark brown distally, pale yel- low spot behind large distal tooth. Ambu- latory legs with faint light dorsal stripe continuous from carapace, distal ends of segments darker, merus with darker areas; dactyli with distal '4 white, apex translucent with penultimate tan band and then white. Legs with faint dark/light bands. Range.— Stenorhynchus yangi has been collected from Martha’s Vineyard south through the Gulf of Mexico to Suriname in depths from 31 to 365 m. Variation.—Within S. yangi, variation has been noted in the shape of postorbital spines, characteristic robustness of the car- apace and relative lengths of the rostrum and moveable finger of the first leg. Wil- liams (1965:244) reported the postorbital spine of S. seticornis as ““occasionally bifid.” During the course of this study, no speci- mens of Stenorhynchus seticornis were ob- served which possessed bifid postorbital spines. However, it is common to come across large specimens of S.. yangi with this condition. Several large individuals of S. yangi were examined which had single and double spines on alternate sides of the car- apace. Specimens with trifid spines were also VOLUME 102, NUMBER 3 observed, and it is probable that the indi- viduals reported by Williams (1984) with bifid spines represent S. yangi, as the gon- opod figured in that work (fig. 2410) cor- responds with S. yangi, not S. seticornis. The characteristic robustness or swelling of the various regions of the carapace also var- ies within S. yangi. This and the relative lengths of the pereopods are probably re- lated to maturity of the individual. Consid- erable variation was also noted in spination of the pereopods. Perhaps the two features which exhibit the greatest degree of variation are the ratio of total carapace length to postorbital car- apace length and the ratio of finger to palm length of the first pereopod. Only individ- uals with the rostrum intact (n = 29) were used for carapace length measurements, with individuals from the extreme ends of the geographic range included. Among males (n = 19), the postorbital carapace length ac- counted for one-fourth to one-half of the total length of the individual (0.27-—0.49); little difference from this ratio was noted among females (0.31—0.48). However, the range ratio of finger to palm length was no- tably different among males (n = 22:0.24— 0.39) and females (n = 13:0.36-0.49). A single anomaly was noted in a specimen from the Caribbean (USNM Accession #42869). This individual is a mature male with a bifid rostrum, but is normal in all other respects. Etymology.—It is my pleasure to name this species for Dr. Won Tack Yang (Texas Biomedical Institute) who, on the basis of larval characters, first recognized the pos- sibility of its distinctness. Remarks.—Yang (1967, 1976) per- formed much of the complicated work in- volved in reviewing the confused history of this genus and gave definitive proof that at least two species of Stenorhynchus are pres- ent in western Atlantic waters. His work was the first to correct the mistaken records of S. seticornis in the eastern Atlantic and showed S. /anceolatus Brullé, 1837, to be 633 the correct name for the West African species. Yang (1967) indicated that S. yangi may be a deep water species and presented data which appeared to show that the species was most often collected in waters deeper than 65 m. He noted that in the collections at the University of Miami Marine Laboratory and those examined from the U.S. National Mu- seum of Natural History, this species’ bathymetric range was from 31-119 m. He suggested the possibility of an isotherm di- viding the preferred habitats of the two species. Abele (1970) indicated a restricted occurrence of S. yangi to waters over 50 m. However, because I have not re-examined that material, that report of the species en- countered is unconfirmed. The robust nature of the carapace of this species, the absence of the interantennular spine, the lack of rostral setation, the shape of the first male pleopod and the form of the spermathecae of the female easily dis- tinguish S. yangi from S. seticornis. Differ- ences in the color patterns of the two species also aid in their identification; however, this difference may be quickly obscured by pres- ervation techniques and be of little use to the researcher studying museum specimens. Differences in relative lengths of the am- bulatory legs may also be of aid in the sep- aration of S. yangi from S. seticornis. How- ever, the fragile nature of members of this genus makes studies of this type very dif- ficult as legs are rarely collected intact. Discussion.—Taxonomic confusion sur- rounding the identity of S. seticornis and S. yangi is partly due to the great amount of variation found within the two taxa. The range of variation in the ratio of postorbital to total carapace lengths differs little from S. seticornis (0.27—-0.49) to S. yangi (0.27- 0.42). A broad range of variation is also evident in the ratio of finger to palm length (0.24—0.49 in S. seticornis and 0.24—0.45 in S. yangl). Because of the taxonomic problems, eco- logical works that have dealt with this group 634 must be used with reserve. Bathymetric rec- ords (Rathbun 1925), distributional records (Holthuis 1959; Williams 1965), substrate preferences (Rathbun 1925) and ecological and behavioral works (Hartnoll 1965; Barr 1971, 1975) must all be viewed with caution and records re-examined where feasible. Al- though the specific identity of the arrow crabs used in the reproductive studies may be of little significance in the understanding of the group’s behavior, it must be recog- nized that doubt exists as to which taxon was being studied by Schone (1968). Barr’s (1971, 1975) field work suggests that S. se- ticornis is a facultative filter feeder which climbs to the top of an outcropping or reef at dusk. By sitting motionless, the crab al- lows the passing debris to collect on the se- tae of its body during the night, and the following day is spent cleaning off the en- trapped food. No data were given by that author to indicate the frequency of this feed- ing mode. R. H. Gore (pers. comm.) noted that in aquaria, S. seticornis will snip off and eat the protruding siphon of the gastro- pod Nassarius vibex. Mary K. Wicksten (pers. comm.) has indicated that S. seticor- nis may perform an unusual decorating act by storing food gathered from the substra- tum on the rostral setae to be eaten later. This 1s a modification of the usual decora- tion behavior used for concealment and camouflage (Wicksten 1980). An alternative feeding behavior is dictated for S. yangi be- cause of the lack of rostral setae. No study has been accomplished to define the niche requirements of these two species and how these requirements may differ. The complete larval development of S. seticornis form A was described by Yang (1967, 1976). He detailed three zoeal stages and the megalopa obtained from females collected in Biscayne Bay, Florida. Yang (1967) specifically mentioned the hiatus at the base of the fingers, a character previ- ously listed as useful in separating the two forms of S. seticornis. A single zoeal stage of S. yangi (Stenorhynchus seticornis of Yang 1967, 1976) was described from a specimen PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON collected in about 225 m of water. The lar- vae of the two species are sufficiently dis- tinct to allow quick separation of the species because of a large lateral bifurcation on the carapace of S. yangi. This situation is un- usual in that the adults are similar enough to have been united under a single name for many years, but the larvae are quite differ- ent at the first zoeal stage. Acknowledgments This study has benefitted from the help of many individuals over several years. However, because of the diversity of opin- ions offered (often conflicting), not all of the reviewers’ suggestions have been incorpo- rated into this report. Any oversights or shortcomings of this report are the sole re- sponsibility of the author. John S. Garth and Jurgen Sieg helped gather obscure orig- inal descriptions. Robert H. Gore, Joseph Fitzpatrick, Richard W. Heard, Jr., Austin B. Williams, Raymond B. Manning, Robin M. Overstreet, Lipke B. Holthuis, and Dar- ryl L. Felder all provided helpful discus- sions, comments or reviews. Thomas S. Hopkins, David K. Camp, Walter Zwink, Darryl L. Felder and Torben Wolff provid- ed material. Darryl L. Felder and the staff of the University of Southwestern Louisi- ana Electron Microscopy Center provided the SEM micrographs. The reviewers of the journal provided much needed guidance in a very gracious manner. Linda Lutz pre- pared Fig. 1 and Cynthia B. Dickens typed the original manuscript. My gratitude is ex- tended to each of these for their help. Literature Cited Abele, L. G. 1970. The marine decapod Crustacea of the northeastern Gulf of Mexico. MS Thesis, March, 1970. Florida State University, Talla- hassee, Florida, 137 pp. Barr, L. 1971. Observations on the biology of the arrow crab, Stenorhynchus seticornis (Herbst) in Lameshur Bay, St. John, Virgin Islands. Pp. 213- 220 inJ. Miller, J. van Derwalker, & R. Walkers, eds., Scientists-in-the-sea. Department of the Interior, Washington, D.C. VOLUME 102, NUMBER 3 . 1975. Biology and behavior of the arrow crab, Stenorhynchus seticornis (Herbst), in Lameshur Bay, St. John, Virgin Islands. Jn Results of the Tektite Program.— Bulletin of the Museum of Natural History, Los Angeles County 20:47-56. Bosc, L.A. G. 1802. Histoire naturelle des crustacés, contenant leur description et leur moeurs; avec figures desinés d’aprés nature. Chez Deterville, Paris. Vol. 1:258 pp. Brulleé, M. 1837, 1839. Crustacés. Jn P. Barker-Webb & S. Berthelot— Histoire Naturelle des Iles Ca- naries, Zoologie, 2(2:Entomologie):13—18 (1839). Atlas (1837), unnumbered plate. Fabricius, J. C. 1793. Entomologia systematics emendata et aucta secundum classes, ordines, genera, species adjectis synonimis, loci, obser- vationbus, descriptionbus. Vol. 2:519 pp. Haf- niae, C. G. Proft. Felder, D. L. 1973. An annotated key to crabs and lobsters (Decapoda, Reptantia) from coastal waters of the northwestern Gulf of Mexico. Lou- isiana State University Sea Grant Publication Number LSU-SG-73-02. 103 pp. —., & A. H. Chaney. 1979. Decapod crustacean fauna of Seven and One-Half Fathom Reef, Texas: Species composition, abundance and species diversity.—Contributions to Marine Science 22:1-29. Garth, J. S., & L. Holthuis. 1963. Stenorhynchus La- marck, 1818 (Crustacea, Decapoda): Proposed validation under the plenary powers with des- ignation of Cancer seticornis Herbst, 1788, as type-species.—Z.N.(S.) 751 Bulletin of Zoolog- ical Nomenclature 20(6):424—428. Goeldi, E. A. 1886. Studien tiber neue und wen- iger bekannte Podophthalmen Brasiliens. — Ar- chives fur Naturglijke 52(1):19—46, pls. 2, 3, figs. 1-36. Guilding, L. 1825. An account of some rare West Indian Crustacea.— Transactions of the Linnean Society of London 14:334—338. Hartnoll, R.G. 1965. The biology of spider crabs: A comparison of British and Jamaican species. — Crustaceana 9:1-16. Hay, W. P., & C. A. Shore. 1918. The decapod crus- taceans of Beaufort, NC, and surrounding re- gion.— Bulletin of the U.S. Bureau of Fisheries 35:369-475. Herbst, J. F. W. 1788-1803. Versuch einer Natur- geschichte der Krabben und Krebse nebst einer systematischen Beschricibung ihrer verschie- denen Arten, vol. 1:1—274; vol. 2:1-—225, Berlin, G. A. Lange. Holthuis, L. 1959. The Crustacea Decapoda of Su- riname (Dutch Guiana).— Zoologische Verhan- delling, Leiden 44:1-296. Kingsley, J. S. 1880. On a collection of Crustacea from Virginia, North Carolina and Florida, with 635 a revision of the genera of Crangonidae and Pa- laemonidae.— Proceedings of the Academy of Natural Sciences of Philadelphia, 31 (for 1879): 383-427. Lamarck, J. B. de. 1818. Histoire naturelle des ani- maux sans vertébres, présentant les caractéres généraux et particuliers de ces animaux, leur distribution, leurs classes, leur familles, leur genres, et la citation des principales espéces qui s’y rapportent; précédée d’une introduction of- frant la determination des caractéres essentiels de l’animal, sa distinction du végetal et des autres corps naturelles, enfin, l’éxposition des prin- cipes fondamentaux de la zoologie. Ist ed., Vol. 5:612 pp. Paris. Leach, W. E. 1814. Crustaceology. Jn D. Brewster, ed., Edinburgh Encyclopaedia, vol. 7:383-437. Edinburgh. 1815. The zoological miscellany; being de- scriptions of new, or interesting animals. Vol. 2:154 pp. London, Nodder and Son. Linnaeus, C. 1758. Systema naturae per regna tris naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synon- ymis, locis. ed. 10, Vol. 1:824 pp. Stockholm, L. Salvius. Manning, R. B., & L. Holthuis. 1981. West African brachyuran crabs (Crustacea: Decapoda). — Smithsonian Contributions to Zoology 306:1- 379. Milne-Edwards, A. 1875. Etudes sur les Xiphosures et les Crustacés de la région Mexicicaine. Jn Mission Scientifique au Mexique et dans l’Amé- rique Centrale, No. 3, pp. 57-120, Paris. International Commission on Zoological Nomencla- ture. 1966. Opinion 763. Stenorhynchus La- marck, 1818 (Crustacea, Decapoda): Validated under the plenary powers with designation of Cancer seticornis Herbst, 1788, as type-species. — Bulletin of Zoological Nomenclature 23(1):19- oA Paula, J. 1987. The first zoeal stage of Stenorhynchus lanceolatus Brullé, 1837 (Decapoda, Brachyura, Mayjidae).—Crustaceana 53(3):276—-280. Rathbun, M. J. 1897. A revision of the nomencla- ture.— Proceedings of the Biological Society of Washington 1 1:153-167. 1900. The Brachyura and Macrura of Porto Rico.— U.S. Fisheries Commission 20:1—127. . 1925. The spider crabs of America. — Bulletin of the U.S. National Museum 129:1-613. Schone, H. 1968. Agonistic and sexual display in aquatic and semi-terrestrial brachyuran crabs. — American Zoologist 8:641-654. Slabber, M. 1778. Natuurkundige Verlustigingen, be- helzende microscopise Waarneemingen van in- en uitlandse Water- und Land-Dierren. Harr- lem, p. 162. 636 Weber, F. 1795. Nomenclator entomologicus secun- dum entomologiam systematicum ill. Fabricii, adjectis speciebus recens detectis et varietatibus. viii + 171 pp. Chilonii (Kiel) et Hamburgi. Wicksten, M. K. 1980. Decorator crabs.—Scientific American 242:146-154. Williams, A. B. 1965. The decapod crustaceans of the Carolinas.— U.S. Fishery Bulletin, 65(1):1- 298. 1984. Shrimps, lobsters, and crabs of the Atlantic coast of the eastern United States, Maine to Florida, Smithsonian Institution Press, Washington, D.C., 550 pp. Yang, W. T. 1967. A study of zoeal megalopal, and early crab stages of some oxyrhynchous crabs (Crustacea, Decapoda). Ph.D. dissertation, Uni- versity of Miami, Coral Gables, Florida. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 1976. Studies on the western Atlantic arrow crab genus Stenorhynchus (Decapoda, Brachy- ura, Majidae). 1. Larval characters of two species and comparison with other larvae of the Ina- chinae.—Crustaceana 31:157-177. Zimsen, E. 1964. The type material of I. C. Fabricius. 656 pp., 2 pls. Copenhagen. Gary D. Goeke, Minerals Management Service, Gulf of Mexico OCS Region, Office for Leasing and Environment, 1201 Elm- wood Park Boulevard, New Orleans, Lou- isiana 70123-2394. PROC. BIOL. SOC. WASH. 102(3), 1989, pp. 637-643 HOBBSEUS YALOBUSHENSIS, A NEW CRAWFISH FROM CENTRAL MISSISSIPPI (DECAPODA: CAMBARIDAE) J. F. Fitzpatrick, Jr. and Craig A. Busack Abstract.—A new crawfish, Hobbseus yalobushensis, is described from the headwaters of the Yalobusha River, ultimately a tributary of the Mississippi River. This is the first record of the genus from the Mississippi drainage. The new species is most closely related to H. prominens (Hobbs) and H. petilus Fitzpatrick. It is distinguished from them by the relative lengths of the terminal elements of the first pleopod of the first form male; females of H. yalobushensis are the only members of the genus to have a deep, long trough in the anterior part of the annulus ventralis. In 1987, one of us (CAB) received a grant from the Mississippi Natural Heritage Pro- gram to determine the current status of the rare crawfish Procambarus (Pennides) lylei Fitzpatrick & Hobbs, 1971. During the study, several specimens of a crawfish as- signable to the genus Hobbseus were col- lected. These proved to represent an un- described species and provided the opportunity for the first published record of the genus from the Yalobusha River drain- age, and thus, from the Mississippi River basin. Hobbseus yalobushensis, new species Fig. 1 Diagnosis. —Pigmented; eyes normal. Rostrum spatulate, without marginal spines; acumen reduced but usually obvious, not sharply delimited basally by strong rostral shoulders. Areola from 34.6 to 41.9% (av. 42.0%) of total carapace length (43.5—51.0%, av. 49.5% of postorbital carapace length) and from 1.75 to 2.73 (av. 2.43) times lon- ger than wide; punctations widely scattered and poorly developed, 2 to 5 across nar- rowest part. Cervical spines absent. Post- orbital ridges strong, terminating cephali- cally in rounded knob or small tubercle. Branchiostegal spine obsolete; suborbital angle lacking. Antennal scale broadest distal to midlength. Dorsal surface of palm of che- liped studded with squamous tubercles. Is- chia of only third pereiopods of males with hooks; bosses lacking on all pereiopodal coxae, but small, sparsely setose, obliquely oriented eminence on caudomesial corner of third, and ventromesial margin of fifth with obvious tubercle bearing one or two long setae. First pleopods of males sym- metrical, apices reaching just beyond caudal margin of coxae of third pereiopods and hooded by dense mat of long setae origi- nating mostly from area of lateral margin of sharply arched sternites; terminating in two parts, rami subparallel with apex of each directed at angle of about 115° to main axis of appendage; mesial process only slightly longer than central projection and tapering from base to acute tip; central projection of first form male corneous, with obscure rounded eminence at proximomesial base, and with acute tip. Annulus ventralis mov- able, subovate in outline; deep, broad trough in cephalic third overhung through most of length of one side by prominent cepha- lolateral tubercle; sinus, originating in fun- dus located beneath caudalmost part of aforementioned overhang, moving trans- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 638 ODS Lean Yt seeteet tote tatees seneces, wecesee ss Lye Ha ep casecesesesnsste® eye Petree e eA VOLUME 102, NUMBER 3 versely past midline and then turning sharp- ly caudad following gently undulant path to or nearly to caudal margin. Hand of female sparsely hirsute with comparatively few punctations above and below. Holotypic male, Form I. —Cephalothorax (Fig. 1b, J) subovate, slightly compressed laterally, deeper than wide at level of cau- dodorsal margin of cervical groove (8.0 and 7.5 mm, respectively). Abdomen longer than carapace (16.0 and 15.4 mm). Areola 2.43 times longer than wide with 3 or 4 irregular rows of poorly delineated punctations, 3 across narrowest part, constituting 36.4% of entire length of carapace (44.1% of postor- bital length). Rostrum slightly depressed an- teriorly, only weakly excavate dorsally, un- thickened elevated margins flanked mesially by deep punctations only near base; acumen slightly upturned, poorly set off from ros- trum and not reaching distal margin of pen- ultimate podomere of antennule. Subrostral ridge weak and barely visible in dorsal as- pect. Postorbital ridge strong, grooved dor- solaterally, terminating cephalically in rounded, unexpanded knob. Suborbital an- gle lacking. Branchiostegal spine obsolete. Cervical spine absent; very few deep punc- tations on carapace, deepest in vague row leading caudomesially from base of post- orbital ridge; few low squamous granula- tions in extreme cephalolateral part. Cephalic lobe of epistome (Fig. 1k) broadly subtriangular, distinctly convex with elevated margins, obtuse fovea in main body. Antennules of usual form with small spine near ventromesial margin of basal ar- ticle. Antennae extending caudally to mid- length of third abdominal tergum; antennal scale (Fig. 1h) 1.40 times longer than wide, widest distal to midlength, lateral part — Fig. 1. 639 thickened, terminating in strong acute spine and overreaching distal margin of ultimate podomere of antennal peduncle. Cephalic section of telson with single im- movable spine in each caudolateral corner and slightly movable spine just mesial to it. Chela (Fig. 17) somewhat depressed, slightly rotated mesially, subovate in cross section. Upper surface with comparatively prominent tubercles and only sparsely punctate, punctations deep only on fingers and in cluster of three proximal to base of immovable finger. Both fingers with median longitudinal ridge above and below, setif- erous punctations more common on dorsal flank of opposable margins. Opposable margin of immovable finger with three prominent tubercles in basal half, single row of minute denticles along distal half. Op- posable margin of dactyl with tubercle near midlength and second subequal tubercle near proximal one-tenth, both smaller than tu- bercles of immovable finger; single row of crowded minute denticles in distal half. Lat- eral margin of fixed finger with obtuse keel along most of its length and four tufts of setae in distal one-fourth; corresponding margin of palm only slightly undulant; inner margin of palm with single, nearly cristi- form row of 13 tubercles, flanked above by three tiny squamous tubercles and, in distal third, below by four. Low but broad squa- mous tubercle near base of dactyl. Outer margin of dactyl entire with row of sub- marginal, conspicuously setose punctations in distal half. Carpus of cheliped slightly longer than wide; dorsal surface with shallow nearly longitudinal furrow toward mesial margin and few scattered setiferous punctations, mostly in distal half; mesial margin with Hobbseus yalobushensis, all figures of holotype except d, e, morphotypic male; i, allotypic female. a, Mesial view of first pleopod; b, Lateral view of carapace; c, Lateral view of first pleopod; d, Mesial view of first pleopod; e, Lateral view of first pleopod; f Caudal view of tip of first pleopod; g, Ventral aspect of basal podomeres of pereiopods; 4, Antennal scale; i, Annulus ventralis; 7, Dorsal aspect of distal podomeres of cheliped (carpus rotated about 20° laterally to visualize stout mesial spine); k, Epistome; /, Dorsal view of carapace. 640 nine irregularly placed squamous to low spi- niform tubercles; lower submesial margin with strong, acute spine slightly distal to midlength; lower laterodistal corner with acute spine, lower mesiodistal corner lack- ing ornamentation. Merus with row of eight subacute spines along ventromesial margin and another of five along ventrolateral mar- gin, with row of four spiniform tubercles on dorsal margin, row terminating in two more small but stout subacute, adjacent spines. Ischium with three small spines along distal two-thirds of mesial margin. Hooks on ischia of third pereiopods only (Fig. lg); hooks strong, simple, slightly arched, and overhanging basioischial artic- ulation but opposing structure on basis lack- ing. Coxae of all pereiopods lacking bosses, but third with slightly globose expansion of caudomesial corner bearing irregular ar- rangement of from seven to nine long, coarse setae; fifth with typical ventromesial setose eminence and penile orifice. Sternum of third through fifth pereiopods deeply excavate and with dense tufts of setae arising from lateral margins, setae (with mi- nor contributions from proximal parts of coxae) obscuring distal half of pleopods when latter held parallel to body. First pleopods (Fig. la, c, f) as described in ‘“‘Diagnosis’’; central projection cor- neous. Allotypic female.—Except in secondary sexual characteristics, differing from holo- type in following respects: abdomen sub- equal in length to carapace; mesial margin of palm with row of 12 tubercles; opposable margin of fixed finger with proximalmost tubercle reduced to scarcely more than prominent undulation; mesial margin of carpus with 2 small tubercles and 1 small spine proximal to stout spine, no ornamen- tation distally; merus with only 3 small spines in ventrolateral row, row ending dis- tally in strong acute spine at corner; ven- tromesial row of 3 large, 1 small, followed by 3 large, 1 small, and distally, 4 large spines, dorsal surface with 2 spines on dis- talmost margin. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Annulus ventralis (Fig. 17) movable, sub- ovate in outline, highest (ventrally) just cau- dal to midlength; deep broad trough arising on cephalomedian margin and progressing obliquely to left of midpoint of annulus, high crescent-shaped eminence (or cephalo- lateral tubercle) overhanging lateral and caudal extremes of trough; sinus arising in deep fossa located in caudolateral corner of trough, progressing transversely to right of median line, there turning sharply caudad, following gently undulant path to just short of caudal margin. Postannular sclerite prominent, about half width of annulus, and not obscured by caudal margin of latter. Morphotypic male, Form IT.— Differing from holotype in following respects: areola with 4 punctations across narrowest part; proportions of chela more like allotype and inner margin of palm with row of 12 tu- bercles; mesial surface of carpus with 3 small spines proximal to stout spine, entire dor- sally; ventromesial margin of merus with row of 3 large, 1 small, and 3 large spines, lateral row represented by single quite prominent acute spine at base of distal fourth; dorsal surface of same podomere with 3 triangularly arranged stout spines near distal margin. Both terminal elements of first pleopod (Fig. 1d, e) noncorneous, less slender, sub- equal in length, and apices subacute. Hooks on ischia of third pereiopods conspicuous but markedly less developed, rounded at apex. Left first pleopod with incomplete, obscure suture delimiting basal tenth. Type locality.—The holotype and allo- type were collected from Topashaw Creek, 3.8 airmi. (6.1 km) SW of Mantee (jct. of St. Rtes. 15 and 46), T21N, R11E, at the boundary of Secs. 2 and 11, Webster Coun- ty, Mississippi. Here the creek flows through agricultural land, but it is shaded by decid- uous trees on both banks. The banks have been severely eroded by inflow from two metal culverts draining the fields and lo- cated approximately 15 m upstream of the collection site. The creek was from 1 to 2 m wide, and up to 0.7 m deep, with slow VOLUME 102, NUMBER 3 flow. The animals were collected by dip net from the streambed of bare mid-phase Por- ters Creek clay. No plant material other than leaf litter was apparent in the shaded area where the specimens were taken. Other crawfish present were large numbers of an undescribed Orconectes species and Cam- barus (Depressicambarus) striatus Hay, 1902. The morphotype was taken from Dry Creek, a Topashaw Creek tributary, 0.5 roadmi. (0.8 km) W of Hohenlinden, T15S, RIE, NE/4 SW/4 Sec. 36, also in Webster County. Disposition of the types.—The holotype, allotype, and morphotype are in the collec- tions of the National Museum of Natural History, Smithsonian Institution (USNM 219513, 219514, and 219515, respectively); the same museum also has three lots of paratypes (141,56 II, 2 2). Other paratypes are 1n the collections of the Mississippi Mu- seum of Natural Science, Jackson (MMNS; moles 15> 2 3 6 1mm., 2 2imm., 12 unsexed imm.); of the Royal Ontario Mu- seum, Toronto (ROM; 1 61, 1¢IHI,12,7¢ imm., 1 2imm.); and of the Milwaukee Pub- lic Museum (MPM;; 1 ¢II, 1 2). Range and specimens examined. — Hobbseus yalobushensis has been collected on nine occasions from six localities, all in the headwaters of the Yalobusha River, tributary to the Yazoo River, tributary to the Mississippi River: Calhoun County. (1) Small unmapped tributary of Bear Creek, 7.5 airmi. (12.2 km) SE of Calhoun City (jct. St. Rtes. 8 and 9), T22N, R1OE, center Sec. 12, (MMNS; 1 2, 1 6 imm., 2 2 imm.), 13 Feb 1987, C. Busack, M. Belk, and N. Hunt, colls.; Chickasaw County. (2) Topashaw Creek at St. Rte. 340, 4.5 roadmi. (7.3 km) W of St. Rte. 15, T15S, R2E, boundary of Secs. 20 and 21, (ROMIZ 13881;1¢1, 164 II, 7¢imm., 1 2 imm.), 14 Mar 1988, M. Belk, coll.; (3) Topashaw Creek at St. Rte. 340, 4.5 roadmi. (7.3 km) W of St. Rte. 15, T15S, R2E, boundary of Secs. 20 and 21, (MMNS; 2 ¢ I, 1 4 Il, 1 9, 1 6 imm., 1 2 imm., 12 unsexed imm.), 26 Mar 1988, C. B. and M. B., colls.; (4) Topashaw Creek at 641 county road, 2.5 roadmi. (4.0 km) W of St. Ric wot Tios: RIE, Sec" 235"°Nw/4. (MMNS; 1 ¢I, 2 4 II, 1 2, 1 ¢imm.), 26 Mar 1988, C. B. and M. B., colls. (5) roadside ditch, 3.4 roadmi. (5.5 km) NW of St. Rte. 9 on St. Rte. 404, (tributary to Sabougla Creek), (USNM 218641; 161, 17 Apr 1967, C. Craig, coll.; (6) 3.4 roadmi. (5.5 km) N of St. Rte. 9 at Bellefontaine on St. Rte. 404, (USNM 207124; 1 6 II, 1 9), 14 May 1969, T. D. Thornhill, coll.; (7) 3.4 roadmi. (5.5 km) NW of St. Rte. 9 at Bellefontaine on St. Rte. 404, (USNM 207112; 4 4 II, 1 9), 14 May 1969, C. C., coll.; (8) Dry Creek, 0.5 roadmi. (0.8 km) W of Hohenlinden, RIE, T15S, NE/4 SW/4 Sec. 36, (USNM 219515; 1 6 ID (ROMIZ 13882; 1 9) (MMNS; 2 4 II, 2 2) (MPM IZ 1988-30; 1 6 II, 1 2), 24 Apr 1987, M. B. and K. Bald- win, colls.; (9) type locality, (USNM 219513, 219514; 1 6 I, 1 2) (MMNS; 1 ¢ DJ, 5 Jun 1987, C. B. and N. Baldwin, colls. In ad- dition, two immature females collected by C. B., M. B., and C. Hill on 30 Jan 1987 from Sabougla Creek, 1 roadmi. (1.6 km) NW of Bellefontaine, T21N, R9E, SW/4 SE/4 Sec. 24, Webster County, are probably assignable to this species; this collection is at MMNS. Variations. — Most of the limits of vari- ation seen in this limited number of spec- imens is reflected in the description of the primary types, above. The second Form I male from the type locality has the apex of the rostrum broadly rounded and is without an acumen; perhaps it was broken early in life; also the mesial margin of the nght palm has arow of 15 tubercles. In one of the Form II males (12.5 mm carapace length) col- lected with the morphotype, the terminal elements of the first pleopod are adpressed throughout their length, and the central pro- jection is markedly the longer element; the pleopod also has a distinct juvenile suture proximally. The rostra of two immature specimens (1 4, 1 2) have sharply converging margins so that the acumen is acute and clearly delineated. Size.—The largest animal collected is a 642 Table 1.— Measurements (in mm) of types of Hobbs- eus yalobushensis. Morpho- Holotype Allotype type Carapace Total length 15.4 16.5 14.1 Postorbital length 27) NZ, EIS) Width ILS) 8.0 7.0 Height 8.0 8.6 7.4 Areola Length 5.6 5.8 Sell Width 23 MJ 2.1 Antennal scale Length 3.5 3.6 3.4 Width 2.5 Dee) 2.5 Rostrum Length 6.2 6.6 40) Width 4.2 4.1 3.9 Chela Length, mesial margin palm 4.6 5.0* 4.0 Width, palm 4.6 4.5* 4.0 Length, lateral margin propodus 9.6 8.8* 7.4 Length, dactyl S55) 5.4% 4.8 Abdomen Length 16.0 16.5 15.4 Width 6.5 ee 6.4 * Left chela; right regenerated. female 21.8 mm in cephalothorax length. The largest first form male is 18.0 mm, and the smallest 14.8 mm. No ovigerous fe- males or females carrying young were col- lected. For measurements of the primary types see Table 1. Color notes.—The color patterns are ex- tremely variable, ranging from medium brown with frequent irregular black splotch- es dorsally, and becoming nearly concol- orous black laterally, to uniform medium tan, except for two poorly defined dorso- laterally placed dark lines extending from the mid-cephalic area to the cephalic margin of the telson. The abdomen is pale brown dorsally, except for the aforementioned lines which degenerate to a series of irregular splotches at the tergal margins. A dark red- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON dish brown line marks the boundary be- tween pleura and terga. The dorsal surface of the chela is basically orange brown, deep- ening to very dark brown on the fingers. The fingers are nearly white at the tips. The car- - pus, except for a light oblique groove, and the merus of the cheliped are dark brown dorsally. The pereiopods are dark brown dorsally and distally, fading to pink ven- trally and proximally. The undersides of the abdomen and cephalothorax are white; those of the proximal pereiopodal segments are white but deepen to pink at the ischia or meropodites. A varying intensity of pink coloration imparts a striped appearance to the pereiopods, especially in juveniles. Associates. —H. yalobushensis has been collected in association with Cambarus (De- pressicambarus) striatus, Procambarus (Ortmannicus) hayi (Faxon, 1884), Procam- barus (Pennides) vioscai Penn, 1946, and an undescribed species of Orconectes. Relationships. —The nearest relatives of Hobbseus yalobushensis are H. prominens (Hobbs, 1966) and H. petilus Fitzpatrick, 1977. It can be distinguished from the for- mer by its less spatulate rostrum and that in H. prominens the terminal elements are slightly divergent. In both of the previously described species the mesial process is markedly longer than the central projection; the mesial process is, at best, only slightly longer in H. yalobushensis. Hobbseus yal- obushensis is unique in the genus in having such a well-developed, broad, long trough in the annulus ventralis and in having com- paratively heavy, non-uniform tuberculate ornamentation on the dorsal surface of the palm of the chela. Acknowledgments We thank all those individuals listed among the collectors, especially Marion Belk, for their assistance. We also thank Horton H. Hobbs, Jr., of the Smithsonian Institution, who compared specimens with the types of other species in the genus and VOLUME 102, NUMBER 3 offered many useful suggestions concerning the manuscript. Finally, we are grateful to the Mississippi Natural Heritage Program for funding the project which led to the ser- endipitous discovery of this new species. This paper is contribution 13 of the Uni- versity of Mississippi Freshwater Biology Research Program. The senior author was supported, in part, by the Research Com- mittee of the University of South Alabama. Literature Cited Faxon, W. 1884. Descriptions of new species of Cam- barus to which is added a synonymical list of the known species of Cambarus and Astacus. — Proceedings of the American Academy of Arts and Sciences 20:107-158. Fitzpatrick, J. F., Jr. 1977. A new crawfish of the genus Hobbseus from northeast Mississippi, with notes on the origin of the genus (Decapoda, Cambaridae).— Proceedings of the Biological Society of Washington 90:367-374. , & H. H. Hobbs, Jr. 1971. A new crawfish of the Spiculifer Group of the genus Procambarus 643 (Decapoda, Astacidae) from central Mississip- pi.—Proceedings of the Biological Society of Washington 84:95-—102. Hay, W. P. 1902. Observations on the crustacean fauna of Nickajack Cave, Tennessee, and vicin- ity. — Proceedings of the United States National Museum 25(1292):417-439. Hobbs, H.H.,Jr. 1966. Anewcrayfish from Alabama with observations on the Cristatus Section of the genus Cambarus (Decapoda, Astacidae).— Proceedings of the Biological Society of Wash- ington 79:109-116. Penn, G. H., Jr. 1946. A new crawfish of the genus Procambarus from Louisiana.—Journal of the Washington Academy of Sciences 36:27-29. (JFF) 207 North Wacker Lane, Mobile, Alabama 36608; (CAB) Freshwater Biology Research Program, Department of Biology, University of Mississippi, University, Mis- sissippi 38677. Present address (CAB): State of Washington Department of Fisheries, Room 115, General Administration Build- ing, Olympia, Washington 98504. PROC. BIOL. SOC. WASH. 102(3), 1989, pp. 644-645 HIPPOLYTE ZOSTERICOLA (CRUSTACEA: DECAPODA) IN THE EASTERN PACIFIC Mary K. Wicksten Abstract.—Specimens of Hippolyte zostericola have been taken in western Colombia. These animals fall well within the range of variation found in spec- imens from the Atlantic and Caribbean regions, and can be distinguished readily from H. williamsi, the other common hippolytid shrimp of the tropical eastern Pacific. During a survey of caridean shrimp of western Colombia, Gabriel Ramos of the Universidad del Valle sent me 18 specimens of an unidentified hippolytid shrimp, taken at San Antonio, Municipio de Robles, Tu- maco (about 2°N, 79°W), on a mud bottom, 3 Aug 1984, by Henry von Prohl. All but one were ovigerous females, no functional males were collected. The shrimp fall within the range of variation for Hippolyte zoster- icola (Smith), previously reported from Massachusetts, U.S.A. to Yucatan, the Ber- mudas, and south to Trinidad, Curacao, and Ceara, Brazil (Chace 1972, Williams 1984). The specimens from Colombia have been placed in the collections of the National Museum of Natural History, Smithsonian Institution, and the Allan Hancock Foun- dation, University of Southern California and Los Angeles County Museum of Nat- ural History. Another five specimens, ex- amined by Gabriel Ramos, have been added to the collections of the Universidad del Valle in Cali, Colombia (catalog number CRBMUV 84013). Hippolyte zostericola is variable: the ros- trum usually overreaches the antennular pe- duncle in adult females, and bears from one to three dorsal and from one to four ventral teeth. The basal article of the antennular peduncle is long and broad, without ter- minal spines. The rostrum is slightly shorter than the blade of the antennal scale. The third pereopod has a stout dactyl armed with three large, terminal spines grading into a series of smaller spines on the flexor margin. There is only one spine on the carpus and merus of the third pereopod. Chace (1972) and Williams (1984) re- marked on the similarity between H. Zos- tericola and H. pleuracanthus (Stimpson), reported from Connecticut to North Caro- lina. The latter species has a shorter ros- trum, not overreaching the antennular pe- duncle, with from one to three dorsal and from one to three ventral teeth. Chace (Fig. 48) shows H. pleuracanthus as having a ros- trum terminating in a sharp point, with two ventral subterminal teeth close to the tip, while H. zostericola is shown as having a rostrum with a nearly bifurcate tip and two ventral teeth well removed from the tip. H. pleuracanthus is shown having four spines on the merus of the third pereopod. Spec- imens of H. zostericola from the Gulf of Mexico tend to have the longest rostrum within the species, while some from Mas- sachusetts have a rostrum not exceeding the antennular peduncle. The specimens from western Colombia vary considerably. All but two have a ros- trum barely exceeding the antennular pe- duncle, with two or three dorsal and two ventral teeth and a bifurcate tip. In two, the rostrum is shorter than the antennular pe- duncle, barely exceeding the cornea of the VOLUME 102, NUMBER 3 eye. One has one dorsal and one ventral tooth, while the other has two dorsal teeth and a single apical point, not a bifurcate tip. As in H. zostericola, the basal segment of the antennular peduncle lacks spines, and the merus of the third pereopod bears one spine. The dactyls are similar in both species. I compared the specimens from Colom- bia with individuals of H. zostericola from Redfish Bay, Texas. In the latter specimens, the rostrum exceeded the antennular pe- duncle, and had two dorsal and three ven- tral teeth with a bifurcate tip. Except that the rostrum was proportionally longer, the specimens were similar. One cannot rule out the possibility that the specimens from Colombia, isolated from populations in the western Atlantic and Ca- ribbean, belong to a distinct species. Gem- inate pairs of species living on opposite sides of the Panamic land mass are known among other decapods, such as the spider crabs Pe- lia pacifica (Pacific) and P. mutica (Atlantic and Caribbean) and many others (Garth 1958). Although the specimens from the eastern Pacific are indistinguishable from H. zostericola on the basis of morphology, there may be genetic, behavioral or ecolog- ical differences that warrant their future des- ignation as a separate species. One other species of Hippolyte, H. wil- liamsi Schmitt, can be common in the trop- 645 ical eastern Pacific. It is readily distinguish- able from H. zostericola by the presence of three spines on the basal segment of the antennular peduncle. Mature females are about 25 mm in total length, while the larg- est adult H. zostericola from Colombia is 10.7 mm long. Williams (1984) gives a total length of 15.5 mm for females of H. zos- tericola. In H. williamsi, the rostrum of the female greatly exceeds the antennular pe- duncle. The two species may differ in hab- itat; H. zostericola lives among sea grasses and in sheltered bays, while H. williamsi can be common in tidepools and rocky areas with algae. Literature Cited Chace, F. A., Jr. 1972. The shrimps of the Smith- sonian-Bredin Caribbean expeditions with a summary of the West Indian shallow-water species (Crustacea: Decapoda: Natantia).— Smithsonian Contributions to Zoology 98:1—179. Garth, J. S. 1958. Brachyura of the Pacific coast of America: Oxyrhyncha.— Allan Hancock Pacific Expeditions 21(1):1—499. Williams, A. B. 1984. Shrimps, lobsters, and crabs of the Atlantic coast of the eastern United States, Maine to Florida. Washington D.C., Smithson- ian Institution Press, 550 pp. Department of Biology, Texas A&M Uni- versity, College Station, Texas 77843. PROC. BIOL. SOC. WASH. 102(3), 1989, pp. 646-650 CHACEON RAMOSAE, A NEW DEEP-WATER CRAB FROM BRAZIL (CRUSTACEA: DECAPODA: GERYONIDAE) Raymond B. Manning, Marcos Siqueira Tavares, and Elaine Figueiredo Albuquerque Abstract. — Chaceon ramosae, a species with depressed dactyli on the walking legs previously identified with C. quinquedens (Smith, 1879), is described as new, based upon material from Brazil. Almost no attempt has been made to study the deep-water fauna off the Brazilian coast since the Challenger Expedition (1873- 1876). Collections made by subsequent oceanographic expeditions such as those carried out aboard the Albatross in 1887 and the Calypso (1961-1962) were largely re- stricted to areas of the Brazilian continental shelf. Information on the Brazilian deep- water decapod crustaceans is scarce and is largely confined to studies by Miers (1886), Bate (1888), Henderson (1888) (all based on the Challenger collections), Moreira (1901), and Rathbun (1937). Knowledge of the deep-water species of Brazil has been broadened as a result of the cruise in 1987 of the Marion Dufresne, an oceanographic ship of the TAAF (Terres Australes et Antarctique Francaises). The cruise was conducted off southeastern Brazil (Fig. 1) as part of a formal agreement be- tween the Muséum National d’Histoire Na- turelle, Paris, and the Universidade Santa Ursula, Rio de Janeiro. Among the mate- rials collected were four large specimens of the deep-sea crab genus Chaceon which proved to represent an undescribed species. That species is named below. The holotype has been deposited in the Museu Nacional, Rio de Janeiro (MNRJ). Other specimens, all paratypes, are in the collections of the Museu de Zoologia, Uni- versidade de Sao Paulo (MZUSP), the Mu- seum National d’Histoire Naturelle, Paris (MNHN), and the National Museum of Natural History, Smithsonian Institution, Washington (USNM). The following abbreviations are used be- low: cb, carapace width (including lateral spines); cl, carapace length, along midline; fm, fathoms; m, meters; mm, millimeters. Chaceon ramosae, new species Figs. 2-3 Geryon quinquedens. — Rathbun, 1937:270, 271 [part, specimen from Brazil only].— Scelzo & Valentini, 1974:561 [part, spec- imens from Brazil only]. [Not Geryon quinquedens Smith, 1879.] Previous records. — Brazil: 24°17'S, 42°48’ 30” W, 671 fm (1228 m) (Rathbun 1937).— 24°28'S, 43°43'W, 800 m, and 25°13’S, 44°33'W, 1200 m (Scelzo & Valentini 1974). Material. — Brazil: 19°38’S, 38°43’W, 960 m, sta. 55 CB 95, 30 May 1987: 1 male (MZUSP 9363).—21°31'S, 40°07'W, 750- 785 m, sta. 4 CP 7, 10 May 1987: 1 male (holotype, MNRJ-MD-1381).—23°46'S, 42°09'W, 592-610 m, sta. 64 CB 105, 2 Jun 1987: 2 males (MNRJ-MD-1382; MNHN).—24°17’S, 42°48'30"W, 671 fm (1228 m), Albatross sta. 2763, 31 Dec 1887: 1 male (USNM 22072). Description. —A large Chaceon, cl to 143 mm, cb to 158 mm in adults, with 5 an- terolateral teeth on the carapace and dor- soventrally depressed dactyli on the walking VOLUME 102, NUMBER 3 SOUTH AMERICA AFRICA _ TROPIC OF CAPRICORN Fig 3) legs. Carapace 1.1 to 1.2 times broader than long. Median pair of frontal teeth separated by U-shaped sinus. Distance between sub- median frontal teeth less than distance be- tween them and lateral frontal teeth. Sec- ond, third, and fourth anterolateral teeth obsolete in adults, second and fourth small- est of all; distance from first to third tooth less than that from third to fifth tooth. Car- apace with distinct raised ridge mesial to fifth anterolateral tooth, carapace surface finely granular, especially posterolaterally, smooth only at hepatic regions. Suborbital tooth short and broad in adults, not ex- tending to level of lateral frontal teeth. Che- liped merus with sharp tooth subdistally, lacking distal tooth or angled lobe; carpus roughened dorsally, usually with irregular, curved granular ridge extending from mid- dle of proximal margin to inner spine, an- terior margin of carpus with at most an an- gled lobe but no spine; propodus with at most distal angled projection dorsally. Meri of walking legs with at most indistinct distal, dorsal tooth. Dactyli of walking legs de- pressed, height at midlength less than width. Area sampled by the Marion Dufresne in 1987. Fifth leg: merus usually less than 5.0 times (range 4.64.9 in adults, 6.4 in juvenile male) times longer than high, length 0.65 to 0.66 cb in adult; carpus with line of sharp gran- ules dorsally; propodus length 4.1 to 4.3 times height in adult, 5.1 times height in juvenile, longer than dactylus. Size.—Males only known, cl 28 to 127 mm, cb 36 to 146 mm in material exam- ined. Rathbun’s specimen is the smallest specimen reported here. Scelzo & Valentini (1974) studied two males with cl 133 to 143 mm, cb 146 to 158 mm. Color. —The specimens taken by the Marion Dufresne were cream colored in life. Scelzo & Valentini (1974) reported that the color of their two specimens was “‘cremo- so.” Depth range. —Our specimens were taken in depths between 601 (592-610) and 1228 meters, with all records from depths in ex- cess of 600 meters. Scelzo & Valentini (1974) reported two specimens from 800 m and 1200 m. The smallest specimen studied, a male with cl 28 mm, came from 1228 me- ters. 648 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON AE Fig. 2. Chaceon ramosae, male paratype, cl 107 mm, sta. 64 CB 105: a, Dorsal view; b, Carapace. Remarks. —This species resembles C. quinquedens (Smith, 1879) in having de- pressed dactyli on the walking legs, but dif- fers in numerous features: the carapace is much more granular posterolaterally, the suborbital tooth is less developed, the car- pus of the chela completely lacks an outer spine in adults, the propodus of the chela lacks a distal dorsal spine, the meri of the walking legs lack a distinct distal dorsal spine, and the propodus of the fifth leg is less than 5 times as long as high. VOLUME 102, NUMBER 3 Ol oe /d In Fig. 3. Chaceon ramosae, male paratype, cl 107 mm, sta. 64 CB 105: a, Suborbital margin; b, Dactylus of fifth leg, posterior view; c, Dactylus of fifth leg, dorsal view. Chaceon maritae (Manning & Holthuis, 1981), from West Africa, also has depressed dactyli on the walking legs, but differs from C. ramosae in several features: the carapace granulation is much coarser, the suborbital spine is smaller, and the walking legs are shorter. The merus and propodus of the fifth leg are about 4 times as long as high. The anterolateral spines of the carapace are more distinct in the smallest specimen; in very large specimens the second and fourth teeth are scarcely discernible, and the third tooth is very low, an obtuse lobe. The merus of the fifth leg is more than 6 times 649 longer than high only in the smallest spec- imen, cl 28 mm. Scelzo & Valentini (1974) reported two specimens taken off Brazil in addition to numerous specimens taken off Uruguay and Argentina. Their material from Brazil dif- fered from the other specimens they iden- tified as Geryon quinquedens in being cream rather than reddish in color, and in the length/height ratios of the propodi of the walking legs, 4.3 to 4.5, which is similar to our findings. Their material from south of Brazil has been referred to C. notialis Man- ning & Holthuis (1989), who also trans- ferred all large geryonids with 5 anterolat- eral spines on the carapace to the genus Chaceon Manning & Holthuis (1989). Etymology. —We take pleasure in naming this species for Jeanete Maron Ramos, Uni- versidade Santa Ursula, one of the organ- izers of the cruise of the Marion Dufresne off the Brazilian coast. Acknowledgments We thank Alain Guille, Muséum Natio- nal d’Histoire Naturelle, Paris, and Janete Maron Ramos, Universidade Santa Ursula, for making available the material of C. ra- mosae. Enrique Macpherson of the Instituto de Ciencias del Mar in Barcelona informed one of us (RBM) of the Chaceon taken by the French-Brazilian expedition, and he is responsible for our establishing contact. The photographs were taken by Roy Kropp and Marilyn Schotte. Lilly King Manning made the line drawings, and prepared all of the figures for publication. Literature Cited Bate, C. S. 1888. Report on the Crustacea Macrura collected by H.M.S. Challenger during the years 1873-76.—Report on the Scientific Results of the Voyage of H.M.S. Challenger during the years 1873-76, Zoology 24:xc + 942 pp., 157 pls. Henderson, J. R. 1888. Report on the Anomura col- lected by H.M.S. Challenger during the years 1873-76.—Report on the Scientific Results of 650 the Voyage of H.M.S. Challenger during the years 1873-76, Zoology 27:xi + 221 pp., 21 pls. Manning, R. B., & L. B. Holthuis. 1989. Two new genera and nine new species of geryonid crabs (Crustacea, Decapoda, Geryonidae).— Proceed- ings of the Biological Society of Washington 102(1):50-77. Miers, E. J. 1886. Report on the Brachyura collected by H.M.S. Challenger during the years 1873- 76.—Report on the Scientific Results of the Voyage of H.M.S. Challenger during the years 1873-76, Zoology 17:xli + 362 pp., 29 pls. Moreira, C. 1901. Crustaceos do Brasil. Contribui- ¢Oes para o conhecimento da fauna brasileira. — Archivos do Museu Nacional, Rio de Janeiro ji Shai ea sii Rathbun, M. J. 1937. The oxystomatous and allied PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON crabs of America.— United States National Mu- seum Bulletin 166:vi + 278 pp. Scelzo, M. A., & A. Valentini. 1974. Presencia de Geryon quinquedens Smith en aguas del Oceano Atlantico sudoccidental (Decapoda, Brachyura, Geryonidae).— Physis, Buenos Aires (A)33(87): 557-567. (RBM) Department of Invertebrate Zo- ology, National Museum of Natural His- tory, Smithsonian Institution, Washington, DC 20560; (MST and CFA) Departamento de Biologia Animal, Universidade Santa Ursula, Rua Fernando Ferrari, 75, 22131 Rio de Janeiro, RJ, Brazil. PROC. BIOL. SOC. WASH. 102(3), 1989, pp. 651-697 ON THE CRAYFISH GENUS FALLICAMBARUS (DECAPODA: CAMBARIDAE) IN ARKANSAS, WITH NOTES ON THE FODIENS COMPLEX AND DESCRIPTIONS OF TWO NEW SPECIES Horton H. Hobbs, Jr. and Henry W. Robison Abstract.—The genus Fallicambarus, embracing 16 species of burrowing crayfishes, is represented in Arkansas by eight of them, five of which are mem- bers of the nominate subgenus and three assigned to the subgenus Creaserinus. A key is provided for the identification of the 16, the combined ranges of which extend from Ontario to southwestern Texas and southwestern Georgia, and from South Carolina to Maryland. Fallicambarus (F.) petilicarpus, a close rel- ative of F. (F.) dissitus, is described from Union County, Arkansas, and F. (C.) gilpini, related to F. (C.) caesius, from Jefferson County, Arkansas. Data are presented for placing Fallicambarus (C.) uhleri (Faxon) and F. (C.) hedgpethi (Hobbs) in the synonymy of F. (C.) fodiens (Cottle). For each of the species occurring in Arkansas, as a complete a synonymy as possible is offered along with a diagnosis of the species; also included are a statement of the range, a list of localities (also noted on spot maps) and specimens examined, color notes, size ranges of adults, and life history and ecological notes. Among the more interesting and least known of the crayfishes of Arkansas are eight species assigned to the genus Fallicambarus. Like all of their congeners, those occurring in Arkansas are seldom found in permanent bodies of water, and, as adults, only after rains or during floods do they frequent tem- porary pools or runoff. Throughout most of their lives, these crayfishes inhabit burrows that are excavated in areas where, for most of the year, the water table does not drop more than a meter or so beneath the surface (in Arkansas, such areas exposed to the sun often may be recognized by the presence of hydrophilic sedges). Turrets, sometimes in the form of slender chimneys, but more often irregular mounds of earthen pellets of a size proportional to that of the crayfish, mark the scattered burrows which, in many areas, may be seen in, or adjacent to, roadside ditches. Less frequently, clusters of mounds, or even entire fields studded with turrets signify the presence of large colonies of these crayfishes. Although seldom seen during daylight hours, on warm evenings individuals, with chelipeds extended, may be observed at the mouths of their lairs, sometimes perched at the top of a chimney, or, when the humidity is sufficiently high, and particularly follow- ing a Shower, walking over the ground. Dur- ing the early spring and after rains, evidence of the presence of a crayfish in its domocile is provided by damp-to-wet pellets of soil recently deposited at openings to the bur- rows. Such signs of recent excavation are seen most frequently early in the day, before the pellets have become dry, but, if the weather is overcast, recent deposits may be found at almost any hour. At one time or another, there exist for most burrows constructed by members of the genus Fallicambarus more than one opening at the surface that lead into the sim- 652 ple or complex system of galleries. The sim- plest burrows constructed by these crayfish- es in Arkansas are those of F. (C.) fodiens (Cottle, 1863). Often they consist of a single subvertical tunnel with a slight enlargement at the fundus, but more often they are made more complex by the addition of a side pas- sage leading to or toward the surface, and on rare occasions, when a second adult shares the domicile with an ovigerous fe- male or one bearing young, there may exist a secondary gallery leading downward. Thus in its simplest form, the burrow consists of passages that are subvertical and are dis- posed in the forms of an “I,”’ “Y”’ (some- times inverted), or “X.’’ The more complex patterns usually occur in areas where during much of the year the water table lies very near or almost at the surface. The galleries of these burrows are disposed largely sub- parallel to the surface, and except for one or two that descend, presumably to the maximum depth to which the water table drops, lie relatively close to the surface. Usually there are at least two exits, one of them nearly always topped by a turret that is Sometimes open, but during dry weather may not only be closed but also the passage leading to it is often “back-filled.”’ If the burrow is that of an ovigerous female or one carrying young, all openings to the surface are frequently plugged. During wet seasons, all of the galleries may be water-filled, but at other times water may be found only at the bottom of the one or two deep passages. A few burrows have been excavated in which no standing water was present. The range of the genus Fallicambarus is a discontinuous one in which the larger seg- ment extends from Ontario southward to Arkansas County, Texas, and eastward to the Apalachicola River basin of southwest- ern Georgia. The more eastern segment en- compasses the Coastal Plain and lower Piedmont provinces from New Jersey to South Carolina. In Arkansas, the genus is confined to the Coastal Plain and foothills of the Ozark and Ouachita mountains. There PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON the nominate subgenus, which is not known to occur east of the Ouachita River basin in Arkansas and Louisiana or west of the Bra- zos Basin in Texas, is restricted to the Ouachita and Red river basins. The much more widespread subgenus Creaserinus, however, occurs in all of the major drainage basins in the state, overlapping much of the range of the subgenus Fallicambarus. Of the eight species present in Arkansas, five are members of the nominate subgenus: Fallicambarus (F.) dissitus (Penn, 1955), F. (F.) harpi Hobbs & Robison (1985), F. (F.) Jeanae Hobbs (1973), F. (F.) petilicarpus, new species, and F. (F.) strawni (Reimer, 1966); and three are assigned to the sub- genus Creaserinus: F. (C.) caesius Hobbs (1975), F. (C.) fodiens Cottle (1863), and F. (C.) gilpini, new species. Little is known about any of them except F. (C.) fodiens, and no investigation has involved a com- parison of representatives of populations throughout the range of the species. More- Over, considerations of its relationship to F. (C.) uhleri and F. (C.) hedgpethi have been only cursory, the principal reason being a lack of specimens from much of the sus- pected or assumed ranges of the three species. In many, if not most, of the collec- tions that were available, there were no first form males. The latter problem persists to date, for in less than half of the collections that we have examined is even one such specimen present. As a result we cannot be confident that the one or two males in a collection exhibit secondary sexual char- acters, the most useful that we have en- countered, that are typical of the local pop- ulation. In defense of the conclusions offered, however, we point out that such features appear to be remarkably uniform in the first form males in most localities represented in our material by two or more such males. This observation has prompted the ques- tion as to why so few of these males have been collected. The following represents primarily an ac- count of our knowledge of the genus Falli- VOLUME 102, NUMBER 3 cambarus in Arkansas, although we have attempted to summarize all available data on those species that range beyond the state boundaries. Indeed, the diversity noted in the populations of F. (F.) fodiens occurring in Arkansas provoked us into a study of representatives of the species throughout its range, one of the largest of any crayfish in North America. In presenting the synony- mies, diagnoses, bibliographic references, and summaries of published biological data for each of the species occurring in the state, we have included all of which we are aware, and the following key encompasses all of the species that have been assigned to the genus. Abbreviations used herein are as follows: AMNH, American Museum of Natural History, New York; ANSP, Academy of Natural Sciences of Philadelphia; cl, cara- pace length; CM, Carnegie Museum, Pitts- burgh; pol, postorbital carapace length; TU, Tulane University, New Orleans; USNM, National Museum of Natural History, Smithsonian Institution. Genus Fallicambarus Hobbs (1969a) Diagnosis.—“‘Adults with rostrum de- void of marginal spines. Mesial margin of palm of chela with row of fewer than 12 tubercles, opposable margin of dactyl usu- ally with prominent excision. Areola linear or obliterated [along part of its length]. An- tennal scale more than twice as long as broad. First pleopods of first form male symmet- rical and terminating in two or three distinct parts (mesial process, central projection and, occasionally, cephalic process; caudal ele- ment always absent) bent caudally or cau- domesially at angle of 90 degrees or more to main shaft or forming broad arc; central projection corneous, blade-like or tapering (but flattened laterally) and [frequently] lacking ... subapical notch; non-corneous mesial process never bulbiform but often appearing twisted and usually with emi- nence on cephalic (morphological) border 653 slightly distal to base; cephalic process, when present, small, at least partially corneous, situated mesially at base of mesial process, and directed caudally or caudodistally” (slightly modified from Hobbs 1973:463). Type species: Cambarus strawni Reimer, 1966:11. Gender: masculine. This taxon was proposed by Hobbs (1969a:111) to receive an assemblage of eight species that had been assigned formerly to the genus Cambarus. Subsequently, the di- agnosis of this assemblage was revised by him (1973:462), and speculations were of- fered on the interrelationships of the 11 members recognized at the time. In point- ing out more formally the existence of two species groups among these crayfishes, he proposed the adoption of two subgenera: the nominate subgenus comprising five species, and the subgenus Creaserinus, six. In this summary the former subgenus is considered to consist of seven species, two of which have been described since 1973 and one herein; F. (F.) spectrum has been relegated to the synonymy of F. (F.) jeanae (Hobbs 1989). The subgenus Creaserinus as pres- ently constituted consists of nine species, four of which were described since 1973 and one is added in this study. Evidence for placing Fallicambarus (C.) hedgpethi and F. (C.) uhleri in synonymy with F. (C.) fodiens is detailed herein. Key to Species of Genus Fallicambarus (Based on First Form Males) i First pleopod with proxi- momesial spur (Fig. 37) . (Subgenus Fallicambarus) . 2 ie First pleopod lacking proxi- momesial spur (Fig. 11h) ... ...(Subgenus Creaserinus) .. 8 2: Mesial ramus of uropod with distinct distolateral spine (Fig. lc); abdomen not conspicu- ously narrower than thorax . 3 on Mesial ramus of uropod lack- ing distinct distolateral spine Bi 4(3’). ia PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON (Fig. 1d); abdomen conspicu- ously narrower than thorax . 5 Mesial ramus of uropod with distomedian spine projecting beyond margin of ramus (Fig. lc); cheliped with sufflamen (Fig. 10z); central projection disposed proximocaudally, never overlapping that of cor- responding pleopod (Fig. 11h) SSE COST, Aes macneesel Mesial ramus of uropod with distomedian spine never pro- jecting beyond margin of ra- mus (Fig. 11d); cheliped with- out suffamen (Fig. 10a’); central projection disposed proximomesially, its distal ex- tremity frequently overlap- ping that of corresponding pleopod (Fig. 32) First pleopod with cephalic process (Fig. 35); length of car- pus of cheliped distinctly greater than width of palm of chela (Fig. 3h) ...... petilicarpus First pleopod lacking cephalic process (Fig. le); length of car- pus of cheliped subequal to or less than width of palm of che- Far ES SPE ES I dissitus First pleopod without cephal- ic process (Fig. lf) ....... jJeanae First pleopod with cephalic process (like Fig. lg) ....... 6 Ischia of third and fourth pe- reiopods with hooks (like Fig. a1). Sr are devastator Ischia of third pereiopods only with hooks (like Fig. 1lm).. 7 Cephalic process of first pleo- pod extending caudodistally (Fig. 1g); mesial ramus of uro- pod without distomedian spine (like Pigtidje: sols. : harpi Cephalic process of first pleo- pod closely paralleling basal part of mesial process (Fig. 10y), apical part if free direct- 8(1’). 8". 9(8). oF 10(8’). 10’. 11(10). ale: 12(11’). joe 13(10’). ed caudally; mesial ramus of uropod with distomedian pre- Marginal spine ......... strawni Ventral surface of merus of cheliped with one row of tu- bercles (Figvilig) 1G Aa 9 Ventral surface of merus of cheliped with two rows of tu- bercles (Fie ii p)t 2a ae 10 Mesial surface of dactyl of chela with tubercles in basal half (Fig. 17); mesial ramus of uropod lacking distolateral Spineo ts a2 Rete caesius Mesial surface of dactyl of chela lacking tubercles (Fig. 110); mesial ramus of uropod with distolateral spine (Fig. Ld) Oe ei Oe a gilpini Mesial surface of palm of che- la of second pereiopod not bearded (Fig. 1a); shaft of first pleopod strongly reflexed (Fig. LA) OPE ee 11 Mesial surface of palm of che- la of second pereiopod beard- ed (Fig. 15); shaft of first pleo- pod straight or only slightly curved (Figs. li, k, 1 8, 9, 10GS=x Oe ee 13 First pleopod with central projection not conspicuously long and slender (Fig. 1h) byersi First pleopod with central projection conspicuously long and slender (Fig. lj) ...... 12 Antennal scale tapering dis- tally to strong distolateral spine (Fig. 17); boss on coxa of fourth pereiopod ridgelike (Figs OE) Note ae gordoni Antennal scale rounded to subtruncate distally, lacking distolateral spine (Fig. 17); boss on coxa of fourth pereio- pod bulbiform (10b’)..... burrisi First pleopod with central projection straight and trun- cate distally (Fig. 17) .... hortoni VOLUME 102, NUMBER 3 13% First pleopod with central projection arched, and taper- ing at distal extremity or bear- ing subapical notch (Figs. 1k, sO. NOG ya Pe 14 Antennal scale with some- what distinct distal and mesial margins, their junction sub- angular (Fig. lo); abdomen broadly joined to cephalotho- PED. Re RT ae RR, Maen Sore a oe fodiens Antennal scale with distome- sial margin strongly inclined abdomen narrowly joined to BONA ss of a ra. so eS 15 First pleopod with mesial pro- cess conspicuously overreach- ing central projection (Fig. 1k) RE OA Re danielae First pleopod with mesial pro- cess overreaching central pro- jection little if any (Fig. 1/) . oryktes 14(13’). 14’. 15(14’). 15%, ee © © © © © © © © © © ee ee we ee ew ee Subgenus Fallicambarus Hobbs (1973) Diagnosis.— First pleopod of male with proximomesial spur and sometimes with cephalic process. Cheliped without suffla- men, except in F. macneesei, chela with tu- bercles scattered over most of dorsal sur- face, lateral margin of palm and basal part of fixed finger rounded, more often subser- rate or serrate, never smoothly costate. Sec- ond pereiopod of male with mesial face of chela and carpus lacking dense mats of plu- mose setae. Coxa of fourth pereiopod usu- ally with conspicuously large boss. Fallicambarus (Fallicambarus) dissitus (Penn) Figs. le, 2 Cambarus dissitus (Penn, 1955:73-80, figs. 1-13 [Types: holotype, allotype, and morphotype, USNM 98125, 98126, 98127; paratypes, ANSP, AMNH, CM, TU, USNM. Type locality: three miles east of Choudrant, Lincoln Parish, Lou- 655 istana-| 1959:5,.629. 11. 15,.16,, P7 pigs. 25, 48, 66, 79.—Penn & Marlow, 1959: 202.— Hobbs, 1959:896; 1962:274; 1967: 13.—Black, 1967:173, 178.— Fitzpatrick & Payne, 1968:14.—Hobbs III, 1969:19, 2A tab 2: Fallicambarus dissitus. —Hobbs, 1969a:111, 124, 173, fig. 20d; 1972:15, 99, figs. 81c, 82d; 1974b:23, 102, fig. 88.— Feinberg, 1971:26.—Hart & Hart, 1974:26, 30.— Bouchard, 1978:432; 1980:432.—Bou- chard & Robison, 1981:26, 29. Fallicambarus (Fallicambarus) dissitus.— Hobbs, 1973:463, 477-479, fig. 4.—Bou- chard & Robison, 1981:28.— Fitzpatrick, 1983:167.—Hobbs & Robison, 1985: 1035. Diagnosis.—Cheliped without sufflamen; ventral surface of merus with mesial and lateral rows of tubercles; length of carpus less than, or subsequal to, width of palm of chela. Chela with lateral margin at least weakly serrate, dorsal surface with scattered tubercles in lateral half, ventrolateral sur- face lacking arched row of prominent setif- erous punctations; opposable margin of dactyl with distinct excision in basal half, mesial margin with longitudinal row of tu- bercles along at least proximal fourth. Me- sial surface of palm of chela of second pe- reiopod lacking conspicuous tufts of plumose setae. First pleopod with proxi- momesial spur, lacking cephalic process; central projection strongly arched, inclined laterally at base, its distal part directed prox- imomesially and often crossing that of cor- responding pleopod. Hooks on ischia of third and fourth pereiopods. Boss on coxa of fourth pereiopod moderately strong and compressed. Mesial ramus of uropod with distolateral and distomedian spine, latter premarginal. Telson divided and with spine/s flanking anterolateral flank of su- ture. Range and specimens examined.—This crayfish is known from only a few localities in the Red and Ouachita river basins of southern Arkansas (Columbia County) and 656 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig... Same of F. (C.) fodiens; c, Dorsal view of telson and uropods of F. (F.) macneesei; d, Same from F. (C.) jeanae; e-l, Mesial view of first left pleopod; e, F. (F.) dissitus; f, F. (F.) jeanae; g, F. (F.) harpi; h, F. (C.) byersi; i, F. (C.) hortoni; j, F. (C.) burrisi; k, F. (C.) danielae, 1, F. (C.) oryktes; m—o, Antennal scale: m, F. (C.) burrisi; n, F. (C.) gordoni; 0, F. (C.) fodiens; p, Ventrolateral view of merus of cheliped of F. (C.) byersi; q, Same, F. (C.) gilpini; r, Dorsal view of chela of F. (C.) caesius; s—u, Mesial view of chela of F. (C.) fodiens; s, from Ohio; t, from Maryland; u, from Florida. Characters used in key: a, Distal podomeres of second pereiopod of Fallicambarus (C.) byersi; b, VOLUME 102, NUMBER 3 == > . bs ai ~ St ad oh Mie o ol s ae eee jal Wis Vie i} Ss 1 657 Fig. 2. Distribution of Fallicambarus (F.) dissitus (encircled stars), F. (F.) jeanae (encircled dots), F. (F.) petilicarpus (triangle) and F. (F.) strawni (dots) in Arkansas. (Some localities listed in text too close to others to be shown.) north central Louisiana (Caldwell, Grant, and Lincoln parishes). It has been found in the following localities in Arkansas. Colum- bia County: (1) Southern Arkansas Univer- sity Campus at Magnolia, 1 61, 23 Apr 1985, HWR. (2) 1 mi SW of Macedonia on St Rte 160, specimens not available, R. W. Bou- chard and HWR. These localities lie in the Dorcheat Bayou-Red River basin. Size.— The largest specimen that we have examined is a first form male from Lincoln Parish, Louisiana, having a cl of 32.7 (pol 29.1) mm. Corresponding lengths of the smallest first form males are 23.1 (20.2) mm. Life history notes. — First form males have been collected in February, March, April, and May. Neither ovigerous females nor ones carrying young have been reported. Ecological notes.—Specimens were col- lected in Grant Parish, Louisiana (8.2 miles, 13.2 km, south of the Winn Parish line on U.S. Highway 167), from moderately com- plex burrows, approximately two feet deep, in a roadside seepage area; this locality is in a rolling terrain where the soil is a sandy clay. The specimen from locality 1 was col- lected as it crawled across a lawn where there were chimneys 18 to 24 centimeters in height. Fallicambarus (Fallicambarus) harpi Hobbs & Robison Figs. lg, 5 Fallicambarus (Fallicambarus) harpi Hobbs & Robison, 1985:1035-1041, fig. 1 658 [Types: holotype, allotype, and morpho- type, USNM 217946, 217947, 217948; paratypes, MHNP, RMHL, USNM. Type locality: see page area 0.2 mi (0.32 km) east of Glenwood, Pike County, Arkan- sas]. Diagnosis.—Cheliped without sufflamen; ventral surface of merus with mesial and lateral rows of tubercles; length of carpus less than, or subequal to, width of palm of chela. Chela with lateral margin strongly serrate, dorsal surface with scattered tuber- cles in lateral half, ventrolateral surface lacking arched row of prominent setiferous punctations; opposable margin of dactyl with distinct excision in basal half, mesial margin with longitudinal row of tubercles extending almost complete length of finger. Mesial surface of palm of chela of second pereiopod lacking conspicuous tufts of plu- mose setae. First pleopod with proxi- momesial spur and freely-projecting (not adnate to mesial process), caudodistally-di- rected cephalic process; central projection strongly arched, but not inclined laterally at base, and never crossing that of correspond- ing pleopod. Hooks on ischia of third pe- reiopods only. Boss on coxa of fourth pe- reiopod very strong and compressed. Mesial ramus of uropod lacking spines. Telson di- vided but lacking spines. Inasmuch as no information has been added to our knowledge of this crayfish since it was described, the following data have been extracted from Hobbs & Robison (1985). Range and specimens examined.—The two nearby localities in the Ouachita River basin cited here are the only ones known for the species. Pike County: (1) Type lo- cality, 36 61, 13 4II, 8 2, 1 j4, 16 Apr 1982, K. Dillard; 2 61, 4 2, 21 Apr 1982, KD. (2) 0.3 mi (0.5 km) E of Glenwood on US Hwy 70,661, 3 2, 18 Mar 1982, KD. Color notes.—‘‘Dorsum of cephalic re- gion of carapace, including rostrum very PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON dark, almost black, that of thoracic region greenish tan except for small black trian- gular spot in open anterior section of areola, narrow band bordering cervical groove, and another band adjacent to posterior margin of carapace; hepatic and mandibular ad- ductor regions also black; orbital, antennal, mandibular, and branchiostegal regions greenish tan. Abdomen with broad median longitudinal, dark red stripe extending from second through fifth segment, becoming pale and disappearing before reaching caudal margin of sixth; lateral part of latter seg- ment, telson, and uropods greenish tan with dark markings; mesial ramus of uropods with black median longitudinal rib. Dorsal surface of cheliped dark green with black suffusion becoming intense mesially so that dorsodistal border of merus, mesial part of carpus, tubercular area of palm, all of dor- sum except tips of fingers and proximome- sial part of fixed finger almost black; tips of fingers and lateral and ventral surfaces of entire cheliped pinkish to yellowish cream. Remaining pereiopods cream with greenish suffusion marking dorsal parts of podo- meres from ischium through propodus, more intense on merus and propodus. Ster- nal elements and ventral surfaces of pereio- pods cream to pinkish cream” (Hobbs & Robison 1985:1039). Size.—‘“‘The largest specimen examined is a female having a carapace length of 39.6 (postorbital carapace length 35.8) mm. The smallest and largest first form males have corresponding lengths of 29.0 (25.8) mm, and 35.4 (31.5) mm, respectively” (Hobbs & Robison 1985:1039-1040). Life history:notes. — First form males were obtained in March and April. Ovigerous fe- males or ones carrying young have not been observed. Ecological notes.—The type locality con- sists of a seep “located in a pasture . . . spec- imens were collected in the early evening as they crawled about in the wet grassy areas. Soil consisted of a sandy clay with some VOLUME 102, NUMBER 3 organic material. Grasses and sedges were abundant....” Fallicambarus (Fallicambarus) jeanae Hobbs Figs. ld, f 2 Fallicambarus (Fallicambarus) jeanae Hobbs, 1973:463-469, 477, 478, 480, figs. 1, 4 [Types: holotype and allotype, USNM 144672, 144673 (61, 2); paratypes USNM. Type locality: seepage area 1.8 mi (2.9 km) E of Clark County line, Hot Spring County, Arkansas, on St Rte 84.].— Bou- chard, 1978:451; 1980:451.— Bouchard & Robison, 1981:28.—Flitzpatrick, 1983: 167, 168.—Hobbs & Robison, 1985:1035, 1040.— Hobbs, 1989. Fallicambarus (Fallicambarus) spectrum Hobbs, 1973:463, 469-478, 480, figs. 2, 4 [Types: holotype and allotype, USNM 144674, 144675 (41, 2); paratypes USNM. Type locality: 2 mi (3.2 km) E of Daisy, Pike County, Arkansas, on US Hwy 70.].—Bouchard, 1978:451; 1980:451.— Bouchard & Robison, 1981:28.—Fitz- patrick, 1983:167, 168.—Hobbs & Ro- bison, 1985:1035. Fallicambarus jeanae.—Hobbs, 1976:550, fig. 1b, d, 1.— Bouchard & Robison, 1981: 26.—Huner & Barr, 1981:50, fig 24b, d, ]; 1984:45, fig. 24b, d, 1.—Robison & Smith, 1982:53 Fallicambarus sp.— Hobbs, 1979:804. Fallicambarus spectrum.—Bouchard & Robison, 1981:26.—Robison & Smith, 1982:53 Diagnosis. —Cheliped without suflamen; ventral surface of merus with mesial and lateral rows of tubercles; length of carpus less than, or subequal to, width of chela. Chela with lateral margin strongly serrate, dorsal surface with scattered tubercles in lateral half, ventrolateral surface lacking arched row of prominent setiferous punc- tations; opposable margin of dactyl with 659 distinct excision in basal half, mesial margin with longitudinal row of tubercles extending almost complete length of finger. Mesial surface of palm of chela of second pereiopod lacking conspicuous tufts of plumose setae. First pleopod with proximomesial spur, lacking cephalic process; central projection moderately arched, not inclined laterally at base, its distal part disposed caudoproxi- mally, never crossing that of corresponding pleopod. Hooks on ischia of third pereio- pods only. Boss on coxa of fourth pereiopod very strong and compressed. Mesial ramus of uropod lacking distolateral spine; disto- median spine, if present, situated premar- ginally. Telson incised laterally but lacking spines Range and specimens examined;—This crayfish, which seems to be endemic to the upper Ouachita River basin in southwestern Arkansas, has been found in the following localities (those for which no first form males are listed should be confirmed). Clark County: (1) 1 mi (1.6 km) NE of Amity Center on St Rte 84, Sec 27NE, T5S, R23 W, DiGi Or ody 2.423) 2 1) Apr1973,"Ge B. Hobbs, J. E. Pugh, HHH; 1 ¢ II, 2 2, 2 jé, 2 j2, 29 Apr 1976, M. Kearney, HHH; 1 ¢ II, 1 2, 1 j2, 14 Apr 1979, D. D. Koym, HWR; 1 ¢4 II, 3 2, 3 j4, 15 Apr 1982, HWR, DDK. (2) roadside ditch 7.4 mi (11.8 km) E of Amity Center on St Rte 84, 1 41, 1 Q, 3 36, 30 Apr 1976, MK, HHH. (3) Richland Cemetery, 3 mi (4.8 km) NW of Gum Springs, 1 j¢, 1 j2, 24 Feb 1980, W. Laird. (4) Rest Haven Cemetery 4 mi (6.4 km) W of Arkadelphia on St Rte 8, 1 2, 1 jd, 1 j9, 24 Sep 1977, E. Laird; 1 jé, 27 Dec 1979, EL; 3 2, 6 May 1980, EL; 2 9, 1 jé, 1 j2, 12 Feb 1981; 2 6 II, 4 2, 636, 5 j2, 9 Mar 1981, EL, HWR; 1 2, 19 Mar 1981, EL. Hemp- stead County: (5) Blevins, Sec 24, T9S, R24W, 3 2, 1 j6, 29 Apr 1983, P. Lee. (6) Blevins, Sec 6, T10S, R24W, 1 jé, 1 j2, 10 May 1983, B. Hill. (7) Blevins, Sec 35, T9S, R25W, 1 2, 16 May 1983, T. Chambers. Hot Spring County: (8) roadside ditch 1.8 660 mi (2.9 km) E of Clark Co line on St Rte 84 (Type locality), 5 6 I, 2 2, 3 jé, 2 j2, 21 Apr 1973, GBH, JEP, HHH. (9) roadside ditch 3.7 mi (5.9 km) E of jct of St Rtes 7 and 84 on latter, 2 4 I, 1 j2, 30 Apr 1976, MK, HHH. (10) roadside ditch 5.2 mi (8.3 km) E of Clark Co line on St Rte 84, 1 j9, 30 Apr 1976, MK, HHH. (11) roadside ditch 4.5 mi (7.2 km) W of Marcus, 3 ¢4I, 1 4 II, 2 2, 234, 5j2, 14 Apr 1979, HWR et al. (12) 2.1 mi (3.3 km) E of Point Cedar, 2 2, 1 jé, 1 j2, 13 Mar 1981, HWR; 1 41, 2 2, 3 jé, 15 Apr 1982, HWR, DDK. Pike County: (13) roadside ditch 2 mi (3.2 km) E of Daisy on US Hwy 70, 1 46 I, 1 2, 21 Apr 1973, GBH, JEP, HHH. (14) roadside ditch 2.8 mi (4.5 km) W of Amity Center on St Rte 84, 2 6 Il, 3 2, 2 j4, 14 Apr 1979, HWR et al. (15) roadside ditch 8.2 mi (13.1 km) W of Amity Center on St Rte 84, 1 j2, 14 Apr 1979, HWR et al. Remarks.— With the acquisition of con- siderably more material than was available to him when Fallicambarus (F.) spectrum was described, Hobbs (1989) concluded that except by the color pattern this crayfish can- not be distinguished from F. (F.) jeanae. The pattern of the former, which is de- scribed below, has been observed only in specimens from the type locality. He there- fore treated spectrum as a color morph of F. (F.) jeanae. Color notes.— Fallicambarus (F.) jeanae exhibits two distinctive color morphs; that occurring in specimens from the type lo- cality was described by Hobbs (1973:468) as follows: ““Dominant color of carapace pale mauve; rostral margins, postorbital ridges, and paired subtriangular areas caudal to postorbital ridges dark brown; latter joining along cervical groove and in caudal gastric area where almost black; caudal margin of carapace dark brown. First abdominal ter- gum dark brown, remaining ones pale yel- lowish tan with paired dorsolateral cream splotches and each edged caudally with ver- milion. Telson and uropods cream with pale tan suffusion basally. Antennule and anten- na with dark yellowish-brown peduncles; PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON flagella with each article yellowish tan ba- sally and dark brown distally; lateral margin of antennal scale almost black. Cheliped mostly yellowish tan dorsally with dark bluish-brown tubercles and bluish-brown suffusion on dorsal margin of merus, dor- somesial surface of carpus, and dorsomesial part of dactyl. Ventral surface of cheliped yellowish cream. Remaining pereiopods with coloration similar to that of cheliped but lacking dark brown tubercles.” The color of the morph that was described under the name of Fallicambarus (F.) spec- trum (Hobbs, 1973:472-473) was recorded as follows: “Dominant color of carapace pale mauve gray; rostral margins and postorbital ridges almost black; paired subtriangular reddish-brown markings extending caudal- ly from caudal extremity of postorbital ridges, their basal portions coalescing in me- dian posterior gastric region, slightly over- lapping cervical groove, and filling cephalic triangular vestige of areola. Hepatic region with pale reddish-tan suffusion; caudal mar- gin of carapace edged with black. [Dorso- lateral parts of branchiostegites provided with conspicuous dark bluish purple (some almost black) spots.] First abdominal ter- gum reddish brown with succeeding terga progressively fading to uniformly reddish- tan telson and uropods; tergum also fading laterally, and pleuron concolorous with tel- son. Peduncle of antennule and antenna dark mauve gray, flagella with each article pale tan proximally, becoming dark gray distally; antennal scale with lateral thickened por- tion almost black. Chelipeds grayish blue dorsally with dark blue tubercles; ventral surface cream; remaining pereiopods simi- lar to cheliped but lacking dark blue tuber- cles.”’ Size. — The largest specimen examined by us is a female from Clark County having a cl of 40.6 (pol, 35.5) mm. The smallest and largest first form males have corresponding lengths of 30.0 (26.4) mm and 35.7 (31.7) mm, respectively. Life history notes.—All of the first form males that we have examined were collected VOLUME 102, NUMBER 3 in April. Neither ovigerous females nor ones carrying young have been observed. A first form male and female were found occupy- ing a single burrow at locality 1 on 21 Apr 1973. Ecological notes.—The largest colony of this crayfish that we have observed was in the type locality. Persistent rains for at least several days while one of us (HHH) was in the area had brought the water table throughout the seep to the surface, and the mounds marking the entrances to the cray- fish burrows were considerably eroded. They did not appear to have been neatly con- structed, and the pellets of which subse- quently observed turrets were constructed by F. (F.) jeanae seemed to have been rather haphazardly affixed to the rim, many having rolled down the side of the crude, vaguely cone like structures. The soil was primarily a sandy clay with gravel and pockets of or- ganic material. The burrows were rather shallow (most of the horizontal passages coursed at depths of 10 to 30 cm; the one or two deeper passages no doubt penetrated the soil for 70 cm to one meter), highly branching, and had two or three openings to the surface. In the immediate vicinity, grasses and sedges were growing in the water- logged soil, and nearby were trees belonging to the genera Acer, Juniperus, and Pinus. In April of 1973, the soil was so lacking in firmness that no tool was needed to aid the hand in searching for the crayfish in their burrows. In localities 1 and 13, where the “spectrum color morph”’ was found, the soil was better drained (a small, clear brook flowed through locality 1), the burrows were deeper (but extended no more than a meter beneath the surface), and the horizontal gal- leries were restricted to a smaller surface area. Fallicambarus (Fallicambarus) petilicarpus, new species Figs. 2, 3, 4a Diagnosis.—Eyes small but pigmented and with faceted cornea. Cheliped without 661 suffamen; ventral surface of merus with mesial and lateral rows of tubercles; length of carpus greater than width of palm of che- la. Chela with lateral margin weakly serrate, dorsal surface with scattered tubercles in lateral half, ventrolateral surface lacking arched row of prominent setiferous punc- tations; opposable margin of dactyl lacking pronounced excision in basal half, mesial margin with row of tubercles along at least proximal third. Mesial surface of palm of chela of second pereiopod lacking conspic- uous tufts of plumose setae. First pleopod with proximomesial spur, and free, caudally projecting cephalic process; central projec- tion strongly arched, inclined laterally at base but projecting mesially and crossing that of corresponding pleopod distally. Hooks on ischia of third and fourth pereiopods. Boss on coxa of fourth pereiopod moderately strong and compressed. Mesial ramus of uropod with distolateral spine, distomedian spine, if present, situated premarginally. Telson divided and with spine/s on an- terolateral flank of suture. Holotypic male, form I.—Body suboval, weakly compressed laterally (Figs. 3a, /, 4a). Abdomen distinctly narrower than thorax (9.6 and 14.5 mm). Greatest width of car- apace near midlength of areola where sub- equal to height (14.5 and 14.2 mm). Areola linear over most of length and comprising 37.4 percent of entire length of carapace (42.9 percent of postorbital carapace length). Rostrum with convergent, slender margins contracting anteriorly, setting off base of in- distinctly delimited acumen, apex of which corneous, upturned, and slightly overreach- ing midlength of penultimate podomere of antennular peduncle. Dorsal surface of ros- trum strongly concave, with submarginal rows of setiferous punctations and scattered ones between. Subrostral ridges rather weak but evident in dorsal aspect to base of acu- men. Postorbital ridges also weak but some- what swollen posteriorly. Branchiostegal and cervical spines absent. Suborbital angle ab- sent. Carapace punctate dorsally and very weakly and sparsely tuberculate laterally; 662 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 3. Fallicambarus (F.) petilicarpus holotype except d, h, j from allotype, and e, f from morphotype): a, Lateral view of carapace; b, c, e, Mesial view of first pleopod, cp cephalic process; d, Annulus ventralis and associated sclerites; f, g, Lateral view of first pleopod; h, 0, Distal podomeres of cheliped; i, Caudal view of first pleopods; j, Antennal scale; k, Epistome; 1, Dorsal view of carapace; m, Dorsal view of caudal part of abdomen; n, Basal podomeres of third, fourth, and fifth pereiopods. (cp, cephalic process; ps, proximomesial spur.) VOLUME 102, NUMBER 3 extreme anteroventral branchiostegal re- gion inflated, with irregular group of tuber- cles, seven or eight forming row on ventral flank of cervical groove; fewer tubercles present on dorsal flank of groove. Abdomen (Fig. 4a) little shorter than car- apace (28.0 and 30.5 mm); pleura moder- ately deep and broadly rounded ventrally, only sixth with angular caudoventral mar- gin; pleuron of first abdominal segment clearly overlapped by that of second. Telson (Fig. 3m) distinctly divided, and caudolat- eral angles of anterior section with two pairs of spines, more mesial pair movable. Prox- imal podomere of uropod with both lobes bearing distal spine, spine on more mesial lobe much stronger than that on lateral lobe; mesial ramus with well developed distolat- eral spine and smaller premarginal disto- median spine. Cephalomedian lobe of epistome (Fig. 3) broadly subtriangular with well defined cephalomedian prominence extending cephalodorsally; cephalolateral margins rather smooth; main body of epistome with cephalomedian area depressed but lacking distinct fovea. Ventral surface of proximal podomere of antennule with median spine slightly distal to midlength. Antennal pe- duncle without spines except on lateral sur- face of basis at proximal base of antennal scale, flagellum reaching second abdominal tergum. Both antennal scales of holotype with distal extremities missing, but distinct- ly reduced: blade little broader than thick- ened lateral portion and apex probably reaching only slightly beyond midlength of penultimate podomere of antennule (see Fig. 3j, 1). Mandible essentially like that of Fal- licambarus (F.) devastator Hobbs & White- man (1987: fig. 11). Ventral surface of is- chium of third maxilliped with lateral row of short, plumose setae, and mesial half studded with clusters of long, stiff setae. Right chela (Fig. 30) approximately 2.4 times as long as broad, not strongly de- pressed; width of palm slightly more than 1.2 times length of mesial margin, latter bearing row of 7 tubercles subtended dor- solaterally by row of six (left with 6 and 7, 663 respectively); dorsal surface of palm and basal part of fingers studded with squamous tubercles; those along lateral margin form- ing subserrate row extending from near proximal extremity to about midlength of fixed finger; ventral surfaces of palm, basal third of fixed finger, and mesial part of prox- imal fourth of dactyl tuberculate; more dis- toventral part of fingers with punctations, in part, flanking weak median ridges; prom- inent spiniform tubercle present on oblique distoventral ridge of palm; ventrolateral surface lacking curved row of long setae. Opposable margin of fixed finger with row of 7 tubercles (fourth from base largest) in proximal three-fifths and another (corre- sponding to usually more ventrally-placed tubercle on the chelae of most cambarids) at base of distal third; minute denticles pres- ent between distal 4 tubercles, and, except at base of distalmost tubercle, continuing to corneous tip of finger. Opposable margin of dactyl with row of 5 tubercles (third from base largest; this tubercle marking end of sharp proximal excavation in dacty] of other members of Fallicambarus) in proximal three-fourths; mesial margin of dactyl with tubercles forming subserrate row. Dorsal surface of both fingers with well defined me- dian longitudinal ridge. Carpus of cheliped conspicuously slender and long, almost 1.6 times as long as broad, and longer than either width or mesial mar- gin of palm of chela. Dorsal surface sparsely punctate and bearing poorly delimited and very shallow, oblique, longitudinal furrow; mesial surface with only one prominent subspiniform tubercle, cluster proximo- ventral to it consisting of much smaller ones; except for ventrodistal extremity bearing usual 2 moderately large tubercles, ventral and lateral surfaces punctate. Merus with single, dorsal, slightly curved row of tuber- cles, increasing in size distally; lateral sur- face punctate, and mesial one with polished and granular areas; ventral surface with both mesial and lateral rows of 15 tubercles joined by oblique row of 3. Mesioventral margin of basioischial podomere with only 2 tu- bercles corresponding to row in other cray- 664 fishes; compound podomere otherwise with scattered punctations. Chela of second pe- reiopod with marginal row of setae on palm, and carpus with dorsal row of long setae; mesial surface of carpus and propodus lack- ing tufts of plumose setae. Ischia of third and fourth pereiopods (Fig. 3n) with simple hooks, neither of which overreaching basioischial articulation and neither opposed by tubercle on correspond- ing basis. Coxa of fourth pereiopod with prominent, compressed caudomesial boss disposed somewhat in longitudinal axis of body; mesial and lateral surfaces of boss with setiferous punctations. Coxa of fifth pereiopod with small tuberculiform boss extending ventrally from caudomesial angle of podomere; ventral membrane setiferous. First pleopods (Fig. 30, c, g, i) reaching coxae of third pereiopods, carried deeply in sternum, and largely concealed by setae ex- tending from ventral margin of sternum and from coxae of third and fourth pereiopods. Proximomesial spur well developed. Shaft of appendage only slightly inclined caudal- ly; plumose setae arising from mesial sur- face of shaft forming feathery plume hiding all or part of each of three terminal ele- ments: mesial process, most proximal of three, non-corneous, somewhat tapering but with rounded apical region, disposed at slightly more than right angle to shaft of appendage; cephalic process smallest, cor- neous, subtriangular, situated between ce- phalic process and central projection, and directed caudally; and central projection most conspicuous of three, consisting of long, tapering, bladelike structure reflexed through arc of at least 150 degrees. Allotypic female.— Differing from holo- type other than in secondary sexual char- acteristics as follows: acumen even less dis- tinctly delimited basally; subrostral ridges evident in dorsal aspect for no more than one-fifth distance from caudal margin of or- bit to base of acumen; rudiment of bran- chiostegal spine present; about same num- ber of tubercles (7 or 8) on both dorsal and ventral flanks of anteroventral segment of PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON cervical groove; spine on lateral lobe of proximal podomere of uropod rudimen- tary; spine on basis of antenna tuberculi- form; flagellum of antenna extending cau- dally over no more than three-fourths length of areola; chela (Fig. 3h) proportionally shorter and broader; mesial row of tubercles on palm of chela subtended dorsally by row of only 5 tubercles; opposable margin of fixed finger with row of 5 tubercles, third from base largest; opposable margin of dac- tyl with row of 4 tubercles, second from base largest; ventral surface of merus of cheliped with mesial and lateral rows of 14 tubercles; mesioventral margin of basioischial podo- mere with 3 tubercles (left chela regener- ated). Annulus ventralis (Fig. 3d) firmly fused to sternum cephalically, approximately twice as broad as long, and strongly asymmetrical. Cephalic and cephalomedian areas de- pressed, latter distinctly excavate; dextral side of excavation elevated in massive prominence along mesial margin of which C-shaped sinus marking junction of prom- inent transverse ridge and prominence; no clearly defined sulcus evident, and fossa hidden. Postannular sclerite about 1.7 times as broad as long and less than half as wide as annulus; lateral margins weakly converg- ing toward broadly rounded anterior ex- tremity; caudal margin irregularly trans- verse. First pleopods present but not reaching anterior to postannular sclerite. Morphotypic male, form ITI.—Differing from holotype as follows: Apex of rostrum reaching base of ultimate podomere of an- tennule; right branchiostegal spine repre- sented by very small tubercle; proximal podomere of uropod lacking spine on both lobes; antennal peduncle lacking spine on basis, flagellum reaching first abdominal tergum; (as in holotype, distolateral part of both antennal scales broken); chela approx- imately 2.2 times as long as broad; left chela with only 6 tubercles in mesialmost row on palm; merus of left cheliped with lateral row of only 12 tubercles; mesioventral margin of basioischial podomere with row of 3 or VOLUME 102, NUMBER 3 4 tubercles; hooks on ischia of third and fourth pereiopods and boss on coxa of latter all clearly defined but weaker than those in holotype. First pleopods (Fig. 3e, f) reaching coxae of third pereiopods, symmetrical, and with well defined proximomesial spur; markedly similar in form to pleopod of first form male, but lacking cephalic process, and base of central projection not nearly so distinctly delimited from mesial process. Color notes.—Basic coloration olive- brown to tan. Carapace dark olive-brown; rostrum and posterior gastric area very dark; lateral areas slightly paler. First abdominal tergum dark olive-brown, second through fifth paler olive-tan and with narrow pink- ish tan arc on posteromedian margins; sixth tergum, telson, and uropods dark olive. An- tennules and antennae with peduncles olive on brown; flagella reddish brown. Chelipeds with basal three podomeres and proximal part of merus pinkish cream, latter suffused dorsally and laterally with olive, becoming dark olive distally; dorsal tubercles on mer- us green with white tips. Dorsal and lateral surfaces of carpus bright olive to forest green on brown, tubercles green and largest ones tipped with cream. Chela olive-brown dor- sally; distal ridge on palm suffused with green, and green on dorsal flank of oppos- able borders of both fingers; palm and fixed finger fading ventrolaterally to pinkish or- ange, ventral surface of all podomeres of cheliped pinkish to lavender cream. Re- maining pereiopods with olive suffusing dis- tal part of merus, carpus, and, except that of second pereiopod, proximal part of mer- us, Otherwise pale pinkish cream. Size.—The largest specimen examined is a female having a cl of 31.8 (pol 27.6) mm. The smaller of the two known first form males, the holotype, has corresponding lengths of 30.5 and 26.6 mm, respectively. Neither ovigerous females nor ones carrying young are available for determining mea- surements. Type locality. — Roadside seepage 0.2 mile east of the Columbia County line on State 665 Route 57, Union County, Arkansas (T16S, R18W, Sec 21). A field of young planted trees (Pinus) was adjacent to the seep and ditch which supported a moderate growth of sedges and grasses. Specimens were col- lected from relatively shallow, but complex burrows which, constructed in a sandy loam and topped by chimneys from 8 to 10 cm high, descended to depths of approximately 0.5 m. Some of them penetrated tangled, dense root mats of grasses growing in and on the banks of the ditch. No other crayfish was found in the immediate area. Disposition of types.—The holotype, al- lotype, and morphotype (USNM 219507, 219508, and 219509, respectively) are de- posited in the National Museum of Natural History, Smithsonian Institution, as are the paratypes consisting of 1 ¢I, 1 é II, 32, and 9 juveniles. Range and specimens examined.—All of the specimens available were collected at the type locality by the second author on 30 Apr 1982 (1 61, 1 4 Il, 2 2, and 3 juv) and 28 Mar 1988 (1 41, 2 4 II, 2 2, 6 juv). Variations.— Among the adult speci- mens, the areola constitutes from 35.2 to 38.5 percent of the total length of the car- apace, and from 39.8 to 43.6 percent of the postorbital carapace length. In one small male with a carapace length of 21.5 mm, the corresponding ratios are 34.0 and 39.0 percent, respectively. Most of the variations noted fall within the range of those noted in the descriptions of the primary types. The greatest range of differences, barring regen- erated appendages, occurs in the numbers of tubercles on the chelipeds: the opposable margin of the fixed finger may have from 4 to 8 tubercles and that of the dactyl 4 to 7; the ventromesial row on the merus ranges from 14 to 16 and the mesiolateral, from 12 to 15. In the female the annulus ventralis occurs in mirrored images of that in the allotype, and the cephalic margin may or may not be firmly fused to the sternum im- mediately anterior to it. All of the females have a linear series of 3 to 5 long setae closely associated with the distal part of the ven- 666 Table 1.—Measurements (mm) of Fallicambarus (F.) petilicarpus. Morpho- Holotype Allotype type Carapace: Entire length 30.5 pA is 29.5 Postorbital length 26.6 25.8 26.6 Width 14.5 E59 13.0 Length 14.2 b3:3 12.6 Areola: Width 0 0 0 Length 11.4 | 10.2 Rostrum: Width 4.4 4.2 4.6 Length Sef 4.5 4.8 Right chela: Length, palm mesial margin a5 4.6 73 Palm width 10.9 8.0 8.9 Length, lateral margin 24.5 16.1 19.4 Dactyl length 14.6 10.5 je Abdomen: Width 9.6 10.1 9.0 Length 28.0 21 26.0 trolateral row of tubercles on the merus; such seem not to be present in the available males. (See Table. 1 for other morphometric dif- ferences.) Life history notes.—The only collections of this crayfish available are two lots col- lected at the type locality in March and April. A first form male was obtained in both sam- ples. Ovigerous females and ones carrying young have not been observed. Ecological notes.—See ““Type locality.” Relationships. — Fallicambarus (F.) peti- licarpus is more closely allied to F. (F.) dis- situs than to any other member of the genus. The unusual first pleopods of the males of the two species with caudomesially dis- posed, crossing (at least sometimes) central projections, are so nearly alike that only af- ter a detailed re-examination of specimens of the former was the distinctive feature, the presence of a small, but well-defined ce- phalic process, observed. Subsequent com- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON parisons of the two revealed, among the many similarities, two other readily ob- served features that set F. (F.) petilicarpus apart: the very long slender carpus of the first cheliped and the unique absence in Fal- licambarus of a distinct concavity on the opposable margin at the base of the dactyl of the chela. The absence of this concavity, a feature the presence of which has served in keys until now to distinguish members of the genus Fallicambarus from Cambarus, necessitated the slight modifications in the generic diagnosis included herein. Etymology.—The name describes the slender carpus of the cheliped: Petilus (L. = slender) + carpus (L. = wrist). Fallicambarus (Fallicambarus) strawni (Reimer) Figs. 2, 10y Cambarus strawni Reimer, 1966:9, 11-14, figs. 9-18 [Types: holotype, allotype, and morphotype, USNM 116675, 116676, 116677 (61, 2, IL); paratypes, R. D. Rei- mer. Type locality: small marshy area in the Saline River basin, 2.7 mi (4.3 km) north of Dierks, Howard Co., Arkan- sas.].— Hobbs, 1967:12; 1968:K16, fig. 32; 1969a:111.—Black, 1967:173, 178.— Bouchard, 1972:61.—Hobbs III et al., 1976:24. Fallicambarus strawni.— Hobbs, 1969a:103, 111, 124, 151, 173, figs. 26, 13), 20: ae 99, 147, fig. 81b; 1974b:24, 100, fig. 81; 1976:551, fig. 1b, e.-—Bouchard & Ro- bison, 1981:26.— Robison & Smith, 1982: ao: Fallicambarus (Fallicambarus) strawni.— Hobbs, 1973:461-479, figs. 3c, h, 4.— Bouchard & Robison, 1981:28.—Fitz- patrick, 1983:168.—Hobbs & Robison, 1985:1035. Diagnosis. —Cheliped without sufflamen; ventral surface of merus with mesial and lateral rows of tubercles; length of carpus less than, or subequal to, width of palm of VOLUME 102, NUMBER 3 chela. Chela with lateral margin strongly serrate, dorsal surface with scattered tuber- cles in lateral half, ventrolateral surface lacking arched row of prominent setiferous punctations; opposable margin of dactyl with distinct excision in basal half, mesial margin with longitudinal row of tubercles extending almost complete length of finger. Mesial surface of palm of chela of second pereiopod lacking conspicuous tufts of plu- mose setae. First pleopod with proxi- momesial spur, and cephalic process adnate to mesial process, if free distally, then di- rected caudally, parallel to mesial process; central projection comparatively weakly arched, not inclined laterally at base, and disposed caudally, never crossing that of corresponding pleopod. Hooks on ischia of third pereiopods only. Boss on coxa of fourth pereiopod very strong and compressed. Me- sial ramus of uropod lacking distolateral spine, distomedian spine, if present, situ- ated premarginally. Telson entire, lacking spines. Range and specimens examined.—This crayfish seems to be endemic to southwest- ern Arkansas (although there is every reason to believe that it will be found in eastern Oklahoma) where it has been collected only in the Little and Saline watersheds in the Red River basin and in the Little Missouri watershed in the Ouachita River basin. We have examined all of the following material except that collected in locality 6. Howard County: (1) Type locality, 1 61, 1 ¢ II, 1 Q, 77 ham 1963) KR. D. Reimer: 2 6], 16 Il, 1 2, 1 jo, 1 j2, 21 Apr 1973, GBH, JEP, & HHH; 1 2, date ?, R. W. Bouchard. (2) road- side ditch 5.1 mi (8.2 km) W of Athens on St Rte 84, 1 6 I, 1 9, 29 Apr 1976, MK & HHH. (3) seep 1.8 mi (2.9 km) E of Polk Co line on St Rte 4, 3 2, 3 6 II (one ¢ later molted to form I), 29 Apr 1976, MK & HHH. (4) roadside ditch 1.8 mi (2.9 km) E of Sevier Co line on US Hwy 70, 3 jé, 6 j2, 10 Apr 1982, HWR. (5) Nashville, 1 ¢ II, 10 Apr 1986, L. Morris. (6) Reimer (1966: 4) cited this crayfish from 4 mi W of Umpire 667 on St Rte 4. Pike County: (7) roadside ditch 1.3 mi (2.1 km) E of Little Missouri River on St Rte 84, 1 2, 1 j¢, 29 Apr 1976, MK, HHH. (8) roadside ditch 0.9 mi (1.4 km) NE of Howard Co line on US Hwy 70, 1 6 I, 1 9,21 Apr 1973, GBH, JEP, HHH. Sevier County: (9) seep and ditch 0.1 mi (16 km) NE ofjct of US Hwys 71-59 and 70 on latter, A 6A 446 I, 13 9,2; 36; 5 32,20 Apr.1973, GBH, JEP, HHH. (10) seep 8.3 mi (13.3 km) E of jet of US Hwys 59-71 and 70 on latter, 1 j2, 28 Apr 1976, MK, HHH. (11) seep 0.2 mi (0.32 km) E of jct of US hwys 71 and 59 on latter, 1 j6, 1 9, 26 Apr 1976, RWB. (12) 5.8 mi (9.3 km) E of jct on St Rtes 41 and 24 on latter, 1 j¢, 9 Apr 1982, HWR. (13) seep 5.0 mi (8.0 km) NE of ject of US Hwys 59-71 and 70, 1 61, 20 Apr 1973, GBH, JEP, HHH. Color notes.—(Based primarily on first form male from locality 8.) Dominant color of carapace pinkish cream to purplish tan overlain by various shades of gray and ver- milion. Most of dorsum of cephalic region pinkish tan fading laterally to buff, often with lavender suffusion; rostrum and post- orbital ridges very dark gray margined in almost black; caudal gastric area and cer- vical groove pale to dark gray; cephalic tri- angle of areola dark gray to almost black, and linear part and caudal triangle of areola vermilion; paired pale gray longitudinal stripes flanking linear areola; remainder of branchiostegites, excluding dark bluish gray caudal margin, very pale cream tan. Tergum of first abdominal segment and cephalic part of that of second dark bluish red, otherwise yellowish tan with vermilion splotches dor- sally, fading caudally, and all pleura lighter tan ventrally. Telson largely translucent but with vermilion to brick red splotches ce- phalically, laterally, and along caudal mar- gin; uropod similarly translucent, but pe- duncle reddish tan, lateral ramus with reddish splotches lateral to median rib and Over entire distal section, and mesial ramus with red pigment largely restricted to me- dian rib and distal third. Chelipeds basically 668 tan but with dense reticulations of slate blue on dorsum of distal half of merus, that of carpus, and most of that of chela; lateral surface of palm pinkish cream. Exposed parts of peduncles of antennule and antenna mostly gray, and flagella with each article buff proximally, becoming dark gray dis- tally. Lateral margin of antennal scale very dark gray. Remaining pereiopods similar to cheliped, although with more red and less blue pigment on fifth. Ventral surface of body and pereiopods cream, latter with blue pigment toward distal ends of merus and carpus. In first form male from locality 2 (How- ard County), dorsal cephalic region darker orange tan, and with dark brownish-gray area extending across posteromedian gastric region abutting cervical groove; thoracic re- gion more apricot-colored dorsally, fading to cream tan ventrally; anterior triangle of areola almost black, branchiocardiac suture vermilion to scarlet, and posterior trian- gular area dark reddish brown. Abdomen much darker than carapace, terga of first two segments almost black anteriorly fading rapidly to brick red caudally; succeeding segments with paired subrectangular red- dish black splotches (gradually narrowing on posterior segments) dorsolaterally, flank- ing median glossy brick red longitudinal stripe, red spreading laterally on posterior part of each segment and spilling ventrally onto dorsal part of pleura, which mostly pale pinkish cream with posterior maroon spot. Telson, uropods, and chelae as de- scribed above. Size.—The largest specimen available is a female from Sevier County (locality 9, above) having a cl of 37.2 (pol 32.4) mm. The smallest and largest first form males have corresponding lengths of 24.8 (22.2) mm and 31.9 (28.5) mm, respectively. Life history notes. — First form males have been collected in April and June. Neither ovigerous females nor ones carrying young have been observed. Ecological notes.—This crayfish, like F. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON (F.) jeanae, constructs highly branching (complex) burrows in sandy clay soil. The largest colony visited by us was found at locality 9. There the roadbed is somewhat elevated above the adjoining wooded area (chiefly Pinus), and on the northern shoul- der of the elevated area, there is a seep in which scores of turrets mark the burrows of this crayfish. When one of us (HHH) first visited this locality in April 1973, the sandy clay soil was water-logged, and one could easily open and follow the complex system of galleries with one’s bare hands. On a visit three years later, following a period without rain, there were few turrets in the hard, dry ground, and, even with the aid of a shovel and considerable effort, only one juvenile was found! Subgenus Creaserinus Hobbs (1973) First pleopod never with proximomesial spur or cephalic process. Cheliped with suf- flamen; chela with tubercles on mesial sur- face of palm but sparse or lacking dorsolat- erally and laterally, lateral margin costate. Second pereiopod of male with mesial face of chela and carpus often bearing dense mats of plumose setae (lacking in F. (C.) burrisi, F. (C.) byersi, F. (C.) caesius, F. (C.) gilpini, and F. (C.) gordoni). Type species: Astacus fodiens Cottle, 1863:217. Gender: mascu- line. Fallicambarus (Creaserinus) caesius Hobbs Figs. Ir, 5 Fallicambarus (Creaserinus) caesius Hobbs, 1975:24—28, 33, fig. 7 [Types: holotype, allotype, and morphotype, USNM 144921, 133922, 133923 G I, 2, 6 I); paratypes, USNM. Type locality: Road- side ditch at Hot Spring-Saline county line, Arkansas, on St Rte 67.]; 1981:269.— Bouchard, 1978:451; 1980:451.—Bou- chard & Robison, 1981:28.— Fitzpatrick, 1983:168; 1987:439. VOLUME 102, NUMBER 3 669 Fig. 4. Dorsolateral views of Fallicambarus: a, F. (C.) petilicarpus; b, F. (F.) gilpini; c—e, F. (C.) fodiens from: c, Independence County, Arkansas; d, Santa Rosa County, Florida; e, Orangeburg County, South Carolina. Fallicambarus caesius.— Hobbs, 1975:28.— Bouchard & Robison, 1981:26. Diagnosis.— Ventral surface of merus of cheliped with mesial row of tubercles, lat- eral one never represented by more than 2. Lateral margin of chela strongly costate, never serrate, dorsal surface without scat- tered tubercles in lateral half, ventrolateral surface with arched row of prominent punc- tations bearing long setae; opposable mar- gin of dactyl with longitudinal row of tu- bercles extending along at least proximal third of finger. Mesial surface of palm of chela of second pereiopod lacking conspic- uous tufts of plumose setae. First pleopod without proximomesial spur, and lacking cephalic process; central projection weakly arched, its base not inclined laterally, its distal part directed caudoproximally, bear- ing well defined subapical notch, and never crossing central projection of corresponding pleopod. Hooks on ischia of third pereio- pods only. Boss on coxa of fourth pereiopod somewhat rounded, neither strongly com- pressed nor conspicuously protruding ven- trally. Mesial ramus of uropod with or with- out distolateral spine; distomedian spine premarginal. Telson incompletely divided and with or without spine on anterolateral flank of suture. Range and specimens examined.—Inso- far as is known, this crayfish is endemic to Arkansas where it is confined to the Oua- 670 pee = 7 Oe Ly, “By Pig. 5: PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ML OS See OL. UE Re an = op ¢ emi i coi pt eet ee ees ees ee -—— +--+ = S555 $Y} Le ae >) , = eh ey | Phy See toe ee 9 ; ees we ahh - a fea --i- bey ; ARE { oth 4 ' “ { “a 4 \ = o ih 5 . Ree At jx te os ee =-—y —~)-. LOAN : RS <———. ia — — Sei} 3 nA Distribution of Fallicambarus (F.) harpi (doubly encircled star); F. (C.) caesius (encircled dot); and F. (C.) gilpini (encircled star) in Arkansas. (Some localities listed in text for F. (C.) gilpini are too close to be noted on map.) chita and Dorcheat Bayou basins in the southern part of the state. We have exam- ined material from the following localities. Clark County: (1) Rose Hedge Cemetery at Gurdon, 1 2, 12 Mar 1983, HWR; 1 ¢ II, 1 Oo Apr 1982, WE:tld iy) bo93 46, 4jeke Apr 1983, DDK, HWR. Columbia County: (2) in Magnolia city limits, 1 2, 19 Mar 1984, J. Pesses. (3) Waldo, 1 9, 24 Aug 1981, EL. (4) seep 0.5 mi (0.8 km) W of Waldo at US Hwy 82, 1 2 with young, 17 Mar 1983, HWR. (5) seep 2 mi (3.2 km) W of Waldo at jct of Hwys 82 & 98, 4 9, 9 Apr 1983, HWR; 1 2, 3 jd, 1 j2, 2 ovig 2, 11 Feb 1984, HWR: Pop Ws Wy 9°O) Soh jes Feb 1984, HWR. (6) behind Impson Whitehead Veterinary Clinic in Magnolia, 1 61, 10 Nov 1979, M. Bryan. (7) Beene residence in Magnolia, 1 ¢ II, 22 May 1983, L. Robison. Dallas County: (8) 0.4 mi (0.64 km) N of Dallas-Ouachita Co line on St Rte 7, 1 jé, 16 Apr 1983, HWR. Hempstead County: (9) Blevins, 1 ovig 2, 11 Apr 1984, J. Tucker; 1 j4, 1j2, 20 Apr 1982, EL; 5j2, 8 Mar 1984, E. McMullen. (10) Blevins, Sec 15, T10S, R24W, 2 2, 15 Mar 1984, B. Scott. (11) Blevins, Sec 11, T10S, R24W, 3 2, 15 Mar 1984, T. Taylor. (12) Bollins Bayou near Blevins, Sec 26, T9S, R24W, 1 juv, 25 Apr 1983, HWR. (13) Blevins, Sec 6, T10S, R24W, 1 4 II, 2 2, 20 May 1983, C. Webb. (14) Blevins, Sec 20, T10S, R23W, 1 41, 29 VOLUME 102, NUMBER 3 Apr 1983, T. Winn. (15) Blevins, Sec 16, T10S, R24W, 1 41, 2 2, 19 May 1983, B. Stephens. Hot Spring County: (16) roadside ditch 2.0 mi (3.2 km) W of Grant Co line on US Hwy 270, 3 2, 1 j6, 30 Apr 1976, HHH & MK. Nevada County: (17) DeAnn Cemetery in Prescott, 2 2, 24 Nov 1980, K. W. Williams; 1 2, 10 Sep 1979, EL; 1 4], 16 Feb 1982, KWW; 1 2 I, 12 Apr 1980, KWW. (18) 3 mi (4.8 km) E of Rosston on St Rte 4, 2 4 II, 4 2, 6 jd, 2 j2, 8 Mar 1984, DDK. (19) 0.4 mi (6.4 km) from jct of St Rte 19 and Cale Rd, between Laneburg and Rosston, Sec 7, 8, T13S, R21W, 5 4], 4 9, 4 j2, 15 Apr 1983, DDK. Ouachita County: (20) 0.9 mi (1.44 km) N of jct of US Hwy 79 and St Rte 203, 1 6 II, 26 Apr 1986, HWR. Saline County: (21) roadside ditch at Hot Spring County line on US Hwy 67 (Type locality), 1 61, 2 6 II, 1 2, 2 32 (Type series), 22 Apr 1973, GBH, JEP, HHH. Color notes.—‘“‘(Based on freshly molted holotypic male.) Carapace bluish gray; dor- sal thoracic region and large arrow-shaped area (with base between origins of mandib- ular [adductor] muscles and extending to apex of rostrum) darker and more bluish than lateral surfaces of branchiostegites, he- patic, and posterior gastric regions where more olive than blue. Cephalic section of tergum of first abdominal segment midnight blue, and caudal section slate blue; succes- sive terga also slate blue but becoming pro- gressively lighter in color posteriorly to tip of telson. Second through fifth terga with reticulate, but almost symmetrical, pattern involving oblique sublinear, dorsolateral grayish cream markings. Sixth tergum and telson with ornate symmetrical light mark- ings. Uropods mostly very pale gray, but proximolateral parts somewhat darker with dark bluish splotches and dark median ribs. Antennae and pereiopods with powder blue reticulations. Antennular peduncle dark, antennal peduncle dark mesially and later- ally, but broad submedian area of penulti- mate podomere and lamellar part of anten- 671 nal scale very pale, lateral margin of scale dark. Cheliped with dorsodistal surface of merus, dorsal surface of carpus, dorsome- sial surface of palm, dorsal surfaces of fixed finger, and dactyl powder blue; both fingers with white tubercles on opposable margin and yellowish cream along distal portion; lateral costa cream, and fingers terminating in brownish cornified tips; bluish color on all podomeres fading ventrally to very pale pinkish cream; articular membranes with dark pink suffusion. Dorsal surface of re- maining pereiopods blue from merus dis- tally; basal podomeres and ventral surfaces of all pereiopods and sternum cream. Distal end of dorsal side of merus and dorsum of carpus and propodus of third maxilliped with blue reticulations’” (Hobbs 1975:27- 28). Size.—The largest specimen is a female from Nevada County having a cl of 31.4 (pol, 27.5) mm. The smallest and largest first form males have corresponding lengths of 21.1 (18.8) mm and 29.5 (26.4) mm, re- spectively. The smallest female carrying eggs or young has corresponding lengths of 27.2 (24.4) mm. Life history notes. — First form males have been collected in February, April, May, and November. Ovigerous females were found in February and April; only one, having a carapace length of 28.4 mm, seemed to be carrying anything like a full complement of eggs: 35 with diameters of 2.1 to 2.3 mm; the diameters of the few eggs carried by the other two females were 2.0 or 2.1 mm. Ecological notes. —In the type locality, this crayfish was collected from highly branch- ing burrows in “rain soaked soil consisting of clay, organic material, and some gravel” (Hobbs 1975:28). As in many, if not most, of the other known localities, sedges and grasses were present in the immediate vi- cinity of the burrows or nearby, and none was taken from burrows more than one me- ter deep, most, if not all, of which opened to the surface through two or three chim- 672 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 6. Distribution of Fallicambarus (C.) fodiens. Dots mark localities represented by at least one first form male. neys, and occasionally there was an opening to one of the galleries that was not marked by a turret. Fallicambarus (Creaserinus) fodiens (Cottle) Figs. 1b, 0, s, t, u, 6-9, 10a—x Astacus fodiens (Cottle, 1863:217) [Types: not extant. Type locality: ““Upper Cana- da,’ probably Ontario.].— Hobbs, 1969a: 111; 1973:463.— Bouchard, 1976b:586.— Page, 1985:422. Cambarus obesus Hagen, 1870:82 [in part].— Faxon, 1885:71.—Osborn & Wil- liamson, 1898:21. Cambarus argillicola Faxon, 1884:115, 116, 144; 1885:56, 72, 76-78, 160, 174, pl. IV: fig. 2; 1890:624-625; 1898:650, 690; 1914:391, 400, 424, 426.— Underwood, 1886:366.—Hay, 1891:147; 1896:478, 491-493; 1899:959, 962; 1919:232; 1920: 83.—Stebbing, 1893:208.—Osborn & Williamson, 1898:21.— Williamson, 1899:48.—Harris, 1901:191; 1903a:59, 71-72, 105, 137, 139-140, 142-144, 146, 147, 150-155; 1903b:603, 605, 608.— Ortmann, 1902:277, 280, 283; 1905:120, 123, 136; 1907:712.—Pearse, 1910a:10, 11, 15, 19, 20, pl. VU; 1910b: 732519 130.—Huntsman, 1915:158.—Cahn, 1915:136, 174.—Cummins, 1921:28- 30.—Engle, 1926:89, 93, 94, 97, 98.— Turner, 1926:146, 154, 156, 160-163, 168, 169, 178, 186-188, 192.—Creaser, 1931:263; 1932:336.—Lyle, 1937:2, 16; 1938:76.—Brimley, 1938:503.—Bou- vier, 1940:71.—Hobbs, 1942:165; 1948: 223, 224, 229, 230.—Rhoades, 1944: 98.—Bovbjerg, 1952:34.—Eberly, 1955: 283.—Crocker, 1957:90.—Hobbs & Hart, 1959:187.—Penn & Marlow, 1959:195. Cambarus uhleri Faxon, 1884:116—117, 145 [Types: holotype, MCZ 3,624; paratypes, MCZ 3,633, 3,635, 3,636. Type locality: “Swamp on Eastern Road near Felsbury, Somerset County, Maryland”’ (restricted by Faxon, 1914:426).]. Faxon, 1885:22, 59, 77-78, 160, 166, 173, pl. VIII: figs. 8, 8’, 8a, 8a’.—Underwood, 1886:373. VOLUME 102, NUMBER 3 Ml) StS) Ma Sas Poe ad Fig. 7. Distribution of Fallicambarus (C.) fodiens in Arkansas. Cambarus diogenes.—Faxon, 1885:71 [in part]. Cambarus uhleri.—Hay, 1899:959, 962; 1904:165.—Ortmann, 1902:277, 315; 1905:119, 123, 128.—Harris, 1903a:59, Bevis. 138, 141, 142, 152, 158: 1903b: 606.— Williamson, 1907:755.— Fowler, 1912:568.—Faxon, 1914:400, 426.—Hay & Shore, 1918:401, pl. 28: fig. 6.—Creas- er, 1931:269.—Brimley, 1938:503.— Hobbs, 1942:165; 1948:229; 1955:95, 98; 1959:896; 1966a:68, 70, 71; 1966b:115; 1968:K16; 1981:270.—Penn, 1955:73.— Crocker, 1957:69, 90.—Crawford, 1959: 150, 151, 177.—Meredith & Schwartz, fe so-2; 1960-4. 5,21, 23, 27,2830; 1962: 2.—Hoffman, 1963:330.—Miller, 1965: 43.—Hobbs III, 1969:42.—Hart & Hart, 1974:73, 91.—Holt, 1973:93.— Pickett & Sloan, 1979:26.—Andolshek & Hobbs, 1986:18. Cambarus (Bartonius) argillicola. —Ort- mann, 1905:120. Cambarus (Bartonius) uhleri.—Ortmann, 1905:120. Bartonius argillicola.—Williamson, 1907: PAS. 152; 753; 158; 1627 163. Cambarus (Cambarus) uhleri.— Fowler, 1912:341 [by implication]. Cambarus fodiens.—Huntsman, 1915: 158.—Creaser, 1931:263; 1932:336.— Hobbs, 1941:121; 1942:165, 167; 1948: 223, 224) 226, 229: 230: 195593" US: 1959:896; 1966b:115; 1968:K16.— Penn, 1941:8; 1955:73, 80, 81.— Rhoades, 1942: 3; 1944:98: 1948:18; 1950:2, 3, 5; 1961: 2, 4.—Hobbs & Marchand, 1943:6.— Bovbjerg, 1952:34-36, 40-54; 1970: 674 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 8. Mesial view of first pleopods of Fallicambarus (F.) fodiens from following counties in Michigan: a, Saginaw; b, c, Washtenaw. Indiana: d, e, Wells; f, Marion. Illinois: g, Cook; h, Jasper; i, Clark; j, Efhngham; k, 1, Richland. Ohio: m, Erie; n, Franklin. West Virginia: 0, Mason. Arkansas: p, q, Lawrence; r, Sharp; s, Clay; t, u, Greene; v, Craighead; w, x, Independence; y, Cross; z, St. Francis; a’, Phillips; b’, Grant; c’, Jefferson; d’, e’, f’, Dallas. 232.—Pennak, 1953:464.—Eberly, 1954: 482.—Hobbs & Hart, 1959:149, 151, 59; 1955:283.— Williams, 1954:810, 900, 159-161, 164, 169, 171, 185, 187-188, 902, 912, 918.—Spoor, 1955:77.— fig. 11.—Hart, 1959:204.—Penn & Mar- Crocker, 1957:90.— Penn & Hobbs, 1958: low, 1959:195, 202.— Wiens & Armitage, VOLUME 102, NUMBER 3 675 Fig. 9. Mesial view of first pleopods of Fallicambarus (F.) fodiens from following counties or parishes in Arkansas: a, Ashley; b, Bradley; c, Columbia; d, Miller; e, Little River; f, Sevier. Oklahoma: g, McCurtain; Texas: h, Upshur; i, Angelina; j, k, Jasper; 1, Brazos; m, Brazoria; n, Madagorda; 0, Victoria; p, Aransas. Louisiana: q, De Soto; r, Ouachita; s, Calciseau; t, East Baton Rouge. Tennessee: u, Tipton; v, Crockett; w, Shelby; x, y, Hardeman; z, Cheatham. Mississippi: a’, Lee; b’, Clay; c’, Oktibbeha; d’, Lowndes; e’, Noxubee; f’, Jones. 1961:39-54.— Bowler, 1963:128.—Mob- 135, 139, figs. 28, 37, 46, 57, 62, 85.— berly, 1965:45.—Judd, 1968:1-4, 6, 8.— Fitzpatrick & Payne, 1968:14.—Jaspers Crocker é Barr, 1968:VIL, 12, 15, 29, & Avault, 1969:637.—Waywell & Corey, 33, 35-37, 40, 56, 58-60, 125, 127, 129- 1970:1462-1464; 1972:294—298.—Bell, 676 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON f 2 Fig. 10. (All illustrations from first form males.) a—x, Mesial view of first pleopods of Fallicambarus (F.) fodiens from following counties in Alabama: a, Lauderdale; b, Limestone; c, d, Tuscaloosa; e, Perry; f, Choctaw; g, Maringo; h, Butler. Florida: i, Santa Rosa. South Carolina: j, Beaufort; k, Bamberg; 1, Colleton; m, Richland; n, Clarendon; o, Dillon; p, Marion; q, Horry. North Carolina: r, Columbus; s, Sampson; t, Hyde; u, Perquimines. Virginia: v, Norfolk; w, Warwick. Maryland: x, Dorchester. y, Same of F. (F.) strawni (cp, cephalic process); z, Lateral view of basal podomeres of cheliped of F. (C.) fodiens (su, sufflamen); a’, Same of F. (F.) jeanae; b’, Basal podomeres of fourth pereiopod of F. (C.) burrisi (cb, coxal boss); c’, Same of F. (C.) gordoni (cb, coxal boss). VOLUME 102, NUMBER 3 1971:17.—Hobbs & Hall, 1974:200.— Terman, 1974:33, 34.—Williams, Wil- liams, & Hynes, 1974:365-369, figs. 1, 2.—Caine, 1974:2.— Becker, Genoway, & Merrill, 1975:384.— Gladwell, Bowler, & Duncan, 1975:89.—Lake, 1977:59.— Berrill, 1978:166.—Momot, Gowing, & Jones, 1978:18.—Radaj, 1978:1.—Rei- nert, 1978:8.—Lawton, 1979:6.— Pickett & Sloan, 1979:26.—Bousfield, 1979: 292.—Crenshaw, Lemons, & Russo, 1980:245.—Grow & Merchant, 1980: 234.—Grow, 1981:355.—Kiley & Dji- neen, 1982:212.—Maude & Williams, 1983:68, 74, 76, figs. 6, 7.—McMahon & Wilkes, 1983:133. Cambarus (Bartonius) fodiens. —Creaser, 1931:260, 261, 263, 269-272, fig. 37. Cambarus hedgpethi Hobbs, 1948:224—230, figs. 17a—-f, h-j, 1 [Types: holotype, mor- photype, USNM 85146 (6 I, 6 II), and allotype, USNM 85147 (8); paratypes, USNM. Type locality: lower middle part of Aransas National Wildlife Refuge, Aransas Co, Texas.].— Washburn, 1953: 6.—Penn, 1953:74; 1955:73, 80; 1959:8, 14-17, figs. 9, 27, 46, 64, map 9.—Penn & Hobbs, 1958:454, 462, 465, 467, 471, 473, 476-478, figs. 11, 28, 42, 55.— Hobbs, 1959:896; 1966b:115; 1968: K16.—Penn & Marlow, 1959:195-197.— Hobbs & Barr, 1960:13.—Reimer, 1966: 14; 1969:50, 51, 53, 60, 61, figs. 2, 39.— Black, 1967:176; 1969:197.—Walls & Black, 1967:60.—Fitzpatrick & Payne, 1968:14, 20.—Walls, 1968:417.—Hobbs III, 1969:19, 21, 26, 49, 64, tab. 2. 677 336, 341, 343, 344, 350, 353, 422-426, 436, figs. 155-158.— Phillips, 1980:84.— Grow, 1981:355.— Bouchard & Robison, 1981:26, 27.—Huner & Barr, 1981:47; 1984:42.—Berrill & Chenoweth, 1982: 199.—Burr & Hobbs, 1984:14, 15, 16.— Norrocky, 1983:3.—Fitzpatrick, 1986: 126:137. Fallicambarus uhleri.—Hobbs, 1969a:111, 112, fig. 20); 1972:102, 147, figs. 82a, 83b, 84a, 85a; 1973:463, 480, figs. 3d, 4; 1974b: 24, 101, fig. 84; 1976:551, fig. la.— Hobbs & Fitzpatrick, 1970:835.—Peters, 1971: 100; 1974:74; 1975:7, 22, 23.—Hart & Hart, 1974:22, 28, 33, 73, 129.—Hobbs III, Thorp, & Anderson, 1976:24.— Hobbs &uPeterspt97 7629.12" 15494 20921; 33, 43, 46, 49, 53, 54, 60, 61, 62.— Whar- ton, 1978:50.—Page, 1985:422.—Fitz- patrick, 1986:126, 137, 138. Fallicambarus hedgpethi.—Hobbs, 1969a: 111, 112, 173, fig. 20f; 1969b:335; 1972: 102, 147, figs. 82c, 83d; 1974b:23, 100, fig. 83.— Hobbs & Hobbs, 1970:12, 14.— Hobbs & Fitzpatrick, 1970:835.—Bou- ehards3197 2:56 3062563) 4073.1977-11.— Albaugh, 1973:6, 11, 12, 25, 103.—Al- baugh & Black, 1973:183, 184, 185.— Payne & Riley, 1974:125-127.—Hart & Hart, 1974:23, 93, 94, 97.—Reimer & Clark, 1974:168, 175, figs. 27-30.—Rei- mer, 1975:24.—Lahser, 1976:278, 279, 281-—284.—Huner, Meyers, & Avault, 1976:150, 152.—Bouchard & Robison, 1981:26, 27.—Huner & Barr, 1981:57, 58; 1984:50.—Rogers & Huner, 1983:79; 1984:37; 1985:23, 24, 26-28, figs. 3, 4, Cambarus hedgepethi.—Walls & Black, 1967:60 [erroneous spelling]. Fallicambarus fodiens.—Hobbs, 1969a:111, 112, fig. 20e; 1972:102, 137, figs. 5u, 82b, 5.—Burr & Hobbs, 1984:15, 16.— Walls, 1985:193.—Page, 1985:424.—Fitzpat- rick, 1986:137. Fallicambarus (Creaserinus) fodiens. — 83c, 84b, 85b; 1974a:12; 1974b:23, fig. 82; 1976:551, fig. 2d.—Hobbs & Fitz- patrick, 1970:835.—Bouchard, 1972:52, 62, 63, 107; 1976a:14; 1976b:585, 586.— Hobbs & Barr, 1972:9.—Hart & Hart, 1974:30, 31, 128.—Hobbs & Hall, 1974: 200, 201, 203.— Page, 1974:99; 1985:335, Hobbs, 1973:463, 480, figs. 3g, 4.—Bou- chard, 1976a:14; 1976b:586.—Clark & Rhoades, 1979:238, fig. 1.—Bouchard & Robison, 1981:28.—Fitzpatrick, 1983: 168, 169, fig. 175.—Thoma & Jezerinac, 1982:136, 137.—Jezerinac & Thoma, 1984:120-124, fig. 1.—Norrocky, 1984: 678 65.—Page, 1985:335.—Jezerinac, 1986: 178.—Jezerinac & Stocker, 1987:46, fig. 1.—Hobbs III & Jass, 1988:3, 23, 39-43, 141, 142, figs. 30, 31.—Mansell, 1989. Fallicambarus (Creaserinus) uhleri.— Hobbs, 1973:463, 480, figs. 3d, 4. Hobbs & Peters, 1977 °6;-95. 127 13,198 20972 i 33, 43, 46, 49, 53, 54, 60, 61, 62.—Fitz- patrick, 1983:168.—Cooper & Ashton, 1985:9, 10.—Andolshek & Hobbs, 1986: 24. Fallicambarus (Creaserinus) hedgpethi.— Hobbs, 1973:463, 480, fig. 4.— Bouchard & Robison, 1981:28.— Fitzpatrick, 1983: 169. P{rocambarus] Fallicambarus fodiens. — Hart & Hart, 1974:88 [lapsus]. Fallicambarus hedgepethi.—Huner, 1977:11 [photo in color]. Fallicambarus sp.—Huner, 1978:621. Fallicambarus (Creserineus) fodiens.—Je- zerinac, 1983:5 [erroneous spelling]. Diagnosis.— Ventral surface of merus of cheliped with mesial and lateral rows of tu- bercles; length of carpus less than, or sub- equal to, width of palm of chela. Chela with lateral margin costate to rounded, never ser- rate, dorsal surface without scattered tuber- cles in lateral half, ventrolateral surface lacking arched row of prominent setiferous punctations; opposable margin of dactyl with distinct excision in basal half, mesial margin with longitudinal row of tubercles extending along at least basal third of finger. Mesial surface of chela of second pereiopod with conspicuous tufts of plumose setae. First pleopod without proximomesial spur and lacking cephalic process; central pro- jection comparatively weakly arched, base not inclined laterally, distal part directed caudally, with or without subapical notch, but never crossing central projection of cor- responding pleopod. Hooks on ischia of third pereiopods only. Boss on coxa of fourth pe- reiopod somewhat rounded, neither strong- ly compressed nor conspicuously protrud- ing ventrally. Mesial ramus of uropod with PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON distolateral spine; distomedian spine pre- marginal. Telson divided and with spines on anterolateral flank of suture. Range and specimens examined.— From southern Ontario southwestward to Aran- sas County, Texas, and southeastward to the Apalachicola Basin, also present in the Coastal Plain and lower Piedmont from Beaufort County, South Carolina, to Som- erset and Dorchester counties, Maryland (Figs. 6, 7). Engel (1926:93) recorded it from Lincoln, Nebraska, but this locality is so far removed from any other reported for the species that it must be confirmed. We have examined some 2500 specimens from Michigan (45), Indiana (58), Illinois (42), Missouri (20), Arkansas (420), Oklahoma (13), Texas (223), Ohio (50), West Virginia (2), Kentucky (58), Tennessee (130), Loui- siana (29), Mississippi (143), Alabama (207), Florida (14), Georgia (15), South Carolina (451), North Carolina (386), Virginia (209), and Maryland (30). Color notes.—In this crayfish, there seem to exist two basic colors and three color patterns, with a wide range of variation link- ing them. Most of the animals that we have seen are either predominantly tan with brown markings or olive-green with dark grayish to brownish green markings. The lighter ventral and ventrolateral areas may be almost white, cream, yellowish, pink, or lavender. Occasionally we have encoun- tered individuals that are predominantly blue. As for the color patterns, the striped (Fig. 4c, e) is the most common. In it, clearly defined longitudinal stripes, that are usually most obvious on the abdomen, make this pattern conspicuously different from the least common concolorous one. The latter is typified by the absence of stripes, splotch- es, or specks. Decidedly more common than the concolorous pattern 1s the speckled one (Fig. 4d; often the specks and splotches are much more abundant, especially dorsally, than depicted) in which the carapace and abdomen are marked dorsally by specks and VOLUME 102, NUMBER 3 irregular dark spots scattered over a lighter background. Paired dorsolateral concentra- tions of darker pigment, which are often more diffuse than in Fig. 4d, on the abdom- inal terga no doubt represent remnants of the dark stripes that are so conspicuous in the striped pattern. In specimens exhibiting concolorous patterns, which are not illus- trated, the dorsum of the cephalothorax is an almost uniform dark brown, green, or blue, fading laterally to cream or very light gray. The abdominal terga, too, are almost uniformly dark brown, green, or blue almost to their junctures with the pleura, which, together with the uropods, are of a much diluted hue of the color of the dorsum. All three patterns have been observed re- peatedly among specimens collected in a single locality, the most recently seen were in a collection from western Sevier County, Arkansas, less than a mile from the Okla- homa line. Hay (1904:165) remarked that the spec- imens from Maryland were “‘a dirty green- ish brown, the tips of the chelae alone being somewhat reddish.” He reported further that Uhler, who first collected the species in Maryland, related to him the presence of beautiful yellow spots. While we have not observed the latter, we have encountered specimens with cream, yellow, orange, and red tipped chelae, but we have not associ- ated any of these colors with a particular geographic region. Size.— The largest specimen of this species that we have examined 1s a first form male from Richland County, Illinois, which has a cl of 42.8 (pol 36.7) mm. The smallest first form male (from Perquimens County, North Carolina) that we have seen has correspond- ing lengths of 19.5 (16.0) mm. Those of the largest and smallest from Arkansas are 36.4 (32.4), from St. Francis County, and 22.8 (19.2) mm, from Dallas County. Compa- rable measurements of the smallest female carrying eggs or young that we have seen are 26.5 (22.6) mm; this specimen was col- lected in Columbia County, Arkansas. 679 Life history notes.—Considering popula- tions throughout the range of the species, first form males have been collected during every month of the year; ovigerous females were found from January to June and in September, October, and November, and females carrying young from January to April and in September. In Maryland, this crayfish was reported by Hay (1904) to leave its burrows in the spring when it becomes common in ditches and small streams. Finding both first and second form males in September led him to conclude that the transition in males from form II to form I must occur in the late fall. The data for Arkansas are indeed inade- quate; except for a total of 11 members of the species collected in June, July, and No- vember, all of the material from the state available to us was collected during Feb- ruary (11), March (31), April (250), and May 15). Among the collections from the state, there are first form males collected in Feb- ruary, March, April, May, and November, Ovigerous females in February, April, and November, and others carrying young in January, February, and March. The egg complements of three of the ovig- erous females appear to be reasonably com- plete and are as follows: cl 28.2, pol 24.8 mm, 190 eggs; cl 28.2, pol 24.2 mm, 196 eggs; cl 26.5, pol 22.6 mm, 177 eggs. The diameters of the eggs were 1.9 and 2.0 mm. Ecological notes.—Ecological data that have been recorded pertaining to F. (F.) fo- diens were recently summarized by Hobbs III & Jass (1988:41-43). The following dis- cussion is therefore limited to a few record- ed observations on the eastern facies of Fal- licambarus (C.) fodiens (formerly F. (C.) uhleri) and to those made by us on the pres- ence of this crayfish in Arkansas, where it frequents temporary pools and burrows from the floodplains of the major rivers to the foothills of the Ozark and Ouachita moun- tains. According to Faxon (1884:117), his C. uhleri occurs in “‘salt marshes, covered twice daily by the tides, and also in brackish 680 and fresh-water ditches... .”” On the eastern shore of Maryland, Hay (1904:165) found it to be “rather abundant in burrows in low- lying areas not far from the bay but always near ponds or ditches of freshwater. In near- ly every case the area selected was in dense pine woods.”’ Hay also learned from local inhabitants of the area that in the spring the crayfish ““emerge from their burrows and are common in ditches and small streams.” The original collection of the species taken by Dr. Philip Uhler was made partly in water that was distinctly brackish. In Arkansas, this crayfish is primarily an inhabitant of temporary bodies of water and burrows, although occasionally it ventures into more permanent lentic and lotic hab- itats. Apparently wherever it occurs in the State the water table must be within range of its burrowing capabilities, for we have no evidence that any members of the species undergo a life-span devoid of periods of what is an apparently voluntary fossorial exis- tence. Whereas we have retrieved a few specimens from burrows that did not pen- etrate the water table, most were taken from pockets of water in them at depths of 0.5 to 1.5 meters. The external appearance of these burrows is described in the introductory re- marks where it was also pointed out that those excavated by members of this species are comparatively simple. Most consist of a subvertical passageway, in clay, sandy clay, or sandy loam, opening to the surface through one or two apertures that may or may not be surrounded by low, for the most part poorly formed, chimneys, and at the fundus of the passageway there is a slight enlargement. Occasionally we have encoun- tered a burrow with a second subvertical gallery leading downward from the main one. With little doubt, these simple I-, Y- or X-shaped patterns reflect, in part, phys- ical features of the environment involving the availability and retention of water. In habitats where there is evidence (based on the presence of hydrophytic plant commu- nities) that during much of the year the water PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON table is close to the surface, this species is largely confined to the marginal areas, and another (e.g., F. (C.) caesius or F. (C) gilpini) dominates, with its shallow, dendritic sys- tems of galleries, the more permanently wet parts of the seasonal seeps. The burrows of those individuals that invade the wetter areas do branch more than those we have regarded as more typical. Clumps of sedges frequently mark the sites of temporary stands of water, and areas in which a not- too-deep retreat of the water table occurs, features that together provide what appears to be the favorite habitat of the species in Arkansas. Most of the burrows that we have dis- sected contained a single individual, occa- sionally one inhabited by a female and a first form male, and sometimes one housing a female with young either clinging to her abdomen or sometimes with young that are presumed to have abandoned their mother. We have no information as to the perma- nency of habitation of a domocile by an individual. Until the current study of J. Norrocky (manuscript in preparation) in- volving marking and recapturing members of the species in Ohio, we were almost con- vinced that a female was a nearly permanent resident of the burrow she occupied, never wandering far away. This assumption was made, at least in part, on the basis of the size of the turrets marking the entrance/s. There can be little doubt that the larger tur- rets mark the burrows inhabited by the larg- er females, and there is no question that the more massive chimneys reflect more spa- cious and deeper domiciles. These obser- vations suggested to us the probability that the older, larger, females had spent a longer time enlarging and reworking their domi- ciles than had the smaller females and males, probably spending the better part of their entire lives hauling soil pellets to the sur- face. In light of some of Norrocky’s data (personal communication) there is good reason to question our conclusion as to the tenure of an individual in a single lair. While VOLUME 102, NUMBER 3 the males must have their own burrow at the time of their presumed biannual molts, there is good reason to believe that when in form I they abandon their burrows, at least temporarily, for sojourns in the domains of one or more females that might be visited. The question as to whether or not they re- turn to their homestead following the breed- ing season or seek seclusion elsewhere re- mains unanswered. | Following rains, when pools flood the mouths of the burrows, the juvenile element of the population emerges in numbers and small crayfish may be observed wandering hither and yon, even during daylight hours. An occasional adult also appears in the open water, and, no doubt, many more adults leave their lairs for short forays in the pools at night, but most return to burrows by at least the early morning hours. We are much puzzled by the paucity of colonies of this crayfish throughout most of the rice-growing areas of the state. Surely when the region was still wooded, and shal- low temporary pools were common features of the landscape, numerous colonies of F. (F.) fodiens must have existed between the Arkansas and Mississippi rivers. Now, one must search rather diligently to find even an isolated burrow in much of the area un- der cultivation. It is understandable that a crayfish might have difficulty in remaining well established in fields that are subjected to the treatment accorded the cultivation of rice, but why should they not be present in the roadside ditches that border these vast tilled and alternately flooded and drained lands? In Cross County, for example, a care- ful search of the ditches along highways and secondary roads for miles revealed only one colony of this crayfish, though the lawn around one of the churches in Hickory Grove was pitted by scores of burrows that must have been constructed by members of this species. Not only is this crayfish largely absent where it should occur in numbers, but other species (Procambarus (Ortman- nicus) acutus acutus (Girard, 1852), Cam- 681 barus (Lacunicambarus) diogenes Girard, 1852, and C. (L.) ludovicianus Faxon, 1914) are also encountered infrequently. It seems likely that something associated with the production of rice is affecting adversely the exploitation of the area by crayfishes. We are further puzzled by our failure to find a single member of this species between the Arkansas and White rivers downstream from Pope County. We do not claim to have exhausted possibilities of the existence of overlooked colonies, but considerable effort has been expended along several routes tra- versing the area between the two rivers. Remarks.— United here are three for- merly recognized species which for a num- ber of years have presented difficulties to one of us (HHH) in searching for charac- teristics that might be used in their differ- entiation (e.g., Hobbs 1959, 1972, 1973, 1981). In preparing an account of the Fal- licambarus from the Apalachicola basin in his study of the crayfishes of Georgia, Hobbs (1981) assigned the specimens that he had previously identified as members of Cam- barus fodiens (Hobbs & Hart, 1959:187) to F. hedgpethi. But before doing so he had vacillated between assigning them to one of the two and to describing them as new! Had the material subsequently collected in east- ern Arkansas and from a number of other localities, as well, been available to him, it is likely that he would have arrived at the conclusions that have been reached in the current study. Given specimens from the vicinity of the type localities of these three crayfishes, we do not believe that anyone would have dif- ficulty in distinguishing between them: the first pleopod of the first form males of F. fodiens exhibits a much shorter central pro- jection than do those of either F. hedgpethi or F. uhleri, and the latter has an areola that constitutes less than 39 percent of the length of the carapace whereas that of hedgpethi is more, and the opposable margin of the dac- tyl of the chela with two instead of one ma- jor tubercle is typical only of F. hedgpethi: 682 too, in the latter the arrangement of the tu- bercles on the mesial surface of the dactyl of the chela in two well developed rows dif- fers from the usual single well developed row in the other two. The limited known range of fodiens when uhleri was described from Maryland by Faxon in 1884, and the existing poor concept of the distribution of the two when hedgpethi was found in south- western Texas, gave neither Faxon nor Hobbs reason to question the validity of the seemingly distinctive characters that they chose in naming what we now believe to be peripherally located populations of a single species. These occur at the angles of a large, distorted, triangular range which in Plio- cene, and probably in part of Pleistocene, times must have been continuous. The range appears even now to be unbroken except for a gap apparently existing across the south- ern part of Georgia where members of the subgenera Hagenides and Leconticambarus of the genus Procambarus are probably vi- cariating for F. fodiens (see Hobbs 1981: 317, 348). Comparisons of the materials from throughout the range of the species have been made, and we have discovered only a few characters (those associated with the first pleopod of the first form male) that are geo- graphically or ecologically restricted to a limited part of the range of the species. The measurements made of the carapace and chelipeds that have been translated into ra- tios suggest that some local populations are rather distinctive, but they, too, are not con- sistent for large segments of the range, and there are no indications of clinal trends. Un- til now, the Fallicambarus ranging along the Atlantic versant from South Carolina to Maryland has been identified as F. (C.) uh- leri. In southern South Carolina the areola of this crayfish spans from 36 to 41 percent of the carapace length, in North Carolina from 34.8 to 38.7 percent, and in Virginia and Maryland 35.2 to 38.5 percent, and in several localities in South Carolina the ratio is distinctly above 39 percent. When these PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON specimens from South Carolina are com- pared with others from the state, we find nothing else that will set them apart, and in the midst of their range there are specimens with areolae occupying as little as 36 percent of the carapace length. If the comparisons are extended to specimens from more west- ern localities including those from Arkansas and Texas the ratios range from 32.7 in Cheatham County, Tennessee to 41.5 in Tuscaloosa County, Alabama, and Brazos County, Texas. Moreover, there seems to be no distributional pattern in the varia- tions between these extremes. Even the three color patterns (concolor- ous, speckled, and striped) that have been noted appear in a single population. Most of the specimens that we have examined from Illinois to Ohio and West Virginia northward possess chelae in which the op- posable margin of the fixed finger bears one tubercle that is slightly to distinctly larger than the others, and the tubercles on the mesial surface of the dactyl are largely aligned in a single row. Specimens from the Atlantic versant exhibit, for the most part, similarly adorned chelae. In contrast, how- ever, in most of those in the lower gulf coast- al area, the fixed finger bears two large tu- bercles and those on the mesial margin of the dactyl form two well developed rows. In eastern Arkansas, Tennessee, and Ala- bama, these features appear in a haphazard fashion. First pleopods of first form males from throughout the range of the species are de- picted in mesial aspect in Figs. 8-10. On the basis of variations noted in specimens from Ontario (see Crocker & Barr 1968: fig. 28), Michigan, and Ohio to Aransas Coun- ty, Texas (from Missouri southward only west of the Mississippi River), and were there no populations occurring east of the river, we should not hesitate to conclude that two subspecies of F. fodiens should be recognized. The nominate subspecies (dis- tinguishable almost solely on features of the first pleopod of the first form male in which VOLUME 102, NUMBER 3 the comparatively short central projection usually bears a subapical notch (Fig. 8a—y)) could be considered to occupy the northern sector, extending as far south as northern Arkansas, where throughout the eastern part of the state it intergrades (Figs. 8z—9g) with the more southwestern populations, ranging from southwestern Arkansas and south- eastern Oklahoma southward. These more southern populations exhibit the facies that has been associated with F. hedgpethi (rec- ognized by the possession of a long central projection lacking a subapical notch, Fig. 9h—s). It should be noted that an occasional influence of the fodiens genome surfaces in specimens occurring south of the Arkansas border (see Fig. 97, g). East of the Mississippi River, we fail to find any such regular dis- tribution pattern in the variation of pleo- podal features (Figs. 97-107) except along the Atlantic versant from South Carolina to Maryland (Fig. 10j—x). But the same type pleopod that characterizes those popula- tions occurring along the eastern seaboard may be found in specimens from Alabama and Texas (Fig. 97, t). Thus, we have been unable to discover a single character that serves consistently to distinguish between the formerly recognized F. (C.) fodiens, F. (C.) uhleri, and F. (C.) hedgpethi. The two diagnostic features that Faxon (1884:117) mentioned as setting his Cambarus uhleri apart from C. argillicola (=F. fodiens) were its ““‘plane rostrum [and] shape of the hand. ... Many specimens, particularly those from the Carolinas, have concave rostra, and while we are not certain as to which features in the “‘shape of the hand”’ Faxon was referring, there seems to exist as much variation within specimens from South Car- olina to Maryland as we have noted in in- dividuals from the rest of the range of the species, and we have recognized no feature as being unique. As noted above, the same applies to characteristics pointed out by Hobbs as typifying his ““Cambarus hedg- pethi.” Notes on sex ratio.—In all of the studies 683 of which we are aware that have yielded data on the sex ratios of cambarids except that of Creaser (1934) (e.g., Andrews 1904, Penn 1943, Smith 1953:92, and Smart 1962:94), all have revealed a near 1:1 ratio. Insuffi- cient numbers of individuals of any popu- lation of members of the genus Fallicam- barus in Arkansas have been available that might permit an estimate of the sex ratio in any population, but, in the samples at hand there are many more females than males. Most of our adult specimens of Fallicam- barus (F.) fodiens were removed from bur- rows, and of 293 adults, less than half, only 98, are males. In order to support the belief that our data are little biased, we have re- peatedly attempted to discover some way in which to determine whether a burrow to be excavated contains a male, female, or pair, but we have been unsuccessful. (As will become evident below, the importance of obtaining first form males from throughout the range of the species is paramount.) On 21 April 1973, in a seepage area 0.4 mile east of the Oklahoma State line on USS. Highway 70, in Sevier County, Jean Pugh and HHH removed 23 females from bur- rows before they found a male. In Phillips County, on 17 April 1985, Robert Gilpin and HHH retrieved females from 14 bur- rows in one locality without finding a male, and, at another nearby, they unearthed five females before taking a male. Except for their apparent rarity, we are aware of no evidence that the males of F. (F.) fodiens might be more secretive than are the females; and, we suggest that perhaps two of their habits are responsible for the real or apparent ab- sence of half of them (assuming the sex ratio at hatching 1s near 1:1) from the adult pop- ulation. To a minor extent, perhaps our data are biased, for the burrows of the males seem sometimes to be less elaborate than those of the female, and this is reflected in the often smaller, open, and eroded turrets marking their domiciles. Admittedly, such burrows offer less temptation to the collec- tor, who, for good reason, prefers to explore 684 one over which the turret is capped, or, if open, adorned with comparatively recently deposited, not abraded, pellets. Well-formed soil pellets offer evidence of the presence of a crayfish instead of perhaps a snake (Ag- kistrodon piscivoris or an ill-tempered Ne- rodia sipidon), an Amphiuma, or some other invader. At one time we suspected that of more importance in skewing the apparent sex ratio of adults than failure of collectors to excavate burrows harboring males are the supposed more frequent wanderings of first form males in seeking mates. In their forays from one lair to another, they place them- selves in jeopardy of becoming prey to rac- coons, skunks, owls, and other predators, and indeed scat from owls and raccoons has been observed to contain fragments of the exoskeleton of crayfishes within the range of the species in Arkansas. Thus by their being more frequently exposed to predators than are females, we reasoned that they are being passively selected, and perhaps effec- tively so. What significance, if any, attaches to our having observed more carcasses of first form males than females of this species in areas where their burrows are located es- capes us. We had placed considerable confidence in the above suggestions as possible expla- nations for the apparent skewed sex ratio existing in the adult populations of F. fo- diens until, on 22 April 1988, one of us (HWR) collected 30 juveniles (cl 5.5 to 11.5 mm) of this species from a pool in the ditch at the locality mentioned above where Pugh and Hobbs had collected in 1973. Twenty- nine of the specimens were females! The question remains as to whether or not the sex ratio at hatching in this species is 1:1, and if it is what factor/s (cannibalism of the perhaps slower growing juvenile males—the male is the smallest of the 30 juveniles) are responsible for the subsequent alteration of the ratio. That our suppositions might well be suspect seem possible when we remem- bered Creaser’s (1934) study revealing pos- sible seasonal changes in populations of the PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON lake- and stream-dwelling Faxonius (=Or- conectes) propinquus in which from August to January 66 to 73 percent of the popula- tion was female; during the rest of the year the percentage ranged from 43 to 56 per- cent. Creaser was apparently as puzzled by these data as are we by the information we have on F. (C.) fodiens. Fallicambarus (Creaserinus) gilpini, new species Figs. 1g, 5, 11 Diagnosis.—Cheliped with sufflamen; ventral surface of merus with mesial row of tubercles, lateral one never represented by more than two; length of carpus less than or subequal to width of palm of chela. Chela with lateral margin strongly costate, never serrate, dorsal surface lacking scattered tu- bercles in lateral half, ventrolateral surface with arched row of prominent punctations bearing long setae, opposable margin of dac- tyl with distinct excision 1n basal half, me- sial margin without tubercles. Mesial sur- face of palm of second pereiopod lacking conspicuous tufts of plumose setae. First pleopod lacking proximomesial spur, and lacking cephalic process; central projection weakly arched, its base not inclined later- ally, its distal part directed caudoproximally with well defined subapical notch, never crossing central projection of corresponding pleopod. Hooks present on ischia of third pereiopods only. Boss on coxa of fourth pe- reiopod somewhat rounded, not distinctly compressed. Mesial ramus of uropod with distolateral spine; distomedian spine pre- marginal. Telson incompletely divided, with spine on anterolateral flank of suture. Holotypic male, form I.—Eyes small but pigmented and with faceted cornea. Body subcylindrical, very weakly compressed (Figs. 4b, 11a, 1). Abdomen distinctly nar- rower than thorax (7.3 and 10.1 mm). Greatest width of carapace at level about one-third length of areola from cervical groove where subequal to height (10.1 and VOLUME 102, NUMBER 3 685 Fig. 11. Fallicambarus (Creaserinus) gilpini (all from holotype except c, e, from morphotype, and i, n from allotype): a, Lateral view of carapace; b, c, Mesial view of first pleopod; d, Dorsal view of caudal part of abdomen; e, f, Lateral view of first pleopod; g, Postaxial view of mandible; h, Caudal view of first pleopods: i, Annulus ventralis and associated sclerites; j}, Antennal scale; k, Epistome; 1, Dorsal view of carapace; n, Basal podomeres of third, fourth, and fifth pereiopods; n, 0, Distal podomeres of cheliped. 686 10.0 mm). Areola linear over most of length and comprising 40.5 percent of entire length of carapace (46.6 percent of postorbital car- apace length). Rostrum with convergent, slender margins gently contracted anterior- ly, marking base of poorly delimited acu- men, apex corneous, slightly upturned, and reaching base of ultimate podomere of an- tennular peduncle. Dorsal surface of ros- trum concave with submarginal rows of se- tiferous punctations and others between, especially dense and conspicuous in basal part. Subrostral ridges weak but evident in dorsal aspect to base of acumen. Postorbital ridges slender but well defined and merging almost imperceptibly with carapace above posterior margin of orbit. Branchiostegal and cervical spines absent. Suborbital angle ab- sent, cephalolateral margin of carapace sloping caudoventrally from base of ros- trum without excrescence or excision. Car- apace comparatively densely punctate dor- sally and laterally; row of few small tubercles flanking anterolateral segment of cervical groove. Abdomen (Fig. 45) shorter than carapace (19.8 and 22.0 mm); pleura small and broadly rounded ventrally, none with an- gular caudoventral angle; pleuron of first segment clearly overlapped by that of sec- ond. Telson (Fig. 11d) not divided but deep- ly incised laterally and caudolateral angles of cephalic section bearing two pairs of spines, more mesial pair smaller and mov- able. Proximal podomere of uropod with mesial lobe bearing acute angle, lateral one rounded; mesial ramus with distolateral spine and smaller premarginal distomedian spine. Cephalomedian lobe of epistome (Fig. 11k) subtriangular with cephalomedian prominence; cephalolateral margins elevat- ed ventrally and only slightly undulant; main body of epistome depressed but lacking fo- vea. Ventral surface of proximal podomere of antennule lacking spine. Antennal pe- duncle without spines, flagellum falling short of caudodorsal margin of carapace. Anten- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON nal scale (Fig. 11/) small, just reaching base of acumen and penultimate podomere of antennular peduncle; lamella broadly rounded distomesially, broadest distal to midlength, but only slightly wider than thickened lateral part. Mandible (Fig. 11g) with cornified subtriangular area of caudal molar process proportionately much small- er than that of F. (F.) devastator and farther removed from corneous tuberculiform ce- phalic molar process. Ventral surface of is- chium of third maxilliped with lateral row of short, plumose setae, and mesial half bearing clusters of longer stiff setae; basis with conspicuous cluster of long setae ob- scuring proximal part of ischium. Right chela (Fig. 110) about 2.2 times as long as broad, rather strongly depressed; width of palm about 1.7 times length of mesial margin, latter bearing row of seven tubercles and one prominent one lying slightly dorsal to row between third and fourth from proximal end; irregular row of six much smaller tubercles on dorsal flank of mesial row; dorsal surface of palm and fingers bearing setiferous punctations, those on and adjacent to base of fixed finger and on proximal half of dactyl conspicuous; lat- eral margin of chela rounded proximally, but largely costate; ventral surface punctate except for single prominent tubercle oppo- site base of dactyl, ventrolateral arc of punc- tations, each made prominent by long stiff seta, extending from base of palm to base of distal third of fixed finger. Both fingers with well defined submedian ridge flanked by punctations dorsally; ridges on ventral surface less well defined. Opposable margin of fixed finger with row of four tubercles (third from base largest) along proximal third and one projecting from lower level slightly distal to midlength; single row of minute denticles extending from third tubercle from base to corneous tip of finger. Opposable margin of dactyl with obvious excavation proximally, two tubercles borne on margin of excavation and one marking its distal ex- tremity, all subequal in size; single row of VOLUME 102, NUMBER 3 minute denticles extending from distalmost tubercle to corneous tip of finger. Mesial margin of dactyl bearing setiferous punc- tations, lacking even basal tubercles. Carpus of cheliped about 1.4 times as long as broad and approximately 1.7 times as long as mesial margin of palm. Dorsal sur- face with deep submedian longitudinal sul- cus flanked by setiferous punctations; me- sial surface of podomere tuberculate, that on distal margin acute and much larger than more proximal ones; lateral and ventral sur- faces punctate; ventrodistal margin with two acute, corneous tubercles: one on ventro- lateral condyle and other mesial to it. Merus with few squamous to rounded tubercles near dorsodistal extremity, two somewhat larger than others; mesial and lateral sur- faces finely punctate; ventral surface with mesial row of eight tubercles (nine on left); usual lateral row absent but single tubercle present on left member; lateral row of tu- bercles characteristic of most crayfishes ab- sent; longitudinal row of long, stiff setae present. Ischium punctate, lacking tubercles ventromesially. Chela of second pereiopod with marginal row of setae on palm, and carpus with dorsal row of long setae; mesial surface of carpus and propodus lacking tufts of plumose setae. Ischium of third pereiopod only with hook (Fig. 11m); latter simple, not overreaching basioischial articulation, and not opposed by tubercle on corresponding basis. Coxa of fourth pereiopod with knoblike caudome- sial boss which, if leg extended laterally, projecting mesially. Coxa of fifth pereiopod devoid of boss, ventral membrane setifer- ous. First pleopods (Fig. 11b, f h) reaching coxae of third pereiopods and largely hid- den by setae extending mesially from ven- tral margin of sternum. When abdomen flexed, however, apices of terminal ele- ments protruding ventrally beyond setal curtain. Proximomesial spur lacking. Shaft of appendage only slightly bowed, with two terminal elements disposed caudally at about 687 90 degrees. Mesial process somewhat spat- ulate, shallowly chamfered, and tilted lat- erally. Corneous central projection blade- like, with distinct subapical notch, arched and reaching caudally to about same level as mesial process. Allotypic female.— Differing from holo- type in other than secondary sexual char- acters as follows: acumen not quite reaching base of ultimate podomere of antennular peduncle; postorbital ridges not reaching level of posterior margin of orbit. Abdomen subequal in length to carapace. Distolateral spine on mesial ramus of uropod greatly reduced (perhaps abraded), that on right represented by no more than angle; because of encrustation, dorsal surface of telson and uropods appearing much more strongly se- tose than that of holotype; both lobes of proximal podomere of uropod rounded. Antennal scale slightly overreaching acu- men and base of penultimate podomere of antennular peduncle. Mandible with ce- phalic molar process strongly abraded, cau- dal molar process lacking corneous ele- ments. Mesial margin of palm of right chela (Fig. 117) with row of 5 tubercles (left with 6) and row of 4 on dorsal flank (left lacking second row but with single tubercle, be- tween fourth and fifth, ventral to row); dacytl with 2 small tubercles distal to 3 associated with proximal excavation; ischium with ventromesial row of 12 tubercles (11 on left). (See Table 2 for mensural features.) Annulus ventralis (Fig. 117), 1.5 times as broad as long, situated deeply in sternum; cephalic region immovable, but caudal two- thirds capable of hingelike motion. Sulcus shallow and narrow cephalically, becoming deeper and broader caudosinistrally; high, prominent caudal wall cut by C-shaped si- nus arising from fossa at caudosinistral side of sinistrally projecting tongue. Postannular sclerite less than half as long and approxi- mately half as wide as annulus with punc- tate, oval, ventrally elevated median area. Morphotypic male, form II.—Differing from holotype in following respects: Tip of 688 Table 2.— Measurements (mm) of Fallicambarus (F.) gilpini. Morpho- Holotype Allotype type Carapace: Entire length 22.0 24.9 28.3 Postorbital length 9et DAT PL 5).8) Width LO 11.6 13.0 Length 10.0 10.9 be Areola: Width — — — Length 8.9 10.0 11.4 Rostrum: Width 35 35 4.2 Length 321 Sit 3.9 Right chela: Length, palm mesial margin ee 323 4.8 Palm width 5.4 6.5 7.4 Length, lateral margin 11.8 (2-7 12 Dactyl length 8.1 8.5 10.1 Abdomen: Width os | 93 Length 19.8 25a 25.0 rostrum abraded but acumen reaching mid- length of penultimate podomere of anten- nular peduncle. Postorbital ridges termi- nating slightly posterior to caudal margin of orbit; lateral surface of branchiostegites granular; telson divided; both lobes of prox- imal podomere of uropod rounded; main body of epistome with cephalomedian fo- vea; left antennal scale as in allotype, right with regenerated distolateral spine; right chela with only five tubercles in row on dor- sal flank of mesialmost row; opposable mar- gin of fingers armed as in allotype; oppos- able margin of dactyl of left chela with seven tubercles, four associated with excavation and three distal to it. Except for terminal elements of first pleo- pod (Fig. 1 1c, e), no noteworthy differences noted between morphotype and holotype. Mesial process more robust with less con- spicuous groove, but projecting caudally much beyond tip of central projection; lat- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ter, in addition to being stouter and non- corneous, also lacking subapical notch. Color notes.—Basic color pale greenish blue, but cephalic region more lavender than blue; rostral margins and postorbital ridges distinctly dark green. Mandibular adductor and posterior part of gastric area lavender with faint greenish suffusion. Lateral ce- phalic region fading to white ventrally, but with paired small navy blue spots midway between tip of rostrum and caudal extrem- ity of cervical groove and another less well defined pair abutting cervical groove. Tho- racic area dark greenish blue dorsally, sud- denly changing to white laterally, and at least half of branchiostegite white. First abdom- inal tergum dark greenish blue, second slightly paler, and third through sixth yet more pale with faint hint of very pale dor- somedian longitudinal stripe; white margin of all pleura partly separated from blue to bluish green tergum by series of short dark greenish blue bars. Telson with cephalic sec- tion mostly bluish, caudal section colorless and translucent; lateral section of lateral ra- mus of uropod pale bluish green, keels of both rami and margins of basal podomere dark greenish blue. Antennules and anten- nae with lateral and mesial borders of pe- duncles darker blue than dorsal and ventral surfaces; flagella greenish blue basally fad- ing to pale tan. Cheliped with dorsal surface of distal part of merus, carpus and chela dark bluish green; tips of fingers yellowish with corneous brown tip; ventrolateral part of palm and fixed finger fading to cream; ridges and tubercles on carpus and palm of chela very dark blue. Venter and basal pod- omeres of remaining pereiopods white; dor- sum of merus, carpus, and propodus of sec- ond through fifth pereiopods bluish green, dactyl with some blue but more cream to yellowish. Size.— The largest specimen available is a second form male having a carapace length of 28.8 (postorbital carapace length, 26.0) mm. Corresponding measurements of the smallest first form male were 21.1 and 18.8 VOLUME 102, NUMBER 3 mm, and those of the smallest ovigerous female, the allotype, are 24.9 and 21.7 mm, respectively. Type locality.—Roadside seepage 3.1 mi south of southern junction of State Route 54 and U.S. Highway 79 at junction of latter with Pepperridge Road (T7S, R1OW, Sec 19), approximately 11 miles south of Pine Bluff and about 3 miles north of Cleveland County line, Jefferson County, Arkansas. Disposition of types.—The holotype, al- lotype, and morphotype (USNM 219511, 219512, and 218944, respectively) are de- posited in the National Museum of Natural History, Smithsonian Institution, as are the paratypes consisting of 1 46 I, 1 6 II, 8 9, 2 juv 6, 2 juv 2, 2 ovigerous 2. Range and specimens examined.— All of the specimens were collected from burrows in roadside seepages in Jefferson County, Arkansas: (1) type locality, 1 6 II, 26 Apr 1986, HWR; 5 2, 2 juv 4, 2 juv 2, 18 Mar 1987, HWR (two additional juvenile males were maintained in aquaria until they molt- ed to first form, one in late Feb 1988, and the other, the holotype, on 9 or 10 Apr 1988). (2) 0.2 mi S of Pine Bluff city limits, 1 2, 11 Apr 1986, HWR, coll. (3) 3.6 mi N of Cleve- land Co. line on US Hwy 79, 2 2, 7 Nov 1987, B. F. Kensley, HWR, HHH; 2 ¢ II, 2 2, 3 ovig 2, 11 Mar 1988, HWR. Variations.— Among the adult speci- mens, the areola constitutes from 38.1 to 40.6 percent of the total length of the car- apace, and from 42.1 to 46.6 percent of the postorbital carapace length. Most of the dif- ferences noted in the specimens may be at- tributed to abrasion and to regeneration of chelipeds. The former is reflected in the ab- sence of an upturned tip on the rostrum, reduced and/or truncated tubercles and spines, and broken or missing setae (partic- ularly noticeable on the third maxillipeds of specimens in late intermolt stages). Re- generated chelipeds can usually be recog- nized by the absence or reduction of the excavation on the proximal part of oppos- able margin of the dactyl of the chela, but 689 also by the smaller, and frequent increase in number of tubercles on the opposable margins of both fingers. The number of tu- bercles in the ventromesial row on the me- rus of the cheliped ranges from 7 to 11, on the mesial margin of the palm of the chela, from 5 to 7 with 0-6 on the dorsal flank; on the opposable margin of both fingers, there are from 4 to 6 tubercles (in one specimen 2, instead of 1, lie at a lower level on the fixed finger). There is no noteworthy vari- ation in the secondary sexual features of the two available first form males, and in the female, the only conspicuous difference ob- served is the anticipated occurrence of a mirrored image of features of the annulus described in the allotype. Life history notes.—No first form male has been collected, and the only two avail- able were reared from very small juveniles collected in March 1987. They were main- tained in the laboratory in Washington, D.C., and molted to first form in late Feb- ruary and early April 1988. Three ovigerous females were dug from burrows on 11 March 1988: one with carapace length of 22.3 mm carried 18 eggs along with several empty ““capsules,’’ another with cl of 24.9 mm, 20 eggs, and the third with cl of 25.5 mm, 35 eggs. All of the eggs were about 2 mm in diameter. The two juvenile females collect- ed on 18 March 1987 have carapace lengths of 11.9 and 12.1 mm, and the two juvenile males, 11.4 and 11.9 mm. Ecological notes.— This crayfish has been found only in complex burrows consisting of branching galleries, several of which, ex- cept in dry seasons, reach the surface, some of their openings marked by rather crudely constructed turrets. Where Fallicambarus (C.) gilpini has been collected in the same locality with F. (C.) fodiens, the burrows of the former were frequently, if not usually, situated higher on the seepage slope, sug- gesting that like the partitioning of a habitat in South Carolina by Cambarus (J.) caro- linus (Erichson, 1846) and Distocambarus (Fitzcambarus) carlsoni Hobbs, 1983 (see 690 latter, page 437), gi/pini might prefer areas in which the groundwater is moving, where- as fodiens more frequently occurs in areas in which the water is more static. In general, the burrows of gi/pini are more complex, exhibiting more subhorizontal galleries than do those of fodiens that we have excavated in Arkansas. Relationships.— Fallicambarus (C.) gil- pini has its closest affinities with F. (C.) cae- sius. In addition to the many features the two species share in common, including being the only typically blue members of the genus, they are the only ones that lack a ventrolateral row of tubercles on the me- rus of the first cheliped. The most readily observed features that distinguish the two species are the absence of tubercles on the mesial surface of the dactyl of the chela and the presence of a distolateral spine on the mesial ramus of the uropod in F. (C.) gi/pini. Etymology.—This crayfish is named in honor of our mutual friend Robert H. Gil- pin, of Cumberland, Maryland, in token of his interest and assistance in collecting much of the material we have from the eastern part of Arkansas. Acknowledgments For their assistance in collecting some of the crayfishes on which this study is based, thanks are extended to Robert H. Gilpin of Cumberland, Maryland; Michael F. Kear- ney of Louisiana State University, Brian F. Kensley of the Smithsonian Institution: Raymond F. Jezerinac of Ohio State Uni- versity, Newark; M. James Norrocky of Vickery, Ohio; John Dempsey of Arkansas High School, and the following students at Southern Arkansas University: Wanda Hobson, Daryl Koym, Elaine Laird, Beth Lovorn, Patrick Robison, and Linda Tate. For the loan of specimens from the Illinois Natural History Survey, we are grateful to Lawrence M. Page. For their criticisms of the manuscript we are indebted to Thomas E. Bowman of the Smithsonian Institution, PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Joseph F. Fitzpatrick, Jr., of the University of South Alabama, and H. H. Hobbs III, Wittenberg University. The Southern Ar- kansas University Faculty Research Fund provided travel funds to H. W. Robison to collect crayfishes. Literature Cited Albaugh, Douglas W. 1973. Life histories of the cray- fishes Procambarus acutus and Procambarus hinei in Texas. 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Comparison of burrows and burrowing behavior of five species of cambarid crawfish (Crustacea, Decapoda) from the Southern Uni- versity campus, Baton Rouge, Louisiana. — Pro- ceedings of the Louisiana Academy of Sciences 48:23-29. Smart, GroverC. 1962. The life history ofthe crayfish Cambarus longulus longulus.— American Mid- land Naturalist 68(1):83-94. Smith, Elsie Wayne. 1953. The life history of the crawfish Orconectes (Faxonella) clypeatus (Hay) (Decapoda, Astacidae).— Tulane Studies in Zo- ology 1(7):79-96. Spoor, W. A. 1955. Loss and gain of heat-tolerance by the crayfish. — Biological Bulletin 108(1):77— 87. Stebbing, Thomas R. R. M. A. 1893. A history of Crustacea, Recent Malacostraca. D. Appleton & Company, New York, xvii + 466 pp. Terman, Max R. 1974. The ecology of crayfish: Some observations on behavioral factors.—The Bi- ologist 56(1):32-39. Thoma, Roger F., & Raymond F. Jezerinac. 1982. New distributional records of crayfishes (Cam- barus and Fallicambarus) from Ohio including a new subspecies record.— Ohio Journal of Sci- ence 82(3):136-138. Turner, Clarence L. 1926. The crayfishes of Ohio. — Ohio Biological Survey Bulletin 3(3):145-195. Underwood, Lucien M. 1886. List of the described species of fresh water Crustacea from America, VOLUME 102, NUMBER 3 north of Mexico. —Bulletin of the Illinois State Laboratory of Natural History 2(5):323-386. Walls, Jerry G. 1968. A new Faxonella from north- east Louisiana (Decapoda, Astacidae).—Pro- ceedings of the Biological Society of Washington 81(41):413-418. . 1985. Distribution and natural history of the crawfish Orconectes difficilis (Decapoda: Asta- cidae) in Louisiana.—Southwestern Naturalist 30(2):189-194. , & Joe B. Black. 1967. New variations and records of the crawfish Procambarus jaculus Hobbs (Crustacea, Decapoda, Astacidae).— Proceedings of the Louisiana Academy of Sci- ences 30:60-62. Washburn, Mel. 1953. Mudbugging.—Louisiana Conservationist 13(3):5-7, 19. Waywell, E. B., & S. Corey. 1970. The presence of pteridines in the hypodermis as a taxonomic tool in crayfish.— Canadian Journal of Zoology 48:1462-1464. 1972. The occurrence and distribution of pteridines and purines in crayfish.—Crusta- ceana 22(3):294—302. Wharton, Charles H. 1978. The natural environ- ments of Georgia. Atlanta, Georgia: Depart- ment of Natural Resources, 227 pp. Wiens, Wayne, & Kenneth B. Armitage. LOGI, The 697 oxygen consumption of the crayfish Orconectes immunis and Orconectes nais in response to temperature and oxygen saturation.—Physio- logical Zoology 34:39-54. Williams, Austin B. 1954. Speciation and distribu- tion of the crayfishes of the Ozark Plateaus and Ouachita Provinces. — University of Kansas Sci- ence Bulletin 36(12):803-918. Williams, D. Dudley, N. E. Williams, & H. B. N. Hynes. 1974. Observations on the life history and bur- row construction of the crayfish Cambarus fod- iens (Cottle) in a temporary stream in southern Ontario.—Canadian Journal of Zoology 52(3): 365-370. Williamson, E. B. 1899. Notes on Ohio Astacidae. — Seventh Annual Report of the Ohio State Acad- emy of Science, pp. 47-48. . 1907. Notes on the crayfish of Wells County, Indiana, with description of new species. — 31st Annual Report of the Department of Geology and Natural Resources, Indiana 1906:749-763. (HHH) Department of Invertebrate Zo- ology, Smithsonian Institution, Washing- ton, D.C. 20560; (HWR) Department of Bi- ology, Southern Arkansas University, Magnolia, Arkansas 71753. PROC. BIOL. SOC. WASH. 102(3), 1989, pp. 698-700 SANQUERUS, A REPLACEMENT NAME FOR POSIDON HERKLOTS, 1851 (CRUSTACEA, DECAPODA, PORTUNIDAE) Raymond B. Manning Abstract. — Sanquerus is proposed as a replacement name for the preoccupied portunid genus Posidon Herklots, 1851, and the name is removed from the synonymy of Portunus Weber, 1795. Sanquerus is a monotypic genus contain- ing only the West African Sanquerus validus (Herklots, 1851). Manning & Holthuis (1981:104, 105) pointed out that the West African “Portu- nus validus shows little similarity to any of the many Indo-West Pacific species of the genus, and it shows little affinity with any of the American species of the genus... .” They also enumerated differences between Portunus validus and Portunus pelagicus (Linnaeus, 1758), the type species of Por- tunus. I take this opportunity to remove Posidon from the synonymy of Portunus and to propose a replacement name for it. Sanquerus, new name Figs. 1-2 Posidon Herklots, 1851:3. [Invalid junior homonym of Posidon Iliger, 1801 (Crus- tacea). Type species: Portunus (Posidon) validus Herklots, 1851, by monotypy. Gender masculine.] Diagnosis. —Size very large, carapace widths in adults exceeding 20 centimeters. Carapace wide, breadth about two times Fig? 1: Sanquerus validus (Herklots) (from Milne Edwards 1861, pl. 29, fig. 1). VOLUME 102, NUMBER 3 699 Fig. 2. Sanquerus validus (Herklots). a, front; b, third maxilliped; c, merus and carpus of cheliped: d, fifth leg; e, abdomen; f, gonopod; g, apex of gonopod. (a, b, d—g from male, 151 mm wide, Nigeria, USNM 121034; c from male, 61 mm wide, Liberia, USNM 97864) (c, f, g from Manning & Holthuis 1981). length. Surface of carapace minutely tuber- culate, appearing smooth to the naked eye, evenly convex, lacking distinct grooves or ridges, except for ridge extending across posterior margin between bases of last walking legs; posterolateral angles of cara- pace unarmed; conspicuous white spot pres- ent posterolaterally on each side. Front with three pairs of spiniform teeth, including in- ner orbitals, median pair largest. Interan- tennular projection low, not visible in dor- sal view. Anterolateral margin of carapace with nine spiniform teeth, lateralmost larg- est. Antenna with free access to orbit. Palate with longitudinal ridge. Merus of third max- illiped lacking produced anterior lobe. Che- liped robust; merus with two posterior spines, one subdistal, three inner spines, and one smaller vental spine distally; carpus with inner and outer spine; palm prismatic and costate, with proximal spine at articulation with wrist, distal dorsal spine, and smaller distal spine on inner carina. Merus of fifth leg with posterodistal margin unarmed, rounded; swimming paddle notched distal- ly. Abdomen of male triangular, 5-seg- mented, third to fifth segments fused; ter- minal segment longer than broad. Male pleopod stout, curved laterally, unarmed. Etymology. — This name 1s in recognition of the enormous contribution made to the knowledge of the West African fauna by Mr. Robert Sanquer of Vouhé, France, former Trawling Master for the Guinean Trawling Survey. The gender is masculine. Remarks. —Sanquerus is a very distinc- tive portunid, easily distinguished from all known portunid genera by the smooth, unornamented carapace in combination with the prismatic and costate chelae. It keys to Portunus in Rathbun (1930:13), Crosnier (1962:34), Garth & Stephenson (1966:10), 700 and to the couplet containing Portunus and Scylla in Stephenson (1972:8). Sanquerus validus keys to Portunus pe- lagicus in Stephenson & Campbell (1959: 90) and to the couplet containing P. pelagi- cus in Crosnier (1962:42). It keys to Por- tunus convexus de Haan, 1833 in Stephen- son (1972:13) because of the paired white spots on the carapace, but that species has distinct ornamentation on the carapace and only one posterior spine on the merus of the cheliped. On morphological grounds it keys to the couplet containing Portunus pelagicus in Stephenson (1972:15). It differs from Portunus pelagicus in numerous features, some of which were mentioned by Manning & Holthuis (1981:105): the carapace lacks surface sculpture and is minutely rather than distinctly tuberculate; the submedian fron- tal teeth are the largest of the frontal teeth; the interantennular spine is not visible in dorsal view; the third maxilliped extends far less forward; the cheliped is more massive, and the merus 1s ornamented with two strong posterior spines; the abdomen of the male is broader; and the gonopod is much stout- er. As in P. pelagicus, the dactylus of the fifth leg is notched distally, not produced into a spine, and the anterior eight antero- lateral teeth of the carapace are not alter- nately large and small. Sanquerus resembles the Indo-West Pa- cific genus Scy//la de Haan, 1833 in having a relatively smooth carapace, but differs from Scy/la in having the chelae prismatic and costate; in Scy//a the chelae are robust and smooth (Stephenson & Campbell 1960: 111; Stephenson 1972:8, 44). In frontal view, the carapace of Sanquerus shows very low paired swollen prominences on the protogastric, mesogastric, and me- sobranchial regions, and distinct but small paired branchial lobes. Sanquerus validus is a commercial species occurring off the West African coast, from PROCEEDINGS OF THE BIOLOGICAL SOCIETY CF WASHINGTON Senegal to Angola. References to the species may be found in Monod (1956:196) and Manning & Holthuis (1981:103). Acknowledgments I thank Fenner A. Chace, Jr., and Austin Williams for reviewing the manuscript. Literature Cited Crosnier, A. 1962. Crustacés Décapodes, Portuni- dae.—Faune de Madagascar 16:1—-154, pls. 1- 13: Garth, J. S., & W. Stephenson. 1966. Brachyura of the Pacific coast of America, Brachyrhyncha: Portunidae.—Allan Hancock Monographs in Marine Biology 1:1-154. Herklots, J. A. 1851. Additamenta ad Faunam Car- cinologicam Africae occidentalis, sive descrip- tiones specierum novarum e crustaceorum or- dine, quas in Guinea collegit vir strenuus H.S. Pel, praefectus residentiis in littore guineae. Lugduni-Batavorum, Leiden, 28 pp., 2 pls. Manning, R. B., & L. B. Holthuis. 1981. West African brachyuran crabs. —Smithsonian Contributions to Zoology 306:x1i + 379 pp. Monod, Th. 1956. Hippidea et Brachyura ouest-af- ricains.— Mémoires de I’Institut frangais d’Af- rique noire 45:1-674. Rathbun, M. J. 1930. The cancroid crabs of America of the families Euryalidae, Portunidae, Atele- cyclidae, Cancridae and Xanthidae.— United States National Museum Bulletin 152:xvi + 609 pp., pls. 1-230. Stephenson, W. 1972. An annotated check list and key to the Indo-West-Pacific Swimming Crabs (Crustacea: Decapoda: Portunidae).— Royal So- ciety of New Zealand, Bulletin 10:1-64. Stephenson, W., & B. Campbell. 1959. The genus Portunus. The Australian portunids (Crustacea: Portunidae), IIJ.— Australian Journal of Marine and Freshwater Research 10(1):84—124, pls. 1-5. =. 2 . 1960. Remaining genera. The Aus- tralian portunids (Crustacea: Portunidae), IV.— Australian Journal of Marine and Freshwater Research 11(1):73-122, pls. 1-6. Department of Invertebrate Zoology, Na- tional Museum of Natural History, Smith- sonian Institution, Washington, D.C. 20560. PROC. BIOL. SOC. WASH. 102(3), 1989, pp. 701-715 RECTIFICATION OF HALIRAGES REGIS AND H. HUXLEYANUS (CRUSTACEA: AMPHIPODA), FROM MARINE ANTARTICA, WITH DESCRIPTION OF A NEW GENUS, AUSTROREGIA J. L. Barnard Abstract. — Halirages huxleyanus and H. regis are removed to the new ant- arctic genus, Austroregia. This leaves Halirages with 6 species entirely confined to arctic waters. Austroregia is a perplexing genus because it is furnished with calceoli of the same kind found in Chosroes and Gammarellus; a family Gam- marellidae is available for these two genera which would be removed from such families as Eusiridae, Pontogeneiidae and Calliopiidae, in which the genera would have been classified previously. Problems remain on speciation within Austroregia. Halirages huxleyanus (Bate) and H. regis (Stebbing) are improperly placed in Halira- ges and are removed to a new genus Aus- troregia. Austroregia huxleyana possesses distinctive calceoli of a kind also present in Gammarellus, Chosroes and Gondogeneia, adequate to justify the resurrection of the family Gammarellidae within the super- family Eusiroidea. This reinforces the dis- covery by Lincoln & Hurley (1981:111) that both the high arctic and high antarctic con- tain taxa with common ancestry in the gam- marellid group, a family first raised by Bousfield (1977), but soon after merged within the family Calliopiidae. Gammarellidae, revived Diagnosis. —Characterized by a type 6 calceolus of Lincoln & Hurley (1981) in which the proximal element forms a dis- crete cup separated from the small 2 to 3-plate distal element by a second smaller cup-shaped element (Fig. 1). Also unique is the arrangement of the calceoli in transverse rows that extend all around the distal mar- gin of the flagellar articles. Remarks.—The family Gammarellidae was established by Bousfield (1977) to con- tain two carinate genera, Gammarellus Herbst and Weyprechtia Stuxberg, separat- ed from other kinds of pontogeneiids by a combination of characters that included a well developed accessory flagellum, lanceo- late weakly setose third uropods, and lam- inar, apically emarginate telson. However, the distinction from other eusiroids was far from clear-cut and in a later updated and revised version of his classification, Bous- field (1983) synonymized the Gammarelli- dae with the Calliopiidae. The present use of calceolus morphology and arrangement as a shared apomorphy to resurrect the Gammarellidae produces a family of quite a different complexion. Brought together are four eusiroid genera, Gammarellus, Gondogeneia, Chosroes, and Austroregia, that would not have been rec- ognized as belonging to a natural and in- dependent group on the basis of traditional morphological characterizations. Thus, Gammarellus possesses a well developed multiarticulate accessory flagellum which is at best small to vestigial in Gondogeneia and 702 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Calceoli: a, Austroregia huxleyana, holotype “a”; b, Chosroes decoratus. Magnifications unknown; courtesy of Dr. Roger J. Lincoln. Austroregia and absent in Chosroes. Fur- Chosroesand Gammarellus is weakly emar- ther, Gammarellus is the only member hav-__ ginate, but has a short to moderate cleft in ing a coxal gill on pereonite 7, a lobate car- Gondogeneia and Austroregia. A lanceolate pus on gnathopods 1-2, and facial setae on _setose uropod 3 is shared by all four genera the inner plate of maxilla 2. The telson of but is also widespread outside the group. VOLUME 102, NUMBER 3 The single feature uniting the family is the type 6 calceolus. This could be treated as yet another instance of convergence, as has to be argued for many eusiroid characters, but in view of the marked complexity of the calceolus sensory receptor and its novel dis- tribution on the antennal articles, there is adequate justification for recognizing the group as a separate unit within the Eusi- roidea. There is a superficial resemblance of the gammarellid group to the Pleustidae, but the latter family has distinctive labia bear- ing outer lobes tilted across fused inner lobes; pleustids lack calceoli and further relation- ships cannot be struck. The lower lip of Me- sopleustes and Chosroes bridges the gap be- tween the two groups. Families based entirely on calceolar structure are very difficult to handle by tax- onomists because many species and genera that obviously are analogous to the calceo- liferous members of various families have lost their calceoli. This has been a major problem with Crangonyctidae and will be a problem with Gammarellidae. Only 18 out of 91 families of Gammaridea have one or more species with calceoli. Fortunately, 16 out of the 18 families can be recognized by characters other than calceoli. At best the two families here mentioned are “ghost families” for the identificatory taxonomist because not all of their species and genera are recognizable by ordinary morphological characters. Just as with the Crangonyctidae, there may be species lacking calceoli which are descendants of an ancestor common to the known members of Gammarellidae. For the moment, those species, plus all other members of Eusiridae, Calliopiidae and Pontogenelidae have to remain in a pool that is identifiable through laborious keys that contain little relationship to the phy- logenetic positions of their genera. This is another case of the difference between prac- tical classification involving the identifica- tion of species and genera versus phyloge- netic classification where species and genera 703 may be arranged on the basis of characters not readily available from preserved mu- seum specimens. Austroregia, new genus Type species.—Atylus huxleyanus Bate, 1862, here selected. Etymology.—From ‘“‘austro,” southern, and “‘regia,”’ kingly. Diagnosis. —Eusiroid-like amphipods with type 6 calceolus; body carinate; ros- trum small; eyes circular; basal article of flagellum on antenna | not elongate; anten- na 2 as long as antenna 1; upper lip entire; inner lobes of lower lip absent; inner plate of maxilla 1 with only six or fewer setae, mostly terminal; palps disymmetrical or not (type); inner plate of maxilla 2 with only medial setae, none facial; outer plate of maxilliped of basic form and size; gnatho- pods small, carpus shorter than propodus, nonlobate, gnathopod | slightly larger than 2; pereopods 3—4 with nonlobate basis; coxa 4 excavate or not posteriorly; gills unpleat- ed, gill 7 lacking; at least pereopod 7 elon- gate; outer rami of uropods 1-2 shortened; peduncle of uropod 3 slightly elongate, rami extending equally, poorly setose; telson elongate, cleft about 10-20 percent of its length. Composition. — Austroregia huxleyana (Bate, 1862) and A. regis (Stebbing, 1914). Relationship. —Dijffers from the confa- milial genera, Gondogeneia in the presence of large dorsal carinae on pereonite 7 and pleonites 1—3; Chosroes in the enlarged gnathopod 1, broader outer plate of the maxilliped and the lack of an anterior lobe on the basis of pereopods 3-4; Gamma- rellus in the nonlobate carpal articles of gnathopods 1-2, lack of coxal gill 7, vestigial accessory flagellum, nonpleated gills, ab- sence of facial setae on the inner plate of maxilla 2, poorly setose inner plate of max- illa 1, and cleft telson. In addition to the type 6 calceolus, Aus- troregia differs from the eusiroid genera, 704 Halirages in the fewer setae on the inner plate of maxilla 1, the lack of facial setae on maxilla 2, and the enlarged gnathopod 1; Cleippides in the short carpus of the gnatho- pods; Haliragoides in the lack of inner lobes on the lower lip, the fewer and nonfacial setae of maxillae 1-2 and the regular sized uropod 3; Whangarusa Barnard & Kara- man (1987), in the carinate body, poorly setose maxillae, and lack of inner lobes on the lower lip; Paracalliopiella in the elon- gate telson, and nonreniform eyes; C/eonar- dopsis in the lack of inner lobes on the lower lip, and lack of carpal lobes on the gnatho- pods; Harcledo in the carinate body, longer anterior coxae and weakly cleft telson; Amathillopsis in the vestigial accessory fla- gellum, nonlobed carpus of the gnathopods, slightly enlarged gnathopod 1, fewer setae on the inner plate of maxilla 1 and the smaller dactyl of the maxilliped; Meteusi- roides in the carinate body, nonreniform eyes, nonelongate article 1 on the flagellum of antenna 1, nonlobate carpus of the gnathopods, and poorly cleft telson; from Paramphithoe in the broader articles 5—6 of the gnathopods, especially the more inflated article 6, the nonbifid coxa 4, the enlarged gnathopod 1, the confinement of body teeth to the dorsal midline and the fewer setae on the inner plate of maxilla 1; from Ponto- geneia in distinctive calceoli, poorly cleft telson, absence of inner lobes on the labium, enlarged setae on inner plate of maxilla 2 which are fully marginal (versus facial) and the bluntly rounded anteroventral margins of the anterior coxae; and finally, Ca/liopius, in the nonlobate carpus of the gnathopods, nonreniform eyes, absence of inner lobes on the lower lip, and the distinctive calceoli. Coxa 4 takes different forms. In juveniles it is usually excavate weakly and because in some adults it curls outward it seemingly is deeply excavate when illustrated without flattening. In some large adults this coxa assumes a diamond-shape (Fig. 6d). Presence of calceoli is vagarious. Gener- ally, adults of A. huxleyana bear calceoli but PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON most of A. regis do not. We have in hand several adults of A. huxleyana without cal- ceoli and Stebbing (1888: pl. 73) shows an adult of A. regis with calceoli. This fits the vagarious occurrence of calceoli classically known in many species of amphipods where presence or absence occurs at a demal level. Note on other species. — Atylus? batei Cun- ningham, 1871, from Magellan Strait, ap- parently is neither of the species studied herein as it has five dorsal body teeth, whereas the two discussed herein have four or fewer. Austroregia huxleyana (Bate) Figs. 2-4 Atylus Huxleyanus Bate, 1862:135, pl. 25, fig. 4.—Cunningham, 1871:498. Acanthozone Huxleyana. —Della Valle, 1893-612, pl. 595 fie. 23: Halirages Huxleyanus. —Stebbing, 1906: 291; 1914:362.—Schellenberg, 1931:176, pl. 1, fig. k.—K. H. Barnard 932-155" fig. 93. Halirages stebbingi. — Alonso, 1980:10, fig. 7 (not Schellenberg, 1931). Diagnosis. —Calceoli abundantly present; posterior pereonites not formed into pos- terolateral sharp wings in adults; outer rami or uropods 1-2 with several marginal spines in two rows; dorsal tooth of pleonite 3 usu- ally as large as tooth on pleonite 2. Material.—BMNH Holotype Hermit Is- land, Magellan Strait, Chile, Antarctic Ex- pedition, female “‘a’’ 19.8 mm, lacking an- tenna 2 (illustrated). BMNH 1928.12.1: 2122-27, Syntype Falklands, R. Vallentin Expedition, formerly identified as regis, one specimen. BMNH 1936.11.2: 1381-84, st. 53, 12.5.1926, Discovery 0-2 m, female “b”’ 14.80 mm, identified as huxleyanus by K. H. Barnard. BMNH 1936.11.2: 1381-84 (Part), st. 56, 16.5.1926, Discovery BtS 10.5-16 m, identified as huxleyanus by K. H. Barnard, five specimens partly frag- mented. VOLUME 102, NUMBER 3 705 Fig. 2. Austroregia huxleyana: holotype, female “‘a’’ 19.8 mm, a, Body; b, Apex of right mandible; c, Antenna 1 lateral; d, Pleon; e, Apex of left mandible; f Head; g, Right lacinia mobilis; h, Prebuccal, anterior view, dorsal to right. Female “b” 14.8 mm: i, Head; j, Antenna 2, medial; k, Pleonite 7 (left) to pleonite 4 (right). 706 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Hd) ! N By ' ' mn Fig. 3. Austroregia huxleyana: holotype, female “a” 19.8 mm, a, Region of accessory flagellum; 5, Gill 6; c, Coxa 1; d, Coxa 2 and oostegite; e, Coxa 3 and gill: f Coxa 4 and oostegite; g, Coxa 5 and oostegite (small) and gill (large); 4, Lower lip; i, Right mandible; 7, Outer plate of maxilla 1; k, Maxilla 2; /, Palp of right mandible; m, Maxilla 1; n, Maxilliped. VOLUME 102, NUMBER 3 Ger KAA 107 Fig. 4. Austroregia huxleyana: holotype, female ‘“‘a” 19.8 mm, a, Gnathopod 1, lateral; b, Gnathopod 2, medial; c, Pereopod 7; d, e, Telson; f—h, Uropods 1, 2, 3; i, Pereopod 4. Description of holotype female “a’’ 19.8 mm.—Antenna 2 missing, nee 5-7 broken, not available from better specimen, no enlarged views of pereopods 4—6 possi- ble. Rostrum small, blunt; eyes small, cir- cular, formed of ommatidia surrounding dark core of pigment, lateral cephalic lobes not protruding, truncate, antennal sinus weak, concave, bounded by tooth below. Antenna | about 50 percent as long as body, peduncle short, articles successively shorter, articles 2 and 3 with weak apicoventral tooth, each with 8+ ventral calceoli, acces- sory flagellum fused to peduncle, formed of small squared boss bearing one long and 3 short setae, primary flagellum with 44 ar- ticles, some basal articles of bead form, first slightly enlarged and subrectangular, others following of varying sizes in sets of 2 and 3, narrower to broader, calceoli present, ter- minal member of each set with groups of aesthetascs, broad articles with aesthetascs = 2.40.0, 12,1 5,18.22.2529.55,00, aes- thetascs on these articles posterior, numbers 708 eS ee he eee for the most part calceoli occurring in com- plex positions similar to complex setal dis- tributions of following species, regis, for ex- ample (‘‘a’’ = anterior, = inner, O° 66599 1 0” = outer, ““‘p’’ = posterior, from lateral views), article 1 with 0, article 2 = lp, article 3 = 2a,1-0,1p, article 4 = Ip, article 5 = Ip, articles 6,9,13 = 2a,2-0,31,1p, articles 7,8, PO212.,14,15:17,18'20,2272495 27229 333: and 35—44 = Ip, articles 11,16,19,21,23,26, 28,30,34 = 1la,1-2-0,2-31,1-2p, article 32 = 2a,2-0,11,2p. Antenna 2 missing (see next specimen). Body weakly depressed, pereonites shal- low, lacking lateral ridges above coxae, not produced strongly posterolaterally, but weakly on 7, from dorsal view these seg- ments not forming lateral wings, pereonite 7 with posterodorsal tooth, pleonites 1—2 with larger tooth, pleonite 3 with equally long thicker tooth, urosomites 1—3 rounded above. Coxae shorter than depth of pereo- nites, coxae 1-4 softly quadrate, with rounded distal corners, almost subcircular, coxae 3-4 weakly excavate behind, poste- rior lobe of coxa 5 as long as coxa 4, coxa 6 not shorter than 5, coxa 7 shortest, un- lobed; coxae poorly armed. Epistome and labrum rounded truncate anteriorly, see illustration for anterior view. Right and left incisors multitoothed, right lacinia mobilis with about 3 teeth, left with 6 teeth, about 8 right and 9 left rakers, each with extra interrakers, molar strongly tritu- rative, palp article 2 densely setose medi- ally, article 3 weakly bent, with no outer basal setae (=A-setae), inner margin with several C-setae and many D-setae, apex with 8+ E-setae. Lower lip lacking inner lobes. Inner plate of maxilla 1 subrectangular, with 6 apicomedial setae, outer plate with 11 mostly denticulate spines, palp article 2 symmetrical on both sides, with 7 thick api- cal spines, 2 thinner apicolateral spines, no facial subdistal setae. Inner plate of maxilla 2 as long as and scarcely narrower than outer plate, lacking facial row of setae, medial PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON margin with three or four setae stouter and longer than apical cluster. Inner plate of maxilliped with 3 stout apical spines, one ventromedial coupling spine (not shown in illustration), outer plate broad and squat, with six apical setae, medially with thin ta- pering setae occasionally in pairs, palp ar- ticles 1-2 broad, 1—2 with few apicolateral and 2 with many medial setae, article 3 weakly geniculate, moderately setose me- dially, dorsolateral face with groups of setae, dactyl stubby, with short apical nail and many accessory inner setules. Gnathopods small, first scarcely larger than second, wrists slender, barely lobate, hands much longer than wrists, ovatorect- angular, palm of gnathopod 1 very oblique, defined by group of two facial spines, with two secondary groups outside dactylar apex on posterior margin of hand, dactyl simple, curved, lined with bent setules; palm of gnathopod 2 slightly better defined, rela- tively shorter than on gnathopod 1. Pereo- pods 3-4 as illustrated, locking spines 3 di- verse members, posterior margins of article 6 with five armament sets each with two or three spines and spinules, dactyls simple, each with inner marginal and outer facial setule. Article 2 of pereopods 5-7 diverse, weakly ovate to more sharply trapezoidal successively, subrectangular distally, weak to strong posteroventral lobe present, limb lengths increasing successively. Gills pres- ent on coxae 2-6; oostegites poorly devel- oped but of slightly expanded form on coxae 2-5, slightly pointed on coxa 2, paddle shaped on coxae 3-4, small and paddle shaped on coxa 5, setae absent but marked by humps or sockets. Epimera 1-3 alike, each with convex pos- terior margin, sharp and weakly turned pos- teroventral tooth connected to lateral ridge, otherwise naked. Outer rami of uropods 1— 2 shortened, all rami bearing apical spines and two marginal rows of spines, both upper margins of peduncle on uropods 1-2 with row of spines, those on uropod 1 lateral margin confined to distal half. Peduncle of VOLUME 102, NUMBER 3 uropod 3 slightly elongate, with five medial spines, rami extending equally, slightly curved apically, sharp, each with two rows of marginal spines, these spines doubled on outer margin of outer ramus, inner margin of inner ramus also strongly setose, other margins with sparse setae mostly apical. Telson elongate, basally broadened, weakly tapering, cleft about 20 percent of its length, each side of apex with four setules in tan- dem. Female “‘b’”’ 14.80 mm. —Enlargement of head illustrated; antenna 2 illustrated, ar- ticles 3—5S of peduncle furnished with ventral calceoli, articles 4-5 very short, flagellum thick, with 52 articles, last vestigial, aesthe- tascs absent, articles short and bead-like, calceoli present in relatively uniform pat- tern (“‘a” = anterior, ““m”’’ = medial, “‘p” = posterior), lp present on articles 1,3,4,6,8, 11,13,16,19,23,25,29,31,34,36,39,41,43,45, 48, none present on articles 50-51, articles 9,10,12,14,15,17,18,20,24,26,30,32, with 2a,3m,2p, articles 2,5 with 2a,4m,2p, article 7 with 2a,3m,1p, articles 21,22,27,28,33,37 with 2a,2m,2p, article 35 with 2a,2m,3p, articles 38,40,42,44,.46 with 2a,1m,3p, ar- ticle 47 with 2a,3p, article 49 with 2a,2p. Dorsal body cuspidation of 2-toothed form, only pleonites 1—2 each with medium sized dorsoposterior tooth; pereopods 5—7 and uropods 1-3 badly broken. Distribution. — Boreal South America and Falkland Islands, 0-55 m. Austroregia regis (Stebbing) Figs. 5-7 Halirages huxleyanus. —Stebbing, 1888: 902, pl. 73 (not Bate, 1862). Bovallia regis Stebbing, 1914:362, pl. 8. Halirages regis. —K. H. Barnard, 1932:161, fig. 94. Halirages stebbingi Schellenberg, 1931:176, pl ie. 1. Diagnosis. —Calceoli usually absent; pos- terior pereonites formed into posterolateral sharp wings in adults; outer rami of uropods 709 1—2 with two or fewer marginal spines; dor- sal tooth of pleonite 3 usually smaller than tooth of pleonite 2. Material. —Syntypes, BMNH 1928.12.1: 2122-27, Roy Cove, Falkland Islands, low spring tide, R. Vallentin Expedition: Lec- totype, here selected, male “‘c” 6.81 mm (illustrated); syntype male “‘d” 6.28 mm; syntype juvenile “e”’ 5.75 mm and one other juvenile. Following specimens all misiden- tified formerly as H. huxleyanus: BMNH 1928.12.11: 2019-26 (part), Stanley Har- bour, Falkland Islands, coll. Stebbing, three giant specimens, male “f’’? 19.80 mm (il- lustrated body), female “g’? 23.81 mm (il- lustrated head and coxae 1-4), female “‘h,” unmeasured. BMNH 1928.12.1: 2019-26 (part), Rock pools, Falkland Islands, 20.11.1910, R. Vallentin Expedition, coll. Stebbing, three specimens, one large, one medium, one small. BMNH 1928.12.1: 2019-26 (part), Falkland Islands, 20.vi.1910, coll. Stebbing, nine juveniles. BMNH 1936.11.2: 1381-84 (part), st. 55, 16.v.1926. BtS 10-16 m, Discovery det. K. H. Barnard, one small form. BMNH 1936.11.2: 1385, st. 56, 16.v.1926, Net: BtS, 10.5-16 m, “white, very heavily mottled with deep crimson,” Discovery det. K. H. Barnard, female “‘i,”” unmeasured, with long head tooth. Description of lectotype male “‘c’’ 6.81 mm.—Antenna 2 missing, pereopods 3-4, 6—7 broken, pereopods 5-7 illustrated from better specimen, left mandible only de- scribed, no enlarged views of head, epi- stome, labium, pereopods 3-7, dactyls. Rostrum small, blunt; eyes circular, with dark core of pigment, lateral cephalic lobes not protruding, truncate, antennal sinus weak, straight, oblique. Antenna | almost 55 percent as long as body, peduncle short, articles successively shorter, accessory fla- gellum fused to peduncle, formed of small squared boss bearing | long and 3 short se- tae, primary flagellum with 30 articles, some basal articles of bead form, first slightly en- larged and rectangular, others following of 710 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON i/ Y ee Ee. ¥ \WA Zz ie ea LF} = ve oe: Etec sace I) if CES = va \ oe a Fig.5. Austroregia regis: lectotype male “‘c’’ 6.81 mm, a, Body; c, Left uropod 3; d, Right molar; e, Maxilliped; g, Lower lip; h, Maxilla 2. Male ‘“‘d” 6.28 mm, b, Head. Male ‘“‘c’”’ 6.81 mm, f, Right mandibular palp. VOLUME 102, NUMBER 3 711 Fig. 6. Austroregia regis: lectotype male “‘c’’ 6.81 mm, a, Maxilla 1; e, Other palp of opposite maxilla 1; f Medial antenna 1; g, Dorsal body, head at bottom; 4, Coxa 4 with gill; 7, Telson. Female “g” 23.81 mm, 5, Head and pereonites 1-5. Male “‘f”’ 19.8 mm, c, Dorsal body from pereonite | (top) to pleonite 1 (bottom); d, Body. 712 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 7. Austroregia regis: lectotype male “‘c” 6.81 mm, a, Region of accessory flagellum; b, Coxa 1; c, Coxa 2 with gill; d, Coxa 3 with gill; e, Coxa 5 with gill; £ Coxa 6 with gill; g, Coxa 7; h, Gnathopod 1, medial; 7, Gnathopod 2, medial; k, /,) Uropods 2 and 1; m, Pleon. Male ‘“‘d” 6.28 mm, j, Pereopod 3. VOLUME 102, NUMBER 3 varying sizes in sets of 2 and 3, narrower to broader, calceoli absent, terminal member of each set with groups of aesthetascs, broad articles with aesthetascs = 1,3,5,8,11,14,17, 20,22,24,26,28, aesthetascs on these articles posterior, numbers = 4,2,2+,3+,4,3,3,1,2, 2,1,1, setal distributions complex, for ex- ample (“‘a’”’ = anterior, “1” = inner, “m” = marginal, “‘s’” = submarginal, from lateral views), article 1 with 2a,2p, article 2 = 0a, Ip, article 3 = 2a,5p, article 4 = 3m,2p, article 5 = 2a,4+p, articles 6,9,12,15 = 2-3s,31, articles 7,10,13,16 = 2—3m,3—4p, articles 8,11,14,17 = 2-3a,3-5p, etc. Antenna 2 missing (see next specimen). Body weakly depressed, pereonites shal- low, pereonites 3-7 with weak lateral ridge above coxae, pereonites 4—7 produced pos- terolaterally, weakly on 7, from dorsal view these segments forming lateral wings more rudimentarily expressed on pereonites 1-3, pereonite 7 with posterodorsal tooth, ple- onites 1—2 with larger tooth, pleonite 3 with short weakly acute dorsal projection, uro- somites 1-3 rounded above. Coxae shorter than depth of pereonites, coxae 1-4 softly quadrate, with rounded distal corners, cox- ae 3—4 weakly excavate behind, posterior lobe of coxa 5 as long as coxa 4, coxa 6 scarcely shorter than 5, coxa 7 shortest, un- lobed; coxae poorly armed. Epistome and labrum as shown for A. huxleyanus. Incisors and laciniae mobiles of mandibles heavily encrusted, similar to A. huxleyanus, thus right incisor with about eight teeth, right lacinia mobilis with three teeth, about six rakers, molar strongly trit- urative, palp article 2 with most setae form- ing apicolateral row, article 3 weakly bent, with one (right) or two (left) outer basal setae (=A-setae), inner margin with several C-se- tae and many D-setae, apex with two E-se- tae. Lower lip lacking inner lobes. Inner plate of maxilla 1 subrectangular, with four apicomedial setae, outer plate with nine mostly denticulate spines, palp article 2 asymmetrical on both sides, with five thick apical spines, one thinner apicolateral spine, 713 two facial subdistal setae on left side, right side with five thin apical spines and two subapical setae. Inner plate of maxilla 2 slightly longer and narrower than outer plate, lacking facial row of setae, medial margin with three setae stouter and longer than api- cal cluster. Inner plate of maxilliped with three stout apical spines, one ventromedial coupling spine, outer plate broad and squat, with six apical setae, medially with thin blades occasionally in pairs, palp articles 1— 2 broad, 2 with few apicolateral and many medial setae, article 3 weakly geniculate, poorly setose medially, dorsolateral face with groups of setae, dactyl stubby, with short apical nail and three accessory inner setules. Gnathopods small, first larger than sec- ond, wrists slender, barely lobate, hands much longer than wrists, subrectangular, palm of gnathopod 1 very oblique, defined by group of three or four marginal and two facial spines, with secondary group outside dactylar apex on posterior margin of hand, dactyl simple, curved, lined with bent set- ules; palm of gnathopod 2 better defined, relatively shorter than on gnathopod 1. Pe- reopods 3—4 as illustrated for specimen “‘d”’ below, locking spines 2 diverse members, posterior margins of article 6 with three or four armament sets each with one spine and from zero to two short setae, dactyls simple, each with inner marginal and outer facial setule. Article 2 of pereopods 5-7 narrow, subrectangular distally, weak posteroven- tral lobe present, limb lengths increasing successively. Gills present on coxae 2-6. Epimera 1-3 alike, each with convex pos- terior margin, sharp and strongly turned posteroventral tooth connected to lateral ridge, otherwise naked. Outer rami of uro- pods 1—2 shortened, each bearing only api- cal spines, inner rami with two rows of mar- ginal spines besides apical cluster, peduncle of uropod 1 with only one small apicolateral spine, medial margin with five spines, pe- duncle of uropod 2 with three dorsolateral spines and one basal setule, medial margin with five spines. Peduncle of uropod 3 714 slightly elongate, with two medial spines and one basal seta, rami extending equally, curved apically, sharp, each with two rows of marginal spines, inner also with seta in last two apical spine sets, outer also with seta in last spine set medially, each ramus with subapical setule. Telson elongate, ba- sally broadened, weakly tapering, cleft about 10 percent of its length, each side with three setules in tandem. . Male ‘d’’ 6.28 mm.—Pereopod 3 en- larged to show details. Male ‘“f’ 19.80 mm.—Supposed termi- nal adult form, body illustrated: pereonites more alate posterolaterally, pereonites 2—7 with shelf above coxae, pointed on pereo- nite 7; coxae more diverse, coxa 2 bluntly pointed below, coxae 3-4 relatively larger than in juvenile form; antenna 2 as illus- trated, articles 4-5 of equal length, flagellum “proliferate,” basal article with new articles forming inside; dorsal tooth of pereonite 3 enlarged; apices of pereopods 6-7 illustrat- ed on body. Female “g’’ 23.81 mm. — Head with large anteroventral tooth below antennal sinus (illustrated). Coxae 3-4 sinuous postero- ventrally. Outer ramus of uropod 2 with two marginal spines. Female “h” large, unmeasured. —Outer ramus of uropod | with one marginal spine, of uropod 2 with one or two marginal spines (right and left); coxae acuminate as in fe- male “g”’ but head lacking tooth. Female “i”? unmeasured.—Like female “gs” but smaller, coxae more strongly acu- minate; each outer ramus of uropods 1-2 with one marginal spine. Discussion.—The lectotype is unfortu- nately a small male less than one-third as long as the largest adults known (as mea- sured by parabolic method). It clearly has the flared-out posterolateral margins of the pereonites but not to the exaggerated degree of larger specimens. In large specimens the pereonal margins flare out laterally until they form horizontally projecting teeth. In some specimens the posterior margins of the flared PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON teeth are incised posteriorly and have very extended lateral wings which look like sep- arate processes from lateral view but which are simply more knife-like. Large adults oc- cur in two forms: (1) head with large pos- teroventral tooth coupled with strongly acu- minate anterior coxae and (2) head without strong posteroventral tooth coupled with poorly acuminate anterior coxae. Small specimens lack dorsally marginal spines on the outer rami of uropods 1-2 whereas larg- er adults have as many as three and two spines on the outer rami of uropods 1 and 2 respectively. No calceoli have been dis- covered in our specimens of this species, although Stebbing shows calceoli in his orig- inal description. Relationship. — Differing from A. huxley- anus in the presence of flared margins on the pereonites, the poorly spinose outer rami of uropods 1-2 and the usual absence of calceoli. The specimen figured by Stebbing (1888: pl. 73) has calceoli. The condition of coxae in adult huxleyana resembles that of juvenile regis so that coxal forms are not necessarily definitive in comparing the two species. There remains the question of whether A. regis 1s congeneric with A. huxleyana be- cause the two species differ in the setal for- mulas on article 3 of the mandibular palp, in the spine numbers on the outer plate of maxilla 1, the difference in setal presence on the palps of maxilla 1, and the symmet- ricity of those palps. Article 3 of the man- dibular palp lacks A-setae in huxleyanus; the latter has 11 spines on the outer plate of maxilla 1 whereas A. regis has 9; and the palps are symmetrical and lack subdistal se- tae in A. huxleyana. There also remains the problem of speciation in this group and be- cause of the extensive demal variations al- ready noted, very large and widespread col- lections of antarctic materials should be assembled to study the life history and vari- ations in the group. Distribution. — Tierra del Fuego and Falk- land Islands, 0-55 m. VOLUME 102, NUMBER 3 Acknowledgments Dr. Roger J. Lincoln of the British Mu- seum of Natural History made extensive contributions to this work and kindly sup- plied the SEM photographs. I thank Linda B. Lutz of Vicksburg, Mississippi, for ink- ing our illustrations and Patricia B. Crowe and Kimberly R. Cleary of the Smithsonian Institution for their assistance in the labo- ratory. Literature Cited Alonso, G. 1980. Amfipodos de la Ria Deseada (San- ta Cruz-Argentina).—Centro de Investigacion de Biologia Marina (CIBIMA) del sistema de Centros del Instituto Nacional de Tecnologia Industrial (INTI), Contribucion Cientifica 175: 3-15. Barnard, J. L.,& G.S. Karaman. 1987. Revisions in classification of gammaridean Amphipoda (Crustacea), Part 3.— Proceedings of the Biolog- ical Society of Washington 100:856-875. Barnard, K. H. 1932. Amphipoda.— Discovery Re- ports 5:326 pp. Bate, C. S. 1862. Catalogue of the specimens of am- phipodous Crustacea in the collection of the British Museum. London, British Museum [of Natural History], 399 pp. Bousfield, E. L. 1977. A new look at the systematics of gammaroidean amphipods of the world.— Crustaceana, Supplement 4:282-316. . 1983. An updated phyletic classification and paleohistory of the Amphipoda. Pp. 257-277 in AIS F. R. Schram, ed., Crustacean phylogeny, A. A. Balkema, Rotterdam, 365 pp. Cunningham, R.O. 1871. Notes on the reptiles, Am- phibia, fishes, Mollusca, and Crustacea obtained during the voyage of H.M.S. ‘Nassau’ in the years 1866-69.—Transactions of the Linnean Society of London 27:465-502. Della Valle, A. 1893. Gammarini del Golfo di Na- poli.— Fauna und Flora des Golfes von Neapel und der angrenzenden Meeres-Abschnitte, Monographie 20:xi and 948 pp. Lincoln, R. J., & D. E. Hurley. 1981. The calceolus, a sensory structure of gammaridean amphipods (Amphipoda: Gammaridea).— Bulletin of the British Museum of Natural History (Zoology) 40:103-116. Schellenberg, A. 1931. Gammariden und Caprelliden des Magellangebietes, Sudgeorgiens und der Westantarktis.—Further Zoological Results of the Swedish Antarctic Expedition 1901-1903 2(6):290 pp. Stebbing, T. R. R. 1888. Report on the Amphipoda collected by H.M.S. Challenger during the years 1873-76.—Report on the Scientific Results of the Voyage of H.M.S. Challenger During the Years 1873-1876, Zoology 29:xxiv and 1737 Pp 1906. Amphipoda I: Gammaridea.— Das Tierreich 21:806 pp., Berlin. 1914. 1. South African Crustacea (Part VII. of S.A. Crustacea, for the marine investigations in South Africa).— Annals of the South African Museum 15:1-55. Department of Invertebrate Zoology, NHB-163, Smithsonian Institution, Wash- ington, D.C. 20560. PROC. BIOL. SOC. WASH. 102(3), 1989, pp. 716-725 TWO NEW SPECIES OF WOOD-BORING LIMNORIA (CRUSTACEA: ISOPODA) FROM NEW ZEALAND, L. HICKSI AND L. RENICULUS Marilyn Schotte Abstract.—Limnoria hicksi is described and distinguished from L. indica Becker & Kampf, 1958 by the structures on the fifth pleonite and the pleotelson. Limnoria reniculus, similar to Limnoria foveolata Menzies, 1957, L. saseboensis Menzies, 1957, L. sexcarinata Kiihne, 1975 and Limnoria sublittorale Menzies, 1957, differs from all four in the structure of the fifth pleonite, pleotelson and morphology of the lacinioid seta of the right mandible. At present only two reliably identified wood-boring isopods of the genus Limnoria have been reported from New Zealand. These are L. guadripunctata Holthuis from Portobello (Hurley 1961), Auckland Har- bour (McQuire 1964), and Port Nicholson (Ralph & Hurley 1952), and L. tripunctata Menzies, also from Auckland Harbour (McQuire 1964). Menzies (1959) listed L. quadripunctata site records for isopods he examined from Auckland, Wellington, and Napier, previously thought to be specimens of L. lignorum (Rathke). Chilton (1914) had assigned the latter name to wood-borers found in timbers in Auckland and Lyttleton harbors and in piles from Akaroa Harbour. The latter isopod, whose name was assigned to most limnoriids before Menzies’ 1957 monograph, has an Arctic-boreal distribu- tion and has been reliably reported only in the Northern Hemisphere from 39° to 58°N and from Iceland (Menzies 1957). Two alga- borers have been found in New Zealand, Phycolimnoria stephenseni Menzies, in floating Lessonia near Auckland Island (also Macquarie Island), and P. segnis (Chilton) from seaweed in Lyttleton and Akaroa har- bors (Menzies 1957) as well as from Por- tobello (Hurley 1961). Across the Tasman Sea, Limnoria quad- ripunctata has been found recently in Aus- tralia at Goat Island and Sydney Harbour in association with L. indica and L. tri- punctata Menzies, as well as in Tasmania (Cookson 1987). Hale (1929) reported L. lignorum from Port Lincoln but this iden- tification is undoubtedly erroneous. L. sub- littorale Menzies has been collected from New South Wales (Menzies 1957), as has Phycolimnoria rugosissima Menzies. An additional alga-borer, P. nonsegnis Menzies from Tasmania, brings to eight the total number of limnoriids thus far known from the Australia-New Zealand region. Material of the new species has been de- posited at the National Museum of New Zealand in Wellington and the National Museum of Natural History, Smithsonian Institution, Washington, D.C. Family Limnoriidae Limnoria (Limnoria) hicksi, new species Figs. 1A—D, 2A—F, 3A-E Material. —Holotype, NMNZ Cr. 5702, 2 tl 4.0 mm, Paratype, USNM 205960, 2 tl 3.3 mm (on slide and SEM stub), stn K6 R/V Kalinovo, 177°39'05’E, 37°23'07’S to 177°36'06”E, 37°23'07’S, off New Zealand, 1075-1100 m, in rotting wood, 23 Nov 1981, coll. G. R. F. Hicks. Description.—Female: Pigment absent. Pleonite 5 with a row of four proximal tu- bercles and two somewhat larger distal tu- 717 VOLUME 102, NUMBER 3 uorneulds ‘ C ‘podieyjus uos[s}09]d Jo oseq 1k so[dsoqn | ‘O ‘uos[a}09;d JOLIOJUR pure d1UON|d YL “¢ ! ‘uos[o}Oo]d JO uld1eW [e1ISIP UO UOS[IN1OI[q “YW USYIY DIMOUUIT “| “BL WHOOT ee A> y Sbay S te. q ae ” - te 718 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON W yi 1] m i " Fig. 2. Limnoria hicksi: A, Fifth pleonite and pleotelson; B, Antennule and antenna; C, First pereopod; D, Seventh pereopod; E, First pleopod of female; F, Uropod. bercles, all bearing short spinules. Lateral crests of pleonite 5 with long setae. Pleo- telsonic margin not evenly rounded but with pair of shallow symmetrical notches in pos- terior third. Base of pleotelson with a row of four large tubercles and pair of smaller tubercles posterior to these, all spinose (Fig. 1C) and bearing at least one long seta. Lat- eral crests with long setae. Surface of pleo- telson regularly spinulose except for bare patches lateral to posterior tubercles. Pos- terior margin not tuberculate but with both simple and sheathed spines (Fig. 1D). Antennular peduncle of three articles, first VOLUME 102, NUMBER 3 Fi Se B Fig. 3. Limnoria hicksi: A, Outer lobe of first maxilla; B, Second maxilla; C, Maxilliped; D, Left mandible; E, Lacinioid seta of right mandible. and third articles subequal in length; fla- gellum consisting of one very short basal article and two subequal articles. Antennal flagellum of four articles, first article longer than three distal articles combined. Mandibular palp of three articles, second article longest, terminal article bearing five distal fringed spines; “‘rasp”’ of left incisor strongly sclerotized; spine row of right man- dible of nine laciniate spines, distal edge of lacinioid seta broad, very finely toothed and acute at ends. Maxilla 1 with nine stout se- tae, four of which provided with blunt teeth and one with setules. Maxilla 2 as figured. Maxillipedal endite with single coupling hook and six setose spines on distal margin; articles 2 and 3 of palp subequal in length; epipod about three times longer than great- est width, not reaching distal margin of basi- pod. Pereopod | with bidentate accessory spine at base of dactylar unguis; propodus with one dentate and one fringed posterodistal spine. Pereopod 7, accessory spine of dactyl small and apparently not bidentate; carpus with many fringed spines on distal margin; merus with anterodistal margin somewhat produced and bearing numerous fringed 720 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 4. Limnoria reniculus: A, Pleotelson and fifth pleonite; B, Pleotelsonic surface enlarged; C, Interior surface of kidney-shaped carina enlarged; D, Pleotelsonic margin. spines visible in lateral view. Endopods of for its collector, Dr. Geoffrey R. F. Hicks, pleopods 1 and 2 of female distally truncate. Structure of male second pleopod unknown. Uropodal exopod less than one-half length of endopod; latter bearing one plumose seta and several terminal setae of varying lengths; peduncle with row of short, simple setae on outer margin and additional row of long plumose setae near margin. Remarks.—The male of the sexually di- morphic L. indica is the only other known limnoriid with six basal tubercles on the pleotelson arranged similarly to those in L. hicksi. Instead of additional tubercles on the fifth pleonite, L. indica in both sexes pos- sesses two subparallel longitudinal carinae bearing setae, thereby distinguishing it from L. hicksi. Etymology. —The new species is named National Museum of New Zealand, Wel- lington, New Zealand. Limnoria (Limnoria) reniculus, new species Figs. 4A—D, 5, 6A—G, 7A—-E Material. —Holotype, NMNZ Cr. 5703, ovig. 2, tl 5.0 mm, Allotype, NMNZ Cr. 5704, 6, 4.4 mm, Paratypes, NMNZ Cr. 5705, 11 ovig. 2, 16 2, 15 6, South Taramaki Bight, North Island, New Zealand, R/V James Cook, sta J20/25/84, 40°56.4'S, 174°44.0’E to 40°59.8’S, 174°43.7’E, in rot- ting wood, 144-182 m, 29 Nov 1984, coll. G. R. F. Hicks. Paratypes, USNM 205961, 11 ovig. 2, 15 2, 14 6, from same locality. Other material: USNM 205962, 1 speci- men, South Taranaki Bight, New Zealand, VOLUME 102, NUMBER 3 m Limnoria reniculus: Different specimen from same lot as specimen in Fig. 4, pleotelson and fifth 400M Fig. 5. pleonite. sta J20/16/84, 40°33.0’'S, 173°04.6’E to 40°31.7’'S, 173°01.2’E, in rotting wood, 47— 52 m, 28 Nov 1984, coll. G. R. F. Hicks. — USNM 205963, 5 specimens, sta J15/40/ 84, 40°33.4'S, 173°04.6’E to 40°32.4’S, 173°02.6’E, in rotting wood, 44-52 m, 28 Nov 1984, coll. G. R. F. Hicks. —USNM 205964, 11 specimens, sta J16/6/84, S751 7 7°19'E to 37°51'S, 177°15’E, in rotting wood, 48-49 m, 17 Sep 1984, coll. G. R. F. Hicks—USNM 205965, J15/22/ 84, 41 specimens, 42°21.6’S, 170°49.5’E to 42°24.4'S, 170°48.1’E, in rotting wood, 179-— 184 m, 3 Sep 1984, coll. G. R. F. Hicks. — USNM 205966, sta J15/40/84, 5 speci- mens, 42°59.4’S, 170°16.5’'E to 42°57.2’'S, 170°19.3’E, in rotting wood, 60-62 m, 5 Sep 1984, coll. G. R. F. Hicks. —USNM 205967, 6 specimens, sta J15/28/84, 42°45.8’S, 170°28.2’E, in rotting wood, 50-64 m, 4 Sep 1984, coll. G. R. F. Hicks.—USNM 205968, 3 specimens, sta J20/12/84, 40°46.7'S, 173°48.0’E to 40°47.9'S, 173°47.3’E, in rot- ting wood, 60-65 m, 27 Nov 1984, coll. G. R. F. Hicks. —USNM 205969, 6 specimens, off North Island, New Zealand, R/V James Cook, sta J9/70/84, 38°02'S, 174°37'E to 38°06'S, 174°36’E, in rotting wood, 50-51 m, 3 Jun 1984, coll. G. R. F. Hicks. —USNM 205970, 3 specimens, sta J16/2/84, 37°39’'S, 17 7°28'E to: 37°39'S, °177°24'E, in ‘rotting wood, 134-137 m, 17 Sep 1984, coll. G. R. F. Hicks.—USNM 205971, 8 specimens, New Zealand, off Cape Runaway, Wanaka, sta WK 3/19/85, 37°29.9'S, 177°47'E, in Log A, 450-481 m, 8 Dec 1985.—USNM 205972, 19 specimens, New Zealand, Bay of Plenty, east of Alderman Island, NUNZ Cr. 5152, from timber, 400-520 m, Jun 1969, coll. R. D. Cooper.—USNM 205973, 14 specimens, New Zealand, WNW White Island, Tumokemoke Knoll, R/V Tangaroa (NZOI sta R 76), 37°29.1'S, 176°54.7'E, in twig, 248-283 m, 20 Jan 1979. Description. —Male: Dorsal integument of pereon and pleon rather foveolate, es- pecially pleonite 5. Pleonite 5 with some- what variable U-shaped carina (Figs. 4A and 722 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON F Fig. 6. Limnoria reniculus: A, Antennule of male; B, Antenna; C, Lacinioid seta of right mandible; D, Left mandible; E, First maxilla; F, Second maxilla; G, Maxilliped. VOLUME 102, NUMBER 3 5). Pleotelson densely spinulose with two rounded submedian ridges basally, flanked on either side by irregular, kidney-shaped Carinae, apparently bare interiorly but ac- tually covered with tiny spinules (Fig. 4C). Submedian ridges becoming obsolete dis- tally. Margins of pleotelson not tuberculate, having multiple rows of simple, unsheathed spines. Female dorsum as in male. Antennular peduncle of three articles; fla- gellum of male consisting of basal article, much shorter than wide, penultimate article with many aesthetascs ringing distal mar- gin, and terminal article half as wide as pre- ceding article. Antennal flagellum of 4 ar- ticles, basal article longer than 3 distal articles together. Mandibular palp of 3 ar- ticles, two proximal articles subequal in length; article 2 with 6 distal fringed spines; terminal article bearing 9 distal fringed spines; spine row of left mandible a single process; spine row of right mandible of 10 laciniate spines, increasing in length proxi- mally; lacinioid seta with two lobes, one elongate and blunt, the other short and jag- ged; incisor with strongly sclerotized cusp. Maxilla 1 and 2 as figured. Maxillipedal en- dite with single coupling hook, eight spines on distal margin, five of which setulose; pal- pal article 3 longest and widest; epipod more than three times longer than greatest width, distally rounded and not reaching base of palp. Pereopod 1, bidentate accessory spine at base of dactylar unguis; propodus with 1 dentate, plus one fringed posterodistal spine. Carpus and merus with short rows of round- ed scales on posterior surface. Pereopod 7, accessory spine of dactyl barely bidentate; carpus with many fringed spines on distal margin; merus produced anterodistally, bearing many fringed spines encircling dis- tal margin. Appendix masculina of pleopod 2 of male articulating proximal to mid- length on median margin of endopod, ex- tending beyond ramus. Uropodal endopod elongate, about four times longer than wide and bearing five plumose setae and several lateral and terminal simple setae; uropodal 7123 exopod a slightly curved claw; peduncle with row of setae inserted medially to lateral margin. Remarks. —L. reniculus may be confused with L. saseboensis Menzies, which also has submedian longitudinal ridges on the pleo- telson, especially if debris obscures the kid- ney-shaped carinae in uncleaned specimens of the former. The U-shaped carina on pleonite 5 in reniculus may vary but the subparallel ridges do not join proximally as they appear to do in saseboensis. Menzies’ description notes the posterior edge of the pleotelson and lateral crests as tuberculate with “‘spike-like bristles on the margin.” SEM photos (Fig. 4) do not reveal obvious tubercles in the present species. The two species also differ somewhat in the shape of the lacinioid seta of the right mandible with “two medially curved teeth at apex” in L. saseboensis. L. sublittorale Menzies from Australia also has a pair of longitudinal ca- rinae on the telson but apparently lacks oth- er surface ornamentation there. The lacin- 10id seta of the right mandible has two recurved teeth instead of a blunt lobe as in L. reniculus. L. foveolata Menzies, while also foveolate on the pleotelson and pleonite 5, has carinae on both segments which are more irregular than those in the new species, and it lacks the kidney-shaped features. The la- cinioid seta of L. foveolata as figured by Menzies has three teeth at the apex. The new species also resembles L. sex- carinata Kuihne. Examination of type ma- terial of the latter species reveals the carina of pleonite 5 as ““horseshoe-shaped” and the curving ridges on the pleotelson do not join distally. Kiihne’s (1975) description states that the lacinioid seta of the right mandible, apparently a good character for distinguish- ing members of this genus, is ““branched and furnished at the end with teeth.’’” SEM pho- tos of L. sexcarinata show the presence of tubercles and sheathed spines on the pleo- telsonic margin, neither of which exist in L. reniculus. Etymology.—The Latin reniculus (little 724 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON sR POLL —— Fig. 7. Limnoria reniculus: A, First pereopod; B, Dactyl of first pereopod enlarged; C, Seventh pereopod; D, Uropod; E, Second pleopod of male. VOLUME 102, NUMBER 3 kidney), proposed as a noun in apposition, refers to the shape of the outline of the ca- rinae flanking middorsal ridges on the pleo- telson. Acknowledgments Material described in this paper was kind- ly provided by Dr. Goeffrey R. F. Hicks of the National Museum of New Zealand. Mrs. Susann Braden of the NMNH assisted with preparation of the scanning electron micro- graphs. The manuscript benefitted from comments by Laurie Cookson and Brian Kensley, both of whom reviewed the manu- script. Literature Cited Chilton, C. 1914. The species of Limnoria, a genus of wood-boring Isopoda.—Annals and Maga- zine of Natural History, Ser. 8, vol. 13:380-390. Cookson, L. J. 1987. The occurrence of Limnoria indica Becker & Kampf (Isopoda) on the eastern coast of Australia.—Crustaceana 52(1):85-89. Hale, H. M. 1929. The crustaceans of South Austra- lia. 2:201-380, figs. 268-269. Government Printer, Adelaide. (P25) Hurley, D. E. 1961. A checklist and key to the Crus- tacea, Isopoda of New Zealand and the Sub- antarctic Islands.—Transactions of the Royal Society of New Zealand, Zoology 1(20):259-292. Kihne, H. 1975. Neubeschreibung einer holzzerst6- renden Bohrassel, Limnoria sexcarinata (Crus- tacea, Isopoda).—Zeitschnft fur angewandte Zoologie 62:447-455. McQuire, A. J. 1964. A note on the occurrence of marine borers in New Zealand. — Proceedings of the New Zealand Wood Preservers’ Association 4:35-44. Menzies, R. J. 1957. The marine borer family Lim- noriidae (Crustacea: Isopoda).— Bulletin of Marine Science of the Gulf and Caribbean 7(2): 101-200. 1959. The identification and distribution of the species of Limnoria. Pp. 10-33 in Dixie Lee Ray, ed., Marine boring and fouling organisms. University of Washington Press, Seattle. Ralph, P. M., & D. E. Hurley. 1952. The settling and growth of wharf-pile fauna in Port Nicholson, Wellington, New Zealand.— Zoological Publi- cations, Victoria University College (19):1-22. Department of Invertebrate Zoology, Na- tional Museum of Natural History, Smith- sonian Institution, Washington, D.C. 20560. PROC. BIOL. SOC. WASH. 102(3), 1989, pp. 726-731 SOME ASPECTS OF THE BIOLOGY OF RHOPALOPHTHALMUS TATTERSALLAE PILLAI, 1961 (CRUSTACEA, MYSIDACEA) AND EXTENSION OF RANGE INTO THE KHOR AL SABIYA, KUWAIT (ARABIAN GULF) Stephen A. Grabe Abstract. —Rhopalophthalmus tattersallae Pillai was collected from the Khor al Sabiya, Kuwait, extending its known range from the Indian Ocean to the northern Arabian Gulf. Population density was greatest in early fall when juveniles predominated and was lowest in mid-winter. Spawning activity was greatest in April, May, July, and August, and lowest in mid-winter. Brood size was correlated with body size; largest females and broods occurred January through May and smallest females and broods occurred July through Novem- ber. Sex ratio was close to 1:1 on most dates. A small number of R. tattersallae were parasitized by an unidentified dajid isopod. Rhopalopthalmus tattersallae was de- scribed by Pillai (1961; cited in Pillai 1965) from specimens collected in coastal waters off Kerala State, India. To my knowledge, there have been no subsequent records of its occurrence. Fifteen species of Rhopa- lophthalmus have been described (Mauch- line 1980) and all are inhabitants of neritic or estuarine waters (Tattersall 1957; Hodge 1963; Pillai 1965, 1973; Mauchline & Mu- rano 1977; Wooldridge & Erasmus 1980). This paper presents some basic life history information for R. tattersallae and records its range extension into the northern Ara- bian Gulf. Methods. — Daytime zooplankton collec- tions were taken at stations in Kuwait Bay and the Khor al Sabiya (Fig. 1) from Sep- tember 1981 through September 1982. Du- plicate step-oblique tows were made with a 0.5 m diameter plankton net outfitted with 0.202 mm and 0.505 mm mesh. The sam- pling schedule is summarized in Table 1. Sample volumes were measured with a dig- ital flow meter. Additional samples were collected at Station 614 on 27 May and 21 September 1982 specifically for mysids. A 0.505 mm mesh net was towed near-bottom (horizontally); sample volumes were not re- corded. All mysids were sorted from each sample, identified to life stage, and carapace length (CL) measured (tip of the rostrum to the posterior border of the carapace). Brood sizes (number of larvae) were de- termined only for those females whose mar- supla appeared undisturbed. The presence of an unidentified ectoparasitic isopod (Da- jidae) was recorded. Monthly population density in the Khor al Sabiya (Stations 614 & 615) was based upon the September and October 1981 0.202 mm collections and the November through August 0.505 mm mesh collections. To fa- cilitate interpretation of seasonal changes in population structure and breeding, samples were pooled for each date at the Khor al Sabiya stations. Assumptions were made, then, that the populations at the two sta- tions in the Khor were structurally similar and that the 0.202 and 0.505 mm mesh nets were similarly efficient in sampling all life stages of this species. In studies of mysid populations in which VOLUME 102, NUMBER 3 Fig. 1. E29 failaka island Location of the sampling stations in Kuwait Bay and Stations 614 & 615 in the Khor al Sabiya used to survey the Rhopalophthalmus tattersallae population during 1981-1982. longitudinal differences in population struc- ture have been observed, there have either been marked differences in habitat (e.g., depth) within the water body (Mauchline 1970) or there was evidence for recruitment, transport and maturation of a coastally spawned population (Hulburt 1957). Since the Khor al Sabiya stations were of similar depth, temperature, salinity and dissolved oxygen (Dames & Moore 1983) and since there was no evidence of recruitment from Kuwait Bay (see below), the first assump- tion seems tenable. The second assumption, that of compa- rable sampling abilities of the two gears, is more questionable. Sampling rates of the two gears were generally similar in the Khor al Sabiya (12.6 vs. 12.7 m?/minute for the 0.202 and 0.505 mesh, respectively; Dames & Moore 1983). A limited comparison of sampling efficiencies for Penaeidae mysis and postlarval stages found that abundance estimates of the two gears were comparable: 0.202 mm abundance = — 1.38 + 1.1 (0.505 mm abundance); r,, = 0.986; P < 0.01 (Dames & Moore, unpublished data). ANOVA (Sokal and Rohlf 1981) was used to test for differences in mean CL of mature females by sampling date, and regression analysis was used to evaluate the relation- ship between brood size and CL. Results and discussion. —The Khor al Sa- biya is a river-like channel separating the Sabiya peninsula on the northeastern shore of Kuwait Bay from Kuwait’s Bubiyan Is- land (Fig. 1). Maximum depth is about 18 m but study areas averaged <10 m. Bottom sediments at Station 614 were generally rock-mud and shell debris and at Station 615, muddy sand and shell debris. Extremes in water temperature occurred during Feb- ruary (12.1°C surface, 11.8°C bottom) and 728 Table 1.—Sampling schedule for Rhopalophthalmus tattersallae in the Khor al Sabiya, Kuwait, September 1981 to September 1982. Mesh size Mesh size 0.202 0.505 0.202 0.505 Date mm mm Date mm mm 17 Sep 81 X 26 Apr Xa 27 Sep xX 12 May xX X 14 Oct x 18 May DS 28 Oct x 27 May

Non-quantitative, near-bottom samples. June (29.9°, 27.9°). Extremes in salinity oc- curred during April (32.6%, 32.7%o) and June (40.4, 40.5%) (Dames & Moore 1983). Rhopalophthalmus tattersallae was col- lected throughout the year in the Khor al 276 220 © = bts ies oc nu a tr 110 aa = = Zz 65 0 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Sabiya but occurred in only 26% of the sam- ples from Kuwait Bay. Highest densities oc- curred during autumn. Thereafter density declined through February before it in- creased again from April through August (Fig. 2). Concentrations of mysids were higher at Station 614 near the mouth of the Khor al Sabiya than at Station 615. The relationship between Cl and TL was significant and highly correlated (r,3; = 0.99). The regression equation is: TL,,, = 3.88 Che 04 Spawning activity was probably greatest during April, May, July and August when large (22.5 mm CL) sexually mature my- sids and 0.7-0.8 mm CL juveniles com- posed respectively from 33 to 72% and from 3 to 13% of the population. Recruitment of juveniles from April through October (they composed from 32 to 82% of the population — at this time) coincided with the peak in pop- ulation density (Fig. 2). The presence of small, but sexually identifiable, mysids in- dicated that the summer-fall generation ma- © STA615 @ STA614 PO AM WO! Bh BP wh A WN gH yr pO MONTH Fig. 2. Abundance (numbers/100 m3) of Rhopalophthalmus tattersallae in the Khor al Sabiya, Kuwait, September 1981—August 1982. VOLUME 102, NUMBER 3 Table 2.— Percent of larvigerous Rhopalophthalmus tattersalae in the Khor al Sabiya, Kuwait, September 1981 to September 1982. Numbers in parentheses are numbers of specimens collected.* Date % Larvigerous Date % Larvigerous 17 Sep 81 25.0 (64) 12 May 80.0 (10) 27 Sep 0.0 (9) 18 May 100.0 (15) 14 Oct 16.7 (18) 27 May 14.3 (7) 28 Oct 6.3 (16) 7 Jun 50.0 (2) 10 Nov 3.6 (28) 5 Jul 0.0 (20) 17 Nov 2.4 (84) 27 Jul 730-72) 16 Dec 0.7 (148) 9 Aug 66.7 (3) 11 Jan 82 fi (39) 18 Aug 64.0 (25) 3 Mar 33.3 (15) 21 Sep 25.0 (52) 11 Apr 66.7 (3) 2 Females were not collected on 20 Feb, 26 Apr and 29 Aug. 26 20 15 NUMBER 10 729 tured from September through November. These composed from 9 to 24% of the pop- ulation. A relatively sparse overwintering population was characterized by large (=2.5 mm CL) mature mysids and few juveniles. Brooding females were collected on 19 of the 22 sampling dates (Table 2) and brood- ing activity appeared to be greatest during May and from late July through August. The lowest incidence of brooding females occurred between mid-fall and mid-spring. Brooding females ranged in size from 1.8 to 3.3 mm CL (Fig. 3) and ANOVA showed that there were significant differences be- tween dates for mean CL of mature females (F 17279 = 17.0; P < 0.001). Brood size ranged from 2 to 23 larvae (Fig. 4) and the equa- tion, Number of Larvae = 9.3 CL,,,, — 13.7, was significant (F, ,,, = 64.0; P < 0.001). 18 19 20 21 22 238 24 25 26 27 28 29 30 31 32 33 34 CARAPACE LENGTH (MM) Fig. 3. Size frequency distribution of larvigerous Rhopalophthalmus tattersallae in the Khor al Sabtya, Kuwait, September 1981-1982. 730 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON NUMBER OF LARVAE 1.8 2.2 26 3.0 3.4 CARAPACE LENGTH (MM) Fig. 4. Relationship between number of larvae and carapace length for Rhopalophthalmus tattersallae in the Khor al Sabiya, Kuwait, September 1981-1982. 25 20 ) i) =a] = < 5 0 JUN_DEC 0) M@ JAN_MAY 23 45 6 7 8 9 1011 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 NUMBER OF LARVAE VOLUME 102, NUMBER 3 Brood sizes were generally <10 larvae/fe- male with only 6.6% carrying 20 or more larvae. Larger females and broods occurred from January through May and the smaller females and broods from July through Sep- tember (Fig. 5). This is consistent with ob- servations of other species of subtropical mysids which produce several generations per year (Mauchline 1980). Male: female sex ratios were near 1:1 on most dates, but sample sizes were often small. Greatest departures from 1:1 were on 11 January (0.5:1), 27 July (0.3:1) and 18 August (2:1). An unidentified dajid isopod was at- tached to the first pleonite of the 13 mysids collected. Parasitized mysids ranged in size from 1.0 to 2.2 mm CL. None of the other seven genera of mysids identified from Ku- wait Bay and the Khor al Sabiya during this study (Afromysis?, Dioptromysis/Kaino- matomysis?, Erythrops spp., Mysidopsis, Proneomysis and Siriella) were found par- asitized. Most of these other taxa were quite rare with Proneomysis and Siriella the only other genera that were locally abundant. Acknowledgments Appreciation is extended to U. Jayusi of Dames & Moore for his support and to T. E. Bowman for verifying the identity of R. tattersallae and identifying the other mysid genera. Literature Cited Dames, & Moore. 1983. Aquatic Biology Investiga- tions for Sabiya Area, Kuwait Bay and Devel- opment of Electrical Networks. Prepared for: Government of Kuwait Ministry of Electricity and Water. —_— Fig’ 3: September 1981-1982. qa Hodge, D. 1963. The distribution and ecology of the mysids in the Brisbane River.— University of Queensland Paper, Department of Zoology II: 90-104. Hulburt, E. M. 1957. The distribution of Neomysis americana in the estuary of the Delaware Riv- er.— Limnology and Oceanography 2:1-11. Mauchline, J. 1970. The biology of Schistomysis or- nata [Crustacea, Mysidacea].—Journal of the Marine Biological Association U.K. 50:169-175. 1980. The Biology of Mysids and Euphau- siids.—Advances in Marine Biology, J. H. S. Blaxter, F. S. Russell and M. Yonge, eds., 18: 1-677. Academic Press. London. —., & M. Murano. 1977. World list of the Mys- idacea, Crustacea.—Journal of Tokyo Univer- sity of Fisheries 64:39-88. Pillai, N. K. 1961. Additions to the Mysidacea of Kerala.—Bulletin of the Research Institute, University of Travancore 8:15-35. 1965. A review of the work on the shallow- water Mysidacea of the Indian waters.—Pro- ceedings of the Symposium on Crustacea Held at Ernaklam from January 12 to 15, 1955, 5:1681-1728. Marine Biological Association of India, Mandapam Camp. 1973. Mysidacea of the Indian Ocean.— Handbook to the International Zooplankton Collections 4:1-125. Indian Ocean Biological Centre. Kerala State, India. Sokal, R. R., & R. J. Rohlf. 1981. Francisco, W. H. Freeman. Tattersall, O. S. 1957. Report on a small collection of Mysidacea from the Sierra Leone estuary to- gether with a survey of the genus Rhopaloph- thalmus Illig and a description of a new species of Tenagomysis from Lagos, Nigeria. — Proceed- ings of the Zoological Society of London 129: 81-128. Wooldridge, T., & T. Erasmus. 1980. Utilization of tidal currents by estuarine zooplankton.—Es- tuarine, Coastal and Marine Science 11:107- 114. Biometry. San Collier County Pollution Control De- partment, 3301 Tamiami Trail East, Na- ples, Florida 33962-4977. Distribution of Rhopalophthalmus tattersallae brood sizes by season in the Khor al Sabiya, Kuwait, PROC. BIOL. SOC. WASH. 102(3), 1989, pp. 732-737 PYCNOGONIDA OF THE WESTERN PACIFIC ISLANDS VI. SERICOSURA COCHLEIFOVEA, A NEW HYDROTHERMAL VENT SPECIES FROM THE MARIANAS BACK-ARC BASIN C. Allan Child Abstract. —A new species, Sericosura cochleifovea, is described from sixteen specimens taken by the Research Submersible A/vin in 3660 m at the hydro- thermal Snail Pits Vent, Burke Field, in the Marianas Back-Arc Basin. The new species is compared with the two other known species of this genus and their distribution is discussed. The generic diagnosis is emended to include variation in palp segment numbers from seven to nine. There are only two reports (Child 1987, Turpaeva 1988) on Pycnogonida taken from deep sea hydrothermal vents. With the in- creasing efforts expended in finding and in- vestigating new vent fields, it is not sur- prising that additional pycnogonids have been and will be found. The new species described here was found during the explo- ration and sampling of recently discovered vent fields west of the Marianas Islands. These vent fields are part of a tectonic spreading zone investigated by scientists from Scripps Institution of Oceanography in April and May, 1987. The biological team of the group investigated three active vent sites along part of the spreading zone. Depths of the three sites varied from 3595 to 3660 m and are characterized by pillow basalts, hydrothermal mounds, vent chimneys, and many vent openings of both active and ex- tinct vents. Fauna is dense in and around the vents with “hairy” snails, brachyuran crabs, bresiliid shrimps, and white anem- ones as the dominant observable fauna (Hessler et al. 1988). The team collected a total of 17 speci- mens of a previously unknown pycnogonid from vent sites in the Snail Pits portion of Burke Field (16) and from the Alice Springs Field (1). Burke Field is dominated by dense aggregations of “hairy” snails that clog the vent openings. The temperature of the emerging water was 4—15°C, and the hot water venting from the openings was crystal clear. Water from Anemone Heaven vents nearby was cloudy. Alice Springs vent water was crystal clear. Family Ammotheidae Genus Sericosura Fry & Hedgpeth, 1969 Sericosura cochleifovea, new species Fig. 1 Material examined. — Marianas Back-Arc Basin, Burke Hydrothermal Vent Field, Snail Pits vent site, 18°19.9'N, 144°43.2’E, 3660 m, coll. R/V Alvin, Dive 1835, 26 Apr 1987 (one male with eggs, holotype, USNM 234505, one male with eggs, one male ju- venile, 4 female juveniles, paratypes, USNM 234506). Other material: Dive 1835 (two males with eggs, four males, two females, one ju- venile), Alice Springs Field, 18°12.6'N, 144°42.4'E, 3640 m, coll. R/V Alvin, Dive 1843, 4 May 1987 (one male juvenile). Description. —Size moderately small, leg span 13.1 mm. Trunk moderately slender, fully segmented, posterior rim of anterior three segments flared out in cowl-shape, without dorsomedian tubercles or setae. Neck short, expanded anteriorly at palp in- Fig. 1. Sericosura cochleifovea, holotype male: A, Trunk, dorsal view; B, Trunk, lateral view; > Palp; 19: Third leg, with cement gland tube enlarged; E, Oviger with several eggs attached; F, Oviger terminal segments, enlarged. Paratype female: G, Third leg. Paratype juvenile: H, Chelifore, enlarged. 133 734 sertion, without tubercles or setae. Ocular tubercle short, carried on elevated swelling at anterior of ocular segment, tubercle only slightly taller than basal diameter, rounded at tip, eyes lacking, sensory papillae prom- inent. Oviger implantation at posterior of neck, anterior to but not touching first pair of lateral processes. Lateral processes closely crowded, separated by half their diameters or less, 1.5 times longer than their maxi- mum diameters, armed with stout dorso- distal spine on each, two posterolateral spines on anterior two pairs, single postero- lateral spine on posterior two pairs, and an- terolateral spine on posterior three pairs, spines half as long as segment diameters. Proboscis long, massive, without constric- tions, carried horizontally, oral surface flat with slightly protruding lateral lips. Abdo- men slender, slightly swollen distally, ex- tending to midpoint of second coxae of fourth legs, with basal segmentation line, armed with two dorsal pairs of spines longer than segment diameter and pair of very short laterodistal setae. Chelifores short, robust, two-segmented. Scape only twice as long as maximum di- ameter, armed with five to six short dor- sodistal and laterodistal setae. Chelae small, bulbous, with scant trace of vestigial finger, without setae. Palps nine-segmented, armed with few setae longer than segment diameter proxi- mally, setae increasing in numbers on distal segments. Fourth segment only 0.75 length of second, third only slightly longer than fifth, distal four segments not longer than wide. Second and fourth segments slightly inflated distally. Oviger second and fourth segments sub- equal, second through fifth segments armed with several recurved spines, fourth and fifth with few lateral setae, sixth with 3 endal recurved spines and field of 17—18 ectal se- tae longer than segment diameter, seventh with 3-4 similar setae, 8th with single seta. Short, finely serrate denticulate spines on terminal three segments in the formula 1:1: PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 2. Eggs carried in round aggregations, size only slightly less than adjacent oviger di- ameter. Legs moderately long, very setose. Setae arranged in dorsal, lateral, and ventral rows, some dorsal setae longer than segment di- ameter, others about as long as segment di- ameter, setae increasing in numbers distal- ly. Longer dorsal setae arising from low tubercles. Second coxae almost equal in length to first and third combined. Femorae equal in length to first tibiae, second tibiae slightly shorter. Single femoral cement gland at extreme proximal end of segment, pro- truding as small bulge on anterior surface with a syringe-shaped tube as long as the segment diameter carried pointing dorsally. Bulge and tube invisible from posterior of leg. Tarsus very short, subtriangular, pro- podus slender, slightly curved, almost five times length of tarsus, armed with dorsal and lateral setae similar to those of tibiae. Sole armed with seven to eight short spines of similar size. Claw robust, half propodal length, moderately curved, auxiliaries about 0.7 length and with same curve as main claw. Sex pores on second coxae of posterior 4 legs. Female and juvenile paratypes: female slightly larger in most measurements. Leg setation extremely dimorphic. Coxae with few short setae, femur with two long lateral setae per side, setae twice segment diameter, and single long dorsodistal seta. Tibial setae few on dorsal and ventral surfaces, row of seven extremely long lateral setae per side, up to five times segment diameters, pro- podus with three long lateral setae per side. Main and auxiliary claws slightly longer in relation to propodus than those of male. Sexual pores not evident on subadult fe- males. Juvenile and subadult females with fully developed small chelae without teeth on scape of slightly smaller size than that of adult males. Measurements. — Holotype, in mm: Trunk length (chelifore insertion to tip 4th lateral processes), 1.65; trunk width (across 2nd VOLUME 102, NUMBER 3 lateral processes), 1.04; proboscis length, 1.31; abdomen length, 0.72; third leg, coxa 1, 0.3; coxa 2, 0.64; coxa 3, 0.44; femur, 1.22; tibia 1, 1.22; tibia 2, 1.11; tarsus, 0.14; propodus, 0.65; claw, 0.32. Distribution. — Known from the type lo- cality, Snail Pits Vent in Burke Hydrother- mal Vent Field, Marianas Back-Arc Basin, in 3660 m, and from Alice Springs Field in 3640 m. Etymology.—The specific name is Latin (cochlea = snail, and fovea = pit) and refers to the collecting site. Remarks.—This species is very closely related to another north Pacific species re- cently described, Sericosura venticola Child. The two species would be synonymous were it not for a set of small but taxonomically important differences which serve to sepa- rate them. Each of the differences taken in- dividually would not be sufficient to des- ignate this species as a new taxon, but the set of small differences in total are enough in my opinion. The differences in this new species are: a palp of nine segments which have not co- alesced into the seven of S. venticola, a much longer syringe-shaped cement gland tube, a slightly shorter neck with the oviger im- plantation slightly more anterior, dorsodis- tal spines on the lateral processes and pos- terolateral spines placed more proximally, a shorter abdomen bearing a different spine arrangement, a much shorter fourth palp segment, a longer fourth oviger segment in relation to the second and many more long setae and a different denticulate spine ar- rangement on the terminal segments, dif- ferent coxal length ratios and many less ven- tral setae on the third coxae and proximal femorae of the holotype, and different major leg segment ratios (femur = first tibia in this species while femur = second tibia in S. venticola). The new species is only half the size of S. venticola. While this fact is of little or no value itself in separating species, it contributes to the suite of differences which determine this new species. 735 The chelate subadult females of the type lot contribute to the known sexual dimor- phism feature of this genus. The first species known in this genus, Sericosura mitrata (Gordon), is also quite closely related to the two other species and has sexually dimor- phic features best seen in the legs, as in the new species. Gordon’s species has male legs with relatively few long dorsal and lateral setae while the female legs have many short ventral spines or setae on the major seg- ments along with many very long slender ventral setae on the tibiae. The new species male legs have many dorsal, lateral, and ventral setae of various lengths while the female legs have far fewer of these setae while having many extremely long lateral setae not found on legs of the male. The legs of S. venticola have a ventral field of many moderately long setae on the third coxae and these extend to the proximoventral fe- mur opposite the dorsolateral cement gland. Unfortunately, the female of the latter species remains unknown along with what- ever dimorphism exists for this species. Discussion The three species of Sericosura present an often encountered distributional problem among the pycnogonids. The first species to be described, S. mitrata (Gordon 1944:54— 57, figs. 19a—e, 22b), was found on the coast of Antarctica in slightly over 200 meters, and was subsequently found on the Walvis Ridge off southern Africa in well over 2000 meters (Child 1982:19-21, fig. 6). The sec- ond known species, S. venticola Child (1987: 896-899, fig. 2; Scipiolus thermophilus Tur- paeva=), is from the Juan de Fuca Ridge hydrothermal vent fields in depths of slight- ly more than 2200 meters. This new species, S. cochleifovea, is from the opposite side of the Pacific at the Marianas Back-Arc Basin in the deepest waters known for the genus, 3660 m. It would be convenient to state that the genus is hydrothermal vent-related ex- cept that we know nothing about the two 736 collecting localities for S. mitrata. No hy- drothermal activity is known for the Walvis Ridge locality but it would be surprising to find hydrothermal vent activity in less than 300 meters of depth off the coast of Ant- arctica. The three known species are found in widely disparate localities suggesting that the genus is worldwide in distribution but the species undoubtedly are much more re- stricted. as with almost all pycnogonids, to specific areas and are in most cases asso- ciated with zones of tectonic spreading hav- ing hydrothermal vent fields. The other anomaly among the three species is the discovery of this new species bearing nine palp segments. The other two species have seven with the three distal seg- ments now appearing to have coalesced from a larger number, presumably nine. Other genera such as Achelia, Tanystylum, Am- mothea, and others, have groups of species with palp segments varying in segment numbers, so the feature is not a new dis- covery except in this genus. The generic di- agnosis therefore needs to be revised to in- clude palps having seven or nine segments instead of the previously diagnosed seven. The genus Ammothea, from which this genus presumably split, has palps of eight or nine segments. This seven and nine palp segment character of Sericosura places it nearer the diagnosis of Ammothea. The only remaining major differences between these genera are the lack of a row of dorsomedian tubercles on the trunk of Sericosura species and the shape and placement of the cement glands and tubes. Most of the species of Ammothea have conspicuous dorsomedian tubercles on the posterior trunk segment ridges while none of the three Sericosura species are known to have these. The ce- ment glands and tubes in Ammothea, where known, are placed at the dorsodistal tip, or nearly so, of the femorae and are incon- spicuous. The fact that the cement gland and its conspicuous bulge and tube is placed proximally on the anterior of the femorae in Sericosura is probably a sufficient reason PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON to maintain the genus as separate from Am- mothea. The leg setae dimorphism and oth- er lesser characters only reinforce the sep- aration of this genus from the closely related Ammothea. Acknowledgments I am grateful to Dr. Robert R. Hessler, Scripps Institution of Oceanography, La Jolla, California, for bringing the specimens to my attention, supplying excellent data concerning their collection, and for donat- ing the specimens to the National Museum collections. Investigations of the Marianas Back-Arc Basin and the collection of spec- imens by Dr. Hessler and Dr. Harmon Craig were supported by NSF grant OCE83- 11258, for which appreciation is herein ex- pressed. Seven type specimens are deposited in the National Museum of Natural History. Smithsonian Institution, Washington, D.C. 20560, under the catalog numbers of the old U.S. National Museum. The other speci- mens have been returned to the Scripps In- stitution of Oceanography, deposited in the Los Angeles County Museum, California, the National Museum, Paris, and several added to the non-type collections at the Na- tional Museum of Natural History, Wash- ington, D.C. Literature Cited Child, C. A. 1982. Deep-sea Pycnogonida from the North and South Atlantic Basins.—Smithsoni- an Contributions to Zoology 349:1-iv, 1-54, 15 figs. 1987. Ammothea verenae and Sericosura venticola, two new hydrothermal vent-associ- ated pycnogonids from the Northeast Pacific. — Proceedings of the Biological Society of Wash- ington 100(4):892-901, 2 figs. Fry, W. G., & J. W. Hedgpeth. 1969. Pycnogonida, 1 Colossendeidae, Pycnogonidae, Endeidae, Ammotheidae. The fauna of the Ross Sea, Part 7.—New Zealand Oceanographic Institute Memoir No. 49, New Zealand Department of Scientific and Industrial Research Bulletin 198: 1-139, 206 figs. VOLUME 102, NUMBER 3 737 Gordon, I. 1944. Pycnogonida.—B.A.N.Z.—Antarc- [sic] in hydrothermal fauna.—Zoologiceski tic Research Expedition 1929-1931 Reports, Zhurnal 67(6):950—953. 2 figs. Series B (Zoology and Botany) 5(1):1—72. 27 figs. ee ee Pe ee ae ey Department of Invertebrate Zoology, Na- terns on the ocean floor.—New Scientist : 117(1605):47-51, 9 figs. tional Museum of Natural History, Smith- Turpaeva, E. P. sonian Institution, Washington, D.C. 20560. 1988. The finding of Picnogonida PROC. BIOL. SOC. WASH. 102(3), 1989, pp. 738-741 A NEW SPECIES OF THE CAMBARINCOLID GENUS SATHODRILUS FROM MISSOURI, WITH THE PROPOSAL OF A REPLACEMENT NAME FOR ADENODRILUS HOLT, 1977 (CLITELLATA: BRANCHIOBDELLIDA) Perry C. Holt Abstract. —Sathodrilus nigrofluvius is described and illustrated. Its relation- ships, distribution and possibly primitive status within the genus are described. Uglukodrilus 1s proposed as a replacement for the preoccupied name Adenodri- lus Holt, 1977. The description of a new branchiobdel- lidan presented below is offered as a part of an ongoing effort to describe the members of the genera Oedipodrilus Holt, 1967, and Sathodrilus Holt, 1968, from the southern Appalachian and Ozarkian regions of the southeastern United States. In addition, this opportunity is taken to propose a replace- ment name for Adenodrilus Holt, 1977, a junior homonym. Sathodrilus nigrofluvius, new species Fig. | Type specimens. —Holotype, USNM 118199, and three paratypes, USNM 118200-118202, taken from a tributary of the Black River, 2 mi NE of Lesterville, Reynolds County, Missouri, on State Road 21, on unknown host, 22 Aug 1961, by Per- ry CAO: Diagnosis. —Slender, small worms (ho- lotype 1.7 mm in length); dorsal ridge on segment VIII; lips entire; no oral papillae; jaws slight, triangular in lateral aspect, very light in color, dental formula (?) 5/4; one prominent pharyngeal sulcus, no corre- sponding exterior one; bursa large, ap- proaching diameter of segment VI in length, penial sheath greater in diameter than atrial region, penis a straight, cuticular, eversible tube; ejaculatory duct short, slender, thin- walled; spermiducal gland without prostate or prostatic protuberance or deferent lobes; spermatheca with thick ectal duct, median bulb, ental process. Etymology. —Latin, Black River. Description. —The members of Sathodri- lus nigrofluvius are small and relatively slen- der worms. The holotype and four para- types have the following mean dimensions: total length, 1.6 mm; greatest diameter, 0.3 mm; head length, 0.3 mm; head diameter, 0.2 mm; diameter, segment I, 0.2 mm; di- ameter, sucker, 0.2 mm. The lips are entire and there are no oral papillae. The eighth body segment bears a low, but distinct dorsal ridge, the others lack dorsal supernumerary muscles. The head tapers slightly towards the peristomium. The clitellum, on segments VI and VII is distinct but not prominent. The anterior nephridio- pore opens dorsally on the anterior margin of segment III. The jaws are small, about one twenty-fifth that of the head in length, delicate, light in color. The dental formula appears to be the common one of 5/4, but the teeth are small, uncolored and difficult to detect. The pau- city of material (the types) makes it inad- visable to destroy it in order to verify this point. The gut contents consist of detritus and diatoms. VOLUME 102, NUMBER 3 Fig. 1. 39 D Sathodrilus nigrofluvius: A, Lateral view of holotype; B, Lateral view of reproductive systems of holotype; C, Optical section of bursa; D, Lateral view of jaws. Abbreviations: af, atrial fold; ba, bursal atrium; ed, ectal duct of spermatheca; ejd, ejaculatory duct; ep, ental process of spermatheca; p, penis; ps, penial sheath of bursa; sb, bulb of spermatheca; spg, spermaducal gland. The spermiducal gland lacks deferent lobes, is about two-thirds the diameter of segment VI in length and three-eighths its own length in diameter and is slightly ta- pered at each end. It lacks even the rudi- ment of a prostate, and often lies longitu- dinally above the gut. The ejaculatory duct is a short, thin- walled, obscure tube that is more nearly in- ferred than seen in the available specimens. In one paratype it appears to be greatly ex- panded and the penial sheath collapsed at the place of its entry into the latter. Since serial sections are unavailable, it is possible, but unlikely, that the spermiducal gland opens directly into the penial sheath region of the bursa. The bursa is a cylindrical sac, subequal to the body diameter in length. About one- third of its length consists of the atrium and atrial fold with a greatly reduced lumen. The penis is a straight, cuticular tube attached by relatively thick strands to the inner wall of the penial sheath which is set off exter- nally from the bursal atrium by an encir- cling constriction (Fig. 1C). The spermatheca is characterized by a thick, muscular, irregularly bent ectal duct 740 that constitutes at least one-half the total length of the organ, a relatively short sper- mathecal bulb that is no greater in diameter than the ectal duct and a short, narrowed ental process (Fig. 1B). Variations. —In one specimen the antero- dorsal quadrat of the bursa is collapsed and the ejaculatory duct may be greatly expand- ed. Otherwise, no differences were noted. Affinities. —Among the species presently assigned to the genus Sathodrilus, S. vera- cruzicus Holt, 1968; S. hortoni Holt, 1973; S. okaloosae Holt, 1973; S. shastae Holt, 1981; and S. nigrofluvius lack a prostate or rudiment thereof (1.e. how many do have a prostate; how big is this genus?). Of this coherent group, S. nigrofluvius most nearly resembles S. veracruZicus. Unlike S. nigrofluvius, S. veracruzicus lacks dorsal ridges, appears to have oral pa- pillae, and has a proportionately shorter and more slender spermiducal gland, a longer and more slender bursa, a penis that is looped (and hence longer than its sheath), and a spermatheca with an ectal duct that is less in diameter than the spermathecal bulb and lacks an ental process. Sathodrilus hortoni consists of larger worms with a parasitic mien: a thin body wall and a gut filled with “globules of fat.” The “‘oesophagus”’ is attached by strands of muscle to the body wall of segments I and II (Holt 1973:97—98). The upper lip is lobed. The secondary reproductive organs are un- usual: the male efferent apparatus is pro- portionately small, the spermiducal gland relatively long and slender; the spermatheca is composed of a large muscular spermathe- cal bursa and a slender spermatozoa storing “bulb”? with a thick muscular wall and no ental process. Sathodrilus okaloosae has low dorsal ridges on the body segments. The jaws are unusual: broad and thick with a “dental ridge”? bearing the teeth and a dental for- mula of doubtfully 3/4, possibly 1/4 (Holt 1973:101). The bursa is less than the body diameter in length. The spermatheca has a PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON long, slender ectal duct and a long ental pro- cess, both proportionately less in diameter than those of S. nigrofluvius. Sathodrilus shastae is composed of larger animals up to 4 mm in length with two pha- ryngeal sulci and no dorsal ridges. The jaws are prominent; the dental formula 1/1. The ejaculatory duct is long and thick and the spermiducal gland is remarkably long (about twice the body diameter in length) and slen- der. The bursa, including penial sheath and penis, is proportionately small. The ectal duct of the spermatheca is short, the ental process narrow, the median bulb fusiform and flattened between the gut and body wall. Host. — Unknown. Distribution. —Members of the genus are distributed widely over the continent, but it may be worthy of note that the most struc- turally similar relative of S. nigrofluvius is from southern Mexico (the state of Vera- cruz) with its other close relatives from Florida (two) and the Pacific northwest, a distribution that suggests, as does the ab- sence of a prostate, that these species are ‘“‘among the least dervied”’ within the genus. Material examined. —The types. Dr. Stuart R. Gelder has informed me that the name Adenodrilus which I proposed (Holt 1977) for a genus later (Holt 1986) assigned to the family Bdellodrilidae is a junior secondary homonym of Adenodrilus Chekanovskaya, 1959, based on a haplo- taxid oligochaete from central Asia. The genus-name Uglukodrilus is hereby proposed as a replacement name for Ade- nodrilus Holt, 1977. It is to be considered as masculine and is derived from that of the leader of a fictional band of Orcs (see Tol- kien, J. R. R. The Lord of the Rings, v. 2, 1954) whose feeding habits were considered suspect. Acknowledgments Drs. Horton H. Hobbs, Jr. and Brent D. Opell have read a first draft of this paper. VOLUME 102, NUMBER 3 As always, I am grateful. Also, I wish to thank Dr. Stuart R. Gelder for calling to my attention the homonymy of Adenodrilus Holt, 1977. Dr. Ernest R. Stout, Head, De- partment of Biology, Virginia Polytechnic Institute and State University, and his sec- retaries have been unsparing in their sup- port and help of which I am greatly appre- Clative. Literature Cited Chekanovskaya, O. V. 1959. On Oligochaeta from the bodies of water in Central Asia (Ferghana Valley and the River Nurgab).—Zoologicheskii Zhurnal 38:1151-1162, figs. 1-5. Holt, Perry C. 1967. Oedipodrilus oedipus, n. g., n. sp. (Annelida: Clitellata: Branchiobdellida).— Transactions of the American Microscopical Society 86(1):58-—60, figs. 1-4. . 1968. New genera and species of branchiob- dellid worms (Annelida: Clitellata).— Proceed- ings of the Biological Society of Washington 81: 291-318, figs. 1-9. 741 1973. Epigean branchiobdellids (Annelida: Clitellata) from Florida. — Proceedings of the Bi- ological Society of Washington 86(7):79-104, figs. 1-8. . 1977. A gill-inhabiting new genus and species of the Branchiobdellida (Annelida: Clitellata). — Proceedings of the Biological Society of Washington 90(3):726-—734, figs. 1-5. 1981. New species of Sathodrilus Holt, 1968 from the Pacific drainage of the United States, with the synonymy of Sathodrilus virgiliae Holt, 1977.— Proceedings of the Biological Society of Washington 94(3):848-862, figs. 1-3. 1986. Newly established families of the order Branchiobdellida (Annelida: Clitellata) with a synopsis of the genera.— Proceedings of the Bi- ological Society of Washington 99(4):676-702, figs. 1-20. Department of Biology, Virginia Poly- technic Institute and State University, Blacksburg, Virginia 24061. Mailing ad- dress: 1308 Crestview Drive, Blacksburg, Virginia 24060. PROC. BIOL. SOC. WASH. 102(3), 1989, pp. 742-752 THE SECOND ANNUAL RISER LECTURE: ECLECTICISM AND THE STUDY OF POLYCHAETES Kristian Fauchald Abstract. —The study of polychaetes has involved two very different research programs: the morphological and systematic descriptions on one hand and the biological and physiological traditions on the other hand. The two traditions each represent two systems of two different approaches to the study of nature: on one hand structural versus procedural studies and on the other hand process- oriented versus taxon-oriented studies. None of the paradigmatic approaches common in biology (e.g., ecological, physiological, genetic or evolutionary) can solve their own problems by using only one of the four approaches. Under- standing the biology of the group can only come from a carefully managed eclectic approach to the study of the group. During the early part of my career the theory of science always appeared to exist totally independent of what I was doing as a biologist: Biology was something to be done, not thought about. A paper published about 25 years ago (Platt 1964) demonstrat- ed that I had been very wrong; the quality of a study depends crucially on the manner in which it is planned and performed. Since then I have examined my own and my fellow workers output for signs of an awareness of theoretical issues associated with the study of biology. I have concen- trated on the polychaete literature with which I am most familiar. By now more than 200 years worth of papers on poly- chaete morphology, systematics, phyloge- ny, physiology and ecology have accumu- The Riser Lecture Series.—In 1985 the annual Riser Lecture was initiated by members, alumni and friends of the Marine Science Center, Northeastern University at Nahant, Massachusetts. The occasion was the official retirement of Professor Nathan W. Riser. As teacher, biologist and founder of the facility, ““Pete’’ Riser en- dowed the laboratory with a legacy—the importance of considering the whole organism regardless of one’s special focus. We dedicate these annual lectures to that principle. lated, representing more than 10,000 individual papers and books. In this paper I will review, very briefly, the development of the study of poly- chaetes. I will then attempt to put this over- view into a minimal theoretical context. The results are some rather trivial admonitions. I believe these recommendations to be worthwhile because most of my colleagues still behave as if their activities were theory- independent. If I can set them thinking about these issues, then the purpose of this paper will have been fulfilled. Early Studies of Polychaetes Aristoteles reported what might be inter- preted as scaleworms in the ocean; Pliny the Older gave a much more convincing de- scription of ““marine scolopenders”’ (Gillet 1988) and this latter report was expanded on by both Rondelet and Gesner (Williams 1851); these “‘scolopenders”’ have tradition- ally been identified as nereidid polychaetes. For all practical purposes polychaetes were first described in 1758 in the 10th edition of Linnaeus Systema Naturae. These early reports and the transition into a scientific VOLUME 102, NUMBER 3 study of the polychaetes is described by Gil- let (1988). Names and descriptions.—The Linnean nomenclature separated names as labels from descriptions and definitions of the or- ganisms studied. For the first time logical procedures known since antiquity could be applied to the description of the living world. One could name an organism and define that label by descriptive terms, independent of the names themselves. The process has been taught as part of introductory classes in logic for a long time, nevertheless, the importance of this first application to bi- ology was overwhelming. The new nomen- clatural system made possible intelligible discourse about Nature in a way that no other device, before or after, has done. The practices of the scientists of the period re- flected an awareness of the different lan- guage levels involved in descriptive pro- cesses (Popper 1979). For example, I believe that the use of names of gods and goddesses for genera of various organisms reflects an awareness of the importance of the sepa- ration of names from definitions and de- scriptions. The trivial names, what we now call the species names, often were simple mnemonics: Nereis virens for example: the green nereid. Nereis diversicolor is another example of this naming tradition. The descriptions and definitions included morphological features. Microscopes were SO primitive that not much more than gross morphological features could be distin- guished. However, early illustrations may be remarkably accurate and detailed. Writ- ten descriptions uniformly are far less de- tailed. The early zoologists did exactly what we do: Include sufficient detail to distin- guish new taxa from previously known ones. One can hardly blame Linnaeus and his contemporaries for not appreciating how many different kinds of worms would even- tually be found, or for not developing the complete terminology for describing their wealth of morphological detail. The first major describers of polychaetes were Danes, 743 Otto Friedrich Muller (Muller 1776) and Otto Fabricius (Fabricius 1780), Russians, such as Peter Paul Pallas (Pallas 1766) and by the turn of the century the famous French scientists Cuvier, Lamarck, and Savigny. Reviews and classifications. —Lamarck and Cuvier, independently and in compe- tition, reviewed all polychaetes described, sorted out, and named a whole series of new higher taxa, especially genera and families (Lamarck 1816, Cuvier 1817). Another fa- mous French worker, Savigny, had made most of the new observations and descrip- tions. He was a careful observer with a fine eye for finding differences among similar forms (Savigny 1820). Lamarck added con- siderably to our understanding of the rela- tionships among the polychaetes. Also his separation of the polychaetes into two ma- jor groups, those with red blood and those with white blood, revealed an interest in physiological properties of the organisms. Nevertheless, more of Cuvier’s morpholo- gy-based system has been retained than of Lamarck’s. Detailed descriptions of newly discovered polychaetes became divorced from the time in which they were penned. The descrip- tions have increased in detail and length from one or two lines to several printed pages, but we still use most of the termi- nology and the overall pattern of descrip- tions established by Audouin and Milne Ed- wards in a study of the French fauna in the early 1830’s (summarized in Audouin & Milne Edwards 1834). The system used by Audouin and Milne Edwards closely resembled the Cuvierian system and formed the base for all workers over the next 20 years. By 1850 however, the emphasis of exploration shifted to Ger- many: Adolph-Eduard Grube (1850) issued a major review of the polychaete families and this paper was the standard for the next 15 years. Two scientists working in Stockholm made the next major advances in the mid 1860’s. Kinberg reported on his worldwide 744 travels and Malmgren detailed the North Atlantic and Arctic Ocean faunas. These two scientists represent two very different ap- proaches to descriptive science. Kinberg briefly described species collected on the cruise of the Eugenie around the globe and added numerous new taxa at all levels (Kin- berg 1865, 1910). Malmgren’s (1867) stud- ies were intensive; he focussed his attention on a much smaller area and carefully re- viewed all previous work before commit- ting himself to describing a new taxon. This difference in approach closely matches a pe- rennial difference among descriptive biol- ogists; among modern systematists Gesa Hartmann-Schroder and Olga Hartman both have used Kinberg’s approach, where- as Marian H. Pettibone more closely match- es Malmgren. I have done a bit of both. Kinberg and especially Malmgren did their best to increase the consistency in use of terms and in the amount of detail re- quired for adequate descriptions. Quatre- fages (1866) issued a large-scale review of the whole annelid fauna as he knew it. Per- haps more pedestrian a systematist than the others mentioned, he nevertheless became extremely influential, due in part I believe to his location: he was in Paris, and had a long history of publications on polychaetes by the time he issued his magnum opus. Kinberg had published a few earlier papers, but neither he nor Malmgren ever issued any additional major contributions to the study of polychaetes. They both left science shortly after the papers mentioned were published. Ludwig Schmarda is one of the more col- orful persons in the history of polychaete studies. He travelled around the world in the 1850’s, not in an exploring vessel, but by hitch-hiking on commercial sailing ves- sels. His description of his trip from South Africa to Australia is singularly harrowing, including very bad weather, seasickness, scurvy and assorted other diseases. In Chile he lost his collections to a fire on board; in Panama he was robbed by some rather un- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON savory characters who made their living by preying on people going from the U.S. east coast to the west coast via the Isthmus. De- spite the loss of his collections, he published a large report (Schmarda 1861) that appar- ently was largely overlooked by his contem- poraries. This was probably in part due to the increasing standards of descriptions and illustrations. Schmarda’s effort was, how- ever, the earliest worldwide tropical survey of polychaetes. He described a large number of new species for which there are few types available and poor locality information. At that time, there was no requirement that types should be deposited anywhere: De- scriptions were considered adequate evi- dence for the presence of a new taxon. How- ever, the first Nomenclature Code, and perhaps just as importantly, the first volume of Zoological Record, was issued in 1864. The morphological tradition. —The mor- phological tradition, outlined above, has continued through the work of McIntosh (1885), Fauvel (1923, 1927), and Augener (1918), and is now followed by most prac- ticing systematists. The total focus of this tradition is very limited in the kind of evi- dence deemed acceptable. Most system- atists will accept only features that can be seen either with the naked eye or with stereo or compound microscopes as valid taxo- nomic characters. Furthermore, a tradition among polychaete systematists suggests that all reasonably well preserved specimens, es- pecially anterior ends, should be identifiable to species. I have more than once heard complaints from well known systematists that a published description was too difficult to use, or was impractical, because it used information not readily available using minimal technical equipment, or required the presence of complete specimens. This tradition is clearly at odds with, for exam- ple, students of isopod crustaceans who for years have accepted limits on the identifi- ability of all specimens. The biological tradition.— Another tra- dition in the study of polychaetes dates back VOLUME 102, NUMBER 3 to about 1850. Thomas Williams (1851) published a major review of the biology and physiology of the polychaetes. This sum- mary is now rarely quoted; it has been su- perceded by more recent reviews, but it was important historically because Williams re- viewed all information available about the life of all worms known to science. Some of the data quoted by Williams date back to Lamarck and are speculative rather than ob- servational in nature and some rather quaint notions were paraded only eight years be- fore the publication of Darwin’s Origin of Species. Williams made some original phys- iological observations on various English polychaetes. The most impressive of the early poly- chaete biologists was Eduard Claparéde, a rather tubercular-looking Swiss, who did most of his work in France and Italy (Cla- paréde 1854). By 1865 he had gotten into a rather virulent quarrel with Quatrefages over all of Quatrefages’ new taxa, defined in many cases without access to any mate- rial (Quatrefages 1865a, Claparéde 1865, Quatrefages 1865b). Claparéde emphasized the importance of observations on live or- ganisms; Quatrefages by that time had be- come very collections-oriented. This differ- ence in approach formed the background for the disagreement. Claparéde, true to his principle, deposited no specimens in any museum, making many of his new taxa dif- ficult to define accurately. The second tradition was biological in na- ture: studying live organisms and making observations of the live processes, such as reproduction, development and feeding. These kinds of observations were difficult to quantify in an age of poor mechanical recording devices, no photography to speak of, and certainly no electronic recording de- vices. Additionally, statistics had not yet developed to the point where repeated sam- ples were taken. The studies were therefore often episodic in nature, and observations were only rarely organized into tables or other means of presenting large, easily sur- 745 veyed data. The kinds of observations at- tempted by Claparéde are still difficult to document for theoretical reasons that I will touch on below. Claparéde combined his studies of live organisms with a detailed study of microan- atomical structures. These studies are ex- cellent and are still the best starting point for any anatomical studies in the groups he covered. Claparéde’s illustrations are among the best ever published on polychaetes. The most important aspect of Claparédes work was that he demonstrated that a remarkable amount of information could be gained by looking at live organisms. He also demon- strated that detailed anatomical and histo- logical studies yielded systematically dis- tributed information, which could be potentially useful in systematics. Ehlers tried to combine the two traditions in his massive publication “Die Borsten- wurmer’’ issued in two parts (Ehlers 1864— 1868). Some of his descriptions of new taxa run 10-15 printed pages, accompanied by one or two full packed plates of illustrations. Consequently, Ehlers succeeded in going through less than '4 of the then known poly- chaete taxa in roughly 700 pages of text, but for the groups he covered, his volume is absolutely indispensible. Ehlers’ research later devolved to thoroughly traditional, morphological descriptions. I can find no evidence in any of his publications that he attempted to complete the massive study he had started. The study of live polychaetes eventually developed into a tradition of physiological studies, based usually on members of rela- tively few families with highly characteris- tic, often unusual physiological patterns. These studies are often performed by pro- cess-oriented rather than by comparative scientists. Reproductive studies, while cov- ering in part members of most groups, have been focussed on eunicids, nereidids and syllids (Schroeder & Hermans 1975); stud- ies of respiratory and blood physiology on glycerids, terebellids and scattered other 746 groups (Dales 1969, Florkin 1969). Studies of regeneration have focussed on sabellids with few glances in other directions (Need- ham 1969). Genetic studies have been done on dorvilleids and little else (Akesson 1982). Neurophysiologists have studied the prop- erties of the giant nerve fibers in sabellids of the genus Myxicola with very little con- cern for the biology of the organism at all. There are about 80 families of polychaetes and of these at least 60 are common in shal- low water and relatively readily available; nevertheless live studies have focussed on a few popular groups and usually on only one or a few species in each group at that. The results of the biological and physio- logical studies have been very valuable, but less as a comparative study of polychaetes than as an exploration of various biological and physiological mechanisms. Theory and the Study of Polychaetes The rather conservative descriptive tra- dition continues among polychaete system- atists; for each advance in morphological or anatomical technique, traditionalists hang back, not wanting to get involved with new methods or add new features to the descrip- tions. Often the young turks among poly- chaetologists are traditionalists in the study of other groups of organisms, especially ver- tebrates. Very few of the scientists closely associated with the study of polychaetes have demonstrated strong theoretical interests. For example, it is difficult to find any ref- erence to evolution, or to Darwinian or anti- Darwinian thinking anywhere. Ehlers’ pub- lication from 1864-1868 gave no indication of a major revolution in biological thinking taking place at the time. McIntosh (1885) mentioned nothing about phylogeny in his treatment of the Challenger polychaetes. One outstanding exception is E. Meyer, who in his studies of polychaetes indicated a good, often anticipatory understanding of biological theory (Meyer 1890). This paper is frequently quoted in the literature on phy- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON logeny of the invertebrates, but not often by polychaete taxonomists. Some of the developmental biologists as- sociated with the study of spiral cleavage at Woods Hole Marine Biological Laboratory used polychaetes for their studies. These sci- entists had deep theoretical interests and showed great skill in using the polychaete material in clarifying theoretical problems (Wilson 1898, Treadwell 1901). The reason for the lack of theoretical and one might say scholarly interest in the study of polychaetes is relatively easily found. Most scientists published only a single pa- per on polychaetes and very few made the study of these animals their lifetime occu- pation (Reish 1958). Through about 1950, the study of polychaetes was a relatively lei- surely pursuit. Even in most early benthic ecology studies (Petersen 1911, Blegvad 1930), few polychaetes are mentioned or named, except to family. In morphological studies, the annelids were considered a step- ping stone to the arthropods (Hanstrom 1928, Binard & Jenner 1928, and the dis- cussion of the anterior nervous system of the annelids and arthropods) and thus of interest insofar as they showed the step-wise advance to the conditions present in the ar- thropods. Parenthetically, papers that treat polychaetes well from a theoretical point of view were, with few exceptions, written by scientists with a limited experience in the group (Hanstrom 1928, Hatschek 1893). This generalization is far less true today than it was before WWII. The rapid development of interest in ben- thic ecology following the publication of Thorson’s (1957) review of the topic lead to considerable change in attitude. Poly- chaetes have turned out to be extremely common in the marine benthos; benthos ecologists have changed their attitudes to- wards the importance of polychaetes with the mesh-size of their screens. Further, modern ecologists are aware that no ques- tions can be answered by studying only a few “‘representative’’ organisms, usually se- VOLUME 102, NUMBER 3 lected among “easily identified’’ organisms, such as some crustaceans, echinoderms and mollusks, as done in the early days of ben- thic ecology. Simple thoughts on theory. —Organisms may be studied in four different ways, which may be organized into two systems of two. First, one may either attempt to describe the structural characteristics of an organ- ism, or one may study interactions among structures in time or space. The other sys- tem of classifying observations describes the investigational intent. One may study the same process in a variety of organisms; or, alternatively, one may study a variety of processes and structures in the same kinds of organisms. Structural descriptions historically start- ed with external morphology, and pro- ceeded via internal anatomy to microscopic anatomy in all its phases. Structural de- scriptions deal with the material presence of anything, including atoms and subatomic particles. In gross morphological descrip- tions the unaided eye is used; all other de- scriptions are based on interpretation of im- ages created by various pieces of gear: microscopes of all kinds, meters and dials and color-reactions, spectrophotometers, or small patches of color on a starch gel. The more highly magnified the analysis be- comes, the more remote the interpretation of the findings become from normal human experiences, but, at least in theory, no dif- ferent from observations of gross morphol- ogy. In some sense, interpretation becomes easier with increasing magnification, since the higher magnification allows a far more precise use of language in describing limit- ing conditions than do observations of a morphological or anatomical nature. Natural historians and some physiolo- gists (a subgroup of the comparative and ecological physiologists) seek a completely different kind of information about organ- isms, information which we have had a great deal of difficulty entering into our structur- ally derived patterns. All organisms change 747 with time and all structural landmarks change in relation to each other during on- togenesis, presumably in an organized fash- ion, but not necessarily in the same pattern even in genetically similar organisms. In- formation derived from these changes is as much an expression of the genome of the organism as is the structural information. I am aware of the problems including this kind of information in our descriptions will create, but I believe that until we do, we will fall short of understanding the organisms we are studying. Computerized modelling may offer help in creating testable predictions for such studies. The other system of groupings of study is familiar to most scientists, especially in technically more complex fields. Scientists become experts on the use of a single tech- nique: transmission and scanning electron microscopes, enzyme electrophoresis, DNA hybridization and so forth and will inves- tigate the limits of what the technique can do. The results of this approach have been excellent and have lead to major advances in our understanding of microstructures and varlous processes. The other major way of looking at the organisms is as a specialist on a single an- imal group; a taxon-oriented person. Such a person may be eclectic in their use of tech- niques, but will rarely add to the develop- ment of new techniques. These biologists often have a better understanding of the evolutionary significance of differences in processes among the organisms studied than the process-oriented scientists, but are usu- ally rather parochial in their view of the world. A polychaete’s-eye view of the globe is limiting in many ways. These four ways of studying organisms do not agree with the traditional breakdown of specialities among biologists. Taxono- mists, while primarily concerned with de- scription of structure, frequently resort to adaptive explanations. Physiologists, while exploring functional issues, base themselves in knowledge of the structures involved in 748 the particular processes studied. Perhaps most confused are the activities that are now subsumed under the heading of ecology. In part, ecologists describe structure in their case patterns of distribution of organisms in nature, but usually use functional expla- nations for the patterns demonstrated. The separation of the two modes of thinking is not trivial, but is built into the language. Ideally a language describing structure should use only shape and position words; in practice we use such words as “‘bran- chiae”’ and “‘notopodial cirri.’’ For trained taxonomists and morphologists the usual meanings of these words have become triv- ial: they are using both words as shape and position markers. However, notopodial cir- ri, usually slender, often very long cirri pro- jecting from the dorsolateral sides of the worms, often appear to be as much respi- ratory as sensory in function. Eclecticism and the study of poly- chaetes.—Thus an adequate description of any polychaete would require a rather eclec- tic collection of pieces of information, both static and dynamic. Most structural descriptions of poly- chaetes now include a minimal mention of major morphological features. At least one species of most families have been studied anatomically, at least at the light micro- scope level. Very few truly comparative studies have been performed within each family. Comparative studies among the families are rather common, but without knowledge of how much variation to expect within each family, the interpretation of such comparative studies will always be difficult. Microanatomical studies are becoming rather more common, but again, with some very Salutary exceptions, have focussed more on the relations among the families. Other studies, with both structural and functional components, are mentioned below. Studies of comparative physiology have given us important information about the interactions among the structures, e.g., studies of mechanisms of respiration among polychaetes. However, most physiological PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON studies have been focussed more on eluci- dating process and are for that reason usu- ally not very useful for comparative pur- poses. Most life history studies published so far include an account of parts of the larval development and metamorphosis into a postlarvae, but little about the rest of the life of the organisms, including longevity (Fauchald 1983). The bits we have are in- teresting, but are insufficient for all species. I am advocating eclecticism because I be- lieve that this approach will force us to change our approach to our studies. Cur- rently we learn one, or perhaps a few, tech- niques and then proceed to apply these to all problems, whether the application can solve the problems posed or not. The in- vestigative technique and the detail sought must depend on the question asked, rather than the other way around. For example, it is not always useful or necessary to identify organisms to the species level in a benthic investigation. The first step in planning a study therefore must be to question the pur- pose of the investigation. If the purpose is an exploration of the area—a study of which organisms are present in what quantities— then identification to species is not only de- sirable, but the only way such information should be presented. But if the purpose is to investigate feeding biology or perhaps trophic structure, in addition to giving a listing of taxa present, at the very least as much effort must be put into investigation of gut contents and mechanisms of feeding, as into the identification of the specimens. Most investigators now identify their or- ganisms (more or less accurately) and then quote some authority for the other infor- mation needed, e.g., feeding physiology. For the polychaetes, most quote Fauchald & Ju- mars (1979), an inappropriate source of in- formation for this purpose. The Diet of Worms was written as a summary of what little information was available in the mid 1970’s and was intended to spur investi- gations: It has apparently done so, but suf- ficient information is still not available for any species to my knowledge. VOLUME 102, NUMBER 3 Polychaetes are valuable for a variety of studies. Polychaetes are ubiquitous and common in all marine environments. The numbers of polychaete taxa is large enough to allow the use of the statistical data re- duction, but is not as overwhelming as in some other groups. Most major subgroups have morphologically very strict body plans and can be identified to family by rank ty- ros. The group is very old (Fauchald 1984) and the major body plans were laid down a long time ago: We can in the polychaetes investigate current evolution of ancient body plans. For example, the eunicids are very uniform in general morphological appear- ance; in fact, the jaws have not changed much since Palaeozoic times. Nevertheless, a preliminary numerical study of about 300 individuals of approximately 12 species (Fauchald 1989) demonstrate several dif- ferent patterns of growth and of control of the body proportions, implying rather dif- ferent physiological properties, perhaps re- lated to the maximum absolute size of each species. The consequences of the studies of Gras- sle & Grassle (1976) and Eckelbarger & Grassle (1987), to mention only two of a series, are fascinating. They have given us a view of a worldwide group of small, ever- changing populations of capitellids becom- ing isolated, perhaps going extinct locally, perhaps meeting up again before, or after, completing a speciating process—in short, a complex mosaic. Chromosome studies of various poly- chaetes indicate that ploidy relations may play a more important part in evolution in polychaetes than previously expected; per- haps leading to a reconsideration of the im- portance of the various processes in the evo- lution of animals. An eclectic approach may thus complete the transformation of the study of poly- chaetes from an intellectual backwater to the forefront of biology. Some final notes. —I agree with my alter ego of 25+ years ago that theory of science 749 exists with little reference to what I do on a day to day basis. I have come to the re- alization that this is perhaps the way it ought to be. If the theory of science was strictly a description of what scientists do, then one could not expect discussion of normative rules. We all use theoretical constructs in even the simplest observations. The belief in theory-independent observations ap- pears now on the wane. Philosophers of sci- ence study and perhaps build into systems the theories behind our observations and make us as working scientists aware of these constructs. Without the precision in think- ing and data definition theory enforces, very little advance 1s possible. A significant fraction of current papers are routine descriptions of a few new taxa, usu- ally with a review paper as authority for the separate status of the new taxa; the mate- rial examined is minimal and comparison with types of previously described species is rare. If current theory and methods were applied to these studies, Iam convinced that the deluge of new taxa would slow down. Most of the new taxa are collected during quantitative investigations and the authors do not have the luxury of performing a com- plete and detailed review of the family or genus of interest before publishing a new taxon or two. Detailed and rigorously per- formed reviews of previously described taxa are lacking for nearly all polychaete families and very few are now on the horizon. Most of the investigations in which the bulk of new material is collected have poorly, or inappropriately defined, goals: however, one requirement runs through most of them: No matter what the stated purpose of the in- vestigation is, the organisms collected must be identified to species. This requirement forces the researches to make rapid, often incorrect decisions. A careful definition of study goals would leave both ecologists and polychaetologists happier and the few poly- chaetologists working full time on poly- chaete taxonomy less overwhelmed. There is little support for all the other kinds of studies needed to describe and study 750 polychaetes adequately. The result is that most of the polychaetologists are limping along, without being able to do even the necessary revisory work, and certainly with- out being able to apply theory or attempt to add truly new information to our de- scriptions of polychaetes. A rather sad con- clusion, but I believe one in which experts on other groups also would concur. Acknowledgments This paper is based on the second Riser Lecture, given at The Marine Science Cen- ter, Northeastern University, October 30, 1986. It is dedicated to Dr. Nathan W. Ris- er, one of the finest polychaete experts liv- ing; Pete Riser combines a morphologist’s eye with a focus on live organisms. I would like thank the committee, Drs. M. Patricia Morse, Joan Ferraris and Jon Norenburg for giving me the opportunity to give the lecture. Dr. Leonard P. Hirsch read and rejected several versions of the paper, for which I am now very grateful; what makes sense in this paper is due to my friends; what appears hare-brained is my own responsibility. Literature Cited Akesson, B. 1982. A life table study on three genetic strains of Ophryotrocha diadema (Polychaeta, Dorvilleidae).— International Journal of Inver- tebrate Reproduction 5:59-69. Audouin, J. V., & H. Milne Edwards. 1834. 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Report on the British Annelida. — D ; ice epartment of Invertebrate Zool - Report of the British Association for the Ad- Pp ate Zoology, Na vancement of Science 21:159-272. tonal Museum of Natural History, Smith- Wilson, E. B. 1898. Considerations on cell lineage SONaN Institution, Washington, D.C. 20560. and ancestral reminiscence, based on a reex- amination of some points in the early devel- PROC. BIOL. SOC. WASH. 102(3), 1989, pp. 753-760 A NEW SPECIES OF EUCHONE (POLYCHAETA: SABELLIDAE) FROM THE NORTHWEST ATLANTIC WITH COMMENTS ON ONTOGENETIC VARIABILITY R. Eugene Ruff and Betsy Brown Abstract. —Euchone banséi, a new species of the polychaete family Sabellidae, is described from the continental slope and rise between Cape Cod, Massa- chusetts, and Cape Lookout, North Carolina. Juvenile and adult specimens are examined and ontogenetic variability is discussed. It is demonstrated that the number of abdominal depression setigers, the shape of the collar, and the number of radioles are not valid diagnostic characters for the identification of juveniles. In response to interest in offshore oil and gas development, the Minerals Manage- ment Service (MMS) of the U.S. Depart- ment of the Interior sponsored research on benthic communities on the continental slope and rise (SO0—3000 m) off the eastern United States in three regions: (1) the U.S. North Atlantic near Georges Bank off Mas- sachusetts, (2) the U.S. Mid-Atlantic off New Jersey, and (3) the U.S. South Atlantic off the Carolinas. As is typical in marine soft- bottom environments (Knox 1977), the macrofaunal communities in these regions are dominated by polychaetous annelids (Maciolek et al. 1987a, b; Blake et al. 1987). Because of the numerous samples collected and the small sieve mesh (300 wm) used throughout this sampling program, juvenile growth stages of many polychaete species were routinely collected. This paper de- scribes a new species of Euchone (Poly- chaeta: Sabellidae) collected between 1345- 2495 m depth along the U.S. Atlantic coast and examines some of the ontogenetic vari- ability exhibited by this species. Euchone bansei, new species Figs./T."2 Euchone spp. Hartman & Fauchald, 1971: 179 [partim]. Euchone sp. 3. Maciolek et al., 1987a, b.— Blake et al., 1987. Material examined. —off Martha’s Vine- yard, 4 May 1966, Chain station Ch 103, 39°43.6'N, 70°37.4’W, 2022 m, 8 speci- mens; 7 Sep 1963, Atlantis station A 58, 3634.5 NY 12°55.0'W.,¥ 2000: 75 m3 specimens; near Baltimore Canyon, 19 May 1985, cruise MID-4 station 13-2, 37°53.29'N, 73°45.30'W, 1607 m, clayey mud, Holotype (USNM 115738); 19 May 1985, cruise MID-4 station 10-3, 37°51.73'N, 73°20.01'W, 2095 m, silty mud, 5 paratypes (USNM 115739); 16 Nov 1985, cruise MID-6 station 10-1, 37°51.77'N, 73°20.01'W, 2104 m, silty mud, 9 paratypes (BMNH ZB 1987.620-628); near Linden- koehl Canyon, 16 May 1985, cruise MID-4 station 3-2, 38°36.75'N, 72°51.57'W, 2055 m, silty mud, 6 paratypes (BMNH ZB 1987.629-634); 17 May 1985, cruise MID-4 Station 3-3, 38°36.75'N, 72°51.60’W, 2052 m, silty mud, 9 paratypes (USNM 115740); 17 May 1985, cruise MID-4 station 11-1, 38°40.10'N, 72°56.43'W, 1510 m, clayey mud, 22 paratypes (USNM 115741); 7 Aug 1985, cruise MID-5 station 12-2, 38°29.25'N, 72°42.22’W, 2495 m, sandy mud, 3 paratypes (USNM 115742); off Cape Cod, 25 Jul 1986, cruise NOR-6 station 754 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Euchone bansei (holotype, USNM 115738): A, Entire animal in lateral view showing the right half of branchial crown; B, Collar and anterior region with branchiae not illustrated; C, Posterior region showing the anal furrow. 3-1, 41°01.55'N, 66°20.12'W, 1345 m, silty m,sandy mud, 9 paratypes (USNM 115744); mud, | paratype (USNM 115743); near Ly- off Cape Lookout, 23 May 1985, cruise SA-4 donia Canyon, 29 Apr 1985, cruise NOR-2 | station 4-3, 34°11.29'N, 75°38.67'W, 2015 station 6-2, 40°05.03'N, 67°29.13’W, 2108 mm, silty mud, 2 paratypes (USNM 115745). VOLUME 102, NUMBER 3 recs RS — wt = *. CE eee cey eee Oe A eek =" ZT_S_S SS 0.05mm Fig. 2. Euchone bansei (paratype, BMNH ZB 1987.620): A, Thoracic notosetae from setiger three, including long and short limbate setae and a narrowly limbate bayonet seta; B, Thoracic neuroseta from setiger three; C, Uncini from abdominal setiger four in profile and in frontal view. Diagnosis. —Small Euchone species with 17 abdominal setigers, last 6 associated with anal depression. Branchial crown with 4 pairs of radioles united with palmate mem- brane for half their length; radioles with long, filiform, pinnule-free ends. Collar entire lat- erally and ventrally, separated by mid-dor- sal gap, of even height all around, extending to branchial basis. Ventral shields absent. Thoracic notopodia with long and short limbate setae and pointed bayonet setae. Abdominal uncini with quadrate base, small 755 756 main fang, and crest of numerous smaller teeth. Description. —Adult holotype 4.0 mm in length (excluding radioles), 0.5 mm in width along posterior thorax (Fig. 1A). Other adult specimens 2.2—5.9 mm in length, 0.3-0.8 mm in width; branchial crown contributing additional 4.4 mm in largest individuals. Four pairs of radioles connected for half their length by delicate palmate membrane; each radiole with about 20 subequal pin- nules alternating along axis and a pinnule- free filiform tip contributing up to half of total radiole length. Dorsal lips with elon- gate-triangular radiolar appendages, about one-half pinnule length, densely ciliated; pinnular appendages not observed. Paired ventral lips shorter, rounded. Peristomium produced into triangular projection ventral to radioles. Collar entire ventrally and of nearly equal height laterally and ventrally in adult (Fig. 1B), reaching approximately to base of branchiae; dorsal gap very narrow; otocysts not observed. First setiger one-half length and narrower than following thoracic setigers. Thin postsetal glandular girdle encircling second setiger. Ventral shields absent. Abdomen with 17 setigers, last 6 associ- ated with anal depression; wings of depres- sion flaring, connected anteriorly by thin membrane (Fig. 1C). Pygidium triangular, slightly longer than wide. First setiger with single bundle of long, narrow limbate setae and shorter bayonet setae; other thoracic notosetae of three kinds: superior arc of long limbate setae, and in- ferior group of short limbate setae and slen- der pointed bayonet setae with narrow wings (Fig. 2A). Thoracic neuropodia with about 12 long-handled acicular uncini in each fas- cicle (Fig. 2B). Abdominal notopodial tori with 10-17 uncini, each with quadrate base and small main fang surmounted by nu- merous rows of smaller teeth (Fig. 2C); shape of uncini nearly constant within each torus, but main fang progressively smaller in pos- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON terior setigers. Abdominal neuropodia with 3-4 narrowly limbate setae per fascicle. Sexes separate with gametes occurring laterally between dorsal and ventral longi- tudinal muscle bundles in posterior thoracic and anterior abdominal segments. Eggs ir- regular in outline, up to 0.1 mm in diameter. Sperm short-headed, with blunt acrosomes, approximately 5 um long excluding flagella. Methyl green stain in 70 percent ethanol (Banse 1970) readily accepted both dorsally and ventrally on most of body including pygidium and wings of anal depression. Staining cells absent in radioles, at anterior margin of collar, in glandular girdle, in in- tersegmental furrows, along fecal groove, around parapodia, and in narrow band of cells encircling each segment at level of se- tae. Distribution. —Euchone bansei is found from the southern flank of Georges Bank off Cape Cod, Massachusetts, southward to Cape Lookout, North Carolina. The species occurs in sandy to clayey muds in depths of 1345-2495 m and is found most often at about 2000 m in abundances occasionally exceeding 100/m7?. Etymology.—This species is named in honor of Dr. Karl Banse in recognition of his significant contributions to the knowl- edge and taxonomy of the Sabellidae. Ontogenetic Variability The routine use of 300-wm mesh sieves throughout the MMS sampling program re- sulted in the collection of a large number of sub-adult and juvenile specimens of Eu- chone bansei. After selection of the type ma- terial, an additional 185 specimens from across the sampling region were examined in detail. Ocular micrometer measurements were made on the total length from the top of the collar to the tip of the pygidium and on the width of the last thoracic setiger. Counts were made on the number of tho- racic and abdominal setigers, on the number VOLUME 102, NUMBER 3 6.0 2 i= D 5.0 = O Q