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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 


542 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 4. The coracoid of Megapodius in dorsal (A, B) and ventral (C, D) aspects. A, C, M. freycinet freycinet, 
male, Halmahera, Northern Moluccas, USNM 557015; B, D, M. molistructor, Lifuka, Tonga, BPBM 165682. 


Scale bar = 10 mm. 


Remarks. —This specimen is referred to 
Megapodius molistructor on the basis of its 
being much larger than any other species of 
Megapodius (Table 4), which is the only ge- 
nus of Megapodiidae that occurs in Ocea- 
nia. Intergeneric comparisons were not 
made. BPBM 165682 differs further from 
the coracoids of M. pritchardi, M. freycinet, 
and M. wallacei in having a less concave 
facies articularis humeralis. Although ad- 
ditional material is needed to determine with 
certainty whether the coracoid from Lifuka 
is conspecific with that of MM. molistructor, 
this specimen does demonstrate that an ex- 
tremely large species of megapode, approx- 
imately the size of M. molistructor, once 
lived on Lifuka. 

The type series of Megapodius molistruc- 
tor, an extinct species known otherwise only 


from New Caledonia (Balouet & Olson, 
1989), does not include a coracoid. Never- 
theless, BPBM 165682 is much larger than 
in M. freycinet and larger than would be 
expected for the coracoid of M. alimentum. 
In coracoidal measurements (Table 4), /. 
cf. molistructor from Lifuka is from 1.30+ 
to 1.45 times larger (x = 1.35+, n = 5) than 
the means for M. freycinet freycinet. This 
corresponds with ratios of the measure- 
ments of the scapula and ulna from the type 
series of M. molistructor, which are from 
1.25 to 1.40 times larger (X = 1.33, 
n = 7) than the means for M. freycinet frey- 
cinet (Balouet & Olson, 1989:Table 3). No 
leg elements of M. cf. molistructor are avail- 
able from Lifuka. The tarsometatarsus in 
M. molistructor from New Caledonia is 
much more massive than in M. alimentum 


VOLUME 102, NUMBER 3 543 

Table 4.— Measurements (in mm) of the coracoid in Megapodius, giving mean, range, and sample size. For 
sample sizes larger than 10, the mean is rounded to the nearest 0.05. CS = cotyla scapularis. FAC = facies 
articularis clavicularis. FAH = facies articularis humeralis. ILA = impressio hgamentum acrocoracoideum. 
F = female. M = male. U = sex unknown. 


Depth between Length of Minimum width 
Depth of FAC ILA & FAH Width of FAH FAH & CS of shaft 

M. pritchardi 4.1 2.4 3.8 7.8 3.0 

Niuafo ou, Tonga (2U) 4.04.2 2.2-2.5 3.64.0 7.6—7.9 2.9-3.1 
2 2 2 2 2 

M. cf. molistructor 7.9+ 4.2 6.3+ 13.6 5.7 
BPBM 165682 1 1 1 1 
Lifuka, Tonga (U) 

M. freycinet freycinet 5.95 2.90 4.85 10.20 4.20 
Halmahera, Moluccas 5.4-6.4 2.6—3.4 4.6—5.2 9.5-11.2 3.64.6 
(8M, 8F) 16 16 16 16 

M. freycinet pusillus 6.4 2.9 4.8 10.3 4.5 
Philippines (M) 1 I I 

M. freycinet gilberti BH 2.6 4.3 9.2 3.8 
Celebes (1M, 1F) 5.6-6.2 2.4—2.7 1 9.1-9.2 3.8-3.9 

2 2 2 

M. freycinet abbotti 6.2 3.0 4.8 10.1 4.2 

Nicobar Islands (2M) 6.0-6.3 2.8-3.1 1 l 4.14.4 
2 2 

M. wallacei 5.6 ae | 4.5 9.6 3 

Halmahera, Moluccas (M) 1 1 1 1 


(Fig. 2). Measurements of the tibiotarsus and 
tarsometatarsus of M. alimentum from L1- 
fuka are, respectively, from 0.98 to 1.21 
times larger (X = 1.10, n = 7) and 1.05 to 
1.39 times larger (X = 1.16, n = 6) than the 
means for M. freycinet freycinet, while mea- 
surements of the femur and tarsometatarsus 
of M. molistructor from New Caledonia are, 
respectively, from 1.32 to 1.54 times larger 
(x = 1.39, n = 3) and 1.14 to 1.43 times 
larger (x = 1.31, n = 4) than the mean values 
for M. freycinet freycinet. The validity of 
these calculations is not likely to be com- 
promised by a sexual dimorphism in size, 
which is extremely slight or non-existent in 
species of Megapodius (Mayr 1938, Ama- 
don 1942). 


Order Columbiformes 
Family Columbidae 
Genus Ducula 


Two specimens are referred to the genus 
Ducula rather than other genera of pigeons 


from Polynesia or eastern Melanesia (Co- 
lumba, Ptilinopus, Caloenas, Gallicolumba, 
Goura, Didunculus) because of these char- 
acters: coracoid—medio-ventral side of hu- 
meral end of shaft rounded, sulcus musculo 
supracoracoidei smooth and shallow, facies 
articularis sternalis medio-laterally expand- 
ed but dorso-ventrally compressed, impres- 
sio musculo sternocoracoidei deepest in 
medio-sternal corner; tibiotarsus—size and 
placement of prominent muscle scar on me- 
dio-distal surface of shaft, degree of con- 
cavity on the distal portion of shaft. 


Ducula, undescribed species 
Fig. 5 


Material. —Tibiotarsus lacking both ends, 
BPBM 165685, Pit ONOW, Layer IV (CU- 
III), Tongoleleka archeological site (To-L1i), 
Lifuka, Ha‘apai Group, Tonga. Tom Dye 
and field party Aug 1984. 

Remarks. — This tibiotarsus is larger than 
in any other species of Ducula (Table 5). 


544 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


A B C D 


Fig. 5: 


E F G H 


The tibiotarsus of Ducula in cranial (A—D) and caudal (E-H) aspects. A, E, Undescribed species, 


Lifuka, Tonga, BPBM 165685; B, F, D. galeata, Hanatekua Shelter No. 2 Archeological Site, Hiva Oa, Marquesas, 
BPBM 166055; C, G, D. aurorae, male, captive (original stock presumably from Makatea Island, Tuamotus), 
USNM 344776; D, H, D. pacifica, male, Rarotonga, Cook Islands, USNM 559586. Scale bar = 10 mm. 


The proximo-ventral and disto-lateral por- 
tions of the shaft have smoother, more 
rounded surfaces than other species. This 
tibiotarsus is too fragmentary to be named, 
yet it represents one of the largest of all 
columbids, being exceeded in size only by 
the crowned pigeons of New Guinea (Goura 
Spp.). 


Ducula cf. david Balouet & Olson 
Fig. 6 


Referred material. —Nearly complete 
coracoid, lacking processus acrocoracoid- 
eus and part of facies articularis clavicularis, 
BPBM 165692, Pit ONOW, Layer IV (CU- 
III), Tongoleleka archeological site (To-L1), 


Lifuka, Haapai Group, Tonga. Tom Dye 
and field party Aug 1984. 

Remarks.—Ducula david, recently de- 
scribed from Wallis Island, was character- 
ized mainly by being larger than any extant 
congeners (Balouet & Olson, 1987). Among 
living species of Ducula, the largest of which 
occurs on oceanic islands, only D. galeata 
of eastern Polynesia and D. goliath of New 
Caledonia approach the size of D. david, 
although even these species are slightly 
smaller and less robust (Table 6). 

Ducula david is exceeded in size only by 
the huge extinct species noted above. Al- 
though comparable elements are not avail- 
able for the undescribed species and D. da- 
vid, comparisons of measurements of these 


VOLUME 102, NUMBER 3 


Table 5.—Measurements (in mm) of the tibiotarsus in Ducula and Goura, giving mean, range, and sample 


size. F = female. M = male. U = unknown. 


Length from distal end of Length of Least width Least depth 
fibular crest to distal knob fibular crest of shaft of shaft 
LLL Ss 
Ducula, undescribed sp. 57.4 23.3 5.5 4.8 
Lifuka, Tonga (U) l 1 1 1 
D. galeata 39.7+ _ 3.9 323 
Nuku Hiva, Marquesas (M) (est. 41-42) 1 1 
| 
D. galeata 40.5 — 4.3 3.3 
Henderson Island (U) (composite) 1 1 
BPBM 160464, 160267 
D. galeata 41.5 — 4.2 3.6 
Hiva Oa, Marquesas (U) 1 1 1 
BPBM 166055 
D. goliath wife 12.7 4.3 3.8 
New Caledonia (1M, 2F) 26.2—28.1 11.7-13.6 4.1-4.7 3.6—4.1 
3 3 3 3 
D. aurorae 26.6 10.9 3.9 3.0 
Captive (M) l 1 | 1 
D. pacifica 30.9 11.0 3.6 2.9 
Niuafo ou, Rarotonga 30.2-31.8 10.9-11.2 3.3-3.7 2.8-3.0 
(1M, 1F, 1U) 3 3 3 3) 
D. oceanica 30.0 ED 3.0 2.5 
Palau, Ponape (2U) 28.5-31.4 11.0-11.4 3.0-3.1 2.42.6 
D ) 2 D 
D. aenea 28.4 14.2 4.1 308) 
Philippines (F) ] 1 1 1 
D. perspicillata 28.9 13.6 3.9 Sr2 
Halmahera, Moluccas 28.2—29.6 13.2-14.1 3.9 3.0-3.4 
(1M, 1F) D 2 2 2 
D. bicolor 38.5 13.0 37 3.0 
Halmahera, Moluccas (M) 1 1 1 1 
D. luctuosa 28.2 13.9 3.5 Sul 
Celebes (F) 1 1 l 1 
D. spilorrhoa D522 14.2 Sed 2.9 
Australia (U) 1 1 1 l 
D. badia WB) 13.1 3.4 2.8 
Thailand (M) l 1 1 1 
D. pinon 29.4 14.0 4.0 Sul 
Captive (M) 1 1 l l 
D. radiata 19.4 10.7 3.1 2.4 
Celebes (M) 1 1 1 1 
Goura victoria 68.0 24.7 6.7 5.9 


Captive (M) 


1 


1 


species with those of D. galeata and D. go-_ ticular, the tibiotarsus of the undescribed 
liath indicate that the undescribed species species is 1.40 times longer than that of D. 
is larger than D. david (Tables 5 and 6herein; galeata, whereas the holotypical tarsometa- 
Balouet & Olson, 1987: Table 1). In par- tarsus of D. david from Wallis Island is only 


546 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


VOLUME 102, NUMBER 3 


1.01 times longer than that of D. galeata 
and 1.12 times longer than that of D. goli- 
ath. These ratios correspond well with those 
of the coracoid in D. cf. david from Lifuka, 
which is 1.07 times longer than that in D. 
galeata and 1.12 times longer than that in 
D. goliath. The amount of sexual dimor- 
phism in size is poorly understood in ocean- 
ic species of Ducula owing to the worldwide 
scarcity of skeletons. In measurements of 
D. goliath, the male is consistently larger 
than the female (Tables 5 and 6), although 
these are captive individuals that may not 
be reliable indicators of the size of wild birds. 

No skeletons were available for D. /atrans 
of Fiji. Based upon measurements of skins 
(duPont 1976:83-85), D. latrans is approx- 
imately the same size as D. aurorae or D. 
pacifica, and thus would be much smaller 
than D. david. 


Ducula pacifica (Gmelin) 


Referred material.—Ulna lacking both 
ends (BPBM 165676), Pit 126NOW, Layer 
III (CU-IIb), Tongoleleka archeological site 
(To-Li), Lifuka, Haapai Group, Tonga. 
Tom Dye and field party Aug 1984. 

Remarks. — The curvature of the shaft and 
the prominent papillae remigiales caudales 
refer this ulna to the Columbidae. The spec- 
imen agrees in size and other features with 
the ulna of Ducula pacifica. Each of the oth- 
er three species of columbids reported here 
is much larger than D. pacifica, which is the 
only species of columbid (other than Gal- 
licolumba stairii and species of Ptilinopus, 
which are very small) surviving on Lifuka 
or anywhere else in Tonga. 


Genus Caloenas 


Among the bird bones from the Tongo- 
leleka Site is another coracoid of a large 


— 


547 


columbid. This specimen, slightly smaller 
than in D. cf. david (Table 6), is referred to 
the genus Caloenas rather than to Ducula 
or other pertinent genera of columbids be- 
cause of these characters: greater pneuma- 
ticity in humeral end of sulcus musculo su- 
pracoracoidei; facies articularis humeralis 
protrudes more ventrad from surface of 
shaft; sharp medio-ventral edge of humeral 
end of shaft; in medial aspect, portion of 
shaft between cotyla scapularis and facies 
articularis clavicularis faces more perpen- 
dicularly (less diagonally); facies articularis 
clavicularis deeper. 


Caloenas cf. canacorum Balouet & Olson 
Fige o/ 


Referred material. —Humeral end of cor- 
acoid, including facies articularis humeralis 
and cotyla scapularis, BPBM 165678, Pit 
ON20E, Layer II (CU-II), Tongoleleka ar- 
cheological site (To-Li), Lifuka, Haapai 
Group, Tonga. Tom Dye and field party 
Aug 1984. 

Remarks.—Caloenas canacorum is an 
extinct species recently described from late 
Holocene fossils (sternum, coracoids, scap- 
ula, and humerus) from New Caledonia 
(Balouet & Olson, 1989). Although direct 
comparison of the holotype coracoid of 
C. canacorum with BPBM 165678 was not 
possible except in photographs (Fig. 7), 
BPBM 165678 is referred to C. cf. cana- 
corum because of similarity in qualitative 
generic characters and in size, being signif- 
icantly larger than in C. nicobarica, the only 
living species in this distinctive genus (Ta- 
ble 6). 


Discussion 


Although the detailed implications of 
these findings will be reported elsewhere, a 


Fig. 6. The coracoid of Ducula in dorsal (A—D) and ventral (E—-H) aspects. A, E, D. cf. david, Lifuka, Tonga, 
BPBM 165692; B, F, D. galeata, Hanatekua Shelter No. 2 archeological site, Hiva Oa, Marquesas, BPBM 
166056; C, G, D. aurorae, male, captive (original stock presumably from Makatea Island, Tuamotus), USNM 
344776; D, H, D. pacifica, male, Rarotonga, Cook Islands, USNM 559586. Scale bars = 10 mm. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


548 


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550 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 7. The coracoid of Caloenas in dorsal (A-C) and ventral (D-F) aspects. A, D, C. nicobarica, female, 
Halmahera, Northern Moluccas, USNM 557089; B, E, Caloenas cf. canacorum, Lifuka, Tonga, BPBM 165678; 
C, F, C. canacorum, paratype, New Caledonia, MNHN 300. Scale bar = 10 mm. 


few comments are pertinent here. The un- 
described species of Ducula is of unknown 
interspecific relationships. The other extinct 
birds from Lifuka are related to species 
found west or north of Tonga. It seems like- 
ly that all of these species, or closely related 
ones, once occurred through much of the 
region of New Caledonia, Vanuatu, Fiji, 
Tonga, and Samoa. Megapodius molistruc- 
tor and Caloenas canacorum are known only 
from New Caledonia and tentatively from 
Lifuka. Megapodius alimentum and the un- 
described species of Ducula are known only 
from Lifuka, although there is no reason to 
believe that they were confined to that is- 
land. Aside from Lifuka, D. david is known 
from Wallis (Uvea) Island, which is north 
of Lifuka, between Samoa and Fiji. 

The late Holocene extinction of two 
species of megapodes and three species of 
pigeons on Lifuka shows that there has been 


a significant loss of birds in western Poly- 
nesia since the arrival of man. Numerous 
extinctions have been documented by fossil 
records from more remote parts of Poly- 
nesia, such as Hawaii (Olson & James 1982a, 
b), Marquesas (Steadman, in press), Hen- 
derson Island (Steadman & Olson 1985), 
Cook Islands (Steadman 1985, in press), and 
New Zealand (Cassels 1984). More exca- 
vation is needed in Tonga and Samoa. Based 
upon the limited record available, the de- 
gree of avian extinction in western Poly- 
nesia may have been just as severe as that 
from elsewhere in Polynesia. 

The fossils from Lifuka indicate that two 
species of Megapodius once occurred there. 
Megapodius pritchardi, restricted to the iso- 
lated Tongan island of Niuafo ou, is the only 
species of megapode that survives anywhere 
in Polynesia, although we presently do not 
know to what extent the natural range of 


VOLUME 102, NUMBER 3 


megapodes has been reduced by human im- 
pact. The widespread M. freycinet reaches 
the eastern limit of its range in Vanuatu 
(New Hebrides). That the absence of mega- 
podes in the Fijian region may be an artifact 
of human disturbance was noted by Olson 
(1980) and confirmed several years ago by 
our examination of bones of Megapodius 
(species undetermined) from an archeolog- 
ical site on Lakeba, Lau Group, Fiji (re- 
ported in Gibbons & Clunie 1986). Else- 
where in the western Polynesian and Me- 
lanesian region, extinct megapodes (species 
undetermined) have been reported from as 
yet unconfirmed historical accounts in the 
Kermadec Islands (Lister 1911), archeolog- 
ical sites on Tikopia (Kirch & Yen 1982: 
282; Green 1976), an egg collected in 1847 
from Samoa (island undetermined; Gray 
1862), and an egg collected before 1862 from 
an undetermined island in the Haapai 
Group of Tonga (Gray 1862, 1864). Oates 
(1901) referred the last two specimens to M. 
pritchardi, a determination that should be 
reconfirmed. 

A better understanding of the systematics 
and natural distribution of megapodes in 
Oceania depends upon the reexamination 
of historic specimens and documents, and 
more fully upon the discovery and study of 
bones from prehistoric sites on many ad- 
ditional islands. It now seems likely that one 
to three species of megapode occurred on 
most or all islands of eastern Melanesia and 
western Polynesia before the arrival of hu- 
mans. Four species of megapodes still exist, 
for example, on the Papuan island of Misool 
(Ripley 1960). 

Columbids also have suffered much ex- 
tinction in Oceania. The hunting of pigeons 
by prehistoric Tongans was extensive and 
highly organized (McKern 1929:19-27). 
Ducula pacifica is the largest pigeon known 
historically from anywhere in Tonga, where 
it is found essentially throughout the group. 
Fossils from Tongoleleka represent three 
additional species of columbids, each ex- 
tinct and larger than D. pacifica. Although 


551 


it may seem remarkable that four large 
species of columbids, including three species 
of Ducula, once lived on Lifuka, we really 
do not yet know the natural (=pre-human) 
distribution and diversity of Pacific co- 
lumbids. From Mangaia in the Cook Is- 
lands, for example, late Holocene fossils 
represent five species of columbids where 
none exists today (Steadman 1985, 1989). 


Acknowledgments 


I thank T. Dye for his cooperation and 
generosity in making available the Lifukan 
specimens and associated information. S. L. 
Olson and J. C. Balouet kindly allowed ac- 
cess to specimens and data on extinct birds 
from New Caledonia, including photo- 
graphs of Megapodius molistructor and Ca- 
loenas canacorum. For access to specimens, 
I thank J. P. Angle, S. L. Olson, C. A. Ross, 
and R. L. Zusi (USNM) and A. Allison, C. 
H. Kishinami, and G. Wine (BPBM). D. 
Pahlavan, S. E. Schubel, and M. C. Zarriello 
assisted in other curatorial matters. D. Shi- 
deler, L. Woodward, and A. Ziegler sorted 
and preliminarily identified the bones. The 
photographs are by T. Beblowski and C. 
Supkis. The manuscript was improved by 
comments from T. Dye, N. G. Miller, S. L. 
Olson, and G. K. Pregill. The research was 
supported by National Science Foundation 
Grant BSR-8607535. This paper is contri- 
bution number 538 of the New York State 
Science Service. 


Literature Cited 


Amadon, D. 1942. Birds collected during the Whit- 
ney South Sea Expedition. XLIX. Notes on some 
non-passerine genera, 1.—American Museum 
Novitates 1175:1-11. 

Baker, R. H. 1951. The avifauna of Micronesia, its 
origin, evolution, and distribution. — University 
of Kansas Publications, Museum of Natural 
History 3:1-359. 

Balouet, J. C., & S. L. Olson. 1987. A new extinct 
species of giant pigeon (Columbidae: Ducula) 
from archeological deposits on Wallis (Uvea) 


352 


Island, South Pacific.— Proceedings of the Bio- 
logical Society of Washington 100:769-775. 
,& 1989. Fossil birds from late Qua- 
ternary deposits in New Caledonia. —Smithson- 
ian Contributions to Zoology 469:1-38. 
Baumel, J. J., A. S. King, A. M. Lukas, J. E. Breazile, 
& H. E. Evans (eds.). 1979. Nomina anato- 
mica avium. Academic Press, London, 664 pp. 

Cassels, R. 1984. The role of prehistoric man in the 
faunal extinction of New Zealand and other Pa- 
cific islands. Pp. 741-767 in P. S. Martin and 
R. G. Klein, eds., Quaternary extinctions. Uni- 
versity of Arizona Press, Tucson. 

duPont, J. E. 1976. South Pacific birds.— Delaware 
Museum of Natural History Monograph Series 
3:1-218. 

Gibbons, J. R. H., & F. G. A. U. Clunie. 1986. Sea 
level changes and Pacific prehistory.—Journal 
of Pacific History 21:58-82. 

Gray,G.R. 1862. List of species composing the Fam- 

ily Megapodiidae, with descriptions of new 

species, and some account of the habits of the 
species. — Proceedings of the Zoological Society 

of London for 1861:288-296. 

1864. Ona new species of megapode.— Pro- 
ceedings of the Zoological Society of London for 
1864:41-44. 

Green, R. C. 1976. Lapita sites in the Santa Cruz 
Group. Pp. 245-265 in R. C. Green and M. M. 
Cresswell, eds., Southeast Solomon Islands cul- 
tural history.—Royal Society of New Zealand 
Bulletin 11. 

Karch, P. V., & D.E. Yen... 1982. Tikopia= The: pre- 
history and ecology of a Polynesian outlier. — 
Bernice P. Bishop Museum Bulletin 238:1-396. 

Lister, J.J. 1911. The distribution of the avian genus 
Megapodius in the Pacific Islands.— Proceed- 
ings of the Zoological Society of London for 
1911:749-759. 

Mayr, E. 1938. Birds collected during the Whitney 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


South Sea Expedition. XX XIX. Notes on New 
Guinea birds.—American Museum Novitates 
1006:1-16. 

McKern, W. C. 1929. Archaeology of Tonga.—Ber- 
nice P. Bishop Museum Bulletin 60:1-123. 

Oates, E. W. 1901. Catalogue of the collection of 
birds’ eggs in the British Museum (Natural His- 
tory). British Museum (Natural History), Lon- 
don, 252 pp. 

Olson, S.L. 1980. The significance of the distribution 
of the Megapodiidae.—Emu 80:21-24. 

—, & H. F. James. 1982a. Fossil birds from the 
Hawaiian Islands: Evidence for wholesale ex- 
tinction by man before Western contact.—Sci- 
ence 217:633-635. 

——., & 1982b. Prodromus of the fossil 
avifauna of the Hawaiian Islands.—Smithson- 
ian Contributions to Zoology 365:1-59. 

Ripley, S. D. 1960. Distribution and niche differ- 
entiation in species of megapodes in the Mo- 
luccas and western Papuan area.— Proceedings 
XII International Ornithological Congress 2: 
631-640. 

Steadman, D. W. 1985. Fossil birds from Mangaia, 
southern Cook Islands.— Bulletin of the British 
Ornithologists’ Club 105:58-66. 

1989. Fossil birds and biogeography in Po- 
lynesia.—Acta XIX Congressus Internationalis 
Ornithologici II:1526-1534. 

—, & S. L. Olson. 1985. Bird remains from an 
archaeological site on Henderson Island, South 
Pacific: Man-caused extinctions on an “unin- 
habited”’ island.— Proceedings of the National 
Academy of Sciences USA 81:4448-4451. 


Biological Survey, New York State Mu- 
seum, The State Education Department, Al- 
bany, New York 12230. 


PROC. BIOL. SOC. WASH. 
102(3), 1989, pp. 553-554 


GEOGRAPHIC VARIATION IN THE YELLOW-RUMPED 
TANAGER (AVES: THRAUPINAE) 


Robert W. Storer 


Abstract.—Females and young males of the Yellow-rumped Tanager (the 
icteronotus group of the Flame-rumped Tanager, Ramphocelus flammigerus) 
from Panama to west central Colombia are grayer and less green on the back 
with less black on the crown than are those from Ecuador. Birds from south- 
western Colombia and northeastern Ecuador are intermediate between those 
populations, the latter being more similar to those of the other Ecuadorean 
populations. The name Ramphocelus flammigerus varians Lafresnaye should 
be applied to the northern populations and R. f. icteronotus Bonaparte to those 


from Ecuador. 


Yellow-rumped Tanagers (the icteronotus 
group of the Flame-rumped Tanager, Ram- 
Dhocelus flammigerus) are widely distrib- 
uted from western Panama to southwestern 
Ecuador (American Ornithologists’ Union 
1983). Wetmore et al. (1984:449) comment 
that “preliminary examination reveals that 
females and immatures from Ecuador are 
much greener, less brownish or gray, above 
than are most birds from Colombia and 
Panama. Thus it is possible that the Pana- 
manian birds should take the name varians 
Lafresnaye 1847 (type locality, Buenaven- 
tura Colombia).” 

To determine the correct name for the 
Panamanian birds, it was necessary to de- 
termine first, if there is sufficient geographic 
variation to recognize more than one race, 
and second, if so, whether the birds from 
near the type locality of varians are closer 
to birds from Panama or western Ecuador 
(the restricted type locality of icteronotus, 
Berlepsch, 1912). Should the latter situation 
apply, a new name would be needed for the 
Panamanian birds. 

I examined 143 specimens of females and 
immature males of the Yellow-rumped 
Tanager for analysis. (Adult males do not 
vary in the deep black of the back.) Wing 
length (chord), tail length, and length of bill 


from nostril to tip of females were measured 
with dial calipers, the first two to the nearest 
0.5 mm and the last to the nearest 0.1 mm. 
(The sample of males was too small to treat 
Statistically.) The geographic range was di- 
vided into five areas with gaps between them 
to avoid comparisons between contiguous 
populations and to reduce overlap between 
samples. The selection of the areas was based 
primarily on the distribution of the mate- 
rial. These areas were: Western Panama 
(Bocas del Toro Province and the Canal 
Zone), Eastern Panama (Darien Province), 
Central Colombia (provinces of Cauca, To- 
lima, and Valle), Northern Ecuador (prov- 
inces of Esmeraldas, Imbabura, and Pichin- 
cha), and Southern Ecuador (provinces of 
Canar, El Oro, Guayas, and Loja). 

Color comparisons were made by ex- 
amination of specimens in north light. Bad- 
ly worn or faded specimens, approximately 
one half of those examined, were not used 
in the comparisons. 

The measurement data for females (Table 
1) show a cline in increasing wing and tail 
length from western Panama to southern 
Ecuador. Bill length is greatest in birds from 
eastern Panama and least in those from 
western Panama and central Colombia, but 
mean differences are small. In none of the 


554 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Table 1.—Sample size, mean, and standard deviation of measurements of female Yellow-rumped Tanagers. 


Measurement Western Panama Eastern Panama 


Central Colombia 


Northern Ecuador Southern Ecuador 


Wing length fT WW 0) 9 S29" aale9 23) OS te 1.8 10 80.4 + 2.6 12) 83:4 27189 
Tail length 6 68.2, 4233.2 $68.9: 3.2 21 GOR say 2yi 1Q; 69.7 22.23 12. 138.4537) 
Bill length A AZ EOS 9) 13:4,-©.0:6 22. 12:8 2106 10, 32.203 12, 13,125)0e 


measurements is the difference between the 
largest and smallest mean greater than the 
sum of the standard deviations. Therefore, 
none of the differences approaches a degree 
of difference useful in separating subspecies. 
In wing and tail measurements, the birds 
from central Colombia are closer to those 
from eastern Panama than to those from 
northern Ecuador. The data for bill length 
are equivocal. 

In color, specimens from Panama are de- 
cidedly grayer (less black) on the back than 
those from Ecuador, the crowns are less 
black, the yellow edging on the back feathers 
is paler and contrasts less with the dark col- 
or of the rest of the feather, and the yellow 
of the underparts is paler, especially on the 
abdomen. The brighter yellow edgings and 
darker central parts of the feathers combine 
to produce a greener tone to the back feath- 
ers of the Ecuadorean birds. Five specimens 
from the province of Valle, central Colom- 
bia, (including Buenaventura, the type lo- 
cality of varians) are nearer birds from Pan- 
ama in color than those from Ecuador. In 
Ecuador, there is a noticeable difference in 
color between specimens from the northern 
and those from the central and southern 
parts of the country, the latter being bright- 
er. However, birds from northern Ecuador 
are more similar to those from the south 
than to those from Panama and northern 
Colombia. Thus, birds from Panama 
through central Colombia (Valle) vary little 
in color, the major change occurring from 
Cauca to Ecuador, with a smaller change 
occurring between the northern and central 
parts of Ecuador. 


In conclusion, differences in wing, tail, 
and bill lengths are not sufficient to warrant 
recognition of subspecies. Color differences, 
however, are greater than in many recog- 
nized subspecies. I therefore propose that 
the birds from Panama through the prov- 
ince of Valle, Colombia, be called Ram- 
Dhocelus flammigerus varians Lafresnaye, 
those from Ecuador, Ramphocelus flam- 
migerus icteronotus Bonaparte, and those 
from southwestern Colombia, intergrades. 


Acknowledgments 


I acknowledge the assistance of the cu- 
rators of the bird collections of the Amer- 
ican Museum of Natural History, the Field 
Museum of Natural History, and the Na- 
tional Museum of Natural History, Smith- 
sonian Institution, for the loan of speci- 
mens, and Richard C. Banks and Thomas 
R. Howell for reading the manuscript and 
offering helpful suggestions. 


Literature Cited 


American Ornithologists’ Union. 1983. Check-list of 
North American Birds, 6th Edition, 662 pp. 

Berlepsch, H. von. 1912. Bericht ueber den V. In- 
ternationalen Ornithologen-Kongress Berlin 
1910, p. 1061. 

Wetmore, A., R. F. Pasquier, & S. L. Olson. 1984. 
The Birds of the Republic of Panama.—Smith- 
sonian Miscellaneous Collections 150(4):448- 
449. 


Museum of Zoology and Department of 
Biology, University of Michigan, Ann Ar- 
bor, Michigan 48109. 


PROC. BIOL. SOC. WASH. 
102(3), 1989, pp. 555-558 


TWO OVERLOOKED HOLOTYPES OF THE HAWAIIAN 
FLYCATCHER CHASTEMPIS DESCRIBED BY 
LEONHARD STEJNEGER (AVES: MYIAGRINAE) 


Storrs L. Olson 


Abstract.—The holotypes of two of the three taxa of Elepaio (Chasiempis) 
described by Leonhard Stejneger in 1887 have been overlooked for almost a 
century but were located in the collections of the British Museum (Natural 
History) and restudied. That of Chasiempis ridgwayi Stejneger presents no 
problems as it is representative of the subspecies from the windward parts of 
the island of Hawaii currently known as Chasiempis sandwichensis ridgwayi. 
The holotype of Chasiempis ibidis Stejneger, 1887, however, is identified as 
being from the Oahu population and this name thus takes precedence over 


Chasiempis gayi Wilson, 1891. 


In the early literature on systematics and 
nomenclature of the Hawaiian flycatcher 
known as the Elepaio, Chasiempis sand- 
wichensis (Gmelin), there was considerable 
disagreement concerning the number of taxa 
and their distribution. Some proponents 
held out for a single species (e.g. Sclater 1885, 
Berlepsch & Leverkuhn 1890), while others 
recognized as many as five or six (e.g. 
Steyneger 1887, Wilson 1891), with the is- 
land of origin often being completely ig- 
nored, however. It was eventually deter- 
mined that the Elepaio occurred only on 
three of the Hawaiian islands, with the pop- 
ulation of each island coming to be recog- 
nized under a single name: C. sclateri Ridg- 
way, 1882, on Kauai; C. gayi Wilson, 1891, 
on Oahu; and C. sandwichensis (Gmelin, 
1789) on Hawaii. These taxa are now gen- 
erally regarded as subspecies of C. sand- 
wichensis. Geographic variation within the 
island of Hawaii caused Henshaw (1902) to 
recognize two forms there, with the second 
taking the name C. ridgwayi Stejneger, 1887. 
Pratt (1979, 1980) recognized these and de- 
scribed a third subspecies from Hawaii, C. 
s. bryani. 

Leonhard Stejneger was in the thick of 
the early confusion surrounding the system- 


atics of Chasiempis and proposed no less 
than three new taxa (Stejneger 1887). The 
only actual specimens available to him, 
however, were those taken on Kauai by Val- 
demar Knudsen and forwarded to the 
Smithsonian Institution. Part of the prolif- 
eration of taxa resulted from the two dis- 
tinct plumage types found on each island, 
now generally regarded as adult and “im- 
mature.”’ Thus, Stejneger’s name Chasiem- 
pis dolei, the type of which (USNM 110040) 
is a gray-backed adult bird from Kauai, is 
a pure synonym of C. sclateri Ridgway, 
1882, the cotypes of which (USNM 41955, 
41956) are brown-backed immature birds 
from Kauai (see Deignan 1961:460). 
Stejneger’s other two names, C. ridgwayi 
and C. ibidis, were based on a color plate 
published in Jbis (hence the latter name) by 
Sclater (1885). Although Stejneger attempt- 
ed to forestall criticism by saying that if C. 
ibidis were not distinct from C. sclateri “‘then 
I can only say that the published figure of 
the former is worse than useless” (Stejyneger 
1887:88), his contemporaries nevertheless 
roundly excoriated him. “It is a pity that 
Dr. Stejneger, with so much good material 
before him, should think it necessary to 
manufacture ‘new species’ out of other peo- 


556 


ple’s figures without seeing the specimens” 
(Sclater 1888:143). “On the genus Chasiem- 
pis | would offer only one remark, and that 
is a word of caution to those who would, on 
the evidence of from a couple to half-a-doz- 
en of specimens, or perhaps even on the 
evidence of a badly-coloured plate, attempt 
to break it up into definable ‘species’ ”’ 
(Newton 1892:469). With the benefit of 
hindsight, it is fair to note that the plate in 
question is actually quite accurate, and that 
both of Stejneger’s taxa based on it are now 
seen to be valid. 

Although both figures were stated to be 
based on particular specimens, these have 
been overlooked in the general collections 
of the British Museum (Natural History) for 
nearly a century and have long gone unrec- 
ognized as types (neither is mentioned in 
Warren & Harrison 1971). They still exist, 
however, and I was able to examine and 
compare them with the series of Chasiempis 
in the National Museum of Natural History, 
Smithsonian Institution (USNM), and with 
a selection of specimens from the American 
Museum of Natural History that were in 
plumage comparable to that of C. ibidis (see 
Material Examined). 

One of Stejneger’s names was applied to 
‘the brown and chestnut colored bird from 
Hawaii, Ch. ridgwayi, figured on plate 1, Ibis, 
1885” (Stejneger 1887:87). Sclater (1885: 
18) had mentioned two specimens collected 
by the Challenger Expedition at Hilo, Ha- 
wail, in August, 1875, and stated that “‘the 
figure (Plate I fig. 1) has been taken from 
one of them.” Both specimens are in nearly 
identical plumage but one of them (BMNH 
80.11.18.445, original no. 529) is in much 
worse condition, with most of the feathers 
of the rump missing and many of the rec- 
trices broken off, so that only one remains 
that has a white tip. Because the plate shows 
a bird with a large white rump patch and 
white tips on most of the rectrices, if it were 
drawn from a single specimen, as stated by 
Sclater, this would have to be BMNH 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


80.11.18.444 (original no. 528), which I here 
affirm to be the holotype of Chasiempis 
ridgwayi Stejneger (culmen, 13.1 mm; wing, 
67.7; tail 54.5, tarsi not measurable). No 
nomenclatural problems attach to this iden- 
tification, as the specimen is of known 
provenance and is clearly the bird from 
windward Hawaii currently known as Cha- 
siempis sandwichensis ridgwayi. 

The specimen from which Sclater’s re- 
maining figure was drawn, the type of Cha- 
siempis ibidis, has a much more curious and 
enigmatic history. Sclater (1885:18) re- 
ceived the specimen, labelled “‘Chili,” prior 
to 1862 from the dealer Verreaux and mis- 
took it for some undetermined species of 
Tyrannidae. Thus he once listed it as “Cni- 
polegus __?” (Sclater 1862:203), an error 
that he later corrected to Chasiempis sand- 
wichensis (Sclater 1873) after comparing the 
specimen with material in the Berlin Mu- 
seum (Sclater 1885), where the only speci- 
mens of Chasiempis then were those col- 
lected on Oahu by Deppe in 1837. The same 
specimen was listed by Sharpe (1879) as 
Chasiempis sandvicensis (sic), at which time 
it was the only example of Elepaio in the 
British Museum. It is the only Elepaio from 
the Sclater collection ex Verreaux in the 
British Museum collections or elsewhere and 
is thus certainly identifiable as the holotype 
of Chasiempis ibidis Stejneger (BMNH 
73.8.6.3; culmen broken; wing, 62.9 mm; 
tail, 59.3; tarsus, 25.1). Itisin fresh, unworn 
plumage and agrees perfectly with the figure 
in Sclater’s (1885) plate except that the 
ochraceous color at the posterior margin of 
the throat is somewhat darker than depict- 
ed. 

Circumstantial evidence alone would 
suggest that this specimen came from Oahu, 
as most commerce then, as now, was through 
Honolulu. That it had been labelled “Chili” 
is understandable as Chile was then a dis- 
patch point for cargo going “around the 
Horn” to Europe. In the Senckenberg Mu- 
seum, Frankfurt, are at least five specimens 


VOLUME 102, NUMBER 3 


of Hawaiian birds labelled as received or 
exchanged from Chile in 1842, the notation 
“Erh[halten] von Chili’? having been mis- 
taken by Banko (1979:31, 32, 80) for the 
collector’s name. Of these, three are of the 
Oahu form of Loxops virens; the other two 
belong to species that show no interisland 
variation (Psittirostra psittacea and Ves- 
tiaria coccinea). Itis uncertain who was sup- 
plying specimens from Oahu through Chile 
at that time, but it was very likely the same 
source whence Verreaux obtained the spec- 
imen of Chasiempis sold to Sclater. 

As far as the actual characters of the ho- 
lotype of Chasiempis ibidis are concerned, 
Stejneger (1887:88) was perfectly correct in 
describing it as differing from C. sclateri of 
Kauai, which is “much deeper and richer 
tawny color. . . and this color extends much 
further on breast, flanks, and tibiae than in 
Ch. ibidis.” 

Wilson (1891) considered C. ibidis to have 
come from Oahu. The new form from Oahu 
that he called Chasiempis gayi he described 
as a second species from that island in ad- 
dition to C. ibidis. Why then has the Oahu 
bird come to be known as C. gayi rather 
than C. ibidis? This results entirely from 
Rothschild’s (1893:71) statement that the 
type of C. ibidis ‘agrees best with the young 
Hawaiian bird, so there is no doubt it really 
came from Hawaii.” This is erroneous. 
Rothschild’s conclusion appears to have 
been unduly influenced by some exceptional 
specimens (e.g. AMNH 607136, 607138) 
collected by Palmer on the Kona coast of 
Hawaii, the first mentioned having served 
as the model for the immature of C. sand- 
wichensis in Rothschild’s accompanying 
plate. These birds are much more rufescent, 
especially on the throat and breast, than typ- 
ical immature birds from Hawaii, which are 
dark brownish above, with a grayish crown, 
and white lores and underparts. The light 
tawny ochraceous color of the type of C. 
ibidis is very unlike this and is matched only 
by specimens from Oahu. The exceptionally 


557 


rufescent specimens from Hawaii are still 
much darker, more chestnut, above, espe- 
cially on the rump, than in C. ibidis. Another 
overlooked difference is in the shape of the 
bill, which in birds from Oahu and Kauai 
appears broader and flatter than in birds 
from Hawaii. Although the bill in the type 
of C. ibidis is damaged and lacks the tip of 
the upper part, its shape agrees better with 
birds from Oahu than with those of Hawaii. 

The holotype of Chasiempis ibidis is un- 
questionably representative of the Oahu 
population of Elepaio, as Wilson (1891) 
himself recognized. Because Chasiempis 
ibidis Stejneger, 1887, has priority over 
Chasiempis gayi Wilson, 1891, the Oahu 
Elepaio should now be known as Chasiem- 
pis ibidis or Chasiempis sandwichensis ibi- 
dis. 

Material examined.—Chasiempis s. 
sandwichensis:. AMNH 607118, AMNH 
607125, AMNH 607136, AMNH 607138. 
C. s. ridgwayi: BMNH 80.11.18.444 (ho- 
lotype), BMNH 80.11.18.445, AMNH 
193362, AMNH 193366, AMNH 193368, 
plus about 30 USNM specimens in imma- 
ture plumage. C. ibidis: BMNH 73.8.6.3 
(holotype), AMNH 193354, AMNH 
193355, AMNH 193357, AMNH 168638, 
AMNH 199353, AMNH 607160, USNM 
301122. C. sclateri: AMNH 168639, 
AMNH 193347, AMNH 607188, AMNH 
607189, AMNH 607190, AMNH 607198, 
USNM 41955 and 41956 (cotypes), USNM 
110040 (type of C. dolei), USNM 110037, 
USNM 110038, USNM 116782, USNM 
116783, USNM 493863, USNM 493864, 
USNM 591935, USNM 591936. 


Acknowledgments 


I am exceedingly grateful to Graham 
Cowles, Sub-Department of Ornithology, 
British Museum (Natural History) (BMNH), 
Tring, for lending the specimens that proved 
to be the types of C. ridgwayi and C. ibidis, 
and to Richard A. Sloss and Mary LeCroy, 
American Museum of Natural History 


558 


(AMNH), New York, for supplying addi- 
tional comparative material. I also thank D. 
S. Peters for access to specimens of Hawai- 
ian birds in the Senckenberg Museum, 
Frankfurt. 


Literature Cited 


Banko, W. E. 1979. History of endemic Hawaiian 
birds [sic] specimens in museum collections. — 
Cooperative National Park Resources Study 
Unit, University of Hawaii, Avian History Re- 
port 2:1-80. 

Berlepsch, H. von, & P. Leverkiihn. 1890. Studien 
iiber stidamerikanische Vogel nebst Beschrei- 
bungen neuer Arte.—Ornis 6:1-32. 

Deignan, H. G. 1961. Type specimens of birds in the 
United States National Museum.— United States 
National Museum Bulletin 221:1-718. 

Gmelin, J.F. 1789. Systema Naturae. Volume 1, part 
2. Lipsiae, G. E. Beer, pp. 501-1032. 

Henshaw, H. W. 1902. The Elepaio of Hawaii.— Auk 
19:225-232. 

Newton, A. 1892. Omnithology of the Sandwich Is- 
lands. — Nature 45:465-469. 

Pratt, H. D. 1979. A new subspecies of the Elepaio, 

Chasiempis sandwichensis, from the island of 

Hawatii.— Bulletin of the British Ornithologists’ 

Club 99:105-108. 

1980. Intra-island variation in the ‘Elepaio 
on the Island of Hawai’i.— Condor 82:449-458. 
Ridgway, R. 1882. Description of a new fly-catcher 

and a supposed new petrel from the Sandwich 
Islands. — Proceedings of the United States Na- 
tional Museum 4:337-338. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Rothschild, W. 1893. The avifauna of Laysan and 
the neighbouring islands: With a complete his- 
tory to date of the birds of the Hawaiian pos- 
sessions. Part 2. London, R. H. Porter, pp. 59- 
126. 

Sclater, P.L. 1862. Catalogue ofa collection of Amer- 
ican birds. London, N. Trubner and Co., 368 
pp. 

1873. [Corrections to Sclater’s Catalogue of 

American Birds.]— Proceedings of the Zoolog- 

ical Society of London 1873:554—555. 

1885. On the muscicapine genus Chasiem- 

pis. —Ibis, series 5, 3:17—19, plate 1. 

. 1888. [review] Stejneger on Hawaiian birds. — 

Ibis, series 5, 6:143-144. 

Sharpe, R. B. 1879. Catalogue of the birds in the 
British Museum. London, British Museum, 494 
pp. 

Stejneger, L.H. 1887. Birds of Kauai Island, Hawai- 
ian Archipelago, collected by Mr. Valdemar 
Knudsen, with descriptions of new species.— 
Proceedings of the United States National Mu- 
seum 10:75-102. 

Warren, R. L. M., & C. J. O. Harrison. 1971. Type- 
specimens of birds in the British Museum (Nat- 
ural History). Volume 2, passerines. London, 
British Museum (Natural History), 628 pp. 

Wilson, S. B. 1891. On the muscicapine genus Chasi- 
empis, with a description of a new species.— 
Proceedings of the Zoological Society of London 
1891:164-166. 


Department of Vertebrate Zoology, Na- 
tional Museum of Natural History, Smith- 
sonian Institution, Washington, D.C. 20560. 


PROC. BIOL. SOC. WASH. 
102(3), 1989, pp. 559-567 


A NEW LIZARD OF THE GENUS LEPIDODACTYLUS 
(REPTILIA: GEKKONIDAE) FROM 
BATAN ISLAND, PHILIPPINES 


Hidetoshi Ota and Ronald I. Crombie 


Abstract. — A new species of Lepidodactylus, L. balioburius, is described from 
Batan Island in the northernmost archipelago of the Philippines. It is most 
closely related to the recently described L. yami from Lanyu Island, Taiwan. 
Morphological variation in both species is analyzed. 


The herpetofauna of mountainous north- 
ern Luzon and the island groups stretching 
north toward Taiwan remains poorly known, 
despite intensive field work in other parts 
of the Philippines during the past 20-30 
years. The Batan Island group, the north- 
ernmost archipelago in the Philippines, is 
of considerable zoogeographic interest since 
it is located almost midway between the 
Philippines and Taiwan (220 km north of 
Luzon and 200 km south of Lanyu Island, 
see Fig. 1). A few specimens from Batan 
reported in the literature perished when the 
Bureau of Science collection in Manila was 
destroyed during World War II. During May 
and June 1985, a multidisciplinary team of 
biologists, coordinated by Angel C. Alcala 
(Silliman University) and Charles A. Ross 
(Smithsonian Institution), collected on sev- 
eral islands in the Batan Group. Their col- 
lections contained a small series of a dis- 
tinctive new species of Lepidodactylus, a 
genus unreported from the extreme north- 
ern Philippines. The new species is super- 
ficially similar to the recently described L. 
yami from Lanyu Island, Taiwan (Ota 1987). 


Materials and Methods 


Data were taken from the series of Lep- 
idodactylus from Batan Island (n = 14) and 


all other Philippine/Taiwanese species of the 
genus (see Specimens Examined). Addition- 
al information was taken from Brown & 
Alcala (1978). Nine meristic and 21 mor- 
phometric characters were used for com- 
parisons. The meristic characters are the 
number of: upper labials (UL), lower labials 
(LL), internasal scales (INS), interorbital 
scales (IOS), midbody scale rows (MSR), 
enlarged preanal and femoral scales (PFS), 
preanal and femoral pores in males (PFP), 
toe I scansors (TIS), toe IV scansors (TIVS). 
The mensural characters are: snout to vent 
length (SVL), head length (HL), head width 
(HW), head depth (HD), snout to eye length 
(SEL), eye diameter (ED), eye to ear length 
(EEL), internasal distance (IND), interor- 
bital distance (IOD), snout to arm length 
(SAL), axilla to groin length (AGL), body 
width (BW), body depth (BD), thigh length 
(THL), tibia length (TBL), toe I length (TIL), 
toe IV length (TIVL), toe IV width (TIVW), 
the length of scansor series beneath toe IV 
(SL), tail width (TW), and tail depth (TD). 
All morphometric characters were mea- 
sured to the nearest 0.1 mm with dial cal- 
ipers. Meristics were compared using Wil- 
coxon’s 2-sample test and morphometric 
characters were examined by principal com- 
ponent analysis, using the PRICOMP pro- 
cedure of SAS (1985) with correlation ma- 


560 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


t.. Philippines 2" 
8) 


Fig. 1. 


Map of the Philippines and Taiwan, showing the type locality of Lepidodactylus balioburius sp. nov. 


(Batan Island, inset), in relation to the distribution of its close relative L. yami (Lanyu Island, inset) and L. 
planicaudus (shaded portions in the central and the southern Philippines). 


trix. Skeletal characters were examined in 
radiographs. Museum acronyms follow 
Leviton et al. (1985). 


Lepidodactylus balioburius, new species 
Fis2 


Holotype. — Philippine National Museum 
(PNM) 984 (Original number USNM-FS 
121559), an adult male collected 2 km (by 
road) SE of Mahatao, Mahatao Municipal- 
ity, Batan Island, Batanes Province, Phil- 
ippines, on 6 Jun 1985, by Angel C. Alcala, 
Ven Samarita, and Braulio Gargar. 

Paratypes.—(n = 13, all from Batan Is- 
land). USNM 266559, 3 km NE of Basco, 
collected by Charles A. Ross & B. Gargar 


on 28 May 1985; OMNH 2349 (USNM-FS 
121200), 1-2 km E of Basco along road to 
Balugdh Bay, A. C. Alcala & D. Catada, 27 
May 1985; USNM 266560-61, 2.5 km ENE 
of Basco on W slope of Mt. Iraya, 150 m, 
Robert S. Kennedy & Fred G. Thompson, 
30 May 1985; OMNH 2348 (USNM-FS 
121372), Basco, C. A. Ross, 31 May 1985; 
USNM 266562, 1.5 km N of Basco, near 
airstrip, C. A. Ross & A. C. Alcala, 4 Jun 
1985; USNM 266563, Itbud, C. A. Ross & 
B. Gargar, 5 Jun 1985; USNM 266564, 3 
km ENE of Basco, W slope Mt. Iraya, 150 
m, C. A. Ross & R. S. Kennedy, 7 Jun 1985; 
CAS 162489, Mahatao, A. C. Alcala, 8 Jun 
1985; USNM 266565-67, CAS 162490, 2 
km E of Mahatao, C. A. Ross & A. C. Alcala, 


VOLUME 102, NUMBER 3 


561 


D 


Fig. 2. Holotype (PNM 984) of Lepidodactylus balioburius, adult male. A) dorsal view (scale = 10 mm), B) 
lateral and C) ventral views of snout (scale = 1 mm), D) ventral view of the preanal and femoral region, showing 


pores and enlarged scales (scale = 2 mm). 


6 Jun 1985. (five males and eight females, 
all adults). 

Etymology.—The specific name is de- 
rived from the Latin roots balius (brown) 
and burius (beast), an appropriate descrip- 
tor for this species. The name also acknowl- 
edges the significant contributions to Phil- 
ippine herpetology by Walter C. Brown and 
Charles A. Ross. 

Diagnesis.—A small (males 27.2-34.9, 


females 33.5—38.7 mm), bisexual, Group III 
(sensu Brown & Parker 1977) species of 
Lepidodactylus, characterized by slight but 
distinct digital webbing, relatively few en- 
larged scales and pores in the femoral/ 
preanal region, the rostral separated from 
the nostril by a scale, presence of lateral 
serration on the tail, and the absence of dis- 
tinctive pattern elements (see Figs. 2, 3). 
Description of holotype.—Habitus mod- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 3. 
& D). 


erately depressed, SVL 30.4 mm. Snout ta- 
pering, rounded at tip, length 3.9 mm. Eye 
diameter 2.1 mm. Internasal distance 1.6 
mm. Rostral separated from nostril by a 
small quadrangular scale. Nostril surround- 
ed by two supranasals, first upper labial, one 
small scale anteriorly and a slightly enlarged 
scale posteriorly. Anterior supranasals sep- 
arated by two small scales that border the 
rostral. Eleven upper labials on the right, 12 
on the left, the 9th beneath the center of the 
orbit; the last 2 only about twice as large as 
the surrounding scales. Eleven lower labials. 
Mental triangular, smaller than adjacent la- 
bials. Three to 5 rows of slightly enlarged 
scales on anterior part of chin. Scales on the 
snout larger than those on the dorsal surface 
of the body. Dorsal and lateral body scales 
very small, granular, with no enlarged tu- 
bercles. Forty-one interorbital scale rows at 
the midpoint of orbits, 131 scale rows at 
midbody. Ventral scales flat, cycloid, dis- 
tinctly larger than dorsals. 


Ventral view of toes and lateral tail margins of Lepidodactylus balioburius (A & C) and L. yami (B 


Length of extended hind limb 11.0 mm. 
Digits moderately dilated, distal three-fifths 
to three-quarters of undersurface bearing 
scansors as follows: fingers—I 7, II 8, III 10, 
IV 11 (ight) or 12 (left), V 8; toes—I 8, I 
9, II 11 eft) or 12 (ight), IV 9 (ight) or 
10 (left), V 8 (left) or 9 (right). Distal two 
to three scansors, including the terminal one, 
divided on all digits except the first. 
First digit with complete terminal and two 
divided subterminal scansors. All digits ex- 
cept the first clawed. Compressed claw- 
bearing phalanges arising from distal mar- 
gin of the dilated part and extending only a 
short distance beyond. Phalangeal formula 
of hand and foot 2-3-4-5-3. Webbing slight 
but evident between toes III and IV, ex- 
tending to about one-eighth to one-fifth 
length of toe IV. 

Twenty-three enlarged preanal and fem- 
oral scales bearing a continuous series of 21 
pores, extending over proximal 60% of thigh. 
Series of pore-bearing scales followed by one 


VOLUME 102, NUMBER 3 


row of slightly enlarged scales on thigh, and 
four to five rows of enlarged scales in pre- 
anal region. Two pairs of cloacal spurs on 
both sides of vent. Tail unregenerated, 
moderately depressed; its depth just pos- 
terior to the basal swollen area 73% of its 
width; lateral flange of skin lacking, but en- 
larged, spine-like scales present every five 
to seven marginal scales. Scales on the ven- 
tral surface of the tail slightly larger than 
those on dorsal surface. Interclavicle dag- 
ger-shaped, without lateral projections. 
Clavicles perforated. Twenty-six presacral 
vertebrae. Nasals fused at midline. 

Color in alcohol.—Dorsal ground color 
light grayish tan, with numerous minute dark 
dots; slightly darker areas on snout and be- 
tween orbits; a wide, indistinct, dark band 
from the tip of the snout, through the nos- 
tril, eye, along the dorsal margin of the ear, 
and fading out between the ear and fore- 
limb. Several dark spots on upper and lower 
labial regions. Indistinct rusty gray mark- 
ings forming vague dorso-lateral lines. Ven- 
ter creamy white, with minute blackish dots, 
much sparser than those on dorsum. Tail 
with 10 dark gray annular bands. 

Variation. — Variation in counts and 
measurements of the type series is presented 
in Tables 1 and 2. In the five specimens 
with regenerated tails the enlarged, spine- 
like lateral scales are absent. 

Coloration in the series is variable. In two 
specimens, the dorsal ground color is much 
darker and more rusty than the holotype, 
with a lighter middorsal region forming a 
broad longitudinal stripe. The dark dorso- 
lateral markings and annular bands on the 
tail are indistinct or absent in seven speci- 
mens. In two others, however, these mark- 
ings are more distinct than in the holotype. 
Four specimens have black spots on the lat- 
eral region of the original tail and on the 
neck. 

Natural history.—Specimens were col- 
lected in both disturbed and forested hab- 
itats from sea level to 150 m on Mt. Iraya. 
The field notes of C. A. Ross indicate that 


Table 1.—Comparison of nine meristic characters in L. balioburius sp. nov., L. yami, and other Philippine congeners. See the text for abbreviations. Data 


marked with an * taken from Brown and Alcala (1978:82-101 + table 6), but note that numbers in the text do not always agree with those in the table. In these 


cases, inclusive values were used. 


IOS MSR PFS PFP TIS TIVS 


INS 


LL 


UL 


Species 


10.00 


Jd 
0.83 
6-9 


24.33 


22.79 


40.00 138.57 


4.71 
0.47 


10.07 


yal 


L. balioburius 


0.68 


9-11 
HS Bee) 


Loy 
19-23 


a 
20-25 


S32 
131-151 


3.57 
34-46 
43.27 


0.83 


9-11 
10.93 


0.94 


10-14 


SD 


4-5 
4.20 


0.68 


range 


8.40 
0.74 
7-9 


pe) 19.63 


139.60 


AKO) 


L. yami 


1.28 
10-15 


1.92 
15-21 


1.51 
19-24 
18-26* 
20-28* 
30-40* 


32-40* 
28-35* 


oo 


126-151 


3.81 
36-49 


0.88 
10-12 


0.80 
11-14 
10-13* 
10-13* 
11-13* 
10-13* 
10-13* 


SD 


3-5 


range 


7-12* 
7-9* 
11-16* 
12-17* 
12-18* 


7-9* 
6-9* 


18-34* 
20-27* 
26-40* 


28-46* 
21-32* 


135-145* 
165-168 


34-42* 
40-41 
38-42 


3-5* 
4-5* 
3-5* 


3-5* 


10-12* 
10-12* 


range 


L. planicaudus 
L. christiani 


range 


9-10* 
9-11* 
9-13* 


120-140* 
78-108* 


120-140* 


9-13* 
9-13* 
9-12* 


range 


L. aureolineatus 


L. herrei 


24-34* 
32-40* 


range 


3-5* 


range 


L. lugubris 


563 


564 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Table 2.—Comparison of 21 morphometric characters of Lepidodactylus balioburius sp. nov. and L. yami, 
and the factor loadings on the first three principal components. See text for abbreviations. 


L. balioburius L. yami 
Characters x SD Range x SD Range PRIN I PRIN II PRIN III 
SVL 34.08 3.81 27.2-38.7 36.16 20 31.7-42.1 0.27 0.08 —0.94 
HL 8.69 0.67 7.6—9.6 9.47 0.69 8.6-10.9 O27 0-01 —0.03 
HW 6.36 0.47 5.6-7.1 6.76 0.95 6.0-9.8 0.25 0.15 0.10 
HD 3.59 0.28 3.1-3.9 4.23 Erg 3.4-7.1 0.25 0.04 =O313 
SEL 4.11 0:35 3.5-4.6 4.19 0.28 3.9-4.8 0.25 0.11 0.02 
ED 20> 0.20 2.0—2.6 2.29 0.23 2.0-2.6 0.23 0.02 0.13 
EEL 2:83 TOL 2.5-3.0 5.02. «uON3 2.8-3.4 0.24 —0.07 0.00 
IND 163 ©.13 1.4-1.8 1.68 0.14 1.5-2.0 0.23 0.08 0.06 
IOD 4.16 0.29 3.7-4.7 4.59 0.36 4.0-5.0 O:17, 023 =@.11 
SAL [2259 1.30 10.6-14.8 13.44 1.05 11.5-15.1 0.24 0.10 0.15 
AGL 16.19 23 12.7-19.3 17.81 1.82 14.9-21.5 0.24 0.04 —0.38 
BW 8.23 1.25 6.2-9.9 7.04 1.23 5.6-9.4 0.05 0.48 =0:35 
BD 4.23 0.86 2.8-5.4 5.25 Ipallig/ 4.0-8.6 0.23 0.01 —0.36 
THL 4.43 0.60 3.5-5.3 4.80 0.68 3.5-5.8 0.24 0.06 —0.10 
TBL 4.38 0.38 3.64.8 4.90 0.36 4.3-5.4 0.26" —0: 1 —0.09 
Te 1.65 0.21 1.42.1 1.95 0.16 1.7—2.2 O22- = =—O0H9 0.12 
TIVL 3305 0.37 2.4-3.7 3.72 0.34 3.0-4.2 O22 =" O71 0.29 
TIVW eal Osh 1.0-1.6 1.19 0.14 0.9-1.5 0.20 0.00 0.51 
SL BMD 0.36 1.6-2.9 2.10. “O21 2.2-2.9 Onl 94 —0:29 —0.00 
TW 4.51 0.64 2.9-5.6 3.50 0.45 2.7-4.1 0.02 0.53 0.11 
TD Zo 0.32 2.3-3.4 2.82) 10:37 2.1-3.6 0.06 0.44 0.34 
Eigenvalue 12.34 3.20 E15 
Difference oS 2.06 O23 
Proportion 0.59 0.15 0.05 
Cum. prop. 0.59 0.74 0.79 


the species was commonly found under loose 
bark on trees during the day, in disturbed 
areas of fields and gardens, in coastal vege- 
tation, and along a forested stream. Of those 
found after dark, one was active on the guest 
house in Basco, and another was on the un- 
derside of a banana leaf in forest on Mt. 
Iraya. Two eggs (USNM 266568), four 
hatched eggshells (not collected), and an 
adult were found under bark in a ravine 
running through a garden area, eventually 
leading to forest. Other gekkonids collected 
with L. balioburius include Hemidactylus 
frenatus, Gehyra mutilata, and Gekko po- 
rosus. Since L. balioburius is ecologically 
tolerant and not restricted to forest, it is 
likely that it will be found on other islands 


in the Batan group, none of which has been 


adequately collected to date. 
Distribution.—Known only from Batan 
Island, Batanes Province, Philippines, but 


expected on other islands in the group (see 
above). 

Remarks. —Brown & Alcala (1978) con- 
veniently separated the Philippine species 
of Lepidodactylus into two species com- 
plexes (=Sections), based largely on digital 
morphology, habitus, and caudal scalation. 
Section A (consisting of aureolineatus, her- 
rei and lugubris) was characterized by high 
scansor counts (11-18, usually more than 
12), the scansors covering most of the mod- 
erately to broadly dilated digits, slight dig- 
ital webbing, moderately depressed habitus, 
and a slightly to moderately flattened tail 
with lateral denticulation. Section B species 
(christiani, planicaudus) have fewer scan- 
sors (7-10, usually less than 10), confined 
to the distal half of broadly dilated and 
strongly webbed digits. The body and tail 
are strongly flattened, the latter with a broad 
flange of skin. 


VOLUME 102, NUMBER 3 


Ota (1987) placed his new species, L. yami 
from Lanyu Island, Taiwan, in Section A, 
but with some reservations. He also re- 
stressed the importance of the nostril po- 
sition relative to the rostral (in contact in 
Section A species, separated by a scale in 
christiani and yami, separated or in contact 
in planicaudus). Brown & Alcala (1978:81) 
dismissed this character as having “‘little 
significance at the species level,’ due to the 
variation in planicaudus, but our data in- 
dicate that it may be more useful than Brown 
& Alcala thought. 

Lepidodactylus balioburius shares the na- 
sal-rostral separation character with L. yami 
and the Section B species, but has little else 
in common with the latter. Consequently, 
it requires comparison only with Section A 
species. Within this group, L. balioburius 
has lower PFS, PFP, TIS, and TIVS counts 
than all species except L. yami (see Table 
1). Lepidodactylus aureolineatus and herrei 
are further distinguished by the presence of 
bright head stripes beginning on the snout 
and extending to near the ear. Lepidodac- 
tylus lugubris is a functionally all-female, 
parthenogenetic species; the few males re- 
ported to date have been sterile (Cuellar & 
Kluge 1972, Pasteur et al. 1987). In color 
and pattern, L. /ugubris is often very pale, 
almost white, with a variable pattern of dark 
spots, occasionally resembling those in the 
much darker L. balioburius. Lepidodactylus 
lugubris is capable of color change, how- 
ever. When in the dark phase, a more com- 
plex, ladder-like pattern becomes obvious, 
but this pattern is distinctly different from 
the plain brown, spotted balioburius. 

Lepidodactylus balioburius and L. yami 
share a number of characters unique in Sec- 
tion A species. Both are small, brown, un- 
specialized species isolated on small islands 
far to the north of their Philippine congeners 
(reports of L. /ugubris from Taiwan and as- 
sociated islands are possibly recent intro- 
ductions [Ota 1986, Cheng 1987]). Besides 
the similar habitus, coloration, and nostril- 
rostral separation, UL, MSR, PFS, and PFP 


565 


showed no statistically significant differ- 
ences between the two species (P = 0.05). 
Although the ranges overlapped somewhat, 
the means of LL, INS, IOS, TIS, and TIVS 
were significantly different (Table 1); INS of 
L. balioburius was larger than yami (P < 
0.05), whereas LL (P = 0.05), IOS (0.05), 
TIS (0.05), and TIVS (0.001) were larger in 
L. yami. Principal component analysis of 
morphometric characters revealed approx- 
imately 80% of the total variation in shape 
as expressed in the first three components— 
PRINs I, II and III. Of these, PRIN I was 
of little use in separating L. balioburius from 
L. yami. This component consists wholly 
of positive variable loadings, and is pri- 
marily a size component (Table 2). PRIN 
II, accounting for about 15% of the total 
variance, tends to discriminate L. balio- 
burius from L. yami more strongly. Variable 
loadings on this component revealed sev- 
eral characters chiefly contributing to the 
shape. The greatest proportion of the vari- 
ance on PRIN II was expressed by differ- 
ences in TW. The BW and TD were also 
heavily loaded, and followed by SL, IOD, 
TIVL, TIL and HW in descending order. 
Outlines of scatter plots of the component 
scores on PRINs I and II separated L. ba- 
lioburius from L. yami without overlap (Fig. 
4). The two species can also be distinguished 
by toe webbing and lateral caudal scalation, 
although these two characters are more sub- 
jective. Lepidodactylus balioburius has 
slightly more extensive webbing than L. 
yami (Fig. 3A, B) and the lateral denticu- 
lation of the original tail of balioburius con- 
sists of strongly enlarged, spine-like scales 
(Fig. 3C). In L. yami, slightly enlarged scales 
are present along the lateral tail edge, but 
they are neither projecting nor spinose (Fig. 
3D). We originally intended to summarize 
and discuss the relationships of Philippine- 
Taiwanese Lepidodactylus in this paper. 
Unfortunately, the unprecedented variation 
in L. planicaudus requires further re-eval- 
uation so we defer our taxonomic summary 
and key to a later paper. 


566 


9 -8 -7 6-5 -4 3 
PRIN | 


Fig. 4. Two-dimensional plots of scores of Lepidodactylus balioburius (closed circles) and L. yami (open 
circles) on principal components (PRINs) I and II. See Table 2 for the factor loadings of each component. 


The apparent absence of Lepidodactylus 
from the large island of Luzon is zoogeo- 
graphically puzzling. Although L. planicau- 
dus is found on small islands both east and 
southwest of Luzon (Brown & Alcala 1978, 
see also Fig. 1), the genus remains unre- 
ported from the entire large island; even the 
widespread human commensal L. lugubris 
is absent from its towns and cities. Although 
Luzon has been reasonably well collected 
(Taylor 1922; Brown & Alcala 1970, 1978), 
this hiatus may not be real. Many Philippine 
Lepidodactylus are ecologically restricted 
and difficult to collect. They may inhabit 
axils of palms or aerial ferns 10 meters or 
more from the ground. The montane areas 
of northern Luzon have never been ade- 
quately sampled and we would not be sur- 
prised if a new species allied to L. balio- 
burius and yami is eventually discovered in 
that area. 

Specimens examined.—L. yami: Osaka 
Museum of Natural History (OMNH) 
R2291 (holotype), 690, 691, R2855-61, 
California Academy of Sciences (CAS) 
158254 (paratypes), USNM 267943-44; L. 
planicaudus: CAS 60570, 128566, 139930, 


2-10 4 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


12 +3 +4 +5 +6 


139931, L. christiani: CAS 128877, 128878. 
L. aureolineatus: CAS-SU 28411, 26127, 
CAS 60226, 139941. L. h. herrei: CAS-SU 
24228, 26342. L. h. medianus: CAS 125239, 
131856 (paratypes). L. lugubris: OMNH 
R1772, 2201, 2202, 2320, 232 ers 
158255, 60595, 137835, Australian Mu- 
seum (AMS) R82602-82610, 82724-82730, 
109804—109809, 110141-110146, 110238- 
110242. 


Acknowledgments 


We are greatly indebted to Angel C. AI- 
cala, Walter C. Brown, and Charles A. Ross 
for their unstinting cooperation in provid- 
ing notes, unpublished data, and specimens 
for our use. We have enjoyed stimulating 
discussions on the Philippine fauna with the 
above individuals, Lawrence R. Heaney, 
Robert S. Kennedy, and Fred G. Thomp- 
son. Yasuhiko Shibata (OMNH), Jens V. 
Vindum (CAS), Harold Cogger and Allen 
E. Greer (AM) kindly loaned specimens for 
comparative purposes. Special thanks are 
due Tsutomu Hikida for computing the data, 
using the facilities of the Data Processing 


VOLUME 102, NUMBER 3 


Center, Kyoto University. Radiographs were 
made with facilities of Biological Labora- 
tory, Yoshida College, Kyoto University 
through the courtesy of Masafumi Matsui. 

The Batan field work was made possible 
by a Smithsonian Research Opportunity 
Fund award to C. A. Ross. Crombie’s mu- 
seum work on the Pacific herpetofauna was 
also supported by two ROF grants, for which 
David Challinor has our thanks. Ota’s work 
was supported in part by a Grant-in-Aid for 
Special Project Research on Biological As- 
pects of Optimal Strategy and Social Struc- 
ture from the Japan Ministry of Education, 
Science and Culture. 

We also thank W. Ronald Heyer and 
George R. Zug for their critical reading of 
the manuscript and Linda K. Gordon for 
valuable assistance in producing the final 
manuscript copy. 


Literature Cited 


Brown, W. C., & A. Alcala. 1970. The zoogeography 
of the herpetofauna of the Philippine Islands, a 
fringing archipelago.— Proceedings of the Cali- 
fornia Academy of Sciences 38:105-—130. 

—,, & 1978. Philippine lizards of the 
Family Gekkonidae. Silliman University Nat- 
ural Science Monograph Series (1), Dumaguete 
City, Philippines, 146 pp. 

—., & F. Parker. 1977. Lizards of the genus Lep- 
idodactylus from the Indo-Australian archipel- 
ago and the islands of the Pacific, with descrip- 
tions of new species.— Proceedings of the 
California Academy of Sciences 41:253-265. 


567 


Cheng, H.-Y. 1987. The record of a gekkonid lizard 
Lepidodactylus lugubris (Dumeril and Bibron, 
1836) from Taiwan.—Journal of the Taiwan 
Museum 40(1):85-89. 

Cuellar, O., & A. G. Kluge. 1972. Natural partheno- 
genesis in the gekkonid lizard Lepidodactylus 
lugubris.—Journal of Genetics 61:14—26. 

Leviton, A. E., R. H. Gibbs, Jr., E. Heal, & C. E. 
Dawson. 1985. Standards in herpetology and 
ichthyology: Part I. Standard symbolic codes for 
institutional resource collections in herpetology 
and ichthyology.—Copeia (3):802-832. 

Ota, H. 1986. The mourning gecko Lepidodactylus 

lugubris (Dumeril and Bibron, 1836); an addi- 

tion to the herpetofauna of Taiwan. —Journal of 
the Taiwan Museum 39(1):55—58. 

. 1987. Anew species of Lepidodactylus (Gek- 

konidae: Reptilia) from Lanyu Island, Tai- 

wan.—Copeia (1):164-169. 

Pasteur, G., J.-F. Agnese, C. P. Blanc, & N. Pasteur. 

1987. Polyclony and low relative heterozygos- 

ity in a widespread unisexual vertebrate, Lepi- 

dodactylus lugubris (Sauria).—Genetica 75:71- 

79. 

1985. SAS user’s guide: Statistics, version 5. 

SAS Institute Inc., Cary, North Carolina. 

Taylor, E. H. 1922. The lizards of the Philippine 
Islands. — Philippine Bureau of Science Publ. No. 
17:1-269. 


SAS. 


(HO)Department of Zoology, Faculty of 
Science, Kyoto University, Kitashirakawa, 
Sakyo, Kyoto, 606 Japan (Present address, 
Department of Biology, University of the 
Ryukyus, Nishihara-cho, Okinawa 903-01, 
Japan); (RIC) Department of Vertebrate 
Zoology (Amphibians & Reptiles), National 
Museum of Natural History, Smithsonian 
Institution, Washington, D.C. 20560. 


PROC. BIOL. SOC. WASH. 
102(3), 1989, pp. 568-576 


A NEW SPECIES OF EUPSOPHUS 
(AMPHIBIA: ANURA: LEPTODACTYLIDAE) FROM 
SOUTHERN CHILE 


J. Ramon Formas 


Abstract.—Eupsophus emiliopugini, a new species of leptodactylid frog, is 
described from southern Chile. This species is distinguished from its congeners 
by the olive-green cephalic pigmentation. The mating call is composed of two 
notes and the tadpole is typically an inhabitant of ground water-filled cavities. 
This frog occurs from eastern Osorno Province through Llanquihue, Chiloé, 


and Aisén provinces. 


The genus Eupsophus Fitzinger, 1843 is 
restricted to the temperate Nothofagus for- 
est of the southwestern border of South 
America. At present five species have been 
described (E. roseus, E. vittatus, E. calca- 
ratus, E. migueli, and E. insularis) (Formas 
& Vera 1982, Formas 1985), and for this 
reason Eupsophus can be considered the 
most diverse genus within the reduced an- 
uran fauna of the temperate forest system 
of southern Chile and Argentina. Among 
the genera of the sub-family Telmatobiinae, 
Eupsophus is remarkable by having tad- 
poles (E. roseus, E. vittatus, and E. calca- 
ratus) which live in small water-filled cav- 
ities in the ground. The larval mouthparts 
(denticles) are reduced, and the larvae feed 
only upon yolk reserves (Formas & Pugin 
1978a, b; Formas 1989). 

Between September 1975 and December 
1987, Carlos Varela, Lila Brieva, Gonzalo 
Aguilar and I collected specimens of a new 
species of Eupsophus in the temperate Noth- 
ofagus forest of southern Chile. In this paper 
the new taxon is described on the basis of 
a series of materials (adults, juveniles, mat- 
ing call, eggs, and tadpoles) collected from 
seventeen localities. 


Eupsophus emiliopugini, new species 
Fig. 1 


Holotype. —IZUA (Instituto de Zoologia, 
Universidad Austral de Chile) 1587, adult 


male; Ramon Formas, Nov 28, 1975, at La 
Picada, 23 km NE (by road) of Ensenada 
(Fig. 2), Osorno Province, Andean Range, 
41°04’S, 72°26’W, 480 m. 

Paratypes. —Seven adult males from the 
type locality: IZUA 1585-86, 1593, 1596, 
1602, 1607-8. 

Diagnosis.—Eupsophus emiliopugini is a 
medium-sized frog which is characterized 
by having a distinctive olive-green band be- 
tween the eyes, which is absent in E. roseus, 
E. calcaratus, E. vittatus, E. migueli and E. 
insularis. Furthermore it differs from E. vit- 
tatus in snout—vent length (Table 1). The 
mating call of E. emiliopugini has two notes 
while that of E. vittatus possesses five (4—6) 
notes (Table 2). 

Adult description. —(Based on 25 living 
frogs and 29 fixed specimens.) Head wider 
than long. Snout pointed in dorsal view, 
slightly sloping in lateral profile; canthus 
rostralis indistinct and rounded; loreal re- 
gion slightly concave; nostrils dorso-lateral, 
closer to tip of snout than to the orbit; length 
of the eye greater than distance between eye 
and nostril; inter-orbital distance narrower 
than length of eye, greater than internarial 
distance. Tympanic membranae medium 
and well defined, tympanum diameter three 
fourths the distance between eye and nostril. 
Well developed supratympanic fold extend- 
ing posteriorly from the corner of the eye 
to the posterior part of the tympanum, cov- 


VOLUME 102, NUMBER 3 


569 


Figc ft. 


ering upper part of tympanic annulus but 
not reaching insertion of arm. Tongue large, 
ovoid, with notch at the tip. Choanae small, 
round dentigerous process of vomers lying 
slightly below choanae; each process bear- 
ing 3-4 slightly oblique teeth close to me- 
dian line. Forelimbs thin. First finger equal 
in length to second; third finger much longer 
than fourth; digital length in decreasing or- 
der 3-4-2-1. Palmar webbing absent; tips of 
fingers rounded and slightly prominent. In- 
ner median palmar tubercle ovoid; outer 
palmar tubercle horseshoe-shaped; subar- 
ticular tubercles globular and moderate in 
size; supernumerary palmar tubercles pres- 
ent. Hind limbs slender. Toes long, slender, 
and moderately fringed; tips of toes round- 
ed; third and fifth toes equal in length; toes 
in decreasing order of length 4-(3,5)-2-1. In- 
ner metatarsal tubercle ovoid and promi- 
nent, outer rudimentary. Subarticular tu- 
bercles ovoid; supernumerary tubercles 
absent. Tarsal fold absent. Rudiment of web 
among toes. Anal opening oriented trans- 
versely and directed postero-ventrally at 
dorsal level of thighs. Dorsal and ventral 
skin smooth. Two weakly developed para- 


Eupsophus emiliopugini, new species. Holotype (IZUA 1587). 


vertebral folds extending from posterior part 
of head, converging behind it. Post-tym- 
panic areas and flanks with scarce minute 
granules. Some specimens with slight gran- 
ular area around vent and posterior part of 
thighs. External measurements of males and 
females of this species shown in Table 1. 

Coloration in preservative. —Dorsal 
ground color light gray with few whitish mi- 
nute spots; a delicate vertebral line extend- 
ing from the tip of snout to the vent. A dark 
greenish band on the eyes. Venter whitish 
and gular area gray. Lips gray with two or 
three irregular spots reaching the tympanic 
region. Arms light gray and crossed by two 
or three irregular transverse bars of dark 
gray color in dorsal area; arms and legs whit- 
ish ventrally. 

Coloration in life. —Dorsal ground color 
grayish brown to leaden; vertebral line lem- 
on-yellow. Band on eyes olive-green col- 
ored. Some specimens with bright yellowish 
reticulations on the thighs. Belly whitish and 
the gular area of mature males bright or- 
ange. 

Distribution. —The presently known range 
of E. emiliopugini extends from northeast- 


570 


Table 1.—Comparative measurements (mm) of Eupsophus emiliopugini and E. vittatus. Means, SD, and 


ranges (parenthesis). 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


E. emiliopugini E. vittatus 

Character Females Males Females Males 

n 14 40 19 19 
Snout—vent length SOLO == 5:01 46.71 + 24.00 59 Olne=s6. 58 55.44 + 8.68 
(41.0-64.0) (42.9-50.0) (47.0-71.8) (44.5-66.6) 
Head length PES See yl 282 16.45 + 0.69 20.43 + 1.50 £3.36 2 23350 
(13.4—20.8) (14.8-18.1) (16.7-23.9) (15.4-21.3) 
Head width PGES Sect Deng 18.48 + 0.33 23:66 = 2:02 DAT 2 E5238 
(15.9-25.6) (16.9-19.1) (19.9-26.6) (18.1-26.0) 
Femur length 23:38 2247 2S6 == 06 28.45 + 1.89 26.37 = 3:85 
(17.1-29.2) (16.6-24.3) (24.7-31.2) (21.7-31.6) 
Tibia length DAD = 2) 24.25 + 4.43 D8: 985 2.05 27.2 + 4.18 
(19.9-29.6) (19.30-33.1) (25.0-32.2) (22.8-33.1) 
Foot length Soe ee S50 365222110 41.83 + 3.56 Al .26 225337 
(29.1-44.9) (29.7-49.1) (34.4-48.2) (33.8-49.1) 


ern Osorno Province (Termas de Puyehue) 
to Aisén Province (Caleta Vidal) (Fig. 2). 
This area is covered by humid and cool 
Nothofagus forests, which are found at the 
Andes Cordillera (below 1000 m), the 
Coastal Range, the Central Valley, and on 
Chiloé Island. The altitudinal distribution 
of E. emiliopugini ranges between the sea 
level (Caleta Vidal) and 700 m in the Andes 
Cordillera (Termas de Puyehue). Over much 
of its range, E. emiliopugini occurs sym- 
patrically with E. calcaratus. It has never 
been found in sympatry with E. vittatus. 

Habitat. —La Picada (type locality) (Fig. 
2) is a small subandean valley surrounded 
by forests (Nothofagus, Aextoxicum, and 
Eucryphia). During winter and summer, 
frogs were found there under logs, however 
during the reproductive period (springtime) 
the animals were collected at the border of 
a small stream shaded by ferns (Dryopteris) 
and mosses (Sphagnum and Hygroambly- 
stegium). 


From the ecological point of view this 
area 1s situated in the oceanic region of med- 
iterranean influence (di Castri 1968). The 
annual mean temperature of this region is 
10.5°C the relative humidity is 84% and the 
rainfall ranges between 2000 to 2500 mm. 
Of seventeen examined localities, fifteen are 
included in the preceding region; however 
the southernmost localities (Rio Cisnes and 
Caleta Vidal) are situated in the oceanic cold- 
temperate region (di Castri 1968). The an- 
nual mean temperature is 8.8°C, the relative 
humidity is 87% and the rainfall ranges from 
2500 to 3000 mm. 

At the type locality the following species 
of amphibians were also collected: Rhino- 
derma darwinii, Bufo variegatus, Batrachyla 
leptopus, B. antartandica, Alsodes monti- 
cola, Pleurodema thaul, Hylorina sylvatica 
and Eupsophus calcaratus. 

Breeding sites and breeding season. —Ten 
breeding sites were examined (Termas de 
Puyehue, La Picada, El Traiguén, Rio Rol- 


— 


Fig. 2. Distribution of Eupsophus emiliopugini (dots) and E. vittatus (triangles) in Southern Chile. Arrow 
indicates location of the type locality (La Picada) of E. emiliopugini. 1) Ramadillas, 2) Contulmo, 3) Mafil, 4) 
Linguento, 5) Mehuin, 6) San Martin, 7) Los Molinos, 8) Valdivia, 9) Huellelhue, 10) Tres Chiflones, 11) 
Cordillera Pelada, 12) Bahia Mansa, 13) Pucatrihue, 14) Puyehue, 15) Piedras Negras, 16) El Traiguén, 17) La 
Picada, 18) Rio Rollizo, 19) Rio Lenca, 20) Camino Maullin, 21) Ancud, 22) Lechagua, 23) Chepu, 24) Cucao, 
25) Puntra, 26) Tepuhueico, 27) Yaldad, 28) Quellon, 29) Rio Cisnes, 30) Caleta Vidal. 


VOLUME 102, NUMBER 3 


@ Frutillar 
Llanqu/hue 


Lake 


© Valdivia 
At0 


<a 
2 
= 
2 
Lid 
©) 
a 
<q 


Bas 


571 


ah2 


a 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


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 


VOLUME 102, NUMBER 3 


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, 


573 


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 


574 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


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 


VOLUME 102, NUMBER 3 


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, 


575 


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- 


576 


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 


578 


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. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


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). 


VOLUME 102, NUMBER 3 


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. 


580 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


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 


VOLUME 102, NUMBER 3 


581 


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 


VOLUME 102, NUMBER 3 


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. 


VOLUME 102, NUMBER 3 


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. 


VOLUME 102, NUMBER 3 


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. 


591 


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. 


392 


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]. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


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 


VOLUME 102, NUMBER 3 


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. 


594 


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 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


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 


VOLUME 102, NUMBER 3 


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 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Figs. 22-27. 
and different localities, lateral views: 22, Belize, Stann Creek District, Carrie Bow; 23, USA, North Carolina, 
Wrightsville; 24, USA, North Carolina, Wrightsville; 25, Mexico, Tabasco, Paraiso (5 km N); 26, Antigua, 
Dutchman Bay; 27, Dominica, David Bay. 


type 2.15 mm). Coloration generally brown 
dorsally and gray laterally. 

Head: Frons generally brown to oliva- 
ceous or slightly charcoal brown, coloration 
of frons uniform or ocellar triangle more 
grayish, and fronto-orbits charcoal gray. 
Ocellar setae divergent, lateroclinate; in- 
trafrontal setae anteroclinate, slightly con- 
vergent; ocellar area with three or four 
smaller setulae. Face, clypeus, and gena (to 
an extent) whitish, with faint tinges of blue 
or gray, gena becoming darker posteriorly, 
more charcoal gray. Orientation and size of 
genal setae as follows (anterior to posterior 
setae): lst seta large, anteroclinate; two large 
anaclinate setae with one, rarely zero or two 
anaclinate setulae between larger anaclinate 
setae; anaclinate setula about one-third 
length of larger setae. 


External male terminalia of Nocticanace texensis: Variation in shape of the surstylus from same 


Thorax: Mesonotum from dorsal view 
mostly brown to olivaceous brown, darker 
around anterior margins; postpronotum, 
anterior half of notopleuron, and to a lesser 
degree the posterior fourth of scutum, and 
scutellum gray; coloration of thorax in lat- 
eral view brown dorsally, becoming gray 
ventrally around area of notopleuron (in 
some specimens just above, others at or be- 
low notopleuron), remainder of pleural areas 
mostly unicolorous, gray. Notopleuron 
bearing only one seta, inserted posteriorly; 
anepisternum with three large setae and from 
three to five smaller setulae between larger 
ones, larger setae with insertions and ori- 
entations as follows: posterior seta, usually 
the largest, with posteroclinate orientation, 
ventral seta, usually the next largest, with 
declinate orientation, dorsal seta smallest 


VOLUME 102, NUMBER 3 


(not much larger than setulae) and with an- 
aclinate orientation; one large katepisternal 
seta and one or two smaller setulae inserted 
ventrad or anteroventrad. Fore femur with 
four or five long setae along posteroventral 
margin and four or five shorter setae pos- 
terodorsally; mid tibia bearing apicoventral 
spinelike seta, length about equal to tibial 
width; hind basitarsomere lacking basoven- 
tral pair of larger setae; legs mostly concol- 
orous, gray to dark gray apically. Wing gen- 
erally lightly infumate, brownish; costal vein 
ratio averaging 0.13 (holotype 0.15); M vein 
index averaging 0.43 (holotype 0.49); apical 
section of vein CuA, slightly more than twice 
length of crossvein dm-cu. 

Abdomen: Dorsum mostly olivaceous 
brown, gray laterally and ventrally. Male 
terminalia (Figs. 22—27) as follows: poste- 
rior margin of surstylus densely setulose, 
posterior half of surstylus in lateral view 
roughly triangular, distinctly angulate to 
form a large and wide posterior projection 
and a tapered, more slender process ven- 
trally (Sometimes curved inward), anterior 
margin with a subapical emargination (see 
remarks section below for discussion of 
variation). 

Type material.—The holotype male is la- 
beled “‘Galveston[,] Tex[as] 9.13.50 [hand- 
written|/M. R. Wheeler Collectors/Holo- 
type Canaceoides texensis Wheeler 1952 
[red; handwritten].’’ The allotype female (not 
examined) is reported to bear the same lo- 
cality label data as the holotype. The ho- 
lotype is double mounted (glued to a paper 
point), is in excellent condition (the abdo- 
men has been removed, dissected, and the 
parts are stored in an attached plastic mi- 
crovial), and is deposited in the California 
Academy of Sciences. 

Other specimens examined.— Antigua. 
Dutchman Bay, 7 Jan 1965, W. W. Wirth 
(4 6; USNM). Belize. Stann Creek District: 
Carrie Bow, 4 Mar 1984, 2 Jun 1985, 15- 
27 Jan 1987, W. N. Mathis, C. Feller (18 4, 
9 2; USNM). Dominica. Calibishie (sea 
shore), 27 Feb—22 Mar 1965, 1989, W. N. 


597 


Mathis, W. W. Wirth (15 6, 3 2; USNM); 
Coulibistri, beach, 21 Mar 1989, W. N. 
Mathis (23 6, 10 2; USNM); Layou River 
(mouth), 9 Jan 1965, W. W. Wirth (1 2; 
USNM); Macoucheri (sea shore), 1 Feb 
1965, W. W. Wirth (1 2; USNM); Pagua 
Bay, 18 Feb 1965, W. W. Wirth (6 4, 5 2; 
USNM); Rosalie (cobble beach), 23 Mar 
1989, W. N. Mathis (7 3, 4 2; USNM); Sou- 
friére Bay, 24 Mar 1989, W. N. Mathis (9 
6, 6 2; USNM); St. David Bay (sea shore), 
23 Jan 1965; W. Ws Wirth (15. 6, 10.9; 
USNM). Mexico. Tabasco Province: Para- 
iso (5 km E), 6 May 1985, W. N. Mathis, 
A. Freidberg (26 6, 25 2; USNM). St. Vin- 
cent. Buccament Bay, 25-28 Mar 1989, W. 
N. Mathis (7 6, 4 2; USNM); Cumberland 
Bay, 28 Mar 1989, W. N. Mathis (4 34; 
USNM); Wallilabou (beach), 27 Mar 1989, 
W. N. Mathis (8 6, 4 2; USNM). United 
States. North Carolina: Wrightsville Beach 
(light trap), 3-7 Sep 1953, R. H. Foote (9 
6, 23 2; USNM). 

Distribution. —East (North Carolina 
south) and Gulf coasts of North America, 
West Indies (Antigua, Dominica, and St. 
Vincent), and Caribbean coast of Mexico 
and Belize. 

Natural history.—All of the specimens. 
from Belize were collected on Carrie Bow 
Cay, formerly Ellen Cay, which is a highly 
disturbed, vegetated sand cay that is about 
18 km off the coast of Hopkins, Stann Creek 
District. The cay is small, 100 by 40 m, and 
the former woodland, mostly red man- 
grove, has been cleared. Twenty-two species 
of plants have been recorded from Carrie 
Bow, including a few that have been intro- 
duced. Stoddart et al. (1982) and a more 
recent paper by Rutzler and Ferraris (1982) 
provide more extensive information on 
Carrie Bow and its habitats. 

The specimens of N. texensis were col- 
lected by sweeping an aerial net with a fine- 
meshed bag immediately over and between 
rocks and exposed coral on the surrounding 
beach, which is mostly sand covered, es- 
pecially on the north, east, and south sides. 


598 


Remarks.—Externally, this species is very 
similar to N. wirthi but can be distinguished 
from it and other congeners by the following 
characters: anterior notopleural seta absent; 
acrostichal setae absent; apical scutellar se- 
tae nearly straight in lateral view, not dis- 
tinctly curved dorsally (anaclinate); and 
shape of the male terminalia, especially the 
unique surstylus (see figures). 

The shape of the surstylus is quite similar 
to that found in males of N. arnaudi Wirth 
(the galapagos group), but specimens of the 
latter are usually larger (3-3.5 mm), are 
much darker brown dorsally with the dark 
coloration extended ventrally to the dorsal 
third of the anepisternum, and the noto- 
pleuron usually bears an anterior and pos- 
terior seta, although the latter seta is usually 
much better developed. 

The shape of the surstylus also differs 
slightly from locality to locality (see Figs. 
22-27 from different localities), and I in- 
terpret this to be intraspecific variation, per- 
haps clinal in nature. The differences found 
in populations from the Lesser Antilles, 
however, especially those from Dominica, 
could represent interspecific variation. The 
surstylus from a male collected in Antigua 
(Fig. 26), for example, appears to be some- 
what intermediate between that found on 
Dominica (Fig. 27) and the more typical 
shape, as found in males from North Car- 
olina, Texas, Mexico, and Belize (Figs. 22- 
25). Unfortunately no specimens are avail- 
able from the Greater Antilles that could 
provide important evidence to help resolve 
this issue, and for the present, I prefer to 
recognize a single species. 


Nocticanace wirthi, new species 
Fig. 28 


Nocticanace texensis [of authors, not 
Wheeler].— Wirth, 1954:61-62 [illustra- 
tion of male terminalia]; 1965:734 
[nearctic catalog]. 


Diagnosis.—As in the preceding species 
except as follows: small to moderately small 
beach flies, length 1.75 to 2.25 mm. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Abdomen: Male terminalia as follows 
(Fig. 28): surstylus in lateral view with two 
large, ventral lobes; posteroventral lobe 
wide, pointed anteroventrally, and densely 
setulose, especially along outer margin; an- 
terior lobe blunt, thumblike, and setulose. 

Type material.— The holotype male is la- 
beled “‘Boyton Beach, Fl[orid]a [Palm Beach 
Co.]/WWWirth Collector/10 VIII 51 [10 
Aug 1951] Intertidal rocks/é/Nocticanace 
texensis (Wheeler) det WWirth [species 
name handwritten; black sub-border]/HO- 
LOTYPE 4 Nocticanace wirthi Mathis 
USNM [species name handwritten; red].” 
The allotype female and 14 paratypes (7 4, 
7 2; CAS, USNM) bear the same locality 
label data as the holotype. Other paratypes 
are as follows: Cuba. Matanzas Province: 
Playa Larga, 1 May 1983, W. N. Mathis (3 
6, 9 2; IZAC, USNM). Bermuda. Smith’s 
Parish: Spittal Pond (intertidal rocks), 18 
Nov 1987, N. E. Woodley (21 6, 25 2; BDAF, 
USNM). The holotype is double mounted 
(minute nadel in a cork block), is in good 
condition, and is in the Smithsonian Insti- 
tution (USNM). 

Distribution.—Bermuda, Cuba, 
southern Florida. 

Natural history.—Wirth (1954:62) re- 
ported that the type series was “... col- 
lected from a shelf of limestone rock about 
a hundred yards long on the Atlantic Ocean 
beach. The rock projected from the water 
only at low tide and was covered with a 
scanty growth of filamentous green algae.” 
The specimens collected in Cuba were swept 
from a large limestone outcrop that extend- 
ed into the sea. The limestone was deeply 
eroded, and the exposed surface had nu- 
merous sharp ridges. 

Etymology.—It is a pleasure to name this 
species after Dr. Willis W. Wirth, who has 
contributed substantially to our knowledge 
of beach flies and who first illustrated this 
species. 

Remarks. —This species is very similar to 
N. texensis, and the only distinguishing 
characters that I have found between these 
two species are those of the male terminalia, 


and 


VOLUME 102, NUMBER 3 


599 


Figs. 28-29. Lateral view of male terminalia: 28, Nocticanace wirthi; 29, Noticanace panamensis. 


especially the bilobed shape of the surstylus 
(see description and figures). Externally the 
specimens are virtually impossible to sep- 
arate from those of N. texensis. 


Nocticanace panamensis, new species 
Fig. 29 


Diagnosis.—As in N. texensis except as 
follows: moderately small beach flies, length 
2.0 mm (based on the single pinned 2 para- 
type). 

Abdomen: Male terminalia as follows 
(Fig. 29): surstylus in lateral view more or 
less rectangular, posterior margin conspic- 
uously setulose; inner lobe not developed as 
a process, bearing several setulae along me- 
dian surface; outer lobe a large, thumblike 
posteroventral process that bears several 
setulae. 

Type material.— The holotype male is la- 


beled ‘“[Panama] Mojinga Swampf,] Ft. 
Sherman, C. Z. Jan. 1953[,] F. S. Blanton 
ct. [collector] Nocticanace texensis (Whee- 
ler) 6 5 [°5” is circled][handwritten].’’ The 
allotype female (double mounted) and three 
paratypes (3 2; USNM;; slide mounted) bear 
the same label data as the holotype. The 
holotype is slide mounted in balsam (most 
body parts have been dissected and are sep- 
arated) and is in the Smithsonian Institution 
(USNM). 

Distribution.— Panama. Fort Sherman is 
located at the Caribbean mouth of the Pan- 
ama Canal. 

Etymology.—The specific epithet, pana- 
mensis, refers to the country of Panama, 
where the type series was collected. 

Remarks.—Like N. wirthi, this species dif- 
fers from N. texensis in characters of the 
male terminalia, especially the shape of the 
surstylus (see figures and description above). 


600 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


ey» 
NY 


wy, 
Af 

ACA 
, 


~ 


Figs. 30-38. Scanning electron micrographs of Paracanace hoguei: 30, Head, lateral view; 31, Gena and 
setae, lateral view; 32, Head, dorsal view; 33, Antenna, lateral view; 34, Ocellar triangle, dorsal view; 35, Fronto- 
orbital setae, dorsal view; 36, Notopleuron and setae, lateral view; 37, Katepisternum and setae, lateral view; 


38, Scutellum, dorsal view. 


Genus Paracanace Mathis & Wirth 


Paracanace Mathis & Wirth, 1978:524. 
Type species: P. hoguei Mathis & Wirth, 
by original designation. 


Diagnosis.—Small to moderately small 
beach flies, 1.40 to 2.60 mm; general col- 
oration whitish gray to brownish black. 

Head: Intrafrontal setae two pairs; post- 
ocellar setae well developed, proclinate and 
very slightly divergent, subequal in length 
to intrafrontal setae; ocelli arranged to form 
isosceles triangle, with greater distance be- 
tween posterior ocelli. Two to three large 


anaclinate genal setae; anteroclinate genal 
seta well developed, subequal in length to 
larger anaclinate genal setae; epistomal 
margin sinuate; clypeus low, width more 
than 4x height; palpus yellowish. 

Thorax: Acrostichal setulae present, in 
about four rows, with a distinctly larger pre- 
scutellar pair; scutellar disc lacking setae; 
apical scutellar setae not anaclinate; ante- 
rior notopleural seta usually present (very 
weak or absent in one species); proepisternal 
seta(e) present; anepisternum with scattered 
setulae; katepisternal seta present. Femora 
and tibiae grayish black; tarsomeres yellow- 


VOLUME 102, NUMBER 3 


ish brown to dark brown, becoming darker 
apically; mid femur of male with comblike 
row of setae; hind tibia lacking spinelike 
setae apically. Wing with length of apical 
section of vein CuA, twice or more that of 
crossvein dm-cu; vein M index 0.38. 

Discussion.— Like Canacea, all of the de- 
scribed species of Paracanace occur in the 
Western Hemisphere, with primarily trop- 
ical or subtropical distributions (Mathis and 
Wirth 1978). 

I have recognized two species groups 
within Paracanace. This arrangement ad- 
heres to the cladogram for the species of this 
genus that Mathis and Wirth (1978:535) 
published. The two species groups are char- 
acterized in the key that follows. All known 
Caribbean species belong to the hoguei group 
(Figs. 30-38). 


Key to Species Groups of Paracanace 


1. Fore femur of male with from three 
to four long setae along posteroven- 
tral surface, setal length subequal to 
2x femoral width; mid femur of 
male bearing a posteroventral, 
comblike row of setae along entire 
length, setae at proximate one-fourth 
pale; costal vein between humeral 
crossvein and subcostal break usu- 
ally bearing a row of long spinelike 
setae, setal length subequal or great- 
er than width of Ist costal cell 

oy ae The hoguei Group 

— Fore femur of male lacking three or 
four setae as described above; mid 
femur of male bearing a postero- 
ventral comblike row of setae along 
distal one-half only; setae along an- 
terior margin of wing much shorter, 
not more than one-half width of Ist 
costal cell The maritima Group 


Key to Species of the hoguei Group 


1. Three, dorsally curved, genal setae 
subequal in length; body strongly se- 


tose (Figs. 30-38) (Cocos Island. 
Costa Rica) 
cy ae eee ee ae P. hoguei Mathis & Wirth 
— Middle, dorsally curved, genal seta 
about one-half length of setae on 
either side; body moderately setose 


2. Surstylus broader on distal half, es- 
pecially evident in lateral view; ven- 
tral, surstylar margin broadly trun- 
cate in lateral and posterior views; 
posterior margin of surstylus bear- 
ing distinct row of longer setae 
(Florida, Jamaica) 48) OF er ee. . 

He Bs a P. lebam Mathis & Wirth 

— Surstylus in lateral view swollen 
along anterior margin near middle, 
tapered ventrally to broadly round- 
ed, ventral margin; posterior margin 
of surstylus lacking distinct row of 
longer setae; posteroventral angle of 
surstylus noticeably produced api- 
cally (Figs. 39, 40) (Belize, Cuba, 
Dominica) .. P. aicen Mathis & Wirth 


Paracanace aicen Mathis and Wirth 
Figs. 39-42 


Paracanace aicen Mathis & Wirth, 1978: 
533. 


Specimens examined.—Belize. Stann 
Creek District: Carrie Bow Cay, 15 Jan—2 
Jun 1984-1987, W. N. Mathis, C. Feller (19 
6, 25 2: USNM); South Water Cay, 1 Jun 
1985, W. N. Mathis (2 6, 6 2; USNM). 

Distribution.— United States (Florida: Lee 
Co., Sanibel Island), Belize, West Indies 
(Cuba, Dominica, and St. Vincent). 

This the first record of Paracanace from 
the United States, although earlier, Mathis 
(1988:330) had included this genus in a key 
to the beach-fly genera from the United 
States with the expectation that the genus 
was likely to be found there. The new lo- 
calities from the western Caribbean sub- 
stantially increase the known distribution 
of this species, which previously was known 
only from Dominica in the Lesser Antilles. 


602 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


4] 


Figs. 39-42. External male terminalia of Paracanace aicen: 39, Epandrium and surstylus (holotype from 
Dominica), lateral view: 40, Epandrium and surstyli (holotype from Dominica), posterior view; 41, Epandrium 
and surstylus (Florida, Lee Co., Sanibel Island), lateral view; 42, Epandrium and surstyli (Florida, Lee Co., 
Sanibel Island), posterior view. 


VOLUME 102, NUMBER 3 


The species may eventually be found to be 
circumcaribbean, but the paucity or absence 
of canacid collections from Colombia, Ven- 
ezuela, and most countries of Central Amer- 
ica precludes assessment of this possibility. 

Natural history.—The specimens from 
Florida were collected on the causeway be- 
tween the Gulf Coast of Florida and Sanibel 
Island (Lee Co.). The sides of the causeway, 
particularly the south side, were partially 
lined with broken pieces of concrete and 
large rocks to moderate the erosive action 
of waves. Much of the surface of the con- 
crete and rocks was covered with algae. In 
addition to this species, beach flies of the 
following two species were also found in this 
habitat: Procanace dianneae (very abun- 
dant; see treatment below) and Canacea 
macateei (uncommon; see treatment above). 

Although this species and Nocticanace 
texensis both occur on Carrie Bow Cay and 
are found in essentially the same habitats, 
I did not collect the two species together 
during a particular season. In the Lesser An- 
tilles (Dominica and St. Vincent), I found 
the two species in the same sweep of the net, 
although in very unequal numbers. Where 
one species was common, the other was not. 
Whether this is an artifact of sampling or is 
indicative of temporal partitioning needs 
further investigation. Indeed, to what de- 
gree these two species may compete for the 
same or similar resources merits closer scru- 
tiny. 

See comments under the treatment of N. 
texensis for further information concerning 
Carrie Bow Cay and the specific habitat 
where this species occurs. 

Remarks.—This species is closely related 
and similar to congeners of the Hoguei group 
but can be distinguished from the latter by 
the following combination of characters: 
middle anaclinate genal seta about one-half 
length of seta on either side; spinelike setae 
along costal margin variable, short, not as 
long as width of Ist costal cell, or long (the 
holotype), length equal or slightly greater 
than width of Ist costal cell; general ap- 


603 


pearance setose but less so than in P. hoguei; 
surstylus of male terminalia as illustrated 
(Figs. 39-42). After study of several males 
from the new material collected in Belize, 
Cuba, Dominica, and Florida, I have noted 
some variation, although slight, in the shape 
of the surstylus. The range in variation is 
as illustrated (Figs. 39-42); the illustrations 
also facilitate identification of this species. 
Based on the sampling available, the shape 
of the ventral surstylar margin in the male 
from Florida, both lateral and posterior 
views (Figs. 41, 42), is more characteristic 
of the species than that of the holotype, 
which is from Dominica (Figs. 39, 40). 


Paracanace lebam Mathis & Wirth 
Figs. 43-44 


Paracanace lebam Mathis & Wirth, 1978: 
530. 


Distribution.—Jamaica. Point Hender- 
son. 

Remarks.— Externally, this species and P. 
aicen are very similar, but it may be distin- 
guished from the latter by the following 
characters from the male terminalia: sur- 
stylus (Figs. 41, 42) broader on distal half, 
especially evident in lateral view; ventral 
margin of surstylus broadly truncate in lat- 
eral and posterior views; posterior margin 
of surstylus bearing distinct row of longer 
setae. 


Genus Procanace Hendel 


Procanace Hendel, 1913:93. Type species: 
Procanace grisescens Hendel, by original 
designation.— Mathis, 1988:329-333 
[first record of genus from Western Hemi- 
sphere]. 


Diagnosis.—General coloration whitish 
gray, olivaceous, to blackish brown. 

Head: Intrafrontal setae absent, but with 
a few setulae inserted anteriorly; fronto-or- 
bital setae three; ocelli arranged to form 
equilateral or isosceles triangle, if isosceles, 


604 
/ 
pele 
\ 

NS a 

dy lle 

43 \ 

Figs. 43-44. 


44, Epandrium and surstyli, posterior view. 


the greater distance is between posterior 
ocelli. Arista pubescent over entire length. 
Two large anaclinate genal setae; anterocli- 
nate genal seta moderately well developed. 
Palpus not bearing long setae. Epistomal 
margin, in lateral view, more or less hori- 
zontal. 

Thorax: Acrostichal setae, especially a 
prescutellar pair of large setae, usually lack- 
ing (setulae present in species of the wil- 
liamsi group); scutellar disc lacking setae 
(one or two pairs of scutellar disc setulae 
occur in P. nakazatoi Miyagi of the william- 
si group); two pairs of marginal scutellar 
setae, apical pair not anaclinate; anterior 
and posterior notopleural setae present, 
length of both subequal, anepisternum with 
scattered setulae. Katepisternal setae usually 
present (lacking in species of the grisescens 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


External male terminalia of Paracanace lebam: 43, Epandrium and surstylus, lateral view; 


group). Hind tibia lacking spine-like setae 
apically. 

Abdomen: Male genitalia as follows: 
Epandrium in posterior view wider than 
high; cerci reduced, poorly sclerotized; sur- 
stylus with an anterior and posterior lobe, 
the latter larger, sometimes markedly so and 
shape unique to species. 

Discussion. — Mathis (1988) first reported 
the occurrence of Procanace in the Western 
Hemisphere from specimens collected in 
Virginia along the tidal shores of the Po- 
tomac River. The species, P. dianneae, was 
then only known from Virginia, but recent 
collecting in North and South Carolina, as 
well as the Gulf Coast of Florida has re- 
sulted in discovery of this species along 
much of the eastern coast of the United 
States. 


VOLUME 102, NUMBER 3 605 


Figs. 45-53. Scanning electron micrographs of Procanace dianneae: 45, Head, lateral view; 46, Gena and 
setae, lateral view; 47, Antenna, lateral view; 48, Mesonotum, dorsal view; 49, Frons and ocellar triangle, dorsal 
view; 50, Fronto-orbital and vertical setae, dorsal view; 51, Scutellum, dorsal view; 52, Notopleuron and setae, 
lateral view; 53, Katepisternum and setae, lateral view. 


Annotated Key to Species Groups of 


Palaearctic, Oceanian, Mala- 
Procanace Hendel 


gasy, Seychelles] 
1. Katepisternal seta absent ........ — Katepisternal seta present ....... 2 
RES inti ced hom Ls dthys the grisescens group 2. Clypeus high, width about twice the 
[four species; Oriental, eastern height; palpus blackish brown; 


606 


54 a7 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Figs. 54-56. External male terminalia of Procanace dianneae: 54, Epandrium and surstylus, lateral view; 
55, Surstyli, posterior view; 56, Fourth and fifth sterna, ventral view. 


proepisternal seta absent 
be See net rad nates the nigroviridis group 
[seven species; Hawaiian Is- 
lands] 

— Clypeus low, width at least four 
times the height; palpus yellowish; 
proepisternal seta(e) present ..... 3 

3. Acrostichal setulae present, in two 
irregular rows .... the williamsi group 

[four species; Hawaiian and 
Ryukyu Islands] 

— Acrostichal setulae absent 

4. Postocellar setae either absent or 
much reduced y.. 3s the fulva group 

[nine species; Oriental and east- 
ern Palaearctic] 

— Postocellar setae present, subequal 
to length of ocellar seta .......... 

ithe WSF lap Dae aie the cressoni group 
[three species; Oriental, Nearc- 
tic] 


Procanace dianneae Mathis 
Figs. 45, 46 


Procanace dianneae Mathis, 1988:330-333. 


Distribution.— Eastern coast of the United 
States from Virginia south through North 


and South Carolina to Florida, including the 
Gulf Coast. 

Natural history.— The type series was tak- 
en along the tidal shore of the Potomac Riv- 
er where the water is only slightly brackish. 
The additional distribution sites reported 
here are oceanic (Atlantic and Gulf of Mex- 
ico), with distinctly saline water. On Kure 
Beach (North Carolina, Brunswick Co.), the 
specimens were extremely abundant on the 
sandy shore within a protected area where 
boats could be launched. The exposed sand 
was largely covered with algae that had 
washed ashore. At Cherry Grove (South 
Carolina, Horry Co.), the specimens were 
found exclusively on the concrete founda- 
tions ofa fishing pier. The foundations near- 
est the shore are exposed at low tide and 
most high tides and were partially covered 
with algae. A species of Tethina (probably 
albula (Loew); Tethinidae) and Fucellia 
(Anthomyiidae) also occurred on the foun- 
dations. The causeway leading to Sanibel 
Island (Gulf Coast side of Florida, Lee Co.) 
had portions of the shore that were lined 
with large chunks of concrete and rock to 
brake the erosive action of waves, and the 
specimens P. dianneae mostly occurred 


VOLUME 102, NUMBER 3 


where the wave action was dampened by 
the rocks and concrete that were also ex- 
tensively covered by algae. Two other beach 
flies, Canacea macateei and Paracanace 
aicen, also occurred on the causeway. 

Remarks.— Externally this species is very 
similar to those of the cressoni group. It 
differs from the two species of that group, 
P. cressoni Wirth and P. taiwanensis Del- 
finado, as well as other congeners by the 
following combination of characters: Post- 
ocellar setae well developed, subequal in 
length to ocellar setae; clypeus low, height 
one-fourth width; palpus yellowish. Scutum 
mostly bluish black, sparsely microtomen- 
tose, scutum densely microtomentose, 
brown; proepisternal seta present, pale; kat- 
episternal seta present; acrostichal setae ab- 
sent. Shape of the male genitalia unique (see 
figs. and description above). 

As noted previously, this species was only 
recently discovered in Virginia, and it has 
now been found to occur along the East Coast 
of the United States from Virginia to Flor- 
ida, including the Gulf Coast. Apparently 
the species either dispersed rapidly along 
the East Coast after its introduction, per- 
haps similar to Brachydeutera longipes 
Hendel (Mathis and Steiner 1986; Ephyd- 
ridae), or it has resided here for some time 
without being detected. 


Acknowledgments 


I am grateful for the assistance in the field 
from Jorge L. Fontenla (IZAC) and the oth- 
er collectors as noted in the text. For the 
loan of the holotype of Canaceoides tex- 
ensis, | thank Paul H. Arnaud, Jr. (CAS). 
For critically reviewing a draft of this paper, 
I thank Amnon Freidberg, Oliver S. Flint, 
Jr., and I. Candida Feller. The pen and ink 
illustrations were skillfully inked by George 
L. Venable, and Susann Braden provided 
technical support for the scanning electron 
micrographs. Funding for this research 
project was provided in part by the Amer- 
ican Philosophical Society (grant number 
9284, Penrose Fund) and a grant from the 


607 


Research Opportunity Fund (Smithsonian 
Institution). Their support is gratefully ac- 
knowledged. 

This is contribution number 257, Carib- 
bean Coral Reef Ecosystems (CCRE), 
Smithsonian Institution, partly supported 
by a grant from the Exxon Corporation. 


Literature Cited 


Cresson, E. T., Jr. 1924. Descriptions of new genera 

and species of the dipterous family Ephydridae. 

Paper VI.—Entomological News 35(5):159-164. 

1936. Descriptions and notes on genera and 
species of the dipterous family Ephydridae. IT. — 
Transactions of the American Entomological 
Society 62:257-270. 

Curran, C.H. 1934. The families and genera of North 
American Diptera. Privately published, New 
York, 512 pp., 235 figs., 2 pls. 

Hendel, F. 1913. Acalyptrate Musciden (Dipt.). II.— 
Supplementa Entomologica 2:77-112, 7 figs. 

Johnson, C. W. 1910. Order Diptera. Jn J. B. Smith, 

ed., The insects of New Jersey. New Jersey State 

Museum Annual Report 1909:703-814. 

1925. 15. List of the Diptera or two-winged 
flies. Jn Fauna of New England. Occasional Pa- 
pers of the Boston Society of Natural History 7: 
1-326, 1 fig. 

Malloch, J.R. 1924. A new species of Canacea from 

the United States (Diptera: Ephydridae).—Pro- 

ceedings of the Entomological Society of Wash- 
ington 26(3):52-53. 

1933. Some Acalyptrate Diptera from the 
Marquesas Islands.— Bulletin of the Bernice P. 
Bishop Museum 114:3-31, 9 figs. 

Mathis, W. N. 1982. Studies of Canacidae (Diptera), 

I: Suprageneric revision of the family, with re- 

visions of new tribe Dynomiellini and new ge- 

nus Jsocanace.—Smithsonian Contributions to 

Zoology 347:1-29. 

1988. The first record of Procanace Hendel 
from North America, with the description of a 
new species. — Proceedings of the Entomological 
Society of Washington 90(3):329-333. 

, & W.E. Steiner. 1986. An adventive species 

of Brachydeutera Loew in North America (Dip- 

tera: Ephydridae).—Journal of the New York 

Entomological Society 94(1):56-61. 

——., & W. W. Wirth. 1978. A new genus near 
Canaceoides Cresson, three new species and 
notes on their classification (Diptera: Canaci- 
dae).— Proceedings of the Entomological Soci- 
ety of Washington 80(4):524—537, 12 figs. 

Ritzler, K., & J. D. Ferraris. 1982. Terrestrial en- 
vironment and climate, Carrie Bow Cay. Pp. 


608 


77-91 in K. Riitzler & I. G. MacIntyre, eds., 
The Atlantic barrier reef ecosystem at Carrie 
Bow Cay.—Smithsonian Contributions to the 
Marine Sciences 12:1-539. 

Stoddart, D., F. R. Fosberg, & D. L. Spellman. 1982. 
Cays of the barrier reef and lagoon.— Atoll Re- 
search Bulletin 256:1076. 

Teskey, H. J., & I. Valiela. 1977. The mature larva 
and puparium of Canace macateei (Diptera: 
Canaceidae).— The Canadian Entomologist 109: 
345-347, 7 figs. 

Tomlinson, P. B. 1986. The botany of mangroves. 
Cambridge Tropical Biology Series, Cambridge 
University Press, Cambridge, 413 pp. 

Wheeler, M. R. 1952. The dipterous family Cana- 
ceidae in the United States.—Entomological 
News 63(4):89-94. 

Wirth, W. W. 1951. A revision of the dipterous fam- 

ily Canaceidae.— Occasional Papers of Bernice 

P. Bishop Museum 29(14):245-275, 6 figs. 

1954. A new intertidal fly from California, 
with notes on the genus Nocticanace Malloch 

(Diptera: Canaceidae).—The Pan-Pacific Ento- 

mologist 30(1):59-62. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


. 1965. Family Canaceidae. Pp. 733-734 in A. 
Stone et al., eds. A catalog of the Diptera of 
America north of Mexico. United States De- 
partment of Agriculture, Agriculture Handbook 
276. Washington, D.C. 

. 1970. The American beach flies of the Cana- 
ce snodgrassii group.— Proceedings of the En- 
tomological Society of Washington 72(3):397- 
403, 4 figs. 

1975. 76. Family Canaceidae. Pp. 1-5 in N. 
Papavero, ed., A catalogue of the Diptera of the 
Americas south of the United States. Museu de 
Sao Paulo, Universidade de Sao Paulo, Brasil. 

1987. Camnacidae [chapter 102]. Pp. 1079- 
1083 in J. F. McAlpine, ed., Manual of Nearctic 
Diptera. Volume 2. Monograph 28, Research 
Branch, Agriculture Canada. Hull, Quebec. 


Department of Entomology, NHB 169, 
Smithsonian Institution, Washington, D.C. 
20560. 


PROC. BIOL. SOC. WASH. 
102(3), 1989, pp. 609-612 


A SMALL COLLECTION OF HETEROPTERA FROM THE 
GALAPAGOS ISLANDS, WITH THE DESCRIPTION OF 
THE NEW SPECIES NIESTHREA ASHLOCKI AND A 
LIST OF NIESTHREA SPECIES (RHOPALIDAE) 


Richard C. Froeschner 


Abstract.—Report on seven species of Heteroptera from the Galapagos Is- 
lands: Five confirm reported occurrences on Santa Cruz Island; one new island 
record of Harmostes disjunctus Barber from Fernandina Island; and description 
of one new species, Niesthrea ashlocki taken from Sida acuta Burmann [Mal- 
vaceae] on Santa Cruz Island, most closely allied to the Brazilian N. digna 
Chopra. Description of the new species accompanied by dorsal habitus drawing 
and sketch of male genital capsule. Included is a checklist of the species of 


Niesthrea Spinola. 


Subsequent to the appearance of my syn- 
opsis of the Heteroptera of the Galapagos 
Islands (1985) Dr. Peter D. Ashlock (Uni- 
versity of Kansas, Lawrence) submitted for 
my study a small collection of Heteroptera 
he made on the Galapagos Islands during 
the period of January to May of 1964. All 
specimens but one were from Santa Cruz 
Island and included the following: Beryti- 
dae: Metacanthus galapagoensis (Barber) [in 
abandoned garden]; Coreidae: Anasa ob- 
scura Dallas [from ‘“‘“Mamortlca” (probably 
a misspelling for the Cucurbitaceae genus 
Momordica) indica; in abandoned garden]; 
Miridae: Horcias lacteiclavus Distant [part 
of the population discussed by Carvalho 
(1968:200); Pentatomidae: Acrosternum 
viridans (Stal) [at light; in abandoned gar- 
den]; Podisus sordidus (Stal) [from Psidium 
sp.]; Rhopalidae: Harmostes disjunctus Bar- 
ber, including nymphs; Niesthrea ashlocki, 
new species described below [from Sida 
acuta Burmann]. One new island record was 
included based on a broken specimen of 
Harmostes disjunctus taken in the Miconia 
Belt at 1300-2100’ on the SW side of Fer- 
nandina Island, 4 Feb 1964, P. D. Ashlock. 


The New World genus Niesthrea Spinola 
(1837:245), in the tribe Niesthrini, was not 
previously reported for the Galapagos Is- 
lands. In my (1985) synopsis it would key 
to Liorhyssus in couplet 2 on page 49. 

Niesthrea’s pronotum lacks the subapical, 
calloused, impunctate, transverse ridge that 
is characteristic of Liorhyssus. Chopra 
(1973) presented a revision of the genus 
Niesthrea based principally on male geni- 
talic characters. 


Niesthrea ashlocki, new species 
Figs. 1-2 


Diagnosis.— Males are easily recognized 
to species by the shape of the medioventral 
lobe on the genital capsule plus the length 
of the claspers (Fig. 2): Medioventral lobe 
broad, lateral expansion subangularly con- 
vex; exposed part of clasper gently curved, 
apically more strongly incurved to a sub- 
acute tip, exposed part of clasper elongate, 
extending by half its length beyond apex of 
medioventral lobe. 

Description (measurements in millime- 
ters).—Holotype male, length 5.35; general 


610 


2 


Figs. 1-2. Niesthrea ashlocki, new species: 1, dorsal 
view; natural length 5.3 mm; 2, male genital capsule, 
ventral view, left clasper omitted. 


color yellowish; head clouded with fuscous 
on midline between eyes, with a pair of di- 
verging, deep-black lines between ocelli. 
Antenna yellow, segment I mesally and lat- 
erally with an oblique, fuscous line; seg- 
ments II and III with a blackened line ex- 
tending almost full length, II blackened 
apically. Pronotum with median fuscous 
area divided by pale median carina; pos- 
terior lobe with humeri, subbasal margin, 
and diagonal row of a few fuscous spots. 
Scutellum with subbasal pair of brown spots. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


or 
S f 
cA 
Vi S 
i i V3) 
Y 
: 


Hemelytral veins with a few reddish-brown 
dots. Dorsal disc of abdomen (viewed 
through hyaline hemelytra) mostly black; 
connexival segments apically pale, visible 
segments III-—V basally broadly black, each 
with an included pale dot, the black ex- 
tending onto margin of venter. Legs yellow, 
appearing annulate due to transverse dark 
marking extending more or less around fe- 
mur and tibia. 

Head.—Length 0.90, width across eyes 
1.15; preocular part convex, tylus distinctly 
produced anteriorly; antennal tubercles 
short, apex transverse. Antennal, segment 
lengths I-IV, 0.36:0.87:0.87:0.95, segment 
I reaching apex of tylus. Labium reaching 
basal segment of abdomen, lengths of seg- 
ments J-IV, 0.53:0.71:0.53:0.76, segment I 
reaching hind margin of eye. 

Pronotum, length 0.90, width 1.77. 

Genital capsule (Fig. 2) with medioven- 
tral lobe broader than long, constricted ba- 
sally, apical margin subangularly concave, 
laterally subangularly convex; dorsolateral 
lobe (best viewed from above) distinctly in- 
curved, markedly surpassing apex of medio- 
ventral lobe. Exposed part of clasper gently 
curved, apex projecting mesally as an acute 
tip, inner margin near apex of capsule with 
a small tooth. 

Female.—Length 5.58. General appear- 
ance similar to male but more abundantly 
dotted on pronotum; ventrally with red dots 
on thoracic pleurae. Last abdominal tergum 
rounded. Last abdominal sternum not 
notched. Head, length 1.15, width 1.23. An- 
tennal segment lengths, I-IV 0.37:1.50:1.50: 
1.20. Labial segment lengths, I-IV, 0.57: 
0.76:0.62:0.82. Pronotum, length 1.25, 
width 2.06. 

Holotype male.—“‘Galapagos, 4 mi. [6.4 
km] N Academy Bay, Santa Cruz Is., 21 Feb 
1965, P. D. Ashlock, Sida acuta Burm. 
[Malvaceae].’’ Paratype: one female, Gala- 
pagos Arch.[ipelago], Santa Cruz Is., 2.4 km 
N Academy Bay, 25 Feb 1964, P. D. Ash- 
lock. Holotype and lone paratype in the Na- 
tional Museum of Natural History. 

Placement of this new species in the phy- 


VOLUME 102, NUMBER 3 


logeny (based solely on males) proposed by 
Chopra (1973) in his revision of the genus 
Niesthrea may be made as follows. Three 
modifications of the male genital structures 
(Fig. 2) place it on the same branch of Cho- 
pra’s (1973:457) “‘Phylogenetic tree’? with 
N. digna Chopra: genital capsule with dor- 
solateral lobes incurved apically, medio- 
ventral lobe broader than long, and clasper 
apically incurved to form an acute angle. It 
differs from N. digna in having both the 
dorsolateral lobes and the clasper greatly 
surpassing the apex of the medioventral lobe, 
the claspers by almost half their own length 
and the dorsolateral lobe extending beyond 
them. The Galapagos Islands occurrence of 
N. ashlocki is geographically remote from 
the Brazilian homeland of N. digna. 

The species name is a dedication to Dr. 
Peter D. Ashlock, the collector whose many 
contributions to heteropterology have 
soundly advanced that science. 


Checklist of the Species of Niesthrea 
Spinola 


The following list is an expanded version 
of that given on pages 52-56 of Gollner- 
Scheiding’s (1983) catalog of the family 
Rhopalidae. 


agnes Chopra, 1973:455 Argentina 
ashlocki, new species Galapagos Islands 
brevicauda Chopra, 1973:455 Peru 
dentata Chopra, 1973:454 Brazil 
digna Chopra, 1973:453 Brazil 
fenestrata (Signoret), 1859:93 Chile 
flava Grillo & Alayo, 1978:43 Cuba 
louisianica Sailer, 1961:297 U.S.A.; 
Mexico 
parasidae Grillo & Alayo, Cuba 
1978:46 
pictipes (Stal), 1859:239 Argentina; 
Brazil; 
Paraguay 
subsp. pictipes (Stal), see 
species entry 
subsp. casinii Gdllner- 
Scheiding, 1984:116 Argentina; 
Uruguay 
sidae (Fabricius), 1794:169 Greater and 
Lesser 
Antilles; 


611 


Colombia; 
Mexico; 
United States; 
Venezuela 
Argentina; 
Brazil 
Guatemala; 
Mexico; 
United States 
Greater and 
Lesser 
Antilles; 
Argentina; 
Brazil; 
Paraguay; 
Venezuela 


similis Chopra, 1973:453 


ventralis (Signoret), 1859:89 


vincentii (Westwood), 1842:6 
and 26 


Acknowledgments 


My appreciation is expressed to Dr. John 
J. Wurdack, Smithsonian Institution De- 
partment of Botany, for his help in deci- 
phering the misspelled host genus for Anasa 
obscura; to Ms. Silver B. West for help in 
preparing the manuscript; to Ms. Elsie 
Froeschner for the drawings, and to Dr. 
Oliver S. Flint and Mr. Thomas J. Henry 
for helpful reviews of the manuscript. 


Literature Cited 


Carvalho, J. C. M., & W. C. Gagne. 1986. Miridae 
of the Galapagos Islands (Heteroptera).—Pro- 
ceedings of the California Academy of Sciences, 
Series 4 36(7):147-219. 

Chopra, N. P. 1973. Arevision of the genus Niesthrea 
Spinola (Rhopalidae: Hemiptera).—Journal of 
Natural History 7:441-459. 

Fabricius, J.C. 1794. Ryngota. Jn Entomologia sys- 
tematica emendata et aucta, secundum classes, 
ordines, genera, species, adjectus synonymis, lo- 
cis, observationibus, descriptionibus 4:[I-IV], 
1-229. 

Froeschner, R. C. 1985. Synopsis of the Heteroptera 
or true bugs of the Galapagos Islands. —Smith- 
sonian Contributions to Zoology 407:1-84. 

Gollner-Scheiding, U. 1983. General-Katalog der 

Familie Rhopalidae (Heteroptera). — Mitteilun- 

gen aus dem Zoologischen Museum in Berlin 

59:37-189. 

1984. Erganzungen zu den Gattungen Lio- 
rhyssus Stal, 1870, Niesthrea Spinola, 1837, und 
Rhopalus Schilling, 1827 (Heteroptera, Rho- 
palidae).— Mitteilungen aus dem Zoologischen 
Museum in Berlin 60:115-121. 


612 


Grillo R., H. & P. Alayo. 1978. La Familia Rho- 
palidae (Heteroptera:Coreidae) en Cuba. Centro 
Agricola. Facultad de Ciencias Agricoles, Uni- 
versidad Central de Las Villa, Septiembre-Di- 
ciembre 1978:41-64. 

Sailer, R. I. 1961. The identity of Lygaeus sidae Fa- 
bricius, type species of the genus Niesthrea (He- 
miptera: Coreidae).— Proceedings of the Ento- 
mological Society of Washington 63:293-299. 

Signoret, V. 1859. Monographie du genre Corizus.— 
Annales de la Societe Entomologique de France. 
Series 3 7:75-105. 

Spinola, M. 1837. Essai sur les genres d’insects ap- 
partenants a l’ordre des Hemipteres, Lin. ou 
Rhynchotes, Fab., et a la section des Heterop- 


Note: The honoree of the new species, Dr. Peter D. 
Ashlock, died 26 January 1989. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


teres, Dufour. Geneva, Yves Graviers. 383 pp., 
15 tabs. 

Stal, C. 1859. Hemiptera: Species novas descripsit. 
Kongliga Svenska Fregattens Eugenies Resa 
Omkring Jorden, III (Zoologi, Insekter). Pp. 219- 
298, pls. 3-4. 

Westwood, J.O. 1842. A catalogue of Hemiptera in 
the collection of the Rev. F. W. Hope, with short 
Latin descriptions of the new species. 2:1—26. 


Department of Entomology, NHB Stop 
127, National Museum of Natural History, 
Washington, D.C. 20560 


PROC. BIOL. SOC. WASH. 
102(3), 1989, pp. 613-619 


CANCER JOHNGARTHI, N. SP. AND 
CANCER PORTERI (BELL) 
(CRUSTACEA, DECAPODA): 
COMPARISONS AND HYPOTHESIS 


Alberto Carvacho 


Abstract.— Cancer johngarthi, long mistaken for C. porteri, is described. It 
is known in the eastern Pacific from Isla Guadalupe, Mexico (29°N), south to 
Panama (7°N), on soft bottoms at depths exceeding 90 m. Differences from C. 
porteri are discussed with an emphasis on biological aspects: C. johngarthi 
shows an isometric growth of chelipeds in relation to carapace width, while in 
C. porteri a strong positive allometry is evident after the puberal molt. These 
two conditions may reflect different mating systems. 


The species Cancer porteri, described by 
Bell (1835) as C. longipes, was transferred 
to the genus Platycarcinus, synonym of 
Cancer, by H. Milne-Edwards & Lucas 
(1844), and given its present name by Rath- 
bun (1930). Nations (1975) included it in 
the subgenus Cancer s.s. The holotype was 
collected in Valparaiso, Chile. Faxon (1895) 
recorded it from Panama Bay as deep as 
523 m (Albatross). These two eastern Pacific 
localities, 33°S and 7°30’N, respectively, 
were long considered as the geographic dis- 
tributional limits of the species. 

Garth (1957) cited a continuous distri- 
bution “from Callao, Peru to Valparaiso, 
Chile, 0-24 fms” and an extralimital record 
from Panama. Nevertheless, he also includ- 
ed in the list of examined material one male 
collected by the Lund University Chile Ex- 
pedition at Talcahuano (36°41'S), some 450 
km south of Valparaiso. This latter record 
has been confirmed by Retamal & Yanez 
(1973). Garth (1961) recorded C. porteri 
from the coast of Sinaloa in the Gulf of 
California, between 108 and 128 m, and 
mentioned that the species ““may now be 
reported as a bi-temperate species that 
transgresses the tropics by submergence, 
being found in the Gulf of California, the 


Bay of Panama in 210 to 286 fathoms, and 
from Peru to Chile in the Sublittoral.”’ 

Information gathered during almost 150 
years supported the idea that C. porteri was 
a eurybathic species with a wide geographic 
distribution. In fact, the case has been used 
as a paradigm of the peculiar tropical sub- 
mergence distribution pattern (Ekman 1953, 
Garth 1961). 

Careful study of several specimens re- 
cently collected off Baja California Sur and 
the reexamination of virtually all specimens 
identified with Cancer porteri from the 
Northern Hemisphere, along with several 
specimens from Chile and Peru, leads to the 
conclusion that they belong to two different 
species. The morphological differences, 
scarcely evident in young specimens, may 
express divergence in their mating systems. 


Cancer johngarthi, new species 
Figs. 1, 3A, 4B 


Cancer longipes, Faxon, 1895:16; Rathbun, 
1930:199 (in part). 

Cancer porteri Rathbun, 1930:199 (in part); 
Garth, 1957:50 (in part); 1961:122; Par- 
ker, 1964:173; Chirichigno, 1970:45 (in 
part); Retamal & Yanez, 1973:12 (in part); 


614 


Nations, 1975:43 (in part); 1979:154, 156, 
178 (in part); Retamal, 1981:30 (in part). 


Carapace granulated, widely oval, very 
convex and moderately areolated, with pro- 
tuberances on proto- and mesogastric re- 
gions and on borders of epi- and meso- 
branchial regions. Both mesobranchial 
regions swollen and nearly meeting in me- 
dian line. Frontal region convex. Front pro- 
jected and furnished with 3 teeth, median 
slightly longer and narrower than others. In- 
ner orbital tooth pointed and slightly short- 
er than frontal teeth. Anterolateral margin 
finely granulated, cut into 9 teeth; granu- 
lations enhanced posteriorly. Posterolateral 
margin granulated and furnished with 2 
teeth, first small and second vestigial, some- 
times imperceptible. Pterygostomial region 
swollen and coarsely granulated towards 
outer edge. Whole carapace remarkably thin; 
epi- and subbranchial and pterygostomial 
regions may be easily flexed. 

Buccal cavity well delimited anteriorly by 
projections of pterygostomial border, with 
2 strong vaults separated by a longitudinal 
keel. 

Maxillipeds granulated, with ischium and 
merus widened distally. Merus with outer 
face concave and a notch on distal half of 
inner margin where palp inserts. 

Chelipeds: fingers with tips and cutting 
edges dark, starting from proximal tooth. 
Palm granulated, with 4 longitudinal cari- 
nae on lower half of outer face. Propodus 
2.7 times as long as wide in adult males. 
Carpus rough, with irregular granulated ca- 
rinae and anterosuperior pyramidal tooth. 
Merus with subtriangular section and upper 
distal margin granulated. 

Walking legs long and slender, without 
spines or setae on proximal articles. Prop- 
odus with scarce setae on distal end of lower 
margin. Dactylus with 4 symmetrical lon- 
gitudinal rows of setae and a deep groove 
along inner and outer faces, respectively. 

Abdomen in adult males with terminal 
segment narrowly rounded distally, lateral 
margins slightly concave and 1.1 times as 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


long as wide. Adult males with second pleo- 
pod slightly overreaching second segment 
of thorax; first pleopod slightly overreach- 
ing third segment of thorax. 

Holotype.—In the collection of the Allan 
Hancock Foundation: male (carapace 14 cm 
wide, 8.7 cm long); Isla Guadalupe, Mexico; 
183 m; 15 Nov 1968, Velero IV sta. 12460- 
68; catalog number AHF 6815. 

Material examined.—Paratypes: Isla 
Guadalupe, Mexico (29°N); 183 m, 15 Nov 
1968, Velero IV sta. 12460-68; 4 males, 1 
female, AHF 6816. Off Rio San Lorenzo, 
Sinaloa, Mexico (24°15'N), 108-128 m; May 


1959; 2 males, 6 females; AHF 5929. Off 


Bahia Magdalena, B.C., Mexico (24°15'N), 
90-125 m; Jul 1987, 1 male, 3 females, CIB, 
La Paz. Bahia de Panama; 384 m; Mar 1891; 
Albatross sta. 3389; 1 female, MCZ, Har- 
vard University. 

Distribution.—Eastern Pacific from Isla 
Guadalupe, Mexico to Bahia de Panama. 
Southern Gulf of California. 

Habitat.—Soft bottoms, 90-523 m. 

Etymology.— Named in honor of Dr. John 
S. Garth, Chief Curator Emeritus, Allan 
Hancock Foundation, University of South- 
ern California, Los Angeles, California. 


Comparison with Cancer porteri 


The observations listed below and also 
data for Figs. 3 and 4 resulted from the ex- 
amination of 17 specimens of C. johngarthi 
(carapace widths from 35 to 140 mm) and 
43 specimens of C. porteri (c.w. 22.8 to 123 
mm). These latter came from Valparaiso, 
Chile and from the following localities in 
Peru: Bahia Independencia, Bahia San Juan, 
Isla San Lorenzo, Bahia San Nicolas, Ca- 
llao, Isla Lobos de Afuera. 

1. Chelipeds of adult males noticeably 
stronger in C. porteri (Fig. 2a). As shown in 
Fig. 3B this allometric character is better 
expressed after the molt of puberty. 

2. Darkening in cutting edges of cheliped 
fingers starts proximally in C. porteri but in 
C. johngarthi it starts at first tooth. 

3. The most remarkable difference at any 


VOLUME 102, NUMBER 3 615 


= 
negra 


pa i ; 


pert haere 


i 


\ 


sre 


iyfig. LCR ose 
SOOT 


L 
¢ “is 
GH 


Fig. 1. Cancer johngarthi, male: a, Carapace, dorsal; b, Abdomen; c, Left cheliped; d, Second pleopod, distal 
end; e, First and second pleopods; f, First pleopod, distal end; g, Third maxilliped. 


616 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 2. Cancer porteri, male: a, Left cheliped; b, First pleopod, distal end; c, Abdomen. 


age is consistency of the carapace. C. john- 
garthi seems to be a paper shell species. 

4. Terminal segment in abdomen of males 
with straight lateral margins in C. porteri 
and slightly concave margins in C. john- 
garthi (Figs. 1b and 2c). 

5. Pereopods proportionately longer and 
dactyli of walking legs longer in relation to 
propodi in C. johngarthi. 

6. Width/length relation of carapace sig- 
nificantly greater in C. johngarthi (Fig. 4). 

7. As shown in Figs. 1f and 2b, apex of 
first male pleopods quite different in each 
species. 

Habitat preferences of each species are 
not sufficiently documented, but C. john- 
garthi seems to prefer deeper waters; it has 
not been collected shallower than 90 m while 
C. porteri inhabits waters from the intertidal 
to more than 350 m (Yanez 1974). This 
scheme agrees with the well known relation 
of Cancer and water temperature: species in 
this genus usually live at latitudes greater 
than those where the surface isotherm of 
20°C is to be found (MacKay 1943, Nations 
1979). At lower latitudes depth compen- 
sates thermal needs; such is the case of C. 
borealis and C. irroratus in Florida, and es- 
pecially C. guezei in Madagascar (Crosnier 
1976). 


The Hypothesis 


The genus Cancer originated in the north- 
eastern Pacific (Ekman 1953, Nations 1975, 
1979) and dispersed southward along the 
west coast of the Americas. Four species 
may be found in Peru and Chile; one of 
these is C. porteri. This species, now sepa- 
rated from C. johngarthi, ranges from Isla 
Lobos de Afuera (6°57’S) to Talcahuano 
(36°41’S), covering most of the Peruvian- 
Chilean province (sensu Briggs 1974). The 
morphological afiinity and geographic dis- 
tribution of the two species suggest the ex- 
istence of a common ancestor that trav- 
eled—perhaps in the Miocene—between 
North and South America (Nations 1979). 

Morphological divergence between C. 
johngarthi and C. porteri may have resulted 
from different mating systems. Orensanz & 
Gallucci (1988) explain some differences 
among four sympatric species of Cancer, 
such as dimorphic development of che- 
lipeds, according to the models of polygyny 
established by Emlen & Oring (1977). Mat- 
ing systems of species with precocious de- 
velopment of a strong cheliped may be in- 
terpreted as a case of resource defense 
polygyny: in C. oregonensis each male holds 
a refuge area—limited resource—which al- 


VOLUME 102, NUMBER 3 


Cheliped height (cm) 


Carapace width (cm) 


Cheliped height (cm) 


2 4 6 8 10 - 12 14 


Carapace width (cm) 
Fig. 3. Cheliped height plotted against carapace 
width in males: A, Cancer johngarthi; B, Cancer por- 
teri. 


lows him to monopolize females. Preco- 
cious development of strong chelipeds is re- 
quired for an early appropriation of adequate 
refuges. The case of harem defense polygyny 
involves direct access to females; therefore, 
defense strategies are only needed once re- 
productive size is reached. Positive allo- 
metric growth of chelipeds, consequently, 
starts just after the molt of puberty. This 
seems to be the case for C. porteri, as in- 
dicated by allometric growth of male che- 
lipeds (Fig. 3B) and also by data in Antezana 


617 


y 


L57 


l 


Fig. 4. Frequencies of width to length ratios in car- 
apaces: A, C. porteri; B, C. johngarthi. 


L; 


54 
1.56 


Uh 


UY, 


1.5I 
1.53 


SS 


1.45 
1.47 


1.48 
L50 


160 
162 


W/L RATIO 


et al. (1965) who determined a figure of 5 
females per each male after one year of 
monthly sampling. 

In the third model, male dominance 
polygyny, mates or critical resources are not 
economically monopolizable. Males aggre- 
gate during the breeding season and females 
select males from these aggregations. Sexual 
dimorphism in the development of che- 
lipeds is not expected here. Orensanz & Gal- 
lucci (1988) included C. magister in this 
category, stressing the fact that C. magister 
is the only species in the genus Cancer in 
which chelipeds are of the same size in males 
and females. I suggest that this is also the 
case for C. johngarthi, in absence of di- 
morphic development of chelipeds. Several 
additional arguments uphold this hypoth- 
esis. As expected, sex ratio is almost 1:1 and 
sexual selection nearly null; from a total of 


618 


27 known specimens of C. johngarthi, 15 
are males and the rest females. Since breed- 
ing assemblages are not permanent, breed- 
ing season should be normally restricted to 
a short period of time; this seems a reason- 
able explanation for the lack of ovigerous 
females in the collected material. 

On the other hand, there is a relation be- 
tween size of cheliped and quality of sub- 
strate. Species inhabiting soft and homog- 
enous bottoms of fine sand have chelipeds 
proportionately weaker than those from ir- 
regular rocky substrates. Lawton & Elner 
(1985) stated that these differences account 
basically for the type of feeding, but they 
left aside an important element of analysis, 
the role of chelipeds in sexual selection, 
which is common to most decapod Crus- 
tacea. Evidently, differences in size and 
shape of chelipeds of dimorphic species are 
not due to differences in diet of each sex; 
male chelipeds fulfill other functions such 
as the defense of a territory or of a harem. 
Sudden development of cheliped dimor- 
phism at molt of puberty in many species 
is strong evidence of cheliped morphology 
depending primarily upon sex require- 
ments. Open soft-bottom environments, 
compared to complex rocky substrates, al- 
low fewer possibilities of delimitation and 
defense of a territory; therefore, different 
mating systems are involved. Following this 
idea, lack of dimorphism in chelipeds of C. 
Johngarthi may be interpreted as a conse- 
quence of a male dominance polygyny mat- 
ing system that in turn results from inhab- 
iting open soft bottoms. 


Acknowledgments 


This paper has been enriched with critical 
comments by John Garth, Janet Haig, Ray- 
mond Manning, José Orensanz and Rubén 
Rios. I also thank Dr. Mario Monteforte 
(Centro de Investigaciones Bioldgicas, La 
Paz, Baja California Sur) and Ardis B. John- 
ston (Department of Invertebrates at the 
Museum of Comparative Zoology, Harvard 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


University), who kindly loaned specimens. 
Illustrations were prepared by Clara Yanez 
and Carlos Sepulveda. 


Literature Cited 


Antezana, T., E. Fagetti, & M. T. Lopez. 1965. Ob- 
servaciones bioecologicas en Decapodos co- 
munes en Valparaiso. — Revista de Biologia Ma- 
rina (Chile) 12:1-60. 

Bell, T. 1835. Observations on the genus Cancer of 
Dr. Leach (Platycarcinus Latreille) with descrip- 
tions of three new species. — Proceedings of the 
Zoological Society, London 3:86-88. 

Briggs, J.C. 1974. Marine zoogeography. New York, 
McGraw-Hill, 475 pp. 

Chirichigno, N. F. 1970. Lista de Crustaceos del Peru 
(Decapoda, Stomatopoda) con datos sobre su 
distribucion geografica.—Informe del Instituto 
del Mar, Pert 35:1-115. 

Crosnier, A. 1976. Donnés sur les Crustacés Déca- 
podes capturés par M. Paul Gueze a Vile de la 
Réunion lors d’essais de péche en eau pro- 
fonde.— Travaux et Documents, ORSTOM 47: 
225-256. 

Ekman, S. 1953. Zoogeography of the sea. London, 
Sidgwick & Jackson, 417 pp. 

Emlen, S. T., & L. W. Oring. 1977. Ecology, sexual 
selection and the evolution of mating systems. — 
Science 197:215-223. 

Faxon, W. 1895. The stalk-eyed Crustacea: Report 
on an exploration off the west coast of Mexico, 
Central and South America, and off the Gala- 
pagos Islands, in charge of Alexander Agassiz 
by the U.S. Fish Commission Steamer ALBA- 
TROSS during 1891. XV.—Memoirs of the 
Museum of Comparative Zoology, Harvard 18: 
1-212. 

Garth, J. S. 1957. Reports of the Lund University 

Chile Expedition 1948-49, No. 29. The Crus- 

tacea Decapoda Brachyura of Chile.—Lund 

University Arsskrifter, (2)53:1-128. 

1961. Distribution and affinities of the 
brachyuran Crustacea.—Systematic Zoology 9: 
105-123. 

Lawton, P., & R. W. Elner. 1985. Feeding in relation 
to morphometrics within the genus Cancer: 
Evolutionary and ecological considerations. Pp. 
357-379 in B. R. Melteff, ed., Proceedings of 
the Symposium on Dungeness Crab Biology and 
Management. — University of Alaska, Alaska Sea 
Grant Reports. 

MacKay, D. C. G. 1943. Temperature and world 
distribution of crabs of the genus Cancer.— 
Ecology, 24:113-115. 

Milne-Edwards, H., et H. Lucas. 1844. Crustacés. Pp. 


VOLUME 102, NUMBER 3 


1-37 in A. D’Orbigny, ed., Voyage dans l’Amé- 
rique Meridionale. 

Nations, J. D. 1975. The genus Cancer (Crustacea: 

Brachyura): Systematics, biogeography and fos- 

sil record.—Scientific Bulletin of the Natural 

History Museum, Los Angeles County 23:1-104. 

1979. The genus Cancer and its distribution 
in time and space.—Bulletin of the Biological 

Society of Washington 3:153-187. 

Orensanz, J. M., & V. F. Gallucci. 1988. A compar- 
ative study of postlarval life-history schedules 
in four sympatric Cancer species (Decapoda: 
Brachyura: Cancridae).— Journal of Crustacean 
Biology 8:187-—220. 

Parker, R. H. 1964. Zoogeography and ecology of 
some macroinvertebrates, particularly mol- 
lusks, in the Gulf of California and the Conti- 
nental Slope of Mexico. — Videnskabelige Med- 
delelser fra Dansk Naturhistorisk Forening 126: 
1-178. 

Rathbun, M. J. 1910. The stalk-eyed Crustacea of 
Peru and the adjacent coast.— Proceeding of the 
United States National Museum 38:531-620. 


619 


. 1930. The cancroid crabs of America.—Bul- 
letin of the United States National Museum 152: 
1-609. 

Retamal, M.A. 1981. Catalogo ilustrado de los Crus- 
taceos Decapodos de Chile.—Gayana (Zoolo- 
gia) 44:7-110. 

,& L. A. Yafiez. 1973. Analisis cuali y cuan- 
titativo de los decapodos de los fondos subli- 
torales blandos de la Bahia de Concepcion, 
Chile. —Gayana (Zoologia) 23:1-47. 

Yanez, E. 1974. Distribucion y abundancia relativa 
estacional de los recursos disponibles a un arte 
de arrastre camaronero frente a la costa de Val- 
paraiso (Invierno y Primavera 1972).—Inves- 
tigaciones Marinas (Chile) 5:125-138. 


Centro de Investigacion Cientifica y de 
Educacion Superior de Ensenada, Apartado 
Postal 2732, Ensenada, B.C., México. Pres- 
ent address, Instituto Professional de Osor- 
no, Casilla 933, Osorno, Chile. 


PROC. BIOL. SOC. WASH. 
102(3), 1989, pp. 620-636 


STENORHYNCHUS YANGI, A NEW WESTERN 
ATLANTIC SPECIES OF ARROW CRAB 
(CRUSTACEA, BRACHYURA, MAJIDAE) AND A 
REDESCRIPTION OF S. SETICORNIS 
(HERBST, 1788) 


Gary D. Goeke 


Abstract.—The common arrow crab of the western Atlantic has a complicated 
taxonomic history as two species have long been confused under the name 
Stenorhynchus seticornis (Herbst, 1788). Through study of living specimens 
and preserved material in museums of North and South America, as well as 
Europe, the genus Stenorhynchus Lamarck, 1818, is reviewed, S. seticornis is 
restricted, its synonyms discussed, and a previously undescribed species, S. 
yangl, is recognized. Adults of these two species are described and illustrated, 
their morphologic variations analyzed, colors differentiated, known geographic 
and bathymetric ranges recorded, and their larvae compared. The species differ 
in rostral setation, shape of male first pleopod, and in other characters. More- 
over, two ecophenotypes related to substrate can be recognized within S. se- 


ticornis. 


The common shallow-water arrow crab, 
Stenorhynchus seticornis (Herbst, 1788), of 
the western North Atlantic is such an ob- 
vious component of the marine fauna that 
it has been described under several different 
scientific names over the past 200 years. It 
has become evident through the work of 
Yang (1967, 1976) on larval development 
in majids and on taxonomic problems in 
the genus Stenorhynchus Lamarck, 1818, 
that two species are confused under the name 
Stenorhynchus seticornis (Herbst, 1788) (Fig. 


1). 


Historical Review 


The group of spider crabs currently as- 
signed the generic name Stenorhynchus has 
a wide distribution in warm and temperate 
waters of the Atlantic and eastern Pacific 
oceans. Species now grouped within this ge- 
nus were previously assigned to various 
genera until taxonomic consistency was 


reached with the use of the generic name 
Leptopodia Leach, 1814. Arrow crabs are 
very common and led Milne-Edwards (1875: 
173) to state, ““Cette espéce est si bien con- 
nue, et elle a été si souvent figurée, qu’il est 
inutile d’en donner ici une description.” 
Leptopodia is now known to be a junior 
synonym of Inachus Weber, 1795, and not 
a valid generic name for this group. Cancer 
sagittarius Fabricius, 1793 (=Stenorhyn- 
chus seticornis) was transferred to the genus 
Leptopodia by Leach in 1815 and was con- 
sidered the type for the genus. However, 
Leptopodia was erected for Cancer pha- 
langium Pennant, 1777, and Leptopodia 
tenuirostris Leach, 1814. Because Cancer 
sagittarius was not mentioned in the origi- 
nal description of Leptopodia, it could not 
serve as the type species. Lamarck (1818) 
erected the genus Stenorhynchus for Cancer 
seticornis Herbst, 1788, and Cancer pha- 
langium. The latter species, however, is a 
member of the genus /nachus. Since the des- 


VOLUME 102, NUMBER 3 621 


ir .43 
. bd 
5 
J 
re 


- 
TAN 


Ly, AUMLMES MAU 


Fig. 1. Stenorhynchus seticornis, a, b, d, e, f, and g; Stenorhynchus yangi, c. 


622 


ignation of the name Stenorhynchus by 
Rathbun (1897) as an available name for 
the group, it has been widely accepted. Garth 
& Holthuis (1963) petitioned the Interna- 
tional Commission on Zoological Nomen- 
clature (ICZN) to designate Cancer seticor- 
nis Herbst, 1788, the type species of the 
genus and to officially emend the spelling of 
the generic name from Stenorynchus to 
Stenorhynchus. These recommendations 
were followed in opinion 763 of the ICZN. 

Five binomials may bear on western At- 
lantic species of Stenorhynchus, the earliest 
of which is the description of the “‘Oost- 
Indische Zee-Krabbe” by Slabber (1778). 
This “East Indies Sea Crab’ was described 
in very general terms which dealt primarily 
with the gross morphology of the carapace 
and legs. The specimen, a female from the 
description of the abdomen, was character- 
ized as having small setae on the sides of 
the rostrum. The very generalized illustra- 
tion accompanying Slabber’s (1778: pl. 18, 
fig. 2) description includes a single useful 
morphologic feature for specific taxonomic 
purposes, and even that is of limited diag- 
nostic value for taxa within this group. The 
figure indicates a rostrum twice the length 
of the postorbital region of the body. How- 
ever, the drawing of the original figure is 
questionable because the crab is not accu- 
rately depicted, as evidenced by the lack of 
spination on the ambulatory legs. Although 
no doubt exists as to the genus with which 
Slabber dealt, the morphological characters 
considered useful for specific identifications 
by today’s standards were not detailed by 
that author. 

The type locality of the “East Indies Sea 
Crab” as given by Slabber is incorrect. Hol- 
thuis (1959:185) noted that the genus Ste- 
norhynchus is not represented in the Indo- 
West Pacific and that material from which 
the description was drawn was apparently 
mislabeled. Holthuis (1959) restricted the 
type locality to Guadeloupe because mate- 
rial examined by Herbst, following Slab- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


ber’s description, was collected from Gua- 
deloupe. 

The binomial nomenclature introduced 
by Linnaeus (1758) was used by Herbst 
(1788:229) to designate the species Cancer 
seticornis. His abbreviated description was 
based on the work and material of Slabber 
and even reproduced the figure used by 
Slabber (Herbst, 1788: tabl. XVI, fig. 91). 
As noted by Yang (1967:211), this figure 
was also reproduced by Bosc (1802). A more 
detailed and accurate representation of a 
male Stenorhynchus species was later given 
by Herbst (1803: tabl. LV, fig. 2) from 
Guadeloupe material. This latter figure 
shows the rostrum nearly twice the length 
of the postorbital carapace. The margins of 
the rostrum are nearly parallel, and the rel- 
ative length of the dactyl of the cheliped 
suggests that the specimen figured repre- 
sents S. seticornis. However, the very reg- 
ular placement of spines and absence of 
postocular spines indicate that the figure is 
somewhat stylized and not an accurate rep- 
resentation. 

The type material for S. seticornis has not 
been located and must be assumed lost or 
destroyed. Slabber’s original material was 
offered for sale to the ““Zeeuwsch Genoot- 
schap van Wetenschappen”’ (Society of Sci- 
ences of the Province of Zeeland) in Mid- 
delburg, the Netherlands. The price asked 
was not agreeable and the offer was de- 
clined. The collection later reputedly was 
sold to the Leiden Museum. No record of 
this acquisition exists and no material is 
present which can be attributed to Slabber’s 
“Sea Crab.’ The type specimens must be 
assumed lost (L. B. Holthuis, pers. comm.). 

Because the original type locality is in error 
and the original description lacks diagnostic 
features, a neotype is proposed for the re- 
description of S. seticornis. The specimen 
on which the redescription is based was col- 
lected from the Dutch West Indies, Curacao 
(USNM 42956). The previously designated 
type locality of Guadeloupe (Holthuis 1959) 


VOLUME 102, NUMBER 3 623 


ee en A IT IIL UA UU DN a UMA AU au a aU 
a ee ee ee es 


baa HK KK KK KL KK 
Soe iene ae ee Poe OO... HB 


Fig. 2. Cancer sagittarius Fabricius, 1793: Above, ““Type” specimen from the Copenhagen Museum: Below, 
the specimen originally deposited in the Kiel Museum. 


624 


is superseded by Curacao with the selection 
of the male neotype. Curacao is a location 
from which collectors are known to have 
sent Slabber material for examination (Hol- 
thuis, pers. comm.). 

The second name to be considered is 
Cancer sagittarius (Fabricius, 1793). The 
original description of Cancer sagittarius 1s 
inconclusive. However, from an examina- 
tion of the syntypes, it has been confirmed 
the Fabrician material is conspecific with 
Stenoyrhynchus seticornis. A syntype was 
originally deposited in the Zoological Mu- 
seum, Copenhagen, and a second syntype, 
along with the remainder of the Fabrician 
material, has been transferred from the Kiel 
Museum to Copenhagen. The syntype of 
Cancer sagittarius transferred from the Kiel 
Museum has deteriorated to fragments that 
are very nearly unrecognizable. This spec- 
imen (Fig. 2) was listed by Rathbun (19235: 
14) as type material. However, it was not 
listed by Zimsen (1964) as a syntype. Prof. 
Torben Wolff (Zoological Museum, Copen- 
hagen) knows no reason why one specimen 
was listed as the type by Zimsen and not 
both (pers. comm.). This female syntype 
possesses all the characters listed below as 
typical of S. seticornis. 

The eastern Atlantic species of Steno- 
rhynchus was considered conspecific with 
the western Atlantic species until Yang 
(1967, 1976) showed it to be distinct. Apart 
from the unavailable Leptopodia vittata 
Kingsley, two names have been applied to 
the eastern Atlantic form, Leptopodia lan- 
ceolata Brullé, 1837, and L. canariensis 
Brullé, 1839, both described from the Ca- 
nary Islands. As Yang pointed out, the cor- 
rect name for the eastern Atlantic taxon is 
Stenorhynchus lanceolatus and L. canarien- 
sis must be considered a junior synonym. 
Yang (1967) showed how S. lanceolatus dif- 
fers from both S. seticornis and the pro- 
posed new species and Manning & Holthuis 
(1981) presented considerable data on the 
biology and distribution of S. /anceolatus. 
Paula (1987) has most recently described 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


the first zoeal stage of S. lanceolatus and 
compared it with the works of Yang (1967, 
1976). 

The next name possibly available is Lep- 
topodia ornata Guilding, 1825, described 
from St. Thomas in the Caribbean Sea. The 
original description in Latin is very brief. It 
treats a majid crab with a serrate rostrum, 
and includes few details as to general shape 
of the legs or other features. Guilding also 
corrected the definition of Leptopodia by 
detailing the arrangement of the abdominal 
segments. By today’s standards, however, 
there are no characters described for L. or- 
nata that are of diagnostic value. Rathbun 
(1925) listed the type specimen of L. ornata 
as not in the British Museum and perhaps 
not extant. Yang (1967) requested Dr. Isa- 
bella Gordon to make another search for 
the specimen, but that search was also fruit- 
less and the type must be assumed lost. Lep- 
topodia ornata Guilding, 1825, is herein 
designated a junior synonym of Stenorhyn- 
chus seticornis (Herbst, 1788). ; 

Another name in the literature which may 
have precedence for the proposed new 
species is Leptopodia vittata. Kingsley (1880) 
reported the presence of a specimen within 
the collections of the Museum of the Acad- 
emy of Natural Sciences of Philadelphia la- 
beled “‘Leptopodia vittata Guer., Senegal.” 
He stated that it might represent a manu- 
script name and that no published descrip- 
tion could be found of the species. Lepto- 
podia vittata Kingsley, 1880, is not available 
because it was published only in synonymy 
and was not adopted before 1961 as a name 
for a species. Manning & Holthuis (1981) 
listed this species as synonymous with the 
West African S. /anceolatus Brullé, 1837. 

Goeldi (1886) described Leptopodia lin- 
eata from Rio de Janeiro and Cabo Frio, 
Brazil. The description is somewhat vague, 
not detailed enough to determine which of 
the 2 western Atlantic species it represented, 
and the illustrations are not sufficiently de- 
tailed to accurately identify the taxon by 
today’s standards. An attempt to locate the 


VOLUME 102, NUMBER 3 


type series of L. /ineata at the major zoo- 
logical museums of Europe and Brazil was 
fruitless; the type must be assumed lost. 
Stenorhynchus material obtained from Cabo 
Frio and Rio de Janeiro revealed only S. 
seticornis. A search by Dr. W. Zwink (Mu- 
seu Nacional, Rio de Janeiro) of holdings 
of local specimens of Stenorhynchus showed 
no representatives of a second species. It 
may be conclusively presumed that this tax- 
on 1s conspecific with S. seticornis and is 
herein designated a junior synonym. 

The only remaining specific name that 
needs to be mentioned here is Pactolus bos- 
ci, Leach, 1815. This species was based on 
a single specimen of unknown origin found 
in the holdings of the British Museum (Nat- 
ural History). As the name has been sup- 
pressed by the ICZN (opinion 763) it is not 
available for use. 


Systematics 


Due to the extremely common nature of 
Stenorhynchus species and their wide geo- 
graphic range, it is nearly impossible to de- 
tail all of the workers who have dealt with 
western Atlantic members of the genus in 
the past 200 years. It is not attempted here. 
In most cases, it is not feasible to determine 
accurately from the literature which of the 
two species was the subject of each report. 
A great many of the citations are species 
listings and not accompanied by diagnoses, 
ecological data, or illustrations which would 
help to determine the identity of the taxon 
reported. However, a few records are suf- 
ficiently detailed (e.g., Hay & Shore 1918; 
Williams 1965, 1984) or have illustrations 
which help to clear up some of the confu- 
sion. For these reasons, the synonymies that 
follow are abbreviated and by no means de- 
tailed accounts of references to Stenorhyn- 
chus species in the literature. 

The materials examined in the following 
species accounts are housed at the United 
States National Museum of Natural History 
(USNM), Florida Department of Natural 


625 


Resources, Marine Research Laboratory 
(FSBC), Museu Nacional, Rio de Janeiro, 
Gulf Coast Research Laboratory (GCRL), 
Dauphin Island Sea Lab (MESC), and the 
University of Southwestern Louisiana 
(USLZ). 


Stenorhynchus seticornis (Herbst, 1788), 
redescription 
Figs. la, b, d—g, 2, and 3 


Oost-Indische Zee-Krabbe Slabber, 1778: 
162: pli 18; fig. 2. 

Cancer seticornis Herbst, 1788:229, pl. 16, 
hie tee 1803-27, pl 55; fig.. 2: 

Cancer sagittarius Fabricius, 1793:442. 

Leptopodia ornata Guilding, 1825:335. 

Leptopodia lineata Goeldi, 1886:37, pl. 3, 
figs. 24-31. 

Stenorynchus sagittarius.—Rathbun, 1901: 
53.—Hay & Shore, 1918: 455, pl. 37, fig. 
8 (in part). 

Stenorynchus seticornis.—Rathbun, 1925: 
14, pls. 2 and 3 (in part).—Abele, 1970: 
137 p. 

Stenorhynchus seticornis.— Williams, 1965: 
244, figs. 222 and 223K (in part).—Feld- 
er, 1973:48, pl. 7, fig. 1.—Yang, 1976.— 
Felder & Chaney, 1979:27.—Wicksten, 
1980:150.— Williams, 1984:304 (in part). 


Material examined.—Table 1. 

Diagnosis.—Carapace naked, rostrum 
covered with short dense felt and setae, be- 
coming longer and thicker distally. No spines 
at distal end of basal antennal article, single 
inter-antennular spine directed posteriorly. 
Chelipeds hairy, palm from three to four 
times length of fingers in mature males, not 
as stout in females. Merus of third maxil- 
liped normally with small spine on antero- 
distal angle. Pereopods, abdomen, and ster- 
num bearing short pubescence. 

Description of male neotype (USNM 
42956).—Carapace subtriangular, smooth, 
naked, regions slightly defined; intestinal and 
cardiac region inflated and separated by 
shallow furrow from posterior regions; 
branchial region inflated, delimited by shal- 


626 


Table 1.— Material examined; Stenorhynchus seticornis. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Catalog number Location Material Depth (m) 

USNM 42956 (Neotype of 

Stenorhynchus seticornis) Curacao, Suriname 16 = 
USNM 19941 Rio de Janeiro, Brazil 16 — 
USNM 76441 Gallows Bay, St. Croix, Virgin Islands 13 = 
USNM 46697 Aucilla, FL 19 13 
USNM 43065 Montego Bay, Jamaica 26 — 
USNM 43066 Jamaica 1¢ — 
USNM 19580 Kingston Harbor, Jamaica 1¢ — 
USNM 31040 Grolding Cay, Bahamas 16 _ 
USNM 11232 33°34'N, 77°42'W iS. 9 16.5 
USNM 19940 Bay of Bahia, Brazil 19 — 
USNM 22554 Santa Marta, Colombia 1¢ — 
USNM 69601 18°30'N, 66°04’W Puerto Rico 1¢ 366 
USNM 101712 16°05'N, 82°05’'W 16 44 
USNM 137761 Grand Island, Trinidad 36 69 
USNM 137020 00°15'S, 46°45’W 14,22 28 
USNM 171562 Tongue of the Ocean, Bahamas 16 — 
USNM 184122 Palm Beach, FL 16 22. 
USNM 58047 Shoal Banks, Barbados 16 55 
USNM 154711 Grenada 1¢ — 
USNM 321390 Belize 1¢ — 
USNM 103268 06°51'N, 54°53’W Suriname 12 ait 
USNM 43060 Montego Bay, Jamaica 1¢ — 
USNM 105004 08°40'N, 77°10’'W Gulf of Darien 19 61 
USNM 137020 00°15'S, 46°45’W Para, Brazil | row) Xe: 27, 
USNM 107804 off Houma, LA 1¢ _ 
USNM 103267 06°49'N, 55°54’W 12 47 
USNM 103504 07°40'N, 57°34’'W 1¢ — 
USNM 17374 22°18'N, 87°04’W 1¢ 44 
USNM 49082 10 miles south of Key West, FL 12 229 
USNM 42955 Curacao 12 _ 
USNM 101713 01°S50’N, 47°93 1'W 19 85 
USNM 43062 Montego Bay 14 _ 
USNM 7653 St. Thomas 1¢ _ 
USNM 17373 (in part) 35°08'30’N, 75°10’'W 1é 90 
USNM 137019 04°46’N, 51°21’W 16 59 
USNM 103265 06°48'N, 54°54’W 26 46 
MESC 6187-0121 28°26'N, 84°21'W 3: 3.2 ~ 
MESC 6187-0113 28°25'N, 84°19'W 64,42 — 
MESC 6187-0114 30°02'30”N, 86°06'30”W 26 ul! 
MESC 6187-0109 28°36'N, 84°16'W To 52 _ 
GCRL 164:695 29°43'N, 88°26'W 14,32 — 
FSBC I 31042 (in part) 30°30'N, 80°15’W 22 47 
FSBC I 31045 (in part) 30°20'N, 80°14’'W 13 65 
FSBC I 31043 (in part) 30°31'N, 80°10'W Lig 22.9 64 
FSBC I 31044 (in part) 30°20'N, 80°17'W 1¢ 46 
FSBC I 31032 27°40'N, 80°06'W 1¢ 27 
FSBC I 31030 27°10'N, 80°01'W 1¢ 46 
FSBC I 31029 27°10'N, 80°01'W 19 45 
Museu Nacional Rio de Janeiro Cabo Frio, Brazil 14,12 — 
Museu Nacional Rio de Janeiro Rio de Janeiro, Brazil 146,12 — 
Museu Nacional Rio de Janeiro Guanabara, Brazil ivGul-e — 


VOLUME 102, NUMBER 3 


low margin; hepatic region inflated, with 
well-defined ventral margin, but remaining 
ventral margin ill defined. Simple, strong 
deflexed postorbital spine; supra-orbital 
furrow shallow; rostrum flattened basally 
between eyes; short setae from base of ros- 
trum to tip, setae dense and increasingly 
long distally; subhepatic region inflated pos- 
teriorly, margins well defined; Ist rostral 
spine directed laterally, slightly deflexed, 
followed by up to 16 large spines on lateral 
margins, directed forward and occasionally 
downward; rostrum 1.8 times length of car- 
apace behind transverse line connecting base 
of eyestalks dorsally. Basal antennal article 
elongate, with longitudinal ventral furrow, 
strong spine anterolaterally directed on ven- 
tral margin; septum dividing antennular si- 
nuses with posteriorly directed spine; an- 
terolateral margin of sinus defined by 
upturned border; lateral and anterior mar- 
gins of buccal cavity with raised ridge, acute 
small spine at anterolateral angle of mouth 
frame. Exopod of third maxilliped with 
maximum width one-third distance from 
base, narrowing distally; internal margin of 
merus straight, with strong spine at antero- 
mesial angle and small spine on exterior 
margin posterior to articulation with palp. 
First pereopods greatly elongate, covered 
with short dense pile; basis inflated, tuber- 
culate, with spine on interior margin; merus 
tuberculate, with region of few tubercles 
dorsolaterally, six to nine strong spines on 
mesial row, single spine dorsally, two spines 
in lateral row, ventral spine toward distal 
margin, three or four spines at articulation 
with carpus; carpus with three strong spines 
dorsally, three distal spines ventrally and 
numerous tubercles; propodus covered with 


strong tubercles and low pile which becomes 


denser and longer at base of finger, pile ex- 
tends laterally along propodus and onto 
ventral surface at base of finger, propodus 
three times length of dactyl; dactyl stout, 
with long dense setae dorsally and laterally 
at base, thinning distally to become sparse 
long setae, tuberculate dorsally. 


627 


Second pereopod longer than first and 
covered with short dense pile; merus with 
4 longitudinal rows of spines, 5 or 6 spines 
in dorsal row, 5-7 spines in lateral row con- 
centrated in distal '2, 11 mesial spines dis- 
tributed along length of segment, 2 ventral 
spines in distal 12; 3 spines at articulation 
with carpus slightly longer than others; car- 
pus with pair of dorsal spines at midlength 
and 3 spines at articulation with propodus 
longer than others; length of carpus and 
propodus equal to dactyl but shorter than 
merus; propodus with 1 1-13 spines, slightly 
compressed laterally, 2 spines at articula- 
tion ventrolaterally; dactyl elongate, slightly 
curved, little compressed laterally, with 5 
longitudinal rows of setae. 

Third pereopod shorter than second, cov- 
ered with short dense pile; merus with 5 
dorsal spines along length, 8—10 spines along 
inner row, 3—4 spines in distal '2 on external 
row, single spine on ventral margin in distal 
Y), 3 spines at articulation with carpus; car- 
pus with pair of dorsolateral spines at mid- 
length and 3 spines at articulation with 
propodus; propodus with 11-13 small 
spines, slightly compressed laterally, with 2 
spines at articulation; dactyl % length carpus 
and propodus combined, with 5 longitudi- 
nal rows of setae, curved slightly in distal 
i. 

Fourth pereopod shorter than third, with 
covering of short pile; merus with four to 
five spines on internal row and ventrolateral 
spine at articulation slightly enlarged; car- 
pus with pair of dorsal spines at midlength 
and three spines at articulation with prop- 
odus; propodus with eight or nine small 
spines; dactyli damaged, slightly com- 
pressed laterally, and bearing five longitu- 
dinal rows of setae. 

Fifth pereopod shorter than fourth, cov- 
ered with short dense pile; merus with 4 
dorsal spines, single ventrolateral spine and 
2—4 mesial spines; carpus with pair of spines 
dorsolaterally at midlength, 3 terminal 
spines at articulation; propodus with 9-10 
low spines, slightly compressed; dactyl elon- 


628 


gate, curved distally with 5 rows of longi- 
tudinal setae. 

Abdominal segments 5 and 6 fused, with 
sutures indicated; segment 1 slightly longer 
than wide, naked along elevated midlength 
but with setae in depressions; segments 3 
and 4 subequal, longer than 2; 3 widest an- 
teriorly, segments 5 and 6 fused, segment 7 
is 1.75 times long as wide with hollowed 
protuberance for locking mechanism and a 
small tubercle medially in proximal one- 
half. Sternum with 22 large tubercles and 
few low setae; sternite 5 with 2 tubercles at 
articulation with basis of maxilliped; ster- 
nite 4 with 8—9 tubercles at articulation with 
first pereopod; plastron with arcuate ridge 
in anterior '/ lined with setae, lateral margin 
parallel. 

Color.—Carapace with alternating stripes 
of off-white and brown-maroon, white 
stripes on dorsum of carapace reminiscent 
of inverted V’s, large white bands originat- 
ing dorsally between fourth and fifth legs 
join anterior to eyes, every other white stripe 
smaller than preceding; white bands origi- 
nating at tip of dactyl of pereopods, con- 
tinuing dorsally along leg onto carapace, or 
originating at posterior margin of carapace 
between coxae of legs. Carapace with 4 ma- 
jor white longitudinal bands, one running 
along the pterygostomial ridge. Distal *4 of 
fingers on chelae blue. Inner surface of fin- 
gers each with 2 whitish spots, forming rough 
circle when fingers close. Merus of cheliped 
with large orange-yellow spot at base of dis- 
tal spine. White band bordered with ma- 
roon extending obliquely across merus and 
palp of third maxilliped. Broad yellow-white 
band connecting coxae of first pereopods 
with area under third maxilliped. 

Range.— Material assignable to S. seti- 
cornis has been examined from Cape Fear, 
North Carolina, through the Gulf of Mexico 
southward to the mouth of the Amazon 
River, and Cabo Frio, Brazil. Bathymetric 
records are from | to 366 m. 

Remarks.—The above technical descrip- 
tion is drawn from the designated neotype, 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


a mature male collected at Curacao, Dutch 
West Indies. This specimen is the form 
commonly found on the rock outcroppings 
or coral reefs in temperate and tropical 
waters of the western Atlantic, and reported 
by Herbst (1803) from Guadeloupe. 

Variation.— Stenorhynchus seticornis ex- 
hibits considerable morphological variation 
which appears to be related to the substrata 
from which the specimens are collected. One 
variant, herein designated S. seticornis form 
A, is most often collected on hard rocky 
bottoms (i.e., rock outcroppings, reefs, jet- 
ties) or immediately adjacent to these hard- 
bottom types. It is a large heavy bodied form, 
having an extremely long rostrum with sub- 
parallel margins throughout most of the 
length. This form was mentioned by Yang 
(1967) as “atypical.” 

Stenorhynchus seticornis form B differs 
from the above by characters enumerated 
below. Form B is the ecotype most often 
found in the northern Gulf of Mexico on 
mud bottoms and in grass beds. It was listed 
by Yang (1967, 1976) as Stenorhynchus se- 
ticornis and the complete larval develop- 
ment has been described. It is the smaller 
of the two forms. In SEM micrographs, the 
male pleopod of S. seticornis form A (Fig. 
3) shows little variation, other than size, 
from that of S. seticornis form B (Fig. 3). 
The somewhat narrower opening of the apex 
and the slightly shorter apex may represent 
differences in the orientation of the gonopod 
at the point where the photographs were 
taken. This very minor difference, and the 
observed gradation between the two forms, 
are well explained by ecophenotypic vari- 
ation. 

Stenorhynchus seticornis form B is sepa- 
rated from S. seticornis form A by: 1) small- 
er overall body size, 2) females and subadult 
males with large hiatus at base of moveable 
fingers, and 3) setae on the dactyl and prop- 
odus of the first leg not forming thick mat 
continuous on dorsal surface. 

Considerable intergradation is found be- 
tween the two forms of S. seticornis, espe- 


VOLUME 102, NUMBER 3 


Fig. 3. Scanning electron micrographs of the tips of male gonopods: Above, Stenorhynchus seticornis form 
A; Below, S. seticornis form B. 


630 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Table 2.— Material examined; Stenorhynchus yangi. 


eee eee 


Catalog number Location Material Depth (m) 
a 
USNM 211812 (HOLOTYPE) 33°49'N, 76°43'W ihc 81 
USNM 211855 (PARATYPE) 26°45'70"N, 84°00'13"W 18 89 
USNM 211803 (PARATYPES) 33°48'46’N, 76°35'W 346,22 60 
USNM 67800 18°30'30’N, 66°23'50”"W 16 He 
USNM 101715 34°17'N, 76°01'W 19 137 
USNM 120135 Dominican Republic 1¢ pa 
USNM 103269 6°52'N, 54°53'W 14,12 288 
USNM 73401 Barbados ia = 
USNM 73086 Grand Cayman 1é 220 
USNM Uncatalogued Oregon St. 4391 12°33) NE 09 Ww. rote li i: 
USNM Uncatalogued Oregon St. 3605 12° T6IN 82°53 W 16 1D 
USNM 69601 18°30'30’N, 66°04'05”W 16 365 
USNM 92646 2T°-1S NSO 2 TOW 16 64 
USNM 211815 33°48'42”N, 76°34'12"W 1. dyke 102 
USNM 24418 Mayaguez Harbor, Puerto Rico 16 137 
USNM 11379 Key West, FL 12 108 
USNM 11219 Martha’s Vineyard 16 62 
USNM 211807 33°47'36’N, 76°34'24"W 22 116 
USNM 9496 Key West jewel) S _ 
USNM 103504 70°40’N, 57°34'W 12 — 
USNM 6934 9°30'15”N, 76°20'30”W Sid oe — 
USNM 9862 33°18'30’N, 77°07'W 16 174 
USNM Uncatalogued Oregon St. 4391 12°33'N, 71°09'W [Boe LS 73 
USNM Uncatalogued Oregon St. 3605 12°16'N, 82°53'W 16 7h: 
USNM 17373 (in part) 35°08'30’N, 75°10’'W 22 90 
GCRL 160:165 25°35'N, 83°42'’W 5:62 110 
FSBC I 31031 27°40'N, 79°59'W ye toe |e: 64 
FSBC I 31049 30°00'N, 80°15’W 16 91 
FSBC I 31042 (in part) 30°30'N, 80°15'W 12 47 
FSBC I 31048 30°11'N, 80°15’W 16 64 
FSBC I 31033 27°50'N, 79°58’'W seks 92 
FSBC I 31045 (in part) 30°20'N, 80°14’W 104,42 65 
FSBC I 31041 30°40'N, 80°06’W PL 91 
FSBC I 31036 28°40'N, 80°06’W 192 64 
FSBC I 31043 (in part) 30°21'N, 80°10’W leds Ee 64 
FSBC I 31046 30°20’N, 80°12’W 16 92 
FSBC I 31040 30°40'N, 80°07’W 16 64 
FSB@ RSt037 28°50'N, 80°09’W 16 64 
FSBC I 31034 28°30'N, 80°01'W 192 91 
FSBC I 31047 30°10'N, 80°14’W 12 91 
FSBC I 31038 29°00'N, 80°10’W 1¢@ 64 
FSBC I 31044 (in part) 30°20'N, 80°17’W 192 46 


cially in areas of rock outcroppings adjacent 
to muddy bottoms. In this situation, spec- 
imens are larger than those found on muddy 
bottoms, but often possess the hiatus at the 
base of the fingers and the greatly elongated 
rostrum of the form common to reefs. Sim- 
ilarly, specimens from grass beds adjacent 
to coral heads exhibit characteristics of both 
forms A and B. 


Specimens of S. seticornis exhibit great 
variation in a number of characters, includ- 
ing length of the rostrum and the fingers. 
To document this variation, I measured the 
total postorbital carapace lengths and total 
carapace lengths on specimens for which the 
rostrum was unbroken on specimens which 
represented the entire geographic range. The 
postorbital measurement was determined 


VOLUME 102, NUMBER 3 


from the posterior margin of the carapace 
to a point midway between the base of the 
eyestalks. Additional measurements of the 
moveable finger and palm length were tak- 
en. The range of the ratio of postorbital 
length to total carapace length is slightly dif- 
ferent in males (n = 20:0.27-0.40) and fe- 
males (n = 5:0.33-0.42), but no real di- 
morphism in carapace dimensions is evident 
which is attributable to the sex of the spec- 
imens. These differences bear no relation- 
ship to either the size of the individual or 
the location from which it was collected. 
However, differences in the range of ratios 
of the finger to palm lengths do seem at- 
tributable to sexual dimorphism. Although 
there is some overlap in the range, males (n 
= 22) tend to have shorter fingers, relative 
to the palm length (0.24—0.39) than do fe- 
males (n = 7:0.37—0.45). This trend is also 
evident in S. yangi. 

A detailed examination of the variation 
found in S. seticornis has not produced suf- 
ficient data to warrant the establishment of 
separate subspecies. Until the larval devel- 
opment of S. seticornis form A has been 
documented, I prefer the designation “‘form”’ 
to a questionable subspecies status. 


Stenorhynchus yangi, new species 
Fig. lc 


Stenorynchus sagittarius.—Hay & Shore, 
1918:455, pl. 37, fig. 8 (in part). 

Stenorynchus seticornis.—Rathbun, 1925: 
14, pls. 2 and 3 (in part). 

Stenorhynchus seticornis.— Williams, 1965: 
244, figs. 222 and 223K (in part).— 1984: 
304 (in part). 

Stenorhynchus sp. A.— Yang, 1967; 459 p.— 
1976:158. 


Material examined.—Table 2. 

Diagnosis.—Carapace naked, rostrum 
devoid of setae or felt; no spines at distal 
end of basal antennal article; interanten- 
nular septum without posteriorly directed 
spinous process; chelipeds hairy, palm only 
twice length of moveable fingers in males, 
1.5 to 2 times length of dactyl in females. 


631 


Merus of third maxilliped with vestigial 
spine at anteromedial angle. Ambulatory 
legs, abdomen and sternum without pubes- 
cence. Regions of carapace well defined and 
inflated. 

Description of holotypic male (USNM 
211812).—Carapace subtriangular, smooth, 
naked, regions well defined; branchial re- 
gions inflated; intestinal and cardiac regions 
elevated in midline and separated from 
branchial and cardiac regions by sulcus; gas- 
tric region inflated; hepatic region moder- 
ately inflated and defined by furrows on all 
sides. Strong postorbital tooth bifid, de- 
flexed and directed slightly forward. Shal- 
low supraorbital furrow; rostrum flattened 
between eyestalks, devoid of setae; subhe- 
patic region inflated posteriorly with well 
defined margins; first rostral spine directed 
slightly forward; rostrum about equal to 
length of postorbital carapace, naked, broad 
basally and tapering to acute apex, bearing 
17 to 19 large lateral spines; basal antennal 
article with shallow longitudinal furrow and 
strong anteriorly directed spine on ventral 
margin; septum dividing antennular sinuses 
well developed, with rounded ventrum; an- 
terolateral margin of sinus simple. Lateral 
and anterior margins of buccal frame with 
raised margin and weak spine at anterolat- 
eral angle; exopod of third maxilliped with 
maximum width at one-half length from 
base, narrowing distally, internal margin of 
merus concave, occasionally with small 
spine at antero-internal angle, small spine 
posterior to articulation with palp. 

All pereopods with few scattered setae. 
First pereopod greatly elongate; ischium 
slightly inflated, smooth; merus with nu- 
merous spines in longitudinal rows; two 
spines in distal one-half of lateral row; three 
spines in dorsal row evenly spaced, enlarged 
terminal spine at articulation with carpus; 
six spines on interior row; carpus with two 
dorsal spines at midlength and pair of ter- 
minal spines ventrally; propodus with small 
scattered tubercles. Felt and dense setae lat- 
erally at base of finger in small patch with 
longer sparse setae on immoveable finger; 


632 


dactyl with small patch of setae at base, with 
few long scattered setae on surface, small 
hiatus at base. 

Second pereopod elongate, longer than 
first; ischium very short, tuberculate dor- 
sally; merus greatly elongate, with about 20 
spines arranged in 4 longitudinal rows, 3 
larger terminal spines; carpus short, about 
. length of merus, with 3 dorsal and 3 ter- 
minal spines; propodus elongate, *4 length 
of merus, with numerous spinules and tu- 
bercles, | lateral terminal spine; dactyl elon- 
gate, 44 length of merus, slightly compressed 
laterally, lined with spinules and with cor- 
neous tip. 

Third pereopod shorter than second; is- 
chium very short, tuberculate; merus elon- 
gate, with approximately 15 spines arranged 
primarily in 2 dorsal rows, 3 terminal spines; 
carpus short, %4 length of merus, 3 dorsal 
and 3 terminal spines; propodus elongate, 
*3; length of merus, with approximately 10 
dorsal and 2 terminal spines; dactyl elon- 
gate, ¥, length of merus, with numerous spi- 
nules, slightly compressed laterally with 
corneous tip. 

Fourth pereopod shorter than third, is- 
chium tuberculate, very short; merus elon- 
gate, roughly 20 spines arranged in 2 pri- 
mary dorsal rows, 3 terminal spines; carpus 
short, 4 length of merus with 3 dorsal and 
3 terminal spines; propodus elongate, * 
length of merus with about 10 spines dor- 
sally and 3 terminal spines; dactyl elongate, 
47, length of merus with numerous spinules 
and corneous tip, slightly compressed lat- 
erally. 

Fifth pereopod shorter than fourth, is- 
chium very short, slightly tuberculate; me- 
rus elongate, with 10 spines in 2 dorsal rows, 
and 3 terminal spines; carpus short, 14 length 
of merus, with 3 dorsal and 3 terminal 
spines; propodus elongate, equal in length 
to merus, with spinules mainly in single dor- 
sal row, slightly compressed laterally, nu- 
merous spinules along length; dactyl with 
corneous tip. 

First abdominal segment little longer than 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


wide, elevated along midline, segments 2 
and 3 very short but broad; segments 4 and 
5 somewhat longer and constricting to nar- 
rowest part of abdomen, segments 6 and 7 
fused, no external indication of fusion vis- 
ible. Tubercles scattered on sternum with 
very few setae apparent; anterior extension 
of sternum with ridge below mouth frame 
bearing setal row and two spines at center 
of process. | 

Color.—Carapace with four triangular 
white stripes, yellow and orange-reddish- 
brown areas surrounding white stripes. Lat- 
eral white stripe passes through orbital area 
and is continued onto lateral edge of ros- 
trum. Lateral teeth of rostrum white; tip of 
rostrum dark brown. Fingers of cheliped 
purple for distal two-thirds to three-fourths 
of length, teeth white, remainder of fingers 
tan; carpus tan with brown stripe proxi- 
mally; merus dark brown distally, pale yel- 
low spot behind large distal tooth. Ambu- 
latory legs with faint light dorsal stripe 
continuous from carapace, distal ends of 
segments darker, merus with darker areas; 
dactyli with distal '4 white, apex translucent 
with penultimate tan band and then white. 
Legs with faint dark/light bands. 

Range.— Stenorhynchus yangi has been 
collected from Martha’s Vineyard south 
through the Gulf of Mexico to Suriname in 
depths from 31 to 365 m. 

Variation.—Within S. yangi, variation 
has been noted in the shape of postorbital 
spines, characteristic robustness of the car- 
apace and relative lengths of the rostrum 
and moveable finger of the first leg. Wil- 
liams (1965:244) reported the postorbital 
spine of S. seticornis as ““occasionally bifid.” 
During the course of this study, no speci- 
mens of Stenorhynchus seticornis were ob- 
served which possessed bifid postorbital 
spines. However, it is common to come 
across large specimens of S.. yangi with this 
condition. Several large individuals of S. 
yangi were examined which had single and 
double spines on alternate sides of the car- 
apace. Specimens with trifid spines were also 


VOLUME 102, NUMBER 3 


observed, and it is probable that the indi- 
viduals reported by Williams (1984) with 
bifid spines represent S. yangi, as the gon- 
opod figured in that work (fig. 2410) cor- 
responds with S. yangi, not S. seticornis. 
The characteristic robustness or swelling of 
the various regions of the carapace also var- 
ies within S. yangi. This and the relative 
lengths of the pereopods are probably re- 
lated to maturity of the individual. Consid- 
erable variation was also noted in spination 
of the pereopods. 

Perhaps the two features which exhibit 
the greatest degree of variation are the ratio 
of total carapace length to postorbital car- 
apace length and the ratio of finger to palm 
length of the first pereopod. Only individ- 
uals with the rostrum intact (n = 29) were 
used for carapace length measurements, with 
individuals from the extreme ends of the 
geographic range included. Among males (n 
= 19), the postorbital carapace length ac- 
counted for one-fourth to one-half of the 
total length of the individual (0.27-—0.49); 
little difference from this ratio was noted 
among females (0.31—0.48). However, the 
range ratio of finger to palm length was no- 
tably different among males (n = 22:0.24— 
0.39) and females (n = 13:0.36-0.49). 

A single anomaly was noted in a specimen 
from the Caribbean (USNM Accession 
#42869). This individual is a mature male 
with a bifid rostrum, but is normal in all 
other respects. 

Etymology.—It is my pleasure to name 
this species for Dr. Won Tack Yang (Texas 
Biomedical Institute) who, on the basis of 
larval characters, first recognized the pos- 
sibility of its distinctness. 

Remarks.—Yang (1967, 1976) per- 
formed much of the complicated work in- 
volved in reviewing the confused history of 
this genus and gave definitive proof that at 
least two species of Stenorhynchus are pres- 
ent in western Atlantic waters. His work was 
the first to correct the mistaken records of 
S. seticornis in the eastern Atlantic and 
showed S. /anceolatus Brullé, 1837, to be 


633 


the correct name for the West African 
species. 

Yang (1967) indicated that S. yangi may 
be a deep water species and presented data 
which appeared to show that the species was 
most often collected in waters deeper than 
65 m. He noted that in the collections at the 
University of Miami Marine Laboratory and 
those examined from the U.S. National Mu- 
seum of Natural History, this species’ 
bathymetric range was from 31-119 m. He 
suggested the possibility of an isotherm di- 
viding the preferred habitats of the two 
species. Abele (1970) indicated a restricted 
occurrence of S. yangi to waters over 50 m. 
However, because I have not re-examined 
that material, that report of the species en- 
countered is unconfirmed. 

The robust nature of the carapace of this 
species, the absence of the interantennular 
spine, the lack of rostral setation, the shape 
of the first male pleopod and the form of 
the spermathecae of the female easily dis- 
tinguish S. yangi from S. seticornis. Differ- 
ences in the color patterns of the two species 
also aid in their identification; however, this 
difference may be quickly obscured by pres- 
ervation techniques and be of little use to 
the researcher studying museum specimens. 
Differences in relative lengths of the am- 
bulatory legs may also be of aid in the sep- 
aration of S. yangi from S. seticornis. How- 
ever, the fragile nature of members of this 
genus makes studies of this type very dif- 
ficult as legs are rarely collected intact. 

Discussion.—Taxonomic confusion sur- 
rounding the identity of S. seticornis and S. 
yangi is partly due to the great amount of 
variation found within the two taxa. The 
range of variation in the ratio of postorbital 
to total carapace lengths differs little from 
S. seticornis (0.27—-0.49) to S. yangi (0.27- 
0.42). A broad range of variation is also 
evident in the ratio of finger to palm length 
(0.24—0.49 in S. seticornis and 0.24—0.45 in 
S. yangl). 

Because of the taxonomic problems, eco- 
logical works that have dealt with this group 


634 


must be used with reserve. Bathymetric rec- 
ords (Rathbun 1925), distributional records 
(Holthuis 1959; Williams 1965), substrate 
preferences (Rathbun 1925) and ecological 
and behavioral works (Hartnoll 1965; Barr 
1971, 1975) must all be viewed with caution 
and records re-examined where feasible. Al- 
though the specific identity of the arrow 
crabs used in the reproductive studies may 
be of little significance in the understanding 
of the group’s behavior, it must be recog- 
nized that doubt exists as to which taxon 
was being studied by Schone (1968). Barr’s 
(1971, 1975) field work suggests that S. se- 
ticornis is a facultative filter feeder which 
climbs to the top of an outcropping or reef 
at dusk. By sitting motionless, the crab al- 
lows the passing debris to collect on the se- 
tae of its body during the night, and the 
following day is spent cleaning off the en- 
trapped food. No data were given by that 
author to indicate the frequency of this feed- 
ing mode. R. H. Gore (pers. comm.) noted 
that in aquaria, S. seticornis will snip off 
and eat the protruding siphon of the gastro- 
pod Nassarius vibex. Mary K. Wicksten 
(pers. comm.) has indicated that S. seticor- 
nis may perform an unusual decorating act 
by storing food gathered from the substra- 
tum on the rostral setae to be eaten later. 
This 1s a modification of the usual decora- 
tion behavior used for concealment and 
camouflage (Wicksten 1980). An alternative 
feeding behavior is dictated for S. yangi be- 
cause of the lack of rostral setae. No study 
has been accomplished to define the niche 
requirements of these two species and how 
these requirements may differ. 

The complete larval development of S. 
seticornis form A was described by Yang 
(1967, 1976). He detailed three zoeal stages 
and the megalopa obtained from females 
collected in Biscayne Bay, Florida. Yang 
(1967) specifically mentioned the hiatus at 
the base of the fingers, a character previ- 
ously listed as useful in separating the two 
forms of S. seticornis. A single zoeal stage 
of S. yangi (Stenorhynchus seticornis of Yang 
1967, 1976) was described from a specimen 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


collected in about 225 m of water. The lar- 
vae of the two species are sufficiently dis- 
tinct to allow quick separation of the species 
because of a large lateral bifurcation on the 
carapace of S. yangi. This situation is un- 
usual in that the adults are similar enough 
to have been united under a single name for 
many years, but the larvae are quite differ- 
ent at the first zoeal stage. 


Acknowledgments 


This study has benefitted from the help 
of many individuals over several years. 
However, because of the diversity of opin- 
ions offered (often conflicting), not all of the 
reviewers’ suggestions have been incorpo- 
rated into this report. Any oversights or 
shortcomings of this report are the sole re- 
sponsibility of the author. John S. Garth 
and Jurgen Sieg helped gather obscure orig- 
inal descriptions. Robert H. Gore, Joseph 
Fitzpatrick, Richard W. Heard, Jr., Austin 
B. Williams, Raymond B. Manning, Robin 
M. Overstreet, Lipke B. Holthuis, and Dar- 
ryl L. Felder all provided helpful discus- 
sions, comments or reviews. Thomas S. 
Hopkins, David K. Camp, Walter Zwink, 
Darryl L. Felder and Torben Wolff provid- 
ed material. Darryl L. Felder and the staff 
of the University of Southwestern Louisi- 
ana Electron Microscopy Center provided 
the SEM micrographs. The reviewers of the 
journal provided much needed guidance in 
a very gracious manner. Linda Lutz pre- 
pared Fig. 1 and Cynthia B. Dickens typed 
the original manuscript. My gratitude is ex- 
tended to each of these for their help. 


Literature Cited 


Abele, L. G. 1970. The marine decapod Crustacea 
of the northeastern Gulf of Mexico. MS Thesis, 
March, 1970. Florida State University, Talla- 
hassee, Florida, 137 pp. 

Barr, L. 1971. Observations on the biology of the 
arrow crab, Stenorhynchus seticornis (Herbst) in 
Lameshur Bay, St. John, Virgin Islands. Pp. 213- 
220 inJ. Miller, J. van Derwalker, & R. Walkers, 
eds., Scientists-in-the-sea. Department of the 
Interior, Washington, D.C. 


VOLUME 102, NUMBER 3 


. 1975. Biology and behavior of the arrow crab, 
Stenorhynchus seticornis (Herbst), in Lameshur 
Bay, St. John, Virgin Islands. Jn Results of the 
Tektite Program.— Bulletin of the Museum of 

Natural History, Los Angeles County 20:47-56. 

Bosc, L.A. G. 1802. Histoire naturelle des crustacés, 
contenant leur description et leur moeurs; avec 
figures desinés d’aprés nature. Chez Deterville, 
Paris. Vol. 1:258 pp. 

Brulleé, M. 1837, 1839. Crustacés. Jn P. Barker-Webb 
& S. Berthelot— Histoire Naturelle des Iles Ca- 
naries, Zoologie, 2(2:Entomologie):13—18 (1839). 
Atlas (1837), unnumbered plate. 

Fabricius, J. C. 1793. Entomologia systematics 
emendata et aucta secundum classes, ordines, 
genera, species adjectis synonimis, loci, obser- 
vationbus, descriptionbus. Vol. 2:519 pp. Haf- 
niae, C. G. Proft. 

Felder, D. L. 1973. An annotated key to crabs and 
lobsters (Decapoda, Reptantia) from coastal 
waters of the northwestern Gulf of Mexico. Lou- 
isiana State University Sea Grant Publication 
Number LSU-SG-73-02. 103 pp. 

—., & A. H. Chaney. 1979. Decapod crustacean 
fauna of Seven and One-Half Fathom Reef, 
Texas: Species composition, abundance and 
species diversity.—Contributions to Marine 
Science 22:1-29. 

Garth, J. S., & L. Holthuis. 1963. Stenorhynchus La- 
marck, 1818 (Crustacea, Decapoda): Proposed 
validation under the plenary powers with des- 
ignation of Cancer seticornis Herbst, 1788, as 
type-species.—Z.N.(S.) 751 Bulletin of Zoolog- 
ical Nomenclature 20(6):424—428. 

Goeldi, E. A. 1886. Studien tiber neue und wen- 
iger bekannte Podophthalmen Brasiliens. — Ar- 
chives fur Naturglijke 52(1):19—46, pls. 2, 3, figs. 
1-36. 

Guilding, L. 1825. An account of some rare West 
Indian Crustacea.— Transactions of the Linnean 
Society of London 14:334—338. 

Hartnoll, R.G. 1965. The biology of spider crabs: A 
comparison of British and Jamaican species. — 
Crustaceana 9:1-16. 

Hay, W. P., & C. A. Shore. 1918. The decapod crus- 
taceans of Beaufort, NC, and surrounding re- 
gion.— Bulletin of the U.S. Bureau of Fisheries 
35:369-475. 

Herbst, J. F. W. 1788-1803. Versuch einer Natur- 
geschichte der Krabben und Krebse nebst einer 
systematischen Beschricibung ihrer verschie- 
denen Arten, vol. 1:1—274; vol. 2:1-—225, Berlin, 
G. A. Lange. 

Holthuis, L. 1959. The Crustacea Decapoda of Su- 
riname (Dutch Guiana).— Zoologische Verhan- 
delling, Leiden 44:1-296. 

Kingsley, J. S. 1880. On a collection of Crustacea 

from Virginia, North Carolina and Florida, with 


635 


a revision of the genera of Crangonidae and Pa- 
laemonidae.— Proceedings of the Academy of 
Natural Sciences of Philadelphia, 31 (for 1879): 
383-427. 

Lamarck, J. B. de. 1818. Histoire naturelle des ani- 
maux sans vertébres, présentant les caractéres 
généraux et particuliers de ces animaux, leur 
distribution, leurs classes, leur familles, leur 
genres, et la citation des principales espéces qui 
s’y rapportent; précédée d’une introduction of- 
frant la determination des caractéres essentiels 
de l’animal, sa distinction du végetal et des autres 
corps naturelles, enfin, l’éxposition des prin- 
cipes fondamentaux de la zoologie. Ist ed., Vol. 
5:612 pp. Paris. 

Leach, W. E. 1814. Crustaceology. Jn D. Brewster, 

ed., Edinburgh Encyclopaedia, vol. 7:383-437. 

Edinburgh. 

1815. The zoological miscellany; being de- 
scriptions of new, or interesting animals. Vol. 
2:154 pp. London, Nodder and Son. 
Linnaeus, C. 1758. Systema naturae per regna tris 

naturae, secundum classes, ordines, genera, 
species, cum characteribus, differentiis, synon- 
ymis, locis. ed. 10, Vol. 1:824 pp. Stockholm, 
L. Salvius. 

Manning, R. B., & L. Holthuis. 1981. West African 
brachyuran crabs (Crustacea: Decapoda). — 
Smithsonian Contributions to Zoology 306:1- 
379. 

Milne-Edwards, A. 1875. Etudes sur les Xiphosures 
et les Crustacés de la région Mexicicaine. Jn 
Mission Scientifique au Mexique et dans l’Amé- 
rique Centrale, No. 3, pp. 57-120, Paris. 

International Commission on Zoological Nomencla- 
ture. 1966. Opinion 763. Stenorhynchus La- 
marck, 1818 (Crustacea, Decapoda): Validated 
under the plenary powers with designation of 
Cancer seticornis Herbst, 1788, as type-species. — 
Bulletin of Zoological Nomenclature 23(1):19- 
oA 

Paula, J. 1987. The first zoeal stage of Stenorhynchus 
lanceolatus Brullé, 1837 (Decapoda, Brachyura, 
Mayjidae).—Crustaceana 53(3):276—-280. 

Rathbun, M. J. 1897. A revision of the nomencla- 

ture.— Proceedings of the Biological Society of 

Washington 1 1:153-167. 

1900. The Brachyura and Macrura of Porto 
Rico.— U.S. Fisheries Commission 20:1—127. 

. 1925. The spider crabs of America. — Bulletin 

of the U.S. National Museum 129:1-613. 

Schone, H. 1968. Agonistic and sexual display in 
aquatic and semi-terrestrial brachyuran crabs. — 
American Zoologist 8:641-654. 

Slabber, M. 1778. Natuurkundige Verlustigingen, be- 
helzende microscopise Waarneemingen van in- 
en uitlandse Water- und Land-Dierren. Harr- 
lem, p. 162. 


636 


Weber, F. 1795. Nomenclator entomologicus secun- 
dum entomologiam systematicum ill. Fabricii, 
adjectis speciebus recens detectis et varietatibus. 
viii + 171 pp. Chilonii (Kiel) et Hamburgi. 

Wicksten, M. K. 1980. Decorator crabs.—Scientific 
American 242:146-154. 

Williams, A. B. 1965. The decapod crustaceans of 

the Carolinas.— U.S. Fishery Bulletin, 65(1):1- 

298. 

1984. Shrimps, lobsters, and crabs of the 
Atlantic coast of the eastern United States, Maine 
to Florida, Smithsonian Institution Press, 
Washington, D.C., 550 pp. 

Yang, W. T. 1967. A study of zoeal megalopal, and 
early crab stages of some oxyrhynchous crabs 
(Crustacea, Decapoda). Ph.D. dissertation, Uni- 
versity of Miami, Coral Gables, Florida. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


1976. Studies on the western Atlantic arrow 
crab genus Stenorhynchus (Decapoda, Brachy- 
ura, Majidae). 1. Larval characters of two species 
and comparison with other larvae of the Ina- 
chinae.—Crustaceana 31:157-177. 

Zimsen, E. 1964. The type material of I. C. Fabricius. 
656 pp., 2 pls. Copenhagen. 


Gary D. Goeke, Minerals Management 
Service, Gulf of Mexico OCS Region, Office 
for Leasing and Environment, 1201 Elm- 
wood Park Boulevard, New Orleans, Lou- 
isiana 70123-2394. 


PROC. BIOL. SOC. WASH. 
102(3), 1989, pp. 637-643 


HOBBSEUS YALOBUSHENSIS, A NEW CRAWFISH 
FROM CENTRAL MISSISSIPPI 
(DECAPODA: CAMBARIDAE) 


J. F. Fitzpatrick, Jr. and Craig A. Busack 


Abstract.—A new crawfish, Hobbseus yalobushensis, is described from the 
headwaters of the Yalobusha River, ultimately a tributary of the Mississippi 
River. This is the first record of the genus from the Mississippi drainage. The 
new species is most closely related to H. prominens (Hobbs) and H. petilus 
Fitzpatrick. It is distinguished from them by the relative lengths of the terminal 
elements of the first pleopod of the first form male; females of H. yalobushensis 
are the only members of the genus to have a deep, long trough in the anterior 


part of the annulus ventralis. 


In 1987, one of us (CAB) received a grant 
from the Mississippi Natural Heritage Pro- 
gram to determine the current status of the 
rare crawfish Procambarus (Pennides) lylei 
Fitzpatrick & Hobbs, 1971. During the 
study, several specimens of a crawfish as- 
signable to the genus Hobbseus were col- 
lected. These proved to represent an un- 
described species and provided the 
opportunity for the first published record of 
the genus from the Yalobusha River drain- 
age, and thus, from the Mississippi River 
basin. 


Hobbseus yalobushensis, new species 
Fig. 1 


Diagnosis. —Pigmented; eyes normal. 
Rostrum spatulate, without marginal spines; 
acumen reduced but usually obvious, not 
sharply delimited basally by strong rostral 
shoulders. Areola from 34.6 to 41.9% (av. 
42.0%) of total carapace length (43.5—51.0%, 
av. 49.5% of postorbital carapace length) 
and from 1.75 to 2.73 (av. 2.43) times lon- 
ger than wide; punctations widely scattered 
and poorly developed, 2 to 5 across nar- 
rowest part. Cervical spines absent. Post- 
orbital ridges strong, terminating cephali- 
cally in rounded knob or small tubercle. 


Branchiostegal spine obsolete; suborbital 
angle lacking. Antennal scale broadest distal 
to midlength. Dorsal surface of palm of che- 
liped studded with squamous tubercles. Is- 
chia of only third pereiopods of males with 
hooks; bosses lacking on all pereiopodal 
coxae, but small, sparsely setose, obliquely 
oriented eminence on caudomesial corner 
of third, and ventromesial margin of fifth 
with obvious tubercle bearing one or two 
long setae. First pleopods of males sym- 
metrical, apices reaching just beyond caudal 
margin of coxae of third pereiopods and 
hooded by dense mat of long setae origi- 
nating mostly from area of lateral margin 
of sharply arched sternites; terminating in 
two parts, rami subparallel with apex of each 
directed at angle of about 115° to main axis 
of appendage; mesial process only slightly 
longer than central projection and tapering 
from base to acute tip; central projection of 
first form male corneous, with obscure 
rounded eminence at proximomesial base, 
and with acute tip. Annulus ventralis mov- 
able, subovate in outline; deep, broad trough 
in cephalic third overhung through most of 
length of one side by prominent cepha- 
lolateral tubercle; sinus, originating in fun- 
dus located beneath caudalmost part of 
aforementioned overhang, moving trans- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


638 


ODS Lean Yt 


seeteet tote tatees seneces, 
wecesee ss Lye 


Ha 
ep casecesesesnsste® 


eye 


Petree e eA 


VOLUME 102, NUMBER 3 


versely past midline and then turning sharp- 
ly caudad following gently undulant path to 
or nearly to caudal margin. Hand of female 
sparsely hirsute with comparatively few 
punctations above and below. 

Holotypic male, Form I. —Cephalothorax 
(Fig. 1b, J) subovate, slightly compressed 
laterally, deeper than wide at level of cau- 
dodorsal margin of cervical groove (8.0 and 
7.5 mm, respectively). Abdomen longer than 
carapace (16.0 and 15.4 mm). Areola 2.43 
times longer than wide with 3 or 4 irregular 
rows of poorly delineated punctations, 3 
across narrowest part, constituting 36.4% of 
entire length of carapace (44.1% of postor- 
bital length). Rostrum slightly depressed an- 
teriorly, only weakly excavate dorsally, un- 
thickened elevated margins flanked mesially 
by deep punctations only near base; acumen 
slightly upturned, poorly set off from ros- 
trum and not reaching distal margin of pen- 
ultimate podomere of antennule. Subrostral 
ridge weak and barely visible in dorsal as- 
pect. Postorbital ridge strong, grooved dor- 
solaterally, terminating cephalically in 
rounded, unexpanded knob. Suborbital an- 
gle lacking. Branchiostegal spine obsolete. 
Cervical spine absent; very few deep punc- 
tations on carapace, deepest in vague row 
leading caudomesially from base of post- 
orbital ridge; few low squamous granula- 
tions in extreme cephalolateral part. 

Cephalic lobe of epistome (Fig. 1k) 
broadly subtriangular, distinctly convex with 
elevated margins, obtuse fovea in main 
body. Antennules of usual form with small 
spine near ventromesial margin of basal ar- 
ticle. Antennae extending caudally to mid- 
length of third abdominal tergum; antennal 
scale (Fig. 1h) 1.40 times longer than wide, 
widest distal to midlength, lateral part 


— 
Fig. 1. 


639 


thickened, terminating in strong acute spine 
and overreaching distal margin of ultimate 
podomere of antennal peduncle. 

Cephalic section of telson with single im- 
movable spine in each caudolateral corner 
and slightly movable spine just mesial to it. 

Chela (Fig. 17) somewhat depressed, 
slightly rotated mesially, subovate in cross 
section. Upper surface with comparatively 
prominent tubercles and only sparsely 
punctate, punctations deep only on fingers 
and in cluster of three proximal to base of 
immovable finger. Both fingers with median 
longitudinal ridge above and below, setif- 
erous punctations more common on dorsal 
flank of opposable margins. Opposable 
margin of immovable finger with three 
prominent tubercles in basal half, single row 
of minute denticles along distal half. Op- 
posable margin of dactyl with tubercle near 
midlength and second subequal tubercle near 
proximal one-tenth, both smaller than tu- 
bercles of immovable finger; single row of 
crowded minute denticles in distal half. Lat- 
eral margin of fixed finger with obtuse keel 
along most of its length and four tufts of 
setae in distal one-fourth; corresponding 
margin of palm only slightly undulant; inner 
margin of palm with single, nearly cristi- 
form row of 13 tubercles, flanked above by 
three tiny squamous tubercles and, in distal 
third, below by four. Low but broad squa- 
mous tubercle near base of dactyl. Outer 
margin of dactyl entire with row of sub- 
marginal, conspicuously setose punctations 
in distal half. 

Carpus of cheliped slightly longer than 
wide; dorsal surface with shallow nearly 
longitudinal furrow toward mesial margin 
and few scattered setiferous punctations, 
mostly in distal half; mesial margin with 


Hobbseus yalobushensis, all figures of holotype except d, e, morphotypic male; i, allotypic female. a, 


Mesial view of first pleopod; b, Lateral view of carapace; c, Lateral view of first pleopod; d, Mesial view of first 
pleopod; e, Lateral view of first pleopod; f Caudal view of tip of first pleopod; g, Ventral aspect of basal 
podomeres of pereiopods; 4, Antennal scale; i, Annulus ventralis; 7, Dorsal aspect of distal podomeres of cheliped 
(carpus rotated about 20° laterally to visualize stout mesial spine); k, Epistome; /, Dorsal view of carapace. 


640 


nine irregularly placed squamous to low spi- 
niform tubercles; lower submesial margin 
with strong, acute spine slightly distal to 
midlength; lower laterodistal corner with 
acute spine, lower mesiodistal corner lack- 
ing ornamentation. Merus with row of eight 
subacute spines along ventromesial margin 
and another of five along ventrolateral mar- 
gin, with row of four spiniform tubercles on 
dorsal margin, row terminating in two more 
small but stout subacute, adjacent spines. 
Ischium with three small spines along distal 
two-thirds of mesial margin. 

Hooks on ischia of third pereiopods only 
(Fig. lg); hooks strong, simple, slightly 
arched, and overhanging basioischial artic- 
ulation but opposing structure on basis lack- 
ing. Coxae of all pereiopods lacking bosses, 
but third with slightly globose expansion of 
caudomesial corner bearing irregular ar- 
rangement of from seven to nine long, coarse 
setae; fifth with typical ventromesial setose 
eminence and penile orifice. 

Sternum of third through fifth pereiopods 
deeply excavate and with dense tufts of setae 
arising from lateral margins, setae (with mi- 
nor contributions from proximal parts of 
coxae) obscuring distal half of pleopods 
when latter held parallel to body. 

First pleopods (Fig. la, c, f) as described 
in ‘“‘Diagnosis’’; central projection cor- 
neous. 

Allotypic female.—Except in secondary 
sexual characteristics, differing from holo- 
type in following respects: abdomen sub- 
equal in length to carapace; mesial margin 
of palm with row of 12 tubercles; opposable 
margin of fixed finger with proximalmost 
tubercle reduced to scarcely more than 
prominent undulation; mesial margin of 
carpus with 2 small tubercles and 1 small 
spine proximal to stout spine, no ornamen- 
tation distally; merus with only 3 small 
spines in ventrolateral row, row ending dis- 
tally in strong acute spine at corner; ven- 
tromesial row of 3 large, 1 small, followed 
by 3 large, 1 small, and distally, 4 large 
spines, dorsal surface with 2 spines on dis- 
talmost margin. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Annulus ventralis (Fig. 17) movable, sub- 
ovate in outline, highest (ventrally) just cau- 
dal to midlength; deep broad trough arising 
on cephalomedian margin and progressing 
obliquely to left of midpoint of annulus, 
high crescent-shaped eminence (or cephalo- 
lateral tubercle) overhanging lateral and 
caudal extremes of trough; sinus arising in 
deep fossa located in caudolateral corner of 
trough, progressing transversely to right of 
median line, there turning sharply caudad, 
following gently undulant path to just short 
of caudal margin. Postannular sclerite 
prominent, about half width of annulus, and 
not obscured by caudal margin of latter. 

Morphotypic male, Form IT.— Differing 
from holotype in following respects: areola 
with 4 punctations across narrowest part; 
proportions of chela more like allotype and 
inner margin of palm with row of 12 tu- 
bercles; mesial surface of carpus with 3 small 
spines proximal to stout spine, entire dor- 
sally; ventromesial margin of merus with 
row of 3 large, 1 small, and 3 large spines, 
lateral row represented by single quite 
prominent acute spine at base of distal 
fourth; dorsal surface of same podomere 
with 3 triangularly arranged stout spines near 
distal margin. 

Both terminal elements of first pleopod 
(Fig. 1d, e) noncorneous, less slender, sub- 
equal in length, and apices subacute. Hooks 
on ischia of third pereiopods conspicuous 
but markedly less developed, rounded at 
apex. Left first pleopod with incomplete, 
obscure suture delimiting basal tenth. 

Type locality.—The holotype and allo- 
type were collected from Topashaw Creek, 
3.8 airmi. (6.1 km) SW of Mantee (jct. of 
St. Rtes. 15 and 46), T21N, R11E, at the 
boundary of Secs. 2 and 11, Webster Coun- 
ty, Mississippi. Here the creek flows through 
agricultural land, but it is shaded by decid- 
uous trees on both banks. The banks have 
been severely eroded by inflow from two 
metal culverts draining the fields and lo- 
cated approximately 15 m upstream of the 
collection site. The creek was from 1 to 2 
m wide, and up to 0.7 m deep, with slow 


VOLUME 102, NUMBER 3 


flow. The animals were collected by dip net 
from the streambed of bare mid-phase Por- 
ters Creek clay. No plant material other than 
leaf litter was apparent in the shaded area 
where the specimens were taken. Other 
crawfish present were large numbers of an 
undescribed Orconectes species and Cam- 
barus (Depressicambarus) striatus Hay, 
1902. The morphotype was taken from Dry 
Creek, a Topashaw Creek tributary, 0.5 
roadmi. (0.8 km) W of Hohenlinden, T15S, 
RIE, NE/4 SW/4 Sec. 36, also in Webster 
County. 

Disposition of the types.—The holotype, 
allotype, and morphotype are in the collec- 
tions of the National Museum of Natural 
History, Smithsonian Institution (USNM 
219513, 219514, and 219515, respectively); 
the same museum also has three lots of 
paratypes (141,56 II, 2 2). Other paratypes 
are 1n the collections of the Mississippi Mu- 
seum of Natural Science, Jackson (MMNS; 
moles 15> 2 3 6 1mm., 2 2imm., 12 
unsexed imm.); of the Royal Ontario Mu- 
seum, Toronto (ROM; 1 61, 1¢IHI,12,7¢ 
imm., 1 2imm.); and of the Milwaukee Pub- 
lic Museum (MPM;; 1 ¢II, 1 2). 

Range and specimens examined. — 
Hobbseus yalobushensis has been collected 
on nine occasions from six localities, all in 
the headwaters of the Yalobusha River, 
tributary to the Yazoo River, tributary to 
the Mississippi River: Calhoun County. (1) 
Small unmapped tributary of Bear Creek, 
7.5 airmi. (12.2 km) SE of Calhoun City (jct. 
St. Rtes. 8 and 9), T22N, R1OE, center Sec. 
12, (MMNS; 1 2, 1 6 imm., 2 2 imm.), 13 
Feb 1987, C. Busack, M. Belk, and N. Hunt, 
colls.; Chickasaw County. (2) Topashaw 
Creek at St. Rte. 340, 4.5 roadmi. (7.3 km) 
W of St. Rte. 15, T15S, R2E, boundary of 
Secs. 20 and 21, (ROMIZ 13881;1¢1, 164 
II, 7¢imm., 1 2 imm.), 14 Mar 1988, M. 
Belk, coll.; (3) Topashaw Creek at St. Rte. 
340, 4.5 roadmi. (7.3 km) W of St. Rte. 15, 
T15S, R2E, boundary of Secs. 20 and 21, 
(MMNS; 2 ¢ I, 1 4 Il, 1 9, 1 6 imm., 1 2 
imm., 12 unsexed imm.), 26 Mar 1988, C. 
B. and M. B., colls.; (4) Topashaw Creek at 


641 


county road, 2.5 roadmi. (4.0 km) W of St. 
Ric wot Tios: RIE, Sec" 235"°Nw/4. 
(MMNS; 1 ¢I, 2 4 II, 1 2, 1 ¢imm.), 26 Mar 
1988, C. B. and M. B., colls. (5) roadside 
ditch, 3.4 roadmi. (5.5 km) NW of St. Rte. 
9 on St. Rte. 404, (tributary to Sabougla 
Creek), (USNM 218641; 161, 17 Apr 1967, 
C. Craig, coll.; (6) 3.4 roadmi. (5.5 km) N 
of St. Rte. 9 at Bellefontaine on St. Rte. 404, 
(USNM 207124; 1 6 II, 1 9), 14 May 1969, 
T. D. Thornhill, coll.; (7) 3.4 roadmi. (5.5 
km) NW of St. Rte. 9 at Bellefontaine on 
St. Rte. 404, (USNM 207112; 4 4 II, 1 9), 
14 May 1969, C. C., coll.; (8) Dry Creek, 
0.5 roadmi. (0.8 km) W of Hohenlinden, 
RIE, T15S, NE/4 SW/4 Sec. 36, (USNM 
219515; 1 6 ID (ROMIZ 13882; 1 9) 
(MMNS; 2 4 II, 2 2) (MPM IZ 1988-30; 1 
6 II, 1 2), 24 Apr 1987, M. B. and K. Bald- 
win, colls.; (9) type locality, (USNM 219513, 
219514; 1 6 I, 1 2) (MMNS; 1 ¢ DJ, 5 Jun 
1987, C. B. and N. Baldwin, colls. In ad- 
dition, two immature females collected by 
C. B., M. B., and C. Hill on 30 Jan 1987 
from Sabougla Creek, 1 roadmi. (1.6 km) 
NW of Bellefontaine, T21N, R9E, SW/4 
SE/4 Sec. 24, Webster County, are probably 
assignable to this species; this collection is 
at MMNS. 

Variations. — Most of the limits of vari- 
ation seen in this limited number of spec- 
imens is reflected in the description of the 
primary types, above. The second Form I 
male from the type locality has the apex of 
the rostrum broadly rounded and is without 
an acumen; perhaps it was broken early in 
life; also the mesial margin of the nght palm 
has arow of 15 tubercles. In one of the Form 
II males (12.5 mm carapace length) col- 
lected with the morphotype, the terminal 
elements of the first pleopod are adpressed 
throughout their length, and the central pro- 
jection is markedly the longer element; the 
pleopod also has a distinct juvenile suture 
proximally. The rostra of two immature 
specimens (1 4, 1 2) have sharply converging 
margins so that the acumen is acute and 
clearly delineated. 

Size.—The largest animal collected is a 


642 


Table 1.— Measurements (in mm) of types of Hobbs- 
eus yalobushensis. 


Morpho- 
Holotype Allotype type 
Carapace 
Total length 15.4 16.5 14.1 
Postorbital length 27) NZ, EIS) 
Width ILS) 8.0 7.0 
Height 8.0 8.6 7.4 
Areola 
Length 5.6 5.8 Sell 
Width 23 MJ 2.1 
Antennal scale 
Length 3.5 3.6 3.4 
Width 2.5 Dee) 2.5 
Rostrum 
Length 6.2 6.6 40) 
Width 4.2 4.1 3.9 
Chela 
Length, mesial 
margin palm 4.6 5.0* 4.0 
Width, palm 4.6 4.5* 4.0 
Length, lateral 
margin propodus 9.6 8.8* 7.4 
Length, dactyl S55) 5.4% 4.8 
Abdomen 
Length 16.0 16.5 15.4 
Width 6.5 ee 6.4 


* Left chela; right regenerated. 


female 21.8 mm in cephalothorax length. 
The largest first form male is 18.0 mm, and 
the smallest 14.8 mm. No ovigerous fe- 
males or females carrying young were col- 
lected. For measurements of the primary 
types see Table 1. 

Color notes.—The color patterns are ex- 
tremely variable, ranging from medium 
brown with frequent irregular black splotch- 
es dorsally, and becoming nearly concol- 
orous black laterally, to uniform medium 
tan, except for two poorly defined dorso- 
laterally placed dark lines extending from 
the mid-cephalic area to the cephalic margin 
of the telson. The abdomen is pale brown 
dorsally, except for the aforementioned lines 
which degenerate to a series of irregular 
splotches at the tergal margins. A dark red- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


dish brown line marks the boundary be- 
tween pleura and terga. The dorsal surface 
of the chela is basically orange brown, deep- 
ening to very dark brown on the fingers. The 
fingers are nearly white at the tips. The car- - 
pus, except for a light oblique groove, and 
the merus of the cheliped are dark brown 
dorsally. The pereiopods are dark brown 
dorsally and distally, fading to pink ven- 
trally and proximally. The undersides of the 
abdomen and cephalothorax are white; those 
of the proximal pereiopodal segments are 
white but deepen to pink at the ischia or 
meropodites. A varying intensity of pink 
coloration imparts a striped appearance to 
the pereiopods, especially in juveniles. 

Associates. —H. yalobushensis has been 
collected in association with Cambarus (De- 
pressicambarus) striatus, Procambarus 
(Ortmannicus) hayi (Faxon, 1884), Procam- 
barus (Pennides) vioscai Penn, 1946, and an 
undescribed species of Orconectes. 

Relationships. —The nearest relatives of 
Hobbseus yalobushensis are H. prominens 
(Hobbs, 1966) and H. petilus Fitzpatrick, 
1977. It can be distinguished from the for- 
mer by its less spatulate rostrum and that 
in H. prominens the terminal elements are 
slightly divergent. In both of the previously 
described species the mesial process is 
markedly longer than the central projection; 
the mesial process is, at best, only slightly 
longer in H. yalobushensis. Hobbseus yal- 
obushensis is unique in the genus in having 
such a well-developed, broad, long trough 
in the annulus ventralis and in having com- 
paratively heavy, non-uniform tuberculate 
ornamentation on the dorsal surface of the 
palm of the chela. 


Acknowledgments 


We thank all those individuals listed 
among the collectors, especially Marion 
Belk, for their assistance. We also thank 
Horton H. Hobbs, Jr., of the Smithsonian 
Institution, who compared specimens with 
the types of other species in the genus and 


VOLUME 102, NUMBER 3 


offered many useful suggestions concerning 
the manuscript. Finally, we are grateful to 
the Mississippi Natural Heritage Program 
for funding the project which led to the ser- 
endipitous discovery of this new species. 
This paper is contribution 13 of the Uni- 
versity of Mississippi Freshwater Biology 
Research Program. The senior author was 
supported, in part, by the Research Com- 
mittee of the University of South Alabama. 


Literature Cited 


Faxon, W. 1884. Descriptions of new species of Cam- 
barus to which is added a synonymical list of 
the known species of Cambarus and Astacus. — 
Proceedings of the American Academy of Arts 
and Sciences 20:107-158. 

Fitzpatrick, J. F., Jr. 1977. A new crawfish of the 
genus Hobbseus from northeast Mississippi, with 

notes on the origin of the genus (Decapoda, 

Cambaridae).— Proceedings of the Biological 

Society of Washington 90:367-374. 

, & H. H. Hobbs, Jr. 1971. A new crawfish of 

the Spiculifer Group of the genus Procambarus 


643 


(Decapoda, Astacidae) from central Mississip- 
pi.—Proceedings of the Biological Society of 
Washington 84:95-—102. 

Hay, W. P. 1902. Observations on the crustacean 
fauna of Nickajack Cave, Tennessee, and vicin- 
ity. — Proceedings of the United States National 
Museum 25(1292):417-439. 

Hobbs, H.H.,Jr. 1966. Anewcrayfish from Alabama 
with observations on the Cristatus Section of 
the genus Cambarus (Decapoda, Astacidae).— 
Proceedings of the Biological Society of Wash- 
ington 79:109-116. 

Penn, G. H., Jr. 1946. A new crawfish of the genus 
Procambarus from Louisiana.—Journal of the 
Washington Academy of Sciences 36:27-29. 


(JFF) 207 North Wacker Lane, Mobile, 
Alabama 36608; (CAB) Freshwater Biology 
Research Program, Department of Biology, 
University of Mississippi, University, Mis- 
sissippi 38677. Present address (CAB): State 
of Washington Department of Fisheries, 
Room 115, General Administration Build- 
ing, Olympia, Washington 98504. 


PROC. BIOL. SOC. WASH. 
102(3), 1989, pp. 644-645 


HIPPOLYTE ZOSTERICOLA 
(CRUSTACEA: DECAPODA) IN THE 
EASTERN PACIFIC 


Mary K. Wicksten 


Abstract.—Specimens of Hippolyte zostericola have been taken in western 
Colombia. These animals fall well within the range of variation found in spec- 
imens from the Atlantic and Caribbean regions, and can be distinguished readily 
from H. williamsi, the other common hippolytid shrimp of the tropical eastern 


Pacific. 


During a survey of caridean shrimp of 
western Colombia, Gabriel Ramos of the 
Universidad del Valle sent me 18 specimens 
of an unidentified hippolytid shrimp, taken 
at San Antonio, Municipio de Robles, Tu- 
maco (about 2°N, 79°W), on a mud bottom, 
3 Aug 1984, by Henry von Prohl. All but 
one were ovigerous females, no functional 
males were collected. The shrimp fall within 
the range of variation for Hippolyte zoster- 
icola (Smith), previously reported from 
Massachusetts, U.S.A. to Yucatan, the Ber- 
mudas, and south to Trinidad, Curacao, and 
Ceara, Brazil (Chace 1972, Williams 1984). 
The specimens from Colombia have been 
placed in the collections of the National 
Museum of Natural History, Smithsonian 
Institution, and the Allan Hancock Foun- 
dation, University of Southern California 
and Los Angeles County Museum of Nat- 
ural History. Another five specimens, ex- 
amined by Gabriel Ramos, have been added 
to the collections of the Universidad del 
Valle in Cali, Colombia (catalog number 
CRBMUV 84013). 

Hippolyte zostericola is variable: the ros- 
trum usually overreaches the antennular pe- 
duncle in adult females, and bears from one 
to three dorsal and from one to four ventral 
teeth. The basal article of the antennular 
peduncle is long and broad, without ter- 
minal spines. The rostrum is slightly shorter 
than the blade of the antennal scale. The 


third pereopod has a stout dactyl armed with 
three large, terminal spines grading into a 
series of smaller spines on the flexor margin. 
There is only one spine on the carpus and 
merus of the third pereopod. 

Chace (1972) and Williams (1984) re- 
marked on the similarity between H. Zos- 
tericola and H. pleuracanthus (Stimpson), 
reported from Connecticut to North Caro- 
lina. The latter species has a shorter ros- 
trum, not overreaching the antennular pe- 
duncle, with from one to three dorsal and 
from one to three ventral teeth. Chace (Fig. 
48) shows H. pleuracanthus as having a ros- 
trum terminating in a sharp point, with two 
ventral subterminal teeth close to the tip, 
while H. zostericola is shown as having a 
rostrum with a nearly bifurcate tip and two 
ventral teeth well removed from the tip. H. 
pleuracanthus is shown having four spines 
on the merus of the third pereopod. Spec- 
imens of H. zostericola from the Gulf of 
Mexico tend to have the longest rostrum 
within the species, while some from Mas- 
sachusetts have a rostrum not exceeding the 
antennular peduncle. 

The specimens from western Colombia 
vary considerably. All but two have a ros- 
trum barely exceeding the antennular pe- 
duncle, with two or three dorsal and two 
ventral teeth and a bifurcate tip. In two, the 
rostrum is shorter than the antennular pe- 
duncle, barely exceeding the cornea of the 


VOLUME 102, NUMBER 3 


eye. One has one dorsal and one ventral 
tooth, while the other has two dorsal teeth 
and a single apical point, not a bifurcate tip. 
As in H. zostericola, the basal segment of 
the antennular peduncle lacks spines, and 
the merus of the third pereopod bears one 
spine. The dactyls are similar in both species. 

I compared the specimens from Colom- 
bia with individuals of H. zostericola from 
Redfish Bay, Texas. In the latter specimens, 
the rostrum exceeded the antennular pe- 
duncle, and had two dorsal and three ven- 
tral teeth with a bifurcate tip. Except that 
the rostrum was proportionally longer, the 
specimens were similar. 

One cannot rule out the possibility that 
the specimens from Colombia, isolated from 
populations in the western Atlantic and Ca- 
ribbean, belong to a distinct species. Gem- 
inate pairs of species living on opposite sides 
of the Panamic land mass are known among 
other decapods, such as the spider crabs Pe- 
lia pacifica (Pacific) and P. mutica (Atlantic 
and Caribbean) and many others (Garth 
1958). Although the specimens from the 
eastern Pacific are indistinguishable from 
H. zostericola on the basis of morphology, 
there may be genetic, behavioral or ecolog- 
ical differences that warrant their future des- 
ignation as a separate species. 

One other species of Hippolyte, H. wil- 
liamsi Schmitt, can be common in the trop- 


645 


ical eastern Pacific. It is readily distinguish- 
able from H. zostericola by the presence of 
three spines on the basal segment of the 
antennular peduncle. Mature females are 
about 25 mm in total length, while the larg- 
est adult H. zostericola from Colombia is 
10.7 mm long. Williams (1984) gives a total 
length of 15.5 mm for females of H. zos- 
tericola. In H. williamsi, the rostrum of the 
female greatly exceeds the antennular pe- 
duncle. The two species may differ in hab- 
itat; H. zostericola lives among sea grasses 
and in sheltered bays, while H. williamsi can 
be common in tidepools and rocky areas 
with algae. 


Literature Cited 


Chace, F. A., Jr. 1972. The shrimps of the Smith- 
sonian-Bredin Caribbean expeditions with a 
summary of the West Indian shallow-water 
species (Crustacea: Decapoda: Natantia).— 
Smithsonian Contributions to Zoology 98:1—179. 

Garth, J. S. 1958. Brachyura of the Pacific coast of 
America: Oxyrhyncha.— Allan Hancock Pacific 
Expeditions 21(1):1—499. 

Williams, A. B. 1984. Shrimps, lobsters, and crabs 
of the Atlantic coast of the eastern United States, 
Maine to Florida. Washington D.C., Smithson- 
ian Institution Press, 550 pp. 


Department of Biology, Texas A&M Uni- 
versity, College Station, Texas 77843. 


PROC. BIOL. SOC. WASH. 
102(3), 1989, pp. 646-650 


CHACEON RAMOSAE, A NEW 
DEEP-WATER CRAB FROM BRAZIL 
(CRUSTACEA: DECAPODA: GERYONIDAE) 


Raymond B. Manning, Marcos Siqueira Tavares, and 
Elaine Figueiredo Albuquerque 


Abstract. — Chaceon ramosae, a species with depressed dactyli on the walking 
legs previously identified with C. quinquedens (Smith, 1879), is described as 


new, based upon material from Brazil. 


Almost no attempt has been made to study 
the deep-water fauna off the Brazilian coast 
since the Challenger Expedition (1873- 
1876). Collections made by subsequent 
oceanographic expeditions such as those 
carried out aboard the Albatross in 1887 and 
the Calypso (1961-1962) were largely re- 
stricted to areas of the Brazilian continental 
shelf. Information on the Brazilian deep- 
water decapod crustaceans is scarce and is 
largely confined to studies by Miers (1886), 
Bate (1888), Henderson (1888) (all based on 
the Challenger collections), Moreira (1901), 
and Rathbun (1937). 

Knowledge of the deep-water species of 
Brazil has been broadened as a result of the 
cruise in 1987 of the Marion Dufresne, an 
oceanographic ship of the TAAF (Terres 
Australes et Antarctique Francaises). The 
cruise was conducted off southeastern Brazil 
(Fig. 1) as part of a formal agreement be- 
tween the Muséum National d’Histoire Na- 
turelle, Paris, and the Universidade Santa 
Ursula, Rio de Janeiro. Among the mate- 
rials collected were four large specimens of 
the deep-sea crab genus Chaceon which 
proved to represent an undescribed species. 
That species is named below. 

The holotype has been deposited in the 
Museu Nacional, Rio de Janeiro (MNRJ). 
Other specimens, all paratypes, are in the 
collections of the Museu de Zoologia, Uni- 
versidade de Sao Paulo (MZUSP), the Mu- 
seum National d’Histoire Naturelle, Paris 


(MNHN), and the National Museum of 
Natural History, Smithsonian Institution, 
Washington (USNM). 

The following abbreviations are used be- 
low: cb, carapace width (including lateral 
spines); cl, carapace length, along midline; 
fm, fathoms; m, meters; mm, millimeters. 


Chaceon ramosae, new species 
Figs. 2-3 


Geryon quinquedens. — Rathbun, 1937:270, 
271 [part, specimen from Brazil only].— 
Scelzo & Valentini, 1974:561 [part, spec- 
imens from Brazil only]. [Not Geryon 
quinquedens Smith, 1879.] 


Previous records. — Brazil: 24°17'S, 42°48’ 
30” W, 671 fm (1228 m) (Rathbun 1937).— 
24°28'S, 43°43'W, 800 m, and 25°13’S, 
44°33'W, 1200 m (Scelzo & Valentini 1974). 

Material. — Brazil: 19°38’S, 38°43’W, 960 
m, sta. 55 CB 95, 30 May 1987: 1 male 
(MZUSP 9363).—21°31'S, 40°07'W, 750- 
785 m, sta. 4 CP 7, 10 May 1987: 1 male 
(holotype, MNRJ-MD-1381).—23°46'S, 
42°09'W, 592-610 m, sta. 64 CB 105, 
2 Jun 1987: 2 males (MNRJ-MD-1382; 
MNHN).—24°17’S, 42°48'30"W, 671 fm 
(1228 m), Albatross sta. 2763, 31 Dec 1887: 
1 male (USNM 22072). 

Description. —A large Chaceon, cl to 143 
mm, cb to 158 mm in adults, with 5 an- 
terolateral teeth on the carapace and dor- 
soventrally depressed dactyli on the walking 


VOLUME 102, NUMBER 3 


SOUTH 
AMERICA 


AFRICA 


_ TROPIC OF CAPRICORN 


Fig 3) 


legs. Carapace 1.1 to 1.2 times broader than 
long. Median pair of frontal teeth separated 
by U-shaped sinus. Distance between sub- 
median frontal teeth less than distance be- 
tween them and lateral frontal teeth. Sec- 
ond, third, and fourth anterolateral teeth 
obsolete in adults, second and fourth small- 
est of all; distance from first to third tooth 
less than that from third to fifth tooth. Car- 
apace with distinct raised ridge mesial to 
fifth anterolateral tooth, carapace surface 
finely granular, especially posterolaterally, 
smooth only at hepatic regions. Suborbital 
tooth short and broad in adults, not ex- 
tending to level of lateral frontal teeth. Che- 
liped merus with sharp tooth subdistally, 
lacking distal tooth or angled lobe; carpus 
roughened dorsally, usually with irregular, 
curved granular ridge extending from mid- 
dle of proximal margin to inner spine, an- 
terior margin of carpus with at most an an- 
gled lobe but no spine; propodus with at 
most distal angled projection dorsally. Meri 
of walking legs with at most indistinct distal, 
dorsal tooth. Dactyli of walking legs de- 
pressed, height at midlength less than width. 


Area sampled by the Marion Dufresne in 1987. 


Fifth leg: merus usually less than 5.0 times 
(range 4.64.9 in adults, 6.4 in juvenile male) 
times longer than high, length 0.65 to 0.66 
cb in adult; carpus with line of sharp gran- 
ules dorsally; propodus length 4.1 to 4.3 
times height in adult, 5.1 times height in 
juvenile, longer than dactylus. 

Size.—Males only known, cl 28 to 127 
mm, cb 36 to 146 mm in material exam- 
ined. Rathbun’s specimen is the smallest 
specimen reported here. Scelzo & Valentini 
(1974) studied two males with cl 133 to 143 
mm, cb 146 to 158 mm. 

Color. —The specimens taken by the 
Marion Dufresne were cream colored in life. 
Scelzo & Valentini (1974) reported that the 
color of their two specimens was “‘cremo- 
so.” 

Depth range. —Our specimens were taken 
in depths between 601 (592-610) and 1228 
meters, with all records from depths in ex- 
cess of 600 meters. Scelzo & Valentini (1974) 
reported two specimens from 800 m and 
1200 m. The smallest specimen studied, a 
male with cl 28 mm, came from 1228 me- 
ters. 


648 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


AE 


Fig. 2. Chaceon ramosae, male paratype, cl 107 mm, sta. 64 CB 105: a, Dorsal view; b, Carapace. 


Remarks. —This species resembles C. 
quinquedens (Smith, 1879) in having de- 
pressed dactyli on the walking legs, but dif- 
fers in numerous features: the carapace is 
much more granular posterolaterally, the 
suborbital tooth is less developed, the car- 


pus of the chela completely lacks an outer 
spine in adults, the propodus of the chela 
lacks a distal dorsal spine, the meri of the 
walking legs lack a distinct distal dorsal 
spine, and the propodus of the fifth leg is 
less than 5 times as long as high. 


VOLUME 102, NUMBER 3 


Ol 


oe /d 


In 


Fig. 3. Chaceon ramosae, male paratype, cl 107 
mm, sta. 64 CB 105: a, Suborbital margin; b, Dactylus 
of fifth leg, posterior view; c, Dactylus of fifth leg, dorsal 
view. 


Chaceon maritae (Manning & Holthuis, 
1981), from West Africa, also has depressed 
dactyli on the walking legs, but differs from 
C. ramosae in several features: the carapace 
granulation is much coarser, the suborbital 
spine is smaller, and the walking legs are 
shorter. The merus and propodus of the fifth 
leg are about 4 times as long as high. 

The anterolateral spines of the carapace 
are more distinct in the smallest specimen; 
in very large specimens the second and 
fourth teeth are scarcely discernible, and the 
third tooth is very low, an obtuse lobe. The 
merus of the fifth leg is more than 6 times 


649 


longer than high only in the smallest spec- 
imen, cl 28 mm. 

Scelzo & Valentini (1974) reported two 
specimens taken off Brazil in addition to 
numerous specimens taken off Uruguay and 
Argentina. Their material from Brazil dif- 
fered from the other specimens they iden- 
tified as Geryon quinquedens in being cream 
rather than reddish in color, and in the 
length/height ratios of the propodi of the 
walking legs, 4.3 to 4.5, which is similar to 
our findings. Their material from south of 
Brazil has been referred to C. notialis Man- 
ning & Holthuis (1989), who also trans- 
ferred all large geryonids with 5 anterolat- 
eral spines on the carapace to the genus 
Chaceon Manning & Holthuis (1989). 

Etymology. —We take pleasure in naming 
this species for Jeanete Maron Ramos, Uni- 
versidade Santa Ursula, one of the organ- 
izers of the cruise of the Marion Dufresne 
off the Brazilian coast. 


Acknowledgments 


We thank Alain Guille, Muséum Natio- 
nal d’Histoire Naturelle, Paris, and Janete 
Maron Ramos, Universidade Santa Ursula, 
for making available the material of C. ra- 
mosae. Enrique Macpherson of the Instituto 
de Ciencias del Mar in Barcelona informed 
one of us (RBM) of the Chaceon taken by 
the French-Brazilian expedition, and he is 
responsible for our establishing contact. The 
photographs were taken by Roy Kropp and 
Marilyn Schotte. Lilly King Manning made 
the line drawings, and prepared all of the 
figures for publication. 


Literature Cited 


Bate, C. S. 1888. Report on the Crustacea Macrura 
collected by H.M.S. Challenger during the years 
1873-76.—Report on the Scientific Results of 
the Voyage of H.M.S. Challenger during the years 
1873-76, Zoology 24:xc + 942 pp., 157 pls. 

Henderson, J. R. 1888. Report on the Anomura col- 
lected by H.M.S. Challenger during the years 
1873-76.—Report on the Scientific Results of 


650 


the Voyage of H.M.S. Challenger during the years 
1873-76, Zoology 27:xi + 221 pp., 21 pls. 

Manning, R. B., & L. B. Holthuis. 1989. Two new 
genera and nine new species of geryonid crabs 
(Crustacea, Decapoda, Geryonidae).— Proceed- 
ings of the Biological Society of Washington 
102(1):50-77. 

Miers, E. J. 1886. Report on the Brachyura collected 
by H.M.S. Challenger during the years 1873- 
76.—Report on the Scientific Results of the 
Voyage of H.M.S. Challenger during the years 
1873-76, Zoology 17:xli + 362 pp., 29 pls. 

Moreira, C. 1901. Crustaceos do Brasil. Contribui- 
¢Oes para o conhecimento da fauna brasileira. — 
Archivos do Museu Nacional, Rio de Janeiro 
ji Shai ea sii 


Rathbun, M. J. 1937. The oxystomatous and allied 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


crabs of America.— United States National Mu- 
seum Bulletin 166:vi + 278 pp. 

Scelzo, M. A., & A. Valentini. 1974. Presencia de 
Geryon quinquedens Smith en aguas del Oceano 
Atlantico sudoccidental (Decapoda, Brachyura, 
Geryonidae).— Physis, Buenos Aires (A)33(87): 
557-567. 


(RBM) Department of Invertebrate Zo- 
ology, National Museum of Natural His- 
tory, Smithsonian Institution, Washington, 
DC 20560; (MST and CFA) Departamento 
de Biologia Animal, Universidade Santa 
Ursula, Rua Fernando Ferrari, 75, 22131 
Rio de Janeiro, RJ, Brazil. 


PROC. BIOL. SOC. WASH. 
102(3), 1989, pp. 651-697 


ON THE CRAYFISH GENUS FALLICAMBARUS 
(DECAPODA: CAMBARIDAE) IN ARKANSAS, WITH 
NOTES ON THE FODIENS COMPLEX AND 
DESCRIPTIONS OF TWO NEW SPECIES 


Horton H. Hobbs, Jr. and Henry W. Robison 


Abstract.—The genus Fallicambarus, embracing 16 species of burrowing 
crayfishes, is represented in Arkansas by eight of them, five of which are mem- 
bers of the nominate subgenus and three assigned to the subgenus Creaserinus. 
A key is provided for the identification of the 16, the combined ranges of which 
extend from Ontario to southwestern Texas and southwestern Georgia, and 
from South Carolina to Maryland. Fallicambarus (F.) petilicarpus, a close rel- 
ative of F. (F.) dissitus, is described from Union County, Arkansas, and F. (C.) 
gilpini, related to F. (C.) caesius, from Jefferson County, Arkansas. Data are 
presented for placing Fallicambarus (C.) uhleri (Faxon) and F. (C.) hedgpethi 
(Hobbs) in the synonymy of F. (C.) fodiens (Cottle). For each of the species 
occurring in Arkansas, as a complete a synonymy as possible is offered along 
with a diagnosis of the species; also included are a statement of the range, a 
list of localities (also noted on spot maps) and specimens examined, color notes, 
size ranges of adults, and life history and ecological notes. 


Among the more interesting and least 
known of the crayfishes of Arkansas are eight 
species assigned to the genus Fallicambarus. 
Like all of their congeners, those occurring 
in Arkansas are seldom found in permanent 
bodies of water, and, as adults, only after 
rains or during floods do they frequent tem- 
porary pools or runoff. Throughout most of 
their lives, these crayfishes inhabit burrows 
that are excavated in areas where, for most 
of the year, the water table does not drop 
more than a meter or so beneath the surface 
(in Arkansas, such areas exposed to the sun 
often may be recognized by the presence of 
hydrophilic sedges). Turrets, sometimes in 
the form of slender chimneys, but more often 
irregular mounds of earthen pellets of a size 
proportional to that of the crayfish, mark 
the scattered burrows which, in many areas, 
may be seen in, or adjacent to, roadside 
ditches. Less frequently, clusters of mounds, 
or even entire fields studded with turrets 


signify the presence of large colonies of these 
crayfishes. 

Although seldom seen during daylight 
hours, on warm evenings individuals, with 
chelipeds extended, may be observed at the 
mouths of their lairs, sometimes perched at 
the top of a chimney, or, when the humidity 
is sufficiently high, and particularly follow- 
ing a Shower, walking over the ground. Dur- 
ing the early spring and after rains, evidence 
of the presence of a crayfish in its domocile 
is provided by damp-to-wet pellets of soil 
recently deposited at openings to the bur- 
rows. Such signs of recent excavation are 
seen most frequently early in the day, before 
the pellets have become dry, but, if the 
weather is overcast, recent deposits may be 
found at almost any hour. 

At one time or another, there exist for 
most burrows constructed by members of 
the genus Fallicambarus more than one 
opening at the surface that lead into the sim- 


652 


ple or complex system of galleries. The sim- 
plest burrows constructed by these crayfish- 
es in Arkansas are those of F. (C.) fodiens 
(Cottle, 1863). Often they consist of a single 
subvertical tunnel with a slight enlargement 
at the fundus, but more often they are made 
more complex by the addition of a side pas- 
sage leading to or toward the surface, and 
on rare occasions, when a second adult 
shares the domicile with an ovigerous fe- 
male or one bearing young, there may exist 
a secondary gallery leading downward. Thus 
in its simplest form, the burrow consists of 
passages that are subvertical and are dis- 
posed in the forms of an “I,”’ “Y”’ (some- 
times inverted), or “X.’’ The more complex 
patterns usually occur in areas where during 
much of the year the water table lies very 
near or almost at the surface. The galleries 
of these burrows are disposed largely sub- 
parallel to the surface, and except for one 
or two that descend, presumably to the 
maximum depth to which the water table 
drops, lie relatively close to the surface. 
Usually there are at least two exits, one of 
them nearly always topped by a turret that 
is Sometimes open, but during dry weather 
may not only be closed but also the passage 
leading to it is often “back-filled.”’ If the 
burrow is that of an ovigerous female or one 
carrying young, all openings to the surface 
are frequently plugged. During wet seasons, 
all of the galleries may be water-filled, but 
at other times water may be found only at 
the bottom of the one or two deep passages. 
A few burrows have been excavated in which 
no standing water was present. 

The range of the genus Fallicambarus is 
a discontinuous one in which the larger seg- 
ment extends from Ontario southward to 
Arkansas County, Texas, and eastward to 
the Apalachicola River basin of southwest- 
ern Georgia. The more eastern segment en- 
compasses the Coastal Plain and lower 
Piedmont provinces from New Jersey to 
South Carolina. In Arkansas, the genus is 
confined to the Coastal Plain and foothills 
of the Ozark and Ouachita mountains. There 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


the nominate subgenus, which is not known 
to occur east of the Ouachita River basin in 
Arkansas and Louisiana or west of the Bra- 
zos Basin in Texas, is restricted to the 
Ouachita and Red river basins. The much 
more widespread subgenus Creaserinus, 
however, occurs in all of the major drainage 
basins in the state, overlapping much of the 
range of the subgenus Fallicambarus. 

Of the eight species present in Arkansas, 
five are members of the nominate subgenus: 
Fallicambarus (F.) dissitus (Penn, 1955), F. 
(F.) harpi Hobbs & Robison (1985), F. (F.) 
Jeanae Hobbs (1973), F. (F.) petilicarpus, 
new species, and F. (F.) strawni (Reimer, 
1966); and three are assigned to the sub- 
genus Creaserinus: F. (C.) caesius Hobbs 
(1975), F. (C.) fodiens Cottle (1863), and F. 
(C.) gilpini, new species. Little is known 
about any of them except F. (C.) fodiens, 
and no investigation has involved a com- 
parison of representatives of populations 
throughout the range of the species. More- 
Over, considerations of its relationship to F. 
(C.) uhleri and F. (C.) hedgpethi have been 
only cursory, the principal reason being a 
lack of specimens from much of the sus- 
pected or assumed ranges of the three 
species. In many, if not most, of the collec- 
tions that were available, there were no first 
form males. The latter problem persists to 
date, for in less than half of the collections 
that we have examined is even one such 
specimen present. As a result we cannot be 
confident that the one or two males in a 
collection exhibit secondary sexual char- 
acters, the most useful that we have en- 
countered, that are typical of the local pop- 
ulation. In defense of the conclusions offered, 
however, we point out that such features 
appear to be remarkably uniform in the first 
form males in most localities represented in 
our material by two or more such males. 
This observation has prompted the ques- 
tion as to why so few of these males have 
been collected. 

The following represents primarily an ac- 
count of our knowledge of the genus Falli- 


VOLUME 102, NUMBER 3 


cambarus in Arkansas, although we have 
attempted to summarize all available data 
on those species that range beyond the state 
boundaries. Indeed, the diversity noted in 
the populations of F. (F.) fodiens occurring 
in Arkansas provoked us into a study of 
representatives of the species throughout its 
range, one of the largest of any crayfish in 
North America. In presenting the synony- 
mies, diagnoses, bibliographic references, 
and summaries of published biological data 
for each of the species occurring in the state, 
we have included all of which we are aware, 
and the following key encompasses all of 
the species that have been assigned to the 
genus. 

Abbreviations used herein are as follows: 
AMNH, American Museum of Natural 
History, New York; ANSP, Academy of 
Natural Sciences of Philadelphia; cl, cara- 
pace length; CM, Carnegie Museum, Pitts- 
burgh; pol, postorbital carapace length; TU, 
Tulane University, New Orleans; USNM, 
National Museum of Natural History, 
Smithsonian Institution. 


Genus Fallicambarus Hobbs (1969a) 


Diagnosis.—“‘Adults with rostrum de- 
void of marginal spines. Mesial margin of 
palm of chela with row of fewer than 12 
tubercles, opposable margin of dactyl usu- 
ally with prominent excision. Areola linear 
or obliterated [along part of its length]. An- 
tennal scale more than twice as long as broad. 
First pleopods of first form male symmet- 
rical and terminating in two or three distinct 
parts (mesial process, central projection and, 
occasionally, cephalic process; caudal ele- 
ment always absent) bent caudally or cau- 
domesially at angle of 90 degrees or more 
to main shaft or forming broad arc; central 
projection corneous, blade-like or tapering 
(but flattened laterally) and [frequently] 
lacking ... subapical notch; non-corneous 
mesial process never bulbiform but often 
appearing twisted and usually with emi- 
nence on cephalic (morphological) border 


653 


slightly distal to base; cephalic process, when 
present, small, at least partially corneous, 
situated mesially at base of mesial process, 
and directed caudally or caudodistally” 
(slightly modified from Hobbs 1973:463). 
Type species: Cambarus strawni Reimer, 
1966:11. Gender: masculine. 

This taxon was proposed by Hobbs 
(1969a:111) to receive an assemblage of eight 
species that had been assigned formerly to 
the genus Cambarus. Subsequently, the di- 
agnosis of this assemblage was revised by 
him (1973:462), and speculations were of- 
fered on the interrelationships of the 11 
members recognized at the time. In point- 
ing out more formally the existence of two 
species groups among these crayfishes, he 
proposed the adoption of two subgenera: the 
nominate subgenus comprising five species, 
and the subgenus Creaserinus, six. In this 
summary the former subgenus is considered 
to consist of seven species, two of which 
have been described since 1973 and one 
herein; F. (F.) spectrum has been relegated 
to the synonymy of F. (F.) jeanae (Hobbs 
1989). The subgenus Creaserinus as pres- 
ently constituted consists of nine species, 
four of which were described since 1973 and 
one is added in this study. Evidence for 
placing Fallicambarus (C.) hedgpethi and F. 
(C.) uhleri in synonymy with F. (C.) fodiens 
is detailed herein. 


Key to Species of Genus Fallicambarus 
(Based on First Form Males) 


i First pleopod with proxi- 
momesial spur (Fig. 37) 
. (Subgenus Fallicambarus) . 2 


ie First pleopod lacking proxi- 
momesial spur (Fig. 11h) ... 
...(Subgenus Creaserinus) .. 8 
2: Mesial ramus of uropod with 


distinct distolateral spine (Fig. 

lc); abdomen not conspicu- 

ously narrower than thorax . 3 
on Mesial ramus of uropod lack- 

ing distinct distolateral spine 


Bi 


4(3’). 


ia 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


(Fig. 1d); abdomen conspicu- 
ously narrower than thorax . 5 
Mesial ramus of uropod with 
distomedian spine projecting 
beyond margin of ramus (Fig. 

lc); cheliped with sufflamen 
(Fig. 10z); central projection 
disposed proximocaudally, 
never overlapping that of cor- 
responding pleopod (Fig. 11h) 

SSE COST, Aes macneesel 
Mesial ramus of uropod with 
distomedian spine never pro- 
jecting beyond margin of ra- 
mus (Fig. 11d); cheliped with- 
out suffamen (Fig. 10a’); 
central projection disposed 
proximomesially, its distal ex- 
tremity frequently overlap- 
ping that of corresponding 
pleopod (Fig. 32) 
First pleopod with cephalic 
process (Fig. 35); length of car- 
pus of cheliped distinctly 
greater than width of palm of 
chela (Fig. 3h) ...... petilicarpus 
First pleopod lacking cephalic 
process (Fig. le); length of car- 
pus of cheliped subequal to or 
less than width of palm of che- 

Far ES SPE ES I dissitus 
First pleopod without cephal- 
ic process (Fig. lf) ....... jJeanae 
First pleopod with cephalic 


process (like Fig. lg) ....... 6 
Ischia of third and fourth pe- 
reiopods with hooks (like Fig. 
a1). Sr are devastator 
Ischia of third pereiopods only 
with hooks (like Fig. 1lm).. 7 


Cephalic process of first pleo- 
pod extending caudodistally 
(Fig. 1g); mesial ramus of uro- 
pod without distomedian 
spine (like Pigtidje: sols. : harpi 
Cephalic process of first pleo- 
pod closely paralleling basal 
part of mesial process (Fig. 
10y), apical part if free direct- 


8(1’). 


8". 


9(8). 


oF 


10(8’). 


10’. 


11(10). 


ale: 


12(11’). 


joe 


13(10’). 


ed caudally; mesial ramus of 
uropod with distomedian pre- 
Marginal spine ......... strawni 
Ventral surface of merus of 
cheliped with one row of tu- 


bercles (Figvilig) 1G Aa 9 
Ventral surface of merus of 
cheliped with two rows of tu- 
bercles (Fie ii p)t 2a ae 10 


Mesial surface of dactyl of 
chela with tubercles in basal 
half (Fig. 17); mesial ramus of 
uropod lacking distolateral 
Spineo ts a2 Rete caesius 
Mesial surface of dactyl of 
chela lacking tubercles (Fig. 
110); mesial ramus of uropod 
with distolateral spine (Fig. 
Ld) Oe ei Oe a gilpini 
Mesial surface of palm of che- 
la of second pereiopod not 
bearded (Fig. 1a); shaft of first 
pleopod strongly reflexed (Fig. 
LA) OPE ee 11 
Mesial surface of palm of che- 
la of second pereiopod beard- 
ed (Fig. 15); shaft of first pleo- 
pod straight or only slightly 
curved (Figs. li, k, 1 8, 9, 
10GS=x Oe ee 13 
First pleopod with central 
projection not conspicuously 
long and slender (Fig. 1h) byersi 
First pleopod with central 
projection conspicuously long 
and slender (Fig. lj) ...... 12 
Antennal scale tapering dis- 
tally to strong distolateral 
spine (Fig. 17); boss on coxa 
of fourth pereiopod ridgelike 
(Figs OE) Note ae gordoni 
Antennal scale rounded to 
subtruncate distally, lacking 
distolateral spine (Fig. 17); 
boss on coxa of fourth pereio- 
pod bulbiform (10b’)..... burrisi 
First pleopod with central 
projection straight and trun- 
cate distally (Fig. 17) .... hortoni 


VOLUME 102, NUMBER 3 
13% First pleopod with central 
projection arched, and taper- 

ing at distal extremity or bear- 

ing subapical notch (Figs. 1k, 
sO. NOG ya Pe 14 
Antennal scale with some- 
what distinct distal and mesial 
margins, their junction sub- 
angular (Fig. lo); abdomen 
broadly joined to cephalotho- 

PED. Re RT ae RR, Maen Sore a oe fodiens 
Antennal scale with distome- 

sial margin strongly inclined 
abdomen narrowly joined to 
BONA ss of a ra. so eS 15 
First pleopod with mesial pro- 

cess conspicuously overreach- 

ing central projection (Fig. 1k) 
RE OA Re danielae 
First pleopod with mesial pro- 

cess overreaching central pro- 

jection little if any (Fig. 1/) . 

oryktes 


14(13’). 


14’. 


15(14’). 


15%, 


ee © © © © © © © © © © ee ee we ee ew ee 


Subgenus Fallicambarus Hobbs (1973) 


Diagnosis.— First pleopod of male with 
proximomesial spur and sometimes with 
cephalic process. Cheliped without suffla- 
men, except in F. macneesei, chela with tu- 
bercles scattered over most of dorsal sur- 
face, lateral margin of palm and basal part 
of fixed finger rounded, more often subser- 
rate or serrate, never smoothly costate. Sec- 
ond pereiopod of male with mesial face of 
chela and carpus lacking dense mats of plu- 
mose setae. Coxa of fourth pereiopod usu- 
ally with conspicuously large boss. 


Fallicambarus (Fallicambarus) dissitus 
(Penn) 
Figs. le, 2 


Cambarus dissitus (Penn, 1955:73-80, figs. 
1-13 [Types: holotype, allotype, and 
morphotype, USNM 98125, 98126, 
98127; paratypes, ANSP, AMNH, CM, 
TU, USNM. Type locality: three miles 
east of Choudrant, Lincoln Parish, Lou- 


655 


istana-| 1959:5,.629. 11. 15,.16,, P7 pigs. 
25, 48, 66, 79.—Penn & Marlow, 1959: 
202.— Hobbs, 1959:896; 1962:274; 1967: 
13.—Black, 1967:173, 178.— Fitzpatrick 
& Payne, 1968:14.—Hobbs III, 1969:19, 
2A tab 2: 

Fallicambarus dissitus. —Hobbs, 1969a:111, 
124, 173, fig. 20d; 1972:15, 99, figs. 81c, 
82d; 1974b:23, 102, fig. 88.— Feinberg, 
1971:26.—Hart & Hart, 1974:26, 30.— 
Bouchard, 1978:432; 1980:432.—Bou- 
chard & Robison, 1981:26, 29. 

Fallicambarus (Fallicambarus) dissitus.— 
Hobbs, 1973:463, 477-479, fig. 4.—Bou- 
chard & Robison, 1981:28.— Fitzpatrick, 
1983:167.—Hobbs & Robison, 1985: 
1035. 


Diagnosis.—Cheliped without sufflamen; 
ventral surface of merus with mesial and 
lateral rows of tubercles; length of carpus 
less than, or subsequal to, width of palm of 
chela. Chela with lateral margin at least 
weakly serrate, dorsal surface with scattered 
tubercles in lateral half, ventrolateral sur- 
face lacking arched row of prominent setif- 
erous punctations; opposable margin of 
dactyl with distinct excision in basal half, 
mesial margin with longitudinal row of tu- 
bercles along at least proximal fourth. Me- 
sial surface of palm of chela of second pe- 
reiopod lacking conspicuous tufts of 
plumose setae. First pleopod with proxi- 
momesial spur, lacking cephalic process; 
central projection strongly arched, inclined 
laterally at base, its distal part directed prox- 
imomesially and often crossing that of cor- 
responding pleopod. Hooks on ischia of third 
and fourth pereiopods. Boss on coxa of 
fourth pereiopod moderately strong and 
compressed. Mesial ramus of uropod with 
distolateral and distomedian spine, latter 
premarginal. Telson divided and with 
spine/s flanking anterolateral flank of su- 
ture. 

Range and specimens examined.—This 
crayfish is known from only a few localities 
in the Red and Ouachita river basins of 
southern Arkansas (Columbia County) and 


656 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig... 


Same of F. (C.) fodiens; c, Dorsal view of telson and uropods of F. (F.) macneesei; d, Same from F. (C.) jeanae; 
e-l, Mesial view of first left pleopod; e, F. (F.) dissitus; f, F. (F.) jeanae; g, F. (F.) harpi; h, F. (C.) byersi; i, F. 
(C.) hortoni; j, F. (C.) burrisi; k, F. (C.) danielae, 1, F. (C.) oryktes; m—o, Antennal scale: m, F. (C.) burrisi; n, F. 
(C.) gordoni; 0, F. (C.) fodiens; p, Ventrolateral view of merus of cheliped of F. (C.) byersi; q, Same, F. (C.) 


gilpini; r, Dorsal view of chela of F. (C.) caesius; s—u, Mesial view of chela of F. (C.) fodiens; s, from Ohio; t, 
from Maryland; u, from Florida. 


Characters used in key: a, Distal podomeres of second pereiopod of Fallicambarus (C.) byersi; b, 


VOLUME 102, NUMBER 3 


== 
> 


. 
bs 


ai 
~ 


St 
ad 


oh Mie o ol s 

ae eee 
jal 

Wis Vie 


i} 


Ss 
1 


657 


Fig. 2. Distribution of Fallicambarus (F.) dissitus (encircled stars), F. (F.) jeanae (encircled dots), F. (F.) 
petilicarpus (triangle) and F. (F.) strawni (dots) in Arkansas. (Some localities listed in text too close to others to 


be shown.) 


north central Louisiana (Caldwell, Grant, 
and Lincoln parishes). It has been found in 
the following localities in Arkansas. Colum- 
bia County: (1) Southern Arkansas Univer- 
sity Campus at Magnolia, 1 61, 23 Apr 1985, 
HWR. (2) 1 mi SW of Macedonia on St Rte 
160, specimens not available, R. W. Bou- 
chard and HWR. These localities lie in the 
Dorcheat Bayou-Red River basin. 

Size.— The largest specimen that we have 
examined is a first form male from Lincoln 
Parish, Louisiana, having a cl of 32.7 (pol 
29.1) mm. Corresponding lengths of the 
smallest first form males are 23.1 (20.2) mm. 

Life history notes. — First form males have 
been collected in February, March, April, 
and May. Neither ovigerous females nor 
ones carrying young have been reported. 


Ecological notes.—Specimens were col- 
lected in Grant Parish, Louisiana (8.2 miles, 
13.2 km, south of the Winn Parish line on 
U.S. Highway 167), from moderately com- 
plex burrows, approximately two feet deep, 
in a roadside seepage area; this locality is 
in a rolling terrain where the soil is a sandy 
clay. The specimen from locality 1 was col- 
lected as it crawled across a lawn where there 
were chimneys 18 to 24 centimeters in 
height. 


Fallicambarus (Fallicambarus) harpi 
Hobbs & Robison 
Figs. lg, 5 


Fallicambarus (Fallicambarus) harpi Hobbs 
& Robison, 1985:1035-1041, fig. 1 


658 


[Types: holotype, allotype, and morpho- 
type, USNM 217946, 217947, 217948; 
paratypes, MHNP, RMHL, USNM. Type 
locality: see page area 0.2 mi (0.32 km) 
east of Glenwood, Pike County, Arkan- 
sas]. 


Diagnosis.—Cheliped without sufflamen; 
ventral surface of merus with mesial and 
lateral rows of tubercles; length of carpus 
less than, or subequal to, width of palm of 
chela. Chela with lateral margin strongly 
serrate, dorsal surface with scattered tuber- 
cles in lateral half, ventrolateral surface 
lacking arched row of prominent setiferous 
punctations; opposable margin of dactyl 
with distinct excision in basal half, mesial 
margin with longitudinal row of tubercles 
extending almost complete length of finger. 
Mesial surface of palm of chela of second 
pereiopod lacking conspicuous tufts of plu- 
mose setae. First pleopod with proxi- 
momesial spur and freely-projecting (not 
adnate to mesial process), caudodistally-di- 
rected cephalic process; central projection 
strongly arched, but not inclined laterally at 
base, and never crossing that of correspond- 
ing pleopod. Hooks on ischia of third pe- 
reiopods only. Boss on coxa of fourth pe- 
reiopod very strong and compressed. Mesial 
ramus of uropod lacking spines. Telson di- 
vided but lacking spines. 

Inasmuch as no information has been 
added to our knowledge of this crayfish since 
it was described, the following data have 
been extracted from Hobbs & Robison 
(1985). 

Range and specimens examined.—The 
two nearby localities in the Ouachita River 
basin cited here are the only ones known 
for the species. Pike County: (1) Type lo- 
cality, 36 61, 13 4II, 8 2, 1 j4, 16 Apr 1982, 
K. Dillard; 2 61, 4 2, 21 Apr 1982, KD. (2) 
0.3 mi (0.5 km) E of Glenwood on US Hwy 
70,661, 3 2, 18 Mar 1982, KD. 

Color notes.—‘‘Dorsum of cephalic re- 
gion of carapace, including rostrum very 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


dark, almost black, that of thoracic region 
greenish tan except for small black trian- 
gular spot in open anterior section of areola, 
narrow band bordering cervical groove, and 
another band adjacent to posterior margin 
of carapace; hepatic and mandibular ad- 
ductor regions also black; orbital, antennal, 
mandibular, and branchiostegal regions 
greenish tan. Abdomen with broad median 
longitudinal, dark red stripe extending from 
second through fifth segment, becoming pale 
and disappearing before reaching caudal 
margin of sixth; lateral part of latter seg- 
ment, telson, and uropods greenish tan with 
dark markings; mesial ramus of uropods 
with black median longitudinal rib. Dorsal 
surface of cheliped dark green with black 
suffusion becoming intense mesially so that 
dorsodistal border of merus, mesial part of 
carpus, tubercular area of palm, all of dor- 
sum except tips of fingers and proximome- 
sial part of fixed finger almost black; tips of 
fingers and lateral and ventral surfaces of 
entire cheliped pinkish to yellowish cream. 
Remaining pereiopods cream with greenish 
suffusion marking dorsal parts of podo- 
meres from ischium through propodus, 
more intense on merus and propodus. Ster- 
nal elements and ventral surfaces of pereio- 
pods cream to pinkish cream” (Hobbs & 
Robison 1985:1039). 

Size.—‘“‘The largest specimen examined 
is a female having a carapace length of 39.6 
(postorbital carapace length 35.8) mm. The 
smallest and largest first form males have 
corresponding lengths of 29.0 (25.8) mm, 
and 35.4 (31.5) mm, respectively” (Hobbs 
& Robison 1985:1039-1040). 

Life history:notes. — First form males were 
obtained in March and April. Ovigerous fe- 
males or ones carrying young have not been 
observed. 

Ecological notes.—The type locality con- 
sists of a seep “located in a pasture . . . spec- 
imens were collected in the early evening as 
they crawled about in the wet grassy areas. 
Soil consisted of a sandy clay with some 


VOLUME 102, NUMBER 3 


organic material. Grasses and sedges were 
abundant....” 


Fallicambarus (Fallicambarus) jeanae 
Hobbs 
Figs. ld, f 2 


Fallicambarus (Fallicambarus) jeanae 
Hobbs, 1973:463-469, 477, 478, 480, figs. 
1, 4 [Types: holotype and allotype, USNM 
144672, 144673 (61, 2); paratypes USNM. 
Type locality: seepage area 1.8 mi (2.9 
km) E of Clark County line, Hot Spring 
County, Arkansas, on St Rte 84.].— Bou- 
chard, 1978:451; 1980:451.— Bouchard 
& Robison, 1981:28.—Flitzpatrick, 1983: 
167, 168.—Hobbs & Robison, 1985:1035, 
1040.— Hobbs, 1989. 

Fallicambarus (Fallicambarus) spectrum 
Hobbs, 1973:463, 469-478, 480, figs. 2, 
4 [Types: holotype and allotype, USNM 
144674, 144675 (41, 2); paratypes USNM. 
Type locality: 2 mi (3.2 km) E of Daisy, 
Pike County, Arkansas, on US Hwy 
70.].—Bouchard, 1978:451; 1980:451.— 
Bouchard & Robison, 1981:28.—Fitz- 
patrick, 1983:167, 168.—Hobbs & Ro- 
bison, 1985:1035. 

Fallicambarus jeanae.—Hobbs, 1976:550, 
fig. 1b, d, 1.— Bouchard & Robison, 1981: 
26.—Huner & Barr, 1981:50, fig 24b, d, 
]; 1984:45, fig. 24b, d, 1.—Robison & 
Smith, 1982:53 

Fallicambarus sp.— Hobbs, 1979:804. 

Fallicambarus spectrum.—Bouchard & 
Robison, 1981:26.—Robison & Smith, 
1982:53 


Diagnosis. —Cheliped without suflamen; 
ventral surface of merus with mesial and 
lateral rows of tubercles; length of carpus 
less than, or subequal to, width of chela. 
Chela with lateral margin strongly serrate, 
dorsal surface with scattered tubercles in 
lateral half, ventrolateral surface lacking 
arched row of prominent setiferous punc- 
tations; opposable margin of dactyl with 


659 


distinct excision in basal half, mesial margin 
with longitudinal row of tubercles extending 
almost complete length of finger. Mesial 
surface of palm of chela of second pereiopod 
lacking conspicuous tufts of plumose setae. 
First pleopod with proximomesial spur, 
lacking cephalic process; central projection 
moderately arched, not inclined laterally at 
base, its distal part disposed caudoproxi- 
mally, never crossing that of corresponding 
pleopod. Hooks on ischia of third pereio- 
pods only. Boss on coxa of fourth pereiopod 
very strong and compressed. Mesial ramus 
of uropod lacking distolateral spine; disto- 
median spine, if present, situated premar- 
ginally. Telson incised laterally but lacking 
spines 

Range and specimens examined;—This 
crayfish, which seems to be endemic to the 
upper Ouachita River basin in southwestern 
Arkansas, has been found in the following 
localities (those for which no first form males 
are listed should be confirmed). Clark 
County: (1) 1 mi (1.6 km) NE of Amity 
Center on St Rte 84, Sec 27NE, T5S, R23 W, 
DiGi Or ody 2.423) 2 1) Apr1973,"Ge B. 
Hobbs, J. E. Pugh, HHH; 1 ¢ II, 2 2, 2 jé, 
2 j2, 29 Apr 1976, M. Kearney, HHH; 1 ¢ 
II, 1 2, 1 j2, 14 Apr 1979, D. D. Koym, 
HWR; 1 ¢4 II, 3 2, 3 j4, 15 Apr 1982, HWR, 
DDK. (2) roadside ditch 7.4 mi (11.8 km) 
E of Amity Center on St Rte 84, 1 41, 1 Q, 
3 36, 30 Apr 1976, MK, HHH. (3) Richland 
Cemetery, 3 mi (4.8 km) NW of Gum 
Springs, 1 j¢, 1 j2, 24 Feb 1980, W. Laird. 
(4) Rest Haven Cemetery 4 mi (6.4 km) W 
of Arkadelphia on St Rte 8, 1 2, 1 jd, 1 j9, 
24 Sep 1977, E. Laird; 1 jé, 27 Dec 1979, 
EL; 3 2, 6 May 1980, EL; 2 9, 1 jé, 1 j2, 12 
Feb 1981; 2 6 II, 4 2, 636, 5 j2, 9 Mar 1981, 
EL, HWR; 1 2, 19 Mar 1981, EL. Hemp- 
stead County: (5) Blevins, Sec 24, T9S, 
R24W, 3 2, 1 j6, 29 Apr 1983, P. Lee. (6) 
Blevins, Sec 6, T10S, R24W, 1 jé, 1 j2, 10 
May 1983, B. Hill. (7) Blevins, Sec 35, T9S, 
R25W, 1 2, 16 May 1983, T. Chambers. 
Hot Spring County: (8) roadside ditch 1.8 


660 


mi (2.9 km) E of Clark Co line on St Rte 
84 (Type locality), 5 6 I, 2 2, 3 jé, 2 j2, 21 
Apr 1973, GBH, JEP, HHH. (9) roadside 
ditch 3.7 mi (5.9 km) E of jct of St Rtes 7 
and 84 on latter, 2 4 I, 1 j2, 30 Apr 1976, 
MK, HHH. (10) roadside ditch 5.2 mi (8.3 
km) E of Clark Co line on St Rte 84, 1 j9, 
30 Apr 1976, MK, HHH. (11) roadside ditch 
4.5 mi (7.2 km) W of Marcus, 3 ¢4I, 1 4 II, 
2 2, 234, 5j2, 14 Apr 1979, HWR et al. (12) 
2.1 mi (3.3 km) E of Point Cedar, 2 2, 1 jé, 
1 j2, 13 Mar 1981, HWR; 1 41, 2 2, 3 jé, 
15 Apr 1982, HWR, DDK. Pike County: 
(13) roadside ditch 2 mi (3.2 km) E of Daisy 
on US Hwy 70, 1 46 I, 1 2, 21 Apr 1973, 
GBH, JEP, HHH. (14) roadside ditch 2.8 
mi (4.5 km) W of Amity Center on St Rte 
84, 2 6 Il, 3 2, 2 j4, 14 Apr 1979, HWR et 
al. (15) roadside ditch 8.2 mi (13.1 km) W 
of Amity Center on St Rte 84, 1 j2, 14 Apr 
1979, HWR et al. 

Remarks.— With the acquisition of con- 
siderably more material than was available 
to him when Fallicambarus (F.) spectrum 
was described, Hobbs (1989) concluded that 
except by the color pattern this crayfish can- 
not be distinguished from F. (F.) jeanae. 
The pattern of the former, which is de- 
scribed below, has been observed only in 
specimens from the type locality. He there- 
fore treated spectrum as a color morph of 
F. (F.) jeanae. 

Color notes.— Fallicambarus (F.) jeanae 
exhibits two distinctive color morphs; that 
occurring in specimens from the type lo- 
cality was described by Hobbs (1973:468) 
as follows: ““Dominant color of carapace pale 
mauve; rostral margins, postorbital ridges, 
and paired subtriangular areas caudal to 
postorbital ridges dark brown; latter joining 
along cervical groove and in caudal gastric 
area where almost black; caudal margin of 
carapace dark brown. First abdominal ter- 
gum dark brown, remaining ones pale yel- 
lowish tan with paired dorsolateral cream 
splotches and each edged caudally with ver- 
milion. Telson and uropods cream with pale 
tan suffusion basally. Antennule and anten- 
na with dark yellowish-brown peduncles; 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


flagella with each article yellowish tan ba- 
sally and dark brown distally; lateral margin 
of antennal scale almost black. Cheliped 
mostly yellowish tan dorsally with dark 
bluish-brown tubercles and bluish-brown 
suffusion on dorsal margin of merus, dor- 
somesial surface of carpus, and dorsomesial 
part of dactyl. Ventral surface of cheliped 
yellowish cream. Remaining pereiopods 
with coloration similar to that of cheliped 
but lacking dark brown tubercles.” 

The color of the morph that was described 
under the name of Fallicambarus (F.) spec- 
trum (Hobbs, 1973:472-473) was recorded 
as follows: “Dominant color of carapace pale 
mauve gray; rostral margins and postorbital 
ridges almost black; paired subtriangular 
reddish-brown markings extending caudal- 
ly from caudal extremity of postorbital 
ridges, their basal portions coalescing in me- 
dian posterior gastric region, slightly over- 
lapping cervical groove, and filling cephalic 
triangular vestige of areola. Hepatic region 
with pale reddish-tan suffusion; caudal mar- 
gin of carapace edged with black. [Dorso- 
lateral parts of branchiostegites provided 
with conspicuous dark bluish purple (some 
almost black) spots.] First abdominal ter- 
gum reddish brown with succeeding terga 
progressively fading to uniformly reddish- 
tan telson and uropods; tergum also fading 
laterally, and pleuron concolorous with tel- 
son. Peduncle of antennule and antenna dark 
mauve gray, flagella with each article pale 
tan proximally, becoming dark gray distally; 
antennal scale with lateral thickened por- 
tion almost black. Chelipeds grayish blue 
dorsally with dark blue tubercles; ventral 
surface cream; remaining pereiopods simi- 
lar to cheliped but lacking dark blue tuber- 
cles.”’ 

Size. — The largest specimen examined by 
us is a female from Clark County having a 
cl of 40.6 (pol, 35.5) mm. The smallest and 
largest first form males have corresponding 
lengths of 30.0 (26.4) mm and 35.7 (31.7) 
mm, respectively. 

Life history notes.—All of the first form 
males that we have examined were collected 


VOLUME 102, NUMBER 3 


in April. Neither ovigerous females nor ones 
carrying young have been observed. A first 
form male and female were found occupy- 
ing a single burrow at locality 1 on 21 Apr 
1973. 

Ecological notes.—The largest colony of 
this crayfish that we have observed was in 
the type locality. Persistent rains for at least 
several days while one of us (HHH) was in 
the area had brought the water table 
throughout the seep to the surface, and the 
mounds marking the entrances to the cray- 
fish burrows were considerably eroded. They 
did not appear to have been neatly con- 
structed, and the pellets of which subse- 
quently observed turrets were constructed 
by F. (F.) jeanae seemed to have been rather 
haphazardly affixed to the rim, many having 
rolled down the side of the crude, vaguely 
cone like structures. The soil was primarily 
a sandy clay with gravel and pockets of or- 
ganic material. The burrows were rather 
shallow (most of the horizontal passages 
coursed at depths of 10 to 30 cm; the one 
or two deeper passages no doubt penetrated 
the soil for 70 cm to one meter), highly 
branching, and had two or three openings 
to the surface. In the immediate vicinity, 
grasses and sedges were growing in the water- 
logged soil, and nearby were trees belonging 
to the genera Acer, Juniperus, and Pinus. In 
April of 1973, the soil was so lacking in 
firmness that no tool was needed to aid the 
hand in searching for the crayfish in their 
burrows. In localities 1 and 13, where the 
“spectrum color morph”’ was found, the soil 
was better drained (a small, clear brook 
flowed through locality 1), the burrows were 
deeper (but extended no more than a meter 
beneath the surface), and the horizontal gal- 
leries were restricted to a smaller surface 
area. 


Fallicambarus (Fallicambarus) 
petilicarpus, new species 
Figs. 2, 3, 4a 


Diagnosis.—Eyes small but pigmented 
and with faceted cornea. Cheliped without 


661 


suffamen; ventral surface of merus with 
mesial and lateral rows of tubercles; length 
of carpus greater than width of palm of che- 
la. Chela with lateral margin weakly serrate, 
dorsal surface with scattered tubercles in 
lateral half, ventrolateral surface lacking 
arched row of prominent setiferous punc- 
tations; opposable margin of dactyl lacking 
pronounced excision in basal half, mesial 
margin with row of tubercles along at least 
proximal third. Mesial surface of palm of 
chela of second pereiopod lacking conspic- 
uous tufts of plumose setae. First pleopod 
with proximomesial spur, and free, caudally 
projecting cephalic process; central projec- 
tion strongly arched, inclined laterally at base 
but projecting mesially and crossing that of 
corresponding pleopod distally. Hooks on 
ischia of third and fourth pereiopods. Boss 
on coxa of fourth pereiopod moderately 
strong and compressed. Mesial ramus of 
uropod with distolateral spine, distomedian 
spine, if present, situated premarginally. 
Telson divided and with spine/s on an- 
terolateral flank of suture. 

Holotypic male, form I.—Body suboval, 
weakly compressed laterally (Figs. 3a, /, 4a). 
Abdomen distinctly narrower than thorax 
(9.6 and 14.5 mm). Greatest width of car- 
apace near midlength of areola where sub- 
equal to height (14.5 and 14.2 mm). Areola 
linear over most of length and comprising 
37.4 percent of entire length of carapace 
(42.9 percent of postorbital carapace length). 
Rostrum with convergent, slender margins 
contracting anteriorly, setting off base of in- 
distinctly delimited acumen, apex of which 
corneous, upturned, and slightly overreach- 
ing midlength of penultimate podomere of 
antennular peduncle. Dorsal surface of ros- 
trum strongly concave, with submarginal 
rows of setiferous punctations and scattered 
ones between. Subrostral ridges rather weak 
but evident in dorsal aspect to base of acu- 
men. Postorbital ridges also weak but some- 
what swollen posteriorly. Branchiostegal and 
cervical spines absent. Suborbital angle ab- 
sent. Carapace punctate dorsally and very 
weakly and sparsely tuberculate laterally; 


662 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 3. Fallicambarus (F.) petilicarpus holotype except d, h, j from allotype, and e, f from morphotype): a, 
Lateral view of carapace; b, c, e, Mesial view of first pleopod, cp cephalic process; d, Annulus ventralis and 
associated sclerites; f, g, Lateral view of first pleopod; h, 0, Distal podomeres of cheliped; i, Caudal view of first 
pleopods; j, Antennal scale; k, Epistome; 1, Dorsal view of carapace; m, Dorsal view of caudal part of abdomen; 
n, Basal podomeres of third, fourth, and fifth pereiopods. (cp, cephalic process; ps, proximomesial spur.) 


VOLUME 102, NUMBER 3 


extreme anteroventral branchiostegal re- 
gion inflated, with irregular group of tuber- 
cles, seven or eight forming row on ventral 
flank of cervical groove; fewer tubercles 
present on dorsal flank of groove. 

Abdomen (Fig. 4a) little shorter than car- 
apace (28.0 and 30.5 mm); pleura moder- 
ately deep and broadly rounded ventrally, 
only sixth with angular caudoventral mar- 
gin; pleuron of first abdominal segment 
clearly overlapped by that of second. Telson 
(Fig. 3m) distinctly divided, and caudolat- 
eral angles of anterior section with two pairs 
of spines, more mesial pair movable. Prox- 
imal podomere of uropod with both lobes 
bearing distal spine, spine on more mesial 
lobe much stronger than that on lateral lobe; 
mesial ramus with well developed distolat- 
eral spine and smaller premarginal disto- 
median spine. 

Cephalomedian lobe of epistome (Fig. 3) 
broadly subtriangular with well defined 
cephalomedian prominence extending 
cephalodorsally; cephalolateral margins 
rather smooth; main body of epistome with 
cephalomedian area depressed but lacking 
distinct fovea. Ventral surface of proximal 
podomere of antennule with median spine 
slightly distal to midlength. Antennal pe- 
duncle without spines except on lateral sur- 
face of basis at proximal base of antennal 
scale, flagellum reaching second abdominal 
tergum. Both antennal scales of holotype 
with distal extremities missing, but distinct- 
ly reduced: blade little broader than thick- 
ened lateral portion and apex probably 
reaching only slightly beyond midlength of 
penultimate podomere of antennule (see Fig. 
3j, 1). Mandible essentially like that of Fal- 
licambarus (F.) devastator Hobbs & White- 
man (1987: fig. 11). Ventral surface of is- 
chium of third maxilliped with lateral row 
of short, plumose setae, and mesial half 
studded with clusters of long, stiff setae. 

Right chela (Fig. 30) approximately 2.4 
times as long as broad, not strongly de- 
pressed; width of palm slightly more than 
1.2 times length of mesial margin, latter 
bearing row of 7 tubercles subtended dor- 
solaterally by row of six (left with 6 and 7, 


663 


respectively); dorsal surface of palm and 
basal part of fingers studded with squamous 
tubercles; those along lateral margin form- 
ing subserrate row extending from near 
proximal extremity to about midlength of 
fixed finger; ventral surfaces of palm, basal 
third of fixed finger, and mesial part of prox- 
imal fourth of dactyl tuberculate; more dis- 
toventral part of fingers with punctations, 
in part, flanking weak median ridges; prom- 
inent spiniform tubercle present on oblique 
distoventral ridge of palm; ventrolateral 
surface lacking curved row of long setae. 
Opposable margin of fixed finger with row 
of 7 tubercles (fourth from base largest) in 
proximal three-fifths and another (corre- 
sponding to usually more ventrally-placed 
tubercle on the chelae of most cambarids) 
at base of distal third; minute denticles pres- 
ent between distal 4 tubercles, and, except 
at base of distalmost tubercle, continuing to 
corneous tip of finger. Opposable margin of 
dactyl with row of 5 tubercles (third from 
base largest; this tubercle marking end of 
sharp proximal excavation in dacty] of other 
members of Fallicambarus) in proximal 
three-fourths; mesial margin of dactyl with 
tubercles forming subserrate row. Dorsal 
surface of both fingers with well defined me- 
dian longitudinal ridge. 

Carpus of cheliped conspicuously slender 
and long, almost 1.6 times as long as broad, 
and longer than either width or mesial mar- 
gin of palm of chela. Dorsal surface sparsely 
punctate and bearing poorly delimited and 
very shallow, oblique, longitudinal furrow; 
mesial surface with only one prominent 
subspiniform tubercle, cluster proximo- 
ventral to it consisting of much smaller ones; 
except for ventrodistal extremity bearing 
usual 2 moderately large tubercles, ventral 
and lateral surfaces punctate. Merus with 
single, dorsal, slightly curved row of tuber- 
cles, increasing in size distally; lateral sur- 
face punctate, and mesial one with polished 
and granular areas; ventral surface with both 
mesial and lateral rows of 15 tubercles joined 
by oblique row of 3. Mesioventral margin 
of basioischial podomere with only 2 tu- 
bercles corresponding to row in other cray- 


664 


fishes; compound podomere otherwise with 
scattered punctations. Chela of second pe- 
reiopod with marginal row of setae on palm, 
and carpus with dorsal row of long setae; 
mesial surface of carpus and propodus lack- 
ing tufts of plumose setae. 

Ischia of third and fourth pereiopods (Fig. 
3n) with simple hooks, neither of which 
overreaching basioischial articulation and 
neither opposed by tubercle on correspond- 
ing basis. Coxa of fourth pereiopod with 
prominent, compressed caudomesial boss 
disposed somewhat in longitudinal axis of 
body; mesial and lateral surfaces of boss 
with setiferous punctations. Coxa of fifth 
pereiopod with small tuberculiform boss 
extending ventrally from caudomesial angle 
of podomere; ventral membrane setiferous. 

First pleopods (Fig. 30, c, g, i) reaching 
coxae of third pereiopods, carried deeply in 
sternum, and largely concealed by setae ex- 
tending from ventral margin of sternum and 
from coxae of third and fourth pereiopods. 
Proximomesial spur well developed. Shaft 
of appendage only slightly inclined caudal- 
ly; plumose setae arising from mesial sur- 
face of shaft forming feathery plume hiding 
all or part of each of three terminal ele- 
ments: mesial process, most proximal of 
three, non-corneous, somewhat tapering but 
with rounded apical region, disposed at 
slightly more than right angle to shaft of 
appendage; cephalic process smallest, cor- 
neous, subtriangular, situated between ce- 
phalic process and central projection, and 
directed caudally; and central projection 
most conspicuous of three, consisting of 
long, tapering, bladelike structure reflexed 
through arc of at least 150 degrees. 

Allotypic female.— Differing from holo- 
type other than in secondary sexual char- 
acteristics as follows: acumen even less dis- 
tinctly delimited basally; subrostral ridges 
evident in dorsal aspect for no more than 
one-fifth distance from caudal margin of or- 
bit to base of acumen; rudiment of bran- 
chiostegal spine present; about same num- 
ber of tubercles (7 or 8) on both dorsal and 
ventral flanks of anteroventral segment of 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


cervical groove; spine on lateral lobe of 
proximal podomere of uropod rudimen- 
tary; spine on basis of antenna tuberculi- 
form; flagellum of antenna extending cau- 
dally over no more than three-fourths length 
of areola; chela (Fig. 3h) proportionally 
shorter and broader; mesial row of tubercles 
on palm of chela subtended dorsally by row 
of only 5 tubercles; opposable margin of 
fixed finger with row of 5 tubercles, third 
from base largest; opposable margin of dac- 
tyl with row of 4 tubercles, second from base 
largest; ventral surface of merus of cheliped 
with mesial and lateral rows of 14 tubercles; 
mesioventral margin of basioischial podo- 
mere with 3 tubercles (left chela regener- 
ated). 

Annulus ventralis (Fig. 3d) firmly fused 
to sternum cephalically, approximately twice 
as broad as long, and strongly asymmetrical. 
Cephalic and cephalomedian areas de- 
pressed, latter distinctly excavate; dextral 
side of excavation elevated in massive 
prominence along mesial margin of which 
C-shaped sinus marking junction of prom- 
inent transverse ridge and prominence; no 
clearly defined sulcus evident, and fossa 
hidden. Postannular sclerite about 1.7 times 
as broad as long and less than half as wide 
as annulus; lateral margins weakly converg- 
ing toward broadly rounded anterior ex- 
tremity; caudal margin irregularly trans- 
verse. First pleopods present but not 
reaching anterior to postannular sclerite. 

Morphotypic male, form ITI.—Differing 
from holotype as follows: Apex of rostrum 
reaching base of ultimate podomere of an- 
tennule; right branchiostegal spine repre- 
sented by very small tubercle; proximal 
podomere of uropod lacking spine on both 
lobes; antennal peduncle lacking spine on 
basis, flagellum reaching first abdominal 
tergum; (as in holotype, distolateral part of 
both antennal scales broken); chela approx- 
imately 2.2 times as long as broad; left chela 
with only 6 tubercles in mesialmost row on 
palm; merus of left cheliped with lateral row 
of only 12 tubercles; mesioventral margin 
of basioischial podomere with row of 3 or 


VOLUME 102, NUMBER 3 


4 tubercles; hooks on ischia of third and 
fourth pereiopods and boss on coxa of latter 
all clearly defined but weaker than those in 
holotype. 

First pleopods (Fig. 3e, f) reaching coxae 
of third pereiopods, symmetrical, and with 
well defined proximomesial spur; markedly 
similar in form to pleopod of first form male, 
but lacking cephalic process, and base of 
central projection not nearly so distinctly 
delimited from mesial process. 

Color notes.—Basic coloration olive- 
brown to tan. Carapace dark olive-brown; 
rostrum and posterior gastric area very dark; 
lateral areas slightly paler. First abdominal 
tergum dark olive-brown, second through 
fifth paler olive-tan and with narrow pink- 
ish tan arc on posteromedian margins; sixth 
tergum, telson, and uropods dark olive. An- 
tennules and antennae with peduncles olive 
on brown; flagella reddish brown. Chelipeds 
with basal three podomeres and proximal 
part of merus pinkish cream, latter suffused 
dorsally and laterally with olive, becoming 
dark olive distally; dorsal tubercles on mer- 
us green with white tips. Dorsal and lateral 
surfaces of carpus bright olive to forest green 
on brown, tubercles green and largest ones 
tipped with cream. Chela olive-brown dor- 
sally; distal ridge on palm suffused with 
green, and green on dorsal flank of oppos- 
able borders of both fingers; palm and fixed 
finger fading ventrolaterally to pinkish or- 
ange, ventral surface of all podomeres of 
cheliped pinkish to lavender cream. Re- 
maining pereiopods with olive suffusing dis- 
tal part of merus, carpus, and, except that 
of second pereiopod, proximal part of mer- 
us, Otherwise pale pinkish cream. 

Size.—The largest specimen examined is 
a female having a cl of 31.8 (pol 27.6) mm. 
The smaller of the two known first form 
males, the holotype, has corresponding 
lengths of 30.5 and 26.6 mm, respectively. 
Neither ovigerous females nor ones carrying 
young are available for determining mea- 
surements. 

Type locality. — Roadside seepage 0.2 mile 
east of the Columbia County line on State 


665 


Route 57, Union County, Arkansas (T16S, 
R18W, Sec 21). A field of young planted 
trees (Pinus) was adjacent to the seep and 
ditch which supported a moderate growth 
of sedges and grasses. Specimens were col- 
lected from relatively shallow, but complex 
burrows which, constructed in a sandy loam 
and topped by chimneys from 8 to 10 cm 
high, descended to depths of approximately 
0.5 m. Some of them penetrated tangled, 
dense root mats of grasses growing in and 
on the banks of the ditch. No other crayfish 
was found in the immediate area. 

Disposition of types.—The holotype, al- 
lotype, and morphotype (USNM 219507, 
219508, and 219509, respectively) are de- 
posited in the National Museum of Natural 
History, Smithsonian Institution, as are the 
paratypes consisting of 1 ¢I, 1 é II, 32, and 
9 juveniles. 

Range and specimens examined.—All of 
the specimens available were collected at 
the type locality by the second author on 30 
Apr 1982 (1 61, 1 4 Il, 2 2, and 3 juv) and 
28 Mar 1988 (1 41, 2 4 II, 2 2, 6 juv). 

Variations.— Among the adult speci- 
mens, the areola constitutes from 35.2 to 
38.5 percent of the total length of the car- 
apace, and from 39.8 to 43.6 percent of the 
postorbital carapace length. In one small 
male with a carapace length of 21.5 mm, 
the corresponding ratios are 34.0 and 39.0 
percent, respectively. Most of the variations 
noted fall within the range of those noted 
in the descriptions of the primary types. The 
greatest range of differences, barring regen- 
erated appendages, occurs in the numbers 
of tubercles on the chelipeds: the opposable 
margin of the fixed finger may have from 4 
to 8 tubercles and that of the dactyl 4 to 7; 
the ventromesial row on the merus ranges 
from 14 to 16 and the mesiolateral, from 
12 to 15. In the female the annulus ventralis 
occurs in mirrored images of that in the 
allotype, and the cephalic margin may or 
may not be firmly fused to the sternum im- 
mediately anterior to it. All of the females 
have a linear series of 3 to 5 long setae closely 
associated with the distal part of the ven- 


666 


Table 1.—Measurements (mm) of Fallicambarus (F.) 
petilicarpus. 


Morpho- 
Holotype Allotype type 
Carapace: 
Entire length 30.5 pA is 29.5 
Postorbital length 26.6 25.8 26.6 
Width 14.5 E59 13.0 
Length 14.2 b3:3 12.6 
Areola: 
Width 0 0 0 
Length 11.4 | 10.2 
Rostrum: 
Width 4.4 4.2 4.6 
Length Sef 4.5 4.8 
Right chela: 
Length, palm 
mesial margin a5 4.6 73 
Palm width 10.9 8.0 8.9 
Length, lateral 
margin 24.5 16.1 19.4 
Dactyl length 14.6 10.5 je 
Abdomen: 
Width 9.6 10.1 9.0 
Length 28.0 21 26.0 


trolateral row of tubercles on the merus; such 
seem not to be present in the available males. 
(See Table. 1 for other morphometric dif- 
ferences.) 

Life history notes.—The only collections 
of this crayfish available are two lots col- 
lected at the type locality in March and April. 
A first form male was obtained in both sam- 
ples. Ovigerous females and ones carrying 
young have not been observed. 

Ecological notes.—See ““Type locality.” 

Relationships. — Fallicambarus (F.) peti- 
licarpus is more closely allied to F. (F.) dis- 
situs than to any other member of the genus. 
The unusual first pleopods of the males of 
the two species with caudomesially dis- 
posed, crossing (at least sometimes) central 
projections, are so nearly alike that only af- 
ter a detailed re-examination of specimens 
of the former was the distinctive feature, 
the presence of a small, but well-defined ce- 
phalic process, observed. Subsequent com- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


parisons of the two revealed, among the 
many similarities, two other readily ob- 
served features that set F. (F.) petilicarpus 
apart: the very long slender carpus of the 
first cheliped and the unique absence in Fal- 
licambarus of a distinct concavity on the 
opposable margin at the base of the dactyl 
of the chela. The absence of this concavity, 
a feature the presence of which has served 
in keys until now to distinguish members 
of the genus Fallicambarus from Cambarus, 
necessitated the slight modifications in the 
generic diagnosis included herein. 

Etymology.—The name describes the 
slender carpus of the cheliped: Petilus (L. = 
slender) + carpus (L. = wrist). 


Fallicambarus (Fallicambarus) strawni 
(Reimer) 
Figs. 2, 10y 


Cambarus strawni Reimer, 1966:9, 11-14, 
figs. 9-18 [Types: holotype, allotype, and 
morphotype, USNM 116675, 116676, 
116677 (61, 2, IL); paratypes, R. D. Rei- 
mer. Type locality: small marshy area in 
the Saline River basin, 2.7 mi (4.3 km) 
north of Dierks, Howard Co., Arkan- 
sas.].— Hobbs, 1967:12; 1968:K16, fig. 32; 
1969a:111.—Black, 1967:173, 178.— 
Bouchard, 1972:61.—Hobbs III et al., 
1976:24. 

Fallicambarus strawni.— Hobbs, 1969a:103, 
111, 124, 151, 173, figs. 26, 13), 20: ae 
99, 147, fig. 81b; 1974b:24, 100, fig. 81; 
1976:551, fig. 1b, e.-—Bouchard & Ro- 
bison, 1981:26.— Robison & Smith, 1982: 
ao: 

Fallicambarus (Fallicambarus) strawni.— 
Hobbs, 1973:461-479, figs. 3c, h, 4.— 
Bouchard & Robison, 1981:28.—Fitz- 
patrick, 1983:168.—Hobbs & Robison, 
1985:1035. 


Diagnosis. —Cheliped without sufflamen; 
ventral surface of merus with mesial and 
lateral rows of tubercles; length of carpus 
less than, or subequal to, width of palm of 


VOLUME 102, NUMBER 3 


chela. Chela with lateral margin strongly 
serrate, dorsal surface with scattered tuber- 
cles in lateral half, ventrolateral surface 
lacking arched row of prominent setiferous 
punctations; opposable margin of dactyl 
with distinct excision in basal half, mesial 
margin with longitudinal row of tubercles 
extending almost complete length of finger. 
Mesial surface of palm of chela of second 
pereiopod lacking conspicuous tufts of plu- 
mose setae. First pleopod with proxi- 
momesial spur, and cephalic process adnate 
to mesial process, if free distally, then di- 
rected caudally, parallel to mesial process; 
central projection comparatively weakly 
arched, not inclined laterally at base, and 
disposed caudally, never crossing that of 
corresponding pleopod. Hooks on ischia of 
third pereiopods only. Boss on coxa of fourth 
pereiopod very strong and compressed. Me- 
sial ramus of uropod lacking distolateral 
spine, distomedian spine, if present, situ- 
ated premarginally. Telson entire, lacking 
spines. 

Range and specimens examined.—This 
crayfish seems to be endemic to southwest- 
ern Arkansas (although there is every reason 
to believe that it will be found in eastern 
Oklahoma) where it has been collected only 
in the Little and Saline watersheds in the 
Red River basin and in the Little Missouri 
watershed in the Ouachita River basin. We 
have examined all of the following material 
except that collected in locality 6. Howard 
County: (1) Type locality, 1 61, 1 ¢ II, 1 Q, 
77 ham 1963) KR. D. Reimer: 2 6], 16 Il, 1 
2, 1 jo, 1 j2, 21 Apr 1973, GBH, JEP, & 
HHH; 1 2, date ?, R. W. Bouchard. (2) road- 
side ditch 5.1 mi (8.2 km) W of Athens on 
St Rte 84, 1 6 I, 1 9, 29 Apr 1976, MK & 
HHH. (3) seep 1.8 mi (2.9 km) E of Polk 
Co line on St Rte 4, 3 2, 3 6 II (one ¢ later 
molted to form I), 29 Apr 1976, MK & 
HHH. (4) roadside ditch 1.8 mi (2.9 km) E 
of Sevier Co line on US Hwy 70, 3 jé, 6 j2, 
10 Apr 1982, HWR. (5) Nashville, 1 ¢ II, 
10 Apr 1986, L. Morris. (6) Reimer (1966: 
4) cited this crayfish from 4 mi W of Umpire 


667 


on St Rte 4. Pike County: (7) roadside ditch 
1.3 mi (2.1 km) E of Little Missouri River 
on St Rte 84, 1 2, 1 j¢, 29 Apr 1976, MK, 
HHH. (8) roadside ditch 0.9 mi (1.4 km) 
NE of Howard Co line on US Hwy 70, 1 6 
I, 1 9,21 Apr 1973, GBH, JEP, HHH. Sevier 
County: (9) seep and ditch 0.1 mi (16 km) 
NE ofjct of US Hwys 71-59 and 70 on latter, 
A 6A 446 I, 13 9,2; 36; 5 32,20 Apr.1973, 
GBH, JEP, HHH. (10) seep 8.3 mi (13.3 
km) E of jet of US Hwys 59-71 and 70 on 
latter, 1 j2, 28 Apr 1976, MK, HHH. (11) 
seep 0.2 mi (0.32 km) E of jct of US hwys 
71 and 59 on latter, 1 j6, 1 9, 26 Apr 1976, 
RWB. (12) 5.8 mi (9.3 km) E of jct on St 
Rtes 41 and 24 on latter, 1 j¢, 9 Apr 1982, 
HWR. (13) seep 5.0 mi (8.0 km) NE of ject 
of US Hwys 59-71 and 70, 1 61, 20 Apr 
1973, GBH, JEP, HHH. 

Color notes.—(Based primarily on first 
form male from locality 8.) Dominant color 
of carapace pinkish cream to purplish tan 
overlain by various shades of gray and ver- 
milion. Most of dorsum of cephalic region 
pinkish tan fading laterally to buff, often 
with lavender suffusion; rostrum and post- 
orbital ridges very dark gray margined in 
almost black; caudal gastric area and cer- 
vical groove pale to dark gray; cephalic tri- 
angle of areola dark gray to almost black, 
and linear part and caudal triangle of areola 
vermilion; paired pale gray longitudinal 
stripes flanking linear areola; remainder of 
branchiostegites, excluding dark bluish gray 
caudal margin, very pale cream tan. Tergum 
of first abdominal segment and cephalic part 
of that of second dark bluish red, otherwise 
yellowish tan with vermilion splotches dor- 
sally, fading caudally, and all pleura lighter 
tan ventrally. Telson largely translucent but 
with vermilion to brick red splotches ce- 
phalically, laterally, and along caudal mar- 
gin; uropod similarly translucent, but pe- 
duncle reddish tan, lateral ramus with 
reddish splotches lateral to median rib and 
Over entire distal section, and mesial ramus 
with red pigment largely restricted to me- 
dian rib and distal third. Chelipeds basically 


668 


tan but with dense reticulations of slate blue 
on dorsum of distal half of merus, that of 
carpus, and most of that of chela; lateral 
surface of palm pinkish cream. Exposed 
parts of peduncles of antennule and antenna 
mostly gray, and flagella with each article 
buff proximally, becoming dark gray dis- 
tally. Lateral margin of antennal scale very 
dark gray. Remaining pereiopods similar to 
cheliped, although with more red and less 
blue pigment on fifth. Ventral surface of 
body and pereiopods cream, latter with blue 
pigment toward distal ends of merus and 
carpus. 

In first form male from locality 2 (How- 
ard County), dorsal cephalic region darker 
orange tan, and with dark brownish-gray 
area extending across posteromedian gastric 
region abutting cervical groove; thoracic re- 
gion more apricot-colored dorsally, fading 
to cream tan ventrally; anterior triangle of 
areola almost black, branchiocardiac suture 
vermilion to scarlet, and posterior trian- 
gular area dark reddish brown. Abdomen 
much darker than carapace, terga of first two 
segments almost black anteriorly fading 
rapidly to brick red caudally; succeeding 
segments with paired subrectangular red- 
dish black splotches (gradually narrowing 
on posterior segments) dorsolaterally, flank- 
ing median glossy brick red longitudinal 
stripe, red spreading laterally on posterior 
part of each segment and spilling ventrally 
onto dorsal part of pleura, which mostly 
pale pinkish cream with posterior maroon 
spot. Telson, uropods, and chelae as de- 
scribed above. 

Size.—The largest specimen available is 
a female from Sevier County (locality 9, 
above) having a cl of 37.2 (pol 32.4) mm. 
The smallest and largest first form males 
have corresponding lengths of 24.8 (22.2) 
mm and 31.9 (28.5) mm, respectively. 

Life history notes. — First form males have 
been collected in April and June. Neither 
ovigerous females nor ones carrying young 
have been observed. 

Ecological notes.—This crayfish, like F. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


(F.) jeanae, constructs highly branching 
(complex) burrows in sandy clay soil. The 
largest colony visited by us was found at 
locality 9. There the roadbed is somewhat 
elevated above the adjoining wooded area 
(chiefly Pinus), and on the northern shoul- 
der of the elevated area, there is a seep in 
which scores of turrets mark the burrows of 
this crayfish. When one of us (HHH) first 
visited this locality in April 1973, the sandy 
clay soil was water-logged, and one could 
easily open and follow the complex system 
of galleries with one’s bare hands. On a visit 
three years later, following a period without 
rain, there were few turrets in the hard, dry 
ground, and, even with the aid of a shovel 
and considerable effort, only one juvenile 
was found! 


Subgenus Creaserinus Hobbs (1973) 


First pleopod never with proximomesial 
spur or cephalic process. Cheliped with suf- 
flamen; chela with tubercles on mesial sur- 
face of palm but sparse or lacking dorsolat- 
erally and laterally, lateral margin costate. 
Second pereiopod of male with mesial face 
of chela and carpus often bearing dense mats 
of plumose setae (lacking in F. (C.) burrisi, 
F. (C.) byersi, F. (C.) caesius, F. (C.) gilpini, 
and F. (C.) gordoni). Type species: Astacus 
fodiens Cottle, 1863:217. Gender: mascu- 
line. 


Fallicambarus (Creaserinus) caesius 
Hobbs 
Figs. Ir, 5 


Fallicambarus (Creaserinus) caesius Hobbs, 
1975:24—28, 33, fig. 7 [Types: holotype, 
allotype, and morphotype, USNM 
144921, 133922, 133923 G I, 2, 6 I); 
paratypes, USNM. Type locality: Road- 
side ditch at Hot Spring-Saline county 
line, Arkansas, on St Rte 67.]; 1981:269.— 
Bouchard, 1978:451; 1980:451.—Bou- 
chard & Robison, 1981:28.— Fitzpatrick, 
1983:168; 1987:439. 


VOLUME 102, NUMBER 3 


669 


Fig. 4. Dorsolateral views of Fallicambarus: a, F. (C.) petilicarpus; b, F. (F.) gilpini; c—e, F. (C.) fodiens from: 
c, Independence County, Arkansas; d, Santa Rosa County, Florida; e, Orangeburg County, South Carolina. 


Fallicambarus caesius.— Hobbs, 1975:28.— 
Bouchard & Robison, 1981:26. 


Diagnosis.— Ventral surface of merus of 
cheliped with mesial row of tubercles, lat- 
eral one never represented by more than 2. 
Lateral margin of chela strongly costate, 
never serrate, dorsal surface without scat- 
tered tubercles in lateral half, ventrolateral 
surface with arched row of prominent punc- 
tations bearing long setae; opposable mar- 
gin of dactyl with longitudinal row of tu- 
bercles extending along at least proximal 
third of finger. Mesial surface of palm of 
chela of second pereiopod lacking conspic- 
uous tufts of plumose setae. First pleopod 
without proximomesial spur, and lacking 


cephalic process; central projection weakly 
arched, its base not inclined laterally, its 
distal part directed caudoproximally, bear- 
ing well defined subapical notch, and never 
crossing central projection of corresponding 
pleopod. Hooks on ischia of third pereio- 
pods only. Boss on coxa of fourth pereiopod 
somewhat rounded, neither strongly com- 
pressed nor conspicuously protruding ven- 
trally. Mesial ramus of uropod with or with- 
out distolateral spine; distomedian spine 
premarginal. Telson incompletely divided 
and with or without spine on anterolateral 
flank of suture. 

Range and specimens examined.—Inso- 
far as is known, this crayfish is endemic to 
Arkansas where it is confined to the Oua- 


670 


pee = 


7 Oe 


Ly, “By 


Pig. 5: 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


ML OS See OL. UE Re an 
= op ¢ emi i coi pt eet ee ees ees ee -—— +--+ = S555 
$Y} Le ae >) , 
= eh ey | 
Phy See toe ee 9 ; ees we ahh 
- a fea --i- bey ; 
ARE 
{ oth 4 ' “ { 
“a 4 \ = 
o 


ih 5 . 
Ree At jx te 
os ee 
=-—y —~)-. LOAN : 
RS <———. ia 
— — Sei} 
3 nA 


Distribution of Fallicambarus (F.) harpi (doubly encircled star); F. (C.) caesius (encircled dot); and 


F. (C.) gilpini (encircled star) in Arkansas. (Some localities listed in text for F. (C.) gilpini are too close to be 


noted on map.) 


chita and Dorcheat Bayou basins in the 
southern part of the state. We have exam- 
ined material from the following localities. 
Clark County: (1) Rose Hedge Cemetery at 
Gurdon, 1 2, 12 Mar 1983, HWR; 1 ¢ II, 1 
Oo Apr 1982, WE:tld iy) bo93 46, 4jeke 
Apr 1983, DDK, HWR. Columbia County: 
(2) in Magnolia city limits, 1 2, 19 Mar 1984, 
J. Pesses. (3) Waldo, 1 9, 24 Aug 1981, EL. 
(4) seep 0.5 mi (0.8 km) W of Waldo at US 
Hwy 82, 1 2 with young, 17 Mar 1983, 
HWR. (5) seep 2 mi (3.2 km) W of Waldo 
at jct of Hwys 82 & 98, 4 9, 9 Apr 1983, 
HWR; 1 2, 3 jd, 1 j2, 2 ovig 2, 11 Feb 1984, 
HWR: Pop Ws Wy 9°O) Soh jes Feb 
1984, HWR. (6) behind Impson Whitehead 


Veterinary Clinic in Magnolia, 1 61, 10 Nov 
1979, M. Bryan. (7) Beene residence in 
Magnolia, 1 ¢ II, 22 May 1983, L. Robison. 
Dallas County: (8) 0.4 mi (0.64 km) N of 
Dallas-Ouachita Co line on St Rte 7, 1 jé, 
16 Apr 1983, HWR. Hempstead County: 
(9) Blevins, 1 ovig 2, 11 Apr 1984, J. Tucker; 
1 j4, 1j2, 20 Apr 1982, EL; 5j2, 8 Mar 1984, 
E. McMullen. (10) Blevins, Sec 15, T10S, 
R24W, 2 2, 15 Mar 1984, B. Scott. (11) 
Blevins, Sec 11, T10S, R24W, 3 2, 15 Mar 
1984, T. Taylor. (12) Bollins Bayou near 
Blevins, Sec 26, T9S, R24W, 1 juv, 25 Apr 
1983, HWR. (13) Blevins, Sec 6, T10S, 
R24W, 1 4 II, 2 2, 20 May 1983, C. Webb. 
(14) Blevins, Sec 20, T10S, R23W, 1 41, 29 


VOLUME 102, NUMBER 3 


Apr 1983, T. Winn. (15) Blevins, Sec 16, 
T10S, R24W, 1 41, 2 2, 19 May 1983, B. 
Stephens. Hot Spring County: (16) roadside 
ditch 2.0 mi (3.2 km) W of Grant Co line 
on US Hwy 270, 3 2, 1 j6, 30 Apr 1976, 
HHH & MK. Nevada County: (17) DeAnn 
Cemetery in Prescott, 2 2, 24 Nov 1980, K. 
W. Williams; 1 2, 10 Sep 1979, EL; 1 4], 
16 Feb 1982, KWW; 1 2 I, 12 Apr 1980, 
KWW. (18) 3 mi (4.8 km) E of Rosston on 
St Rte 4, 2 4 II, 4 2, 6 jd, 2 j2, 8 Mar 1984, 
DDK. (19) 0.4 mi (6.4 km) from jct of St 
Rte 19 and Cale Rd, between Laneburg and 
Rosston, Sec 7, 8, T13S, R21W, 5 4], 4 9, 
4 j2, 15 Apr 1983, DDK. Ouachita County: 
(20) 0.9 mi (1.44 km) N of jct of US Hwy 
79 and St Rte 203, 1 6 II, 26 Apr 1986, 
HWR. Saline County: (21) roadside ditch 
at Hot Spring County line on US Hwy 67 
(Type locality), 1 61, 2 6 II, 1 2, 2 32 (Type 
series), 22 Apr 1973, GBH, JEP, HHH. 
Color notes.—‘“‘(Based on freshly molted 
holotypic male.) Carapace bluish gray; dor- 
sal thoracic region and large arrow-shaped 
area (with base between origins of mandib- 
ular [adductor] muscles and extending to 
apex of rostrum) darker and more bluish 
than lateral surfaces of branchiostegites, he- 
patic, and posterior gastric regions where 
more olive than blue. Cephalic section of 
tergum of first abdominal segment midnight 
blue, and caudal section slate blue; succes- 
sive terga also slate blue but becoming pro- 
gressively lighter in color posteriorly to tip 
of telson. Second through fifth terga with 
reticulate, but almost symmetrical, pattern 
involving oblique sublinear, dorsolateral 
grayish cream markings. Sixth tergum and 
telson with ornate symmetrical light mark- 
ings. Uropods mostly very pale gray, but 
proximolateral parts somewhat darker with 
dark bluish splotches and dark median ribs. 
Antennae and pereiopods with powder blue 
reticulations. Antennular peduncle dark, 
antennal peduncle dark mesially and later- 
ally, but broad submedian area of penulti- 
mate podomere and lamellar part of anten- 


671 


nal scale very pale, lateral margin of scale 
dark. Cheliped with dorsodistal surface of 
merus, dorsal surface of carpus, dorsome- 
sial surface of palm, dorsal surfaces of fixed 
finger, and dactyl powder blue; both fingers 
with white tubercles on opposable margin 
and yellowish cream along distal portion; 
lateral costa cream, and fingers terminating 
in brownish cornified tips; bluish color on 
all podomeres fading ventrally to very pale 
pinkish cream; articular membranes with 
dark pink suffusion. Dorsal surface of re- 
maining pereiopods blue from merus dis- 
tally; basal podomeres and ventral surfaces 
of all pereiopods and sternum cream. Distal 
end of dorsal side of merus and dorsum of 
carpus and propodus of third maxilliped 
with blue reticulations’” (Hobbs 1975:27- 
28). 

Size.—The largest specimen is a female 
from Nevada County having a cl of 31.4 
(pol, 27.5) mm. The smallest and largest 
first form males have corresponding lengths 
of 21.1 (18.8) mm and 29.5 (26.4) mm, re- 
spectively. The smallest female carrying eggs 
or young has corresponding lengths of 27.2 
(24.4) mm. 

Life history notes. — First form males have 
been collected in February, April, May, and 
November. Ovigerous females were found 
in February and April; only one, having a 
carapace length of 28.4 mm, seemed to be 
carrying anything like a full complement of 
eggs: 35 with diameters of 2.1 to 2.3 mm; 
the diameters of the few eggs carried by the 
other two females were 2.0 or 2.1 mm. 

Ecological notes. —In the type locality, this 
crayfish was collected from highly branch- 
ing burrows in “rain soaked soil consisting 
of clay, organic material, and some gravel” 
(Hobbs 1975:28). As in many, if not most, 
of the other known localities, sedges and 
grasses were present in the immediate vi- 
cinity of the burrows or nearby, and none 
was taken from burrows more than one me- 
ter deep, most, if not all, of which opened 
to the surface through two or three chim- 


672 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 6. Distribution of Fallicambarus (C.) fodiens. Dots mark localities represented by at least one first form 


male. 


neys, and occasionally there was an opening 
to one of the galleries that was not marked 
by a turret. 


Fallicambarus (Creaserinus) 
fodiens (Cottle) 
Figs. 1b, 0, s, t, u, 6-9, 10a—x 


Astacus fodiens (Cottle, 1863:217) [Types: 
not extant. Type locality: ““Upper Cana- 
da,’ probably Ontario.].— Hobbs, 1969a: 
111; 1973:463.— Bouchard, 1976b:586.— 
Page, 1985:422. 

Cambarus obesus Hagen, 1870:82 [in 
part].— Faxon, 1885:71.—Osborn & Wil- 
liamson, 1898:21. 

Cambarus argillicola Faxon, 1884:115, 116, 
144; 1885:56, 72, 76-78, 160, 174, pl. IV: 
fig. 2; 1890:624-625; 1898:650, 690; 
1914:391, 400, 424, 426.— Underwood, 
1886:366.—Hay, 1891:147; 1896:478, 
491-493; 1899:959, 962; 1919:232; 1920: 
83.—Stebbing, 1893:208.—Osborn & 
Williamson, 1898:21.— Williamson, 
1899:48.—Harris, 1901:191; 1903a:59, 


71-72, 105, 137, 139-140, 142-144, 146, 
147, 150-155; 1903b:603, 605, 608.— 
Ortmann, 1902:277, 280, 283; 1905:120, 
123, 136; 1907:712.—Pearse, 1910a:10, 
11, 15, 19, 20, pl. VU; 1910b: 732519 
130.—Huntsman, 1915:158.—Cahn, 
1915:136, 174.—Cummins, 1921:28- 
30.—Engle, 1926:89, 93, 94, 97, 98.— 
Turner, 1926:146, 154, 156, 160-163, 
168, 169, 178, 186-188, 192.—Creaser, 
1931:263; 1932:336.—Lyle, 1937:2, 16; 
1938:76.—Brimley, 1938:503.—Bou- 
vier, 1940:71.—Hobbs, 1942:165; 1948: 
223, 224, 229, 230.—Rhoades, 1944: 
98.—Bovbjerg, 1952:34.—Eberly, 1955: 
283.—Crocker, 1957:90.—Hobbs & Hart, 
1959:187.—Penn & Marlow, 1959:195. 


Cambarus uhleri Faxon, 1884:116—117, 145 


[Types: holotype, MCZ 3,624; paratypes, 
MCZ 3,633, 3,635, 3,636. Type locality: 
“Swamp on Eastern Road near Felsbury, 
Somerset County, Maryland”’ (restricted 
by Faxon, 1914:426).]. Faxon, 1885:22, 
59, 77-78, 160, 166, 173, pl. VIII: figs. 
8, 8’, 8a, 8a’.—Underwood, 1886:373. 


VOLUME 102, NUMBER 3 


Ml) StS) Ma Sas Poe ad 


Fig. 7. Distribution of Fallicambarus (C.) fodiens in Arkansas. 


Cambarus diogenes.—Faxon, 1885:71 [in 
part]. 

Cambarus uhleri.—Hay, 1899:959, 962; 
1904:165.—Ortmann, 1902:277, 315; 
1905:119, 123, 128.—Harris, 1903a:59, 
Bevis. 138, 141, 142, 152, 158: 1903b: 
606.— Williamson, 1907:755.— Fowler, 
1912:568.—Faxon, 1914:400, 426.—Hay 
& Shore, 1918:401, pl. 28: fig. 6.—Creas- 
er, 1931:269.—Brimley, 1938:503.— 
Hobbs, 1942:165; 1948:229; 1955:95, 98; 
1959:896; 1966a:68, 70, 71; 1966b:115; 
1968:K16; 1981:270.—Penn, 1955:73.— 
Crocker, 1957:69, 90.—Crawford, 1959: 
150, 151, 177.—Meredith & Schwartz, 
fe so-2; 1960-4. 5,21, 23, 27,2830; 1962: 
2.—Hoffman, 1963:330.—Miller, 1965: 
43.—Hobbs III, 1969:42.—Hart & Hart, 
1974:73, 91.—Holt, 1973:93.— Pickett & 


Sloan, 1979:26.—Andolshek & Hobbs, 
1986:18. 

Cambarus (Bartonius) argillicola. —Ort- 
mann, 1905:120. 

Cambarus (Bartonius) uhleri.—Ortmann, 
1905:120. 

Bartonius argillicola.—Williamson, 1907: 
PAS. 152; 753; 158; 1627 163. 

Cambarus (Cambarus) uhleri.— Fowler, 
1912:341 [by implication]. 

Cambarus fodiens.—Huntsman, 1915: 
158.—Creaser, 1931:263; 1932:336.— 
Hobbs, 1941:121; 1942:165, 167; 1948: 
223, 224) 226, 229: 230: 195593" US: 
1959:896; 1966b:115; 1968:K16.— Penn, 
1941:8; 1955:73, 80, 81.— Rhoades, 1942: 
3; 1944:98: 1948:18; 1950:2, 3, 5; 1961: 
2, 4.—Hobbs & Marchand, 1943:6.— 
Bovbjerg, 1952:34-36, 40-54; 1970: 


674 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 8. Mesial view of first pleopods of Fallicambarus (F.) fodiens from following counties in Michigan: a, 
Saginaw; b, c, Washtenaw. Indiana: d, e, Wells; f, Marion. Illinois: g, Cook; h, Jasper; i, Clark; j, Efhngham; k, 
1, Richland. Ohio: m, Erie; n, Franklin. West Virginia: 0, Mason. Arkansas: p, q, Lawrence; r, Sharp; s, Clay; t, 


u, Greene; v, Craighead; w, x, Independence; y, Cross; z, St. Francis; a’, Phillips; b’, Grant; c’, Jefferson; d’, e’, 
f’, Dallas. 


232.—Pennak, 1953:464.—Eberly, 1954: 482.—Hobbs & Hart, 1959:149, 151, 
59; 1955:283.— Williams, 1954:810, 900, 159-161, 164, 169, 171, 185, 187-188, 
902, 912, 918.—Spoor, 1955:77.— fig. 11.—Hart, 1959:204.—Penn & Mar- 
Crocker, 1957:90.— Penn & Hobbs, 1958: low, 1959:195, 202.— Wiens & Armitage, 


VOLUME 102, NUMBER 3 675 


Fig. 9. Mesial view of first pleopods of Fallicambarus (F.) fodiens from following counties or parishes in 
Arkansas: a, Ashley; b, Bradley; c, Columbia; d, Miller; e, Little River; f, Sevier. Oklahoma: g, McCurtain; 
Texas: h, Upshur; i, Angelina; j, k, Jasper; 1, Brazos; m, Brazoria; n, Madagorda; 0, Victoria; p, Aransas. Louisiana: 
q, De Soto; r, Ouachita; s, Calciseau; t, East Baton Rouge. Tennessee: u, Tipton; v, Crockett; w, Shelby; x, y, 
Hardeman; z, Cheatham. Mississippi: a’, Lee; b’, Clay; c’, Oktibbeha; d’, Lowndes; e’, Noxubee; f’, Jones. 


1961:39-54.— Bowler, 1963:128.—Mob- 135, 139, figs. 28, 37, 46, 57, 62, 85.— 
berly, 1965:45.—Judd, 1968:1-4, 6, 8.— Fitzpatrick & Payne, 1968:14.—Jaspers 
Crocker é Barr, 1968:VIL, 12, 15, 29, & Avault, 1969:637.—Waywell & Corey, 
33, 35-37, 40, 56, 58-60, 125, 127, 129- 1970:1462-1464; 1972:294—298.—Bell, 


676 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


f 2 


Fig. 10. (All illustrations from first form males.) a—x, Mesial view of first pleopods of Fallicambarus (F.) 
fodiens from following counties in Alabama: a, Lauderdale; b, Limestone; c, d, Tuscaloosa; e, Perry; f, Choctaw; 
g, Maringo; h, Butler. Florida: i, Santa Rosa. South Carolina: j, Beaufort; k, Bamberg; 1, Colleton; m, Richland; 
n, Clarendon; o, Dillon; p, Marion; q, Horry. North Carolina: r, Columbus; s, Sampson; t, Hyde; u, Perquimines. 
Virginia: v, Norfolk; w, Warwick. Maryland: x, Dorchester. y, Same of F. (F.) strawni (cp, cephalic process); z, 
Lateral view of basal podomeres of cheliped of F. (C.) fodiens (su, sufflamen); a’, Same of F. (F.) jeanae; b’, 
Basal podomeres of fourth pereiopod of F. (C.) burrisi (cb, coxal boss); c’, Same of F. (C.) gordoni (cb, coxal 
boss). 


VOLUME 102, NUMBER 3 


1971:17.—Hobbs & Hall, 1974:200.— 
Terman, 1974:33, 34.—Williams, Wil- 
liams, & Hynes, 1974:365-369, figs. 1, 
2.—Caine, 1974:2.— Becker, Genoway, & 
Merrill, 1975:384.— Gladwell, Bowler, & 
Duncan, 1975:89.—Lake, 1977:59.— 
Berrill, 1978:166.—Momot, Gowing, & 
Jones, 1978:18.—Radaj, 1978:1.—Rei- 
nert, 1978:8.—Lawton, 1979:6.— Pickett 
& Sloan, 1979:26.—Bousfield, 1979: 
292.—Crenshaw, Lemons, & Russo, 
1980:245.—Grow & Merchant, 1980: 
234.—Grow, 1981:355.—Kiley & Dji- 
neen, 1982:212.—Maude & Williams, 
1983:68, 74, 76, figs. 6, 7.—McMahon & 
Wilkes, 1983:133. 

Cambarus (Bartonius) fodiens. —Creaser, 
1931:260, 261, 263, 269-272, fig. 37. 
Cambarus hedgpethi Hobbs, 1948:224—230, 
figs. 17a—-f, h-j, 1 [Types: holotype, mor- 
photype, USNM 85146 (6 I, 6 II), and 
allotype, USNM 85147 (8); paratypes, 
USNM. Type locality: lower middle part 
of Aransas National Wildlife Refuge, 
Aransas Co, Texas.].— Washburn, 1953: 
6.—Penn, 1953:74; 1955:73, 80; 1959:8, 
14-17, figs. 9, 27, 46, 64, map 9.—Penn 
& Hobbs, 1958:454, 462, 465, 467, 471, 
473, 476-478, figs. 11, 28, 42, 55.— 
Hobbs, 1959:896; 1966b:115; 1968: 
K16.—Penn & Marlow, 1959:195-197.— 
Hobbs & Barr, 1960:13.—Reimer, 1966: 
14; 1969:50, 51, 53, 60, 61, figs. 2, 39.— 
Black, 1967:176; 1969:197.—Walls & 
Black, 1967:60.—Fitzpatrick & Payne, 
1968:14, 20.—Walls, 1968:417.—Hobbs 
III, 1969:19, 21, 26, 49, 64, tab. 2. 


677 


336, 341, 343, 344, 350, 353, 422-426, 
436, figs. 155-158.— Phillips, 1980:84.— 
Grow, 1981:355.— Bouchard & Robison, 
1981:26, 27.—Huner & Barr, 1981:47; 
1984:42.—Berrill & Chenoweth, 1982: 
199.—Burr & Hobbs, 1984:14, 15, 16.— 
Norrocky, 1983:3.—Fitzpatrick, 1986: 
126:137. 


Fallicambarus uhleri.—Hobbs, 1969a:111, 


112, fig. 20); 1972:102, 147, figs. 82a, 83b, 
84a, 85a; 1973:463, 480, figs. 3d, 4; 1974b: 
24, 101, fig. 84; 1976:551, fig. la.— Hobbs 
& Fitzpatrick, 1970:835.—Peters, 1971: 
100; 1974:74; 1975:7, 22, 23.—Hart & 
Hart, 1974:22, 28, 33, 73, 129.—Hobbs 
III, Thorp, & Anderson, 1976:24.— Hobbs 
&uPeterspt97 7629.12" 15494 20921; 
33, 43, 46, 49, 53, 54, 60, 61, 62.— Whar- 
ton, 1978:50.—Page, 1985:422.—Fitz- 
patrick, 1986:126, 137, 138. 


Fallicambarus hedgpethi.—Hobbs, 1969a: 


111, 112, 173, fig. 20f; 1969b:335; 1972: 
102, 147, figs. 82c, 83d; 1974b:23, 100, 
fig. 83.— Hobbs & Hobbs, 1970:12, 14.— 
Hobbs & Fitzpatrick, 1970:835.—Bou- 
ehards3197 2:56 3062563) 4073.1977-11.— 
Albaugh, 1973:6, 11, 12, 25, 103.—Al- 
baugh & Black, 1973:183, 184, 185.— 
Payne & Riley, 1974:125-127.—Hart & 
Hart, 1974:23, 93, 94, 97.—Reimer & 
Clark, 1974:168, 175, figs. 27-30.—Rei- 
mer, 1975:24.—Lahser, 1976:278, 279, 
281-—284.—Huner, Meyers, & Avault, 
1976:150, 152.—Bouchard & Robison, 
1981:26, 27.—Huner & Barr, 1981:57, 58; 
1984:50.—Rogers & Huner, 1983:79; 
1984:37; 1985:23, 24, 26-28, figs. 3, 4, 


Cambarus hedgepethi.—Walls & Black, 
1967:60 [erroneous spelling]. 

Fallicambarus fodiens.—Hobbs, 1969a:111, 
112, fig. 20e; 1972:102, 137, figs. 5u, 82b, 


5.—Burr & Hobbs, 1984:15, 16.— Walls, 
1985:193.—Page, 1985:424.—Fitzpat- 
rick, 1986:137. 

Fallicambarus (Creaserinus) fodiens. — 


83c, 84b, 85b; 1974a:12; 1974b:23, fig. 
82; 1976:551, fig. 2d.—Hobbs & Fitz- 
patrick, 1970:835.—Bouchard, 1972:52, 
62, 63, 107; 1976a:14; 1976b:585, 586.— 
Hobbs & Barr, 1972:9.—Hart & Hart, 
1974:30, 31, 128.—Hobbs & Hall, 1974: 
200, 201, 203.— Page, 1974:99; 1985:335, 


Hobbs, 1973:463, 480, figs. 3g, 4.—Bou- 
chard, 1976a:14; 1976b:586.—Clark & 
Rhoades, 1979:238, fig. 1.—Bouchard & 
Robison, 1981:28.—Fitzpatrick, 1983: 
168, 169, fig. 175.—Thoma & Jezerinac, 
1982:136, 137.—Jezerinac & Thoma, 
1984:120-124, fig. 1.—Norrocky, 1984: 


678 


65.—Page, 1985:335.—Jezerinac, 1986: 
178.—Jezerinac & Stocker, 1987:46, fig. 
1.—Hobbs III & Jass, 1988:3, 23, 39-43, 
141, 142, figs. 30, 31.—Mansell, 1989. 

Fallicambarus (Creaserinus) uhleri.— 
Hobbs, 1973:463, 480, figs. 3d, 4. Hobbs 
& Peters, 1977 °6;-95. 127 13,198 20972 i 
33, 43, 46, 49, 53, 54, 60, 61, 62.—Fitz- 
patrick, 1983:168.—Cooper & Ashton, 
1985:9, 10.—Andolshek & Hobbs, 1986: 
24. 

Fallicambarus (Creaserinus) hedgpethi.— 
Hobbs, 1973:463, 480, fig. 4.— Bouchard 
& Robison, 1981:28.— Fitzpatrick, 1983: 
169. 

P{rocambarus] Fallicambarus fodiens. — 
Hart & Hart, 1974:88 [lapsus]. 

Fallicambarus hedgepethi.—Huner, 1977:11 
[photo in color]. 

Fallicambarus sp.—Huner, 1978:621. 

Fallicambarus (Creserineus) fodiens.—Je- 
zerinac, 1983:5 [erroneous spelling]. 


Diagnosis.— Ventral surface of merus of 
cheliped with mesial and lateral rows of tu- 
bercles; length of carpus less than, or sub- 
equal to, width of palm of chela. Chela with 
lateral margin costate to rounded, never ser- 
rate, dorsal surface without scattered tuber- 
cles in lateral half, ventrolateral surface 
lacking arched row of prominent setiferous 
punctations; opposable margin of dactyl 
with distinct excision in basal half, mesial 
margin with longitudinal row of tubercles 
extending along at least basal third of finger. 
Mesial surface of chela of second pereiopod 
with conspicuous tufts of plumose setae. 
First pleopod without proximomesial spur 
and lacking cephalic process; central pro- 
jection comparatively weakly arched, base 
not inclined laterally, distal part directed 
caudally, with or without subapical notch, 
but never crossing central projection of cor- 
responding pleopod. Hooks on ischia of third 
pereiopods only. Boss on coxa of fourth pe- 
reiopod somewhat rounded, neither strong- 
ly compressed nor conspicuously protrud- 
ing ventrally. Mesial ramus of uropod with 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


distolateral spine; distomedian spine pre- 
marginal. Telson divided and with spines 
on anterolateral flank of suture. 

Range and specimens examined.— From 
southern Ontario southwestward to Aran- 
sas County, Texas, and southeastward to the 
Apalachicola Basin, also present in the 
Coastal Plain and lower Piedmont from 
Beaufort County, South Carolina, to Som- 
erset and Dorchester counties, Maryland 
(Figs. 6, 7). Engel (1926:93) recorded it from 
Lincoln, Nebraska, but this locality is so far 
removed from any other reported for the 
species that it must be confirmed. We have 
examined some 2500 specimens from 
Michigan (45), Indiana (58), Illinois (42), 
Missouri (20), Arkansas (420), Oklahoma 
(13), Texas (223), Ohio (50), West Virginia 
(2), Kentucky (58), Tennessee (130), Loui- 
siana (29), Mississippi (143), Alabama (207), 
Florida (14), Georgia (15), South Carolina 
(451), North Carolina (386), Virginia (209), 
and Maryland (30). 

Color notes.—In this crayfish, there seem 
to exist two basic colors and three color 
patterns, with a wide range of variation link- 
ing them. Most of the animals that we have 
seen are either predominantly tan with 
brown markings or olive-green with dark 
grayish to brownish green markings. The 
lighter ventral and ventrolateral areas may 
be almost white, cream, yellowish, pink, or 
lavender. Occasionally we have encoun- 
tered individuals that are predominantly 
blue. 

As for the color patterns, the striped (Fig. 
4c, e) is the most common. In it, clearly 
defined longitudinal stripes, that are usually 
most obvious on the abdomen, make this 
pattern conspicuously different from the 
least common concolorous one. The latter 
is typified by the absence of stripes, splotch- 
es, or specks. Decidedly more common than 
the concolorous pattern 1s the speckled one 
(Fig. 4d; often the specks and splotches are 
much more abundant, especially dorsally, 
than depicted) in which the carapace and 
abdomen are marked dorsally by specks and 


VOLUME 102, NUMBER 3 


irregular dark spots scattered over a lighter 
background. Paired dorsolateral concentra- 
tions of darker pigment, which are often 
more diffuse than in Fig. 4d, on the abdom- 
inal terga no doubt represent remnants of 
the dark stripes that are so conspicuous in 
the striped pattern. In specimens exhibiting 
concolorous patterns, which are not illus- 
trated, the dorsum of the cephalothorax is 
an almost uniform dark brown, green, or 
blue, fading laterally to cream or very light 
gray. The abdominal terga, too, are almost 
uniformly dark brown, green, or blue almost 
to their junctures with the pleura, which, 
together with the uropods, are of a much 
diluted hue of the color of the dorsum. 

All three patterns have been observed re- 
peatedly among specimens collected in a 
single locality, the most recently seen were 
in a collection from western Sevier County, 
Arkansas, less than a mile from the Okla- 
homa line. 

Hay (1904:165) remarked that the spec- 
imens from Maryland were “‘a dirty green- 
ish brown, the tips of the chelae alone being 
somewhat reddish.” He reported further that 
Uhler, who first collected the species in 
Maryland, related to him the presence of 
beautiful yellow spots. While we have not 
observed the latter, we have encountered 
specimens with cream, yellow, orange, and 
red tipped chelae, but we have not associ- 
ated any of these colors with a particular 
geographic region. 

Size.— The largest specimen of this species 
that we have examined 1s a first form male 
from Richland County, Illinois, which has 
a cl of 42.8 (pol 36.7) mm. The smallest first 
form male (from Perquimens County, North 
Carolina) that we have seen has correspond- 
ing lengths of 19.5 (16.0) mm. Those of the 
largest and smallest from Arkansas are 36.4 
(32.4), from St. Francis County, and 22.8 
(19.2) mm, from Dallas County. Compa- 
rable measurements of the smallest female 
carrying eggs or young that we have seen 
are 26.5 (22.6) mm; this specimen was col- 
lected in Columbia County, Arkansas. 


679 


Life history notes.—Considering popula- 
tions throughout the range of the species, 
first form males have been collected during 
every month of the year; ovigerous females 
were found from January to June and in 
September, October, and November, and 
females carrying young from January to 
April and in September. In Maryland, this 
crayfish was reported by Hay (1904) to leave 
its burrows in the spring when it becomes 
common in ditches and small streams. 
Finding both first and second form males 
in September led him to conclude that the 
transition in males from form II to form I 
must occur in the late fall. 

The data for Arkansas are indeed inade- 
quate; except for a total of 11 members of 
the species collected in June, July, and No- 
vember, all of the material from the state 
available to us was collected during Feb- 
ruary (11), March (31), April (250), and May 
15). Among the collections from the state, 
there are first form males collected in Feb- 
ruary, March, April, May, and November, 
Ovigerous females in February, April, and 
November, and others carrying young in 
January, February, and March. 

The egg complements of three of the ovig- 
erous females appear to be reasonably com- 
plete and are as follows: cl 28.2, pol 24.8 
mm, 190 eggs; cl 28.2, pol 24.2 mm, 196 
eggs; cl 26.5, pol 22.6 mm, 177 eggs. The 
diameters of the eggs were 1.9 and 2.0 mm. 

Ecological notes.—Ecological data that 
have been recorded pertaining to F. (F.) fo- 
diens were recently summarized by Hobbs 
III & Jass (1988:41-43). The following dis- 
cussion is therefore limited to a few record- 
ed observations on the eastern facies of Fal- 
licambarus (C.) fodiens (formerly F. (C.) 
uhleri) and to those made by us on the pres- 
ence of this crayfish in Arkansas, where it 
frequents temporary pools and burrows from 
the floodplains of the major rivers to the 
foothills of the Ozark and Ouachita moun- 
tains. According to Faxon (1884:117), his 
C. uhleri occurs in “‘salt marshes, covered 
twice daily by the tides, and also in brackish 


680 


and fresh-water ditches... .”” On the eastern 
shore of Maryland, Hay (1904:165) found 
it to be “rather abundant in burrows in low- 
lying areas not far from the bay but always 
near ponds or ditches of freshwater. In near- 
ly every case the area selected was in dense 
pine woods.”’ Hay also learned from local 
inhabitants of the area that in the spring the 
crayfish ““emerge from their burrows and are 
common in ditches and small streams.” The 
original collection of the species taken by 
Dr. Philip Uhler was made partly in water 
that was distinctly brackish. 

In Arkansas, this crayfish is primarily an 
inhabitant of temporary bodies of water and 
burrows, although occasionally it ventures 
into more permanent lentic and lotic hab- 
itats. Apparently wherever it occurs in the 
State the water table must be within range 
of its burrowing capabilities, for we have no 
evidence that any members of the species 
undergo a life-span devoid of periods of what 
is an apparently voluntary fossorial exis- 
tence. Whereas we have retrieved a few 
specimens from burrows that did not pen- 
etrate the water table, most were taken from 
pockets of water in them at depths of 0.5 to 
1.5 meters. The external appearance of these 
burrows is described in the introductory re- 
marks where it was also pointed out that 
those excavated by members of this species 
are comparatively simple. Most consist of 
a subvertical passageway, in clay, sandy clay, 
or sandy loam, opening to the surface 
through one or two apertures that may or 
may not be surrounded by low, for the most 
part poorly formed, chimneys, and at the 
fundus of the passageway there is a slight 
enlargement. Occasionally we have encoun- 
tered a burrow with a second subvertical 
gallery leading downward from the main 
one. With little doubt, these simple I-, Y- 
or X-shaped patterns reflect, in part, phys- 
ical features of the environment involving 
the availability and retention of water. In 
habitats where there is evidence (based on 
the presence of hydrophytic plant commu- 
nities) that during much of the year the water 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


table is close to the surface, this species is 
largely confined to the marginal areas, and 
another (e.g., F. (C.) caesius or F. (C) gilpini) 
dominates, with its shallow, dendritic sys- 
tems of galleries, the more permanently wet 
parts of the seasonal seeps. The burrows of 
those individuals that invade the wetter 
areas do branch more than those we have 
regarded as more typical. Clumps of sedges 
frequently mark the sites of temporary 
stands of water, and areas in which a not- 
too-deep retreat of the water table occurs, 
features that together provide what appears 
to be the favorite habitat of the species in 
Arkansas. 

Most of the burrows that we have dis- 
sected contained a single individual, occa- 
sionally one inhabited by a female and a 
first form male, and sometimes one housing 
a female with young either clinging to her 
abdomen or sometimes with young that are 
presumed to have abandoned their mother. 
We have no information as to the perma- 
nency of habitation of a domocile by an 
individual. Until the current study of J. 
Norrocky (manuscript in preparation) in- 
volving marking and recapturing members 
of the species in Ohio, we were almost con- 
vinced that a female was a nearly permanent 
resident of the burrow she occupied, never 
wandering far away. This assumption was 
made, at least in part, on the basis of the 
size of the turrets marking the entrance/s. 
There can be little doubt that the larger tur- 
rets mark the burrows inhabited by the larg- 
er females, and there is no question that the 
more massive chimneys reflect more spa- 
cious and deeper domiciles. These obser- 
vations suggested to us the probability that 
the older, larger, females had spent a longer 
time enlarging and reworking their domi- 
ciles than had the smaller females and males, 
probably spending the better part of their 
entire lives hauling soil pellets to the sur- 
face. In light of some of Norrocky’s data 
(personal communication) there is good 
reason to question our conclusion as to the 
tenure of an individual in a single lair. While 


VOLUME 102, NUMBER 3 


the males must have their own burrow at 
the time of their presumed biannual molts, 
there is good reason to believe that when in 
form I they abandon their burrows, at least 
temporarily, for sojourns in the domains of 
one or more females that might be visited. 
The question as to whether or not they re- 
turn to their homestead following the breed- 
ing season or seek seclusion elsewhere re- 
mains unanswered. | 

Following rains, when pools flood the 
mouths of the burrows, the juvenile element 
of the population emerges in numbers and 
small crayfish may be observed wandering 
hither and yon, even during daylight hours. 
An occasional adult also appears in the open 
water, and, no doubt, many more adults 
leave their lairs for short forays in the pools 
at night, but most return to burrows by at 
least the early morning hours. 

We are much puzzled by the paucity of 
colonies of this crayfish throughout most of 
the rice-growing areas of the state. Surely 
when the region was still wooded, and shal- 
low temporary pools were common features 
of the landscape, numerous colonies of F. 
(F.) fodiens must have existed between the 
Arkansas and Mississippi rivers. Now, one 
must search rather diligently to find even 
an isolated burrow in much of the area un- 
der cultivation. It is understandable that a 
crayfish might have difficulty in remaining 
well established in fields that are subjected 
to the treatment accorded the cultivation of 
rice, but why should they not be present in 
the roadside ditches that border these vast 
tilled and alternately flooded and drained 
lands? In Cross County, for example, a care- 
ful search of the ditches along highways and 
secondary roads for miles revealed only one 
colony of this crayfish, though the lawn 
around one of the churches in Hickory 
Grove was pitted by scores of burrows that 
must have been constructed by members of 
this species. Not only is this crayfish largely 
absent where it should occur in numbers, 
but other species (Procambarus (Ortman- 
nicus) acutus acutus (Girard, 1852), Cam- 


681 


barus (Lacunicambarus) diogenes Girard, 
1852, and C. (L.) ludovicianus Faxon, 1914) 
are also encountered infrequently. It seems 
likely that something associated with the 
production of rice is affecting adversely the 
exploitation of the area by crayfishes. 

We are further puzzled by our failure to 
find a single member of this species between 
the Arkansas and White rivers downstream 
from Pope County. We do not claim to have 
exhausted possibilities of the existence of 
overlooked colonies, but considerable effort 
has been expended along several routes tra- 
versing the area between the two rivers. 

Remarks.— United here are three for- 
merly recognized species which for a num- 
ber of years have presented difficulties to 
one of us (HHH) in searching for charac- 
teristics that might be used in their differ- 
entiation (e.g., Hobbs 1959, 1972, 1973, 
1981). In preparing an account of the Fal- 
licambarus from the Apalachicola basin in 
his study of the crayfishes of Georgia, Hobbs 
(1981) assigned the specimens that he had 
previously identified as members of Cam- 
barus fodiens (Hobbs & Hart, 1959:187) to 
F. hedgpethi. But before doing so he had 
vacillated between assigning them to one of 
the two and to describing them as new! Had 
the material subsequently collected in east- 
ern Arkansas and from a number of other 
localities, as well, been available to him, it 
is likely that he would have arrived at the 
conclusions that have been reached in the 
current study. 

Given specimens from the vicinity of the 
type localities of these three crayfishes, we 
do not believe that anyone would have dif- 
ficulty in distinguishing between them: the 
first pleopod of the first form males of F. 
fodiens exhibits a much shorter central pro- 
jection than do those of either F. hedgpethi 
or F. uhleri, and the latter has an areola that 
constitutes less than 39 percent of the length 
of the carapace whereas that of hedgpethi is 
more, and the opposable margin of the dac- 
tyl of the chela with two instead of one ma- 
jor tubercle is typical only of F. hedgpethi: 


682 


too, in the latter the arrangement of the tu- 
bercles on the mesial surface of the dactyl 
of the chela in two well developed rows dif- 
fers from the usual single well developed 
row in the other two. The limited known 
range of fodiens when uhleri was described 
from Maryland by Faxon in 1884, and the 
existing poor concept of the distribution of 
the two when hedgpethi was found in south- 
western Texas, gave neither Faxon nor 
Hobbs reason to question the validity of the 
seemingly distinctive characters that they 
chose in naming what we now believe to be 
peripherally located populations of a single 
species. These occur at the angles of a large, 
distorted, triangular range which in Plio- 
cene, and probably in part of Pleistocene, 
times must have been continuous. The range 
appears even now to be unbroken except for 
a gap apparently existing across the south- 
ern part of Georgia where members of the 
subgenera Hagenides and Leconticambarus 
of the genus Procambarus are probably vi- 
cariating for F. fodiens (see Hobbs 1981: 
317, 348). 

Comparisons of the materials from 
throughout the range of the species have 
been made, and we have discovered only a 
few characters (those associated with the first 
pleopod of the first form male) that are geo- 
graphically or ecologically restricted to a 
limited part of the range of the species. The 
measurements made of the carapace and 
chelipeds that have been translated into ra- 
tios suggest that some local populations are 
rather distinctive, but they, too, are not con- 
sistent for large segments of the range, and 
there are no indications of clinal trends. Un- 
til now, the Fallicambarus ranging along the 
Atlantic versant from South Carolina to 
Maryland has been identified as F. (C.) uh- 
leri. In southern South Carolina the areola 
of this crayfish spans from 36 to 41 percent 
of the carapace length, in North Carolina 
from 34.8 to 38.7 percent, and in Virginia 
and Maryland 35.2 to 38.5 percent, and in 
several localities in South Carolina the ratio 
is distinctly above 39 percent. When these 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


specimens from South Carolina are com- 
pared with others from the state, we find 
nothing else that will set them apart, and in 
the midst of their range there are specimens 
with areolae occupying as little as 36 percent 
of the carapace length. If the comparisons 
are extended to specimens from more west- 
ern localities including those from Arkansas 
and Texas the ratios range from 32.7 in 
Cheatham County, Tennessee to 41.5 in 
Tuscaloosa County, Alabama, and Brazos 
County, Texas. Moreover, there seems to 
be no distributional pattern in the varia- 
tions between these extremes. 

Even the three color patterns (concolor- 
ous, speckled, and striped) that have been 
noted appear in a single population. Most 
of the specimens that we have examined 
from Illinois to Ohio and West Virginia 
northward possess chelae in which the op- 
posable margin of the fixed finger bears one 
tubercle that is slightly to distinctly larger 
than the others, and the tubercles on the 
mesial surface of the dactyl are largely 
aligned in a single row. Specimens from the 
Atlantic versant exhibit, for the most part, 
similarly adorned chelae. In contrast, how- 
ever, in most of those in the lower gulf coast- 
al area, the fixed finger bears two large tu- 
bercles and those on the mesial margin of 
the dactyl form two well developed rows. 
In eastern Arkansas, Tennessee, and Ala- 
bama, these features appear in a haphazard 
fashion. 

First pleopods of first form males from 
throughout the range of the species are de- 
picted in mesial aspect in Figs. 8-10. On 
the basis of variations noted in specimens 
from Ontario (see Crocker & Barr 1968: fig. 
28), Michigan, and Ohio to Aransas Coun- 
ty, Texas (from Missouri southward only 
west of the Mississippi River), and were 
there no populations occurring east of the 
river, we should not hesitate to conclude 
that two subspecies of F. fodiens should be 
recognized. The nominate subspecies (dis- 
tinguishable almost solely on features of the 
first pleopod of the first form male in which 


VOLUME 102, NUMBER 3 


the comparatively short central projection 
usually bears a subapical notch (Fig. 8a—y)) 
could be considered to occupy the northern 
sector, extending as far south as northern 
Arkansas, where throughout the eastern part 
of the state it intergrades (Figs. 8z—9g) with 
the more southwestern populations, ranging 
from southwestern Arkansas and south- 
eastern Oklahoma southward. These more 
southern populations exhibit the facies that 
has been associated with F. hedgpethi (rec- 
ognized by the possession of a long central 
projection lacking a subapical notch, Fig. 
9h—s). It should be noted that an occasional 
influence of the fodiens genome surfaces in 
specimens occurring south of the Arkansas 
border (see Fig. 97, g). East of the Mississippi 
River, we fail to find any such regular dis- 
tribution pattern in the variation of pleo- 
podal features (Figs. 97-107) except along 
the Atlantic versant from South Carolina to 
Maryland (Fig. 10j—x). But the same type 
pleopod that characterizes those popula- 
tions occurring along the eastern seaboard 
may be found in specimens from Alabama 
and Texas (Fig. 97, t). Thus, we have been 
unable to discover a single character that 
serves consistently to distinguish between 
the formerly recognized F. (C.) fodiens, F. 
(C.) uhleri, and F. (C.) hedgpethi. The two 
diagnostic features that Faxon (1884:117) 
mentioned as setting his Cambarus uhleri 
apart from C. argillicola (=F. fodiens) were 
its ““‘plane rostrum [and] shape of the hand. 
... Many specimens, particularly those 
from the Carolinas, have concave rostra, 
and while we are not certain as to which 
features in the “‘shape of the hand”’ Faxon 
was referring, there seems to exist as much 
variation within specimens from South Car- 
olina to Maryland as we have noted in in- 
dividuals from the rest of the range of the 
species, and we have recognized no feature 
as being unique. As noted above, the same 
applies to characteristics pointed out by 
Hobbs as typifying his ““Cambarus hedg- 
pethi.” 

Notes on sex ratio.—In all of the studies 


683 


of which we are aware that have yielded data 
on the sex ratios of cambarids except that 
of Creaser (1934) (e.g., Andrews 1904, Penn 
1943, Smith 1953:92, and Smart 1962:94), 
all have revealed a near 1:1 ratio. Insuffi- 
cient numbers of individuals of any popu- 
lation of members of the genus Fallicam- 
barus in Arkansas have been available that 
might permit an estimate of the sex ratio in 
any population, but, in the samples at hand 
there are many more females than males. 
Most of our adult specimens of Fallicam- 
barus (F.) fodiens were removed from bur- 
rows, and of 293 adults, less than half, only 
98, are males. In order to support the belief 
that our data are little biased, we have re- 
peatedly attempted to discover some way 
in which to determine whether a burrow to 
be excavated contains a male, female, or 
pair, but we have been unsuccessful. (As will 
become evident below, the importance of 
obtaining first form males from throughout 
the range of the species is paramount.) On 
21 April 1973, in a seepage area 0.4 mile 
east of the Oklahoma State line on USS. 
Highway 70, in Sevier County, Jean Pugh 
and HHH removed 23 females from bur- 
rows before they found a male. In Phillips 
County, on 17 April 1985, Robert Gilpin 
and HHH retrieved females from 14 bur- 
rows in one locality without finding a male, 
and, at another nearby, they unearthed five 
females before taking a male. Except for their 
apparent rarity, we are aware of no evidence 
that the males of F. (F.) fodiens might be 
more secretive than are the females; and, 
we suggest that perhaps two of their habits 
are responsible for the real or apparent ab- 
sence of half of them (assuming the sex ratio 
at hatching 1s near 1:1) from the adult pop- 
ulation. To a minor extent, perhaps our data 
are biased, for the burrows of the males seem 
sometimes to be less elaborate than those 
of the female, and this is reflected in the 
often smaller, open, and eroded turrets 
marking their domiciles. Admittedly, such 
burrows offer less temptation to the collec- 
tor, who, for good reason, prefers to explore 


684 


one over which the turret is capped, or, if 
open, adorned with comparatively recently 
deposited, not abraded, pellets. Well-formed 
soil pellets offer evidence of the presence of 
a crayfish instead of perhaps a snake (Ag- 
kistrodon piscivoris or an ill-tempered Ne- 
rodia sipidon), an Amphiuma, or some other 
invader. At one time we suspected that of 
more importance in skewing the apparent 
sex ratio of adults than failure of collectors 
to excavate burrows harboring males are the 
supposed more frequent wanderings of first 
form males in seeking mates. In their forays 
from one lair to another, they place them- 
selves in jeopardy of becoming prey to rac- 
coons, skunks, owls, and other predators, 
and indeed scat from owls and raccoons has 
been observed to contain fragments of the 
exoskeleton of crayfishes within the range 
of the species in Arkansas. Thus by their 
being more frequently exposed to predators 
than are females, we reasoned that they are 
being passively selected, and perhaps effec- 
tively so. What significance, if any, attaches 
to our having observed more carcasses of 
first form males than females of this species 
in areas where their burrows are located es- 
capes us. 

We had placed considerable confidence 
in the above suggestions as possible expla- 
nations for the apparent skewed sex ratio 
existing in the adult populations of F. fo- 
diens until, on 22 April 1988, one of us 
(HWR) collected 30 juveniles (cl 5.5 to 11.5 
mm) of this species from a pool in the ditch 
at the locality mentioned above where Pugh 
and Hobbs had collected in 1973. Twenty- 
nine of the specimens were females! The 
question remains as to whether or not the 
sex ratio at hatching in this species is 1:1, 
and if it is what factor/s (cannibalism of the 
perhaps slower growing juvenile males—the 
male is the smallest of the 30 juveniles) are 
responsible for the subsequent alteration of 
the ratio. That our suppositions might well 
be suspect seem possible when we remem- 
bered Creaser’s (1934) study revealing pos- 
sible seasonal changes in populations of the 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


lake- and stream-dwelling Faxonius (=Or- 
conectes) propinquus in which from August 
to January 66 to 73 percent of the popula- 
tion was female; during the rest of the year 
the percentage ranged from 43 to 56 per- 
cent. Creaser was apparently as puzzled by 
these data as are we by the information we 
have on F. (C.) fodiens. 


Fallicambarus (Creaserinus) gilpini, 
new species 
Figs. 1g, 5, 11 


Diagnosis.—Cheliped with sufflamen; 
ventral surface of merus with mesial row of 
tubercles, lateral one never represented by 
more than two; length of carpus less than 
or subequal to width of palm of chela. Chela 
with lateral margin strongly costate, never 
serrate, dorsal surface lacking scattered tu- 
bercles in lateral half, ventrolateral surface 
with arched row of prominent punctations 
bearing long setae, opposable margin of dac- 
tyl with distinct excision 1n basal half, me- 
sial margin without tubercles. Mesial sur- 
face of palm of second pereiopod lacking 
conspicuous tufts of plumose setae. First 
pleopod lacking proximomesial spur, and 
lacking cephalic process; central projection 
weakly arched, its base not inclined later- 
ally, its distal part directed caudoproximally 
with well defined subapical notch, never 
crossing central projection of corresponding 
pleopod. Hooks present on ischia of third 
pereiopods only. Boss on coxa of fourth pe- 
reiopod somewhat rounded, not distinctly 
compressed. Mesial ramus of uropod with 
distolateral spine; distomedian spine pre- 
marginal. Telson incompletely divided, with 
spine on anterolateral flank of suture. 

Holotypic male, form I.—Eyes small but 
pigmented and with faceted cornea. Body 
subcylindrical, very weakly compressed 
(Figs. 4b, 11a, 1). Abdomen distinctly nar- 
rower than thorax (7.3 and 10.1 mm). 
Greatest width of carapace at level about 
one-third length of areola from cervical 
groove where subequal to height (10.1 and 


VOLUME 102, NUMBER 3 685 


Fig. 11. Fallicambarus (Creaserinus) gilpini (all from holotype except c, e, from morphotype, and i, n from 
allotype): a, Lateral view of carapace; b, c, Mesial view of first pleopod; d, Dorsal view of caudal part of abdomen; 
e, f, Lateral view of first pleopod; g, Postaxial view of mandible; h, Caudal view of first pleopods: i, Annulus 
ventralis and associated sclerites; j}, Antennal scale; k, Epistome; 1, Dorsal view of carapace; n, Basal podomeres 
of third, fourth, and fifth pereiopods; n, 0, Distal podomeres of cheliped. 


686 


10.0 mm). Areola linear over most of length 
and comprising 40.5 percent of entire length 
of carapace (46.6 percent of postorbital car- 
apace length). Rostrum with convergent, 
slender margins gently contracted anterior- 
ly, marking base of poorly delimited acu- 
men, apex corneous, slightly upturned, and 
reaching base of ultimate podomere of an- 
tennular peduncle. Dorsal surface of ros- 
trum concave with submarginal rows of se- 
tiferous punctations and others between, 
especially dense and conspicuous in basal 
part. Subrostral ridges weak but evident in 
dorsal aspect to base of acumen. Postorbital 
ridges slender but well defined and merging 
almost imperceptibly with carapace above 
posterior margin of orbit. Branchiostegal and 
cervical spines absent. Suborbital angle ab- 
sent, cephalolateral margin of carapace 
sloping caudoventrally from base of ros- 
trum without excrescence or excision. Car- 
apace comparatively densely punctate dor- 
sally and laterally; row of few small tubercles 
flanking anterolateral segment of cervical 
groove. 

Abdomen (Fig. 45) shorter than carapace 
(19.8 and 22.0 mm); pleura small and 
broadly rounded ventrally, none with an- 
gular caudoventral angle; pleuron of first 
segment clearly overlapped by that of sec- 
ond. Telson (Fig. 11d) not divided but deep- 
ly incised laterally and caudolateral angles 
of cephalic section bearing two pairs of 
spines, more mesial pair smaller and mov- 
able. Proximal podomere of uropod with 
mesial lobe bearing acute angle, lateral one 
rounded; mesial ramus with distolateral 
spine and smaller premarginal distomedian 
spine. 

Cephalomedian lobe of epistome (Fig. 
11k) subtriangular with cephalomedian 
prominence; cephalolateral margins elevat- 
ed ventrally and only slightly undulant; main 
body of epistome depressed but lacking fo- 
vea. Ventral surface of proximal podomere 
of antennule lacking spine. Antennal pe- 
duncle without spines, flagellum falling short 
of caudodorsal margin of carapace. Anten- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


nal scale (Fig. 11/) small, just reaching base 
of acumen and penultimate podomere of 
antennular peduncle; lamella broadly 
rounded distomesially, broadest distal to 
midlength, but only slightly wider than 
thickened lateral part. Mandible (Fig. 11g) 
with cornified subtriangular area of caudal 
molar process proportionately much small- 
er than that of F. (F.) devastator and farther 
removed from corneous tuberculiform ce- 
phalic molar process. Ventral surface of is- 
chium of third maxilliped with lateral row 
of short, plumose setae, and mesial half 
bearing clusters of longer stiff setae; basis 
with conspicuous cluster of long setae ob- 
scuring proximal part of ischium. 

Right chela (Fig. 110) about 2.2 times as 
long as broad, rather strongly depressed; 
width of palm about 1.7 times length of 
mesial margin, latter bearing row of seven 
tubercles and one prominent one lying 
slightly dorsal to row between third and 
fourth from proximal end; irregular row of 
six much smaller tubercles on dorsal flank 
of mesial row; dorsal surface of palm and 
fingers bearing setiferous punctations, those 
on and adjacent to base of fixed finger and 
on proximal half of dactyl conspicuous; lat- 
eral margin of chela rounded proximally, 
but largely costate; ventral surface punctate 
except for single prominent tubercle oppo- 
site base of dactyl, ventrolateral arc of punc- 
tations, each made prominent by long stiff 
seta, extending from base of palm to base 
of distal third of fixed finger. Both fingers 
with well defined submedian ridge flanked 
by punctations dorsally; ridges on ventral 
surface less well defined. Opposable margin 
of fixed finger with row of four tubercles 
(third from base largest) along proximal third 
and one projecting from lower level slightly 
distal to midlength; single row of minute 
denticles extending from third tubercle from 
base to corneous tip of finger. Opposable 
margin of dactyl with obvious excavation 
proximally, two tubercles borne on margin 
of excavation and one marking its distal ex- 
tremity, all subequal in size; single row of 


VOLUME 102, NUMBER 3 


minute denticles extending from distalmost 
tubercle to corneous tip of finger. Mesial 
margin of dactyl bearing setiferous punc- 
tations, lacking even basal tubercles. 

Carpus of cheliped about 1.4 times as long 
as broad and approximately 1.7 times as 
long as mesial margin of palm. Dorsal sur- 
face with deep submedian longitudinal sul- 
cus flanked by setiferous punctations; me- 
sial surface of podomere tuberculate, that 
on distal margin acute and much larger than 
more proximal ones; lateral and ventral sur- 
faces punctate; ventrodistal margin with two 
acute, corneous tubercles: one on ventro- 
lateral condyle and other mesial to it. Merus 
with few squamous to rounded tubercles 
near dorsodistal extremity, two somewhat 
larger than others; mesial and lateral sur- 
faces finely punctate; ventral surface with 
mesial row of eight tubercles (nine on left); 
usual lateral row absent but single tubercle 
present on left member; lateral row of tu- 
bercles characteristic of most crayfishes ab- 
sent; longitudinal row of long, stiff setae 
present. Ischium punctate, lacking tubercles 
ventromesially. Chela of second pereiopod 
with marginal row of setae on palm, and 
carpus with dorsal row of long setae; mesial 
surface of carpus and propodus lacking tufts 
of plumose setae. 

Ischium of third pereiopod only with hook 
(Fig. 11m); latter simple, not overreaching 
basioischial articulation, and not opposed 
by tubercle on corresponding basis. Coxa of 
fourth pereiopod with knoblike caudome- 
sial boss which, if leg extended laterally, 
projecting mesially. Coxa of fifth pereiopod 
devoid of boss, ventral membrane setifer- 
ous. 

First pleopods (Fig. 11b, f h) reaching 
coxae of third pereiopods and largely hid- 
den by setae extending mesially from ven- 
tral margin of sternum. When abdomen 
flexed, however, apices of terminal ele- 
ments protruding ventrally beyond setal 
curtain. Proximomesial spur lacking. Shaft 
of appendage only slightly bowed, with two 
terminal elements disposed caudally at about 


687 


90 degrees. Mesial process somewhat spat- 
ulate, shallowly chamfered, and tilted lat- 
erally. Corneous central projection blade- 
like, with distinct subapical notch, arched 
and reaching caudally to about same level 
as mesial process. 

Allotypic female.— Differing from holo- 
type in other than secondary sexual char- 
acters as follows: acumen not quite reaching 
base of ultimate podomere of antennular 
peduncle; postorbital ridges not reaching 
level of posterior margin of orbit. Abdomen 
subequal in length to carapace. Distolateral 
spine on mesial ramus of uropod greatly 
reduced (perhaps abraded), that on right 
represented by no more than angle; because 
of encrustation, dorsal surface of telson and 
uropods appearing much more strongly se- 
tose than that of holotype; both lobes of 
proximal podomere of uropod rounded. 
Antennal scale slightly overreaching acu- 
men and base of penultimate podomere of 
antennular peduncle. Mandible with ce- 
phalic molar process strongly abraded, cau- 
dal molar process lacking corneous ele- 
ments. Mesial margin of palm of right chela 
(Fig. 117) with row of 5 tubercles (left with 
6) and row of 4 on dorsal flank (left lacking 
second row but with single tubercle, be- 
tween fourth and fifth, ventral to row); dacytl 
with 2 small tubercles distal to 3 associated 
with proximal excavation; ischium with 
ventromesial row of 12 tubercles (11 on left). 
(See Table 2 for mensural features.) 

Annulus ventralis (Fig. 117), 1.5 times as 
broad as long, situated deeply in sternum; 
cephalic region immovable, but caudal two- 
thirds capable of hingelike motion. Sulcus 
shallow and narrow cephalically, becoming 
deeper and broader caudosinistrally; high, 
prominent caudal wall cut by C-shaped si- 
nus arising from fossa at caudosinistral side 
of sinistrally projecting tongue. Postannular 
sclerite less than half as long and approxi- 
mately half as wide as annulus with punc- 
tate, oval, ventrally elevated median area. 

Morphotypic male, form II.—Differing 
from holotype in following respects: Tip of 


688 


Table 2.— Measurements (mm) of Fallicambarus (F.) 
gilpini. 


Morpho- 
Holotype Allotype type 
Carapace: 
Entire length 22.0 24.9 28.3 
Postorbital length 9et DAT PL 5).8) 
Width LO 11.6 13.0 
Length 10.0 10.9 be 
Areola: 
Width — — — 
Length 8.9 10.0 11.4 
Rostrum: 
Width 35 35 4.2 
Length 321 Sit 3.9 
Right chela: 
Length, palm 
mesial margin ee 323 4.8 
Palm width 5.4 6.5 7.4 
Length, lateral 
margin 11.8 (2-7 12 
Dactyl length 8.1 8.5 10.1 
Abdomen: 
Width os | 93 
Length 19.8 25a 25.0 


rostrum abraded but acumen reaching mid- 
length of penultimate podomere of anten- 
nular peduncle. Postorbital ridges termi- 
nating slightly posterior to caudal margin of 
orbit; lateral surface of branchiostegites 
granular; telson divided; both lobes of prox- 
imal podomere of uropod rounded; main 
body of epistome with cephalomedian fo- 
vea; left antennal scale as in allotype, right 
with regenerated distolateral spine; right 
chela with only five tubercles in row on dor- 
sal flank of mesialmost row; opposable mar- 
gin of fingers armed as in allotype; oppos- 
able margin of dactyl of left chela with seven 
tubercles, four associated with excavation 
and three distal to it. 

Except for terminal elements of first pleo- 
pod (Fig. 1 1c, e), no noteworthy differences 
noted between morphotype and holotype. 
Mesial process more robust with less con- 
spicuous groove, but projecting caudally 
much beyond tip of central projection; lat- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


ter, in addition to being stouter and non- 
corneous, also lacking subapical notch. 

Color notes.—Basic color pale greenish 
blue, but cephalic region more lavender than 
blue; rostral margins and postorbital ridges 
distinctly dark green. Mandibular adductor 
and posterior part of gastric area lavender 
with faint greenish suffusion. Lateral ce- 
phalic region fading to white ventrally, but 
with paired small navy blue spots midway 
between tip of rostrum and caudal extrem- 
ity of cervical groove and another less well 
defined pair abutting cervical groove. Tho- 
racic area dark greenish blue dorsally, sud- 
denly changing to white laterally, and at least 
half of branchiostegite white. First abdom- 
inal tergum dark greenish blue, second 
slightly paler, and third through sixth yet 
more pale with faint hint of very pale dor- 
somedian longitudinal stripe; white margin 
of all pleura partly separated from blue to 
bluish green tergum by series of short dark 
greenish blue bars. Telson with cephalic sec- 
tion mostly bluish, caudal section colorless 
and translucent; lateral section of lateral ra- 
mus of uropod pale bluish green, keels of 
both rami and margins of basal podomere 
dark greenish blue. Antennules and anten- 
nae with lateral and mesial borders of pe- 
duncles darker blue than dorsal and ventral 
surfaces; flagella greenish blue basally fad- 
ing to pale tan. Cheliped with dorsal surface 
of distal part of merus, carpus and chela 
dark bluish green; tips of fingers yellowish 
with corneous brown tip; ventrolateral part 
of palm and fixed finger fading to cream; 
ridges and tubercles on carpus and palm of 
chela very dark blue. Venter and basal pod- 
omeres of remaining pereiopods white; dor- 
sum of merus, carpus, and propodus of sec- 
ond through fifth pereiopods bluish green, 
dactyl with some blue but more cream to 
yellowish. 

Size.— The largest specimen available is 
a second form male having a carapace length 
of 28.8 (postorbital carapace length, 26.0) 
mm. Corresponding measurements of the 
smallest first form male were 21.1 and 18.8 


VOLUME 102, NUMBER 3 


mm, and those of the smallest ovigerous 
female, the allotype, are 24.9 and 21.7 mm, 
respectively. 

Type locality.—Roadside seepage 3.1 mi 
south of southern junction of State Route 
54 and U.S. Highway 79 at junction of latter 
with Pepperridge Road (T7S, R1OW, Sec 
19), approximately 11 miles south of Pine 
Bluff and about 3 miles north of Cleveland 
County line, Jefferson County, Arkansas. 

Disposition of types.—The holotype, al- 
lotype, and morphotype (USNM 219511, 
219512, and 218944, respectively) are de- 
posited in the National Museum of Natural 
History, Smithsonian Institution, as are the 
paratypes consisting of 1 46 I, 1 6 II, 8 9, 2 
juv 6, 2 juv 2, 2 ovigerous 2. 

Range and specimens examined.— All of 
the specimens were collected from burrows 
in roadside seepages in Jefferson County, 
Arkansas: (1) type locality, 1 6 II, 26 Apr 
1986, HWR; 5 2, 2 juv 4, 2 juv 2, 18 Mar 
1987, HWR (two additional juvenile males 
were maintained in aquaria until they molt- 
ed to first form, one in late Feb 1988, and 
the other, the holotype, on 9 or 10 Apr 1988). 
(2) 0.2 mi S of Pine Bluff city limits, 1 2, 11 
Apr 1986, HWR, coll. (3) 3.6 mi N of Cleve- 
land Co. line on US Hwy 79, 2 2, 7 Nov 
1987, B. F. Kensley, HWR, HHH; 2 ¢ II, 2 
2, 3 ovig 2, 11 Mar 1988, HWR. 

Variations.— Among the adult speci- 
mens, the areola constitutes from 38.1 to 
40.6 percent of the total length of the car- 
apace, and from 42.1 to 46.6 percent of the 
postorbital carapace length. Most of the dif- 
ferences noted in the specimens may be at- 
tributed to abrasion and to regeneration of 
chelipeds. The former is reflected in the ab- 
sence of an upturned tip on the rostrum, 
reduced and/or truncated tubercles and 
spines, and broken or missing setae (partic- 
ularly noticeable on the third maxillipeds 
of specimens in late intermolt stages). Re- 
generated chelipeds can usually be recog- 
nized by the absence or reduction of the 
excavation on the proximal part of oppos- 
able margin of the dactyl of the chela, but 


689 


also by the smaller, and frequent increase 
in number of tubercles on the opposable 
margins of both fingers. The number of tu- 
bercles in the ventromesial row on the me- 
rus of the cheliped ranges from 7 to 11, on 
the mesial margin of the palm of the chela, 
from 5 to 7 with 0-6 on the dorsal flank; on 
the opposable margin of both fingers, there 
are from 4 to 6 tubercles (in one specimen 
2, instead of 1, lie at a lower level on the 
fixed finger). There is no noteworthy vari- 
ation in the secondary sexual features of the 
two available first form males, and in the 
female, the only conspicuous difference ob- 
served is the anticipated occurrence of a 
mirrored image of features of the annulus 
described in the allotype. 

Life history notes.—No first form male 
has been collected, and the only two avail- 
able were reared from very small juveniles 
collected in March 1987. They were main- 
tained in the laboratory in Washington, 
D.C., and molted to first form in late Feb- 
ruary and early April 1988. Three ovigerous 
females were dug from burrows on 11 March 
1988: one with carapace length of 22.3 mm 
carried 18 eggs along with several empty 
““capsules,’’ another with cl of 24.9 mm, 20 
eggs, and the third with cl of 25.5 mm, 35 
eggs. All of the eggs were about 2 mm in 
diameter. The two juvenile females collect- 
ed on 18 March 1987 have carapace lengths 
of 11.9 and 12.1 mm, and the two juvenile 
males, 11.4 and 11.9 mm. 

Ecological notes.— This crayfish has been 
found only in complex burrows consisting 
of branching galleries, several of which, ex- 
cept in dry seasons, reach the surface, some 
of their openings marked by rather crudely 
constructed turrets. Where Fallicambarus 
(C.) gilpini has been collected in the same 
locality with F. (C.) fodiens, the burrows of 
the former were frequently, if not usually, 
situated higher on the seepage slope, sug- 
gesting that like the partitioning of a habitat 
in South Carolina by Cambarus (J.) caro- 
linus (Erichson, 1846) and Distocambarus 
(Fitzcambarus) carlsoni Hobbs, 1983 (see 


690 


latter, page 437), gi/pini might prefer areas 
in which the groundwater is moving, where- 
as fodiens more frequently occurs in areas 
in which the water is more static. In general, 
the burrows of gi/pini are more complex, 
exhibiting more subhorizontal galleries than 
do those of fodiens that we have excavated 
in Arkansas. 

Relationships.— Fallicambarus (C.) gil- 
pini has its closest affinities with F. (C.) cae- 
sius. In addition to the many features the 
two species share in common, including 
being the only typically blue members of 
the genus, they are the only ones that lack 
a ventrolateral row of tubercles on the me- 
rus of the first cheliped. The most readily 
observed features that distinguish the two 
species are the absence of tubercles on the 
mesial surface of the dactyl of the chela and 
the presence of a distolateral spine on the 
mesial ramus of the uropod in F. (C.) gi/pini. 

Etymology.—This crayfish is named in 
honor of our mutual friend Robert H. Gil- 
pin, of Cumberland, Maryland, in token of 
his interest and assistance in collecting much 
of the material we have from the eastern 
part of Arkansas. 


Acknowledgments 


For their assistance in collecting some of 
the crayfishes on which this study is based, 
thanks are extended to Robert H. Gilpin of 
Cumberland, Maryland; Michael F. Kear- 
ney of Louisiana State University, Brian F. 
Kensley of the Smithsonian Institution: 
Raymond F. Jezerinac of Ohio State Uni- 
versity, Newark; M. James Norrocky of 
Vickery, Ohio; John Dempsey of Arkansas 
High School, and the following students at 
Southern Arkansas University: Wanda 
Hobson, Daryl Koym, Elaine Laird, Beth 
Lovorn, Patrick Robison, and Linda Tate. 
For the loan of specimens from the Illinois 
Natural History Survey, we are grateful to 
Lawrence M. Page. For their criticisms of 
the manuscript we are indebted to Thomas 
E. Bowman of the Smithsonian Institution, 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Joseph F. Fitzpatrick, Jr., of the University 
of South Alabama, and H. H. Hobbs III, 
Wittenberg University. The Southern Ar- 
kansas University Faculty Research Fund 
provided travel funds to H. W. Robison to 
collect crayfishes. 


Literature Cited 


Albaugh, Douglas W. 1973. Life histories of the cray- 

fishes Procambarus acutus and Procambarus 

hinei in Texas. Ph.D. Dissertation, Texas A&M 

University, xii + 135 pp. 

, & Joe B. Black. 1973. A new crawfish of the 

genus Cambarellus from Texas, with new Texas 

distributional records for the genus (Decapoda, 

Astacidae). —Southwestern Naturalist 18(2):177-— 

185. 

Andolshek, Margaret D.,& Horton H. Hobbs, Jr. 1986. 
The entocytherid ostracod fauna of southeastern 
Georgia. — Smithsonian Contributions to Zool- 
ogy 424:3 + 43 pp., 20 figs. 

Andrews, E. A. 1904. Breeding habits of crayfish.— 
American Naturalist 38(447):165—206. 

Becker, C. Dale, Robert G. Genoway, & J. A. Merrill. 
1975. Resistance of a northwestern crayfish, 
Pacifastacus leniusculus (Dana), to elevated 
temperatures.— Transactions of the American 
Fisheries Society 1042(2):374-387. 

Bell, Ross T. 1971. Handbook of the Malacostraca 
of Vermont and neighboring regions (crayfish, 
sowbugs and their relatives), Privately printed, 
Vermont, 65 pp. 

Berrill, Michael. 1978. Distribution and ecology of 
crayfish in the Kawartha Lakes region of south- 
ern Ontario.— Canadian Journal of Zoology 56: 
166-177. 

Berrill, Michael, & Brian Chenoweth. 1982. The bur- 
rowing ability of nonburrowing crayfish. — 
American Midland Naturalist 108(1):199-201. 

Black, Joe B. 1967. Anew crawfish of the genus Cam- 
barus from Southwest Louisiana (Decapoda; 
Astacidae).— Proceedings of the Biological So- 

ciety of Washington 80:173-178. 

. 1969. A new crawfish of the genus Hobbseus 

from Mississippi (Decapoda, Astacidae).— Pro- 

ceedings of the Biological Society of Washington 
82(14):193-199. 

Bouchard, Raymond W. 1972. A contribution to the 
knowledge of Tennessee crayfish. Ph.D- Disser- 
tation, University of Tennessee, Knoxville, vi 

+ 113 pp. 

. 1976a. Crayfishes and shrimps. Pp. 13-20 in 

Herbert Boschung, ed., Endangered and threat- 


VOLUME 102, NUMBER 3 


ened plants and animals of Alabama. Alabama 

Museum of Natural History, 2. 

1976b. Geography and ecology of crayfishes 
of the Cumberland Plateau and Cumberland 
Mountains, Kentucky, Virginia, Tennessee, 
Georgia and Alabama, Part II: The genera Fal- 
licambarus and Cambarus. Pp. 585-605 in James 
W. Avault, Jr., ed.. Freshwater crayfish. Baton 
Rouge, Louisiana, Louisiana State University 
Division of Continuing Education. 

1978. Morphology of the mandible in hol- 
arctic crayfishes (Decapoda: Astacidae and 
Cambaridae): Ecological and phylogenetic im- 
plications. Pp. 425-452 in Ossi V. Lindqvist, 
ed., Freshwater crayfish. University of Kuopio, 
Kuopio, Finland. 

1980. Morphology of the mandible in hol- 
arctic crayfishes (Decapoda: Astacidae and 
Cambaridae): Ecological and phylogenetic im- 
plications. [Reprinted, with same pagination 
from Bouchard 1978.] 

. & Henry W. Robison. 1981. An inventory 

of the decapod crustaceans (crayfishes and 

shrimps) of Arkansas with a discussion of their 
habitats. — Proceedings of the Arkansas Acade- 

my of Science 34:22-30. 

Bousfield, E. L. 1979. Crustacea. Pp. 291-294 in H. 
V. Danks, ed., Canada and its insect fauna.— 
Memoirs of the Entomological Society of Can- 
ada, 108. 

Bouvier, E.L. 1940. Faune de France, 37, Décapodes 
marcheurs. Paul LeChevalier et Fils, Paris, 404 
pp. 

Bovbjerg, Richard V. 1952. Comparative ecology and 
physiology of the crayfish Orconectes propin- 

quus and Cambarus fodiens.— Physiological Zo- 

ology 25(1):34—-56. 

1970. Ecological isolation and competitive 
exclusion in two crayfish (Orconectes virilis and 
Orconectes immunis).— Ecology 51(2):225-236. 

Bowler, K. 1963. A study of the factors involved in 
acclimatization to temperature and death at high 
temperatures in Astacus p. pallipes.—Journal of 
Cellular and Comparative Physiology 62:119- 
132. 

Bnmley, C. S. 1938. The insects of North Carolina, 
being a list of the insects of North Carolina and 
their close relatives. North Carolina Depart- 
ment of Agriculture, Raleigh, 560 pp. 

Burr, Brooks M., & Horton H. Hobbs, Jr. 1984. Ad- 
ditions to the crayfish fauna of Kentucky, with 
new locality records for Cambarellus shu- 
feldtii.—Transactions of the Kentucky Acade- 
my of Science 45(1—2):14—18. 

Cahn, Alvin Robert. 1915. An ecological survey of 
the Wingra Springs region, near Madison, Wis- 
consin, with special reference to its ornitholo- 


691 


gy.— Bulletin of the Wisconsin Natural History 
Society 13(3):123-177. 

Caine, Edsel A. 1974. Adaptations to species-specific 
habitats by epigean and troglobitic crayfishes 
(Decapoda: Astacidae). Ph.D. Dissertation, 
Florida State University, iii + 85 pp. 

Clark, Clarence F., & Rendell Rhoades. 1979. Notes 
on the crayfishes of Auglaize County, Ohio, 
1941-1943.—Ohio Journal of Science 79(5):236— 
239, 1 fig. 

Cooper, John E., & Ray E. Ashton, Jr. 1985. The 
Necturus lewisi study: Introduction, selected lit- 
erature review, and comments on the hydrologic 
units and their faunas.— Brimleyana 10:1-12. 

Cottle, T. J. 1863. On the two species of Astacus 
found in upper Canada.— Canadian Journal of 
Industry, Science, and Arts, new series 45:216— 
219. 

Crawford, E. A. 1959. Five new ostracods of the ge- 
nus Entocythere (Ostracoda, Cytheridae) from 
South Carolina.— University of South Carolina 
Publications, Biology, Series III 2(4):149-189. 

Creaser, Edwin P. 1931. The Michigan decapod crus- 

taceans.— Papers of the Michigan Academy of 

Science, Arts, and Letters 13:257-—276. 

1932. The decapod crustaceans of Wiscon- 
sin.— Transactions of the Wisconsin Academy 
of Sciences, Arts, and Letters 27:321-338. 

. 1934. Age, growth, and sex ratios in the cray- 

fish Faxonius propinquus.— Papers of the Mich- 

igan Academy of Science, Arts, and Letters 19: 

581-585. 

Crenshaw, Larry I., Daniel E. Lemons, & Karen E. 
Russo. 1980. Crayfish behavior in relation to 
power plants. Pp. 241—259 in Charles H. Hocutt 
et al., ed., Power plants, effects on fish and shell- 
fish behavior. Academic Press, New York. 

Crocker, Denton W. 1957. The crayfishes of New 

York State (Decapoda, Astacidae).— New York 

State Museum and Science Service Bulletin 355: 

1-97. 

, & David W. Barr. 1968. Handbook of the 

crayfishes of Ontario. Royal Ontario Museum, 

University of Toronto Press, Tronto, xiii + 158 

pp. 

Cummins, H. 1921. Spring migration in the cray- 
fish.—Twenty-sixth Annual Meeting of the 
North American Benthological Society 40:28- 
30. 

Eberly, William R. 1954. The use of crayfish to il- 

lustrate some ecological principles.— Turtox 

News 32(3):58-60. 

. 1955. Summary ofthe distribution of Indiana 

crayfishes, including new state and county rec- 

ords.— Proceedings of the Indiana Academy of 

Science 64:28 1-283. 

Engle, Earl Theron. 1926. Crayfishes of the genus 


692 
Cambarus in Nebraska and eastern Colorado. — 
Bulletin of the Bureau of Fisheries 42:87—104. 
Erichson, W. F. 1846. Uebersicht der Arten der Gat- 


tung Astacus.—Archiv fiir Naturgeschichte, 
Berlin 12(1):86—103. 
Faxon, Walter. 1884. Descriptions of new species of 
Cambarus, to which is added a synonymical list 
of the known species of Cambarus and Asta- 
cus. — Proceedings of the American Academy of 
Arts and Sciences 20:107-158. 
. 1885. A revision of the Astacidae, Part I. The 
genera Cambarus and Astacus.— Memoirs of the 
Museum of Comparative Zodlogy at Harvard 
College 10(4):vi + 186 pp. 

1890. Notes on North American crayfishes, 
Family Astacidae.— Proceedings of the United 
States National Museum 12(785):619-634. 

1898. Observations on the Astacidae in the 
United States National Museum and in the Mu- 
seum of Comparative Zoology, with descrip- 
tions of new species. — Proceedings of the United 
States National Museum 20(1136):643-694. 

1914. Notes on the crayfishes in the United 
States National Museum and the Museum of 
Comparative Zodlogy with descriptions of new 
species and subspecies to which is appended a 
catalogue of the known species and subspe- 
cies.— Memoirs of the Museum of Comparative 
Zoology at Harvard College 40(8):35 1-427. 
Feinberg, Harold S. 1971. A catalogue of type spec- 

imens in the Department of Living Inverte- 
brates, American Museum of Natural History, 
New York, New York, U.S.A. Phylum Arthrop- 
oda; Class Crustacea, Part One: Order Decap- 
oda. Department of Living Invertebrates, 
American Museum of Natural History, ii + 75 
pp. 
Fitzpatrick, J. F., Jr. 1983. How to know the fresh- 
water Crustacea. Dubuque, Iowa, Wm. C. Brown 
Company, 227 pp. 
. 1986. The Pre-Pliocene Tennessee River and 
its bearing on crawfish distribution (Decapoda: 
Cambaridae).—Brimleyana 12:123-146. 
. 1987. Fallicambarus (Creaserinus) burrisi and 
F. (C.) gordoni, two new burrowing crawfishes 
associated with pitcher plant bogs in Mississippi 
and Alabama (Decapoda: Cambaridae).—Pro- 
ceedings of the Biological Society of Washington 
100(3):433-446. 
, & James F. Payne. 1968. A new genus and 
species of crawfish from the southeastern United 
States (Decapoda, Astacidae).— Proceedings of 
the Biological Society of Washington 81(2):11- 
ae: 
Fowler, Henry W. 1912. The Crustacea of New Jer- 
sey. Pp. 29-650 in Annual Report of the New 
Jersey State Museum for 1911. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Girard, Charles. 1852. A revision of the North Amer- 
ican Astaci with observations on their habits 
and geographical distribution. — Proceedings of 
the Academy of Natural Sciences of Philadel- 
phia 6:87-91. 

Gladwell, R. T., K. Bowler, & C. J. Duncan. 1975. 
Heat death in the crayfish Austropotamobius 
pallipes. lon movements and their effects on ex- 
citable tissues during heat death.—Journal of 
Thermal Biology 1:79-94. 

Grow, Linda. 1981. Burrowing behaviour in the cray- 
fish Cambarus diogenes diogenes Girard.—An- 
imal Behaviour 29:351-—356. 

———, & Henry Merchant. 1980. The burrow habit 
of the crayfish, Cambarus diogenes diogenes 
(Girard).— American Midland Naturalist 103(2): 

231-237. 

Hagen, Hermann A. 1870. Monograph of the North 
American Astacidae.—Illustrated Catalogue of 
the Museum of Comparative ZoGlogy at Har- 
vard College 3:viii + 109 pp. 

Harris, J. Arthur. 1901. Notes on the habits of Cam- 

barus immunis Hagen.—American Naturalist 

35:187-191. 

1903a. An ecological catalogue of the cray- 
fishes belonging to the genus Cambarus.— Kan- 
sas University Science Bulletin 2(3):51-187. 

. 1903b. The habits of Cambarus.— American 
Naturalist 37(441):601-608. 

Hart, C. W., Jr. 1959. The ostracods of the genus 
Entocythere from the lower Chattahoochee-Flint 
Basin.— Bulletin of the Florida State Museum 
4(6):193-204. 

Hart, Dabney G., & C. W. Hart, Jr. 1974. The os- 
tracod family Entocytheridae.—Academy of 
Natural Sciences of Philadelphia Monograph 18: 
1x + 239 pp. 

Hay, William P. 1891. The Crustacea of Indiana.— 

Proceedings of the Indiana Academy of Science 

1891:147-150. 

. 1896. The crawfishes of the State of Indiana. 

Pp. 475-507 in 20th Annual Report of the De- 

partment of Geology and Natural Resources of 

Indiana. 

1899. Synopses of North American inver- 
tebrates, VI. The Astacidae of North Ameri- 
ca.—American Naturalist 33(396):957-966. 

. 1904. On the habits of Cambarus uhleri Fax- 

on.—Proceedings of the Biological Society of 

Washington 17(33):165-168. 

1919. The crawfishes. Jn B. W. Evermann 
and H. W. Clark, The crustaceans of Lake Max- 
inkuckee.— Proceedings of the Indiana Acade- 
my of Science (1919):230—235. (Also published 
in Indiana Department of Conservation Publi- 
cation 2(7):83-86 (1920).) 

, & C. A. Shore. 1918. The decapod crusta- 


VOLUME 102, NUMBER 3 


ceans of Beaufort, N. C., and the surrounding 
region.— Bulletin of the Bureau of Fisheries 35: 
371-475. 
Hobbs, Horton H., Jr. 1941. Three new Florida cray- 
fishes of the subgenus Cambarus (Decapoda, 
Astacidae).—American Midland Naturalist 
26(1):110-121. 
. 1942. The crayfishes of Florida.— University 
of Florida Publications, Biological Science Se- 
ries 3(2):v + 179 pp. 
. 1948. Anew crayfish of the genus Cambarus 
from Texas, with notes on the distribution of 
Cambarus fodiens (Cottle). — Proceedings of the 
United States National Museum 98(3230):223- 
231: 
. 1955. Anew crayfish of the genus Cambarus 
from Mississippi.— Proceedings of the Biologi- 
cal Society of Washington 68(15)95-100. 
. 1959. Pp. 883-901 in W. T. Edmondson, ed., 
Fresh-water biology, Second edition. John Wi- 
ley & Sons, New York. 
. 1962. Notes on the affinities of the members 
of the Blandingii section of the crayfish genus 
Procambarus (Decapoda, Astacidae).— Tulane 
Studies in Zoology 9(5):273-293. 

1966a. An illustrated key to the species of 
the genus Ankylocythere with a description of a 
new species from Louisiana (Ostracoda, Ento- 
cytheridae).— Proceedings of the Louisiana 
Academy of Sciences 29:67-75. 

1966b. A new crayfish from Alabama with 
observations on the Cristatus Section of the ge- 
nus Cambarus (Decapoda, Astacidae).—Pro- 
ceedings of the Biological Society of Washington 
79(15):109-116. 

1967. A new crayfish from Alabama Caves 
with notes on the origin of the genera Orconectes 
and Cambarus (Decapoda: Astacidae).—Pro- 
ceedings of the United States National Museum 
123(3621):1-17. 

1968. Crustacea: Malacostraca. Pp. K1-K36 
in Fred K. Parrish, ed., Keys to water quality 
indicative organisms (Southeastern United 
States). Federal Water Pollution Control 
Administration, Department of the Interior. 

. 1969a. On the distribution and phylogeny of 
the crayfish genus Cambarus. Pp. 93-178 in Per- 
ry C. Holt, Richard L. Hoffman, & C. Willard 
Hart, Jr., eds., The distributional history of the 
biota of the southern Appalachians, Part I: In- 
vertebrates. Virginia Polytechnic Institute, Re- 
search Division Monograph 1. 

. 1969b. Two new species of the crayfish genus 
Procambarus (Decapoda, Astacidae) with keys 
to the members of the Spiculifer group. —Pro- 
ceedings of the Biological Society of Washington 
83(24):329-348. 


693 


1972. Crayfishes (Astacidae) of North and 
Middle America, identification manual 9: x + 
173 pp. Jn Biota of freshwater ecosystems. 
United States Environmental Protection Agen- 
cy, Water Pollution Research Control Series. 

1973. New species and relationships of the 
members of the genus Fallicambarus.—Pro- 
ceedings of the Biological Society of Washington 
86(40):46 1-482. 

1974a. Synopsis of the families and genera 
of crayfishes (Crustacea, Decapoda).—Smith- 
sonian Contributions to Zoology 164:1-—32. 

1974b. A checklist of the North and Middle 

American crayfishes (Decapoda: Astacidae and 

Cambaridae).—Smithsonian Contributions to 

Zoology 166:1ii + 161 pp. 

1975. New crayfishes (Decapoda: Cambari- 
dae) from the southern United States and Mex- 
ico.—Smithsonian Contributions to Zoology 
201:1-34. 

. 1976. Adaptations and convergence in North 

American crayfishes. Pp. 541-551 in James W. 

Avault, Jr., ed., Freshwater crayfish. Papers from 

the Second International Symposium on Fresh- 

water Crayfish, Baton Rouge, Louisiana, Divi- 

sion of Continuing Education, Louisiana State 

University. 

. 1979. Anewcrayfish from the Ouachita Riv- 

er basin in Arkansas (Decapoda: Cambari- 

dae).— Proceedings of the Biological Society of 

Washington 92(4):804-811. 

. 1981. The crayfishes of Georgia.—Smithson- 

ian Contributions to Zoology 318:viii + 549 pp. 

1983. Distocambarus (Fitzcambarus) carl- 
soni, anew subgenus and species of crayfish (De- 
capoda: Cambaridae) from South Carolina. — 
Proceedings of the Biological Society of Wash- 
ington 96(3):429-439. 

1989. An illustrated checklist of the Amer- 
ican crayfishes (Decapoda: Astacidae, Camba- 
ridae, and Parastacidae). —Smithsonian Contri- 
butions to Zoology (in press). 

—., & Thomas C. Barr, Jr. 1960. The origins and 
affinities of the troglobitic crayfishes of North 
American (Decapoda, Astacidae), I. The genus 
Cambarus.—American Midland Naturalist 
64(1):12-33. 

——., & 1972. Origins and affinities of the 

troglobitic crayfishes of North America (Decap- 

oda: Astacidae), II. Genus Orconectes.—Smith- 

sonian Contributions to Zoology 105:1i1 + 84 

pp. 

, & J. F. Fitzpatrick, Jr. 1970. A new crayfish 

of the genus Fallicambarus from Tennessee (De- 

capoda, Astacidae).— Proceedings of the Bio- 
logical Society of Washington 82(64):829-836. 

——,, & Edward T. Hall, Jr. 1974. Crayfishes (De- 


694 


capoda: Astacidae). Pp. 195-214 in C. W. Hart, 

Jr., and S. L. H. Fuller, eds., Pollution ecology 

of freshwater invertebrates. Academic Press, 

New York. 

,&C. W. Hart, Jr. 1959. The freshwater deca- 

pod crustaceans of the Apalachicola drainage 

system in Florida, southern Alabama, and 

Georgia. — Bulletin of the Florida State Museum 

4(5):145-191. 

—, & H. H. Hobbs III. 1970. New entocytherid 

ostracods with a key to the genera of the subfam- 

ily Entocytherinae.—Smithsonian Contribu- 

tions to Zoology 47:1-19. 

, & Lewis J. Marchand. 1943. A contribution 

toward a knowledge of the crayfishes of the Reel- 

foot Lake area.—Journal of the Tennessee 

Academy of Science 18(1):6-35. 

, & Daniel J. Peters. 1977. The entocytherid 

ostracods of North Carolina.—Smithsonian 

Contributions to Zoology 247:iv + 73 pp. 

—., & Henry W. Robison. 1985. A new burrow- 
ing crayfish (Decapoda: Cambaridae) from 
southwestern Arkansas. — Proceedings of the Bi- 
ological Society of Washington 98(4):1035-1041. 

—., & Mike Whiteman. 1987. A new economi- 
cally important crayfish (Decapoda: Cambari- 
dae) from the Neches River Basin, Texas, with 
a key to the subgenus Fallicambarus.—Pro- 
ceedings of the Biological Society of Washington 
100(2):403-411. 

Hobbs III, H. H. 1969. Studies in ecological and host 

specificity in entocytherid ostracods (Ostracoda: 

Entocytheridae). Ph.D. Thesis, Mississippi State 

University, vii + 93 pp. 

, & Joan P. Jass. 1988. The crayfishes and 

shrimp of Wisconsin. Miwaukee Public Mu- 

seum, Milwaukee, vili + 178 pp. 

, James H. Thorp, & Gilbert E. Anderson. 1976. 

The freshwater decapod crustaceans (Palae- 

monidae, Cambaridae) of the Savannah River 

Plant, South Carolina. Savannah River Plant, 

National Environment Research Park Program, 

63 pp. 

Hoffman, Richard L. 1963. A revision of the North 
American annelid worms of the genus Cam- 
barincola (Oligochaeta: Branchiobdellidae).— 
Proceedings of the United States National Mu- 
seum | 14(3470):27 1-371. 

Holt, Perry C. 1973. Epigean branchiobdellids (An- 
nelida: Clitellata) from Florida.— Proceedings of 
the Biological Society of Washington 86(7):79- 
104. 

Huner, Jay V. 1977. Ditch bugs, good bait . . . good 

to eat!— Bayou State Sportsman 2(2):10-13. 

1978. Crawfish population dynamics as they 
affect production in several small, open com- 
mercial crawfish ponds in Louisiana.— Ninth 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Annual Meeting, World Mariculture Society, pp. 
619-640. : 

——.,, & J. E. Barr. 1981. Red swamp crawfish: 

Biology and exploitation. Sea Grant Publication 

Number LSU-T-80-001. Baton Rouge: Louisi- 

ana State University Center for Wetland Re- 

sources, xi + 148 pp. 

ee Be 1981. Red swamp crawfish: Bi- 

ology and exploitation. Sea Grant Publication 

Number LSU-T-80-001. Baton Rouge: Louisi- 

ana State University Center for Wetland Re- 

sources, xi + 148 pp. (Revised edition, 1984). 

Huner, J. V., Samuel P. Meyers, & James W. Avault, 
Jr. 1976. Response and growth of freshwater 
crawfish to an extruded, water-stable diet. Pp. 
149-157 in James W. Avault, Jr., ed., Fresh- 

’ water crayfish. Baton Rouge, Louisiana, Divi- 
sion of Continuing Education, Louisiana State 
University. 

Huntsman, A.G. 1915. The fresh-water Malacostra- 
ca of Ontario.— Contributions to Canadian Bi- 
ology 1911-1914:145-163. 

Jaspers, Edmonde, & James W. Avault, Jr. 1969. En- 
vironmental conditions in burrows and ponds 
of the red swamp crawfish, Procambarus clarki 
(Girard), near Baton Rouge, Louisiana.—Pro- 
ceedings of the 23rd Annual Conference of the 
Southeastern Association of Game and Fish 
Commissioners, pp. 634-648. 

Jezerinac, Raymond F. 1983. Possible correlations 

of present distributions of Ohio crayfishes (De- 

capoda: Cambaridae) with Teays-age drainages. 

In Anonymous, Teays-age drainage effects on 

present distributional patterns of Ohio biota— 

An Ohio biogeography conference [Abstracts and 

Supplements].—Ohio Biological Survey Infor- 

mative Circular 11:4—S. 

1986. Endangered and threatened crayfishes 
(Decapoda: Cambaridae) of Ohio.—Ohio Jour- 
nal of Science 86(4):177—180. 

, & G. Whitney Stocker. 1987. Fallicambarus 

(Creaserinus) fodiens (Cottle, 1863) (Decapoda: 

Cambaridae) in West Virginia: A new state rec- 

ord.—Ohio Journal of Science 87(1):46—47. 

Jezerinac, Raymond F., & Roger F. Thoma. 1984. 
An illustrated key to the Ohio Cambarus and 
Fallicambarus (Decapoda: Cambaridae) with 
comments and a new subspecies record. — Ohio 
Journal of Science 84(3):120-125. 

Judd, W.W. 1968. Crayfish in the vicinity of London, 
Ontario.— National Museum of Canada, Nat- 
ural History Papers 41:1-9. 

Kiley, Ann, & Clarence F. Dineen. 1982. Crayfishes 
of Lake Wawasee. — Proceedings of the Indiana 
Academy of Science 91:211-212. 

Lahser, Carl W., Jr. 1976. Epizodites of crayfish I. 
Ectocommensals and parasites of crayfish of 


VOLUME 102, NUMBER 3 


Brazos County, Texas. Pp. 277-285 in James 
W. Avault, Jr., ed., Freshwater crayfish. Papers 
from the Second International Symposium on 
Freshwater Crayfish, Baton Rouge, Louisiana: 
Division of Continuing Education, Louisiana 
State University. 

Lake, P.S. 1977. Pholeteros— The faunal assemblage 
found in crayfish burrows.— Australian Society 
of Limnology Newsletter 15(1):57-60. 

Lawton, Steven M. 1979. A taxonomic and distri- 
butional study of the crayfishes (Decapoda 
Cambaridae) of West Virginia with diagnostic 
keys to species of the genera Cambarus and Or- 
conectes. Ph.D. Thesis, Marshall University, 107 
+ 8 unnumbered pp. 

Lyle, Clay. 1937. The crawfishes of Mississippi, with 

special reference to the biology and control of 

destructive species. Ph.D. Thesis, Iowa State 

College, 140 pp. 

1938. The crawfishes of Mississippi, with 
special reference to the biology and control of 
destructive species. — Iowa State College Journal 

of Science 13:75-77. 

Mansell, Barry W. 1989. The occurrence of the cray- 
fish Fallicambarus fodiens in Florida.— Florida 
Scientist (in press). 

Maude, S. H., & D. D. Williams. 1983. Behavior of 
crayfish in water currents: Hydrodynamics of 
eight species with reference to their distribution 
patterns in southern Ontario.— Canadian Jour- 
nal of Fisheries and Aquatic Sciences 40(1):68- 
V4: 

McMahon, B. R., & P. R. H. Wilkes. 1983. Emer- 
gence responses and aerial ventilation in nor- 
moxic and hypoxic crayfish Orconectes rusti- 
cus.— Physiological Zoology 56(2):133-141. 

Meredith, W. G., & F. J. Schwartz. 1959. The cray- 
fishes of Maryland.— Maryland Tidewater News. 

15(1) Supplement 12:1-2. 

1960. Maryland crayfishes.— Maryland De- 
partment of Research and Education, Educa- 
tional Series 46:32 pp. 

1962. The crayfishes of Maryland.— Mary- 
land Tidewater News 15(1), Supplement 12:2 
pp. 

Miller, George C. 1965. Western North American 
crawfishes (Pacifastacus) in brackish water en- 
vironments.— Research Briefs, Oregon Fish 
Commission | 1(1):42—50. 

Mobberly, William C., Jr. 1965. Lethal effect of tem- 
perature on the crawfish Faxonella clypeata.— 
Proceedings of the Louisiana Academy of Sci- 
ences 28:45-51. 

Momot, Walter T., Howard Gowing, & Patricia D. 
Jones. 1978. The dynamics of crayfish and 
their role in ecosystems.—American Midland 
Naturalist 99(1):10-35. 


695 


Norrocky, M. James. 1983. Untitled note on col- 
lecting burrowing crayfishes. — International As- 
sociation of Astacology Newsletter 6(2):3. 

1984. Burrowing crayfish trap.—Ohio Jour- 
nal of Science 84(1):65-66. 

Ortmann, A. E. 1902. The geographical distribution 

of fresh-water decapods and its bearing upon 

ancient geography. — Proceedings of the Amer- 
ican Philosophical Society 41(171):267—-400. 

1905. The mutual affinities of the species of 
the genus Cambarus, and their dispersal over 
the United States.— Proceedings of the Ameri- 
can Philosophical Society 44(180):9 1-136. 

1907. Grabende Krebse in Nordamerika.— 
Aus der Natur (Zeitschrift ftir alle Natur- 
freunde), II Jahrgang, 1906/7, Heft 23:705-711, 
743-748. 

Osburn, R. C., & E. B. Williamson. 1898. The cray- 
fish of Ohio. Sixth Annual Report of the Ohio 
State Academy of Science, p. 21. 

Page, Lawrence M. 1974. Aquatic Malacostraca re- 

corded for Illinois, with notes on their distri- 

butions and habitats within the state.—Trans- 
actions of the Illinois State Academy of Science 
67(1):89-104. 

1985. The crayfishes and shrimps (Decapo- 
da) of Illinois. —Illinois Natural History Survey 
Bulletin 33(4):xi1 + 335-448. 

Payne, J. F.,& L.A. Riley. 1974. Notes on crayfishes 
from the Chickasaw Basin. — Journal of the Ten- 
nessee Academy of Science 49(4):125-128. 

Pearse, A. S. 1910a. The crawfishes of Michigan.— 

Michigan State Biological Survey 1:9-22. 

. 1910b. A preliminary list of the Crustacea of 

Michigan.— Twelfth Report of the Michigan 

Academy of Science, 1910, pp. 68-76. 

. 1911. Notes on Michigan Crustacea. I. Thir- 

teenth Report of the Michigan Academy of Sci- 

ence, 1911, p. 130. 

Penn, George H., Jr. 1941. Preliminary report of a 
survey of the crawfishes of Louisiana. Abstracts 
of Papers Presented at the Eighty-eighth Annual 
Meeting of the New Orleans Academy of Sci- 
ences, p. 8. 

1943. A study of the life history of the Lou- 
isiana Red-Crawfish, Cambarus clarkii Gi- 
rard.— Ecology 24(1):1-18. 

. 1953. Anew burrowing crawfish of the genus 

Procambarus from Louisiana and Mississippi 

(Decapoda, Astacidae).— Tulane Studies in Zo- 

ology 1(6):71-76. 

1955. A new Cambarus of the Diogenes Sec- 
tion from North Louisiana (Decapoda, Astaci- 
dae).— Tulane Studies in Zoology 3(4):73-81. 

1959. An illustrated key to the crawfishes of 
Louisiana with a summary of their distribution 


696 


within the state (Decapoda, Astacidae).—Tu- 
lane Studies in Zoology 7(1):3—20. 

— ., & Horton H. Hobbs, Jr. 1958. Acontribution 
toward a knowledge of the crayfishes of Texas 
(Decapoda, Astacidae).— Texas Journal of Sci- 
ence 10(4):452-483. 

—., & Guy Marlow. 1959. The genus Cambarus 
in Louisiana.—American Midland Naturalist 
61(1):191-203. 

Pennak, Robert W. 1953. Fresh-water invertebrates 
of the United States. The Ronald Press Com- 
pany, New York. 1x + 769 pp. 

Peters, Daniel J. 1971. Entocytherid ostracods of the 

lower James-York Peninsula, Virginia.— Vir- 

ginia Journal of Science 22(3):100. 

1974. The ecological and geographical dis- 
tribution of some Virginia ostracods.—ASB 
Bulletin 21(2):74. 

. 1975. The entocytherid ostracod fauna of the 

James and York river basins with a description 

of a new member of the genus Entocythere. — 

Virginia Polytechnic Institute and University, 

Research Division Bulletin 93:11 + 50 pp. 

Phillips, Gary S. 1980. The decapod crustaceans of 
Iowa.— Proceedings of the Iowa Academy of 
Science 87(3):81-—95. 

Pickett, Joseph F., Sr., & Ronald Sloan. 1979. The 
hidden world of the crayfish.—The Conserva- 
tionist [New York] 33(6):22-26. 

Radaj, Richard H. 1978. Key to the Wisconsin cray- 
fish (Astacidae). University of Wisconsin, Wau- 
kesha, Mimeographed, 4 pp. [not seen]. 

Reimer, Rollin D. 1966. Two new species of the ge- 

nus Cambarus from Arkansas (Decapoda, As- 

tacidae).— Tulane Studies in Zoology 13(1):9- 

tS). 

. 1969. A report on the crawfishes (Decapoda, 

Astacidae) of Oklahoma.— Proceedings of the 

Oklahoma Academy of Sciences 48:49-65. 

. 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. 

Smith, Elsie Wayne. 1953. The life history of the 
crawfish Orconectes (Faxonella) clypeatus (Hay) 
(Decapoda, Astacidae).— Tulane Studies in Zo- 
ology 1(7):79-96. 

Spoor, W. A. 1955. Loss and gain of heat-tolerance 
by the crayfish. — Biological Bulletin 108(1):77— 
87. 

Stebbing, Thomas R. R. M. A. 1893. A history of 
Crustacea, Recent Malacostraca. D. Appleton & 
Company, New York, xvii + 466 pp. 

Terman, Max R. 1974. The ecology of crayfish: Some 
observations on behavioral factors.—The Bi- 
ologist 56(1):32-39. 

Thoma, Roger F., & Raymond F. Jezerinac. 1982. 
New distributional records of crayfishes (Cam- 
barus and Fallicambarus) from Ohio including 
a new subspecies record.— Ohio Journal of Sci- 
ence 82(3):136-138. 

Turner, Clarence L. 1926. The crayfishes of Ohio. — 
Ohio Biological Survey Bulletin 3(3):145-195. 

Underwood, Lucien M. 1886. List of the described 

species of fresh water Crustacea from America, 


VOLUME 102, NUMBER 3 


north of Mexico. —Bulletin of the Illinois State 
Laboratory of Natural History 2(5):323-386. 

Walls, Jerry G. 1968. A new Faxonella from north- 

east Louisiana (Decapoda, Astacidae).—Pro- 

ceedings of the Biological Society of Washington 
81(41):413-418. 

. 1985. Distribution and natural history of the 

crawfish Orconectes difficilis (Decapoda: Asta- 

cidae) in Louisiana.—Southwestern Naturalist 
30(2):189-194. 

, & Joe B. Black. 1967. New variations and 

records of the crawfish Procambarus jaculus 

Hobbs (Crustacea, Decapoda, Astacidae).— 

Proceedings of the Louisiana Academy of Sci- 

ences 30:60-62. 

Washburn, Mel. 1953. Mudbugging.—Louisiana 
Conservationist 13(3):5-7, 19. 

Waywell, E. B., & S. Corey. 1970. The presence of 

pteridines in the hypodermis as a taxonomic 

tool in crayfish.— Canadian Journal of Zoology 

48:1462-1464. 

1972. The occurrence and distribution of 
pteridines and purines in crayfish.—Crusta- 
ceana 22(3):294—302. 

Wharton, Charles H. 1978. The natural environ- 
ments of Georgia. Atlanta, Georgia: Depart- 
ment of Natural Resources, 227 pp. 

Wiens, Wayne, & Kenneth B. Armitage. 


LOGI, The 


697 


oxygen consumption of the crayfish Orconectes 
immunis and Orconectes nais in response to 
temperature and oxygen saturation.—Physio- 
logical Zoology 34:39-54. 

Williams, Austin B. 1954. Speciation and distribu- 
tion of the crayfishes of the Ozark Plateaus and 
Ouachita Provinces. — University of Kansas Sci- 
ence Bulletin 36(12):803-918. 

Williams, D. Dudley, N. E. Williams, & H. B. N. Hynes. 
1974. Observations on the life history and bur- 
row construction of the crayfish Cambarus fod- 
iens (Cottle) in a temporary stream in southern 
Ontario.—Canadian Journal of Zoology 52(3): 
365-370. 

Williamson, E. B. 1899. Notes on Ohio Astacidae. — 

Seventh Annual Report of the Ohio State Acad- 

emy of Science, pp. 47-48. 

. 1907. Notes on the crayfish of Wells County, 

Indiana, with description of new species. — 31st 

Annual Report of the Department of Geology 

and Natural Resources, Indiana 1906:749-763. 


(HHH) Department of Invertebrate Zo- 
ology, Smithsonian Institution, Washing- 
ton, D.C. 20560; (HWR) Department of Bi- 
ology, Southern Arkansas University, 
Magnolia, Arkansas 71753. 


PROC. BIOL. SOC. WASH. 
102(3), 1989, pp. 698-700 


SANQUERUS, A REPLACEMENT NAME 
FOR POSIDON HERKLOTS, 1851 
(CRUSTACEA, DECAPODA, PORTUNIDAE) 


Raymond B. Manning 


Abstract. — Sanquerus is proposed as a replacement name for the preoccupied 
portunid genus Posidon Herklots, 1851, and the name is removed from the 
synonymy of Portunus Weber, 1795. Sanquerus is a monotypic genus contain- 
ing only the West African Sanquerus validus (Herklots, 1851). 


Manning & Holthuis (1981:104, 105) 
pointed out that the West African “Portu- 
nus validus shows little similarity to any of 
the many Indo-West Pacific species of the 
genus, and it shows little affinity with any 
of the American species of the genus... .” 
They also enumerated differences between 
Portunus validus and Portunus pelagicus 
(Linnaeus, 1758), the type species of Por- 
tunus. I take this opportunity to remove 
Posidon from the synonymy of Portunus and 
to propose a replacement name for it. 


Sanquerus, new name 
Figs. 1-2 


Posidon Herklots, 1851:3. [Invalid junior 
homonym of Posidon Iliger, 1801 (Crus- 
tacea). Type species: Portunus (Posidon) 
validus Herklots, 1851, by monotypy. 
Gender masculine.] 


Diagnosis. —Size very large, carapace 
widths in adults exceeding 20 centimeters. 
Carapace wide, breadth about two times 


Fig? 1: 


Sanquerus validus (Herklots) (from Milne Edwards 1861, pl. 29, fig. 1). 


VOLUME 102, NUMBER 3 


699 


Fig. 2. Sanquerus validus (Herklots). a, front; b, third maxilliped; c, merus and carpus of cheliped: d, fifth 
leg; e, abdomen; f, gonopod; g, apex of gonopod. (a, b, d—g from male, 151 mm wide, Nigeria, USNM 121034; 
c from male, 61 mm wide, Liberia, USNM 97864) (c, f, g from Manning & Holthuis 1981). 


length. Surface of carapace minutely tuber- 
culate, appearing smooth to the naked eye, 
evenly convex, lacking distinct grooves or 
ridges, except for ridge extending across 
posterior margin between bases of last 
walking legs; posterolateral angles of cara- 
pace unarmed; conspicuous white spot pres- 
ent posterolaterally on each side. Front with 
three pairs of spiniform teeth, including in- 
ner orbitals, median pair largest. Interan- 
tennular projection low, not visible in dor- 
sal view. Anterolateral margin of carapace 
with nine spiniform teeth, lateralmost larg- 
est. Antenna with free access to orbit. Palate 
with longitudinal ridge. Merus of third max- 
illiped lacking produced anterior lobe. Che- 
liped robust; merus with two posterior 
spines, one subdistal, three inner spines, and 
one smaller vental spine distally; carpus with 
inner and outer spine; palm prismatic and 
costate, with proximal spine at articulation 


with wrist, distal dorsal spine, and smaller 
distal spine on inner carina. Merus of fifth 
leg with posterodistal margin unarmed, 
rounded; swimming paddle notched distal- 
ly. Abdomen of male triangular, 5-seg- 
mented, third to fifth segments fused; ter- 
minal segment longer than broad. Male 
pleopod stout, curved laterally, unarmed. 
Etymology. — This name 1s in recognition 
of the enormous contribution made to the 
knowledge of the West African fauna by Mr. 
Robert Sanquer of Vouhé, France, former 
Trawling Master for the Guinean Trawling 
Survey. The gender is masculine. 
Remarks. —Sanquerus is a very distinc- 
tive portunid, easily distinguished from all 
known portunid genera by the smooth, 
unornamented carapace in combination 
with the prismatic and costate chelae. It keys 
to Portunus in Rathbun (1930:13), Crosnier 
(1962:34), Garth & Stephenson (1966:10), 


700 


and to the couplet containing Portunus and 
Scylla in Stephenson (1972:8). 

Sanquerus validus keys to Portunus pe- 
lagicus in Stephenson & Campbell (1959: 
90) and to the couplet containing P. pelagi- 
cus in Crosnier (1962:42). It keys to Por- 
tunus convexus de Haan, 1833 in Stephen- 
son (1972:13) because of the paired white 
spots on the carapace, but that species has 
distinct ornamentation on the carapace and 
only one posterior spine on the merus of the 
cheliped. On morphological grounds it keys 
to the couplet containing Portunus pelagicus 
in Stephenson (1972:15). It differs from 
Portunus pelagicus in numerous features, 
some of which were mentioned by Manning 
& Holthuis (1981:105): the carapace lacks 
surface sculpture and is minutely rather than 
distinctly tuberculate; the submedian fron- 
tal teeth are the largest of the frontal teeth; 
the interantennular spine is not visible in 
dorsal view; the third maxilliped extends far 
less forward; the cheliped is more massive, 
and the merus 1s ornamented with two strong 
posterior spines; the abdomen of the male 
is broader; and the gonopod is much stout- 
er. As in P. pelagicus, the dactylus of the 
fifth leg is notched distally, not produced 
into a spine, and the anterior eight antero- 
lateral teeth of the carapace are not alter- 
nately large and small. 

Sanquerus resembles the Indo-West Pa- 
cific genus Scy//la de Haan, 1833 in having 
a relatively smooth carapace, but differs 
from Scy/la in having the chelae prismatic 
and costate; in Scy//a the chelae are robust 
and smooth (Stephenson & Campbell 1960: 
111; Stephenson 1972:8, 44). 

In frontal view, the carapace of Sanquerus 
shows very low paired swollen prominences 
on the protogastric, mesogastric, and me- 
sobranchial regions, and distinct but small 
paired branchial lobes. 

Sanquerus validus is a commercial species 
occurring off the West African coast, from 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY CF WASHINGTON 


Senegal to Angola. References to the species 
may be found in Monod (1956:196) and 
Manning & Holthuis (1981:103). 


Acknowledgments 


I thank Fenner A. Chace, Jr., and Austin 
Williams for reviewing the manuscript. 


Literature Cited 


Crosnier, A. 1962. Crustacés Décapodes, Portuni- 
dae.—Faune de Madagascar 16:1—-154, pls. 1- 
13: 

Garth, J. S., & W. Stephenson. 1966. Brachyura of 
the Pacific coast of America, Brachyrhyncha: 
Portunidae.—Allan Hancock Monographs in 
Marine Biology 1:1-154. 

Herklots, J. A. 1851. Additamenta ad Faunam Car- 
cinologicam Africae occidentalis, sive descrip- 
tiones specierum novarum e crustaceorum or- 
dine, quas in Guinea collegit vir strenuus H.S. 
Pel, praefectus residentiis in littore guineae. 
Lugduni-Batavorum, Leiden, 28 pp., 2 pls. 

Manning, R. B., & L. B. Holthuis. 1981. West African 
brachyuran crabs. —Smithsonian Contributions 
to Zoology 306:x1i + 379 pp. 

Monod, Th. 1956. Hippidea et Brachyura ouest-af- 
ricains.— Mémoires de I’Institut frangais d’Af- 
rique noire 45:1-674. 

Rathbun, M. J. 1930. The cancroid crabs of America 
of the families Euryalidae, Portunidae, Atele- 
cyclidae, Cancridae and Xanthidae.— United 
States National Museum Bulletin 152:xvi + 609 
pp., pls. 1-230. 

Stephenson, W. 1972. An annotated check list and 
key to the Indo-West-Pacific Swimming Crabs 
(Crustacea: Decapoda: Portunidae).— Royal So- 
ciety of New Zealand, Bulletin 10:1-64. 

Stephenson, W., & B. Campbell. 1959. The genus 
Portunus. The Australian portunids (Crustacea: 
Portunidae), IIJ.— Australian Journal of Marine 
and Freshwater Research 10(1):84—124, pls. 1-5. 

=. 2 . 1960. Remaining genera. The Aus- 
tralian portunids (Crustacea: Portunidae), IV.— 
Australian Journal of Marine and Freshwater 
Research 11(1):73-122, pls. 1-6. 


Department of Invertebrate Zoology, Na- 
tional Museum of Natural History, Smith- 
sonian Institution, Washington, D.C. 20560. 


PROC. BIOL. SOC. WASH. 
102(3), 1989, pp. 701-715 


RECTIFICATION OF HALIRAGES REGIS AND 
H. HUXLEYANUS (CRUSTACEA: AMPHIPODA), 
FROM MARINE ANTARTICA, WITH 
DESCRIPTION OF A NEW GENUS, 
AUSTROREGIA 


J. L. Barnard 


Abstract. — Halirages huxleyanus and H. regis are removed to the new ant- 
arctic genus, Austroregia. This leaves Halirages with 6 species entirely confined 
to arctic waters. Austroregia is a perplexing genus because it is furnished with 
calceoli of the same kind found in Chosroes and Gammarellus; a family Gam- 
marellidae is available for these two genera which would be removed from 
such families as Eusiridae, Pontogeneiidae and Calliopiidae, in which the genera 
would have been classified previously. Problems remain on speciation within 


Austroregia. 


Halirages huxleyanus (Bate) and H. regis 
(Stebbing) are improperly placed in Halira- 
ges and are removed to a new genus Aus- 
troregia. Austroregia huxleyana possesses 
distinctive calceoli of a kind also present in 
Gammarellus, Chosroes and Gondogeneia, 
adequate to justify the resurrection of the 
family Gammarellidae within the super- 
family Eusiroidea. This reinforces the dis- 
covery by Lincoln & Hurley (1981:111) that 
both the high arctic and high antarctic con- 
tain taxa with common ancestry in the gam- 
marellid group, a family first raised by 
Bousfield (1977), but soon after merged 
within the family Calliopiidae. 


Gammarellidae, revived 


Diagnosis. —Characterized by a type 6 
calceolus of Lincoln & Hurley (1981) in 
which the proximal element forms a dis- 
crete cup separated from the small 2 to 
3-plate distal element by a second smaller 
cup-shaped element (Fig. 1). Also unique is 
the arrangement of the calceoli in transverse 
rows that extend all around the distal mar- 
gin of the flagellar articles. 


Remarks.—The family Gammarellidae 
was established by Bousfield (1977) to con- 
tain two carinate genera, Gammarellus 
Herbst and Weyprechtia Stuxberg, separat- 
ed from other kinds of pontogeneiids by a 
combination of characters that included a 
well developed accessory flagellum, lanceo- 
late weakly setose third uropods, and lam- 
inar, apically emarginate telson. However, 
the distinction from other eusiroids was far 
from clear-cut and in a later updated and 
revised version of his classification, Bous- 
field (1983) synonymized the Gammarelli- 
dae with the Calliopiidae. 

The present use of calceolus morphology 
and arrangement as a shared apomorphy to 
resurrect the Gammarellidae produces a 
family of quite a different complexion. 
Brought together are four eusiroid genera, 
Gammarellus, Gondogeneia, Chosroes, and 
Austroregia, that would not have been rec- 
ognized as belonging to a natural and in- 
dependent group on the basis of traditional 
morphological characterizations. Thus, 
Gammarellus possesses a well developed 
multiarticulate accessory flagellum which is 
at best small to vestigial in Gondogeneia and 


702 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 1. Calceoli: a, Austroregia huxleyana, holotype “a”; b, Chosroes decoratus. Magnifications unknown; 
courtesy of Dr. Roger J. Lincoln. 


Austroregia and absent in Chosroes. Fur- Chosroesand Gammarellus is weakly emar- 
ther, Gammarellus is the only member hav-__ ginate, but has a short to moderate cleft in 
ing a coxal gill on pereonite 7, a lobate car- Gondogeneia and Austroregia. A lanceolate 
pus on gnathopods 1-2, and facial setae on _setose uropod 3 is shared by all four genera 
the inner plate of maxilla 2. The telson of but is also widespread outside the group. 


VOLUME 102, NUMBER 3 


The single feature uniting the family is the 
type 6 calceolus. This could be treated as 
yet another instance of convergence, as has 
to be argued for many eusiroid characters, 
but in view of the marked complexity of the 
calceolus sensory receptor and its novel dis- 
tribution on the antennal articles, there is 
adequate justification for recognizing the 
group as a separate unit within the Eusi- 
roidea. 

There is a superficial resemblance of the 
gammarellid group to the Pleustidae, but 
the latter family has distinctive labia bear- 
ing outer lobes tilted across fused inner lobes; 
pleustids lack calceoli and further relation- 
ships cannot be struck. The lower lip of Me- 
sopleustes and Chosroes bridges the gap be- 
tween the two groups. 

Families based entirely on calceolar 
structure are very difficult to handle by tax- 
onomists because many species and genera 
that obviously are analogous to the calceo- 
liferous members of various families have 
lost their calceoli. This has been a major 
problem with Crangonyctidae and will be a 
problem with Gammarellidae. Only 18 out 
of 91 families of Gammaridea have one or 
more species with calceoli. Fortunately, 16 
out of the 18 families can be recognized by 
characters other than calceoli. At best the 
two families here mentioned are “ghost 
families” for the identificatory taxonomist 
because not all of their species and genera 
are recognizable by ordinary morphological 
characters. Just as with the Crangonyctidae, 
there may be species lacking calceoli which 
are descendants of an ancestor common to 
the known members of Gammarellidae. For 
the moment, those species, plus all other 
members of Eusiridae, Calliopiidae and 
Pontogenelidae have to remain in a pool 
that is identifiable through laborious keys 
that contain little relationship to the phy- 
logenetic positions of their genera. This is 
another case of the difference between prac- 
tical classification involving the identifica- 
tion of species and genera versus phyloge- 
netic classification where species and genera 


703 


may be arranged on the basis of characters 
not readily available from preserved mu- 
seum specimens. 


Austroregia, new genus 


Type species.—Atylus huxleyanus Bate, 
1862, here selected. 

Etymology.—From ‘“‘austro,” southern, 
and “‘regia,”’ kingly. 

Diagnosis. —Eusiroid-like amphipods 
with type 6 calceolus; body carinate; ros- 
trum small; eyes circular; basal article of 
flagellum on antenna | not elongate; anten- 
na 2 as long as antenna 1; upper lip entire; 
inner lobes of lower lip absent; inner plate 
of maxilla 1 with only six or fewer setae, 
mostly terminal; palps disymmetrical or not 
(type); inner plate of maxilla 2 with only 
medial setae, none facial; outer plate of 
maxilliped of basic form and size; gnatho- 
pods small, carpus shorter than propodus, 
nonlobate, gnathopod | slightly larger than 
2; pereopods 3—4 with nonlobate basis; coxa 
4 excavate or not posteriorly; gills unpleat- 
ed, gill 7 lacking; at least pereopod 7 elon- 
gate; outer rami of uropods 1-2 shortened; 
peduncle of uropod 3 slightly elongate, rami 
extending equally, poorly setose; telson 
elongate, cleft about 10-20 percent of its 
length. 

Composition. — Austroregia huxleyana 
(Bate, 1862) and A. regis (Stebbing, 1914). 

Relationship. —Dijffers from the confa- 
milial genera, Gondogeneia in the presence 
of large dorsal carinae on pereonite 7 and 
pleonites 1—3; Chosroes in the enlarged 
gnathopod 1, broader outer plate of the 
maxilliped and the lack of an anterior lobe 
on the basis of pereopods 3-4; Gamma- 
rellus in the nonlobate carpal articles of 
gnathopods 1-2, lack of coxal gill 7, vestigial 
accessory flagellum, nonpleated gills, ab- 
sence of facial setae on the inner plate of 
maxilla 2, poorly setose inner plate of max- 
illa 1, and cleft telson. 

In addition to the type 6 calceolus, Aus- 
troregia differs from the eusiroid genera, 


704 


Halirages in the fewer setae on the inner 
plate of maxilla 1, the lack of facial setae on 
maxilla 2, and the enlarged gnathopod 1; 
Cleippides in the short carpus of the gnatho- 
pods; Haliragoides in the lack of inner lobes 
on the lower lip, the fewer and nonfacial 
setae of maxillae 1-2 and the regular sized 
uropod 3; Whangarusa Barnard & Kara- 
man (1987), in the carinate body, poorly 
setose maxillae, and lack of inner lobes on 
the lower lip; Paracalliopiella in the elon- 
gate telson, and nonreniform eyes; C/eonar- 
dopsis in the lack of inner lobes on the lower 
lip, and lack of carpal lobes on the gnatho- 
pods; Harcledo in the carinate body, longer 
anterior coxae and weakly cleft telson; 
Amathillopsis in the vestigial accessory fla- 
gellum, nonlobed carpus of the gnathopods, 
slightly enlarged gnathopod 1, fewer setae 
on the inner plate of maxilla 1 and the 
smaller dactyl of the maxilliped; Meteusi- 
roides in the carinate body, nonreniform 
eyes, nonelongate article 1 on the flagellum 
of antenna 1, nonlobate carpus of the 
gnathopods, and poorly cleft telson; from 
Paramphithoe in the broader articles 5—6 of 
the gnathopods, especially the more inflated 
article 6, the nonbifid coxa 4, the enlarged 
gnathopod 1, the confinement of body teeth 
to the dorsal midline and the fewer setae on 
the inner plate of maxilla 1; from Ponto- 
geneia in distinctive calceoli, poorly cleft 
telson, absence of inner lobes on the labium, 
enlarged setae on inner plate of maxilla 2 
which are fully marginal (versus facial) and 
the bluntly rounded anteroventral margins 
of the anterior coxae; and finally, Ca/liopius, 
in the nonlobate carpus of the gnathopods, 
nonreniform eyes, absence of inner lobes on 
the lower lip, and the distinctive calceoli. 
Coxa 4 takes different forms. In juveniles 
it is usually excavate weakly and because in 
some adults it curls outward it seemingly is 
deeply excavate when illustrated without 
flattening. In some large adults this coxa 
assumes a diamond-shape (Fig. 6d). 
Presence of calceoli is vagarious. Gener- 
ally, adults of A. huxleyana bear calceoli but 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


most of A. regis do not. We have in hand 
several adults of A. huxleyana without cal- 
ceoli and Stebbing (1888: pl. 73) shows an 
adult of A. regis with calceoli. This fits the 
vagarious occurrence of calceoli classically 
known in many species of amphipods where 
presence or absence occurs at a demal level. 

Note on other species. — Atylus? batei Cun- 
ningham, 1871, from Magellan Strait, ap- 
parently is neither of the species studied 
herein as it has five dorsal body teeth, 
whereas the two discussed herein have four 
or fewer. 


Austroregia huxleyana (Bate) 
Figs. 2-4 


Atylus Huxleyanus Bate, 1862:135, pl. 25, 
fig. 4.—Cunningham, 1871:498. 

Acanthozone Huxleyana. —Della Valle, 
1893-612, pl. 595 fie. 23: 

Halirages Huxleyanus. —Stebbing, 1906: 
291; 1914:362.—Schellenberg, 1931:176, 
pl. 1, fig. k.—K. H. Barnard 932-155" 
fig. 93. 

Halirages stebbingi. — Alonso, 1980:10, fig. 
7 (not Schellenberg, 1931). 


Diagnosis. —Calceoli abundantly present; 
posterior pereonites not formed into pos- 
terolateral sharp wings in adults; outer rami 
or uropods 1-2 with several marginal spines 
in two rows; dorsal tooth of pleonite 3 usu- 
ally as large as tooth on pleonite 2. 

Material.—BMNH Holotype Hermit Is- 
land, Magellan Strait, Chile, Antarctic Ex- 
pedition, female “‘a’’ 19.8 mm, lacking an- 
tenna 2 (illustrated). BMNH 1928.12.1: 
2122-27, Syntype Falklands, R. Vallentin 
Expedition, formerly identified as regis, one 
specimen. BMNH 1936.11.2: 1381-84, st. 
53, 12.5.1926, Discovery 0-2 m, female “b”’ 
14.80 mm, identified as huxleyanus by K. 
H. Barnard. BMNH 1936.11.2: 1381-84 
(Part), st. 56, 16.5.1926, Discovery BtS 
10.5-16 m, identified as huxleyanus by K. 
H. Barnard, five specimens partly frag- 
mented. 


VOLUME 102, NUMBER 3 705 


Fig. 2. Austroregia huxleyana: holotype, female “‘a’’ 19.8 mm, a, Body; b, Apex of right mandible; c, Antenna 
1 lateral; d, Pleon; e, Apex of left mandible; f Head; g, Right lacinia mobilis; h, Prebuccal, anterior view, dorsal 
to right. Female “b” 14.8 mm: i, Head; j, Antenna 2, medial; k, Pleonite 7 (left) to pleonite 4 (right). 


706 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Hd) 
! N By 


' 


' mn 


Fig. 3. Austroregia huxleyana: holotype, female “a” 19.8 mm, a, Region of accessory flagellum; 5, Gill 6; c, 
Coxa 1; d, Coxa 2 and oostegite; e, Coxa 3 and gill: f Coxa 4 and oostegite; g, Coxa 5 and oostegite (small) and 
gill (large); 4, Lower lip; i, Right mandible; 7, Outer plate of maxilla 1; k, Maxilla 2; /, Palp of right mandible; 
m, Maxilla 1; n, Maxilliped. 


VOLUME 102, NUMBER 3 


Ger 


KAA 


107 


Fig. 4. Austroregia huxleyana: holotype, female ‘“‘a” 19.8 mm, a, Gnathopod 1, lateral; b, Gnathopod 2, 
medial; c, Pereopod 7; d, e, Telson; f—h, Uropods 1, 2, 3; i, Pereopod 4. 


Description of holotype female “a’’ 19.8 
mm.—Antenna 2 missing, nee 5-7 
broken, not available from better specimen, 
no enlarged views of pereopods 4—6 possi- 
ble. Rostrum small, blunt; eyes small, cir- 
cular, formed of ommatidia surrounding 
dark core of pigment, lateral cephalic lobes 
not protruding, truncate, antennal sinus 
weak, concave, bounded by tooth below. 
Antenna | about 50 percent as long as body, 
peduncle short, articles successively shorter, 
articles 2 and 3 with weak apicoventral 


tooth, each with 8+ ventral calceoli, acces- 
sory flagellum fused to peduncle, formed of 
small squared boss bearing one long and 3 
short setae, primary flagellum with 44 ar- 
ticles, some basal articles of bead form, first 
slightly enlarged and subrectangular, others 
following of varying sizes in sets of 2 and 
3, narrower to broader, calceoli present, ter- 
minal member of each set with groups of 
aesthetascs, broad articles with aesthetascs 
= 2.40.0, 12,1 5,18.22.2529.55,00, aes- 
thetascs on these articles posterior, numbers 


708 


eS ee he eee 
for the most part calceoli occurring in com- 
plex positions similar to complex setal dis- 
tributions of following species, regis, for ex- 
ample (‘‘a’’ = anterior, = inner, O° 


66599 
1 


0” = 
outer, ““‘p’’ = posterior, from lateral views), 
article 1 with 0, article 2 = lp, article 3 = 
2a,1-0,1p, article 4 = Ip, article 5 = Ip, 
articles 6,9,13 = 2a,2-0,31,1p, articles 7,8, 
PO212.,14,15:17,18'20,2272495 27229 333: 
and 35—44 = Ip, articles 11,16,19,21,23,26, 
28,30,34 = 1la,1-2-0,2-31,1-2p, article 32 = 
2a,2-0,11,2p. Antenna 2 missing (see next 
specimen). 

Body weakly depressed, pereonites shal- 
low, lacking lateral ridges above coxae, not 
produced strongly posterolaterally, but 
weakly on 7, from dorsal view these seg- 
ments not forming lateral wings, pereonite 
7 with posterodorsal tooth, pleonites 1—2 
with larger tooth, pleonite 3 with equally 
long thicker tooth, urosomites 1—3 rounded 
above. Coxae shorter than depth of pereo- 
nites, coxae 1-4 softly quadrate, with 
rounded distal corners, almost subcircular, 
coxae 3-4 weakly excavate behind, poste- 
rior lobe of coxa 5 as long as coxa 4, coxa 
6 not shorter than 5, coxa 7 shortest, un- 
lobed; coxae poorly armed. 

Epistome and labrum rounded truncate 
anteriorly, see illustration for anterior view. 
Right and left incisors multitoothed, right 
lacinia mobilis with about 3 teeth, left with 
6 teeth, about 8 right and 9 left rakers, each 
with extra interrakers, molar strongly tritu- 
rative, palp article 2 densely setose medi- 
ally, article 3 weakly bent, with no outer 
basal setae (=A-setae), inner margin with 
several C-setae and many D-setae, apex with 
8+ E-setae. Lower lip lacking inner lobes. 
Inner plate of maxilla 1 subrectangular, with 
6 apicomedial setae, outer plate with 11 
mostly denticulate spines, palp article 2 
symmetrical on both sides, with 7 thick api- 
cal spines, 2 thinner apicolateral spines, no 
facial subdistal setae. Inner plate of maxilla 
2 as long as and scarcely narrower than outer 
plate, lacking facial row of setae, medial 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


margin with three or four setae stouter and 
longer than apical cluster. Inner plate of 
maxilliped with 3 stout apical spines, one 
ventromedial coupling spine (not shown in 
illustration), outer plate broad and squat, 
with six apical setae, medially with thin ta- 
pering setae occasionally in pairs, palp ar- 
ticles 1-2 broad, 1—2 with few apicolateral 
and 2 with many medial setae, article 3 
weakly geniculate, moderately setose me- 
dially, dorsolateral face with groups of setae, 
dactyl stubby, with short apical nail and 
many accessory inner setules. 

Gnathopods small, first scarcely larger 
than second, wrists slender, barely lobate, 
hands much longer than wrists, ovatorect- 
angular, palm of gnathopod 1 very oblique, 
defined by group of two facial spines, with 
two secondary groups outside dactylar apex 
on posterior margin of hand, dactyl simple, 
curved, lined with bent setules; palm of 
gnathopod 2 slightly better defined, rela- 
tively shorter than on gnathopod 1. Pereo- 
pods 3-4 as illustrated, locking spines 3 di- 
verse members, posterior margins of article 
6 with five armament sets each with two or 
three spines and spinules, dactyls simple, 
each with inner marginal and outer facial 
setule. Article 2 of pereopods 5-7 diverse, 
weakly ovate to more sharply trapezoidal 
successively, subrectangular distally, weak 
to strong posteroventral lobe present, limb 
lengths increasing successively. Gills pres- 
ent on coxae 2-6; oostegites poorly devel- 
oped but of slightly expanded form on coxae 
2-5, slightly pointed on coxa 2, paddle 
shaped on coxae 3-4, small and paddle 
shaped on coxa 5, setae absent but marked 
by humps or sockets. 

Epimera 1-3 alike, each with convex pos- 
terior margin, sharp and weakly turned pos- 
teroventral tooth connected to lateral ridge, 
otherwise naked. Outer rami of uropods 1— 
2 shortened, all rami bearing apical spines 
and two marginal rows of spines, both upper 
margins of peduncle on uropods 1-2 with 
row of spines, those on uropod 1 lateral 
margin confined to distal half. Peduncle of 


VOLUME 102, NUMBER 3 


uropod 3 slightly elongate, with five medial 
spines, rami extending equally, slightly 
curved apically, sharp, each with two rows 
of marginal spines, these spines doubled on 
outer margin of outer ramus, inner margin 
of inner ramus also strongly setose, other 
margins with sparse setae mostly apical. 
Telson elongate, basally broadened, weakly 
tapering, cleft about 20 percent of its length, 
each side of apex with four setules in tan- 
dem. 

Female “‘b’”’ 14.80 mm. —Enlargement of 
head illustrated; antenna 2 illustrated, ar- 
ticles 3—5S of peduncle furnished with ventral 
calceoli, articles 4-5 very short, flagellum 
thick, with 52 articles, last vestigial, aesthe- 
tascs absent, articles short and bead-like, 
calceoli present in relatively uniform pat- 
tern (“‘a” = anterior, ““m”’’ = medial, “‘p” = 
posterior), lp present on articles 1,3,4,6,8, 
11,13,16,19,23,25,29,31,34,36,39,41,43,45, 
48, none present on articles 50-51, articles 
9,10,12,14,15,17,18,20,24,26,30,32, with 
2a,3m,2p, articles 2,5 with 2a,4m,2p, article 
7 with 2a,3m,1p, articles 21,22,27,28,33,37 
with 2a,2m,2p, article 35 with 2a,2m,3p, 
articles 38,40,42,44,.46 with 2a,1m,3p, ar- 
ticle 47 with 2a,3p, article 49 with 2a,2p. 

Dorsal body cuspidation of 2-toothed 
form, only pleonites 1—2 each with medium 
sized dorsoposterior tooth; pereopods 5—7 
and uropods 1-3 badly broken. 

Distribution. — Boreal South America and 
Falkland Islands, 0-55 m. 


Austroregia regis (Stebbing) 
Figs. 5-7 


Halirages huxleyanus. —Stebbing, 1888: 
902, pl. 73 (not Bate, 1862). 

Bovallia regis Stebbing, 1914:362, pl. 8. 

Halirages regis. —K. H. Barnard, 1932:161, 
fig. 94. 

Halirages stebbingi Schellenberg, 1931:176, 
pl ie. 1. 


Diagnosis. —Calceoli usually absent; pos- 
terior pereonites formed into posterolateral 
sharp wings in adults; outer rami of uropods 


709 


1—2 with two or fewer marginal spines; dor- 
sal tooth of pleonite 3 usually smaller than 
tooth of pleonite 2. 

Material. —Syntypes, BMNH 1928.12.1: 
2122-27, Roy Cove, Falkland Islands, low 
spring tide, R. Vallentin Expedition: Lec- 
totype, here selected, male “‘c” 6.81 mm 
(illustrated); syntype male “‘d” 6.28 mm; 
syntype juvenile “e”’ 5.75 mm and one other 
juvenile. Following specimens all misiden- 
tified formerly as H. huxleyanus: BMNH 
1928.12.11: 2019-26 (part), Stanley Har- 
bour, Falkland Islands, coll. Stebbing, three 
giant specimens, male “f’’? 19.80 mm (il- 
lustrated body), female “g’? 23.81 mm (il- 
lustrated head and coxae 1-4), female “‘h,” 
unmeasured. BMNH 1928.12.1: 2019-26 
(part), Rock pools, Falkland Islands, 
20.11.1910, R. Vallentin Expedition, coll. 
Stebbing, three specimens, one large, one 
medium, one small. BMNH 1928.12.1: 
2019-26 (part), Falkland Islands, 
20.vi.1910, coll. Stebbing, nine juveniles. 
BMNH 1936.11.2: 1381-84 (part), st. 55, 
16.v.1926. BtS 10-16 m, Discovery det. K. 
H. Barnard, one small form. BMNH 
1936.11.2: 1385, st. 56, 16.v.1926, Net: BtS, 
10.5-16 m, “white, very heavily mottled 
with deep crimson,” Discovery det. K. H. 
Barnard, female “‘i,”” unmeasured, with long 
head tooth. 

Description of lectotype male “‘c’’ 6.81 
mm.—Antenna 2 missing, pereopods 3-4, 
6—7 broken, pereopods 5-7 illustrated from 
better specimen, left mandible only de- 
scribed, no enlarged views of head, epi- 
stome, labium, pereopods 3-7, dactyls. 
Rostrum small, blunt; eyes circular, with 
dark core of pigment, lateral cephalic lobes 
not protruding, truncate, antennal sinus 
weak, straight, oblique. Antenna | almost 
55 percent as long as body, peduncle short, 
articles successively shorter, accessory fla- 
gellum fused to peduncle, formed of small 
squared boss bearing | long and 3 short se- 
tae, primary flagellum with 30 articles, some 
basal articles of bead form, first slightly en- 
larged and rectangular, others following of 


710 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


i/ 
Y ee 


Ee. ¥ 


\WA Zz 
ie 


ea 
LF} 


= 
ve 


oe: 
Etec sace 
I) 


if 
CES 


= 


va \ 
oe 


a 


Fig.5. Austroregia regis: lectotype male “‘c’’ 6.81 mm, a, Body; c, Left uropod 3; d, Right molar; e, Maxilliped; 
g, Lower lip; h, Maxilla 2. Male ‘“‘d” 6.28 mm, b, Head. Male ‘“‘c’”’ 6.81 mm, f, Right mandibular palp. 


VOLUME 102, NUMBER 3 711 


Fig. 6. Austroregia regis: lectotype male “‘c’’ 6.81 mm, a, Maxilla 1; e, Other palp of opposite maxilla 1; f 
Medial antenna 1; g, Dorsal body, head at bottom; 4, Coxa 4 with gill; 7, Telson. Female “g” 23.81 mm, 5, 
Head and pereonites 1-5. Male “‘f”’ 19.8 mm, c, Dorsal body from pereonite | (top) to pleonite 1 (bottom); d, 
Body. 


712 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 7. Austroregia regis: lectotype male “‘c” 6.81 mm, a, Region of accessory flagellum; b, Coxa 1; c, Coxa 
2 with gill; d, Coxa 3 with gill; e, Coxa 5 with gill; £ Coxa 6 with gill; g, Coxa 7; h, Gnathopod 1, medial; 7, 
Gnathopod 2, medial; k, /,) Uropods 2 and 1; m, Pleon. Male ‘“‘d” 6.28 mm, j, Pereopod 3. 


VOLUME 102, NUMBER 3 


varying sizes in sets of 2 and 3, narrower to 
broader, calceoli absent, terminal member 
of each set with groups of aesthetascs, broad 
articles with aesthetascs = 1,3,5,8,11,14,17, 
20,22,24,26,28, aesthetascs on these articles 
posterior, numbers = 4,2,2+,3+,4,3,3,1,2, 
2,1,1, setal distributions complex, for ex- 
ample (“‘a’”’ = anterior, “1” = inner, “m” = 
marginal, “‘s’” = submarginal, from lateral 
views), article 1 with 2a,2p, article 2 = 0a, Ip, 
article 3 = 2a,5p, article 4 = 3m,2p, article 
5 = 2a,4+p, articles 6,9,12,15 = 2-3s,31, 
articles 7,10,13,16 = 2—3m,3—4p, articles 
8,11,14,17 = 2-3a,3-5p, etc. Antenna 2 
missing (see next specimen). 

Body weakly depressed, pereonites shal- 
low, pereonites 3-7 with weak lateral ridge 
above coxae, pereonites 4—7 produced pos- 
terolaterally, weakly on 7, from dorsal view 
these segments forming lateral wings more 
rudimentarily expressed on pereonites 1-3, 
pereonite 7 with posterodorsal tooth, ple- 
onites 1—2 with larger tooth, pleonite 3 with 
short weakly acute dorsal projection, uro- 
somites 1-3 rounded above. Coxae shorter 
than depth of pereonites, coxae 1-4 softly 
quadrate, with rounded distal corners, cox- 
ae 3—4 weakly excavate behind, posterior 
lobe of coxa 5 as long as coxa 4, coxa 6 
scarcely shorter than 5, coxa 7 shortest, un- 
lobed; coxae poorly armed. 

Epistome and labrum as shown for A. 
huxleyanus. Incisors and laciniae mobiles 
of mandibles heavily encrusted, similar to 
A. huxleyanus, thus right incisor with about 
eight teeth, right lacinia mobilis with three 
teeth, about six rakers, molar strongly trit- 
urative, palp article 2 with most setae form- 
ing apicolateral row, article 3 weakly bent, 
with one (right) or two (left) outer basal setae 
(=A-setae), inner margin with several C-se- 
tae and many D-setae, apex with two E-se- 
tae. Lower lip lacking inner lobes. Inner 
plate of maxilla 1 subrectangular, with four 
apicomedial setae, outer plate with nine 
mostly denticulate spines, palp article 2 
asymmetrical on both sides, with five thick 
apical spines, one thinner apicolateral spine, 


713 


two facial subdistal setae on left side, right 
side with five thin apical spines and two 
subapical setae. Inner plate of maxilla 2 
slightly longer and narrower than outer plate, 
lacking facial row of setae, medial margin 
with three setae stouter and longer than api- 
cal cluster. Inner plate of maxilliped with 
three stout apical spines, one ventromedial 
coupling spine, outer plate broad and squat, 
with six apical setae, medially with thin 
blades occasionally in pairs, palp articles 1— 
2 broad, 2 with few apicolateral and many 
medial setae, article 3 weakly geniculate, 
poorly setose medially, dorsolateral face with 
groups of setae, dactyl stubby, with short 
apical nail and three accessory inner setules. 
Gnathopods small, first larger than sec- 
ond, wrists slender, barely lobate, hands 
much longer than wrists, subrectangular, 
palm of gnathopod 1 very oblique, defined 
by group of three or four marginal and two 
facial spines, with secondary group outside 
dactylar apex on posterior margin of hand, 
dactyl simple, curved, lined with bent set- 
ules; palm of gnathopod 2 better defined, 
relatively shorter than on gnathopod 1. Pe- 
reopods 3—4 as illustrated for specimen “‘d”’ 
below, locking spines 2 diverse members, 
posterior margins of article 6 with three or 
four armament sets each with one spine and 
from zero to two short setae, dactyls simple, 
each with inner marginal and outer facial 
setule. Article 2 of pereopods 5-7 narrow, 
subrectangular distally, weak posteroven- 
tral lobe present, limb lengths increasing 
successively. Gills present on coxae 2-6. 
Epimera 1-3 alike, each with convex pos- 
terior margin, sharp and strongly turned 
posteroventral tooth connected to lateral 
ridge, otherwise naked. Outer rami of uro- 
pods 1—2 shortened, each bearing only api- 
cal spines, inner rami with two rows of mar- 
ginal spines besides apical cluster, peduncle 
of uropod 1 with only one small apicolateral 
spine, medial margin with five spines, pe- 
duncle of uropod 2 with three dorsolateral 
spines and one basal setule, medial margin 
with five spines. Peduncle of uropod 3 


714 


slightly elongate, with two medial spines and 
one basal seta, rami extending equally, 
curved apically, sharp, each with two rows 
of marginal spines, inner also with seta in 
last two apical spine sets, outer also with 
seta in last spine set medially, each ramus 
with subapical setule. Telson elongate, ba- 
sally broadened, weakly tapering, cleft about 
10 percent of its length, each side with three 
setules in tandem. . 

Male ‘d’’ 6.28 mm.—Pereopod 3 en- 
larged to show details. 

Male ‘“f’ 19.80 mm.—Supposed termi- 
nal adult form, body illustrated: pereonites 
more alate posterolaterally, pereonites 2—7 
with shelf above coxae, pointed on pereo- 
nite 7; coxae more diverse, coxa 2 bluntly 
pointed below, coxae 3-4 relatively larger 
than in juvenile form; antenna 2 as illus- 
trated, articles 4-5 of equal length, flagellum 
“proliferate,” basal article with new articles 
forming inside; dorsal tooth of pereonite 3 
enlarged; apices of pereopods 6-7 illustrat- 
ed on body. 

Female “g’’ 23.81 mm. — Head with large 
anteroventral tooth below antennal sinus 
(illustrated). Coxae 3-4 sinuous postero- 
ventrally. Outer ramus of uropod 2 with two 
marginal spines. 

Female “h” large, unmeasured. —Outer 
ramus of uropod | with one marginal spine, 
of uropod 2 with one or two marginal spines 
(right and left); coxae acuminate as in fe- 
male “g”’ but head lacking tooth. 

Female “i”? unmeasured.—Like female 
“gs” but smaller, coxae more strongly acu- 
minate; each outer ramus of uropods 1-2 
with one marginal spine. 

Discussion.—The lectotype is unfortu- 
nately a small male less than one-third as 
long as the largest adults known (as mea- 
sured by parabolic method). It clearly has 
the flared-out posterolateral margins of the 
pereonites but not to the exaggerated degree 
of larger specimens. In large specimens the 
pereonal margins flare out laterally until they 
form horizontally projecting teeth. In some 
specimens the posterior margins of the flared 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


teeth are incised posteriorly and have very 
extended lateral wings which look like sep- 
arate processes from lateral view but which 
are simply more knife-like. Large adults oc- 
cur in two forms: (1) head with large pos- 
teroventral tooth coupled with strongly acu- 
minate anterior coxae and (2) head without 
strong posteroventral tooth coupled with 
poorly acuminate anterior coxae. Small 
specimens lack dorsally marginal spines on 
the outer rami of uropods 1-2 whereas larg- 
er adults have as many as three and two 
spines on the outer rami of uropods 1 and 
2 respectively. No calceoli have been dis- 
covered in our specimens of this species, 
although Stebbing shows calceoli in his orig- 
inal description. 

Relationship. — Differing from A. huxley- 
anus in the presence of flared margins on 
the pereonites, the poorly spinose outer rami 
of uropods 1-2 and the usual absence of 
calceoli. The specimen figured by Stebbing 
(1888: pl. 73) has calceoli. The condition of 
coxae in adult huxleyana resembles that of 
juvenile regis so that coxal forms are not 
necessarily definitive in comparing the two 
species. 

There remains the question of whether A. 
regis 1s congeneric with A. huxleyana be- 
cause the two species differ in the setal for- 
mulas on article 3 of the mandibular palp, 
in the spine numbers on the outer plate of 
maxilla 1, the difference in setal presence 
on the palps of maxilla 1, and the symmet- 
ricity of those palps. Article 3 of the man- 
dibular palp lacks A-setae in huxleyanus; 
the latter has 11 spines on the outer plate 
of maxilla 1 whereas A. regis has 9; and the 
palps are symmetrical and lack subdistal se- 
tae in A. huxleyana. There also remains the 
problem of speciation in this group and be- 
cause of the extensive demal variations al- 
ready noted, very large and widespread col- 
lections of antarctic materials should be 
assembled to study the life history and vari- 
ations in the group. 

Distribution. — Tierra del Fuego and Falk- 
land Islands, 0-55 m. 


VOLUME 102, NUMBER 3 


Acknowledgments 


Dr. Roger J. Lincoln of the British Mu- 
seum of Natural History made extensive 
contributions to this work and kindly sup- 
plied the SEM photographs. I thank Linda 
B. Lutz of Vicksburg, Mississippi, for ink- 
ing our illustrations and Patricia B. Crowe 
and Kimberly R. Cleary of the Smithsonian 
Institution for their assistance in the labo- 
ratory. 


Literature Cited 


Alonso, G. 1980. Amfipodos de la Ria Deseada (San- 
ta Cruz-Argentina).—Centro de Investigacion 
de Biologia Marina (CIBIMA) del sistema de 
Centros del Instituto Nacional de Tecnologia 
Industrial (INTI), Contribucion Cientifica 175: 
3-15. 

Barnard, J. L.,& G.S. Karaman. 1987. Revisions in 
classification of gammaridean Amphipoda 
(Crustacea), Part 3.— Proceedings of the Biolog- 
ical Society of Washington 100:856-875. 

Barnard, K. H. 1932. Amphipoda.— Discovery Re- 
ports 5:326 pp. 

Bate, C. S. 1862. Catalogue of the specimens of am- 
phipodous Crustacea in the collection of the 
British Museum. London, British Museum [of 
Natural History], 399 pp. 

Bousfield, E. L. 1977. A new look at the systematics 

of gammaroidean amphipods of the world.— 

Crustaceana, Supplement 4:282-316. 

. 1983. An updated phyletic classification and 

paleohistory of the Amphipoda. Pp. 257-277 in 


AIS 


F. R. Schram, ed., Crustacean phylogeny, A. A. 
Balkema, Rotterdam, 365 pp. 

Cunningham, R.O. 1871. Notes on the reptiles, Am- 
phibia, fishes, Mollusca, and Crustacea obtained 
during the voyage of H.M.S. ‘Nassau’ in the 
years 1866-69.—Transactions of the Linnean 
Society of London 27:465-502. 

Della Valle, A. 1893. Gammarini del Golfo di Na- 
poli.— Fauna und Flora des Golfes von Neapel 
und der angrenzenden Meeres-Abschnitte, 
Monographie 20:xi and 948 pp. 

Lincoln, R. J., & D. E. Hurley. 1981. The calceolus, 
a sensory structure of gammaridean amphipods 
(Amphipoda: Gammaridea).— Bulletin of the 
British Museum of Natural History (Zoology) 
40:103-116. 

Schellenberg, A. 1931. Gammariden und Caprelliden 
des Magellangebietes, Sudgeorgiens und der 
Westantarktis.—Further Zoological Results of 
the Swedish Antarctic Expedition 1901-1903 
2(6):290 pp. 

Stebbing, T. R. R. 1888. Report on the Amphipoda 
collected by H.M.S. Challenger during the years 
1873-76.—Report on the Scientific Results of 
the Voyage of H.M.S. Challenger During the 
Years 1873-1876, Zoology 29:xxiv and 1737 


Pp 


1906. Amphipoda I: Gammaridea.— Das 
Tierreich 21:806 pp., Berlin. 

1914. 1. South African Crustacea (Part VII. 
of S.A. Crustacea, for the marine investigations 
in South Africa).— Annals of the South African 
Museum 15:1-55. 


Department of Invertebrate Zoology, 
NHB-163, Smithsonian Institution, Wash- 
ington, D.C. 20560. 


PROC. BIOL. SOC. WASH. 
102(3), 1989, pp. 716-725 


TWO NEW SPECIES OF WOOD-BORING LIMNORIA 
(CRUSTACEA: ISOPODA) FROM NEW ZEALAND, 
L. HICKSI AND L. RENICULUS 


Marilyn Schotte 


Abstract.—Limnoria hicksi is described and distinguished from L. indica 
Becker & Kampf, 1958 by the structures on the fifth pleonite and the pleotelson. 
Limnoria reniculus, similar to Limnoria foveolata Menzies, 1957, L. saseboensis 
Menzies, 1957, L. sexcarinata Kiihne, 1975 and Limnoria sublittorale Menzies, 
1957, differs from all four in the structure of the fifth pleonite, pleotelson and 
morphology of the lacinioid seta of the right mandible. 


At present only two reliably identified 
wood-boring isopods of the genus Limnoria 
have been reported from New Zealand. 
These are L. guadripunctata Holthuis from 
Portobello (Hurley 1961), Auckland Har- 
bour (McQuire 1964), and Port Nicholson 
(Ralph & Hurley 1952), and L. tripunctata 
Menzies, also from Auckland Harbour 
(McQuire 1964). Menzies (1959) listed L. 
quadripunctata site records for isopods he 
examined from Auckland, Wellington, and 
Napier, previously thought to be specimens 
of L. lignorum (Rathke). Chilton (1914) had 
assigned the latter name to wood-borers 
found in timbers in Auckland and Lyttleton 
harbors and in piles from Akaroa Harbour. 
The latter isopod, whose name was assigned 
to most limnoriids before Menzies’ 1957 
monograph, has an Arctic-boreal distribu- 
tion and has been reliably reported only in 
the Northern Hemisphere from 39° to 58°N 
and from Iceland (Menzies 1957). Two alga- 
borers have been found in New Zealand, 
Phycolimnoria stephenseni Menzies, in 
floating Lessonia near Auckland Island (also 
Macquarie Island), and P. segnis (Chilton) 
from seaweed in Lyttleton and Akaroa har- 
bors (Menzies 1957) as well as from Por- 
tobello (Hurley 1961). 

Across the Tasman Sea, Limnoria quad- 
ripunctata has been found recently in Aus- 
tralia at Goat Island and Sydney Harbour 


in association with L. indica and L. tri- 
punctata Menzies, as well as in Tasmania 
(Cookson 1987). Hale (1929) reported L. 
lignorum from Port Lincoln but this iden- 
tification is undoubtedly erroneous. L. sub- 
littorale Menzies has been collected from 
New South Wales (Menzies 1957), as has 
Phycolimnoria rugosissima Menzies. An 
additional alga-borer, P. nonsegnis Menzies 
from Tasmania, brings to eight the total 
number of limnoriids thus far known from 
the Australia-New Zealand region. 

Material of the new species has been de- 
posited at the National Museum of New 
Zealand in Wellington and the National 
Museum of Natural History, Smithsonian 
Institution, Washington, D.C. 


Family Limnoriidae 
Limnoria (Limnoria) hicksi, new species 
Figs. 1A—D, 2A—F, 3A-E 


Material. —Holotype, NMNZ Cr. 5702, 2 
tl 4.0 mm, Paratype, USNM 205960, 2 tl 
3.3 mm (on slide and SEM stub), stn K6 
R/V Kalinovo, 177°39'05’E, 37°23'07’S to 
177°36'06”E, 37°23'07’S, off New Zealand, 
1075-1100 m, in rotting wood, 23 Nov 
1981, coll. G. R. F. Hicks. 

Description.—Female: Pigment absent. 
Pleonite 5 with a row of four proximal tu- 
bercles and two somewhat larger distal tu- 


717 


VOLUME 102, NUMBER 3 


uorneulds 


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718 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


W 
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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 


VOLUME 102, NUMBER 3 


Fi Se 


B 


Fig. 3. Limnoria hicksi: A, Outer lobe of first maxilla; B, Second maxilla; C, Maxilliped; D, Left mandible; 


E, Lacinioid seta of right mandible. 


and third articles subequal in length; fla- 
gellum consisting of one very short basal 
article and two subequal articles. Antennal 
flagellum of four articles, first article longer 
than three distal articles combined. 
Mandibular palp of three articles, second 
article longest, terminal article bearing five 
distal fringed spines; “‘rasp”’ of left incisor 
strongly sclerotized; spine row of right man- 
dible of nine laciniate spines, distal edge of 
lacinioid seta broad, very finely toothed and 
acute at ends. Maxilla 1 with nine stout se- 
tae, four of which provided with blunt teeth 
and one with setules. Maxilla 2 as figured. 


Maxillipedal endite with single coupling 
hook and six setose spines on distal margin; 
articles 2 and 3 of palp subequal in length; 
epipod about three times longer than great- 
est width, not reaching distal margin of basi- 
pod. 

Pereopod | with bidentate accessory spine 
at base of dactylar unguis; propodus with 
one dentate and one fringed posterodistal 
spine. Pereopod 7, accessory spine of dactyl 
small and apparently not bidentate; carpus 
with many fringed spines on distal margin; 
merus with anterodistal margin somewhat 
produced and bearing numerous fringed 


720 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 4. Limnoria reniculus: A, Pleotelson and fifth pleonite; B, Pleotelsonic surface enlarged; C, Interior 
surface of kidney-shaped carina enlarged; D, Pleotelsonic margin. 


spines visible in lateral view. Endopods of for its collector, Dr. Geoffrey R. F. Hicks, 


pleopods 1 and 2 of female distally truncate. 
Structure of male second pleopod unknown. 
Uropodal exopod less than one-half length 
of endopod; latter bearing one plumose seta 
and several terminal setae of varying lengths; 
peduncle with row of short, simple setae on 
outer margin and additional row of long 
plumose setae near margin. 

Remarks.—The male of the sexually di- 
morphic L. indica is the only other known 
limnoriid with six basal tubercles on the 
pleotelson arranged similarly to those in L. 
hicksi. Instead of additional tubercles on the 
fifth pleonite, L. indica in both sexes pos- 
sesses two subparallel longitudinal carinae 
bearing setae, thereby distinguishing it from 
L. hicksi. 

Etymology. —The new species is named 


National Museum of New Zealand, Wel- 
lington, New Zealand. 


Limnoria (Limnoria) reniculus, new species 
Figs. 4A—D, 5, 6A—G, 7A—-E 


Material. —Holotype, NMNZ Cr. 5703, 
ovig. 2, tl 5.0 mm, Allotype, NMNZ Cr. 
5704, 6, 4.4 mm, Paratypes, NMNZ Cr. 
5705, 11 ovig. 2, 16 2, 15 6, South Taramaki 
Bight, North Island, New Zealand, R/V 
James Cook, sta J20/25/84, 40°56.4'S, 
174°44.0’E to 40°59.8’S, 174°43.7’E, in rot- 
ting wood, 144-182 m, 29 Nov 1984, coll. 
G. R. F. Hicks. Paratypes, USNM 205961, 
11 ovig. 2, 15 2, 14 6, from same locality. 
Other material: USNM 205962, 1 speci- 
men, South Taranaki Bight, New Zealand, 


VOLUME 102, NUMBER 3 


m 


Limnoria reniculus: Different specimen from same lot as specimen in Fig. 4, pleotelson and fifth 


400M 


Fig. 5. 
pleonite. 


sta J20/16/84, 40°33.0’'S, 173°04.6’E to 
40°31.7’'S, 173°01.2’E, in rotting wood, 47— 
52 m, 28 Nov 1984, coll. G. R. F. Hicks. — 
USNM 205963, 5 specimens, sta J15/40/ 
84, 40°33.4'S, 173°04.6’E to 40°32.4’S, 
173°02.6’E, in rotting wood, 44-52 m, 28 
Nov 1984, coll. G. R. F. Hicks. —USNM 
205964, 11 specimens, sta J16/6/84, 
S751 7 7°19'E to 37°51'S, 177°15’E, in 
rotting wood, 48-49 m, 17 Sep 1984, coll. 
G. R. F. Hicks—USNM 205965, J15/22/ 
84, 41 specimens, 42°21.6’S, 170°49.5’E to 
42°24.4'S, 170°48.1’E, in rotting wood, 179-— 
184 m, 3 Sep 1984, coll. G. R. F. Hicks. — 
USNM 205966, sta J15/40/84, 5 speci- 
mens, 42°59.4’S, 170°16.5’'E to 42°57.2’'S, 
170°19.3’E, in rotting wood, 60-62 m, 5 Sep 
1984, coll. G. R. F. Hicks. —USNM 205967, 
6 specimens, sta J15/28/84, 42°45.8’S, 
170°28.2’E, in rotting wood, 50-64 m, 4 Sep 
1984, coll. G. R. F. Hicks.—USNM 205968, 
3 specimens, sta J20/12/84, 40°46.7'S, 
173°48.0’E to 40°47.9'S, 173°47.3’E, in rot- 
ting wood, 60-65 m, 27 Nov 1984, coll. G. 


R. F. Hicks. —USNM 205969, 6 specimens, 
off North Island, New Zealand, R/V James 
Cook, sta J9/70/84, 38°02'S, 174°37'E to 
38°06'S, 174°36’E, in rotting wood, 50-51 
m, 3 Jun 1984, coll. G. R. F. Hicks. —USNM 
205970, 3 specimens, sta J16/2/84, 37°39’'S, 
17 7°28'E to: 37°39'S, °177°24'E, in ‘rotting 
wood, 134-137 m, 17 Sep 1984, coll. G. R. 
F. Hicks.—USNM 205971, 8 specimens, 
New Zealand, off Cape Runaway, Wanaka, 
sta WK 3/19/85, 37°29.9'S, 177°47'E, in Log 
A, 450-481 m, 8 Dec 1985.—USNM 
205972, 19 specimens, New Zealand, Bay 
of Plenty, east of Alderman Island, NUNZ 
Cr. 5152, from timber, 400-520 m, Jun 
1969, coll. R. D. Cooper.—USNM 205973, 
14 specimens, New Zealand, WNW White 
Island, Tumokemoke Knoll, R/V Tangaroa 
(NZOI sta R 76), 37°29.1'S, 176°54.7'E, in 
twig, 248-283 m, 20 Jan 1979. 
Description. —Male: Dorsal integument 
of pereon and pleon rather foveolate, es- 
pecially pleonite 5. Pleonite 5 with some- 
what variable U-shaped carina (Figs. 4A and 


722 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


F 


Fig. 6. Limnoria reniculus: A, Antennule of male; B, Antenna; C, Lacinioid seta of right mandible; D, Left 
mandible; E, First maxilla; F, Second maxilla; G, Maxilliped. 


VOLUME 102, NUMBER 3 


5). Pleotelson densely spinulose with two 
rounded submedian ridges basally, flanked 
on either side by irregular, kidney-shaped 
Carinae, apparently bare interiorly but ac- 
tually covered with tiny spinules (Fig. 4C). 
Submedian ridges becoming obsolete dis- 
tally. Margins of pleotelson not tuberculate, 
having multiple rows of simple, unsheathed 
spines. Female dorsum as in male. 
Antennular peduncle of three articles; fla- 
gellum of male consisting of basal article, 
much shorter than wide, penultimate article 
with many aesthetascs ringing distal mar- 
gin, and terminal article half as wide as pre- 
ceding article. Antennal flagellum of 4 ar- 
ticles, basal article longer than 3 distal 
articles together. Mandibular palp of 3 ar- 
ticles, two proximal articles subequal in 
length; article 2 with 6 distal fringed spines; 
terminal article bearing 9 distal fringed 
spines; spine row of left mandible a single 
process; spine row of right mandible of 10 
laciniate spines, increasing in length proxi- 
mally; lacinioid seta with two lobes, one 
elongate and blunt, the other short and jag- 
ged; incisor with strongly sclerotized cusp. 
Maxilla 1 and 2 as figured. Maxillipedal en- 
dite with single coupling hook, eight spines 
on distal margin, five of which setulose; pal- 
pal article 3 longest and widest; epipod more 
than three times longer than greatest width, 
distally rounded and not reaching base of 
palp. Pereopod 1, bidentate accessory spine 
at base of dactylar unguis; propodus with 1 
dentate, plus one fringed posterodistal spine. 
Carpus and merus with short rows of round- 
ed scales on posterior surface. Pereopod 7, 
accessory spine of dactyl barely bidentate; 
carpus with many fringed spines on distal 
margin; merus produced anterodistally, 
bearing many fringed spines encircling dis- 
tal margin. Appendix masculina of pleopod 
2 of male articulating proximal to mid- 
length on median margin of endopod, ex- 
tending beyond ramus. Uropodal endopod 
elongate, about four times longer than wide 
and bearing five plumose setae and several 
lateral and terminal simple setae; uropodal 


7123 


exopod a slightly curved claw; peduncle with 
row of setae inserted medially to lateral 
margin. 

Remarks. —L. reniculus may be confused 
with L. saseboensis Menzies, which also has 
submedian longitudinal ridges on the pleo- 
telson, especially if debris obscures the kid- 
ney-shaped carinae in uncleaned specimens 
of the former. The U-shaped carina on 
pleonite 5 in reniculus may vary but the 
subparallel ridges do not join proximally as 
they appear to do in saseboensis. Menzies’ 
description notes the posterior edge of the 
pleotelson and lateral crests as tuberculate 
with “‘spike-like bristles on the margin.” 
SEM photos (Fig. 4) do not reveal obvious 
tubercles in the present species. The two 
species also differ somewhat in the shape of 
the lacinioid seta of the right mandible with 
“two medially curved teeth at apex” in L. 
saseboensis. L. sublittorale Menzies from 
Australia also has a pair of longitudinal ca- 
rinae on the telson but apparently lacks oth- 
er surface ornamentation there. The lacin- 
10id seta of the right mandible has two 
recurved teeth instead of a blunt lobe as in 
L. reniculus. L. foveolata Menzies, while also 
foveolate on the pleotelson and pleonite 5, 
has carinae on both segments which are more 
irregular than those in the new species, and 
it lacks the kidney-shaped features. The la- 
cinioid seta of L. foveolata as figured by 
Menzies has three teeth at the apex. 

The new species also resembles L. sex- 
carinata Kuihne. Examination of type ma- 
terial of the latter species reveals the carina 
of pleonite 5 as ““horseshoe-shaped” and the 
curving ridges on the pleotelson do not join 
distally. Kiihne’s (1975) description states 
that the lacinioid seta of the right mandible, 
apparently a good character for distinguish- 
ing members of this genus, is ““branched and 
furnished at the end with teeth.’’” SEM pho- 
tos of L. sexcarinata show the presence of 
tubercles and sheathed spines on the pleo- 
telsonic margin, neither of which exist in L. 
reniculus. 

Etymology.—The Latin reniculus (little 


724 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


sR 


POLL 
—— 


Fig. 7. Limnoria reniculus: A, First pereopod; B, Dactyl of first pereopod enlarged; C, Seventh pereopod; 
D, Uropod; E, Second pleopod of male. 


VOLUME 102, NUMBER 3 


kidney), proposed as a noun in apposition, 
refers to the shape of the outline of the ca- 
rinae flanking middorsal ridges on the pleo- 
telson. 


Acknowledgments 


Material described in this paper was kind- 
ly provided by Dr. Goeffrey R. F. Hicks of 
the National Museum of New Zealand. Mrs. 
Susann Braden of the NMNH assisted with 
preparation of the scanning electron micro- 
graphs. The manuscript benefitted from 
comments by Laurie Cookson and Brian 
Kensley, both of whom reviewed the manu- 
script. 


Literature Cited 


Chilton, C. 1914. The species of Limnoria, a genus 
of wood-boring Isopoda.—Annals and Maga- 
zine of Natural History, Ser. 8, vol. 13:380-390. 

Cookson, L. J. 1987. The occurrence of Limnoria 
indica Becker & Kampf (Isopoda) on the eastern 
coast of Australia.—Crustaceana 52(1):85-89. 

Hale, H. M. 1929. The crustaceans of South Austra- 
lia. 2:201-380, figs. 268-269. Government 
Printer, Adelaide. 


(P25) 


Hurley, D. E. 1961. A checklist and key to the Crus- 
tacea, Isopoda of New Zealand and the Sub- 
antarctic Islands.—Transactions of the Royal 
Society of New Zealand, Zoology 1(20):259-292. 

Kihne, H. 1975. Neubeschreibung einer holzzerst6- 
renden Bohrassel, Limnoria sexcarinata (Crus- 
tacea, Isopoda).—Zeitschnft fur angewandte 
Zoologie 62:447-455. 

McQuire, A. J. 1964. A note on the occurrence of 
marine borers in New Zealand. — Proceedings of 
the New Zealand Wood Preservers’ Association 
4:35-44. 

Menzies, R. J. 1957. The marine borer family Lim- 

noriidae (Crustacea: Isopoda).— Bulletin of 

Marine Science of the Gulf and Caribbean 7(2): 

101-200. 

1959. The identification and distribution of 
the species of Limnoria. Pp. 10-33 in Dixie Lee 
Ray, ed., Marine boring and fouling organisms. 
University of Washington Press, Seattle. 
Ralph, P. M., & D. E. Hurley. 1952. The settling and 

growth of wharf-pile fauna in Port Nicholson, 
Wellington, New Zealand.— Zoological Publi- 
cations, Victoria University College (19):1-22. 


Department of Invertebrate Zoology, Na- 
tional Museum of Natural History, Smith- 
sonian Institution, Washington, D.C. 20560. 


PROC. BIOL. SOC. WASH. 
102(3), 1989, pp. 726-731 


SOME ASPECTS OF THE BIOLOGY OF 
RHOPALOPHTHALMUS TATTERSALLAE PILLAI, 1961 
(CRUSTACEA, MYSIDACEA) AND 
EXTENSION OF RANGE INTO THE 
KHOR AL SABIYA, KUWAIT (ARABIAN GULF) 


Stephen A. Grabe 


Abstract. —Rhopalophthalmus tattersallae Pillai was collected from the Khor 
al Sabiya, Kuwait, extending its known range from the Indian Ocean to the 
northern Arabian Gulf. Population density was greatest in early fall when 
juveniles predominated and was lowest in mid-winter. Spawning activity was 
greatest in April, May, July, and August, and lowest in mid-winter. Brood size 
was correlated with body size; largest females and broods occurred January 
through May and smallest females and broods occurred July through Novem- 
ber. Sex ratio was close to 1:1 on most dates. A small number of R. tattersallae 
were parasitized by an unidentified dajid isopod. 


Rhopalopthalmus tattersallae was de- 
scribed by Pillai (1961; cited in Pillai 1965) 
from specimens collected in coastal waters 
off Kerala State, India. To my knowledge, 
there have been no subsequent records of 
its occurrence. Fifteen species of Rhopa- 
lophthalmus have been described (Mauch- 
line 1980) and all are inhabitants of neritic 
or estuarine waters (Tattersall 1957; Hodge 
1963; Pillai 1965, 1973; Mauchline & Mu- 
rano 1977; Wooldridge & Erasmus 1980). 
This paper presents some basic life history 
information for R. tattersallae and records 
its range extension into the northern Ara- 
bian Gulf. 

Methods. — Daytime zooplankton collec- 
tions were taken at stations in Kuwait Bay 
and the Khor al Sabiya (Fig. 1) from Sep- 
tember 1981 through September 1982. Du- 
plicate step-oblique tows were made with a 
0.5 m diameter plankton net outfitted with 
0.202 mm and 0.505 mm mesh. The sam- 
pling schedule is summarized in Table 1. 
Sample volumes were measured with a dig- 
ital flow meter. Additional samples were 
collected at Station 614 on 27 May and 21 
September 1982 specifically for mysids. A 


0.505 mm mesh net was towed near-bottom 
(horizontally); sample volumes were not re- 
corded. 

All mysids were sorted from each sample, 
identified to life stage, and carapace length 
(CL) measured (tip of the rostrum to the 
posterior border of the carapace). 

Brood sizes (number of larvae) were de- 
termined only for those females whose mar- 
supla appeared undisturbed. The presence 
of an unidentified ectoparasitic isopod (Da- 
jidae) was recorded. 

Monthly population density in the Khor 
al Sabiya (Stations 614 & 615) was based 
upon the September and October 1981 0.202 
mm collections and the November through 
August 0.505 mm mesh collections. To fa- 
cilitate interpretation of seasonal changes in 
population structure and breeding, samples 
were pooled for each date at the Khor al 
Sabiya stations. Assumptions were made, 
then, that the populations at the two sta- 
tions in the Khor were structurally similar 
and that the 0.202 and 0.505 mm mesh nets 
were similarly efficient in sampling all life 
stages of this species. 

In studies of mysid populations in which 


VOLUME 102, NUMBER 3 


Fig. 1. 


E29 


failaka 
island 


Location of the sampling stations in Kuwait Bay and Stations 614 & 615 in the Khor al Sabiya used 


to survey the Rhopalophthalmus tattersallae population during 1981-1982. 


longitudinal differences in population struc- 
ture have been observed, there have either 
been marked differences in habitat (e.g., 
depth) within the water body (Mauchline 
1970) or there was evidence for recruitment, 
transport and maturation of a coastally 
spawned population (Hulburt 1957). Since 
the Khor al Sabiya stations were of similar 
depth, temperature, salinity and dissolved 
oxygen (Dames & Moore 1983) and since 
there was no evidence of recruitment from 
Kuwait Bay (see below), the first assump- 
tion seems tenable. 

The second assumption, that of compa- 
rable sampling abilities of the two gears, is 
more questionable. Sampling rates of the 
two gears were generally similar in the Khor 
al Sabiya (12.6 vs. 12.7 m?/minute for the 
0.202 and 0.505 mesh, respectively; Dames 
& Moore 1983). A limited comparison of 
sampling efficiencies for Penaeidae mysis 


and postlarval stages found that abundance 
estimates of the two gears were comparable: 
0.202 mm abundance = — 1.38 + 1.1 (0.505 
mm abundance); r,, = 0.986; P < 0.01 
(Dames & Moore, unpublished data). 

ANOVA (Sokal and Rohlf 1981) was used 
to test for differences in mean CL of mature 
females by sampling date, and regression 
analysis was used to evaluate the relation- 
ship between brood size and CL. 

Results and discussion. —The Khor al Sa- 
biya is a river-like channel separating the 
Sabiya peninsula on the northeastern shore 
of Kuwait Bay from Kuwait’s Bubiyan Is- 
land (Fig. 1). Maximum depth is about 18 
m but study areas averaged <10 m. Bottom 
sediments at Station 614 were generally 
rock-mud and shell debris and at Station 
615, muddy sand and shell debris. Extremes 
in water temperature occurred during Feb- 
ruary (12.1°C surface, 11.8°C bottom) and 


728 


Table 1.—Sampling schedule for Rhopalophthalmus 
tattersallae in the Khor al Sabiya, Kuwait, September 
1981 to September 1982. 


Mesh size Mesh size 


0.202 0.505 0.202 0.505 
Date mm mm Date mm mm 
17 Sep 81 X 26 Apr Xa 
27 Sep xX 12 May xX X 
14 Oct x 18 May DS 
28 Oct x 27 May <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 
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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. 


VOLUME 102, NUMBER 3 


Fig. 1. 


39 


D 


Sathodrilus nigrofluvius: A, Lateral view of holotype; B, Lateral view of reproductive systems of 


holotype; C, Optical section of bursa; D, Lateral view of jaws. Abbreviations: af, atrial fold; ba, bursal atrium; 
ed, ectal duct of spermatheca; ejd, ejaculatory duct; ep, ental process of spermatheca; p, penis; ps, penial sheath 
of bursa; sb, bulb of spermatheca; spg, spermaducal gland. 


The spermiducal gland lacks deferent 
lobes, is about two-thirds the diameter of 
segment VI in length and three-eighths its 
own length in diameter and is slightly ta- 
pered at each end. It lacks even the rudi- 
ment of a prostate, and often lies longitu- 
dinally above the gut. 

The ejaculatory duct is a short, thin- 
walled, obscure tube that is more nearly in- 
ferred than seen in the available specimens. 
In one paratype it appears to be greatly ex- 
panded and the penial sheath collapsed at 
the place of its entry into the latter. Since 
serial sections are unavailable, it is possible, 


but unlikely, that the spermiducal gland 
opens directly into the penial sheath region 
of the bursa. 

The bursa is a cylindrical sac, subequal 
to the body diameter in length. About one- 
third of its length consists of the atrium and 
atrial fold with a greatly reduced lumen. The 
penis is a straight, cuticular tube attached 
by relatively thick strands to the inner wall 
of the penial sheath which is set off exter- 
nally from the bursal atrium by an encir- 
cling constriction (Fig. 1C). 

The spermatheca is characterized by a 
thick, muscular, irregularly bent ectal duct 


740 


that constitutes at least one-half the total 
length of the organ, a relatively short sper- 
mathecal bulb that is no greater in diameter 
than the ectal duct and a short, narrowed 
ental process (Fig. 1B). 

Variations. —In one specimen the antero- 
dorsal quadrat of the bursa is collapsed and 
the ejaculatory duct may be greatly expand- 
ed. Otherwise, no differences were noted. 

Affinities. —Among the species presently 
assigned to the genus Sathodrilus, S. vera- 
cruzicus Holt, 1968; S. hortoni Holt, 1973; 
S. okaloosae Holt, 1973; S. shastae Holt, 
1981; and S. nigrofluvius lack a prostate or 
rudiment thereof (1.e. how many do have a 
prostate; how big is this genus?). Of this 
coherent group, S. nigrofluvius most nearly 
resembles S. veracruZicus. 

Unlike S. nigrofluvius, S. veracruzicus 
lacks dorsal ridges, appears to have oral pa- 
pillae, and has a proportionately shorter and 
more slender spermiducal gland, a longer 
and more slender bursa, a penis that is 
looped (and hence longer than its sheath), 
and a spermatheca with an ectal duct that 
is less in diameter than the spermathecal 
bulb and lacks an ental process. 

Sathodrilus hortoni consists of larger 
worms with a parasitic mien: a thin body 
wall and a gut filled with “globules of fat.” 
The “‘oesophagus”’ is attached by strands of 
muscle to the body wall of segments I and 
II (Holt 1973:97—98). The upper lip is lobed. 
The secondary reproductive organs are un- 
usual: the male efferent apparatus is pro- 
portionately small, the spermiducal gland 
relatively long and slender; the spermatheca 
is composed of a large muscular spermathe- 
cal bursa and a slender spermatozoa storing 
“bulb”? with a thick muscular wall and no 
ental process. 

Sathodrilus okaloosae has low dorsal 
ridges on the body segments. The jaws are 
unusual: broad and thick with a “dental 
ridge”? bearing the teeth and a dental for- 
mula of doubtfully 3/4, possibly 1/4 (Holt 
1973:101). The bursa is less than the body 
diameter in length. The spermatheca has a 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


long, slender ectal duct and a long ental pro- 
cess, both proportionately less in diameter 
than those of S. nigrofluvius. 

Sathodrilus shastae is composed of larger 
animals up to 4 mm in length with two pha- 
ryngeal sulci and no dorsal ridges. The jaws 
are prominent; the dental formula 1/1. The 
ejaculatory duct is long and thick and the 
spermiducal gland is remarkably long (about 
twice the body diameter in length) and slen- 
der. The bursa, including penial sheath and 
penis, is proportionately small. The ectal 
duct of the spermatheca is short, the ental 
process narrow, the median bulb fusiform 
and flattened between the gut and body wall. 

Host. — Unknown. 

Distribution. —Members of the genus are 
distributed widely over the continent, but 
it may be worthy of note that the most struc- 
turally similar relative of S. nigrofluvius is 
from southern Mexico (the state of Vera- 
cruz) with its other close relatives from 
Florida (two) and the Pacific northwest, a 
distribution that suggests, as does the ab- 
sence of a prostate, that these species are 
‘“‘among the least dervied”’ within the genus. 

Material examined. —The types. 


Dr. Stuart R. Gelder has informed me 
that the name Adenodrilus which I proposed 
(Holt 1977) for a genus later (Holt 1986) 
assigned to the family Bdellodrilidae is a 
junior secondary homonym of Adenodrilus 
Chekanovskaya, 1959, based on a haplo- 
taxid oligochaete from central Asia. 

The genus-name Uglukodrilus is hereby 
proposed as a replacement name for Ade- 
nodrilus Holt, 1977. It is to be considered 
as masculine and is derived from that of the 
leader of a fictional band of Orcs (see Tol- 
kien, J. R. R. The Lord of the Rings, v. 2, 
1954) whose feeding habits were considered 
suspect. 


Acknowledgments 


Drs. Horton H. Hobbs, Jr. and Brent D. 
Opell have read a first draft of this paper. 


VOLUME 102, NUMBER 3 


As always, I am grateful. Also, I wish to 
thank Dr. Stuart R. Gelder for calling to my 
attention the homonymy of Adenodrilus 
Holt, 1977. Dr. Ernest R. Stout, Head, De- 
partment of Biology, Virginia Polytechnic 
Institute and State University, and his sec- 
retaries have been unsparing in their sup- 
port and help of which I am greatly appre- 
Clative. 


Literature Cited 


Chekanovskaya, O. V. 1959. On Oligochaeta from 
the bodies of water in Central Asia (Ferghana 
Valley and the River Nurgab).—Zoologicheskii 
Zhurnal 38:1151-1162, figs. 1-5. 

Holt, Perry C. 1967. Oedipodrilus oedipus, n. g., n. 

sp. (Annelida: Clitellata: Branchiobdellida).— 

Transactions of the American Microscopical 

Society 86(1):58-—60, figs. 1-4. 

. 1968. New genera and species of branchiob- 

dellid worms (Annelida: Clitellata).— Proceed- 

ings of the Biological Society of Washington 81: 

291-318, figs. 1-9. 


741 


1973. Epigean branchiobdellids (Annelida: 

Clitellata) from Florida. — Proceedings of the Bi- 
ological Society of Washington 86(7):79-104, 
figs. 1-8. 
. 1977. A gill-inhabiting new genus and species 
of the Branchiobdellida (Annelida: Clitellata). 
— Proceedings of the Biological Society of 
Washington 90(3):726-—734, figs. 1-5. 

1981. New species of Sathodrilus Holt, 1968 
from the Pacific drainage of the United States, 
with the synonymy of Sathodrilus virgiliae Holt, 
1977.— Proceedings of the Biological Society of 
Washington 94(3):848-862, figs. 1-3. 

1986. Newly established families of the order 
Branchiobdellida (Annelida: Clitellata) with a 
synopsis of the genera.— Proceedings of the Bi- 
ological Society of Washington 99(4):676-702, 
figs. 1-20. 


Department of Biology, Virginia Poly- 
technic Institute and State University, 
Blacksburg, Virginia 24061. Mailing ad- 
dress: 1308 Crestview Drive, Blacksburg, 
Virginia 24060. 


PROC. BIOL. SOC. WASH. 
102(3), 1989, pp. 742-752 


THE SECOND ANNUAL RISER LECTURE: 
ECLECTICISM AND THE STUDY OF 
POLYCHAETES 


Kristian Fauchald 


Abstract. —The study of polychaetes has involved two very different research 
programs: the morphological and systematic descriptions on one hand and the 
biological and physiological traditions on the other hand. The two traditions 
each represent two systems of two different approaches to the study of nature: 
on one hand structural versus procedural studies and on the other hand process- 
oriented versus taxon-oriented studies. None of the paradigmatic approaches 
common in biology (e.g., ecological, physiological, genetic or evolutionary) can 
solve their own problems by using only one of the four approaches. Under- 
standing the biology of the group can only come from a carefully managed 
eclectic approach to the study of the group. 


During the early part of my career the 
theory of science always appeared to exist 
totally independent of what I was doing as 
a biologist: Biology was something to be 
done, not thought about. A paper published 
about 25 years ago (Platt 1964) demonstrat- 
ed that I had been very wrong; the quality 
of a study depends crucially on the manner 
in which it is planned and performed. 

Since then I have examined my own and 
my fellow workers output for signs of an 
awareness of theoretical issues associated 
with the study of biology. I have concen- 
trated on the polychaete literature with 
which I am most familiar. By now more 
than 200 years worth of papers on poly- 
chaete morphology, systematics, phyloge- 
ny, physiology and ecology have accumu- 


The Riser Lecture Series.—In 1985 the annual Riser 
Lecture was initiated by members, alumni and friends 
of the Marine Science Center, Northeastern University 
at Nahant, Massachusetts. The occasion was the official 
retirement of Professor Nathan W. Riser. As teacher, 
biologist and founder of the facility, ““Pete’’ Riser en- 
dowed the laboratory with a legacy—the importance 
of considering the whole organism regardless of one’s 
special focus. We dedicate these annual lectures to that 
principle. 


lated, representing more than 10,000 
individual papers and books. 

In this paper I will review, very briefly, 
the development of the study of poly- 
chaetes. I will then attempt to put this over- 
view into a minimal theoretical context. The 
results are some rather trivial admonitions. 
I believe these recommendations to be 
worthwhile because most of my colleagues 
still behave as if their activities were theory- 
independent. If I can set them thinking about 
these issues, then the purpose of this paper 
will have been fulfilled. 


Early Studies of Polychaetes 


Aristoteles reported what might be inter- 
preted as scaleworms in the ocean; Pliny the 
Older gave a much more convincing de- 
scription of ““marine scolopenders”’ (Gillet 
1988) and this latter report was expanded 
on by both Rondelet and Gesner (Williams 
1851); these “‘scolopenders”’ have tradition- 
ally been identified as nereidid polychaetes. 
For all practical purposes polychaetes were 
first described in 1758 in the 10th edition 
of Linnaeus Systema Naturae. These early 
reports and the transition into a scientific 


VOLUME 102, NUMBER 3 


study of the polychaetes is described by Gil- 
let (1988). 

Names and descriptions.—The Linnean 
nomenclature separated names as labels 
from descriptions and definitions of the or- 
ganisms studied. For the first time logical 
procedures known since antiquity could be 
applied to the description of the living world. 
One could name an organism and define 
that label by descriptive terms, independent 
of the names themselves. The process has 
been taught as part of introductory classes 
in logic for a long time, nevertheless, the 
importance of this first application to bi- 
ology was overwhelming. The new nomen- 
clatural system made possible intelligible 
discourse about Nature in a way that no 
other device, before or after, has done. The 
practices of the scientists of the period re- 
flected an awareness of the different lan- 
guage levels involved in descriptive pro- 
cesses (Popper 1979). For example, I believe 
that the use of names of gods and goddesses 
for genera of various organisms reflects an 
awareness of the importance of the sepa- 
ration of names from definitions and de- 
scriptions. The trivial names, what we now 
call the species names, often were simple 
mnemonics: Nereis virens for example: the 
green nereid. Nereis diversicolor is another 
example of this naming tradition. 

The descriptions and definitions included 
morphological features. Microscopes were 
SO primitive that not much more than gross 
morphological features could be distin- 
guished. However, early illustrations may 
be remarkably accurate and detailed. Writ- 
ten descriptions uniformly are far less de- 
tailed. The early zoologists did exactly what 
we do: Include sufficient detail to distin- 
guish new taxa from previously known ones. 
One can hardly blame Linnaeus and his 
contemporaries for not appreciating how 
many different kinds of worms would even- 
tually be found, or for not developing the 
complete terminology for describing their 
wealth of morphological detail. The first 
major describers of polychaetes were Danes, 


743 


Otto Friedrich Muller (Muller 1776) and 
Otto Fabricius (Fabricius 1780), Russians, 
such as Peter Paul Pallas (Pallas 1766) and 
by the turn of the century the famous French 
scientists Cuvier, Lamarck, and Savigny. 

Reviews and classifications. —Lamarck 
and Cuvier, independently and in compe- 
tition, reviewed all polychaetes described, 
sorted out, and named a whole series of new 
higher taxa, especially genera and families 
(Lamarck 1816, Cuvier 1817). Another fa- 
mous French worker, Savigny, had made 
most of the new observations and descrip- 
tions. He was a careful observer with a fine 
eye for finding differences among similar 
forms (Savigny 1820). Lamarck added con- 
siderably to our understanding of the rela- 
tionships among the polychaetes. Also his 
separation of the polychaetes into two ma- 
jor groups, those with red blood and those 
with white blood, revealed an interest in 
physiological properties of the organisms. 
Nevertheless, more of Cuvier’s morpholo- 
gy-based system has been retained than of 
Lamarck’s. 

Detailed descriptions of newly discovered 
polychaetes became divorced from the time 
in which they were penned. The descrip- 
tions have increased in detail and length 
from one or two lines to several printed 
pages, but we still use most of the termi- 
nology and the overall pattern of descrip- 
tions established by Audouin and Milne Ed- 
wards in a study of the French fauna in the 
early 1830’s (summarized in Audouin & 
Milne Edwards 1834). 

The system used by Audouin and Milne 
Edwards closely resembled the Cuvierian 
system and formed the base for all workers 
over the next 20 years. By 1850 however, 
the emphasis of exploration shifted to Ger- 
many: Adolph-Eduard Grube (1850) issued 
a major review of the polychaete families 
and this paper was the standard for the next 
15 years. 

Two scientists working in Stockholm 
made the next major advances in the mid 
1860’s. Kinberg reported on his worldwide 


744 


travels and Malmgren detailed the North 
Atlantic and Arctic Ocean faunas. These two 
scientists represent two very different ap- 
proaches to descriptive science. Kinberg 
briefly described species collected on the 
cruise of the Eugenie around the globe and 
added numerous new taxa at all levels (Kin- 
berg 1865, 1910). Malmgren’s (1867) stud- 
ies were intensive; he focussed his attention 
on a much smaller area and carefully re- 
viewed all previous work before commit- 
ting himself to describing a new taxon. This 
difference in approach closely matches a pe- 
rennial difference among descriptive biol- 
ogists; among modern systematists Gesa 
Hartmann-Schroder and Olga Hartman 
both have used Kinberg’s approach, where- 
as Marian H. Pettibone more closely match- 
es Malmgren. I have done a bit of both. 

Kinberg and especially Malmgren did 
their best to increase the consistency in use 
of terms and in the amount of detail re- 
quired for adequate descriptions. Quatre- 
fages (1866) issued a large-scale review of 
the whole annelid fauna as he knew it. Per- 
haps more pedestrian a systematist than the 
others mentioned, he nevertheless became 
extremely influential, due in part I believe 
to his location: he was in Paris, and had a 
long history of publications on polychaetes 
by the time he issued his magnum opus. 
Kinberg had published a few earlier papers, 
but neither he nor Malmgren ever issued 
any additional major contributions to the 
study of polychaetes. They both left science 
shortly after the papers mentioned were 
published. 

Ludwig Schmarda is one of the more col- 
orful persons in the history of polychaete 
studies. He travelled around the world in 
the 1850’s, not in an exploring vessel, but 
by hitch-hiking on commercial sailing ves- 
sels. His description of his trip from South 
Africa to Australia is singularly harrowing, 
including very bad weather, seasickness, 
scurvy and assorted other diseases. In Chile 
he lost his collections to a fire on board; in 
Panama he was robbed by some rather un- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


savory characters who made their living by 
preying on people going from the U.S. east 
coast to the west coast via the Isthmus. De- 
spite the loss of his collections, he published 
a large report (Schmarda 1861) that appar- 
ently was largely overlooked by his contem- 
poraries. This was probably in part due to 
the increasing standards of descriptions and 
illustrations. Schmarda’s effort was, how- 
ever, the earliest worldwide tropical survey 
of polychaetes. He described a large number 
of new species for which there are few types 
available and poor locality information. At 
that time, there was no requirement that 
types should be deposited anywhere: De- 
scriptions were considered adequate evi- 
dence for the presence of a new taxon. How- 
ever, the first Nomenclature Code, and 
perhaps just as importantly, the first volume 
of Zoological Record, was issued in 1864. 

The morphological tradition. —The mor- 
phological tradition, outlined above, has 
continued through the work of McIntosh 
(1885), Fauvel (1923, 1927), and Augener 
(1918), and is now followed by most prac- 
ticing systematists. The total focus of this 
tradition is very limited in the kind of evi- 
dence deemed acceptable. Most system- 
atists will accept only features that can be 
seen either with the naked eye or with stereo 
or compound microscopes as valid taxo- 
nomic characters. Furthermore, a tradition 
among polychaete systematists suggests that 
all reasonably well preserved specimens, es- 
pecially anterior ends, should be identifiable 
to species. I have more than once heard 
complaints from well known systematists 
that a published description was too difficult 
to use, or was impractical, because it used 
information not readily available using 
minimal technical equipment, or required 
the presence of complete specimens. This 
tradition is clearly at odds with, for exam- 
ple, students of isopod crustaceans who for 
years have accepted limits on the identifi- 
ability of all specimens. 

The biological tradition.— Another tra- 
dition in the study of polychaetes dates back 


VOLUME 102, NUMBER 3 


to about 1850. Thomas Williams (1851) 
published a major review of the biology and 
physiology of the polychaetes. This sum- 
mary is now rarely quoted; it has been su- 
perceded by more recent reviews, but it was 
important historically because Williams re- 
viewed all information available about the 
life of all worms known to science. Some of 
the data quoted by Williams date back to 
Lamarck and are speculative rather than ob- 
servational in nature and some rather quaint 
notions were paraded only eight years be- 
fore the publication of Darwin’s Origin of 
Species. Williams made some original phys- 
iological observations on various English 
polychaetes. 

The most impressive of the early poly- 
chaete biologists was Eduard Claparéde, a 
rather tubercular-looking Swiss, who did 
most of his work in France and Italy (Cla- 
paréde 1854). By 1865 he had gotten into 
a rather virulent quarrel with Quatrefages 
over all of Quatrefages’ new taxa, defined 
in many cases without access to any mate- 
rial (Quatrefages 1865a, Claparéde 1865, 
Quatrefages 1865b). Claparéde emphasized 
the importance of observations on live or- 
ganisms; Quatrefages by that time had be- 
come very collections-oriented. This differ- 
ence in approach formed the background for 
the disagreement. Claparéde, true to his 
principle, deposited no specimens in any 
museum, making many of his new taxa dif- 
ficult to define accurately. 

The second tradition was biological in na- 
ture: studying live organisms and making 
observations of the live processes, such as 
reproduction, development and feeding. 
These kinds of observations were difficult 
to quantify in an age of poor mechanical 
recording devices, no photography to speak 
of, and certainly no electronic recording de- 
vices. Additionally, statistics had not yet 
developed to the point where repeated sam- 
ples were taken. The studies were therefore 
often episodic in nature, and observations 
were only rarely organized into tables or 
other means of presenting large, easily sur- 


745 


veyed data. The kinds of observations at- 
tempted by Claparéde are still difficult to 
document for theoretical reasons that I will 
touch on below. 

Claparéde combined his studies of live 
organisms with a detailed study of microan- 
atomical structures. These studies are ex- 
cellent and are still the best starting point 
for any anatomical studies in the groups he 
covered. Claparéde’s illustrations are among 
the best ever published on polychaetes. The 
most important aspect of Claparédes work 
was that he demonstrated that a remarkable 
amount of information could be gained by 
looking at live organisms. He also demon- 
strated that detailed anatomical and histo- 
logical studies yielded systematically dis- 
tributed information, which could be 
potentially useful in systematics. 

Ehlers tried to combine the two traditions 
in his massive publication “Die Borsten- 
wurmer’’ issued in two parts (Ehlers 1864— 
1868). Some of his descriptions of new taxa 
run 10-15 printed pages, accompanied by 
one or two full packed plates of illustrations. 
Consequently, Ehlers succeeded in going 
through less than '4 of the then known poly- 
chaete taxa in roughly 700 pages of text, but 
for the groups he covered, his volume is 
absolutely indispensible. Ehlers’ research 
later devolved to thoroughly traditional, 
morphological descriptions. I can find no 
evidence in any of his publications that he 
attempted to complete the massive study he 
had started. 

The study of live polychaetes eventually 
developed into a tradition of physiological 
studies, based usually on members of rela- 
tively few families with highly characteris- 
tic, often unusual physiological patterns. 
These studies are often performed by pro- 
cess-oriented rather than by comparative 
scientists. Reproductive studies, while cov- 
ering in part members of most groups, have 
been focussed on eunicids, nereidids and 
syllids (Schroeder & Hermans 1975); stud- 
ies of respiratory and blood physiology on 
glycerids, terebellids and scattered other 


746 


groups (Dales 1969, Florkin 1969). Studies 
of regeneration have focussed on sabellids 
with few glances in other directions (Need- 
ham 1969). Genetic studies have been done 
on dorvilleids and little else (Akesson 1982). 
Neurophysiologists have studied the prop- 
erties of the giant nerve fibers in sabellids 
of the genus Myxicola with very little con- 
cern for the biology of the organism at all. 
There are about 80 families of polychaetes 
and of these at least 60 are common in shal- 
low water and relatively readily available; 
nevertheless live studies have focussed on 
a few popular groups and usually on only 
one or a few species in each group at that. 

The results of the biological and physio- 
logical studies have been very valuable, but 
less as a comparative study of polychaetes 
than as an exploration of various biological 
and physiological mechanisms. 


Theory and the Study of Polychaetes 


The rather conservative descriptive tra- 
dition continues among polychaete system- 
atists; for each advance in morphological or 
anatomical technique, traditionalists hang 
back, not wanting to get involved with new 
methods or add new features to the descrip- 
tions. Often the young turks among poly- 
chaetologists are traditionalists in the study 
of other groups of organisms, especially ver- 
tebrates. Very few of the scientists closely 
associated with the study of polychaetes have 
demonstrated strong theoretical interests. 
For example, it is difficult to find any ref- 
erence to evolution, or to Darwinian or anti- 
Darwinian thinking anywhere. Ehlers’ pub- 
lication from 1864-1868 gave no indication 
of a major revolution in biological thinking 
taking place at the time. McIntosh (1885) 
mentioned nothing about phylogeny in his 
treatment of the Challenger polychaetes. 
One outstanding exception is E. Meyer, who 
in his studies of polychaetes indicated a 
good, often anticipatory understanding of 
biological theory (Meyer 1890). This paper 
is frequently quoted in the literature on phy- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


logeny of the invertebrates, but not often by 
polychaete taxonomists. 

Some of the developmental biologists as- 
sociated with the study of spiral cleavage at 
Woods Hole Marine Biological Laboratory 
used polychaetes for their studies. These sci- 
entists had deep theoretical interests and 
showed great skill in using the polychaete 
material in clarifying theoretical problems 
(Wilson 1898, Treadwell 1901). 

The reason for the lack of theoretical and 
one might say scholarly interest in the study 
of polychaetes is relatively easily found. 
Most scientists published only a single pa- 
per on polychaetes and very few made the 
study of these animals their lifetime occu- 
pation (Reish 1958). Through about 1950, 
the study of polychaetes was a relatively lei- 
surely pursuit. Even in most early benthic 
ecology studies (Petersen 1911, Blegvad 
1930), few polychaetes are mentioned or 
named, except to family. In morphological 
studies, the annelids were considered a step- 
ping stone to the arthropods (Hanstrom 
1928, Binard & Jenner 1928, and the dis- 
cussion of the anterior nervous system of 
the annelids and arthropods) and thus of 
interest insofar as they showed the step-wise 
advance to the conditions present in the ar- 
thropods. Parenthetically, papers that treat 
polychaetes well from a theoretical point of 
view were, with few exceptions, written by 
scientists with a limited experience in the 
group (Hanstrom 1928, Hatschek 1893). 
This generalization is far less true today than 
it was before WWII. 

The rapid development of interest in ben- 
thic ecology following the publication of 
Thorson’s (1957) review of the topic lead 
to considerable change in attitude. Poly- 
chaetes have turned out to be extremely 
common in the marine benthos; benthos 
ecologists have changed their attitudes to- 
wards the importance of polychaetes with 
the mesh-size of their screens. Further, 
modern ecologists are aware that no ques- 
tions can be answered by studying only a 
few “‘representative’’ organisms, usually se- 


VOLUME 102, NUMBER 3 


lected among “easily identified’’ organisms, 
such as some crustaceans, echinoderms and 
mollusks, as done in the early days of ben- 
thic ecology. 

Simple thoughts on theory. —Organisms 
may be studied in four different ways, which 
may be organized into two systems of two. 
First, one may either attempt to describe 
the structural characteristics of an organ- 
ism, or one may study interactions among 
structures in time or space. The other sys- 
tem of classifying observations describes the 
investigational intent. One may study the 
same process in a variety of organisms; or, 
alternatively, one may study a variety of 
processes and structures in the same kinds 
of organisms. 

Structural descriptions historically start- 
ed with external morphology, and pro- 
ceeded via internal anatomy to microscopic 
anatomy in all its phases. Structural de- 
scriptions deal with the material presence 
of anything, including atoms and subatomic 
particles. In gross morphological descrip- 
tions the unaided eye is used; all other de- 
scriptions are based on interpretation of im- 
ages created by various pieces of gear: 
microscopes of all kinds, meters and dials 
and color-reactions, spectrophotometers, or 
small patches of color on a starch gel. The 
more highly magnified the analysis be- 
comes, the more remote the interpretation 
of the findings become from normal human 
experiences, but, at least in theory, no dif- 
ferent from observations of gross morphol- 
ogy. In some sense, interpretation becomes 
easier with increasing magnification, since 
the higher magnification allows a far more 
precise use of language in describing limit- 
ing conditions than do observations of a 
morphological or anatomical nature. 

Natural historians and some physiolo- 
gists (a subgroup of the comparative and 
ecological physiologists) seek a completely 
different kind of information about organ- 
isms, information which we have had a great 
deal of difficulty entering into our structur- 
ally derived patterns. All organisms change 


747 


with time and all structural landmarks 
change in relation to each other during on- 
togenesis, presumably in an organized fash- 
ion, but not necessarily in the same pattern 
even in genetically similar organisms. In- 
formation derived from these changes is as 
much an expression of the genome of the 
organism as is the structural information. I 
am aware of the problems including this 
kind of information in our descriptions will 
create, but I believe that until we do, we will 
fall short of understanding the organisms we 
are studying. Computerized modelling may 
offer help in creating testable predictions for 
such studies. 

The other system of groupings of study is 
familiar to most scientists, especially in 
technically more complex fields. Scientists 
become experts on the use of a single tech- 
nique: transmission and scanning electron 
microscopes, enzyme electrophoresis, DNA 
hybridization and so forth and will inves- 
tigate the limits of what the technique can 
do. The results of this approach have been 
excellent and have lead to major advances 
in our understanding of microstructures and 
varlous processes. 

The other major way of looking at the 
organisms is as a specialist on a single an- 
imal group; a taxon-oriented person. Such 
a person may be eclectic in their use of tech- 
niques, but will rarely add to the develop- 
ment of new techniques. These biologists 
often have a better understanding of the 
evolutionary significance of differences in 
processes among the organisms studied than 
the process-oriented scientists, but are usu- 
ally rather parochial in their view of the 
world. A polychaete’s-eye view of the globe 
is limiting in many ways. 

These four ways of studying organisms 
do not agree with the traditional breakdown 
of specialities among biologists. Taxono- 
mists, while primarily concerned with de- 
scription of structure, frequently resort to 
adaptive explanations. Physiologists, while 
exploring functional issues, base themselves 
in knowledge of the structures involved in 


748 


the particular processes studied. Perhaps 
most confused are the activities that are now 
subsumed under the heading of ecology. In 
part, ecologists describe structure in their 
case patterns of distribution of organisms 
in nature, but usually use functional expla- 
nations for the patterns demonstrated. The 
separation of the two modes of thinking is 
not trivial, but is built into the language. 
Ideally a language describing structure 
should use only shape and position words; 
in practice we use such words as “‘bran- 
chiae”’ and “‘notopodial cirri.’’ For trained 
taxonomists and morphologists the usual 
meanings of these words have become triv- 
ial: they are using both words as shape and 
position markers. However, notopodial cir- 
ri, usually slender, often very long cirri pro- 
jecting from the dorsolateral sides of the 
worms, often appear to be as much respi- 
ratory as sensory in function. 

Eclecticism and the study of poly- 
chaetes.—Thus an adequate description of 
any polychaete would require a rather eclec- 
tic collection of pieces of information, both 
static and dynamic. 

Most structural descriptions of poly- 
chaetes now include a minimal mention of 
major morphological features. At least one 
species of most families have been studied 
anatomically, at least at the light micro- 
scope level. Very few truly comparative 
studies have been performed within each 
family. Comparative studies among the 
families are rather common, but without 
knowledge of how much variation to expect 
within each family, the interpretation of such 
comparative studies will always be difficult. 
Microanatomical studies are becoming 
rather more common, but again, with some 
very Salutary exceptions, have focussed more 
on the relations among the families. Other 
studies, with both structural and functional 
components, are mentioned below. 

Studies of comparative physiology have 
given us important information about the 
interactions among the structures, e.g., 
studies of mechanisms of respiration among 
polychaetes. However, most physiological 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


studies have been focussed more on eluci- 
dating process and are for that reason usu- 
ally not very useful for comparative pur- 
poses. Most life history studies published so 
far include an account of parts of the larval 
development and metamorphosis into a 
postlarvae, but little about the rest of the 
life of the organisms, including longevity 
(Fauchald 1983). The bits we have are in- 
teresting, but are insufficient for all species. 

I am advocating eclecticism because I be- 
lieve that this approach will force us to 
change our approach to our studies. Cur- 
rently we learn one, or perhaps a few, tech- 
niques and then proceed to apply these to 
all problems, whether the application can 
solve the problems posed or not. The in- 
vestigative technique and the detail sought 
must depend on the question asked, rather 
than the other way around. For example, it 
is not always useful or necessary to identify 
organisms to the species level in a benthic 
investigation. The first step in planning a 
study therefore must be to question the pur- 
pose of the investigation. If the purpose is 
an exploration of the area—a study of which 
organisms are present in what quantities— 
then identification to species is not only de- 
sirable, but the only way such information 
should be presented. But if the purpose is 
to investigate feeding biology or perhaps 
trophic structure, in addition to giving a 
listing of taxa present, at the very least as 
much effort must be put into investigation 
of gut contents and mechanisms of feeding, 
as into the identification of the specimens. 
Most investigators now identify their or- 
ganisms (more or less accurately) and then 
quote some authority for the other infor- 
mation needed, e.g., feeding physiology. For 
the polychaetes, most quote Fauchald & Ju- 
mars (1979), an inappropriate source of in- 
formation for this purpose. The Diet of 
Worms was written as a summary of what 
little information was available in the mid 
1970’s and was intended to spur investi- 
gations: It has apparently done so, but suf- 
ficient information is still not available for 
any species to my knowledge. 


VOLUME 102, NUMBER 3 


Polychaetes are valuable for a variety of 
studies. Polychaetes are ubiquitous and 
common in all marine environments. The 
numbers of polychaete taxa is large enough 
to allow the use of the statistical data re- 
duction, but is not as overwhelming as in 
some other groups. Most major subgroups 
have morphologically very strict body plans 
and can be identified to family by rank ty- 
ros. 

The group is very old (Fauchald 1984) 
and the major body plans were laid down 
a long time ago: We can in the polychaetes 
investigate current evolution of ancient body 
plans. For example, the eunicids are very 
uniform in general morphological appear- 
ance; in fact, the jaws have not changed 
much since Palaeozoic times. Nevertheless, 
a preliminary numerical study of about 300 
individuals of approximately 12 species 
(Fauchald 1989) demonstrate several dif- 
ferent patterns of growth and of control of 
the body proportions, implying rather dif- 
ferent physiological properties, perhaps re- 
lated to the maximum absolute size of each 
species. 

The consequences of the studies of Gras- 
sle & Grassle (1976) and Eckelbarger & 
Grassle (1987), to mention only two of a 
series, are fascinating. They have given us 
a view of a worldwide group of small, ever- 
changing populations of capitellids becom- 
ing isolated, perhaps going extinct locally, 
perhaps meeting up again before, or after, 
completing a speciating process—in short, 
a complex mosaic. 

Chromosome studies of various poly- 
chaetes indicate that ploidy relations may 
play a more important part in evolution in 
polychaetes than previously expected; per- 
haps leading to a reconsideration of the im- 
portance of the various processes in the evo- 
lution of animals. 

An eclectic approach may thus complete 
the transformation of the study of poly- 
chaetes from an intellectual backwater to 
the forefront of biology. 

Some final notes. —I agree with my alter 
ego of 25+ years ago that theory of science 


749 


exists with little reference to what I do on 
a day to day basis. I have come to the re- 
alization that this is perhaps the way it ought 
to be. If the theory of science was strictly a 
description of what scientists do, then one 
could not expect discussion of normative 
rules. We all use theoretical constructs in 
even the simplest observations. The belief 
in theory-independent observations ap- 
pears now on the wane. Philosophers of sci- 
ence study and perhaps build into systems 
the theories behind our observations and 
make us as working scientists aware of these 
constructs. Without the precision in think- 
ing and data definition theory enforces, very 
little advance 1s possible. 

A significant fraction of current papers are 
routine descriptions of a few new taxa, usu- 
ally with a review paper as authority for the 
separate status of the new taxa; the mate- 
rial examined is minimal and comparison 
with types of previously described species 
is rare. If current theory and methods were 
applied to these studies, Iam convinced that 
the deluge of new taxa would slow down. 
Most of the new taxa are collected during 
quantitative investigations and the authors 
do not have the luxury of performing a com- 
plete and detailed review of the family or 
genus of interest before publishing a new 
taxon or two. Detailed and rigorously per- 
formed reviews of previously described taxa 
are lacking for nearly all polychaete families 
and very few are now on the horizon. Most 
of the investigations in which the bulk of 
new material is collected have poorly, or 
inappropriately defined, goals: however, one 
requirement runs through most of them: No 
matter what the stated purpose of the in- 
vestigation is, the organisms collected must 
be identified to species. This requirement 
forces the researches to make rapid, often 
incorrect decisions. A careful definition of 
study goals would leave both ecologists and 
polychaetologists happier and the few poly- 
chaetologists working full time on poly- 
chaete taxonomy less overwhelmed. 

There is little support for all the other 
kinds of studies needed to describe and study 


750 


polychaetes adequately. The result is that 
most of the polychaetologists are limping 
along, without being able to do even the 
necessary revisory work, and certainly with- 
out being able to apply theory or attempt 
to add truly new information to our de- 
scriptions of polychaetes. A rather sad con- 
clusion, but I believe one in which experts 
on other groups also would concur. 


Acknowledgments 


This paper is based on the second Riser 
Lecture, given at The Marine Science Cen- 
ter, Northeastern University, October 30, 
1986. It is dedicated to Dr. Nathan W. Ris- 
er, one of the finest polychaete experts liv- 
ing; Pete Riser combines a morphologist’s 
eye with a focus on live organisms. 

I would like thank the committee, Drs. 
M. Patricia Morse, Joan Ferraris and Jon 
Norenburg for giving me the opportunity to 
give the lecture. Dr. Leonard P. Hirsch read 
and rejected several versions of the paper, 
for which I am now very grateful; what 
makes sense in this paper is due to my 
friends; what appears hare-brained is my 
own responsibility. 


Literature Cited 


Akesson, B. 1982. A life table study on three genetic 
strains of Ophryotrocha diadema (Polychaeta, 
Dorvilleidae).— International Journal of Inver- 
tebrate Reproduction 5:59-69. 

Audouin, J. V., & H. Milne Edwards. 1834. Re- 
cherches pour servir a |’Histoire Naturelle du 
Littoral de la France, ou recueil de Mémoires 
sur l’anatomie la physiologie, la classification et 
les moeurs des animaux de nos cétes; ouvrage 
accompagné du planches faites d’aprés nature. 
Paris, Crochard, Libraire 2: Annélides Premiére 
Partie 1-290, 18 pls. 

Augener, H. 1918. Polychaeta. In W. Michaelsen, 
ed., Beitrage zur Kenntnis der Meeresfauna 
Westafrikas. 2(2):67-625, 6 pls. 

Binard, A., & R. Jenner. 1928. Morphologie du lobe 
préoral des Polychétes.—Receuil des Institut 
Zoologique Torley-Rousseau, Bruxelles 2:118- 
240, 8 pls. 

Blegvad, H. 1930. Quantitative investigations of bot- 
tom invertebrates in the Kattegat with special 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


reference to the plaice food.—Reports of the 
Danish Biological Station 36:3—-56. 

Claparéde, E. 1854. Résumé des travaux les plus ré- 
cents sur la generation alternante et sur les mé- 
tamorphoses des animaux inférieurs. —Ar- 
chives des Sciences Physiques et Naturelles, 
Genéve 25:229-263, 313-330. 

[Claparéde, E.] 1865. Revew of: Quatrefages, A. de. 
Notes sur la classification des Annelides 
(Comptes Rendus des Séances de l’Academie 
des Sciences, Paris, 27 mars 1865).—Biblio- 
théque Universelle et Revue Suisse— Archives 
des Sciences Physiques et Naturelles, Genéve, 
series 2 22:346-355. 

Cuvier, G. 1817. La Régne Animal distribué d’aprés 
son organisation, pour servir de base a l’histoire 
naturelles des animaux et d’introduction a 
lanatomie comparée. Paris, Deterville 2:X VIII 
and 1-532. 

Dales, R. P. 1969. Respiration and energy metabo- 
lism in annelids. Jn M. Florkin & B. T. Scheer 
eds.—Chemical Zoology 4(3):93-110. 

Eckelbarger, K. J., & J. F. Grassle. 1987. Interspecific 
variation in genital spine, sperm, and larval 
morphology in six sibling species of Capitella. — 
Bulletin of the Biological Society of Washington 
7:62-76. 

Ehlers, E. 1864-1868. Die Borstenwiirmer (Annelida 
Chaetopoda) nach systematischen und anato- 
mischen Untersuchungen dargestellt. Wilhelm 
Engelmann, Leipzig, XX and 1-722, 24 pls. (pp. 
1-268 issued in 1864; the rest in 1868). 

Fabricius, O. 1780. Fauna Groenlandica, systematice 
sistens, Animalia Groenlandiae occidentalis 
hactenus indagata, quad nomen specificum, 
triviale, vernaculumque; synonyma acutorum 
plurium, descriptionem, locum, victum, genera- 
tionem, mores, usum, capturamque singuli; 
prout detegendi occasio fuit, maximaque parti 
secundum proprais observationes. Havniae, X VI 
and 1-452. 

Fauchald, K. 1983. Life diagram patterns in benthic 

Polychaetes. — Proceedings of the Biological So- 

ciety of Washington 96(1):160-177. 

1984. Polychaete distribution patterns, or: 
Can animals with Palaeozoic cousins show large- 
scale geographical patterns. Pp. 1-6 in P. A. 
Hutchings, ed., Proceedings of the First Inter- 
national Polychaete Conference, Sydney, Aus- 
tralia, July 1983, published by The Linnean So- 
ciety of New South Wales. 

1989. Variability of morphological features 
in eunicid polychaetes. — Ophelia (special issue, 
in press). 

——., & P.A.Jumars. 1979. The diet of worms: A 
study of polychaete feeding guilds.—Oceanog- 
raphy and Marine Biology: An Annual Review 
17:193-284. 


VOLUME 102, NUMBER 3 


Fauvel, P. 1923. Polychétes errantes.—Faune de 

France 5:1-488, 188 figs. 

1927. Polychétes sedentaires. Addenda aux 
Errantes, Archiannélides, Myzostomaires.— 
Faune de France 16:1-494, 152 figs. 

Florkin, M. 1969. Respiratory proteins and oxygen 
transport. Jn M. Florkin and B. T. Scheer eds. — 
Chemical Zoology 4(4):111-134. 

Gillet, P. 1988. Ongine et histoire des Annélides 
Polychétes (vers marine).—Impacts 1:55-62. 

Grassle, J. F., & J. P. Grassle. 1976. Sibling species 
in the marine pollution indicator Capitella.— 
Science 192:567-569. 

Grube, A.-E. 1850. Die Familien der Anneliden.— 
Archiw ftir Naturgeschichte, Berlin 16(1):249- 
364. 

Hanstrom, B. 1928. Die Beziehungen zwischen dem 
Gehirn der Polychaeten und dem der Arthro- 
poden.— Zeitschrift fiir Morphologie und Oe- 
kologie der Tiere, Berlin 11:152-160. 

Hatschek, B. 1893. System der Anneliden, ein vor- 
laufiger Bericht. Lotos, Prag 13:123-126. 

Kinberg, J. G. H. 1865. Annulata nova.—6fversigt 

af Kongliga Svenska Vetenskaps-Akademiens 

Forhandlingar 21:559-574. 

1910. Annulater. Kongliga Svenska Fregat- 
ten Eugenies resa omkring jorden under befal 
af C. A. Virgin Aren 1851-1853. Vetenskapliga 
Iakttagelser pa Konung Oscar, Den Forstes be- 
fallning utgifna af K. Svenska Vetenskapsaka- 
demien.—2. Zoologi 3:1—78, 29 pls. (pp. 1-32, 
pls. 1-8 issued in 1858). 

Lamarck, J. B. de. 1816. Histoire naturelle des An- 
imaux sans Vertébres présentant les caractéres 
généraux et particuliers de ces animaux, leur 
distribution, leurs classes, leur families, leur 
genres, et la citation des principales espéces qui 
s’y rapportent; précédée d’une Introduction of- 
frant la détermination des caractéres essentiels 
de l’Animal, sa distinction du végétal et des autres 
corps naturels, enfin, l’°exposition des principes 
fondamenteux de la Zoologie. Paris 2:1—568. 

Linnaeus, C. 1758. Systema naturae per regna tria 
naturae, secundum classes, ordines, genera, 
species, cum characteribus, differentiis, synon- 
ymis, locis. Editio Decima, Reformata. Vol. 1: 
1-824. Holmiae, Laurentii Salvii. 

Malmgren, A. J. 1867. Annulata Polychaeta Spets- 
bergiae, Groenlandiae, Islandiae et Scandina- 
viae hactenus cognita.—Ofversigt af Kongliga 
Svenska Vetenskaps-Akademiens Forhandlin- 
gar 24:127-235, pls. 2-15. 

McIntosh, W.C. 1885. Report on the Annelida Poly- 
chaeta collected by H.M.S. Challenger during 
the years 1873-—76.—Challenger Reports 12:1- 
554, pls. 1-55, la—39a. 

Meyer, E. 1890. Die Abstammung der Anneliden. 
Der Ursprung der Metamerie und die Bedeu- 


751 


tung des Mesoderms.—Biologische Central- 
blatt, Leipzig 10:296-308. 

Miller, O. F. 1776. Zoologica Danicae. Prodromus 
seu Animalium Daniae et Norvegiae indigena- 
rum charactens, nomine et synonyma imprimis 
popularium. Havnae, XXXII and 1-274. 

Needham, A. E. 1969. Growth and development. Jn 
M. Florkin & B. T. Scheer, eds.— Chemical Zo- 
ology 4(11):377-442. 

Pallas, P.S. 1766. Miscellanea Zoologica, quibus no- 
vae imprimis atque obsurae animalium species 
describuntur et observationibus iconibusque il- 
lustrantur. Hagae Comitum XII and 1-224. 

Petersen, C.G. J. 1911. Valuation of the sea. Animal 
life of the sea-bottom, its food and quantity. — 
Reports of the Danish Biological Station 20:1- 
80. 

Platt, J. R. 1964. Strong inference.—Science 146 
(3642):347-353. 

Popper, K. 1979. Objective knowledge. An Evolu- 
tionary approach. Revised Edition. Oxford at 
the Clarendon Press X and 1-395. 

Quatrefages, A. de. 1865a. Note sur la classification 

des Annélides.— Comptes Rendu des Séances de 

l’Academie des Sciences Naturelles, Paris 40(1 3): 

586-600. 

1865b. Note sur la classification des anné- 
lides, et reponse aux observations de M. Cla- 
paréde.— Annales des Sciences Naturelles, Paris, 
series 5 3:253-296. 

1866 [erroneously dated 1865]. Histoire na- 
turelle des Annéles marins et d’eau douce. An- 
nélides et Gephyriens. Paris, Libraire Ency- 
clopedique de Roret 1:VII and 1-588. 

Reish, D. J. 1958. An analysis of a sample of system- 
atic literature, 1758 to 1954.—Systematic Zo- 
ology 7:174-179. 

Savigny, J.C. 1820. Systéme des Annélides, princi- 
palement de celles des cdtes de l’Egypte et de la 
Syrie, offrant les caractéres tant distinctifs que 
natureles des ordres, familles et genres, avec la 
description des especes.—Description de 
l’Egypte. Histoire Naturelles 21:325-472 (front 
page of volume dated 1809, 1(3) first issued in 
1820; 2nd edition in 1826). 

Schmarda, L. 1861. Neue Wirbellose Thiere beo- 
bachtet und gesammelt auf einer Reise um die 
Erde 1853 bis 1857, I. Neue Turbellarian, Ro- 
tatorien und Anneliden beobachtet und gesam- 
melt auf einer Reise um die Erde 1853 bis 1857. 
(2):1-164, pls. 16-37. 

Schroeder, P. C., & C.O. Hermans. 1975. Annelida: 
Polychaeta. Jn A. C. Giese & J. S. Pearse eds., 
Reproduction of marine invertebrates. Aca- 
demic Press 3:1-213. 

Thorson, G. 1957. Bottom communities. Jn J. Hedg- 
peth, ed. Treatise of marine ecology and paleo- 
ecology 1.—Ecology (17):461-534. 


752 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Treadwell, A. L. 1901. The cytogeny of Podarke ob- opment of annelids and polyclades.— Annals of 
scura.—Journal of Morphology, Boston 17:399- the New York Academy of Science 11:1-27. 
486. 


Williams, T. 1851. Report on the British Annelida. — D 
; ice epartment of Invertebrate Zool - 
Report of the British Association for the Ad- Pp ate Zoology, Na 


vancement of Science 21:159-272. tonal Museum of Natural History, Smith- 
Wilson, E. B. 1898. Considerations on cell lineage SONaN Institution, Washington, D.C. 20560. 


and ancestral reminiscence, based on a reex- 
amination of some points in the early devel- 


PROC. BIOL. SOC. WASH. 
102(3), 1989, pp. 753-760 


A NEW SPECIES OF EUCHONE (POLYCHAETA: SABELLIDAE) 
FROM THE NORTHWEST ATLANTIC WITH COMMENTS 
ON ONTOGENETIC VARIABILITY 


R. Eugene Ruff and Betsy Brown 


Abstract. —Euchone banséi, a new species of the polychaete family Sabellidae, 
is described from the continental slope and rise between Cape Cod, Massa- 
chusetts, and Cape Lookout, North Carolina. Juvenile and adult specimens are 
examined and ontogenetic variability is discussed. It is demonstrated that the 
number of abdominal depression setigers, the shape of the collar, and the 
number of radioles are not valid diagnostic characters for the identification of 


juveniles. 


In response to interest in offshore oil and 
gas development, the Minerals Manage- 
ment Service (MMS) of the U.S. Depart- 
ment of the Interior sponsored research on 
benthic communities on the continental 
slope and rise (SO0—3000 m) off the eastern 
United States in three regions: (1) the U.S. 
North Atlantic near Georges Bank off Mas- 
sachusetts, (2) the U.S. Mid-Atlantic off New 
Jersey, and (3) the U.S. South Atlantic off 
the Carolinas. As is typical in marine soft- 
bottom environments (Knox 1977), the 
macrofaunal communities in these regions 
are dominated by polychaetous annelids 
(Maciolek et al. 1987a, b; Blake et al. 1987). 
Because of the numerous samples collected 
and the small sieve mesh (300 wm) used 
throughout this sampling program, juvenile 
growth stages of many polychaete species 
were routinely collected. This paper de- 
scribes a new species of Euchone (Poly- 
chaeta: Sabellidae) collected between 1345- 
2495 m depth along the U.S. Atlantic coast 
and examines some of the ontogenetic vari- 
ability exhibited by this species. 


Euchone bansei, new species 
Figs./T."2 


Euchone spp. Hartman & Fauchald, 1971: 
179 [partim]. 


Euchone sp. 3. Maciolek et al., 1987a, b.— 
Blake et al., 1987. 


Material examined. —off Martha’s Vine- 
yard, 4 May 1966, Chain station Ch 103, 
39°43.6'N, 70°37.4’W, 2022 m, 8 speci- 
mens; 7 Sep 1963, Atlantis station A 58, 
3634.5 NY 12°55.0'W.,¥ 2000: 75 m3 
specimens; near Baltimore Canyon, 19 May 
1985, cruise MID-4 station 13-2, 
37°53.29'N, 73°45.30'W, 1607 m, clayey 
mud, Holotype (USNM 115738); 19 May 
1985, cruise MID-4 station 10-3, 
37°51.73'N, 73°20.01'W, 2095 m, silty mud, 
5 paratypes (USNM 115739); 16 Nov 1985, 
cruise MID-6 station 10-1, 37°51.77'N, 
73°20.01'W, 2104 m, silty mud, 9 paratypes 
(BMNH ZB 1987.620-628); near Linden- 
koehl Canyon, 16 May 1985, cruise MID-4 
station 3-2, 38°36.75'N, 72°51.57'W, 2055 
m, silty mud, 6 paratypes (BMNH ZB 
1987.629-634); 17 May 1985, cruise MID-4 
Station 3-3, 38°36.75'N, 72°51.60’W, 2052 
m, silty mud, 9 paratypes (USNM 115740); 
17 May 1985, cruise MID-4 station 11-1, 
38°40.10'N, 72°56.43'W, 1510 m, clayey 
mud, 22 paratypes (USNM 115741); 7 Aug 
1985, cruise MID-5 station 12-2, 
38°29.25'N, 72°42.22’W, 2495 m, sandy 
mud, 3 paratypes (USNM 115742); off Cape 
Cod, 25 Jul 1986, cruise NOR-6 station 


754 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 1. Euchone bansei (holotype, USNM 115738): A, Entire animal in lateral view showing the right half 
of branchial crown; B, Collar and anterior region with branchiae not illustrated; C, Posterior region showing 
the anal furrow. 


3-1, 41°01.55'N, 66°20.12'W, 1345 m, silty m,sandy mud, 9 paratypes (USNM 115744); 
mud, | paratype (USNM 115743); near Ly- off Cape Lookout, 23 May 1985, cruise SA-4 
donia Canyon, 29 Apr 1985, cruise NOR-2 | station 4-3, 34°11.29'N, 75°38.67'W, 2015 
station 6-2, 40°05.03'N, 67°29.13’W, 2108 mm, silty mud, 2 paratypes (USNM 115745). 


VOLUME 102, NUMBER 3 


recs RS — 


wt 


= *. CE eee cey eee Oe A eek =" 
ZT_S_S SS 


0.05mm 


Fig. 2. Euchone bansei (paratype, BMNH ZB 1987.620): A, Thoracic notosetae from setiger three, including 


long and short limbate setae and a narrowly limbate bayonet seta; B, Thoracic neuroseta from setiger three; C, 
Uncini from abdominal setiger four in profile and in frontal view. 


Diagnosis. —Small Euchone species with 
17 abdominal setigers, last 6 associated with 
anal depression. Branchial crown with 4 
pairs of radioles united with palmate mem- 
brane for half their length; radioles with long, 
filiform, pinnule-free ends. Collar entire lat- 


erally and ventrally, separated by mid-dor- 
sal gap, of even height all around, extending 
to branchial basis. Ventral shields absent. 
Thoracic notopodia with long and short 
limbate setae and pointed bayonet setae. 
Abdominal uncini with quadrate base, small 


755 


756 


main fang, and crest of numerous smaller 
teeth. 

Description. —Adult holotype 4.0 mm in 
length (excluding radioles), 0.5 mm in width 
along posterior thorax (Fig. 1A). Other adult 
specimens 2.2—5.9 mm in length, 0.3-0.8 
mm in width; branchial crown contributing 
additional 4.4 mm in largest individuals. 
Four pairs of radioles connected for half 
their length by delicate palmate membrane; 
each radiole with about 20 subequal pin- 
nules alternating along axis and a pinnule- 
free filiform tip contributing up to half of 
total radiole length. Dorsal lips with elon- 
gate-triangular radiolar appendages, about 
one-half pinnule length, densely ciliated; 
pinnular appendages not observed. Paired 
ventral lips shorter, rounded. 

Peristomium produced into triangular 
projection ventral to radioles. Collar entire 
ventrally and of nearly equal height laterally 
and ventrally in adult (Fig. 1B), reaching 
approximately to base of branchiae; dorsal 
gap very narrow; otocysts not observed. First 
setiger one-half length and narrower than 
following thoracic setigers. Thin postsetal 
glandular girdle encircling second setiger. 
Ventral shields absent. 

Abdomen with 17 setigers, last 6 associ- 
ated with anal depression; wings of depres- 
sion flaring, connected anteriorly by thin 
membrane (Fig. 1C). Pygidium triangular, 
slightly longer than wide. 

First setiger with single bundle of long, 
narrow limbate setae and shorter bayonet 
setae; other thoracic notosetae of three kinds: 
superior arc of long limbate setae, and in- 
ferior group of short limbate setae and slen- 
der pointed bayonet setae with narrow wings 
(Fig. 2A). Thoracic neuropodia with about 
12 long-handled acicular uncini in each fas- 
cicle (Fig. 2B). Abdominal notopodial tori 
with 10-17 uncini, each with quadrate base 
and small main fang surmounted by nu- 
merous rows of smaller teeth (Fig. 2C); shape 
of uncini nearly constant within each torus, 
but main fang progressively smaller in pos- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


terior setigers. Abdominal neuropodia with 
3-4 narrowly limbate setae per fascicle. 

Sexes separate with gametes occurring 
laterally between dorsal and ventral longi- 
tudinal muscle bundles in posterior thoracic 
and anterior abdominal segments. Eggs ir- 
regular in outline, up to 0.1 mm in diameter. 
Sperm short-headed, with blunt acrosomes, 
approximately 5 um long excluding flagella. 

Methyl green stain in 70 percent ethanol 
(Banse 1970) readily accepted both dorsally 
and ventrally on most of body including 
pygidium and wings of anal depression. 
Staining cells absent in radioles, at anterior 
margin of collar, in glandular girdle, in in- 
tersegmental furrows, along fecal groove, 
around parapodia, and in narrow band of 
cells encircling each segment at level of se- 
tae. 

Distribution. —Euchone bansei is found 
from the southern flank of Georges Bank off 
Cape Cod, Massachusetts, southward to 
Cape Lookout, North Carolina. The species 
occurs in sandy to clayey muds in depths of 
1345-2495 m and is found most often at 
about 2000 m in abundances occasionally 
exceeding 100/m7?. 

Etymology.—This species is named in 
honor of Dr. Karl Banse in recognition of 
his significant contributions to the knowl- 
edge and taxonomy of the Sabellidae. 


Ontogenetic Variability 


The routine use of 300-wm mesh sieves 
throughout the MMS sampling program re- 
sulted in the collection of a large number of 
sub-adult and juvenile specimens of Eu- 
chone bansei. After selection of the type ma- 
terial, an additional 185 specimens from 
across the sampling region were examined 
in detail. Ocular micrometer measurements 
were made on the total length from the top 
of the collar to the tip of the pygidium and 
on the width of the last thoracic setiger. 
Counts were made on the number of tho- 
racic and abdominal setigers, on the number 


VOLUME 102, NUMBER 3 


6.0 
2 
i= 
D 5.0 
= 
O 
Q 
<x 
em 
O 4.0 
z 
ra 
==) 
= 
2 30 
x 3. 
2 
= 
O 
z 
2.0 
x 
S 
Ee 


4ST 


eo © 00 eg ge ame pp fageee § 0 co co 


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 


758 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


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 


VOLUME 102, NUMBER 3 


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. 


762 


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 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


764 


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VOLUME 102, NUMBER 3 


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766 


BUCCAL APPARATUS 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


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 


: <a wi 


TA Maes 
6 . 


773 


a se 


Twin Cays: 


4 
i 


uv 


4 
, 


4 
2 


Fig. 1. Study area in Belize. The box to the left indicates the position of the study areas in relation to Belize; 
the study area is enlarged on the right. The two arrows in the enlargement indicates West Bay, Twin Cays and 
the un-named cay in the Blue Ground Range at which the collections were made. 


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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Fig. 3. Relationship between length through setiger 
15 and number of thoracic setigers in N. setosa from 
Belize and Bermuda. 


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 


— 


783 


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. 


784 


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. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


786 


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 


787 


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. 


788 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


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- 


VOLUME 102, NUMBER 3 


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- 


VOLUME 102, NUMBER 3 


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- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


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 


a99 


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 


800 


i , x 
f a BS 
* rf . + i x 


= 


oo 
aS 
a = 
SS << = = 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


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 


VOLUME 102, NUMBER 3 


801 


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- 


802 


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 


803 


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 


804 


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- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


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- 


806 


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). 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


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 


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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 
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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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INFORMATION FOR CONTRIBUTORS 


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Eigenmann, C.H. 1915. The Cheirodontidae, a subfamily of minute characid fishes of South 

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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 

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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 
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Plants: David B. Lellinger Raymond B. Manning 

Insects: Wayne N. Mathis Vertebrates: G. David Johnson 


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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- 


816 


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. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


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 


VOLUME 102, NUMBER 4 


to its own family. Also, Carlgren (1949) used 


817 


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 


818 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


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. 


VOLUME 102, NUMBER 4 


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 


819 


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 


822 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


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. 


VOLUME 102, NUMBER 4 


A B 


823 


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, 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


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. 


Literature Cited 


Carlgren, O. 1921. Actiniaria Part 1.—The Danish 

Ingolf-Expedition 5:1-241. 

. 1949. A survey of the Ptychodactiaria, Cor- 

allimorpharia and Actiniaria.—Kungl. Svenska 

Vetenskapsakademiens Handlingar, Series 4, 

1(1):1-121. 

. 1956. Actiniaria from depths exceeding 6000 

meters.—Galathea Report 2:9-16. 

. 1959. Corallimorpharia and Actiniaria with 

description of a new genus and species from 

Peru.—Lunds Universitets Arsskrift, n.f. Avd. 

2, 56(6):1-38. 

Childress, J. J., H. Felbeck, & G. N. Somero. 1987. 
Symbiosis in the deep sea. — Scientific American 
255:115—120. 

Danielssen, D.C. 1890. Actinida.— Norwegian North- 
Atlantic Expedition 1876-1878, Zoology 19:1- 
184. 

Doumenc, D., & M. Van-Praét. 1988. Actinies abys- 
sales d’un site hydrothermal du Pacifique ori- 
ental.—Oceanologica Acta special volume 8:61- 
68. 

Hertwig, R. 1882. Report on the Actiniaria dredged 
by H.M.S. Challenger during the years 1873- 
1876.—Scientific Results of the Voyage of 
H.M.S. Challenger, Zoology 6(1):1-136. 

Hessler, R. R., P. Lonsdale, & J. Hawkins. 1988. Pat- 
terns on the ocean floor.— New Scientist 1605: 
47-51. 

——., & W. M. Smithey, Jr. 1983. The distribution 


VOLUME 102, NUMBER 4 


and community structure of megafauna at the 
Galapagos Rift hydrothermal vents. Pp. 735- 
770 in P. A. Rona, K. Bostrom, L. Laubier, & 
K. L. Smith, Jr., eds., Hydrothermal processes 
at seafloor spreading centers. Plenum Press, New 
York and London. 

Lewis, J. B. 1984. Photosynthetic production by the 
coral reef anemone, Lebrunia coralligens Wil- 
son, and behavioral correlates of two nutritional 
strategies. — Biological Bulletin 167(3):601-612. 

Muscatine, L. 1974. Endosymbiosis of cnidarians and 
algae. Pp. 359-395 in L. Muscatine & H. M. 
Lenhoff, eds., Coelenterate biology: Reviews and 
new perspectives. Academic Press, New York. 

Parry, G. 1952. The Actiniaria of New Zealand. A 
check-list of recorded and new species, a review 
of the literature and a key to the commoner 
forms. Part 2.—Records of the Canterbury Mu- 
seum 6(2):121-141. 

Riemann-Zurneck, K. 1978. Tuiefsee-Aktinien der 
Familie Actinoscyphiidae aus dem Nordatlan- 
tik (Actiniaria, Mesomyaria).— Zoologica Scrip- 
ta 7:145-153. 


825 


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 


VOLUME 102, NUMBER 4 


Table 1.—Continued. 


829 


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 


830 


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- 


832 


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., 


VOLUME 102, NUMBER 4 


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 


839 


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 & 


840 


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. 


VOLUME 102, NUMBER 4 


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- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


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 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


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. 


VOLUME 102, NUMBER 4 


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. 


846 


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- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


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 


VOLUME 102, NUMBER 4 


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. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


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- 


VOLUME 102, NUMBER 4 


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 


850 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


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 


VOLUME 102, NUMBER 4 


(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 


852 


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 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


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. 


VOLUME 102, NUMBER 4 


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 


VOLUME 102, NUMBER 4 


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 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


& 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 


Literature Cited 


Akesson, B. 1958. A study of the nervous system of 
the Sipunculoideae, with some remarks on the 
development of the two species Phascolion 
strombi Montagu and Golfingia minuta Kefer- 
stein. — Undersékningar 6ver Oresund 38:1-249. 

Andrew, M., & N. V. Andrew. 1953. Some annelid 
and sipunculid worms of the Bimini region.— 
American Museum Novitates 1617:1-16. 

Amor, A. 1975. Notas sobre Sipuncula de la Argen- 
tina, Brasil y Peru.—Physis 34(88):113-—120. 

Augener, H. 1903. Beitrage zur Kenntnis der Ge- 
phyreen nach Untersuchung der im Gottinger 
zoologischen Museum befindlichen Sipunculi- 
den und Echiuriden.— Archiv ftir Naturgesch- 
ichte 69:297-371. 

Baird, W. 1868. Monograph on the species of worms 

belonging to the subclass Gephyreae. — Proceed- 

ings of the Zoological Society of London 1868: 

77-114. 

1873. Descriptions of some new species of 
Annelida and Gephyrea in the collections of the 
British Museum.—Journal of the Linnean So- 
ciety of London, Zoology 11:94—97. 

Benham, W. B. 1912. Report on sundry invertebrates 
from the Kermadec Islands. — Transactions and 
Proceedings of the New Zealand Institute 44: 
135-138. 

Blainville, H. M. D. de. 1827. Dictionnaire des sci- 
ences naturelles.— Vers 49:305-313. 

Bovier, E. L. 1894. Un nouveau cas de commensal- 
isme: association de vers du genre Aspidosiphon 
avec des Polypes madréporaires et un Mol- 
lusque bivalve.—Comptes Rendus Academié des 
Sciences, Paris 119:96-98. 

Chamisso, A. de, & C. G. Eysenhardt. 1821. De an- 
imalibus quisbusdam e classe vermium lin- 
neana.—Nova Acta Academia Caesarea Leo- 
poldino 10:343-374. 

Chapman, G. 1955. Aspects of the fauna and flora 
of the Azores. III. Gephyrea. — Annals and Mag- 
azine of Natural History 8(12):351-352. 

Collin, A. 1892. Gephyreen gesammelt von Herrn 
Stabsarzt Dr. Sander auf der Reise S.M.S. Prinz 
Adalbert.—Archiv fur Naturgeschichte 58(1): 
177-182. 

Cordero, E. H., & A. Mello-Leitao. 1952. Duas navas 
especies do genero Aspidosiphon da Ihla Trin- 
idade.—Memorias do Instituto Oswaldo Cruz 
50:277-295. 

Cuénot, L. 1922. Sipunculiens, Echiuriens, Priapu- 
liens.—Faune de France, Paris 4:1—30. 

Cutler, E. B. 1965. Sipunculids of Madagascar. —Ex- 

trait des cahiers ORSTOM-—Océanographie 3(4): 

51-63. 

1969. New species of Sipuncula from the 


VOLUME 102, NUMBER 4 


western North Atlantic. — Proceedings of the Bi- 

ological Society of Washington 82:209-218. 

. 1973. Sipuncula of the western North Atlan- 

tic.— Bulletin of the American Museum of Nat- 

ural History 152(3):105—204. 

. 1977a. The bathyal and abyssal Sipuncula.— 

Galathea Report 14:135-156. 

1977b. Shallow water sipunculans collected 
by the Galathea Expedition (Sipuncula).— 

Steenstrupia 4:151-155. 

. 1981. A new species of Aspidosiphon (Sipun- 

cula) from the western Atlantic Ocean.—Pro- 

ceedings of the Biological Society of Washington 
94(2):445-449. 

—, & N. J. Cutler. 1979a. Madagascar and In- 
dian Ocean Sipuncula.— Bulletin du Muséum 
National d’Histoire Naturelle, Paris 4e(1):941- 
990. 

——., & 1979b. Campagnes de la Calypso 
au large des cétes Atlantiques Africaines (1956 
et 1959) et Sud-Ameéricaines (1960-1961). 23. 
Sipuncula.—Campagnes de la Calypso XI:103- 
109. 

—. & 1980a. Sipincula from southern 
Brazil.— Boletim do Institut Oceanografico, Sao 

Paulo 29(1):1-8. 

~& . 1980b. Deep water Sipuncula from 

the Gulf of Gascogne.—Journal of the Marine 

Biological Association of the United Kingdom 

60:449-459. 

——, & . 1980c. Sipuncula collected by the 

R/V “Vema.”— Journal of Zoology, London 190: 

193-209. 

,& . 1981. A reconsideration of Sipun- 

cula named by I. Ikeda & H. Sato.— Publications 

of the Seto Marine Biological Laboratory 26: 

51-93. 

——, & 1985a. A revision of the genera 
Sipunculus and Xenosiphon (Sipuncula).— Zoo- 
logical Journal of the Linnean Society 85:219- 
246. 

——, & 1985b. A revision of the genera 
Phascolion Theel and Onchnesoma Koren and 
Danielssen (Sipuncula). — Proceedings of the Bi- 
ological Society of Washington 98(4):809-850. 

——,& . 1987a. Deep-water Sipuncula from 
the eastern Atlantic Ocean.—Sarsia 72:71-89. 

~ & 1987b. A revision of the genus 

Golfingia (Sipuncula: Golfingiidae).— Proceed- 

ings of the Biological Society of Washington 

100(4):735-761. 

——.,.& . 1988. A revision of the genus The- 
miste (Sipuncula).— Proceedings of the Biolog- 
ical Society of Washington 101(4):741-766. 

: , & P. E. Gibbs. 1983. A reconsider- 

ation of the Gol/fingia subgenera Golfingiella and 


861 


Siphonoides (Sipuncula).—Proceedings of the 
Biological Society of Washington 96(4):669-694. 
; , & T. Nishikawa. 1984. The Sipun- 
cula of Japan: Their systematics and distribu- 
tion.— Publications of the Seto Marine Biolog- 
ical Laboratory 29(4/6):249-322. 

, & V. V. Murina. 1977. On the sipunculan 

genus Golfingia Lankester, 1885.—Zoological 

Journal of the Linnean Society 60:173-189. 

, & P.E. Gibbs. 1985. A phylogenetic analysis 

of higher taxa in the phylum Sipuncula.—Sys- 

tematic Zoology 34:162-173. 

Cutler, N. J., & E. B. Cutler. 1986. A revision of the 
genus Nephasoma (Sipuncula, Golfingiidae).— 
Proceedings of the Biological Society of Wash- 
ington 99(4):547-573. 

Danielssen, D. C., & J. Koren. 1880. New northern 
Gephyrea.—Annals and Magazine of Natural 
History (5)6:462-465. 

——.,& . 1881. Fra den norske Norhavsex- 
pedition (1876-1878)-Gephyreer.—Nytt Ma- 
gasin for Naturvidenskapene 26:44-66. 

DeRochebrune, A. T. 1881. Matériaux pour la faune 
de l’Archipel du Cap Vert.—Nouveux Archivs 
Muséum National d’Histoire Naturelle, Paris, 
2éme sér. 4:215-340. 

Diesing, K. M. 1851. Systema helminthum.— 

Braumiller, Vindobonae, 1-558 pp. 

. 1859. Revision der Rhyngodeen.—Sitzungs- 

berichte de mathematisch-naturwissenschaft- 

liche Klasse, Akademie der Wissenschaften in 

Wien 37:719-782. 

Ditadi, A. S. 1975. Aspidosiphon schnehageni (Si- 
puncula) inhabiting Tornatina shells.— The Ve- 
liger 18(2):200-—202. 

Edmonds, S. J. 1956. Australian Sipunculoidea II. 

The genera Phascolosoma, Dendrostomum, 

Golfingia, Aspidosiphon, and Cloeosiphon.— 

Australian Journal of Marine and Freshwater 

Research 7(2):281-315. 

. 1971. Some sipunculans and echiurans, chief- 

ly from Guam (Sipuncula and Echiura).—Mi- 

cronesica 7(1—2):137-151. 

1974. A new species of Sipuncula (Aspido- 
siphon exiguus) belonging to the interstitial fau- 
na of marine beaches collected by Mr. Botosa- 
neanu during the second Cuban-Romanian 
Biospeleological Expedition to Cuba 1973.—In- 
ternational Journal of Speleology 6:187-192. 

1980. A revision of the systematics of Aus- 
tralian sipunculans (Sipuncula).— Records of the 
South Australian Museum (Adelaide) 18(1):1- 
74. 

. 1987. Phyla Sipuncula and Echiura. Pp. 185- 

212 in D. M. Devaney and L. E. Eldridge, eds., 

Reef and shore fauna of Hawaii, sec. 2 and 3. 

Bishop Museum Press, Honolulu. 


862 

Fischer, J. 1914. Die Sipunculiden der Nord und 
Ostsee, unter Beruchsichtigung von Formen des 
nordatlantischen Gebietes. — Wissenschaftliche 
Meeresuntersuchungen Abteilung Kiel 16:85- 
L25% 

Fischer, W. 1892. Ubersicht der von Herr Dr. F. 


Stuhlmann auf Sanzibar und an der gegeniiber- 
liegenden Festlands-kuste gesammelten Gephy- 
reen.—Jahrbuch der Hamburgischen Wissen- 
schaftlichen Anstalten 9(2):79-89. 

1895. Die Gephyreen des Naturhistorischen 
Museums zu Hamburg.—Abhandlungen aus 
dem Gebiet der Naturwissenschaften, Hamburg 
13:1-24. 

1896. In Semon, Zoologische Forschungs- 
reisen in Australien und Malayischen Archi- 
pel. — Medizinisch-naturwissenschaftliche Ge- 
sellschaft zu Jena, Denkschriften 8:337-339. 

1913. Uber einige Sipunculiden des Natur- 
historischen Museums zu Hamburg. — Jahrbuch 
der Hamburgischen Wissenschaftlichen Anstal- 
ten 30(2):93-101. 

1914a. Gephyrea. Beitrage zur Kenntnis der 
Meeresfauna Westafrikas. — Beitrage zur Kennt- 
nis der Meeresfauna Westafrikas 1:57-84. 

1914b. Weitere Meitteilungen Uber die Ge- 
phyreen des Naturhistorischen (Zoologischen) 
Museums zu Hamburg.—Jahrbuch der Ham- 
burgischen Wissenschaftlichen Anstalten 31:1- 
28. 

1919. Gephyreen der Siid-westkiiste Aus- 
traliens.— Zoologischer Anzeiger 50:277-285. 

. 1920. Gephyreen der antarktischen und sub- 
antarktischen Meere.— Deutsche Sudpolar Ex- 
pedition 16, Zoologie 8:409-430. 

1921. Aspidosiphon pygmaeus n. sp., eine 
neue Gephyree aus Juan Fernandez.— Natural 
History of Juan Fernandez and Easter Islands, 
Uppsala 3:45-47. 

1922a. Gephyreen des Reichsmuseums zu 
Stockholm. — Arkiv fiir Zoologi 14(19):1-39. 

. 1922b. Gephyreen der deutschen Tiefsee-Ex- 
pedition, auf des Dampfer “Valdivia” 1898- 
1899.—Wissenschaftliche Ergebnisse der 
deutschen Tiefsee-Expedition, Valdivia 22(1): 
1-26. 

1922c. Westindische Gephyreen.—Zoolo- 
gischer Anzeiger 55:10-18. 

1923. Gephyreen des Golfes von Siam.— 
Videnskabelige Meddelelser Dansk fra Natur- 
historisk Forening 76:21-27. 

1925. Echuiriden, Sipunculiden, und Pria- 
puliden. Pp. 1-55 in G. Grimpe and E. Wagler, 
eds., Die Tierwelt der Nord-und Ostsee. Aka- 
demische Verlagsgesellschaft, Leipzig. 

1926. Sipunculiden und Echiuriden der 
Hamburger Sidsee-Expedition 1908-1909.— 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Mitteilungen aus dem Zoologischen Staatsinsti- 
tut und Zoologischen Museum in Hamburg 42: 
104-117. 

1931. Sipunculidae.— Mitteilungen aus dem 
Zoologischen Museum in Hamburg 44:130-140. 
Gerould, J. H. 1913. The sipunculids of the eastern 

coast of North America.—Proceedings of the 
United States National Museum 44:373-437. 

Gibbs, P.E. 1977. British sipunculans.— Synopses of 

the British Fauna No. 12. The Linnean Society 

of London 12:1-35. 

. 1978. Macrofauna of the intertidal sand flats 

on low wooded islands, northern Great Barrier 

Reef. — Philosophical Transactions of the Royal 

Society of London 284:8 1-97. 

—., & E. B. Cutler. 1987. A classification of the 
phylum Sipuncula.— Bulletin of the British Mu- 
seum Natural History (Zoology) 52(1):43-58. 

, E. B. Cutler, & N. J. Cutler. 1983. A review 

of the genus 7hysanocardia Fisher (Sipuncu- 

la).— Zoologica Scripta 12(4):295-304. 

Graveley, F.H. 1927. The littoral fauna of Krusadai 
Islands in the Gulf of Manaar. Gephyrea and 
Phoronis. — Bulletin of the Madras Government 
Museum 1:87-88. 

Grtibe, E. 1868. Beschreibungen einiger von Georg 
Ritter von Frauenfeld gesammelter Anneliden 
und Gephyreen des rothen Meeres.—Zoolo- 
gisch-Botanische Gesellschaft in Wien. Ver- 
handlungen der kaiserlich-khoniglichen Zoolo- 
gisch-Botanischen Gesellschaft in Wien 18:629- 
650. 

Haldar, B. P. 1976. Sipuncula from the Andaman 

and Nicobar Islands.—Records of the Zoolog- 

ical Survey of India 70:109. 

1978. Aspidosiphon (Paraspidosiphon) havy- 
elockensis, a new Sipuncula from the Anda- 
mans, India. — Bulletin of the Zoological Survey 
of India 1(1):37-41. 

Hammerstein, O. 1915. Geophyreen von Madagas- 
car gesammelt van W. Kaudern 1911-1912.— 
Arkiv fiir Zoologi 9(10):1-3. 

Hérubel, M. A. 1904. Liste des Sipunclides et des 

Echiurides rapportés par M. Ch. Gravier du 

Golfe de Tadjourah (mer Rouge).— Bulletin du 

Muséum National d’Histoire Naturelle, Paris 10: 

562-565. 

. 1924. Quelques echiurides et sipunclides des 

Cétes du Maroc et du Mauritane.— Bulletin du 

Société des Sciences Naturelles et Physiques du 

Maroc 4:108-112. 

Horst, R. 1899. Aspidosiphon cylindricus, n. sp.— 
Notes from the Leyden Museum (Rijks Museem 
van natuurlijke historie) 20:195-198. 

Ikeda, I. 1904. The Gephyrea of Japan.—Journal of 
the College of Science, Imperial University of 
Tokyo 20(4):1-87. 


VOLUME 102, NUMBER 4 


1924. Further notes on the Gephyrea of Ja- 
pan, with descriptions of some new species from 
the Marshall, Caroline, and Palau Islands.— 

Japanese Journal of Zoology 1:23-44. 

Johnson, P. 1964. Two new species of Aspidosiphon 
(Sipunculoidea). — Annals and Magazine of Nat- 
ural History 7(13):331-335. 

Keferstein, W. 1867. Untersuchungen uber einige 
amerikanische Sipunculiden.— Zeitschrift fiir 
Wissenschaftliche Zoologie 17:44—-55. 

Krohn, A. 1851. Uber die Larve des Sipunculus nu- 
dus.—Archiv fir Anatomie und Physiologie 
1851:368-379. 

Lanchester, W. F. 1905a. Gephyrea in the marine 

fauna of Zanzibar and British East Africa from 

the collections made by C. Crossland in 1901- 

1902.— Proceedings of the Zoological Society of 

London 1:28-35. 

. 1905b. On the Sipunculids and Echiurids col- 

lected during the ‘““Skeat”’ Expedition to the Ma- 

lay Peninsula.—Proceedings of the Zoological 

Society of London 1:35-41. 

Leroy, P. 1936. Les sipunculiens du Muséum d’His- 

toire Naturelle de Paris.— Bulletin du Muséum 

National d’Histoire Naturelle, Paris, Série 2 8: 

423-426. 

1942. Sipunculiens d’Indochine. Notes sta- 
tion maritime de Cauda. — Notes of the Institute 
Oceanography Nhatrang 40:1-51. 

LiGreci, F. 1980. Sul rinvenimento di un Sipuncu- 
lidae del genere Aspidosiphon, Diesing 1851, sui 
fondali batiali della Sicilia occidentale. —Riv- 
ista di Parassitologia 41(1):123-134. 

Longhurst, A. R. 1958. A ecological survey of the 
West African marine benthos. — Colonial Office 
of Fisheries Publications London 11:1-102. 

Lopez, R. J. S., J. I. Saiz-Salinas, & M. C. Diaz-Vilela. 
1984. Sobre la fauna de sipunculidos (Sipun- 
cula) del archipelago canario. Pp. 189-196 in V. 
Monteiro-Marques, ed., Actas do 4th Simpo- 
sio Iberico de Estudos do Benthos Marinho, As- 
sociacao des Estudantes da Faculdade Cien-Cias 
des Lisboa. 

Migotto, A. E., & A. S. F. Ditadi. 1988. The Aspi- 
dosiphonidae (Sipuncula) from the northern and 
northeastern coasts of Brazil.—Revista Brasi- 
leira de Biologia 48(2):245-263. 

Monro, C. A. 1931. Polychaeta, Oligochaeta, Echiu- 
roidea and Sipunculoidea.—Scientific Report of 
the Barrier Reef Expedition 4:1037. 

Miller, J. 1844. Uber einem neuen Wurm Sipunculus 
(Phascolosoma) scutatus.—Archiv fir Natur- 
geschichte 10:166-168. 

Murina, V. V. 1964a. Report on the sipunculid worms 
from the coast of South Chinese Sea.— Trudy 
Instituta Okeanologii Academii Nauk SSSR 69: 
254-270. 


863 


1964b. Sipunculid fauna of the Mediterra- 
nean Sea.—Trudy Sevastapol Biologicheskoi 
Statsil 17:51-76. 

1967a. New facts about the distribution of 
marine sipunculid worms in the Red Sea and 
the Gulf of Aden. Pp. 52-56 in Some results of 
Expedition III to the Red Sea Inbuma.— Express 
Information, Academy of Sciences Ukrainian 
Soviet Socialist Republic. 

. 1967b. On the sipunculid fauna of the littoral 

of Cuba. — Zoologicheskii Zhurnal 46:35-47. 

. 1967c. Report on the sipunculid worms from 

the sublittoral zone of Cuba and the Gulf of 

Mexico. —Zoologicheskii Zhurnal 54(9):1329- 

1339. 

. 1968. New species of Sipuncula from the Red 

Sea.— Zoologicheskii Zhurnal 47:1722-1725. 

1970. Contributions to the fauna of sipun- 
culid worms from the Aden Gulf.—Vestnik 
Zoologii 2:65-71. 

1971. On the species composition and ecol- 
ogy of sipunculids of the Red Sea. Pp. 76-88 in 
A. O. Kovalevski, ed., Shelf benthos of the Red 
Sea. Academii Nauk SSSR, Moscow. 

1972. Contribution of the sipunculid fauna 
of the Southern Hemisphere.—Zoologii Insti- 
tuta Akademii Nauk SSSR 11(19):294-314. 

. 1974. New subspecies of sipunculid worms. — 

Zoologicheskii Zhurnal 53:1713-1717. 

1978. On the sipunculid and priapulid fau- 
na of the southern Pacific.—Trudy Instituta 
Okeanologii Academii Nauk SSSR 113:120-131. 
. 1981. First report of the sea worms Sipuncula 
and Echiura from littoral Somali.— Vestnik 

Zoologi 1:11-18. 

Ocharan, F. J. 1980. Primeros datos sobre los sipun- 
ulides del litoral N.Y. N.W. de Espana (Santan- 
der, Asturias y Galicia).—Boletin de Ciencias 
de la Naturaleza I 26:111-119. 

Quatrefages, A. 1865. Gephyrea inermia.— Histoire 
naturelle des Annelés marins et d’eau douce, 
Paris 2:599-632. 

Rice, M. E. 1970. Asexual reproduction in a sipun- 

culan worm.—Science 167:1618-1620. 

1975. Survey of the Sipuncula of the coral 
and beach rock communities of the Caribbean 
Sea. Pp. 35-49 in M. E. Rice & M. Todorovic, 
eds., Proceedings of the International Sympo- 
sium of Sipuncula and Echiura, Naucno Delo 
Press, Belgrade. 

1976. Larval development and metamor- 
phosis in Sipuncula.—American Zoologist 16: 
563-571. 

— ., &I.G. Macintyre. 1972. A preliminary study 
of sipunculan burrows in rock thin sections. — 
Caribbean Journal of Science 12(1—2):41-44. 

——, & 1979. Distribution of Sipuncula 


864 


in the coral reef community, Carrie Bow Cay, 
Belize.—Smithsonian Contributions to Marine 
Sciences 12:311-320. 

—, & A.C. Stephen. 1970. The type specimens 
of Sipuncula and Echiura described by J. E. Gray 
and W. Baird in the collections of the British 
Museum of Natural History.—Bulletin of the 
British Museum of Natural History 20(2):49- 
TD 

Saiz Salinas, J. I. 1984. Gusanos sipunculidos (Si- 

puncula) de variaslocalidades de la costa de Por- 

tugal. Pp. 177-188 in V. Monteiro-Marques, ed., 

Actas do 4th Simposio Iberico de Estudos do 

Benthos Marinho, Associacao des Estudantes da 

Faculdade Cien-Cias des Lisboa. 

1986a. Los gusanos sipunculidos (Sipuncula) 
de los fondos litorales y circalitorales de las cos- 
tas de la peninsula Iberica, Islas Baleares, Cana- 
rias y mares adyacentes. — Monografias Instituto 

Espanol de Oceanografia, Madrid 1:1-84. 

1986b. The overlooked species of sipuncu- 
lans commensal of solitary corals (Sipuncula). — 

Bulletin du Muséum National d’Histoire Na- 

turelle, Paris 4e sér 8, sec. A 3:551-565. 

Sato, H. 1935. Sipunculoidea and Echiuroidea of the 

West Caroline Islands.— Tohoku Daigaku, Sen- 

dai, Japan, Science Reports series 4, 10:299- 

329. 

. 1939. Studies on the Echiuridea, sipunculids, 

and priapulids of Japan.—Science Reports of 

Tohoku Imperial University 14(4):339-459. 

Schmidt, O. 1865. Uber den Bau un die systema- 
tische Stellung von Aspidosiphon muelleri Dies- 
ing (Lesinia farcimen Schmidt).— Mitteilungen 
des Naturwissenschaftlichen Vereines fur 
Steiermark 3:56-66. 

Selenka, E. 1885. Report on the Gephyrea collected 

by H.M.S. Challenger during 1873—76.— Report 

of scientific results of the Voyage of Challenger 

Zoology 13(36):1-25. 

, J. G. de Man, & C. Bulow. 1883. Die Sipun- 

culiden, eine systematische Monographie. — 

Semper Reisen in Archipel Phillippinen 2 4:1- 

13%. 

Shipley, A.E. 1898. Report on the Gephyrean worms 
collected by Mr. J. Stanley Gardiner at Rotuma 
and Funafuti.—Journal of Zoology 1898:468- 
473. 

1899a. On Gephyrean worms from Christ- 
mas Island.— Proceedings of the Zoological So- 

ciety of London 22:54—57. 

1899b. A report on the Sipunculoidea col- 
lected by Dr. Willey at the Loyalty Islands and 
in New Britain. Pp. 151-160 in A. Willey, ed., 

Zoological results based on material from New 

Britain, New Guinea, Loyalty Islands and else- 

where collected 1985-1897 by A. Willey. Uni- 

versity Press, Cambridge. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


1902. Sipunculoidea, with an account of a 
new genus Lithacrosiphon. Pp. 131-140 in J. S. 
Gardiner, ed., Fauna and geography of the Mal- 
dive and Laccadive Archipelagoes. University 
Press, Cambridge. 

1903. Report on the Gephyrea collected by 

Prof. Herdman at Ceylon in 1902.—Ceylon Pearl 
Oyster Fisheries Supplementary Reports 3:169- 
176. 
Sluiter, C. P. 1881. Beitrage zu der Kenntnis der 
Gephyreen aus dem Malayischen Archipel.— 
Natuurkundig Tijdschrift voor Nederlandisch- 
Indie 41:84-108. 

1884. Beitrage der Gephyreen aus dem Ma- 
layischen Archipel.—Natuurkunding Tijdsch- 
rift voor Nederlandisch-Indie 43:26-88. 

1886. Beitrage zu der Kenntnis der Gephy- 
reen aus dem Malayischen Archipel.—Natu- 
urkundig Tijdschrift voor Nederlandisch-Indie 
45:472-517. 

1891. Die Evertebraten aus der Sammlung 
des KOniglichen naturwissenschaftlichen Ver- 
eins in Nederlandisch-Indien in Batavia. — Na- 
tuurkunding Tijdschrift voor Nederlandisch-In- 
die 50:102-125. 


1898. Gephyreen von Sud Africa.—Zoolo- 
gische Jahrbucher Abteilung fiir Systematik 11: 
442-450. 


. 1900. Gephyriens (Sipunculides et Echiurides) 

provenant des campagnes de |’Hirondelle et de 

la Princesse Alice, 1886—-1897.—Résultats des 

Campagnes Scientifique accomplies sur son yacht 

sur Albert I, Prince Souverain de Monaco 15: 

1-29. 

1902. Die Sipunculiden und Echiuriden der 
““Siboga’”’ Expedition, nebst Zusammenstellung 
der Uberdies aus den indischen Archipel be- 
kannten Arten.—Siboga-Expeditie, Monogra- 
phie 25:1-53. 

1912. Géphyriens (Sipunculides et Echiu- 
rides) provenant des campagnes de la Princesse 

Alice, 1898-1910.—Résultats des Campagnes 
Scientifique accomplies sur son yacht sur Albert 
I, Prince Souverain de Monaco 36:1-—36. 

Southern, R. 1913. Gephyrea of the coasts of Ire- 
land.—Scientific Investigations of the Fisheries 
Branch, Department of Agriculture for Ireland 
3:1-46. 

Stephen, A. C. 1934. The Echiuridae, Sipunculidae, 
and Priapulidae of Scottish and adjacent 
waters. — Proceedings of the Royal Society of 
Edinburgh 22:159-186. 

1941. The Echiuridae, Sipunculidae, and 
Priapulidae collected by the ships of the Dis- 
covery Committee during the years 1926- 
1937.— Discovery Reports 21:235-260. 

. 1942. The South African intertidal zone and 


VOLUME 102, NUMBER 4 


its relation to ocean currents. Notes on the in- 

tertidal sipunculids of Cape Province and Na- 

tal.—Annals of the Natal Museum 10(2):245- 

256. 

1958. The sipunculids of Haifa Bay and 
neighborhood. — Bulletin of the Research Coun- 
cil of Israel 7b(3-4):129-136. 

. 1960a. Echiuroidea and Sipunculoidea from 

Senegal, West Africa.—Bulletin de I’Institut 

Francais d’Afrique Noire 22a(2):512-520. 

1960b. British echiurids (Echiuroidea), si- 
punculids (Sipunculoidea), and priapulids (Pri- 
apuloidea). Synopses of the British Fauna No. 

12. The Linnean Society London, 27 pp. 

1964. A revision of the classification of the 
phylum Sipuncula.—Annals and Magazine of 
Natural History 7(13):457-462. 

,& S.J. Edmonds. 1972. The phyla Sipuncula 

and Echiura. Trustees British Mus. (Nat. Hist.), 

London, 528 pp. 

—., & J.D. Robertson. 1952. A preliminary re- 
port on the Echiuridae and Sipunculidae of Zan- 
zibar.—Proceedings of the Royal Society of 
Edinburgh 64(4):426-444. 

Steuer, A. 1936. Sipunculidea, Phoronidea, Brachi- 

opoda, Enteropneusta und Acrania von Alex- 

andrien in Agypten.—Note dell’Instituto Bio- 

logia marina di Rovigno 23:1-18. 

1939. The fishery grounds near Alexandria. 
XVIII. Sipunculoidea, Phoronidea, Brachiop- 
oda, Enteropneusta and Acrania.— Fouad I. In- 
stitute of Hydrobiology and Fisheries Notes and 
Memoirs 30:1-3. 
ten Broeke, A. 1925. Westindische Sipunculiden und 

Echiuriden. — Bijdragen tot de Dierkunde 24:8 1- 
96. 

Théel, H. 1875. Etudes sur les géphyriens inermes 
des mers de la Scandinavie, du Spitzberg et du 

Groenland.— Bihang till Kungliga Svenska Ve- 

tenskaps-Akademiens Handlingar 3(6):1-30. 

1905. Northern and Arctic invertebrates in 
the collection of the Swedish State Museum I. 
Sipunculids.—Kungliga Svenska Vetenskaps- 
Akademiens Handlingar 39(1):1-130. 

Vaillant, L. 1871. Note sur quelques Gephyriens de 

Saint-Vincent du Cap Vert. Pp. 268-276 in Les 

fonds de la mer. Etude internationale sur les 

particularités nouvelles des régions sous-ma- 
rines (Vol. 1), L. De Folin & L. Périer (dirs.), 

Paris, Savy. 

. 1875. (Planche IV, Géphyriens). Jn Les fonds 

de la mer. Etude internationale sur les particu- 

larités nouvelles des regions sous-marines (Vol. 

2), L. De Folin & L. Périer (dirs.), Paris, Savy, 

pl. IV. 

Voss-Foucart, M. F., S. Barzin, C. Jeuniaux, & J. C. 
Bussers. 1977. Etude comparée de la com- 


865 


position chimique des regions souples et durcies 

de la cuticule de quatre espéces de sipuncu- 

liens. — Cahiers de Biologie Marine 18:135-145. 
Wesenberg-Lund, E. 1937. Gephyrea.— Bulletin Mu- 
sée royal d’Histoire naturelle de Belgique 13(36): 
1-23. 

1954. Priapuloidea, Sipunculoidea and 
Echiuroidea.— Bulletin Institut royal des Sci- 
ences naturelles de Belgique 30(16):1-18. 

. 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 


870 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


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 


ceedings of the United States National Museum 
14:277-302. 

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 

Coast with a few new records from the shore. — 

Journal of the Linnean Society of London 44: 

463-560. 

. 1971. Polychaeta. Pp. 384-390 in E. M. van 

Zinderen Baker, Sr., J. M. Winterbottom, & 

R. A. Dyer, eds., Marion and Prince Edward 

Islands. A. A. Balkema, Capetown, 427 pp. 

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. 


871 


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. 


VOLUME 102, NUMBER 4 


(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 


VOLUME 102, NUMBER 4 


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. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


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 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


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. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


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. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


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 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


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. 


VOLUME 102, NUMBER 4 


. 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 


\ 


AN 


\ 


\) 


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. 


VOLUME 102, NUMBER 4 897 


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). 


898 


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- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


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- 


VOLUME 102, NUMBER 4 


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, 


899 


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 


900 


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. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


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). 


VOLUME 102, NUMBER 4 


903 


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. 


VOLUME 102, NUMBER 4 


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 


905 


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 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


906 


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 


— 


907 


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. 


910 


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. 


914 


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. 


917 


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- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


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. 


919 


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. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


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 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


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). 


934 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


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- 


VOLUME 102, NUMBER 4 


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 


936 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


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- 


VOLUME 102, NUMBER 4 937 


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. 


938 


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 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


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 


VOLUME 102, NUMBER 4 


/ 
(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. 


VOLUME 102, NUMBER 4 


941 


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 


942 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


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; 


943 


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 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


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- 


VOLUME 102, NUMBER 4 


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. 


Literature Cited 


Bou, C. 1970. Observations sur les Ingolfiellides 
(Crustacés Amphipodes) de Grece.—Biologia 
Gallo-Hellenica 3(1):57-70, pl. 1. 

Bousfield, E. L. 1979. A revised classification and 
phylogeny of amphipod crustaceans.—Trans- 
actions of the Royal Society of Canada (4)16: 
343-390. 

Bowman, T. E. 1984. Stalking the wild crustacean: 
the significance of sessile and stalked eyes in 
phylogeny. — Journal of Crustacean Biology 4(1): 
7-11. 


945 


—, & L. G. Abele. 1982. Classification of the 
Recent Crustacea. Pp. 1-27 in L. G. Abele, ed., 
The biology of Crustacea, vol. 1, Systematics, 
the fossil record, and biogeography. Academic 
Press, New York. 

—, & T.M. Iliffe. 1985. Mictocaris halope, a new 

unusual peracaridan crustacean from marine 

caves on Bermuda.—Journal of Crustacean Bi- 
ology 5(1):58-73. 

,& . 1986. Halosbaena fortunata, anew 

thermosbaenacean crustacean from the Jamios 

del Agua marine lava cave, Lanzorote, Canary 

Islands. —Stygologia 2(1/2):84-89. 

,& 1988. Tulumella unidens, a new 

genus and species of thermosbaenacean crus- 

tacean from the Yucatan Peninsula, Mexico. — 

Proceedings of the Biological Society of Wash- 

ington 101(1):221-226. 

Coineau, N. 1963. Presence du sous-ordre des In- 

golfiellidea Reibisch (Crustacea Amphipoda) 

dans les eaux souterraines continentales de 

France.—Comptes Rendus Hebdomadires des 

Séances de l’Academie des Sciences 256:4729- 

4731. 

. 1968. Contribution a l’étude de la faune in- 

terstitielle Isopodes et Amphipodes.—Mém- 

oires du Muséum National d’Histoire Naturelle 

(n.s.) (A, Zoologie) 55(3):145-216. 

Dahl, E. 1977. The amphipod functional model and 
its bearing upon systematics and phylogeny. — 
Zoologica Scripta 6:221-228. 

Dojiri, M., & J. Sieg. 1987. Ingolfiella fuscina, new 
species (Crustacea: Amphipoda) from the Gulf 
of Mexico and the Atlantic coast of North 
America, and partial redescription of J. atlantisi 
Mills, 1967.— Proceedings of the Biological So- 
ciety of Washington 100(3):494—50S. 

Gordon, I. 1957. On Spelaeogriphus, a new caver- 
nicolous crustacean from South Africa.—Bul- 
letin of the British Museum (Natural History) 
Zoology 5(2):31-47. 

Hansen, H.J. 1903. The Ingolfiellidae, fam. n., a new 
type of Amphipoda.—Journal of the Linnean 
Society of London (Zoology) 29:117-133. 

Karaman, S. L. 1957. Eine neue Jngolfiella aus Ju- 

goslawien, Ingolfiella petkovskii n. sp.—Folia 

Balcanica, Skopje 1(7):35-38. 

. 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 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


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- 


948 


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- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


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 


VOLUME 102, NUMBER 4 


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). 


949 


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 


950 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


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. 


VOLUME 102, NUMBER 4 951 


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. 


952 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


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 


VOLUME 102, NUMBER 4 


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- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


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.) 


956 


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- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


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- 


VOLUME 102, NUMBER 4 


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. 


Literature Cited 


Akatsuka, K., & T. Komai. 1922. Pseudocrangonyx, 
a new genus of subterranean amphipods from 
Japan.—Annotationes Zoologicae Japonenses 
10:119-126. 

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 

gammarid Amphipoda.—Crustaceana, supple- 

ment 6:5-16. 

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. — 

Oklahoma Underground 6(1):15-—17. 

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- 

390. 

. 1982. Amphipoda. Pp. 254-293 inS. S. Park, 

ed., Synopsis and classification of living organ- 

isms. McGraw-Hill Book Company, New York. 

. 1983. An updated phyletic classification and 

palaeohistory of the Amphipoda. Pp. 257-277 

in F. R. Schram, ed., Crustacean phylogeny. A. 

A. Balkema, Rotterdam. 


VOLUME 102, NUMBER 4 


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. 

Pp. 85-106 in R. H. Gore and K. L. Heck, eds., 

Crustacean biogeography. A. A. Balkema, Rot- 

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. 


959 


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 

Japan.—Annotationes Zoologicae Japonenses 

13(1):21-23. 

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. 

Wilson. J. H. (ed.). 1984. Rare and endangered species 
of Missouri. Missouri Department of Conser- 
vation Publication, 171 pp. 


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- 


961 


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- 


VOLUME 102, NUMBER 4 


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 


964 


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 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


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 


974 


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 


VOLUME 102, NUMBER 4 


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- 


975 


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. 


976 


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, 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


VOLUME 102, NUMBER 4 


979 


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. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


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VOLUME 102, NUMBER 4 


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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


VOLUME 102, NUMBER 4 


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. 


— 


983 


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. 


984 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


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- 


VOLUME 102, NUMBER 4 


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 


VOLUME 102, NUMBER 4 


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 


990 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


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 


VOLUME 102, NUMBER 4 


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, 


992 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


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 


VOLUME 102, NUMBER 4 


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 


994 


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 


VOLUME 102, NUMBER 4 


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 


995 


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- 


996 


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 


VOLUME 102, NUMBER 4 


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- 


998 


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 


999 


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. 


1000 


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. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


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- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


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). 


VOLUME 102, NUMBER 4 


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 


1003 


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- 


1004 


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. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


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- 


VOLUME 102, NUMBER 4 


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 


1006 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


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 


1007 


VOLUME 102, NUMBER 4 


nn ne LEE SEass ssn L TST 


Ic 81 *f 8 IG 9 WNJIUINWAS *H 

90°61 G I I *9 v G AIQNAOASIU “HY 

Seen ——eeE ee eee 
x CC IZ 0c 61 81 LI so1oads 


(quay Jomo] + quit] soddn) [R10], 


eee ee 8. oo 


Nee ee eee 


coe! 49 86. ov. 19'P C x61 LI CV's «Ll KG WUNJOUIIIWAS *H 
py cl (6 I *f 9 v vv L I I + *V 9 v 00°9 (4 «CI C AIGNdOA31U “FY 
cc ee ee 

x 91 cI vI tl Gi x IT Ol 6 8 L 9 S 4 x 6 8 L 9 S sa1seds 

(‘pni + “Aop) ung AieyUSWIIpNyY podojeasq 
quit] IoMOT 

Nee ee ee eee 
91'S «8 8¢ C «8 8C C 3 *8€ WNJIUIIIVAAS "HT 
c9'S Ol *9 Ol *9 *9 I ADQnAOASIU * FT 
rr ee ee 

x 9 iS v ¢ v t I sa1sedg 

(‘pni + “‘Aop) wing AieyuSWIpny padojsasqd 
quit] odd 

*SYSLIONSe 


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 


1008 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


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. 


1010 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


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 


VOLUME 102, NUMBER 4 


1011 


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 


1012 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


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- 


VOLUME 102, NUMBER 4 


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 


1013 


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.) 


VOLUME 102, NUMBER 4 


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. 


Literature Cited 


Allen, G. R. 1976. Descriptions of three new fishes 
from Western Australia.—Journal of the Royal 
Society of Western Australia 59(1):24—30. 

—, & J. T. Moyer. 1980. Ellerkeldia wilsoni, a 
new species of serranid fish from southwestern 
Australia. — Japanese Journal of Ichthyology 26: 
329-333. 

Anderson, W. D., Jr., & P. C. Heemstra. 1980. Two 
new species of western Atlantic Anthias (Pisces: 
Serranidae), redescription of A. asperilinguis and 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


review of Holanthias martinicensis.—Copeia 
1980:72-87. 

Bauchot, M. L., M. Desoutter, & J. E. Randall. 1984. 
Catalogue critique des types de poissons du Mu- 
séum National d’Histoire Naturelle (famille des 
Serranidae).— Bulletin Muséum National 
d’Histoire Naturelle, Paris, 4th Sér., 6, section 
A, No. 3, supplément:3-82. 

Bleeker, P. 1873. Surles espéces indo-archipelagiques 
d’Odontanthias et de Pseudopriacanthus. — 
Nederlandsch Tijdschrift voor de Dierkinde 4: 
235-240. 

Boulenger, G. A. 1895. Catalogue of the perciform 
fishes in the British Museum, volume 1. 2nd ed. 
London, xix + 394 pp. 

Buen, F.de. 1959. Lampreas, tiburones, rayas y peces 
en la Estacion de Biologia Marina de Montemar, 
Chile.— Revista de Biologia Marina 9:3—200. 

Castelnau, F. de. 1875. Researches on the fishes of 
Australia.—Intercolonial Exhibition Essays, 
1875-6, No. 2:3-52. 

Cossman, M. 1889. Catalogue illustré des coquilles 
fossiles de l’Ecoéne des environs de Paris. —An- 
nales de la Société Royale Malacologique de Bel- 
gique 24:3-381. 

Cuvier, G. 1828. In G. Cuvier & A. Valenciennes, 
Histoire naturelle des poissons, tome 2. Paris, 
xxiv + 490 pp. 

Fourmanoir, P. 1982. Trois nouvelles espéces de Ser- 
ranidae des Philippines et de la Mer du Corail 
Plectranthias maculatus, Plectranthias barroi, 
Chelidoperca lecromi.—Cybium 6(4):57-64. 

—, & J. Rivaton. 1980. Plectranthias randallin. 
sp., un nouveau serranidé (anthiiné) du sud-ouest 
Pacifique.— Revue Frangaise d’Aquariologie 7: 
27-28. 

Gill, T. 1862. Remarks on the relations of the genera 
and other groups of Cuban fishes. — Proceedings 
of the Academy of Natural Sciences of Phila- 
delphia 14:235-242. 

Gosline, W. A. 1966. The limits of the fish family 

Serranidae, with notes on other lower per- 

coids.— Proceedings of the California Academy 

of Sciences, 4th Ser., 33:91-111. 

1968. The suborders of perciform fishes. — 
Proceedings of the United States National Mu- 
seum 124(No. 3647):1-78. 

1971. Functional morphology and classifi- 
cation of teleostean fishes. Honolulu, The Uni- 
versity Press of Hawaii, ix + 208 pp. 

Gunther, A. 1859. Catalogue of the acanthopterygian 
fishes in the collection of the British Museum, 
volume |. London, xxxii + 524 pp. 

Hector, J. 1875. Notes on New Zealand ichthyolo- 
gy.— Transactions and Proceedings of the New 
Zealand Institute 7(for 1874):239-250. 

Heemstra, P. C., & W. D. Anderson, Jr. 1983. Anew 
species of the serranid fish genus Plectranthias 
(Pisces: Perciformes) from the southeastern Pa- 


VOLUME 102, NUMBER 4 


cific Ocean, with comments on the genus Eller- 
keldia. —Proceedings of the Biological Society 
of Washington 96:632-637. 

Hughes, D. R. 1981. Development and organization 
of the posterior field of ctenoid scales in the 
Platycephalidae.—Copeia 1981:596-606. 

International Commission on Zoological Nomencla- 
ture. 1985. International code of zoological 
nomenclature. 3rd ed. University of California 
Press, Berkeley, California. xx + 338 pp. 

Johnson, G. D. 1975. The procurrent spur: An un- 

described perciform caudal character and its 

phylogenetic implications.— Occasional Papers 

of the California Academy of Sciences 121:1- 

D3: 

1983. Niphon spinosus: A primitive epi- 
nepheline serranid, with comments on the mon- 
ophyly and intrarelationships of the Serrani- 
dae.—Copeia 1983:777-787. 

. 1984. Percoidei: Development and relation- 

ships. Pp. 464-498 in H. G. Moser et al., eds., 

Ontogeny and systematics of fishes. American 

Society of Ichthyologists and Herpetologists, 

Special Publication No. 1. 

. 1988. Niphon spinosus, a primitive epinephe- 
line serranid: Corroborative evidence from the 
larvae.— Japanese Journal of Ichthyology 35:7— 
18. 

Jordan, D.S. 1891. In D. S. Jordan & C. H. Eigen- 
mann, A review of the genera and species of 
Serranidae found in the waters of America and 
Europe.—Bulletin of the United States Fish 
Commission 8(for 1888):329-441. 

Katayama, M. 1954. A new serranid fish found in 

Japan. Japanese Journal of Ichthyology 3:56- 

61. 

. 1959. Studies on the serranid fishes of Japan 

(1).— Bulletin of the Faculty of Education, Ya- 

maguchi University 8(Pt. 2):103-180. 

1960. Fauna japonica Serranidae (Pisces). 
Biogeographical Society of Japan. Tokyo Elec- 
trical Engineering College Press, Tokyo. viii + 
189 pp. 

—_., & H. Masuda. 1980. Two new anthiine fishes 
from Sagami Bay, Japan.—Japanese Journal of 
Ichthyology 27:185-190. 

Leis, J. M., & D.S. Rennis. 1983. The larvae of Indo- 
Pacific coral reef fishes. New South Wales Uni- 
versity Press, Sydney, Australia. 269 pp. 

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 1985:802-832. 

Ogilby, J. D. 1908. New genera and species of fish- 
es.—Proceedings of the Royal Society of 
Queensland 21:1-26. 


1017 


Olmi, C. B. 1986. Morphology of the larvae of Amer- 
ican Anthiinae (Pisces: Serranidae) with com- 
ments on relationships within the subfamily. 
Unpublished M.S. Thesis, College of Charles- 
ton, South Carolina. viii + 119 pp. 

Paulin, C. D., & C. D. Roberts. 1987. A new species 
of the anthiine fish genus Plectranthias (Perco- 
morpha; Serranidae) from the Kermadec Is- 
lands offnorthern New Zealand. — National Mu- 
seum of New Zealand Records 3(2):13-16. 

Raj, U., & J. Seeto. 1983. A new species of the an- 
thiine fish genus Plectranthias (Serranidae) from 
the Fiji Islands.—Japanese Journal of Ichthy- 
ology 30:15-17. 

Randall, J. E. 1980. Revision of the fish genus Plec- 
tranthias (Serranidae: Anthiinae) with descrip- 
tions of 13 new species.— Micronesica 16:101- 
187. 

—,& A.CeaEgania. 1984. Native names of East- 
er Island fishes, with comments on the origin of 
the Rapanui people.—Occasional Papers of 
Bernice Pauahi Bishop Museum 25(12):1—16. 

Rendahl, H. 1921. The fishes of the Juan Fernandez 
Islands. Pp. 49-58 in C. Skottsberg, ed., The 
natural history of Juan Fernandez and Easter 
Island, volume 3, Zoology, Part 1. Almqvist & 
Wiksells Boktryckeri-A. -B., Uppsala. 

Scott, E. O. G. 1979. Observations on some Tas- 
manian fishes: Part XXV.—Papers and Pro- 
ceedings of the Royal Society of Tasmania 113: 
99-148. 

Sepulveda Vidal, J. I.,& G. Pequeno R. 1985. Fauna 
ictica del archipiélago de Juan Fernandez. Pp. 
81-91 in P. Arana, ed., Investigaciones marinas 
en el Archipiélago de Juan Fernandez. Valpa- 
raiso, Universidad Catolica de Valparaiso. 

Valenciennes, A. 1833. In G. Cuvier & A. Valen- 
ciennes, Histoire naturelle des poissons, tome 
9. Paris. xxxii + 512 pp. 

Whitley, G. P. 1927. Studies in ichthyology. No. 1.— 

Records of the Australian Museum 15:289-304. 

1929. Studies in ichthyology. No. 3.—Rec- 

ords of the Australian Museum 17:101-143. 

Yanez-Arancibia, L.A. 1975. Zoogeografia de la fau- 
na ictidlogica de la Isla de Pascua (Easter Is- 
land).—Anales del Centro de Ciencias del Mar 
y Limnologia de la Universidad Nacional Au- 
tonoma de México 2:29-51. 


(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, 


VOLUME 102, NUMBER 4 


1019 


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). 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


1020 


‘azIjog ‘ABD MOg SLLUeD “TS WU 6°¢T ‘oTeW ‘CEQTOE INNSN ‘edAjojoy ‘sotoods mou ‘njnNnd visDWMalquiayjUuvIp 


‘| ‘Sy 


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 


1021 


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 


1022 


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- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


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 


VOLUME 102, NUMBER 4 


1023 


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 


1024 


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 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


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 


1025 


(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- 


1026 


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 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


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- 


1027 


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 


1028 


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 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


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. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


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 


1035 


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 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


& 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 


1043 


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, 


1044 


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 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


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). 


1046 


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 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


{ 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 


VOLUME 102, NUMBER 4 


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). 


VOLUME 102, NUMBER 4 


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. 


Literature Cited 


Abbott, I. A. 1970. Yamadaella, a new genus in the 
Nemaliales (Rhodophyta).—Phycologia 9(2): 
115-123. 

—., & M. Yoshizaki. 1981. A second species of 
Dotyophycus (Nemaliales, Rhodophyta) that 
emphasizes the distinctness of a diffuse goni- 
moblast.— Phycologia 20(3):222-227. 

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 


1065 


(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. 

Littler, M. M. 1972. The crustose Corallinaceae. — 

Oceanography and Marine Biology Annual Re- 

view 10:311-347. 

1976. Calcification and its role among the 
macroalgae.— Micronesica 12:27-41. 

,&D.S. Littler. 1984. Models of tropical reef 

biogenis: The contribution of algae. Pp. 323- 

364 in F. E. Round and D. J. Chapman, eds., 

Progress in phycological research, vol. 3. Bio- 

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. 

Pueschel, C. M. 1979. Ultrastructure of tetrasporo- 

genesis in Palmaria palmata (Rhodophyta).— 

Journal of Phycology 15:409-424. 

1980. A reappraisal of the cytochemical 
properties of rhodophycean pit plugs.— Phyco- 
logia 19:210-217. 

1989. An expanded survey of the ultrastruc- 


1066 


ture of red algal pit plugs. — Journal of Phycology 
25 (in press). 

—.,&K.M.Cole. 1982. Rhodophycean pit plugs: 
An ultrastructural survey with taxonomic im- 
plications.—American Journal of Botany 69: 
703-720. 

Ramus, J. 1969. The developmental sequence of the 
marine red alga Pseudogloiophloea in culture. — 
University of California Publications in Botany 
52:1-28. 

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. 

Sheath, R. G. 1984. The biology of freshwater red 
algae. Pp. 89-157 in F. E. Round and D. J. 
Chapman, eds., Progress in phycological re- 
search, vol. 3. Biopress Ltd., Bristol. 

Silva, P. C., & H. W. Johansen. 1986. A reappraisal 
of the Order Corallinales (Rhodophyceae).— 
British Phycological Journal 21:245-254. 

Svedelius, N. 1939. Uber den Bau und die Entwick- 

lung der spermatangiengruben bei der Flori- 

deengattung Galaxaura. —Botaniska Notiser 

1939:59 1-606. 

1943. Zytologisch-Entwicklungsgeschicht- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


liche Studien tiber Galaxaura eine diplobion- 
tische Nemalionales-Gattung. — Nova Acta Re- 
giae Societatis Scientiarum Upsaliensis, IV, 
13(4):1-154. 

Swale, E. M. F. 1962. The development and growth 

of Thorea ramosissima Bory.— Annales of Bot- 

any 26:105-117. 

. 1963. Notes on the morphology and anatomy 

of Thorea ramosissima Bory.—Journal of the 

Linnean Society (Botany) 58:429-435. 

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. 

Wittmann, W. 1965. Aceto-iron-haematoxylin- 
chloral hydrate for chromosome staining. — Stain 
Technology 40:161-164. 

Yoshizaki, M. 1986. The morphology and reproduc- 
tion of Thorea okadai (Rhodophyta).—Phyco- 
logia 25(4):476—-481. 


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. 

Review. —One of the Society’s aims is to give its members an opportunity for prompt pub- 
lication of their shorter contributions. Manuscripts are reviewed in order of receipt by a board 
of Associate Editors and appropriate referees. 

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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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