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VOLUME 102
18 OCTOBER 1989
ISSN 0006-324X
NUMBER 3
THE BIOLOGICAL SOCIETY OF WASHINGTON
1988-1989 Officers President: Kristian Fauchald Secretary: G. David Johnson President-elect: Leslie W. Knapp Treasurer: Don E. Wilson Elected Council
Gary R. Graves Meredith L. Jones
W. Ronald Heyer Raymond B. Manning
W. Duane Hope Wayne N. Mathis
Custodian of Publications: Austin B. Williams
PROCEEDINGS
Editor: C. Brian Robbins
Associate Editors Classical Languages: George C. Steyskal Invertebrates: Stephen D. Cairns Frank D. Ferrari Plants: David B. Lellinger Raymond B. Manning Insects: Wayne N. Mathis Vertebrates: G. David Johnson
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PROC. BIOL. SOC. WASH. 102(3), 1989, pp. 537-552
NEW SPECIES AND RECORDS OF BIRDS (AVES: MEGAPODIIDAE, COLUMBIDAE) FROM AN ARCHEOLOGICAL SITE ON LIFUKA, TONGA
David W. Steadman
Abstract. — A new species of megapode, Megapodius alimentum, is described from bones excavated at the Tongoleleka archeological site, Lifuka, Haapai Group, Tonga. Two coracoids from the same site are referred to Megapodius cf. molistructor and Caloenas cf. canacorum, both extinct species otherwise known from late Holocene fossils on New Caledonia. Another coracoid from Tongoleleka is referred to Ducula cf. david, an extinct species recently described from an archeological site on Wallis (Uvea) Island. A single tibiotarsus, too fragmentary to be named, represents an extinct, undescribed species of Ducula that is larger than any congener, living or extinct. The first four extinct species from Lifuka are more closely related to Melanesian species than to those of eastern Polynesia. The occurrence of five extinct species at the Tongoleleka site indicates that late Holocene losses of land birds in western Polynesia may have
been as severe as those in eastern Polynesia.
In 1984, Tom Dye of Yale University excavated a rich archeological deposit on Lifuka, Haapai Group, Tonga, known as the Tongoleleka Site. Situated in an ancient sand dune, the site consists of three strata (Layers II, III, and IV) that bear pottery. The lowest and oldest stratum is strati- graphic Layer IV, which mainly represents Cultural Unit III. Layer IV, buried by 1 to 1.5 m of more recent sand, yielded deco- rated Lapita pottery, believed on the basis of sites excavated elsewhere to date at 3500 to 3000 years B.P. (T. Dye, pers. comm.). Two radiocarbon dates on charcoal from the upper portion of the overlying strati- graphic Layer III are 2260 + 60 years B.P. (Beta-14171) and 1370 + 70 years B.P. (Beta-11243).
Among the 20 identifiable bird bones from Tongoleleka are 11 bones of land birds, all but two of which are from stratigraphic Lay- er IV. Six of these bones are from two species of extinct megapodes, four others are from three extinct and one extant species of col-
umbids, and one is from the extant starling Aplonis tabuensis. Also recovered were shellfish and bones of fish, reptiles, mam- mals, and marine birds. All of the bones probably represent food remains of early Tongans.
The purpose of this paper is to describe the morphology and systematics of the ex- tinct species of birds from Tongoleleka. Ad- ditional details of the chronological, strati- graphic, zoogeographic, and cultural con- texts of these extinct species will be presented in a future publication.
Materials and methods. —The fossil spec- imens are cataloged in the vertebrate zo- ology collections of the Bernice P. Bishop Museum (BPBM). Modern skeletal speci- mens are from the Division of Birds, Na- tional Museum of Natural History, Smith- sonian Institution (USNM). Fossils from New Caledonia are from the Institut de Pa- léontologie, Muséum National d’Histoire Naturelle, Paris (MNHN). Osteological ter- minology mainly follows that of Baumel et
538
>
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
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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
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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).
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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
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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
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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.
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548
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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
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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
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(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.
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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-
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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,
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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-
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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
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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
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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
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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).
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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.
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@ Frutillar Llanqu/hue
Lake
© Valdivia At0
<a 2 = 2 Lid ©) a <q
Bas
571
ah2
a
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sec.
SEC.
0.5
Fig. 3. Audiospectrograms (left) and sections (right) of the mating call of Eupsophus emiliopugini (a) and E.
vittatus (b), band filter 300 Hz.
lizo, Rio Lenca, Maullin, Puntra, Cucao, Yaldad, and Caleta Vidal) primarily at La Picada (type locality) and Puntra. Calls have been heard from September to December. Mature females with white oocytes were collected in October (La Picada and Cucao) and tadpoles have been found in December
(Puntra). Males with nuptial asperities were collected from September to December. The breeding sites are series of small water-filled Cavities at the border of the sireams near the forests.
Mating call. —The mating call of E. emi- liopugini was recorded at seven localities
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Table 2.—Characteristics of the mating call of Eup- sophus emiliopugini and E. vittatus. Observed ranges in parenthesis below means; E. emiliopugini recorded at Puntra (Chiloé Province) and E. vittatus recorded at Mehuin (Valdivia Province). (n = number of analyzed calls.)
Character E. emiliopugini E. vittatus n 10 10 Notes per 2 5 call (4-6) Notes 0.203 0.089 duration (0.132-0.250) (0.062-0.187) (sec) Pulses per 25.45 15.90 note (17-34) (11-23) Dominant 1132.08 1154.34 frequency (500-2000) (600-1680) (Hz)
(Termas de Puyehue, La Picada, El Trai- guen, Rio Rollizo, Maullin, Puntra, and Yaldad), and 37 frogs were recorded (5° 16°C). The description of the mating call is based on 10 calls from five animals from Puntra. Males of this species were observed and collected while they were calling from cavities in the ground, at the border of a stream. At this locality males were isolated; however, in the other sites a moderate ag- gregation was observed. The mating call of E. emiliopugini is characteristically com- posed of two notes (Table 2, Fig. 3). Both are similar in duration (x = 0.20 + 0.027 sec) and have 27 pulses (mean) per note. The fundamental frequency ranges between 85 and 633 Hz and the dominant frequency is spread between 729 and 1320 Hz. Anal- ysis of numerous field cecordings demon- strated only minor variations in call char- acteristics among individuals from different populations. In three localities (La Picada, Rio Rollizo, and Puntra) E. emiliopugini and E. calcaratus were collected calling in close physical association.
Eggs.—A female collected at the type lo- cality (La Picada), 24 Sep 1983, had 148 mature oocytes. At the same place (29 Oct 1984 and 21 Nov 1985) two clutches,
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Table 3.—Measurements (mm) (x + SD) of 30 tad- poles of Eupsophus emiliopugini at developmental stage 37 (Gosner 1960).
Total length 23.2) + O74 Body length 9.6 + 0.24 Body deep Or O21 Fin deep 6:5. ..0.36 Snout-nostril distance Le? 62002 Interocular distance 1.7
Eye diameter 1.0 + 0.009 Mouth width Pi Nea (J 2
containing 166 and 131 eggs (5.78-6.9 mm diameter) respectively, were found in water- filled cavities covered by mosses (Sphag- num). Eggs were stuck to one another and were creamy-white in color.
Tadpole. —131 larvae of E. emiliopugini were collected in a water-filled cavity near (20 cm) a cold stream (12°C) at Puntra (6 Dec 1984). The measurements of the tad- poles are indicated in Table 3. Gosner’s (1960) developmental stages are used in the following description. Larvae in stage 37 (Fig. 4a, b) with body ovoid in lateral view, two times longer than deep; contour of the snout flattish. Nostrils small, situated be- tween eye and snout tip. Eyes lateral, di- ameter 1.7 times the interocular distance. Mouth small and anteromedial, labial pa- pillae interrupted anteriorly, few denticles and tooth formula 2/2; beak well developed. Anal tube medial; no spiracle but small ven- trolateral fissure on the left side of the body. Caudal musculature robust, dorsal and ven- tral fins well developed; end of tail rounded. Color in life creamy-white abdomen and fins transparent; internal organs visible. Mi- nute melanophores among the nearest mus- cular septa to the body. Dorsal areas of head and trunk scarcely pigmented. On comple- tion of metamorphosis (10 Mar 1985) frogs were 10.1—10.6 mm in snout—vent length. Froglets have the dorsum gray with two well developed paravertebral folds also found in the adults.
Juvenile. —Juveniles (five fixed speci- mens from La Picada) exhibit the same
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2.5mm
0.5 mm
Fig. 4. Tadpole of Eupsophus emiliopugini, lateral view (a) and mouth (b). Gosner stage 37.
characteristics of the adults; however, some differences were observed. The two para- vertebral folds are well developed and the dorsum and flanks are covered by minute granules. The gular area is dark gray and the olive-green interocular band is not evident.
Variation. — Measurements of 54 adults (40 males and 14 females) (Table 1) show sexual size dimorphism. Of the 99 speci- mens examined (54 adults and 45 juve- niles), 59 specimens (60.2%) have a mid- dorsal line, 38 specimens lack a mid-dorsal line (37.7%) and two specimens (2.1%) from Puntra (IZUA 2060, 2062) exhibit scattered
whitish irregular spots on a dark back- ground.
Etymology. —The trivial name honors the memory of my friend and colleague Prof. Dr. Emilio Pugin, for his contribution to knowledge of the reproductive biology and development of the Chilean frogs.
Comparisons
When the adult snout—vent length of E. emiliopugini (males x = 46.7 mm; females X = 50.6 mm) is compared with that of other Eupsophus species, it is observed that this
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frog is as large as E. vittatus (males X = 54.4 mm; females X = 59.6 mm) (Table 1). Both taxa differ from their smaller congeners E. roseus (36.6 mm in snout-vent length), £. migueli (35.5 mm), E. calcaratus (35.1 mm), and E. insularis (39.9 mm) (Formas and Vera 1982).
In habitus and dorsal pattern, E. emilio- puginiis similar to Eupsophus vittatus; how- ever, both species differ in external features. Eupsophus emiliopugini has an olive-green band between the eyes, which is absent in E. vittatus. Furthermore, the latter has the vertebral line wide and creamy-white in col- or while in E. emiliopugini it is narrow and lemon-yellow colored.
Remarkable differences are found be- tween the mating calls of E. emiliopugini and E. roseus, E. migueli, and E. calcaratus. The mating call of E. emiliopugini has two notes while the other species presents only one note (Formas 1985). Differences be- tween the voices of E. emiliopugini and E. vittatus are presented in Table 2 and Fig. 3.
From the geographical point of view E. emiliopugini and E. vittatus demonstrate a different range of distribution (Fig. 2). Both taxa are allopatric, and EF. vittatus is dis- tributed in the Coastal Range and the Cen- tral Valley between Ramadillas (37°18'S; 73°14’'W) and Bahia Mansa (40°33'S; 73°46'W).
Specimens Examined
Abbreviations. —Instituto de Zoologia, Universidad Austral de Chile (IZUA); Field Museum of Natural History (FMNH); Mu- seo de Zoologia, Universidad de Concep- cion (MZUC); Juan Carlos Ortiz (personal collection) (J.C.O.); Nelson Diaz (personal collection) (N.D.).
Eupsophus emiliopugini. (99) Osorno: Puyehue (40°42'S; 72°18’W), IZUA 1931, 1933-34, 2012 (tape); Piedras Negras (40°53’S; 72°27'W), IZUA 1951-53; La Pi- cada (41°04'S; 72°26’'W), FMNH 218584- 85, IZUA 1585-88, 1950, 1953, 1608,
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1955-75, 2101 (tape). Llanquihue: Rio Len- ca (41°37'S; 72°40’W), IZUA 1939, 1948- 50, 1939-45; El Traiguén (41°11’S; 73°25'W), IZUA 2103 (tape); Rio Rollizo (41°27'S; 72°20'W), IZUA 2104 (tape); Road to Maullin (41°41’S; 73°21'W), IZUA 2102 (tape). Chiloé: Ancud (41°52’S; 73°50’W), J.C.O. 61/1-61/3, 68/13, 33/3; Lechagua (41°53’S; 73°51'W), FMNH_ 154829-30; Chepu (42°03’S; 74°02'W), MZUC 11939; Puntra (42°07’S; 73°49’W), IZUA 2059-64, 2104 (tape); Tepuhueico (42°47'S; 73°58'W), (2 adults and 6 juveniles untagged); Cucao (42°37'S; 74°07'W), IZUA 1627-28; Quel- lon (43°07'S; 73°37'W), FMNH 3715; Yal- dad (43°07'S; 73°43'W) 2078-95, 2105 (tape). Aisén: Puerto Cisnes (43°30’'S; 71°19'W), FMNH 132050-52, 132317, 132659; Caleta Vidal (45°16’S; 73°27'W), (1 adult untagged).
Eupsophus vittatus. (103) Arauco: Ra- madillas (37°18'S; 73°16’W), MZUC 11482; Contulmo (38°S; 73°13’W), IZUA 272/80. Valdivia: Mehuin (39°26’S; 73°10'W), IZUA 832-36, 1644-46, N.D. 1-6, IZUA 2096; San Martin (39°33’S; 72°59’W), IZUA 1716- 20, 1724, 2097 (tape), 2028-52; Linguento (39°33'S; 72°59'W), IZUA 1937-38, 2098 (tape); Mafil (39°39'S; 72°57'W), FMNH 3825-27; Ciudad de Valdivia (39°48’S; 73°14'W), IZUA 225-28; Huellelhue (39°44'S; 73°06’W), IZUA 1929-30, 2099 (tape); Los Molinos (39°46’S; 73°18'W), IZUA 1936; Tres Chiflones (40°03’S; 73°10’W), IZUA 2009-27; Cordillera Pe- lada (40°03'S; 73°10’W), IZUA 1935. Osor- no: Pucatrihue (40°26’'S; 73°47’'W), MZUC 12402, IZUA 1988-2008, 2100 (tape); Ba- hia Mansa (40°33’S; 73°46’W), J.C.O. 69/ 74.
Acknowledgments
The author would like to give special thanks to Carlos Varela; Lila Brieva, Gon- zalo Aguilar and Jorge Oporto for their field assistance. My gratitude also goes to Hymen Marx (FMNH) who kindly provided spec-
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imens under his control. Corina Zuniga typed the manuscript. This study was sup- ported by Fondo de Investigacion Cientifica y Tecnoldégica (Proyecto Fondecyt N° 1225) and Direccion de Investigacion Universi- dad Austral de Chile (Proyecto RS 85-25).
Literature Cited
di Castri, F. 1968. Esquisse écologique du Chile. Bio- logie de l’ Amerique australe. Cl. Deboutteville and E. Rapaport (eds). — Editions du Centre Na- tional de la Recherche Scientifique. Paris [V:7— 52.
Formas, J. R. 1985. The voices and relationships of the Chilean frogs Eupsophus migueli and E. cal- caratus (Amphibia: Anura: Leptodactylidae).— Proceedings of the Biological Society of Wash- ington 98:411-415.
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
1989. The tadpole of Eupsophus calcaratus in Southern Chile.—Journal of Herpetology (in press).
, & E. Pugin. 1978a. The tadpoles of Hylorina sylvatica, Eupsophus vittatus, and Bufo varie- gatus in Southern Chile.— Herpetologica 34:355-— 358.
,& . 1978b. Tadpoles of Eupsophus ro- seus and Bufo variegatus in Southern Chile.— Journal of Herpetology 12:243-246.
,& M.I. Vera. 1982. The status of two Chilean frogs of the genus Eupsophus (Anura: Lepto- dactylidae).— Proceedings of the Biological So- ciety of Washington 95:594-601.
Gosner, K. L. 1960. A simplified table for staging anuran embryos and larvae with notes on iden- tification. — Herpetologica 16:183-190.
Instituto de Zoologia, Universidad Aus- tral de Chile, Casilla 567, Valdivia, Chile.
PROC. BIOL. SOC. WASH. 102(3), 1989, pp. 577-585
A REDESCRIPTION OF PSEUDORHOMBUS MEGALOPS, WITH COMMENTS ON CEPHALOPSETTA VENTROCELLATA (OSTEICHTHYES: PLEURONECTIFORMES: PARALICHTHYIDAE)
Dannie A. Hensley and Kunio Amaoka
Abstract. —Pseudorhombus megalops has not been recorded in the literature since first described by Fowler (1934) from the Philippine Islands. The species is redescribed from the type specimens and additional material from the Phil- ippine Islands, eastern Indian Ocean, Bali Strait, and Arafura Sea. Pseudo- rhombus megalops has a very distinct black spot or ocellus on the left pelvic fin. The only other Indo-Pacific paralichthyid with a similar character is Cepha- lopsetta ventrocellata. These species are compared and characters are given for their separation. Cephalopsetta ventrocellata, previously known from the east and west coasts of India and Pakistan, is shown to range to the Andaman Sea and Gulf of Oman. Cephalopsetta has been considered a close relative of An- cylopsetta (western Atlantic and eastern Pacific) and Gastropsetta (western At- lantic) because they share an elongate left pelvic fin. Osteological characters of the caudal fin, however, support placement of Cephalopsetta in with the Indo-
Pacific genera Pseudorhombus and Tarphops.
Fowler (1934) described many new flat- fish species collected mainly from the Phil- ippine Islands and adjacent regions. Most authors have overlooked Fowler’s publi- cation, apparently because it appeared dur- ing the same year as, and thus was not cited in, Norman’s (1934) monograph on flatfish- es. Most of Fowler’s descriptions and figures of the new flatfishes were inadequate. In ad- dition, he based many of his interpretations upon the older classification of Weber & de Beaufort (1929). Thus, the status of most of Fowler’s (1934) genera and species was un- certain. One species described in this work was Pseudorhombus megalops. The descrip- tion and figure of this species were poor, and there have been no other published rec- ords of P. megalops. Additional specimens _ were recently collected from the eastern In- dian Ocean, Bali Strait, and Arafura Sea during the Joint Eastern Tropical Indian Ocean Fishery Survey (JETINDOFISH; see
Gloerfelt-Tarp & Kailola 1984). We origi- nally could not identify the JETINDOFISH specimens to species and left them as ““Pseu- dorhombus sp. 1”’ in Gloerfelt-Tarp & Kai- lola (1984), stating that the species appeared very close to P. megalops. Since that time we have found additional specimens from the Philippine Islands and have had the op- portunity to make the necessary compari- sons with Fowler’s types for a positive iden- tification as P. megalops.
Dutt & Rao (1965) described Cepha- lopsetta ventrocellata from the east coast of India. The only other published records of the species are those of Kotthaus (1977) from the west coast of India and Pakistan. We have examined additional material from the Andaman Sea, west coast of India, and Gulf of Oman. Cephalopsetta ventrocellata shows some similarity to P. megalops and can be confused with that species.
In this paper we redescribe P. megalops
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Fig. 1.
from the type specimens and additional ma- terial and discuss some of the characters and possible phylogenetic position of C. ventro- cellata and compare it with P. megalops.
Materials and Methods
Methods of counts and measurements follow those of Hubbs & Lagler (1949) with two changes. Because all dorsal- and anal- fin rays are unbranched, all ray elements are counted as individual rays. Length of the pelvic fin is the length of the longest ray of that fin. Measurements were made with dial calipers to the nearest 0.1 mm. For regres- sion analysis all variates were transformed to natural logarithms. Standard length was treated as the independent variable. Tests for allometry were performed with the geo- metric-mean-functional-regression model of Ricker (1973). In this model 95% con- fidence limits are determined for the slope (v). If unity is outside of these limits allom- etry is assumed (positive if below, negative if above); isometry is assumed if unity is within the limits.
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Pseudorhombus megalops, holotype, USNM 93082, 152.0 mm SL.
Caudal-fin drawings were made from ra- diographs or specimens cleared and stained according to the method of Taylor (1967).
The institutional abbreviations are as fol- lows: ANSP—Academy of Natural Sci- ences, Philadelphia; CSIRO—Common- wealth Science and Industrial Research Organization, Hobart, Tasmania; FONR— Florida Department of Natural Resources, St. Petersburg, Florida; HUMZ— Hokkaido University, Laboratory of Marine Zoology, Faculty of Fisheries, Hakodate; NTIM— Northern Territory Museum of Arts and Sciences, Darwin, Australia; UPRM— Uni- versity of Puerto Rico-Mayaguez; and USNM-— National Museum of Natural His- tory, Smithsonian Institution, Washington, D.C. Standard length and total length are abbreviated SL and TL.
Pseudorhombus megalops Fowler Figs. 1, 2A—B, 3A—B, 5B, Tables 1-2
Pseudorhombus megalops Fowler, 1934:329, fig. 83 (Philippine Islands).
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imm EEE
imm
379
' imm
Fig. 2. Scales from near mid-length of body immediately above lateral line: A, left side of Pseudorhombus megalops, HUMZ 111769, 146.5 mm SL; B, right side of P. megalops, HUMZ 111769, 146.5 mm SL; C, left side of Cephalopsetta ventrocellata, ANSP 153379, 161.7 mm SL.
Pseudorhombus sp. 1.—Gloerfelt-Tarp & Kailola, 1984:272 (unnumbered plate), 273, 357 (eastern Indian Ocean, Bali Strait, and Arafura Sea).
Diagnosis. —Dorsal-fin rays 67—70; anal- fin rays 50-53; lateral-line scales 70-77. Gill rakers elongate, 15—18 on lower limb. Low- er jaw with 12-20 teeth on blind side, 2—4 large canines near symphysis. Scales on ocu- lar side ctenoid, most scales on blind side ctenoid (Fig. 2A—B). Tip of first interhemal spine stout, usually projecting through body wall anterior to first anal-fin ray. Pelvic fin of ocular side with distinct black spot over fourth or fifth ray (Fig. 3A—B); pelvic fin of blind side longer than that of ocular side.
Description. — Morphometrics as % SL are presented in Table 1. Dorsal-fin rays 67—70; anal-fin rays 50-53; pectoral-fin rays ocular side 11-13, blind side 1 1—12; pelvic-fin rays ocular side 6, blind side 6; lateral line scales 70-77; gill rakers ocular side 5—9 + 15-18; teeth on blind side of lower jaw 12-20.
Head length 3.0—3.3, body depth 1.9-2.4, both in SL. Measurements in head length are as follows: Snout length 3.9—4.5; upper- jaw length ocular side 2.0—2.2; lower-jaw length ocular side 1.6—1.8; lower-eye length 3.4—4.8; pectoral-fin length ocular side 1.5— 1.9, blind side 2.1—2.9; pelvic-fin length ocular side 3.04.2, blind side 2.7—3.8; length of first dorsal-fin ray 2.6-—3.5, second 3.1-
4.0, third 3.6-4.7, fourth 3.6—5.4; length of caudal peduncle 3.6-5.2, depth of caudal peduncle 2.8-—3.4.
Anterior profile of head with indentation anterior to upper eye. Anterior margins of eyes at same transverse levels. Posterior end of maxilla below middle to posterior one- third of lower eye. Nostrils of ocular side at same horizontal level as upper margin of lower eye. First dorsal-fin ray variable in position, above either nostril or interspace between nostrils. Tip of isthmus below pos- terior one-quarter or posterior margin of lower eye. Teeth of upper jaw similar on ocular and blind sides, small and closely spaced laterally with from four to six widely spaced, large canines anteriorly; teeth of lower jaw similar on ocular and blind sides, large and widely spaced laterally with from two to four (usually two) very large canines anteriorly. Lower jaw with prominent sym- physial knob. Gill rakers elongate, pointed, with small teeth.
Scales on ocular side ctenoid; most scales on blind side ctenoid, cycloid scales prob- ably being replacement scales (Fig. 2A—B). Supratemporal branch of lateral line reach- ing one-half to three-quarters of distance to dorsal fin base.
First interhemal spine stout, usually pro- jecting through body wall immediately an- terior to first anal-fin ray.
First few dorsal-fin rays slightly elongate.
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Fig. 3:
Left pelvic fins: A, Pseudorhombus megalops, CSIRO CA2526, 129.8 mm SL; B, P. megalops,
holotype, USNM 93082, 152.0 mm SL; C, Cephalopsetta ventrocellata, ANSP 153383, 66.0 mm SL; D, C. ventrocellata, ANSP 153382, 102.4 mm SL; E, C. ventrocellata, ANSP 153379, 161.7 mm SL.
Pelvic fin of blind side slightly longer than that of ocular side.
Color in alcohol. —Ocular side tan to dark grey. Some specimens with dark rings ar- ranged in two longitudinal series above and below lateral line, scattered, smaller dark rings and blotches, and dark streaks on dor- sal and anal fins (see unnumbered plate of “*Pseudorhombus sp. 1” in Gloerfelt-Tarp & Kailola 1984:272). All preserved specimens with distinct black spot on fourth or fifth
ray of pelvic fin of ocular side (Fig. 3A—B); some indication that black spot may have white margin in life. Blind side tan or whit- ish.
Discussion
Norman (1934) recognized 21 species of Pseudorhombus Bleeker as valid and three as being of doubtful validity. Subsequently, Fowler (1934) described P. megalops and
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Table 1.— Morphometric proportions expressed as percentage of SL for specimens of Pseudorhombus megalops.
Holoty
pe (USNM 93082)
Standard length (mm) 152.0 Head length 31.3 Body depth 48.9 Upper-jaw length (ocular side) 15.3 Lower-jaw length (ocular side) 18.9 Eye length (lower) 8.6 Snout length Ye Depth of caudal peduncle 9.5 Length of caudal peduncle 6.8 Pectoral-fin length (ocular side) 19.3 Pectoral-fin length (blind side) 14.6 Pelvic-fin length (ocular side) 8.8 Pelvic-fin length (blind side) 11.4 First dorsal-fin ray length 9.9 Second dorsal-fin ray length 8.5 Third dorsal-fin ray length 7.8 Fourth dorsal-fin ray length 7.0
Amaoka (1969) P. oculocirris. Characters given in the “Diagnosis” will distinguish P. megalops from all known species of Pseu- dorhombus.
Dutt & Rao (1965) described a new genus and species of paralichthyid, Cephalopsetta ventrocellata, from the east coast of India (Visakhapatnam). This species resembles P. megalops in having a very distinct black spot enclosed by a light-colored ring be- tween the third and fifth ray of the pelvic fin of the ocular side (Figs. 3, 4). These species are the only known Indo-Pacific par- alichthyids with a distinctive dark spot or ocellus on this fin. The South African species Pseudorhombus natalensis Gilchrist has a small dark spot on this fin but the spot is relatively diffuse and frequently absent, at least in preserved specimens. Other paral- ichthyids with an ocellus or distinctive dark spot on the pelvic fin of the ocular side are western Atlantic (Ancylopsetta kumperae Tyler, Paralichthys oblongus [Mitchill], Paralichthys isosceles Jordan) or eastern Pa- cific species (Lioglossina tetrophthalmus Gilbert). C. ventrocellata and P. megalops can be distinguished by characters pre- sented in Table 2.
Philippine Islands, including paratypes Eastern Indian Ocean, Bali (USNM 93550) Strait, and Arafura Sea (n = 7) (n = 5)
131.7-174.7 116.8-154.2 31.9-33.4 30.2-33.1 47.4-51.9 40.9-48.4 15.0-16.1 13.9-16.6 18.4-19.5 17.6—20.1 7.9-8.9 6.2-9.2 7.4-8.4 6.7-7.7 9.5-10.1 9.6-11.0 6.4-7.2 5.8-8.7 17.9-22.0 15.8-20.8 13.0-15.8 10.4-14.5 8.0—-9.3 7.5-10.9 8.6—10.5 8.1-11.5 10.0-11.6 8.7-12.7 7.9-9.1 7.6-10.8 7.0-8.7 6.5-9.2 6.1-8.1 7.0-9.2
Kotthaus (1977) examined specimens of C. ventrocellata from the west coast of India and Pakistan and described some additional characters not mentioned by Dutt & Rao (1965). We have examined specimens from the Andaman Sea, eastern Arabian Sea, and Gulf of Oman. Our specimens agree with the descriptions of Dutt & Rao (1965) and Kotthaus (1977) with some exceptions. Dutt & Rao (1965) state that the ocular side has “‘a few irregular dark blotches.”’ Most of our specimens show distinct dark spots ar- ranged in about five longitudinal rows. The most distinctive dark spots are those im- mediately below the bases of the dorsal and anal fins and usually a series of three spots along the lateral line. In some specimens there is a faint pattern of several broad, dark transverse bars. Kotthaus (1977) describes the dorsal-fin origin as being immediately above the posterior nostril on the blind side. This character is variable in our specimens, the base of the first dorsal-fin ray being above either nostril or the space between them. According to Dutt & Rao (1965), C. ven- trocellata has scales with very weak ctenti on the ocular side of the body and cycloid scales on the head and blind side. Kotthaus
582 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig. 4. Cephalopsetta ventrocellata: A, adult, ANSP 153379, 168.4 mm SL; B, juvenile, ANSP 153383, 66.0 mm SL.
(1977) expanded on this by saying the scales appear to be cycloid except for some very are covered by skin. Scales on the ocular small ctenii proximal to the exposed edges side of our specimens are covered by skin (Fig. 2C). The specimens we examined have with only their posterior edges visible. They the elongate left pelvic fin described by Dutt
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583
Table 2.—Characters useful for distinguishing Pseudorhombus megalops and Cephalopsetta ventrocellatus.
P. megalops C. ventrocellatus Standard length/head length 3.0-3.3 2.3-3.1 (usually 2.3-2.9) Head length/length of pelvic fin of ocular side 3.0-4.2 1.4-2.3 Head length/snout length 3.9-4.5 4.7-5.9 Length of pelvic fin of ocular side/length of 0.8-0.9 1.2-1.8 pelvic fin of blind side Number of teeth on lower jaw on blind side 12-20 23-31
Morphology of lower-jaw teeth
Scales on ocular side Scales on blind side
& Rao (1965) and Kotthaus (1977). In ad- dition, our specimens showed negative al- lometric growth of this fin (v = 0.5492 + 0.1437). Thus, our smallest specimen (66.0 mm SL) had a left-pelvic-fin length of 28.9% SL, specimens of 100.7—168.4 mm SL 16.8—- 22.2% SL, and a 207.5-mm-SL specimen 15.6% SL (Figs. 3C-E, 4).
The major characters used by Dutt & Rao (1965) to define Cephalopsetta are a large head (2.3-3.1 in SL) and an elongate left pelvic fin. Ancylopsetta Gill (western Atlan- tic and eastern Pacific) and Gastropsetta Bean (western Atlantic) also have the pelvic fin of the ocular side longer than that of the blind side, and were thus treated by Nor- man (1934) as being closely related and dis- tinct from other paralichthyid genera. For the same reason Dutt and Rao considered Cephalopsetta closely related to Ancylop- setta and Gastropsetta and restricted their comparative statements to these genera.
Current knowledge of relationships with- in the Paralichthyidae was recently re- viewed by Ahlstrom et al. (1984) and Hens- ley & Ahlstrom (1984). These authors regard Cephalopsetta as a member of the Pseudor- hombus group (along with the Indo-Pacific genera Pseudorhombus and Tarphops Jor- dan & Thompson), a group they considered as probably monophyletic. Ancylopsetta and Gastropsetta were left in a group (referred to as the Paralichthys group) composed of Paralichthys Girard, Hippoglossina Stein- dachner, Lioglossina Gilbert, Verecundum
Widely spaced, large canines anteriorly Well-developed ctenii
Most ctenoid
Closely spaced, no large canines
Feeble ctenii
Cycloid
Jordan, and Xystreurys Jordan & Gilbert; the authors could find no current evidence for monophyly of this group.
Much of the evidence Ahlstrom et al. (1984) and Hensley & Ahlstrom (1984) used for placing Cephalopsetta in the Pseudo- rhombus group and excluding Ancylopsetta and Gastropsetta involved caudal-fin struc- ture. Species of the Paralichthys group have 18 caudal-fin rays, at least one free epural (except in one species of Hippoglossina [Sumida et al. 1979]), and a splinter ray on the base of the ventralmost caudal-fin ray (Fig. SC). The splinter ray is probably a rem- nant of a ray lost through fusion with an adjacent ray (Okiyama 1974). Amaoka (1969) and Hensley & Ahlstrom (1984) con- sidered these characters as probably being plesiomorphic. The Pseudorhombus group usually has 17 caudal-fin rays, the epural fused to the fifth hypural and no splinter ray (Fig. 5A—-B). These authors regarded these characters as probably derived and indica- tive of monophyly.
Although we tentatively treat Cephalop- setta as a member of the Pseudorhombus group, it should be noted that Gutherz (1966) found that juveniles of Ancylopsetta antil- larum Gutherz and Gastropsetta frontalis Bean have greater relative lengths of left pelvic fins than adults, a growth pattern similar to that seen in C. ventrocellata. However, other paralichthyids are known to have elongate left pelvic fins at some stage of development. Several species of what
584
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig. 5.
Caudal skeletons: A, Cephalopsetta ventrocellata, ANSP 153379, 161.7 mm SL; B, Pseudorhombus
megalops, HUMZ 111769, 146.5 mm SL; C, Ancylopsetta quadrocellata, FDNR 6115, 52.9 mm SL; abbrevi- ations: EP = epural; HY 1-5 = hypurals 1-5; PHY = parhypural; PU2 = preural centrum 2; SR = splinter ray;
THC = terminal half-centrum.
Ahlstrom et al. (1984) and Hensley & Ahl- strom (1984) called the Cyclopsetta group have elongate pelvic fin rays on the ocular side as larvae, while adults have short pelvic fins of approximately equal length (see Ahl- strom etal. 1984). Nielsen (1963) has shown that post-metamorphic individuals of at least one species of Cyclopsetta (named Dor- sopsetta norma in Nielsen 1963) have elon- gate rays in the left pelvic fin. A more de- tailed comparative analysis of pelvic-fin growth patterns is needed before they can be used for phylogenetic inference. Material examined. —Pseudorhombus megalops: Philippine Islands: USNM 93082, holotype, 152.0 mm SL; USNM 93550, paratypes, 2 specimens, 141.1-147.8; USNM 93551, 2, 131.8-136.4; USNM 93548, 2, 144.3-174.7; USNM 93549, 131.7. Indian Ocean (south coasts of Su- matra, Java, and Lombok): HUMZ 111768, 154.0; HUMZ 111769, 146.5; NIM 10760- 006, 154.2. Bali Strait: NTM S.11022-002, 116.8. Arafura Sea: CSIRO CA2526, 129.8. Cephalopsetta ventrocellata: Gulf of Oman: ANSP 153383, 66.0. India (west coast): ANSP 153379, 6, 100.7-168.4; ANSP 153380, 207.5; ANSP 153382, 102.4; Andaman Sea: ANSP 153381, 138.0.
Acknowledgments
The senior author thanks V. G. Springer and S. L. Jewett (USNM), and W. F. Smith-
Vaniz, B. Chernoff, and W. G. Saul (ANSP) for space and assistance during visits to their museums. We thank P. Kailola and T. Gloerfelt-Tarp for providing us with spec- imens of P. megalops from the JETIND- OFISH Survey, L. P. Norrod (USNM) for helping us to solve a problem with Fowler’s type specimens, W. G. Saul for radiographs of C. ventrocellata, and D. L. Ballantine (UPRM) for help with photography. This work was partially supported by the L. P. Schultz Fund (USNM), Smithsonian Insti- tution Short-Term Visitors Program, Office of Research Coordination (UPRM), and the National Science Foundation U.S.—Japan Cooperative Science Program (NSF INT-86 35526):
Literature Cited
Ahlstrom, E. H., K. Amaoka, D. A. Hensley, H. G. Moser, & B. Sumida. 1984. Pleuronecti- formes: Development. Pp. 640-670 in H. G. Moser, W. J. Richards, D. M. Cohen, M. P. Fahay, A. W. Kendall, Jr., & S. L. Richardson, eds., Ontogeny and systematics of fishes. Amer- ican Society of Ichthyologists and Herpetolo- gists, Special Publication 1.
Amaoka, K. 1969. Studies on the sinistral flounders found in the waters around Japan. Taxonomy, anatomy, and phylogeny.—Journal of the Shi- monoseki University of Fisheries 18(2):65—340.
Dutt, S., & K. H. Rao. 1965. A new bothid flatfish Cephalopsetta ventrocellatus gen. et sp. nov. from
VOLUME 102, NUMBER 3
Bay of Bengal.—Proceedings of the Indian Academy of Sciences 62(4):180-187.
Fowler, H. W. 1934. Descriptions of new fishes ob- tained 1907 to 1910, chiefly in the Philippine Islands and adjacent seas.— Proceedings of the Academy of Natural Sciences of Philadelphia 85:233-367.
Gloerfelt-Tarp, T., & P. J. Kailola. 1984. Trawled fishes of southern Indonesia and northwestern Australia. Australian Development Assistance Bureau, Directorate General of Fisheries, In- donesia, and German Agency for Technical Co- operation, xx + 406 pp.
Gutherz, E. J. 1966. Revision of the flounder genus Ancylopsetta (Heterosomata: Bothidae) with de- scriptions of two new species from the Antilles and the Caribbean Sea. — Bulletin of Marine Sci- ence 16(3):445-479.
Hensley, D. A., & E. H. Ahlstrom. 1984. Pleuronec- tiformes: Relationships. Pp. 670-687 in H. G. Moser, W. J. Richards, D. M. Cohen, M. P. Fahay, A. W. Kendall, Jr., & S. L. Richardson, eds., Ontogeny and systematics of fishes. Amer- ican Society of Ichthyologists and Herpetolo- gists, Special Publication 1.
Hubbs, C. L., & K. F. Lagler. 1949. Fishes of the Great Lakes region.—Cranbrook Institute of Science Bulletin 26:1-186.
Kotthaus, A. 1977. Fische des Indischen Ozeans. Er- gebnisse der ichthyologischen Untersuchungen wahrend der Expedition des Forschungsschiffes “Meteor” in den Indischen Ozean, Oktober 1964 bis Mai 1965. A. Systematischer Teil, XX. Pleu- ronectiformes (Heterosomata).—‘‘Meteor’’ Forschungs-Ergebnisse Reihe D 26:1-—20.
Nielsen, J. G. 1963. Notes on some Heterosomata (Pisces) from N-W South America with the de- scription of a new genus and species and a new subspecies of Paralichthinae.— Videnskabelige Meddelelser fra dansk naturhistorisk Forening 125:377-400.
585
Norman, J. R. 1934. A systematic monograph of the flatfishes (Heterosomata). Vol. I. Psettodidae, Bothidae, Pleuronectidae. British Museum, London, vii + 459 pp.
Okiyama, M. 1974. Studies on the early life history ofa flounder, Paralichthys olivaceus (Temminck et Schlegel). II. Descriptions of juveniles and the comparison with those of related species. — Bul- letin of the Japan Sea Regional Fisheries Re- search Laboratory 25:39-61.
Ricker, W. E. 1973. Linear regression in fishery re- search. — Journal of the Fisheries Research Board of Canada 30(3):409-434.
Sumida, B., E. H. Ahlstrom, & H. G. Moser. 1979. Early development of seven flatfishes of the east- ern North Pacific with heavily pigmented larvae (Pisces, Pleuronectiformes).— Fishery Bulletin 77(1):105-145.
Taylor, W. R. 1967. An enzyme method of clearing and staining small vertebrates. — Proceedings of the United States National Museum 122(3596): 1-17.
Weber, M., & L. F. de Beaufort. 1929. The fishes of the Indo-Australian Archipelago. V. Anacan- thini, Allotriognathi, Heterosomata, Beryco- morphi, Percomorphi: Families Kuhliidae, Apogonidae, Plesiopidae, Pseudoplesiopidae, Priacanthidae, Centropomidae. E. J. Brill, Ltd., Leiden, xiv + 458 pp.
(DAH) Department of Marine Sciences, University of Puerto Rico, P.O. Box 5000, Mayaguez, Puerto Rico 00709-5000, U.S.A.; (KA) Laboratory of Marine Zoology, Fac- ulty of Fisheries, Hokkaido University, Hakodate, Hokkaido 041, Japan.
PROC. BIOL. SOC. WASH. 102(3), 1989, pp. 586-589
PARASPADELLA ANOPS, NEW SPECIES, FROM SAGITTARIUS CAVE GRAND BAHAMA ISLAND, THE SECOND TROGLOBITIC CHAETOGNATH
Thomas E. Bowman and Robert Bieri
Abstract. — Paraspadella anops is described from a single specimen collected in Sagittarius Cave, Grand Bahama Island. It lacks eyes and body pigmentation, and is the first fully troglobitic chaetognath discovered. In morphological details it closely resembles P. schizoptera (Conant), but is much slenderer than the latter. The separation of Paraspadella Salvini-Plawen, with two pairs of lateral fins, and Gephyrospadella Salvini-Plawen, with one pair, is rejected, since in the former the two pairs of fins are continuous in young specimens and only separated at sexual maturity by lateral growth of the vaginas. Gephyrospadella is reduced to a synonym of Paraspadella.
Until the discovery of the species de- scribed herein, the only chaetognath re- ported to be troglobitic was Spadella ledo- yeri Casanova, 1986, from a marine cave about 15 km east of Marseille, France. In contrast to Paraspadella anops, however, S. ledoyeri has well developed eyes having ““srand développement de la tache pigmen- taire,’”’ which Casanova considered an ad- aptation to the reduction or absence of light in the interior of the cave. Casanova said nothing about the presence or absence of body pigment in S. ledoyeri.
Spadella equidentata Casanova, 1987, from 452 m in the Gulf of Cadiz, south of Portugal, was described as having not a trace of eyes, even after staining with methylene blue. However, Casanova (1987) remarked that the absence of eyes required confir- mation.
The new species described herein is the first truly troglobitic chaetognath that lacks both eyes and body pigment.
Spadellidae Tokioka, 1965 Paraspadella Salvini-Plawen, 1986
Type species.—Spadella schizoptera Co- nant, 1895a (original designation).
Paraspadella anops, new species Figs. 1-3
Material. —Bahamas, Grand Bahama Is- land, Sweeting’s Cay, Sagittarius Cave (26°37'N, 77°53'W), leg. Dennis Williams and Jill Yager, 17 Dec 1987, holotype, USNM 120108.
Etymology. —From the Greek ‘“‘an-”’ (without) + “ops” (eye), alluding to the ab- sence of eyes.
Description. — Total length, excluding tail fin, 3.55 mm. Length of tail 1.75 mm = 49.3% of total length. Eyes and body pig- ment totally lacking. Head about 1.4 as wide as long, about twice as wide as trunk and 2.7X as wide as tail. Grasping spines 10 on each side, each with tip set off by suture. Anterior teeth two on each side, elongate. Posterior teeth lacking. Corona ciliata pyriform, length about 1.6 x greatest width, which is near posterior end. Collar- ette extending from posterior end of head to anterior insertion of tail fin. Lateral mar- gins of collarette of trunk with four pairs of sensory tufts, of tail with three pairs. An- terior intestinal diverticula not evident in the undissected specimen. One pair of lat- eral fins, completely rayed, extending from
VOLUME 102, NUMBER 3
distance equal to body width anterior to be- ginning of tail to short distance anterior to attachment of adhesive organs; fin width increasing posteriorly to maximum slightly posterior to fin midlength, thereafter de- creasing. Tail fin beginning just posterior to insertion of adhesive organs; lateral margins nearly straight; posterior margin, which ap- pears somewhat frayed, slightly concave.
Adhesive organs inserted on ventral sur- face of tail, each with four slender fingers. Outer two and inner two fingers each with common base. Outer two fingers subequal, about half length of tail fin; inner two fingers with inner member slightly longer, about two-thirds length of tail fin, reaching pos- terior margin of tail. Fingers longitudinally striated; tips of most fingers eroded, but one relatively undamaged tip covered with small papillae (Fig. 3).
Ventral ganglion oval, occupying middle one-fifth trunk. Ovaries immature, extend- ing slightly anterior to 4th pair of sensory tufts. Transverse musculature appears to extend from posterior end of head to pos- terior margin of ventral ganglion.
Comparisons. —Of the known species of Paraspadella, P. schizoptera (Conant, 1895) most closely resembles P. anops. The two species agree in the numbers of grasping spines and anterior teeth; both have pyri- form coronae ciliatae, and the number, form, and arrangement of the fingers of the ad- hesive organs are identical. However, P. schizoptera has well developed eyes and is heavily pigmented compared with most other chaetognaths (Feigenbaum 1976). It is also a stockier species than the very slen- der P. anops, whose length (excluding the tail fin) is about 9.5 x its greatest trunk width. The length : width ratio of P. schizoptera in published illustrations is as follows: 6.5
—
Figs. 1-3. Paraspadella anops: 1, Habitus, ventral (not all 10 pairs of grasping spines shown; many fin rays omitted); 2, Head, dorsal; 3, Tip of finger of ad- hesive organ.
L
—
SPEAR Te APSR LES a es BEREAN et Se Oy that ——- AT et
ey GEN G) Eon im Tecate are
ay
Viiipe, yy
TER
Ezz ey j
‘Re
CFM ZA
587
588
(Conant 1895a, fig. 6, 3.5 mm specimen), 7.8 (Owre 1963, fig. la, 1.7 mm specimen), 7.9 (Owre 1972, pl. 2, fig. 1, 3.8 mm spec- imen), 5.3 (Feigenbaum 1976, fig. 2f, 1.4 mm specimen).
Because of these differences, it 1s our sub- jective opinion that P. anops merits recog- nition as specifically distinct from P. schi- zoptera, but the latter is clearly the ancestor of our new species.
The genera of Spadellidae. — Prior to Sal- vini-Plawen’s two new genera, the family Spadellidae Tokioka, contained only the ge- nus Spadella, although three species-groups corresponding to the three genera recog- nized by Salvini-Plawen had been pro- posed. Salvini-Plawen (1986) assigned the 13 then known species of Spadella to 3 gen- era: Spadella Langerhans, 1880 (type species Sagitta cephaloptera Busch, 1851), one pair of lateral fins, no adhesive organs; Gephy- rospadella, new genus (type species Spadella pulchella Owre, 1963), one pair of lateral fins, adhesive organs present; Paraspadella, new genus (type species Spadella schizop- tera Conant, 1895a), two pairs of lateral fins, adhesive organs present.
We agree that the presence or absence of adhesive organs is significant at the generic level, but we have reservations about the supposed two pairs of lateral fins. Illustra- tions of Paraspadella schizoptera show specimens with two pairs (Conant 1895a, Owre 1972) and one pair (Owre 1963, Fei- genbaum 1976) of lateral fins. Where there are two pairs, the gap between them is much smaller than in species of Sagitta, in which the anterior and posterior fins are widely separated. In P. schizoptera the fins are either continuous or separated only by the width of the female gonopore and vagina. It ap- pears that the lateral fins are single in sex- ually immature individuals, but with the development of sexual maturity they be- come separated into anterior and posterior parts by lateral growth of the vagina. This slight separation scarcely warrants the rec- ognition of two pairs of fins.
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
This observation leads us to combine Sal- vini-Plawen’s proposed new genera, Gephy- rospadella and Paraspadella, since they dif- fer only in the supposed number of paired lateral fins. Acting as first revisors (ICZN Article 24), we may select one of the names. Salvini-Plawen did not give the derivation of his new generic names, but it is obvious the ““Gephyro” means “‘bridge” and “‘Para”’ means “near.”’ Salvini-Plawen’s fig. 3 shows Geophyrospadella bridging the evolutionary pathway between Spadella and Paraspadel- la. Combining the new genera eliminates the bridge, hence we select Paraspadella rather than Gephyrospadella. This act re- duces the number of genera of Spadellidae to two: Spadella, without adhesive organs, and Paraspadella, with adhesive organs.
Habitat.—A detailed description of Sag- ittarius Cave is given by Cunliffe (1985). It is an anchialine cave beneath the small is- land of Sweeting’s Cay at the east end of Grand Bahama Island. The single specimen of P. anops was found swimming in the water column in an area completely devoid of light, more than 100 m inward from the surface entrance to the cave. The salinity was 35%o. Associated fauna include remipedians, os- tracodes, amphipods, isopods, mysida- ceans, thermosbaenaceans, and cave fishes.
Acknowledgments
Our sincere thanks go to the collector, Jill Yager, who sent us the specimen, together with collection data.
Literature Cited
Busch, W. 1851. Beobachtungen liber Anatomie und Entwicklung einiger wirbellosen Seethiere. Ber- lin, vill + 143 pp., 17 plates [Chaetognatha, pp. 93-100].
Casanova, Jean-Paul. 1986a. Spadella ledoyeri, chae- tognathe nouveau de la grotte sous-marine ob- scure des Trémies (calanques de Cassis).—Rap- ports et Procés Verbaux des Réunions Commission Internationale pour |’Exploration Scientifique de la Mer Méditerranée 30(2):196.
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1986b. Découverte en Méditerranée d’un
chaetognathe nouveau du genre archaique pro-
fond Archeterokrohnia: Description et signifi- cation biogéographique.—Rapports et Procés
Verbaux des Réunions Commission Interna-
tionale pour l’Exploration Scientifique de la Mer
Méditerranée 30(2):196.
1987. Deux chaetognathes benthiques nou- veaux du genre Spadella des parages de Gi- braltar: Remarques phylogénétiques. — Bulletin du Muséum National d’Histoire Naturelle, Paris, series 4, 9 (section A, no. 2):375-390.
Conant, F. S. 1895a. Description of two new chae- tognaths: Spadella schizoptera and Sagitta his- pida. —The Johns Hopkins University Circulars
14(119):77-78.
. 1895b. Description of two new chaetognaths
(Spadella schizoptera and Sagitta hispida).—
Annals and Magazine of Natural History, series
6, 16:288-292. (Verbatim reprint of Conant
1895a).
Cunliffe, Sarah. 1985. The flora and fauna of Sagit- tarius, an anchialine cave and lake in Grand Bahama.—Cave Science 12(3):103-109.
Feigenbaum, David L. 1976. Development of the adhesive organ in Spadella schizoptera (Chae- tognatha) with comments on growth and pig-
589
mentation.— Bulletin of Marine Science 26(4): 600-603.
Langerhans, P. 1880. Die Wurmfauna von Madeira III.— Zeitschrift fiir Wissenschaftliche Zoologie 34:132-136.
Owre, Harding B. 1963. The genus Spadella (Chae-
tognatha) in the western North Atlantic Ocean,
with descriptions of two new species. — Bulletin of Marine Science of the Gulf and Caribbean 13(3):378-390.
. 1972. Marine biological investigations in the
Bahamas 18. The genus Spadella and other
Chaetognatha.—Sarsia 49:49-58.
Salvini-Plawen, L.v. 1986. Systematic notes on Spa- della and on the Chaetognatha in general. — Zeit- schrift fiir Zoologische Systematik und Evolu- tionsforschung 24(2):122-128.
Tokioka, Takasi. 1965. The taxonomical outline of Chaetognatha.— Publications of the Seto Ma- rine Biological Laboratory 13(3):335—357.
(TEB) Department of Invertebrate Zool- ogy, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560; (RB) 175 Brookside Drive, Yellow Springs, Ohio 45387.
PROC. BIOL. SOC. WASH. 102(3), 1989, pp. 590-608
A REVIEW OF THE BEACH FLIES OF THE CARIBBEAN AND GULF OF MEXICO (DIPTERA: CANACIDAE)
Wayne N. Mathis
Abstract.—The beach flies (Diptera: Canacidae) of the Caribbean and Gulf of Mexico, with focus on the species of Belize (Central America), are reviewed. The faunas from the Caribbean and Gulf of Mexico comprise four genera and seven species as follows: Canacea Cresson (C. macateei Malloch), Nocticanace Malloch (N. texensis (Wheeler); N. wirthi, new species; N. panamensis, new species), Paracanace Mathis & Wirth (P. aicen Mathis & Wirth; P. lebam Mathis & Wirth), and Procanace Hendel (P. dianneae Mathis). Keys and a diagnosis are provided for each genus and most species, and for Nocticanace, Paracanace, and Procanace a key to the species groups on a world basis is furnished. In addition, the texensis group of the genus Nocticanace 1s revised. Several struc- tures of each new species are illustrated to facilitate their identification.
In the published literature on the dipter- ous family Canacidae, more commonly known as beach flies, no mention is made of collection records from the Caribbean or Gulf of Mexico before 1950. Since then, the record is limited primarily to descriptions of a few new species (Wirth 1975). For in- dividual countries that are a part of this region, the paucity of information is even more acute—usually none exists. Thus for a country like Belize, which is a focus of this paper, there are no records of beach flies, elther under that geopolitical name or its predecessor, British Honduras. Although no species or collection records have been pub- lished from Belize, the occurrence of a few species was expected because genera such as Canacea Cresson and Nocticanace Mal- loch include species with distributions in adjacent countries (Wirth 1975). Like most projects on the systematics of Diptera, how- ever, especially those concerned with the neotropics, many of the observations and discoveries reported here were not antici- pated. To give greater meaning and per- spective to these findings, the results are reported within the context of a faunal re-
view of the beach flies of the Caribbean and Gulf of Mexico.
The impetus for this project resulted from field work on the insects of mangroves, sometimes called the mangal (Tomlinson 1986), that are associated with Belizean cays, especially those within the Stann Creek Dis- trict. With funding from the Caribbean Cor- al Reef Ecosystems Program (CCRE), field work has been conducted on the mangrove habitats of Twin Cays, with reconnaissance work on several of the nearby cays as well. On five field trips to these cays, I have made particular effort to collect specimens of the family Canacidae. This study also includes recently collected specimens from field work that I conducted in Cuba (1984), Dominica and St. Vincent (1989), and Florida (1989). In addition, W. W. Wirth made special ef- fort to collect beach flies on visits to Anti- gua, Dominica, Jamaica, and Puerto Rico and has graciously made these specimens available. A total of three species was col- lected and is reported here from Belize. This brings the total number of beach-fly species from the Caribbean and Gulf of Mexico to seven.
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Most of the specimens that were studied in conjunction with this paper are in the collections of the Smithsonian Institution (USNM). Others, especially primary types, were borrowed from other institutions. These institutions and their acronyms, as used in the text, and their respective cura- tors are as follows: CAS— California Acad- emy of Sciences, San Francisco, California (Dr. Paul H. Arnaud, Jr.); BDAF—Depart- ment of Agriculture and Fisheries, Botani- cal Garden, Paget, Bermuda (Dr. Daniel J. Hilburn); IZAC— Intituto de Zoologia, Aca- demia de Ciencias, Havana, Cuba (Mr. Jorge L. Fontenla).
For each genus and species treated in this review, a synonymy, diagnosis, the known distribution, and a remarks section are pro- vided. In the synonymies, only the literature that is pertinent to the Caribbean and Gulf regions or the species’ nomenclatural his- tory is cited. Specific collection data, except for the texensis group of the genus Nocti- canace, are cited only for Belizean speci- mens. The texensis group is more compre- hensively revised to include complete descriptions and a listing of all specimens examined. The descriptive format for the new species follows Mathis & Wirth (1978) and Mathis (1982, 1988). The illustrations of the male terminalia, especially the lateral views of the surstylus, were drawn from flat- tened, slide-mounted structures to ensure uniform views. In some specimens, the apex of the surstylus has a tendency to curve in- ward ventrally, partially obscuring the shape of that structure from a lateral view. A more detailed account of the morphology and higher classification of Canacidae can be found in Mathis (1982) and Wirth (1987). Two venational ratios are used commonly in the descriptions. Costal Vein Ratio: The straight line distance between the apices of R,,, and R,,,;/distance between the apices of R, and R,,;. M Vein Index: The straight line distance along vein M between cross- veins (dm-cu and r-m)/distance apicad of dm-cu.
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Key to Genera of Canacidae from the Caribbean and Gulf of Mexico
1. Lateroclinate fronto-orbital setae four or more; fore femur bearing row of usually from four to five spinelike setae along apical one-half of an- teroventral surface; apical one-third of arista bare; two supra-alar setae Hep h een ey ee ane S25, Canacea Cresson — Lateroclinate fronto-orbital setae three; fore femur lacking row of spinelike setae; arista evenly haired throughout length; one supra-alar seta 2. Intrafrontal setae absent, although anterior one-third of frons occa- sionally with scattered setulae .... 5 A A eae Procanace Hendel — Intrafrontal setae present, one or more pairs in addition to any setulae
3. One intrafrontal seta present; post- ocellar setae either much reduced or Packie. =.45. 3... Nocticanace Malloch
— Two intrafrontal setae present; post- ocellar setae well developed, procli- nate and slightly divergent
erage ee Paracanace Mathis & Wirth
Genus Canacea Cresson
Canacea |lapsus].— Malloch, 1924:52 [un- available; see Mathis, 1982, for discus- sion].
Canacea Cresson, 1924:164. Type species: C. macateei Malloch, by original desig- nation [Cresson validated Malloch’s name as an indication].— Mathis, 1982:4—7 [re- view].
Canace [in part].—Wirth, 1951:259-265 [review]; 1965:733 [nearctic catalog]; 1970:397-403 [revision]; 1975:1 [neo- tropical catalog].— Wheeler, 1952:90-91 [discussion].
Diagnosis.—Moderately small to mod- erately large beach flies, length 2.0 to 4.5 mm.
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Head: Mesofrons distinct from parafrons, shiny, very thinly microtomentose, bearing from three to five large, proclinate setae along lateral margin, mesofrons bare of se- tae; parafrons microtomentose, appearing dull; postocellar setae subequal in length to ocellar setae and with similar orientation; four large, lateroclinate, fronto-orbital se- tae; arista variable, usually with apical one- fourth to one-third bare of setulae, stylelike, some species with minute setulae extended nearly to apex but not appearing plumose; anaclinate genal setae two; anteroclinate ge- nal seta one.
Thorax: Dorsocentral setae 4 (1 + 3), all subequal in size; acrostichal setae conspic- uous, arranged in 4 irregular rows anterior- ly, becoming more regular posteriorly, mid- dle rows with setulae slightly larger and with large pair of prescutellar setae; lateral scu- tellar setae 2 pairs and with several setae dorsally; supra-alar setae 2, anterior seta only slightly smaller than posterior seta; 2 no- topleural setae; color of pleural setulae vari- able, usually black, all large setae black; propleuron bare of setulae; 1-2 large, anepisternal setae; katepisternal setae pres- ent; fore femur with row of 3-12 stout, spinelike setae anteroventrally; hind tibia lacking apical seta anteroventrally; apical section of vein M straight.
Abdomen: Female epiproct broad basal- ly, roughly triangular to nearly ellipsoidal, cerci as two parallel-sided narrow processes, each with one long, stout, apical seta; male surstylus with anteriorly curved hook.
Discussion.— Canaceais a New World ge- nus of four species that are primarily trop- ical in distribution. Mathis (1982) last re- viewed the genus.
Canacea macateei Malloch Figs. 1-12
Canacea macateei Malloch, 1924:52.— Cresson, 1924:164 [discussion]; 1936:265 [discussion].—Johnson, 1925:276 [list].— Mathis, 1982:7 [review].
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Canace snodgrassii [misidentification, in part].—Johnson, 1910:807 [list]. — Wirth, 1951:260 [synonymy]; 1965:733 [nearc- tic catalog].
Canace macateei.—Malloch, 1933:5 [note].—Curran, 1934:356 [status].— Wheeler, 1952:90-91 [distribution, key].— Wirth, 1970:399 [resurrection from synonymy, figure of male termi- nalia].—Teskey and Valiela, 1977:545- 547 [description of larva and puparium, natural history].
Specimens examined.—Belize. Stann Creek District: Twin Cays (Aanderaa Flats), 7-19 Nov 1987, W. N. & D. Mathis (28 6, 14 9; USNM); Twin Cays (S end of West Island), 17 Nov 1987, W. N. & D. Mathis (1 6; 1 9; USNM).
Distribution.—Canada (New Brunswick, Prince Edward Island) and United States (Atlantic and Gulf coasts from Maine to Texas), south to Cuba and Belize.
Natural history.—Teskey & Valiela (1977) successfully reared this species from larvae and puparia that were collected on mats of blue-green algae in Great Sippewissett marsh on Cape Cod, Massachusetts. Larvae of C. macateei, along with those of Dolichopod- idae, were the principal contributors to the biomass of invertebrates in the algal mats. Pupae were quite common, and the pupal stage is very brief, perhaps no more than two to three days. Adults were observed to ageregate, especially at night, on the tops of grass-covered dunes.
In Belize, we collected adults by sweeping an aerial net just above mats of blue-green algae that occurred along the mud banks of brackish pools. The surface of the mud banks, which is water covered only at higher tides, is broken by deep cracks that have divided the bank into irregularly shaped mud plates that are 15—25 cm in diameter. The algal mats were comparatively thick, and as a protective covering, they helped keep the underlying mud moist.
Remarks.—The species of Canacea are
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593
WS
ee =: \ M50 Yh. Nh ig : Yi Ks AN
2 ic 3
‘y J
SS (BMG, Wee) ate
Figs. 1-12. Scanning electron micrographs of Canacea macateei: 1, Head, lateral view; 2, Head, fronto- oblique view; 3, Head, anterior view; 4, Frons, dorsal view; 5, Fronto-orbital setae, dorsal view; 6, Ocellar triangle, dorsal view; 7, Antenna, lateral view; 8, Notopleuron and setae, lateral view; 9, Scutellum, dorsal view; 10, Katepisternum and setae, lateral view; 11, Left foreleg, anterior view; 12, Enlargement of left fore femur and tibia showing anteroventral, spinelike setae, anterior view.
quite similar externally but can be distin- guished by the structures of the male ter- minalia and the position, number, and color of certain setae. Specimens of C. macateei can be distinguished from congeners by the following combination of characters: fore
femur with a row of from three to five stout, anteroventral setae; the setulae of the pleu- ral sclerites are black; specimens are gen- erally larger, over 2.50 mm in length; and the surstylus of the male terminalia has the stem of the hook wider than long.
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Genus Nocticanace Malloch
Nocticanace Malloch, 1933:4. Type species: N. peculiaris Malloch, by original desig- nation.— Wirth, 1951:269-274 [revi- sion]; 1965:734 [nearctic catalog]; 1975: 2-3 [neotropical catalog].
Diagnosis. —Small to medium-sized beach flies, length 1.8 to 3.7 mm; general coloration grayish black.
Head: Intrafrontal setae one pair; post- ocellar setae either absent or much reduced, less than one-fourth length of ocellar setae; ocelli arranged to form an isosceles triangle, distance between posterior ocelli greater than that between either posterior ocellus and the anterior ocellus. Epistomal margin sinuate; clypeus low, width subequal to length of antenna. Two long anaclinate genal setae; anteroclinate genal seta moderately well de- veloped, at least one-half length of larger anaclinate genal setae. Palpus grayish black, bearing one to several long setae, each seta two to three times greatest width of palpus.
Thorax: Anepisternum with scattered setulae; proepisternal seta absent; katepi- sternal seta present, well developed. Legs entirely dark colored, grayish black; fore fe- mur bearing from four to six long and evenly spaced setae along posteroventral margin, length of setae at least equal to and usually greater than width of femur.
Discussion.—In preparing the following key to species groups, I have examined most species of the genus. The key, however, should still be considered preliminary, and I invite the critique and comment of users, as eventually I plan to review the remaining genera of Canacidae in a paper similar to Mathis (1982). The new species that are de- scribed below belong to the texensis group, which is also characterized and its species revised.
Annotated Key to Species Groups of the Genus Nocticanace
1. Anterior notopleural seta absent... 2 — Anterior notopleural seta present... 3
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2. Apical scutellar setae distinctly
curved anaclinate ...the pacifica group [19 species; Old World and Oceania]
— Apical scutellar setae straight to very
slightly curved . the fexensis group [3 species; Caribbean, Gulf of Mexico, Atlantic coast of south- eastern United States]
3. Length of apical section of vein CuA, twice or more length of crossvein dm-cu the galapagos group
[8 species; Galapagos Islands and southwestern North America]
— Length of apical section of vein cuA, subequal to length of crossvein dm-
CU .5.428.000... 2a 4
4. Apical scutellar setae distinctly an- aclinate the ashlocki group
[1 species, N. ashlocki Wirth; Galapagos Islands] — Apical scutellar setae not anaclinate Soe ee the chilensis group [1 species, NV. chilensis (Cresson); Chile (there are numerous unde- scribed species in this group)]
a6 0 = ae} ae fetes
The texensis Group
Diagnosis. —Acrostichal setae lacking; apical scutellar setae nearly straight in lat- eral view, slightly convergent in dorsal view, but not distinctly curved upward; anterior notopleural seta lacking, only a posterior seta present; proepisternal seta(e) present; mid femur of male lacking comblike row of setae; hind basitarsomere lacking spinelike basoventral setae. Wing with length of api- cal section of vein CuA, long, about twice length of crossvein dm-cu; M vein index 0.42-0.49.
Discussion.— Until now, the only includ- ed species in the texensis group was N. tex- ensis (Wheeler). The two new species added here are very similar and are very closely related to N. texensis. The three species to- gether are the only representatives of the
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PRO AWA
Figs. 13-21. view; 15, Head, dorsal view; 16, Gena and setae, lateral view; 17, Antenna, lateral view; 18, Fronto-orbital setae, dorsal view; 19, Right notopleuron and setae, lateral view; 20, Left katepisternum and setae, lateral view; 21, Scutellum, dorsal view.
genus Nocticanace that occur on beaches of the Atlantic Ocean (primarily within the Caribbean Sea and Gulf of Mexico). Other species of Nocticanace are found along coasts of the Indian and Pacific oceans. The species of the texensis group probably arose from a single lineage that originally came from the Pacific Ocean and then penetrated into and radiated within the Caribbean Sea and Gulf of Mexico.
As there are only three known species in the texensis group, I have not provided a key to distinguish between them. The di- agnoses, descriptions (only N. texensis is completely described; the others are com- pared to it), and figures (Figs. 13-29) ade-
595
& > : ' 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
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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
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(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.
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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.
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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
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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
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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
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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.
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. 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.
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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
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= 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.
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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-
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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.
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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.
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. 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-
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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-
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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)
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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
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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
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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-
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Table 1.— Material examined; Stenorhynchus seticornis.
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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 —
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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.
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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-
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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,
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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-
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Fig. 3. Scanning electron micrographs of the tips of male gonopods: Above, Stenorhynchus seticornis form A; Below, S. seticornis form B.
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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
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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.
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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;
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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
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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
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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
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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.
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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
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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.
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. 1925. The spider crabs of America. — Bulletin
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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.
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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-
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638
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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.
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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.
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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
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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
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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-
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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-
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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,
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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-
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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
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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
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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-
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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
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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
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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
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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.
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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-
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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
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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-
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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.
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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,
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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).
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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.
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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:
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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
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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.
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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
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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
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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
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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-
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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:
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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
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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
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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
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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
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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
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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
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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.
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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
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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
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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
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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-
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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
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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
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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
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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,
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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.
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. 1975. Procambarus (Girardiella) curdi, a new
crawfish from Arkansas, Oklahoma, and Texas
(Decapoda, Astacidae).— Tulane Studies in Zo-
ology and Botany 19(1, 2):22—25.
——., & William J. Clark. 1974. Decapod crusta- ceans of the Navasota River system in Central Texas. —Southwestern Naturalist 19(2):167-178.
Reinert, Howard Keith. 1978. The ecology and mor- phological variation of the Massasauga rattle- snake, Sistrurus catenatus. Ph.D. Thesis, Clar- ion State College, 193 pp.
Rhoades, Rendell. 1942. A list of the crawfishes of
Ohio.— Mimeographed “Leaflet”? of the Divi-
sion of Conservation and Natural Resources,
Department of Agriculture, State of Ohio, pp.
1-3.
. 1944. Further studies on the distribution and
taxonomy of Ohio crayfishes and the description
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
of a new subspecies. —Ohio Journal of Science 44(2):95-99.
1948. Notes on the crayfishes and their re- lation to farm ponds and agriculture.—Ohio Conservation Bulletin 12(1):18.
1950. Remarks concerning Lake Erie cray- fishes and a discussion of the origin of the Lake Erie crayfish fauna.— Wilmington Wildlife Re- search Project, Report No. 23:1—S.
1961. Lake Ene crayfishes and the origin of the crayfish fauna.—Bio-Briefs, Short Notes on Natural History 1:14.
Robison, H. W., & K. L. Smith. 1982. The endemic flora and fauna of Arkansas.— Proceedings of the Arkansas Academy of Science 36:52—57. Rogers, Robert L., & Jay V. Huner. 1983. Obser- - vations of life histories of crawfish on Southern University Baton Rouge campus.—ASB Bulletin 30(2):79.
1984. Observations on the life histories of crawfishes on the Southern University campus, Baton Rouge, Louisiana. P. 37 in Per Brinck, ed., Programme of the 6th International Sym- posium of Astacology, August 13-15, 1984, Lund, Sweden.
. 1985. 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.
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Stebbing, Thomas R. R. M. A. 1893. A history of Crustacea, Recent Malacostraca. D. Appleton & Company, New York, xvii + 466 pp.
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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.
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Underwood, Lucien M. 1886. List of the described
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, & Joe B. Black. 1967. New variations and
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1972. The occurrence and distribution of pteridines and purines in crayfish.—Crusta- ceana 22(3):294—302.
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LOGI, The
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oxygen consumption of the crayfish Orconectes immunis and Orconectes nais in response to temperature and oxygen saturation.—Physio- logical Zoology 34:39-54.
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(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).
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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
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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.
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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
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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.
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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.
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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.
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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
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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
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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
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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,
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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 <P 10 Nov xX 7 Jun xX xX 17 Nov xX 5 Jul X? 16 Dec xX 27 Jul xX xX 11 Jan 82 xX 9 Aug X? 20 Feb xX 18 Aug X X 3 Mar 4 x 29 Aug VS 11 Apr xX xX 21 Sep 2B
4 Station 614 only. > 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] <L BW Te Le C) rc 10 faa) > = < 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.
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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
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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-
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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
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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.
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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
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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.
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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).
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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
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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-
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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 <x em O 4.0 z ra ==) = 2 30 x 3. 2 = O z 2.0 x S Ee
4ST
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we we ods atpruf bom +4 ee
51 "6
Zines 39
10 11 12 13 14 15 16 17
NUMBER OF ABDOMINAL SETIGERS
Fig..3. 185 specimens of Euchone bansei.
of segments in the anal depression, and on the number of radioles. Observations were made on the shape and height of the collar.
The smallest individuals retained by the sieves are less than 1.0 mm in length and about 0.1 mm wide. At this size, all thoracic setigers and five or more abdominal setigers are present. The first segment in the anal depression usually appears as abdominal se- tiger 12, but in three of the 185 specimens measured it appears as abdominal setiger 11. Depression segments are added through abdominal setiger 17. No specimens were found with more than six depression seg- ments or 17 total abdominal setigers. In the three cases where the anal depression begins at setiger 11, the total abdominal setiger count is 16.
There is a Statistically significant corre-
Relationship between number of abdominal setigers and total body length (excluding radioles) for
lation between total body length and the width of the posterior thoracic segment (r? = 0.79, P < 0.005), revealing that E. bansei exhibits a very constant pattern of growth throughout its life. When total length is plot- ted as a function of the number of abdom- inal setigers, it is evident that most growth is accomplished after the majority of the segments are in place (Fig. 3). Sexual ma- turity is attained after the individual has grown considerably in total size and added most or all of the setigers. Gametes are found only in a few of the largest specimens having 16 abdominal setigers, and in many of the larger individuals having 17 abdominal se- tigers.
The height and shape of the collar de- pends on the size of the specimen. In the smallest individuals it is short, oblique, and
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0.2mm
*o ! Sat yf kt
yaA\EENY
Fig. 4. Collar development in juvenile specimens of Euchone bansei in lateral and dorsal view: A, Six abdominal setiger stage: B, Twelve abdominal setiger stage; C, Sixteen abdominal setiger stage.
relatively undeveloped on the dorsal side, wrapping closely against the anterior body (Fig. 4A). The collar lengthens dorsally and laterally, becoming more foliaceous as the worm adds abdominal setigers (Fig. 4B, C). By the time the specimen has become sex- ually mature the collar extends to the base
of the branchiae and is of nearly equal height all around (Fig. 1B).
Only two pairs of radioles are present on the smallest specimens examined. How- ever, even at this stage the branchiae have the long filiform free ends characteristic of the adults. By the time the abdomen has
Table 1.—Comparison of small species of Euchone: (1) Pre-depression setigers in the abdomen; (2) Abdominal depression setigers in adult specimens; (3) Pairs of radioles; (4) Length:width ratio of the blades of the inferior series of limbate thoracic notosetae; (5) Occurrence of ventral shields; (6) Ratio of the length of the free tips of the radioles relative to the total length of the radioles. Information is derived from the original descriptions and from Banse (1970).
Species ] 2 3 4 5 6 Euchone arenae Hartman, 1966 6-9 6 5-7 4:1 Present 1:6 Euchone elegans Verrill, 1873 12-15 8—10 6-8 S21 Present 1:6 Euchone hancocki Banse, 1970 5 3 4 10:1 Absent 3 Euchone incolor Hartman, 1965 6 3 g 5:1 Absent 1:4 Euchone rosea Langerhans, 1884 10-12 5-7 5-8 6:1 Present 1:10 Euchone bansei New species 11 6 4 9:1 Absent 1:2
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added the tenth setiger, a third pair of ra- dioles is developing ventral to the first two pairs. The fourth and final pair appears about the time the fifteenth abdominal setiger is developing.
Discussion
Euchone bansei is most similar to E. ro- sea Langerhans, 1884, from the eastern At- lantic off Europe and North Africa (Table 1), but it differs in lacking distinct ventral shields and in having fewer radioles, each with long rather than short filiform tips. Eu- chone bansei shares several characters with E. hancocki Banse, 1970, including four pairs of radioles with long free ends, a short and narrow first thoracic setiger, an inferior group of thoracic notosetae with narrow wings, and the absence of ventral shields. Euchone bansei, however, has a total of 17 abdominal setigers instead of eight as in E. hancocki, and it lacks a girdle of glands on the first abdominal setiger.
No seasonality could be detected in the growth stages of this species throughout the year. Sexually mature adults, juveniles with no depression setigers, and all intermediate stages were found in samples taken in the spring, summer, and winter months.
As noted by Banse (1970) for other small species of Euchone, all of the pre-depression setigers are in place before the first depres- sion setiger is formed. Sub-adult specimens have fewer depression setigers, and the smallest individuals entirely lack the depression, negating the utility of this char- acter for the specific determination of ju- venile specimens. Ontogenetic differences between the juveniles and adults of E. ban- sei also occur in the shape of the collar and the number of branchial radioles. These tra- ditional diagnostic characters, therefore, can be considered only when dealing with adults of the species of Euchone.
Several other characters, however, re- main constant as the specimens grow. The inferior series of thoracic notosetae have
759
narrow wings rather than being broadened, an unusual feature within the genus which is helpful in confirming the identity of small Euchone bansei. The elongated free ends of the radioles, present during all observed stages of growth, is another useful character. Staining with methyl green also affords a high degree of accuracy in distinguishing this species from other small sabellids occurring in the same sample. The juveniles have few- er stain-accepting cells per unit area than the adults, and some variability in the dis- tribution of individual gland cells 1s evident. However, the lack of stain-accepting cells in the intersegmental furrows and at the level of the parapodia results in a distinctly bian- nulate appearance to each setiger, and the pattern and uniform intensity of the staining regions when viewed in total are distinctive.
Acknowledgments
We wish to thank Drs. James A. Blake, Nancy Maciolek, Brigitte Hilbig, and two anonymous reviewers for valuable discus- sions and for critically reviewing the manu- script. Ms. Jennifer Taylor brought the ex- cellent holotype to our attention, and Ms. Barbara Greene assisted with typing and production of the manuscript. This work was partially supported by Contract No. 14- 12-0001-30064 from the U.S. Department of the Interior, Minerals Management Ser- vice, to Battelle Memorial Institute.
Literature Cited
Banse, K. 1970. The small species of Euchone Malm- gren (Sabellidae, Polychaeta).— Proceedings of the Biological Society of Washington 83(35):387— 408.
Blake, J. A., B. Hecker, J. F. Grassle, B. Brown, M. Wade, P. D. Boehm, E. Baptiste, B. Hilbig, N. Maciolek, R. Petrecca, R. E. Ruff, V. Starczak, & L. Watling. 1987. Study of biological pro- cesses on the U.S. South Atlantic slope and rise. Phase 2. Final Report prepared for the U.S. De- partment of the Interior, Minerals Management Service, Washington, D.C., under Contract No. 14-12-0001-30064. 11 + 414 pp. and 13 appen- dices. NTIS PB87-214359.
760
Hartman, O. 1965. Deep-water benthic polychaetous
annelids off New England to Bermuda and other
North Atlantic areas.—Allan Hancock Foun-
dation Occasional Paper 28:1-—378.
1966. Quantitative survey of the benthos of San Pedro Basin, Southern California. II. Final results and conclusions. — Allan Hancock Pacif- ic Expeditions 19(2):1-456.
, & K. Fauchald. 1971. Deep-water benthic
polychaetous annelids off New England to Ber-
muda and other North Atlantic areas. Part 2.—
Allan Hancock Monographs in Marine Biology
6:1-327.
Knox, G. A. 1977. The role of polychaetes in benthic soft-bottom communities. Pp. 547-604 In D. J. Reish & K. Fauchald eds., Essays on polychae- tous annelids in memory of Dr. Olga Hartman. Allan Hancock Foundation, Los Angeles.
Langerhans, P. 1884. Die Wurmfauna von Madeira. IV.—Zeitschrift fiir wissenschaftliche Zoologie 40:247-285.
Maciolek, N., J. F. Grassle, B. Hecker, P. D. Boehm, B. Brown, B. Dade, W. G. Steinhauer, E. Bap- tiste, R. E. Ruff, & R. Petrecca. 1987a. Study of biological processes on the U.S. Mid-Atlantic slope and rise. Final Report prepared for the
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
U.S. Department of the Interior, Minerals Man- agement Service, Washington, D.C., under Con- tract No. 14-12-0001-30064. ii + 314 pp. and 13 appendices. NTIS PB88 183090.
, ; , B. Brown, J. A. Blake, P. D. Boehm, R. Petrecca, S. Duffy, E. Baptiste, & R. E. Ruff. 1987b. Study of biological pro- cesses on the U.S. North Atlantic slope and rise. Final Report prepared for the U.S. Department of the Interior, Minerals Management Service, Washington, D.C., under Contract No. 14-12- 0001-30064. 358 pp. and 12 appendices. NTIS PB88 196514/AS.
Verrill, A. E. 1873. Report upon the invertebrate animals of Vineyard Sound and the adjacent waters, with an account of the physical char- acters of the region.— Report of the U.S. Com- mission of Fisheries 1871-1872 (I):295-747.
(RER) Battelle Ocean Sciences, 397 Washington Street, Duxbury, Massachu- setts 02332; (BB) Battelle Marine Sciences Laboratory, 439 W. Sequim Bay Road, Se- quim, Washington 98382.
PROC. BIOL. SOC. WASH. 102(3), 1989, pp. 761-767
PARALVINELLA HESSLERI, NEW SPECIES OF ALVINELLIDAE (POLYCHAETA) FROM THE MARIANA BACK-ARC BASIN HYDROTHERMAL VENTS
Daniel Desbruyéres and Lucien Laubier
Abstract. —A new alvinellid polychaete, Paralvinella hessleri, from the Mar- jana back-arc basin hydrothermal area is described. It is morphologically close to P. bactericola, P. palmiformis and P. grasslei, from active vents in the eastern Pacific. Biogeographical implications are discussed herein.
The first alvinellid polychaete, Alvinella pompejana, was collected by D.S.R.V. Alvin from honeycomb-like structures on the chimney walls of active hot vents at 21°N Eastern Pacific Rise hydrothermal site (Des- bruyéres & Laubier 1980). Since that time, specimens collected on American, French and Canadian submersible dives on the east Pacific ridges have expanded our knowledge of alvinellids. These worms until now have been exclusively sampled associated with warm and hot deep-sea vents in the eastern Pacific ridge system. Six species and one sub-species, belonging to two genera, have been described from the East Pacific Rise, Galapagos Ridge, Guaymas Basin, Explorer and Juan de Fuca Ridges (Desbruyéres & Laubier 1982, 1986, 1989). The examina- tion of all these taxa led us recently (1986) to propose the erection of a new family Al- vinellidae, which seems to be primitive within the order Terebellida.
Recently, an A/vin study of the Mariana Back-Arc Basin resulted in the discovery of two large vent areas at 3600-3700 m. The vent fields lie on the flank of active axial volcanoes (Craig et al. 1987) with a striking hydrothermal fauna. According to Hessler et al. (1987), patterns of faunal distribution at the vents are similar to those seen on the East Pacific Rise although the dominant or- ganisms belong to different and new taxa.
The alvinellid species described here were sent to us by R. R. Hessler of the Scripps Institution of Oceanography. The speci- mens were collected by submersible Alvin on the rocks directly exposed to venting water whose temperature was recorded up to: 25°C)
Order: Terebellida Family: Alvinellidae Desbruyéres & Laubier, 1986 Paralvinella hessleri, new species
Type locality, material examined. —Thir- ty three specimens collected during Alvin dives 1831 (04/16/87) at Ilium vent site (depth 3595 m, 18°12.8'N and 144°42.4’E), 1843 (05/04/87) and 1845 (05/06/87) at Al- ice Springs (depth 3640 m, 18°12.6'N and 144°42.4’E). Holotype (dive 1831) depos- ited in the collections of the Division of Worms, National Museum of Natural His- tory, Smithsonian Institution, Washington, D.C. (USNM 119431). Paratypes (number 5) from the same dive deposited in the col- lections of the “‘Laboratoire des Vers,’’ Mu- séum national d’Histoire naturelle de Paris (UC 90/A 908).
Etymology.—This species is named for Dr. Robert R. Hessler, Scripps Institution of Oceanography, leader of the biological study of the Mariana vents.
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Description. —Holotype 22 mm in length and 2.1 mm in greatest width with 60 se- tigerous segments (Fig. 1A). Paratypes range from 52 to 61 setigerous segments with the majority having 58 to 61. Color pinkish af- ter preservation in ethanol, with capillary setae yellow, modified setae brown. Body gradually tapering from about setigerous segment 30 to the end of the body. Ventral shields in the anterior third. .
Prostomium medially reduced with a me- dian incision and two well developed lateral lobes ventrally enclosing peristomium (Fig. 2A). Buccal apparatus complex, comprising a ventral globular bulky organ (a), two lat- eral, large and strong pointed tentacles (b) bearing a deep groove without ciliation, and many grooved and ciliated smaller tentacles (c) inserted in two groups on a quadrilobed upper lip (Fig. 3A). The whole apparatus or each part (e.g. ciliated tentacles, large ten- tacles and globular organ) eversible.
First segments (II and III) achaetous and fused to the first three setigerous; these five segments are not discernible ventrally. The first 15 to 20 setigerous segments with no- topodia only.
First three notopodia smaller than others and dorsally elevated. Branchial region formed by segments III to VI (an achaetous segment plus three setigerous segments). Branchiae four pairs, all similar, strong and regularly attenuated, and arranged in two adjacent groups. Branchial stem with large number of slender filaments inserted on two opposite narrow areas; branchial filaments cylindrical with small secondary filaments arising on two opposite lines. Setiger 4 (seg- ment VII) with a median dorsal expansion which protrudes forward (Fig. 1B).
- Notopodia, from setiger 4 to 13-17 (7 excepted), cylindrical with a dorsal digiti- form lobe (Fig. 3C) bearing two groups of capillary setae, one with short and the other with long. Setigerous segment 7 strongly modified, lacking cylindrical notopodia but bearing, on each side, 4 to 5 very large aci- cular notopodial-hooks directed posteriorly. Setigerous segment 8 with cylindrical no-
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
topodium and very strong digitiform lobe directed forward (Fig. 2B and 3C).
Cylindrical notopodia and uncinigerous neuropodial tori on each segment from seg- ment 15-20, both to end of body. Uncini numerous (20 to 50 per torus) in single rows, with teeth facing anteriorly (retrogressive situation). Uncini with one main tooth sur- mounted by one smaller secondary tooth (Fig. 3B) as in all other alvinellids. Pygidi- um rounded with inconspicuous papilla- tions.
Tubes whitish and corneous in aspect, amoeba-like in shape with long anchor fil- aments (Fig. 2D). Tube walls thick and mul- tilayered. Inner surface bearing huge fila- mentous bacterial mats.
Ecology. — All the specimens have been found in tubes on rocks directly exposed to venting water whose temperature was re- corded up to 25°C. The holotype was col- lected inside Ilium vent site from Whelk’s Club hot vent area near a hot smoker whose temperature is 282°C.
Discussion
Since the discovery of the first Alvinel- linae, another genus, five species and one sub-species have been described, all sam- pled from active hydrothermal vent areas of the eastern Pacific; a new family was erected within the order Terebellida (see Desbruyeéres & Laubier 1986, Holthe 1986) to accomodate these unusual polychaetes.
Specimens collected from the Mariana Back-Arc Basin hydrothermal vents un- doubtedly belong to Paralvinella and are morphologically close to Paralvinella gras- slei Desbruyéres & Laubier, 1982, P. pal- miformis Desbruyéres & Laubier, 1986, and P. bactericola Desbruyéres & Laubier, 1989. All share the following features: prostomi- um medially reduced with two lateral ex- pansions, four pairs of bipinnate branchiae with secondary slender filaments in oppo- site arrangement, uncini present posterior to the modified segment and buccal organ complex with large paired, grooved non-cil-
VOLUME 102, NUMBER 3 763
Fig. 1. Paralvinella hessleri: A, Habitus in latero-ventral view. Buccal apparatus invaginated; B, Anterior end in dorso-lateral view, branchiae removed.
iated tentacles. Paralvinella hessleri differs of the first five segments as in A/vinella spp. from these three species by the shape of the (Figs. 2B and 4).
buccal structures, the first appearance of the Paralvinella grasslei and P. palmiformis uncinigerous tori, the presence of a stouter are closely related, while P. bactericola dif- notopodial lobe on setiger 8 and the fusion fers from both by the structure of its buccal
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Fig. 4. Distinctive characters of the seven species and subspecies of Alvinellidae.
apparatus and the posterior position of the neuropodia. Due to its buccal apparatus with two large and strong pointed tentacles, P. hessleri has close relationship with P. bac- tericola. On the contrary, the pair of sibling species P. grasslei and P. palmiformis is characterized by two buccal structures end- ing in three rounded lobes. The ventral globular bulky organ of P. hessleri resembles the ventral lobe of P. grasslei and P. pal- miformis. Still, its absence in P. bactericola is not definitely established due to the small size of the sample and the possibility of in- vagination. The first neuropodium of P. hessleri (setigerous segment 18) appears in an intermediate position between P. grasslei (setigerous segment 13) and P. palmiformis (setigerous segment 20). In P. hess/leri, the fusion of anterior segments and the tenden-
cy towards reduction and constancy in number of segments can be considered as apomorphous conditions within Alvinelli- dae. At present, we can consider P. hessleri as a “recent’”’ species within the genus; it exhibits clear relationship with the group P. bactericola and P. grasslei and P. palmifor- mis.
The morphological likeness between P. bactericola and P. hessleri led us to assume a common ancestor. The Mariana Back-Arc Basin is located about 5000 nautical miles west from the eastern Pacific ridge system where all other Paralvinella have been found. This young back-arc basin, probably less than 10 million years old, has no physical connection with the mid-oceanic ridge sys- tem and is strongly isolated. Such habitat isolation, combined with close relationship
VOLUME 102, NUMBER 3
between P. hessleri and P. bactericola, led us to assume 1) the existence of very efficient dispersal mechanisms, or 2) the possibility that submarine volcanoes or organic matter patches could act as stepping stones for Par- alvinella dispersal. The youngest stage of Paralvinella, presently known, is a twelve- segmented erpochaeta of P. pandorae irlan- dei (Desbruyéres & Laubier 1986). By com- parison with a few examples of ampharetid species, we recently concluded that alvinel- lids should probably have a short larval life (Desbruyéres & Laubier 1986), while the opposite hypothesis could be supported by comparison with several terebellids. The hypothesis of submarine volcanoes acting as stepping stones for the dispersal of ben- thic species is supported by recent results on branchiate polynoids (Desbruyéres & Laubier 1988). Concerning the two sibling species P. palmiformis and P. grasslei, it is though that they arise from a common ancestor by allopatric speciation after the original area was separated into two distinct hydrothermal districts, north and south of the Oregon subduction zone some 26 mil- lion years ago by the overlapping American plate (Tunnicliffe 1988). At the opposite P. bactericola and P. grasslei dwell close to- gether in Guaymas Basin, demonstrating a strong sympatric speciation: the first asso- ciated with bacterial mats, the second with vestimentiferans and sulfides.
Acknowledgments
We wish to thank R. R. Hessler of the Scripps Oceanographic Institution for the material on which this study is based as well as scientists and crew members of the Mar- iana Back-Arc Basin expedition. We thank, too, M. A. Boudrias for his help and the chief scientist of the first leg H. Craig.
767
Literature Cited
Craig, H., Y. Horibe, K. A. Farley, J. A. Welhan, K. R. Kim, & R. N. Ney. 1987. Hydrothermal vents in the Mariana trough: Results of the first Alvin dives.— Eos 68(44):1531.
Desbruyéres, D., & L. Laubier. 1980. Alvinella pom- pejana gen. sp. nov. Ampharetidae aberrant des
sources hydrothermales de la ride Est-Paci-
fique.— Oceanologica Acta 3:267-274.
Oe 1982. Paralvinella grasslei, new
genus, new species of Alvinellinae (Polychaeta:
Ampharetidae) from the Galapagos rift geo-
thermal vents.—Proceedings of the Biological
Society of Washington 95:484—-494.
——.,& . 1986. Les Alvinellidae, une famille nouvelle d’annélides polychétes inféodées aux sources hydrothermales sous-marines: Systé- matique, biologie et écologie.— Canadian Jour- nal of Zoology 64(10):2227-2245.
——,, & 1988. Exploitation d’une source de matiére organique concentrée dans |’océan profond: Intervention d’une annélide polychéte nouvelle.—Comptes rendus de |’Académie des
Sciences de Paris, série III 307(6):329-335.
, & 1989. Systematics, phylogeny, biological cycle and ecology of the Alvinellidae from deep-sea hydrothermal vents.—Proceed- ings of the 2nd International Polychaete Con- ference, Ophelia.
Hessler, R. R., S. C. France, & M. A. Boudrias. 1987. Hydrothermal vent communities of the Mar- iana back-arc basin.— Eos 68(44):1531.
Holthe, T. 1986. Evolution, systematics and distri- bution of the Polychaete Terebellomorpha, with a catalogue of the taxa and a bibliography. — Gunneria 55:1-236.
Tunnicliffe, V. 1988. Biogeography and evolution of hydrothermal vent fauna in the eastern Pacific Ocean.—Proceedings of the Royal Society of London, ser. B 233:347-366.
(DD) Département Environnement Pro- fond, Centre de Brest de 1 TFREMER, B.P. 70, 29263 Plouzané, France; (LL) IFRE- MER, 66 avenue d’Iena, 75116 Paris, France.
PROC. BIOL. SOC. WASH. 102(3), 1989, pp. 768-771
A NEW SPECIES OF ODONTOSYLLIS (POLYCHAETA: SYLLIDAE) FROM TWIN CAYS, BELIZE
David E. Russell
Abstract. —Odontosyllis twincayensis is illustrated and described from Twin Cays, a mangrove island located on the Belizean barrier reef. The species is distinguished by setal blade shape and a unique combination of features that includes a distinct occipital flap, a trepan with 6 teeth, a large bulbous proventri- cle with 35 to 44 muscle cell rows, and bidentate setal blades of only one type.
An investigation of the distribution of syllid polychaetes in mangrove and adjacent shallow-water habitats at Twin Cays, Be- lize, revealed a multitude of species (Russell 1987). Forty-three syllid species were re- corded from 24 benthic cores. Among the over 7800 specimens examined were two individuals of a new species of Odontosyllis described below.
The material examined has been depos- ited in the National Museum of Natural History, Smithsonian Institution (USNM), Washington, D.C.
The generic definition is that of San Mar- tin (1984).
Odontosyllis Claparéde, 1863 Odontosyllis twincayensis, new species Fig. 1
Material examined.—West Bay, Twin Cays, Belize, Caribbean Sea; 20 cm depth, rootmat of Rhizophora mangle covered with a dense growth of Halimeda opuntia f. triloba; Nov 1983: holotype, core H-8U (USNM 102372); paratype, core H-1U (USNM 102373).
Comparative material examined. — Hutchinson Island, Florida, North Atlantic; 11.8 m; May 1972: Odontosyllis longigulata Perkins, 1981; holotype (USNM 60445); 2 paratypes (USNM 60447). Off North Car-
olina, North Atlantic; 130 m; Apr 1965: Odontosyllis longiseta Day, 1973; holotype (USNM 43120); 25 paratypes (USNM 43121). Barceloneta, Puerto Rico, North Atlantic; 23 m; Sep 1974: Odontosyllis lon- giseta, 1 specimen (USNM 52255) id. by C. Long.
Description. —Body pale yellow without color markings, pharynx pale amber. Body more or less cylindrical anteriorly. Holo- type a gravid female, 2.4 mm long, poste- riorly incomplete, with 16 setigers, 0.7 mm wide across proventricle without parapodia. Paratype a mature male (?), 0.64 mm long, posteriorly incomplete, with 7 setigers, 0.20 mm wide across proventricle without para- podia. Body length, width, and number of setigers in complete specimens unknown.
Prostomium oval, about twice as long as wide, with two pairs of large garnet eyes each with several small lens-like elements, posterior pair slightly smaller and closer to- gether than anterior pair (Fig. 1A). Paratype with third pair of smaller lensed eyes on anterior margin of prostomium; eyes of pair well separated, each immediately lateral to a lateral antenna. Antennae short, digiti- form, similar in size; median antenna orig- inating midway between anterior pair of eyes; lateral antennae arising from anterior margin of prostomium. Palps about as long as prostomium, directed ventrally, fused basally. Pair of C-shaped, ciliated nuchal
VOLUME 102, NUMBER 3
A dtc [
769
0.01 mm
E 0.1 mm
Fig. 1. Odontosyllis twincayensis, holotype (USNM 102372): A, Anterior end, dorsal view (slightly com- pressed dorsoventrally, prostomium and palps curving ventrally), setae omitted; B, Parapodium from setiger 13, posterior view, setae omitted; C, Aciculae from setiger 16; D, Inferior compound seta from setiger 16, scale same for C and D; E, Pharyngeal teeth and plates; dc = dorsal cirrus; dtc = dorsal tentacular cirrus; Ip = lateral plate; pa = palp; vt = ventral teeth; vtc = ventral tentacular cirrus.
organs along posterior margin of prosto- mium, medial portion of each extending an- teriorly, terminating between eyes; laterally, nuchal organs forming part of deep groove between prostomium and peristomium. Peristomium reduced dorsally, with clearly defined occipital flap extending anteriorly, partially covering nuchal organs. Two pairs of tentacular (peristomial) cirri, digitiform, smooth or with a few indistinct articula- tions; ventral tentacular cirri about as long as antennae, dorsal tentacular cirri about 1.5 times length of ventral tentacular cirri.
Parapodia divided distally into presetal and postsetal lobes (Fig. 1B). Two aciculae per parapodium in setigers 13 and 16, both dorsal to setal fascicle; each tapering distally to pointed tip with serrated subterminal ex- pansion or rim (Fig. 1C). Dorsal cirri mostly smooth, or with a few partial, indistinct ar- ticulations most common distally (Fig. 1A, B); longest dorsal cirrus on setiger 1, there- after dorsal cirri similar in length. Ventral cirri short, fusiform, partially fused to ven- tral surfaces of parapodia, not extending be- yond tips of parapodia. Dorsal and ventral
770
cirri of middle and posterior setigers, anal cirri, and pygidium unknown.
Simple setae not observed. Seventeen to 19 compound setae per parapodium in se- tigers 13 and 16; all blades on these para- podia similar in size and shape (Fig. 1D), approximately 19 wm long, maximum width about 4.4 um, blade-length to blade-width ratio 4.0—4.4, bidentate with similar, small terminal and subterminal teeth. Blade cut- ting edge straight to slightly convex, bearing short, coarse serrations along entire length. Proximal oblique edge of blade not discern- ible near blade cutting edge. Shaft heads of compound setae all similar, slender, about 5.3 wm wide, bearing short serrations.
Pharynx extending to setiger 6, trepan with ventral row of 6 teeth and 2 lateral plates (Fig. 1E). Proventricle with about 44 nar- row, indistinct muscle cell rows, occupying setigers 7 through 14, 0.98 mm long, about twice as long as wide. Paratype with phar- ynx extending to setiger 3, proventricle with about 35 narrow muscle cell rows, occu- pying setigers 3 through 5, 0.17 mm long, about twice as long as wide.
Etymology. —The specific name refers to the pair of mangrove islands, Twin Cays, where the type locality, West Bay, is located.
Remarks. —The description is of the ho- lotype unless otherwise indicated. Odonto- syllis twincayensis is distinguished from all previously described species of the genus by the unique combination of an occipital flap, a trepan with six teeth, a large bulbous pro- ventricle with about 35 to 44 muscle cell rows, and serrated, bidentate setal blades of only one type. Furthermore, the setal blades are longer than those of most Odontosyllis species, and are unique in having the shape ofa low triangle, a straight to slightly convex cutting edge with coarse serrations, and a bidentate tip with small, nearly equal ter- minal and subterminal teeth.
Odontosyllis twincayensis resembles O. longigulata Perkins, 1981, by having a tre- pan with six teeth and similar bidentate se- tal blades, but differs by having a well-de-
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
veloped, clearly defined occipital flap rather than a slight anterior peristomial fold, a proventricle with about 44 muscle cell rows rather than 60-70, and aciculae with point- ed rather than blunt truncate tips. The new species is similar to O. /ongiseta Day, 1973, in having an occipital flap, a trepan with six teeth, and long serrated, bidentate setal blades, but differs by having a proventricle with about 44 muscle cell rows, rather than 60, and setal blades that are somewhat tri- angular with a length-to-width ratio of about 4.0, rather than elongate with nearly parallel edges and a length-to-width ratio of 7.8.
Odontosyllis twincayensis also resembles O. glandulosa Augener, 1913, and O. gray- elyi Fauvel, 1928, particularly with regard to the length and bidentate aspect of the longest setal blades in these species, but dif- fers by having a much more triangular setal blade bearing coarse serrations, rather than a slender elongate blade with nearly parallel edges and a smooth cutting edge or one bearing very fine serrations. Odontosyllis twincayensis is further distinguished from these two species by having a distinct well- developed occipital flap and only one type of bidentate setal blade.
Acknowledgments
I wish to thank Dr. Kristian Fauchald, National Museum of Natural History (NMNH), Smithsonian Institution, Wash- ington, D.C., for kindly reviewing the manuscript. This paper represents a portion of a dissertation submitted to the Graduate School of Arts and Sciences of The George Washington University, Washington, D.C., in partial satisfaction of the requirements for the Ph.D. degree. I wish to thank Dr. Robert Knowlton, Department of Biologi- cal Sciences, for his guidance and encour- agement. I gratefully acknowledge financial support from The George Washington Uni- versity, the Lerner-Gray Fund for Marine Research (American Museum of Natural History), the Caribbean Coral Reef Ecosys-
VOLUME 102, NUMBER 3
tems (CCRE) Program (NMNH, Dr. Klaus Rutzler, Director), and the Exxon Corpo- ration. This paper is CCRE Contribution IiNe. 255.
Literature Cited
Augener, H. 1913. Polychaeta I: Errantia. Pp. 65- 304 in H. Michaelsen & R. Hartmeyer, eds., Die Fauna Sudwest-Australiens: Ergebnisse der Hamburger sudwest-australischen Forschungs- reise 1905, volume 4. Jena.
Claparéde, E. 1863. Beobachtungen uber Anatomie und Entwicklungsgeschichte wirbelloser Thiere an de Kiiste von Normandie angestellt. Leipzig, vii + 120 pp.
Day,J.H. 1973. New Polychaeta from Beaufort, with a key to all species recorded from North Car- olina.— NOAA Technical Report, National Ma- rine Fisheries Service Circular No. 375, 140 pp.
Fauvel, P. 1928. Annélides Polychétes nouvelles de l’Inde, part 2.—Bulletin du Muséum National d’Histoire naturelle 34(2):159-165.
Tet
Perkins, T. H. 1981. Syllidae (Polychaeta), princi- pally from Florida, with descriptions of a new genus and twenty-one new species.—Proceed- ings of the Biological Society of Washington 93(4):1080-1172.
Russell, D. E. 1987. The taxonomy and distribution of Syllidae (Annelida: Polychaeta) inhabiting mangrove and adjacent shallow-water habitats of Twin Cays, Belize. Ph.D. Dissertation. The George Washington University, Washington, D.C xv + 3889p:
San Martin, G. 1984. Estudio biogeografico, faunisti- co y sistematico de los poliquetos de la familia silidos (Polychaeta: Syllidae) en Baleares. Tesis Doctoral, Publicaciones de la Universidad Complutense de Madrid, ii + 529 pp.
Department of Biological Sciences, George Washington University, Washing- ton, D.C. (Present address) Department of Biological Sciences, Goucher College, Tow- son, Maryland 21204.
PROC. BIOL. SOC. WASH. 102(3), 1989, pp. 772-792
ORBINIIDAE (ANNELIDA: POLYCHAETA) FROM MANGROVE ROOT-MATS IN BELIZE, WITH A REVISION OF PROTOARICIIN GENERA
Vivianne Solis-Weiss and Kristian Fauchald
Abstract. — Benthic samples taken in root-mats of Rhizophora mangle contain four species of orbiniid polychaetes including Naineris setosa, and three new taxa, Protoaricia pigmentata, new species, Pettibonella multiuncinata, new ge- nus and new species and Pararicia belizensis, new genus and new species. Recognition of the new genera lead to a preliminary phylogenetic analysis of the genera of the subfamily Protoariciinae, brief characterizations of all genera in the subfamily and the development of a key to genera of the Protoariciinae
from world-wide areas.
The mangrove fauna of Twin Cays, Be- lize, has been under study for several years by a team of scientists under the direction of Dr. Klaus Rutzler of the Smithsonian Institution. As part of this overall program, a study of the fauna of the root-mats of red mangroves, especially where these are cov- ered by the green alga, Caulerpa verticillata was undertaken by Brian F. Kensley and Kristian Fauchald. The ecological findings will be reported elsewhere (Kensley & Fau- chald, in preparation). This paper is the first report on the polychaetes collected during the study; several additional papers are in preparation.
Members of the family Orbiniidae have been reported from sandy and muddy en- vironments world-wide. The subfamily Or- biniinae has been revised repeatedly (Day 1977, and references therein). The subfam- ily Protoariciinae has been less comprehen- sively treated. The presence of three new protoariciin taxa in material collected in Be- lize, including two that did not belong to any known genus, caused us to review the generic subdivision of the subfamily, to up- date and clarify as much as possible defi- nitions and to attempt a phylogenetic anal- ysis of the subfamily. The subfamily is here
considered monophyletic; a dubious as- sumption, but without access to very much larger materials than was currently available a more detailed study is not possible. Materials and methods.—The material was collected by K. Fauchald and B. F. Kensley as part of SWAMP (Smithsonian Western Atlantic Mangrove Program), di- rected by Dr. Klaus Rutzler. The sample localities include West Bay, Twin Cays, and the mainland side of the middle islands in Blue Ground Range (Fig. 1). The habitat sampled was covered with red mangrove forest (Rhizophora mangle) varying in height from approximately 1.5 to 5 m and in den- sity from open, isolated trees to dense forest with complete canopies. The microhabitat sampled was the root-mat where this mat was covered with a mat of Caulerpa verti- cillata. Part of the study includes a series of quantitative samples taken over a two-year period from 1979-1981. The samples were taken with a 10.4 cm diameter corer to a depth of approximately 10 cm in the sub- strate. The resulting core of the peat-like root-mat was gently broken up and screened through a 0.5 mm screen and preserved 1m- mediately in 10% neutralized formalin to which had been added Rose Bengal. After
VOLUME 102, NUMBER 3
| BELIZE
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24 hours in fixative, the samples were re- screened, and the specimens were sorted out in sea water, washed in freshwater and transferred to 70% alcohol. The samples were later sorted to family and counted.
All illustrations were made with the aid of a camera lucida attached to a stereo or compound microscope.
The morphological terminology is de- rived from Hartman (1957), Pettibone (1957) and Fauchald (1977). Any new terms used are explained in context. The literature cited include only papers directly used in this study; other papers can be found in the literature sections of the three papers men- tioned above. The cladistic analysis was run using the IBM microcomputer version of PAUP 2.4; details are indicated below in the section on cladistic analysis of the gen- era. The terminology follows the one estab-
lished by Wiley (1981). The character-list is given in Appendix | and the original data table in Table 2.
Station list. —As indicated above, all sta- tions were taken in two locations; each sam- ple consisted of a single numbered core; the core numbers for each of the two localities are given below.
West Bay, Twin Cays, Belize, 10-50 cm water depth; root-mat of Rhizophora man- gle, covered with Caulerpa verticillata core numbers M-1, M-2, M-3, M-5, M-9, M-10, M-11, M-12, M-27, M-32, M-35, M-SO0, M-51, M-55, M-58, M-59, M-70, M-71, M-88, M-90, M-95, M-96, M-102, M-105, M-107, M-133, M-136, M-137, M-139, M-140, M-142, M-143, M-144, M-145, M-146, M-147 and M-148.
West side of middle cay, Blue Ground Range, Belize, 10-50 cm water depth; root-
774
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Table 1.— Variability of selected morphological features of Pettibonella multiuncinata. All specimens included.
Range Mean SD Length in mm 1-20 8.01 Zi Number of setigers 31-110 66.83 OG Number of thoracic setigers 7-18 12.34 3.36 Branchiae from setiger number 6-16 Pe 4.18 Maximum number of rows of thoracic neuropodial uncini 2-5 3.3 0.79 Maximum number of uncini per row, thoracic neuropodia 2-7 4.16 1.39 Abdominal thin hooks first present from setiger number 7-19 13.20 4.63 Abdominal large hooks first present from setiger number 6-22 14.57 3.69
mat of Rhizophora mangle, covered with Caulerpa verticillata, core numbers M-23, M-24, M-78 and M-79.
In addition to the material newly iden- tified from the collections in Belize, we also examined type material and other materials as needed to verify our identifications and to clarify taxonomic uncertainties. This ma- terial is listed as previously identified ma- terial for each species.
Systematic Results Family Orbiniidae Hartman, 1942
The two subfamilies, Orbiniinae and Pro- toariciinae are currently separated only by the presence of one or two asetigerous an- terior segments (Fauchald 1977).
Subfamily Orbiniinae Hartman, 1957
Key to genera of this subfamily can be found in Day (1977).
Genus Naineris Blainville, 1828 Naineris setosa (Verrill, 1900) Figs. 2-3
Aricia setosa Verrill, 1900:651-653.
Anthostoma latacapitata Treadwell, 1901: 203-205, figs. 61-65.
Naineris setosa.—Hartman, 1942:61, figs. 116-118.—Hartman 1951:67-70, pl. 17, figs. 1-6.—Hartman, 1957:305, pl. 41, figs. 1-6.
Material examined.—Previously identi- fied material: Bermuda, Platts Inlet, 1898,
coll. A. E. Verrill and party (1 incomplete syntype, YPM 1242). Bermuda, 1901, coll. A. E. Verrill and party; id. M. Pettibone, 1962 (one incomplete specimen, YPM 1384). Bermuda, 1903, coll. W. R. Coe; id. M. Pettibone, 1962 (one complete speci- men, YPM 1303). Bermuda; id. M. Petti- bone (one incomplete specimen, USNM 34092). Bermuda, SE of Causeway, 1979, coll. & id. S. Gardiner (12 specimens).
Newly identified material: Belize, West Bay, Twin Cays and Blue Ground Range, 1979-1981, coll. K. Fauchald and B. F. Kensley M-11 (5, USNM 120928); M-12 (1, USNM 120932); M-23 (2, USNM 120935); M-24 (1, USNM 120938); M-90 (1, USNM 120955).
Description. — The description is based on the specimen referred as the syntype above, supplemented by notes on other material. No additional type material is currently available. The syntype is an incomplete fragment of 150 segments measuring 58 mm. It is widest at midthorax, 1.63 mm without, 3.83 mm with parapodia; the anterior ab- domen is 1.53 mm without and 2.83 mm with parapodia; the posterior abdomen is 1.33 mm without and 2.4 mm with para- podia. Color as preserved, brown.
The prostomium is broadly truncate to T shaped (Fig. 2a). Two diffuse, deeply embedded eyespots are present. Two shal- low, comma-shaped grooves are present dorsally at the posterior end of the prosto- mium.
The peristomium is a broad, asetigerous
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segment; the mouth is more than *3 of the total width of the peristomium. The ever- sible pharynx is not everted in the syntype; it is eversed and illustrated as present in another Bermudian specimen (YPM 1303, Fig. 2b).
The thorax consists of 20 biramous se- tigers, fully developed by setiger 5. The no- topodial postsetal lobes are foliaceous, broadest at two-thirds distance from the base. The neuropodial postsetal lobes are shorter, broader and rounder than corre- sponding notopodial lobes. They bear an upper digitiform papilla which is longer an- teriorly than posteriorly (Fig. 2c). Statocysts are visible as oval spots dorsally, antero- medial to the branchiae. Setigers 21 to 25 are transitional, characterized by a dimin- ishing number of neuropodial setae both in number of rows and in number of setae in each row. The parapodia become gradually more dorsal. The abdomen begins at setiger 25. The notopodial postsetal lobes become slenderer and progressively shorter towards the posterior abdomen. The corresponding neuropodial lobes become sharply reduced in size and foliaceous in shape (Fig. 2d), instead of round. Low dorsal ridges are pres- ent from the beginning of the abdomen, be- coming less conspicuous towards the end of the fragment. No neuropodial subpodial lobe is present either in anterior or middle ab- dominal segments.
Branchiae appear, in all specimens stud- ied, in setiger 6. However, in the syntype, a stout, unpaired bifid structure is found on one side in the position where branchiae are located on later setigers. This feature seems to be an abnormality of the specimen rather than a feature normally associated with the
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species (Fig. 2a). From setiger 6, normal paired branchiae appear. They are digiti- form, elongate, held erect over the body or recumbent along the dorsum. They are sim- ilar in length to the notopodial lobes but slenderer in the thoracic region, whereas in the abdominal region they are longer and broader than the corresponding notopodial lobes. They are present to the end of the fragment, and to the end of the body in all the complete specimens.
The thoracic notopodia have long cren- ulate capillary setae aligned in about three irregular rows and totalling approximately 25 or 30 setae per notopodium. The neu- ropodial thoracic setae are all crenulate cap- illaries. They are shorter than the notopo- dial setae and positioned in two bundles: 1). Approximately eight irregular, longitudinal palisaded rows, each bearing about 25 such setae, and 2). Approximately four irregular
Fig. 2. Naineris setosa: a, Anterior end of syntype, dorsal view; b, Anterior end of syntype, YPM 1303, showing evaginated pharynx; c, Right parapodium setiger 12, syntype, anterolateral view; d, Left parapodium, setiger 49, syntype, anterolateral view; e, Right parapodia, setigers 17-18, syntype, dorsal view; f, Abdominal uncinus, setiger 49, syntype; g, Furcate seta, abdominal setiger, CBC-M-11; h, Posterior end, CBC-M-11; i, Anterior end, dorsal view, CBC-M-11: j, Anterior end, dorsal view, CBC-M-90. Scales: a, b, e, 1 mm; c, d, h-
j, 100 um; f, g 10 um.
778
diagonal palisaded rows, posterior to the first bundle, each bearing 14 to 18 setae (Fig. 2e). In the abdomen, the number of setae decreases to about a dozen or less in both rami, although they are more abundant in the notopodium than in the neuropodium. Furcate setae are present in some abdomi- nal notopodia but are difficult to observe. They have a delicately spinous shaft and two distal tines of different length (Fig. 2g). In the abdominal neuropodia, in addition to the crenulate setae two or three straight, bluntly pointed uncini appear (Figs. 2d, f).
The syntype is incomplete, and the only complete specimen from YPM, has a dam- aged pygidium. The pygidium in a complete specimen from Belize (CBC-M-11, Fig. 2h) is elongate with three (probably originally four) elongate distally tapering anal cirri. Anal aperture 1s terminal and central.
Comparison of specimens from Belize and Bermuda. — The specimens newly identified agree fairly well with both type material and earlier descriptions. The main differences noted are:
1) The prostomium can be either T shaped or rounded.
2) In specimens with rounded prostomia numerous eyes are present, scattered be- tween the middle and the posterior end of the prostomium (Fig. 21). In specimens with T shaped prostomia, the eyespots are formed into two to four sickle shaped dark areas located at the posterior end of the prosto- mium; occasionally some additional small isolated spots are present (Fig. 2).
3) The number of thoracic setigers varies from! 3540 23%
4) The two separate groups of thoracic neurosetae are distinct only in larger spec- imens. Smaller specimens have only one bundle with fewer rows and fewer setae per row than in the syntype.
In order to determine if the number of thoracic setigers and the shape of the pro- stomium are size-related features, we did a least squares regression correlating the
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
number of thoracic setigers of the specimens with the length of the 15 first setigers. The results show a high correlation between the two (r = 0.91). There is no significant cor- relation between the different prostomial shapes and the size of the organism (r = 0.37).
Figure 3 shows that in general, the Belize specimens reach maximum number of tho- racic setigers at a smaller size than do the Bermuda specimens. There is no consistent trend in relation between numbers of tho- racic setigers and length in the material from Bermuda. Especially the syntype is very long in relation to the number of thoracic seti- gers. The differences among the populations are not sufficient to recognize them, even at the subspecies level, but are useful in allow- ing us to expand and quantify the descrip- tion of the species.
Hartman (1957) stated that neuropodial subpodial lobes should be present in this species; a feature not mentioned by Verrill (1900) in the original description nor men- tioned in any other review. We examined part of the material listed by Hartman (1957) including the specimens used to make the illustrations for that paper and failed to find subpodial lobes in any of the specimens. We assume that the subpodial lobes as men- tioned and illustrated by Hartman (1957) represent a /apsus calami, and that such lobes are normally absent in the species.
Habitat. —Subtidal, probably euryhaline species associated with vegetation (Thalas-
sia testudinum beds, algal mats and Rhi-
zophora mangle root-mats). Substrate may be sandy, sandy mud, or mangrove root- mats with minimal sediment. Locally pres- ent both in West Bay, Twin Cays and at Blue Ground Range (Fig. 1).
Distribution. —N. setosa has been report- ed from Bermuda (type locality), various localities in the Gulf of Mexico (Perkins & Savage 1975; Hernandez-Alcantara & Solis- Weiss 1989), Puerto Rico (Treadwell 1901), and Acapulco, Mexico (Hartman 1957).
VOLUME 102, NUMBER 3
Subfamily Protoariciinae Hartman, 1957 Preliminary Phylogenetic Analysis of the Protoariciin Genera
This analysis of putative relations among the protoariciin genera is based on several assumptions. First, the subfamily is as- sumed to be monophyletic; this assumption cannot be justified without a complete anal- ysis of the whole family, or indeed the order to which the family will eventually be re- ferred (its current assignment is unsatisfac- tory).
A second major set of assumptions can be summarized by the choice of the genus Leitoscoloplos among the Orbiniinae as out- group. Members of this genus are charac- terized first and foremost by lacking all modified setae in the thorax, and by the extreme simplicity of the acicular spines in the abdomen, in addition to the simple structure of the parapodial lobes and bran- chiae. The choice thus polarizes all more complex features, such as the presence of complex parapodial lobes, the presence and structure of various kinds of thoracic hooks and even the loss of certain features, such as branchiae, as apomorphic features. The procedure has the advantage of simplicity: without information to the contrary, it ap- peared simpler to assume that all more com- plex features were apomorphic, rather than randomly select some as being plesio- morphic and others as apomorphic. Reso- lution of this issue cannot come until pos- sible relations among all orbiniids and between the orbiniids and the related fam- ilies have been analyzed in detail.
The features used to characterize the gen- era are those traditionally used in orbiniid systematics (Day 1954, Fauchald 1977). The initial list contained 41 characters; the list was reduced to 31 characters by exclusion of features invariant among the taxa con- sidered (including the outgroup) and of cer- tain features that were so poorly known for most members of the group that they could
779
not be coded (numbers of abdominal seg- ments present for example). Appendix | lists the characters and character-states included in the analysis. Multistate characters are listed as transformation-series.
The character matrix was run on PAUP using the ALLTREES option (cfr. docu- mentation for PAUP as issued with the pro- gram).
Four trees were found, the consistency index was 0.670 and length was 88 for all four. All four trees plus a consensus tree is presented in Fig. 4. The four trees have sev- eral features in common. Protoariciella, Schroederella and Scoloplella are grouped together in all four trees. Orbiniella which is mainly characterized by the loss of var- ious features, nevertheless is defined by unique autapomorphies. In three of the four trees, Protoaricia and Pararicia show a unique synapomorphy and emerge jointly; in the last tree this character-state is inter- preted as having being a reversal. In all four trees the two genera emerge next to each other. The presence of the curved hooks (called swan-shaped in Proscoloplos) is a unique synapomorpy joining Pettibonella and Proscoloplos.
All internal nodes are supported by var- ious apomorphies in all four trees. All four trees are defined by synapomorphies. None of the nodes is exclusively supported by re- versals or parallellisms, or exclusively by synapomorphies created by various states in transformation series. All tree-topologies and a strict consensus tree are shown in Fig. 4. All taxa, including the two genera are supported by autapomorphies.
The consensus tree demonstrates that the summary given above cannot be expanded upon. The character-sequence used to de- fine the four trees differ and different trans- formation-series have been reversed in each tree. Without additional information the “correct” reading of this series cannot be confirmed.
The analysis was undertaken to examine
780 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Leitoscoloplos Leitoscoloplos Proscoloplos Proscoloplos Pettibonella Pettibonella Orbiniella Orbiniella Pararicia Protoaricia Leitoscoloplos TENET Pararicia Proscoloplos Protoariciella Protoariciella Pettibonella Scoloplella Scoloplella Orbiniella Schroederella Schroederella Protoaricia Leitoscoloplos Leitoscoloplos
Pararicia Orbiniella Proscoloplos Protoariciella Protoaricia Pettibonella Scoloplella Pararicia Orbiniella Schroederella Proscoloplos Protoaricia CONSENSUS DIAGRAM Pewbenele eee Scoloplella Scoloplella Protoariciella Protoariciella Schroederella Schroederella
Fig. 4. Cladograms showing possibly phylogenetic relations among the genera of the Protoariciinae. Further explanation in the text.
VOLUME 102, NUMBER 3
the level of support for previously described genera and compare them to the newly de- scribed genera. We feel justified in erecting the new genera: They represent unique com- binations of features otherwise not present in the subfamily, but recognize that the va- lidity of all genera may again be tested when the whole family is being analyzed.
Key to Genera of Protoariciinae
1. Branchiae absent Orbiniella SeGanenige Present ........... 4.4. 2
2. Transition between thorax and ab- domen indistinct Protoariciella
— Transition between thorax and ab-
domen distinct, transitional seg- ments may be present
3. Only crenulated capillaries present 200). Scoloplella
— Crenulated capillaries and other
sre] ae! (sae,
kemi@ssof Setae present ........... 4 4. Abdominal hooks acicular ....... 5 Abdominal hooks otherwise ..... i 5. Prostomium acutely pointed; anus SCSI” 6a a Schroederella — Prostomium distally rounded; or bluntly conical; anus terminal .... 6 6. Thorax with mucronate setae and subuluncini in addition to crenulat- eoueapillanicS .......:... Protoaricia — Thorax with crenulated capillaries CLS! . eee Pararicia 7. Abdominal hooks of a single kind ee aici 5.5 8 2 Proscoloplos — Abdominal hooks of two different L2SAGIS. pee Pettibonella
Brief Generic Characterizations
The new genera are defined in place in the text.
Orbiniella Day, 1954, type species O. minuta Day, 1954. Prostomium rounded or pointed. Branchiae absent. All thoracic se- tae crenulate. Thoracic notopodial setal lobes indistinct; notopodial postsetal lobes reduced; neuropodial postsetal lobes single, rounded. Two transitional segments pres-
781
ent. Abdominal setae crenulated capillaries, acicular setae and sometimes furcate setae. Anus terminal.
Proscoloplos Day, 1954, type species P. cygnochaetus Day, 1954. Prostomium rounded. Eyes absent. Branchiae from se- tiger 8. All thoracic setae crenulated capil- laries. Thoracic notopodial setal lobes in- distinct. Notopodial and neuropodial postsetal lobes tapering. Abdominal setae crenulated capillaries and one or two swan- shaped hooks. Anus terminal with four ta- pering anal cirri.
Protoaricia Czerniawsky, 1881, type species Aricia oerstedi Claparéde, 1864. Prostomium rounded. Two eyes. Branchiae limited to abdominal segments. Thoracic setae crenulated capillaries, hooks and sub- uluncini. Thoracic notopodial setal lobes distinct. Notopodial and neuropodial post- setal lobes tapering. No transitional seg- ments present. Abdominal setae crenulated capillaries and neuropodial uncini. Anus terminal with four blunt anal papillae or anal cirri absent.
In the original description of the type species, Claparéde (1864), stated that the dorsal (notopodial) rami in the abdomen were bifurcate; no types are available of any of Claparéde’s species (cfr. Fauchald, in prep.). Specimens from the Mediterranean Sea (off Malaga, Spain and off Marseille, France lack bifurcate abdominal notopodia (see discussion below).
Protoariciella Hartmann-Schroder 1962a, type species P. uncinata Hartmann-Schro- der, 1962a. Prostomium rounded. Two eyes. Branchiae from setiger 6 or 8. Thoracic no- tosetae all crenulated capillaries. Abdom1- nal notosetae crenulated capillaries and acicular setae. Neurosetae include crenulat- ed capillaries, thick, tridentate hooks, slen- der acicular setae with flattened teeth and thick, smooth spines. Separation between thorax and abdomen indistinct. Anus ter- minal; anal cirri absent.
Schroederella Laubier, 1962, type species S. pauliani Laubier, 1962. Prostomium
782 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
= eee “ee ere eC Le
Fig. 5. Protoaricia pigmentata: a, Anterior end, holotype, dorsal view; b, Left parapodium, setiger 6, holotype, anterolateral view; c, Hooded hook, setiger 5, holotype; d, Subuluncinus, setiger 5, holotype; e, Mucronate seta,
VOLUME 102, NUMBER 3
acutely pointed. Two eyes. Branchiae on ab- domen only. Parapodia poorly developed; thoracic notopodial postsetal lobes digiti- form, increasing in size posteriorly; thoracic neuropodial postsetal lobes single, rounded. Thoracic setae crenulated capillaries and straight neuropodial uncini. Abdominal no- topodial and neuropodial postsetal lobes ta- pering. Abdominal setae crenulated capil- laries and slender, pointed notopodial aciculae and hooded, thick neuropodial aciculae. Transitional segments present. Anus distinctly dorsal with 4 anal lobes.
Scoloplella Day, 1963, type species S. ca- pensis Day, 1963. Prostomium pointed. Eyes absent. Branchiae present from mid-ab- dominal segments. All postsetal lobes rounded. Parapodial rami reduced. All setae crenulated capillaries. Anus terminal.
The genus Scoloplosia proposed by Rul- lier, 1972, was synonymized with Protoar- icla by Ben-Eliahu (1976). This synonymy is here accepted.
Genus Protoaricia Czerniawsky, 1881 Protoaricia pigmentata, new species Fig. 5
Material examined.—M-78 (one, holo- type, USNM 120950, two paratypes, USNM 120951); M-79 (three paratypes, USNM 120952, one paratype Australian Museum; three paratypes British Museum (NH) and two paratypes Zoological Museum Ham- burg).
Description.—Holotype with 8 thoracic and 55 abdominal setigers for a total of 63; transitional setigers absent. Total length 5.4 mm; width at midthorax 0.8 mm and 0.54 mm in posterior abdomen. Length of other types 3 to 6.5 mm. Body somewhat flat- tened dorsoventrally; widest at midthorax. Color, as preserved, white with scattered
—
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brown dorsal pigmentation in branchial re- gion to mid-abdomen in some specimens. Brown, circular postsetal patch on each no- topodial thoracic lobe about two thirds from base, through mid-abdomen in most spec- imens (Fig. 5a, b).
Prostomium frontally round and wider at base. Two small round eyes deeply embed- ded, occasionally very difficult to see; lo- cated towards peristomial boundary (Fig. 5a). Peristomium and asetigerous segment clearly defined on all sides. Mouth, with lat- eral lips more than two thirds of ventral peristomial width. Pharynx not everted in any specimen.
All parapodia biramous. Thoracic noto- podial postsetal lobes elongate, cirriform; widest at proximal two-thirds of length (Fig. 5b). Thoracic neuropodial postsetal lobes shorter, wider and rounder than corre- sponding notopodial lobes; upper digiti- form papillae present on lobes (Fig. 5b). Ab- dominal parapodia located slightly more dorsally than thoracic ones (Fig. 51). Ab- dominal notopodial postsetal lobes similar to thoracic notopodial postsetal lobes; be- coming reduced in far posterior setigers. Abdominal notopodial and neuropodial postsetal lobes reduced last one to four se- tigers; upper papillae of neuropodia elon- gate; cirriform in anterior and mid-abdo- men; shorter in far posterior setigers.
Branchiae in holotype from setiger 8; missing on last two setigers; in paratypes from setigers 6—9 and missing in last two to four setigers; foliaceous, spionid-like, elon- gate, never overlapping; recumbent (Fig. 51). First pair shorter and slenderer than other branchiae. Branchiae longer and wider than notopodial lobes through mid-abdomen; thereafter distinctly reduced and more cir- riform.
Thoracic notopodial setae distinctly long-
setiger 5, holotype; f, Abdominal furcate notoseta, holotype; g, Abdominal neuropodial uncinus, holotype; h, Posterior end, dorsal view, holotype; i, Right parapodium, setiger 36, from one of the paratypes, anterolateral
view. Scales: a, b, h, i, 100 um; c—g, 10 wm.
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er than abdominal notopodial setae; capil- lary crenulate setae long, slender and more abundant in thorax than in abdomen; fur- cate setae present from thorax (Fig. 5f); sin- gle or at most two in a notopodium. Tho- racic neuropodial capillary crenulate setae shorter than corresponding notosetae; three or four mucronate setae, up to five subu- luncini and up to three thinly hooded, dis- tally tapering hooks present in thoracic neu- ropodia (Figs. 5c-e). Mucronate setae in upper end of setal bundles; subuluncini in middle and hooks in lower end of bundles (Fig. 5b). In the abdomen only three to five crenulate and one or two furcate neurosetae present; subuluncini, mucronate setae and hooded hooks absent; two to three slightly sigmoid, distally tapering hooks without hoods present (Figs. 51, g).
Pygidium elongate with four large pa- pillae; each terminated by a slender digiti- form cirrus of variable length (Fig. 5h). The anal aperture 1s central and terminal. Tubes absent.
Etymology. —The specific name refers to the characteristic brown color patterns pres- ent in specimens of this species.
Discussion. — Among the described species of Protoaricia, this species resembles P. oer- stedi (Claparéde) and P. capsulifera (Bob- retzky) more than P. minima (Rullier). The types of P. oerstedi and P. capsulifera are unavailable. The discussion is based on the original descriptions and illustrations and in the case of P. oerstedi, on observations on specimens from the Mediterranean Sea (Cap Couronne, near Marseille, France and near Malaga, Spain).
According to the literature (Claparéde 1864, Bobretzky 1870, Eisig 1914, and Fau- vel 1927), P. oerstedi and P. capsulifera are much larger (13 to 15 mm) than P. pig- mentata. P. capsulifera and P. pigmentata have very short segments, up to 77 for 6 mm in length; in contast P. oerstedi has only 52 segments for the same length.
In descriptions of P. oerstedi, branchiae are said to begin on the first abdominal se-
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
tiger, reported as setiger 12, the abdomen is flattened posteriorly, notopodial lobes are bifurcate and two or three straight aciculae are present in the posterior notopodia; only one to two uncini are reported present in each abdominal neuropodium (Claparéde 1864, Eisig 1914, Fauvel 1927). In the Med- iterranean material however, branchiae be- gin well after the first abdominal setiger (12— 14, but the thorax has only six to nine se- tigers).
In P. pigmentata the abdomen is nearly cylindrical; the notopodial lobes are never bifurcate; distinct abdominal aciculae are absent and we commonly found three ab- dominal neuropodial uncini, even in small specimens. Branchiae are present from the last thoracic setiger.
Possible differences in pygidial structures present a problem: Bobretsky’s illustration of P. capsulifera, shows the pygidium to be very similar to that of P. pigmentata. Fau- vel’s illustration of the pygidium of P. oer- stedi is in lateral view, making it impossible to determine accurately the distribution and length of various papillae; Fauvel’s descrip- tion is uninformative in that it only refers to the pygidium as having four short round cirri. The Mediterranean specimens have all short anal papillae rather than distinct cirri. Statocysts are present in both P. oerstedi and P. capsulifera and absent in P. pigmentata. In the description of P. capsulifera, no men- tion is made of the mucronate setae or sub- uluncini, nor are they illustrated.
Rullier (1972) did not mention the num- ber of thoracic setigers, the shape of the tho- racic region, or the shape of the pygidium for P. minima. Rullier (1972) reported branchiae absent and on the strength of this feature created a new genus, Scoloplosia for it: Ben-Eliahu (1976) synonymized it with Protoaricia since her largest specimen of the Same species had branchiae “from setigers 13 to 16’; thatis, from one of the abdominal setigers as in the other species of Protoari- cia. In addition, P. minima differs from our specimens in the following characters: Eyes
VOLUME 102, NUMBER 3
are absent in P. minima. The branchiae, are fingerlike from the start in P. minima, not foliaceous as in P. pigmentata. Ben-Eliahu (1976) did not mention presence of mu- cronate setae for P. minima and Rullier (1972) specifically stated that subuluncini and mucronate setae were absent in his ma- terial. P. minima also has one or two ab- dominal neuropodial uncini rather than three as present in P. pigmentata. Distribution. —The species is known only from Blue Ground Range, Belize (Fig. 1).
Pettibonella, new genus
Diagnosis. —Prostomium rounded or conical, usually with two eyespots. Two an- terior asetigerous segments. Branchiae de- ciduous, present from thoracic region, be- coming longer than notopodial postsetal lobes in abdominal region. Notopodial postsetal lobes well developed in thorax and abdomen, neuropodial postsetal lobes well
eveloped only in thorax. Notosetae in- clude crenulate capillaries only. Neurosetae in thorax and abdomen include crenulate capillaries (shorter than notosetae) and un- cini in thorax; a few crenulate capillaries and two different kinds of dentate hooks in abdomen. The pygidium with four digiti- form anal cirri.
Because of obvious close similarities be- tween Proscoloplos and Pettibonella, we compared Proscoloplos cygnochaetus, the type species, and P. confusus Hartmann- Schroder, 1962b, the only other species in the genus, to our new species. The type ma- terial of P. cygnochaetus (British Museum (Natural History), ZK 1955.3.20.1-6) was examined as were the types of P. confusus.
In Proscoloplos eyes are absent, rather than present. Branchiae are rounded, with glan- dular cells and much shorter than in Petti- bonella. Only a few crenulate capillaries are present in the thoracic setigers in Proscolop- los; these setae are abundant in Pettibonella, and in the latter there are, in addition, sev- eral neuropodial thoracic uncini. The dis-
785
tinctive swan-shaped hooks are present sin- gly or at most paired in Proscoloplos and they differ little in size or shape where paired; in Pettibonella two kinds of hooks, differing in size and shape are present.
Etymology.—This genus is named in honor of Dr. Marian H. Pettibone, Emeritus Zoologist of the Smithsonian Institution, in recognition of her excellent work on poly- chaete systematics.
Type species. —Pettibonella multiuncina- ta, new species.
Pettibonella multiuncinata, new species Fig. 6, Table 1
Material examined.—M-1 (one speci- men); M-2 (1); M-3 (1); M-9 (one paratype, USNM 120926); M-11 (1); M-12 (8); M-23 (13); M-24 (1); M-27 (5); M-32 (1); M-35 (4); M-50 (2); M-51 (2); M-55 (8); M-59 (1); M-70 (1); M-71 (1); M-88 (4); M-90 (2); M-95 (3); M-96 (6); M-102 (1); M-105 (1); M-107 (1); M-139 (4); M-140 (3); M-142 (1); M-143 (3); M-144 (4); M-145 (1, ho- lotype, USNM 120971, one paratype USNM 120972); M-146 (three paratypes, USNM 120973); M-147 (4); M-148 (1).
Description. —Holotype with 15 thoracic and 75 abdominal setigers, for a total of 90; total length 16mm, greatest width (in tho- rax) approximately 0.8 mm excluding para- podia. Body somewhat flattened dorsoven- trally, especially in anterior region. Color as preserved, white.
Prostomium conical, with two deeply embedded eyespots near peristomial boundary (Fig. 6a). Peristomium partially fused ventrally to next segment. Mouth two- thirds of ventral width of peristomium; lips lateral (Fig. 6b). Pharynx not seen in any specimen. Division between asetigerous segments distinct laterally, indistinct dor- sally or ventrally, but never simultaneously on both sides.
Branchiae from setiger 9; missing in last two setigers; elongate, flattened, broad based.
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Pans Bai," ms 5
nr
VOLUME 102, NUMBER 3
Some branchiae with middle constrictions (Fig. 6d). Branchiae deciduous with no ob- vious glandular cells; branchial surface with minute digitiform papillae (Fig. 6f). First branchiae shorter than notopodial lobes; in- creasing in length towards posterior end, be- coming between three and five times longer than notopodial lobes (Fig. 6e). Most bran- chiae held erect over the dorsum; some re- cumbent. All parapodia biramous.
First notopodium with filiform postsetal papillae; first neuropodium with similar, somewhat shorter postsetal lobes. Follow- ing notopodial postsetal lobes increasing in length. From setiger 2 neuropodial postsetal lobes unequally bilobed; superior part dis- tally rounded; inferior part digitiform and longer than superior part (Fig. 6c). Bilobed neuropodial postsetal lobes best developed at about setiger 5—6, decreasing towards end of thorax. Inferior part disappearing grad- ually towards end of thorax (Fig. 6d); absent in abdomen.
Thoracic setae of two kinds: crenulate capillary noto- and neurosetae, and neuro- podial uncini. First setiger with a bundle of approximately 10 notopodial crenulate cap- illary setae, and 15 or more neuropodial crenulate setae in a fan-shaped array. No- topodial crenulate setae increasing in length, but numbers remaining roughly constant through thorax. Crenulate neurosetae in- creasing dramatically in numbers; becom- ing arranged in irregular rows forming fan- shape total arrays. Four rows of crenulated neurosetae present in setigers 5—6. In seti- gers 13-15 number of crenulate neurosetae decreasing to six or seven. In abdomen two short crenulate neurosetae present. From setiger 2 to end of thorax, up to five uncini
Se
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in a vertical row ventralmost in each neu- ropodium. Uncini yellow, shafts straight; distally bent, blunt tipped; with 7-12 flat- tened transverse scales (Fig. 6g); tips some- times worn resulting in tooth-like structures being formed (Fig. 6h). In transitional seg- ments (13-15) uncini with two terminal teeth.
Transition from thorax to abdomen marked at setiger 16 by reduction in number of neuropodial crenulate setae and replace- ment of thoracic uncini by two tridentate hooks (Fig. 61). Transition region also with reduction of postsetal lobes and progres- sively more dorsal position of parapodia.
Tridentate hooks without hoods located ventrally with cutting edges facing dorsally (Fig. 61). Hooks of setiger 16 intermediate in shape between thoracic uncini and fully formed hooks of middle abdomen (i.e. teeth present but hooks more elongate than in following setigers). From setiger 17, another kind of hook present, facing the other two hooks in a vis-a-vis position. In setigers 1 7— 20, all hooks increasing gradually in width, especially subterminally. Where fully de- veloped, large hooks with large main fangs surmounted by four denticles in a rhomboid arrangement (Fig. 6j, k); shafts distinctly in- flated below rostrum. Usually one large hook and two slender hooks in a setiger; occa- sionally and scattered, some setigers with two large hooks parallel to each other; facing either two or three slender hooks. Branchiae distinctly reduced and hooks are absent in last three parapodia and last two parapodia asetigerous.
Pygidium elongated with four slender, digitiform cirri; two dorsal cirri longer than ventral ones. Pygidial cirri retracted or
Fig. 6 Pettibonella multiuncinata: a, Anterior end, holotype, dorsal view; b, Anterior end, holotype, ventral view; c, Left parapodium, setiger 6, holotype, anterolateral view; d, Left parapodium, setiger 12, holotype, anterolateral view; e, Left parapodium, setiger 60, holotype, anterolateral view; f, Branchial edge, setiger 60, holotype; g, Thoracic uncinus, setiger 6, holotype; h, Thoracic uncinus, setiger 12, holotype; i, Slender neuropodial abdominal hook, setiger 82, holotype; j, Large neuropodial abdominal hook, setiger 82, holotype; k, Large neuropodial abdominal hook, setiger 82, holotype, view from distal end; 1, Posterior end, ventral view, holotype; m, Pygidium, CBC-M-9. Scales: a—e, 1, m, 100 wm; f-k, 10 um.
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Fig. 7. Pararicia belizensis: a, Anterior end, holotype, dorsal view; b, Right parapodium, setiger 8, holotype, anterolateral view; c, Right parapodium, setiger 33, holotype, anterolateral view; d, Posterior end, holotype, dorsal view; e, Abdominal uncini, setiger 33, holotype; f, Furcate notoseta, setiger 9, holotype. Scales: a—d, 100
um: e, f, 10 um.
damaged (Fig. 61) in some specimens. Anal aperture terminal, central; surrounded by about nine papillae (Fig. 6m).
Holotype without tube; some paratypes are covered with fragments of tubes in mid- dle abdominal region. Fragments thin, transparent and covered with sand grains of varying sizes; very small shell fragments and vegetal debris.
Etymology. —The specific name refers to the very distinctive hooks present in this species.
Discussion. —The occurrence of some fea-
tures is size dependent: The total number of setigers can vary from 32 to 106. The branchiae usually appear at setiger 9 but may be present from setiger 6 to 8. The number of thoracic segments and hence the first appearance of the abdominal hooks, is also size dependent. The variation of these and other characters is summarized in Ta- ble?
Habitat. —The species is equally well rep- resented both at Twin Cays and Blue Ground Range (Fig. 1), taking the relative sample density into account.
VOLUME 102, NUMBER 3
Pararicia, new genus
Diagnosis. —Prostomium rounded, with two to numerous eyes, two asetigerous tho- racic segments present; branchiae from one of the thoracic setigers. Notopodial setae include crenulate capillaries and furcate se- tae. Thoracic neuropodial setae shorter than crenulate capillary setae; thoracic un- cini absent. Abdominal neuropodial setae, a few crenulate capillaries in addition to smooth acicular uncini. Four fingerlike anal cirri present.
Etymology.—Derived from the old ge- neric name Arvicia used in this family.
Type species. —Pararicia belizensis, new species.
Pararicia belizensis, new species Fig. 7
Material examined.—M-3 (one speci- men); M-5 (4); M-10 (1); M-11 (one, ho- lotype, USNM 120930, one paratype, USNM 120931); M-12 (2); M-23 (5); M-24 (one paratype, USNM 120939); M-35(1); M-88 (two paratypes, USNM_ 120953); M-133 (one paratype, USNM 120962); M-135 (1); M-136 (1); M-147 (1).
Description. —Holotype complete with 68 setigers; length 4 mm, greatest width, in tho- rax, approximately 470 wm, without para- podia. Body slightly more flattened dorso- ventrally and wider in thoracic than in abdominal region. Abdomen tapering pos- teriorly with reduced parapodia in last 15 setigers. Length of other complete speci- mens from 2 to 9 mm; number of setigers from about 30 to 70. Color as preserved white. Thorax with 10 setigers; abdomen with 58 setigers, including first four tran- sitional setigers.
Prostomium rounded with many eye- spots. Two of those are round, clearer and present at the posterior end of the prosto- mium. The rest are divided in two roughly comma-shaped groups of eyespots at the middle region of the prostomium. No ap-
789
pendages are present (Fig. 7a). In smaller specimens, only two small, round, widely separated eyes are present, near the poste- rior boundary with the peristomium.
The first two asetigerous segments are dis- tinctly separated from each other and from the prostomium. Mouth about '2 of peri- stomial width, with lateral lips. Pharynx not everted in any specimens.
Branchiae from setiger 6 in all specimens; becoming reduced in last 15 setigers and absent in last 2-3 setigers; flattened, elon- gate, triangular, widely separated, never overlapping and recumbent. Mid-abdomi- nal branchiae somewhat larger than other branchiae; otherwise all branchiae similar in size; slightly shorter than notopodial lobes in thorax (Fig. 7b); not deciduous. The last few branchiae are rudimentary and are not visible in the illustration.
All parapodia biramous. Thoracic noto- podial postsetal lobes elongate, cirriform, broader in proximal 7% of length; slightly increasing in length in first setigers. Tho- racic neuropodial postsetal lobes shorter, wider, distally more rounded than corre- sponding notopodial lobes. Median papillae present; tapering distally (Fig. 7b). Setigers 11 to 14 transitional, characterized by grad- ual reduction in number of neurosetae and by dorsal shift in neuropodia. Abdominal notopodial postsetal lobes similar to tho- racic notopodial postsetal lobes. Abdomi- nal neuropodial postsetal lobes increasingly reduced in length; retaining the same shape (Fig. 7c). Low, dorsal transverse ridges pres- ent on abdominal segments.
Both rami with bundles of capillary cren- ulate setae; thoracic notopodia with about 8-10 setae; thoracic neuropodia with usu- ally 12-18 up to 30 setae; neuropodial fas- cicles in rows of spreading setae. Notopodial crenulate capillaries longer than neuropo- dial ones throughout. Uncini absent in tho- racic region. Abdominal setigers with re- duced numbers of crenulate_ setae; neuropodia with only four or five setae. Some abdominal notopodia with furcate se-
790
tae (Fig. 7f). Each abdominal neuropodium with one or two stout hooks. Hooks straight to slightly sigmoid, bluntly pointed, without hoods (Fig. 7e). Small specimens (M-133, M-136, M-147) with hooks from first ab- dominal setiger. Larger specimens with hooks from the first post-transitional seti- ger.
Pygidium elongate with four cirriform, distally tapering anal cirri; all anal cirri sim- ilar in size. Anal aperture central; terminal (Fig. 7d).
Tubes absent.
Discussion. —The species differs from re- lated taxa as indicated in the discussion of the protoariciin genera.
Etymology. —The specific name refers to the country of origin of the type material.
Habitat.—The species was found mainly in Twin Cays with the exception of M23 and M24 at Blue Ground Range (Fig. 1).
Acknowledgments
This paper is Contribution number 265 from the Caribbean Coral Reef Ecology Program and the SWAMP, both programs under the direction of Dr. Klaus Rutzler. We would like to thank Dr. Rutzler for all his help during the collection of materials. The junior author would also like to thank Dr. Brian F. Kensley, his collaborator in the benthic studies in Belize, for many years of collaboration on collecting, sorting and pre- serving benthic samples in Belize and else- where. We would like to acknowledge the curators in charge of the collections at Yale Peabody Museum; Zoologisches Museum und Staatsinstitut, Hamburg; British Mu- seum (Natural History); and Allan Hancock Foundation, University of Southern Cali- fornia. Dr. Gerard Bellan, Station Marine d’Endoume, Marseille, France and Dr. Guillermo San Martin, Fac. Ciencias, Univ. Aut. Madrid, Spain both donated material of orbiniids from the Mediterranean Sea. Dr. Meredith L. Jones lent us material of Naineris setosa from Bermuda. The senior
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
author would like to thank Instituto de Ciencias del Mar y Limnologia, UNAM, Mexico, and its Director, Dr. A. Ayala, DGAPA (UNAM) and the Organisation of American States for financial support, mak- ing her stay at the Smithsonian Institution possible.
Literature Cited
Ben-Eliahu, M. N. 1976. Polychaete cryptofauna from rims of similar intertidal vermetid reefs on the Mediterranean coast of Israel and in the Gulf of Elat: Sedentaria.—Israel Journal of Zoology 25: 121-155.
Bobretsky, N. 1870. [On the Fauna of the Black Sea] (In Russian).— Kiev odschestva estest. Zapisky 1:188-274.
Claparéde, E. 1864. Glanures zoomotiques parmi les Annélides de Port-Vendres (Pyrénées Orien- tales). — Mémoires de la Société de Physique et d’Histoire Naturelle de Genéve 17(2):463-600.
Czerniavsky, V. 1881. Materialia ad Zoographiam ponticam (continuatio).— Bulletin de la Societé Impériale des Naturalistes de Moscou 56(1):338- 420.
Day, J. H. 1954. The Polychaeta of Tristan da
Cunha.—Results of the Norwegian Scientific
Expedition to Tristan da Cunha 1937-1938, Det
Norske Videnskaps-Akademi, Oslo 29:1-35.
. 1963. The Polychaete fauna of South Africa.
Part 8. New species and records from grab sam-
ples and dredgings.— Bulletin British Museum
(Natural History) (Zoology) 10(7):384-445.
1977. A review of the Australian and New
Zealand Orbiniidae (Annelida: Polychaeta).—
Pp. 217-246 Jn D. J. Reish & K. Fauchald eds.,
Essays on polychaetous annelids in memory of
Dr. Olga Hartman. Allan Hancock Foundation,
University of Southern California.
Eisig, H. 1914. Zur Systematik, Anatomie und Mor- phologie der Ariciiden nebst Beitragen zur ge- nerellen Systematik.— Mitteilungen aus der Zoologische Station zu Neapel 21(6):153-600.
Fauchald, K. 1977. The Polychaete worms. Defini- tions and keys to the orders, families and gen- era.— Natural History Museum of Los Angeles County, Science Series 28:1-188.
Fauvel, P. 1927. Polychétes Sédentaires. Addenda aux Errantes, Archiannélides, Myzosto- maires.—Faune de France 16:1-495.
Hartman, O. 1942. A review of the types of poly- chaetous annelids at the Peabody Museum of Natural History, Yale University.— Bulletin of the Bingham Oceanographic Collection, Pea-
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body Museum of Natural History, Yale Uni-
versity 8(1):1-98.
1951. The Littoral marine annelids of the Gulf of Mexico.— Publications of the Institute of Marine Science, 2(1):7—124.
1957. Orbiniidae, Apistobranchidae, Para- onidae and Longosomidae. — Allan Hancock Pacific Expeditions 15(3):183-393, pls. 21-44. Hartmann-Schroder, G. 1962a. Zweiter Beitrag zur Polychaetenfauna von Peru.—Kieler Meeres- forschungen 18(1):109-147.
. 1962b. Die Polychaeten des Eulitorals. Jn G.
Hartmann-Schroder, & G. Hartmann: Zur
Kenntnis des Eulitorals der chilenischen Pazi-
fikkiiste und der argentinischen Kiste Siidpa- tagoniens unter besonderer Berucksichtigung der Polychaeten und Ostracoden.— Mitteilungen aus dem Hamburgischen Zoologischen Museum und Institut 60:57-270.
Hernandez-Alcantara, P. & V. Solis-Weiss. 1989. Ecological aspects of the polychaete populations associated to the red mangrove Rhizophora mangle at Terminos Lagoon, southeastern part of the Gulf of Mexico.— Ophelia (in press).
Laubier, L. 1962. Schroederella pauliani gen. nov., sp. nov., un nouvel orbiniide (Polychétes Sé- dentaires) de la faune interstitielle d’Afrique. — Annals of the Transvaal Museum 24:231-238.
Perkins, T. H., & T. Savage. 1975. A Bibliography and checklist of polychaetous annelids of Flor- ida, the Gulf of Mexico, and the Caribbean Re-
791
gion.—Florida Marine Research Publications, Florida Department of Natural Resources, Ma- rine Research Laboratory 14:1-62.
Pettibone, M. H. 1957. North American genera of the family Orbiniidae (Annelida: Polychaeta), with descriptions of new species.—Journal of the Washington Academy of Sciences 47(5):159- 167, figs. 1-4.
Rullier, F. 1972. Annélides polychétes de Nouvelle- Calédonie recueillies par Y. Plessis et B. Sal- vat.— Expédition Frangaise sur les récifs coral- liens de la Nouvelle-Calédonie 6:1-169.
Treadwell, A. L. 1901. The polychaetous annelids of Porto Rico. — Bulletin of the United States Fish- eries Commission 20:181-210.
Verrill, A. E. 1900. Additions to the Turbellaria, Nemertina, and Annelida of the Bermudas. — Transactions of the Connecticut Academy of Arts and Sciences 10(2):595-671.
Wiley, E. O. 1981. Phylogenetics. The Theory and practice of phylogenetic systematics. John Wiley and Sons, New York, XV and 439 pp.
(VSW) Instituto de Ciencias del Mar y Limnologia— UNAM Apdo Postal 70-305. Mexico, D.F. 04510, México; (KF) De- partment of Invertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington D.C. 20560.
af
792 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Appendix 1.—Character-list for protoariciine genera.
1. Prostomial shape 1. pointed or conical 2. rounded or truncate 3. both 2. Branchial start 0. branchia absent 1-8. setiger on which branchia begin 3. Shape of branchiae 0. branchia absent 1. triangular (flattened) 2. fusiform 3. foliaceous 4. cirriform 4. Anterior and posterior branchiae 0. all branchiae absent 1. similar 2. dissimilar 5. Glandcells in branchial walls 0. all branchiae absent 1. present 2. absent 6. Thoracic furcate notosetae 1. present 2. absent 7. Thoracic acicular notosetae 1. present 2. absent 8. Thoracic acicular neurosetae 1. present 2. absent 9. Thoracic neuropodial subuluncini 1. present 2. absent 10. Thoracic neuropodial uncini @ 1. present 2. absent 11. Thoracic notopodial setal lobes 1. distinct 2. indistinct 12. Thoracic notopodial postsetal lobes 1. tapering 2. fusiform
3. digitiform (increasing in length through tho-
rax) 13. Thoracic neuropodial postsetal lobes 1. single 2. double 14. Thoracic neuropodial postsetal lobes 1. rounded 2. tapering
ey
16.
17.
18.
20.
ZAG
pipe
23:
24.
ye
26.
Die
28.
DS
30.
31
Number of transitional segments
0-7. number of transitional segments Abdominal setal lobes
1. distinct
2. indistinct
Abdominal neuropodial postsetal lohes 1. tapering
2. rounded
Abdominal notopodial furcate setae 1. present
2. absent
. Abdominal notopodial acicular setae
1. present
2. absent
Abdominal neuropodial crenulate setae 1. present
2. absent
Abdominal neuropodial acicular setae 1. present
2. absent
Abdominal neuropodial subuluncini 1. present
2. absent
Abdominal neuropodial uncini
1. present
2. absent
Abdominal swan-shaped hooks
1. present
2. absent
Abdominal crested hooks
1. present
2. absent
Abdominal hooks
1. of a single kind
2. of two kinds in vis-a-vis rows Pygidium
1. with short, blunt projections
2. with distinct pygidial cirri Number of anal projections or cirri 0-6. number of anal cirri
All anal cirri
1. similar
2. of two or more different kinds Eyes
1. absent
2. paired
3. more than a pair
Distinct nuchal organs
1. present
2. absent
er
PROC. BIOL. SOC. WASH. 102(3), 1989, pp. 793-804
THREE NEW SPECIES OF MYZOSTOMA (MYZOSTOMIDA)
Mark J. Grygier
Abstract. —Myzostoma armatae, new species, is an ectocommensal of Analci- dometra armata (Pourtalés) in the Bahamas and Jamaica, and of Davidaster discoidea (Carpenter) in Jamaica. Myzostoma attenuatum, new species, is a widespread ectocommensal of diverse Indo-Pacific comatulids. Myzostoma divisor, new species, is an Antarctic and sub-Antarctic ectocommensal of Pro- machocrinus kerguelensis Carpenter and of Notocrinus mortenseni John. The post-settlement ontogeny of M. divisor is documented by scanning electron
microscopy.
Myzostoma Leuckart is the largest genus of the Myzostomida, a group of obligately echinoderm-associated worms of unsettled zoological affinities. Currently 115 species of Myzostoma are recognized; most of them are ectocommensals of crinoids, although a few form galls on crinoids or infest ophiu- roids. The last significant taxonomic paper on this genus was by Jagersten (1940a). My recent work on myzostomes isolated from several major crinoid collections has un- covered many undescribed species. Most of these are represented by few specimens, but the three new species described here are no- table for their abundance.
Abbreviations for museums mentioned in the text include: National Museum of Natural History (USNM), British Museum (Natural History) (BMNH), Northern Ter- ritory Museum of Arts and Sciences (NTMAS).
Class Myzostomida Graff, 1877 Order Proboscidea Jagersten, 1940b Family Myzostomatidae Graff, 1884
Genus Myzostoma Leuckart, 1836 Myzostoma armatae, new species Fig. 1
Diagnosis. —Small species with flat, oval body, narrower toward rear, posterior end
truncate in uninjured specimens. Marginal zone narrow or poorly evident. About 10 pairs of short marginal cirri, gaps between them larger behind pair 8; supernumerary cirri common. Parapodia acirrate, restricted to anterior 70% of length. Manubrium of parapodial support rods truncate with distal lobes. Lateral organs about halfway from parapodia to margin. Cloacal opening sev- eral times farther from margin than pro- boscis opening. Proboscis unarmed.
Etymology. —Named for the usual host, Analcidometra armata (Pourtalés).
History. —McClendon (1907) included one myzostome from Analcidometra ar-* ~ mata in his type lot of Myzostoma cerrife- roidum, which otherwise occurred only on Crinometra brevipinna (Pourtalés). This type lot has since been pooled (USNM 5780), so the anomalous individual is no longer iden- tifiable, but one very small, damaged spec- imen 0.90 mm long and 0.86 mm wide might actually be M. armatae.
Material.—Five Bahamian samples as- sociated with Analcidometra armata: ho- lotype (USNM 118208) and 16 paratypes (USNM_ 118209) with over 34 crinoids (USNM E17939), 5 Sep 1973, 1 km N350°E of South Bight IV, Goldring Cay, Andros Ts) 2422329 NG 7362 WW, 1417 m:-one paratype (USNM 118210) with two cri-
794
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig’ bs
Myzostoma armatae. A, Holotype (USNM 118208), ventral view; B, Paratype from type locality
(from USNM 118209), ventral view; C, Large Jamaican paratype (from USNM 118216), dorsal view; D, small Jamaican paratype (from USNM 118220), ventral view; E, Aberrant paratype from type locality, ventral view, missing right parapodium 5 and lateral organ 4, numerous supernumerary cirri on abnormally short and rounded (due to injury?) rear margin of body; F, G, Parapodial hook and support rod, respectively, from a third paratype from type locality. Key: some parapodia numbered from the front in A and E; c, cloacal opening; lo, lateral organ; mz, marginal zone; 0, proboscis opening; p, penis; pr, proboscis; scale bars in mm.
noids (USNM E17987), 25 Aug 1973, Ma- rine Farm, Crooked Is., 22°50'N, 73°21'W, 40-52 m; one paratype (USNM 118211) with one crinoid (USNM E17749), 28 Aug L97 3.0? South). Long \ Cayo 22236sIN, 74°22.2'W, 15-18 m; six paratypes (USNM 118212) with five crinoids (USNM E178 18), 9 Sep 1973, 2 km S118°E of South Bight IV, Goldring Cay, Andros Is., 24°12.4’N, 77°34.9'W, 43 m; 16 paratypes (USNM 118213) with over 20 crinoids (USNM E17951), 9 Sep 1973, same collection as previous entry except 11-12 m.
Seven samples collected by D. B. Macur- da, Jr., at Discovery Bay, Jamaica, from Analcidometra sp. unless otherwise stated (presumably A. armata, since that is the only species currently recognized in the genus; Meyer et al. 1978): 24 paratypes (USNM
118214), W moat and knob in front of Dancing Lady Reef, 9 Jul 1974, 23-27 m; 5 paratypes (USNM 118215), LTS W of Dancing Lady Reef, and Pinnacle II, 10 Jul 1974, 21-27 m; 3 paratypes (USNM 118216), forereef escarpment, Dancing Lady Reef, 8 Jul 1974; 20 paratypes (USNM 118217) on A. armata, no detailed collec- tion data; 21 paratypes (USNM 118218), NW edge of forereef terrace, Lynton’s Mine, 7 Jul 1974, 21-24 m; 1 paratype (USNM 118219), SE side of Pinnacle I, front of Lyn- ton’s Mine, 6 Jul 1974, 24-27 m; 2 para- types (USNM 118220) on Davidaster dis- coidea (Carpenter), E slope and sand channel E of Lynton’s Mine, 9 Jul 1974, 12-17 m. Description. —Holotype 2.25 mm long, 1.61 mm wide (Fig. 1A). Bahamian para- types 0.90-—2.31 mm long, mean of 16 from
VOLUME 102, NUMBER 3
USNM 118213 1.51 mm, of 15 from USNM 118209 1.80 mm, average length : width 1.7— 1.8 but range 1.2—2.5 depending on con- traction or enrollment. Jamaican paratypes larger (0.73—3.06 mm long, mean of 17 from USNM 118217 2.28 mm), but with same body shape (average length : width 1.7, range 1.2—2.6). Bahamian specimens (Fig. 1A, B, E) uncolored or yellowish, lateral edges of body often downturned; nearly smooth, rather thick cuticle; marginal zone poorly distinguished. Jamaican specimens (Fig. 1C) mostly light brown, sometimes with darker dorsal patches, body usually flat or saddle- shaped, dorsal texture leathery or minutely pebbled; narrow but distinct, translucent marginal zone present.
Following idealized description applies in full to minority of examined specimens; variability described afterwards.
Body oval with rounded front and nar- rower, often truncate rear, usually with dis- tinct narrowing at level of cloacal opening or of ninth pair of marginal cirri. Funda- mentally 10 pairs of marginal cirri, first 8 pairs equally spaced around anterior and lateral margins to level of last parapodia, last 2 pairs progressively father apart, ninth just behind cloacal opening, tenth on rear corners (Fig. 1A—C). In small specimens tenth pair often at least twice as long and thick as other pairs (Fig. 1D), and first pair sometimes also long, but this distinction mostly lost in large specimens. Cirri usually equilaterally or acutely triangular, but lon- ger and thinner in a few specimens of all sizes, or, especially in Jamaican specimens, reduced to little more than marginal thick- enings.
Five pairs of parapodia forming oval in anterior two-thirds of ventral side, equally spaced and positioned about halfway from center of oval to anterior and lateral mar- gins, third pair closest to edge (Fig. 1A, B, D). Parapodia small, with conical, acirrate base and stubby, finger-like distal part; lat- ter extensible enough to overreach body margin, even in rear parapodia (only re- tracted ones illustrated). Parapodial hooks
795
moderately stout, nearly straight, tips evenly rounded and tapered, bending more than 90° (Fig. 1F). Support rods thinner and a little longer than hooks, manubrium ex- panded very little on rear side, front side truncate about 5 small, distal lobes (Fig. 1G). One or 2 replacement hooks present. Penes present as short, broad, cylindrical nozzles arising from lateral bases of third parapodia and sometimes reaching as far as body mar- gin, diameter greater than that of distal part of parapodium (Fig. 1A, B, E). Four pairs of lateral organs alternating with parapodia and forming arcs parallel to body margin about halfway between parapodia and mar- gin (Fig. 1A); lateral organs small, round or more usually radially oval, resembling short, cylindrical tubes when protruded. Proboscis opening ventral, about one-third of way from front margin to first parapodia; ex- tended proboscis cylindrical, twice as long as thick, lacking papillae (Fig. 1B). Cloacal opening at level of ring of lateral organs, much closer to last parapodia than to rear body margin.
Variability. —Body outline and marginal cirri subject to much variability, apparently due to injury from predators; lateral edges and more often rear end often showing ev1- dence of healing of large wounds and some- times missing parapodia (Fig. 1E). Rear often deeply notched on right or left side, sometimes entire postanal region missing. In a few cases, rear split lengthwise. Ten marginal cirri on each side of body in only 30% of 70 specimens from 4 lots, unilateral cirral count 4—21, but 8-12 86% of the time. Low cirral counts due to injury except in a few apparently undamaged Jamaican spec- imens simply lacking cirri towards rear. Commonly from one to two supernumerary cirri of usual form. If more cirri present, extra ones most often found in healed pos- terior areas, less often laterally (Fig. 1E); such cirri smaller than normal and generally appearing in closely spaced rows.
Remarks. —Of described Caribbean my- zostomes, only Myzostoma rotundum Graff, 1883, has an oval body with an eccentric
796
ring of parapodia (Graff 1884). The body form and size (1.7 X 1 mm), the small para- podia excluded from the rear part of the body, and the position of the proboscis opening match M. armatae. But the wide marginal zone of M. rotundum ('% of body diameter), and its much larger hemispher- ical lateral organs that are very close to the margin (in the marginal zone itself), are dif- ferent. So are the high number (22 along one side of the body), equal spacing, and rather filiform shape of its marginal cirri and the wider spacing of the parapodia to- wards the rear.
Except for two specimens found with Davidaster discoidea, Myzostoma armatae seems to be restricted to Analcidometra (presumably always A. armata) as a host, and this new species is currently known from the Bahamas and Jamaica at 12-52 m. The specimens from D. discoidea co-occurred with another undescribed species of My- zostoma.
Myzostoma attenuatum, new species Fig. 2
Diagnosis. — Body elongate, up to 4.5 mm long, with tapered, postanal, caudal region in undamaged specimens. No translucent marginal zone. Numerous short, irregularly sized, closely spaced marginal cirri back to level of cloacal opening, sparser in caudal region. Parapodia acirrate, usually confined to front half of body, a little closer to margin than midline. Parapodial hooks very thick, manubrium of support rods hatchet-shaped. Round lateral organs halfway from para- podia to margin, half parapodial diameter when protruded. Proboscis opening closer to first parapodia than margin, proboscis unarmed.
Etymology.—Named for the drawn out, attenuated caudal region of undamaged specimens.
Type material.—Unless otherwise speci- fied, all specimens loose in jars with sup-
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
posed hosts. Holotype (USNM 118221) with one Capillaster sentosa (Carpenter) (USNM 34844), Albatross stn. 5146, 5.7 km SE of E Sulade Is., near Siasi, Sulu Archipelago, Philippines, 5°46’40’N, 120°48'50’E, 44 m, 16 Feb 1908; 2 paratypes (USNM 118222) with 3 Comaster multifida (J. Miller) (USNM E34538), collected by D. L. Meyer, Lizard Is., Queensland, Australia; 3 para- types (USNM 118223) with 2 Colobometra perspinosa (Carpenter) (USNM E34765), collected by D. L. Meyer, E side of Ta Bui Is., Indonesia, 6-18 m, 29 Mar 1975; 10 paratypes (USNM 118224) with 2 Oxj- metra finschi (Hartlaub) (USNM E34861), collected by D. L. Meyer, Malaysia; 20 para- types (USNM 118225) with 2 O. finschi (USNM E34576), collected by D. L. Meyer, Singapore; 1 paratype (USNM_ 118226) found halfway out along arm of Pontio- metra andersoni Carpenter (USNM 35222), same collection as holotype; 3 paratypes (USNM 118227) with 1 P. andersoni (USNM E3130), New Harbour, Singapore, 1899; 9 paratypes (USNM 118228) with 2 Pontiometra sp. (USNM G2605), collected by D. L. Meyer, Lizard Is., Queensland, Australia, 1975; 1 paratype (USNM 118229) with 2 Decametra mylitta A. H. Clark (USNM E11628), International Indian Ocean Expedition, R/V Anton Bruun cr. 1, stn. 47B, northern Bay of Bengal, 19°50’N, 92°55’E, 22—30 m, 5 Apr 1963; 10 paratypes (NTMAS Ref. no. W326), collected from Cenometra cornuta A. H. Clark by R. Lock- yer, Cootamundra Shoal, Timor Sea, Northern Territory, Australia, stn. 2/43, 10°50’S, 129°13’E, 20 m, 10 May 1982; 1 paratype (author’s collection), host un- known, collected by A. Pietsch, Maldives, 1985.
Additional material. —3 partial speci- mens (USNM 118230) with Stephanometra oxyacantha (Hartlaub) (USNM E34854), collected by D. L. Meyer, Singapore; 1 spec- imen (USNM 118231) with 1 Comaster gracilis (Hartlaub) (USNM_ E35356), col- lected by D. L. Meyer, Lizard Is., Queens-
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797
G 01 GH
<a
Fig. 2. Myzostoma attenuatum. A, Holotype (USNM 118221), ventral view, some lateral organs not visible due to upturned body margin; B, Paratype, ventral view (personal collection); C, Unusually short paratype (from USNM 118228), ventral view; D, Posterolaterally injured paratype (from USNM 118227), ventral view; E, Paratype (from USNM 118225) showing abnormal development of caudal part of body, presumably due to injury, dorsal view; F, Longest paratype (from USNM 118223), body twisted, lateral organs not shown; G, H, Parapodial hook and support rod, respectively, from specimen in same lot as C. Key: as in Fig. 1, except w, lump apparently caused by endoparasitic worm; scale bars in mm.
land, Australia; 1 specimen (USNM 118232), host unknown, from International Indian Ocean Expedition, R/V Anton Bruun cr. 1, stn. 47B (details above); 15 specimens
(USNM 118233), host unknown, from In- ternational Indian Ocean Expedition, R/V Anton Bruun cr. 1, stn. 18A, Andaman Sea off Phuket, Thailand, 7°34’N, 98°00’E, 77
798
m, 21 Mar 1963; 1 damaged specimen (NIMAS W325), collected from Petaso- metra helianthoides A. H. Clark, same col- lection data as NIMAS W326 above except stn. 2/50.
Description. —Body usually elongate and posteriorly tapered (Fig. 2A, B, F), but sub- ject to injury-induced malformations (Fig. 2D, E). Front half either convex dorsally and concave ventrally with downturned margins or slightly convex dorsally and flat ventrally, with parallel, extended lateral margins. Dorsum smooth, fine-textured, sometimes with low, longitudinal ridge over proboscis and gut. Color usually dark brown, but poorly preserved specimens colorless or with dark dorsal speckling. No marginal zone. Holotype 2.50 mm long, 1.11 mm wide, with upturned sides (Fig. 2A), other undamaged specimens 0.88—4.46 mm long, length: width from 1.6—3.5 (all measure- ments excluding cirri). Marginal cirri very numerous and closely spaced anterior of level of cloacal opening (about 40 in holo- type, varying widely from about 25 to nearly 50 in other specimens), caudal part of body lined by smaller and more widely spaced cirri. Cirri bluntly triangular to digitiform with 3-fold difference in length and thick- ness in a single specimen. Alternating size pattern seen in many specimens (Fig. 2A). Caudal region usually symmetrical, but in many specimens secondarily shortened (Fig. 2C), or split longitudinally due to injury (predation?). Entire caudal region some- times lost or bizarrely asymmetrical (Fig. 2E).
In undamaged specimens, five pairs of parapodia arranged in two nearly parallel rows or shallow arcs on anterior half or one- -third of ventral side, slightly closer to margin than to midline (Fig. 2A—D, F). Obliquely conical base of parapodium lack- ing medial cirrus; distal part of parapodium a stubby, rounded process up to twice as long as thick. Parapodial hooks very stout, evenly tapered, tip bent at 90° (Fig. 2G). Support rods of same length but much thin-
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ner than hooks, manubrium hatchet-shaped, produced on both front and rear sides (Fig. 2H). Two replacement hooks. Pair of penes laterally on swollen basal parts of third parapodia (Fig. 2B, D), thicker than distal part of parapodium when protruded. Four pairs of lateral organs alternating with para- podia halfway between them and margin; when withdrawn, minute apertures sur- rounded by slightly raised annuli (Fig. 2B, C); expanded lateral organs with wide ap- erture and more prominent annulus, di- ameter about half that of distal part of para- podium (Fig. 2A, D). Proboscis opening ventral, closer to first parapodia than to an- terior margin, proboscis cylindrical, slightly tapering, unarmed (tip visible in Fig. 2A, F). Cloacal opening a little farther posterior of fifth parapodia than latter from fourth pair.
Longest specimen with oval dorsal hump presumably indicating presence of internal parasite (Fig. 2F). Injured specimens some- times lacking one or two parapodia on one or both sides.
Remarks.—Myzostoma attenuatum re- sembles two described species, M. dentatum Graff, 1884, and M. moebianum Graff, 1884. The first of these was described on the basis of one specimen, which has not been locat- ed, from the Torres Straits (Graff 1884). A second specimen of M. dentatum, which also has not been located, was found in a jar with seven species of Moluccan crinoids (Graff 1887). The elongate, oval body and super- numerary marginal cirri recall M. attenu- atum, but M. dentatum has a wide, distinct marginal zone, the proboscis opening closer to the front margin, the parapodia closer to the midline and not particularly confined to the front half of the body, and a rounded, untapered rear.
Myzostoma moebianum, a species de- scribed on the basis of collector’s notes and nearly useless microscopical preparations, is from an unknown host at Fouquet Is. southeast of Mauritius (Graff 1884). Its body outline and parapodial placement are sim-
VOLUME 102, NUMBER 3
ilar to small specimens of M. attenuatum; the rear is tapered and the parapodia, with very thin support rods, are restricted to the anterior two-thirds. It differs from the pres- ent species in having untapered parapodial hooks, somewhat differently shaped sup- port rod manubria, and most importantly a pair of lateral organs behind the fifth pair of parapodia. This last feature is unusual, otherwise occurring only in M. costatum sensu Boulenger (19 13a), and given the unor- thodox way the original description was prepared, it might be a mistake. If this could be proved, then M. attenuatum might be a synonym of M. moebianum.
Myzostoma attenuatum shows little host specificity. It infests 10 species of oligo- phreatan comatulids that belong to two su- perfamilies, three families (Comasteridae, Mariametridae, Colobometridae), and nine genera, one of the widest host ranges of the myzostomes I have investigated. Geograph- ically it ranges from the Maldives and the Bay of Bengal east through Singapore, Ma- laysia, and Indonesia to the southern Phil- ippines and northern Australia, and its re- corded depth range is 6—30 m. The holotype co-occurred with M. furcatum Graff, 1887, and M. longimanum (Jagersten, 1937); USNM 118222 with M. furcatum and M. ambiguum Graff, 1887; and USNM 118226 with M. sp. cf. triste Graff, 1877.
Myzostoma divisor, new species Figs. 3, 4
Diagnosis. — Body a round disc with pos- terior pair of cylindrical caudal processes up to 1.75 times longer than body diameter. Nine pairs of moderately long, equal cirri around disc, additional pair at ends of cau- dal processes. Parapodia two-thirds of way from center of disc to margin, basal part with pointed medial cirrus, distal part in two sections, extremely extensible. Para- podial hooks and support rods very long and slender, manubrium of latter digiti- form. Four pairs of relatively large, round
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lateral organs abutting margin. Proboscis opening terminal, proboscis unarmed. Clo- acal opening terminal, on papilla between caudal processes.
Etymology. — From Latin divisor, a divid- er, since larger animals in dorsal view re- semble a pair of dividers.
Type material.—Holotype (BMNH ZB 1980.460) and 12 paratypes (BMNH ZB 1980.461-472) from pinnules of Promacho- crinus kerguelensis Carpenter, Discovery sin. liGD2, 75°36:2.SPb78"35.5' W; 567m, 23-I-1936. 3 lots collected by W. H. Little- wood, Deep Freeze II, R/V Staten Island, hosts unknown: 1 paratype (USNM 118234), Weddell Sea, 77°32’S, 44°45’W, 284 m, 21-I-1957; 1 paratype (USNM 118235), stn. 24, 77°21'S, 44°30'W, 300 m, 20-I-1957; 5 paratypes including 2 early ju- veniles (USNM 118236), Weddell Sea, TS 20S. oT IW, S49 om, -17-1-1957. 1 paratype (USNM 118237), host unknown, Hero cr. 824, stn. 4-1, 65°13.60—13.67'S, 64°14.72-15.07'W, 49-58 m, 16-III-1982; 7 paratypes (USNM 118238), host un- known, Hero cr. 691, stn. 2A, 64°49.5’S, 63°47'W, 70 m, 1-II-1969; 107 intact, 272 damaged, and 26 early juvenile paratypes (USNM 118239), some used for SEM, host unknown, Herocr. 824, stn. 26-1, 64°14.30—- 13.80'S, 61°57.60-58.30'W, 238-285 m, 24- ITI-1982.
Additional material. —One specimen (BMNH ZB 1980.540) from pinnules of P. kerguelensis, Discovery stn. 1658, off Franklin Is., 76°9.6’S, 168°40’E, 520 m, 26- I-1936; one specimen (BMNH ZB 1980.547) from genital pinnules of female Notocrinus mortenseni John, Discovery stn. 187, NeuMayr Channel, Palmer Archipelago, 64°48'30"S, 63°31'30”W, 259-354 m, 18- III-1927; five specimens (BMNH ZB 1980.543-546) from pinnules of P. kergue- lensis, Discovery stn. 156, 53°51'00"S, 36° 21'30”W, 200-236 m, 20-I-1927; two spec- imens (BMNH ZB 1980.541-542) free on lower parts of arms of P. kerguelensis, Dis- covery stn. 42, off mouth of Cumberland
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i , x f a BS * rf . + i x
=
oo aS a = SS << = =
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Fig. 3. Myzostoma divisor. A, Holotype (BMNH ZB 1980.460), ventral view; B, Paratype from type locality (from BMNH ZB 1980.461-472), dorsal view; C, Parapodial hook and distal part of another from paratype (from USNM 118239); D, Parapodial support rod from same specimen as C. Key: as in Fig. 1, except cp, caudal
process; pc, parapodial cirrus; scale bars in mm.
Bay, South Georgia, 10.5 km N89°E of Ja- son Lt. to 6.7 km N39°E, 120-204 m, 1-IV- 1926; two specimens (USNM 118240), host unknown but one specimen from gorgonian washings, Hero cr. 824, stn. 14-1, 64°48.63— 48.20'S, 64°4.00’W, 70-150 m, 19-III-1982; one specimen (USNM 118241) from cri- noid washings, Hero cr. 824, stn. 26-1, 64°14.30-13.80'S, 61°57.60—58.30'W, 238- 285 m, 24-III-1982.
Description of adults. —Body a round, flat disc with pair of cylindrical caudal processes and nine pairs of marginal cirri. Specimens from type locality larger than others. Ho- lotype disc diameter 1.79 mm, right and left caudal processes (omitting terminal cirri) 2.44 and 3.12 mm long, respectively (Fig. 3A). Largest paratype with disc diameter 2.12 mm, longer caudal process 2.82 mm. Maximum ratio of caudal process length to disc diameter 1.75 in specimen 1.10 mm across; caudal processes generally, but with
many exceptions, relatively longer in larger individuals (Fig. 4; see ontogeny section be- low).
Color yellowish brown, no translucent marginal zone. Dorsum smooth or with lon- gitudinal swelling, and sometimes with ra- dial grooves between parapodial muscle masses (Fig. 3B). Nine pairs of evenly spaced, equally long marginal cirri, similar pair at ends of caudal processes, though these often broken off; in holotype marginal cirri about 0.14 mm long, terminal ones on cau- dal processes about 0.16 mm long. Caudal processes cylindrical or somewhat flattened dorsoventrally, diameter about one-fifth that of body disc (Fig. 3A, B).
Five pairs of parapodia equally spaced in arcs at least two-thirds of way from center of body disc to margin, members of first pair and especially fifth pair widely sepa- rated compared to spacing within arcs (Figs. 3A, 4C, D). Basal part of parapodium an
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Fig.4. Myzostoma divisor. A-D, Successively later stages in early, post-settlement ontogeny, scanning electron micrographs of paratypes (from USNM 118239). Key: as in Fig. 1 except mc, marginal cirrus; cp, caudal process; pc, parapodial cirrus; scale bars 0.2 mm.
obliquely truncate, radially inclined cylin- der with slender cirrus near apex of medial side (Figs. 3A, 4D). Long, slender, appar- ently bipartite distal part of parapodium projecting radially, either straight or me- dially curved, extremely extensible and ca- pable of doubling length. Parapodial hooks long, slender, weakly sigmoid, tips exhib- iting range of curvatures (Fig. 3C), appar- ently most broadly rounded in third pair (cf. Fig. 4A). Support rods same length, slightly thinner than hooks, manubrium a long, digitiform process (Fig. 3D). One or two replacement hooks. Penes present as small buttons at lateral bases of third para- podia. Four pairs of lateral organs alternat- ing with parapodia, outer edges nearly or actually abutting body margin, inner edges just inside outer edges of parapodial bases
(Figs. 3A, 4B). Lateral organs round, with same or greater diameter as distal parts of parapodia, low mounds with radially elon- gate, stellate apertures when retracted, round pads with depressed centers when protrud- ed. Proboscis opening on anterior margin, proboscis a short, unarmed cylinder almost as thick as caudal processes (Figs. 3B, 4D). Large cloacal papilla on rear margin of body between bases of caudal processes (Fig. 3A). Ontogeny. —Seven minute to small para- types from USNM 118239 were examined by SEM (Fig. 4) after critical point drying and sputter coating with carbon and gold. Earliest stage (Fig. 4A) with oval body 200 um long not counting proboscis, widest behind third parapodia, tapering more to- wards front than rear. No marginal cirri or caudal processes developed. Third para-
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podia largest, others smaller towards front and rear, last pair much smaller than first, no parapodial cirri present. Lateral organ apertures small, on sides of body. Appar- ently non-retractile proboscis 60-80 um long. Cloacal opening not seen (nor in any other SEM specimen).
Next stage (Fig. 4B) represented by spec- imen 430 um long (310 wm of main body plus extended proboscis); body 0.21 mm wide at level of third pair of lateral organs. Most marginal cirri present as blunt, ta- pered processes, last pair (caudal process rudiments) thicker than others. Articulation groove between basal and distal part of parapodium deeper and more nearly cir- cular than in preceding stage, parapodial cirri now clearly present. First pair of parapodia much smaller than other 4 pairs. Lateral organs visible as raised annuli around small pores.
In later stages (Fig. 4C, D) body round, marginal cirri and distal parts of parapodia elongate. Caudal processes first thickened into cones as long as marginal cirri (Fig. 4C); illustrated specimen the smallest of its type, wider but slightly shorter than specimen in Fig. 4B. Caudal processes becoming elon- gate (Fig. 4D) and proboscis retractable.
Remarks.—The only described species similar to M. divisor are M. bicaudatum Graff, 1883, and the species described under the name M. filicauda by Graff (1884). My- zostoma bicaudatum was collected in the Gulf of Mexico, and the unique specimen, which has been lost, was most fully de- scribed by Graff (1884). It has a round body 0.45 mm across with a pair of caudal pro- cesses. However, there are 10 pairs of mar- ginal cirri, not 9, the front and rear cirri are considerably longer than the lateral ones, the caudal processes lack a “terminal thread”’ (=terminal cirrus), and the proboscis open- ing is ventral; no parapodial cirri are re- ported, but in such a small animal they may have been overlooked.
Myzostoma filicauda was most fully de- scribed by Graff (1884) from specimens col-
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
lected by the Corvin off Sandkey; these spec- imens have also been lost. Graff (1883, 1884) has accidentally caused a nomencla- tural confusion between this species and M. filiferum Graff, 1884 (from the Torres Straits), because his 1883 original diagnosis of M. filicauda agrees with the 1884 descrip- tion only in terms of size and host, while the characterization of the caudal processes fits the 1884 original description of M. fili- ferum (terminal threads longer than body diameter). Since the type specimen of the latter species is also missing, it is now 1m- possible to resolve this apparent lapsus of Graff’s, so assume that Graff’s actual con- ceptions of the two species are those pre- sented in the Challenger Report (Graff 1884), from which the 1883 paper was ab- stracted. Myzostoma filicauda, as here understood, differs from M. divisor in hav- ing 10 pairs of marginal cirri as well as the pair on the caudal processes, and the lateral cirri are distinctly shorter than the anterior and posterior ones. The proboscis may have papillae, although the latter may be folds in the wall of the proboscis opening, to judge from the illustration. M. filicauda also has large, radially oval, lateral organs instead of round ones. Finally, it is hard to imagine that a single species of myzostome could range from Antarctica to the subtropical and tropical shallows of the Atlantic or Pacific.
Few studies of myzostome post-settle- ment ontogeny have been conducted. Ja- gersten (1940b) has given the most com- plete account, based on Myzostoma cirriferum Leuckart, 1836, and Kato (1952) studied a species identified as M. ambiguum Graff, 1887; both authors reviewed the scanty literature. The earliest observed stage in MM. divisor corresponds to Jagersten’s (1940b) fig. 4 of M. cirriferum and fig. 7 of M. alatum Graff, 1884, and Kato’s (1952) fig. 41 of M. ambiguum in having an oval body, all 5 pairs of parapodia developed, the proboscis unretracted, and no marginal cirri. There are no obvious identifying fea- tures at this stage, and the present minute
VOLUME 102, NUMBER 3
juveniles are identified as M. divisor on the basis of their association with hundreds of older, positively identifiable specimens. In M. cirriferum the first pair of parapodia is apparently the last to arise; in M. ambiguum the first and last pairs appear after the mid- dle three. The first pair is small compared to the middle three pairs in the present ju- veniles, but in the youngest ones the sixth pair is even smaller, suggesting that it may have appeared last, and the first pair second to last. The progressive envelopment of the proboscis (actually the anterior part of the body; Jagersten 1940b) by the larger pos- terior part of the body is well exhibited by M. divisor.
Until now, only three species of myzo- stomes have been reported from Antarctic waters, one each of Myzostoma (M. antarc- ticum Stummer-Traunfels, 1908), Cysti- myzostomum (C. cysticolum Graff, 1883), and an unnamed Asteromyzostomum (cf. Stummer-Traunfels 1908, Boulenger 1913b, Grygier 1988). Of these, only C. cysticolum is known from Promachocrinus kerguelen- sis, where it occupies soft cysts on the oral disc, and none are known from M™. divisor’s other identified host, Notocrinus morten- seni. Promachocrinus and Notocrinus be- long to different suborders of the Comatu- lida, so M. divisor is probably not at all host specific. Its currently known geographical and depth range includes the Ross Sea, Weddell Sea, waters northwest of the Ant- arctic Peninsula, and South Georgia, at 49-— 567 m.
Acknowledgments
I thank Mr. Alex Muir (BMNBH), Mr. R. Hanley (NTMAS), and Dr. L. Knapp (Smithsonian Oceanographic Sorting Cen- ter) for loans of specimens; Dr. K. Fauchald, Dr. D. L. Pawson, Ms. C. Ahearn, Ms. L. Ward, and Mr. T. Coffer (USNM) for as- sistance in amassing and documenting my- zostomes from the USNM crinoid collec- tion; Dr. M. Pettibone (USNM) for access
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to literature; and the SEM laboratory (USNM) for technical assistance. This work was supported by a Smithsonian Institution Postdoctoral Fellowship.
Literature Cited
Boulenger, C. L. 1913a. Report on the Myzostomida
collected by Mr. Cyril Crossland in the Red Sea
in 1905.— Proceedings of the Zoological Society
of London 1913:85-108, Pl. V-VII.
1913b. Myzostomida.—British Antarctic
(Terra Nova) Expedition 1910, Zoology 2:135-
140, Pl. 1.
Graff, L. 1877. Das Genus Myzostoma (F. S. Leuck- art). Verlag von Wilhelm Engelmann, Leipzig, vat + 82 pp., 11 Laf.
Graff, L. v. 1883. Verzeichniss der von den United
States Coast Survey steamers ‘Hassler’? und
“Blake” von 1867 bis 1879 gesammelten My-
zostomiden.— Bulletin of the Museum of Com-
parative Zoology 11(7):125-133.
. 1884. Report on the Myzostomida collected
during the voyage of H.M.S. Challenger during
the years 1873-76.—Challenger Reports, Zo-
ology 10:1-82, Pl. I-X VI.
. 1887. Report on the Myzostomida collected
during the voyage of H.M.S. Challenger during
the years 1873-76. Supplement.—Challenger
Reports, Zoology 20:1-16, Pl. I-IV.
Grygier, M. J. 1988. Unusual and mostly cysticolous crustacean, molluscan, and myzostomidan as- sociates of echinoderms. Pp. 775-784 in R. D. Burke, P. V. Mladenov, P. Lambert, and R. L. Parsley, eds., Echinoderm biology, Proceedings of the Sixth International Echinoderm Confer- ence, Victoria, 23-28 August 1987, A. A. Bal- kema, Rotterdam and Brookfield.
Jagersten,G. 1937. Myzostomiden von Prof. Dr. Six-
ten Bocks Expedition nach Japan und den Bo-
nin-Inseln 1914.—Arkiv for Zoologi 29A(17):
1-35, Taf. 1-2.
1940a. Neue und alte Myzostomum-Arten aus dem Zoologischen Museum Kopenhagen. —
Videnskabelige Meddelelser fra Dansk Natur-
historisk Forening i Kobenhavn 104:103-125,
|e lege
1940b. Zur Kenntnis der Morphologie, Ent- wicklung und Taxonomie der Myzostomida. —
Nova Acta Regiae Societatis Scientiarum Up-
saliensis (4)11(8):1-84, Taf. 1-7.
Kato, K. 1952. Onthe development of myzostome. — Science Reports of Saitama University, Series B (Biology and Earth Sciences) 1:1—16, Pl. I-III.
Leuckart, F.S. 1836. In Beziehung auf den Haarstern (Comatula) und Pentacrinus europaeus, so wie
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auf der Schmarotzerthier auf Comatula.—No- tizen aus dem Gebiete der Natur- und Heil- kunde, gesammelt und mitgetheilt von Dr. L. G. v. Froriep 50(9, no. 1087):129-131.
McClendon, J. F. 1907. New marine worms of the genus Myzostoma.— Proceedings of the United States National Museum 32:63-65.
Meyer, D. L., C. G. Messing, & D. B. Macurda, Jr. 1978. Zoogeography of tropical western Atlan- tic Crinoidea (Echinodermata).— Bulletin of Marine Science 28:412-441.
Stummer-Traunfels, R. R. v. 1908. Myzostomi-
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dae.— National Antarctic Expedition, Natural History 4:1-26, 1 Pl.
Department of Invertebrate Zoology, Na- tional Museum of Natural History, Smith- sonian Institution, Washington, D.C. 20560; (present address) Sesoko Marine Science Center, University of the Ryukyus, Sesoko, Motobu-cho, Okinawa 905-02, Japan.
PROC. BIOL. SOC. WASH. 102(3), 1989, pp. 805-807
SYSTEMATIC STATUS OF LEPIDOTEUTHIS, PHOLIDOTEUTHIS, AND TETRONYCHOTEUTHIS (CEPHALOPODA: OEGOPSIDA)
Clyde F. E. Roper and C. C. Lu
Abstract. — The status of the nomenclature and systematics at species, generic, and familial levels for Lepidoteuthis, Pholidoteuthis, and Tetronychoteuthis is reviewed and current knowledge consolidated based on examination of spec-
imens and analysis of literature.
The systematics and nomenclature of the squid genera Lepidoteuthis Joubin, 1895, Pholidoteuthis Adam, 1950, and Tetronych- oteuthis Pfeffer, 1900 have become so con- fused that it is difficult to discuss any of the species without raising doubts concerning the entities involved. During the course of a study of the comparative functional mor- phology of dermal structures in several species of oceanic squids (Roper & Lu 1989), we were able to make some conclusions about relationships in this group. We pre- sent here the results of our examination of numerous specimens and the literature in an effort to clarify the situation.
Tetronychoteuthis had been placed for “convenience” in the Lepidoteuthidae with Lepidoteuthis and Pholidoteuthis (Roper et al. 1969, Voss 1977) because of their com- mon character of dermal “‘scales,”’ but the actual relationships of these genera have re- mained obscure because of a lack of speci- mens to support research. Clarke (1980) and Clarke & Trueman (1988) separated these genera into the monotypic family Lepido- teuthidae (Lepidoteuthis grimaldii Joubin, 1895) and the family Pholidoteuthidae, containing Pholidoteuthis boschmai Adam, 1950, P. adami Voss, 1956, and Tetro- nychoteuthis massyae Pfeffer, 1912. The status of Tetronychoteuthis dussumieri (Or- bigny) sensu Pfeffer, 1900, in relation to Ony- choteuthis dussumieri Orbigny, 1839, needs clarification.
Our review suggests that the following sit- uations exist.
1) Onychoteuthis dussumieri (Orbigny, 1839 [in 1834-1848] (p. 335, Onychoteuthis pl. 13, figs. 1-6; type locality — Mauritius; type depository— Museum National d’His- toire naturelle, Paris?) is a species of Mo- roteuthis Verrill, 1881, based on the pres- ence of two rows of hooks on the tentacular clubs, the dermal structures that are larger and fewer than in Tetronychoteuthis mas- syae Pfeffer, 1912 (pp. 102-104, pl. 14, figs. 15-19), smooth sucker rings on the arm suckers, and an onychoteuthid shape of the gladius, fins and body. Therefore, this species belongs in the family Onychoteuthidae and bears the name Moroteuthis dussumieri (Or- bigny, 1839 [in 1834—1848]). Furthermore, future research may show it to be a senior synonym of a currently recognized species of Moroteuthis Verrill, 1881.
2) The specimen that Pfeffer (1900) re- ferred to Tetronychoteuthis dussumieri (Or- bigny, 1839) when he established the genus cannot be conspecific with Orbigny’s species because it has no hooks on the tentacular clubs and has a gladius very different from that of O. dussumieri Orbigny (see Pfeffer, 1912:98-102, pl. 13, figs. 1-3, pl. 14, figs. 10-14). Furthermore, it is so different from any onychoteuthid that it belongs to a dif- ferent genus and family as well. Pfeffer’s specimen, therefore, is a misidentification of the type species of his genus Tetronycho-
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teuthis. So, what is the type species of Tetro- nychoteuthis - the real dussumieri of Orbig- ny or the species that Pfeffer actually had in hand? The case must be referred to the International Commission on Zoological Nomenclature for a decision under Article 70 of the International Code. In such cases, the Commission usually makes a decision based on subsequent usage of the names, with present opinion and usage of active workers weighing heavily (pers. comm., F. M. Bayer). Toll’s (1982:247) placement of “T. dussumieri”’ of Pfeffer into the synon- ymy of Pholidoteuthis boschmai Adam, 1950 is premature from the standpoint of the Code (ICZN 1985).
Rees and Clarke (1963:853-854, fig. 1) recorded as 7. dussumieri (Orbigny) five specimens from the Northwest Atlantic Ocean. We point out for the record that this is a misidentification and that the specimen in the photograph is a Brachioteuthis sp. Presumably the other four specimens were Brachioteuthis as well.
3) Pholidoteuthis boschmai Adam, 1950 (pp. 1592-1598, pls. 1-3, figs. 1-6; type lo- cality— Flores Sea: type depository — Rijks- museum van Natuurlijke Historie, Leiden) was erected as the type species of a new genus Pholidoteuthis and new family, Phol- idoteuthidae. It bears ““hinged”’ suckers and no hooks on the tentacular club. The gladi- us, the dermal structures (Roper and Lu 1989) and the club structure of P. boschmai are similar to those of Pfeffer’s “7. dussu- mierl,” and Adam (1950), Clarke (1980: 129-138, pl. II, figs. 1, 2, text-fig. 94), and Toll (1982:247-252, pl. 28C) suggested that these two taxa are conspecific. We, however, feel that currently there is insufficient in- formation about Pfeffer’s species to verify this assertion. If examinations of Pfeffer’s specimen of “dussumieri’ and Adam’s type of boschmai proves them to be conspecific, the correct generic and specific names will have to be determined by a submission to the International Commission on Zoologi- cal Nomenclature (see 2 above).
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4) Pholidoteuthis adami Voss, 1956 (pp. 132-136, fig. 9; type locality — Gulf of Mex- ico; type depository — National Museum of Natural History, Washington) shows close familial affinity to Lepidoteuthis grimaldii Joubin, 1895 (pp. 1172-1173, 1 fig.; type locality— Azores Islands; type depository — V’Institut Oceanography, Monaco) in the structure of the dermal cushions (formerly called “‘scales’’; see Roper & Lu 1989), the gladius and the shape, conformity, and con- sistency of the mantle and fins. Therefore, we recommend that P. adami Voss, 1956 be placed in the family Lepidoteuthidae.
However, the correct generic designation cannot be determined until the status of P. boschmai, the type of the genus, is estab- lished. Nor do we know if adami and bosch- mai are, in fact, congeneric. Toll (1982:250- 251) demonstrated that the gladius of P. adami (and L. grimaldii) is so different from P. boschmai that a congeneric relationship of currently aligned Pholidoteuthis species appears untenable.
5) Tetronychoteuthis massyae Pfeffer, 1912 (pp. 102-104, pl. 14, figs. 15-19; type locality —48°N15°W (Atlantic); type deposi- tory —unknown). This species has to be re- tained in the genus Tetronychoteuthis until the nomenclatural status of “7. dussumieri” of Pfeffer (1900) is resolved (see 2 above) and the relationship between the two species is established. Several authors (Pfeffer 1912; Clarke 1966, 1980; Rancurel 1970) have suggested that 7. massyae is the juvenile of “T. dussumieri’”’ Pfeffer, 1900, but we be- lieve this is not so, especially if “7. dussu- mieri’” Pfeffer and P. boschmai Adam are conspecific. The specimen of “7. dussu- mieri’” Pfeffer, 1900 had a mantle length (ML) of 162 mm (sex and stage of maturity unknown). Clarke (1980) recorded nine specimens of P. boschmai from 240 to 580 mm ML, seven of which were mature or spent females. We have a male specimen of T. massyae of 105 ML (Museum of Victoria collections) that has developing testis, sper- matophoric apparatus, and Needham’s sac
VOLUME 102, NUMBER 3
(without spermatophores), so it must be ap- proaching maturity and maximum size. At present too few specimens, of sufficient size range, of the two taxa of Tetronychoteuthis have been examined to enable us to be cer- tain, but we believe that they are distinct species.
The ultimate solution of these problems lies in an examination of all extant type ma- terial, of voucher specimens, and of addi- tional specimens. In the meantime, this summary should provide a basis from which future work can proceed.
Acknowledgments
We thank Richard E. Young, University of Hawaii, Michael J. Sweeney and Fred- erick M. Bayer, National Museum of Nat- ural History for reviewing the manuscript.
Literature Cited
Adam, W. 1950. Un céphalopode nouveau: Pholi- doteuthis boschmai gen. et sp. nov.—Konink- lijke Nederlandse Akademie van Wetenschap- pen 53(10):1-8.
Clarke, M. R. 1966. A review of the systematics and
ecology of oceanic squids.—Advances in Ma-
rine Biology 4:91-—300.
. 1980. Cephalopoda in the diet of sperm whales
of the southern hemisphere and their bearing on
sperm whale biology.—Discovery Reports 37:
1-824.
Clarke, M.R., & E.R. Trueman. 1988. Introduction,
Pp. 1-10 Jn M. R. Clarke and E. R. Trueman,
eds., The Mollusca, volume 12, Academic Press,
San Diego, California, 355 pp.
1985. International Code of Zoological No- menclature. Ed. 3. Adopted by the 20th General Assembly of the International Union of Biolog- ical Sciences. University of California Press, Berkeley, California, 338 pp.
Joubin, L. 1895. Céphalopodes recueillis dans |’es- tomac d’un cachalot, capturé aux Iles Acores. — Comptes rendus des séances de l’Académie des Sciences, Paris 121:1172-1174.
ICZN.
807
Orbigny, A. d’. 1834-1848. Jn A. de Ferussac and A. @Orbigny. Histoire naturelle générale et par- ticuliere céphalopodes acétabuliféres vivants et fossiles. Paris, 96 pages + LVI + 361 pages, Atlas of 144 pls.
Pfeffer, G. 1900. Synopsis der oegopsiden Cephalo- poden.— Mitteilungen Naturhistorischen Mu- seum, Hamburg 17:147-198.
1912. Die Cephalopoden der Plankton-Ex- pedition.—Ergebnisse der Plankton-Expedition der Humboldt-Stiftung 2:1-815, Atlas of 48 pls. Rancurel, P. 1970. Les contenus stomacaux d’Alepi-
saurus ferox dans le sudouest Pacifique (Céph- alopodes).— Cahier ORSTOM, Séries Océano- graphie 8(4):5-87.
Rees, E. I. S., & M. R. Clarke. 1963. First records of Tetronychoteuthis dussumieri (d’Orbigny) (Cephalopoda; Onychoteuthidae) from the Northwest Atlantic.—Journal of the Fisheries Research Board of Canada 20(3):853.
Roper, CC. FYE. & €.'C. Lu. 1989. * Comparative
morphology and function of dermal structures
in oceanic squids (Cephalopoda). —Smithsoni- an Contributions to Zoology (in press).
, R. E. Young, & G. L. Voss. 1969. An illus-
trated key to the families of the Order Teu-
thoidea (Cephalopoda).—Smithsonian Contri-
butions to Zoology 13:1-32.
Toll, R. B. 1982. The comparative morphology of the gladius in the Order Teuthoidea (Mollusca: Cephalopoda) in relation to systematics and phylogeny. PhD. Dissertation, University of Miami, 390 pp.
Voss, G. L. 1956. A review of the cephalopods of the
Gulf of Mexico.— Bulletin of Marine Science of
the Gulf and Caribbean 6(2):85-178.
1977. Classification of Recent cephalo- pods.—Symposia of the Zoological Society of
London 38:575-579.
(CFER) Department of Invertebrate Zo- ology—Molluscs, National Museum of Nat- ural History, Smithsonian Institution, Washington, D.C., 20560 U.S.A.; (CCL) Department of Invertebrate Zoology, Mu- seum of Victoria, Melbourne 3000, Victo- ria, Australia.
PROC. BIOL. SOC. WASH. 102(3), 1989, pp. 808-811
RHYSSOPLAX BALIENSIS, A NEW SPECIES OF CHITON FROM INDONESIA (MOLLUSCA: POLYPLACOPHORA: CHITONIDAE)
Robert C. Bullock
Abstract. — Rhyssoplax baliensis is described from Bali, Indonesia. It differs from R. burmana (Pilsbry, 1893) by its smooth jugum, central mucro on valve VIII, and slightly convex valve I; from R. densilirata (Pilsbry, 1893) by having a banded girdle and by its narrower longitudinal ribs which bend medially and are more nodulose and widely spaced; and from R. vauclusensis (Hedley & Hull, 1909) by its smooth jugum, more numerous and smoother ribs on the lateral triangle, and smoother, less inflated girdle scales.
A single example of a large chiton from Bali, Indonesia, in the collection of the Aus- tralian Museum, Sydney, represents an un- described species of Rhyssoplax. While the study of many chiton species is made easier by the examination of numerous examples, I do not hesitate to describe the present species due to the specimen’s distinctive- ness, size, and excellent condition which al- lows a detailed comparison with related Rhyssoplax species.
Abbreviations used in the text: AMS, Australian Museum, Sydney; ANSP, Acad- emy of Natural Sciences of Philadelphia; BMNH, British Museum (Natural History), London; DMNH, Delaware Museum of Natural History, Greenville; MCZ, Mu- seum of Comparative Zoology, Harvard University, Cambridge; and ZMK, Univ- ersitetets Zoologiske Museum, Copenha- gen.
Class Polyplacophora Gray, 1821 Family Chitonidae Rafinesque, 1815 Genus Rhyssoplax Thiele, 1893
Use of Rhyssoplax at the generic level is not accepted by all malacologists. Van Belle (1978, 1983) and Kaas & Van Belle (1980) treated the group as a subgenus of Chiton Linnaeus, 1758. Morphological features of
the valves of both genera exhibit great plas- ticity and often are unreliable as taxonomic characters at the generic level. However, substantial radular differences exist between Chiton, which is primarily a New World group, and Rhyssoplax, which is represent- ed abundantly in the Indo-Pacific region (Bullock 1988a, b).
Rhyssoplax baliensis, new species Figs. 1-4, 8-11
Holotype. — Australian Museum, Sydney, C.60874, collected by T. Dranga.
Type locality.—Bali Island, Indonesia. Depth not recorded.
Description.— Animal moderately large, 47 mm in length, 24 mm in width. Valves subcarinate, angle about 115°. Valve I slightly convex; postmucronal slope of valve VIII concave anteriorly, convex posteriorly. Mucro somewhat blunt, central on valve VIII. Jugal region smooth; central areas with about 23 thin, longitudinal ribs which are more numerous and occasionally joined to- ward jugum. Lateral triangle raised, with seven or eight faintly nodular, radiating ribs; nodules more pronounced along posterior margin of lateral triangle. Terminal areas with numerous nodular, sometimes bifur- cate radial ribs; anterior valve with about
VOLUME 102, NUMBER 3
43 ribs; posterior valve with about 33 ribs. Valve color very light cream orange; jugum pinkish with reddish brown speckles; seven small, dark brown splotches on longitudinal ribs along anterior edge of lateral triangle, additional splotches along posterior margin; longitudinal and radial ribs speckled with reddish brown. Girdle yellowish white with several splotches or bands of dull green or brown. Interior of valves white with fleck of dull reddish brown on each side of pos- terior slope of callus near mucro.
Tegmentum: Ventral layer of suprateg- mentum laterally with wedgelike, nearly transparent shelf. Subtegmentum not de- veloped toward jugum, present laterally as thin layer of small canals (Fig. 8).
Esthete pores: Megalopores, each with surrounding micropores, scattered across central area; round to slightly ovate megalo- pores 1.6 to 2.6 times as large as round to ovate micropores (Fig. 9).
Articulamentum: Central depression of intermediate valves with numerous trans- verse slits in jugal tract. Primary slit-ray consisting of series of very small holes. Sec- ondary slit-ray present laterally. Insertion teeth finely grooved, not deeply pectinate; pectination of intermediate and posterior valves proceeding anteriorly to lateral mar- gins of apophyses. Slit formula 8/1/15.
Radula: [not available].
Girdle elements: Scales moderately large, roundly triangular, moderately inflated. Ex- tensive ventrolateral reticular sculpture mostly hidden by overlapping scales. Cen- tral area with 15-16 thin, moderately pro- nounced ribs. Apical region with pustules which may be smaller and linearly arranged distally and at sides. Apical shelf barely ev- ident (Figs. 10, 11). Ventral scales closely packed, rectangular, length 104-120 um, width 23-28 wm. Marginal spicules oblong, bluntly pointed distally, rounded proxi- mally, length 119-135 um, width 35—43 um.
Remarks. —Morphological features of the valves and girdle scales indicate that Rhys- soplax baliensis is related to R. vauclusensis
809
e {they =: Y sy ~ SRE AE SY mS % YL AO NGS eee Y Ae Cees WS Se ¢ We ANS Aes otk ~ Begrate RAN #3 c Py, 5 5$ ry Sic SPL eee SAVE He MA = ip pA Se xa tS, 4 “ABN “7 ¢ * Sa re S >y Z 4. ~ a ee ae 1 c 2 > - - < = z = oy “Zz: c > S ets i
Fig. 1. Rhyssoplax baliensis, new species, holotype, AMS C.60874: Top, Anterior valve (I); Middle, Inter- mediate valve (IV); Lower, Posterior valve (VIII). Scale bar = 0.5 cm.
(Hedley & Hull, 1909) from southern Queensland and New South Wales, Austra- lia (Iredale & Hull 1926); R. densilirata (Pilsbry, 1893), which occurs from the Phil- ippine Islands [Ang (1967) as Ischnochiton (Lepidozona) luzonicus (Sowerby, 1841); and specimens in DMNH and ZMK] south to the Admiralty Islands (BMNH:; see Fig. 6); and R. burmana (Pilsbry, 1893) from Waltair, India (BMNH; see Fig. 5), east to Burma. Rhyssoplax baliensis is differentiat- ed from the sympatric R. densilirata by: (1) the narrower longitudinal ribs of the central area which bend medially, not laterally; (2) the ribs of the central area and lateral tri- angle which are more nodulose and widely spaced; (3) the fewer radial ribs on the end valves (about 43 on valve I and about 33 on valve VIII compared with up to 69 on valve I and up to 57 on valve VIII of R. densilirata), and (4) the banded girdle. The
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
810
te oe
a Ee
VOLUME 102, NUMBER 3
girdle of examples of R. densilirata exam- ined was never banded.
Rhyssoplax vauclusensis (Fig. 7) appears to be the closest known relative of R. bal- iensis. The former differs by having longi- tudinal ribs on the jugum, fewer, smoother radial ribs on the lateral triangle, and smoother, less inflated girdle scales with a cluster of elongate nodules distally on the apex.
Rhyssoplax burmana, which has conspic- uous, narrow ribs on the central areas, dif- fers from R. baliensis by its smaller size, ribbed jugum, slightly posteriorly acentric mucro on the posterior valve, fewer (4-6) ribs on the lateral triangle, and prominently concave anterior valve and postmucronal slope of the posterior valve.
Chiton vangoethemi Leloup, 1981, a re- cently described species from Madang Province, Papua New Guinea, differs from R. baliensis in its coloration, smooth jugal and central areas, and evenly rounded pos- terior outline of valve VIII. The smooth insertion teeth described by Leloup indicate that this species belongs in the Ischnochi- tonidae, not the Chitonidae.
Acknowledgments
I am grateful to W. Ponder (AMS), for the opportunity to study the sample of Rhyssoplax baliensis. I thank R. T. Abbott and R. Jensen (formerly DMNH), K. Boss (MCZ), G. Davis and R. Robertson (ANSP), and J. Taylor, J. Peake, and K. Way (BMNH) for the loan of comparative material. The scanning electron microscopy was done in the Department of Zoology and at the Grad-
_
811
uate School of Oceanography, University of Rhode Island. The drawings of valves were provided by D. DeCarlo.
Literature Cited
Ang, E. Z. 1967. Loricates of the Philippines. — Nat- ural and Applied Science Bulletin 20(4):383- 464, 21 pls.
Bullock, R. C. 1988a. Notes on some Rhyssoplax
from the Pacific Ocean (Mollusca: Polyplacoph-
ora: Chitonidae).— Proceedings of the Biological
Society of Washington 101(3):682-692.
1988b. The genus Chiton in the New World
(Polyplacophora: Chitonidae).— Veliger 31(3/4):
141-191, 144 figs.
Hedley, C., & A. F. B. Hull. 1909. Descriptions of new and notes on other Australian Polyplacoph- ora.— Records of the Australian Museum 7:260- 266, pls. 73, 74.
Iredale, T., & A. F. B. Hull. 1926. A monograph of the Australian loricates. VI.— Australian Zool- ogist 4:164-185, pls. 18-20.
Kaas, P., & R.A. Van Belle. 1980. Catalogue of living chitons. W. Backhuys: Rotterdam. 144 pp.
Leloup, E. 1981. Chitons de Papua New Guinea.— Bulletin du Institut Royal Sciences Naturelles de Belgique 53(15):1-4, 2 figs.
Pilsbry,H. A. 1892-1894. Polyplacophora.— Manual of Conchology 14:1—128 [1892], 129-350 [1893]; 15:1-133 [1894].
Van Belle, R. A. 1978. Sur la classification des Poly-
placophora: V. Classification systématique des
Chitonidae (Neoloricata: Chitonina).—Infor-
mations de la Société Belge de Malacologie, Se-
rie 6, No. 1, pp. 19-28, pl. 8.
. 1983. The systematic classification of the chi-
tons (Mollusca: Polyplacophora).—Informa-
tions de la Société Belge de Malacologie, Serie
11, Nos. 1-3, pp. 1-178.
Department of Zoology, University of Rhode Island, Kingston, Rhode Island 02881.
Figs. 2-7. Rhyssoplax baliensis new species, R. densilirata (Pilsbry), R. burmana (Pilsbry), and R. vauclusensis (Hedley & Hull): 2-4, Anterior, intermediate, and posterior valves, respectively, of R. baliensis, holotype, AMS C.60874, width valve IV, 17.5 mm; 5, R. densilirata, Damma Is., Admiralty Is., BMNH 1899.4.12.3, 45 mm (slightly curled); 6, R. burmana, Waltair, India, BMNH 1952.11.19.62, 23 mm; 7, R. vauclusensis, Shellharbour,
New South Wales, Australia, MCZ 204376, 27 mm. Figs. 8-11.
Scanning electron micrographs of valve and girdle scale morphology of Rhyssoplax baliensis new
species: 8, Anterior tegmental innervation, <x 70; 9, Esthete pores of central area, x 280; 10, Dorsal surface of girdle scale, x 137; 11, Ventrolateral reticular sculpture of girdle scale, x 665.
812 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
ANNOUNCEMENT
Copies of a “List of parasitic Copepoda in the collection of the Atlantic Reference Centre, St. Andrews, New Brunswick, Canada,” are now available for distribution to interested individuals and institutions. The collection comprises lots representing 62 species of parasitic copepods from 13 families, ranging from commonly encoun- tered to extremely rare and paratype specimens. The list includes data on hosts,
localities, site of infection and number of specimens of each species. Copies are available by contacting:
W. E. Hogans
Systematics Laboratory
Huntsman Marine Science Centre
St. Andrews, New Brunswick, EOG 2X0 CANADA
VOLUME 102, NUMBER 3 813
APPLICATIONS PUBLISHED IN THE BULLETIN OF ZOOLOGICAL NOMENCLATURE
The following applications were published on 29 March 1989 in Vol. 46, Part 1 ofthe Bulletin of Zoological Nomenclature. Comment or advice on these applications is invited for publication in the Bu//etin and should be sent to the Executive Secretary, 1.C.Z.N., % British Museum (Natural History), Cromwell Road, London SW7 S5BD, U.K.
Case
2668 Drepanites Mojsisovics, 1893 and Hyphoplites Spath, 1922 (Mollusca, Ceph- alopoda): proposed conservation.
2452 Aphrodita imbricata Linnaeus, 1767 (currently Harmothoe imbricata) and Aphrodita minuta Fabricius, 1780 (currently Pholoe minuta) (Annelida, Polychaeta): proposed conservation of the specific names.
2603 GRYLLACRIDOIDEA Stal, 1874 (Insecta, Orthoptera): proposed prece- dence over STENOPELMATOIDEA Burmeister, 1838.
2646 Ptochus Schonherr, 1826 (Insecta, Coleoptera): proposed conservation by confirmation of Marshall’s (1916) designation of Ptochus porcellus Boheman in Schonherr, 1834 as the type species.
2680 Euribiajaceana Hering, 1935 (currently Urophora jaceana; Insecta, Diptera): proposed precedence over Euribia conyzae Hering, 1933.
2674 Monograptus exiguus (Graptolithina): proposed conservation of accepted usage by the citation of Lapworth (1876) as author.
2681 Heliastes ovalis F. Steindachner, 1900 (currently Chromis ovalis; Osteich- thyes, Perciformes): proposed conservation of the specific name.
2527 Heteronota pelagica Girard, 1857 (currently Gymnodactylus, Crytodactylus or Nactus pelagicus; Reptilia, Sauria): proposed conservation of the specific name.
814 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
OPINIONS PUBLISHED IN THE BULLETIN OF ZOOLOGICAL NOMENCLATURE
The following Opinions were published on 29 March 1989 in Vol. 46, Part 3 of the Bulletin of Zoological Nomenclature.
Opinion No.
1518 MHarpa articularis Lamarck, 1822 (Mollusca, Gastropoda): specific name con- served.
1519 Ammonites neubergicus Hauer, 1858 (Cephalopoda, Ammonoidea): to be given precedence over Ammonites chrishna Forbes, 1846.
1520 Chagrinichnites brooksi Feldmann, Osgood, Szmuc & Meinke, 1978 and Chagrinichnites osboodi Hannibal & Feldmann, 1983 (Trace fossil; arthropod): conserved.
1521 Eriophyes von Siebold, 1851 and Phytoptus Dujardin, 1851 (Arachnida, Acarina): Phytoptus pyri Pagenstecher, 1857 and Phytoptus avellanae Nalepa, 1889 designated as the respective type species.
1522 Callianidea H. Milne Edwards, 1837 (Crustacea, Decapoda): conserved.
1523 Corisa germari Fieber, 1848 (currently Arctocorisa germari; Insecta, He- miptera): neotype designated.
1524 Corisa distincta Fieber, 1848 (currently Sigara (Subsigara) distincta; Insecta, Hemiptera): specific name conserved.
1525 Phymatodes Mulsant, 1839 and Phymatestes Pascoe, 1867 (Insecta, Coleop- tera): conserved.
1526 Nanophyes Schoenherr, 1838 (Insecta, Coleoptera): conserved.
1527 Polyommatus emolus Godart, [1824] (currently Anthene emolus; Insecta, Lepidoptera): specific name conserved.
1528 Pyralis nigricana Fabricius, 1794 (currently in Cydia or Laspeyresia; Insecta, Lepidoptera): specific name conserved.
1529 Ceutorhynchus Germar, 1824, Rhinoncus Schoenherr, 1825 and Curculio assimilis Paykull, 1792 (Insecta, Diptera): conserved, and Curculio as- similis Paykull, 1792 and Curculio pericarpius Linnaeus, 1758 desig- nated as the type species of Ceutorhynchus and Rhinoncus respectively.
Coeloides Wesmael, 1838 (Insecta, Hymenoptera): Coeloides scolyticida Wesmael, 1838 designated as the type species.
Disophrys Foerster, 1862 (Insecta, Hymenoptera): Agathis caesa Klug, 1835 designated as the type species.
Siphonosoma vastum Selenka, De Man & Biilow, 1884, Phascolosoma steph- ensoni Stephen, 1942, Phascolosoma scolops Selenka, De Man & Bu- low, 1884 and Phascolosoma pacificum Keferstein, 1866 (Sipuncula): specific names conserved.
Holothuria arenicola Semper, 1868 (Echinodermata, Holothuroidea): specific name conserved.
Sternotherus Gray, 1825 and Pelusios Wagler, 1830 (Reptilia, Testudines): conserved.
Halianassa studeri von Meyer, 1838 (Mammalia, Sirenia): neotype desig- nated; and Halitherium Kaup, 1838 (Mammalia, Sirenia): Pugmeodon schinzii Kaup, 1838 designated as the type species.
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CONTENTS
New species and records of birds (Aves: Megapodiidae, Columbidae) from an archeological site on Lifuka, Tonga David W. Steadman Geographic variation in the Yellow-rumped Tanager (Aves: Thraupinae) Robert W. Storer Two overlooked holotypes of the Hawaiian flycatcher Chasiempis described by Leonhard Steyneger (Aves: Myiagrinae) Storrs L. Olson A new lizard of the genus Lepidodactylus (Reptilia: Gekkonidae) from Batan Island, Philippines Hidetoshi Ota and Ronald I. Crombie A new species of Eupsophus (Amphibia: Anura: Leptodactylidae) from southern Chile J. Ramon Formas A redescription of Pseudorhombus megalops, with comments on Cephalopsetta ventrocellata (Osteichthyes: Pleuronectiformes: Paralichthyidae) Dannie A. Hensley and Kunio Amaoka Paraspadella anops, new species, from Sagittarius Cave, Grand Bahama Island, the second troglobitic chaetognath Thomas E. Bowman and Robert Bieri A review of the beach flies of the Caribbean and Gulf of Mexico (Diptera: Canacidae) Wayne N. Mathis 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 Cancer johngarthi, n. sp. and Cancer porteri (Bell) (Crustacea, Decapoda): comparisons and
hypothesis Alberto Carvacho Stenorhynchus yangi, a new western Atlantic species of Arrow Crab (Crustacea, Brachyura, Majidae) and a redescription of S. seticornis (Herbst, 1788) Gary D. Goeke
Hobbseus yalobushensis, a new crawfish from central Mississippi (Decapoda: Cambaridae) : J. F. Fitzpatrick, Jr., and Craig A. Busack Hippolyte zostericola (Crustacea: Decapoda) in the eastern Pacific Mary K. Wicksten Chaceon ramosae, a new deep-water crab from Brazil (Crustacea: Decapoda: Geryonidae) Raymond B. Manning, Marcos Siqueira Taveres, and Elaine Figueiredo Albuquerque 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 Sanquerus, a replacement name for Posidon Herklots, 1851 (Crustacea, Decapoda, Portunidae) Raymond B. Manning Recitification of Halirages regis and H. huxleyanus (Crustacea: Amphipoda), from marine
Antarctica, with description of a new genus, Austroregia J. L. Barnard Two new species of wood-boring Limnoria (Crustacea: Isopoda) from New Zealand, L. hicksi and L. reniculus Marilyn Schotte
Some aspects of the biology of Rhopalophthalmus tatersallae Pillai, 1961 (Crustacea, Mysi- dacea) and extension of range into the Khor Al Sabiya, Kuwait (Arabian Gulf) Stephen A. Grabe Pycnogonida of the Western Pacific Islands VI. Sericosura cochleifovea, a new hydrothermal vent species from the Marianas Back-Arc Basin C. Allan Child 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 The Second Annual Riser Lecture: Eclecticism and the study of Polychaetes Kristian Fauchald A new species of Euchone Polychaeta: Sabellidae) from the northwest Atlantic with comments
on ontogenetic variability R. Eugene Ruff and Betsy Brown Paralvinella hessleri, new species of Alvinellidae (Polychaeta) from the Mariana Back-Arc Basin hydrothermal vents Daniel Desbruyéres and Lucien Laubier
A new species of Odontosyllis (Polychaeta: Syllidae) from Twin Cays, Belize David E. Russell Orbiniidae (Annelida: Polychaeta) from mangrove root-mats in Belize, with a revision of protoariciin genera Vivianne Solis-Weiss and Kristian Fauchald Three new species of Myzostoma (Myzostomida) Mark J. Grygier Systematic status of Lepidoteuthis, Pholidoteuthis, and Tetronychoteuthis (Cephalopoda: Oegopsida) Clyde F. E. Roper and C. C. Lu Rhyssoplax baliensis, a new species of chiton from Indonesia (Mollusca: Polyplacophora: Chi- tonidae) Robert C. Bullock International Commission on Zoological Nomenclature: Applications and Opinions
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PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY a
VOLUME 102 NUMBER 4
19 DECEMBER 1989 ©
ISSN 0006-324X
THE BIOLOGICAL SOCIETY OF WASHINGTON
1988-1989 Officers President: Kristian Fauchald Secretary: G. David Johnson President-elect: Leslie W. Knapp Treasurer: Don E. Wilson Elected Council
Gary R. Graves Meredith L. Jones
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Custodian of Publications: Austin B. Williams
PROCEEDINGS
Editor: C. Brian Robbins
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Classical Languages: George C. Steyskal Invertebrates: Stephen D. Cairns Frank D. Ferrari
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PROC. BIOL. SOC. WASH. 102(4), 1989, pp. 815-825
MARIANACTIS BYTHIOS, A NEW GENUS AND SPECIES OF ACTINOSTOLID SEA ANEMONE (COELENTERATA: ACTINIARIA) FROM THE
MARIANA VENTS
Daphne G. Fautin and Robert R. Hessler
Abstract.— We describe a new genus and species of sea anemone from the vicinity of hydrothermal vents in the Mariana back-arc basin, western North Pacific. Belonging to family Actinostolidae, Marianactis bythios is one of the most conspicuous animals around low temperature vents, at a depth of about 3600 m. Its mesenteries are not arrayed according to the Actinostola rule, it lacks basal tentacular thickenings, it has six pairs of complete mesenteries, and all its stronger mesenteries (members of the first three cycles) are gametogenic. It is further distinguished by having microbasic amastigophores in its tentacles. The final portion of Carlgren’s (1949) key to the Actinostolidae is revised to accommodate Marianactis and other recent taxonomic changes in the family.
In April and May 1987, scientists discov- ered deep-sea hydrothermal vents and an associated faunal community in the Mar- iana back-arc basin, NNW of Guam, at a depth of about 3600 m. Far from the pre- viously studied East Pacific Rise, these vents are unconnected to it by any intermediate spreading center.
Not surprisingly, much of the Mariana vent fauna is new (Hessler et al. 1988), in- cluding the relatively large sea anemone we describe here. One of the most conspicuous animals around low temperature vents, this member of family Actinostolidae belongs to a new genus and species, Marianactis by- thios (Fig. 1). Actinians found on the Ga- lapagos Spreading Center (Hessler & Smith- ey 1983) were too poorly preserved for detailed taxonomic study. The first species of actinian described from deep sea vents, Cyananthea hydrothermala Doumenc & Van-Praet, 1988, was collected on the French Biocyatherm 1 expedition around vents at 11°N in the Pacific, at 2000 m depth.
We detected no specializations to this un- usual habitat in M. bythios, an anatomically unremarkable member of a predominantly
deep water family. This ordinariness con- trasts with many new species of other groups from vent habitats. Marianactis is distin- guished by a suite of characters that indi- vidually occur in other of the approximately 20 genera comprising the Actinostolidae, as well as by microbasic amastigophores in its tentacles.
Materials and Methods
Field work relied on the submarine A/vin. Photographs were taken with a hand-held camera through the view ports, and with a Photosea M2000 stereocamera held by one of the mechanical arms. Video recordings were made with an Osprey video camera mounted on the same arm.
Eight specimens were collected using A/- vin’s mechanical arm. Placed in an insulated container for transport to the surface, they were not exposed to temperatures higher than 10°C. Once on the surface, seven spec- imens were preserved in either 10% for- malin or alcohol; one was initially frozen, and later formalin-preserved.
Paraffin sections 8 um thick were stained with hematoxylin and eosin. Cnidae mea-
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surements were on undischarged capsules in squash preparations. In the section ““Dis- tribution and size of cnidae,”’ “‘n’’ refers to the number of capsules measured, and ““N”’ is the proportion of animals examined in which that type of cnida was present. A measurement in parentheses was from a sin- gle capsule falling considerably outside the range of the others. Not all tissues of each specimen were studied. Microscopy, in- cluding photomicrography, was with a Reichert Ultrastar equipped with Nomarski interference contrast optics, and an auto- matic exposure camera.
Marianactis, new genus
Definition. — Actinostolidae with well de- veloped pedal disc. Column smooth; di- ameter about equal to height; mesoglea firm. Sphincter moderately strong; tentacles can be completely covered in retraction. Ten- tacles of uniform thickness entire length; outer much shorter than inner; arrayed in several cycles on marginal half of oral disc. Longitudinal tentacle and oral disc circular musculature ectodermal; that of tentacles equally well developed on all sides. Micro- basic amastigophores in tentacles. Fewer tentacles than mesenteries. Mesenteries not arrayed according to Actinostola rule; six pairs complete; all stronger ones fertile; two symmetrically arrayed siphonoglyphs at- tach to directive mesenteries; retractor mus- cles diffuse, parietobasilar muscles present. Cnidom: spirocysts, basitrichs, microbasic p-mastigophores, microbasic amastigo- phores.
Type species. —Marianactis bythios, new species.
Etymology and gender.—The name Mar- lanactis 1s a composite of ““Mariana,”’ geo- graphical locality of the first known occur- rence of this taxon, and “‘actis,’’ Greek literally for ray or beam, a term that is ap- plied to many taxa of sea anemones. The gender of “‘actis,”” and hence of Marianctis, is feminine.
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Differential diagnosis. —Most members of Actinostolidae inhabit deep water and are therefore known primarily from preserved specimens. The genus Marianactis resem- bles many other genera of this family in being colorless (in preservation), in having relatively thick mesoglea, and in possessing tentacles that are wrinkled in preservation. It has the following combination of diag- nostic characters: 1) mesenteries not ar- rayed according to the Actinostola rule, 2) no basal tentacular thickenings, 3) six pairs of complete mesenteries, and 4) all the stronger mesenteries (that is, members of the first three cycles) gametogenic. Marian- actis is distinguished from all other actino- stolids by its tentacular microbasic amas- tigophores.
Marianactis most resembles Anthosactis Danielssen, 1890, which has microbasic b-mastigophores in its stinging batteries, and unequally developed longitudinal tentacle musculature. It is also quite similar to Tea- lidium Hertwig, 1882, which is papillose. The definition of genus Jsoparactis greatly re- sembles that of Marianactis. Carlgren (1949) included it among the Actinostolidae, using the definition of Stephenson (1920), who created it for Paractis ferax Stuckey, 1909. Parry (1952) demonstrated that Stuckey’s species is actually an acontiate anemone be- longing to family Bathyphelliidae.
All taxa were diagnosed initially with Carlgren (1949) and subsequently from the original literature. In Carlgren’s (1949:77— 78) key to the Actinostolidae, Marianactis falls under option II (““mesenteries not ar- rayed according to the Actinostola-rule’’), B (“all or all stronger mesenteries fertile’’). Be- yond that, the key requires revision not only to accommodate Marianactis. Carlgren’s descriptions of two additional genera of ac- tinostolids were published posthumously — Hadalanthus Carlgren, 1956, and Cnidan- thea Carlgren, 1959. In addition to Jsopar- actis, Actinoscyphia must be eliminated, be- cause Riemann-Ziurneck (1978) restored it
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to its own family. Also, Carlgren (1949) used
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lets. The last two-thirds of the key is revised
the letter ““h’’ in two separate couplets/trip- to read as follows:
B)
BB)
BBB) BBBB)
All or all stronger mesenteries fertile (with the possible exception of directives) f) Longitudinal tentacle muscles mesogleal; at least 12 pairs of mesenteries complete g) Outer tentacles with basal battery of microbasic b-mastigophores on aboral side. Submarginal collar. Directives may be sterile
ond Lie long Be YASS SS Ady Sue aa Aint ae an Ae Hormosoma gg) Microbasic b-mastigophores of tentacles scattered, not arrayed in Batches PCCMVESMEIANG 220 9x0 ce es a oes ee ee es Cnidanthus
ff) Longitudinal tentacle muscles ectodermal h) Microbasic b-mastigophores in tentacles; arrayed in batteries and may also be scattered
a) Columm with mesogical papillae...) 2s. eee ee he se Tealidium 11) Column smooth; longitudinal tentacle muscles strongest on oral Se Sets.) RS ee hs ene Anthosactis
hh) No microbasic b-mastigophores in tentacles j) Microbasic amastigophores in tentacles; six pairs of mesenteries SULTS a. BS eee ok ser ee Marianactis jj) Microbasic p-mastigophores in tentacles; six pairs of mesen- teries complete; column divided into scapus and scapulus ... ee ee eee a Ce ee. ns sence os Hadalanthus jij) No tentacular mastigophores k) At least 12 pairs of mesenteries perfect ......... Paranthus kk) Six pairs of mesenteries perfect 1) Sphincter very strong, forming a projecting wall. Mes- enteries not hexamerously arrayed. Few, stout ten- tacles. May have distal papillae ......... Bathydactylus ll) Sphincter strong but not forming a wall. Mesenteries hexamerously arrayed. Column with papillose ne-
MiatOcyse Daltcrics + REY See eT Cnidanthea lll) Column smooth. More than 48 tentacles, closely packed at the rim, in at least 2 cycles ...... Epiparactis
The oldest six pairs of mesenteries sterile, the other stronger mesenteries fertile m) Column divisible into scapus and capitulum. Possibly two sphincters
Pseudoparactis mm) Column not divisible into scapus and capitulum ........... Antiparactis ia Gide | pains OF diescmerics Siete... ........---...---:- Pycnanthus
The three oldest cycles of mesenteries sterile. Mesenteries not divided into filament-free fertile and filament-bearing sterile ones. Oral disc lobed. Sphinc- fie eee ree wer ee es. A eee Fee Soe eas Antholoba
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Fig. 1.
Marianactis bythios, new species
Description.—Body form and size: Ex- panded preserved specimens columnar and colorless (white to yellowish); column di- ameter approximately equal to height. Con- tracted preserved specimens dome-shaped; height one-half to two-thirds diameter. Field notes, and measurements on photos and video recordings, indicate that expanded animals (Fig. 1) were up to “‘five inches [1.e., about 130 mm] across” the tentacle crown, but preserved ones 15-50 mm diameter. Column appears rugose in some specimens, probably due to contraction; ectoderm thin relative to body wall (Figs. 2, 4, 5); often entirely sloughed off.
Base: Flat; equal to column width or somewhat larger; 15-50 mm. Adherent in life.
Tentacles and oral disc: Tentacles color-
Bed of Marianactis bythios in situ. Probe diameter 1 cm.
less in preservation, but photos and field observations note crowns pastel colored— mainly pinks and yellows. Tentacles ar- rayed in several cycles on peripheral half of oral disc; arise from indistinct margin where microbasic amastigophores may be densely packed (Fig. 6). Outer tentacles much short- er than inner; some outer tentacles merely stubs, inner to 15 mm length in animal 50 mm basal diameter. Tentacles taper slightly from base 1—2 mm in diameter to blunt point, but some swollen mid-way along length; transversely ridged in contraction; of equal thickness on all sides; lack basal thickenings. Fewer tentacles than mesen- teries, but more than number of mesentery pairs; commonly about 60 tentacles. Oral disc capable of covering tentacles complete- ly. Other details obscure due to contraction of most individuals examined.
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Fig. 2. Cross section of Marianactis bythios at mid- column. Note regular array of mesenteries and diffuse retractor muscles. Fourth cycle mesenteries (X) are minimally developed, secondary mesenteries (S) are fertile, column mesoglea (M) is thick, and ectoderm (arrow) is thin. CAS 065172 (holotype).
Internal anatomy: Mesenteries regularly arrayed, not according to Actinostola rule (Fig. 2). Four cycles in most specimens; only first order complete and with small oral and marginal stomata; highest (fourth) order barely developed but seemingly wider proximally than distally, sterile, lacking fil- aments; filaments of penultimate (third) order absent from distal half. All except highest order (with possible exception of penultimate order in some individuals) fer- tile, including directives (Fig. 3). Only males seen; sexes presumably separate. Retractor muscles strong, diffuse, with increasingly wider processes centrally (Figs. 2, 3). Pari- etobasilar muscles with short detached pen- non (Fig. 2); not evident at mid-body.
Sphincter muscle mesogleal, reticulate; composed of very small alveoli of uniform (Fig. 4) or slightly irregular (Fig. 5) size. Best
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a
AB “=? / +
:
Fig.3. Cross section through pair of fertile directive mesenteries, Marianactis bythios. CAS 065170.
developed at margin, tapering proximally; hugs endodermal side. Mesoglea on ecto- dermal side fibrous (Fig. 4).
Longitudinal musculature of tentacles strong, ectodermal; circular muscles not ap- parent. Oral disc circular muscles ectoder- mal, disrupted where tentacles insert on oral disc (Fig. 7).
Actinopharynx of typical actinostolid length and rugosity; white in color or rarely violet-brown (as is common in deep-water actinians); two symmetrical siphonoglyphs attach to directive mesenteries; siphono- glyphs not especially prolonged.
Cnidom: basitrichs, microbasic p-masti- gophores, spirocysts, microbasic amasti- gophores.
820 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig. 5. Longitudinal section of mesogleal sphincter of Marianactis bythios. CAS 065172.
VOLUME 102, NUMBER 4
Fig. 6. bythios. CAS 065172.
Distribution and size of cnidae. —(Letter corresponds to illustration in Fig. 8.)
Tentacles: spirocysts (A) (26.1) 27.5—49.2 (54.9) xX 2.5-4.7 umn = 41 N= 4/4 robust spirocysts (B) 43.6-73.1 x 4.6-7.9 um n = 35 N = 4/4 basitrichs (C) 35.3-42.3 x 2.5-4.9 um n= 34N = 4/4 microbasic amastigophores (D) 29.8— 36.0 x 4.3-5.0 (5.4) umn = 18N = 2/3 Actinopharynx: microbasic p-mastigophores (E) (27.3) 30.4-45.9 x 4.3-6.0 umn = 36 N = 3/3 Mesenterial filaments: microbasic p-mastigophores (F) 28.4-44.3 (48.0) x (3.9) 4.2-6.2 umn = 44N = 4/4 Column: basitrichs (G) (19.1) 20.3-—27.3 (27.8) x 2.3-3.7 um n = 44 N = 4/4
Intersection of column and tentacles. Arrow indicates layer of microbasic amastigophores, Marianactis
Discussion. — Habitat, range, and natural history: The eight specimens of Marianactis bythios examined were collected from the Anemone Heaven portion of the Burke hy- drothermal field (18°10.9'N, 144°43.2’E, 3660 m) and from the Alice Springs site (18°12.6’N, 144°42.4’E, average depth 3640 m). All were originally attached to rocks, although some were detached during col- lection.
Members of this species were the domi- nant inhabitants of the region peripheral to the vent openings. Population density was high in places, but tentacles of adjacent in- dividuals generally did not make contact; the animals seemed to be evenly spaced. At both sites, the plume of vent water appeared quite ““smoky.”’ Where emerging vent water was Clear, such as at the Ilium field and the Snail Pits portion of the Burke field, this anemone occurred in smaller numbers. We can offer no explanation for this correlation.
Even individuals attached to rock near
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Fig. 7. Longitudinal section of Marianactus bythios at intersection of tentacles and oral disc. T = coelenteric space within tentacle; M = mesentery; arrows = region lacking ectodermal muscles. MNHN specimen.
the lip of the openings were never posi- tioned where they were exposed to undi- luted emerging vent water, unlike some crabs, snails, and shrimps, which occurred in the vent openings themselves. Therefore, it is unlikely that the actinians contacted water in excess of a degree or so Over am- bient, which is 1.6°C; more than a meter from the vent, water temperature was es- sentially ambient. At both sites, abundance of actinians declined with distance from the vent opening. They extended tens of meters from the emerging water, being found fur- ther from vent openings than organisms of any other taxa associated with the vents. The outermost edge of their distribution op- erationally defined the periphery of the vent field.
Many shallow water sea anemones pos- sess intracellular algal symbionts that pro- vide fixed carbon to their hosts (e.g., Mus- catine 1974). Morphological and behavioral adaptations to them have evolved in some species (e.g., Lewis 1984). Several inverte- brate taxa associated with hydrothermal vents bear endosymbiotic bacteria that ox- idize reduced compounds (sulfide, meth- ane), providing an energy source for their hosts (Childress et al. 1987; Stein et al. 1988). Therefore, we were especially alert for mor- phological evidence of symbionts in this anemone, but found none. Doumenc & Van- Praét (1988) concluded that the diet of the vent actinian Cyananthea hydrothermala includes bacteria, but that they are not in symbiosis.
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A B
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20 Am
Fig. 8. Cnidae signature of Marianactis bythios. See text for explanation.
Nematocyst batteries: Whether the mi- crobasic amastigophores of the tentacles are organized into batteries is uncertain. That type of cnida seems absent altogether from some individuals, and scattered in the ten- tacles of others. But the holotype has a dense layer of them (Fig. 6) in what is either the distalmost column or basalmost tentacles— with no distinct margin, a dividing line can- not be drawn. Carlgren’s (1949) catalog re- fers repeatedly to nematocyst batteries, but contains no definition of the term. At least
for actinostolids, the implication is that the microbasic b-mastigophores in stinging bat- teries are extraordinarily large (e.g., Carl- gren 1921, in the description of Tealidium jJungerseni). The microbasic amastigo- phores of M. bythios are of unremarkable size.
Comparison with other vent species: The description of Cyananthea hydrothermala Doumenc & Van-Praét, 1988, the only pre- viously described vent actinian, is incom- plete and tentative, being based on a portion
824
of one poorly preserved specimen. Thus, some of its anatomy had to be inferred, and its attribution to family Actinostolidae was mainly due to the absence of acontia. In- deed, critical features such as mesenterial arrangement and whether nematocyst bat- teries are present seemingly could not be determined; hence our omission of it from our revised key. Marianactis bythios clearly differs from this species in cnidae (specifi- cally tentacle basitrichs and spirocysts of our species are larger, and ours lacks colum- nar microbasic p-mastigophores), in tenta- cle arrangement, and in color.
Phylogenetic relationships within the Ac- tinostolidae: Many genera of actinostolids, as is true in some other actinian families, are defined by unique combinations of char- acters rather than by singular features. It is this mosaic nature of diagnostic characters that makes inferences about evolution dif- ficult.
Etymology: The specific epithet bythios means “‘of the deep”’ in Greek.
Type Locality and Specimens
Holotype.—Department of Invertebrate Zoology, California Academy of Sciences (CAS), catalog #065172; male; from Alice Springs, Mariana back-arc basin. Includes 10 microscope slides from it.
Paratypes.—CAS #065171; two speci- mens, one sectioned (male); from Anemone Heaven, Mariana back-arc basin. Includes 10 microscope slides from sectioned spec- imen.
CAS #065170; one specimen; male; from Alice Springs, Mariana back-arc basin. In- cludes 10 microscope slides from it.
National Museum of Natural History (USNM), catalog #84401; one specimen; from Burke field, Mariana back-arc basin; includes 10 microscope slides from it.
USNM #84402; one specimen; from AI- ice Springs, Mariana back-arc basin.
Museum Nationale d’Histoire Naturelle,
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Paris (MNHN); one specimen; from Burke field, Mariana back-arc basin.
MNHN; one specimen; from Alice Springs, Mariana back-arc basin; includes 10 microscope slides from it.
Acknowledgments
Michel Boudrias and Scott France aided in collecting the material. Collection was supported by NSF grant OCE83-11258 and a grant from the National Geographic So- ciety. Sections were cut and stained courtesy of Andromeda Systems. We thank Cadet Hand, Michael Kellogg, and Karen Rie- mann-Zumeck for critical and constructive reading of an earlier version of the manu- script, Verena Tunnicliffe for a copy of a relevant paper, and Jean de Mouthe for pre- paring the figures.
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Stein, J. L., S: C. Gary, R. R- Hessler, S. Ohta, R. D. Vetter, J. J. Childress, & H. Felbeck. 1988. Chemoautotrophic symbiosis in a hydrothermal vent gastropod.— Biological Bulletin 174:373- 378:
Stephenson, T. A. 1920. On the classification of Ac- tiniaria. Part I.— Forms with acontia and forms with a mesogloeal sphincter.— Quarterly Jour- nal of Microscopical Science 64 (new series): 425-574.
Stuckey, F.G. A. 1909. A review of the New Zealand Actiniaria known to science, together with a de- scription of twelve new species.— Transactions of the New Zealand Institute 41(for 1908):374— 398.
(DGF) Department of Invertebrate Zo- ology, California Academy of Sciences, Golden Gate Park, San Francisco, Califor- nia CA 94118; (RRH) Scripps Institution of Oceanography, La Jolla, California 92093.
PROC. BIOL. SOC. WASH. 102(4), 1989, pp. 826-865
A REVISION OF THE GENUS ASPIDOSIPHON (SIPUNCULA: ASPIDOSIPHONIDAE)
Edward B. Cutler and Norma J. Cutler
Abstract. —The 64 putative species of the sipunculan genus Aspidosiphon and the morphological characters used to differentiate them are critically reviewed. The monograph of Stephen & Edmonds (1972) is used as a starting place and all changes made in the intervening years are reiterated here. All available type material was studied and new collections of Hawaiian and Caribbean material are used to analyze within-deme variation. Hook and anal shield morphology are determined to be broadly useful at the species level, four characters (lon- gitudinal muscle layer, retractor muscle origins, caudal shield, nephridia length) in a more restricted manner to separate subgroups, and three (introvert/trunk angle, bifurcated anterior spindle muscle, loosely wound gut coil) are useful in special cases. A new subgenus, Aspidosiphon (Akrikos), is proposed for those five species lacking hooks in rings. A key to, and a discussion of, the 19 remaining species (plus one reduced to subspecies) with the newly designated synonyms are presented. A brief statement of the distribution of each species is given. An overall summary of the zoogeography and habitat shows more endemic species are found in the warm water regions of the Atlantic Ocean than in the Indo-West Pacific, and that only 42% of Aspidosiphon species live
in coral or rock.
This continues our revisionary series on the species of sipunculan worms (e.g., Cutler & Cutler 1985a, b, 1986, 1987b, 1988). With this work we complete our examination of all the genera in this phylum except Phas- colosoma, which is in preparation. The monograph of Stephen & Edmonds (1972) is the starting place for this work (48 species names). Also included (Table 1) are the 11 species erected since that time, the two species transferred into this genus, and the three resurrected names.
The genus Aspidosiphon was erected by Diesing in 1851, and was placed in its own family, Aspidosiphonidae, by Baird (1868) in the order Aspidosiphoniformes Cutler & Gibbs (1985). The name Paraspidosiphon was proposed by Stephen in 1964 as a genus for those species with the longitudinal mus- cle layer separated into bundles. Cutler (1973) reduced it to subgeneric rank and this
has been followed by most other authors (see discussion below).
Whenever possible we have obtained type material to verify the original descriptions. In several cases we have made detailed ob- servations on series of recently collected in- dividuals to evaluate better the traditionally used morphological characters. Recent col- lecting trips to Hawaii, Curacao and Ven- ezuela (Cumana and Isla de Los Roques) have greatly facilitated this effort. The op- - portunity to observe living material is in- valuable. Parts of these collections will be deposited in the National Museum of Nat- ural History, Washington, D.C. as reference material.
We first discuss the morphological char- acters in light of our recent analyses, then discuss those taxa not clearly belonging to this genus. Following are a key to all the species we consider valid, a section where
VOLUME 102, NUMBER 4
each of these species is discussed including a synonomy, a discussion of any newly added junior synonyms, and a summary of the known distribution of each species. A short zoogeographical summary of the genus con- cludes this work.
For clarity in the Morphological Char- acters section, the recent work of Saiz Sa- linas (1984) needs to be mentioned here. His redescription of Quatrefages’ 1865 species from the Paris Museum has led to the elevation of A. coyi and A. /aevis as se- nior synonyms of the more familiar A. trun- catus for the former and the large A. cum- ingii/klunzingeri complex for the latter. In both cases holotypes are now available to science, which is not the case for the more familiar names. In some ways this action 1s analogous to that of Rice & Stephen (1970) where they resurrected the older and long unused names of Gray and Baird.
The following abbreviations are used in the text for the museums from which we borrowed material: American Museum of Natural History, New York (AMNH); Bnit- ish Museum (Natural History), London (BMNH);: Muséum National d’Histoire Na- turelle, Paris (MNHN); Museum fur Na- turkunde der Humboldt-Universitat zu Berlin (MNHU): Musée Océanographique Monaco (MOMYV); Naturhistoriska Riks- museet, Stockholm (NHRS); Royal Scottish Museum, Edinburgh (RSME); National Museum of Natural History, Washington (USNM); Zoologisk Museum, Copenhagen (UZMK); Zoological Institute, Academy of Science, Leningrad (ZIAS); Zoological In- stitute, Tohoku University, Sendai (ZITU); ZoOdlogisch Museum, Universiteit van Am- sterdam (ZMUA); Zoological Museum, University Bergen (ZMUB): Zoologisches Museum, Universitat Hamburg (ZMUH).
Morphological Characters
1. Introvert hooks and spines.— As in most genera, the introvert bears (in all but three species) some array of specialized structures
827
usually referred to as hooks. Voss-Foucart et al. (1977) have shown these to lack chitin but consist of a horny protein. Many hooks are arranged in regular rings around the dis- tal portion of the introvert and may have either one or two points (uni- or bidentate). On some species scattered hooks also are found proximally and in two species only scattered hooks are found. Additionally, epidermal structures of varying sizes and shapes called spines are arranged in a ran- dom manner on the proximal portion of the introvert. Examination of the literature re- veals that the term “spine’’ meant different things to different authors, sometimes being used as a synonym for what others would term “‘unidentate hook.” Since hooks and spines come in a wide variety of arrange- ments, sizes, and shapes, and often grade into one another, it is easy to understand the genesis of this problem that is unique to Aspidosiphon. We will attempt some clar- ification and definitions.
In the phylum Sipuncula, the term “hook” has been applied to structures having a wide variety of shapes, sizes, and arrangement. It is clear that a Themiste hook is different from a Nephasoma hook and that both dif- fer from Phascolosoma hooks. Structures that have been called “spines” are similar to “hooks” of Themiste and of some Phas- colion species. We now propose calling all of these introvert structures hooks. The term “spine” will be restricted to conical pointed anal shield units. The hook’s apex points posteriorly (away from the mouth) with the convex curvature being anterior.
The different types of hooks are defined as follows:
Type A: Compressed hooks. —Usu- ally arranged in rings, occasionally scattered, laterally compressed, and having, in a side view, a distinct pos- terior curve. These may be unidentate or bidentate (Fig. 1A, B). When a sec- ondary tooth is present it may be vari- able in size, sometimes reduced to a
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Table 1.—Original and proposed names of the Aspidosiphon species.
Subgenus Aspidosiphon s.s.
Aspidosiphon albus Murina, 1967
Aspidosiphon brocki Augener, 1903 Aspidosiphon carolinus Sato, 1935
Aspidosiphon cylindricus Horst, 1899 Aspidosiphon elegans (Chamisso & Eysenhardt, 1821) Aspidosiphon exhaustus Sluiter, 1912 Aspidosiphon exiguus Edmonds, 1974 Aspidosiphon exilis Sluiter, 1886
Aspidosiphon gerouldi ten Broeke, 1925 Aspidosiphon gosnoldi Cutler, 1981
Aspidosiphon gracilis (Baird, 1868)
Aspidosiphon hartmeyeri Fischer, 1919 Aspidosiphon hispitrofus LiGreci, 1980 Aspidosiphon homomyarius Johnson, 1964 Aspidosiphon imbellis Sluiter, 1902 Aspidosiphon inquilinis Sluiter, 1902 Aspidosiphon jukesii Baird, 1873
Aspidosiphon kovaleskii Murina, 1964 Aspidosiphon longirhyncus Cutler & Cutler, 1980 Aspidosiphon macer (Sluiter, 1891)
Aspidosiphon mexicanus (Murina, 1967) Aspidosiphon misakiensis Ikeda, 1904 Aspidosiphon muelleri Diesing, 1851 Aspidosiphon ravus Sluiter, 1886
Aspidosiphon spinalis Ikeda, 1904
Aspidosiphon spinosus Sluiter, 1902 Aspidosiphon spiralis Sluiter, 1902
Aspidosiphon thomassini Cutler & Cutler, 1979 Aspidosiphon tortus Selenka, de Man & Biilow, 1883 Aspidosiphon venabulum Selenka, de Man & Biilow, 1883 Aspidosiphon zinni Cutler, 1969
Subgenus Paraspidosiphon
Aspidosiphon ambonensis Augener, 1903 Aspidosiphon angulatus Ikeda, 1904
Aspidosiphon brasiliensis Cordero & Mello-Leitao, 1952 Aspidosiphon coyi Quatrefages, 1865
Aspidosiphon cumingii Baird, 1868
Aspidosiphon exostomus Johnson, 1964 Aspidosiphon fischeri ten Broeke, 1925
Aspidosiphon formosanus Sato, 1939
Aspidosiphon gigas Sluiter, 1884
Aspidosiphon grandis Sato, 1939
Aspidosiphon havelockensis Haldar, 1978 Aspidosiphon insularis Lanchester, 1905 Aspidosiphon johnstoni Edmonds, 1980 Aspidosiphon klunzingeri Selenka, de Man & Biilow, 1883 Aspidosiphon laevis Quatrefages, 1865
Aspidosiphon levis Sluiter, 1886
Aspidosiphon major Vaillant, 1871
Aspidosiphon makoensis Sato, 1939
Aspidosiphon ochrus Cutler & Cutler, 1979 Aspidosiphon pachydermatus Wesenberg-Lund, 1937 Aspidosiphon parvulus Gerould, 1913
no change*
A. elegans
A. elegans species inquirenda no change
A. muelleri no change
A. elegans
A. misakiensis no change
no change
. misakiensis . muelleri
. elegans
. muelleri
. muelleri
. muelleri
. muelleri
. mexicanus* species inquirenda no change* no change
no change
A. elegans
A. elegans
A. elegans
no change
no change*
A. muelleri no change* no change*
A RAPA AA A Pw
A. tenuis
A. laevis
A. laevis
no change
A. laevis
A. steenstrupli no change
A. tenuis
A. laevis
A. laevis
A. tenuis Phascolosoma perlucens A. laevis
A. laevis
no change
A. tenuis
A. laevis
A. steenstrupli A. steenstrupii A. laevis
no change
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Table 1.—Continued.
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Aspidosiphon planoscutatus Murina, 1968 Aspidosiphon quatrefagesi Saiz Salinas, 1984 Aspidosiphon pygmaeus Fischer, 1921 Aspidosiphon schnehageni Fischer, 1913 Aspidosiphon semperi ten Broeke, 1925 Aspidosiphon speciosus Gerould, 1913 Aspidosiphon speculator Selenka, 1885 Aspidosiphon spinososcutatus Fischer, 1922 Aspidosiphon steenstrupii Diesing, 1859 Aspidosiphon tenuis Sluiter, 1886
Aspidosiphon trinidensis Cordero & Mello-Leitao, 1952
Aspidosiphon truncatus (Keferstein, 1867) Golfingia mokyevskii Murina, 1964
* Now in new subgenus 4. (Akrikos).
small knob. A transition zone in some species exists at the proximal end of the rings of hooks where one may find a gradual widening of the anterior base of the unidentate hooks. Sometimes these scattered hooks are rounded at the anterior-lateral corners but still compressed posteriorly looking like a ship’s stout mast and sail.
Type B: Pyramidal hooks.—Have triangular bases, the anterior side of which is shorter than the lateral sides, are usually less curved than Types A or C, are variably pigmented (dark to light), and translucent (Fig. 1E, F). The borderline between Types A and B is not clear in all species.
Type C: Conical hooks.—Have a nearly circular cross section (cone shape), a gentle posterior curve, and are usually opaque and dark colored (Fig. 1C, D). This type is found on the dorsal side of A. elegans’ introverts.
When introvert skin is removed and placed on a slide in a drop of glycerin for closer examination, the orientation of these hooks can add to the confusion. If viewed from the anterior or posterior (instead of laterally), scattered unidentate compressed hooks look very much like pyramidal hooks (Fig. 1G). However, a compressed hook has
no change
. laevis
. muelleri
. gracilis schnehageni . Steenstrupli
. laevis
. Steenstrupii
. parvulus
no change
no change
A. steenstrupli
A. coyl
Antillesoma antillarum
am wm Pw w AP
a narrower base. Further distortion can be caused if the hooks are not lying flat on the slide. Scanning electron micrographs can help reveal the natural configurations and the three dimensionality of these structures.
The shape of the clear area (less dense to transmitted light) in the hook has limited taxonomic value. In most species there is an ill-defined triangular area, but in 4A. steenstrupli and A. elegans there is a thin, posteriorly directed, tongue-like extension (Fig. 2).
The height of the hook has sometimes been used as a diagnostic character, but this has limitations. The unidentate A. /aevis complex (as defined below) shows a clear correlation between trunk size and hook size (5-10 mm worms have 20-40 um hooks, 20-30 mm worms have 40-60 um hooks, and 40-70 mm worms have 60-80 um hooks). The same pattern is shown by an analysis of 32 specimens of A. steenstrupii ranging in trunk length from 7-50 mm. Hook size varied from 30-90 um with larger worms having larger hooks (a linear regres- sion of these data gave a positive slope of 0.89). Therefore, hook size should not be considered in isolation from trunk size. De- spite this a clear pattern does appear with certain species (e.g., the members of the new subgenus proposed here) always having small hooks (under 30 um) and some species
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Fig. 1.
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Introvert hooks: A, Compressed, bidentate (Type A) from A. muelleri, posterior view; B, Compressed
unidentate (Type A) from A. misakiensis; C-D, Conical (Type C) from A. elegans; E, Pyramidal (Type B) from A. parvulus; F, Pyramidal (Type B) from A. steenstrupii viewed from above; G, Pyramidal (Type B) from A. steenstrupli, different angles on light microscope. Scale line = 10 um, for G = 20 um.
with only larger hooks. The central problem is that many species have both large and small hooks.
In general, some species lack unidentate hooks, some lack bidentate hooks, some have both Type A and B hooks while others have only Type A hooks. Certain species will have both kinds of compressed and Type B hooks. In summary, if one examines the most distal rings of hooks and differentiates between unidentate Type A and Type B
hooks, hook morphology can be useful to the systematist in almost all cases and hook size can help in some cases.
2. Anal shield (degree of development, na- ture of units, grooves). — At the anterior end of the trunk, horny protein (not chitin) forms an array of cuticular units varying in degree of development (Voss-Foucart et al. 1977). At one extreme is A. mexicanus or A. tho- massini with a collection of small scattered units sometimes looking more like an area
VOLUME 102, NUMBER 4
Fig. 2. extension in C. Scale lines = 20 um.
of rough skin. At the other end of the con- tinuum is A. /aevis or most A. muelleri where the units are compacted to form a thick, dark, solid mass. These two species are among those that have shields with well de- veloped longitudinal and/or transverse grooves. When a shield has aggregations of units separated by grooves we refer to these aggregations as plates. The shield nearest
the mid-dorsal anus is the dorsal part while
that nearest the introvert is the ventral part.
The nature of the shield units may undergo slight changes with age (see A. jukesii in Cut- ler & Cutler 1979a:970) and may be mod- ified by the size/shape of the shell (gastro- pod vs. scaphopod) or other space occupied by the worm. When the introvert is retract- ed this shield functions as an operculum. While some within-deme variation does ex- ist, the morphology of the anal shield is con- sistent enough and distinct enough to be useful to the systematist.
3. Caudal shield (degree of develop- ment). — At the posterior end of the trunk there is a epidermal structure (horny pro- tein) present in most (but not all) species. This shield assumes various forms in living worms and, therefore, when preserved, can also vary from rather flat to pointed to pa- goda shaped. They have a variable number of radially arranged grooves or furrows, but this attribute is not species specific. Even within a deme the degree of development
831
Internal hook structure: A, A. elegans, B, A. steenstrupii, C, A. tenuis. Note absence of tongue-like
(thickness) varies (Fig. 3). In a species with extensive historical data indicating a “‘nor- mal”’ shield, one may find individuals with very weakly developed shields. In such an- imals the shield may be reduced to some papillae and only a vague suggestion of a chitinoid layer. It is probable that in many species the genetic potential is there, but its expression is variable and responds to en- vironmental stimuli or age. Some species (A. laevis) always have a shield and others (e.g., A. mexicanus or A. zinni) never do. When a shield is present, there is little to distinguish one species from another using this character. Therefore, aside from pres- ence/absence, the caudal shield has limited value to the systematist.
4. Introvert retractor muscles (origin, de- gree of fusion).—The single pair of long muscles used to retract the introvert insert at the anterior end and originate from the ventral trunk wall in the posterior third of the worm. The placement of these origins (attachment) is usually included in species descriptions, often with a significant lack of precision (near the posterior end, in the last quarter), but sometimes it is used as a di- agnostic (or key) character. While in other genera we have found that retractor origins appear to shift anteriad as the animal grows (zone of trunk growth being posterior to the origins), it is difficult to generalize about this in Aspidosiphon. If one calculates the dis-
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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig. 3. Caudal shields of A. elegans. Three 17 mm worms from one deme in Ponape to show variation in
degree of development. Scale line = 1 mm.
tance as a percentage of trunk length, there do seem to be some species where this value remains reasonably constant over a wide range of sizes (e.g., A. muelleri, 95—100%). However, other species exhibit more vari- ation (see Table 2 for A. elegans, 65-85% and A. steenstrupii, 70-90%). This is not clearly correlated with size. Therefore, while this attribute can be useful for differentiat- ing two subsets, it does not have value at the species level.
The second aspect of this complex that is sometimes noted is the degree of fusion be- tween these two muscles. In the older lit- erature there is considerable confusion on this matter, such that some descriptions al- lege there to be only one or as many as four, or one muscle with one, two or four “‘roots.”’ From what we know about the ontogeny of this functional complex in sipunculans, it is probable that they all begin life with two pairs of retractors (Rice 1976). The dorsal pair is lost and the ventral pair fuses to vary-
ing degrees. However, quantifying this is ex- tremely difficult because extension of the introvert will stretch out the muscle and the point of fusion will move away from the posterior end of the trunk. While one gets the impression that there are different sized muscles and different degrees of fusion, no pattern can be discerned. We are unable to suggest how this information can be uni- versally applied even if species specificity was evident.
5. Spindle muscle attachment. — This thin muscle extends from the body wall at the posterior end of the trunk, through the gut coils, and along the rectum to the body wall just anterior to the anus. While there have been a few reports of this muscle merging with the body wall posterior to the anus, the only cases of this kind we have been able to confirm are in A. Jaevis and A. coyi in which the muscle sends a large branch to the mid-dorsal body wall posterior to the anus. It is not always easy to trace the course
VOLUME 102, NUMBER 4
833
Table 2.—Selected morphological attributes of A. elegans and A. steenstrupii.
Aspidosiphon elegans
Aspidosiphon steenstrupii
Retractor Nephnidia Nephridia Retractor Nephridia Nephridia
Trunk length origin; length; attachment; Trunk length origin; length; attachment;
in mm % t % trunk % nephridium in mm % trunk % trunk % nephridium 6 83 fie 89 7 fal 42 67 7 86 70 95 7 79 57 iP) 7 _ vi 95 10 is 50 50 8 63 63 99 11 82 45 TS 9 67 50 99 11 73 99 67 10 70 60 67 12 vie. 50 67 10 70 70 93 12 15 99 99 11 82 64 99 14 79 54 IS 14 86 vi 88 15 Ty 50 — 15 60 67 50 15 80 60 56 15 80 40 83 15 73 67 50 16 63 53 85 15 73 60 50 16 81 63 a9 17 82 41 71 16 88 88 95 19 79 58 50 17 76 35 99 19 68 47 67 17 65 65 82 19 84 47 90 19 79 58 86 20 85 75 40 19 74 47 99 21 76 57 _ 21 90 52 64 22 77 68 50 22 86 86 95 24 71 42 75 23 78 57 99 25 80 48 58 25 84 60 67 26 81 69 50 32 78 56 99 27 70 81 99 28 75 65 99 33 79 67 50 a3 85 52 53 34 85 50 50 50 76 70 33 50 80 54 75
of this muscle, especially in smaller worms or where the spindle muscle seems to merge with the wing muscle. Except for the above species, the muscle originates anterior to the anus and, therefore, in this genus, it is of limited usefulness as a diagnostic character.
6. Fixing muscle number.—In most si- punculan taxa fine thread-like muscles an- chor some part of the anterior intestine to the body wall. The maximum number re- corded in this genus is one and its presence (or absence) has been considered system- atically important by some biologists. Our review of the literature reveals a lack of con- sistency on this point. Our own studies strongly suggest that either: A. Genetic poly-
morphism exists within populations or B. This fragile structure is placed where it can be easily damaged during dissection. While it may be true that some species totally lack this muscle and others usually have one, the possibility of any one worm deviating from the norm is too great to give weight to this character.
7. Nephridia (length, attachment, level of nephridiopore). —Both the nephrostome and the nephridiopores are located at the ante- rior end of the nephridia, a pair of tubular sac-like ventro-lateral organs. These open to the outside at the anterior end of the trunk. Three attributes have been recorded and are sometimes treated as diagnostic.
834
First is the position of the nephridiopores relative to the anus. In the literature most species of A. (Aspidosiphon) and a few A. (Paraspidosiphon) are reported as having nephridia at the level of the anus but slightly posterior to it in the remaining species. In this latter group our own data show this distance to be 3—8% of the trunk length. However, even in these populations a few animals have the nephridiopores and anus at the same level. While there do seem to be other species where this distance is al- most always zero, careful examination of more than ten worms will probably reveal one or two where these openings are not at the same level. It may be possible to say: “80-90% of species X have these openings at the same level while in species Y only 5— 10% do.” But as these kinds of data are not always available (small sample sizes), the information has limited value to the sys- tematist.
A second attribute is the length of these organs. The literature includes statements like “‘very long, long, reach to the base of the retractors, two-thirds/half/one fourth as long as the trunk,’ etc. Our observations show a range of lengths within a population, not correlated with trunk size (see Table 2), but there are some differences among species. Six species have nephridia half the trunk length or less. Many exhibit a broad range (e.g., 25-100%, 45-85%, 50-100%) and a few have only long nephridia (over 85% of the trunk length). With a few ex- ceptions, nephridia length can only be used in a limited manner.
A third attribute often mentioned is the attachment of the nephridia to the body wall by a membranous sheet of connective tis- sue. Occasionally this takes the form of fil- amentous strands of tissue scattered along the nephridia that bind them less tightly. While nephridial attachment often appears in keys to Aspidosiphon, its constancy is overrated. While the original description may state that the nephridia are attached for a particular length (one-half, two-thirds,
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
100%, etc.), subsequent authors either ig- nore it or do not verify it critically. The connective tissue is easily torn, and within a single worm the attachment can differ for each organ. In most species a wide range of values is observed, commonly in the 50- 100% range (see Table 2 for A. steenstrupii, 50-80% and A. elegans, 80—100%) while A. muelleri ranges from O0-100% attached in different reports. Therefore, this attribute has restricted systematic value.
8. Rectal caecum (presence, complexi- ty). —In many sipunculans there is a small caecum on the rectum near the intestinal coil. Certain species of Aspidosiphon are re- ported to lack a caecum while others are said to have one. Two problems exist: If one looks at all of the published reports of any frequently recorded species, one finds inconsistencies, e.g., in A. elegans several authors say it lacks a caecum, others make no mention (this could mean that it is absent or possibly that they did not look for it), and a few report seeing the caecum. Alter- natively, A. muelleri has been said to have a caecum by many authors but a few assert that it lacks one. In the A. muelleri we have recently examined, 25% have a caecum. In A. albus the original describer (Murina 1967c) asserts that the caecum is absent. Subse- quent workers looking at other members of the population (Cutler 1973, Migotto & Di- tadi 1988) saw one.
The second problem is related to this and it is illustrated by our finding a caecum in two of ten A. elegans we examined. Here we have one observer looking at one deme and finding dimorphism. In 30 worms that Mig- otto & Ditadi (1988) dissected they found a caecum in 18 (two of these being “large, villous’’). In other words, 40% of this pop- ulation of a species described as having a caecum was found to lack one. As this is true for one population, it may be true for all. We conclude that it is an error to con- sider the presence or absence of a rectal cae- cum as species specific.
In this genus there is one special case, 1.e.,
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a rectal structure described as: “‘. . . the last part of the rectum densely bordered with long villi-like structures” (Selenka et al. 1883 in A. cumingii) or “rectum with caecum plus many blind tubes attached to both sides” (Sato 1939 for A. grandis) or “‘rectum with a larger caecum, many lobed on both sides” (Selenka et al. 1883 in A. klunzingeri) or “rectal caecum with lappets” (Edmonds 1956 in A. klunzingeri) or “a large cluster of blind sacs as ramified intestinal appendages”’ (Wesenberg-Lund 1937 in A. pachyderma- tus). Only one specimen of A. pachydermatus and two of A. grandis (100, 70 and 100 mm trunks) exist. Of the 12 records of A. klun- zingeri that include morphological com- ments, only the original material (3 worms) has this type of rectal elaboration. For sub- sequent authors, its absence seemed incon- sequential. Similarly, in the nine reports of A. cumingii with morphological comments, only Selenka et al. (1883) record this struc- ture in five worms (the words are used by other authors but only when referring to Selenka et al.). So, while these species sup- ' posedly have an elaborate rectum, several authors have used this name for animals lacking said structure. In museum collec- tions, fewer than 10 worms exist (Baird’s two have been lost) with this condition. The four worms in our 1985 Hawaii material that we are calling A. Jaevis have trunk lengths of 7, 10, 11 and 14 mm. The largest and the smallest bear a single structure with 6-8 short branches or lobes coming off each side. The 10 and 11 mm worms show small lobes, but the precise structure is less clear due to the fragility of the rectum. In our 1988 Venezuela collections we have five A. laevis (10-30 mm trunks), but none of these has a caecum of any kind. Migotto & Ditadi (1988) report large villous, simple and no caeca in a single population.
The question posed above reappears here: If an author did not mention this structure, does that mean it was not present? In those cases when an author specifically says that the complex caecum was not present, what
835
significance does that have? Historically the position that this is insignificant and vari- able within a population has implicitly pre- vailed. Our recent data confirm that and suggest that A. /aevis has the genetic poten- tial (perhaps polygenic), but this potential is not always expressed, and when it is ex- pressed it may be overlooked by an observ- ee
9. Intestinal coils (tightness).—In some other genera the number of gut coils has been used as a systematic character, but we have elsewhere shown this to be size de- pendent and not useful. In Aspidosiphon the interest 1s restricted to the nature of the coil- ing. In most species a regular, compact dou- ble helix is present, but A. misakiensis ex- hibits a loose, less regular helix (Fig. 4). This has also been reported by a few authors for A. elegans. The helix is maintained by the fine strands of muscle linking the coils to the axial spindle muscle. In those animals having a looser gut coil the linkage is not continuous and the strands may be longer.
10. Longitudinal muscle (bundles, anas- tomosing, fracturing).—The body wall of sipunculans has an internal layer of lon- gitudinal muscle. In one subgenus (4. Aspidosiphon) this is an undivided sheet, almost. In the other subgenus (A. Paraspi- dosiphon) this layer is divided into separate bundles. However, it is not always a clear- cut dichotomy. Intermediate conditions of two general types exist.
First, ten putative species of A. (Aspido- siphon) have been described as having frac- turing of the muscle layer in the anterior dorsal trunk. Commonly this is restricted to the area under the anal shield, but in some species the fractures continue out be- yond these borders for a small (10-20% of the trunk length) but variable distance.
The second variation occurs in species of A. (Paraspidosiphon) where the longitudinal muscle bands are not distinct. The degree of variation is much greater than in other genera with these bundles. While several have distinct rarely anastomosing bundles,
836
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig. 4.
Intestinal coil of A. misakiensis showing irregular loose arrangement (A—anal shield, M—retractor
muscle, N—nephridium, R—rectum). Scale line = 2 mm.
many exhibit a modest degree of anasto- mosing and others show frequent cross link- ages. In these latter the layer appears like a continuous sheet that has split or fractured and not like distinct bundles (e.g., A. fi- scheri).
The number of muscle bands varies con- siderably within a population and within an individual (25-35 anteriorly and 15-25 posteriorly). It is difficult in a few cases, especially in small worms, to know whether one is looking at an A. (Paraspidosiphon) with much anastomosing of bundles or at an A. (Aspidosiphon) with some fracturing of a layer.
While this character state may be used for separating subgenera, it is neither species
specific nor discriminating at that level. Since the subdivision of a continuous layer appears to be an homoplastic, apomorphic condition (Cutler & Gibbs 1985) it is pos- sible that it has arisen more than once with- in this genus. Therefore, using it as the single attribute to separate subgenera may mask actual phylogenetic relationships.
11. Angle of introvert to trunk. —In most Aspidosiphon the extended introvert pro- trudes at an angle of 75—90° to the main axis of the trunk at the ventral edge of the anal shield. However, in at least three species, all with very weakly developed anal shields, this angle is reduced to 45-60°. While this is not broadly useful, it can help in these special cases.
VOLUME 102, NUMBER 4
Summary.—Two characters that have broad taxonomic usefulness are the hook and anal shield morphology. Four charac- ters useful for separating the genus into dif- ferent subsets are: (1) longitudinal muscle layer continuous or divided, (2) retractors originating in the most posterior 5% of the trunk or in the 70-80% range, (3) caudal shield developed or not, and (4) nephridia length (less than 50% of trunk length, more than 75%, or a broad range). An introvert/ trunk angle of less than 75° separates three species, while a bifurcated anterior spindle muscle and a loosely wound gut coil each characterizes one species. The presence/ab- sence of fixing muscles or caecum, the place- ment of the nephridiopores, and the attach- ment of the nephridia to the body wall are too variable to have any systematic value in this genus.
Systematic Section Aspidosiphon Diesing, 1851
Diagnosis. — Introvert usually longer than trunk. Recurved hooks in numerous rings (absent in three, only scattered in two species). Trunk with anal shield composed of hardened units (occasionally inconspic- uously developed). Introvert protrudes from ventral margin of shield. Body wall either with continuous longitudinal muscle layer or with longitudinal muscle layer gathered into anastomosing, sometimes ill-defined, bundles. Oral disk with tentacles enclosing dorsal nuchal organ but not mouth. Con- tractile vessel without villi. Two introvert retractor muscles sometimes almost com- pletely fused. Spindle muscle attached pos- teriorly. Two nephridia. One species may exceed 100 mm but most less than 40 mm long.
We herein create a new subgenus for a set of five species as defined below. The major character state we focus on is the absence of compressed hooks in rings, a plesio- morphic character state for this entire class (Cutler & Gibbs 1985). One could argue that
837
these species represent primitive transition forms from a very early ancestral stock, but we propose instead that this trait has been secondarily lost through subsequent evo- lution, 1.e., a type of reversal. In support of this, note the atypical ecology of these taxa (e.g., interstitial or abyssal, none boring in coral or rock) suggesting an ecological spe- cialization.
The following four taxa are not consid- ered valid members of this genus and are discussed first. After the key, the remaining species are presented alphabetically within subgenera.
Aspidosiphon cylindricus Horst, 1899
Aspidosiphon cylindricus Horst, 1899:195— 198, text-ugs. 3-4.—Stephen & Ed- monds, 1972:222-—223.—Not Sluiter, 1902:18-19.
Material examined.—ZMUA, Sluiter’s material (V. Si. 26.8).
Discussion. — This species was based on a single worm that disappeared from the Lei- den museum prior to 1930 (van der Lund, pers. comm.). Enough questionable but now unverifiable features exist (especially the hook morphology) that we place this name on the list of species inquirenda pending fu- ture clarification. Sluiter’s (1902) material was reexamined and is herein referred to A. elegans.
Aspidosiphon insularis (Lanchester, 1905)
Aspidosiphon insularis Lanchester, 1905b: 40, pl. 2, fig. 4.—Gibbs & Cutler, 1987: 56.
Paraspidosiphon insularis. —Stephen & Ed- monds, 1972:247.
Material examined. —BMNH, (1924.3.1.80).
Discussion. —This worm is in poor con- dition, but, as noted by Gibbs & Cutler (1987), it is clearly a Phascolosoma and based on hooks and papillae is a junior syn- onym of P. perlucens.
type
838
Aspidosiphon macer (Sluiter, 1891)
Phascolosoma macer Sluiter, 1891:114—115, pl. 2, figs. 13-14; 1902:34.
Golfingia macra.—Stephen & Edmonds, 1972:149.—Cutler & Murina, 1977:183.
Aspidosiphon macer. —Cutler & Cutler, 1986:568.
Material examined.—ZMUA, type and only specimen (V. Si. 65).
Discussion. — This putative taxon is based on a single specimen that has been thor- oughly dissected over the years. The generic status is not altogether firm since the anal shield is very poorly developed and the in- trovert does not appear to be ventrally dis- placed. The gut is missing and while Sluiter asserted that the spindle muscle is not at- tached posteriorly, there is a muscle coming from the center of the caudal shield that we interpret as the broken posterior portion of this muscle. Sluiter said there were no hooks but there are about ten distinct rings of sharply pointed, unidentate hooks. The lon- gitudinal muscle layer is undivided. While Sluiter asserted that the tentacles surround the mouth, the introvert is not extended so one cannot really tell how the tentacles are arranged. They appear to us to be clustered towards one side as in this genus. Therefore, until additional material is obtained to clar- ify this uncertainty, we place this name on the list of species inquirenda.
Golfingia mokyevskii Murina, 1964
Golfingia mokyevskii Murina, 1964a:256— 259, figs. 4—5.
?Aspidosiphon mokyevskii.—Gibbs et al., 1983:302.
Material examined.—ZIAS, type mate- rial.
Discussion. —Gibbs et al. (1983) suggest- ed that Murina’s species might be an As- pidosiphon; however, it is now clear that this was an error. The 50 long tentacles, absence of hooks, anastomosing longitudinal muscle bands, large dark papillae at the base of the
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
introvert, and especially the presence of contractile vessel villi all support placing this name in the synonomy of Antillesoma antillarum.
Key to Aspidosiphon species
1. Hooks not present, or if present, NOLIN MNES: . se A. (Akrikos) 2 — Hooks arranged in rings on distal
Portion of MtLOVEFE (>. -B-10 ee 6
2. ‘Introvert hooks absent |/)..9gae 3 — Scattered introvert hooks present
128 Je Gee a ee 5
3. Anal shield of tightly packed, uni- form sized, pale, flat units with dis- tinct angular margin A. albus Murina
— Anal shield of dispersed, often dark units, sometimes very poorly de- veloped, with indistinct margin... 4
4. Anal shield units distinct, dark; those around margin usually pointed cones
... A. venabulum Selenka, de Man & Bulow
— Anal shield units indistinct, widely spaced, flat, sometimes arranged in indistinct rows
BOEHES A. thomassini Cutler & Cutler
5. Anal shield ill defined and diffuse, trunk usually more than 5 mm, shallow warm water
Ae, “eas nah wera te A. mexicanus Murina
— Anal shield well defined and com- pact, trunk usually less than 5 mm, deep cold water ..... A. zinni Cutler
6. Longitudinal muscles in continu- ous layer (except under anal shield)
bapeleonst A. (Aspidosiphon)
— Longitudinal muscle layer divided into separate (or anastomosing) bundles .. A. (Paraspidosiphon) .. 13
7. Anal shield with extensive array of furrows present, not just around
Marpim! 6). Hi): AVG... Se 8 — Anal shield with randomly distrib-
uted hardened units, lacking ex-
tensive grooves/furrows ........ 9
VOLUME 102, NUMBER 4
8.
10.
iM:
12s
NS:
14.
IS:
16.
Individual units form into longi- tudinal ridges over dorsal half of anal’shield 2... A. muelleri Diesing Individual units arranged in offset squares or rectangles «wel geek ee a A. spiralis Sluiter All hooks unidentate, ill-defined analsSmicld 2... . A. gracilis (Baird) Distal hooks bidentate, anal shield distinct All compressed hooks bidentate followed by dark conical hooks ..
... A. elegans Chamisso & Eysenhardt Distal bidentate compressed hooks followed by proximal unidentate ones Interstitial, introvert 2—5 times the trunk length, nephridia 25-33% monk so. A. exiguus Edmonds Occupies coral or shells often sub- tidal, introvert 1-3 times trunk, nephridia more than 50% of trunk LES 25 ee re 12 Normal gut helix, lives in gastro- pod shells, anal shield units square, arranged in rows, and each made up of smaller granular subunits
sag oe A. gosnoldi Cutler Gut coils loose or absent, does not occupy gastropod shells, anal shield units more solid and randomly ar- AMC CO secs esl A. misakiensis Ikeda Anal shield ungrooved or, if pres-
ent, only as short marginal ones
oe © © © © © © we we
CFLS SVE) a eae ns 14 Anal shield with extensive grooves
or furrows present (Fig. 5B) .... 18 Distal hooks bidentate ......... 15
All hooks unidentate ...........
» oo Bae es Sees A. planoscutatus Murina Compressed hooks bidentate fol- lowed by dark pyramidal hooks .
A. steenstrupii Diesing Compressed hooks of both types, pyramidal hooks pale, if present 16 No pyramidal hooks, longitudinal muscle bands distinct, compressed hooks over 30 um tall, retractor
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origins 75-88% ..... A. tenuis Sluiter Pale pyramidal hooks present, lon- gitudinal muscle bands indistinct, hooks less than 30 um tall, retrac- tor ongins 95—100% 222... 35... We, 17. Anal shield marginally becomes diffuse forming cones or spikes, nephridia more than '2 trunk ... | Seno rnp if! tay be A. parvulus Gerould — Anal shield with distinct margins, no cones or spikes, nephridia less than % trunk ...A. fischeri ten Brocke 18. All hooks unidentate, retractor origins not at posterior end (60- SOCOM ney cas on A. laevis Quatrefages — Distal hooks usually have very small secondary tooth, retractor origins at posterior end (95—100%) Mere aeereite Wo orale A. coyi Quatrefages
Aspidosiphon (Akrikos), new subgenus
Diagnosis. —Aspidosiphon with com- pressed hooks not in rings, 1.e., either scat- tered and small (less than 30 wm), or absent; caudal shield absent or very diffuse; longi- tudinal muscle layer continuous. Not known to bore in coral or rock.
The name is from the Greek meaning “without rings.’’ The spelling is a literal transliteration according to the classical method.
Type species: Aspidosiphon albus Murina, 1967.
Aspidosiphon albus Murina, 1967
Aspidosiphon albus Murina, 1967a:1330- 1331, fig. 2 (1)-(3).—Stephen & Ed- monds, 1972:219-—221.—Cutler, 1973: 174-175.—Cutler & Cutler, 1980a:4.— Migotto & Ditadi, 1988:247-248.—Not Cutler et al., 1984:307.
Aspidosiphon hartmeyeri. —Wesenberg- Lund, 1957a:7—8; 1959a:197; 1959b:212.
Material examined. —ZIAS, type; USNM, cataloged as A. cumingii 066214-222 off Louisiana; material from Brazil (Cutler &
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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Cutler 1980a), U.S. east coast (Cutler 1973), and new Ivory Coast material from 5°N, 4°W, at 20 m; A. hartmeyeri, UZMK, Wes- enberg-Lund’s Niger worms.
Discussion.—This hookless species also seems to lack tentacles. The recorded body length ranges from 2 to 45 mm, the introvert is 3-5 times the trunk length, and the anal shield is fine grained with small furrows around the margin but without grooves. Often there is a median stripe made up of darker units. The nephridia are 50-75% of the trunk.
The Cutler et al. (1984) record was based
Fig. 5. Anal shields: A, Ungrooved type of A. elegans; B~C, Grooved type as in A. muelleri; D, Ventral cone- shaped units near the ventral margin present in some A. muelleri. Scale lines on A, B, C = 0.5 mm, D = 0.1 mm.
on one incomplete worm and was not a pos- itive identification. This should not be in- cluded within this species unless additional material is found in Japanese waters.
Distribution.—Cape Hatteras, northern Gulf of Mexico (unpublished), Cuba, Brazil continental shelf (10-123 m), and the east Atlantic from the Gulf of Guinea. It is the most common member of this genus on the Brazilian shelf.
Aspidosiphon mexicanus (Murina, 1967)
Golfingia mexicana Murina, 1967c:1333- 1334, fig. 3.
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Aspidosiphon mexicana Cutler et al., 1983: 673.
Aspidosiphon longirhyncus Cutler & Cutler, 1980a:4—6, figs. 4—5.
Material examined. —ZIAS, type; A. lon- girhyncus, AMNH, type (4022); other ma- terial of both species from the type locali- ties. Also, 12 unpublished specimens from the Azores (33°N, 16°W, 320 m) and seven from three stations off Florida and North Carolina (25—34°N, 85-190 m).
Discussion. —This species does not have the typical aspidosiphonid appearance in that the anal shield is very weakly devel- oped with scattered, ill-defined platelets. Also, the introvert is on an atypical angle (45—60°) with the main trunk axis. The cau- dal shield is almost nonexistent. The intro- vert is 4—5 times the trunk length, bearing scattered, small (less than 30 wm), uniden- tate compressed hooks. The nephridia are 35-75% of the trunk.
The decision to combine these two names was discussed in Cutler et al. (1983) as part ofa review ofthe taxon containing Murina’s species.
Distribution. —Southern Brazil, Cuba, and southeastern U.S. at shelf depths (80—200 m), and the Azores at 320 m.
Aspidosiphon thomassini Cutler & Cutler, 1979
Aspidosiphon thomassini Cutler & Cutler, 1979a:971-973, figs. 3-14.
Material examined. —MNHN, type ma- terial (AH 406-408).
Discussion. — This is another small (1.5- 7 mm) hookless species whose tentacles Gif present) have yet to be observed. The re- tractor muscles are fused for most of their length and the nephridia are around 50% of the trunk. The introvert is 2—4 times the trunk length. These have no caudal shield and the anal shield is very poorly devel- oped. This, together with the smaller angle between the trunk and introvert axis (40- 45°) make it possible to mistakenly identify this as a Nephasoma species.
841
Distribution. —Intertidal coral sands in Madagascar and French Polynesia.
Aspidosiphon venabulum Selenka, de Man & Bulow, 1883
Aspidosiphon venabulum Selenka et al., 1883:123, pl. 14, figs. 202—204.— Fischer, 1895:18; 1914a:68-69.— Wesenberg- Lund, 1957c:5—7; 1959a:196—-197; 1959c: 212.—Longhurst, 1958:85.—Stephen, 1960a:519.—Cutler, 1977a:148.
Aspidosiphon venabulus.—Stephen & Ed- monds, 1972:237.—Cutler & Cutler, 1979a:971.
Material examined. —MNHU, type (644); UZMK, Wesenberg-Lund’s Atlantide ma- terial; worms from Madagascar (Cutler & Cutler 1979a) and West Africa (Cutler 1977a).
Discussion. — This species lacks hooks and the anal shield is ungrooved, being made up of dark, pointed, conical units more widely scattered than in many species. It resembles the anterior end of certain Phascolion species that have large anterior papillae. The re- tractor muscles extend to the posterior end and under the anal shield, the longitudinal muscle layer splits into a few bundles. The nephridia are 60—95% of the trunk, the latter being reported from 5-30 mm.
Distribution. —Subtidal depths (10-55 m with one intertidal report and one at 960 m). Most records are off West Africa with one report from southern Madagascar.
Aspidosiphon zinni Cutler, 1969
Aspidosiphon zinni Cutler, 1969:209-211, fig. 1.—Cutler, 1973:176-178.—Cutler & Cutler, 1979a:968; 1980b:457; 1987a:73.
Material examined.—USNM, type ma- terial (38242, 38243); Atlantic Ocean ma- terial (Cutler & Cutler 1987a).
Distribution. — This small (most less than 5 mm), deep-water worm commonly lives in foraminiferan tubes. The anal shield is made up of very fine grained, pale units and the caudal shield is nonexistent. The small
842
(15-20 wm), scattered, unidentate, com- pressed hooks and introvert not longer than the trunk are unusual in this genus. The nephridia are less than 25% of the trunk.
Distribution. —Common in north Atlan- tic Ocean (1100-4400 m), two stations around 9°S off the Congo River (1535 and 2700 m), and one in the Mozambique Channel (25°S) at 132 m.
Subgenus Aspidosiphon (Aspidosiphon) —
Diagnosis. —Introvert with compressed hooks in rings, longitudinal muscle layer continuous except near anal shield. Most do not bore in coral or rock.
Aspidosiphon elegans (Chamisso & Eysenhardt, 1821)
Sternaspis elegans Chamisso & Eysenhardt, 1821:351-352, pl. 24, figs. Sa—e.
Sipunculus elegans.—de Blainville, 1827, pl. 26, fig. 2.
Loxosiphon elegans. —Diesing, 1851:70.— Quatrefages, 1865:605.
Phascolosoma (Aspidosiphon) elegans. — Grube, 1868a:645-647.
Aspidosiphon elegans. —Selenka et al., 1883: 124-126.—Shipley, 1898:471; 1899b: 153.—Whitelegge, 1899:393.—Sluiter, 1891:116; 1902:19.—Hérubel, 1904: 564.—Lanchester, 1905a:33; 1905b:40.— Fischer, 1914b:14.— Gravely, 1927:87.— Sato, 1935:316; 1939:426—-427.— Wes- enberg-Lung, 1954:10-11; 1957a:198- 199; 1957b:7-8; 1959c:68.—Cutler, 1977b:154.—Cutler & Cutler, 1979a: 968.—Edmonds, 1980:44—46.— Cutler et al., 1984:304.—Migotto & Ditadi, 1988: 248-250.
Aspidosiphon elegans elegans. —Stephen & Edmonds, 1972:223.
Aspidosiphon elegans var. yapense Sato, 1935:316-318, pl. 4, fig. 18, text-figs. 12— IS
Aspidosiphon elegans yapensis. —Stephen & Edmonds, 1972:224.
Aspidosiphon brocki Augener, 1903:328-
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330, figs. 9-13.—Murina, 1967b:42.— Rice, 1970:1618-1620; 1975:44-45.— Stephen & Edmonds, 1972:221.—Rice & Macintyre, 1979:311-319.
Aspidosiphon carolinus Sato, 1935:318-319, pl. 4, fig. 19, text-figs. 16—17.—Stephens & Edmonds, 1972:222.—Cutler & Cutler, 1981:77-78.
Aspidosiphon cylindricus. —Sluiter, 1902: 18-19.
Aspidosiphon exilis Sluiter, 1886:497, pl. 3, figs. 11-12; 1891:116; 1902:18.—Leroy, 1942:39-40.—Stephen & Edmonds, 1972: 224—225.—Edmonds, 1980:44—46.
Aspidosiphon homomyarium Johnson, 1964: 332-334, pl. 8.
Aspidosiphon homomyarius. —Stephen & Edmonds, 1972:227.
Aspidosiphon ravus Sluiter, 1886:495-—496, pl. 3, figs. 9-10; 1891:116; 1902:18.— Shipley, 1899a:56; 1899b:153.—Stephen & Edmonds, 1972:234.
Aspidosiphon spinalis ikeda, 1904:47—49, text-figs. 12, 81-85; 1924:37.—Sato, 1939:428.—Stephen & Edmonds, 1972: 234—235.—Cutler & Cutler, 1981:79-81.
Aspidosiphon spinosus Sluiter, 1902:28, pl. 2, figs. 17-19.—Stephen & Edmonds, 197222355.
Material examined.—ZMUA, Sluiter’s from Indonesia (V. Si. 3); worms from the western Pacific Ocean (Cutler et al. 1984), 1988 collections from the southern Carib- bean, a few from Brazil (Migotto & Ditadi 1988); A. brocki, MNHU, type material (6954-5); USNM, Caribbean material iden- tified by M. Rice (USNM 48924-5); A. cy- lindricus, ZMUA, two of Sluiter’s (26.8); A. exilis, BMNH and ZMUA (V. Si. 4), parts of type material in both places, all of it badly dried out; A. homomyarius, RSME, two worms from India presumably deposited by Johnson (1965.32.1); A. ravus, ZMUA (V. Si. 13) and BMNH, type material.
Discussion. —Like A. (Paraspidosiphon) steenstrupii in its subgenus, A. elegans is the most common and widespread tropical
VOLUME 102, NUMBER 4
member of this subgenus with many junior synonyms. Additionally, both species have ungrooved anal shields, bidentate com- pressed hooks in rings, and dark scattered hooks. Edmonds (1980:44—45) presented a detailed description of A. elegans and some interesting comments on this complex of related (or identical) taxa. What he illus- trates as introvert spines (his figs. 77-78) are, by our definition, conical hooks. The variation in the shape (real and due to ori- entation on slide) of the bidentate hooks has led to the creation of different species. Fig- ure 6 illustrates some of this variation (de- gree the hook is bent and sharpness of the point) that we now know to be within-deme variation. Six to twelve short stubby nuchal tentacles are present. The caudal shield is weakly developed, barely discernable in many worms.
Internally the longitudinal muscle layer may subdivide in the area of the anal shield. Less than 20% of the worms dissected have a caecum and a fixing muscle was seen in only 4% of the worms dissected. The ne- phridia open at the level of the anus or just posterior to it. Table 2 shows our obser- vations on the nephridia and retractor origins. The gut has the normal helical coil, but about half the worms show a degree of looseness in part of the coil.
The putative species A. carolinus and A. spinalis were reduced to junior synonyms by Cutler & Cutler (1981) and A. exilis by Edmonds (1980). We reaffirm those deci- sions.
Aspidosiphon brocki (Augener 1903) is a name used by only two authors since it was described: Murina (1967b) and Rice (1970, 1975). Neither of these authors has used the name A. elegans in their writings. Augener’s paper was the first of only two articles he wrote about this phylum, and he made no reference to any other member of this sub- genus nor was there a differential diagnosis. The original report was from Malaya but the four subsequent records are all Carib- bean, and all of this latter material is less
843
Fig. 6. Variation in shape of bidentate compressed hooks within a single specimen of A. elegans. Note difference in degree of bend and sharpness of point. Scale line = 20 um.
than 10 mm long. Our recent Caribbean collections (340 worms) ranged from 4 to 22 mm long. The pictures of hooks in Ste- phen & Edmonds (1972:230, fig. 27B, F, G) illustrate the extremes of a continuum and could be misleading. This much variation occurs within demes. Comparison of the type, Rice’s material, and our own Carib- bean and Pacific material convinces us that this putative species 1s conspecific with A. elegans.
Rice (1970) reported asexual reproduc- tion by budding in A. brocki, something not recorded elsewhere in this phylum. Our Ca- ribbean material showed this (even in 4 mm worms), but it is also present in our collec- tions from Majuro, Marshall Islands, and in French Polynesian worms collected by Peyrot-Clausade.
Sluiter’s (1902) Siboga report included A. cylindricus, Horst, but not A. elegans. Our examination of his material showed no dif- ferences from A. elegans.
When Johnson (1964) described 4. ho- momyarlus, he did not include a differential diagnosis except to contrast it with the other new species in that paper, so we do not know how he thought it differed from A. elegans.
844
He seemed to think that the dorsal array of nuchal tentacles was unique in this genus. Our examination of his two worms in Edin- burgh confirmed our analysis of his article, i.e., meaningful differences from A. elegans are lacking.
Aspidosiphon ravus was described by Slui- ter (1886) as having unidentate hooks. Our reexamination of the material (the Amster- dam worms are in good condition but the London worm is not) revealed bidentate compressed and conical hooks. The intro- vert is retracted in all three worms. This material is clearly conspecific.
Aspidosiphon spinosus Sluiter, 1902, was based on a single worm that has subse- quently been lost. His description is indis- tinguishable from A. elegans. He compared it to A. exilis differentiating it by the form of the hooks and a few other features. Plac- ing this name on the list of incertae sedis was considered since it cannot be examined (the hypothesis cannot be tested) but re- ducing it to ajunior synonym of this species is a rational alternative given what we now know about within-deme variation.
The subspecies A. elegans yapensis, which Sato (1935) described as a variety, was dif- ferentiated on the basis of hook morphol- ogy: sharply pointed apex, not blunt. The within-deme variation we find makes it clear that this subspecies lacks biological signif- icance.
Distribution.— Widespread and common in the Indian and western Pacific Ocean (from southern Japan to northern Australia out to Hawaii); the Red Sea and Israel. In the Caribbean from northern Brazil to the Florida keys and Bermuda.
Aspidosiphon exiguus Edmonds, 1974
Aspidosiphon exiguus Edmonds, 1974:187— 192, figs. 1-7.
Material examined.—BMNH, holotype (197 5.22.1):
Discussion. —The largest worm reported is less than 4.5 mm long, but it does bear bidentate hooks in rings as well as a few
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scattered unidentate compressed hooks. The introvert is very long (2—5 times the trunk) and no tentacles or gametes have yet been observed. The anal shield consists of small, pale units with no grooves. At the anterior/ ventral border 1-4 cone-like papillae are present. It is similar to A. albus but has hooks and shorter nephridia (25-33%). In the anterior 10% of the trunk the longitu- dinal muscle layer is divided into bands vis- ible through the body wall. Aspidosiphon (Paraspidosiphon) parvulus is common in this area and is similar in several ways de- spite having longitudinal muscle bands (weakly developed).
Distribution. —Cuba, intertidal, intersti- tial.
Aspidosiphon gosnoldi Cutler, 1981
Aspidosiphon gosnoldi Cutler, 1981:445- 449, figs. 1-4.—Migotto & Ditadi, 1988: 253-254.
Aspidosiphon spinalis.—Cutler, 1973:175- 176.—Cutler & Cutler, 1979b:107.
Material examined. —USNM, type (61624-5) and additional material from the western Atlantic Ocean (Cutler 1981).
Discussion.—The anal shield is com- posed of randomly arranged flat units of relatively uniform size. The borders are usually distinct but dark skin papillae may be present at the anterior end of the trunk. The introvert is 1.5—3 times the trunk length and bears distal rings of 20-30 um, biden- tate hooks (the secondary tooth may be small). Scattered, pale, pyramidal hooks cover much of the proximal part of the in- trovert. This 1s in the group of species where the longitudinal musculature commonly splits into irregular bundles under the anal shield and the retractors originate from the caudal shield. The intestine forms a normal helical coil, and the nephridia are 50-90% of the trunk length.
Distribution. —Shelf waters (7-185 m) from Cape Hatteras to Florida and Brazil (to 23°S) living in gastropod shells.
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Aspidosiphon gracilis gracilis (Baird, 1868)
Pseudoaspidosiphon gracile Baird, 1868:103, pl. 10, figs. 1, la.
Aspidosiphon gracilis —Selenka et al., 1883: 122-123.—Sluiter, 1902:17.—Augener, 1903:319-321.—Heérubel, 1904:564.— Rice & Stephen, 1970:69.—Stephen & Edmonds, 1972:225—226.— Gibbs, 1978: 85.—Edmonds, 1980:46-47.
Material examined.—BMNH, syntypes (43.5.15.58a/b); MNHN, two of Hérubel’s specimens (V20).
Discussion. —This species has an under- developed anal shield composed of non- contiguous brown papillae surrounded by darker platelets. These units are arranged in irregular longitudinal rows. The introvert comes off at about a 60° angle, is up to 1.5 times the trunk length, and the slender trunk (up to 15 times the width) is coarsely pap- illated all over. The assertion that bidentate hooks are present (Stephen & Edmonds 1972) must be a flawed translation since we cannot find any other reference to them. The unidentate hooks are in rings, are broader than high (up to 40 wm), and are followed by a proximal area of pyramidal hooks. The retractor muscles originate very near the posterior end and the nephridia are as long as the trunk.
Hérubel (1904) had three worms from the Gulf of Aden that look different but are damaged and desiccated such that we can- not confirm or refute his identification.
Distribution. — Australia, Indonesia, Phil- ippines, Gulf of Aden and an unpublished record from the Andaman Islands.
Aspidosiphon gracilis schnehageni Fischer, 1913, new status
Aspidosiphon schnehageni Fischer, 1913:99- 100, figs. 4-6; 1914b:15.—Wesenberg- Lund, 1955:13.—Ditadi, 1975:200-—202.
Paraspidosiphon schnehageni. —Stephen & Edmonds, 1972:252.
845
Material examined. —ZMUH, type spec- imen (V2127).
Discussion. — The type specimen is 1n very poor condition, missing, among other parts, the distal end of the introvert and the re- tractor muscles. Ditadi (1975) redescribed this taxon, but his material cannot be lo- cated at the Los Angeles County Museum or the Hancock Foundation where it had been deposited. The anal shield was de- scribed as furrowed by Fischer but as ran- domly arranged plates by Ditadi. This is another situation where the units may sometimes appear arranged in rows giving an impression of indistinct ridges and fur- rows. The longitudinal musculature is par- tially separated into 10-14 anastomosing bundles in the anterior part but is contin- uous elsewhere. In this regard, it is one of the borderline taxa not having distinct bun- dies, and, therefore, we have moved it into this subgenus.
The decision to reduce this to a subspecies rather than a junior synonym was based on ill-defined differences from the nominate form; habitat (mollusc shells), hook shape (more triangular), nephridia (shorter, 33- 50%), trunk size and shape (length less than 8 times the width), and longitudinal muscle layer (splitting extends beyond the anal shield). These are not clear or distinct dif- ferences by themselves, but if one adds the geographical gap between the populations (most of the Pacific Ocean), this status may be appropriate pending additional material for better comparison. The west coast of Central and South America are very poorly represented in curated, accessible collec- tions.
Distribution. —Chile and Pacific coast of Guatemala.
Aspidosiphon misakiensis Ikeda, 1904
Aspidosiphon misakiensis Ikeda, 1904:41- 43, text-figs. 9, 68—72.—Sato, 1939:428.— Stephen & Edmonds, 1972:229-231.— Cutler & Cutler, 1981:78—79.—Cutler et al., 1984:305-—306.
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Aspidosiphon hartmeyeri Fischer, 1919:281- 282, text-figs. 1-3; 1926:204—205.—Ed- monds, 1956:306—307; 1980:47.—Muri- na, 1967c:1332.—Stephen & Edmonds, 1972:226-227.—Cutler, 1977a:147-148. Not Wesenberg-Lund, 1957a:7—-8; 1959a: 197; 1959b:212.
Aspidosiphon gerouldi ten Broeke, 1925:93, text-figs. 23-25.—Stephen & Edmonds, 1972:225.—Cutler & Cutler, 1979b:106—- 107.—Migotto & Ditadi, 1988:251-253.
Aspidosiphon speculator. —Saiz Salinas, 1986a:11-14.
Material examined.—USNM, paraneo- type; other material from the type locality (Cutler & Cutler 1981); A. gerouldi, ZMUA, type (V. Si. 7), material from Azores (Cutler & Cutler 1987), and Brazil (Migotto & Di- tadi 1988); A. hartmeyeri, ZMUH (V8913- 14) and MNHU (6036), co-types; UZMK, Wesenberg-Lund’s Niger material (=A. al- bus), A. speculator, three specimens iden- tified by Saiz Salinas.
Discussion.—The anal shield is com- posed of closely packed, irregular, granular units, but it has borders that are not sharply defined, i.e., widely spaced, square blocks of shield material are around the anterior quarter of the trunk. The caudal shield is granular but does have vague radial grooves present. Bidentate hooks are present (25—40 um) in distal rings, then proximally the hooks are scattered, unidentate com- pressed, 25-60 um tall (Fig. 1B). The sec- ondary tooth has normal dimensions on distal hooks but becomes very small in proximal ones. The introvert is up to three times the trunk length (largest known worm is 25 mm). The longitudinal muscle layer exhibits some fractures/splits in some in- dividuals, and the gut coils are ill defined or only loosely wound. In 6-7 mm worms the intestine has a few loose folds, is an- chored at the posterior end, then has a straight tube to the anus. Sheets of connec- tive tissue link the ascending and descend-
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ing loops. The nephridia are 50-100% of the trunk and the retractors originate very close to the caudal shield.
When Fischer (1919) described A. hart- meyeri from Australia, he made no refer- ence to Ikeda’s A. misakiensis even though it is clear he knew of Ikeda’s paper. Stephen & Edmonds’ (1972) key uses the nephridia being mostly free to separate it from Ikeda’s. Our examination of five type specimens showed considerable variation (up to 80% attached and length up to 100% of the trunk, not 0% and 50% as stated). The anal shield was described as having 5-6 flat grooves. These are not grooves but irregular units arranged in indistinct rows in a few worms (see also Edmonds 1956:306). Wesenberg- Lund’s four worms from West Africa are clearly part of the A. albus population. She never recorded hooks, and her drawings of one, plus our examination of another, con- firm this.
The decision to reduce the status of A. gerouldi came after examination of unpub- lished material from the Azores and com- parison to the Japanese worms. The original description was based on a single worm, but several hundred have been subsequently collected. The bidentate hooks in the Azores population are at the small end of the range (25-30 um) as is the introvert (only up to twice the trunk in worms up to 20 mm long).
Saiz Salinas (1986a) used the name A. speculator for 18 worms off Spain and the Canary Islands. This population is partic- ularly troublesome as indicated by his plac- ing it in A. (Paraspidosiphon). The longi- tudinal musculature is neither continuous nor divided into distinct bands. It varies from worm to worm in degree, but we in- terpret it to be continuous with fracturing in the anterior part, especially on the dorsal side. In other respects (hooks, shield, gut) his worms fit A. misakiensis better than any other taxon.
Distribution. —In the Pacific from both sides of central Japan at 1-50 m depth, South
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and West Australia, and Kermadec Island. In the eastern Atlantic from the Azores, Cape Verde, and Canary Islands to the Gulf of Guinea, at depths down to 75 m, and the Spanish Mediterranean. In the western At- lantic from Brazil (14—-16°N), Haiti, and Cuba.
Aspidosiphon muelleri Diesing, 1851
Aspidosiphon muelleri Diesing, 1851:68.— Quatrefages, 1865:609-610.—Schmidt, 1865:56-66.— Baird, 1868:101.—Selen- ka et al., 1883:120-121.—Fischer, 1895: 18; 1914a:69-70; 1914b:13-14; 1922a:22- 23; 1925:25—26.—Sluiter, 1900:14; 1902: 18; 1912:19.—Hérubel, 1904:564.— Southern, 1912:31-34.—J. Fischer, 1914: 105-106.—Ikeda, 1924:38.—Stephen, 1934:173; 1941:257; 1958:133-134; 1960a:518-519; 1960b:22—23.—Steuer, 1936:5; 1939:2.—Sato, 1939:428.— Chapman, 1955:351.— Wesenberg-Lund, 1957a:4—-5; 1957b:197-198; 1959a:194— 196; 1959c:68.—Longhurst, 1958:1.— Stephen & Edmonds, 1972:231-233.— Zavodnik & Murina, 1975:127; 1976:81- 82.—Cutler, 1977a:148.—Gibbs, 1977: 30-31.—Cutler & Cutler, 1979b:107; 1987a:73.—Ocharan, 1980:114-115.— Cutler et al., 1984:306-307.—Saiz Sali- nas, 1984:177-178; 1986a:9-11.
Sipunculus scutatus J. Miller, 1844:166—-168 (not scutatum J. Miller, 1843).—Kefer- stein, 1867:52.—Selenka et al., 1883:120.
Phascolosoma scutatum. —Krohn, 1851: 371.—Selenka et al., 1883:120.
Aspidosiphon clavatus. —Diesing, 1851: 68.—Cuénot, 1922:12-—13.—Heérubel, 1924:111.—Leroy, 1936:426.—Akesson, 1958:206.—Voss-Foucart et al., 1977: N35:
Pseudaspidosiphon clavatum.—Baird, 1868: 103.
Sipunculus cochlearius Valenciennes, 1854: 640.—Saiz Salinas, 1986b:554.
Lesinia farcimen Schmidt, 1854:2.—Selen- ka et al., 1883:120.
847
Aspidosiphon eremita Diesing, 1859:768 (not Phascolosoma eremita Sars, 1851).
Phascolosoma radiata Alder, 1860:75.— Southern, 1913:32.
Sipunculus heterocyathi McDonald, 1862: 78-8 1.—Saiz Salinas, 1986b:554.
Aspidosiphon jukesii Baird, 1873:97.— Rice & Stephen, 1970:68-69.—Stephen & Ed- monds, 1972:228.—Cutler & Cutler, 1979a:969-970.—Edmonds, 1980:49.— Saiz Salinas, 1986b:551.
Aspidosiphon mirabilis Théel, 1875:17; 1905:91-—92.—Selenka et al., 1883:121.— Southern, 1913:31-33.
Aspidosiphon armatum Danielssen & Ko- ren, 1880:464; 1881:64.—Selenka et al., 1883:124.—Théel, 1905:93.—Southern, 1913:31-33.
Aspidosiphon tortus Selenka et al., 1883:119- 120, pl. 14, figs. 196-201.—Hérubel, 1904:564.—Fischer, 1923:21-22.—Ste- phen & Edmonds, 1972:236-237.— Gibbs, 1978:85.
Aspidosiphon heteropsammiarum Bovier, 1894:98.—Saiz Salinas, 1986b:555-557.
Aspidosiphon michelini Bovier, 1894:98.— Saiz Salinas, 1986b:557-559.
Aspidosiphon corallicola Sluiter, 1902: 19-22.—Shipley, 1903:169-171.—Ste- phen & Robertson, 1952:441-442.—Cu- tler, 1965:58.
Aspidosiphon imbellis Sluiter, 1902:29, pl. 2, fig. 20.—Stephen & Edmonds, 1972: 227-228.
Aspidosiphon inquilinus Sluiter, 1902:29-30, pl. 2, figs. 21-22.—Stephen & Edmonds, 1972:227.—Edmonds, 1980:47-49.
Aspidosiphon exhaustum Sluiter, 1912:20- 21, pl. 1, fig. 11.—Murina, 1971:78.
Aspidosiphon exhaustus.—Stephen & Ed- monds, 1972:224.—Murina, 1972:295- 296; 1978:120.—Cutler & Cutler, 1979a: 969; 1980a:4.—Edmonds, 1980:46.— Cutler et al., 1984:305.
Aspidosiphon exhaustus mirus Murina, 1974:1715-1716, fig. 2.
848
Aspidosiphon pygmaeus Fischer, 1921:45- 47, text-figs. 1—-7.—Murina, 1967a:54; 1971:78.
Paraspidosiphon pygmaeus. —Stephen & Edmonds, 1972:251-—252.
Aspidosiphon kovaleskii Murina, 1964b:5 1— 55, figs. 1-5; 1970:66.—Stephen & Ed- monds, 1972:229.—Zavodnik & Murina, 1975:127.—Cutler & Cutler, 1979a:970- 971. |
Aspidosiphon hispitrofus LiGreci, 1980:123- 134, figs. 1-4.
Material examined.—ZMUA, Sluiter’s Indonesian worm (V. Si. 253); material from the Azores, 1-600 m depth collected by Zi- browius and CENTOB; Celtic Sea from P. Gibbs; French Mediterranean from Voss- Fouchart; Japan (Cutler, Cutler & Nuishi- kawa 1984); A. armatus, ZMUB, type (1745); A. corallicola, ZMUA, types (V. Si. 1); A. exhaustus, MOMV, type; Cutlers’ Brazilian, Japanese and Indian Ocean worms; A. imbellis, ZMUA, type (V. Si. 25/5); A. inquilinis, ZMUA, type (V. Si. 25/6); A. jukesii, BMNH, type (1965.25.3); Indian Ocean material (Cutler & Cutler 1979a); A. kovaleskii, ZIAS, type; Indian Ocean worms (Cutler & Cutler 1979a); A. mirabilis, ZMUB, type (15957); A. pyg- maeus, NHRS, type specimens (279).
Discussion.—This species is the wide- spread, eurytopic, polymorphic member of this genus and in this way is comparable to Golfingia margaritacea, Phascolion strom- bus, Sipunculus nudus, etc. Each genus seems to have one such species with a long list of junior synonyms and a morphology difficult to define with precision. Stephen & Ed- monds (1972:232—233) discuss some of the early confusion about the proper name for this taxon. Most records are from the north Atlantic Ocean and Mediterranean Sea from abandoned mollusc shells. When biologists found similar worms from different parts of the world or from different habitats, new names were put forth.
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Our present concept of A. muelleri has an anal shield made up of very small units ar- ranged into variable sized plates, partially separated by longitudinal furrows dorsally, in the midsection by transverse furrows, and made up of raised wart-like or cone-shaped units ventrally (Fig. SC). The possibility that two species exist, one with pointed, cone- shaped spines on the ventral part of the anal shield and the other with only flat wart-like units, was considered. It is our conclusion that this species has the capability to re- spond to some environmental stimulus (e.g., pressure, temperature, or host shell shape), or that random allelic frequency shifts can occur, to produce anal shields with ventral units varying in degree of cone develop- ment.
The trunk may be straight or coiled de- pending on its habitat. The introvert is 1-3 times the trunk length, and there are 6-12 small nuchal tentacles. Hook morphology has been a long standing point of confusion (see Southern 1913, Stephen & Edmonds 1972:233). Our analysis of within-deme variation of compressed hooks clearly sug- gests that A. muelleri has the genetic poten- tial for producing only unidentate hooks or only bidentate hooks or some of both on a single worm. Proximal to the rings (covering about one-third of the introvert), the scat- tered hooks quickly change from com- pressed unidentate to pyramidal. The SEM photomicrographs reveal the existence of small comb-like structures at the posterior base of the compressed hooks (Fig. 1A).
Internally the pair of introvert retractor muscles originates from the edge of the cau- dal shield. Under the anal shield the lon- gitudinal muscle layer divides into separate bands. The nephridia open at or just pos- terior to the anus, are from 25—100% of the trunk length, and the gut forms a regular helical coil. The rectal caecum and fixing muscle are only present in some individu- als.
Aspidosiphon jukesii (Baird 1873) and its several junior synonyms must be moved into this species. When one considers each char-
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acter and looks not just at one author’s per- ception but at the collected writings and the many specimens at hand, the presumed ““‘bridgeless gap”’ ceases to exist. If a young worm occupies a gastropod shell in an area where solitary corals live, it is possible that this shell will be settled upon by a coral larva establishing a mutualistic relationship. This fact does not make the worm a different species. While it is true that most A. jukesii had anal shields with some number of cone- shaped spines, this is not universal in, or restricted to, any particular population.
The type of A. tortus (Selenka et al. 1883) is not with their other type specimens in the Berlin museum. They had one 25 mm worm with a longitudinal muscle layer continuous except for the anterior dorsal area where it had some separation without being clearly separate. This worm also had just one ne- phridium, a condition Fischer (1923) as- serted was only an anomaly when he de- scribed his two worms. This has been presumed to be a separate species because it lacks unidentate hooks, now not a valid basis for separation, even if true. Their con- cept of spines (Stacheln) may well overlap with our idea of unidentate hooks (see their fig. 200, 201). Hérubel (1904) recorded one worm from Djibouti with no comments. When Gibbs (1978) listed this species from the Great Barrier Reef he did indicate some doubt with the (cf.) notation. Our analysis of the literature (including figures) con- vinces us that A. tortus is a junior synonym of A. muelleri.
When Sluiter (1902) erected A. imbellis he asserted that this single 13 mm worm with retracted introvert lacked hooks. Our reexamination showed this to be an error. Rings of 25-35 wm hooks exist, some with a small secondary point. The spindle muscle is attached posteriorly and it is clearly con- specific with A. muelleri.
Aspidosiphon inquilinis was also based on a single specimen that Sluiter (1902) differ- entiated from A. muelleri on the basis of hook and skin body morphology plus a mis- understanding of retractor origins. His spec-
849
imen had lived in a scaphopod shell and the anal shield has a peculiar slant. The intro- vert skin is folded back over the ventral edge of the anal shield. Edmonds (1980) pointed out that Sluiter had overlooked the small cone-shaped units along the shield margin just as Edmonds’ four worms exhibited. These five worms have only compressed and pyramidal unidentate hooks. They are con- specific with A. muelleri.
In 1912 Sluiter erected A. exhaustus for a single 17 mm worm taken from a scapho- pod shell in the east Atlantic on the slope off Morocco. No differential diagnosis was presented and examination of the type shows it to be like the other A. muelleri of this region with unidentate hooks. The name went unused for 59 years, after which Mu- rina used it four times followed by the Cut- lers who used it three times for cold water worms from diverse locations. This usage was predicated on the false assumption that the real A. muelleri must bear some biden- tate hooks.
The name A. pygmaeus has been used by two authors. Fischer (1921) asserted that the longitudinal muscle layer consisted of bands anastomosing so frequently that he could not count them. He did say that these were most numerous at the anterior end and ran together at the posterior end. This statement caused Stephen & Edmonds (1972) to place the species in the subgenus 4. (Paraspido- siphon). When we examined the type ma- terial (four of his seven worms in good con- dition with introverts out and dorsal tentacles showing), we saw a continuous muscle layer with some fracturing under the anal shield, a condition common to many worms in this taxon. The coast of Chile is not a common location for A. muelleri, but Fischer’s worms do fit this construct as do Murina’s (1967a, 1971). Her second record was a repeat of the first for two worms from 150 m in the Gulf of Aden.
Aspidosiphon kovaleskii Murina, 1964, was presumed to differ from A. muelleri be- cause it lacked bidentate hooks. Several populations (see Fischer 1895, Southern
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C
Fig. 7. A. spiralis: A-B, Bidentate and unidentate compressed hooks with broad bases (both 20 um high); C, Anal shields with irregular squares in offset rows. Scale line in C = 2 mm.
1913, Gibbs 1977) of A. muelleri from the eastern Atlantic are reported to lack biden- tate hooks. By Murina’s definition these should be A. kovaleskii. Recently Saiz Sa- linas (1984:177-180) provided a detailed account of these two taxa and proposed that in both species both kinds of compressed hooks exist. He suggested that A. kovaleskii, if valid, might differ by having cone-shaped spines on the anal shield. He did express strong reservations about the validity of these taxa as separate and distinct biological entities; we conclude that they are not. Aspidosiphon hispitrofus LiGreci, 1980, was based on a series of worms from Sicily. None had their introverts all the way ex- tended, and he reported these to have no tentacles and only unidentate hooks. He did not mention the name A. muelleri in his
paper but compared his material to A. cla- vatus. It is clear that his material is conspe- cific with A. muelleri and his unfamiliarity with the phylum led him to misinterpret the distal end of the introvert.
Distribution. —Common in the northeast Atlantic from Norway through the British Isles, the Azores and Canary Islands and West Africa (to 10°N). It extends through the Mediterranean and Red Sea into the Gulf of Aden and along the coast of east Africa to Madagascar and South Africa. The rec- ords then skip to Ceylon and sparse reports up to Japan through Thailand, Vietnam, In- donesia, and down to Australia, New Guinea, and Kermadec Island. Most of the Pacific Ocean is unpopulated by this species. One record from Juan Fernandez Island off Chile (33°S) and one from southern Brazil
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(34°S) exist, but nothing else from the American hemisphere. This southern hemi- sphere distribution is intriguing. Through- out most of its range this species inhabits shelf depths (10-300 m), but there are sev- eral records down to 1000 m with the deep- est at 2930 m.
Aspidosiphon spiralis Sluiter, 1902
Aspidosiphon spiralis Sluiter, 1902:25-26, pl. 2, figs. 9-13.—Shipley, 1903:171.— Stephen & Edmonds, 1972:236.
Material examined. —ZMUA, type spec- imens (V. Si. 171-174).
Discussion. —While Sluiter said this species has unidentate hooks, rings of small bidentate hooks are also present. These are about 20 um high and the secondary point is not large (Fig. 7A, B). The unidentate hooks are about the same size and in rings. A gradual transition may exist during growth as there are some intermediate hooks. The anal shield does not have regular furrows, but neither is it made up of uniform gran- ules. It is divided into irregular squares overlain with horny protein (Fig. 7C). The retractor muscles originate from the pos- terior end of the trunk. This species does not have a firm foundation (four worms), but we propose no change in its status at this time.
The three worms Shipley identified can- not be located, and we question whether he was looking at the same entity. His descrip- tion of the agglutinated sand packed around the opening of the gastropod shell is much more typical of Phascolion species. Also, he provided no morphological information (other than color) or station data.
Distribution. —Indonesia, gastropod shells, 14—91 m.
Subgenus Aspidosiphon (Paraspidosiphon)
Diagnosis. —Introvert with compressed hooks in rings, longitudinal muscle layer di- vided into anastomosing bands. All bore in coral or rock.
851
Aspidosiphon coyi Quatrefages, 1865
Aspidosiphon coyi Quatrefages, 1865:608— 609 (partim).— Baird, 1868:101.—Ste- phen & Edmonds, 1972:340.—Saiz Sali- nas, 1984:42—49.
Phascolosoma truncatum Keferstein, 1867: 50-53, pl. 6, figs. 15-18.
Aspidosiphon truncatus. —Selenka et al., 1883:118-119.—Selenka, 1885:20.— Sluiter, 1898:444; 1902:17.—Shipley, 1899b:154; 1902:132.—Herubel, 1904: 564.—Ikeda, 1904:38-39.— Lanchester, 1905a:34.— Fischer, 1914b:15.—Ben- ham, 1912:136.—Hammerstein, 1915: 2.—Sato, 1939:428.—Cutler & Cutler, 1979a:976.—Cutler et al., 1984:309-310.
Paraspidosiphon truncatus. —Stephen & Edmonds, 1972:258.
Material examined. —-MNHN, type ma- terial (V20); A. truncatus, ZMUA, Sluiter’s Indonesian worm (V. Si. 25.1); from Mo- zambique (Cutler & Cutler 1979a) and Ja- pan (Cutler et al. 1984); USNM, unpub- lished material from the Gulf of Panama (21477), and Gulf of California, Puerto Pen- asco, Mexico (26443).
Discussion. — A. coyi is one of two species in this subgenus with a grooved anal shield. The primary distinction from A. /aevis is the presence of distal rings of bidentate hooks 25-35 um tall. However, the second- ary tooth on these is very small and not consistently present. There can be uniden- tate compressed hooks as well as thin py- ramidal hooks, and the presence of both has led to some confusion in the literature. The skin papillae towards the two ends of the trunk are large and rugose. The longitudinal muscle layer can exhibit much anastomos- ing and is not always clearly banded. The retractors originate from the posterior end (95—100%), the spindle muscle may bifur- cate near the anus, and the wing muscle is well developed extending down to near the ventral nerve cord. The nephridia are 40- 95% of the trunk length and attached for most of their length. This has not been a
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well defined species and the size of the data base is small. Striking similarities to A. (As- pidosiphon) muelleri exist (e.g., shield, re- tractors, hooks), and show another example of the less than clear boundary between the two subgenera.
The careful redescription of this species by Saiz Salinas (1984) brought the name A. coyi back from its earlier dubious status (Stephen & Edmonds 1972) and these taxa are clearly conspecific. The location of Qua- trefages’ specimens is not clear but is prob- ably somewhere in the Indian Ocean.
Distribution. —Several locations in the western Indian Ocean and the western Pa- cific from Japan through Okinawa, Philip- pines, Indonesia, and Kermadec Islands. Two Latin American records of uncertain location exist: The type of A. truncatus from Panama (east or west coast?) and a single 3 mm worm from San Salvador (Bahamas or Galapagos?). Eastern Pacific locations for these two are supported by the USNM ma- terial from the Gulf of Panama and the Gulf of California. The former was collected in 1866 and may be part of Keferstein’s type collection.
Aspidosiphon fischeri ten Broeke, 1925
Aspidosiphon fischeri ten Broeke, 1925:92- 93, figs. 21-22.— Migotto & Ditadi, 1988: 250-251.
Paraspidosiphon fischeri fischeri. —Stephen & Edmonds, 1972:244-245.—Amor, 1975:118-119.—Rice, 1975:38-44.— Rice & Macintyre, 1979:311-319.
Aspidosiphon fischeri cubanus Murina, 1967b:39-42, figs. 5-7, 1967c:1331.
Paraspidosiphon fischeri cubanus. —Ste- phen & Edmonds, 1972:245.
Material examined.—ZMUA, type ma- terial (V. Si. 5); USNM, Venezuelan spec- imens identified by M. Rice (100901); un- published specimens labeled A. truncatus (20726, 20725, 20727, 20836); our 1988 collections from the Caribbean.
Discussion. — The body wall of these small
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worms (trunks 4—16 mm) is smooth, thin, and white, but the longitudinal muscles bands cannot be easily seen through it, mak- ing it easy to misplace these worms in A. (Aspidosiphon) during preliminary sorting, 1.e., they are similar to A. A. misakiensis or A. A. gosnoldi. Few, thin, ill-defined muscle
bands exist that anastomose frequently and
seem to fuse in the posterior part of the trunk in some worms. The long introvert (one or two times the trunk) bears rings of compressed, 18-27 um, bidentate hooks. The proximal rings may include hooks with a very small secondary point and unidentate hooks mixed together or a few rings of just unidentate hooks. Following these are scat- tered, pale, pyramidal hooks 15-50 um tall. The retractor muscles are thin and originate at or very near the posterior end of the trunk (95-100%). The nephridia are short; 33-50% of the trunk length.
When ten Broeke described this species she provided no differential diagnosis, and the differences from other species, e.g., A. parvulus, or those mentioned above, are not always distinct. One might make a case for this being a subspecies or an incipient or sibling species, but we propose no change now. Migotto & Ditadi (1988) question the basis for the subspecies A. fischeri cubanus Murina, 1967, and we agree that it only rep- resents some of the variation within the species.
Distribution.—Numerous southern Ca- ribbean locations from Cuba to Sao Paulo, Brazil, in shallow coral rock. Also from the Pacific coast of Panama, Ecuador, plus James and Hood Islands, Galapagos.
Aspidosiphon laevis Quatrefages, 1865
Aspidosiphon laeve Quatrefages, 1865: 609.—Baird, 1868:102.— Vaillant, 1871: 272-273; 1875: pl. 4, figs. C1-4.—De- Rochebrune, 1881:233.—Saiz Salinas, 1984:55-62.
Aspidosiphon laevis. —Stephen & Edmonds, 1972:340.
Aspidosiphon cumingii Baird, 1868:102, pl.
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11, fig. 2.—Selenka et al., 1883:113- 115.—Fischer, 1892:85; 1922b:12.— Collin, 1892:177.—Sluiter, 1898:444; 1902:17.—Augener, 1903:321-322.— Hérubel, 1904:564.— Monro, 1931:34.— Leroy, 1936:426.—Andrew & Andrew, 1953:1.—Rice & Stephen, 1970:67.— Cutler, 1973:179-180.
Paraspidosiphon cumingii. —Stephen & Ed- monds, 1972:243-244.—Edmonds, 1980: 50.
Aspidosiphon major Vaillant, 1871:270- 271; 18752 pl. 4, figs. Al-6.—De- Rochebrune, 1881:232.
Aspidosiphon klunzingeri Selenka et al., 1883:115-116, pl. 13, figs. 187-189.— Fischer, 1896:338; 1914a:70.—Sluiter, 1898:444: 1912:20.—Shipley, 1898:471; 1899b:153.—Heérubel, 1904:564.— Mon- ro, 1931:34.—Edmonds, 1956:308.— Wesenberg-Lund, 1957b:8-9; 1959a:196; 1959b:211-212; 1963:138.—Cutler & Cutler, 1979a:974—-975; 1979b:107.
Paraspidosiphon klunzingeri.—Stephen & Edmonds, 1972:247-—249.—Rice & Mac- intyre, 1972:42.—Rice, 1975:40-41.— Haldar, 1976:7.—Gibbs, 1978:85.
Aspidosiphon gigas Sluiter, 1884:39-—57, pl. 2, figs. 1-11, pl. 2a, figs. 12-25; 1886:473; 1891:116; 1902:19.
Paraspidosiphon gigas. —Stephen & Ed- monds, 1972:246.
Aspidosiphon angulatus Ikeda, 1904:45—47, figs. 11, 78-80; 1924:37.—Sato, 1939: 428.—Cutler & Cutler, 1979a:974; 1981: 81.—Cutler et al., 1984:308.
Paraspidosiphon angulatus. —Stephen & Edmonds, 1972:241.
Aspidosiphon speciosus Gerould, 1913:426- 427, text-fig. 16, pl. 62, fig. 22.— Fischer, 1922c:13.—Migotto & Ditadi, 1988:254— 25ST.
Paraspidosiphon speciosus. —Stephen & Ed- monds, 1972:253.—Rice, 1975:38-45.— Rice & Macintyre, 1979:314.
Aspidosiphon grandis Sato, 1939:414-419, pl. 21, fig. 21, text-figs. 46-50.— Cutler & Cutler, 1981:83-84.
853
Paraspidosiphon grandis.—Stephen & Ed- monds, 1972:246-247.
Aspidosiphon grandis obliquoscutatus Mu- rina, 1974:1713-1715, fig. 1.
Aspidosiphon pachydermatus Wesenberg- Lund, 1937:9-16, text-figs. 4-9.
Paraspidosiphon pachydermatus. —Stephen & Edmonds, 1972:250-251.
Aspidosiphon brasiliensis Cordero & Mello- Leitao, 1952:277-282, 288-292, text-figs. 1-5.
Paraspidosiphon brasiliensis.—Stephen & Edmonds, 1972:241-243.
Aspidosiphon johnstoni Edmonds, 1980:51- 53, figs. 91, 100—102.— Lopez et al., 1984: 194-196.
Aspidosiphon quatrefagesi Saiz Salinas, 1984:49-55, fig. 4.
Material examined. —MNHN, type ma- terial (V20); our 1985 Hawaii and 1988 Venezuela material; A. cumingii, type no longer at BMNH; 4. angulatus, Madagascar and Polynesia (Cutler & Cutler 1979a); A. brasiliensis, type cannot be located; A. gigas, ZMUA, holotype (V. Si. 8); A. grandis, ZITU, type specimen (24); A. johnstoni, two specimens from Edmonds’ original mate- rial; A. klunzingeri, type cannot be located; ZMUA, Sluiter’s Durban worm (V. Si. 9); UZMK, Wesenberg-Lund’s Cape Verde material; South Africa and Cape Verde (Cutler & Cutler 1979a, b); USNM, 26437 from Saipan identified by W. K. Fisher; A. pachydermatus, the type cannot be located; USNM, two specimens identified by W. K. Fisher (from Saipan, 24645 and Philip- pines, 21480); A. speciosus, USNM, type material (16820, 16391, 4088); two speci- mens from Brazil (Migotto & Ditadi, 1988).
Nomenclatural note. —Quatrefages’ orig- inal spelling is incorrect (Steyskal, pers. comm.) and when the correct ending is ap- pended the name A. /evis Sluiter becomes a junior homonym according to ICZN Art. 58.
Discussion. —A. laevis is a widespread but low density population of worms that has
854
INV fF TR
Fig. 8. Unidentate, compressed, Type A hooks of several A. laevis from different populations to show differences in shape and size, the latter being roughly correlated with trunk size. Scale line = 40 um.
been given several names over the past cen- tury. One of the characters that has been weighted very differently by various authors is the nature of the caecum and its elabo- rations as discussed above. Setting this aside, a second issue has been the presence/ab- sence of introvert spines. It is clear that when a few authors made reference to spines, they were either looking at scattered unidentate compressed hooks, or had a different species in hand. These worms have a solid anal shield bearing 10-15 longitudinal grooves. They have unidentate, compressed hooks, sharply pointed or blunt, in many rings (Fig. 8). These are 20-80 um tall, hook size being roughly correlated with trunk size. Also present are a few scattered compressed hooks (referred to as spines by some earlier au- thors). Up to 24 tentacles surround the nu- chal organ.
Internally, the pair of retractor muscles are fused for most of their length, some- times giving the impression of a single broad muscle with the ventral nerve cord running through a notch in the base. These muscles
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
usually have their origins from the body wall about 65-80% of the distance towards the posterior end of the trunk, 1.e., well in front of the caudal shield. Another distinc- tive feature is the bifurcation towards the anterior end of the spindle muscle. One branch continues along the rectum into the connective tissue and wing muscle to join the body wall just anterior to the anus. The second, and often larger branch, leaves the posterior rectum going to the dorsal body wall well posterior to the anus. In many specimens the contractile vessel is not a smooth tube but has vesicular pouches or swellings along the part of its length united with the retractors (not unlike that seen in some Golfingia specimens, and sometimes confused with contractile vessel villi [see Cutler & Cutler 1987:750]). The rectum usu- ally bears a caecum that may be simple or complex (see part 8, Morphological Char- acters section). The 25-35 longitudinal muscle bands anastomose frequently, and the circular muscle layer also subdivides into anastomosing bundles of varying degree of development. The nephridia open at, or just posterior to, the anus, are attached to the body wall for about half to two-thirds their length, and are usually more than half the trunk length.
The unpublished USNM material (21480 and 24645) consists of two worms, 44 and 123 mm long. They both have complex multiple caeca, a grooved anal shield, uni- dentate hooks (in the larger one these are blunt triangular and up to 60 um tall), and in all ways match A. J/aevis.
This new arrangement reduces ten puta- tive species to the rank of junior synonyms of A. laevis, a decision reached only after extended analysis. We shall not detail each case, but in a few instances there were ob- servational errors made by the original au- thor (for exarnple, A. gigas does have rows of unidentate hooks, a spindle muscle and a complex rectal caecum, all overlooked by Sluiter). However, in most cases the differ- ent conclusions are based on judgments
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about within-deme variation, or simply lack of either critical comparison or differential diagnoses (e.g., Gerould [1913:427] when describing A. speciosus simply says: “This species resembles A. klunzingeri from the Red Sea’”’ and nothing more as to how it differed). In another case Saiz Salinas (1984) erected A. quatrefagesi based on a suite of characters he assumed to be absent in older species while actually they were present but simply not mentioned by earlier authors.
As discussed in the Morphological Char- acters section, the variability in rectal ap- pendages is great, therefore, to use such a variable character to differentiate species is unwise. A more difficult issue for us, partly because of statements by Edmonds (1980) and Rice (1975), centers on the hook mor- phology. We have examined many hooks from museum specimens and newly col- lected material, comparing hooks from small worms to those from large ones, and hooks from anterior to posterior rings in the same worm (Fig: 8). We conclude that while pop- ulations do exhibit among-deme differ- ences, much of the alleged between species variation in shape is present within demes. Therefore, hook shape alone cannot be used to separate these putative species.
Distribution. —Widespread, but low den- sity, in warm water Indo-West Pacific Ocean (Durban to the Red Sea, Andaman Islands, Malaya to southern Japan, Indonesia, Great Barrier Reef and several islands out to Ha- wail). Also present in the Caribbean and west Atlantic (from 20°S to 31°N), then in the east Atlantic from the Canary and Cape Verde Islands to the Gulf of Guinea. In- habits shallow water coral rock.
Aspidosiphon tenuis Sluiter, 1886
Aspidosiphon tenuis Sluiter, 1886:49 1-492, pl. 3, fig. 7; 1891:116; 1902:19.
Paraspidosiphon tenuis.—Stephen & Ed- monds, 1972:257.
Aspidosiphon levis Sluiter, 1886:493-494, pl. 3, fig. 8; 1891:116; 1902:18.
855
Paraspidosiphon levis. —Stephen & Ed- monds, 1972:249-—250.
Aspidosiphon ambonensis Augener, 1903: 325-328, figs. 5-8.
Aspidosiphon steenstrupii var. ambonensis Fischer, 1922a:24—26; 1923:21.
Paraspidosiphon ambonensis.—Stephen & Edmonds, 1972:240-241.
Aspidosiphon formosanus Sato, 1939:421- 424, pl. 21, fig. 23, text-figs. 55—57.— Cut- ler & Cutler, 1981:81-83.
Paraspidosiphon formosanus.—Edmonds, 1971:144—-146; 1980:50-51.—Stephen & Edmonds, 1972:245.
Aspidosiphon havelockensis Haldar, 1978: 37-41, figs. 1-2.
Aspidosiphon speculator.—Cutler & Cutler, 1979a:975—-976 (partim).
Material examined. —BMNH, (1889.6.15. 42/44) syntype; A. levis, ZMUA, type ma- terial (V. Si. 11); A. ambonensis, MNHU, type material (6956-6958); A. formosanus, type material cannot be located, but we have examined material from Guam and Austra- lia identified by S. J. Edmonds; A. speculator, Pacific Ocean (Cutler & Cutler 1979a).
Discussion. —A. tenuis 1s here defined as having an anal shield made up of very fine dark units with a smooth overall appear- ance; a few very short grooves may appear around the margin. The distal rings of bi- dentate hooks (30-60 um tall) lack the dis- tinct tongue on the clear streak (Fig. 2C) and are followed by scattered, unidentate hooks (25-60 um tall) with an internal clear streak in the distal ones. More proximally, these scattered unidentate structures have lateral reinforcing ridges. Dark pyramidal or con- ical hooks are absent. Internally this species is very much like A. steenstrupii except that only two of the ten worms dissected has a rectal caecum and the nephridia are less than 50% of the trunk length.
When Sluiter (1886) described this species he overlooked the distal rings of bidentate hooks and posterior attachment of the spin-
856
dle muscle. He overemphasized the few small grooves around the margin of the anal shield. The caudal shields are not all dis- tinctly grooved. Therefore, this is clearly conspecific with A. /evis from the same lo- cation, and becomes the senior synonym by virtue of their position in the text.
Fischer (1922a) reduced Aspidosiphon ambonensis to a variety of A. steenstrupii, but Stephen & Edmonds (1972) elected to elevate it back to species rank since the shape of the clear area in the hook “seems to be different.’’ We assume they based their con- clusion on Augener’s (1903) picture (their fig. 29K is Augener’s fig. 6). Our examina- tion of the type material (19 worms) con- firms Augener’s perceptions and supports our position that this population is conspe- cific with A. tenuis, but different than A. steenstrupil in accordance with Stephen & Edmonds (1972).
Sato (1939) erected A. formosanus, but Cutler & Cutler (1981) reduced it to a junior synonym of A. steenstrupii. Given our cur- rent understanding we would move A. for- mosanus into the synonomy of A. tenuis as it shares its attributes.
In 1978 Haldar described Aspidosiphon havelockensis. In his unpublished disserta- tion (pers. comm.) it is reduced to a junior synonym of A. steenstrupii ambonensis dif- ferentiated from the nominate form on the basis of hook, spine, and papillae structure. We agree with his conclusions as far as they go. However, we now consider both these names to be junior synonyms of A. tenuis.
Upon reexamination, the specimens Cut- ler & Cutler (1979a) identified as A. spec- ulator from the Solomon Islands and Thai- land belong in this taxon.
Distribution. — Andaman Islands to Thai- land, Formosa, and Guam, out to the east- ern Caroline Islands and down through the Solomon Islands to the Great Barrier Reef and Indonesia.
Aspidosiphon parvulus Gerould, 1913
Aspidosiphon parvulus Gerould, 1913:425— 426, pl. 61, fig. 17, text-fig. 15.—Stephen
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& Edmonds, 1972:233-234.—Cutler, 1973:178-179.
Aspidosiphon spinoso-scutatus Fischer, 1922c:13-14, text-figs. 2—3.— Murina, 1967b:42; 1967¢:1332.
Paraspidosiphon spinososcutatus. —Ste- phen & Edmonds, 1972:254.—Rice, 1975:38-45.
Material examined. —USNM, type (15118); western north Atlantic (Cutler 1973); our 1988 Venezuelan worms; A. spinoso-scu- tatus, MNHU, type (6053).
Discussion.—This species shares many attributes with A. fischeri, but the shield morphology seems consistent and distinc- tive. Centrally it is made up of larger flat plates; these are sometimes arranged in rows giving an impression of ridges and grooves. Ventrally and laterally the units become smaller, scattered, wart- or cone-shaped. The shields have a diffuse boundary where the units grade into coarse trunk papillae. At both ends of the trunk these darker papillae are located in rectangles that remind one of Sipunculus skin. The hooks (bidentate and unidentate in rings, scattered unidentate, and pyramidal, Fig. 1E) are 25-35 um tall. The 10-12 short tentacles may appear webbed together with connective tissue, and there are about 24 anastomosing longitudinal muscle bands. These bundles are generally quite distinct in worms over 5 mm long, more so towards the anterior end (see Cutler 1973:178 where Gerould’s 3 and 4 mm specimens are discussed). The nephridia are 50-75% of the trunk length.
Distribution.—Western Atlantic Ocean from Cape Hatteras through the Caribbean to Venezuela; often found together with Themiste alutacea and Nephasoma pellu- cidum in branching corals.
Aspidosiphon planoscutatus Murina, 1968
Aspidosiphon planoscutatus Murina, 1968: 1722-1724, figs. 1-2; 1971:78.
Material examined.—ZIAS, type mate- rial.
VOLUME 102, NUMBER 4
Discussion. —This taxon is based on two specimens and is very similar to A. steen- strupii but has only unidentate compressed hooks on the introvert. Also, the shield units are smaller (fine grained like A. zinni) and the trunk is densely covered with more ob- vious skin bodies. The nephridia are 85% of the trunk length.
The Red Sea is a marine habitat with un- usual abiotic conditions (e.g., high salinity, low oxygen, high temperature) that may re- strict gene flow and favor selection of dif- ferent allelic frequencies. The absence of bi- dentate hooks on the introvert may be a real difference, but we cannot verify this. De- spite our reservations we are leaving the name because of the habitat. It is hoped that more collecting in the area will produce ad- ditional material for analysis.
Distribution. —Red Sea at 40 m.
Aspidosiphon steenstrupii Diesing, 1859
Aspidosiphon steenstrupii Diesing, 1859:767, pl. 2, figs. 1-6.— Quatrefages, 1865:610.— Selenka et al., 1883:116—118.—Sluiter, 1886:489-490; 1891:115; 1902:18.— Whitelegge, 1899:394.—Shipley, 1899b: fo 4 1902:131—132: 1903:171.— Ikeda, 1904:40-41; 1924:38.—Heérubel, 1904:564.—Lanchester, 1905b:39.— Fischer, 1914a:70—71; 1914b:13; 1922a: Beet 2 2-1 3: 1923-21; 1926:108; 1931: 139.—ten Broeke, 1925:93-94.— Monro, 1931:34.—Sato, 1935:315-316; 1939: 424-426.—Leroy, 1936:426; 1942:36- 38.—Stephen, 1942:253.—Stephen & Robertson, 1952:441.—Edmonds, 1956: 307-308.— Wesenberg-Lund, 1959a:197- 198: 1963:138.—Murina, 1967b:42; 1981:12-13.—Cutler, 1977a:148.—Cut- ler & Cutler, 1979a:976; 1979b:107- 108.—Cutler et al., 1984:308-309.— Migotto & Ditadi, 1988:259-260.
Paraspidosiphon steenstrupii steenstrupil. — Stephen & Edmonds, 1972:254—-255.— Rice, 1975:38-—45.—Haldar, 1976:8.— Rice & Macintyre, 1972:42; 1979:311- 319.—Edmonds, 1980:51.
857
Aspidosiphon steenstrupii var. faciatus Au- gener, 1903:322-325, figs. 1-4.
Paraspidosiphon steenstrupii fasciatus. — Stephen & Edmonds, 1972:255-256.
Aspidosiphon fuscus Sluiter, 1881:86-108; 1886:474; 1891:116; 1902:19.—Selenka et al., 1883:116.
Aspidosiphon semperi ten Broeke, 1925:92, text-figs. 18—20.—Gibbs & Cutler, 1987: 56.
Paraspidosiphon semperi.—Stephen & Ed- monds, 1972:252.
Aspidosiphon speculator Selenka, 1885:19- 20, pl. 4, figs. 24~27.— Fischer, 1914b:71; 1920:413.—Wesenberg-Lund, 1959b: 213.—Cutler & Cutler, 1979a:975-976 (partim).— Not Saiz Salinas, 1986a:11-14.
Paraspidosiphon speculator.—Stephen & Edmonds, 1972:253-254.
Aspidosiphon makoensis Sato, 1939:419- 421, pl. 21, fig. 22, text-figs. 51-54. —Cut- ler & Cutler, 1981:82-83.
Paraspidosiphon makoensis. —Stephen & Edmonds, 1972:250.
Aspidosiphon trinidensis Cordero & Mello- Leitao, 1952:283-286, 292-294, figs. 6— 10.—Cutler & Cutler, 1979b:108; 1980c: 206.
Paraspidosiphon trinidensis. —Stephen & Edmonds, 1972:257-—258.
Aspidosiphon exostomum Johnson, 1964: 331-332, pl. 7.
Paraspidosiphon exostomus. —Stephen & Edmonds, 1972:244.
Aspidosiphon ochrus Cutler & Cutler, 1979a: 976-979, figs. 15—17.—Edmonds, 1987: 204.
Material examined.—ZMUA, Sluiter’s 1902 specimens (V. Si. 21); our Pacific and Caribbean material; Brazil (Migotto & Di- tadi 1988); A. semperi, ZMUA, types (V. Si. 14); A. exostomum, RSME, type (1965.32.2); A. speculator, BMNH, 1885. 12.3.28, syntype; specimens from Canary Islands and Spain identified by J. Saiz Sa- linas; Madagascar (Cutler & Cutler 1979a).
Discussion. —We came to the present un- derstanding of A. steenstrupii only after
858 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
looking at many worms from diverse lo- cations, and carefully reexamining the lit- erature. The color of the anal shield was a confusing element. It now seems clear that a range of colors is possible, from almost white to very dark brown, and that addi- tional calcareous material may be deposited externally, thus masking the underlying units. Geographic variation seems present with the Atlantic Ocean populations being dark, the mid Pacific Ocean populations being pale and the Indian Ocean popula- tions exhibiting a mixture (a higher fre- quency of dark shields in populations near continents, rare in island populations).
The other major issue centers around the hooks. We are defining this species as hav- ing bidentate hooks in rings (30-60 um tall; up to 90 wm in worms over 25 mm long), most with a tongue-like extension on the internal clear streak (Fig. 2B), and no uni- dentate compressed hooks. The proximal introvert does bear many scattered, dark, pyramidal hooks about 30-60 um tall.
Internally the pair of retractor muscles originate about 70-85% of the distance to the posterior end of the trunk, not correlated with trunk size (see Table 2). The nephridia are commonly 50-80% of the trunk length and attached to the body wall for 50-75% of their length (over 90% in three worms). A simple rectal caecum was observed in 21 of the 30 worms dissected. The longitudinal muscle bands anastomose (14—22 anteriorly and 20-28 posteriorly), and the number is not correlated with trunk length.
J. Silverstein (pers. comm.) determined the karyotype on a Japanese population. The 2N number is 20 with five pairs of meta/ submetacentric and five pairs of telo/sub- telocentric chromosomes.
When Selenka (1885) erected A. specu- lator he made no reference to any other species (no differential diagnosis or key). In Stephen & Edmonds (1972) the key sepa- rates these two species based on the location of the retractor origins that were imprecisely stated in the original. In the syntype, the
origins are at 75% of the distance to the posterior end of the trunk, well within the A. steenstrupii range. The internal structure of the compressed hooks and the nature of the pyramidal hooks, shield, and other char- acters all match this species. The part of Cutler & Cutler (1979a) collection that be- longs here are the Madagascar worms, the remainder are A. tenuis. Saiz Salinas (1986a) used A. speculator for a collection that we consider to be A. misakiensis.
Fischer (1922a) reduced Augener’s 1903 A. steenstrupii fasciatus to a junior synonym of the nominate form where it remained until Stephen & Edmonds (1972) resurrect- ed its subspecies rank. They based their de- cision on the clear area in the hook and color differences on the shield and mid-trunk. Our examination of the type, within the context of this study, convinces us that Fischer’s action was correct.
When ten Broeke (1925) described A. semperi from Curacao she asserted that it had four retractor muscles, but Gibbs & Cutler (1987:56) determined that there are only two. Despite the pale colored anal shield A. semperi is clearly conspecific with A. steenstrupii, a conclusion confirmed by our recent collections in Curacao.
Sato’s two species, Aspidosiphon formo- sanus and A. makoensis were reduced to the status of junior synonym in Cutler & Cutler (1981). We reaffirm that action for the latter but not the former (see below).
Aspidosiphon trinidensis was described from a single worm that cannot be located. The two subsequent reports were also based on single worms (in Cutler & Cutler 1980c, it should have read 25 mm trunk, not 125 mm). Reexamination of the two available worms revealed a few distal rings of biden- tate hooks that had been overlooked. Those structures reported as unidentate hooks are now interpreted as pyramidal hooks. Cor- dero & Mello-Leitao’s worm (1952) had its introvert entirely withdrawn and the objects they described as unidentate hooks were not in rings. From their words and drawings we
VOLUME 102, NUMBER 4
interpret these as pyramidal hooks. With this understanding (and the assumption that they too overlooked the bidentate hooks) nothing separates A. trinidensis from A. steenstrupii, thus we place it in synonomy.
Aspidosiphon exostomus (Johnson 1964) was alleged to be different because of the dorsal crown of tentacles. This was clearly visible as the esophagus was protruding through the mouth. However, we now know that all members of the Aspidosiphonidae have dorsal tentacles and this taxon is clear- ly not unique.
When Cutler & Cutler (1979a) described A. ochrus we were working within a different experiential framework. As a result of the present analysis, it clearly should be reduced to ajunior synonym. In that same paper we
identified a specimen from Madagascar as |
A. speculator that, upon reexamination, we now consider to be an A. steenstrupii. Ed- monds (1987) used the name A. ochrus after consulting with us, but these also are A. steenstrupii with pale anal shields.
Distribution.—Throughout the western and northern Indian Ocean, Queensland through Indonesia and the South China Sea to southern Japan, out through the western Pacific islands to Hawaii. Also collected from numerous Caribbean locations, in the eastern Atlantic only from the Cape Verde Islands and the Gulf of Guinea. It lives in shallow water coral rocks.
Zoogeographical Summary
Ten of the 19 species live in the tropical/ subtropical western Atlantic Ocean and Ca- ribbean Sea, an area bounded by Cape Hat- teras on the north and the Amazon delta on the south (A. albus, A. exiguus, A. gosnoldi, A. parvulus, A. fischeri, A. mexicanus, A. elegans, A. laevis, A. steenstrupii, and A. mi- sakiensis). The first four of these are endem- ic to the region. The fifth extends into the eastern Pacific (Panama to Galapagos). The sixth extends its range in the other direction, to the eastern Atlantic (between Iberia and
859
the Gulf of Guinea) but nowhere else. The next three species are circum-tropical while the last is found on both sides of the Atlantic Ocean and off Japan and Australia.
Two species are found in the eastern At- lantic and elsewhere that do not live in the western part (A. venabulus from both sides of Africa, and A. muelleri, see below). Also in the north Atlantic (plus one record from the Mozambique Channel) is A. zinni, the one bathyal/abyssal member of this genus.
Of special note is A. muelleri, since it has the most widespread distribution, almost cosmoplitan in temperate waters. Two ap- parent gaps occur; in the western Atlantic (except for one record off southern Brazil), and in the eastern Pacific there is only one record off Chile. This is the most eurytopic Aspidosiphon living in a much wider variety of temperatures and depths than other species.
Six species are widely distributed within the Indo-West Pacific area. Aspidosiphon gracilis schnehageni and A. coyi extend into the eastern Pacific Ocean. Three are also found in the Caribbean (as above). The re- maining two do not get to Hawaii or the Atlantic (A. gracilis gracilis and A. tenuis). Two species (A. thomassini and A. spiralis) are more restricted within the Indian Ocean and A. planoscutatus is known only from a single collection in the Red Sea.
Of particular interest is the number of endemic species in the warm water Atlantic/ eastern Pacific Ocean (six) when compared to the Indo-West Pacific area (five). Of the 19 species, 13 live somewhere in the Atlan- tic Ocean while 11 occupy some part of the Indo-West Pacific (six of these are in both areas). These data have interesting evolu- tionary implications suggesting that the tra- ditional “‘center of origin’’ hypothesis for marine invertebrates (Indo-West Pacific) may not fit Aspidosiphon.
While it is true that common, widespread species bore in coral or soft rock, 11 species (58%) do not occupy this stereotypical as- pidosiphonid niche, i.e., they live in dis-
860
carded mollusc shells (8), arenaceous fora- miniferan tests (1), or interstitially (2):
Acknowledgments
The encouragement and exchange of ideas and specimens with S. J. Edmonds, Ade- laide, has contributed greatly to the com- pletion of this work. A working visit from J. Saiz Salinas, Bilbao, including an open exchange of perceptions, was very fruitful. The opportunity to read unpublished manuscripts and to examine some speci- mens of A. E. Migotto and A. S. F. Ditadi, Sao Paulo, and B. P. Haldar, Calcutta, has added significantly to our understanding. Our work in Hawaii was accomplished with the assistance of B. Burch, Bishop Museum and M. Hadfield, Kewalo Marine Labora- tory. The field work in Curacao was depen- dant on the cooperation of W. Bakhuis, Car- mabi Foundation. In Cumana J. Perez and associates at the Universidad de Oriente as- sisted us in many ways. The visit to the Fundacion Cientifica Los Roques was made possible by the cooperation of R. Laughlin and B. Alverez. Lynn Cutler, Mountain View, and J. Silverstein, Seattle, provided invaluable assistance in the field and lab. Correct spelling of species’ names was kind- ly provided by G. Steyskal, Washington, and K. Bart, Clinton, assisted with the scanning electron microscope. Financial support was provided by the National Science Founda- tion (BSR 86-15315).
The cooperation of the following persons and institutions in the loan of reference material and/or providing access to their collections was essential to the completion of this project and greatly appreciated: E. Easton (BMNH); J. Rénaud-Mornant (MNHN); G. Hartwich (MNHU); C. Car- pine (MOMV); R. Olerod (NHRS); S. Chambers (RSME); M. Rice (USNM); J. Kirkegaard (UZMK); A. Ivanov (ZIAS); Zoological Institute Tohoku University, Sendai; A. Pierrot-Bults (ZMUA); E. Wil- lassen (ZMUB); M. Dzwillo (ZMUBH).
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
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. 1955. Reports of Lund University Chile Ex- pedition, 1948-49. 19. Gephyrea from Chile. — Lunds Universitets Arsskrift 51(10):1-23. . 1957a. Sipunculoidea and Echiuroidea from West Africa together with a bibliography on Ge- phyrea after 1920.—Bulletin Institut royal des Sciences naturelles de Belgique 33(42): 1-24. 1957b. Sipunculoidea from the coast of Is- rael.— Bulletin of the Research Council of Israel 6b(3-4): 193-200. . 1957c. Sipunculoidea and Echiuroidea from the Red Sea.—Contributions to the knowledge of the Red Sea No. 3, The Sea Fisheries Re- search Station, Haifa Bulletin 14:1-15. . 1959a. Sipunculoidea and Echiuroidea from tropical West Africa.— Atlantide Report 5:177- DVO:
1959b. Sipunculoidea and Echiuroidea.—
Resultats Scientifiques des Campagnes de la “Calypso” 4:207-217. . 1959c. Sipunculoidea and Echiuroidea from Mauritius. — Videnskabelige Meddelelser Danske fra Naturhistorisk Forening 1 Kjobenhavn 121: 53-73.
1963. South African sipunculids and echiu- roids from coastal waters. — Videnskabelige Meddelelser Danske fra Naturhistorisk Foren- ing 1 Kjobenhavn 125:101-146.
Whitelegge, T. 1899. The Hydrozoa, Scyphozoa, Ac- tinozoa and Vermes of Funafuti.— Memoirs of the Australian Museum, Vermes 3:37 1-394.
Zavodnik, D., & V. G. Murina. 1975. Contribution to Sipuncula of North Adriatic insular region. — Rapports et procés-verbaux des reunions com- mission internationale pour |’Exploration Scientifique de la Mer Méditerranée 23(2):127- 128.
—_, & 1976. Sipuncula of the region of Rovinj (North Adriatic Sea).—Biosistematika 2(1):79-89.
(EBC) Biology Department, Utica College of Syracuse University, Utica, New York 13502; (NJC) Biology Department, Ham- ilton College, Clinton, New York 13323.
PROC. BIOL. SOC. WASH. 102(4), 1989, pp. 866-871
AXIOTHELLA CROZETENSIS, A NEW SPECIES OF MALDANID POLYCHAETE FROM CROZET ISLANDS (INDIAN OCEAN)
Patrick Gillet
Abstract.—A new species of Annelida Polychaeta, Axiothella crozetensis, is described from Crozet Islands (46°46.6’'S and 50°28.4’E, Indian Ocean). Spec- imens were collected from depths of 1350 to 1440 m during the MD.08/Benthos Expedition (7 Mar 1976-26 Apr 1976). Axiothella crozetensis is compared with all described species of the genus Axiothella.
Maldanid polychaetes, called bamboo- worms, are capitelliform, usually long and truncate; the prostomium with nucal organs and a cephalic keel, without appendages, is fused to the buccal segment. Reviews of the family are found in Arwidsson (1907). Ax- lothella crozetensis 1s the first species of the genus Axiothella collected at a great depth. Holotype and paratypes are deposited in the Institute of Fundamental and Applied Re- search Museum (I.R.F.A.-MAL-047).
Genus Axiothella Verrill, 1900
Type species. —Axiothella catenata (Malmgren, 1865:190-191).
Diagnosis. —Eighteen to 20 setigers pres- ent. Rostrate uncini present in all neuro- setigerous segments including the first three neuropodia. Each uncinus with lateral series of teeth above main fang without barbules. Acicula absent. Rim of cephalic plate entire or incised. Anal plaque funnel-shaped; anal cirri of different lengths or as crenulations. Mangum (1962) reviewed the status of the genus Axiothella which she placed as a sub- genus of Clymenella Verrill (Mangum 1962, 1966). This decision did not gain acceptance
Fig. 1.
and authors now recognize the original ge- neric status.
Axiothella crozetensis, new species Figs. 1-2
Material examined. —Holotype Crozet Islands (19 Apr 1976), Station 70: Sampling DC 280, 46°46.6’S and 50°28.4’E; Paratypes (2) Crozet Islands (19 Apr 1976), Station 70: DC 280, 46°46.6’S, 50°28.4’E. Materials deposited to I.R.F.A. Museum IRFA MAL 047.
Description.—Holotype has 18 setigers and two achaetous preanal segments. Body without color pattern in alcohol. Prosto- mium longer than wide (1.4 mm long; 0.70 to 0.90 mm wide) with the cephalic plate 0.60 mm long (Fig. 1F). First setiger 1.4 mm long, second setiger 2 mm long, diameter ranging from 0.50 to 0.75 mm. Prostomium well developed with entire rim, not incised. Nucal organs long and parallel; ocelli ab- sent. Buccal segment as long as the first se- tiger (Figs. 1A, 2A). First setiger depressed anteriorly near fascicle of capillary setae. Notosetae numbering about 15 per fascicle, short and long, are all capillary smooth (Fig.
~
Axiothella crozetensis: A, Anterior region lateral view; B, Posterior region; C, Uncinus from setiger
2; D, Uncinus from setiger 4; E, Capillary setae from setiger 4; F, Cephalic plate frontal view.
VOLUME 102, NUMBER 4 867
3
L Nase Bene
0.05mm
868 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
LSE ASEHE ABBE
Fig. 2. Scanning electron micrographs of Axiothella crozetensis: A, Anterior region lateral view; B, Posterior region; C, Uncinus from setiger 3; D, Row of uncini from setiger 4; E, Uncinus from setiger 4.
VOLUME 102, NUMBER 4
1E). Spinulose, barbelate and bipennate no- tosetae are absent. Neurosetae of the first setiger are 1-2 uncini per side, each with 4—6 apical teeth without barbules (Figs. 1C, 2C). The second setiger has two or three uncini, the third setiger has three uncini and the fourth setiger six uncini. Uncini from setiger 4 to the end of the worm have bar- bules below the main fang and one row of s1x Or seven vertical teeth and many smaller ones on each side (Figs. 1D, 2D, 2E). Glan- dular area absent. The posterior region has two achaetous preanal segments. Pygidium is prolonged posteriorly as a cone with three long anal cirri, each of equal length and nearly the same as the length of the pygid- ium (Figs. 1B, 2B). The anus is subdorsal with a ventral value.
Discussion. —Axiothella crozetensis be- longs to the subfamily Euclymeninae, genus Axiothella Verrill, 1900, because of the presence of rostrate uncini on the first three neuropodia. Other genera without acicular spines are Gravierella Fauvel, 1919 and Macroclymenella Augener, 1926, but the number of segments in these genera is more than 30. Species of the genus Maldanella McIntosh, 1885 have the first setiger with notosetae only, and those of the genus C/y- menura Verrill, 1900 have a flanged collar on setiger 4 and a large triangular glandular field on setiger 8.
Axiothella crozetensis differs from other species of Axiothella in that it has only one to three uncini on the first three setigers and a prolonged anal cone with three long anal cirri as in the genus Clymenura (Table 1). Among the species living in the Antarctic and Subantarctic regions Axiothella jarli Kirkegaard, 1959 (Angola, Cape) differs in having from three to seven hooks on the first three setigers. The structure of the py- gidium is different with 18 anal cirri alter- nately long and short. Axiothella antarctica Monro, 1930 differs in having three uncini on the first setiger and numerous on the second and third setigers, pygidium with a
869
long single anal cirrus and fifteen shorter cirri. Axiothella quadrimaculata Augener, 1914 has numerous uncini on the first three setigers, only one achaetous preanal seg- ment and also differs in lacking anal cirri. Hartmann-Schroder & Hartmann (1984) found two different species of Axiothella sp. at Murat Bay, Australia: Axiothella sp. A has uncini with five teeth and short barbules and Axiothella sp. B has six teeth and long barbules. This species was also found at Blanche Harbour, Australia (Hartmann- Schroder 1985).
Axiothella crozetensis has been found at depths exceeding most records given for other species of Axiothella living in the in- tertidal zone except Axiothella constricta Claparéde, 1868 at 35-750 m (Tarente) and Axiothella catenata Malmgren, 1865 at 10— 900 m (Arctic) (Table 1). Axiothella quad- rimaculata lives in southwest Australia at 3 to 12.5 m on rocks or sand (Augener 1914), at Kerguelen Islands at 20-50 m (Monro 1939), in South Africa at 0-54 m (Day 1961) and at Marion and Prince Edward islands on the shore with ascidians and amphipods (Day 1971). Axiothella crozetensis was dredged from 1350 to 1440 m in mud flats with associated fauna of Annelida Poly- chaeta including Nephtys hombergii, Try- panosyllis gigantea, Ampharete kerguelensis and Melinna cristata. Morphologically and ecologically these two species differ accord- ingly: Axiothella quadrimaculata lives on rocky bottoms near the shore from 0 to 54 m and Axiothella crozetensis lives in mud flats at a depth of about 1400 m. These two species differ from others species of Axio- thella by the structure of the pygidium. Based on this character, the genus Axiothella is in need of revision. Axiothella catalinia (Hart- man 1969) was transferred to Maldanella by Kudenov and Read (1978) because of the absence of uncini on the first setiger, and Clymenura brasiliensis (Mangum 1966) must be referred to Axiothella.
Habitat. —Subantarctic, Crozet Islands
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Table 1.—Species of the genus Axiothella Verrill, 1900 (N: numerous uncini).
Species
Distribution, depth (m)
A. antarctica Monro, 1930
A. australis Augener, 1914
A. brasiliensis Mangum, 1966
A. catenata Malmgren, 1865
A. cirrifera Langerhans, 1880
A. constricta Claparéde, 1868
A. jarli Kirkegaard, 1959
A. mucosa Andrews, 1891
A. obockensis Gravier, 1905
A. quadrimaculata Au- gener, 1914
A. rubrocincta Johnson, 1901
A. serrata Kudenov and Read, 1978
A. somersi Verrill, 1900
A. tambalagamensis Pil- lai, 1961
A. zetlandica McIntosh, 1915
A. crozetensis, present work
A. sp. Wesenberg-Lund, 1949
A. sp. Imajima, 1963
A. sp. Wolf, 1984
A. sp. Hartmann-Schro- der and Hartmann, 1984
Antarctica, 200-344
Australia, India, 3—12.5
Brazil, 1-12
Arctic, 10-900
Madeira, ?
Italy, 35 to 750 m
Angola, Cape, 35-100
United States, Intertidal
Red Sea, India, Intertid- al
Subantarctic, 11—50
United States, Intertidal
New Zealand, Intertidal
Bermudas, Intertidal
Ceylon, ?
North Sea, 110-183
Crozet Island, 1350- 1440
Iranian Gulf, 71
Kamchatka, 68 Gulf of Mexico, 19-43
Australia, ?
Achae- Number of setae tous setigers 1 — 2 —3 segments Pygidial cirri
3-N-N 5 1 long cirrus, 15 short cirri
N-N-N 2 7-11 long, 23-30 short cirri
4/6 — 4/6 — 3/5 2 alternately short and long
8-9-15 4 22 alternately short and long
1 to 6 2 1 median cirrus, 14 pa- pillae
N-N-N 3 1 long cirrus, numerous short
3-6-7 D 18 alternately short and long
12-N-N 3/4 20 to 30 short and long
N-N-N D 1 long cirrus numerous short
N-N-N 0/1 without cirri
N-N-N 2/3 1 long cirrus, 18/30 short/long
6-—7-10 0 1 long cirrus, 21 to 42 cirri
3/4 — 4/5 -— 6/8 2 24 alternately short and long
N-N-N 0 numerous cirri, 13 pa- pillae
2/3 -4-4 & 44 alternately short and long
1/2 — 2/3 -—3 2 3 long cirri
25 — 25 — 25 U u
no accessory teeth
10-N-N ? v
2/3 with barbules 1 1 long cirrus, 20-24 short
1 to 4 D 1 long cirrus, 15 short
C1IT1
(South of Penguins Island) from 1350 to 1440 m in mud flats.
Acknowledgments
I am grateful to Patrick M. Arnaud, Sta- tion Marine d’Endoume, Marseille; to the South Africa Museum; and to Dr. Renaud- Mornant, Muséum d’Histoire Naturelle,
Paris, for collections. Special thanks to Dr. Gruet, University of Nantes and to Dr. Fil- mon, University of Angers, for their help with scanning electron microscopy.
Literature Cited
Andrews, E. A. 1891. Report upon the Annelida Polychaeta of Beaufort, North Carolina.—Pro-
VOLUME 102, NUMBER 4
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Arnaud, P. M., & G. Hureau. 1979. Compte-rendu de la campagne MD.08/Benthos (7 mars—26 avril 1976).—Comité National Frangais des Re- cherches Antarctiques 44:1-38.
Arwidsson, I. 1907. Studien iiber die scandinavisch- en und arktischen Maldaniden nebst Zummer- stellung der ubrigen bisher bekannten Arten dieser Familie.— Zoologischen Jahrbiichern Suppl. 9:1-308.
Augener, E. A. 1914. Polychaeta II. Sedentaria. Die Fauna Siidwest— Australiens. — Herausgeg. von W. Michaelsen und R. Hartmeyer 5:1-170.
Claparéde, C. 1868. Les Annélides Chétopodes du Golfe de Naples.— Mémoires Société Physique, Genéve 19:313-584.
Day, J. H. 1961. The Polychaete fauna of South Af-
rica. Part 6. Sedentary species dredged off Cape
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463-560.
. 1971. Polychaeta. Pp. 384-390 in E. M. van
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Gravier, C. 1905. Annélides Polychétes de la Mer Rouge. Cirratuliens Amphicténiens, Terebel- liens, Maldaniens.— Bulletin du Museum His- toire Naturelle de Paris 7:319-327.
Hartman, O. 1969. Atlas of sedentariate polychae- tous annelids from California. Allan Hancock Foundation, University of Southern California, Los Angeles, 812 pp.
Hartmann-Schroéder, G. 1985. Zur kenntnis des Eu-
litorals des australischen kisten unter beson-
derer Beriicksichtigung des Polychaeten und Os- tracoden. Teil 11. Die Polychaeten der antiborealen Siidkiiste Australiens (zwischen
Port Lincoln im Westen und Port Augusta im
Osten). — Mitteilungen aus dem Hamburgischen
Zoologischen Museum und Institut 82:61-99.
,&G. Hartmann. 1984. Zur kenntnis des Eu-
litorals der australischen kiisten unter beson-
derer Bericksichtigung des Polychaeten und Os- tracoden. Teil 10. Die Polychaeten der antiborealen Siidkiiste Australiens (zwischen
Albany im Westen und Ceduna im Osten).—
Mitteilungen aus dem Hamburgischen Zollo-
gischen Museum und Institut 81:7-62.
Imajima, M. 1963. Polychaetous annelids collected off the west coast of Kamchatka. II. Notes on species found in the collection of 1959.—Pub- lications of the Seto Marine Biological Labo- ratory 11(2):345-372.
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Johnson, H. P. 1901. The Polychaeta of the Puget Sound region.— Proceedings of the Boston So- ciety of Natural History 29:38 1-437.
Kirkegaard, J. B. 1959. The Polychaeta of West Af- rica.—Atlantide Report 5:7-117.
Kudenov, J. D., & G. B. Read. 1978. Axiothella ser- rata n. sp. a maldanid polychaete from Porirua Harbour, New Zealand. — New Zealand Journal of Marine and Freshwater Research 1 1(4):697- 702.
Langerhans, P. 1880. Die Wurmfauna von Madei- ra.— Zeitschrift fur Wissenschaften Zoologie, Leipzig 34:513-592.
Malmgren, A. J. 1865. Nordiska Hafs Annulater.— Ofversigt af Konglia Vetenskaps Akademiens Forhandlingar Stockholm 2:181-192.
Mangum, C. P. 1962. Studies on speciation in mal-
danid polychaetes of the North American At-
lantic coast. I. A taxonomic revision of three species of the subfamily Euclymeninae.— Pos-
tilla 65:1-12.
. 1966. Two new species of Clymenella (Poly-
chaeta: Maldanidae) from Brazil.— Postilla 104:
1-10.
McIntosh, W. C. 1915. A monograph of the British marine annelids.— Ray Society, Dulau and Co. Ed. 3:217-229.
Monro, C. C. A. 1930. Polychaete worms. — Discov-
ery Reports, Cambridge 2:1-122.
1939. B.A.N.Z. Antarctic Research Expedi- tion 1929-1931.—Part 4 Polychaeta. B.A.N.Z.A.R. Expedition Committee 4(4):89- 156.
Pillai, T. G. 1961. Annelida Polychaeta of tambala- gam Lake, Ceylon.—Ceylon Journal of Sciences 4:1-40.
Vermrill, A. E. 1900. Additions to the Turbellaria, Nemertia and Annelida of the Bermudas with revisions of some New England genera and species.— Transactions of the Connecticut Academy of Arts and Sciences 4(2):595-672.
Wesenberg-Lund, E. 1949. Polychaetes of the Iranian Gulf.— Danish Scientific Investigations in Iran 4:347-400.
Wolf, P. S. 1984. Maldanidae. Pp. 1-21 in J. M. Uebelacker and P. G. Johnson, eds., Taxonomic guide to the polychaetous annelids of the north- ern Gulf of Mexico. Barry A. Vittor and Asso- ciated, Mobile, Alabama 2(15):1-21.
Institute of Fundamental and Applied Research, Zoology and Marine Ecology De- partment, BP 808, 49005 Angers Cedex, France.
PROC. BIOL. SOC. WASH. 102(4), 1989, pp. 872-877
PARAPROCERASTEA CROCANTINAE, A NEW GENUS AND SPECIES (POLYCHAETA: SYLLIDAE: AUTOLYTINAE) FROM THE SPANISH MEDITERRANEAN
Guillermo San Martin and Carmen Aloés
Abstract. —Paraprocerastea crocantinae, a new genus and species of Auto- lytinae (Syllidae: Polychaeta) from off Cabo de Creus and Cabo de Gata both on the Mediterranean coasts of Spain, is described. The new genus is discussed and compared with the remaining genera of Autolytinae. Finally, a key to the
genera of Autolytinae is given.
In a study of Alos (1988) on the poly- chaetous annelids from the Cabo de Creus coast (Spain), two specimens ofa syllid were found. These specimens were originally identified as an unknown species of Proce- rastea Langerhans, 1884. Another speci- men, longer than the others, was collected by the first author in a study of the poly- chaetes from Posidonia oceanica beds along the southern coast of Spain. This latter study was part of a project on the invertebrates from this habitat in Southeast Spain.
A new detailed study of all these speci- mens reveals the presence of small dorsal cirri on all segments. This character defin- itively separates these specimens from the genus Procerastea which only has dorsal cir- ri on the first setiger. This new genus is named Paraprocerastea and is overall very similar to Procerastea.
Two other genera of Autolytinae seem also to be related to Paraprocerastea: Alluau- della Gravier, 1905, from Madagascar and the Arabian Gulf, and Phyllosyllis Ehlers, 1897, from Antarctica. Paraprocerastea dif- fers from both genera by having a pharynx with a trepan. It differs further from Phyl- losyllis by having a different disposition of anterior segments and different setae, and from Alluaudella by having antennae and dorsal cirri of a different shape.
Observations and measurements were
made by means of a microscope with in- terference contrast optics. Drawings were made by means of a drawing tube. Length measurements exclude antennae and anal cirrl; width measurements were taken at proventricular level, excluding dorsal cirri and setae. The holotype and paratype from off Cabo de Creus are preserved in 70% eth- anol. The other paratype is in a permanent microscopical preparation made with glyc- erin jelly. All type material is deposited in the Museo Nacional de Ciencias Naturales de Madrid (MNCNM), Spain.
Family Syllidae Grube, 1850 Subfamily Autolytinae Rioja, 1925 Paraprocerastea, new genus
Diagnosis. —Body small, short, without segmentarial ciliation, small number of se- tigers. Prostomium relatively large, partial- ly covered by an occipital flap from tentacu- lar segment. Four dorsal eyes and two ventral eyespots; three thick, club-shaped antennae. Palps reduced, completely fused. Two pairs of tentacular cirri. Tentacular cirri and dor- sal cirri of first setiger club-shaped. Re- maining dorsal cirri minute, egg-shaped. Two subrectangular, relatively long, anal cirri. Simple setae of only two kinds: bay- onet-shaped setae and falcate setae. Phar- ynx slender, with a distal trepan.
VOLUME 102, NUMBER 4
Remarks.—Gidholm (1967:177) classi- fied the genera of Autolytinae in two natural groups. One group, composed of Proceraea Ehlers, 1864, Procerastea Langerhans, 1884, and Virchowia Langerhans, 1879 (=Um- bellysyllis Sars, 1869; according to Hartman 1959), is characterized by having internal parapodial glands, segmental ciliation re- duced or absent, bayonet setae of the thick type, and stolonization by anterior scissi- parity. The other group, composed of Au- tolytus Grube, 1850, and Myrianida Milne Edwards, 1845, is characterized by external parapodial glands, well developed segmen- tarial ciliation, bayonet setae of the slender type, and stolonization, if present, of a type other than anterior scissiparity. Gidholm did not include two, poorly known, genera of this subfamily, Phyllosyllis Ehlers, 1897 and Alluaudella Gravier, 1905. These two gen- era have unarmed pharynges, lack segmen- tal ciliation, and apparently lack bayonet- shaped setae. The parapodial glands have not been described; they could constitute a third natural group.
Another genus, Odontoautolytus Hart- mann-Schroder, 1979, has an uncertain po- sition between the subfamilies Eusyllinae and Autolytinae, because it is very similar to Odontosyllis Claparéde, 1863, but lacks ventral cirri.
Imajima & Hartman (1964) described another genus, Autosyllis, on the basis of a solitary Polybostrichus stolon. Consequent- ly, itis very difficult to know the characters of the atokous form and determine to which group it belongs. According to Gidholm, the specimen of Autosyllis is very similar to a male Polybostrichus of Umbellysyllis.
Paraprocerastea differs from Procerastea in having minute dorsal cirri on all seg- ments, whereas Procerastea has dorsal cirri only on setiger 1. However, the stolons of Procerastea have dorsal cirri on all setigers (Fauvel 1923:326, Allen 1921:135-137) similar to those of Paraprocerastea, and the segments in regeneration of adults have very small dorsal cirri (see Allen 1921:135, 140).
873
These observations indicate a very close re- lationship between both genera. Another dif- ference is the peculiar, slender, bayonet se- tae of Paraprocerastea, whereas the bayonet setae of Procerastea are thicker, with several short spines and a long filament, very sim- ilar to those of Proceraea (see Gidholm 1967: 207). On the other hand, the setae of Para- procerastea crocantinae are very similar to the simple setae of Procerastea halleziana Malaquin, 1893 (see Malaquin 1893:81, Fauvel 1923:326, Gidholm 1967:207, Alos 1988:244), P. nematodes Langerhans, 1884 (see Langerhans 1884:249, Fauvel 1923:326, Gidholm 1967:207) and P. australensis Hartmann-Schroder, 1987 (see Hartmann- Schroder 1987:65). These simple setae probably originated by shaft and blade fu- sion of compound setae; however, no com- pound setae or slightly fused setae have been found on Paraprocerastea crocantinae. Fi- nally, Procerastea has nuchal epaulettes and Paraprocerastea has an occipital flap.
The genus Phyllosyllis is also very similar to Paraprocerastea in having cylindrical to club-shaped anterior appendages, similar in length to body width, and very small, pyr- iform to egg-shaped dorsal cirri on the re- maining segments. However, Phyllosyllis has an unarmed pharynx, only compound setae and setae on the tentacular segment (Hart- man 1964:83); this last character is unusual in the family Syllidae and, in our opinion, another interpretation is possible; according to the drawings of Ehlers (1897) and Hart- man (1964), P. albida has only a pair of short tentacular cirri on a reduced tenta- cular segment, and the longer cirri are ac- tually the dorsal cirri of the second segment (first setiger).
Finally, Al/uaudella is also very close to Paraprocerastea but has shorter antennae, an unarmed pharynx, well developed dorsal cirri similar throughout the body, whereas the latter genus has long antennae, a phar- ynx with a trepan, and short, egg-shaped, dorsal cirri from the second setiger. On the other hand, P. crocantinae is similar to A.
874
madagascarensis in having an occipital flap and in the shape of the setae.
Type species.—Paraprocerastea crocan- tinae.
Etymology.—The generic name is re- ferred to the close relation with the genus Procerastea; gender is feminine.
Paraprocerastea crocantinae, new species Fig. 1
Material examined.—Cala Taballera, off Cabo de Creus, Gerona, Spain; calcareous concretions of Mesophyllum lichenoides and Lithophyllum expansum; 12 m depth; ho- lotype, complete, but left dorsal cirri of first setiger lacking. Isle Massa d’or, off Cabo de Creus; calcareous concretions of Lithophyl- lum expansum; 20 m depth; one paratype, complete, but median antenna lacking. Off Los Genoveses Inlet, Almeria, Spain; rhi- zomes of Posidonia oceanica; 3 m depth; one paratype, complete, but antennae lack- ing.
Etymology.—The name of the species is dedicated to the research vessel Crocantina of the Departamento de Biologia (Zoolo- gia), Universidad Autonoma de Madrid.
Description. — Body relatively short and thick, cylindrical, 3.4 mm length, 0.32 mm width, 33 setigers, without color marking, without segmentarial ciliation. Prostomium (Fig. 1A, B) proportionally large, circular; four large eyes in rectangular arrangement and two small ventral eyespots. Palps very reduced, practically nonexistent, complete- ly fused to one another. Three thick anten- nae, cylindrical to club-shaped, lateral an- tenna more than twice prostomium length, originating on anterior margin of prosto- mium; median antenna approximately three times longer than lateral antennae, origi- nating between anterior eyes (Fig. 1A). Ten- tacular segment ventrally reduced, dorsally prolonged in an occipital flap, covering pos- terior half of prostomium; two pairs of ten- tacular cirri, similar in shape to antennae, dorsal ones approximately half of lateral an- tennae length, ventral ones somewhat short-
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
er (Fig. 1A, B). Dorsal cirri of first setiger club-shaped, similar in length to dorsal ten- tacular cirri; dorsal cirri of remaining seg- ments minute, egg-shaped to oval (Fig. 1A, B). Parapodia short, conical, without ven- tral cirri (Fig. 1E). Setae simple and of only two kinds: bayonet-shaped setae and falcate setae; similar on all segments, without dor- soventral gradation in shape (Fig. 1E); para- podia each with one dorsal bayonet-shaped seta of a peculiar kind, relatively thin, end- ing in four short spines and a long filiform filament (Fig. 1F, J), and from three to five thick simple falcate setae, bifid, with two acute, curved, unequal teeth, provided with a subterminal thickened crown of a com- plete ring of small spines (Fig. 1G, H, K, L). Aciculae numbering 1-2 per parapo- dium, thin, with rounded tip (Fig. 11). In- ternal glands with granular material in each side of post-proventricular setigers, two to three per parapodium. Pygidium small, anal cirri longer than dorsal cirri, thick, rectan- gular to oval (Fig. 1D). Pharynx thin, with distinct sinuation, through about 3 seg- ments (Fig. 1A, B); trepan with 20 similar acutely triangular teeth (Fig. 1C). Proven- triculum shorter than pharynx, through about 1! segments, with 22 muscle cell rows (Fig. 1A, B). Reproduction unknown.
Remarks.—The only species of Autoly- tinae in the Mediterranean with only simple setae is Procerastea pori Ben-Eliahu, 1977 (Ben-Eliahu 1977, San Martin 1984); how- ever, this species has setae very different in shape, the trepan has only five large teeth, lacks of bayonet setae and nuchal epaulettes and, finally, the real number of tentacular cirri is unknown.
Key to the genera of Autolytinae
The genera Autosyllis and Odontoauto- lytus are not included in the key for the reasons given in the remarks.
1. Pharynx unarmed — Pharynx provided with atrepan ... 3 2. Two pairs of tentacular cirri. Occip-
VOLUME 102, NUMBER 4 875
Fig. 1. Paraprocerastea crocantinae, gen. and sp. n. A, Anterior end, dorsal view, holotype; B, Anterior end, dorsal view, paratype from Almeria; C, Trepan; D, Posterior end, dorsal view, paratype from Almeria; E, Middle-posterior parapodium, paratype from Almeria; F, J, Bayonet-shaped setae; G, H, K, L, Simple setae; I, aciculum. Scale: A, B, D: 0.13 mm; E: 20 um; C, F, G, H, I, J, K, L: 10 wm.
ital flap present. Dorsal cirri well de- small dorsal cirri on all segments)
veloped, similar throughout. Fal- ——........ Procerastea Langerhans, 1884 cate simple setae or compound setae — Dorsal cirri throughout in the ato-
.. 2 Alluaudella Gravier, 1905 Kkous Stocksow.) wustewmiil..cc...s 4 — One pair of tentacular cirri (?). No 4. Anterior appendages club-shaped, occipital flap. Dorsal cirri from se- dorsal cirri from setiger 2 minute, tiger 2 small, pyriform. Compound egg-shaped. Only simple setae. Oc- SEtGG . haces Phyllosyllis Ehlers, 1897 cipital flap present .... .. 2b-Jd.ale-
Ss» Dorsal cirti only om first setiger Of sence eccss Paraprocerastea n. gen.
the atokous stock (stolons with very — Anterior appendages cylindrical,
876
club-shaped or foliaceous, dorsal cirri longer, not egg-shaped. No oc- cipital flap; two nuchal epaulettes. Compound setae and simple bayo-
netesciac..0. Mee ee 5 5 Dorsalreirsi cylindrical, ci shee 6 — Dorsal cirri club-shaped or folia-
ECE OUS 2 cn cc Be Me A ee a
6. Bayonet setae as thick as shafts of compound setae. Without body Giliation’. ==. ©. Proceraea Ehlers, 1864
— Bayonet setae more slender than shafts of compound setae. Segmen- talciliary bands
ete cden We Autolytus Grube, 1850
7. Dorsal cirri foliaceous. Two short
ciliated nuchal epaulettes en Myrianida Milne Edwards, 1845
— Dorsal cirri club-shaped. Two very long, ciliated, foliaceous nuchal epaulettes .... Umbellisyllis Sars, 1869
eo © © © © © © © © © © © © ee 8
Acknowledgments
This paper has been partially supported by a grant of the “Comision Asesora de In- vestigacion Cientifica y Tecnologica”’ of the Spanish Ministry of Science and Education (project number Pr 84041) and the “Pro- grama de Bentos Cap de Creus”’ of the Fa- cultad de Biologia of the Universidad de Barcelona.
We would like to express our deep grat- itude to Dr. David Russell, Cove Corpo- ration, Lusby, Maryland, for his revision of the manuscript and valuable suggestions, as well as an anonymous referee. Dr. Mary E. Petersen, Zoologisk Museum, Copenhagen, provided us with some literature not acces- sible to us.
Literature Cited
Allen, E. J. 1921. Regeneration and reproduction of the syllid Procerastea.—Philosophical Trans- actions of the Royal Society (B) 211:131-177, pls. 11-16.
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Alos,C. 1988. Anélidos Poliquetos del Cabo de Creus (Alt Emporda). Unpublished doctoral thesis, Universidad de Barcelona, 838 pp.
Ben-Eliahu, M. N. 1977. Polychaete cryptofauna from rims of similar intertidal vermetid reefs on the Mediterranean coast of Israel and in the Gulf of Elat. Exogoninae and Autolytinae (Polychaeta, Errantia, Syllidae).—Israel Journal of Zoology 26:59-99.
Ehlers, E. 1897. Polychaeten. in Ergebnisse der Ham- burger magalhaenis-che Sammelreise, Ham- burg 3:1-148, pls. 1-9.
Fauvel, P. 1923. Faune de France, 5. Polychétes Er- rantes. Le Chevalier eds., Paris, 488 pp. Gidholm, L. 1967. A revision of the Autolytinae (Syl- lidae, Polychaeta) with special reference to Scan- dinavian species and with notes on external and internal morphology, reproduction and ecolo-
gy.—Arkiv for Zoologi 19(7):157-213.
Gravier, L. 1905. Sur un nouveau genre de Syllidien, Alluaudella nov. gen. madagascarensis nov. sp.— Comptes rendues du 6°™* Congrés International de Zoologie 6:372-376.
Hartman, O. 1959. Catalogue of the polychaetous
annelids of the world. Pt. I.—Allan Hancock
Foundation Occasional Papers 23:1—353.
1964. Polychaeta errantia of Antarctica.—
Antarctic Research series 3:1-131.
Hartmann-Schroder, G. 1979. in: Hartmann-Schro-
der, G. & Hartmann, G. eds.,: Zur Kenntnis des
Eulitorals der Australischen ktisten unter be-
sonderer Berucksichtigung der Polychaeten und
Ostracoden. Teil 2 und 3.— Mitteilungen aus dem
Hamburgischen Zoologischen Museum und In-
stitut 79:75-218.
. 1987. in: G. Hartmann-Schroder & G. Hart-
mann, eds., Zur Kenntnis des Eulitorals der aus-
tralischen kiisten unter besonderer Beriicksich- tigung der Polychaeten und Ostracoden. Teil. 13.—Mitteilungen aus dem Hamburgischen
Zoologischen Museum und Institut 84:27-66.
Imajima, M. & Hartman, O. 1963. The polychaetous annelids of Japan. — Allan Hancock Foundation Occasional Papers 26:1-—452.
Langerhans, P. 1884. Die Wiirmfauna von Madei- ra.— Zeitschrift fur Wissenschafthiche Zoologie 40(2):247-285.
Malaquin, A. 1893. Recherches sur les Syllidiens. Morphologie, anatomie, reproduction, devel- opement.— Mémoires de la Societé des Sciences et Arts de Lille 18, 447 pp.
San Martin, G. 1984. Estudio biogeografico, faunisti- co y sistematico de los poliquetos de la familia Silidos (Syllidae: Polychaeta) en Baleares. Tesis doctoral, Ediciones de la Universidad Complu- tense de Madrid, 529 pp.
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(GSM) Departamento de Biologia, Uni- dad de Zoologia, Laboratorio de Inverte- brados Marinos, Facultad de Ciencias, Uni- versidad Autonoma de Madrid, Canto
877
Blanco, E-28049, Madrid, Spain. (CA) De- partamento de Zoologia, Facultad de Cien- cias Biologicas, Universidad de Barcelona, Diagonal 645, E-0828, Barcelona, Spain.
PROC. BIOL. SOC. WASH. 102(4), 1989, pp. 878-886
FOUR NEW WEST ATLANTIC SPECIES OF TUBIFICOIDES (OLIGOCHAETA, TUBIFICIDAE)
Christer Erséus
Abstract. —Two species without hair setae are described from the Caribbean area: Tubificoides panamensis, from Panama and Bahamas, and T. inops, from Panama, Barbados and the Gulf of Mexico. The former is closely related to T. motei Brinkhurst, 1986 from Florida, but differs in setal numbers, shape of penis sheaths and morphology of spermathecal ducts. The latter lacks true penes and penis sheaths, but is included in Tubificoides as its genitalia in all other respects share apomorphies with those of all other species of that genus. Tubificoides bruneli, and T. crinitus, both with hair setae, are described from Lower St. Lawrence estuary (Québec, Canada) and the Gulf of Mexico, re- spectively. The first species has very stout spermatozeugmata, and ventral setae in anterior segments with upper teeth much longer than lower. Tubificoides crinitus is a small species occurring on the continental slope (down to at least about 550 m); its dorsal setae are all hair-like (alternating long and short within bundles). The generic definition of Tubificoides Lastockin, 1937 is emended,
partly to accommodate the aberrant 7. inops.
The marine tubificid genus 7ubificoides Lastockin, 1937 has been subject to recent revision by Brinkhurst (1985, 1986), with subsequent additions to the list of species by Rasmark & Erséus (1986), Helgason & Erséus (1987), Milligan (1987), Erséus & Milligan (1989), Erséus (1989), and Erséus & Davis (1989). The present contribution describes four additional species of this large genus. Two of them possess, the other two lack hair setae in their dorsal bundles.
The material comes from (1) the author’s own collection in Barbados (while at the Bellairs Research Institute of McGill Uni- versity); (2) material from R.V. Alpha Helix cruise “CARIB. I.” in 1977 (sorted by Smithsonian Oceanographic Sorting Cen- ter, SOSC, Washington, D.C.); (3) a collec- tion from Andros Island, Bahamas [cour- tesy Dr. M. L. Jones, United States National Museum of Natural History (USNM), Washington, D.C.]; (4) samples from off Pascagoula, Mississippi, in the northern Gulf of Mexico (courtesy Mr. M. R. Milligan,
Mote Marine Laboratory, Sarasota, Flori- da); (5) oligochaetes from the St. Lawrence estuary, Québec (courtesy Prof. P. Brunel, Université de Montréal, Montréal, Québec, Canada); and (6) benthos from a study of the Northern Gulf of Mexico Continental Shelf (LGL Ecological Research Associates, Inc., Bryan, Texas, for Minerals Manage- ment Service).
Material collected by the author was fixed in Bouin’s fluid, that collected by colleagues probably in formalin for most parts. Some specimens of 7. inops were longitudinally sectioned and stained in Heidenhain’s he- matoxylin, all other worms were stained in paracarmine and mounted whole in Canada balsam. The type series are all deposited in the USNM.
Tubificoides panamensis, new species Fig. 1
Holotype. —USNM_ 119870, whole- mounted specimen from N coast of
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Fig. 1.
f (fa VUE
879
,__-100nm =" pr
A-E, Tubificoides panamensis: A, Anterior seta; B, Posterior setae; C, Lateral view of spermatheca
and male genitalia in segments (IX)X—XI; D, Penis sheaths, specimens from Panama; E, Penis sheath, specimen from Bahamas; F, Tubificoides motei Brinkhurst: penis sheaths, specimens from Miami, Florida (coll. by author in 1977). Abbreviations: a, atrium; p, penis; pr, prostate gland; s, spermatheca; sz, spermatozeugma; vd, vas
deferens.
Panama, Caribbean Sea, 90°34'36’N, 78°43'12’W, O-4 m (30 Jun 1977); CARIB. I. Alpha Helix Stn. no. 23-333/149.
Paratypes. -USNM 119871-119873, three specimens from type locality. USNM 119874, one specimen from High Ridge Cay, Andros Island, Bahamas, 24°17'42’N, 77°45'12”"W, subtidal, associated with Tha- lassia (5 Mar 1966); M. L. Jones Stn. no. 56.
Description. —Length 4.7—-8.7 mm, up to 49 segments (some specimens not fully dif- ferentiated posteriorly). Width at XI 0.35- 0.44 mm. Prostomium small, triangular. Postclitellar segments often elongate. Body wall naked anteriorly, with cover of parti- cles tending to form papillae on small cu- ticular projections in postclitellar segments. Clitellum extending over XI—'/XII. Setae all bifid, 45-65 um long, two (rarely three) per bundle throughout most of body (gen- erally reduced to one per “‘bundle”’ in cli-
tellar region and near posterior end), with upper tooth thinner and slightly shorter than lower; teeth more diverging in postclitellar (Fig. 1B) than in anterior setae (Fig. 1A), with lower tooth occasionally much longer than upper. Ventral, and generally also dor- sal, setae absent from XI. Spermathecal pores in middle of X, between lateral lines and lines of ventral setae. Male pores in line with ventral setae in middle of XI. Pharyngeal glands in IV—-V. Esophagus somewhat enlarged in IX. Male genitalia (Fig. 1C) paired. Vas deferens 17-21 um wide, thin-walled and ciliated, about 1.5 times longer than atrium, entering latter subapically opposite to entrance of prostate gland. Atrium small and curved, histolog- ically tri- or bipartite (third, most ectal re- gion not always distinct), about 165-175 um long, entally 40-47 wm wide, at middle 23- 33 um wide, ectally 40-47 um wide, with very thin outer lining of muscles. Ental part
880
of atrium heavily granulated; granulation similar to that of prostate gland. A short, granulated portion generally also discern- ible in ectal part of atrium, at base of penis. Prostate gland small, without discrete stalk. Penis (Figs. 1C, p; D—-E) with cuticularized, conical penis sheath, with somewhat irreg- ular, wrinkled outline and large oval lateral opening. Penis 65—95 um long, basally 35- 55 um wide, at opening 16-28 um wide. Spermathecae (Fig. 1C, s) with slender, 80- 105 wm long, 23-20 wm wide ducts, and elongate ampullae; ectal half of ducts thick- ened and somewhat glandular, ampullae with slender spermatozeugmata.
Remarks. — Tubificoides panamensis ap- pears closely related to 7. motei Brinkhurst, 1986, known from Florida, but differs from that species by: (1) its lower number of setae (two, only rarely three, per bundle, as op- posed to three, or even four, per bundle in anterior segments of motei); (2) its less smoothly funnel-shaped penis sheaths, which exhibit very distinct lateral openings [penis sheath openings not as distinct in specimens of motei from Miami in my pos- session (see Fig. 1F); it should be noted, however, that according to Brinkhurst (1986: 1274, fig. 3), the narrow distal end of the penis sheath of motei is “possibly with an oblique opening”’]; and (3) the thickened ec- tal halves of the spermathecal ducts (ducts ectally provided with a very short, round and hollow swelling in my material of mo- tel).
Distribution and habitat. —Caribbean side of Panama, Bahamas. Subtidal to about 4 m depth. In Panama, the species was found together with 7. inops.
Tubificoides inops, new species Fig. 2
Holotype. —USNM_ 119875, whole- mounted specimen from N coast of Pana- ma. -| Caribbean,, «Sea, .<)09°34'362N: 78°43'12"W, 0-4 m (30 Jun 1977); CARIB I. Alpha Helix Stn. no. 23-333/149.
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Paratypes.—USNM 119876-119880, five whole-mounted specimens from type local- ity. USNM 119881-119885, one sectioned and four whole-mounted specimens from “‘the hole,’ at Holetown, St. James, Bar- bados, 5-9 m, muddy silt (28 Oct to 1 Nov 1979; C. Erséus).
Other material. — Author’s collection: two sectioned and seven whole-mounted spec- imens from Carlisle Bay, Barbados, 3.5—6.5 m, muddy silt and sand (24—25 Oct 1979; C. Erséus); and three sectioned and five whole-mounted specimens from Holetown (see paratypes). M. R. Milligan collection: two whole-mounted specimens (one of which sexually immature) from off Pasca- goula, Mississippi, northern Gulf of Mexi- co, 30°11'42”’N, 88°37'18”W, 13 m, medi- um to fine sand with silt and clay (22 Apr 1987).
Etymology. —The name inops is Latin for ““noor, lacking,”’ alluding to the species’ lack of penes.
Description. — Length 3.1-6.6 mm, 24—45 segments; width at XI 0.23—0.42 mm. Pro- stomium small, generally rounded and shorter than wide; posterior segments often elongate. Body wall naked anteriorly, in postclitellar segments with cover of fine par- ticles, generally aggregated into discrete, pointed papillae. Clitellum extending over XI—'»XII. Setae all bifid, 50-75 um long, two to five per bundle anteriorly, one to three per bundle in postclitellar segments, with upper tooth thinner than lower; upper tooth somewhat longer than lower and low- er tooth bearing indistinct subdental liga- ment in anterior setae (Fig. 2A), upper tooth shorter than lower in postclitellar setae (Fig. 2B). Ventral setae sometimes absent from XI. Spermathecal pores immediately ante- rior to ventral setae, in middle of X; male pores paired in line with ventral setae in middle of XI.
Pharyngeal glands in IV-V. Esophagus somewhat enlarged in IX. Male genitalia (Fig. 2D-E) paired; vas deferens 8-13 um wide, thin-walled and ciliated, about 1.5
VOLUME 102, NUMBER 4
50 um
881
‘ \ t ‘ &
! K
Y
2
100 um
Fig. 2. Tubificoides inops: A, Anterior seta; B, Posterior seta; C, Spermatheca, specimen from Barbados; D, Lateral view of spermatheca and male genitalia in segments X—XI, whole-mounted specimen from Panama; E, Lateral section through male genitalia in segment XI, sectioned specimen from Barbados. pp, pseudopenis; se,
seta; other abbreviations as in Fig. 1.
times longer than atrium, entering latter subapically opposite to entrance of prostate gland; atrium erect or curved, histologically tri- or bipartite (third, most ectal region not always distinct), 80-145 um long, entally 23—40 um wide, at middle 11-35 wm wide, ectally 21-39 um wide, with about 1-2 wm thick outer lining of muscles; ental part of atrium granulated in a way different from that of other parts, sometimes some distinct granules also discernible in ectal part of atrium, at base of pseudopenis. Prostate gland compact, small, with smooth outline. True penis absent, instead copulatory sac, 35-65 um long, 33-45 um wide, modified into an (eversible?) pseudopenis; inner sur- face of sac lined with cuticle and folded in an irregular way. Spermathecae (Fig. 2C; D,
s) slender, with 55-115 wm long, 20-34 um wide ducts, and thin-walled ampullae, ducts with roundish ectal swelling, ampullae con- taining slender spermatozeugmata.
Remarks. —The specimens from the type locality (Panama) have genitalia that are clearly smaller (Fig. 2D) than those of the worms from Barbados (Fig. 2C, E) and Gulf of Mexico.
The copulatory organs of 7. inops are dif- ferent from those of all congeners; they are pseudopenes rather than true, pendant penes. In fact, this feature disqualifies it for inclusion in the genus as currently defined (Brinkhurst & Baker 1979:1554): “. . . pen- ial structure bearing a penis sheath of vary- ing form.” However, its body wall papilla- tion, characteristic atria and spermathecae
882
appear to be good synapomorphic charac- ters shared with all “‘typical’’ members of Tubificoides. Then T. inops is either a very primitive member of the group (true penes had still not evolved when inops, or its ancestor, was split off from the rest of the lineage), or it is an advanced species within Tubificoides in the sense that its copulatory structure has “‘regressed” to an eversible pseudopenis. In the first case, T. Inops could be hypothesized as the plesiomorphic sister group to the others and could be regarded as a monotypic genus, separate from Tubifi- coides. In the second case, it should (cla- distically) be classified as a Tubificoides. Since most Tubificinae possess true penes, but do not possess the particular (advanced) atria fround in T. inops (and in Tubificoides as a whole), the latter alternative seems the most parsimonious one.
Accordingly, the generic definition of Tubificoides is modified below to accom- modate also 7. Inops.
Distribution and habitat. —Caribbean coast of Panama, Barbados, Gulf of Mexico. Subtidal silt and sand, to at least 13 m depth. In Panama, the species was found together with 7. panamensis.
Tubificoides bruneli, new species Fig. 3
Holotype. —USNM_ 119886, whole- mounted specimen from off Ile du Bic, Lower St. Lawrence estuary, Québec, Can- ada, 48°26'24”N, 69°00’00’W, 242 m, silt (31 Jub 19707, Ps Branel):
Paratypes. -USNM 119887-119890, four specimens from type locality.
Other material. —Author’s collection: three specimens from type locality.
Etymology.—The species is named for Prof. P. Brunel (Département de Biologie, Université de Montréal), who provided the material.
Description. —Length more than 7.6 mm, more than 63 segments (no complete spec-
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
imens available); width at XI 0.37—0.50 mm. Prostomium variable in shape, rounded or triangular. Body wall with some scattered particles in postclitellar segments. Clitellum extending over XI—'2XII. Dorsal bundles of IJ-VIII containing two to three bifid setae (Fig. 3A), up to 85 wm long and with par- allel, more or less equally long, slender teeth, and two hair setae, up to 200 um long; from IX, dorsal bundles containing one bifid or single-pointed seta (Fig. 3B), up to 85 um long, if bifid with teeth small and upper tooth longer than lower, and one hair, up to 145 um long. Ventral bundles of I—VIII with two to three bifid or single-pointed setae (Fig. 3C), up to 105 um long; if bifid, teeth slender and upper tooth much longer than lower; from IX, ventral setae single-point- ed, up to 85 um long, one to two per bundle; ventral setae absent from XI. Spermathecal pores anterior to, and at some distance from, ventral setae, anterior to middle of X; male pores in line with ventral setae, posterior to middle of XI.
Pharyngeal glands in IV—V. Esophagus not particularly enlarged in IX. Male genitalia (Fig. 3E) paired; vas deferens 16-22 um wide, thin-walled and ciliated, several times longer than atrium, but exact length not es- tablished. Entrance of vas into atrium not observed, but probably subapical as in all congeners; atrium curved, histologically tri- partite, 340-430 um long, entally 60-70 wm wide, at middle 50-65 um wide, ectally 50- 90 wm wide, with thin outer lining of mus- cles; ental and ectal parts of atrium granu- lated in a way different from that of middle part. Prostate gland large, lobed. Penis (Fig. 3D; E, p) with cuticularized, smooth, fun- nel-shaped penis sheath with ectal ends characteristically distended and curved (opening terminal), 105-120 um long, ba- sally 50-65 um wide, at distended ectal ends 28-42 um wide. Spermathecae (Fig. 3E, s) with 70-80 um long, 41—47 um wide ducts, and thin-walled, roundish ampullae; ducts with oval ectal swelling, lumen of which
VOLUME 102, NUMBER 4
(10
‘ iy A Ay cy v3 (4 CN 7 Wik, <= 200 um Fig. 3.
883
Tubificoides bruneli: A, Dorsal anterior seta; B, Dorsal postclitellar setae; C, Ventral anterior setae;
I), Penis sheaths; E, Lateral view of spermatheca and male genitalia in segments X—XI. Abbreviations as in
Figs. 1-2.
hollow near pore, ampullae with very stout, or (in one paratype only) slender, sperma- tozeugmata (Fig. 3E, sz).
Remarks.—Only two additional species in the genus have single-pointed ventral se- tae in postclitellar segments, viz., 7. pala- coleus Baker, 1983 and T. cuspisetosus Bak- er, 1983. The new species is, however, easily distinguished from both of these by its very characteristic penis sheaths; the penes are straight in the other two, those of JT. pala- coleus being provided with a strongly flared tip, those of 7. cuspisetosus being very slen- der. The shape of the penis sheaths (Fig. 3D), as well as the very long upper tooth of at least some of the anterior ventral setae (Fig. 3C), in fact distinguish 7. bruneli from all congeners. The similarities in the outline of the penis sheaths with that found in T. crenacoleus Baker, 1983 is superficial; whereas the tip of the sheath is properly
curved in J. bruneli, it is more or less straight, but bears a lateral, blunt projec- tion, in 7. crenacoleus.
The very stout spermatozeugmata found in all but one of the postcopulatory speci- mens in the available material (spermato- zeugmata of “normal,” slender type found in one worm only) are also noteworthy; spermatozeugmata with a similar shape have previously only been reported for 7. acu- leatus (Cook, 1970) (cf. Cook 1970: fig. 2D).
Distribution and habitat.—Known only from the type locality in the St. Lawrence estuary, Canada. Subtidal, silty sediment, 242 m depth.
Tubificoides crinitus, new species Figs. 4—5
Holotype. —USNM_ 119891, whole- mounted specimen from S of Louisiana,
884 A
10 pm B
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100 um
Fig. 4. Tubificoides crinitus, holotype: A, Ventral anterior seta; B, Lateral view of spermatheca and male genitalia in segments X—XI; C, Penis sheaths. Abbreviations as in Fig. 1.
27°43'07’"N, 92°53'18”W, 349 m (Minerals Management Service, Northern Gulf of Mexico Continental Shelf Study; 7 Jun 1985).
Paratypes. -USNM 119892-119895, four specimens: one from type locality; one from 27°43'23’N, 92°53'23"W, 335 m (7 Juneri 985)5 .ienes! fromat 27°50) 297.Ne 90°44'06”"W, 547 m (13 Jun 1985); and one (immature) from 28°34'42’”N, 90°14’06’W, 320 m (26 Nov 1983).
Etymology. —The name crinitus is Latin for “having hair, hairy,” referring to the species’ possession of hair-like setae throughout the dorsal bundles.
Description. —Length and segment num- ber unknown (specimens not complete), but worms appear very small. Width at XI 0.13- 0.22 mm. Prostomium small, blunt, at least partially retractable. Body wall naked an- teriorly, with some fine particles in postcli- tellar segments (but very few postclitellar segments remain on available worms). Cli- tellum poorly developed. Anterior dorsal bundles with one to three hair-like, single- pointed crotchets, 65-85 wm long, and one to three hair setae, 125-175 um long; post- clitellar dorsal bundles with two short hair- like setae, one slightly longer than the other;
ventral bundles with (one) two to three bifid setae (Fig. 4A), 60-80 um long, with very slender, thin and almost parallel teeth; teeth equally long. Ventral setae of XI absent. Spermathecal pores in line with (and near) ventral setae in middle of X; male pores in line with ventral setae slightly posterior to middle of XI.
Pharyngeal glands in IV-V. Esophagus somewhat enlarged in IX. Male genitalia (Figs. 4B, 5) paired; vas deferens 8-11 wm wide, thin-walled and ciliated, at least about three times longer than atrium, entering lat- ter subapically. Atrium of varying shape and length, histologically bi- or tripartite, 80- 185 um long, 30-42 um wide, with up to 3 um thick lining of (circular) muscles; ental part of atrium heavily granulated, remain- ing part less so. Prostate gland small, ap- pears to be attached to atrium at some dis- tance from apical end (see Fig. 5). Penis (Figs. 4B, p; C; 5, p) with cuticularized, cone- shaped penis sheath possessing large sub- terminal opening; penis 37-47 um long, ba- sally 23-26 um wide, ectally 16-19 um wide. Spermathecae (Fig. 4B, s) with 65-95 ym long, 28-36 wm wide ducts, and roundish to oval, small ampullae; ducts appear glan- dular, each with a roundish and hollow ectal
VOLUME 102, NUMBER 4
swelling; ampullae empty in holotype, with a few poorly preserved, slender, spermato- zeugmata in one paratype.
Remarks.—Two other species of Tubifi- coides lack bifid setae throughout the dorsal bundles: 7. aguadillensis Milligan, 1987, from Puerto Rico, and 7. uncinatus Hel- gason & Erséus, 1987, from off the east coast of the U.S.A. These species, as well as 7. crinitus, are also characterized by the some- what aberrant location of the prostate gland on the atrium; the prostate is not attached to atrium opposite to entrance of vas def- erens, but at a considerable distance from the apical end of the atrium (cf. Milligan 1987:fig. 4C; Helgason & Erséus 1987:fig. 6A). Both 7. aguadillensis and T. uncinatus differ, however, from T. crinitus by having bifid (ventral) setae with short, 1.e., “nor- mal,” teeth, and the penis sheaths of T. aguadillensis are evenly conical with a small terminal opening (penis sheaths of crinitus not as evenly tapering and with a large sub- terminal opening), those of 7. uncinatus are long and cylindrical and provided with a large subapical spur.
The spermathecae of the available ma- terial of 7. crinitus are smaller and with stouter ducts and ampullae than those of most other species of the genus. However, as most of the studied specimens are pre- copulatory, this feature should be used with some caution; other, postcopulatory, worms of T. crinitus may have larger spermathecae than those described here.
A species very similar to 7. crinitus is simultaneously described from a hydrocar- bon seep area in the northern Gulf of Mex- ico by Erséus & Milligan (1989).
Distibution and habitat.—Known only from the northern Gulf of Mexico. Subtidal on outer part of continental shelf, 320-547 m depth.
Tubificoides Lastockin, 1937
Definition (emended).—A species-rich group of marine and brackish-water tubif- icids (subfamily Tubificinae). Body wall na-
885
_0um , Pp PF Fig. 5. Tubificoides crinitus, paratype: male geni-
talia. Abbreviations as in Fig. 1.
ked, or with fine particles adhering to cu- ticle, often forming distinct papillae. In most heavily papillated species, prostomium often retractable. Hair setae absent or present in dorsal bundles; when present, generally to- gether with single-pointed, bifid or pectinate crotchets. Posterior dorsal setae, at least, frequently reduced to a single-pointed form barely distinguishable from the short hair setae where these are present. Modified gen- ital setae absent. Male pores paired, more or less in line with ventral setae in posterior part of segment XI. Spermathecal pores paired more or less in middle of X. Coelomocytes, if present, small and sparse, not of the “rhyacodriline-type.”’ Male efferent ducts paired in XI. Vas deferens ciliated, thin-walled, entering atrium sub- apically, generally opposite to large prostate gland. Rounded inner end of atrium cap- like, very heavily granulated and histolog- ically different from rest of atrium. Main body of atrium cylindrical, generally curved: terminal part often again histologically dis- tinct. Atrium terminating in a copulatory sac, which in all but one species contains a pendant penis bearing a cuticular penis sheath (in 7. inops the copulatory organ is
886
modified into a complex, probably eversi- ble, pseudopenis). Spermathecae consisting of a cylindrical duct, which often bears a distinct swelling near the ectal pore, and a round or oval-to-pear-shaped ampulla. Lat- ter with spindle-shaped, often very slender, spermatozeugmata in postcopulatory spec- imens.
Type species. — Tubificoides heterochaetus Lastockin, 1937 = T. swirencowi Jaroschen- ko, 1948 [not T. heterochaetus (Michaelsen, 1926)] (see Brinkhurst & Baker 1979).
Remarks. — This definition is an extended and partly revised version of the latest one by Brinkhurst & Baker (1979:1554). The shape of the atrium (with the cap-like apical part and the subapical entrance of the vas deferens) should probably be regarded as the single most important synapomorphy for the genus. This is why 7. inops, which lacks proper penes, is included (see further Remarks for 7. inops above).
Acknowledgments
I am indebted to Dr. F. Sander (former Director, Bellairs Research Institute, Bar- bados) for working facilities; to Dr. G. L. Hendler (former supervisor for benthos, SOSC), Dr. M. L. Jones, Mr. M. R. Milligan, Prof. P. Brunel, and Dr. L. H. Pequegnat (LGL Ecol. Res. Ass.), for providing spec- imens; to Ms. B. Lofnertz and Mrs. A. Falck- Wahlstrom, for technical assistance; to the Swedish Natural Science Research Council, for financial support; and Mr. M. R. Mil- ligan for very constructive criticism on the manuscript.
The LGL specimens reported on in this publication were collected through funding by the U.S. Department of Interior, Min- erals Management Service, Gulf of Mexico Regional OCS Office under Contract Num- ber 14-12-0001-30046 and 14-12-0001- 30212.
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Literature Cited
Brinkhurst, R.O. 1985. A further contribution to the
taxonomy of the genus Tubificoides Lastockin
(Oligochaeta: Tubificidae).—Canadian Journal
of Zoology 63:400-410.
1986. Taxonomy of the genus Tubificoides Lastockin (Oligochaeta, Tubificidae): species with bifid setae.— Canadian Journal of Zoology 64:1270-1279.
—.,& H.R. Baker. 1979. A review of the marine Tubificidae (Oligochaeta) of North America. — Canadian Journal of Zoology 57:1553-1569.
Cook, D. G. 1970. Peloscolex dukei n. sp. and P. aculeatus n. sp. (Oligochaeta, Tubificidae) from the North-west Atlantic, the latter being from abyssal depths.— Transactions of the American Microscopical Society 88:492—-497.
Erséus, C. 1989. Marine Oligochaeta of Hong Kong. Pp. 000-000 in B. Morton, ed., Proceedings: Second International Marine Biological Work- shop on the Marine Flora and Fauna of Hong Kong and Southern China, Hong Kong 1986. Vol. 1. The University of Hong Kong Press, Hong Kong (in press).
—, & D. Davis. 1989. The marine Tubificidae (Oligochaeta) of Hawaii.—Asian Marine Biol- ogy (in press).
—, & M. R. Milligan. 1989. Three new species of Tubificidae (Oligochaeta) from an oil seepage area on the continental slope of the northern Gulf of Mexico.— Proceedings of the Biological Society of Washington 102:887-893.
Helgason, G. V., & C. Erséus. 1987. Three new species of Tubificoides (Oligochaeta, Tubificidae) from the North-west Atlantic and notes on geographic variation in the circumpolar T. kozloffi.—Sarsia 72:159-169.
Milligan, M. R. 1987. Marine Tubificidae (Oligo- chaeta) from Puerto Rico with descriptions of two new species, Tubificoides aguadillensis and Heterodrilus paucifascis.—Proceedings of the Biological Society of Washington 100:480-489.
Rasmark, B., & C. Erséus. 1986. A new species of Tubificoides Lastochkin (Oligochaeta: Tubifici- dae) from Bermuda and Bahamas.—Proceed- ings of the Biological Society of Washington 99: 612-615.
Zoo-tax, Swedish Museum of Natural History, Stockholm, and (postal address): Department of Zoology, University of Go- teborg, Box 25059, S-400 31 Goteborg, Sweden.
PROC. BIOL. SOC. WASH. 102(4), 1989, pp. 887-893
THREE NEW SPECIES OF TUBIFICIDAE (OLIGOCHAETA) FROM AN OIL SEEPAGE AREA ON THE CONTINENTAL SLOPE OF THE NORTHERN GULF OF MEXICO
Christer Erséus and Michael R. Milligan
Abstract. —Limnodriloides olearius (subfamily Limnodriloidinae), Tubifi- coides paracrinitus, and T. pequegnatae (Tubificinae) are described from a hydrocarbon seep area of vestimentiferan growth at 540 m depth south of Louisiana. The tubificids are associated with vestimentiferans, mussels and bacterial mats. Limnodriloides olearius belongs to the winckelmanni-group within the genus, but is unique by its possession of transverse patches of epidermal glands ventrally in segments III—X. Tubificoides paracrinitus is closely related to T. crinitus Erséus, 1989, but is larger and possesses postclitellar body wall papillae, a greater number of setae, and different penes and spermathecae; T. pequegnatae resembles 7. bakeri Brinkhurst, 1985 in its setal distribution, but differs in the morphology of its penis sheaths.
Five samples of oligochaetes, from an area of natural hydrocarbon seepage on the Gulf of Mexico continental slope about 128 km south of Louisiana, were sent to the senior author for identification. They represented three new species of Tubificidae (one species of Limnodriloides Pierantoni, two of Tubifi- coides Lastockin) described in the present paper.
The samples were collected from the sub- mersible Johnson Sea Link I, Dive No. 1878 (28 Sep 1986), at “Bush Hill,” an area of thick growths of tube worms, mussel beds and bacterial mats in Blocks 184 and 185 of the Green Canyon offshore oil leasing area (cf. Brooks et al. 1987); oil leasing blocks being set by the U.S. Department of Inte- rior, Minerals Management Service. The worms were provided by LGL Ecological Research Associates, Inc. (Bryan, Texas). Only two or three sexually mature individ- uals of each species were found. The spec- imens were stained in paracarmine and mounted whole in Canada balsam, and have been deposited as type specimens in the U.S. National Museum of Natural History
(USNM), Smithsonian Institution, Wash- ington, D.C.
Subfamily Limnodriloidinae
Limnodriloides olearius, new species Fig. |.
Holotype. —USNM 119904, 2.4 mm long, consisting of first 15 segments only, from S of Louisiana, 27°46'56”N, 91°30'20’” W, 540 m (28 Sep 1986), in area of vestimentiferan growth.
Paratype. —USNM 119905, from type lo- cality.
Etymology. —The epithet olearius is Lat- in for “‘of oil,’ here alluding to the species’ occurrence in an area of oil seepage.
Description. —Length more than 4.2 mm, more than about 35 segments (no specimen complete); width at XI in whole-mounted, compressed specimens about 0.25 mm. Prostomium rounded triangular. Clitellum not distinct in available specimens. Very distinct, transverse, elongated patches of epidermal glands present ventrally in most of preclitellar segments, at least in IJJ—-X
888
Figs.
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palit Ix X
Limnodriloides olearius, n. sp.: A, Ventral view of anterior end of worm (semi-schematical); B, Free-
hand drawing of somatic seta; C, Three (probably all) parts of one atrium, holotype; D, Spermatheca and spermathecal seta. Abbreviations: aa atrial ampulla; ad atrial duct; b brain; d oesophageal diverticula; o oes- ophagus; pe patch of epidermal glands; pg pharyngeal glands; pp pseudopenis; ppa prostatic pad; pr prostate gland; s spermatheca; sp spermathecal pore; ss spermathecal seta; Roman numerals denoting segment numbers.
(Fig. 1A, pe); bundles of ventral setae and (in X) spermathecal pores located within these glandular patches. Somatic setae (Fig. 1B) bifid, with upper tooth as long as but thinner than lower; bifids 40-70 um long, about 2.5 wm thick, (two) three (four) per bundle anteriorly, two per bundle in po- stclitellar segments. In holotype and para- type, one spermathecal seta (Fig. 1D, ss), partly enclosed in narrow (glandular?) sac, present immediately posterior to one of the two spermathecal pores in X; at other side of X setal sac empty (spermathecal seta lacking; see Fig. 1A). Spermathecal seta, sin- gle-pointed, shaped like a walking-stick, about 100-110 um long, about 3.5 um at node, with node at about middle, and with part ectal to node grooved; large gland as- sociated with sac of spermathecal seta not observed, but may be present (as in some congeners; see, e.g., Erséus 1982:figs. 15, 20—- 21). Penial setae absent; male pores paired, somewhat ventral to lines of ventral setae, in middle-to-posterior part of XI; sper-
mathecal pores paired, in line with ventral setae, in middle of X.
Pharyngeal glands (Fig. 1A, pg) present in IV—V. Large, somewhat conical esopha- geal diverticula (Fig. 1A, d) present in IX. Male genitalia (Fig. 1C) paired, not well pre- served in available material; vas deferens 9-14 um wide, but length and junction with atrium not established. Atrial ampulla 75- 80 um long, about 45 wm wide, with ectal half filled with large, conspicuous prostatic pad; muscular lining of ampulla 2-3 wm thick in paratype, thinner in holotype (Fig. 1C); prostate gland lobed. Atrial duct (bro- ken into pieces in holotype; not visible in paratype) slender, about 135 um long, 21- 28 um wide, granulated for most parts, ter- minating in simple pseudopenis with some- what folded inner wall; inner structure of pseudopenis not clear, but no distinct pseu- dopenial papilla appears to be present. Sper- mathecae (Fig. 1D, s) large, consisting of very short, triangular ducts, and oval, up to about 170 um long, 75 um wide, ampullae;
VOLUME 102, NUMBER 4
sperm arranged as distinct, curved bundles in spermathecae; possibly they are (poorly preserved) spermatozeugmata.
Remarks. —The function of the conspic- uous patches of epidermal glands in the an- terior segments is unknown. It appears like- ly, however, that they are in some ways related to reproduction as such patches are not developed in four juvenile specimens of Limnodriloides found at the type locality, and presumably belonging to the same species. Patches of epidermal glands have not been reported for Limnodriloides be- fore, and they are in fact distinguishing L. olearius from all other members of the subfamily Limnodriloidinae, but similar glands are present dorsally (largely in post- clitellar segments) in some species of Bathy- drilus Cook, in the subfamily Phallodrili- nae (Erséus 1986:figs. 9A—B).
The new species is a member of the win- kelmanni-group within Limnodriloides, i.e., the species with spermathecal setae (Erséus 1982). Both type specimens lack the sper- mathecal seta at one side of segment X, but this asymmetric arrangement is not neces- sarily a specific character; other species within the group occasionally lack one or both spermathecal setae (e.g., L. victoriensis Brinkhurst & Baker, 1979; see Brinkhurst & Baker 1979; Erséus 1982).
Among those species in the winckelman- ni-group that have: (1) a pair of esophageal diverticula in segment IX, and (2) well sep- arated male pores, L. olearius appears most closely related to L. barnardi Cook, 1974, acommon species in the Northwest Atlantic and the Caribbean, and also known from the Pacific coast of Mexico; both species have elongate atrial ampullae with the pros- tatic pads located in the ectal half of these ampullae (Fig. 1C; Erséus 1982:fig. 13A). Limnodriloides olearius is distinguished from the latter by its shorter spermathecal ducts, and its very simple pseudopenes. In L. barnardi, the atrial ducts terminate in distinct pseudopenial papillae (Erséus 1982: fig. 13 A—B), which in fact were erroneously interpreted as proper penes by the original
889
author (Cook 1974). The male genitalia are not very well preserved in the material of L. olearius, but there 1s no indication of such discrete papillae in the copulatory organs (Fig. 1C, pp).
Distribution and habitat.—Only known from the type locality, N Gulf of Mexico. Upper continental slope, 540 m depth.
Subfamily Tubificinae
Tubificoides paracrinitus, new species Figo 2
Holotype. —USNM_ 119901, whole- mounted specimen from south of Louisi- ana, 27°47'01’N, 91°30'03”W, 582.5 m (28 Sep 1986), in area free of vestimentiferan growth, but not very far from the hydro- carbon seep.
Paratypes. —USNM 119902, 119903, two whole-mounted specimens from 27°46’ 56”N, 91°30’20”W, 540.1 m (28 Sep 1986), in area of vestimentiferan growth.
Etymology. —Named paracrinitus for its resemblance to T. crinitus Erséus, 1989.
Description.—Length of holotype 27.1 mm, 53 segments; paratypes not complete; width at XI in whole-mounted, compressed specimens 0.25-0.27 mm. Prostomium pointed triangular. Body wall naked ante- riorly, but covered with very small papillae in most of postclitellar segments (papilla- tion not beginning immediately behind cli- tellar region). Clitellum poorly developed. Anterior dorsal bundles with up to six hair- like, generally single-pointed, crotchets, 55— 95 um long (occasionally such a seta bifid with minute teeth), alternating with about the same number of long hair setae, 200— 250 um long; postclitellar dorsal bundles similar to anterior ones, but setal number and length generally not as great; anterior ventral bundles with two to four bifid setae (Fig. 2A), 65-80 um long, with very slender, thin and almost parallel teeth, upper tooth tending to be longer than lower; postclitellar ventral bundles with two to three setae, sim- ilar to anterior ventrals or sharply single- pointed. Ventral setae of XI, and sometimes
890
5
Fig. 2. D, Penis sheaths; E, Tubificoides crinitus Erséus, penis sheaths. Abbreviations: a atrium; p penis; pr prostate gland; sz spermatozeugma; vd vas deferens.
of X, absent. Spermathecal pores in line with (and near) ventral setae (if present) in middle of X; male pores in line with ventral setae slightly posterior to middle of XI.
Pharyngeal glands in IV-V. Esophagus somewhat enlarged in IX. Male genitalia (Fig. 2B) paired; vas deferens 14-18 um wide, thin-walled and ciliated, at least about three times longer than atrium, entering lat- ter subapically. Atrium cylindrical, up to about 290 um long, 33-52 um wide, his- tologically tripartite, with up to about 2 um thick lining of muscles; ental part of atrium more heavily granulated than remaining part; prostate gland lobed, attached to atrium opposite to entrance of vas deferens. Penis (Fig. 2D) with conical, thimble-shaped pe- nis sheath, 70-90 um long, basally 40-47 um wide, ectally 27-35 wm wide, with in- distinct, terminal or somewhat subtermi- nal(?) opening. Spermathecae (Fig. 2C) with ducts 130-210 um long, 35-50 um wide, somewhat glandular and with a hollow ectal swelling; ampullae and spermatozeugmata slender in postcopulatory specimens; sperm trap present(?).
Remarks. — Tubificoides paracrinitus is
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A-D, Tubificoides paracrinitus, n. sp.: A, Ventral anterior seta; B, Male genitalia; C, Spermatheca;
closely related to J. crinitus Erséus, 1989, which is simultaneously described from non- seep areas in the same part of the Gulf of Mexico (Erséus 1989b), but it differs from the latter in several respects: (1) it is dis- tinctly larger (0.25-—0.27 mm wide at seg- ment XI, as opposed to 0.13-—0.22 mm for crinitus; length of latter unknown); (2) it has discrete body wall papillae in postclitellar segments (crinitus has fine particles scat- tered over postclitellar segments, but no pa- pillae are formed); (3) it has up to six crotch- ets and about the same number of hairs in the dorsal bundles (setae only half as many in crinitus); (4) occasionally its dorsal crotchets are bifid (well visible in segment V of holotype)(all dorsal crotchets of cri- nitus hair-like and single-pointed); (5) its penis sheaths (Fig. 2D) appear to have smaller and not as lateral openings, as those of the sheaths of crinitus (Fig. 2E, shown here for comparison); and (6) its much more slender spermathecae (Fig. 2C) (the sper- mathecal ampullae of crinitus are small and oval). Some of these differences could per- haps, per se, be regarded as intraspecific rather than interspecific, but taken together
VOLUME 102, NUMBER 4
7
( ( ( L We”
W/
Fig. 3. ventral seta; D, Posterior ventral seta; E, Male genitalia; F, Spermatheca. Abbreviations as for Fig. 2.
they certainly indicate that the two forms are separate species.
Distribution and habitat.—Known only from the Gulf of Mexico. Upper continental slope, 540-582 m depth.
Tubificoides pequegnatae, new species Fig. 3
Holotype. —USNM_ 119896, whole- mounted specimen from 27°46'56’N, 91°30'20” W, 540.1 m (28 Sep 1986), in area of vestimentiferan growth.
Paratypes.—USNM 119897-119900, four whole-mounted specimens (three of which sexually immature) from type locality.
Etymology.—Named for Dr. Linda H. Pequegnat, who very kindly provided the present material.
Description. — Length of holotype 3.5 mm, 13+ segments (posterior end partially re- generating); sexually mature paratype 2.6 mm, 17 segments, but incomplete. Width at XI in whole-mounted, compressed spec- imens, 0.32—0.46 mm. Prostomium round- ed, somewhat broadly triangular. Body wall naked anteriorly, but coated with fine par-
891
% ey pe,
are
\Y) "a.
ean
Saat
0 Be Ae 5 1}
Tubificoides pequegnatae, n. sp.: A, Anterior dorsal bifid seta; B, Posterior dorsal bifid seta; C, Anterior
ticles with a tendency to form papillae in postclitellar segments. Clitellum poorly de- veloped. Anterior dorsal bundles with (one) two to three bifid setae, 45-75 um long, upper slightly longer than lower (Fig. 3A), and (one) two to three hair setae, 115-140 um long; postclitellar dorsal bundles gen- erally with two bifid setae, upper tooth long- er and thinner than lower (Fig. 3B), and occasionally in a few segments immediately posterior to clitellum one hair seta may be present; anterior ventral bundles with three to four bifid setae, 60-75 um long, upper tooth tending to be longer and thinner than lower (Fig. 3C); postclitellar ventral bundles with two setae (Fig. 3D), similar to anterior ventrals. Ventral setae of X and XI absent. Spermathecal and male pores in line with ventral setae in middle of X and XI, re- spectively.
Pharyngeal glands in IV—VI. Male geni- talia (Fig. 3E) paired; vas deferens about 20 um wide, thin-walled and ciliated, at least three times longer than atrium, entering lat- ter subapically. Atrium cylindrical, up to about 235 um long, histologically tripartite, with up to 2 um thick lining of muscles,
892
ental part of atrium more heavily granulat- ed than remaining part, about 70 um wide, middle part about 40 um wide. Prostate gland lobed, attached to atrium opposite to entrance of vas deferens; penis with elon- gate, funnel-shaped sheath, 75—105 um long, basally 37 um wide, with somewhat sub- terminal opening. Spermathecae (Fig. 3F) with ducts 130 wm long, 20-25 um wide, ectal region slightly bulbous with a thick- ened muscular covering; ampullae ovoid, 180-190 um long, 135-150 um wide; sper- matozeugmata long and slender; sperm trap not seen.
Remarks. —This species is similar to the Northeast Pacific 7. bakeri Brinkhurst, 1985 with regard to its setal distribution; no other described species in the genus with hairs and bifid setae anteriorly lacks the hair setae in the posterior segments. Tubificoides bakeri has a slightly greater number of anterior dorsal setae (up to four bifids, plus as many hairs) than 7. pequegnatae, and its atria are distinctly smaller than those of the new species, but the most important difference between the two species is the shape of the penis sheaths: the penis of 7. bakeri has a bulbous tip (Brinkhurst 1985:fig. 7B), whereas the penis of the new species con- tinuously tapers ectally (see Fig. 3E).
Discussion
Seven species of Tubificidae were previ- ously recorded from continental slope depths (about 300 m and deeper) in the northern Gulf of Mexico (Erséus 1988, 1989a, b): Phallodrilus constrictus Erséus, P. grasslei Erséus, P. vescus Erséus, Bathydrilus con- nexus Erséus, B. longiatriatus Erséus, Lim- nodriloides monothecus Cook and Tubifi- coides crinitus Erséus. The samples from the “Bush Hill’ hydrocarbon seeps yielded three separate species, which is noteworthy con- sidering that the area is located in the same part of the Gulf. It thus seems possible that the three new species, or at least some of them, are endemic to these hydrocarbon seep
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
situations. The extensive bacterial mats (often containing separate phase oil lying between them) probably provides a rich food supply to the associated macrofauna (see Brooks et al. 1987), including these (possi- bly specialized) tubificids.
Acknowledgments
The LGL specimens reported in this pub- lication were collected through funding by the U.S. Department of Interior, Minerals Management Service, Gulf of Mexico Re- gional OCS Office under Contract Number 14-12-0001-30046 and 14-12-0001-30212. We are indebted to Dr. Linda H. Pequegnat (LGL) for providing this interesting mate- rial, and for valuable information about the collecting site, and to Ms. Barbro Lofnertz and Mrs. Aino Falck-Wahlstrom for tech- nical assistance.
Literature Cited
Brinkhurst, R.O. 1985. A further contribution to the taxonomy of the genus Tubificoides Lastockin (Oligochaeta, Tubificidae).—Canadian Journal of Zoology 63:400-410.
—., & H.R. Baker. 1979. A review of the marine Tubificidae (Oligochaeta) of North America. — Canadian Journal of Zoology 57:1553-1569.
Brooks, J. M., etal. 1987. Hydrates, oil seepage, and chemosynthetic ecosystems on the Gulf of Mex- ico slope: an update. — Eos, Transactions, Amer- ican Geophysical Union 68(18):498-499.
Cook, D. G. 1974. The systematics and distribution of marine Tubificidae (Annelida: Oligochaeta) in the Bahia de San Quintin, Baja California, with descriptions of five new species. — Bulletin of the Southern California Academy of Sciences 73:126-140.
Erséus, C. 1982. Taxonomic revision of the marine
genus Limnodriloides (Oligochaeta: Tubifici-
dae). — Verhandlungen des naturwissenschaftli-
chen Vereins in Hamburg (Neue Folge) 25:207-
PA iS i
1986. Marine Tubificidae (Oligochaeta) at Hutchinson Island, Florida.— Proceedings of the Biological Society of Washington 99:286-315.
1988. Deep-sea Tubificidae (Oligochaeta) from the Gulf of Mexico.—Proceedings of the
Biological Society of Washington 101:67-71.
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. 1989a. Phallodrilus vescus, new species (Oli- gochaeta, Tubificidae) from the Gulf of Mexi- co.—Proceedings of the Biological Society of Washington 102:134-136.
1989b. Four new West Atlantic species of Tubificoides (Oligochaeta, Tubificidae).—Pro- ceedings of the Biological Society of Washington 102:877-886.
893
(CE) Zoo-tax, Swedish Museum of Nat- ural History, Stockholm, and (postal ad- dress): Department of Zoology, University of Goteborg, Box 25059, S-400 31 Gote- borg, Sweden; (MRM) Mote Marine Lab- oratory, 1600 City Island Park, Sarasota, Florida 33577, USA.
PROC. BIOL. SOC. WASH. 102(4), 1989, pp. 894-900
EULIMNADIA OVILUNATA AND E. OVISIMILIS, NEW SPECIES OF CLAM SHRIMPS (CRUSTACEA, BRANCHIOPODA, SPINICAUDATA) FROM SOUTH AMERICA
Joel W. Martin and Denton Belk
Abstract.—Two new species of the clam shrimp genus Eulimnadia are de- scribed from South America. Eulimnadia ovilunata, a species with spherical eggs bearing circular crater-like depressions, is described from Argentina, and Eulimnadia ovisimilis, a species with stout cylindrical eggs that closely resemble those of the North American species E. texana (Packard), is described from
Paraguay.
Species of the clam shrimp genus Eulim- nadia Packard (sometimes considered syn- onymous with Limnadia; see Webb & Bell 1979) in North, Central, and South America were the subject of two recent reviews by Belk (1989) and Martin (1989). Although many traditionally employed characters were shown by those authors to be variable and unreliable, an additional character, morphology of the external egg shell, ap- pears to be conservative and species-spe- cific. In his comparison of described Central and South American species, Martin (1989) examined two South America forms that did not conform to any known species. Those species, described from samples in the collections of Denton Belk in San An- tonio, Texas, were referred to by Martin as Eulimnadia sp. A (DB 305) and Eulimna- dia sp. B (DB 632). In this paper we describe the two new species.
Materials and Methods
Illustrations were made from ethanol- preserved specimens with a Wild M-SAPO stereoscope with camera lucida. All speci- mens are catalogued in the Natural History Museum of Los Angeles County, abbrevi- ated LACM. The abbreviation DB refers to the cataloging system of the private collec-
tion of the junior author in San Antonio, Texas; lots from that private collection formed the basis for this report.
The highway designations in Catamarca Province, Argentina, changed between the time Arthur Hulse made clam shrimp col- lections in 1973-1974 and today. We cite the current nomenclature according to the 14 April 1986 map published by Automovil Club Argentino in addition to including the location information originally supplied by Dr. Hulse (in parentheses).
Eulimnadia ovilunata, new species Figs. 1, 3A—C
Eulimnadia sp. A.—Martin, 1989, fig. 5d (eggs).
Material. —DB 303, paratypes, LACM 74- 107.1, 3 males, 45 females (22 ovigerous), Argentina, Catamarca Province, highway 46 W of Andalgala in flood plain of Rio Ama- nao (formerly Route 62, km 1508, at time of collection), 4 Mar 1974, coll. A. Hulse; DB 304, paratypes, LACM 74-108.1, 11 fe- males (1 ovigerous), Argentina, Catamarca Province, highway 46 S of Andalgala (for- merly Route 1, km 104, at time of collec- tion), 8 Mar 1974, coll. A. Hulse; DB 305, holotype female (ovigerous), LACM 73- 180.1; paratypes, LACM 73-180.2, 19 fe-
VOLUME 102, NUMBER 4 895
Fig. 1. Eulimnadia ovilunata: A, Right valve of holotype female; B, Holotype female, right valve removed, with egg enlarged above; C, Frontal region of holotype female; D, Rostrum and eye of one of three poorly preserved males; E, Caudal region of holotype female; F, Caudal region of male. Scale bars indicate 1.0 mm.
males (15 ovigerous), Argentina, Catamarca Measurements. —Male carapace 5.5 to 6.0 Province, highway 46 S of Andalgala (for- mm length, 3.5 to 4.1 mm height (but see merly Route 1, km 45, at time ofcollection), Remarks below) (n = 3; DB 303, no males 30 Dec 1973, coll. A. Hulse, 1 female de- in other collections). Female carapace 5.1 stroyed in SEM preparation (Fig. 3). to 6.8 mm length, 3.3 to 4.7 mm height (n
896 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
\ NY NM
we :
NY WIR BERS ;
\\w MIN
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Ny oe a
Yt AINE: AN
Fig. 2. Eulimnadia ovisimilis: A, Carapace (right valve) of holotype female; B, Holotype female with right valve removed; C, Allotype male with right valve removed; D, Head region of holotype female; E, Head region of allotype male; F, Second male clasper of allotype male; G, Caudal region of allotype male; H, Caudal region of holotype female. Scale bars indicate 1.0 mm.
= 76, DB 303, DB 304, DB 305; smallest apaces all shriveled; condition indetermi- and largest females both ovigerous). nate (see below).
Carapace. — Female carapace (Fig. 1A, B) Head region.—Female (Fig. 1C) with broadly oval, with hinge border domed and broadly triangular rostrum; male (Fig. 1D) with three or four lines of growth. Male car- with very prominent acute rostrum.
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Fig. 3. Scanning electron micrographs of eggs of Eulimnadia ovilunata (A—C) and E. ovisimilis (D-F): A, Eggs of E. ovilunata (paratype) from Catamarca, Argentina (DB 305), x 200; B, Single egg of E. ovilunata (from same female as in A), x520; C, Higher magnification of crater-like depression of egg in B showing mound at bottom of depression (arrow), <x 1,180; D, Eggs of E. ovisimilis (paratype) from Paraguay (DB 632), x75; E, Cluster of eggs of E. ovisimilis on epipod of female, x 120; F, Higher magnification of eggs shown in E showing nature of end pieces, x 250.
Antennae. —First antennae pseudoseg- on ventral border of each segment; number mented with aesthetascs on anterior border of segments varies from seven to nine. of each lobe. Second antennae natatory, with Male thoracopods.—Not examined (see spines on dorsal border and plumose setae _ below).
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Caudal region.—Female (Fig. 1E) and male (Fig. 1F) caudal regions similar, with 8 to 10 stout downward curved spines on posterior borders and with telsonal fila- ments arising from between second and third such spines.
Eggs.—Spherical, with oval or circular depressions that appear fringed with small fingerlike projections of the shell or tertiary envelope, each depression with a slightly convex and relatively smooth floor (Figs. 1B, 3A-C).
Type locality. —Argentina, Catamarca, highway 46, 45 km S of Andalgala.
Range. —Known from three localities in Catamarca, Argentina.
Etymology. —From the Latin ‘“‘luna’’ (moon) and “‘ova’”’ (egg), because the depres- sions on the eggs are reminiscent of craters on the lunar surface.
Remarks.—Eulimnadia ovilunata does not differ appreciably from many other species of the genus except by virtue of the egg morphology. Of the South American species described or reviewed by Martin (1989), only the egg of E. brasiliensis Sars is spherical. However, the E. brasiliensis egg lacks the minute projections fringing each indentation, and the indentations do not ap- pear as regularly formed and do not bear the clearly defined oval “‘floor’’ of the crater (compare Martin’s (1989) fig. 4c to his fig. 5d and to Fig. 3A, this paper). The North American species Eulimnadia diversa Mat- tox, E. agassizii (Packard), and E. antlei (Mackin) also have spherical eggs, but the eggs of all three of these species clearly differ from E. /unaova eggs under high magnifi- cation (see Belk 1989). The eggs of Eulim- nadia antillarum (Baird), a species that oc- curs in North and South America, remain undescribed, but the caudal region of that species is nearly devoid of spines (see Mar- tin 1989:fig. SA), whereas the caudal region in E. ovilunata bears relatively large, well developed spines in both sexes (Fig. 1E, F).
The only males in the series are in very poor condition, apparently having com-
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pletely dried out at some time in the past. Structures traditionally described for males could not be ascertained, and the illustra- tion of the male rostral region (Fig. 1D) should be verified when males in good con- dition are collected. Because of the condi- tion of the males no allotype was designated among the three male paratype specimens in DB 303.
Eulimnadia ovisimilis, new species Figs. 2, 3D-F
Eulimnadia sp. B.— Martin, 1989, fig. Se-g (eggs).
Material. —DB 632, holotype female (ovigerous), LACM 84-204.1; allotype male, LACM 84-204.2; paratypes (2 males, 48 fe- males (21 of which are ovigerous)), LACM 84-204.3, Paraguay, Chaco Departamento, Parque Nacional Defensores del Chaco, Tribo Nuevo, “encontrado en regiones ba- jas de laguna recién inundada,”’ from shal- low regions with submerged grass about one week after heavy rains filled a formerly dry pond-marsh (Terry Bonace, personal letter to D. Belk), 23 Nov 1984, coll. T. Bonace and D. Drenner.
Measurements. — Male carapace 6.9 to 7.5 mm length, 4.5 to 4.7 mm height. Females (including ovigerous females) 6.0 to 10.0 mm length, 4.2 to 6.9 mm height.
Carapace. —Femaie carapace (Fig. 2A, B) broadly oval, distinctly elevated along dor- sal border, with four lines of growth. Male carapace (Fig. 2C) usually smaller, oval but not elevated along dorsal border, instead somewhat flattened along hinge line, with two or three lines of growth.
Head region. — Female (Fig. 2D) with very short bluntly rounded rostrum; male (Fig. 2E) with short but acute rostrum.
Antennae. —Similar to that described for E. ovilunata (and many other members of the genus); second antennae with eight or nine segments.
Male thoracopods.—Typical for the ge- nus, with long 2-segmented palp, stout dis-
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tal spines on clasper border, and small cup- like projection on distal border of clasper finger (Fig. 2F).
Caudal region.—Similar in males (Fig. 2G) and females (Fig. 2H), with 13 to 17 stout caudal spines and with telsonal fila- ments arising from about level of fourth posterior spine.
Eggs.—Stout and cylindrical, with par- allel grooves separating rounded ridges along cylinder axis and with perpendicular grooves on end pieces (Fig. 3D-F; see also Martin 1989, fig. 5e—g).
Type locality. — Paraguay, Chaco Depart- amento, Parque Nacional Defensores del Chaco, Tribo Nuevo, “encontrado en re- giones bajas de laguna recién inundada,”’ shallow region with submerged grass.
Range.—Known only from the type lo- cality.
Etymology. —The name refers to the marked similarity that eggs of this species bear to eggs of the North American species E. texana (see Belk 1989).
Remarks. —The eggs of Eulimnadia ovi- similis are virtually identical to those of E. texana, a predominantly North American species that has been reported, probably mistakenly, as far south as Sao Paulo, Brazil (Lutz 1929, Daday 1926). This is disturbing in light of our recent findings that egg mor- phology is often species-specific in the Lim- nadiidae and is in any case a more conser- vative taxonomic character than any previously employed feature. Indeed, we at first thought that we had encountered a southern hemisphere population of E. tex- ana rather than an undescribed form. How- ever, there are several salient differences be- tween the two species. In E. texana males and females have a similar rostral mor- phology. In contrast, sexual dimorphism is marked in E. ovisimilis. In females of E. ovisimilis the rostrum is rounded and slight- ly protruding, whereas in males the rostrum is attenuated and sharply produced (see Fig. 2D, E). Although there is known to be vari- ation in rostral morphology in E. texana,
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there is never sexual dimorphism as marked as is seen in E. ovisimilis (Sissom 1971, Belk 1989). Additionally, the eggs of E. ovisimilis appear less domed on the end pieces than are eggs of E. texana.
Discussion
Morphology of the external egg shell has been shown to be a useful and reliable char- acter in identifying species of the genus Eu- limnadia (Belk 1989). Some caution is ad- vised when consulting previous published accounts of egg morphology where scanning electron microscopy was not used. Specifi- cally, some of the illustrations or verbal ac- counts given by Daday (1926) were shown by Belk (1989) to be erroneous or lacking in sufficient detail to be of taxonomic value, perhaps a result of limitations of the mi- croscopy available at that time. Even when SEM is applied, however, there may be cases where egg morphology will fail to distin- guish between two otherwise recognizable species. This was pointed out by Mura (1986) for distinct species of anostracans that have identical egg morphologies, and the present paper demonstrates that this scenario also occurs in some species of clam shrimp (Spinicaudata only; the Laevicau- data do not have sculptured eggs; Martin & Belk 1988). This need for caution was noted by Belk (1989) in anticipation of this sort of problem. We emphasize again the need for using SEM on branchiopod egg mor- phology, but advise against using egg mor- phology to the exclusion of other characters.
Finally, as mentioned briefly by Belk (1989) and Martin (1989), the present status of the genera Limnadia Brongniart and Eu- limnadia Packard is unclear. Most mor- phological characters previously used for separating the two genera are variable, and there is some overlap, leading several work- ers to suggest that the two genera should be synonymized (Webb & Bell 1979). How- ever, there are at least two characters that serve to separate the two genera for species
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in the Americas. First, the well developed spine on the posteroventral border of the caudal somite is always easily discerned in Eulimnadia, whereas in the only species of Limnadia known from the Americas (L. lenticularis) this spine is absent (although a small lobe is present in the same location). Webb & Bell (1979:fig. 1) show a morpho- logical gradation from one state to the other based on drawings in existing literature, and suggest that this character is therefore un- reliable. Second, the telsonal filaments arise from between the paired spinose postero- caudal borders in all American Eulimnadia, whereas in Limnadia lenticularis the fila- ments arise from a location anterodorsal to the point where these borders become fused. We will address these two characters and the status of Eulimnadia vs. Limnadia ina future paper.
Acknowledgments
Much of this paper was written or planned during a brief visit by JWM to San Antonio, Texas, during which time the hospitality and assistance of Mary Shug Belk was greatly appreciated by both authors. Captain J. Trent Collier, U.S. Air Force, assisted in the collection of specimens used later for com- parative work and provided useful discus- sion. We thank Arthur C. Hulse and Terry Bonace for collecting and giving to Denton Belk the specimens used in these descrip- tions. This work was supported by the Na- tional Science Foundation, grant no. BSR- 8615018 to J. W. Martin and L. G. Abele.
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Literature Cited
Belk, D. 1989. Identification of species in the con- chostracan genus Eulimnadia by egg shell mor- phology. — Journal of Crustacean Biology 9:115— 125%
Daday, E. 1926. Monographie systématique des Phyllopodes Conchostraces. Troisiéme Partie (suite). —Annales des Sciences Naturelles, Zool- ogie, 10e série, 1926 9:1-81 (505-580).
Lutz, A. 1929. Dous phyllopodos, observados no Rio Grande do Norte.—Instituto Oswaldo Cruz, Supplemento das Memorias 5:3-9, 3 plates.
Martin, J. W. 1989. Eulimnadia belki, a new clam shrimp from Cozumel, Mexico (Conchostraca, Limnadiidae), with a review of Central and South American species of the genus Eulimnadia.— Journal of Crustacean Biology 9:104—114.
—.,&D. Belk. 1988. Areview of the clam shrimp family Lynceidae Stebbing, 1902 (Branchiopo- da: Conchostraca), in the Americas. — Journal of Crustacean Biology 8:451-482.
Mura, G. 1986. SEM morphological survey on the egg shell in the Italian anostracans (Crustacea, Branchiopoda).—Hydrobiologia 134:273-286.
Sissom, S. L. 1971. Morphological variation in Eu- limnadia texana, Texas’ most common eulim- nadian Conchostraca.—Texas Journal of Sci- ence 23:295-297.
Webb, J. A., & G. D. Bell. 1979. A new species of Limnadia (Crustacea: Conchostraca) from the granite belt in southern Queensland and north- ern New South Wales. — Proceedings of the Lin- naean Society of New South Wales 103:237- 245.
(JWM) Life Sciences Division, Natural History Museum of Los Angeles County, 900 Exposition Boulevard, Los Angeles, California 90007; (DB) Biology Depart- ment, Our Lady of the Lake University of San Antonio, San Antonio, Texas 78285.
PROC. BIOL. SOC. WASH. 102(4), 1989, pp. 901-911
DANTYA FEROX, A NEW SPECIES OF MYODOCOPID OSTRACODE FROM NIUE, CENTRAL SOUTH PACIFIC (CRUSTACEA: OSTRACODA: SARSIELLIDAE)
Louis S. Kornicker and Thomas M. Iliffe
Abstract.—Dantya ferox, a new species of myodocopid ostracode in the subfamily Dantyinae from a marine cave in the island of Niue, central South Pacific, is described and illustrated. The genus had not been reported previously from the Pacific. A key is presented to the species of Dantya.
The genus Dantya, proposed in 1978, is now known from five species, but speci- mens are sparse, with only one species known from more than two specimens (Dantya benthedi—50 specimens, D. mag- nifica—2 specimens, and D. fossula, D. piercei, and D. ferox—1 specimen for each species). The adult male is not known for any of the species, but this is not unusual in the Sarsiellidae, because the ratio of fe- males to males is high; as a consequence systematic discrimination in the family is based mostly on female characters.
The island of Niue is a raised limestone atoll encompassing 259 square km located 386 km east of Vava’u, Tonga, in the central South Pacific Ocean (Fig. 1). A 20 m sea cliff at the inner edge of a narrow reef plat- form surrounds the island. Inland, a second terrace rises to a central plateau about 60 m above sea level. Faulting during uplift has produced many deep chasms which run par- allel to the coastline. Well developed karst relief is present around the margins of the island.
An adult female of a new species, Dantya ferox, described herein, was collected inside a cave with direct connection to the sea along the west side of the island. The species is not considered to be a troglobite because of the open connection of the cave to the sea; this conclusion is supported by the species having normal eyes. This is the first report
of the genus in the Pacific Ocean; previ- ously, the genus was known only from the Caribbean Sea (one species) and Indian Ocean (three species).
Sarsiellidae Brady & Norman, 1896
Composition.—The Sarsiellidae include two subfamilies: Sarsiellinae Brady & Nor- man, 1986, and Dantyinae Kornicker & Co- hen, 1978.
Dantyinae Kornicker & Cohen, 1978
Composition. —The Dantyinae include two genera: Dantya Kornicker & Cohen, 1978, and Nealella Kornicker & Caraion, 1980.
Dantya Kornicker & Cohen, 1978
Type species.—Dantya magnifica Kor- nicker & Cohen, 1978.
Composition. —The new species de- scribed herein increases the number of known species of the genus to five: D. mag- nifica Kornicker & Cohen, 1978, D. ben- thedi Kornicker, 1983, D. piercei Kornicker, 1983, D. fossula Kornicker, 1983, and D. ferox, new species.
Distribution. — D. magnifica is known only from a coral reef fringing Carrie Bow Cay, Belize, Caribbean Sea, where it was col- lected at a depth of 20 m. D. fossula and D.
902 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
O kilometers 15
Fig. 1. Map showing location of the island of Niue, central South Pacific Ocean. Cave from which Dantya ferox was collected is near Alofi.
benthedi were collected in the Mozambique Channel, Indian Ocean, at depths of 24 m and 250-550 m, respectively. D. piercei was also collected in the Indian Ocean, on the continental shelf east of the Somali Repub- lic at a depth of 60-70 m. The new species, D. ferox, was collected at a depth of 2 m in a sea cave on the island of Niue, central South Pacific Ocean.
Key to the Species of Dantya (females)
1. Ventral margin of rostrum forming right angle with anterior margin of Valve ventral (0 rosimum? -.. 3. 2
— Ventral margin of rostrum forming acute angle with anterior margin of valve ventral to rostrum ......... )
2. Surface of valves with numerous minute knob-like processes; dorsal margin of second joint of first an- tenna with one bristle ... D. magnifica
— Surface of valves without knob-like processes; dorsal margin of second joint of first antenna without bristle
3 pe) sive eer: D. ferox, new species
3. Longest ventral claw of first endo-
podial joint of mandible with three
stout teeth, without slender teeth and SRICSpyg econ ea ee D. benthedi
— Longest ventral claw of first endo-
podial joint of mandible with slen-
der teeth and spines and without
three stout teeth 4. Second endopodial joint of mandi-
ble with five claws ......... D. piercei
— Second endopodial joint of mandi-
ble with two claws ......... D. fossula
Dantya ferox, new species Figs. 2—6
Etymology. — From the Latin ferox (wild, spirited, fierce) in reference to the name Savage Island by which the island of Niue is also known.
Holotype. —USNM 193645, adult female on slide and in alcohol, unique specimen.
Type locality. —PWD (Public Works De- partment) Cave (Stn. 88-012), Alofi, leg. T. M. Iliffe, 23 Feb 1988; unique specimen col- lected in a plankton net from gravel bottom of cave in 2 m depth and 5 m inside the cave from the sea.
Description of cave. —PWD Cave is a sea cave located on the west coast of Niue and behind the Public Works Department depot at Alofi. It is reached by a tourist footpath south of the depot which leads to the coast. The cave is at the back of a small sheltered bay. It consists of a sea water filled fissure approximately 10 m in length with a small dry section at the inner end. Water depth in the cave is 2 m with a gravel bottom. The water temperature was 29°C on 23 Feb 1988. The walls of the cave were relatively barren despite the direct connection with the sea, and had only a few small sponges and other encrusting organisms. A sea snake was ob- served in the rear of the cave. Amphipods, tanaidaceans and isopods were also collect- ed.
Description of adult female (Figs. 2—6).— Carapace elongate with prominent rostrum with pointed tip, and elongate caudal pro- cess with truncate tip (Figs. 2, 3a).
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Fig. 2. Dantya ferox, holotype, adult female, USNM 193645: Lateral view of complete specimen from right side, length 0.99 mm. All over tone for shape, contour and lighting done with airbrush.
Ornamentation: Surface with numerous oval fossae with crenulate or papillate edges and papillate bottoms, and two low hori- zontal ribs formed of platelets having cren- ulate posterior edges (Figs. 2, 3b); platelets closer together at anterior and posterior ends of ribs than at midlength (Fig. 2); surface of platelets with minute pores or papillae (dif- ficult to resolve) (Fig. 3b). Upper rib with anterior end at tip of rostrum and posterior end at posterodorsal corner of valve; lower rib passing over lower half of central ad- ductor muscle attachments, with anterior end at anteroventral corner of valve and posterior end at vertical ridge anterior to caudal process; upper and lower ribs weakly connected by low vertical rib at about one- fourth length of valve from anterior end (Fig. 2); vertical rib extends dorsally and branch- es near dorsal edge of valve.
Carapace bristles: Outer surface with widely scattered medium and long bare bristles, some with broad base, most with bases in shallow round fossae (Fig. 2). Inner side of rostrum with two bristles forming row near ventral margin (Fig. 3c); inner side of anteroventral margin with about 12 bris- tles forming row close to valve edge and five bristles forming distal row closer to edge (Fig. 3d); inner side of ventral margin with eight bristles forming row.
Infold: Anterior edge of infold at poste- rior end of rostrum with three spinous bris- tles forming verticle row, and one shorter bare bristle near inner corner of incisur (Fig. 3c). Broad anteroventral infold with three parallel ridges, one small bare bristle at mid- width ventral to rostrum, and three similar bristles near outer edge at anteroventral cur- vature of valve (Fig. 3d). Infold of caudal
904 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig. 3. Dantya ferox, holotype, adult female, USNM 193645: a, Dorsal view of complete specimen, anterior to left, length 0.99 mm (specimen slightly oblique and with valves open slightly); b, Detail of surface of right valve at midlength of lower rib; c, Inside view of anterior of right valve; d, Inside view of anteroventral margin of right valve; e, Inside view of caudal process of right valve; f, Outside view of central adductor muscle attachments of right valve, anterior to right. Abbreviations: i.m., inner margin of infold; s, selvage. Scale bar represents 0.1 mm for d, and f, and 0.05 mm for b, ¢, e.
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process with four bristles along inner mar- gin, and “pocket” with six or seven flat frond-like bristles forming row along ante- rior edge of pocket, and two small indistinct spines forming row near posterior edge of caudal process (Fig. 3e).
Selvage (Fig. 3d): Lamellar prolongation of selvage with anterior end near ventral spinous bristle of rostral infold and poste- rior end at ventral end of caudal process; prolongation between inner end of incisur and anteroventral corner of valve with short marginal spines; short section of prolonga- tion posterior to anteroventral corner with long streamers along margin; posterior to that section prolongation broader, either bare or with minute marginal spines; pro- longation narrow and bare along ventral margin of caudal process and absent along posterior edge (not shown); dorsal edge of rostrum and caudal process with narrow prolongation (not shown).
Central adductor muscle attachments (Fig. 3f): Consisting of about 17 discrete oval at- tachments.
Carapace size: Holotype: length 0.99 mm, height 0.58 mm.
First antenna (Fig. 4a): First joint bare. Second joint with minute medial spines forming row in distal dorsal corner. Third and fourth joints and also fifth and sixth joints fused but each joint identified by dis- tribution of bristles. Third joint with two bristles: dorsal bristle with long proximal hairs and few minute spines at tip; ventral bristle longer and with short marginal spines. Fourth joint with three bristles: single dorsal bristle with few indistinct short spines; shorter of two ventral bristles medial and with indistinct short spines; longer of ven- tral bristles lateral, bare. Ventral bristle of long fifth joint with five small filaments and minute process at tip. Sixth joint with spi- nous medial bristle longer than fused fifth and sixth joints. Seventh joint: a-bristle longer and stouter than bristle of sixth joint, with few indistinct short spines; b-bristle
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slightly longer than a-bristle, with short dis- tal filament and minute process at tip; c-bristle about same length as bristle of fifth joint, with three small marginal filaments and minute process at tip. Eighth joint: d- and e-bristles same length as c-bristle, bare with blunt tips; f-bristle shorter than c-bris- tle, with two short proximal filaments, one minute subterminal filament or spine, and minute process at tip; g-bristle same length as c-bristle, with two short proximal fila- ments, one minute filament or spine near midlength, and minute process at tip. All bristles ringed (rings not shown).
Second antenna: Protopodite bare (Fig. 4b). Endopodite two-jointed (Fig. 4b): first joint with two small ringed proximal ante- rior bristles; second joint small, with long ringed bristle with short marginal spines. Exopodite (left limb): first joint with small recurved medial bristle near midwidth of distal margin (Fig. 4c); bristles of joints 2— 8 long, with six proximal ventral spines (dis- tal spine longest) followed by natatory hairs (Fig. 4e); bristles of joints 4—7 with dorsal hairs proximal to spines; ninth joint small, with two bristles (ventral bristle shorter and slenderer than bristle of eighth joint, with three small proximal ventral spines fol- lowed by one longer dorsal spine, then na- tatory hairs; dorsal bristle of ninth joint short with few small hair-like marginal spines). Joint 2 with spines forming two distal rows (Fig. 4c); joints 3-6 with spines forming one distal row (Fig. 4c, d); spines mostly on me- dial side but rows curving around dorsal edge of joint resulting in few spines being on lateral side near dorsal margin. Joints 4— 8 with basal spines increasing in size on distal joints (spine on eighth joint about twice length of small ninth joint (Fig. 4d). Exopodite of mght limb aberrant, with only seven joints: small medial terminal bristle of first joint straight, not bent as on left limb; bristles of joints 2—5 similar to those of left limb; bristle of sixth joint with only three ventral spines and distal part invaginated
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VOLUME 102, NUMBER 4
into base (Fig. 4f); terminal seventh joint larger than terminal ninth joint of left limb, with two bristles (longest bristle ventral, with three ventral spines and without natatory hairs; dorsal bristle minute, about half length of seventh joint); joints 2-6 with long me- dial spines forming one to three rows ar- ranged differently than on left limb; joints 2-6 with basal spines. (Exopodial bristles not shown in Fig. 4c, d.)
Mandible (Fig. 4g): Coxale endite with stout terminal spine with two or three small- er marginal spinules, a subterminal spine with two smaller marginal spines, and long slender spines forming two proximal rows; ventral edge of coxale with slender spines forming two or three rows. Basale: medial side near ventral margin with three small ringed bristles (distal stouter and with mar- ginal spines); medial side near proximal margin with two rows of spines; lateral side near ventral margin with row of three mi- nute bare ringed bristles; dorsal margin with short ringed bristle distal to midlength and two terminal ringed bristles (longest reach- ing midlength of first endopodial joint) (rings on bristles not shown). Exopodite cylindri- cal, about one-third length of dorsal margin of first endopodial joint; tip with short di- aphanous triangular flap and ringed bristle (rings not shown). First endopodial joint: medial side with rows of distal spines; dor- sal margin with row of terminal spines (spines stouter than those of medial side); ventral margin with small medial ringed ter- minal bristle (rings not shown) and two ter- minal claws (proximal medial claw with
—
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slender marginal spines, other stouter and with four or five stout ventral spines at mid- length (spines increasing in length and stoutness distally along claw). Second en- dopodial joint: dorsal margin with three bristles (one bare, two with spines); ventral margin with stout claw (with ventral and dorsal spines) distal to midlength, and stouter terminal claw with few proximal ventral spines; medial surface with slender spines forming rows; lateral surface with small indistinct terminal bristle near mid- width. Third endopodial joint with one short spinous dorsal claw, one small unringed spine-like bristle adjacent to dorsal claw (in- distinct and observed only on right limb), two small ringed lateral bristles near ventral margin, two stout bare unequal terminal claws, and one small medial spine just ven- tral to shortest stout terminal claw (spine could be on claw).
Maxilla (Fig. 4h, 1): Coxale with stout dor- sal bristle. Endites I, II, and III each with five bristles (Fig. 41). (Note: in Fig. 41 an- terior bristle of endite II is behind endite III making it appear to be on that endite.) Ba- sale with spinous dorsal bristle and distal medial bristle (not shown in Fig. 4h). Ex- opodite with three terminal ringed spinous bristles (one longer than others). First en- dopodial joint: alpha-bristle with distal rings, long proximal hairs and short distal spines; beta-bristle stouter than alpha-bris- tle, ringed only near tip, with short distal hairs; anterior margin and medial surface near anterior margin with long spines (not all shown). Second endopodial joint with
Fig. 4. Dantya ferox, holotype, adult female, USNM 193645: a, Medial view of right first antenna. b-e, Medial views of left second antenna: b, Protopodite, endopodite, and exopodial joints 1 and 2; c, Joints 1-3 of exopodite; d, Joints 5—9 of exopodite; e, Bristle of second exopodial joint; f, Tip of invaginated bristle of sixth joint of exopodite of aberrant right second antenna; g, Medial view of right mandible; h, Lateral view of night maxilla (endites not shown); i, Medial view of endites of right maxilla. Abbreviations: a-g, letters assigned to bristles; s, sensory bristle of fifth joint of first antenna; Arabic numbers, numbers assigned to individual joints; Roman numerals, numbers assigned to endites. Scale bar represents 0.05 mm for a, b, e, g—i, and 0.02 mm for
cod, t.
908 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig. 5. Dantya ferox, holotype, adult female, USNM 193645: a, Anterior view of right fifth limb; b, Posterior bristles of right fifth limb as seen through limb; c, Posterior view of right fifth limb as seen attached to body (only 1 epipodial bristle shown); d, Posterior bristles of exopodial joints 2—5 of left fifth limb as seen through
VOLUME 102, NUMBER 4
two spinous lateral a-bristles (one missing on illustrated right limb), one smaller and more slender ringed spinous medial c-bris- tle, and five terminal bristles: anterior bris- tle ringed distally and with slender teeth along margins (teeth longer and stouter along unringed part); other bristles stouter, un- ringed, with teeth along margins proximal to midlength, and with narrow transparent velum along each edge (Fig. 4h). (Note: a dorsal bristle was observed on a coxale when the maxilla was attached to the body, but was absent on each mounted limb; I assume that the bristle was broken off during dis- section; a visible empty socket on the right limb supports the assumption.)
Fifth limb (Fig. 5a—d): Epipodite with about 40 bristles (Fig. 5c). Endite I with two bristles; endite II with three bristles; endite III with six bristles (Fig. 5a). Exopodite: first joint: anterior side with two short pectinate bristles at midwidth and one closer to inner edge (Fig. 5a). Second joint: large square tooth with proximal round tooth on inner edge (Fig. 5a); posterior side with three bris- tles forming row (middle bristle stout pec- tinate) (Fig. 5b, d) (teeth not shown on mid- dle bristle in Fig. 5d). Third joint with two short bristles on outer lobe and one long bristle on inner lobe (Fig. 5b, d) (bristle of inner lobe could be on first or second joints; bristle observed only on left limb). Fourth and fifth joints fused, with total of five bris- tles (Fig. 5b, d).
Sixth limb (Fig. Se-g): Limb partly frag- mented during dissection, with four endites. Endite I with three short bristles; endite II with two bristles (one missing in Fig. 5e); endites III and IV each with four bristles. End joint with eight or nine bristles (pos- terior two bristles hirsute, others mostly with short stout spines, but some with proximal hairs and slender distal spines). A single bare
909
bristle on posterior margin interpreted herein to be epipodial bristle. Limb hirsute medially.
Seventh limb (Fig. 6a—c): Each limb with four proximal bristles (two on each side), each with two to four bells, and six terminal bristles (three on each side), each with three to seven bells; all bristles without marginal spines. Terminus with comb of about five teeth opposite two small pegs (one straight, one slightly longer and curved) (Fig. 6c).
Furca: Each lamella with six claws (Fig. 6d, e); 1, 2, and 4 primary claws; claws 3, 5 and 6 secondary claws; claw 4 stouter than claw 3; claw 1 nonarticulated on both la- mellae; claw 2 of left lamella articulated (Fig. 6d), of right lamella nonarticulated (prob- ably aberrant because claw 2 of all known species of genus are articulated) (Fig. 6e); remaining claws articulated; claw 1 with teeth forming two rows along proximal two- thirds, some teeth slightly longer than oth- ers; claw 2 with few proximal teeth; claw 3 with slender teeth along most of posterior margin; no teeth observed on claw 4; few indistinct teeth on claws 5 and 6; anterior edge and medial surface of right lamella with
_ long hairs; anterior edge of left lamella with
few proximal spines; claw 1 or right lamella slightly anterior to claw 1 or left lamella (Fig. 6d).
Bellonci organ (Fig. 6f, g): Elongate, bare, with five proximal segments, broadening distally and with unevenly rounded tip.
Eyes: Lateral eyes with five amber-col- ored ommatidia (Fig. 6f). Medial eye larger than lateral eye, bare, with scattered brown pigment (Fig. 6f, g).
Upper lip (Fig. 61): Rounded with minute spines.
Genitalia (Fig. 6h, j): Oval sclerotized ring on each side of body anterior to furca.
_
limb; e, Medial view of right sixth limb; f, Lateral view of left sixth limb without endites I and II; g, Medial view of endites II and III of left sixth limb. Abbreviations: Arabic numbers, numbers assigned to joints; Roman numerals, numbers assigned to endites. Scale bar represents 0.05 mm for c, and 0.02 mm for a, b, d-g.
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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
— Fa a) a omer gs
Fig. 6. Dantya ferox, holotype, adult female, USNM 193645: a, Seventh limb; b, Detail from a; c, Tip of seventh limb viewed from peg side (this limb not limb shown in a); d, Medial view of claw 1 of right lamella of furca and lateral view of left lamella; e, Posterior of body with right lamella of furca; f, Anterior of body showing medial eye, Bellonci organ, left lateral eye, and joints 1 and 2 of right first antenna; g, Medial eye and Bellonci organ; h, Posterior of body showing girdle, Y-sclerite, left genital organ, and proximal part of left lamella of furca; i, Upper lip, anterior to right; j, Ventral view of both genitalia; k, Brush organ of right side. Abbreviations: gen, genital organ; m.e., medial eye. Scale bar represents 0.1 mm for e, 0.05 mm for a, c, d, f, h-k, and 0.02
mm for b, g.
VOLUME 102, NUMBER 4
Brush-like organ (Fig. 6k): Four or five minute bristles anterior to Y-sclerite.
Posterior of body (Fig. 6e): Evenly round- ed, bare.
Y-sclerite (Fig. 6h): Branching distally.
Comparisons. —D. ferox differs from pre- viously described species of Dantya in lack- ing a dorsal bristle on the second joint of the first antenna. In lateral view the cara- pace of D. ferox resembles that of D. mag- nifica, but lacks the knob-like processes present on the surface of that species. Also, the exopodite of the mandible of D. ferox is about one-third the length of the dorsal margin of the first endopodial joint, and is minute on D. magnifica.
Acknowledgments
Collections of specimens from caves in Niue was part of a year-long expedition studying the biology of marine caves in the South Pacific. This research was supported by grants from the National Science Foun- dation (BSR-8700079) and the National Geographic Society (3412-86) to Iliffe. We thank Drs. John Maciolek and L. Eldredge for providing information on the island of Niue and its caves, T. Coe and E. Meili of Niue Adventures for assisting with diving logistics in Niue, and Yolanda Iliffe for helping with cave collections. We also thank Molly Ryan for rendering the illustration of the holotype in Fig. 2, Jack Schroeder for
911
inking penciled camera lucida drawings of appendages, and Elizabeth Harrison-Nel- son for general assistance in preparing the manuscript for publication. This paper is Contribution No. 1232 of the Bermuda Biological Station for Research. Our thanks to Dr. Anne C. Cohen for reviewing the manuscript and to Dr. Thomas E. Bowman for suggesting the specific name.
Literature Cited
Brady, G. S.,& A. M. Norman. 1896. A monograph of the marine and freshwater Ostracoda of the North Atlantic and of northwestern Europe. — Scientific Transactions of the Royal Dublin So- ciety, series 2 5:621-784.
Kornicker, Louis S. 1983. New species of Dantya from the Indian Ocean (Ostracoda: Sarsiellidae: Dantyinae).—Smithsonian Contributions to Zoology 383:1-18.
—., & Francisca Elena Caraion. 1980. Nealella, a new genus of myodocopid Ostracoda (Sar- siellidae: Dantyinae).—Smithsonian Contribu- tions to Zoology 309:1-27.
—, & Anne C. Cohen. 1978. Dantyinae, a new subfamily of Ostracoda (Myodocopina: Sarsiel- lidae). — Proceedings of the Biological Society of Washington 91(2):490—508.
(LSK) Department of Invertebrate Zo- ology, National Museum of Natural His- tory, Smithsonian Institution, Washington, D.C. 20560; (TMI) Bermuda Biological Sta- tion for Research, Ferry Reach GE 01, Ber- muda.
PROC. BIOL. SOC. WASH. 102(4), 1989, pp. 912-915
LAMELLIFORM STRUCTURES ON THE PROBOSCIS OF PENICULUS AND METAPENICULUS (COPEPODA: PENNELLIDAE)
Raul Castro Romero and Hernan Baeza K.
Abstract.—Two pairs of laminae, at the proboscis base ventral surface were discovered in adult and chalimus IV female stage of Peniculus specimens. One pair of these structures is present at the proboscis base of the premetamor-
phosing female of Metapeniculus.
The laminae of Peniculus are narrow, those of Metapeniculus wide. These laminae are described and illustrated; their importance for the copepod and
its taxonomy are discussed.
The taxonomy of the Pennellidae Bur- meister, 1835 (Copepoda: Siphonostoma- toida), which includes parasites on teleosts and some on Balaenoptera (e.g., Pennella Oken, 1816) has some problems at both the generic and specific levels. The problems are due to the lack of good discriminant char- acters, and to the fact that we do not possess accurate and detailed description of the ap- pendages for the majority of pennellid species. The characters used at present (e.g., buccal appendages, leg armament) are more or less uniform throughout the family, and have little, if any discriminant value. The morphology of the anterior part of the body, one of the most commonly used taxonomic characters, shows great intraspecific vari- ability according to the specific attachment site, as has been shown by Hogans (1986) for Pennella instructa Wilson, 1917 and by Bellwood (1981) for Cardiodectes spiralis Bellwood, 1981.
The taxonomy of Peniculus von Nord- mann, 1832 is not clear, and is made more difficult by the lack of external characters that facilitate differentiation of the species and that indicate their relationships with other genera of the family, especially with Metapeniculus Castro & Baeza, 1985, which is very close morphologically. Kabata & Wilkes (1977) suggest that P. fissipes Wil-
son, 1917 is probably a synonym of P. fistula von Nordmann, 1832, this being only one example of the taxonomic problems within this genus.
We searched for new features that could be useful in clarifying the interspecific and intergeneric differences among the pennel- lids, as well as in establishing intergeneric relationships. We studied some specimens of Peniculus von Nordmann, 1832 and Metapeniculus antofagastensis Castro & Baeza, 1985, and discovered lamelliform structures on the ventral surface of the buc- cal cone of Peniculus and M. antofagasten- sis, which are illustrated, described, and their importance for the copepods and their tax- onomy discussed.
Methods. —Peniculus specimens were taken from different host species; adult fe- males from Hemilutjanus macrophthalmus (Tschudi, 1845), Mugiloides chilensis (Mol.), and Sciaena fasciata (De Buen); chalimus IV from Eleginops maclovinus (Val.). The premetamorphosing female of M. antofa- gastensis was collected from the type host Anisotremus scapularis (Tschudi).
The material was fixed and preserved in formalin (5%) and/or alcohol (70%). Some specimens, fixed as described above, were dehydrated by critical point drying, coated with gold, and examined under SEM Auto-
VOLUME 102, NUMBER 4 913
Figs. 1-5. Peniculus sp.: 1, Adult 2 buccal area, lateral view, x 400; 2, Adult 2 cephalothorax, ventrofrontal view, x 200; 3, Adult 2 detail of laminae, x 800; 4, Chalimus 4 2, ventral view, showing position of laminae on proboscis ventral surface, x98; 5, Metapeniculus antofagastensis, premetamorphosing 2, ventral view, x 400. Abbreviations: bt— buccal tube; fm—first maxilla; is—intrabuccal stylet; sm—second maxilla. Laminae indicated by arrows and asterisks.
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scan at 20 Kv acceleration. To prevent pos- sible damage to the copepod surface no spe- cial cleaning agents were used.
Results. —Peniculus sp. (Figs. 1-4): The female specimens examined, regardless of their stage of development (adult and chal- imus IV), and the identity of their hosts, bear on the ventral surface, near the base of the buccal cone, two pairs of smooth, nar- row laminae with rounded margins. The two pairs are separated from each other by a short gap, and the two members of each pair are slightly separated from each other.
Metapeniculus antofagastensis (Fig. 5): Free living, not metamorphosed, females examined bear on the ventral surface, near the base of the buccal cone, a single pair of laminae, smooth, wide, and with entire margins. The two members of the pair are separated from each other near the mid- ventral line.
Discussion. —The existence of armament on the ventral surface of the pennellid buc- cal cone is not widely known. Kabata (1963) reported the presence of “‘serrated lamellae”’ on the buccal cone of Lernaeenicus sprattae, and later Kabata (1979) modified this de- scription to “denticles’’. These findings were corroborated by Schram (1979). Similar denticles were found by Kabata (1965), in Lernaeocera centropristi. In the same paper, Kabata mentioned the presence of trans- verse “‘wrinkles” on the buccal cone of Ler- naeocera branchialis (possibly long lami- nae). Thomsen (1949) found small ““denticles”’ (“‘finisimos dientecitos’’) in 77i- fur tortuosus, and Castro & Baeza (1985) confirmed his finding of ““small squamiform sclerites’’. We add to those records two pairs of long laminae in Peniculus and one such pair in Metapeniculus. |
In view of these findings it is reasonable to assume that structures of this kind may be present in other genera of Pennellidae. The differences among them, in number, shape etc. might be adaptations to their re- spective microhabitats and to the function they have evolved to fulfil. The two pairs
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
of laminae in Peniculus might assist in food intake by collecting solid particles and liq- uid and facilitating their ingestion. They might also be of some assistance in the pro- cess of penetration of the host tissues by the buccal complex.
The same is true of Metapeniculus, a ge- nus with microhabitat similar to that of Peniculus.
The reduction in number of laminae, and the increase in their size, could be indicative of a functional improvement of those struc- tures in Metapeniculus, according to their similar microhabitat shared with Peniculus.
The lamelliform structures show a clear difference between Metapeniculus (with one pair of laminae) and Peniculus (with two pairs of laminae). This adds to the differ- ences between these genera in their thora- copod number (four pairs in Peniculus and three in Metapeniculus).
Without any doubt the discovery of la- mellae in these two genera will be of great help in separating specimens of Peniculus and Metapeniculus which are very close in external gross morphology, and will help in determining generic relationships within the family. The presence or absence of this structure and their possible variability must be studied for all the species assigned to Peniculus, which can give us a clue for species differentiation, and the validity of some Peniculus species whose taxonomic status is not clear.
Literature Cited
Bellwood, R. 1981. Two new species of Cardiodectes Wilson (Copepoda: Siphonostomatoida).—Sys- tematic Parasitology 2:149-156.
Castro, R., & H. Baeza. 1985. Metapeniculus anto- fagastensis gen. et sp. nov. (Copepoda, Pennel- lidae) parasitic on two inshore fishes of Anto- fagasta, Chile, South Pacific. —Crustaceana 49(1): 22-29.
Hogans, W. 1986. Redescription of Pennella instruc- ta Wilson, 1917 (Copepoda: Pennellidae) from the swordfish (Xiphias gladius L.).—Canadian Journal of Zoology 64:727-730.
Kabata, Z. 1963. The free swimming stage of Ler-
VOLUME 102, NUMBER 4 915
naeenicus (Copepoda parasitica). — Crustaceana got of the sprat, Lernaeenicus sprattae (Sowerby)
5:181-187 (Copepoda: Lernaeoceridae).—Sarsia 64:279- 1965. Systematic position of the copepod 316.
Lernaeocera centropristi.—Proceedings of the Thomsen,R. 1949. Copépodos parasitos de los peces
Zoological Society of London 144:351-360. marinos del Uruguay.— Communicaciones
. 1979. Parasitic Copepoda of British fishes. — Zoologicas del Museo de Historia Natural de
The Ray Society, No. 152:1-468. London. Montevideo 3(54):1-41.
—., & S.N. Wilkes. 1977. Peniculus asinus (Co- pepoda: Pennellidae), a new species of copepod
parasitic on fishes of the genus Sebastes along Universidad de Antofagasta, Instituto de the west coast of North America.—Canadian Investigaciones Oceanologicas, Casilla 170, Journal of Zoology 55:1988-1991. Antofagasta, Chile.
Schram, T. A. 1979. The life history of the eye-mag-
PROC. BIOL. SOC. WASH. 102(4), 1989, pp. 916-923
ACONTIOPHORUS EXCAVATUS, A NEW SPECIES (COPEPODA: SIPHONOSTOMATOIDA) ASSOCIATED WITH THE SOFT CORAL DENDRONEPHTHYA (ALCYONACEA) IN THE INDO-PACIFIC
Arthur G. Humes
Abstract. —A new species of siphonostomatoid asterocherid copepod, Acon- tiophorus excavatus, is described from Madagascar, the Philippines, and the Moluccas, where it is associated with the alcyonaceans Dendronephthya mu- cronata and D. koellikeri. The new copepod, though closely related to Acon- tiophorus bracatus from the Mediterranean, may be distinguished by the ex- cavated outer margin of the first exopod segment of leg 1 and by the presence of only two small dentiform projections between the terminal setae on the free
segment of leg 5.
Many poecilostomatoid and siphono- stomatoid copepods are associated with shallow-water cnidarians in the Indo-Pacif- ic. While the poecilostomatoid associates of Alcyonacea have received considerable at- tention in recent years (e.g., Humes 1975, 1980, 1982; Humes & Dojiri 1979a, 1979b, 1979c; Humes & Stock 1973), the siphono- stomatoid copepods associated with these hosts are poorly known. The purpose of this work is to describe a new widely distributed asterocherid copepod living as an associate of soft corals belonging to the genus Den- dronephthya in the Indo-Pacific.
Materials and Methods
The host alcyonaceans were isolated in plastic bags immediately after collection. Later they were soaked for 1-2 hours in sea water with 5% ethanol, rinsed thoroughly, and the sea water passed through a fine net (about 120 holes per 2.5 cm). The copepods were then recovered from the sediment re- tained in the net.
The copepods were studied using the wooden slide/lactic acid technique de- scribed by Humes & Gooding (1974). Mea- surements were made on specimens in lactic
acid. All figures were drawn with the aid of a camera lucida. The letter after the expla- nation of each figure refers to the scale at which it was drawn. The abbreviations used are: A, = first antenna, A, = second anten- na, and P, = leg 1.
Order Siphonostomatoida Thorell, 1859 Family Asterocheridae Giesbrecht, 1899 Genus Acontiophorus Brady, 1880 Acontiophorus excavatus, new species Figs. 1-23
Type material. —43 92, 70 66 from Den- dronephthya mucronata (Pitter), in 25 m, N of Ankazoberavina, near Nosy Be, NW Madagascar, 13°27.6’S, 47°58.2’E, 24 Aug 1967. Holotype female, allotype, and 103 paratypes (38 99, 65 44) deposited in the National Museum of Natural History, Smithsonian Institution, Washington, D.C.
Other specimens. — 18 22, 43 66 from Den- dronephthya koellikeri Kikenthal, in 25 m, southern shore of Goenoeng Api, Banda Is- lands, Moluccas, 04°32’05”’S, 129°52’'30’E, 26 Apr 1975; 3 99, 2 46 from same host, locality, and date; 5 99, 9 4¢, and 9 cope- podids from Dendronephthya koellikeri, in 10 m, Poelau Gomumu, S of Obi, Moluccas,
VOLUME 102, NUMBER 4
01°50’00”S, 127°30'45”E, 30 May 1975; 5 92 from Dendronephthya sp., in 17 m, southwestern shore of Goenoeng Api, Ban- da Islands, 04°31'45”S, 129°51'55”E, 30 Apr 1975; 1 2 from unidentified alcyonacean, probably Dendronephthya, in 30 m, Bohol Island, Philippines, 10°17.9’N, 124°10.9’E, 21 Aug 1975.
Female. —Body (Fig. 1) with broad pro- some. Length 1.10 mm (0.99-1.19 mm) and greatest width 0.56 mm (0.51-0.59 mm), based on 10 specimens. Greatest dorsoven- tral thickness 0.35 mm. Epimeral areas of segments bearing legs 1-3 pointed, those of segment bearing leg 3 especially so. Segment bearing leg 4 much smaller than preceding segment and rounded laterally, with only slight point. Ratio of length to width of pro- some 1.22:1. Ratio of length of prosome to that of urosome 2.0:1.
Segment bearing leg 5 (Fig. 2) 78 x 101 pm, with small scalelike spines along both lateral margins. Genital segment 172 wm iong, 200 um wide at small anterior rounded expansions, and 185 wm wide posteriorly. Genital areas situated dorsolaterally in front of middle of segment. Each area (Fig. 3) with two small setae, 12 wm and 4 um. Two post- genital segments from anterior to posterior 49 x 123 um and 161 X 126 um. Elongate anal segment, more than twice as long as preceding segment, with small scalelike spines along both lateral margins. Postero- ventral border of anal segment smooth.
Caudal ramus (Fig. 4) 47 x 57 wm, wider than long, ratio 1:1.22. Outer lateral seta, lightly feathered, placed dorsally and sub- terminally, 209 wm. Dorsal seta 55 um, smooth, with proximal third broader than distal two-thirds. Outermost terminal seta 265 wm, innermost terminal seta 308 um, and two median terminal setae 297 um (out- er) and 374 um (inner), all feathered. Inner median terminal seta swollen. Ramus with outer margin having several small scalelike spines and few distal setules, inner margin with several distal setules.
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Dorsal surface of body without visible sensilla.
Egg sac empty or incomplete in most specimens. Two egg sacs with single egg 195 <x 164 um (Fig. 5).
Rostrum (Fig. 6) weakly developed. First antenna (Fig. 7) slender, 475 um long, 17- segmented. Lengths of its segments (mea- sured along their posterior nonsetiferous margins): 26 (68 um along anterior margin), 25, BOR26 1 e895 29129-99: 30, 30,34, 36, 15, 15, and 20 um, respectively. For- pana: pa 2110, 2.5) tebe ere 22. 2. 1, 2 + 1 aesthete, 2, 3, and 5 + 1 aesthete. Aes- thete on segment 14 172 um long. Certain setae on segments 1-4 subspiniform and having lateral setules. First segment with small spinules along anterior edge.
Second antenna (Fig. 8) with short coxa and elongate basis (greatest length 99 um) with small spinules on inner margin. Exo- pod 1l-segmented, slender, length 47 um, with small inner smooth seta and long ter- minal barbed seta 71 um long, and having setules along outer side as shown. Endopod 2-segmented, first segment 60 um long and unarmed, second segment 40 um long, bear- ing one seta proximally, one seta near midregion, and two terminal setae very un- equal in length, longer of these 138 wm. Fine ornamentation as in Fig. 8.
Siphon (Fig. 9) long and slender, 590 um long, reaching nearly to posterior rim of in- tercoxal plate of leg 3.
Mandible (Fig. 10) with 1-segmented palp bearing one small smooth subterminal seta and very long feathered terminal seta. Mas- ticatory part of mandible smooth, elongate, styliform. First maxilla (Fig. 11) with two lobes, outer smaller lobe with three setae, larger inner lobe with five setae, two plu- mose, two elongate smooth, and one smaller smooth seta. Few setules on inner angle of outer lobe. Second maxilla (Fig. 12) with unarmed basal segment, bearing recurved claw armed with two setae near its midre- gion and having recurved tip (Fig. 13). Max-
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918
genital
, egg, ventral (E); 6, cephalosome, ventral (E); 7, first antenna,
>
Figs. 1-7. Acontiophorus excavatus, new species, female: 1, dorsal (scale A); 2, urosome, dorsal (B); 3
area, dorsal (C); 4, caudal ramus, dorsal (D); 5
dorsal (F).
VOLUME 102, NUMBER 4
illiped (Fig. 14) with two short proximal segments, distal of these with one minute inner seta. Third segment elongate and un- armed. Segments 4, 5, and 6 forming part of “‘claw’’, fourth segment having one seta, fifth segment two setae, and sixth segment one seta. Claw elongate, 112 um, recurved, its proximal third swollen.
Ventral region between maxillipeds and first pair of legs not protuberant and having widely diverging U-shaped sclerotization in front of intercoxal plate of leg 1 (Fig. 6).
Legs 1-4 (Figs. 15-18) biramous, with 3-segmented rami throughout. Formula for armature as follows (Roman numerals in- dicating spines, Arabic numerals represent- ing setae):
P, coxa 0-1 basis 1-I exp I-1; I-1; I1,2,3 enp 0-1; 0-2; 1,2,3
P, coxa 0-1 basis 1-0 exp I-1; I-1; II,1,4 enp 0-1; 0-2; 1,1 + 1,3
P; coxa Q-1 basis 1-0 exp I-1; I-1; ITI,1,3 enp 0-1; 0-2; 1,1,3
P, coxa Q-1 basis 1-0 exp I-1; I-1; III,1,3 enp 0-1; 0-2; 1,1,2
Basis of leg 1 with inner barbed spine 36 um, small spinules adjacent to its insertion. First segment of exopod of leg 1 with outer margin excavated (Fig. 15). Coxa of leg 2 with outer pectinate fringe (Fig. 16). En- dopod of leg 4 (Fig. 18) with inner margins of segments having slender spinules rather than hairlike setules as in legs 1-3.
Leg 5 (Fig. 19) with oval free segment 76 x 50 um, placed ventrally, armed from in- ner to outer with two smooth spines 30 um and 25 um, short smooth seta 26 wm, and two longer weakly feathered setae 60 um and 52 um. Adjacent “‘dorsal”’ seta, here insert- ed ventrally, smooth, 40 um. Pair of small dentiform processes between two terminal setae. Outer margin of segment with small scalelike spines.
Leg 6 represented by two small setae on genital area (Fig. 3).
Color of living specimens in transmitted light pinkish red, eye red.
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Male. —Body (Fig. 20) with prosome less broad than in female. Length 0.71 mm (0.69-0.74 mm) and greatest width 0.32 mm (0.30-0.33 mm), based on 10 specimens. Greatest dorsoventral thickness 0.21 mm. Epimera of segment bearing leg 1 rounded, those of segments bearing legs 2 and 3 some- what pointed but less prominent than in female. Segment bearing leg 4 relatively wider than in female and more pointed. Ra- tio of length to width of prosome 1.62:1. Ratio of length of prosome to that of uro- some 2.14:1.
Segment bearing leg 5 (Fig. 21) 60 x 120 um. Genital segment 65 x 122 um, with well-rounded lateral margins. Three post- genital segments from anterior to posterior 39 x 94,24 x 78, and 75 x 73 um.
Caudal ramus 30 X 37 wm, resembling that of female.
Body surface as in female.
Rostrum like that of female. First antenna (Fig. 22) geniculate, 400 wm long, 12-seg- mented. Lengths of its segments (measured along their posterior nonsetiferous mar- gins): 26 (55 um along anterior margin), 22, 49. 29: 2. 5; 8, 66; 29, 52, 42, and:39 pam, respectively. Formula: 2,.2;10, 2, 5, 1, 1, 6, 2, 1, 1 + 1 aesthete, and 6. Second an- tenna, siphon, mandible, first maxilla, sec- ond maxilla, maxilliped, and ventral area between maxillipeds and first pair of legs as in female.
Legs 1-4 as in female.
Leg 5 similar to that of female but free segment smaller, 42 <x 28 um.
Leg 6 (Fig. 23) represented by 2 unequal setae 10 wm and 31 um.
Spermatophore unknown.
Color of living specimens in transmitted light opaque light tan, eye red.
Etymology. —The specific name excava- tus, Latin meaning hollowed out, refers to the excavated outer margin of the first seg- ment of the exopod of leg 1.
Remarks.—Eight species are presently recognized in the genus Acontiophorus (not including the very insufficiently described Acontiophorus angulatus Thompson, 1888).
920 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Figs. 8-16. Acontiophorus excavatus, new species, female: 8, second antenna, inner (scale C); 9, siphon, ventral (F); 10, mandible, anteroventral (F); 11, first maxilla, posterodorsal (C); 12, second maxilla, anterodorsal (F); 13, second maxilla, posteroventral (F); 14, maxilliped, posterior (C); 15, leg 1 and intercoxal plate, anterior (F); 16, leg 2 and intercoxal plate, anterior (F).
921
a N
(/ = 4
, dorsal
me
VOLUME 102, NUMBER 4
Sipe
7:
A \
IN
wis a
a ae IT e
al plate, anterior (scale F);
Female: 17, leg 3 and intercox
al plate, anterior (F); 19, leg 5, ventral (C). Male: 20, body, dorsal (E); 21, uroso
(F); 22, first antenna, dorsal (C); 23, leg 6, ventral (D).
Figs. 17-23. Acontiophorus excavatus, new species.
18, leg 4 and intercox
922
Acontiophorus excavatus differs from seven congeners (A. antennatus Hansen, 1923, A. brevifurcatus Stock, 1966, A. ornatus (Brady & Robertson, 1876), A. maldivensis Sewell, 1949, A. scutatus (Brady & Robertson, 1873), A. tynani Eiselt, 1965, and A. zea- landicus Nicholls, 1944) in that the last two postgenital segments in these species are nearly equal in length.
The new species resembles Acontiophorus bracatus Stock & Kleeton, 1963, in having the anal segment much longer than the pre- ceding segment, and in the presence of small scalelike spines along the sides of this seg- ment. However, A. excavatus differs from A. bracatus (and from all congeners as far as can be determined from published de- scriptions) in having the outer margin of the first segment of the exopod of leg 1 distinctly excavated (see Fig. 15), and in having two small dentiform processes between the two terminal setae on the free segment of leg 5 (instead of four as in A. bracatus).
Both Acontiophorus excavatus and A. bra- catus are associated with alcyonaceans, the former with the nephtheids Dendronephthya mucronata and D. koellikeri in the Indo- Pacific and the latter with the alcyoniid Par- erythropodium coralloides (and also the gor- gonians Eunicella stricta and Leptogorgia sarmentosa) in the Mediterranean.
Acknowledgments
The collection of the copepods was made possible in 1967 in Madagascar by a grant (GB 5838) from the National Science Foun- dation and in 1975 in the Moluccas and the Philippines during the Alpha Helix East Asian Bioluminescence Expedition, which was supported by the National Science Foundation under grants OFS 74 01830 and OFS 74 02888 to the Scripps Institution of Oceanography and grant MBS 74 23242 to the University of California, Santa Barbara.
I am indebted to Dr. J. Verseveldt, now deceased, for the identification of the al- cyonacean hosts.
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
The study of the copepods in the labo- ratory was supported by a grant (BSR- 8514561) from the National Science Foun- dation.
Literature Cited
Brady, G. S., & D. Robertson. 1873. Contributions to the study of the Entomostraca. VIII. On ma- rine Copepoda taken in the west of Ireland.— Annals and Magazine of Natural History (4)12: 126-142.
——, & 1876. Report on dredging off the coast of Durham and north Yorkshire in 1874.— Report of the 45th Meeting of the British As- sociation for the Advancement of Science 1875: 185-197.
Eiselt, J. 1965. Revision und Neubeschreibungen weiterer siphonostomer Cyclopiden (Copepoda, Crust.) aus der Antarktis. —Sitzberichte der Os- terreichischer Akademie der Wissenschaften in Wien, mathematisch-naturwissenschaftliche Klasse, Abteilung I, 174:151-169.
Hansen, H. J. 1923. Crustacea Copepoda. II. Cope- poda parasita and hemiparasita. — Danish Ingolf Expedition 3(7):1—92.
Humes, A. G. 1975. Cyclopoid copepods (Licho-
molgidae) associated with alcyonaceans in New
Caledonia. —Smithsonian Contributions to Zo-
ology 191:1-27.
1980. Copepoda (Cyclopoida, Lichomolgi- dae) associated with the alcyonacean Nephthea in the Moluccas.— Hydrobiologia 68:49-71.
. 1982. Copepoda (Poecilostomatoida, Licho-
molgidae) associated with the genus Sarcophy-
ton in the Indo-Pacific. — Publications of the Seto
Marine Biological Laboratory 27:25-76.
—, & M. Dojiri. 1979a. Poecilostome copepods (Lichomolgidae) from the alcyonacean coral Cespitularia multipinnata in the Moluccas.— Proceedings of the Biological Society of Wash- ington 92:51-69.
——, & 1979b. Poecilostome copepods (Cyclopoida, Lichomolgidae) from the alcyo- nacean Lobophytum crassum in the Moluc- cas.— Bulletin of Marine Science 29:554—-571.
——, & 1979c. Poecilostome copepods (Lichomolgidae) associated with the alcyona- cean Litophyton in the Moluccas.—Transac- tions of the American Microscopical Society 98: 337-352.
—.,&R.U. Gooding. 1974. A method for study- ing the external anatomy of copepods.—Crus- taceana 6:238-240.
—., & J.H. Stock. 1973. A revision of the family Lichomolgidae Kossmann, 1877, cyclopoid co-
VOLUME 102, NUMBER 4
pepods mainly associated with marine inver- tebrates.— Smithsonian Contributions to Zool- ogy 127:i-v, 1-368.
Nicholls, A. G. 1944. Littoral copepods from South Australia (II) Calanoida, Cyclopoida, Notodel- phyoida, Monstrilloida and Caligoida.—Rec- ords of the South Australian Museum 8(1):1- 62.
Sewell, R. B. S. 1949. The littoral and semiparasitic Cyclopoida, the Monstrilloida and Notodel- phyoida.—John Murray Expedition, 1933-34, Scientific Reports 9:17-199.
Stock, J. H. 1966. Cyclopoida siphonostoma from Mauritius (Crustacea, Copepoda).—Beaufortia 13:145-194.
923
—,&G. Kleeton. 1963. Copépodes associés aux invertébrés des cétes du Roussillon 3.—Acon- tiophorus bracatus n. sp. un cyclopoide siphon- ostome associé aux octocoralliaires. — Vie et Mi- lieu 14:551-559.
Thompson, I.C. 1888. Copepoda of Madeira and the Canary Islands, with descriptions of new genera and species. — Journal of the Linnean Society of London (Zoology) 20:145-156.
Boston University Marine Program, Ma- rine Biological Laboratory, Woods Hole, Massachusetts 02543.
PROC. BIOL. SOC. WASH. 102(4), 1989, pp. 924-932
AN UNUSUAL SPECIES OF THE BALANUS AMPHITRITE DARWIN COMPLEX (CIRRIPEDIA, BALANIDAE) FROM THE ANCESTRAL COLORADO RIVER DELTA IN WESTERN ARIZONA AND SOUTHEASTERN CALIFORNIA
Victor A. Zullo and Anna V. Buising
Abstract. —Shells and opercular plates of Balanus canabus, new species, form shell hashes in the upper Cenozoic Bouse Formation of western Arizona and southeastern California. The thin, columnar shells and morphologically vari- able opercular plates indicate growth of large, crowded colonies in low salinity waters. The vesicular sheath and the overall morphology of the opercular and compartmental plates are characteristic of the genus Fistulobalanus Zullo, but the lack of multiple rows of parietal tubes precludes assignment to this genus. Balanus canabus 1s not clearly related to any extant eastern Pacific species, but does resemble the western Pacific species Fistulobalanus albicostatus (Pilsbry) and F. kondakovi (Tarasov & Zevina) and the Atlantic and Indian Ocean species
F. pallidus (Darwin).
Barnacle hashes in the Bouse Formation of western Arizona and southeastern Cali- fornia (Fig. 1) are formed by the shells of a single species of balanid barnacle. The hash- es consist of partially crushed, extremely thin-walled, columnar to tulipiform shells with intact opercular pyramids, together with disarticulated compartmental and opercular plates (Fig. 4e). The new species is remarkable in the thinness of its plates and, particularly, in the unusual growth modifications of its scuta and terga. The carinal margins of the terga and the upper occludent margins of the scuta are reflexed inward through secondary growth, creating an expanded compartment at the apex of the opercular pyramid.
The tubiferous shell wall with numerous transverse septa, the tubiferous basis, the narrow radii with finely denticulate sutural edges, and the well developed scutal ad- ductor ridge, serve to identify this species with the diverse and widespread Balanus amphitrite Darwin complex (see Henry & McLaughlin 1975). Although the overall
morphology of the opercular plates and the vesicular sheath of the new species are char- acteristic of the genus Fistulobalanus Zullo, the lack of multiple rows of parietal tubes precludes assignment to this genus.
Stratigraphy and Paleoenvironments of the Bouse Formation
The outcrop belt of the Mio-Pliocene Bouse Formation includes more than 3000 km? of discontinuous exposure in western Arizona and southeastern California (Fig. 1). Strata now assigned to the Bouse For- mation were recognized early in the twen- tieth century (e.g., Blanchard 1913), but were not formally named until later regional work by Metzger (1968). The Bouse Formation was recently re-examined by Buising (1988) on whose study the following discussion of stratigraphy and sedimentology is based.
Strata of the Bouse Formation are inter- preted as documenting transgression of what is now the lower Colorado River trough by waters of the proto-Gulf of California, a tec-
VOLUME 102, NUMBER 4
CALIFORNIA
925
MEXICO
Fig. 1.
CALIFORNIA
Distribution of Bouse Formation (cross-hatched) in relation to inferred extent of proto-Gulf outcrop
belt (stippled). Inset shows mountain ranges (stippled) of the lower Colorado River region and location of Mesquite Mountain, type locality of B. canabus (after Buising 1988).
tonically complex marine embayment that occupied the Gulf of California physio- graphic province prior to onset of modern spreading- and transform-generated subsi- dence in that region (Fig. 1). Basal trans- gressive carbonate of the Bouse Formation overlies pre-Bouse fanglomerate, reflecting inundation of an internally drained alluvial system during proto-Gulf subsidence. The basal carbonate is overlain by fine-grained terrigenous clastic strata recording the southward progradation of the ancestral Colorado River delta into the proto-Gulf. These deltaic facies interfinger laterally with a shoreline complex that includes algal tufa,
clastic limestone and barnacle coquina, as well as coarse-grained, terrigenous clastic units derived from the erosion of local is- land and basin-margin highs. The Bouse Formation is overlain by, and interfingers with trough cross-bedded cobble conglom- erates, informally called the Colorado River gravels, that are characterized by a clast as- semblage derived from the Colorado Pla- teau, and are interpreted as recording arriv- al of the through-going Colorado fluvial channel in approximately its modern po- sition. Although the age of the Bouse For- mation remains problematic, recent re- evaluation suggests that it was deposited
926
between eight and four million years ago (Buising 1988).
Paleoecology
With the exception of channel lag occur- rences, such as those at Mesquite Mountain, and of isolated occurrences in distributary channel siltstone, the barnacles are usually associated with coarse terrigenous clastic units and algal tufa that mark the shoreline of the proto-Gulf (Buising 1988). The proto- Gulf shoreline, which is preserved in essen- tially its original depositional configuration, was extremely rugged and rocky and, pre- sumably, provided the principal substratum for the barnacles. However, barnacles were not found attached to rocks in the shoreline facies, and the only examples found in life position were attached to algal tufa at a shoreline locality northeast of the type channel lag deposits at Mesquite Mountain (Buising 1988).
The abundance of barnacles, and thinness and morphological variability of their plates, indicate rapid, seasonal growth in waters of lowered salinity. Low salinity waters are also suggested by algal tufa morphology and 87/6 Sr values of tufa and barnacle shell samples (Buising 1988). The crowded, columnar to tulipiform growth habit suggests that hard substrata suitable for settlhement were lim- ited, and that vertical growth was a neces- Sary prerequisite for competition under crowded conditions. In addition, some de- positional environments of the Bouse For- mation suggest that rapid vertical growth was essential to maintaining the individual above the terrigenous clastic detritus being deposited in the delta system.
It is likely that cyprid larvae were carried into the delta by the advancing salt wedge during the dry season, and settled in dense masses on every available hard substratum. Adult size was probably attained within a single year, as at least two size classes are recognized, appearing to represent separate yearly settlement. The parietes and scuta of
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
the larger size class, which attained an av- erage adult height of one cm, are overgrown by smaller individuals averaging about one- half cm in height. Although the limited use of calcium carbonate in the shell is indica- tive of growth in lowered salinities, neither size class shows any evidence of the shell corrosion common in such brackish water species as Balanus eburneus Gould and Fis- tulobalanus pallidus (Darwin). This is par- ticularly interesting as the larger size class appears to have lived for more than a single year and, thus, long enough for corrosion to occur.
Systematics
Family Balanidae Leach, 1817 (Newman & Ross 1976) Subfamily Balaninae Leach 1817 (Newman 1980)
Genus Balanus Da Costa, 1778 Balanus canabus, new species Figs. 2-4
Holotype. —Partially crushed opercular pyramid, USNM 423910.
Type locality.—Bouse Formation, west flank of Mesquite Mountain, La Paz Coun- ty, Arizona.
Diagnosis. —Shell thin, cylindric to tulip- iform, with smooth parietes; radii narrow, with steeply sloping summits and denticu- late sutural edges; alae extremely broad, with horizontal summits and denticulate sutural edges; sheath vesicular; parietal tubes rect- angular, in single row. Scutum very thin, extremely convex or medially sulcate, much taller than broad; external longitudinal striae very fine or absent; articular ridge about one-half length of tergal margin; adductor ridge short, well separated from articular ridge; depressor muscle pit shallow and tri- angular or absent. Tergum very thin, broad, externally convex, with convex carinal mar- gin and narrow, sometimes deep, spur fur- row broadening toward spur; upper carinal margin reflexed inward to form broad in-
VOLUME 102, NUMBER 4 927
Fig. 2. Balanus canabus: a, Interior of lateral plate, paratype USNM 423920, x 27; b, Interior of carinolateral plate, paratype USNM 423921, x11; c, Exterior of tergum, paratype USNM 423922, x11; d, Exterior of tergum (lacking spur), paratype USNM 423923, x12; e, Exterior of scutum, paratype USNM 423924, x 12; f, Articulated opercular pyramid, holotype USNM 423910, x12.
928 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig. 3. Opercular plates of Balanus canabus: a, Exterior of tergum, paratype USNM 423911, x12; b, Exterior of apical half of scutum showing secondary extension of occludent margin (on right side), paratype USNM 423912, x18; c, Interior of scutum, paratype USNM 423913, x12; d-e, Interiors of scuta, paratypes USNM 423914 through 423915, x16; f, Interior of tergum, paratype USNM 423916, x12; g, exterior of scutum, paratype USNM 423917, x12;h, Interior of tergum (lacking spur) showing apical compartment, paratype USNM 423918, x12; i, Interior of scutum, paratype USNM 423919, x14.
VOLUME 102, NUMBER 4 929
Fig. 4. Balanus canabus: a, Articulated lateral and carinolateral plates, paratype USNM 423925, x25; b, Basis of second generation individual attached to scutum, paratype USNM 423926, x19; c, Exterior of cari- nolateral showing distal extension of ala, paratype USNM 423927, x11; d, Carinolateral of cylindric individual, paratype USNM 423928, x11; e, Barnacle hash from channel lag deposit of type locality, paratype lot USNM 423929, x1.5. '
930
ternal shelf; spur slightly longer than wide, basally subtruncate; distance from basiscu- tal angle to spur less than or equal to spur width; basal margin straight to concave on both sides of spur, not deeply excavated.
Material. —Twenty-seven shells with or without opercular plates; over 1000 disar- ticulated compartmental plates; 77 whole or partial scuta; 65 whole or partial terga.
Disposition of types. —Holotype USNM 423910, paratypes USNM 423911 through 423928, and paratype lot USNM 423929 are deposited in the collection of the De- partment of Paleobiology, National Mu- seum of Natural History, Washington, D.C.
Geologic and geographic range.—Late Cenozoic (probably latest Miocene or Plio- cene), Bouse Formation, southeastern Cal- ifornia and western Arizona.
Etymology.—The specific name is de- rived from the Greek kanabos, meaning “‘a mere skeleton,” and refers to the extremely thin shell of this species.
Description. —Shell thin, cylindric to tu- lipiform in adults, high conic in juveniles, with slightly toothed orifice and smooth pa- rietes; a few specimens show traces of broad, longitudinal color stripes on parietes; radii narrow, not sunken, transversely striate, with steeply sloping summits (70°) and thin, finely crenate, sutural edges; alae very broad, composed of proximal and distal segments separated by incised diagonal line and change in growth pattern; horizontal to con- vex summits of alae formed by distal seg- ments; alar sutural edges finely crenate; sheath about one-third length of compart- mental plate, vesicular, without vesicles in furrow below dependent lower margin; in- ternal parietal ribs prominent, regularly spaced, flat-topped, extending from base to sheath, finely crenate basally; longitudinal parietal tubes large, rectangular, in single row, crossed by numerous, closely spaced, transverse septa; basis calcareous, thin in center and thickening appreciably toward margin, with fine radial tubes bearing few transverse septa.
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Scutum thin, narrow, with basal margin two-thirds to three-fourths length of tergal margin; exterior either flat with median lon- gitudinal sulcus or broadly convex; apex often twisted slightly toward tergum; exte- rior ornamented by broadly spaced major growth ridges forming denticles on occlu- dent margin and numerous, fine, minor growth ridges between major ridges; exte- rior growth ridges usually crossed by ex- tremely fine longitudinal striae in central part of plate; tergal margin straight to slight- ly concave, broadly reflected from 45° to 60° from plane of plate; occludent margin con- vex, sharply inflected in upper half; inflected area of occludent margin often enlarged by secondary growth in apical region; basal margin straight to slightly sinuous; articular ridge prominent, usually about one-half length of tergal margin, forming a broad, flat shelf in upper half, becoming narrow and sharp in lower half, and ending in sharp, downturned hook, partially reflected over deep, narrow, articular furrow; adductor ridge sharp, erect, highest adjacent to ad- ductor muscle pit, variable in length, but usually short, centrally located, and well re- moved from articular ridge; adductor mus- cle pit shallow, small, round to oval; a shal- low, triangular pit often present on tergal side of adductor ridge; depressor muscle pit either moderately large, shallow and trian- gular or absent; interior of apical half of scutum rugose.
Tergum thin, convex, broad, with basal margin one-half to two-thirds length of plate; carinal margin convex, usually protuberant apically, sharply inflected in apical half forming internal apical chamber; scutal margin straight to concave, sharply inflect- ed to form flat shelf; basal margin not deeply embayed, straight to concave on either side of spur; exterior ornamented by closely spaced major, and a few interspersed minor, growth ridges; broad area along carinal mar- gin marked by sharply upturned growth lines; carinal side of exterior bearing fine radial striae partially reflecting position of
VOLUME 102, NUMBER 4
internal depressor muscle crests; spur fur- row narrow in apical half, widening toward spur, often very deep; sides of spur furrow not marked by incised lines and not infold- ed; spur with nearly parallel sides and sub- truncate base, placed between one-half its width and distance equal to its width from basiscutal angle, moderately long, about two- ninths length of plate, and narrow, less than one-third width of basal margin; articular ridge nearly erect, high, thin, concave on scutal side, little more than one-half length of scutal margin, and continuous with in- flected part of carinal margin forming apical chamber; articular furrow broad, moder- ately deep; depressor muscle crests short, well developed, closely spaced, usually five in number; interior or tergum markedly ru- gose in apical half.
Discussion. — Aside from the marked thinness of the compartmental and oper- cular plates, the features that set the new species apart from other members of the Balanus amphitrite complex are those ap- parently associated with the broadening of the orifice with continued vertical growth under crowded conditions. Development of the tulipiform shell is accomplished through secondary distal growth of the alae rather than widening of the radii. In fact, the steep- ly sloping radius is in contact with the paries of the adjacent compartment only in the basal third of the shell wall. The opercular pyramid, which occupies most of the orifice, accommodates the increase in orifice di- ameter through secondary horizontal growth on either side of the aperture formed by the occludent margin of the scutum and the car- inal margin of the tergum. This horizontal growth produces a chamber in the apex of the opercular pyramid which is particularly apparent on the interior of the tergum. A similar chamber 1s seen in terga of the extant western Pacific species Fistulobalanus al- bicostatus (Pilsbry) and F. kondakovi (Tar- asov & Zevina). The growth form and re- sulting morphology of B. canabus appear to be related both to competition for living
931
space under crowded conditions in an area of limited substrata, and to maintenance of the individual above the sediment that was being deposited in the delta environment.
Although B. canabus shares many mor- phological features with fossil and extant species of Fistulobalanus, it lacks multiple rows of parietal tubes in the shell wall and, therefore, cannot be included in the genus. The similarity of the new species to Fistu- lobalanus is striking, however. All of the extant species of the genus are found in brackish waters (Henry & McLaughlin 1975; the habitats of F. abeli(Lamy & André) and of F. patelliformis (Bruguiére) are un- known). Some specimens of F. pallidus and F. kondakovi lack subsidiary parietal tube rows, and these species often occur in crowded, thin-walled, cylindric masses in brackish water environments (Stubbings 1963; Henry & McLaughlin 1975). The morphology of the opercular and compart- mental plates of B. canabus is typical of Fistulobalanus, especially of F. pallidus, F. albicostatus and F. kondakovi, and bears less resemblance to species of Balanus. The ab- sence of subsidiary parietal tubes may be related to the crowded, columnar growth habit and the conservative use of calcium carbonate in shell construction.
It is possible that B. canabus was derived from a Fistulobalanus ancestor. The genus extends back to the middle Miocene in the North Atlantic basin (Zullo 1984), and is known from the Pleistocene of Japan (Ya- maguchi 1980). However, the two extant species of Fistulobalanus known from the eastern Pacific, F. dentivarians (Henry) and F. suturalis (Henry), bear less resemblance to Balanus canabus than do the previously mentioned western Pacific and Atlantic Ocean species.
Among the species of the B. amphitrite complex with a single row of parietal tubes, only the extant western Atlantic species B. eburneus and B. subalbidus Henry possess a vesicular sheath. Like the species of Fis- tulobalanus, both are inhabitants of brack-
952
ish waters. Balanus eburneus differs in hav- ing radii with broader, gently sloping summits; prominent, deeply incised exter- nal radial striae on the scutum; a scutal ad- ductor ridge that is almost confluent with the articular ridge; and a tergum with a spur fasciole and a markedly concave and usually deeply embayed basal margin on the carinal side of the spur. Balanus subalbidus differs in having a broader scutum with the ad- ductor and articular ridges nearly confluent, and a tergum with a spur fasciole and a broader spur.
Locality Descriptions
The type lot of B. canabus was obtained from the fine-grained, terrigenous-clastic, deltaic facies of the Bouse Formation on the west flank of Mesquite Mountain, La Paz County, Arizona. At this locality, trans- ported barnacles and barnacle plates occur as a lag deposit in a northerly-trending channel approximately 5 m wide and slight- ly less than 1 m deep. The channel, filled with green mud, is located at the western terminus of a pink siltstone bed with per- vasive, westward-migrating, meter-high, trough cross-beds. The entire complex is in- terpreted as representing progressive west- ward migration of a tidally-influenced dis- tributary channel, followed by channel abandonment and infilling by green hypoxic mud (Buising 1988).
Other specimens of B. canabus examined for this study were collected in 1961 by Blakemore E. Thomas, San Diego State University, from Bouse Formation out- crops on the north end of the Riverside Mountains, Riverside County, and the Palo Verde Mountains, Imperial County, Cali- fornia.
Acknowledgments
We thank the people and council of the Colorado River Indian Tribes for allowing
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access to tribal lands in the Mesquite Moun- tain area for mapping and sampling pur- poses. Special thanks are extended to Charles Lamb, Weldon Johnson, Curtis Martin and the Colorado River Indian Tribes Museum.
Literature Cited
Blanchard, R. C. 1913. The geology of the western Buckskin Mountains, Yuma County, Arizo- na.— Columbia University Contributions of the Geology Department 26:1-80.
Buising, A. V. 1988. Deposition and tectonic evo- lution of the northern proto-Gulf of California and lower Colorado River, as documented in the Mio-Pliocene Bouse Formation and brack- eting units, southeastern California and western Arizona. Unpublished Ph.D. dissertation, Uni- versity of California at Santa Barbara, 196 pp.
Henry, D. P., & P. A. McLaughlin. 1975. The bar- nacles of the Balanus amphitrite complex (Cir- ripedia, Thoracica). — Zoologische Verhandelin- gen 141, 254 pp.
Metzger, D. G. 1968. The Bouse Formation (Plio- cene) of the Parker-Blythe-Cibola area, Arizona and California.— United States Geological Sur- vey Professional Paper 600-D:D126-D136.
Stubbings, H.G. 1963. Cirripedia of the tropical south Atlantic coast of Africa.—Expédition Océano- graphique Belge dan las Eaux Cotiéres Afn- caines de l’Atlantique Sud (1948-1949), Institut Royal des Sciences Naturelles de Belgique, Ré- sultats Scientifiques 3(10), 39 pp.
Yamaguchi, T. 1980. A new species belonging to the Balanus amphitrite Darwin group (Cirripedia, Balanomorpha) from the late Pleistocene of Ja- pan.—Journal of Paleontology 54:1084-1101.
Zullo, V. A. 1984. New genera and species of bal- anoid barnacles from the Oligocene and Mio- cene of North Carolina.— Journal of Paleontol- ogy 58:1312-1338.
(VAZ) Department of Earth Sciences, University of North Carolina at Wilming- ton, North Carolina 28403; (AVB) Depart- ment of Geological Sciences, California State University, Hayward, California 94542.
PROC. BIOL. SOC. WASH. 102(4), 1989, pp. 933-946
FIRST INGOLFIELLIDS FROM THE SOUTHWEST PACIFIC (CRUSTACEA: AMPHIPODA) WITH A DISCUSSION OF THEIR SYSTEMATICS
James K. Lowry and Gary C. B. Poore
Abstract. —Two new species of ingolfiellid amphipods, [ngolfiella australiana and J. bassiana are described from the continental shelf of Bass Strait, south- eastern Australia. Ingolfiella australiana is similar to some species of the sub- genus Trianguliella Stock, 1976. Ingolfiella bassiana cannot be placed easily into any known subgenus, but shows some similarities to species described from the West Indies and the Canary Islands in the subgenus Gevgeliella Kar- aman, 1959. A re-analysis of sexually dimorphic characters casts doubt on current generic and subgeneric concepts. It is concluded that the ingolfiellidean ““eye-lobe”’ is not homologous with the dorsal pedunculate eyestalk of other peracaridans because of its lateral position. This and other evidence from the Metaingolfiellidae place the ingolfiellidean families within the Gammaridea. The ingolfiellidean maxilliped, carpochelate gnathopod 2, and the entire telson suggest similarities to the leucothooid gammarideans. Retention of shared ple- siomorphic characters such as a maxilliped without an ischial endite and an
entire telson indicates an early derivation from the amphipodan stem.
The ingolfiellidean amphipods comprise about 30 species in 2 families, Ingolfiellidae and Metaingolfiellidae. Ruffo (1970), Stock (1976, 1977, 1979), Ronde-Broekhuizen & Stock (1987), and Dojiri & Sieg (1987) have reviewed the systematics and zoogeography of the group. Although widely distributed from the deep sea to fresh water and hy- pogean habitats, ingolfiellideans have not previously been recorded from the South- west Pacific.
Two new species of small ingolfiellids, de- scribed herein, were discovered in 2 of over 200 lots of amphipods sorted from benthic samples taken from the continental shelf and slope of Bass Strait, southeastern Australia. Material is lodged in the Museum of Vic- toria, Melbourne (NMV) and the Australian Museum, Sydney (AM).
The Ingolfiellidae is a conservative family of three genera (Stock 1976, Ruffo 1985). Few specific differences occur in overall body shape, antennae, mouthparts or peraeo-
pods. Nevertheless, Stock (1976) erected five subgenera within /ngolfiella, the largest ge- nus. These are separated largely on the basis of sexually dimorphic differences in the sec- ond gnathopod and pleopods. We discuss some of the taxonomic problems associated with these subgenera. We also discuss the phylogenetic placement of the ingolfiellid- ean group.
Family Ingolfiellidae Hansen, 1903
Ingolfiella australiana, new species Figs. 1-4
Type specimens.—Holotype, male, 2.3 mm, NMV J12851 with 2 slides, paratype, lemale.. 2.2 mim INMY 12850, with, 2 slides, 34 km SW of King Island, 40°26.7'S 143°41.4’E, Bass Strait, Australia, 85 m, sandy shell, Smith-MclIntyre grab, R. Wil- son et al. on RV Jangaroa, 22 Nov 1981 (NMV station BSS 198).
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Fig. 1.
Ingolfiella australiana, holotype male, NMV J12851; southwest of King Island, Bass Strait, Australia.
Abbreviations are as follows: Al, A2 = Antennae 1-2; D = Penes; G1, G2 = Gnathopods 1-2; H = Head; LL = Lower lip; 1 = Left; MD = Mandible; MX1, MX2 = Maxillae 1-2; MP = Maxilliped; P3—P7 = Peraeopods 3-7; PLI-PL3 = Pleopods 1-3; r = Right; T = Telson; U1-U3 = Uropod 1-3; UR3 = Urosomite 3.
Description of holotype. — Body elongate, all segments laterally compressed. Head, anterodorsal margin angular, without ros- trum; “‘eye-lobe’’ semicircular, small. Pe- raeonite 1 about half as long as head; pos- teroventral margin oblique; much deeper anteriorly than posteriorly such that perae- onites 1 and 2 are separated by a waist. Peraeonites 2 to 7 increasing in depth pos- teriorly. Pleonites 1 to 3 with posteriorly rounded epimera. Urosomites 1 and 2 not markedly differentiated from pleonites, of
similar length; urosomite 3 with lateral plates enclosing base of telson and uropod 3.
Antenna 1, peduncular article 1 as long as head; article ratio 1.0:0.4:0.3; flagellum slightly less than half length of peduncle, five articles, last minute; accessory flagel- lum just longer than article 1 of flagellum, three articles, last minute. Antenna 2, pe- duncle as long as peduncle of antenna 1; flagellum of five articles, about one-third length of peduncle.
Left mandible, incisor with four teeth; la-
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Fig. 2. Ingolfiella australiana, holotype male, NMV J12851 (circle star); paratype “‘female,’””» NMV J12850 (star); southwest of King Island, Bass Strait, Australia. For Abbreviations see Fig. 1.
cinia mobilis as broad as incisor, with five teeth; spine row of three curved, denticulate spines; molar a long triangular blade with minutely denticulate margin. Maxilla 1, in- ner plate subquadrate, with four setae; outer plate with three strong cuspidate spines in anterior row, two denticulate spines in pos- terior row, and one well developed curved comb-spine medially; palp of two articles, with three apical plumose setae. Maxilla 2, inner plate with one subterminal seta and four terminal setae; outer plate with five setae. Maxilliped, basal endite with one sub- apical and one apical seta; palp articles 1 to 4 with three, one, one, and one mesial setae respectively, article 5 with long falcate un- guis, seta at midlength and at base of unguis.
Gnathopod | carpochelate, palm strongly
oblique; coxa inserted at anteroventral cor- ner of peraeonite; carpus 2.2 times as long as wide, palm with three proximal spines and eight setae, without teeth; dactylus with four serrations. Gnathopod 2 carpochelate, palm slightly oblique; carpus 1.6 times as long as wide; palm defined by strong curved spine, distally with triangular tooth and quadrate tooth separated by narrow notch, with one spine and four setae (one seta prox- imal to definitive palm spine); propodus with two triangular blades posteriorly, distal blade larger; dactylus with three teeth on posterior margin, dactylus longer than palm, extend- ing Over carpus.
Peraeopods 3 and 4, dactylus with two distal setae and cylindrical bifid unguis. Pe- raeopods 5 to 7, basis slightly broadened in
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Fig. 3. Ingolfiella australiana, holotype male, NMV J12851 (circle star); paratype “female,” NMV J12850 (star); southwest of King Island, Bass Strait, Australia. For Abbreviations see Fig. 1.
peraeopod 5, becoming progressively nar- rower in peraeopods 6 and 7; dactylus stout, curved, unguis not defined.
Pleopod 1 cylindrical with two setae; pleopods 2 and 3 subtriangular.
Uropod 1, peduncle 1.3 times as long as inner ramus; inner ramus with lateral row of 10 long setae, 4 apical spines in dished
tip, edges of apex finely denticulate; outer ramus 2.2 times as long as inner ramus, with 2 distal setae. Uropod 2, peduncle with 5 oblique rows of (proximal to distal) 11, 9, 17, 15, 12 spines mesially (2 spines between rows 4 and 5); inner ramus % length of pe- duncle, with 3 proximal setae; outer ramus shorter than inner with 2 setae near mid-
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Fig. 4. Ingolfiella australiana, holotype male, NMV J12851 (circle star); paratype “female,” NMV J12850; southwest of King Island, Bass Strait, Australia. For Abbreviations see Fig. 1.
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length. Uropod 3 with one ramus; peduncle with two setae, ramus short, broad, with distal seta. Telson subtriangular, with pair of long dorsal setae.
Variation. —Paratype, 2.2 mm. “‘Eye- lobe”’ reaches to article 2 of antenna 2. Gnathopod 2, palm convex, triangular and blade-like teeth poorly defined, with five se- tae (no seta proximal to definitive palm spine); propodus, teeth blunt. Pleopod 1 subtriangular with one seta. Uropod 1, pe- duncle 1.6 times as long as inner ramus, with longitudinal-oblique row of four setae, ventrally with a dense brush of stout, short setae; inner ramus with two setae at mid- point; outer ramus longer than inner ramus, with four setae. Uropod 2, peduncle with 5 oblique rows of (proximal to distal) 2, 10, 13, 15, 13 spines mesially.
Etymology. —For Australia.
Remarks.—The holotype of Ingolfella australiana is most similar to the South Af- rican species J. (Trianguliella) berrisfordi Ruffo, 1974 and the West Indian species J. (T.)grandispina Stock, 1979. Ingolfiella ber- risfordi has two setae on the inner plate and five spine-teeth on the outer plate of maxilla 1; pleopod 1 is subtriangular; pleopods 2 and 3 are broader and distally truncate; uro- pod 1| hasa shorter inner ramus; and uropod 2 has a basofacial hook on the peduncle. Ingolfiella grandispina has the dactylus and unguis separate on peraeopods 5 to 7; pleo- pods broad and distally truncate, and a slightly better developed ramus on uro- pod 3.
Ingolfiella australiana is different from the other three species assigned to Trianguliel- la: I. (T.) manni Noodt, 1961 which has long, slender endites on the maxilliped and three spine-rows on the peduncle of uropod 2; I. (T.) macedonica Karaman, 1959 in which the palmar spine and the dactylus of gnathopod 2 are not enlarged; and J. (7.) thibaudi Coineau, 1968 in which males have only pleopod 1 and females have no pleo- pods.
Uropod 1 of the paratype of this species is unusual in the possession of a ventral
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peduncular row of stout setae not reported in any other ingolfiellidean.
Ingolfiella bassiana, new species Figs. 5-8
Type specimens.—Holotype male, 1.8 mm, NMV J13124 with 2 slides, paratype “female,” 1.8 mm, NMV J13119 with 2 slides; 3 paratype males, 1.7 to 1.9 mm J13125, J13126, AM P38458; 6 paratype “females” (without oostegites), 1.1 to 1.8 mm NMV J13120, with 2 slides, to J13123, AM P38459, 75 km WSW of Cape Otway, 39°02.4’S 142°37.8’E, Bass Strait, Australia, 121 m, shelly sand, dredge, G.C.B. Poore on HMAS Kimbla, 9 Oct 1980 (NMYV sta- tion BSS 64).
Description. — Based on holotype, male, 1.8 mm, and paratypes, “females,” 1.8 mm, NMV J13119 and 1.7 mm, NMV J13120. Body elongate, all segments laterally com- pressed. Head, anterodorsal margin round- ed, without rostrum; “eye-lobe”’ semicir- cular, small. Peraeonite 1 about half as long as head; posteroventral margin oblique; deeper anteriorly than posteriorly such that peraeonites 1 and 2 only weakly separated. Peraeonites 2 to 7 increasing in depth pos- teriorly. Pleonites 1 to 3 with posteriorly rounded epimera. Urosomites 1 and 2 not markedly differentiated from pleonites, of similar length; urosomite 3 with lateral plates enclosing base of telson and uropod 3.
Antenna 1, peduncular article 1 as long as head; article ratio 1.0:0.4:0.3; flagellum of four articles, slightly less than half length of peduncle; accessory flagellum of two ar- ticles, last longer, reaching midlength of ar- ticle 2 of flagellum. Antenna 2, peduncle as long as peduncle of antenna 1; flagellum of five articles, about one-third length of pe- duncle.
Mouthparts of juvenile (NMV J13120) (Fig. 6). Left mandible, incisor with three cusps; lacinia mobilis as broad as incisor, with five cusps; spine row of three curved, denticulate spines; molar a long triangular blade with minutely serrate margin. Right
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/ (i \
959
REG
Fig. 5. JIngolfiella bassiana, paratype, “female,” 1.8 mm, NMV J13119; southwest of Cape Otway, Bass
Strait, Australia.
mandible, incisor with four cusps on two overlapping blades; lacinia mobilis almost as broad as incisor with denticulate margin; two denticulate spines, and molar same as left. Maxilla 1, inner plate distorted, with three setae apparent; outer plate with three strong cuspidate spines in anterior row, two denticulate spines in posterior row, and one well developed curved comb-spine mesial- ly; palp of two articles, with one naked and two plumose apical setae. Maxilla 2 un- known. Maxilliped, basal endite with two apical setae; palp articles 1 to 4 with two, one, one, and one mesial setae respectively, article 4 with oblique row of slender setae, article 5 with long falcate unguis, seta at midlength and distally.
Gnathopod | carpochelate, palm oblique; coxa at anterior of peraeonite; carpus 2.2 times as long as wide, palm with three prox- imal spines, three weaker spines and three setae, without teeth; dactylus with three spines along posterior margin. Gnathopod 2 carpochelate, palm nearly transverse; car- pus 1.6 times as long as wide; palm defined
by one reversed pectinate seta and three strong complex spines, (mesial spine com- plexly bifurcate, two lateral spines simpler), palm obliquely transverse, with a triangular tooth at midlength, with three stout setae and two finer setae laterally and three setae mesially, propodus with a triangular blade posteriorly; dactylus with three teeth on in- ner margin; dactylus as long as palm, not extending over carpus.
Peraeopods 3 and 4, dactylus with two distal setae and cylindrical bifid unguis. Pe- raeopods 5 to 7 becoming narrower poste- riorly; dactylus stout, curved, unguis not de- fined.
Pleopods subtriangular, with notch on oblique margin; pleopods 1 and 2 each with two terminal setae.
Uropod 1, peduncle 1.4 times as long as inner ramus with lateral, plumose setae; in- ner ramus with a mesial row of four long setae and four distal spines; outer ramus 0.6 times as long as inner ramus, with one seta. Uropod 2, peduncle with proximoventral hook and 4 obliquely transverse rows of
940 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
és zy,
xU1 a. Soman are te =
rT A ETE Ss ogy
Fig. 6. Jngolfiella bassiana, holotype, male, 1.8 mm, NMV J13124 (closed star); paratype, “female,” 1.7 mm, NMV J13120 (closed circle); paratype, “female,” 1.8 mm, NMV J13119 (circle star); southwest of Cape Otway, Bass Strait, Australia. For Abbreviations see Fig. 1.
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Fig. 7.
Ingolfiella bassiana, holotype, male, 1.8 mm, NMV J13124 (closed star); paratype “female,” 1.8 mm
NMV J13119 (circle star); paratype, male, 1.8 mm, NMV J13125 (open star); southwest of Cape Otway, Bass
Strait, Australia. For Abbreviations see Fig. 1.
(proximal to distal) 12, 16, 13, 8 spines (those ventrally in third and fourth rows apically complex); rami 1-articulate, equal, 0.6 times length of peduncle, inner ramus with 4 setae; outer ramus with transverse row of 3 spines plus 3 setae. Uropod 3 with one ramus; peduncle with two lateral setae,
ramus short, broad, with long distal seta. Telson subtriangular, with pair of long dor- sal setae.
“Females.” —As in male but: gnathopod 2 carpus palm defined by two spines, palm oblique and with distal notch; propodus with straight posterior margin; dactylus with three
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Fig. 8. Australia. For Abbreviations see Fig. 1.
oblique proximal teeth. Pleopods without distal setae. Uropod 1 peduncle without lat- eral plumose seta. Uropod 2 peduncle with- out proximoventral hook, with 4 mesial transverse rows of (proximal to distal) 9, 15, 12, 8 spines.
Etymology. —For Bass Strait.
Remarks. —The possession of triangular pleopods and a basofacial hook on uropod 2 allies I. bassiana to the five West Indian and Canary Islands species in the subgenus Gevgeliella discussed by Ronde-Broekhu- izen & Stock (1987). All differ from the type species I. petkovskii Karaman, 1957, and the two other Mediterranean species of Gev- geliella (I. catalanensis Coineau, 1963 and I. vandeli Bou, 1970) which do not have
Ingolfiella bassiana, holotype, male, 1.8 mm, NMV J13119; southwest of Cape Otway, Bass Strait,
these characters. The complexity of the pal- mar spines on male gnathopod 2 and apical setae on pleopods 1 and 2 distinguish J. bassiana from all of these species.
Stock (1979) has suggested that the re- versed seta on gnathopod 1, the apical setae on pleopod 1 and the basofacial hook on uropod 2 are all male characters of Gevge- liella. Our holotype has all of these char- acters and paired penial processes. Our “‘fe- male” specimens do not, thus confirming Stock’s presumptions.
The complex nature of the mesial spines on uropod 2 (shown here in female) have been illustrated (for J. fuscina) previously only by Dojiri and Sieg (1987). Males of J. bassiana possess a long plumose seta on the
VOLUME 102, NUMBER 4
peduncle of uropod 1, a character never be- fore reported.
Discussion
Sexual dimorphism and subgenera. — Sexual dimorphism was used to help define two of the five subgenera recognized by Stock (1976). It was not used to define the three other subgenera because the data were, and still are, not available. In the absence of information about sexual dimorphism, dif- ferences between some subgenera are un- convincing (for example, between /ngolfiel- la and Hansenliella). Dojiri & Sieg (1987) also questioned the value of subgenera for the same reason. Stock (1976) used only two dimorphic characters, gnathopod 2 and pleopod 1. We have examined 8 characters which show sexual dimorphism, and sur- veyed the literature for 25 species. For about half of these species males are not known or cannot be distinguished within the avail- able material. Most of these species fall into subgenera for which sexually dimorphic characters are not considered to be diag- nostic.
Some characters are unique or not widely distributed and thus of little phylogenetic value. These apomorphic forms include:
1. Complex palmar spines in the male gnathopod 2 (only J. bassiana);
2. Loss of pleopods 2 and 3 in males and 1 to 3 in females (only J. thibaudi Coineau, 1968 and I. catalanensis Coineau, 1963, Ruffo, pers. comm);
3. Complex distal seta on the peduncle of male uropod 1 (in J. fuscina Dojiri & Sieg, 1987 and Trogloleleupia eggerti Ruffo, 1951);
4. Row of stout ventral setae in ‘‘female”’ uropod 1 (only J. bassiana).
Other characters are more widespread and may indicate phylogenetic affinities. These include:
5. Reversed seta on carpus of gnathopod 2 (occurs in all species assigned to Gevge- liella and Trogloleleupia eggerti);
6. Male pleopod | digitiform (most species of Trianguliella and G. catalanensis;
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7. Male apical setae on pleopod 1 (all species for which males are known);
8. Uropod 2 with male peduncular ba- sofacial hook (most, but not all species of Gevegeliella, I. bassiana, Trianguliella ber- risfordi and Trogloleleupia eggerti).
Among the widespread sexually dimor- phic characters there is overlap between genera and subgenera and the existing ar- rangement is not well supported by this in- vestigation. Until more males are known and more species are discovered subgeneric classification within the Ingolfiellidae is un- stable.
It is probable that some form of sexual dimorphism is plesiomorphic in ingolfiel- lideans (Dojiri and Sieg 1987). When the plesiomorphic type is established and the derived types are understood, then these forms may be used to define species groups.
Subordinal status. — Recently Bowman & Abele (1982), based on the extensive ar- guments of Dahl (1977), abandoned the suborder Ingolfiellidea and placed both of its families in the suborder Gammaridea. Dahl looked at several characters from ear- lier works by Hansen (1903), Ruffo (1951), and Siewing (1963). He concluded that of these characters only the “eye-lobe”’ could be used to define the group at the subordinal level.
The presence of an “‘eye-lobe”’ in ingolfi- ellideans was first noted by Hansen (1903). Dahl (1977) seemed unconvinced that this structure, a small scale on the side of the head of some species, is a rudimentary stalked eye. He noted: “Its functional sig- nificance 1s unknown, and it contains no dioptric and apparently also no nervous ele- ments. Its location, however, corresponds well with that of the lobate rudiment of the compound eye in Gammarus...” Bowman (1984), citing Dahl, was equally equivocal: ‘“‘Whether these lobes represent eyestalks is questionable.”’
Only three of seven peracaridan orders contain species with unambiguous eye- stalks. In the Mysidacea (numerous species) the eyestalks carry well developed terminal
944
eyes. In the Spelaeogriphacea, where only two living species are known (Spelaeogri- phus lepidops and Potiicoara brasiliensis), the lobe has no pigment or optic structure (Gordon 1957, Pires 1987). In Mictacea one of the three species known (Mictocaris hal- ope) has pyriform eyestalks lacking visual elements (Bowman & Iliffe 1985). In the only pancaridan order, four of the six Ther- mosbaenacea genera have plate-like eye- stalks but lack pigment (Bowman & Iliffe 1986, 1988). Eyestalks in the syncarid fam- ily Anaspididae are similar to those in pera- caridans. The spelaeogriphaceans, the ther- mosbaenaceans and the mictacean are troglobitic so it is not surprising that they are blind. What is notable is that the eye- stalks in all these groups attach obliquely above the first antenna near the base of the rostrum, a position and attitude very dif- ferent from that of the ingolfiellidean scale.
It seems probable that the hinged ingol- fiellidean “‘eye-lobe’’ is the anterolateral margin of the head, often produced at this point in amphipods, whether it bears a ses- sile eye or not. A similar situation is seen in the tanaidacean genus Heterotanais in which the eyes occur on hinged lobes (Sars 1896). These are in the same position, ven- tral to the antennae, as the normally sessile eyes of other tanaidaceans.
We think that the so-called “‘eye-lobe’’ of ingolfiellideans is not homologous to that of other peracaridans or of other malacos- tracans. The “‘eye-lobe”’ therefore is not of subordinal importance.
Dahl (1977) did not take into account the single species of the second ingolfiellidean family, Metaingolfiella mirabilis (Ruffo, 1969). The plesiomorphic characters of this species provide further evidence for the placement of the group within the Gam- maridea. The pleopods of M. mirabilis are biramous, typical of most Gammaridea, and quite unlike the reduced form of the Ingolfi- ellidae. On peraeopods 3 and 4 the unguis is undifferentiated, as in Gammaridea, and not specialized as in Ingolfiellidae. The
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mandibular molar is a fixed process, not articulating as in Ingolfiellidae. The exis- tence of Metaingolfiella removes any argu- ment for retaining a separate suborder.
Relationships within the Gammaridea. — Many of the unusual characters of the in- golfiellideans are associated with their in- terstitial way of life and are paralleled by other crustaceans in similar habitats. The maxilliped, gnathopods | and 2, and the telson may help place the group within the Gammaridea.
In considering the polarity of character states in the following discussion we have use the isopods for outgroup comparison. Most peracaridan groups, including the iso- pods, have a maxilliped with a basal endite and a palp of five articles, a condition which we consider plesiomorphic among peracar- idans. This state also occurs in ingolfiellid- eans and in all members of the gammarid- ean families Cressidae, Leucothoidae, Pagetinidae, Stenothoidae and Thaumatel- sonidae. This is in contrast to the more widely held view that a maxilliped with both basal and ischial endites (the usual gam- maridean state) is plesiomorphic (Bousfield 1979). The small ischial endite which occurs in some of these families is thus a develop- ing rather than a reducing condition.
Carpochelate gnathopods are an impor- tant character defining the ingolfiellideans. A similar condition is known in leucoth- oids, some corophioids, the deep sea par- daliscid Eperopeus abyssicola Mills, 1967, and is widespread among the hyperiideans. There can be little doubt that carpochely has arisen more than once, but the leucoth- oids are the only group which share the primitive maxilliped with the ingolfielli- deans.
The telson is entire in all peracaridan groups. The only exception is some Am- phipoda. We consider it to be the plesio- morphic condition among the isopods and the amphipods. It is entire in the ingolfiel- lideans, leucothoids, corophioids and rep- resentatives within other families. We rec-
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ognize that within the Amphipoda the entire telson has been secondarily derived many times. This contrasts with the view of Bous- field (1979) that the laminate cleft telson is plesiomorphic.
The evidence for phylogenetic relation- ships of the ingolfiellideans within the Gam- maridea is meager. The ingolfiellideans and the leucothoids are the only living amphi- pods without an ischial endite on the max- illiped. This is the usual form in all other peracaridans, and indicates that the evolu- tion of ischial endites occurred after the am- phipods arose as a distinct group. The in- golfiellideans and the leucothoids both have carpochelate gnathopods. These observa- tions may be interpreted as a phylogenetic link between the ingolfiellideans and: the leucothoids and may indicate that both groups were derived early and close from the amphipodan stem.
Acknowledgments
We thank Jean Just who first drew our attention to the presence of ingolfiellids in the Bass Strait material. We thank Alan Myers and Tom Bowman for recent dis- cussions on the higher classification of the Amphipoda. We thank Sandro Ruffo, Tom Bowman and an anonymous referee for many helpful comments on the manuscript. We thank Roger Springthorpe for compos- ing and inking our drawings. The Bass Strait Survey was funded in part by a Marine Sci- ences and Technologies Grant to the Mu- seum of Victoria.
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. 1959. Uber die Ingolfielliden Jugoslawiens. —
Bioloski Glasnik 12:63-80.
Mills, E. L. 1967. Deep-sea Amphipoda from the western North Atlantic Ocean. I. Ingolfiellidea and an unusual new species in the gammaridean family Pardaliscidae.— Canadian Journal of Zo- ology 45:347-355.
Noodt, W. 1961. Estudios sobre Crustaceos Chilenos de aquas subterraneas, 2. Nueva Jngolfiella de aquas subterraneas limnicas de las Lomas de
946 Paposo en el Norte de Chile (Crustacea, Am- phipoda).—Investigaciones Zooldgicas Chile- nas 7:7-15.
Pires, A. M. S. 1987. Potiicoara brasiliensis: a new
genus and species of Spelaeogriphacea (Crus- tacea: Peracarida) from Brazil with a phyloge- netic analysis of the Peracarida.—Journal of Natural History 21:225-238. Ronde-Broekhuizen, B. L. M., & J. H. Stock. 1987. Stygofauna of the Canary Islands, 1. A new in- golfiellid (Crustacea, Amphipoda) with West In- dian affinities from the Canary Islands. — Archiv fiir Hydrobiologie 110(3):441-450. Ruffo, S. 1951. Ingolfiella Leleupi n. sp. nuovo an- fipodo troglobio del Conga Belga (Amphipoda- Ingolfiellidae).—Revue de Zoologie et Bota- nique Africaines 44(2):189-209.
1969. Descrizione di Metaingolfiella mira- bilis n. gen. n. sp. (Crustacea, Amphipoda, Metaingolfiellidae fam. nova) delle acque sot- terranee del salento nell’Italia Meridionale. — Memorie del Museo Civico di Storia Naturale di Verona 16:239-260.
1970. Considerations a propos de la syste-
matique et de la biogeographie des Ingolfielles (Crustacea Amphipoda).—Livre du Centenaire Emile G. Racovitza, pp. 223-230. . 1974. Studi sui Crostacei anfipodi LX XVII. Nuovi Anfipodi interstiziali delle coste del Sud Africa.—Atti dell’Istituto Veneto di Scienze, Lettere ed Arti 1973-74, (Classe di Scienze Ma- tematiche e Naturali) 132:399-419.
1985. Un nuovo Ingolfiellideo delle acque
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sotterranee della Namibia: Stygobarnardia ca- prellinoides n. gen. n. sp.— Atti della Societa It- aliana di Scienze Naturali, edel Museo Civico di Storia Naturale di Milano 126(1/2):43-53.
Siewing, R. 1963. Zur Morphologie der aberranten Amphipodengruppe Ingolfiellidae und zur Be- deutung extremer Kleinformen fiir die phylo- genie. — Zoologischer Anzieger 171:76—91.
Sars, G. O. 1896. An account of the Crustacea of Norway with short descriptions and figures of all the species. Vol. 2 Isopoda. Parts I, II, pp. 1-40, pls. I-X VI. Bergen Museum, Bergen.
Stock, J. H. 1976. A new member of the crustacean
suborder Ingolfiellidea from Bonaire with a re- view of the entire suborder.—Studies on the
Fauna of Curacao and other Caribbean Islands
50(164):56-75.
1977. The zoogeography of the crustacean suborder Ingolfiellidea with descriptions of new West Indian taxa. — Studies on the Fauna of Cu- racao and other Caribbean Islands 55:131-146.
1979. New data on taxonomy and zooge- ography of ingolfiellid Crustacea. — Bijdragen tot de Dierkunde 45(2):181-190.
(JKL) Division of Invertebrate Zoology, Australian Museum, P.O. Box A285, Syd- ney South, NSW 2000, Australia; (GCBP) Department of Crustacea, Museum of Vic- toria, Swanston Street, Melbourne, Victoria 3000, Australia.
PROC. BIOL. SOC. WASH. 102(4), 1989, pp. 947-959
ALLOCRANGONYCTIDAE AND PSEUDOCRANGONYCTIDAE, TWO NEW FAMILIES OF HOLARCTIC SUBTERRANEAN AMPHIPOD CRUSTACEANS (GAMMARIDEA), WITH COMMENTS ON THEIR PHYLOGENETIC AND ZOOGEOGRAPHIC RELATIONSHIPS
John R. Holsinger
Abstract.—Two new families of gammaridean amphipod crustaceans, AI- locrangonyctidae and Pseudocrangonyctidae, are described. The former is com- posed of a single genus that occurs in south-central United States; the latter is composed of two genera that occur in northeastern Asia. Members of both families inhabit subterranean freshwaters and are of stygobiont facies. Although the phylogenetic relationship of the allocrangonyctids is unclear, they are ap- parently allied with the genus Pseudoniphargus and may be aberrant hadzioids. The pseudocrangonyctids are allied with the Holarctic family Crangonyctidae and are assigned to the superfamily Crangonyctoidea.
The North American amphipod genus A/- locrangonyx Schellenberg, and the east Asian genera Pseudocrangonyx Akatsuka & Ko- mai and Procrangonyx Schellenberg were originally assigned to the Crangonyx group of the old family Gammaridae (s.l.) by Schellenberg (1936). In recent years, how- ever, the Gammaridae have been split into a number of separate families, and many of the genera placed in the Crangonyx group by Schellenberg have been assigned to dif- ferent families. Some of these genera are now placed in the family Crangonyctidae (see Holsinger 1977, 1986a, b), with which Allocrangonyx, Pseudocrangonyx and Pro- crangonyx have sometimes been associated (see Bousfield 1983, Holsinger 1986a, b). Although these genera may be somewhat “crangonyctid-like” in overall similarity, they possess unique character state combi- nations that preclude their membership in this family (see Holsinger 1986a, b). Both Allocrangonyx and Pseudocrangonyx/Pro- crangonyx have been referred to informally as separate family groups by several workers (Bousfield 1977, 1978, 1982; Holsinger 1977, 1986b), but heretofore neither group
has been given formal family status. In this paper, I will show that each group represents a distinct family of gammaridean amphi- pods. Their phylogenetic, zoogeographic and super-familial relationships will also be ex- amined.
Allocrangonyctidae, new family
Type genus (and only known genus).—Al- locrangonyx Schellenberg, 1936.
Diagnosis. —Without eyes or pigment, of stygobiont facies. Body smooth except for few dorsal spines on uronites. Sexually ma- ture male larger than female, with sexually dimorphic uropod 3. Interantennal lobe of head rounded anteriorly, inferior antennal sinus shallow. Antenna 1 longer than 2, ac- cessory flagellum 2-segmented. Antennae lacking calceoli. Mandibles well developed; left lacinia mobilis 4-dentate; molar tritur- ative; palp 3-segmented. Lower lip with thick inner lobes. Inner plate of maxilla 1 with one apical seta, outer plate with eight or nine unmodified (non-serrate/pectinate) apical spines. Apical margin of outer plate of max- illa 2 uneven (weakly bilobed), with two dis-
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tinct sets of unequal setae; inner plate with- out oblique row of facial setae. Inner plate of maxilliped short, much smaller than out- er; inner margin of outer plate with row of bladelike spines.
Coxae 1-4 rather shallow, posterior mar- gin of 4 weakly excavate. Propods of gnathopods powerful, subchelate, 2nd near- ly twice size of Ist; palms bearing double row of tiny spines, many distally notched. Carpus (segment 5) of gnathopod 1 subequal in length to propod, that of gnathopod 2 proportionately much shorter. Pereopods 5-7 increasing in length posteriorly but oth- erwise generally similar; dactyls of 6 and 7 with several sets of anterior and posterior marginal spines. Coxal gill of gnathopod 2 large and bilobed; coxal gills of pereopods 3-6 ovate or subovate, with very short (ru- dimentary) stalks. Sternal gills absent. Brood plates sublinear. Distoposterior corners of pleonal plates not acuminate. Pleopods sub- equally biramous. Uronite 3 without ec- dysial spine on ventral margin. Margins and apices of uropods 1 and 2 with spines, pe- duncle of 1 with basofacial spine. Uropod 3 elongate, biramous (parviramous); inner ramus greatly reduced (scale-like); outer ra- mus well developed, bearing tiny 2nd seg- ment, becoming greatly elongate and sec- ondarily segmented in larger males. Telson short, with shallow apical notch, apical lobes with spines.
Relationship. —In comparison with a number of potentially related outgroups of gammaridean amphipods, including the Crangonyctidae, Hadziidae, Niphargidae, Pseudocrangonyctidae (new family de- scribed below), and Pseudoniphargus (prob- able family group, but not formally named to date), the Allocrangonyctidae possess at least five, presumably autapomorphic, character states that, in full combination, make them unique: (1) large, bilobed coxal gill on gnathopod 2; (2) non-serrate (or non- pectinate) spines on apex of outer plate of maxilla 1; (3) stalks of coxal gills vestigial; (4) posterior marginal spines on dactyls of pereopods 6 and 7; and (5) positive allo-
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metric growth in combination with second- ary segmentation of outer ramus of male uropod 3.
Although the allocrangonyctids share a number of important characters with other genera, such as Pseudoniphargus, Niphar- gus, and members of the Crangonyctidae (see Holsinger 1971, Bousfield 1977, Bar- nard & Karaman 1980, Notenboom 1988), their phylogenetic relationship has never been clearly demonstrated. Allocrangonyx shares at least 14 apomorphic characters with the peri-Mediterranean, amphi-Atlan- tic subterranean genus Pseudoniphargus Chevreux (and its satellite genus Parapseu- doniphargus Notenboom) (Table 1). At least seven of these characters are also shared with the western Palearctic subterranean ge- nus Niphargus (characters 1, 3, 4, 6, 7, 10, 13) but there are also a number of important differences between this genus and A/lo- crangonyx (see Holsinger 1971, Noten- boom 1988). In addition, five of these char- acters are shared with genera in the Holarctic family Crangonyctidae (characters 1, 8, 11, 13, 14), but character 8 of the crangonyctids may be different (i.e., the palmar spines are proportionately much larger), and character 11 is variable among species in several gen- era. Based on the significantly greater num- ber of apomorphic characters shared by A/- locrangonyx and Pseudoniphargus, the allocrangonyctids are obviously more closely related phylogenetically to the latter than they are to either Niphargus or the cran- gonyctid genera.
Allocrangonyx Schellenberg
Allocrangonyx Schellenberg, 1936:33 (type species by original designation, Niphar- gus pellucidus Mackin, 1935).—Holsin- ger, 1971:318-319.—Barnard & Barnard, 1983:447-448.
Remarks. — Many of the important char- acters of the genus are clearly stated in the literature (see above) and need not be re- peated here. Some omissions and mistakes
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in earlier descriptions should be pointed out, however. Segment 3 of the mandibular palp is heavily setose and bears A, B, C, D and E setae. Carpus of gnathopod | is relatively long, approximately as long as the propod; merus bears a small, semihyaline posterior lobe. Carpus of gnathopod 2 is short, less than 2 the length of propod, and bears a distinct (narrow) posterior lobe. A majority of palmar margin spines on the gnathopods in the outer row are distally notched and not simply “‘spinate”’ or setule tipped as in- dicated by Holsinger (1971). Coxal gills (on pereopods 3-6) have vestigial peduncles or stalks. Coxal gill 1 shown by Barnard & Bar- nard (1983:211, fig 91) is mislabelled a ster- nal gill. The small basofacial spine on pe- duncle of uropod 1 was inadvertently omitted on plate 107 (fig. 4m) in Holsinger (1971) and also in the diagnosis by Barnard & Barnard (1983:448). Uropod 3 of the male shown in Holsinger (1986a:540, fig. 1) is drawn too short; it should be about twice as long as indicated.
At present the genus is composed of two troglobitic species from south-central United States, the geographic distributions of which are shown on a number of range maps (see Holsinger 1971, Barnard & Barnard 1983, Holsinger 1986a, b). The distributions shown in Holsinger (1986b, fig. 6) encom- pass all known localities recorded to date, including those given below.
Allocrangonyx pellucidus (Mackin) Fig. 1
Allocrangonyx pellucidus (Mackin).—Hol- singer, 1971:320—322 (with references). — Black, 1971:7.—Holsinger, 1972:77, fig. 32b.— Black, 1973:15.—Reisen, 1975:28, 30.—Pennak, 1978:460, fig. 317H, K.— Barnard & Barnard, 1983:447-448, fig. 18D—Fitzpatrick, 1983:151.—Holsin- ger, 1986a:540, 542, fig. 1; 1986b:97.
Range.—Caves and springs of the Ar- buckle Mountains in south-central Okla- homa (Murray and Pontotoc counties).
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Table 1.—List of 14 apomorphic character states shared by Allocrangonyx and Pseudoniphargus/Par- apseudoniphargus. Outgroups used to determine char- acter polarity include: Crangonyctidae, Gammaridae, Hadziidae, Niphargidae and Pseudocrangonyctidae.
. Accessory flagellum of first antennae 2-segmented.
. Lacinia mobilis of left mandible 4-dentate.
. Lower lip with thick inner lobes.
. Apical setae of inner plate of maxilla 1 reduced in number (typically less than 3) and often non-plu- mose, or only weakly so.
5. Apical margin of outer plate of maxilla 2 uneven and bearing two distinct groups of setae.
6. Inner plate of maxilla 2 without oblique row of facial setae.
7. Inner plate of maxilliped reduced in size relative to outer plate.
8. Distally notched spines on palms of gnathopod propods.
9. Merus of gnathopod 1 with posterior lobe (some- times semihyaline, sometimes pubescent).
10. Absence of sternal gills.
11. Brood plates narrowly sublinear.
12. Peduncle of uropod 1 with basofacial spine.
13. Inner ramus of uropod 3 reduced to scalelike plate.
14. Telson relatively short and not deeply notched or
cleft (lobes nearly completely fused).
hWN
New locality record (since Holsinger 197 1) based on material in author’s collection. — Oklahoma, Murray County: spring on Hon- ey Creek near Davis (W. K. Reisen, collec- tor).
Allocrangonyx hubrichti Holsinger Figs. 2,.3
Allocrangonyx hubrichti Holsinger, 1971: 324-326, pls. 107-109 (with refer- ences).—Holsinger, 1972:77-78, fig. 32a.—Pflieger, 1974:36.—Craig, 1975:4; 1977:83.—Nordstrom et al., 1977:8.— Barnard & Barnard, 1983:447-448, fig. 9I.—Fitzpatrick, 1983:151.— Wilson, 1984:26.—Gardner, 1986:17-18.—Hol- singer, 1986a:542; 1986b:97.
Range. — Caves and spring(s) of the Ozark Plateau in east-central Missouri (Phelps, Pulaski and Washington counties).
New locality records (since Holsinger
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Fig. 1. Allocrangonyx pellucidus (Mackin). Male (21.75 mm), seep, 0.5 km NW of Turner Falls, Murray County, Oklahoma: A, Left mandible; B, Dentate part of right mandible; C, Maxilla 1; D, Maxilla 2. Max- illae drawn to larger scale than mandibles. Female (18.00 mm) from same locality: E, Pleopod 1 (in part), F, Uropod 1.
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Fig. 2. Allocrangonyx hubrichti Holsinger. Female (17.00 mm), Saltpeter Cave, Phelps County, Missouri: A, Gnathopod | (palm enlarged); B, Pereopod 4 (dactyl enlarged). Gnathopod and pereopod drawn to same scale.
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Fig. 3. Male specimen (16.0 mm) of Allocrangonyx hubrichti Holsinger from Kaintuck Hollow Natural Bridge, Phelps County, Missouri. Elongate 3rd uropods indicated by white arrows. Note that last three segments of pereopods 6 and 7 are broken off.
1971); based on material in author’s collec- tion). — Missouri, Phelps County: Kaintuck Hollow Natural Bridge (in stream pool), 18 km SW of Rolla (J. E. Gardner, coll.).— Meramec Spring, 8 km SE of St. James (J. E. Gardner, coll.). Pulaski County: Killam Cave (in pool), 14 km S of Waynesville (J. E. Gardner, coll.). Washington County: Hamilton Springs Cave (in pool), 12 km SE of Sullivan (J. L. Craig and T. Cravens, coll.).— Mossy Spring Cave (in stream), 16 km E of Richwoods (J. E. Gardner, coll.). Remarks. —The remarkable develop- mental changes in the third uropod of both species of Allocrangonyx were discussed at some length by Holsinger (1971). During growth, the second segment of the outer ra- mus decreases in proportion to an increase in size of the first segment in both sexes. In males, the first segment of the outer ramus
increases allometrically in relation to both the length of the peduncle and the body and, in concert with increase in size, the outer ramus differentiates into secondary seg- ments. This unusual secondary sexual di- morphism is even more pronounced in A. hubrichti. For example, as reported earlier (Holsinger 1971, 1972) uropod 3 was 45% as long as the body, with 9 secondary seg- ments, in a 15.0 mm-long male, and 65% as long as the body, with 16 secondary seg- ments, in a 18.0 mm-long male. In a more recently collected male specimen, 16.0 mm in length, uropod 3 was as long (or slightly longer) than the body, with 30 secondary segments (see Fig. 3).
Elongation of uropod 3 in larger males of many species Pseudoniphargus has also been reported (Stock 1980). But in this genus, the peduncle may also become elongate and the
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outer ramus neither differentiates into sec- ondary segments nor reaches the extraor- dinary length seen in A. hubrichti (Stock 1980, Notenboom 1986, 1988). Aithough there are no observations on the reproduc- tive behavior of these organisms, the pos- sibility that the hyperextended third uropod of the male of Allocrangonyx is utilized either in sex recognition or manipulation of the female during copulation, or even ago- nistic behavior between males, should be investigated.
Pseudocrangonyctidae, new family
Type genus.—Pseudocrangonyx Akatsu- ka & Komai, 1922.
Diagnosis.—Typically without eyes (ex- cept one species) and pigment, of stygobiont facies. Body generally smooth, except last seven body segments bearing dorsal setae and uronite 2 bearing few small dorsal spines. Sexually mature females larger than males. Interantennal lobe rounded ante- riorly, inferior antennal sinus shallow. An- tennae | longer than 2, accessory flagellum 2-segmented. Antennae without calceoli. Apical margin of upper lip rounded, un- notched. Mandibles well developed; molar weakly triturative, bearing single seta or not; left lacinia mobilis 5-dentate; segment 3 of palp equal in length to segment 2, with A (sometimes), D and E setae. Inner lobes of lower lip small or vestigial. Inner plate of maxilla 1 with apical plumose setae, outer plate with typically seven serrate (or pectin- ate) spines. Inner plate of maxilla 2 with oblique row of facial setae. Inner margin of outer plate of maxilliped with setae and few small spines, but lacking bladespines.
Coxae shallow, barely touching or typi- cally discontiguous; posterior margin of 4 without excavation. Propods of gnathopods relatively large (crangonyctid-like), subche- late; propod of 1 a little larger than 2; palms rather long, oblique, armed with double row of distally notched spines (possibly vari- able). Carpus of gnathopod 1 short, with
953
small posterior lobe; that of 2 longer, with- out lobe. Pereopods 3 and 4 normal, sub- equal in length. Pereopods 5-7 increasing in overall length posteriorly; bases with small distoposterior lobes. Stalked coxal gills on gnathopod 2 and pereopods 3-6. Single me- dian sternal gills on pereonites 2-4 or 2-5. Brood plates small, sublinear. Distoposter- ior corners of pleonal plates not acuminate. Pleopods normal, subequally biramous; pe- duncles with 2 coupling spines each. Uro- pods 1 and 2 biramous, with marginal and apical spines; peduncle of 1 with basofacial spine(s). Uropod 3 uniramous; ramus elon- gate, bearing spines and few setae, 2nd seg- ment present and short, or absent. Telson longer than broad; apical margin with notch of variable depth but typically rather shal- low (not deeper the '2 length of telson); api- cal lobes with few spines.
Remarks. — At present this family is com- posed of two northeast Asian genera, Pseu- docrangonyx and Procrangonyx. Their geo- graphic distribution is shown on maps in Barnard & Barnard (1983) and Holsinger (1986b). Unfortunately, previous descrip- tions of taxa assigned to this family have been very uneven. Thus important taxo- nomic details are available for some species (e.g., Pseudocrangonyx asiaticus and P. co- reanus) but are lacking or unclear for others (e.g., Procrangonyx and other species of Pseudocrangonyx). It may therefore be nec- essary to amend the family diagnosis given above as these missing taxonomic details become available.
Relationship. — Although the Pseudo- crangonyctidae are allied with the Holarctic family Crangonyctidae as indicated below, they differ from this group in a number of important characters and warrant recogni- tion as a distinct family: (1) segment 3 of mandibular palp equal in length to segment 2; (2) molar of mandible weakly triturative (or perhaps not triturative in some species); (3) gnathopods and pereopods tending to be more setose, especially segment 2 (basis) of the gnathopods and pereopods 3 and 4, and
954
the coxae of pereopods 5-7; (4) coxae gen- erally discontiguous (a character shared with many members of the family Bogidiellidae); (5) abdominal segments (pleonites and uronites) and 7th pereonite with clusters of setae dorsodistally; (6) uronites with small spines (Ist with ecdysial spine on ventral margin, 2nd with few dorsodistal spines, 3rd with several spines ventrodistally near base of peduncle of uropod 3); and (7) uropod 3 uniramous and elongate, with ramus 3 to 6 times length of peduncle.
A relatively close phylogenetic relation- ship between the families Pseudocrango- nyctidae and Crangonyctidae is indicated by similarity of the following characters, most of which are apparently synapomor- phies: (1) 2-segmented accessory flagellum; (2) structure of mouthparts, except that mandibular palp segment 3 is proportion- ately a little longer and the molar is not as strongly developed in the Pseudocrango- nyctidae (see above); (3) similar shape of, and proportionately large, gnathopod pro- pods (in combination with short carpi); (4) palms of gnathopod propods with double row of thick, distally notched spine teeth (although possibly variable in Pseudocran- gonyctidae); (5) rastellate setae on carpus of one or both gnathopods; (6) median sternal gills; (7) loss of inner ramus of uropod 3 (cf. Stygobromus and Synurella); and (8) rela- tively short telson with shallow apical notch (variable).
Pseudocrangonyx Akatsuka & Komai
Pseudocrangonyx Akatsuka & Komai, 1922: 120 (type species not designated there- in).—Uéno, 1966:504—505 (with refer- ences).— Barnard & Barnard, 1983:442- 443 (type species Pseudocrangonyx shi- kokunis Akatsuka & Komai, designated therein).
Remarks.—Nine species are included in this genus at present; detailed range maps are found in Birstein (1955), Uéno (1966) and Barnard & Barnard (1983). They in-
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habit subterranean waters (e.g., caves, springs, wells) in northeastern China, east- ern Siberia (including the Kamchatka Pen- insula), Korea and the Japanese Islands (see Holsinger 1986a). Further details on one of these species are given below.
Pseudocrangonyx asiaticus Uéno Fig. 4
Pseudocrangonyx asiaticus Uéno.— Ueno, 1966:506—-518, figs. 2-8 (with ref- erences); 1971:198.— Barnard & Barnard, 1983:443, figs. 8B, 9G, 11B, 18C, 20A.— Holsinger, 1986a:542, fig. 4.
Material examined. —South Korea: Gosu- gul (cave), 2 males collected by B. A. Lee (in Zodlogisch Museum of Amsterdam col- lection); Simbog-gul (cave) (location in Uéno 1966:502—503), 12 females, 6 males, | juv. collected by K. S. Lee (in author’s collec- tion).
Range. —Subterranean waters in Korea, northeastern China and the Tsushima Is- lands of Japan.
Remarks. — Although Uéno’s (1966) re- description of this species (and also the de- scription of P. coreanus in the same paper) was very thorough, my recent examination of the above material revealed some taxo- nomic details that were either omitted or should be further emphasized as follows.
Segment 3 of mandibular palp equal in length to segment 2, bearing several short A setae on outer margin, row of short D setae on distal half of inner margin, and 7 to 8 longish E setae of unequal length on apex. Mandible: molar weakly triturative, bearing | seta; left lacinia mobilis 5-dentate. Dactyls of gnathopods with row of blade- like processes (spines?) on inner margin; un- gues relatively long. Propod of gnathopod 1: palm with uneven double row of distally notched spine teeth and row of long setae on outside; medial setae present, in sets of 2s and 4s. Propod of gnathopod 2: palm with double row of 5 distally notched spine teeth; defining angle with 2 spine teeth on
VOLUME 102, NUMBER 4 955
Fig. 4. Pseudocrangonyx asiaticus Uéno. Female (10.2 mm), Simbog-gul (cave), South Korea: A, Gnathopod 1 (rastellate setae and palmar margin enlarged); B, Gnathopod 2 (dactyl, palmar spines and rastellate setae enlarged). Male (6.9 mm) from same locality: C, Urosome (uronites 1, 2, 3) (Gnathopods and urosome drawn to same scale.)
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outside; inferior medial setae in sets of mostly 2s, superior medial setae in sets of 4s. Segment 5 of gnathopods bearing 2 ras- tellate setae on posterior margin at distal end. Pereonites 5—7 with sternal blisters (small ventral humps) (cf., Sternophysinx from South Africa and species of the hubbsi group of Stygobromus from western United States). Upper half of posterior margins of pleonites (of larger specimens) with short row of fine setae. Pereonite 7 and abdominal segments each with cluster of 4 to 8 fine setae dorsodistally. Uronite 1 with ecdysial spine(s) on ventral margin, uronite 2 with 2 short spines dorsodistally, uronite 3 with few spines ventrodistally near base of pe- duncle of uropod 3. Uropod 2 of male sex- ually dimorphic: inner ramus with 2 or 3 distally serrate, apical spines in cluster with several unmodified spines (sexually mature females lack modified apical spines).
Procrangonyx Schellenberg
Procrangonyx Schellenberg, 1934:217 (type species by monotypy, Eucrangonyx ja- ponicus Uéno, 1930).—Barnard & Bar- nard, 1983:444445.
Eocrangonyx Schellenberg, 1936:37 (objec- tive junior synonym).
Remarks.—Procrangonyx is based on a single species, P. japonicus, which was de- scribed by Uéno (1930) from two male spec- imens taken from a subterranean stream in the suburbs of Tokyo (see also Holsinger 1977, 1986a). Except for the absence of a 2nd segment on the ramus of uropod 3, this genus appears to be closely allied with Pseu- docrangonyx. However, Uéno’s original de- scription was incomplete, and several im- portant taxonomic details were omitted or are unclear. For example, it cannot be de- termined from the description whether the palmar margin spines of the gnathopod pro- pods are distally notched or not.
Although Uéno (1930) stated that the type specimens were deposited in the collection of the Otsu Hydrobiological Station, my at-
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tempts to borrow this material or further study have been unsuccessful. Unfortu- nately, there is a good possibility that the type specimens no longer exist.
Discussion
Both mosaic evolution and convergence are common phenomena in the Amphipoda and have resulted in many taxa that are difficult to interpret phylogenetically or to classify with any degree of certainty. Nu- merous taxa display mosaic patterns, re- sulting from differential rates of evolution and reflected in curious mixtures of plesio- morphic and apomorphic characters. In many instances a single taxonomically im- portant structure, such as the gnathopod or uropod 3, may display a combination of both primitive and advanced character states. Convergence has led to the evolution of homoplastic structures in taxa that oth- erwise are apparently only distantly related. Frequently, these homoplasies involve lost or reduced structures (e.g., one or both rami of uropod 3) that are often difficult to in- terpret cladistically. For these reasons there has been considerable confusion regarding the phylogenetic relationship of the allo- crangonyctids with other families of gam- maridean amphipods, as well as their place- ment in a superfamily that most clearly reflects their taxonomic affinities. It is doubtful if morphology alone can ever pro- vide a wholly satisfactory solution to this problem.
Because of their morphological similarity and potential relationship to Niphargus, Al- locrangonyx and Pseudoniphargus were originally aligned in a single, unnamed fam- ily group and assigned to the superfamily Niphargoidea by Bousfield (1977, 1978). Barnard & Karaman (1980:13), however, suggested that these two genera “only have in common a few coincidental characters” and therefore should not be assigned to the same family group. They also strongly ad- vocated abandoning the superfamily Ni-
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phargoidea and suggested placing both Pseudoniphargus and the niphargids in the superfamily Hadzioidea. Subsequently, Bousfield (1982) reassigned A/locrangonyx to the superfamily Crangonyctoidea and also suggested that Pseudoniphargus is more closely allied to the superfamily Melitoidea (=Hadzioidea). In their treatise ““Freshwa- ter Amphipoda of the World,” Barnard & Barnard (1983) referred to the “allocran- gonyctids”’ as a member of their “‘Sterno- branchiate Groups (Crangonyctoids),” but they did not propose any formal taxonomic designation.
Despite significant differences in geo- graphic distribution and ecology, which are pointed out below, the allocrangonyctids are probably more closely related phylogenet- ically to the pseudoniphargids than to any other group of amphipods. Although sev- eral workers, including Stock (1980), Bar- nard & Karaman (1980), and Notenboom (1988), attribute much of the similarity be- tween these two groups to convergence (ho- moplasy) or as being overvalued, I believe that the high number of apomorphic char- acters they share suggests otherwise. It is unlikely that so many detailed similarities in the mouthparts and gnathopods of these genera would have resulted from conver- gence.
The geographic distribution of Al/locran- gonyx is restricted to subterranean fresh- waters in the central interior of North America and is far removed from coastal areas at present. Its range does not extend into areas exposed to marine embayments in the Tertiary or even the Cretaceous, but a part of it in southern Oklahoma would have been less than 100 kilometers from marine embayments in the Late Cretaceous (Holsinger 1971). This distribution pattern suggests that A//ocrangonyx represents a rel- ict lineage, long removed from marine ancestors. In contrast, species of Pseudo- niphargus occupy a wide range of marine to fresh water, subterranean habitats in the cir- cum-Mediterranean region of southern Eu-
957
rope and North Africa and on several is- lands in the Atlantic, including the Azores, Madeira and Bermuda (Stock 1980; Stock et al. 1986; Notenboom 1986, 1987a, b, 1988). These species occur at present in coastal areas or in areas that were directly exposed to marine embayments in the Ter- tiary.
Neither the difference in geographic dis- tribution between A//ocrangonyx and Pseu- doniphargus, nor the fact that some species of the latter live in brackish (or even marine) water, rule out the origin of these two groups from a common ancestor, however. The many synapomorphies between these groups indicate a common ancestry. On this basis, I suggest that the allocrangonyctid and pseu- doniphargid lineages could have been de- rived from a widespread ancestor that in- habited the old Tethyan seaway in Mesozoic times. At that time the areas presently oc- cupied by these groups would have been much closer geographically. Subsequent continental movements combined with widening of the Atlantic, regression of shal- low inland seas, and various other geolog- ical changes would have severely isolated these groups from each other. Divergence during the long period of geographic isola- tion that followed has produced some major morphological differences, but enough im- portant similarities remain to support an obvious phylogenetic relationship.
Notenboom (1988) has recently pointed out a number of important similarities, pre- sumably synapomorphies, between Pseu- doniphargus and the monotypic genus A/- lomelita, which occurs in brackish waters and sometimes in interstitial habitats along the coast of Europe from Norway to Por- tugal. Allomelita, in turn, is closely allied with Melita and is thus a bona fide member of the superfamily Hadzioidea as presently understood. The cladistic relationship be- tween Allocrangonyx and Pseudoniphargus- is certainly as strong as that of the latter with A/lomelita, suggesting, ipso facto, that both the allocrangonyctids and pseudoni-
958
phargids may also be members of the Had- zioidea. Although some problems regarding the phylogenetic affinities of Al/ocrangonyx and Pseudoniphargus remain unresolved, there is a good possibility that both are high- ly divergent hadzioids. Their morphological character combinations make it highly im- probable that either is a crangonyctoid or niphargoid as some workers have previ- ously suggested.
The phylogenetic relationship of the fam- ily Pseudocrangonyctidae is less problem- atic. The number of apomorphies shared by this family and the Crangonyctidae suggest a relatively close phylogenetic relationship of these two groups. These two families, in turn, can be placed in the superfamily Cran- gonyctoidea, which by definition also in- cludes the freshwater families Neoniphar- gidae and Paramelitidae of the Southern Hemisphere (see Bousfield 1978, 1982, 1983; Holsinger 1986a, b; Williams & Bar- nard 1988).
Both the pseudocrangonyctids and cran- gonyctids are known only from freshwater habitats in the Northern Hemisphere and lack close morphological affinities with any group of marine amphipods. They are there- fore believed to represent very old groups of freshwater amphipods that originated on the Laurasian paleocontinent prior to the separation of Eurasia and North America (Holsinger 1986a, b). Geographically, the pseudocrangonyctids replace the crango- nyctids in extreme eastern Asia, where the latter are almost entirely absent (see Hol- singer 1986b:fig. 1). The present range of Pseudocrangonyx, which encompasses parts of the northeastern Asian mainland and the Japanese Islands, probably reflects an ear- lier, continuous distribution of this genus in freshwater habitats throughout the region. Separation of the Japanese Islands as slivers from the Asian continent by tectonic activ- ity beginning in the middle Tertiary (see Dott & Batten 1976) would have isolated populations in Japan from those on the mainland. Based on the assumption that
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Pseudocrangonyx is an old subterranean freshwater inhabitant, it is highly unlikely that any of these insular populations were established by recent invasions from marine waters.
Acknowledgments
I am grateful to H. S. Kim and K. S. Lee of Seoul National University of South Ko- rea, J. E. Gardner, formerly with the Mis- sourl Department of Conservation, J. L. Craig, and W. K. Reisen for providing me with some of the specimens used in this study. I thank J. L. Barnard of the Smith- sonian Institution, E. L. Bousfield of the Royal Ontario Museum, and D. E. Peterson of Old Dominion University for comment- ing on earlier drafts of the manuscript.
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Barnard, J. L., & C. M. Barnard. 1983. Freshwater Amphipoda of the World (Parts I & II). Hayfield Associates, Mt. Vernon, Virginia, 830 pp.
, & G. S. Karaman. 1980. Classification of
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Birstein, J. A. 1955. Genus Pseudocrangonyx Akat- suka & Komai (Crustacea, Amphipoda) in SSSR.— Bulletin Moscow Society of Naturalists (Section Biology) 60(5):77—84 (in Russian).
Black, J.H. 1971. The cave life of Oklahoma.—Okla-
homa Underground 4(1—2):2-49.
. 1973. A checklist of the cave fauna of Okla-
homa—corrections in the order Amphipoda. —
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Bousfield, E.L. 1977. A new look at the systematics
of gammaroidean amphipods of the world.—
Crustaceana, supplement 4:282-316.
. 1978. A revised classification and phylogeny
of amphipod crustaceans. — Transactions of the
Royal Society of Canada (4th series) 16:343-
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. 1982. Amphipoda. Pp. 254-293 inS. S. Park,
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. 1983. An updated phyletic classification and
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Craig, J.L. 1975. Achecklist of the invertebrate species recorded from Missouri subterranean habi- tats.— Missouri Speleology 15(2):1—10.
1977. Invertebrate faunas of caves to be in- undated by the Meramec Park Lake in eastern Missouri. — National Speleological Society Bul- letin 39(3):81-89.
Dott, R. H., & R. L. Batten. 1976. Evolution of the earth. McGraw-Hill Book Company, New York, 504 pp.
Fitzpatrick, J. F., Jr. 1983. How to know the fresh- water Crustacea. Wm. C. Brown Company, Du- buque, IA, 227 pp.
Gardner, J. E. 1986. Invertebrate fauna from Mis- souri caves and springs. Natural History Series No. 3, Conservation Commission of the State of Missouri, 72 pp.
Holsinger, J. R. 1971. A new species of the subter- ranean amphipod genus A/l/ocrangonyx (Gam- maridae), with a redescription of the genus and remarks on its zoogeography.—International Journal of Speleology 3:317-331.
. 1972. The freshwater amphipod crustaceans
(Gammaridae) of North America.—Biota of
freshwater ecosystems, Identification Manual U.S.
Environmental Protection Agency 5:1-89.
. 1977. A review of the systematics of the Hol-
arctic amphipod family Crangonyctidae. —
Crustaceana, supplement 4:244—281.
1986a. Holarctic crangonyctid amphipods. Pp. 535-549 in L. Botosaneanu, ed., Stygofauna Mundi. E. J. Brill/Dr. W. Backhuys, Leiden.
1986b. Zoogeographic patterns of North
American subterranean amphipod crustaceans.
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terdam.
Mackin, J.G. 1935. Studies on the Crustacea of Okla- homa, III, subterranean amphipods of the gen- era Niphargus and Boruta. — Transactions of the American Microscopical Society 54(1):41—-51.
Nordstrom, G. R., W. L. Pflieger, K. C. Sadler, & W. H. Lewis. 1977. Rare and endangered species of Missouri. Missouri Department of Conser- vation and U. S. Department of Agriculture Soil Conservation Publication, 129 pp.
Notenboom, J. 1986. The species of the genus Pseu-
doniphargus Chevreux, 1901 (Amphipoda) from
northern Spain.—Bijdragen tot de Dierkunde 56(1):75-122.
. 1987a. Species of the genus Pseudoniphargus
Chevreux, 1901 (Amphipoda) from the Betic
Cordillera of southern Spain. — Bijdragen tot de
Dierkunde 57(1):87-—150.
1987b. Lusitanian species of the amphipod Pseudoniphargus Chevereux, 1901 witha key to all known Iberian species.—Bijdragen tot de Dierkunde 57(2):191-—206.
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1988. Parapseudoniphargus baetis, new ge- nus, new species, a stygobiont amphipod crus- tacean from the Guadalquivir River basin (southern Spain), with phylogenetic implica- tions. — Journal of Crustacean Biology 8(1):110- ‘21.
Pennak,R.W. 1978. Fresh-water invertebrates of the United States. John Wiley & Sons, New York, 803 pp.
Pflieger, W. L. 1974. Fauna of Missouri springs. Pp. 33-42 inJ. D. Vineyard and G. L. Feder, Springs of Missouri. Missouri Geological Survey and Water Resources, Water Resources Report No. 29.
Reisen, W. K. 1975. The ecology of Honey Creek, Oklahoma: Spatial and temporal distributions of the macroinvertebrates. — Proceedings of the Oklahoma Academy of Science 55:25-31.
Schellenberg, A. 1934. Eine neue Amphipoda-Gat-
tung aus einer belgischen Hohle, nebst Bermerk-
ungen uber die Gattung Crangonyx.—Zoolo-
gischer Anzeiger 106:215-218.
1936. Die Amphipodengattungen um Cran- gonyx, ihre Verbreitung und ihre Arten.— Mit- teilungen aus dem Zoologisches Museum in Ber- lin 22:31-44.
Stock, J. H. 1980. Regression model evolution as exemplified by the genus Pseudoniphargus (Am- phipoda).—Bijdragen tot de Dierkunde 50(1): 105-144.
—., J. R. Holsinger, B. Sket, & T. M. Tliffe. 1986. Two new species of Pseudoniphargus (Amphi- poda), in Bermudian groundwaters. — Zoologica Scripta 15(3):237-249.
Uéno, M. 1930. A new subterranean amphipod from
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1966. Results of the speleological survey in South Korea 1966, 2, gammarid Amphipoda found in subterranean waters of South Korea. — Bulletin of the National Science Museum, To- kyo 9(4):501-535.
. 1971. Subterranean Amphipoda from the is-
lands of Tsushima.—Bulletin of the National
Science Museum, Tokyo 14(2):195-199.
Williams, W. D., & J. L. Barnard. 1988. The tax- onomy of crangonyctoid Amphipoda (Crusta- cea) from Australian fresh waters: Foundation studies. Records of the Australian Museum Sup- plement 10:1-180.
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Department of Biological Sciences, Old Dominion University, Norfolk, Virginia 23529-0266.
PROC. BIOL. SOC. WASH. 102(4), 1989, pp. 960-967
NEW GENERA IN THE THALASSINIDEAN FAMILIES CALOCARIDIDAE AND AXIIDAE (CRUSTACEA: DECAPODA)
Brian Kensley
Abstract. — The family Calocarididae Ortmann, 1891, is resurrected and re- diagnosed. Three characters distinguish members of the family: hermaphro- ditism, eye reduction, and second pleopods having enlarged appendices mas- culinae along with loss of the distal endopod. The first pleopods are spatulate and somewhat plow-shaped, a character also seen in some axiids. Four genera are included in the Calocarididae: Calastacus, Calocaris, Callistocaris, and Lophaxius, the latter two being described as new. Two new genera in the Axiidae are described: Posthonocaris and Sakaiocaris. A scheme for the derivation of the Calocarididae from more generalized axiids such as the two newly-described
genera, 1S proposed.
The families and many of the genera of the Thalassinidea are extremely poorly de- fined, with little sense of phylogeny in the currently accepted classification. The Axi- idae in particular contains several more or less well diagnosed and probably polyphy- letic genera. In the course of a phylogenetic study of the thalassinideans, it became clear that a group of four genera of axiids were more closely related to each other than to the rest of the family. Almost 100 years ago, Ortmann (1891), placed one of these genera in a separate family, the Calocaridae, but this taxon received no recognition and has not been used in the last 50 years. This fam- ily is now resurrected and rediagnosed for this group of four probably monophyletic genera.
Calocarididae Ortmann, 1891
Calocaridae Ortmann, 1891:47, 50.—Ste- phensen, 1910:75, 77, 189.—Runnstrom, 1925:14.
Type genus.—Calocaris Bell, 1853.
Diagnosis. —Hermaphroditic, generally deepwater forms, with eyes showing some reduction and loss of both pigment and om- matidial facets. Linea thalassinica absent.
Maxilla 2 scaphognathite bearing spinulose whip. Maxilliped 3 bearing dentate ischial crest. Pereopods lacking exopods. Pereo- pods | and 2 chelate. Pleopodal rami elon- gate-slender. Pleopod 1 always present, of 2 articles, directed anteromesially along posterior thoracic sternites, with distal ar- ticle broadened and flattened, and bearing proximomesial cluster of hooks (fused ap- pendix interna). Pleopod 2 present, showing loss of distal endopod, enlargement of ap- pendix masculina, and eventual fusion of appendix interna with appendix masculina. Uropodal outer ramus with transverse su- ture.
Genera. —Calastacus Faxon, 1893; Cal- listocaris, new genus; Calocaris Bell, 1853;
_Lophaxius, new genus.
Remarks.—Ortmann (1891) designated the new family Calocaridae, characterized by four features: rostrum flat, triangular; pleopodal rami narrow; outer uropodal ra- mus with suture; podobranchs and mas- tigobranchs present on pereopods. Ort- mann further suggested that eventually two subfamilies, the Calocarinae and the Lao- midiinae, would need to be distinguished. Stephensen (1910), ina rather obscure sem1-
VOLUME 102, NUMBER 4
popular handbook, used the name Calocar- idae as a family of the Anomura, though not designating an author. His diagnosis, loose- ly translated, reads: ‘““The large chelae have straight long fingers, but no tubercles. Two pairs of legs have chelae. Suture across outer branches of uropod. (*Footnote— By a mis- fortune this suture was dropped from fig. 51). Only one genus and species with us”’ [presumably meaning ‘in Denmark’]. Fol- lowing this diagnosis, is a short discussion and figure of Calocaris macandreae Bell. The only other use of the name Calocaridae I have been able to trace, is that of Runn- strom (1925). This family name (in corrected form), is now resurrected, and its diagnosis expanded, to include several related genera, but not the genera of the Laomidiidae, which are markedly different.
Calastacus Faxon, 1893
Calastacus Faxon, 1893:194.—Borradaile, 1903:539.—de Man, 1925:8, 116.—Balss, 1957:1580.—de Saint Laurent, 1972:353, 354.
Type species. —By monotypy, Calastacus Sstilirostris Faxon, 1893:194.
Diagnosis. —Hermaphroditic. Carapace: supraocular spine present; post-cervical ca- rina and spines lacking; rostrum at lower level than anterior carapace; rostral margins unarmed (except for supraocular spine); median carina entire; submedian carina lacking; lateral carina present only as short posterior extension of lateral rostral mar- gins. Eye lacking pigment, stalk equal to or shorter than cornea. Antennal acicle a slen- der, elongate spike.
Maxillipeds: exopods on 1-3; epipods on 1-3, with small podobranch on 3; 2 arthro- branchs on 3.
Pereopods: lacking exopods; epipods on 1-4, small podobranchs on 1-3; 2 arthro- branchs on 1-4; pleurobranchs lacking; pro- podi and dactyli of 3—5 cylindrical, not ex- panded; pereopod 1 chelipeds slightly asymmetrical, not sexually dimorphic; pe-
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reopod 2 chelate; pereopods 3-5, dactyli simple.
Pleopods: rami elongate-slender; appen- dix interna present on 2-5; pleopod 1 of 2 articles, distal article broad, plow-shaped, appendix interna represented by small me- sial lobe bearing hooks; pleopod 2 with small appendix interna fused with basal article of appendix masculina; both articles of latter with double row of setae on mesial margin.
Uropod: outer ramus with transverse dentate suture.
Telson: lacking dorsal spines, longer than wide.
Species. —
Calastacus laevis de Saint Laurent, 1972. Bay of Biscay, north-east Atlantic, 950- 1000 m.
Calastacus stilirostris Faxon, 1893. Pacific Mexico, 1208 m.
Callistocaris, new genus
Type species.—By present designation, Calocaris alcocki (McArdle, 1900).
Etymology.—The generic name is de- rived from the Greek “kallistos’’—most beautiful, plus “‘karis’—a shrimp. Gender: feminine.
Diagnosis. —Hermaphroditic. Carapace: strong supraocular spine present; postcer- vical carina and spines lacking; rostrum set slightly lower than anterior carapace; rostral margins armed; median carina entire; sub- median carina entire; lateral carina entire posterior to supraocular spine. Eye lacking pigment, anteriorly flattened, with mesio- distal tubercle; eye and stalk not differen- tiated. Antennal acicle fairly well devel- oped, spike-like, but considerably less than half length of peduncle article 4.
Maxillipeds: exopods and epipods (no podobranchs) on 1-3; 2 arthrobranchs on 3.
Pereopods: exopods lacking; epipods on 1-4, podobranchs lacking; 2 arthrobranchs on 1-4; pleurobranchs lacking; propodi and dactyli of 3—5 not expanded; pereopod 1 chelae symmetrical, lacking gape between
962
fingers; pereopod 2 chelate; dactyli of 3-5 simple. Branchiae simple, lacking pinnae.
Pleopods: rami elongate-slender, appen- dix interna lacking on 3-5, pleopod 1 uni- ramous, biarticulate, distal article bilobed, with small mesial patch of hooks; pleopod 2 appendix masculina with indication of fu- sion of 2 articles, with double row of spines along mesial margin, and with appendix in- terna fused basally.
Uropod: outer ramus with transverse non- dentate suture.
Telson: longer than wide, lacking dorsal spines.
Species.—
Callistocaris aberrans (Bouvier, 1905). Off St. Lucia, Lesser Antilles, 809 m.
Callistocaris alcocki (McArdle, 1900). Bay of Bengal, 992 m.
Callistocaris cf. alcocki (McArdle, 1900). SW Indian Ocean, 1000 m.
Calocaris Bell, 1853
Calocaris Bell, 1853:231.—Ortmann, 1891: 50, pl. 1, fig. 5d-—i.—Borradaile, 1903: 539.—de Man, 1925:7, 114.—Balss, 1957: 1580.—de Saint Laurent, 1972:353, 354. Type species. —By monotypy, Calocaris
macandreae Bell, 1853:233.
Diagnosis. —Hermaphroditic. Carapace: supraocular spine part of lateral rostral spine series; post-cervical carina entire; rostrum at slightly lower level than anterior cara- pace; rostral margins armed; median carina entire; submedian carina absent; lateral ca- rina armed. Eye lacking pigment, stalk and cornea not differentiated; anteriorly flat- tened and contiguous along midline. An- tennal acicle reduced to barely visible scale.
Maxillipeds: exopods and epipods on 1- 3; reduced podobranch on 2 and 3; 2 ar- throbranchs on 3.
Pereopods: lacking exopods; epipods on 1-4, small podobranchs on 1-3; 2 arthro- branchs on 1-4; pleurobranchs lacking; pro- podi and dactyli of 3-5 not expanded; pere- opod 1, chelae subsimilar, symmetrical, not sexually dimorphic, broad gape between fin-
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
gers; pereopod 2 chelate; pereopods 3-5 dactyli simple.
Pleopods: pleopod 1 of 2 articles, distal article expanded, lobed, with patch of hooks on small mesiodistal lobe; pleopod 2, en- dopod lacking distal portion, appendix masculina slender, mesially setose, of single article, appendix interna articulating at its base.
Uropod: outer ramus with transverse non- dentate suture.
Telson: longer than wide, with or lacking two rows of submedian non-articulating dorsal spines.
Species. —
Calocaris barnardi Stebbing, 1914. Off Sal- danha Bay, South Africa, 89-180 m. Off Namibia, 338 m.
Calocaris granulosus Grebenyuk, 1975. Gulf of Alaska.
Calocaris macandreae Bell, 1853. Mediter- ranean; North-east Atlantic, 25—1072 m.
Calocaris sp. Indian ‘form’ of C. macan- dreae (see Alcock, 1901): Bay of Bengal, Arabian Sea.
Calocaris templemani Squires, 1965. Northwest Atlantic, Newfoundland, Gulf of Maine, 260 m.
Lophaxius, new genus
Type species.—By present designation, Lophaxius rathbunae, new species (=Ca- lastacus investigatoris Rathbun, 1904, non Anderson, 1896).
Etymology.—The generic name is de- rived from the Greek ‘lophos,’ a crest or mane, referring to the post-cervical mid- dorsal ridge of the carapace, plus the fre- quently-used ‘axius.’ Gender: masculine.
Diagnosis. —Hermaphroditic. Carapace: supraocular spine (part of lateral rostral se- ries) present; post-cervical carina with ir- regular tubercles present; rostrum at slightly lower level than anterior carapace; rostral margins armed; median carina entire; sub- median carina absent; lateral carina only extending short distance posterior to ros- trum, with one or two spines. Cornea un-
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pigmented, not flattened; stalk subequal to cornea in length. Antennal acicle short. Maxillipeds 1-3 with exopods and epipods; large podobranch on 2 and 3; 2 arthro- branchs on 3.
Pereopods: exopods lacking; epipods on 1-4, with large podobranch on 1-3; 2 ar- throbranchs on 1-4; pleurobranchs lacking; pereopod 1 symmetrical, fingers of chelae gaping basally; pereopod 2 subchelate; dac- tyli of 3-5 simple.
Pleopods: rami elongate-slender; appen- dix interna present on 2-5; pleopod 1 of 2 articles, distal article spatulate, with small mesial patch of hooks; pleopod 2, appendix masculina slender, tapering, setose, with ap- pendix interna articulating at its base.
Uropod: outer ramus with transverse non- dentate suture.
Telson: longer than wide, with dorsal non- articulating spines.
Species. —
Lophaxius investigatoris (Anderson, 1896). Arabian Sea, 1733 m.
Lophaxius rathbunae, new species. North- eastern Pacific, Alaska to California, 549— 1190 m.
Remarks.—Lophaxius resembles Calas- tacus in having non-pigmented eyes, and with the cornea not flattened as in Calo- caris. It differs from typical Calastacus in having spines on the rostrum, pleopod | not as broadly plow-shaped, the appendix mas- culina of pleopod 2 not as modified, the appendix interna free, in having a well de- veloped postcervical carina, and in having the chela of pereopod | with a gap between fingers as in Calocaris.
Lophaxius rathbunae, new species
Calastacus investigatoris Rathbun, 1904: 151, non Anderson, 1896.—Schmitt, 1921:112, fig. 75, non Anderson, 1896.
Diagnosis. —Carapace, palm of first che- la, abdominal somites, uropods, and telson tuberculate. Merus of pereopod 1 with about 10 spines on anterior (upper) margin, eight
963
spines on posterior (lower) margin. Outer uropodal ramus with six spines on outer margin; inner uropodal ramus with two or three spines on outer margin.
Material. —Syntypes, USNM 28316, cl 19.5 mm, Albatross sta 3347, off Cascade Head, Oregon, 631 m.—USNM 28317, cl 19.1 mm, Albatross sta 3210, south of Dan- nakh Islands, Alaska, 884 m.—USNM 28318, ovig. cl 18.4 mm, 17.1 mm, Alba- tross sta 2928, off San Diego, California, 763 m.—USNM 155734, cl 17.1 mm, A/- batross sta 4352, off San Diego, California, 979-1005 m.
Remarks. — The species Calastacus inves- tigatoris Anderson, 1896, from 1733 m in the Arabian Sea is, from the description and figure (Alcock & Anderson 1896:pl. 25, fig. 1) remarkably similar to the north-eastern Pacific species, but differs in having weaker spination on the anterior and posterior mar- gins of the merus of the first chelipeds, and in lacking marginal teeth on the inner uro- podal ramus. No doubt further differences would be apparent, were material of the In- dian Ocean species available. Indeed, Rath- bun (1904:151) mentioned that not all the eastern Pacific specimens agreed with Al- cock’s description; she also noted some variability in the specimens at her disposal. Given the vast distance between the Indian and Pacific Ocean records, and the differ- ences noted, the two species cannot be re- garded as conspecific.
Key to genera of the Calocarididae
1. Post-cervical carina or ridge present
OM, CARAPACE, tise a ek So Bynum 2
— Carapace lacking post-cervical ca- i ATE eu Ce ne ne ee 3
2. Eyes flattened, mesially contiguous ERO Re ee Calocaris
— Eyes rounded, not mesially contig- Lie oe ee Lophaxius
3. Eyes rounded; appendix masculina MIESIAlIY SEUOSE. x. ifs ae ween. Calastacus
— Eyes flattened; appendix masculina mesially spinose ........ Callistocaris
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Axiidae Huxley, 1878 Posthonocaris, new genus
Type species.—By present designation, Axius rudis Rathbun, 1906.
Etymology. —The generic name is a com- bination of the Greek ‘posthon’—one hav- ing a large penis (referring to the large ap- pendix masculina), plus ‘karis’—a shrimp. Gender: feminine.
Diagnosis. —Gonochoristic, but with her- maphroditic forms occurring in the popu- lation. Carapace: cervical groove present; postcervical carina and spines absent; ros- tral margins armed; median carina entire; submedian carina dentate; lateral carina en- tire (apart from supraocular spine). Eyes well pigmented; eyestalk rounded, longer than or subequal to cornea. Antennal acicle a well developed spike.
Maxillipeds: exopods on 1-3; epipod present on | and 2; 3 with epipod plus po- dobranch, two arthrobranchs.
Pereopods: exopods absent; pereopods 1— 3 with epipod plus podobranch; pereopod 4 with epipod only; two arthrobranchs on 1—4; one pleurobranch on 2—4. Pereopod 1 chelae asymmetrical. Pereopods 3-5, dac- tyli simple.
Pleopods: Rami elongate-slender. Pleo- pod 1 in female slender-elongate, of 2 ar- ticles, distal article bearing marginal setae; pleopod 1 of male or hermaphrodite spat- ulate, of 2 articles, distal article bearing proximomesial clump of hooks; pleopod 2 of male or hermaphrodite with distal setose portion of endopod somewhat reduced, ap- pendix masculina and appendix interna ar- ticulating at about midlength of endopod, appendix masculina elongate, setose, reach- ing well beyond apices of endopod and ex- opod; pleopods 3-5 lacking appendices in- ternae.
Uropod: Outer ramus with transverse dentate suture.
Telson: With non-articulating dorsal spines; single posterolateral articulating spine.
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Species. —
Lophaxius longipes (Bouvier, 1905), off Bar- bados, 225 m.
Lophaxius rudis (Rathbun, 1906), off Ha- wall, 73—426 m.
Sakaiocaris, new genus
Type species.—By present designation, Axiopsis brucei Sakai, 1986.
Etymology. —The generic name is a com- bination of ‘sakai,’ for Dr. Katsushi Sakai, plus the Greek ‘karis’—a shrimp. Gender: feminine.
Diagnosis. —Males, females, and her- maphroditic forms occurring in same species. Carapace: cervical groove present; post-cervical carina a low rounded ridge lacking spines or tubercles; rostral margins dentate; median carina dentate; submedian carina dentate; lateral carina dentate. Eye with reduced pigmentation; eyestalk round- ed. Antennal acicle a well developed spike.
Maxillipeds: 1—3 with epipod and exo- pod, 2—3 with reduced podobranch; two ar- throbranchs on 3.
Pereopods: exopods lacking; epipod plus podobranch on 1-3; epipod only on 4; two arthrobranchs on 1—4; one pleurobranch on 2—4. Pereopod 1, chelae asymmetrical. Pe- reopod 2 chelate. Pereopods 3-5, dactyli simple.
Pleopods: Pleopod 1 of male of 2 articles, distal article spatulate, with proximomesial clump of small hooks. Pleopod | in female (and in only hermaphrodites seen), slender, elongate, of 2 articles, distal article bearing marginal setae; pleopod 2 in male with large exopod and endopod, setose appendix mas- culina and appendix interna both articulat- ing proximally on endopod. Pleopods 3-5 with free appendix interna.
Uropod: outer ramus with dentate trans- verse suture.
Telson: with non-articulating dorsal spines; with two articulating postero-lateral spines.
VOLUME 102, NUMBER 4
y) fh [A ' (N | f\ ' V a t l | og \\% Wie? \4 [ : } \ 17 \\ \ 4 Y ) f\ : | \g eee | ) \ \ |
H \ \ / | | . \ |
|] \ \/ | } | NJ by
g1 dt oe 91) d1- -c'2 Si— ot ci2 are er 4 Gy 2 2 Oe Coralaxius Sakaiocaris Posthonocaris Lophaxius Calocaris Calastacus Callistocaris
eyes flattened eyes flattened app. mas. spinose post-cervical carina pl. 3-5 app. int. lost app. int. fused with app. mas. eye pigment lost pl. 2 distal endopod lost hermaphroditic pl. 2 app. mas. enlarged some ¢ forms gonochoristic pl. 10° or ¢ spatulate Fig. 1. Hypothetical scheme for derivation of the Calocarididae from more generalized Axiidae, illustrating
pleopods 1 and 2, and changes in character-states. Endopod of pleopod 2 shaded. (Abbreviations: app. int.— appendix interna; app. mas.—appendix masculina; pl.—pleopod.)
Species. —
Sakaiocaris brucei (Sakai, 1986); off West- ern Australia, in hexactinellid sponges, 296-458 m.
Discussion
Three synapomorphies separate the Cal- ocarididae from the Axiidae (s.].). 1. Invar- iable hermaphroditism. 2. Enlargement of the appendix masculina of pleopod 2, along with loss of the setose distal element of the endopod. 3. Eye reduction and loss of eye pigment.
A possible pathway in the development of hermaphroditism, from purely gono- choristic forms (e.g., Coralaxius), through forms having some hermaphrodites in the population (e.g., Posthonocaris, Sakaiocar- is), to purely hermaphroditic forms (Calo-
caris, Calastacus, Callistocaris, Lophaxius), is illustrated in Fig. 1. Coralaxius Kensley & Gore, 1981, is purely gonochoristic, and possesses sexually dimorphic first pleopods. In the male, the first pleopod is uniramous and biarticulate, the distal article being spatulate and having a clump of mesial hooks. These latter are presumed to come from the appendix interna which has fused with the endopod; the exopod has either been lost or has fused with the endopod. It is postulated that the genera Posthonocaris and Sakaiocaris have hermaphroditic forms in the population. In these, the first pleopod of the female or protandrous hermaphrodite is a slender setose, uniramous, biarticulate structure, while in the males, a spatulate first pleopod very similar to those found in the Calocarididae is seen. The presence of pro-
966
tandrous hermaphrodites in populations of decapod species has been well documented. Policansky (1982) mentions a variety of decapod crustaceans in which protandry oc- curs, including Calocaris macandreae in which the biology, and especially reproduc- tion, has been well examined (see Wolle- baek 1909, Runnstrom 1925, Buchanan 1963). Bauer (1986) reported the presence of primary males, primary females, and pro- tandric hermaphrodites that pass through a male phase, a transitional phase, and then become breeding females, in the hippolytid caridean shrimp Thor manningi. While ten specimens of Sakaiocaris brucei and four specimens of the two species of Posthono- caris form too small a sample on which to state with certainty that protandrous her- maphrodites occur in these populations, the dimensions of the specimens (at least of the former species) would seem to suggest this. The only ovigerous female of S. brucei seen has a carapace length of 21.9 mm; the three hermaphrodites with their female first pleo- pods measure 16.1, 19.6, and 20.0 mm (sug- gesting that these are either in the transi- tional phase or approaching the breeding female phase), while five males range from 13.8—23.3 mm.
Posthonocaris seems to represent a more advanced stage than Sakaiocaris, in this supposed trend towards hermaphroditism. This is seen in the reduction of the setose distal portion of endopod of pleopod 2, an elongation of the proximal non-setose por- tion, and marked enlargement of the ap- pendix masculina. In the fully hermaphro- ditic calocaridids Calastacus and Callistocaris, the final stage of this trend, with complete loss of the distal endopod, and enlargement and specialization of the appendix masculina, with which the appen- dix interna has fused, can be seen.
There would seem to be a correlation be- tween development of hermaphroditism and depth distribution in this group of thalas- sinideans. Coralaxius, with its spatulate first pleopod in the male, is a shallow (1 1-76 m) coral reef inhabitant. Sakaiocaris inhabits hexactinellid sponges in 296-456 m. Per-
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
haps there is a reproductive advantage in having a hermaphroditic phase in this species, given its cryptic habit. Species of Posthonocaris have been recorded in depths of 73-426 m; Calocaris from 89-1072 m; Callistocaris from 809-1000 m; Calastacus from 970-1208 m; Lophaxius from 1733 m.
Loss of eye pigment and corneal facets, along with a breakdown of the distinction between cornea and stalk, can be seen in the four genera of the Calocarididae, suggesting loss of function linked to increased depth distribution. The hypothetical scheme pro- posed in Fig. 1 requires that anteriorly flat- tened eyes arose independently in Calocaris and Callistocaris, which is not unfeasible. Similar flattening of eyes can be seen in sev- eral decapod groups such as cave-dwelling hippolytid shrimps, deepsea bresiliid shrimp, as well as in the stomatopod genus Bathysquilla (R. B. Manning, pers. comm.).
Acknowledgments
Iam very grateful to the following for the loan of material: A. J. Bruce of the Northern Territory Museum of Arts and Sciences, Australia; A. Johnson of the museum of Comparative Zoology, Harvard University; R. Springthorpe of the Australian Museum, Sydney; M. van der Merwe of the South African Museum, Cape Town. I thank F. A. Chace, Jr., A. B. Williams, and R. B. Man- ning, all of the Smithsonian Institution, for their (as always) perceptive comments and improvements to the manuscript.
Literature Cited
Alcock, A. 1901. A descriptive catalogue of the In- dian deep-sea Crustacea Decapoda Macrura and Anomala, in the Indian Museum. Being a re- vised account of the deep-sea species collected by the Royal Indian Marine Survey Ship In- vestigator. Calcutta, Trustees of the Indian Mu- seum. 286 pp, 3 pls.
, & A. R. S. Anderson. 1896. Illustrations of the zoology of the Royal Indian Marine Survey Ship Investigator under the command of Com- mander T. H. Heming, R.N. Crustacea, Part 4, plates 16-27; Calcutta. Office of the Superin- tendent of Government Printing, India.
VOLUME 102, NUMBER 4
Anderson, A. R. S. 1896. Natural history notes from the R.I.M. Survey Steamer ‘Investigator,’ Com- mander C. F. Oldham, R.N., commanding. Se- ries II, No. 21. An account of the deep sea Crus- tacea collected during the season 1894—95.— Journal of the Asiatic Society of Bengal 65(2): 88-106.
Balss, H. 1957. Decapoda.—Dr. H. G. Bronns Klas- sen und Ordnungen des Tierreichs 5(1)7(12): 1505-1672.
Bauer, R.T. 1986. Sex change and life history pattern in the shrimp Thor manningi (Decapoda: Car- idea): A novel case of partial protandric her- maphroditism.— Biological Bulletin 170(1):11- ah.
Bell, T. 1853. A history of the British stalk-eyed crus- tacea. John Van Voorst, London, lxv, 386 pp, 174 figs.
Borradaile, L. A. 1903. On the classification of the Thalassinidea.— Annals and Magazine of Nat- ural History (7)12:534—-551.
Bouvier, E. L. 1905. Sur les Thalassinides recueillis par le Blake dans la mer des Antilles et le golfe du Mexique.— Comptes Rendus Hebdomadaire des Séances de l’Académie des Sciences 141(21): 802-806.
Buchanan, J. B. 1963. The biology of Calocaris ma- candreae (Crustacea: Thalassinidea).— Journal of the Marine Biological Association of the United Kingdom 43(3):729-747.
Faxon, W. 1893. Reports on the dredging operations off the west coast of Central America to the Galapagos, to the west coast of Mexico, and in the Gulf of California, in charge of Alexander Agassiz, carried on by the U.S. Fish Commis- sion Steamer “Albatross,” during 1891, Lieut. Commander Z. L. Tanner, U.S.N., command- ing.— Bulletin of the Museum of Comparative Zoology at Harvard College 24(7):149-220.
Grebenyuk, L. P. 1975. Two new species of the su- perfamily Thalassinidea. —Zoologichesckii Zhurnal 54(2):299-304 [In Russian].
Huxley, T. H. 1878. On the classification and the distribution of the crayfishes.— Proceedings of the Zoological Society of London for the year 1878:752-788.
Kensley, B., & R. H. Gore. 1981. Coralaxius abelei, new genus and new species (Crustacea: Deca- poda: Thalassinidea: Axiidae): A coral-inhab- iting shrimp from the Florida Keys and the west- ern Caribbean Sea.—Proceedings of the Biological Society of Washington 93(4):1277- 1294.
Man, J. G. de. 1925. The Decapoda of the Siboga- Expedition. Part 6. The Axiidae collected by the Siboga-Expedition. —Siboga-Expeditie Mono- graphie 39a5:1-127.
967
McArdle, A. F. 1900. Natural history notes from the Royal Indian Marine Survey Ship ‘Investigator,’ Commander T. H. Heming, R.N., command- ing. Series III, No. 4. Some results of the dredg- ing season 1899-1900.—Annals and Magazine of Natural History (7)6:471-478.
Ortmann, A. 1891. Die Decapoden-Krebse des Strassburger Museums. 3. Die Abtheilungen der Reptantia Boas: Homaridae, Loricata und Thal- assinidea. —Zoologischen Jahrbiichern 6:1-58.
Policansky, D. 1982. Sex change in plants and ani- mals.— Annual Review of Ecology and System- atics 13:471-495.
Rathbun, M. J. 1904. Decapod crustaceans of the northwest coast of North America. — Harriman Alaska Expedition 10:1-—190.
1906. The Brachyura and Macrura of the Hawaiian Islands.—Bulletin of the U.S. Fish Commission for 1903, part 3:829-930. Runnstrém, S. 1925. Beitrag zur Kenntnis einiger
hermaphroditischen dekapoden Crustaceen.— Bergens Museums Skrifter, n.s., 3(2):1—115.
Saint Laurent, M.de. 1972. Un thalassinide nouveau du golfe de Gascogne, Calastacus laevis sp. nov. Remarques sur le genre Calastacus Faxon (Crus- tacea Decapoda Axiidae).—Bulletin du Mu- seum National d’Histoire Naturelle (3)35, Zool- ogie 29:347-356.
Sakai, K. 1986. Axiopsis brucei sp. nov., anew sponge- inhabiting axiid (Crustacea: Decapoda: Thal- assinidea), from north-west Australia.—The Beagle. Occasional Papers of the Northern Ter- ritory Museum of Arts and Sciences 3(1):11—20.
Schmitt, W. L. 1921. The marine decapod crustacea of California.— University of California Publi- cations in Zoology 23:1-359.
Squires, H.J. 1965. Anew species of Calocaris (Crus- tacea: Decapoda, Thalassinidea) from the northwest Atlantic.—Journal of the Fisheries Research Board of Canada 22(1):1-11.
Stebbing, T. R. R. 1914. South African Crustacea (Part 7 of S. A. Crustacea, for the marine in- vestigations in South Africa).—Annals of the South African Museum 15(1):1-55.
Stephensen, K. 1910. Storkrebs. 1. Skjoldkrebs. [Danmarks Fauna 9]. G. E. C. Gads, Copen- hagen, 193 pp., 108 figs.
Wollebaek, A. 1909. Effektiv hermaphroditisme hos en decapod Crustace, Calocaris Macandreae, Bell.—Nyt Magazin for Naturvidenskaberne, Christiania 47(3):25 1-268.
Department of Invertebrate Zoology, Na- tional Museum of Natural History, NHB- 163, Smithsonian Institution, Washington, D.C,20560.
PROC. BIOL. SOC. WASH. 102(4), 1989, pp. 968-972
THREE NEW SPECIES OF COLOMBIAN LACE BUGS OF THE GENERA LEPTODICTYA AND LEPTOPHARSA (HETEROPTERA: TINGIDAE)
Richard C. Froeschner
Abstract. —Three new species of lace bugs from Colombia are described and illustrated by habitus drawings: Leptodictya fuscipes collected on an unidentified species of Poaceae; Leptopharsa madrigali found on Phaseolus species (Faba- ceae); and Leptopharsa reflexa taken on Roupala “glabriflora’’ (Proteaceae).
A small collection of lace bugs repre- senting samplings of environments in the vicinity of crop fields in Colombia was sub- mitted by Dr. Raul Velez-Angel, Univer- sidad Nacional, Medellin, Colombia, for identification. In it were three species new to science. Neither of the two genera in- volved here has had a recent key to aid in identification of the numerous species in- cluded in the Drake & Ruhoff (1965) cata- log: Leptodictya Stal with 52 species and Leptopharsa Stal with 105 species. Con- struction of such keys at this time is im- practical, but as names are needed for mak- ing known more information about these insects, they are described here.
Leptodictya fuscipes, new species Fig. 2
Diagnosis.— This species belongs among those forms of Leptodictya in which the cos- tal area has embrowned veins everywhere except in a broad band adjacent to and along the full length of the discoidal area; veins in this area are milky white and together form a sort of halo around the discoidal areas. L. fuscipes can be distinguished most readily from all other members of the group by the strongly embrowned limiting veins of the discoidal area plus the nearly black femora.
Description. —Length 4.5—5.1 mm. Head, surface of pronotum except apical half of posterior projection, and under surface of
body black, these parts usually coated with a dense white pruinosity. Antenna black- ened except for the slightly paler apex of segment III. Bucculae, except narrow mar- gins, black. Legs deep brown to black. Par- anota and anteromedian cyst mostly milky white, outermost vein (along fold) of par- anotum and mediodorsal vein of cyst light brown. Longitudinal pronotal carinae light brown. Hemelytral cells hyaline; veins de- limiting discoidal area, of costal area (except for the broad band along discoidal area), and of most of sutural area fuscous to black; discoidal area with a faintly but distinctly embrowned diagonal line across midlength. Sternal laminae yellow.
Head deflexed, with 5 elongate pale spines directed forward or slightly obliquely up- ward: occipital pair; pair above base of clyp- eus; and one spine on midline of dorsum of head. Antennal segment I about as long as head, twice as long as II; III 4.5 times as long as I + II; IV slightly less than half as long as III. Labium attaining posterior end of mesosternal laminae.
Pronotum with anteromedian cyst tec- tate, as high as median carina, anteriorly produced medially as a right angle attaining medlength of eyes, posteriorly extending al- most to apex of interhumeral convexity. Lateral longitudinal carinae very low, with a single row of punctiform cells; median ca- rina twice as high as lateral carinae, unise- riate.
VOLUME 102, NUMBER 4
Figs. 1-3.
969
1, Leptopharsa reflexa new species, actual length 3.5 mm. 2, Leptodictya fuscipes new species,
actual length 4.5 mm. 3, Leptopharsa madrigali new species, actual length 3.8 mm.
Discoidal area confined to basal half of hemelytron, with 4—5 cells across greatest width. Costal area with 4 usually promi- nent, regular, nearly straight cross veins be- tween which are numerous irregular cells; cells in milky area along discoidal area dis- tinctly reduced. Hypocostal laminae uni- seriate.
Peritreme obliquely transversely auricu- late, reaching hypocostal lamina. Sternal laminae absent from prosternum, present and subparallel on mesosternum, present and forming a cordate outline on metaster- num. Abdomen convex, impunctate.
Holotype male, Colombia; “‘Cocorna (Ant. [ioquia]), Agt. 1981, A. Madrigal, en Graminea,” deposited in the National Mu- seum of Natural History. Paratypes (depos-
ited in Universidad Nacional, Medellin, Colombia, and National Museum of Nat- ural History): 1 male, 2 females with same data as holotype.
The species name directs attention to the dark fuscous to black legs.
Leptopharsa madrigali, new species Fig. 3
Diagnosis. —Among those species of Lep- topharsa with divergent hemelytra (apices, at rest, not completely overlapping, see Fig. 3) this species may be recognized by the combination of 2 regular rows of subequal cells in the costa along the basal three-fourths of the discoidal area; the short occipital spines which do not surpass the antennal
970
insertions; and the wholly blackened fem- ora.
Description. —Length 3.4—4.0 mm. Head, pronotal surface (except pale posterior apex), and body ventrally black. Antennal seg- ments I and IV black, II brown, III yellow. Bucculae black except for yellow ventral row of cells. Femora, extreme ends of tibiae, and tarsi black; most of tibia yellow. Pronotal outgrowths (paranota, longitudinal carinae, and anteromedian cyst) mostly yellow with hyaline cells. Discoidal and subcostal areas with most veins black. Costal area, includ- ing veins, pale except for fuscous band ra- diating posterolaterally from darkened base of membranal area. Sternal laminae yellow.
Head vertically deflexed, with 5 forward directed short spines: occipital spines de- cumbent, not or just reaching antennal in- sertions; a pair of supraclypeal spines, a sin- gle spine above bases of latter pair. Antennal segment I about as long as width of head across eyes; II almost a third as long as I and twice as long as wide; III about 4 times as long as I + II; IV about two-thirds as long as III. Labium reaching posterior ends of mesosternal laminae.
Pronotum with anteromedian cyst inflat- ed, slightly higher than median carina, ex- tending almost to apex of head, extending less than half way up anterior slope of in- terhumeral convexity. Longitudinal carinae very low, composed of a single row of punc- turelike cells. Paranotum biseriate, outline convex around humerus, thence straight or weakly convex almost to rounded anter- oapical angle.
Hemelytron with costal margin weakly convexly diverging on basal fifth, thence straight and diverging to apical fourth. Dis- coidal area confined to basal two-fifths of hemelytron, with 5 cells across greatest width. Subcostal area with 4—5 cells across greatest width. Costal area biseriate along basal three-fourths of discoidal area, irreg- ularly 3-4 cells wide beyond apex of dis- coidal area. Hypocostal lamina uniseriate.
Peritreme obliquely transversely auricu- late, reaching hypocostal lamina. Sternal
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laminae present on all 3 sterna, low unise- riate, more widely separated on meta- than on pro- or mesosterna; sternal groove not interrupted by transverse carina. Abdomen convex, impunctate.
Holotype, male, Colombia; ““Urrao (Ant. fioquia]), Mar. 1977, A. Madrigal C., en Phaseolus sp.” (Fabaceae), deposited in the United States National Museum of Natural History. Paratypes deposited in the Uni- versidad Nacional, Medellin, Colombia, and the National Museum of Natural History; 2 females with same data as holotype; 3 females, La Estrella (Ant. [ioquia]) Colom- bia, April 1985, A. Madrigal on Fabaceae; 1 male and | female, Coldos (Ant. [ioquia]), Colombia, Nov. 1973, A. Madrigal, on Fa- baceae.
The species name dedicates this lace bug to the Colombian entomologist, A. Mad- rigal C., who obtained this type series and has done much to increase knowledge of Heteroptera in Colombia.
Leptopharsa reflexa, new species Fig. 1
Diagnosis. — Among the species of Lep- topharsa in which the sutural areas overlap at rest (apices not divergent, see Fig. 1), this new species belongs to a group characterized by the following features: 1) 3-5 long ce- phalic spines; 2) low, tectate (in no way swollen) anteromedian cyst separated from the higher median longitudinal pronotal ca- rina by a deep angulation; 3) a narrow (1— 2 rows of cells) paranotum continued at same width around humerus and then incurved and abruptly terminated to form a distinct angle with the posterior pronotal projection; and 4) a biseriate costal area.
The 7 species in the group diagnosed with the above enumeration of characters can be separated by the following couplets:
1. Pronotum with prominent, project- ing anterolateral angles, its anterior margin as wide or wider than head across eyes
VOLUME 102, NUMBER 4
— Pronotum without distinct antero- lateral angles, its anterior margin narrower than width of head across eyes
2. Height of longitudinal pronotal ca- rinae over crest of inter-humeral convexity greater than diameter of FORME 6 kd, tenuis (Champion)
— Height of longitudinal pronotal ca- rinae over crest of interhumeral convexity less than diameter of a fe- mur
3. Occipital spines decurved, decum- bent, cylindrical, apices blunt, not surpassing antennal insertions ....
oc Oe er albella Drake
— Occipital spines long, tapering to acute apices, projecting obliquely away from head and extended be- yond antennal insertions
4. Head with 5 spines: a pair of short spines immediately below the me- dian supraclypeal spine plus a pair of occipital spines
— Head with 3 spines: a median su- praclypeal spine and a pair of oc- €ipital spines ............. avia Drake
5. Antennal segment I short, about twice as long as length of eye. Dorsal aspect darker: head, pronotal sur- face, and generally veins of discoidal area lightly but distinctly em- maomoned: =. 1s. Laisa) Jack. fica Drake
— Antennal segment I long, about 2 and a half times as long as length of eye. Dorsal aspect whitish to faintly yellowed (a few veins sometimes @ackened) ............ delicata Monte
6. Paranotum vertically reflexed, free Margin straight from anterior end of prominent humerus
a. he) reflexa, new species
— Paranotum weakly obliquely ele- vated, free margin convex from hu- merus to anterior end.. elegantula Stal
2 © © © © © we we we ew ew
Description. —Length 3.3—3.6 mm. Head, pronotal surface (except anterior margin and
971
most of posterior projection) black, these surfaces may be covered with a light coating of white pruinosity. Antennal segments I and IV black, II and III reddish yellow. Buc- culae mostly white. Legs, except black tarsi, reddish yellow. Pronotum with anterome- dian cyst, longitudinal carinae, and para- nota yellow, cells hyaline to lightly en- fumed. Hemelytral veins mostly yellow, those in discoidal and subcostal areas some- times brown. Sternal laminae pale yellow.
Head vertically deflexed, with 5 cephalic spines: pair of decurved, decumbent occip- itals reaching antennal insertions; supra- clypeal pair and median spine directed an- teriorly. Antennal segment I slightly shorter than length of head, twice as long as II, III nearly 5 times as long asI + II, IV one third as long as III. Labium reaching posterior ends of mesosternal laminae.
Pronotum with anteromedian cyst small, lower than median longitudinal carina, forming a short angulation above base of head, posteriorly terminated between calli, not ascending interhumeral convexity. Lon- gitudinal carinae low, uniseriate, median one slightly higher posteriorly. Paranotum ver- tically reflexed, wholly biseriate or in part uniseriate; free margin straight from ante- rior end to projecting humerus.
Hemelytra with apices overlapping, axes and costal margins parallel. Discoidal area 4—5 cells wide, confined to basal two-fifths of hemelytron. Subcostal area 3-4 cells wide. Costal area biseriate along basal three- fourths, uniseriate on apical fourth. Hypo- costal lamina uniseriate.
Peritreme small, weak, transversely au- riculate, reaching hypocostal lamina. Lam- inae present on all 3 sterna, straight and parallel on pro- and mesosterna, more widely separated and bowed convexly out- ward on metasternum; sternal groove not interrupted by transverse carina. Abdomen convex, impunctate.
Holotype male, Colombia, ““Guarne (Ant. [ioquia]), Jun 1978, A. Madrigal C., en Rou- pala glabriflora”’ (Proteaceae), deposited in the United States National Museum of Nat-
O72
ural History. Paratypes: 4 males, same data as holotype, deposited in Universidad Na- cional, Medellin, Colombia, and in Nation- al Museum of Natural History.
The species name was suggested by the vertically reflexed paranota.
Acknowledgments
The author’s appreciation 1s expressed to Ms. Silver B. West for assistance in pre- paring the manuscript; to Thomas J. Henry and Paul J. Spangler for constructive re-
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
views of the manuscript; and to Young Sohn for the fine habitus drawings.
Literature Cited
Drake, C. J., & F. A. Ruhoff. 1965. Lacebugs of the world: A catalog (Hemiptera: Tingidae).—Bul- letin of the United States National Museum 243: i-vill, 1-634, pls. 1-56.
Department of Entomology, NHB 127, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560.
PROC. BIOL. SOC. WASH. 102(4), 1989, pp. 973-976
KROHNITTELLIDAE AND BATHYBELIDAE, NEW FAMILIES IN THE PHYLUM CHAETOGNATHA; THE REJECTION OF THE FAMILY TOKIOKAISPADELLIDAE AND THE GENERA TOKIOKAISPADELLA, ZAHONYA, AND ABERROSPADELLA
Robert Bieri
Abstract.—New monogeneric families are proposed for the genera Krohnit- tella and Bathybelos (Krohnittellidae and Bathybelidae). The family Tokio- kaispadellidae is considered invalid because the type species is a misidentified Pterosagitta draco. Morphologic changes in poorly preserved chaetognaths are described, and the genera Zahonya and Aberrospadella, based on poorly pre- served specimens, are considered invalid.
After compiling a comprehensive list of approximately 100 species in 22 genera cur- rently included in the phylum Chaetogna- tha, it became clear to me that the genus Kronittella has not been assigned to any ex- tant family and the genus Bathybelos is so exceptional that it cannot be included in any extant family. Therefore, to correct these inconsistencies I propose the following two new families.
Krohnittellidae, new family
Diagnosis. — The family lacks both ante- rior and posterior teeth. A single pair of lateral fins, completely rayed, partly on the trunk and partly on the tail do not reach the ventral ganglion. No ventral transverse musculature.
Discussion. —The family is monogeneric containing the single genus, Krohnittella Germain & Joubin, 1912. The authors based their new genus on two specimens taken in a 4000 m net tow that possibly struck bot- tom in the eastern North Atlantic. Because more than 60 years passed before the genus was reported again, considerable doubt ex- isted about its validity. Tokioka in his 1965 revision retained it as valid, but did not
assign it to any known family. Bieri (1974) rediscovered the genus in the eastern North Pacific where a single, nearly mature spec- imen in good condition was caught with an opening-closing sled trawl at 2000 m in the San Clemente Basin. The lack of ventral transverse musculature and teeth clearly dif- ferentiate the family from the Eukrohniidae and Spadellidae. Salvini-Plawen (1986), un- aware of Bieri’s (1974) paper, suggested that the genus may belong to Bathyspadella. The lack of transverse musculature in Krohni- tella is irreconcilable with Bathyspadella which in any case is the junior genus. AI- though the names are similar, there is no confusion with the family Krohnittidae whose members have a single pair of fla- belliform teeth among other distinguishing characteristics.
Bathybelidae, new family
Diagnosis. —No ventral ganglion but rather a dorsal ganglion. Two pairs of lateral fins completely lack fin rays. One paired row of teeth which are highly curved. Eyes ves- tigial.
Discussion. —The family is monogeneric containing the single genus, Bathybelos. In
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her paper describing the genus, Owre (1973) (=Michel) referred to the dorsal nerve gan- glion as a cerebral ganglion but in her later description (Michel 1984) she referred to it as a rectangular nerve center. I think dorsal ganglion or nerve center is more accurate than cerebral because it is posterior to the apparent location of the vestigial eyes and is more in the position of the corona ciliata. In all known chaetognaths the cerebral gan- glion is well anterior to the eyes (Goto & Yoshida 1987). The second unique feature of this species is the complete lack of rays in the lateral fins. All other known chaeto- gnaths, except the problematic Aberrospa- della discussed below, have some rays in the lateral fins. These characters and the presence of only one paired row of teeth differentiate it from the families Sagittidae, Eukrohniidae, and Spadellidae.
Status of the Family Tokiokaispadellidae
In 1986 Salvini-Plawen erected a new family, Tokiokaispadellidae to contain To- kiokaispadella lata Kassatkina, 1980. The original type genus of the family is incor- rectly given as Tokiospadella, apparently a typographical error (Salvini-Plawen 1986: 126). It is clear that the single specimen which formed the basis of Kassatkina’s new genus and species is nothing more than a poorly preserved specimen of the common species Pterosagitta draco, ubiquitous in the tropical and semitropical oceanic waters of the world.
In Kassatkina’s (1980) description of the new species, taken in a 0-14 m net tow in a lagoon north of New Guinea, the posterior fin reaches only to the transverse septum and is completely rayed as in P. draco. There are two paired rows of teeth as in P. draco. The tooth and hook formula agrees with that given by Alvarino (1967) for P. draco. The collarette is narrow and extends from the neck to the tail fin. In perfectly preserved specimens of P. draco, the collarette extends onto the tail fin and is nearly as broad on
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
each side as the body width. But as E. L. Michael (1919) pointed out long ago, the collarette is usually eroded to varying de- grees. Sometimes it is completely missing. It is very rare to find P. draco with a com- pletely intact collarette and almost never are the “wings” of the collarette preserved (Bieri 1966). In Kassatkina’s specimen the collar- ette is partly stripped away, the usual situ- ation when relatively coarse meshed nets are used or the net is towed faster than about 25 cm per second.
The ovaries are short and immature and there is no indication of seminal vesicles. Her figure shows large gut diverticulae but Alvarino (1967) stated that in P. draco the diverticulae are small and incipient. Kas- satkina’s drawing shows the corona ciliata extending well up onto the head between the eyes, but in the text states that it is en- tirely on the trunk. Aida (1897) and Grassi (1883) show the corona entirely on the trunk whereas Ghirardelli (1952) showed it ex- tending onto the head. Tokioka (1940) showed the corona in three figures. In two of the figures it is on the trunk, but in the largest figure it is partly on the head.
Thus there are no characters which dif- ferentiate this species from P. draco except the ventral transverse musculature, the very short tail segment, and the relatively great body width. None of these characters were listed in Salvini-Plawen’s extremely short diagnosis of the new family.
Tokioka (1952:312) in discussing the transverse musculature of the enigmatic ge- nus Zahonya, wrote, “Itis a noteworthy fact that some species with strongly developed musculature assume an appearance, when they are in a slightly contracted state, as if they were provided with a transverse mus- culature along the whole body.” Nagasawa & Nemoto (1985) discussed the distortion of chaetognaths probably due to attack by bacteria. They documented the shortening of the body length by 50% but did not de- scribe the appearance of pseudo-transverse musculature. My own experience convinces me that if chaetognaths are trawled for more
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than half an hour, some specimens will be seriously damaged and distorted not just by physical abrasion but also by bacterial at- tack as discussed by Nagasawa and Nemoto, and by biochemical lysis after death. If the plankton sample is not preserved immedi- ately upon retrieval from the sea, distorted individuals may compose from a few to all of the plankton sample. Chaetognaths that die before preservation and are left unpre- served for more than a few minutes develop contracted longitudinal muscles that have a series of interuptions that look superficially like transverse muscles as Tokioka report- ed. I have seen this condition in almost every species of chaetognath from samples throughout the Pacific and from the Arctic Ocean. The specimens lose their character- istic crystalline transparency and become chalky-white in reflected light or opaque- grey in transmitted light. The body loses its tubular shape and becomes flattened dorso- ventrally, sometimes to an extreme degree. The body is often distorted into an arc. The tail segment often shortens extremely and becomes relatively wider before the trunk deteriorates. The shortening of the body length can be extreme, specimens of Flac- cisagitta hexaptera that would normally be 40 mm long can shorten to 20 mm. The head may become greatly shortened and thus appear relatively wider, with the jaws thrown widely apart, and the neck constricted.
Thus the three characters which might possibly distinguish Kassatkina’s Tokio- kaispadella from Pterosagitta draco, trans- verse musculature, short tail segment, and relatively wide body, in my opinion, are artifacts. The genus Tokiokaispadella and the family Tokiokaispadellidae based upon it are both invalid.
Status of the Genera Zahonya Oye, 1918 and Aberrospadella Kassatkina, 1971
In 1918 van Oye described a new species and genus from the Java Sea, Zahonya ces- toda. No one has found an example of this species since the original description; how-
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ever, it has been mentioned in review ar- ticles, for example, Tokioka (1965). It seems remarkable that almost no one (Tokioka 1952 excepted) has recognized this species for what it is, a poorly preserved individual showing all the characteristics of the genus Sagitta (sensu Ritter-Zahony, 1911) except for the transverse musculature which clearly is an artifact of morbidity before killing with a preservative. Oye’s description and draw- ing are not adequate to distinguish the spec- imen from the more than 25 epiplanktonic species of sagittids that occur in that region, although Tokioka’s supposition (1952) that it might be a small individual of Sagitta robusta or Sagitta ferox is a reasonable one. Thus, Zahonya cestoda is best left as a no- men dubium with the genus Zahonya a ju- nior synonym of Sagitta.
In 1971 Kassatkina described a new ge- nus and species, Aberrospadella verruculosa from Possjet Bay in the Sea of Japan near Vladivostok. The description is based on two contorted specimens 1.4 mm and 3.4 mm long. No seminal vesicles nor ovaries are present. The specimens almost certainly are poorly preserved or diseased individuals of Parasagitta elegans. Only the rayless lat- eral fins, rayless tail fin and the supposed transverse musculature differentiate the species from juvenile Parasagitta elegans (see Kotori 1975 for detailed descriptions of early larval stages of Parasagitta elegans). The species has two sets of teeth and the number of hooks agrees with P. elegans at that size. As Kotori showed, P. elegans emerges from the egg with one pair of lateral fins. In juvenile P. elegans the collarette is massive, the ventral ganglion is relatively very large, and the tail segment relatively large, all characteristics of Aberrospadella verruculosa. Gut diverticulae are present in both species. Although the absence of rays in the fins is something of an enigma, until the species is redescribed based on mature specimens that are clearly in an excellent state of preservation, this species is anomen dubium and the genus a junior synonym of Parasagitta.
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Acknowledgments
I express my grateful thanks to Dr. Thom- as E. Bowman for his advice, encourage- ment, and critical reading of the manu- script.
Literature Cited
Aida, T. 1897. The Chaetognatha of. Misaki Har- bor.—Annotationes Zoologicae Japonenses 1: 13-21, pl. 3.
Alvarifo, A. 1967. The Chaetognatha of the NAGA expedition (1959-1961) in the South China Sea and the Gulf of Thailand. Part 1-Systematics. — Naga Report 4, pt. 2:1-197.
Bieri, R. 1966. The function of the “wings” of Prer-
osagitta draco and the so-called tangoreceptors
in other species of Chaetognatha. — Publications
of the Seto Marine Biological Laboratory 14:
23-26.
1974. First record of the chaetognath genus Krohnittella in the Pacific and description of a new species. — Wasmann Journal of Biology 32: 297-301.
Germain, L., & L. Joubin. 1912. Note sur quelques Chétognaths nouveaux des croisieres de S. A. S. le Prince de Monaco.—Bulletin de I’Institut Océanographique 228:1-15.
Ghirardelli, E. 1952. Osservazioni biologiche e sis- tematiche sui Chetognati del Golfo di Napoli. — Publicazioni della Stazione Zoologica di Napoli. 23:296-312.
Goto, T., & M. Yoshida. 1987. Nervous system in Chaetognatha. Pp. 461-481 in M. A. Ali, ed., Nervous systems in invertebrates. Plenum Press, New York.
Grassi, B. 1883. I Chaetognati.—Fauna und Flora des Golfes von Neapel, Monograph 5:1-126, 13 pls.
Kassatkina, A. P. 1971. New neritic species of chae- tognaths from Possjet Bay in the Sea of Japan. Pp. 265-294 in Fauna and flora of the Possjet Bay of the Sea of Japan, explorations of the Fauna of the Seas 8, Zoological Institute Acad-
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
emy of Sciences of the U.S.S.R., Nauka Press,
Leningrad (in Russian).
. 1980. A new genus Tokiokaispadella and its
position in the system of Chaetognatha.—Zoo-
logichesii Zhurnal 52:1202-1207 (in Russian).
Kotori, M. 1975. Morphology of Sagitta elegans (Chaetognatha) in early larval stages. — Journal of the Oceanographic Society of Japan 31:139- 144.
Michael, E. L. 1919. Report on the Chaetognatha collected by the United States fisheries steamer ‘“‘Albatross”’ during the Philippine Expedition, 1907-1910.—Bulletin 100, United States Na- tional Museum, Smithsonian Institution 1:235- 277, pls. 34-38.
Michel, H. B. 1984. Chaetognatha of the Caribbean Sea and adjacent areas. NOAA Technical Re- port NMFS 15:1-33.
Nagasawa, S., & T. Nemoto. 1985. The decay of chaetognaths.—La Mer, Bulletin de la Société Franco-Japonaise d’Océanographie 23:56-63.
Owre, H. B. 1973. A new chaetognath genus and species with remarks on the taxonomy and dis- tribution of others. — Bulletin of Marine Science 23:948-963.
Oye, P. van. 1918. Untersuchungen tiber die Chae- tognathen des Javameeres.— Contributions a la Fauna des Indes Néerlandaises 4:1-61.
Ritter-Zahony, R. von. 1911. Revision der Chato- gnathen.— Deutsche Siidpolar Expedition 1901- 1903, 13:1-71.
Salvini-Plawen, L.,V. 1986. Systematic notes on Spa- della and on the Chaetognatha in general. — Zeit- shrift fiir zoologische Systematik und Evolu- tionsforschung 24:122-128.
Tokioka, T. 1940. Phylum Chaetognatha.— Fauna
Nipponica 5(2):1-129.
1952. Chaetognaths of the Indo-Pacific. — Annotationes Zoologicae Japonenses 25:307— 316.
1965. The taxonomical outline of Chaetog- natha.— Publications of the Seto Marine Bio- logical Laboratory 12:335-357.
175 Brookside Drive, Yellow Springs, Ohio 45387.
PROC. BIOL. SOC. WASH. 102(4), 1989, pp. 977-986
HANSENOTHURIA BENTI, NEW GENUS, NEW SPECIES (ECHINODERMATA: HOLOTHUROIDEA) FROM THE TROPICAL WESTERN ATLANTIC: A BATHYAL, EPIBENTHIC HOLOTHURIAN WITH SWIMMING ABILITIES
John E. Miller and David L. Pawson
Abstract. — Hansenothuria benti, new genus, new species, is described and illustrated from material collected with the Research Submersibles Johnson- Sea-Link I and II on bathyal slopes of the Bahama Islands at depths of 639- 904 m. This species was also observed at St. Vincent, Lesser Antilles, at depths of 363-447 m. On several occasions, H. benti has been observed to swim in
response to physical disturbance.
From 1983 to 1988, in collaboration with Dr. Gordon Hendler (Los Angeles County Museum— LACM) and Porter M. Kier (Di- rector Emeritus, National Museum of Nat- ural History, Smithsonian Institution—SJ), we studied the composition and ecology of the echinoderm fauna of the bathyal slopes of the Bahama Islands using the research submersibles Johnson-Sea-Link (JSL) I and II (Harbor Branch Oceanographic Institu- tion, Inc.—HBOl]). A total of 84 dives was made to depths of 200-904 m, and the di- verse echinoderm fauna, comprising at least 180 conspicuous species, was sampled, pho- tographed, and videotaped. Of the 10 bathyal holothurian species encountered, one of the most common proved to be a new genus and species of the Family Syn- allactidae. In April 1989, after this paper was accepted for publication, the authors participated in a JSL expedition to the Less- er Antilles, and this new species was again observed, on seven of 10 dives conducted off the west coast of St. Vincent. It is of particular interest for it has the ability to swim when disturbed. It is a facultative swimming species (sensu Miller & Pawson 1990) whose swimming behavior is de- scribed in detail elsewhere (Miller & Paw- son 1990).
The Family Synallactidae has a world- wide distribution in bathyal and abyssal depths, and at present it comprises approx- imately 15 genera and 120 species. Synal- lactid genera and species are usually difficult to characterize, and the family is in urgent need of a thorough revision. Under these circumstances we are reluctant to erect a new genus for the species described here, but its unique diagnostic features preclude referral of the species to any of the existing confamilial genera.
Methods
The JSL submersibles carry four passen- gers to a maximum depth of 904 m for 3- 5 h per dive. Each submersible is equipped with an array of sophisticated tools that al- low the occupants to sample, photograph, and videotape benthic or pelagic organisms. During a dive, several environmental pa- rameters, for example temperature, con- ductivity and depth, are automatically monitored and recorded at intervals pre- selected by the scientist. Additional infor- mation on the submersibles can be found in Askew (1984).
For in situ photography, we used a laser- aiming device (Fig. 2, U.S. Patent #4,
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FILL-IN STROBE
~ Pal / et oy
CAMERA FIELD OF| VIEW: 7” | 4 | a
J “4 ?
Fig. 2. Diagram of laser-aimed camera system (U.S. Patent #4,777,501) developed for the Johnson-Sea- Link submersibles to photograph benthic organisms at bathyal depths.
777,501) developed by HBOI engineers to aim and focus a Benthos camera system (Model 372) on the animals being studied. A detailed description of the laser-aiming system is found in Caimi & Tusting (1987). Because the distance between the film plane and the subject photographed remained constant throughout a dive, we were able to calculate the exact field of view in each pho- tograph and thereby determine the precise length of living specimens prior to collec- tion.
— Fig. 1.
Order Aspidochirotida Grube, 1840 Family Synallactidae Ludwig, 1894 Hansenothuria, new genus
Diagnosis. —Body fragile, translucent, semi-cylindrical, tapering gently anteriorly and posteriorly, four to five times as long as broad. Ventral surface flat, body arched, middorsal interradial area depressed. Ven- trolateral margin fringed with short to long papillae joined by continuous brim; brim broadest anteriorly, where component po-
Hansenothuria benti, new genus and species: Jn situ photographs—A, French Bay, San Salvador Is.,
JSL-I-2005, 694 m, TL = 20.3 cm; B, off Plana Cays, JSL-I-2274, 829 m, TL = 17.2 cm; C-E, swimming postures, off Cockburn Town, San Salvador Is., JSL-I-2331, 649 m, TL = 23.0 cm, bright streak to left of posterior end of specimen in E is sediment-laden mucus strand shed by active swimming movements; F, Fernandez Bay, San Salvador Is., JSL-I-2048, 759 m, TL = 16.5 cm; G, Fernandez Bay, San Salvador Is., JSL- I-2048, 759 m, TL = 21.5 cm. Photo credit for C-G: C. Young and L. Cameron, HBOI.
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dia may reach 25 mm length. Midventral radius naked; ventrolateral radii with nu- merous, minuscule hair-like feet in two or three rows. Each lateral dorsal radius with 10-31 short (15 mm), sharply pointed pa- pillae arranged in slightly zigzag row. Ex- cepting endplates in ventral feet, ossicles ex- clusively simple rods, occurring only in tentacles, papillae and feet. Gonad as two tufts of tubules, one to each side of dorsal mesentery.
Remarks.—Hansenothuria superficially resembles some species of the genera Bathy- plotes Ostergren, 1898 and Paelopatides Théel, 1886 in body shape and in possessing a marginal brim that aids in swimming but it differs from these genera, and most other synallactid genera, in possessing very sim- ple body wall ossicles. Hansenothuria fur- ther differs from Bathyplotes in having a dorsally positioned anus, and from Paelo- patides in lacking feet along the midventral radius. Ossicles appear to be consistently absent from Benthothuria Perrier, 1898 and Paroriza Herouard, 1923; neither of these genera seems to bear any close relationship to Hansenothuria.
Type species. —Hansenothuria benti, new species, designated herein.
Etymology. —The genus name is of fem- inine gender. This holothurian is named for our friend and colleague, the late Dr. Bent Hansen, Zoological Museum, Denmark, in recognition of his superb contributions to our knowledge of deep-sea holothurians.
Hansenothuria benti, new species Figs. 1, 3; Tables 1, 2
Diagnosis. — As for the genus. Color in life variable, light blue to pale purple; dorsal papillae translucent to black.
—
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Material examined.—Collection and lo- cality data for the Bahama Islands speci- mens/photographs studied during our dives aboard Johnson-Sea-Link I and II are pre- sented in Table 1. Due to the delicate nature of this species, it was difficult to preserve; accordingly only a few specimens collected preserved well enough to be catalogued as museum material. The following specimens have been deposited at the National Mu- seum of Natural History, Smithsonian In- stitution (USNM), the Indian River Coastal Zone Museum, HBOI (IRCZM) or the Los Angeles County Museum of Natural His- tory (LACM).
Holotype: USNM E38201, 12.5 cm total length (TL), JSL-II-808, 699 m.
Paratypes: one specimen, USNM E38202, 10.8 cm TL, JSL-II-808, 700 m; one spec- imen, USNM E38203, 10.0 cm TL, JSL-I- 2009, 694 m; one specimen, USMN E38204, 8.0 cm TL, JSL-I-2264, 903 m; one speci- men, IRCZM 71:00447, 12.0 cm TL, JSL- I-2264, 904 m; two specimens, USNM E38205, too damaged to measure, JSL-I- 2274, 814 m; one specimen, USNM E38206, 20.3 cm TL, JSL-I-2274, 821 m; USNM E38207, 21.5 cm TL, JSL-I-2274, 817 m; one specimen, LACM 88-205.1, 15.0 cm TL, JSL-I-2264, 895 m.
External morphology. —Subcylindrical species of moderate size; 13-23 cm TL in life, approximately four to five times as long as broad (Fig. 1D). Body fragile; body wall thick, gelatinous, transparent; internal structures (intestine, Fig. 1A, G; gonad, Fig. 1C-E) usually visible through body wall. Anterior and posterior ends gently tapering; anterior end high, bluntly rounded (Fig. 1B), posterior end low, narrowed to form short “*tail” (Fig. 1A). Ventral surface a flattened sole with minute, hairlike tube feet along
Fig. 3. Scanning electron micrographs of skeletal ossicles from Hansenothuria benti: A-C (holotype of 12.5 cm TL, measured preserved) A, Rods from dorsal papillae; B, Rods from ventrolateral papillae; C, Rods from tentacles; D-F (paratype of 10 cm TL, measured from seafloor photograph) D, Rods from dorsal papillae; E, Rods from ventrolateral papillae; F, Rods from tentacles.
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Table 2.—Length of skeletal rods in appendages of Hansenothuria benti. n, number of ossicles measured; X, mean; SD, standard deviation; ossicles taken from holotype, 12.5 cm TL (preserved length).
Length (um) of rods from: n xe SD Range Lateral papillae of brim 25 155.6 34.4 109.3 — 230.2 Dorsal papillae 2S 180.5 41.1 109.3 — 260.5 Ventrolateral feet 25 226.6 59.3 114.0 — 376.7 Tentacles 25 266.6 67.6 178.8 — 423.5
lateral radii (Fig. 1C—E); midventral radius naked. Ventrolateral margin composed of brim of webbed podia (Fig. 1 F). Brim widest at anterior and posterior ends of body (Fig. 1D). Bodywall high with steep-sloping sides; dorsal interradius distinctly sunken, form- ing shallow trough between radii (Fig. 1A). Each dorsal radius with 10-31 slender, hair- like papillae with pointed tips (Fig. 1A, B, E-G). Papillae regularly spaced in slightly zigzag rows, extending along length of body. Number of papillae per radius tending to increase with size of specimen. Papillae completely contractile, varying in length up to 15 mm. Mouth ventral, at extreme an- terior end (Fig. 1C—E), surrounded by 20 peltate tentacles. Tentacles, short, with thick trunks, terminating in flat, circular disks. Anus dorsal, situated just above posterior brim (Fig. 1©),
Internal anatomy. —Holotype (USNM E38201) and one paratype (USNM E38202) dissected to examine internal features; both specimens lacking intestinal tract and res- piratory trees. Holotype (male): Two bul- bous polian vesicles attached to circum-oral water ring. Gonad in 2 tufts, one on each side of dorsal mesentery; each tuft with up- wards of 10 tubules, some branching 2-3 times, others unbranched. Tubules extend- ing posteriorly for two-thirds length of body cavity. Radial longitudinal muscles un- equally developed; dorsal muscles larger than ventral.
Paratype (female): Gonad apparently ful- ly developed, tubules up to 30 mm TL. Tu- bules swollen throughout length, filled with
eggs in germinal vesicle stage. Eggs dense, yolky, subspherical; measuring up to 600 um in diameter. Longitudinal muscle bands divided; those to either side of dorsal mes- entery with four bundles of strands; re- maining three bands with two bundles each.
Ossicles (see Table 2 for measure- ments).—Only two ossicle types occur in body wall. Simple rods present in dorsal papillae (Fig. 3A, D), papillae of lateral brim (Fig. 3B, E), tentacles (Fig. 3C, F), and ven- tral feet. Rods straight to curved, with one or more enlarged areas near middle, and few to several spinules near ends. Average length of rods from tentacles and ventral feet con- siderably longer than those from dorsal and lateral papillae (Table 2). Delicate end plate ossicles, composed of two or more pieces, in tube feet; plates have polygonal perfo- rations and irregular, scalloped margins.
Coloration. —Bodywall coloration vary- ing from light blue to pale purple; smaller specimens generally lighter colored than larger individuals. Marginal brim of webbed papillae appears transparent (Fig. 1F), usu- ally lacking pigment. Black flecks scattered on dorsal surface, especially along mid-dor- sal interradius and on dorsal papillae. Fully extended dorsal papillae grey; contracted papillae appear black. Ventrally, bodywall transparent to white with scattered black flecks along each lateral radius and on some tube feet (Fig. 1C, D). Tentacles white; go- nadal tubules, visible through body wall posterior to mouth on ventral surface (Fig. 2C-E), also white.
Habitat. — All specimens collected or ob-
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served at the Bahamas (see Table 1) oc- curred in the upper bathyal zone (639-904 m) on slopes ranging from ca. 5° — 60°. Sed- iment composition at collecting sites varied from a thin layer of biogenic sand covering a limestone pavement to a thick layer of silty sand with scattered coral and algal (Halimeda spp.) rubble. Off the west coast of St. Vincent, H. benti was encountered at depths of 363-447 m, on fairly steep slopes (ca. 30°-50°) covered with terrigenous silty sand.
Distribution. —Hansenothuria benti is at present known from the Bahama Islands at depths of 639-904 m, and from St. Vincent in the Lesser Antilles at depths of 363-447 m. Our submersible observations were re- stricted to depths of less than 904 m; it is likely that the species occurs at greater depths and on bathyal slopes of other Caribbean islands.
Behavior. —Swimming specimens of Hansenothuria benti were observed and videotaped on several occasions. Swim- ming activity was at first observed after three specimens were captured by the submers- ible and placed in collection bins for the ascent to the surface. While in the bin, the holothurians swam by flexing and curling the anterior and posterior ends of their bod- ies. This behavior continued for at least two hours. On several subsequent dives, swim- ming H. benti were videotaped; this species is a facultative swimmer (sensu Miller & Pawson 1990) which spends the majority of its time on the seafloor, venturing upwards into the water column only in response to physical disturbance, e.g., prodding. No in- dividuals of H. benti were swimming when first encountered. Swimming behavior in this species is most likely an adaptation to avoidance of predators or physical hazards, such as sediment slumping. A detailed ac- count of active swimming movements in H. benti is provided in Miller & Pawson (1990).
Remarks. —Of the 10 holothurian species encountered during our dives in the Baha-
985
mas, only the pelagothuriid elasipod En- ypniastes eximia Théel was found to occur in greater abundance than Hansenothuria benti at the depths studied. C. Young and L. Cameron (HBOI, pers. comm.) report seeing aS Many as Six or seven individuals within an area of approximately 225 m7? (the estimated area visible to the occupants of the JSL’s forward compartment when vis- ibility is 15 m). At St. Vincent, H. benti is a common component of the megabenthos; it was the most frequently observed holo- thurian on our dives, with densities reach- ing a peak of four to six individuals/10 m7?.
Because H. benti is a relatively large and common component of the epibenthic in- vertebrate fauna, it might seem surprising that no individuals of this species were tak- en by earlier workers sampling the seafloor with dredges and trawls. The explanation for this may lie in the fact that even spec- imens collected with care by the submers- ible and carried to the surface in seawater- filled containers arrive aboard ship in very poor condition. During the journey from the seafloor to the surface, the holothurian’s epidermal and dermal tissues begin to slough off, and frequently the entire external body- wall is completely autotomized, leaving in- ternal organs surrounded by the longitudi- nal and circular muscle layers.
Acknowledgments
The authors wish to express their grati- tude to their colleagues in the Bahamas echinoderm project, Drs. Gordon Hendler and Porter M. Kier for their assistance. We thank the crews of the Johnson-Sea-Link submersibles and the Research Vessels Johnson, Seward Johnson, and Edwin Link for their dedicated cooperation during sev- eral research missions to the Bahama Is- lands and the Lesser Antilles. We are grate- ful to C. Young and L. Cameron and to researchers of the Division of Marine Bio-
986
technology at HBOI for supplying us with specimens, photographs, videotapes, and locality data of Hansenothuria benti from their submersible programs. Funding for submersible dives was granted through the following organizations (principal investi- gators listed in parentheses): HBOI (J. Mil- ler, C. Young, S. Pomponi); National Sci- ence Foundation (C. Young, L. Cameron); Smithsonian Institution (J. Miller, D. Paw- son). The following HBOI employees are gratefully acknowledged for their patience and perseverence in developing the laser- aimed camera system used in this research: F. Caimi, C. Chulamanis, T. Couture, C. Tietze, R. Tusting. T. Smoyer, HBOI, was most helpful in producing the color plate (Fig. 1). Figure 2 was rendered by J. Has- tings, HBOI. P. Linley, HBOI, assisted with photography of the ossicles (Fig. 3). This paper is HBOI Contribution No. 714 and Smithsonian Institution Marine Station at Link Port Contribution No. 247. Contri- bution No. 14—Studies on bathyal echi-
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
noderms of the Bahama Islands, J. E. Miller (HBOJ), Principal Investigator.
Literature Cited
Askew, T. M. 1984. Submersibles for science—John- son-Sea-Link Land II. Pp. 612-616 in OCEANS °84 Conference Record. Institute of Electrical and Electronic Engineers, Piscataway, New Jer- sey.
Caimi, F. M., & R. F. Tusting. 1987. Applications of lasers to ocean research and image recording systems.— Proceedings of the International Conference on LASERS ’86, Society for Optical and Quantum Electronics, pp. 518-524.
Miller, J. E., & D. L. Pawson. 1990. Swimming sea cucumbers (Echinodermata: Holothuroidea): A survey, with analysis of swimming behavior in four bathyal species.—Smithsonian Contribu- tions to the Marine Sciences (in press).
(JEM) Harbor Branch Oceanographic In- stitution, 5600 Old Dixie Highway, Ft. Pierce, Florida, 34946; (DLP) Department of Invertebrate Zoology, National Museum ~ of Natural History, Smithsonian Institu- tion, Washington, D.C. 20560.
PROC. BIOL. SOC. WASH. 102(4), 1989, pp. 987-1000
ETHEOSTOMA (NOTHONOTUS) WAPITI (OSTEICHTHYES: PERCIDAE), A NEW DARTER FROM THE SOUTHERN BEND OF THE TENNESSEE RIVER SYSTEM IN ALABAMA AND TENNESSEE
David A. Etnier and James D. Williams
Abstract. —A new percid fish, Etheostoma wapiti, a member of the E. mac- ulatum species group, is described from two tributaries to the Tennessee River in northwest Alabama and south central Tennessee. This rare darter, known from 55 specimens, is most closely related to E. vulneratum from which it differs in details of pigmentation and squamation. Etheostoma vulneratum, formerly treated as a subspecies of E. maculatum or E. sanguifluum, is elevated to species level based on phylogenetic analysis of the maculatum species group and subgenus Nothonotus. Comments on the former distribution and the re- maining habitat of E. wapiti are presented.
In our (Williams & Etnier 1978) descrip- tion of Etheostoma aquali we briefly con- sidered the identity of three juvenile Notho- notus, one specimen collected in 1963 from Elk River at Fayetteville, Lincoln County, Tennessee, and two specimens collected in 1884 from Shoal Creek, Florence, Lauder- dale County, Alabama. We suspected at that time that these specimens represented an undescribed species of the E. maculatum species group. No additional specimens were taken in a survey of the Elk River fishes by Jandebeur (1972) or in the several addi- tional collections from the Elk River and Shoal Creek systems prior to 1980. Our hopes of eventually locating an extant pop- ulation and seeing adults were renewed when Charles F. Saylor provided a juvenile from a Tennessee Valley Authority collection from Elk River Mile 41, Giles County, Ten- nessee, and another from Elk River Mile 40.8. A University of Tennessee Regional Faunas Class located a population in lower Richland Creek, a major Elk River tribu- tary, in May 1981 where an adult female and a juvenile were collected. We revisited the Richland Creek locality in June 1982,
in anticipation of finally capturing an adult male in or near breeding condition. We were most interested in whether the red pigment patterns of adult males would be drastically different from those of other members of the E. maculatum species group. We were able to collect five adult males which to our great surprise completely lacked the bright red body spots and red fin markings typical of other members of the species group. Sub- sequent to discovery of the Richland Creek population, main channel Elk River popu- lations have been located at one site in Lin- coln County and two sites in Giles County, Tennessee, associated with anthropogenic limestone rubble, and at one site in Lincoln County, Tennessee and two sites in Lime- stone County, Alabama, where natural limestone bedrock formations abut or cross the river. At present there are only 55 known specimens of this rare but distinctive darter.
Phylogenetic analysis of subgenus Notho- notus, particularly the Etheostoma macu- latum species group, indicates that both E. sanguifluum (Cope) and E. vulneratum (Cope) should be treated as species rather than as subspecies.
988
.
: eee ne ee. ’ ‘ meas
Fig. 1.
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Ce Boe OE eee Ve we
t.® ~~. +) okey ea weet * —-—s i tao ee ee™__j
aS
Etheostoma wapiti, new species, holotype, USNM 288069, male, 71 mm SL (upper specimen) and
allotype, USNM 288070, female, 55 mm SL (lower specimen).
Type material of Etheostoma wapiti is de- posited in the collections of the American Museum of Natural History (AMNH), Academy of Natural Sciences of Philadel- phia (ANSP), California Academy of Sci- ences (CAS), Cornell University (CU), II- linois Natural History Survey (INHS), University of Kansas (KU), Tulane Uni- versity (TU), University of Alabama (UAIC), Florida State Museum (UF), Uni- versity of Michigan (UMMZ), National Museum of Natural History, Smithsonian Institution (USNM), and the University of Tennessee (UT). Comparative material of other species of Nothonotus from the col- lections at USNM and UT were utilized in the preparation of this description. Methods described by Hubbs & Lagler (1958) were used in obtaining counts and measurements
except diagonal scale counts were made ac- cording to the techniques of Raney & Sutt- kus (1964). Vertebral counts were made us- ing the methods of Bailey & Gosline (1955). Swofford’s (1984) PAUP program, version 2.3, was used for phylogenetic analysis. Outgroups utilized include genus Percina and Etheostoma subgenera Allohistium and Litocara for polarizing characters within-ge- nus Etheostoma, with subgenus Oligoceph- alus, the hypothesized sister group of Noth- onotus, used as outgroup for assessing polarity of additional characters within Nothonotus. As our primary objective was to assess relationships within the E. mac- ulatum species group (aquali, maculatum, microlepidum, moorei, rubrum, sanguti- fluum, vulneratum, wapiti), we treated each outgroup as a single taxon, utilizing the most
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989
Table 1.—Standard length (mm), and proportional measurements (in thousandths of SL) of the holotype (male) and four other males and five females of Etheostoma wapiti, new species. X = mean.
USNM
288069
holotype Standard length 70.8 Body depth at dorsal-fin origin Z15 Caudal peduncle depth 138 Caudal peduncle length 220 Pelvic fin length 168 Pectoral fin length 201 Head length 266 Snout length fi Orbit length 66 Upper jaw length 71 Longest dorsal-fin spine 130 Longest dorsal-fin ray 145 151
lst anal-fin spine 95 Longest anal-fin ray
widespread character state as typical for the taxon. For instance, in Oligocephalus only E. radiosum has dark margins on median fins and only E. pottsi and E. swaini have caudal peduncle depth approximating that of Nothonotus. In these cases, the character coded for Oligocephalus was, respectively, median fins lacking dark margins, and cau- dal peduncle slender (see Phylogenetic Analysis, characters 2, 8).
Etheostoma wapiti, new species Boulder Darter Fig. 1, Tables 1-3
Etheostoma rufilineatum Gilbert, 1891:151. Two of five specimens from Shoal Creek, Florence, Alabama.
Etheostoma microlepidum Raney & Zo- rach, 1967:93. Specimens from Shoal Creek, Florence, Alabama.
Etheostoma (Nothonotus) sp. Starnes & Et- nier, 1986. Endemic in Tennessee River drainage.— Biggins, 1987, 1988. Endan- gered status proposed and finalized, re- spectively, under U.S. Endangered Species Act.
Holotype. — Adult male, USNM 288069, 70.8 mm standard length (SL), Richland
Males Females
Range x Range 57.2-70.8 32:2 46.0-57.7 213-223 232 217-251 133-145 129 126-135 213-228 ath 213-222 168-189 191 185-200 199-215 220 212-231 253-278 pp 262-276
67-74 2 68-75
66-69 72 69-74
71-83 74 73-76 120-134 126 120-132 145-156 146 142-150
95-105 101 97-105 144-156 150 137-167
Creek at County Road 4209 crossing, 11.2 air miles south-southeast of Pulaski, Giles Co., Tennessee, 7 Jun 1982, W. C. Dick- inson, D. A. Etnier, M. A. Etnier, C. E. Lou- ton, J. A. Louton, W. C. Pennington.
Allotype. — Adult female, USNM 288070, 55 mm SL, taken with holotype.
Paratopotypes. —Paratypes taken with primary type are TU 148010 (2), UMMZ 213950 (2), and UT 91.3469 (1). Additional paratypes taken at the type locality are UT 91.2203 (2), 22 May 1981.
Other paratypes. —Elk River system, Tennessee: TU 30271 (1), 11 Apr 1963, and UT 91.3459 (1), 13 Sep 1988, Elk River Mile 89.7, Fayetteville, Lincoln Co., 0.25 miles below U.S. Highway 231 bridge. AMNH 58257 (2), ANSP 162645 (2), CAS 64178 (2), CU 71707 (2), and KU 22122 (5), Elk River Mile 61.0, Hamilton Mill, Lincoln Co., 13 Sep 1988. INHS 68229 (3), 14 Apr 1985, and UT 91.3075 (1), 25 Jun 1986, mouth of Indian Creek at Elk River Mile 52.5, Giles Co. UT 91.80 (1), Elk River Mile 41.0, Giles Co., 7 Oct 1980. UT 91.2628 (1), Elk River Mile 40.8, Giles Co.., 2 Aug 1983. UT 91.3072 (4) and UF 44921 (4), Elk River Mile 52.5, at mouth of Indian Creek, Giles Co., 25 Jun 1986. UT 91.3084
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Table 2.—Frequency distribution, sample size, and means of selected scale counts in Etheostoma wapiti, E. aquali, E. vulneratum, and Elk River system E. camurum. Counts for E. aquali from Williams & Etnier (1978)
and E. vulneratum from Zorach & Raney (1967).
Lateral line scales
50 51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 on x E. wapiti 3 125° 7 3-7 FT? AQ = 2- = a de Searowos E. aquali Polls SSF ee Or 86 AP 6) 3103 57 26221 E. vulneratum 1 2. i 32s OANA OF Oe a Sin Sh 5 ame Le 76 57.87 E. camurum 2) a" y= “D) D2 Ree 582 ln al jee | 25) 55:88 Anal-fin origin to first dorsal fin
13 14 15 16 17 18 19 20 n x E. wapiti 1 1 8 18 18 7 53 18.36 E. aquali 5 13 23 13 3 57 17.93 E. vulneratum 1 3 14 23 16 10 6 3 76 KS S7/ E. camurum 3 13 5 3 24 16.33
Second dorsal-fin origin to anal fin
13 14 15 16 17 18 19 n x E. wapiti 4 9 23 a 8 1 52 16.17 E. aquali 1 16 26 13 1 57 14.95 | E. vulneratum 8 11 30 23 4 74 15.19 E. camurum T 8 1 24 15.08
Caudal peduncle scales
18 19 20 21 oD) 23 24 25 n x E. wapiti 6 14 20 10 3 53 22.81 E. aquali D, 28 13 10 3 56 2271 E. vulneratum 3 6 15 26 18 8 76 21.97 E. camurum D 9 3 8 2 24 19.96
(7), Elk River Mile 36.7, Giles Co., 17 Oct 1986.
Elk River system, Limestone Co., Ala- bama: UAIC 7851.01 (4), Elk River Mile 30.7, about 3 miles above Smith Hollow Road (Alabama highway 127) bridge, 24 Oct 1986. UT 91.3097 (1), Elk River Mile 29.7, 24 Oct 1986.
Other material not designated as types. — USNM 36670 (2) Shoal Creek, Florence [Lauderdale Co.], Alabama, 1884. A male taken with the holotype was frozen at UT for electrophoretic analysis; counts for this specimen are included in the tables. Two additional specimens, Elk River Mile 52.5,
Giles Co., Tennessee, 14 Sep 1988, were frozen for electrophoretic analysis at UAIC, and are not included in the tables. Diagnosis. —Etheostoma wapiti is a species of subgenus Nothonotus as defined by Page (1981). Within Nothonotus, E. wa- piti belongs to the EF. maculatum species group (Williams & Etnier, 1978), all of which have a few scales associated with the post- orbital spot on the upper cheek (cheek com- pletely naked in other Nothonotus). Within Nothonotus, E. wapiti differs from acuticeps, aquali, jordani, juliae, maculatum, moorei, rubrum, sanguifluum, and tippecanoe in having the combination of dark marginal
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991
Table 3.—Frequency distribution, sample size, and means of selected fin-ray counts and vertebral counts in Etheostoma wapiti, E. aquali, E. vulneratum, and Elk River system E. camurum. Counts for E. aquali from Williams & Etnier (1978) and E. vulneratum from Zorach & Raney (1967). Vertebral counts for E. camurum
not available.
First dorsal fin Second dorsal fin 11 12 13 14 15 n x 11 12 13 14 n x E. wapiti 10 33 10 53 13.00 1 35 17 53 12.51 E. aquali a 49 3 60 12.90 1 pal 33 3 58 12.66 E. vulneratum 4 11 55 6 2 78 12.89 6 39 31 76 12.33 E. camurum 3 18 3 24 12.00 1 12 11 24 12.42 Total dorsal-fin rays Anal-fin soft rays TO ek TA DS ite I? IS n x 7 8 9 10 n x E. wapiti 8 27 16 2 53 25.23 8 ee 8 1 52 8.04 E. aquali 4 OA 5 DS 5 58° PSSA zy” AD 15 59 9.22 E. vulneratum 1 7 As. et 3 1 1G 22S 16 50 9 1 76 7.93 E. camurum 1 2 9 10 2 2A 2A 4S 8 15 l 24 cae Pectoral-fin rays Vertebrae 12 13 14 15 16 n x 38 39 40 4] n x E. wapiti 2 23 28 53 13.49 4 1 10 39.70 E. aquali 1 25 31 2 1 60 13.63 2 na | 1 30 38.97 E. vulneratum 2 23 47 4 76 13.70 6 pas. 1 32 39.84 E. camurum 6 16 Z 24 13.83
bands on median fins of both sexes, scales on the opercles, horizontal dark lines be- tween scale rows on posterior half of body, a naked nape, and a completely scaled belly. Differs from rufilineatum in lacking large basicaudal pale spots, orange lips, and hor- izontal dark markings on the cheek. Differs from bellum, camurum, and chlorobran- chium in lacking well defined pale submar- ginal bands on soft dorsal, caudal, and anal fins, and in having sexual dimorphism in soft dorsal and caudal fins throughout the year (pale yellow with small brown spots in female wapiti). Most similar to E. vulner- atum, but differing from it (and all other Nothonotus except acuticeps, chlorobran- chium, and occasional maculatum) in lack- ing red or orange on fins or body of nuptial males, and in having higher scale counts (Table 2). Other distinguishing character- istics of the species include moderately de-
veloped subocular bar, cheeks without wavy copper-colored lines.
In numerous adult and juvenile males a pale to yellowish submarginal band is pres- ent on the median fins that closely resem- bles that of male and female E. camurum and E. chlorobranchium. Etheostoma cam- urum and E. wapiti are sympatric in Elk River, and subadults and juveniles can be very difficult to separate. The former lacks any trace of a suborbital bar, never has scales associated with the postorbital dark spot, has a more blunt snout, and (Table 3) mod- ally has 12 (vs. modally 13) dorsal spines.
Description. —Etheostoma wapiti is a moderately large species of the subgenus Nothonotus, the five adult (holotype and four paratopotypes) males from the type locality averaging 63.2 mm SL, the largest 70.8 mm SL. The three adult females from this col- lection are smaller, averaging 52.2 mm SL,
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Table 4.—Characters useful in differentiating between five similar species in the Etheostoma maculatum
species group.
Character E. wapiti eee E. maculatum weer E. aquali
Dark margins on anal, soft dorsal,
and caudal fins present present absent absent absent Shape of caudal fin truncate truncate rounded rounded rounded Red on anal and pelvic fins, adult
male absent absent absent present present Red on margin of spinous dorsal
fin, adult male absent present absent present present Copper colored lines on cheek absent absent absent absent present Dark suborbital bar present present often present present absent Modal vertebral number 39 or 40 40 38 39 39 Mean lateral-line scales 61.65 S737, 60.91 5655 6221 Mean and (mode) of total dorsal-
fin rays 25.23 (25) 25.28(25) 24.65 (24-25) 24.97(25) 25.53 (25-26) Mean and (mode) of anal-fin soft
rays 8.04 (8) 7.93 (8) 8.54 (8-9) 8.18 (8) 9.229)
the largest 57.7 mm SL. Proportional mea- surements of the holotype and nine para- types of E. wapiti are given in Table 1. The general body shape of E. wapiti is illustrated jPod yy sas be
Frequency distributions of scale, fin-ray, and vertebral counts for E. wapiti are given in Tables 2, 3. Body scaled except for the breast, prepectoral, and nape areas. Cheeks naked except for the typical presence of 2— 5 embedded to partially exposed cycloid and/or ctenoid scales behind eye. Opercles scaled. Lateral-line complete with 55-69 (x = 61.6) scales. Transverse scales, anal-fin origin to first dorsal fin 15-20 (x = 18.4) and origin of second dorsal fin to anal fin 14-19 (x = 16.2). Caudal peduncle scale rows 21-25 (X = 22.8). Dorsal fin with 12- 14 (x = 13.0) spines and 11-13 (* = 12.5) soft rays. Total dorsal fin elements 24—27 (x = 25.2). Anal fin with 2 spines and 7-9 (x = 8.0) soft rays. Pectoral-fin rays 12-14 (x = 13.5). Vertebrae 39-41 (X = 39.7). Branchiostegal rays 6-6, branchiostegal membranes separate. Frenum broad. Table 4 contains a summary of characters useful in differentiating between Etheostoma
aquali, E. maculatum, E. sanguifluum, E. vulneratum, and E. wapiti.
Cephalic sensory canal pores are as fol- lows: lateral canal pores 5-6, usually 5; preoperculomandibular canal pores 10-11, usually 10; infraorbital canal pores 7-9, usually 8; supraorbital canal pores 3; su- pratemporal canal pores 3; coronal pore sin- gle.
Coloration. — Following color description is based on the series collected at type lo- cality on 7 Jun 1982. Body of males olive to grayish without red spots. Posterior half of body with 10-14 dark horizontal stripes between scale rows. Stripes are absent on belly and become pale dorsally and ven- trally along caudal peduncle. Humeral scale black. Belly, breast, and prepectoral area grayish. Head grayish with distinct dark gray to black suborbital bar and postorbital spot. Iris dark with some yellow pigment. Cheeks uniform gray without chromatic markings. Gular area pale blue. Spinous dorsal fin dark gray with thin black marginal band and nar- row pale yellow submarginal (marginal an- teriorly) band. Soft dorsal fin gray with black marginal band and pale yellow submarginal
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band. Black marginal band on spinous and soft dorsal fins is wider and darker posteri- ad. Anal fin gray with narrow dark gray to black marginal band on posterior third of fin. Membrane between anal spines pale bluegreen. Caudal fin grayish with faint bluegreen base, pale yellow submarginal band, and black marginal band. Pelvic fins grayish with faint bluegreen membranes near base. Pectoral fins grayish with dusky mar- gin posteriorly. Subadult males and unsexed juveniles collected during June and October 1986 had orange to red on caudal fin, nar- row red margin and occasional red anterior interradial membranes of spinous dorsal fin, and occasional dirty red spots on posterior sides. Larger males from these collections lacked red or orange colors, except one Oc- tober adult male had about ten obscure dirty red spots along posterior sides and reddish caudal fin. In the 13 adults (all males 43- 62 mm SL) collected on 13 Sep 1988 and examined on 7 Oct, 1 of the smaller spec- imens had marginal red-orange on interra- dial membranes 1, 2, 4, and 6 of the spinous dorsal fin, and an additional 3 had marginal red-orange on membrane 1. Five of these had a few dirty red spots on the sides and five had orange on dorsal and ventral prin- cipal caudal fin rays in contrast to the pale yellow middle rays. The five largest males in this series (55-62 mm SL) were notice- ably less colorful—none had red in the spi- nous dorsal fin, one had the red-orange cau- dal fin color described above, and four had pale yellow rays in either the soft dorsal or anal fin.
In females, coloration of head and body similar to that of males except paler. Spi- nous dorsal fin dusky gray, margin of first interradial membrane red. Soft dorsal and caudal fin pale yellow with dark brown speckles. Anal fin dusky gray, not speckled with brown. Median fins with narrow black marginal band. Pectoral and pelvic fins were colorless.
Distribution and habitat.—Etheostoma
993
Fig. 2. Geographic distribution of Etheostoma wapiti (stars), E. vulneratum (dots), E. aquali (dia- monds) and E. sanguifluum (triangles).
wapiti is known from the Elk River system in Tennessee and the Shoal Creek system in Alabama, both northern tributaries of the Tennessee River, along the southern por- tion of the Highland Rim physiographic province (Fig. 2). It was first collected in Shoal Creek in 1884 by C. H. Gilbert and J. Swain and subsequently reported (as E. rufilineatum) by Gilbert (1891). No habitat data were given and the only locality infor- mation was “Shoal Creek, Florence, Ala- bama.” Shoal Creek, located east-northeast of Florence, Lauderdale Co., enters the Ten- nessee River about seven miles east of Flor- ence. The lower portion is flooded by back- waters of Wilson Dam which extend upstream to the Lauderdale County Road 64 crossing. The two specimens of E. wapiti collected by Gilbert and Swain were most likely collected in the lower portion of Shoal Creek. While recent extensive efforts to col- lect E. wapiti in the unimpounded portion of Shoal Creek have been unsuccessful, it may still occur there since the remaining stream habitat appears to be in good con- dition and supports a large variety (50 species) of fishes.
The boulder darter is currently known from eight localities in the main channel of Elk River and from the lower reaches of two Elk River tributaries, Richland and Indian creeks. Adults have been found only in areas
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of boulder substrate, and we consider this strong circumstantial evidence that it shares egg-clumping reproductive habits with oth- er members of the maculatum species group. In Elk River proper the most upstream rec- ord (two specimens, River Mile 89.7, Fay- etteville, 1963, 1988) is in an area strongly influenced by cold water releases from Tims Ford Reservoir, completed in 1970. The 12 adult males and | juvenile from Elk River Mile 61.0, 13 Sep 1988, were collected in a high energy bend of the river where boulders eroded from an adjacent bluff provide hab- itat. Single juveniles from River Miles 40.8 and 41.0 are likely waifs that dispersed downstream from the Richland Creek pop- ulation (mouth at River Mile 42.6). Else- where in Elk River, Tennessee, the popu- lation at the mouth of Indian Creek (River Mile 52.5) is apparently utilizing limestone slabs from a collapsed mill dam for spawn- ing substrate; juveniles from this population enter lower Indian Creek, but no adults have been taken there and suitable spawning hab- itat is virtually absent. At River Mile 36.7, adults were taken in association with boul- ders from a former stone bridge. In Elk Riv- er, Alabama, natural outcrops of limestone occur above and below the Smith Hollow Road (Alabama Highway 127) bridge, and boulder darter populations are presumably rather continuous from River Mile 30.7 (ca. 400 m above bridge) downstream to about River Mile 29. At River Mile 28.1 (up- stream end of Gallus Island) Elk River is impounded when Wheeler Reservoir is at full pool, and boulder darters were not col- lected.
Historically the boulder darter probably occurred in the main channel of the Ten- nessee River between Muscle Shoals and the mouth of the Elk River, a distance of ap- proximately 45 miles. It is likely that it also occurred upstream from the mouth of Elk River and in the lower portion of other northern tributaries of the southern bend of the Tennessee River such as the Flint and Paint Rock rivers.
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Conservation.—There are presently 55 known specimens of Etheostoma wapiti. Its habitat is difficult to sample with standard kick-seining methods which may account, at least in part, for its rarity. However, most of its potential habitat of deep, rocky, flow- ing pools in rivers and lower portions of large tributaries has been altered by im- poundment, pollution, and siltation. It is likely that additional reproducing popula- tions could be established in Elk River by merely providing suitable spawning sub- strate in areas with adequate current and good water quality. The restricted distri- bution of E. wapiti and loss of potential habitat in other large rivers of the area prompted the U.S. Fish and Wildlife Ser- vice to list it as an Endangered species (Big- gins, 1988).
Etymology. —The species name, wapiti, is an American Indian name for the American elk and calls attention to the Elk River sys- tem, the only habitat presently known to support the species. The common name, boulder darter, is in reference to the habitat of adults.
We suggest the vernacular names of wounded darter for E. vulneratum and bloodfin darter for E. sanguifluum. These names are suggested by the meanings of the Latin vulneratum (=wounded) and sangut- fluum (=flowing blood).
Phylogenetic analysis. —Following are characters utilized in assessing relationships within subgenus Nothonotus, with a brief survey of their distribution in outgroups and in Nothonotus. Our polarity judgements are included, but the PAUP program was run with polarity unfixed.
1. Horizontal dark lines on sides of body above and below lateral line. This character does not appear in any of the outgroups, and is essentially a subgeneric synapomor- phy, secondarily lost only in jordani and tippecanoe as suspected autapomorphies.
2. The deep-bodied, slab-sided form of Nothonotus species (expressed as caudal pe- duncle depth divided by SL, data from Page
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1981, and UT specimens) is a presumed synapomorphy for the subgenus. The cau- dal peduncle is slender in genus Percina (above ratio = 0.065-0.100) and in Etheo- stoma subgenera Allohistium (0.098) and Litocara (0.088—0.092). In subgenus Oligo- cephalus Page’s (1981) values range from 0.089 to 0.116, with only grahami (0.115), pottsi (0.116), and swaini (0.112) having ra- tios higher than 0.108. Ten adults of both grahami and swaini from the UT collection were measured, and we get a value identical to Page’s for swaini, but a value of 0.103 for grahami. In Nothonotus, Page’s values range from 0.111 to 0.128. We rechecked three of Page’s lower values with ten UT specimens, and get an identical value for bellum (0.114), but higher ratios for jordani (0.120 vs. 0.111) and chlorobranchium (0.133 vs. 0.118).
3. The darkened anterior interradial membranes of the spinous dorsal fin occur throughout subgenus Nothonotus, with sim- ilar pigment occurring in outgroups only in four of nine species of Percina subgenus A/- vordius and in Percina (Imostoma) shu- mardi. We consider it to be a synapomor- phy for Nothonotus.
4. Males with distinctive nuptial colors of blue, green, gray, or brown on breast. Nothonotus is often accorded the subgeneric vernacular of “‘bluebreast darters,”’ and this presumed synapomorphy is consistent throughout the subgenus except in juliae (males darken on throat and breast, James & Taber 1986), and in jordani where green of the breast may extend to adjacent bran- chiostegal membranes and/or belly. In out- groups, nuptial breast color of this nature is absent (Percina, Allohistium, Litocara) or occasionally present only in /epidum (green branchiostegal membranes and breast) and whipplei (blue breast). In some other Oli- gocephalus (caeruleum, radiosum, specta- bile) the orange-red of the branchiostegal membranes may extend onto the breast.
5. Naked nape. The consistent and ex- cessive loss of nape squamation, typical of
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all Nothonotus except juliae, does not occur in outgroups, and is considered a synapo- morphy uniting Nothonotus other than ju- liae. We interpret the scaly nape of juliae as symplesiomorphic with outgroups.
6. United gill membranes, an autapo- morphy occurring in ju/iae, are absent from outgroups and from other Nothonotus.
7. Naked opercles. Autapomorphy, acu- ticeps. Opercles are consistently scaled in outgroups and other Nothonotus.
8. Marginal dark bands on caudal, soft dorsal, and anal fins. A presumed synapo- morphy for Nothonotus except juliae, acu- ticeps, and tippecanoe. The character is absent in outgroups except in E. (Oligoceph- alus) radiosum. Characters 9, 10, 11, and 14 support our assuming this to be a shared reversal in aquali, maculatum, and sangui- fluum. Character 10 suggests a possible ho- moplasy in tippecanoe.
9. Red spots on body of nuptial males. A synapomorphy for all species of Nothonotus except acuticeps, juliae, tippecanoe, and wapiti. Character 10 suggests a possible re- versal in tippecanoe. Presence of red spots in juvenile males plus characters 10, 11, 14, and 15 indicate an obvious reversal in nup- tial male wapiti. In outgroups, similar pig- ment occurs only in E. (Oligocephalus) whipplei.
10. Sexual dimorphism of fin pigment pattern throughout the year. The darkly speckled median fins of females of rufiline- atum, jordani, tippecanoe, and the macu- latum species group are markedly different from the pattern in males and persist throughout the year and in preservative. Ex- cept for tippecanoe, where it is tentatively treated as an autapomorphy, we consider it a synapomorphy for these species. In Per- cina, sexual pattern dimorphism in median fins is essentially absent (cannot be sexed in preservative) and sexual dichromatism is absent or weakly expressed during the breeding season except in subgenera Eri- cosma, Hypohomus, and P. (Alvordius) cras- sa and roanoka. In genus Etheostoma out-
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groups and other Nothonotus, sexual dichromatism may be spectacular, espe- cially during the breeding season, but these are alcohol-soluble pigments that are lost in preservative, again making accurate pat- tern-based sexing of museum specimens dif- ficult or impossible.
11. Presence of scales on upper cheek, near postorbital spot. Although scaly cheeks are widespread in outgroups, and naked cheeks are routinely and, we suspect, cor- rectly considered the derived condition in percids, we note that these scales, absent from Nothonotus except the maculatum species group (aquali, maculatum, micro- lepidum, moorei, rubrum, sanguifluum, vul- neratum, wapiti) must be considered de- rived under the most parsimonius phylogenetic hypothesis.
12. Reduced belly squamation. A syn- apomorphy for moorei and rubrum. In other Nothonotus and in Etheostoma outgroups the belly is fully scaled or occasionally and variably with a small naked area anteriad. In genus Percina, a small portion of the an- terior belly may be naked, or there may be seasonally or sexually variable naked areas corresponding to the position of the mod- ified midventral scales.
13. Reduced vertebral number. A syn- apomorphy for rubrum and moorei and homoplastic in juliae. Modal vertebral counts as low as 36 also occur commonly in subgenus Oligocephalus, but not in other outgroups or in other Nothonotus.
14. Egg clumping. A synapomorphy known or inferred to be shared by six species of the maculatum species group, absent in outgroups, and absent in other Nothonotus. Voirs (1988) confirms that E. moorei buries its eggs in the gravel as is typical of other Nothonotus. It seems likely that E. rubrum will also be an egg-burier given the presence of adult males in gravel riffle areas through- out the breeding season.
15. Anal fin lacking bright colors in males, lacking spots in females. In genus Percina, anal fins of males lack bright colors except
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
in nuptial P. (Ericosma) evides and P. (Hy- pohomus) aurantiaca where it is or may be iridescent blue. In genus Etheostoma out- groups, the anal fin of males is blue, green, or red, or combinations of these colors. In Nothonotus, the anal fin of males is orange, red, or green except in vu/neratum and wapi- ti. In wapiti, a trace of iridescent blue was noted between the anal spines in nuptial males, but the remainder of the fin was gray, while in vu/neratum the entire fin is gray. Failure of vu/neratum and wapiti males to develop brightly colored anal fins during the breeding season (a presumed synapomor- phy) implies significantly different court- ship behavior. Correlated with this, the anal fin of female vu/neratum and wapiti is un- spotted, whereas in the remainder of the maculatum species group plus jordani, rufi- lineatum, and tippecanoe, females have the anal fin spotted (see character 10). While we do not feel justified in considering these as separate characters (anal fin pigmentation of males and females is likely to have the same genetic basis), our confidence in this character’s validity as an indicator of the sister-species relationship between vu/ner- atum and wapiti is certainly enhanced by its deviation from the maculatum species group norm in both sexes.
16. Absence (loss) of dark margins on soft- rayed median fins. In genus Percina and in Etheostoma subgenera Allohistium and Li- tocara, median fins other than the spinous dorsal lack dark margins. In Oligocephalus, a dark margin on the soft dorsal is frequent- ly present, but only in radiosum do all three soft-rayed median fins have dark margins. In Nothonotus, darkly margined median fins are absent only in acuticeps, juliae, and tip- pecanoe (presumed symplesiomorphy), and as a presumed synapomorphy in aquali, maculatum, and sanguifluum of the mac- ulatum species group. Other Nothonotus have dark margins on these fins consistently evident in males and often apparent in fe- males.
17. Rounded caudal fin. This character is
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597
Jjuliae acuticeps tippecanoe bellum camurum chlorobranchium Jjordani rufilineatum moorei rubrum microlepidum vulneratum Wapiti
aquali maculatum
ist) 7/ Sanguifluum
Fig. 3. Hypothesized phylogenetic relationships within subgenus Nothonotus. Characters 1-17 are discussed
in the text. Asterisks indicate suspected homoplasies.
somewhat subjective, but we and Zorach & Raney (1967) find it useful, if somewhat variable, within the maculatum species group. The caudal fin is truncate to slightly forked in outgroups, and Nothonotus except for aquali, maculatum, and sanguifluum. We do note some sexual and size dimorphism in this character, with extremely large males of vulneratum occasionally having the cau- dal fin rounded, and occasional sangui- fluum, especially females and smaller males, having the caudal fin truncate.
Discussion. —While characters 1-17 above are not sufficient to resolve relationships within subgenus Nothonotus, the concensus tree generated (Fig. 3, consistency index = 0.654) indicates that (1) Nothonotus is monophyletic; (2) Etheostoma juliae is sis- ter to all other Nothonotus; (3) Etheostoma acuticeps, E. tippecanoe, and all other Noth- onotus (except juliae) form an unresolved trichotomy; (4) although E. bellum, E. ca- murum, and E. chlorobranchium share all
17 characters analyzed, none is a synapo- morphy for the three species, and their monophyly is not established; (5) the E. maculatum species group is monophyletic as are its branches of moorei + rubrum, aquali + maculatum + sanguifluum, and vulneratum + wapiti; (6) Etheostoma moorei and E. rubrum are sister to remain- ing members of the maculatum species group; and (7) Etheostoma microlepidum is included in an unresolved trichotomy with aquali + maculatum + sanguifluum and vulneratum + wapiti.
Prior to availability of sufficient material of Etheostoma aquali and E. wapiti, Zorach & Raney (1967) recognized E. maculatum as polytypic, including E. m. sanguifluum and E. m. vulneratum. They pointed out (p. 296) that “‘Three allopatric forms are rec- ognized as subspecies, but might properly be considered as species.’’ Our data indicate that vu/neratum and wapiti are sister taxa and that aquali + maculatum + sangui-
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fluum comprise a monophyletic branch, rendering their polytypic E. maculatum paraphyletic.
Our hypothesis concerning relationships between the taxa aguali, maculatum, mi- crolepidum, sanguifluum, vulneratum, and wapiti differs from Page’s (1985) hypothe- sized relationships between the species aquali, maculatum, sanguifluum, and mi- crolepidum in which he treated vulneratum as a subspecies of sanguifluum. In his scheme, the polytypic E. sanguifluum and E. aquali are considered sister species based on the presumed synapomorphy of adult males with “two red spots at the front and one red spot at the rear”’ of the spinous dor- sal fin. Although this pattern is distinctive, there is considerable variation within aquali, sanguifluum, and vulneratum in the extent of red margination, and a complete or near- ly complete marginal band may be present. Marginal red pigment can occur in spinous dorsal fins of both maculatum and wapiti in subnuptial males (description, this paper, and Zorach & Raney 1967), but is absent in nuptial males. Red to orange margination in dorsal fins of male Nothonotus is var- iously expressed in all species except acu- ticeps, and perhaps chlorobranchium, and is often brighter anteriad. Since red margin- ation of the spinous dorsal fin of males is present in subgenera Allohistium and Lito- cara, and often present although more sub- marginal in subgenus Oligocephalus, we treat this character as a symplesiomorphy within Nothonotus, and anterior and posterior con- centration of this pigment, also variously expressed in several Nothonotus, is very sus- pect as asynapomorphy uniting aquali, san- guifluum, and vulneratum. Page’s scheme conflicts with additional characters (round- ed caudal fin, lack of bright colors in pelvic and anal fins of males, lack of dark spots on anal fin of females and lack of dark margins on median fins) that are supported by out- group analysis as being synapomorphies within more derived Nothonotus species. We elevate vu/neratum to species status based
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
on our contention that wapiti and vulnera- tum are sister species, with either microlepi- dum or the unresolved trichotomy of aquali, sanguifluum, and maculatumas sister to that pair. The presumed synapomorphy uniting wapiti and vulneratum is secondary loss of bright colors in the anal and pelvic fins of adult males plus absence of spots in anal fin of females. The clade of aquali, sangui- fluum, and maculatum share the presumed synapomorphies of rounded caudal fin and loss of dark margination on the median fins. Additional characters, whose polarity is dif- ficult to evaluate, support the above rela- tionships. Vertebrae are modally 40 in both vulneratum and wapiti, but modally 39 or fewer in other Nothonotus except chloro- branchium (39 or 40). High vertebral counts in chlorobranchium are likely related to its being the largest and most cold-adapted member of the subgenus, but neither vu- neratum nor wapiti is larger than nor more tolerant of cold water than most other Noth- onotus species. Mean anal fin ray counts are 7.93 and 8.04 for vulneratum and wapiti, respectively, but 8.18 or more for other egg- clumping Nothonotus. Simon et al. (1987) noted that egg diameter (2.8—3.1 mm) and hatchling length (8.3 mm SL) for vulnera- tum (treated as E. sanguifluum vulneratum) were larger than for other Nothonotus stud- ied (egg diameter range 1.3-2.1 mm, hatch- ling length range 3.8—7.2 mm). Etheostoma aqualiand E. maculatum were the only oth- er members of the maculatum group in- cluded in their study. Our examination of egg diameter in gravid females of the mac- ulatum species group (their data were based on fertilized, water-hardened eggs) confirms the large egg size in E. vulneratum, but the two available gravid females of wapiti had eggs within the range of sizes seen in the other species.
Zoogeography. — Dr. R. E. Jenkins (in litt.) has noted that dark marginal bands on me- dian fins are less well developed in Clinch/ Powell/Emory river E. vulneratum speci- mens than elsewhere. He has suggested that
VOLUME 102, NUMBER 4
these might represent intergrades with san- guifluum, but this supposition demands a major headwater piracy event (members of the maculatum species group inhabit streams of order three or larger) between those systems and the Cumberland River drainage. Such a piracy has been alluded to (Ross 1971, Starnes et al. 1977), and sim- ilarities between fish faunas of the upper Cumberland River below the Falls and the adjacent Clinch/Powell/Emory systems cer- tainly suggest such an event. Starnes & Et- nier (1986) rethought and refuted this hy- pothesis based on the complete absence of geological evidence — Cumberland Gap was formed by faulting, and does not represent an extinct watercourse. They attribute fau- nal similarities to physiographic conditions (abundance of Silurian and/or Carbonifer- ous sandstones in the watersheds and sim- ilar stream size and gradient) coupled with formerly more continuous distribution of ancestral forms throughout the Tennessee and Cumberland drainages. We accept this view as most likely, and any similarities be- tween Cumberland sanguifluum and Clinch/ Powell/Emory vulneratum are treated as homoplasies rather than as the result of re- cent contact. Assuming that main channel dispersal rather than headwater piracy has been responsible for present distribution, the ranges of sanguifluum and vulneratum, rather than abutting, are at maximum sep- aration for the four taxa being considered (Fig. 2), with aquali and wapiti occupying geographically intermediate areas. These distributions and our proposed relation- ships of the four taxa conform nicely to the Starnes & Etnier (1986) hypothesis that, based on considerable geological evidence, the upper and middle portions of the Ten- nessee River drainage had a remote outlet from that of the present lower Tennessee and Duck river systems during the late Ter- tiary, and that the Tennessee achieved its present configuration in the Pleistocene. This development may have effected vicariance of the ancestral stock into a middle/upper
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Tennessee component (vulneratum/wapiti precursor) and a Duck/lower Tennessee/ Cumberland river component (aquali/san- guifluum precursor). Further speciation into the four taxa considered here may be at- tributable to strict habitat requirements and physiographic fidelity of these darters (see discussion in Starnes & Etnier 1986). Since vulneratum/wapiti and aquali/maculatum/ sanguifluum are supported as being mono- phyletic groups with aquali and sangui- fluum likely sister species in the latter, a polytypic maculatum including vulneratum and/or sanguifluum would be paraphyletic. Monophyly could be retained by consider- ing all five taxa as subspecies of a polytypic Etheostoma maculatum, a move we con- sider unwarranted. The unusual situation of two species (aquali and wapiti) evolving in- dependently from two taxa that maintain their subspecies status (sanguifluum and vulneratum, respectively) could occur if (a) a founder effect were present, or (b) the al- lopatric ranges of aquali and wapiti pre- sented drastically different selective pres- sures from those prevailing in the total range of sanguifluum and vulneratum, or (c) if the original subspecies continue to have or more recently have had a more continuous dis- tribution than their vicariates. Since none of these, perhaps not exclusive, conditions appears likely, we are uncomfortable in ac- cepting the derivation of two relatively dis- tinct species (aquali and wapiti) from two different subspecies of a polytypic species.
Acknowledgments
During 1986, status surveys were funded by the U.S. Department of Interior, Office of Endangered Species. Discussions with Wayne Starnes, Bob Jenkins, and Larry Page provided valuable input on the biology and evolution of the subgenus Nothonotus. Con- structive suggestions from Wayne C. Starnes, Richard L. Mayden, and three anonymous reviewers resulted in considerable improve- ment over the original manuscript. Jennifer L. Etnier provided the graphics for Fig. 3.
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Literature Cited
Bailey, R. M., & W. A. Gosline. 1955. Variation and systematic significance of vertebral counts in the American fishes of the family Percidae.— Mis- cellaneous Publications, Museum of Zoology University of Michigan 93:1—44.
Biggins, R. G. 1987. Endangered and threatened
wildlife and plants; proposal to list the boulder
darter as an endangered species.— U.S. Federal
Register 52(221):43921-43923.
. 1988. Endangered and threatened wildlife and
plants; determination of endangered species sta-
tus for the boulder darter.—U.S. Federal Reg- ister 53(170):33996-33998.
Gilbert, C. H. 1891. Report of explorations made in Alabama during 1889, with notes on the fishes of the Tennessee, Alabama, and Escambia rivers. — Bulletin of the United States Fish Com- mission 9:143-160.
Hubbs, C. L., & K. F. Lagler. 1958. Fishes of the Great Lakes Region.—Cranbrook Institute of Science Bulletin 26:1-213.
James, P. W., & C. A. Taber. 1986. Reproductive biology and age and growth of the yoke darter, Etheostoma juliae.—Copeia 1986:536—-540.
Jandebeur, T. S. 1972. A study of the fishes of the Elk River drainage system in Alabama and Ten- nessee. Unpublished M.S. Thesis, University of Alabama, Tuscaloosa. 153 pp.
Page, L.M. 1981. The genera and subgenera of dart-
ers (Percidae, Etheostomatini).— Occasional
Papers of the Museum of Natural History of the
University of Kansas 78:1-69.
1985. Evolution of reproductive behaviours in percid fishes. — Bulletin of the Illinois Natural History Survey 33:275-295.
Raney, E. C., & R. D. Suttkus. 1964. Etheostoma moorei, anew darter of the subgenus Nothonotus from the White River system, Arkansas.— Copeia 1964:130-139.
—, & T. Zorach. 1967. Etheostoma micro- lepidum, a new percid fish of the subgenus Nothonotus from the Cumberland and Tennes- see river systems.—American Midland Natu- ralist 77:93-103.
Ross, R. D. 1971. The drainage history of the Ten- nessee River. Pp. 11-42 in P. C. Holt, R. A.
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Patterson, & J. P. Hubbard, eds., The distri- butional history of the biota of the southern Appalachians. Virginia Polytechnic Institute and State University Research Division Monograph No. 4, Blacksburg, Virginia.
Simon, T. P., R. D. Wallus, & K. B. Floyd. 1987. Descriptions of protolarvae of seven species of the subgenus Nothonotus (Percidae: Etheosto- matini) with comments on intrasubgeneric char- acters. Pp. 179-190 in R. D. Hoyt, ed., 10th Annual Larval Fish Conference, American Fisheries Society Symposium 2, Bethesda, Maryland.
Starnes, W. C., & D. A. Etnier. 1986. Drainage evo- lution and fish biogeography of the Tennessee
and Cumberland river drainages. Pp. 325-361
in C. H. Hocutt, & E. O. Wiley, eds., Zooge-
ography of North American freshwater fishes.
Wiley-Interscience, New York.
— , L. B. Starnes, & N. H. Douglas. 1977. Zoogeographic implications of the rediscovery of the percid genus Ammocrypta in the Tennes- see River drainage.—Copeia 1977:783-786. Swofford, D. S. 1984. PAUP. Phylogenetic analysis
using parsimony. Version 2.3. Illinois Natural History Survey, Champaign, Illinois.
Voirs, W. D. 1988. Agonistic and reproductive be- havior of the yellowcheek darter Etheostoma moorei. American Society of Ichthyologists and Herpetologists 68th Annual Meeting, Program and Abstracts, p. 185 (abstract).
Williams, J. D., & D. A. Etnier. 1978. Etheostoma aquali, a new percid fish (subgenus Nothonotus) from the Duck and Buffalo rivers, Tennessee. — Proceedings of the Biological Society of Wash- ington 91:463-471.
Zorach, T., & E. C. Raney. 1967. Systematics of the percid fish, Etheostoma maculatum Kirtland, and related species of the subgenus Nothono- tus.—American Midland Naturalist 77:296—322.
(DAE) Department of Zoology, Univer- sity of Tennessee, Knoxville, Tennessee 37996-0810; and (JDW) National Fisheries Research Center, U.S. Fish and Wildlife Service, Gainesville, Florida 32606.
PROC. BIOL. SOC. WASH. 102(4), 1989, pp. 1001-1017
ELLERKELDIA, A JUNIOR SYNONYM OF HYPOPLECTRODES, WITH REDESCRIPTIONS OF THE TYPE SPECIES OF THE GENERA (PISCES: SERRANIDAE: ANTHIINAE)
William D. Anderson, Jr. and Phillip C. Heemstra
Abstract. —Characters that may prove useful in defining the serranid subfam- ily Anthiinae are briefly discussed, and a single synapomorphy, vertebral num- ber, that unites the species of Hypoplectrodes is recognized. Ellerkeldia is con- sidered a junior synonym of Hypoplectrodes; the relationships of Hypoplectrodes are discussed; the type species (Plectropoma semicinctum and P. nigrorubrum) of the two nominal genera are redescribed; and Scopularia rubra is demon- strated to be a junior synonym of H. semicinctum. Hypoplectrodes semicinctum is known from shallow waters off Juan Fernandez and San Félix islands, and has been reported from Easter Island; H. nigroruber has been collected from shallow Pacific and Indian ocean waters off southeastern, southern, and south-
western Australia.
Some years ago, after examining the orig- inal descriptions of Plectropoma semicinc- tum and Scopularia rubra, one of us (PCH) concluded that the two species are synon- ymous. More recently the senior author ex- amined the holotype of P. semicinctum, compared it with the original description of S. rubra, and arrived at the same conclu- sion. In view of the similarities of the de- scriptions in the literature of species of E/- lerkeldia and of Hypoplectrodes nigroruber, the senior author examined the syntypes of H. nigroruber and determined that this species 1s congeneric with P. semicinctum. Because H. nigroruber is the type (and until now the only) species of Hypoplectrodes and P. semicinctum is the type species of Eller- keldia, it follows that Hypoplectrodes and Ellerkeldia are subjective synonyms. The purposes of this paper are to redescribe Hy- poplectrodes nigroruber and H. semicinctum and to document the assertions of synony- my made above.
Abbreviations and Methods
Institutional abbreviations are as listed in Leviton et al. (1985); ICZN denotes the In- ternational Code of Zoological Nomencla- ture (International Commission on Zoolog- ical Nomenclature 1985); SL signifies standard length and TL, total length.
Methods for making counts and mea- surements are those of Anderson & Heem- stra (1980), except as noted below. Scales below the lateral line were counted oblique- ly, both in posterodorsal and anterodorsal directions from the origin of the anal fin (the posterodorsal direction is apparently the di- rection used by de Buen (1959) on Scopi:- laria rubra).
Instead of scales in the lateral line, de Buen (1959) gave counts of scales in a lon- gitudinal line. We interpret this to mean scales along the body in a mid-lateral line to base of caudal fin. It is difficult to get repeatable counts in a single longitudinal
1002
line of scales along the body; therefore our counts of “‘scales in a longitudinal line’’ are of oblique rows of scales along mid-body from cleithrum to base of caudal fin.
De Buen (1959) gave the lengths of spec- imens of S. rubra as total lengths, but the body proportions as percentages of standard length or head length. Based on the rela- tionship of standard and total lengths in the specimens of Hypoplectrodes semicinctum examined, we have estimated the standard lengths of the types of S. rubra. Some of de Buen’s measurements of S. rubra require interpretation; we have construed them as follows: height of body as greatest depth of body, width of body as greatest width of body, preorbital as length of snout, preven- tral as prepelvic length (premaxillary sym- physis to origin of pelvic fin), and pectoral base as width of base of fin.
In the text some measurements are pre- sented as quotients of the standard length, length of head, length of snout, or diameter of orbit. These quotients are rounded off to the nearest 0.05.
Anthiinae
Johnson (1983) defined the family Ser- ranidae with respect to the Percichthyidae (sensu Gosline 1966) on the basis of three reductive specializations, and demonstrat- ed that members of the Serranidae share at least one innovative specialization—thus demonstrating the monophyly of the fam- ily. Johnson (1983, 1988) followed Gosline (1966) in recognizing three subfamilies in the Serranidae, the Serraninae, Epinephel- inae, and Anthiinae, but was able to define only the Epinephelinae on a character that can be interpreted as being uniquely de- rived. Olmi (1986) found a reductive char- acter in the branchial skeleton that may prove to be a synapomorphy uniting the members of the Anthiinae. In all of the At- lantic and eastern Pacific species of an- thiines that she examined and in all of the Indo-Pacific species for which she could ob-
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tain data the second epibranchial lacks a tooth plate, whereas it is present in all ser- ranines and epinephelines observed in her study. She concluded that the absence of this tooth plate in the Anthiinae appears to be the derived condition in the Serranidae.
As pointed out by Johnson (1983) it is difficult to evaluate the importance of ver- tebral number in determining relationships among the Percoidei; nevertheless this char- acter may ultimately prove useful in cir- cumscribing the limits of the Anthiinae. Members of the Serraninae and Epinephel- inae almost always have 24 vertebrae, but species of Anthiinae have 25 to 28, usually 26 (see section on relationships of Hypo- plectrodes).
Until additional studies have been con- ducted the Anthiinae will continue to be an inadequately defined group. Despite this shortcoming the recognition of the Anthii- nae as a distinct taxon will continue to serve a useful purpose because the concept an- thiine unites a plethora of look-alike species that share at some level within the Serran- idae uniquely derived characters.
Hypoplectrodes Gill, 1862
Hypoplectrodes Gill, 1862:236 (type species Plectropoma nigrorubrum Cuvier, 1828, by monotypy).
Gilbertia Jordan, 1891:346 (type species Plectropoma semicinctum Valenciennes, 1833, by original designation; preoccu- pied by Gilbertia Cossman, 1889, a genus of Mollusca).
Ellerkeldia Whitley, 1927:298 (type species Plectropoma semicinctum Valenciennes, 1833, by virtue of the facts that E/lerkel- dia was proposed as a replacement name for Gilbertia Jordan, 1891, preoccupied by Gilbertia Cossman, 1889, and that a replacement name retains the type of the prior name [ICZN, Article 67h]; Whitley, 1927, incorrectly considered Plectro- poma annulatum Gunther, 1859, as the type species).
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Scopularia de Buen, 1959:95 (type species Scopularia rubra de Buen, 1959 [=Plec- tropoma semicinctum Valenciennes, 1833], by original designation).
Diagnosis. —A genus of anthiine serranid fishes characterized by the following: 27 or 28 vertebrae (usually 27, very rarely 26), three predorsal bones, 17 principal caudal- fin rays (15 branched), one to three antrorse spines on preopercle, supramaxilla typically present, and maxilla without scales.
Gender. —Generic names such as Hypo- plectrodes, with the suffix “‘“—odes,”’ are sub- stantivated adjectives and are masculine (ICZN, Article 30b). Accordingly, adjecti- val specific names in combination with Hy- poplectrodes must have the masculine ter- mination (ICZN, Article 31b).
Species of Hypoplectrodes.—Allen & Moyer (1980:329) recognized six species in the genus Ellerkeldia (herein considered as species of Hypoplectrodes), presented a key for their identification, and stated that they ‘“‘are confined to shallow temperate seas of
New Zealand and southern Australia.” They —
overlooked the type species of the genus E/- lerkeldia, Plectropoma semicinctum (=H. semicinctum), from the eastern Pacific, pre- sumably because Whitley (1927) mistak- enly considered Plectropoma annulatum Gunther, 1859, as the type species.
In addition to H. semicinctum and the type species of Hypoplectrodes, H. nigro- ruber, from waters off Australia, the other nominal species of the genus are: H. an- nulatus (Gunther, 1859), H. huntii (Hector, 1875), H. jamesoni Ogilby, 1908, H. mac- cullochi (Whitley, 1929), H. ruber (Allen, 1976), which is in need of a replacement name because it is a junior secondary hom- onym of Scopularia rubra de Buen, 1959 [=H. semicinctum], and H. wilsoni (Allen & Moyer, 1980). John R. Paxton informed us (in litt., 23 Sep 1987) that he and Gerald R. Allen are in the process of revising E/- lerkeldia (=~Hypoplectrodes) and that they recognize two undescribed species of that
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genus from eastern Australia and New Zea- land.
Relationships of Hypoplectrodes. — Ran- dall (1980:102) considered Ellerkeldia (=Hypoplectrodes) to be “‘closely related to Plectranthias’’ Bleeker, 1873, and gave characters for separating the two genera. Heemstra & Anderson (1983) pointed out that Randall’s characters would not distin- guish these genera, but suggested that ver- tebral number might be useful (Plectran- thias with 26 vertebrae, Ellerkeldia with 27). Doubt about the utility of vertebral number in distinguishing the genera is cast by the discovery of a new species of Plectranthias (P. bilaticlavia) from the Kermadec Islands off northern New Zealand by Paulin & Rob- erts (1987). The holotype of their new species has 27 vertebrae; the two paratypes (and only other specimens known) each have 26. Radiographs of the types of P. bilaticlavia revealed no indications of fusions or de- formities of the vertebral columns. It is pos- sible that a count of 27 vertebrae is rare for this species.
In order to evaluate the relationship of Hypoplectrodes with Plectranthias, detailed comparative studies of the species of the two genera are needed. This will be a for- midable task because there are 10 species (8 with names, two undescribed) of Hypo- plectrodes (see previous section) and 37 species of Plectranthias (Randall 1980, Fourmanoir & Rivaton 1980, Katayama & Masuda 1980, Fourmanoir 1982, Raj & Seeto 1983, Heemstra & Anderson 1983, Paulin & Roberts 1987). Further study will likely lead to the recognition at the generic level of one or more of the eight genera subsumed by Randall (1980) into Plectran- thias. One of those yet to be resurrected genera is a logical candidate for recognition as the sister genus of Hypoplectrodes. A more precise estimation of these generic relation- ships is beyond the scope of this work.
In an attempt to provide a character anal- ysis for Hypoplectrodes we consider other anthiines as the first outgroup, other ser-
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ranids (serranines plus epinephelines) as the second, and other percoids as the third. In this analysis the only character whose states we feel confident in polarizing is vertebral number. Species of Hypoplectrodes have 27 or 28 vertebrae (one of 33 specimens of H. maccullochi examined with only 26; see Heemstra & Anderson 1983, and the ge- neric diagnosis); other anthiines usually have 26 (one of three known specimens of Plec- tranthias bilaticlavia with 27, see above; Gi- ganthias immaculatus Katayama, 1954, if indeed it is an anthiine, with 25; Boulenger 1895; Katayama 1959, 1960; Gosline 1966; Anderson & Heemstra 1980; Heemstra & Anderson 1983; Johnson 1983; our unpub- lished data). Other members of the Serran- idae (serranines and epinephelines) almost always have 24 vertebrae (Niphon, a prim- itive epinepheline, with 30; Pseudogramma with 26, Suttonia with 26 or 27, Aporops with 27 or 28—these last three genera being highly derived grammistin epinephelines; Boulenger 1895; Katayama 1959, 1960; Gosline 1966; Johnson 1983; Leis & Rennis 1983; Carole C. Baldwin, pers. comm.). Forty-five of the 91 groups of percoids listed by Johnson (1984, Table 120) have 24 or 25 vertebrae lending support to Gosline’s (1968, 1971) assertion that “‘the basal per- coid number” is 24 or 25. In view of the preceding we interpret 24 or 25 as the most primitive character state for vertebral num- ber in the Serranidae and 26, 27, and 28 as progressively more derived states. Accord- ingly, then, we consider the number of ver- tebrae (27 or 28) as a synapomorphy uniting the species of Hypoplectrodes.
Hypoplectrodes nigroruber and HT, semicinctum
Because Hypoplectrodes nigroruber and H. semicinctum are very similar, it is ap- propriate to characterize those two species under a single heading and then to elaborate as necessary under the respective species ac- counts.
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Single dorsal fin (not divided to base be- tween spinous- and soft-rayed parts). Mar- gin of anal fin broadly rounded to squared off posteriorly. Second spine of anal fin more robust than first or third, considerably lon- ger than first, usually slightly longer than third. Pectoral fin symmetrical, middle rays longest; dorsalmost ray unbranched, the others usually branched. Pelvic-fin rays I, 5; pelvic fin inserted at vertical from base of pectoral fin, falling short of anal fin. Cau- dal fin truncate; principal rays 9 + 8; branched rays 8 + 7. Procurrent spur (John- son 1975) absent. Parhypural and five au- togenous hypurals present; epurals three. No dorsal trisegmental pterygiophores. For- mula for predorsal bones, anterior neural spines, and anterior dorsal pterygiophores 0/0+0/2/1+1/.
Scales ctenoid, resembling those of ser- ranine serranids (i.e., with rows of ctenial bases [Hughes 1981] present proximal to marginal cteni); no secondary squamation. Most of head covered with scales; dorsum and lateral aspect of snout, maxilla, supra- maxilla, lower jaw, membranes between branchiostegals, and most of branchioste- gals without scales; gular region usually without scales; squamation variously de- veloped on interopercle, but usually con- fined to posterior part. No axillary process at base of pelvic fin. Squamation well de- veloped on bases of all fins and continuing for some distance onto fins. Lateral line complete, extending to at least base of cau- dal fin (running parallel to dorsal body con- tour below dorsal fin, curving to near mid- lateral axis of body on caudal peduncle).
Supramaxilla present. Premaxillae pro- trusile. Posterodorsal border of maxilla not covered by elements of circumorbital series when mouth closed. Mouth terminal. Pos- terior margin of preopercle serrate; one to three antrorse spines on preopercle (one spine usually at angle or on ventral margin near angle, other spine(s) on ventral mar- gin). Posterior margin of bony opercle with three spinous processes, middle one best de-
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1005
Fig. 1.
veloped. Distal margins of interopercle and subopercle usually smooth, occasionally with a few serrae or slightly roughened. On each side of snout, two closely set nares near eye. Snout usually longer than diameter of orbit. Diameter of bony orbit considerably greater than bony interorbital width. Bran- chiostegals seven. Gill arches four, with a slit behind fourth. Well developed gillrakers rather short (longest gillrakers usually short- er than longest gill filaments), anterior low- er-limb rakers and most of upper-limb rak- ers rudimentary. Vomer and palatines with teeth: vomerine tooth patch chevron shaped, without a backward prolongation; palatine teeth in a longitudinal band. No teeth on tongue or pterygoids.
Hypoplectrodes nigroruber (Cuvier, 1828) Figs. 1, 2; Tables 1-5
Plectropoma nigrorubrum Cuvier, 1828:402 (original description; lectotype, herein designated, MNHN 7776, 189 mm SL; type locality Port du Roi Georges [=King George Sound, Western Australia}).
Diagnosis.—This species appears to be distinguishable from all other species of Hy- poplectrodes in morphology of the lateral-
Lectotype of Plectropoma nigrorubrum, MNHN 7776, 189 mm SL; Western Australia.
line scales (lateral-line tubes reaching pos- terior borders of scales; tubes of anterior lateral-line scales highly branched, becom- ing less so posteriorly, tubes of posterior- most scales bifurcate or unbranched) and in having an area of very small scales (on dor- sum and dorsolateral part of body dorsal to lateral line) beginning at anterior end of dor- sal fin and extending anteriorly to become continuous with scaly regions of head. Pos- teroventral corner of maxilla usually with- out prominent extension. Ventral margin of preopercle with one to three, usually two, antrorse spines; spines sometimes covered by skin. Vertebrae usually 27 (10 precaudal + 17 caudal), occasionally 28 (10 + 18). Pleural ribs on vertebrae 3 through 10 (3- 11 in one of 15 specimens). Dorsal fin rays X, 16 to 18. Anal fin rays III, 8. Pectoral- fin rays 13 to 15, usually 14. Gillrakers, in- cluding rudiments, on first gill arch 5 or 6 + 12 to 16—total 17 to 22; developed gill- rakers on lower limb 5 to 7. Tubed lateral- line scales 55 to 65, most frequently 57 to 63. Scales from anal-fin origin to lateral line 19 to 23 (counted posterodorsally), 23 to 28 (counted anterodorsally). Scales on cheek quite small; rows of cheek scales very dif- ficult to count; number of cheek scale rows
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Fig. 2. Hypoplectrodes nigroruber, AMNH 31307, 114 mm SL; Western Australia.
ca. 21 to ca. 29, usually ca. 22 to ca. 26. Pseudobranch with 26 to 36 filaments, tend- ing to increase in number with increase in SL. Length of second anal spine 10.3 to 15.1% SL. Body encircled by four darkly pigmented bands.
Description. —Characters included in the combined description of H. nigroruber and H. semicinctum and those presented in the species diagnosis form part of the species description. Frequency distributions for a number of meristic traits are in Tables | to 4; morphometric data appear in Table 5.
Procurrent caudal-fin rays 8 to 10 (usually 8) dorsally, 6 to 9 (usually 8) ventrally. Epi- pleural ribs associated with first 9 or 10 ver- tebrae (infrequently with 10th). Anal triseg- mental pterygiophores 0 to 5 (most frequently 4). Rows of scales between lateral line and mid-base of spinous dorsal fin 3 or
4 (usually 3). Scales from dorsal-fin origin to lateral line 4 to 7 (usually 5 or 6). Circum- caudal-peduncle scales 30 to 34 (most fre- quently 30 or 31).
Depth of body (at origin of dorsal fin) 2.80 to 3.15, length of head 2.25 to 2.45 in SL. Horizontal diameter of bony orbit 4.40 to 6.70 in length of head, 1.10 to 1.95 in length of snout. Bony interorbital width 13.20 to 19.45 in length of head, 2.30 to 4.40 in di- ameter of bony orbit. Lower jaw exceeding upper when mouth closed. Maxilla reaching vertical through posterior part of orbit to slightly beyond orbit. Anterior naris at dis- tal end of short tube; posterior border of tube produced into a flap which reaches or falls just short of posterior naris when re- flected. Premaxilla with wide band of small conical teeth; band narrower posteriorly; posterior teeth at anterior end of band (near
Table 1.—Frequency distributions of numbers of fin rays in two species of Hypoplectrodes. Separate counts from both left and right pectoral fins included. Counts of name-bearing types are indicated by asterisks. Pectoral-fin rays
Dorsal soft rays Anal soft rays
Species 16 7 IS IO Oe or x 7T wR 97 iat aS RST a6 i7 ~* 18 x H. nigroruber Digat ee Wey 16* |e aa | 14.00 H. semicinctum 67 WS 416" ol 2020'S Gall 1 16% 57, 2. tows
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VOLUME 102, NUMBER 4
nn ne LEE SEass ssn L TST
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Aq poieorput oie sodA} 3utreoq-oweu Jo sjuNOD ‘sapo1joajdoda] Jo soroods OM} Ul YOIe [13 ISIY UO SIOYeIT[IS JO S19quINU JO SUOTINGLYSIp Aouonbelj—"Z 3qeL
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Table 3.—Frequency distributions of numbers of tubed lateral-line scales in two species of Hypoplectrodes.
Counts of name-bearing types are indicated by asterisks.
Species 48 49 50 Si. 52> 53 54) 55 S657 SE 58 60 61 62 63 64 65 x H. nigroruber Ll = 2 1 4. =] 4 Bot] ieee H. semicinctum Ss: & 202" 8 2 50.16
symphysis) larger and recurved; usually one to a few teeth at anterior end of jaw enlarged into small canines; no teeth at symphysis. Dentary with band of small conical teeth; band considerably widened near symphysis; one to a few small canines present about one-third way from symphysis to posterior end of band; teeth at anterior end of band near symphysis mostly recurved—more posterior ones somewhat enlarged; one to a few small canines at anterior end of jaw; no teeth at symphysis. Villiform to small con- ical teeth on vomer and palatines. Fourth, fifth, or sixth (usually fourth or fifth) dorsal spine longest. First anal spine 1.50 to 2.05 in second anal spine. Pectoral fin reaching vertical through anterior part of anal fin or pectoral fin appreciably shorter. Coloration. —In alcohol head mottled dorsally and laterally, lighter ventrally. Dark bar just anterior to dorsal fin descending from dorsal contour to terminate where opercle joins body dorsally. Rather weakly pigmented bar ventral to anterior part of spinous dorsal fin which descends for vari- able distances ventrally. Four darkly pig- mented bands, usually narrower than lightly pigmented interspaces, encircling body: an- teriormost ventral to middle of spinous dor- sal fin, second extending from anterior end of soft dorsal fin to anterior end of anal fin, third ventral to posterior end of soft dorsal
fin, posteriormost on caudal peduncle; area in anteriormost band just ventral to lateral line more heavily pigmented. Fins without distinctive pigmentation except where weakly pigmented bar and three anterior- most bands encroach upon dorsal and anal fins. Cuvier (1828) wrote that the body is a very vivid red-orange and is crossed by five black bands: the first is faint and originates beneath the first rays of the dorsal; the other four are very dark; the last encircles the base of the tail. Castelnau (1875:8) stated that this species has “five broad transverse black bands on a reddish ground colour.”
Distribution. —We have examined speci- mens collected in Pacific and Indian ocean waters of Australia off New South Wales, Victoria, South Australia, and Western Australia. Scott (1979) reported two speci- mens from Tasmanian waters (one from off the northeastern coast in Banks Strait; the other off the southwestern coast, off Port Davey). Depths of capture are available for only a few collections; they range from 3.5 to 15 m.
Gender. —The only published spelling that we have seen for the specific name in com- bination with Hypoplectrodes is nigroru- brum. The compound nigrorubrum is an adjective and in association with Hypoplec- trodes (which is masculine, see section on gender of Hypoplectrodes) must have the
Table 4.— Frequency distributions of numbers of pseudobranchial filaments in two species of Hypoplectrodes.
Counts of name-bearing types are indicated by asterisks.
Species 145 505 16 ids 18. 897-20) 21,922 H. nigroruber H. semicinctum 2. =) = FHF Ae aassysS
26 27 28 29. 30.31 32. 33° 342733
23 1 =| Se
VOLUME 102, NUMBER 4 1009
Table 5.—Data on morphometric characters for two species of Hypoplectrodes. Standard lengths are in mm; other measurements, in percentages of standard length.
H. nigroruber H. semicinctum
Character n Range n Range
Standard length Is 84.8-203 17 77.9-177 Head, length 15 40.4-44.3 17 38.6-45.1 Snout, length 14 9.4-12.7 17 9.6-13.4 Orbit, diameter 15 6.6—9.3 iF 6.5-10.0 Postorbital length of head 15 21.0-26.8 j§ 2132357 Upper jaw, length 15 16.1-19.3 17 17.7-19.8 Maxilla, width 15 4.9-6.4 17 5.5-6.7 Interorbital width 15 2.1-3.3 7 2.9-3.8 Body, depth at origin of dorsal fin iS 32.0-35.5 he 32.9-37.1 Predorsal length 15 37.4-41.0 17 38.2-43.9 Preanal length 15 62.6-73.1 iV 65.4-73.0 Caudal peduncle, length 15 18.8-22.0 i 9/ 18.7—21.7 Caudal peduncle, depth 15 11.0-13.0 17 10.0—12.4 Pectoral fin, length 15 25:I=31-5 Iba 27.0-34.2 Pelvic fin, length 15 19.7-22.8 V7 21.0-25.8 Anal fin, depressed length 15 25.4—30.9 17 28.7—32.2 Upper caudal-fin lobe, length 1 19.2-25.0 16 20.9-25.3 Lower caudal-fin lobe, length 14 19.6—25.0 17 20.9-25.7 Third dorsal spine, length 13 12.3->15.4 Ly 11.9-> 14.6 Fourth dorsal spine, length 15 13.9-17.2 17 13.1-17.3 Longest dorsal spine, length 15 13.9-18.2 16 13.5-17.6 First anal spine, length 15 5.8-8.3 16 7.3-9.8 Second anal spine, length 15 10.3-15.1 i) 14.6-19.7 Third anal spine, length 14 9.5-14.0 16 123-771
masculine termination (ICZN, Article 31b); consequently the correct binomen is Hy- poplectrodes nigroruber.
Remarks.—Through the courtesy of M. L. Bauchot we have examined the two syn- types (MNHN 7776) of Plectropoma nigro- rubrum. Both are in poor condition, but the larger is in a better state of preservation. We hereby designate as the lectotype of Plec- tropoma nigrorubrum Cuvier, 1828, the syntype of 189 mm SL, which retains MNHN 7776 as its catalog number; the paralectotype (142 mm SL) has been as- signed a new number (MNHN 1988-799).
Material examined. —Sixteen specimens, 85 to 203 mm SL.
Lectotype: MNHN 7776 (189 mm SL); King George Sound, Western Australia; J. Quoy & P. Gaimard.
Paralectotype: MNHN 1988-799 (142 mm SL); same data as for lectotype.
Other material: USNM 42015 (one spec- imen, 193 mm SL), Port Jackson, New South Wales; USNM 42019 (1, 198), Port Jack- son, New South Wales; CAS-SU 9189 (1, 203), Maroubra, New South Wales; CAS- SU 20797 (1, 199), Port Hacking, New South Wales; NMV A2554 (1, 174), Cape Wel- lington, Wilson’s Promontory, Victoria, 39°4.1'S, 146°28.6'’E,<10 m, R. Kuiter and M. McDonald, 9 Feb 1982; NMV A2588 (1, 163), western shore of Brown Head, Wil- son's’ Promontory; |’Victoria,: 39°2:7’S, 146°28.3’E, 15 m, T. Cochrane, R. Kuiter, and M. Larsen, 9 Feb 1982: NMV A3007 (1, 134), northern shore of Horn Point, Wil- son’s Promontory, Victoria, 39°1.6’S, 146°28.2’E, <10 m, R. Kuiter and M.
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Fig. 3. Holotype of Plectropoma semicinctum, MNHN 7777, 146 mm SL; Juan Fernandez Islands.
McDonald, 9 Feb 1982; USNM 177114 (1, 157), around Kangaroo Island and St. Ste- phens Bay, South Australia, Howard, Mar-— Apr 1952; NMV A289 (1, 86), Cape Cas- sini, Kangaroo Island, South Australia, 35°35’S, 137°19’E, W. Gosline and J. Glov- er, 17 Aug 1966; AMNH 31307 (4, 85-128), northeast and southwest sides of North Point, south of Boulder Hill, Western Aus- tralia, ca. 34°56’S, ca. 118°13’E, 3.5 m, Nel- son, Butler, and Rosen, 14 Mar 1969; NMV A5061 (1, 167), Champion Bay, Western Australia, 28°46’S, 114°36’E.
Hypoplectrodes semicinctum (Valenciennes, 1833) Figs. 3-5; Tables 1-6
Plectropoma semicinctum Valenciennes, 1833:442 (original description; holotype MNHN 7777, 146 mm SL; type locality Juan Fernandez Islands, eastern Pacific Ocean).
Scopularia rubra de Buen, 1959:95 (original description and illustration; holotype EBMC 123-124,174 mm TL, apparently
lost; type locality Cumberland Bay, Mas a Tierra Island, Juan Fernandez Islands, eastern Pacific Ocean).
Diagnosis.—Lateral-line tubes reaching posterior borders of scales; tubes of anterior lateral-line scales bifurcate; those of poste- rior scales unbranched. Scales on body an- terior to dorsal fin not greatly reduced in size (except one of 38 specimens with small area of reduced scales adjacent to anterior end of dorsal fin). Posteroventral corner of maxilla usually with prominent extension. Ventral margin of preopercle with two or three, usually three, antrorse spines; spines frequently covered by skin. Vertebrae 27 (10 precaudal + 17 caudal). Pleural ribs on vertebrae 3 through 10. Dorsal fin rays X, 19 to 22. Anal fin rays III, seven to nine (usually eight). Pectoral-fin rays 15 to 18 (usually 16 or 17). Gillrakers, including ru- diments, on first gill arch 4 to 6 + 12 to 14—total 17 to 20; developed gillrakers on lower limb 8 or 9. Tubed lateral-line scales 48 to 55, most frequently 48 to 51. Scales from anal-fin origin to lateral line 16 to 20
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Fig. 4. Hypoplectrodes semicinctum, MCZ 46165, 108 mm SL; Juan Fernandez Islands.
(counted posterodorsally), 19 to 24 (count- ed anterodorsally). Rows of cheek scales 14 to 19. Pseudobranch with 14 to 28 fila- ments, tending to increase in number with increase in SL. Length of second anal spine 14.6 to 19.7% SL. Body usually with 9 dark- ly pigmented bars (including one on nape); bars wider than lightly pigmented inter- spaces.
Description. —Characters included in the combined description of H. nigroruber and
H. semicinctum and those presented in the species diagnosis form part of the species description. Frequency distributions for a number of meristic traits are in Tables | to 4; morphometric data appear in Table 5. Procurrent caudal-fin rays 8 to 10 (very rarely 10) dorsally, 6 to 9 (usually 7 or 8) ventrally. Epipleural ribs associated with first 9 or 10 vertebrae (usually first 9). Anal trisegmental pterygiophores 0 to 3 (most frequently 1). Rows of scales between lateral
Fig.-5. Islands.
Holotype of Scopularia rubra, EBMC 123-124, 174 mm TL (from de Buen, 1959); Juan Fernandez
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Table 6.—Comparisons of data on Hypoplectrodes semicinctum and data from de Buen (1959) on holotype and paratype of Scopularia rubra (EBMC 123-124). Standard lengths and total lengths are in mm; other mea- surements in percentages of standard length (% SL) or percentages of head length (% HL). L = left; R = right; ruds. = rudimentary gillrakers; est. = standard lengths estimated (see text); > = slight damage to caudal fin.
H. semicinctum S. rubra Character Range Holotype Holotype and paratype Dorsal-fin rays X, 19-22 X, 19 X, 20 Anal-fin rays III, 7-9 III, 8 III, 8 Pectoral-fin rays 15-18 16 16 Gillrakers: Total 17-20 20 16 Upper limb 4-6 6 5 (1+3-5 ruds.) (1+5 ruds.) (1+4 ruds.) Lower limb 12-14 14 11 (8 or 9+ 4-6 ruds.) (9+5 ruds.) (8+3 ruds.) Lateral-line scales 48-55 50 (L), 49 (R) — Scales in longitudinal line (see text) — ca. 50 (L), ca. 53 (R) 49-52 Scales above lateral line 5-7 6 5 or 6 Scales below lateral line (see text) 16-20 ca. 16 14 or 15 Standard length 77.9-177 146 143 and 107 (est.) Total length 97.3-214 >179 174 and 131 % SL % SL % SL Head, length 38.6—-45.1 38.6 39.3-41.5 Body, depth (greatest) 33.3-38.8 3357 34.5-34.9 Body, width (greatest) 15.3-20.5 15.9 18.6-18.8 Predorsal length 38.2-43.9 38.2 40.0-41.5 Preanal length 65.4—73.0 (AP 72.4-76.4 Prepelvic length 37.5-45.8 44.9 46.2-52.8 Dorsal-fin base 51.9-58.1 53.8 50.0—-53.8 % HL % HL % HL Snout, length 23.1-31.1 27.8 34.0-36.5 Orbit, diameter 14.9-24.0 19.9 17.5—22.7 Postorbital length of head 50.9-55.6 55.1 50.0-52.2 Interorbital width 6.8—9.6 9.6 11.3-14.0 Caudal peduncle, depth 23.2—-29.8 27.0 29.7-29.8 Pectoral-fin base, width 16.8—21.3 202 20.4—22.8 Pectoral fin, length 65.7—80.6 70.9 63.6-64.9 Pelvic fin, length 50.4-61.8 56.4 43.8-50.0
line and mid-base of spinous dorsal fin 3 or 4 (most frequently 4). Scales from dorsal- fin origin to lateral line 5 to 7 (most fre- quently 6). Circum-caudal-peduncle scales 27 to 32 (usually 28 to 30).
Depth of body (at origin of dorsal fin) 2.70 to 3.05, length of head 2.20 to 2.60 in SL. Horizontal diameter of bony orbit 4.15 to 6.75 in length of head, 0.95 to 2.05 in length
of snout. Bony interorbital width 10.45 to 14.75 in length of head, 1.70 to 3.30 in di- ameter of bony orbit. Jaws nearly equal or lower jaw exceeding upper when mouth closed. Maxilla usually falling short of ver- tical through posterior margin of orbit. An- terior naris at distal end of tube; posterior border of tube elongated slightly, but falling short of posterior naris when reflected. Pre-
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maxilla with band of small conical teeth; band expanded anteriorly; posterior teeth in expanded part of band (adjacent to sym- physis) enlarged and posteriorly directed; one or two canines at anterior end of jaw; no teeth at symphysis. Dentary with band of small conical teeth; band somewhat ex- panded adjacent to symphysis; one to three canines at about middle of band; numerous enlarged posteriorly directed conical teeth at anterior end of band near symphysis; one or two canine teeth (may be exserted) at anterior end of jaw; no teeth at symphysis. Small conical teeth on vomer and palatines. Fourth, fifth, or sixth (usually fifth) dorsal spine longest. First anal spine 1.75 to 2.25 in second anal spine. Pectoral fin usually reaching vertical through anterior part of anal fin.
Coloration. —In alcohol dorsum of head mostly darkly mottled; cheek and opercular series with several mostly horizontal stripes, narrower than lighter interspaces. Body usually with nine darkly pigmented bars; bars evenly spaced, wider than lightly pig- mented interspaces; anteriormost bar on nape (saddle-like, extending over dorsum to join bar from other side); second bar begin- ning on nape and beneath anterior part of spinous dorsal fin; third through fifth bars beneath spinous dorsal fin; sixth through eighth bars beneath soft dorsal fin; eighth bar also extending onto and over caudal pe- duncle to become continuous with corre- sponding bar from other side; eighth bar surrounding small lightly pigmented area dorsally just posterior to base of soft dorsal fin; ninth bar on caudal peduncle; second through fifth bars usually extending about 60 to 70% of distance from dorsum to ven- tral midline (on specimens more than ca. 100 mm SL; on smaller specimens these bars may extend further ventrally); on many specimens sixth through eighth bars becom- ing very narrow ventrally, sometimes reach- ing anal fin or ventral border of caudal pe- duncle (eighth); very frequently eighth and ninth bars becoming narrowly confluent with
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corresponding bars from other side; bars frequently showing various anastomoses, often bars three and four, four and five, and six and seven uniting broadly. Fins mostly straw colored except where dark bars extend onto dorsal and anal fins.
Valenciennes (1833) described the col- oration of the holotype of Plectropoma semicinctum. He wrote that the colors of this fish are a beautiful vermilion red, tra- versed by eight half bands of a bright red brown, that descend on the back and stop on the middle of the sides, so as to form half belts on the sides. Only the last almost encircles the entire tail. Some paler and oblique brown bars cross the cheeks, and form on the opercle indistinct rivulations. The dorsal and caudal are reddish. The pec- torals, ventrals, and anal are olive, mixed with the red that forms the general back- ground color. De Buen (1959) stated that Scopularia rubra is red with black bands.
Distribution. —We have examined speci- mens of H. semicinctum collected in the eastern South Pacific off the Juan Fernandez Islands and San Félix Island in shallow waters with a maximum depth of 20 m. Yanez-Arancibia (1975) illustrated a spec- imen identified as Scopularia rubra that was collected at Easter Island. This drawing is a good representation of H. semicinctum; accordingly, then, it would appear that H. semicinctum can be considered as reliably reported from Easter Island. Randall & Cea Egana (1984) included Ellerkeldia rubra (de Buen), based on Yanez-Arancibia’s (1975) report of Scopularia rubra, in their paper on native names of Easter Island fishes. Randall has not observed or collected H. semicinctum at Easter Island, despite the fact that he has collected fishes extensively there on three separate occasions, and he has not met any fishermen or divers there who are familiar with this species (J. E. Ran- dall, pers. comm.). Consequently, Randall believes that there is no breeding population of H. semicinctum at Easter Island (at least not in shallow water) and that Yanez-Aran-
1014
cibia’s report of a specimen from Easter Is- land was probably of a stray or possibly of a specimen for which the locality was in- correctly recorded.
Orthography.—The correct termination for the specific name semicinctum is debat- able. Valenciennes (1833) proposed the name in the genus Plectropoma. The sufhx ‘“‘poma”’ 1s a neuter Greek noun, whereas the suffix “‘cinctum”’ is either a neuter Latin noun or a verbal adjective, the perfect pas- sive participle of the Latin verb “‘cingo.”’ If a species-group name is a noun in apposi- tion, it keeps the same termination without regard to the gender of the generic name with which it is associated (ICZN, Article 31b[1i]), but a species-group name that ends in a Latin participle in the nominative sin- gular ““must agree in gender with the generic name with which it is at any time combined, and its termination must be changed ac- cording to Latin inflection” (ICZN, Article 31b).
It can be argued that Valenciennes (1833) did not indicate whether he meant semi- cinctum to be a verbal adjective or a noun in apposition to Plectropoma because there is nothing in the original description per se to show his intent. Jordan (1891) described the genus Gi/bertia and designated Plectro- poma semicinctum as the type species. Fur- ther on he used the binomen Gilbertia semi- cincta and gave the etymology of semicincta as “‘semi’’—half, “‘cinctus’’—belted, indi- cating that he considered the second part of the compound to be a participle. The com- bination Gilbertia semicincta has been used by a number of other authors (including Boulenger 1895, Rendahl 1921, de Buen 1959, Sepulveda Vidal & Pequeno 1985). Bauchot et al. (1984) used the binomen E/- lerkeldia semicincta—semicincta agreeing in gender with Ellerkeldia. It appears that the evidence of usage could be considered as decisive in the sense of the Code (ICZN, Article 31b[1i]), and that the specific name is a verbal adjective (spelled semicinctus in combination with Hypoplectrodes).
On the other hand it can be asserted that
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Valenciennes did intend semicinctum as a noun, because it is essentially the Latin equivalent of the last part of the French vernacular name, Le PLECTROPOME A DEMI-CEINTURES (=the plectropome with half girdles), which precedes the orig- inal description. Support for this view is given by the facts that the very next species described by Valenciennes (1833), Meso- prion isodon, is preceded by the French name Le MESOPRION A DENTS EGALES, that Plectropoma nigrorubrum, described by Cu- vier (1828), is preceded by Le PLECTRO- POME ROUGE ET NOIR, and that in each of these cases the specific name is a trans- lation of the last part of the French name and the same part of speech (nouns in the first instance, adjectives in the second). We prefer this latter interpretation—that Valen- ciennes did indeed indicate that he regarded the name semicinctum as a noun, and con- sider the correct binomen for this species to be Hypoplectrodes semicinctum.
Remarks.—As mentioned in the intro- duction, the junior author was convinced some years ago after comparing the original descriptions that Plectropoma semicinctum and Scopularia rubra are synonymous. Be- cause of a few discrepancies between Valen- ciennes’ (1833) description of P. semicinc- tum and de Buen’s (1959) description of S. rubra, the senior author disagreed. Valenci- ennes described the presence of three strong antrorse spines on the lower limb of the preopercle and gave the anal- and pectoral- fin ray counts as III, 7 and 15, respectively; in contrast, de Buen did not mention the presence of any preopercular spines (al- though he recorded the occurrence of serrae on the upper limb of the preopercle) and gave the anal- and pectoral-fin ray counts as III, 8 and 16, respectively. In de Buen’s illustration of the holotype of S. rubra (see Fig. 5) the upper limb of the preopercle is serrate, but the lower limb is smooth. (Al- though de Buen mentioned H. semicinctum, as Gilbertia semicincta, in a list near the beginning of his paper, he did not compare it with S. rubra.)
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Despite a number of attempts over a pe- riod of more than 15 years, we have been unable to find de Buen’s type material of S. rubra; the types are apparently lost. How- ever, we have examined the holotype of P. semicinctum and find that the discrepancies noted above between the two original de- scriptions can be easily resolved. Valen- ciennes’ (1833) counts of III, 7 (anal-fin rays) and 15 (pectoral-fin rays) are in error. The holotype of P. semicinctum has an anal-fin ray count of III, 8 and pectoral-fin ray count of 16 (in each fin). (Valenciennes was also inaccurate when he recorded the dorsal-fin
ray count of the holotype of P. semicinctum
as X, 20; the correct count is X, 19. This difference is probably the result of counting the last soft ray, which is split to the base, as two elements rather than as one.) Fre- quently in specimens of species of Hypo- plectrodes the antrorse spines on the ventral margin of the preopercle are covered by skin and easily overlooked, despite the fact that they are typically well developed. In view of the overall close similarity between spec- imens of H. semicinctum and de Buen’s de- scription of S. rubra (see Table 6), it is rea- sonable to assume that the preopercular spines on de Buen’s specimens were ob- scured by skin.
In Table 6 data taken by us on specimens of H. semicinctum are compared with those given by de Buen on the holotype and para- type of S. rubra. De Buen gave total lengths, but did not give standard lengths for his material. We have estimated the standard lengths of his specimens based on our mea- surements of total and standard lengths of 15 specimens of H. semicinctum (SL =a + b [TL], where a = —5.2094, b = 0.8541, r = 0.9995). The meristic data are in close agreement; with the exception of two char- acters (gillrakers and scales below the lateral line) de Buen’s counts fall within the ranges we obtained for H. semicinctum, and de Buen’s counts for those two characters are just outside our ranges. De Buen’s ranges for several morphometric characters fall outside our ranges. Because our morpho-
1015
metric data are based on a relatively small number of specimens (16 or 17), de Buen’s ranges may be reasonable extensions of ours. Alternatively, in some cases de Buen’s methods of measuring may have been dif- ferent from ours or we may have misinter- preted his methods (see section on abbre- viations and methods), perhaps as a result of not adequately translating his Spanish into English (although we had our transla- tion edited by Dr. José Escobar, Spanish faculty, College of Charleston). In any event we consider our lack of complete agreement with de Buen’s morphometric data to be relatively minor in view of the general sim- ilarity we find between de Buen’s descrip- tion of S. rubra and the specimens of H. semicinctum that we examined. The strik- ing resemblance between H. semicinctum and de Buen’s S. rubra can be seen by com- paring Figs. 3 and 4 with Fig. 5 and by comparing the colorations of the two nom- inal species as described by Valenciennes and de Buen. Accordingly, then, we consid- er Scopularia rubra de Buen, 1959, to be a junior synonym of Hypoplectrodes semi- cinctum (Valenciennes, 1833). (G. R. Allen and J. E. Randall, pers. comm., have ar- rived at the same conclusion regarding the synonymy of S. rubra and H. semicinctum.)
Material examined. — Thirty-eight speci- mens, 38-177 mm SL.
Holotype: MNHN 7777 (146 mm SL); Juan Fernandez Islands; C. Gay.
Other material: MCZ 4827 (two speci- mens, 134-141 mm SL), Juan Fernandez Islands, Hassler Expd., 1872; USNM 176414 (1, 142), Cumberland Bay, Juan Fernandez Islands, 33°38.0'S, 78°50’W, M. J. Lobell, 20 Feb 1945; SIO65-634 (17, 38- 160), Cumberland Bay, Juan Fernandez Is- lands, 33°38'20’S, 78°48'50’W, 6-11 m, W. Baldwin et al., 11 Dec 1965; MCZ 46165 (7, 78-177), West Bay, Mas a Tierra Island, Juan Fernandez Islands, 0-20 m, R/V An- ton Bruun, cr. XIII, coll. 15, Jan 1966; CAS 24143 (4, 88-115), data as for MCZ 46165; SI1065-624 (4, 86-157), San Félix Island, NW side, 26°17'30”’S, 80°05'40”W, 0-9 m,
1016
W. Baldwin et al., 5 Dec 1965; SIO65-628 (2, 106-156), locality as for SIO065-624, 0- 8 m, W. Baldwin et al., 6 Dec 1965.
Homonymy
Allen (1976) described Ellerkeldia rubra from Western Australia. As a result of our synonymizing Scopularia rubra with Hy- poplectrodes semicinctum, Ellerkeldia rubra Allen, 1976 (=Hypoplectrodes ruber) be- comes a junior secondary homonym of S. rubra de Buen, 1959. Gerald R. Allen and John E. Randall plan to propose a new name to replace Hypoplectrodes ruber.
Acknowledgments
M. L. Bauchot, D. Catania, M. Desoutter, W.N. Eschmeyer, M. N. Feinberg, M. F. Gomon, K. E. Hartel, S. L. Jewett, M. Nor- man, R. H. Rosenblatt, and H. J. Walker, Jr., allowed us to examine specimens in their care; W. I. Follett and G. C. Steyskal gave us advice on nomenclature; C. C. Baldwin, W. F. Hoffman, and J. F. McKinney pro- vided the radiographs used in this study; P. Coleman made the photograph for Fig. 5, and J. F. McKinney made those for Figs. 1 to 4; N. A. Chamberlain and J. Escobar im- proved our translations of passages in French and Spanish, respectively; G. D. Johnson sent us a copy of the pertinent part of the paper by Cossman; and F. Brigman typed the manuscript. G. R. Allen, C. C. Baldwin, W. F. Smith-Vaniz, and J. R. Paxton read the manuscript and made suggestions for improving it. This is GMBL contribution number 80.
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(WDA) Grice Marine Biological Labo- ratory, College of Charleston, 205 Fort Johnson, Charleston, South Carolina 29412; (PCH) J. L. B. Smith Institute of Ichthyol- ogy, Private Bag 1015, Grahamstown 6140, Republic of South Africa.
PROC. BIOL. SOC. WASH. 102(4), 1989, pp. 1018-1030
ACANTHEMBLEMARIA PAULA, A NEW DIMINUTIVE CHAENOPSID (PISCES: BLENNIOIDEI]) FROM BELIZE, WITH COMMENTS ON LIFE HISTORY
G. David Johnson and Edward B. Brothers
Abstract. —Acanthemblemaria paula is described from the barrier reef and outlying islands of Belize. The new species is the smallest member of the Chaenopsidae, reaching sexual maturity at 11.5 mm SL or smaller and attaining a maximum size of about 18 mm SL. In addition to its small size, it differs from all other Acanthemblemaria by its low number of dorsal-fin spines and distinctive head spination. Confusion of A. paula with A. spinosa led previous authors to erroneous conclusions about life history parameters of A. spinosa and larval recruitment in Acanthemblemaria.
In March 1987, we made shallow rote- none collections of fishes at the north end of Carrie Bow Cay, Belize, the site of a Smithsonian Institution field station since 1972. The purpose of these collections was to obtain otoliths for analysis of daily growth increments, and, thus, all specimens were fixed in 95% ethanol. Among the fishes col- lected were two tiny specimens of the chae- nopsid genus Acanthemblemaria Metzelaar that we were unable to identify using the most recent generic revision (Smith-Vaniz & Palacio 1974). The ethanol-fixed material was less than ideal for systematic study, but led us to examine the extensive holdings of Belize Acanthemblemaria at the Field Mu- seum of Natural History, collected and re- ported on by Greenfield & Johnson (1981). There, among 12 lots identified as A. spi- nosa, we found 50 additional specimens representing what we were then able to rec- ognize as a previously undescribed species. Returning to Carrie Bow in March 1988, we discovered that the new species is quite common on the reef flat and has habits sim- ilar to those of A. greenfieldi. The new species is a diminutive form, the smallest known member of the Chaenopsidae, and this may partially explain how it has gone unrecog-
nized, despite its residence in the environs of the very active Smithsonian research fa- cility. As currently known its geographic distribution is restricted to Belize, and spec- imens were not present in material exam- ined by Smith-Vaniz & Palacio (1974). In this paper we describe the new species and discuss its bearing on the conclusions of Greenfield & Greenfield (1982) concerning life history parameters of A. spinosa and habitat partitioning and larval recruitment in Acanthemblemaria.
Methods. —Counts and measurements largely follow the methods of Stephens (1963). Pore terminology is that of Smith- Vaniz & Palacio (1974), as modified by Johnson & Greenfield (1976) and Rosen- blatt & McCosker (1988). Measurements were made with an ocular micrometer. All lengths are standard length (SL) unless stat- ed otherwise. Vertebral and median fin ray counts were made from radiographs. Tooth and gill-raker counts were determined on cleared and stained specimens. Pore counts and head spine distribution were deter- mined from scanning electron micrographs and whole specimens. Institutional abbre- viations are as follows: ANSP, Academy of Natural Sciences of Philadelphia; FMNH,
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Table 1.—Frequency distributions of fin-ray and vertebral counts in three species of Acanthemblemaria.
Dorsal-fin spines
Dorsal-fin soft rays
Total dorsal-fin elements
Srecies, GEN 0, DL en a aS 4 CTS spinosa’ — — 24 111 If 209 4 71 53 spinosa — — 2 11 20:95" = 40; ..4 aspera) "— |i 59 53 8 206 = 5 29 aspera® — 3 19 1 — 199 — 8 14 aspera — — 6 13 — 20.77 — 2 14 paula 13 72 29 b= 19.2) =
Anal-fin soft rays
Species Soi) 2 2s soe) oS ox 12 spinosa! 1 10 ~68— -67 6 23.4 10 spinosa? — 3 8 Se) == 23.0 — aspera! — i?) SOME AS 3 11 aspera 5 18 — — — 21.8 aspera? — 3: 53 Say = 9230 — paula? — 6 60: 38 33, 23.4
Precaudal vertebrae
Species 11 12 13 x 28 29 spinosa! 81 4 11.0 50 ES aspera? (ey a WR Dn 25 paula? 81 fee 1251 14 59
' Data from Smith-Vaniz & Palacio (1974).
Caudal vertebrae
[ikea Mea GRONIan os sen 37 38 ox —- — — 14.5 1 8) 67 66° 9 — 35.5 — — — 143 — 1 9 4 — — 35.2 18 — — 15.8 — — 16 49 48 7 36.4 — — — 147 — 9 14 — — — 34.6 ——— ee £50) co 1D oe 35:7 34012. 1 16:8 —. =| 18 60°25 — 36:1
Pectoral-fin rays Segmented caudal-fin rays
13 14 x 11 12 13 14 iG 153 3 13.0 — — 81 — 13.0 14 — F330 — — 14 — 13.0 5 5 13:0 2 92 34 1 12:3 18 ] 13.0 _ 11 8 — 12.4 60 4 13.0 4 32° 36 _ 12.4
Total vertebrae
31 x 39 40 41 42 43 x 12 29:9 1 RS ea. -k2 41.4 9. 2907 Zh 30). 20 e420 29.1 ish 454 (27 41.1
? Data from Cartagena, Colombia population (Acero 1984).
3 Original data based on specimens from Belize.
Field Museum of Natural History; USNM, United States National Museum of Natural History.
Acanthemblemaria paula, new species Figs. 1, 2
Acanthemblemaria spinosa (in part, not of Metzelaar, 1919) Greenfield & Johnson, 1981.—Greenfield & Greenfield, 1982.
Diagnosis.—An Acanthemblemaria dif- fering from all other members of the genus by its small size (sexually mature at <12 mm, largest specimen 18.4 mm) and low number of dorsal-fin spines (18-21, X= 19.2) and further distinguished by the following combination of characters: supraorbital cir- rus complexly branched; head spines well developed, those on frontals extending pos- teriorly beyond orbits in a triangular patch
reaching about two-thirds distance to dor- sal-fin origin; anterior infraorbital with a few spines anteriorly, posterior infraorbital smooth.
Counts and measurements in mm of ho- lotype.—Dorsal fin XX, 17; anal fin II, 24; pectoral fin 13. Standard length 15.9; head length 3.4; head depth 2.0; upper-jaw length 1.4; orbit length 0.6; snout length 0.5; in- terorbital width 0.5; predorsal length 1.9; preanal length 6.8; caudal-peduncle length 0.8; caudal-peduncle depth 0.9; orbital cir- rus length 0.5; pectoral-fin length 3.3; lon- gest dorsal-fin spine length 1.6.
Description. —Frequency distributions of fin-ray and vertebral counts are given in Ta- ble 1. Dorsal-fin spines 18-21 (* = 19.2), soft rays 15-19 (X = 16.8); total elements 35-37 (x = 36.1). Anal fin II, 22-25 (% = 23.4). Pectoral-fin rays 12-14 (X = 13.0).
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VOLUME 102, NUMBER 4
Segmented caudal-fin rays 11-13 (¥ = 12.4). Pelvic-fin rays I, 3. Vertebrae: precaudal 12- 13 (¥ = 12.1); caudal 28-30 (x = 29.1); total 40-42 (x = 41.2).
Body very slender and elongate, compa- rable to that of Acanthemblemaria chaplini, the most elongate and slender species of the genus; head depth 7.3—8.0 in SL (7.8 in ho- lotype of A. chaplini, 5.7—7.2 in holotypes of remaining species); head length 4.5-5.0 in SL (3.0-4.4 in other species).
Cephalic sensory pores: mandibular 4; common |; preopercular 5; posttemporal 2— 3; lateral supratemporal 2—3; median su- pratemporal 1-2; supraorbital 1-3; poste- rior infraorbital 3; anterior infraorbital 3; frontal 0-1; median interorbital 1-3; ante- rior frontal 1.
Supraorbital cirrus complexly arbores- cent; main stalk usually comprising three palmate, multifid branches—a large central branch with two smaller branches arising proximally on either side of it; medial to main portion of cirrus, a smaller multifid branch arises at posterolateral margin of stalk base; cirrus generally longer in males (1-1.8 in eye) than in females (1.5—2.0 in eye). Anterior nostril on short tube, with posterior rim extended as multifid cirrus (length 1.2—2.2 in eye of males, 2.0-2.6 in eye of females).
Dorsal fin with fleshy flap on anterior margin of first spine. Membranes between adjacent spines and soft rays of dorsal fin not notably incised, continuous between tips of elements so that margin of fin is smooth; last spine slightly shorter than first soft ray, resulting in slight notch at junction of spi- nous and soft portions of fin. Membranes between adjacent rays of anal fin incised, extending posteriorly from tip of each ray of anterior margin of succeeding ray slightly proximal to tip, so that tip of each ray ap- pears slightly exerted; exerted tips of all rays curved posteriorly; condition exaggerated in males wherein tissue surrounding ray tips is notably thickened. Ultimate dorsal- and anal-fin rays connected along caudal pe- duncle by membrane that extends to distal
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tips of posteriormost dorsal and ventral procurrent caudal rays. Margin of caudal fin rounded. Pectoral fin large, rounded, with weakly scalloped margin; ventralmost three rays thicker than others and with tips slight- ly exerted, more so in males.
Vomerine teeth 9-13, arranged in ring, frequently with one tooth in center. Each palatine with two rows of teeth, 3-7 in outer row, 1-4 in inner row (one specimen has only 3 in single row on one palatine). Each premaxilla with outer row of 8—12 teeth, flattened and pointed at tips, becoming less so posteriorly as they also decrease in size, and inner band of smaller, curved, conical teeth. Anterior expanded portion of each dentary with outer row of 7 relatively in- cisiform teeth, similar in configuration to outer row on premaxillae, and inner band of smaller, robust, conical teeth that con- tinue as single row onto straight, posterior portion, where they become considerably larger. Gill rakers 3+5.
Anterior and dorsal areas of head orna- mented with anteriorly directed spines (Fig. 2). Supraorbital rim of each frontal bearing row of five to six spines that extends from posterodorsal corner of orbit anteriorly to junction with lateral ethmoid and nasal where a row of spines continues on each of these bones. Each lateral ethmoid bears two spines on anterior orbital rim. Each nasal bears four spines, a vertical row of three and one lateral to the ventralmost of these. Each lacrimal bears four to five spines along an- terior half of dorsolateral margin, these de- creasing substantially in size posteriorly. Dorsum of cranium (frontals) with a rough- ly diamond-shaped patch of 17-23 spines, its anterior apex extending just anterior to a point about midway between posterior or- bital rim and pupil, or slightly beyond, and its posterior apex reaching posteriorly about two-thirds distance between posterior or- bital rim and dorsal-fin origin.
Color in alcohol. —Background color- ation over head and body is uniformly pale cream to straw. Pigmentation is sparse and extremely variable among individuals. In
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addition to the intrinsic individual vari- ability, it appears that there is substantial loss of dark pigment with time in alcohol. The FMNH specimens (all in alcohol for at least ten years) exhibit only a sparse distri- bution of black pigment. Fresher specimens (USNM material, preserved for about one year) exhibit the dark black melanophores that characterize the older material, but also have large numbers of fine, purplish-red pigment cells. The latter are apparently lost with long-term storage in alcohol. Due to the marked differences, FMNH and USNM specimens are described separately. FMNH specimens: Posterior to the anus, the body surface is immaculate, although in some specimens a few internal clusters of melanophores can be detected along the vertebral column. Anterior to the anus, the amount and arrangement of external me- lanophores varies considerably; some in- dividuals appear almost completely devoid of pigment, and there is no consistent pat- tern of melanophores and no consistent sex- ual dichromatism. The following descrip- tion emphasizes the most consistent pigmentary features. There are frequently two sparse clusters of internal melano- phores in the ventrolateral area of the ab- dominal cavity just anterior to the anus; melanophores also may be scattered var- iously along the mid-ventral and ventrolat- eral surfaces of the anterior portion of the abdominal cavity, in the area below the pec- toral fin. The abdominal pigmentation fre- quently consists of two posterodorsally di- rected crescents, one just anterior to the anus and the other just ventral to the pectoral fin. Small individual melanophores occasion- ally occur on the dorsal half of the body surface, usually anterior to a vertical from the anus. Branchiostegal rays and mem- branes may be unpigmented, or covered with scattered or densely packed melanophores; ventrally, the anterior rays are more heavily pigmented, but in some specimens mela- nophores also occur at the dorsal tips of the most posterior rays, where they curve an-
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teriorly above the opercle. In occasional specimens, there are a few melanophores on the cheek just anterior to the angle of the preopercle. The iris is uniformly black, oth- erwise the head lacks pigment. The pectoral fin may be immaculate, have a sparse sprin- kling of melanophores, or have a more reg- ular arrangement of melanophores along the dorsal and ventral edges of many of the rays, the latter usually being denser on the medial side of the fin; in a few specimens the ventral portion of the pectoral-fin base is pigment- ed. The pelvic-fin base and rays range from immaculate to relatively heavily pigment- ed. Frequently there are several sparse clus- ters of melanophores irregularly placed along the spinous dorsal fin, more or less associ- ated with the more distal portions of indi- vidual spines; these sparse clusters begin posterior to the fifth spine, but in occasional specimens there is a larger, denser concen- tration of pigment on the membrane be- tween the third and fourth spines. Dorsal soft rays are more consistently pigmented, most specimens having many rays with sev- eral mostly dash-like melanophores lying along their anterior and/or posterior edges. Similar pigment occurs much less frequent- ly on the anal-fin rays. Most caudal-fin rays have dash-like melanophores along their dorsal and ventral edges.
USNM specimens: The melanophore patterns described above also characterize the fresher USNM specimens and will not be repeated here, where we describe only the additional purple-red pigment cells, re- ferred to, for convenience, as ““P-phores.”’ Both the persistent black melanophores and the transient P-phores are evident in the black-and-white photograph of the holo- type (Fig. 1), where the dichotomy is evi- denced to some extent in relative intensity of the spots. In fresh, unpreserved speci- mens, there is no detectible dichotomy, that is, all dark pigment cells appear black. Soon after formalin fixation, the majority of these dark spots fade to purplish-red and, with long-term storage in alcohol, apparently
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Fig. 2. Scanning electron micrographs illustrating head spines and cephalic sensory pores of three species of Acanthemblemaria in lateral (left) and dorsal (right) views; all males, supraorbital cirri removed. (A) A. paula, FMNH 90876, 15.4 mm SL; (B) A. aspera, USNM 276052, 19.1 mm SL; (C) A. spinosa, USNM 198276, 21.6
mm SL.
continue fading, eventually disappearing to leave only the black melanophores de- scribed above. (In rare specimens some pig- ment cells with distributions characteristic of P-phores appear black.) In all specimens, a fine speckling of P-phores begins in the pectoral region of the body at about the level of the sixth to eighth dorsal spine and ex-
tends anteriorly to cover much of the head, including the opercular series, branchi- ostegal membranes, pectoral-fin base and about half the surface of the fin, pelvic-fin base and rays and, in some specimens, the membrane between the first four or five dor- sal spines. The densest concentration of P-phores is seen on the branchiostegal
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membranes of some males; females tend to have this area more lightly pigmented, but this sexual difference is inconsistent. On the head, the most sparsely pigmented areas are the cheek, anterior portion of the cranium, and the jaws, which are frequently immac- ulate; denser concentrations are found just posterior and ventral to the orbit, forming one or two oblique bars. Superficial and subcutaneous P-phores may also be appar- ent on the abdomen, where they may be associated with the two distinctive crescents described for the melanophores. Dorsal to these, one or two small clusters of super- ficial P-phores occur on the upper body in some specimens. Deep, internal blocks of P-phores may be present on six to eight cen- tra along the length of the body, and, in such specimens, there are usually clusters of P-phores at the bases of every second or third dorsal- and anal-fin ray. These clusters begin internally around the pterygiophores and may rise to the surface at the fin-ray bases, sometimes extending a short distance along the rays. P-phores are also found along the bases of some caudal-fin rays.
In USNM specimens that were retained in formalin for several months before trans- fer to ethanol, white pigmentation (leuco- phores) is evident in discrete patches on the head, fins, and body. There is a prominent white patch or oblique bar, bordered by the two purple-red bars, on the lower portion of the cheek immediately posterior to the orbit. There is a triangular white patch on the dorsum of the cranium between and im- mediately posterior to the orbits, the iris is white with a speckling of P-phores, and the supraorbital cirrus is white. Several discrete white patches occur on the opercular series and branchiostegal membranes, and the en- tire gular region is white. Leucophores are also prominent on the pelvic-fin rays and the pectoral-fin base and proximal one-third or more of the fin. Large areas of white pig- ment may be seen internally on the abdo- men, and smaller superficial clusters may be seen on the body dorsal to this. The
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membrane between the first four to seven dorsal spines is white. Internal and super- ficial leucophores at the bases of dorsal- and anal-fin rays alternate with the similarly placed P-phores described above.
Live color.—The following description is based on specimens that were placed on ice while still alive, painted with full strength formalin to fix the fins, and photographed immediately thereafter. All pigment de- scribed above is evident, but, as noted, me- lanophores and P-phores are indistinguish- able. The head and anterior body (to about the anus), including the pelvic and proximal portion of the pectoral fins, appear predom- inantly white with prominent black and greenish-yellow (xanthophores) markings. Two distinctive black crescents are evident on the silvery-white background of the ab- domen. Xanthophores are in close associ- ation with melanophores (and P-phores), so that their distribution is essentially com- pletely overlapping. The large white blotch covering the anterior portion of the spinous dorsal fin is a very prominent feature. Pos- terior to the anus, the body is transparent with alternating white and dark markings along the lateral midline and median fin bases. Most distinctive is a series of seven to eight rectangular blocks of white pigment lying internally along the vertebral column, each encompassing two to four centra. These white blocks are bordered on either side by narrower, less circumscribed, dark greenish marks that frequently extend to the bases of the dorsal and anal fins.
Sex. —Specimens are easily sexed based on genital morphology. Males have a single papilla at the posterior margin of the anus, whereas in females the anus is surrounded by a papillar fringe. The ratio of males to females in both the USNM and FMNH col- lections is about two to one. Because most specimens were collected individually with quinaldine, we cannot discount the possi- bility that the sex ratio is partially a reflec- tion of collecting bias. Nonetheless it seems likely that males significantly outnumber fe-
VOLUME 102, NUMBER 4
males in the natural population. Greenfield & Greenfield (1982) reported that males of Acanthemblemaria spinosa and A. green- fieldi significantly outnumbered females in their collections, and Rosenblatt & Ste- phens (1978) reported that sex ratios in their collections of Mccoskerichthys sandae was strongly biased in favor of males.
There appears to be no pronounced sex- ual dimorphism; however, as described above, males tend to have slightly larger supraorbital and nasal cirri and usually have the tips of the anal-fin rays notably thick- ened and more strongly curved. Males also tend to have more intense and extensive dark pigmentation on the branchiostegal membranes than do females, but this is not a consistent difference. Males ranged in length from 10.1 mm to 18.4 mm with a mean of 14.8 mm, whereas females ranged from 10.3 mm to 16.7 mm with a mean of 13.1 mm. Size at sexual maturity and fe- cundity are discussed below.
Etymology. — Paula, Latin for little, in reference to the diminutive size of this fish, the smallest member of the Chaenopsidae, here used as a noun 1n apposition. The name was chosen to honor Paula Keener, who participated in the collection that resulted in recognition of this species.
Habitat. —A. paula occupies burrows made by invertebrates (usually those of si- punculids) in dead coral in shallow water (<5 m) on both sides of reef crest and on pavement zone of reef flat.
Distribution. —Known only from the bar- rier reef and outlying atolls of Belize.
Material examined. —114 specimens (10.1-18.4 mm) in 20 collections, all from Belize.
Holotype. —USNM 301835, 15.9 mm male taken with quinaldine from dead coral on reef flat at south end of Carrie Bow Cay, Belize, at depth of 1-2 m on 22 Mar 1988, by G. D. Johnson, and party.
Paratypes.—Carrie Bow Cay. South end of island, depth 1.0—2.5 m, coral rubble and pavement zone on reef flat: FMNH 90876
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(17, 13.8-18.4, 3 prepared for SEM), 4 Jan 1978, D. W. and T. A. Greenfield, and C. Rakocinski; FMNH 90869 (8, 12.9-16.6), 17 May 1977, D. W. and T. A. Greenfield; USNM 301831 (3, 13.1-16.7), 21 Mar 1988, G. D. Johnson; USNM 301836 (4, 13.4— 15.1), 22 Mar 1988, E. B. Brothers, R. A. Fritzsche, and G. D. Johnson; _USNM 301832 (10, 11.6-17.0), 25 Mar 1988, G. D. Johnson; ANSP 162806 (6, 11.8—16.0), 26 Mar 1988, G. D. Johnson; USNM 301833 (23, 12.1-16.7), 28 Mar 1988, R. A. Fritzsche and G. D. Johnson; SIO 89-14 (1, 15.4), 19 Nov 1988, R. A. Fritzsche; SIO 89-13 (5, 11.6-15.3), 19 Mar 1989. North end of island, coral and coral rubble just outside reef crest, 2-5 m: USNM 290669 (2, 12.4-14.9), 24 Mar 1987, E. B. Brothers, G. D. Johnson, and P. Keener; USNM 301834 (9, 10.3-15.2, cleared and stained), 17 Mar 1988, E. B. Brothers, R. A. Fritzsche, G. D. Johnson, and P. Keener. East side of island, just inside reef crest, 1-2 m: FMNH 98298 (3, 13.9-15.0), 16 May 1977, D. W. and T. A. Greenfield; USNM 301830 (3, 10.8-14.7), 27 Mar 1988, E. B. Brothers, R. A. Fritzsche, and G. D. Johnson. Collection site unspecified: FMNH 89372 (1, 14.8), 2 Jan 1978, D. W. Greenfield et al.; FMNH 89328 (2, 15.2-15.8), 14 May 1977, D. W. Greenfield.
Glover’s Reef. Cay in shallow water be- hind cabin no. 9: FMNH 90497 (12, 12.0- 155); 14% Jun £97 8s.» Glodck, «BD. (W: Greenfield, and R. K. Johnson. West side of Long Cay, about 200 yards south of cabin no. 9: FMNH 98296 (1, 14.2), 9 Jun 1978, G. Glodek and T. Murphy. Collection site unspecified: FMNH 77558 (1, 13.5), 29 Jul 1973, D. W. and T. A. Greenfield.
Buttonwood Cay. One mile south of is- land: FMNH 86085 (3, 11.7—13.6), 23 Jul 1974; FMNH 86088 (1, 10.1), 23 Jul 1974.
Sargeant’s Cay. FMNH 98297 (2, 11.6- 12.4).
Ambergris Cay. 2.5 miles north of San Pedro: FMNH 98295 (1, 14.8).
Discussion. —A cladistic analysis of rela-
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tionships within Acanthemblemaria is be- yond the scope of this study. Furthermore, although Stephens’ (1963) surmise that Ekemblemaria is the closest relative of Acanthemblemaria seems reasonable, char- acter polarity for Acanthemblemaria re- mains problematic in the absence of a well- corroborated cladistic hypothesis of generic relationships for the Chaenopsidae. Based on unpolarized similarities, we believe that A. paula is probably most closely related to A. aspera and medusa, two species that Smith-Vaniz & Palacio (1974) hypothesized to be closely related, but this remains to be tested cladistically. These three species (paula, aspera and medusa) share complex, deeply branched supraorbital cirri not found in other members of the genus. Head spines in A. paula are long as in A. maria and spinosa but not as numerous. Distribution of spines on top of the head is more like that in A. aspera and medusa, wherein there is a triangular patch on the frontals extend- ing posteriorly from the orbits. In paula this area is completely covered with spines, whereas in aspera and medusa it 1s largely smooth (Fig. 2). The extreme development of fleshy papillae that uniquely character- izes A. medusa is lacking in paula, and sev- eral characters suggest that paula and aspera are most closely related. They share the low- est dorsal-fin spine count in the genus (X = 19.2 and 20.5, respectively; 21.7 in medusa) and the lowest segmented caudal-fin ray count among Atlantic species (x = 12.4 and 12.3; 13.0 in medusa; data not available for eastern Pacific species). Finally, the color patterns of A. paula and aspera resemble one another more closely than they do that of any other species. Preserved specimens lack any regular external pigment pattern consisting of repeated bars, stripes or spots.
Acanthemblemaria paula may mature at sizes smaller than any known blennioid, al- though data in size at sexual maturity is lacking for most species. Females of A. pau- la may be ripe at 11.5 mm and individuals smaller than 11.0 mm may have developing
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eggs. Among the smallest known blennioids are the blenniid, Medusablennius chani (Springer, 1966) and the labrisomid, Stark- sia nannodes. Medusablennius chani is known from ten specimens ranging in size from a 12.1 mm ripe female to a 15.4 mm immature male (Smith-Vaniz & Springer 1971). Specimens of S. nannodes examined by Bohlke & Springer (1961) ranged from 10.5 to 17.0 mm with a 13.1 mm ripe fe- male. There is little published information on size at sexual maturity in chaenopsids, but all species attain maximum sizes larger than A. paula (18.4 mm). Within Acan- themblemaria, specimens 30 mm or longer (some >50 mm) are known for all 15 species. Whatever the precise relationships of A. paula to other species of the genus, its small size 18 unquestionably derived.
Weitzman & Vari (1988) listed 85 species representing five orders and 11 families of miniature Neotropical freshwater fishes. Miniature species were considered by Weitzman and Vari to be those that either are known to mature sexually at less than 20 mm and may reach slightly larger sizes (<25-—26 mm), or for which maturity data are unknown but which are not reported to exceed 25—26 mm in the wild. By those cri- teria, Acanthemblemaria paula is unequiv- ocally a miniature species. The elongate body and relatively small head further em- phasize its miniature stature; wet weights of 15 specimens, 13.4-16.4 mm in length, ranged from a mere 15 to 25 mg.
Weitzman & Vari (1988) noted that all species in their list of miniatures that had been examined in detail exhibit numerous apparently paedomorphic morphological reductions, particularly in the degree of de- velopment of the laterosensory canal sys- tem of the head and body, sculpturing on the bones of the head and total numbers of fin rays and body scales. The state of these and other characters in A. paula indicates that miniaturization does not always in- volve obvious reductive paedomorphic expression. The laterosensory ossifications
VOLUME 102, NUMBER 4
and pores (Fig. 2) of A. paula appear no less developed than in other species of the genus. Head spination is among the most extensive in the genus and considerably more pro- nounced than that of A. aspera, the putative closest relative, which reaches a maximum size of about 35 mm. Total numbers of dor- sal- and anal-fin rays are lower than those of some species but higher than others and not reduced compared to those of aspera. Stephens (1963) also noted the lack of cor- relation between reduced body size and numbers of dorsal- and anal-fin rays among other species of Acanthemblemaria. Pec- toral-fin ray counts do not differ substan- tially among species of Acanthemblemaria, and all chaenopsids lack scales, so no intra- generic comparisons of states of these char- acters can be made. We have identified only two features (neither absolute reductions) of A. paula that might be interpreted as pae- domorphic. Although the total number of dorsal-fin elements is equal to that of as- pera, paula has one more soft ray and one fewer spine; in fact, the mean dorsal-fin spine number (19.2) is the lowest in the genus. If transformation of the posteriormost dorsal- fin spines from soft rays occurs ontogenet- ically in chaenopsids, as it does in many percomorphs, the dorsal-fin ray composi- tion of A. paula could be interpreted as pae- domorphic with respect to the other species. The very slender, elongate body of A. paula, equalled in only one other species, A. chap- lini, could also be seen as paedomorphic, because chaenopsid larvae are relatively more elongate than adults. It seems clear that the marked reductive features that characterize miniaturization in Neotropical freshwater fishes and at least some marine fishes (e.g., some gobioids, Springer 1983, 1988), are not universally associated with extreme size reduction.
In a study of habitat and resource parti- tioning between Acanthemblemaria spinosa and A. greenfieldi, Greenfield & Greenfield (1982) compared ecological and life history parameters of the two species. All subse-
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quent references herein to G&G pertain to that paper. They found that A. spinosa oc- curs only on horizontal surfaces of dead cor- al whereas A. greenfieldi occurs only on ver- tical surfaces, but they noted an apparent exception in their collections. One collec- tion made in a typical A. greenfieldi habitat (coral rubble and pavement zone of the back reef) contained, in addition to 36 specimens of A. greenfieldi, 17 specimens identified as “young A. spinosa.’’ Catalog numbers were not given, but based on the date, locality and number of specimens, we conclude that that collection is FMNH 90869, which con- sists of 17 specimens of A. paula.
A total of 90 specimens identified as A. spinosa, collected at Carrie Bow Cay 10-17 May 1977 and 1-4 Jan 1978, were analyzed by G&G for length frequency, stomach con- tents and fecundity. Of these, 30 are actually A. paula. Consequently, most of G&G’s conclusions about life history parameters of A. spinosa are invalid. We did not attempt to reanalyze their data, because all 90 spec- imens were not used in determining each parameter and we were unable to determine how many A. paula were included in each analysis. Most profoundly affected are the data on size at sexual maturity and fecun- dity. Their observation that A. spinosa fe- males mature as small as 12.4 mm is clearly based on specimens of A. paula. We ex- amined 25 female A. spinosa, ranging in size from 11.6 to 20.4 mm; the smallest ripe individual was 15.6 mm and the mean size of ripe individuals was 18.4 mm. Of 22 fe- male A. paula, 10.3-16.7 mm, the smallest ripe specimen was 11.5 mm, and the mean size of ripe specimens was 13.6 mm.
Conclusions regarding average brood size in A. spinosa were also affected by inclusion of several specimens of A. paula. Based on data presented in fig. 218 of G&G, the num- ber of eggs per brood in A. spinosa ranged from 5 to 25 with a mean of 12.8. If spec- imens smaller than 15 mm (presumably all A. paula) are removed from the data, brood size ranges from 8 to 25, with a mean of
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14.8. This probably still includes some specimens of A. paula, in which the brood size is substantially smaller, but is in better agreement with our own data from ten spec- imens of A. spinosa with a range of 9 to 30 and a mean of 17.2. Among sixteen speci- mens of A. paula, numbers of eggs per brood ranged from 1 to 10, with a mean of only 4.7. This conforms with our single obser- vation of eggs in a burrow of a male A. paula; the clutch consisted of only four eggs, all apparently close to hatching. The egg size of A. paula and spinosa is roughly the same with the largest eggs averaging 0.7—-0.8 mm in diameter.
Misidentification of A. paula led G&G to the following speculation about larval re- cruitment in Acanthemblemaria:
The 4 January 1978 collection from a typ- ical Acanthemblemaria greenfieldi habi- tat, which also yielded young of A. spi- nosa, provides valuable information concerning the interactions of these two species. Although the adults exhibit com- plete habitat separation, apparently planktonic larval Acanthemblemaria species settle and occupy any available hole or crevice. Because individuals of A. spinosa found here were only 18.4 mm or less, it may be assumed that when they outgrow their holes and search for a larger hole, they are excluded by A. greenfieldi from this habitat.
Because the young A. spinosa referred to above are actually adult A. paula, these con- clusions are unsubstantiated. There is no evidence that individuals of A. spinosa oc- cupy the specific A. greenfieldi habitat at any stage of their lives. In contrast, individuals of A. paula occupy this habitat throughout their lives and are not excluded by A. green- fieldi as they grow; in fact, the specific col- lection referred to by G&G contains the two largest known specimens of A. paula. Co- existence is undoubtedly facilitated by the fact that the diminutive A. paula utilizes small burrows that are unavailable to the much larger adult A. greenfieldi; according
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to G&G, A. greenfieldi matures at 19.0 mm (almost twice the size for A. paula) and reaches a maximum size of 32.5 mm with mean sizes of 25.5 and 23.5 for males and females, respectively. Thus, although adults of A. paula may compete for space with newly settled or juvenile A. greenfieldi, there is fine-grain habitat (refuge) partitioning be- tween adults of the two species. This would be worth further investigation with respect to the findings of Stephens et al. (1970) on Hypsoblennius jenkinsi. Those authors found that maximum size of individuals was controlled by size of available tubes and hy- pothesized that individuals that outgrew the available tubes were subject to a greater risk of predation.
Our observations at Carrie Bow Cay on the microhabitat distribution of A. paula and greenfieldi failed to establish a clear-cut specificity to either vertical or horizontal surfaces; in areas where both surfaces were abundant, both were occupied. In a single collection (19 Mar 1989) using quinaldine just inside the reef crest we took: 7 speci- mens of greenfieldi and 8 of paula from ver- tical surfaces; 6 of greenfieldi and 13 of pau- la from oblique surfaces; and 2 of greenfieldi and 8 of paula from horizontal surfaces. We found these two species limited to horizon- tal surfaces only far back on the reef flat, where vertical surfaces are much less prev- alent. Acanthemblemaria spinosa does not occur far inside the reef crest and appears to live primarily on vertical surfaces. Dis- cussion by G&G of competitive interaction and displacement among Acanthemblemar- la species is speculative and not supported by evidence that shelter is a resource lim- iting population size. Hastings (1984) dem- onstrated the latter for A. crockeri, and sim- ilar experimental field manipulations of the habitat and/or fishes will be necessary to elucidate the nature of interspecific inter- actions between the species in question here. G&G made a valuable contribution by em- phasizing and reviewing the potential sig- nificance of very subtle differences in hab- itat utilization in small, cryptic reef fishes.
VOLUME 102, NUMBER 4
Fish community ecologists have shown an understandable bias for larger and more conspicuous fishes such as wrasses, dam- selfishes, surgeonfishes, butterflyfishes, etc. Population and behavioral characteristics of such groups may be entirely different from those of the diminutive, cryptic and fre- quently speciose blennies, gobies, eels, ophidioids, dactyloscopids, etc. Although some work has been done on chaenopsids and other blennies (e.g., Lindquist 1985, Stephens et al. 1966, Wirtz 1983), we re- main largely ignorant of the demography, life history and recruitment biology of such fishes.
The very small size of Acanthemblemaria paula and some of its congeners raises a number of interesting questions, particular- ly with respect to longevity, fecundity and larval ecology. How can such species main- tain populations when females apparently produce so few eggs in a lifetime? Acan- themblemaria species lay demersal eggs that presumably hatch to a planktonic larval stage. Based on otolith microstructure (Brothers et al. 1983) we have made prelim- inary determinations of the planktonic du- ration of the larvae of four species in the genus (paula, aspera, greenfieldi, spinosa). All show stereotypic “settlement marks” at presumptive ages of 22 to 25 days. A larval period of this magnitude is equal to or lon- ger than that exhibited by many larger reef species having instantaneous fecundities hundreds to many thousands of times great- er (e.g., Brothers & Thresher 1986). Do these chaenopsid blennies have any special eco- logical and/or behavioral adaptations that reduce larval mortality in the plankton? Do they bypass the planktonic phase complete- ly? Detailed life history studies of small, short-lived reef species will undoubtedly yield many surprises.
Acknowledgments
Barry Chernoff (FMNH) and William F. Smith-Vaniz (ANSP) assisted with the loan of specimens from their respective institu-
1029
tions. Donna Henriques (University of Washington, Seattle) prepared the radio- graphs and assisted with some of the counts. SEM facilities were made available by Spe- cialty Testing and Equipment of Ithaca, New York. The manuscript benefitted from the comments of David W. Greenfield (Uni- versity of Hawaii at Manoa), Philip A. Has- tings (University of Arizona, Tucson), Pau- la Keener (South Carolina Wildlife and Marine Resources Department), Richard H. Rosenblatt (SIO), William F. Smith-Vaniz, and Victor G. Springer (USNM). We are particularly grateful to Stanley H. Weitz- man for providing the excellent photo- graphs of the holotype. This is Contribution Number 263, Caribbean Coral Reef Eco- systems Program, National Museum of Natural History, Smithsonian Institution, Washington, D.C., partly supported by the Exxon Corporation.
Literature Cited
Acero P., A. 1984. The chaenopsine blennies of the southwestern Caribbean (Pisces: Clinidae: Chaenopsinae). II. The genera Acanthemble- maria, Ekemblemaria and Lucayablennius. — Revista de Biologia Tropical 32(1):35-44.
Bohlke, J. E., & V. G. Springer. 1961. A review of the Atlantic species of the clinid fish genus Starksia. — Proceedings of the Academy of Nat- ural Sciences of Philadelphia 1 13(3):29-60.
Brothers, E. B., & R. E. Thresher. 1986. Larval du-
ration, dispersal and biogeography of Indo-Pa-
cific coral reef fishes. Pp. 53-69 in M. Reaka, ed., The ecology of coral reefs. NOAA Sym- posium Series for Undersea Research, Wash-
ington, D.C. 3(2).
, D. M. Williams, & P. F. Sale. 1983. Length
of larval life in 12 families of fishes at One Tree
Lagoon, Great Barrier Reef, Australia. — Marine
Biology 76:319-324.
Greenfield, D. W., & T. A. Greenfield. 1982. Habitat and resource partitioning between two species of Acanthemblemaria (Pisces: Chaenopsidae), with comments on the chaos hypothesis. Pp. 499-507 in K. Ruetzler and I. G. Macintyre, eds., Atlantic Barrier Reef ecosystems at Carrie Bow Cay, Belize, 1: Structure and communities. Smithsonian Contributions to the Marine Sci- ences 12.
—,&R.K. Johnson. 1981. The blennioid fishes of Belize and British Honduras, Central Amer-
1030
ica, with comments on their systematics, ecol- ogy, and distribution (Blenniidae, Chaenopsi- dae, Labrisomidae, Tripterygiidae). — Fieldiana, Zoology, new series 8:1—106.
Hastings, P. A. 1984. Reource limitation in a coral- dwelling blennioid fish.—American Zoologist 24:128A [Abstract].
Johnson, R. K., & D. W. Greenfield. 1976. A new chaenopsid fish, Emblemaria hyltoni, from Isla Roatan, Honduras.—Fieldiana, Zoology 70(2): 13-27.
Lindquist, D. G. 1985. Depth zonation, microhabi- tat, and morphology of three species of Acan- themblemaria (Pisces: Blennioidei) in the Gulf of California, Mexico.—PSZNI: Marine Ecol- ogy 6(4):329-344.
Metzelaar, J. 1919. Report on the fishes collected by Dr. J. Boeke in the Dutch West Indies 1904— 1905. With comparative notes on marine fishes of tropical West Africa.— Rapport Visscher en de industrie van Zeeprodukten in de Kolonie Curacao, Gravenhage. 315 pp.
Rosenblatt, R. H., & J. E. McCosker. 1988. A new species of Acanthemblemaria from Malpelo Is- land, with a key to the Pacific members of the genus (Pisces: Chaenopsidae).— Proceedings of the California Academy of Sciences 45(7):103- 110.
—., & J. S. Stephens, Jr. 1978. Mccoskerichthys sandae, a new and unusual chaenopsid blenny from the Pacific coast of Panama and Costa Rica.— Contributions in Science of the Natural History Museum of Los Angeles County 293: 1-22.
Smith-Vaniz, W. F., & F. J. Palacio. 1974. Atlantic fishes of the genus Acanthemblemaria, with comments on Pacific species (Clinidae: Chae- nopsinae).— Proceedings of the Academy of Natural Sciences of Philadelphia 125(11):197- 224.
—., & V.G. Springer. 1971. Synopsis of the tribe Salariini, with description of five new genera and three new species (Pisces: Blenniidae).— Smithsonian Contributions to Zoology 73:1—72.
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Springer, V. G. 1966. Medusablennius chani, a new genus and species of blennioid fish from Tua- motu Archipelago: Its implication on blennioid classification.—Copeia 1966(1):56—60.
1983. Tyson belos, new genus and species of western Pacific fish (Gobiidae, Xenisthminae), with discussions of gobioid osteology and clas- sification.—Smithsonian Contributions to Zo- ology 390:1-40.
1988. Rotuma lewisi, new genus and species of fish from the southwest Pacific (Gobioidei, Xenisthmidae).— Proceedings of the Biological Society of Washington 101(3):530-539. Stephens, J. S., Jr. 1963. A revised classification of the blennioid fishes of the American family Chaenopsidae.— University of California Pub- lications in Zoology 68:1-133.
, E. S. Hobson, & R. K. Johnson. 1966. Notes on distribution, behavior, and morphological variation in some chaenopsid fishes from the tropical eastern Pacific, with descriptions of two new species, Acanthemblemaria castroi and Coralliozetus springeri.—Copeia 1966(3):424— 438. , R. K. Johnson, G. S. Key, & J. E. McCosker. 1970. The comparative ecology of three sym- patric species of California blennies of the genus Hypsoblennius Gill (Teleostomi, Blennidae).— Ecological Monographs 40(2):213-233. Weitzman, S.H.,&R.P. Vari. 1988. Miniaturization in South American freshwater fishes; an over- view and discussion.— Proceedings of the Bio- logical Society of Washington 101(2):444-465. Wirtz, P. 1983. The reproductive behavior of three blennioid fish endemic to the Galapagos Is- lands.— Noticias de Galapagos 37:26-27.
(GDJ) Division of Fishes, National Mu- seum of Natural History, Smithsonian In- stitution, Washington, D.C. 20560; (EBB) 3 Sunset West, R.D. 7, Ithaca, New York 14850.
PROC. BIOL. SOC. WASH. 102(4), 1989, pp. 1031-1035
A NEW SPECIES OF EUPSOPHUS (ANURA: LEPTODACTYLIDAE) FROM CONTULMO, NAHUELBUTA RANGE,
SOUTHERN CHILE
Juan Carlos Ortiz, Héctor Ibarra-Vidal, and J. Ramon Formas
Abstract. —Eupsophus contulmoensis, a new species of leptodactylid frog is named from the locality of Contulmo, Nahuelbuta Range, Southern Chile. This species is distinguished from its congeners by the dark-purple dorsum and the bright yellow coloration of its belly. This frog displays lumbar amplexus and
is known only from the type locality.
The Nahuelbuta Range is that part of the Chilean Coastal Range bounded to the northeast by the Biobio river and to the south by the Imperial river (see Fig. 1). This mountainous area of southern Chile is ap- proximately 175 km in length and reaches 1530 m at its greatest elevations (Alto de Nahuelbuta). The Nahuelbuta Range is cov- ered by Nothofagus temperate forest and at elevations of more than 1000 m the conifer Araucaria araucana 1s present.
In this area three endemic anuran species are found (Telmatobufo bullocki, Alsodes vanzolinii, and Alsodes barrioi) (Schmidt 1952, Formas 1981, Veloso et al. 1981). Nonendemic species include: Bufo rubro- punctatus, Pleurodema thaul, Rhinoderma darwinii, R. rufum, Hylorina sylvatica, Ba- trachyla leptopus, B. taeniata, Eupsophus roseus, and E. vittatus.
The Nahuelbuta Range shows a high de- gree of human-induced disturbance (pine groves of Pinus radiata) and little original forest remains. One of these areas is the Natural Monument of Contulmo (37°02’S; 78°12’W), where a series of herpetological collections were made between 1986 and 1987. As a result of this fieldwork a new species of frog of the genus Eupsophus was collected.
Eupsophus contulmoensis, new species Fig. 2
Holotype.—MZUC (Museo de Zoologia, Universidad de Concepcion, Chile) 17141, adult female collected by Hector Ibarra-Vi- dal, 10 Jul 1987 at Contulmo, Malleco Province, Nahuelbuta Range, alt. 700 m, 15 km W (by road) of Purén, Chile (Fig. 1).
Paratypes.— Four adults (MZUC 17142, 17145, 17148, 17149) and one subadult (MZUC 17144) collected at the type local- ity.
Diagnosis. —A medium-sized species of Eupsophus (34.0-42.5 mm SVL), distin- guished from its congeners (E. roseus, E. migueli, E. calcaratus, E. insularis and E. vittatus) by the dark purple dorsal pigmen- tation and bright yellow belly; upper part of the iris bronze-yellow in life and inner pal- mar tubercle prominent.
Description of adult (based on five fixed specimens). — Head slightly wider than long. Snout rounded in dorsal and lateral view, canthus rostralis concave, loreal area slight- ly concave, nostrils located laterally, at middistance between snout tip and orbit; eye length greater than distance between eye and nostril; interorbital distance smaller than eye length but greater than internarial
1032
OCONCEPCION
we 7
° 3 ;
Polo\/ Sur |!
CONTULM Q. ¢ QPUREN \
&) ] VICTORIA
Fig. 1. Situation of the type locality (Contulmo) of E. contulmoensis.
distance. Tympanic membranes present and well developed. Supratympanic fold absent. Tongue round, notched at tip. Choanae rounded, dentigerous processes of vomers lying below the choanae; each process bear- ing five or six sharp teeth.
Forelimbs slender, 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 protuberant. Inner palmar tubercle prominent; outer pal- mar tubercle ovoid and well developed; subarticular tubercles rounded and mod- erate in size; supernumerary palmar tuber- cles absent. Toes long, slender; tips of toes round; third and fifth equal in length; toes in decreasing order of length 4-(3,5)-2-1. In- ner metatarsal tubercle ovoid and promi- nent; supernumerary tubercles absent; outer metatarsal tubercle tiny. Rudiment of web between toes.
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Fig. 2. Eupsophus contulmoensis, new species. Ho- lotype (MZUC 17141), female.
Anal opening rounded and directed pos- tero-ventrally at dorsal level of thighs.
Dorsal and ventral skin smooth. Two faintly marked folds extending from pos- terior part of eye to the midlateral part of body.
Pectoral girdle arciferal; omosternum cartilaginous with short thin stem; tip sharply acute; sternum rectangular in shape with slight median constriction, its tip rounded and almost completely calcified. External measurements are shown in Ta- ble 1.
Color and color patterns. —In life dorsal ground color of head and body dark purple, two specimens (MZUC 17142, 17148) with yellow vertebral line (Fig. 3); dorsal limb surfaces dark purple with small yellow ir- regular spots; throat dark brown with mi- nute irregular yellow spots; ventral surface brown and marbled with yellow; two spec- imens (MZUC 17141, 17144) with immac- ulate bright yellow abdomen; ventral sur- face of limbs with yellow irregular spots; yellowish irregular marks on side of head and body; upper part of iris bronze-yellow.
In alcohol dorsal surfaces dark brown and
VOLUME 102, NUMBER 4 1033 Table 1.—Measurements of the type series of Eupsophus contulmoensis (mm). Holotype MZUC MZUC MZUC MZUC MZUC MZUC 17141 17142 17145 17148 17149 17144 female female female female male male subadult Snout—vent length 45.2 39.4 43.0 44.4 38.1 34.0 Tibia length 24.2 24.5 2305 23.4 21.4 19.5 Foot length 34.1 33.3 30.8 33.8 31.6 26.5 Head length 1523 13.8 13.9 14.8 15.4 11.9 Head width 16.3 16.2 16.2 ie E37 27 Interorbital distance ae 4.5 4.7 Sail 5.5 Sh Internarial distance 3 Be 4.3 4.3 4.1 3:3 Diameter of eye 6.1 502 3.1 5.4 5.6 4.1 Diameter of tympanum 2.6 2.6 Dee, Zan 21 1.6 Eye-nostril 4.4 4.1 2.9 3.4 2 23
vertebral line whitish; ventral areas brown and whitish spotted.
Distribution. —Known from the type lo- cality.
Etymology.—The specific name of this frog is after the type locality.
Natural history. — The type locality, Con- tulmo, is a small natural reserve (approxi- mately 1 km7), in the Nahuelbuta Range, where the original Nothofagus forest yet sur- vives. Contulmo is situated in the mediter- ranean perhumid region (di Castri 1968). The annual mean temperature is 12.6°, the relative humidity is 82% and the annual mean rainfall is 1896 mm (Hajek and di Castri 1975). The following trees occur there: Nothofagus oblicua, N. dombeyi (Fagaceae), Eucryphia cordifolia (Eucryphiaceae), Per- sea lingue (Lauraceae), Laurelia phillipiana (Monomiaceae), and Aetoxicum punctatum (Aetoxicaceae). The climber Lapageria ro- sea (Phileseaceae) was observed on logs. Ferns (Lophosoria quadripinnata, and Cte- nitis spectabilis) and the moss Dendroligo- trichum dendroides were collected on the ground. During winter, frogs were collected under decaying logs and stones near streams.
The following species of amphibians were also collected at the type locality: Eupsophus roseus, E. vittatus, Batrachyla leptopus, and Rhinoderma darwinii.
A female collected in spring (Nov 1987) had 65 white oocytes (1.14—2.28 mm di-
ameter) in its ovaries, and a male had testes 4.6 mm in length. Mature males did not have nuptial asperities in winter, however the gular areas were darker than in the an- imals collected. In the laboratory inguinal amplexus was observed.
The stomach contents of two Eupsophus contulmoensis collected on 7 Nov 1987, were examined. Both specimens were killed just after capture. The following food items were identified: Oligochaeta (4), Aranea (3), Di- plopoda (2), Coleoptera (2), Diptera (1), Collembola (1), and Formicidae (1). Two specimens collected in 5 Sep 1987 had emp- ty stomachs.
Comparisons
Eupsophus contulmoensis is a frog of moderate size (xX = 42.0 mm snout—vent length) as are E. calcaratus (X = 35.1 mm, Formas & Vera 1982), E. migueli (x = 35.5 mm, Formas 1978), FE. roseus (X = 36.0 mm; Cei 1962), and E. insularis (X = 39.3 mm, Formas & Vera 1982). These species are notably smaller than E. vittatus (xX = 59.4, mm Grandison 1961). Eupsophus contulmoensis and E. insularis differ in the dorsal color and in the shape of the tip of the sternum. The latter species is dark brown with yellow irregular spots on the dorsum and the sternum is truncated whereas E.
1034
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig. 3. Color pattern variation in E. contulmoensis (not to scale). Dorsal patterns (above) and ventral patterns
(below) of the specimens MZUC 17144, 17148.
contulmoensis is dark purple dorsally with a rounded sternum. The upper part of the iris is bronze yellow in E. contulmoensis and the dorsal area is unmarked, while in E. roseus the upper part of the iris is orange and an hour-glass shaped mark is present on its dorsum. Eupsophus contulmoensis, E. migueli and E. calcaratus have a similar colored upper iris (bronze yellow), but these species differ in the ventral color and pat- tern. The belly of E. migueliis wine red with irregular white spots whereas the ventral area of E. contulmoensis is dark brown with bright yellow irregular spots. In E. calcar- atus the spots are also present. On the other hand the dorsum of FE. migueli and E. cal-
caratus exhibits a typical hour-glass pattern that is absent in E. contulmoensis.
Acknowledgments
Collecting permits were graciously issued by Carlos Sepulveda, Director of Corpora- cion Nacional Forestal (CONAF, VIII Re- gion). We are grateful to Ismael Matamala, ranger of the Natural Monument of Con- tulmo, for his field assistance. Roberto Ro- driguez identified the plants. Fieldwork was supported by Direccion de Investigacion Universidad de Concepcion (Proyecto 20.38.04). Corina Zuniga typed the manu- script.
VOLUME 102, NUMBER 4
Literature Cited
Cei, J. M. 1962. El género Eupsophus en Chile.— Investigaciones Zoolégicas Chilenas 8:7-42.
di Castri, F. 1968. Esquisse écologique du Chili. Pp. 7-52 in Doutteville and Rapaport, eds., Biologie de l’Amerique Australe vi. Editions du Centre National de la Recherche Scientifique.
Formas, J. R. 1978. A new species of leptodactylid frog (Eupsophus) from the Coastal Range in Southern Chile. —Studies on Neotropical Fauna and Environment 13:1-19.
1981. The identity of the frog Eupsophus
vanzolinii from Ramadillas, Nahuelbuta Range,
Southern Chile.— Proceedings of the Biological Society of Washington 93:920-927.
—. & M.I. Vera. 1982. The status of two Chilean frogs of the genus Eupsophus (Anura: Lepto- dactylidae).— Proceedings of the Biological So- ciety of Washington 95:594-601.
Grandison, A.G. 1961. Chilean species of the genus
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Eupsophus (Anura: Leptodactylidae).— Bulletin of the British Museum (Natural History) 9:111- 149.
Hajek, E. R., & F. di Castn. 1975. Bioclimatografia de Chile. Universidad Catolica de Chile, San- tiago, 107 pp.
Schmidt, K. P. 1952. A new leptodactylid frog from Chile. —Fieldiana, Zoology 34:11-15.
Veloso, A., P. Iturra, & M. Penna. 1981. Descripcion de una nueva especie de telmatobino del género Alsodes (Amphibia, Leptodactylidae) de la cor- dillera de Nahuelbuta (Sur de Chile).— Medio Ambiente 5:115-121.
(JCO and HIV) Departamento de Zool- ogia, Universidad de Concepcion, Casilla 2407-10, Concepcion, Chile; (JRF) Insti- tuto de Zoologia, Universidad Austral de Chile, Casilla 567, Valdivia, Chile.
PROC. BIOL. SOC. WASH. 102(4), 1989, pp. 1036-1044
A NEW SPECIES OF COLUBRID SNAKE OF THE GENUS CONIOPHANES FROM THE HIGHLANDS OF CHIAPAS, MEXICO
Jonathan A. Campbell
Abstract.—A new species of colubrid snake, Coniophanes alvarezi, is de- scribed from the Meseta Central of Chiapas, Mexico. This snake inhabits tem- perate mesic pine-oak forest at elevations of 2012 to 2134 m. Unlike C. alvarezi, most other members of this genus have essentially lowland, tropical distribu- tions. Coniophanes alvarezi 1s easily distinguished from its congeners by its uniformly brown dorsal coloration in adults (weakly striped in juveniles), im- maculate venter, and features of lepidosis and dentition. This species appears to be most closely related to the wide-ranging C. fissidens, and a detailed comparison of these two species is presented.
Over 15 years have now passed since I first discovered an unusual colubrid snake of the genus Coniophanes near Teopisca in Chiapas, Mexico. On subsequent trips through the region Gin 1976 and 1983) ad- ditional material representing this taxon was collected, including three eggs that were al- lowed to hatch. In all, a total of seven spec- imens are now available. I propose that this snake, apparently endemic to the Mexican state of Chiapas, be known as:
Coniophanes alvarezi, new species Figs. 1-3
Holotype.—The University of Texas at Arlington (UTA) R-12256 (original field number, JAC 9389), an adult male from 11.3 km ESE Teopisca, 2073 m elevation, Chiapas, Mexico, collected by J. A. Camp- bell on 7 Aug 1983.
Paratypes. —Six specimens, all from Chiapas, Mexico: UTA R-2793, an adult female from 12.1 km ESE Teopisca, 2134 m elevation, collected 14 Aug 1973; UTA R-5766-67, adult females from 10.3 km ESE Teopisca, 2012 m elevation, collected 16 Jun 1976; and UTA R-6111-13, two neo- nate males and a female, respectively, from
10.3 km ESE Teopisca, 2012 m elevation, that hatched on 18 Aug 1976 from eggs dis- covered beneath a log on 29 May 1976.
Diagnosis.—A relatively large species of Coniophanes in which females are known to reach 521 mm in total length; dorsal scales disposed in 19-19-17 rows; adults differing from all congeners in having a uniformly brown dorsal coloration and immaculate yellow venter (except for a few tiny black stipples anteriorly). All other members of the genus have at least traces of longitudinal body striping, except C. /ateritius and some specimens of C. meridanus, both of which have an orange or red dorsal coloration and a black head or collar.
Description of holotype. —An adult male, 375 mm in total length; tail length 93 mm (24.8% of total); head length 14.9 mm from front face of rostral to posterior end of man- dible; head width 8.5 mm at broadest point (level of angle of mouth); head moderately distinct from neck; snout acutely rounded in dorsal view; snout 2.3 times as long as horizontal distance across eye; pupil round; rostral about 1.7 times broader than high; head scutellation of generalized colubrid type; internasals 1.4 times wider than long, laterally contacting anterior and posterior
VOLUME 102, NUMBER 4
1037
Fig. 1.
nasals; prefrontals large, slightly longer than wide, laterally contacting posterior nasal and loreal, forming upper anterior margin of or- bit; median prefrontal suture about half as long as frontal; frontal 1.6 times longer than wide; parietals about 1.6 times longer than wide, median suture slightly less than fron- tal length; nostrils located in central pos- terior portion of anterior nasals; loreal about half of combined length of nasals; two post- oculars; temporals 1 + 2, separating supra- labials 6 and 7 from parietal; supralabials 7/7, 1st contacting nasals and loreal, 2nd contacting loreal and preocular, 3rd con- tacting preocular and orbit, 4th contacting orbit and lower postocular, 5th contacting lower postocular and anterior temporal, 6th contacting primary and lower secondary temporals, 7th contacting lower secondary temporal; mental two times broader than long, separated from chinshields by first pair of infralabials which contact each other along the midline; anterior chinshields well de- veloped, about twice as long as wide; pos-
Dorsal aspect of Coniophanes alvarezi, holotype, UTA R-12256, 375 mm TL.
terior chinshields well differentiated from gulars, separated from Ist ventral by two gulars plus two preventrals; infralabials 9/9, 1-4 contacting anterior chinshields, 4th largest; dorsal scales disposed in 19 smooth rows, reduced to 17 posteriorly; dorsal scales in six rows at level of 10th subcaudal; no apical pits; ventrals 134; supra-anal tuber- cles (keels) present; anal divided; subcau- dals 64, paired.
In life, dorsal ground color uniformly brown; venter immaculate pale yellow, ex- cept on the anterior third of the body where a small amount of fine black peppering pres- ent, especially on head and gular region; a black line running across upper portion of rostral and extending posteriorly just below eye to angle of mouth, thence posteriorly for two or three scales, bordered ventrally by white; supralabials mostly white, heavily speckled with black; iris copper-colored, with heavy suffusion of black ventrally.
In preservative (alcohol after formalin) ground color of dorsum brown, except where
1038
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Fig. 2. Ventral aspect of Coniophanes alvarezi, holotype.
the stratum corneum sloughed off, where it is gray; venter creamy white.
Everted hemipenis five subcaudals in length; organ simple with sulcus spermati- cus bifurcating at about level of subcaudal 3, but extending distally on capitulum for a length less than half of everted capitulum; asulcate, basal portion of organ possesses three large basal hooks, distal to which are some 7—9 large spines; sulcate side of organ also bears some 13-15 spines, but these are considerably smaller than those on the asul- cate side; organ distinctly capitate; margin of capitulum covered with spinulate ca- lyces, toward apex calyces become papillate; capitulum extends proximally almost half the length of organ on sulcate side; on asul- cate side a pair of naked grooves extend distally from several large spines almost to apex (1.e., to overhang of the capitulum).
Variation. —The paratypic series is com- posed of two males and four females. The holotype is the largest male; the largest fe-
male has a total length of 521 mm, and a tail length of 111 mm (21.3% of total). Three juveniles, preserved on the date of hatching, were 154-162 mm in total length, with tail lengths of 34—38 mm (21.7—24.7% of total). Only one snake in the type series, an adult female, has an incomplete tail. The supra- labials are invariably 7/7; the infralabials are 9/10 in one specimen, 9/9 in all others. The number of dorsal scale rows is 19-19- 17 in all but one specimen which has 16 rows posteriorly. There are 134-136 and 140-143 ventrals, and 59-64 and 54-57 subcaudals, in males and females, respec- tively. The top of the head in the juveniles (UTA R-6111-13) is dark brown and con- trasts with the pale brown of the dorsum. Juveniles have a faint middorsal stripe, darker than the ground color, involving only the vertebral scale row, extending the length of the body and tail, and have almost in- distinguishable indications of a dark lateral stripe on scale row 4.
VOLUME 102, NUMBER 4
1039
Fig. 3. Lateral aspect of Coniophanes alvarezi showing details of head pattern.
Etymology. —The name alvarezi is a pat- ronym in honor of Miguel Alvarez del Toro, Director of the Instituto de Historia Natural of Chiapas. His noble love for the flora and fauna of southern Mexico has led to a better understanding of the natural history of Chiapan wildlife and forests and to the first significant conservation efforts taken in this state.
Distribution. —This species is known only from the Meseta Central of Chiapas at 2012 to 2134 m elevation. These highlands are covered by a temperate mesic pine-oak for- est with abundant epiphytes. All specimens of Coniophanes alvarezi were taken under rocks or logs at the edges of clearings.
Comparison with Coniophanes fissi- dens.—Owing to characters of lepidosis, color pattern, hemipenes, and dentition, Coniophanes alvarezi appears to be most closely related to C. fissidens. The geograph- ical distribution of C. a/varezi is more closely approached by that of C. fissidens than by
any other species of Coniophanes. A com- parison of selected features between C. al- varezi and C. fissidens is presented in Ta- ble 1.
Throughout most of its range C. fissidens usually has 21 dorsal scale rows at midbody (Bailey 1939, Minton & Smith 1960, Smith 1941). However, several isolated popula- tions of C. fissidens in the northern part of the range have a modal number of 19 dorsal scale rows at midbody. The populations of C. fissidens inhabiting eastern San Luis Po- tosi and west-central Veracruz, described as C. f. proterops (Smith, 1941) and C. f. con- vergens (Shannon & Smith, 1949), resemble C. alvarezi in usually having 19 midbody dorsal scale rows and 7 supralabials. These snakes also have a relatively high number of ventrals for the species (males, 121-133: females, 126-133) and low number of sub- caudals (males, 63-80; females, 59-74) (Smith 1941; Smith & Laufe 1945; Taylor 1949, 1953), but the counts do not overlap
1040
the range of variation known for C. alvarezi. The pattern of longitudinal body striping is relatively subdued, but always present, at least posteriorly, in adults (Smith 1941; Taylor 1949, 1953), and is distinct in ju- veniles and subadults (Smith 1941). Con- iophanes f. proterops was reported from high elevations of the Atlantic versant of Chiapas by Smith & Williams (1963) on the basis of three specimens from two localities, 13 miles east of Las Rosas and Monserrat. I have not re-examined these specimens, housed in the University of Illinois Museum of Natural History, but the description provided by these authors is similar in some respects to C. alvarezi, although the ventral and sub- caudal counts suggest they may be all males rather than females as stated.
Specimens of C. fissidens from the Pacific slopes of Michoacan and Guerrero, de- scribed as C. f. dispersus (Smith, 1941), also have 19 midbody scale rows (Bailey 1939, Davis & Dixon 1959, Peters 1954, Smith 1941), but in most other respects of lepi- dosis and pattern do not closely resemble C. alvarezi.
Along the Atlantic versant of Middle America, Coniophanes fissidens occurs in well drained mesic forests from eastern San Luis Potosi and west-central Veracruz (Mexico) eastward across east-central Belize and northern Guatemala through Hondu- ras, Nicaragua, Costa Rica, and Panama (Alvarez del Toro 1983, Henderson & Hoevers 1975, Lee 1980, Minton & Smith 1960, Myers 1969, Schmidt 1941, Shreve 1957, Stuart 1963, Wilson & Meyer 1985, Zug et al. 1979), but it is absent from most of the Yucatan Peninsula. Coniophanes fis- sidens also occurs along the Pacific versant from Michoacan and Guerrero (Mexico) eastward through Central America to Ec- uador (Bailey 1939, Myers 1969, Peters 1954, Peters & Orejas-Miranda 1970, Smith & Taylor 1945, Zug et al. 1979). On the Pacific slopes this species has been reported from up to 1463 m in Mexico (Landy et al. 1966) and 1432 m in Guatemala (Campbell
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& Vannini 1988). I have taken two speci- mens of this species (UTA R-16015, 20744) on the Atlantic side of Guatemala in cloud forest at elevations of 1500-1600 m. The range of C. fissidens surrounds the highlands of the Meseta Central of Chiapas, but in no” place is this species known to be sympatric with C. alvarezi or to attain similar eleva- tions.
Ina sample of 71 specimens of C. fissidens from southern Mexico and Guatemala (62 of Pacific slope origin, 9 from the Atlantic versant; Appendix), a number of salient dif- ferences are apparent between this species and C. alvarezi (Table 1). Coniophanes al- varezi differs from populations of C. fissi- dens in southern Mexico and Guatemala in having 19 midbody dorsal scale rows (vs. usually 21 in C. fissidens); 7 supralabials (vs. usually 8—9); usually 9 infralabials (vs. 10- 11); 134-136 and 140-143 ventrals in males and females, respectively (vs. 114-124 and 119-131, respectively); 59-64 and 54—57 subcaudals in males and females, respec- tively (vs. 67-86 and 59-86, respectively); a uniformly brown or brownish olive dor- sum in adults (vs. dark lateral and usually middorsal stripes); an essentially immacu- late yellow venter (vs. a white or cream col- ored venter, heavily marked with dark spots, speckling, or mottling); and a relatively short tail that accounts for 22.8—24.8% and 20.9— 21.7% of the total length in males and fe- males, respectively (vs. 29.3-35.4% and 25.5-—33.8%, respectively).
Coniophanes fissidens from the Atlantic versant of southern Mexico and Guatemala often possess large prominent dark spots on the lateral edges of the ventrals; the venter of snakes from Pacific Chiapas and Gua- temala is heavily strippled or flecked with black anteriorly, less so posteriorly.
The vague dark stripes in juvenile C. al- varezi are not positioned the same as in C. fissidens, but may have been derived from the pattern evident in the latter species. The
color pattern in juvenile C. a/varezi consists
of a poorly indicated dark strip confined to
VOLUME 102, NUMBER 4 1041
Table 1.—Comparison of Coniophanes alvarezi and C. fissidens from southern Mexico and Guatemala for selected features of lepidosis, color pattern, and proportion. These data are based on specimens examined (see Appendix).
Coniophanes alvarezi Coniophanes fissidens
Dorsal scale rows 19-19-17(16)! 21(19}-21(19}-17(15)? Supralabials 7 (100)% 7 (0.7%) 8 (97.9%) 9 (1.4%) Infralabials 9 (93.0%) 9 (12.7%) 10 (7.0%) 10 (86.6%) 11 (0.7%)
134-136 (x = 135.3) 140-143 (x = 141.3) 59-64 (X = 62.3) 54-57 (x = 55.7) Uniformly brown or brownish olive
114-124 (« = 119.2) 119-131 (* = 124.7) 67-86 (X = 77.5) 59-86 (x = 73.0) With distinct dark lateral stripes; often with dark dorsal stripe
Ventrals (males) (females) Subcaudals (males) (females) Adult dorsal color pattern
Usually white or cream, with dark spots or mottling on lateral portion of ventrals
29.3-35.4% (X = 33.1%)
25.5-33.8% (X = 30.9%)
Immaculate yellow, except for a few dark flecks in gular region
Ventral color pattern
22.8-24.8% (X = 24.1%) 20.9-21.7% (% = 21.3%)
Tail/total length (males) (females)
! The dorsal scale rows are reduced posteriorly to 16 in one paratype (UTA R-2793).
2 In 5 of 62 specimens examined from the Pacific versant of Guatemala and southern Mexico, the number of dorsal scale rows is reduced to 19 anteriorly and/or at midbody; one of nine specimens from the Atlantic slopes of southern Mexico and Guatemala has 19 dorsal scale rows at midbody. Only one specimen (UTA R-20711) from a large series collected at Finca El Faro, Quezaltenango, Guatemala, has 15 dorsal scale rows posteriorly.
the vertebral scale row and extremely faint darker lateral stripes on scale row 4. The dark coloration on the dorsum of C. fissi- dens extends over 3 to 7 dorsal scale rows, including the vertebral row, and the lateral stripes are broad, extending from scale rows 1 to 5, with particularly dark pigment on the upper portion of scale row 4 and the lower portion of scale row 5.
The hemipenes of Coniophanes fissidens differ from those of C. al/varezi in that on the asulcate side of the organ the distal most spines are nearly as large or larger than the
basal hooks, and in that there is no pair of}
naked grooves.
In a paratype of C. alvarezi (UTA R-5767) the right maxillary bone bears 9 teeth (counting sockets), followed by a broad di- astema and two enlarged fangs with deep anterolateral grooves extending four-fifths of their length. The right palatine and pter-
ygoid bones have 7 and 21 teeth, respec- tively. Examination of the dentition and as- sociated bones of ten specimens of C. fissidens from Quezaltenango (Guatemala), only some 150 km frm the type locality of C. alvarezi, reveals a number of differences. In comparison with C. fissidens, the teeth are relatively shorter, stouter, and not so strongly recurved; the choanal process of the palatine and the ectopterygoid process of the maxilla are more broadly expanded; and the ectopterygoid is more robust in C. alvarezi. There are 13-14 maxillary teeth (including the enlarged posterior teeth), 9-11 palatal teeth, and usually more than 21 pter- ygoid teeth in C. fissidens. The two posterior maxillary teeth of C. fissidens have strik- ingly different grooves from C. alvarezi. In C. fissidens the groove is very wide at the base of the tooth and tapers distally; in C. alvarezi the groove is no wider proximally
1042
than distally. Further, the tips of the rear maxillary teeth in C. fissidens become dis- tinctly compressed and bladelike, whereas in C. alvarezi the tips of these teeth are only slightly modified.
Comparisons with other species of Con- iophanes. — Other than C. alvarezi, only two species of Coniophanes may lack any trace of longitudinal body striping. Coniophanes lateritius occurs in the Pacific Mexican low- lands and foothills from Sinaloa (Hardy & McDiarmid 1969) to Oaxaca (Smith & Tay- lor 1945). This species has a black head or collar and a red dorsum, often becoming dark posteriorly (Hardy & McDiarmid 1969, Smith & Grant 1958). Coniophanes lateri- tius further differs from C. alvarezi in hav- ing usually more ventrals (140-146) and subcaudals (84-99). Coniophanes meridan- us has a uniclor reddish dorsum, pale tem- poral stripes, an incomplete black collar, often a trace of a middorsal stripe, and a bifurcate spineless hemipenis that extends 11 subcaudals (Bailey 1939). Pale temporal stripes are present in C. imperialis, C. pi- ceivittis, and C. schmidti, and the latter two species have a broad dorsal black band. A double row of conspicuous black spots is present on the ventrals of C. bipunctatus and C. quinquevittatus. Coniophanes bipuncta- tus, C. piceivittus, C. quinquevittatus, and C. schmidti usually have 21 or more dorsal scale rows at midbody, C. meridanus and C. Joanae have 17. The number of ventrals is higher in C. piceivittus (153-174), C. quin- quevittatus (152-163), and C. schmidti (158— 175), and lower in C. joanae (131-132). Coniophanes alvarezi has fewer subcaudals (54-64) than any congener except C. joanae (47+ 53) and, very rarely, specimens of C. imperialis (62-94) and C. quinquevittatus (63-70). Coniophanes dromiciformis is a striped South American species and is not considered here.
The only other named species of Conio- phanes with an essentially upland distri- bution is C. joanae, known from mesic
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
montane forests in Panama east of the canal at elevations of 500-1440 m (Myers 1966, 1969). This species differs from C. alvarezi in having a brown venter and supralabials, longitudinal body striping, and 17 dorsal scale rows at midbody. Coniophanes bi- punctatus, C. imperialis, C. meridanus, C. quinquevittatus, and C. schmidti are restrict- ed to lowland habitats and their range in- cludes the northern portion of the Yucatan Peninsula (Duellman 1965, Lee 1980, McCoy 1969, Smith 1960). At least two of these species, C. bipunctatus and C. quin- quevittatus, are semiaquatic and found in swampy or coastal lagoon habitats (Conant 1965, McCoy 1969, Myers 1969).
Resumen
Se describe una nueva especies de colu- brido, Coniophanes alvarezi, de la Meseta Central de Chiapas, México. Esta especies de culebra occure en los bosques humedos de pino y encino a elevaciones de 2012 a 2134 msnm. Al contrario de C. alvarezi, los otros miembros del género tiene rangos tropicales en tierras bajas. Se distingue fa- cilmente C. alvarezi de sus congéneres por su coloracion, color dorsal uniforme, su vientre sin manchas, y caracteristicas de es- camacion y denticion. Parece que esta es- pecie esta relacionada con C. fissidens que tiene un rango muy extensivo, y se presenta una comparacion detallada de estas dos es- pecies.
Acknowledgments
For their assistance in the field, I thank Barry L. Armstrong, David M. Hillis, and William W. Lamar, who also took the pho- tographs for Figs. 1-3. Iam grateful to John E. Cadle and Charles W. Myers for many helpful comments on the manuscript. Of- ficials of the Direccion General de la Fauna Silvestre issued collecting permits for Mex- ico.
VOLUME 102, NUMBER 4
Literature Cited
Alvarez del Toro, M. 1983 [1982]. Los reptiles de Chiapas. Third edition. Publicacion del Insti- tuto de Historia Natural, Tuxtla Gutiérrez, Chiapas, México. 248 pp.
Bailey, J.R. 1939. Asystematic revision of the snakes of the genus Coniophanes.—Papers from the Michigan Academy of Science, Arts, and Letters 24:1-48.
Campbell, J. A., & J. P. Vannini. 1988. Preliminary checklist of the herpetofauna of Finca El Faro, El] Palmer, Quezaltenango, Guatemala.—Pub- licacion Ocasional, Fundacion Interamericana de Investigacion Tropical 1:1—10.
Conant, R. 1965. Miscellaneous notes and comments on toads, lizards, and snakes from Mexico.— American Museum Novitates 2205:1-38.
Davis, W. B., & J. R. Dixon. 1959. Snakes of the Chilpancingo region, Mexico.— Proceedings of the Biological Society of Washington 72:79-92.
Duellman, W. E. 1965. Amphibians and reptiles from the Yucatan Peninsula, México.— University of Kansas Publications, Museum of Natural His- tory 15(12):577-614.
Hardy, L. M., & R. W. McDiarmid. 1969. The am- phibians and reptiles of Sinaloa, México. — Uni- versity of Kansas Publications, Museum of Nat- ural History 18(3):39-252.
Henderson, R. W., & L. G. Hoevers. 1975. A check- list and key to the amphibians and reptiles of Belize, Central America.— Contributions in Bi- ology and Geology, Milwaukee Public Museum 5:1-63.
Landy, M. J., D. A. Langebartel, E. O. Moll, & H. M. Smith. 1966. Acollection of snakes from Vol- can Tacana, Chiapas, Mexico.—Journal of the Ohio Herpetological Society 5(3):93-101.
Lee, J. C. 1980. An ecogeographic analysis of the herpetofauna of the Yucatan Peninsula. — Mis- cellaneous Publication, Museum of Natural History, University of Kansas 67:1-75.
McCoy, C.J. 1969. Snakes of the genus Coniophanes (Colubridae) from the Yucatan Peninsula, Mex- ico.—Copeia 1969(4):847-849.
Minton, S. A., & H. M. Smith. 1960. A new subspe- cies of Coniophanes fissidens and notes on Cen- tral American amphibians and reptiles. — Her- petologica 16(2):103-111.
Myers, C. W. 1966. A new species of colubrid snake,
genus Coniophanes, from Darién, Panama.—
Copeia 1966(4):665-668.
1969. Snakes of the genus Coniophanes in Panama.— American Museum Novitates 2372: 1-28. Peters, J. A.
1954. The amphibians and reptiles of
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the coast and coastal sierra of Michoacan, Mex- ico.— Occasional Papers of the Museum of Zo- ology, University of Michigan 554:1-37.
—, & B. Orejas-Miranda. 1970. Catalogue of Neotropical squamata. Part 1. Snakes.—Bulle- tin of the United States National Museum 297(1): 1-347.
Schmidt, K. P. 1941. The amphibians and reptiles of British Honduras. — Field Museum of Natural History Publications, Zoological Series 22(8): 475-510.
Shannon, F. A.,& H.M.Smith. 1949. Herpetological results of the University of Illinois field expe- dition, spring 1949. I. Introduction, Testudines, Serpentes.— Transactions of the Kansas Acad- emy of Science 52(4):494—-509.
Shreve, B. 1957. Reptiles and amphibians from the Selva Lacandona. in R. A. Paynter (ed.), Bio- logical investigations in the Selva Lacandona, Chiapas, Mexico.— Bulletin of the Museum of Comparative Zoology 116(4):242-248.
Smith, H. M. 1941. The Mexican subspecies of the
snake Coniophanes fissidens.—Proceedings of
the United States National Museum 91:103-
|
. 1960. Herpetozoa from Tabasco.—Herpe-
tologica 16:222-223.
,&C. Grant. 1958. Noteworthy herptiles from
Jalisco, Mexico.—Herpetologica 14(1):18-23.
—, & L. E. Laufe. 1945. Mexican amphibians and reptiles in the Texas Cooperative Wildlife Collections. — Transactions of the Kansas Acad- emy of Science 48(3):325-354.
—,&E.H. Taylor. 1945. An annotated checklist and key to the snakes of Mexico.— Bulletin of the United States National Museum 187:1-239.
—_,&K.L. Williams. 1963. Newand noteworthy amphibians and reptiles from southern Mexi- co.—Herpetologica 19(1):22-27.
Stuart, L. C. 1963. A checklist of the herpetofauna of Guatemala.— Miscellaneous Publications of the Museum of Zoology, University of Michigan 122:1-150.
Taylor, E. H. 1949. A preliminary account of the
herpetology of the state of San Luis Potosi, Mex-
ico.— The University of Kansas Science Bulletin 33(2):169-215.
. 1953. Fourth contribution to the herpetology
of San Luis Potosi.—The University of Kansas
Science Bulletin 35(13):1587-1614.
Wilson, L. D., & J. R. Meyer. 1985. The snakes of Honduras, second edition.— Milwaukee Public Museum. 150 pp.
Zug, G. R.,S. B. Hedges, & S.Sunkel. 1979. Variation in reproductive parameters of three Neotropical snakes, Coniophanes fissidens, Dipsas catesbyi,
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and Jmantodes cenchoa. —Smithsonian Contri- butions to Zoology 300:1-20.
Department of Biology, The University of Texas at Arlington, Arlington, Texas 76019.
Appendix
Specimens of Coniophanes fissidens ex- amined. — All are in the University of Texas at Arlington Collection of Vertebrates.
Guatemala: Baja Verapaz; vicinity of La Union Barrios, 1500-1600 m (UTA R-16015, 20744); Escuintla: S slope Volcan
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de Agua, Finca Rosario Vista Hermosa (UTA R-4468, 16016, 20691); Izabal: 5.1 km WSW Puerto Santo Tomas, 152m (UTA R-20684); 7.0 km SW Puerto Santo Tomas, 400 m (UTA R-20685); Quezaltenango: S slope Volcan Santa Maria, ca. 4.0 km N El Palmar, 875 m (UTA R-20692-743); Finca El Carmen (UTA R-20686-90).
Mexico: Oaxaca; Cerro Batl, 19 km NW Rizo de Oro (UTA R-12257); 2-3 mi S Ta- panatepec (UTA R-4337, 4339); Veracruz; 2.1 mi NW Sontecomapan (UTA R-3069); 7.7 mi NW Sontecomapan (UTA R-9457, 9468); 5.6 mi ESE Tebanca (UTA R-3067).
PROC. BIOL. SOC. WASH. 102(4), 1989, pp. 1045-1049
THE KARYOTYPE OF EXILIBOA PLACATA BOGERT (TROPIDOPHEIDAE), AND COMPARISONS WITH THE FAMILY BOIDAE (REPTILIA: SERPENTES)
Laurence M. Hardy
Abstract.—The first karyotype for any member of the Tropidopheidae is described from one male and one female of the dwarf boa, Exiliboa placata Bogert, from Oaxaca. The diploid number is 36, composed of 16 macrochro- mosomes and 20 microchromosomes. A possible secondary constriction is present on the second pair of macrochromosomes, and this constitutes the only distinctive difference between this species and some members of the Boidae. The significance of possible differences in centromere positions between ExI- liboa and boids with 36 chromosomes is unknown.
The Tropidopheidae includes four genera according to several recent authors (Under- wood 1976, McDowell 1987): Tropidophis (15 species), Trachyboa (2 species), Unga- liophis (2 species), and Exiliboa (monotyp- ic). Nothing is known of the chromosome morphology for any member of the family. This paper reports the karyotype of the monotypic genus Exiliboa.
Two specimens of Exiliboa placata Bo- gert were available for study: an adult fe- male (UTA R-4731) and an adult male (UTA R-4732). These specimens were col- lected by Jonathan A. Campbell at 7.1 mi. (UTA R-4731) and 6.5 mi. (UTA R-4732), respectively, north of the crest of Cerro Pe- lon, Oaxaca, México, on 28 June 1975.
Chromosomes were prepared by the hy- potonic citrate method of Patton (1967), us- ing the modification by Cole & Leavens (1971). Velban was used instead of colchi- cine. Each macrochromosome was mea- sured (to the closest 0.01 mm) with dial calipers directly on the 4x5’ negative. Chromosome terminology follows Cole (1970). The arrangement of the chromo- somes within the karyotype is based on size, from the largest pair (number one) to the smallest. In addition to the karyotypes pre- sented (Figs. 1, 2), I subjected the measure-
ments of the best 19 cells (seven cells from the female and twelve cells from the male) to computer analysis using the program Ka- rypak (ver. 1.0) by William H. LeGrande (pers. comm.). The macrochromosome means were calculated from each arm of each chromatid. In this analysis only mac- rochromosomes were measured and, for purposes of the karyotype percentages and arm ratio (centromeric index) estimations, they were treated as the entire complement (i.e., microchromosomes were not included as part of the karyotype). This process does not allow for the detection of differences among the microchromosomes nor for the contribution of the microchromosomes to the entire karyotype. However, for most snakes such information on the microchro- mosomes is rarely available and any differ- ences in size are suspect, in most cases, be- cause of the small sizes and poor resolution. Therefore, omission of the microchromo- somes is practically the same as assignment of a constant. I believe that this approach is most effective and reasonable for the crit- ical examination of the macrochromo- somes. Since no sexual dimorphism was de- tected, the male cells were combined with the female cells for the construction of the composite idiogram (Fig. 3).
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qs
g 10 I l2 13
Gn 24 am jt Af A* we Karyotype of an adult male Exiliboa placata (UTA R-4732), 2n = 36.
Fig. 1.
Fifteen cells each from the male and the female were photographed. The karyotype consists of eight pairs of macrochromo- somes and ten pairs of microchromosomes for a diploid number of 36 (Figs. 1, 2). The fundamental number is 56 (30 from macro- chromosomes and 26 from microchromo- somes). The largest macrochromosome pair is metacentric, the second largest pair is sub- metacentric, and pair three is metacentric. These three pairs are clearly distinguishable from all of the other chromosomes. Pairs four and seven are subtelocentric and sim- ilar in morphology, but pair seven is slightly smaller and the short arms are slightly lon- ger (proportionally; Table 1) than the short arms of pair four. Pair five is metacentric, pair six is telocentric, and pair eight is sub- metacentric. All of the macrochromosomes
2 13
Ge 4a @* ae 4&* BF AH
C ) ¢) AK as
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{ GS RR gq ba Ae
M
14 15 16 17 18 e@ At ”&
are easily distinguishable from each other and from all of the microchromosomes (Fig. 3). At least three pairs of the microchro- mosomes appear to be bi-armed; the re- mainder appear to be telocentric, or nearly so. No morphologically distinguishable sex chromosomes are apparent; however, pair five is probably homologous to the ZZ sex chromosomes because it is the only pair of metacentric macrochromosomes that is ap- proximately the same size as the ZZ sex chromosomes identified in members of the Boidae by other workers (Mengden & Stock 1980). All other macrochromosomes of Ex- iliboa are distinctly different in centromere position or in size.
Comparisons with the Boidae. —All species of the Boidae for which karyotypes are known have 36 chromosomes (2n) ex-
BK gp AAAS
14 5 I) 17 18 a & ~ * »*)
Fig. 2. Karyotype of an adult female Exiliboa placata (UTA R-4731), 2n = 36. The arrow indicates a possible
secondary constriction.
VOLUME 102, NUMBER 4
2 | 2 3 4 5 6 7 8
MACROCHROMOSOME NUMBER
PERCENT TOTAL LENGTH
Fig. 3. Composite idiogram of the macrochromo- somes of Exiliboa placata, based on mean measure- ments and arm ratios from 19 cells. Percent total length is calculated from the total length of the macrochro- mosomes in each cell, excluding the ten pairs of mi- crochromosomes. Macrochromosome number is the pair number.
cept Sanzinia madagascarensis (Branch 1980), Acrantophis dumerili (Mengden & Stock 1980), Eryx johni (Singh et al. 1968), and Gonglyophis conicus (Singh et al. 1970), all with 34, Corallus caninus with 44 (Becak 1965), and C. enhydris with 40 (Gorman & Gress 1970).
Diploid numbers other than 36 among boids probably represent derived condi- tions since 36 is the modal number for known boids and is also represented in the primitive Boa (McDowell 1979). In San- zinia madagascarensis there are nine pairs of macrochromosomes, including an extra metacentric (pair four in Mengden & Stock 1980:fig. 10), but only eight pairs of micro- chromosomes, versus ten in Exiliboa. Acrantophis dumerili differs from Exiliboa by having only nine pairs of microchro- mosomes; the macrochromosomes appear indistinguishable except for the telocentric W chromosome in Acrantophis (Mengden & Stock 1980). Eryx johni differs from Exi- liboa by having pair eight telocentric, not submetacentric, and by having only nine pairs of microchromosomes, all of which
1047
Table 1.—A comparison of the macrochromosomes of Exiliboa placata, both sexes combined; n = 19. Per- cent of total is the percent of the total length (in mm, measured from the 4 x 5” negatives) of macrochro- mosomes, excluding the microchromosomes.
Pair Arm lengths
num- Percent
ber Short Long’ Ratio of total Centromere position 1 19D) — 143117 el 262) Metaceninic 2 9.6 14.1 1.47 11.29 Submetacentric 3 135 S143 7.62 Metacentric 4 pe | 6.8 3.24 4.24 Subtelocentric 5 40 4.4 1.10 4.00 Metacentric 6 0.0 78:20:00, +, 3:71 »~Telocentrc 1 1.8 525', 3:06 3.48 Subtelocentric 8 2h 43. 2.05 3.05 Submetacentric
are telocentric (Singh et al. 1968); at least three pairs of microchromosomes in ExI- liboa are bi-armed. The species of Corallus have telocentric macrochromosomes, prob- ably due to centric fission of the first two (C. enhydris) or four (C. caninus) macro- chromosomes. Gonglyophis conicus has one fewer microchromosome (2n = 34; Singh et al. 1970).
All of the remaining boids for which chro- mosome morphology is known have diploid numbers of 36. The macrochromosomes of Exiliboa placata are similar to those re- ported for Liasis by Mengden & Stock (1980) except that pair six of Exiliboa is clearly telocentric and distinguishable from all oth- er pairs, whereas pairs six, seven, and eight of Liasis are telocentric; Mengden & Stock (1980) also identified pair five as ZZ of the sex chromosomes. Python molurus differs from Exiliboa only in the arm ratios of some macrochromosomes (Singh et al. 1968). The karyotype of Xenopeltis unicolor (some- times included in the Boidae) is similar to Exiliboa in number and morphology of chromosomes except that pairs four, seven, and eight have longer short arms than do the apparent homologues in Xenopeltis (Cole & Dowling 1970). Also similar in number and morphology is Loxocemus bicolor (Fischman et al. 1972). In Charina bottae
1048
and Lichanura roseofusca the karyotypes (Gorman & Gress 1970) are extremely sim- ilar to Exiliboa except that the telocentric macrochromosome (pair six in Exiliboa) appears homologous to the smallest macro- chromosome pair in Charina and Lichan- ura and pairs five through seven in Charina and Lichanura are telocentric rather than subtelocentric (pairs four and seven) or even submetacentric (pair eight) as in Exiliboa. The microchromosome morphology was not given by Gorman & Gress (1970) although they did report a fundamental number of 44, which only would result from all of the microchromosomes being treated as telo- centric. Eunectes murinus, Epicrates cen- chria, and Boa constrictor differ from Exi1- liboa mainly by having relatively shorter short arms on macrochromosomes four (pair five in Becak 1965:figs. 1-12), seven, and eight (Becak 1965). Examination of addi- tional tropidopheids is necessary to deter- mine the significance of the above differ- ences.
Even though heteromorphic sex chro- mosomes are not evident in Exiliboa, pair five is probably homologous to the ZZ sex chromosomes identified in some boids (Acrantophis and Liasis by Mengden & Stock 1980). The lack of apparent difference among the cells studied here suggests that the W sex chromosome has undergone little, if any, morphological change if homo- morphic sex chromosomes are primitive in snakes (Becak et al. 1966). This supports the position that Exi/iboa is relatively prim- itive, among snakes in general, and its close relationship to the boids is not unreason- able.
In at least five (three from the male, two from the female) of the photographs of Ex- iliboa chromosomes there is a consistent discontinuity in the basal part of the short arm of chromosome pair two (Fig. 2). That discontinuity is possibly a secondary con- striction; if so, it is the first reported for any member of either the Tropidopheidae or the Boidae.
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
The karyotype of Exiliboa placata is not distinctively different from several species of the Boidae, nor is it distinctive from sev- eral non-boids (i.e., Xenopeltis in the Xen- opeltidae [Cole & Dowling 1970]; several colubrids, except for heteromorphic sex chromosomes in some colubrids). How- ever, consistent differences in centromere positions (i.e., pairs 4, 7, and 8 with longer short arms in Exiliboa) might exist between Exiliboa and some boids with 36 chromo- somes. Based on the karyotype alone, the evolutionary relationships of Exiliboa with- in the Tropidopheidae and the separation of the Tropidopheidae from the Boidae is neither refuted nor supported.
Acknowledgments
I thank John L. Darling and Jonathan A. Campbell for the opportunity to karyotype these specimens. I also thank Charles J. Cole for helpful comments and criticisms during this study. George Steyskal and Hobart M. Smith are gratefully acknowledged for their advice and counsel on taxonomic nomen- clature.
Literature Cited
Becak, Willy. 1965. Constituicao chromoss6mica e mechanismo de determinacao do sexo em ofi- dios Sul-Americanos. I. Aspectos cariotipi- cos.—Memorias do Instituto Butantan (Sao Paulo) 32:37-78.
———.,, Maria Luiza Becak, & Heleneide Nazareth. 1966. Evolution and sex chromosomes in ser- pentes.—Memorias do Instituto Butantan (Sao Paulo) Simposium Internacional 33(1):151—152.
Branch, W. R. 1980. Chromosome morphology of the Madagascar tree boa Sanzinia madagascar- iensis. —South African Journal of Zoology 15(2): 79-82.
Cole, Charles J. 1970. Karyotypes and evolution of the spinosus group of lizards in the genus Sce- loporus.— American Museum Novitates (2431): 1-47.
—, & Herndon G. Dowling. 1970. Chromo- somes of the sunbeam snake, Xenopeltis uni- color Reinwardt (Reptilia: Xenopeltidae).— Herpetological Review 2(2):35.
VOLUME 102, NUMBER 4
, & Carol R. Leavens. 1971. Chromosome preparations of amphibians and reptiles: im- proved technique. — Herpetological Review 3(6): T-102.
Fischman, H. K., J. Mitra, & H. Dowling. 1972. Chromosome characteristics of 13 species in the order Serpentes.—Mammalian Chromosomes Newsletter 13:7-9.
Gorman, George C., & Franklin Gress. 1970. Chro- mosome cytology of four boid snakes and a var- anid lizard, with comments on the cytosystem- atics of primitive snakes. — Herpetologica 26(3): 308-317.
McDowell, S. B. 1979. A catalogue of the snakes of New Guinea and the Solomons, with special reference to those in the Bernice P. Bishop Mu- seum. Part III. Boinae and Acrochordoidea (Reptilia, Serpentes).—Journal of Herpetology 13(1):1-92.
McDowell, Samuel B. 1987. Systematics. Pp. 3-50. in Richard A. Seigel, Joseph T. Collins, and Su- san S. Novak, eds., Snakes: ecology and evo- lutionary biology. MacMillan Publ. Co., New York, 529 pp.
Mengden, Gregory A., & A. Dean Stock. 1980. Chro-
1049
mosomal evolution in serpentes: A comparison of G and C chromosome banding patterns of some colubrid and boid genera.— Chromosoma (Berlin) 79:53-64.
Patton, James L. 1967. Chromosome studies of cer- tain pocket mice, genus Perognathus (Rodentia: Heteromyidae).— Journal of Mammalogy 48(1): 27-37.
Singh, L., T. Sharma, & S. P. Ray-Chaudhuri. 1968.
Chromosomes and the classification of the snakes
of the family Boidae. — Cytogenetics 7:161-168.
; ,& . 1970. Chromosome num-
bers and sex chromosomes in a few Indian species
of amphibia and reptiles. —Mammalian Chro-
mosomes Newsletter 11:91-94.
Underwood, Garth. 1976. A systematic analysis of boid snakes. Pp. 151-175 in A. d’A. Bellairs and C. Barry Cox, eds., Morphology and biology of reptiles. Academic Press, London, 290 pp.
Museum of Life Sciences, Louisiana State University in Shreveport, One University Place, Shreveport, Louisiana 71115.
PROC. BIOL. SOC. WASH. 102(4), 1989, pp. 1050-1066
RHODOGORGON, AN ANAMOLOUS NEW RED ALGAL GENUS FROM THE CARIBBEAN SEA
James N. Norris and Katina E. Bucher
Abstract.— Rhodogorgon (Rhodophyta) a new genus with two species, R. carriebowensis and R. ramosissima, superficially resembling some gorgonian soft corals, is described from the Caribbean Sea. Studies of vegetative mor- phology, male reproductive structure, pigment composition, nature of calcium carbonate, and ultrastructure reveal a combination of characters that is excep- tional among the red algae: thallus with a sharply demarcated cortex of laterally interconnected cortical fascicles and rhizoidal filamentous medulla; three types of cortical cells which develop from the base ofa cortical fascicle— 1, pigmented, assimilatory filaments with inflated, hyaline apical cells, 2, unusual elongate, hyaline, hair-like calciferous cells with inflated tips, and 3, basal hair cells; uninucleate vegetative cells that lack secondary pit-connections and cell fusions; pit-plugs with two cap layers on either side of plugs, the outer caps dome- shaped; and calcite (among the red algae previously known only in the Coralli- nales) confined to single “‘husklike” structures (unique among all algae) that distally surround a calciferous cell. Thalli are apparently dioecious; sperma- tangial parent cells are borne bilaterally on subterminal cells of the cortical filaments, with each cutting off a single spermatangium by oblique division.
Possible taxonomic affinities of the new genus are discussed.
A relatively large, cartilaginous, and pe- culiar red alga has been collected at many Caribbean localities over the past 16 years. Plants are usually rare or sparse in occur- rence and grow in shallow to mid-subtidal depths on rocks or coral heads, in patch reefs, fringing reefs and barrier reefs throughout the year. This alga could have been easily overlooked because of its resem- blance to some gorgonians (Gorgonacea; Anthozoa) in shape, color, and thick car- tilaginous texture (see color photograph of “mystery alga” in Littler et al. 1989:184). It has been noted by F. M. Bayer (octocoral systematist) that the living plants of Rhodo- gorgon carriebowensis superficially resem- ble some species of Carijoa F. Miller (a gorgonian cosmopolitan in subtropical- tropical oceans), and the dried herbarium specimens of R. ramosissima resemble Plu-
migorgia Nutting (a gorgonian from the Indo-Pacific). Initial examination by our phycological colleagues suggested speci- mens could be confused with gorgonians. The presence of pit plugs between the ex- ceedingly small cells, documented by TEM studies (S. Brawley, pers. comm.), demon- strated that the specimens were plants! Materials and methods. —Specimens were collected from the Caribbean Sea, from 1973 to 1989 by skin or SCUBA diving, at depths from 1-25 m. For morphological studies, thalli were pressed fresh or preserved in 5% buffered Formalin/seawater. Collection numbers cited with the prefix JN- or KB- refer to the field notebooks of J. N. Norris or K. E. Bucher, respectively. Live speci- mens were studied in the field at the Smith- sonian Institution’s Carrie Bow Cay Labo- ratory on the barrier reef of Belize, the Galeta
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Marine Laboratory of the Smithsonian Tropical Research Institute (STRI) on the Caribbean coast of the Republic of Panama, and aboard the NSF ships OR/V Cape Flor- ida and OR/V Columbus Iselin.
Microscope slides for anatomical studies were prepared from living or liquid-pre- served specimens by hand sectioning with single- or double-edged razor blades, or by using a Reichert Histostat cryostat micro- tome to make both transverse and longi- tudinal sections of the main axes, branches and apices. Some were acidified with 2—5% HCL to remove calcium carbonate, and then stained with aniline blue and mounted in serial dilutions of clear Karo Syrup with phenol added (as a preservative) following techniques of Tsuda and Abbott (1985), or acidified with 1—5% acetic acid and stained with aceto-iron-hematoxylin-chloral-hy- drate (Wittmann 1965) and mounted in 50: 50 Hoyer’s mounting medium according to the procedure of Hommersand and Fred- ericq (1988). Other preparations of fresh or liquid-preserved specimens were not acidi- fied or stained prior to mounting on micro- scope slides in order to observe the unique calciferous cells.
Living specimens of Rhodogorgon col- lected from patch-reefs in Belize (Carrie Bow Cay) and the Bahamas (Chub Cay) were re- turned to our laboratory and grown in the 511-liter algal reef tank, at 11D:13L pho- toperiod (under 6 Sylvania, 6 ft. VHO-160w fluorescent lights), a water temperature range of 26—-29°C, and salinity range of 35.5- 36.7%. Field collected plants from Carib- bean Panama (San Blas Islands) were grown in the outdoor holding tanks under natural conditions at STRI’s Galeta Marine Labo- ratory from 1979-1984, and in the living coral reef exhibit (7570 liter) at the National Museum of Natural History under ten-1000w multi-vapor halide lights (Adey 1983) with other physical conditions similar to those described above in the 511 liter reef tank.
For ultrastructure studies, branches and main axes of freshly collected specimens
1051
were cut into 0.5—1.0 mm thick pieces and preserved in 5 dram vials of either 4% glu- taraldehyde/seawater or 4% glutaralde- hyde/cacodylate buffer, and then 2% os- mium tetroxide (see previously described procedures in Pueschel 1979, 1980).
Freshly collected plants were frozen and transported in dark bottles covered with aluminum foil for pigment studies. Phyco- biliproteins were extracted in 0.03 M Po- tassium-phosphate (pH 6.8) following the methods described by Gantt et al. (1979).
For calcification analysis, trans-sections 0.3-0.5 mm thick of the axes and branches of living and preserved specimens were placed on microscope slides. Some sections were rinsed in distilled water and others were not, then they were placed in glass petri- plates and dried at 40°C in a Thomas drying oven or air-dried at room temperature. Cal- cium carbonate was analyzed by powder x-ray diffraction studies.
Comparative thin-layer chromatograms (TLC’s) of 90% ETOH extracts of freshly collected specimens of Rhodogorgon carrie- bowensis from Carrie Bow Cay, Belize and R. ramosissima from Antigua were done at the Smithsonian’s Carrie Bow Cay Lab. and aboard the OR/V Cape Florida, following methods of Norris & Fenical (1985). As part of a field research project aboard NSF’s OR/V Columbus Iselin fresh homogenates of R. carriebowensis, collected from Chub Cay, Bahamas, were analyzed in collabo- ration with J. Burgess and R. Jacobs, for the presence of enzymes capable of producing bioactive compounds (i.e., phospholipase A and lipoxygenase). Lipoxygenase activity was measured both polarigraphically using a Clark type electrode and spectrophoto- metrically determining olefin conjugation utilizing arachidonic acid as substrate (Red- dana et al. 1988). Phospholipase activity was measured directly based on procedures of Dagan and Yedgar (1987) using the flu- orescent substrate 1-acyl-2-(N-4-nitroben- zo-2-oxa-1,3-diazole) aminocaproylphos- phatidycholine, and separating the free fatty
1052 P ROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
y
= Re eF% \ jis
{ Pag VV yes a 2 sy
e ‘ \
4 a Ys
ae a i]
fa sie
J
, SS
Figs. 1-3. Rhodo gorgon carriebowensis, f: lax form with br sis, from Carrie Bow Cay, Beli anches to 40 cm in length (US-098364). 3, A ae ee vs Pw oe ee 2, A long ; , lax form (US-098363). i
VOLUME 102, NUMBER 4
acid product by solid phase extraction on C,, cartridges.
Type specimens, including microscope slides and liquid preserved material are de- posited in the Algal Collection of the U.S. National Herbarium, National Museum of Natural History, Smithsonian Institution. Additional cited specimens are deposited in ADU, BISH, MELU, MICH, UC and US (following herbarium designations of Holmgren et al. (1981).
Rhodogorgon J. Norris et Bucher, gen. nov.
Description. — Thallus erectus rosealus vel atroruber, cartilaginogelatinosus, per hap- teron tenue discoideum affixus, laxe vel abunde ramosus; ramis teretibus vel com- pressis. Frondes solitariae aut 2 aut 3 erec- tae orti stipite brevi; axibus principalibus 0.3—0.5 cm crassis, ramulis 2—3 mm crassis. Ramificatio strati corticalis ex fasciculis fi- lorum pseudotrichomatorum aut pseudo- dichomatorum constans; medulla ex filis oblongatis eramosis rhizoidibus constans; rhizoidibus medullaris ab cellulis interior- ibus corticalibus ortis. Cellulae corticales et medullosae uninucleatae, conjunctionibus vegetativis et synapsibus secundis absenti- bus. Synapses primariae obturamentae cum duobus stratis capitularibus continentes, exterioribus tholiformibus. Extensio cellu- larum calcifera protoplasmica hyalina es- egmentata cellulis interioribus fasciculorum corticalium orta, ad apicem cylindrica cal- cifera (calcite). Cellulae parentes sperma- tangiorum bilateraliter cellula subterminali filorum corticalium ortae, spermatangio singulo ferentes.
The thalli are erect, cartilaginous, slip- pery, locally lightly calcified, with terete to compressed upright main axes and side branches. Usually a single (rarely more) short stipe arises from a small, discoid holdfast. The thalli may be stringy and sparsely branched to a few orders, or compact and densely branched to several orders (Figs. 1- 7). Branching is mostly irregular, or alter-
1053
nate, or more or less radial, or occasionally pinnate, and tends to become secund to- ward the apices.
The thalli apparently are multiaxial, con- sisting of two distinct regions, with a sharp boundary (Figs. 8, 9) between the pigment- ed, fasciculate cortical layer and the medulla of unpigmented, intertwined rhizoidal fil- aments (Figs. 8, 9). Both cortical cells (Figs. 11-13) and medullary cells (Fig. 17) are uni- nucleate and lack secondary pit-connec- tions and cell fusions. The assimilatory fil- aments of the cortex are organized into fascicles that are radially interconnected at their base (Figs. 10, 11). The filamentous medulla is composed of hyaline, thick- walled rhizoidal filaments that interlace. One or two unpigmented medullary rhizoidal fil- aments are cut off secondarily from the in- nermost cortical cell bearing a cortical fas- cicle (Figs. 10-13). These rhizoidal filaments continue to grow inward, contributing to the structure of the medulla.
The branching pattern of a cortical fas- cicle is typically pseudotrichotomous, and there are three types of cortical structures. The first are filaments composed of pig- mented, granular, cylindrical cells that ter- minate in an inflated hyaline cell (Figs. 16, 18, 21, 23-26). The second type are basal hair cells (Figs. 18-20), and the third are unique, elongate, unsegmented, hyaline, hair-like calciferous cells, which are distally surrounded by a brownish, “husk-like”’ cal- careous structure (Figs. 21-26). A new branch within a cortical fascicle originates when a cell of a pigmented filament buds part of its cytoplasm distally (Figs. 13, 14). Subsequent septation of the protruded cy- toplasm (Fig. 15) followed by cell division towards the thallus surface results in a new branch. The septum develops horizontally (Fig. 15) or slightly obliquely. The pseu- dotrichotomy of the cortical fascicle results when a cell that bears a pseudodichotomy buds (Fig. 13), septates, and undergoes cell division. Concordantly, a pseudodichoto- my originates when the bearing cell of an unbranched cortical filament protrudes and
1054 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
VOLUME 102, NUMBER 4
divides (Figs. 14, 15). Well developed branches have a central branching point that has cut off rhizoids and bears two orders of trichotomies (Fig. 13). An unbranched cor- tical filament usually consists of 4—6 cells (Figs. 10, 11).
Cell division does not always produce a new side branch, but may stop after cutting off one cell that becomes the basal cell of a hair (Figs. 18, 19). Cytological transfor- mation of such a basal cell is often accom- panied by breakdown of its cell wall ma- terial, revealing persistent putative cellulose fibrils (Figs. 18, 19). Once the basal cell pri- mordium becomes dense with cytoplasm and increases in size (Fig. 19), it cuts off a similarly darkly staining hair (Fig. 20) that protrudes beyond the cortical surface. If the hair is broken off, the pit-connection be- tween the cells persists.
The specialized, brownish, “‘husk-like”’ structures of the calciferous cells are the lo- calized sites of extracellular calcite precip- itation. When dilute acid is pipetted under- neath the coverslip, the calcite readily dissolves, revealing a hyaline, small to large, inflated tip on the elongate, unsegmented, hair-like cell. These hair-like cells are very thin, vacuolate protoplasmic extensions that are cut off and pit-connected to the base of a pseudotrichotomy (Figs. 24, 27) or pseu- dodichotomy (Fig. 26). The calcite com- pletely surrounds the apical portion of the calciferous hair-like cell when it is narrow (Fig. 21), but as its tip inflates a longitudinal furrow (Figs. 22, 23) forms, and the narrow extension slightly expands in width (Figs. 24—26). Occasionally a very narrow channel forms within the cell wall of the hyaline, calciferous extension (Fig. 22). The calcite ““husk-like”’ structures are scattered among
— Figs. 4—S.
1055
and located just beneath the zone of the terminal inflated cells of the cortical fila- ments (Figs. 8, 9, 21, 23, 26). These unique calcite structures vary in abundance within the cortical layer and give a greyish sheen to the thallus. The calciferous cells are com- monly one per fascicle, but up to three may be cut off. Segmentation within these cal- ciferous cells was never observed, although their apical portions were occasionally sep- arated from the remaining portion of the cell by a lenticular wall (Fig. 28).
Plants are apparently dioecious. In sper- matangium bearing gametophytes, the cor- tex consists of a continuous zone of sper- matangial parent cells (Fig. 29). Spermatangial parent cells are borne bilat- erally on the subterminal cell of a cortical filament (Fig. 30), each cutting off a single spermatangium (Fig. 31) by oblique divi- sion.
Remarks. — Rhodogorgon is named for its resemblance to the branching soft corals, the gorgonians (Gorgonacea). Rhodo- means red and -gorgon refers to Gorgon, a figure in Greek mythology (Genaust 1976), and the other gorgons, Euryale, Steno and Me- dusa, who had hair of snakes.
Type species.—Rhodogorgon carriebow- ensis.
Key to the species of Rhodogorgon
1. Plants loose, sparingly branched to 2-3 orders, with determinate and indeterminate ultimate branches . . Maik Seach eee R. carriebowensis — Plants more compact, densely branched to 3—5 orders, with deter- minate ultimate branchlets R. ramosissima
ee, ee fee Ye me oe eee fe, woe (a 0 ey es
Variation in branching of Rhodogorgon carriebowensis. 4, Specimen from Islas San Blas, Panama,
with both long and short branches (US-098366). 5, Specimen from Passe du Marin, off Pte. Borgnesse, Martinique
(US-098367).
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
1056
Ob. : «
4 ie, 2 .. Ve Y — af nt aS -® WwW we’ .' ee we i RS.
gg ware % v
; “>
-
ye? \ -@ e oe. ‘ WN a” Vath a jen 2754 es
| 7 a
——_ 2 oe
varricuriuticitvicitutlinalantTinlunt
Figs.6-7. Rhodogorgon ramosissima. 6, Holotype from Carlisle Bay, Antigua (US-098361). 7, A more openly
branched form, from Pte. Borgnesse, Martinique (US-098365).
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Rhodogorgon carriebowensis J. Norris et Bucher, sp. nov. Figs. 1-5, 8-31
Diagnosis. —Thallus usque ad 50 cm, in- ferne atropurpureus, superne aurantio-ro- sealus. Frondes solitariae aut raro 2 aut 3, sparse vel profuse 1-2(-3)-plo ramosae; ax- ibus ad 1.0 cm diam.; ramis saepe longis teretibus ad 6 mm diam. et 9.0—13.5(-40) cm longis; cellulis medullosis plusminusve 4.5 um diam. et ad 90 um longis; exten- sionibus calciferis 1-4 wm diam. et 40-65 um longis, ad apicem calciferum 10 wm diam. et 25 um longis.
The thalli are cartilaginous, slippery, to 50 cm tall, dark purple to light peach, usu- ally darker below and lighter above, with a greyish tint throughout. A single (rarely two or three), short stipe grades into terete to compressed axes, to 1.0 cm diam. The axes branch irregularly, alternately, or occasion- ally more or less radially up to three orders. The branches are terete, short to long, to 6 mm diam. and up to 40 cm long (Fig. 2), with blunt ultimate branches to 4 mm diam.
The cortex in cross section is 70-105 wm wide and distinctly separate from the me- dulla. Cortical filaments are composed of pigmented, granular, cylindrical cells, 1.5— 6.0 wm diam. by 12-18 um long, that ter- minate in bulbose, hyaline cells, 6-10 wm diam. by 9-14 um long. The hyaline, cal- ciferous cells are 14 um diam. by 40-65 um long, with a swollen tip, and bear unique, calcite structures distally, 10 um diam. by 25 wm long. The medulla is composed of intertwined, hyaline, thin cells, 4 wm diam. and mostly to 90 um long. All other char- acteristics are given above in the generic description.
Remarks. — Rhodogorgon carriebowensis resembles some species of the gorgonian Carijoa. It is named after Carrie Bow Cay, the type locality and site of the Smithson- ian’s Caribbean Coral Reef Ecosystem Pro- gram (CCRE) on the Belizean Barrier Reef. This species differs from R. ramosissima in
1057
being less densely branched and only to two or three orders, with the upper branches in some tending to be secund. The branches, including the ultimate branchlets, are either determinate or indeterminate, and some- times are very long, to 40 cm. See R. ra- mosissima for other differences.
Type. —Carrie Bow Cay, Belizean Barrier Reef, Belize, spur and groove zone, 4.6—12.2 m depth, 1 May 1979, K. E. Bucher, JN- 7520 (holotype: Alg. Coll. #US-098360).
Distribution. —Caribbean Sea: Bahamas, St. Croix, Belize, Martinique, Panama.
Paratypes. —Caribbean Sea: BAHA- MAS.—Chub Cay, 4 m depth, among cor- als, 12 Jun 1989, R. Sims, s. n. (US). U.S. VIRGIN ISLANDS. —St. Croix: Boiler Bay, growing on fore-pavement in front of boiler, 3—4 m depth, 19 Aug 1978, W. Adey s. n. (US); Tague Bay, patch reef, 2.4 m depth, 6 Jan 1973, C. Bowman, IAA-11592 (MELU), 5 Apr 1973, P. Adey, IAA-11447a & b (BISH, US), and patch reef, 25 Jan 1974, R. Burke & R. Steneck, IAA-11783 (MICH, US). BELIZE. — W of Carrie Bow Cay, patch reef, among gorgonians and corals, 4.6-6.1 m depth, 25 Nov 1980, K. Bucher & R. Sims, JN-10670 (UC); off S end of Carrie Bow Cay, on coral head, 4.5 m depth, 30 Mar 1980, M. Hay, RHS-80-275, and 6 m depth, 9 Apr 1980, R. Sims, s. n. (MICH); SW of Carrie Bow Cay, 3.0 m depth, 25 Mar 1980, M. Hay, s. n. (UC, US), and on coral rubble, patch reef, 6.1—9.1 m depth, 1 Apr 1985, M. Littler, JN-12477 (US), and under coral head, patch reef, 5—8 m depth, 5 Apr 1989, J. Norris, K. Bucher & C. Pues- chel, JN-16241 (US); SE of Carrie Bow Cay, patch reef, 4.6 m depth, 28 Apr 1980, R. Sims, s. n. (US); vic. of Wee Wee Cay, patch reef, 1.5—7.6 m depth, 25 Nov 1980, K. Bucher, R. Sims, P. Taylor & M. Littler, JN-10138 (US); Blue Ground Range, on Ac- ropora palmata, 0.3-—1.5 m depth, 10 Apr 1985, C. Tanner, JN-14830 (US). MAR- TINIQUE. — between Ilet au Chiens and Pte. Ferre, 8 m depth, 25 Aug 1985, K. Bucher & B. Brooks, KB-1687 (US); Pte. Borgnesse,
1058 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
-f if 4 WANs / bs
WI % Dy? 50 um
Figs. 8-13. Anatomy of Rhodogorgon carriebowensis. 8, Transverse section through a third order branch showing cortical cells (c) sharply demarcated from the medullary region (m) of intertwined rhizoidal filaments (unstained, Nomarski). 9, Close-up of Fig. 8 showing junction between cortex and medulla (unstained, Nomarski). 10-11, Assimilatory filaments organized into cortical fascicles (cf) radially interconnected at their base (arrow), and rhizoidal filaments (arrowhead) cut off from innermost cortical cell bearing fascicle (Nomarski). 12, Close- up of base of cortical fascicle, showing pseudotrichotomous branching (arrow), uninucleate cortical cells and
VOLUME 102, NUMBER 4
3.0-18.3 m depth, 23 Aug 1985, M. Hay, KB-1264 (US), to 12.2 m depth, 24 Aug 1985, M. Hay & L. Fisher, KB-1645 (US), and 5 m depth, 29 Aug 1985, M. Hay & K. Gustafson, KB-768 (US); Passe du Marin, off Pte. Borgnesse, 3.0-12.2 m depth, 28 Aug 1985, K. Bucher & B. Brooks, KB-1438 (ADU, MELU, UC, US), and 12 m depth, 28 Aug 1985, M. Hay & L. Fisher, KB-1447 (BISH, MICH, US). PANAMA.—Galeta Reef, N of STRI’s Galeta Lab., 1 Jul 1978, M. Hay, MH-193 (US); San Blas Islands, SW side of Sail Rock, 6.1 m depth, 8 Aug 1979, J. Norris, s. n. (US).
Rhodogorgon ramosissima J. Norris et Bucher, sp. nov. Figs. 6—7
Diagnosis. —Thallus compactus ad 30 cm, obscure canescens vel atrovirens, plusmi- nusve radialiter et profuse ad 5-plo ramo- sus; ramis distaliter tenuiorious ultimis brevis in diametro uniformibus. Cortex as 105 wm diam.; cellulis medullosis 3—4 um latis et plerumque 140 um longis; cylindricis calciferis plusminusve 27 wm longa et 12 pm diam.
The thalli of R. ramosissima are usually more compact and densely, more or less radially, branched to five orders, with the branches becoming progressively smaller outwards. The ultimate branchlets are short, of uniform diameter and have blunt apices. The color of R. ramosissima is generally darker, dark grey to blackish-green; the me- dulla and cortex are distinct, with the cor- tical layer to 105 um wide; medullary cells 3-4 wm in diam. and mostly 110-150 um long; the calcite structures of the calciferous cells are 12 um diam. by 27 um long.
Remarks.—This species differs from R. carriebowensis primarily in habit, being
_
1059
more densely branched, up to five orders. Branching tends to be radial, with the branches becoming progressively smaller and shorter outwards. The ultimate branch- es are short, apparently determinate, and of uniform diameter with blunt apices. The specific epiphet, ramosissima, is derived from ramosus, full of branches, and -issimus (adjectival superlative) meaning very or most. The air-dried herbarium specimens of R. ramosissima are similar to the gor- gonian, Plumigorgia.
Type. —Carlisle Bay, Antigua, Lesser An- tilles, on rocks, 0.5-6.1 m depth, 21 Aug 1985, K. E. Bucher & B. L. Brooks KB- 1271a (holotype: Alg. Coll. US-098361; iso- type: Alg. Coll. US-098362).
Distribution. —Caribbean Sea: Antigua and Martinique, Lesser Antilles.
Paratypes —Caribbean Sea: ANTI- GUA.—Carlisle Bay, 0.5-6.0 m depth on rocks, 21 Aug 1985, K. Bucher & B. Brooks, KB-1271b (ADU, BISH, MELU, MICH); vic. Cade Reef, 10.7 m depth, 21 Aug 1985, K. Bucher, B. Brooks, & W. Fenical, KB- 1551 (US). MARTINIQUE—near Petite Martinique in Havre du Robert, 1.5-3.0 m depth on rocks, 25 Aug 1985, W. Fenical, KB-1691 (MICH, UC, US); Pte. Borgnesse, 12.2 m depth, 24 Aug 1985, M. Hay & L. Fisher, KB-1647 (US); Ilet Rainville, 6.1 m depth, 26 Aug 1985, M. Hay, KB-699 (US).
Results. —The absorption spectrum peaks of the red algal phycobilisomes (Gantt 1981) of R. carriebowensis are 497 nm and 565 nm, indicating R-phycoerythrin as known only in the red algae. The fluorescence spec- trum peak is 578 nm, indicating phycoer- ythrins as found in both red and blue-green algae. Comparative thin layer chromato- graphs revealed no apparent unusual sec- ondary metabolites in the lipid extract. This suggests that Rhodogorgon is not chemically
medullary rhizoidal filaments (arrowhead) (hematoxylin stained). 13, Cell within cortical filament budding off part of its cytoplasm distally (arrow) leading to pseudotrichotomy formation, and medullary rhizoidal filaments (arrowhead) cut off from base of cortical fascicle (hematoxylin stained).
1060 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Figs. 14-22. Anatomy of Rhodogorgon carriebowensis. 14, Cell within cortical filament budding off part of its cytoplasm distally (arrow) leading to pseudotrichotomy formation (hematoxylin stained). 15, Septation (arrow) of protruded cytoplasm leading to new filament is laid down horizontally (hematoxylin stained). 16, Thick-
VOLUME 102, NUMBER 4
defended (sensu Norris & Fenical 1982) against herbivory. Because of its resem- blance to certain soft corals, we suggest it may elude predation as a gorgonian mimic.
Recently, hydroxy fatty acids chemically related to mammalian prostaglandins and leukotrienes have been isolated from the tropical red alga Platysiphonia miniata (Moghaddam et al. 1989). Studies on fresh homogenates of R. carriebowensis showed it apparently contains significant lipoxygen- ase activity resulting in the formation of a product with a UV absorbance spectrum indicative of conjucated diene. Rhodogor- gon also appeared to contain measurable phospholipase A activity. The presence of these two enzymes suggests that Rhodogor- gon may be capable of producing biologi- cally active eicosanoid-like compounds (metabolites of arachidonic acid), thus far only known in these red algae.
The life history of Rhodogorgon is pres- ently unknown. Because it is infrequently encountered and usually sparse where found, it may have a microscopic, filamentous or other hetermorphoric alternate not yet rec- ognized. No tetrasporophytes of Rhodogor- gon have been field collected, although they and its life history would likely have sys- tematic implications.
Discussion.—A striking feature of Rho- dogorgon is the presence of localized calcite deposits that envelop unsegmented, elon- gate, hair-like cells with inflated tips. These calciferous cells are cut off from the base of cortical fascicles and are not known to occur in any other alga. In the calcareous red algae, calcite is an unusual mineral form of cal- cium carbonate. The Corallinales (Silva &
—
1061
Johansen 1986) are the only other red algae known to precipitate calcium carbonate in this form; all other known calcified red algae possess aragonite (Borowitzka et al. 1974, Littler 1976). The Corallinales are consid- ered to be an ancient group, having been found in limestone deposits from the late Cretaceous (Littler 1972), and extending as far back as the Jurassic (Johansen 1981). If calcite is the ancestral mode of calcium car- bonate precipitation, perhaps Rhodogorgon is also a very old taxon.
The function of the calcite structures borne on the calciferous cells of the cortical fas- cicles is still unknown. Because the calcite is localized and only in the apical region, the cells may be involved in secondary branch formation and contribute to thallus structure. They could also play a role in nutrient boundary layer breakdown, or may be herbivore deterrents.
Ultrastructural features of pit-plugs have been useful in postulating phylogenetic af- finites at ordinal levels (Pueschel & Cole 1982, Pueschel 1989). Presence of a dome- shaped outer cap layer on the pit plug is, besides Rhodogorgon, only reported in the Corallinales, Batrachospermales and some Acrochaetiales (Pueschel 1989). Since Rho- dogorgon shares morphological character- istics with the former two orders, it will be briefly compared with them for possible taxonomic affinities. Although Rhodogor- gon shares pit plug characteristics and cal- cite with the Corallinales, their vegetative and reproductive morphologies are very dif- ferent. Rhodogorgon is unique among all the algae in the location and specialized struc- tures of calcium carbonate deposits (for
walled, vacuolate, inflated mature terminal cells of assimilatory filaments; chloroplasts of intercalary cortical cells are parietal (unstained, Nomarski). 17, Close-up of thick-walled, medullary rhizoidal filament showing pit- connection and pit ring (arrowhead) (hematoxylin stained). 18, Basal cell (bh) of hair erupting cellulose fibrils from cell wall (unstained, Nomarski). 19, Darkly stained basal cell (bh) that will develop a hair cell (hematoxylin stained, Nomarski). 20, Densely staining hair cell (h) pit-connected to its basal cell (bh) (hematoxylin stained, Nomarski). 21, Calcite surrounded apices (arrowheads) of elongate, calciferous cells embedded in cortex (un- stained, Nomarski). 22, Narrow channel within wall of a calciferous cell (unstained).
1062 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
1
Figs. 23-31. Anatomy of Rhodogorgon carriebowensis. 23, Hyaline, elongate, calciferous cells with calcite structures (arrowhead) located among the cortical filaments (unstained, Nomarski). 24, Elongate calciferous cell, bearing calcite structure (arrowhead), that is pit-connected (arrow) to base of pseudotrichotomy (unstained). 25, Formation of a longitudinal furrow within calcite covered apex (arrowhead), and emergence of inflated tip of a calciferous cell (unstained). 26, Calciferous cell at base of a pseudodichotomy, and partially dissolved calcite deposit (arrowhead) (unstained, Nomarski). 27, Pseudotrichotomy with basally pit-connected (arrowhead) rem- nant of a calciferous cell (hematoxylin stained). 28, Apical portion of a calciferous cell which is separated from
VOLUME 102, NUMBER 4
summary of calcification in the algae, see Littler & Littler 1984). The domoid expan- sion of the outer cap layer on the pit plug may be a primitive trait. If possession of a dome-shaped outer cap is ancestral, it may again indicate that Rhodogorgon is a very old taxon.
Among the Batrachospermales, Rhodo- gorgon is morphologically most similar to Thorea in the Thoreaceae, a family recently transferred from the Nemaliales to the Ba- trachospermales (Pueschel & Cole 1982). In contrast to the Thoreaceae, other members of the Batrachospermales are uniaxial and composed of axial filaments of indetermi- nate growth surrounded by whorled lateral filaments of limited growth (Aghajanian & Hommersand 1980). The Thoreaceae are multiaxial (Swale 1962, 1963; Yoshizaki 1986). In both Rhodogorgon and Thorea the basal cells of assimilatory branches produce rhizoidal filaments that contribute to the structure of the medulla. However, their cortical fascicles differ, being basically pseu- dotrichotomous in Rhodogorgon and pseu- dodichotomous in Thorea. Also, Rhodo- gorgon is marine and calcified, whereas the Thoreaceae are exclusively freshwater (Sheath 1984) and lack calcification. Inter- estingly, two different types of carpospo- rophyte development have been described for two species currently placed in Thorea. In T. bachmannii Pujals ex Pujals from Bra- zil, Necchi (1987) illustrated a compact car- posporophyte, whereas Yoshizaki (1986) observed a diffuse carposporophyte in T. okadai Yamada from Japan. These differ- ences could indicate that the Thoreaceae may be polyphyletic, and that some mem- bers of the family (e.g., 7. okadai) may be
—_—
1063
more closely related to some of the Nemali- ales, such as Dotyophycus (Abbott & Yoshi- zaki 1981) and Yamadaella (Abbott 1970), and others, such as 7. bachmannii, to the Batrachospermales. So far, only J. riekei Bischoff (1965) has been investigated for pit plug morphology, and if 7. okadai lacks an outer dome pit plug cap, it would be more related to the Nemaliales. The systematic position of Thorea needs to be critically reinvestigated.
Rhodogorgon shares certain anatomical similarities with some members of the Ga- laxauraceae Parkinson and Liagoraceae Kutzing of the Nemaliales. Although both families contain some calcareous members, they possess only aragonite (Borowitzka et al. 1974, Littler 1976, Okazaki et al. 1982). Apical depressions containing an apical cell that directs cell growth in members of Ga- laxauraceae were not seen in Rhodogorgon, but the presence of intertwined medullary filaments is reminiscent of Galaxaura and Scinaia. In the latter genera, basal cells of the cortical filaments cut off rhizoids and the process of cortex differentiation 1s main- ly one of vacuolization accompanied by in- flation in the terminal utricles (see Ramus 1969, for Scinaia, as ‘Pseudogloiophloea’). One could envision that appression of the utricles in the Galaxauraceae is a more ad- vanced trait than the non-appressed ter- minal inflations of Rhodogorgon. There are also reproductive differences. The maie re- productive structures are very simple in Rhodogorgon, while in some Galaxauraceae (i.e., Galaxaura, Actinotrichia, Nothogenia) the spermatangial parent cells are organized within specialized conceptacles (Svedelius 1939, 1943; Magruder 1984); the other gen-
the remaining portion by a lenticular wall (arrowhead), observed after dissolution of the calcite (hematoxylin stained). 29, Cortex with spermatangial parent cells and spermatangia (hematoxylin stained, Nomarski). 30, Pair of spermatangial parent cells (arrows) borne bilaterally on subterminal cell of cortical filament (hematoxylin stained). 31, Spermatangium (arrowhead) cut off singly from a spermatangial parent cell (arrow) (hematoxylin
stained).
1064
era produce spermatangia at the thallus sur- face (e.g., Huisman 1986 for Scinaia). The spermatangial configuration of Rhodogor- gon is certainly simpler than in the calcified members of the Galaxauraceae.
In the Nemaliaceae (Farlow) DeToni et Levi and the Liagoraceae, main axes and branches are of the multiaxial ‘Springbrun- nen-type’ (Oltmanns 1922), and thus very different from the vegetative anatomy of Rhodogorgon which lacks a central core of axial medullary filaments. The origin of spermatangial parent cells in Rhodogorgon is nevertheless reminiscent of that of Ya- madaella (Liagoraceae), where spermatan- gial parent cells are borne on subterminal cells of cortical filaments (Abbott 1970). Spermatangial parent cells in Yamadaella frequently cut off a short chain of sperma- tangia (Abbott 1970), whereas those of Rhodogorgon were only seen to cut off a single spermatangium. Yamadaella and Rhodogorgon both have inflated terminal cells on their assimilatory filaments.
Conclusions. — Based on vegetative and reproductive anatomy, our light-micro- scope studies suggest that Rhodogorgon ex- hibits similarities with some Thoreaceae, Galaxauraceae and Liagoraceae, whereas based on pit plug morphology, it shows re- lationships with Thorea and Nemalionopsis of the Thoreaceae (Pueschel & Cole 1982, Pueschel 1989). However, it differs in sev- eral characteristics from all of these fami- lies, and based on its combination of un- usual characters, we are of the opinion that the new genus is only ancestrally related to the families mentioned above, or via con- vergence has evolved some similiar char- acteristics. Rhodogorgon 1s hypothesized to be a primitive red alga that may have orig- inated before the families of the Nemaliales were present and diversified, and so we be- lieve it would be better placed in a family of its own. However, this assessment must await more detailed studies on the devel- opment of the female reproductive system, life history, and ultrastructural character- istics.
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Acknowledgments
We thank D. E. Appleman for the powder x-ray diffraction analysis, E. Gantt for pig- ment analysis, S. Brawley for the initial TEM preparations, C. M. Pueschel for TEM stud- ies documenting the pit-plug types, W. Fen- ical for the comparative thin-layer chro- matograms, and J. R. Burgess and R. S. Jacobs for the enzyme analysis (NOAA-Sea Grant Prog. No. NA-85AA-D-SG140). We are grateful for the collections and/or diving assistance of I. A. Abbott, P. J. Adey, W. H. Adey, C. Bowman, B. L. Brooks, J. R. Burgess, R. Burke, L. D. Coen, J. L. Connor, W. H. Fenical, L. Fisher, K. Gustafson, M. E. Hale, M. E. Hay, B. E. LaPointe, D. S. Littler, M. M. Littler, O. J. McConnell, J. A. Norris, R. H. Sims, R. S. Steneck, C. E. Tanner, and P. L. Taylor; and to D. L. Bal- lantine, G. Bula Meyer, P. S. Dixon, M. H. Hommersand, B. Santelices, R. B. Searles, W.R. Taylor, C. K. Tseng and J. A. West who in addition to those above, examined specimens and shared comments with us over the years. We thank S. Fredericq who has assisted us with observations, photo- micrography, and with D. B. Lellinger the Latin diagnoses. Our thanks to W. H. Adey, D. L. Ballantine, S. Fredericq, M. H. Hom- mersand, M. M. Littler, D. S. Littler and H. E. Robinson for their review and comments on various stages of the manuscript. F. M. Bayer and T. E. Coffer provided interesting discussions on the gorgonians and their re- semblance to our new genus. Support for the STRI R/V Benjamin cruise to the San Blas Islands (Caribbean Panama) came from a Smithsonian Environmental Science Pro- gram grant to J. Cubit and J. Norris. Travel support from CCRE enabled us to partici- pate on the NSF Lesser Antilles cruise of OR/V Cape Florida (NSF fund #CHE-86- 20217; W. Fenical, Chief Scientist). The Smithsonian’s Research Opportunity Fund supported our participation on the NSF Ba- hamas expeditions aboard the OR/V Co- lumbus Iselin (both NSF fund #CHE-86- 20217; W. Fenical, Chief Scientist). We
VOLUME 102, NUMBER 4
thank K. Ritzler (Program Coordinator, CCRE) for his interest and support of this study at the Smithsonian’s Carrie Bow Cay Lab., Belize. This study represents contri- bution number 276 of the Smithsonian In- stitution’s Caribbean Coral Reef Ecosystem Program (CCRE), partially supported by the Exxon Corporation.
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Adey, W. H. 1983. The microcosm: A new tool for reef research.—Coral Reefs 1:193-201.
Aghajanian, J. G., & M. H. Hommersand. 1980. Growth and differentiation of axial and lateral filaments in Batrachospermum sirodotii. —Jour- nal of Phycology 16:15-28.
Bischoff, H. W. 1965. Thorea riekei sp. nov. and related species.—Journal of Phycology 1:111- Ti.
Borowitzka, M. A., A. W. D. Larkum, & C. E. Nock- olds. 1974. A scanning electron microscope study of the structure and organization of the calcium carbonate deposits of algae.—Phyco- logia 13:195—203.
Dagan, A., & S. Yedgar. 1987. A facile method for direct determination of phospholipase A, activ- ity in intact cells.—Biochemistry International 15:801-808.
Gantt, E. 1981. Phycobilisomes.— Annual Review of Plant Physiology 32:327-347.
—.,C. A. Lipschultz, J. Grabowski, & B. K. Zim- merman. 1979. Phycobilisomes from blue- green and red algae.— Plant Physiology 63:615— 620.
Genaust, H. 1976. Etymologisches Worterbuch der botanischen Pflanzennamen. Birhauser, Stutt- gart, 390 pp.
Holmgren, P. K., W. Keuken, & E. K. Schofield. 1981. Index Herbariorum, I. The herbaria of the world, 7th ed. Bohn, Scheltema, & Holkema, Utrecht, 452 pp.
Hommersand, M. H., & S. Fredericq. 1988. An in- vestigation of cystocarp development in Gelid- ium pteridifolium with a revised description of the Gelidiales (Rhodophyta).—Phycologia 27: 254-272.
Huisman, J. M. 1986. The red algal genus Scinaia
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(Galaxauraceae, Nemaliales) from Australia.— Phycologia 25(3):27 1-296.
Johansen, H. W. 1981. Coralline algae, a first syn- thesis. CRC Press, Boca Raton, Florida, 239 pp.
Littler, D. S., M. M. Littler, K. E. Bucher, & J. N. Norris. 1989. Marine plants of the Caribbean. Smithsonian Institution Press, Washington, D. C., villi + 263 pp.
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view 10:311-347.
1976. Calcification and its role among the macroalgae.— Micronesica 12:27-41.
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biogenis: The contribution of algae. Pp. 323-
364 in F. E. Round and D. J. Chapman, eds.,
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press Ltd., Bristol.
Magruder, W.H. 1984. Reproduction and life history of the red alga Galaxaura oblongata (Nemali- ales, Galaxauraceae).—Journal of Phycology 20(3):402-409.
Moghaddam, M. F., W. H. Gerwick, & D. L. Ballan- tine. 1989. Discovery of 12-(S)-hydroxt- 5,8, 10,14-icosatetraenoic acis [12-(S)-HETE] in the tropical red alga Platysiphonia miniata.— Prostaglandina 37:303-—308.
Necchi, O. 1987. Sexual reproduction in Thorea Bory (Rhodophyta, Thoreaceae).—Japanese Journal of Phycology (Sérui) 35:106-112.
Norris, J. N., & W. H. Fenical. 1982. Chemical de- fense in tropical marine algae. Pp. 417-431 in K. Ritzler and I. G. Macintyre, eds., The At- lantic Barrier Reef ecosystem at Carrie Bow Cay, Belize, I: Structure and communities. —Smith- sonian Contributions to the Marine Sciences, b2:
——,, & . 1985. Natural products chemistry: Uses in ecology and systematics. Pp. 121-145 in M. M. Littler and D. S. Littler, eds., Hand- book of phycological methods, vol. 4, Ecological field methods: Macroalgae. Cambridge Univer- sity Press, Cambridge/New York.
Okazaki, M., K. Ichikawa, & F. Furuya. 1982. Stud- ies on calcium carbonate deposition of algae, IV. Initial calcification site of the calcareous red alga Galaxaura fastigiata Decaisne.— Botanica Marina 25:511-517.
Oltmanns, F. 1922. Morphologie und Biologie der Algen. Vol. 2. Gustav Fischer, Jena, iv + 439 pp.
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Reddanna, P., J. Whelan, P. S. Reddy, & C. C. Reddy. 1988. Isolation and characterization of 5-lipox- ygenase from tulip bulbs.—Biochemical and Biophysics Research Communications 157: 1348-1351.
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liche Studien tiber Galaxaura eine diplobion- tische Nemalionales-Gattung. — Nova Acta Re- giae Societatis Scientiarum Upsaliensis, IV, 13(4):1-154.
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Tsuda, R. T., & I. A. Abbott. 1985. Collection, han- dling, preservation, and logistics. Pp. 67-86 in M. M. Littler and D. S. Littler, eds., Handbook of phycological methods, Vol. 4, Ecological field methods: Macroalgae. Cambridge University Press, Cambridge/New York.
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Department of Botany, National Mu- seum of Natural History, Smithsonian In- stitution, Washington, D.C. 20560.
PROC. BIOL. SOC. WASH. 102(4), 1989, pp. 1067-1068
THE HIGH FREQUENCY OF ERRORS IN FORMAT IN MANUSCRIPTS SUBMITTED TO PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Thomas E. Bowman
Abstract. — Failures to conform to 25 items of Proceedings style are enumer- ated for 40 manuscripts submitted for publication. Up to 15 errors/manuscript were found, with a mean of 4.85 errors/manuscript. Authors are urged to
improve this record.
Having recently reached the 7th decade of my life and having served as Associate Editor for Invertebrates for 14 years, I have decided to give up that position and spend more time on research projects to which I am committed. Hoping to make my suc- cessor’s task easier, I wish to call attention to the high frequency of failures by authors to conform to the format of the Proceedings. Such failures increase unnecessarily the bur- dens of the editors and in most cases could easily be avoided.
I chose 25 items of format and analyzed 40 randomly selected manuscripts for con- formity with these items. The items are list- ed below, arranged in 10 categories. Follow- ing each format item is the number, and in parentheses, the percentage of manuscripts containing that error of format.
I. Address(es) of author(s) 1. Address(es) omitted—2 (5%) 2. Address(es) in wrong place—13 (32.5%) II. Abstract 3. Abstract missing—2 (5%) 4. “‘Abstract”’ a center instead of a side heading— 12 (30%) Center headings (e.g., name of a fam- ily, Acknowledgments, Literature Cit- ed) 5. Side heading used instead of cen- ter heading—4 (10%) 6. Center heading in italics—2 (5%)
Il.
IV.
VI.
VIL.
VU.
IX.
7. Center heading in all capitals— 17
(42.5%) Side headings
8. Side heading not indented—16 (40%)
9. Side heading in wrong format (e.g., in all capitals, all words capital- ized, colon used instead of period and dash)— 13 (32.5%)
. Synonymies
10. Punctuation incorrect—3 (7.5%) 11. Names ofjournals included—2 (5%) Citations of references in text 12. Punctuation incorrect, e.g., “Smith, 1960, p. 10” instead of “Smith 1960:10’—6 (15%) References in text to figures i>" -Figure’” instead of “Fig.” —) (12.5%) 14. “fig.” instead of “‘Fig.” —5 (12.5%) Citations of dates in Materials sections 15. Failure to use day-month-year and 3-letter abbreviation without pe- riod for month, e.g., 15 Jun 1960— 7 (17.5%) Literature Cited 16. Literature Cited single spaced—3 (7.5%) . Names of journals abbreviated — LV 7.5%) . Names of journals in italics—3 (7.5%) . Words in titles of articles capital- ized— 3 (7.5%)
1068
20. Punctuation of authors’ names in- correct— 14 (35%)
21. Pagination lacking or incom- plete—2 (5%)
22. Indentation incorrect—4 (10%)
X. Legends for illustrations
23. ‘Figure’ used instead of “‘Fig.”’— 10 (25%)
24. Indentation incorrect— 19 (47.5%)
25. Punctuation incorrect— 17 (42.5%)
A total of 194 of the 25 kinds of format errors was found in the 40 manuscripts. The number of errors/manuscript ranged from 0-15, with a mean of 4.85 and a mode of 4. Thus the average manuscript contained errors in about ' of the format items. Only Y,) of the manuscripts were free of format errors.
PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Of the 25 format items, directions are giv- en for 8 in “Information for Contributors”’ on the inside back cover of each issue of the Proceedings. The other 17 should be ap- parent to any author who examines a recent issue. Proceedings editors are uncompen- sated volunteers. Please be kind to them by doing your best to follow Proceedings for- mat; it does not require great effort or sac- rifice to do so, and it will speed up the ed- itorial processing of your manuscript.
Department of Invertebrate Zoology, Na- tional Museum of Natural History NHB- 163, Smithsonian Institution, Washington, D.C. 20560.
VOLUME 102, NUMBER 4 1069
INTERNATIONAL COMMISSION ON ZOOLOGICAL NOMENCLATURE
Call for nominations for New Members of the International Commission on Zoological Nomenclature
The following members of the Commission reach the end of their terms of service at the close of the XXIV General Assembly of the International Union of Biological Sciences to be held in Amsterdam, in July 1991: Dr. H. G. Cogger (Australia, Herpetology); Prof Dr O. Kraus (Fed. Rep. Germany, Arachnology); Dr M. Mrocz- kowski (Poland, Coleoptera); Dr W. D. L. Ride (Australia, Mammalia). A further vacancy arises from the resignation of Dr G. C. Gruchy (Canada, Ichthyology).
The addresses and specialist fields of the present members of the Commission may be found in the Bulletin of Zoological Nomenclature, 46(1) (March 1989). Under Article 3b of the Commission’s Constitution a member whose term of service has terminated is not eligible for immediate re-election unless the Council of the Com- mission has decided to the contrary.
The Commission now invites nominations, by any person or institution, of can- didates for membership. Article 2b of the Constitution prescribes that:
“The members of the Commission shall be eminent scientists, irrespective of nationality, with a distinguished record in any branch of zoology, who are known to have an interest in zoological nomenclature.”
(It should be noted that “‘zoology”’ here includes the applied biological sciences (medicine, agriculture, etc.) which use zoological names).
Nominations made since September 1987 will be reconsidered automatically and need not be repeated. Additional nominations, giving the date of birth, nationality and qualifications (by the criteria mentioned above) of each candidate should be sent by 15 June 1990 to: The Executive Secretary, International Commission on Zoological Nomenclature, % British Museum (Natural History), Cromwell Road, London SW7 5BD, U.K.
1070 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON
INTERNATIONAL COMMISSION ON ZOOLOGICAL NOMENCLATURE
Applications Published in the Bulletin of Zoological Nomenclature
Tze iis igs applications were published on 23 June 1989 in Vol. 46, Part 2 of the Bulletin of Zoological Nomenclature. Comment or advice on these applications is invited for publication in the Bulletin and should be sent to the Executive Secretary, I.C.Z.N., % British Museum (Natural History), Cromwell Road, London SW7 5BD, U.K.
Case No.
2657 Marssonopora Lang, 1914 (Bryozoa, Cheilostomata): proposed designation of Membranipora densispina Levinsen, 1925 as the type species.
2403 Valanginites Sayn in Kilian, 1910 (Cephalopoda, Ammonoidea): confirma- tion of the author of the genus, and of Ammonites nucleus Roemer, 1841 as its type species.
2642 POLYGYRIDAE Pilsbry, 1894 (Mollusca, Gastropoda): proposed precedence Over MESODONTIDAE Tryon, 1866.
2666 Lucicutia Giesbrecht in Giesbrecht & Schmeil, 1898: proposed conservation, and Pseudaugaptilus longiremis Sars, 1907: proposed conservation of the specific name (both Crustacea, Copepoda).
2624 Ranguna Bott, 1966 and Larnaudia Bott, 1966 (Crustacea, Decapoda): pro- posed fixation of Thelphusa longipes A. Milne Edwards, 1869 and Thelphusa larnaudii A. Milne Edwards, 1869 as the respective type species.
2542/2 Trapezia Latreille, 1828 (Crustacea, Decapoda): proposed conservation.
2656 Chira Simon, 1902 (Arachnida, Araneae): proposed conservation.
2647 Heliophanus kochi Simon, 1868 (Arachnida, Araneae): proposed conserva- tion of the specific name.
2648 Attus penicillatus Simon, 1875 (currently Sitticus penicillatus; Arachnida, Araneae): proposed conservation of the specific name.
2649 Thyene Simon, 1885 (Arachnida, Araneae): proposed conservation.
2677 Saissetia Déplanche, 1859 (Insecta, Homoptera): proposed designation of Lecanium coffeae Walker, 1852 as the type species.
2695 Fonscolombia Lichtenstein, 1877 (Insecta, Homoptera): proposed designa- tion of Fonscolombia graminis Lichtenstein, 1877 as the type species.
2665 Rosema Walker, 1855 (Insecta, Lepidoptera): proposed conservation.
2658 Protocalliphora Hough, 1899 (Insecta, Diptera) and its type species Musca azurea Fallén, 1817: proposed conservation of usage by designation of a replacement lectotype.
2659 Osteoglossum Cuvier, 1829 (Osteichthyes, Osteoglossiformes): proposed fix- ation of O. bicirrhosum (Cuvier, 1829 as the name of the type species.
VOLUME 102, NUMBER 4 1071
INTERNATIONAL COMMISSION ON ZOOLOGICAL NOMENCLATURE
Opinions published in the Bulletin of Zoological Nomenclature Opinion No.
1536 Sorites Ehrenberg, [1839] (Foraminiferida): Nautilus orbiculus Forsskal, 1775 designated as the type species.
1537 Discocyclina Gimbel, 1870 (Foraminiferida): Orbitolites prattii Michelin, 1847 designated as the type species.
1538 Disculiceps Joyeux & Baer, 1935 (Cestoidea): conserved.
1539 Conus floridanus Gabb, 1869 (Mollusca, Gastropoda): not to be given pre- cedence over Conus anabathrum Crosse, 1865.
1540 Avicula gryphaeoides J. de C. Sowerby, 1836 (Mollusca, Bivalvia): specific name conserved.
1541 Loxoconchella Triebel, 1954 (Crustacea, Ostracoda): Loxoconcha honolu- liensis Brady, 1880 confirmed as the type species.
1542 Chelifer Geoffroy, 1762 (Arachnida, Pseudoscorpionida): conserved.
1543 Dytiscus cinereus Linnaeus, 1758 (currently Graphoderus cinereus; Insecta, Coleoptera): neotype replaced.
1544 ETHMIIDAE Busck, 1909 (Insecta, Lepidoptera): given precedence over AZINI- DAE Walsingham, 1906.
1545 Glabellula Bezzi, 1902 (Insecta, Diptera): Platygaster arcticus Zetterstedt, 1838 designated as the type species.
1546 Chelonus Panzer, 1806 (Insecta, Hymenoptera) and Anomala Samouelle, 1819 (Insecta, Coleoptera): names conserved.
1547 Silurus felis Linnaeus, 1766 (currently Ariopsis felis; Osteichthyes, Siluri- formes): neotype designated.
1548 Sarotherodon melanotheron Rippell, 1852 (Osteichthyes, Perciformes): spe- cific name conserved.
PROC. BIOL. SOC. WASH. 102(4), 1989, p. 1072
BIOLOGICAL SOCIETY OF WASHINGTON
116th Annual Meeting, 25 May 1989
The meeting was called to order by Kris- tian Fauchald, President, at 12:00 noon in the Waldo Schmitt Room, National Mu- seum of Natural History.
Don Wilson, Treasurer, reported that in- come from dues in 1988 was about equal to that of 1987, and that income from sub- scriptions, sales of past issues of the Pro- ceedings, and page charges was substantially higher than in 1987, the latter showing an increase of about $10,000. Total income for 1988 was $100,185 and total expenditures were $87,224. The proposed budget for 1989 balances an estimated income of $94,000 with equal estimated expenditures. The Treasurer's Report was unanimously ac- cepted.
Brian Robbins, Editor, then reported on Volume 101 of the Proceedings. Ninety-nine papers, totaling 977 pages, were published in 1988. Brian noted that unpaid papers cur- rently make up about 300 pages of the Pro- ceedings per year and are subject to a pub- lication delay of approximately one year. Papers with full funding generally have less than a year (about three issues) delay. All manuscripts with full funds submitted in 1988 and some from 1989 will be published
in 1989. Brian announced several changes in Associate Editors during the past year. For Vertebrate Zoology, G. David Johnson replaced Richard P. Vari; for Invertebrate Zoology, Frank D. Ferrari and Raymond B. Manning replaced Thomas E. Bowman; for Entomology, Wayne N. Mathis replaced Robert D. Gordon. Brian noted that sub- stantial progress has been made on the One- Hundred Year Index. Once family coding for each paper has been completed, Phyllis Spangler will produce camera-ready copy for publication as a special issue of the Pro- ceedings, to appear sometime in 1990. The Editor’s Report was accepted without com- ment.
Kristian announced that we will need a new Custodian of Publications to replace Dave Pawson who will be stepping down at the end of May (Austin B. Williams has accepted those duties).
A motion was made and seconded that the meeting be adjourned; Kristian ad- journed the meeting at 12:25 p.m.
Respectfully submitted, G. David Johnson Secretary
INFORMATION FOR CONTRIBUTORS
Content.—The Proceedings of the Biological Society of Washington contains papers bearing on systematics in the biological sciences (botany, zoology, and paleontology), and notices of business transacted at meetings of the Society. Except at the direction of the Council, only manuscripts by Society members will be accepted. Papers are published in English (except for Latin diagnoses/descriptions of plant taxa), with a summary in an alternate language when appropriate.
Submission of manuscripts. —Submit manuscripts to the Editor, Proceedings of the Biological Society of Washington, National Museum of Natural History NHB-108, Smithsonian Insti- tution, Washington, D.C. 20560.
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Presentation. — Manuscripts should be typed double-spaced throughout (including tables, leg- ends, and footnotes) on one side of 8’ x 11 inch sheets, with at least one inch of margin all around. Submit three copies complete with tables, figure captions, and figures (plus originals of the illustrations), and retain an author’s copy. Pages must be numbered consecutively. Underline singly scientific names of genera and lower categories; leave other indications to the editor.
The sequence of material should be: Title, Author(s), Abstract, Text, Acknowledgments, Literature Cited, Author’s(s’) Address(es), Appendix, Figure Legends, Figures (each numbered and identified), Tables (double-spaced throughout, each table numbered with an Arabic numeral and with heading provided).
Clarity of presentation, and requirements of taxonomic and nomenclatural procedures ne- cessitate reasonable consistency in the organization of papers. Telegraphic style is required for descriptions and diagnoses. Literature citations in the text should be in abbreviated style (author, date, page), except in botanical synonymies, with unabbreviated citations of journals and books in the Literature Cited sections. Direct quotations in the text must be accompanied by author, date, and pagination. The establishment of new taxa must conform with the requirements of the appropriate international codes of nomenclature. When appropriate, accounts of new taxa must cite a type specimen deposited in an institutional collection.
Examples of journal and book citations:
Eigenmann, C.H. 1915. The Cheirodontidae, a subfamily of minute characid fishes of South
America.— Memoirs of the Carnegie Museum 7(1):1-99.
Ridgely, R. S. 1976. A guide to the birds of Panama. Princeton, New Jersey, Princeton
University Press, 354 pp.
Olson, S. L. 1973. The fossil record of birds. Pp. 79-238 in D. Farner, J. King, and K. Parkes, eds., Avian biology, volume 8. Academic Press, New York.
Figures and tables with their legends and headings should be self-explanatory, not requiring reference to the text. Indicate figure and table placement in pencil in the margin of the manu- script. Plan illustrations in proportions that will efficiently use space on the type bed of the Proceedings. Original illustrations should not exceed 15 x 24 inches. Figures requiring solid black backgrounds should be indicated as such when the manuscript is submitted, but should not be masked.
CONTENTS
Marianactis bythios, a new genus and species of actinostolid sea anemone (Coelenterata: Ac-
tiniaria) from the Mariana vents Daphne G. Fautin and Robert R. Hessler A revision of the genus Aspidosiphon (Sipuncula: Aspidosiphonidae)
Edward B. Cutler and Norma J. Cutler
Axiothella crozetensis, anew species of maldanid polychaete from Crozet Islands (Indian Ocean)
Patrick Gillet Paraprocerastea crocantinae, a new genus and species (Polychaeta: Syllidae: Autolytinae) from the Spanish Mediterranean Guillermo San Martin and Carmen Aldés
Four new west Atlantic species of Tubificoides (Oligochaeta, Tubificidae) Christer Erséus Three new species of Tubificidae (Oligochaeta) from an oil seepage area on the continental slope of the northern Gulf of Mexico Christer Erséus and Michael R. Milligan Eulimnadia ovilunata and E. ovisimilis, new species of clam shrimps (Crustacea, Branchiopoda, Spinicaudata) from South America Joel W. Martin and Denton Belk Dantya ferox, a new species of mydocopid ostracode from Niue, central South Pacific (Crus- tacea: Ostracoda: Sarsiellidae) Louis S. Kornicker and Thomas M. Iliffe Lamelliform structures on the proboscis of Peniculus and Metapeniculus (Copepoda: Pennel- lidae) Raul Casto Romero and Hernan Baeza K. Acontiophorus excavatus, a new species (Copepoda: Siphonostomatoida) associated with the soft coral Dendronephthya (Alcyonacea) in the Indo-Pacific Arthur G. Humes An unusual species of the Balanus amphitrite Darwin complex (Cirripedia, Balanidae) from the ancestral Colorado River delta in western Arizona and southeastern California Victor A. Zullo and Anna V. Buising First ingolfiellids from the southwest Pacific (Crustacea: Amphipoda) with a discussion of their systematics James K. Lowry and Gary C. B. Poore Allocrangonyctidae and Pseudocrangonyctidae, two new families of holarctic subterranean amphipod crustaceans (Gammaridea), with comments on their phylogenetic and zoogeo- graphic relationships John R. Holsinger New genera in the thalassinidean families Calocarididae and Axiidae (Crustacea: Decapoda) Brian Kensley Three new species of Colombian lace bugs of the genera Lepodictya and Leptopharsa (Het- eroptera: Tingidae) Richard C. Froeschner Krohnittellidae and Bathybelidae, new families in the Phylum Chaetognatha; the rejection of the family Tokiokaispadellidae and the genera Tokiokaispadella, Zahonya, and Aberro- spadella Robert Bieri Hansenothuria benti, new genus, new species (Echinodermata: Holothuroidea) from the tropical western Atlantic: a bathyal, epibenthic holothurian with swimming abilities John E. Miller and David L. Pawson Etheostoma (Nothonotus) wapiti (Osteichthyes: Percidae), a new darter from the southern bend of the Tennessee River system in Alabama and Tennessee David A. Etnier and James D. Williams Ellerkeldia, a junior synonym of Hypoplectrodes, with redescription of the type species of the genera (Pisces: Serranidae: Anthiinae) William D. Anderson, Jr. and Phillip C. Heemstra Acanthemblemaria paula, a new diminutive chaenopsid (Pisces: Blennioidei) from Belize, with comments on life history G. David Johnson and Edward B. Brothers A new species of Eupsophus (Anura: Leptodactylidae) from Contulmo, Nahuelbuta Range, southern Chile Juan Carlos Ortiz, Héctor Ibarra-Vidal, and J. Ramén Formas A new species of colubrid snake of the genus Coniophanes from the highlands of Chiapas, Mexico Jonathan A. Campbell The karyotype of Exiliboa placata Bogert (Tropidopheidae), and comparisons with the family Boidae (Reptilia: Serpentes) Laurence M. Hardy Rhodogorgon, an anamolous new red algal genus from the Caribbean Sea James N. Norris and Katina E. Bucher The high frequency of errors in format in manuscripts submitted to Proceedings of the Biological Society of Washington Thomas E. Bowman International Commission on Zoological Nomenclature Biological Society of Washington: 116th Annual Meeting Table of Contents, Volume 102 L< Index to new taxa, Volume 102 Errata—corrected map for Fig. 2 (p. 657), Hobbs & Robison 102(3):651-697
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