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THE BIOLOGICAL SOCIETY OF WASHINGTON 
1999-2000 
Officers 


President: Richard P. Vari Secretary: Carole C. Baldwin 
President-elect: Brian F. Kensley Treasurer: T. Chad Walter 


Elected Council 


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PROCEEDINGS 


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EDITOR 

BIOLOGICAL SOCIETY OF WASHINGTON 
NATIONAL MUSEUM OF NATURAL HISTORY 
WASHINGTON, D.C. 20560, U.S.A. 


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This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper). 


This number is dedicated to 
Dr. Austin B. Williams 
Systematic Zoologist 
National Systematics Laboratory 
National Marine Fisheries Service 
National Museum of Natural History 
Smithsonian Institution 


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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


113(1):1—12. 2000. 


Austin Beatty Williams (17 October 1919-27 October 1999). 
Biographical summary 


Rafael Lemaitre and Bruce B. Collette 


(RL) Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian 
Institution, Washington, D.C. 20560-0163, U.S.A.; 

(BBC) Bruce B. Collette, National Marine Fisheries Service Systematics Laboratory, National 

Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560-0153, U.S.A. 


Abstract.—The professional career and aspects of the life of Austin Beatty 
Williams (17 October 1919-27 October 1999), Systematic Zoologist for the 
Systematics Laboratory of the U.S. National Marine Fisheries, are summarized. 
Included is a bibliography with the 118 papers published by Williams, and a 
list of all the new names he proposed along with the holotype repository and 
catalogue number of species and subspecies. 


Austin B. Williams (Fig. 1) was System- 
atic Zoologist at the Systematics Laborato- 
ry, National Marine Fisheries Service 
(NMES), based at the National Museum of 
Natural History, Smithsonian Institution, 
Washington, D.C. After a valiant fight with 
cancer, Austin passed away at his home in 
Falls Church, Virginia. Shortly after his 
death, the Council of the Biological Society 
of Washington unanimously voted to dedi- 
cate this issue, the first of the new millen- 
ium, to his memory. This dedication is most 
fitting as Austin generously served the So- 
ciety 1n many capacities during the last 
quarter of the 20th century, and gave luster 
to the Proceedings by using it to publish 
many of his important papers. He served 
the Society as a Editor of the Proceedings 
(1974-1977), Vice-president (1983-1986), 
President (1986-1988), Past-President 
(1989-1999), Custodian of Publications 
(1989-1995), and contributed significantly 
to its financial soundness as a member of 
the Finance Committee (1995-1999). He 
was editor of Bulletin No. 3: ““Symposium 
on the Composition and Evolution of Crus- 
taceans in the Cold and Temperate Waters 
of the World Oceans”’ (1979), based on the 
results of a U.S.-U.S.S.R. Cooperative Pro- 
gram. He also provided the summary chap- 


ter for Bulletin No. 6: ‘““The hydrothermal 
vents of the eastern Pacific: An overview” 
(Williams 1985b). 

Austin had a distinguished career span- 
ning five decades during which he pub- 
lished 118 papers (see bibliography). Born 
in Plattsburg, Missouri in 1919, he was the 
first child of Oliver Perry Williams and 
Lucy Sell; his siblings are brothers Hillis 
and Oliver. He married Jean McNicol with 
whom he had their only child, David (mar- 
ried to Anita Kyle, with two children, Lau- 
ren and Kyle). His family had only modest 
means so he had to work to support his ed- 
ucation, first at McPherson College (A.B. 
1943), and then at the University of Kansas 
(Ph.D. 1951), where he studied Ozark cray- 
fishes. His studies on these crayfishes re- 
main among the key references to identify 
these decapods in the region. From 1951 to 
1955 he was with the University of North 
Carolina Institute of Fisheries Research, 
studying the life history and ecology of 
penaeid shrimps. He then worked at the 
University of Illinois from 1956 to 1963, 
after which he returned to the North Caro- 
lina Institute of Fisheries Research to con- 
tinue his studies on marine and estuarine 
decapods. In the mid 1960s, Donald F 
Squires, Chairman of the Department of In- 


i) 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 1. 
at Heart Tail Ranch, Butte County, South Dakota, one of his favorite fossil collecting sites, 31 July 1997, in 
back is son David, in front left to right are David’s wife Anita with grandchildren Kyle and Lauren, and Norma 
Samuels (Norma Samuels); recording observations in his notebook on fossil Cretaceous decapods collected from 
the Pierre Shale at Heart Tail Ranch, Butte County, South Dakota, 6 August 1997 (Gale A. Bishop); at Mount 
Rushmore, South Dakota, August 1997 (Norma Samuels). 


vertebrate Zoology and then Deputy Direc- 
tor of the National Museum of Natural His- 
tory, Smithsonian Institution, considered 
him for a job; however, his interview with 
Secretary Dillon Ripley did not go well 
(most probably because of Austin’s non-as- 
suming personality), and he turned him 


Austin B. Williams. Clockwise from upper left: from church directory, 1995; with family and friends 


down. This was a loss for the Museum but 
fortunately he was hired by the NMFS’ 
Systematics Laboratory in 1971. 

In addition to his crustacean work, Aus- 
tin served NMFS and the Museum in a 
number of different ways. For example, he 
represented the Allied Agencies (NMFS, 


VOLUME 113, NUMBER 1 


Agriculture, and what was then the Fish and 
Wildlife Service) on the Senate of Scientists 
in the Museum. During his tenure in this 
position, a question arose as to whether or 
not the administrative staff of the Museum 
had increased significantly in the several 
preceding years. Discussion went back and 
forth between the Senate and the Director 
of the Museum. Finally, Austin volunteered 
to get some real data on the issue. In a typ- 
ical A. B. Williams way, he systematically 
went through the entire telephone directory 
and counted museum administrators at 5- 
year intervals, and demonstrated that the 
type of positions that the Senate considered 
as ‘“‘administrative’’ had in fact increased 
significantly. 

He was the acknowledged expert on and 
leader in studies of the systematics of east- 
ern American decapod crustaceans. He is 
probably best known for his widely used 
monograph “Shrimps, lobsters, and crabs 
of the Atlantic coast of the eastern United 
States’’ published by the Smithsonian in 
1984. His earlier study on the decapods of 
the Carolinas published in 1965, a precursor 
to his 1984 monograph, was selected as a 
Science Citation Classic in 1983, a rare 
honor for a systematist. His invaluable pa- 
per (Williams 1987a) on the identification 
of spiny lobsters by color patterns of the 
tails grew into a book co-authored with I. 
Dore, entitled ‘‘Lobsters of the world—an 
illustrated guide’? (Williams & Dore 
1988e); these two publications are indis- 
pensable for anybody interested in this 
group of economically significant decapods. 

The primary focus of his research was 
the taxonomy, systematics, biogeography 
and evolution of various decapod groups, 
both fossil and Recent. He named 101 new 
decapod taxa (see list), including one su- 
perfamily, 2 families, 16 genera, 80 species, 
and 2 subspecies. Occasionally he also 
worked on other groups such as cirripeds, 
mysids, amphipods, and euphausiids, and 
even bird ecology. His publications provide 
us with a standard of excellence, and are 
well known for attention to detail, accuracy, 


and usefulness in the identification of spec- 
imens while at the same time giving insight 
into phylogenetic relationships. No major 
group of decapods escaped Austin’s atten 
tion. He published important works on 
crayfishes, peneaeoids, carideans, thalassin- 
ideans, lobsters, anomurans, and brachyu- 
rans. His landmark studies on swimming 
crabs of the genus Callinectes, mud shrimps 
of the family Upogebiidae, commercial lob- 
sters, xanthid crabs, and deep-sea hydro- 
thermal vent decapods, among others, have 
earned him a place in the history of Zool- 
ogy. His contributions to the systematics of 
hydrothermal vent decapods inspired other 
colleagues, and one genus and species of 
the crab family Bythograeidae Williams, 
1980, was named after him (Austinograea 
williamsi Hessler & Martin, 1989, Journal 
of Crustacean Biology 9(4):645—-661). In 
March of 1996 he traveled to Kumamoto, 
Japan, on a Japanese fellowship program to 
join Keiji Baba (Kumamoto University Fac- 
ulty of Ecucation) in the study of galatheids 
and other vent decapods from hydrother- 
mally active sites in the western Pacific. Al- 
though his work concentrated on aspects of 
systematics and evolution, he also pub- 
lished key studies on the biology of com- 
mercial penaeid shrimps, ecology of mero- 
plankton, larval genetics, and crustacean 
fisheries and mariculture. At the time of his 
death he had completed work but unfortu- 
nately left unpublished, an important revi- 
sion of the crab family Latreilliidae. 
Austin’s impact on carcinology is not 
limited to that derived from his publica- 
tions. Throughout his exemplary career he 
actively participated in many professional 
societies in addition to the Biological So- 
ciety of Washington. He was president of 
the Atlantic Estuarine Research Society 
(1960-1961); co-founder, secretary (1971-— 
1973), and president (1983-1985) of the 
Estuarine Research Federation; secretary 
(1985-1988) for the Society of Systematic 
Biology; associate editor for The Crusta- 
cean Society (1986-1991); and vice-presi- 
dent (1990-1991) and president (1991-— 


+ PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


1992) of the American Association for Zoo- 
logical Nomenclature. He was also a mem- 
ber of the American Association for the 
Advancement of Science, American Fish- 
eries Society, American Institute of Biolog- 
ical Sciences, American Institute of Fishery 
Research Biologists, American Society of 
Limnology and Oceanography, Society for 
Integrative and Comparative Biology (for- 
merly American Society of Zoologists), As- 
sociation of Systematic Collections, Ecolog- 
ical Society of America, Kansas Academy 
of Science, and Society for the Study of Evo- 
lution. His editorial activities with various 
journals and symposia proceedings pro- 
duced many significant contributions, and 
his involvement with doctoral and master- 
level students at various academic institu- 
tions led to the development of outstanding 
carcinologists who now must continue his 
legacy. His skills as reviewer were highly 
regarded by editors and grant program man- 
agers who were assured of a detailed, un- 
biased evaluation. 

Several of Austin’s papers won important 
awards or honorable mentions. His lobster 
identification paper (Williams 1987a), for 
example, won the highly regarded ‘*Nation- 
al Marine Fisheries Service Outstanding 
Publication Award” for best paper in the 
1997 Marine Fisheries Review. In recogni- 
tion of Austin’s life-time work, The Crus- 
tacean Society presented him in 1997 with 
their ““Excellence in Research Award”’ dur- 
ing a ceremony at the National Museum of 
Natural History, Smithsonian Institution, 
Washington, D.C. [see Lemaitre, R., 1998, 
Journal of Crustacean Biology 18(3):619-— 
620]. He was also honored with the ‘‘1999 
Elton Sette Award” from the Marine Fish- 
eries Section of the American Fisheries So- 
ciety. 

Austin will be remembered not only for 
his impressive scientific accomplishments 
but also for his human qualities. Unselfish 
almost to a fault, he made every effort to 
help colleagues and students alike. His 
height of 1.85 m (6'1”) gave him a towering 
physique which combined with his deep 


knowledge of decapods made him an im- 
posing figure to both students and junior 
colleagues; however, his modesty was such 
that he made sure to treat everyone as a 
friend or colleague of similar stature. He 
accepted life-time honors bestowed upon 
him by his peers only hesitantly, and main- 
tained until his end that he was undeserving 
of such attention. His personality, working 
habits, discipline, and inspirations date back 
to his early life experiences which he often 
mentioned to friends during casual conver- 
sations. One of his first jobs prior to enter- 
ing college was at his family’s farm in Ster- 
ling, Colorado, where he helped string fenc- 
es, some of which had to be modified to 
‘first class communications grade’”’ by in- 
sulating the top strand of barbed wire to 
carry telephone signals. This worked fine 
until it rained and the circuits became 
grounded by water. During his Sterling days 
he also taught high school. One of Austin’s 
scientific strengths was his observational 
and note taking abilities. He had been 
trained at Kansas to write reflective notes 
each night as if they were to be published. 
Those who have examined his field and of- 
fice notebooks are struck by how remark- 
ably clear and detailed they are. 

Austin often mentioned the impact of the 
Great Plains of Kansas and Colorado on his 
psyche. One of his closest friends, Gale A. 
Bishop (Georgia Southern University), has 
said that he was impressed with Austin’s 
collegiality when he first met him during a 
visit to the Smithsonian to study fossil 
decapods. Gale suggested that he might 
want to join him in the field in South Da- 
kota to collect fossils. Austin did so with 
much enthusiasm, and the two worked to- 
gether almost every summer from 1980 un- 
til the year of his death. Nancy Brannen 
Marsh (Science Department, Portal High 
School, Georgia) also joined them, and the 
three collaborated in studies of decapods of 
the Western Interior Cretaceous, collecting 
numerous fossil crabs, lobsters, and shrimps 
from the Carlile Shale and the Pierre Shale 
of South Dakota, Wyoming and Colorado. 


VOLUME 113, NUMBER 1 


These fossil collections have been donated 
to the Museum of Geology at the South Da- 
kota School of Mines, Rapid City. He con- 
fessed to Gale that the timing of their col- 
laboration was most appropriate as his wife 
Jean had passed away (1983) after a diffi- 
cult illness (with Austin as major care giv- 
er), and his return to the Great Plains was 
just what he needed to gain closure and 
healing from the loss. This theme, Gale 
says, ““came up many times and we con- 
cluded that getting back to our roots was an 
extremely healing process, both for Austin’s 
loss and for the loss of both of my parents; 
it was an annual ‘rehealing’ as we came 
back into harmony with our roots and 
Mother Earth. When working with Austin 
on fossils in the Western Interior his intel- 
lect and collegiality were always apparent. 
His interests spanned the sciences, arts, ed- 
ucation, and humanities. He often would 
visit the Rapid City Astronomy club to par- 
ticipate in telescopic observations, take us 
all out dancing at the Broken Boot Saloon 
in Rapid City, climb Bear Butte or Harney 
Peak, or head us up to Rushmore for the 
evening patriotic lighting program. While 
in the field, he brought new insights to pa- 
leontology, often seeing things we took for 
granted or forcing clearer explanation of 
our mutual deductions. Our collaboration 
was clearly very beneficial to Austin as 
well as to paleontologists Nancy Brennan 
Marsh, the late Reinhard Forster of Munich, 
and Georgia Southern students Mike Klug, 
Mehmet Samiratedu, and Amy Samiratedu. 
These insights were carried over into the 
laboratory and into collaborations on papers 
and research comparing Recent and fossil 
decapods.”’ 

The multi-faceted personality of Austin 
included a deep appreciation of the simple 
things of life, his family and friends. During 
the last decade or so of his life he was for- 
tunate to share many moments with Norma 
Samuels, of Fairfax, Virginia, whose com- 
panionship undoubtedly enriched his life. 
Austin developed a passion for ballroom 
and international dancing, and a love for 


choral music. He actively participated in 
several choral groups, including the Wash- 
ington Cathedral Choral Society which per- 
formed in the National Cathedral, Washing- 
ton, D.C. One of his performances is pre- 
served on an audio CD-ROM entitled ‘‘Mil- 
lenium; Russian Choral Music’’ (1990 
Centaur Record Inc.). He worshipped, sang 
and was an active member of the New York 
Avenue Presbyterian Church, of Washing- 
ton, D.C., where well-attended and emo- 
tional services were held for him on Octo- 
ber 30, officiated by The Rev. Robert H. 
Craig. 

Austin’s remains are buried in Marion, 
Kansas, alongside those of his wife. 


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, & A. B. Leonard The crayfishes of 

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. Identification of juvenile shrimp (Pen- 

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Ogyrides (Caridea: Ogyrididae).—Chesapeake 
Science 13(2):145—-148. 

, & K. H. Bynum. A ten-year study of 
meroplankton in North Carolina estuaries: Am- 
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. A ten-year study of meroplankton in 
North Carolina estuaries: Mysid shrimps.— 
Chesapeake Science 13(4):254—262. 

. Decapod crustaceans of the Chesa- 
peake Bay.—Chesapeake Science 13 (Supple- 
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. Allactaea lithostrota, a new genus and 
species of crab (Decapoda: Xanthidae) from 
North Carolina, U.S.A.—Proceedings of the Bi- 
ological Society of Washington 87(3):19—26. 

. Two new axiids (Crustacea: Decapoda: 
Thalassinidea: Calocaris) from North Carolina 
and the Straits of Florida.—Proceedings of the 
Biological Society of Washington 87(39):451— 
464. 


1970. 


LST La. 


197 1b. 


1972a. 


1972b. 


1972c. 


972d" 


1972e. 


19721. 


1974a. 


1974b. 


1974c. 


. A new species of Hypsophrys (Deca- 
poda: Homolidae) from the Straits of Florida, 
with notes on related crabs.—Proceedings of 
the Biological Society of Washington 87(42): 
485-492. 

, T. E. Bowman, & D. M. Damkaer. 
Distribution, variation, and supplemental de- 
scription of the opossum shrimp, Neomysis 
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Bulletin 72(3):835-842. 

. The swimming crabs of the genus Cal- 


1974d. 


1974e. 


VOLUME 113, NUMBER 1 


linectes (Decapoda: Portunidae).—Fishery Bul- 
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. Marine flora and fauna of the north- 
eastern United States. Crustacea: Decapoda.— 
NOAA Technical Report NMFS Circular 389: 
1-50. 


1974f. 


1976a. . Distinction between a Gulf of Mexico 
and a Carolinian Atlantic species of swimming 
crab Ovalipes (Decapoda: Portunidae).—Pro- 
ceedings of the Biological Society of Washing- 
ton 89(14):205—214. 

. Integumental organs of unknown func- 
tion on chelipeds of deep-sea crabs, genus Hyp- 
sophrys.—Journal of Morphology 150(4):889— 
899. 


1976b. 


1977a. 


, J. K. Shaw, & T. S. Hopkins. Stilbom- 
astax, anew genus of spider crab (Mayjidae: Ty- 
chinae) from the West Indies region, with notes 
on American relatives.—Proceedings of the Bi- 
ological Society of Washington 90(4):884—893. 
, & R. L. Wigley. Distribution of deca- 
pod Crustacea off northeastern United States 
based on specimens at the Northeast Fisheries 
Center, Woods Hole, Massachusetts—-NOAA 
Technical Report NMFS Circular 407:i—i1i, 1— 
44. 


1977b. 


1978a. . Transfer to Pseudomedaeus of the xan- 

thid crab Micropanope distinctus (Rathbun).— 

Proceedings of the Biological Society of Wash- 

ington 91(2):546—557. 

. True crabs. (Unpaginated) in W. Fi- 
scher, ed., FAO Species Identification Sheets for 
Fishery Purposes. Western Central Atlantic 
(Fishing Area 31), vol. 6, Food and Agriculture 
Organization of the United Nations, Rome. 

1979a. Herbst, G. N., 3 6B By Boothe, Jr. 
Reassessment of northern geographic limits for 
decapod crustacean species in the Carolinian 
Province, USA; some major range extensions 
itemized.—Proceedings of the Biological Soci- 
ety of Washington 91(4):989—-998. 

, & T. W. Duke. Chapter 6. Crabs (Ar- 

thropoda: Crustacea: Decapoda: Brachyura). Pp. 

171—233 in C. W. Hart and S. M.N. Fuller, eds., 

Pollution ecology of estuarine invertebrates. 

Academic Press, N.Y., 406 pp. 

. A new crab family from shallow wa- 

ters of the West Indies (Crustacea: Decapoda: 

Brachyura).—Proceedings of the Biological So- 

ciety of Washington 92(2):399—413. 

. A new crab family from the vicinity of 

submarine thermal vents on the Galapagos Rift 

(Crustacea: Decapoda: Brachyura).—Proceed- 

ings of the Biological Society of Washington 

93(2):443-—472. 

. Western Atlantic species of the cari- 

dean shrimp genus Ogyrides.—Journal of Crus- 

tacean Biology 1(1):143—147. 


1978b. 


1979b. 


S72. 


1980. 


198 1a. 


198 1b. , & D. McN. Williams. Carolinian re- 

cords for American lobster, Homarus american- 

us, and tropical swimming crab, Callinectes bo- 
courti. Postulated means of dispersal.—Fishery 

Bulletin 79(1):192—198. 

. Revision of the genus Latreillia Roux 

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boratorio de Tecnologia della Pesca, Ancona, It- 

aly 3(2—5):227-255. 

, & E A. Chace, Jr. A new caridean 

shrimp of the family Bresiliidae from thermal 

vents of the Galapagos Rift.—Journal of Crus- 
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. Marine decapod crustaceans of the 

Carolinas. Fishery Bulletin 65:1-298.—Current 

Contents 14(17):20. 

, & C. L. Van Dover. A new species of 

Munidopsis from submarine thermal vents of 

the East Pacific Rise at 21°N (Anomura: Gal- 

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. The mud crab, Panopeus herbstii, s.1. 
Partition into six species (Decapoda: Xanthi- 
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1984b. Sullivan, B., K. Miller, K. Singleton, A. G. 
Scheer, & . Electrophoretic analyses of 
hemocyanins from four species of mud crabs, 
genus Panopeus, with observations on the ecol- 
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885. 

1984c. Reams, R. C., & . Mud crabs of the 
Panopeus herbstii H. M. Edw., s.1., complex in 
Alabama, U.S.A.—Fishery Bulletin 81(4):885— 
890. 

1984d. Van Dover, C. L., , & J. R. Factor. The 
first zoeal stage of a hydrothermal vent crab 
(Decapoda: Brachyura: Bythograeidae).—Pro- 
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ton 97(2):413-418. 

1984e. Millikin, M. R., & . Synopsis of bio- 
logical data on the blue crab, Callinectes sapi- 
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1, FAO Fisheries Synopsis 138:1—39. 

. Shrimps, lobsters, and crabs, of the At- 

lantic coast of the eastern United States, Maine 

to Florida. Smithsonian Institution Press, 550 


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1982a. 


1982b. 


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1983b. 


1984a. 


1984f. 


1984¢g. . Review: Shallow-water crabs. By R. 

W. Ingle, 1983, Linnean Society of London and 

the Estuarine and Brackish-Water Sciences As- 

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206 pp.—Estuaries 7(3):266—267. 

. Review: Crustacean Phylogeny. F R. 
Schram, ed., 1983, Crustacean Issues 1. Based 
on a symposium, Dallas, Dec. 1981, Balkema, 
Rotterdam, 372 pp.—American Scientist 72(5): 
514-515. 

1985a. Van Dover, C. L., J. R. Factor, 


1984h. 


, &76. 


8 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


J. Berg, Jr. Reproductive patterns of decapod 
crustaceans from hydrothermal vents of the 
Eastern Pacific. In M. L. Jones, ed., The hydro- 
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. Summary comments. /n M. L. Jones, 

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cific: An overview.—Bulletin of the Biological 

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. Mud shrimps, Upogebia, from the 
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Memoir 14:1—60. 

1986b. Bishop, G. A., & . The fossil lobster 
Linuparus canadensis, Carlile Shale (Creta- 
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search 2(3):372—387. 

, & P A. Rona. Two new caridean 

shrimps (Bresiliidae) from a hydrothermal field 

on the Mid-Atlantic Ridge.—Journa! of Crus- 
tacean Biology 6(3):446—462. 

, & R. D. Turner. Squat lobsters (Gal- 

atheidae: Munidopsis) associated with mesh-en- 

closed wood panels submerged in the deep 

sea.—Journal of Crustacean Biology 6(3):617— 

624. 


1985b. 


1986a. 


1986c. 


1986d. 


1986e. 


, & D. L. Felder. Analysis of stone 
crabs: Menippe mercenaria (Say), restricted, 
and a previously unrecognized species de- 
scribed (Decapoda: Xanthidae).—Proceedings 
of the Biological Society of Washington 99(3): 
517-543. 

. Foreword. Jn D. A. Wolfe, ed., Estu- 
arine variability. Academic Press, New York, 
509 pp. 


1986f. 


1987a. . Lobsters—identification, world distri- 

bution, and U.S. trade.—Marine Fisheries Re- 

view 48(2):1—36. 

. More records for shrimps of the genus 
Rimicaris (Decapoda: Caridea: Bresiliidae) 
from the Mid-Atlantic Rift—Journal of Crus- 
tacean Biology 7(1):105. 

1987c. Wetherbee, D. K., & . The late eigh- 
teenth century paintings by Rabie of Haitian 
mollusks and crustaceans. Further Contributions 
on the History of Zoology in Hispaniola, Art. 
3:18—48. Privately published, Shelburne, Mas- 
sachusetts. 


1987b. 


1987d. 


. Upogebia synagelas, new species, a 
commensal mud shrimp from sponges in the 
western central Atlantic (Decapoda: Upogebi- 
idae).—Proceedings of the Biological Society 
of Washington 100(3):590—595. 

. Indo-Pacific spiny lobsters in the U.S. 
National Museum of Natural History collected 
from 1963 to 1981 (Crustacea: Palinuridea).— 
Crustaceana 55(3):313-316. 


1988a. 


1988b. 


. Notes on decapod and euphausiid 
crustaceans, continental margin, western Atlan- 
tic, Georges Bank to western Florida, USA.— 
Fishery Bulletin 86(1):67—76. 

. New marine decapod crustaceans from 
waters influenced by hydrothermal discharge, 
brine, and hydrocarbon seepage.—Fishery Bul- 
letin 86(2):263—287. 

. Conjoined twin adult shrimp (Deca- 
poda: Penaeidae).—Fishery Bulletin 86(3):595— 
597 


1988c. 


1988d. 


1988e. 


, & I. Dore. Lobsters of the world - An 
illustrated guide; Lobsters of the world in U.S. 
trade. Osprey Books, Huntington, New York, 
186 pp. 

1989a. 


, & C. A. Child. Comparison of some 
genera and species of box crabs (Brachyura: 
Calappidae), southwestern North Atlantic, with 
description of a new genus and species.—Fish- 
ery Bulletin 87(1):105—121. 
, & P. J. B. Scott. Upogebia corallifora, 
a new species of coral-boring shrimp from the 
West Indies (Decapoda: Upogebiidae).—Pro- 
ceedings of the Biological Society of Washing- 
ton 102(2):405—410. 
» Le Ge Abele;-D. L. Feldes eat 
Hobbs, Jr., R. B. Manning, P- A. McLaughlin, 
& I. Pérez Farfante. Common and scientific 
names of aquatic invertebrates from the United 
States and Canada: decapod crustaceans.— 
American Fisheries Society Special Publication 
No. 17:1-77. 
, & J. J. McDermott. An eastern United 
States record for the western Indo-Pacific crab, 
Hemigrapsus sanguineus (Crustacea: Decapo- 
da: Grapsidae).—Proceedings of the Biological 
Society of Washington 103(1):108—109. 
, & K. Baba. New squat lobsters (Gal- 
atheidae) from the Pacific Ocean: Mariana Back 
Arc Basin, East Pacific Rise, and Cascadia Ba- 
sin.—Fishery Bulletin 87(4):899-910. 
1990c. Kensley, B., & . Axlopsis eximia, a new 
thalassinidian shrimp (Crustacea, Decapoda, 
Axiidae) from the Middle Eocene of South Car- 
olina.—Journal of Paleontology 64(5):798—802. 
, & E. E. Boschi. Panopeus margentus, 
a new crab from the Argentine warm temperate 
subregion (Decapoda: Xanthidae).—Proceed- 
ings of the Biological Society of Washington 
103(3):598—-601. 
, & N. Ngoc-Ho. Pomatogebia, a new 
genus of thalassinidean shrimps from western 
hemisphere tropics (Crustacea: Upogebiidae).— 
Proceedings of the Biological Society of Wash- 
ington 103(3):614—616. 
, & R. W. Heard. Upogebia spinistipula, 
a new burrowing shrimp from the Florida shelf, 
northeastern Gulf of Mexico (Decapoda: Upo- 


1989b. 


1989c. 


1990a. 


1990b. 


1990d. 


1990e. 


199 1a. 


VOLUME 113, NUMBER 1 


gebiidae).—Proceedings of the Biological So- 

ciety of Washington 104(1):49—54. 

, & R. B. Moffitt. Crabs from the Mar- 
iana Archipelago: Bothromaia griffini new ge- 
nus and species (Brachyura: Majidae), and re- 
marks on Poupinia hirsuta Guinot (Homolo- 
idea, Poupiniidae).—Proceedings of the Biolog- 
ical Society of Washington 104(3):569-582. 

1991c. Bishop, G. A., & 
sonorum, new species, a crab (Decapoda: Ca- 
lappidae) from the Cretaceous Carlile Shale 
(Turonian), Western Interior United States.— 
Journal of Crustacean Biology 11(3):451—459. 

. Comments on the proposed conser- 
vation of the specific name of Artemia francis- 
cana Kellogg, 1906 (Crustacea, Branchiopoda). 
(Case 2728; see BZN 47:178—-183; 48:57).— 
Bulletin of Zoological Nomenclature 48(3): 
246-247. 

1991le. Van Dover, C. L., & . Egg size in squat 
lobsters (Galatheoidea): Constraint and free- 
dom. Pp. 143-156 in A. Wenner & A. Kuris, 
eds., Crustacean Egg Production. Crustacean Is- 
sues, vol. 7, A. A. Balkema/Rotterdam, 401 pp. 

, & R. Wahle. Distinguishing juvenile 

stages of Jonah and Atlantic rock crabs, Cancer 

borealis and C. irroratus (Crustacea: Decapoda: 

Cancridae).—Journal of Crustacean Biology 

12(3):464—466. 

. Review: Decapod Crustacea of the At- 

lantic Coast of Canada. By H. J. Squires, 1990, 

Canadian Bulletin of Fisheries and Aquatic Sci- 

ences 221:532 pp.—Journal of Crustacean Bi- 

ology 12(2):329. 

. Review: Marine Lobsters of the World. 

An Annotated and Illustrated Catalog of Species 

of Interest Known to Date. FAO Species Cata- 

log. By L. B. Holthuis, 1991, FAO Fisheries 

Synopsis No. 125, vol. 13:292 pp. Rome.—The 

Lobster Newsletter 5:14. 

. Comments on the proposed conser- 

vation of the specific names of Gebia major ca- 

pensis Krauss, 1843 and G. africana Ortmann, 

1894 (currently Upogebia capensis and U. af- 

ricana; Crustacea, Decapoda) by the designa- 

tion of a replacement neotype for U. capensis. 

(Case 2827; BZN 49(3):187—190.).—Bulletin 

of Zoological Nomenclature 50(2):143—144. 

. Mud shrimps, Upogebiidae, from the 

western Atlantic (Crustacea: Decapoda: Thal- 

assinidea).—Smithsonian Contributions to Zo- 

ology 544:i-iii, 1-77. 

. Reflections on crab research in North 

America since 1758. Pp. 259-273 in E Trues- 

dale, ed., History of Carcinology, Crustacean Is- 

sues, vol. 8., A. A. Balkema/Rotterdam, 445 pp. 

, & L. G. Eldredge. A new species of 

spider crab from Guam, Rochinia decipiata 


199 1b. 


. Necrocarcinus ol- 


199 1d. 


1992a. 


1992b. 


L9D2c. 


1993a: 


1993b. 


1593¢. 


1994. 


(Brachyura: Majidae).—Crustacean Research 
23:1-4. 

1995a. Rodriguez, G., & . Epilobocera weth- 
erbeei, a new species of freshwater crab (De- 
capoda: Brachyura: Pseudothelphusidae) from 
Hispaniola.—Proceedings of the Biological So- 
ciety of Washington 108(1):76—-83. 

1995b. Kornfield, [., , & R. S. Steneck. As- 
signment of Homarus capensis (Herbst, 1792), 
the Cape lobster of South Africa, to the new 
genus Homarinus (Decapoda: Nephropidae).— 
Fishery Bulletin 93(1):97—102. 

, & EC. Dobbs. A new genus and spe- 

cies of caridean shrimp (Crustacea: Decapoda: 

Bresiliidae) from hydrothermal vents on Loihi 

Seamount, Hawaii.—Proceedings of the Biolog- 

ical Society of Washington 108(2):228—237. 

. Chapter 2, Taxonomy and evolution. 
Pp. 13-21 in J. R. Factor, ed., Biology of the 
lobster Homarus americanus. Academic Press, 
New York, 529 pp. 

1995e. Pequegnat, L. H., & . Two new species 
of Munidopsis (Decapoda: Anomura: Galathei- 
dae) from the Western Atlantic Ocean.—Journal 
of Crustacean Biology 15(4):786—792. 

. Comments on the proposed conser- 

vation of the generic names Monstrilla Dana, 

1849 and Thaumaleus Kréyer, 1849 (Crustacea, 

Copepoda) (Case 2894; see BZN 52(3):245-— 

249).—Bulletin of Zoological Nomenclature 

53(2):122-123. 

. Comments on the proposed conser- 

vation of the generic name Glomeris Latreille, 

1802 (Diplopoda) and the specific name of Ar- 

madillo vulgaris Latreille, 1804 (Crustacea, Is- 

opoda), and the application for a ruling on the 

status of the name Armadillo Latreille, 1802 

(Crustacea, Isopoda). (Case 2909; see BZN 

52(3):236—244).—Bulletin of Zoological No- 

menclature 53(2):121—122. [Resolved in Opin- 
ion 1897, Bulletin of Zoological Nomenclature 

55(2):124—128, June 1998.] 

Bythograea Williams, 
1980. Arthropoda, Crustacea, Decapoda, Brach- 
yura, Brachyrhyncha, Bythograeoidea, Bytho- 
graeidae. p. 212 in D. Desbruyéres and M. Se- 
gonzac, eds., Handbook of deep-sea hydrother- 
mal vent fauna. Editions IFREMER, Brest, 279 
PP- 

1997b. Curran, H. A., & . Ichnology of an in- 
tertidal carbonate sand flat: Pigeon Creek, San 
Salvador Island, Bahamas. Pp. 33—46 in J. L. 
Carew, ed., Proceedings of the 8th Symposium 
on the Geology of the Bahamas: San Salvador, 
Bahamian Field Station, June 9—13, 1995. 

. Occurrence of three species of mud 

shrimps in aquaculture ponds on Caribbean 

coasts of Venezuela and Colombia, with a re- 


1995¢: 


1995d. 


1996a. 


1996b. 


1997a. 


thermydron 


1997c. 


10 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


description of Upogebia omissago Williams, 

1993 (Decapoda: Upogebiidae).—Proceedings 

of the Biological Society of Washington 110(3): 

412-416. 

. Two new species and a range exten- 
sion of mud shrimps, Upogebia, from Pacific 
Costa Rica and Mexico (Decapoda: Thalassin- 
idea: Upogebiidae).—Proceedings of the Bio- 
logical Society of Washington 110(4):617- 623. 

1998a. Baba, K., & . New Galatheoidea (Crus- 
tacea, Decapoda, Anomura) from hydrothermal 
systems in the West Pacific Ocean: Bismarck 
Archipelago and Okinawa Trough.—Zoosyste- 
ma 20(2):143—156. 

, & J. L. Hernandez-Aguilera. A new 

species of mud shrimp, Upogebia toralae, from 

Veracruz, Mexico (Decapoda: Thalassinidea: 

Upogebiidae).—Proceedings of the Biological 

Society of Washington 111(4):908—911. 

, & R. Vargas. A new species of mud 
shrimp, Upogebia cortesi, from Pacific Costa 
Rica (Decapoda: Thalassinidea: Upogebi- 
idae).—Proceedings of the Biological Society 
of Washington 113(1):13—16. 

2000b. Bishop, G. A., & . Fossil crabs from 
Tepee Buttes, submarine seeps of the Late Cre- 
taceous Pierre Shale, South Dakota and Colo- 
rado, U.S.A.—Journal of Crustacean Biology 
(Special no. 2):(in press). 

In press. . Comment on the proposed designa- 
tion of Scottia pseudobrowniana Kempf, 1971 
as the type species of Scottia Brady & Norman, 
1889 (Crustacea: Ostracoda) (Case 2896; see 
BZN 51(4):304—305).—Bulletin of Zoological 
Nomenclature. 

In press. . Comment on the proposed conser- 
vation of both the generic and specific names 
Cubaris murina Brandt, 1833 (Case 2910; see 
BZN 52(2):153—156).—Bulletin of Zoological 
Nomenclature. 


1997d. 


1998b. 


2000a. 


List of taxa named by Austin B. Williams 


Taxa are listed alphabetically within each 
major decapod group. Holotype deposition 
and number is indicated for all species and 
subspecies. Asterisk indicates fossil taxa. 
Abbreviations for repositories are as fol- 
lows: AHKE Allan Hancock Foundation, 
University of Southern California (now 
Natural History Museum of Los Angeles 
County); AMNH, American Museum of 
Natural History, New York; MCZ, Museum 
of Comparative Zoology, Harvard Univer- 
sity, Cambridge, Massachusetts; MZUSP, 
Museu de Zoologia, Universidade de Sao 


Paulo, Brazil; SDSM, Museum of Geology, 
South Dakota School of Mines, Rapid City; 
SDSNH, San Diego Society of Natural His- 
tory, California; UKMNH, University of 
Kansas Museum of Natural History, 
Lawrence; USNM, National Museum of 
Natural History, Smithsonian Institution, 
Washington, D.C. 


Decapoda 


Caridea 

Alvinocaris Williams & Chace, 1982b. 

Alvinocaris lusca Williams & Chace, 
1982b. USNM 184534. 

Alvinocaris markensis Williams, 1988c. 
USNM 234286. 

Alvinocaris muricola Williams, 1988c. 
USNM 234288. 

Alvinocaris stactophila Williams, 1988c. 
USNM 234291. 

Leptalpheus Williams, 1965b. 


Leptalpheus forceps Williams, 1965b. 
USNM 111084. 
Ogyrides hayi Williams, 198la. USNM 


47958. 

Ogyrides limicola Williams, 1955a. USNM 
96675. 

Opaepele Williams & Dobbs, 1995c. 

Opaepele loihi Williams & Dobbs, 1995c. 
USNM 251447. 

Rimicaris Williams & Rona, 1986c. 

Rimicaris chacei Williams & Rona, 1986c. 
USNM 228452. 

Rimicaris exoculata Williams & Rona, 
1986c. USNM 228443. 


Astacidea 


Homarinus Kornfield, Williams & Steneck, 
1995b. 

Orconectes eupunctus Williams, 1952a. 
UKMNH T4250. 

Orconectes meeki brevis Williams, 1952a. 
UKMNH T8140. 

Orconectes nana marcus Williams, 1952a. 
UKMNH T4970. 

Orconectes nana 
UKMNH T6640. 


Williams, 1952a. 


VOLUME 113, NUMBER 1 


Orconectes neglectus chaenodactylus Wil- 
liams, 1952a. UKMNH T4420. 

Orconectes ozarkae Williams, 
UKMNH T6150. 


LO524. 


Thalassinidea 


Aethogebia Williams, 1993b. 

Aethogebia gorei Williams, 1993b. USNM 
251425. 

*Axiopsis eximia Kensley & Williams, 
1990c. USNM 219431. 

Calocaris (Calastacus) jenneri Williams, 
1974b. USNM 150472. 

Calocaris (Calastacus) oxypleura Williams, 
1974b. USNM 101651. 

Pomatogebia Williams & Ngoc-Ho, 1990e. 

Upogebia acanthops Williams, 1986a. 
USNM 213194. 


Upogebia aestuari Williams, 1993b. 
USNM 251407. 

Upogebia aquilina Williams, 1993b. 
USNM 251426. 

Upogebia baldwini Williams, 1997d. 
USNM 251486. 

Upogebia bermudensis Williams, 1993b. 
MCZ 12873. 

Upogebia burkenroadi Williams, 1986a. 
SDSNH 3985. 

Upogebia careospina Williams, 1993b. 


USNM 138899. 

Upogebia casis Williams, 1993b. USNM 
251224. 

Upogebia cocosia Williams, 1986a. USNM 
213268. 

Upogebia coralliflora Williams & Scott, 
1989b. USNM 230075. 

Upogebia cortesi 2000a. USNM 291186. 

Upogebia dawsoni Williams, 1986a. 
AHF2566. 

Upogebia felderi Williams, 1993b. USNM 
251430. 

Upogebia galapagensis Williams, 1986a. 
USNM 213223. 

Upogebia inomissa Williams, 
USNM 251396. 

Upogebia jonesi Williams, 1986a. USNM 
213195. 


1993b. 


| 


Upogebia lepta Williams, 1986a. USNM 
213270. 

Upogebia maccraryae Williams, 1986a. 
USNM 213202. 

Upogebia macginitieorum Williams, 1986a. 
USNM 213219. 

Upogebia molipollex Williams, 1993b. 
AMNH 6820. 

Upogebia omissago 
USNM 222057. 

Upogebia onychion Williams, 1986a. AHF 
4133. 

Upogebia paraffinis Williams, 
MZUSP 8049 

Upogebia_ pillsbury 
USNM 251435. 

Upogebia ramphula Williams, 
USNM 213446. 

Upogebia schmitti Williams, 1986a. AHF 
3933: 

Upogebia spinistipula Williams & Heard, 
199la. USNM 239251. 

Upogebia synagelas Williams, 
USNM 233572. 

Upogebia tenuipollex Williams, 1986a. 
USNM 213236. 

Upogebia thistlei Williams, 1986a. USNM 
21S251% 

Upogebia toralae Williams & Hernandez- 
Aguilera, 1998b. USNM 285522. 


Williams, 1993b. 
1993b. 
1993b. 


Williams, 


1986a. 


1987d. 


Upogebia vargasae Williams, 1997d. 
USNM 251484. 
Upogebia veleronis Williams, 1986a. 
(USNM 213272) 
Anomura 
Munidopsis alvisca Williams, 1988c. 


USNM 234294. 

Munidopsis glabra Pequegnat & Williams, 
1995e. USNM 251455. 

Munidopsis granosicorium Williams & 
Baba, 1990b. USNM 240205. 

Munidopsis lentigo Williams & Van Dover, 
1983b. USNM 191160. 

Munidopsis lignaria Williams & Baba, 
1990b. USNM 240202. 

Munidopsis marianica Williams & Baba, 
1990b. USNM 240198. 


{[F9, 


Shinkaia Baba & Williams, 1998a. 

Shinkaia crosnieri Baba & Williams, 
1998a. USNM 251480. 

Uroptychus edisonicus Baba & Williams, 
1998a. USNM 251479. 


Brachyura 


Allactaea Williams, 1974a. 

Allactaea lithorostrata Williams, 
USNM 143770. 

Bothromaia Williams & Moffit, 1991b. 

Bothromaia  griffini Williams & Moffit, 
1991b. USNM 250884. 

Bythograeoidea Williams, 1980. (Super- 
family). 

Bythograeidae Williams, 1980. (Family). 

Bythograea Williams, 1980. 

Bythograea mesatlantica Williams, 1988c. 
USNM 234300. 

Bythograea thermydron Williams, 
USNM 172830. 

Callinectes similis Williams, 1966b. USNM 
113341. 

Cyclozodion Williams & Child, 1989a. 

Cyclozodion tuberatum Williams & Child, 
1989a. USNM 234462. 

Epilobocera wetherbeei Rodriguez & Wil- 
liams, 1995a. USNM 268832. 

Eplumula Williams, 1982a. 

*Heus Bishop & Williams, 2000b. 

*Heus foersteri Bishop & Williams, 2000b. 
SDSM 11016. 

Hypsophrys noar Williams, 1974c. USNM 
150816. 


1974a. 


1980. 


Latreillia manningi Williams, 1982a. 
USNM 57071. 

Latreillia metanesa Williams, 1982a. 
USNM 74570. 


Menippe adina Williams & Felder, 1986e. 
USNM 228862. 

Mimilambridae Williams, 1979c. (Family). 

Mimilambrus Williams, 1979c. 

Mimilambrus wileyi Williams, 
USNM 172222. 

*Necrocarcinus olsonorum Bishop & Wil- 
liams, 1991c. SDSM 11000. 


19796. 


2 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Ovalipes stephensoni Williams, 1976a. 
USNM 155110. 

Panopeus austrobesus Williams, 1984a. 
USNM 59462. 

Panopeus margentus Williams & Boschi, 
1990d. USNM 239191. 

Panopeus meridionalis Williams, 
USNM 99846. 

*Plagiophthalmus bjorki Bishop & Wil- 
liams, 2000b. SDSM 11021. 

*Raninella manningi Bishop & Williams, 
2000b. SDSM 11018. 

Rochinia decipiata Williams & Eldredge, 
1994. USNM 251434. 

Stilbomastax Williams, Shaw & Hopkins, 
1977a. 


1984a. 


Acknowledgments 


We would like to thank David McN. Wil- 
liams for providing valuable family facts, 
and also several of Austin’s colleagues and 
friends for contributing photographs and 
other information. In particular, we would 
like to mention Norma Samuels who kindly 
provided photographs and insight on as- 
pects of Austin private life; Gale A. Bishop, 
for providing valuable and often emotional 
testimony on his experiences in the field; 
Raymond B. Manning and Brian Kensley 
for reminiscing on their interactions with 
Austin; and Keiji Baba for information and 
photographs. The invaluable help of Mi- 
chael Vecchione and the dedicated staff of 
the Systematics Laboratory, National Ma- 
rine Fisheries Service, at the Smithsonian, 
in locating and compiling files and data on 
Austin’s professional life, is gratefully ac- 
knowledged; these include Lara Cooper, 
Ruth Gibbons, David Hardy, Keiko Hira- 
tsuka Moore, Roosevelt McMillan, Mary 
Mickevich, Tom Munroe, and Martha Ni- 
zinski. Thanks also to Rose A. Gulledge for 
processing electronic images of the photo- 
graphs, and compiling bibliographic infor- 
mation; Chad Walter for proof-reading the 
bibliography; and Molly K. Ryan for put- 
ting together the figure. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


113(1):13—16. 2000. 


A new species of mud shrimp, Upogebia cortesi, from Pacific Costa 
Rica (Decapoda: Thalassinidea: Upogebiidae) 


Austin B. Williams} and Rita Vargas 


(ABW) National Marine Fisheries Service Systematics Laboratory, National Museum of 
Natural History, Smithsonian Institution, Washington, D.C. 20560, U.S.A.; (RV) Museo de 
Zoologia, Escuela de Biologia, Universidad de Costa Rica, 2060 Costa Rica (+: deceased 
October 27, 1999) 


Abstract.—Upogebia cortesi, a new species of mud shrimp from the Pacific 
side of Costa Rica is described and illustrated. The type series was dredged 
parallel to shore in water of 30—40 m depth. The species shares lack of a strong 
proximal mesioventral spine on the merus of the second pereopod with several 
members of the genus from the western hemisphere; moreover, the species 
stands alone in having spineless articles on pereopods 1—5 except for fingers 
of the chelae. The rostrum has no ventral spines. Similarities between this and 
related species are emphasized in a partial abridgment of the key to upogebiid 


species in the eastern Pacific. 


Eight species of the family Upogebiidae 
have been reported for the Pacific coast of 
Costa Rica (Vargas & Cortés 2000). Of 
these Pomatogebia cocosia (Williams 
1986) and Upogebia vargasae Williams, 
1997, were described based on material 
from Costa Rica. During the 1998 Mollusk 
Workshop organized by INBio at the “‘Re- 
serva Absoluta de Cabo Blanco’’, Peninsula 
de Nicoya, a dredge sample conducted be- 
tween the mainland and Isla Cabo Blanco, 
at 30—40 m depth, yielded three specimens 
of a new species of Upogebia Leach, 1814, 
described here. The substrate where the 
specimens were obtained included calcare- 
ous algae and rock fragments; at least 10 
species of brachyuran crabs and several 
mollusk species were also found in the sam- 
ple. 

Specimens are deposited in the National 
Museum of Natural History, Smithsonian 
Institution, Washington, D.C. (USNM), and 
in Museo de Zoologia, Escuela de Biologia, 
Universidad de Costa Rica, San José 
(UCRMZ), as indicated in the species ac- 
count. 


Upogebia cortesi, new species 
Fig. 1 


Material.—Costa Rica, NW side of Isla 
Cabo Blanco, dredged parallel to coast, 30— 
40 m, 16-17 May 1998: USNM 291186, 
male holotype; UCRMZ 2220-06, male par- 
atype; USNM 291187, female ovigerous 
paratype. 

Diagnosis.—Rostrum with sides convex 
in dorsal view, obsolescent pair of spines 
on submedian anterior margin; projections 
to either side of rostrum slender and spine- 
like; no spine on postocular margin; ante- 
rior gastric region bearing many spines 
nearly hidden in patch of dense setae. Ab- 
dominal sternites unarmed. Telson subrect- 
angular, sides slightly crenulate. Merus of 
cheliped lacking subdistal dorsal spine and 
spines on ventral margin; carpus essentially 
spineless; palms spineless. Pereopods 2—5 
spineless; pereopod 2 without proximal me- 
sioventral spine on merus. 

Description.—Rostrum (Fig. la,b) hori- 
zontal in lateral view with tip slightly ex- 
ceeding eyestalks; convex in dorsal aspect, 
with pair of submedian obsolescent blunted 


14 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig? tl. 


Upogebia cortesi, new species, male holotype, USNM 291187: a, Carapace and cephalic region, 


lateral; b, Anterior carapace, dorsal; c, Cheliped, right lateral; d, Chela and carpus, right mesial; e, Right pereopod 
2; f, Right pereopod 3; g, Right pereopod 4; h, Right pereopod 5; i, Telson, left uropod, and part of abdominal 


segment 6, dorsal. Scale = 3 mm. 


spines on anterior margin; 2 subdistal erect 
acute dorsal spines followed on each side 
by about 6 smaller marginal spines along 
either side of field of small spines on an- 
terior gastric region of carapace, spines on 
anterior part of field hidden in dense patch 
of setae. Lateral ridge on either side of an- 
terior gastric region extended anteriorly to 
spine similar in size to dorsal spines on ros- 
trum, ridge bearing crest of 18 spines, an- 
terior 4 spines larger than succeeding 


spines. Posterior region of carapace gla- 
brous. Shoulder lateral to cervical groove 
bearing no spines below intersection with 
thalassinidean line, latter continuing to pos- 
terior margin of carapace without interrup- 
tion; postocular margin of carapace un- 
armed. 

Abdominal sternites unarmed. 

Telson (Fig. 17) subrectangular, wider 
than long, posterior margin nearly straight, 
smooth; transverse proximal ridge promi- 


VOLUME 113, NUMBER 1 


nent, rather broad, lateral ridge at each side 
also rather broad; lateral margins almost 
imperceptibly crenulate. 

Eyestalk stout, horizontal, shorter than 
rostrum; lower margin slightly convex; cor- 
nea narrower than diameter of stalk and di- 
rected laterally. 

Antennular peduncle (Fig. la) reaching 
to about base of terminal article of antennal 
peduncle, combined length of proximal 2 
articles subequal to length of terminal arti- 
cle. 

Antennal peduncle (Fig. la) with distal 
article and distal half of penultimate article 
extending beyond tip of rostrum; moderate 
compressed scale bearing obsolescent an- 
teroventral spine. 

Maxilliped 3 bearing epipod. 

Epistomial projection rather broad in lat- 
eral view, bearing prominent, acuminate 
apical spine. 

Chelipeds (Fig. lc,d) lacking spine on 
ventral margin of merus. Carpus trigonal, 
with barely perceptible longitudinal lateral 
groove, obsolescent spine at anterior ven- 
trolateral corner. Chela length about 2.7 
times chela height. Fixed finger slender, 
with extended tip. Dactyl thick, longer than 
fixed finger, drawn to corneous tip, and at 
midlength bearing small tooth on occlusive 
edge opposing tip of fixed finger. 

Pereopod 2 (Fig. le) reaching about to 
distal edge of cheliped palm; pereopods 2-— 
4 (Fig. 1f-/) spineless. Pereopod 5 of usual 
form, with cleaning brush on propodus. 

Uropods (Fig. 17) with acute spine on 
protopod above base of mesial ramus; both 
rami slightly exceeding telson, and with 
distal margins smooth. 

Measurements (in mm).—Male holotype, 
anterior carapace length 6.1, carapace 
length 9.1, length of chela including fixed 
finger 4.1, mid-length height of chela 1.8; 
male paratype, same, 4.8, 7.8, 3.8, 1.8; fe- 
male ovigerous paratype, same, 5.4, 7.9, 
42, 1:5: 

Known range.—Known only from the 
type locality. 

Etymology.—The species is named for 


15 


Dr. Jorge Cortés, Curator of Cnidaria, Mu- 
seo de Zoologia, Escuela de Biologia, 
Universidad de Costa Rica, in recognition 
of his life-long efforts to advance knowl- 
edge of the marine fauna from Costa Rica. 

Remarks.—Upogebia cortesi, new spe- 
cies, shares with several eastern Pacific and 
western Atlantic members of the genus a 
second pereopod on which the merus bears 
no proximal mesioventral spine (see keys to 
species in Williams 1986, 1993). The ab- 
dominal sternites and pleura bear no ventral 
spinules. 

The species stands alone, however, with 
respect to several other characters. The ros- 
trum with rounded anterior margin has no 
ventral spines. The anterior gastric region 
and rostrum bear an extremely dense patch 
of setae obscuring spines on this surface. 
The pereopods are virtually spineless. This 
condition is most easily demonstrated by 
emendation of the key to species of the 
eastern Pacific (Williams 1986) in which 
the comparative relationship to other spe- 
cies in the region becomes apparent. Al- 
though the new species has spineless pereo- 
pods, it appears to be related to U. tenui- 
pollex Williams, 1986 which has a well de- 
veloped distodorsal spine on the merus of 
the cheliped and a cluster of spines on the 
merus of pereopod 3. 


Emended part of Williams (1986:7—10) 
key distinguishing U. cortesi, new species, 
and U. tenuipollex 


4. Merus of pereopod 2 lacking proximal 


MESIOVERINAL SPINE. 6). cies own dee eens os 5 

Merus of pereopod 2 bearing proximal 

TUE STOMEMELON SOHNE gg 5p Sc a ch ig sr wenn 12 
DS: Posltoctiar Spine @OSEML «6... 4s... > 6a 


Postocular spine present and well de- 
WER Cner teh reer ee Ss at oe 8 
6a. Merus of cheliped and pereopod 3 
SpINCIESS 7. PF, U. cortesi, new species 
Merus of cheliped and pereopod with 
SS). Were esa Re ee TI eal 6 
6. Merus of chelipeds bearing well devel- 
oped distodorsal spine; merus of pereo- 
pod 3 bearing cluster of proximoventral 


16 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


SHINES 24.2)! Se. eee U. tenuipollex 
Williams Merus of chelipeds lacking 
distodorsal spine or with spine tiny; 
merus of pereopod 3 bearing few ob- 


solescent proximoventral spines .... 7 


Acknowledgments 


This study was made possible, in part, by 
Short-Term Visitor grant awarded to RV by 
the Office of Fellowships and Grants, 
Smithsonian Institution, Washington, D.C. 
We thank Michael Vecchione for critical 
reading of the manuscript and Keiko Hira- 
tsuka Moore for preparing the illustrations. 
This is a contribution of the Museo de 
Zoologia, Escuela de Biologia, Universidad 
de Costa Rica. 


Literature Cited 


Leach, W. E. 1814. Crustaceology. /n Edinburgh En- 
cyclopaedia, 2nd edition, 7:383—437, plate 221. 
New York: Samuel Whiting and John L. Tiffany. 

Vargas, R., & J. Cortés. 2000. Biodiversidad marina 
de Costa Rica: Crustacea: Decapoda (Penaeo- 
idea, Sergestoidea, Caridea, Astacidea, Thalas- 
sinidea, Palinura) del Pacifico.—Revista de 
Biologia Tropical 47 (in press). 

Williams, A. B. 1986. Upogebia, from the eastern Pa- 
cific (Thalassinoidea: Upogebiidae)—Memoir 
of the San Diego Society of Natural History 14: 
1-60. 

. 1993. Mud shrimps, Upogebiidae, from the 

western Atlantic (Crustacea: Decapoda: Thal- 

assinidae).—Smithsonian Contributions to Zo- 

ology 544:1-77. 

. 1997. Two new species and a range extension 

of mud shrimps, Upogebia from Pacific Costa 

Rica and Mexico (Decapoda: Thalassinidea: 

Upogebiidae).—Proceedings of the Biological 

Society of Washington 110:617—623. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


113(1):17—23. 2000. 


Periclimenes murcielagensis, a new species of shrimp 
(Crustacea: Decapoda: Palaemonidae) living on black coral from the 
Pacific coast of Costa Rica 


Rita Vargas 


Escuela de Biologia, Universidad de Costa Rica, 2060 San José, Costa Rica 


Abstract.—Periclimenes murcielagensis, new species, a pontoniine shrimp 
living on black coral colonies, occurs at depths of 25 m at Isla San Pedrito, 
Archipiélago de las Islas Murciélago, Guanacaste, Costa Rica. This new species 
is most similar to P. veleronis, Holthuis, and P. americanus Kingsley, and is 
distinguished from these by characters from the rostrum, basal antennular seg- 
ment, antennular flagellum, and incisor process of the mandible. 


Shrimps of the genus Periclimenes Cos- 
ta, 1844, belong to the subfamily Pontoni- 
ine, and are distributed widely in subtropi- 
cal and tropical marine waters worldwide. 
Some species live freely but the majority 
are associated with other marine inverte- 
brates. Of the species of this genus known 
from the eastern Pacific, only P. infraspinis 
(Rathbun 1902), P. lucasi Chace, 1937, and 
P. veleronis Holthuis, 1951, have not been 
reported in association with other inverte- 
brates (Holthuis 1951). Known associates 
of these shrimps in the western Atlantic are 
sponges, gorgonians, actinians, corallimor- 
pharians, rhizostome scyphozoans, hy- 
droids, antipatharians, bivalves, ophiuroids 
and crinoids (Heard & Spotte 1991, Spotte 
et al. 1994). 

While conducting ecological studies of 
soft corals, specimens of an undescribed 
species of Periclimenes were found living 
on colonies of black coral (Antipathes pan- 
amensis Verrill, 1869) at Isla San Pedrito, 
Archipiélago de las Islas Murciélago, 
Guanacaste, Costa Rica. This new species 
is described herein. 

Specimens were collected during SCU- 
BA dives at 25 m. Plastic bags were placed 
over black coral colonies and closed with a 
rubber band. The shrimps and Antipathes 
panamensis colonies were fixed in a mix- 


ture of 10% formalin and seawater, the col- 
onies washed in the laboratory with fresh 
water, and all the liquid passed through a 
0.5 mm mesh. The shrimps were preserved 
in 70% ethanol. 

The material is deposited in the National 
Museum of Natural History, Smithsonian 
Institution, Washington, D.C. (USNM), and 
Museo de Zoologia, Universidad de Costa 
Rica (UCRMZ). Carapace length (CL) was 
measured from the tip of the rostrum to the 
posterodorsal margin of the carapace. 


Periclimenes murcielagensis, new species 
Figs. 1-5 


Material.—Holotype: ovigerous female 
(CL 7.75 mm), USNM 260931, Isla San 
Pedrito, Archipiélago de las Islas Murcié- 
lago, Guanacaste, Costa Rica, 25 m, coll. 
Odalisca Breedy, 24 May 1996.—Para- 
types: 4 females (CL 7.5—7.9 mm), 2 males 
(CL 5.256.5), USNM 260932; 9 females 
(CL 6.25—-8.4 mm), 2 males (CL 5.6—6.25 
mm), UCRMZ 2247-01, same collection 
data as holotype. 

Diagnosis.—Carapace armed with both 
hepatic and antennal spine; rostrum well 
developed, reaching end of antennular pe- 
duncle, usually armed with 8—10 teeth dor- 
sally and 2—4 teeth on distal half of ventral 
margin. Antennular peduncle having basal 


18 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 1. 


yes BESS 


— oe 
= ———— 
—— = —— 

eS oS Se ee as Ss ee SS 


= 


an 


e, 
2 
v 
SB 
Ss 
eee 
a 
a = oe 


Periclimenes murcielagensis, new species, holotype ovigerous female (CL 7.75 mm), lateral view 


USNM 260931. Left antennule, antenna and pereopods 1—5 not shown. Scale equals 5 mm. 


article armed with 2—4 distolateral spines. 
Pereopods 3—5 with dactylus simple. 

Description.—Rostrum (Figs. 1, 5a—f) 
nearly straight, slightly reaching end of an- 
tennular peduncle; dorsal margin armed 
with 8—10 teeth interspaced with setae, pos- 
teriormost tooth placed on carapace poste- 
rior to level of orbital margin, second tooth 
on carapace usually even with, or anterior 
to, orbital margin. Distance between first 
and second tooth, sometimes slightly larger 
than distance between other teeth, which 
are regularly spaced over rostrum; ventral 
margin slightly convex, armed with 2-4 
teeth on distal half. No postorbital ridges or 
supra orbital spines. Antennal spine well 
developed, acute. Hepatic spine well devel- 
oped, slightly larger and more robust than 
antennal spine. Lower orbital angle pro- 
duced into small, blunt lobe. 

Abdominal pleura broadly rounded. 
Sixth somite (Fig. 1) nearly twice as long 
as fifth and slightly longer than telson. Dor- 
sal spines of telson (Fig. 2c) distinct, prox- 
imal pair situated near midlength of telson, 
distal pair of spines closer to proximal pair 
than posterior margin of telson; intermedi- 
ate marginal spines at posterior end of tel- 
son not quite twice as long as mesial pair. 

Cornea as broad as peduncle (eyestalk), 
constricted at junction with eyestalk, acces- 


sory pigment spot and associated ommatid- 
ia present on dorsoproximal margin of cor- 
nea. 

Antennular peduncle (Figs. 2b, 4g—h) 
with stylocerite sharp and slender, reaching 
to about midlength of basal segment; dis- 
tolateral margin of basal segment armed 
with 2—4 spines, second and third segment 
subequal in length and width. Lateral an- 
tennular flagellum with 2 branches fused 
for about 8 joints; portion of shorter branch 
not fused consisting of about 6 joints, and 
about 0.5 times shorter than fused portion. 

Antennal scale (Fig. 2a) reaching distal 
margin of third antennular segment, more 
than 3 times as long as broad; lateral mar- 
gin nearly straight, distal tooth falling far 
short of strongly produced anteromesial an- 
gle of blade. Antennal peduncle reaching 
about to midlength of scale; basal segment 
with sharp lateral spine near base of scale. 

Mouthparts as figured (Fig. 3a—f). Man- 
dible (Fig. 3a) lacking palp; incisor process 
ending in 4 distinct teeth, distal tooth larg- 
est; molar process dentate. Maxilla 1 (Fig. 
3b) with upper endite (lacinia) possesing 
stout apical spine-setae crown, distal to, 10 
or more smaller subapical setae; endite with 
4 or more stout apical spine-setae and 6 or 
more subapical setae on each side. Maxilla 
2 (Fig. 3c) with entire endite. Maxilliped 1 


VOLUME 113, NUMBER 1 


4 


Fig. 2. Periclimenes murcielagensis, new species. Paratype male (CL 5.6 mm) UCRMZ 2247-01. a, anterior 
part of carapace; b, right antennule; c, telson; d, appendix masculina. Scale equal 2 mm (a, c), 1 mm (b), 0.25 
mm (qd). 


(Fig. 3d) with well developed exopodal fla- 
gellum (lash) possessing 4 terminal plu- 
mose spine-setae; epipod slightly bilobed; 
palp slender and laching terminal spine se- 
tae. Maxilliped 2 (Fig. 3e) possessing well 
developed exopod with terminal plumose 
setae; epipod rectangular. Maxilliped 3 
(Fig. 3f) extending for proximal 0.25 of 
scaphocerite; exopod exceeding midlength 
of proximal segment; with broad, round 
epipod. 

First pereopod (Fig. 4a) reaching end of 
antennal scale; fingers unarmed, shorter 
than palm; carpus distinctly longer than 
chela, subequal to merus. Second pereopod 
distinctly unequal. Major cheliped (Fig. 4b, 
Cc) overreaching antennal scale by approxi- 
mately length of chela; fingers armed with 
teeth, distinctly shorter than palm, carpus 
distinctly shorter than chela, subequal to 
merus; ischium and merus equal in length. 


Minor cheliped (Fig. 4d) of second pair 
overreaching antennular scale by approxi- 
mately length of fingers; fingers unarmed, 
shorter than palm; chela, carpus, merus and 
ischium subequal in length. Pereopods 3—5 
(Fig. 4e—g) nearly equal in size and shape; 
dactyls entire (not bifid), propodi with sin- 
gle spine on distal flexor margin. Third pe- 
reopod just reaching end of antennal scale; 
propodus 4 times length of dactyl, slightly 
more than twice length of carpus or ischi- 
um; merus subequal in length to propodus, 
with single distal spine on flexor margin. 
Fourth pereopod extending to distal end of 
second segment of antennular peduncle; 
propodus 4 times length of dactyl, slightly 
more than twice length of carpus or ischi- 
um; merus subequal in length to propodus. 
Fifth pereopod extending to distal end of 
second segment of antennular peduncle; 
propodus 4 times length of dactyl, subequal 


20 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Big. 3: 
dible; b, maxilla 1; c, maxilla 2; d, maxilliped 1; e, maxilliped 2; f, maxilliped 3. Scale equal 0.5 mm (a-e), 1 
mm (f). 


to merus; carpus and ischium subequal in 
length. 

Males with appendix masculina (Fig. 2d) 
armed with 3 apical spine-like setae, midle 
smaller and weakly serrate, and subapical 
spine-like setae. Eggs size ranging in max- 
imum length from 0.3 to 0.5 mm (hatching 
Stage). 

Uropodal exopods extending beyond tel- 
son for about 0.8 times length of exopod; 
with strong movable spine between disto- 
lateral tooth and blade; movable spine dis- 
tinctly longer than distolateral spine. 

Color.—Orange after preservation. 

Habitat.—Periclimenes murcielagensis, 
new species, was found living on colonies 


Fe 
> N 
cs y~ 

: 


Periclimenes murcielagensis, new species. Paratype male (CL 5.6 mm) UCRMZ 2247-01. a, man- 


of black coral Antipathes panamensis. Also 
found on the same colonies with the pon- 
toniin Waldola schmitti Holthuis, 1951, cir- 
ripeds, mollusks and polychaete worms. 
Depth: 25 m. 

Distribution.—Known only from type lo- 
cality, Archipiélago de las Islas Murciélago, 
Guanacaste, Costa Rica. 

Etymology.—The species is named for 
the type locality, Archipiélago de las Islas 
Murciélago. 

Remarks.—Adult males and females 
differ only in the size of the major cheli- 
ped of the second pereopod; in males the 
major cheliped is markedly smaller than 
in females. The major cheliped in adult 


VOLUME 113, NUMBER 1 21 


G 


é€ 


rr 


Wf 


Fig. 4. Periclimenes murcielagensis, new species. Paratype male (CL 5.6 mm), lateral view, UCRMZ 2247- 
01. a, distal portion of chela of first pereopod; b, right second major pereopod; c, enlargement of chela of mayor 
pereopod; d, distal portion of chela of minor pereopod; e, f, g, third, fourth and fifth pereopods. Scale equal 4 
mm (b), 2 mm. (e, f, g), and 1 mm (a, c, d). 


males is similar to that of immature fe- cies, is most similar to Periclimenes vele- 
males. ronis, from La Libertad, Ecuador (Holthuis 

Among the eastern Pacific species of 1951). The new species can be distin- 
Periclimenes, P. murcielagensis, new spe- guished from P. veleronis by the slender 


22 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


7 oe 4 


+ ae 


ge oe 


yf 


—_———S Ss 


Fig. 5. 


Periclimenes murcielagensis, new species. Variations 1n rostrum (a—f) and basal segment of antennular 


peduncle (g—j). (a, f, males; b, c, females; g, i, j, females; h, male). Scale equals 5 mm. 


shape and number of teeth on the dorsal (8— 
10) and ventral (2—4) margins of the ros- 
trum; the presence of two to four subequal 
spines on the distolateral margin of basal 
segment of antennular peduncle; the two 
branches of the antennular flagellum are 
fused for about eight joints (five joints in 
P. veleronis); the non-fused portion consists 
of about six joints and is about one-half 
times shorter than fused portion, whereas in 
P. veleronis the non-fused portion has three 
joints and is more than half as long as the 
fused portion. 

Periclimenes murcielagensis, new spe- 
cies, also resembles P. americanus, from 
Florida (Kinsgley 1878). The two have a 
similarly shaped and armed rostrum. The 
new species can be distinguished from P. 
americanus by the presence of two to four 


subequal spines on the distolateral margin 
of the basal segment of antennular peduncle 
(one in P. americanus); no postorbital ridge 
is present in P. murcielagensis; the two 
branches of the antennular flagellum are 
fused for about eight joints (eight to 12 
joints in P. americanus), and the non-fused 
portion consists of about six joints (three or 
four in P. americanus); the incisor process 
of the mandible ends in four distinct teeth 
in P. murcielagensis (three in P. american- 
us); the second pereopods are distinctly un- 
equal in P. murcielagensis (equal in P. 
americanus). 


Acknowledgments 


This study was possible thanks to a 
Short-Term Visitor award, granted by the 


VOLUME 113, NUMBER 1 


Office of Fellowships and Grants, Smith- 
sonian Institution, Washington, D.C. I thank 
““Area de ConservaciOn Guanacaste, Min- 
isterio del Ambiente y Energia’”’ for arrang- 
ing a visit to Islas Murciélago. I also thank 
R. Lemaitre and R. Heard for their critical 
comments on the manuscript; J. Cortés for 
his constant support, and O. Breedy for 
sending me the specimens. This is a contri- 
bution of the Museo de Zoologia, Escuela 
de Biologia, Universidad de Costa Rica. 


Literature cited 


Chace, FE A., Jr. 1937. Caridean decapod crustacea 
from the Gulf of California and the west coast 
of Lower California. Part VII. The Templeton 
Crocker Expedition.—Zoologica (New York) 
22:109-138. 

Costa, O. G. 1844. Su due nuovi generi di Crostacei 
Decapodi Macrouri Nota.—Annali delle Acca- 
demia degli Aspiranti Naturalisti, Napoli 2: 
290-291. 

Heard, R. W., & S. Spotte. 1991. Pontoniinae shrimps 
(Decapoda: Caridea: Palaemonidae) of the 
northwest Atlantic. II. Periclimenes patae new 


23 


species, a gorgonian associate from shallow reef 


areas off Turks and Caicos Islands and Florida 
Keys.—Bulletin of Marine Sciences 55(1):212 
DLT. 


Holthuis, L. B. 1951. A general revision of the Palae- 
monidae (Crustacea Decapoda Natantia) of the 
Americas. I. The subfamilies Euryrhynchinae 
and Pontoniinae.—Occasional Papers, Allan 
Hancock Foundation Publications 11:1—331. 

Kingsley, J. S. 1878. Notes on the North American 
Caridea in the Museum of the Peabody Acad- 
emy of Science at Salem, Mass.—Proceedings 
of the Academy of Natural Sciences of Phila- 
delphia 1878:89-98. 

Rathbun, M. J. 1902. Description of new decapod crus- 
taceans from the west coast of North Ameri- 
ca.—Proceeding of United States National Mu- 
seum 24:885-—905. 

Spotte, S., R. Heard, & P. Bubucis. 1994. Pontoniinae 
shrimps (Decapoda: Caridea: Palaemonidae) of 
the northwest Atlantic. IV. Periclimenes anti- 
pathophilus new species, a black coral associate 
from the Turks and Caicos Islands and eastern 
Honduras.—Bulletin of Marine Sciences 55(1): 
212-227. 

Verrill, A. E. 1869. Review of the corals and polyps 
of the west coast of America.—Transactions of 
Connecticut Academy of Science 1:377—558. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


113(1):24—29. 2000. 


A new squat lobster of the genus Munidopsis Whiteaves, 1874 
(Crustacea: Decapoda: Galatheidae) from Taiwan 


Ming-Feng Wu and Tin-Yam Chan 


Institute of Marine Biology, National Taiwan Ocean University, Keelung, Taiwan, R.O.C. 


Abstract.—A new species of the squat lobster of the genus Munidopsis Whi- 
teaves, M. formosa, is described from deep-water off the northeastern coast of 
Taiwan. The new species is most similar to M. camelus Ortmann, but differs 
in having a much broader and less curved rostrum, the posterior carapace ridge 
generally armed with a pair of submedian spines, only the chelipeds bearing 
epipods, and having a lighter coloration. The relationships of this new species 
with some other similar species are also discussed. 


There are about 150 known species of 
Munidopsis Whiteaves, 1874 (see Baba 
1988), most of which inhabit deep waters 
(to 5330 m deep) and generally have small 
eyes as well as a triangular rostrum. To 
date, only three species, namely M. anda- 
manica MacGilchrist, 1905, M. cylindro- 
phthalma (Alcock, 1894) and M. latimana 
Miyake & Baba, 1966, of this genus have 
been reported from Taiwan (Wu et al. 
1998). Recently, several specimens of an 
undescribed species of Munidopsis from 
Taiwan were collected off the northeastern 
coast at a depth of about 500 m. Careful 
examinations of these specimens showed 
that they are most similar to M. camelus 
Ortmann, 1892 from Japan, and differ from 
other members of the genus in the rostrum 
being trifid, as well as the second and the 
third abdominal tergites each armed only 
with a pair of large submedian spines. De- 
tailed comparison with M. camelus revealed 
several slight but constant differences be- 
tween the Taiwanese and Japanese material. 
The Taiwanese specimens represent a new 
species described herein. 

Specimens of the new species are depos- 
ited in the National Taiwan Ocean Univer- 
sity, Keelung (NTOU), and those used of 
M. camelus in the personal collection of 
Prof. K. Baba, Kumamoto University, Ja- 


pan (KB). The measurements given are of 
carapace length (cl) excluding rostrum. 


Munidopsis formosa, new species. 
Figs. -la,.c=e, (2a. ¢. .d.5 


Material examined.—Holotype: Taiwan, 
northeastern coast, Tai-Shi fishing port, I- 
Lan County, commercial trawlers, about 
500 m, soft bottoms, Aug 1998, 1 6, cl 
20.4 mm (NTOU-H 1998-08). 

Paratypes: Taiwan, northeastern coast, 
Tai-Shi fishing port, I-Lan County, com- 
mercial trawlers, about 500 m, soft bottoms, 
Apr 1997, 1 6, cl 27.8 mm (NTOU-P 
1997-04); 15 May 1998, 1 6, cl 25.3 mm, 
1 ovigerous 2, cl 22.5 mm (NTOU-P 1998- 
05-15); 28 Apr 1999, 1 ovigerous @, cl 
17.4 mm (NTOU-P 1999-04-28). 

Description.—Body entirely covered 
with fine short setae. Rostrum broad, about 
2.5 times as long as wide and % as long as 
carapace; more or less horizontal, with tip 
gently curving dorsad; carinate dorsally, 
with trifid tip. Carapace (Figs. la, 2a) 
slightly longer than wide; frontal margin 
with 1 spine between rostrum and antero- 
lateral spines; lateral margin feebly convex, 
armed with 4 stout spines (including an- 
terolateral spine) at distal half; 1 pair of epi- 
gastric spines present; gastric region mod- 
erately convex, bearing row of 1-3 longi- 


VOLUME 113, NUMBER 1 


tudinal spines; cervical region moderately 
excavated; cardiac region raised, bearing 
large median spine; posterior transverse 
ridge elevated and generally armed only 
with pair of large submedian spines. 

Abdomen (Fig. 1a) with second and third 
tergites each having pair of large subme- 
dian spines. Telson (Fig. 2d) subdivided 
into 10 plates; lateral margins fringed with 
dense setae (very thick in male), posterior 
margin with plumose setae. 

Eyes small, lacking dark pigments, mov- 
able, extending to about middle of rostrum. 
A large spine present between eye and an- 
tenna. Basal antennular segment (Fig. Ic) 
bearing 2 strong distolateral spines, with 
distal one slightly larger. Antennal peduncle 
(Fig. 1d) with basal segment bearing dis- 
tomesial and distolateral spines (distomesial 
one very strong and long), other segments 
spineless. 

Merus of third maxilliped (Fig. le) lon- 
ger than ischium, flexor margin bearing 3 
spines, diminishing in size anteriorly, ex- 
tensor margin armed with large distal spine. 

Third thoracic sternite narrowing poste- 
riorly, anterior margin concave; fourth tho- 
racic sternite much wider, about 3 times as 
wide as preceding (Fig. 2c). 

Chelipeds subequal, long and robust 
(massive in largest male), surface covered 
with long setae; merus long and with some 
large spines; carpus short, with spines 
mainly restricted at distal margin; palm lon- 
ger than finger and without large spines (but 
sharply granular in the largest male); fin- 
gers not perfectly gaping, inner margins 
bearing some intermeshing teeth, outer 
margins nearly straight except at tips (that 
of fixed finger becoming convex in largest 
male). 

Walking legs robust, similar, all covered 
with setae. First walking leg having merus 
with large distodorsal and distolateral 
spines, and some dorsal spines; carpus gen- 
erally armed with 2 distodorsal spines; pro- 
podus nearly straight, more than 5 times as 
long as wide; dactylus much shorter than 


25 


propodus, distally curving ventrad, ventral 
margin minutely dentate. 

Epipod present only on chelipeds. 

Eggs subspherical, about 0.5 mm in di- 
ameter. 

Coloration (Fig. 3).—Body pale orange 
to orange, with color of females generally 
deeper. Rostrum except tip, pale orange or 
whitish. Fourth or fifth abdominal segments 
to tailfan from pale orange to pale white 
posteriorly. Eyes pale orange or nearly 
whitish. Antennules same color as body but 
antennal flagella orange red. Cervical 
groove and cardiac depressions sometimes 
whitish. Ventral surface whitish except 
mouth parts and chelipeds pale orange. 

Size.—Largest male and female cl 27.8 
mm and 24.6 mm respectively. Smallest 
ovigerous female cl 17.4 mm. 

Distribution.—So far known only from 
the northeastern coast of Taiwan, at depths 
of about 500 m. 

Type locality.—Taiwan, northeastern 
coast. 

Remarks.—The present form is closely 
related to Munidopsis camelus Ortmann, 
1892 from Japan and can be readily sepa- 
rately from the other species of the genus 
by the rostrum being trifid, as well as the 
second and third abdominal tergites each 
armed with a pair of large submedian 
spines. Careful comparisons between M. 
camelas (1.6 ¢ch23.2 mm. 1-2. cl 21.5 mm, 
off Hayama, Sagami Bay, Japan, lobster 
pot, Oct 1987, H. Ikeda coll. deposited at 
Kumamoto University Faculty of Educa- 
tion; also see Miyake & Baba 1967) and the 
Taiwanese material revealed the following 
differences. The anterior three pereiopods 
bear distinct epipods in the Japanese ma- 
terial but usually only the chelipeds have 
epipods in the Taiwanese specimens. How- 
ever, in one of the Taiwanese specimens 
(NTOU-P 1997-04), a distinct epipod is 
also present on the left first walking leg. It 
seems that the presence or absence of epi- 
pod are not always consistent as previously 
thought for galatheids. Nevertheless, further 
differences between the Taiwanese and Jap- 


26 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


PU UES 


Wiad At sidatili 
\s! anes IV, Ga ara 


qu’ 


\ 
r { { 


ta 

= me ety j 

acs OO Sa aii 
= ii 

AUT si 1) 

Cty 1) galley 


i iy 
bay 
Uy 


4 iy i 
Leh 


Fig. 1. a, c-e: Munidopsis formosa, new species, d holotype cl 20.4 mm, N. E. Taiwan (NTOU-H 1998- 
08). b: M. camelus Ortmann, 1892, 3d cl 23.2 mm, Sagami Bay, Japan (KB). a, carapace and anterior abdominal 
somites, dorsal view; b, carapace; c, left basal antennular segment, ventral view; d, left antennal peduncle, 
ventral view; e, basal segments of endopod of right maxilliped, ventral view. Scale bars = 5 mm. 


VOLUME 113, NUMBER 1 27 


Fig. 2. a, c—d: Munidopsis formosa, new species, 6 holotype cl 20.4 mm, N. E. Taiwan (NTOU-H 1998- 
08). b: M. camelus Ortmann, 1892, d cl 23.2 mm, Sagami Bay, Japan (KB). a—b, carapace, lateral view; c, 
anterior part of thoracic sternum, ventral view; d, telson, dorsal view. Scale bar = 5 mm. 


anese material can be found. The posterior In the Taiwanese form, the posterior cara- 
carapace ridge always bear a large median pace ridge generally armed with a pair of 
spine which is accompanied with several submedian spines only (except in one spec- 
large lateral spines in M. camelus (Fig. 1b). imen, NTOU-P 1999-1-19, there is one 


28 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Bige 3: 
05-15). 


more small spine present between the large 
submedian spines) with the lateral parts al- 
ways devoid of spines (Fig. la). Moreover, 
the rostrum is distinctly narrower (near 4 
times as long as wide) and with the tip 
abruptly curving upward in the Japanese 
material (Fig. 2b). The rostrum in the Tai- 
wanese form, however, is rather broad (2.5 
times as long as wide) and only gently 
bending upward at tip (Fig. 2a). The col- 
oration of the Japanese material also ap- 
pears to be more reddish. Color photo- 
graphs of a live specimen collected off 
Boso Penisula (180—250 m) showed that 
the body color of the Japanese material is 


Munidopsis formosa, new species, ovigerous 2 paratype cl 22.5 mm, N. E. Taiwan (NTOU-P 1998- 


orange-red. All the above differences show 
that the Taiwanese form is distinct from M. 
camelus and it is hereby described as new. 

The present new species is also similar 
to M. regia Alcock & Anderson, 1894, and 
M. plumatisetigera Baba, 1988. However, 
M. formosa new species, can be readily dis- 
tinguished from M. plumatisetigera by the 
epipods on the chelipeds and the less spiny 
body. M. regia differs considerably from M. 
formosa in having a much narrower and 
longer rostrum, and a different spination on 
the abdomen. Moreover, Alcock (1901) 
mentioned that the color in life of M. regia 
is “‘chalky pink’’. 


VOLUME 113, NUMBER 1 


Munidopsis formosa was collected from 
about 500 m deep. This is probably the 
main reason for this species being found 
only recently. The fishing depth of local 
deep-water trawlers have extended their 
trawling depths down to 500—600 m, and 
many deep-sea animals unknown to Taiwan 
have been collected, including the present 
new species. 

Etymology.—This species is named after 
its type-locality Taiwan since it is so far 
only known from there. Formosa was the 
old name of Taiwan and is used here as a 
noun in apposition. 


Acknowledgments 


Grateful acknowledgment is extended to 
Dr. K. Baba of the Kumamoto University 
Faculty of Education for sending us on loan 
the specimens of M. camelus and providing 
us with many valuable comments; Dr. T. 
Komai of Natural History Museum and In- 
stitute, Chiba, for sending us color photo- 
graphs of M. camelus, and S. H. Wu of our 
laboratory for collecting most of the present 
specimens. This work is supported by a re- 
search grant from the National Science 
Council, Taiwan, R.O.C. 


Literature Cited 


Alcock, A. 1894. Natural history notes from H. M. 
Indian Marine Survey Steamer “Investigator,” 
commander R. F Hoskyn, R. N., commaning. 
Series II, no. 1. On the results of deep-sea 
dredging during the season 1890-91 (contin- 
ued).—Annals and Magazine of Natural Histo- 
ry, series 6, 13:321—334. 

. 1901. A descriptive catalogue of the India 

deep-sea Crustacea Decapoda, Macrura and An- 


29 


omala in the Indian Museum. Being a revised 

account of the deep-sea species collected by the 

Royal Indian Marine Survey Ship Investigator. 

Indian Museum, Calcutta. 286 pp. 

, & A. R. S. Anderson. 1894. Natural history 
notes from H. M. Indian Marines Survey 
Steamer “Investigator”, commander C. F. Old- 
ham, R. N., Commanding. Series I, No. 14. An 
account of a recent collection of deep sea Crus- 
tacea from the Bay of Bengal and Laccadive 
Sea.—Journal of Asiatic Society of Bengal, 63 
(II:3):141-185. 

Baba, K. 1988. Chirostylid and Galatheid Crustacean 
(Decapoda: Anomura) of the “‘Albatross”’ Phil- 
ippine Expedition, 1907—1910.—Researches on 
Crustacea, spec. no. 2:1—203. 

MacGilchrist, A. C. 1905. Natural history notes from 
the R. I. M.S. “Investigator’’. Capt. T. H. Hem- 
ing, R. N. (retired), commanding. Series III. no. 
6. An account of the new and some of the rarer 
decapod Crustacea obtained during the survey- 
ing seasons 1901—1904.—Annals and Magazine 
of Natural History, series 7, 15:233-268. 

Miyake, S., & K. Baba. 1966. Two new species of the 
family Galatheidae from the Tosa Bay, Japan.— 
Journal of the Faculty of Agriculture, Kyushu 
University, Fukuoka 14(1):81—88. 

Wis4 . 1967. New and rare species of the 
family Galatheidae (Crustacea, Anomura) from 
the Sagami Bay in the collection of the biolog- 
ical Laboratory, Imperial Household, Japan.— 
Journal of the Faculty of Agriculture, Kyushu 
University 14(2):213-—224. 

Ortmann, A. 1892. Die Decapoden-Krebse des Strass- 
burger Museums. IV. Die Abtheilungen Galath- 
eidea und Paguridea.—Zoologischen Jahrbuch- 
ern, Abtheilung fiir Systematik, Geographie und 
Biologie der Tiere, 6:241—326. 

Whiteaves, J. EF 1874. On the recent deep-sea dredging 
operations in the Gulf of St. Lawrence.—Amer- 
ican Journal of Science (3)7:210—219. 

Wu, M.F,.T. Y¥.Chan, & TH PB Yu. 1998.’On the Chi- 
rostylidae and Galatheidae (Crustacea: Deca- 
poda: Anomura) of Taiwan.—Annual of Taiwan 
Museum 40:75—153. [In Chinese, with English 
abstract] 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


113(1):30—38. 2000. 


A new freshwater crab of the genus Geothelphusa Stimpson, 1858 
(Crustacea: Decapoda: Brachyura: Potamidae) from Yakushima 
Island, southern Kyushu, Japan 


Hiroshi Suzuki and Tomokazu Okano 


Marine Biological Laboratory, Faculty of Fisheries, Kagoshima University, 4-50-20 Shimoarata, 
Kagoshima 890-0056, Japan 


Abstract.—A new freshwater crab, Geothelphusa marmorata, is described 
from Yakushima Island of Kagoshima Prefecture, southern Kyushu, Japan. It 
is differentiated from congeners by possession of distally narrowed eyes, choc- 
olate brown or dark red carapace with scattered black speckles in life, stout 
gastric cristae, a deep H-shaped median gastro-cardiac depression, and laterally 
curved penultimate segment of the male first gonopod with a mesially curved 
ultimate segment. This is the second species of Geothelphusa Stimpson known 
from Yakushima Island, and the twelfth species from Japan. Electrophoretic 
analysis of 15 gene loci suggests that G. marmorata, new species, G. exigua 
Suzuki & Tsuda and G. dehaani (White) are reproductively isolated. 


Freshwater crabs of the genus Geothel- 
phusa Stimpson, 1858 are distributed from 
Taiwan, through the Ryukyu Islands, to the 
Japanese mainland. Recently, a large num- 
ber of species were reported from Taiwan 
and Ryukyu Islands, and 39 species are 
now recognized in the genus (Shy et al. 
1994, Shy & Ng 1998, Tan & Liu 1998). 
In Japan, 11 species are currently known, 
1.e., G. dehaani (White, 1847), G. obtusipes 
Stimpson, 1858, G. sakamotoana (Rathbun 
1905), G. aramotoi Minei, 1973, G. tenui- 
mana (Miyake & Minei 1965), G. levicer- 
vix (Rathbun 1898), G. candidiensis Bott, 
1970, G. miyazakii (Miyake & Chiu 1965), 
G. exigua Suzuki & Tsuda, 1994, G. shok- 
itai Shy & Ng, 1998, and G. minei Shy & 
Ng, 1998 (see also de Haan 1835, Rathbun 
1904, Bott 1967, Minei 1974b, Shy et al. 
1994). Of these, G. dehaani is widely dis- 
tributed on the Japanese mainland (north of 
Honshu southward to Nakano-shima of the 
Tokara Islands, south of Kyushu). The other 
10 species are restricted to the southern 
Kyushu or the Ryukyu Islands, including 
Amami-ohshima. 


During our current study of the popula- 
tion genetics and geographic distribution of 
G. dehaani and G. exigua in southern Kyu- 
shu, unusual specimens of Geothelphusa 
species were found on Yakushima Island in 
Kagoshima Prefecture. The unusual eyes, 
coloration of body, and structure of the 
male first gonopods of these crabs indicate 
that they represent a new species that is 
herein described and illustrated. In addition, 
a genetic analysis based on electrophoresis 
is included. 


Materials and Methods 


For electrophoretic analysis, a total of 
201 specimens of G. dehaani were collect- 
ed from three populations (174 specimens 
from Kotsuki River of the Kagoshima 
mainland, 20 from Hitotsutani River of the 
Kagoshima mainland, and seven from An- 
boh River of the Yakushima Island); 40 
specimens of G. exigua were collected from 
two populations (20 specimens from Hitot- 
sutani River and 20 from Kamiharai River 
of the Kagoshima mainland); and 17 spec- 
imens of the new species were collected 


VOLUME 113, NUMBER 1 


3] 


Table 1.—List of enzymes and protein, and buffer systems used in electrophoretic analysis. CAPM 6.0; Citric 
acid-aminopropyl morpholine, pH 6.0: CAPM 7.0; Citric acid-aminopropy! morpholine, pH 7.0: and CT 8.0; 


Tris-citric acid, pH 8.0. 


Enzyme and protein Symbol 
(Abbreviation and E. C. number) for locus Buffer system 

Aspartate aminotransferase (AAT, 2.6.1.1) AAT* CAPM 6.0 

Adenylate kinase (AK, 2.7.4.3) AK* CAPM 7.0 

Glyceraldehyde-3-phosphate dehydrogenase (GAPDH, 1.2.1.12) GAPDH* CAPM 6.0 

Glucose-6-phosphate isomerase (GPI, 5.3.1.9) Grr CAPM 6.0, CAPM 7.0 

Hexokinase (HK, 2.7.1.1) HkK= CTr-s:0 

Isocitrate dehydrogenase (IDHP, 1.1.1.42) IDHP-I* CAPM 7.0 
IDHP-2* CAPM 7.0 

Lactate dehydrogenase (LDH, 1.1.1.27) IBD) « buy Cr sv 

Malate dehydrogenase (MDH, 1.1.1.37) MDH- 1* CAPM 7.0 
MDH-2* CAPM 7.0 

Malic enzyme (ME, 1.1.1.38) ME* Cr s.6 

Mannose-6-phosphate isomerase (MPI, 5.3.1.8) MPI* CT-E:0 

Phosphogluconate dehydrogenase (PGDH, 1.1.1.44) PGDH* CAPM 7.0 

Phosphoglucomutase (PGM, 5.4.2.2) PGM* CT 3.0 

General protein (PROT) PROT* CAPM 7.0 


from the Anboh River during 1997. Speci- 
mens used for electrophoretic analysis were 
stored at —35°C with a small volume of 
freshwater. Muscles were extracted and ho- 
mogenized with an equal volume of cold 
distilled water. Horizontal starch gel elec- 
trophoresis was performed for the detection 
of enzyme and protein variations (Table 1). 
Locus and gene nomenclature follows that 
of Shaklee et al. (1990). Multiple loci for a 
given enzyme were distinguished by nu- 
merals, with “‘—/*’’ representing the most 
anodally-migrating isozyme. All alleles 
studied are designated alphabetically. The 
genetic distance was calculated using Nei’s 
formula (Nei 1972). All the specimens were 
collected by the junior author. 

The holotype and a paratype are deposited 
in the Kitakyushu Museum of Natural His- 
tory, Kitakyushu (KMNH), and additional 
paratypes in the National Museum of Natu- 
ral History, Smithsonian Institution, Wash- 
ington, D.C. (USNM), and Marine Biologi- 
cal Laboratory, Faculty of Fisheries, Kago- 
shima University, Kagoshima (KUMB). 
Measurements shown in parentheses under 
*‘Material examined”’ indicate the maximum 
carapace width in millimeters. Abbreviations 
used include: M, male; F female. 


Family Potamidae Ortmann, 1896 
Genus Geothelphusa Stimpson, 1858 
Geothelphusa marmorata, new species 
Figs. 1-3, Tables 1-3 


Material examined.—River Anboh: Ar- 
akawa, 1280 m alt., 24 Oct 1998: holotype, 
M (29.4), KMNH-IvR 900005, paratype, F 
(21.1), KMNH-IvR 900006, M (23.2), F 
28.9), USNME 263857 %> 1120 m alt., $ Jul 
19975 3) MEG05., 26,6, 21.8), KUMBcr 
1053 (used for the electrophoretic analysis). 

Diagnosis.—Penultimate segment of 
male first gonopod slightly curved laterally, 
ultimate segment strongly curved mesially, 
cone-shaped, with terminal aperture. Ocular 
peduncle swollen proximally, cornea small. 
Gastric cristae stout, H-shaped median gas- 
tro-cardiac depression distinct. In life, car- 
apace and pereopods chocolate brown or 
dark red with scattered black speckles. 

Description.—Carapace much broader 
than long, smooth, devoid of setae (Fig. 
la); faint, short oblique striae on epibran- 
chial and posterolateral regions; epi- and 
uro-gastric regions distinct, former divided 
into 2 stout gastric cristae by deep median 
groove; H-shaped median gastro-cardiac 
depression deep, wide; deep transverse 


S2 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 1. 
view; c, ventral view. Male paratype (USNM 268571): d, ventral view; e, dorsal view. Scales indicate 10 mm. 


groove between cardiac and intestinal re- 
gions; cervical groove obsolete on epibran- 
chial region. Anterolateral margin of cara- 
pace cristate, lined with fine rounded gran- 
ules; epibranchial notch rudimentary. Pos- 
terior margin of epistome divided into 3 
parts by 2 deep notches (Fig. 1b), granules 
present on lower edge of epistome, absent 
medially. Lower orbital margin and groove 
between subhepatic and pterygostomian re- 
gions lined with faint granules. 


Geothelphusa marmorata, new species, male holotype (KMNH-IvR 900005): a, dorsal view; b, frontal 


Eyestalk short, proximally swollen, distally 
slender. Cornea small, slightly wider than dis- 
tal portion of ocular peduncle (Fig. la, b). 

Merus of third maxilliped broad, squar- 
ish, with deep depression (Fig. 1b). Palp 3- 
segmented, connected on inner distal angle 
of merus, tip of palp not below distal mar- 
gin of ischium. Exopod slender, longer than 
ischium, with small 5-segmented flagellum 
(Figs. lc, d, 2a; exopodal flagellum dam- 
aged in holotype). 


VOLUME 113, NUMBER 1 


33 


Fig. 2. 


Geothelphusa marmorata, new species, male paratype (USNM 268571): a, exopod of third maxil- 


liped, frontal view; b—g, male holotype (KMNH-IvR 900005): b, left mandibular palp, ventral view; c, left first 
gonopod, dorsal view; d, same, ventral view; e, left second gonopod, dorsal view; f, same, ventral view; g. tip 
of left second gonopod, dorsal view. Scales indicate | mm. 


Chelipeds asymmetrical in males, sym- 
metrical in females; movable finger of large 
cheliped strongly curved in large male (not 
so in small male) (Fig. la, c—e); palm smooth, 
outer surfaces convex (Fig. la, c, d). Carpus 
of large cheliped almost smooth, with stout 


inner tooth, below which is a low projection 
(Fig. la, e). Carpus of small cheliped without 
any projection below stout inner tooth. 

Palp of mandible 3-segmented (Fig. 2b); 
distal segment uniramous, sickle-shaped; 
median segment longer than wide, distal 


34 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


half expanded; proximal segment short, 
stout. 

Penultimate segment of adult male first 
gonopod (G 1) gently curved laterally (Fig. 
2c, d), synovial membrane short, about 3 
times as long as broad (Fig. 2c), ultimate 
segment strongly curved mesially (Fig. 2c, 
d), cone-shaped, with terminal aperture. 
Male second gonopod (G 2) slender, flat, 
weakly convex at proximal part, a small 
cup-like structure on distal one-fourth, tip 
of GZ coneave (Fis: Ze: f.1¢). GZ shorter 
than G 1, tip of G 2 not protruding from 
aperture of G 1 when coupled. 

Color in life-—Carapace and pereopods 
(Fig. la, e) chocolate brown or dark red 
with scattered black speckles. Lower part of 
palm and inmovable finger of both cheli- 
peds white in large male, movable finger 
and upper part of palm of chelipeds choc- 
olate brown. In small males and females, 
lower part of palm of both chelipeds white, 
fingers and upper part of palm dark red. 
Otherwise, no color variation observed be- 
tween sexes and sizes. 

Genetic characteristics—Among fifteen 
gene loci coding for twelve enzymes and 
one protein, allelic substitution was ob- 
served between G. marmorata, new spe- 
cies, and G. exigua at Lactate dehydroge- 
nase (LDH*), Isocitrate dehydrogenase-1 
(IDHP-1*), and Phosphogluconate dehy- 
drogenase (PGDH*) loci (Table 2), and be- 
tween G. marmorata and G. dehaani at 
PGDH* locus. Nei’s genetic distances (D) 
were calculated between all samples based 
on the fifteen loci (Table 3). The D values 
of G. dehaani populations and G. exigua 
populations were low (0.016—0.066 and 
0.037, respectively). However, the D values 
between G. marmorata and G. dehaani 
were high, ranging from 0.155 to 0.2, and 
the values between G. marmorata and G. 
exigua were higher (0.473 and 0.534). 

Etymology.—The specific name is de- 
rived from the Latin marmoratus, marbled, 
alluding to the characteristic color pattern 
of the new species. 

Remarks.—The saber-like G 1 and the 3- 


segmented mandibular palp with uniramous 
distal segment present in the new species 
are characteristics of the genus Geothelphu- 
sa (see Bott 1970). The medium-sized car- 
apace, anterolateral margin lined with small 
rounded granules and absence of an epi- 
branchial tooth ally G. marmorata with G. 
dehaani, G. exigua, G. bicolor, G. miya- 
zakii, G. candidiensis, G. ferruginea, G. 
tali, G. shokitai, and G. minei, from which 
it is distinguished by several features. 

The life color of the carapace with scat- 
tered black speckles easily distinguishes G. 
marmorata from G. dehaani, G. bicolor, G. 
miyazakii, G. ferruginea, and G. tali. The 
most definitive differences are in the struc- 
ture of the G 1. The G 1 ultimate segment 
in eight related species, except for G. exi- 
gua, is Straight or slightly curved mesially 
(Bott 1967, 1970; Minei 1973, 1974a; Su- 
zuki & Tsuda 1994, Shy et al. 1994, Shy & 
Ng 1998). The ultimate segment in G. exi- 
gua is curved laterally and tapering, and 
has a subterminal aperture. However, in G. 
marmorata, this segment is strongly curved 
mesially, ending in a papilla-like tip with a 
terminal aperture. The eyestalks in the eight 
related species are constricted medially, and 
the cornea and proximal part of the ocular 
peduncle are swollen. In G. marmorata, 
only the proximal part of the ocular pedun- 
cle is swollen, while the cornea and the dis- 
tal part of the peduncle are proportionately 
narrower as in G. exigua. 

Previous genetic studies revealed that the 
different populations of G. dehaani in Japan 
exhibit varying allele frequencies in some 
gene loci in Japan (Sugawara & Gamo 
1984, Nakajima & Masuda 1985, Aotsuka 
et al. 1995, Ikeda et al. 1998). If allelic sub- 
stitution is observed at any gene locus 
among the different morphological groups 
based on coloration, shape of G 1, and so 
on, there is a possibility of the groups ac- 
tually representing different species. In the 
previous studies, there was no allelic sub- 
stitution in any gene locus among the dif- 
ferent morphological groups. However, Ike- 
da et al. (1998) observed an allelic replace- 


VOLUME 113, NUMBER 1 35 


Table 2.—Allele frequencies at 15 loci for 3 populations of G. dehaani, 2 populations of G. exigua and | 
population of G. marmorata in Kagoshima Prefecture. 


G. dehaani G. exigua 
G. marmorata 


Locus Allele Koutsuki Hitotsutani Anboh Hitotsutani Kamiharai Anboh 
AAT* he 0.027 0.025 0.000 0.975 1.000 0.059 
*b 0.922 0.975 0.929 0.025 0.000 0.941 

Fc 0.051 0.000 0.071 0.000 0.000 0.000 

AK* ia 0.961 1.000 1.000 1.000 0.750 1.000 
al 2 0.028 0.000 0.000 0.000 0.250 0.000 

sa 0.011 0.000 0.000 0.000 0.000 0.000 

GAPDH* *q 1.000 1.000 1.000 1.000 1.000 1.000 
GPI* *q 0.253 0.025 0.000 0.600 0.200 0.059 
*b 0.726 0.475 0.000 0.000 0.275 0.647 

=C 0.018 0.050 0.000 0.400 D525 0.000 

*d 0.003 0.425 1.000 0.000 0.000 0.235 

ye 0.000 0.025 0.000 0.000 0.000 0.059 

HK* *a 0.085 0.000 0.000 0.000 0.000 0.000 
=D 0.755 1.000 0.786 0.917 0.975 0.735 

KG 0.160 0.000 0.214 0.083 0.025 0.265 

IDHP- 1* *q 0.891 1.000 1.000 0.000 0.000 1.000 
*D 0.070 0.000 0.000 0.000 0.000 0.000 

at 3 0.039 0.000 0.000 0.000 0.000 0.000 

a 7) 0.000 0.000 0.000 1.000 1.000 0.000 

IDHP-2* re) 1.000 1.000 1.000 0.050 0.000 1.000 
*b 0.000 0.000 0.000 0.950 1.000 0.000 

EDH* na 1.000 1.000 1.000 0.000 0.000 1.000 
2 0) 0.000 0.000 0.000 1.000 1.000 0.000 

MDH-1* *q 0.964 0.900 1.000 0.975 0.950 1.000 
*b 0.036 0.100 0.000 0.025 0.050 0.000 

MDH-2* a 0.994 0.975 1.000 1.000 1.000 1.000 
2) 0.006 0.025 0.000 0.000 0.000 0.000 

nC 0.000 0.000 0.000 0.000 0.000 0.000 

ME* “a 1.000 1.000 1.000 1.000 1.000 1.000 
MPI* ne 1.000 1.000 1.000 1.000 1.000 1.000 
PGDH* ta 0.000 0.000 0.000 0.000 0.000 1.000 
4 0.991 1.000 1.000 1.000 1.000 0.000 

ss 0.009 0.000 0.000 0.000 0.000 0.000 

PGM* wa 0.875 0.925 Oey | 0.925 0375 0.971 
=D 0.018 0.075 0.214 0.075 0.625 0.000 

=G 0.108 0.000 0.214 0.000 0.000 0.029 

PROT* a 1.000 1.000 1.000 0.000 0.000 0.059 
*D 0.000 0.000 0.000 1.000 1.000 0.941 


Table 3.—Genetic distance among 3 populations of G. dehaani, 2 populations of G. exigua and | population 
of G. marmorata in Kagoshima Prefecture. 


G. dehaani G. exigua 
Koutsuki Hitotsutani Anboh Hitotsutani Kamiharai 
(1) (II) (IIT) (IV) (V) 
G. dehaani (11) 0.016 
(IIT) 0.032 0.066 
G. exigua (IV) 0.469 0.462 0.519 
(V) 0.520 0.518 0.559 0.037 


G. marmorata Anboh (VI) 0.156 0.155 0.200 0.473 0.534 


36 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


130° 30 E 


ips, 3: 


130° 40 E 


30° 20° N 


Distribution and abundance of G. marmorata, new species (solid circle) and G. dehaani (White, 


1847) (white circle) in Yakushima Island, Kagoshima, Japan. Arabic numerals correspond to size of circles in 
the square at bottom left, and indicate the number of crabs captured per ten minutes by one person. Broken line 


shows 1000 m contour line. 


ment at three loci between color morphs 
representing allopatric populations. No 
morphological differences in G 1 structure 
have been reported among those popula- 
tions thus far. In this study, allelic substi- 
tution was observed at three loci among G. 
marmorata, G. dehaani, and G. exigua. In 
addition, G. marmorata, and G. dehaani are 
sympatric in Anboh River, Yakushima Is- 
land. The observed allelic substitutions, 
therefore, strongly indicate the presence of 


reproductive isolation among three species. 
The genetic distance (D values) further sug- 
gest that G. marmorata, belongs to a dif- 
ferent evolutionary lineage from the Anboh 
population of G. dehaani. 
Distribution.—The specimens of G. mar- 
morata, examined have been obtained only 
in the area above 950 m altitude on Yaku- 
shima Island, Kagoshima Prefecture (Fig. 
3). Geothelphusa marmorata, and G. de- 
haani are sympatric, having been taken to- 


VOLUME 113, NUMBER 1 


gether at some locations from 950 m to 
1350 m altitude. 


Acknowledgments 


We thank C. L. McLay of the University 
of Canterbury, and P. K. L. Ng of National 
University of Singapore for their critical 
reading of the manuscript. Thanks are also 
extended to R. Lemaitre for his valuable 
comments on the manuscript. We are also 
indebted to Y. Hiwatashi for his technical 
help with the electrophoretic analysis. 


Literature cited 


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Genetic differentiation in Japanese freshwater 
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gawa Prefecture and Tokyo.—Zoological Sci- 
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Bott, R. 1967. Potamiden aus Ost-Asien (Parapotamon 
De Man, Sinopotamon n. gen., Candidiopota- 
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. 1970. Die Susswasserkrabben von Europa, 
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Eine Revision der Potamoidea und der Parath- 
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Haan, W. de 1833-1850. Crustacea. Jn P. F von Sie- 
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legit, Notis, observationibus et adumbrationibus 
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J, L-Q + 1-55; Lugduni-Batavorum, Leiden. 

Ikeda, M., T. Suzuki, & Y. Fujio. 1998. Genetic dif- 
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with reference to the body color variation.— 
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Minei, H. 1973. Potamoid crabs of the Ryukyu Islands, 
with descriptions of five new species (Crusta- 
cea, Decapoda, Potamoidea).—Journal of the 
Faculty of Agriculture, Kyushu University 17: 
203-226. 

. 1974a. Potamoid crabs of Taiwan, with de- 

scription of one new species (Crustacea, Deca- 

poda).—Journal of the Faculty of Agriculture, 

Kyushu University 18:239-251. 

. 1974b. Studies on the freshwater crabs of 


tf 


Japan I. Genus Geothelphusa Stimpson.— 
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Miyake, S., & J. K. Chiu. 1965. A new potamonid 
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, & H. Minei. 1965. A new fresh-water crab, 
Potamon (Geothelphusa) tenuimanus sp. nov., 
from Okinawa-jima, the Ryukyu Islands.—Sci- 
ence Bulletin of the Faculty of Agriculture, 
Kyushu University 21:377—-382 (in Japanese 
with English summary). 

Nakajima, K., & T. Masuda. 1985. Identification of 
local populations of freshwater crab Geothel- 
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Les SIE 

Nei, M. 1972. Genetic distance between popula- 
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292. 


Ortmann, A. E. 1896. Das System der Decapoden- 
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fur Systematik, Geographie und Biologie der 
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Rathbun, M. J. 1898. Descriptions of three new species 
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. 1904. Les crabes d’eau douce.—Nouvelles 

Archives du Muséum d’Histoire Naturelle 6: 

225-312. 

. 1905. Les crabes d’eau douce.—Nouvelles 
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Shaklee, J. B., E W. Allendorf, D. C. Morizot, & G. 
S. Whitt. 1990. Gene nomenclature for protein- 
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Shy, J.-Y., & P K. L. Ng. 1998. On two new species 
of Geothelphusa Stimpson, 1858 (Decapoda, 
Brachyura, Potamidae) from the Ryukyu Is- 
lands, Japan.—Crustaceana 71:778—784. 

, & H.-P. Yu. 1994. Crabs of the genus 
Geothelphusa Stimpson, 1858 (Crustacea: De- 
capoda: Brachyura: Potamidae) from Taiwan, 
with descriptions of 25 new species.—Raffles 
Bulletin of Zoology 42:781—846. 

Stimpson, W. 1858. Prodromus descriptions animalium 
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um Pacificum Septentrionalem, a Republica 
Federata missa, Cadwaladaro Ringgold et Jo- 
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38 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


local populations of the Japanese freshwater 
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Biogeographic Society of Japan 39:33-37 (in 
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ings of the Biological Society of Washington 
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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


113(1):39-—47. 2000. 


A new species of the genus Neostylodactylus Hayashi & Miyake, 1968 
(Crustacea: Decapoda: Stylodactylidae) from southern Japan 


Junji Okuno and Hiroyuki Tachikawa 


Coastal Branch of Natural History Museum and Institute, Chiba, 123 Yoshio, Katsuura, Chiba 
299-5242, Japan 


Abstract.—A new species of stylodactylid shrimp, Neostylodactylus litoralis, 
is described and illustrated on the basis of four ovigerous females collected from 
sublittoral zones on the Ogasawara and Ryukyu Islands at depths of 3—10.5 m. 
This new species is readily distinguished from the five described species of 
Neostylodactylus Hayashi & Miyake in having a non-produced posterior margin 
of the telson, the noticeably elongate mesialmost pair of posterior spines on the 
posterior margin of the telson, and unarmed meri of the third to fifth pereiopods. 


Members of the caridean shrimp family 
Stylodactylidae are classified in five genera 
(Hanamura & Takeda 1996), with species 
known to occur in deep waters exceeding 
100 m (Chace 1983; Cleva 1990, 1994, 
1997). Only one stylodactylid, Neostylodac- 
tylus amarynthis (De Man, 1902), has been 
recorded in shallow waters less than 100 m 
(Kemp 1925, Chace 1983, Cleva 1990). 

While sampling the sublittoral zone (3.0— 
10.5 m) of the Ogasawara and Ryukyu Is- 
lands, southern Japan, we obtained four sty- 
lodactylid specimens referable to the genus 
Neostylodactylus Hayashi & Miyake, 1968. 
Morphologically, our specimens differed 
from the five previously described species 
of Neostylodactylus, and clearly represent a 
new species described here. 

Specimens were collected by using a hand 
net, and are deposited in the Coastal Branch 
of Natural History Museum and _ Institute, 
Chiba (CMNH). The method of measure- 
ments follows Cleva (1990). The abbrevia- 
tion CL indicates postorbital carapace length. 


Family Stylodactylidae 
Genus Neostylodactylus Hayashi & 
Miyake, 1968 
Neostylodactylus litoralis, new species 
Figs. 1—4 
Type series.—Holotype: ovig. 2 CL 
29mm (CMNH-Z2C 00071), 27°11.5'N, 


142°07.0'E, Takinoura, Ani-jima Island, 
Ogasawara Islands, 5 m, Aug 1996, coll. Y. 
Morita. Paratypes: 1 ovig. 2 CL 2.8 mm 
(CMNH-ZC 00103), 27°04.6'N, 142°07.1’E, 
Hyotan-jima Islet, NW of Chichi-jima Is- 
land, Ogasawara Islands, 8 m, May 1996, 
coll. -T. Gomi. 1 ovig 2 CL 24 mm 
(CMNH-ZC 00119), 26°13.7'N, 127°27.4’E, 
Gahi-jima Islet, Kerama Group, Ryukyu Is- 
lands, 3m, 3, Sep: 1998, coll A. Ono. 

Non-type material.—|1 2 CL~2.2 mm 
(molting) (CMNH-ZC 00137), 26°42.0’N, 
127°27.4'E, Ie-shima Island, Ryukyu Is- 
lands, 10.5 m,,14 Jun 1996, coll T. No- 
mura. 

Diagnosis.—Small sized stylodactylid 
species (known specimens CL 2.4-2.9 
mm). Carapace armed with supraorbital 
spine. Rostrum well developed, falling 
short of distal end of scaphocerite, armed 
with movable spines on both dorsal and 
ventral margins. Pleuron of third to fifth ab- 
dominal somite each armed with postero- 
lateral spine. Telson armed with 2 pairs of 
dorsal and dorsolateral spines, posterior 
margin ending roundly, armed usually with 
3 pairs of spines, mesialmost pair longest, 
with 4 plumose setae between spines. Sca- 
phocerite armed laterally with 5 or 6 mov- 
able spines. Third to fifth pereiopods with 


40 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 1. Neostylodactylus litoralis, new species. Holotype (CMNH-ZC 00071). A, left carapace with cephalic 
appendages, in lateral view. B, right first to sixth abdominal somites, in lateral view. C, telson and uropods, in 
dorsal view. D, tip of telson, in dorsal view. E, right antennular peduncle, in ventral view. FE right antenna, in 
dorsal view. Scales equal 1.5 mm (A, B), 1 mm (C, E, F), 0.5 mm (D). 


VOLUME 113, NUMBER 1 4] 


< 


ari 


NN wat 


S 


VALLE 
\ 
pe, 
ig 


Z 
C D — 
eS 
= Se ae 
| Ty 
fy, 
qfeaBi® ifs, / E 
F ~ 
a ee ae: > 


Fig. 2. Neostylodactylus litoralis, new species. Holotype (CMNH-ZC 00071). A, right mandible. B, right 
maxillule. C, right maxilla. D, right first maxilliped. E, right second maxilliped. E right third maxilliped. Scales 
equal 0.5 mm (A, B, C, D), 1 mm (E, F). 


uniunguiculate dactyli, armed posteriorly postorbital region slightly concave; infra- 
with 3 movable spines; meri unarmed. orbital margin anteriorly produced in tri- 

Description.—Carapace (Fig. 1A) _ angular process, distinctly overreaching tip 
smooth, glabrous; supraorbital spine acute, of antennal spine; hepatic depression indis- 
continuous with feeble postorbital ridge; tinct; antennal spine submarginal, directed 


42 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Table 1.—Branchial formula of Neostylodactylus 
litoralis, new species (female only). 


Maxillipeds 
I II Ill I II OE IN NY 


Pereiopods 


Pleurobranchs —_—- — — ] ] ] l ] 
Arthrobranchs — 
Podobranchs -— 
Epipods | 
Exopods ] 


] 
|. . 3 a eae 
| 


anteriorly; pterygostomian angle armed 
with spine. Rostrum (Fig. 1A) straight, 
slightly descending, 0.71—0.79 times as 
long as carapace; dorsal margin armed with 
6—12 articulated teeth, 3—9 teeth anterior to 
level of postorbital margin, more or less 
equidistant, interspace with simple long 
plumose setae, 1 tooth just above level of 
postorbital margin; dorsal carina with 2 
teeth posterior to level of postorbital mar- 
gin, and with 2 or 3 robust plumose setae; 
ventral margin armed with 1-3 articulated 
teeth. 

Abdominal somites (Fig. 1B) smooth, 
glabrous; first somite with pleuron sparsely 
fringed with simple setae; pleuron of sec- 
ond somite fringed with robust plumose se- 
tae along anterolateral and ventromesial 
margins, with simple setae ventrally; pleu- 
ron of third somite armed with small acute 
tooth posterolaterally, densely fringed with 
robust plumose setae ventrally; pleuron of 
fourth somite armed with acute tooth pos- 
terolaterally, sparsely fringed with robust 
plumose setae marginally; pleuron of fifth 
somite armed with acute tooth posterolat- 
erally, with plus 2 or 3 elongate stout setae 
bearing setules, and simple setae ventrally; 
sixth somites 0.33—0.46 times as long as 
carapace, unarmed posteroventrally. 

Telson (Fig. 1C) 0.52—0.68 times as long 
as carapace, dorsal surface armed with 2 
pairs of spines, posterior pair situated more 
laterally than anterior pair, midline with 
transverse row of long plumose setae prox- 
imally; posterior margin (Fig. 1D) feebly 
rounded, not produced in acute median pro- 
cess, armed with 3 pairs of spines (an extra 


spine on left side in holotype), mesialmost 
pair noticeably elongate, flanking 4 stout 
plumose setae. 

Eye with lightly pigmented cornea, and 
plumose setae, without ocellus; corneal di- 
ameter 0.21—0.29 times as long as carapace; 
stalk slender. 

Antennular peduncle (Fig. 1E) slender, 
slightly overreaching or falling slightly 
short of rostral apex; proximal segment 
armed with ventromesial spine and long 
spiniform seta posterior to spine, dorsodis- 
tal margin fringed with short simple setae; 
stylocerite reaching midlength of proximal 
segment, tapering distally in acute point; in- 
termediate segment armed dorsolaterally 
with 2 spiniform setae; distal segment short, 
about half length of intermediate segment. 
Upper flagellum ventrally with short setae, 
seventh article with long spiniform seta dis- 
tolaterally; seventh article of lower flagel- 
lum with long, plumose seta. 

Antenna (Fig. 1F) with scaphocerite with 
lateral margin distinctly concave, slightly 
overreaching rostral apex, 0.69—0.75 times 
as long as carapace, lateral margin armed 
with 5 or 6 acute movable spines, distolat- 
eral tooth distinctly overreaching rounded 
distal blade; carpocerite fringed with long 
simple setae distomesially; basicerite armed 
with spine distolaterally; antennal flagellum 
armed with long spiniform setae, articula- 
tions much indistinct. 

Mandible (Fig. 2A) without palp; incisor 
process well developed, slightly rounded, 
distal margin armed with 5 blunt teeth; mo- 
lar process truncated distally, with short 
Sparse setae. 

Maxillule (Fig. 2B) with feebly bilobed 
palp, inner lobe with long simple seta, 
armed dorsally with small spine proximal 
to outer lobe; upper lacinia fringed with nu- 
merous setae, distal 3 setae considerably 
longer, plumose; lower lacinia distally with 
numerous simple setae, midlength of mesial 
margin with single short seta. 

Maxilla (Fig. 2C) with palp distally 
fringed with long plumose setae; basal en- 
dite bilobed, with numerous setae on mesial 


VOLUME 113, NUMBER 1 43 


tc —— 
> 
Spe) 
= hia 
LL 


i) 
(é 
) 


6 
TY 


/ 


5M 
ii eA 


= 
ai atin eras 5) 


Fig. 3. Neostylodactylus litoralis, new species. Holotype (CMNH-ZC 00071). A, left first pereiopod. B, left 
second pereiopod. C, left third pereiopod. D, same, dactylus. Scales equal 1.5 mm (A, B), | mm (C), 0.5 mm 
(D). 


44 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 4. Neostylodactylus litoralis, new species. A, holotype (CMNH-ZC 00071), fresh specimen, lateral view 
(photo by H. Tachikawa); B, paratype (CMNH-ZC 00119), alive in aquarium (photo by A. Ono). 


margin, upper lobe slightly overreaching First maxilliped (Fig. 2D) with well de- 
level of distal margin of lower lobe; coxal veloped exopod; caridean lobe well devel- 
endite feebly rounded, with numerous setae. oped, rounded, palp slender, setose; basal 
Scaphognathite broad, rounded, marginally endite distally truncate, with 3 rows of nu- 
with numerous setae. merous setae; coxal endite distinct, with 2 


VOLUME 113, NUMBER 1 


rows of numerous setae mesially. Epipod 
bilobed. 

Second maxilliped (Fig. 2E) with well 
developed exopod; terminal 2 segments im- 
planted side by side at distal end of ante- 
penultimate segment, flexor segment slight- 
ly longer and narrower than extensor seg- 
ment, distally fringed with numerous sim- 
ple setae, flexor margin with 3 simple setae, 
extensor segment rounded, marginally with 
numerous long simple setae; antepenulti- 
mate segment protruded dorsodistally, with 
long simple setae, external and mesial mar- 
gins with long simple setae; carpal segment 
short, naked; ischiomeral segment with dor- 
sal margin sparsely with setae with setules, 
mesially with similar setae. Epipod oblong, 
with well developed podobranch. 

Third maxilliped (Fig. 2F) slender, over- 
reaching distal margin of scaphocerite by 
full length of ultimate segment, without ex- 
opod; flexor margins of distal 3 segments 
densely fringed with long setae with nu- 
merous setules and sparsely with short plu- 
mose setae; penultimate segment |.21—1.22 
times as long as ultimate segment, armed 
with row of 5-8 articulated spines; ante- 
penultimate segment with lateral row of 9- 
12 articulated spines, and | spine on dorsal 
surface. Small arthrobranch present. 

Branchial formula as indicated in Table 1. 

First pereiopod (Fig. 3A) overreaching 
distal end of scaphocerite by distal margin 
of carpus; chela with reduced palm about 
one-fifth of movable and fixed fingers, 
proximally convex, cutting edges of both 
fingers entire, without tooth, movable finger 
with flexor margin fringed with long setae 
with numerous setules, fixed finger fringed 
ventrally with similar setae; carpus 0.83-— 
0.93 times as long as carapace, dorsal mar- 
gin armed with row of spiniform setae, lat- 
eral surface armed with row of spines, ven- 
tral margin fringed with long setae with nu- 
merous setules and with a few plumose 
setae; ischiomerus armed laterally with row 
of spines, with short simple setae distola- 
terally, ventral margin fringed with long se- 


45 


tae with numerous setules and with a few 
plumose setae. 

Second pereiopod (Fig. 3B) overreaching 
distal end of scaphocerite by proximal mar- 
gin of movable finger; chela with reduced 
palm about 0.20 times as long as fingers, 
proximally convex, cutting edges of both 
fingers entire, without tooth, movable finger 
with flexor margin fringed with long setae 
with numerous setules, fixed finger fringed 
ventrally with similar setae; carpus 0.67— 
0.79 times as long as carapace, dorsal mar- 
gin armed with row of spiniform setae, lat- 
eral surface armed with row of spines, ven- 
tral margin densely fringed with long setae 
with numerous setules and with a few plu- 
mose setae; ischiomerus armed laterally 
with row of spiniform setae, distolaterally 
with spine and short plumose setae, ventral 
margin densely fringed with long setae with 
numerous setules and sparsely with long 
plumose setae. 

Third pereiopod (Fig. 3C) slightly over- 
reaching midlength of scaphocerite; dacty- 
lus (Fig. 3D) uniunguiculate, armed with 3 
movable spines posteriorly; propodus 0.75— 
0.76 times as long as carapace, 2.20—2.25 
times as long as carpus, distomesial margin 
concave, ventrally armed with short spines; 
carpus unarmed, with robust plumose setae 
distolaterally; ischiomerus unarmed, 0.76— 
0.83 times as long as carapace, 2.20—2.50 
times as long as carpus, laterally with long 
robust plumose setae, dorsodistal margin 
with short plumose setae, ventrally with 
short plumose setae. Fourth and fifth pe- 
reiopods similar to third pereiopod. Fourth 
pereiopod falling slightly short of mid- 
length of scaphocerite. Fifth pereiopod 
overreaching basicerite by full length of 
dactylus. 

Uropod (Fig. 1C) slightly overreaching 
distal margin of telson. Protopodite armed 
with strong lateral spine; exopod armed 
with acute fixed tooth distolaterally, mesi- 
ally with stout movable tooth; endopod 
with prominent lateral lobe proximally. 

Color in life (Fig. 4).—Carapace and ab- 
dominal somites reddish-violet, posterolat- 


46 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


eral part of carapace and dorsolateral parts 
of the first to fourth somites darker than 
other parts, covered with numerous fine red 
spots. Antennular peduncle transparent-red- 
dish, flagella transparent-whitish. Scapho- 
cerite transparent, antennal basicerite red- 
dish-violet. Telson, third maxilliped, ante- 
rior two pereipopods and uropods transpar- 
ent. Ambulatory pereiopods with coxae and 
basis reddish-violet with numerous red 
spots; meri, carpi, propodi and dactyli 
transparent. Pleopods transparent. 

Etymology.—litoralis, from Latin Jitus, 
the shore, in allusion to the habitat in a re- 
markably shallow depth in the vertical dis- 
tributional range of Stylodactylidae. 

Distribution.—Known only from the 
Ogasawara and Ryukyu Islands, southern 
Japan. 

Remarks.—Based on the distally acute 
stylocerite, the prominent supraorbital 
spine, and the absence of the mandibular 
palp and arthrobranchs on all pereiopods, 
the specimens clearly belong in the genus 
Neostylodactylus. 

The rostral armature does show marked 
intraspecific variations. The holotype is 
armed dorsally with six teeth anterior to the 
level of the postorbital margin. The dorsal 
margin anterior to the orbital margin has 
nine teeth in the paratype from the Ogasa- 
wara Islands (CMNH-ZC 00103), and three 
teeth in that from the Ryukyu Islands 
(CMNH-ZC 00119). 

The non-type material (CMNH-ZC 
00137) was molting when captured. 

Collectors of the present specimens ob- 
served that WN. litoralis appeared to face up- 
wards. The carapace was bent strongly 
backwards towards the abdominal somites, 
with the dorsal surface of the carapace al- 
most in contact with the somites (see Fig. 
4B). 

In recent taxonomic studies dealing with 
this genus (Chace 1983; Cleva 1990, 1994, 
1997; Komai 1997), five Neostylodactylus 
species were recognized: N. affinis Hayashi 
& Miyake, 1968; N. amarynthis (De Man, 
1902); N. hayashii Komai, 1997; N. inves- 


tigatoris (Kemp, 1925); and N. sibogae (De 
Man, 1918). 

The non-produced posterior margin of 
the telson and the unarmed meri of the am- 
bulatory pereiopods readily distinguish WN. 
litoralis from all other congenerics. In other 
congeneric species, the posterior margin of 
the telson is acutely produced mesially, and 
the meri of ambulatory pereiopods are each 
armed with one or more lateral spines (see 
De Man 1918, 1920; Kemp 1925, Hayashi 
& Miyake 1968, Chace 1983, Komai 1997). 
Moreover, Dr. R. Cleva (in litt.) kindly in- 
formed us that the long mesialmost pair of 
spines on the posterior margin of the telson 
is also a characteristic of this new species. 
Furthermore, N. litoralis differs from N. 
amarynthis, the type species of the genus, 
and N. affinis, in having the rostrum falling 
short of the distal end of the scaphocerite, 
and the telson armed with two pairs of 
spines instead of five and three respectively. 
The smooth pleura of the first and second 
abdominal somites and the lack of the spine 
on the lateral surface of the sixth abdominal 
somite also separate the present new species 
from N. amarynthis. The rostral formula 
separate N. litoralis from N. affinis. From 
N. investigatoris and N. sibogae, the present 
new species is readily distinguished by hav- 
ing an armed ventral margin of the rostrum, 
and the presence of marginal spines on the 
third to fifth abdominal somites. Neostylo- 
dactylus litoralis differs from N. hayashii 
by having a much shorter rostrum, which 
falls short of the distal end of the scapho- 
cerite; the rostral armature; the lateral mar- 
gin of the scaphocerite armed with spines; 
and the number of spines on the dorsal sur- 
face of the telson. 


Acknowledgements 


We thank the Japanese skillful divers, 
Messrs. T. Gomi, Y. Morita, T. Nomura, and 
A. Ono, for making the material available 
for this study. Mr. A. Ono kindly permitted 
us to use his color photograph. Drs. A. J. 
Bruce and R. Cleva kindly read an early 


VOLUME 113, NUMBER 1 


draft, and provided valuable comments. The 
manuscript was benefited from reviews by 
Drs. T. Komai and R. Lemaitre. 


Literature Cited 


Chace, FE A., Jr. 1983. The caridean shrimps (Crusta- 
cea: Decapoda) of the Albatross Philippine Ex- 
pedition, 1907-1910, part 1: Family Stylodac- 
tylidae.—Smithsonian Contributions to Zoolo- 
gy 381:1-21. 

Cleva, R. 1990. Crustacea Decapoda: Les genres et les 
espéces indo-ouest pacifiques de Stylodactyli- 
dae. Pp. 71-136 in A. Crosnier, ed., Résultats 
des Campagnes MUSORSTOM, 6.—Mémoires 
du Muséum National d’ Histoire Naturelle, Paris 
(A) 145. 

. 1994. Some Australian Stylodactylidae (Crus- 

tacea: Decapoda), with descriptions of two new 

species.—The Beagle, Records of the Museums 

and Art Galleries of the Northern Territory 11: 

53-64. 

. 1997. Crustacea Decapoda: Stylodactylidae 
récoltés en Indonésie, aux iles Wallis et Futuna 
et au Vanuatu (Campagnes KARUBAR, MU- 
SORSTOM 7 et 8). Données complémentaires 
sur les Stylodactylidae de Nouvelle-Calédonie. 
Pp. 385—407 in A. Crosnier & P. Bouchet, eds., 
Résultats des Campagnes MUSORSTOM, 
16.—Mémoires du Muséum National d’ Histoire 
Naturelle, Paris 172. 

De Man, J. G. 1902. Die von Herrn Professor Kiiken- 
thal im Indischen Archipel gesammelten De- 


47 


kapoden und Stomatopoden. /n W. Kiikenthal, 

Ergebnisse einer zoologischen Forschungsreise 

in den Molukken und Borneo.—Abhandlungen 

Herausgegeben von der Senckenbergischen Na- 

turforschenden Gesellschaft 25:467—929 pls. 

19-27. 

. 1918. Diagnoses of new species of macrurous 

decapod crustacea from the Siboga-Expedi- 

tion.—Zoologische Mededelingen, Leiden 4: 

159-166. 

. 1920. The Decapoda of the Siboga Expedi- 
tion, part IV. Families Pasiphaeidae, Stylodac- 
tylidae, Hoplophoridae, Nematocarcinidae, 
Thalassocaridae, Pandalidae, Psalidopodidae, 
Gnathophyllidae, Processidae, Glyphocrangon- 
idae, and Crangonidae.—Siboga-Expeditie 
39a3:1-318, pls. 1-25. 

Hanamura, Y., & M. Takeda. 1996. Establishment of a 
new genus Bathystylodactylus (Crustacea: De- 
capoda: Stylodactylidae), with description of a 
new species from northwestern Pacific.—Zoo- 
logical Science 13:929—934. 

Hayashi, K.-I., & S. Miyake. 1968. Notes on the fam- 
ily Stylodactylidae with the description of a 
new genus Neostylodactylus.—Journal of the 
Faculty of Agriculture, Kyushu University 14: 
583-611. 

Kemp, S. 1925. Notes on Crustacea Decapoda in the 
Indian Museum, XVII: On various Caridea.— 
Records of the Indian Museum 27:249-—343. 

Komai, T. 1997. A new species of the shrimp genus 
Neostylodactylus Hayashi and Miyake (Crusta- 
cea: Decapoda: Stylodactylidae) from Japan.— 
Natural History Research 4:125—133. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


113(1):48-53. 2000. 


On the male of Scutumara enodis Ng & Nakasone, 1993 
(Crustacea: Decapoda: Brachyura: Grapsidae) 


N. K. Ng and T. Komai 


(NKN) Department of Biological Sciences, National University of Singapore, Lower Kent Ridge 
Road, Singapore 119260, Republic of Singapore; (TK) Natural History Museum & Institute 
Chiba, 955-2 Aoba-cho, Chuo-ku, Chiba 260, Japan 


Abstract.—The male of the unusual grapsid, Scutumara enodis Ng & Na- 
kasone, is described for the first time, and the species is compared with its 
congeners, S. laniger (Tesch) and S. miyakei (Nakamura & Takeda). The male 
characters of S. enodis, the type species of the genus, provide further evidence 
to support the establishment of Scutumara Ng & Nakasone, for these three 
species. The male abdomen and male first gonopods of S. enodis are, however, 
distinctly different from those of S. laniger and S. miyakei. 


A male specimen of the coral reef grap- 
sid crab, Scutumara enodis Ng & Naka- 
sone, 1993, was recently collected by the 
second author in Ishigaki Island, Yaeyama 
Group, the Ryukyus. This species was orig- 
inally described from only a single subadult 
female. This study describes the male spec- 
imen of this species, and gives information 
on the male abdomen and male first pleo- 
pod (G1). Our observations provide addi- 
tional evidence in support of the establish- 
ment of the genus Scutumara Ng & Naka- 
sone, 1993. 

Measurements are given in mm in the or- 
der: carapace width by length. Specimens 
used here are deposited in the Zoological 
Reference Collection (ZRC), Department of 
Biological Sciences, National University of 
Singapore; Natural History Museum & In- 
stitute (CBM), Chiba, Japan; and National 
Museum of Natural History, Smithsonian 
Institution, Washington, D.C., U.S.A. 
(USNM). 


Family Grapsidae MacLeay, 1838 
Subfamily Varuninae Alcock, 1900 


Scutumara enodis Ng & Nakasone, 1993 
(Figse2d32) 


Scutumara enodis Ng & Nakasone, 1993: 
1, figs. 1-2. 


Material examined.—Holotype, female 
(5.8 X 5.9 mm) (ZRC.1993.1), under litto- 
ral coral sand, Kunri-Hama Beach, Sesoko 
Island, Okinawa, Ryukyus, Japan, coll. 
PK.L. Ng & Y. Nakasone, Apr 1992. Oth- 
ers—l1 male (4.9 X 5.1 mm) (CBM-ZC 
3604), beach near Fukido-gawa river- 
mouth, Ishigaki Island, Yaeyama group, Ja- 
pan, coll. T. Komai, 24 Mar 1997. 

Description of male.—Carapace slightly 
longer than broad; dorsal surface smooth, 
glabrous, without setae; regions not defined, 
strongly convex transversely and longitu- 
dinally, gastric region most convex (Fig. 
1A). Frontal margin slightly convex, entire; 
lateral lobes not visible dorsally, slightly 
deflexed downwards; lateral edges appar- 
ently confluent with supraorbital margin 
from dorsal view, but not confluent from 
frontal view; inner edges of smooth, entire 
supraorbital margins strongly deflexed 
downwards, much more than outer edges of 
front, forming slight crimp at junction be- 
tween frontal and supraorbital margins. In- 
fraorbital margin not distinctly cristate, 
slightly raised; not granulated or striated. 
Anterolateral margin slightly arcuate, sub- 
cristate, very faintly trilobed, lobes separat- 
ed by very broad, shallow clefts; external 
orbital lobe most distinct, very broad; sec- 


VOLUME 113, NUMBER 1 


ond, third lobes more like weak undula- 
tions. Posterolateral margins not sharply de- 
marcated from anterolateral margin, ap- 
pearing almost straight, distinctly converg- 
ing. Orbits small, eyes completely filling 
orbit, corneae well developed. Pterygosto- 
mial, suborbital, branchial regions smooth. 
Orbital hiatus completely filled by large 
basal anntennal segment; flagellum com- 
pletely enclosed within orbit. Antennules 
folding obliquely, fossae very large; basal 
segment large, subtriangular in shape, 
broader than long. Posterior margin of ep- 
istome weakly sinuous, with 3 distinct ridg- 
es (1 median, 2 lateral); ridges separated 
from each other by narrow gap. Endosto- 
mial region with 3 well developed longi- 
tudinal palatal ridges (1 median, 2 lateral); 
lateral palatal ridges joining inner edges of 
lateral ridges of posterior epistomial mar- 
gin. 

Third maxilliped with foliaceous merus, 
broader than long; anterolateral angle 
strongly produced, auriculiform; distal mar- 
gin distinctly bilobed, outer lobe larger, its 
base with small median cleft. Ischium lon- 
ger than broad, sulcus not discernible. 
Small, distinct rhomboidal gape formed be- 
tween inner margins of meri and ischia of 
third maxillipeds when closed. Exopod with 
obtuse, blunt inner subdistal angle, flagel- 
lum longer than width of merus (Fig. 1B). 

Chelipeds small, subequal, outer, inner 
surfaces smooth, glabrous; merus, carpus 
without spines or teeth; inner distal angle 
of carpus with broad, low, rounded lobe. 
Outer surface of chela with low but distinct 
ventral ridge running from near proximal 
part of palm to almost tip of pollex (Fig. 
1C); inner surface without setae at base of 
fingers. Fingers distinctly longer than palm; 
cutting edge of both fingers with numerous 
denticles, ending in recurved, sharp tips; no 
gape discernible when fingers closed. 

Ambulatory legs with second pair lon- 
gest (Fig. 1D). All segments smooth, with- 
out spines or setae. Dorsal and ventral mar- 
gins of merus sub-cristate, dorsal margin 


49 


with blunt subdistal tooth. Dactylus taper- 
ing to slender, acute tip (Fig. 1E). 

Male abdomen triangular (Fig. 1F); lat- 
eral margins sparsely setose; first abdomi- 
nal segment weakly arched, with very weak 
transverse ridge; second segment narrow, 
short; third segment broad, slightly swollen 
laterally but medially depressed with prox- 
imal margin broader than distal margin, lat- 
eral margins rounded; fourth segment 
broader but shorter than fifth segment; fifth 
segment with proximal and distal margins 
straight, lateral margins slightly concave; 
sixth segment quadrate, lateral margins 
trapezoidal, weakly convex, proximal mar- 
gin slightly concave, convex medially. Tel- 
son sub-triangular, lateral margins weakly 
concave, distal margin rounded. 

Lateral margins of first 2 thoracic ster- 
nites finely granulated; suture between ster- 
nites 2 and 3 slightly convex towards ab- 
domen; lateral margins of sternites 3 and 4 
sinuous, with deep, broad notch demarcat- 
ing edge of suture; median groove between 
sternites 5 and 6 narrow; space between 
Ssternites 8 very narrow throughout length 
(Fig. 2A). 

Penis located at base of eighth sternite 
(Fig. 2A). G1 relatively slender, weakly 
curving outwards, reaching to anterior mar- 
gin of fifth sternite (Fig. 2B); terminal lobe 
elongate, apparently 2-articulated, dorsally 
curved; genital opening lateral to base of 
terminal lobe; subterminal lobe chitinous, 
rounded, narrower than terminal lobe, 
densely setose (Figs. C—E). G2 short, small. 

Distribution.—Southern Japan (Okinawa 
south to Ishigaki Islands, Ryukyus). 

Remarks.—Ng & Nakasone (1993) es- 
tablished Scutumara and transferred two 
species previously placed in Pseudograp- 
sus, S. laniger (Tesch, 1918), and S. miyakei 
(Nakamura & Takeda, 1972), to this new 
genus. Although the type specimen of S. 
enodis was represented only by a subadult 
female, Ng & Nakasone (1993) argued that 
on the basis of differences on the carapace 
and ambulatory legs, S. enodis, S. laniger 
and S. miyakei are distinct enough to war- 


50 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


E 


Fig. 1. Scutumara enodis Ng & Nakasone, 1993 (male, 4.9 by 5.1 mm, CBM-ZC 3604). A, carapace; B, 
third maxilliped; C, dorsal view of left chela; D, second ambulatory leg; E, fourth ambulatory leg; K abdomen. 
Scales: A, C = 1.0 mm; B, E, F = 0.5 mm. 


VOLUME 113, NUMBER | 5] 


Fig. 2. Scutumara enodis Ng & Nakasone, 1993 (male, 4.9 by 5.1 mm, CMB-ZC 3604). A, sternum; B, natural 
position of male first gonopod; C, male first gonopod in different view; D, enlarge view of the G1; E, different 
views of the shaven enlarged distal region of Gl. Scales: A, B = 1.0 mm, C = 0.5 mm, D, E = 0.25 mm. 


D2 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


rant their placement in Scutumara. The dis- 
covery of the male S. enodis provide further 
evidence to the establishment of Scutu- 
mara. We have also found that the exopod 
of the third maxillipeds of all three species 
reaches two-thirds the length of the merus, 
while the exopod of the third maxillipeds 
in Pseudograpsus species reach to half the 
length of the merus, a character that further 
seperates these two genera. In addition, all 
known Scutumara species are ivory-cream 
white when alive, while all Pseudograpsus 
species are chestnut in color. The lateral 
margins of the sixth abdominal segment are 
generally rounded in Scutumara, however, 
in Pseudograpsus, the lateral margins are 
angular at the distal region. The sixth ab- 
dominal segment in Scutumara (length to 
width ratio between 3.5 and 5.0) is narrow 
compared to Pseudograpsus (length to 
width ratio between 1.5 to 2.0). Noteworthy 
is that the dentation on the anterolateral 
margin of the male specimen of S. enodis 
is less prominent than that of the female 
holotype. We interpret this as intraspecific 
variation. 

Scutumara enodis is separated from S. 
laniger and S. miyakei by the different 
structure of the frontal margin, the absence 
of gastric and cardiac grooves, and absence 
of setae on the inner surface of the palm 
(Ng & Nakasone 1993). Based on pub- 
lished descriptions and figures of S. laniger 
and S$. miyakei, the male abdomen and gon- 
opod of S. enodis are also different. The 
third abdominal segment is narrower in S. 
enodis (length to width ratio: ca. 4.1) than 
for its congeners (ca. 3.5 in S. laniger, ca. 
3.1 in S. miyakei). In the two species of 
Pseudograpsus examined for this character, 
the length to width ratio of the third abdom- 
inal segment ranges from ca. 3.8 (P. elon- 
gata) to ca. 4.0 (P. albus), which is narrow- 
er than S. laniger and S. miyakei, but only 
slightly broader than S. enodis. In S. enodis, 
the lateral margins of the sixth abdominal 
segment are weakly rounded, while in S. 
laniger, the posterior part of the lateral mar- 
gins are distinctly convex; in S. miyakei the 


lateral margins are even more convex. The 
sixth abdominal segment of S. enodis is 
narrower (length to width ratio: ca. 5.0) 
than that of S. laniger (ca. 3.5) and S. mi- 
yakei (ca. 3.8). The length to width ratio of 
the telson in S. enodis is 1.0, which is sim- 
ilar to S. /aniger, but is relatively longer in 
S. miyakei (ca. 1.4). Nakamura & Takeda 
(1972) commented that the G1 of S. miyak- 
ei differs from S. laniger and Pseudograp- 
sus elongata (A. Milne-Edwards, 1873) in 
having a short distal part and obtuse sub- 
terminal lobe, while the latter two have 
long beaks and small subterminal lobes 
(Nakamura & Takeda 1972: 438). This im- 
plies that the G1 of S. laniger is also longer 
and more slender than in S. enodis and S. 
miyakei. The G1 of S. enodis is character- 
ised by having an elongated tube, and nar- 
row subterminal lobe. The length of the dis- 
tal part of the G1 of Scutumara seem to be 
variable within the genus. Likewise, in the 
three species of Pseudograpsus examined 
(P. elongata, P. albus and P. crassus) for 
this character, the length of the distal part 
of the G1 varies greatly, from long (P. elon- 
gata) to short (P. albus), indicating that the 
G1 is useful character at the species level 
but not at the generic level. 

Specimens of Scutumara enodis are 
small, as are the other two known Scutu- 
mara species. The holotype female (a sub- 
adult) is only 5.8 by 5.9 mm (Ng & Na- 
kasone 1993). The smaller adult male spec- 
imen, is only 4.9 by 5.1 mm. Similarly, the 
holotype of S. laniger is only 8.3 mm in 
carapace length, and that of S. miyakei is 
3.6 mm. On the other hand, Pseudograpsus 
species are generally larger but it is not con- 
sistent. The two smallest species are P. 
elongata and P. albus: P. elongata ranges 
from 8.8 by 7.8 mm (USNM 33411) to 9.7 
by 8.6 mm (Crosnier 1965); and P. albus 
ranges from 8.2 by 7.3 mm (USNM 81732) 
to 9.5 by 8.6 mm (Crosnier 1965). P. cras- 
sus (42.6 by 36.3 mm) is a very large spe- 
cies (USNM 93152). 


VOLUME 113, NUMBER 1 


Acknowledgments 


The authors are most grateful to Dr. Peter 
K. L. Ng and Mr. Yixiong Cai for help in 
checking the manuscript. Thanks are due to 
Dr. Hiroshi Suzuki (Kagoshima University) 
kindly obtained some key references for the 
first author, and Mrs. C. M. Yang (ZRC) for 
the loan of the holotype. Thanks are due to 
the Associate Editor and the two referees 
for laboriously correcting the manuscript. 
This is contribution 34/98 from the Ecology 
and Systematics Laboratory, Department of 
Biological Sciences, National University of 
Singapore. Support from research grant RP 
950324 to Dr. Peter K. L. Ng from the Na- 
tional University of Singapore is acknowl- 
edged. 


Literature Cited 


Alcock, A. 1900. Materials for a carcinological fauna 
of India. No. 6. The Brachyura Catometopa or 
Grapsoidea.—Journal of the Asiatic Society of 
Bengal 69(2):279—456. 

Crosnier, A. 1965. Faune de Madagascar. Publiée sous 
les auspices du Gouvernement de la République 
Malgache. XVIII. Crustacés decapodés, Grap- 
sidae et Ocypodidae. O.R.S.T.O.M., 143 pp, 11 
pl. 

Latreille, P A. 1817. Gélasime, Gelasimus (Buffon). 
In Nouveau dictionnaire d’histoire naturelle, ap- 
pliquée aux arts, 4 l’agriculture, 4 1’économie 


a 


rurale et domestique, 4 la médicine, etc., edition 
2, 12:91 1-920, 

MacLeay, W. S. 1838. Illustrations of the Annulosa of 
South Africa; being a portion of the objects of 
natural history chiefly collected during an ex- 
pedition into the interior of South Africa, under 
the direction of Dr. Andrew Smith, in the years 
1834, 1835 and 1836; fitted out by “‘The Cape 
of Good Hope association for exploring Central 
Africa”. In A. Smith, Illustrations of the Zool- 
ogy of South Africa; consisting chiefly of fig- 
ures and descriptions of the objects of natural 
history chiefly collected during an expedition 
into the interior of South Africa, under the di- 
rection of Dr. Andrew Smith, in the years 1834, 
1835 and 1836; fitted out by “The Cape of 
Good Hope association for exploring Central 
Africa,” (Invertebrates). 75 pp., 4 pls., London. 

Milne-Edwards, A. 1873. Recherches sur la faune car- 
cinologique de la Nouvelle-Calédonie, Deuxie- 
me Partie-——Nouvelles Archives du Muséum 
d’Histoire naturelle (Paris), 9:155—332, pls. 4— 
18. 

Nakamura, I., & M. Takeda. 1972. A new species of 
the Grapsidae (Crustacea, Brachyura) from Bali 
Island in the collection of the National Science 
Museum, Tokyo.—Bulletin of the National Sci- 
ence Museum, Zoology, Tokyo 15(3):435—439. 

Ng, P. K. L., & Y. Nakasone. 1993. Scutumara enodis, 
a new genus and species of grapsid crab (De- 
capoda, Grapsidae) from Okinawa, Ryukyus, 
Japan.—Crustacean Research 22:1-—6. 

Tesch, J. J. 1918. The Decapoda Brachyura of the Si- 
boga Expedition. I. Hymenosomidae, Retro- 
plumidae, Ocypodidae, Grapsidae and Gecar- 
cinidae.—Siboga-Expeditie 39c, 2:1—148, pls. 
1-6. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


113(1):54-—65. 2000. 


Larval development of Cryptolithodes expansus Miers 
(Decapoda: Anomura: Lithodidae) reared in the laboratory 


Mi Hyang Kim and Sung Yun Hong 


Department of Marine Biology, Pukyong National University, Pusan 608-737, Korea 
E-mail: syhong @ dolphin.pknu.ac.kr 


Abstract.—The complete larval development of Cryptolithodes expansus 
Miers is described based on laboratory rearing. The species has four zoeal 
stages and a megalopa. The larvae are described and illustrated, and detailed 
comparisons are made with other lithodid larvae, particularly those of Cryp- 


tolithodes typicus Brandt. 


Cryptolithodes expansus Miers, 1879 is 
found along the Korean coast (personal ob- 
servation) and in Japan (Makarov 1938, 
1962) on rocky bottoms in the sublittoral 
region to a depth of about 50—60 m. The 
family Lithodidae is represented by 16 gen- 
era and 95 species (Dawson 1989), but the 
larval development of only 16 species, rep- 
resenting 9 genera, has been described 
(Konishi 1986; Haynes 1984, 1993; Koni- 
shi & Taishaku 1994). 

Sars (1890) described all zoeal stages up 
to the megalopa of Lithodes maja Linnaeus, 
1758. Subsequent studies of Lithodidae lar- 
vae have been described, at least in part, for 
15 species (Konishi 1986; Haynes 1984, 
1993; Konishi & Taishaku 1994): Crypto- 
lithodes typicus Brandt, 1848, Dermaturus 
mandtii Brandt, 1850, Hapalogaster greb- 
nitzkii Schalfeew, 1892, H. dentata (De 
Haan 1844), H. mertensii Brandt, 1850, 
Lithodes aequispina Benedict, 1895, L. an- 
tarctica Jacquinot, 1853, Lopholithodes 
manatii Brandt, 1848, Paralithodes brevi- 
pes (H. Milne Edwards & Lucas, 1841), P. 
camtschaticus (Tilesius 1815), P. platypus 
(Brandt 1850), Paralomis granulosa (Jac- 
quinot 1852), P. hystrix (De Haan 1846), 
Placetron wosnessenskii Schalfeew, 1892, 
and Rhinolithodes wonessenskii Brandt, 
1848. 

Hart (1965) described all zoeal stages, 
megalopa and crab 1 stage of Cryptolitho- 


des typicus Brandt, 1848 reared in the lab- 
oratory. This is the only species of Cryp- 
tolithodes for which larvae are known so 
far. The purpose of the present study is to 
describe complete larval stages of C. ex- 
pansus reared in the laboratory, and to com- 
pare them with those of C. typicus. 


Materials and Methods 


On 20 August 1995 a local fisherman 
caught three berried females of Cryptolitho- 
des expansus with a commercial octopus 
trap in the vicinity of Pusan (35°10'N, 
129°10’E), Korea. The ovigerous females 
were brought to the laboratory, and kept in 
a container filled with running sea-water. 
On 10 March 1996, about 250 larvae 
hatched from one of the females. Of these, 
60 were individually reared in 50 ml glass 
bottles placed in an incubator at 9.5— 
10.5°C. Some larvae were kept in 11 glass 
beakers for mass culture. 

Larvae were fed with newly hatched Ar- 
temia nauplii. Moulting and mortality were 
checked daily. After checking, the larvae 
were transferred to freshly prepared bottles 
and beakers. At each developmental stage, 
dead larvae and exuviae were fixed and pre- 
served with 3% neutralized formalin solu- 
tion. At least five specimens of each stage 
were dissected in ethylene glycol for mi- 
croscopic observation. Drawings were 
made with the aid of a drawing tube. 


VOLUME 113, NUMBER 1 


Number of Larvae 


Zoea 1 
Zoea 2 
Zoea 3 
Zoea 4 


Megalopa 


Days after Hatching 


Fig.l. 
ditions of 9.5—10.5°C and 31.80—32.65%o. 


Measurements taken were: carapace 
length (CL), from the anterior tip of the ros- 
trum to the postero-median margin of the 
carapace; total length (TL), from the ante- 
rior tip of the rostrum to the postero-median 
margin of the telson excluding telson pro- 
cesses. At least five specimens were mea- 
sured using a calibrated ocular micrometer. 


Results 


Development and duration of the lar- 
vae.—Cryptolithodes expansus passed 


Table 1.—Cryptolithodes expansus Miers. Duration 
of each larval stage reared at 9.5~10.5°C and 
31.80~32.65%o. 


Duration of larval stages (day) 


Larval stages Mean Range n 
Zoea I 4.8 3-7 47 
Zoea II 8.9 6-18 42 
Zoea III 8.2 6-17 29 
Zoea IV 14.8 13-16 18 
Megalopa 10.5 7-15 , 


Cryptolithodes expansus Miers. Survival and duration of larval stages reared under laboratory con- 


through four zoeal stages and a megalopa 
(Table 1, Fig. 1). Of the 60 larvae reared 
individually, only three molted to the me- 
galopa stage within approximately 37 days. 


Descriptions 
First Zoea. 


Size: CL = 1.6—1.8 mm (mean 1.6 mm); 
TL = 4.0—4.4 mm (mean 4.1 mm). 

Duration: 3-7 days. 

Color: dark brown color uniformly dif- 
fused over the whole carapace. 

Carapace (Fig. 2A, B): rostrum well-de- 
veloped, tapering from broad base to acute 
tip; ventral and posterior carapace margins 
with submarginal furrow and raised rim; 
postero-lateral margin smoothly rounded. 

Eyes (Fig. 2A, B): sessile. 

Abdomen (Fig. 2 A, B): narrow and slen- 
der; 5 somites plus telson; 3rd—Sth somites 
with pair of lateral spines; pair of minute 
spines on postero-dorsal margin of 2nd—Sth 
somites. 


56 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Telson (Fig. 2L): armed with 7 pairs of 
marginal processes; outermost a naked 
spine; 2nd a fine hair; 3rd—7th denticulate 
spines. 

Antennule (Fig. 2C): biramous; an en- 
dopodal bud naked, a delineated exopod 
with 10 aesthetascs. 

Antenna (Fig. 2D): biramous; scaphocer- 
ite of the protopod with 6 long plumose se- 
tae and 2 minute naked setae; endopod na- 
ked, tapering to sharp terminal point, a 
spine adjacent to the endopod. 

Mandible (Fig. 2E): stout and well armed 
with median teeth; mandibular palp present 
as a small bud. 

Maxillule (Fig. 2F): coxal endite with 7 
serrated and 1 naked spinnule; basial endite 
with 7 stout denticulate spines and 2 minute 
setae; endopod 3-segmented with 2+1+3 
setae. 

Maxilla (Fig. 2G): proximal and distal 
lobes of coxal endite with 7 and 4 setae 
respectively; proximal and distal lobes of 
basial endite with 4 setae each; endopod un- 
segmented with 3 subterminal and 5 ter- 
minal setae; scaphognathite with 10 plu- 
mose setae. 

First maxilliped (Fig. 2H): coxa naked; 
basis with 2, 2, 3, 3 setae, endopod 5-seg- 
mented with 3, 2, 1, 2, 4+1 setae; exopod- 
ite with 4 terminal plumose natatory setae. 

Second maxilliped (Fig. 21): coxa naked; 
basis with 1, 2 setae; endopod 4-segmented 
with 2, 2, 2, 4+1 setae; exopod with 4 plu- 
mose natatory setae. 

Third maxilliped (Fig. 2J): biramous; en- 
dopod unsegmented; exopod two-segment- 
ed. 

Pereiopods (Fig. 2K): elongate but not 
functional appendages; first pair subchelate. 


Second Zoea 


Size: CL = 2.1—2.2 mm (mean 2.2 mm); 
TL = 4.4-5.0 mm (mean 4.8 mm). 

Duration: 6—18 days. 

Carapace (Fig. 3A, B): slight increase in 
size, but no change in armature. 

Eyes (Fig. 3A, B): stalked and movable. 


Telson (Fig. 3L): unchanged. 

Antennule (Fig. 3C): similar to first zoea 
exopod with 9 aesthetascs of varying size. 

Antenna (Fig. 3D): exopod with 7 plu- 
mose setae and | minute naked seta. 

Mandible (Fig. 3E): some teeth added. 

Maxillule (Fig. 3F): setation of endites 
and endopod unchanged. 

Maxilla (Fig. 3G): proximal and distal 
lobes of coxal endite with 9 and 4 setae 
respectively; proximal and distal lobes of 
basial endite with 5 and 4 setae respective- 
ly; endopod unsegmented, with 3 subdter- 
minal and 5 terminal setae; scaphognathite 
with 22 plumose setae. 

First maxilliped (Fig. 3H): setation of ba- 
sis unchanged; | plumose lateral seta added 
on lIst—3rd segments of endopod; exopod 
with 8 plumose natatory setae. 

Second maxilliped (Fig. 31): setation of 
basis unchanged; 1 plumose lateral seta 
added to Ist to 3rd segments of endopod; 
exopod with 8 plumose natatory setae. 

Third maxilliped (Fig. 3J): biramous; en- 
dopod bud with 3 setae; exopod with 8 plu- 
mose natatory setae. 

Pereiopods (Fig. 3K): unchanged in ar- 
mature; slight increase in size. 

Pleopods (Fig. 3B): rudimentary unira- 
mous buds on somites 2nd—Sth. 


Third Zoea 


Size: CL = 2.2—3.2 mm (mean 2.6 mm); 
TL = 4.9-5.9 mm (mean 5.4 mm). 

Duration: 6—17 days. 

Carapace (Fig. 4A, B): similar to second 
zoea. 

Telson (Fig. 4L): unchanged. 

Antennule (Fig. 4C): exopod with 9 
aesthetascs. 

Antenna (Fig. 4D): endopod much longer 
than scaphocerite. 

Mandible (Fig. 4E): some teeth added. 

Maxillule (Fig. 4F): setation and shape 
unchanged. 

Maxilla (Fig. 4G): proximal and distal 
lobes of coxal endite with 8 and 4 setae 
respectively; basial endite unchanged; en- 


VOLUME 113, NUMBER 1 57 


Fig. 2. Cryptolithodes expansus Miers. First zoea. A, Dorsal view; B, Lateral view; C, Antennule; D, An- 
tenna; E, Mandible; K Maxillule; G, Maxilla; H, First maxilliped; I, Second maxilliped; J, Third maxilliped; K, 
Pereiopods; L, Telson. Scale bars = 0.2 mm. 


58 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 3. Cryptolithodes expansus Miers. Second zoea. A, Dorsal view; B, Lateral view; C, Antennule; D, 
Antenna; E, Mandible; K Maxillule; G, Maxilla; H, First maxilliped; I, Second maxilliped; J, Third maxilliped; 
K; Pereiopods; L, Telson. Scale bars = 0.2 mm. 


VOLUME 113, NUMBER 1 


dopod unsegmented, with 3 subterminal 
and 4 terminal setae; scaphognathite with 
24 plumose setae. 

First maxilliped (Fig. 4H): setation un- 


changed. 

Second maxilliped (Fig. 41): setation un- 
changed. 

Third maxilliped (Fig. 4J): setation un- 
changed. 


Pereiopods (Fig. 4K): unchanged. 
Pleopods (Fig. 4B): well developed but 
not functional appendages. 


Fourth Zoea 


Size: CL = 3.0—3.4 mm (mean 3.2 mm); 
TL = 6.1—6.9 mm (mean 6.4 mm). 

Duration: 13—16 days. 

Carapace (Fig. 5A, B): similar to third 
zoea. 

Telson (Fig. 5T): unchanged. 

Antennule (Fig. 5C): similar to previous 
stage. 

Antenna (Fig. 5D): endopod 2-segment- 
ed and much longer than exopod. 

Mandible (Fig. 5E): some teeth added. 

Maxillule (Fig. 5F): setation of coxal en- 
dite and endopod unchanged; basial endite 
with 8 stout denticulate spines and 2 setu- 
les. 

Maxilla (Fig. 5G): setation of coxal and 
basal endite unchanged; scaphognathite 
with 26 plumose setae. 

First maxilliped (Fig. 5H): setation un- 
changed. 

Second maxilliped (Fig. 51): setation un- 
changed. 

Third maxilliped (Fig. 5J): more devel- 
oped than in third zoea. 

Pereiopods (Fig. 5K—O): uniramous; Ist 
pair unchange; 2nd—4th with apical spine 
except on Sth. 

Pleopods (Fig. 5P—S): unchanged. 


Megalopa 


Size: CL = 2.4—2.5 mm (mean 2.5 mm); 
TL = 3.6—3.8 mm (mean 3.7 mm). 

Duration: 7—15 days. 

Color: intense scarlet. 


39 


Carapace (Fig. 6A, B): triangular in dor- 
sal view, with conical rostrum, convex in 
transverse section; cervical groove distinct; 
angle between lateral carapace and pleuron 
acute, with well-marked keel projecting lat- 
erally. 

Abdomen (Fig. 6A, B, O): 6 somites plus 
telson; 2nd—Sth segments with pleopods; 
6th segment with 3 minute hairs on the dis- 
tal margin. 

Telson (Fig. 6O): naked; more elongate 
and conical; twice as long as 6th abdominal 
segment. 

Antennule (Fig. 6C): biramous; lower ra- 
mus 2-segmented with 0, 3+3 setae; upper 
ramus 4-segmented with 0, 4, 3, 3 aesthe- 
tascs plus 3 terminal setae. 

Antenna (Fig. 6D): scale naked; flagel- 
lum 8-segmented with 0, 2, 0, 0, 4, 1, 4, 4 
setae. 

Mandible (Fig. 6E): strongly chitinized, 
smooth and not toothed as in zoeal stages, 
with broad blade-like process; palp 2-seg- 
mented. 

Maxillule (Fig. 6F): endites with minute 
spines; endopod unsegmented. 

Maxilla (Fig. 6G): setae of coxal and ba- 
sial endite reduced and tooth-like; endopod 
unsegmented; enlarged scaphognathite with 
47 plumose setae. 

First maxilliped (Fig. 6H): basis with 8 
setae; endopod unsegmented and with 5 se- 
tae; exopod with 4 terminal setae. 

Second maxilliped (Fig. 61): basis with 2 
setae; endopod 4-segmented with 0, 0, 0, 2 
setae; exopod 2-segmented with 0, 4 setae. 

Third maxilliped (Fig. 6J): basis with 1, 1 
setae; endopod 5-segmented with 4, 6, 5, 14, 
9 setae; exopod with 3 long plumose setae. 

Pereiopods (Fig. 7A—E): well developed 
and armed with spines; functional cheli- 
peds. 

Pleopods (Fig. 6K—N): present on 2nd— 
5th abdominal somites; endopod small and 
naked; exopods with 9-11 plumose nata- 
tory setae. 


Discussion 


Morphological characteristics of lithodid 
larvae have been discussed by Gurney 


60 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 4. Cryptolithodes expansus Miers. Third zoea. A, Dorsal view; B, Lateral view; C, Antennule; D, 
Antenna; E, Mandible; K Maxillule; G, Maxilla; H, First maxilliped; I, Second maxilliped; J, Third maxilliped; 
K, Pereiopods; L, Telson. Scale bars = 0.2 mm. 


(1942), MacDonald et al. (1957), Pike & to that of the Paguridae, except for the re- 
Williamson (1960), Kurata (1964), and duction or disappearance of uropods. 

Konishi (1986). They implied that larval Although the genus Cryptolithodes be- 
morphology of this family was very similar longs to the family Lithodidae, it differs 


VOLUME 113, NUMBER 1 61 


Fig. 5. Cryptolithodes expansus Miers. Fourth zoea. A, Dorsal view; B, Lateral view; C, Antennule; D, 
Antenna, D,-D,, Endopods of antenna; E, Mandible; F Maxillule; G, Maxilla; H, First maxilliped; I, Second 
maxilliped; J, Third maxilliped; K, First pereiopod; L, Second pereiopod; M, Third pereiopod; N, Fourth pe- 
reiopod; O, Fifth pereiopod; P, Pleopod of second abdominal segment; Q, Pleopod of third abdominal segment; 
R, Pleopod of fourth abdominal segment; S, Pleopod of fifth segment; T, Telson. Scale bars = 0.2 mm. 


62 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 6. Cryptolithodes expansus Miers. Megalopa. A, Dorsal view; B, Lateral view; C, Antennule; D, An- 
tenna; E, Mandible; K Maxillule; G, Maxilla; H, First maxilliped; I, Second maxilliped; J, Third maxilliped; K, 
Pleopod of second abdominal segment; L, Pleopod of third abdominal segment; M, Pleopod of fourth abdominal 
segment; N, Pleopod of fifth abdominal segment; O, Telson. Scale bars = 0.2 mm. 


VOLUME 113, NUMBER 1 63 


“i oe ; ea = 
ee 
vee Se 


Fig. 7. Cryptolithodes expansus Miers. Megalopa. A, First pereiopod; B, Second pereiopod; C, Third pe- 
reiopod; D, Fourth pereiopod; E, Fifth pereiopod. Scale bars = 0.2 mm. 


Table 2.—Comparison of morphological characters between C. typicus and C. expansus 


Species 
C. typicus C. expansus 
Characters (Hart 1965) (Present study) 
First Zoea 
Size* carapace length 1.4 mm 1.6 mm 
total length 3.0 mm 4.1 mm 
Antennule aesthetascs 8 10 
Antenna endopod sharp tip, minute subterminal tooth. _ sharp tip 
Maxillule basial endite 7 setae 9 setae 
endopod TVisetae 9 setae 
Maxilla proximal lobes 8, 6 setae 7, 4 setae 
Telson posterior margin round straight 
Second Zoea 
Antennule aesthetascs 8 2) 
Antenna endopod segmented unsegmented 
Maxillule basial endue 8 setae 9 setae 
endopod 1-1-3 setae 24715-3' setae 
Third Zoea 
Antenna endopod segmented unsegmented 
Fourth Zoea 
Antenna endopod segmented segmented 
Maxillule basial endue 9 setae 10 setae 
Megalopa 
Size* Carapace length 2.0 mm 2.5 mm 
Total length 2.9 mm 3.7 mm 
Carapace rostrum wide and flattened narrow and pointed 
Abdomen 5th segment 2 spines 3 spines 
Antenna flagellum 2+1+ 1+-04+-3+-44+2+7 setae 0+2+0+0+4+1+4+4 setae 
Third maxil- 
liped exopod 4 plumose setae 3 plumose setae 
Telson shape triangular spaculate 


* Measured from Hart (1965, fig. 1 I, fig. 3) 


64 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


morphologically from other lithodid larvae. 
The larvae of Cryptolithodes spp. resemble 
those of some Diogenidae and Coenobitidae 
in that they lack lateral carinae on the pos- 
tero-lateral margins of the carapace. Major 
morphological characteristics of lithodid 
larvae as reported by Konishi (1986) are as 
follows: ‘‘Uropods, if present, lack an en- 
dopod, even in the final zoeal stage; the 
third maxilliped has an endopod in the first 
zoeal stage; abdominal somites lack medio- 
dorsal spines; the telson without an anal 
spine.” 

Comparative morphological features of 
Cryptolithodes typicus and C. expansus are 
summarized in the Table 2. The larvae of 
C. expansus differed from those of C. typ- 
icus in body size, number of antennular 
aesthetascs, setation of the maxillule, and 
segmentation of the second maxilliped in 
the zoeal stages. 

The zoeal stages of Cryptolithodes typi- 
cus and C. expansus differ from those of 
the other lithodid larvae in the following: 
The abdomen is narrower and more slender, 
and composed of five segments plus telson 
in all the zoeal stages; uropods are absent 
in all the zoeal stages; a postero-lateral ca- 
rinae on the carapace is absent in zoeal 
stages; a mandibular palp is present at the 
first zoeal stage. 


Acknowledgements 


We are very grateful for the helpful com- 
ments of Drs. D. I. Williamson (Port Erin 
Marine laboratory, Isle of Man, England) 
and C. Sankarankutty (Universidade Fed- 
eral do Rio Grande do Norte, Natal, Brazil). 
We thank Mrs. D. N. Kim for the collection 
of the ovigerous females. We also thank 
Patsy A. McLauglin (Shannon Point Marine 
Center, Washington, U.S.A.) and two other 
anomymous reviewers for their valuable 
criticisms and suggestions in improving the 
manuscript. 


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turae, secundum classes, ordines, genera, spe- 
cies, cum characteribus, differentiis, synonymis, 
locis, (ed. 10), I:i-iv, 1-824 (Laurentii Salvii, 
Holmiae). 

MacDonald, J. D., R. B. Pike, & D. I. Williamson. 
1957. Larvae of British species of Diogenes, 
Pagurus, Anapagurus and Lithodes (Crustacea, 
Decapoda).—Proceedings of the Zoological So- 
ciety of London 128:209-—257. 

Makarov, V. V. 1938. Rakoobraznyey, vol. 10, no. 3, 
Anomura. (Crustacés Décapodes Anomures). 
Fauna SSSR, n. ser., 16:i-x + 1-324, pls. 1-5. 
Akademii Nauk SSSR, Moscow. 

. 1962. Crustacea, vol. 10, no. 3, Anomura [En- 

glish translation of Makarov, 1938].—Fauna of 

USSR, n. ser. 16:1—4 + 1-283. Israel Program 

for Scientific Translation, Jerusalem. [Published 

for the National Science Foundation and Smith- 

sonian Institution, Washington, D.C.]. 


65 


Miers, E. J. 1879. On a collection of Crustacea made 
by Capt. H. C. St. John, R. N., in the Corean 
and Japanese Seas. Part 1, Podophthalmia. With 
an appendix by Capt. H. C. St. John. On the 
method of dredging and separating the speci- 
mens, &c., with remarks on temperature, &c.— 
Proceedings of the Zoological Society of Lon- 
don 1879:18—61, pls. I-III. 

Milne Edwards, H., & H. Lucas. 1841. Description des 
Crustacés nouveaux Ou peu connus conservés 
dans la collection du Muséum.—Archives du 
Muséum national d’histoire naturelle (Paris) 2: 
461-483. 

Pike, R. B., & D. I. Williamson. 1960. Larvae of deca- 
pod Crustacea of the families Diogenidae and 
Paguridae from the Bay of Naples.—Pubblicato 
il Stazione Zoology, Napoli 31:493-—552. 

Sars, G. O. 1890. Bidrag Kundskaben om Decapod- 
ernes Forvandlinger. II: Lithodes-Eupagurus- 
Spiropagurus-Galathodes-Munida-Porcellana- 
(Nephros).—Archiv for Mathematik og Naturv- 
idenskab 13:133-—201. 

Schalfeew, P. 1892. Carcinologische Bemerkungen aus 
dem Zoologischen Museum der Kaiserlichen 
Akademie der Wissenschaften.—Bulletin de 
lV’ Académie Impériale des Sciences de Saint-Pé- 
tersbourg 35:33 1-342. 

Tilesius, W. C. 1815. De cancris camtschaticis, oniscis, 
entomostracis et cancellis marinis microscopicis 
noctilucentibus, cum tabulis IV: aenaeis et ap- 
pendice adnexo de acaris et ricinis Camtscha- 
ticis. Conventui exhibuit die 3 Februarii 
1813.—Mémoires de |’ Académie Impériale de 
Sciences de St Pétersbourg, 5:331—405. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


113(1):66—69. 2000. 


A new genus of pinnotherid crab from the Indian Ocean 
(Crustacea: Decapoda: Brachyura) 


Raymond B. Manning and Bella Galil 


(RBM) Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian 
Institution, Washington, D.C. 20560-0163, U.S.A.; (BG) Israel Oceanographic and Limnological 
Research Ltd., RO. Box 8030, Haifa 31080, Israel 


Abstract.—Abyssotheres, new genus, is recognized for Pinnotheres abyssi- 
cola Alcock & Anderson, 1899, known from a single female taken in a bivalve 
shell from a depth of 787 m off Travancore, India. In this new genus the dactyli 
of walking legs 1 and 2 are longer than those of walking legs 3 and 4. This is 
the eighth genus of pinnotherids that shares a two-segmented palp on the third 


maxilliped. 


Alcock & Anderson (1899) described 
Pinnotheres abyssicola from a single ovig- 
erous female found in a lamellibranch taken 
off Travancore, India at a depth of 430 fm 
(787 m). It is known only from the holotype 
kept in the Zoological Survey of India and 
remains the deepest recorded occurrence of 
a pinnotherid crab. 

One of us (BG) recently visited the Zoo- 
logical Survey of India (formerly the Indian 
Museum) and examined the holotype of P. 
abyssicola. It is in very poor condition, pos- 
sibly having dried out. However, she was 
able to examine the third maxilliped (Fig. 
la) and determined that the palp comprises 
only two segments. Campos (1996) re- 
viewed the pinnotherid genera with a two- 
segmented palp on the third maxilliped and 
reported that the two-segmented palp was 
found only on members of six genera. Man- 
ning (1993) added a seventh, Epulotheres. 
A combination of characters distinguishes 
P. abyssicola from all pinnotherid genera 
with a two-segmented palp. We recognize a 
new genus here for it. 

We use the following abbreviations in the 
account below: fm, fathom(s); m, meter(s); 
MXP3, third maxilliped; WL, walking leg(s). 


Abyssotheres, new genus 


Diagnosis.—Size medium, carapace 
length and width described as less than 10 


mm in adult. Carapace length and width 
subequal, front prominent, transverse, pro- 
jecting anteriorly beyond eyes. Eyes visible 
in dorsal view. MXP3 with ischium and 
merus indistinguishably fused, arched, in- 
ner margin projecting at about distal third. 
Palp 2-segmented (Fig. la), terminal seg- 
ment spatulate, shorter than preceding seg- 
ment. Chela with dactylus slightly less than 
half of propodus. Walking legs (Fig. 1b) 
slender, equal right and left; WL1-2 with 
dactyli longer than dactyli of WL3-4. Ab- 
domen unknown. 

Male.—Unknown. 

Type species.—Pinnotheres abyssicola 
Alcock & Anderson, 1899, by present des- 
ignation and monotypy. 

Etymology.—From the Latin abyss, 
depth, and the ending theres. 

Host.—A large bivalve, Acesta indica 
(Smith) (originally described in Lima) (AlI- 
cock & Anderson 1899). 

Distribution.—Known only from off the 
coast of Travancore, India, at a depth of 787 
m. This is the deepest record for a pinnoth- 
erid. 

Remarks.—Alcock & Anderson’s origi- 
nal account, based on an ovigerous female 
8 mm wide, is: “‘Carapace as long as broad, 
circular, smooth; front rather prominent, 
about one-fifth the greatest breadth of the 


VOLUME 113, NUMBER 1 


67 


es 


Eig: I. 
mim: a MXP3; b, WL. 


carapace. The whole of the eyes and eye- 
stalks and almost the whole of the orbit are 
visible in dorsal view. The eyes are well 
developed, but very pale. The dactylus of 
the external maxillipeds is styliform and is 
inserted at the end of the preceding joint. 
The lower border of the thumb is fringed 
with fine hairs. The legs are slender; the 
second and third pair are both about 1% 
times as long as the carapace, and have the 
dactylus slightly longer than it is in the oth- 
er two pairs” (Alcock & Anderson 1899: 
14). 

This account provides few diagnostic 
features. First, the carapace is round, as 
broad as long, and the front is fairly prom- 
inent. 

Second, the terminal segment of the 
MXP3 palp is articulated terminally on the 
subdistal segment. Among those Pinnoth- 
erinae with a two-segmented palp, the dac- 
tylus of the palp is articulated terminally in 
members of Orthotheres Sakai, 1969 (see 
Campos 1989, Manning 1993) and in three 
species described by Birger (1895), Pin- 
notheres glaber, P. impressus, and P. lae- 


Abyssotheres abyssicola (Alcock & Anderson, 1899). Ovigerous female holotype, carapace length 8 


vis, all presumably shore species known 
from the Pacific Ocean. In Biirgers’s species 
the dactyli of the walking legs are equally 
long. 

A third distinguishing character of A. 
abyssicola is that the dactyli of WL1-2 are 
longer than those of WL3-4. This may be 
an unique feature within the Pinnotheridae. 

A fourth characteristic feature of A. abys- 
sicola is the arched MXP3, a feature shared 
with the unrelated Limotheres nasutus Hol- 
thuis, 1975, from the Caribbean. It also is 
a commensal of a species of Lima, but oc- 
curs in shallow water. Limotheres has a 
three-segmented mandibular palp, and dif- 
fers from Abyssotheres in numerous other 
features. 

Campos (1996) studied six genera of pin- 
notherids that have a two-segmented palp 
on the MXP3: Calyptraeotheres Campos, 
1990; Dissodactylus Smith, 1870; Gem- 
motheres Campos, 1996; Ostracotheres H. 
Milne Edwards, 1853; Tunicotheres Cam- 
pos, 1996; and Xanthasia White, 1846. 
Members of Abyssotheres can be distin- 
guished from members of these genera as 


68 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


follows: the carapace lacks the sharp lateral 
margins and the dorsal sulci characteristic 
of Calyptraeotheres; the apices of WL dac- 
tyli are simple, rather than bifurcated as in 
Dissodactylus; MXP3 with the inner distal 
margin of the merus obtusely angled rather 
than curved as in Gemmotheres, Ostracoth- 
eres, and Tunicotheres; carapace lacking 
upturned lateral margins and median mush- 
room-shaped tubercle as in Xanthasia. All 
of these other genera comprise shore rather 
than slope species, and all but Ostracoth- 
eres and Xanthasia are restricted to the 
Americas. 

A seventh genus, Epulotheres, was added 
by Manning (1993), who incorrectly re- 
ported that it had a MXP3 with a three- 
segmented palp. This was corrected by 
Manning & Felder (1996) who reillustrated 
the palp, which has the usual three seg- 
ments. Nannotheres Manning & Felder, 
1996 does have a two-segmented palp. In it 
WL4 is the longest walking leg and the dac- 
tyli of the walking legs are similar and 
equal in length. 

The obtuse projection on the dorsal sur- 
face of the dactylus of the WL (Fig. 1b) 
may prove to be an unique feature of A. 
abyssicola. 

Only one other pinnotherid, Alain cros- 
nieri Manning, 1998, is known from depths 
greater than 400 m. An associate of holo- 
thurians, it was taken in depths of 399-461 
m off Indonesia. 

We are pleased to have the opportunity 
to include this report in a volume dedicated 
to our late colleague Austin B. Williams, 
whose many studies on decapods have con- 
tributed much to our knowledge of the 


group. 


Acknowledgments 


We acknowledge with thanks the coop- 
eration of the Director, Zoological Survey 
of India, Calcutta, for permission for one of 
us (BG) to examine material in the care of 
that institution. The figure was prepared by 
Lilly King Manning. We thank Ernesto 


Campos and Roy K. Kropp for their very 
helpful reviews of the manuscript. Man- 
ning’s studies of pinnotherid crabs are sup- 
ported by the Smithsonian Marine Station 
at Fort Pierce. This is contribution no. 487 
from that facility. 


Literature Cited 


Alcock, A., & A. R. S. Anderson. 1899. An account 
of the deep-sea Crustacea dredged during the 
survey season of 1897-98. Natural History 
Notes from H. M. Royal Indian Marine Survey 
Ship “Investigator,” Commander T. M. Heming, 
R. N., commanding, series 3, no. 2.—Annals 
and Magazine of Natural History, series 7, 3:1— 
27, 278-292. 

Birger, O. 1895. Ein Beitrag zur Kenntniss der Pin- 
notherinen.—Zoologische Jahrbiicher, Abthei- 
lung fiir Systematik, Geographie und Biologie 
der Thiere 8:361—390. 

Campos, E. 1989. Comments on taxonomy of the ge- 
nus Orthotheres Sakai 1969 (Crustacea, Deca- 
poda, Pinnotheridae).—Bulletin of Marine Sci- 
ence 44:1123-—1128. 

. 1990. Calyptraeotheres, a new genus of Pin- 

notheridae for the limpet crab Fabia granti 

Glassell, 1933 (Crustacea, Brachyura).—Pro- 

ceedings of the Biological Society of Washing- 

ton 106:92-101. 

. 1996. Partial revision of pinnotherid crab gen- 
era with a two-segmented palp on the third 
maxilliped (Decapoda: Brachyura).—Journal of 
Crustacean Biology 16(3):556—563. 

Holthuis, L. B. 1975. Limotheres, a new genus of pin- 
notherid crab, commensal of the bivalve Lima, 
from the Caribbean Sea.—Zoologische Mede- 
delingen 48(25):291—295. 

Manning, R. B. 1993. West African pinnotherid crabs, 
subfamily Pinnotherinae (Crustacea, Decapoda, 
Brachyura).—Bulletin du Muséum national 
d’ Histoire naturelle, Paris, série 4, 15 (A, 1-4): 
125-177. 

. 1993. Epulotheres angelae, new genus, new 

species, a pinnotherid crab from the Caribbean 

Sea (Decapoda: Pinnotheridae).—Journal of 

Crustacean Biology 13(4):801—804. 

. 1998. A new genus and species of pinnotherid 

crab (Crustacea, Decapoda, Brachyura) from In- 

donesia.—Zoosystema 20(2):357-—362. 

, & D. L. Felder. 1996. Nannotheres moorei, a 
new genus and species of minute pinnotherid 
crab from Belize, Caribbean Sea (Crustacea: 
Decapoda: Pinnotheridae).—Proceedings of the 
Biological Society of Washington 109(2):311-— 
oh We 

Milne Edwards, H. 1853. Mémoire sur la famille des 


VOLUME 113, NUMBER | 69 


Ocypodiens, suite.—Annales des Sciences Na- ican Crustacea, I.—Transactions of the Con- 

turelle, série 3, Zoologie, 20:163—228. necticut Academy of Arts and Sciences 2:1] 13— 
Sakai, T. 1969. Two new genera and twenty-two new 176. 

species of crabs from Japan.—Proceedings of the | White, A. 1846. Notes on four genera of Crustacea. 

Biological Society of Washington 82:243—280. Annals and Magazine of Natural History 18: 


Smith, S. I. 1870. Ocypodidea. Notes on North Amer- 176-178. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


113(1):70-—76. 2000. 


A new genus and species of ghost shrimp from Tobago, West Indies 
(Crustacea: Decapoda: Callianassidae) 


Richard Heard and Raymond B. Manning 


(RH) Gulf Coast Research Laboratory, East Beach Road, Ocean Springs, Mississippi 39564, 
U.S.A.; (RBM) Department of Invertebrate Zoology, National Museum of Natural History, 
Smithsonian Institution, Washington, D.C. 20560-0163, U.S.A. 


Abstract.—The new genus Pseudobiffarius is the seventh genus of Calli- 
anassinae recognized from the Americas. This monotypic genus is based on its 
type species from Tobago, West Indies, Pseudobiffarius caesari, new species. 
It differs from all members of the subfamily in having a very short, stout upper 


flagellum on the Al peduncle. 


Collections of a very small callianassid 
were made on Tobago by two individuals 
at different times, Lois Nickell in 1989 and 
Richard Heard in 1992 and 1993. The spe- 
cies was initially identified as a member of 
Biffarius Manning & Felder, 1991, but clos- 
er examination after Heard’s collections of 
1993 revealed that it represented not only a 
new species but also a new genus as well. 
The new taxa are named here. 

The types have been deposited in the Na- 
tional Museum of Natural History, Smithson- 
ian Institution, Washington, D.C. (USNM). 

Carapace length (cl) measured as post- 
orbital carapace length and total length (t1), 
measured on the midlineare in millimeters 
(mm). Other abbreviations: Al, antennule; 
A2, antenna; leg., collector; m, meter(s); 
Max1-—2, maxillae 1—2; Mxp1-3, maxilli- 
peds 1—3; P1—5, pereopods 1—5 (P1 and P2 
are chelipeds, P3—5 walking legs); Plp1—S, 
pleopods 1-5. 


Family Callianassidae Dana, 1852 
Subfamily Callianassinae Dana, 1852 
Pseudobiffarius, new genus 


Type species.—Pseudobiffarius caesari, 
new species, by present designation and 
monotypy. 

Etymology.—The generic name _ is 
formed by combining the Latin prefix pseu- 


do-, false, with the generic name Biffarius, 
alluding to the resemblance of the present 
genus to the latter. The gender is masculine. 
Diagnosis.—Size relatively small, tl of 
adults less than 30 mm. Carapace with acute 
rostral projection, lacking rostral carina or 
spine. Cornea dorsal, subterminal, disk- 
shaped, corneal elements distinct. Al pedun- 
cle longer than A2 peduncle; dorsal flagel- 
lum of Al peduncle short and stout, shorter 
than peduncle. Mxp3 without exopod, ischi- 
um-merus operculiform; inner face of ischi- 
um with cristate ridge of strong teeth; merus 
projecting slightly beyond articulation with 
carpus; last 3 segments slender. Chelipeds 
unequal in male, equal in female, with meral 
hook. Plpl uniramous in both sexes; Plp2 
absent in male, biramous in female; Plp3—5 
foliaceus, with stubby, projecting appendices 
internae in both sexes. 
Remarks.—Manning & Felder (1991) 
recognized and named three American gen- 
era in this subfamily, Biffarius, Neotrypaea, 
and Notiax, and in 1992 added a fourth, 
Gilvossius. Rodrigues & Manning (1992) 
added a fifth, Poti. Heard & Manning 
(1998) added a sixth, Necallianassa. Poti 
and the type genus of the family, Calli- 
anassa Leach, 1814, differ from the re- 
mainder of these genera by having slender 
Mxp3, which are pediform rather than op- 
erculiform; Poti also differs from all other 


VOLUME 113, NUMBER 1 


genera in the subfamily by having an in- 
complete linea thalassinica on the carapace. 
Members of Neotrypaea and Notiax are 
known only from the eastern Pacific; mem- 
bers of the other genera are known only 
from the western Atlantic. 

Members of Pseudobiffarius can be dis- 
tinguished at once from the American gen- 
era with broad Mxp3 by the short, stout dor- 
sal flagellum of Al. They further differ from 
both Neotrypaea and Gilvossius in having 
the appendix internae of the Plp3—5 embed- 
ded in the edge of the pleopod, whereas they 
are projecting in Pseudobiffarius, as in Bif- 
farius, Necallianassa and Notiax. In Notiax 
the second male pleopod is present, whereas 
it is absent in Pseudobiffarius; members of 
Notiax also have a strong rostral spine ex- 
tending almost beyond the cornea and a me- 
dian distal spine on the telson. Members of 
Biffarius lack the strong ridge of teeth pre- 
sent on the inner margin of the Mxp3 in 
members of Pseudobiffarius. 


Pseudobiffarius caesari, new species 
Figs. 1, 2, 3a—k, n—o, 4, 5a 


Material.—Tobago: Lover’s Beach, 
northwest corner of Man O’War Bay 
[11°19’N, 60°34’W], protected beach and 
Shallow reef, depth ca. 2 m, leg. R. Heard, 
sta 3, 6 Apr 1992: 1 male, cl 3.2 mm (par- 
atype, USNM 260965). 

Pirate’s Cove, east side of Man O’ War 
Bay, depth 2—3 m, leg. R. Heard, 12 Jan 
1992: 3 females, 2 non-ovigerous, cl 2.6 
and 3.8, 1 ovigerous, cl 4.1 (paratypes, 
USNM 260970). 

Buccoo Reef [11°11’N, 60°49’W], back 
reef area with sand bottom, depth ca. 2 m, 
leg. R. Heard, sta 10A (1992 sta 2), 11 Jan 
1993: 1 male, cl 4.7 mm (holotype, USNM 
260966). 

Coral Gardens, Buccoo Reef, depth 5 m, 
leg. Lois Nickell, 6 Jul 1989: 1 female, cl 
3.6 mm (paratype, USNM 260968).—Coral 
Gardens, Buccoo Reef, depth 2 m, leg. Lois 
Nickell, 2 Aug 1989: 1 female, cl 6.0 mm, 
tl 26 mm (paratype, USNM 260969). 


71 


Pigeon Point [11°10’N, 60°51’W], depth 
1 m, on sand, leg. R. Heard, 15 Jan 1993: 
1 female, cl 4.4 mm (paratype, USNM 
260967). 

Lowlands Lagoon [= Petit Trou; 
13°50'N, 61°05’W], leg. R. Heard, sta. 9, 7 
Apr 1992: 1 male, cl 5.4 mm (paratype, 
USNM 260971). 

Diagnosis.—Size very small, cl 6.0 mm 
or less and tl 30 mm or less in adults. Tel- 
son subrectangular, with small submarginal 
spines posteriorly and distinct median and 
posterolateral spines. Mxp3 ischium-merus 
operculiform, without exopod, inner surface 
of ischium with strong crest of teeth. Major 
cheliped not dimorphic in males; merus of 
both chelipeds with ventral hook. Male 
lacking Plp2. Uropods unarmed. 

Description.—Carapace (Figs. 1, 4a—c) 
smooth, with dorsal oval, without cardiac 
prominence; cervical groove distinct; linea 
thalassinica distinct, parallel to longitudinal 
axis of body; rostrum short, acute, not pro- 
duced into distinct spine; lateral frontal pro- 
jections present, low, obtuse. 

Abdominal somites (Figs. 1, 4a, d) 
smooth, somite 1 saddle-like, shortest; so- 
mite 2 longest, almost twice as long as so- 
mite 6, with small tuft of setae posterolat- 
erally, set in submarginal longitudinal 
crease; somites 3-5 subequal in length, 
each with small lateral row of setae set in 
submarginal crease; somite 6 about as long 
as telson, with median carina. Telson (Fig. 
4e, f) subrectangular, tapering posteriorly, 
slightly longer than wide, with anterior tuft 
of setae mid-dorsally and some isolated se- 
tae and small spinules dorsally, submarginal 
row of spinules present posteriorly; poste- 
rior margin setose, emarginate, with median 
spine and 2 pairs of posterolateral spines. 

Eyes (Figs. 1, 4a—c, 5a) flattened, slightly 
convex, about as long as first segment of 
Al peduncle, external margins moderately 
convergent, mesial margins parallel, tips 
pointed; cornea rounded, darkly pigmented, 
subterminal. 

Al peduncle (Figs. 4a, b, 5a) extending 
beyond A2 peduncle; segment 3 more than 


72 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Figi 
Reef. Lateral view. Scale = 1 mm. 


twice as long as segment 2, segment 1 sub- 
equal to segment 2; upper flagellum stout, 
much shorter than distal segment of pedun- 
cle. A2 peduncle (Figs. 4a, b, 5a) with seg- 
ment 4 slightly shorter than segment 3 and 
as long as segment | and 2 together; anten- 
nal scale not evident; flagellum much lon- 
ger than peduncle. 

Mandible (Fig. 3a, b) with numerous 
small teeth on incisor process; molar pro- 
cess unarmed; palp 3-segmented, segment 
3 elongate, longer than segments 1 and 2 
combined, tapering distally. Max1 (Fig. 3c) 
with broad basal endite, distal lobe almost 
forming right angle; coxal endite elongate, 
triangular; palp slender, with curved tip. 
Max2 (Fig. 3d) with protopodal endites bi- 
lobed, distalmost lobe elongate, triangular; 
palp slender; exopod broad, with crenulated 
margins. Mxp1 (Fig. 3e) with protopodal 
endites very slender, bilobed; palp very 
small but distinct, longer than wide; exopod 
bilobed, mesial margin densely setose with 
short, spatulate setae (Fig. 3f); epipod sub- 
rectangular, elongate. Mxp2 (Fig. 3g) with 


Pseudobiffarius caesari, new genus and species, male holotype (USNM 260966), cl 4.7 mm, Buccoo 


5-segmented endopod, terminal segment 
short, spatulate; exopod unsegmented, 
shorter than endopodal merus; protopod 
with small setose epipod, apparently lack- 
ing arthrobranch. Mxp3 (Fig. 3h—k) ischi- 
um-merus sub-operculiform; ischium about 
as wide as long, mesial surface with dentate 
crest of large teeth; merus more than 1.5 
times wider than long, almost 4 times wider 
than carpus and propodus, distal margin of 
outer face projecting beyond articulation 
with carpus; propodus much longer than 
wide, tapering distally, only slightly wider 
than digitiform dactylus. 

Males with very unequal chelipeds. In 
larger cheliped of male (Figs. 1, 2a, b): is- 
chium with ventral margin spinulose in 
largest of 3 males; merus with dorsal mar- 
gin spinulose in proximal fourth, smooth 
distally, posterior margin with spinules 
proximal to distally-directed hook, with dis- 
tal portion serrated; carpus distinctly longer 
than wide, smooth dorsally and ventrally, 
broadly rounded proximally; palm shorter 
than carpus, length less than height; fingers 


VOLUME 113, NUMBER 1 


73 


Fig. 2. Pseudobiffarius caesari, new genus and species. a, Male major Pl, outer aspect; b, Male major P1, 
inner aspect; c, Female P2; d, Male P2; e, Male minor P1; f, Juvenile male minor P1, setae omitted; g, Juvenile 
male major Pl; h, Male P3; i, Female P4; j, Female P5; k, Female Plp1; 1, Female Plp2. a, b, d, male holotype 
(USNM 260966), cl 4.7 mm, Buccoo Reef; c, g, h, k, 1, female paratype (USNM 260967), cl 4.4 mm, Pigeon 
Point; e, f, juvenile male paratype (USNM 260965), cl 3.2 mm, Lover’s Beach. Scale: a—j, 0.5 mm; k, 1, 1.0 mm. 


about as long as palm, gaping, tips acutely 
pointed, crossing; cutting edge of movable 
finger minutely serrated; gape conspicuous, 
base with dorsal convex lobe, edge serrate, 
above smoother, deep invagination in distal 
border of propodus. 

Smaller cheliped of male (Fig. 2d, e) and 
both chelipeds of female (Fig. 2c) similar in 
size (chelipeds symmetrical in female): is- 
chium smooth, unarmed, about as long as 
merus; merus with small, acute hook at mid- 
length of ventral margin; carpus longer than 


Table 1.—Gill formula of Pseudobiffarius caesari, 
new genus and species. 


Maxillipeds Pereopods 
vie. 3 TP a eee ee 
Pleurobranchs = 2 aap AeA LE ES. 
Arthrobranchs - — 2 aS 2 Qe 
Podobranchs ey pees ee. eee 
Epipods a aa 
Exopods ee Se 


other joints, length about 2.5 times height; 
palm less than half as long as merus, length 
and height subequal; fingers subequal, 
curved, slightly longer than palm, sharply 
pointed, with some serrations on cutting 
edge of fixed finger, tip of fingers crossing. 

P2 (Fig. 2c, d) with fingers not gaping, cut- 
ting edges straight, smooth. P3 (Fig. 2h) with 
ischium and merus of about same width, mer- 
us longer than ischium; carpus broadening 
distally; propodus wider than carpus, anterior 
and posterior margins convex, posterior mar- 
gin much longer than anterior; dactylus as 
long as wide, spatulate. P4 (Fig. 2j) lacking 
subchelar projection. PS (Fig. 21) distinctly 
chelate, fingers small, tips curved. 

Branchial formula, differing from normal 
callianassid formula in complete absence of 
arthrobranch on Mxp2, as shown in Table 1. 

Plpl of male uniramous, 2-segmented. 
Male lacking Plp2. Plp1 of female (Fig. 2k) 
uniramous, 2-segmented, proximal segment 


74 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig: 3: 


Pseudobiffarius caesari, new genus and species: a—k, n, 0, male holotype (USNM 260966), cl 4.7 


mm, Buccoo Reef. Pseudobiffarius caesari, new genus and species: h, i, female paratype (USNM 260968), cl 
3.6 mm, Buccoo Reef.—Biffarius biformis (Biffar, 1971): 1, m, male (USNM 266071), cl 5.5 mm, Fort Pierce, 
Florida (USNM 266071): a—b, Mandible; c, Max1; d, Max2, e, Mxp1; f, Marginal setae of Mxpl1, enlarged; g, 
Mxp2; h, Mxp3, inner face; i, Mxp3, outer face; j, Mxp3, inner face); 1, Mxp3, outer face; m, Mxp3, inner face; 
n, 0, Plp3 appendix interna. Upper scale, a—e, g-m = 0.5 mm; lower scale, f, n, o, = 1.0 mm. 


with subterminal and terminal patch of short 
setae; distal joint about as long as proximal, 
with a patch of small setae at midlength and 
4 spiniform apical setae. Plp2 of female 
(Fig. 21) biramous; endopod straight, with 
long terminal setae and a few setae scattered 
along its length; exopod similar to endopod 
but curved and lacking longer apical setae. 
Plp3—5 with stubby, projecting appendices 
internae (Fig. 3n, 0) in both sexes. 


Uropodal endopod (Fig. 4a, e) slightly 
longer than wide, margins densely setose; 
exopod slightly longer than and much wider 
than endopod, upper plate more than half as 
wide as lower. 

Size.—cl of males (n = 3), 3.2 to 5.4 mm; 
tl of largest male, 20 mm; cl of females (n 
= 6), 2.6 to 6.0 mm, of ovigerous female, 
4.1 mm; tl of largest female, 26 mm. 

Remarks.—Although this species superfi- 


VOLUME 113, NUMBER 1 


75 


i; 
¥, 


Uf 


4, VAT at ATPase cS 
APRN 


| 


Fig. 4. Pseudobiffarius caesari, n. gen., n. sp. a, Dorsal view (pereopods omitted); b, Carapace and anterior 
appendages, dorsal view; c, Carapace and anterior appendages, lateral view; d, Abdomen, lateral view; e, Sixth 
abdominal somite, telson, and left uropod, dorsal view; f, Telson, dorsal view, enlarged; g, Details of telson 
margin, enlarged. a—d, f—g, female paratype (USNM 260967), cl 4.4 mm, Pigeon Point; e, female paratype 
(USNM 260968), cl 3.6 mm, Buccoo Reef. Scale: a, c-e, 2 mm; b, f, 1 mm; g, 0.5 mm. 


cially resembles the type species of Biffarius, 
B. biformis (Biffar 1971), it differs in many 
features. In contrast to B. biformis, males 
have only one type of cheliped rather than 
two types, as reported by Biffar (1971) in B. 
biformis. Pseudobiffarius caesari also differs 
from B. biformis in having the A2 peduncle 
shorter than the Al peduncle, the upper Al 
flagellum much shorter, stouter, and with 9 or 
10 rather than 18 to 20 segments. In B. bi- 
formis the inner face of the ischium of Mxp3 
(Fig. 3m) lacks spinules, whereas there is a 
line of strong spinules on the inner surface of 
Mxp3 (Fig. 3h, j) in P. caesari. 


Examination of the telson under high 
magnification (Fig. 4f, g) yielded a very 
different picture of its marginal ornamen- 
tation than can be obtained under the mag- 
nification available from a dissecting micro- 
scope. In addition to being lined with short, 
plumose setae and some much longer ones, 
especially laterally, the posterior margin of 
the telson has a distinct median spine and 
a series of small, thick marginal spines and 
shorter submarginal spines, with two larger 
spines posterolaterally. 

The shore fauna of Tobago is very rich 
in species of burrowing decapods. Among 


76 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Pip... 
female paratype (USNM 260967), cl 4.4 mm, Pigeon Point; b, Biffarius biformis (Biffar 1971), male, (USNM 
266071, cl 5.5 mm, Fort Pierce, Florida. Scale = 1 mm. 


the new species discovered by us (RH) are 
another callianassid, Neocallichirus nickel- 
lae Manning, 1993 and the commensal pin- 
notherid crab, Austinixa hardyi Heard & 
Manning, 1997. 

We are pleased to acknowledge our re- 
spect for our late colleague, Austin B. Wil- 
liams, by being able pubiish this report in 
a volume of the Proceedings of the Biolog- 
ical Society of Washington dedicated to 
him. 

Etymology.—Named for Errol Caesar of 
the Fisheries Division, Ministry of Agri- 
culture, Land, and Marine Resources, To- 
bago. His enthusiastic support and interest 
in Heard’s fieldwork on Tobago materially 
improved collecting opportunities. 


Acknowledgments 


We thank both Errol Caesar and J. David 
Hardy for their support of Heard’s investi- 
gations on Tobago; Lois Nickell, then with 
the University Marine Biological Station, 
Miullport, Scotland, for the gift of her col- 
lections; and Lilly King Manning for pre- 
paring the figures. Manning’s studies on 
callianassid systematics are supported by 
the Smithsonian Marine Station at Fort 
Pierce; this is contribution no. 455 from 
that facility. 


Carapace and anterior appendages in lateral view. a, Pseudobiffarius caesari, new genus and species, 


Literature Cited 


Biffar, T. A. 1971. New species of Callianassa (De- 
capoda, Thalassinidea) from the western Atlan- 
tic.—Crustaceana 21:225-—236. 

Heard, R. W., & R. B. Manning. 1997. Austinixa, a 
new genus of pinnotherid crab (Crustacea: De- 
capoda: Brachyura), with the description of A. 
hardyi, a new species from Tobago, West In- 
dies.—Proceedings of the Biological Society of 
Washington 110:393-398. 

,& . 1998. A new genus and species of 
ghost shrimp (Crustacea: Decapoda: Callianas- 
sidae) from the Atlantic Ocean.—Proceedings 
of the Biological Society of Washington 111(4): 
883-888. 

Leach, W. E. 1814. Crustaceology. /n D. Brewster, ed., 
Edinburgh Encyclopaedia 7(2):385—437, Edin- 
burgh. 

Manning, R. B. 1993. Two new species of Neocalli- 
chirus from the Caribbean Sea (Crustacea: De- 
capoda: Callianassidae).—Proceedings of the 
Biological Society of Washington 106:106—114. 

, & D. L. Felder. 1991. Revision of the Amer- 

ican Callianassidae (Crustacea: Decapoda: 

Thalassinidea).—Proceedings of the Biological 

Society of Washington 104:762—790. 

sates . 1992. Gilvossius, a new genus of 
callianassid shrimp from the eastern United 
States (Crustacea: Decapoda: Thalassinidea).— 
Bulletin of Marine Science 49(1—2)[for 1991]: 
558-561. 

Rodrigues, S. de A., & R. B. Manning. 1992. Poti 
gaucho, a new genus and species of ghost 
shrimp from southern Brazil (Crustacea: Deca- 
poda: Callianassidae).—Bulletin of Marine Sci- 
ence 51:9—13. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


113(1):77-87. 2000. 


Griceus buskeyi, a new genus and species of calanoid copepod 
(Crustacea) from benthopelagic waters off Hawaii 


Frank D. Ferrari and E. L. Markhaseva 


(FDF) Department of Invertebrate Zoology; National Museum of Natural History; Smithsonian 
Institution; Washington, D.C. 20560-0534, U.S.A.; (ELM) Academy of Sciences, Zoological 
Institute; Universitetskaya nab. 1; 199034 St. Petersburg, Russia 


Abstract.—The new genus and species of deep-sea, benthopelagic calanoid 
copepod, Griceus buskeyi, differs from other calanoid copepods in the follow- 
ing derived character states: dorsal interlocking extensions of abdominal so- 
mites; antenna 2, mandible, and maxilla 1 originate posterior to the labrum and 
paragnaths; coxal endite of the mandible outside of the labral-paragnathal en- 
velope; praecoxal and coxal endites of maxilla 1 unarmed; praecoxal endites 
of maxilla 2 unarmed; endopod of maxilliped with four articulating segments. 
Two setae on middle endopodal segment of swimming legs 3 and 4 suggests 
the new genus and species belongs to the superfamily Augaptiloidea. A man- 
dibular endopod which is reduced in size to a small unarmed lobe and absence 
of an endopod on maxilla | suggests an affinity with the arietellid genus Par- 


augaptilus Wolfenden. 


Samples from deep-sea, benthopelagic 
habitats continue to reveal a diverse fauna 
of calanoid copepods which often express 
unusual morphologies. Here we describe an 
unusual species of calanoid which we be- 
lieve can be placed in the family Arietelli- 
dae as it is presently understood (Ohtsuka 
et al. 1994), but not in any of its known 
genera. 


Methods 


This adult female calanoid copepod was 
collected 6 July 1997 from seawater flow- 
ing from a flexible plastic pipe maintained 
by Natural Energy Laboratory of Hawaii 
Authority near Kona, Island of Hawaii, 
19°43'27.01"N, 156°04'35.46”"W. The intake 
of the pipe (1 m diameter) is located at 675 
m, about 30 m from the bottom, and draws 
in about 132 cubic m/min. Because of its 
flexibility, the position of the intake may 
vary vertically + 10 m with the movement 
of the tide. Water from one outflow pipe (10 


cm in diameter) drains through a 53 micron 
mesh net placed in a large, dark tank. The 
sample was collected after 12 hrs; the ani- 
mal reported here may have been dead prior 
to sample fixation with 4% formaldehyde. 
The specimen was later cleared in steps 
through 50% lactic acid/50% water to 
100% lactic acid, stained by adding a so- 
lution of chlorazol black E dissolved in 
70% ethanol/30% water, and examined with 
bright-field and with differential interfer- 
ence optics. 

Cephalic appendages are abbreviated Al 
= antenna 1; A2 = antenna 2; Mn = man- 
dible; Mx1 = maxilla 1; Mx2 = maxilla 2. 
Appendages on thoracic somites are Mxp = 
maxilliped (thoracopod 1); Pl-5 = swim- 
ming legs (thoracopods 2—6). The caudal 
ramus is CR. Designations of appendage 
segments are according to Ferrari (1995) as 
follows: medial lobe of a segment = hi, lat- 
eral lobe = le; rami are exopod = Re and 
endopod = Ri; ramal segments of Mx2 are 
exopodal; the Mxp has a basopod with a 


78 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


distal medial lobe, and with at most five 
endopodal segments in calanoids (Ferrari & 
Dahms 1998). Armament elements of ap- 
pendages are termed setae regardless of 
their position or degree of rigidity. Setules 
are epicuticular extensions of a seta; denti- 
cles are epicuticular extensions of an ap- 
pendage segment; spinules are epicuticular 
extensions of a somite. 

Calanoid superfamily names follow An- 
dronov (1974) with the exception of Clau- 
socalanoidea for Pseudocalanoida (Bow- 
man 1987, International Commission on 
Zoological Nomenclature 1988). The 
changes proposed by Andronov (1991) are 
not followed here because there is no re- 
quirement that a family group name be de- 
rived from the oldest included genus name, 
or by coordination here the oldest included 
family name (International Code of Zoolog- 
ical Nomenclature 1985, articles 62—64). 
The correct names in question are Augap- 
tiloidea, Centropagoidea and Megacalano- 
idea. 


Griceus, new genus 


Diagnosis.—The following derived char- 
acter states are assumed to be shared by all 
species of this genus discovered subse- 
quently: dorsal interlocking extensions of 
abdominal somites; A2, Mn and Mx! orig- 
inating posterior to the labrum and parag- 
naths; coxal endite of Mn outside of the la- 
bral-paragnathal envelope; praecoxal and 
coxal endites of Mx1 unarmed; praecoxal 
endites of Mx2 unarmed; endopod of Mxp 
with four articulating segments. 

Type species.—Griceus buskeyi, by mon- 
otypy. 

Etymology.—The name honors Dr. 
George D. Grice for his contributions to the 
exploration of the deep-sea benthopelagic 
fauna, and to the taxonomy of calanoid co- 
pepods. Dr. Grice’s family name is latinized 
to form Griceus. The gender of the name is 
masculine. 


Griceus buskeyi, new species 


Specimens.—Holotype (National Muse- 
um of Natural History, Smithsonian Insti- 
tution USNM 288058) a dissected female, 
0.90 mm in length; prosome—0.74 mm and 
urosome—O.16 mm. 

Female.—Pr (Fig. 1C): 6 segments; Ist a 
complex of 5 cephalic somites plus Thl; 
Th2—4 simple and articulated; arthrodial 
membrane between Th5 and 6 incomplete; 
reduced in width dorsally. Rostral area a 
simple, bilobe plate (Fig. 1A, C) without 
spinules or attenuations of cephalon. Lo- 
cation of base of A2, Mn, Mx1, Mx2 and 
Mxp as in Fig. 1A, B. Base of A2 posterior 
to labrum; base of Mn outside of labral- 
paragnathal envelope. Base of Mxp lateral 
to base of Mx2; base of Mx1 lateral and 
posterior to base of Mx2. Rostrum, labrum, 
and paragnaths as in Fig. 2D. 

Ur (Fig. 1D): 4 segments; anterior seg- 
ment a genital complex of Th7 and Ab1; as 
viewed dorsally, genital complex symmet- 
rical (Fig. 2A). Viewed laterally (Fig. 2B, 
C), genital complex with a thickened ridge 
laterally on right side. Right laterally and 
dorsad, posterior margin of abdominal so- 
mites 1—3 asymmetrical, each with attenu- 
ate, finger-like process which engages sock- 
et-like depression on anterior margin of fol- 
lowing somite (Fig. 2E). 

Al (Fig. 3): A large segmental complex 
proximally (Fig. 4A) of 19 setae separated 
by incomplete ventral (anterior) arthrodial 
membranes into 8 groups of 3, 2, 2+1, 3, 
2, 3, 1, 2 setae+aesthetascs with setae of 
same group laterally displaced relative to 
each other; aesthetasc of the 3rd group 
short, triangular with distal sensilla. Distal 
to the complex an incompletely articulating 
segment with 2 setae and short, triangular 
aesthetasc with distal sensilla followed by 
13 completely articulating segments with 
Zt hig, Qine2 pe Did sp Dole Lawton Dee wea cee 
6+1 setae+aesthetascs. 

A2 (Fig. 4B, C): coxa and basis without 
setae. Re 8 articulating segments with 0, 1, 


VOLUME 113, NUMBER 1 


79 


Fig. 1. 


Griceus buskeyi new genus, new species. A, Oral area, ventral, showing location of only one of 


antenna | (1), antenna 2 (2), mandiblar gnathobase (3), maxilla 1 (4), maxilla 2 (5), maxilliped, obscuring 
mandibular palp (6); both swimming legs | (7); B, same, showing location of origin of antenna | (1), antenna 
2 (2), mandible (3), maxilla | (4), maxilla 2 (5), maxilliped (6); C, animal, left lateral; D, Th5—6 and urosome, 


dorsal; E, leg 5. All scale lines are 0.01 mm. 


1, 1, 1, 1, 0, 3 setae. Ri 2-segmented with 
1 and 8 (4 terminal, 4 subterminal) setae. 

Mn (Fig. 4D): coxa elongate with lobe at 
mid-length and attenuations medially; basis 
unarmed. Re 5-segmented with 1, 1, 1, 1, 1 
setae. Ri apparently a lobe on the basis me- 
dial to the exopod. 

Mx1 (Fig. 4E): Praecoxa, coxa and basis 
with poorly-developed, unarmed lobes. Ri 


not distinguished. Re 1l-segmented with 1 
medial and 3 terminal setae. 

Mx2 (Fig. 4F): Proximal and distal prae- 
coxal endites poorly-developed, unarmed; 
no arthrodial membrane separating prae- 
coxa and coxa. Proximal coxal endite with 
2 setae, distal coxal endite with 2 setae. En- 
dites of basis indistinct with 1 and 2 setae. 
Re unsegmented with 7 setae. 


80 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 2. 


= 


Griceus buskeyi new genus, new species. A, Genital complex, dorsal; B, Genital complex, left lateral, 


C, Genital complex, right lateral; D, Mouth area, ventral, showing location of rostrum (1), labrum (2), paragnath 
(3), tip of mandibular gnathobase (4) and origin of antenna 2 (5); E, urosome, dorsal and slightly lateral, with 
attenuate finger-like processes and socket-like depressions darkened. All scale lines are 0.01 mm. 


Mxp (Fig. 5A): syncoxa with 1 seta but 
without distinct lobes. Basis with 3 setae (2 
on a distal medial lobe which is poorly de- 
veloped and weakly sclerotized) and ante- 
rior denticles not organized into a clear pat- 
tern. Ri 4-segmented proximal to distal with 
1, 2, 3, 4 setae (4, 3, 1, 2 by segmental age). 

P1 (Fig. 5B): coxa with medial seta. Ba- 
sis with medial denticles; medial seta 


curved only toward its tip with setules 
along the proximal straight part; lateral seta 
absent. Re apparently 3-segmented with 
distal segment missing; proximal segment 
with 2 (medial and lateral) setae; middle 
segment with 2 (medial and lateral) setae. 
Ri 3-segmented; proximal segment with 
medial seta, a pore on anterio-distal margin 
(with the inner seta of the basis comprising 


VOLUME 113, NUMBER 1 


Fig. 3. 


Von Vaupel Klein’s organ) and a lateral at- 
tenuation; middle segment with 2 medial 
setae; distal segment with | medial, 2 ter- 
minal and 2 lateral setae. 

P2 (Fig. 5C): coxa with medial seta. Ba- 
sis unarmed. Re apparently 3-segmented 
with distal segment missing; proximal seg- 
ment with 2 (medial and lateral) setae; mid- 
dle segment with 2 (medial and lateral) se- 
tae. Ri 3-segmented; proximal segment 
with medial seta and middle segment with 
2 medial setae; distal segment with 3 me- 
dial, 2 terminal and 2 lateral setae. 

P3 (Fig. 5D): coxa with medial seta. Ba- 
sis unarmed. Re apparently 3-segmented 
with distal segment missing; proximal seg- 
ment with 2 (medial and lateral) setae; mid- 
dle segment with 2 (medial and lateral) se- 
tae. Ri apparently 3-segmented; proximal 
segment with medial seta and middle seg- 
ment with 2 medial setae; distal segment 
missing. 

P4 (Fig. 5E): coxa with medial seta. Ba- 
sis with lateral seta. Re apparently 3-seg- 
mented with distal segment missing; prox- 
imal segment with 2 (medial and lateral) 
setae; middle segment with 1 medial seta 
(lateral seta apparently missing). Ri appar- 
ently 3-segmented; proximal segment with 
a medial seta and middle segment with 2 
medial setae; distal segment missing. 

Leg 5 (Fig. 1E): coupler uniting limbs 
which each bear a terminal seta. 


Griceus buskeyi new genus, new species. A, 


81 


antenna |, left. Scale line is 0.01 mm. 


CR (Fig. 1D): 4 thick apical setae; dorsal 
seta small and thin. 

Male.—unknown. 

Etymology.—The name recognizes Ed- 
ward Buskey for collecting the sample 
which contained this copepod. The specific 
epithet is a noun in the genitive singular. 

Remarks.—The unknown, but possibly 
extensive, period of time this specimen may 
have remained unfixed in the collecting net 
makes difficult the determination of several 
possible synapomorphies involving setae or 
spinules. For example, a rostrum without 
spinules (filaments), antenna 2 basis with- 
out setae, antenna 2 endopod with four ter- 
minal setae and mandibular exopod with 
one seta on the terminal segment all rep- 
resent potential apomorphies that should be 
verified when better preserved specimens 
are available. For this reason, we have cho- 
sen to emphasize the apparently derived na- 
ture of the following changes in shape of 
somites and appendage segments which are 
presumed apomorphies for species of Gri- 
ceus: interlocking extensions of abdominal 
somites dorsally; antenna 2, mandible and 
maxilla 1 originating posterior to the la- 
brum and paragnaths; coxal endite of the 
mandible lying outside of the labral parag- 
nathal envelope; praecoxal and coxal en- 
dites of maxilla 1 unarmed; praecoxal en- 
dites of maxilla 2 unarmed; endopod of 
maxilliped with four articulating segments. 


82 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 4. Griceus buskeyi new genus, new species. A, proximal segmental complex plus following incompletely 
articulating segment of left antenna 1, ventral view, distal is down, wavy line cutoff on broken seta, circle with 
X is setal scar, broken lines are incomplete arthrodial membranes (scale line as for B); B, exopod of right antenna 
2 with, segments |, 2—4 and 9 indicated; C, left antenna 2, with exopod broken; D, left mandible; E, left maxilla 


1; E left maxilla 2. All scale lines are 0.01 mm. 


The effect of the changes in location of 
the cephalic appendages may be general- 
ized as follows: A2, Mn and Mx1 retain 
their ancestral positions relative to one an- 


other but as a group they have been drawn 
back posteriorly and laterally from the usu- 
al calanoid locations. The penultimate seg- 
ment of the maxilliped may be a complex 


VOLUME 113, NUMBER 1 


Fig: 5. 


83 


Griceus buskeyi new genus, new species. A, left maxilliped; B, swimming leg 1, anterior; C, swim- 


ming leg 2, posterior; D, swimming leg 3, posterior; E, swimming leg 4, posterior. All scale lines are 0.01 mm. 


of 2nd and 5th segments with the arthrodial 
membrane missing because it bears more 
setae than the two segments proximal to it 
(see Ferrari 1995, Ferrari & Dahms 1998). 
The endopod of the maxilliped of derived 
centropagoideans is poorly sclerotized and 
without arthrodial membranes; setal addi- 
tions during development of Acartia tonsa 
Dana, 1849, Tortanus dextrilobatus Chen & 
Zhang, 1965, and Epilabidocera longipe- 
data (Sato 1913) suggest these endopods 
are 4-segmented with one seta on proximal, 
antepenultimate and penultimate segments, 


and two setae on the distal segment. The 
endopod of G. buskeyi is well-sclerotized; 
there is one seta on the proximal, two setae 
on the antepenultimate, three setae on the 
penultimate and four setae on the distal seg- 
ments suggesting that this 4-segmented ra- 
mus is not convergent with the centropa- 
goideans. The ramus of maxilla | is inter- 
preted as an exopod because its quadrate 
morphology and terminal crown of setae is 
similar to the exopod of many heterorhab- 
dids and augaptilids. We know of no cal- 
anoid copepod with a maxilla 1 in which 


84 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


the endopod is the only ramus. Segmental 
homologies of leg 5 are based on the lo- 
cation of the terminal seta toward the lateral 
side of the appendage. This seta is inter- 
preted as the terminal seta of the distal ex- 
opodal segment; the proximal segment is a 
fused coxa plus basis. 

We have reconsidered what kinds of in- 
formation the morphology of the exopod of 
A2 provides about the superfamily of Gri- 
ceus. We do not follow the hypothesis of 
Park (1986) that the exopod of antenna 2 
of species of Augaptiloidea have only nine 
segments, none of which are complexes, 
while among the remaining superfamilies 
there are up to 10 segments, and except for 
the Eucalanoidea with the 2nd always fused 
to the 3rd and 9th always fused to the 10th. 
Which of the ten segments present in the 
remaining superfamilies fails to form in 
Augaptiloidea is not indicated by Park 
(1986). Here, homologies of the exopod are 
derived from the following assumption: all 
segments, with the exception of the distal 
segment, are serial homologues bearing at 
most one medial seta. The female of Rhin- 
calanus gigas Brady, 1883 has nine medial 
setae, each proximal to an arthrodial mem- 
brane, and a distal segment with a terminal 
and two sequentially arranged medial setae 
(Fig. 6A). An incomplete, distal arthrodial 
membrane extends in part along the anterior 
and posterior surfaces between the second 
and third medial setae, while an incomplete, 
distal arthrodial membrane fails to extend 
to the lateral surface separating the 3rd and 
4th medial setae. Giesbrecht (1892: plate 
11, figs. 16, 17) shows the exopod of Eu- 
calanus attenuatus (Dana 1849) and E. 
crassus Giesbrecht, 1888 which can be in- 
terpreted as 11-segmented, a proximal com- 
plex of four segments represented by four 
medial setae, followed by six articulating 
segments, each with a seta, and a distal seg- 
ment with a crown of three setae. We have 
been able to verify only a crown of three 
setae and nine medial setae each with at 
least part of a distal arthrodial membrane 
for these species. 


Males of Temora longicornis (Miiller, 
1785) have a more common calanoid mor- 
phology (Fig. 6B). An elongate segment 
distally with a medial seta near its mid- 
length is assumed to correspond to the pen- 
ultimate segment which is elongate distad 
from the seta in this species. The distal seg- 
ment has a crown of three setae correspond- 
ing to the distal segment of R. gigas. The 
2nd articulating segment with three medial 
setae is assumed to be a complex of the 
2nd, 3rd, and 4th segments in which the 
incomplete arthrodial membranes of R. gi- 
gas fail to form on T. longicornis. The five 
remaining articulating segments with a me- 
dial setae and distal arthrodial membrane 
are the Ist and 5th—8th of R. gigas. There 
are many calanoids in which the distal ar- 
throdial membrane of three or more proxi- 
mal segments fails to form; in these cases, 
a medial setae is assumed to represent the 
location of each segment. For example, in 
Calanus finmarchicus (Gunnerus, 1765) the 
elongate second segment shows no trace of 
an arthrodial membrane on its dorsal sur- 
face but four medial setae suggest this is a 
complex of the 2nd—5th segments; the re- 
maining distal segments, each with a medial 
seta, articulate distally (Fig. 6C). 

In contrast, in augaptiloidean species like 
Phyllopus bidentatus Brady, 1883 and Het- 
erorhabdus spinifrons (Claus 1863), a me- 
dial seta on each of a set of proximal seg- 
ments fails to form while the distal arthro- 
dial membrane of each of those segments is 
present (Giesbrecht 1892: plate 18, fig. 29 
and plate 20, fig. 9). Ohtsuka et al. (1994) 
describe arietellids with an elongate, prox- 
imal segmental complex in which both the 
seta and distal arthrodial membrane of a set 
of segments apparently fail to form. Gri- 
ceus buskeyi has its 1st segment with a dis- 
tal arthrodial membrane but no medial seta; 
the following segmental complex is inter- 
preted as composed of the 2nd and 3rd seg- 
ments which have neither an arthrodial 
membrane nor a medial seta, plus the 4th 
segment with a medial seta and a distal ar- 
throdial membrane. The next four articulat- 


VOLUME 113, NUMBER 1 


85 


Fig. 6. Rhincalanus gigas antenna 2 exopod; Temora longicornis antenna 2 exopod; Calanus finmarchicus 
antenna 2 exopod. Presumed segments are numbered proximal to distal; scale lines for A and C are 0.01 mm; 


B is 0.05 mm. 


ing segments each with a seta are the 5th— 
8th segments. The following elongate seg- 
ment is an unarmed 9th segment and the 
10th segment has a crown of three setae. 
Our interpretation assumes that all seg- 
ments homologous to those of R. gigas are 
present in these calanoids. The alternate hy- 
pothesis of Park (1986) that one of the ten 
segments fails to form, has not been con- 
sidered because we are unsure of the loca- 
tion on the antennal exopod that new seg- 
ments are patterned, and whether there is a 
single location for segment patterning, like 
the copepod maxilliped (Ferrari & Dahms 
1998), or more than one location, like the 
calanoid antenna | (Ferrari & Benforado 
1998). 

Loss of segments on some of the swim- 
ming legs complicates the assignment of 
this specimen to the correct calanoid family. 
However, two character states suggest that 


G. buskeyi belongs to the superfamily Au- 
gaptiloidea. Two setae on middle segment 
of endopod of swimming legs 3 and 4, 
probably an ancestral calanoid state, are 
present only on Epacteriscioidea, Pseudo- 
cyclopoidea, Augaptiloidea, Centropago- 
idea, Megacalanoidea, and some Bathypon- 
tioidea (Andronov 1974, Suarez-Morales & 
Iliffe 1996). We note in passing a misprint 
in Table 1 of Suarez-Morales & Iliffe 
(1996) in the number of inner setae on the 
terminal segment of swimming legs 3 and 
4 (column E); Pseudocalanoidea (Clauso- 
calanoidea) are unique in having four setae 
while the Spinocalanoidea with five setae 
are identical to the remaining superfamilies. 
The Arietellidae and Heterorhabdidae are 
the only families among the above six su- 
perfamilies with species in which the en- 
dopod of maxilla 1 does not develop (Oht- 
suka et al. 1994, Park 2000), so an assign- 


86 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


ment to the Augaptiloidea seems reason- 
able. 

Among the eight families and 39 genera 
of Augaptiloidea, a set of derived characters 
states have been proposed only for the fam- 
ily Heterorhabdidae and its genera (Park 
2000), so placement of G. buskeyi in a fam- 
ily is difficult. With respect to the affinity 
of G. buskeyi, a small unarmed mandibular 
endopod and maxilla 1 without an endopod 
are character states shared only with Par- 
augaptilus Wolfenden, 1904 (Arietellidae) 
and a few species of Augaptilus Giesbrecht, 
1889 (Augaptilidae). The absence of setae 
on the praecoxal endites of maxilla 2 sug- 
gests a relationship with Paraugaptilus 
which has only one seta on each praecoxal 
endite. All species of Augaptilus have two 
setae on these endites; this is a widespread 
and presumably ancestral state for the Au- 
gaptiloidea. Given our limited knowledge 
the Augaptiloidea and its families, the 
shared similarities of G. buskeyi to species 
of Paraugaptilus provides a reasonable jus- 
tification for placing this new genus in the 
Arietellidae. 


Literature Cited 


Andronovy, V. N. 1974. Filogeneticheskie otnosheniya 
krypnykh taksonov podotryada Calanoida 
(Crustacea, Copepoda).—Zoologicheskii Zhyr- 
nal 53:1002—1012. [Phylogenetic relationships 
of the large taxa within the suborder Calanoida 
(Crustacea, Copepoda).—translated, 1980, by 
Alahram Center for Scientific Translations] 

. 1991. On renaming of some taxa in Calanoida 
(Crustacea).—Zoologicheskii Zhyrnal 70:133- 
134 [in Russian, English summary]. 

Bowman, T. E. 1987. Comment on the proposed pre- 
cedence of Pseudocalanidae Sars, 1901 (Crus- 
tacea, Copepoda) over Clausocalanidae Gies- 
brecht, 1892.—Bulletin of Zoological Nomen- 
clature 44:129. 

Brady, G. S. 1883. Report on the Copepoda. Part 23.— 
Report on the Scientific Results of the voyage 
of H.M.S. “‘Challenger’’ during the years 1873-— 
1876, Zoology 8:1—142, 55 pls. 

Claus, C. 1863. Die freilebenden Copepoden mit be- 
sonderer Beriicksichtigung der Fauna Deutsch- 
lands, der Nordsee und des Mittelmeeres. W. 
Engelmann, Leipzig, 230 pp., 37 pls. 

Chen, Q., & S. Zhang. 1965. The planktonic copepods 


of the Yellow Sea and the East China Sea I. 
Calanoida.—Studia Marina Sinica 7:20—-131 + 


53 pls. 
Dana, J. D. 1849. Conspectus crustaceorum, quae in 
orbis terrarum circumnavigatione, Caroli 


Wilkes, e classe Reipublicae foederatae duce, 
lexit et descripsit Jacobus D. Dana.—Ameri- 
can Journal of Science (2) 8:276—285, 424- 
428. 

Ferrari, F D. 1995. Six copepodid stages of Ridge- 
wayia klausruetzleri, a new species of copepod 
crustacean (Ridgewayliidae, Calanoida) from 
the barrier reef in Belize, with comments on 
appendage development.—Proceedings of the 
Biological Society of Washington 108:180— 
200. 

, & A. Benforado. 1998. Setation and setal 

groups on antenna | of Ridgewayia klausruet- 

zleri, Pleuaromamma xiphias, and Pseudocalan- 
us elongatus (Crustacea: Copepoda: Calanoida) 
during the copepodid phase of their develop- 
ment.—Proceedings of the Biological Society 

of Washington 111:209-—221. 

, & H.-E. Dahms. 1998. Segmental homologies 
of the maxilliped of some copepods as inferred 
by comparing setal numbers during copepodid 
development.—Journal of Crustacean Biology 
18:298-307. 

Giesbrecht, W. 1888. Elenco dei Copepodi pelagici 
raccolti dal Tenete di vascello Gaetano Chier- 
chia durante il viaggio della R. Corvetta “‘Vettor 
Pisani’? negli anni 1882—1885 e dal Tenete de 
vascello Francesco Orsini nel Mar Rosso, nel 
1884.—Rendiconti della Reale Accademia dei 
Lincei Classe di Scienze fisiche, matematiche e 
naturali 4, semestre 2:284—287, 330-338. 

. 1889. Elenco dei Copepodi pelagici raccolti 

dal Tenete di vascello Gaetano Chierchia dur- 

ante il viaggio della R. Corvetta “‘Vettor Pisa- 
ni” negli anni 1882—1885 e dal Tenete de vas- 
cello Francesco Orsini nel Mar Rosso, nel 
1884.—Rendiconti della Reale Accademia dei 

Lincei Classe di Scienze fisiche, matematiche e 

naturali 5, semestre 1:811—815, semestre 2:24— 

20: 

. 1892. Systematik und Faunistik der pelagisch- 
en Copepoden des Golfes von Neapel und der 
angrenzenden Meeres-abschnitte.—Fauna und 
Flora des Golfes von Neapel und der angren- 
zenden Meeres-abschnitte 19:1—831 + 54 pls. 

Gunnerus, J. E. 1770. Nogle smaa rare og meesten- 
deelen nye Norske Sgedyr.—Skrifter, som udi 
det Kigbenhavnske Selskab af Laerdoms og Vi- 
denskabers Elskere, 1765-1769, 10:166—176. 

International Code of Zoological Nomenclature. 1985. 
Articles 62—64. P. 119 in W. D. L. Ride et al., 
eds., 3rd Edition. University of California Press, 
Berkeley, 338 pp. 


VOLUME 113, NUMBER 1 


International Commission on Zoological Nomencla- 
ture 1988. Opinion 1503: Pseudocalanidae Sars, 
1901 (Crustacea, Copepoda) not to be given 
precedence over Clausocalanidae Giesbrecht 
1892.—Bulletin of Zoological Nomenclature 
43:228-229. 

Miiller, O. EK 1785. Entomostraca seu Insecta Testacea 
quae in aquis Daniae et Norvegicae reperit, des- 
cripsit et iconibus illustravit Otho Fridericus 
Miiller. EF W. Thiele, Lipsiae & Havniae, 134 
pp. + 21 pls. 

Ohtsuka, S., G. A. Boxshall, & H. S. J. Roe. 1994. 
Phylogenetic relationships between §arietellid 
genera (Copepoda: Calanoida), with the estab- 
lishment of three new genera.—Bulletin of the 
Natural History Museum London (Zoology) 60: 
105-172. 


87 


Park, T. 1986. Phylogeny of calanoid copepods.—Syl- 
logeus 58:191—196. 

. 2000. Taxonomy and distribution of the ma- 
rine calanoid copepod family Heterorhabdi- 
dae.—Scripps Institution of Oceanography Bul- 
letin 31 (in press). 

Sato, C. 1913. Fuyusei-Tokyakurui.—Suisan Chosa 
Hokoku 1:28-—29. 

Suarez-Morales, E., & T. M. Iliffe. 1996. New super- 
family of Calanoida (Copepoda) from an an- 
chialine cave in the Bahamas.—Journal of 
Crustacean Biology 16:754—762. 

Wolfenden, R. N. 1904. Notes on the Copepod of the 
North Atlantic Sea and the Farée Channel.— 
Journal of the Marine Biological Association of 
the United Kingdom (new series) 7:110—146, 
pee. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


113(1):88—-94. 2000. 


Studies on the Crustacea of the Turks and Caicos Islands, British 
West Indies. IV. Heteromysis (Heteromysis) spottet, a new species 
(Peracarida: Mysidacea: Mysidae) from Pine Cay 


W. Wayne Price and Richard W. Heard 


(WWP) Department of Biology, University of Tampa, Tampa, Florida 33606, U.S.A.; 
(RWH) University of Southern Mississippi, Institute of Marine Sciences, 
Ocean Springs, Mississippi 39566-7000, U.S.A. 


Abstract.—During a survey of the marine Crustacea in the vicinity of Pine 
Cay, Turks and Caicos Islands, an undescribed mysid, Heteromysis spottei, new 
species, was collected at depths ranging from 1 to 18 m. The new species 
belongs to the subgenus Heteromysis Bacescu and can be distinguished from 
the five other members of that subgenus known from the western Atlantic by 
the setation of the telson. In Heteromysis spottei the telson has spine-setae 
along the entire length of the lateral margins, spinules along the entire margins 
of the cleft, and a pair of spine-setae on each apical lobe, the outer being nearly 
three times longer than the inner. A key and diagnostic table to the species of 
the subgenus Heteromysis from the Northwest Atlantic is presented. 


Excluding the commensal palaemonid 
shrimps, which have been addressed in a 
separate series of publications (Heard & 
Spotte 1991, Heard et al. 1993, Spotte et al. 
1994, Spotte & Bubucis 1996, Heard & 
Spotte 1997), this is the fourth contribution 
in a series on the crustacean fauna of the 
Turks and Caicos Islands. The first three re- 
ports dealt with marine isopods (Kensley & 
Heard 1991, Schotte & Heard 1991, Schotte 
et als 199): 

This report presents the description of a 
new species of mysid belonging to the ge- 
nus Heteromysis Smith, 1873. The new spe- 
cies, which is referred to the subgenus Het- 
eromysis (Heteromysis) Smith 1873 sensu 
Bacescu, 1968, was collected in water 
depths ranging from 1 to 18 m in the vicin- 
ity of Pine Cay. 

The type material was deposited in the 
National Museum of Natural History 
(USNM), Smithsonian Institution, and the 
Gulf Coast Research Laboratory (GCRL). 
We follow the setal classification of Watling 
(1989) in which a seta is defined as an ar- 
ticulated cuticular outgrowth of the integu- 


ment. The term “‘spine-seta’’ as used here 
refers to any seta with spine-like character- 
istics; a spine is defined as a non-articulated 
extension of the cuticle. 


Heteromysis (Heteromysis) spottei, new 
species 
Figs. 1-2 


Material examined (all material from 
Turks and Caicos).—Holotype: adult @ 
(Length ,[L] 3:1 mm), JUSNM) 282712: 
fringing reef off Pine Cay, live bottom/ 
sand, depth 18 m, J. A. McLelland (coll.), 
suction device, 12 Nov 1989. Paratypes: 
adult ¢ (L3.5mm), USNM 282720; adult 
6, damaged, GCRL 1346, same collection 
data as holotype.—1 adult ¢d, 2 29, (all 
damaged) Pine Cay, shallow inner reef, 
sand-coral rubble substratum with associ- 
ated sponges and cnidarians, depth 4 m, 1 
mm mesh dredge net, R. Heard, J. Mc- 
Lelland, P. Bubucis, & S. Spotte (colls.), 5 
Nov 1988.—16d6 (damaged), Pine Cay 
(Rock-a-Wash Cay), depth 1 m, Neogoniol- 
ithon & sponge washings, R. Heard (coll.), 


VOLUME 113, NUMBER 1 


30 Oct 1988.—2¢ 6, 1 ¢ (all damaged), 
same data as holotype. 
Diagnosis.—Article 3 of antennular pe- 
duncle without distomedial flagellated 
spine-seta; thoracic endopod 3 with 3 flag- 
ellated spine-setae on medial margin of 
merus, medial margin of carpo-propodus 
without flagellated spine-setae; pleopods 
uniramous, reduced to simple setose plates 
with no modified spine-setae in either sex; 
endopod of uropod armed with 1 small 
spine-seta near statocyst; lateral margins of 
telson armed along entire length with 10— 
12 spine-setae per margin (including apical 
spine-setae), posterior-most lateral spine- 
seta 1.6—-1.7 times length of preceding 
spine-seta, extending beyond posterior end 
of telson; outer apical spine-seta nearly 3 
times longer than inner; cleft completely 
armed with 18—20 spinules. 
Description.—General body form (Fig. 
1A): moderately robust; carapace with an- 
terior margin produced into pointed trian- 
gular rostrum; posterior dorsal margin 
emarginate, partly exposing thoracic seg- 
ment 8; anterolateral lobes rounded. 
Antennule peduncle (Fig. 1B): article 1 
slightly shorter than article 3, with 3 plu- 
mose and 2 simple spine-setae on distolat- 
eral process, dorsomedial longitudinal ridge 
with 4 strong simple distal spine-setae; ar- 
ticle 2 compressed with 2 distomedial 
spine-setae, | simple and 3 plumose spine- 
setae near middle of distal margin; article 3 
with simple spine-seta on medial margin, 3 
simple distomedial spine-setae, 3 plumose 
spine-setae near distolateral margin, | sim- 
ple spine-seta on dorsolateral surface, lack- 
ing flagellated spine-seta, males with small 
moderately setose lobe on ventral surface. 
Antenna (Fig. 1C): scale slightly shorter 
than peduncle, 3.0—3.2 times as long as 
maximum width, medial margin strongly 
convex, lateral margin straight, all margins 
setose, lacking distal article; antennal pe- 
duncle having 3 articles; article 1 incon- 
spicuous; article 2 just over 1.5 times lon- 
ger than article 3, short plumose spine-seta 
near distolateral border, 2 simple and 2 plu- 


89 


mose distomedial spine-setae; article 3 with 
1 plumose and 3 simple distomedial spine- 
setae, 3 plumose spine-setae along lateral 
margin. 

Eyes (Fig. 1A): large, oval, directed lat- 
erally, distal part of eye stalk wider than 
cornea, lacking ocular tooth; cornea large, 
oval. 

Mandibles (Fig. 1D—F): molar, incisor, 
and lacina mobilis as illustrated. Palp 3-seg- 
mented; article | small, inconspicuous; ar- 
ticle 2 expanded, medial margin with 5—10 
simple spine-setae, lateral margin with 4—5 
proximal plumose spine-setae and | simple 
distolateral spine-seta; article 3 half as long 
as 2, medial margin with | plumose spine- 
seta, distal part armed with | simple and 9— 
10 barbed spine-setae, 1 simple spine-seta 
on medial surface. 

Labrum and paragnaths (Fig. 1D, G): as 
illustrated. 

Maxillule (Fig. 1H): outer lobe with 11 
stout apical and 3 subapical spine-setae; in- 
ner lobe with 3 long, distally curved, serrate 
spine-setae, 2 plumose and 2 simple spine- 
setae distally, and 1 plumose and | simple 
spine-seta on distomedial margin. 

Maxilla (Fig. 11): as illustrated; exopod 
with 13—15 plumose spine-setae on disto- 
lateral margin. 

Thoracic endopods 1-8 (Fig. 2A-—G): 
thoracic endopods 1 and 2 as illustrated. 
Thoracic endopod 3, merus approximately 
1.6 length of ischium and equal in length 
to carpo-propodus, medial margin with 3 
flagellated and 3 simple spine-setae, lateral 
margin with | distal simple spine seta; me- 
dial margin of carpo-propodus with 4—5 
simple spine-setae, 2 simple and 2 stout ser- 
rate spine-setae on distomedial margin; dac- 
tyl small, with long, slightly curved claw 
on distal end surrounded by several simple 
spine-setae. Thoracic endopod 4, merus 
about 1.5 length of ischium; carpo-propo- 
dus about %4 length of merus, with 3 articles 
(distal 2 subequal, combined length slightly 
longer than proximal article); dactyl mi- 
nute, papillate, lacking terminal claw. Tho- 
racic endopod 5, ischium and merus sub- 


90 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 1. 


= 
Sy 
Zp ~ 


Heteromysis spottei, new species. A, adult 2, dorsal view; B, antennular peduncle; C, antennal 


peduncle and scale; D, labrium & right mandibular palp; E, KE right mandible, inner and upper aspects, respec- 
tively; G, paragnaths; H, maxillule; I, maxilla. Scale 1, A = 0.5 mm; C = 0.2 mm; B, D-E I = 0.1 mm; A = 


0.05 mm; Scale 2, G = 0.1 mm. 


equal in length; carpo-propodus with 4 ar- 
ticles, distal 3 subequal in length, each half 
as long as proximal article, ultimate article 
with 6—7 simple spine-setae (2 sickle- 
shaped with strong articulated bases); dac- 
tyl small, papillate with long distal, slender 
claw. Thoracic endopod 6, carpo-propodus, 
ischium and merus subequal in length; car- 
po-propodus with 5 articles, distal 4 sub- 
equal in length, each about % as long as 
proximal article; dactyl small, papillate with 
distal serrate slender claw. Thoracic endo- 
pod limb 7, ischium slightly longer than 


merus, merus slightly longer than carpo- 
propodus; carpo-propodus with 3 articles, 
proximal distinctly longer than each of dis- 
tal articles; dactyl small, papillate with dis- 
tal serrate slender claw. Thoracic endopod 
8 (1 damaged limb available for study), 
long, attenuated, at least 4% longer than other 
thoracic endopods; carpo-propodus with at 
least 5 articles. 

Thoracic exopods: exopod 1 with 8 arti- 
cles; exopods 2—8 with 9 articles. 

Thoracic sternal processes: median spi- 
niform processes on sterna 3—7 in males. 


VOLUME 113, NUMBER 1 


9] 


Fig, 2. 


Heteromysis spottei, new species. A-—G endopods of thoracic limbs 1—7; H, telson; I, uropod. Scale 


1, H = 0.1 mm; Scale 2, A = 0.1 mm, B—G = 0.2 mm; Scale 3, I = 0.2 mm. 


Pleopods: without sexual dimorphism; 
reduced to uniramous plates with no mod- 
ified spine-setae, similar in form. 

Uropods (Fig. 21): exopod about 1.2 
times longer than endopod, lateral margin 
straight, medial margin slightly convex, all 
margins setose; endopod linguiform with 1 
spine-seta on medial margin in region of 
statocyst, all margins setose. 

Telson (Fig. 2H): 0.8 times length of ex- 
opod of uropod, 1.4—1.5 times as long as 


maximum width, lateral margins slightly 
concave, armed along entire length with 10— 
12 spine-setae per margin (apical spine-setae 
included), increasing in length posteriorly, 
most posterio-lateral spine-seta 1.6—1.7 
times length of preceding spine-seta, extend- 
ing beyond posterior lobes of telson; outer 
apical spine-seta 2.8—3.0 times longer than 
inner; cleft, depth 0.25 length of telson, 
completely armed with 18—20 small spines. 

Etymology.—This species is named for 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Neen nner nnn rr nn reese SSS 


Cenvic ERS OG EC OiGaL Cc OVr-8e [eos [BUUDIUR JO ORI YIPIM:yIsUsT 

juasqe juasoid juasoid juaso.id juasqe juosoid gyeos [RUUDJUR UO JOTI [eISIG 
Yjsud] JOUUT:19}NO 

O'€-8'7 ‘1Ied O'7 ‘aed CQ ‘aed o[suls o[ SUIS o[ SUIS ‘9qo] Jod ovjas-auids uosja} jeoidy 
Elev ie orp) OSE (pepnyout ovjos ouids 

TI-OI “OAMUD = p—€_‘JTBY AOLID}sod Q ‘yyey 10110}sod ‘yyey 1011a}sod ‘yyey 10119)sod ‘yvey 10113a}sod jeoide) uosya} uo oejas-outds [ese] 

SI-Ol 

OT-8I ‘anus OZ ‘a1uS QI ‘Jey jso11ajue €7-7T ‘arus O€-9] ‘ornUS ‘yyey 1O19}uUe We]O UOSTa} UT saynuIds 

I 0 I €7-0Z 6I-+rI CT-SI podopus [epodoin uo oejas-auids ‘oN 

ouou ouou é ouou ouou ouou spodogjd ayeu poyipoyy 
Gipe ¢€ podopud s1dv10Yy) Jo sn13W 

-jel[osey ‘¢ ‘ojduns L ‘oyduuts ZT ‘oydurts ouou €[-8 ‘o[duits L ‘oydurts JO UISIeU [eIpsu UO avRjas-suIds 
Z ‘oyelas ‘1op ZT poeyjosey yno}s 9 jNO}s Q ‘Jap OI ynoqe ¢€ podopus s1ov10y) Jo snpodo.id-od 

-ug[s /-9 ‘ajduiis = ‘ropusys 6 ‘ajduris = ‘tapusys / ‘aydurts ouou = -UaTS 6-8 ‘ayduuts ‘Jopus[s ‘ojduts = -1v9 JO UTSIeU [eIpoW UO deIaS-9UIdS 
ajyounpod 

juasqe juasqe juosqe juasqe juasqe juasqe Jejnuusjue uo vjas-oulds payey[aseyy 


canner nr SSS Ss 


1ajJods inuvasainu 1jaanou DUDIIXIU psowusof suds] Jquoevieyo 


See 


sa1oadg 
anne nnn nn SSS 


“ONUL[IY UI9ISOMYIOU IY) Ul SISdMOsLIJAF] SNUISQNS dy} JO S1d}dvIeYO J[Npe oytoods puke uoWWIOD—'| 2IqQR I], 


VOLUME 113, NUMBER 1 


Stephen Spotte in recognition of his many 
contributions to marine science. 

Habitat.—Heteromysis spottei was col- 
lected from habitats having a variety of cal- 
careous algae, sponges, anthozoans, and 
other sessile forms. It occurred in depths of 
less than 1 m on the Caicos Banks, 3—4 m 
in the back reef area, and 18 m on the outer 
fringing reef. Unfortunately, our collecting 
techniques were too general to determine if 
the species was associated with a specific 
sessile host. 

Distribution.—This species is presently 
known only from waters immediately ad- 
jacent to Pine Cay, Turks and Caicos Is- 
lands, British West Indies. 

Remarks.—The dichotomous key and Ta- 
ble 1 present some of the salient characters 
that distinguish these western Atlantic spe- 
cies of the subgenus Heteromysis. 


Key to western Atlantic species of the 
subgenus Heteromysis Smith, 1873 sensu 
Bacescu, 1968 
(species lacking flagellated spine on 
segment 3 of antennular peduncle and 
sexually dimorphic pleopods). 


1. Endopod of uropod with 12 or more spines 
along medial margin from region of statocyst 
1) EL HAIRS ARS rd ee ara Z 
—Endopod of uropod with no spines or one 
spine medially, near statocyst ......... = 
2. Margins of telsonic cleft with spinules along 
entire length; antennal scale 3 times or less 
as long as maximum width .......... 3 
—Margins of telsonic cleft with spinules 
only in apical part; antennal scale at least 3.5 
times as long as maximum width......... 
BNA 6103s Heteromysis elegans Brattegard, 1974 
3. Carpo-propodus of third thoracic endopod 
with 3 pairs of stout spine-setae on distome- 
SCS WiLL Ee ee Re Re ce na a 
.... Heteromysis formosa S. 1. Smith, 1873 
—Carpo-propodus of third thoracic endo- 
pod with no stout spine-setae on medial 
MURS PMN Rese ctl a Nag Mine es Oe A gers 'y. gs 
Heteromysis mexicana Escobar-Briones & 
Soto, 1990 
4. Endopod of uropod with no spine-setae me- 
dially, near statocyst 


Aas Ouse ee ere le,)ew te =, sw We fa, ee 


93 


....Heteromysis mureseanui Bacescu, 1986 
—Endopod of uropod with | medial spine- 
Sever ar SIMLOCYSE oo. vee ke ee ees 5 
5. Posterior half of lateral margins of telson 
armed with spine-setae; margins of telsonal 
cleft with spinules in apical half only ..... 
hy ast Heteromysis nouveli Brattegard, 1969 
—Lateral margins of telson armed with 
spine-setae along entire length; margins of 
telsonal cleft with spinules along entire 
length 


eis Wan ee eke 'e Shia” Gee bi Wen. fe ete) We “EOS eee ts se fw 


Heteromysis spottei appears most similar to 
H. nouveli, but differs in several aspects, espe- 
cially in the setation of the telson and third tho- 
racic endopod (see Key, Table 1). Except for H. 
(Heteromysis) waitei W. Tattersall, 1927, a 
South Australian species, the distinctive telson 
setation of H. spottei differs from all other de- 
scribed species of the subgenus Heteromysis. 
Several characters distinguish H. spottei from H. 
waitei including the presence of a single, instead 
of 3—4, spine-setae adjacent to the statocyst on 
the uropodal endopod and the absence of ocular 
teeth (see W. Tattersall 1927). 


Acknowledgments 


Support for the collection of specimens 
used in this study was sponsored by the 
Oakleigh L. Thorne Foundation through a 
grant to Stephen Spotte. We thank Oakleigh 
B. Thorne, members and employees of the 
Meridian Club, and the Turks and Caicos 
government for support and encourage- 
ment. We are grateful to Steve Spotte for 
his help, interest, and overall coordination 
of the field work for the faunal survey. Pa- 
tricia Bubucis, Jerry McLelland, and Steve 
Spotte assisted in the collection of speci- 
mens and provided other helpful assistance. 
The first author was supported through a 
University of Tampa Faculty Development 
Grant. 


Literature Cited 


Bacescu, M. 1968. Heteromysini nouveaux des eaux 
Cubaines: trois especes nouvelles de Hetero- 
mysis et Heteromysoides spongicola n.g., 

n.sp.——Revue Roumaine de Biologie, Serie de 


Zoologie 13:221-—237. 


94 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


. 1986. Heteromysis mureseanui n.sp. and Kal- 
liapseudes viridis, ssp. brasiliensis n. ssp., from 
the Brazilian littoral waters—Revue Roumaine 
de Biologie, Serie de Biologie Animale 31:93- 
OTs 

Brattegard, T. 1969. Marine biological investigations 
in the Bahamas. 10. Mysidacea from shallow 
water in the Bahamas and Southern Florida. 
Part I.—Sarsia 39:17—106. 

1974. Additional Mysidacea from shallow 
water on the Caribbean coast of Colombia.— 
Sarsia 57:47-86. 

Escobar-Briones, E., & L. A. Soto. 1990. Heteromysis 
mexicana, a new species from Campeche Bank, 
Gulf of Mexico (Crustacea: Mysidacea).—Pro- 
ceedings of the Biological Society of Washing- 
ton 103:131-139. 

Heard, R. W., & S. Spotte. 1991. Pontoniine shrimps 
(Decapoda: Caridea: Palaemonidae) of the 
northwest Atlantic. II. Periclimenes patae, new 
species, a gorgonian associate from shallow reef 
areas off the Turks and Caicos Islands and Flor- 
ida Keys.—Proceedings of the Biological So- 
ciety of Washington 104:40—48. 

, & . 1997. Pontoniine shrimps (Deca- 

poda: Caridea: Palaemonidae) of the northwest 

Atlantic. V. Periclimenes mclellandi, new spe- 

cies, a gorgonian associate from Pine Cay, 

Turks and Caicos Islands.—Proceedings of the 

Biological Society of Washington 110:39—48. 

, & P.M. Bubucis. 1993. Pontoniine 
shrimps (Decapoda: Caridea: Palaemonidae) of 
the northwest Atlantic. III. Neopericlimenes 
thornei, new genus, new species, from Pine 
Cay, Turks and Caicos Islands, British West In- 
dies.—Journal of Crustacean Biology 13:793-— 
800. 

Kensley, B., & R. W. Heard. 1991. Studies on the 
Crustacea of the Turks and Caicos Islands, Brit- 
ish West Indies. I. Four new marine isopod crus- 
taceans from the vicinity of Pine Cay.—Gulf 
Research Reports 8:237—246. 

Schotte, M., & R. W. Heard. 1991. Studies on the 


Crustacea of the Turks and Caicos Islands, Brit- 
ish West Indies. II. A new species, Armadillon- 
iscus stepus, (Isopoda: Oniscidea: Scyphacidae) 
from Pine Cay.—Gulf Research Reports 8:247— 
250. 


, & B. Kensley. 1991. Studies on the 
Crustacea of the Turks and Caicos Islands, Brit- 
ish West Indies. HI. Records of marine Isopoda 
from Pine Cay, Ft. George Cay, Water Cay, and 
adjacent waters.—Gulf Research Reports 8: 
251-257. 

Smith, S. I. 1873. Systematic catalogue of the inver- 
tebrates of the southern New England and ad- 
jacent waters. Crustacea: Pp. 545-580 in A. E. 
Verrill, Report upon the invertebrate animals of 
Vineyard Sound and the adjacent waters, with 
an account of the physical characters of the re- 
gion: Pp. 295-778 in S. FE Baird, Report on the 
condition of the sea fisheries of the south coast 
of New England in 1871 and 1872.—United 
States commission of Fish and Fisheries 7:1— 
852. 

Spotte, S., & P. M. Bubucis. 1996. Diversity and abun- 
dance of caridean shrimps associated with the 
slimy sea plume (Pseudopterogorgia ameri- 
cana) at Pine Cay, Turks and Caicos Islands, 
British West Indies.—Marine Ecology Progress 
Series 113:229-—232. 

, R. W. Heard, & P. M. Bubucis. 1994. Ponto- 
niine shrimps (Decapoda: Caridea: Palaemoni- 
dae) of the northwest Atlantic. IV. Periclimenes 
antipathophilus new species, a black coral as- 
sociate from the Turks and Caicos Islands and 
Eastern Honduras.—Bulletin of Marine Science 
55:212—227. 

Tattersall, W. M. 1927. Australian opossum shrimps 
(Mysidacea).—Records of the South Australian 
Museum 3:235-—257. 

Watling, L. 1989. A classification system for crusta- 
cean setae based on the homology concept. Pp. 
15-26 in B. E. Felgenhauer, L. Watling & A. 
B. Thistle, eds., Crustacean Issues 6, Functional 
morphology of feeding and grooming in Crus- 
tacea, A. A. Balkema, Rotterdam, 225 pp. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


113(1):95-103. 2000. 


Gynodiastylis laciniacristatus, a new species (Crustacea: Cumacea) 
from Australia 


Sarah Gerken and Jennifer Gross 


Darling Marine Center, School of Marine Sciences, University of Maine 
Walpole, Maine 04573, U.S.A. 


Abstract.—A new species, Gynodiastylis laciniacristatus (Crustacea: Cuma- 
cea) is described from two disparate localities on the Australian shelf, the Bass 
Strait (Southeast) and the Dampier Archipelago (West), from depths of 48—85 
m. The ovigerous female is completely figured. No males were collected. The 
new species is distinguished from all other species of Gynodiastylis by a large 
blunt process on the ischium of pereopod 4 and by the pattern of toothed ridges 
on the carapace. G. mutabilis Hale, 1946 and G. ornata Hale, 1946 have similar 
ridge patterns, however G. laciniacristatus has a distinct sharp corner on the 
anterolateral prominence of the carapace and rudimentary exopods on pereopods 
3 and 4, in addition to the large blunt process on the ischium of pereopod 4. 


The genus Gynodiastylis is known pri- 
marily from the Southern hemisphere. Hale 
(1946) described the great majority of the 
species in the genus, from the environs of 
Australia. A new and unusual species of 
Gynodiastylis was identified from the ex- 
tensive cumacean collections of the Muse- 
um of Victoria and is described here, as part 
of an NSF PEET (Partnerships for Enhanc- 
ing Expertise in Taxonomy) project. 


Methods 


Samples were collected with the WHOI 
epibenthic sled or a Smith-MclIntyre grab. 
Drawings were prepared using a camera lu- 
cida on a Wild compound microscope. 
Body length was measured from the tip of 
the pseudorostral lobes to the posterior bor- 
der of the last somite. One ovigerous fe- 
male was prepared for SEM (scanning elec- 
tron microscope) photography. 


Family Gynodiastylidae Stebbing 1912 
Gynodiastylis Calman 1911 
Gynodiastylis laciniacristatus, new species 
Figs. 1-5 

Type material.—Holotype (NMV J45433) 
1 ovigerous female, Bass Strait, Australia, 


39°49.0'S, 143°24.0’E, 56 m, 20 Nov 1981. 
Paratype (NMV J45434) | ovigerous °, 
Bass-> “Strait, — Australia,’ °39°49;0’S, 
143°24.0'E, 56 m, 20 Nov 1981. Paratypes 
(NMV J45435) 1 ovigerous 2, 1 subadult 
oo Bass, Strait) “Austral! 39°49 O'S, 
143°24.0’E, 56 m, 20 Nov 1981. Paratype 
(NMV J45436) 1 ovigerous °, Bass Strait, 
Australia, 38°38.2’S, 142°35.0’E, 59 m, 20 
Nov 1981. Paratype (NMV J45437) 1 ovig- 
erous 2, Bass Strait, Australia, 39°06.3’S, 
142°55.6’E, 81 m, 21 Nov 1981. Paratype 
(NMV J45438) 1 ovigerous ¢, Bass Strait, 
Australia, 40°00.0’S, 144°20.9’E, 48 m, 22 
Nov 1981. Paratype (NMV J45439) 1 ovig- 
erous 2, Bass Strait, Australia, 39°00.2'S, 
144°33.9'E, 74 m, 23 Nov 1981. Paratype 
(NMV J45440) 1 subadult 2, Bass Strait, 
Australia, 39°13:6 S; 143°55:6'E; ‘85m, 23 
Nov 1981. Paratype (NMV J45441) 1 sub- 
adult ¢, Western Australia, (damaged), 
20°1.00’S, 117°11.00’E, 48 m, 11 Jun 1983. 
Paratype (NMV J45442) 1 subadult 9°, 
Western Australia, 19°38.00’S, 118°6.00’E, 
49 m, 13 Jun 1983. 
Diagnosis.—Carapace with depression 
sweeping dorsally from anterior edge, an- 
terolateral prominence produced as sharp 


96 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


corner, with dorsal ridge bounding promi- 
nence toothed, dorsal depression bounded 
by sharp lateral ridges, continuing through 
dorsum of pereon 5, anterior ventral margin 
toothed, antennal notch oblique. Pereopod 
4 with large blunt process on ischium. 
Adult females less than 4 mm. 

Description.—Ovigerous female, 3.5 
mm. Carapace with dorsal depression, eye- 
lobe present, no lenses; pseudorostral lobes 
0.5 carapace length; antenna 1 visible, ex- 
tending just past pseudorostral lobes. Per- 
eonites 1—3 expanded ventrally as anteriorly 
directed flap, pereonites 4 and 5 directed 
posteriorly dorsally (Figs. 1A, 1B, 2A). 

Antennule article 1 longer than articles 2 
and 3 together, bearing 1 plumose and 2 
simple setae proximally and 1 simple seta 
distally; distomedial margin with fine hair- 
like setae; article 2 bearing 1 simple and 1 
plumose setae distally, medial margin bear- 
ing fine hair-like setae; articles 2 & 3 sub- 
equal in length, article 3 bearing 2 simple 
setae; main flagellum of 2 articles bearing 
1 long annulate seta, 2 long simple setae, 
and 2 short simple setae terminally; acces- 
sory flagellum of 1 article with 2 terminal 
setae (Fig. 3A). 

Mandible navicular, with 10 lifting setae; 
left side with stout lacinia mobilis, incisor 
quadridentate, right side with slender laci- 
nia mobilis, incisor bidentate (Fig. 3B). 

Maxillule of 2 lobes, outer broad lobe 
bearing double row of stout setae terminal- 
ly, medial margin bearing fine hair-like se- 
tae; inner lobe bearing 2 tridentate, | long, 
2 short setae terminally, medial margin 
bearing fine hair-like setae (Fig. 3C). 

Maxilla of 3 lobes; broad lobe 4 times as 
wide as narrow lobe, medial margin bearing 
9 simple setae, medial-distal corner bearing 
2 plumose setae, distal margin bearing 
many simple setae; distal-lateral corner 
bearing | forked and 1 microserrate setae; 
inner narrow lobe bearing 2 microserrate 
and 2 simple setae apically; outer narrow 
lobe bearing 5 simple setae apically (Fig. 
3D). 

Maxilliped 1 basis as long as all other 


articles together, produced as large disto- 
medial lobe, lobe bearing several short sim- 
ple setae distally, lobe bearing several plu- 
mose setae, setulose on distal 0.3 only; is- 
chium not present, merus half as long as 
carpus, medial margin bearing fine hair-like 
setae; carpus bearing 3 blade-like, 3—4 bi- 
dentate, several simple setae medially, 1 
long plumose seta distolaterally; propodus 
as long as carpus, half as broad, bearing 2 
tridentate and 2 long plumose setae distally, 
margins with many fine hair-like setae; dac- 
tyl half as long and half as wide as propo- 
dus, bearing 4 small simple setae terminally 
(Fig. 5C). 

Maxilliped 2 basis as long as next 2 ar- 
ticles together, bearing 3 long plumose se- 
tae distally; ischium not present; merus 0.5 
basis length, bearing 2 plumose setae dis- 
tally; carpus slightly longer than propodus, 
bearing 3 plumose setae medially; propodus 
bearing 2 long plumose setae distally and 3 
plumose setae medially; dactyl half length 
of propodus, bearing 4 simple setae termi- 
nally; endite bearing 5 stout annulate setae 
and 2 simple setae (Fig. 2B). 

Maxilliped 3 basis 2 times as long as 
next 4 articles together, medial margin bear- 
ing 8 plumose setae, lined with short hair- 
like setae, single plumose seta distally, dis- 
tomedial corner produced as 2 teeth, lateral 
margin lined with fine hair-like setae, dis- 
tolateral corner bearing 4 long plumose se- 
tae; ischium 0.5 basis width, slightly longer 
than merus, 1 plumose setae medially, oth- 
erwise margins lined with fine hair-like se- 
tae; merus 0.5 length carpus, produced as 
tooth at distolateral corner and bearing 1 
plumose seta, medial margin lined with fine 
hair-like setae and bearing | plumose and 1 
simple setae distally; carpus bearing 1 plu- 
mose seta on distolateral corner, medial 
margin lined with fine hair-like setae and 
bearing 1 plumose and 1 simple setae; pro- 
podus subequal to dactyl, bearing | simple 
seta on distolateral corner, medial margin 
bearing 2 simple setae; dactyl bearing 4 
simple setae terminally (Fig. 3E). 

Pereopod 1 basis as long as all other ar- 


VOLUME 113, NUMBER 1 97 


eee B 


Fig. 1. Gynodiastylis laciniacristatus, new species. Ovigerous ¢ (not the same individual as Figs. 2—5); A, 
habitus; B, dorsal view. Scale bars are 200 microns in length. 


ticles together, posterior margin bearing fine hair-like setae, anterodistal corner bear- 
many simple and 2 plumose setae, postero- ing 2 plumose setae; ischium 0.5 length of 
distal corner produced as 3 teeth, anterior merus, posterodistal corner produced as 
margin bearing 1 simple seta, lined with tooth; merus produced as 3 small teeth on 


98 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


a 
a 
A 
a 
A 
a 
A 
a 
a 
a 
A 
A 
a 
a 
a 
a 
a 
a 
a 
a 
a 


\t 
Pa 
) 


1 

wee 
so 

mo 


viv 
D earn nn null 


Fig 2. 


illiped 2; C, telson and uropods. 


Gynodiastylis laciniacristatus, new species. Ovigerous 2 paratype NMV 45434; A, habitus; B, max- 


VOLUME 113, NUMBER 1 99 


A, C,D 0.2mm 
B,E 0.5mm 


Fig. 3. Gynodiastylis laciniacristatus, new species. Ovigerous 2 paratype NMV 45434; A, antennule; B, 
mandible; C, maxillule; D, maxilla; E, maxilliped 3. 


i00 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


posterior margin; carpus subequal to pro- 
podus; propodus twice dactyl length, bear- 
ing | long and 1 short simple setae distally, 
single small simple seta proximally; dactyl 
bearing 1 long and 1 short simple setae ter- 
minally; exopod 0.5 length basis, basal ar- 
ticle bearing 2 simple setae, produced as 
small tooth distally; flagellum bearing 6 
long stout plumose setae (Fig. 4A). 

Pereopod 2 basis longer than all other ar- 
ticles together, 3 times ischium width, mar- 
gins lined with many simple setae, anterior 
margin produced as multiple teeth; ischium 
reduced, unarmed; merus twice carpus 
length, anterior margin produced as multi- 
ple teeth, bearing 2 simple setae; carpus an- 
terior margin produced as 2 teeth, bearing 
2 simple setae; propodus subequal to car- 
pus, unarmed; dactyl slightly longer than 
propodus, bearing 3 simple setae terminal- 
ly; exopod subequal to basis, basal article 
unarmed, flagellum bearing 4 long stout 
plumose setae (Fig. 4B). 

Pereopod 3 basis as long as next 2 arti- 
cles together, anterior margin produced as 
multiple teeth, bearing 5 plumose, 1 annu- 
late and 2 annulate plumose setae, posterior 
margin bearing 2 simple setae; ischium 0.2 
length merus, bearing 1 simple seta; merus 
twice carpus length, bearing 2 annulate and 
4 simple setae; carpus twice propodus 
length, bearing 6 annulate and 2 simple se- 
tae distally; propodus twice dactyl length, 
bearing single annulate seta distally; dactyl 
bearing 2 simple setae terminally; exopod 
biarticulate, rudimentary, bearing 1 simple 
seta apically (Fig. 4C). 

Pereopod 4 coxa bearing several stout 
simple setae; basis as long as next 2 articles 
together, anterior margin bearing 2 plu- 
mose, 1 annulate plumose, 7 annulate, and 
3 simple setae, produced as multiple teeth 
distally, 1 plumose seta mid article; ischium 
0.3 length merus, unarmed, produced as 
large posteriorly directed blunt lobe; merus 
3 times carpus length, bearing 7 annulate 
setae, produced as multiple scales, posterior 
margin produced as 3 teeth; carpus slightly 
longer than propodus, bearing 7 annulate 


setae, produced as 2 teeth proximally; pro- 
podus slightly longer than dactyl, bearing 2 
annulate setae; dactyl bearing single stout 
seta with single setule; exopod biarticulate, 
rudimentary, bearing | simple seta apically 
(Fig. 5A). 

Pereopod 5 basis as long as next 2 arti- 
cles together, bearing 6 plumose, | annulate 
plumose and | annulate setae, produced as 
small blunt processes on posterior margin; 
ischium 0.5 length merus, bearing | simple 
seta; merus slightly longer than carpus, 
bearing 5 simple and 1 annulate setae, pos- 
terior margin produced as scales; carpus 
bearing 7 annulate setae; propodus subequal 
to carpus, bearing 1 annulate seta; dactyl 
0.5 length propodus, bearing 1 stout and 1 
slender setae terminally (Fig. 5B). 

Telson equal in length to pleonite 6, post- 
anal section negligible, unarmed (Fig. 2C). 

Uropod peduncles equal in length to tel- 
son, bearing single seta at distomedial cor- 
ner; rami shorter than peduncles; endopod 
biarticulate, article 1 bearing 1 short stout 
seta at distomedial corner, article 2 bearing 
2 short stout setae medially, single long seta 
terminally; exopod biarticulate, article 1 
half length article 2, unarmed, article 2 
bearing 2 short setae laterally, single long 
stout seta terminally; all setae on rami bear- 
ing single thick setule apically (Fig. 2C). 

Etymology.—laciniacristatus from the 
Latin lacinia, meaning jagged and crista, 
meaning ridge. 

Remarks.—Gynodiastylis laciniacrista- 
tus is distinguished from all other Gyno- 
diastylis by the large blunt process on the 
ischium of pereopod 4. Gynodiastylis laci- 
niacristatus is superficially similar to both 
G. ornata Hale, 1946 and G. mutabilis 
Hale, 1946. However, there are obvious dif- 
ferences in addition to the process on pe- 
reopod 4. Neither G. ornata nor G. muta- 
bilis have exopods on pereopods 3 and 4 of 
the female, while they are present in G. la- 
ciniacristatus. The uropod endopod in G. 
mutabilis is uniarticulate, while in G. laci- 
niacristatus the uropod endopod is biartic- 
ulate. The lateral margins of the telson in 


VOLUME 113, NUMBER | 10] 


A,B, C 0.5mm 


Fig. 4. Gynodiastylis laciniacristatus, new species. Ovigerous paratype NMV 45434; A, pereopod 1; B, 
pereopod 2; C, pereopod 3. 


102 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


C 0.2 mm 
A, B, 0.5 mm 


Fig. 5. Gynodiastylis laciniacristatus, new species. Ovigerous 2 paratype NMV 45434; A, pereopod 4; B, 
pereopod 5; C, maxilliped 1. 


VOLUME 113, NUMBER 1 


G. mutabilis are produced as at least one 
pair of teeth, while in G. laciniacristatus 
the telson lateral margins are entire. The 
carapace of G. laciniacristatus has a dis- 
tinct anterolateral prominence produced as 
a sharp corner, while G. ornata has no such 
prominence. 


Acknowledgments 


The Museum of Victoria kindly loaned 
the material used in the study. The National 
Science Foundation (Grant DEB95-21783, 
L. Watling, I. Kornfield PI.’s) supported 
this research as part of the Partnerships to 
Enhance Expertise in Taxonomy (PEET) 


103 


program. The Gulf of Maine Foundation 
supported this research as a Summer Un- 
dergraduate Research Experience (SURE) 
for Jennifer Gross. 


Literature Cited 


Calman, W. T. 1911. On new or rare Crustacea of the 
Order Cumacea from the collection of the Co- 
penhagen Museum, Part IJ.—Transactions of 
the Zoological Society of London 18:341—399. 

Hale, H. 1946. Australian Cumacea, No. 12, the Fam- 
ily Diastylidae (part 2) Gynodiastylis and relat- 
ed genera.—Records of the South Australian 
Museum 8(3):357—444. 

Stebbing, T. R. R. 1912. The sympoda, Part 6.—An- 
nals of the South African Museum 10:129—176. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


113(1):104—123. 2000. 


Revision of the subterranean amphipod genus Spelaeogammarus 
(Bogidiellidae) from Brazil, including descriptions of three new 
species and considerations of their phylogeny and biogeography 


Stefan Koenemann and John R. Holsinger 


Department of Biological Sciences, Old Dominion University, 
Norfolk, Virginia 23529-0266, U.S.A. 


Abstract.—Three new subterranean amphipods of the genus Spelaeogam- 
marus da Silva Brum, 1975, are described from eastern Brazil, bringing the 
total number of species in the genus to four. Based on the examination of type- 
material of Spelaeogammarus bahiensis, a comparative diagnosis of all four 
species of Spelaeogammarus, including the new species Spelaeogammarus spi- 
nilacertus, Spelaeogammarus trajanoae, and Spelaeogammarus santanensis, 1s 
given. A diagnosis for the genus is also provided. The occurrence of these 
species in caves that are separated from each other in discontinuous karst areas 
is biogeographically significant for the family Bogidiellidae in continental 


South America. 


Exploration of caves in eastern Brazil in 
the early 1970s resulted in the discovery of 
the new bogidiellid genus and species Spe- 
laeogammarus bahiensis (da Silva Brum, 
1975). The specimens were collected from 
a cave near Curaca, capital of the district 
Matamuté, in the state of Bahia. Subse- 
quently, between 1989 and 1993, many ad- 
ditional specimens from various caves in 
Bahia were collected by Brazilian speleol- 
ogists and sent to us for identification. Be- 
cause these specimens differed morpholog- 
ically from the description of Spelaeogam- 
marus bahiensis, we borrowed paratypes of 
this species from the Museu Nacional in 
Rio de Janeiro, for a comparison with the 
new material. The paratypes enabled us to 
identify and describe three new species and 
also to diagnose the genus Spelaeogam- 
marus. In addition, a key to the four species 
of the genus is provided as well as a table 
detailing morphological differences. 

The holotypes of the new species are de- 
posited in the Museu Nacional in Rio de 
Janeiro (MNRJ), Brazil, as indicated. 


Genus Spelaeogammarus da Silva Brum, 
1975 


Spelaeogammarus da Silva Brum, 1975: 
125-128. 

Type species (by monotypy): Spelaeo- 
gammarus bahiensis da Silva Brum, 1975. 

Diagnosis.—Eyes absent. Body smooth, 
unpigmented. Coxal plates 1—2 small, wider 
than long; plates 3-6 longer than wide, 
overlapping. Antenna 1 about 45-50% of 
body length, primary flagellum longer than 
peduncle, with 16—20 segments. Accessory 
flagellum with 4—5 segments. Antenna 2 
flagellum bearing 7—10 segments. Mandib- 
ular palp 3-segmented. Maxilla 1 with sym- 
metrical, 2-segmented palp; inner plate with 
3 plumose setae; outer plate bearing 6—7 
serrate spines. Inner plate of maxilliped 
bearing apically 2 bifid (y-shaped) spines; 
outer plate with 3 or 4 blade-like spines api- 
cally and subapically. Propodus of gnatho- 
pod 1 larger than that of gnathopod 2. Dac- 
tyls of both gnathopods distinctly serrate 
along inner margins. Pereopods without any 
trace of lenticular organs; pereopods 5—7 


105 


VOLUME 113, NUMBER 1 


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106 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


bases broad, propodus and/or carpus with 
long, bifurcate setae. Pleopods and uropods 
unmodified. Pleopods biramous, with 3- 
segmented outer ramus and 1-segmented in- 
ner ramus, rami subequal in length. Uro- 
pods biramous: peduncle of uropod | with 
3 or 4 large basiofacial (ventrolateral) 
spines; uropod 3 with subequal, 1-segment- 
ed rami, outer ramus bearing a row of long, 
bifurcate setae along medial margin. Telson 
typically longer than wide, apex with shal- 
low excavation, bearing apical and subapi- 
cal spines. Coxal gills present on pereopods 
4—6. Oostegites linear and elongate, on pe- 
reopods 2—5. No sexual dimorphism in any 
characters. 


Key to the species of Spelaeogammarus 
(based on males and females) 


1. Accessory flagellum of antenna 1 with 4 seg- 
ments; flagellum of antenna 2 with 7 seg- 
ments; propodus of gnathopod | slightly larg- 
Cr ‘thai DasiS <a ges epee ee oe 2 

Accessory flagellum of antenna 1 with 5 
segments; flagellum of antenna 2 with 8—10 
segments; propodus of gnathopod 1 much 
larger than basis’ #yn.-. 6... .. S. santanensis 

2. Coxal plate 6 bearing 1 spine, setae absent; 
outer plate of maxilla 1 with 6 serrate spines 
and | plumose seta; inner plate of maxilliped 
with. 4; plumose.setae... . 2: .Naajun. 52 3 

Coxal plate 6 bearing 1 spine and about 20 
setae; outer plate of maxilla 1 with 7 serrate 
spines; inner plate of maxilliped without plu- 
MOSE Sta 92 4. Bon eke kh eee S. bahiensis 

3. Anterior margin of basis of gnathopod 1 with 
5—9 short setae; coxal plate 5 bearing 1 spine 
and 17-18 setae S. trajanoae 

Anterior margin of basis of gnathopod 1 
with 2—4 spines (or longer setae) and 1 short 
seta; coxal plate 5 bearing 1 spine and about 
O SSCAAG oa aa oes Oo eae S. spinilacertus 


© wr 6; © 16 @ 6 te 8, ae 


Spelaeogammarus spinilacertus, new 
species 
Figs. 1-5, 6c 


Material examined.—Estado do Bahia, 
Brazil: holotype male (6.1 mm) and 1 para- 
type male (7.5 mm) from Baixa do Salitre 


Cave, Iraquara, collected by J. A. Cardoso, 
Sep 1993; 4 paratypes (3 females, 1 male) 
from Baixa do Salitre Cave, collected by L. 
Mendes & E. Rubbioli, 24 May, 1997; al- 
lotype female (8.1 mm) and 2 paratypes (1 
male, 1 female), from Jaburu Cave, Ira- 
quara, collected by J. A. Cardoso, Sep 
1993" 

The holotype and allotype are dissected 
and mounted on microscope slides in 
Faure’s medium. Holotype (MNRJ 13340) 
and allotype are deposited in the Museu Na- 
cional. The remaining 7 paratypes are re- 
tained in the research collection of the sec- 
ond author. 

Diagnosis.—Spelaeogammarus spinila- 
certus is easily distinguished from other 
species in the genus by 2—4 spines or setae 
of corresponding length on the distoanterior 
margin of basis of gnathopod 1. Largest 
male 10.5 mm, largest female 10 mm. 

Description.—Antenna | (Fig. 2a) about 
45-50% of body length. Peduncular seg- 
ments 1—3 gradually decreasing in length; 
peduncular segment 1 with 3—4 spines; pe- 
duncular segment 2 with 2—3 short spines; 
peduncular segment 3 bearing O-—2 spines. 
Primary flagellum longer than peduncle, 
with 17—20 segments; aesthetascs on most 
segments; accessory flagellum with 4 seg- 
ments, terminal segment vestigial. 

Antenna 2 (Fig. 2b) about 85% length of 
antenna 1. Peduncular segment 4 longer 
than peduncular segment 5; peduncular seg- 
ment 5 with 3—4 ventral and 2—4 medial 
spines. Flagellum with 7 segments, Ist seg- 
ment twice as long as average length of 
segments 2-6. 

Upper lip (Fig. 2c) as long as broad, trap- 
ezoidal, with few apical setules. 

Mandible (Fig. 2h, 1): Palp 3-segmented, 
with 3—4 terminal setae; 2nd segment bear- 
ing 2 apical and 2-3 subapical setae. Molar 
rounded and well developed, with 1 long 
lateral seta. Both incisor and lacinia mobilis 
on left mandible with 5 irregular, rounded 
cusps (Fig. 2h); 3 long and 3 short plumose 
spines between lacinia and molar. Right 
mandible (Fig. 21): lacinia apically serrated, 


VOLUME 113, NUMBER 1 


consisting of irregularly pointed denticles; 
2 long and 2 short plumose spines between 
lacinia and molar. 

Lower lip (Fig. 2d) bearing setules on 
outer lobes, outer and mandibular lobes 
with rounded corners. 

Maxilla 1 (Fig. 2e): Palp 2-segmented, 
with 5—6 apical setae and few lateral set- 
ules. Outer plate with 6 serrate spines and 
1 plumose spine; inner plate with 3 plumose 
setae. 

Maxilla 2 (Fig. 2f): Outer plate apically 
with 1—2 comb-like setae, +15 medium- 
sized plumose setae and 2 large plumose 
spines (slightly subapical); medial margin 
with few fine setules; apical margin of inner 
plate bearing +18 long comb-like setae and 
+5 short naked setae (seta/spine types in 
Fig. 2g). 

Maxilliped (Fig. 2j, 6c): Segment 1 of 
palp with 1—2 medial setae; segment 2 bear- 
ing 12-13 medial setae; dactyl long and 
slender, bearing a row of marginal setules. 
Inner plates apparently fused along medial 
margins. 

Gnathopod | (Fig. 3a): Posterior margin 
of basis with 6—8 long setae (some doubly 
inserted); anterior margin bearing 2—4 
spines plus 1 short seta. Carpus with 8 setae 
on posterior lobe (4—6 comb-like and 2-3 
naked). Propodus ovate, almost twice as 
long as broad, larger than gnathopod 2 pro- 
podus; palm uneven, serrate with minute se- 
tules at corner (Fig. 3b); palmar margin 
bearing 7—8 normal spines and 15—19 short 
bifid spines on lateral margin; medial mar- 
gin with 6 short setae, 1 normal angular 
spine, and 4—6 oblique subangular spines 
(1-2 relatively long). Dactyl about 70% 
length of propodus; inner margin with dis- 
tinct row of denticles (Fig. 3c). 

Gnathopod 2 (Fig. 3d): Posterior margin 
of basis bearing 9-10 long setae. Carpus 
posteriorly with fine setules and 7 sets of 
setae (1—5 setae per set). Propodus ovate, 
almost twice as long as broad; palm 
oblique, with 5—6 corner spines (Fig. 3e), 
11-14 short lateral bifid spines, and 7-8 
short medial setae; palmar margin finely 


107 


serrate at whole margin, with minute setules 
at corner. Dactyl about 50% length of pro- 
podus; inner margin with distinct row of 
denticles. 

Pereopods 3 and 4 subequal (Fig. 4a). 
Basis without spines, anterodistal margin 
even (pereopod 4 basis with O—1 spine plus 
1 seta at anterodistal margin). Posterior 
margin of carpus bearing 4—5 spines. Pro- 
podus with 8—9 spines along posterior mar- 
gin and 2 apical spines. Dactyl about 24% 
length of propodus (Fig. 4b). 

Pereopod 5 (Fig. 4c): Basis with 10-11 
spines at posterior margin (distal and prox- 
imal group of spines separated by a gap); 
anterior margin bearing 13-15 spines; an- 
terior lateral surface with 6 short setae; 10— 
11 short setae at posterior margin and pos- 
terior lateral surface. Ischium with 1 spine 
and 3 setae. Anterior margin of carpus with 
a row of long, bifurcate setae (Fig. 4f) and 
6—12 spines (some doubly inserted). Pro- 
podus anteriorly with a row of long, bifur- 
cate setae, occurring progressively shorter 
distally, with slightly thicker bases; lateral 
margin with 17—19 spines (some doubly in- 
serted); proximal part with 3—4 spines. Dac- 
ty] 14—20% length of propodus. 

Pereopod 6 subequal to pereopod 5 but 
slightly longer. 

Pereopod 7 (Fig. 4d): Basis ovate, bear- 
ing 8 spines on anterior margin and 7—9 
spines on posterior margin. Ischium with 2 
spines. Merus with 3 spines on posterior 
margin and 5 spines at anterior margin (1 
singly and 2 doubly inserted). Carpus with 
13 spines, occurring in 6—7 sets (with 1-3 
spines per set) on anterior margin, 4 spines 
(2 doubly inserted) plus 4 setae on posterior 
margin, and 10—11 terminal spines. Propo- 
dus bearing 12 slender spines on anterior 
margin and rows of long, bifurcate setae 
along posterior and anterior margins (Fig. 
4f). Dactyl about 27% length of propodus. 

Pereopods 1—7 without any trace of len- 
ticular organs; pereopods 5—7 with broad 
bases. 

Coxal gills ovate, present on pereopods 
4-6. 


108 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fie: 
Brazil. 


Coxal plates 1 and 2 (Fig. 3f, g) about 
equal in size, subrectangular to ovate in 
shape, wider than long; plates 3—6 longer 
than wide, overlapping; plates 3 and 4 
about the same size, plate 5 largest; plate 5 
and 6 with distinct anterodistal lobes (Fig. 
4c, g); plate 7 (Fig. 4e) subtriangular to ir- 
regular in shape, with single long spine at 
tapered posterior corner. 

Oostegites linear and elongate, on pereo- 
pods 2-5. 

Pleopods 1—3 morphologically alike (Fig. 
5a), decreasing slightly in size posteriorly, 
with subequal outer and inner rami. Outer 
rami 3-segmented, with 2 terminal plumose 
setae per segment; segment 1 bearing 6—10 
lateral plumose setae and 5—8 medial plu- 
mose setae. Inner rami 1-segmented, with 
4—5 medial plumose setae. 

Epimeral (pleonal) plates subquadrate, 
with small, subacute posterior margins, 
bearing 1 setule each (Fig. 4h). 

Uropod 1 (Fig. 5b): Rami subequal in 
length, slightly shorter than peduncle. Pe- 
duncle with 3 spines on dorsolateral, dor- 


Spelaeogammarus spinilacertus n. sp., allotype female (8.1 mm) from Jaburu Cave, Estado do Bahia, 


somedial, ventrolateral (basofacial), and 
apical margin, respectively. Outer ramus 
bearing 3—4 dorsolateral and 4 apical 
spines. Inner ramus with 5 apical and 4—5 
dorsolateral spines, the latter occurring as 
3—4 singly and 1—2 doubly inserted. 

Uropod 2 (Fig. 5c): Peduncle bearing 1 
dorsomedial spine, 2 dorsolateral spines, 
and 2 apical spines. Outer ramus slightly 
shorter than inner ramus, as long as pedun- 
cle, bearing 2—3 spines dorsolaterally and 4 
spines apically. Inner ramus with 5 dorso- 
lateral and 4—5 apical spines. 

Uropod 3 (Fig. 5d) with subequal, lan- 
ceolate rami, both 1-segmented. Peduncle 
about 46% length of rami, with 2 apical 
spines, 1 subapical spine, and 1 small dor- 
soproximal spine. Outer ramus bearing 3 
apical spines, 6 sets of spines (with 2-3 
spines per set) along lateral margin, and 
about 20 bifurcate long setae along medial 
margin (Fig. 5e). Inner ramus with 3 apical 
spines; lateral margin with 5 spines; medial 
margin bearing 9 spines (some doubly in- 
serted). 


VOLUME 113, NUMBER 1 


109 


Pig...2. 
lower lip, e) maxilla 1, f) maxilla 2, g) spine and seta types (on maxilla | and 2, maxilliped, and gnathopod | and 
2), from left: serrate spine, comb-like seta, plumose seta, and plumose spine, h) left mandible, 1) right incisor and 
lacinia mobilis, j) maxilliped, k) spine types of maxilliped outer plate (far left 2) and inner plate (far right 2). 


Telson (Fig. 5f) width about 84% of 
length, with shallow excavation (10% of 
length); each side bearing 3 setae, 2 apical 
and 2 (sometimes 3) subapical spines. 

Etymology.—tThe epithet spinilacertus is 
a noun in apposition, alluding to the pres- 


& 


VODA 


Spelaeogammarus spinilacertus n. sp., allotype female: a) antenna 1, b) antenna 2, c) upper lip, d) 


ence of spines on the anterior margin of the 
basis of gnathopod 1. It is formed by com- 
bining spini, from Latin meaning thorn or 
spine, with lacertus, from Latin meaning 
upper arm (~basis). 

Remarks.—Two of the three specimens 


110 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 3. Spelaeogammarus spinilacertus n. sp., holotype male: a) left gnathopod 1, b) detail of right gnathopod 
1, c) detail of right gnathopod | dactyl, d) left gnathopod 2, e) detail of left gnathopod 2; allotype female: f) 
coxal plate of gnathopod 1, g) coxal plate of gnathopod 2. 


from Jaburu cave showed variation in some 
characters, one of which is diagnostic for 
the species: instead of 2—4 spines, the basis 
of the female gnathopod 1 had 2-3 setae 


along the anterodistal margin (compare Fig. 
3a with Fig. 7a and c). These setae had the 
same length as the spines they replaced and 
could be clearly distinguished from corre- 


VOLUME 113, NUMBER 1 


11] 


Fig. 4. Spelaeogammarus spinilacertus n. sp., allotype female: a) left pereopod 4, b) right pereopod 4 dactyl, 
c) left pereopod 5, d) right pereopod 7, e) coxal plate of right pereopod 7, f) bifurcate seta (on carpus and 
propodus of pereopod 5-7), g) coxal plate of left pereopod 6, h) epimeral plates. 


sponding setae in S. santanensis and S. tra- 
jJanoae (described below). Furthermore, 
both specimens (10 mm male and 7 mm 
female) had 3 subapical spines on each lobe 
of the telson. The male from the Jaburu 
sample also showed morphological vari- 


ability in its appendages, e.g., the bases of 
both gnathopods and pereopods 3—6 were 
relatively narrow and elongate; similarly, 
the propods of both gnathopods appeared 
relatively longer and larger, with a conspic- 
uously sinusoid palmar margin. 


Hi PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


f 


Fig: 5: 


Spelaeogammarus spinilacertus n. sp., holotype male: a) left pleopod 3, b) left uropod 1, c) left 


uropod 2, d) left uropod 3, e) bifurcate seta of outer ramus of uropod 3, f) telson. 


Spelaeogammarus trajanoae, new species 
Fics. 6b;1, 7c)'d, Sa, b 


Material examined.—Campo Formoso, 
Estado do Bahia, Brazil: holotype female 
(10.4 mm) and 5 paratypes (2 males, 3 fe- 
males) from Toca do Pitu Cave (=Gruta do 
Pitu?), collected by A. Auler & M. Martins, 
Sep 1989; 1 male and | female paratype 
from Toca do Pitu Cave, collected by E. 
Rubbioli, Jan 1992; 3 paratypes (1 female, 
2 juveniles) from Buraco do Teodoro Cave, 
collected by J. A. Cardoso (no date given); 
1 female paratype from Toca do Gongalo 
Cave, collected by P. Gernhard, 4 Jul, 1997; 
1 fragmented specimen from Convento 
Cave, collected by S. Larizotti, 1986 (?). 

The holotype is dissected and mounted 
on microscope slides in Faure’s medium. 
Holotype (MNRJ 13341) and 2 paratypes 
from the type locality are deposited in the 
Museu Nacional. The allotype and the re- 


maining paratypes are retained in the col- 
lection of the second author. 

Diagnosis.—Spelaeogammarus traja- 
noae is morphologically closely allied with 
S. spinilacertus but can be distinguished 
from that species by 5—9 short setae instead 
of spines or spine-length setae on the dis- 
toanterior margin of the basis of gnathopod 
1 and coxal plate 5 with approximately 18 
setae plus | spine on margins instead of 9 
setae plus 1 spine. Largest males 10.0 mm, 
largest female 10.5 mm. S. trajanoae is 
moreover distinguished from S. spinilacer- 
tus as indicated in the following descrip- 
tion. 

Description.—Antenna 1: Peduncular 
segment 1 with 5—7 spines; peduncular seg- 
ment 2 with 3—4 short spines; peduncular 
segment 3 bearing 2 spines. 

Lower lip (Fig. 61) bearing setules on 
medial margin of outer lobes. 


VOLUME 113, NUMBER | 


Maxilliped: Segment 2 with +17 setae 
along medial margin. Inner and outer plate 
shown in Fig. 6b. 

Gnathopod 1 (Fig. 7c): Posterior margin 
of basis bearing 9-10 singly inserted, long 
setae; anterior margin with 5—9 short setae. 
No setules on posterior margin of ischium. 
Carpus bearing 6—9 setae on pointed pos- 
terior lobe (4—6 comb-like and 2—3 naked). 
Palm of propodus even; lateral margin with 
5—7 normal spines and 16—17 short bifid 
spines; medial margin with 6—8 short and 6 
long setae. 

Gnathopod 2: Posterior margin of basis 
with 8—9 long setae. Carpus posteriorly 
with 5—6 rows of setae (1—5 setae per row). 
Propodus with 6—8 rows of setae (1—3 setae 
per row) at proximoposterior margin; pal- 
mar margin with 3—4 lateral corner spines, 
16-17 short lateral bifid spines, and 8—9 
short medial setae. 

Pereopods 3 and 4 (Fig. 8a): Basis bear- 
ing | spine at posterodistal margin; antero- 
distal margin sinusoid, with 1 spine and 1 
seta (pereopod 3) or 2 spines and 1 seta 
(pereopod 4). Carpus bearing 6—7 spines 
posteriorly. Propodus with 13-15 spines 
along posterior margin. 

Pereopod 5: Coxal plate with 17—18 mar- 
ginal setae on anterior lobe. Anterior mar- 
gin of basis bearing 10—12 spines; 15 short 
setae on both posterior and anterior lateral 
surfaces. Dactyl about 22% length of pro- 
podus. 

Pereopod 6: Ischium with 2 spines and 
1-2 setae. Dactyl about 26% of propodus 
length. 

Pereopod 7: Basis with 11 spines on an- 
terior margin and 10 spines on posterior 
margin. Merus with 3 sets of doubly in- 
serted spines at posterior margin and 3 sets 
of spines at anterior margin (with 3—4 
spines per set). 

Pleopods: Outer ramus with 8—10 lateral 
plumose setae and 8—9 medial plumose se- 
tae on first segment. Inner ramus bearing 5— 
7 medial plumose setae. 

Epimeral plates (Fig. 7d) with produced, 
bluntly rounded distoposterior corners. 


113 


Uropod 1: Peduncle with 3—4 spines on 
dorsolateral margin, 4—5 spines on dorso- 
medial margin. Inner ramus with 4—5 mar- 
ginal spines, occurring as 4 dorsomedial 
and O—1 dorsolateral spines. 

Uropod 2: Peduncle with 1—2 dorsome- 
dial spines, 2—3 dorsolateral spines. 

Uropod 3: Outer ramus bearing 3-5 api- 
cal spines, 4—6 rows of spines (with 1-3 
spines per row) along lateral margin. Inner 
ramus bearing 10—11 rows of spines (with 
5 singly and 5 doubly inserted) at medial 
margin. 

Telson (Fig. 8b) width about 81% of 
length, with u-shaped apical excavation 
(19% of length); each side with 3 apical 
setae, 2 apical spines, and 3—4 subapical 
spines. 

Etymology.—The species is named in 
honor of Professor Dr. Eleonora Trajano of 
the University of Sao Paulo, who has made 
important contributions to Brazilian bio- 
speleology. 

Remarks.—Out of a total of 12 speci- 
mens examined, 2 individuals were found 
with 2 and 3 subapical spines on each tel- 
sonic lobe, respectively. The majority (10 
specimens) had 4 subapical spines on one 
lobe and 3 subapical spines on the other 
lobe. 


Spelaeogammarus santanensis, new 
species 
Figs. 6a, e—g, 7a, b, 8c—e 


Material examined.—Padre Cave, San- 
tana, Estado do Bahia, Brazil: holotype 
male (13.6 mm), 3 male and 3 female para- 
types (11.5—13.6 mm), and | juvenile para- 
type (10.6 mm), collected by KE Chaimow- 
icz, July 1987. 

The holotype is dissected and mounted 
on microscope slides in Faure’s medium. 
Holotype (MNRJ 13342) and 2 paratypes 
are deposited in the Museu Nacional. The 
remaining paratypes are retained in the col- 
lection of the second author. 

Diagnosis.—A comparatively large cav- 
ernicolous species, easily distinguished 


114 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 6. Spelaeogammarus santanensis n. sp., holotype male: a) inner and outer plate of maxilliped, e) detail 
of antenna 1, f) left mandible, g) detail of right mandible; S. trajanoae n. sp., paratype female (10.36 mm): b) 
inner and outer plate of maxilliped, i) lower lip; S. spinilacertus n. sp., holotype male: c) inner and outer plate 
of maxilliped; S. bahiensis paratype (8 mm juvenile): d) inner and outer plate of maxilliped, h) left incisor and 
lacinia mobilis. 


VOLUME 113, NUMBER 1 


115 


Fig. 7. 


Spelaeogammarus santanensis n. sp., holotype male: a) left gnathopod 1, b) right gnathopod 2; S. 


trajanoae n. sp., paratype female (10.36 mm): c) left gnathopod 1, d) epimeral plates. 


from other species in the genus by having: 
5-segmented accessory flagellum; 20—23 
setae on posterior margins of the bases of 
gnathopods 1 and 2; propodus of gnathopod 
1 proportionally larger. Largest male 13.6 
mm, largest female 10.5 mm. S. santanensis 


is furthermore distinguished from S. spini- 
lacertus according to the following descrip- 
tion. 

Description.—Antenna | about 40—45% 
length of body. Primary flagellum bearing 
20-—21 segments, some of which with mul- 


116 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 8. 


Spelaeogammarus trajanoae n. sp., paratype female (10.36 mm): a) left pereopod 3, b) holotype 


female: telson; S. santanensis n. sp., holotype male: c) right pereopod 3, d) coxal plate of pereopod 7, e) telson; 
S. bahiensis, paratype (8 mm juvenile): f) coxal plate of pereopod 7, g) telson. 


tiple inserted aesthetascs (Fig. 6e). Pedun- 
cular segment 1 with 5—6 spines; pedun- 
cular segment 3 with 2 ventromedial spines. 
Accessory flagellum 5-segmented. 

Antenna 2 slightly shorter than antenna 
1. Peduncular segment 5 with 5 ventral 
spines. Flagellum as long as peduncular 
segment 5, with 8—10 segments. 

Mandible (Fig. 6f, g): Palp with 3 ter- 
minal setae. Palp segment 2 of left mandi- 
ble with 3 setae and 1 spine; 3 short and 
1(—2?) long plumose spines between lacinia 


and molar. Palp segment 2 of right mandi- 
ble with 4 setae and 2 spines. 

Lower lip with few thin setules on medial 
margin of outer and inner lobes; corner of 
outer lobes with slightly pointed corners. 

Maxilla 1: Palp with 8—9 apical and sub- 
apical setae and a few lateral setules. Outer 
plate with 7 serrate spines apically. 

Maxilla 2: Outer plate apically with 0-1 
comb-like setae, +21 plumose setae, and 1— 
2 plumose spines (slightly subapical). Inner 
plate bearing +22 long comb-like setae and 


VOLUME 113, NUMBER 1 


several short naked setae (two of which on 
medial margin). 

Maxilliped: Palp segment 1 with 2 me- 
dial and 2 proximomedial setae; segment 2 
with +25 medial setae; segment 3 bearing 
+17 medial and 3 dorsomedial setae. Dac- 
tyl with 8 setae on outer margin. Inner and 
outer plate shown in Fig. 6a. 

Gnathopod | (Fig. 7a): Posterior margin 
of basis bearing 20 long, singly inserted se- 
tae; anterior margin with 4 setae. Carpus 
with distinctly pointed posterior lobe, 
which bears 13—14 plumose setae. Propo- 
dus bearing 7—8 setae on medial surface; 
palmar margin sinusoid, with 10 short me- 
dial setae, 4 long lateral setae, 8—9 normal 
lateral spines, 19-20 short bifid lateral 
spines, 1 normal angular spine medially, 2— 
3 oblique subangular spines (1—2 relatively 
long), +13 subangular lateral setae, and 6— 
7 subangular medial setae. Inner margin of 
dactyl with row of blunt denticles. 

Gnathopod 2 (Fig. 7b): Posterior margin 
of basis bearing 21—23 long setae. Carpus 
posteriorly with 8—9 rows of setae and 5 
short setae distolaterally. Propodus twice as 
long as broad; palm with 6 lateral spines 
and 1 medial spine, 18 short lateral bifid 
spines, 1 long lateral seta, and 12 short me- 
dial setae; palmar margin with blunt serra- 
tion of whole margin, distinct serration and 
minute cilia at corner. Dactyl about 53% 
length of propodus, inner margin with row 
of blunt denticles. 

Pereopod 3: Basis (Fig. 8c) with 2 spines 
at posteriodistal margin. Carpus bearing 5— 
6 spines posteriorly and 2-3 spines plus 1— 
2 setae anteriorly. Propodus with 11-12 
spines along posterior margin (some doubly 
inserted) and 2 spines plus 4 setae apically. 
Dactyl about 25% length of propodus. 

Pereopod 4 subequal to pereopod 3, ex- 
cept for the following differences: coxal 
plate with 1 spine and 8 setae along distal 
margin, 2 setae at proximal margin, and 3 
setae on lateral surface. Basis with 2—4 
spines at posterodistal margin. 

Pereopod 5: Coxal plate with 12 setae on 
anterior lobe and 2 setae on lateral surface. 


i 


Basis bearing 12 spines on anterior margin; 
anterior and posterior lateral surface with 4 
short setae respectively. Ischium with 4 se- 
tae. Anterior margin of carpus with 5—6 sets 
of spines (2-3 singly and 2-3 doubly in- 
serted). Lateral margin of propodus with 13 
spines (S—6 doubly inserted). Dactyl about 
18% length of propodus, bearing | plumose 
seta posteriorly. 

Pereopod 6 subequal to pereopod 5 ex- 
cept for the following characters: coxal 
plate with 1 spine and | seta on anterior 
lobe. Basis without setae on posterolateral 
surface; anterolateral surface with 2—6 se- 
tae; posterior margin bearing 14 spines (pe- 
reopod 5: 11 spines). Merus posteriorly 
with 3 spines and anteriorly with 3—4 spines 
plus 3—4 setae (see S. spinilacertus for pe- 
reopod 5). Anterior margin of carpus with 
8 spines (3 doubly inserted). Dactyl about 
20% length of propodus, bearing 1 plumose 
seta and 1 spine posteriorly. 

Pereopod 7: Coxal plate (Fig. 8d) irreg- 
ular in shape. Basis with 10—11 spines on 
anterior margin. Ischium bearing 1 spine 
and 1—2 setae. Merus with 6 spines (3 dou- 
bly inserted) on anterior margin. Carpus 
with 3 sets of spines (with 3—4 spines per 
set) on anterior margin and 4 sets of spines 
(with 1—3 spines per set) on posterior mar- 
gin. Anterior margin of propodus addition- 
ally with 10 sets of 1—3 slender spines. Dac- 
tyl about 21% length of propodus, with | 
plumose seta at posterior margin. 

Pleopods: Outer rami bearing 12-13 lat- 
eral plumose setae and 7—8 medial plumose 
setae. Inner rami with 7—8 medial plumose 
setae. 

Epimeral plates subquadrate, with small, 
subacute posterior margins, bearing | setule 
each. 

Uropod 1: Peduncle with 4 spines on 
dorsolateral, 4 on dorsomedial, 3 on ventro- 
lateral, and 3 on apical margin, respectively. 
Inner ramus with 7 dorsolateral spines (3-— 
4 dorsomedial and 1—2 dorsolateral spines). 

Uropod 2: Outer ramus bearing 4 dor- 
solateral spines (doubly inserted). 

Uropod 3: Outer ramus bearing 2 spines 


118 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


and 1 seta apically, 5 sets of spines (with 
1-3 spines per set) along lateral margin; 
medial margin with +8 bifurcate setae 
proximally and +6 slender setae distally. 
Medial margin of inner ramus with 8 sets 
of singly and doubly inserted spines. 

Telson (Fig. 8e) width about 76% of 
length; apex with shallow excavation (5% 
of length); each lobe bearing 1 spine plus 
1 seta apically and 3 spines plus 1 seta sub- 
apically. 

Etymology.—The proposed epithet San- 
tanensis is a toponym, referring to the cap- 
ital city Santana, which is situated near the 
type-locality. 

Remarks.—In marked contrast to the 3 
other species of the genus, the preserved 
specimens of S. santanensis appeared whit- 
ish and almost transparent. Specimens of S. 
spinilacertus and S. trajanoae were yellow- 
ish-grey, whereas S. bahiensis showed a 
dark, brownish tone. Since these variations 
appeared to be interspecific (i.e., consistent 
for species from different localities) it is 
possible that they were caused by structural 
differences of the exoskeletons. 


Spelaeogammarus bahiensis da Silva 
Brum, 1975 
Figs: 6dhy Tiss 


Material examined.—Patamute Cave 
(type locality), Curaga, Distrito de Mata- 
muté, Estado do Bahia, Brazil: 1 male par- 
atype, 11.1 mm, and 1| juvenile paratype, 8 
mm (Museu Nacional catalogue no. MN 
5725), collected by P Magalhaes, 1972-— 
1973 @): 

Diagnosis.—Spelaeogammarus bahien- 
sis 1s morphologically more similar to S. 
spinilacertus and S. trajanoae than to S. 
santanensis but differs from the former two 
species in having 7 serrate spines on the 
outer plate of maxilla 1, 20—21 setae on 
coxal plate 5, and the presence of dorsal 
setules on pereonite 7 (1 setule), pleonites 
1—3 (2-10 setules), and uronites 1—2 (1-7 
setules). In addition, it is distinguished from 
S. spinilacertus by short setae instead of 


spines on the anterior margin of the basis 
of gnathopod 1. Male specimen 11.1 mm in 
length. Corresponding to the original de- 
scription by da Silva Brum (1975) with the 
additions and modifications given below. 

Redescription.—Antenna 1: Peduncular 
segment 1—3 gradually decreasing in length; 
peduncular segment 1 with 6 ventral spines. 
Aesthetascs on most segments of primary 
flagellum (as long as segments); accessory 
flagellum with 4 segments, terminal seg- 
ment vestigial. 

Antenna 2: Peduncular segment 3 with 
3—4 spines. Flagellum with 7 segments. 

Mandible: Left lacinia mobilis distinctly 
toothed. Long and short plumose spines be- 
tween lacinia and molar more dissimilar 
than in other species (Fig. 6h). 

Maxilla 1: Palp without lateral setules. 
Outer lobe with 7 apical spines (3 different 
types) and distinct row of setules at medial 
margin, subapically. 

Maxilliped: Dactyl bearing 3—4 relatively 
long setae along inner margin. Inner plate 
bearing apically 2 bifid (y-shaped) spines; 
outer plate with 3 blade-like spines sub- 
apically and 2 strong setae apically (Fig. 
6d). 

Gnathopod 1: Posterior margin of basis 
with 7—9 long setae (all singly inserted); an- 
terior margin bearing 3—5 short setae. Car- 
pus with rectangular posterior lobe, bearing 
3-5 long, sickle-shaped, naked setae, 3—6 
comb-like setae, and 3—4 long, naked setae. 
Palmar margin of propodus bearing 5-9 
normal spines and 13—18 short bifid spines 
on the lateral margin; medial margin with 
8—10 short setae, 1 normal angular spine, 
and 3—5 corner spines (1—2 relatively long). 
Dactyl with 9-10 denticles, each with 1 
short seta. 

Gnathopod 2 basically like that of S. spi- 
nilacertus. Palp with 2 normal spines and 4 
spines at corner. Dactyl with 6—7 denticles. 

Pereopods 3 and 4 subequal. Pereopod 3 
basis with | distoanterior spine, and 1 dis- 
toposterior seta. Pereopod 4 basis with 1—2 
distoanterior spines and | seta plus 1 spine 


VOLUME 113, NUMBER | 


at distoposterior margin; dactyl with | spine 
similar to S. santanensis. 

Pereopods 5—7 missing in specimens ex- 
amined. 

Pleopods basically like those of S. spi- 
nilacertus. Segment 1 bearing 7-8 lateral 
plumose setae and 5—6 medial plumose se- 
tae. Inner ramus with 7 plumose setae on 
medial margin. 

Coxal plates 3—4 with 9-10 setae and 1 
spine; plates 5—6 identical, bearing 20—21 
setae and 1 spine, respectively; plate 7 (Fig. 
8f) irregular, slightly lobed. 

Uropod 1: Peduncle with 4 spines on 
dorsolateral, dorsomedial, and ventrolateral 
margins, respectively, and 2 on apical mar- 
gin. Outer ramus with up to 5 apical spines. 
All spines on inner ramus singly inserted. 

Uropod 2: Peduncle bearing 1—2 dorso- 
medial, 1—2 dorsolateral, and 2 apical 
spines. 

Uropod 3 missing in specimens exam- 
ined. 

Telson (Fig. 8g) width about 80% of 
length; each lobe bearing 2 setae, 2 apical 
and 3—4 subapical spines. 


Discussion 


The four species described above are re- 
corded from eight caves in a series of dis- 
continuous karst areas that extend over a 
linear distance of ca. 1200 km from north 
to south in eastern Brazil (Fig. 9). The rec- 
ords for each species per karst area are as 
follows: S. santanensis—1 cave, S. bahien- 
sis—1 cave, S. spinilacertus—2 caves, and 
S. trajanoae—4 caves. Each species is ob- 
viously restricted to one or more caves in a 
separate karst area. These areas are disjunct 
and apparently physically isolated from 
each other. The greatest distance between 
caves with two species is roughly 650 km 
(S. bahiensis and S. santanensis), whereas 
the shortest distance is only about 135 km 
(S. bahiensis and S. spinilacertus). Caves 
situated within a single, continuous karst 
area that are inhabited by the same species 
are never more than ca. 100 km apart. 


119 


Both the elongate coxal plates and ae- 
quiramus pleopods found in the genus Spe- 
laeogammarus are characters usually con- 
sidered plesiomorphic for the family Bogi- 
diellidae (Stock 1981, Barnard & Barnard 
1983). Apart from Spelaeogammarus, coxal 
plates that are longer than wide are known 
only for the genus Artesia Holsinger, 1980 
(in Holsinger & Longley 1980), and this ge- 
nus, like Spelaeogammarus, also has pleo- 
pods with aequiramus inner rami. However, 
they are 5-segmented in Artesia and only 1- 
segmented in Spelaeogammarus. Artesia 
can also be distinguished from Spelaeo- 
gammarus by 1-segmented accessory fla- 
gellum, fewer flagellar segments in both an- 
tennae, l-segmented palp of maxilla 2, 6- 
segmented pleopodal exopodite, unlobed 
coxal plates 5 and 6, and the telson, which 
is deeply cleft and bears 4—6 apical spines 
on each lobe. Equally long rami are also 
found in Aequigidiella Botosaneanu & 
Stock, 1989, Kerguelenicola Ruffo, 1974, 
and Parabogidiella Holsinger, 1980 (in 
Holsinger & Longley 1980). However, in 
these three genera the coxal plates are typ- 
ically wider than long. Moreover, Aequigi- 
diella differs from Spelaeogammarus by 
sexually dimorphic inner rami and spines of 
the second uropods of the male and a telson 
that is much longer than wide. Although to 
date only one specimen of the genus Ker- 
guelenicola is known, there are several 
characters that distinguish it from Spelaeo- 
gammarus: 1-segmented accessory flagel- 
lum, distinct shape of and lack of armature 
on the telson, large mandibular molar, and 
reduced number of spines and setae on the 
outer and inner lobes of maxilla 1. Para- 
bogidiella differs from Spelaeogammarus 
by l-segmented accessory flagellum, 5-seg- 
mented flagellum of antenna 2, 1-segment- 
ed palp of maxilla 2, characteristically elon- 
gated pereopod 7, 5 pairs of coxal gills, and 
armature of the telson. 

The most closely related bogidiellid tax- 
on to Spelaeogammarus described to date 
may be Bogidiella gammariformis Sket 
(1985) from a cave in Equador. This species 


120 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


VENEZUELA 
bd San —_ 
> 


COLOMBIA Bi 
ia . 


Pacific 
Ocean 


Fig. 9: 


BRAZIL 


’ Salvador 


2 mi 
Rio De Janeiro 


Sao Paulo 


Atlantic Ocean 


Distribution of species of Spelaeogammarus in eastern Brazil: 1) S. bahiensis (1 cave); 2) S. trajanoae 


(4 caves); 3) S. spinilacertus (2 caves); 4) S. santanensis (1 cave). Shading indicates principal cave and/or karst 


areas. Map adapted from Trajano & Sanchez (1994). 


features some interesting characters that 
might be interpreted as intermediate states 
between the relatively primitive Spelaeo- 
gammarus and the more derived Bogidiella 
s. str. For example, B. gammariformis has 
enlarged, bilobed coxal plates 5 and 6, 
which are longer than wide, therefore show- 
ing a strong resemblance to the coxae of 


Spelaeogammarus. The inner rami of the 
pleopods of B. gammariformis are also 1- 
segmented, but show the same reduction as 
in most other species of Bogidiella s. str. 
(i.e., shorter than segment 1 of the outer 
ramus). 

The concentration of the four morpho- 
logically closely similar species of Spelaeo- 


a 


Fig. 10. Distribution of bogidiellid genera in continental South America: 1) Bogidiella cooki Grosso & 
Ringuelet, 1979; 2) B. gammariformis Sket, 1985; 3) B. neotropica Ruffo, 1952; 4) B. (Dycticogidiella) ringueleti 
Grosso & Fernandez, 1988; 5) B. (Dyct.) talampayensis Grosso & Claps, 1985; 6) B. (Mesochthongidiella) 


VOLUME 113, NUMBER 1 121 


sein Pa ss 


pie 


10S 


- “St 


20S 


Pacific Ocean 


30S 


ARGENTINA Atlantic Ocean 


40S 


50S 


80W 7OW 60W SOW 40W 


tucumanensis Grosso & Fernandez, 1985; 7) B. (Stygogidiella) hormocollensis Grosso & Fernandez, 1988; 8) 
B. (Styg.) lavillai Grosso & Claps, 1984; 9) Eobogidiella purmamarcensis Karaman, 1982; 10) Marigidiella 
brasiliensis Stock, 1981; 11) Megagidiella azul Koenemann & Holsinger, 1999; 12) Patagongidiella danieli 
Grosso & Fernandez, 1993 and P. mauryi Grosso & Fernandez, 1993 (sympatric species); 13) Pseudingolfiella 
chilensis Noodt, 1965; 14) Spelaeogammarus bahiensis da Silva Brum, 1975, S. santanensis n. sp., S. spinila- 
certus n. sp., and S. trajanoae n. sp. 


[22 


22 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


gammarus in a series of disjunct caves is 
unique for South America (see Fig. 10). 
Particularly interesting are the relatively 
subtle morphological differences between 
the four species that, in turn, appear to be 
correlated with the interspecific spatial dis- 
tance as well as the abundance of species 
per area: S. spinilacertus and S. trajanoae 
occur in the central part of the range of Spe- 
laeogammarus and show the highest mor- 
phological resemblance. In contrast are the 
more obvious differences between the rel- 
atively large S. santanensis and the smaller 
S. bahiensis, which occur on opposite ends 
of the generic range. 

In South America there is a second con- 
centration of species in northern Argentina, 
which occur exclusively in hyporheic (7?) 
habitats along the Rio Grande (see Fig. 10). 
This cluster is also strictly endemic but it 
has a higher generic diversity, with six spe- 
cies in two genera and three subgenera, 
possibly reflecting the time of divergence 
from a very old freshwater precursor and 
the subsequent radiation into a region of 
isolated inland habitats. However, the dis- 
tribution pattern of Spelaeogammarus 
seems to indicate quite a different historical 
scenario, inasmuch as the species of this ge- 
nus show an exceptionally close morpho- 
logical relationship with each other when 
compared with all other South American 
bogidiellids. If we assume, for the sake of 
argument, the same evolutionary rate for all 
South American bogidiellids, the cluster of 
species belonging to Spelaeogammarus ap- 
pears to have originated from a common 
ancestor far more recently as opposed to 
other bogidiellids in South American fresh- 
water habitats. 

The morphological appearance of the 
four species as well as their distribution 
Over a relatively wide range of disjunct 
karst “‘islands’”’ characterize Spelaeogam- 
marus as a distinct genus within the family 
Bogidiellidae. The apparent isolation of 
these species may well reflect a sequence of 
allopatric (geographic) speciation events 
over a relatively short period of time. 


Acknowledgments 


We are grateful to Professor Eleonora 
Trajano for providing us with some of the 
specimens examined in this study and for 
helpful comments on Brazilian caves and to 
Dr. Paolo S. Young, curator of the Museum 
Nacional, Rio de Janeiro, for making the 
paratypes of Spelaeogammarus bahiensis 
available to us. This study was supported 
by a PEET grant from the National Science 
Foundation to JRH (DEB-9521752). We 
thank the Graphic Office at Old Dominion 
University for assistance with preparation 
of the distribution maps. 


Literature Cited 


Barnard, J. L., & C. M. Barnard. 1983. Freshwater 
Amphipoda of the world.—I. Evolutionary pat- 
terns. Hayfield Associates: Mt. Vernon (Virgin- 
ia). 

Botosaneanu, L., & J. H. Stock. 1989. A remarkable 
genus of cavernicolous Bogidiellidae (Crusta- 
cea, Amphipoda) from Thailand.—Studies in 
honour of Dr. Pieter Wagenaar Hummelinck. 
Foundation for Scientific Research in Surinam 
and the Netherlands Antilles, Amsterdam, no. 
123; 

da Silva Brum, I. N. 1975. Spelaeogammarus bahien- 
sis g.n. sp.n. de Anfipodo Cavernicola do Brasil. 
(Amphipoda-Bogidiellidae).—Atas da Socieda- 
de de Biologia di Rio de Janeiro 17:125—128. 

Grosso, L. E., & G. L. Claps. 1984. Tercer Bogidiel- 
lidae (Crustacea Amphipoda) de la cuenca del 
Rio Grande (Jujuy, Argentina).—Neotropica 
30:223-231. 

ee ss . 1985. Distribucién geogrdafica de 

la familia Bogidiellidae (Crustacea, Amphipo- 

da) en la Republica Argentina, con la descrip- 
cidn de un nuevo subgénero y una nueva es- 

pecie.—Physis (Buenos Aires), Secc. B 43:49— 

523 

, & H. R. Fernandez. 1985. Una nueva Bogi- 

diella (Amphipoda Bogidiellidae) hiporreica de 

la provincia de Tucuman (Argentina).—Neotro- 

pica 31:201—209. 

pide . 1988. Un caso de simpatria de tres 

especies del género Bogidiella (Crustacea, Am- 

phipoda) en el noroeste Argentino, con la de- 
scription de dos nuevoas especies.—Stygologia 

4:64-78. 

Pee . 1993. Nuevo género cavernicola 

austral de Bogidiellidae; Patagongidiella n. 

gen. del noroeste Patagonico (Neuquén, Argen- 


VOLUME 113, NUMBER 1 


tina).—Bolletino del Museo Civico di Storia 

Naturale Verona 17:357-372. 

, & R. A. Ringuelet. 1979. Fauna subterranea 
de las aguas dulces de la Republica Argentina. 
I. Dos nuevas especies de Amphipodos del gé- 
nero Bogidiella.—Limnobios 1:381—394. 

Holsinger, J. R., & G. Longley. 1980. The subterranean 
amphipod crustacean fauna of an artesian well 
in Texas.—Smithonian Contributions to Zoolo- 
gy 308:1-62. 

Karaman, G. S. 1982. Critical remarks to the recent 
revisions of bogidiella-group of genera with 
study of some taxa (fam. Gammaridae). Contri- 
bution to the knowledge of Amphipoda 126.— 
Poljoprivreda I Sumarstvo Titograd 28:31—57. 

Koenemann, S., & Holsinger, J. R. 1999. Megagidiella 
azul n. gen., n. sp., a cavernicolous amphipod 
crustacean of the family Bogidiellidae from 
Brazil, with remarks on its biogeographic and 
phylogenetic relationships.—Proceedings of the 
Biological Society of Washington 112:572—580. 


123 


Noodt, W. 1965. Interstitielle Amphipoden der kon- 
vergenten Gattungen I/ngolfiella Hansen und 
Pseudingolfiella n. gen. aus Suedamerika.— 
Crustaceana 9:17—30. 

Ruffo, S. 1952. Bogidiella neotropica n. sp., nuovo 
Anfipodo dell’ Amazonia.—Rivista Svizzera di 
Idrologia 14/1:129-—134. 

. 1974. Nuovi Anfipodi interstiziali delle coste 
del Sud Africa.—Atti dell’ Istituto Veneto di 
Scienze, Lettere ed Arti 132:399-—419. 

Sket, B. 1985. Bogidiella (s. 1.) gammariformis sp. n. 
(Amphipoda) from Equador.—BioloSk, Vestnik 
33:81-88. 

Stock, J. H. 1981. The taxonomy and zoogeography 
of the family of Bogidiellidae (Crustacea, Am- 
phipoda), with emphasis on the West Indian 
taxa.—Bijdragen tot de Dierkunde 51:345-—374. 

Trajano, E., & L. E. Sanchez, 1994. Bresil. Pp. 527- 
540 in C. Juberthie, & V. Recu eds., Encyclo- 
paedia Biospeologica, Tome I. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


113(1):124—128. 2000. 


Eudendrium bathyalis, a new species of hydroid 
(Hydrozoa: Anthomedusae: Eudendriidae) from Bermuda 


Antonio C. Marques and Dale R. Calder 


(ACM) Departamento de Biologia - Faculdade de Filosofia, Ciéncias e Letras de Ribeirao Preto - 
Universidade de Sao Paulo. Av. Bandeirantes 3900, 14040-9001, Ribeirao Preto, SP, Brazil. 
E-mail: marques @ ffclrp.usp.br; 

(DRC) Centre for Biodiversity and Conservation Biology, Invertebrate Zoology, Royal Ontario 
Museum, 100 Queens Park, Toronto, Ontario, MS5S 2C6, Canada. E-mail: dalec@rom.on.ca 


Abstract.—Eudendrium bathyalis, a new species, is described from relatively 
deep-water (283 m) on the offshore slope of the oceanic island of Bermuda. 
Its distinctive characters are the small size of the colony, the cnidome, com- 
prising two sizes of microbasic euryteles and the unreduced female blastostyle, 
bearing eggs supported by long peduncles. 


The shallow-water hydroids of Bermuda 
were recently revised in a series of studies 
by Calder (1988, 1991, 1997). These: re- 
ports complemented several earlier studies 
on the fauna of the region (viz. Allman 
1888, Congdon 1907, Ritchie 1909, Bennitt 
1922, among others). 

Combining records from these studies 
with collections of hydroids from deeper 
waters (Calder 1996, 1998), a total of 110 
species has been reported from this small 
oceanic island. The relatively large number 
of species from a small geographic area 
likely reflects the extensive sampling un- 
dertaken in the area, from the intertidal 
zone to abyssal bottoms, including bays, 
grassbeds, caves, reefs, ponds, mangroves, 
and pelagic seaweeds. 

Within the Eudendriidae, four species be- 
longing to the two known genera of the 
family (Myrionema amboinense Pictet, 
1893, Eudendrium bermudense Calder, 
1988, Eudendrium capillare Alder, 1856, 
and Eudendrium carneum Clarke, 1882) are 
known from Bermuda. 

These four species appear to be wide- 
spread in the warm western Atlantic. An- 
other species recorded from the Caribbean 
region is Eudendrium ramosum (Linnaeus 
1758) (Wedler 1975, for Colombia; Wedler 


& Larson 1986, for Puerto Rico); this spe- 
cies is also present in the fauna of Bermuda 
(pers. obs.). Other species of Eudendrium 
have also been described or reported for the 
region (Allman 1877, Fraser 1944). These 
species are poorly known taxonomically, 
being characterized on gross morphology 
alone; in some cases, hydranths were lack- 
ing, and no information exists on their com- 
plement of nematocysts. 

The purpose of this report is to describe 
a new species of Eudendrium, collected by 
submersible from bathyal waters on the 
slope of the Bermuda Pedestal. 


Methods 


Material was collected by submersible 
(SDL-1) on 3 Mar 1997, depth 283 m, from 
the Bermuda Pedestal. The study area is 
discussed more fully elsewhere (Calder 
1998). The holotype was examined, mea- 
sured, and drawn under microscope and ste- 
reomicroscope, both with camara lucida. 
The cnidome terminology follows Weill 
(1934) and Mariscal (1974), and measures 
of nematocysts were made on non-dis- 
charged capsules. The L/W ratio (Kubota 
1976) and S/C ratio (Watson 1987) are also 
provided. Other study methods for Euden- 


VOLUME 113, NUMBER 1 


driidae are from Marques (1995) and 
Marques & Migotto (1998). 


Systematics 


Genus Eudendrium Ehrenberg, 1834 
Eudendrium bathyalis, new species 
Fig. 1 

Examined material.—Holotype, one fe- 
male colony, Bermuda Pedestal 32°16.6'N 
ma 44.5 W, 283 m,; 3 Mar 1997, on a 
sponge, ROMIZ B3034. 

Description.—Colonies dioecious, frag- 
ile, up to 18 mm in height; main stems 
slightly fascicled basally or up to the half 
of the colony, sometimes formed only by a 
couple of tubes, fascicled region up to 0.18 
mm in diameter. Hydrocauli arising from 
stolonal hydrorhiza growing over a sponge; 
branches few, irregular, occurring over en- 
tire hydrocaulus, branches up to third order, 
in radiate planes; pedicels arising from 
main stem or branches of first and second 
order. Perisarc of main stem weakly devel- 
oped, single tubes 0.08—0.10 mm in diam- 
eter, unfascicled region with scarce annu- 
lations, in sets of 2—3 rings. Branches with 
2-5 rings at origin, 0.06—0.08 mm in di- 
ameter. Pedicels obscurely annulated at or- 
igin, with 2—4 rings, very delicate, 0.05— 
0.06 mm in diameter. 

Hydranths 0.10—0.25 mm in height, 
0.10—0.29 mm in diameter (measured in the 
body region just below the tentacles), with 
a distinct deep groove in the aboral region; 
hypostome large; tentacles 18—24 in num- 
ber, occurring in a whorl below hypostome. 

Gonophores styloids, arising from body 
of hydranth. Immature styloids placed in a 
circle around body of hydranth. Female 
blastostyles styloids without a characteristic 
Spadix over a single egg. Tentacles and hy- 
postome not reduced during ontogeny of fe- 
male gonophores. Eggs almost circular, ma- 
turity undeterminable, encapsulated by a 
thin gelatinous layer, linked by long pedun- 
cles to body of hydranth, distal part of pe- 
duncle broadened for egg support. Eggs 3-— 


125 


5 in number, 0.14—0.15 mm in diameter. 
Male gonophores not observed. 

Nematocysts of one category, heterotri- 
chous microbasic euryteles in two size clas- 
ses. 

Small microbasic euryteles (not seen dis- 
charged), 6.0-6.3 by 2.8-3.2 wm, L/W = 
1:2.0—2.1, oval, abundant; distributed over 
hydranth body, hypostome, peduncle coe- 
nosarc of female gonophore, and tentacles. 

Large microbasic euryteles (seen dis- 
charged), 22.1—23.4 by 8.2—-8.9 um, L/W = 
1:2.6—2.7, bean-shaped; shaft crossing 
about from 0.5 to 0.75 of the whole length 
of undischarged capsule, discharged shaft 
heavily armed, ca. 19.8 ym in length, pro- 
portion S/C = 1.4; nematocysts distributed 
over hydranth body (sometimes common), 
egg peduncle (rare), and coenosarc (not 
seen on hypostome). 

Etymology.—bathyalis, adj. from Greek 
bathys = deep, in allusion to the bathyal 
depth from which material of this species 
was collected. 

Remarks.—The distinctive character of 
this species is the pedunculated gonophore, 
which we presumed would be a female one. 
A similar female gonophore has only been 
observed in Eudendrium vervoorti Marques 
& Migotto, 1998, a recently described spe- 
cies based on scarce material, without hy- 
drorhiza and covered by debris, from the 
coast of the Netherlands. In E. vervoorti, 
the gonophores apparently arose from the 
hydranth body, without a conspicuous spa- 
dix; some other gonophores (presumably 
older ones) are linked to the body of hy- 
dranth or to the pedicel by long stalks 
(Marques & Migotto 1998). Another simi- 
lar feature between both species is the ex- 
tremely delicate branches (widths barely 
greater than 0.1 mm), but this character is 
also shared by several other species of the 
genus [e.g., Eudendrium album Nutting, 
1896, Eudendrium fragile Motz-Kossows- 
ka, 1905, Eudendrium generale von Len- 
denfeld, 1885, Eudendrium tottoni Ste- 
chow, 1932 (=Eudendrium antarcticum 
Totton, 1930, see Stechow 1932) and Eu- 


126 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


A 


Fig. 1. 


y 


C D E 


Eudendrium bathyalis, new species; all from holotype ROMIZ B3034. A, hydranth with the distri- 


bution of the large microbasic euryteles represented over the body; B, female blastostyle; C, capsule of the large 
microbasic eurytele; D, discharged large microbasic eurytele; E, capsule of the small microbasic euryteles. Scale 


bar, A-B = 0.1 mm; C—E = 10 pm. 


dendrium motzkossowskae Picard, 1951], 
but none of those has a female gonophore 
as in E. vervoorti and E. bathyalis. 

The cnidome is the distinctive character 
between E. vervoorti and E. bathyalis. The 
Netherlands species has only small micro- 
basic euryteles and the species from Ber- 
muda has both small and large nematocysts. 
The material from Bermuda is slightly fas- 
cicled, but as the Netherlands material is 
represented only by a small fragment, even 
without hydrorhiza (Marques & Migotto 


1998), this difference could be only an ar- 
tifact. 


Key to the species of Eudendriidae 
recorded from Bermuda 


la. More than 40 tentacles arranged in two 
whorls, presence of abundant zooxan- 
thellac-4c nn. Ss Myrionema amboinense 
lb. Usually fewer than 35 tentacles ar- 
ranged in a unique whorl, zooxanthel- 
las absent th) 3-4": Se eee 2 


VOLUME 113, NUMBER 1 


2a. Nematocysts of only one size class, mi- 
crobasic eurytele type 
“65 OT SS Cee ee Eudendrium capillare 
2b. Nematocysts in two size classes .... 3 
3a. Cnidome including small and large mi- 
Sea CUR YUCICS 5 ox. ys. spite w wy etailers + 
3b. Cnidome including small microbasic 
euryteles and large nematocysts differ- 
ent from microbasic euryteles ...... 5 
4a. Female gonophore encircled by un- 
mumimed Spagix'. 27 1. 8S 70S 


4b. Female gonophore supported by long 
peduncles, no characteristic spadix . 
ae Eudendrium bathyalis, new species 
5a. Large nematocysts heterotrichous ani- 
sorhiza, female immature spadix bifid 
Rete. kak h ans Eudendrium carneum 
5b. Large nematocysts macrobasic euryte- 


les, female immature spadix  un- 

ie i eee 

ofl Eudendrium bermudense 
Acknowledgments 


The authors wish to thank to Prof. Wim 
Vervoort (Nationaal Natuurhistorisch Mu- 
seum, Leiden, The Netherlands), in which 
laboratory part of the study was done, to 
Dr. Alvaro E. Migotto (University of Sao 
Paulo, Sao Sebastiao, Brazil) and Dr. Jean- 
ette E. Watson (Museum of Victoria, Aus- 
tralia) for suggestions concerning the man- 
uscript, and to Charles M.D. Santos for his 
help with some drawings. ACM has finan- 
cial support from the Fundagao de Amparo 
a Pesquisa do Estado de Sao Paulo (FA- 
PESP 96/10544-0, 97/04572-4, and 98/ 
14822-0). DRC acknowledges the Canadian 
Navy for the opportunity to dive in the 
SDL-1 submersible, used in collection of 
this hydroid. Thanks are also due to the 
Natural Sciences and Engineering Research 
Council of Canada for financial support. 


Literature Cited 


Alder, J. 1856. A notice of some new genera and spe- 
cies of British hydroid zoophytes.—Annals and 
Magazine of Natural History, 2nd series 18: 
353-362. 


127 


Allman, G. J. 1877. Report on the Hydroida collected 
during the exploration of the Gulf Stream by 
L.F de Pourtalés, assistant United States Coast 
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. 1888. Report on the Hydroida dredged by 
H.M.S. Challenger during the years 1873-76. 
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Report on the Scientific Results of the Voyage 
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76, Zoology 23:1—90. 

Bennitt, R. 1922. Additions to the hydroid fauna of the 
Bermudas. Contributions from the Bermuda bi- 
ological Station for Research. No. 136.—Pro- 
ceedings of the American Academy of Arts and 
Sciences 57(10):241—259. 

Calder, D. R. 1988. Shallow-water hydroids of Ber- 
muda: the Athecatae.—Royal Ontario Museum, 
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. 1991. Shallow-water hydroids of Bermuda: 

the Thecatae, exclusive the Plumularioidea.— 

Royal Ontario Museum, Life Sciences Contri- 

butions 154:1—140. 

. 1996. Hydroids (Cnidaria: Hydrozoa) record- 

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ern North Atlantic.—Canadian Journal of Zo- 

ology 74:1721-—1726. 

. 1997. Shallow-water hydroids of Bermuda: 

superfamily Plumularioidea.—Royal Ontario 

Museum, Life Sciences Contributions 161:1— 

85. 

. 1998. Hydroid diversity and species compo- 
sition along a gradient from shallow waters to 
deep sea around Bermuda.—Deep-sea Research 
I, 45:1843-—1860. 

Clarke, S. F 1882. New and interesting hydroids from 
Chesapeake Bay.—Memoirs of the Boston So- 
ciety of Natural History 3(4):135—142. 

Congdon, E. D. 1907. The hydroids of Bermuda.— 
Proceedings of the American Academy of Arts 
and Sciences 42:461—485. 

Ehrenberg, C. G. 1834. Beitrage zur physiologischen 
Kenntniss der Corallenthiere im allgemeinen, 
und besonders des Rothen Meeres, nebst einem 
Versuche zur physiologischen Systematik der- 
selben.—Abhandlung der KOniglichen Akade- 
mie der Wissenschaften 1:225—380. 

Fraser, C. M. 1944. Hydroids of the Atlantic coast of 
North America. The University of Toronto 
Press, Toronto, 1-451, pls. 1-94. 

Kubota, S. 1976. Notes of nematocysts of Japanese 
hydroids, I.—Journal of the Faculty of Science 
of the Hokkaido University 20(2):230-—243. 

Mariscal, R. N. 1974. Chapter 3. Nematocysts. Pp. 
129-178, in L. Muscatine & H. M. Lenhoff, 
eds., Coelenterate Biology. Academic Press, 
New York. 


128 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Marques, A. C. 1995. Eudendrium pocaruquarum Nn. 
sp. (Hydrozoa, Eudendriidae) from the south- 
eastern coast of Brazil, with remarks on taxo- 
nomic approaches to the family Eudendri- 
idae.—Contributions to Zoology 65(1):35—40. 

, & A. E. Migotto. 1998. A new species of 
Eudendrium (Hydrozoa, Anthomedusae, Euden- 
driidae) from the Netherlands.—Zoologische 
Verhandelingen 323:149—-154. 

Motz-Kossowska, S. 1905. Contribution a la coinnaiss- 
ance des hydraires de la Méditerranée Occiden- 
tale. I. Hydraire Gimnoblastiques.—Archives 
de Zoologie Expérimentale et Générale, 4me 
série 3:39—98. 

Nutting, C. C. 1896. Notes on Plymouth Hydroids.— 
Journal of the Marine Biological Association of 
United Kingdom, n. ser. 4(2):146—154. 

Picard, J. 1951. Note sur les hydraires littoraux de 
Banyuls-sur-Mer. Vie et Milieu.—Banyuls-sur- 
Mer 2:338-349. 

Pictet, C. 1893. Etude sur les hydraires de la Baie 
d’Amboine.—Revue Suisse de Zoologie 1:1-— 
64. 

Ritchie, J. 1909. Two unrecorded “‘Challenger’’ hy- 
droids from the Bermudas, with a note on the 
synonymy of Campanularia insignis.—Zoolo- 
gist, 4th Ser. 13:260-—263. 

Stechow, E. 1932. Neue Hydroiden aus dem Mittel- 


meer und dem Pazifischen Ozean, nebst Be- 
merkungen liber einige wenig bekannte For- 
men.—Zoologischer Anzeiger 100(3—4):81—92. 

Totton, A. K. 1930. Coelenterata. Part V.-Hydroida.— 
British Antarctic (‘Terra Nova’) Expedition 
1910, Natural History Report, Zoology 5(5): 
131-252. 

von Lendenfeld, R. 1885. The Australian Hydrome- 
dusae. I].—Proceedings of the Linnean Society 
of New South Wales 9:345-—363. 

Watson, J. E. 1987. Records of Eudendrium (Hydro- 
zoa: Hydroida) from New Zealand.—Proceed- 
ings of the Linnean Society of New South 
Wales 109(4):325—330. 

Wedler, E. 1975. Okologische Untersuchungen an Hy- 
droiden des Felslitorals von Santa Marta (Kolum- 
bien).—Helgolander Wissenschaftliche Meere- 
suntersuchungen 27:324—363. 

Wedler, E., & R. Larson. 1986. Athecate hydroids from 
Puerto Rico and the Virgin Islands.—Studies of 
Neotropical Fauna and Environment 21(1-2): 
69-101. 

Weill, R. 1934. Contribution a 1l’étude des cnidaires et 
de leurs nématocystes. I. Recherches sur les né- 
matocystes (morphologie, physiologie, dével- 
opment). II. Le valeur taxonomique du cni- 
dome.—Travaux de la Station Zoologique de 
Wimereux 11:1—351. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


113(1):129-144. 2000. 


Cnidae of two species of Discosomatidae 
(Cnidaria: Anthozoa: Corallimorpharia) from Brazil 


Suzana Machado Pinto and Maria Julia da Costa Belém 


(SMP) Depto. de Invertebrados, Museu Nacional, Universidade Federal do Rio de Janeiro/ 
UFRJ, Quinta da Boa Vista, 20940-040, Rio de Janeiro, RJ.; Current address: 
Depto. de Zoologia, Instituto de Biociéncias, Universidade de Sao Paulo (USP), Caixa Postal 
11461, 05422-970. Sao Paulo, SP, Brasil. e-mail: suzanamp @ib.usp.br; 
(MJCB) Caixa Postal 24.030, 20.522-970. Rio de Janeiro, RJ, Brasil. 
e-mail: belcosta@ mandic.com.br 


Abstract.—The cnidae of Discosoma carlgreni (Watzl, 1922) and Discosoma 
sanctithomae (Duchassaing & Michelotti, 1860) are described and illustrated. 
Five types (sensu Schmidt 1969, 1974) were observed: spirocysts, b-rhabdoids, 
p-rhabdoids D, holotrichs I and holotrichs II. An unusual variety of holotrich 
occurred in the tentacles and column that had not been reported previously. 
These species are distinguished based on the distribution and size of the types 
of cnidae. The importance of qualitative studies of corallimorpharians cnidae 


is discussed. 


This paper investigates the diversity of 
cnidae that occur in the corallimorpharians 
Discosoma carlgreni (Watzl, 1922) and D. 
sanctithomae (Duchassaing & Michelotti, 
1860) from Brazil, as well as their taxo- 
nomic value. 

The diagnostic value of cnidae in differ- 
ent structures of cnidarians has been ex- 
amined by several authors (e.g., Weill 1934, 
Russell 1938, Carlgren 1949, Cutress 1955, 
Schmidt 1972, 1974, Mariscal 1974, den 
Hartog 1980, Fautin 1986, England 1991, 
Williams 1996, Pires 1997, Ostman & Hyd- 
man 1997). 

Corallimorpharia is a relatively small or- 
der of skeletonless Anthozoa, morphologi- 
cally intermediate between sea anemones 
and corals. Many authors consider them 
most closely related to corals (e.g., Duerden 
1904, Stephenson 1921, Schmidt 1974, den 
Hartog 1980, Fautin & Lowenstein 1992, 
Chen et al. 1996, Pinto & Belém 1997, Pi- 
res & Castro 1997). Contributions to the 
systematics of the Discosomatidae, based 
on morphological and anatomical charac- 
ters, have been made by several authors 


(e.g., Duchassaing & Michelotti 1864, 
Duerden 1900, Stephenson 1921, Carlgren 
1940, 1949, Corréa 1964, Schmidt 1972, 
1974, den Hartog 1980, Schlenz & Belém 
1982). The revision of the Caribbean shal- 
low-water Corallimorpharia, as set forth by 
den Hartog (1980), provided a comprehen- 
sive review of the morphology, anatomy, 
histology and classification of this group. 
den Hartog (1980) rearranged the family 
Discosomatidae Duchassaing & Michelotti, 
1864 on the basis of an array of material 
from the Caribbean. He united the five gen- 
era of Actinodiscidae (Carlgren, 1949) in 
the genus Discosoma Riippell & Leuckart, 
1828. den Hartog (1980) concluded that the 
order Corallimorpharia does not fundamen- 
tally differ from corals and should be in- 
cluded as a separate sub-order in the Scler- 
actinia. 

The cnidom of the Discosomatidae was 
previously studied by Watzl (1922), Carl- 
gren (1927, 1949), Corréa (1964), den Har- 
tog (1980) and Schlenz & Belém (1982), 
but these studies were not sufficiently de- 
tailed to provide a good understanding of 
the cnidae in this group. 


130 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Materials and Methods 


Specimens of Discosoma carlgreni (Fig. 
1A) and D. sanctithomae (Fig. 1B) were 
collected from 1990 to 1993 and deposited 
in the Cnidaria collection of the Museu Na- 
cional/Universidade Federal do Rio de Ja- 
neiro, Rio de Janeiro (MNRJ). For this 
study, 57 specimens of D. carlgreni and 42 
specimens of D. sanctithomae were exam- 
ined. Both species were also acquired by 
exchange with Mexico and Cuba in order 
to carry out a comparative study. Speci- 
mens of D. carlgreni were found at the 
South Coast of the state of Espirito Santo 
and Bahia. So far, D. sanctithomae has been 
found only in Abrolhos Archipelago, be- 
tween the channel of Redonda and Siriba 
Islands (see Appendix for details). 

Material was collected by snorkel and 
SCUBA diving at depths between | m and 
16 m, using a hammer and chisel. Living 
animals were placed in plastic sacks with 
water from their site of collection and trans- 
ported on ice. Specimens were anesthetized 
by slowly adding 1:1 solution of 8% MgCl, 
and sea water. After anesthetization, they 
were fixed in 4% formalin. Cnidae were 
studied in squash preparations of living and 
preserved specimens. A fragment of tissue 
was squashed and spread on a slide. Five 
structures were examined: discal tentacle, 
marginal tentacle, column, stomodaeum 
and mesenterial filament. Measurements 
were made only on undischarged capsules. 
At least 40 length/width measurements 
were made of each nematocyst type in each 
structure examined. Measurements were 
made with light microscopy at magnifica- 
tions up to 1250 with differential inter- 
ference contrast optics (Nomarski), using 
an eyepiece micrometer. Drawings were 
made with a camera lucida. Cnidae were 
classified according to the nomenclatures of 
Weill (1934, later modified by Carlgren 
1940), Schmidt (1969, 1972, 1974) and den 
Hartog (1980). 


Results 


Five types of cnidae were observed: spi- 
rocysts, b-rhabdoids, p-rhabdoids D, holo- 
trich I and holotrich II. The cnidae were 
classified as follows. 

Spirocysts (Figs. 3A, 6E).—Elongate and 
thin-walled capsule with a long tube coiled 
in numerous spirals, tubule without spines. 

Remarks.—This variety was sparse but 
typical of the marginal tentacles of both 
species. The capsules reached up to 18 wm 
in length and 3.1 wm in width. Eight un- 
discharged capsules in each species were 
observed. They were not included in Table 
1 due to their sparseness. 

B-rhabdoids.—Varied in shape and size, 
capsules generally oval in aspect. The 
thread of the b-rhabdoids is always armed 
with spines, not having a clear difference 
between the width of the proximal part of 
the tubule and that of the distal portion. We 
observed two morphological varieties of b- 
rhabdoids in undischarged capsules: 

B-rhabdoids' (Figs. 2B, 3C, 4B, 6K 7B, 
Table 1).—Capsule oval to cylindrical in 
shape, of refractive contrast. In the undis- 
charged state, the basal portion of the tu- 
bule is shorter and thinner than that of b- 
rhadoids’, up to 0.33 of the length of the 
capsule. Sometimes the basal part appears 
slightly curved, the tubule being arranged 
in few irregular coils. 

Remarks.—The undischarged capsules 
from the column may be occasionally con- 
fused with holotrich II, due to little refrac- 
tive contrast. 

B-rhabdoids* (Figs. 2A, 3B, 4A, Table 
1).—Capsule transparent, elongate, very 
small and of refractive contrast. Its basal 
portion is short, refractive and clear. 

P-rhabdoids D.—Characterized by a no- 
tably wide tubule ending with a funnel- 
shaped “‘V”’ with two distinct parts: large 
basal shaft and tapered distal tubule. The 
thread is long, bearing spines. We observed 
two morphological varieties of p-rhabdoids 
D in undischarged capsules: 

P-rhabdoids D' (Figs. 2C, D, 3D, 4C, D, 


VOLUME 113, NUMBER 1 131] 


Fig. 1. A, Discosoma carlgreni. Specimens collected among Zoanthus sp. at Santa Cruz, Aracruz, ES. Scale: 
5 cm. B, D. sanctithomae. Aggregation from study site at Abrolhos Archipelago, between the channel of Redonda 
and Siriba Islands. Scale: 2 cm. 


132 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


G, 5B, C, 6A, B, G, H; 7A, H; 8A, B; Table 
1).—Capsule cylindrical, slightly curved, of 
refractive contrast. The shaft has obvious 
turns of spines that are up to 0.5 of the cap- 
sule length. 

Remarks.—This variety was rather com- 
mon and also occurred in all structures ex- 
amined in both species. It also occurred in 
two size classes in the discal tentacles, col- 
umn and mesenterial filaments. In the fila- 
ments of D. sanctithomae some transparent 
thin-walled capsules, with rich contrast in 
the shaft, were observed (Fig. 8B). 

P-rhabdoids D? (Fig. 7D, Table 1).—Rel- 
atively large elongate capsule, cylindrical, 
slightly curved and of refractive contrast. 
Shaft with clear turns of spines can reach 
up to 0.5 length of the undischarged cap- 
sule. A long tubule is irregularly arranged 
inside the capsule. 

Holotrichs—We observed two morpho- 
logical varieties of holotrich in both spe- 
cies: 

Holotrichs I (Figs. 2E, G, 3E G, 4EF H, 
I, 5A,.Ds 6G. D, 1 Ko IE, BGI se. 
Table. 1).—Capsules of various sizes, filled 
with a tubule having a long figure eight 
form, of rich contrast in several turns (Figs. 
2G, 6K). In the discharged state, the tubule 
contains conspicuous spines of equal size 
distributed along almost its entire length, 
except for a short, naked basal portion. The 
distal end of the tubule presents an abrupt 
tapered distal tip, completely devoid of 
spines and measuring up to 30 pm (Figs. 
2G, 6D). 

Remarks.—The distal end of the tubule 
of these nematocysts was previously de- 
scribed by den Hartog (1980) as a terminal 
tubule. The terminal tubule can be seen 
only when the tubule is totally discharged. 
Holotrichs I, which are common, occurred 
in two size classes throughout all structures 
examined, except for the presence of only 
small capsules in the column of D. carl- 
greni (Fig. 4E Table 1). This type ranges in 
size from the small (e.g., in marginal ten- 
tacles—25-—34 by 9.4—18 ym) to large (e.g., 
in filaments—76.9-168.1 by 25-75 wm). 


According to den Hartog (1980), these 
nematocysts are the most voluminous an- 
thozoan cnidae, reaching up to 250 by 80 
jxm. We also observed a particular shape of 
the holotrichs I in the stomodaeum. They 
varied from oblong to cylindrical (Figs. 41; 
7G). 

Holotrichs II (Figs. 3E, 6J, Table 1).— 
Sharply distinguished from holotrich I in 
capsule shape and in having small spines 
with little contrast. The capsule is opaque, 
cylindrical, with coiled tubule in small 
tums, filling the entire undischarged cap- 
sule. The spines are smaller than those of 
holotrichs I and of little contrast and diffi- 
cult to view with light microscopy. 

Remarks.—These were exclusive to the 
marginal tentacles in both species. An un- 
usual variety of this category is smaller and 
cylindrical, with a tubule filling the whole 
capsule and with spines smaller than those 
of the typical holotrich II. These capsules 
seem to differ from the holotrich II, except 
for their shape, with spines so reduced that 
they appear like spots distributed inside the 
capsule with irregularly coiled small turns. 
Presently, we consider it a holotrich II, oc- 
curring in the discal tentacles and in the 
column of D. carlgreni (Figs. 2K 4E); in 
the column of D. sanctithomae, they were 
large (Fig. 7C). This variety has never been 
reported from the discal tentacles and col- 
umn in the Discosomatidae. 


Discussion 


We identified seven types of cnidae in the 
Discosomatidae. The presence of spirocysts 
in this group was discussed by Carlgren 
(1949) and den Hartog (1980). Carlgren 
(1949) recorded the sparseness of spirocysts 
in the Discosomatidae, claiming that they 
were found in the tentacles. den Hartog 
(1980:36) characterized the family by the 
absence of spirocysts and stated “‘not only 
are spirocysts absent in the tentacles of Dis- 
cosomatidae, other ectodermal cnidae too 
are) very “scarse” .. ithe “tentacles: . <2 are 
non-retractile, non-motile ... are either re- 


VOLUME 113, NUMBER 1 


WF 
D. carlgreni ‘ ‘ 
a) 


A 
o 
Cc 
B 
DISCAL 
TENTACLE 


‘“N 


x 


Oks 
7 


eens 


Fig. 2. Cnidome of Discosoma carlgreni, Discal tentacles. A, b-rhabdoids?; B, b-rhabdoids'; C, D, p-rhab- 
doids D'; E, G, holotrichs I; E holotrichs If. Abbreviation: t = terminal tubule. Scale: 10 wm. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


D. carlgrent 


MARGINAL 
TENTACLE 


=> 


SLL. 
CLE — 


\ 


SSA a5 
y— 


\ 
NAN 
NAN 
SS ANS AARNE 


SSSA SD 
Ce Sa ae a 
LAT Or ire 
N 


~ 
~— = 


Nes 


, b-rhabdoids'!: 


D, p-rhabdoids D'; E, holotrichs II; EK G, holotrichs I. Abbreviation: t = terminal tubule. Scale: 10 wm. 


2. 
’ 


Cnidome of Discosoma carlgreni, Marginal tentacles. A, spirocysts; B, b-rhabdoids 


Bipg3) 


a 
om 


VOLUME 113, NUMBER 1 


D. carlgreni 


COLUMN 


STOMODAEUM 


Cnidome of Discosoma carlgreni, Column A—F; Stomodaeum G— 


: B, b-rhabdoids': 


5 


I. A, b-rhabdoids 


Fig. 4. 
C, D, G, p-rhabdoids D'; 


10 ym. 


= terminal tubule. Scale: 


E, holotrichs II; E H, I, holotrichs I. Abbreviation: t 


? 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


2 


FILAMENTS 


D. carlgreni 


| SN 
~ Se oe 


eee 
te 
ANS 


<< 


SE TNS 


! 


! 
NA 
=a At 


= 


Ww, 
FW swans ay LIES pep OEE 
Tea77 70! ‘\ wns Ss 


rhabdoids D!. Abbreviation: 


p 


> 


Cnidome of Discosoma carlgreni, Filaments. A, D, holotrichs I; B, C 


t = terminal tubule. Scale: 10 wm. 


Figs: 


137 


VOLUME 113, NUMBER 1 


D. sanctithomae 


MARGINAL 
TENTACLE 


DISCAL 


Ww 
—_/ 
< 
Ee 
z= 
Lu 
be 


i eae ae ie ae I I 


G, H, 


B, 
terminal 


> 


Fig. 6. Cnidome of Discosoma sanctithomae, Discal tentacles A-—D, Marginal tentacles E-K. A 


p-rhabdoids D'; 


C, D, I, K, holotrichs I; E, spirocysts; EK b-rhabdoids'; J, holotrichs Il. Abbreviation: t 


tubule. Scale: 10 xm. 


° 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


3 


D. sanctithomae 


COLUMN 


Sp PPE SS SSIS 


> ss 
we We %, “y Dy, “7 
TTT, 
SS Spam re 
\ 
ol! 
OTN 


STOMODAEUM 


Cnidome of Discosoma sanctithomae, Column A—F, Stomodaeum G-I. A, H, p-rhabdoids D!; B, b- 


Fig. 7: 
rhabdoids!; 
10 pm. 


terminal tubule. Scale: 


—G, I, holotrichs I. Abbreviation: t 


2. 
> 


holotrichs II; D, p-rhabdoids D 


° 


C 


139 


VOLUME 113, NUMBER 1 


D. sanctithomae 


~ 


FILAMENTS 


SU iy FS 9 


“Bh in Wee re 


Z SLE 


Iam 


= 


SS 


Dopp 7Y 


SSS > 
GOD COCA a 


=> 


~ 
—_— 


pee 


POSAANA SASS 


ML 


-rhabdoids D!; C, D, holotrichs I. Abbre- 


Cnidome of Discosoma sanctithomae, Filaments. A, B, p 


Fig. 8. 
viation: t 


terminal tubule. Scale: 10 ym. 


140 


Table 1.—Distribution measurements of cnidae. Sp = species, A = Discosoma carlgreni, B = D. sanctith- 


omae. n = number of capsules measured, F = figure. 


Structure 


Average and range of length 
and width of nematocyst capsules 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Cnidae type Sp Length Width n jm 
Discal tentacles 
b-rhabdoids' A 14.4 (12.5-17.5) 4.7 (1.9-6.3) 41 2B 
b-rhabdoids? A O57 .5=1 1-3) 1.8 (1.3-3.0) 41 2A 
p-rhabdoids D'! A 12.9 (8.8-17.5) 3.5 (1.9-5.6) 66 ae 
25.4 (18.8-30.6) 6.6 (5.6-8.8) 54 2D 
B 13.3 (10.6—17) 3.5 (2.5-5.0) 40 6A 
33.2 (20-46.9) 7.7 (4.4-11.3) 50 6B 
holotrichs I A 37.2 (28-41.3) 13.3 (6.9-18.1) 76 2E 
72.1 (45—123.1) 31.2 (18.1-55.6) 45 2G 
B 34 (26.9-43.8) 14.8 (11.3-—20) 83 6C 
69 (57.5-85) 32.4 (21.9-45.8) 92 6D 
holotrichs I A 16.5 (11.3-—21.9) 4.9 (3.1-6.9) 42 2F 
Marginal tentacles 
b-rhabdoids' A 15.5 (12.2—23.8) 4.9 (3.8-6.9) 65 6) & 
B 15.6 (11.3—20) 4.0 (2.5—5.6) 72 6F 
b-rhabdoids? A 8.9 (6.3-10.9) 1.9 (0.9-3.1) 40 3B 
p-rhabdoids D'! A 13.2 (8.8-16.3) 4.2 (1.9-6.9) 79 3D 
B 13.5 (10-20) 3.8 (2.5—5.6) 50 6G 
33.7 (23.1-49) 8.0 (5.6—11.9) 45 6H 
holotrichs I A 35.6 (27.5—40.6) 12.7 (9.6—20.6) 65 3F 
82.2 (50.6—125) 34.2 (20.6—-49.4) 60 3G 
B 31.6 (25—34.4) 13.6 (9.4—-18.1) a2 6! 
68.0 (54.4—-83.1) 30.0 (17.5—-40.3) 77 6K 
holotrichs I A 29.7 (8.8-43.8) 5.4 (2.5—10.6) 159 3E 
B 24.8 (13.1-—37) 5.1 (3.1-13.8) 110 6J 
Column 
b-rhabdoids'! A 15.7 (13.1—20) 5.5 (3.8-6.3) 63 4B 
B 19.6 (16.3-25) 5.5 (3.8-7.5) 50 7B 
b-rhabdoids? A 9.6 (5.6—12.5) 2.0 (1.3-3.8) 40 4A 
p-rhabdoids D! A 13.9 (10-17.5) 4.6 (2.5-5.3) 60 4C 
24.1 (19.1—28.8) 6.9 (6.2—8.1) 40 4D 
B 14.5 (10—20.6) 4.0 (2.5-6.3) 45 TA 
p-rhabdoids D? B 41.1 (25-58.8) 8.2 (4.4-13.1) 64 7D 
holotrichs I A 34.6 (30.6—37.5) 1257 (73=16.3) 50 4F 
B 34.1 (28.1—38.1) 15.4 (11.3-18.8) 58 TE 
76.1 (61.9-94.4) 34 (25-42.5) 53 TF 
holotrichs II A 20 (13.8-19) 6.3 (3.8—9.4) 65 4E 
B 33.7 (26.3-40.6) 4.0 (2.5-6.3) 40 Ze 
Stomodaeum 
p-rhabdoids D' A 13.5 (8.8-19) 3.9 (1.9-6.3) 50 4G 
B 17.9 (10.6—32.5) 4.7 (2.5—9.4) 52 7H 
holotrichs I A 37.3 (32.5—40.6) 14.1 (10.6—17.5) 35 4H 
54.6 (45.6—70) 18.1 (10.9—22.5) 65 4I 
B 33.7 (25.640) 13.3 (5.6—18.8) a7 7 
58.8 (44.4-78.8) 18.9 (11.9-35.6) 65 7G 
Filaments 
p-rhabdoids D'! A 18.2 (11.9-23.7) 4.9 (4.5-8.8) 63 5B 
27 (25-33.1) 6.9 (5.0—10) 79 SC 
B 14.6 (8.8—20) 3.6 (1.9-6.3) 40 8A 
31.5 (24.4-38.1) 8.2 (5.0—-15) 78 8B 


VOLUME 113, NUMBER 1 14] 
Table 1.—Continued. 
Average and range of length 
and width of nematocyst capsules 
Structure 
Cnidae type Sp Length Width n F 
holotrichs I A 36.9 (26.2—53.8) 13.6 (7.5—-18.8) 50 SA 
117.6 (90.6—175.2) 49 (31.2-72) 69 5D 
B 31.8 (23.8—37.5) 13.1 (5.0-18.1) 69 8C 
146.5 (76.9—168.1) 60.3 (25-75) 73 8D 


duced to insignificant, wartlike protuber- 
ances or developed into vesicle-like struc- 
tures...” den Hartog (1980) pointed out 
the difficulty of accepting that these tenta- 
cles were functional catching devices be- 
cause the Discosomatidade have been as- 
sociated with zooxanthellae that provide 
nutrition to the animals. However, several 
Actiniaria, such as Stichodactyla haddoni 
(Saville-Kent, 1893) and Stichodactyla 
duerdeni (Carlgren, 1900), with a large 
number of spirocysts, have similar tentacles 
and zooxanthellae. In this study, we found 
spirocysts in the marginal tentacles of both 
species, thus confirming Carlgren’s finding. 

Although den Hartog (1980) accepted 
Schmidt’s system (1972, 1974) of termi- 
nology for corallimorpharian cnidae, he 
preferred to adopt Stephenson’s system of 
classification. den Hartog (1980) considered 
Schmidt’s b- and p-rhabdoids (=b and p- 
mastigophores sensu Carlgren, 1940) as 
synonyms of spirulae and penicilli sensu 
Stephenson (1928), respectively. Therefore, 
he recorded four types of cnidae for the 
Discosomatidae: spirulae, penicilli D, pen- 
icilli E and homotrich. In agreement with 
Belém & Schlenz (1982), we adopted 
Schmidt’s classification because it provides 
descriptions and illustrations of the differ- 
ent types of corallimorpharian nematocysts, 
based on many specimens. 

The comparative study carried out with 
specimens of Discosoma carlgreni from 
Mexico and D. sanctithomae from Cuba has 
confirmed that they are conspecific with the 
Brazilian species, respectively. 

The varieties of b-rhabdoids' and b-rhab- 
doids? qualitatively separate Discosoma 


carlgreni from the D. sanctithomae. The 
first variety occurred particularly in the ten- 
tacles as well as in the column of both spe- 
cies, except in the discal tentacles of D. 
sanctithomae. Such absence was also veri- 
fied by den Hartog (1980). The b-rhab- 
doids? occurred particularly in the tentacles 
and column of D. carlgreni. Schlenz & Be- 
lém (1982) recorded two size-classes of b- 
rhabdoids in the stomodaeum of D. carl- 
greni, neither of them observed by den Har- 
tog (1980) nor in this study. 

The p-rhabdoids D' were easily seen in 
all structures of the body. The distribution 
and occurrence of these nematocysts 
seemed uniform in both species, except for 
the presence of only one small size class in 
the marginal tentacles of D. carlgreni as 
well as in the column of D. sanctithomae. 

Contrary to the view of den Hartog 
(1980) concerning the total absence of p- 
rhabdoid D in the stomodaeum in Coralli- 
morpharia, we observed a variety p-rhab- 
doid D' in this structure in both species. 
These nematocysts were rather common, 
also being larger in Discosoma sanctitho- 
mae. Our results are in agreement with 
those of Corréa (1964), who observed the 
microbasic p-mastigophore (=p-rhabdoid D 
sensu Schmidt) in the stomodaeum of D. 
sanctithomae. We also observed the variety 
p-rhabdoid D? in the column of D. sanctith- 
omae. None of the previous works have 
registered this variety before. 

Another feature which distinguishes Dis- 
cosoma sanctithomae from D. carlgreni is 
the size of holotrich I. In the column of D. 
sanctithomae, both large and small ones oc- 
cur, whereas in D. carlgreni they are small. 


142 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


We noticed in all discharged capsules of 
holotrich I, an abruptly tapered distal tube 
called terminal tube by den Hartog (1980). 
This terminal portion is flat, hardly refrac- 
tive and spineless, being also observed by 
Schlenz & Belém (1982). This portion can 
be observed in squash preparations, though 
sometimes it is not fully discharged so the 
terminal tube of most remains unevaginat- 
ed. The presence of this tube was previous- 
ly recognized in the macrobasic p-masti- 
gophores [=holotrich sensu Schmidt (1974) 
and penicilli E sensu den Hartog (1980)] by 
Cutress (1955:134). In his words: ‘“‘these 
nematocysts have a shaft which is more 
than three times the length of the capsule 
and which is abruptly reduced to a 
thread .. .”’. Cutress (1955), therefore, sug- 
gested that corallimorpharian holotrichs 
should be termed macrobasic p-mastigo- 
phores. Nevertheless, some authors (e.g., 
Werner 1965, Mariscal 1974) did not accept 
Cutress’ proposal. An alternative and in- 
dependently derived system, using Stephen- 
son’s term penicilli E, was provided by den 
Hartog (1980). Schmidt (1972, 1974) iden- 
tified a large holotrich I, commonly occur- 
ring in the filaments, stomodaeum and ten- 
tacles in the Corallimorpharia, as well as in 
the Scleractinia, as having the most distinc- 
tive spines possessed by anthozoan nema- 
tocysts. As indicated by Schmidt (1972, 
1974), this type presented a gradual taper- 
ing of the tubule. 

Studying the cnidae of four species of 
Brazilian Mussidae, Pires & Pitombo 
(1992) observed a holotrich I in the mes- 
enterial filaments. However, they did not re- 
cord in this type the abrupt end of the tube 
as a vestigial thread. In short, Pires & Pi- 
tombo (1992) observed a gradual tapering 
of the tubule, as mentioned by Schmidt 
(1974). 

den Hartog et al. (1993), who studied the 
corallimorpharians from the CANCAP ex- 
pedition, found penicilli E, especially in the 
filaments, as well as in the tentacles, of five 
species of Corallimorphidae. However, they 
did not observe the terminal tube in dis- 


charged capsules of penicilli E from the fil- 
aments of Corynactis sp. den Hartog et al. 
(1993) added that the previous observations 
on this type by Cutress (1955) and espe- 
cially by den Hartog (1980) needed confir- 
mation, considering the fact that the peni- 
cilli E were based on few occasions. 

In spite of the divergence among termi- 
nologies adopted in previous works, we still 
consider the type holotrich I sensu Schmidt 
(1972, 1974) the best term to be employed 
in this paper. To avoid further misunder- 
standing, the holotrichs I found in Disco- 
somatidae here has an abrupt tapered and 
spineless distal tip into a terminal tubule 
(the reason that den Hartog (1980) consid- 
ered them as penicilli E). 

The holotrichs II of the marginal tenta- 
cles of Discosoma carlgreni are in variably 
larger than those of D. sanctithomae. This 
nematocyst was also found by den Hartog 
(1980). According to den Hartog (1980) 
and Belém & Schlenz (1982), this type oc- 
curs only in the marginal tentacles. Never- 
theless, an unusual variety of holotrich, ob- 
served in the column and discal tentacles, 
showed size differences, especially in the 
column of D. sanctithomae. This variety 
had never been found before. We tentative- 
ly classify it as Schmidt’s holotrich II; fur- 
ther studies of its ultrastructure will provide 
a more comprehensive description. 

The results of this study demonstrate the 
importance of the nematocysts in distin- 
guishing Discosoma carlgreni from D. 
sanctithomae. The varieties of b-rhabdoids 
and p-rhabdoids D allow us to separate the 
species. 


Acknowledgments 


We are grateful to Dr. E. Schlenz and Dr. 
E L. da Silveira (Universidade de Sao Pau- 
lo, Sao Paulo) for their helpful comments 
to the manuscript. Thanks to Dr. A. Herrera 
(Instituto de Oceanologia, Academia de 
Ciencias de Cuba, ACC), Dr. E. Jordan 
(Universidade Nacional Autonoma de Méx- 
ico), Dr. E M. Amaral (Universidade Fed- 


VOLUME 113, NUMBER 1 


eral Rural de Pernambuco) and Dr. FE B. 
Pitombo (Universidade Federal Rural do 
Rio de Janeiro) for providing some of the 
samples used in this study. We also owe 
thanks to Dr. E Pitombo for the photograph 
of Figure 2B. Thanks to Parque Nacional 
Marinhos dos Abrolhos who provided fa- 
cilities and assistance during the collection 
of specimens. We wish to thank Conselho 
Nacional de Desenvolvimento Cientifico e 
Tecnol6gico (CNPq), Brazil for financial 
support. Part of this study was also sup- 
ported by FAPESP (Fundacgao de Amparo 
a Pesquisa do Estado de Sao Paulo), by a 
grant to the first author (no 98/3222-2). Fi- 
nally, we also grateful to Dr. Daphne Fautin 
(University of Kansas) and an anonymous 
reviewer who made invaluable suggestions 
that improved the manuscript. 


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Appendix 


List of specimens from the Museu Nacional do Rio 
de Janeiro Cnidaria collection used for this study. 
Discosoma carlgreni (Watzl, 1922) 

MNRJ. 1796—Brazil, Espirito, Santo Santa Cruz 
(19°49'08"S and 40°16'43”W), Aracruz, Estagao de 
Biologia Marinha, coll. S. M. Pinto, E B. Pitombo 
& EM. Amaral, 8 Aug 1990, 14 specimens, det. S. 
M. Pinto Aug 1990. 

MNRJ. 1866 and 1867—Brazil, Espirito Santo, Santa 
Cruz (19°49'08"S and 40°16'43”"W), Aracruz, Esta- 
¢ao de Biologia Marinha, coll. M. J. C. Belém & E. 


Schlenz, 8 Sep 1991, 14 specimens, det. M. J. C. 
Belém & E. Schlenz Sep 1991. 

MNRJ. 1868—Brazil, Espirito Santo, Guarapari 
(20°40'16”S and 40°28’5”W), Trés Praias, coll. M. J. 
C. Belém & E. Schlenz, 5 Sep 1991; 1 specimen, 
det. S. M. Pinto Sep 1991. 

MNRJ. 1878—Brazil, Espirito Santo, Santa Cruz 
(19°49'08"S and 40°16'43”W), Aracruz, Estagao de 
Biologia Marinha, coll. M. J. C. Belém, E. Schlenz 
& C. C. Ratto, 8 Sep 1991, 1 specimen, det: M. J. 
C. Belém Aug 1991. 

MNRJ. 2075—Brazil, Bahia, Abrolhos Archipelago 
(17°20’—18°10'S and 38°35’—39°20'W), Siriba Island, 
coll. E B. Pitombo & C. G. Fonseca, 15 Dec 1992, 
5 specimens, det. S. M. Pinto Dec 1992. 

MNRJ. 2250—Brazil, Bahia, Abrolhos Archipelago 
(17°20'—18°10'S and 38°35’—39°20'W), Siriba Island, 
coll. FE B. Pitombo,& C. C., Ratto, 19 Dec. 199358 
specimens, det. S. M. Pinto Dec 1993. 

MNRJ. 2259—Brazil, Bahia, Abrolhos Archipelago 
(17°20’-18°10'S and 38°35’—39°20'W), Chapeirao, 
coll. E B. Pitombo, 23 Dec 1993, 13 specimens, det. 
S. M. Pinto Dec 1993. 

MNRJ. 1540—México, Puerto Morelos, Quintana 
Roo, coll. E. Jordan Dahlgren & E D. Amaral, 24 
Oct 1989, 1 specimen, det: S: M- Pinto Dec 1989: 
Discosoma sanctithomae (Duchassaing & Michelotti, 
1860) 

MNRJ. 2076—Brazil, Bahia, Abrolhos Archipelago 
(17°20'—18°10'S and 38°35’—39°20'W), Siriba island, 
coll. EK B. Pitombo & C. G. Fonseca, 15 Dec 1992, 
22 specimens, det. S. M. Pinto Dec 1992. 

MNRJ. 2251 and 2252—Brazil, Bahia, Abrolhos Ar- 
chipelago (17°20’—18°10’S and 38°35’—39°20'W), 
Siriba island, coll. EK B. Pitombo & C. C. Ratto, 19 
Dec 1993, 20 specimens, det. S. M. Pinto Dee 1993. 

MNRJ. 2109—Cuba, La Habana, Playa, Playa Jaiman- 
itas, coll. A. Herrera, 26 Aug 1992, 4 specimens, det. 
M. J. C. Belém Feb 1994. 

MNRJ. 2110—Cuba, Recife de Punta del Este, Isla de 
Juventud, coll. A Herrera, 26 Aug 1992, 4 speci- 
mens, det. M. J. C. Belém Dec “1992. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


113(1):145—154. 2000. 


Additions to the cancellariid (Mollusca: Neogastropoda) fauna of 


South Africa 


Richard E. Petit and M. G. Harasewych 


Department of Invertebrate Zoology, National Museum of Natural History, 
Smithsonian Institution, Washington, D.C. 20560-0118, U.S.A. 


Abstract.—Four new species of Cancellariidae are described from the con- 
tinental shelf and upper continental slope off eastern South Africa. Admetula 
afra is distinguished from all congeners by its combination of small adult size 
(<11 mm), rounded shoulder, evenly reticulate sculpture, and distinct varix at 
the juncture of protoconch and teleoconch. Trigonostoma kilburni differs from 
all other Trigonostoma in its distinctive shell outline, unornamented peripheral 
keel, rounded rather than tabulate shoulder, and very narrow umbilicus. Nip- 
ponaphera wallacei differs from N. paucicostata (Sowerby, 1894), its geo- 
graphically closest congener from the Arabian Sea area, in being umbilicate 
and in having a more rounded shoulder, and more numerous and finer spiral 
cords. Its frequent association with the turbinid Bolma andersoni suggests that 
it may be an ectoparasite of this species. Zeadmete verheckeni is most similar 
to Zeadmete subantarctica Powell, 1933, from off New Zealand, from which 
it can be distinguished by its lower spire and weaker surface sculpture. Among 
the South African taxa, Z. verheckeni most closely resembles ‘‘Cancellaria”’ 
eutrios Barnard, 1959, from which it is easily distinguished by its tabulate 
shoulder and lower spire. The geographic and bathymetric ranges of Admetula 
epula Petit & Harasewych, 1991, a species previously known only from ‘“‘ex 


pisces”’ material, have been expanded based on live-collected specimens. 


The Cancellariidae comprises a family of 
diverse and highly specialized, suctorial 
neogastropods that inhabit soft bottom, sub- 
tidal to bathyal habitats throughout tropical 
and temperate seas. The cancellariid fauna 
of South Africa was reviewed comprehen- 
sively by Barnard (1959) and Kensley 
(1973), and to a limited extent more re- 
cently in popular works by Richards (1981) 
and Steyn & Lussi (1998). 

This paper describes four new species of 
cancellariids collected in South African wa- 
ters by SCUBA and by the vessels R/V 
Meiring Naudé (1984-1988) and NMPD 
Africana (1995). These species are assigned 
to the genera Admetula, Trigonostoma, Nip- 
ponaphera, and Zeadmete. The new taxa 
are compared to related species from the 
Indian Ocean, New Zealand, and Australia. 


All type specimens are housed in the col- 
lections of the Natal Museum (NM), Pie- 
termaritzburg, Republic of South Africa. 


Family Cancellariidae Forbes & Hanley, 
1851 
Genus Admetula Cossmann, 1889 


Type species: Cancellaria evulsa (Solan- 
der, 1766) (=Buccinum evulsum Solander, 
1766) by original designation. 


Admetula epula Petit & Harasewych, 1991 
Figs. 1-2, 18 


Admetula epula Petit & Harasewych, 
1991:181, figs. 1-3. 


Diagnosis.—A small species with an 
ovately conical shell. Transition from pro- 


146 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


toconch to teleoconch gradual, indicated by 
appearance first of spiral, then axial sculp- 
ture. Teleoconch sculpture of strong spiral 
cords and less pronounced, rounded axial 
ribs. Outer lip thin, smooth within. 

Gross anatomy.—Preserved animal yel- 
lowish tan, foot long, narrow, tapering pos- 
teriorly. Mantle cavity spanning 0.67 whorl. 
Osphradium slightly broader than ctenid- 
ium. Pericardium very small. Tentacles 
symmetrical, bluntly cylindrical, flanking 
central rostrum. Eyes small, black. Probos- 
cis short, about 0.67 length of mantle cav- 
ity. Buccal mass large, nearly filling retract- 
ed proboscis. Salivary glands and accessory 
salivary glands in cephalic haemocoel, not 
contained within proboscis. Penis long, nar- 
row, dorsoventrally compressed, distal end 
bluntly rounded. 

Remarks.—This species was originally 
described based on seven specimens taken 
from the stomachs of fish. Its bathymetric 
range was inferred to overlap with that of 
Congiopodus spinifer (Smith) (55—146 m), 
one of the fish from which it was taken. 
Among the material collected by the 
NMPD Africana [sta. A17419D] was a live 
collected specimen of A. epula (Figs. 1-2, 
18) trawled at a depth of 210 m off the 
mouth of the Tsitsikamma River. 

We have examined a specimen of Ad- 
metula from deeper water (450—500 m) off 
the Mbashe River, Transkei (R/V Meiring 
Naudé sta. Q14), between the type localities 
of A. epula and the new species. This spec- 
imen (NM C9050) is too worn for the tran- 
sition from protoconch to teleoconch to be 
clearly discerned, but there is no indication 
of a pronounced varix. Because it also has 
secondary spiral cords between all primary 
cords, this specimen is tentatively identified 
as A. epula. This record expands the geo- 
graphic range of A. epula from Cape St. 
Blaize to Transkei, off of the mouth of the 
Mbashe River (32°22.8’S, 29°00.8’E). The 
bathymetric range is extended well into the 
bathyal zone (450 m). 


Admetula afra, new species 
Figs. 3—4, 18 


Diagnosis.—A small species with a 
broadly conical shell. Protoconch demar- 
cated from teleoconch by broad, rounded 
varix. Teleoconch with sharply reticulated 
sculpture. Outer lip reflected posteriorly, 
with weak lirae beneath spiral cords. 

Description.—Shell (Fig. 3) small, reach- 
ing 9.0 mm, ovately conical with rounded 
anterior. Protoconch (Fig. 4) of 1.67 
smooth, inflated whorls deflected from coil- 
ing axis of teleoconch by about 7°. Transi- 
tion to teleoconch (Fig. 4, arrow) demar- 
cated by prominently rounded varix, fol- 
lowed immediately by onset of both spiral 
and axial sculpture. Teleoconch of 2.75—3 
evenly rounded whorls. Suture weakly im- 
pressed. Shoulder rounded, weakly defined. 
Axial sculpture consists of narrow, regular- 
ly spaced, weakly prosocline ribs (14-16 
on body whorl). Spiral sculpture of narrow, 
sharply defined primary cords (12-13 on 
body whorl, 6 on penultimate whorl). Sin- 
gle, weak, secondary cords may be present 
between primary cords on posterior half of 
body whorl. Spiral and axial sculpture in- 
tersect to form sharply reticulated pattern 
with small nodes at intersections of ribs and 
cords. Aperture broadly ovate, deflected 
from coiling axis by 16—19°. Outer lip thin, 
slightly flared posteriorly, weakly lirate be- 
neath spiral cords. Parietal callus thin, 
translucent, overlying 5—6 spiral cords from 
previous whorl. Columella forming angle of 
128-130° with parietal region, thick, 
straight, with two weak columellar folds 
and broad siphonal fold. Siphonal canal 
shallow but well-defined. Shell color white. 

Type locality.—SE of Port Durnford, 
South Africa (29°01.5'S, 32°11.8’E), dredged 
in 310-320 m, glutinous sandy mud. R/V 
Meiring Naudé sta. ZQ9, 6 Jul 1985. 

Type material.—Holotype, NM_ E3189/ 
T1382, 8.0 mm; Paratype 1, 7.8 mm, and 
Paratype 2, 6.9 mm, NM V7006/T1383 from 
the type locality; Paratype 3, 9.0 mm, NM 
E3764, from off Cape Vidal (28°08.4'S, 


VOLUME 113, NUMBER | 


Figs. 1-4. 1-2; Admetula epula Petit & Harasewych, 1991. Off Tsitsikamma River, South Africa (34°45'S, 
24°47'E) in 210 m, sand, old shell grit and shell debris. NMPD Africana sta. A17419D. 1. Apertural view of 
shell. 2. Apical view of protoconch. 3—4. Admetula afra new species. Holotype, NM E3189/T1382, SE of Port 
Durnford, South Africa (29°01.5’S, 32°11.8’'E), dredged in 310—320 m, glutinous sandy mud. R/V Meiring Naudé 
sta. ZQ9, 6 Jul 1985. 3. Apertural view of holotype. 4. Apical view of protoconch. 


32°36.4'E), dredged in 165 m, moderately Comparative remarks.—This new spe- 
fine sand, R/V Meiring Naudé sta. ZM8, 11 cies appears most closely related to the 
June 1988. more southern Admetula epula Petit & Har- 

Etymology.—Feminine form of the Latin asewych, 1991, from which it differs in 
afer, African. having uniformly rectangular sculpture 


148 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


formed by equally-sized axial ribs and spi- 
ral cords, and a posteriorly reflected outer 
lip with weak denticles beneath the spiral 
cords. The most striking difference between 
these two species is the presence of a strong 
varix at the termination of the protoconch 
in A. afra. In contrast, the transition from 
protoconch to teleoconch is indistinct and 
gradual in Admetula epula. 


Genus Trigonostoma Blainville, 1827 


Type species: Delphinula trigonostoma 
Lamarck, 1822 (?=Buccinum scalare Gme- 
lin, 1791) by monotypy. 


Trigonostoma kilburni, new species 
Figs. 5-10, 18 


Diagnosis.—A small species with a thin, 
angular, narrowly umbilicate shell. Shoul- 
der rounded, not tabulate, lacking pro- 
nounced spines. Axial sculpture of numer- 
ous scabrous varices. Outer lip smooth, 
lacking lirae. 

Description.—Shell (Fig. 5) small, reach- 
ing 14.3 mm, thin, angular, biconical, 
strongly shouldered, with deep, narrow um- 
bilicus. Spire high (spire angle 55°), com- 
prising over half of shell length. Protoconch 
(Figs. 6—7) of 2 smooth whorls, offset from 
coiling axis of shell by about 5°. Transition 
to teleoconch abrupt, marked by a slightly 
flared varix and the onset of spiral sculp- 
ture. Teleoconch of up to 5 sharply angular 
whorls. Suture deeply impressed behind the 
evenly rounded shoulder delineated by a 
sharp keel along the periphery of the shell. 
Axial sculpture consists of numerous, 
flared, weakly prosocline varices (Fig. 8), 
regularly spaced on early whorls (about 16— 
18 on first teleoconch whorl), increasing in 
number and becoming more irregularly 
spaced in subsequent whorls. Spiral sculp- 
ture of broad, crisply demarcated primary 
spiral cords (Fig. 8, p), with 1-3 slightly 
narrower secondary cords (Fig. 8, s) be- 
tween adjacent primary cords, and much 
finer spiral threads (Fig. 8, t) between some 
cords. Aperture roundly triangular, deflect- 


ed from coiling axis by 18—19°. Siphonal 
canal short, broad, barely discernible except 
externally as the siphonal fasciole. Outer lip 
thin, smooth within, slightly reflected with 
spiral cords visible through edge of lip. 
Posterior portion of inner lip adpressed 
against siphonal fasciole. Short parietal re- 
gion forms angle of 150° with long, slightly 
concave columella that bears 2 weak, wide- 
ly spaced columellar folds and 1 siphonal 
fold. Umbilicus deep, narrow. Shell white, 
sometimes with a yellowish cast. 

Radular teeth (Figs. 9-10) extremely 
long, ribbon-like, tricuspid. Central cusp 
smooth, with recurved rim. Lateral cusps 
long, folded toward central cusp, each with 
four, anteriorly directed secondary cusps. 
Second most proximal secondary cusp bi- 
fid. 

Type locality.—Off East London, South 
Africa (33°04.7'S, 28°07.2’E) dredged in 90 
m, associated with coarse sand, sponges, 
gorgonians. R/V Meiring Naudé sta. XX46, 
17 Jul 1984. 

Type material.—Holotype, NM D679, 
12.9 mm, from type locality. Paratype, NM 
D680, 14.4 mm, off Kidd’s Beach, South 
Africa (33°11.8'S, 28°03.2’E) dredged in 90 
m, associated with coarse sand, sponges. R/ 
V Meiring Naudé sta. XX50, 17 Jul 1984. 

Etymology.—This species honors Dr. Ri- 
chard N. Kilburn, Natal Museum, Pieter- 
maritzburg, Republic of South Africa, for 
his many contributions to malacology. 

Comparative remarks.—This new spe- 
cies differs from all other Trigonostoma in 
its distinctive shell outline. It may be dis- 
tinguished from Trigonostoma scalare 
(Gmelin, 1791) and T. thysthlon (Petit & 
Harasewych, 1987) by its unornamented 
peripheral keel, its rounded rather than tab- 
ulate shoulder, and by its very narrow um- 
bilicus. The only other South African Tri- 
gonostoma is the common shallow-water T. 
semidisjuncta (Sowerby, 1849), which has 
a heavier, more rounded shell with strong 
spiral cords. 


VOLUME 113, NUMBER I 


149 


Figs. 5-8. 
(33°04.7'S, 28°07.2'E), dredged in 90 m, coarse sand, sponges, gorgonians. R/V Meiring Naudé sta. XX46, 17 
Jul 1984. 5. Apertural and lateral views of holotype. 6. Apical and 7. Lateral views of protoconch. 8. Detail of 
surface sculpture on body whorl. p, primary spiral cords; s, secondary spiral cords; t, spiral threads. 


Genus Nipponaphera Habe, 1961 


Type species: Nipponaphera habei Petit, 
1972 by I.C.Z.N. Opinion 1052. 


Nipponaphera wallacei new species 
Figs. 11-13, 16 


Diagnosis.—A small species with a 
heavy, strongly sculptured, narrowly um- 
bilicate shell. Aperture sharply triangular, 
outer lip with strong teeth along inner edge 


Trigonostoma kilburni new species. Holotype, NM D679, Off east London, South Africa 


of varix. Columella with two prominent, 
Sharply keeled columellar folds and a 
strongly reflected siphonal fold. 
Description.—Shell (Fig. 11) small for 
genus, to 12.4 mm, ovately conical, with 
rounded anterior. Spire relatively short 
(spire angle 70°), comprising less than half 
of shell length. Protoconch (Fig. 13) of 1.67 
smooth, inflated whorls. Transition to teleo- 
conch marked by onset of spiral cords fol- 
lowed immediately by strong axial ribs. Te- 


150 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Figs. 9-10. Trigonostoma kilburni new species. Distal ends of radular teeth of holotype. 9. Lateral view. 


10. End-on view. 


leoconch of up to 4 angular whorls. Suture 
impressed behind sloping shoulder that is 
delineated by peripheral keel. Axial sculp- 
ture of strong, broad, prosocline ribs (10 on 
penultimate whorl) that become more wide- 
ly spaced on body whorl where they appear 
as varices with flared edges. Spiral sculp- 
ture of broad, flattish primary cords, 3 sec- 
ondary cords between adjacent primary 
cords, with fine threads between some 
cords. Aperture sharply triangular, deflected 
from coiling axis by 20°. Siphonal canal 
short, well-defined. Outer lip reflected an- 
teriorly, with spiral sculpture visible 
through thin edge, 9 strong teeth along in- 
ner edge of varix, small, angular indenta- 
tion at shoulder. Parietal region short, col- 
umella straight, with small, thin callus, 2 
Sharply keeled columellar folds and strong- 
ly reflected siphonal fold. Umbilicus nar- 
row, inconspicuous, bordered by well-de- 
veloped, cord-like siphonal fasciole. Shell 
color chestnut brown, with lighter spiral 
bands along shoulder and middle of body 
whorl. 

Type locality.—Off Phumula, Natal, 
South Africa, in 45 m, on reef. Jun 1997. 

Type material.—Holotype, NM V4689, 
6.6 mm, from type locality. Paratype 1, NM 
V3889, Off Phumula, Natal, South Africa, 
in 35 m, taken by SCUBA, living on shell 
of Bolma andersoni (E. A. Smith, 1902). 4 


Sep 1996. Paratype 2, NM V303, 12.4 mm, 
Off Park Rynie, Natal, South Africa, in 53 
m. Taken by SCUBA, living on the spire of 
the turbinid gastropod Bolma andersoni. 19 
May 1990. 

Etymology.—This species is named for 
Mr. Martin Wallace, who collected the type 
material and generously made it available 
for study. 

Comparative remarks.—The genus Nip- 
ponaphera is distinguished from Trigonos- 
toma primarily by the presence, in the latter, 
of an open umbilicus that extends back to 
the protoconch. Also, Nipponaphera has a 
wide and flat siphonal fold. Although the 
new species here described has an umbili- 
cus, it is not profound. It differs from N. 
paucicostata (Sowerby, 1894) of the Ara- 
bian Sea area in being umbilicate, in having 
a more rounded shoulder, and more and fin- 
er spiral cords. Specimens of N. paucicos- 
tata figured by Verhecken (1986, Figs. 4— 
6) lack varix-like axial ribs on the body 
whorl, but the type specimen has very 
strong, unevenly spaced ribs on the body 
whorl. The Japanese N. teramachii (Habe, 
1961) has a sharp keel, is umbilicate, with 
a cord-like siphonal fasciole, but lacks the 
squarish, finely imbricated spiral sculpture 
of this new species. Melvill and Standen 
(1901:451) reported N. paucicostata from 
the Gulf of Aden ‘adhering to the upper 


VOLUME 113, NUMBER 1 


Figs. 11-13. 
SCUBA in 45 m, on reef. Jun 1997. 11. Apertural view of holotype. 12. Detail of sculpture on body whorl. 13. 
Apical view of protoconch. 


part of Rapana bulbosa, 30—50 fathoms.” 
The alimentary system of cancellariids is 
adapted to feed on body fluids of prey/host 
organisms (Petit & Harasewych, 1986; Har- 
asewych & Petit, 1986). The association of 
species of Nipponaphera with other gastro- 
pods suggests that this group of cancellar- 
lids may be specialized ectoparasites of 
large gastropods. 


Genus Zeadmete Finlay, 1926 


Type species: Cancellaria trailli Hutton, 
1973 by original designation. 


Zeadmete verheckeni, new species 
Figs. 14-18 


Diagnosis.—A small species with an 
ovate shell. Shoulder tabulate. Surface 
sculpture dominated by spiral cords. Pseu- 
do-umbilicus narrow. Outer lip thin, smooth 
within. Buccal mass minute. Radula absent. 

Description.—Shell (Fig. 14) small, 
reaching 7.9 mm, thin, with stepped spire, 
rounded anterior. Protoconch (Figs. 15—16) 
erect, smooth, of 1.5 whorls. Transition to 


Nipponaphera wallacei new species. Holotype, NM V4689, Off Phumula, Natal, South Africa, 


teleoconch abrupt, marked by onset of 
weak, closely-spaced axial ribs that become 
stronger, more regularly spaced, when spi- 
ral cords first appear within 0.25 whorl. Te- 
leoconch of 2 rounded, strongly tabulate 
whorls. Suture strongly impressed. Axial 
sculpture of evenly spaced, well-defined, 
axially aligned ribs (30 on body whorl) as 
broad as intervening spaces. Spiral sculp- 
ture of sharply demarcated evenly spaced 
cords (2 between suture and shoulder, 17 
below shoulder) diminishing in strength 
abapically, forming small nodules as they 
cross axial ribs. Aperture elongated, nar- 
rowly elliptical. Outer lip faintly sinuate, 
smooth within. Columella with two broad, 
weak, columellar folds (Fig. 17, pcf, acf) 
and siphonal fold (Fig. 17, sf). Siphonal ca- 
nal small but distinct, axially aligned, not 
forming siphonal fasciole. Anterior portion 
of inductura bordered by weak parietal 
wash, partially covering narrow pseudo- 
umbilicus. Shell color white. Periostracum 
thin, finely lamellate, straw-colored. 
Preserved animal white, with short, nar- 
row, posteriorly rounded foot. Tentacles tu- 


[52 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


1 mm 


sf 
500 um 


“ 
og ge 
3 a 
_ 


Figs. 14-17. 


Zeadmete verheckeni new species. Holotype, NM C6800, Off Stony Point, Transkei, South 
Africa (32°37.5'S, 28°45.8’E), dredged in 390—400 m, muddy sand, small stones. R/V Meiring Naudé sta. V11, 
12 Jul 1984. 14. Apertural and dorsal views of holotype. 15. Apical and 16. Lateral views of protoconch. 17. 


Columella. pcf, posterior columellar fold; acf, anterior columellar fold; sf, siphonal fold. 


bular, symmetrical, with very large black 
eyes at their bases. Penis long, narrow, dor- 
so-ventrally flattened, with small terminal 
papilla. Osphradium very broad, strongly 
asymmetrical, dorsal leaflets twice as broad 
as ventral leaflets. Ctenidium less than half 
as wide and twice as long as osphradium. 
Hypobranchial gland large, glandular. Re- 
tracted proboscis occupies anterior two- 


thirds of cephalic haemocoel, the rest oc- 
cupied by long, convoluted mid-esophagus. 
Proboscis strongly coiled within proboscis 
sheath. Extended proboscis likely exceeds 
shell length. Proboscis thin, with minute 
buccal mass in anteriormost 0.125 of pro- 
boscis. Radula absent. 

Type locality.—Off Stony Point, Trans- 
kei, South Africa (32°37.5’S, 28°45.8’E), 


VOLUME 113, NUMBER 1 


015° E 


020° E 


Fig. 18. 


025° E 


ee 


030° E 035° E 


Geographic distributions of South African Cancellariidae. Admetula epula Petit & Harasewych, 


1991, vertically hatched area = type locality; open squares = new records. Admetula afra new species, star in 
circle = type locality; filled circle = additional record. Trigonostoma kilburni new species, black star = type 
locality; open circle = additional record. Nipponaphera wallacei new species, triangle = type locality. Zeadmete 


verheckeni new species, diamond = type locality. 


dredged in 390—400 m, muddy sand, small 
stones. R/V Meiring Naudé sta. V11, 12 Jul 
1984. 

Type material.—Holotype, NM C6800, 
5.1 mm, from type locality. 

Etymology.—Named for Mr. André Ver- 
hecken, Mortsel, Belgium, in recognition of 
his contributions to the study of the Can- 
cellariidae. 

Comparative remarks.—Placement of 
this new species in the genus Zeadmete is 
tentative. Zeadmete verheckeni is concho- 
logically most similar to a group of species 
from off the southern coasts of New Zea- 
land and Australia that have been assigned 
to the genera Oamaruia Finlay, 1924 and 
Zeadmete Finlay, 1926. Powell (1979:224) 
treated Zeadmete as a subgenus of Oama- 
ruia. Several New Zealand and Australian 
species presently assigned to Zeadmete 
agree with this South African species in 
Shell form and sculpture (see Garrard 1975, 
Powell 1979). Zeadmete subantarctica 


Powell, 1933, from 50 fathoms off Snares 
Islands, New Zealand, has the same shell 
shape and columellar structure as Z. ver- 
heckeni but has deeply cancellated sculp- 
ture on the posterior half of the body whorl 
and only spiral cords on the anterior por- 
tion. 

In the course of a study on Australian 
cancellariids, we found that the abyssal 
Zeadmete kulanda Garrard, 1975 has a rad- 
ula similar to that of Nothoadmete tumida 
(Oliver, 1982:figs. 3, 5). The fact that no 
radula was found in Z. verheckeni argues 
against these species being congeneric. 
However, we are reluctant to introduce an- 
other genus-level taxon until more data on 
the species with this shell form are avail- 
able. The problems of generic placement of 
small, deep-water cancellariids was briefly 
discussed by Verhecken (1997:296). 

Among the South African taxa, this spe- 
cies most closely resembles ‘‘Cancellar- 
ia’ eutrios Barnard, 1959, from which it is 


154 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


easily distinguished by its tabulate shoulder 
and lower spire. 


Acknowledgments 


We thank Dr. Richard N. Kilburn for 
making these specimens available and for 
his patience in waiting for their description. 
Correspondence and discussion with Mr. 
André Verhecken of Mortsel, Belgium add- 
ed to our knowledge and was helpful in pre- 
paring this paper. 


Literature Cited 


Barnard, K. H. 1959. Contributions to the knowledge 
of South African marine Mollusca. Part II. Gas- 
tropoda: Prosobranchiata: Rachiglossa.—An- 
nals of the South African Museum 45:1—237. 

Garrard, T. A. 1975. A revision of Australian Cancel- 
lariidae (Gastropoda: Mollusca).—Records of 
the Australian Museum 30(1):1—62. 

Harasewych, M. G., & R. E. Petit. 1986. Notes on the 
morphology of Admete viridula (Gastropoda: 
Cancellariidae).—The Nautilus 100(3):85—91. 

L.C.Z.N. 1976. Opinion 1052. Nipponaphera Habe, 
1961 (Gastropoda): Designation of a type-spe- 
cies under the plenary powers.—Bulletin of 
Zoological Nomenclature 32(4):242—243. 

Kensley, B. 1973. Sea-Shells of southern Africa, Gas- 
tropods. Maskew Miller Ltd., Cape Town, 236 


PP. 


Melvill, J. C., & R. Standen. 1901. The Mollusca of 
the Persian Gulf, Gulf of Oman, and Arabian 
Sea, as evidenced mainly through the collec- 
tions of Mr. EF W. Townsend, 1893-1900; with 
descriptions of new species.—Proceedings of 
the Zoological Society of London for 1901: 
327-460, pls. 21-24. 

Oliver, P. G. 1982. A new species of cancellariid gas- 
tropod from Antarctica with a description of the 
radula.—British Antarctic Survey Bulletin 57: 
15-20. 

Petit, R. E., & M. G. Harasewych. 1986. New Philip- 
pine Cancellariidae (Gastropoda: Cancellari- 
acea), with notes on the fine structure and func- 
tion of the Nematoglossan radula.—The Veliger 
28(4):436-443. 

Powell, A. W. B. 1979. New Zealand Mollusca. Wil- 
liam Collins, Auckland, xiv + 500 pp. 

Richards, D. 1981. South African seashells. A collec- 
tor’s guide. C. Struik Publishers, Cape Town. 98 
pp. + 60 pls. 

Steyn, D. G., & M. Lussi. 1998. Marine shells of South 
Africa. Ekogilde Publishers, Hartebeespoort. 
264 pp. 

Verhecken, A. 1986. A revision of the Cancellariidae 
(Neogastropoda: Cancellariacea) of the Red Sea 
and the Gulf of Aden.—Gloria Maris 25(4): 
133-153. 

Verhecken, A. 1997. Mollusca Gastropoda: Arafura 
Sea Cancellariidae collected during the KA- 
RUBAR Cruise. Pp. 295-323 in A Crosnier & 
P. Bouchet, eds., Résultats des Campagnes MU- 
SORSTOM, vol. 16. Mémoirs du Muséum Na- 
tional d’ Histoire Naturelle 172. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


113(1):155—-161. 2000. 


Erpobdella lahontana (Annelida: Hirudinea: Arhynchobdellida: 
Erpobdellidae), a new species of freshwater leech from 
North America 


Peter Hovingh and Donald J. Klemm 


(PH) 721 Second Avenue, Salt Lake City, Utah 84103, U.S.A.; 
(DJK) U.S. Environmental Protection Agency, National Exposure Research Laboratory, 
Ecological Exposure Research Division, Ecosystems Research Branch, 
26 W. Martin Luther King Drive, Cincinnati, Ohio 45268, U.S.A. 


Abstract.—new species of a leech, Erpobdella lahontana, is described from 
the Lahontan Basin in California and Nevada of the western United States. 
This species has four pairs of eyes, the preatrial loops of male paired ducts 
extend to ganglion XI, and the male and female gonopores are located in 
furrows of the annuli, separated by five annuli. 


The zoological history of the Great Basin 
of the western United States has resulted in 
an area of high endemism of fishes (Hubbs 
& Miller 1948) and hydrobiid snails 
(Hershler 1998). However, amphibians have 
not morphologically evolved into endemics 
and may be the result of very recent im- 
migration to the region (Hovingh 1997). 

A general survey of the Great Basin and 
adjacent regions of the western United 
States was undertaken to determine if 
leeches had drainage specific distributions. 
Over 2300 aquatic sites were surveyed in 
the Great Basin including some 600 sites 
within the Lahontan Basin. Leeches show 
an entirely different pattern than fishes, hy- 
drobiid snails, and amphibians with respect 
to basin specific distribution without mor- 
phological evolution, suggesting evolution- 
ary stasis and an inhabitant of the Great Ba- 
sin since its geological formation in the 
Miocene (Hovingh, unpublished data). 
Within this study, an erpobdellid leech was 
identified with the gonopores separated by 
five annuli. This leech was found in two 
separate drainages of the Honey Lake Sub- 
basin (Eagle Lake and one other location 
out of the 2300 sites in the survey) in the 
Lahontan Basin of northeastern California. 
This paper describes this new leech species 


and is the first endemic species of leech to 
be found in the Great Basin. 


Materials and Methods 


Collection methods consisted of exam- 
ining the underside of substrates such as 
rocks, logs, and anthropogenic debris in the 
periphery of aquatic systems (i.e., springs, 
streams, and lakes) up to 100 cm deep. 
Leeches were relaxed with dilute ethanol, 
wiped clean of mucous, fixed in 10% for- 
malin overnight, and preserved in 70% eth- 
anol. Histological examination of serial sec- 
tions of the clitellum region, after staining 
with hematoxylin and eosin, was used for 
detailed morphological analysis of the gen- 
ital atrium, cornua, and preatrial loops of 
the ejaculatory ducts. A model of the gen- 
ital atrium, cornua, and preatrial loops was 
constructed from photographs taken of the 
serial sections. 


Systematics 


Family Erpobdellidae Blanchard, 1894 
Genus Erpobdella Blainville, 1918 
Erpobdella lahontana, new species 


Type material.—Holotype, United States 
National Museum (USNM 186409) and 8 


156 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


paratypes (USNM 186410), deposited in 
the National Museum of Natural History, 
Division of Worms, Smithsonian Institu- 
tion, Washington, D.C., collected 15 Sep 
1997, same locality. 

Type locality.—Eagle Lake, Lassen 
County, California; latitude 40°33.5’N and 
longitude 120°48.8’W (selected by the 
abundance of leeches at this locality and 
public access to the lake). 

Diagnosis.—Dorsal surface dark gray, 
heavily mottled with light gray spots; white 
or yellow irregular minute, transverse rows 
of papillae on the dorsal side of every an- 
nulus with some living specimens having 
prominent papillae on both dorsal and ven- 
tral sides; dorsum with a light black mid- 
dorsal stripe or plain; five-annulate, each 
annulus of approximately equal size; four 
pairs of eyes: first pair of labial eyes large, 
second pair of smaller labial eyes behind 
first pair; buccal eyes two pairs, small; male 
and female gonopores separated by five an- 
nuli, located in furrows of segments XII 
and XIII and male gonopore large, raised, 
and especially glandular (Fig. 1); atrium 
with cornua (horns), with sperm ducts 
forming paired preatrial loops extending an- 
teriorly to ganglion XI (Figs. 2, 3). Preatrial 
loops angle laterally and anteriorly from 
cornua. Cornua and preatrial loops dip ven- 
trally to join each other. Ovisacs extend 
posteriorly to ganglion XV. Five annuli sep- 
arate the segmental ganglia. 


Description of Holotype 


External anatomy.—(based on holotype, 
USNM 186409): Body elongate, flattened, 
sides of body narrowing along most of 
length to pointed head (length 30 mm, max- 
imum width 3 mm); body wall uniformly 
smoke-gray, dorsal surface darker gray than 
ventral surface, dorsal surface with a faint 
middorsal dark line, no black pigmentation; 
dorsal surface dark gray, heavily mottled 
with light gray spots; somites five-annulate; 
white irregular minute, transverse rows of 
papillae on every annulus; clitellum con- 


<— 5mm —P 


Fig. 1. Erpobdella lahontana. Ventral view, male 
and female gonopores (holotype). 


spicuous, 15 annulate; nephridiopores on b, 
annulus of XI and XII (other nephridio- 
phores not determined); mouth small (width 
of oral sucker 1 mm); eyes four pair, one 
large labial pair, second pair of small labial 
eyes behind first pair and two smaller buc- 
cal pairs on fourth annulus (further poste- 
rior); anus surrounded by papillae, located 
dorsally at the base of the caudal sucker; 
caudal sucker small (width 2 mm), less than 
one-half maximum body width; male gon- 
opore large, raised, cylindrical and glandu- 
lar, (Fig. 1) located in furrow of the clitel- 
lum region between annuli XIIb, and XIIb,; 
distance from mouth to male gonopore, 14 
mm; female gonopore inconspicuous, lo- 
cated in furrow between annuli XIIIb, and 
XIIb,. 

Internal anatomy.—(based on dissection 
of paratypes): Atrium wider than long; atri- 
al horns projecting anteriorly; preatrial 
loops of vas deferens extending to ganglion 
XI; ovisacs extend from segments XIII to 
XV. 

Additional observations of paratypes 
(USNM_ 186410).—External anatomy. 
Paratypes resemble the holotype with the 
following additional observations: length 


VOLUME 113, NUMBER 1 


157 


Fig. 2. 
Ganglion XI (denoted A) and Ganglion XII (denoted D) showing the preatrial loops (denoted B) and the genital 
atrium (denoted C) with cornua. The angle in which the preatrial loops join the cornua is indicated by short 
bars: 30° for the left side and 45° for the right side as viewed from the dorsum. Note that the right cornua 
displays an oval cross-section representing the section that has turned ventrally. Long bar = | mm. 


(42—55 mm) and width (3.5—5 mm) of some 
individuals varies from the holotype; clitel- 
lum is inconspicuous in some individuals; 
female gonopore difficult to locate in some 
individuals; anus small to large in some 
paratypes. 

Remarks.—The eyes in four pairs are ar- 
ranged in two transverse rows; mouth with 
muscular ridges but no jaws; body seg- 
ments (somites) five-annulate with all an- 
nuli of approximately equal in width. The 
structure of the reproductive system shows 
that this species belongs to the genus Er- 
pobdella Blainville, 1918, with preatrial 
loops of the male paired ducts and testes in 
small, numerous bunched sacs. 

Distribution, habitat, and ecology.—Er- 
pobdella lahontana is found in two subba- 
sins of the western Lahontan Basin (Fig. 4): 
Eagle Lake and Grasshopper Valley in Las- 


Photograph of horizontal (parallel to dorsal and ventral surfaces) section of the region between 


sen County, California. The Pleistocene 
precursor of Eagle Lake and Grasshopper 
Valley (Madeline Plains) drained into Hon- 
ey Lake Subbasin, the latter being an arm 
of Pleistocene Lake Lahontan. The new 
species was found in Buck Bay in the north 
part of Eagle Lake and along the south 
shore of Eagle Lake. In Eagle Lake, the 
leech was often found on vesicular basaltic 
rocks. When removing the leech, the pos- 
terior sucker would remain on the rocks. 
This occurred with five leeches after which 
sampling was from smoother artificial sub- 
strate. The leech was found in the outflow 
of springs in Grasshopper Valley. Eleva- 
tions varied from 1555 to 1625 m above 
mean sea level (msl). Eagle Lake and, to a 
lesser extent, the Grasshopper Valley were 
associated with ponderosa pine (Pinus pon- 
derosa) forest and both belong to the Mo- 


158 


Fig: 3. 
ejaculatory ducts. Legend same as in Figure 2. Draw- 
ing made from a model constructed from photographs 
of serial sections of this region. The arrows denote 
where the cornua (paired horns) and the preatrial loops 
both turn ventrally before they are joined. The right 
cornua also makes a turn to the center. 


Dorsal view of male genital atrium and 


doc Plateau geomorphic province, a region 
of numerous basalt-andesite flows which 
isolated these two basins from Honey Lake 
(Gester 1962). Eagle Lake (12,150 ha) has 
three subbasins of which two (north and 
central) basins do not stratify due to their 
Shallow nature and strong winds. Conduc- 
tivity of Eagle Lake is near 800 pMbhos, 
probably as a result of the fact that 40% of 
Eagle Lake’s inflow is derived from ground 
water (Huntsinger & Maslin 1976). Eagle 
Lake contains five species and Madeline 
Plains contains one species of native fish. 
The ecology of Eagle Lake is further de- 
scribed by Huntsinger & Maslin (1976). 
Two other locations in the Lahontan Basin 
(Winnemucca and Granite Springs Subba- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


sins, east of Pyramid Lake, Nevada) con- 
tained leeches in which the gonopores were 
separated by five annuli, and these popula- 
tions are still under investigation. 

Associated leech species.—Erpobdella 
lahontana was associated with the leeches 
Erpobdella punctata (Leidy, 1870), Helob- 
della stagnalis (Linnaeus, 1758), and Hae- 
mopis marmorata (Say, 1824). In Grass- 
hopper Valley, H. marmorata was associ- 
ated with E. lahontana. Eagle Lake speci- 
mens at the National Museum of Natural 
History, Worm Division, include: E. punc- 
tata (USNM 42502, USNM 60061, USNM 
60062), Glossiphonia complanata (USNM 
2572), Placobdella ornata (USNM 60055), 
Theromyzon trizonare (USNM 42500), and 
H. marmorata (USNM 42570). The leech 
specimen (USNM 42571) was classified as 
Dina fervida, but this specimen is most 
likely E. lahontana. The gonopore separa- 
tion was not unresolved, but the pigmenta- 
tion and number of eyes suggested this spe- 
cies. 

Variations.—All Eagle Lake specimens 
(22) were identical with respect to eyes and 
the number of annuli between gonopores. 
Some specimens contained dark annuli in 
which the papillae became conspicuous 
with an even darker dorsal strip, and this 
varied to uniform light color with invisible 
papillae and with no dorsal strip. Sizes of 
leech specimens were up to 55 mm long 
and 3.5 mm wide. 

Similar species.—Erpobdella lahontana 
is distinguished from E. dubia (Moore & 
Meyer, 1951), E. parva (Moore, 1912), and 
E. punctata (Leidy, 1870) by the five annuli 
separating the gonopores. Erpobdella dubia 
(dorsum greenish, heavily mottled, usually 
with a variable dark mid-dorsal stripe; gon- 
opores separated by 3.5—4 annuli) and E. 
parva (dorsum unpigmented or color uni- 
formly smokey gray; gonopores separated 
by 2.5—3.5 annuli) have not been observed 
in the Great Basin collections with the gon- 
opores separated by five annuli. If the gon- 
opores are not distinguishable, the pigmen- 
tation patterns of E. lahontana could lead 


VOLUME 113, NUMBER 1 159 


LL a a a a a a a LF SBT IES ae: | 


Lassen 5 


Madeline Plains 


ho” 


Fig. 4. Map of the western Lahontan Basin in California and Nevada. Lassen County is enclosed and the 
communities of Susanville, California and Reno, Nevada are designated by diamonds. The heavy solid line 
denotes the Lahontan Basin with arrows showing Pleistocene drainages from Eagle Lake and Madeline Plains 
Subbasins. The light solid line shows the extent of Pleistocene lakes with the patterned horizontal lines showing 
the present Eagle, Honey, and Pyramid Lakes. The three solid circles show location of Erpobdella lahontana in 
Eagle Lake and in Grasshopper Valley of the Madeline Plains Subbasin. Inset shows the locations in relation to 
the Great Basin and the western United States. Map derived from Snyder et al. (1964). 


160 


to some confusion with E. dubia and E. 
parva, neither of which has been found in 
the Lahontan Basin. E. punctata has been 
found in the Great Basin, but this species 
can be distinguished easily from E. lahon- 
tana by having only two annuli separating 
the male and female gonopores, and having 
three pairs of eyes. Taxonomic discussion 
and keys to the family Erpobdellidae and 
these North American species can be found 
in Sawyer (1972, 1986b), Klemm (1985, 
1990, 1995) and Davies (1991). 


Discussion 


Similar species, Erpobdella dubia and E. 
parva, to the new species, E. lahontana, 
have been historically classified within the 
genus Dina (Soos 1963, 1966, 1968; Saw- 
yer 1972, Klemm 1985,,Davies 1991):-The 
genus (or subgenus) Dina is identified by 
the fact that every fifth annulus (b,) of a 
somite is distinctly wider or more broad 
than the other four annuli and is subdivided 
by a faint transverse furrow (Lukin 1976, 
Mann 1982, Sawyer 1986a, 1986b; Davies 
1991, Nesemann 1995, Neubert & Nese- 
mann 1995). Also, the genus Dina is re- 
ported from only the southern, western, and 
central areas of the Palaearctic region (Lu- 
kin 1976, Sawyer 1986b). Therefore, Saw- 
yer (1986a, 1986b) revised the genus Er- 
pobdella to include the two species of Dina 
from the Neartic region, including North 
America, based on the equal width of all 
the body annuli and distribution of E. dubia 
and E. parva. It was concluded that the ge- 
nus Dina was not found in North America. 
This was later confirmed by Klemm (1990, 
1995). The North America (Neartic region) 
genera Nephelopsis, Mooreobdella, and the 
Palearctic region genus Dina were grouped 
together by the feature that, of any five an- 
nuli of the body somites, there is one an- 
nulus that is distinctly wider than the other 
four annuli and usually subdivided by a 
faint transverse furrow (Mann 1962, Lukin 
1976, Sawyer 1986b, Davies 1991). This 
anatomical feature was not seen in Great 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Basin species of Erpobdellidae. All the spe- 
cies of erpobdellids in North America have 
annuli of similar width. 
Etymology.—Louis Armand de Lom 
d’ Arce, Baron de Lahontan was a 17th cen- 
tury French explorer of the upper Missis- 
sippi River basin who suggested that the 
western continental flows drained into a sa- 
line lake and then into the ocean. Although 
Baron de Lahontan may never have entered 
the western drainages or the Great Basin, 
the large western basin, Lahontan Basin, 
within the Great Basin was named after the 
Baron (Cline 1963). The leech is, thus, 
named after the basin in which it is found. 
The anatomical tradeoff in erpobdellid 
leeches having preatrial loops and having 
the gonopores separated by more annuli is 
that the preatrial loops and the cornua may 
become shorter. An evolutionary compen- 
sation would be for the cornua and the pre- 
atrial loop to form contortions. This may be 
what is happening in Erpobdella lahontana 
with the cornua and the preatrial loops both 
taking a ventral dip to their junction. 


Acknowledgments 


The authors wish to thank Teresa Puste- 
jovsky (U.S.ES., Eagle Lake, Susanville, 
California) for a wealth of ecological infor- 
mation and Cathy Sanderson, Wasatch His- 
to Consultants, Winnemucca, Nevada for 
making the histological slides of the repro- 
ductive system. 


Literature Cited 


Cline, G. C. 1963. Exploring the Great Basin.—Uni- 
versity of Oklahoma Press, Norman, Oklahoma, 
254 pp. 

Davies, R. W. 1991. Annelida: leeches, polychaetes, 
and acanthobdellids. Pp. 427-479 in J. H. 
Thorp and A. P. Covich, eds. Ecology and clas- 
sification of North American freshwater inver- 
tebrates. Academic Press, Inc., New York, 911 
Pp. 

Gester, G. C. 1962. The geological history of Eagle 
Lake, Lassen County, California.—Occasional 
Papers of California Academy of Sciences 34: 
1-29. 

Hershler, R. 1998. A systematic review of the hydro- 


VOLUME 113, NUMBER 1 


biid snails (Gastropoda: Rissoida) of Great Ba- 
sin, western United States. Part 1. Genus Pyr- 
gulopsis.—Veliger 41:1—132. 

Hovingh, P. 1997. Amphibians in the eastern Great Ba- 
sin (Nevada and Utah, USA): a geographical 
study with paleozoological models and conser- 
vation implications.—Herpetological Natural 
History 5(2):97-134. 

Hubbs, C. L., & R. R. Miller. 1948. The zoological 
evidence. The Great Basin, with emphasis on 
glacial and post-glacial times.—Bulletin Uni- 
versity of Utah Biological Series 38:18—166. 

Huntsinger, K. R., & P. E. Maslin. 1976. A limnolog- 
ical comparison of the three basins of Eagle 
Lake, California.—California Fish and Game 
62:232-245. 

Klemm, D. J. 1985. Freshwater leeches (Annelida: Hir- 
udinea). Pp. 70-194 in D. J. Klemm, ed., A 
guide to the freshwater Annelida (Polychaeta, 
naidid and tubificid Oligochaeta, and Hirudinea) 
of North America. Kendall/Hunt Publishing 
Company, Dubuque, Iowa, 198 pp. 

. 1990. Hirudinea. Pp. 398—415 in B. L. Peck- 

arsky, P. R. Fraissinet, M. A. Penton, & D. J. 

Conklin Jr. eds. Freshwater macroinvertebrates 

of northeastern North America. Cornell Univer- 

sity Press, Ithaca, New York, 442 pp. 

. 1995. Identification guide to the freshwater 
leeches (Annelida: Hirudinea) of Florida and 
other southern states.—Bureau of Surface Water 
Management, Florida Department of Environ- 
mental Protection, 2600 Blair Stone Road, Tal- 
lahassee, Florida 32399-2400. 

Lukin, E. I. 1976. Fauna of the USSR. Leeches, vol. 
1. Leeches of fresh and brackish water bod- 
ies._—Academy of Science, Zoological Institute 
of the Soviet Union. Nauka Publishers, Lenin- 
grad Branch. (467 pp., Translated from Russian 
by Literature Research Company for the U.S. 
EPA (in part) and the Canadian Translation Bu- 


16] 


reau for the Canadian Museum of Nature (in 
part). 

Mann, K. H. 1962. Leeches (Hirudinea) their structure, 
physiology, ecology, and embryology. Perga- 
mon Press, New York, 201 pp. 

Nesemann, H. 1995. On the morphology and taxono- 
my of the Asian leeches (Hirudinea: Erpobdel- 
lidae, Salifidae)—Acta Zoologica Academiae 
Hungaricae 41:165—182. 

Neubert, E., & H. Nesemann. 1995. Contribution to 
the knowledge of the genus Dina Blanchard, 
1892 (Hirudinea: Erpobdellidae).—Hydrobio- 
logia 315:89-94. 

Sawyer, R. T. 1972. North American freshwater leech- 
es, exclusive of the Piscicolidae with a key to 
all species.—Illinois Biological Monographs 
46:1-154. 

. 1986a. Leech Biology and Behavior. volume 

I. Anatomy, physiology, and behavior. Oxford 

University Press, Oxford, 417 pp. 

. 1986b. Leech Biology and Behavior. volume 
II. Feeding Biology, Ecology, and Systematics. 
Oxford University Press, pp. 419-793. 

Snyder, C. T., G. Hardman, & F. F Zdenek. 1964. Pleis- 
tocene Lakes in the Great Basin. U.S.G.S. Mis- 
cellaneous Geological Investigations Map _ I- 
416. 

Soos, A. 1963. Identification key to the species of the 
genus Dina R. Blanchard, 1892 (Emend. Mann 
1952, Proceedings Zoological Society, London 
122) (Hirudinea: Erpobdellidae).—Acta Uni- 
versitatis Szedgediensis, Szeged, Hungaricae 
IX:253-261. 

. 1966. Identification key to the leech (Hirudi- 

noidea) genera of the world, with a catalogue 

of the species. III. Family Erpobdellidae.—Acta 

Zoologica Academiae Scientiarum Hungaricae 

XII. (3—4):37 1-407. 

. 1968. Identification key to the species of the 

genus Erpobdella de Blainville, 1818 (Hirudi- 

noidea: Erpobdellidae).—Annales Historico- 

Naturales Musei Nationalis Hungaricae 60:141— 

145. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 
113(1):162—209. 2000. 


A cladistic analysis of Sciomyzidae Fallén (Diptera) 


Luciane Marinoni and Wayne N. Mathis 


(LM) Department of Zoology, Universidade Federal do Parana, Caixa Postal 19020, 81531-990, 


Curitiba, Parana, Brazil; 
(WNM) Department of Entomology, NHB 169, Smithsonian Institution, 
Washington, D.C. 20560, U.S.A. 


Abstract.—A preliminary cladistic analysis of adult characters is presented 
that illustrates the phylogenetic relationships among the genera of the family 
Sciomyzidae. The monophyly of Sciomyzidae is based primarily on larval char- 
acters: the habit of malacophagy and the presence of a serrate ventral arch that 
articulates with the lower margin of the mouth hooks. A reduction in the num- 
ber of spermathecae, from three to two, is also likely to be a synapomorphy 
for Sciomyzidae. The analysis was done using Hennig&6, and 37 morphological 
characters were arranged among 50 genera. After using successive weighting, 
six cladograms were produced, and from these a consensus cladogram was 
obtained. The subfamilies Salticellinae and Sciomyzinae are confirmed to be 
monophyletic, as are the tribes Sciomyzini and Tetanocerini. Eutrichomelina 
Steyskal, which has been placed in the tribe Sciomyzini, is transferred to the 
tribe Tetanocerini. The genus Antichaeta Haliday is confirmed to be in the tribe 


Tetanocerini. Illustrations of antenna and male terminalia are presented. 


Among families of Acalyptrate Diptera, 
the Sciomyzidae Fallén (1820), more com- 
monly known as marsh or snail-killing flies, 
are comparatively well studied, especially 
the biology of their immature stages (Berg 
& Knutson 1978, Ferrar 1987, Knutson 
1987, Rozkosny 1997). Sciomyzid larvae 
are primarily parasitoids or predators on 
aquatic or terrestrial mollusks. This feeding 
proclivity may be of considerable impor- 
tance to the biological control of certain 
parasitic, mostly tropical diseases, such as 
fascioliasis and schistosomiasis (Knutson 
1976). The trematodes causing both diseas- 
es parasitize many of the same aquatic mol- 
lusks, as intermediate hosts, that are also 
fed upon by sciomyzid larvae. 

Although the natural history and ecology 
and to a degree the descriptive taxonomy 
and cytology (Boyes et al. 1972) of the 
Sciomyzidae are relatively well known, no 
cladistic analysis at the generic level is 
available. As a step toward filling that void 


in our knowledge, this cladistic study was 
undertaken and is reported here. To provide 
perspective, we begin this report with a 
brief overview of the higher-level classifi- 
cation. 

In the first comprehensive treatment of 
Palearctic Sciomyzidae, Hendel (1900) di- 
vided the family into two subfamilies: Scio- 
myzinae and Tetanocerinae. Hendel char- 
acterized these subfamilies by the proepi- 
sternal seta (present in Sciomyzinae, absent 
in Tetanocerinae) and the frontal vitta (well 
developed and shiny in most Tetanocerinae, 
absent or reduced in most Sciomyzinae). 

Cresson (1920), in a study limited to the 
Nearctic fauna of Sciomyzidae, recognized 
Sciomyzinae, as characterized by Hendel, 
and described two additional subfamilies: 
Dryomyzinae and Euthycerinae. Cresson 
also proposed five tribes that were divided 
among two of the subfamilies as follows: 
(1) Sciomyzinae with Oidematopsini and 
Sciomyzini; and (2) Euthycerinae with 


VOLUME 113, NUMBER | 


Chaetomacerini, Euthycerini, and Sepedon- 
tini. 

Hendel (1924) published a key to the Pa- 
learctic genera and distinguished the genus 
Tetanura Fallén as a separate subfamily, 
Tetanurinae, based on the following com- 
bination of characters: arista subapical, 
forefemur bare, and ovipositor telescoped. 
In the same paper and within the subfamily 
Tetanocerinae, Hendel also proposed the 
tribe Salticellini for the genus Salticella Ro- 
bineau-Desvoidy. 

Over 30 years ago, Steyskal (1965) pro- 
posed a classification for the Sciomyzidae 
that has been the most extensive treatment 
for the family from the standpoint of higher 
categories. Steyskal’s classification, which 
has been adopted by most subsequent work- 
ers (Knutson et al. 1976, Barnes 1979, Roz- 
kosny & Elberg 1984, Knutson 1987, 
Barnes & Knutson 1989, McAlpine 1989), 
recognized five subfamilies: Huttonininae, 
Salticellinae, Helosciomyzinae, Phaeomyi- 
inae, and Sciomyzinae. The subfamily Scio- 
myzinae included two tribes, Sciomyzini 
and Tetanocerini. Although Steyskal’s char- 
acterization of Sciomyzidae did not identify 
apomorphic characters, he distinguished the 
family from related families by the follow- 
ing set of morphological characters: costal 
vein (C) without breaks; subcostal vein (Sc) 
complete, free from vein R,; vein A, com- 
plete; oral vibrissae absent; postvertical se- 
tae divergent to parallel; midfemur bearing 
a seta on the anterior surface; and at least 
one tibia with a preapical seta (Knutson 
1987, McAlpine 1989). The tribes Tetano- 
cerini and Sciomyzini are distinguished by 
the presence (Sciomyzini) or absence (Te- 
tanocerini) of a proepisternal seta. 

Griffiths (1972), who incorporated many 
characters of the male genitalia in his high- 
er level phylogenetic study, introduced the 
prefamily as a category between the super- 
family and the family categories. Using this 
classificatory structure, the prefamily Scio- 
myzoinea comprises the families Coelopi- 
dae, Phaeomyiidae, Dryomyzidae, Scio- 
myzidae, Helosciomyzidae, Ropalomeridae, 


163 


Sepsidae, Megamerinidae, and Cremifani- 
dae. Griffiths’ studies of Sciomyzidae were 
based on the following species: Pherbellia 
guadrata Steyskal, Pherbellia griseola 
(Fallén), Sciomyza simplex Fallén, Pterom- 
icra apicata (Loew), Elgiva sundewalli 
Kloet & Hincks, and Tetanocera robusta 
(Loew). The family Sciomyzidae, as char- 
acterized by Griffiths (1972), includes Sal- 
ticellinae + Sciomyzinae, and the family’s 
monophyly is based primarily on the ma- 
lacophagous habits of the larvae and the 
presence of a ventral arch in the cephalo- 
pharyngeal skeleton (Knutson et al. 1970, 
Barnes 1981, McAlpine 1989). The reduced 
number of spermathecae, from three to two, 
is likely to be another synapomorphy for 
the Sciomyzidae, although in Salticellinae, 
the number of spermathecae is four. We in- 
terpret the latter condition to be secondarily 
derived from two and to be an autapomor- 
phy for the subfamily Salticellinae. 

The subfamily Salticellinae has three 
species in two genera, the extant Salticella 
Robineau-Desvoidy and the fossil Prosal- 
ticella Hennig. The Sciomyzinae, which in- 
clude 505 recent species, have 57, mostly 
widespread genera. 

The purpose of this paper is to present a 
classification for the genera of Sciomyzidae 
sensu Griffiths that is based on a cladistic 
analysis of primarily morphological char- 
acters. Our analysis is intended more spe- 
cifically to test the hypotheses that the sub- 
families Salticellinae and Sciomyzinae and, 
within the latter subfamily, the tribes Scio- 
myzini and Tetanocerini are monophyletic. 


Material and Methods 


Fifty of the 57 genera belonging to the 
Salticellinae and Sciomyzinae were exam- 
ined and analyzed (Appendix 1). The seven 
genera not examined are: Ditaeniella Sack, 
Eulimnia Tonnoir & Malloch, Neodictya 
Elberg, Oligolimnia Mayer, Pseudomelina 
Malloch, Tetanoptera Verbeke, and Verbek- 
aria Knutson. We also studied a new genus 
and species from India, Steyskalina picta 


164 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Ghorpadé & Marinoni, that was recently 
described (Ghorpadé et al. 1999). Since it 
was virtually impossible to examine all spe- 
cies of each genus and suspecting that some 
genera may be polyphyletic (e.g., Pherbel- 
lia Robineau-Desvoidy with 81 species; Se- 
pedon Latreille with 75 species; and Tetan- 
ocera Duméril with 49 species) the analysis 
was done with the type species serving as 
exemplars of each genus. The type species 
are always linked to the appropriate generic 
name. 

Pelidnoptera, represented by P. fuscipen- 
nis (Meigen 1830), was used as the out- 
group. This genus, which is in the family 
Phaeomylidae (Griffiths 1972), was select- 
ed because it is morphologically similar and 
closely related, perhaps the sister group 
(Steyskal 1965, included it as a subfamily) 
of the Sciomyzidae. Its characters are di- 
rectly comparable to those of Sciomyzidae. 
The family Helosciomyzidae (Griffiths 
1972), represented by Helosciomyza aliena 
Malloch (1928); Huttonina abrupta Tonnoir 
& Malloch (1928); Huttonina furcata Ton- 
noir & Malloch (1928), and Huttonina scu- 
tellaris Tonnoir & Malloch (1928), was 
studied to further confirm the polarization 
of characters. 

A matrix with 37 morphological charac- 
ters of adults and one of the larval mor- 
phology was produced (Table 1). Autapo- 
morphies for particular genera were not in- 
cluded in the analysis. 

Multistate characters in the analysis were 
first treated as unordered (Carvalho 1989, 
Pape 1992, Marinoni & Carvalho 1993). 
The ordination of the characters was done 
in accordance with standard procedures for 
cladistic analysis (Wiley 1981). The polar- 
ization was done using the outgroup com- 
parison (Watrous & Wheeler 1981, Wiley 
1981, Brooks 1989). 

The analysis was facilitated with the 
computer program Hennig86, version 1.5 
(Farris 1988). To find cladograms supported 
by the most consistent characters, the fol- 
lowing command sequence was used: 
‘“‘mhennig”’ (mh), “‘branch and swapping” 


(*) and “‘successive weighting’ (xs w) 
(Carpenter 1988, Dietrich & McKamey 
1995). The option “‘nelsen’’ (ne) was used 
to construct a strict, consensus cladogram 
(Pape 1992, Marinoni & Carvalho 1993). 


Characters and Character States Used in 
the Analysis 


The characters used in the analysis are 
listed and discussed in the same sequence 
as they appear in the cladogram (Figs. 323— 
324). The letters A and P represent the rel- 
ative apomorphic (derived) and plesio- 
morphic (primitive) conditions respectively. 

la. Larval feeding behavior: P (0) feed- 
ing on Diplopoda; A (1) feeding as a par- 
asitoid or predator on terrestrial or aquatic 
Mollusca. 

This is one of just a few characters in the 
matrix that establishes the monophyly of 
Salticellinae + Sciomyzinae. The genus 
Pelidnoptera is a parasitoid on Diplopoda, 
and numerous larval characters distinguish 
it from the larvae of Sciomyzidae (Vala et 
al. 1990). Although malacophagy charac- 
terizes nearly all Sciomyzidae and is a syn- 
apomorphy for the family, there is at least 
one species, Sepedonella nana Verbeke, 
that feeds on oligochaetes (Vala et al 2000) 
as a secondary departure from the ground- 
plan of the more basal clades of the family. 

1b. Ventral arch in the larval cephalo- 
pharyngeal skeleton: P (0) absent; A (1) 
present. 

The serrated ventral arch articulates with 
the ventral margin of the larval mouth 
hooks. This is the only larval structural 
character that is an autapomorphy for Sal- 
ticellinae + Sciomyzinae and may be relat- 
ed with the malaphagous feeding behavior. 

2. Number of spermathecae: P (0) three; 
A (1) two; A (2) four. 

According to McAlpine (1989) the im- 
mediate ancestor of Muscomorpha had 
three sclerotized spermathecae. This is the 
basic number in the main sections of Mus- 
comorpha: Aschiza, Schizophora, Acalyp- 
tratae, and Calyptratae. In Pelidnoptera, the 


VOLUME 113, NUMBER 1 


Table 1.—Matrix of taxa and character states. 


Pelidnoptera 
Salticella 
Sciomyza 
Oidematops 
Atrichomelina 
Tetanura 
Colobaea 
Calliscia 
Parectinocera 
Pteromicra 
Pherbellia 
Eutrichomelina 
Ectinocera 
Renocera 
Antichaeta 
Chasmacryptum 
Shannonia 
Perilimnia 
Hoplodictya 
Dictya 
Hydromya 
Neolimnia 
Tetanoceroides 
Euthycerina 
Tetanocera 
Trypetolimnia 
Psacadina 
Steyskalina 
Dictyodes 
Ilione 
Pherbina 
Trypetoptera 
Limnia 
Poecilographa 
Pherbecta 
Protodictya 
Guatemalia 
Elgiva 

Hedria 
Dichetophora 
Coremacera 
Dictyacium 
Euthycera 
Ethiolimnia 
Teutoniomya 
Thecomyia 
Sepedoninus 
Sepedonella 
Sepedon 
Sepedomerus 
Sepedonea 


000 
123 


001 
120 
LLO 
110 


000 
456 


010 
010 
120 
020 
120 
iy oP 
110 
v6 
010 
010 
010 
OEE 
101 
Od. 
101 
10,2 
101 
FOP 
101 
101 
101 
DOs 
COL 
102 
G2 
102 
102 
nO 
Oe 
101 
00. 
101 
101 
101 
1.1 
TOL 
101 
nO 
FO 
£04 
101 
£02 
Jo 
101 
101 
102 
102 
102 
102 
LZ 
1 02 


000 
789 


a 
012 


000 
020 
101 
101 
100 
110 
£10 
000 
000 
000 
000 
000 
000 
000 
020 
000 
000 
000 
000 
000 
000 
000 
010 
000 
000 
010 
000 
000 
010 
010 
010 


ili BL 
345 


000 
000 
000 
000 
000 
000 
000 
100 
100 
000 
000 
001 
011 
011 
010 


111 
678 
000 
000 
000 
000 
000 
000 
000 
010 
000 
000 
000 
000 
000 
000 
100 
110 
112 
TAZ 
ai 2 
2 
12 
12 
142. 
142 
eee 
{ise 
MEY 
Es 
cag 
wi 
ele 
ce 
aia 
LAB 
142 
Ti2Z 
ly 
qb2 
ited 
dl: 
ie i 
101 
a ba 
003 
a Se 
pee 
isl s 
113 
TVS 
113 
13 


LZ 
901 


000 
000 
000 
000 
000 
000 
000 
000 
000 
000 
000 
000 
001 
000 
000 
000 
001 
001 


222 
234 
000 
000 
000 
000 
000 
000 
000 
000 
000 
000 
000 
000 
000 
000 
100 
000 
000 
000 
000 
000 
000 
000 
000 
000 
000 
000 
000 
000 
140 
100 
100 
100 
100 
100 
aaa ad 
ie eh 
000 
000 
000 
000 
000 
000 
000 
000 
000 
000 
000 
000 
000 
000 
000 


222 
567 
000 
000 
000 
000 
000 
010 
000 
000 
000 
000 
000 
000 
000 
000 
000 
000 
000 
000 
000 
000 
000 
000 
000 
000 
000 
000 
000 
000 
000 
000 
000 
000 
000 
000 


165 


a eo Crono eS SOO oS Of OO. © GC) Oso oS © Goro. 16 2 o O-O,. oo: ©) © CG, O20; Oo 2 Oo. OOO .© ©'@ 


166 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


number of spermathecae is also three, this 
number being considered the plesiomorphic 
condition for the Sciomyzidae. Only in Sal- 
ticella is the number of spermathecae four, 
which we interpret to be a secondary con- 
dition, probably being derived from two. 
Thus, four spermathecae are an autapomor- 
phy for the Salticellinae, and two sperma- 
thecae remain a synapomorphy for Scio- 
myzinae. 

3. Position of sixth left abdominal spi- 
racle of the male (Figs. 115-159): P (O) in 
membrane; A (1) in sclerotized tergite. 

Within the Sciomyzidae the abdominal 
spiracles of males may occur in the mem- 
brane or the sclerotized tergite. Primitively 
in Muscomorpha, the spiracles are in the 
membrane, which is considered the ple- 
siomorphic condition for the Sciomyzidae. 
The position of the sixth spiracle in the 
sclerotized portion of the tergite is a syna- 
pomorphy for the Tetanocerini. Males of 
Colobaea Zetterstedt have the spiracle in 
the sclerotized tergite, a condition that 
probably represents a secondary reversal. 

4. Sixth abdominal tergite of the male 
(Figs. 69, 74-76): P (O) present; A (1) ab- 
sent. 

Having all abdominal sclerites present is 
undoubtedly the plesiomorphic condition. 
According to McAlpine (1989) the reduc- 
tion of the sixth abdominal tergite in males 
is an apomorphy for the superfamily Scio- 
myzoidea. Griffiths (1972) considered this 
reduction to be a synapomorphy, confirm- 
ing the monophyly of Sciomyzidae. In 
Sciomyzidae, however, there is a complete 
absence of this tergite in most genera. Only 
in Salticella and in four genera of Scio- 
myzini, Oidematops Cresson, Parectino- 
cera Becker, Pherbellia, and Pteromicra 
Lioy, is there a sixth tergite, which, how- 
ever, is reduced. 

5. Subepandrial plate (Figs. 161-169, 
197): A (QO) absent; P (1) vestigial; A (2) 
well developed. 

The common, plesiomorphic condition, 
is the presence of a vestigial subepandrial 
plate found in Pelidnoptera, Salticella, Co- 


lobaea, Calliscia Steyskal, Parectinocera, 
Pteromicra, and Pherbellia. It is lacking in 
all genera of the Tetanocerini. In Sciomyza 
Fallén, Oidematops, and Atrichomelina 
Cresson, the subepandrial plate is well de- 
veloped and is a synapomorphic condition 
that characterizes these three genera. In Te- 
tanura (tribe Sciomyzini), the plate is also 
absent, a condition we consider to be ho- 
moplastic. 

6. Anterior surstylus (Figs. 161—203): P 
(O) well developed; A (1) vestigial; A (2) 
absent. 

The transformation series for this char- 
acter is linear: 0 ~ 1 ~ 2. Pelidnoptera 
has two pairs of well-developed, paired sur- 
Styli, and thus the presence of an anterior 
well-developed surstylus is the plesiom- 
orphic condition for this character. Within 
the Sciomyzidae, the tribe Sciomyzini also 
has an anterior, well-developed surstylus. 
The genera Tefanura (tribe Sciomyzini) and 
Elgiva Meigen; the assemblage formed by 
the genera Euthycerina Malloch, Tetano- 
cera, Trypetolimnia Mayer, Psacadina En- 
derlein, and Steyskalina Knutson; and the 
group of Thecomyia Perty, Sepedoninus 
Verbeke, Sepedonella Verbeke, Sepedon, 
Sepedomerus Steyskal + Sepedonea Steys- 
kal; lack an anterior surstylus, having only 
the posterior one. The remaining genera of 
Tetanocerini have a vestigial, anterior sur- 
stylus. 

7. Shape of aedeagus (Figs. 61—63, 66— 
67, 204-322): P (O) asymmetrical; A (1) 
symmetrical. 

Asymmetry in the postabdomen of male 
Acalyptratae may involve internal struc- 
tures, including the aedeagus. Within gen- 
era of Sciomyzidae, the aedeagus demon- 
strates great variation in shape and sym- 
metry. Pelidnoptera has an asymmetrical 
aedeagus, a condition that is considered to 
be plesiomorphic. In most Tetanocerini, ex- 
cept for Antichaeta, the aedeagus is a com- 
plex structure with several completely 
asymmetrical sclerites. In the Sciomyzini 
and in the group of Sepedoninus, Sepedo- 


VOLUME 113, NUMBER 1 


nella, Sepedon, Sepedomerus + Sepedonea, 
the aedeagus is completely symmetrical. 

8. Attachment of gonopod (Figs. 204— 
322): P (O) fused to the hypandrium; A (1) 
free. 

McAlpine (1989) postulated the fusion of 
the gonopod to the hypandrium as a ple- 
siomorphic condition for the Acalyptratae. 
All groups examined that are closely related 
to the Sciomyzidae have the gonopod fused 
to the hypandrium. The presence of a gon- 
opod that is well developed and free is a 
condition that supports the monophyly of 
the tribe Sciomyzini. In Tetanoceroides 
Malloch, a free gonopod is apparently ho- 
moplastic. 

9. Paramere (Figs. 204—322): A (QO) not 
digitiform, elongate and well developed; P 
(1) digitiform and well developed; A (2) ab- 
sent. 

Pelidnoptera has a digitiform, well-de- 
veloped paramere that is considered to be 
plesiomorphic. In Salticella, the paramere is 
elongate and well developed, as in the Scio- 
myzini and in the genera Antichaeta, Psa- 
cadina, Neolimnia, Ethiolimnia, and Steys- 
kalina. Two genera, Sepedonea and Sepe- 
donella, jack a paramere, a condition that is 
considered homoplastic. 

10. Shape of aedeagal apex (Figs. 204— 
213): P (O) variously shaped, but not flat- 
tened; A (1) flattened. 

The presence of an aedeagus with a flat- 
tened apex is a synapomorphy for the group 
of Sciomyza, Oidematops, Atrichomelina, 
Tetanura, and Colobaea within the Scio- 
myzini. 

11. Basiphallus (Figs. 203-322): P (0) 
well developed; A (1) present but poorly 
developed; A (2) absent. 

This character has the greatest number of 
steps in the matrix and analysis, indicating 
a large number of homoplasies and rever- 
sals. The taxa that have the intermediate 
apomorphic condition, the basiphallus 
poorly developed, are Colobaea, Tetanura, 
Trypetolimnia, Tetanoceroides, Dictyodes 
Malloch, Jlione Verbeke, Pherbina Robi- 
neau-Desvoidy, Trypetoptera Hendel, Gua- 


167 


temalia Steyskal, Hedria Steyskal, Diche- 
tophora Rondani, Poecilographa Melander, 
Protodictya Malloch + Pherbecta Steyskal 
(the later two genera are sister groups, as 
indicated by the “‘+”’’ connection). The gen- 
era Salticella, Antichaeta, Sepedonella, Se- 
pedon, Sepedomerus, and Sepedonea \ack a 
basiphallus. The remaining genera have a 
well-developed basiphallus. 

12. Setal investment of foretibia: P (0) 
without preapical setae; A (1) bearing a pair 
of preapical setae. 

The presence of a pair of preapical setae 
on the foretibia is a synapomorphy for Oi- 
dematops and Sciomyza. 

13. Vestiture of the aedeagus (Figs. 214-— 
217): P (O) without scalelike structures; A 
(1) with scalelike structures. 

In Calliscia and Parectinocera, the ae- 
deagus is covered with small scalelike 
structures, a synapomorphy for these gen- 
era. 

14. Proepisternal seta: P (0) present; A 
(1) absent. 

Steyskal (1965) used the presence of the 
proepisternal seta to distinguish the tribe 
Sciomyzini from the Tetanocerini. Most 
genera of Tetanocerini lack this seta, al- 
though Shannonia Malloch and Perilimnia 
Becker possess one. For the latter genera 
this condition is considered to be homo- 
plastic. 

15. Male terminalia (Figs. 57-59, 64-65, 
68-160): P (O) with the 6th, 7th, and 8th 
sternites separated, the 6th and 7th asym- 
metrical; A (1) with the synsternite 7+8 
and 6th sternite separated and asymmetri- 
cal; A (2) with the synsternite 6+7+8 sym- 
metrical. 

According to Steyskal (1957) the asym- 
metry of the male Acalyptratae postabdo- 
men, including segments beyond the fifth, 
is due to three basic movements within the 
pupa: circumversion, reflection, and “‘stro- 
phe.”’ The symmetry may occur as a result 
of the obliteration of these movements. 
Asymmetry in Sciomyzidae is restricted to 
the sixth, seventh, and eighth segments. 
There is a modification gradient of these 


168 


segments, changing from completely asym- 
metrical to completely symmetrical, asso- 
ciated to the fusion of the segments men- 
tioned above. 

16. Placement of sixth right abdominal 
spiracle of the male (Figs. 68-114): P (0) 
in membrane; A (1) in sclerotized tergite. 

Sciomyzini and Eutrichomelina, Reno- 
cera Hendel, and Ectinocera Zetterstedt 
have the plesiomorphic condition of this 
character, the spiracle is in the membrane. 
From Antichaeta to Sepedonea, the spiracle 
is in the tergite. In Ethiolimnia, there is a 
reversal to the plesiomorphic condition. 

17. Placement of seventh right abdominal 
spiracle of the male (Figs. 68—114): P (O) 
in membrane; A (1) in sclerotized tergite. 

The plesiomorphic condition occurs in 
most Sciomyzini except for Calliscia. From 
Chasmacryptum Becker to Sepedonea most 
genera have the spiracle in the tergite, ex- 
cept for Dictyacium Steyskal and Ethiol- 
imnia. 

18. Shape of head: P (O) as in Fig. 52a; 
A()-asin Figs s2bie’ @)as im Biesis2c; 
A(8)-as inePiss 52d. 

The head, from the ancestor to Chasma- 
cryptum, has the shape shown in Fig. 52a. 
Beyond the group of Shannonia + Perilim- 
nia the intermediate apomorphic condition, 
represented by Fig. 52c, is present. From 
this state the other two apomorphic states 
evolved. The pattern of head shape, as in 
Fig. 52b, is present in Trypetolimnia and 
Limnia Robineau-Desvoidy as homoplasies 
and in Hedria, Dichetophora, Coremacera 
Rondani, Dictyacium, and Euthycera La- 
treille. The shape, as in Fig. 52d, defines 
the group of Ethiolimnia, Thecomyia, Se- 
pedoninus, Sepedonella, Sepedon, Sepe- 
domerus, and Sepedonea. Because Teuto- 
niomyia Hennig has a very different head, 
compared to other genera, this character 
was coded as missing in the matrix (Table 
i): 

19. Proportion between the length of the 
pedicel and first flagellomere (Figs. 1—51): 
P (QO) pedicel approximately half of first fla- 
gellomere; A (1) pedicel subequal to the 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


length of the first flagellomere; A (2) ped- 
icel approximately twice the length of the 
first flagellomere. 

The plesiomorphic condition is present 
from Sciomyzini until Shannonia + Peri- 
limnia. From this group to Ethiolimnia the 
intermediate apomorphic condition (1) is 
present. The apomorphic condition (2) is 
present in the following genera: Teutoni- 
omyia, Thecomyia, Sepedoninus, Sepedo- 
nella, Sepedon, Sepedomerus, and Sepedo- 
nea. 

20. Condition of sternites of female ter- 
minalia (Figs. 53-56): P (O) with sternites 
6th, 7th, and 8th separate; A (1) with 6th 
separate and 7th and 8th fused; A (2) 6th, 
7th, and 8th incompletely fused; A (3) 6th, 
7th, and 8th completely fused. 

The transformation series was_ linear. 
Two lineages arose from state 1. One of 
these goes through the third state and the 
second goes through the fourth state. The 
evolution occurred from separated sternites 
passing through steps of fusion to the com- 
plete fusion of 6th, 7th, and 8th. All Scio- 
myzini and the Tetanocerine genera until 
Hoplodictya Cresson + Dictya Meigen 
have sternites 6th, 7th, and 8th separate. 
From the polytomy of Hydromya Robi- 
neau-Desvoidy to the ancestor of Ethiol- 
imnia the 7th and 8th sternites are fused. 
Protodictya + Pherbecta and Poecilogra- 
pha have the three sternites completely 
fused as Sepedoninus, Sepedon, Sepedom- 
erus, and Sepedonea. In Sepedonella, a ge- 
nus belonging to this group, the three ster- 
nites are separate. The monophyletic group 
formed by Coremacera, Dictyacium, and 
Euthycera, and the genera Limnia, Teuto- 
niomyia, and Thecomyia have the sternites 
6, 7, and 8 almost fused. Lines separating 
these sternites are perceptible. 

21. Aedeagus with lobed apex (Figs. 
222-224, 230-238): P (O) absent; A (1) 
present. 

Ectinocera, Hoplodictya, and Perilimnia 
+ Shannonia have the aedeagus with the 
distal end lobed. This character apparently 
appeared independently in these genera. In 


VOLUME 113, NUMBER | 


the other genera, the distal end of the ae- 
deagus has several different shapes. 

22. Subalar setae: P (0) absent; A (1) 
present. 

The subalar setae are present in Dictyo- 
des, Ilione, Pherbina, Trypetoptera, Limnia, 
Poecilographa, Pherbecta + Protodictya. 
The apomorphic state apparently evolved 
independently in Antichaeta. 

23. Aedeagus convoluted as in Figs. 
263-265, 281-286: P (0) absent; A (1) 
present. 

A convoluted aedeagus occurs in Proto- 
dictya, Pherbecta, and Dictyodes. For Pro- 
todictya and Pherbecta this character de- 
fines the common ancestor. 

24. Fourth abdominal sternite of the male 
fused to fifth, forming a medioapical pro- 
cess: P (OQ) absent; A (1) present. 

The presence of a well-developed, me- 
dioapical process in the fifth abdominal 
sternite of the male as a complex structure 
is a synapomorphy for the species of Pro- 
todictya (Marinoni & Knutson 1992, Mar- 
inoni & Carvalho 1993). In Pherbecta, 
however, there is, at the same position, a 
process that is less well developed and 
which may be homologous to that of Pro- 
todictya. 

25. Lunula: P (0) not exposed; A (1) ex- 
posed; A (2) greatly exposed. 

In the outgroup, Pelidnoptera, genera of 
Sciomyzini, and genera of Tetanocerini 
from Eutrichomelina to Protodictya, the lu- 
nula is not exposed. From Guatemalia to 
Sepedonella, the intermediate state (1) is 
present. In Ethiolimnia, there is a reversal 
to the plesiomorphic state. In the apical 
genera, Sepedon, Sepedomerus, and Sepe- 
donea, the lunula is greatly exposed, which 
is interpreted as an even more derived con- 
dition from the intermediate state. In Teu- 
toniomyia, which also has a greatly exposed 
lunula, this character state is homoplastic. 

26. Presutural supra-alar setae: P (0) 
present; A (1) absent. 

The plesiomorphic condition of this char- 
acter is the presence of presutural supra-alar 
setae. The absence of these setae is a syn- 


169 


apomorphy for the group Elgiva, Hedria, 
Dichetophora, Ethiolimnia, Teutoniomyia, 
Thecomyia, Sepedoninus, Sepedonella, Se- 
pedon, Sepedomerus, and Sepedonea. In 
Coremacera, Dictyacium, and Euthycera, 
there is a reversal to the plesiomorphic 
state. Tetanura has this seta, a condition 
that is considered homoplastic. 

27. Seventh and 8th abdominal tergites 
of the female: P (0) separate; A (1) fused. 

The fusion of the 7th and the 8th abdom- 
inal tergites of the female is a character 
state that links the genera Dichetophora, 
Coremacera, Dictyacium, and Euthycera. 
In Elgiva and Sepedonella, the fusion is ho- 
moplastic. 

28. Prominent eyes: (0) absent; (1) pres- 
ent. 

Ethiolimnia, Thecomyia, Sepedoninus, 
Sepedonella, Sepedon, Sepedomerus, and 
Sepedonea have prominent eyes, a syna- 
pomorphy for these genera. As Ethiolimnia 
is in a polytomy with Teutoniomyia, prom- 
inent eyes are a confirmed synapomorphy 
for Thecomyia, Sepedoninus, Sepedonella, 
Sepedon, Sepedomerus, and Sepedonea and 
only perhaps for Ethiolimnia. 

29. Number of postalar setae: P (0) 2; A 
(1) O-1. 

The presence of two postalar setae is the 
plesiomorphic condition. In Ectinocera and 
in the genera beyond Ethiolimnia, the num- 
ber of postalar setae is reduced to one or 
none. 

30. Ocellar setae: P (QO) present; A (1) 
absent. 

The presence of a pair of strong ocellar 
setae has been considered a plesiomorphic 
condition for the Muscomorpha (McAlpine 
1989). In Pelidnoptera, these setae are pres- 
ent, as they are in most Sciomyzidae, in- 
cluding Salticella. These setae are absent in 
Hedria and in the group of Thecomyia, Se- 
pedoninus, Sepedonella, Sepedomerus + 
Sepedonea. 

31. Postpronotal setae: P (0) present; A 
(1) absent. 

Most Sciomyzidae have postpronotal se- 
tae. The absence of these setae character- 


170 


izes the group of genera beyond Thecomyia. 
In Dichetophora and Tetanura, the absence 
of postpronotal setae is considered to be ho- 
moplastic. 

32. Number of ‘scutellar setae?“ PP")? 2 
pairs; A (1) 1 pair. 

Primitively the family Sciomyzidae has 
two pairs of scutellar setae. One pair of scu- 
tellar setae occurs in Dichetophora and the 
group of genera beyond Thecomyia. 

33. Number of notopleural setae: P (0) 2; 
A (ty 

Two notopleural setae are present in most 
Sciomyzidae and in Pelidnoptera. Only the 
genera Thecomyia, Sepedoninus, and Sepe- 
donella have a single notopleural seta. A 
reversal of this characters to the plesio- 
morphic state of two notopleural setae oc- 
curs in the genera Sepedon, Sepedomerus, 
and Sepedonea. 

34. Hindtibia arch-shaped: P (0) absent; 
A (1) present. 

The genera Teutoniomyia, Sepedoninus, 
Sepedonella, Sepedon, Sepedomerus, and 
Sepedonea have an arch-shaped hindtibia. 
Only Thecomyia does not have the hindtibia 
arch-shaped. 

35. Head with sutures: P (0) distinct; A 
(1) indistinct. 

The head sutures of Thecomyia, Sepe- 
doninus, Sepedonella, Sepedon, Sepedome- 
rus, and Sepedonea are indistinct. This con- 
dition is considered a synapomorphy for 
this group. 

36. Epandrium (Figs. 68—160): P (0) free; 
A (1) fused to the 8th sternite. 

The fusion of the epandrium to the 8th 
sternite is a synapomorphy for the genera 
Sepedonella, Sepedon, Sepedomerus, and 
Sepedonea. 

37. Hindtibia with a spinelike projection: 
P (O) absent; A (1) present. 

In most Sciomyzidae, there is no projec- 
tion from the hindtibia. Sepedonea and Se- 
pedomerus have a hindtibia that bears a 
spinelike projection, which is a synapomor- 
phy that links these two genera. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Discussion 


Relationships.—The monophyly of the 
Sciomyzidae is defined by a _ behavioral 
character, the larvae are predators or para- 
sitoids on aquatic or terrestrial Mollusca 
(Berg 1953, Knutson et al. 1970, Griffiths 
1972, Barnes 1979) and the presence of a 
serrate ventral arch below the mouth hooks 
in the larvae (Griffiths 1972). The serrate 
ventral arch could be closely correlated 
with the malacophagous feeding behavior. 

The genus Salticella, representing the 
Salticellinae, remains at the base of the 
cladogram, being distinguished from the 
subfamily Sciomyzinae and its monophyly 
confirmed by the following three autapo- 
morphies: four spermathecae; cell M nar- 
rowed apically (this character was not used 
in the analysis because of its variability 
among genera of Sciomyzidae); and three 
pairs of surstyli. Knutson et al. (1970), 
working on the biosystematics of Salticella 
fasciata (Meigen), concluded that this ge- 
nus is more closely related to Sciomyzini 
than to the Tetanocerini in morphological 
and biological aspects. This relationship is 
corroborated in the present analysis. 

Among genera of the subfamily Scio- 
myzinae, two main monophyletic clades 
were formed, the first including Pherbellia, 
Pteromicra, Calliscia, Parectinocera, Te- 
tanura, Colobaea, Atrichomelina, Oidema- 
tops, and Sciomyza; and the second includ- 
ing the 40 remaining genera. The two 
clades are concordant with the tribes Scio- 
myzini and Tetanocerini of Steyskal (1965). 
The genus Eutrichomelina, which tradition- 
ally has been placed in the Sciomyzini, is 
shown to be allied to the Tetanocerini in 
this analysis. 

Behavioral aspects of the larvae have 
been discussed as possible synapomor- 
phies for the tribes (Knutson & Lyneborg 
1965, Abercrombie 1970). Toward one end 
of the scale, where the Sciomyzini are 
placed, the species that have been reared 
are parasitoids on stranded aquatic or ter- 
restrial mollusks (Colobaea (Knutson 


VOLUME 113, NUMBER 1 


0,5 mm 


10 


Figs. 1-11. 


0,5 mm 
2 ae es ? 


7a 


11 


Right antenna, right lateral view: 1, Pelidnoptera; 2, Salticella; 3, Sciomyza; 4, Oidematops; 5, 


Atrichomelina; 6, Tetanura;,; 7, Colobaea; 8, Calliscia; 9, Parectinocera, 10, Pteromicra; 11, Pherbellia. Fig. 


11, scale = 1.0 mm; remaining Figs. scale = 0.5 mm. 


1973); Pherbellia (Bratt et al. 1969); Pter- 
omicra (Rozkosny & Knutson 1970); Scio- 
myza (Foote 1959); Tetanura (Knutson 
1970)). Toward the other end, most Tetan- 
ocerini are predaceous on aquatic or ter- 
restrial mollusks (Coremacera (Knutson 
1973); Dichetophora (Vala et al. 1987); 
Dictya (Valley & Berg 1977); Dictyodes 
(Abercrombie 1970, Abercrombie & Berg 
1978); Elgiva (Knutson & Berg 1964a, 
Orth & Knutson 1987); Euthycera (Vala et 


al. 1983, Vala & Caillet 1985); Hoplodic- 
tya (Neff & Berg 1962); Hydromya (Knut- 
son & Berg 1963); /lione (Knutson & Berg 
1964b); Limnia (Vala & Knutson 1990); 
Perilimnia (Kaczynski et al. 1969); Pher- 
becta (Knutson 1972); Pherbina (Knutson 
et al. 1975, Vala & Gasc 1990); Protodic- 
tya (Abercrombie 1970, Neff & Berg 
1961); Psacadina (Knutson et al. 1975); 
Renocera (Foote & Knutson 1970); Sepe- 
don (Neff & Berg 1966, Knutson et al. 


172 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Ee 
SM 


eG a ai 


12 13 


0,5 mm 


LEER OR 


GA 


24 


0,5 mm 


28 


Figs. 12-29. Right antenna, right lateral view: 12, Eutrichomelina; 13, Ectinocera; 14, Renocera; 15, An- 
tichaeta;,; 16, Chasmacryptum; 17, Shannonia; 18, Perilimnia; 19, Hoplodictya,; 20, Dictya; 21, Hydromya; 22, 
Neolimnia; 23, Tetanoceroides; 24, Euthycerina; 25, Tetanocera; 26, Trypetolimnia; 27, Psacadina; 28, Steys- 
kalina; 29, Dictyodes. Figs. 18 and 23 with scale 1.0 mm; the remaining Figs. 0.5 mm. 


VOLUME 113, NUMBER 1 173 


0,5 mm 


1mm 


1mm 


wu G‘O 


Figs. 30-43. Right antenna, right lateral view: 30, /lione; 31, Pherbina; 32, Trypetoptera; 33, Limnia; 34, 
Poecilographa; 35, Pherbecta; 36, Protodictya; 37, Guatemalia; 38, Elgiva; 39, Hedria; 40, Dichetophora; 41, 
Coremacera; 42, Dictyacium; 43, Euthycera. Figs. 30-35 and 40 with scale 0.5 mm; Figs. 37-39 and 41—43 
with scale 1.0 mm. 


1967); Sepedonea (Abercrombie 1970, 1970); Thecomyia (Abercrombie & Berg 
Knutson & Valley 1978); Shannonia (Kac- 1975); Trypetoptera (Vala 1986)). 

zynski et al. 1969); Tetanocera (Knutson A few species of Sciomyzini and Tetan- 
et al. 1965); Tetanoceroides (Abercrombie  ocerini, however, have an intermediate be- 


174 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Figs. 44-51. 


Right antenna, right lateral view: 44, Ethiolimnia; 45, Teutoniomyia; 46, Thecomyia; 47, Se- 


pedoninus,; 48, Sepedonella; 49, Sepedon; 50, Sepedomerus; 51, Sepedonea. Fig. 45 with scale 0.5 mm; the 


remaining Figs. with scale 1.0 mm. 


havior between these two behaviors, 1.e., 
sometimes they have parasitoid trends, 
sometimes predaceous. Examples of these 
kinds of behavior are evident in the species 
Atrichomelina pubera Loew and Antichaeta 
analis Zetterstedt. Atrichomelina pubera 
belongs to the Sciomyzini. Depending on 
circumstances this species has a predatory, 
parasitoid or even saprophagous behavior 
(Berg et al. 1959, Foote et al. 1960). Sap- 
rophagy may be the primitive condition, 
similar to that found in Salticella (Knutson 
et al. 1970) and the ancestor of Sciomyzi- 
dae (Abercrombie 1970). Antichaeta analis 
has some parasitoid characteristics, such as 
host specificity and oviposition on the host, 
but its behavior is predatory with the larvae 
feeding on snail eggs (Fisher & Orth 1964, 
Knutson 1966, Knutson & Abercrombie 
LOFT): 

More studies related to the evolution of 
the larval behavior are necessary to confirm 
that the larval habit is responsible for re- 
lationships among the genera within the 
family. 


Most genera in the analysis remained in 
unresolved polytomies due to the great 
number of homoplasies and low number of 
synapomorphies. 

Within the Sciomyzini, the relationships 
of Pteromicra and Pherbellia are not well 
resolved. RozkosSny & Knutson (1970), 
studying the biology and immature stages 
of Pteromicra, discovered that this genus is 
closely related to Pherbellia. However, 
even in our analysis, where these two gen- 
era are in close proximity on the cladogram, 
there is no adult character to corroborate 
this relationship. 

Knutson et al. (1990) considered Colo- 
baea, Pherbellia, and Pteromicra to be re- 
lated but did not elaborate any synapomor- 
phies to support their grouping. In the same 
paper they also suggest that Pherbellia is 
paraphyletic. 

Parectinocera and Calliscia appear in the 
analysis as having the same ancestor. These 
two genera are the only two within the 
Sciomyzini with Neotropical distributions. 

The genera Tetanura, Colobaea, Atricho- 


VOLUME 113, NUMBER 1 


52 


0,5 mm 


©» 


Figs. 52—56. 


syn. 6+7+8 


Ce) 
5 


6 


52. Shape of head: (a), State 0; (b), state 1; (c) state 2; (d) state 3. 53-56. Female terminalia, 


ventral view: 53, Perilimnia; 54, Elgiva; 55, Coremacera; 56, Protodictya. Sternite 6 (st. 6); sternite 7 (st. 7); 


sternite 8 (st. 8); synsternite 6+7+8 (syn. 6+7+8). 


melina, Sciomyza, and Oidematops form a 
clade established by characters 4 and 10 
(Figs. 323-324). In this group Atrichome- 
lina, Sciomyza + Oidematops have their re- 
lationships well defined. The synapomorph- 
ic character that links the three genera is the 


presence of a well-developed male sub- 
epandrial plate (Character 5). The other two 
genera remain in polytomy. 

In the Tetanocerini, Eutrichomelina, Ec- 
tinocera, Renocera, Antichaeta, and Chas- 
macryptum are at the base of the clade. Ec- 


176 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


1mm 


bas. 


( | 0,5 mm 


62 63 


Figs. 57-63. Male terminalia. Pelidnoptera. Aedeagal apodeme (aed. ap.); basiphallus (bas.); cerci (cer.); 
aedeagal complex (aed. c.); epandrium (ep.); sternite 6 (st. 6); sternite 7 (st. 7); sternite 8 (st. 8); hypandrium 
(hip.); paramere (par.); subepandrial plate (sub. pl.); anterior surstylus (a. s.); posterior surstylus (p. s.). Figs. 
57—60 with scale 1.0 mm; Figs. 61—63 with scale 0.5 mm. 


tinocera and Renocera remain in polytomy. already discussed by Fisher & Orth (1964) 
Steyskal (1959), using intuitive analysis, & Knutson (1966). Adult characters suggest 
considered Renocera, Chasmacryptum, and that this genus belongs to the Tetanocerini, 
Tetanocera to be morphologically related. | while larval characters indicate its position 

The uncertain position of Antichaeta was _ to be within the Sciomyzini. Boyes et al. 


VOLUME 113, NUMBER 1 177 
st. 8 
/ Low cer. 
ep. 
oe 
st. 7 
= a.s 
st. 6 a } 
p.s 
\ s.m 
cs 
p.s. 65 
64 
aed. 
Figs. 64-67. Male terminalia. Salticella. Aedeagal apodeme (aed. ap.); cerci (cer.); aedeagal complex (aed. 


c.); epandrium (ep.); sternite 6 (st. 6); sternite 7 (st. 7); sternite 8 (st. 8); hypandrium (hip.); paramere (par.); 
anterior surstylus (a. s.); medial surstylus (s. m.); posterior surstylus (p. s.). Figs. 64—65 with scale 1.0 mm; 


Figs. 66—67 with scale 0.5 mm. 


(1969), studying the karyotype of Antichae- 
ta melanosoma Melander, observed its sim- 
ilarity with those of the Tetanocerini gen- 
era. In the present analysis, Antichaeta is 
placed in the Tetanocerini group of genera 
near Chasmacryptum. 

After Chasmacryptum, the clade of Shan- 
nonia + Perilimnia appears as the sister 
group of the other genera. Kaczynski et al. 


(1969) studied the biosystematics of the im- 
mature stages of these two genera and ob- 
served morphological and behavioral affin- 
ities between them. As observed by these 
authors, the main similarities between the 
larvae of Shannonia and Perilimnia are in 
the cephalopharyngeal skeleton. They also 
suggested the position of both genera with- 
in the Tetanocerini. 


178 


Figs. 68-76. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Male terminalia, ventral view: 68, Sciomyza; 69, Oidematops; 70, Atrichomelina; 71, Tetanura; 


72, Colobaea;, 73, Calliscia; 74, Parectinocera; 75, Pteromicra; 76, Pherbellia. Cerci (cer.); epandrium (ep.); 
sternite 6 (st. 6); sternite 7 (st. 7); sternite 8 (st. 8); anterior surstylus (a. s.); posterior surstylus (p. s.); tergite 6 
(terg. 6). Fig. 68 with scale 1.0 mm; the remaining Figs. with scale 0.5 mm. 


Hoplodictya + Dictya form the next 
clade. Boyes et al. (1969) compared its kar- 
yotypes and recognized the proximity of 
these two genera. 

Hydromya, Neolimnia, and Tetanocero- 
ides remain as a polytomy with three other 
groups formed. The components of one of 
these groups are: Euthycerina, Tetanocera, 
Trypetolimnia, Psacadina, and Steyskalina. 
The other group is formed by Dictyodes, 
Ilione, Pherbina, Trypetoptera, Limnia, 


Poecilographa, Pherbecta + Protodictya. 
Within the latter group the genera Limnia, 
Poecilographa, Pherbecta + Protodictya 
are monophyletic. In the third group, which 
comprises the last fifteen genera, we rec- 
ognize two subgroups. One is formed by 
Hedria, Dichetophora, Coremacera, Dic- 
tyacium, and Euthycera, and the other by 
Ethiolimnia, Teutoniomyia, Thecomyia, Se- 
pedoninus, Sepedonella, and Sepedomerus 
+ Sepedonea. The monophyly of the first 


VOLUME 113, NUMBER I 179 


0,5 mm 


85 


1mm 


Figs. 77-87. Male terminalia, ventral view: 77, Ectinocera; 78, Renocera; 79, Antichaeta; 80, Chasmacryp- 
tum; 81, Shannonia; 82, Perilimnia; 83, Hoplodictya; 84, Dictya; 85, Hydromya; 86, Neolimnia; 87, Euthycerina. 
Cerci (cer.); epandrium (ep.); sternite 6 (st. 6); sternite 7 (st. 7); sternite 8 (st. 8); synsternite 7+8 (syn. 7+8); 
synsternite 6+7+8 (syn. 6+7+8); anterior surstylus (a. s.); posterior surstylus (p. s.). Fig. 86 with scale 1.0 mm; 


the remaining Figs. with scale 0.5 mm. 


180 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


89 90 
syn. 6+7+8 
(So 
QZG 93 


0,5 mm 


99 


0,5 mm 


Figs. 88-99. Male terminalia, ventral view: 88, Tetanocera; 89, Trypetolimnia; 90, Psacadina; 91, Steys- 
kalina; 92, Dictyodes; 93, Ilione; 94, Pherbina; 95, Trypetoptera; 96, Limnia; 97, Poecilographa; 98, Pherbecta; 
99, Protodictya. Cerci (cer.); epandrium (ep.); sternite 6 (st. 6); synsternite 7+8 (syn. 7+8); synsternite 6+7+8 
(syn. 6+7+8); anterior surstylus (a. s.); posterior surstylus (p. s.). Figs. 90 and 98 with scale 0.5 mm; the 
remaining Figs. with scale 1.0 mm. 


VOLUME 113, NUMBER 1 181 


0,5 mm 1mm 
ep. 


cer. Oy (s 


103-7 >< 
<a 102 ee 


syn.7+8 


110 
109 


pe 6+7+8 es 
111 112 ain 113 AQ 
0,5 mm 


Figs. 100-114. Male terminalia, ventral view: 100-101, Guatemalia; 102, Elgiva; 103, Hedria; 104, Diche- 
tophora; 105, Coremacera; 106, Dictyacium; 107, Euthycera; 108, Ethiolimnia; 109, Thecomyia; 110, Sepedon- 
inus; 111, Sepedonella; 112, Sepedon; 113, Sepedomerus; 114, Sepedonea. Cerci (cer.); epandrium (ep.); sternite 
6 (st. 6); synsternite 7+8 (syn. 7+8); synsternite 6+7+8 (syn. 6+7+8); anterior surstylus (a. s.); posterior 
surstylus (p. s.). Figs. 100, 101, 104-106, 111—114 with scale 0.5 mm; the remaining Figs. with scale 1.0 mm. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


120 


0,5 mm 


123 


cer. 


Figs. 115-123. Male terminalia, left lateral view: 115, Sciomyza; 116, Oidematops; 117, Atrichomelina; 118, 
Tetanura; 119, Colobaea; 120, Calliscia; 121, Parectinocera; 122, Pteromicra; 123, Pherbellia. Cerci (cer.); 
epandrium (ep.); sternite 6 (st. 6); sternite 7 (st. 7); sternite 8 (st. 8); anterior surstylus (a. s.); posterior surstylus 
(p. s.); tergite 6 (terg. 6). Fig. 115 with scale 1.0 mm; the remaining Figs. with scale 0.5 mm. 


VOLUME 113, NUMBER 1 183 


124 


0,5 mm 


Z 134 135 


Figs. 124-135. Male terminalia, left lateral view: 124, Ectinocera; 125, Renocera; 126, Antichaeta; 127, 
Chasmacryptum; 128, Shannonia; 129, Perilimnia; 130, Hoplodictya; 131, Dictya; 132, Hydromya; 133, Neo- 
limnia; 134, Euthycerina; 135, Tetanocera. Cerci (cer.); epandrium (ep.); sternite 6 (st. 6); synsternite 7+8 (syn. 
7+8); synsternite 6+7+8 (syn. 6+7+8); anterior surstylus (a. s.); posterior surstylus (p. s.). Figs. 133 with scale 
1.0 mm; the remaining Figs. with scale 0.5 mm. 


184 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Figs. 136-146. Male terminalia, left lateral view: 136, Trypetolimnia; 137, Psacadina; 138, Steyskalina; 139, 
Dictyodes; 140, Ilione; 141, Pherbina; 142, Trypetoptera; 143, Limnia; 144, Poecilographa; 145, Pherbecta; 
146, Protodictya. Cerci (cer.); epandrium (ep.); sternite 6 (st. 6); synsternite 7+8 (syn. 7+8); synsternite 6+7+8 


(syn. 6+7+8); anterior surstylus (a. s.); posterior surstylus (p. s.). Figs. 138 and 146 with scale 0.5 mm; the 
remaining Figs. with scale 1.0 mm. 


VOLUME 113, NUMBER | 


0,5 mm 


syn. 6+7+8 7 
mm 
i ase 2 
syn. 6+7+8 
<a ae 


Figs. 147-160. Male terminalia, left lateral view: 147, Guatemalia; 148, Elgiva; 149, Hedria; 150, Diche- 


tophora; 151, Coremacera; 152, Dictyacium; 153, Euthycera; 154, Ethiolimnia; 155, Thecomyia; 156, Sepedon- 
inus; 157, Sepedonella; 158, Sepedon; 159, Sepedomerus; 160, Sepedonea. Cerci (cer.); epandrium (ep.); sternite 
6 (st. 6); synsternite 7+8 (syn. 7+8); synsternite 6+7+8 (syn. 6+7+8); posterior surstylus (p. s.). Figs. 148— 
150, 154, 155 with scale 1.0 mm; the remaining Figs. 0.5 mm. 


186 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Figs. 161-163. Male terminalia, posterior view: 161, Sciomyza; 162, Oidematops; 163, Atrichomelina. Cerci 
(cer.); subepandrial plate (sub. pl.); anterior surstylus (a. s.); posterior surstylus (p. s.). 


subgroup, from Thecomyia to Sepedonea, is group comprising the genera Thecomyia, 
established by eight characters: 6, 19, 28, Sepedon, Sepedonella, Sepedoninus, Sepe- 
30, 31, 32, 33 and 35. It is the most cor- domerus, and Sepedonea. Steyskal (1973: 
roborated lineage within this analysis. 143) stated that “I do not believe that this 

Steyskal (1973) placed Sepedon in a_ group is sufficiently distinct from more typ- 


VOLUME 113, NUMBER 1 


0,5 mm 


164 


187 


Figs. 164-169. 


Male terminalia, posterior view: 164, Tetanura; 165, Colobaea; 166, Calliscia; 167, Parec- 


tinocera; 168, Pteromicra; 169, Pherbellia. Cerci (cer.); subepandrial plate (sub. pl.); anterior surstylus (a. s.); 


posterior surstylus (p. s.). 


ical Tetanocerini, especially from such gen- 
era as Hedria and Dichetophora, to be giv- 
en the rank of tribe or even subtribe. We 
would prefer to call it merely “‘the Sepedon 
group.” ”’ This statement agrees with the 
result of the present analysis, but with the 
addition the genera Ethiolimnia and Teu- 
toniomyia to the Sepedon group. 

Proposal of classification.—The follow- 
ing natural classification was done using the 


sequencing convention (Wiley 1981). The 
subfamilies Salticellinae and Sciomyzinae 
and the tribes Sciomyzini and Tetanocerini 
are confirmed. 
Sciomyzidae Fallén, 1820 
Salticellinae Hendel, 1924 

Salticella Robineau-Desvoidy, 1830 
Sciomyzinae Fallén, 1820 

Sciomyzini Fallén, 1820 

Pherbellia Robineau-Desvoidy, 1830 


188 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


p.s. 
| 171 
, 173 
= : 
175 
174 176 


Figs. 170-176. Male terminalia, posterior view: 170, Antichaeta; 171, Shannonia; 172, Perilimnia; 173, Hoplod- 
ictya; 174, Dictya (lateral view), 175, Hydromya; 176, Neolimnia. Anterior surstylus (a. s.); posterior surstylus (p. s.). 


0,5 mm 


a.S. 


Pteromicra Lioy, 1864 Eutrichomelina Steyskal, 1975 
Calliscia Steyskal, 1975 Ectinocera Zetterstedt, 1838 
Parectinocera Becker, 1919 Renocera Hendel, 1900 
Colobaea Zetterstedt, 1837 Antichaeta Haliday, 1838 
Tetanura Fallén, 1820 Chasmacryptum Becker, 1907 
Atrichomelina Cresson, 1920 Shannonia Malloch, 1933 
Oidematops Cresson, 1920 Perilimnia Becker, 1919 
Sciomyza Fallén, 1820 Hoplodictya Cresson, 1920 


Tetanocerini Hendel, 1900 Dictya Meigen, 1803 


VOLUME 113, NUMBER 1 189 


177 


0,5 mm 


” 181 


182 


Figs. 177-186. Male terminalia, posterior view: 177, Euthycerina; 178, Tetanocera; 179, Trypetolimnia; 
180, Psacadina; 181, Steyskalina; 182, Dictyodes; 183, Ilione; 184, Pherbina; 185, Trypetoptera; 186, Limnia. 


Anterior surstylus (a. s.); posterior surstylus (p. s.). 


190 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


189 
ip [ 
a.s. Nace | 
‘ic 
(‘ 7 
p.s 
191 
190 
. 7 mn 
192 193 
194 


: =—ias 
p.s. 
195 Wee 


Figs. 187-196. Male terminalia, posterior view: 187, Poecilographa; 188, Pherbecta; 189, Protodictya; 190, 
Guatemalia; 191, Elgiva; 192, Hedria; 193, Dichetophora; 194, Coremacera; 195, Dictyacium; 196, Euthycera. 


Anterior surstylus (a. s.); posterior surstylus (p. s.). 


Hydromya Robineau-Desvoidy, 1830 Tetanocera Duméril, 1800 
Neolimnia Barnes, 1976 Trypetolimnia Mayer, 1953 
Tetanoceroides Malloch, 1933 Psacadina Enderlein, 1939 


Euthycerina Malloch, 1933 Dictyodes Malloch, 1933 


VOLUME 113, NUMBER 1 19] 


198 


Ke ‘ 


Figs. 197-203. Male terminalia, posterior view: 197, Ethiolimnia; 198, Thecomyia; 199, Sepedoninus, 200, Se- 
pedonella; 201, Sepedon; 202, Sepedomerus; 203, Sepedonea. Anterior surstylus (a. s.); posterior surstylus (p. s.). 


Ilione Verbeke, 1964 Ethiolimnia Verbeke, 1950 
Pherbina Robineau-Desvoidy, 1830 Teutoniomyia Hennig, 1952 
Trypetoptera Hendel, 1900 Thecomyia Perty, 1833 
Limnia Robineau-Desvoidy, 1830 Sepedoninus Verbeke, 1950 
Poecilographa Melander, 1913 Sepedonella Verbeke, 1950 
Pherbecta Steyskal, 1956 Sepedon Latreille, 1804 
Protodictya Malloch, 1933 Sepedomerus Steyskal, 1973 
Guatemalia Steyskal, 1960 Sepedonea Steyskal, 1973 
Elgiva Meigen, 1838 Steyskalina Knutson, 1999 


Hedria Steyskal, 1954 
Dichetophora Rondani, 1868 
Coremacera Rondani, 1856 
Dictyacium Steyskal, 1920 The subfamilies Salticellinae and Scio- 
Euthycera Latreille, 1829 myzinae are monophyletic. In the Sciomy- 


Conclusions 


192 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


205 207 


209 210 


« 


208 


212 
213 

Figs. 204-213. Male terminalia. 204, 208, 206, 210, 212, Right lateral view; 205, 207, 209, 211, 213, frontal 
view. 204, 205, Sciomyza; 206, 207, Oidematops; 208-209, Atrichomelina; 210-211, Tetanura; 212—213, Co- 
lobaea. Aedeagal apodeme (aed. ap.); basiphallus (bas.); aedeagus (aed.); gonopod (gon.); hypandrium (hip.); 
paramere (par.). 


VOLUME 113, NUMBER 1 


> 
> 

. 
. 


“e 
ve 
vee 


eve 


7 We | 


216 


218 
219 


221 


Male terminalia. 214, 216, 218, 220, Right lateral view; 215, 217, 219, 221, frontal view. 


Figs. 214-221. 
214, 215, Calliscia; 216, 217, Parectinocera; 218, 219, Pteromicra; 220, 221, Pherbellia. Aedeagal apodeme 


(aed. ap.); basiphallus (bas.); aedeagus (aed.); gonopod (gon.); hypandrium (hip.); paramere (par.). 


194 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


par. 


aed. 


0,5 mm 


227 228 
: 229 


aed. 


>, 
< 33 
ee Youd vis 
¥ 
"weve vec 
Othe? 1 2 
ae 3 1 


230 
Figs. 222—232. Male terminalia. 222, 225, 227, 230, Right lateral view; 223, 226, 228, 231, frontal view; 
224, 229, 232, left lateral view. 222-224, Ectinocera; 225—226, Antichaeta; 227—229, Chasmacryptum; 230- 


232, Shannonia. Aedeagal apodeme (aed. ap.); basiphallus (bas.); aedeagus (aed.); paramere (par.). 


VOLUME 113, NUMBER 1 


aed. 


aed. 


0,5 mm 


Figs. 233-241. 


195 


234 


241 


Male terminalia. 233, 236, 239, Right lateral view; 234, 237, 240, frontal view; 235, 238, 


241, left lateral view. 233-235, Perilimnia; 236—238, Hoplodictya,; 239-241, Dictya. Aedeagal apodeme (aed. 


ap.); basiphallus (bas.); aedeagus (aed.); paramere (par.). 


zinae the tribes Sciomyzini and Tetanocer- 
ini are monophyletic. 

The tribe Sciomyzini includes Pherbel- 
lia, Pteromicra, Calliscia, Parectinocera, 
Colobaea, Tetanura, Atrichomelina, Oide- 
matops, and Sciomyza. 


The tribe Tetanocerini includes Antichae- 
ta, Chasmacryptum, Coremacera, Dicheto- 
Phora, Dictya, Dictyacium, Dictyodes, Ec- 
tinocera, Elgiva, Ethiolimnia, Euthycera, 
Euthycerina, Eutrichomelina, Guatemalia, 
Hedria, Hydromya, Hoplodictya, Ilione, 


196 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


0,5 mm 
NO 
BSS 
NO 


aed.ap. 
< =e 
247 


245 246 


249 250 


253 


252 


Figs. 242-253. Male terminalia. 242, 245, 248, 251, Right lateral view; 243, 246, 249, 252, frontal view; 
244, 247, 250, 253, left lateral view. 242-244, Hydromya; 245-247, Neolimnia; 248-250, Euthycerina; 251-— 
253, Tetanocera. Aedeagal apodeme (aed. ap.); aedeagus (aed.); paramere (par.). 


VOLUME 113, NUMBER 1 197 


0,5 mm 


par. 


257 258 259 


es 
261 ES) Fe 


aed. 


265 


263 


Figs. 254-265. Male terminalia. 254, 257, 260, 263, Right lateral view; 255, 258, 261, 264, frontal view; 
256, 259, 262, 265, left lateral view. 254—256, Trypetolimnia; 257-259, Psacadina; 260—262, Steyskalina; 263-— 


265, Dictyodes. Aedeagal apodeme (aed. ap.); basiphallus (bas.); aedeagus (aed.); paramere (par.). 


198 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


ew 


273 274 


Figs. 266-274. Male terminalia. 266, 269, 272, Right lateral view; 267, 270, 273, frontal view; 268, 271, 


274, left lateral view. 266—268, Ilione; 269-271, Pherbina; 272-274, Trypetoptera. Aedeagal apodeme (aed. 
ap.); basiphallus (bas.); aedeagus (aed.); paramere (par.). 


Limnia, Neolimnia, Perilimnia, Pherbecta, nea, Shannonia, Steyskalina, Tetanocera, 
Pherbina, Poecilographa, Protodictya, Tetanoceroides, Trypetoptera, Trypetolim- 
Psacadina, Renocera, Sepedoninus, Sepe- nia, Teutoniomyia, and Thecomyia. 

donella, Sepedon, Sepedomerus, Sepedo- Within the Sciomyzini a monophyletic 


VOLUME 113, NUMBER | 199 


SP 
279 


par. 


284 286 


Figs. 275—286. Male terminalia. 275, 278, 281, 284, Right lateral view; 276, 279, 282, 285, frontal view; 
377, 380, 383, 386, left lateral view. 275-277, Limnia; 278-280, Poecilographa; 281-283, Pherbecta; 284-286, 
Protodictya. Aedeagal apodeme (aed. ap.); aedeagus (aed.); paramere (par.). 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


aed.ap. 
aed. 
£ 
E 
287 288 3 | 4 


par. 


200 


298 


y aed.ap. 
bas. OJ 
eR 29 , 295 


Figs. 287-298. Male terminalia. 287, 290, 293, 296, Right lateral view; 288, 291, 294, 297, frontal view 
289, 292, 295, 298, left lateral view. 287-289, Guatemalia; 290-292, Elgiva; 293-295, Hedria; 296-298, Di- 


chetophora. Aedeagal apodeme (aed. ap.); basiphallus (bas.); aedeagus (aed.); paramere (par.) 


VOLUME 113, NUMBER 1 


0,5 mm 


aed. 


201 


301 
300 


Figs. 299-307. Male terminalia. 299, 302, 305, Right lateral view; 300, 303, 305, frontal view; 301, 304, 
306, left lateral view. 299-301, Coremacera; 302-304, Dictyacium; 305-307, Euthycera. Aedeagal apodeme 
(aed. ap.); basiphallus (bas.); aedeagus (aed.); paramere (par.). 


group was formed by the genera Colobaea, 
Tetanura, Atrichomelina, Oidematops + 
Sciomyza. In this group there is the sub- 
group formed by Atrichomelina, Oidema- 
tops + Sciomyza. 


The groups of Shannonia + Perilimnia 
and Hoplodictya + Dictya, at the base of 
Tetanocerini, are monophyletic. Three 
other major groups of genera are formed 
within the Tetanocerini: the first by Eu- 


202 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


> 


> 
Dias 
SOS 


2B 
> 
ie 


? 


2r5? 
>? 


0,5 mm 


312 313 


Figs. 308-315. Male terminalia. 308, 311, 314, Right lateral view; 309, 312, 315, frontal view; 310, 313, 
left lateral view. 308-310, Ethiolimnia; 311-313, Thecomyia; 314, 315, Sepedoninus. Aedeagal apodeme (aed. 
ap.); basiphallus (bas.); aedeagus (aed.); paramere (par.). 


thycerina, Psacadina Enderlein (1933), dictya, and the third by Guatemalia, El- 
Steyskalina, Tetanocera, and Trypetolim- giva, Dichetophora, Hedria, Coremacera, 
nia, the second by the genera Dictyodes,- Dictyacium, Euthycera, Ethiolimnia, Teu- 
Ilione, Pherbina, Trypetoptera, Limnia, toniomyia, Thecomyia, Sepedoninus, Se- 
Poecilographa, and Pherbecta + Proto-  pedonella, Sepedon, Sepedomerus + Se- 


VOLUME 113, NUMBER 1 203 


316 317 


0,5 mm 


—— aed.ap. 


ee par. 


aed. 


‘318 


322 


Figs. 316-322. Male terminalia. 316, 318, 320, 322, Right lateral view; 317, 319, 321, frontal view. 316, 
317, Sepedonella; 318, 319, Sepedon; 320, 321, Sepedomerus; 322, Sepedonea. Aedeagal apodeme (aed. ap.); 
aedeagus (aed.); paramere (par.). 


pedonea. Within the last group the genera _ ships; and the group of Ethiolimnia, Teu- 
Coremacera, Dictyacium, and Euthycera  toniomyia, Thecomyia, Sepedoninus, Se- 
form a monophyletic clade without any pedonella, Sepedon, Sepedomerus + Se- 
resolution of their phylogenetic relation- pedonea is monophyletic. 


nN 
C= ie) 
nN a Nn 
— nN = 
= = 
oe 
= ez 
_ 
©-2 On Rw rm) 
= Mm-0 2 = =~ 
— 
— 
ey 
— 
—_ 
ra) 
g 
_ 
~ 
_ 
_ 
GC} 
XN =a = 
BA me YES 
© 
_ 
_ 
~ At 
=) 
_ 
o 
nN 
— 
— 
= 
~ 
SS 
-_ 
N 
a 
_ 
™~ 
So 
Le) 
N 
=) 
wo 
NN 
ID 
o = 
Sy 
—_ 
—_ 
S 
-e as 
—_ 
—_= oO 
C) 
RROD Ses el 
O©ONSOO& N 
- SOON |] W& 
=- Nn N 
NI 
= 
OWWWWWN AD 
AOn20800N 
=- = = -—- = _ 
gRrex 
on 
2 NN 
eo @o 
a — 
= N 
wONN 
oud 
on;,DM WY 
w 
“ 
_ 
Fig 2323: 


bpe '€z 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Pelidnoptera PA 
Salticella PA AT 
Sciomyza NA PA 
Oidematops NA 
Atrichomelina NA NT 
Tetanura PA 
Colobaea NA PA AT 
Calliscia NT 
Parectinocera NT 
Pteromicra NA PA 
Pherbellia NA PA NT AT OR AS 
Eutrichomelina NT 
Ectinocera PA 
Renocera NA PA 
Antichaeta NA PA 
Chasmacryptum PA 
Shannonia NT 
Perilimnia NT 


. Hoplodictya NA NT 


Dictya NA PA NT 
Hydromya PA 
Neolimnia AS 
Tetanoceroides NT 
Euthycerina NT 
Tetanocera NA PA OR 
Trypetolimnia PA 
Psacadina PA 
Steyskalina OR 
Dictyodes NT 

llione PA NA OR 
Pherbina PA 
Trypetoptera NA PA 
Limnia NA PA 
Poecilographa NA 
Pherbecta NA 
Protodictya NT 
Guatemalia NT 
Elgiva NA PA 

Hedria NA 
Dichetophora PA OR AS 
Coremacera PA 
Dictyacium NA 
Euthycera NA PA NT 
Ethiolimnia AT 
Teutoniomyia NT 
Thecomyia NT 
Sepedoninus AT 
Sepedonella AT 
Sepedon NA PA NT AT OR AS 
Sepedomerus NT OR 
Sepedonea NT 


Consensus cladogram. Outgroup Pelidnoptera. 37 characters. Consistency index: 0.60. Retention 


index: 0.88. Regions: AT, Afrotropical; AS, Australian; NA, Nearctic; NT, Neotropical; OR, Oriental; PA, Pa- 


learctic. 


VOLUME 113, NUMBER 1 205 


OL 
db} 
BL 
6L 


Pelidnoptera 
1} {| tt | | | Satticelta 


| | | Sciomyza 


Atrichomelina 


tT } | t—4+--4 } 
| | | 
| 4h a feel Tetanura 
} 


Colobaea 


|_| 
| | | | | Oidematops 


were IT TT] 111) camiseta 


| | | Parectinocera 
tied Seles | | Pteromicra 
Pherbellia 
Eutrichomelina 
g | | Ectinocera 

| | | Renocera 

| | Antichaeta 


ey Chasmacryptum 
| Shannonia 


hele Perilimnia 
Hoplodictya 

Pee Dictya 

| | Hydromya 


‘z 


Neolimnia 


Tetanoceroides 


Euthycerina 


Tetanocera 


Trypetolimnia 


Psacadina 


Steyskalina 

| | | Dictyodes 

| llione 
Pherbina 
Trypetoptera 


| 
Limnia 


| | Poecilographa 
Pherbecta 
Protodictya 
Guatemalia 


ROMER RT BASE me! = = Ta Re ae ae es as a 


Elgiva 
Hedria 
Dichetophora 


- 


| 
| 
+ 


= 


Coremacera 


1 
| 


Dictyacium 


Euthycera 


Ethiolimnia 


| 


Teutoniomyia 


1 
i 


Thecomyia 


Sepedoninus 


Sepedonelia 


Sepedon 


=|a 
SEIMESESaIeE 


Sepedomerus 


Sepedonea 


Fig. 324. Arrangement of character states among genera of Sciomyzidae. White = plesiomorphic condition; 
gray (light and dark) = intermediate and relatively more apomorphic conditions for multistate characters (pro- 
gressing from light gray to dark gray); black = apomorphic condition. 


Acknowledgments 


Many individuals and institutions have 
contributed to this analysis. We thank the 
following institutions and respective cura- 
tors for their cooperation in lending speci- 
mens. 
BMNH _ The Natural History Museum, 
London, England 
Colecao de Entomologia Pe. J. 
S. Moure, Department of Zool- 
ogy, Universidade Federal do 
Parana, Curitiba, Parana, Brazil 
Instituto Nacional de Pesquisas 
da Amazonia, Manaus, Ama- 
zonas, Brazil 
Museu Nacional, Universidade 
Federal do Rio de Janeiro, Rio 
de Janeiro, Rio de Janeiro, Bra- 
zil 
Museu de Zoologia da Univer- 
sidade de Sao Paulo, Sao Paulo, 
Sao Paulo, Brazil 
University of California, River- 
side, California, U.S.A. 
former United States National 
Museum, collections in the Na- 
tional Museum of Natural His- 
tory, Smithsonian Institution, 
Washington, D.C., U.S.A. 

We thank Dr. Claudio J. B. de Carvalho, 
Dr* Marcia S. Couri, Dr* Dilma S. Napp, & 
Dr. Angelo P. do Prado for reviewing the 
manuscript and Dr. John Kingsolver for re- 
viewing the English version. We also thank 
Drs. Rudolf RozkosSny & Lloyd V. Knutson 
for very constructive discussions on nu- 
merous aspects and the conclusions that are 
presented in this paper. 


DZUP 


INPA 


MNRJ 


MZSP 


UCR 


USNM 


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nana eats worms.—Science (in press). 

Valley, K., & C. O. Berg. 1977. Biology, immature 
stages, and new species of snail-killing Diptera 
of the genus Dictya (Sciomyzidae).—Search of 
Agricultural Entomology 7(2):1—45. 

Watrous, L. E., & Q. D. Wheeler. 1981. The out-group 
comparison method of character analysis.— 
Systematic Zoology 30(1):1-11. 

Wiley, E. O. 1981. Phylogenetics: the theory and prac- 


VOLUME 113, NUMBER 1 


tice of phylogenetic systematics. John Wiley 
and Sons, New York. 

, D. Siegel-Causey, D. R. Brooks, & V. A. 
Funk. 1991. The compleat cladist: a primer of 
phylogenetic procedures. Special Publication 
19, The University of Kansas, Museum of Nat- 
ural History, Lawrence, Kansas, 158 pp. 


Appendix 1. 


List of genera and type species used in 
the analysis. 
Antichaeta analis Meigen, 1830 
Atrichomelina pubera (Loew), 1862 
Calliscia calliscelles (Steyskal), 1963 
Chasmacryptum seriatimpunctatum Becker, 

1907 
Colobaea bifasciella (Fallén), 1820 
Coremacera marginata (Fabricius), 1775 
Dichetophora obliterata (Fabricius), 1805 
Dictya umbrarum (Linnaeus), 1758 
Dictyacium ambiguum (Loew), 1864 
Dictyodes dictyodes (Wiedemann), 1830 
Ectinocera borealis Zetterstedt, 1838 
Elgiva cucularia (Linnaeus), 1767 
Ethiolimnia platalea Verbeke, 1950 
Euthycera chaerophylli (Fabricius), 1798 
Euthycerina vittithorax Malloch, 1933 
Eutrichomelina fulvipennis (Walker), 1837 
Guatemalia straminata (Wulp), 1897 
Hedria mixta Steyskal, 1954 
Hoplodictya setosa (Coquillett), 1901 
Hydromya dorsalis (Fabricius), 1775 


209 


Ilione albiseta (Scopoli), 1763 
Limnia unguicornis (Scopoli), 1763 
Neolimnia repo Barnes, 1976 
Oidematops ferrugineus Cresson, 1920 
Parectinocera neotropica Becker, 1919 
Perilimnia albifacies Becker, 1919 
Pherbecta limenitis Steyskal, 1956 
Pherbellia punctata (Fabricius), 1791 
Pherbina coryleti (Scopoli), 1763 
Poecilographa decora (Loew), 1864 
Protodictya chilensis Malloch, 1933 
Psacadina disjecta Enderlein, 1939 
Pteromicra glabricula (Fallén), 1820 
Renocera pallida (Fallén), 1820 
Salticella fasciata (Meigen), 1830 
Sciomyza simplex Fallén, 1820 
Sepedomerus macropus (Walker), 1849 
Sepedon sphegea (Fabricius), 1775 
Sepedonea lindneri (Hendel), 1932 
Sepedonella nana Verbeke, 1950 
Sepedoninus planifrons Verbeke, 1950 
Shannonia meridionalis Zuska, 1969 
Steyskalina picta Ghorpadé and Marinoni, 
1999 
Tetanocera elata (Fabricius), 1781 
Tetanoceroides mesopleuralis Malloch, 
1933 
Tetanura pallidiventris Fallén, 1820 
Teutoniomyia plaumanni Hennig, 1952 
Thecomyia longicornis Perty, 1833 
Trypetolimnia rossica Mayer, 1953 
Trypetoptera punctulata (Scopoli), 1763 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 
113(1):210—217. 2000. 


Review of the chewing louse genus Abrocomophaga 
(Phthiraptera: Amblycera), with description of two new species 


Roger D. Price and Robert M. Timm 


(RDP) Department of Entomology and Plant Pathology, Oklahoma State University, 
Stillwater, Oklahoma 74078-0464, U.S.A. (Current address) 4202 Stanard Circle, 
Fort Smith, Arkansas 72903-1906, U.S.A.; 

(RMT) Natural History Museum and Department of Ecology & Evolutionary Biology, 
University of Kansas, Lawrence, Kansas 66045-2454, U.S.A. 


Abstract.—The South American chewing louse Abrocomophaga chilensis 
Emerson & Price has, since its description, remained the sole known member 
of the amblyceran family Abrocomophagidae. We herein provide a redescrip- 
tion of the species as well as descriptions and illustrations for two new species 
of Abrocomophaga: A. emmonsae off Cuscomys ashaninka Emmons from Pert 
and A. hellenthali off the degu, Octodon degus (Molina), from Chile. After our 
reevaluation of the status of the family Abrocomophagidae, we consider it a 
synonym of the family Gyropidae. 

Resumen.—Desde su descripcion original, el piojo sudamericano Abrocom- 
ophaga chilensis Emerson & Price, ha permanecido como el inico miembro 
conocido de la familia Abrocomophagidae, perteneciente a los ambliceros. En 
este trabajo presentamos una redescripcion de dicha especie y ademas descrip- 
ciones e ilustraciones de dos especies nuevas del género Abrocomophaga: A. 
emmonsae parasito de Cuscomys ashaninka Emmons proveniente del Pert y 
A. hellenthali parasito del degu, Octodon degus (Molina), de Chile. Después 
de reevaluar el status de la familia Abrocomophagidae, la consideramos como 


sin6nimo de la familia Gyropidae. 


Emerson & Price (1976) described a dis- 
tinctive new taxon of South American 
chewing louse, Abrocomophaga chilensis, 
and placed it in a new monotypic family, 
the Abrocomophagidae. All known speci- 
mens of this louse were found on a single 
chinchilla rat, Abrocoma bennetti Water- 
house, from Chile. Since the original de- 
scription, little has been written about this 
enigmatic and phylogenetically important 
amblyceran (Emerson & Price 1985), al- 
most certainly because no new information 
was available. We recently obtained series 
of Abrocomophaga from two other South 
American rodent hosts; these lice represent 
two new species in this genus. With the ad- 
dition of these two new species, we reeval- 
uate the status of the family Abrocomo- 


phagidae and the genus Abrocomophaga. 
Our purpose here is to describe and illus- 
trate these new species and to establish the 
synonymy for the family Abrocomophagi- 
dae. 


Family Gyropidae Kellogg 


Gyropidae Kellogg, 1896:68. Type genus: 
Gyropus Nitzsch. 

Abrocomophagidae Emerson & Price, 
1976:425. Type genus: Abrocomophaga 
Emerson & Price. New synonymy. 


The features associated with the species 
of Gyropidae and the relationship of this 
family to other members of the suborder 
Amblycera have been accurately treated by 
Clay (1970) and will not be repeated here. 


VOLUME 113, NUMBER 1 


In her treatment of the amblyceran Phthir- 
aptera, Clay (1970) followed the organiza- 
tion presented by Ewing (1924) in recog- 
nizing the Gyropidae as being divided into 
three subfamilies: (1) the Gyropinae with 
six pairs of abdominal spiracles (on III-— 
VIII) and at least one pair of legs having a 
modified tarsal claw strongly adapted for 
clasping hair; (2) the Gliricolinae with only 
five pairs of abdominal spiracles (on II-— 
VII) and legs with a single greatly reduced 
tarsal claw; and (3) the Protogyropinae with 
six pairs of abdominal spiracles (on II-— 
VIII) and all legs having a single unmodi- 
fied tarsal claw. 

The establishment of the family Abro- 
comophagidae by Emerson & Price (1976) 
was based primarily on the fact that Abro- 
comophaga chilensis, the sole member of 
this new family, had an unmodified tarsal 
claw on each leg and only five pairs of ab- 
dominal spiracles, a combination of char- 
acters seen in no other gyropid louse. How- 
ever, as our understanding of character evo- 
lution has matured over the years, we have 
come to the conclusion that these differenc- 
es are of generic-level rather than familial- 
level significance. In fact, the main differ- 
ence between the Abrocomophagidae and 
the monotypic Protogyropinae was that the 
former had only five pairs of abdominal 
spiracles (on III—VII), whereas the latter 
had six pairs (on III—VIII). Because the 
number of abdominal spiracles in the Gy- 
ropidae varies from the primitive six pairs 
to a reduced state of five pairs, it is our 
feeling now that this difference is not of 
familial significance and that the Abrocom- 
ophagidae should be considered a synonym 
of the family Gyropidae (subfamily Proto- 
gyropinae) as delineated by Clay (1970). A 
discussion of other features associated with 
the Abrocomophagidae and Abrocomopha- 
ga may be found in Emerson & Price 
(1976). Lakshminarayana (1976) presented 
a thorough review of the suprageneric clas- 
sification of the Phthiraptera and gave a list- 
ing of the superfamily, family, subfamily, 
and tribe names proposed to that time. 


211 


Genus Abrocomophaga Emerson & Price 


Abrocomophaga Emerson & Price, 1976: 
425. Type species: Abrocomophaga chi- 
lensis Emerson & Price. 


This genus is separated from others in the 
family Gyropidae (and suborder Ambly- 
cera) in having known representatives with 
all legs having an unmodified tarsal claw 
and the abdomen with only five pairs of spi- 
racles. 


Abrocomophaga chilensis Emerson & 
Piice 


Abrocomophaga chilensis Emerson & 
Price, 1976:426. Type host: Abrocoma 
bennetti bennetti Waterhouse. 


Male.—Emerson & Price (1976) provide 
a full illustration in Fig. 5 and two aspects 
of the male genitalia in Figs. 3 and 4. Head 
with numerous medium-length setae on 
dorsal and ventral surfaces; without heavy 
dorsal seta near antennal base. Pronotum 
with 10—14 setae; prosternum with 5-7 se- 
tae, mesosternum with 8—10. Metanotum 
with 10 setae, including 6 longer median 
setae and each side laterally with 1 long, 1 
short seta; metasternal plate with 12—14 se- 
tae. Abdominal tergal setae: I, 4; II, 6—8; 
III, 8—10; IV, 10-12; V, 11-13; VI, 11-14; 
VII, 10-12; VIII, 6-9. Terga III—VII each 
with small faint median pigmentation; V— 
VIII each with median pair of shorter setae 
recessed posterior to main row. Posterior 
margin of each of pleura IIJ—VIII with 1 
very long and 1 adjacent short heavy seta. 
Last tergum with 2 short, 1 very long seta 
on each side. Abdominal sternal setae: II, 
7-9; Il, 9-12; IV, 11-15; V, 12-14; VI, 
10—13; VII, 9—11. Sterna IV—VII each with 
small faint median pigmentation. Subgeni- 
tal plate with 7 or 8 medium to short me- 
dian setae. Genitalia with only ill-defined 
weak mesosomal sclerites associated with 
sac and with blunt apical tip of parameres. 
Dimensions (in mm): temple width, 0.19— 
0.20; head length, 0.16—0.18; prothorax 
width, 0.15—0.16; metathorax width, 0.18— 


212 


0.20; abdomen width at segment V, 0.30— 
0.31; genitalia width, 0.08; genitalia length, 
0.17—0.20; genitalia paramere length, 0.06— 
0.07; total length, 0.94—0.98. 

Female.—Emerson & Price (1976) pro- 
vide a full illustration in Fig. 2 and one of 
the ventral terminalia in Fig. 1. Head and 
thorax as for male, abdomen differing as 
follows. Tergal setae: IV, 9-11; V, 10-12; 
VI, 10-13; VII, 9-12. All setae on terga V— 
VIII aligned in row. Last tergum with 2 
very long setae on each side, in addition to 
2 shorter setae. Sternal setae: II, 8—10; III, 
10-13; IV, 11-13; V, 11-14; VI, 11-13; 
VII, 8-11. Subgenital plate with 9-11 me- 
dioanterior setae and convex posterior mar- 
gin bearing median fringe of small spicules. 
Anal fringe ventrally of 5 + 5 and dorsally 
of 3 + 3 longer lateral setae, with few mi- 
nute median setae in each row. Dimensions 
(an mm): temple width, 0.20—0.22; head 
length, 0.17—0.20; prothorax width, 0.16— 
0.17; metathorax width, 0.19—0.20; abdo- 
men width at segment V, 0.32—0.35; anus 
width, 0.07—0.08; total length, 1.07—1.14. 

Material.—10 male, 10 female paratypes 
of Abrocomophaga chilensis, ex Abrocoma 
bennetti bennetti, Chile. 

Remarks.—This species is recognized by 
both sexes with consistently small dimen- 
sions, including narrow head and abdomen, 
and their abdominal tergal and sternal setal 
counts; the female with a narrow anus; and 
the male with unique genitalia and a small 
recessed pair of abdominal tergal setae on 
each of V-—VIII. Type deposited in the 
NMNH. 

Emerson & Price described Abrocomo- 
phaga chilensis on the basis of a large se- 
ries of specimens from a single host indi- 
vidual of Abrocoma bennetti bennetti from 
Chile collected by Robert E. Martin. We 
can now provide more specific information 
on the type host and type locality. The type 
host collected by Robert E. Martin on 2 No- 
vember 1974 bears his field number 1105 
and is deposited in the Recent mammal col- 
lection at the Field Museum in Chicago as 
FMNH 119794 with locality information as 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


follows: Chile: Santiago Province; 10 km 
W of Til Til [Tiltil; 1000 m]. The host was 
an adult female. 


Abrocomophaga emmonsae, new species 
Figs. 1—4 


Type host.—Cuscomys ashaninka Em- 
mons. 

Male.—As in Fig. 1. Differing from 
Abrocomophaga chilensis as follows. Ab- 
dominal tergal setae: IV, 14; V, 18; VI, 14; 
VI, 16. Terga I-VII each with small me- 
dian faintly pigmented area; V—VII each 
with median pair of shorter setae recessed 
posterior to main row. Abdominal sternal 
setae: IT, 10; III, 14; IV, 16; V, 15; VI-VII, 
12. Sterna I—VII each with lightly pig- 
mented median area. Genitalia (Fig. 3) with 
prominent inwardly curved sharply pointed 
parameres and conspicuous associated me- 
sosomal sclerites as shown. Dimensions (in 
mm): temple width, 0:26; head length, 0.20; 
prothorax width, 0.19; metathorax width, 
0.26; abdomen width at segment V, 0.40; 
genitalia width, 0.09; genitalia length, 0.20; 
genitalia paramere length, 0.06; total 
length, 0.98. 

Female.—As in Fig. 2. Differing from 
Abrocomophaga chilensis as follows. Ab- 
dominal tergal setae: IV, 12—13; V, 12-16; 
VI, 13-18; VII, 13-16. Abdominal sternal 
setae: II, 8-11; III, 11-15; IV, 13-17; V, 
13—16; VI, 12—18; VII, 11-14. Ventral ter- 
minalia as in Fig. 4. Dimensions (in mm): 
temple width, 0.26—0.27; head length, 
0.20—0.22; prothorax width, 0.19—0.20; 
metathorax width, 0.23—0.26; abdomen 
width at segment V, 0.41—0.47; anus width, 
0.09—0.10; total length, 1.06—1.16. 

Type material.—Holotype male, ex Cus- 
comys ashaninka, Peré: Cuzco, Cordillera 
Vilcabamba: (11°39'36'S5/73°38'3 1 WwW) sek 
3350 m, 15 June 1997, coll. Louise H) Em- 
mons #1359; in collection of the Museo de 
Historia Natural, Universidad Nacional de 
San Marcos, Lima, Peri, MUSM 12715 °. 
Paratypes: 6 females, same data as holo- 
type; in collections of the Universidad Na- 


VOLUME 113, NUMBER 1 


213 


Lae 


Figs. 1-4. Abrocomophaga emmonsae: (1) Dorsal-ventral male; (2) Dorsal-ventral female; (3) Male geni- 


talia; (4) Ventral female terminalia. 


cional de San Marcos and the National Mu- 
seum of Natural History, Washington, D.C. 

Etymology.—This new species is named 
in honor of Louise H. Emmons, National 
Museum of Natural History, Smithsonian 
Institution, who collected the host and en- 


thusiastically encouraged the junior author 
to examine it for ectoparasites. Her survey 
efforts in South America have greatly con- 
tributed to our understanding of this fauna 
and her field guides to Neotropical mam- 
mals have created a broader understanding 


214 


of the rainforest for both the public and stu- 
dents of all ages, and have certainly con- 
tributed to conservation efforts. 
Remarks.—Although this new species is 
close to Abrocomophaga chilensis, both 
sexes are readily separable by their consis- 
tently larger dimensions, including broader 
head and abdomen, and their larger number 
of abdominal tergal and sternal setae; the 
female with the broader anus; and the male 
with the recessed pair of setae on only ab- 
dominal terga V—VII and with unique gen- 
italia that have the distinctive mesosomal 
sclerites and acutely pointed parameres. 


Abrocomophaga hellenthali, new species 
Figs. 5-8 


Type host.—Octodon degus (Molina). 
Male.—As for Abrocomophaga chilensis 
except as follows. Head (Fig. 6) with heavy 
dorsal seta near antennal base. Abdomen 
(Fig. 5) with tergal setae: II, 5—6; III, 6—7; 
IV-—V, 7-8; VI-VII, 8; VUI, 5—6. Terga I- 
VII each with small faint median pigmen- 
tation; V and VII each with median pair of 
shorter setae recessed posterior to main 
row. Posterior margin of each of pleura II-— 
VIII with short slender seta adjacent to very 
long seta. Last tergum with | long, 3-4 
short setae on each side. Abdominal sternal 
setae: II, 6; II, 8—9; IV, 9-10; V, 8-10; VI, 
7-8; VII, 6. Sterna II-VII each with small 
faint median pigmentation. Genitalia (Fig. 
7) with numerous conspicuous spinelike 
mesosomal sclerites associated with sac and 
with sharply pointed apical tip of parameres. 
Dimensions (in mm): temple width, 
0.23—0.26; head length, 0.18—0.20; protho- 
rax width, 0O.17—0.18; metathorax width, 
0.22—0.23; abdomen width at segment V, 
0.38—0.41; genitalia width, 0.08—0.09; gen- 
italia length, 0.19—0.21; genitalia paramere 
length, 0.07; total length, 0.83—0.88. 
Female.—Head and thorax as for male, 
abdomen (Fig. 8) differing as follows. Ter- 
gal setae: VIII, 6—7. All setae on terga V 
and VII aligned in row. Last tergum with 2 
very long setae on each side, in addition to 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


2 shorter setae. Sternal setae: II, 6—8; VI, 
8-9; VII, 6—7. Subgenital plate with 9-11 
medioanterior setae and convex posterior 
margin bearing median fringe of small spic- 
ules. Anal fringe as for Abrocomophaga 
chilensis. Dimensions (in mm): temple 
width, 0.24—0.26; head length, 0.19—0.20; 
prothorax width, 0.17—0.19; metathorax 
width, 0.22—0.26; abdomen width at seg- 
ment V, 0.43—0.47; anus width, 0.09—0.10; 
total length, 0.94—0.98. 

Type material.—Holotype male, ex Oc- 
todon degus, Chile: Santiago, Santiago, 2.5 
km NE of Cerro Manquehue, Trappist Mon- 
astery, 9 July 1975, coll. Robert E. Martin 
#1222 (FMNH 119614 &); in collection of 
the K. C. Emerson Entomology Museum, 
Oklahoma State University, Stillwater. 
Paratypes: 2 males, 3 females, same data as 
holotype; 1 female, same except 1 August 
1975, Robert E. Martin #1235 (FMNH 
119756 3); 1 male, 1 female, same except 
27 April 1976, Robert E. Martin #1432 
(FMNH 119639 <); all in same collection 
as holotype. 

Etymology.—This species is named for 
Ronald A. Hellenthal, University of Notre 
Dame, in recognition of his strong interest 
in chewing louse taxonomy and his many 
years of cooperative studies with RDP. 

Remarks.—This species is separated 
from the other two of the genus by both 
sexes having large dimensions, including a 
broad head and abdomen, a heavy dorsal 
preantennal head spine, a short slender seta 
adjacent to the very long seta on each of 
pleura IHI—VIII, and a consistently small 
number of abdominal tergal and sternal se- 
tae; the female with a broad anus; and the 
male with a recessed pair of setae only on 
abdominal terga V and VII and with unique 
genitalia that have a distinctive assemblage 
of spinous mesosomal sclerites and acutely 
pointed parameres. 


Discussion 


The caviomorph rodent family Octodon- 
tidae contains 6 genera and 11 living spe- 


VOLUME 113, NUMBER 1 


6 ae SS ees SE \ 


et 

thats ele sey 
Li ee eee ry 
Sian | 
“76 


Figs. 5-8. Abrocomophaga hellenthali: (5) Dorsal-ventral male abdomen; (6) Dorsal male head; (7) Male 


genitalia; (8) Dorsal-ventral female abdomen. 


cies, all with very restricted Andean or pre- 
Andean ranges; Octodon degus is the most 
widely distributed species. With the discov- 
ery of Cuscomys ashaninka, the cavio- 
morph family Abrocomidae now contains 
two Recent genera and a total of five spe- 
cies (Emmons 1999). Both families occupy 
a diverse array of high-elevation habitats, 
and their geographic ranges overlap broadly 
in southern South America. The relation- 
ships between the Abrocomidae and Octo- 
dontidae have been the subject of debate. 
The abrocomids were historically treated as 
a subfamily of the Echimyidae and more 
recently as either a subfamily of the Octo- 
dontidae or as a closely related family in 
the superfamily Octodontoidea following 
Reig (1986) and Carleton (1984). However, 
Glanz & Anderson (1990) suggested that 
the abrocomids are more closely aligned 
with the chinchillas and may belong in the 
superfamily Chinchilloidea. 

Where accurate records are available, we 
have found gyropid lice to be extremely 


host-specific ectoparasites (Price & Timm 
1997), with speciation of lice closely par- 
alleling speciation in their mammalian 
hosts. The various species in a louse genus 
are almost always restricted to the various 
species of a host genus or to very closely 
related genera of hosts. Our discovery of 
the two new species of Abrocomophaga de- 
scribed herein, although clearly supporting 
the host specificity we observe in gyropids, 
is odd in that closely related species of lice 
are found on different families of rodents. 
Two of the species of Abrocomophaga now 
known—A. chilensis and A. emmonsae— 
are apparently host-specific parasites of 
abrocomid rodents (Abrocoma bennetti and 
Cuscomys ashaninka, respectively) and one 
species—A. hellenthali—is apparently a 
host-specific parasite of an octodontid ro- 
dent (Octodon degus). Because only three 
species are known in the genus Abrocom- 
ophaga and the genus is only known from 
five separate host individuals in three sep- 


216 


arate genera, it is premature to attempt to 
reconstruct a phylogeny for the group. 
Naturally occurring transfers (host 
switching) of parasitic lice between nonre- 
lated hosts is not common, but has been 
documented in both mammals and birds 
(Paterson et al. 1999). Nest sharing between 
different species of mammals is rare in na- 
ture; however, sequential use of tree hole 
nests by birds is widespread. Timm (1983) 
postulated that the broad host distributions 
of species and genera seen in many lineages 
of bird lice are in part due to host transfers 
that might occurred during the evolutionary 
history of the groups. Mammal lice exhibit 
greater host specificity because opportuni- 
ties for lice to colonize new host taxa are 
rare. Although nest sharing by different 
species of rodents is exceedingly rare, it has 
been well documented for Octodon degus 
and Abrocoma bennetti in Chile. In a field 
study with marked individuals, Fulk (1976) 
found O. degus and A. bennetti regularly to 
share burrows and even the same nests. In 
excavating nests, he found female A. ben- 
netti with their own infants as well as infant 
degus. Correspondingly, female O. degus 
also had both species of young in their 
nests. Fulk (1976:504) considered nest 
sharing a common behavior in these two 
species, and suggested “‘This practice may 
be mutualistic, each animal contributing to 
the survival of the other’s young.” 
Although we don’t know that nest shar- 
ing occurs between Octodon degus and 
Abrocoma bennetti throughout their range, 
the fact that it does occur demonstrates how 
an ancestral Abrocomophaga could be 
transferred from one host family to another. 
With only three species of Abrocomophaga 
known to date, we cannot differentiate be- 
tween the hypotheses that the current dis- 
tribution of species we see on the rodent 
families Abrocomidae and Octodontidae 
are historically shared or relatively recent 
host transfers. When additional species of 
Abrocomophaga from other abrocomid and 
octodontid hosts are known, we may be 
able to address whether the Abrocomidae or 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


the Octodontidae, or perhaps a common an- 
cestor, represent the ancestral host for this 
enigmatic and poorly known group of lice. 


Acknowledgments 


We thank Louise Emmons and Linda 
Gordon, National Museum of Natural His- 
tory, Washington, D.C., for allowing us to 
remove specimens of lice from mammal 
specimens housed at the National Museum, 
thereby making the discovery of Abrocom- 
ophaga emmonsae and other new species 
possible. Larry Heaney and Bruce Patterson 
made our work at the Field Museum pos- 
sible and productive. Robert Anderson ex- 
amined specimens of Abrocoma for us as 
well as translated our Abstract into Spanish 
for the Resumen included herein. Rob An- 
derson, Barbara Clauson, Louise Emmons, 
Marcela Gomez-Laverde, and Ricardo Pal- 
ma provided valuable assistance on earlier 
drafts of this manuscript. This manuscript 
has been approved for publication by the 
Director, Oklahoma Agricultural Experi- 
ment Station, Stillwater. 


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ropidae.—Proceedings of the United States Na- 
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Fulk, G. W. 1976. Notes on the activity, reproduction, 


VOLUME 113, NUMBER 1 


and social behavior of Octodon degus.—Journal 
of Mammalogy 57:495—505. 

Glanz, W. E., & S. Anderson. 1990. Notes on Bolivian 
mammals. 7. A new species of Abrocoma (Ro- 
dentia) and relationships of the Abrocomidae.— 
American Museum Novitates 2991:1—32. 

Kellogg, V. L. 1896. New Mallophaga, I, with special 
reference to a collection made from maritime 
birds of the Bay of Monterey, California.—Pro- 
ceedings of the California Academy of Sciences 
(Series 2) 6:31—168. 

Lakshminarayana, K. V. 1976. Nomenclatural changes 
in Phthiraptera—some suggestions.—Ange- 
wandte Parasitologie 17:160—167. 

Paterson, A. M., R. L. Palma, & R. D. Gray. 1999. 
How frequently do avian lice miss the boat? 


217 


Implications for coevolutionary studies.—Sys- 
tematic Biology 48:214—223. 

Price, R. D., & R. M. Timm. 1997. A new subgenus 
and four new species of Gliricola (Phthiraptera: 
Gyropidae) from the Caribbean hutias (Roden- 
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ical Society of Washington 110:285-—300. 

Reig, O. A. 1986. Diversity patterns and differentiation 
of high Andean rodents. Pp. 404—440 in F. Vuil- 
leumier and M. Monasterio, eds., High altitude 
tropical biogeography. Oxford University Press, 
New York, 649 pp. 

Timm, R. M. 1983. Fahrenholz’s rule and resource 
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Pp. 225-265 in M. H. Nitecki, ed., Coevolution. 
University of Chicago Press, Chicago, 392 pp. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


113(1):218—223. 2000. 


A new species of the genus Cubacubana 
(Insecta: Zygentoma: Nicoletiidae) 
from a Mexican cave 


Luis Espinasa 


CEAMISH, Universidad Autonoma del Estado de Morelos, Av. Universidad 1001 Col. Chamilpa, 
Cuernavaca, Morelos CP 62210, Mexico 


Abstract.—A new cavernicolous species of the genus Cubacubana (Zygen- 
toma: Nicoletiidae) is described. This species is probably more distant from 
any other species of this genus described until now. 


The genus Cubacubana was originally 
described with three cavernicolous species 
from Cuba (Wygodzinsky & Hollinger 
1977), although Espinasa (1999b) has pro- 
posed that one of the species is of the genus 
Anelpistina (Silvestri, 1905). In Wygodzin- 
sky’s paper, he asserted that the Mexican 
fauna of free-living and cavernicolous Ni- 
coletiines is very rich and quite possibly 
species of Cubacubana will also be found 
on the mainland. Later another Cubacubana 
species was described from Aruba island 
(Mendes 1986) and finally a species was 
described from México, C. mexicana (Es- 
pinasa, 1991), thus fulfilling Wygodzin- 
sky’s prediction. Further exploration of 
caves in México has shown that organisms 
of the genus Cubacubana are actually abun- 
dant (Espinasa 1989). 

The new species has a series of charac- 
ters that permit us to define it as a cave 
adapted troglobite such as the overall large 
size, enlarged appendages, and surface 
crawling as opposed to occurring under 
rocks. Its presence in at least two different 
cave localities can be explained because 
they are within the same karstic area and 
probably are a single underground system 
(Diamant-Adler 1991). 


Materials and Methods 


Detailed descriptions of the caves can be 
found in the “‘Tepeyollotli’’ bulletins of the 


SMES speleological society (Diamant-Ad- 
ler 1991). Organisms collected were crawl- 
ing on the cave’s floor. They were placed 
into a vial with 96% ethanol. Dissections 
were made with the aid of a stereo micro- 
scope and the different parts of the body 
were mounted in fixed preparations with 
Hoyer’s solution. All illustrations were 
made with aid of a camera lucida attached 
to a compound microscope. 

Types were deposited in the following 
collection: LESM-DB-MEX (Laboratory of 
Ecology and Systematic of Microarthro- 
pods. Department of Biology, Faculty of 
Sciences, UNAM. México D.E). Catalog 
number: ZYG-5. 


Cubacubana asymmetrica, new species 
Figs. 1[A—G, 2A—F 


Type material.—México, Puebla State, 
Coyomeapan municipality, Tepepa, ““TP4- 
13” Cave (780 meters deep and more than 
5 kilometers long). Ex soil, 23 Dec 1990, 
L. Espinasa col. Male holotype, two male 
paratypes and five female paratypes. 

Other localities: México, Puebla State, 
Coyomeapan municipality, Huitzilatl, 
‘““Xaltégoxtl’”> Cave (60 meters deep and 
more than 2 kilometers long). Ex soil, 3 Jan 
1991, R. Espinasa col. Two males and two 
females. 

Description.—Maximum body length 
22.5 mm. Maximum length of antennae 34 


VOLUME 113, NUMBER 1 


mm, of caudal appendages 35 mm. When 
complete, antennae measure 3X length of 
body and caudal appendages 2 length of 
body. General color light yellow to white. 
Macrochaetae simple or forked. 

Head with approximately 5 + 5 macro- 
chaetae on border of insertion of antennae. 
Antennae of male with 83 preserved arti- 
cles; distal articles very slender and long, 
with internal sensory organs similar to C. 
negreai (Wygodzinsky & Hollinger, 1977). 
Basal article of antennae in males without 
projections. Pedicellus of antennae of male 
elongate as shown in Fig. 1B, with clusters 
of unicellular glands arranged in 5 groups. 
In adults, right pedicellus bigger than left 
and longer than basal article (Fig. 1B—C). 
Female basal articles of antennae simple. 
Mouthparts very elongate (Figs. 1A and 
2A); galea almost as long as lacinia (Fig. 
2A), with 2 apical conules of different 
width (Fig. 2B). Two teeth on lacinia and 
on bigger tooth’s base, an extra very small 
hyaline tooth. Labial palp long and slender, 
apical article one and a half times longer 
than wide and subtriangular (Fig. 1A). La- 
bium and Ist article of labial palp with 
macrochaetae. Mandibles with 4 macro- 
chaetae, without small pegs on the bigger 
tooth. Legs elongate; hind tibia approxi- 
mately 8—9 times longer than wide. Leg 
chaetotaxy as in C. mexicana (Espinasa, 
1991). Claws of normal size. 

Cerci of male with a very short basal ar- 
ticle, a very long 2nd one bearing numerous 
spines, followed by numerous short articles 
of simple chaetotaxy. In adults, the very 
long article is slightly curved and with 
spines, all inserted on tubercles and roughly 
of the same size and form (Fig. 1D—E). The 
spine row does not reach base of article. 
Female cerci simple. 

Thorax with short macrochaetae: 2 + 2 
on anterior border of pronotum, 3 + 3 mac- 
rochaetae on nota’s lateral borders, and | + 
1 submedian distinct macrochaetae apart 
from several setae of varied sizes on nota’s 
posterior borders. 

Urotergite X of both sexes with a small 


219 


degree of prominence and posterior part 
short in length, approximately one fifth its 
width, with a shallow emargination (Fig. 
1G), posterior angles with a few macro- 
chaetae of varied sizes, length of inner ma- 
crochaetae almost equal to distance be- 
tween them. 

Abdominal terga and sterna as in other 
members of genus. Abdominal sterna H— 
VII subdivided into coxites and sternite. 
Sterna VIII and [X of male entire. Uroster- 
num III and IV of male without modified 
coxites. Urosternum VIII of male shallowly 
emarginate on posterior margin, angles of 
emargination pointed to slightly rounded. 
Urosternum IX of male straight behind, 
without modifications (Fig. 1F). Stylets H-— 
VIII with two macrochaetae and an extra 
subapical pair. Stylets [X larger than others, 
with four macrochaetae and the extra sub- 
apical pair. Terminal spine with small teeth. 
In males and females styles IX without 
spines. 

Penis and parameres as shown in Fig. 1E 
Parameres very short, broaden slightly at 
base on inner face, and attaining only % of 
stylets IX. Surface of parameres with short 
setae, apical portion with numerous very 
short setae. Subgenital plate of female sub- 
elliptic or parabolic slightly rounded, slight- 
ly longer than wide. Ovipositor surpassing 
apices of stylets IX by a distance equal to 
% the length of stylets (Fig. 2F). Gonapo- 
physes with approximately 17 articles. 

Post-embryonic development as in Ta- 
bles 1—2, Figures 1D—E and 2C—EF Length 
of body can be obtained from the length of 
hind tibia according to the next formula: 


Length of body = (5.57) length of hind 
Gbiaate 65 3 


Etymology.—asymmetrica = Asymmet- 
ric. Makes reference to the asymmetric size 
of pedicellus in males, the right one being 
longer than the left. 

Remarks.—Cubacubana asymmetrica 
can be differentiated from other members 
of the subfamily Cubacubaninae by the fol- 
lowing characters: Species of genus Texo- 


Nw 
N 
o 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


eS 
t. set 
L | \ Pay | Lee 
Wore ell | = 
va if == 4 f |! ot = z 
ea | 
bi fs “is peste) x: eae 
! A ! ote 24 - 
Fl bi aX 
1 
! Y 
wotant 
! 
ue 
2, 
i} 
1 
1 
' 
! 
' 
Ot 
cin 
i] 
at 
i] 
L at ae 
DiI IPN es 
yan ne 
v Ase vx 
aE ection Ne 
Moris “i ic 
ae iN si 
ant Bid Viti 
! WAY 
uw TaN ES 
rig j+\ 
14 iW 
ml a ary 
ii) D aS 


1 


x 
S42 


Fig. 1. Cubacubana asymmetrica n. Sp.: A, labium with palp; B, Male. Basal portion of left antenna; C, 
Male. Basal portion of right antenna; D, Juvenile male. Basal portion of cercus with five spines; E, Adult male. 
Basal portion of cercus with seven spines; K Male. Genital area; G, Urotergum X. 


reddellia (Wygodzinsky, 1973) or Squami-_ icoletia (Paclt, 1979) has a rugged endo- 
gera (Espinasa, 1999a), have scales, which podium, absent in Cubacubana. Prostheci- 
are lacking in Cubacubana. Allonicoletia na (Silvestri, 1933), has a submentum with 
(Mendes, 1992) lacks stylets in urosternite conspicuous lateral lobes bearing numerous 
II, which are present in Cubacubana. Neon- glandular pores, absent in Cubacubana. 


VOLUME 113, NUMBER I 


Xk | 


ee | 


Pint. 


[onennnann-==] 


B 


ames 
5 
ee 


O 


Fig) 2: 


221 


| 


-1 mm. 


[cet tr ere 


Cubacubana asymmetrica n. Sp.: A, Maxilla; B, Apical portion of maxilla; C-K Post-embryonic 


development of female. Ovipositor and subgenital plate. Length of ovipositor in Table 2 as follows: C = —%, 


DS sal, E. = 0,.Fi— hh. 


Most species of Anelpistina, have articulat- 
ed submedian appendages in urosternite IV 
of males, which are absent in Cubacubana. 
From the two species of Anelpistina that 
lack such appendages; A. decui (Wygodzin- 
sky & Hollinger, 1977) and A. inappendi- 
cata (Espinasa, 1999b), and from the de- 
scribed species of Cubacubana; C. ramosi 
(Wygodzinsky, 1959), C. negreai (Wygod- 
zinsky & Hollinger, 1977), C. arubana 
(Mendes, 1986), and C. mexicana (Espina- 


sa, 1991), C. asymmetrica differs by the 
posterior part of urotergite X, which is short 
in length (one fifth the width of the uroter- 
gite), while in the other species it is longer 
(approximately one half to one third). 
Males are also differentiated by long pedi- 
cellus (twice or more long than wide) and 
also by the asymmetry of adults, in which 
right pedicellus is bigger than left. 
Cubacubana asymmetrica can be further 
differentiated from C. mexicana, the only 


N 


N 


Table 1.—Post-embryonic development of male Cubacubana asymmetrica. 


MALES: 


(Fig. 1D-E) 


Spines in stylets 
IX 


Spines in Cerci 


Parameres vs. 
stylets IX 


Sensory cones 
urosternum IX 


Emargination 
urosternum VIII 


Spines in III 
urosternum 


Length both 
pedicellus 


Length 
Hind 
tibia 


3/4 No 


1/2 


No 1/3 


Yes 


Yes 


Unequal No NOES No Small 


Equal 


mm 


Cave of -“iR4-137" 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


+++ +4 


++ +4 +4 


1.50 
2.74 
Sa10 
Sn8 


oo 


Cave of ‘*Xaltégoxtl’’ 


2.74 
2.88 


a 


Table 2.—Post-embryonic development of female 
Cubacubana asymmetrica. 


FEMALES: 
Ovipositor surpassing apex stylets IX 
: (Fig. 2C—F) 
Lenethvhind. ‘= Se ees 
tibia mm = Sy? =i (0) 1/4 


Caver or, “TP4213~ 


1.74 45 

3.45 ae 

3.45 ae 

3.49 il 
Cave of *‘Xaltégoxtl’’ 

1.76 == 

2.89 = 

3.16 at 


other Mexican Cubacubana species de- 
scribed, by its longer antenna (three times 
the length of the body instead of one and a 
half), caudal appendages (two times the 
length of the body instead of slightly more 
than one), maxillary palp (distal article 11 
or more times longer than wide instead of 
less than ten) and legs (hind tibia approxi- 
mately eight to nine times longer than wide 
instead of six). Furthermore males do not 
have spines or cones on urosternum III and 
IX or on styles IX. Gonapophyses of fe- 
males have fewer articles (approximately 
17 instead of 22). 

Cubacubana asymmetrica shares char- 
acteristics present in other genera, such as 
the curvature of cerci of Prosthecina ad- 
dititia (Wygodzinsky, 1951), or some type 
of asymmetry in the pedicellus as in Cole- 
tinia subterranea (Silvestri, 1902) and Col- 
etinia asymetrica (Bach de Roca et al., 
1985). These characters are probably the re- 
sult of convergent evolution. 


Acknowledgments 


I thank Dr. José G. Palacios-Vargas, di- 
rector of “‘Laboratorio de Ecologia y siste- 
matica de Microartr6podos”’, where most of 
the descriptive work was done and the di- 
rector of CEAMISH-UAEM,, Dr. Oscar Do- 
rado, for support this research. Thanks are 
also due to Dr. Luis EK Mendes and Monika 
Baker for reviewing the manuscript. 


VOLUME 113, NUMBER 1 


Literature Cited 


Bach de Roca, C., L. E Mendes, & M. Gaju Ricart. 
1985. Sur une nouvelle espéce et une nouvelle 
citation de Nicoletiinae de Sierra Morena (Cor- 
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Diamant-Adler, R. 1991. Participacién de la S.M.E.S. 
en MEXPE IV, expedicion de la S.Q.S. a la Si- 
erra Negra.—Tepeyollotli: Gaceta de la Socie- 
dad Mexicana de Exploraciones Subterraneas 5: 
22-28. 

Espinasa, L. 1989. Bioespeleologia.—Tepeyolloth: 
Gaceta de la Sociedad Mexicana de Explora- 
ciones Subterraneas 4:45—46. 

. 1991. Descripci6n de una nueva especie del 

género Cubacubana (ZYGENTOMA: Nicoleti- 

idae) y registro del género para América Con- 

tinental.—Folia Entomolégica Mexicana 82:5— 

16. 

. 1999a. A new genus of the subfamily Cuba- 

cubaninae (Insecta: Zygentoma: Nicoletiidae) 

from a Mexican cave.—Proceedings of the Bi- 

ological Society of Washington 112:52-58. 

. 1999b. Two new species of the genus Anel- 
pistina (Insecta: Zygentoma: Nicoletiidae) from 
Mexican caves, with redescription of the ge- 
nus.—Proceedings of the Biological Society of 
Washington 112(1):59-69. 

Mendes, L. FE 1986. Nouvelles données sur le Zygen- 
toma (Insecta) de 1’ Amerique centrale et du Me- 
xique.—Bulletin du Muséum national d’ Histoire 
naturelle,. Paris (4) 8 (A) (2):333-—342. 

. 1992. Novos dados sobre os tisanuros (Micro- 


Ze5 


coryphia e Zygentoma) da América do Norte.— 
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Paclt, J. 1979. Neue Beitrage zur Kenntnis der Apter- 
ygoten-Sammlung des Zoologischen Instituts 
und Zoologischen Museums der Universitat 
Hamburg. VI Weitere Doppel- und Borstensch- 
wanze (Diplura: Campodeidae. Thysanura: Lep- 
ismatidae und Nicoletiidae).—Entomologische 
Mitteilungen aus dem zoologischen Museum 
Hamburg 6(105):221-—228. 

Silvestri, EK 1902. Materiali per lo studio dei Tisanuri. 
III. Nuove specie di Nicoletia.—Bolletino della 
Societa entomologica italiana 33:223-—227. 

. 1905. Materiali per lo studio dei Tisanuri. VI. 

Tre nuovo sottogenero.—Redia (Firenze) 2: 

111-120. 

. 1933. Nuovo contributo alla conoscenza dei 
tisanuri de Mesico.—Bolletino del Laboratorio 
di Zoologia general e agraria di Portici 27:127— 
144. 

Wygodzinsky, P. 1951. Apuntes sobre “‘Thysanura”’ 
americanas.—Acta zoologica Lilloana 11:435-— 
458. 

. 1959. Contribution to the knowledge of the 

“Thysanura” and ‘“‘Machilidae’’ (Insecta).— 

Revista Brasileira de Biologia 19(4):441—457. 

. 1973. Description of a new genus of cave 

Thysanura from Texas (Nicoletiidae. Thysanu- 

ra. Insecta).—American Museum Novitates 

2518:1-8. 

, & A. M. Hollinger. 1977. A Study of Nico- 

letiidae from Cuba (Thysanura).—Resultats des 

Expéditions Bioespeleologiques Cubano-rou- 

maines 4 Cuba 2:313-324. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


113(1):224—237. 2000. 


Three new species of bathyal cidaroids (Echinodermata: Echinoidea) 
from the Antarctic region 


Rich Mooi, Bruno David, F Julian Fell, and Thérése Choné 


(RM) Department of Invertebrate Zoology & Geology, California Academy of Sciences, 
Golden Gate Park, San Francisco, California 94118-4599; 
(BD & TC) UMR CNRS 5561, Université de Bourgogne, 6, bd. Gabriel, F-21000, Dijon, France; 
(FJF) Box 222, Errington, British Columbia, Canada VOR 1V0O 


Abstract.—Two new species of Aporocidaris, A. eltaniana and A. usarpi, 
and one new species of Notocidaris, N. lanceolata, are described from material 
collected during United States Antarctic Research Program expeditions in the 
1960’s. All three species occur in the bathyal zone near or south of the Sub- 
antarctic Convergence. Gonopore sizes and peristomial morphology suggest 
that these species are sexually dimorphic and that the females can brood direct- 
developing young. However, only one of the Aporocidaris species was found 
to harbor embryos on the peristomial membrane. 


In the mid- to late 1960’s, Dr. Richard H. 
Chesher, at the Museum of Comparative 
Zoology at Harvard (MCZ), studied echi- 
noid material collected under the auspices 
of the United States Antarctic Research 
Program (USARP). The USARP, which be- 
gan in 1955, included trawling and dredg- 
ing expeditions by ships in the vicinity of 
Antarctica (Fig. 1). Material collected by 
the U.S. National Science Foundation Re- 
search Vessel R/V Eltanin is of particular 
importance to this paper. Dr. Chesher iden- 
tified numerous USARP echinoids, in the 
process discovering several new species. In 
some cases, he proposed manuscript names 
and indicated type specimens for new spe- 
cies (for example, see David & Mooi 1990), 
but he was unable to continue systematic 
work after 1967, thereby leaving these 
names and manuscripts unpublished. 
Chesher’s cidaroid research was incorporat- 
ed into a dissertation (Fell 1976), and two 
of the species described below were cov- 
ered in this work. As a result, several pu- 
tative types for which no names have been 
published are deposited in the National Mu- 
seum of Natural History (NMNH)-the ma- 
jor repository of echinoid collecting during 


the USARP, and in the MCZ. Here, we pub- 
lish these names for the first time. 

Antarctic cidaroids are represented by 
approximately 20 species in 5 genera, and 
all belong to the subfamily Ctenocidarinae. 
With one or two exceptions, the ctenoci- 
darines are restricted to the Antarctic and 
Subantarctic regions and are among the 
most diverse of the Antarctic echinoids. Al- 
though their taxonomy was hugely ad- 
vanced in Fell (1976), an exact count of 
valid taxa is difficult, given the uncertain 
specific and subspecific status applied to the 
variants. We are currently engaged in a da- 
tabasing project that will result in a com- 
prehensive overview of all Antarctic echi- 
noids, including cidaroids, and it was this 
review that prompted the work on the taxa 
described herein. 

Much of the Antarctic cidaroid fauna still 
requires revision in a modern context. For 
example, the only recent attempt (Smith & 
Wright 1988) to resolve phylogenetic rela- 
tionships among any of the cidaroids con- 
centrated on Cretaceous forms, and did not 
deal with Antarctic taxa. Phylogenetic re- 
lationships among the different genera and 
species are still speculative and will remain 


VOLUME 113, NUMBER 1 


- 4 
90° 


New Zealand 0 


180° (0 


Fip- 1. 


TOT 


225 


@ Aporocidaris eltaniana 

A Aporocidaris usarpi 

¥% Notocidaris lanceolata 
7 NUMBER OF 


DEPTH STATIONS 
, (m) » ea SAR ae | 


\ (0) 
2: 
a L 
: 
\ 
Sr 
1000 @ 
90° 
\ 
\ 
\ 
2000 
vt 
S 
xv 


4000 


Collecting localities and recorded depths (graph at right) for Aporocidaris eltaniana new species, A. 


usarpi new species, and Notocidaris lanceolata new species. Number of stations refers to the number of R/V 
Eltanin stations that for a given species fall within the depth range indicated by the height of the shaded box. 


so until a more complete knowledge of the 
taxonomy is established. The lack of con- 
sistent, well-delineated features as well as 
detailed analyses of morphological varia- 
tion makes the Antarctic cidaroids a partic- 
ularly difficult group, as evidenced by the 
almost unusable keys provided by previous 
major revisions such as Mortensen (1928). 
In addition, material is often rare, and sev- 
eral taxa are represented only by juveniles, 
or have not been collected since they were 
first described from single or very few in- 
dividuals. Therefore, determination of ter- 
minal taxa to be used in phylogenetic anal- 
ysis is in its infancy, and any effort to place 
the taxa described here in an evolutionary 
context is premature. 

Jackson (1912) and Smith (1984) provid- 
ed excellent comparative overviews of ci- 


daroid morphology. Cidaroids are very easy 
to separate from other types of regular ur- 
chins. Each interambulacral plate bears a 
single, large, perforate primary tubercle 
supporting a primary spine that is strongly 
differentiated from the secondary spines 
covering the rest of the test. Primaries are 
many times larger than secondaries, and 
much more robust. The shafts of adult ci- 
daroid primary spines are unique not only 
among all spines on a cidaroid, but also 
among echinoids in general in completely 
lacking an epithelial layer. These spines are 
often invested with various epizoans such 
as sponges, serpulids, bryozoans, and small 
mollusks. Each primary is encircled by a 
palisade of much shorter secondary spines 
called scrobiculars that can be appressed to 
the base of the primary spine. 


226 


In contrast to other regular echinoids, ci- 
daroid ambulacra are very narrow relative 
to the interambulacra. The ambulacral 
plates are arranged in two simple columns, 
each of which bears a single tube foot pore 
(usually bipartite), and one to several small 
secondary spines. The ambulacral series 
continue to the mouth on the peristomial 
membrane, and the number of peristomial 
ambulacrals can be helpful in discriminat- 
ing certain Antarctic taxa. The perignathic 
girdle consists of interambulacral processes 
called apophyses. The Aristotle’s lantern of 
Antarctic species is typical for cidaroids, 
and does not vary enough to form the 
source of characters that could be used to 
distinguish the taxa. 

Although it is relatively easy to recog- 
nize a cidaroid, a confounding feature of 
the taxonomy within the group is conser- 
vatism in plate architecture and overall 
form of the test. However, the cidaroids 
more than compensate for this conservatism 
in diversity of primary spine morphology. 
The taxa described in this paper exemplify 
the fact that many cidaroid species cannot 
be identified without information on the pri- 
mary spines. Both scrobicular and non- 
scrobicular secondary spines can also be 
useful in diagnosing Antarctic cidaroids. 

Cidaroids are epibenthic and inhabit var- 
ious environments from sandy and stony 
bottoms in littoral zones to gravels and 
muds of sub-littoral regions. They are also 
common in the deep-sea on muddy bottoms 
of bathyal slopes or abyssal plains to 5000 
meters, and they constitute a significant part 
of the benthic community at these depths. 
Many, if not all, of the Antarctic cidaroids 
are direct developers, as suggested by the 
fact that in almost every species, a subset 
of adults presumed to be females have 
greatly enlarged gonopores. Fell (1976) and 
Lockhart et al. (1994) recorded that many 
species also brood their young in and 
around “‘marsupia”’ created by the more or 
less sunken peristome, a feature of which 
we make special note in the following de- 
scriptions. Because the presence of brooded 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


young is correlated with enlarged gono- 
pores, and because large gonopores are 
known to signify females in other echi- 
noids, we also use the presence of enlarged 
gonopores and marsupia to infer gender in 
adult specimens. 

Upon occasion, primary spines of Ant- 
arctic cidaroids also support a very poorly 
understood parasite, Echinophyces mirabi- 
lis Mortensen & Rosenvinge, 1909. The oc- 
currence of this parasite is correlated with 
some modifications of the infected echi- 
noid’s morphology and a delay in the ap- 
pearance and change in position of the gon- 
opores. The mechanism by which a parasite 
of a more or less biologically inert external 
feature such as the cidaroid spine can affect 
the expression of gonads and gonopores is 
totally unknown and clearly deserves fur- 
ther study. It is surprising that virtually no 
research beyond the original description 
and that of Mortensen & Rosenvinge 
(1910) has been done on this unusual phe- 
nomenon. Fell (1976), Jangoux (1987), and 
Pearse & Cameron (1991) have summa- 
rized what few data exist, finding that 
Echinophyces is found only in Ctenocidaris 
Mortensen, 1910 and Rhynchocidaris Mor- 
tensen, 1909. We report on the results of 
our as yet unsuccessful search for evidence 
of this parasite in the new species, in the 
hope that future studies might be able to use 
even these negative data. 


Order Cidaroida L. Agassiz, 1835 
Family Cidaridae Gray, 1825 
Genus Aporocidaris A. Agassiz & Clark, 
1907 


Diagnosis.—Ctenocidarine cidarids with a 
large apical system which can be as much 
as 75% of the horizontal diameter of the 
test. Apical system often significantly 
domed or convex. Mid-interambulacral re- 
gions usually with a slightly depressed, na- 
ked interradial suture. 

Remarks.—Mortensen (1928) considered 
four nominal species in the genus Aporo- 
cidaris, but he emphasized the “‘exceeding- 


VOLUME 113, NUMBER 1 


ly slight” differences between 3 of them: 
A. antarctica Mortensen, 1909, A. fragilis 
A. Agassiz & Clark, 1907, and A. milleri 
(A. Agassiz, 1898). Fell (1976) proposed 
synonymy of these 3 species under A. mil- 
leri but suggested a new species, A. eltan- 
iana. In an unpublished manuscript, Chesh- 
er recognized yet another species, A. usarpi. 
We introduce the latter names to the pub- 
lished literature for the first time. 


Aporocidaris eltaniana, new species 
Fig. 2 


Aporocidaris eltaniana.—ex Fell, 1976: 
211, figs. 7, 8j-k, name used in unpub- 
lished thesis. 


Diagnosis.—Apical system as much as 
68% of test diameter. Aboral primary tu- 
bercles large and markedly transversely 
oval at ambitus. Aboral primary spines cy- 
lindrical, 1 to 1.5 times test diameter, most- 
ly smooth or with small bumps arranged in 
longitudinal rows; tip blunt, or slightly con- 
cave. Oral primary spines with conspicuous 
thorns along lateral edges of shaft. Pre- 
served specimens dark brown, primary 
spines whitish, secondary spines beige to 
light brown. Other characters as for Apo- 
rocidaris. 

Description.—The test is about half the 
height of the horizontal diameter, but the 
height of the apical system is quite variable 
and can add as much as 20% to the overall 
height of the coronal part of the test. The 
largest recorded size of the specimens ex- 
amined is about 45 mm in horizontal di- 
ameter of the test. The holotype is 29.3 mm 
in diameter, 16.9 mm high, has an apical 
system diameter of 15.6 mm and a peri- 
stome diameter of 12.5 mm. 

The ambulacra are almost straight (Fig. 
2A, B), particularly on the oral surface, and 
their tuberculation is of the usual Aporoci- 
daris pattern (Fig. 2E). There is no naked 
area along the perradial suture. The inner 
and outer pore of each podial pore pair are 
divided by a well-developed ‘“‘bridge”’ of 
stereom (Fig. 2E). 


227 


The interambulacra have about one more 
plate in each series than A. milleri of similar 
size. The crowding of the plates into the 
relatively flat corona, and the size of the 
tubercles between the ambitus and the peri- 
stome cause the tubercles to be compressed 
into horizontally oriented ovals (Fig. 2B). 
The scrobicular tubercles are more than 
twice the diameter of the other secondary 
tubercles, and surround each primary tu- 
bercle except where the oval outlines of the 
primaries touch (Fig. 2D). The interradial 
suture 1S conspicuously sunken in larger 
specimens (Fig. 2D). 

The apical system is large (about 60% of 
the horizontal diameter on average), and al- 
most flat or only slightly domed in most 
specimens. However, a few exceptional in- 
dividuals have a strongly arched apical re- 
gion that can account for over a fifth of the 
total height of the test. The genital plates 
are large, with scattered secondary tuber- 
cles. In males, the gonopores are small and 
close to the outer edges of the genital plates 
(Fig. 2A). In several of the specimens, the 
gonopores are large, forming notches that 
in extreme cases can extend part way along 
the interradial suture separating the inter- 
ambulacral plates adjacent to the genital. 
Like Fell (1976), we have assumed these 
specimens to be females. The gonopores 
first appear in specimens as small as 16 mm 
in test diameter. Ocular plates are only 
about a third the area of the genitals, and 
the ocular pores are surrounded by a cir- 
cular ridge that seems to be most pro- 
nounced in the males (Fig. 2A). Tubercu- 
lation is not dense, and restricted to the cen- 
tral parts of the plates over the majority of 
the apex (Fig. 2A). 

The peristome is just over 55% of the test 
diameter in small specimens, but becomes 
smaller in larger individuals (approximately 
40% of the test diameter), relative to test 
diameter. The peristome itself is somewhat 
sunken around its periphery, notably in the 
interambulacral regions. Seven to 8 pairs of 
ambulacral plates continue onto the peri- 
stome in larger specimens, and there are 


228 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


==eee' 
(NTR 

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Wipe 


f 
fl 


f iit A\\\ | @yY: 
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(A 


a 3 

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S Zp 
SSS 


(y 


Aboral plate pattern Oral plate pattern eh 
Aboral }!1 
primary (y 


Scrobicular j 
3 secondary 
Side view of holotype spine 


Non-scrobicular } 
secondary § 


Fig. 2. Aporocidaris eltaniana new species. C drawn from holotype (NMNH E48122); A, B, D—I drawn 
from paratypes in lot NMNH E11161. For A and B: mouth, anus, and gonopores in solid black; podial pores 
omitted; secondary tubercles shown only for single genital, ocular, and periproctal plates, but omitted elsewhere, 
including coronal plates. 


VOLUME 113, NUMBER | 


only 2 or 3 small scales in each interradial 
region (Fig. 2B). 

The longest aboral primary spines found 
(on the almost intact specimens in NMNH 
E11161) are approximately 1.5 times the 
test diameter and rod-like with blunt or 
slightly concave tips and large milled rings 
(Fig. 2F). The surface of the shaft is smooth 
proximally, but bears small, longitudinally 
aligned, blunt spinules for more than 80% 
of its length out to the tip (Fig. 2F). The 
flat areas between the longitudinal rows are 
covered with a variously developed fibrous 
and anastomosing calcareous meshwork. 
The oral primaries are strongly curved to- 
wards the peristome, and bear well devel- 
oped thorns that project laterally, but be- 
come smaller towards the tip of the spine. 
There is often another row of shorter thorns 
along the distal part of the shaft (Fig. 2G). 
The more or less cylindrical scrobicular 
spines are blunt, but not club-shaped (Fig. 
2H). The non-scrobicular secondaries are 
much shorter than the scrobicular spines 
and sometimes slightly curved near the al- 
most imperceptibly swollen tip (Fig. 21). 

The small globiferous pedicellariae are 
rare, but appear simply to be half-sized ver- 
sions of the larger type. The stem of a glob- 
iferous pedicellaria is thick, and straight. 
Neither the valves nor stem are otherwise 
distinctive among Aporocidaris species 
(Mortensen 1928). 

Types.—Holotype NMNH E48122, ex- 
tracted from NMNH E11161, R/V Eltanin 
Cruise 6, Station 432 (Fig. 2C). Paratypes 
NMNH E11161 (3 dry specimens, one dis- 
sected lantern), E11188 (17 dry specimens, 
including 3 less than 10 mm in horizontal 
test diameter), E11212 (dry spines only), 
E11290 (more than 60 dry specimens), 
E14597 (2 specimens greater than 28 mm 
in test diameter, the largest of which is a 
female with more than 12 young urchins 
brooded on the peristome, 1 specimen 14.3 
mm in horizontal test diameter, and 16 
specimens less than 10 mm in test diameter, 
all in alcohol), MCZ 8406 (1 dry speci- 
men). The holotype was chosen because it 


229 


retained significant spine cover (Fig. 2C). 
The paratypes in NMNH E11161 were used 
to make the drawings of plate architecture 
so as not to disturb what spination remains 
on the holotype. 

Etymology.—Named for U.S. National 
Science Foundation R/V Eltanin. The spe- 
cies name was first coined in an unpub- 
lished doctoral thesis by Fell (1976), and 
we introduce the name here in recognition 
of that first usage. 

Distribution.—A. eltaniana is known 
only from the type series, which was col- 
lected from two R/V Eltanin stations 
(Cruise 4, Station 138; Cruise 6, Station 
432) at two distinct localities, both in the 
region of Livingston Island in the South 
Shetland Islands, between 884 and 1437 m 
depth (Fig. 1). 

Biology.—The gonopores in females 
may reach 1.8 mm in diameter. One female 
(NMNH E14597), 43.4 mm in horizontal 
test diameter, had more than a dozen juve- 
nile urchins in the marsupium along the 
sunken edge of the peristomial margin. An 
exact count of the juveniles was impossible 
without damaging the adult specimen. Most 
of the young were obscured by oral primary 
and secondary spines that folded over to 
screen the deepest parts of the marsupium 
in the interambulacral areas proximal to the 
edge of the coronal plates. One of the larg- 
est of the juveniles was 2.2 mm in horizon- 
tal test diameter, and all of the brooded 
young possessed primary spines, the lon- 
gest of which were approximately 2.0 mm 
long. These observations suggest that ju- 
veniles of A. eltaniana develop directly 
from relatively large eggs which are moved 
into, and then develop in a peristomial mar- 
supium. None of the specimens appear to 
be infected by Echinophyces. Nothing is 
known of the preferred bottom type, but the 
gut contents of one specimen are made up 
of light gray silt containing small fragments 
of thin mollusk shells. 

Remarks.—The greatly enlarged apical 
system and the naked, sunken area along 
the interradius indicate that the new species 


230 


is an Aporocidaris. A. eltaniana is most 
similar to A. milleri, but is distinct from all 
Aporocidaris in possessing strongly devel- 
oped thorns on the oral primary spines. The 
dramatic difference between the relatively 
smooth, cylindrical aboral primaries and the 
thorny oral primaries is unmatched in the 
genus, if not within the Antarctic cidaroids 
in general. 


Aporocidaris usarpi, new species 
Fig. 3 


Diagnosis.—Apical system as much as 
70% of test diameter in adults. Aboral pri- 
mary spines cylindrical, 1 to 1.5 times test 
diameter, with small, irregularly distributed 
or only slightly aligned bumps distally. 
Neck of aboral primaries glossy, porcella- 
neous, and markedly swollen in larger spec- 
imens. Oral primary spines flattened with 
lateral, diametrically opposed, serrated 
keels that are widest just distal to neck and 
taper towards spine tip. Scrobicular spines 
fine, small, closely spaced, and not strongly 
differentiated from the non-scrobicular 
spines. Both types of secondary spines 
slightly club-shaped. Preserved specimens 
beige, primary spines white, secondary 
spines light beige. Other characters as for 
Aporocidaris. 

Description.—The test is only moderate- 
ly flattened. The height of small specimens 
is Just over 50% of the horizontal diameter, 
but in larger specimens, the height is greater 
(up to 75% of the horizontal diameter), al- 
most 15% of which can be in the form of 
the domed apical system (Fig. 3C). The 
largest specimen is just over 40 mm in hor- 
izontal diameter. The holotype is 35.2 mm 
in diameter, 23.4 mm high, has an apical 
system diameter of 23.0 mm and a peri- 
stome diameter of 15.2 mm. 

The ambulacra are almost straight (Fig. 
3A, B), particularly on the oral surface, and 
their tuberculation is of the usual Aporoci- 
daris pattern, with smaller secondary tuber- 
cles adjacent to the perradial suture, and 
Slightly larger ones just distal to those (Fig. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


3E). There is no naked region along the per- 
radius. In the holotype and the largest of the 
paratypes (NMNH E11059), the “bridge” 
between the pore pairs on each ambulacral 
plate is incomplete so that the pores are con- 
nected via a constricted passage between the 
pores (Fig. 3E). This condition is rare, if not 
unique among Antarctic cidaroids. 

The interambulacral plates are high. Ab- 
orally, the primary tubercles do not occupy 
the entire height of the plate so that the ab- 
oral primary tubercles retain their round 
outlines. However, adjacent to the peri- 
stome, the tubercles are more crowded, and 
tend to compress slightly into transverse 
ovals (Fig. 3B). The scrobicular tubercles 
are not strongly differentiated in size from 
the non-scrobicular secondaries, which are 
small and closely spaced everywhere out- 
side the primary tubercle except along the 
interradius, where tubercles are lacking 
(Fig. 3D). This distinct naked area is adja- 
cent to each interradial suture and occupied 
by a shallow furrow that, in larger speci- 
mens, sends branches circumferentially 
along sutures separating plates in each of 
the two interambulacral columns (Fig. 3D). 

The apical system is large, ranging from 
just over 50% of the test diameter in juve- 
niles to over 70% of the diameter in the 
largest adults. In the juveniles, the apical 
region tends to be almost flat. It is distinctly 
domed in adults, with considerable irregular 
inflation of the centers of some of the larger 
apical plates (particularly the genitals), es- 
pecially in the specimen identified as a fe- 
male. The genital plates are large relative 
to the other plates in the apical system. In 
the largest paratype, which appears to be a 
female, the gonopores can be as much as 
2.4 mm in diameter and in some cases oc- 
cupy shallow clefts along the interradial su- 
ture (Fig. 3A). The gonopores are present 
in the holotype, which appears to be a male, 
but are not evident in the larger of the 2 
small paratypes in NMNH E11059, which 
is 13.9 mm in diameter. Ocular plates are 
only about a quarter the area of the genitals, 
and the ocular pores are surrounded by a 


VOLUME 113, NUMBER 1 


prominent circular ridge (Fig. 3A). The tu- 
berculation on both oculars and genitals is 
fine and dense, and tends to cover the entire 
plate surface almost to the suture (Fig. 3A). 

The peristome is sunken around its pe- 
riphery, and in the region of half the test 
diameter. Each perradial region bears 8 or 
9 plates in each ambulacral column, and 
there are two relatively large scales in each 
interradial portion of the membrane (Fig. 
3D). 

The longest aboral primary spines are ap- 
proximately 1 to 1.5 times the horizontal 
diameter of the test, although even the best 
examples are worn or broken so that the 
nature of the tips cannot be ascertained. 
Distal to the neck, the shaft is cylindrical 
and rod-like, and invested with irregularly 
spaced to only partially aligned, short, blunt 
spinules or bumps that arise from _ the 
smooth, porcellaneous surface. The neck is 
most peculiar in that it is greatly swollen in 
primary spines of larger specimens, and 
completely unadorned with spinules. The 
surface of the swelling is smooth, porcel- 
laneous and glossy, and increases the di- 
ameter of the spine by a factor of 2 at its 
widest point (Fig. 3F). In juvenile speci- 
mens, the neck is also porcellaneous but 
only slightly swollen, suggesting that the 
vase-like shape of the swelling becomes 
more prominent with age. The oral primary 
spines are only slightly curved towards the 
peristome, and bear diametrically opposing 
flanges or keels along the lateral sides of 
the shaft. The edges of these keels are finely 
serrated, and the keels themselves are wid- 
est proximally, tapering towards the rela- 
tively blunt spine tip to give the entire spine 
a dagger-like appearance (Fig. 3G). The 
surface of the oral primaries is smooth and 
shiny, but not as glossy as the aboral pri- 
maries. The almost cylindrical scrobicular 
spines are blunt and very slightly swollen 
towards the tip (Fig. 3H). Scrobicular sec- 
ondaries are only slightly longer than the 
non-scrobicular secondary spines (Fig. 3I). 
Both types of secondaries are club-shaped. 

The valves of the large globiferous ped- 


2351 


icellariae are less than 1 mm in length, and 
very similar to those of other Aporocidaris 
(Mortensen 1928). The smaller globiferous 
were not detected. 

Types.—Holotype NMNH E11134, R/V 
Eltanin Cruise 14, Station 1212 (Fig. 3C, 
F-I). Paratypes NMNH E11059 (3 dry 
specimens, one dissected lantern, and one 
almost perfect, loose spine), E14603 (1 
specimen in alcohol). The holotype was 
chosen because it retained some spines 
(Fig. 3C). Plate architecture was drawn 
from one of the largest of the paratypes in 
NMNH E11059 to avoid further damage to 
the holotype. 

Etymology.—The name “‘A. usarpi’’ was 
originally used by Dr. Richard Chesher in 
his identifications of USARP material now 
housed the NMNH (see above), and it is 
clear from notes placed with that material 
that he intended to publish the species in a 
manuscript that never saw press. We retain 
Chesher’s suggested name in honor of his 
recognition of the distinctiveness of this 
taxon. 

Distribution.—A. usarpi is known only 
from a single R/V Eltanin locality (Cruise 
14, station 1212) from a mid-ocean point 
far to the southeast of New Zealand and 
north of the Ross Sea (Fig. 1) at a depth of 
between 3678 and 3935 m. 

Biology.—Nothing is known of the hab- 
itat preferences of A. usarpi, and appropri- 
ate specimens could not be sacrificed for 
gut content observations. There is some 
sexual dimorphism. The putative female 
has greatly enlarged gonopores (up to 2.4 
mm in diameter) relative to the putative 
male, suggesting direct development in this 
species. None of the specimens had young 
retained on the test, although the peristo- 
mial edge is slightly sunken in such a way 
as to suggest the same type of marsupia 
seen in other species bearing young on the 
peristome. None of the specimens show ob- 
vious signs of Echinophyces infection, but 
the paucity and condition of the material 
prevent us from ruling out the occurrence 
of the parasite in A. usarpi. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


[SN 


Aboral plate fr C7 ‘ 
pattern by i ‘ ES 6 © 
0° et e 7 NN \ 


EZ @y 


ay 


rd LE 
: [Pr bo% 
((uauatte 


1D) 
ies 


ONS 
ree 
S 


o \ fe 
Side view of G a H primnanyt 
holotype Oral ("8 spine}: 
< primary (. 23 Scrobicular 
pe spine(® #4 secondary 
‘} a spine 


Non- 
scrobicular 
secondary 

spine 


Ambulacrum at ambitus 


Fig. 3. Aporocidaris usarpi new species. C, F—I drawn from holotype (NMNH E11134); A, B, D, E drawn 
from paratypes in lot NMNH E11059. Conventions for A and B as in Fig. 2. 


VOLUME 113, NUMBER 1 


Remarks.—The greatly enlarged apical 
system and the naked, sunken area along 
the interradius indicate that the new species 
is an Aporocidaris. The species is readily 
distinguished from all other Aporocidaris 
by the peculiar swelling of the neck in the 
primary spines, and the extreme glossiness 
of both this swelling, and the shaft of the 
spines between the distal spinules. The ab- 
sence of stereom bridges separating the 
members of the podial pore pairs has not 
been previously reported for any ctenoci- 
darine taxa, let alone other Aporocidaris. 
Both the holotype and the largest of the 
paratypes exhibit this feature, but the two 
smallest specimens (below 14 mm horizon- 
tal diameter) have clearly divided pores, 
suggesting that the unification of the pores 
progresses with ontogeny, and is a derived 
feature of adult A. usarpi. 


Genus Notocidaris Mortensen, 1909 


Diagnosis.—Oral primary spines spear- 
or dagger-shaped. Aboral primaries rod-like 
and cylindrical to flattened and spatulate, 
with large lateral expansions distally. 
Spines smooth or spiny, but when present, 
spinules restricted to proximal part of shaft. 
Proximal surface of spines occasionally 
with coating of anastomosing hairs. Scrob- 
icular secondary spines simple and slender 
(but not as fine or densely distributed as in 
Aporocidaris), not distinct in shape from 
other secondary spines. Interradial sutures 
naked, but not sunken. Apical system about 
60% of horizontal diameter. 

Remarks.—Mortensen (1928) listed 4 
species in the genus: N. gaussensis Morten- 
sen, 1909, N. hastata Mortensen, 1909, N. 
mortenseni (Koehler, 1900), and N. platya- 
cantha (H. L. Clark, 1925). Another spe- 
cies, N. remigera Mortensen, 1950 was de- 
scribed after Mortensen’s monograph of the 
cidaroids was published. We do not consid- 
er the entity N. platyacantha var. contracta 
Koehler, 1926 to be a valid taxon. There- 
fore, we attribute 5 full species to Notoci- 
daris, all from Antarctic and Subantarctic 


233 


waters, and all distinguished by features of 
the aboral primary spines. It should be not- 
ed that the range of variation recorded for 
each species can be extremely large and 
spine characteristics can overlap for species 
such as N. remigera and N. mortenseni. 
Therefore, biogeographic and bathymetric 
data, as well as other parts of the descrip- 
tions must also be used to make positive 
determinations. 


Notocidaris lanceolata, new species 
Fig. 4 


Notocidaris lanceolata.—ex Fell, 1976: 
195; figs. 7, 8f-g, name used in unpub- 
lished thesis 


Diagnosis.—Aboral primary spines 
coarsely thorned at their base, with thorns 
arranged irregularly, and becoming greatly 
reduced in size and aligned towards tip. 
Spines keeled in two orthogonal planes in 
cross section, yielding lance-like appear- 
ance, spine tapering slightly towards rela- 
tively blunt tip. Longest spines about 2 
times horizontal diameter of test in most 
specimens. Apical system varying from flat 
to arched. Test of preserved specimens 
beige to ochre, primary spines beige with 
shiny purplish-beige neck, secondary spines 
beige. Other characters as for Notocidaris. 

Description.—The largest recorded hori- 
zontal test diameter is 36 mm. The height 
is between 50% and 60% of the test diam- 
eter. The holotype is 32.5 mm in horizontal 
diameter and 19.4 mm high. 

The ambulacra are straight on the oral 
surface, with some slight sinuousness ab- 
orally (Fig. 4A, B). There are one or two 
secondary tubercles perradial to the podial 
pores, and two or three much smaller sec- 
ondary tubercles next to the perradial suture 
(Fig. 4E). There is no naked region along 
the perradius. The inner and outer pore of 
each podial pore pair are divided by a well- 
developed “‘bridge’’ of stereom (Fig. 4E). 

The aboral interambulacral plates are 
high and the primary tubercles are circular. 
Orally, the plates are not as high and the 


234 


tubercles are compressed into transverse 
ovals (Fig. 4B). There are 7 or 8 plates in 
each column in a specimen about 50 mm in 
diameter. The scrobicular tubercles are in 
general only slightly differentiated from the 
non-scrobicular secondaries, which are 
abundant and closely spaced except along 
the interradial suture, where tubercles are 
lacking (Fig. 4D). There is no sunken area 
along the interradial suture. 

The apical system is usually about 60% 
of the test diameter in larger specimens, and 
can range from being flat, as in the holo- 
type, to domed. The gonopores are restrict- 
ed to the genital plates and do not signifi- 
cantly invade the adjoining interambulacra. 
The females have enlarged gonopores. The 
oculars are large and circumferentially 
elongated so that they are much wider than 
they are high, and about half the surface 
area of the genital plates (Fig. 4A). The oc- 
ular pore is encircled by a prominent ridge 
and the tuberculation is fine and evenly dis- 
tributed across each plate in the apical sys- 
tem except directly adjacent to the sutures 
(Fig. 4A). 

The peristome is only slightly sunken at 
its periphery, and approximately 45% of the 
test diameter. There are 7 or 8 plates in each 
ambulacral column on the peristome of 
larger specimens, and as many as 5 small, 
irregularly shaped scales in each interradial 
portion of the membrane (Fig. 4B). 

The ambital and aboral primary spines 
can be up to 2 times the diameter of the test 
and tapered. Several complete examples re- 
main attached to the holotype (Fig. 4C). 
Near the base, the spine is adorned with 
irregularly arranged, prominent thorns and 
serrations which diminish sharply in size 
about 30% of the spine’s length away from 
the base, giving way to strongly aligned, 
distal rows of spinules (Fig. 4F). Although 
the cross-section of the spine is broadly cir- 
cular, usually there are also well-developed 
keels set almost orthogonally to each other 
along the distal part of the spine. The de- 
gree to which these keels are developed 
varies among specimens and even on an in- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


dividual, but the basal thorns seem to be a 
constant feature. Basally, the spine can bear 
a surface coat of anastomosing hairs similar 
to that seen in some Aporocidaris. The oral 
primaries are differ from the aboral prima- 
ries in lacking strongly developed keels ex- 
cept laterally, where they can also be 
strongly serrated (Fig. 4G), particularly in 
specimens with thorny aboral primaries. 
The oral primaries adjacent to the peristome 
are very small and dagger- or arrowhead- 
shaped, with somewhat less distinctly ser- 
rated keels. In specimens with reduced 
keels on the aboral primaries, the oral pri- 
maries are less dagger-shaped, and more 
oval in outline. The scrobicular spines are 
slightly pointed (Fig. 4H). The non-scrobi- 
cular secondary spines are smaller, partic- 
ularly in the ambulacra (Fig. 41). 

As in most ctenocidarines, the globifer- 
ous pedicellariae come in two sizes, both of 
which are quite long and slender in N. Jan- 
ceolata, though not diagnostically so. The 
valves can be just over 1 mm long in the 
larger form. 

Types.—Holotype MCZ 8336, R/V EI- 
tanin Cruise 32, Station 2110 (Fig. 4C). 
Paratypes NMNH E21865 (3 dry speci- 
mens), E21866 (1 dry specimen), E22004 
(1 specimen in alcohol), E22005 (5 speci- 
mens in alcohol), E22006 (3 specimens in 
alcohol). We decided to retain the holotype 
designated in Fell (1976) to avoid confu- 
sion, and also because the specimen has ex- 
emplary spine cover (Fig. 4C). Drawings of 
spines and plate architecture were made 
from the paratypes to avoid damaging the 
holotype. 

Etymology.—In his thesis, Fell (1976) at- 
tributed the name to Richard Chesher, who 
recognized the distinctiveness of the new 
taxon in a manuscript that was never pub- 
lished. Chesher based the name on the 
keeled, lanceolate spines. 

Distribution.—Known from 7 R/V El- 
tanin stations (Cruise 27, Stations 1867, 
1926, 1929; Cruise 32, Stations 2002, 2108, 
2110, 2129) at three distinct localities in the 


VOLUME 113, NUMBER 1 to 


A a F 
Aboral plate LEZ = Aboral /', 
pattern primary {'f 


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Side view 
of holotype fj 


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s {| | 
NG GAS 
~ GY NV, Ss H fi 
We... ee Scrobicular | | 
secondary 1 
spine 


2! Interambulacrum at ambitus ea ern at ainbhGs 


Fig. 4. Notocidaris lanceolata new species. C drawn from holotype (MCZ 8336); A, B, D, E drawn from 
paratypes in lot NMNH E21865; F—I drawn from paratype (NMHH E 21866). Conventions for A and B as in 
Big: 2. 


236 


Ross Sea between 2005 and 2421 m (Fig. 
1): 

Biology.—The spines seem remarkably 
free of epizoans. The preferred habitat is 
unknown. Some specimens, which could be 
females, have enlarged gonopores, but no 
broods have been observed. No evidence of 
Echinophyces infection could be detected. 

Remarks.—As noted by Fell (1976:197), 
the species was based on a specimen res- 
cued by Dr. Merrill Foster from ‘material 
intended for disposal overboard”’ and later 
turned over to Chesher, who selected it as 
a potential type. The holotype, in having 
primaries that are thorny and strongly lan- 
ceolate, is close to one end of a range that 
includes forms in which the keels can be 
almost absent. As Fell (1976:197) indicat- 
ed, “‘were it not for the range of interme- 
diates between fluteless [unkeeled] and ful- 
ly fluted [keeled] specimens obtained at two 
stations, one would not believe they were 
of a single species”’. Typical N. lanceolata 
may be most easily confused with particu- 
larly thorny N. hastata, especially since 
they inhabit the same environments, while 
variants of N. lanceolata with greatly re- 
duced keels on the spines may be confused 
with N. gaussensis. In addition, the surface 
coating of anastomosing hairs between the 
thorns and keels near the base of the spine 
can cause some confusion with Ctenocidar- 
is, particularly if this feature alone is used 
to sort material. Because N. lanceolata oc- 
curs, on average, almost 1000 m deeper 
than N. gaussensis and virtually all Cteno- 
cidaris, available depth data should help to 
provide initial clues in identification. 


Acknowledgments 


We would like to thank Cynthia Ahearn 
(National Museum of Natural History) for 
her patience and helpfulness throughout our 
endless inquiries and loan requests and 
Chris Mah of the California Academy or 
Sciences (California Academy of Sciences) 
for digging out distant data. Fred Collier 
(Museum of Comparative Zoology) provid- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


ed invaluable information concerning the 
type of N. lanceolata. We are also grateful 
for the editorial comments of John Pearse 
and Dave Pawson. Mooi’s research was 
supported by an American Philosophical 
Society General Research grant. This paper 
is a contribution of the theme ‘‘Signal mor- 
phologique de Evolution” of the UMR 
CNRS 5561 “‘Biogéosciences.”’ 


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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 
113(1):238—248. 2000. 


A new species of Pristigaster, with comments on the 
genus and redescription of P. cayana 
(Teleostei: Clupeomorpha: Pristigasteridae) 


Naércio A. Menezes and Mario C. C. de Pinna 


(NAM) Museu de Zoologia, Universidade de Sao Paulo, Av. Nazaré 481, 
Sao Paulo-SP 04263-000, Brazil; (MCCdeP) Departamento de Zoologia, 
Universidade de Sao Paulo, Caixa Postal 11461, Sao Paulo-SP 05422-970, Brazil 


Abstract.—A new species of the hitherto monotypic genus Pristigaster (Clu- 
peiformes: Pristigasteridae) is described for the Amazon basin. Pristigaster 
whiteheadi, new species, is distinguished from its only congener, P. cayana, 
by the presence of pelvic fins; the lack of caudal-fin filaments; the different 
angle between the predorsal bones and the vertebral column; the presence of 
36—39 vertical scale rows (40—47 in P. cayana); and the presence of 18—20 
horizontal scale rows (21—26 in P. cayana). Pristigaster cayana is redescribed, 
and its occurrence in French Guyana is questioned. 

Resumo.—Uma nova espécie do género até entao monotipico Pristigaster 
(Clupeiformes: Pristigasteridae) é descrita para a bacia Amaz6nica. Pristigaster 
whiteheadi, espécie nova, distingue-se de P. cayana pela presenga de nadad- 
eiras pélvicas; auséncia de filamentos nos lobos da nadadeira caudal; o 4ngulo 
diferente entre os ossos pré-dorsais e a coluna vertebral; a presenga de 36—39 
fileiras verticais de escamas (40—47 em P. cayana); e a presenga de 18—20 
fileiras horizontais de escamas (21—26 em P. cayana). Pristigaster cayana é 


redescrita e sua ocorréncia na Guiana Francesa é questionada. 


The Clupeomorpha is a highly diverse 
group of teleosts, containing over 350 re- 
cent and over 150 fossil species (Grande 
1985, Nelson 1994). The group is one of 
the most important fisheries resources 
worldwide, and their phylogenetic relation- 
ships within teleosts have been a matter of 
intense debate in recent years. For the past 
two decades, following the original sugges- 
tion by Patterson & Rosen (1977), Clupeo- 
morphs were placed as the sister group to 
the Euteleostei. More recently, Clupemor- 
pha have been proposed as sister group to 
Ostariophysi, a hypothesis supported by 
molecular data (Van Le et al. 1993, Patter- 
son 1994) and morphological characters 
(Lecointre & Nelson 1996, Johnson & Pat- 
terson 1996, Arratia 1997). 

Clupeomorphs are a demonstrably mono- 


phyletic group (Grande 1985) including 
basal fossils as the extinct genera Diplo- 
mystus and Armigatus, and the extinct order 
Ellimmichthyiformes. Recent clupeo- 
morphs are all in the order Clupeiformes, 
itself divided into suborders Denticipitoidei 
(with a single species from African fresh- 
waters) and Clupeoidei (all other recent clu- 
peiforms). Clupeoids comprise three super- 
families: Engrauloidea (with a single fami- 
ly, Engraulididae), Clupeoidea (with fami- 
lies Chirocentridae and Clupeidae) and 
Pristigasteroidea (with families Pristigaster- 
idae and Pellonidae). 

The pristigasteroid family Pellonidae in- 
cludes the central and South American gen- 
era Chirocentrodon, Neoopisthopterus, Pel- 
lona, and Pliosteostoma. The Pristigasteri- 
dae, in turn, comprises the Central and 


VOLUME 113, NUMBER 1 


South American Odontognathus and Pris- 
tigaster, the South American and Indo-Pa- 
cific Opisthopterus and the Indo-Pacific Ra- 
conda. The genus /lisha could not be dem- 
onstrated monophyletic by Grande (1985), 
most of its species forming a polytomy at 
the base of Pristigasteroidea. 

Pristigaster is the most peculiarly-shaped 
of all pristigasteroids, with an extremely 
deep body resembling characiforms of the 
genera Gasteropelecus and Thoracocharax. 
So far a single species, P. cayana, is rec- 
ognized in the genus. All other proposed 
names have been shown to be either invalid 
or junior synonyms of that species (White- 
head 1973, 1985). However, Whitehead 
(1985) suggested that a second species 
might exist. Stimulated by Whitehead’s 
original suggestion, we undertook a de- 
tailed examination of available material of 
Pristigaster, and concluded that indeed 
there is a second diagnosable species in the 
genus, still undescribed. In this paper, we 
formally name and diagnose the new spe- 
cies and redescribe P. cayana. 


Methods and materials. 


Morphometric measurements were all 
point-to-point, taken with calipers, recorded 
to the nearest 0.1 mm and expressed as per- 
centages of standard length, except for sub- 
units of the head, expressed as percentages 
of head length. Counts and measurements 
were made on the left side of the speci- 
mens, whenever possible, according to 
Whitehead (1985), except for horizontal 
rows of scales (counted between dorsal-fin 
origin and anal-fin origin), vertical rows of 
scales (counted from origin of pectoral fin 
to caudal base), and scales around caudal 
peduncle (number of horizontal scale rows). 
Principal caudal-fin rays included all 
branched rays plus one unbranched ray in 
each lobe. Counts for each lobe, upper first, 
are separated by a slash. Vertebral counts 
were taken from radiographs and cleared 
and stained specimens and the terminal 
“half centrum” is included. Specimens 


239 


were dissected to determine sex by an in- 
cision on the right side of the abdomen to 
expose the gonads. Tooth counts include 
sockets in cases where the actual tooth has 
fallen off. Within the meristic information 
given here, figures for holotype are provid- 
ed in parentheses. 

Figures associated with specimen lists in 
species descriptions are, first, number of 
specimens examined in respective lot, and 
second, range of SLs in mm. 

Specimens cleared and counterstained for 
bone and cartilage were prepared by a mod- 
ified version of the method of Taylor & Van 
Dyke (1985). Descriptive accounts follow 
the general organization in Whitehead & 
Teugels (1985), the most complete anatom- 
ical survey of a clupeomorph available to 
date. Synonymic lists include only those 
references in which the species referred to 
can be reliably identified as either P. cay- 
ana or P. whiteheadi. 

Specimens examined in this work are 
deposited in the following institutions: 
AMNH, American Museum of Natural 
History, New York; BMNH, The Natural 
History Museum, London; FMNH, Field 
Museum of Natural History, Chicago; 
INPA, Instituto Nacional de Pesquisas da 
Amazonia, Manaus; MZUSP, Museu de 
Zoologia, Universidade de Sao Paulo, Sao 
Paulo. 


Pristigaster whiteheadi, new species 
Fisssf) 2B 


Pristigaster cayana (not Cuvier); White- 
head, 1985:301 (in part, only specimens 
with pelvic fins); Whitehead & Bauchot, 
1985:24 (in part, only specimens with 
pelvic fins); Stewart, Barriga & Ibarra, 
1987:21 (specimen examined). 


Holotype.-—MZUSP 52963 (female, 83.4 
mm SL). BRAZIL: Amapa, Rio Araguari, 
Ferreira Gomes, collected by M. Goulding, 
January—February, 1984. 

Paratypes.—Brazil: MZUSP 30341 (2, 
76.5—83.4), same data as holotype. Ama- 
zonas; MZUSP 11391 (8, 57.5—67.7, 2 of 


240 


which cleared and counterstained), AMNH 
227329: (15'52:2); USNIM351306°G1; 6220); 
FMNH 107783 (1, 56.5), Rio I¢a, Santo 
Anténio do I¢a4; MZUSP 11392-393 (2, 
69.4—73.0), Rio Solim6es, above mouth of 
Jutai; MZUSP 11394—403 (10, 43.5—76.2), 
Rio Solimdes, Fonte Boa; MZUSP 27597 
(1, 62.00, Rio Solimdes, Municipio de Ben- 
jamin Constant; MZUSP 18694 (3, 29.4— 
40.0), Rio Solimdes, Lago Janauaca and vi- 
cinity; MZUSP 52950 (2, 22.0—25.0), Rio 
SolimGes, 3°10'57”S, 67°56'31"W; MZUSP 
6600 (1, 70.0), Lago Manacapuru; MZUSP 
18512 (1, 67.0), mouth of Rio Ituxi; 
MZUSP 18516 (1, 70.0), mouth of the Pa- 
cia; INPA 8555 (21, 18.0—66.6, 3 of which 
cleared and stained), Parana do Tapura, near 
mouth of Rio Madeira; MZUSP 6220 (1, 
84.0), Rio Negro, Igarapé Jaraqui, above 
Manaus; MZUSP 52951 (1, 39.0), Rio 
Jauaperi, 1°34'54"S, 61°28'48"W; MZUSP 
52952 "(25 '57-7 ‘and 167.0) R10" Neero; 
1°33'48”"S, 61°33'02"W; MZUSP 49597 (3, 
28.3—37.3), Rio Acre, above Boca do Acre; 
MZUSP 7625 (1, 67.0), Rio Amazonas, Pa- 
rana do Mocambo, above Parintins; BMNH 
1897.12.1.197—-199 (3, 62.8—-65.0), Rio Ju- 
rua; MZUSP 52949 (21, 47.0—86.0), Rio 
Japurad, Parana do Japurd, 3°09'12"S, 
64°46'54"W; MZUSP 52962 (1, 63.3), Rio 
Amazonas, 1°54’S, 55°31’W; MZUSP 
52948 (5, 35.5—76.0), Rio Madeira, below 
Nova Olinda; MZUSP 52958 (1, 36.0), Rio 
Madeira, 3°33'37"S, 58°59'49"W; MZUSP 
52957. (1, 34.0)23° 3348S. 585957 Ww 
MZWSP »529592 Gs 4120)53 29) 2S, 
538°5.1°338" W, .MZUSP.. 52960. (1... 30:5): 
3°26'44"S, 58°49’49"W, MZUSP 52961 (4, 
30.5—41.0), 3°33’S, 58°55'W, Rio Madeira, 
Parana do Urucurituba. Roraima: MZUSP 
11404 (1, 33.0), Rio Branco, 20 kilometers 
below Boiacu. Paraé: MZUSP 52953 (1, 
28.0); 1535" S,.5271 1 WAMZUSP#52954. (7, 
50.0—55.0), 1°27’S, 52°03’W, Rio Amazon- 
as, Furo do Urucuricaia; MZUSP 5493 (1, 
85.0), Rio Trombetas, Oriximina; MZUSP 
529558(1;. 53.7),4 1529'S 52°35 Wa MZUSP 
52956 (8, 22.3—66.5), 1°36’S, 52°12'W, Rio 
Amazonas, below Rio Xingu. ECUADOR. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


FMNH 101946 (1, 89.0) 0°49’S, 75°31'W, 
Rio Tiputini, near mouth in Rio Napo and 
Quebradas). 

Diagnosis.—Distinguished from its only 
congener, P. cayana, by the following fea- 
tures: 1—presence of pelvic fins; 2—ab- 
sence of filaments on upper and lower 
lobes of caudal fin; 3—-vertical scale rows 
36-39 (40—47 in P. cayana); 4—horizon- 
tal scale rows 18—20 (21-26 in P. cayana); 
5—supraneurals (predorsal bones) gradu- 
ally less sloped posteriorly, posterior one 
nearly perpendicular to vertebral column 
(supraneurals all equally sloped in P. cay- 
ana). Most specimens of the new species 
can also be distinguished from P. cayana 
by lower gill raker counts (18—21, versus 
21-25 in P. cayana) and by lower anal-fin 
ray counts (41—48 versus 44—53 in P. cay- 
ana). 

Description.—Meristic and morphomet- 
ric data are presented in Tables 1 and 2. For 
a general aspect of the fish, refer to Fig. 1. 
Body highly compressed, ventral profile of 
body extremely expanded and convex, its 
anterior region (at isthmus) almost perpen- 
dicular to longitudinal axis of fish. Five 
protruding predorsal supraneurals strongly 
inclined anteriorly. The entire abdominal 
region, from isthmus to anal-fin origin, bor- 
dered by a series of 29-34 (holotype 31) 
abdominal scutes, gradually more promi- 
nent posteriorly. Scutes anterior to vertical 
through pectoral-fin base mostly imbedded 
in soft tissue, those posterior to that point 
protruding markedly beyond abdominal 
profile resembling a series of translucent 
hooks. Pelvic-fin origin usually over 25th 
scute (as in holotype), rarely over 24th or 
26th scutes. 

Snout blunt, always shorter than orbital 
diameter. Mouth subterminal and turned 
dorsally, its lower jaw protruding beyond 
upper. Maxilla extending slightly posterior 
to vertical through anterior margin of eye. 
Teeth conical, minute, disposed in a single 
irregular row in both jaws and highly var- 
iable in number, becoming more numerous 
with growth. Premaxillary teeth 8 (in spec- 


VOLUME 113, NUMBER 1 


Table 1.—Selected meristic features of Pristigaster cayana and P. whiteheadi. 


Pristigaster cayana Pristigaster whiteheadi 
n Range Mean SD n Holotype Range Mean SD 
Branched dorsal-fin rays 79 12-13 12.177 0.384 95 12 11-13 12.053 0.305 
Branched pectoral-fin rays 79 10-12 10.848 0.622 95 10 oI) 710.3537 0,522 
Branched pelvic-fin rays 95 3 3-4 3.137 0.346 
Branched anal-fin rays 73 44-53 48.096 2.076 93 44 41-48 44.258 1.436 
Gill rakers TT Y23=25+° 22,208 - 0:8468 95 Ps 18-21 19.895 0.722 
Horizontal rows of scales 55 21-26 23.182 1.090 a, — 18-20 19.114 0.631 
Vertical rows of scales 39 40-47 43.077 1.797 18 — 36-39 38.222 0.878 
Scales around caudal peduncle 18 15-16 ~13,833- 0.383 9 — 13-15", 14.556. 0.726 
Premaxillary teeth 58 9-23 16.862 3.322 78 14 8-18 11.962 1.970 
Maxillary teeth 62 18-80 53.871 14.140 84 63 14-64 40.000 11.600 
Dentary teeth 49 5-14 9.408 2.188 53 5-9 ye lw: 1.301 
Ventral scutes Ta. 30535 32,013 IEP | 92 31 29-34 31.065 0.849 
Vertebrae 7 42-44 43.428 0.728 | — 43-44 43.286 0.488 


imens <44 mm SL) to 18 in larger speci- 
mens (holotype 14). Maxillary teeth 14 (in 
22 mm SL specimen) to 64 (in 79 mm SL 
specimen) (holotype 63). Dentary teeth 6 
(at 43 mm SL) to 9 in larger specimens 
(holotype 9). Eyes very large, round in ex- 
ternal aspect. Pupil extremely large rela- 
tive to eye size, its diameter almost 60% 
that of orbit. Adipose eyelid weakly de- 
veloped in young, but covering most of iris 


in adults. 


Gill cover rounded in profile, with con- 
tinuous membranous margin concealing a 
marked depression on posterior margin of 
opercle. Dorsal limit of gill opening slightly 
ventral to horizontal at dorsal margin of 
eye. Gill rakers slender, closely set and con- 
spicuous, their number increasing with 
growth, ranging from 18 (at 22 mm SL) to 
21 (at 83 mm SL) (holotype 21) on lower 
part of first branchial arch and from 8 to 12 
(holotype 11) on upper part. 


Table 2.—Morphometrics of Pristigaster cayana and P. whiteheadi. Standard length is expressed in mm; 
measurements | to 12 are proportions of standard length; 13 to 16 are proportions of head length. 


Pristigaster cayana 


Character n Range Mean 
Standard length 79 =—-.23.5-142.0 66.68 
1. Body depth 78 0.38-0.65 0.550 
2. Predorsal length 79 ~=0..39-0.48 0.421 
3. Prepectoral length [Bie o0.23-031 0.271 
4. Prepelvic length 
5. Preanal length 78  0.57-0.70 0.649 
6. Caudal peduncle depth 79  0.08-0.11 0.096 
7. Head length 79 ~=—-0.25-0.31 0.287 
8. Dorsal-fin base 79 ~=—-0..12-0.16 0.138 
9. Dorsal-fin length 72 ~=0.24—0.33 0.283 
10. Anal-fin base 78 0.42-0.53 0.486 
11. Pectoral-fin length 79 = 0.17-0.28 0.250 
12. Pelvic-fin length 
13. Snout length 79 ~=0.18-0.26 0.223 
14. Eye diameter 79 ~=0.33-0.44 0.393 
15. Interorbital width 79 = -0.12-0.21 0.176 
16. Upper jaw length 79 ~=0.41-0.57 0.486 


Pristigaster whiteheadi 


SD n Holotype Range Mean SD 
— oy ‘S3o.4 22.0—86.0 56.12 — 
0.041 95 0.56 0.42-0.58 0.534 0.031 
0.014 95 043 0.40-0.48 0.425 0.017 
O.0l2., O35 0:27 , 30.24-0.31,, 0272. >, 0.013 

95 O59. “OAS-O.6Ok O57) . 0025 
O;025: -95 0.67 0.54-0.71 0.657 0.024 
0.008 95 0.09 0.07-0.12 0.098 0.007 
DOIG" -95. 9 1029". O25-0:32-' 0.286 0:012 
D009 95 euOil2s 0:12-0:17 ~ 0.145) 0012 
0.020 94 0.26 0.24-0.32 0.289 0.018 
0.026 95 0.44 0.42-0.50 0.467 0.016 
UG. 3s 0.28  U.16-0°26° “0.230 ~“OOT 

94 0.04 0.02-0.07 0.048 0.006 
O02) “95 0.25 0.21-0.29 0.247 0.015 
0.022 95 0.40 0.34—-0.45 0.399 0.023 
0.016) +956» 0.20. -018-0.25.... 0:205.., 0.013 
0.033: 95, , O49 ©.43-0:53 0:479. . 0.019 


242 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Le: 


Bigeile 


Dorsal fin triangular when extended, 
pointed dorsally; origin located on anterior 
half of SL, and anterior to vertical through 
anal-fin origin. Dorsal-fin rays 11+(12)—13, 
third ray (unbranched) longest, twice as 
long as fin-base. Pectoral-fin origin located 
at middepth of body, slightly dorsal to ven- 
tral margin of gill cover and slightly ante- 
rior to its posterior margin. Pectoral-fin 
length shorter than HL, its posterior tip, 
when adpressed to body, reaching beyond 
vertical through dorsal-fin origin. Pectoral- 
fin rays 1+(10)—11. Axillary scale present 
on region dorsal to pectoral-fin base, ex- 
tending for approximately 20% of fin. Pel- 
vic fins minute, length spanning approxi- 
mately the space of three consecutive ab- 
dominal scutes, with origin located nearly 
at ventral margin of body, at vertical 
through posterior tip of pectoral fin. Pelvic- 
fin rays i+3+i. Anal fin long and low, its 
origin slightly posterior to vertical through 
posterior end of dorsal-fin base. Anal-fin 
progressively shorter posteriorly. Tips of 
last anal-fin rays reaching base of inferior 
caudal-fin basal fulcra. Caudal fin deeply 


Pristigaster whiteheadi, new species, holotype, MZUSP 52963, 83.4 mm SL. 


forked, lower lobe slightly longer than up- 
per. Caudal-fin rays 10/9. Vertebrae 43 or 
44. 

Scales large (approximately same size as 
pupil) and cycloid, covering all of body, 
and easily detachable, rarely preserved in 
their entirety. Vertical scales rows 36-39. 
Horizontal rows 18—20. Scale covering ex- 
tending to midlength of middle caudal-fin 
rays. Entire anal-fin base rimmed by row of 
small scales (about half as large as remain- 
ing body scales). 

Pigmentation in alcohol.—Dorsum, 
snout and upper part of sides brown. Re- 
mainder of body silvery due to heavy de- 
posits of guanine (which tend to disappear 
after extended preservation). Narrow con- 
centrations of dark melanophores on upper 
and lower lips, and sometimes anterior por- 
tion of chin. Region corresponding to neu- 
rocranium dark, due to brain pigment visi- 
ble through translucent skull bones. Scat- 
tered dark chromatophores along entire dor- 
sum and upper sides, more concentrated 
along dorsal-fin base and dorsal part of cau- 
dal peduncle. A middorsal dark spot is 
sometimes present anterior to dorsal fin (as 


VOLUME 113, NUMBER 1 


avec 


7 ee 


Fig. 2. 


/ 


ave 


B 


Predorsal bones and corresponding neural spines in lateral view, anterior to left. A—Pristigaster 


cayana, MZUSP 30338; B—P. whiteheadi, n. sp., paratype, INPA 8555; Abbreviations: pb—predorsal bone, 
ns—neural spine, avc—main axis of vertebral column. Scale bars = | mm. 


in holotype). Myosepta visible but not out- 
lined by dark pigmentation. Fins hyaline, 
except for faint rows of melanophores 
along outer rays of caudal fin and a few 
scattered isolated melanophores on dorsal 
fin. 

Etymology.—The name of this species is 
a posthumous tribute to Peter J. P. White- 
head, who contributed more than any other 
individual to the knowledge of clupeo- 
morph fishes, also for first suggesting that 
there might be two different species in Pris- 
tigaster. 

Distribution.—Brazil and Ecuador, in the 
following river basins: Rio Solim6es/Ama- 
zonas, Rio Trombetas, Rio Branco, Rio Ja- 
pura, Rio Negro, Rio Madeira, Rio Jurua, 
Rio Araguari (State of Amapa, Brazil) and 
Rio Napo. 


Pristigaster cayana Cuvier, 1829 
Figs. 3, 4 


Pristigaster cayanus Cuvier, 1829: pl. 10, 
fig. 3; 1829:321 (name only, American 
seas); Valenciennes, 1847 (redescription 
of holotype). 


Pristigaster argenteus Schinz, 1822:300 


(based on Cuvier’s figure; nomen obli- 
tum). 

Pristigaster lichtensteinii Jarocki, 1822: 
322, fig. 3 (based on Cuvier’s figure, re- 
produced; nomen oblitum). 

Pristigaster triangularis Stark, 1828:408 
(based on Cuvier’s figure; nomen obli- 
tum). 

Pristigaster martii Agassiz, in Spix & Ag- 
assiz, 1829:55, pl. 24a (Amazon); no pel- 
vic fins; post-dates cayanus by a month 
or so (see Whitehead & Bauchot, 1985); 
Amaral-Campos, 1941:187; Whitehead 
é& Myers, 1971;-Le Bail et al.,. 1983. 

Pristigaster americanus Guérin-Menéville, 
1844:33, pl. 57, fig. 3 (Atlantic coast of 
South America). 

Pristigaster phaeton Valenciennes, 1847: 
338 (Amazon); no pelvic fins (see White- 
head & Bauchot, 1985). 

Pristigaster cayana; Myers, 1956 (validity 
of P. cayanus Cuvier as of 1829; em- 
mendation of specific name to agree with 
feminine gender of genus); Hildebrand, 
1964 (synonymy of cayana, phaeton and 
martii); Whitehead, 1964:428, fig. 108 
(synopsis); 1967:100, 102 (types of cay- 
ana and phaeton); 1973a:85 (triangularis 


244 


a nomen oblitum; synonymy); 1985:301 
(in part, only specimens without pelvic 
fins; synonymy, diagnosis; distribution; 
habitat and biology); Whitehead & My- 
ers, 1971:487 (validity of martii); White- 
head & Bauchot, 1985:24 (in part, only 
specimens without pelvic fins; types of 
cayana, phaeton); Eschmeyer, 1998:347 
(catalog and suggestion—not accepted 
here—that name is not available from 
Cuvier, 1829). 


Material examined.—95_ specimens 
(23.5-142.0 mm SL). Brazil. Amazonas: 
MZUSP 31032 (1, 90.0), Lago do Prato, 
Rio Negro, Anavilhanas; MZUSP 11389 (1, 
142.0), Lago Puraquequara; MZUSP 52947 
(1, 64.0), Lago Manacapuru; MZUSP 7023 
(20, 52.5—-74.0, 2 of which cleared and 
counterstained), Rio Madeira, 25 kilometers 
below Nova Olinda; MZUSP 11405 (1, 
37.5); Rio, Solimoes,. Ilha do -Xibeco; 
MZUSP 9568 (1, 96.0), Manaus; MZUSP 
18696 (1, 92.0), Rio Solimdes, Lago Jan- 
auaca; MZUSP 52943 (1, 23.5), Rio Negro, 
1°58: 167S3261- 115/A22W:MZUSP52944.(6, 
30.0—36.0), Rio Icd, 3.:O3.45,09, 
68°04'26"W; MZUSP 52945 (1, 34.0), Rio 
SolimGes, 2°40'15”S, 66°39'14"W; MZUSP 
52946 (1, 62.2), Rio Japura, Parana do Ja- 
pura, 3°08'20"S, 64°46'52”W; MZUSP 1388 
(1-11 1.0),and 11390<(1,;36.0); Rio: Jurua: 
BMNH 1925.10.28.5 (1, 88.9), Rio Soli- 
moes, Manacapuru. Roraima: MZUSP 
30334 (1, 137.0), Rio Branco, below Xe- 
ruini; MZUSP 30335 (1, 112.0) and 30337 
(1, 103.0), Rio Branco, Marara; MZUSP 
30339 (2, 100.0 and 110.0), Rio Branco, 
Lago do Maguari; MZUSP 30340 (6, 68.3— 
82.5), Rio Branco, Xeruini; MZUSP 52942 
(1,. 105-0):,. Rio 26Branceo,. 1 7:6.59"S, 
61°50'52”W. Rondonia: MZUSP 30336 (1, 
81.0) and 30338 (29, 33.0—66.0, 3 of which 
cleared and counterstained), Rio Madeira, 
Calama. Para: MZUSP 5560 (2, 72.0 and 
73.5), Lago Ururié, Oriximina; MZUSP 
5668 (1, 82.5), Lago Puru, Oriximina; 
MZUSP 8280 (1, 97.0), Rio Trombetas, 
Oriximina; MZUSP 5689 (1, 72.0), Rio 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Trombetas, mouth of Lago Paru. Mato 
Grosso: MZUSP 17030 (1, 29.0), Rio Ar- 
aguaia, Santa Terezinha; MZUSP18627 (2, 
116.0 and 126.0), Rio Araguaia, Lago 
Dumba. PERU. Loreto: FMNH 71264 (1, 
100.0), mouth of Rio Tigre, 80 miles SW 
of Iquitos; MZUSP 15217 (1, 83.0), Cocha 
Aguajal, Rio Amazonas, Iquitos: MZUSP 
15216 (2, 83.0 and 88.0), Rio Marafion, 
Nauta; Pucallpa: MZUSP 18557 (1, 47.0), 
Rio Ucayali. 

Diagnosis.—See diagnosis of P. white- 
headi. 

Description.—Meristics and morphomet- 
rics are presented in Tables 1 and 2. Body 
form and disposition, shape and arrange- 
ment of abdominal scutes (30-35 in num- 
ber) as in P. whiteheadi. 

Shape, size and position of snout, eye, 
pupil, adipose eyelid, mouth, maxilla and 
also shape, arrangement and number of 
teeth as in P. whiteheadi. Number of teeth 
also increasing with growth. Premaxillary 
teeth 9 (in specimens <25 mm SL) to 23 
in larger specimens. Maxillary teeth 18 (in 
23.5 mm SL specimen) to 80 Gin 105 mm 
SL specimen). Dentary teeth 7 (at 54 mm 
SL) to 14 in larger specimens. 

Shape of gill cover, gill membrane and 
shape and disposition of gill rakers and po- 
sition of dorsal limit of gill opening as in 
P. whiteheadi. Number of gill rakers also 
increasing with growth, ranging from 21 (at 
45 mm SL) to 25 (at 110 mm SL) on lower 
part of first branchial arch and from 9—12 
on upper part. 

Pelvic fins absent. Position, shape and 
size of all other fins and axillary scale iden- 
tical to those of P. whiteheadi. Dorsal-fin 
rays 111+12—13. Pectoral-fin rays i+10—12. 
10/9. Tips of dorsalmost two branched up- 
per lobe caudal-fin rays elongated into fil- 
ament twice as long as first principal ray 
(unbranched) in a specimen 100 mm SL. 
Tips of eighth and ninth lower lobe caudal- 
fin rays also prolonged into a shorter lower 
lobe filament about one-third as long as 
tenth (unbranched) ray in same specimen. 


VOLUME 113, NUMBER 1 


25 
20 A A At 
ds, 
A A A 
= A A AA 
2 AA A 
¢ 15 A oOo 
* Oo Oo 
& Oo 
2 
Qa 
= 
3 10 Oo 
oOo 

= 
e) 
2 

=) 

i¢) ca = all a RJ a - er — ee ee mee ee 

0 20 40 60 


Fig. 3. 


Upper and lower caudal-fin filaments fre- 
quently broken in preserved specimens. 
Vertebrae 43-44. 

Size, shape and distribution of cycloid 
scales on body and fins as in P. whiteheadi. 
Vertical scale rows 40—47. Horizontal rows 
21-26. 

Pigmentation in alcohol.—As in P. 
whiteheadi. 

Distribution.—Nearly coincident with 
that of P. whiteheadi, with which it is sym- 
patric in most localities of the Amazon Ba- 
sin, but extending further south into Rio 
Araguaia, State of Mato Grosso, Brazil. 
The locality associated with the holotype 
of the species is reported as Cayenne, 
French Guiana (Whitehead 1967). This in- 
formation is not provided in the original 
accounts on the species by Cuvier, which 
State simply “‘seas of America’’. Other re- 
ports on the type-locality have inferred it 
from the species name (e.g., Myers 1956). 
Representatives of Pristigaster have never 
been collected again in French Guiana, in 
any of the surveys of the area (e.g., Le Bail 


245 


o o 
o o 
Oo o 
A oo Oo 
ao oo Oo 
Oo 
Oo Oo 
o 8a 5 
o 
O Other samples of P. cayana 
4 Rio Madeira sample 
| 
| 
80 100 120 140 160) 


Standard lenght - mm 


Size-dependent variation of the number of premaxillary teeth in P. cayana. 


et al. 1983, Planquette et al. 1996, also P. 
Keith, pers. comm.). The genus has also 
not been reported from surrounding areas, 
like Guyana, Suriname or the Orinoco ba- 
sin. We strongly suspect that the Cayenne 
locality is erroneous, and may simply re- 
flect a port of shipment or an intermediate 
post en route between South America and 
France, in which the material studied by 
Cuvier may have remained temporarily. 

Remarks.—The sample from Rio Madei- 
ra differs from remaining ones in number 
of premaxillary teeth (Fig. 3), but no other 
meristic or morphometric difference was 
found between that population and others 
throughout the range of the species. In the 
absence of additional significant differences 
we prefer to consider the higher number of 
premaxillary teeth in the Rio Madeira sam- 
ple as populational variation. 

Notes on the name P. cayana.—The his- 
tory of the names associated with the spe- 
cies referred to here as Pristigaster cayana 
is rather complicated. The first reference to 
the species was done in Cuvier (1816), as 


246 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 4. Pristigaster cayana, MZUSP 30340; caudal fin in lateral view, showing filaments. Scale bar = 1 mm. 


an illustration associated with the genus 
name only. In 1829, Cuvier proposed the 
name cayanus for the species figured in 
1816, in what is clearly a valid species de- 
scription as was shown by Myers (1956). 
The species name therefore must date from 
1829, and it precedes the validly described 
P. martii by a month or so (Whitehead 
1985:301). Various other names (argen- 
teus, lichtensteinii and triangularis) were 
proposed between 1816 and 1829, all on 
the basis of the same illustration in Cuvier 
(1816). We consider all of these names to 
qualify as nomina oblita according to the 
International Code of Zoological Nomen- 
clature (ICZN, 1985, Art. 79c). They have 
never been used subsequently as the valid 
names for the species, and the junior syn- 
onym, cayanus, has certainly been used in 
more than 10 publications by more than 
five authors in the intervening period (e.g., 
Valenciennes 1847, Giinther 1868, Jordan 
& Evermann 1896, Norman 1923, Ihering 
1930, Myers 1956, Whitehead 1967, 1973, 
1985; Whitehead & Bauchot 1985). 


Discussion 


As demonstrated in this paper, in spite of 
the overall similarity in body shape and pig- 
mentation, Pristigaster cayana and P. whi- 
teheadi are clearly distinct and diagnosable 
through several morphological features. 
When suggesting the recognition of a sec- 
ond species of the genus, Whitehead (1985) 
and Whitehead & Bauchot (1985) based 
their observations on just a few specimens 
and did not notice some of the striking dif- 
ferences we found. In the diagnostic fea- 
tures of P. cayana, Whitehead (1985) de- 
scribed the pelvic fins as usually absent, in- 
dicating that presence or absence could 
vary within the species. At the same time, 
however, the author considered that one 
Species without pelvic fins and high gill 
raker counts (22—24, usually 23) could pos- 
sibly be different from a species bearing 
pelvic fins with lower gill raker counts (19— 
20, usually 20). Results of our study reveal 
that presence of pelvic fins is indeed diag- 
nostic. They exist only in P. whiteheadi and 


VOLUME 113, NUMBER 1 


are consistently present from the smallest to 
the largest individual of both sexes. Gill 
raker counts overlap to a certain extent, but 
there is a significant mean difference (see 
Table 1) between the two species. Other dif- 
ferences pointed out in the diagnosis of P. 
whiteheadi leave little doubt that there are 
indeed two separate species in Pristigaster. 
None of the various other names applied to 
P. cayana (see synonymy above) could 
possibly have been based on the new spe- 
cies. They either were based on Cuvier’s 
1816 illustration or clearly mentioned the 
absence of pelvic fins in specimens exam- 
ined. 

The two Pristigaster species share the 
highly peculiar expanded morphology of 
the abdomen, to a degree which distinguish- 
es them from all other recent clupeiforms. 
Not only the depth, but also the shape of 
the abdominal expansion (abruptly emerg- 
ing nearly vertically from gular region) are 
obviously apomorphic conditions not seen 
elsewhere in other recent clupeomorphs, 
and strongly suggest that Pristigaster is 
monophyletic. These characteristics are as- 
sociated with a host of internal-anatomical 
modifications not yet studied in detail, and 
which will be the subject of a forthcoming 
paper. 

Sexing of most of the specimens of both 
species examined did not reveal any obvi- 
ous sexual dimorphism. It also tested the 
validity of our interspecific diagnostic char- 
acters by showing that the differences were 
not simply due to sexual dimorphism. 


Acknowledgments 


We thank Efrem Ferreira and Paulo Petry 
(INPA), Tony Gill and Oliver Crimmens 
(BMNH), and Barry Chernoff and Mary- 
Anne Rogers (FMNH), for the loan of ma- 
terial under their care. We are also grateful 
to Philippe Keith for information regarding 
collections in French Guyana. The manu- 
Script benefited from reviews by Fabio Di 
Dario, Lance Grande, Thomas Munroe and 
an anonymous reviewer. Research funding 


247 


for both authors is provided by the Conse- 
Iho Nacional de Desenvolvimento Cientifi- 
co e Tecnolégico (CNPq; individual grants) 
Projeto PRONEX, and Fundagao de Am- 
paro a Pesquisa do Estado de Sao Paulo 
(FAPESP). 


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. 1829. Le Réne Animal Distribué d’ Apres son 
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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


113(1):249-—263. 2000. 


A new species of Apogon (Perciformes: Apogonidae) from the Saya 
de Malha Bank, Indian Ocean, with redescriptions of Apogon regani 


Whitley, 1951, A. gardineri Regan, 1908, and A. heraldi 
(Herre, 1943). 


Thomas H. Fraser 


W. Dexter Bender and Associates, Inc., 2052 Virginia Avenue, Fort Myers, Florida 33901, U.S.A. 


Abstract.—A new species of fish, Apogon quartus, known only from the 
Mascarene Plateau, is described. It is related to Apogon poecilopterus Cuvier 
in Cuvier and Valenciennes, 1828, Apogon carinatus Cuvier in Cuvier & Va- 
lenciennes, 1828, and Apogon queketti Gilchrist, 1903, recently treated by Gon 
(1996) as members of the subgenus Jaydia Smith, 1961. Apogon quartus can 
be distinguished from these three species by having a single predorsal scale, 
18 total (13 well developed) gill rakers on the first arch, and a single dark spot, 
perhaps appearing as an ocellus in life, on the body just behind the opercle 
flap. Six additional nominal species are added to the 10 valid (19 nominal) 
species treated by Gon in Jaydia. Three of these names, Apogon argyrogaster 
Weber, 1909, formerly in Siphamia, Apogon melanopus Weber, 1911, and Apo- 
gon fuscomaculatus Allen & Morrison, 1996, are valid species. The holotype 
of Apogon heraldi (Herre, 1943) redescribed here, is a synonym of A. poeci- 
lopterus. Apogon fuscovatus Allen, 1985 was determined to be a synonym of 
A. melanopus by Allen & Morrison (1996) The other name, Apogon tchefouen- 
sis Fang, 1942, may be synonymous with one of the species Gon treated but 
is not placed with certainty. Character overlaps between the subgenus Jaydia 
and other Apogon subgenera, particularly the largest subgenus Ostorhinchus 
Lacépeéde, 1802, are briefly examined. Virtually all derived characters of Jaydia 
grade into Ostorhinchus. Two rare species Apogon regani Whitley, 1951 and 
Apogon gardineri Regan, 1908 only known from the Mascarene Plateau are 
redescribed from new material. Both belong with the Apogon nigripinnis Cu- 
vier in Cuvier and Valenciennes, 1828, complex of species. 


The Russian vessel Vityaz made trawl 
stations along the Saya de Malha Bank dur- 
ing 1989 collecting fishes. Two of these 
collections contained a new species and a 
specimen of Apogon regani Whitley, 1951. 
The new species was first identified as an 
Apogonichthys Bleeker, 1854c, based on the 
mostly smooth preopercle edge and the 
slightly rounded caudal fin. An examination 
of internal and external characteristics led 
to the conclusion that this specimen is an 
Apogon Lacépéde, 1802. Some characters 
are held in common with species in the sub- 


genus Jaydia recently treated by Gon 
(1996). The ten species of Jaydia have the 
fourth dorsal spine longer than any other 
dorsal spine, variable serrations on pre- 
opercular edges, rounded or truncated cau- 
dal fins, and some have bioluminescent or- 
gans. Gon and Allen (1998) included an- 
other new species, Apogon photogaster in 
Jaydia. A review of additional nominal spe- 
cies with possible relationships to the new 
species yielded six names, Apogon argyr- 
ogaster Weber, 1909, (previously in Si- 
phamia Weber, 1909, see Lachner 1953, p. 


250 


416), Apogon melanopus Weber, 1911, 
Apogon tchefouensis Fang, 1942, Apogon 
heraldi (Herre, 1943), Apogon fuscovatus 
Allen, 1985 and Apogon fuscomaculatus 
Allen & Morrison, 1996. 

Specimens of all the above species have 
been most frequently taken by trawls, often 
in deeper waters. Fishes in Jaydia have 
body shapes ranging from elongate more 
slender forms, Apogon truncatus Bleeker, 
1854e, to A. melanopus a deeper body form 
with an emarginate caudal fin typical of 
most Apogon. Smith (1961) based Jaydia 
on Apogon ellioti Day, 1875, now recog- 
nized as a synonym of A. truncatus a slen- 
der species with a rounded caudal fin and 
having bioluminescent activity. 

In the process of trying to locate more 
material of the new species, specimens of 
two deep-dwelling species from the Naza- 
reth Bank near Cargados Carajos Shoals 
(St. Brandon) described by Regan (1908) 
were found. Randall and van Egmond 
(1994) reported Apogon punctatus Regan, 
1908, (=Apogon regani Whitley, 1951) tak- 
en in a trawl from the Seychelles. No ma- 
terial of Apogon gardineri Regan, 1908, has 
been reported since the original description, 
but deep dives by J. E. Randall during 1979 
in Mauritius yielded two specimens. It is 
likely that more material will be only slow- 
ly forth coming. Both of these species are 
regarded as endemics of the Mascarene Pla- 
teau. 


Methods 


Methods of taking and recording meristic 
data and measurements are given in Fraser 
& Lachner (1985). All measurements are in 
millimeters to the nearest 0.1 mm. All pro- 
portions are based on standard length and 
all material is reported by standard length 
rounded to the nearest millimeter, except 
for the primary type material. All x-ray 
photographs are in data files maintained by 
the author. The acronyms used in the lists 
of materials to designate institutions and 
collections cited, follow general usage giv- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


en in Leviton et al. (1985) and Eschmeyer 
(1998). 


Apogon quartus, new species 
Figs. [dé 2 


Material examined.—Holotype: USNM 
307688; (49.8); Indian Ocean, Saya de Mal- 
ha Bank, 11°05’00"S, 62°02’00"E; Vityaz 
Cr. 17; Sta 2808; 8 Jan 1989; 58—61 m.; X- 
ray. 

Comparative material.—Amia albomar- 
ginata Holotype: USNM 68402; (83.6); 
Philippines, Cavite; X-ray. Apogon arafur- 
ae Holotype: BMNH 1879.5.11.141; (90.0); 
Arafura Sea. Apogon argyrogaster Syn- 
types: ZMA 101075; (34.9—47.5); New 
Guinea, west coast; Siboga sta 164; 32 m. 
BPBM 32628; (15.5—48.3); New Guinea, 
Nagada Harbor; 18 Nov 1987; 30 m; X-ray. 
Apogon bilaciniatus Lectotype: ZMA 
101280; (46.2); Indonesia, Lombok. Apo- 
gon ellioti Paralectotype?; ZSI 1905; (77.7, 
96.0 mm TL); India, Madras; X-ray. Para- 
lectotype?; AMS B.8226; (77.3, 98.1 mm 
TL); India, Madras; X-ray. Apogon fusco- 
vatus Holotype: WAM P14397; (94); Aus- 
tralia, Darwin; 4 Sep 1965. Paratypes: 
WAM P14516—-17; (83-85); Australia, N. 
of Darwin; 9 Sep 1965. WAM P 28316- 
001; (66); Australia, Darwin; 10 Sep 1965. 
Apogon glaga Syntype: RMNH 5614 (71.9, 
~93 mm TL); Indonesia. Apogon hungi 
Neotype: USNM 340009; (76.4); Mozam- 
bique Channel. Jaydia hungi Holotype: 
MNHN 1965-711 (94.3); Egypt, Gulf of 
Suez. Apogon lineatus Lectotype: RMNH 
70a; (62.4); Japan. USNM 71240; (64.6); 
Japan, Shimizu Suruga; Albatross; 1906; 
female. USNM 32586: (66.3): China: 
35°55'49"N,. 120°2129°E; 25 Jule 1993 -ie- 
male. Amia melas [=Apogonichthys melan- 
opterus| Holotype: ANSP 47491; (~35.2 
head distorted); Philippine Islands. Apogon 
melanopus WAM P. 14963; (100); Austra- 
lia, Darwin; 4 Sep 1965. Apogon modestus 
Holotype: RMNH 5579; (61.9); Indonesia, 
Java. Apogon nigricans Syntype: ZSI 1872; 
(55.0, 71+ TL) India, Madras; X-ray. Apo- 


VOLUME 113, NUMBER 1 


Hire... 


gon novaeguineae Holotype: MNHN 8695; 
(56.5); New Guinea. Apogon photogaster 
Paratype: USNM 348214; (43.3); Papua 
New Guinea, Madang Lagoon; 21 Oct 
1996; 18—23 m; X-ray. Apogon poecilop- 
terus Holotype: RMNH 214; (79.0); Indo- 
nesia, Java. Apogon queketti Syntypes: 
SAM 11657; (75.1); South Africa, Natal; 
X-ray. SAM 11658; 5(44.1-—77.4); same 
data; X-ray. Jaydia smithi Paratypes: ZMH 
5034; 5(12.7—46.3); Amia striata Holotype: 
USNM 68403; (67.2); Philippines, Luzon; 
female; X-ray. Paratypes: USNM 93410; 
11(39.2—66.2); same data; X-ray. Somalia, 
Gulf of Aden; X-ray. Apogon striatodes 
Holotype: USNM 213408; (55.5); Indian 
Ocean, Thailand. Paratypes: China, Hong 
Kong: CAS 160877; (57); CAS 161015; 
(55). Philippines, Luzon: CAS 85669; (45); 
Corregidor I.: CAS 32723 (46). Thailand, 
Gulf of Thailand: CAS 79648; (36-37); 
CAS 79652; (44); (CAS 82205; 8(18—-59); 
CAS 85676; (26-30); CAS 85666; 3(28— 
34); CAS 85667; (19); CAS 82202; (17- 
47); CAS 82208; (54). Apogon tchefouensis 
Paratypes: MNHN 1941-148; (45.7); China, 
Shandong Prov., Tché-Fou, Ho Ting Chieh; 
X-ray. MNHN 1941-149; (35.0); same data; 
X-ray. Apogon truncatus Holotype: RMNH 
5582; (55.4); Indonesia, Batavia (=Jakarta). 


The holotype of Apogon quartus from the Saya de Malha Bank, Indian Ocean, 49.8 mm SL. 


Apogon hoevenii Syntype: RMNH 5581; 
(40:2; 50m“ TE; one ‘of: 6"'spec: imisize 
range, 24 spec. 25—49 mm SL); Indonesia, 
Ambon. USNM 261058; 8(35—39); Philip- 
pines, Siquoijor I. Apogon sp. USNM 
349199; (48.8); Philippines, Palawan, 
Puerto Princesa City. 

Diagnosis.—A species of Apogon in the 
subgenus Jaydia with 15 pectoral rays, gen- 
erally smooth to crenulate preopercle edge, 
one predorsal scale, 18 total (13 well de- 
veloped) gill rakers on the first arch, fourth 
dorsal spine the longest, and a single dark 
spot, perhaps appearing as an ocellus in life, 
on the body just behind the opercle flap. 

Description.—For general body shape 
see Fig. 1. Proportions (as percentage of 
standard length): greatest body depth 33.5; 
head length 42.2; eye diameter 11.4; snout 
length 9.2; bony interorbital width 10.2; up- 
per-jaw length 20.5; caudal-peduncle depth 
14.8; caudal-peduncle length 21.5; first dor- 
sal-spine length 3.4; second dorsal-spine 
length 9.0; third dorsal-spine length 15.7; 
fourth dorsal-spine length 17.9; spine in 
second dorsal fin 14.8; first-anal-spine 
length 2.4; second-anal-spine length 11.4; 
pectoral-fin length 21.2; pelvic-fin length 
22. 

Dorsal fin VII-I,9; anal fin 1,8; pectoral 


252 


fin 15-15; pelvic fin I,5; principal caudal 
rays 9 + 8; number of simple pored lateral- 
line scales unknown, series extending from 
posterior edge of posttemporal to caudal 
fin; transverse scale rows above lateral line 
2: transverse scale rows below lateral line 
~4; median predorsal scales 1, the lateralis 
system extending well onto the nape as a 
raised semi-translucent fleshly area of about 
seven striations or rows with many small 
papillae; number of circumpeduncular scale 
rows unknown; total gill rakers 18, well de- 
veloped 13, 342 upper arch,. 11--2 lower 
arch. 

Villiform teeth in wide band on premax- 
illa; wide band grading to two rows on den- 
tary; one to two rows on palatine and one 
to three rows on vomer; none on ectopter- 
ygoid, endopterygoid or basihyal. 

Vertebrae 10 + 14. Five free hypurals, 
one pair of slender uroneurals, three epur- 
als, a free parhypural. Three supraneurals, 
two supernumerary spines on first dorsal 
pterygiophore. Basisphenoid present. Su- 
pramaxilla absent. Posttemporal smooth on 
posterior margin. Preopercular ridge 
smooth, posterior vertical edge smooth, 
ventral horizontal edge slightly crenulate 
with one or two spinelets at angle. Infra- 
orbital with irregular edges, but without ser- 
rations. Scales ctenoid. Caudal fin slightly 
rounded. Nose and pre-interorbital area 
semi-translucent and slightly bulbous. No 
indication of bean-like bioluminescent or- 
gans associated with intestine near anus. 

Life colors.—Unknown. 

Preserved color pattern.—In 70% ethyl 
alcohol: peritoneum pale, stomach and in- 
testine pale. Dark mark from ventral margin 
of eye ending near posterior edge of pre- 
maxilla, two dark lines from posterior mar- 
gin of eye, upper to posttemporal, lower 
onto opercle ending in a ventrally directed, 
short mark; dark spot ocellus-like on side 
just behind opercle flap, with about five 
smaller spots positioned anteriorly and ven- 
trally close to large spot; other melano- 
phores on pectoral-fin base and abdomen, 
scattered diffuse spots on caudal peduncle 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


behind second dorsal and above anal fin; 
scales above the lateral-line scales outlined 
with melanophores; lower third of snout 
dusky between upper jaw and eye; pelvic 
fins with spine and distal third of rays dark 
to tips; anal fin with dark band on spines 
and first three fin-rays to tip, fourth fin-ray 
mostly dark, distal third of next two fin-rays 
dark, last two fin-rays pale from base to 
tips; caudal fin with tips of rays dark; sec- 
ond dorsal-fin spine and first four fin-rays 
dark to tips, remaining fin-rays pale; first 
dorsal fin with the anterior four spines and 
membranes dark to their tips, remaining 
spines and membranes pale. 

Distribution.—Known from the Indian 
Ocean on the Saya de Malha Bank (Fig. 2). 

Etymology.—The Latin word for fourth, 
quartus in reference to the longest dorsal 
spine. 

Remarks.—The fourth dorsal spine is the 
longest dorsal spine in this specimen. With 
additional characters including a high num- 
ber of pectoral-fin rays, rounded caudal fin 
and preserved color pattern, this species can 
be placed within the broad limits of sub- 
genus Jaydia revised by Gon (1996). Apo- 
gon quartus is the first species of this sub- 
genus to be found on the extensive isolated 
shallow banks the Central Indian Ocean. 
Most of the species treated by Gon (1996) 
have a continental distribution pattern. Only 
a few specimens of Apogon hungi Four- 
manior & Nhu-Nhung, 1965, and Apogon 
smithi (Kotthaus, 1970) have been reported 
from insular localities (Gon, 1996). 

Apogon quartus appears to be most 
closely related to Apogon poecilopterus Cu- 
vier in Cuvier & Valenciennes, 1828, Apo- 
gon carinatus Cuvier in Cuvier & Valen- 
ciennes, 1828, and Apogon queketti Gil- 
christ, 1903. The new species differs from 
A. queketti and A. carinatus by not having 
an ocellus in either dorsal fin. A. poecilop- 
terus has more predorsal scales (4—5) and a 
darkish gill chamber. The ocellus-like spot 
on the body is unique to A. quartus among 
these species. 

Apogon argyrogaster is unique among 


VOLUME 113, NUMBER 1 


40° 60° 


Arabian Sea 


Lakshadweep Is. 


Africa 


Mauritius 


Madagasgar 


Maldive ‘3 
Is. 2 


80° 100° 


Re Bay of Bengal age 
“3 Andaman 


vt ay 
Sri acess ‘s 
Lanka S. 


Jawa Sea 
26 
Cocos (Keeling) ; . 
Is. . Christmas 


~ Rodrigues 


* Apogon quartus 
@ Apogon regani 
@ Apogon gardineri 


40° 60° 
Fig... 2. 


species of Apogon with its large ventral or- 
gans (probably bioluminescent) extending 
on each side of the body from the breast 
area forward into the lower part of the 
mouth and backward to past the origin of 
the anal fin. Gon & Allen (1998) recently 
described Apogon photogaster with an ex- 
ternally similar bioluminescent system to A. 
argyrogaster but made no comparisons. 
Their name appears to be a synonym of A. 
argyrogaster. They placed A. photogaster 
in Jaydia. Apogon argyrogaster has faint 
bars on the side of its body, two predorsal 
scales, 8—9 well developed gill rakers in ad- 
dition to the luminous organs. All are char- 
acters differing from A. guartus. 

Apogon melanopus, with nine anal rays 


80° 100° 120° 


Distribution of collection sites for Apogon quartus, Apogon gardineri, and Apogon regani. 


and an emarginate caudal fin unlike all oth- 
er Jaydia species, is a member of Jaydia 
with the fourth dorsal spine the longest and 
16-17 pectoral-fin rays. This species was 
described by Allen (1985) as A. fuscovatus. 
Allen & Morrison (1996) later synony- 
mized Allen’s species when describing an- 
other new species, A. fuscomaculatus, 
which they compared with Apogon striatus 
(Smith & Radcliffe in Radcliffe 1912). 
Apogon melanopus and A. fuscomaculatus 
differ in having higher well-developed gill 
rakers (15—16) and lower well-developed 
gill rakers (8—9) respectively and general 
color pattern (no ocellus-like spot in either 
species) from A. quartus. 

Paratypes of A. tchefouensis from the 


254 


East China Sea examined by me have no 
discernable color patterns because of the 
strong brown stain on both specimens. Fang 
(1942) described the color in alcohol as 
5 uniformément brun-noiratre.”’ Fang 
(1942) described the preopercle as “Bord 
libre du préopercule nettement denticulé 
avec des dents assez fortes surtout a son 
angle, mais plus fines a la partie supérieure 
du bord postérieur.’? The condition of the 
preopercle with a smooth ridge and prom- 
inent serrations, stronger at the angle and 
finer ones on the posterior edge and ventral 
edge, suggest a different group, probably 
type B (but serrations more like type A on 
edges) of Gon’s categories than with the 
group (type C) containing A. quartus. The 
posttemporal is serrated. Total gill raker and 
rudiment count on the first gill arch is 18, 
well-developed rakers 14 (3+2 on upper 
arch, 12+1 on lower arch) of the larger 
paratype. The smaller paratype is in poor 
shape and has a badly damaged head. The 
pectoral ray count was 15 on one side of 
both paratypes and 14 for the other side for 
the larger paratype. Palatine teeth are in one 
to two short rows for the larger paratype. 
Palatine teeth of the lectotype of Apogon 
lineatus Temminck & Schlegel, 1842, are 
in 4 short rows like the holotype of Apogon 
striatodes Gon, 1996, (3—4 short rows), and 
the holotype A. striatus (3—4 short rows) 
but not Apogon novaeguinae Valenciennes, 
1832 with one row. Additional information 
about the color of the peritoneum and in- 
testine from the holotype and paratypes 
may help determine whether A. tchefouen- 
sis 1S a junior synonym of either A. lineatus 
or A. striatus or the senior synonym of A. 
striatodes. The known distribution of the 
two species recognized by Gon (1996) in 
the East China Sea region do not appear to 
overlap, but he suggests the possibility for 
Taiwan. If distributional patterns are accu- 
rate, then the larger paratype of A. tche- 
fouensis should be identified as A. lineatus. 
What ever the eventual status of A. tche- 
fouensis, the paratypes can be distinguished 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


from A. quartus by having serrated preoper- 
cle and posttemporal. 

Fowler & Bean (1930) compared Apo- 
gonichthys melanopterus, their replacement 
name for Amia melas Fowler, 1918, with 
Apogon carinatus. Fowler (1918) first re- 
lated his new species to Apogon nigricans 
Day, 1875, a nominal species related to, if 
not the same as Apogon melas Bleeker, 
1848, with a forked caudal fin, a trace of a 
dark spot on the midbase of the soft dorsal 
fin and darkish pelvic and vertical fins. In 
his original description, Fowler noted that 
the holotype of Amia melas has the fourth 
dorsal spine longer than the third. My ob- 
servation of the holotype agrees with Fowl- 
er’s statement. However, the third spine is 
noticeably stronger (thicker) than the fourth 
spine unlike other species of Jaydia. Apo- 
gon melanopterus 1s consistent in some oth- 
er characters with Jaydia: gill-raker count 
(3+1-11+3), some serrations on the pre- 
opercle edge and a rounded? (damaged) 
caudal fin, but has 14 pectoral-fin rays ver- 
sus the common 15-17, rarely 14 or 18 pec- 
toral-fin rays of Jaydia. This species differs 
from A. quartus in preserved color pattern 
(no ocellus-like spot; dorsal, pelvic, anal 
and caudal fins all blackish), predorsal 
scales (3), strong third dorsal spine and 14 
pectoral rays. I cannot with certainty place 
A. melanopterus within Jaydia as treated by 
Gon. 

Relationships.—The basis for the pro- 
posed monophyly of Jaydia will need to be 
re-examined because some of Gon’s (1996) 
diagnostic features are present in other spe- 
cies of Apogon. For example, Apogon hoev- 
enii Bleeker, 1854d, a more ‘typical’ Apo- 
gon with an emarginate caudal fin, serrated 
preopercular edge and a low pectoral ray 
count (12), has the fourth dorsal spine var- 
iably as its longest. The holotype of A. me- 
lanopterus has the fourth dorsal spine as the 
longest and a robust thicker third dorsal 
spine. One undescribed species and a relat- 
ed species, Apogon moluccensis Valenci- 
ennes, 1832, have the fourth dorsal spine as 
the longest in nearly all adults but not in all 


VOLUME 113, NUMBER 1 


small individuals. The third spine in these 
two species is about as strong as the fourth 
spine, and both have forked caudal fins. The 
undescribed species has a darkish roof of 
the mouth and gill arches reminiscent of A. 
poecilopterus. An unidentified species of 
Apogon, either new or a juvenile of a spe- 
cies in the Apogon diversus (Smith & Rad- 
cliffe in Radcliffe, 1912) group, has strong 
serrations on the preopercular ridge and 
edges, serrated infraorbitals, a strongly fork 
caudal fin and fourth dorsal spine as the 
longest spine. Characters present in A. me- 
lanopus and the above species suggest that 
an expanded review is needed. A solid basis 
for the subgenus founded on the shape of 
the caudal fin, the length of the fourth dor- 
sal spine, strength of the third dorsal spine, 
high pectoral-fin ray counts, supporting 
structures for the dorsal fin and other char- 
acters elucidated by Gon may grade with 
the examination of more species. Perhaps 
some of the other species of Apogon dis- 
cussed here eventually may be shown to be 
closer to this species complex than to other 
Apogon. Those species treated by Gon that 
have bioluminescent systems are more like- 
ly to be monophyletic and may be worthy 
of the recognition as the subgenus Jaydia. 
There is, as yet, no clear resolution about 
whether these two subgenera represent 
monophyletic groupings within Apogon. 


Status of Apogon heraldi 
Fig. 3 


Apogon heraldi (Herre, 1943) was de- 
scribed in Mionurus, a misspelling of Mio- 
norus Krefft, 1868, as having nine anal 
rays. Herre compared it with two unrelated 
nominal species with six first dorsal spines, 
Apogon bombonensis (Herre, 1925) and 
Apogon mydrus (Jordan & Seale, 1905). 
There is enough information in Herre’s de- 
scription to determine that this species 
should be compared with those treated by 
Gon (1996), but not enough to place it with 
certainty. Herre did not give a figure of the 
type. A redescription of the type, with a fig- 


255 


ure 1s needed to provide comparison with 
the new species and with other related spe- 
cigs, 

Material examined.—Mionurus heraldi 
Holotype: SU 38263 (104.5); Philippines, 
Luzon I., Ragay Gulf, X-ray. 

Comparative material.—See A. quartus. 

Description.—For general body shape 
see Fig. 3, Range of proportions (as per- 
centage of standard length): greatest body 
depth 39.4; head length 42.0; eye diameter 
9.8; snout length 8.5; bony interorbital 
width 9.6; upper-jaw length 20.1; caudal- 
peduncle depth 16.7; caudal-peduncle 
length 20.2; first dorsal-spine length 2.4; 
second dorsal-spine length 7.1; third dorsal- 
spine length 10.9; fourth dorsal-spine 
length 14.1; spine in second dorsal fin 11.3; 
first anal-spine length 2.8; second anal- 
spine length 10.9; pectoral-fin length 25.4; 
pelvic-fin length 26.7. 

Dorsal fin VII-—I,9; anal fin II,8; pectoral 
fin 16-16; pelvic fin I,5; principal caudal 
rays 9 + 8; number of pored lateral-line 
scales unknown; number of transverse scale 
rows above lateral line unknown; number 
of transverse scale rows below lateral line 
unknown; median predorsal scales 4 or 5; 
number of circumpeduncular scale rows un- 
known. Gill rakers, upper arch 2 rudiments 
1 well developed raker, lower arch | rudi- 
ment, 10 well-developed rakers, 11 well de- 
veloped and 14 total. 

Villiform teeth in single row anteriorly, 
slightly larger grading into wide band at 
sides of premaxilla; three rows anteriorly 
with one to two rows at sides of dentary, 
interior row slightly larger from anterior to 
side; one row on palatine; two rows on vo- 
mer; none on ectopterygoid, endopterygoid 
or basihyal. 

Vertebrae 10 + 14. Five free hypurals, 
one pair of slender uroneurals, three epur- 
als, a free parhypural. Three supraneurals, 
two supernumerary spines on first dorsal 
pterygiophore. Basisphenoid present. Su- 
pramaxilla absent. Posttemporal smooth on 
posterior margin. Preopercular ridge 
smooth, vertical margin smooth, horizontal 


256 


The holotype of Mionurus heraldi, SU 38263, 104.5 mm SL from the Philippine Islands, Luzon I., Ragay Gulf. 


Fig. 3. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


margin with smooth undulations. Infraor- 
bitals with crenulate to sharp edges. Scales 
ctenoid, many missing from body. Caudal 
fin slightly rounded. No indication of bean- 
like bioluminescent organs associated with 
intestine near anus. 

Life colors.—Unknown. 

Preserved color pattern.—In 70% ethyl 
alcohol: peritoneum silvery; intestine and 
stomach mostly pale with scattered mela- 
nophores; second dorsal fin with dark stripe 
near base, a narrow pale area above on first 
few rays to about fifth ray, more distally the 
fin-ray membranes dusky to dark with sec- 
ond narrow pale area, then fin rays dusky 
to tips; wide darkish stripe basally in anal 
fin, the membranes pale distally; spinous 
dorsal dusky distally, dark between the sec- 
ond and fifth spines, pale basally; pelvic 
and pectoral fins pale; gill chamber and gill 
arches dusky with melanophores; caudal fin 
membranes dusky with no indication of a 
darker margin. 

Remarks.—Apogon heraldi has the 
fourth dorsal spine longest and can be 
grouped with Gon’s type C preopercle pat- 
tern of Jaydia. Herre (1943) described the 
color of A. heraldi in alcohol as “‘... pale 
brown, with a more or less evident vertical 
dark-brown bar under each scale; there is a 
blackish spot on the upper half of the spi- 
nous dorsal, and a wide dark-brown cross- 
band on the second dorsal and the anal, 
which are otherwise clear; all but the basal 
part of the caudal is dusky; the pectorals 
and ventrals are clear’. Herre’s description 
of the teeth as “The very small conical 
teeth are in a single row in the lower jaw,” 
and lack of comment about color in the oral 
cavity would suggest that this specimen is 
not A. poecilopterus. However, my re-ex- 
amination of the holotype indicates that the 
dentition and the color pattern in the gill 
chamber are consistent with that of A. poe- 
cilopterus. The holotype has 14 gill rakers 
plus rudiments on the first gill arch, 2+1 on 
the upper arch and 10+1 on the lower arch, 
about 4 or 5 predorsal scales and 16 pec- 
toral rays, all consistent with A. poecilop- 


VOLUME 113, NUMBER 1 


terus. Herre (1943) reported on material of 
A. ellioti [=truncatus] and A. poecilopterus 
from the Ragay Gulf and A. striatus from 
Manila Bay in the same paper. Apogon her- 
aldi is considered a synonym of A. poeci- 
lopterus. The type is a ripe female. 


Apogon gardineri Regan, 1908 
Figs. 2 & 4 


Material examined.—Holotype: BMNH 
1908.3.23.93; 39.7 mm SL; Cargados Ca- 
rajos; 55 m. Other material: BPBM 24775; 
(41—42); Mauritius, off Flic en Flac; J. E. 
Randall et al.; 7 April 1979; 57 m; color 
photo; x-ray. 

Comparative material.—Apogon nigri- 
pinnis Syntype: MNHN 8694; 63.4 mm SL; 
India, Pondicherry; Leschenault. Mozam- 
bique: Delagoa Bay: RUSI 3149; (60-76); 
x-ray. RUSI 3151; (67); 1938; x-ray. RUSI 
3148; 4(42—84); x-ray. RUSI 3150; 4(68-— 
74); x-ray. Palma: RUSI 3152; (69); 1 Aug 
1951; x-ray. Aldabra I.: RUSI 3153; (49-— 
53); 14 Nov 1954; x-ray. Apogon striatus 
Syntypes: MNHN 1973-41; 3(16.8—26.2); 
Madagascar, Nosy-Bé; x-ray. Apogon suezii 
Hotetype: MNHN 5137; (50:6); Egypt, 
Suez; Letourneux. Apogon taeniatus Syn- 
types: MNHN 8693; (55.3-—57.4); Red Sea; 
Ehrenberg. Kenya: RUSI 3106; (85); Mal- 
medi; 2Oct 1952; x-ray. RUSI 3111; (94); 
Shimoni; 1950; x-ray. RUSI 3107; (77-83); 
Shimoni; Nov 1952; x-ray. RUSI 3112; 
3(15—66); Shimoni; May 1951; x-ray. RUSI 
3109; (76); Feb 1952; x-ray. Mozambique: 
RUSI 3105; (79); Pinda reef; 1950; x-ray. 
RUSI 3114; (49); Mocimboa de Praia; 3 
Sep 1951; x-ray. RUSI 3110; (81); Inham- 
bane; Sep 1948; x-ray. RUSI 3108; 3(59-— 
82) Jun 1950; x-ray. Aldabra: RUSI 3113; 
(55); Nov. 1954; x-ray. ANSP 63868; (86); 
South Africa. Apogon thurstoni Holotype: 
BMNH 1889.8.17.2; (55.5); India, Madras; 
Day. Apogon timorensis Holotype: RMNH 
5583; (60.8); Timor. QM 1.820; 3(23-31); 
Australia, Darnley I. QM I. 8506; (28); 
Australia, Queensland, Green I. Apogon- 
ichthyoides fraxineus Holotype: RUSI 356; 


pmo 


(57.7); Mozambique, Pinda Reef; 3 Sep 
1956; x-ray. Paratypes (all x-rayed): Moz- 
ambique: RUSI 746; 8(42—65); Pinda Reef; 
2 Sep 1956..RUSI 751; (67); Pinda Reef; 
Sep 1956. RUSI 762; (37-69); Pinda Reef; 
23 Sep 1956. RUSI 758; (48-55); Delagoa 
Bay. RUSI 7409; (33); Ibo I.; Aug 1951. 
Zanzibar: RUSI 757; (41); 20 Sep 1952. 
RUSI 754; (54); 9 Sep 1952. RUSI 748; 
3(52-57); 4 Sep 1952. RUSI 759; (23-24); 
Seychelles, Mahe I.; Sep 1954. 

Diagnosis.—A species of the subgenus 
Ostorhinchus with two saddles on the body 
and one on the caudal peduncle, no cheek 
mark, a pale stomach, intestine and perito- 
neum, 14 pectoral-fin rays, 2 predorsal 
scales and 14—15 total gillrakers. 

Description.—For general body shape 
see Fig. 4. Range of proportions (as per- 
centages of standard lengths), holotype in 
parentheses: greatest body depth 38-39 
(37.3); head length 42 (42.3); eye diameter 
15 (13.8); snout length 8.8—9.5 (9.3); bony 
interorbital width 8.4—8.8 (8.6); upper-jaw 
length 18—20 (19.6); caudal-peduncle depth 
14-16 (14.9); caudal-peduncle length 21-— 
24 (19.4); first dorsal-spine length 3.0—3.6 
(5.0); second dorsal-spine length 8.3—9.5 
(9.3); third dorsal-spine length 19-20 
(18.6); fourth dorsal-spine length 16—19 
(18.6); spine in second dorsal fin 13-15 
(13.8); first anal-spine length 3.1—4.1 (3.3); 
second anal-spine length 12—15 (11.8); pec- 
toral-fin length 21—24 (20.1); pelvic-fin 
length 23-27 (23.4). 

Dorsal fin VII-—I,9; anal fin II,8; pectoral 
fin 14—14; pelvic fin 1,5; principal caudal 
rays 9 + 8; simple pored lateral-line scales 
24; transverse scale rows above lateral line 
2; transverse scale rows below lateral line 
5—6; median predorsal scales 2; circumpe- 
duncular scale rows 12 (5+2+5). Total gill- 
rakers 14-15 (14), well developed 9-10 
(9), upper arch 2+1, lower arch 8—9+2-—3 
(8+°3): 

Villiform teeth in a wide band on the pre- 
maxilla and dentary; two rows on the pal- 
atine and vomer; none on ectopterygoid, 
endopterygoid or basihyal. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 4. Apogon gardineri, BPBM 24775, 42 mm SL, from Mauritius, off Flic en Flac by J. E. Randall. 


Supramaxilla absent. Posttemporal 
smooth on posterior margin. Preopercle ser- 
rate on vertical and horizontal margins, 
smooth on ridge. Infraorbitals smooth, pos- 
terior ones with crenulations. Scales cte- 
noid. Caudal fin weakly emarginate. 

Life colors.—Based on a Ektachrome 
transparency of the Mauritian specimen the 
color pattern is as follows: Iris yellow; head 
(without dark or pale spots, bars or stripes) 
and body with a light tannish background; 
a small brownish spot at the beginning of 
pored lateral-line scales; a darker brownish 
bar beginning at the base of first five dorsal 
spines and extending to about level of pec- 
toral fin, as a chevron, angled posteriorly; 
a second fainter bar or blotch under last 
four or five soft rays of second dorsal fin 
reaching to or just below pored lateral line; 
a full dark brownish mark (spot or bar-like) 
on caudal peduncle; first dorsal fin with a 
dark brownish bar extending over first five 
spines and contiguous with bar on the body, 
membrane behind the fifth to seventh spines 
pale; second dorsal, anal and caudal fins 
pale without spots, bars or stripes; pelvic fin 
pale except for the last third of the first two 
soft rays. 

Preserved color pattern.—In 70% ethyl 
alcohol: The holotype has no color remain- 
ing. Regan (1908) reported: ‘‘Faint traces 


of 3 dark vertical bars, the first below the 
spinous dorsal, the second below the soft 
dorsal, the third on the caudal peduncle. 
Spinous dorsal blackish except posteriorly; 
ventrals blackish at the tips’’. The Mauri- 
tian specimens have a pale intestine, with a 
few melanophores on the stomach, perito- 
neum pale; first dorsal fin dark, with a mark 
extending onto the body to about even with 
the pectoral fin; a second saddle extends 
downward from the posterior part of the 
second dorsal fin base; a diffuse blotch on 
the caudal peduncle; a faint cheek mark and 
a spot behind the eye. 

Distribution.—Known only from the 
Mascarene Plateau. 

Remarks.—Regan (1908) suggested that 
A. gardineri was related to Apogon ban- 
danensis Bleeker, 1854a. Apogon banda- 
nensis has more gill-raker (total gillrakers 
25-29) and has black stomach and intestine. 
Smith (1961) synonymized A. gardineri 
with Apogon nigripinnis Cuvier in Cuvier 
& Valenciennes, 1828. The holotype of A. 
nigripinnis is a different species with 16 
pectoral-fin rays, 16 total rudiments and gill 
rakers, 12 well-developed rakers (3+2 up- 
per arch, 10+1 lower arch), two predorsal 
scales, a deeper body (depth 42.9% stan- 
dard length), soft dorsal and anal fins black- 
ish and a uniform brownish body without 


VOLUME 113, NUMBER 1 


an obvious ocellated spot on the side of the 
body (faded?) or basicaudal spot (faded?). 
The holotype of Apogon thurstoni Day, 
1888, shares all of the above-listed char- 
acters of A. nigripinnis, 16 pectoral-fin rays, 
16 total rudiments and gill rakers, two pre- 
dorsal scales, a body depth of 43.4% of 
standard length, but has 10 well developed 
gill rakers (2+2 upper arch, 8+4 lower 
arch) and an obvious ocellated spot on side 
about midline. All of the Southern African 
material specimens here as A. nigripinnis 
have an ocellus on the side, but otherwise 
have meristics and body depth similar to the 
type: 15-16 pectoral-fin rays, 16—18 total 
rudiments and gill rakers, 11—12 well-de- 
veloped rakers (2—3+2 upper arch, 9- 
10+ 1—4 lower arch), two predorsal scales, 
a body depth of 42—48% of standard length. 

Apogon taeniatus Cuvier in Cuvier & 
Valenciennes, 1828, appears closely related 
to A. nigripinnis and may also be confused 
with A. gardineri. However, A. taeniatus 
variably has an ocellated spot within the 
first dorsal saddle above the pectoral fin, a 
small basicaudal spot, faint stripes on body, 
16—19 total rudiments plus gill rakers, 9— 
12 well-developed rakers (2—4+1—2 upper 
arch, 8—10+3-—5 lower arch), 14—15 pecto- 
ral-fin rays, 3 predorsal scales and a greater 
body depth of 40—45% standard length. 
The holotype of Apogon suezii Sauvage, 
1883, shares with A. taeniatus an ocellated 
spot above the pectoral fin, 17 total rudi- 
ments and gill rakers, 12 well-developed 
rakers (2+2 upper arch, 10+3 lower arch), 
15 pectoral-fin rays, three predorsal scales 
and a body depth of 40.9% standard length, 
but it has a small basicaudal spot within a 
darkish bar. 

The low gill-raker count, body shape and 
color pattern of A. gardineri may cause 
some confusion with Apogon timorensis 
Bleeker, 1854b and two of its synomyms 
Apogon fraxineus (Smith, 1961) and Apo- 
gon striatus Fourmanior & Crosnier, 1964, 
which differs in having a thin, dark cheek 
line from the eye onto the preopercle, a 
black stomach and intestine, 15—17, usually 


259 


16 pectoral rays and fewer developed (6—8) 
and more rudiments (4-7) on the lower 
arch. Fourmanoir and Crosnier’s name is an 
unavailable name as a secondary homonym 
of Apogon striatus (Radcliffe in Smith & 
Radcliffe, 1912). 

Apogon gardineri is found in deeper wa- 
ter. It does not appear to have been reported 
from any of the shallow-water collections 
from Mauritius, Cargados Carajos or the 
Seychelles in the past several decades. 
Many of the species in this complex appear 
to have continental distributions, for ex- 
ample, Gon (1986, map 1). Apogon gardi- 
neri represents an insular species endemic 
to the Mascarene Plateau. 


Apogon regani Whitley, 1951 
Pies) 2 2ac55 


Synonyms: Apogon punctatus Regan, 
1908, preoccupied by Apogon punctatus 
Klunzinger, 1880. 

Material examined.—Syntypes: Apogon 
punctatus BMNH — 1908.3.23.86-—89; 
4(31.9—-65.4); Cargados Carajos; 39-55 m. 
Other material: BPBM 35475; (45); Sey- 
chelles, N. of Aride I., 4°10’S, 55°44’E; 
R/V Tyro, Station 714, 2.4-m Agassiz trawl; 
J. van der Land et al.; 19 Dec 1992; 55 m, 
color photo; USNM 307706; (44); Indian 
Ocean, Saya de Malha Bank, 10°16'00’S, 
61°09’00"E; Vityaz Cr. 17; Sta 2810; 8 Jan 
1989; 50-70 m; x-ray. 

Comparative material.—See A. gardi- 
neri. 

Diagnosis.—A species of Apogon in the 
subgenus Ostorhinchus with 15-17 pectoral 
rays, 4—6 rows of small spots on the body, 
pale stomach, intestine and peritoneum and 
14—16 total gill rakers. 

Description.—For general body shape 
see Fig. 5. Range of proportions (as per- 
centages of standard lengths, syntypes in 
parentheses): greatest body depth 44 (40.7— 
43.0); head length 37—42 (40.4—42.5); eye 
diameter 10—13 (10.8—12.5); snout length 
10-11 (9.4—10.6); bony interorbital width 
9.6-9.8 (8.7—-9.7); upper-jaw length 23 


260 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Bige 5: 


(20.8—22.6); caudal-peduncle depth 17-19 
(15.9-17.0); caudal-peduncle length 20—22 
(18.5—20.8); first dorsal-spine length 3.7— 
3.9 (3.0—4.1); second dorsal-spine length 
10 (6.6—8.1); third dorsal-spine length 22— 
23 (18.5—21.6); fourth dorsal-spine length 
20—22 (17.9—20.3); spine in second dorsal 
fin 16-18 (13.3-—14.5); first anal-spine 
length 4.1—5.0 (3.0—4.1); second anal-spine 
length 15-16 (12.0—13.3); pectoral-fin 
length 21—23 (21.9); pelvic-fin length 29— 
33° (25.7=27.7): 

Dorsal fin VII—I,9; anal fin 1,8; pectoral 
fin 15-15, 15-16, 16—16, or 17-17; pelvic 
fin I,5; principal caudal rays 9 + 8; pored 
lateral-line scales 24—25; transverse scale 
rows above lateral line 2; transverse scale 
rows below lateral line 6—7; median pre- 
dorsal scales 2—3; circumpeduncular scale 
rows 12(5+2+5). Total gillrakers 14-16, 
well-developed 10—11, upper arch 2+2 and 
lower arch 8—9+ 1-3. 

Villiform teeth in a band on the premax- 
illa, the outer row slightly larger; several 
rows anteriorly, two rows on side of the 


Apogon regani, BPBM 35475, 45 mm SL, from the Seychelles, N. of Aride I. by J. E. Randall. 


dentary; one to two rows on the palatine, 
two rows On vomer; none on ectopterygoid, 
endopterygoid or basihyal. 

Vertebrae 10 + 14. Five free hypurals, 
one pair of slender uroneurals, three epur- 
als, a free parhypural. Three supraneurals, 
two supernumerary spines on first dorsal 
pterygiophore. Basisphenoid present. Su- 
pramaxilla absent. Posttemporal serrate on 
posterior margin. Preopercle serrate on ver- 
tical and horizontal margins, smooth on the 
ridge. Infraorbitals smooth, the second 
crenulate and third with a single spine. 
Scales ctenoid. Caudal fin truncate. 

Life colors.—Based on an Ektachrome 
transparency of the Seychelles specimen the 
color pattern is as follows: Iris pale; head 
and body with a pale tannish background 
on the upper parts; a dark brown oblique 
line behind the eye; head silvery below eye 
level; body silvery below the pored lateral 
line on body to about the end of anal-fin 
base, faint brownish spots on some, but not 
all scales above the pored lateral line; larger 
brownish spots on each scale below the 


VOLUME 113, NUMBER 1 


pored lateral line, forming about five lines 
of spots, reducing to two lines on the an- 
terior portion of the caudal peduncle; a 
brownish area on the dorsum below the first 
four or five spines; another mark on dorsum 
below the fifth soft dorsal ray; caudal and 
pectoral fins pale; first dorsal dusky and 
whitish with a spot near the base of the 
fourth to sixth spines; second dorsal fin 
dusky; anal fin with a dark brownish edge 
to the second anal spine and first soft ray, 
the rest of fin dusky, pelvic fins dark 
brownish except the fifth soft ray pale. 

Preserved color pattern.—In 70% ethyl 
alcohol: the syntypes with three to four 
rows of small spots below the lateral line 
on side of body. The pelvic fins and second 
dorsal fin blackish. Stomach, intestine and 
peritoneum pale. The other two specimens 
with six rows of small spots on side of body 
to caudal peduncle, one row above the lat- 
eral line; three marks behind eye, a narrow 
oblique cheek mark, a mark between the 
eye and preopercle just below the mid-line 
of the head, and an oblique mark past the 
top of the preopercle; all vertical fins dark- 
ish, caudal fin pale. 

Distribution.—Known only from the 
Mascarene Plateau. 

Remarks.—Whitley (1951) created the 
replacement name, A. regani for A. punc- 
tatus Regan, 1908, which is preoccupied by 
Apogon punctatus Klunzinger, 1880, a pri- 
mary homonym. Klunzinger’s species is 
presently placed in Vincentia, a mostly 
warm temperate species group from Austra- 
lia and Tasmania. According to the Inter- 
national Commission on Zoological No- 
menclature (1985: Art 59b) Regan’s hom- 
onym is to be treated as permanently re- 
jected. 

Regan suggested a relationship with A. 
nigripinnis. Smith (1961) synonymized A. 
punctatus with A. nigripinnis. However, A. 
regani differs from all known species in the 
A. nigripinnis group by the presence of 
small spots, one per scale, on the body. A. 
regani represents an insular species in this 


group. 


261 


This species is found in deeper water. 
Randall & Egmond (1994) reported A. re- 
gani as A. punctatus from the Seychelles 
and provided a color photograph without 
description. It does not appear to have been 
reported from any of the shallow-water col- 
lections from Mauritius or Cargados Cara- 
jos in the past several decades. Apogon re- 
gani represents the second insular species 
endemic to the Mascarene Plateau in the A. 
nigripinnis complex. 


Acknowledgments 


For the loan of material, the use of mu- 
seum facilities and aid in curatorial pro- 
cesses I thank: Arnold Y. Suzumoto 
(BPBM), David Catania, Tomio Iwamoto, 
William N. Eschmeyer (CAS), Susan L. 
Jewett, David G. Smith, and Jeffery T. Wil- 
liams (USNM). Jeffery T. Williams helped 
with photography and David G. Smith pro- 
vided x-ray photography of some speci- 
mens. Many other colleagues over the past 
two decades at various museums have pro- 
vided valuable assistance in my studies of 
the Apogonidae. Leonard P. Schultz funds 
were provided by Victor G. Springer 
(USNM) for several study trips to the 
Smithsonian. John E. Randall provided col- 
or transparencies of A. gardineri and A. re- 
gani. Ofer Gon and John E. Randall re- 
viewed an earlier draft of this manuscript. 
Two anonymous reviewers provided helpful 
comments. 


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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


113(1):264—268. 2000. 


A new species of Pogonophryne 
(Pisces: Perciformes: Artedidraconidae) from East Antarctica 


Richard R. Eakin and Arcady V. Balushkin 


(RRE) Department of Life Sciences, University of New England, Westbrook College Campus, 
Portland, Maine 04103; (AVB) Zoological Institute, Russian Academy of Sciences, 
Universitetskaya nab. 1, St. Petersburg, 199034 Russia 


Abstract.—A new species of notothenioid fish, Pogonophryne squamibar- 
bata, is described from East Antarctica. This dorsally-spotted species belongs 
to the ‘“‘mentella’’ group of the genus and is characterized by having a relatively 
short mental barbel (16% SL) with bluntly palmate, overlapping, scale-like 
processes on its terminal expansion, a wide interorbital region (6.9% SL), and 
a long opercular-subopercular distance (19.2% SL). 


German research cruises of the R/V Po- 
larstern in Antarctic waters have yielded 
several new species of the artedidraconid 
genus Pogonophryne (Eakin 1987, Balush- 
kin & Eakin 1998, Eakin & Balushkin 
1998). The “‘mentella’’ group is the most 
speciose of the genus, comprising at least 
10 species, three of which have been re- 
cently described (Balushkin & Eakin 1998, 
Eakin & Balushkin 1998, Eakin & Eastman 
1998). This paper continues our descrip- 
tions of new species of Pogonophryne in 
the Zoological Museum collections at Ham- 
burg University. 

Measurements and counts follow those of 
Balushkin & Eakin (1998) and Eakin & 
Eastman (1998). Opercular-subopercular 
distance is measured from the posterior 
edge (hook) of the opercular bone to the 
anterior edge of the subopercular bone. 
Measurements are presented as percentages 
of standard length unless otherwise indicat- 
ed. ISH refers to the Zoological Museum 
collections (formerly Institut fiir Seefischer- 
ei) currently maintained at Hamburg Uni- 
versity (ZMH). 


Pogonophryne squamibarbata, 
new species 
Figs. 1-2 
Holotype.—ISH 65-1991; male, 147 mm 
SL; R/V Polarstern station 211, East Ant- 


arctica (69°58.9'S, 05°8.4’E), 651-742 m; 
10 Mar 1991. 

Diagnosis.—This species of Pogono- 
phryne belongs to the “‘mentella”’ group. 
The relatively short (16% SL) mental bar- 
bel has a terminal expansion composed of 
bluntly palmate, flattened processes ar- 
ranged in an overlapping, scale-like pattern. 
This species also has a wide interorbital re- 
gion (6.9% SL) and a long opercular-sub- 
opercular distance (19.2% SL). 

Description.—Body robust anteriorly, ta- 
pering to low, narrow caudal peduncle. 
Head length 43.5; posttemporal ridges not 
well developed (upper limb of posttemporal 
bone not arched upward to form a promi- 
nent curved ridge), depth of head at this 
level 19.7; head width at preoperculars 
37.4. Body depth at second dorsal fin origin 
17.3; body depth at anal fin origin 13.6; 
body width at anal fin origin 10.5; caudal 
peduncle depth 6.5. Snout broadly rounded 
in dorsal view; internostril distance 9.2; 
snout length 8.6; eye filling orbit; diameter 
of orbit 9.9. Interorbital region wide, bony 
measurement 6.9. Opercular-subopercular 
distance long, 19.2. Postorbital length of 
head 23.7. Jaw width at posterior ends of 
maxillaries 25.5. Upper jaw length 19.3; 
posterior end of maxillary extending slight- 
ly beyond pupil of eye; lower jaw project- 


VOLUME 113, NUMBER 1 


male, 147 mm SL. 


> 


Pogonophryne squamibarbata, holotype, ISH 65-1991 


Fig. 1: 


N 


Nn 


266 


ing beyond upper a distance 7.8 of head 
length. Tongue long, extending anteriorly 
slightly in front of tip of upper jaw. Upper 
jaw with two rows of teeth (largest near 
symphysis) in a broad arc. Lower jaw teeth 
in two rows near symphysis, becoming uni- 
serial posteriorly (nine large teeth on both 
sides) in a V-shaped pattern. Mental barbel 
(Fig. 2) rather short (16% SL) and with a 
terminal expansion composed of bluntly 
palmate, flattened processes arranged in an 
overlapping, scale-like pattern and continu- 
ing onto stalk; barbel length 16.0; terminal 
expansion 34.0 of barbel length and slightly 
wider than stalk. Anterior gillrakers on first 
arch 2 + 0 + 7 = 9; posterior gillrakers O 
+ 1+ 7 =s3tonla7 

First antedorsal distance 37.4; second an- 
tedorsal distance 46.3; anteanal distance 
64.6. Length of second dorsal fin base 52.4; 
length of anal fin base 33.0; length of cau- 
dal fin 25.3;. length of pectoral fin 28:7: 
width of pectoral fin base 10.5; length of 
pelvic fin 20.4. Interdorsal distance 7.5. 
First dorsal fin with two spines of about 
equal length, 9.2. Second dorsal fin with 27 
rays; length of longest (fifth) ray 15.3. Ratio 
of longest first dorsal fin spine to longest 
second dorsal fin ray 0.60. Anal rays 17. 
Pectoral rays 20. Upper lateral line with 
about 26 (left) and 27 (right) pores (tubular 
scales), ending under the twenty-first ray of 
the second dorsal fin. Middle lateral line 
with about 11 tubular scales (13 pores) on 
the left side and 11 tubular scales on the 
right side; tubular scales originate under the 
eighteenth ray of the second dorsal fin. Ce- 
phalic lateral-line pores typical for Pogon- 
ophryne: preoperculo-mandibular canal 
with nine pores; infraorbital canal with sev- 
en pores; supraorbital canals with two nasal 
pores, two interorbital pores, and one un- 
paired coronal pore; temporal canal with six 
pores; and supratemporal canal incomplete 
across occiput (one pore on each side). Ver- 
tebrae 16 + 21 = 37 (second preural ver- 
tebra appears double, being longer than the 
others and possessing two neural and two 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


hemal spines, but is counted as one ele- 
ment). 

Radiographic analysis of skeleton.— 
Number of vertebrae to first pterygiophore 
of first dorsal fin 2; to first pterygiophore 
of second dorsal fin 6; to first pterygiophore 
of anal fin 16. Free caudal vertebrae count- 
ing from last pterygiophore of second dor- 
sal fin 6; counting from last pterygiophore 
of anal fin 6. Four pterygiophores anterior 
to first pterygiophore of second dorsal fin; 
the anterior two support spines of the first 
dorsal fin, while the posterior two are ray- 
less pterygiophores. One pterygiophore an- 
terior to first caudal vertebra under abdom- 
inal section of vertebral column. Four hy- 
pural plates attached to urostyle; parhypural 
(PH) + lower complex hypural (H1 + H2) 
+ upper complex hypural (H3 + H4) + 
minimal hypural (H5). All hypural plates 
except H5 fused to urostyle. Caudal fin with 
21 rays (4 upper procurrent + 13 principal 
+ 4 lower procurrent). Distribution of prin- 
cipal caudal rays with reference to hypural 
plates: 1-6—4—2. Epurals 3, the anterior two 
of which resemble in shape the neural spine 
of the first preural vertebra. One pair of uro- 
neurals. 

Color in alcohol.—Head and body with 
brownish markings dorsally and laterally on 
yellowish background. Markings on head in 
the form of round spots and vermiculations; 
those on body larger and more irregular. A 
broad, dark horizontal marking extends 
along base of anal fin. Ventral surfaces of 
head and body dusky and unmarked. First 
dorsal fin light; second dorsal fin with dark 
oblique stripes. Anal fin light. Caudal fin 
with about six dark vertical stripes. Pectoral 
fins with about six dark vertical stripes. Pel- 
vic fins light. Mental barbel light. 

Etymology.—Named for the overlapping, 
scale-like processes on the mental barbel 
(Latin sqguama: a scale; barb: a beard). 

Comparative notes.—The new species 
belongs to the “‘mentella’’ group of the ge- 
nus which is generally characterized by 
having a well developed mental barbel 
(elongation, complex terminal expansion, 


VOLUME 113, NUMBER 1 267 


Fig. 2. Mental barbel of Pogonophryne squamibarbata, holotype; a, terminal expansion (10); b, detail of 
terminal expansion showing overlapping, palmate processes (X40). 


268 


or both), relatively undeveloped posttem- 
poral ridges (upper limb of posttemporal 
bone not arched upward to form a promi- 
nent curved ridge), and large, sparsely dis- 
tributed, round spots and vermiculations on 
the dorsal and lateral surfaces of the head 
(Balushkin & Eakin 1998). The mental bar- 
bel somewhat resembles that of P. mentella 
in having a terminal expansion composed 
of bluntly palmate, flattened processes. 
However, the barbel of P. squamibarbata is 
much shorter (16% SL) than that of the ho- 
lotype of P. mentella (27.1% SL; Andria- 
shev 1967) and has a relatively longer ter- 
minal expansion (34% of barbel length 
compared to 26.4% of barbel length; mea- 
sured from Andriashev 1967, Fig. 1). The 
scale-like pattern of densely packed, over- 
lapping processes on the mental barbel of 
P. squamibarbata is unlike that of any other 
species of Pogonophryne. This species also 
has a wide interorbital region (6.9% SL) 
and a long opercular-subopercular distance 
(19 29% SE): 


Acknowledgments 


We thank Horst Wilkens of the Zoolog- 
ical Museum, Hamburg University, for his 
kind invitation to examine the ISH collec- 
tions and for his hospitality during our visit 
in June 1997. RRE thanks the University of 
New England/Westbrook College for partial 
financial support from a Bingham Faculty 
Enrichment Grant. AVB is grateful for fi- 
nancial support from the German Scientific 
Society (Grant no. 436 RUS-17/41/97). The 
work was also supported by the Russian 
Foundation for Basic Research (Grant no. 
97-04-48950). We are indebted to Gudrun 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Schulze of the Zoological Museum, Ham- 
burg University, for her help in all aspects 
of our work. We thank Jacob Levenson of 
the University of New England for photo- 
graphing the holotype and Helga Kapp of 
the Zoological Museum, Hamburg Univer- 
sity, for help and use of equipment in pho- 
tographing the mental barbel. We thank An- 
gelika Brandt of the Zoological Museum, 
Hamburg University, for checking collec- 
tion data. We also thank Joseph Eastman of 
Ohio University for critically reading the 
manuscript. 


Literature Cited 


Andriashev, A. P. 1967. A review of the plunder fishes 
of the genus Pogonophryne Regan (Harpagifer- 
idae) with descriptions of five new species from 
the East Antarctic and South Orkney Islands. 
Pp. 389-412 in A. P. Andriashev and P. V. 
Ushakov, eds., Biological results of the Soviet 
Antarctic Expedition (1955-1958), vol. 3. Len- 
ingrad Zoological Institute, Academy of Scienc- 
es of the USSR. 

Balushkin, A. V., & R. Eakin. 1998. A new toad plun- 
derfish Pogonophryne fusca sp. nova (Fam. Ar- 
tedidraconidae: Notothenioidei) with notes on 
species composition and species groups in the 
genus Pogonophryne Regan.—Journal of Ich- 
thyology 38:574—579 (English translation from 
Voprosy Ikhtiologii 38:598—603). 

Eakin, R. R. 1987. Two new species of Pogonophryne 
(Pisces, Harpagiferidae) from the Weddell Sea, 
Antarctica.—Archiv fiir Fischereiwissenschaft 
38:57-74. 

, & A. V. Balushkin. 1998. A new species of 

toadlike plunderfish Pogonophryne orangiensis 

sp. nova (Artedidraconidae, Notothenioidei) 
from the Weddell Sea, Antarctica.—Journal of 

Ichthyology 38:800—803 (English translation 

from Voprosy Ikhtiologii 38:830—833). 

, & J. T. Eastman. 1998. New species of Po- 

gonophryne (Pisces, Artedidraconidae) from the 

Ross Sea, Antarctica.—Copeia 1998:1005-— 

1009. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 
113(1):269—283. 2000. 


A new genus and species of inseminating fish 
(Teleostei: Characidae: Cheirodontinae: Compsurini) from 
South America with uniquely derived caudal-fin dermal papillae 


Luiz R. Malabarba and Stanley H. Weitzman 


(LRM) Departamento de Zoologia - IB, Universidade Federal do Rio Grande do Sul, 

Av. Paulo Gama s/n, 90046-900, Porto Alegre, RS, Brasil, and Museu de Ciéncias e Tecnologia, 
PUCRS, Av. Ipiranga 6681, 90619-900, Porto Alegre, RS, Brazil; (SHW) Division of Fishes, 
Department of Vertebrate Zoology, Smithsonian Institution, Washington, D.C. 20560-0159, 
U.S.A. 


Abstract.—Kolpotocheirodon theloura, a new genus and species of the chei- 
rodontine tribe Compsurini, is described from the uppermost tributaries of the 
rio Sao Francisco and rio Parana in central Brazil. Males of the new species 
bear a caudal-fin organ formed by fin-ray hooks and hypertrophied elongate 
dermal flaps along the fin rays, mostly in the ventral lobe of the caudal fin. 
The largest flap, attached dorsally to the nineteenth principal caudal-fin ray 
(ventral principal fin ray of the ventral caudal-fin lobe), is bordered with a 
series of tabs, and extends dorsally over most of the flaps extending from 
caudal-fin rays 17 and 18. There are also a series of fleshy papillae distributed 
along the border of the remaining ventral caudal-fin ray flaps or along the fin 
rays just dorsal to the flap-bearing fin rays. These tabs and papillae are un- 
known in both other inseminating and all externally fertilizing characids and 
therefore are considered derived. The new species is referred to the Compsurini 
of the Cheirodontinae on the basis of a cladistic diagnosis of these taxa. The 
relationships of K. theloura with other inseminating compsurins bearing mod- 
ified caudal fins are discussed. 

Resumo.—Kolpotocheirodon theloura, género e espécie novos da tribo 
Compsurini de Cheirodontinae, é descrito para os tributarios superiores do rio 
Sao Francisco e rio Parana no Planalto Central do Brasil. Machos da nova 
espécie possuem um 6rgao formado por ganchos nos raios da nadadeira caudal 
e por dobras de pele (‘“‘flaps’’) hipertrofiadas ao longo dos raios, principalmente 
no lobo ventral da nadadeira caudal. O maior “‘flap’’, conectado ao longo da 
face dorsal do 19° raio da nadadeira caudal (raio principal inferior do lobo 
ventral da nadadeira caudal), é marginado por uma série de tabiques, e se 
estende dorsalmente sobre as dobras de pele originadas nos 17° e 18° raios. 
Existe ainda uma série de papilas carnosas distribuidas ao longo das demais 
dobras de pele do lobo ventral da nadadeira caudal ou ao longo dos raios 
imediatamente superiores aos raios com dobras de pele. Estes tabiques e papilas 
sao desconhecidos em outros caracideos, tanto naqueles com inseminagao como 
nos com fertilizagao externa, sendo considerados como caracteres derivados. 
A espécie nova é referida aos Compsurini em Cheirodontinae com base em 
uma diagnose cladistica destes taxons. SAo discutidas as relagdes de K. theloura 
com outros compsurineos de nadadeiras caudais modificadas. 


The new genus and species herein de- with modified, possibly glandular tissues, 
scribed (see Figs. 1-3) is a cheirodontine and hooks on the caudal fin of males. It was 


270 


discovered in the collections of the Museu 
de Zoologia, Universidade de Sao Paulo by 
one of us (L.R.M.) while reviewing chei- 
rodontine characids of southeastern and 
eastern Brazil. The species was first hy- 
pothesized to belong to a cheirodontine 
clade diagnosed by specialized modified 
anal-fin hooks and derived scales, fin rays 
and/or hypertrophied soft tissues on the 
caudal fin. In the unpublished thesis of Mal- 
abarba (1994), this clade includes Sacco- 
derma Schultz, Compsura Eigenmann, Ma- 
cropsobrycon Eigenmann, Acinocheirodon 
Malabarba & Weitzman, and the Central 
American ‘“‘Odontostilbe’’ dialeptura (Fink 
& Weitzman) and “‘O.”’ mitoptera (Fink & 
Weitzman). Later, the new taxon (listed as 
undescribed genus and species A in Burns 
et al. 1997) and all cheirodontine taxa listed 
above (see Burns et al. 1997) were found 
to be inseminating species. Subsequently, 
this putative clade was recognized as a 
cheirodontine tribe, the Compsurini (Mala- 
barba, Weitzman, & Burns in Malabarba 
1998), including the taxon described herein 
(therein referred as new genus and species 
A). The significance of these characters in 
resolving the phylogeny of compsurin chei- 
rodontines, as well as its possible homology 
to similar characters found in glandulocau- 
dine characids, were further discussed by 
Malabarba & Weitzman (1999). 

Males of the new genus and species have 
hooks on the caudal fin (Fig. 4) and hyper- 
trophied dermal flaps along the caudal-fin 
rays (Fig. 5). The largest flap attached dor- 
sally to the 19th ventral lobe caudal-fin ray 
is bordered with a series of tabs, and ex- 
tends dorsally over most of the flaps ex- 
tending from caudal-fin rays 17 and 18. 
There is also a series of fleshy papillae dis- 
tributed along the border of the remaining 
ventral caudal-fin ray flaps or along the fin 
rays just dorsal to the fin rays bearing flaps. 
These tabs and papillae are unknown in the 
remaining inseminating cheirodontines, as 
well as on other inseminating or externally 
fertilizing species of characids and are con- 
sequently hypothesized to be derived. This 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


is the only species of the Compsurini 
known to have aquasperm (a nearly spher- 
ical or spherical sperm nucleus, similar to 
that found in externally fertilizing chara- 
cids; see Burns et al. 1997:434, fig. 1A). 
All other species of the Compsurini so far 
investigated have elongate sperm cell bod- 
ies (see Burns et al. 1997:434, fig. 1B—H). 


Methods 


The systematic methods, counts and 
measurements used here are the same of 
those described and used by Malabarba & 
Weitzman (1999). The following acronyms 
are used for institutions and collections: 
MCP, Museu de Ciéncias e Tecnologia, 
Pontificia Universidade Catdlica do Rio 
Grande do Sul, Porto Alegre; and MZUSP, 
Museu de Zoologia da Universidade de Sao 
Paulo, Sao Paulo. The following abbrevia- 
tions are used in the text or figures: SL = 
standard length; HL = head length; c&s = 
alizarin red s and alcian blue stained spec- 
imens cleared with trypsin; spm(s) = spec- 
imen(s). The comparative material is the 
same listed in Malabarba & Weitzman 
(1999). 


Kolpotocheirodon, new genus 


Type species.—Kolpotocheirodon thel- 
oura, new species. 

Diagnosis.—The following apomorphies 
diagnose Kolpotocheirodon: 

A specialized caudal organ (Fig. 5) is 
present at the proximal region of the ventral 
caudal-fin lobe of the males (character 
number 36 in Malabarba 1998). This con- 
sists of hypertrophied elongate dermal flaps 
attached along the fin rays and a series of 
relatively flat tabs attached along the ex- 
posed border of the largest flap. The base 
of this flap is attached along the anterior 
approximately one-third of the nineteenth 
caudal-fin ray. Visually moving dorsally, 
the flaps decrease in length and width until 
those of the thirteenth or fourteenth fin rays 
are relatively short, narrow, and almost un- 
detectable. Each flap, other than that of the 


VOLUME 113, NUMBER 1 


nineteenth ray, bears papillae in a single se- 
ries along its exposed border. The tabs of 
the nineteenth fin ray flap extend dorsally 
while the papillae of the fourteenth to at 
least the sixteenth or seventeenth flap are 
directed ventrally. 

Hooks on the anal-fin rays of mature 
males (Fig. 6) occur along the posteriola- 
teral border of the posterior unbranched and 
five anterior branched anal-fin rays (Char- 
acter 30 in Malabarba 1998). Although the 
number of hook-bearing anal-fin rays in any 
species of the Characidae with hooks is var- 
iable according to the degree of maturation, 
it seems to be constant in fully mature spec- 
imens of some species. All fully mature 
males of Kolpotocheirodon theloura have 
fully developed hooks restricted to six of 
the anterior anal-fin rays. The remaining 
species of the Compsurini have hooks on a 
larger number of anal-fin rays, with the ex- 
ception of the species of Saccoderma, 
which have anal-fin hooks only along the 
posterior ray of the anterior unbranched 
rays and the four most anterior branched 
anal-fin rays. 

The twelfth and thirteenth caudal-fin rays 
are somewhat curved, being noticeably 
concave along their dorsal borders at about 
their basal half lengths (Fig. 4) and some- 
what convex along their ventral borders 
where the segments are slightly expanded 
longitudinally (character 34, state 2 in Mal- 
abarba 1998). This feature separates K. 
theloura from other cheirodontines. Acin- 
ocheirodon melanogramma Malabarba & 
Weitzman (1999), another compsurin, also 
has the basal halves of the caudal-fin rays 
dorsally concave with ventrally expanded 
segments, but in this species they are thir- 
teenth and fourteenth, rather than twelfth 
and thirteenth rays. 

Etymology.—The first component of the 
name Kolpotocheirodon is from the Greek, 
kolpotos = formed into folds, and refers to 
the caudal organ formed by hypertrophied 
dermal folds along the caudal-fin rays. The 
second component refers to the characid ge- 
nus Cheirodon Girard, in reference to our 


271 


referral of this new genus to the Cheiro- 
dontinae. 


Kolpotocheirodon theloura, new species 
Figs. 1-3 


All specimens from Brazil, Brasilia, Dis- 
trito Federal, except when noted. 

Holotype.-—MZUSP 55194, 1 male, 27.3 
mm SL, small marsh at Curva da Morte, 
Goias, 3 May 1978, E. K. Bastos. 

Paratypes.—Rio Parana tributaries: MCP 
1161, Lt male 25-9 mm SL; eds; MZUSP 
38840, 2 females, 27.9 and 29.7 mm SL, 
and MZUSP 38839, 1 male, 29.9 mm SL, 
ribeirao do Gama, just above the mouth of 
ribeirao Taquara, 1 Oct 1985, J. P Viana 
Mendes. MZUSP 39014, 1 female, 24.0 
mm SL and MCP 11160, 1 female, 22.2 
mm SL, ribeirao Riacho Fundo, tributary of 
rio Sao Bartolomeu, 3 Jul 1985, M. Ribeiro. 
MZUSP 42802, 1 male, 27.8 mm SL, 1 fe- 
male, 24.6 mm SL, collected with the ho- 
lotype. Rio Sao Francisco tributaries: MCP 
12204, 4 spms. (1 alcohol, 3 c&s), 14.0— 
16:0:mm SL, and MZUSP 35722; 12 spms, 
14.7—19.5 mm SL, corrego Vargem de Tras, 
1—2 Apr 1979, N. A. Menezes & E. K. Bas- 
tos. MZUSP 42801, 2 males, 22.9—23.8 
mm SL and 1 female, 20.2 mm SL, lagoa 
Feia, 3 May 1978, E. K. Bastos. 

Diagnosis.—The same as for the genus. 

Distinguishing characters.—The pres- 
ence of spherical sperm nuclei (aquasperm; 
see Burns et al. 1997:434, fig. 1A and tab. 
1, “undescribed genus and species A’’) is 
plesiomorphic for the Compsurini, accord- 
ing to the parsimony analysis in Malabarba 
(1998). Its presence in K. theloura distin- 
guishes this species from all other species 
of the Compsurini so far investigated, 
which have elongated sperm nuclei (see 
Burns et al. 1997:434, fig. 1B—H and tab. 
1). 

Kolpotocheirodon theloura also has an 
atypical color pattern for the Cheirodonti- 
nae of three to five very small vertical bars 
on the sides of the body, crossing the nar- 
row lateral horizontal body stripe. These are 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig, <1: 
at Curva da Morte, Goias, Brazil. 


located dorsal to the visceral cavity, and be- 
tween the pseudotympanum and the area 
ventral to the dorsal fin (Figs. 1-3). 

Kolpotocheirodon theloura can also be 
distinguished from other cheirodontines by 
a conspicuous dark brown band formed of 
a series of spots distributed along all of the 
hook-bearing portion of the anal-fin of the 
males. These pigment spots are placed at 
approximately the midlength of the poste- 
riormost of the anterior unbranched anal-fin 
rays and the five anteriormost branched 
anal-fin rays. The most conspicuous and 
darkest spot is at the anterior surface of the 
first branched ray (Fig. 1; although also pre- 
sent in the paratypes, the dark brown spot 
cannot be seen in the black ground Figs. 2— 
3). A similar, but less conspicuous band of 
spots occurs in females along the midlength 
of the anal-fin rays. These are more strong- 
ly pigmented in the anterior portion of the 
fin. 

Description —Morphometric data sum- 
marized in Table 1. 

Body moderately elongate and com- 
pressed, greatest depth at dorsal-fin origin. 
Predorsal profile convex, slightly concave 
in region of supraoccipital process. Profile 
of body from base of posterior dorsal-fin 
ray to origin of adipose fin straight or mod- 
erately convex. Ventral profile of body con- 
vex from tip of lower jaw to pelvic-fin or- 
igin and moderately concave from there to 
anal-fin origin. Body profile along anal-fin 


Kolpotocheirodon theloura, new species, holotype, male, MZUSP 55194, SL 27.3 mm; small marsh 


base of males moderately concave in ante- 
rior half and convex posteriorly. In female 
entire fin base relatively straight. Dorsal 
and ventral profile of caudal peduncle con- 
cave. Largest mature male examined with 
an expanded and rounded dorsal and ventral 
caudal-peduncle profiles. Caudal peduncle 
about as long as deep or somewhat shorter 
than deep. 

Head small and snout short, rounded. 
Mouth terminal. Maxilla short, positioned 
at an angle of approximately 45 degrees rel- 
ative to long axis of body. Posterior extrem- 
ity of maxilla reaching vertical that passes 
through anterior border of eye. 

Premaxilla with 4 (rarely 5) symmetrical 
teeth having 9-11 small evenly spaced 
cusps all about equal in size (Fig. 7). Cut- 
ting edge arched in most medial tooth and 
almost straight in most lateral tooth. Max- 
illa with 2 (rarely 3) teeth, similar in form 
to those of premaxilla, with 7—11 cusps. 
Cutting edge slightly arched to almost 
straight. Dentary with 4 large teeth with 5, 
7, or 9 cusps; followed by 1 or 2 smaller 
teeth with 3, 5 or 7 cusps. Teeth following 
second tooth asymmetrical with most lateral 
cusp situated towards tooth base and most 
medial cusp more distally located. Cusps 
small and regular and approximately equal 
in size. Cutting edge slightly arched to al- 
most straight. 

Dorsal-fin rays, ii, 9, n = 29. First un- 
branched ray about half length of second. 


VOLUME 113, NUMBER 1 


Dorsal-fin origin approximately at mid- 
length of body. Proximal portion of dermal 
flaps along largest unbranched and first 
branched dorsal-fin rays bordered by series 
of papillae in both sexes. Adipose-fin origin 
at, or slightly anterior to, vertical through 
insertion of posteriormost anal-fin ray. 

Anal-fin rays, iii, 18, Gi-iv, X = 3.3, 17— 
fee = 19.1, mn = 22). Anal-fin origin 
slightly posterior to vertical passing through 
base of posteriormost dorsal-fin ray. Anal 
fin of females with anterior 5—6 branched 
rays very long, forming prominent anterior 
lobe. Rays posterior to anterior lobe abrupt- 
ly shorter and distal border of anal fin con- 
cave posterior to anterior lobe. Distal bor- 
der of male anal fin decreasing in length 
gradually so that although anterior lobe 
larger than but not as abruptly distinct as in 
females. Anal-fin rays of males with slen- 
der, elongate retrorse hooks on longest un- 
branched ray, and anterior first 5 branched 
rays, usually with 4 to 5 segments bearing 
hooks on each ray (Fig. 6). Hooks inserted 
at posterolateral border of fin rays, bent 
over lateral surface of fin ray and anteriorly 
directed. Hooks located on _ posterior 
branches of rays only; never on proximal 
portions of main body of rays. One, rarely 
two, bilateral pair of bony hooks per ray 
segment. 

Pectoral-fin rays, i, 9, (i, 8-11, X = 9.2, 
n = 28). Distal ends of longest rays extend 
slightly beyond pelvic-fin origin. Pelvic-fin 
fet i 78 = F1, ) = (29). Pelvic- 
fin origin anterior to vertical passing 
through dorsal-fin origin. Tip of fin reach- 
ing anal-fin origin in males but not in fe- 
males. Male pelvic fins with unilateral, ven- 
tromedial retrorse bony hooks (Fig. 8); 
branched rays bearing one, rarely two slen- 
der hooks per segment along most of 
branched ray lengths, including both pri- 
mary branches and often posterior second- 
ary branches. Distal sections of unbranched 
part of each branched ray with one to oc- 
casionally few hooks. Number of hooks 
varies among different males, those males 
bearing more hooks appearing more mature 


Kolpotocheirodon theloura, new species, paratype, male, MZUSP 42802, SL 27.8 mm; small marsh at Curva da Morte, Goids, Brazil. 


Fig. 2. 


274 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 3. 
at Curva da Morte, Goias, Brazil. 


when considering other secondary sexual 
characters such as caudal-fin papillae. 
Principal caudal-fin rays 10/9, n = 29. 
Proximal portion of thirteenth or fourteenth 
through nineteenth caudal-fin rays in ven- 
tral caudal fin lobe with hypertrophied der- 
mal flaps. Largest flap extending dorsally 
from nineteenth principal caudal-fin ray 
(Fig. 5). Each flap bordered by one series 
of external papillae, more numerous and 
more developed in males. Proximal portion 


Kolpotocheirodon theloura, new species, paratype, female, MZUSP 42802, SL 24.6 mm; small marsh 


of third caudal-fin ray in dorsal caudal-fin 
lobe with hypertrophied dermal flaps and 
external papillae only in males. Males with 
modified 12th to 14th or 15th principal cau- 
dal-fin rays; these rays bearing (on each 
side) row of 4—6 dorsoanteriorly directed 
hooks along dorsal divisions (Fig. 4). Dor- 
sal and ventral procurrent rays equivalent in 
numbers and structure in both sexes, similar 
to those of most characids. Dorsal procur- 
rent caudal-fin rays, 13 (9-13, X = 10.8, n 


Table 1.—Morphometrics of Kolpotocheirodon theloura, new species. Standard length is expressed in mm; 
measurements through head length are percentages of standard length; the last four entries are percentages of 
head length. Range includes the holotype, MZUSP 55194, and following paratypes MZUSP 38839, MZUSP 
42801, MZUSP 42802, MCP 11161, MZUSP 38840, MZUSP 39014, MCP 11160. 


Males 


Holotype n Low 
Standard length (mm) DS 6 2pe9 
Snout to anal-fin origin 62.3 6 60.1 
Snout to dorsal-fin origin 49.5 6 48.9 
Snout to pelvic-fin origin 45.4 5 41.7 
Dorsal-fin base length hg Ws 5) 7 
Anal-fin base length 29.3 5 26.4 
Caudal peduncle length of 5 2h) 
Caudal peduncle depth 14.7 5) 29 
Depth at dorsal-fin origin 5525 5 32.4 
Dorsal-fin height 28.2 =) 27.0 
Pelvic-fin length 20:9 5 19.4 
Pectoral-fin length 23.8 5 2055 
Bony head length 26.0 5 24.4 
Snout length 13:3 3) 18.3 
Upper jaw length 25.4 4 25.4 
Horizontal eye diameter Sez 5 32.4 
Least interorbital width 29.6 5 29.6 


Females 

High XK nen \Ebote Pauklich ee ose 

259 26.3 5) 2ae2 294] Dae 
63.9 61.8 3) 62:2 66.3 65.2 
S15 50.0 5 48.3 5310 5 kat 
46.5 44.9 5 44.1 45.5 44.6 
13-1 [25 5 12.6 14.0 13,3 
29.4 28.6 5 26.2 28.5 27.4 
12.4 11.0 5 11.0 12-2 11.6 
14.7 Le 5 LAS 13:3 122 
56:1 34.3 5 30.4 37.6 34.4 
30.4 233, 5 Doe 29.3 27.9 
2, Wee 20.4 5 15.0 18.6 ig ee | 
23.8 PMs?) + 19.5 23:3 21.4 
26.8 25-8 5 Play, yada 26.5 
24.1 222 5 2A 24.7 22.9 
29.2 213 2 297 32:2 31.0 
39.1 36.1 5 35.6 38.5 36.8 
36.2 31.3 5) 29.2 IEA 30.6 


VOLUME 113, NUMBER 1 


Fig. 4. Kolpotocheirodon theloura, new species, paratype, MCP 11161, c&s male, SL 25.9 mm. Caudal-fin 
rays eleven (11th) through fifteen (15th) of the ventral caudal-fin lobe, other lower caudal-fin rays only partially 
indicated. Figure illustrates the position and number of caudal-fin ray hooks along fin rays twelve to fourteen. 


Anterior is to left. 


= 5). Ventral procurrent caudal-fin rays, 9 
(n = 5). 

Scales cycloid, moderately large. Lateral 
line poring incomplete, 8, (6-10, X = 7.6, 
n = 21) anterior perforated scales. Scales 
in lateral series 33, (31-34, X = 32.8, n = 
19). Scale rows between dorsal-fin origin 
mueeeneral line 5, (5-6, X — 5.1, n — 22). 
Scale rows between lateral line and pelvic- 
famien 4, G—4, X = 3:6; n = 22). Pre- 
dorsal scales, when in regular series 11 
esti oX = 10.8,.n = 9). Scale. rows 
around caudal peduncle 14 (nm = 5). Scale 
Sheath on anal-fin base consisting of 3—5 
scales covering unbranched rays and ante- 
rior 4—5 branched rays. 

Saoraneurals, 5° (3—5,°X' ="4.3).n' = 8). 
Precaudal vertebrae, 15 (15-16, X = 15.1, 
n = 12). Caudal vertebrae, 17 (17-18, X = 
17.5, n = 12). 

Color in alcohol.—Head and body pale 
brownish yellow. Black lateral body stripe 
very narrow, pale on caudal peduncle and 
anterior to dorsal-fin origin (Figs. 1—3). Lat- 
eral body stripe crossed by 3—5 very small 
vertical bars between pseudotympanum and 
area ventral to dorsal fin. Humeral spot ab- 
sent. Caudal-fin base and posterior termi- 
nation of caudal peduncle bear dark black, 
vertically-elongate, lozenge-shaped spot 
that reaches dorsal and ventral margins of 
caudal peduncle. Caudal spot extends just 
to most proximal portion of middle caudal- 


fin rays otherwise white or pale brownish 
yellow. All other fins hyaline except anal 
fin. Anal fin of males with conspicuous 
dark brown band distributed along all hook- 
bearing portions of fin (approximately mid- 
dle length of last unbranched anal-fin ray 
and five anteriormost branched anal-fin 
rays). Pattern most conspicuous at anterior 
surface of first branched ray. Less conspic- 
uous similar band of spots occurs in fe- 
males, along midlength of most anterior 
branched anal-fin rays. 

Sexual dimorphism.—The females lack 
hooks on the pelvic, anal, and caudal fins, 
while the males bear numerous small hooks 
on the pelvic and anal-fin rays. The male 
holotype and paratypes bear about 4—6 
hooks on caudal-fin rays 12—14 with more 
of these hooks on ray 12 than on ray 14. 
Both sexes bear dermal tabs and papillae on 
the caudal fin, but the females lack these 
papillae in the dorsal caudal-fin lobe al- 
though they are present in males. The fe- 
males have the distal border of the anal fin 
with a distinct anterior lobe that is abruptly 
curtailed at about branched rays 6—7. The 
male distal anal-fin profile is not lobate. In- 
stead the fin rays gradually diminish in 
length from the anterior to posterior termi- 
nations of the fin. The pelvic fins are longer 
in the males (19.4—21.7% of SL) than in 
the females (15.0—18.6% of SL), reaching 
past the anal-fin origin in the males, but not 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 5. 
portion of the ventral caudal-fin lobe. Figure illustrates the derived hypertrophied flaps bearing tabs and papillae 
along the caudal-fin rays. White arrow indicates largest flap (1f) extending dorsally from the nineteenth principal 
caudal-fin ray (19th), bordered by one series of flat tabs. All other flaps are bordered by one series of fleshy 
papillae. 


females. The relative position of the anal fin 
also seems to be sexually dimorphic, with 
the snout to anal-fin origin distance in 
males (60.1—63.9% of SL) shorter than in 
females (62.2—66.3% of SL). However, our 
sample size is small and the examination of 
larger population samples is necessary to 
confirm this pattern of difference between 
males and females. 
Distribution.—Kolpotocheirodon thel- 
oura has an interesting distribution, along 
the uppermost tributaries of both the rio Sao 
Francisco and the upper rio Parana basins, 
near Brasilia, in central Brazil. It is found 
close to an area called Aguas Emendadas 
(joint waters), in allusion to the close prox- 
imity of the head waters of the upper trib- 
utaries of the rio Sao Francisco, alto rio Pa- 
rana and rio Tocantins. This may explain 


Kolpotocheirodon theloura, new species, holotype, male, MZUSP 55194, SL 27.3 mm. Proximal 


the distribution of this species in two of 
these major basins. So far, this species is 
unknown from the rio Tocantins tributaries, 
but samples from that area are too rare to 
assure its absence in that drainage. 

Etymology.—The name theloura is from 
the Greek thele meaning nipple and oura 
meaning tail. The name refers to the pres- 
ence of papillae on the ventral lobe of the 
caudal fin. The name is a noun in apposi- 
tion. 

Discussion.—The relationships of Kol- 
potocheirodon theloura to other cheirodon- 
tine characids is supported by the presence 
of all four synapomorphies diagnosing the 
Cheirodontinae (Malabarba, 1998). These 
are: The presence of a large, nearly trian- 
gular, hiatus among the muscles covering 
the anterior chamber of the swimbladder 


VOLUME 113, NUMBER 1 


LY. 


Fig. 6. Kolpotocheirodon theloura, new species, paratype, MCP 11161, c&s male, SL 25.9 mm. Anterior 
portion of anal fin bearing hooks. Figure shows anal-fin hooks positioned along posterolateral border of anal- 
fin rays, bent anteriorly over lateral surface of anal-fin ray to which it is attached, and distal tip pointing 


anteriorly. 


between the first and second pleural ribs 
(pseudotympanum—Figs. 1—3). This hiatus 
is limited dorsally by the lateralis superfi- 
cialis muscle, posteriorly by a naked ante- 
rior face of the second pleural rib, postero- 
ventrally by the obliquus inferioris muscle, 
and antero-ventrally by the obliquus super- 
ioris muscle. See also Malabarba (1998: 
200-201, figs. 2B and 3A) and Weitzman 
& Malabarba (1999:7 and 16, figs. 5, 6, 16 
and 17). The humeral spot is absent (Figs. 
1—3). The teeth are pedunculated, largely 
expanded and compressed on their distal 
borders (Fig. 7). An unique regular, single 
tooth row is present on the premaxilla. The 
teeth of this row are perfectly aligned and 
similar in shape and cusp number (Fig. 7). 

Kolpotocheirodon theloura is included 
among the members of the cheirodontine 
tribe Compsurini (Malabarba, Weitzman, & 
Burns in Malabarba, 1998) because it has 
the following synapomorphies of that tribe: 
The species is inseminating. The anal-fin 
hooks are positioned along the posterolat- 


eral border of the anal-fin rays, but are bent 
more or less anteriorly over the lateral sur- 
face of the anal-fin ray to which each is 
attached. The distal tip of each anal-fin 
hook is directed towards the anterior border 
of the anal-fin ray to which it is attached 
(Fig. 6), instead of pointing posteriorly or 
dorsally as in most other characids that 
have anal-fin hooks. Hooks are present on 
the dorsal surface of some of the caudal-fin 
rays and inclined towards the caudal-fin 
base (Fig. 4). Hooks are distributed along 
the distal lengths of the principal caudal-fin 
rays 11 to 14 (Fig. 4). The anal fin is more 
strongly pigmented along the distal portion 
of the branched rays (Fig. 1). Kolpotochei- 
rodon theloura was placed as the most bas- 
al species (identified as Species A) of the 
Compsurini in the parsimony analysis of 
Malabarba (1998). We herein further dis- 
cuss and describe the characters of K. thel- 
oura in order to better hypothesize its re- 
lationships to other compsurin species. 
The presence of insemination, where 


278 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Figs 7. 
maxilla and premaxilla and teeth, internal view. 


sperm are introduced into the ovary (noted 
above), is so far found in the species of the 
Glandulocaudinae (Burns et al. 1995), in 
the species of the tribe Compsurini of the 
Cheirodontinae (Burns et al. 1997; Mala- 
barba, Weitzman, & Burns in Malabarba 
1998), and species of uncertain relation- 
ships including Monotocheirodon pearsoni 
Eigenmann in Pearson (1924), Brittan- 
ichthys axelrodi Géry (1965), “‘“Cheirodon”’ 
ortegai Vari & Géry (1980) and “‘Brycon- 
americus’’ pectinatus Vari & Siebert 
(1990), (Weitzman & Malabarba 1998, 
Burns et al. 1999). The absence of cheiro- 
dontine synapomorphies among the taxa 


Kolpotocheirodon theloura, new species, paratype, MCP 11161, c&s male, SL 25.9 mm. Right 


just listed above and the hypothesis that the 
Compsurini is a derived branch arising 
from externally fertilizing cheirodontines 
(see cladogram, fig. 1 of Malabarba 1998) 
suggests a hypothesis that insemination in 
the compsurin cheirodontines arose from 
externally fertilizing ancestral cheirodonti- 
ne stock, and that it probably arose inde- 
pendently of the Glandulocaudinae and of 
the other taxa listed above. At least all chei- 
rodontines, including the compsurins, lack 
the following synapomorphy of the glan- 
dulocaudines used in part to diagnose that 
subfamily by Weitzman & Menezes (1998): 
presence of a sperm storage area in the tes- 


Fig. 8. 
illustrates the pelvic-fin rays and hooks in the left pelvic fin, anterior to left. Ventral view. 


Kolpotocheirodon theloura, new species, paratype, MCP 11161, c&s male, SL 25.9 mm. Figure 


VOLUME 113, NUMBER | 


tes. Although rather tentative because not 
all glandulocaudine taxa have yet been 
sampled, the presence of an elongate cyto- 
plasmic collar binding the flagellum to the 
nucleus at some stage of spermiogenesis or 
in mature sperm cells in glandulocaudines 
(see Burns et al. 1998) is absent in at least 
one compsurin species, Macropsobrycon 
uruguayanae. 

The presence of elongate sperm nuclei 
was independently pointed out by Weitz- 
man & Menezes (1998) as a synapomorphy 
for the Glandulocaudinae, and by Malabar- 
ba et al. in Malabarba (1998) as a synapo- 
morphy for a subclade of the Compsurini 
(excluding K. theloura). This requires some 
explanation. Sperm with spherical or slight- 
ly oval nuclei are often referred to as 
‘“‘aquasperm’”’ and considered to be the ple- 
siomorphic sperm type for teleosts (Jamie- 
son 1991). Elongation of sperm nuclei is a 
derived feature and seems to be related to, 
but not necessarily needed for, sperm trans- 
fer to and storage within ovaries (Jamieson 
1987). Such a modification is found in the 
two inseminating characiform groups listed 
above and in several inseminating or inter- 
nally fertilizing teleosts. Although the 
sperm cells in these groups have undergone 
changes in shape resulting in sperm cells 
different from the spherical or slightly oval 
nuclei of “‘aquasperm,’’ these changes are 
not necessarily homologous. Although we 
still lack information on the ultrastructural 
morphology of the sperm cells of most 
compsurins and several glandulocaudines, 
the evidence so far noted in Burns et al. 
(1998) indicates the elongated sperm cells 
are derived in different ways in the comp- 
surin Macropsobrycon uruguayanae and in 
the glandulocaudines Diapoma speculifer- 
um Cope (1894), Diapoma sp., Corynopo- 
ma riisei Gill (1858), Pseudocorynopoma 
doriae Perugia (1891), Mimagoniates bar- 
beri Regan (1907), and M. microlepis 
(Steindachner, 1876). Thus, elongation of 
the sperm nuclei in glandulocaudines and 
elongation of the sperm nuclei in at least 
part of the compsurins appear to have been 


279 


acquired in different ways and are non-ho- 
mologous, and thus likely constitute differ- 
ent synapomorphies for each clade. How- 
ever, much of the ultrastructural evidence 
from sperm cell morphology in the species 
of both clades remains to be discovered and 
described. 

The anal-fin hook arrangement of the 
Compsurini, including Kolpotocheirodon 
theloura, is also hypothesized to constitute 
a synapomorphy. Compsurin cheirodon- 
tines, except for Acinocheirodon melano- 
gramma, have hooks inserted along the 
posterolateral border of the anal-fin rays 
and bent anteriorly over the lateral surface 
of the anal-fin ray where each is attached 
(Fig. 6). Thus, instead of pointing posteri- 
orly and dorsally as in most characids, the 
distal tip of the anal-fin hooks are directed 
anteriorly. Non-compsurin cheirodontines 
have retrorse hooks positioned along the 
posterolateral border of the anal-fin rays, 
and directed posteriorly and towards the 
anal-fin base. Some glandulocaudines such 
as the xenurobryconins have the anal-fin 
hooks recurved and pointing dorsoanterior- 
ly, but in most cases these differ in shape 
from those of the compsurins, do not bend 
anteriorly over the lateral surface of the 
anal-fin ray, and do not pass across the 
point where it is attached (see Weitzman & 
Fink 1985:figs. 38-43; Weitzman et al. 
1994:fig. 11; and Weitzman & Ortega, 
1995:fig. 4). Among xenurobryconins, Ar- 
gopleura chocoensis Eigenmann has modi- 
fied large anal-fin hooks, anterodorsally and 
laterally directed, on rays 6—7 to 12-14, re- 
sembling those of the compsurins (see 
Weitzman & Fink 1985:fig. 38). However, 
these hooks in A. chocoensis are part of a 
complex pattern of hook development (see 
Weitzman & Fink 1985:29, apomorphy 7) 
absent in compsurin species. In any case, 
parsimony analysis also indicates that the 
condition in the Xenurobryconini and the 
Compsurini are non-homologous. 

The presence of hooks on the dorsal sur- 
face of some of the caudal-fin rays (char- 
acter 7) is also found among inseminating 


280 


glandulocaudines (see Menezes & Weitz- 
man 1990:fig. 5, Mimagoniates microlepis 
and fig. 24, Mimagoniates rheocharis Me- 
nezes & Weitzman (1990); Weitzman & 
Fink 1985:fig. 11, Xenurobrycon macropus 
Myers & Miranda-Ribeiro (1945), fig. 12, 
Xenurobrycon heterodon Weitzman & Fink 
(1985), fig. 16, Corynopoma riisei, fig. 17, 
Gephyrocharax atricaudatus (Meek & Hil- 
debrand 1912), fig. 19, Mimagoniates mi- 
crolepis; Weitzman et al., 1994:fig. 5, 
Ptychocharax rhyacophila Weitzman et al. 
(1994), in the incertae sedis species Brit- 
tanichthys axelrodi (see Malabarba & 
Weitzman 1999:figs. 9-10) and “‘Cheiro- 
don’”’ ortegai. The possible homology of the 
caudal fin modifications of the inseminating 
compsurin cheirodontine Acinocheirodon 
melanogramma with two Mimagoniates 
Regan (1907) species and with B. axelrodi 
were extensively discussed in Malabarba & 
Weitzman (1999) and rejected. Homology 
of the caudal-fin hooks in K. theloura with 
all non-compsurin taxa listed above is also 
rejected because these inseminating chara- 
cids lack the synapomorphies diagnosing 
the Cheirodontinae. 

The putative caudal gland of Kolpoto- 
cheirodon theloura does not have gross 
similarities to those of the inseminating 
glandulocaudines, except possibly Landon- 
ia latidens Eigenmann & Henn in Eigen- 
mann et al. (1914) of the tribe Landonini 
that also has folds of tissue in the caudal- 
fin region. Although a detailed comparison 
of the histological nature of these species 
needs to be done, it is very unlikely the 
folds in the Compsurini and Landonini are 
homologous in a parsimony analysis. The 
caudal organ of K. theloura also does not 
resemble those found in Compsura and 
Saccoderma. Species of these genera have 
derived scales associated with their caudal 
organs. Instead the caudal organ of K. thel- 
oura is formed by derived dermal flaps 
found along the proximal length of the thir- 
teenth or fourteenth to nineteenth caudal-fin 
rays. Although not sharing a tabular or pa- 
pillose margin, the caudal glands of other 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


inseminating compsurins are also partially 
formed by derived hypertrophied dermal 
flaps along the ventral lobe caudal-fin rays, 
as described by Malabarba & Weitzman 
(1999) along the thirteenth through fifteenth 
caudal-fin rays of Acinocheirodon melano- 
gramma, and fifteenth or sixteenth to nine- 
teenth caudal-fin rays in the Saccoderma 
species and Compsura heterura Eigen- 
mann. The caudal glands of compsurin spe- 
cies, although formed in different ways, are 
always derived from modified dermal flaps 
along the ventral lobe caudal-fin rays. This 
seems to reinforce the monophyly of the 
group and the hypothesized homology or 
common origin of insemination among 
compsurin cheirodontines. 

Males of Kolpotocheirodon theloura 
have modified twelfth to the fourteenth or 
fifteenth principal caudal-fin rays. These 
rays bear 4—6 retrorse hooks on each side 
in a row along their dorsal divisions (Fig. 
4). Among the Compsurini, Malabarba & 
Weitzman (1999) listed and described the 
presence of hooks in Acinocheirodon me- 
lanogramma (caudal-fin rays 13-14, rarely 
on ray 15), Saccoderma hastata Eigenmann 
(1915) (caudal-fin rays 13-18), ‘‘Odonto- 
stilbe”’ dialeptura (caudal-fin rays 12—16), 
and in Macropsobrycon uruguayanae (cau- 
dal-fin rays 12—14, plus several spinelets 
along the proximal half of the caudal-fin 
rays 14 to 18). Although these hooks are 
present on the ventral lobe of the caudal fin 
in all these species, we noted (Malabarba & 
Weitzman 1999) that they do not all occur 
on the same caudal-fin rays in all species 
and are of different shapes. Additionally, 
not all compsurin species have caudal-fin 
hooks, as for example Compsura heterura, 
Compsura gorgonae Evermann & Golds- 
borough and ‘‘Odontostilbe’’ mitoptera. We 
also pointed out that the use of the presence 
of caudal-fin hooks in diagnosing a clade 
among the Compsurini species would be 
parsimoniously incongruent with other de- 
rived characters, as for example the pres- 
ence of derived pouch scales in the ventral 
lobe of the caudal fin, found in some hook- 


VOLUME 113, NUMBER |! 


bearing Saccoderma species and some non- 
hook-bearing species such as Compsura 
heterura and C. gorgonae. A parsimony 
analysis of character distribution indicates 
the presence of hooks as a synapomorphy 
for the compsurin cheirodontines, and its 
lack a secondary reversal in some of its spe- 
cies (Malabarba et al. in Malabarba 1998). 
Kolpotocheirodon theloura was placed as 
the most basal species among the Comp- 
surini, in the phylogeny proposed by Mal- 
abarba et al. (in Malabarba 1998). Among 
the characters that placed the new species 
as the most basal compsurin is the presence 
of aquasperm (see Burns et al. 1997:434, 
fig. 1A), as found in externally fertilizing 
cheirodontines. In other words, the presence 
of derived elongate sperm nuclei (see Burns 
et al. 1997:434, table I) in the remaining 
inseminating compsurins represents a syn- 
apomorphy for these derived fishes. 
Another character that seems to corrob- 
orate a basal position for Kolpotocheirodon 
theloura is the lack of enlarged caudal- 
pouch scales as a possible pumping mech- 
anism in the ventral lobe of the caudal fin 
associated with a caudal gland (see Weitz- 
man & Fink 1985:96—-99; Malabarba & 
Weitzman 1999). Compsura heterura, C. 
gorgonae, and Saccoderma species have 
the posterior ultimate scale of the scale row 
immediately ventral to the lateral-line scale 
row enlarged, connected dorsally to liga- 
ments arising from the twelfth and _thir- 
teenth principal caudal-fin rays, and ven- 
trally to a skin flap connected to the nine- 
teenth principal caudal-fin ray, forming a 
pouch with a posterior opening that may 
serve aS a pumping mechanism. Acino- 
cheirodon melanogramma does not have 
any pumping mechanism involving caudal- 
fin scales. Instead it has two large skin flaps 
between the thirteenth and fifteenth princi- 
pal caudal-fin rays (Malabarba & Weitzman 
1999:fig. 6), that occur just ventral to and 
surrounding the hypertrophied soft tissue 
found around the hooks of the thirteenth 
and fourteenth fin rays. Malabarba & 
Weitzman (1999) suggested that these flaps 


281 


might also be part of a mechanism for in- 
creasing the rate of pheromone dissemina- 
tion in the surrounding water during court- 
ship. We found no putative active pumping 
mechanism in Kolpotocheirodon theloura 
that might be related to an increase in pher- 
omone dissemination in water during court- 
ship. The papillose margins of the dermal 
flaps of K. theloura seem to be a passive 
strategy for increasing pheromone dissem- 
ination. 


Acknowledgments 


Financial aid for travel for museum study 
and field work in Brazil were supplied to 
LRM by CAPES and Conselho Nacional de 
Desenvolvimento Cientifico e Tecnolégico 
(CNPq-Proc. 451459/96-2). José Lima de 
Figueiredo and Osvaldo Oyakawa (MZUSP) 
loaned specimens in their care or provided 
cataloging services. Lisa Palmer (USNM) 
and José EF P. Silva provided cataloging and 
other technical services. John R. Burns pro- 
vided useful insights through histological 
and TEM analyses. This paper was im- 
proved by the comments of José Lima de 
Figueiredo in the first version of the descrip- 
tion of this species, and in more recent ver- 
sions by Richard Vari. 


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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


113(1):284—290. 2000. 


Calls and calling behavior of the frog Leptodactylus natalensis 
(Amphibia: Anura: Leptodactylidae) 


W. Ronald Heyer and Celso Morato de Carvalho 


(WRH) Department of Vertebrate Zoology, National Museum of Natural History, 
Smithsonian Institution, Washington, DC 20560-0107, U.S.A.; (CMC) Departamento de Biologia, 
Universidade Federal de Sergipe, Jardim Rosa Elze, s/n, Sao Crist6vao, SE 49100-000, Brazil 


Abstract.—Leptodactylus natalensis has a prolonged calling season, typically 
from May to September. Individuals occur in and call from the most forested 
patch of habitat on the campus of the Universidade Federal de Sergipe. Two 
kinds of calls were recorded and analyzed—advertisement and chirp calls. The 
function of the chirp call is unknown. The advertisement call has two carrier 
frequency bands, known to date for only two other species of Leptodactylus. 
The two carrier bands may correspond to different frequency tuning sensitiv- 
ities in males and females, as females have slightly larger tympani than males, 
corresponding with their larger overall size. 


Published natural history information for 
the frog Leptodactylus natalensis Lutz, 
1930, consists of comments on calling and 
microhabitat made by Lutz (1930) in his 
original description of the species and a re- 
cent study of some reproductive aspects, in- 
cluding the first description of the tadpole 
for the species (Oliveira & Lirio Junior 
1999). Calls of Leptodactylus natalensis 
have not been analyzed previously (Heyer 
1994). One of us (CMC) had the opportu- 
nity to observe calling behavior and record- 
ed the calls of L. natalensis. The purpose 
of this paper is to report these observations 
and describe the calls. 


Methods and Materials 


Observational data and recordings of 
Leptodactylus natalensis were made by 
CMC on the campus of the Universidade 
Federal de Sergipe, SAo Crist6vao, Sergipe, 
Brazil. Incidental information on calls at 
Santo Amaro das Brotas, Sergipe, was also 
obtained by CMC. 

Recordings were made from two individ- 
uals. Five recordings (USNM tape 323, cut 
1) were made from specimen MZUSP 


89945 (MZUSP = Museu de Zoologia, 
Universidade de Sao Paulo collection), 
SVL (snout—vent length) 33.9 mm, on 22 
June 1998, at an air temperature of 25°C, 
water temperature 24°C, the last recording 
completed at. 1955 h. One recording 
(USNM tape 323, cut 2) was made from 
specimen MZUSP 89947, SVL 32.4 mm, 
also on 22 June 1998, air 25°C, water 24°C, 
the recording completed at 2030 h. It was 
not raining when these recordings were 
made, but a brief heavy rain occurred at 
2100 h. 

Calls were recorded with a Sony TM- 
5000 cassette recorder with a M 44 N(C) 
Beyer Dynamic Soundstar II microphone. 

Terminology follows that of Heyer et al. 
(1990), with the following clarifications. 
Carrier frequency is the frequency broad- 
cast by the calling frog to the environment. 
Often, the carrier frequency embraces a 
range of frequencies loud enough to be 
heard by the intended receiver (male or fe- 
male). The dominant frequency as used 
here is the single frequency value (or very 
narrow range of frequencies) that has the 
most sound energy in the call. 

Recordings were analyzed using ‘‘Ca- 


VOLUME 113, NUMBER 1 


nary 2.1”’ software (Charif et al. 1995). Call 
rates and durations were measured from 
wave form displays, beginning and ending 
carrier frequencies from audiospectrogram 
displays, and dominant frequencies from 
spectral displays. Although the initiation of 
calls is clear on the wave form displays, the 
ends of the calls are not very distinct, per- 
haps due to the level of background noise 
and possible microphone ringing. 

Advertisement call rates were based on 
six calling sequences ranging from 7.8 to 
15.2 sec from each of six recordings. Fre- 
quency data were taken from 10 consecu- 
tive calls from each of the six recordings; 
call duration and pulse data were taken on 
the same 10 calls for recordings 2 and 5 of 
MZUSP 89945 and for MZUSP 89947. 

Chirp call data are from the six chirps 
loud enough to analyze in the second re- 
cording of MZUSP 89945. 


Behavior and Calling Site 


Leptodactylus natalensis has a prolonged 
calling season coinciding with the rainy 
season, typically from May to September 
(also see Oliveira & Lirio Junior 1999). 
Calling starts around 1600 h. Calling activ- 
ity is most intense around 1900 and contin- 
ues until 2300 or 2400 h. However, on rainy 
days, especially at the beginning of the 
rainy season, males can be heard calling at 
0800 h, and continuing throughout the day 
and night. Males do not call during hot and 
sunny days during the rainy season. 

The general site the frogs occupy on the 
campus is a small patch (100 <X 400 m) of 
second growth (capoeira) on the right bank 
of the Rio Poxim, a seasonally partially 
flooded area (varzea) with many ponds. The 
site has 5—6 m high trees, many young palm 
trees, a thick grass ground cover, shallow 
litter, and white sand. The Leptodactylus 
natalensis were only heard calling from the 
capoeira—never from the left bank of the 
Rio Poxim, which has more artificial open 
areas and patches of mangrove swamps. 

MZUSP 89945 was calling very close to 


285 


the foot of a young palm tree surrounded 
by a pond with leaves in the water. The frog 
sat in the water, half submerged. A second 
male (MZUSP 89946) was about a half me- 
ter from the calling male. This second male 
was in a small burrow, with half the body 
exposed and uttered some “‘clicks”’ but no 
other calls. The “‘clicks’’ did not record ad- 
equately enough to analyze. 

MZUSP 89947 was near the foot of a 
small tree, at the edge of a little channel of 
rain water, hidden in grass. 

Leptodactylus natalensis infrequently 
jump when disturbed. Usually they take two 
or three steps backward and quietly hide 
under a leaf or other ground cover. 


Advertisement Call 


Calls (Fig. 1) consist of single notes giv- 
en frequently when actively calling. Call 
rates vary from 3.4—4.1 per sec for MZUSP 
89945 and at a rate of 3.0 per sec for 
MZUSP 89947. The call sequence with the 
least background noise is the fifth recording 
sequence for MZUSP 89945. In that se- 
quence, each call (=a single note) has a du- 
ration of 0.06—0.07 (mode = 0.07) sec and 
consists of 5—7 pulses, modally 7. The call 
is partially pulsed (=incompletely ampli- 
tude modulated). In two other recordings 
with greater background noise, it appears 
that the terminal pulses are indistinguish- 
able from the background noise in the wave 
forms. Three to five partial pulses (mode = 
4) can be distinguished in calls from the 
second recording of MZUSP 89945, with 
an average duration of 0.04 sec. Two to 
three partial pulses (mode = 3) are distin- 
guishable in the recording from MZUSP 
89947, with an average duration of 0.02 
sec. The calls are frequency modulated with 
extremely fast rise times (difficult to see be- 
cause of time scale in Fig. 1, more visible 
in the two advertisement calls shown in Fig. 
3). Beginning frequencies of the carrier 
(=fundamental) frequency range from av- 
erages of 550—600 Hz with the highest fre- 
quencies ranging from averages of 1370-— 


286 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 
“ni Wernher 
50 ms 
Sa. Saar ls 

N 

ed 

i 

iP) 

c= 

‘e) 

eas 

v 

e e 
Time in seconds 
Fig. 1. Wave form (above) and audiospectrogram (below) of the advertisement call of Leptodactylus natal- 


ensis, USNM tape 323, cut 1, fifth recording. The wave form is of the first call shown in the audiospectrogram. 


1830 Hz for MZUSP 89945. The average 
beginning dominant frequency for MZUSP 
89947 is 650 Hz, with an average maxi- 
mum of 1600 Hz. The dominant frequen- 
cies most often occur in two peaks in the 
calls of MZUSP 89945. For this frog, the 
first recording series usually has a dominant 
frequency of 760 Hz, with either a co-dom- 
inant or quieter frequency at 1050 Hz; the 
second recording has the dominant frequen- 
cy at 770 Hz, with a secondary dominant 
at 1020 Hz; the third recording has the 
dominant frequency at 760 Hz, with a sec- 
ondary dominant at 1040 Hz; the fourth re- 
cording has the most variation with either 
780 and 1030 Hz being primary or second- 
ary in loudness, 860 and 1030 Hz being 
primary or secondary in loudness, 1030 Hz 
loudest with a secondary dominant at 880 
Hz, or a single dominant frequency at 1030 


Hz; calls in the fifth recording have the 
dominant frequency at 1030—1040 Hz, with 
a secondary dominant at 760—770 Hz (Fig. 
2). The calls of MZUSP 89947 have either 
a dominant frequency at 1030 Hz, or also 
with a secondary dominant at 875 Hz. The 
partial pulsing of the calls appears to be the 
cause of the sidebands weakly visible in the 
audiospectrograms (Fig. 1). Calls of 
MZUSP 89945 range from having no visi- 
ble harmonics, a weak second harmonic, or 
intermediate strength second and third har- 
monics (Fig. 2). The calls of MZUSP 
89947 have a weak second harmonic. 


Chirp Call 


Calls (Fig. 3) consist of either 1 or 2 
notes. The loudest chirp calls are louder 
than the advertisement calls. In the two note 


VOLUME 113, NUMBER 1 


dB 


287 


| | 1980-2070 


\ as 
VV \ 2600 2930 
oe Ch ow ee el aoe 


O 1 2 3 + 5 
e 
Kilohertz 
Fig. 2. Power spectrum of advertisement call of Leptodactylus natalensis, USNM recording 323, cut | (of 


second call in audiospectrogram of Fig. 1). 


calls, the first note is shorter than the sec- 
ond (0.04 and 0.13 with an internote inter- 
val of 0.10 sec; 0.05 and 0.08 with an in- 
ternote interval of 0.16 sec; 0.04 and 0.13 
with an internote interval of 0.14 sec). The 
three single note chirp calls recorded range 
from 0.10—0.14 sec duration. From 3-10 
partial pulses can be discerned in the wave 
forms. The calls are frequency modulated, 
at a noticeably slower rise time than the ad- 
vertisement calls. The lowest carrier (=fun- 
damental) frequency ranges from 530—650 
Hz; the highest carrier frequency ranges 
from 990-1200 Hz. There is a suggestion 
of the carrier frequency briefly falling at the 
end of some calls, back to around 900—950 
Hz. The dominant frequencies vary from 
690-860 Hz. There are weak to strong sec- 
ond, second and third, or second, third, and 
fourth harmonics. 


Discussion 


The function of the chirp calls is un- 
known. Other members of the L. melano- 
notus group have these calls as well, which 
usually initiate calling bouts. In the case of 


the L. natalensis recording, the chirp calls 
were uttered after there was a slowing down 
in the rate of advertisement calls. The chirp 
calls are much more variable than the ad- 
vertisement calls. 

Many species of Leptodactylus have fre- 
quency modulated calls that result in broad- 
casting their calls over a range of frequen- 
cies. Most of these calls have a single (or 
narrow range) dominant frequency, how- 
ever. Leptodactylus natalensis is unusual in 
that there are two peaks of loudness. Only 
L. melanonotus and podicipinus within the 
L. melanonotus group share this condition 
(of those for which the calls are known, Ta- 
ble 1). We hypothesize that these separate 
dominant frequencies correspond to differ- 
ent tuning curves in males and females, as 
demonstrated in Eleutherodactylus coqui 
(Narins & Capranica, 1976), although no 
experimental data are available for L. na- 
talensis. Morphological data do not contra- 
dict this hypothesis. Other things being 
equal, a larger tympanum will be more sen- 
sitive to lower frequencies than a smaller 
tympanum and vice versa. Morphological 


b> 


Kilohertz 
NO 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Time in seconds 


Bigs: 


Wave form (above) and audiospectrogram (below) of the chirp call of Leptodactylus natalensis, 


USNM tape 323, cut 1. In the audiospectrogram, a single-note chirp call is followed by two advertisement 


calls. 


data are available from two localities rep- 
resented by several males and females. The 
data indicate that the tympanum size does 
differ between males and females (Table 2). 
It would seem from the variability of calls 
in the fourth recording of MZUSP 89945 
that an individual frog has control over 
whether more energy is broadcast in the 
lower or higher dominant frequency. 
Sexual difference in tympanum size is the 
expected condition in most frogs, as most 
frog species are sexually dimorphic, with the 
females being larger. Broadcasting of higher 
and lower dominant frequencies, as seen in 
L. natalensis, is one solution to matching 
different receptor sensitivities to the adver- 
tisement call. The frequency modulation ob- 


served in other Leptodactylus species, such 
as L. fuscus, is another solution. 

Our observations apparently conflict with 
those of Adolfo Lutz regarding the call of 
L. natalensis. In the original description of 
the species, Lutz observed (1930:27, En- 
glish version of text): ““This frog has a call 
with the strength of a whistle but sounding 
more like the voice of a cricket. It seems to 
have also another sound, heard during cop- 
ulation, like a soft clucking, sometimes re- 
peated by other males, so as to form a gen- 
eral concert. The specimens were caught 
near to the water but generally well hid- 
den.’? CMC has not heard cricket-like calls 
from L. natalensis at either the campus at 
Sao Crist6vao or Santo Amaro das Brotas, 


VOLUME 113, NUMBER 1 


289 


Table 1.—Advertisement call characteristics of members of the Leptodactylus melanonotus species group. 
Data taken from Heyer (1994), Heyer et al. (1996), and for L. melanonotus, from USNM tape 83, cut 1. Note 
that Leptodactylus diedrus is no longer considered a member of the L. melanonotus species group (Heyer, 1998). 
The duration and pulse data for L. natalensis are from the recording with the least background noise. 


Call duration Number partial 


Carrier Dominant 


Species in sec pulses per call frequency range frequency range 
L. griseigularis 0.01 l 1380-3060 2770 
L. leptodactyloides 0.02—0.04 1 or 3-5 650-1600 1 100—1300 
L. melanonotus 0.07—0.08 4-6 820-2620 1320-1380 or 2330 
L. natalensis 0.06—0.07 5-7 550-1830 760-880 or 1020—1040 
L. nesiotus 0.03 4-5 1500—2000 1800—2000 
L. pallidirostris 0.03-—0.05 1 or 2-5 1500-3500 3000-3500 
L. petersii A 0.04—0.05 3-4 700-1200 750-800 
L. petersii B 0.03—0.05 1 or 2-4 800—1600 or 1800—2800 N.A. 
L. podicipinus 0.02—0.04 3-7 1000—3500 1000—1200 or 3300-3500 
L. sabanensis 0.04—0.06 very weak 900-2300 1400—1800 
L. silvanimbus 0.15 about 20 420-1920 510 
L. validus 0.03—0.06 1 or 2-6 1300-3500 2300-3500 
but the call we describe as the advertise- Acknowledgments 


ment call is a soft clucking call. 

Advertisement calls in the L. melanono- 
tus group are often not as species-specific 
as in other Leptodactylus, perhaps because 
the calls have a strong point-location ori- 
entation function overriding (or in addition 
to) a species-coding function (Table 1 and 
Heyer 1994:106—107). Leptodactylus natal- 
ensis has a disjunct geographic distribution 
relative to all other members of the L. me- 
lanonotus group. The species group mem- 
ber that most closely approaches the distri- 
bution of L. natalensis is L. podicipinus 
(compare Figs. 39 and 43 in Heyer 1994). 
Interestingly, as pointed out above, L. pod- 
icipinus is only one of two other known 
members of the L. melanonotus group be- 
side L. natalensis to have two distinct loud- 
est broadcast peaks in the dominant fre- 
quency band. 


This research was partially supported by 
the Neotropical Lowlands Research Pro- 
gram, National Museum of Natural History, 
Dr. Richard P. Vari, Principal Investigator 
(WRH) and Departamento de Biologia, 
Universidade Federal de Sergipe (CMC). 
We thank these institutions for support and 
Dr. A. Stanley Rand for reviewing drafts of 
this paper for us. 


Literature Cited 


Charif, R. A., S. Mitchell, & C. W. Clark. 1995. Ca- 
nary 1.2 User’s Manual. Ithaca, New York, Cor- 
nell Laboratory of Ornithology. 

Heyer, W. R. 1994. Variation within the Leptodactylus 
podicipinus-wagneri complex of frogs (Am- 
phibia: Leptodactylidae).—Smithsonian Contri- 
butions to Zoology 546:1—124. 

1998. The relationships of Leptodactylus 


Table 2.—Snout-vent length and tympanum diameter measurements for male and female Leptodactylus na- 
talensis from two localities in the State of Sergipe, Brazil. 


Snout—vent length Tympanum diameter 


Locality n Sex Min. Max. Mean Min. Max. Mean 
Areia Branca 8 fe) 28.7 36.5 33.1 Zz 32 2.8 
Areia Branca 9 Q 33.9 42.6 38.8 Pink 3.4 3.0 
Santo Amaro das Brotas iT 3 30.6 34.6 325 2.4 ZF 2.6 
Santo Amaro das Brotas 14 2 33.1 37.0 S| Ps 2.9 Og | 


290 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


diedrus (Anura, Leptodactylidae).—Alytes 16: ras alliadas——Memorias do Instituto Oswaldo 
1294. Cruz 23:1-—20 (Portuguese), 21—34 (English), 
, R. de SA, J. R. McCranie, & L. D. Wilson. +5 plates. 


Narins, P. M., & R. R. Capranica. 1976. Sexual dif- 
ferences in the auditory system of the tree frog 
Eleutherodactylus coqui.—Science 192:378- 
380. 

Oliveira, EF FE, & G. P. Lirio Junior. 1999. Diversidade 
e aspectos reprodutivos de uma comunidade de 


1996. Leptodactylus silvanimbus (Amphibia: 
Anura: Leptodactylidae): natural history notes, 
advertisement call, and relationships.—Herpe- 
tological Natural History 4:169-174. 

, A. St Rand, C, A: G, Cruz, OF} Peixoto, & 


Caples Nelson. fe Frogs of Boracéia.—Ar- anfibios anuros de uma regiao do rio Poxim, 
quivos de Zoologia 31:231—410. Sergipe (Amphibia, Anura).—Publicag6es 
Lutz, A. 1930. Segunda memoria sobre especies bras- Avulsas do Centro Académico Livre de Biolo- 


ileiras do genero Leptodactylus, incluindo out- gia, Universidade Federal de Sergipe (in press). 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


113(1):291-297. 2000. 


A new species of venomous coral snake (Serpentes: Elapidae) from 


high desert in Puebla, Mexico 
Jonathan A. Campbell 


Department of Biology, The University of Texas at Arlington, 
Arlington, Texas 76019, U.S.A. 


Abstract.—A new species of coral snake (Micrurus pachecogili) is described 
from southern Puebla, Mexico. Black body rings in this snake are monadal and 
the species is allopatric from all other coral snakes except perhaps M. laticol- 
laris, a species in which black body rings are in triads. The new species is 
characterized by distinctive, well developed yellow rings, relatively short red 
rings, and a relatively large number of ventral scutes in comparison with M. 
nebularis. 

Resumen.—Se describe una nueva especie de serpiente coral (Micrurus pa- 
checogili) del sur de Puebla, México. En esta serpiente, los anillos corporales 
negros son monodales; la nueva especie es alopatrica de otras serpientes de 
coral, tal vez con excepcién de Micrurus laticollaris, una especie en la que los 
anillos negros se disponen en triadas. La nueva especie se caracteriza por po- 
seer anillos amarillos bien desarrollados, anillos rojos relativamente cortos y 


un numero grande de escamas ventrales en comparacion con M. nebularis. 


Species of coral snakes are widespread, 
if not always conspicuous, components of 
New World herpetofaunal assemblages. 
They occur throughout most of the Neo- 
tropics and reach into both temperate North 
and South America. These snakes are often 
associated with mesic conditions and the 
majority of species occur in rainforest or 
cloudforest habitats. Coral snakes appear to 
be delicate and desiccate quickly when ex- 
posed to direct sunlight. Thus, most species 
are nocturnal or are active only during early 
morning or later afternoon, especially on 
rainy nights or overcast days. However, a 
fair number of coral snakes have become 
adapted to subhumid habitats, occurring in 
desert, thorn scrub, or tropical deciduous 
forest from the Sonoran Desert of the south- 
western U.S.A. southward on the Pacific 
coast of Mexico and Central America and 
in various regions in South America. In Pa- 
cific Mexico, tropical deciduous forest is in- 
habited by Micurus browni, M. distans, M. 


ephippifer, M. bogerti, M. laticollaris, M. 
proximans, and Micruroides euryxanthus, 
with the latter species also extending into 
the Sonoran Desert (Campbell & Lamar 
1939). 

The arid lands of southern Puebla are sit- 
uated in the northern part of the Mesa del Sur 
physiographic region and are just south of the 
Mesa Central (West 1964, fig. 3). Heretofore, 
this area was reported to harbor only a single 
species of coral snake, M. laticollaris (Camp- 
bell & Lamar 1989, Roze 1996), easily dis- 
tinguishable from other Mexican coral snakes 
in being one of two species with black body 
rings in triads (the other triadal species being 
M. elegans of mesic forests on the eastern 
versant). Some years ago, I traversed on mul- 
tiple occasions the countryside surrounding 
Zapotitlan Salinas in southern Puebla, during 
which time I assembled small collections of 
amphibians and reptiles. Among the material 
in these collections are two specimens of a 
species of coral snake that cannot be associ- 
ated with any species known from Mexico. 


292 


Materials and Methods 


Descriptions of characters and terminol- 
ogy of scales in the diagnosis and descrip- 
tion of this new taxon follow Campbell & 
Lamar (1989) and Roze (1996). The sex of 
individuals was confirmed by checking for 
the presence of hemipenes by making a 
small midventral incision on the proximal 
section of the tail. Head measurements were 
taken to the nearest 0.1 mm using digital 
calipers held under a dissecting scope and 
the snout—vent and total length was taken 
to the nearest 1.0 mm using a meter stick. 


Micrurus pachecogili, new species 
(Figs. 1-3) 


Holotype.—An adult male (Figs. 1-2), 
UTA R-12546 (original number JAC 9752), 
from 5.6 km SSW Zapotitlan Salinas, 1494 
m, Puebla, Mexico, collected in November 
1983 by one of the children of E. Pacheco- 
Gil. This locality is in high tropical arid for- 
est at 18°18'N, 97°31'W (Fig. 3). 

Paratype.—A subadult male, UTA R- 
17145, from the type-locality, collected in 
December 1985. 

Diagnosis.—Micrurus pachecogili dif- 
fers most notably from other species of Mi- 
curus in central Mexico in being tricolored 
and having black body rings in monads 
(i.e., a body pattern of red-yellow-black- 
yellow-red), in having 220—223 ventrals in 
males, in possessing a wide pale parietal 
ring that includes all of the parietals and 
portions of adjacent scales, and in having 
non-melanized red body rings that are about 
equal to or shorter than black body rings. 
Indeed, these four characters, in combina- 
tion, distinguish M. pachecogili from all 
other venomous Mexican coral snakes. 

Micrurus bernadi differs in having mostly 
red and black body rings, although a yellow 
parietal ring and, rarely, narrow yellow edg- 
ing may be present along black markings, 
black rings are usually reduced to dorsal 
spots or saddlelike bands, and males have 
198-212 ventrals. Micrurus browni differs 
in having a black nuchal ring that covers the 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


posterior portion of the parietals and adult 
males have supracloacal keels. In M. b. 
browni of Guerrero and Oaxaca, the subspe- 
cies that is geographically most proximate to 
Zapotitlan, males have 204—218 ventrals, the 
black head cap extends to the anterior part 
of the parietals and frontal, and the black 
nuchal ring covers the ultimate and penulti- 
mate supralabials (versus only the posterior 
edge of the ultimate supralabial). Micrurus 
diastema differs in that the red rings are ei- 
ther very long (>5 times wider than the 
black rings) or contain scales that are dis- 
tinctly black-tipped, the black head cap in- 
cludes the anterior portion of the parietals 
and/or the black nuchal ring includes the 
posterior portion of the parietals. Micrurus 
distans of western Mexico differs in having 
no more than 17 black body rings, males 
have <217 ventrals, and red body rings are 
usually at least three times longer than black 
rings. Micrurus ephippifer differs in having 
a black nuchal ring that covers the parietal 
tips, red rings have black-tipped scales, 
spots, or large saddlelike bands, and =219 
ventrals in males. Micrurus laticollaris has 
black body rings arranged in triads (i.e., 
body pattern of red-black-yellow-black-yel- 
low-black-red), has 1 + 2 temporals, and 
males possess $215 ventrals. Micrurus lim- 
batus is bicolored with red and black rings 
and <192 ventrals. Overall, the color pattern 
of M. pachecogili is most similar to that of 
M. nebularis: similarities include red and 
black rings of relatively equal length, scales 
in red rings not black-tipped or distinctly 
spotted, well developed yellow body rings 
(1.0—1.5 dorsal scale lengths long in M. ne- 
bularis versus 2.0 in M. pachecogili), and 
tail with subequal black and yellow rings. 
Micururus nebularis differs by having a 
black nuchal ring that covers the posterior 
ends of the parietals, 203—208 ventrals in 
males, and seven black tail bands. 
Description of holotype.—A single pre- 
nasal and postnasal on each side; nostril sit- 
uated mostly in posterior part of prenasal; an 
elongate preocular on each side contacting 
postnasal; no loreal; anterior scales on head 


VOLUME 113, NUMBER 1 


and snout, including mental and infralabials, 
with numerous tiny tubercles; postoculars 2/ 
2; temporals 1 + 1 + 2; upper tertiary tem- 
poral large, about half size of parietal; su- 
pralabials 7/7; infralabials 7/7 (Fig. 1); ven- 
trals 223: cloacal scute divided; subcaudals 
47, all divided except proximal subcaudals 
2-7; dorsal scale rows smooth, in 15 unred- 
uced rows; no supracloacal keels. 

Dimensions are as follows: head length 
15.0 mm, head width 10.9 mm, total length 
639, tail length 87 mm (comprising 13.6% 
of total). 

The black head cap covers the rostral, first 
three supralabials, the upper and anterior 
edge of Supralabial 4, the anterior edge of 
the upper preocular, slightly more than three- 
fourths of the supraoculars, and about half 
of the frontal; there is no pale spot on snout. 
The yellow parietal ring extends from the 
black head cap to include all of the parietals, 
the primary and secondary temporals, and 
anterior part of the tertiary temporals, and 
all of the ultimate supralabial except for the 
posterior edge (Fig. 1). About half or slight- 
ly more of the lingual portion of the mental 
and the first three pairs of infralabials are 
black; the posterior gular area is immaculate 
yellow and continuous with the yellow pa- 
rietal ring. The black nuchal ring does not 
reach the posterior tips of the parietals and 
involves less than half of the first dorsal 
scale situated partially between the parietals. 
The nuchal ring extends posteriorly along 
ime> middorsum ~1/2 + ° 5 + 1/2 dorsal 
scales; it is shortened ventrally to cover most 
of the first two ventrals and adjacent 1.5 
preventrals. There are 24 black body rings, 
including the nuchal, with the posteriormost 
just anterior to the vent. Throughout most of 
the body black rings are about four dorsal 
scale lengths long, but the anterior three 
rings behind the nuchal ring are slightly lon- 
ger, being about 4.5-5.5 scales in length 
(Fig. 2). On the belly, most black body rings 
are reduced to about three ventrals (one ring 
covers only two ventrals and several cover 
four), with an additional half ventral (divid- 
ed along midventral line) associated with 


293 


many rings. Black body rings tend to have 
vertical edges or, if shortened on the side of 
the body, they are reduced by no more than 
about 0.5 scale lengths. The yellow rings are 
immaculate and relatively wide throughout 
the body, being about two dorsal scales long. 
The cloacal scute is yellow. Most red rings 
are 3—4 scales in length with a few reduced 
to as short as one and one-half to two scale 
lengths on the posterior half of the body. 
Scales in the red rings are mostly immacu- 
late but with slightly dusky free margins and 
there are a few irregular, inconspicuous 
black dots in a few of the red rings. There 
are five black tail rings that are one and one- 
half to three times the width of the yellow 
rings separating them; the tip of the tail is 
yellow. 

Variation.—The paratype (UTA R- 
17145) is a subadult male, 327 mm in TL, 
with a tail length of 36, comprising 11.0% 
of the total. It agrees with the holotype in 
most respects of scutellation and pattern, 
but has 220 ventrals, 43 subcaudals (sub- 
caudals 2—3 not divided); a black nuchal 
ring involving only the first ventral plus ad- 
jacent two and one-half preventrals; 27 
black body rings, most of which are about 
four dorsal scales in length throughout the 
body; red body rings which are two to three 
dorsal scales in length throughout the body; 
and 6 black tail rings, exclusive of the black 
tail tip, which are 1.5—2 times longer than 
the yellow rings separating them. 

Etymology.—The species name is a noun 
in the genitive case, formed in honor of 
Emiglio Pacheco Gil, a good friend and 
longtime resident of the Zapotitlan Valley. 
He, his wife, and about twelve children (I 
never knew the total number and am not 
sure he did either) always were gracious 
and hospitable hosts during my visits, most 
willing to share whatever meager provi- 
sions they had available. The country 
around Zapotitlan is harsh and Emiglio sup- 
plemented his income by working in the 
onyx mines which abound in the region. He 
was killed in a mining accident in 1982. 

Remarks.—The morphology of coral 


fl 


| 


| & 


VOLUME 113, NUMBER 1 


Fig. 2. 


snakes is extremely conservative (Slowin- 
ski 1995) and the numbers and arrangement 
of most head scales, rows of dorsal scales, 
and even skull and dentition is remarkably 
consistent, with minor exceptions, through- 
out Micrurus (Campbell & Lamar 1989, 
Roze 1996). Perhaps, the most dramatic ad- 
aptation within the genus is the conspicuous 
elongation of the body apparent in certain 
coral snakes including the genus Leptomi- 
crurus and various species of Micrurus in 
Lower Central America and northern South 
America. Many species of coral snakes are 
diagnosed primarily on the basis of color 
pattern and this has proven to be a generally 
reliable method for distinguishing most spe- 
cies, although a few species are notoriously 
variable (e.g., Micrurus diastema) which 
has resulted in a tangled taxonomic history 
for several species. 

The habitats of coral snakes also may be 
useful for species identification, although 
many species occur over several ecological 
associations and may have elevational dis- 


Dorsal aspect of Micrurus pachecogili (holotype, UTA R-12546). Total length = 639 mm. 


tributions of over 2000 m. Nevertheless it is 
worth noting that heretofore only a single 
species of coral snake has been reported 
from southern Puebla, namely M. Jaticollar- 
is, a distinctive species and one of only two 
species in Mexico possessing black body 
rings arranged in triads. Micrurus laticollar- 
is occurs on the west coast of Mexico in 
Jalisco, Colima, and Michoacan and through 
much of the Balsas Basin and associated 
tributaries into southern Puebla where it may 
be sympatric with M. pachecogili. 

The distributions of a number of other 
species of Micrurus, all with monadal black 
body rings, approach the Zapotitlan Valley 
within about 150 km, but are characterized 
by markedly different habitats. The region 
around Zapotitlan is dry, being in a rain- 
shadow valley. This region receives about 
250-350 mm of precipitation annually, and 
the valley floor and surrounding slopes are 
covered by arid scrub forest dominated by 
many species of cactus. Micrurus bernadi 
occurs in northern Puebla in tropical ever- 


296 


{a . 
Ue 


ea ee) 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Oaxaca 


A Micrurus pachecoglili 
@ Micrurus nebularis 


Pacific Ocean 


97° 


Bie 33: 


green forest and cloudforest at elevations of 
50 to about 2000 m. There is some evidence 
that this taxon is conspecific with M. dia- 
stema (Pérez-Higareda & Smith 1990), also 
known from the eastern versant of Mexico, 
but this allocation has not been universally 
accepted (Roze 1996). Micrurus browni is 
known from the subhumid coastal tropical 
forest, tropical deciduous forest, and dry 
pine-oak forest of southern Mexico from 
about sea level to over 2000 m. The ex- 
tremely variable M. diastema inhabits trop- 
ical evergreen forest and cloudforest of 
eastern Puebla and adjacent Veracruz and 
Oaxaca, as well as in subhumid forests over 


96° 


Distributions of two species of coral snakes, genus Micrurus, from the highlands of southern Mexico. 


most of the Yucatan Peninsula. It ranges 
from near sea level to about 1250 m. Mi- 
crurus ephippifer zapotecus occurs in high- 
land pine-oak forest of central Oaxaca at 
elevations of 1700—2400 m; another appar- 
ently closely related population, M. e. 
ephippifer, occurs at lower elevations (100— 
1500 m) in tropical deciduous forest and in 
the ecotone of this forest with pine-oak for- 
est. Micrurus nebularis occurs in pine-oak 
forest at elevations of 2100—2300 m on the 
southern slopes of the Sierra de Juarez in 
central Oaxaca; all known specimens have 
been collected in the vicinity of Ixtlan de 
Juarez (Fig. 3). 


VOLUME 113, NUMBER 1 


During the course of my investigations 
around Zapotitlan, I collected a series of 
distinctive Lampropeltis triangulum which 
subsequently became the type-series of a 
new subspecies (Quinn 1983). This snake 
closely resembles M. pachecogili, having 
relatively long yellow body rings and nar- 
row red rings, thus serving as one more 
compelling example of mimicry. 


Acknowledgments 


I thank the Officials from the Direcci6n 
General de Flora y Fauna Silvestre for is- 
suing collecting permits for Mexico. How- 
ard Arnott kindly took the photograph in 
Fig. 2. For help at various times in the field, 
I appreciate the efforts of Barry Armstrong, 
William Lamar, and David Hillis. 


297 


Literature Cited 


Campbell, J. A., & W. W. Lamar. 1989. The venomous 
reptiles of Latin America. Cornell University 
Press, Ithaca, 425 pp. 

Pérez-Higareda, G., & H. M. Smith. 1990. The endem- 
ic coral snakes of the Los Tuxtlas region, south- 
ern Veracruz, Mexico.—Bulletin of the Mary- 
land Herpetological Society 26:5—13. 

Quinn, H. 1983. Two new subspecies of Lampropeltis 
triangulum from Mexico.—Transactions of the 
Kansas Academy of Science 86:113-—135. 

Roze, J. A. 1996. Coral snakes of the Americas: bi- 
ology, identification, and venoms. Krieger Pub- 
lishing Company, Malabar, Florida, 328 pp. 

Slowinski, J. B. 1995. A phylogenetic analysis of the 
New World coral snakes (Elapidae: Leptomicru- 
rus, Micruroides, and Micrurus) based on al- 
lozymic and morphological characters.—Jour- 
nal of Herpetology 29:325—338. 

West, R. C. 1964. Surface configuration and associated 
geology of Middle America. Pp. 33-83 in R. 
Wauchope and R. C. West, eds., Handbook of 
Middle American Indians, vol. 1, Natural En- 
vironment and Early Cultures. University of 
Texas Press, Austin, 570 pp. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


113(1):298-—301. 2000. 


Fossil Red-shouldered Hawk in the Bahamas: Calohierax quadratus 
Wetmore synonymized with Buteo lineatus (Gmelin) 


Storrs L. Olson 


Department of Vertebrate Zoology, National Museum of Natural History, Smithsonian Institution, 
Washington, D.C. 20560, U.S.A. 


Abstract.—A supposedly extinct genus and species of hawk, Calohierax 
quadratus Wetmore, was originally described from a fragmentary tarsometa- 
tarsus from Quaternary deposits in Little Exuma Island in the Bahamas. This 
and a referred tibiotarsus from New Providence, Island, were later assigned to 
the extant genus Buteo, but their specific identity remained uncertain. A pre- 
viously unstudied humerus from a cave deposit on New Providence Island, 
Bahamas, is here identified with the extant Red-shouldered Hawk, Buteo li- 
neatus Gmelin, a species widespread in eastern North America and common 
in peninsular Florida but unknown in the Bahamas. The other fossils are as- 
signed to this species as well. Calohierax quadratus therefore becomes a syn- 
onym of Buteo lineatus, which species has retreated from the Bahamas in the 
late Quaternary for reasons that are unclear. 


Among the severai new species of birds 
that were described by Wetmore (1937) 
from Quaternary cave deposits on Little 
Exuma Island, Bahamas, was a new genus 
and species of hawk, Calohierax quadratus, 
based on the distal end of a tarsometatarsus. 
The site was not on Great Exuma as Wet- 
more originally reported (see Olson and 
Pregill 1982:3). Brodkorb (1959) later re- 
ferred the distal end of a tibiotarsus from a 
cave deposit on New Providence Island, 
Bahamas, to the same species. Restudy of 
these specimens showed that the supposed 
characters of the genus Calohierax were 
founded entirely on artifacts of wear in the 
case of the holotype, or intrageneric varia- 
tion in the case of the referred specimen 
(Olson & Hilgartner 1982). The specimens 
were otherwise considered to be too frag- 
mentary for specific identification, falling 
within the range of size variation of the liv- 
ing Red-shouldered Hawk Buteo lineatus 
(Gmelin) or Gray Hawk B. nitidus (La- 
tham). 

Under present geographical and climatic 
conditions, the Red-shouldered Hawk, 


which occurs through most of eastern North 
America and is abundant in Florida, would 
be considered a more likely candidate for 
having occurred in the Bahamas. The Neo- 
tropical Gray Hawk now occurs only as far 
north and east as Arizona and Texas. Nev- 
ertheless, there is ample precedent for var- 
ious Neotropical and western vertebrates 
having occurred in eastern North America, 
particularly Florida, in the Pleistocene (Em- 
slie, 1998), so the possibility of Buteo ni- 
tidus possibly having once occurred in the 
Bahamas is not unthinkable. 

The Bahaman fossil hawk was thus listed 
simply as “‘“Buteo sp.’’, with the genus Ca- 
lohierax Wetmore falling into the synony- 
my of Buteo Lacepede, 1799 (Olson & Hil- 
gartner 1982). The idea that there was an 
extinct hawk in the Bahamas has lingered, 
however, and Calohierax quadratus has 
even appeared, in a reference that I can no 
longer recall, under the absurd name 
‘‘Quadrate Hawk.’’ Wetmore’s specific 
name ‘“‘guadratus’’, it should be noted, was 
derived from the squared appearance of one 
of the tarsometatarsal trochleae, a feature 


VOLUME 113, NUMBER 1 


that turned out to be only an artifact of wear 
(Olson & Hilgartner 1982). 

Because of the ambiguities surrounding 
the records of a medium-sized species of 
Buteo in the Bahamas, it is fortunate that 
an additional specimen bearing on its iden- 
tity was located in the collections of the 
Florida Museum of Natural History. This is 
a nearly complete right humerus (UF 
41801), lacking only a portion of the pec- 
toral crest. It was collected by J. C. Dick- 
inson and W. Auffenberg in the same “‘Ba- 
nana Hole”? on New Providence Island that 
yielded the fossils that formed the basis of 
Brodkorb’s (1959) study and much of that 
of Olson & Hilgartner (1982). It was col- 
lected in 1958-1960, evidently after Brod- 
korb’s study was completed, and since its 
collection has apparently been overlooked. 

The fossil humerus is in all details and 
proportions identical with that in Buteo li- 
neatus (Fig. 1) and in size falls squarely 
among males from Florida (Fig. 2), which 
belong to the southeastern subspecies B. /. 
alleni Ridgway, which is smaller than the 
nominate subspecies (Crocoll 1994). From 
the small sample of Buteo nitidus exam- 
ined, it is clear not only that B. nitidus is a 
smaller species, with females in the range 
of males of B. lineatus in length of the hu- 
merus and the single male being much 
smaller (Fig. 2), but the humerus is also 
much more robust, the shaft especially be- 
ing much thicker. Assuming that there was 
only one species of Buteo in the Bahamas 
in this size range, then the holotypical tar- 
sometatarsus of Calohierax quadratus from 
Little Exuma would belong to the same spe- 
cies as the humerus from New Providence, 
both islands being on the Great Bahama 
Bank. Therefore the species Calohierax 
quadratus Wetmore, 1937, becomes a ju- 
nior subjective synonym of Falco lineatus 
Gmelin, 1788. 

The Red-shouldered Hawk is normally a 
bird of moist riparian woods or swamp- 
lands, hardly like the dry, scrubby habitats 
that predominate in the Bahamas today. 
Brown and Amadon (1968:578) remark that 


299 


this species “‘seems to be incompatible 
with”’ the larger Red-tailed Hawk, B. ja- 
maicensis (Gmelin), although this is as 
much a reflection of the preference of the 
latter for drier uplands rather than being due 
to competition or antipathy. In any case, it 
is the Red-tailed Hawk that occurs in the 
Bahamas today, although it is an uncom- 
mon resident only on some of the larger 
northern islands and may perhaps be a re- 
cent colonist, as it is absent in the fossil 
record. 

Prior to the arrival of Europeans only a 
single terrestrial mammal lived in the Ba- 
hamas, the hutia Geocapromys ingrahami 
(Allen), which is now extinct on all but a 
single small islet. The adults of this species 
are too large to have served as prey for 
Red-shouldered Hawks, but because these 
hawks are very catholic in their choice of 
food, taking birds, reptiles, amphibians, 
large insects, and even crustaceans in ad- 
dition to mammals (Crocoll 1994), and be- 
cause all of these faunal elements are still 
present in the Bahamas, it would be diffi- 
cult to correlate the extinction of Buteo li- 
neatus there to lack of suitable prey. 

Habitat in most places in the Bahamas is 
not now like that usually considered suit- 
able for Red-shouldered Hawks, but is 
thought to have been even more xeric in the 
past (Pregill & Olson 1981, Olson & Hil- 
gartner 1982). Thus, environmental and cli- 
matic changes are also difficult to invoke as 
an explanation for the disappearance of this 
species from the Bahamas in the late Qua- 
ternary. 

The Bahaman fossil records constitute 
the only evidence of Buteo lineatus any- 
where in the West Indies, although B. ridg- 
wayi of Hispaniola is now usually consid- 
ered to be a derivative of B. lineatus (e.g., 
Sibley & Monroe 1990), so the Bahamas 
may once have provided a stepping-stone 
for this colonization. 


Acknowledgments 


I am most grateful to the staff of the Flor- 
ida Museum of Natural History (UP), 


300 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 1. Right humeri of Buteo in anconal aspect: A, male Red-shouldered Hawk, B. lineatus, (UF 23893); 
B, fossil referred to B. lineatus from New Providence Island, Bahamas (UF 41801); C, female Gray Hawk, B. 
nitidus (UF 33746). Scale bar = 2 cm. 


17 


16 


iis 


14 


humerus, distal width 


1c 
70 80 90 100 


humerus length 


Fig. 2. Scatter diagram showing length of humerus vs. distal width of humerus in male Buteo lineatus 
(squares), female B. lineatus (triangles), the Bahaman fossil assigned to B. lineatus (star), female B. nitidus 
(circles), and male B. nitidus (X). Five obviously missexed specimens have been corrected. All specimens of 
B. lineatus are from Florida and are in the collections of the Florida Museum of Natural History. 


VOLUME 113, NUMBER 1 


Gainesville, for access to and loan of spec- 
imens that made this study possible: David 
W. Steadman and Tom Webber for modern 
birds, and Marc Frank for the fossils. The 
photograph is by John Steiner, Smithsonian 
Photographic Services, and Fig. 2 was pre- 
pared by Helen James. 


Literature Cited 


Brodkorb, P. 1959. Pleistocene birds from New Prov- 
idence Island, Bahamas.—Bulletin of the Flor- 
ida State Museum, Biological Sciences 4(11): 
349-371. 

Brown, L., & D. Amadon. 1968. Eagles, hawks, and 
falcons of the world. 2 vols. McGraw Hill, New 
York, 945 pp. 

Crocoll, S. T. 1994. Red-shouldered Hawk Buteo li- 
neatus.—Birds of North America 107:1—19. 

Emslie, S. D. 1998. Avian community, climate, and 
sea-level changes in the Plio-Pleistocene of the 


301 


Florida Peninsula.—Ornithological Mono- 
graphs 50:1—113. 

Olson, S. L., & W. B. Hilgartner. 1982. Fossil and 
subfossil birds from the Bahamas. Pp. 22—56 in 
S. L. Olson, ed. Fossil vertebrates from the Ba- 
hamas.—Smithsonian Contributions to Paleo- 
biology 48. 

, & G. K. Pregill. 1982. Introduction to the pa- 
leontology of Bahaman vertebrates. Pp. 1—7 in 
S. L. Olson, ed., Fossil vertebrates from the Ba- 
hamas.—Smithsonian Contributions to Paleo- 
biology 48. 

Pregill, G. K., & S. L. Olson. 1981. Zoogeography of 
West Indian vertebrates in relation to Pleisto- 
cene climatic cycles.—Annual Review of Ecol- 
ogy and Systematics 12:75—98. 

Sibley, C. G., & B. L. Monroe, Jr. 1990. Distribution 
and taxonomy of birds of the world. Yale Uni- 
versity Press, New Haven, 1111 pp. 

Wetmore, A. 1937. Bird remains from cave deposits 
on Great Exuma Island in the Bahamas.—Bul- 
letin of the Museum of Comparative Zoology 
80:427—441. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 
113(1):302-—318. 2000. 


Revision of the extant taxa of the genus Notiosorex 
(Mammalia: Insectivora: Soricidae) 


Leslie N. Carraway and Robert M. Timm 


(LNC) Nash 104, Department of Fisheries and Wildlife, Oregon State University, Corvallis, 
Oregon 97331-3803, U.S.A.; (RMT) Natural History Museum & Department of Ecology and 


Evolutionary Biology, University of Kansas, Lawrence, Kansas 66045-2454, U.S.A. 


Abstract.—We evaluated the taxonomic status of three specimens of gray 
shrews, Notiosorex (Insectivora: Soricidae), collected in isolated mountain val- 
leys in Tamaulipas, Mexico, with specimens referable to the Recent taxa N. 
crawfordi crawfordi (n = 229) and N. c. evotis (n = 34). Statistical analyses 
of cranial morphometric data revealed that the specimens from Tamaulipas 
represented a heretofore unknown taxon which is described herein as a new 
species of Notiosorex (N. villai). Also, N. c. evotis was found to be equally 
distinct from N. c. crawfordi, thus was elevated to specific status. 

Resumen.—Se evalu6 el estado taxonémico de tres especimenes de la mu- 
sarana gris Notiosorex (Insectivora: Soricidae), colectados en valles dentro de 
las motanas en Tamaulipas, México y referidos a los taxa recientes N. crawfordi 
crawfordi (n = 229) y N. c. evotis (n = 34). Analisis estadisticos morfométricos 
del craneo indicaron que los especimenes de Tamaulipas representan un taxon 
desconocido, el cual se describe como una especie nueva de Notiosorex (N. 
villai). Asimismo, N. c. evotis se encontr6é distinto a N. c. crawfordi, por lo 


que se elevo al nivel especifico. 


Gray shrews of the genus Notiosorex (In- 
sectivora: Soricidae) are widely distributed 
in the southwestern United States and 
northern and western Mexico (Fig. 1). They 
range from southern California, Nevada, 
Utah, Colorado, Oklahoma, and western 
Arkansas, southward to southern Baja Cal- 
ifornia Sur, Michoacan, and southern Ta- 
maulipas (Sealander 1952, Baker 1966, 
Armstrong & Jones 1972, Woloszyn & Wo- 
loszyn 1982, Carie et al. 1989). Insular re- 
cords are known from Isla San Martin, Baja 
California Norte (Schulz et al. 1970), and 
Isla Palmito del Verde, Sinaloa, Mexico 
(Armstrong & Jones 1971). Gray shrews 
occur in a variety of habitats including de- 
sert shrub (characterized by mesquites, Pro- 
sopis; palo verde, Cercidium; Acacia; Yuc- 
ca; Agave; and scattered Juniperus; Lange 
1959), pine-oak forest (characterized by 
Abies religiosa, Populus tremuloides, Jun- 


iperus flaccida, four species of Pinus, and 
three of Quercus; Alvarez 1963), in grass- 
land with oak chaparral (characterized by 
chamise, Adenostomna fasciculatum; scrub 
oak, Quercus dumosa; California live oak, 
Q. agrifolia; and mountain lilac, Ceano- 
thus) and oak woodland habitats nearby 
(Cunningham 1956), coastal sage scrub 
(characterized by coastal sagebrush, Arte- 
mesia californica; black sage, Salvia mel- 
lifera; laurel sumac, Rhus laurina; and 
grasses), yellow pine forest (Pinus austral- 
is; Lange 1959), alkaline marsh (Stephens 
1906), arid grasslands (containing scattered 
catclaw, juniper, and mesquite; Baker 
1966), and sandy flats (characterized by Ar- 
temisia tridentata, Ephedra viridis, Peuce- 
phyllum schottii, and Chrysothamnus vis- 
cidiflorus; Fisher 1941). Elevations of oc- 
cupied habitats range from 3 to 2618 m 
(Fisher 1941, Lange 1959, Baker 1966, Da- 


VOLUME 113, NUMBER 1 


303 


Fig. 1. 
crawfordi (closed circles); 2, N. c. evotis (closed squares); and 3, Notiosorex unknowns from Tamaulipas (tri- 
angles). Open circles, N. c. crawfordi, and open squares, N. c. evotis, represent collection localities for which 
specimens were not examined (Bailey 1905; von Bloeker 1944; Cunningham 1956; Davis 1960; Baker 1966; 
Dalby & Baker 1967; Armstrong 1972; Armstrong & Jones 1972; Findley et al. 1975; Sealander 1979; Hoff- 
meister 1986; Caire et al. 1989; Rodriguez Vela 1999; Angeles Mendoza Duran, pers. comm.; Philip Myers, 
pers. comm.). 


vis & Sidner 1989). Because of the wide 
range of habitats in which Notiosorex is 
know to occur, the commonly used vernac- 
ular name “‘desert shrew”’ (Hall 1981) is a 
misnomer. Herein, we recommend the use 
of gray shrew. 

In 1953, Gerd H. Heinrich collected two 
specimens of Notiosorex and in 1976 
George D. Baumgardner collected a third 
from western Tamaulipas. Findley (1955: 
616) unequivocally categorized the former 
two specimens as WNotiosorex crawfordi 
crawfordi ‘‘on geographic grounds,”’ 
whereas Alvarez (1963:397) later referred 


Distribution of gray shrews (Notiosorex) based upon specimens examined herein: 1, N. crawfordi 


them to N. crawfordi noting that ““When 
more abundant material is available the No- 
tiosorex crawfordi of northeastern México 
probably will be found to represent a new 
subspecies.’” Schmidly & Hendricks (1984: 
23) examined all three specimens and re- 
ferred them to N. c. crawfordi because it 
was “‘the name currently applied to desert 
shrews in Texas and northern México.”’ As 
these Tamaulipian shrews are isolated geo- 
graphically from the widespread WN. c. craw- 
fordi and are much larger and quantitatively 
different from that taxon, we evaluated their 
taxonomic status. 


304 


Materials and Methods 


Specimens from throughout the distribu- 
tion of Notiosorex (n = 266) were exam- 
ined. Seven cranial and five mandibular 
characters were recorded for each of the 
139 specimens measured (Fig. 2). Relative 
age of specimens was indexed by the lateral 
length of the right I1 (first upper incisor; 
Carraway et al. 1996). Quantitative char- 
acters were analyzed by age to determine if 
age caused a bias. Greatest length of skull, 
rostral breadth, least interorbital breadth, 
and cranial breadth were measured to 0.01 
mm with Mitutoyo Digimatic electronic 
calipers. All other quantitative characters 
were measured by use of an ocular micro- 
meter mounted in a Bausch and Lomb bin- 
ocular microscope. Values were converted 
from number of ocular lines to millimeters 
for multivariate analyses and tabulation of 
reported values. The states of a qualitative 
character, roof of glenoid fossa extending 
laterally from side of cranium, also were 
recorded for each specimen. 

Three a priori groups were formed of in- 
dividuals of Notiosorex crawfordi crawfor- 
di, N. c. evotis, and the three specimens 
from Tamaulipas; assignment of individuals 
to their a priori group was based on the geo- 
graphic location of their collection site. 
Multivariate analyses of the three a priori 
groups were performed on the 12 quanti- 
tative cranial and mandibular characters by 
use of multigroup discriminant-function 
analysis in BIOXTAT II (Pimentel 1995). A 
large sample of N. c. crawfordi (n = 54) 
from Huachuca Mts., Arizona, was exam- 
ined for intraspecific geographic variation 
and a comparison with the remaining N. c. 
crawfordi was performed with multigroup 
discriminant-function analysis. Univariate 
and regression analyses were calculated by 
use of STATGRAPHICS Plus (Statistical 
Graphics Corporation 1995). For all analy- 
ses, P < 0.05 was accepted as statistically 
significant. 

Standardized canonical vectors for the 12 
quantitative variables were plotted on the 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


same graph as the three a priori groups with 
the same canonical variate axes. The plot of 
the variable vectors, when placed at the or- 
igin of the canonical variates plot, provided 
an indication of the direction and magni- 
tude of effect that each variable had in dis- 
criminating the three a priori groups (Joli- 
coeur, 1959). 

Summary statistics (CX SE. range, and 
CV) were calculated for all 12 quantitative 
variables for the three taxa of Notiosorex 
under consideration (Table 1). Color of hair 
in the pelage was determined by compari- 
son with Munsell soil color charts (Munsell 
Color 1975). 


Results 


Multigroup discriminant function analy- 
sis of Notiosorex crawfordi crawfordi, N. c. 
evotis, and Notiosorex unknowns from Ta- 
maulipas, resulted in 99% correct classifi- 
cation of individuals into their a priori 
groups (Fig. 3). The two significant canon- 
ical-variate axes (x7 = 153.61, df. = 24 
and x”? = 37.73, df. = 11, respectively) 
acounted for 81.10 and 18.90% of the var- 
iation, respectively. Variance in the first 
axis (canonical variate I) was accounted for 
by greatest length of skull, 98.19%; cranial 
breadth, 99.03%; length of unicuspid tooth- 
row, 98.35%; width across M2-M2, 
92.69%; length of mandible, 99.39%; 
length of mandibular toothrow, 89.70%; 
height of coronoid process, 96.08%; and 
length of the coronoid process-ventral point 
of upper condylar facet, 97.64%. The sec- 
ond axis (canonical variate II) was affected 
by length of the coronoid process-ventral 
point of lower condylar facet (89.98%). 
Only one individual (KU 105409), from 5 
mi WNW EI Carrizo, Sinaloa, Mexico, was 
not placed into its a priori group, WN. c. ev- 
otis; it was classified as a N. c. crawford. 
This individual also was classified as a N. 
c. crawfordi based on morphological com- 
parisons by Jones et al. (1962). Herein, this 
individual is considered N. c. crawfordi for 
purposes of further analyses. 


VOLUME 113, NUMBER 1 305 


glenoid 
fossa 


ar 


4 
11 
9 
10 
LUC 


Fig. 2. Camera-lucida tracing of skull of a Notiosorex (KU 145262) illustrating skull dimensions measured: 
1, greatest length of skull; 2, rostral breadth; 3, least interorbital breadth; 4, cranial breadth; 5, length of maxillary 
unicuspid toothrow; 6, length of maxillary complex toothrow; 7, width across M2—M2; 8, length of mandible; 
9, length of mandibular toothrow; 10, height of coronoid process; 11, length of coronoid process-ventral point 
of upper condylar facet; and 12, length of coronoid process-ventral point of lower condylar facet. Qualitative 
character recorded is: 13, roof of glenoid fossa extending laterally from the side of the skull. Scale bar equals 
5 mm. 


306 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


CANONICAL VARIATE II 


-6.2 -4.2 


-2.2 


1.8 


-0.2 


CANONICAL VARIATE | 


Big. 3: 


Canonical-variates plot of Notiosorex c. crawfordi (open circles), N. c. evotis (closed circles), and 


Notiosorex new sp. (triangles) based on 12 measured variables. Canonical-variate axis I accounted for 81.10% 
and canonical-variate axis II 18.90% of the variation present among the three taxa. Differences in cranial 
morphology among the three taxa are characterized by the plot of the 12 variable vectors on the canonical- 
variates plot in the upper right corner. Numbers are as in Fig. 2. The relationship of the variables among the 
taxa are indicated by the length and orientation of the vector relative to the canonical-variate axes. From left to 
right, taxa have shorter skulls relative to cranial breadth and taller and narrower coronoid processes. From bottom 
to top, taxa have greater rostral breadth relative to length of maxillary complex toothrow. 


Coefficients of variation for the N. c. 
crawfordi from Huachuca Mts., Arizona, 
ranged from 2.276% to 4.918% with one 
outlier at 6.46%. Whereas, the same values 
for the remaining N. c. crawfordi ranged 
from 3.028% to 8.098%. This indicates 
that among specimens of Notiosorex from 
the Huachuca Mts. the level of variation 
within and among the 12 quantitative char- 
acters examined is very low. The one dis- 
criminant function produced by a multi- 
group discriminant-function analysis of the 


two a priori groups of N. c. crawfordi was 
not significant, at PP .< 0.05. Therefore: 
these two a priori groups are acting as a 
single unit. 

Regression analyses indicated weak re- 
lationships between age and length of uni- 
cuspid toothrow for crawfordi (r? = 0.24) 
and evotis (r? = 0.53) and between age and 
least interorbital breadth (r? = 0.04) and 
length of mandible (7? = 0.04) for crawfor- 
di. When the discriminant analysis was re- 
peated without these three variables, 93.5% 


VOLUME 113, NUMBER 1 


of individuals still were classified correctly 
into their a priori groups. 

For all N. c. crawfordi and N. c. evotis 
the roof of the glenoid fossa extends later- 
ally from the cranium. However, among the 
Notiosorex from Tamaulipas the roof of the 
glenoid fossa does not extend laterally from 
the cranium. 


Discussion 


Morphometrically, the three specimens 
of Notiosorex from Tamaulipas are most 
similar to each other and are distinct from 
all specimens of crawfordi and evotis (Fig. 
3) in terms of size and shape of their skulls. 
We had expected this outcome after making 
careful visual comparisons (with aid of a 
binocular microscope) of the three groups. 
The discovery that evotis was equally dis- 
tinct from the two other groups (Fig. 3) ne- 
cessitated a reevaluation of its taxonomic 
status. Previously, N. c. evotis was recog- 
nized as being larger than N. c. crawfordi, 
but we also found significant differences in 
the shape of skulls and mandibles between 
individuals of the two taxa. 

Based on analyses reported herein we re- 
fer the three specimens from Tamaulipas to 
a new species and elevate N. c. evotis to 
species level. 


Class Mammalia Linnaeus, 1758 
Order Insectivora Bowdich, 1921 
Family Soricidae Fischer von Waldheim, 
1817 
Subfamily Soricinae Fischer von 
Waldheim, 1817 
Genus Notiosorex Coues, 1877 
Notiosorex villai, new species 
Villa’s Gray Shrew 
Fig. 4A 


Notiosorex crawfordi.—Findley 1955:616, 
Ball & Kelson 1959:64 [part], Alvarez 
1963:397, Hall 1981:65 [part], Schmidly 
& Hendricks 1984:22. 


Holotype.—Adult, female, skin and 
skull; KU 54932, University of Kansas, 


307 


Natural History Museum, Mammal Collec- 
tion; from “‘Jaumave, Tamaulipas, Mexico, 
2400 ft.”’; obtained 26 July 1953 by Gerd 
H. Heinrich, original number 7612. Alvarez 
(1963:386) recorded the latitude and lon- 
gitude of Jaumave as 23°34’N, 99°23’W. 

Distribution.—Known only from the Po- 
tosian Biotic Province (=Sierra Madre Ori- 
ental Biotic Province) of the central moun- 
tains of Tamaulipas, Mexico (Fig. 1). 

Diagnosis.—As in all Notiosorex, speci- 
mens of N. villai (Fig. 4A) have a combi- 
nation of a deeply emarginated area be- 
tween the condylar processes, e.g., interar- 
ticular breadth about half the width of the 
superior condylar process; the alveolus of 
il extending posteriorly beneath at least 
part of paraconid of ml (Carraway 1995); 
pigment present of some teeth; and three 
unicuspids. 

Notiosorex villai can be distinguished 
from other Notiosorex by the roof of the 
glenoid fossa not extending laterally from 
the cranium when the skull is viewed from 
the dorsal aspect (Fig. 4A); and usually 
from N. c. crawfordi by greatest length of 
skull 216.97 mm and from N. c. evotis by 
length of maxillary unicuspid toothrow 
=2.0 mm, height of coronoid process <4.1 
mm, length of coronoid process-ventral 
point of upper condylar facet =3.8 mm, and 
length of coronoid process-ventral point of 
lower condylar facet =3.4 mm. All speci- 
mens of N. villai can be separated from 
specimens of N. c. crawfordi and N. c. ev- 
otis by application of the following discrim- 
inant-function equation: discriminant score 
= 1.30722 (cranial breadth) — 0.34104 
(height of coronoid process) — 0.01685 
(ength of maxillary complex toothrow) — 
4.87675 (least interorbital breadth) — 
0.10742 (length of coronoid process-central 
point of lower condylar facet) — 0.16575 
(length of mandible) + 0.22460 (length of 
mandibular toothrow) + 0.88996 (greatest 
length of skull) + 0.22980 (length of max- 
illary unicuspid toothrow) — 0.03089 
(length of coronoid process-ventral point of 
upper condylar facet) + 6.66117 (rostral 


308 


breadth) — 0.23311 (width across M2—M2) 
— 11.0491. Those specimens with scores 
=3.359 are referrable to villai and those 
with scores =2.454 are referrable to craw- 
fordi or evotis. 

Description.—As in all soricids, Notio- 
sorex villai has a double-faceted condylar 
process and a fissident I1 with a large hook- 
shaped anterior cusp and a posterior ven- 
trally directed cusp. The skull is moderately 
large, smooth, and without prominent ridg- 
es and processes. The paroccipital processes 
are small and lie against the exoccipitals 
(Fig. 4A). The upper condylar facets are in- 
flected; the corresponding areas in the su- 
perior portion of the glenoid fossas are de- 
pressed. 

The pelage is composed of multibanded 
hairs. In summer, hairs of the dorsal pelage 
have a narrow band of silver gray (LOYR 
6/1) distally and a wide band of dark gray 
(7.5YR N4/O) proximally. Hairs of the ven- 
ter are the same colors, except that the dis- 
tal band of silver gray is wide and the prox- 
imal band of dark gray is narrow producing 
a silver wash. In winter, hairs of the dorsal 
pelage have a narrow band of very dark 
grayish-brown (1OYR 3/2) distally, a nar- 
row band of pinkish white (7.5YR 8/2) me- 
dially, and a wide band of very dark-gray 
(7.5YR N3/O) proximally. Hairs of the ven- 
tral pelage have a distal wide band of pink- 
ish white (7.5YR 8/2) and a proximal nar- 
row band of very dark-gray. The tail is very 
dark grayish-brown. 

Measurements.—Individuals of Notioso- 
rex villai are smaller than those of N. c. 
evotis, but larger than those of N. c. craw- 
fordi (Table 1). 

Ecology.—The three known specimens 
of Notiosorex villai were collected in dif- 
ferent habitats within the Potasian Biotic 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Province (Alvarez 1963). The area of Jau- 
mave is pine-oak forest, that of Palmillas is 
tropical forest, and the habitat of the col- 
lection site near Rancho Carricitos is ripar- 
ian. The subprovince, within the Potosian 
Biotic Province, from which the three spec- 
imens were collected contains ‘“‘low mese- 
ta-like folded mountains” characterized by 
“closely spaced ridges’”’ (Ferrusquia- Villa- 
franca 1993:33). 

Mammal associates of the holotype were 
Baiomys taylori, Onychomys arenicola, 
Oryzomys couesi, Peromyscus leucopus, 
Reithrodontomys fulvescens, Sigmodon his- 
pidus, Liomys irroratus, Mephitis ma- 
croura, and Mustela frenata; only Pero- 
myscus pectoralis was collected in the vi- 
cinity of the specimen from Palmillas (G. 
H. Heindrich, in litt.). Cryptotis parva, 
Mormoops megalophylla, Desmodus rotun- 
dus, Myotis californicus, Lasiurus borealis, 
L. cinereus, Antrozous pallidus, Tadarida 
brasiliensis, Sylvilagus floridanus, Sciurus 
aureogaster, S. alleni, Liomys irroratus, 
Peromyscus leucopus, P. pectoralis, P. boy- 
lii, Baiomys taylori, Mus musculus, Bassar- 
iscus astutus, and Mephitis mephitis were 
collected in the vicinity of the specimen 
from SW of Rancho Carricitos (Schmidly 
& Hendricks 1984). One of the C. parva 
was collected in the same pitfall as the No- 
tiosorex. The greater species richness noted 
for the latter locality is almost certainly the 
result of greater trapping effort and not nec- 
essarily an indication that the other mam- 
mal associations were depauperate. 

Etymology.—The species epithet is a pat- 
ronymic to honor Bernardo Villa-R., the 
‘‘father’”’ of Mexican mammalogy. 

Remarks.—All three specimens of Notio- 
sorex villai were collected in isolated 
mountain valleys. The extent of the distri- 


— 


Fig. 4. Camera-lucida tracings of dorsal, lateral, and ventral views of the cranium, and lateral view of labial 
side of mandible and oblique view of posterior portion of lingual side of left mandible of taxa of Notiosorex. 
A, Holotype of Notiosorex villai (KU 54932, adult female). Note smooth lateral edge of cranium when viewed 
from dorsal aspect (solid arrow), absence of extension of roof of glenoid fossa, small paroccipital processes 


VOLUME 113, NUMBER 1 309 


lying against the exoccipitals (open arrow), and coronoid process slender relative to height. B, N. evotis (KU 
89214, adult male). Specimen from 17 km SW Choix, Sinaloa. Note prominent ridge on lateral edge of cranium 
caused by extension of roof of glenoid fossa (solid arrow), low-set paroccipital processes extending at an oblique 
angle from skull (open arrow), and coronoid process broad relative to height (dashed arrow). C, N. crawfordi 
(KU 145262, adult male). Specimen from Peloncillo Mts., Guadalupe Canyon, Hidalgo Co., New Mexico. Note 
prominent ridge on lateral edge of cranium caused by extension of roof of glenoid fossa (solid arrow), small 
paroccipital processes lying against the exoccipitals (open arrow), and coronoid process slender relative to height 
(dashed arrow). Scale bar equals 5 mm. 


310 


bution of the species has yet to be deter- 
mined. However, considering the three dif- 
ferent habitat types represented, N. villai 
likely has a reasonably wide distribution in 
Tamaulipas. 

Heinrich, both in his field catalog and on 
the specimen tag of one of the specimens 
included herein, clearly recorded the local- 
ity as Palmilla. However, on a map of Hein- 
rich’s collecting localities drawn on 1 Sep- 
tember 1953 by Hildegarde Heinrich, his 
wife who accompanied him both in 1952 
and 1953, the name is spelled Palmillas. 
The hand-drawn map used by Heinrich in 
1953 is believed to be based on the copy 
of ‘“‘Mapa de la Republica Mexicana—Es- 
tado de los Caminos Federales Estatales y 
Vecinales’’ (1944) on deposit in the Natural 
History Museum at the University of Kan- 
sas. Notations made by E. R. Hall on the 
latter map identify it as a gift to the muse- 
um. The name of the town on this map is 
spelled Palmillas; all subsequent authors 
who have published upon this specimen 
used the spelling Palmillas. 

Specimens examined.—3, as follows. 

Mexico: Tamaulipas: Jaumave, 2400 ft 
(KU 54932, 2); Palmilla [sic], 4400 ft 
[23°18’N, 99°33'W; Alvarez 1963:386] 
(KU 54933, 2); 0.3 mi SW Rancho Carri- 
citos, San Carlos Mts., 1900 ft (TCWC 
30492, <). 


Notiosorex evotis (Coues) 
Large-eared Gray Shrew 
Fig. 4B 


Sorex (Notiosorex) evotis Coues, 1877:652. 

N[otiosorex]. evotis Coues, 1877:652. 

Notiosorex crawfordi evotis Merriam, 1895: 
34; Hall & Kelson 1959:64 [part], Jones 
et al. 1962:151, Hall 1981:65 [part]. 


Holotype.—Adult, no sex given, skin 
only; USNM 9066, National Museum of 
Natural History; from ‘“‘area of Mazatlan, 
Sinaloa, Mexico’”’; obtained February 1868 
by Ferinand Bischoff. The skull is men- 
tioned by Coues (1877), but apparently was 
never cataloged into the National Museum 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


collection. A sketch of the lateral view of 
the right-side rostral area of the cranium 
was included in Dobson (1890). However, 
by the time Merriam (1895) published his 
revision of the genus, the skull was missing. 

Distribution.—Notiosorex evotis is 
known from the states of Colima, Jalisco, 
Michoacan, Nayarit, and Sinaloa, Mexico 
(Fig. 1). 

Diagnosis.—Notiosorex evotis can be 
distinguished from N. villai by the roof of 
the glenoid fossa extending laterally beyond 
the curve of the cranium (Fig. 4B) and usu- 
ally by length of maxillary unicuspid tooth- 
row <2.1 mm, height of coronoid process 
=4.3 mm, length of coronoid process-ven- 
tral point of upper condylar facet =3.8 mm, 
and length of coronoid process-ventral 
point of lower condylar facet =4.0 mm; and 
usually from WN. crawfordi by rostral 
breadth =5.2 mm, width across M2—M2 
=5.1 mm, length of mandible =7.4 mm, 
height of coronoid process 24.3 mm, 
length of coronoid process-ventral point of 
upper condylar facet =4.0 mm, and length 
of coronoid process-ventral point of lower 
condylar facet =3.8 mm. N. evotis usually 
can be distinguished from N. crawfordi by 
use of a combination of height of coronoid 
process and cranial breadth (Fig. 5). Based 
on the results of the discriminant analysis, 
all specimens of N. evotis can be separated 
from specimens of N. crawfordi by appli- 
cation of the following discriminant-func- 
tion equation: discriminant score = 1.73654 
(cranial breadth) + 0.22159 (height of cor- 
onoid process) + 0.06177 (length of max- 
illary complex toothrow) — 0.65656 (least 
interorbital breadth) — 0.00471 (length of 
coronoid process-central point of lower 
condylar facet) + 0.27652 (length of man- 
dible) + 0.08699 (length of mandibular 
toothrow) — 0.86213 (greatest length of 
skull) + 0.23251 (length of maxillary uni- 
cuspid toothrow) — 0.02611 (length of cor- 
onoid process-ventral point of upper con- 
dylar facet) — 0.92248 (rostral breadth) + 
0.10899 (width across M2—M2) — 37.2423. 
Individuals with a score =—2.26 are refer- 


VOLUME 113, NUMBER 1 


rable to evotis and those with a score 
=—1.65 are referrable to crawfordi. 

Description.—Within extant members of 
the genus Notiosorex, evotis has the largest 
skull with prominent ridges (the roof of the 
glenoid fossa) on the lateral sides of the cra- 
nium and the paroccipital processes are set 
low on the mastoid and extend at an oblique 
angle from the skull (Fig. 4B). The coro- 
noid processes are broad relative to their 
height (Fig. 4B; Choate 1969:473, fig. 3b) 
resulting in different placement of the con- 
dyloid processes relative to each other 
(Choate 1969:473, figs. 3b—c) for N. evotis 
and N. crawfordi. 

The pelage is composed of multibanded 
hairs. In summer, hairs of the dorsal pelage 
have a narrow band of very dark grayish- 
brown (1OYR 3/2) distally, a narrow band 
of pinkish white (7.5YR 8/2) medially, and 
a wide band of dark gray (7.5YR N4/0) 
proximally. The hairs of the venter have a 
wide distal band of pinkish white and a nar- 
row proximal band of very dark-gray 
(7.5YR N3/0). The winter pelage consists 
of hairs of the dorsum with a wide distal 
band of very dark grayish-brown (1OYR 3/ 
2) and a proximal band of very dark-gray 
(7.5YR N3/O). The hairs on the venter have 
a wide distal band of light yellowish-brown 
(1OYR 6/4) and a narrow proximal band of 
dark gray (7.5YR N4/O). The hairs on the 
tail are very dark grayish-brown. 

Measurements.—Individuals of Notioso- 
rex evotis are the largest members of extant 
Notiosorex (Table 1). 

Ecology.—Notiosorex evotis is known to 
occur from 3-m elevation along the Pacific 
Coast to 550 m in the Sierra Madre Occi- 
dental, Sinaloa, and to 2317 m in the Sierra 
Moroni, Zacatecas. It is know to occur in 
habitats characterized by scattered cacti and 
dense thornbush, and abandoned agricultur- 
al fields bordered by an area of scattered 
cacti, thornbush, and mesquite (scientific 
names not given—Armstrong & Jones 
1971); communities that “‘consisted mostly 
of dry, dense weeds and short, thorny shrub 
with a few trees” and “‘in low weeds near 


311 


thorn bush’’ (Jones et al. 1962:148—149); 
‘‘*in damp spots under rocky ledges” (Fisher 
& Bogan 1977:826); and in “‘semi-desert 
habitat’? (Schlitter 1973:423). 

Reported small-mammal associates are 
Liomys pictus, Chaetodipus pernix, Sigmo- 
don hispidus, and Mus musculus (Baker 
1962, Jones et al. 1962). 

Etymology.—The species epithet is de- 
rived from the Greek ev, meaning good, and 
otus, Meaning ear, possibly in reference to 
its “‘extremely large’’ ears (Coues 1877: 
652). 

Remarks.—A discussion of the taxonom- 
ic history of Notiosorex evotis is presented 
in Jones et al. (1962). 

Specimens examined.—34, as follows. 

Mexico: Jalisco: 13 mi S, 15 mi W Gua- 
dalajara (KU 33318); 21 mi SW Guadala- 
jara (KU 42583-42585). Michoacan: 2 mi 
E La Palma, SE side of Lago de Chapala 
(KU 42586-42588). Nayarit: El Refilion 
(USNM 508358); Tepic (USNM 314064). 
Sinaloa: 20 km N, 5 km E Badiraguato (KU 
96419); 16 km NNE Choix, 1700 ft (KU 
89210-89213); 1 mi S El Cajoén, 1800 ft 
(KU 100319); 15 mi SE Escuinapa (MSUM 
5691); Isla Palmito del Verde, 6 mi NNW 
Teacapan (KU 98880); Laguna, 17 km SW 
Choix, 500 ft (KU 89214-89216); Maza- 
tlan (KU 85533-85536; USNM 9066); 1 mi 
N Mazatlan, 10 ft (MSUM 8149); 1 mi N 
Mazatlan, 25 ft (MSUM 5690); Rosario, 
500 ft (KU 90581); 10 km S, 38 km E Sin- 
aloa (KU 125476—125479); 44 km ENE 
Sinaloa, 600 ft (KU 89998); HWY 15, 0.25 
mi S Sonora state line (UMMZ 109403). 


Notiosorex crawfordi (Coues) 
Crawford’s Gray Shrew 
Fig. 4C 


Sorex (Notiosorex) crawfordi Coues, 1877: 
651. 

Notiosorex crawfordi Coues, 1877:652, 
True 1884:606, Merriam 1895:32, Hall & 
Kelson 1959:64 [part], Hall 1981:65 


[part]. 
Holotype.—Adult, no sex recorded, skin 


312 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Table 1.—Means (+SE), ranges (in parentheses), and CVs of measurements (in mm) of skull characters of 
Notiosorex crawfordi crawfordi (n = 122), N. evotis (n = 14), and N. villai (n = 3). 


Notiosorex 


Character crawfordi evotis villai 
Greatest length of skull 16.08 + 0.044 F206 O43 LF Le 
(14.95—-17.25) (16.49-18.41) (16.97-17.27)<¢ 
0.03 0.03 
Rostral breadth 4.83 + 0.02 3393" O04 5.19 
(3.70-5.23) (5.0—5.63) (5.01—5.30) 
0.04 0.03 
Least interorbital 3.76 + 001 3.98 + 0.04¢ 3.83 
breadth (3.42-4.14) (3.65-4.16) (3.57-4.01) 
0.04 0.04 
Cranial breadth 796. = 0.022 $567 O07? 8.30 
(7.05—8.52) (8.02—8.99) (8.07—8.53)° 
0.03 0.03 
Length of maxillary 1:9 > 6:01 2.0 + 0.03 ae 
unicuspid toothrow (1.5—2.2) (1.80—2.3) (2.0—2.3) 
0.05 0.05 
Length of maxillary 4.3 + 0.01 ABO =="0.03" 4.6 
complex toothrow (3.9-4.6) (4.5—5.0) (4.44.7) 
0.02 0.02 
Width across M2—M2 4:3 + 0.01 DdyznO: 03° 5.0 
(3.7-S.3) (4.9-5.4) (4.9-5.1) 
0.02 0.02 
Length of mandible 71.0°-= O02" 7.7 = 0.04 7.4 
(6.4—7.7) (7.4-8.2) (7.3-7.5)° 
0.03 0.03 
Length of mandibular 4:7 = 0.014 5.0 2 0:03 5.1 
toothrow (4.4-5.1) (4.9-5.3) (5.0-5.1)¢ 
0.02 0.02 
Height of coronoid AOr= O:02 4720.05 4.1 
process (3.44.9) (4.35.0) (4.0-4.1)° 
0.02 0.04 
Length of coronoid 3:3; HO102¢ 3.9. 0.04 3.8 
process—ventral point (2.7-3.9) (3.84.2) (3.7-3.8)° 
of upper condylar facet 0.06 0.03 
Length of coronoid 347 70102 43° "O05 3.4 
process—ventral point (3.2—-4.3) (4.0-4.7) (3.3-3.4)¢° 
of lower condylar facet 0.05 0.05 


«Sample size reduced by 6. 
> Sample size reduced by 2. 
©‘ Sample size reduced by 1. 
4 Sample size reduced by 3. 


and skull; USNM 2653/4437, National Mu- 
seum of Natural History; from ‘‘near Fort 
Bliss, about 2 miles above El Paso, El Paso 
County, Tex.’’; obtained September 1861 
by S. W. Crawford. 

Distribution.—From southern California, 
Nevada, Utah, Colorado, Oklahoma, and 
western Arkansas, United States, southward 
to southern Baja California Sur, and east- 


ward to northern Sinaloa, and southern Za- 
catecas and Nuevo Leén, Mexico (Fig. 1). 

Diagnosis.—Notiosorex crawfordi can 
be distinguished from WN. villai by the roof 
of the glenoid fossa extending laterally be- 
yond the curve of the cranium (Fig. 4C) and 
by the greatest length of skull usually 
<=16.98 mm. It usually can be distinguished 
from N. evotis by rostral breadth <5.14 


VOLUME 113, NUMBER 1 


mm, width across M2—M2 <=5.1 mm, length 
of mandible =7.5 mm, height of coronoid 
process =4.4 mm, length of coronoid pro- 
cess-ventral point of upper condylar facet 
=3.9 mm, and length of coronoid process- 
ventral point of lower condylar facet =4.1 
mm. N. crawfordi usually can be distin- 
guished from WN. evotis by use of a combi- 
nation of height of coronoid process and 
cranial breadth (Fig. 5). Specimens of N. 
crawfordi can be separated from 100% of 
specimens of N. villai and N. evotis by ap- 
plication of the discriminant-function equa- 
tions presented in the Diagnoses sections of 
those species accounts. 

Description.—The skull of Notiosorex 
crawfordi is much like that of N. evotis, ex- 
cept that it is much smaller, the coronoid 
processes are slender compared to their 
height, and the paroccipital processes are 
small and lie against the exoccipitals (Fig. 
4C). The latter two characters are similar in 
form to N. villai. Although significant ge- 
netic differences (M. B. O’Neill, C. Porter, 
and R. J. Baker, pers. comm.) occur be- 
tween populations of N. crawfordi in Baja 
California and Texas, we found no identi- 
fiable morphological differences. 

The pelage is composed of multibanded 
hairs. In summer, hairs of the dorsal pelage 
have a narrow band of very dark grayish- 
brown (1OYR 3/2) distally and a wide band 
of dark gray (7.5YR N4/O) proximally. The 
hairs of the venter have equal-width bands 
of pinkish white (7.5YR 8/2) distally and 
very dark-gray (7.5YR N3/0) proximally. 
The winter pelage consists of hairs of the 
dorsum with a narrow band of very dark 
grayish-brown distally, a narrow band of 
pinkish white medially, and a wide band of 
dark gray (7.5YR N4/O) proximally. The 
hairs of the venter have a wide band of 
white (7.5YR N8/O) distally and a very nar- 
row band of gray (7.5YR N5/0) proximally 
creating a silvery wash effect over the ven- 
ter. The hairs of the tail are dark grayish- 
brown (1OYR 4/2). 

Measurements.—Individuals of Notioso- 


S13 


rex crawfordi are the smallest extant mem- 
bers of the genus (Table 1). 

Ecology.—wNotiosorex crawfordi is 
known to occur in habitats as diverse as de- 
sert shrub and yellow pine forest (Lange 
1959), pine-oak forest (Alvarez 1963), in 
grassland with oak chaparral and oak wood- 
land habitats nearby (Cunningham 1956), 
alkaline marsh (Stephens 1906), sandy flats 
(Fisher 1941, Yensen & Clark 1986), ‘‘arid 
grasslands with scattered catclaw, juniper 
and mesquite’? (Baker 1966:345), and near 
‘“‘a mesquite tree on a moist mud flat”’ 
(Armstrong & Jones 1971:751). It is known 
to occur at elevations at least as great as 
2618 m (Davis & Sidner 1989). 

From throughout the range of Crawford’s 
gray shrew, known mammal associates are 
members of the genera Cryptotis, Sorex, 
Scapanus, Sylvilagus, Lepus, Tamias, Sciu- 
rus, Spermophilus, Thomomys, Cratogeo- 
mys, Liomys, Dipodomys, Perognathus, 
Chaetodipus, Baiomys, Neotoma, Peromys- 
cus, Onychomys, Reithrodontomys, Sigmo- 
don, Microtus, Mus, Rattus, and Odoco- 
ileus, plus 14 species of bats (Chiroptera; 
Cunningham 1956, Anderson & Long 
1961, Coulombe & Banta 1964, Baker 
1966, Dalby & Baker 1967, Davis & Sidner 
1989). Also, Crawford’s gray shrew is well 
known for its association with woodrat 
(Neotoma) nests throughout its distribution 
(Armstrong & Jones 1972). 

Etymology.—The species epithet is a pat- 
ronymic to honor the collector of the type 
specimen, S. W. Crawford. 

Remarks.—Although trapping success 
for specimens of Notiosorex crawfordi is 
limited, remains thereof commonly occur in 
pellets regurgitated by barn owls (Tyto 
alba) and great horned owls (Bubo virgi- 
nianus) throughout its distribution in the 
United States and Mexico (Twente & Baker 
1951, Baker 1953, Baker & Alcorn 1953, 
Cunningham 1956, Anderson & Ogilvie 
1957, Lange & Mikita 1959, Bradshaw & 
Hayward 1960, Anderson & Long 1961, 
Glass & Halloran 1961, Schaldach 1966, 
Anderson 1972). 


314 


Jones et al. (1962:150—151), in a review 
of Notiosorex from Sinaloa, treated evotis 
and crawfordi as full species, stating that 
‘“‘evotis has a longer body and hind foot 
than crawfordi but a relatively (sometimes 
actually) shorter tail and ear, and a distinct- 


ly larger, heavier skull .... Notiosorex ev- 
otis differs cranially from Notiosorex craw- 
fordi as follows: larger . . . ; mesopterygoid 


fossa squared rather than broadly U-shaped 
anteriorly; rounded process on maxillary at 
posterior border of infraorbital canal well 
developed ...; occipital condyles smaller 
and, in lateral view, elevated above basal 
plane of skull; upper molars slightly more 
crowded in occlusal view.”’ Armstrong & 
Jones (1971:750), in their update on Sina- 
loan mammals, treated the two forms as 
subspecies stating that “‘the population in 
northern Sinaloa is intermediate between 
crawfordi and evotis’’ and that some of the 
cranial differences noted earlier now appear 
to be inconsistent. After examination of the 
263 specimens referrable to N. crawfordi 
and N. evotis included in this study, we 
found that most of the characters presented 
in Jones et al. (1962) are either inconsistent 
or strongly age-related, thus are of limited 
use in distinguishing crawfordi and evotis. 

A discussion of the natural history, fossil 
record, and reproduction of N. crawfordi 
can be found in Armstrong & Jones (1972) 
and Coulombe & Banta (1964). 

Specimens examined.—229, as follows. 

Mexico: Baja California: 9 mi N Catav- 
ina on Mexico HWY 1 (MVZ 159725); 10 
mi SE El Rosario (MVZ 159726); San Mar- 
tin Island, 300 yds. inland from Hassler’s 
Cave (MVZ 136207); San Quintin (CAS 
52); San Quintin, San Simon River (USNM 
139592-—139593); San Tomas (USNM 
137142); Santa Anita (USNM 74550, 
79088-79092, 146693, 146933-146934, 
146936, 147352, 147421). Chihuahua: 3.5 
mi ESE Los Lamentos (KU 76488); 2 mi 
W Minaca (KU 109475). Coahuila: 3 mi 
NW Cuatrociénegas (KU 51571-51572); 
Sabinas (USNM 277621). Durango: 2 km 
SE Atotonilco, 6680 ft (MSUM 13887- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


13888); 4.8 km SE Atotonilco, 6680 ft 
(MSUM_ 13889-13890); 7 mi NNE Bo- 
quilla, 6400 ft (MSUM_ 10260). Nuevo 
Leon: 3 mi SW Galeana, 5100 ft (MSUM 
11238). Sinaloa: On HWY 15, 0.25 mi S 
Sonora line (UMMZ 109403); 5 mi WNW 
El Carrizo (KU 105409); El Fuerte (KU 
75184). Sonora: 14.6 mi E [by road] Ma- 
zocahul (MVZ 148830); 4.1 mi NW [by 
road] Nacori Chico (MVZ 148831). Zaca- 
tecas: Plateado (USNM 90845). 

United States: Arkansas: Crawford Co.: 
Natural Dam (USNM 286549). Arizona: 
Apache Co.: 1 mi N Spigerville (UMMZ 
80236); Cochise Co.: 9.1 mi S Chiricahua 
(USNM 552386); Huachuca Mts. (MSB 
62141-62144, 62146-62148, 62150, 
62154-62155, 62158-62159, 62161-— 
62166, 62168;..62171-62173, 62173 
62177, 62179, 62183-62184, 62188- 
62189, 62210-62215, 62218-62219; 
62223—62238);..20 mi E.. Pearce, -Pinery 
Canyon, 6500 ft (UMMZ 64102); Coconino 
Co.: 10 mi SW Black Falls (USNM 
244129); Grand Canyon (USNM 250676); 
Greenlee Co.: Blue River (USNM 144533); 
Maricopa Co.: Phoenix South Mt. (CAS 
13928); Pima Co.: 36 mi S Tucson (USNM 
2725 15—272516); 40 mi S Tucson (USNM 
272844); Santa Cruz Co.: locality unknown 
(USNM 289955); Yuma Co.: Yuma 
(USNM 120357). California: San Bernar- 
dino Co.: San Bernardino (USNM 187011); 
Inyo Co.: Panamint Range Cottonwood 
Canyon (CAS 23228-23230); Saline Valley 
(CAS 21270); Saline Valley, Grapevine 
Canyon, 4036-5750 ft (CAS 23231-23232, 
21249-21269); Silver Canyon Rd., 4.3 mi 
E [by road] jct. with California HWY 6 
(MVZ 158116); Riverside Co.: Millard 
Canyon (CAS 23240); San Bernardino Co.: 
Cottonwood (CAS 23242); Deep Canyon 
(CAS 23238); Kingston Range (CAS 
21518); San Gorgonia, Cottonwood (CAS 
23233-23237, 23241); San Diego Co.: lo- 
cality unknown (USNM 62619); Lakeside, 
head of Wildcat Canyon, 2100 ft (KU 
92627); Escondido (MVZ 33582); 9 mi S 
Escondido (MVZ 33388); Santer Mts. 


VOLUME 113, NUMBER 1 


(USNM 62919). Colorado: Baca Co.: 14 mi 
N, 4 mi E Springfield (KU 116960); Fre- 
mont Co.: Phantom Canyon, Eightmile 
Creek (KU 125348—125367); Wet Mt. (KU 
125368—125379); Montezuma Co.: Mesa 
Verde National Park (KU 105109); Otero 
Co.: 3 mi NW Higbee (KU 51673). New 
Mexico: Cibola Co.: Juan Tofoya (USNM 
147966); Hidalgo Co.: locality unknown 
(KU 145266, MSB 46468); 7.5 mi W An- 
imas, Antelope Pass (KU 145258—145259); 
Cienega Ranch ruins (KU 144031); 30 mi 
E Douglas, Guadalupe Canyon, Peloncillo 
Mts. (KU 145260—-145265); San Luis 
Spring, Mexican Boundary (USNM 
38250); Lincoln Co.: Capitan Mts. (USNM 
127229); Shafer Ranch (UMMZ 114784); 
Otero Co.: 3 mi N Tularosa (UMMZ 
81380); Union Co.: Tollgate Canyon, 10 mi 
N Folson (MWSU 15900). Nevada: Nye 
Cor: 1 mi N, 5 mi E Grapevine, Peak Mt., 
5500 ft (MVZ 92391). Oklahoma: Cimar- 
ron Co.: 2 mi N Kenton (MWSU 15867— 
15868, 15895); 3 mi N Kenton (MWSU 
15902); 4 mi N Kenton (MWSU 15779); 
Comanche Co.: Wichita Mts. National 
Wildlife Refuge (USNM 271959); Harmon 
Co.: 4 mi S Hollis (OSU 5823). Texas: Ar- 
cher Co.: intersection US 82. and 277, 22 
mi SW Holliday (UMMZ 167208—167209); 
14 mi WNW Archer City (MWSU 8584, 
8586); Lake Kickapoo (MWSU 5614), NW 
side Lake Kickapoo (MWSU 7016, 11106); 
Bexar Co.: San Antonio (USNM 125708); 
Brewster Co.: Burro Mesa, 3500 ft (MVZ 
80281); Briscoe Co.: Tule Canyon (UMMZ 
67277); Cottle Co.: 8 mi ESE Paducah (KU 
64560); Dickens Co.: 1 mi E Dickens 
(MWSU 2543); El Paso Co.: near Fort 
Bliss, about 2 miles above El Paso (USNM 
2653/4437); Garza Co.: locality unknown 
(MMNH_ 12502-12503); 1 mi SE Post 
(PSM 13878); Hansford Co.: 10 mi S, 3 mi 
W Gruver (KU 119395); Howard Co.: Big 
Spring (UMMZ 80248); Jim Wells Co.:: 
near Alice (TCWC 53283); Knox Co.: 4 mi 
E Benjamin (MWSU 16023); Nueces Co.: 
Corpus Christi (USNM_ 120087). Locality 
unknown: (USNM 4437). 


315 


Conclusions 


As presently understood, the geographic 
distributions of these three taxa do not over- 
lap, although those of Notiosorex crawfordi 
and N. evotis are parapatric in northern Sin- 
aloa. We believe the lack of overlap in geo- 
graphic ranges is the result of these taxa 
being low in abundance wherever they oc- 
cur, use of trapping techniques inappropri- 
ate for collecting shrews during surveys of 
small mammals, and lack of collecting ef- 
fort for soricids over large segments of the 
distribution of Notiosorex in Mexico. 

In multivariate space, not only can 100% 
of N. villai, N. crawfordi, and N. evotis be 
separated (Fig. 3), but N. villai can be dis- 
tinguished from N. crawfordi and N. evotis 
by the qualitative character of the roof of 
the glenoid fossa not extending laterally 
from the cranium (Fig. 4). Also, N. craw- 
fordi and N. evotis can be distinguished by 
the qualitative character of the relative 
shape of the coronoid processes (Figs. 4B— 
C; Choate 1969:473, figs. 3b—c) and the 
quantitative relationship of height of coro- 
noid process and cranial breadth (Fig. 5). 

With recognition herein of three species 
in the genus Notiosorex, the Mexican mam- 
malian fauna now includes 28 species in the 
family Soricidae (Ramirez-Pulido et al. 
1996, Woodman & Timm 1999). In Mexi- 
co, the greatest diversity of shrews is in the 
Trans-Mexican Volcanic Belt and the Sierra 
Madre del Sur provinces. Our discovery of 
the distinctive new species Notiosorex villai 
in Tamaulipas, part of the Sierra Madre Ori- 
ental Province, an area not especially 
known for endemism (Fa & Morales 1993), 
and that all three species of Notiosorex oc- 
cur in Mexico, highlights the need for con- 
tinued collection of specimens and study of 
available museum specimens to better un- 
derstand the mammalian fauna of Mexico. 


Acknowledgments 


For the loan of specimens in their care, 
we thank the curators and collection man- 
agers of the California Academy of Scienc- 


eS 
oO 


> 
—~] 


ae 
iN 


oie) 


HEIGHT OF 
> 


CORONOID PROCESS (mm) 


ee 
Co 


ee 
on 


7.0 7.4 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


O@o oO 
o 6€6afa@O On 


O GDOOKIEED ODOGEDD 


@aoo ao 


OQOOCCOGED @ O OO 
00 0@m0 
O qaimq@a0D00 
0. 80: 3020 
O O 
O 


7.8 


@MO O 


8.2 8.6 9.0 


CRANIAL BREADTH (mm) 


Fier: 


Bivariate plot of height of coronoid process and cranial breadth illustrating almost complete separation 


of Notiosorex evotis (closed circles) and N. crawfordi (open circles). 


es (CAS); Michigan State University Mu- 
seum (MSUM); Midwestern University, 
Wichita Falls, Texas (MWSU); Mammal 
Division, National Museum of Natural His- 
tory (USNM); Collection of Vertebrates, 
Oklahoma State University (OSU); Texas 
Cooperative Wildlife Collection, Texas 
A&M University (TCWC); Museum of 
Vertebrate Zoology, University of Califor- 
nia at Berkeley (MVZ); Museum of Zool- 
ogy, University of Michigan (UMMZ); 
James Ford Bell Museum of Natural His- 
tory, University of Minnesota (MMNH); 
the Museum of Southwestern Biology, Uni- 
versity of New Mexico (MSB); and James 
R. Slater Museum of Natural History, Uni- 
versity of Puget Sound (PSM). We thank T. 


Holmes for preparation of problematic 
specimens, V. Sanchez-Cordero for trans- 
lation of our Abstract into Spanish, and N. 
A. Slade for statistical advice. EK Cervantes, 
R. S. Hoffmann, V. Sanchez-Cordero, and 
B. J. Verts read earlier drafts of this man- 
uscript. 


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Rodriguez Vela, H. 1999. Notiosorex crawfordi 
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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 
113(1):319-—333. 2000. 


Status of the name Odocoileus hemionus crooki 
(Mammalia: Cervidae) 


James R. Heffelfinger 


Arizona Game and Fish Department, 555 N. Greasewood Road, Tucson, Arizona 85745, U.S.A. 


Abstract.—The present name of the desert mule deer (Odocoileus hemionus 
crooki) is based on a specimen collected in southwestern New Mexico near 
the Mexican border. This specimen was originally described as a new species 
(Dorcelaphus crooki) of black-tailed deer, not as a mule deer, because many 
of its characteristics are intermediate between mule deer and white-tailed deer. 
In the same publication, Dorcelaphus hemionus eremicus is described from 
western Sonora, Mexico, as a new subspecies of desert mule deer. A number 
of mammalogists believed the type specimen of crooki to be a hybrid between 
desert mule deer and Coues white-tailed deer (O. virginianus couesi), while 
others hypothesized it represented extremes of normal variation in mule deer. 
I have reassessed the type specimen of Dorcelaphus crooki and reaffirm that 
it is a hybrid, invalidating the use of the crooki subspecies name. Consequently, 
the oldest available name for the desert mule deer is O. h. eremicus. 

Resumen.—E]1 nombre actual del venado bura del desierto (Odocoileus hem- 
ionus crooki) se basa en un ejemplar colectado en el suroeste de Nuevo Mexico 
cerca de la frontera con México. E. A. Mearns describi6 este eyjemplar como 
una nueva especie (Dorcelaphus crooki) de venado cola negra, no como un 
venado bura, debido a que muchos de sus caracteres son intermedios entre los 
del venado cola blanca y los del venado bura. En la misma publicaci6n, Mearns 
describio también a Dorcelaphus hemionus eremicus del occidente de Sonora, 
México, como una nueva subespecie de venado bura del desierto. Un gran 
numero de mastozodlogos ha considerado que el ejemplar tipo de crooki es un 
hibrido entre el venado bura del desierto y el venado cola blanca de Coues (O. 
virginianus Ccouesi), mientras que otros han hipotetizado que representaba ex- 
tremos de la variaci6n normal del venado bura. Después de reevaluar el ejem- 
plar tipo de Dorcelaphus crooki concluyo que es en realidad un hibrido. Por 
lo tanto, el nombre valido mas antiguo para el venado bura del desierto es O. 
h. eremicus. 


Desert mule deer (presently known as 
Odocoileus hemionus crooki Mearns, 1897) 
inhabit the southwestern United States from 
West Texas through southern New Mexico 
and southern Arizona, and southward into 
Sonora, Chihuahua, Coahuila, Zacatecas, 
and Durango, Mexico (Leopold 1959, Cow- 
an 1961, Wallmo 1981). In addition, a small 
herd has been translocated to Nuevo Le6én, 
Mexico (Morrison et al. 1992; Fig. 1). 
Since its original description, the subspe- 


cific name of this taxon has been conten- 
tious due to uncertainties regarding the hy- 
brid status of the type specimen. 


Historical Review 


Mearns (1897) described Dorcelaphus 
crooki as a new species of black-tailed deer 
because of its similarity to the Columbian 
black-tailed deer (O. h. columbianus). The 
type specimen (National Museum of Natu- 


320 
118° 
0 100 300 kilometers 
22° 
118° 
Big. 1. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


98° 


ow 

5 
<= 
8 
at 
fe) 


Durango a 


‘ 


* San Luis a 


ee “¢ Potosi 


Current distribution of desert mule deer in the southwestern United States and northern Mexico. Type 


localities identified for a) Dorcelaphus hemionus eremicus Mearns, 1897; b) Dorcelphus crooki Mearns, 1897; 


and c) Odocoileus hemionus canus Merriam, 1901. 


ral History [USNM] 20572/35752) was col- 
lected in 1892 by E. A. Mearns on the sum- 
mit of the Dog Mountains, Hidalgo County 
(formerly part of Grant Co.), New Mexico 
during the survey of the boundary between 
Mexico and the United States. No other 
deer with similar characteristics were col- 
lected at or near that locality. 

In the same publication Mearns (1897) 
described the ‘“‘Burro deer or desert mule 
deer,’ Dorcelaphus hemionus eremicus 
based on a male (USNM 63403, the type) 
collected in 1895 ‘by. W. J..McGee in the 
Sierra Seri, Sonora, Mexico. McGee did 
not keep the skull, but processed the hide 
as a deer skin rug. Mearns (1897) de- 
scribed the subspecies as pale gray in color 
with short pelage, a dark dorsal stripe, pal- 
er forehead markings, and wide, heavy ant- 


lers. This description was based on the 
skin of the male and two sets of antlers, 
each from different deer; one of which 
came from the Sonoyta Valley, Sonora 
(USNM 59910), the other from Black 
Butte, Baja California (USNM 60855). 
Pieces of skin trimmed from the hide as it 
was made into a rug comprised the only 
material representing the type specimen in 
the National Museum until 1902 when the 
rug was acquired from Anita McGee 
(Poole & Schantz 1942). 

In 1901, Merriam described Odocoileus 
hemionus canus based on a male (USNM 
99361) from Sierra en Medio, Chihuahua, 
Mexico. Merriam (1901) distinguished O. 
h. canus from the western subspecies O. h. 
eremicus solely on the basis of antler con- 
formation. The Sierra en Medio lies only 40 


VOLUME 113, NUMBER 1 


km southwest of the type locality for Dor- 
celaphus crooki in New Mexico. 

In his list of big game of North America, 
and replacing Dorcelaphus with the correct 
senior synonym QOdocoileus, Seton (1898: 
286) included O. h. eremicus as a subspe- 
cies of mule deer, but maintained the 
‘Crook black-tailed deer”’ as O. crooki. Ly- 
dekker (1915) also listed this animal as a 
black-tailed deer, O. columbianus crooki. 
Following Merriam’s (1901) description, O. 
h. canus was used for the desert mule deer 
in West Texas, Arizona, New Mexico, and 
northcentral Mexico (Seton 1909, Lantz 
1910, Lydekker 1915, Bailey 1931, Cowan 
1936, Cahalane 1939, Dalquest 1953). 

Several mammalogists believed that 
crooki was based on a hybrid between 
Coues white-tailed deer (O. virginianus 
couesi) and a desert mule deer but lacked 
known hybrids for comparison (Lydekker 
1898, Seton 1929, Bailey 1931, O’Conner 
1939). Goldman & Kellogg (1939) exam- 
ined the holotype of O. h. canus and other 
specimens from the Sierra en Medio, Chi- 
huahua, along with the holotype of Dorce- 
laphus crooki, and concluded that the type 
of crooki was an unusual specimen of mule 
deer and not a hybrid. Because the name 
crooki antedates canus, they adopted O. h. 
crooki as the correct name for desert mule 
deer in the north-central states of Mexico 
and adjacent Arizona, New Mexico, and 
Texas. Hoffmeister (1962) also re-examined 
the type of crooki and compared it with 
specimens of O. virginianus and O. hem- 
ionus. He interpreted the specimen as sim- 
ply a mule deer with some features that 
were intermediate with or shared by white- 
tailed deer. 

Based on the type specimen from Sierra 
Seri, the range of O. h. eremicus was des- 
ignated somewhat arbitrarily as western So- 
nora, southwestern Arizona, and extreme 
southeastern California (Mearns 1907:210). 
Hoffmeister (1962) listed O. h. eremicus as 
a synonym of O. h. crooki because he did 
not consider western Sonoran mule deer 
(burro deer) distinguishable from other pop- 


321 


ulations of desert mule deer. Only Cowan 
(1936, 1961) and Longhurst & Chattin 
(1941) attempted to quantify differences be- 
tween deer within the ranges of eremicus 
and crooki. Cowan’s (1936) interspecific 
cranial distinctions were based on only four 
eremicus Skulls; of the two additional er- 
emicus specimens from Mexico that he 
used to differentiate external characteristics, 
one was from Tiburon Island, Sonora, 
which Cowan (1961) later considered to be 
a different subspecies (O. h. sheldoni). 
Cowan (1961) based his differentiation of 
eremicus On measurements of only one 
male and one female specimen, which may 
have been previously described (Cowan 
1936, Longhurst & Chattin 1941). Long- 
hurst & Chattin (1941) added descriptions 
of pelage variations to differentiate eremi- 
cus, but they had only one crooki skin and 
three skulls for comparison. Cowan’s 
(1936:236) measurements of eremicus from 
southwestern Arizona and California are 
within the normal variation of crooki re- 
ported by Hoffmeister (1986). Hall (1981) 
continued to treat western Sonoran mule 
deer as a separate subspecies (O. h. eremi- 
cus), but provided no supporting informa- 
tion. Hoffmeister (1962) found mule deer 
from southern Arizona and northern Sonora 
within 80 km of the type locality of O. h. 
eremicus (Sierra Seri, Sonora) to be suffi- 
ciently similar to warrant treating eremicus 
as a synonym of crooki. Hoffmeister 
(1986), while remarking that he could not 
confirm that O. h. eremicus was a synonym 
of O. h. crooki, still implied that western 
Sonoran mule deer were not distinguishable 
from those farther east. The purpose of this 
study is to confirm hybrid status of the type 
specimen for O. h. crooki and clarify sub- 
specific nomenclature for desert mule deer. 


Material and Methods 


I re-examine the type specimen of O. h. 
crooki to compare and contrast its qualita- 
tive and quantitative characters with corre- 
sponding features of mule deer, white-tailed 


322 


deer, and their hybrids from southern Ari- 
zona, southern New Mexico, and adjacent 
Mexico. Published data from previous com- 
parisons are supplemented by measure- 
ments (in millimeters) from female O. h. 
crooki (n = 12), female O. v. couesi (n = 
17), the type specimen of Dorcelaphus 
crooki, and a known O. h. crooki X O. v. 
couesi F, hybrid. The type is an adult fe- 
male, thus I included only adult females 
(22 years old), as determined from tooth 
wear and replacement (Robinette et al. 
1957, Severinghaus 1949), for comparison. 
Cranial measurements (Table 1) include the 
six used by Hoffmeister (1962, 1986) in ad- 
dition to depth of lacrimal fossa (deter- 
mined as either shallow, deep, or “‘no de- 
cision” by Hoffmeister 1962). All cranial 
measurements were taken with a metric dial 
caliper and recorded to the nearest 0.1 mm. 
Values for paired measurements (e.g., 
length of right and left nasals) are averages. 
Elsewhere in this report, I use the term hy- 
brid to refer to only verified O. h. crooki X 
O. v. couesi F, hybrids, unless otherwise 
noted. 


Review of Characters 


Hoffmeister (1962) reviewed character- 
istics useful for distinguishing O. virgini- 
anus and O. hemionus in Arizona in his 
evaluation of the type of crooki. These in- 
cluded cranial and external measurements 
(including size of metatarsal gland) along 
with qualitative descriptions of the type of 
antler, lacrimal pit, color pattern of the tail, 
and color and position of the metatarsal 
gland. He relied heavily on size because de- 
sert mule deer are larger than Coues white- 
tailed deer. Nevertheless, Hoffmeister did 
not have adults of known hybrids available 
to evaluate their size characteristics when 
deciding on the taxonomic status of the type 
of crooki. Although acknowledging Ni- 
chol’s (1938) success in producing hybrids 
in captivity, Hoffmeister (1962:52) tended 
to discount the occurrence of hybrids in the 
wild. Today, however, data on dimensions 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


and other characteristics are available for 
mule deer X white-tailed deer hybrids, in- 
cluding both captive produced and geneti- 
cally confirmed, wild-taken animals. Char- 
acteristics of known hybrids produced in 
captivity provide a morphological basis for 
assessing suspected hybrids between mule 
and white-tailed deer encountered in the 
wild. The primary features used to distin- 
guish between desert mule deer and Coues 
white-tailed deer concern the length and 
color pattern of the tail; size, position, and 
color of hair tuft of the metatarsal gland, 
depth of the lacrimal pit, and dimensions of 
the body and cranium (Table 1). My eval- 
uation of the holotype of crooki based on 
these features follows. 

Metatarsal glands.—Metatarsal glands 
of desert mule deer are positioned high on 
the metatarsus, exceed 75 mm in length, 
and are circumscribed with brown hair (Ca- 
ton 1877, Hoffmeister 1986). Those of 
Coues white-tailed deer are positioned be- 
low the midpoint of the metatarsus, mea- 
sure 25 mm or less in length, and are 
rimmed by white hairs (Quay 1971, Hoff- 
meister 1986). Unlike either parent, all 
known F, hybrids have metatarsal glands 
that are intermediate in length, location, and 
appearance (Table 2). As affirmed by 
Mearns (1907) and Bailey (1931), the po- 
sition of the metatarsal gland in the holo- 
type of crooki is intermediate in compari- 
son to its location in white-tailed and mule 
deer (Fig. 2). It is nearly identical in loca- 
tion, length, and appearance to metatarsal 
glands of F, hybrids produced in captivity 
(Day 1980) and to wild mule X white-tailed 
deer hybrids whose status was confirmed 
genetically (Wishart 1980). 

The metatarsal glands of the type of 
crooki (right = 34 mm, left = 42 mm) are 
longer than those of white-tailed deer (25 
mm or less) and well below the range for 
desert mule deer (75-150 mm; Table 1). 
Day’s (1980) measurements of the metatar- 
sal glands of two adult hybrids born in cap- 
tivity were 50 mm for a female and 73 mm 
for a male. Wishart (1980) reported lengths 


VOLUME 113, NUMBER 1 


of glands from two wild-taken mule X 
white-tailed deer hybrids (status confirmed 
by electrophoresis) as 50 mm for a female 
and 62 mm for a male. Halloran & Kennedy 
(1949) and Lang (1957:14) gave lengths of 
metatarsal glands of adult female desert 
mule deer from southern New Mexico that 
averaged 2 to 3 times the length of the 
gland in the holotype of crooki. 

Anderson et al. (1964) included the co- 
rona of hair (circumglandular hair tuft) in 
their measurements of the metatarsal glands 
of 431 adult female mule deer from the 
Sacramento and Guadalupe mountains of 
southern New Mexico. Metatarsal gland 
lengths of females from the Sacramento 
Mountains (identified as O. h. hemionus) 
averaged 138 mm (range, 100—190); those 
of the Guadalupe Mountains (identified as 
O. h. crooki), 131 mm (range, 90-190). The 
shortest (90 mm) is considerably longer 
than the longest circumglandular hair-tuft 
measurement (70 mm) on the type of 
crooki. 

Mearns (1897, 1907) described the hairs 
surrounding the metatarsal gland of the type 
of crooki as “‘sooty at the base and white 
apically.”’ On examination, these hairs are 
“sooty at the base;’’ however, while pale, 
they are not white apically (Fig. 2c). Instead 
they are nearly the same pale color as the 
remainder of the leg, which Mearns (1897, 
1907) described as ‘“‘cream-buff, except 
where new clay colored hair is coming in 
on the anterior border.’’ Metatarsal glands 
of hybrids produced in captivity are either 
circumscribed with white hairs (G. I. Day, 
in litt.) or the hairs match the brown col- 
oration of their mule deer parent (J. C. 
Haigh, in litt.). 

Tail.—Mearns (1897:2) described the tail 
of crooki as “‘colored much as in D. col- 
umbianus, but has a longer terminal switch; 
upper side and extremity of tail all black, 
lower side white medially, and naked to- 
wards the base”’ (Fig. 3c). He gave its ver- 
tebral length as 195 mm, which is longer 
than that of a female hybrid (184 mm; G. 
I. Day, in litt.) and in the range of O. v. 


323 


couesi, but at or exceeding the upper limit 
for desert mule deer (Table 1). The color 
pattern of the tail of the type of crooki re- 
sembles the tails of captive-born hybrids, 
which Day (1980) described as “‘dark red- 
dish-brown or reddish-black above with 
white beneath and along the borders”’ (Fig. 
3d). Tails of some subspecies of mule deer 
(e.g., O. h. fuliginatus) commonly have a 
dark dorsal surface; however, this pattern is 
rare in southern Arizonan and New Mexi- 
can populations. Of 349 desert mule deer 
observed in southeastern Arizona during 
January 1998, no adults had dark tails re- 
sembling the type of crooki. However, in 
areas of sympatry with white-tailed deer, 
two fawns seen in the company of female 
mule deer each had a wide, dark tail stripe. 
I do not know if these were hybrids or pure 
mule deer fawns; yet hybrid fawns are typ- 
ically seen in the company of mule deer 
(Wishart 1980, Kay & Boe 1992), implying 
that the usual hybrid cross is between an 
aggressive white-tailed buck and a mule 
deer doe. 

Length of hind foot.—Hoffmeister (1962, 
1986) allowed that the length of hind foot 
(400 mm) of the crooki type is more char- 
acteristic of white-tailed deer. Based on the 
data at hand (Table 1), 400 mm is at the 
upper extreme of length of hind foot for 
Coues white-tailed deer and at the lower ex- 
treme for desert mule deer. G. I. Day’s (in 
litt.) measurement of the length of hind foot 
in a captive-born hybrid is 405 mm. 

Total length.—The total length of the 
type of crooki (1440 mm; Mearns 1897) 
and that of a female hybrid (1549 mm; G. 
I. Day, in litt.) is within the range for desert 
mule deer, but longer than normal for Coues 
white-tailed deer (Table 1). The ratio of tail 
length to total length for the type of crooki 
is 7.4X which is at the upper extreme for 
Coues white-tailed deer; however, the ratio 
in Day’s female hybrid is 8.4 and within 
the normal range for desert mule deer (Ta- 
ble 1). The range of ratios of tail length to 
total length in Coues white-tailed deer is 
5.6X-—7.5X, whereas the normal range of 


324 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Table 1.—Comparison of Coues white-tailed deer (Odocoileus virginianus couesi), desert mule deer (O. 
hemionus crooki), and an O. virginianus couesi X O. hemionus crooki F, hybrid, and the type specimen of 
Dorcelaphus crooki. All measurements (in millimeters) are from adult females two years of age or older. 
Characteristics of the type of Dorcelaphus crooki from Mearns (1907); cranial measurements from A. L. Gardner 
(in litt.). Measurements of known hybrid (UA 22358) supplemented by morphological information from G. I. 
Day (in litt.). Sample sizes in parentheses. 


Interpretation 
Character O. v. couesi F, hybrid Type of crooki O. h. crooki of crooki type 


Metatarsal gland: 


Location Below midpoint Intermediate Intermediate At or above mid- Hybrid 
of shank? point of shank? 
Length Usually <25? 50 34R, 42L™ 75-150? Hybrid 
14—23 (4)? 102-121 (4)° 
18 (1) X = 132° 
Length hair tuft 70 90-190 (431)! Hybrid 
Color hair tuft White Brown or white Pale brown Buff or Brown; Hybrid 
never white*® 
Tail: 
Length 215-260 (3)8 184 195 170—228 (5) Mule deer or 
Hybrid 
>1888 145-180 (8)? 
170-230 (9) 152-191 (5) 
165-229 (8)! 
127-185 (8) 
Dorsal color White border, dull Dark reddish- Black White with black Hybrid 
cinnamon? black? terminal brush" 
Gray, reddish- Like white- No white border, 
brown, grayish tailed deer‘ usually without 
brown or almost midband? 
black? 
Length of hindfoot 387-390 (3) 405 400 430—464 (5)* Hybrid 
<404° 380-490 (448)! 
X = 409 406-445 (5)° 
332—405 (18)4 406-475 (8)? 
Total length 1410-1450 (3) 1549 1440 1370-1570 (5)* Mule deer or 
1230-1420 (18) 1346-1549 (5)° Hybrid 
1397-1702 (8)! 
1430-1582 (8)! 
Ratio of tail to total 5.6 X-6.6x (3)) 8.4 X TA X 6.0 X-8.9 x (5)* Hybrid 
length 5.9 X-7.5 x (9) 8.2 <X-10.5 x™ 
6.4 X-7.3 x™ 8.1 X-10.2 x (5)<* 
= ST Xe X = 8.0 X (8) 
X = 10.1 x 
Depth of lacrimal 3.0-5.7 (4)" 53 5.9Ro63L 6.4-11.2 (11)" Hybrid 
fossa Shallow?" Deep*" 
Length of ear Gilg pas 188 190 190-193 (2) Mule deer 
145-170 (18) 184.2—203.2 (5) 
175-220 (8)? 
194.7—209.6 (8)! 
Basilar length 205-216 (4)" DID 239-265 (7)" Hybrid 
190.9-216.2 (13)° 229° 230-246 (12) 
Length of nasals 71.3-79.8 (5)" 72.0 79.1—95.6 (8)* 
60.4—76.7 (13)° 80.7° 79-95 (4)h Mule deer 
56.4—61.0 (3) 62.5-81.7 (12) 
Orbital width 55.3-59.7 (5)" 63 65.7—79.8 (10)" Hybrid 


54.5—56.0 (4) 64.2° 68.0—81.0 (12) 


VOLUME 113, NUMBER 1 


Table 1.—Continued. 


325 


Interpretation 


Character O. v. couesi F, hybrid Type of crooki O. h. crooki of crooki type 
31.5=62.2 (13)° 
51.4-60.8 (2)? 
Zygomatic width 87.8—97.7 (6)* 96.3 99.1—113.7 (9)* Mule deer 
90-97 (4) 102.1° 100-110 (13) 
89.4—100.4 (11)° 
96.6—100.7 (2)? 
Length of upper P-M 63.5—67.2 (5)" 70.5 72.1-82.6 (11)? Mule deer 
toothrow 61.1-69.6 (13)° 76.4° 80-89 (4) 
63.4—-66.8 (4)* 70.1—85.5 (12)? 
Length of lower P-M 72.5—76.2 (3)* 80.7 80.9—100.5 (10)" Mule deer 
toothrow 66.1—77.6 (11)° S72" 87-97 (4) 
72.1-—74.4 (2)? 82.4—97.4 (12) 
Usual topographic 1231-21549 above 1800 <1400' White-tailed 
elevation (m) deer 


“Hoffmeister 1986. 

> Quay 1971. 

© Halloran & Kennedy 1949. 
#<a. f. Day, in litt. 
¢Lang 1959. 

f Anderson et al. 1964. 

® Caton 1877. 

Mearns 1907. 

‘Cowan 1961. 
Pruett, im litt. 

K Nichols 1938. 

' Bailey 1931. 

™ Hoffmeister 1962. 

" This study. 

° A. L. Gardner, in litt. 

P Krausman et al. 1978. 

4 Anthony & Smith 1977. 
"Krausman 1978. 


ratios in desert mule deer is 7.7X—10.5X 
(Mearns 1907, Hoffmeister 1962, G. I. Day, 
in litt.). An exception is a desert mule deer 
from west of El Paso, Texas (Mearns 1907), 
with an unusually long tail (228 mm) and 
short total length (1307 mm) yielding a ra- 
tio of 6.0X. 

Length of ear.—Total length of ear for 
the type specimen of O. h. crooki (190 mm) 
is within the normal range for desert mule 
deer and is longer than that of a white-tailed 
deer (Table 1). The length of ear for two 
adult F,; hybrids (O. h. crooki X O. v. coue- 
si) was 188 mm (female), and 209 mm 
(male), also within the normal range for 
mule deer (G. I. Day, in litt.). 

Lacrimal fossa.—Depth of the lacrimal 


fossa is diagnostic; it is deeper (6.4—11.2 
mm) and larger in desert mule deer than in 
Coues white-tailed deer (3.0—5.7 mm). 
Hoffmeister’s (1962:49) “tno decision’’ on 
the depth of the fossa in the type specimen 
of crooki was because both fossae are fe- 
nestrate. However, the floor of each fossa is 
clearly evident and the depth of the right 
fossa measures 5.9 and the left fossa, 6.3. 
These measurements are intermediate be- 
tween the ranges of lacrimal-fossa depths of 
the two species and confirms Wishart’s 
(1980) observation for known mule X 
white-tailed deer hybrids. Depth of lacrimal 
fossa in another hybrid doe measures 5.3 
(Table 1), which is near the upper range of 
that for white-tailed deer. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


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VOLUME 113, NUMBER | 327 


Fig. 2. Metatarsal glands of: a) desert mule deer (Odocoileus hemionus crooki [=eremicus]); b) Coues white- 
tailed deer (O. virginianus couesi); c) holotype of (Dorcelaphus crooki, USNM 20572/35752); d) known F, 
hybrid (O. h. crooki X O. v. couesi). Photographs by J. R. Heffelfinger (a & b), L. M. Snyder (c), and G. I. 
Day (d). 


Fig. 3. Tails of: a) desert mule deer (Odocoileus hemionus crooki [=eremicus]); b) Coues white-tailed deer 
(O. virginianus couesi); c) holotype of (Dorcelaphus crooki, USNM 20572/35752); d) known F, hybrid (O. h. 
crooki X O. v. couesi). Photographs by J. R. Heffelfinger (a & b), L. M. Snyder (c), and G. I. Day (d). 


328 


Cranial measurements.—Hoffmeister’s 
(1962, 1986) decision that the type of 
crooki was a mule deer pivoted on the use 
of skull measurements. Adult desert mule 
deer in the region of the type locality of 
crooki are much larger and may weigh 
twice as much as Coues white-tailed deer 
resulting in some cranial dimensions being 
interspecifically diagnostic. Hoffmeister 
(1962:48) provided generalized interspecif- 
ic limits, but not actual ranges of the six 
measurements (basilar length of Hensel, 
length of nasals, orbital width, zygomatic 
breadth, and length of both upper and lower 
molariform toothrows) he used in separat- 
ing female mule and white-tailed deer. He 
said all six measurements of the type spec- 
imen for crooki were within the lower range 
of these measurements for desert mule deer. 
However, I found that orbital width and 
basilar length were below the correspond- 
ing ranges for this measurement for mule 
deer; the remaining four measurements 
were within the normal range. The discrep- 
ancy between Hoffmeister’s (1962) mea- 
surement of the nasals of the type of crooki 
and the longer measurement in Table 1 
credited to A. L. Gardner results from Hoff- 
meister’s measurement equaling the shortest 
distance from the frontal-nasal suture to the 
proximal margin of its anterior border be- 
tween medial and lateral anterior projec- 
tions. Gardner’s measurement is the average 
of the greatest distance between anterior 
and posterior points of right and left nasals; 
the right nasal measures 80.0 mm and the 
left, 81.5 mm. Hoffmeister’s (1986) princi- 
pal components analysis using 11 cranial 
measurements grouped measurements of 
the type of crooki with those of mule deer. 
Cranial measurements of an adult captive- 
born hybrid doe are either intermediate or 
within the normal range for mule deer (Ta- 
ble 1). 

Cowan (1962) reported four cranial di- 
mensions (width of nasals, interorbital 
width, palatal width, and postpalatal width) 
used to differentiate mule deer and white- 
tailed deer in Alberta, Canada. Three of 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


these four measurements from a male mule 
x white-tailed deer hybrid (as determined 
from metatarsal gland characteristics) were 
within the normal range for mule deer. Wis- 
hart (1980) used five cranial measurements 
in an analysis of a male and a female wild- 
taken hybrid (both confirmed by electro- 
phoresis) in Alberta. All measurements fell 
within the normal range for mule deer with 
the exception of post-palatal width of the 
female, which was within the range for 
white-tailed deer. 

Type locality.—Mearns (1897, 1907:190) 
collected the type of Dorcelaphus crooki on 
9 June 1892, on the summit of ‘Emory 
Peak of the Dog Mountains, where I noted 
its range as from 1500 to 1868 meters.”’ 
Mearns (1907:87) was camped at “Dog 
Spring .. . [which] is about 2 kilometers... 
north of [Boundary] Monument No. 55... 
near the south extremity of the Dog Moun- 
tains, a rugged range of which Emory Peak, 
having an altitude of 1868 meters (6129 
feet), is the highest.’”’ Dog Spring (31°21'N, 
108°19'W) appears on several old maps as 
“Ojo del Perro.’’ The Dog Mountains are 
known today as the Alamo Hueco Moun- 
tains. Apparently, the name Emory Peak 
does not appear on any topographic map of 
the area produced from 1881 through 1983 
(C. Kollen, pers. comm.). A Department of 
the Interior, General Land Office map dated 
1903 shows an “Emory Sp.”’ northwest of 
Ojo del Perro, in the general vicinity of 
Pierce Peak (31°27'N, 108°20’W). As 
Pierce Peak is drained to the north by Emo- 
ry Canyon and 11 km northwest of Dog 
Spring, it is most likely the same peak iden- 
tified by Mearns as Emory Peak. The ele- 
vation of Pierce Peak is given today as 
1877 m; however, a map of the area dated 
1942 has the elevation as 6149 ft (1874 m). 
Mearns’ (1907) elevation of 1868 m for 
Emory Peak is nearly equivalent; further- 
more, Pierce Peak is covered with alligator 
juniper (Juniperus deppeana), which 
matches his description of the area. 

The elevation at which the type of crooki 
was collected, presumed to be above 1800 


VOLUME 113, NUMBER 1 


m, is above the normal upper elevational 
limit for desert mule deer (1400; Krausman 
1978, McCulloch 1972). Coues white-tailed 
deer, however, often occur in highest den- 
sities between 1230 and 2150 m elevation 
(Anthony & Smith 1977). Both species are 
present in the Alamo Hueco Mountains, but 
white-tailed deer are less common today 
than they were earlier in this century (Bai- 
ley 1931; Raught 1967; A. Hurt, pers. 
comm.). 


Discussion 


In the original description of Dorcela- 
phus crooki, Mearns (1897:3) said ‘“The 
skull has very nearly the same conforma- 
tion as that of D. columbianus [black-tailed 
deer], the lacrimal fossa being deeper than 
in the Virginia deer, but shallower than in 
the mule deer. The same intermediate con- 
dition obtains with respect to the vomer, in 
the relationships of the nasal and premax- 
illary bones, in the form and size of the 
teeth; and, in short, the whole animal ap- 
pears to be a compromise between the char- 
acteristics of the white-tailed and mule 
deer’ 

In 1907, Mearns referred to this taxon as 
Odocoileus crooki and explained naming 
the deer for General George Crook. Mearns 
also received a specimen of a 2-year-old 
buck shot in the vicinity of Bill Williams 
Mountain, Arizona, in 1884 by a member 
of General Crook’s hunting party. Mearns 
believed this specimen also represented his 
new species of black-tailed deer. I have not 
examined the male to verify its hybrid sta- 
tus because its identity has no bearing on 
the status of the name crooki because it is 
not the type. Mearns (1907:187) reported 
the length of metatarsal gland on this spec- 
imen as 13 mm, which is typical of a white- 
tailed deer (Table 1). 

For several decades following Mearns’ 
(1897) description of Dorcelaphus crooki, 
several authorities suggested that the type 
was a hybrid (Lydekker 1898, Seton 1929, 
Bailey 1931, O’Conner 1939) and Merri- 


329 


am’s name O. h. canus was used for the 
desert mule deer. Goldman & Kellogg 
(1939), having noted that a mule deer (O. 
h. peninsulae) from lower Baja California, 
Mexico, had a tail color pattern similar to 
that of the type of crooki and reasoning that 
their animal could not be a hybrid because 
white-tailed deer were not in Baja Califor- 
nia, concluded that the type of crooki was 
an abnormal specimen of mule deer. Hoff- 
meister (1962, 1986) acknowledged that the 
type of crooki was intermediate in some 
features, but believed the preponderance of 
evidence, particularly of size, supported his 
assessment that the animal was simply an 
abnormal mule deer. He also said that the 
metatarsal gland may be small or indistinct 
in some populations of mule deer else- 
where, citing Hershkovitz’s (1958:538) ob- 
servation that two mule deer from lower 
Baja California, Mexico, had poorly-devel- 
oped glandular tissue underlying well-de- 
fined metatarsal hair tufts. Nevertheless, the 
type specimen of crooki has _ shortened 
metatarsal glands and circumglandular tufts 
unlike those of any known mule deer, but 
consistent in size, form, and position with 
those of known hybrids (Tables 1 & 2). 
Hoffmeister (1962:52), in his statement 
‘The few ‘hybrids’ that I have been able to 
track down either prove to be clearly O. 
hemionus or O. virginianus,’’ implied that 
hybridization between these species proba- 
bly did not occur in the wild. 

White-tailed deer < mule deer hybrids 
are known to have been produced in cap- 
tivity as early as 1865 (Gray 1972). Other 
examples of hybridization have been doc- 
umented at captive facilities in Arizona (Ni- 
chol 1938, Day 1980), Colorado (Spraker 
et al. 1997), Illinois (Caton 1877), Texas 
(Derr 1990), Wyoming (Guiroy et al. 1991, 
E. S. Williams, in litt.), and Alberta, Canada 
(Lingle 1992, W. D. Wishart, in litt.). 
Whitehead (1972) reported white-tailed x 
black-tailed deer hybrids produced in cap- 
tivity in Tennessee. 

Hybridization between white-tailed deer 
and mule deer has been documented genet- 


330 


ically or on the basis of metatarsal gland 
morphology in the wild in Arizona (Day 
1964, P A. Dratch, in litt., J. A. Holcomb, 
in litt.), Montana (Cronin 1991), Texas 
(Carr et al. 1986, Stubblefield et al. 1986, 
Derr 1990, Ballinger et al. 1992), Washing- 
ton (Gavin & May 1988), Wyoming (Kay 
& Boe 1992), and in Alberta (Wishart 
1980) and British Columbia (Cowan 1962), 
Canada. White-tailed deer and mule deer 
are sympatric in the vicinity of the type lo- 
cality of crooki (Hoffmeister 1962; A. Hurt, 
pers. comm.), and hybrids are documented 
from adjacent areas in Arizona and Texas. 

The only genetic tests that will differen- 
tiate white-tailed deer and mule deer are 
electrophoresis of albumin (Scribner et al. 
1984) and erythrocyte acid phosphatase (P. 
A. Dratch, in litt.), and isoelectric focusing 
of muscle esterase (Oates et al. 1979). All 
of these analyses require fresh or frozen 
samples. No molecular markers are cur- 
rently known that will differentiate these 
species from skin samples from museum 
specimens (P. A. Dratch, in litt.). 

Known hybrids are large and some body 
(ength of ear and total length) and cranial 
measurements (zygomatic breadth, length 
of nasals, and upper and lower molariform 
toothrows) are within the normal range for 
desert mule deer (G. I. Day, in litt.). The 
phenomenon of heterosis (hybrid vigor) in 
F, hybrids is well known in cervids (Krzy- 
winski 1993, Tate et al. 1997). The deer 
farming industry has capitalized on heter- 
osis by crossing the phenotypically diverse, 
but presumed conspecific, red deer and wa- 
piti (Cervus elaphus). F, hybrids show 
higher and faster weight gains making them 
more profitable than either purebred paren- 
tal stock (Pearse 1993). Variability in over- 
all size of F, phenotypes means that most 
cranial and body measurements are poor 
choices for evaluating hybrid status in deer 
(Cowan 1962; Day 1980, in litt.; Wishart 
1980). 

Any mule deer may have an abnormal 
tail, metatarsal gland, lacrimal fossa, cranial 
measurements, or length of hind foot. How- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


ever, many characteristics from the type 
specimen for crooki fall outside the normal 
range of variation for mule deer, and are 
consistent with those of known white-tailed 
x mule deer hybrids. The overwhelming 
concordance of morphologic evidence in 
this comprehensive analysis reveals the ho- 
lotype of Dorcelaphus crooki as a hybrid 
between Coues white-tailed deer and desert 
mule deer. This, then, has serious repercus- 
sions for the current scientific name for de- 
sert mule deer. A scientific name based on 
a type specimen later found to be a hybrid 
is invalid and can not be used for either of 
the parental species even if it has priority 
over all other available names (ICZN 1985: 
Art. 23[{h]). This reanalysis clarifies the 
long-standing confusion regarding the sta- 
tus of this contentious type specimen and 
therefore the correct scientific name for this 
taxon. The oldest available name for desert 
mule deer (formerly known as O. h. crooki 
and O. h. canus) is Odocoileus hemionus 
eremicus Mearns, 1897; an abbreviated 
synonymy follows: 


Odocoileus hemionus eremicus (Mearns) 


Dorcelaphus crooki Mearns, 1897:2; un- 
available name because it is based on a 
hybrid. 

Dorcelaphus hemionus eremicus Mearns, 
1897:4; type locality “Sierra Seri, near 
the Gulf of California, in the most arid 
portion of Sonora, Mexico.” 

Odocoileus hemionus canus Merriam, 
1901:560; type locality “‘Sierra en Media, 
Chihuahua, Mexico.”’ 

Odocoileus hemionus crooki: Goldman & 
Kellogg, 1939:507; name combination. 


Acknowledgments 


I am very much indebted to A. L. Gard- 
ner, USGS Patuxent Wildlife Research Cen- 
ter, Biological Survey Unit, National Mu- 
seum of Natural History (formerly called 
the United States National Museum), for 
measurements of specimens including the 


VOLUME 113, NUMBER 1 


type of Dorcelaphus crooki and for tech- 
nical advice and significant assistance with 
the manuscript. L. M. Snyder of the Na- 
tional Museum of Natural History provided 
photographs of the type specimen of D. 
crooki. Thanks are extended to V. C. 
Bleich, D. E. Brown, M. A. Cronin, P. R. 
Krausman, R. M. Lee, and J. R. Purdue for 
their review of an earlier draft of this man- 
uscript. EF J. Abarca, D. E. Brown, S. Gal- 
lina, R. S. Henry, D. Humphreys, A. Mar- 
tinez, E. Mellink, B. L. Morrison, J. Rom- 
ero, A. Santos, E Tapia, M. Weber, and D. 
L. Weybright provided important informa- 
tion regarding the current range of desert 
mule deer. Arizona Game and Fish Depart- 
ment Wildlife Managers collected data on 
tail color pattern during routine deer sur- 
veys. J. C. deVos provided assistance with 
photographs and M. Alderson generated the 
map of the distribution of desert mule deer. 
N. B. Carmony provided insightful com- 
ments regarding subspeciation. I thank D. 
E. Brown, P K. Devers, Y. Petryszyn, M. 
T. Pruss, and especially R. S. White for al- 
lowing access to specimens and for osteo- 
logical assistance. G. I. Day, P. R. Kraus- 
man... Y. McCulloch, and J. C. Truett 
shared morphological data from their earlier 
work. A. Fisher, A. Hurt, C. Kollen, C. R. 
LeQuiev, E. M. Rominger, and C. G. 
Schmitt provided details on the Dog/Alamo 
Hueco mountains of southwestern New 
Mexico. Thanks are also extended to E. 
Mellink for providing the Spanish version 
of the abstract. Production costs for this 
manuscript were paid by the Federal Aid in 
Wildlife Restoration Act, W-53-M and W- 
78-R, of the Arizona Game and Fish De- 
partment. 


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Journal of Wildlife Management 21:134-153. 

Scribner, K. T., R. J. Warren, & S. L. Beasom. 1984. 
Electrophoretic identification of white-tailed 


VOLUME 113, NUMBER 1 


and mule deer feces: a preliminary assess- 
ment.—Journal of Wildlife Management 48: 
656-658. 

Seton, E. T. 1898. A list of big game of North Amer- 
ica.—Forest and Stream (October) pp. 285—286. 

. 1909. Life histories of northern animals. Chas. 

Scribner’s Sons, New York, 1:1—673. 

. 1929. Lives of game animals. C. T. Branford 
Company, Boston, 412 pp. 

Severinghaus, C. W. 1949. Tooth development and 
wear as criteria of age in white-tailed deer.— 
Journal of Wildlife Management 13:195—216. 

Spraker, T. R., et al. 1997. Spongiform encephalopathy 
in free-ranging mule deer (Odocoileus hemion- 
us), White-tailed deer (Odocoileus virginianus) 
and Rocky Mountain elk (Cervus elaphus nel- 
soni) in northcentral Colorado.—Journal of 
Wildlife Diseases 33:1-—6. 

Stubblefield, S. S., R. J. Warren, & B. R. Murphy. 
1986. Hybridization of free-ranging white-tailed 
and mule deer in Texas.—Journal of Wildlife 
Management 50:688—690. 

aete, mi. ., G. J. Goosen, G. Patene, A. J. Pearse, K. 
M. McEwan, & P. FE Fennessy. 1997. Genetic 
analysis of Pere David’s X red deer interspecies 
hybrids.—Journal of Heredity 88:361—365. 

Wallmo, O. C. 1981. Distribution and habits. Pp. 1-26 
in O. C. Wallmo, ed., Mule and black-tailed 
deer of North America. Wildlife Management 
Institute and University of Nebraska Press, Lin- 
coln, 605 pp. 

Whitehead, Jr, C. J. 1972. A preliminary report on 
white-tailed and black-tailed deer crossbreeding 
studies in Tennessee.—Proceedings of the An- 
nual Conference of the Southeastern Associa- 
tion of Game and Fish Commissions 25:65—69. 

Wishart, W. D. 1980. Hybrids of white-tailed and mule 
deer in Alberta.—Journal of Mammalogy 61: 
716-720. 


Appendix | 
Specimens Examined 


With the exception of the two type specimens of 
mule deer (both males) listed below, specimens used 
in this analysis, including the type of Dorcelaphus 
crooki, are all adult females (=2 years), and deposited 
in the following institutions: Arizona State University 
(ASU); Arizona Game and Fish Department, Tucson 
(AGFD); University of Arizona (UA); National Mu- 
seum of Natural History (USNM). A. L. Gardner mea- 
sured the USNM specimens; I measured all others. 

Odocoileus hemionus crooki [=eremicus| (14).— 
United States. Arizona: Yavapai Co., 8 km NE Horse- 
shoe Dam (ASU 643); Maricopa Co., 3 km E Horse- 
shoe Dam (ASU 637); Pinal Co., N side of Canyon 
Lake (ASU 643), Picacho Mountains (UA 24418-— 
24420, 24429, 24430, 24436, 24478, 25299, 25308). 
Mexico. Chihuahua: Sierra en Medio (USNM 99361, 
type of O. h. canus). Sonora: Sierra Seri (USNM 
63403, type of O. h. eremicus). 

Odocoileus virginianus couesi (17).—United States. 
Arizona: Cochise Co., Chiricahua Mountains (UA 
20340, 20346); Graham Co., Blue River (USNM 
32115); Pima Co., Santa Rita Mountains (AGFD 10, 
UA 23304), Baboquivari Mountains (AGFD 53201); 
Santa Cruz Co., Santa Rita Mountains (USNM 
202931). New Mexico: Catron Co., Mogollon Moun- 
tains (USNM 148574); Grant Co., 32 km W Silver 
City (USNM 286685), head of Mimbres River (USNM 
147476); Hidalgo Co., near Cloverdale (USNM 
35748). Mexico. Chihuahua: Colonia Garcia (USNM 
99347, 99350); Sonora: San Luis Mountains (USNM 
36320); E side of San Luis Mountains (USNM 35751, 
37085); Pozo de Luis (USNM 59229). 

Odocoileus hemionus crooki [=eremicus] X O. vir- 
ginianus couesi hybrids (2).—Arizona: Pima Co., Uni- 
versity of Arizona Captive Facilities (UA 22358). New 
Mexico: Hidalgo Co., Summit of the Dog Mountains 
(USNM 20572/35752, type of Dorcelaphus crooki). 


23) hint 


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ie he" PRET (arth Ree 
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bh Ao ye wo Ay epee Bp Salih 4 4Ondas fA aE) 
i i vio. 4h teen at ATES vo lpti H wViket 4 
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pj CAE tee i> pp einai ied: Lal (kee 
ol te A OA) ee 4c OP earch of ee 

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a) (10D owe etm A ») fore 7h Worl) Fate 
rol dat ite A Teverlduet omeeee et esiidd> iim 


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fire Bebe 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 
113(1):335-—336. 2000. 


INTERNATIONAL COMMISSION ON ZOOLOGICAL NOMENCLATURE 


Applications published in the Bulletin of Zoological Nomenclature 


The following Applications were published on 30 September 1999 in Vol. 56, 
Part 3 of the Bulletin of Zoological Nomenclature. Comment or advice on any of 
these applications is invited for publication in the Bulletin and should be sent to the 
Executive Secretary (I.C.Z.N.), % The Natural History Museum, Cromwell Road, 
London SW7 5BD, U.K. (e-mail: iczn@nhm.ac.uk). 


Case No. 


3116 Gnomulus Thorell, 1890 (Arachnida, Opiliones): proposed designation of G. 
sumatranus Thorell, 1891 as the type species. 

3089 Leucocytozoon (Protista, Haemosporida): proposed adoption of Berestneff, 
1904 as the author and of Leukocytozoen danilewskyi Ziemann, 1898 
as the type species. 

Drosophila rufifrons Loew, 1873 and D. lebanonensis Wheeler, 1949 (cur- 
rently Scaptodrosophila rufifrons and S. lebanonensis; Insecta, Dip- 
tera): proposed conservation of the specific names by the designa- 
tion of a neotype for D. rufifrons. 

Vespertilio pipistrellus Schreber, 1774 and V. pygmaeus Leach, 1825 (cur- 
rently Pipistrellus pipistrellus and P. pygmaeus; Mammalia, Chi- 
roptera): proposed designation of neotypes. 

Tanaecia coelebs Corbet, 1941 (Insecta, Lepidoptera): proposed conservation 
of the specific name. 

Diastylis Say, 1818 (Crustacea, Cumacea): proposed designation of Cuma 
rathkii Krgyer, 1841 as the type species. 

Hybognathus stramineus Cope, 1865 (currently Notropis strmineus; Ostei- 
chthyes, Cypriniformes): proposed conservation of the specific 
name 

Strongylus tetracanthus Mehlis, 1831 (currently Cyathostomum tetracan- 
thum) and C. catinatum Looss, 1900 (Nematoda): proposed con- 
servation of usage by the designation of a neotype for C. fetra- 
canthum 

Musca geniculata De Geer, 1776 and Stomoxys cristata Fabricius, 1805 (cur- 
rently Siphona geniculata and Siphona cristata; Insecta, Diptera): 
proposed conservation of usage of the specific names by the re- 
placement of the lectotype of M. geniculata by a neotype 

Ichthyosaurus cornalianus Bassani, 1886 (currently Mixosaurus cornalianus; 
Reptilia, Ichthyosauria): proposed designation of a neotype 

Mystacina Gray, 1843, Chalinolobus Peters, 1866, M. tuberculata Gray, 1843 
and Vespertilio tuberculatus J. R. Forster, 1844 (currently C. tub- 
erculatus) (Mammalia, Chiroptera): proposed conservation of usage 
of the names 


336 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Holochilus Brandt, 1835, Proechimys J. A. Allen, 1899 and Trinomys Thom- 
as, 1921 (Mammalia, Rodentia): proposed conservation by the des- 
ignation of H. sciureus Wagner, 1842 as the type species of Holo- 


chilus 

Cervus gouazoubira Fischer, 1814 (currently Mazama gouazoubira; Mam- 
malia, Artiodactyla): proposed conservation as the correct original 
spelling 


VOLUME 113, NUMBER 1 aor 


Opinions published in the Bulletin of Zoological Nomenclature 


The following Opinions were published on 30 September 1999 in Vol. 56, Part 3 
of the Bulletin of Zoological Nomenclature. Copies of these Opinions can be ob- 
tained free of charge from the Executive Secretary, I.C.Z.N., % The Natural History 
Museum, Cromwell Road, London SW7 5BD, U.K. (e-mail: iczn@nhm.ac.uk). 


Opinion No. 


1930 Osilinus Philippi, 1847 and Austrocochlea Fischer, 1885 (Mollusca, Gastro- 
poda): conserved by the designation of Trochus turbinatus Born, 
1778 as the type species of Osilinus. 

1931 Campeloma Refinesque, 1819 (Mollusca, Gastropoda): conserved. 

1932 Holospira Martens, 1860 (Mollusca, Gastropoda): Cylindrella goldfussi Men- 
ke, 1847 designated as the type species. 

1933 Androctonus caucasicus Nordmann, 1840 (currently Mesobuthus caucasicus; 
Arachnida, Scorpiones): specific name conserved. 

1934 Paruroctonus Werner, 1934 (Arachnida, Scorpiones): conserved. 

1935 Cicada clavicornis Fabricius, 1794 (currently Asiraca clavicornis; Insecta, 
Homoptera): specific name conserved. 

1936 Thamnotettix nigropictus Stal, 1870 (currently Nephotettix nigropictus; In- 
secta, Homoptera): specific name conserved. 

1937 Corisa propinqua Fieber, 1860 (currently Glaenocorisa propinqua; Insecta, 
Heteroptera): specific name conserved. 

1938 Musca rosae Fabricius, 1794 (currently Psila or Chamaepsila rosae; Insecta, 
Diptera): specific name conserved. 

1939 Trigonocephalus pulcher Peters, 1862 (currently Bothrops pulcher, Bothrie- 
chis pulcher or Bothriopsis pulchra; Reptilia, Serpentes): defined by 
the holotype, and not a neotype; Bothrops campbelli Freire Lascano, 
1991: specific name placed on the Official List. 

Hoplocephalus vestigiatus De Vis, 1884 (Reptilia, Serpentes): specific name 
placed on the Official List. 

Australopithecus afarensis Johanson, 1978 (Mammalia, Primates): specific 
name conserved. 


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qyailt isurmht ot ant ee 


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~~ 


crue) onli} axa Jotoe palissowms hos Fest aati bivatingt 
oss. voy isa oe jo robianiginely sul “oe Lev RG t a oy 
nue Wo sepa eyes cid) agi ore af 

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gine ageh MM 2s Danegash CRT ssa ae 
2D’) BMG ME, wie tica) shal Ai DIV, 9V aio Sains 
bovayies oon ofioggs 3 <anonpood  Abiniam re, 


p ey yee ‘(eammber32 ghlnitaes A £0] sone . moe oe 


iia pertl Savio sorn\s aoa Yinoraiey BOVE aoliormdas “nasi a 


MK. Sop eries. gawt othigade » On a ae 

oft: an 1 Rip Toe, vi scorns at VA. NOE 3 

. ieeY SEY Dede ge uae hie 

union. sonynieet ativan ml Vitis O08 ti ioelei es nie tt 

lbuvdags sagiel, githada acaatiys web. a 

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Ao Gs eh oft oge CSE 

‘Or the Re ee mor, Yilow eo) LOS) Aree vanity ay 

Co boittals. | werryiee pili ad scossl pig eeeieiioe, uo te 1 a 

silky a eyo, ‘eragh Ades, bee oy rola ott > a 
wa ae ie AKO ‘ott ete aperm aaa Stlinaga 210 a = 

SHAM Loa Stengel wciliim he Hn cele : f 

| sett ne bagabg 8 2 eS 


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yi 


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Front cover—from this issue, p. 44. 


CONTENTS 


Austin Beatty Williams (17 October 1919—27 October 1999). Biographical summary 
Rafael Lemaitre and Bruce B. Collette 
A new species of mud shrimp, Upogebia cortesi, from Pacific Costa Rica (Decapoda: Thalassinidea: 


Upogebiidae) Austin B. Williams and Rita Vargas 
Periclimenes murcielagensis, a new species of shrimp (Crustacea: Decapoda: Palaemonidae) living on 
black coral from the Pacific coast of Costa Rica Rita Vargas 
A new squat lobster of the genus Munidopsis Whiteaves, 1874 (Crustacea: Decapoda: Galatheidae) 
from Taiwan Ming-Feng Wu and Tin- Yam Chan 
A new freshwater crab of the genus Geothelphusa Stimpson, 1858 (Crustacea: Decapoda: Brachyura: 
Potamidae) from Yakushima Island, southern Kyushu, Japan Hiroshi Suzuki and Tomokazu Okano 
A new species of the genus Neostylodactylus Hayashi & Miyake, 1968 (Crustacea: Decapoda: 
Stylodactylidae) from southern Japan Junji Okuno and Hiroyuki Tachikawa 


On the male of Scutumara enodis Ng & Nakasone, 1993 (Crustacea: Decapoda: Brachyura: Grapsidae) 
N. K. Ng and T. Komai 
Larval development of Cryptolithodes expansus Miers (Decapoda: Anomura: Lithodidae) reared in the 
laboratory Mi Hyang Kim and Sung Yun Hong 

A new genus of pinnotherid crab from the Indian Ocean (Crustacea: Decapoda: Brachyura) 
Raymond B. Manning and Bella Galil 
A new genus and species of ghost shrimp from Tobago, West Indies (Crustacea: Decapoda: 


Callianassidae) Richard Heard and Raymond B. Manning 
Griceus buskeyi, a new genus and species of calanoid copepod (Crustacea) from benthopelagic waters 
off Hawaii Frank D. Ferrari and E. L. Markhaseva 


Studies on the Crustacea of the Turks and Caicos Islands, British West Indies. IV. Heteromysis (Heteromysis) 
spottei, a new species (Peracarida: Mysidacea: Mysidae) from Pine Cay 
W. Wayne Price and Richard W. Heard 
Gynodiastylis laciniacristatus, a new species (Crustacea: Cumacea) from Australia 
Sarah Gerken and Jennifer Gross 
Revision of the subterranean amphipod genus Spelaeogammarus (Bogidiellidae) from Brazil, including 
descriptions of three new species and considerations of their phylogeny and biogeography 
Stefan Koenemann and John R. Holsinger 
Eudendrium bathyalis, a new species of hydroid (Hydrozoa: Anthomedusae: Eudendriidae) from 
Bermuda Antonio C. Marques and Dale R. Calder 
Cnidae of two species of Discosomatidae (Cnidaria: Anthozoa: Corallimorpharia) from Brazil 
Suzana Machado Pinto and Maria Julia da Costa Belém 
Additions to the cancellariid (Mollusca: Neogastropoda) fauna of South Africa 
Richard E. Petit and M. G. Harasewych 
Erpobdella lahontana (Annelida: Hirudinea: Arhynchobdellida: Erpobdellidae), a new species of fresh- 


water leech from North America Peter Hovingh and Donald J. Klemm 
A cladistic analysis of Sciomyzidae Fallén (Diptera) Luciane Marinoni and Wayne N. Mathis 
Review of the chewing louse genus Abrocomophaga (Phthiraptera: Amblycera), with description of 
two new species Roger D. Price and Robert M. Timm 


A new species of the genus Cubacubana (Insecta: Zygentoma: Nicoletiidae) from a Mexican cave 
Luis Espinasa 
Three new species of bathyal cidaroids (Echinodermata: Echinoidea) from the Antarctic region 
Rich Mooi, Bruno David, F. Julian Fell, and Thérése Choné 
A new species of Pristigaster, with comments on the genus and redescription of P. cayana (Teleostei: 
Clupeomorpha: Pristigasteridae) Naércio A. Menezes and Mario C. C. de Pinna 
A new species of Apogon (Perciformes: Apogonidae) from the Saya de Malha Bank, Indian Ocean, with 
redescriptions of Apogon regani Whitley, 1951, A. gardineri Regan, 1908, and A. heraldi 
(Herre, 1943) Thomas H. Fraser 
A new species of Pogonophryne (Pisces: Perciformes: Artedidraconidae) from East Antarctica 
Richard R. Eakin and Arcady V. Balushkin 
A new genus and species of inseminating fish (Teleostei: Characidae: Cheirodontinae: Compsurini) from 
South America with uniquely derived caudal-fin dermal papillae 
Luiz R. Malabarba and Stanley H. Weitzman 
Calls and calling behavior of the frog Leptodactylus natalensis (Amphibia: Anura: Leptodactylidae) 
W. Ronald Heyer and Celso Morato de Carvalho 
A new species of venomous coral snake (Serpentes: Elapidae) from high desert in Puebla, Mexico 
Jonathan A. Campbell 
Fossil Red-shouldered Hawk in the Bahamas: Calohierax quadratus Wetmore synonymized with Buteo 
lineatus (Gmelin) Storrs L. Olson 
Revision of the extant taxa of the genus Notiosorex (Mammalia: Insectivora: Soricidae) 
Leslie N. Carraway and Robert M. Timm 
Status of the name Odocoileus hemionus crooki (Mammalia: Cervidae) James R. Heffelfinger 
International Commission on Zoological Nomenclature 


88 


95 


104 


124 


129 


145 


155 
162 


210 


218 


224 


238 


249 


264 


269 


284 


291 


298 


302 
S19 
335 


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LIBRARIES 
Taxonofriry~2 


113(2):339-—355. 2000. 


* ROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


fT evolutionary relationships of Phillips’ small-eared 


-Shrew, Cryptotis phillipsii (Schaldach, 1966), from Oaxaca, Mexico 
(Mammalia: Insectivora: Soricidae) 


Neal Woodman and Robert M. Timm 


(NW) Department of Biological Sciences, East Stroudsburg University, 200 Prospect Street, 
East Stroudsburg, Pennsylvania 18301-2999, U.S.A.; (RMT) Natural History Museum and 
Department of Ecology & Evolutionary Biology, 

University of Kansas, Lawrence, Kansas 66045-2454, U.S.A. 


Abstract.—The name Cryptotis peregrina (Merriam, 1895) previously en- 
compassed two separate populations of a small-eared shrew of the Cryptotis 
mexicana-group inhabiting the Sierra de Cuatro Venados and the Sierra de 
Miahuatlan in Oaxaca, Mexico. Analysis of museum specimens from these two 
populations, including the type series of C. peregrina, indicates that they are 
neither conspecific nor even sister taxa. We taxonomically restrict the name C. 
peregrina to populations in the Sierra de Cuatro Venados and adjoining Sierra 
Yucuyacua, whereas the name Cryptotis phillipsii (Schaldach, 1966) is avail- 
able for the population in the Sierra de Miahuatlan. Moreover, we demonstrate 
that C. phillipsii is the most primitive member of the C. mexicana-group, 
whereas, C. peregrina is one of the more derived members of the group. 


Choate’s (1970) comprehensive revision 
of the small-eared shrews of the genus 
Cryptotis divided the Mexican and Central 
American species into three informal 
groupings: the ‘‘Cryptotis mexicana- 
group,” the “‘Cryptotis parva-group,”’ and 
a group of three “relict species.”’ At that 
time, C. mexicana (with four subspecies), 
Cryptotis goldmani (with two subspecies), 
and the monotypic Cryptotis goodwini 
comprised the C. mexicana-group. Based 
on cranial and postcranial morphology (in 
particular, unique modifications of the fore- 
limb) and biogeographic patterns, we 
(Woodman & Timm 1999) recently revised 
the C. mexicana-group, providing evidence 
that it may represent a natural grouping dis- 
tinct from other members of the genus. Our 
revision recognized eight species in the C. 
mexicana-group, many of which were treat- 
ed previously as subspecies (Choate 1970, 
Hall 1981): C. alticola, C. goldmani, C. 


goodwini, C. griseoventris, C. mexicana, C. 
nelsoni, C. obscura, and C. peregrina. 
Cryptotis peregrina, which is endemic to 
Oaxaca, Mexico, was first described by 
Merriam (1895) as a subspecies of C. mex- 
icana based on a series of 24 specimens 
collected by E. W. Nelson and E. A. Gold- 
man in 1894 ‘from mountains 15 miles 
[south] west of city of Oaxaca”’ (Merriam 
1895:24). Binford (1989) identified this 
range of mountains as the Sierra de Cuatro 
Venados. In addition to the type series, 
Choate (1970) referred specimens from the 
Sierra de Miahuatlan in southern Oaxaca, 
including the holotype and two paratypes of 
Notiosorex phillipsii, to C. mexicana pere- 
grina. We initially agreed with Choate’s as- 
sessment, referring material from the Sierra 
de Cuatro Venados and the Sierra de Mia- 
huatlan to the species Cryptotis peregrina 
(Woodman & Timm 1999). Cranially, the 
shrews previously considered by Choate 
(1970) and subsequent authors (e.g., Hall 


340 


1981) to be subspecies of C. mexicana (C. 
mexicana, C. nelsoni, C. obscura, C. pere- 
grina) are quite similar and can be difficult 
to distinguish. However, the foreclaws vary 
noticably in length and breadth among 
members of the C. mexicana-group (Choate 
1970, Woodman & Timm 1999), and we 
showed that other aspects of the forelimb, 
in particular the morphology of the humer- 
us, are distinct as well (Woodman & Timm 
1999). Our phylogenetic analysis indicated 
that C. peregrina was the most plesio- 
morphic member of the C. mexicana-group. 

Our study (Woodman & Timm 1999) fo- 
cused mainly on the larger, broad-clawed 
shrews that we termed the Cryptotis gold- 
mani-group, which are a subset of the C. 
mexicana-group. Initially, we were interest- 
ed in the former subspecies of C. mexicana 
primarily as outgroups to aid in polarizing 
characters for phylogenetic analysis. In 
studying Cryptotis peregrina, we depended 
heavily on specimens from the Sierra de 
Miahuatlan in the University of Kansas 
Natural History Museum collection, be- 
cause they were readily available to us and 
included the only postcranial material then 
known for the species. One inconsistency 
we noted between our study and Choate’s 
revision was that he referred to C. [m.] mex- 
icana as “having less highly developed 
front feet and claws’? (Choate 1970:232) 
than C. [m.] peregrina. However, the spec- 
imens from the Sierra de Miahuatlan had 
distinctly shorter and narrower claws than 
C. mexicana. We also observed that their 
humeri were less modified that those of C. 
mexicana, adding to our view of their being 
more plesiomorphic. 

Recently, we re-studied the holotype and 
type series of Cryptotis peregrina at the Na- 
tional Museum of Natural History and dis- 
covered that Choate’s (1970) observations 
were correct—specimens in the type series 
from Sierra de Cuatro Venados all possess 
longer, broader claws than C. mexicana. In 
addition, we were graciously granted per- 
mission by the American Museum of Nat- 
ural History to remove the humerus from 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


the only known fluid-preserved specimen of 
C. peregrina from the Sierra Yucuyacua, a 
mountain range in western Oaxaca that ad- 
joins the Sierra de Cuatro Venados. Study 
of these specimens indicates that the two 
populations of Cryptotis inhabiting the Si- 
erra de Cuatro Venados\Sierra Yucuyacua 
complex and the Sierra de Miahuatlan, re- 
spectively, are not conspecific. Herein, we 
taxonomically restrict the name Cryptotis 
peregrina to shrews inhabiting the former 
region. The name Cryptotis phillipsii 
(Schaldach, 1966) is available for the pop- 
ulation in the Sierra de Miahuatlan. Much 
of what we reported earlier in regard to C. 
peregrina (Woodman & Timm 1999) actu- 
ally refers to a combination of C. peregrina 
and C. phillipsii. It is our purpose here to 
redescribe C. peregrina and C. phillipsii in 
light of this new information and to clarify 
the distinctions among the species in the 
Cryptotis mexicana-group. In addition, we 
present a refined hypothesis of phylogenetic 
relationships among the species in this 
group of shrews. 


Methods 


Our current work used the techniques 
and methods of our previous studies of 
shrews (Woodman & Timm 1992, 1993, 
1999), and more detailed accounts of meth- 
odology are presented there. Species and 
subspecies synonomies list only published 
uses of names. Regional names, place 
names, and coordinates in Oaxaca derive 
from Binford (1989), Choate (1970), and 
our review of maps and gazetteers of the 
region. Terminology of dentition and dental 
characteristics follows Choate (1970). An- 
atomical terminology of the humerus and 
other aspects of the postcranial skeleton fol- 
lows Reed (1951). Measurements used in 
our analyses follow Woodman & Timm 
(1993, 1999); abbreviations of measure- 
ments are explained in Table 1. Measure- 
ments of the skull were taken to the nearest 
0.1 mm using either an ocular micrometer 
in a binocular microscope or a hand-held 


VOLUME 113, NUMBER 2 


dial caliper (for condylobasal-length and 
cranial-breadth). Univariate statistics in- 
clude mean + standard deviation. 

Multivariate analysis was carried out us- 
ing Minitab 8.0. Our principle components 
analysis (PCA; Fig. 1, Table 2) used ten 
logarithm-transformed cranial measure- 
ments (condylobasal-length, breadth-of-zy- 
gomatic-plate, interorbital-breadth, breadth- 
across-U's, breadth-across-U?s, breadth- 
across-M?’s, palatal-length, length-of-upper- 
toothrow, length-of-unicuspid-toothrow, 
length-of-upper-molariform-toothrow) from 
12 Cryptotis peregrina and 14 C. phillipsii 
with complete crania. 

Phylogeny and character evolution were 
analyzed using PAUP 4.0b2 (see Swofford 
1998) and MCCLADE 3.0 (see Maddison 
& Maddison 1992). Phylogenetic analysis 
(Fig. 2) was carried out using an exhaustive 
search of 32 unordered and equally-weight- 
ed transition series (Table 3, Appendix I), 
with Cryptotis parva parva and C. nigres- 
cens serving as outgroups for polarizing 
characters. 

Specimens from the following institu- 
tions were used in this study: American 
Museum of Natural History, New York 
(AMNH); Natural History Museum, Lon- 
don (BM); California Academy of Scienc- 
es, San Francisco (CAS); Escuela Nacional 
de Ciencias Biolédgicas, Mexico City 
(ENCB); Instituto de Biologia, Universidad 
Nacional Aut6noma de México, Mexico 
City (UNAM); University of Kansas Natu- 
ral History Museum, Lawrence (KU); Nat- 
ural History Museum of Los Angeles 
County, Los Angeles (LACM); Museum of 
Comparative Zoology, Cambridge (MCZ); 
James Ford Bell Museum of Natural His- 
tory, St. Paul (MMNH); Museo de Zoolo- 
gia, Facultad de Ciencias, Universidad Na- 
cional Autonoma de México, Mexico City 
(MZFC); Texas Cooperative Wildlife Col- 
lection, College Station (TCWC); Univer- 
sity of Michigan Museum of Zoology, Ann 
Arbor (UMMZ); National Museum of Nat- 
ural History, Washington (USNM). 


341 


Results of Multivariate Analysis 


Principle components analysis (PCA) of 
cranial variables supports the separation of 
Cryptotis peregrina and Cryptotis phillipsii. 
A plot of factor scores on factor axes | and 
2 shows a clear distinction between these 
two species with minimal overlap (Fig. 1). 
This plot emphasizes the generally larger 
cranial size (factor axis 1—Table 2) of C. 
phillipsii, despite the generally similar ex- 
ternal measurements of the two species (Ta- 
ble 1). In the region of size overlap between 
the two species on the plot, C. peregrina 
and C. phillipsii are separated by the com- 
bined effects of the two axes, suggesting a 
common, but offset trend of decreasing uni- 
cuspid-toothrow-length and palatal-length 
and increasing interorbital-breadth and pal- 
atal-breadth (M2B) with increasing size 
(Table 2). 


Revised Phylogeny of the 
Cryptotis mexicana-group 


Our re-analysis of phylogenetic relation- 
ships within the Cryptotis mexicana-group, 
based on 32 transition series (Table 3), re- 
sulted in a single most parsimonious tree of 
67 steps (Fig. 2). Discounting differences 
such as the exclusion of Cryptotis goodwini 
magnimana and the inclusion of Cryptotis 
phillipsii as a species separate from Cryp- 
totis peregrina, the new tree essentially 
matches three of the nine most parsimoni- 
ous trees from our previous phylogenetic 
analysis of this group of shrews (Woodman 
& Timm 1999, fig. 13, central column of 
trees). The four species that comprised the 
Cryptotis goldmani-group (C. alticola, C. 
goldmani, C. goodwini, C. griseoventris) 
form the crown clade of the C. mexicana- 
group, whereas the species previously sub- 
sumed under the name C. mexicana (C. 
mexicana, C. nelsoni, C. obscura, C. pere- 
grina, C. phillipsii) appear as individual 
branches at different levels of the tree. This 
supports our previous view that C. mexi- 
cana (sensu latu) was paraphyletic with re- 
spect to the C. goldmani-group. 


342 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Table 1.—Measurements of Cryptotis mexicana-group shrews used in this study. Statistics presented are mean 
+ standard deviation of the mean, and observed extremes. 


C. mexicana 
(Oaxaca) C. nelsoni C. obscura C. peregrina C. phillipsii 


Skin measurements 


n= 118 n=9 n= 55 n= 24 n=27 
Head and body length (HB) 
69 + 4 16.23 66 = 4 F2-s03 68 + 6 
55-88 72-79 56-75 66-78 58-78 
Tail length (TL) 
Of Gracies DX NaS 26s o 30) E72 Sas ¥3 
20-34 22-33 20-32 24-33 23-36 
Craniomandibular measurements 
n = 30 n=6 n = 32 n= 12 n= 14 
Condylobasal-length (CBL) 
18.7 + 0.4 L9:5.250)3 18.4 + 0.4 19.1 +-04 1952 05 
17.5-19.3 19.1—20.0 17.4-19.1 18.3-19.8 18.8—20.2 
Breadth-of-braincase (BB) 
9:82 0:3 104 43.033 916. > 02 9:0°> 0.1 10.3 =O 
9.3-10.7 9.9-10.7 9.2—10.2 9.7-10.1 9.8-10.7 
(n = 28) (n = 30) (n = 11) (n = 13) 
Breadth-of-zygomatic-plate (ZP) 
LS oy 220622 (Oe 2.0) 25 Ost 1.6-2,04 20.70% 
1.42.0 1.0—2.2 1.8—2.1 1.5—-1.8 1.8—2.3 
Interorbital-breadth(IO) 
A.J 102 5. OH ry 0.1 A Tea b 419) =O 
4.3-5.0 4.8-5.2 4.44.9 4.64.9 4.75.1 
Breadth-across-first-unicuspids (U1B) 
DAO 28 Of 25041 yes vagal 0) | 26° = Ol 
2.3—2.5 2.6—2.9 2.3—2.6 2.1—2.6 2.5—2.8 
Breadth-across-third-unicuspids (U3B) 
DSpHaOst 8.3 (0.2 3° Nie ze 0 i | Gage ate cel Ya | 655 Wea | 
2.7—3.0 3.0—3.4 2.6—3.2 2.6—3.0 2.9-3.4 
Breadth-across-second-molars (M2B) 
5 4ee 2 oni 02 543 pmueel 0 all Soe OD 5.5. = O11 
5.1-5.8 5.9-6.3 5.1-5.5 5.35.8 5.6-6.0 
Palatal-length (PL) 
S2ra OP? $4 02 FR pause 0 92 S50 = OZ 6.5. = U2 
7.3-8.5 8.1-8.7 7.5—-8.5 8.2-8.9 8.0-8.7 
Length-of-upper-toothrow (TR) 
[2 a0 2 Pie farsume | E02 Pa = (Oe fess | i | dies get BS 
6.6—7.5 7.4-7.8 6.8—7.4 7.2—7.5 7.2-8.0 
Length-of-unicuspid-toothrow (UTR) 
245-10 DAO 2A OM 2 SAO ay 2 Ot 
2.1—2.5 2.42.6 2.1—2.6 2.4—2.6 2.3—2.7 
Length-of-upper-molariform-toothrow (MTR) 
a.2 = Ol SP et (0 Id 2 + Ol 573) = OR 54! I. 2 
4.9-5.5 5.45.8 4.9-5.4 5.1-5.4 5.3-5.8 


VOLUME 113, NUMBER 2 343 


Table 1.—Continued. 


C. mexicana 
(Oaxaca) C. nelsoni C. obscura C. peregrina C. phillipsii 


Posterior-width-of-M! (WM1) 


Lo = 0.1 1.9 eg Onl eBrs=O:1 tc pica rg | 
1.5-1.8 — 1.6-1.8 1.6—2.0 1.6—2.0 
(n = 25) (n = 1) 
Length-of-mandible (LM) 
6.0.+ 0.2 6.4.+ 0.1 aie. 90.2 G2 20:2 64. +.0.3 
5.4—6.3 6.36.6 5.4-6.4 5.86.4 5.8-7.1 
Height-of-coronoid-process (HCP) 
4.3 + 0.1 4.8 + 0.1 44+ 0.1 es se Oo 43 + 0.1 
4.1.6 4.74.8 4.0-4.6 4.3-4.7 4.7-5.1 
Height-of-coronoid-valley (HCV) 
26 +. 0.1 26 = Ol Pe = (4 PS.5051 Deduct Oy] 
2.32.8 2.8-2.9 2.42.8 2.7—3.0 2.72.9 
Height-of-articular-condyle (HAC) 
Be =*0:.2 3.9 ='0.05 So == 10:2 5.8: 0:1 GS ae 
3.43.9 3.84.1 3.2-3.9 3.64.1 3.84.2 
Breadth-of-articular-condyle (BAC) 
mo 0.1 i Ae seee 0| 29) = Oh S30 051 cs ie aa, 6 Bs | 
2.7—3.2 3.0—3.2 2.6—3.0 2.8-3.2 3.0-3.3 
Articular-condyle-to-M, (AC3) 
ae 30.2 a2 =) Ox A. G.==.O:2 Sb cae) 2 3.0.2. 02 
4.4-5.2 5.05.3 4.3-5.0 4.8-5.3 4.8-5.3 
Length-of-lower-toothrow 
so O.1 6.02 0.1 Sp eee 0 Ua SQ) 2=0ek 5.9 = 02 
5.4—-5.9 5.8-6.2 5.45.9 5.8—6.0 5.5-6.2 
Length-of-lower-molar-row (m13) 
fest /(). 1 AS 0). 4 42+ 0.1 44+ 0.1 4.4 + 0.2 
4.14.5 4.44.6 4.0-4.4 4.2-4.5 4.0-4.8 
(n= 6) 
Length-of-m1 (m1L) 
a ae OL 1.8 + 0.05 tec OL Le Ow a fae | 
1.6—1.9 1.8-1.9 1.6—1.8 1.6—-1.8 1.6—2.0 
Weight (g) 
TAL 1D — OF S14 _ $.0)2- 14 
5.0-9.5 5.0—10.0 6.5—10.0 
(n = 37) (n = 23) (n = 19) 


Cryptotis phillipsii appears as the most the more primitive members of the C. mex- 
primitive member of the Cryptotis mexi- icana-group. Based on all but two charac- 
cana-group, matching our supposition _ ters (relative tail length and relative height 
based on the morphology of its forelimb. In of the coronoid process of the mandible) C. 
contrast, Cryptotis peregrina is much more’ peregrina could, in fact, be considered a 
derived and serves as a “‘link’’ between the member of the C. goldmani-group. How- 
Cryptotis goldmani-group of shrews and ever, both characters are much more vari- 


344 


Table 2.—Factor loadings for the first two factor 
axes from principle components analysis of Cryptotis 
peregrina and Cryptotis phillipsii from Oaxaca. The 
two axes accounted for 72% of the variation. Abbre- 
viations as in Table 1. 


Variable PC] EG2 

UTR =0)1;70 0.378 
PL =Orh95 0.523 
IO —0.208 —0.458 
MTR —0.334 SOAMOZ 
TR —=(}335 0.233 
M2B —0:337 33:25 
PL es =O:3)/i1 0.221 
CBE =O:3571 Uiz22 
U1B =O;372 —0.198 
U3B =0;373 —~O.Za9 


factor 2 


O C. peregrina 


@ C phillipsii 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


able among the more primitive members of 
the C. mexicana-group (Table 4) than we 
previously realized (Woodman & Timm 
1999). For this reason, we redefine the C. 
goldmani-group below, excluding these two 
characters. 


Systematic Biology 
“Cryptotis mexicana-group”’ Choate, 1970 


We (Woodman & Timm 1999) recently 
redescribed the Cryptotis mexicana-group 
as small to medium-sized members of the 
genus with long, dark dorsal pelage. All 
members possess variably broadened fore- 
feet and variably broadened and elongated 
foreclaws; posterior border of the zygo- 
matic plate usually positioned equal to, or 


-6 -4 -2 0 2 4 6 
factor 1 
Fig. 1. Plot of factor scores from PCA of ten log-transformed measurements from 12 Cryptotis peregrina 


and 14 Cryptotis phillipsii. Factor axis 1 represents size (Table 2); factor loadings are negatively weighted, so 
the largest individuals have the smallest scores. Factor axis 2 represents a combination of palatal-length, inter- 
orbital-breadth, unicuspid-toothrow-length, and breadth-across-M?s (Table 2). 


VOLUME 113, NUMBER 2 


20 


1 


345 

C. parva 
C. nigrescens 
C. mexicana 

E C. nelsoni 

. peregrina 

. alticola 

C. goldmani 

C. goodwini 

. gnseoventris 


C. obscura 


C. phillipsii 


Fig. 2. Topology of the single most parsimonious tree resulting from phylogenetic analysis of the Cryptotis 


mexicana-group of shrews. Cryptotis nigrescens and C. parva parva served as the outgroups for polarizing 


t—) 


characters. Number of transitions is shown for each branch. Tree length = 67 (minimum possible length = 49; 


maximum possible length = 122); consistency index = 


index = 0.753; homoplasy index = 0.269. 


slightly posterior to, the posterior base of 
the maxillary process; upper toothrow un- 
crowded; dentition not bulbous; anterior 
border of the coronoid process of the man- 
dible joins the horizontal ramus at a rela- 
tively low angle; posterior border of lower 
incisor extends to posterior cingulum of 
P,; relatively long distance from the coro- 
noid process to the posterior border of M;; 
tall, wide articular face of the articular pro- 
cess; deep lower sigmoid notch; relatively 
long, low P,; relatively short, broad meta- 
carpals; shortened and broadened humerus 
with elongated processes and a dorsoven- 
trally elongate head; deeply pocketed pos- 
terior edge of the falciform process of the 
tibia. 

Included species.—Cryptotis alticola, C. 
goldmani, C. goodwini, C. griseoventris, C. 


0.731; rescaled consistency index = 0.551; retention 


mexicana, C. nelsoni, C. obscura, C. pere- 
grina, and C. phillipsii. 


Cryptotis peregrina (Merriam, 1895) 


Blarina mexicana peregrina Merriam, 
1895:24. 

C[ryptotis]. mexicana peregrina: Miller, 
911222. 

Cryptotis mexicana peregrina: Hall & Kel- 
son, 1959:60; Goodwin, 1969:39 (in 
part); Choate, 1970:237 (in part); Hall, 
1981:59 (in part). 

Cryptotis mexicana mexicana: Goodwin, 
1969:39 (in part). 

Cryptotis peregrina: Woodman & Timm, 
1999:35 (in part). 


Holotype.—Skin and skull of male, 
USNM 68317; captured on 12 September 


346 


Table 3.—Complete character matrix for phylogenetic analysis of members of the Cryptotis mexicana-group and two outgroups, Cryptotis parva parva, and Cryptotis 


nigrescens. Transition series (TS) are explained in Appendix 1. 


Transition series 


N 


Taxon 


parva 


nigrescens 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Sc A ce re ce ee ee ee en | 


SKS SES iw i i i oo 


See eee ee ee! 


N 


mexicana 


N 


nelsoni 


obscura 


peregrina 
phillipsii 
alticola 


goldmani 


[oe) 


goodwini 


3 


griseoventris 


1894 by E. W. Nelson and E. A. Goldman 
(collector number 6748). 

Type locality.—‘‘mountains 15 miles 
[south]west of city of Oaxaca [de Juarez], 
Mexico (altitude, 9,500 feet).’’ According 
to Goldman (1951), the collectors spent 
September 10th to 19th, 1894, traveling to 
and from, and working at, this locality. This 
trip took them ca. 8 mi southwest of Oaxaca 
de Juarez to Cuilapan, then west into the 
mountains, passing Santa Inéz [Santa Inés 
del Monte, 16°54’30"N, 96°52’W] and 
climbing over the summit to a wet meadow 
called Neverfa Herrera on the western 
slope. 


“These high mountains, which appeared to have no 
distinctive name, are somewhat isolated, the only 
connection with the high mountains to the east be- 
ing through low hills north of Oaxaca. The upper 
slopes are well watered, and general conditions 
much as on Cerro San Felipe. Specimens were la- 
beled “Mountains 15 miles west of Oaxaca,’ but as 
we later learned the direction was more nearly 
southwest.”’ (Goldman, 1951:218). 


Merriam (1895) reported that the speci- 
mens were obtained from a variety of ele- 
vations between 8800 [2680 m] and the 
summit at 9500 ft [2895 m], indicating that 
the type series of Cryptotis peregrina was 
collected at or near the top of the range. On 
topographic maps, the summit appears to be 
near a landmark called La Plazuela 
[16°56'52”N, 96°53'45"W]. Binford (1989) 
refers to these mountains as the Sierra de 
Cuatro Venados. 

Distribution.—Known from 2680 to 
3200 m in the Sierra de Cuatro Venados, 
central Oaxaca, and the Sierra Yucuyacua, 
western Oaxaca (Fig. 3). 

Description.—Size medium to large for 
the genus (Table 1); tail long, averaging 30 
mm, or ca. 42% (Table 4) of head-and-body 
length; dorsal guard hairs 6—7 mm long; 
forepaws enlarged; foreclaws noticeably 
elongate and broadened; rostrum of mod- 
erate length (PL/CBL = 44.5%, Table 4); 
often two well-developed dorsal foramina 
(44%, Table 4); foramen posterior to dorsal 
articular facet leading to ventral extension 


VOLUME 113, NUMBER 2 


Oaxaca 
re de Juarez 


Fig. 3. 


of the sinus canal typically present on one 
or both sides of the cranium (94%, Table 
4); foramen dorsal to dorsal articular facet 
typically absent (87%, Table 4); zygomatic 
plate narrow (ZP/PL = 19.3%, Table 4), an- 
terior border usually aligned with posterior 
¥%3 of mesostyle-metastyle valley or metas- 
tyle of M'; U* unreduced in size, surface 
area >'% that of U’; U* usually aligned with 
the unicuspid toothrow and partially visible 
in lateral view of the skull; P*, M', and M? 
slightly to moderately recessed on posterior 
border; protoconal basin of M! reduced rel- 


347 


Map of part of Oaxaca, Mexico, illustrating the known distribution of Cryptotis peregrina (*) and 
Cryptotis phillipsii (@). The 2000 m contour is shown. 


ative to hypoconal basin; M? simple: usu- 
ally possessing paracrista, paracone, pre- 
centrocrista (all pigmented), mesostyle, and 
very short postcentrocrista; M?* lacking 
metacone, hypocone absent or poorly-de- 
veloped, and a reduced, typically unpig- 
mented protocone occasionally present. Ar- 
ticular process of the mandible generally 
moderately tall and wide, with a moderately 
broad lower articular facet; moderately high 
coronoid process of the mandible (HCP/ML 
= 73.1%, Table 4); entoconid usually pre- 
sent on M, (73%, Table 4), typically vesti- 


348 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Table 4.—Characteristics among the five species previously synonomized with Cryptotis mexicana. 


C. mexicana 
(Oaxaca) C. nelsoni C. obscura C. peregrina C. phillipsii 

Foramen of sinus canal 

7% O% 19% 94% 24% 

n = 90 n=8 n = 52 n= 16 n= 17 
Foramen dorsal to dorsal articular facet present 

13% 38% 28% 13% 82% 

n = 90 n=8 n = 36 n= 15 n=17 
Two distinct dorsal foramina present 

54% 50% 22% 44% 47% 

n = 95 n=6 n= 58 n= 16 n= 19 
Posteroventral border of unicuspids 

concave concave straight to convex concave concave 
Entoconid of M, present 

100% 100% 96% 73% 93% 

68% well-developed 100% well-developed 96% well-developed 100% vestigal 83% vestigal 

= 77 n=7 n = 26 n = 22 n= 13 

Humerus modified 

slightly slightly extremely slightly 
Relative tail length (TL/HB x 100) 

392 5 38 + 4 40° + 6 42 iS 46 + 6 

26-52 28-44 29-55 36-47 30-59 

n= 118 n=9 n= 55 n = 24 n = 27 
Relative rostrum length (PL/CBL xX 100) 

Asal = 0:8 45.2, = 0.8 ya i use [| 44.5.+ 0.7 433 = 0.6 

41.7-45.1 42.0-44.3 41.0-45.6 43.246.0 42.244.4 

n = 30 n=6 n = 32 n= 12 n= 14 
Relative breadth of zygomatic plate (ZP/PL x 100) 

2166: = V6 23a A PAST ae bal 19.32 .1.0 234° 2-46 

17.9-24.4 21.8—25.6 22.2—26.9 18.1-21.2 21.4—26.4 

n = 30 n=7 n = 32 n= 12 n= 14 
Relative breadth of zygomatic plate (ZP/CBL Xx 100) 

9.4 + 0:7 10:2 = 0:6 10:6: ="65 S67 0'5 LO = Os 

7.9-10.4 9.5—11.2 9.7-11.5 7.8—-9.4 9.2-11.4 

n = 30 n=6 n = 32 n= 12 n= 14 
Relative length of unicuspid toothrow (UTR/CBL X 100) 

12a) 0.4 12 Duct 0,5 13:0) 27054 129° +03 12 EO:6 

11.4-13.3 12.0—-13.4 11.6—13.9 12.1-13.4 11.4—-13.7 

n = 30 n=6 n = 32 n= 12 n= 14 
Relative palatal breadth (M2B/PL Xx 100) 

65:9 = 2.5 T2o = OO 67.4 + 2:2 63.2 se DS 6827 271 

60.0—72.6 70.9-73.3 62.4—70.7 60.9—-68.7 65.1—72.5 

n = 30 n=7 n = 32 n= 12 n= 14 
Relative height of coronoid process (HCP/ML x 100) 

TOS = ScD 73.8 13 74.3-= 2:0 MSA es 16 Tse) tae 

65.1—79.6 72.3-76.2 70.2—83.6 69.4—75.0 71.6—86.2 

n = 30 n=8 n = 32 n= 12 n= 16 


VOLUME 113, NUMBER 2 349 
Table 4.—Continued. 
C. mexicana 
(Oaxaca) C. nelsoni C. obscura C. peregrina C. phillipsii 
Relative posterior length of mandible (AC3/ML X 100) 
80.8 + 2.6 yA aos a 77.0 + 3.0 Seal = yh 1484 = 3.6 
77.4—86.2 77.8-82.5 72.6—85.5 77.8-85.0 73.2-86.2 
n = 30 n= 8 n = 32 n= 12 n= 16 
Relative extension of articular condyle (AC3/HCP x 100) 
Haw = 5.) 108:8-+ 1,7 105.6) 37 iA 20 103.7 = 3.9 
100.0—124.4 106.4—110.6 100.0—114.3 109.9-115.6 96.1—110.6 
n = 30 n= 8 n = 32 n= 12 n= 16 


gial and unpigmented when present. Broad, 
curved humerus, with elongated processes 
(Fig. 4B). 

Comparisons.—Cryptotis peregrina is 
distinctive among the species previously 
considered to be subspecies of Cryptotis 


mexicana. It has a narrow zygomatic plate, 
and it has the broadest forepaws and the 
longest and broadest foreclaws. The hu- 
merus is shorter, more curved, and gener- 
ally more highly derived: the pectoral pro- 
cess is higher, lateral epicondyle more ex- 


Fig. 4 Left humeri of (A) Cryptotis phillipsii (KU 124299) and (B) Cryptotis peregrina (AMNH 149965). 
The humerus labeled “C. peregrina” in Woodman and Timm (1999: Fig. 15B) is from a specimen (KU 124298) 
that we now recognize as C. phillipsii. 


350 


panded, and teres tubercle and medial epi- 
condyle longer and more closely situated to 
each other. Cryptotis peregrina is the only 
species in the C. mexicana-group besides 
Cryptotis goldmani that usually has a well- 
developed foramen of the ventral branch of 
the sinus canal (Table 4; Woodman & 
Timm 1999). 

Cryptotis mexicana: Cryptotis peregrina 
has slightly longer, broader claws and a 
much more derived humerus; M®? simple, 
lacking metacone; entoconid of M, vesti- 
gial (rather than well-developed), when 
present. 

Cryptotis nelsoni: Cryptotis peregrina 
has slightly longer, broader claws and a 
much more derived humerus; skull narrow- 
er overall; longer rostrum and much nar- 
rower palate (Table 4); M? simple, lacking 
metacone; entoconid of M, vestigial (rather 
than well-developed), when present. 

Cryptotis obscura: Cryptotis peregrina 
has much longer, broader claws and a much 
more derived humerus; longer rostrum (Ta- 
ble 4); upper unicuspids concave on pos- 
terior surface (rather than straight or con- 
vex), appearing curved; M> simple, lacking 
metacone; more likely to have two well-de- 
veloped dorsal foramina (44% vs. 22%, Ta- 
ble 4); entoconid of M, vestigial (rather 
than well-developed), when present. 

Cryptotis phillipsii: Cryptotis peregrina 
has much longer and broader claws and a 
much more derived humerus; longer ros- 
trum and much narrower palate (Table 4); 
absolutely and relatively broader zygomatic 
plate (Tables 1, 4); larger U* relative to U’; 
slightly more recessed upper dentition; 
slightly less complex M?; well-developed 
foramen of the ventral extension of sinus 
canal; usually no foramen dorsal to dorsal 
articular facet. 

Cryptotis goldmani-group: Cryptotis per- 
egrina is generally smaller in head-and- 
body-length and has a relatively longer tail 
(Table 1, 4); narrower zygomatic plate; 
more likely to possess a vestigial entoconid 
(rather than no entoconid) of M?. 

Remarks.—Nearly all known specimens 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


of Cryptotis peregrina are part of the type 
series, collected in the Sierra de Cuatro 
Venados ca. 15 mi SW of Oaxaca de Jua- 
rez in 1894. Merriam (1895) stated that 
there were 25 specimens in the type series, 
but he did not list them individually. We 
were able to locate only 24 original spec- 
imens collected by Nelson and Goldman. 
The only other specimen of C. peregrina 
of which we are aware is a fluid-preserved 
individual (with the skull and left humerus 
removed) in poor condition, collected “‘N. 
La Muralla”’ in the Sierra Yucuyacua by 
Thomas B. MacDougall on 18 March 
1952. Locality notes by MacDougall indi- 
cate this locality is at 10,500+ ft at the 
‘“‘top of Cerro Yucunino” (AMNH field 
notes, 1952). We believe (as apparently did 
Choate, 1970:239) that this peak is equiv- 
alent to Cerro Yucuyacua [= Piedra de 
Ollila], based on its elevation and position 
relative to La Muralla and Santa Maria 
Asuncion Tlaxiaco. 

Goldman (1951:218) noted that the type 
locality was moist, with general conditions 
‘“‘much as on Cerro San Felipe.’’ Areas 
above 8000 ft on Cerro San Felipe were 
covered in oak-dominated mixed forest 
with an herbaceous understory, and they 
experienced cold night-time temperatures, 
with frosts recorded in August. Meadows 
(neverias), such as the one in the vicinity 
of the type locality, were said to be em- 
ployed by local people for making ice to be 
used at lower elevations. MacDougall 
(AMNH field notes, 1952) noted the pres- 
ence of pine and oak where he captured C. 
peregrina north of La Muralla. 

Specimens examined (25).—Mexico: Oa- 
xaca: Nforth]. [of] La Muralla [16°58'N, 
97°56'W], top of Cerro Yucunin6 [= Cerro 
Yucuyacua, 17°06’N, 97°40'W], Tlaxiaco 
[Santa Maria Asuncion Tlaxiaco, 17°16'N, 
97°41'W], 10,500+ ft (AMNH 149965); 
mountains 15 mi [south] west Oaxaca de 
Judrez [16°30°N, 96°33 Wi. 9500 aa 
(BMNH 95.11.1.26 [= USNM 68337], 
USNM 68315-68336, 68338, including ho- 
lotype). 


VOLUME 113, NUMBER 2 


‘““Cryptotis goldmani-group’’> Woodman & 
Timm, 1999 


Description.—The C. goldmani-group is 
a subset of the C. mexicana-group that is 
comprised of medium-sized members of the 
genus with greatly broadened forefeet; ex- 
tremely long, broad foreclaws; fourth upper 
unicuspid usually aligned with the unicus- 
pid toothrow and partially visible in labial 
view of the cranium; protoconal basin of 
M! reduced relative to hypoconal basin; M° 
simple, hypocone absent or poorly devel- 
oped and lacking metacone; entoconid of 
M, vestigial or absent; and extremely broad 
humerus with greatly elongated processes. 

Included species.—Cryptotis alticola, C. 
goldmani, C. goodwini, C. griseoventris, 
and C. peregrina. 


Cryptotis phillipsii (Schaldach, 1966) 


Notiosorex (Xenosorex) Phillipsii Schal- 
dach, 1966:289; Goodwin, 1969:43. 

Cryptotis mexicana machetes: Musser, 
1964:6; Schaldach, 1966:288; Goodwin, 
1969:40 (in part). 

Cryptotis mexicana peregrina: Goodwin, 
1969:40 (in part); Choate, 1970:237 (Gn 
part); Hall, 1981:59 (in part). 

Cryptotis peregrina: Woodman & Timm, 
i225) (in part). 


Holotype.—Skin and skull of adult fe- 
male, UNAM 8445; taken 18 Dec 1964 by 
W. J. Schaldach, Jr. (collector number 
13278). Two paratypes, KU 114226 [= 
UNAM 8446] and UNAM 8447. 

Type locality.—*‘the Rio Molino, 3 ki- 
lometers S.W. San Miguel Suchixtepec, al- 
titude 2250 meters, southern Oaxaca, Méx- 
ico”’ (Schaldach, 1966:289). 

Distribution.—Known from 1060 to 
2745 m in the Sierra de Miahuatlan, south- 
ern Oaxaca, Mexico (Fig. 3). 

Description.—Size medium to large for 
the genus (Table 1); tail long, averaging 31 
mm, or ca. 46% (Table 4) of head-and-body 
length; dorsal guard hairs 6-7 mm long; 
forepaws somewhat enlarged; foreclaws 


351 


among the narrowest for members of the 
Cryptotis mexicana-group, but elongate and 
somewhat broadened relative to other mem- 
bers of the genus; rostrum of moderate 
length (PL/CBL = 43.3%, Table 4); often 
two well-developed dorsal foramina (47%, 
Table 4); ventral extension of the sinus ca- 
nal and associated foramen typically lack- 
ing (76%, Table 4), but well-developed 
when present; a foramen dorsal to the dor- 
sal articular facet usually present on one or 
both sides of the skull (82%, Table 4); zy- 
gomatic plate broad (ZP/PL = 23.4%, Table 
4), anterior border usually aligned with pos- 
terior % of mesostyle-metastyle valley or 
metastyle of M'; P*, M', and M? unrecessed 
to slightly recessed on posterior border; sur- 
face area of U* typically =% that of U*; U* 
usually aligned with the unicuspid tooth- 
row, but not typically visible in lateral view 
of the skull; protoconal basin of M' same 
size as hypoconal basin; M? simple, lacking 
metacone, hypocone poorly-developed or 
absent; M? usually possessing paracrista, 
paracone, precentrocrista, and mesostyle 
(all pigmented), and occasionally a very 
short postcentrocrista and a well-developed, 
pigmented protocone. Articular process of 
the mandible generally moderately tall and 
wide, with a moderately broad lower artic- 
ular facet; coronoid process of the mandible 
moderately high relative to mandibular 
length (HCP/ML = 75.7%, Table 4); ento- 
conid usually present on M, (92%, Table 4), 
typically vestigial (83% of those possessing 
entoconids), but occasionally well-devel- 
oped (17%). The tall, broad, slightly curved 
humerus has somewhat elongated process- 
es, but it is among the least modified within 
the C. mexicana-group (Fig. 4). 
Comparisons.—Cryptotis phillipsii has 
the least modified forelimbs of any member 
of the Cryptotis mexicana-group: it has the 
smallest forepaws and the shortest and nar- 
rowest foreclaws; the humerus is among the 
longest and least curved; teres tubercle is 
short, medial epicondyle is short and has a 
Straight proximal edge, and teres tubercle 
and medial epicondyle are broadly separat- 


352 


ed. Cryptotis phillipsii also has the longest 
tail relative to head-and-body-length (Table 
4). There may be a local tendency for some 
C. phillipsii to develop without U*s (12%, 
n = 17) in the region of the type locality, 
resulting in a reduced dentition. 

Cryptotis mexicana: Cryptotis phillipsii 
has shorter, narrower foreclaws and a rela- 
tively longer tail; less modified humerus; 
broader palate (Table 4); simple M?, lacking 
metacone; entoconid of M, usually vestigial 
(rather than well-developed) when present. 

Cryptotis nelsoni: Cryptotis phillipsii has 
shorter, narrower foreclaws and a relatively 
longer tail; narrower palate (Table 4); sim- 
ple M?, lacking metacone; entoconid of M, 
usually vestigial (rather than well-devel- 
oped) when present. 

Cryptotis obscura: Cryptotis phillipsii 
has a relatively longer tail; upper unicus- 
pids concave on posterior surface, appear- 
ing curved, rather than convex and cone- 
shaped; more likely to have two well-de- 
veloped dorsal foramina (Table 4); simple 
M?, lacking metacone; entoconid of M, usu- 
ally vestigial (rather than well-developed) 
when present. 

Cryptotis goldmani-group: Cryptotis 
phillipsii is smaller in head-and-body- 
length, and has much shorter, narrower fo- 
reclaws, a much less derived humerus, and 
a relatively longer tail; higher coronoid pro- 
cess (Table 4); shorter posterior portion of 
mandible (Table 4); much more likely to 
possess vestigial entoconid (rather than no 
entoconid) of M,. 

Remarks.—The type series of Cryptotis 
phillipsii was collected in 1964 by William 
J. Schaldach, Jr. and Allan R. Phillips dur- 
ing a trip to southern Oaxaca to obtain 
specimens of birds and mammals. Schal- 
dach (1966) originally described a new spe- 
cies, Notiosorex phillipsii, on the basis of 
three specimens: a skin with skull and a 
skin without skull from Rio Molino 
(UNAM 4445, 4447) and a skin with skull 
from Rio Guajalote (KU 114226 = UNAM 
8446). Originally identified as Cryptotis 
mexicana, the main consideration in sub- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


sequently referring these three specimens to 
Notiosorex appears to have been the pres- 
ence of only three unicuspids in the upper 
toothrow, rather than the four unicuspids 
typical of Cryptotis. A fourth specimen 
(UNAM 8444), collected at Rio Molino and 
possessing four unicuspids, was reported as 
Cryptotis mexicana (Schaldach 1966). Not- 
ing strong differences in external, cranial, 
and dental characters between the type se- 
ries of N. phillipsii and specimens of Notio- 
sorex crawfordi and Megasorex gigas that 
he used for comparison, Schaldach (1966) 
erected a new subgenus, Xenosorex, to dis- 
tinguish the new shrew. Choate (1969) re- 
examined the type series of Notiosorex 
phillipsii and reported a number of distinc- 
tive cranial, mandibular, and dental char- 
acters that clearly established N. phillipsii 
as a member of the genus Crypfotis, despite 
its reduced dentition. He justifiably placed 
Schaldach’s N. phillipsii as a junior syno- 
nym of Cryptotis [mexicana] peregrina, 
where it has remained until now (Choate 
1970, Hall 1981, Hutterer 1993, Woodman 
& Timm 1999). 

Cryptotis phillipsii is known from higher 
elevation forest, including cloud forest, in 
the Sierra de Miahuatlan of southern Oa- 
xaca. The species is syntopic with Cryptotis 
goldmani at some localities, but the geo- 
graphic distribution of C. phillipsii is not 
nearly as broad as that of C. goldmani, de- 
spite an apparently lower elevational limit. 

Schaldach (1966) reported the humid, 
shaded ravines along the slopes of the val- 
ley of the Rio Molino at the type locality 
contained remnant, pine- and oak-dominat- 
ed cloud forest, with a dense understory of 
ferns, mosses, vines, and shrubs. Musser 
(1964) described the canyon bottom as con- 
taining moist, open forest with bromeliad- 
covered oaks and a dense ground cover of 
shrubs, ferns, and herbaceous vegetation. 
The higher, drier slopes were covered with 
a mixed secondary growth of pines and 
oaks. Musser (1964:6) took one C. phillip- 
sil, two Sorex saussurei oaxacae, and 29 
Peromyscus megalops under a “‘lush her- 


VOLUME 113, NUMBER 2 


baceous ground cover’’ consisting largely 
of ferns along the steep banks of a “‘moist, 
densely vegetated streamside”’ with a “‘deep 
layer of wet humus and leaf litter’? at 7300 
ft [2225 m] near Rio Molino. Paul B. Rob- 
ertson (KU field notes and catalog, 1969, 
1970) captured three C. phillipsii at Rio 
Molino. In December 1969, he trapped one 
C. phillipsii, one Microtus mexicanus, two 
Peromyscus aztecus, two P. levipes, and 
four P. megalops. In April 1970 using a 
trapline “‘along a small rivulet,’ he took 
one C. phillipsii with two Cryptotis gold- 
mani, one Sorex saussurei oaxacae, one 
Microtus mexicanus, one Neotoma mexi- 
cana, two Oryzomys chapmani, 12 Pero- 
myscus megalops, and one Reithrodonto- 
mys mexicanus. Nearby, “‘among & under 
rocks in an overgrown field,’’ he collected 
a C. phillipsii with two Reithrodontomys 
megalotis, three R. sumichrasti, and one 
Sigmodon alleni. He described the stream- 
side vegetation at Rio Molino as “hard 
wood”’ and the slopes as “almost pure 
pine.’ In addition, Schaldach (1966) re- 
ported capturing Sorex mutabilis (as Sorex 
veraepacis) and Liomys pictus at Rio Mol- 
ino. Cryptotis phillipsii and C. goldmani 
were collected in syntopy also at “‘lumber 
camp, km 158” along the Puerto Angel 
Road in January 1970 by John R. Arnold 
and Craig Moe. Cryptotis phillipsii and C. 
parva pueblensis were captured syntopical- 
ly 20 mi S, 5 mi E San Miguel Sola de Vega 
by Percy L. Clifton in June—July 1964. He 
described the habitat there as dense cloud 
forest, with C. phillipsii “‘caught under a 
rock in a low damp situation along a creek”’ 
(KU field notes, 1964). Cryptotis phillipsii 
also was taken in syntopy with C. p. pue- 
blensis at San Miguel Suchixtepec by 
Thomas B. MacDougall in October 1967. 
Specimens examined (36).—Mexico: Oa- 
xaca: 20 mi S, 5 mi E Sola de Vega [San 
Miguel Sola de Vega, 16°31'N, 96°59’W], 
4800 ft (KU 98728); 36 km (by road) N 
San Gabriel Mixtepec [16°06’N, 97°06'’W], 
1680 m (KU 124294); 27.8 km (by road) 
N San Gabriel Mixtepec, 1320 m (KU 


353 


124295, 124296); La Cima, Puerto Escon- 
dido road, km 184.5 [16°12'N, 97°07'W], 
5750 ft (CAS 15473); Puerto Escondido 
road, km 193 [16°10’N, 97°07'W], 4200 ft 
(CAS 15474); Sinai [Finca Sinai, 16°07'N, 
97°08'W], 10 km [by trail] E Nopala [Santo 
Reyes Nopala], 7200 ft (CAS 14940); Rio 
Molino [16°04'N, 96°28'’W], 2250-2745 m 
(AMNH 213758, 213759, 214152, 214803- 
214805; KU 121661, 124298, 124299; 
UNAM 8444, 8445, 8447—includes holo- 
type); near campemento Rio Molino (Hwy 
175), 7300 ft (UMMZ 112572); Puerto An- 
gel road, km 153 [16°04'N, 96°28’W], 7100 
ft (CAS 14068); San Miguel Suchixtepec 
[16°OS’N, 96°28’W], Miahuatlan District 
(AMNH 214806—214808); 16 km SW [San 
Miguel] Suchixtepec, 2000 m (ENCB 3413, 
3414); Lovene [16°02’N, 96°12’W], Mia- 
huatlan (AMNH_ 178739); lumber camp, 
Puerto Angel road, km 158, 8375 ft (CAS 
15478); Rio Guajalote [16°00'N, 96°28’ W], 
2000 m (KU 114226); Rio Jalatengo 
[15°58’N, 96°27'W], Puerto Angel road, km 
178, 4275 ft (CAS 14069, 14071, 14072, 
15475; UNAM 27518); Puerto Angel road, 
km 195, 3475 ft (UNAM 26551); Puerto 
Angel road (UNAM 27517). 


Acknowledgments 


We thank Robert P. Anderson for valu- 
able assistance in tracking down informa- 
tion helpful to our study. We thank the fol- 
lowing curators and collection managers for 
loans and for permission to study important 
specimens under their care: Guy G. Musser 
and Robert Voss (AMNH); Robert C. Dow- 
ler (ASNHOC); Paula Jenkins (BM); Douglas 
J. Long (CAS); Ticul Alvarez (ENCB); 
John E. Heyning (LACM); Maria E. Rutz- 
moser (MCZ); Elmer C. Birney (MMNH); 
Livia Leén P. (MZFC); George D. Baum- 
gardner (TCWC); Phil Myers (UMMZ); 
Fernando Cervantes R. (UNAM); Michael 
D. Carleton, Alfred L. Gardner, Linda K. 
Gordon, Helen L. Kafka, and Richard W. 
Thorington, Jr. (USNM). Two reviewers 


354 


provided helpful comments that improved 
our manuscript. 


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. 1970. Systematics and zoogeography of Mid- 
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Appendix I: Transition Series Used in Phylogenetic 
Analysis 


Transition series (TS) marked by an asterisk (*) 
were modified from Woodman & Timm (1999). TS 
30—32 were added since that paper. 


*1. length of foreclaws: short (0); elon- 
gate (1); more elongate (2); greatly 
elongate (3). 

*2. breadth of foreclaws: narrow (0); 
broad (1); broader (2); extremely 
broad (3). 

3. forefeet: small (0); enlarged, broad- 
ened (1); greatly enlarged and broad- 
eneau(2). 

4. metacarpals: long, narrow (0); short, 
broad (1). 

5. posterior border of zygomatic plate: 
even with or anterior to anterior root 
of maxillary process (0); even with 
posterior root of zygomatic process, 
but separated from it by posterior bor- 
der of palate (1); even with (or pos- 
terior to) and confluent with posterior 
root of zygomatic process (2). 

6. anterior border of coronoid process: 
steep, forming a narrow angle with 
horizontal ramus of mandible (Q); less 
steep, forming a wide angle with hor- 
izontal ramus of mandible (1). 

7. articular condyle: low and broad (0); 
high and narrow (1). 

8. lower sigmoid notch: very shallow 
(O};-deep’ (Pf): 

9. shape of unicuspids (U!—U*): cone- 
shaped, posteroventral border straight- 
edged or convex (0); narrow, poster- 
oventral border concave (1). 

10. protoconal basin of M!: about equal in 


VOLUME 113, NUMBER 2 


i. 


ee 


1S. 


14. 


15. 


*16, 


eat. 


ca > 


£S: 


20. 


DA. 


cpp 24 


size to hypoconal basin (0); reduced 
relative to hypoconal basin (1). 

M? morphology: simple, metacone ab- 
sent (O); complex, metacone present 
Ch). 

shape of P,: short and high (0); long 
and low (1). 

shape of humerus: long, narrow, rela- 
tively straight (0); short, robust, 
curved (1); short, robust, and very 
curved (2). 

head of humerus: rounded (0); dorso- 
ventrally elongate (1). 

ventral edge of proximal face of great- 
er tuberosity of humerus: rounded (0); 
with broad, deep pocket (1). 

pectoral process of humerus: low (0); 
high (1). 

length of teres tubercle: relatively 
short (0); elongate (1); greatly elon- 
gate (2). 

medial epicondyle of humerus: short 
(O); elongate (1); greatly elongate (2). 
lateral epicondyle of humerus: small 
(O); expanded (1). 

teres tubercle and medial epicondyle 
of humerus: far apart (0); close to- 
gether (1); very close (2); extremely 
close (3). 

posterior edge of falciform process of 
tibia: not deeply pocketed (0); deeply 
pocketed (1). 

foramen of sinus canal: absent in 
100% of specimens (0); absent in 
>75% of specimens (1); present in 
>50% of specimens, but vestigial (2); 
present in >90% of specimens, well- 
developed (3). 


24. 


F202; 


26. 


Dike 


28. 


Pas J 


#30; 


pele 


*52. 


. dorsal foramina: 


two in <75% of 
specimens (0); two in >75% of spec- 
imens (1). 

foramen dorsal to articular facet: pre- 
sent in <75% of specimens (0); pre- 
sent in >75% of specimens (1). 
entoconid of M,: present in >90% of 
specimens, typically well-developed 
(O); present in >50% of specimens, 
typically vestigial (1); absent in >80% 
of specimens (2). 

body size (head-and-body length): 
smaller, mean <69 (QO); larger, mean 
> ie), 

relative tail length (% of head-and- 
body length): short, mean <39% (0); 
long, mean >40% (1). 

upper unicuspid toothrow: crowded, 
three unicuspids visible in lateral view 
(0); uncrowded, four unicuspids visi- 
ble in lateral view (1). 

dorsal guard hairs: short, <4 mm (0); 
of intermediate length, 4-5 mm (1); 
long, >5 mm (2). 

distal edge of pectoral process of hu- 
merus relative to teres tubercle: ex- 
tends well distal to teres tubercle (0); 
extends to proximity of distal edge of 
teres tubercle (1). 

bones of hind paws: metatarsals and 
proximal phalanges long narrow (0); 
metatarsals and proximal phalanges 
broad (1); metatarsals and proximal 
phalanges short and very broad (2). 
posterior border of P*, M', and M7’: 
slight to no emargination (0); notica- 
ble emargination (1); strong emargi- 
nation (2). 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


113(2):356—368. 2000. 


Two new species of flightless rails (Aves: Rallidae) from the Middle 


Pleistocene ‘‘crane fauna”? of Bermuda 


Storrs L. Olson and David B. Wingate 


(SLO) Department of Vertebrate Zoology, National Museum of Natural History, 
Smithsonian Institution, Washington, D.C. 20560-0116, U.S.A.; 
(DBW) Bermuda Natural History Museum, P.O. Box FL 145, Flatts, Bermuda 


Abstract.—Two new species of flightless rails are described from a Pleis- 
tocene fauna in Bermuda that also includes an extinct crane (Grus latipes) and 
an extinct duck (Anas pachyscelus). The medium-sized Rallus ibycus, new 
species, was possibly derived from North American populations of Virginia 
Rail (R. limicola), but had a longer bill, much more robust legs, and reduced 
wings and pectoral girdle. The very small Porzana piercei, new species, except 
for the reduced wing and pectoral girdle, is very similar to the extant Yellow- 
breasted Crake (P. flaviventer), which now occurs only in the Neotropics, in- 
cluding the Greater Antilles. The fauna that included these rails developed 
during a long, stable glacial period of lowered sea-levels in the Middle Pleis- 
tocene, during which the entire Bermuda platform was emergent. This was 
followed by an abrupt and extreme interglacial about 400,000 years ago when 
sea-levels rose to 21 m above present levels, obliterating most of Bermuda and 


much of its endemic fauna, including the rails. 


The island of Bermuda, situated in the 
western North Atlantic 1050 km east of 
Cape Hatteras, North Carolina, is composed 
almost entirely of calcareous aeolianite on 
the southeastern rim of a submerged, trun- 
cated summit of an extinct volcano. The 
aeolianite has been modified by solution to 
form numerous caves and fissures that are 
accessible to collectors either through nat- 
ural openings or through limestone quar- 
rying operations. These caves frequently 
contain accumulated fossils of vertebrates 
and invertebrates, mainly birds and terres- 
trial gastropods. Although much of interest 
to avian paleontology has been collected 
from Bermuda, relatively little has been 
published, partly for lack of a better under- 
standing of the complexities of the island’s 
stratigraphy and chronology, which has 
been greatly improved in recent years. 

The first contribution to knowledge of 
Pleistocene birds in Bermuda was that of 


Wetmore (1960) who described a new ge- 
nus and species of endemic crane, Baeo- 
pteryx latipes, and an extinct endemic duck, 
Anas pachyscelus, from the Wilkinson 
Quarry, Hamilton Parish. The genus Baeo- 
pteryx was later synonymized with typical 
cranes of the genus Grus (Fischer & Ste- 
phan 1971). 

Wetmore (1960:10) mentioned that “‘the 
collection contains various bones from four 
species of rails, one very small, two of in- 
termediate size, and one nearly as large as 
the modern clapper rail [Rallus longiros- 
tris]. These are not clearly marked in the 
present collection so that no attempt is 
made to describe them here in detail, par- 
ticularly since complete material for one of 
them is now in other hands for study.’’ Ac- 
tually, among the rail material that Wetmore 
examined, there are remains of only two 
species of rails, one medium-sized and the 
other very small (uncataloged specimens in 


VOLUME 113, NUMBER 2 


USNM). Wetmore’s information on Ber- 
muda rails apparently came in part from 
Pierce Brodkorb, University of Florida, 
who had been in Bermuda collecting fossil 
rails with Wingate only weeks before Wet- 
more’s publication appeared. Later, Olson 
(1977:353-354) briefly mentioned the still 
undescribed fossil rails from Bermuda. 
Brodkorb’s collections containing the Ber- 
muda rails passed to the Florida Museum 
of Natural History after Brodkorb’s death 
in 1992, and we have now belatedly begun 
the process of describing them. 

Some information on the relationships 
and adaptations of the largest species of rail 
to which Wetmore alluded was provided by 
Olson (1997). This species was not contem- 
poraneous with the “‘crane fauna’”’ and does 
not appear to have co-existed with any oth- 
er endemic rail on Bermuda. In February 
1999, we discovered a new deposit in 
which this species was the predominant 
bird. Preliminary results from amino acid 
racemization ratios from associated snail 
shells indicate that it originated during a 
much younger glacial period. We have post- 
poned description of this species until the 
new material can be fully prepared and an- 
alyzed. Here we describe the two species 
that are definitely part of the “‘crane fauna”’ 
that we believe dates back at least to the 
Middle Pleistocene. 


Materials and Methods 


The specimens obtained by Wingate and 
Brodkorb in 1960 were collected in asso- 
ciation with bones of the extinct crane Grus 
latipes, from a vertical fracture on the east- 
ern face of the Bermuda Government quar- 
ry in Hamilton Parish that was filled with 
soil and snail shells of the genus Poecilo- 
zonites and was named the “Crane Crev- 
ice.”’ The fossil birds, formerly part of the 
Pierce Brodkorb collection, are now cata- 
loged in the Florida Museum of Natural 
History, University of Florida, Gainesville, 
and all take the prefix UF PB, which we 
have omitted except in the citations of the 


357 


holotypes and the figure legends. There are 
many more specimens available than are 
listed among the type material. As para- 
types we have listed those specimens that 
were used in the descriptions, are illustrat- 
ed, or were used for any of the cited mea- 
surements. Measurements of long bones of 
the limbs usually do not include broken or 
juvenile specimens, which accordingly are 
not among the paratypes listed. Measure- 
ments were taken with digital calipers to 
0.01 mm and rounded to the nearest 0.1 
mm. 

Comparative material examined:—Skel- 
etons (complete unless otherwise indicated) 
of the following species in the collections 
of the National Museum of Natural History, 
Smithsonian Institution (USNM), unless 
otherwise noted. Amaurolimnas concolor 
613963; Coturnicops noveboracensis 
556931; Laterallus albigularis 611563; L. 
Jamaicensis 492195, 502495; Pardirallus 
maculatus 561272—76; Poliolimnas ciner- 
eus 560913; Porzana flaviventer 501640, 
561276—78, and trunk skeletons 430043, 
430931, 430979, 431339—41; Porzana por- 
zana 552914; P. albicollis 562750; P. atra 
562788; (PS carolina. 501052, 501671; \P. 
pusilla 291704-05; Rallus aquaticus 
431545, 553039, 553041, UF 34461; R. 
limcola. 4899795, S25915; S259 E17, UF 
19598, UF 19769, UF 24324, UF 24322; R. 
elegans 499437, 525886, 610780; R. lon- 
girostris: S258 16529873, 525879; R: Jon- 
girostris X R. elegans 525887. 


Systematics 


Family Rallidae 
Genus Rallus Linnaeus 


The very long, slender bill of the follow- 
ing species clearly places it in the genus 
Rallus in its strict sense (Olson 1973a). No 
flightless species of the restricted genus 
Rallus have been described previously. 
Such characters as are preserved in the fos- 
sils that are not obscured by flightless ad- 
aptations show no evidence of relationship 
to the species of Pardirallus (including Or- 


358 


tygonax), which also have long bills but are 
osteologically quite distinct and not espe- 
cially closely related to Rallus (Olson 
1973a). 


Rallus ibycus, new species 
Figs. 1-5 


Holotype.—Premaxillary symphysis with 
anterior portion of internarial bar (pila su- 
pranasalis) UF PB5403. Collected in May 
1960 by David B. Wingate. 

Type locality.—Bermuda, Hamilton Par- 
ish, Government Quarry, Crane Crevice. 

Chronology.—Middle Pleistocene, pre- 
sumably within Oxygen Isotype Stages 13 
to 20, approximately 800 to 450 kya (see 
discussion). 

Measurements (mm) of holotype.— 
Length of premaxillary symphysis, 15.5. 

Paratypes.—Premaxillary symphyses 
5511, 5521. Pila supranasalis 5526. Crania 
5401, 5512, 5518. Sterna 5402, 5430. Cor- 
acoids 5404, 5415, 5416, 5441, 5456. Hu- 
meri 5405, 5417, 5422, 5425-5426, 5428, 
5457, 6063—6072, 6073, 6077, 6079. Ulnae 
5406, 5423, 5440, 5458. Carpometacarpi 
5407, 5419, 5424, 5439, 5459. Pelvis 5429. 
Femora 5409, 5432-5438, 5451, 5460, 5 
uncataloged specimens. Tibiotarsi 5410, 
5448-5449, 5452-5455, 5461, 5491-5492, 
5498-5502, 5503, 5541-5542, 5551-5560. 
Tarsometatarsi 5442, 5444-5447, 5462, 
5543-5546, 6001—6002, 6028-6031, 6055— 
6057. 

Measurements (mm) of paratypes.—Ros- 
trum: length from nasofrontal hinge esti- 
mated from 5526 and 5403 to be between 
46.5 and 48.0; length of premaxillary sym- 
physis 14.1, 15.5, 16.7. Cranium: length 
from naso-frontal hinge 28.0; width at post- 
orbital processes 15.5, 15.7; width of inter- 
orbital bridge 3.3, 3.6, 3.7. Coracoid: length 
15.4, 17.4, 18.2, 18.4, 19.5. Sternum: length 
along midline (from anterior sternal notch, 
not manubrium) 21.2, 21.3; width across 
coracoidal sulci 10.7, 12.1; depth of carina 
3.7, 4.4. Carpometacarpus: length 14.0, 
L6215.1'6.3,,:16.5,)16.95;proxamal -depthi3 .G; 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


4.1, 4.2, 4.3, 4.3. Ulna: length 22.8, 24.5, 
25.8, 26.9. Pelvis: sacrum length 27.3, 30.0; 
width across antitrochanters 15.4. See also 
Vable *1- 

Etymology.—Greek, m. ibykos, literally 
of a crane, but here a noun in apposition 
referring to Ibycus (i: 528. B.C.),"a lyme 
poet whose murder was revealed by cranes 
(see Anthon 1869). The name is applied to 
the new species from its association with 
the much larger fossils of cranes, the dis- 
covery of which also revealed the extinc- 
tion of the rail. 

Diagnosis.—Similar in overall size to 
Rallus limicola but bill longer and more 
slender, with a longer and more decurved 
premaxillary symphysis. Flightless, with 
the sternum and pectoral elements reduced. 
Hindlimb elements much more robust. 

Description.—In addition to the longer, 
more decurved bill, this species also seems 
to have the orbits reduced so the frontal 
area in lateral view has the appearance of 
sloping more steeply. 

Compared with Rallus limicola, the ster- 
num is shorter, wider, lacks a manubrial 
spine, and has a deep notch between the 
coracoidal sulci. The carina is very low and 
thick anteriorly, extending laterally as a 
rim, which, with the anterior ridge leading 
from the keel, creates a deep depression on 
the sides of the carina. 

The shaft of the coracoid is narrowed, 
forming a sharp ridge from the head to mid- 
shaft. The procoracoid process is smaller, 
more delicate and pointed. The sternocora- 
coidal process is much more pronounced 
and pointed due to the deep circular inci- 
sion in the external margin. 

The humerus has the head lower and 
smaller, the capital groove deeper, the distal 
end narrower, with the tricipital grooves 
deeper. The ulna is shorter and stouter and 
slightly less curved. The carpometacarpus 
is very reduced, short, and stout. 

The pelvis in dorsal view is decidedly 
wider, both anteriorly and posteriorly, al- 
though the ischial area in lateral view is not 
as deep as in R. limicola. 


VOLUME 113, NUMBER 2 


359 


Table 1.—Measurements (mm) of selected skeletal elements of two new species of flightless rails from the 


Middle Pleistocene of Bermuda. 


n Range Mean SD 
Rallus ibycus, n. sp. 
Humerus 
Length 18 28.5—36.3 32.3 ee. 
Proximal width 18 6.0—7.2 6.6 0.4 
Shaft width 18 1.7—2.2 1.9 0.1 
Distal width 18 4.0-4.8 4.4 0.2 
Femur 
Length 16 34.9-41.5 a7 9 2.6 
Proximal width 15 5.7-6.9 6.4 0.3 
Distal width [5 5.77.1 6.3 0.5 
Tibiotarsus 
Length from proximal articular surface 28 45.7-55.5 30/5 3.4 
Distal width 27 4.7-5.8 32 0.3 
Tarsometatarsus 
Length 19 27.1—34.3 30:2 2.4 
Proximal width 18 5.1-5.9 5.4 Ome) 
Distal width 18 5.1-6.1 5.6 0.3 
Porzana pierceéi, n. sp. 
Humerus 
Length 54 19.7—23.2 24 0:7 
Tibiotarsus 
Length from proximal articular surface 8 33.6—38.1 33.5 1.6 
Distal width Gi 3.13.4 fe 0.1 
Tarsometatarsus 
Length DS 19.9—23.9 22.3 0.9 
Proximal width 25 3:1=3:6 3.4 0.1 
Distal width 24 3.43.8 3.6 ot 


All of the elements of the hindlimb are 
extremely robust compared with R. limico- 
la, with heavier shafts and more expanded 
articulations. The head of the femur is pro- 
portionately larger. The tibiotarsus and tar- 
sometatarsus are proportionately shorter. 
The cnemial crests of the tibiotarsus are 
better developed, the fibular crest is longer, 
and the distal tendinal opening is larger. 
The tarsometatarsus has the trochleae more 
splayed, with the inner trochlea less elevat- 
ed and retracted than in R. limicola. 

Remarks.—The most likely progenitor of 
R. ibycus would be the Virginia Rail, Rallus 
limicola, a common migratory species in 
eastern North America that has been found 
as a “frequent but scarce vagrant’’ in Ber- 


muda (Amos 1991:121). Rails described as 
being similar to R. limicola are known from 
the Pliocene and Pleistocene of North 
America (Olson 1977). The Eurasian Water 
Rail, Rallus aquaticus, is a less likely an- 
cestor on geographic grounds, and also be- 
cause of its larger size. 


Genus Porzana Vieillot 


The following new species of flightless 
rail from Bermuda is characterized by very 
small size and a short “‘crake-like”’ bill that 
invites comparison with the New World 
species of Porzana, Laterallus, and Cotur- 
nicops. Of particular concern is the Neo- 
tropical Yellow-breasted Crake, usually 


360 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig, Ie 


Comparison of skulls (a—b) and sterna (c—e) of Rallus in lateral view. a, Rallus ibycus, new species 


(premaxillary symphysis, holotype UF PB5403; pila supranasalis UF PB5526; cranium UF PB 5401); b, e, R. 
limicola female USNM 525915; c, d, R. ibycus UF PB5430, UF PB5402. Scale bar = 2 cm. 


known as Porzana flaviventer, for which 
Ridgway (1920) once erected the monotyp- 
ic genus Hapalocrex (type Rallus flaviven- 
ter Boddaert). This species has little resem- 
blance to the various species of Porzana 
with which it has been placed in most cur- 
rent literature. Using mostly external char- 
acters, Olson (1970) suggested that it be 
placed in the genus Poliolimnas with the 
Australo-Malayan species P. cinereus. Al- 
though a few authors have accepted Olson’s 
conclusion (e.g., Short 1975), it has other- 
wise either been widely ignored or even 
Sharply attacked (Mees 1982). 


In an extensive morphological analysis of 
the Gruiformes emphasizing the Rallidae, 
Livezey (1998) found the relationships 
among the ‘‘crakes’’ and supposed allies to 
be difficult to resolve and even more diffi- 
cult to reconcile with traditional taxonomy. 
The preliminary results of an extensive 
sampling of mitochondrial DNA sequences 
of the Rallidae indicate that the genus Por- 
zana aS now generally construed (e.g., del 
Hoyo et al. 1996), is an unnatural assem- 
blage (Beth Slikas, National Zoological 
Park, pers. comm.). Until the taxonomic 
difficulties among the crakes can be better 


VOLUME 113, NUMBER 2 


361 


Fig.:2. 


Comparison of skulls (a—b) in dorsal view and sterna (c—e) in ventral view of Rallus. a, Rallus ibycus, 


new species (premaxillary symphysis, holotype UF PB5403; pila supranasalis UF PB5526; cranium UF PB 
5401); b, e, R: limicola female USNM 525915; c, d, R. ibycus UF PB5430, UF PB5402. Scale bar = 2 cm. 


resolved, we have chosen to continue with 
general usage in including flaviventer in the 
genus Porzana. 

The fossil species from Bermuda com- 
pares as follows with other crakes from 
which it might have been derived. In La- 
terallus the nostril is shorter and higher, the 
premaxillary symphysis is shorter, and the 
interorbital bridge is wider. In species of 
Porzana except P. flaviventer, the nostril is 
longer and the premaxillary symphysis is 
shorter (except in P. pusilla, which has a 
long symphysis). In Coturnicops the bill is 
extremely short and deep, quite unlike the 


fossil species. In the relatively short nostril, 
long premaxillary symphysis, and narrow 
interorbital bridge, the small Bermuda rail 
agrees perfectly with Porzana flaviventer. 
A striking feature is the proportions of 
the hindlimb. When the hindlimb elements 
of Porzana flaviventer are compared with 
those of Laterallus jamaicensis it is seen 
that whereas the tibiotarsi are of nearly 
equal length, the femur in P. flaviventer is 
much shorter and all the elements are stout- 
er. In Coturnicops noveboracensis the tibi- 
otarsus is shorter yet the femur is longer 
than in P. flaviventer. Compared with other 


362 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 3. Comparison of pectoral and wing elements of Rallus ibycus, new species (a—e) with R. limicola (f- 
i, female USNM 525915). a, f, right coracoids in ventral view (a, UF PB5415); b, c, g, left humeri in palmar 
view (b, UF PB5422; c, UF PB6072); d, h, left ulnae in internal view (d, UF PB5458); e, i, left carpometacarpi 
in internal view (e, UF PB5407). Scale bar = 2 cm. 


Fig. 4. Comparison of hindlimb elements in anterior view of Rallus ibycus, new species (a—f) with R. limicola 
(gi, USNM 525915 female; j-l, USNM 610783 male). a, left femur UF PB5438; b, right femur UF PB5432; 
c, right tibiotarsus UF PB5556; d, left tibiotarsus UF PB5500; e, right tarsometatarsus UF PB5462; f, left 
tarsometatarsus UF PB5544. Scale bar = 2 cm. 


VOLUME 113, NUMBER 2 


363 


Fig. 5. 


Pelves of rails in dorsal view. a, Rallus ibycus, new species (UF PB5429); b, R. limicola (USNM 610783 


male); c, Porzana piercei new species (UF PB5490); d, P. flaviventer (USNM 501640). Scale bar = 2 cm. 


species of Porzana, the hindlimb elements 
of P. flaviventer are more likewise more ro- 
bust, with the femur being proportionately 
shorter. As with the cranial elements, the 
hindlimb in the small Bermuda rail is most 
similar to that of P. flaviventer. The labels 
that Pierce Brodkorb left with the speci- 
mens indicate that he, too, had concluded 
that the small Bermuda rail was derived 
from P. flaviventer. The diagnosis of the 
new species is therefore based on its dif- 
ferences from that species. 


Porzana piercei, new species 
Figs. 5—9 


Holotype.—Complete rostrum UF 
PB5413. Collected in March 1960 by 
Pierce Brodkorb and David B. Wingate. 

Type locality.—Bermuda, Hamilton Par- 
ish, Government Quarry, Crane Crevice. 

Chronology.—Middle Pleistocene, pre- 
sumably within Oxygen Isotype Stages 13 
to 20, approximately 800 to 450 kya (see 
discussion). 

Measurements (mm) of holotype.— 
Length from nasofrontal hinge to tip, 16.8; 
length of premaxillary symphysis, 5.8. 

Paratypes.—Rostra 5421, 5536-5540. 
Crania 5412, 5527-5528, 5533. Mandible 
5481. Sternum 5414. Scapula 5482. Cora- 
coids 5464, 5474, 5483. Humeri 5465, 


5484, 5618, 5620, 5620 bis, 5621-5624, 
5624 bis, 5625-5629, 5631-5633, 5635-— 
5640, 5640 bis?, 5641-5643, 5645-5648, 
5650-5651, 5653, 5654-5671, 5686, 6195. 
Ulnae 5466, 5485. Carpometacarpi 5476, 
5487. Pelvis 5490. Femora 5468, 5478, 
5488, 1 uncataloged. Tibiotarsi 5469-5471, 
5510, 5547-5550. Tarsometatarsi (all rights 
except 5590 & 1 uncataloged with it) 3489, 
5575-5585, 5587-5590 + 1 uncataloged, 
5594, 5606—5607, 5617, 6060. 

Measurements (mm) of paratypes.— 
Scapula: length 17.4. Coracoid: length 10.5, 
hO:7, Biles Ulna: teneth 16:6, 17.5. Carpo- 
metacarpus: length 11.0, 11.2. Pelvis: sa- 
crum length 18.2, 19.0; width across anti- 
trochanters 11.1, 11.6. Femur: length 25.0, 
252, 26.2, 26:2; proximal width 3.8,.4.1, 
42, 4.2; distal width 3.7. 3.9, 3.9, 4.1. See 
also Table 1. 

Etymology.—Dedicated to the late Pierce 
Brodkorb who was instrumental in collect- 
ing rail material on Bermuda and who first 
recognized the affinities of this species. 

Diagnosis.—Very similar to P. flaviven- 
ter but differing in having the skull and bill 
somewhat larger and in being flightless, 
with the sternum and pectoral apparatus re- 
duced. 

Description.—Compared with P. flavi- 
venter the cranium is wider, the foramen 


364 


magnum larger, and the rostrum is wider 
with a shorter, wider premaxillary symphy- 
sis. The single specimen of mandible, a 
symphysis with the left dentary, is abraded 
at the tip but the symphysis is wider and 
deeper and the bone appears larger than in 
P. flaviventer. 

The single available sternum lacks the 
lateral processes but clearly indicates a 
flightless species, as the body of the bone 
is much shorter, but slightly wider, with a 
much reduced carina that is lower and more 
posteriorly situated than in P. flaviventer. 
The sternocoracoidal processes are narrow- 
er and angled less dorsally in anterior view. 
The intercoracoidal notch is wider and the 
manubrial spine lacking. 

The coracoids are reduced with more 
slender shafts and more delicate heads. The 
only available scapula has the articular end 
and anterior portion of the shaft wider, the 
acromion less pointed, and the glenoid facet 
larger. 

The humeri are only slightly smaller than 
in P. flaviventer but have a weaker shaft, a 
wider capital groove, and a reduced bicip- 
ital crest. The ulnae hardly differ except in 
the slightly weaker shafts. The carpometa- 
carpi differ only in slightly smaller size and 
proportionately shorter intermetacarpal 
space. 

The pelvis is wider in dorsal view and 
the postacetabular portion is proportionate- 
ly shorter. Apart from being slightly more 
robust, it is difficult to detect any differenc- 
es in the hindlimb elements from those of 
the living species, particularly given the 
very limited comparative material available 
for the latter. 

Remarks.—This species differs very little 
morphologically from Porzana flaviventer. 
Although clearly flightless, the degree of re- 
duction of the wing and pectoral girdle is 
relatively slight compared with many other 
flightless rails, probably indicating recency 
of derivation. 

The Yellow-breasted Crake, P. flaviven- 
ter, is a very small, delicate rail that typi- 
cally inhabits relatively large bodies of wa- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


ter with dense, emergent vegetation. Similar 
habitats were unlikely to have been present 
in Bermuda, so the birds must have been 
able to adapt to different ecological condi- 
tions, just as other aquatic rails have adapt- 
ed to even harsher insular environments on 
Ascension, St. Helena, and Laysan Islands 
(Olson 1973b). The modern distribution of 
P. flaviventer includes the Greater Antilles 
and the Neotropical mainland from south- 
ern Mexico to Argentina. An extralimital 
record from Antigua suggests that the spe- 
cies may have occurred in the Lesser An- 
tilles at least up until about 4300 years ago 
(Pregill et al. 1988:15). 

The Yellow-breasted Crake has never 
been found historically in Bermuda (Amos 
1991). Although it is possible that P. piercei 
was derived from populations of that species 
in the Greater Antilles, it seems equally 
plausible that the distribution of P. flaviven- 
ter may have extended into North America 
during periods of the Pleistocene, as was 
shown for the Neotropical rail Laterallus ex- 
ilis (Olson 1974), in which case colonization 
of Bermuda may have been directly from the 
North American mainland. 


Discussion 


Of critical importance is determining the 
age of the Bermudian avifauna that includ- 
ed at least a crane and a duck with reduced 
powers of flight and the two species of 
flightless rails described here. Of these fos- 
sils Wetmore (1960:1) could say no more 
than that “‘it is certain that they are old, and 
for the present it is my assumption that they 
date back to the Pleistocene.’’ Brodkorb (in 
Olson 1977:354) thought that they were 
‘‘post-Nebraskan, probably Aftonian or 
Kansan.’’ We now know, however, that the 
periodicity of sea-level and climatic chang- 
es associated with glacial cycles in the 
Pleistocene was much more complex than 
the traditional view of the four Nebraskan 
through Wisconsinan glacial periods, with 
their intervening interglacials (e.g., Hearty 
1998). 


VOLUME 113, NUMBER 2 


365 


Fig. 6. 


Comparison of skulls (a—d, lateral view; b—e, dorsal view) and dorsal views of mandibles (c, f) of 


Porzana piercei, new species (a—c) with a very small female P. flaviventer USNM 501640 (d—f). a—b, rostrum, 
holotype UF PB5413, and cranium UF PB5412; c, UF PB5481. Scale bar = 2 cm. 


There are as yet no direct radiometric 
dates on the “‘crane fauna”’ of Bermuda but 
circumstantial evidence points very strong- 
ly towards a long period of lowered sea lev- 
els in the Middle Pleistocene. The ‘“‘crane 
fauna” certainly did not survive into the 
Holocene as there is no mention of any 


such birds in the accounts of early settlers 
nor are there fossils of these birds in any 
late Quaternary deposits on the island. To 
have evolved and sustained such a diverse 
endemic avifauna, especially including a 
bird as large as a crane, which would have 
required extensive savanna-like habitat, the 


366 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig: 7: 


Comparison of elements of the pectoral girdle of Porzana piercei, new species (a—d) with a very 


small female P. flaviventer USNM 501640 (e—-h). a—b, sternum UF PB5414 in lateral and ventral views; c, right 
scapula in dorsal view; UF PB5482; d, left coracoid in ventral view UF PB5483. Scale bar = 2 cm. 


land area of Bermuda would have had to be 
at a maximum for a long period, indicating 
a glacial period of lowered sea-level of ex- 
ceptional duration. Thus, Brodkorb’s sug- 
gestion of Aftonian, which was an intergla- 
cial period between the Kansan and Ne- 
braskan glacial epochs, can be ruled out. 


Interglacials on Bermuda were periods of 
marine buildup of carbonate sands on the 
flooded Bermuda platform, whereas glacial 
periods were marked by sea-levels well be- 
low the surface of the platform when de- 
position of “terra rossa’”’ soils occured 
(Hearty & Vacher 1994). Thus, we would 


Fig. 8. 


Comparison of wing elements of Porzana piercei, new species (a—d) with a very small female P. 


flaviventer USNM 501640 (e—g). a—b, left humeri in anconal view UF PB5637 and UF PB5632; c, right ulna 
in internal view UF PB5485; d, right carpometacarpus UF PB5487. Scale bar = 2 cm. 


VOLUME 113, NUMBER 2 


367 


Fig.9. 


Comparison of hindlimb elements in anterior view of Porzana piercei, new species (a—f) with P. 


flaviventer (g—i, USNM 501640 small female; j—l, USNM 561276 unsexed). a, right femur UF PB5478; b, right 
femur UF PB5488; c, right tibiotarsus UF PB5470; d, right tibiotarsus UF PB5510; e, right tarsometatarsus UF 
PB9229; f, right tarsometatarsus UF PB5590. Scale bar = 2 cm. 


expect deposition of fossils associated with 
‘‘terra rossa’’ soils in caves and fissure fills 
to have taken place mainly during periods 
of lowered sea levels. The longest and most 
stable period of lowered sea-levels in the 
Quaternary history of Bermuda was from 
Oxygen Isotope Stage 13 back to about 
Stage 21 (Hearty & Vacher 1994:687, table 
1), which spanned a period from about 450 
kya to about 800 kya. Although sea-levels 
fluctuated during this time, as shown by the 
various isotope stages, the amplitude of the 
fluctuations was greatly reduced. Thus, 
even during the highest stands, the sea was 
at or below the level of the Bermuda plat- 
form through the preponderance of this in- 
terval, during which the subaerial portion 
of the platform would have been at its max- 
imum extent. It was during this time that 
deep limestone dissolution (karstification) 
and the accumulation of a massive “‘terra 
rossa’’ soil, the Castle Harbour Geosol, oc- 
curred (Hearty & Vacher 1994). 

This span of several hundred thousand 
years of maximum island emergence and 
relatively stable terrestrial conditions would 
probably have been the only time during 
Bermuda’s Pleistocene history when the 


“crane fauna”’ could have developed. Sub- 
sequent events also provide a very con- 
vincing explanation for the disappearance 
of this fauna, for it is now known that dur- 
ing interglacial Stage 11, 380 to 440 kya, 
sea levels rose much higher than had pre- 
viously been documented (Hearty et al. 
1999), reaching a height of over 20 m 
above present level. This would have re- 
duced the land area of Bermuda by two or- 
ders of magnitude, so that only a few small 
islets would have remained. In terms of the 
terrestrial biota, this event would essentially 
have wiped the slate clean, eliminating the 
crane, duck, and rails. A new period of col- 
onization and adaption during the next gla- 
cial period probably resulted in a substan- 
tially different avifauna. 


Acknowledgments 


We would like to thank the manager and 
staff of the Bermuda Government quarry 
for alerting Wingate to the extraordinary 
deposits of avian fossil material that were 
exposed by quarrying operations in 1959 
and 1960, and for their considerable coop- 
eration and assistance during collection of 


368 


specimens. We would also like to acknowl- 
edge the late Pierce Brodkorb for his part 
in collecting, numbering, and studying the 
fossil rail material from Bermuda. We thank 
Marc Frank and David W. Steadman, Flor- 
ida Museum of Natural History, Gaines- 
ville, for information, arrangements, access 
to fossil and modern collections, lending 
specimens, and other considerations. For an 
understanding and appreciation of aspects 
of the geology of Bermuda we are indebted 
to Paul J. Hearty, who also provided many 
useful comments on the manuscript. Helen 
James generated the statistics in Table 1 
from our measurements; the photographs 
are by John Steiner and Victor E. Krantz, 
Smithsonian Photographic Services, and 
these were made ready for publication by 
Brian Schmidt, Division of Birds, to all of 
whom we are especially grateful. This is 
Contribution #26, Bermuda Biodiversity 
Project (BBP), of the Bermuda Aquarium, 
Natural History Museum and Zoo. 


Literature Cited 


Amos, E. J. R. 1991. A guide to the birds of Bermuda. 
Published by the author, Warwick, Bermuda, 
206 pp. 

Anthon, C. 1869. A classical dictionary. Harper Broth- 
ers, New York, 1451 pp. 

del Hoyo, J., A. Elliot, J. Sargatal (eds.). Handbook of 
birds of the world, vol. 3. Lynx Editions, Bar- 
celona, 821 pp. 

Fischer, K., & B. Stephan. 1971. Ein flugunfahiger 
Kranich (Grus cubensis n. sp.) aus dem Pleis- 
tozin von Kuba—Eine Osteologie der Familie 
der Kraniche (Gruidae).—Wissenschaftlichen 
Zeitschrift der Humboldt-Universitaét zu Berlin 
Mathematisch-Naturwissenschaftliche Reihe 
20:541-592. 

Hearty, P. J. 1998. The geology of Eleuthera Island, 
Bahamas: a rosetta stone of Quaternary stratrig- 
raphy and sea-level history.—Quaternary Sci- 
ence Reviews 17:333-355. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


, & P.L. Vacher. 1994. Quaternary stratigraphy 

of Bermuda: a high-resolution pre-Sangamonian 

rock record.—Quaternary Science Reviews 13: 

685-697. 

, P. Kindler, P Cheng, & R. L. Edwards. 1999. 
A +20 m middle Pleistocene sea-level high- 
stand (Bermuda and the Bahamas) due to partial 
collapse of Antarctic ice.—Geology 27:375— 
S18: 

Livezey, B. C. 1998. A phylogenetic analysis of the 
Gruiformes (Aves) based on morphological 
characters, with an emphasis on the rails (Ral- 
lidae).—Philosophical Transactions of the Roy- 
al Society of London B 353:2077-2151. 

Mees, G. FE 1982. Birds from the lowlands of southern 
New Guinea (Merauke and Koembe).—TZoolo- 
gische Verhandelingen 191:1—188. 

Olson, S. L. 1973a. A classification of the Rallidae.— 
Wilson Bulletin 85:381—416. 

. 1973b. Evolution of the rails of the South At- 

lantic Islands (Aves: Rallidae).—Smithsonian 

Contributions to Zoology 152:1-—53. 

. 1974. The Pleistocene rails of North Ameri- 

ca.—Condor 76:169-175. 

. 1977. A synopsis of the fossil Rallidae. Pp. 

339-373 in S. D. Ripley, Rails of the world: A 

monograph of the family Rallidae. Boston, Da- 

vid R. Godine, 406 pp. 

. 1997. Towards a less imperfect understanding 
of the systematics and biogeography of the 
Clapper and King rail complex (Rallus longi- 
rostris and R. elegans). Pp. 93-111 in R. W. 
Dickerman, compiler, The era of Allan R. Phil- 
lips: A Festschrift. Albuquerque, New Mexico, 
Horizon Communications, 246 pp. 

Pregill, G. K., D. W. Steadman, S. L. Olson, & E V. 
Grady. 1988. Late Holocene fossil vertebrates 
from Burma Quarry, Antigua, Lesser Antil- 
les.—Smithsonian Contributions to Zoology 
463:1-27. 

Ridgway, R. 1920. Diagnoses of some new genera of 
bird—Smithsonian Miscellaneous Collections 
72(4): 1-4. 

Short, L. L. 1975. A zoogeographic analysis of the 
South American chaco avifauna.—Bulletin of 
the American Museum of Natural History 154: 
167-352. 

Wetmore, A. 1960. Pleistocene birds in Bermuda.— 
Smithsonian Miscellaneous Collections 140(2): 
1-11, 3 plates. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


113(2):369-—385. 2000. 


A new species of Anthias (Teleostei: Serranidae: Anthiinae) from the 


Galapagos Islands, with keys to Anthias and eastern 
Pacific Anthiinae 


William D. Anderson, Jr. and Carole C. Baldwin 


(WDA) Grice Marine Biological Laboratory, College of Charleston, 205 Fort Johnson, 

Charleston, South Carolina 29412-9110, U.S.A.; (CCB) Department of Vertebrate Zoology, 

National Museum of Natural History, Smithsonian Institution, 
Washington, D.C. 20560-0159, U.S.A. 


Abstract.—Anthias noeli, new species, is described from 17 specimens col- 
lected off the Galapagos Islands in the eastern Pacific, keys to the species of 
Anthias and to the species of eastern Pacific Anthiinae are provided, and com- 
ments are presented on hermaphroditism in the Anthiinae. The specimens of 
the new species, described herein, constitute the first authentic record of An- 
thias from the Pacific; the genus is otherwise known only from the Atlantic 
where it is represented by seven species. The following characters in combi- 
nation allow the separation of Anthias noeli from its congeners: soft rays in 
the dorsal fin 15 (rarely 16); tubed lateral-line scales 38 to 46; caudal-peduncle 
scales 22 to 25; lower jaw naked or with only a few scales posteriorly; gular 
region naked; total gillrakers on first gill arch 37 to 41; dorsal, anal, pelvic, 
and caudal fins each with some produced soft rays; anal fin angulate posteri- 
orly; depressed anal-fin length 32 to 43% SL; longest dorsal-fin spine (fourth 
or fifth) 14 to 20% SL; upper caudal-fin lobe (39-60% SL) longer than lower 


(38-57% SL); no teeth on tongue. 


During recent collecting trips to the Ga- 
lapagos Islands, investigators aboard sub- 
mersibles from the Harbor Branch Ocean- 
ographic Institution (Fort Pierce, Florida) 
have captured a number of new species, in- 
cluding 17 specimens of the new anthiine 
serranid described herein. Anthiines are 
small to medium size brightly colored fish- 
es that occur worldwide in shallow to mod- 
erate depths of tropical, subtropical, and 
temperate seas. They are usually associated 
with coral reefs or rocky bottoms that pro- 
vide refuge from predators. Individuals of 
most species feed upon zooplankton near 
the bottom and hastily seek shelter when 
approached by predators. Anthiines are of- 
ten seen in aggregations, with males attend- 
ing large harems. As far as is known, they 
are protogynous hermaphrodites, with 


many species being sexually dichromatic 
and some being sexually dimorphic, espe- 
cially in fin structure. 

There are approximately 185 valid de- 
scribed species in the serranid subfamily 
Anthiinae, classified variously in at least 25 
genera; additionally, there are a number of 
other known undescribed species of Anthi- 
inae in museum collections, and there are 
undoubtedly other species that remain to be 
discovered. 

Katayama & Amaoka (1986) restricted 
Anthias to include only Atlantic forms, re- 
moving Indo-Pacific species more appro- 
priately regarded as representatives of 
Pseudanthias, Franzia, and Mirolabri- 
chthys. Although not clearly stated, it ap- 
pears that Katayama & Amaoka (1986:217-— 
219, 221) considered Anthias to include the 


370 


following species: anthias, asperilinguis, 
helenensis, menezesi, nicholsi, salmopunc- 
tatus, tenuis, and woodsi. We modify their 
concept of the genus Anthias to exclude sal- 
mopunctatus and tenuis, which, as shown 
by Baldwin (1990), appear to be sister spe- 
cies and warrant placement in a genus dis- 
tinct from Anthias. In addition, we include 
in Anthias both the new species described 
herein and cyprinoides formerly assigned to 
Holanthias by Katayama & Amaoka 
(1986). 

Although we are not aware of any char- 
acter that is clearly synapomorphic for the 
species of Anthias, all eight are extremely 
similar morphologically and appear to form 
a natural group. Herein we provide a di- 
agnosis and description of the genus that 
distinguishes species of Anthias from all 
other serranid fishes. 


Methods 


Institutional abbreviations follow Leviton 
et al. (1985), except for HBOM (=Harbor 
Branch Oceanographic Museum). The 
methods used here are those of Anderson 
& Heemstra (1980), as modified by Ander- 
son et al. (1990). Some of those methods 
are reiterated or clarified below. Tubed lat- 
eral-line scales were counted on both sides 
of each specimen of the new species; other 
scale counts, except those around the caudal 
peduncle, were made on either side, de- 
pending on condition of the specimen. 
Counts of caudal-peduncle scales were of 
the number of scales around the narrowest 
part of the peduncle. In making counts of 
rows of cheek scales and counts of scale 
rows and scales above and below the lateral 
line, small scales at orbit and at bases of 
dorsal and anal fins were excluded. (Scales 
excluded from those counts are distinctly 
and abruptly smaller than adjacent scales in 
the counted series.) Scales above the lateral 
line were counted in a ventroposterior di- 
rection from origin of dorsal fin to, but ex- 
cluding, the lateral-line scale. Scales below 
the lateral line were counted in a dorsoan- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


terior direction from origin of anal fin to, 
but excluding, the lateral-line scale. Gillrak- 
ers (including rudiments, when present) on 
the first gill arch were counted on the right 
side. We follow Mabee (1988) and use ‘“‘su- 
praneural”’ instead of “‘predorsal’’ to refer 
to the short series of bones anterior to the 
pterygiophores that support the dorsal fin, 
and we conform with Johnson & Patterson 
(1993: 557) and Patterson & Johnson 
(1995) in using the term “‘epineurals”’’ for 
the intermuscular bones that conventionally 
have been called ‘“‘epipleurals’”’ in perci- 
form fishes. Measurements are given as per- 
centages of standard length (SL), except in- 
ternarial distance which is also presented as 
the quotient of the snout length divided by 
the distance between the nares. Osteological 
data were recorded from radiographs. Stan- 
dard methods for wax histology were used 
in preparing the gonadal sections for micro- 
scopic examination. 


Anthias Bloch 


Anthias Bloch, 1792 (type species Labrus 
anthias Linnaeus, 1758, by absolute tau- 
tonomy). 

Aylopon Rafinesque, 1810 (type species La- 
brus anthias Linnaeus, 1758, by virture 
of the fact that a replacement name re- 
tains the type of the prior name; Anthias 
incorrectly regarded as preoccupied by 
Anthia Weber, 1801, a genus of Coleop- 
tera). 


Diagnosis.—A genus of Anthiinae dis- 
tinguishable from all other genera of the 
family Serranidae by the following combi- 
nation of characters. No tooth plate on sec- 
ond epibranchial. Vertebrae 26 (10 precau- 
dal + 16 caudal). Formula for configuration 
of supraneural bones, anterior neural spines, 
and anterior dorsal pterygiophores 0/0/2/ 
1+1/1/ (using notation of Ahlstrom et al., 
1976), except A. nicholsi rarely with slight- 
ly different placement of supraneural bones. 
Principal caudal-fin rays 15 (8 + 7); 
branched rays 13 (7 + 6). Dorsal-fin rays 
X, 13 to 16 (usually X, 14 or 15, most fre- 


VOLUME 113, NUMBER 2 


quently X, 15). Anal-fin rays III, 6 to 8 
(usually III, 7). Pleural ribs on vertebrae 3 
through 10. Epineurals associated with first 
11 to 13 vertebrae. Scales ctenoid, with 
only marginal cteni (i.e., no ctenial bases 
present proximal to marginal cteni—see 
Hughes 1981; this type of scale called pe- 
ripheral ctenoid by Roberts 1993); no 
smaller accessory scales (secondary squa- 
mation) at bases of body scales. Most of 
head, including maxilla scaly. Lateral line 
complete, extending to at least base of cau- 
dal fin (running parallel to dorsal body con- 
tour a few scale rows ventral to dorsal-fin 
base, curving rather abruptly ventral to pos- 
terior end of dorsal-fin base to run near 
midlateral axis of body). No supramaxilla. 
Anterior and posterior nares closely set on 
each side of snout; posterior border of an- 
terior naris produced into a short flap, but 
never into a long filament. No fleshy papil- 
lae on border of orbit. Posterior margin of 
bony opercle with three spinous processes. 
Branchiostegal rays seven. Dorsal fin sin- 
gle, not deeply notched between spinous 
and soft portions. First caudal vertebra 
without parapophyses. Preopercle serrate, 
but without antrorse spines. Vomer and pal- 
atines with teeth; vomerine tooth patch 
without a well-developed posterior prolon- 
gation. 

Description.—Characters included in the 
generic diagnosis form a part of the generic 
description and are not repeated. Premax- 
illae protrusile. Posterodorsal border of 
maxilla not covered by elements of circu- 
morbital series when mouth closed. Outer 
teeth in jaws mostly conical; inner teeth 
mostly villiform or cardiform; some en- 
larged as canines. Endopterygoids usually 
toothless. Tongue usually without teeth (ex- 
cept present on tongue in almost all A. me- 
nezesi and A. cyprinoides). Pectoral fin ap- 
proximately symmetrical, with 16 to 22 
rays. Pelvic-fin rays I, 5. Gillrakers well de- 
veloped, total on first arch 37 to 48. Lateral- 
line scales 31 to 48. Caudal-peduncle scales 
16 to 25. No trisegmental pterygiophores 
associated with dorsal and anal fins. No 


av 


spur on posteriormost ventral procurrent ray 
(see Johnson 1975); penultimate ventral 
procurrent caudal-fin ray not shortened ba- 
sally. Parhypural with well-developed hy- 
purapophysis. Autogenous hypurals 5—no 
hypural fusions. Epurals 3. Uroneurals | 
pair (posterior pair absent). Modified scales 
(interpelvic process) overlapping pelvic-fin 
bases along midventral line. 

Those species of Anthias in which the 
larvae have been described lack several 
characters found in the larvae of some 
American anthiines, viz., specialized larval 
scales and serrae on supraoccipital crest, 
pterotic ridge, articular, frontal ridge, pari- 
etal ridge, and fin spines. In addition, the 
known larvae of Anthias have a single ser- 
rate supraorbital ridge dorsal to each eye, 
in contrast with the larvae of at least two 
species of American anthiines that have 
three serrate supraorbital ridges on each 
side (Baldwin 1990). 


Key to the Species of Anthias 


la. Lateral-line scales 31—34; sum of lat- 
eral-line scales plus total number of 
gillrakers on first gill arch, in individual 
specimens, 71-76; caudal-fin lobes 
moderate (length of upper lobe 31—49% 
ST). ae tee. Anthias nicholsi Firth, 1933 
(western North Atlantic) 

1b. Lateral-line scales 36—48; sum of lat- 
eral-line scales plus total number of 
gillrakers on first gill arch, in individual 
specimens, 75-88; caudal-fin lobes 
moderate to well produced (length of 
upper lobe 32—110% SL) 

2a. Longest dorsal-fin spine (usually the 
third) 13-30% SL, 19-30% SL in spec- 
imens more than ca. 100 mm SL; third 
dorsal-fin spine typically with well de- 
veloped filament which may be up to 
17% SL; lower caudal-fin lobe usually 
longer thamMipperctiink Vite os eee. 

Anthias anthias (Linnaeus, 1758) 
(eastern Atlantic, including the Medi- 
terranean and Adriatic seas) 

2b. Longest dorsal-fin spine 10—20% SL; 
fin membrane usually extending as a 
short filament at tip of each dorsal 


aae 


SD: 


4a. 


4b. 


Dae 


2).0); 


6a. 


6b. 


vey 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


spine, but never produced to the extent 
seen in A. anthias; upper caudal-fin 
lobe usually longer than lower ...... 3 
Soft dorsal-fin rays usually 14 (15 in 1 
of 17 specimens); midline of gular re- 
gion and lower jaw well covered with 
scales; pelvic-fin length 27-41% SL 
Anthias 
1980 


woodsi Anderson & Heemstra, 
(western North Atlantic) 

Soft dorsal-fin rays usually 15 (rarely 
16); gular region naked; lower jaw na- 
ked or only partly covered with scales; 
pelvic-fin length 33-76% SL 
Pectoral-fin rays 19—21 (usually 20 or 
21); longest dorsal-fin spine 10-13% 
SL; upper caudal-fin lobe 32—44% SL; 
pelvic-fin length 33-39% SL; known 
only from the eastern South Atlantic SS) 
Pectoral-fin rays 17—20 (usually 18 or 
19); longest dorsal-fin spine 12—20% 
SL; upper caudal-fin lobe 39-—>70% 
SL; pelvic-fin length 33-76% SL; not 
known from the eastern South Atlantic 


pas vps argh a Are Ce Rae ats eR ONE eh 2, lees as 6 
Caudal-peduncle scales ca. 18; posteri- 
or margin of anal fin rounded; upper 
caudal-fin lobe 37-44% SL; no teeth on 
tonPUesie eRe Teale tens: Anthias 
helenensis Katayama & Amaoka, 1986 


(eastern South Atlantic north of the Is- 
land of St. Helena) 

Caudal-peduncle scales 20—24; poste- 
rior margin of anal fin angular; upper 
caudal-fin lobe 32-37% SL; tongue 
usually with one or two small patches 

Of ME Ctlayes.c4-42 ete Fae Sede Anthias 
cyprinoides Katayama & Amaoka, 1986 
(eastern South Atlantic west of the Is- 
land of Pagalu) 

Total gillrakers on first gill arch 41—48; 
tongue usually with teeth, teeth usually 

in narrow elongated patch ... Anthias 
menezesi Anderson & Heemstra, 1980 
(western South Atlantic) 

Total gillrakers on first gill arch 37-41; 
tongue usually without teeth 
Soft dorsal fin and usually soft anal fin 
without produced rays; caudal-peduncle 
scales 17 or 18; lateral-line scales 36— 
41; two of largest individuals examined 
(out of the 10 known specimens) with 
teeth on tongue 


O:) Bi 56) (6) te) 8: (0;)" enue) 460 10) 26) Ke. (ever ie! jelnle 


ue Anthias asperilinguis Giinther, 1859 

(western North Atlantic) 

7b. Two or more soft dorsal-fin rays and 
one or more soft anal-fin rays pro- 
duced; caudal-peduncle scales 22-25; 
lateral-line scales 38—46; no teeth on 
tongue Anthias noeli, new species 
(Galapagos Islands, eastern Pacific) 


4 je ie je Je) <0 


Anthias noeli, new species 
Rosy Jewelfish 
Figs. 1—5; Tables 1, 2 


Material examined.—Seventeen speci- 
mens, 62.1 to 173 mm SL; all collected off 
the Galapagos Islands in the eastern Pacific 
Ocean in depths of 184 to 351 m. 

Holotype: USNM 353113 (167 mm SL, 
d); seamount SE of Isla San Cristobal— 
01°06.48'S, 89°06.70'W; 202 m; Johnson- 
Sea-Link-I dive no. 3937; Gilmore & San- 
tos, 6 Nov 1995. 

Paratypes: USNM 351335 (2, 127-150; 
®, 6); off NE Usla” Darwin=01-42 ne 
92°00’ W; 351 m; Johnson-Sea-Link-II dive 
no. 3103; McCosker & Pawson, 18 Jul 
1998. CAS 86573 (1, 132; 3); off Isla Dar- 
win—01°41.39'’N, 91°58.88'W; 335 m:; 
Johnson-Sea-Link-I dive no. 3963; Mc- 
Cosker et al., 20 Nov 1995. USNM 351334 
(2, 86-110; 3, 6); off Isla Marchena— 
00°24’'N, 90°26.3'W; 303 m; Johnson-Sea- 
Link-IT dive no. 3109; Baldwin & Mc- 
Cosker, 21 Jul 1998. GMBL 95-34 (1, 139; 
2°) & HBOM 107:08471 (1, 123; 2); off 
Cabo Douglas, Isla Fernandina—00°17.60'S, 
91°39.00'W; 299 m; Johnson-Sea-Link-I 
dive no. 3956; Robison & Santos, 16 Nov 
1995. CAS 201896 (1, 173; 3); off Isla Pla- 
zas (=Plaza Sur in Fig. 2)—00°31'24'S, 
90°09'0"W; 308 m; Johnson-Sea-Link-II 
dive no. 3096; McCosker & Day, 7 Jul 
1998. USNM 351333 (2, 132-170; ?, 3); 2 
miles E of Isla Plazas (=Plaza Sur in Fig. 
2)—00°32.25'S, 90°09.02’W; 308 m; John- 
son-Sea-Link-II dive no. 3096; McCosker 
& Day, 7 Jul 1998. CAS 86807 (1, 163; 
6); seamount SE of Isla San Cristobal— 
01°06.03'S, 89°12.20'W; Johnson-Sea- 


VOLUME 113, NUMBER 2 


373 


Table 1.—Data on morphometric characters for Anthias noeli. Standard lengths are in mm; other measure- 


ments, in percentages of standard length. 


Character n Min Max. Mean Holotype 
Standard length 1? 62.1 173 —— 167 
Head, length 17 36.8 43.4 39.4 37.6 
Snout, length le 6.0 a2 Iso 8.4 
Orbit, diameter 17. el 15.1 135 11.4 
Postorbital length of head 17 17.0 20.4 18.8 18.4 
Upper jaw, length 17 17.0 19.0 17.9 18.2 
Maxilla, width Wi 5.9 7.4 6.6 7.0 
Bony interorbital, width Li? 7.6 oJ 8.3 8.6 
Internarial distance 17 0.7 LZ 1.0 0.9 
Body, depth 17 35.0 42.1 38.5 40.4 
Body, width 17 |e | He 15.9 17.6 
Predorsal length We 31.8 36.6 34.1 55:7 
Prepelvic length LZ 35:7 43.7 39.1 38.2 
Preanal length 17 59.4 66.3 63.3 64.0 
Caudal peduncle, length ile 21.9 26.7 24.6 20:0 
Caudal peduncle, depth 17 11.8 13.6 12.6 13.6 
Dorsal-fin base, length 7, 54.4 le), 570 61.7 
Pectoral fin, length 17 Deo 33.4 31.2 31.4 
Pelvic fin, length 18 32.6 =p 42.7 53.7 
Anal-fin base, length 17 16.1 18.5 te 18.2 
Anal fin, length £5 323 42.9 37.0 41.9 
Upper caudal-fin lobe, length ye 38.8 60.4 49.2 46.9 
Lower caudal-fin lobe, length |e) 33.2 57.4 46.8 46.3 
First dorsal spine, length 14 6.1 8.5 7.0 7.4 
Third dorsal spine, length LS 13s 17.8 153 14.9 
Fourth dorsal spine, length it 14.3 19:9 16.6 14.7 
Tenth dorsal spine, length is) 10:7 12.6 8 Wea Lee 
Longest dorsal spine, length 13 14.3 ee) 16.5 L522 
Longest dorsal soft ray, length 11 26.1 44.7 36.0 33.3 
Pelvic spine, length 15 15.4 19.8 17-3 5 lf | 
First anal spine, length 16 Gad 10.4 8.2 ee | 
Second anal spine, length 14 12.0 20.4 15.6 14.4 
Third anal spine, length 16 12.8 gre 14.7 12.8 
Longest anal soft ray, length 15 24.0 31:8 27.4 31.8 


Link-I dive no. 3934; McCosker et al., 5 
Nov 1995. UF 110990 (2, 68-134; 2, 9); 
seamount SE of Isla San Cristobal— 
01°06.23'S, 89°06.91'W; 184-215 m; John- 
son-Sea-Link-I dive no. 3935; Robison & 
Liberatore, 5 Nov 1995. ANSP 177770 (3 
specimens, 62.1-106 mm SL; ?, 2, 3); off 
Devil’s Crown, Isla Floreana—01°12.50’S, 
90°25.56’W; 232 m; Johnson-Sea-Link-I 
dive no. 3945; McCosker & Liberatore, 9 
Nov 1995. 

Diagnosis.—A species of Anthias distin- 
guishable from all other species of the ge- 
nus by the following combination of char- 
acters. Dorsal-fin rays X, 15 or 16 (usually 


X, 15). Anal-fin rays II], 6 or 7 (usually 
III, 7). Pectoral-fin rays 18 to 20 (most fre- 
quently 19). Lateral-line scales 38 to 46 
(usually 39—44). Total gillrakers on first 
gill arch 37 to 41 (usually 38—40; no ru- 
dimentary rakers). Sum of lateral-line 
scales plus total number of gillrakers, on 
individual specimens, 78 to 85. Caudal-pe- 
duncle scales 22 to 25 (usually 23 or 24). 
Caudal fin lunate to deeply forked (larger 
individuals tending to have more lunate 
fins). Dorsal, anal, pelvic, and caudal fins 
each with some produced soft rays. Upper 
caudal-fin lobe longer than lower. Anal fin 
angulated posteriorly. Gular region without 


374 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Table 2.—Frequency distributions of numbers of fin rays, scales, and gillrakers on first gill arch in Anthias 


noeli. Counts for the holotype are indicated by asterisks. 


Character 15 16 6 


Dorsal soft rays Gs ] 
Anal soft rays 
Pectoral-fin rays, 


Left side 
Right side 


38 39 40 41 


Tubed lateral-line scales, 


Left side l 
Right side l 


i) 
Nw NY 
£ 


Circum caudal-peduncle scales l 


10 1] 12 


Gillrakers 


Upper limb 4 La 
Lower limb 


37 38 


Total gillrakers 


(upper limb + lower limb) 6 


scales. Endopterygoids and tongue without 
teeth. 

Description.—Characters presented in 
the generic and species diagnoses and the 
generic description form part of the species 
description and are not repeated unless nec- 
essary for clarification. Data for morpho- 
metric characters appear in Table 1; those 
for some countable characters in Table 2. 
Mouth nearly terminal, lower jaw exceed- 
ing upper jaw very slightly. Maxilla falling 
short of vertical through posterior border of 
orbit. Maxilla usually widened distally, 
with a small shelf or rostrally directed hook 
on labial border (as in Anatolanthias api- 
omycter—see Anderson et al. 1990:926, fig. 
2). Internarial distance 6—12, usually 7—10, 
times in snout length. Vertical limb of preo- 
percle serrate; horizontal limb smooth or 
with several small serrae near angle; angle 
with a single spine, a serrated spinous pro- 
cess, or relatively large serrae. Distal mar- 
gins of interopercle and subopercle usually 


Mean 


oT, 18 19 20 Mean 
15.06 
i Se 6.94 
ik 14* 2 19.06 
2. 10 qr 9.12 
43 44 45 46 Mean 
4 3%* — 1 41.82 
4* 1 4 1 42.06 
23 24 DS Mean 
i idl y) 2355 
27 28 29 30 Mean 
3) ih 4 1 28.06 
39 40 41 Mean 
5 4 l 38.88 


smooth or nearly so, occasionally with sev- 
eral small serrae. 

Premaxilla with inner band of very small 
teeth and outer series of larger, mostly an- 
teriorly directed, conical teeth; near sym- 
physis, two to several teeth along medial 
margin of inner band enlarged as posteri- 
orly directed conical teeth; outer row of 
larger conical teeth usually preceded by one 
or two enlarged canine teeth. Dentary with 
row of slender conical teeth along lateral 
edge of jaw; this row including one to three 
teeth, usually enlarged into recurved ca- 
nines, at a point approximately one third 
length of row from its anterior end; band of 
very small teeth extending anteriorly from 
this row and reaching to near symphysis; 
one to several teeth on inner edge of band 
near symphysis enlarged and directed pos- 
teriorly; one or two enlarged exserted ca- 
nines near symphysis. 

Branchiostegal rays seven—anterior 
three inserting along ventral edge or ven- 


VOLUME 113, NUMBER 2 


Pip. 1. 
Pacific Ocean. Photograph by Donald Hurlbert, National Museum of Natural History, Smithsonian Institution. 


tromedial aspect of hyoid arch, posterior 
four inserting on lateral surface of arch. 
Pseudobranch well developed, with ca. 22 
to 33 filaments; number of filaments in- 
creasing with increase in SL (No. filaments 
= a+ b[SL]; where a = 17.03, b = 0.08, 
f= ©7522). 

Most of head, including most of dorsum 
of snout, interorbital region, and maxilla, 
scaly. Lateral aspect of snout, lachrymal, 
branchiostegals, branchiostegal membranes, 
and gular region naked (one specimen with 
a few scales on lachrymals). Dentary usu- 
ally naked, but some specimens with a few 
scales posteriorly. Rows of scales on cheek 
8 to 11 (usually 9 or 10; difficult to count). 
Dorsal and anal fins mostly scaleless, ex- 
cept columns of very small scales frequent- 
ly present proximally on interradial mem- 
branes between soft rays, particularly on 
larger specimens. Rows of scales between 
lateral line and mid-base of spinous dorsal 
fin 2 or 3. Scales between origin of dorsal 


Holotype of Anthias noeli, new species, USNM 353113, 167 mm SL; Galapagos Islands, eastern 


fin and lateral line 6 to 8. Scales between 
origin of anal fin and lateral line 17 to 20. 

Longest dorsal-fin spine fourth or fifth. 
Second anal-fin spine more robust than first 
or third. Procurrent caudal-fin rays 9 (rarely 
10) dorsally, 8 or 9 ventrally. Dorsal-fin 
membrane produced into a short filament 
posterior to each spine. Two or more soft 
dorsal-fin rays, one or more soft anal-fin 
rays, first two (and on occasion third) soft 
pelvic-fin rays (second longest), and cau- 
dal-fin lobes produced. Pectoral fin reach- 
ing posteriorly to at least as far as vertical 
through base of third anal spine, frequently 
as far as vertical through base of second 
anal soft ray or beyond; dorsalmost pecto- 
ral-fin ray always unbranched, ray next to 
dorsalmost ray and ventralmost ray occa- 
sionally unbranched. Pelvic fin reaching 
posteriorly at least to base of first anal soft 
ray to as far as well beyond posterior end 
of anal-fin base. Pelvic fin inserted beneath 
anterior part of pectoral-fin base. 


376 


oOo | 
«Darwin 
Wolf 
(\ Pinta 
\ 
0° S 


San Salvador 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


O 100 km 


QGenovesa 


Bartolome 
o : : Seymour 
ernandina “| Rdbida* Daphne. )\ Baltra 
ok) 
Pinzon Plaza Sur 
Oo 
lsabela Sy 


oe AOS 


Pie ’ Hermanos 
O° 


Tortuga 


Santa Fe 
San Cristobal 
] 
& _ Champion 
Floreano/ . Gardner 
Gardner 
Espanola 


Figs 2. 
species. Modified from Grant 1984. 


Coloration.—Description based on color 
photographs, taken shortly after capture, of 
two paratypes (CAS 201896, 173 mm SL; 
USNM 351333, 170 mm SL; see Fiesta); 
and color notes, made by the second author 
in the field, of two other paratypes (USNM 
351335, 127 & 150 mm SL). Head mostly 
rosy, a yellow streak extending across lach- 


Map of Galapagos Islands, eastern Pacific Ocean, showing localities of capture of Anthias noeli, new 


rymal and part of cheek and a broad yellow 
stripe extending from posterior margin of 
eye to posterior tip of opercle. Jaws rosy 
except small patch of yellow on upper lip 
near premaxillary symphysis. Iris yellow. 
Body mostly rosy dorsally, paler ventrally, 
a few yellow stripes or blotches on lateral 
and ventral aspects of body; black blotch 


VOLUME 113, NUMBER 2 


present at anterior base of spinous dorsal 
fin. Membrane covering dorsal-fin spines 
yellow except rosy distally; interradial 
membranes mostly rosy; soft dorsal fin 
mostly rosy, except membranes separating 
three anteriormost dorsal soft rays mostly 
yellow or yellow distally and last four rays 
pale purple. Anal-fin spines and interradial 
membranes yellow; anterior soft anal-fin 
rays and interradial membranes yellow ba- 
sally, yellow and rosy distally; posterior 
portion of soft anal fin rosy to pale purple. 
Pectoral fin rosy. Pelvic fin mostly pale 
rose, some yellow basally and on mem- 
brane between first and second rays. Caudal 
fin mostly rosy, some yellow on outer rays 
of dorsal and ventral lobes; produced distal 
ends of caudal-fin lobes rosy or lilac to pur- 
plish in color. 

Coloration in alcohol.—Dark spot ante- 
rior to base of spinous dorsal fin usually 
persisting; dorsum of head frequently 


3T7 


dusky; fins mostly pallid; remainder of fish 
straw-colored. 

Sexuality.—Histological examination of 
the gonads of all specimens of the new spe- 
cies (except for two for which no gonadal 
tissue could be found) showed that six in- 
dividuals (68.0—-139 mm SL) are females 
and nine (86.0—173 mm SL), including the 
five largest (150-173 mm SL), are males, 
suggesting that Anthias noeli is protogyn- 
ous (see section on hermaphroditism in an- 
thiines). 

Sexual dimorphism.—In specimens more 
than about 120 mm SL, females (4 speci- 
mens, 123—139 mm SL) have shorter pelvic 
fins (33-36% SL vs. 43-55% SL), shorter 
longest dorsal soft rays (31-33% SL vs. 
38-45% SL), shorter longest anal soft rays 
(24-28% SL vs. 28-32% SL), and shorter 
depressed anal-fin lengths (33-37% SL vs. 
37—42% SL) than do males (5 specimens, 
150-173 mm SL). 


Fig: 3: 
Pacific Ocean. 


Paratype of Anthias noeli, new species, USNM 351333, 170 mm SL; Galapagos Islands, eastern 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 4. 
with only marginal cteni, as in species of Anthias; B, 
marginal cteni. 


Comparisons.—Anthias noeli is separa- 
ble from other species of Anthias as fol- 
lows: longest dorsal-fin spine fourth or fifth, 
third dorsal-fin spine with at most a short 
filament, and upper caudal-fin lobe longer 
than lower caudal-fin lobe in A. noeli versus 
longest dorsal-fin spine usually third, third 
dorsal-fin spine typically with well devel- 
oped filament (up to 17% SL), and lower 
caudal-fin lobe usually longer than upper 
caudal-fin lobe in A. anthias; two or more 
soft dorsal-fin rays and one or more soft 
anal-fin rays produced and 22 to 25 caudal- 
peduncle scales in A. noeli versus soft dor- 
sal fin and usually soft anal fin without pro- 
duced rays and 17 or 18 caudal-peduncle 
scales in A. asperilinguis; upper caudal-fin 
lobe length 39 to 60% SL, 18 to 20 (usually 
19) pectoral-fin rays, and tongue without 
teeth in A. noeli versus upper caudal-fin 
lobe length 32 to 37% SE, 20: or 21 pec- 
toral-fin rays, and tongue usually with teeth 
in A. cyprinoides; 22 to 25 caudal-peduncle 
scales and anal fin angulated posteriorly in 
A. noeli versus ca. 18 caudal-peduncle 
scales and anal fin rounded posteriorly in A. 
helenensis; 37 to 41 total gillrakers on first 
gill arch and tongue without teeth in A. noe- 
li versus 41 to 48 total gillrakers on first 
gill arch and tongue usually with teeth in 
A. menezesi; 38 to 46 lateral-line scales in 
A. noeli versus 31 to 34 in A. nicholsi; 15 
or 16 (usually 15) soft dorsal-fin rays and 
gular region and dentary without scales 
(some specimens with a few scales on den- 


Scales of anthiine fishes (posterior fields of scales towards the top of the page). A, ctenoid scale 


ctenoid scale with ctenial bases present proximal to 


tary posteriorly) in A. noeli versus 14 or 15 
(usually 14) soft dorsal-fin rays and midline 
of gular region and dentary well covered 
with scales in A. woodsi. 

In addition, Anthias noeli is distinguish- 
able from the other species of Anthiinae 
found in the eastern Pacific by the follow- 
ing combination of characters: scales cte- 
noid, with only marginal cteni (1.e., no cten- 
ial bases present proximal to marginal cteni; 
see Fig. 4A); maxilla scaly; dentary naked 
or with only a few scales posteriorly; gular 
region without scales; vomer with teeth; no 
teeth on tongue; anterior naris relatively 
close to posterior naris (internarial distance 
6—12, usually 7—10, times in snout length); 
posterior border of anterior naris produced 
into a short flap, but never into a long fil- 
ament; preopercle without antrorse spines; 
urohyal without anteriorly projecting spine; 
dorsal-fin rays X, 15 or 16 (usually X, 15); 
anal-fin rays III, 6 or 7 (usually III, 7); lat- 
eral-line scales 38 to 46; total gillrakers on 
first gill arch 37 to 41; diameter of bony 
orbit 11 to 15% SL. The most recent com- 
parative treatment of the adults of eastern 
Pacific anthiines is that of Fitch (1982), 
which covers only four of those species, all 
found in the northern hemisphere. To facil- 
itate identification of all 13 species of east- 
ern Pacific Anthiinae, we present a key to 
those taxa (see below). 

Distribution.—Anthias noeli is known 
only from the Galapagos Islands, eastern 


VOLUME 113, NUMBER 2 


Pacific Ocean, in depths ranging from 184 
(0/351. meters (see Fig. (2): 

Etymology.—The specific name, noeli, is 
for Noel Archambault, IMAX® cameraman/ 
stereographer, who lost his life on 26 June 
1998 in a tragic plane crash in the Gala- 
pagos during one of the expeditions on 
which the new species was collected. Noel 
was a pioneer of modern 3-D film technol- 
ogy. It is appropriate to name in his mem- 
ory a new species collected using the mod- 
ern submersible technology that is allowing 
the exploration of oceanic regions previ- 
ously inaccessible to scientific study. 


Key to the Eastern Pacific Species of 
Anthiinae 


la. Scales ctenoid, with only marginal 
cteni (i.e., no ctenial bases present 
proximal to marginal cteni; Fig. 4A) 


co lB gi A i ar ea ara 2 
1b. Scales ctenoid with ctenial bases pre- 
sent proximal to marginal cteni (Fig. 
4B) or in one species (Trachypoma 
macracanthus) scales cycloid ..... 1 
ameeeasilascaly 220.0. Re es 3 
Parasillanaked) 2. cae ee eee 5 


3a. Vomer edentate; anterior naris some- 
what remote from posterior naris; in- 
ternarial distance 2.8—3.1 times in 
snout length .. Anatolanthias apiomycter 
Anderson, Parin, & Randall, 1990 
(Nazca Ridge) 

3b. Vomerine dentition well developed; 
anterior naris relatively close to pos- 
terior naris; internarial distance more 
than 5.0 times in snout length 

4a. Posterior border of anterior naris pro- 
duced into a filament, usually reaching 
posterior naris when reflected; ventral 
surface of dentary and midline of gu- 
lar region with scales; bony orbit di- 
ameter 7—11% SL (8-11% SL in spec- 
imens <160 mm SL, 7-9% SL in 
specimens >165 mm SL); teeth usu- 
ay present Of tomeuc .. 7... .......- 

Pronotogrammus multifasciatus 
Gill, 1863 (southern California and 
the Gulf of California to northern Peru 
and the Galapagos Islands) 

4b. Posterior border of anterior naris with- 


© * © © » w 


5a: 


ab: 


6a. 


6b. 


Ta 


7b 


out a filament, but produced into a 
short flap that almost always fails to 
reach posterior naris when reflected; 
dentary usually without scales, occa- 
sionally with a few scales posteriorly; 
gular region without scales; bony orbit 
diameter 11-15% SL (12-15% SL in 
specimens <140 mm SL, 11-13% SL 
in specimens >150 mm SL); no teeth 
on tongue ... 
(Galapagos Islands) 

Lateral-line scales 60—71 (usually 62-— 
68); specimens more than about 70 mm 
SL with sharp spine projecting anteri- 
orly from ventral border of urohyal 


379 


Anthias noeli, new species 


. Hemanthias signifer (Garman, 1899) 


(southern California to northern Peru) 
Lateral-line scales 36-57 (usually 54 
or fewer); urohyal without anteriorly 
PLOjeCline Spine ii yt .4% eels. 2S 

Lateral-line scales 36—40; total gill- 
rakers on first arch 38—43; longest 
dorsal-fin spine most frequently the 
sixth, never the third; third dorsal-fin 
spine with short filament, but filament 
never well produced; middle rays of 
upper and lower lobes of caudal fin 
not longest in fin 


we Se je eo) Vay car a Le lelve “a ef “ate 


Sle “er elise vey eras el Oe se) mt yer ce 


Gilbert, 1890 (mid-Gulf of California 
[28°N; Fitch 1982] to Colombia 
[6.5°N]) 

Lateral-line scales 49-57; total gill- 
rakers on first arch 31-35; longest 
dorsal-fin spine—the third; third dor- 
sal-fin spine with well produced fila- 
ment (at least in larger individuals); 
middle rays of upper and lower lobes 
of caudal fin longest in fin 


a eps a @ el iste) ee 6 oe ees 


(Steindachner, 1875) (Hipolito Bank 
[27°N], Baja California Sur, in the Pa- 
cific and Cabo Lobos [ca. 30°N], So- 
nora, in the Gulf of California to Tru- 
jillo [8°S], Peru, and the Galapagos Is- 
lands [Fitch 1982; Grove & Laven- 
berg 1997]) 

Spines in dorsal fin XI-XIII; ventral 
border of preopercle with strong an- 
trorse spines 
Spines in dorsal fin X; ventral border 
of preopercle without antrorse spines, 
except in Hypoplectrodes semicinctum 


Sa ee ts) eee ee ee a eee a Se oe ee 


Pronotogrammus eos 


Hemanthias peruanus 


380 


8a. 


8b. 


9a. 


Ob. 


10a. 
10b. 
iia: 


1 1b. 


ia: 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


and occasionally in Plectranthias exsul 
eel etegte Sec ieee. tee 10 
Scales cycloid; spines in dorsal fin 
XII; soft rays in anal fin 6 
Trachypoma macracanthus 
Gunther, 1859 (widely distributed in 
the South Pacific, including Easter Is- 
land and the Desventuradas Islands off 
the coast of Chile [Randall & Cea 
Egana 1984; Pequefo & Lamilla 
1996a,b; Rojas & Pequefio 1998a]) 
Scales ctenoid; dorsal spines XI or 
XIII; soft rays in anal fin 8 or 9 ... o 
Dorsal-fin rays XI, 17 or 18 
dee Acanthistius pictus (Tschudi, 1846) 
(coasts of Peru and Chile) 
Dorsal-fin rays XIII, 14 or 15 ...... 
Pattee: Acanthistius fuscus (Regan, 1913) 
(Easter Island and Sala y Gémez Is- 
land [Rojas & Pequeno 1998a]) 
Soft rays in dorsal fin 19-22 ..... it 
Soft. rays m-dorsal’ fin: 1Sror 16242). 12 
Preopercle with 2 or 3 antrorse spines 
(one spine usually at angle or on ven- 
tral margin near angle, other spine[s] 
on ventral margin); total gillrakers on 
first gill arch 17—20; lateral-line scales 
48-55 (most frequently 48-51) ... 
Hypoplectrodes semicinctum 
(Valenciennes, 1833) (Juan Fernandez 
Islands and San Félix Island [in the 
Desventuradas Islands] off the coast 
of Chile, and possibly Easter Island 
[Anderson & Heemstra 1989; Pe- 
queno & Lamilla 1996b; Rojas & Pe- 
queno 1998a]) 
Preopercle serrate, but without an- 
trorse spines; total gillrakers on first 
gill arch 34—37; lateral-line scales 58— 
65, usually 61—64 (counts only from 
eastemPacilic specimens) =. 4. 
.. Caprodon longimanus (Gunther, 1859) 
(widely distributed in the Pacific—in- 
cluding Easter Island and the Nazca 
Ridge and from the Desventuradas 
and Juan Fernandez islands off the 
coast of Chile, and ‘‘along the coast 
of South America”’ [Pequefo & Lam- 
illa 1996a:931, 1996b]) 
Scales on dorsum of head not extend- 
ing anterior to nares (Randall 1996: 
117); longest soft dorsal-fin ray ca. 
26-35% SL; total gillrakers on first 


©. ©) ue) je7ne) lel (07a) 0 ici fe 


eye) 6. mei ©) e; ‘ei: (eo; (67 fe 


gill arch 26—28; in life, two orange- 
red bars on body (one just anterior to 
anal fin, the other terminating ventral- 
ly posteriok toyanal ini a: 5 
Plectranthias parini 
(Anderson & Randall, 1991) (Sala y 
Gomez Ridge and Easter Island 
[Randall 1996]) 

Scales on dorsum of head extending 
anteriorly almost to upper lip, except 

for triangular premaxillary groove 
(Randall 1996:117); longest soft dor- 
sal-fin ray ca. 16—>26% SL; total gill- 
rakers on first arch 26—31 (usually 28 

or 29); in life, brilliant red oblong area 
extending from bases of posterior rays 

of soft dorsal fin ventrally to just below 
midline and then posteriorly over mid- 

dle of caudal peduncle to reach mid- 
ventral line near base of caudal fin .. 
PASUS Midi, MERCER So OE Plectranthias exsul 
Heemstra & Anderson, 1983 (Juan 
Fernandez Islands and Nazca Ridge; 
also reported from Desventuradas Is- 
lands by Rojas & Pequeno 1998a) 


oe) "ee se Je (ol is, ie: co [0 jo, Se 0:0) 0 


1b: 


Plectranthias lamillai 


Rojas & Pequeno (1998b) described a 
new species of Plectranthias, P. lamillai, 
from a single specimen (MNHNC P7055, 
139.6 mm SL) collected off Alejandro Sel- 
kirk Island in the Juan Fernandez Islands 
off the coast of Chile. In the description 
provided by Rojas & Pequefio (1998b), we 
could find no characters that seem to reli- 
ably distinguish it from Plectranthias exsul, 
also known from off the Juan Fernandez Is- 
lands. As a consequence, the first author 
made a detailed examination of the holo- 
type of P. lamillai (now 136 mm SL) and 
compared it closely with two specimens of 
P. exsul (USNM 176577—a paratype, 158 
mm SL, and USNM 312927, 122 mm SL) 
and found that the holotype agrees well 
with the original description of P. exsul 
and, with one exception, with the speci- 
mens of P. exsul with which it was directly 
compared. The interorbital region of the 
paratype of P. exsul examined in compari- 
son is more flattened than that of the ho- 


VOLUME 113, NUMBER 2 


lotype of P. lamillai; this we attribute to 
ontogenetic differences or perhaps to sexual 
dimorphism. In contrast, the 122-mm spec- 
imen of P. exsul is essentially identical in 
head and body shape to the holotype of P. 
lamillai. It should be noted that the drawing 
with the original description of P. lamillai 
(see Rojas & Pequeno 1998b:fig 2) is not a 
particularly good representation of the ho- 
lotype, especially of the head. Plectranthias 
lamillai and P. exsul have somewhat dif- 
ferent patterns of coloration, but we think 
that the different patterns displayed are best 
interpreted as variations on a common 
theme. In view of the preceding and pend- 
ing a more complete investigation, we deem 
it best to consider P. lamillai Rojas & Pe- 
queno, 1998, as a junior synonym of P. 
exsul Heemstra & Anderson, 1983. 


Comments on Hermaphroditism in 
Anthiine Serranids 


Protogyny has been reported in species 
representing a number of anthiine genera, 
including Hypoplectrodes (H. huntii [as El- 
lerkeldia huntii] and H. maccullochi; Jones 
1980 and Webb & Kingsford 1992, respec- 
tively), Hemanthias (H. vivanus and H. per- 
uanus;,; Hastings 1981 and Coleman 1983, 
respectively), Pronotogrammus (P. martin- 
icensis [as Holanthias martinicensis|; Co- 
leman 1981), Pseudanthias (P. squamipin- 
nis [as Anthias squamipinnis|; Fishelson 
1970 and Shapiro in a series of studies on 
the behavioral aspects of sex reversal—e.g., 
Shapiro 1986), Sacura (S. margaritacea; 
Reinboth 1963), and Anthias (A. anthias; 
Reinboth 1964). In addition, Heemstra 
(1973) provided evidence for protogyny in 
Pseudanthias conspicuus (as Anthias con- 
spicuus), and Robins & Starck (1961) stated 
that Plectranthias garrupellus is probably 
protogynous. Thresher (1984) presented a 
summary (current through about 1981) of 
the information available on the reproduc- 
tive biology of anthiines. 

One might assume, from the publications 
on anthiines cited above, that all of them 


381 


are protogynous and monandric. Similarly, 
based on a number of studies, groupers 
(Serranidae: Epinephelinae), in general, 
have been considered to display monandric 
protogyny, but Sadovy & Colin (1995:961) 
found that sexuality in Epinephelus striatus 
(the Nassau grouper) “‘is essentially gono- 
choristic, with potential for sex change” 
and that “‘the juveniles pass through a bi- 
sexual stage of gonadal development,”’ thus 
illustrating the importance of not making a 
priori assumptions about the reproductive 
biology of serranid fishes. In a study of the 
Serranidae of the eastern Gulf of Mexico, 
Bullock & Smith (1991) presented evidence 
for protogyny in the anthiine Hemanthias 
leptus, but, based on finding a ripening 
male of 86 mm SL (p. 21, fig. 8b) and a 
sexually mature male of 61 mm SL (p. 207, 
pl. Id), entertained the idea that H. leptus 
may be diandric, acknowledging, however, 
that additional study is needed to confirm 
this. 

In the subsection entitled Sexuality (un- 
der Anthias noeli, new species), we have 
presented evidence suggesting that Anthias 
noeli is protogynous. To gather a better ap- 
preciation of sexuality in the genus Anthias, 
William A. Roumillat, at our request, ex- 
amined histological sections of the gonads 
of 20 specimens of the western Atlantic A. 
nicholsi and found 12 females (52.0—125 
mm SL), one individual (73.0 mm SL) 
transforming from female to male, and sev- 
en secondary males (99.9-134 mm SL). 
These data strongly suggest that Anthias ni- 
cholsi is protogynous. Anthias noeli dis- 
plays patterns of gonadal morphology sim- 
ilar to those seen in A. nicholsi (Fig. 5), 
lending further support to our contention 
that A. noeli is probably protogynous. 

Sadovy & Shapiro (1987) gave criteria 
for diagnosing various types of hermaph- 
roditism in fishes. Features that they iden- 
tified as strongly indicating protogyny are: 
‘*membrane-lined central cavities in testes; 
transitional individuals; atretic bodies in 
stages 1, 2, or 3 of oocytic atresia within 
testes; sperm sinuses in the gonadal wall; 


eS) 
(o.2) 
N 


ees 


R% 


‘oe 


Rig. : 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Gonadal sections of specimens of Anthias stained with hematoxylin and eosin-Y. A, A. noeli, new 


species, HBOM 107:08471, 123 mm SL, mature female; B, A. noeli, new species, CAS 201896, 173 mm SL, 
secondary male; C, A. noeli new species, CAS 86573, 132 mm SL, secondary male; D, A. nicholsi, GMBL 59- 
32, 73.0 mm SL, individual transforming from female to male. Abbreviations: AR = artifact; AO = atretic 
oocyte; D = dorsal; M = medial; OL = ovarian lumen; OO = oocyte; SS = sperm sinus; TT = testicular 
tissue. Photomicrographs by William A. Roumillat, digital imagery by Karen Swanson, South Carolina Depart- 


ment of Natural Resources, Charleston. 


and experimental production of transitional 
or sex-reversed individuals through manip- 
ulation of the social system’? (Sadovy & 
Shapiro 1987:150). In our studies we found, 
with the assistance of Bill Roumillat, that 
Anthias nicholsi fulfills all of those criteria, 
except the last one which requires live ma- 
terial, and that A. noeli meets all but the 
last and the one involving transitional in- 


dividuals (sensu Sadovy & Shapiro 1987: 
147-148) which were not observed. We 
feel that it is reasonable to conclude that 
both species are protogynous. 


Acknowledgments 


We appreciate the cooperation of officials 
of the Instituto Nacional de Pesca, the 


VOLUME 113, NUMBER 2 


Parque Nacional Galapagos, and the Esta- 
cidn Cientifica Charles Darwin for allowing 
the study and capture of fishes, and we ac- 
knowledge the support in the field of Al 
Giddings of Images Unlimited, David Clark 
of David Clark, Inc., the Discovery Chan- 
nel, Mandalay Media Arts, LLC, IMAX 
Corporation, and the Smithsonian Institu- 
tion. David Catania, R. Grant Gilmore, Jr., 
and John E. McCosker provided specimens 
and collection data, and Gilmore and 
McCosker allowed us to deposit material of 
the new species in several museums. Rob- 
erto Meléndez very kindly sent the holo- 
type of Plectranthias lamillai on loan. Wil- 
liam A. Roumillat examined and interpreted 
histological sections of the gonads of two 
species of Anthias; John McCosker lent us 
color transparencies of the new species; 
Yvonne Sadovy called our attention to an 
important reference; and Antony S. Harold 
and Dorian R. McMillan assisted in a num- 
ber of ways. Special thanks are due to John- 
son-Sea-Link submersible pilots Don Lib- 
eratore and Phil Santos (Harbor Branch 
Oceanographic Institution), whose expertise 
in maneuvering the submersibles and their 
collection gear was instrumental in collect- 
ing the specimens of the new species. The 
black and white photograph of the holotype 
of Anthias noeli (Fig. 1) was made by Don- 
ald Hurlbert; photomicrographs of the go- 
nadal sections of A. noeli and A. nicholsi 
(Fig. 5) were provided by W. A. Roumillat, 
who also commented upon the section on 
hermaphroditism in anthiines. Karen Swan- 
son produced the color plate (Fig. 5) of the 
gonadal tissues from photomicrographic 
transparencies. Support for the publication 
of the color plates (Figs. 3 & 5) was pro- 
vided by IMAX Corporation, Mandalay 
Media Arts, LLC, and the National Muse- 
um of Natural History’s Office of Public 
Programs. John McCosker and Sarah 
Riseman read the entire manuscript and of- 
fered suggestions for its improvement. This 
is contribution number 165 of the Grice 
Marine Biological Laboratory, College of 
Charleston. 


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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


113(2):386—396. 2000. 


Description of two new blenniid fish species: Entomacrodus lemuria 
from the western Indian Ocean and E. williamsi from the western 
Pacific Ocean 


Victor G. Springer and Ronald A. Fricke 


(VGS) Division of Fishes, MRC-159, National Museum of Natural History, 
Washington, D.C. 20560-0159, U.S.A.; (RAF) Staatliches Museum fiir Naturkunde in Stuttgart, 
Rosenstein 1, D-70191, Stuttgart 1, Germany 


Abstract.—Entomacrodus lemuria is described from specimens from Mad- 
agascar, Mauritius, and Renunion, and E. williamsi is described from specimens 
from Halmahera (Indonesia), Madang (Papua New Guinea), and Duff Islands 
(Santa Cruz Islands, Solomon Islands). New distribution records are provided 
for E. sealei, a widely distributed Pacific plate endemic, and E. corneliae, a 
Marquesas Islands endemic. All four species are members of the Nigricans 
Group of Entomacrodus, which group is distinguished from all other blenniids 
in having the lateral thirds of the ventral margin of the upper lip crenulate and 
the middle third entire. Within the group, these four species form a subgroup 
distinguished by having most of the five preopercular pore positions repre- 
sented externally by pairs or multiples of pores, as opposed to rarely having 
more than one or two positions with pairs or multiples of pores. 


Springer (1967) revised the circumtropi- 
cal shorefish genus Entomacrodus. There 
have been no new species described in the 
genus since that publication. Springer 
(1972:13) provided additional information 
on the Indo-Pacific species, including men- 
tion of a variant color pattern in a single 
specimen from Madang Harbor, Papua New 
Guinea. He tentatively identified the variant 
as Entomacrodus sealei Bryan & Herre, 
1903, which is otherwise unknown from the 
island of New Guinea. Springer (1982:19) 
reported a second specimen of the variant 
from Ternate, Indonesia, where typical 
specimens of E. sealei are also unknown. 
He believed that the distribution of the var- 
iant, from two localities near the western 
margin of the Pacific plate, was allopatric 
to that of typical E. sealei, a widely distrib- 
uted Pacific plate endemic (Springer 1982: 
fig. 8). Recently, in a single collection, J. T. 
Williams and associates obtained five spec- 
imens of the putative variant together with 


a large number of typical specimens of E. 
sealei. The sympatry of the two forms con- 
vinces us that two species are involved, and 
we formally describe the unnamed one 
herein. We have also obtained numerous 
specimens recently collected from Mauri- 
tius, Reunion, and Madagascar of another 
undescribed species that also appears to be 
related to E. sealei, and we describe that 
species herein. 

In recent years there has been much in- 
terest in documenting (inventorying) the 
fish faunas of many parts of the world. Con- 
comitantly, there has been much collecting 
and specimens have become available from 
many localities previously unsampled. 
Since 1967, there has been an extensive ac- 
cumulation of Entomacrodus specimens in 
museum collections, and the genus is in 
need of a new revision. We are unable to 
undertake that project in the forseeable fu- 
ture, but believe that when there is reason- 
able certainty that undescribed species ex- 


VOLUME 113, NUMBER 2 
Table 1. 


387 


Frequency distributions for number of preopercular pore positions represented by pairs or multiples 


of pores in specimens =25 mm SL of the Nigricans Species Group of Entomacrodus. There are five pore 


positions. 


Number of positions with pairs or 


Species N 0 i. 
cadenati 84 84 - 

textilis a, 54 l 
caudofasciatus 149 145 4 
vomerinus 113° ,- 409 3 
nigricans 343. 314 826 
chiostictus 487 184 164 
williamsi 7 - - 

lemuria 47 - l 
sealei 99 - - 
corneliae 14 - - 


ist, their descriptions are warranted without 
delay. 

Although the species of Entomacrodus 
have not been analyzed cladistically, some 
of the seven species groups Springer (1967: 
12—13) recognized in the genus are proba- 
bly monophyletic. One of these, the Nigri- 
cans Group, was defined on the basis of the 
morphology of the ventral margin of the 
upper lip: the central third of the margin is 
entire and the lateral third of the margin on 
each side is crenulate. This morphology 
does not occur in any other blennioid and, 
thus, serves as a reasonable basis for an hy- 
pothesis of monophyly of the group. Based 
on this character, the two new species are 
both members of the Nigricans Group. 


Methods 


Including the two new species, there are 
ten species in the Nigricans Group. There 
are very few characters that distinguish 
these species, and primary among them are 
a few particulars of the color pattern and 
sensory pores. For this reason, our descrip- 
tions are brief and limited to little more than 


multiples of pores 


2 3 4 2 X 
: = - - 0.00 
: . - - 0.02 
- - - - 0.03 
] - - - 0.04 
e) - - = 0:09 
DOO ce mah 2 Tee 
it 2 3 = Doe 
2; aged Seri Oi, LS oa SeBO 
- ii Jo. 4.09 
- - “ae gh  D.00 


the essentials necessary to differentiate the 
taxa. 

Unless defined herein, methods are those 
of Springer (1967). All counts of vertical 
fin-ray elements were made from radio- 
graphs. When split to the base and sup- 
ported by a single pterygiophore, the pos- 
teriormost anal-fin ray was counted as | 
(Method A of Springer 1967:5). 

Preopercular pore positions: There are 
five sensory canal foramina on the lateral 
surface of the preopercle (Springer 1968: 
fig. 7; Smith-Vaniz & Springer 1971:fig. 
16). Each of these foramina is represented 
by (connected to) one or more pores in the 
skin immediately external to the foramen. 
We term each such representation a “‘preo- 
percular pore position’’ or simply “‘posi- 
tion’”’ when in context (Fig. 1). Springer 
(1967) recognized six pore positions, but 
his ventroanteriormost position does not 
exit from the lateral surface of the preoper- 
cle and is excluded from our definition. The 
pores are small and often difficult to see; 
however, the number of these positions with 
two or more pores is an important character 


388 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Piece 
Entomacrodus. External limits of preopercle margin 
indicated by dotted line; x below ventralmost preoper- 
cular pore position; pore position types shown from 
ventralmost to dorsalmost: paired, paired, multiples, 
multiples, simple. 


Diagrammatic illustration of head pores in 


for distinguishing species of the Nigricans 
Group (Table 1). In specimens shorter than 
24 mm SL the pore positions may consist 
only or mostly of single pores in all species 
of the Nigricans Group; however, in E. le- 
muria, all of the five specimens 18.2—22.7 
mm SL, we checked for this character, had 
two to five positions with pairs of pores, 
and the smallest specimen had four posi- 
tions with pairs of pores. To be on the con- 
servative side, we restrict our characteriza- 
tion of pore positions to specimens 24 mm 
SL and longer (Table 1). 

Color pattern: Many features of the color 
pattern in the Nigricans Group species are 
highly variable intraspecfically, others are 
reasonably consistent within and among the 
species. Some of the variation is the result 
of poor preservation, but much variation 
exists in well-preserved specimens. Among 
the most variable markings are the presence 
(or absence) and expression of the dusky 
bars on the body sides, and some of the 
dusky markings on the head and dorsal and 
anal fins. These markings, even in speci- 
mens that appear to exhibit full expression 


of color pattern, are of little use for diag- 
nosing species. The more consistent band- 
ing of the caudal fin is also of no use. We 
have found that only aspects of the color 
pattern of the humeral area dorsal to the 
pectoral-fin axil and the upper lip are of im- 
portance. For these reasons, we restrict our 
color-pattern descriptions to these two fea- 
tures. The reader desiring more detail is re- 
ferred to our illustrations and those of the 
Nigricans Group species in Springer (1967: 
pls. 18-27). 

Institutional abbreviations are those list- 
ed in Eschmeyer (1998:16—22). 


Materials 


Unless noted otherwise, data reported are 
taken from specimens cited in Springer 
(1967, 197). 


Nigricans Species Group 


Diagnosis.—Species of Entomacrodus 
with: central third of ventral margin of up- 
per lip entire and lateral thirds crenulate; 
supraorbital cirri branched; nuchal cirri 
simple; posteriormost dorsal-fin spine re- 
duced, not visible externally; segmented 
dorsal-fin rays 13-17 (rarely 13, strongly 
modally 14 or 15 in all but one species: 
modally 16 in E. vomerinus from the south- 
western Atlantic); segmented anal-fin rays 
14-18 (rarely 14 and only in E. nigricans, 
from the Caribbean; strongly modally 16 in 
all but E. vomerinus, modally 17, and 18 
only in E. vomerinus); vertebrae 10+23-—26 
(strongly modally 34 in all but E. vomeri- 
nus, modally 35). Small species, largest 
specimen (always male) ranging from 53— 
105 mm SL, attaining more than 70 mm 
only in E. lemuria (72.1 mm) and E. vom- 
erinus (105 mm). 


Species Subgroups and Species 
Distributions 


The ten species of the Nigricans Group 
appear to fall into three subgroups based on 
the number of preopercular pore positions 


389 


VOLUME 113, NUMBER 2 


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P] uosiapuapyy *. 


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spy sesanbreyy 


ISWDI]]IM “3 
lajvas “J 
DUNUA] “Fy 
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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Figs 3. 
humeral spot); b & c, SMNS 213272, 61.5 and 57.4 mm SL. 


with pairs or multiples of pores (Table 1). 
The monophyly and inter- and intrarelation- 
ships of these subgroups are problematic, 
but the distributions of each subgroup and 
the species within each subgroup seem to 
correlate more strongly with geography 
than might be expected by chance and may 
hint at the naturalness of the groups. 
Subgroup 1 comprises five species, of 
which 91.5—100% of the specimens have all 
pore positions represented by single pores, 
and no specimen has more than two posi- 
tions represented by pairs or multiples of 
pores. Subgroup 2 comprises only one spe- 
cies, in which 37.8% of the specimens have 
only simple pore positions, and based on 


Entomacrodus lemuria, Reunion: a, SMNS 20827, holotype, 59.8 mm SL (pectoral fin obscures 


the mean number of positions with paired 
or multiple positions (Table 1) is more sim- 
ilar to Subgroup 1 than Subgroup 3. Sub- 
group 3 comprises four species, in which 
no specimen has only simple pore positions. 
Subgroup 1 includes all four of the Atlantic 
species and one eastern Indian Ocean-west- 
ern-and-central Pacific species: E. cadenati 
Springer (tropical eastern Atlantic, except 
St. Helena and Ascension), E. textilis (Quoy 
& Gaimard) (Ascension and St. Helena), E. 
nigricans Gill (Caribbean Sea, Bermuda, 
Bahamas), E. vomerinus (Valenciennes) 
(southwestern Atlantic), and E. caudofas- 
ciatus (Regan), eastern Indian Ocean east 
to Henderson Island, Pacific Ocean. 


VOLUME 113, NUMBER 2 


a 


39] 


Fig. 4. Entomacrodus lemuria, Mauritius: a, USNM 341905, 66.3 mm SL (note damage to second dorsal 
fin); b & c, USNM 339747, 43.9 mm SL, ventral and anterior views of head. 


Subgroup 2 comprises only E. chiostictus 
(Jordan & Gilbert) (eastern Pacific). 
Subgroup 3 (Fig. 2) comprises only Pa- 
cific or Indian Ocean species: E. corneliae 
(Fowler) (Marquesas Islands endemic), E. 
sealei (widely destributed Pacific plate en- 
demic, but absent from Marquesas Islands), 
and the two new species, E. williamsi 
(western margin of Pacific plate) and E. le- 
muria (southwestern Indian Ocean). 
Within Subgroups 1 and 3, all the species 
are allopatric except for one co-occurrence 
of E. sealei and E. williamsi near the west- 
ernmost limits of E. sealei. Considering all 
the species together, the only other occur- 
rence of sympatry is that of the widely dis- 
tributed E. caudofasciatus with E. sealei. 
If the sister species of E. lemuria is E. 
sealei, or E. sealei and one or both of the 
other two Group 3 species, the distribution 


pattern shown by E. lemuria and its sister 
taxa is another example of eastern Indian 
Ocean-Pacific plate disjunct distributions 
reported by Springer & Williams (1990). 
Those authors hypothesized that the broad 
gap between the two distributions was the 
result of extinction and explained the gen- 
esis of widely distributed Pacific plate en- 
demic species, such as E. sealei. 


Entomacrodus lemuria, new species 
Figs. 3 & 4 


Entomacrodus sp. Fricke, 1999:478, Ré- 
union; Mauritius. 


Holotype.—SMNS 20827, male, 59.8 
mm SL, Réunion, west coast, 250 m N 
Boucan-Canot, 5 km WSW St. Paul, 
21°01'35"S, 55°13'36"E, intertidal area of 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


ign. 
holotype, 59.8 mm SL lateral and frontal views of head; b & d, USNM 357792, 37.7 mm SL, frontal view of 
head and lateral view. 


narrow fringing reef, near black rocks on 
shore, R. Fricke and S. Ribes, 18 Dec 1998. 

Paratypes.—SMNS 21372, 4 specimens, 
43.4—61.4 mm SL, collected with the ho- 
lotype; AMS 1.39536-001 (5:40.9-61.5 mm 
SL), CAS 209025 (5:25.6—60.0 mm SL), 
ROM 71976 (5:29.7-54.7 mm SL), RUSI 
60472 (5:42.5-63.3 mm SL), USNM 
339747 (57:18.1—67.9 mm SL), all Mauri- 
tius, W coast, Albion, off rocks at end of 
Victoria Avenue, surge zone, O—5 m, A. C. 
Gill, P. C. Heemstra, M. Smale, and D. G. 


Entomacrodus williamsi, Duff Islands, Santa Cruz Islands, Solomon Islands: a & c, USNM 356864, 


Smith, 15 May 1995, field no. PCH 95- 
M23; USNM 341905 (4:23.7-66.3 mm 
SL), same data as preceding, except: tide 
pools, between 25 April and 17 May 1995, 
field no. PCH 95-M9; USNM 357266 (29.5 
mm SL), Madagascar, Nosy Be, near An- 
dilama Beach Hotel, J. Paxton et al., 7 Nov 
1988, Vityaz cruise 17, field no. JP 88-9. 
Diagnosis.—Ventral margin of upper lip 
crenulate on lateral thirds; entire on central 
third; nape cirri present, simple; orbital cirri 
branched; 1 to 5, usually 3 to 5, preoper- 


VOLUME 113, NUMBER 2 


393 


Fig. 6. Entomacrodus corneliae, Marquesas Islands, 


frontal and ventral views of head. 


cular pore positions with pairs or multiples 
of pores; subquadrate dark blotch present in 
humeral area on body (Figs. 3b, c; 4a); up- 
per lip frequently with up to 15 alternating 
dark and 15 pale bands of subequal width, 
up to 7 of each above central, ventrally en- 
tire portion of lip (Fig. 4c, d); prominent, 
irregular dark mark on head just posterior 
to midpostorbital sensory pores (Figs. 3 & 
4a). 

Description (* denotes character state of 
holotype).—Dorsal fin XIII,14 (n = 4), 
Pott. 197> (46), > oF XTVA17 © C1) 2) Anal. fin 
IE16* (G3) or 11,17.(7/), dast: ray split to 
base* (37) or simple (3, of which all have 
17 segmented rays). Vertebrae: precaudal 
10+23 (1), 24 (32), or 25* (6); pleural ribs 
11*-(30); epineural ribs: 13* (9); 14.(6), 15 
(11), 16 (0), 17 (1). Segmented pelvic-fin 
rays 4* (40). Pectoral-fin rays 13/14 (2), 14/ 
14* (37), or 14/15 (1). Dorsal procurrent 
caudal-fin rays/ventral procurrent caudal-fin 
rays 7/6 (2), 7/7* (10), 8/7 (18), 8/8 (4), or 
9/8 (1); segmented caudal-fin rays 13* (27), 


Nuku Hiva: AMS I.21773010, a, lateral view; b & c, 


of which middle 9 are branched, dorsalmost 
and ventralmost 2 are simple. 

Orbital cirrus branched (usually only on 
medial edge of stoutest cirrus), free cirrus 
tips 3-8, number tending to increase with 
size, variable bilaterally (holotype has 6/5). 
Nape cirri 1* on each side, rarely with | or 
2 tiny branches. Nasal cirri palmate, 5 to 10 
on each side (holotype with 8/9). 

Lateral-line a continuous posteroventral- 
ly curving tube extending posteriorly to 
vertical from base of 9th to 13th dorsal-fin 
spine (10th*), continuing along body mid- 
side as series of 1 to 8 (5*) tiny, discon- 
nected, bipored tubes extending to vertical 
from 11th dorsal-fin spine to 5th segmented 
ray (2nd segmented ray*). Preopercular 
pore positions (Table 1) with 1 to 5* posi- 
tions with pairs or multiples of pores. 

Color pattern: Highly variable ranging 
from almost lacking distinctive markings to 
being well marked. Some indication of the 
degree of color-pattern variation is indicat- 
ed in Fig. 3, all specimens taken in the same 


394 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig57. 
and frontal views of head. 


collection. Most noticeable and consistent 
marking a diagnostic subquadrate to ovoid 
dark spot on body just dorsoposterior to 
pectoral-fin axil. Other markings may in- 
clude: irregular dark spot on sensory pore 
series posterior to eye; snout above lip and 
sides of head with irregular pattern of dark 
and pale markings; underside of head with 
up to about 6 dusky chevrons separated by 
narrower pale bands; upper lip often with 
series of up to 15 dark bands alternating 
with up to 15 pale bands (dark lip bands, 
when present, 5 or 6 on central part of lip 
in area over noncrenulate portion of upper 
lip. Body with 4 or 5 pairs of dusky bands 
on sides, with offset dorsal portions. 
Comparisons.—Entomacrodus lemuria 
is the only species of Subgroup 3 that has 
as the darkest marking on the body, a sub- 
quadrate to ovoid spot in the humeral area. 
Entomacrodus sealei occasionally has a 
dark slash in the humeral region, but it is 


Entomacrodus sealei, Eua, Tonga Islands: USNM 335206, 57.8 mm SL, a, lateral view; b & c ventral 


much more slender than the humeral spot 
of E. lemuria. The humeral spot is essen- 
tially the only difference between E. lemu- 
ria and E. williamsi. Among the other two 
members of Subgroup 3, E. lemuria also 
differs from EF. sealei (Fig. 7) in having 
more dark and pale lip bands, including 
more in the region above the noncrenulate 
portion of the upper lip, and in having the 
pale bands only a little less slender than the 
dark bands (versus almost consistently 7 or 
8, each, dark and pale alternating bands, 
midlip dark bands much broader than pale 
bands, no more than 4 of each in area above 
noncrenulate portion of lip). Entomacrodus 
lemuria differs from E. corneliae (Fig. 6) 
in having only one prominent irregular dark 
spot on the head posterior to midpostorbital 
sensory pores (versus 2 such dark marks). 

Etymology.—From Lemuria, a hypothet- 
ical continent, supposed to have existed in 
the Indian Ocean and now represented by 


VOLUME 113, NUMBER 2 


Madagascar and some adjacent islands; 
here used as a noun in apposition. 


Entomacrodus williamsi, new species 
Fig: 5 


Holotype.-—USNM 356864, male, 52.8 
mm SL, Solomon Islands, Santa Cruz Is- 
lands, Duff Islands, Lakao, NW end at Te- 
momoa Pt, 09°47'54”S, 165°05'18”E, small 
cove, surge channels in big boulders, in 
heavy surge, 0-10 m, 24 Sep 1998, J. T. 
Williams et al. 

Paratypes: USNM 357792, 4 specimens, 
36.5—38.1 mm SL, collected with holotype. 
USNM 206400, 37.6 mm SL, Papua New 
Guinea, Madang Harbour, S edge of Massas 
Island, O—2.4 m, 2 Jun 1970, B. B. Collette, 
field no. BBC 1501. USNM 356244, 32.4 
mm SL, Indonesia, Moluccas, Ternate, Ter- 
nate [city], 0.2—-0.5 m, 1-2 July 1979, H. 
Singou, field no. HS-F610. 

Description (* denotes character state of 
holotype).—Dorsal fin XIII,14 (n = 1) or 
Poriet>~ (6). Anal fin 11,15 (1), 11,16* () 
onal.t7.(1), last ray split to base* (5) or 
simple (2, of which | has 15 and 1 has 17 
segmented rays). Vertebrae: precaudal 
10+24* (7); pleural ribs 11* (7); epineural 
ribs 14* (5) or 15 (1). Segmented pelvic- 
fin rays 4* (7). Pectoral-fin rays 13/14 (1) 
or 14/14* (6). Dorsal procurrent caudal-fin 
rays/ventral procurrent caudal-fin rays 8/2 
(hee, 7 (2), 8/8* (2), or 9/8 (1); segmented 
caudal-fin rays 13* (7), of which middle 9 
are branched, dorsalmost and ventralmost 2 
are simple* (6, one specimen damaged). 

Orbital cirrus usually branched (usually 
only on medial edge of stoutest cirrus), free 
cirrus tips 1—6 (1 or 2 probably abnormal), 
variable bilaterally (holotype has 5/6). Nape 
cirri 1* on each side (1 specimen with cir- 
rus forked on one side). Nasal cirri palmate, 
4 to 7 on each side (holotype with 6/7). 

Lateral-line a continuous posteroventral- 
ly curving tube extending posteriorly to 
vertical from base of 9th to 12th dorsal-fin 
spine (11th*), continuing along body mid- 
side as series of 2 to 6 (*5) tiny, discon- 


395 


nected, bipored tubes extending to vertical 
from 13th dorsal-fin spine to 5th segmented 
ray (4th ray*). Preopercular pore positions 
(Table 1) with 2 to 5 positions with pairs 
or multiples of pores (5*; see Table 1). 

Comparisons.—The lack of a dark hu- 
meral spot in E. williamsi is essentially the 
only difference between it and E. lemuria. 
There are very few specimens of E. wil- 
liamsi, and none of these exhibits distinc- 
tive markings on the body, which may have 
been present in life. If lack of pattern on 
the body of preserved specimens is typical, 
this would constitute another difference 
from E. lemuria, and the other two species. 
Among the other two species of Subgroup 
3, E. williamsi also differs from E. sealei 
(Fig. 7) in having more dark and pale lip 
bands, including more in the region above 
the noncrenulate portion of the upper lip 
and in having the pale bands only a litte 
less slender than the dark bands (versus al- 
most consistently 7 or 8, each, dark and 
pale alternating bands, midlip dark bands 
much broader than pale bands, no more 
than 4 of each in area above noncrenulate 
portion of lip), and from E. corneliae (Fig. 
6) in having only one prominent irregular 
dark spot on the head posterior to midpos- 
torbital sensory pores (versus 2 such dark 
marks). 

Etymology.—Named for our colleague 
Jeffrey T. Williams, in recognition of his 
outstanding efforts in collecting Indo-Pacif- 
ic fishes. 


Entomacrodus corneliae (Fowler, 1932) 
Fig. 6 


Additional material (all of the few known 
specimens are from the same locality): 
Marquesas Islands, Nuku Hiva: AMS 
E217 73010) (9). 


Entomacrodus sealei Bryan & Herre, 
1903 
Fig. 7 


New distribution records for E. sealei in- 
clude: Loyalty Islands: Lifou Island, SMNS 


396 


21712 (1 specimen). Vanuatu: Tanna, 
USNM 344230 (13); Epi, USNM 356399 
(7). Fiji: Rotuma, USNM 283060 (1). Ton- 
ga: Eua, USNM 329699 (>30), 335206 
(12); Tongatapu, USNM 337428 (1); 
Vava’u Group, Hunga, USNM 339314 (8); 
Vava’u, USNM 340229 (2). Solomon Is- 
lands: Santa Cruz Islands, Duff Islands, 
Lakao, USNM 356857 (92); Taumako, 
USNM 357098 (12); Russell Islands, 
Mbanika (or Banika), NMNZ P.26004 (1). 
Tuamotu Archipelago: Rangiroa, BPBM 
14007 (8). Pitcairn Islands: Henderson Is- 
land: BPBM 17091 (10). 


Acknowledgments 


For the loan of specimens we thank J. E. 
Randall (BPBM), M. McGrouther (AMS), 
and C. Roberts (NMNZ). Several USNM 
colleagues provided a variety of services: 
K. Darrow prepared the distribution map; 
K. Murphy provided technical assistance; J. 
Clayton, L. Palmer, and S. Raredon handled 
loan and accession actions. T. B. Griswold 
produced the photographs for Figs. 3-7. 
The U.S.N.M. Office of Biodiversity Pro- 
grams Grant to V. G. Springer and J. T. Wil- 
liams provided funds for Williams’s expe- 
dition to the Santa Cruz Islands (1998), dur- 
ing which important material was collected. 
U.S.N.M. Research Opportunities Fund 
erant. no. 1233F57A. to..D.G. Smith 
(USNM) supported his participation in an 
expedition to Mauritius, during which one 
of the new species was collected. A draft 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


of this manuscript was reviewed by J. T. 
Williams. 


Literature Cited 


Bryan, W. A., & A. W. C. T. Herre. 1903. Annotated 
list of the Marcus Island fishes.—Occasional 
Papers of the Bernice P. Bishop Museum 2 (1): 
126-139. 

Eschmeyer, W. N. 1998. Introduction. Pp. 16—22 in W. 
N. Eschmeyer, ed., Catalog of Fishes. Volume 
1. California Academy of Sciences, 958 pp. 

Fowler, H. W. 1932. The fishes obtained by the Pinchot 
South Seas Expedition of 1929, with descrip- 
tions of one new genus and three new spe- 
cies.—Proceedings of the United States Nation- 
al Museum 80 (2906):1—16. 

Fricke, R. 1999. Fishes of the Mascarene Islands (Ré- 
union, Mauritius, Rodriguez), an annotated 
checklist with descriptions of new species. 
Koeltz Scientific Books, K6nigstein, 759 pp. 

Smith-Vaniz, W. FE, & V. G. Springer. 1971. Synopsis 
of the tribe Salariini, with description of five 
new genera and three new species (Pisces: 
Blenniidae).—Smithsonian Contribtions to Zo- 
ology 73:1—72. 

Springer, V. G. 1967. Revision of the circumtropical 
shorefish genus Entomacrodus (Blenniidae: Sa- 
lariinae).—Proceedings of the United States Na- 
tional Museum 122 (3582):1—150, Pls. 1—30. 

. 1968. Osteology and classisfication of the 

fishes of the family Blenniidae.—Bulletin of the 

United States National Museum 284:1-—85, pls. 

1-11. 

. 1972. Additions to Revisions of the blenniid 

fish genera Ecsenius and Entomacrodus, with 

descriptions of three new species of Ecsenius.— 

Smithsonian Contributions to Zoology 134:1— 

13% 

. 1982. Pacific plate biogeography, with special 

reference to shorefishes.—Smithsonian Contri- 

butions to Zoology 367:1—82. 

, & J. T. Williams. 1990. Widely distributed 

Pacific plate endemics and lowered sea level.— 

Bulletin of Marine Science 47(3):631—640. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


113(2):397—400. 2000. 


Ammodytoides leptus, a new species of sand lance 
(Teleostei: Ammodytidae) from Pitcairn Island 


Bruce B. Collette and John E. Randall 


(BBC) National Marine Fisheries Service Systematics Laboratory, 
National Museum of Natural History, Washington, D.C., 20560-0153, U.S.A.; 
(JER) Bernice P. B. Bishop Museum, P.O. Box 190, Honolulu, Hawaii 96817-0916, U.S.A. 


Abstract.—Ammodytoides leptus is described from 23 specimens from Pit- 
cairn Island. It has more lateral-line scales than any other known species of 
Ammodytoides or Bleekeria (119-123 vs. 88-118). It is thinner than A. pylei, 
A. kimurai, and A. gilli (body depth 8.6—9.5% SL vs. 9.5—-11.7%). 


During his expedition to the eastern 
South Pacific on board the schooner West- 
ward (Randall 1978, 1999), the second au- 
thor collected the first sand lance recorded 
from the South Seas at Pitcairn Island in 
January 1971. The purpose of this paper is 
to describe this sand lance and assess its 
relationships to other members of the genus 
Ammodytoides as defined by Ida et al. 
(1994), including two additional species de- 
scribed by Ida & Randall (1993) and Rand- 
all et al. (1994) and A. gilli redescribed by 
Collette & Robertson (2000). 


Materials and Methods 


Type specimens of the new species have 
been deposited in the Bernice P. Bishop 
Museum, Honolulu (BPBM), the National 
Museum of Natural History, Washington, 
D.C. (USNM), the Australian Museum, 
Sydney (AMS), the Museum of Compara- 
tive Zoology, Cambridge (MCZ), and the 
National Science Museum, Tokyo (NSMT). 
Institutional abbreviations for sources of 
comparative material follow Leviton et al. 
(1985). Measurements follow Ida and 
Randall (1993) and Randall et al. (1994). 
Generic nomenclature follows Ida et al. 
(1994). All proportions are presented as 
percent of standard length (SL). 


Ammodytoides leptus, new species 
Fig. 1 


Ammodytes sp. Randall 1999:24 (an undes- 
cribed species from Pitcairn). 


Diagnosis.—A species of Ammodytoides 
with dorsal-fin rays 50—53; anal-fin rays 
24-25; pectoral-fin rays 16—17; pelvic fins 
absent; lateral line incomplete, pored later- 
al-line scales 114-118 + 4-6 unpored 
scales = 119-123; gill rakers on first arch 
(6-7) + (22-25) = 29-32; vertebrae (34-— 
36) + (26-27) = 61-63, including hypural 
plate. 

Description.—Body elongate, depth 8.6— 
9.5%, width 5.6—6.3%; head length 23.3- 
25.1%; snout length 6.7—7.3%; orbit diam- 
eter 4.1—4.5%; fleshy interorbital distance 
3.5—3.7%; upper jaw length 7.8—8.6%; least 
caudal peduncle depth 4.7—5.1%; caudal 
peduncle length 6.6—8.3% SL; predorsal 
distance 24.0—28.5%; preanal distance 
63.3—66.2%; caudal-fin length 13.6—-14.5%; 
caudal fin concavity 6.3—7.1%; pectoral-fin 
length 9.0—10.1% (Table 1). 

Scales small, thin, and cycloid, arranged 
in straight diagonal rows; head naked, no 
row of small scales on upper part of oper- 
cle, scales extending anteriorly to supratem- 
poral lateral-line canal; about 10—12 rows 
of predorsal scales; fins naked except cau- 
dal fin, which has scales extending about 


398 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Table 1.—Proportional measurements of type specimens of Ammodytoides leptus expressed in percent of 


standard length. 


Holotype 
Standard length (mm) 96:2 89.1 86.9 
Body depth 8.6 a2 95 
Body width 6.2 5.8 6.3 
Head length 233 24.8 24.4 
Snout length 6.8 6.7 70 
Orbit diameter 4.2 4.2 4.1 
Interorbital space oy 3] Se, 
Upper jaw length 7.8 8.0 i9 
Caudal peduncle depth 4.7 4.7 4.7 
Caudal peduncle length 8.3 7.4 — 
Predorsal length 262 24.0 24.7 
Preanal length Gna 65.7 66.2 
Caudal fin length 14.4 1326 Shed) 
Caudal concavity eal 6.7 6.7 
Pectoral fin length 10.1 oe wee) 


three-fourths distance to posterior margin; 
lateral line high on body, ascending from 
gill opening to three scales below origin of 
dorsal fin, passing posteriorly parallel to up- 
per edge of body, pored scales ending high 
on body, about 4—6 scales from caudal-fin 
base. Suborbital lateral-line canal interrupt- 
ed, with four preorbital and four postorbital 
pores (as in A. pylei; Randall et al., 1994: 
fig. 2A). 

In life, dorsum grayish green, sides sil- 
very and ventrum iridescent; fins hyaline, 
caudal yellowish. Specimens brown in pre- 
servative. 

Comparisons.—Ammodytoides leptus 
has more lateral-line scales than any other 
known species of Ammodytoides or Bleek- 
eria (119-123 vs. 88-118). It is thinner 


Paratypes 
86.4 85.3 o20 81.3 79.8 75.8 
973 9:0 8.8 8.7 8.9 9:0 
6.1 5.6 5.6 6.0 5.6 Do) 
24.1 24.2 24.6 24.6 24.4 2a 
TA 6.9 7.0 7.0 7.0 LS 
4.1 4.1 4.1 4.2 4.4 4.5 
Sy 3.6 351 Se EFS) 5 
8.1 TS 8.2 8.2 8.0 8.6 
4.9 4.9 4.8 4.8 5:0 Sail 
6.9 6.8 6.6 6.8 6.8 69 
24.3 28.5 PLES 5 24.7 24.4 24.1 
66.2 63:3 65.1 65.7 65.8 65.0 
13:9 14.2 14.5 Nes ei 13%5 14.1 
6.8 6.6 6.6 6.3 6.8 — 
9.6 9.0 9.4 9.6 93 9.8 


than A. pylei, A. kimurai, and A. gilli (body 
depth 8.6—9.5% vs. 9.5—11.7%, Table 2). It 
resembles A. gilli and differs from A. pylei 
and A. kimurai in several morphometric 
characters, larger eye (4.1-5.1% vs. 2.8— 
3.7%), longer upper jaw (7.8—9.5% vs. 6.6— 
7.9%), longer predorsal distance (24.0— 
28.5% vs. 21.8—24.7%), and longer caudal 
fin (12.8-16.0% vs. 10.5—12.9%). There is 
no row of small scales on upper part of op- 
ercle as in A. pylei (Randall et al., 1994). 
Ecology.—The series of ten types was 
collected from a school of about 25 indi- 
viduals seen over sand with high ripple 
marks. Schools were fast swimming and ex- 
hibited rapid changes in direction, all fish 
in perfect unison. When frightened, as by a 
spear shot into the school, a few dove into 


Fig. L: 
20 Jan 1971. Photo by J. E. Randall. 


Ammodytoides leptus. Paratype, USNM 360077, 86.9 mm SL, Pitcairn Island, off Gudgeon Point; 


VOLUME 113, NUMBER 2 


399 


Table 2.—Morphometric comparison of four species of Ammodytoides expressed in percent of standard length. 
(Data for A. kimurai from Ida & Randall 1993; for A. pylei from Randall et al. 1993; and for A. gilli from 


Collette & Robertson 2000). 


A. leptus A. kimurai A. pylei A. gilli 
Min Max Min Max i iMin,’ | (Max. | °F Mis - | "Max? 

Standard length (mm) 75.8 96.2 99.4 121 93.2 168 51.6 84.5 
Body depth 8.6 9.5 10.0 10:3 10.0 1t.7 9.5 i 
Body width 3:0 6.3 8.1 8.4 7 ee 9.5 5.9 7.8 
Head length Pn e&, 25.1 i aes 23:6 22.0 24.6 25.4 27.8 
Snout length 6.7 W 6.7 Vike) 6.4 10 6.9 79 
Orbit diameter 4.1 4.5 2.8 Dis 2.9 3.7 4.1 52] 
Interorbital space 55 Epi 4.1 4.6 Hs) 4.1 ki. 4.9 
Upper jaw length 7.8 8.6 TS 7.9 6.6 13 7.8 9.5 
Caudal peduncle depth 4.7 ail 4.0 5.0 4.5 ms 4.8 5.4 
Caudal peduncle length 6.6 8.3 8.4 9.8 10.6 12.0 3.6 6.0 
Predorsal length 24.0 28.5 de | 24.2 21.8 24.7 2 26.9 
Preanal length G3:3 66.2 61.3 65.3 63.8 66.8 62.5 65.3 
Caudal fin length 13:6 14.5 i I) Re, 10.5 12.0 12.8 16.0 
Caudal concavity 6.3 Wl G1 es 5.3 5.8 6.3 es 
Pectoral fin length 9:0 10.1 9.1 oS 8.4 92 9.9 yee 
N 9 6 10 10 


the sand. A larger school (about 100 indi- 
viduals) was observed in 30 m around an 
offshore rock in the same general area. Am- 
modytoides leptus is eaten by larger pelagic 
fishes such as the jack, Carangoides ferdau, 
as are other species of Ammodytoides. 

Etymology.—Named leptus because it is 
thinner than other species of Ammodyto- 
ides. 

Distribution.—Ammodytoides leptus is 
presently known only from Pitcairn Island. 

Discussion.—Ammodytoides leptus, as 
well as two additional species recently de- 
scribed by Ida & Randall (1993) and Rand- 
all et al. (1994), fits the definition of the 
genus as given by Ida et al. (1994) and dif- 
fers from species of Bleekeria in the follow- 
ing characters: no teeth in jaws; infraorbital 
canal interrupted; lateral line ending high 
on caudal peduncle instead of curving 
downward and continuing onto caudal fin 
base; two predorsal bones present; olfactory 
rosettes absent; neural and haemal spines 
on four posterior caudal vertebrae expanded 
and flattened; dorsal-fin rays 51—53 (three 
more than the previously recorded range for 
the genus), and anal-fin rays 21-25. 

In addition to Ammodytoides leptus, 


known species of the genus include the 
type-species, A. vagus (McCulloch & 
Waite, 1916) from Lord Howe Island and 
New South Wales; A. renniei (Smith, 1957) 
from South Africa, Seychelles Islands, and 
the Chagos Archipelago (Winterbottom et 
al. 1989, Winterbottom & Anderson 1999); 
A. kimurai Ida & Randall, 1993 from the 
Ogasawara Islands; A. pylei Randall et al., 
1994 from the Hawaiian Islands; and A. gil- 
li (Bean, 1895) from the eastern tropical Pa- 
cific (Collette & Robertson 2000). 
Material examined.—23 specimens 
(71.4-127 mm SL) from 3 original lots 
from Pitcairn Island. Holotype USNM 
360076 (1, 96.2), off Gudgeon Harbor, 10.5 
m, rotenone; J. E. Randall and D. B. Can- 
noy; 20 Jan 1971. Paratypes BPBM 16949 
(23° 79°8=89: 1)” MCZ>. 157036 (hb, 981.3), 
AMS 1.39856-001 (1, 82.9), NSMT-P 
59154 (1, 86.4), USNM 360077 (3, 75.8- 
86.9), and USNM 360078 (1, 89.8, cleared 
and stained), same data as holotype. BPBM 
16441 (1, 83.3), dredge haul 2, 48—54 fms; 
16 Oct 1967. Additional material examined 
but not designated as types due to their poor 
condition: BPBM 1660 (12, 71.4—127), off 
W. Harbour, stomach of Carangoides fer- 


400 


dau, 50 ft, J. E. Randall and S. Christian; 
27 Dee. 1970: 

Comparative material examined: Am- 
modytoides kimurai Ida & Randall, 1993. 
Paratype. USNM 324610 (1, 121), Japan, 
Ogasawara Islands, off Minami-shima, 15 
m;.HeIida .& R..L. Pyles) Jun,19972. 

Ammodytoides pylei Randall, Ida, and 
Earle, 1994. Paratype. USNM 316514 (1, 
137), Hawaiian Islands, Oahu, Kahe Point; 
RK. LU: Pyle, A. Y. Suzumoto, J.B. Culp: 19 
May 1989. 

Ammodytoides renniei (Smith, 1957). 
RUSI 8440 (1, 56.4), Seychelles Islands, 
3°57'S, 54°32”E; 25 Jul 1978. ROM 41487 
(1, 60.7), Chagos Archipelago, Peros Ban- 
hos Atoll; Isle -dw—Coin; -5°25°20'S: 
71°46'52”E; 6 Feb 1979; R. Winterbottom. 

Ammodytoides vagus (McCulloch & 
Waite, 1916). Holotype. AMS I-9272 (143 
mm SL), Lord Howe Island. 

Bleekeria mitsukurii Jordan & Evermann, 
1903. USNM 59599 (2, 78.5—151), Japan, 
Kochi; H. M. Smith. UW 21253 (2, 91.6— 
107), Taiwan. 

Bleekeria viridianguilla (Fowler, 1931). 
Paratypes. ANSP 53462-5 (4, 116—137), 
Hong Kong; 1930; G. A. C. Herklots. 

Protammodytes sarisa (Robins & 
Bohlke, 1970). Holotype ANSP 113091 
(115 mm SL) and paratype ANSP 113092 
(99.2), Windward Islands, off east coast of 
St. Vincent713717 12"Ny 61-05 13) We A877. 
m; Pillsbury sta. 874; 6 Jul 1969. 


Acknowledgments 


Funds for support of field work on board 
the Westward were provided to the second 
author by the National Geographic Society 
(Grant No. 821). We thank curators and 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


staff of museums that house the compara- 
tive material for loans and access to the ma- 
terial. Ruth Gibbons X-rayed specimens. 
The manuscript was reviewed by Thomas 
A. Munroe. 


Literature Cited 


Collette, B. B., & D. R. Robertson. 2000. Redescrip- 
tion of Ammodytoides gilli, the eastern Pacific 
sand lance (Teleostei: Ammodytidae).—Revista 
de Biologia Tropical, (in press). 

Ida, H., & J. E. Randall. 1993. Ammodytoides kimurai, 
a new species of sand lance (Ammodytidae) 
from the Ogasawara Islands.—Japanese Journal 
of Ichthyology 40(2):147-151. 

, P. Sirimontaporn, & S. Monkolprasit. 1994. 
Comparative morphology of the fishes of the 
family Ammodytidae, with a description of two 
new genera and two new species.—Zoological 
Studies, Tokyo 33(4):251—277. 

Leviton,. A..E.,..R...H.-Gibbs, Jnj-E. Heald Ga 
Dawson. 1985. Standards in ichthyology and 
herpetology. Part I. Standard symbolic codes for 
institutional resource collections in herpetology 
and ichthyology.—Copeia 1985:805-—832. 

Randall, J. E. 1978. Marine biological and archaeolog- 
ical expedition to southeast Oceania.—National 
Geographic Society Research Reports, 1969 
Projects:473—495. 

. 1999. Report on fish collections from the Pit- 

cairn Islands.—Atoll Research Bulletin No. 

461, 36 p. 

, H. Ida, & J. L. Earle. 1994. Ammodytoides 
pylei, a new species of sand lance (Ammody- 
tidae) from the Hawaiian Islands.—Pacific Sci- 
ence 48(1):80—89. 

Winterbottom, R., and R. C. Anderson. 1999. Fishes 
of the Chagos Archipelago.—Ecology of the 
Chagos Archipelag, C. R. C. Sheppard & M. R. 
D. Seaward, eds., Linnean Society Occasional 
Publications 2:101—117. 

, A. R. Emery, & E. Holm. 1989. An annotated 

checklist of the fishes of the Chagos Archipel- 

ago, central Indian Ocean.—Royal Ontario Mu- 

seum, Life Sciences Contributions No. 145, 226 


Pp- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


113(2):401—405. 2000. 


Neostrengeria binderi, a new species of pseudothelphusid crab from 
the eastern Andes of Colombia (Crustacea: Decapoda: Brachyura) 


Martha R. Campos 


Universidad Nacional de Colombia, Instituto de Ciencias Naturales, Apartado Aéreo 103698, 
Bogota, Colombia, S.A. 


Abstract.—A new species of freshwater crab of the genus Neostrengeria 
Pretzmann, 1965, N. binderi, is described from Alto de Cunday, Tolima De- 
partment, Colombia. The addition of this new species brings to 18 species and 
2 subspecies the total number of taxa known in this genus endemic to the 


Eastern Andes of Colombia. 


The genus Neostrengeria Pretzmann, 
1965, comprising 18 species and 2 subspe- 
cies of pseudothelphusid crabs, is endemic 
to the Eastern Andes of Colombia. On the 
west slope, the genus was known as far 
south as Cundinamarca, but recent collec- 
tions in the Cunday region of eastern Toli- 
ma have resulted in the discovery of a new 
species, which extends the range of the ge- 
nus ca. 50 km further south. Additional ex- 
plorations in the region have failed to locate 
this species west of the Magdalena river, 
confirming the association of the genus 
with the Eastern Andes. 

The systematics of Neostrengeria were 
established by Rodriguez (1982) and have 
been recently reviewed by Campos (1992, 
1994). The geographical distribution of the 
genus has been discussed by Campos & 
Rodriguez (1985), and Campos (1992, 
1994). The general characteristics of the ge- 
nus and a key for the identification of the 
species was presented by Campos & Le- 
maitre (1998). 

The terminology used for the different 
processes of the male first gonopods is that 
established by Smalley (1964) and Rodri- 
guez (1982). The material is deposited in 
Museo de Historia Natural, Instituto de 
Ciencias Naturales, Universidad Nacional 
de Colombia, Bogota (ICN-MHN). The ab- 
breviations cb and cl indicate carapace 


breadth and carapace length, respectively. 
Color nomenclature follows Smithe (1975). 


Family Pseudothelphusidae Rathbun, 1893 
Tribe Strengerianini Rodriguez, 1982 
Genus Neostrengeria Pretzmann, 1965 


Neostrengeria binderi, new species 
Feo 2 


Holotype.—Laguna Los Catorce, Vereda 
Alto de Cunday, Tolima Department, Co- 
lombia, 470 m alt., 22 May 1998, leg. P. 
Binder: 1 male, cl 20.7 mm, cb 37.8 mm 
(ICN-MHN-CR 1702). 

Paratypes.—Same locality data as holo- 
type: 2 females, cl /19:15-17.9 mm, :cb 33.6— 
31.4 mm (ICN-MHN-CR 1703). 

Type locality.—Laguna Los Catorce, 
Vereda Alto de Cunday, Tolima Depart- 
ment, Colombia, 470 m alt. 

Diagnosis.—Carapace with median 
groove deep, reaching upper border of 
front. First male gonopod with mesial bor- 
der slightly convex with strongly subapical 
notch in caudal view; apex compressed ce- 
phalo-caudal, expanded mesially into sub- 
triangular projection; mesial lobe forming 
acute triangle. 

Description of holotype.—Carapace (Fig. 
1A) with cervical groove nearly straight, 
deep, ending some distance from lateral 
margin. Anterolateral margin with shallow 


402 


depression behind external orbital angle, 
followed by series of papillae on anterolat- 
eral half; posterior half smooth. Postfrontal 
lobes small, oval, delimited anteriorly by 2 
depressions; median groove deep, reaching 
upper border of front. Surface of carapace 
in front of postfrontal lobes inclined ante- 
riorly, depressed towards midline. Front bi- 
lobed, lacking distinct upper border in fron- 
tal view; lower margin visible in dorsal 
view, strongly sinuous in frontal view, with 
tubercles. Orbital margins each with row of 
tubercles. Dorsal surface of carapace 
smooth, covered by small papillae; regions 
distinctly marked. Third maxilliped with 
merus having sharp angle on distal half of 
external margin; exognath 0.57 length of is- 
chium (Fig. 1D). Orifice of efferent bran- 
chial channel irregularly ovate (Fig. 1B). 
First pereiopods heterochelous, left che- 
liped larger than the right. Merus with 3 
longitudinal crests as follows: upper one 
with rows of tubercles, internal lower one 
with row of teeth, and external lower one 
with row of tubercles. Carpus with few tu- 
bercles on internal crest, and blunt distal 
spine. Palms of both chelipeds smooth and 
swollen; fingers elongate, each 0.58 the 
length of propodi, tips crossing, and surfac- 
es with rows of small tubercles; finger of 
larger chelae not gaping when closed. 
Walking legs (pereiopods 2—5) slender 
(Fig. 1A). Dactyli elongated, each 1.5 times 
as long as propodi, with papillae and 5 lon- 
gitudinal rows of large spines diminishing 
in size proximally. Number of spines and 
papillae on each dactylus arranged as fol- 
lows: 1 anterolateral row and 1 anteroven- 
tral row each with 6 spines; 1 external row 
with 5 spines and 4 intercalated papillae 
and | pair of proximal papillae; 1 postero- 
lateral row with 4 spines and 1 posterov- 
entral row with 5 spines. First gonopod 
wide in caudal view, mesial border (Fig. 
2A, 1) slightly convex with strongly sub- 
apical notch in caudal view. Accesory lobe 
(Fig. 2A, 2), shorter than lateral lobe (Fig. 
2A, 3), flat caudally, elongate; lateral lobe 
wide and semicircular (Fig. 2A—D). Apex 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


in distal view compressed cephalo-caudally 
(Fig. 2E), expanded cephalically into sub- 
triangular projection; mesial lobe (Fig. 2E, 
4) forming acute triangle; mesocaudal pro- 
jection of spermatic channel (Fig. 2E, 5) bi- 
fid with acute spinules. 

Color.—The holotype preserved in al- 
cohol is light brown (near 121C, Mikado 
Brown) with pale brown (Verona Brown, 
223 B) specks on the dorsal side of the car- 
apace. The walking legs and chelipeds are 
cinnamon brown (Tawny, 38) dorsally and 
ventrally. The ventral surface of the cara- 
pace is buffy-brown (Antique Brown, 37). 

Etymology.—The species is named in 
honor of Dr. Philippe Binder, Colombian 
scientist who collected the specimens, and 
to recognize his efforts in stimulating a new 
generation of scientists at the Universidad 
de los Andes, Bogota. 

Remarks.—This species is most similar 
to Neostrengeria aspera Campos, 1992. 
Both can be distinguished by features of the 
first gonopod. The mesial border of the first 
gonopod (in caudal view) of N. aspera is 
slightly convex, similar to N. binderi, but 
this latter species has a strong subapical 
notch. The elongate accesory lobe of N. 
binderi is shorter than the lateral lobe, 
whereas in N. aspera this lobe is as long as 
the lateral lobe, with the apical portion ir- 
regular in form and densely covered with 
spinules on the distal half (cf., Campos & 
Lemaitre 1998). The lateral lobe is spatu- 
late, rounded in its distal portion, and sep- 
arated from the accesory lobe by a deep 
notch in N. aspera, while in N. binderi the 
lateral lobe is semicircular and is almost ad- 
pressed to the accesory lobe. In the apex of 
the gonopod, N. binderi is most similar and 
probably closely related to N. gilberti Cam- 
pos, 1992. The differences are basically that 
in N. gilberti the apex is oval (not com- 
pressed cephalo-caudally) and there is an 
expansion with a conspicuous cephalic 
spine (cf. Campos 1992), whereas in N. bin- 
deri the apex consists of a subtriangular 
projection. The mesial lobe in N. binderi 
shows an acute subtriangular feature, while 


VOLUME 113, NUMBER 2 403 


Fig. 1. Neostrengeria binderi, new species, male holotype, ICN-MHN-CR 1702. A, dorsal view of carapace 
and pereiopods; B, opening of left efferent branchial channel, external view; C, left chela, external view; D, left 
third maxilliped, external view; E, frontal view of carapace. 


404 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 2. Neostrengeria binderi, new species, male holotype, ICN-MHN-CR 1702. A, left first gonopod, caudal 
view; B, same, lateral view; C, same, cephalic view; D, same, mesial view; E, same, apex, distal view. 1, mesial 
border; 2, accesory lobe; 3, lateral lobe; 4, mesial lobe; 5, mesocaudal projection of spermatic channel. 


VOLUME 113, NUMBER 2 


in N. gilberti it is smaller and subcircular 
with a papilla. The mesocaudal projection 
of the spermatic channel in N. gilberti is 
awl-shaped, with one spinule on the inner 
side, while in N. binderi it is bifid with 
acute spinules. 


Acknowledgments 


The author is grateful to Dr. R. Lemaitre, 
G. Styles and Dr. J. Lynch for their useful 
comments on this manuscript. The illustra- 
tions were prepared by Juan C. Pinzon. 


Literature Cited 


Campos, M. R. 1992. New species of fresh-water crabs 
of the genus Neostrengeria Pretzmann, 1965 
(Crustacea: Decapoda: Pseudothelphusidae) 
from Colombia.—Proceedings of the Biological 
Society of Washington 105:540—554. 

. 1994. Diversidad en Colombia de los cangre- 

jos del género Neostrengeria.—Academia Co- 

lombiana de Ciencias Exactas Fisicas y Natur- 

ales. Col. Jorge Alvarez Lleras No. 5:1—143. 


405 


, & R. Lemaitre. 1998. A new freshwater crab 

of the genus Neostrengeria Pretzmann, 1965, 

from Colombia (Crustacea: Decapoda: Brach- 

yura: Pseudothelphusidae), with a key to the 
species of the genus.—Proceedings of the Bio- 

logical Society of Washington 111:899—907. 

, & G. Rodriguez. 1985. A new species of 
Neostrengeria (Crustacea: Decapoda: Pseudoth- 
elphusidae) with notes on geographical distri- 
bution of the genus.—Proceedings of the Bio- 
logical Society of Washington 98:718—727. 

Pretzmann, G. 1965. Vorlaufiger Bericht tiber die Fam- 
ilie Pseudothelphusidae.—Anzeiger der Oster- 
reichischen Akademie der Wissenschaften 
Mathematische Naturwissenschaftliche Klasse 
(1),1:1-10. 

Rathbun, M. 1893. Descriptions of new species of 
American freshwater crabs.—Proceedings of 
the United States National Museum 16 (959): 
649-661, pl. 73-77. 

Rodriguez. Gi 1982... Les\ceabes d-ean,) douce 
d’Amérique. Famille des Pseudothelphusi- 
dae.—Faune Tropicale 22:1—223. 

Smalley, A. 1964. A terminology for the gonopods of 
the American river crabs.—Systematic Zoology 
13:28-31. 

Smithe, E B. 1975. Nuturalist’s color guide. The 
American Museum of Natural History. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


113(2):406—419. 2000. 


Freshwater crabs (Brachyura: Potamoidea: Potamonautidae) from 
the rainforests of the Central African Republic, Central Africa 


Neil Cumberlidge and Christopher B. Boyko 


(NC) Department of Biology, Northern Michigan University, Marquette, Michigan 49855, U.S.A.; 
(CBB) Division of Invertebrate Zoology, American Museum of Natural History, 
Central Park West @ 79th St., New York, New York 10024, U.S.A., and 
Department of Biological Sciences, University of Rhode Island, 
Kingston, Rhode Island 02881, U.S.A. 


Abstract.—A collection of freshwater crabs of the family Potamonautidae 
from the Central African Republic made recently by the Central African Re- 
public Expedition of the American Museum of Natural History, New York, 
comprised four species in two genera. The collection included two species of 
Potamonautes MacLeay, 1838, P. paecilei A. Milne-Edwards, 1886, and P. 
ballayi A. Milne-Edwards, 1886 and two species of Sudanonautes Bott, 1955, 
S. africanus A. Milne-Edwards, 1869, and S. sangha, new species. Only six 
species of freshwater crabs have been previously reported to occur in the Cen- 
tral African Republic. The addition of S. africanus and S. sangha brings to 
eight the number of species of freshwater crabs reported from that country. 


The freshwater crabs reported on here 
were collected during a systematic survey of 
the freshwater and terrestrial Mollusca of the 
Central African Republic made by the 
American Museum of Natural History. The 
survey aimed to document biodiversity, to 
catalogue introduced species, and to identify 
intermediate hosts that may serve as vectors 
of parasite larvae responsible for disease in 
humans and domestic animals. The fresh- 
water crab collections were made in the vi- 
cinity of the village of Bayanga, Central Af- 
rican Republic, which lies on the banks of 
the Sangha river in a remote and largely 
roadless area close to southern Cameroon, 
the Republic Populaire du Congo (formerly 
French Congo), and Gabon. This region in- 
cludes an area of tropical forest that supports 
elements of both the West African and Zaire 
river basin faunas, and consequently has an 
unusually high biodiversity. 

The collection of freshwater crabs ob- 
tained included two species of Potamonau- 
tes MacLeay, 1838, and two species of Su- 


danonautes Bott, 1955. Both of these genera 
belong to the exclusively African freshwater 
crab family Potamonautidae Bott, 1970. The 
two species of Potamonautes are P. ballayi 
(A. Milne-Edwards, 1886) and P. paecilei 
(A. Milne-Edwards, 1886). Both are little- 
known species and their diagnostic charac- 
ters are redescribed here. Additionally, a lec- 
totype is herein designated for P. paecilei. 
One of the species of Sudanonautes is 
clearly S. africanus (A. Milne-Edwards, 
1869). This is a common and well-known 
Species that was recently redescribed by 
Cumberlidge (1995a, 1999). The second 
species of Sudanonautes in the collection is 
superficially similar in some respects to S. 
africanus, S. chavanesii (A. Milne-Edwards, 
1886), S. faradjensis (Rathbun, 1921), S. 
floweri (de Man, 1901), S. granulatus 
(Balss, 1929) and S. aubryi (H. Milne Ed- 
wards, 1853). However, the new specimen 
from the Central African Republic has a 
number of important characters that do not 
conform to the descriptions of any of these 


VOLUME 113, NUMBER 2 


species (Cumberlidge 1993, 1994, 1995a, 
1995b, 1995c, 1995d; 1999). Although the 
specimen is a subadult female, and ideally 
an adult male is needed to make a definitive 
identification, a preliminary description is 
provided here, based on several unique so- 
matic characters of the specimen. This new 
taxon is the eleventh species of Sudanon- 
autes to be described (Cumberlidge 1999). 
Characters of the gonopods, male abdomen, 
and male chelipeds will be described when 
more material (including an adult male) be- 
comes available. 

Only six species of freshwater crabs have 
been previously reported to occur in the 
Central African Republic (Bott 1955, Cum- 
berlidge 1999). These are: Potamonautes 
ballayi (A. Milne-Edwards, 1886), P. pae- 
cilei (A. Milne-Edwards, 1886), P. dybowski 
(Rathbun, 1904), Sudanonautes faradjensis 
(Rathbun, 1921), S. floweri (de Man, 1901) 
and S. granulatus (Balss, 1929). The addi- 
tion of S. africanus and S. sangha new spe- 
cies in the present work brings to eight the 
number of species of freshwater crab re- 
ported from the Central African Republic. 


Materials and Methods 


Figures were prepared by capturing an 
image with a digital camera and completed 
using the programs Adobe Photoshop™ and 
Adobe Illustrator® (Harvey 1999). The 
specimens are deposited in the American 
Museum of Natural History, New York, 
U.S.A. (AMNH). Abbreviations: Muséum 
national d’Histoire naturelle, Paris, France 
(MNHN); Muséum royale d’ Afrique central, 
Tervuren, Belgium (MRAC); Biology, 
Northern Michigan University, Marquette, 
Michigan, U.S.A. (NMU); Senckenberg Mu- 
seum, Frankfurt, Germany (SMF); cw, dis- 
tance across the carapace at the widest point; 
cl, carapace length measured along the me- 
dian line, from the anterior to the posterior 
margin; ch, carapace height (the maximum 
height of the cephalothorax); fw, front width 
measured along the anterior margin; s, tho- 
racic sternite; e, thoracic episternite; s4/s5, 


407 


S4/s5, s5/s6, s6/s7, s7/s8, sternal sulci be- 
tween adjacent thoracic sternites; s4/e4, s5/ 
e5, s6/e6, s7/e7, episternal sulci between ad- 
jacent thoracic sternites and episternites; P1— 
P5, pereiopods 1-5. 


Systematic Account 
Genus Potamonautes MacLeay, 1838 


Diagnosis.—Postfrontal crest completely 
crossing carapace and meeting anterolateral 
margins at epibranchial teeth. Anterolateral 
margin always lacking intermediate tooth 
between exorbital angle and epibranchial 
tooth. Mandibular palp always two-seg- 
mented. Exopod of third maxilliped always 
with long flagellum. Terminal article of gon- 
opod 1 short, about one-quarter to one-third 
as long as subterminal segment of gonopod 
1. Terminal article of gonopod 2 a long fla- 
gellum about 0.5—0.75 times as long as sub- 
terminal segment of gonopod 2. 

Remarks.—Bott (1955) revised Potamon- 
autes and included 38 species and 14 sub- 
species, and erected 15 subgenera to accom- 
modate these taxa. Since that work, a num- 
ber of other species and subspecies have 
been described. These are Potamonautes 
triangulus (Bott, 1959), P. brincki (Bott, 
1960) (Cumberlidge 1994, 1999; Stewart 
1997a), P. (Isopotamonautes) anchetiae ma- 
chadoi Bott, 1964, P. (Lirrangopotamonau- 
tes) lirrangensis adeleae Bott, 1968, P. (J) 
senegalensis Bott, 1970, P. dentatus Stew- 
art, Coke, & Cook, 1995, P. parvispina 
Stewart, 1997b, P. granularis Daniels, Stew- 
art, & Gibbons, 1998, and P. reidi Cumber- 
hdge; 1999: 

Bott (1955) assigned Potamonautes bal- 
layi and P. paecilei to the subgenus Longi- 
potamonautes Bott, 1955, which also includ- 
ed a number of other species of rainforest 
river crabs from Central Africa in which 
adult males have an elongated, highly arched 
right cheliped and sharp teeth on the antero- 
lateral margins of the carapace: P. vanden- 
brandeni (Balss, 1936), P. schubotzi (Balss, 
1914), P. punctatus Bott, 1955, P. ballayi 
acristatus Bott, 1955, and P. ballayi gono- 


408 


cristatus Bott, 1955. Although Bott (1955) 
recognised numerous subgenera of Pota- 
monautes including Isolapotmonautes, Pla- 
typotamonautes, Lirrangopotamonautes and 
Longipotamonautes, we prefer here to fol- 
low Cumberlidge (1999) and use Potamon- 
autes sensu lato for all species, pending a 
revision of the entire genus. 


Potamonautes ballayi (A. Milne-Edwards, 
1886) 
Figs. 2 


Thelphusa Ballayi A. Milne-Edwards, 1886: 
149.—A. Milne-Edwards, 1887:132, pl. 7, 
figs. 2., 22) 

Potamon (Potamon) ballayi: Rathbun, 1904: 
294, pl. 12, fig. 9—Rathbun, 1921:419- 
422, pl. 27—28, figs. 1, 10. 

Potamon (Potamonautes) ballayi: Balss, 
1936:174-177, figs. 9, 12-13. 

Potamon ballayi: Chace, 1942:206.—Ca- 
part, 1954:827, fig. 3. 

Potamonautes (Longipotamonautes) ballayi 
ballayi: Bott, 1955:244—245, pl. VII, figs. 
2a-d, figs. 23, 73. 


Type material and type locality.—Female 
holotype, Ngancin (=Nganchu = Ngabé), 
Republic Populaire du Congo (formerly 
French Congo), 03°18’S, 16°6’E, on oppo- 
site bank to Kwamouth, Democratic Repub- 
lic of Congo (formerly Zaire), coll. Apr 
1884, MNHN. 

Material examined.—Central African Re- 
public. 1 adult male, cw 18.8 mm (AMNH 
17826), about 19 km from the village of 
Bayanga, Yobei (Yobé) river, depth 0.1 m, 
sandy shore near large dam made entirely of 
closely interwoven branches and vines, coll. 
J. Cordeiro, 18 Jun 1998. 

Diagnosis.—Postfrontal crest not com- 
plete, epigastric lobes significantly separated 
from postorbital crests, and lateral ends of 
postorbital crests not quite meeting antero- 
lateral margins (Fig. 1A). Exorbital angle 
produced into small pointed tooth; epibran- 
chial tooth large, sharp, and pointing for- 
ward; anterolateral margin between exorbital 
angle tooth and epibranchial tooth smooth, 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


curving slightly outward, lacking interme- 
diate tooth; anterolateral margin posterior to 
epibranchial tooth smooth, raised, curving 
inward over mesobranchial surface of cara- 
pace, and not continuous with posterolateral 
margin. Carapace height approximately 
equal to front width (ch/fw 1.07). Front very 
broad, measuring almost one half width of 
carapace (fw/cw 0.45) (Fig. 1B). Sidewall of 
carapace with distinct vertical sulcus, con- 
tinuing downward in pterygostomial region, 
dividing sidewall into four parts. Exopod of 
third maxilliped with a long flagellum, is- 
chium of third maxilliped smooth lacking 
vertical sulcus (Fig. 1D). First thoracic ster- 
nal sulcus sl/s2 absent; second sulcus s2/s3 
deep, running horizontally across sternum; 
third sternal sulcus s3/s4 absent so that ster- 
num in this region completely smooth (Fig. 
1C). Thoracic episternal sulci s4/e4, s5/e5, 
s6/e6 and s7/e7 smooth, none marked by 
visible groove. Major cheliped of adult 
males distinct, with widely arched dactylus 
and propodus longer than carapace width 
(Fig. 1G). First carpal tooth of inner margin 
of carpus of cheliped large, slender, pointed; 
second carpal tooth pointed, half size of first 
tooth. Lateral inferior margin of merus of 
cheliped lined by small teeth, medial inferior 
margin of merus of cheliped smooth, with 
single large pointed distal meral tooth at dis- 
tal end (Fig. 1J); superior surface of merus 
ridged by rows of short carinae (Fig. 11). 
Terminal article of gonopod 1 short (about 
one-third as long as subterminal segment), 
longitudinal groove visible on dorsal and su- 
perior sides (but not on ventral side); entire 
terminal article slim, tubular, and directed 
outward at 45° angle to vertical, ending in 
wide tip forming distinct pointed process on 
medial side; lateral and medial folds on ter- 
minal article of gonopod 1 approximately 
equal (Figs. 2A—C). The adult size range of 
P. ballayi is between cw 28-30 mm. 
Description.—For a detailed description 
and additional illustrations see Rathbun 
(1921) and Bott (1955). For a brief descrip- 
tion of the type, see Capart (1954, fig. 3). 
Remarks.—Rathbun (1921) recorded the 


VOLUME 113, NUMBER 2 409 


Fig. 1. Potamonautes ballayi (A. Milne-Edwards, 1886). Male, cw 17.3 mm, AMNH 17826. A, carapace 
and eyes, dorsal view; B, cephalothorax, carapace and eyes, frontal view; C, anterior sternum; D, left third 
maxilliped; E, left mandible; K abdomen; G, right cheliped, frontal view; H, left cheliped, frontal view; I, carpus 
and merus of right cheliped, lateral view; J, carpus and merus of right cheliped, mesial view; K, left second 
pereiopod, lateral view. Scale = 1.6 mm (EB), 3.3 mm (D, G—J), and 4.4 mm (A-C, E K). 


410 


Fig. 2. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


VL 
mn csc 
oS 


a, eR 
2 Sty SSeS ORO 


Potamonautes ballayi (A. Milne-Edwards, 1886). Male, cw 17.3 mm, AMNH 17826. A, left gonopod 


1, cephalic view; B, left gonopod 1, caudal view; C, distal portion of left gonopod 1, superior view; D, left 
gonopod 2. Scale = 0.80 mm (C), and 1.67 mm (A, B, D). 


mandibular palp as three-segmented. This is 
an error, for it is clearly two-segmented (Fig. 
1E), as is the case for all members of the 
genus, and, indeed all potamonautid African 
freshwater crabs (Cumberlidge 1999). Be- 
cause the type of P. ballayi is a female, the 
gonopods were not illustrated in the first de- 
Scriptive works on the species. Gonopod 1 
of P. ballayi was subsequently illustrated by 
Rathbun (1921) who used a male (cw 26 
mm, AMNH 3356) from Stanleyville (now 
Kisangani, Democratic Republic of Congo) 
and by Bott (1955), who used a male (cw 
33 mm, MRAC 17413) from Karawa, Uban- 
gi, Democratic Republic of Congo. Gono- 
pod 1 of the male from the Central African 
Republic is shown here in more detail (Figs. 
2A-C), and gonopod 2 of P. ballayi is il- 
lustrated for the first time (Fig. 2D). 
Ecology.—The specimen from the Central 
African Republic was caught in shallow wa- 
ter (only 0.1 m deep) near a dam made en- 
tirely of closely interwoven branches and 
vines. Herbert Lang’s field notes (in Rathbun 
1921) record that P. ballayi is common in 
shallow forest streams around Stanleyville 


(now Kisangani). Lang wrote that it is prob- 
able that P. ballayi can live out of water and 
that it is only dependent on a certain amount 
of moisture. When disturbed, crabs were re- 
ported to instantly cover themselves with 
mud and secure protection beneath any ob- 
ject. 

Distribution.—Republic Populaire du 
Congo (formerly French Congo), Democrat- 
ic Republic of Congo, and Gabon. The type 
locality of P. ballayi at Ngabé, Republic Po- 
pulaire du Congo lies on the banks of the 
Zaire river opposite Kwamouth, Democratic 
Republic of Congo. The present study 
showed that P. ballayi is present in the Yobé 
river, a tributary of the Sangha river which 
drains into the Zaire river in a broad marshy 
area at Mossaka, Republic Populaire du 
Congo. For more localities see Rathbun 
(1921), Balss (1936), and Bott (1955). 


Potamonautes paecilei (A. Milne-Edwards, 
1886) 
Fig. 3 
Thelphusa paecilei A. Milne-Edwards, 
1886:149 


VOLUME 113, NUMBER 2 


Parathelphusa paecilei: A. Milne-Edwards, 
1887:143, pl. 7, figs. 1, la; Ortmann, 
1897:300. 

Potamon (Parathelphusa) paecilei: Rath- 
bun, 1905:257, fig. 167. 

Potamon paecilei: Chace, 1942:208.—Ca- 
part, 1954:841—842, figs. 34, 37. 

Potamonautes (Longipotamonautes) paeci- 
lei: Bott, 1955:242-243, pl. VI, figs. 2a— 
event figs. 21, 71. 


Type material and type locality.—Adult 
male lectotype, cw 32 mm (MNHN-B263), 
Central Africa, Republic Populaire du Con- 
go (formerly French Congo), Latéké (=Lék- 
éti), Alima river (14°56’E, 1°36’S), coll. M. 
de Brazzae. 

Material examined.—Central African Re- 
public. 1 adult female, cw 18.6 mm, 
(AMNH 17827), 17.3 km from the village 
of Bayanga (02°45'43"N, 16°14'12”E), Lossi 
creek, depth 1—2 m, bottom of the fine sand 
and mud, caught in net in swiftly moving 
black water, coll. J. Sullivan, 19 Jun 1998.— 
1 adult male, cw 20.7 mm (AMNH 18032), 
17.3 km from the village of Bayanga 
02645/43'"N, 16°14'12’”E),..Lossi creek, 
depth 1—2 m, bottom of fine sand and mud, 
caught in net in swiftly moving black water, 
coll. M. Lawrence, J. Sullivan, and local res- 
idents, 30 Jun 1998. 

Diagnosis.—Postfrontal crest either com- 
plete or almost complete, wherein lateral 
ends of postorbital crests not quite meeting 
anterolateral margins (Fig. 3A). Exorbital 
angle produced into small, pointed, sharp 
tooth; epibranchial tooth large, pointed, di- 
rected forward; anterolateral margin be- 
tween exorbital angle tooth and epibranchial 
tooth smooth, curving outward, lacking in- 
termediate tooth; anterolateral margin pos- 
terior to epibranchial tooth with two sharp, 
forward-pointing teeth; margin otherwise 
smooth, continuous with posterolateral mar- 
gin. Front very broad, measuring almost 
one-half width of carapace (fw/cw 0.45) 
(Fig. 3A, B). Carapace height approximately 
equal to front width (ch/fw 1.07). Exopod of 
third maxilliped with long flagellum, ischi- 


411 


um of third maxilliped smooth, lacking ver- 
tical sulcus (Fig. 3D). First thoracic sternal 
sulcus sl/s2 absent; second sulcus s2/s3 
deep, running horizontally across sternum; 
third sternal sulcus s3/s4 absent; sternum in 
this region completely smooth. Episternal 
sulci s4/e4, s5/e5, s6/e6 and s7/e7 smooth, 
none marked by visible groove. Major che- 
liped of adult males distinct, with widely 
arched dactylus and a propodus longer than 
carapace width. First carpal tooth of inner 
margin of carpus of cheliped large, slender, 
pointed; second carpal tooth half size of first. 
Lateral and medial inferior margins of merus 
of cheliped lined by small teeth; single large 
pointed distal meral tooth at distal end (Fig. 
3J); superior surface of merus ridged by 
rows of short carinae (Fig. 31). Terminal ar- 
ticle of gonopod 1 short (about one-third as 
long as subterminal segment), longitudinal 
groove visible on superior side, but not on 
dorsal and ventral sides; entire terminal ar- 
ticle slim, tubular, curved; directed outward 
at an approximately 45° angle to vertical; 
ending in broadened upcurved tip; lateral 
and medial folds on terminal article of gon- 
opod 1 approximately equal size. The adult 
size range of P. paecilei is between cw 28-— 
30 mm. 

Description.—See Bott (1955), and Ca- 
part (1954). 

Remarks.—One of us (NC) has examined 
an adult male syntype (cw 32 mm) of P. 
paecilei (MNHN-B 263) from the Alima riv- 
er, Latéké French Congo collected by M. de 
Brazzae. We designate this specimen here as 
the lectotype. A different male syntype (now 
paralectotype) of P. paecilei (cw 22.7 mm) 
was figured by Capart (1954). The charac- 
ters of this species (Fig. 3) include a large 
forward-pointing epibranchial tooth; a large 
tooth (or two teeth) behind the epibranchial 
tooth on the anterolateral margin; an en- 
larged major cheliped in adult males with a 
widely arched dactylus and a propodus that 
is longer than the carapace width; a long 
sharp distal meral spine on the merus of the 
cheliped; and the ischium of the third max- 


412 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 3. Potamonautes paecilei (A. Milne-Edwards, 1886). Female, cw 18.6 mm, AMNH 17827. A, carapace 
and eyes, dorsal view; B, cephalothorax, carapace and eyes, frontal view; C, anterior sternum; D, left third 
maxilliped; E, left mandible; EK abdomen; G, right cheliped, frontal view; H, left cheliped, frontal view; I, carpus 
and merus of right cheliped, lateral view; J, carpus and merus of right cheliped, mesial view; K, left second 


pereiopod, lateral view. Scale = 2.2 mm (E), 3.3 mm (D, I, J), 4.4 mm (A, B, F—-H, K), and 5.9 (C). 


VOLUME 113, NUMBER 2 


illiped is smooth and a vertical sulcus is 
lacking. 

The sidewall of the carapace of the spec- 
imens from the Central African Republic has 
a distinct vertical sulcus which continues 
downward across the pterygostomial region, 
dividing the carapace sidewall into four 
parts. This contrasts with the type from the 
Republic Populaire du Congo where the 
sidewall of the carapace is divided into only 
three parts. 

Ecology.—The specimens from the Cen- 
tral African Republic were netted in a small 
stream (1-2 m deep), with swiftly moving 
black water flowing over fine sand and mud. 

Distribution.—Republic Populaire du 
Congo, and Democratic Republic of Congo. 
The Yobé river is a tributary of the Sangha 
river which drains southwestern Central Af- 
rican Republic and forms part of the border 
between the Central African Republic and 
Cameroon, and then between Cameroon and 
Republic Populaire du Congo, before joining 
the Zaire river in a broad marshy area at 
Mossaka, Republic Populaire du Congo. The 
type locality lies on the Alima river which 
flows into the Zaire river just to the south of 
Mossaka and contributes to the same ex- 
panse of marsh and wetlands as the Sangha 
river. 


Sudanonautes Bott, 1955 


Sudanonautes Bott, 1955:295.—Cumberlid- 
ge, 1999:172-176. 


Diagnosis.—Intermediate tooth on antero- 
lateral margin between epibranchial tooth 
and exorbital angle. Postfrontal crest prom- 
inent, almost horizontal, complete, with lat- 
eral ends meeting anterolateral margins. Car- 
apace sidewall divided by two sulci into 
three parts. Mandibular palp two-segmented; 
terminal segment consisting of large oval 
posterior lobe (in three species there a small 
but distinct anterior process at junction be- 
tween segments). Long, plumose flagellum 
on exopod of third maxilliped in all species. 
Sternal sulcus s3/s4 represented only by two 
short notches at sides of sternum. Terminal 


413 


article of gonopod | very long (at least two- 
thirds as long as subterminal segment). Ter- 
minal article of gonopod | either slim and 
needle-like (where longitudinal groove not 
visible) or broadened in middle (the result 
of a higher medial fold) with longitudinal 
groove visible at least for part of length. Ter- 
minal article of gonopod 2 very short, one- 
fifteenth length of subterminal segment. 
Distribution. —The genus is present in 
Céte-d’ Ivoire, Ghana, Togo, Benin, Nigeria, 
Cameroon, Gabon, Bioko (Fernando Po), 
Central African Republic, Congo, Zaire, 
northern Angola, and southwest Sudan. The 
eleven species of Sudanonautes are found in 
the inland waters of West and Central Africa 
in a region bounded by Céte-d’ Ivoire, south- 
west Sudan, and northern Angola. This area 
includes the Upper Guinea rainforests, the 
Lower Guinea forest together with the sa- 
vannas of the eastern part of West Africa, 
and the offshore island of Bioko. In Central 
Africa seven species of Sudanonautes (S. af- 
ricanus, S. aubryi, S. floweri, S. granulatus, 
S. chavanesii, S. faradjensis, and S. sangha) 
share the rivers and forests with species of 
Potamonautes and Erimetopus A. Milne-Ed- 
wards, 1886 (Bott 1955, Cumberlidge 1999). 


Sudanonautes africanus (A. Milne- 
Edwards, 1869) 


Thelphusa africana A. Milne-Edwards, 
1869:186, pl. XI, figs. 2, 2a,b.—A. Milne- 
Edwards, 1887:124—126, pl. IV, fig. 8. 

Potamon (Potamonautes) africanus: de 
Man, 1903:41, pl. [X, figs. 7—9.—Rath- 
bun, 1904, pl. 16, fig. 6—Rathbun, 1905: 
188-190, fig. 47—Balss, 1929:124—125, 
figs. 5—7.—Balss, 1936:166. 

Potamon (Potamonautes) africanum: Colo- 
si, 1920:34.—Colosi, 1924:21, fig. 16.— 
Roux, 1927:237. 

Potamon africanus: Chace, 1942:204.—Ca- 
part, 1954:824, figs. 1, 6. 

Sudanonautes (Sudanonautes) africanus af- 
ricanus: Bott, 1955:295-—298, figs. 61, 93— 
95, 103 a-d, pl. 24, figs. 2a—c, 3.—Bott, 
1959:1004—1005.—Monod,  1977:1216 


414 


(not figs. 93-95, 102).—Monod, 1980: 
384, pl. V, fig. 27. 

Sudanonautes africanus: Cumberlidge, 
1995a:588—598, figs. 1-3, table 1.—Cum- 
berlidge, 1999:181—184, figs. 30B, 32B, 
33B, 34B, 35C, 36E, 37C, 53P, 54-57, 
60B, 67A, table IX. 


Type material.—The holotype used by A. 
Milne-Edwards (1869) to describe Thelphu- 
sa africana was a small juvenile (MNHN) 
(cw 17 mm) collected from Gabon by M. 
Aubry-Lecomte. A more detailed descrip- 
tion by A. Milne-Edwards (1887) was based 
on a larger, but still subadult female 
(MNHN) (cw 53 mm) collected from the 
river Ogoué, Congo (=Gabon). Because nei- 
ther of these specimens was suitable to re- 
describe the species (one is a juvenile and 
the other a subadult female), and because no 
topotypes were available, the species was re- 
described by Cumberlidge (1995a) from an 
adult male (cw 83 mm) from Cross River 
State, Nigeria (NMU 9.IV.1983), and an 
adult female (cw 108 mm) from a tributary 
of the Ikpan river, Cross River State, Nigeria 
(NMU 5.IV.1983). 

Material examined.—Central African Re- 
public. Ten specimens, sub-adults and juve- 
niles (no adults) (AMNH 18033), 19.5 km 
from the village of Bayanga (03°05'27'N, 
16°16’40"E), Mapoyo (Mboyé) creek, depth 
1—1.5 m, either on muddy bottom, or in bur- 
rows in overhanging banks, coll. M. 
Lawrence, J. Sullivan, and local residents, 17 
Jun 1998. Four specimens, 1 adult male and 
3 juveniles (AMNH 18034), about 5 km from 
the village of Bayanga, Mobeya [Moubia?] 
creek, upstream of mouth, depth 0.5—1 m, 
sandy, gravely and muddy pools, in burrows 
in overhanging banks, among roots, under 
logs, coll. M. Lawrence and local residents, 
26 Jun 1998. One subadult male (AMNH 
18035), 19.5 km from the village of Bayanga 
(03°05'27"N, 16°16'40"E), Mapoyo (Mboyé) 
creek, depth 1—1.5 m, on muddy bottom, or 
in burrows in overhanging banks, coll. M. 
Lawrence, J. Sullivan, and local residents, 17 
Jun 1998. Four specimens, all juveniles 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


(AMNH 18036), about 5 km from the village 
of Bayanga, Mobeya [Moubia?] creek, up- 
stream of mouth, depth 0.5—-1 m, sandy, 
gravely and muddy pools, in burrows in over- 
hanging banks, among roots, under logs, coll. 
M. Lawrence, J. Sullivan, and J. B. Kindi- 
moungo, 28 Jun 1998. 

Type locality.—Gabon. 

Diagnosis.—Carapace relatively flat (ch/ 
fw 1.06). Postfrontal crest smooth almost 
straight; spanning entire carapace, meeting 
anterolateral margins at epibranchial teeth; 
posterior surface of carapace in cardiac and 
branchial regions with patches of raised cir- 
cular blisters, lateral parts with fields of 
raised short lines (carinae); semi-circular, 
cardiac, urogastric grooves very deep. Prox- 
imal region of pollex of propodus of major 
cheliped of adult with large, conspicuously 
flattened tooth. Exorbital angle tooth large, 
triangular; intermediate tooth large, triangu- 
lar blunt, as big as exorbital angle tooth. Epi- 
branchial tooth small, about half size of in- 
termediate tooth and exorbital angle tooth. 
Anterolateral margin behind epibranchial 
tooth smooth. Terminal article of gonopod 1 
thin, needle-like, subterminal segment of 
gonopod 1 slim. This is the largest species 
of freshwater crab in Africa. Adult sizes 
range from the size at the pubertal molt (cw 
70—75 mm) to largest the known specimen 
(cw 113 mm). 

Description.—For a detailed description 
see Cumberlidge (1995a, 1999). For a brief 
description of the type see Capart (1954, 
figs: 1546): 

Remarks.—Sudanonautes africanus is a 
common and well-known species that was 
recently redescribed (Cumberlidge 1995a, 
1999): 

Ecology.—This species is restricted to the 
more humid areas of the coastal rainforest 
belt from south-east Nigeria to the mouth of 
the Zaire river. Sudanonautes africanus oc- 
curs in a range of permanent aquatic habitats 
from large rivers and small streams (with 
both fast and slow flowing water) to ponds. 
In the Central African Republic S. africanus 
is found in creeks up to 1.5 m deep with a 


VOLUME 113, NUMBER 2 


sand, gravel or mud bottom. Specimens 
were also taken from burrows in overhang- 
ing banks, among roots and under logs. Else- 
where in its range, this species is also com- 
mon in streams and rivers draining mature 
forest, and has been reported to dig burrows 
near waterways. This crab also occurs in 
temporary water sources such as drainage 
culverts and ditches. For more details see 
Cumberlidge (1995a, 1999). 

Distribution.—Sudanonautes africanus 
occurs in the coastal rainforest regions of Ni- 
geria and Central Africa. In Central Africa 
S. africanus occurs in south Cameroon, the 
Republic Populaire du Congo, and Gabon 
(in the San Benito, Ogoué and Alima rivers), 
and in the lower reaches of the Zaire River 
basin. For more details see Cumberlidge 
(1995a, 1999). The present record is the first 
report of the presence of S. africanus in the 
Central African Republic. 


Sudanonautes sangha, new species 
Fig. 4 


Type material and type locality.—Central 
African Republic. Holotype: 1 subadult fe- 
male, cw 40.5, cl 29.7, ch 12.5, fw 10.7 mm 
(AMNH 17825), a few km upstream from 
the village of Bayanga (02°45'43’N, 
16°14’12"E), Sangha river, depth 1—2 m, in 
fish trap, eating worm, coll. J. Sullivan and 
J. B. Kindimoungo, 18 Jun 1998. 

Diagnosis.—Exorbital tooth large, point- 
ed; intermediate tooth small, low; epibran- 
chial tooth small, low, not directed outward, 
set back behind mid-point of postfrontal 
crest. Postfrontal crest spanning entire cara- 
pace, crest curving backward before meeting 
epibranchial tooth, anterolateral margin pos- 
terior to epibranchial tooth raised, lined by 
small granules. Semi-circular, urogastric, 
transverse branchial grooves very deep. Ver- 
tical suture on carapace sidewall meeting in- 
termediate tooth. Carapace medium height 
(ch/fw = 1.16). Mandibular palp two-seg- 
mented; terminal segment single, undivided, 
with hair at junction between segments. First 
carpal tooth on carpus of cheliped large, 


415 


pointed; second carpal tooth reduced to 
small granule. 

Description.—Carapace (Figs. 4A, B).— 
Ovoid, widest in anterior third (cw/fw 3.79), 
medium height (ch/fw = 1.16), semi-circu- 
lar, urogastric, transverse branchial grooves 
very deep, regions smooth; cardiac region 
weakly marked, cervical grooves present but 
weak. Front slightly bilobed, anterior margin 
indented, relatively narrow, about one-quar- 
ter carapace width (fw/cw = 0.26) (Fig. 4B). 
Postfrontal crest smooth, spanning entire 
carapace, straight part consisting of fused 
epigastric, postorbital crests, then curving 
backward behind intermediate teeth to meet 
anterolateral margins at epibranchial teeth. 
Anterolateral margin smooth posterior to 
epibranchial tooth. Exorbital tooth large, 
sharp, pointed forward. Epibranchial tooth 
low, small, set back behind mid-point of 
postfrontal crest. 

Carapace sidewalls mostly smooth, with 
faint granules in suborbital regions. Each 
sidewall with two sutures, one longitudinal, 
one vertical, dividing sidewall into three 
parts. Longitudinal (epimeral) suture divid- 
ing suborbital, subhepatic regions from pter- 
ygostomial region, beginning medially at 
lower margin of orbit, curving backward 
across flank. Short vertical suture dividing 
suborbital region from subhepatic region; 
vertical suture meeting intermediate tooth. 
First transverse groove on sternum, between 
sternal segments s2 and s3, complete; sec- 
ond groove, between sternal segments s3 
and s4, consisting of two small notches at 
sides of sternum. Third maxillipeds filling 
entire oral field, except for transversely oval 
efferent respiratory openings at superior lat- 
eral corners; long flagellum on exopod of 
third maxilliped; ischium of third maxilliped 
smooth, with clear vertical groove (Fig. 4C). 
Mandibular palp two-segmented; terminal 
segment single, undivided, with hair but no 
hard flap at junction between segments (Fig. 
4D). Segments 1—6 of female abdomen four- 
sided, last segment a broad rounded triangle, 
sides forming a smooth curved, rounded 
margin; segments 5—6 broadest (Fig. 4E). 


416 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 4. Sudanonautes sangha, new species. Holotype subadult female, cw 40.5 mm, AMNH 17825. A, 
carapace and eyes, dorsal view; B, cephalothorax, carapace and eyes, frontal view; C, left third maxilliped; D, 
left mandible; E, abdomen; FE right cheliped, frontal view; G, left cheliped, frontal view; H, carpus and merus 
of right cheliped, lateral view; I, carpus and merus of right cheliped, mesial view; J, left second pereiopod, 


lateral view. Scale = 4.4 mm (D), 6.2 mm (I), 6.7 mm (C), 7.2 mm (H), 8.6 mm (EF G, J), and 12.7 mm (A, 
B, E). 


Dactylus of right cheliped long, slim, granules, cluster of granules surrounding 
straight; palm of propodus swollen; fingers larger pointed distal meral tooth at distal end 
of digits of chelipeds with small even teeth, (Fig. 41). Inner margin of carpus of cheliped 
forming long slim interspace when closed, with large, slender, pointed tooth (first carpal 
fingers almost meeting when shut (Fig. 4F). tooth), second carpal tooth reduced to a 
Inferior margins of merus with rows of small granule (Fig. 41). Pereiopods P2—P5 slender 


VOLUME 113, NUMBER 2 


(Fig. 4J), P4 longest, P5 shortest. Propodus 
of P2—P5 broad, posterior margin of propo- 
dus of P2—P5 serrated, dactyli of P2—P5 ta- 
pering to point, each bearing four rows of 
downward-pointing sharp bristles; dactylus 
of P5 shortest. 

Etymology.—tThe species is named for the 
Sangha river where it was collected. The 
Sangha river drains a large region of the 
Central African Republic, and is a dominant 
natural feature of the area. The species name 
sangha is a noun in apposition. 

Remarks.—It is not normally good prac- 
tice to describe a new species from a sub- 
adult female. However, we have decided to 
establish this taxon in light of the distinct 
nature of the available morphological char- 
acters, and because of the isolated nature of 
the study area which may mean that further 
specimens are unlikely to become available 
for some time. Sudanonautes sangha is the 
eleventh species of this West and Central Af- 
rican genus. Characters of the gonopods, 
adult male chelipeds, abdomen and sternum 
are not available because the only specimen 
of S. sangha is a subadult female. Neverthe- 
less, there are a number of unique characters 
that distinguish S. sangha from other species 
in the genus. 

Sudanonautes sangha is most likely to be 
confused with other large species of fresh- 
water crabs occurring in the rain forests of 
Central Africa such as S$. chavanesii, S. far- 
adjensis, S. africanus, S. aubryi and S. flow- 
Crt: 

Sudanonautes sangha and S. africanus are 
similar in that both species have a small epi- 
branchial tooth, and both lack large teeth on 
the anterolateral margins of the carapace. 
However, there are a number of characters 
which distinguish S. sangha from S. african- 
us. The carapace of S. sangha is not as flat- 
tened as that of S. africanus (ch/fw S. san- 
gha = 1.16, S. africanus = 1.06), and the 
posterior region of the carapace of S. sangha 
is smooth, whereas that of S. africanus is 
rough with warty patches and ridges in the 
posterior region. The cardiac regions of S. 
sangha are flattened and are neither rounded 


417 


nor well marked, whereas these regions in 
S. africanus appear as a pair of distinct 
raised rounded structures that are clearly 
outlined by deep cardiac grooves. The inter- 
mediate tooth on the anterolateral margin be- 
tween the exorbital tooth and the epibran- 
chial tooth of S. sangha is small and low, 
whereas that of S. africanus is large and tri- 
angular. The second carpal tooth on the che- 
liped of S. sangha is very small, only the 
size of a granule, whereas the second carpal 
tooth of S. africanus, while smaller than the 
first carpal tooth, is distinct, pointed, and is 
a tooth rather than a granule. The vertical 
groove on the carapace sidewall of S. san- 
gha meets the anterolateral margin at the 
base of the intermediate tooth, whereas the 
vertical groove of S. africanus meets the an- 
terolateral margin at the base of the epibran- 
chial tooth. 

Sudanonautes sangha can be distin- 
guished from S. chavanesii by the position 
of the postfrontal crest and by the shape of 
the epibranchial teeth. In S. sangha the lat- 
eral ends of the postfrontal crest curve for- 
ward to meet the epibranchial teeth in line 
with the mid groove of the crest, whereas in 
S. chavanesii the lateral ends of the post- 
frontal crest curve sharply backward before 
meeting the epibranchial teeth which are set 
back posterior to the mid groove of the crest. 
In S$. sangha, the epibranchial tooth is small, 
blunt, pointed forward, and positioned in 
line with the mid groove of the crest, where- 
as in S. chavanesii, the epibranchial tooth is 
large, sharp, pointed outward, and positioned 
well behind the mid-groove of the postfron- 
tal crest. Further, in S. sangha the vertical 
suture on the carapace sidewall is simple, 
whereas in S. chavanesii the vertical suture 
on the carapace sidewall forms a Y-shaped 
depression beneath the intermediate tooth. 
Finally, in S. sangha the junction between 
the two segments of the mandibular palp is 
simple, whereas in S. chavanesii there is a 
small hard flap on the mandibular palp at the 
junction between the two segments. 

Sudanonautes sangha can be distin- 
guished from S. faradjensis by the form of 


418 


the anterolateral margin behind the epibran- 
chial tooth: that of S. sangha is raised and 
lined by small granules, whereas in S. far- 
adjensis, there 1s a row of sharp teeth. Su- 
danonautes sangha can be distinguished 
from S. granulatus as follows: the exorbital 
tooth of S. sangha is wide and triangular, 
whereas that of S. granulatus is narrow and 
low, and the anterolateral margin of S. san- 
gha is raised and lined by granules, whereas 
that of S. granulatus is completely smooth. 
Sudanonautes sangha can be distin- 
guished from S. aubryi by differences in the 
postfrontal crest: that of S. sangha meets the 
anterolateral margin at the epibranchial 
tooth, whereas that of S. aubryi meets the 
anterolateral margin behind the epibranchial 
tooth. In addition, the carapace of S. sangha 
is distinctly flatter than that of S. aubryi (ch/ 
fw S. sangha = 1.16, S. aubryi = 1.29). 
Finally, S. sangha can be distinguished 
from S. floweri by differences in the man- 
dibular palp: in S. sangha the junction be- 
tween the two segments of the mandibular 
palp is simple, whereas in S. floweri there is 
a small hard flap on the mandibular palp at 
the junction between the two segments. In 
addition, the carapace of S. sangha is dis- 
tinctly flatter than that of S. floweri (ch/fw 
S. sangha = 1.16, S. floweri = 1.68). 
Ecology.—Collected in a large river (as 
opposed to a small stream), and caught in a 
baited trap. 
Distribution.—Central African Republic, 
a few kilometers upstream from the village 
of Bayanga (02°45'43"N, 16°14'12”E), San- 
gha river. 


Acknowledgments 


Marie Lawrence and James Cordeiro 
(AMNH) are thanked for making this ma- 
terial available to the authors for identifica- 
tion. Alan Harvey (Georgia Southern Uni- 
versity) provided training in computer aided 
biological illustration to CBB. Collection of 
this material was made possible through the 
efforts of the AMNH Center for Biodiversity 
Conservation and the World Wildlife Fund. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


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. 1994. Identification of Sudanonautes aubryi 
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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


113(2):420—425. 2000. 


The freshwater crabs of the Barbilla National Park, Costa Rica 
(Crustacea: Brachyura: Pseudothelphusidae), with notes on the 
evolution of structures for spermatophore retention 


Gilberto Rodriguez and Ingemar Hedstr6m 


(GR) Centro de Ecologia, Instituto Venezolano de Investigaciones Cientificas, Apartado 21827, 
Caracas 1020 A, Venezuela, email:grodrigu @ oikos.ivic.ve; (IH) Mid Sweden University, 


Department of Applied Science, S-871 88 Harnésand, Sweden, email: 
ingemar.hedstrom @tnv.mh.se. 


Abstract.—The Barbilla National Park, a natural area of high biodiversity 
on the Caribbean slope of Costa Rica, possesses two species of freshwater 
crabs that share the same biotope in different localities of the Park. The first 
species, Potamocarcinus magnus (Rathbun, 1896), one of the largest species 
of the family Pseudothelphusidae, is widely distributed in Middle America, 
from Costa Rica to Southern Mexico. The other species is a new species, 
Ptychophallus barbillaensis. This is a species of small crabs, possibly restricted 
to the National Park and neighboring areas. It can be distinguished from all 
other species in the genus by the form of the receptacle formed in the apex of 
the male gonopod, possibly for the retention of spermatophora during copu- 
lation. The species of Ptychophallus Smalley, can be arranged in a morphocline 
according to the relative development of this receptacle, with the present new 
species midway between the ancestral condition and the closed channel found 


in P. goldmanni Pretzmann, 1965. 


The Barbilla National Park covers 12,830 
hectares of humid tropical forest on the Ca- 
ribbean watershed of the Sierra de Tala- 
manca, Costa Rica. This stretch of mostly 
primary vegetation is located between the 
valleys of the Pacuare and Chirrip6o rivers, 
between 200 and 1600 m above sea level. 
The area is noted for its high biodiversity. 

During recent surveys carried out by the 
Mid Sweden University several samples of 
freshwater organisms have been collected 
in the ravines that surround the Nairi Field 
Station, close to the northwestern border of 
the Park. Two species of freshwater crabs 
were discovered among the materials in 
these collections. One of them, a new spe- 
cies belonging to the genus Ptychophallus 
Smalley, 1964a, is described in the present 
contribution. The materials recorded are de- 
posited in the Reference Collection of the 


Instituto Venezolano de Investigaciones 
Cientificas, Caracas (IVIC); the National 
Museum of Natural History, Smithsonian 
Institution, Washington, D.C. (USNM), and 
the Museum of Natural History of Tulane 
University, New Orleans (TU). Other ab- 
breviations used are cl, carapace length, and 
cb, carapace breadth. 


Systematics 


Family Pseudothelphusidae Rathbun, 1893 
Genus Potamocarcinus H. Milne Edwards, 
1555 
Potamocarcinus magnus (Rathbun, 1896) 


Material.—Costa Rica: Rio Cano Seco 
(10°00’N, 83°26'W), affluent of Rio Dantas, 
Barbilla National Park, 7 Jan 1999, leg. I. 
Hedstrom, 300 m above sea level, 1 im- 
mature female, cl 17.3 mm, cb 25.0 mm 


VOLUME 113, NUMBER 2 


(IVIC 1070); Casas Negras, Rio Dantas, 
Barbilla National Park, 8 Jan 1999, leg. I. 
Hedstrém, 200 m above sea level, 1 im- 
mature male, cl 19.8 mm, cb 28.3 mm 
(IVIC 1071); Rio Barbilla, Barbilla Nation- 
al Park, 28 Jan 1999, leg. I. Hedstr6m, 100 
m above sea level, 2 immature males, cl 
17.0 and 12.4 mm, cb 23.7 and 18.5 mm, 
1 immature female, cl 10.0 mm, cb 15.7 
mm (IVIC 1072). 

Remarks.—Although our specimens are 
immature, they can be referred with cer- 
tainty to this species. Potamocarcinus mag- 
nus 1s widely distributed in Central Amer- 
ica, but Rodriguez (1982) distinguish three 
different morphological groups from (a) 
Costa Rica, (b) El Salvador, Guatemala and 
southern Mexico, and (c) Guatemala. The 
characters of the carapace in our specimens 
agree with those given for the Costarican 
form. However, the typical flat tubercles 
present over the surface of the carapace 
cover a greater area, a characteristic possi- 
bly due to the juvenile condition of the 
specimens. The right cheliped has the typ- 
ical long and narrow tubercle over the 
palm, at the base of the fingers, although it 
is relatively litthe developed. In the first 
male gonopods there is not an abnormal in- 
trusion of the marginal process into the dis- 
tal lobe, as was the case in the specimen 
from Costa Rica illustrated by Rodriguez 
(1982, fig. 72b). Potamocarcinus magnus is 
the largest of all species of Pseudothelphu- 
sidae, with a cb of 135 mm reported by 
Rathbun (1896). 


Genus Ptychophallus Smalley, 1964a 
Ptychophallus barbillaensis, new species 
Pig. 1 2C) 


Material.—Costa Rica: Rio Cano Seco 
(10°00’N; 83°26’W), affluent of Rio Dantas, 
Barbilla National Park, 7 Jan 1999, leg. I. 
Hedstr6m, 300 m above sea level, 1 male 
holotype, cl 16.9 mm, cb 27.7 mm (IVIC 
1073), 1 male paratype, cl 13.4 mm, cb 20.7 
mm (IVIC 1074); Nairi Field Station 
(09°59'N; 83°27'W), 3 km north of the bor- 


421 


der of Barbilla National Park, 7 Jan 1999, 
leg. I. Hedstrém, 2 ovigerous females, cl 
15.6 and 13.9 mm, cb 25.0 and 21.1 mm 
(IVIC 1075); Casas Negras, Rio Dantas, 
Barbilla National Park, 8 Jan 1999, leg. I. 
Hedstr6m, 200 m above sea level, 2 males, 
cl 12.2 and 11.9 mm, cb 18.9 and 18.7 mm 
(IVIC 1076); Las Cuevas, Rio Dantas, Bar- 
billa National Park, 11 Jan 1999, leg. I. 
Hedstrom, 150 m above sea level, 2 males, 
cl 14.2 and 11.7 mm, cb 22.5 and 18.6 mm, 
1 mature female, cl 14.1 mm, cb 21.5 mm 
(IVIC 1077); El Recodo, Rio Dantas, Bar- 
billa National Park, 6 Jan 1999, leg. I. Hed- 
strom, 300 m above sea level, 1 male, cl 
15.2 mm, cb 23.3 mm, 1 immature male, cl 
9.9 mm, cb 15.0 mm, 1 male juvenile, cl 
5.8 mm, cb 13.5 mm (IVIC 1078). 

Diagnosis.—First gonopods with large 
lateral lobe divided in 2 subequal rounded 
segments by median notch; proximal seg- 
ment smaller, subcircular; distal segment 
projected anteriorly and bent caudally to 
form cup-shaped receptacle; apex strongly 
bent laterally forming, with distal segment 
of lateral lobe, characteristic sinus; field of 
spines directed toward latero-cephalic side, 
oblong, with deep notch on mesial side; ce- 
phalic end of apex with 2 expansions, distal 
subtriangular in lateral and caudal views, 
with notch on lateral side, proximal round- 
ed in cephalic view, beak-like in mesial 
view. 

Description of holotype.—Carapace 1.6 
times as wide as long, surface smooth and 
polished, except for few granules on pos- 
terior branchial regions, near margins; cer- 
vical grooves recurved backwards, narrow 
and deep, not reaching margins of carapace; 
anterolateral margins with shallow and 
wide postorbital notch bordered by 5 papil- 
lae, rest of borders with small papillae that 
becomes dentiform behind level of cervical 
grooves. Postfrontal lobes low, delimited 
anteriorly by transverse depressions; medi- 
an groove narrow, deep, making incision on 
upper margin of front. Surface of carapace 
between postfrontal lobes and front flat, 
slightly inclined forward and towards mid- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Figw le 


Ptychophallus barbillaensis, new species, holotype from Rio Cafio Seco, Costa Rica (IVIC 1073): 


A, dorsal view of right side of carapace; B, third maxilliped, left; C, chela of largest cheliped (left), external 
view; D, first left gonopod, caudal view; E, same, cephalic view; EK same, apex, mesial view; G, same, apex, 


cephalic view. 


dle line. Upper margin of front in dorsal 
view slightly convex and divided into two 
halves by median notch, thin, well marked, 
with small tubercles; lower margin thin, 
moderately sinuous, advanced in front of 
upper margin; both margins subparallel. 
Exognath of third maxilliped 0.65 length 
of ischium of endognath. Palm of largest 
cheliped moderately swollen, with lower 
and upper margins convex; fingers gaping, 


with rows of small black-brown points on 
external surface. 

First gonopods wide in latero-mesial di- 
rection, narrow in caudo-cephalic direction; 
large lateral lobe divided in 2 subequal 
rounded segments by median notch; proxi- 
mal segment smaller, subcircular; distal 
segment projected anteriorly and bent cau- 
dally to form cup-shaped receptacle semi- 
enclosed by apex of gonopod; apex strongly 


VOLUME 113, NUMBER 2 


bent laterally, forming with distal segment 
of lateral lobe characteristic sinus; field of 
spines directed toward latero-cephalic side, 
oblong, with deep notch on mesial side; ce- 
phalic end of apex forming two expansions; 
distal expansion continuous with margin of 
gonopod, subtriangular, in lateral and cau- 
dal views, with notch on lateral side; prox- 
imal expansion rounded in cephalic view, 
beak-like in mesial view. 

Remarks.—This species can be distin- 
guished from all others in the genus by the 
cupshaped expansion of the distal segment 
of the lateral lobe of the first gonopods and 
by the strong recurvation of the apical pe- 
duncle of gonopod over this expansion. 

Etymology.—The species is named after 
the Barbilla National Park, Costa Rica, 
where the species was collected. 


Evolution of Structures for Spermatophore 
Retention in Ptychophallus 


Although the morphology of the first 
male gonopods are essential for the dis- 
crimination of the species of Pseudothel- 
phusidae, very little is known of the func- 
tion of the different processes of these ap- 
pendages during copulation. In all species, 
the apex is provided with a field of minute 
spines (Smalley 1964b) which surround the 
opening of the spermatic channel and serve 
to hold the spermatophore once it is extrud- 
ed by the piston-like pumping action of the 
second gonopod. 

In the species of Ptychophallus a trend 
can be observed towards a reaccomodation 
of the lateral lobe and the various compo- 
nents of the apex of the gonopod, to form 
a structure that possibly serves to receive 
and keep the spermatophore once it is ex- 
truded. This is fully developed in Ptycho- 
Phallus goldmanni Pretzmann, 1965, but 
the other species can be arranged in a mor- 
phocline comprising several stages of de- 
velopment of this structure, as follows. 

Stage I (Fig. 2A, F).—In the primitive 
condition there is a wide lateral lobe, which 
is foliose, with a shallow depression or 


423 


deeply notched, with the distal portion ad- 
vanced and transverse. The field of spines 
is located distant from the lateral lobe, fac- 
ing caudo-laterally, its long axis forming an 
angle of approximately 60° with the longi- 
tudinal axis of the appendage. Species at 
this stage are P. colombianus (Rathbun, 
1893), P. exillipes (Rathbun, 1898), P. kuna 
Campos & Lemaitre, 1999, and P. tristani 
(Rathbun, 1896). 

Stage II (Fig. 2B, G).—Lateral lobe 
deeply notched as in some species of stage 
I, but he distal portion is more advanced 
and joins the caudal side of the apical pe- 
duncle. The field of spines approaches the 
lateral lobe and is directed laterally, with its 
long axis forming an angle of approximate- 
ly 180° with the longitudinal axis of the ap- 
pendage. Species at this stage are P. micra- 
canthus Rodriguez, 1994, P. montanus 
(Rathbun, 1898), and P. tumimanus (Rath- 
bun, 1898). 

Stage IIIT (Fig. 2C, H).—Lateral lobe as 
in stage II, but the distal portion is cupped, 
and its border is continuous with the caudal 
margin of the apical peduncle. The field of 
spines faces the lateral side, and its long 
axis forms an angle of approximately 230° 
with the longitudinal axis of the appendage. 
The only species at this stage is P. barbil- 
laensis. 

Stage IV (Fig. 2D, I).—The lateral lobe 
has the proximal segment strongly reduced, 
the distal one very advanced and forms, to- 
gether with the border of the field of spines, 
a channel-like receptacle which is wide 
open. The field of spines is directed later- 
ally, its long axis forming an angle of ap- 
proximately 180° with the longitudinal axis 
of the appendage. The only species at this 
stage is P. cocleensis Pretzmann, 1965. 

Stage V (Fig. 2E, J).—The lateral lobe is 
reduced to a distal segment, very advanced 
and forms, together with the border of the 
field of spines, an almost closed channel- 
like receptacle. The field of spines is di- 
rected laterally, its long axis forming an an- 
gle of approximately 180° with the longi- 
tudinal axis of appendage. The only species 


424 


Pier 2. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


a-e 2mm 


iL 


First left gonopod of Ptychophallus species: A, EF P. tristani (Rathbun, 1896), male from ~1.8 km 


E Atenas, San José Province, Costa Rica (TU 4436); B, G, P. montanus (Rathbun, 1898), male from ~3 km N 
San Jer6nimo de Moravia, San José Province, Costa Rica (TU 4442); C, H, P. barbillaensis, new species, 
holotype from Rio Cafio Seco, Costa Rica (IVIC 1073); D, I, P. cocleensis Pretzmann, 1965, holotype from Rio 
Coclé del Norte, Panama (USNM 119869); E, J, P. goldmanni Pretzmann, 1965, holotype from Cana, Panama 
(USNM 54044); A-E, detail of apex, lateral view: F—J, total view, caudal. 


at this stage is P. goldmanni Pretzmann, 
1965. 

A similar morphocline, although not 
leading to the formation of a structure for 
the retention of the spermatophore, has 
been observed in the lateral lobe of species 
of the genera Pseudothelphusa Saussure, 
1857, and Tehuana Rodriguez & Smalley, 
1969, in southern Mexico (Rodriguez 
1986). In this case the morphocline follows 
a general westward direction, suggesting 
an allopatric speciation of primitive demes, 
encompassing a migration along the same 
geographical axis. In the case of Ptycho- 
phallus the pattern is more complex. Spe- 
cies of stages I and II overlap their areas 
of distribution in Costa Rica, and the areas 
of P. kuna (stage I) and P. micracanthus 
(stage II) are relatively close in Central 
Panama (see map in Campos & Lemaitre 
1999). The area of P. barbillaensis (stage 
III) is nested among the species of stages 
I and II. P. cocleensis (Stage IV) is found 


in Central Panama, and P. goldmanni 
(Stage IV) further East, near the Colom- 
bian border. Notwithstanding this complex 
pattern, the general trend of the morphoc- 
line is eastward. This direction contrasts 
with the supposed radiation of the family, 
that proceeded westward from an ancestral 
area in northern Colombia (Rodriguez 
1986). 

Two species of Ptychophallus cannot be 
placed in this morphocline. P. lavallensis 
Pretzmann, 1978, has a field of spines 
strongly upturned cephalically and a very 
wide undivided lateral lobe that does not 
approach the field of spines. P. paraxan- 
thusi Bott, 1968, has a field of spines bent 
mesially and a wide, shallow-notched, lat- 
eral lobe that displays distally several ridg- 
es but does not form a receptacle. These 
species possibly represent phyletic lines 
that differ both among themselves, as with 
those species in the morphocline described 
above. 


VOLUME 113, NUMBER 2 
Literature Cited 


Bott, R. 1968. Fluss-Krabben aus dem 6stlichen Mittel- 
Amerika und von den Grossen Antillen (Crus- 
tacea, Decapoda).—Senckenbergiana Biologica 
49:39—49. 

Campos, M. R., & R. Lemaitre. 1999. Two new fresh- 
water crabs of the genus Ptychophallus Smal- 
ley, 1964 (Crustacea: Decapoda: Brachyura: 
Pseudothelphusidae) from Panama, with notes 
on the distribution of the genus.—Proceedings 
of the Biological Society of Washington 112: 
553-561. 

Milne Edwards, H. 1853. Memoire sur la famille des 
Ocypodien.—Annales des Sciences Naturelles, 
Zoologie, 3e série 20:163—228. 

Pretzmann, G. 1965. Vorlaufiger Bericht tiber die Fam- 
ilie Pseudothelphusidae.—Anzeiger der Mathe- 
matisch Naturwissenschaftliche Klasse der Os- 
terreichischen Akademie der Wissenschaften (1) 
1:1-10. 

. 1978. Neue Potamocarcinini, Poglayen-Neu- 
wall leg 1975 (Vorlaufige Mitteilung).—Sit- 
zungsberichten der Osterreichischen Akademie 
der Wissenschaften, Mathematisch-Naturwis- 
senschaftliche Klasse (1) 1978 (2):51—54. 

Rathbun, M. J. 1893. Descriptions of new species of 
American freshwater crabs.—Proceedings of 
the United States National Museum 16(959): 
649-661. 

. 1896. Descriptions of two species of fresh- 

water crabs from Costa Rica.—Proceedings of 


425 


the United States National Museum 18(1071): 

377-379, fig. 1-3, pl. 29-30. 

. 1898. A contribution to a knowledge of the 
freshwater crabs of America. The Pseudothel- 
phusinae.—Proceedings of the United States 
National Museum 21(1158):507—537. 

Rodriguez, G. 1982. Les crabes d’eau douce 
d’Amérique. Famille des Pseudothelphusi- 
dae.—Faune Tropicale 22:1—223. 

. Centers of distribution of Neotropical fresh- 

water crabs. Jn R. H. Gore & K. L. Heck, eds., 

Biogeography of the Crustacea.—Crustacean 

Issues 3:51—67. 

. 1994. A revision of the type material of some 

species of Hypolobocera and Ptychophallus 

(Crustacea: Decapoda: Pseudothelphusidae) in 

the National Museum of Natural History, Wash- 

ington, D.C. with descriptions of a new species 
and a new subspecies.—Proceedings of the Bi- 

ological Society of Washington 107:296—307. 

,& A. E. Smalley. 1969. Los cangrejos de agua 
de Mexico.—Anales del Instituto de Biologia 
(Mexico) 41:69—112. 

Saussure, H. de. 1857. Diagnoses de quelques Crus- 
tacés nouveaux des Antilles et du Mexique.— 
Revue et Magazine de Zoologie Pure et Appli- 
qué (2)9:304—306. 

Smalley, A. E. 1964a. The river crabs of Costa Rica, 
and the subfamilies of the Pseudothelphusi- 
dae.—Tulane Studies in Zoology 12:5-—13. 

. 1964b. A terminology for the gonopods of the 

American river crabs.—Systematic Zoology 13: 

28-31. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


113(2):426—430. 2000. 


Acanthilia, a new genus of leucosioid crabs (Crustacea: Brachyura) 
from the Atlantic coast of the Americas 


Bella S. Galil 


National Institute of Oceanography, Israel Oceanographic & Limnological Research, P.O.B. 8030, 
Haifa 31080, Israel 


Abstract.—A new monotypic leucosioid genus, Acanthilia, is established 
from the Atlantic coast of the Americas, for one species of the genus /liacantha 
Stimpson, J. intermedia Miers. The new genus differs from /liacantha in its 
tridentate anterior margin of the efferent branchial channel, the shorter cheliped 
fingers, the glabrous pereiopodal dactyls, the convex margins of the sixth ab- 
dominal segment in the male, and the claw-tipped first male pleopod. 


As part of an on-going study of leuco- 
sioidean crabs, the leucosioids of the Atlan- 
tic coast of the Americas at the National 
Museum of Natural History, Smithsonian 
Institution (USNM), were examined. An in- 
vestigation of /liacantha Stimpson, 1871, 
showed that J. intermedia Miers, 1886, dif- 
fers from the five other members of the ge- 
nus in its tridentate anterior margin of the 
efferent channel, the shorter cheliped fin- 
gers, the glabrous pereiopodal dactyls, the 
convex margins of the sixth abdominal seg- 
ment in the male, and the claw-tipped male 
pleopod. J. intermedia was removed from 
the genus /liacantha and placed in a new 
genus herein established. 


Acanthilia, new genus 


Diagnosis.—Carapace subovate, glo- 
bose; regions indistinct. Front narrow, 
prominent, bilobed. Basal antennular seg- 
ment forming an operculum, partially seal- 
ing antennular aperture. Antennular fossa 
not separated from orbit. Orbital margin 
with 3 sutures, basal antennal segment in- 
serted in orbital hiatus proximally on ven- 
tral margin. Postorbital region concave. 
Third maxillipeds fitting closely, sealing 
buccal cavity, efferent branchial channel. 
Inner surface of 3 maxilliped exognath gla- 


brous. Anterior margin of efferent branchial 
channel produced, tridentate, contiguous 
with lower orbital margin. 

Anterolateral margin of carapace medi- 
ally concave, posterolateral margin uni- 
formly curved. Posterior margin of cara- 
pace bidentate. Spine on posterior intestinal 
region upcurved, projecting beyond poste- 
rior denticles. Abdominal sulcus in male 
deep, nearly reaching buccal cavity. 

Chelipeds long; merus_ subcylindrical; 
propodus swollen basally; fingers half as 
long as propodus, opening vertically. Pe- 
reiopods short, dactyls triqueterous, longer 
than propodi, glabrous. 

Male abdomen with segments 3—5 fused, 
tapering distally, basio-lateral regions of 
fused segments somewhat inflated; lateral 
margins of segment 6 convex. Female ab- 
domen with segments 4—6 fused, greatly 
swollen. 

Male first pleopod stout, tip claw-shaped; 
second pleopod short, distally scoop-like. 

Type species.—lIliacantha intermedia 
Miers, 1886. 

Etymology.—Acanthilia is an anagram of 
Iliacantha Stimpson, 1871. 

Remarks.—Stimpson (1871) erected the 
genus Jliacantha for two species, I. subgl- 
obosa and I. sparsa, collected off Florida, 
and characterized by three posterior spines, 


VOLUME 113, NUMBER 2 427 


QoL28 


Fig. 1. Acanthilia intermedia (Miers, 1886).—dé (USNM 274840). A, carapace and right cheliped, dorsal 
view; B, anterior margin of efferent branchial channel; C, sixth abdominal segment and telson, ventral view; D, 
tip of first pleopod, ventral view. A, 5 mm; B,C,D, 0.1 mm. 


428 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 2. Iliacantha subglobosa.—é (USNM 274724). A, carapace and right cheliped, dorsal view; B, anterior 
margin of efferent branchial channel, ventral view; C, sixth abdominal segment and telson, ventral view; D, tip 
of first pleopod, ventral view. A, 5 mm; B,C,D, 0.1 mm. 


VOLUME 113, NUMBER 2 


cheliped fingers opening vertically, and pro- 
jecting pterygostomian channels. When de- 
scribing J. intermedia, Miers (1886:302) 
noted it differed from Stimpson’s species in 
the “‘length of the dactyli of the chelipeds’’. 

Acanthilia, new genus, is easily distin- 
guished from J/liacantha by the tridentate 
margin of the efferent branchial channel 
(Fig. 1B), the cheliped fingers half as long 
as the propodus (Fig. 1A), the glabrous per- 
eiopodal dactyls, the convex margins of the 
male abdominal segment 6 (Fig. 1C), and 
the claw-tipped first male pleopod (Fig. 
1D); whereas in /liacantha the margin of 
efferent channel is bilobate (Fig. 2B), the 
cheliped fingers are longer than the propo- 
dus (Fig. 2A), pereiopodal dactyls are se- 
tose, margins of the male abdominal seg- 
ment 6 are straight (Fig. 2C), and the first 
male pleopod is distally tapering, simple 
(Fig. 2D). 


Acanthilia intermedia (Miers, 1886) 
Fig. 1 


Iliacantha intermedia Miers, 1886:302, pl. 
26, fig. 3—Hay & Shore, 1918:424, pl. 
32, figs. 3, 3a.—Rathbun, 1937:186, pl. 
54, figs. 1, 2.—Williams, 1965:151, fig. 
129; 1984:290, fig. 225.—Coelho, 1970: 
234.—Coelho & Ramos, 1972:184.— 
Fausto-Filho, 1975:81; 1979:51.—G6- 


mez & Ortiz, 1976:8.—Powers, 1977: 
37.—Coelho & Torres, 1980:72.—Ro- 
drigues, 1980:259.—Takeda, 1983: 


118.—Keith, 1985:254, fig. 2f—Abele 
& Kim, 1986:42, fig. 485 b.—Taissoun, 
1988:126.—Werding & Miiller, 1990: 
412, figs. 7a—c.—de Melo, 1996:159, 
fig.—de Melo et al., 1998:450. 

Iliacantha liodactylus Takeda, 
fig. 


1983:159; 


Material examined.—United States. 
North Carolina, 33°20’N, 77°46’W, 25 m, 
16 Aug 1977, 2 66 15.8, 20:0mm, ¢ juv. 
12.2 mm (USNM_ 174250).—Georgia, 
31°41.12’N, 80°20.48’W, 28 m, 30 Jan 
1980, ¢ 21.6 mm (USNM 214909).—Sa- 
pelo Is., 22 m, 5 Mar 1963, coll. M. Gray, 


429 


6 22.5 mm (USNM_ 155484).—Florida, 
25°45.56'N, 82°09.21'W, 19.6 m, 28 Apr 
1981, coll. R. Lemaitre, ovigerous @ 25.1 
mm (USNM 242750). 

Redescription.—Dorsal surface of cara- 
pace granulate, front minutely shagreened. 
Fronto-orbital margin half as wide as pos- 
terior margin, slightly concave. Anterolat- 
eral margin set with prominent pearliform 
granules, subhepatic margin somewhat in- 
flated. Posterior margin bearing pair of dor- 
so-ventrally flattened triangular denticles 
laterally, pearliform granules medially. In- 
testinal spine prominent, granulate, upcur- 
ved. 

Anterior margin of efferent branchial 
channel prominently tridentate, advanced 
beyond orbital margin. Third maxilliped 
bearing fungiform granules, crowded ante- 
riorly, female ischium longitudinally tra- 
versed by row of setae. 

Cheliped merus granulose, granules larg- 
er proximally; carpus and propodus smooth, 
fingers costate. Pereiopodal meri minutely 
granulate, upper margin of propodi promi- 
nently costate, dactyli styliform. 

Thoracic sternites closely granulate. 
Male abdomen minutely granulate, fused 
segments of female abdomen smooth ex- 
cept for granulate basal margin. Male first 
pleopod slightly sinuous, dorso-ventrally 
flattened, subapically setose; cornute tip 
bearing subquadrate plate proximally, claw- 
like appendix distally. 

Color.—“‘gray, without markings of any 
kind”’ (Hay & Shore 1918:424), ‘“‘Amarillo 
crema O gris con puntos pequefnios dispersos 
de color marr6n claro”? (Taissoun 1988: 
126). 

Remarks.—The color plates of J. inter- 
media and I. liodactylus (Takeda, 1983:118, 
119) were erroneously exchanged. 

Distribution.—North Carolina to Brazil; 
10—329 m. 


Acknowledgements 


I am deeply grateful to R. B. Manning 
for the invitation to study the leucosioids in 


430 


the National Museum of Natural History, 
Smithsonian Institution, Washington D.C. 
The visit was supported by the Museum’s 
Collection Improvement funds. 


Literature Cited 


Abele, L. G., & W. Kim. 1986. An illustrated guide to 
the marine decapod crustaceans of Florida. 
State of Florida Department of Environmental 
Regulation, Tallahassee, 760 pp. 

Coelho, P. A. 1970. A distribuigao dos crustaceos de- 
capodos reptantes do norte do Brasil.—Trabal- 
hos do Instituto Oceanograficos, Universidade 
Federal de Pernambuco, Recife, 9/1 1:223—238. 

, & M. A. Ramos. 1972. A constituigao e a 

distribuigao da fauna de decapodos do litoral 

leste da América do sul entre as latitudes de 5° 

N e 39° S.—Trabalhos do Instituto Oceanograf- 

icos, Universidade Federal de Pernambuco, Re- 

cies 132135—2 36: 

, & M. FE A. Torres. 1980. Zoogeografia mar- 
inha do Brazil II. Consideragdes ecolégicas e 
biogeograficas sobre a familia Leucosiidae (De- 
capoda: Brachyura).—Revista nordestina de 
biologia 3(especial):67—77. 

Fausto-Filho, J. 1975. Quinta contribui¢gao ao inven- 
tario dos crustaceos decapodos dos substratos 
de cascalho do Nordeste brasileiro.—Ciencia 
Agronomica 10:109—124. 

. 1979. Crustaceos estomat6podos e decapodos 
dos substratos de areia do nordeste brasileiro.— 
Arquivos de Ciencias do Mar 19:45—56. 

Gomez, O., & M. Ortiz. 1976. Lista de braquiuros cu- 
banos.—Ciencias, Investigaciones Marinas, 
Universidad de la Habana, 25:3—20. 

Hay, W. P, & C. A. Shore. 1918. The decapod crus- 
taceans of Beaufort, N.C. and surrounding re- 
gion.—Bulletin of the United States Bureau of 
Fisheries 35(1915/1916):369—475. 

Keith, D. E. 1985. Shallow-water and terrestrial Brach- 
yuran crabs of Roatan and the Swan Islands, 
Honduras.—Sarsia 70:25 1-278. 

Melo, G. A. S., de. 1996. Manual de identificacgao dos 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Brachyura (caranguejos e siris) do litoral bras- 

ileiro. Pléiade/FAPESP, 604 pp. 

, M. EF A. Torres, & O. Campos. 1998. Mala- 
costraca-Eucarida. Brachyura. Dromiacea and 
Oxystomata. Pp. 439—454 in PS. Young, Cata- 
logue of Crustacea of Brazil. Rio de Janeiro 
Museu Nacional. 

Miers, E. J. 1886. Report on the Brachyura collected 
by H.M.S. Challenger during the years 1873— 
1876.—Report on the scientific results of the 
voyage of H.M.S. “Challenger”, Zoology 
17(49): 1-362. 

Powers, L. W. 1977. A catalogue and bibliography to 
the crabs (Brachyura) of the Gulf of Mexico.— 
Contributions in Marine Science 20(supple- 
ment):1—190. 

Rathbun, M. J. 1937. The oxystomatous and allied 
crabs of America.—United States National Mu- 
seum Bulletin 166:1—278, 86 pls. 

Rodriguez, G. 1980. Los Crustaceos decadpodos de 
Venezuela. Instituto Venezolano de Investiga- 
ciones Cientificas, Caracas, 444 pp., 70 figs. 

Stimpson, W. 1871. Preliminary report on the Crusta- 
cea dredged in the Gulf Stream in the Straits of 
Florida, by L.E de Pourtales, Assist. U. S. Coast 
Survey. Part I. Brachyura.—Bulletin of the Mu- 
seum of Comparative Zoology 2:109—160. 

Taissoun, N. E. 1986-88. Los cangrejos decapodos 
Brachyura de las costas da Venezuela. III. Fam- 
ilias Leucosiidae Dana, 1852 y Geryonidae 
(Beurlen, 1930).—Boletin del Centro de Inves- 
tigaciones Bioldgicas, Universidad del Zulia, 
Maracaibo, 17:121—140. 

Takeda, M. 1983. Crustaceans. In M. Takeda, & T. 
Okutani, Crustaceans and mollusks trawled off 
Suriname and French Guiana. Japan Marine 
Fishery Resource Research Center, Tokyo, 185 
PP. 

Werding, B., & H. G. Miiller. 1990. Leucosiidae von 
der nordkuste Kolumbiens (Crustacea: Deca- 
poda: Brachyura).—Senckenbergiana Biologica 
70:405—417. 

Williams, A. B. 1965. Marine decapod crustaceans of 
the Carolinas.—Fishery Bulletin 65:298 pp. 

. 1984. Shrimps, lobsters, and crabs of the At- 

lantic coast of the eastern United States, Maine 

to Florida. Smithsonian Institution Press, Wash- 
ington, D.C., 550 pp. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


113(2):431—442. 2000. 


Cambarus (Cambarus) davidi, a new species of crayfish 
(Decapoda: Cambaridae) from North Carolina 


John E. Cooper 


North Carolina State Museum of Natural Sciences, Research Lab, 4301 Reedy Creek Road, 
Raleigh, North Carolina 27607, U.S.A. 


Abstract.—Cambarus (Cambarus) davidi is a new species of crayfish from 
the eastern Piedmont Plateau of North Carolina, where it is restricted to inter- 
mittent streams, seepage areas, springs, and burrows. Although the ranges of 
the two species appear to be broadly disjunct, C. (C.) davidi is most closely 
related to the ecologically more tolerant Cambarus (C.) bartonii (s.1.), and 
especially resembles some members of the controversial subspecies C. (C.) b. 
cavatus. Cambarus (C.) davidi is distinguished by a suite of characters that 
includes a vaulted carapace; a deeply excavate, ladlelike rostrum; a very nar- 
row, sparsely punctate areola; and an obtuse to nearly obsolete suborbital angle 
that almost always bears a small tubercle. 


On 18 August 1993, David G. Cooper 
collected several specimens of a Cambarus 
from under large rocks in a shallow, inter- 
mittent tributary of the Neuse River, Wake 
County, North Carolina. In the field they 
appeared to be aberrant individuals of the 
burrowing species, Cambarus (Depressi- 
cambarus) reduncus Hobbs, 1956, which is 
not uncommon in such habitats in the upper 
Neuse River watershed. In the laboratory, 
however, I was surprised to discover that 
the specimens belonged to the subgenus 
Cambarus, and to some species previously 
unknown from the Neuse River basin, 
whose crayfish fauna is well documented 
(Cooper & Ashton 1985, Cooper & Bra- 
swell 1995, Cooper & Cooper 1995). Since 
that time, many additional specimens from 
the Neuse and Cape Fear river basins have 
either been collected or have been recog- 
nized in prior collections. They belong to 
an undescribed species of Cambarus that 
seems to have its closest affinities with con- 
geners that occur in the Tennessee and Ohio 
river drainages. 

Abbreviations used in the text are as fol- 
lows: j, juvenile; NC, North Carolina State 


highway; NCSM, North Carolina State Mu- 
seum of Natural Sciences, Raleigh; PCL, 
postorbital carapace length; R, river; SR, 
State secondary (county) road; TCL, total 
carapace length; US, United States high- 
way; USGS, United States Geological Sur- 
vey; and UTM, Universal Transverse Mer- 
cator coordinates. 


Cambarus (Cambarus) davidi, 
new species 
Fig. 1, Table 1 


Diagnosis.—Body and eyes pigmented, 
eye small (X adult diam 1.7 mm, n = 30). 
Carapace vaulted, thoracic section averag- 
ing 1.3 times wider than deep (n = 52). 
Rostrum acarinate; margins elevated, sub- 
parallel, caudally thickened, strongly to 
moderately constricted at base of acumen, 
lacking marginal spines or tubercles; floor 
(dorsal surface) of rostrum deeply concave, 
ladlelike; acumen 24.5 to 49.1% (X = 
33.6%, n = 52) length of rostrum, latter 
13.0 to 19.3% GG =.16.2% nn: = 52): of 
TCL. Areola 5.2 to 14.5 (X = 8.1, n = 76) 
times longer than wide, constituting 35.4 to 


432 


41.8% (X = 37.6%, n = 76) of TCL and 
42.2 to 47.3% (K = 44.1%, n = 45) of 
PCL; areola sparsely punctate, with 2 (n = 
22) to 3 (n = 64) punctations across nar- 
rowest part. Cervical spines reduced to 
multiple tubercles. Branchiostegal spine re- 
duced to small tubercle; hepatic and sur- 
rounding regions of carapace crowded with 
tubercles. Suborbital angle obtuse to nearly 
obsolete, almost always bearing small tu- 
bercle; postorbital ridge short, cephalic 
margin rounded and usually devoid of tu- 
bercle. Antennal scale 2.0 to 3.6 (X = 2.5, 
n = 50) times as long as broad, widest just 
distal to midlength, lateral margin thick- 
ened and with long distal spine. 

Palm of chela of cheliped 1.5 to 1.8 (X 
= 1.6, n = 51) times wider than deep, width 
1.3 to 1.7 = 1.5, 2 = 51) tames length 
of mesial margin; dorsolateral margin cos- 
tate distally, without impression; mesial 
margin of palm with 2, rarely 3, rows of 
tubercles: mesial row of 6 to 8 (usually 7) 
large, generally adpressed tubercles, sub- 
tended dorsally by row of 1 to 5 (usually 4 
or 5) smaller tubercles. Fixed finger of che- 
la costate laterally, with well defined lon- 
gitudinal ridges dorsally and ventrally; op- 
posable surface of finger with row of 4 to 
11 (usually 5 or 6) tubercles in addition to 
subconical tubercle; dactyl 1.7 to 2.6 (X = 
2.0, n = 51) times as long as mesial margin 
of palm, with strong longitudinal ridge dor- 
sally, weaker ridge ventrally; mesial margin 
with prominent tubercles; opposable sur- 
face with row of 6 to 14 (usually 7 to 9) 
tubercles. Carpus of cheliped generally 
lacking dorsomesial tubercles; merus with 
prominent multiple dorsodistal tubercles 
and often row of small squamous tubercles 
along dorsal ridge. 

Hook on ischium of third pereiopod of 
males, that of form I male (Fig. 1K) uni- 
ramous, overreaching basioischial articula- 
tion and opposed by tubercle on basis; coxa 
of fourth pereiopod of males with vertically 
disposed, caudomesial boss. 

In situ gonopods (Fig. 1G) symmetrical, 
with abutted or slightly separated, tubercle- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


like proximomesial apophyses; proximola- 
teral portion of gonopod set off from rest 
of shaft by weak groove; in lateral aspect 
(Fig. 1B), central projection curved over 
90° to plane of shaft, untapered, with prox- 
imally directed subapical notch; mesial pro- 
cess inflated, symmetrically tapered, slight- 
ly constricted at base of caudal third, with 
subacute, caudally directed apex extending 
slightly farther caudally than apex of cen- 
tral projection; caudal process reduced to 
swelling at caudodistal margin of shaft; in 
mesial aspect (Fig. 1C), distal portion of 
gonopod with essentially flat surface, few 
setae at midlength. 

Annulus ventralis of allotypic female 
(Fig. 1H) 1.3 times broader than long, gen- 
erally symmetrical and subrhomboidal; ce- 
phalic margin convex and fused to sternum, 
caudal margin subangular, free and capable 
of slight movement; cephalic half of annu- 
lus depressed, sloping, with narrow median 
trough, flanked each side by low, poorly de- 
fined ridge; ridges diverging caudally, si- 
nistral ridge continuing obliquely and ter- 
minating before joining caudosinistral wall, 
dextral ridge curving caudodextrally to 
merge with upper arm of heavy, C-shaped 
caudodextral wall; transverse tongue prom- 
inent, originating from sinistral wall near 
caudal midline, continuing cephalically and 
slightly obliquely before turning dextrally 
and plunging into fossa beneath dextral 
wall; sinus dissecting caudal wall near mid- 
line. 

Measurements of type specimens provid- 
ed in Table 1. 

Description of holotypic male, form I._— 
Body and eyes pigmented, eye 1.8 mm 
diam. Cephalothorax (Fig. 1A, D) subcy- 
lindrical, thoracic section 1.2 times wider 
than deep. Areola 9.2 times longer than 
wide, constituting 39.3% of TCL (45.1% of 
PCL), sparsely punctate, with 3 punctations 
across narrowest part. Rostrum acarinate, 
with slightly thickened caudal margins 
moderately converging to base of acumen, 
where moderately constricted; margins of 
acumen slightly concave and converging to 


VOLUME 113, NUMBER 2 


Fig. 1. Cambarus (Cambarus) davidi, new species (all from holotypic male, form I, except E, EK from 
morphotypic male, form II, and H, from allotypic female): A, lateral aspect of carapace; B, E, lateral aspect of 
gonopod (first pleopod); C, EK mesial aspect of gonopod; D, dorsal aspect of carapace; G, caudal aspect of in 
situ gonopods; H, annulus ventralis and postannular sclerite; I, dorsal aspect of distal podomeres of right cheliped; 
J, antennal scale; K, hook on ischium of third pereiopod; L, epistome. 


dorsally directed apical tubercle, which 
reaching just beyond midlength of penulti- 
mate podomere of antennular peduncle; 
acumen comprising 30.2% of rostrum 
length, latter constituting 14.2% of TCL; 


floor of rostrum excavate, ladlelike, mod- 
erately punctate, and ascending caudally 
into broad dorsomedian depression of car- 
apace; subrostral ridge strong, visible to 
base of acumen in dorsal aspect. 


434 


Table 1.—Measurements (mm) of types of Cam- 
barus (Cambarus) davidi, new species. 


Holotyp- Morpho- 
ic Allotypic typic 
male female male 


Carapace 
Total length 30:3. 36:5: 4.3077 
Postorbital length 26:4. 31.6 . 26.6 
Length cephalic section 18.4 22.4 19.0 
Width 19.7 "18:4 15S 
Depth I2AGsl 1529 122 
Length rostrum 4.3 5.6 4.9 
Length acumen ee) 22 i 
Length areola LO 14k ey, 
Width areola 13 jai ino 
Antennal scale 
Length 4.5 5.4 Dye 
Width 1.8 Jecdl 1.8 
Abdomen 
Length 29:6, 37.1 \B50:6 
Width 1326 G4. -g122 
Cheliped 
Length lateral margin chela 24AA 26.7 23:8 
Length mesial margin palm ies 8.8 fis 
Width palm A i ha ad bl | 
Depth palm 6.8 7.8 6.7 
Length dactyl 1555 16:3); .452 
Length carpus LO:0%, 1c) ee) 
Width carpus iy? 8.0 6.7 
Length dorsal margin merus r2:0) 13:87 212.2 
Depth merus afa 8.5 6.9 
Gondopod length 8.4 N/A 8.0 


Postorbital ridge strong, groove essen- 
tially lateral, cephalic margin with vestigial 
tubercle. Branchiostegal spine reduced to 
tubercle; suborbital angle obtuse, without 
tubercle. Thoracic section of carapace dor- 
sally and dorsolaterally punctate, laterally 
with large, scattered granules; cephalic sec- 
tion 1.5 times longer than areola and con- 
stituting 60.7% of TCL, laterally crowded 
with large tubercles and with row of small 
tubercles along ventral margin of cephalic 
section of cervical groove; gastric region 
mostly glabrous. Cervical spine region on 
right with 4 large and 3 smaller tubercles 
(2 large and 2 smaller on left). Abdomen 
slightly narrower and shorter than cepha- 
lothorax. Proximal podomere of uropod 
without spine or tubercle on lateral lobe, 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


with prominent caudomedian spine on me- 
sial lobe; mesial ramus of uropod with me- 
dian keel bearing strong caudal spine, tip of 
which reaching caudal margin of ramus, 
caudolateral margin with small spine; lat- 
eral ramus with submedian keel bearing ter- 
minal spine at transverse flexure, latter with 
total of 17 fixed spines along margin and 1 
long, movable sublateral spine. Telson with 
1 long stationary and 1 long articulated 
spine in each caudolateral corner of ce- 
phalic section; caudal margin domelike. 

Epistome (Fig. 1L) with symmetrical, 
subtriangular cephalic lobe bearing short 
cephalomedian projection; margins of lobe 
uninterrupted, moderately thickened, lateral 
apices thicker than rest; floor (ventral sur- 
face) of lobe punctate and slightly convex, 
concave along lateral margins; transverse 
basal sulcus complete; central depression of 
body broad, moderately deep, with cephal- 
omedian fovea; lamellae punctate, lateral 
margins subtruncate, with 2 large subacute 
tubercles and 1 small rounded tubercle in 
right caudolateral corner (1 large subacute 
and 1 small rounded tubercle on left); zy- 
goma moderately arched, pits elongate. An- 
tennal peduncle with small cephalolateral 
tubercle on basis, small ventral tubercle on 
ischium; antennular peduncle with small, 
laterally displaced subdistal spine on ven- 
tral surface of basal podomere. Antennal 
scale (Fig. 1J) 2.5 times longer than wide, 
broadest just distal to midlength; lateral 
margin thickened, terminating in long distal 
spine, tip of which reaching distal margin 
of penultimate podomere of antennular pe- 
duncle; lamella ca. 1.3 times as wide as 
thickened lateral margin; distal margin of 
lamella subtransverse for most of width, 
then sloping to widest point; mesial margin 
subparallel to lateral margin for most of 
length. 

Third maxilliped with tip of endopodite 
reaching about midlength of penultimate 
podomere of antennal peduncle; exopodite 
hirsute, tip reaching base of distal two- 
thirds of merus of endopodite; cephalola- 
teral corner of ischium slightly produced, 


VOLUME 113, NUMBER 2 


not spinelike; ventrolateral ridge flanked 
mesially by row of punctations bearing 
moderately long setae; lateral half of ischi- 
um with punctations bearing short setae, 
punctations most abundant on proximal 
third; mesial half with long stiff setae large- 
ly obscuring mesial margin, latter with 27 
denticles on right. Right mandible with in- 
cisor ridge bearing 7 denticles (6 on left). 

Right chela (Fig. 11) 2.1 times longer 
than wide; palm 1.7 times broader than 
deep, width 1.6 times length of mesial mar- 
gin; latter 29.9% of chela length, 47.1% of 
dactyl length. Dorsal surface of palm punc- 
tate; distolateral margin of palm and lateral 
margin of fixed finger costate, area at junc- 
ture of palm and finger with aggregation of 
large punctations creating slight impres- 
sion; lateral margin of palm rounded and 
with row of large punctations. Ventral sur- 
face of palm less punctate than dorsal, dis- 
tolateral area with moderate depression and 
aggregation of large punctations; lateral 
eminence of articular ridge with distal tu- 
bercle, none proximal to ridge. Mesial mar- 
gin of right palm with mesial row of 7 ad- 
pressed tubercles, proximal 3 of which with 
elevated distal margins (same on left), me- 
sial row subtended dorsally by row of 4 
smaller tubercles (5 on left) and ventrally 
by 1 small, squamous distal tubercle (3 on 
left). 

Fingers gaping in proximal three-fourths 
of length, greatest width of gape about four- 
fifths width of base of fixed finger; oppos- 
able base of fixed finger with tuft of short 
setae; both fingers slightly curved distov- 
entrally in lateral aspect, dactyl very slight- 
ly bowed in dorsal aspect. Mesial margin of 
right dactyl bearing row of 4 prominent and 
4 weaker tubercles on proximal half, distal 
half of margin punctate; dorsal surface of 
dactyl with low, rounded longitudinal ridge, 
flanked mesially by punctate groove, later- 
ally by row of large, spaced punctations; 
ventral surface with similar longitudinal 
ridge; opposable surface with 8 tubercles, 
fourth from base very large and slightly dis- 
placed ventrally (7 tubercles on left, third 


435 


from base largest); denticles in 2 or 3 rows 
from near tip of finger to sixth tubercle 
from base, single row from there. Fixed fin- 
ger with very strong dorsomedian ridge 
flanked mesially by deep punctate groove 
and second narrower ridge, laterally by 
deep punctate groove; lateral margin with 
irregular row of large punctations; ventral 
surface with very strong longitudinal ridge, 
flanked both sides by row of large puncta- 
tions; opposable margin with subconical tu- 
bercle ventral to denticles at base of distal 
third of finger, and 5 tubercles proximal to 
subconical one, third from base more prom- 
inent than others (4 tubercles on left finger, 
third from base very large); denticles in 2 
or 3 rows from tip of finger to subconical 
tubercle, single row from there. 

Carpus (Fig. 11) 1.4 times as long as 
wide, 1.4 times as long as mesial margin of 
palm; carpus dorsally with long, deep, 
slightly oblique sulcus, lateral and mesial to 
which surface punctate; mesial margin with 
large distal spine and prominent proximal 
tubercle; ventral surface with stout, rounded 
tubercle at lateral articular condyle, similar 
distomedian tubercle, and 1 small tubercle 
proximomesial to latter. Right merus 1.7 
times longer than deep; dorsal surface with 
3 prominent and 2 smaller subdistal tuber- 
cles (2 prominent and 1 smaller on left), 
and small squamous tubercles along much 
of dorsomedian ridge; ventrolateral ridge 
with 2 small acute tubercles and 1 minus- 
cule tubercle near articular condyle (1 small 
acute tubercle and minuscule distal tubercle 
on left); ventromesial ridge with 10 spini- 
form tubercles, distalmost one somewhat 
larger than others; ischium with 3 small 
ventral tubercles (4 on left). 

Palm and fingers of chela of second pe- 
reiopod hirsute. Ventral margins of pleura 
subtruncate or slightly rounded, caudoven- 
tral corners slightly angular, caudal margins 
rounded; terga very punctate, except artic- 
ular cephalic portions glabrous. Sternites 
between third and fourth coxae with very 
dense, matted setae, covering distal ends of 
in situ gonopods. 


436 


Gonopods (Fig. 1B, C, G) as described 
in “‘Diagnosis.’’ Length of gonopod 27.7% 
of: TCL: G8 of PCE): 

Description of allotypic female.—Except 
for secondary sexual characters, differing 
from holotypic male in following respects: 
Areola 6.7 times wider than long, consti- 
tuting 38.6% of TCL (44.6% of PCL). Acu- 
men comprising 39.3% of rostrum length, 
latter constituting 15.3% of TCL. Suborbit- 
al angle nearly obsolete, with small tuber- 
cle. Cervical spine region on both sides of 
carapace with 3 prominent tubercles, largest 
of those on right side subacute. Caudolater- 
al corner of cephalic section of telson with 
2 spines on right, 3 on left. Cephalic lobe 
of epistome subcordiform; lamellae with 
single tubercle each caudolateral corner; zy- 
goma strongly arched. Antennal peduncle 
lacking tubercle on basis; antennal scale 2.6 
times longer than wide, lamella about 1.5 
times as wide as thickened lateral margin. 
Incisor ridge of right mandible bearing 8 
denticles. Right chela 2.0 times longer than 
wide, palm length 33.0% of chela length, 
54.0% of dactyl length. Opposable surface 
of right dactyl with 10 tubercles (9 on left); 
opposable surface of both fixed fingers with 
6 tubercles in addition to subconical one, 
fourth from base largest. Carpus of cheliped 
1.3 times as long as mesial margin of palm; 
merus 1.6 times longer than deep, dorsal 
surface with 2 prominent and 2 smaller sub- 
distal tubercles (same on left); ventrolateral 
ridge with 5 small tubercles (same on left), 
ventromesial ridge with 8 spiniform tuber- 
cles (9 on left). 

Annulus ventralis (Fig. 1H) as described 
in “‘Diagnosis.”’ In addition, first pleopods 
short, reaching caudal margin of annulus 
when abdomen flexed; annulus about 3 
times wider than postannular sclerite, which 
elongate, ventrally domed, punctate. 

Description of morphotypic male, form 
II.—Differing from holotypic male in fol- 
lowing respects: Thoracic section of cara- 
pace 1.3 times wider than deep. Areola 6.2 
times as long as wide, constituting 38.1% 
of TCL (44.0% of PCL). Margins of ros- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


trum less constricted at base of acumen, 
converging at ca. 45°, acumen comprising 
24.5% of rostrum length, latter constituting 
16.0% of TCL. Suborbital angle obtuse and 
with very small tubercle. Cephalic section 
of carapace 1.6 times longer than areola and 
constituting 61.9% of TCL. Cervical region 
on both sides of carapace with 3 prominent 
and 1 moderate tubercle, ventralmost of 
which on right side subacute. Transverse 
flexure of lateral ramus of right uropod with 
16 spines in addition to sublateral one. Lat- 
eral corners of lamellae of epistome with 1 
moderate tubercle on right, 2 on left; zy- 
goma strongly arched. Antennal scale 2.9 
times wider than long, lamella ca. 1.5 times 
width of thickened lateral margin, distal 
margin moderately declivous. Length of 
palm of cheliped 31.5% of chela length, 
49.3% of dactyl length. Mesial margin of 
palm with mesial row of 6 adpressed tu- 
bercles, subtended dorsally on right palm 
by row of 3 moderate tubercles (4 on left) 
and ventrally by 4 small squamous tuber- 
cles (2 on left). Opposable surface of dactyl 
with 9 tubercles on right, 8 on left, basal- 
most largest. Opposable surface of fixed 
finger with 6 tubercles on right in addition 
to subconical one, 7 on left, third from base 
largest. Carpus 1.3 times as long as palm. 
Merus 1.8 times longer than deep; dorsal 
surface of right merus with patch of 5 
prominent subdistal tubercles and 5 smaller 
tubercles proximal to them (2 moderate and 
6 smaller on left); ventrolateral ridge on 
right with 2 subacute and 2 other tubercles 
(2 small, and 1 minuscule articular tubercle, 
on left); ventromesial ridge on right with 10 
small tubercles (9 on left). 

Hook on ischium of third pereiopod 
weak, not overreaching basioischial articu- 
lation, opposed by tubercle on basis; boss 
on coxa of fourth pereiopod moderately de- 
veloped. 

Gonopod (Fig. 1E, F) length 26.1% of 
TCL. In situ gonopods with weak, separat- 
ed proximomesial apophyses; mesial pro- 
cess noncorneous, bulbous, distal surface 
creased near extruded tip; in lateral aspect, 


VOLUME 113, NUMBER 2 


gonopod with weak juvenile suture; central 
projection noncorneous, curved 90° to 
shaft, tapered to subacute tip; mesial pro- 
cess tapered, triangular in outline, tip di- 
rected caudally and inclined slightly disto- 
laterally. Setae on sternites between third 
and fourth coxae not dense. 

Color notes.—Adult ground color usu- 
ally dark olivaceous, sometimes light 
brown with orange or tan overtones. Ce- 
phalic section of carapace often lighter in 
color than thoracic section. Hepatic region 
with orangish midlateral streak just cephalic 
to cervical groove. Margins of rostrum out- 
lined in tan; antennal scale pale orangish- 
tan with dark lateral margin, antennal fla- 
gellae green. Most tubercles, spines, and 
granules of carapace and chelipeds tan to 
orangish. Dorsal surface of cheliped green- 
ish or olivaceous; articular ridges of chela 
pinkish or orangish, ventral surface of chela 
pale grayish-tan. Lateral surface of entire 
propodus (palm and fixed finger) strikingly 
colored, varying from pinkish-tan to 
creamy orange or yellow. Tips of fingers of 
chelipeds pale orange or orangish-tan, color 
not subtended by black band. Proximal po- 
domeres of other pereiopods pale tan to 
light brown with darker mottling, distal po- 
domeres greenish or bluish, fingers of che- 
lae of second and third pereiopods pale 
blue. Cephaloventral structures bluish-gray, 
except epistomal zygoma almost white. An- 
nulus ventralis usually pale, mottled with 
orange in one female. 

Cephalicmost tergite of abdomen with 
transverse dark brown or black rectangular 
band. Lighter colored adults and all juve- 
niles with dark, diagonal blotch each side 
of caudal margin of thoracic section of car- 
apace, blotches extending onto dorsolateral 
surfaces of two adjacent tergites as short, 
curved markings. Series of short, dark bars 
producing interrupted dorsolateral stripe 
each side of abdomen. Other dorsal surfaces 
of abdomen with scattered dark spotting. 
Ventrolateral pleura of abdomen with pale 
pink cephalic area and narrow oblique or 
V-shaped black bar dorsal to it, series of 


437 


these bars producing zigzag lateral stripe 
each side of abdomen. Juveniles generally 
paler than adults, color patterns in most re- 
spects similar but more vivid. 

Type _ locality.—North Carolina, Wake 
County, small intermittent stream entering 
cove along western shore of Falls Lake (im- 
poundment of Neuse River), ca. 1.4 air km 
NW of western end of NC 98 bridge & ca. 
2.6 air km W of Stony Hill (Bayleaf 7.5’ 
USGS quadrangle, UTM zone 17, 3984850/ 
712190). 

The shallow stream, which lies at the 
bottom of a steep ravine in a hardwood for- 
est, is seasonally intermittent and has a 
maximum width of about 1.2 m. All spec- 
imens from this locality were found in the 
mud of shallow residual pools under large 
rocks when water levels were very low. 

Disposition of types.—The holotypic 
male, allotypic female, and morphotypic 
male are in the crustacean collection of the 
NCSM (catalogue numbers NCSM C-4413, 
C-4414, and C-2656, respectively), as are 
paratypes consisting of 2 6 I, 15 ¢ II, 6 j 
S18 iS and 57 <2. 

Range and specimens examined.—Ap- 
parently limited to the upper Neuse and 
Cape Fear river basins in the eastern Pied- 
mont Plateau of North Carolina. Voucher 
specimens (n = 107), all in the crustacean 
collection at NCSM (catalogue numbers in 
parentheses), have been collected at the fol- 
lowing localities. 

Neuse River Basin: Durham Co.; upper 
trib Little R near Durham, 1 ¢ II (C-4742), 
13 Feb 1995, coll. T. Cuffney. Orange Co.; 
small intermittent stream in headwaters 
West Fork Eno R at SR 1358, 2.4 air km E 
of Cam, 3:7 2i(C-3425), 25. Jal! 1995,. coll. 
M.A. Hartman, M.E. Savacool. Wake Co.; 
seep entering small trib New Light Crk 
along SR 1918, 0.3 km SW of jct SR 1909, 
ca. 7.4 air km NNE of Bayleaf, 2 jd (C- 
44), 18 Feb 1976, coll. A.L. Braswell 
(ALB), N. Murdock; type locality, 1 ¢ I 
(C-2656), 1 jd (C-2657), 4 Jul 1994, coll. 
Eee Copper (DGC), JEC, Ligd 5.3. 2)(C- 
2779), 1 j2 (C-2780), 3 jd (C-2781), 18 


438 


Aug 1993, coll. DGC, JEC, 1 6 I (C-4413), 
8. dT 6 G56 “Py Tgln(GE-3795)) 14: Jun 
1997, coll. DGC, 1 2 (C-4553), 20 Jun 
1998, coll. DGC; spring on small trib Low- 
er Barton Crk between SR 1005 & SR 
1844, SW of Bayleaf, 2 32 (C-3055), 6 Apr 
1996;scollS: YirkayDG@asjGlk 2a 248 
(C-3293), 4 Jul 1996, coll. D. DeOliveira 
(DD), DGC; spring on S shore Lower Bar- 
ton Crk, W of SR 1005, ca. 1.6 km NNW 
of ‘center of Bayleaf, 1) d7ie(E-3333), bh! 
Aug 1996, coll. DGC; small spring entering 
Falls Lake, ca. 0.8 km N of entrance of 
Lower Barton Crk into lake, 2 ¢ II, 1 2,3 
j2 (E-5IS Doo twApr: 1999:.-coll...DGE: 
“Raleigh,” 2 2 (C-3143), 28 Nov 1924, 
coll. C.S. Brimley, W.B. Mabee; small in- 
termittent stream entering lake at Schenck 
Forest, Raleigh, 1 ¢ II (C-3471), 25 Oct 
1996, coll. DGC, 1 2 (C-3603), 4 Apr 
1997, coll. DGC; small trib to Richland Crk 
near SW side of Reedy Creek Rd, Schenck 
Forest, Raleigh, 2 2 (C-5077), 10 Apr 
1999, coll. DGC; small stream E of Jeffrey 
Dr off Lake Wheeler Rd, SE of Lake 
Wheeler Raleich, 3.9. Sao 4-371 7).20 
May 1997;:colls DD; DGE;, Ind. 11.546, 4 
j2 (C-3766), 1 2 (C-4414), 24 May 1997, 
coll. DGC; Crabtree Crk below Duraleigh 
Rd, Raleigh, 1 j2 (C-4591), 30 Aug 1998, 
coll. D.A. Jackan. 

Cape Fear River Basin: Alamance Co.; 
spring on trib Toms Crk, Scott farm off SR 
1612, 0.8 km NW of Union Ridge, 1 6 I, 
1 2 in amplexus (C-45), 2 Mar 1976, coll. 
ED. Scott (FDS); burrow at 403 Glen Ra- 
ven Rd, Burlington, 1 6 II (C-3618), 25 Jun 
1993, coll. C. McGrath (CM). Caswell Co.; 
Benton Branch between SR 1103 and 1105, 
S of SR 1100, ca. 1.9 air km SW of town 
of, Stoneycréek; 4)621Ri ij Gein (C789); 
14 May 1975, coll. FDS, JEC. Chatham 
Co.; small, intermittent upper trib New 
Hope Crk at SR 1716, 6.6 km NNE of jet 
US 64, ca. 10.4 air km ENE of center of 
Bynum, | jd (C-3026), 1 Apr 1986, coll. 
D.R. Lenat (DRL), T. MacPherson, 1 j 2 (C- 
3106),,5.-Oct,.1992,.colk \E Miss 7ou (Ez 
3748) U. Feb41993 calls -CMs kid he (C- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


4464), 13 Feb 1998, coll. DRL, D. Penrose 
(DP); upper trib Robeson Crk at US 64, 
Pittsboro, 1 2 (C-4316), 6 Mar 1997, coll. 
DP. Harnett Co.; upper trib Kenneth Crk at 
SR 1447, NE of Rawls, 2 6 II, 1 jd (C- 
2963), 28 Aug 1991, coll. N. Medlin (NM), 
DP. Rockingham Co.; spring on small trib 
Benaja Crk, ca. 0.3 air km ESE of jct SR 
2426 and 242734: Cp iniiss 2) 24252 € 
3104), 1 3 If (C-3105), 3 Jun 1976, coll. 
M.R. Cooper, JEC; drainage ditch in flood- 
plain Haw R, Camp Guilrock, ca. 19.2 air 
km SE of Monroeton, 1 2 (C-4843), Mar 
1998, coll. A.B. Somers & students; Little 
Troublesome Crk at SR 2600, ca. 1.1 air km 
W of Williamsburg, 1 2 (C-4912), 6 Apr 
1998, coll. B. Tracy, NM, L. Eaton, DP. 

Variations.—In addition to those ad- 
dressed in the “Diagnosis,”’ the following 
variations have been recorded. The margins 
of the rostrum are usually abruptly or at 
least moderately constricted at the base of 
the acumen, but in seven specimens the 
margins, while increasing in convergence, 
are not notably constricted. The number of 
prominent cervical tubercles varies from 
one to six (usually three to five), and in 
some specimens at least one of these tuber- 
cles is spiniform. In three individuals, the 
usual small tubercle on the suborbital angle 
is absent, and in several the angle itself is 
subacute. Nearly all specimens have a mi- 
nuscule tubercle or very weak spine on both 
the basis and ischium of the antennal pe- 
duncle, but five lack a tubercle on the is- 
chium and two lack a tubercle on the basis. 
The width of the lamella of the antennal 
scale ranges from approximately 1.1 to 2.0 
(X = 1.4) times the width of the thickened 
lateral margin. The distal margin of the la- 
mella is usually either subtransverse or 
moderately sloping for much of its width, 
but in three females it is strongly declivous 
from the base of the distolateral spine to the 
mesial margin. 

Most individuals have two spines in each 
caudolateral corner of the cephalic section 
of the telson, but seven of them have two 
spines in one corner and three in the other, 


VOLUME 113, NUMBER 2 


and two have two spines in one corner and 
one in the other. The lateral lobe of the 
proximal podomere of the uropod normally 
lacks a spine or tubercle, but 12 specimens 
have a very small spine or acute tubercle 
on this lobe. The spine on the mesial lobe 
of this podomere varies in size from very 
small to moderate, and is absent in one an- 
imal. 

The chela of form I males is longer than 
that of mature form II males and females, 
averaging 81% of TCL. In form II males 
the average is 71.6% and in females it is 
70.8%. The largest tubercle on the oppos- 
able surface of the fixed finger varies from 
the third to the fifth from the base, but in 
most specimens the fourth tubercle is much 
larger than the others. The largest tubercle 
on the comparable surface of the dactyl 
varies from the first to the fifth, but in most 
it is the fourth, and the largest tubercle is 
almost always offset toward the ventral sur- 
face. The dorsal surface of the merus bears 
from one to eight prominent subdistal tu- 
bercles, with the usual number being three 
or four. Most specimens have from two to 
four weaker tubercles just proximal to the 
more prominent ones, and many have squa- 
mous to subsquamous tubercles along at 
least part of the dorsal ridge. The number 
of tubercles on the ventrolateral ridge of the 
merus ranges from two to five (usually 
three), and the distalmost is very small or 
vestigial. The number of tubercles on the 
ventromesial ridge ranges from six to elev- 
en (usually nine or ten), and the distalmost 
is seldom much larger than the largest of 
the others. 

In 17 adult females the width of the an- 
nulus ranges from 1.2 to 1.8 (X = 1.5) 
times its length, and the cephalomedian 
trough varies from moderately wide to 
nearly obliterated. The thick, C-shaped wall 
of the annulus, beneath which lies the deep- 
est part of the fossa, is dextral in 22 fe- 
males, sinistral in eight others. 

The floor (dorsal surface) of the rostrum 
of juveniles and some subadults, while no- 
tably excavate, is often less ladlelike than 


439 


it is in larger, mature animals. Also in these 
smaller individuals, the setae on the oppos- 
able surface of the fixed finger of the che- 
liped are far longer and more dense than 
they are in adults, often filling the space 
between the fingers and obscuring the tu- 
bercles on both. 

Size.—The largest specimens collected 
are two females measuring 50.7 and 50.0 
mm TCL (44.4 and 44.5 mm PCL), both 
from the Haw River subdrainage of the 
Cape Fear River basin. The next largest 
specimen is a form I male, which measures 
42.5 mm TCL (37.3 mm PCL). The largest 
form II male measures 33.4 mm TCL (28.7 
mm PCL). 

Life history notes.—A form I male was 
collected at the type locality on 14 June 
1997, one was taken in Chatham County on 
13 February 1998, and one was found in 
amplexus with a female in Alamance Coun- 
ty on 2 March 1976. No females with at- 
tached ova or young have yet been seen, 
but one measuring 32.5 mm TCL, taken on 
4 April 1997, had all cement glands highly 
developed. 

Crayfish associates.—Seldom have other 
crayfishes been found in the same habitats 
with C. (C.) davidi. At a few localities, 
however, a number of specimens of Cam- 
barus (Depressicambarus) latimanus (Le- 
Conte, 1856), and of at least one of the spe- 
cies in the complex subsumed under Cam- 
barus (Puncticambarus) acuminatus Faxon, 
1884, have been found. At two sites, juve- 
nile C. (D.) reduncus were collected. While 
juvenile Cambarus (Lacunicambarus) di- 
ogenes Girard, 1852, have been taken from 
under cover near the mouth of the stream 
at the type locality, and chimneyed burrows 
of this species may be seasonally abundant 
along the nearby shoreline of Falls Lake, 
this burrowing species has not been found 
with C. (C.) davidi. 

Relationships.—Ilt appears to me that C. 
(C.) davidi has its strongest affinities with 
Cambarus (Cambarus) bartonii (Fabricius, 
1798). Students of American crayfishes, 
however, have long been cognizant of the 


440 


taxonomic perplexities presented by the 
broadly distributed and highly variable pop- 
ulations currently assigned to this species. 
Two subspecies have been described, Cam- 
barus (Cambarus) bartonii cavatus Hay, 
1902, and Cambarus (Cambarus) bartonii 
carinirostris Hay, 1914, but opinions anent 
their validity have for years fluctuated (see 
Faxon 1914, Ortmann 1931, Hobbs 1972, 
1989, Bouchard 1976, Thoma & Jezerinac 
1982, Jezerinac 1985, Fitzpatrick 1983, Jez- 
erinac & Thoma 1984, Jezerinac et al. 
1995, Cooper 2000). The status of C. (C.) 
b. cavatus remains controversial, but C. 
(C.) b. carinirostris has been elevated to 
species status (Thoma & Jezerinac 1999). 
Although it is still difficult at this time to 
establish precise diagnostic parameters for 
these taxa throughout their ranges (what- 
ever those ranges might be), current diag- 
noses must be used in assessing the rela- 
tionships of C. (C.) davidi. 

In his brief description of C. b. cavatus, 
whose type locality is the Powell River 
(Tennessee-Ohio river drainage) near Ta- 
zewell, Claiborne County, Tennessee, Hay 
(1902:435) emphasized its “‘deeply exca- 
vated rostrum,’’ an areola that is “‘narrower 
and more thickly punctate than in C. bar- 
toni bartoni,”’ and a carapace that is “‘more 
nearly cylindrical.’’ Except for the “‘more 
thickly punctate’’ areola, a number of the 
characters displayed by C. (C.) davidi in- 
dicate a possible relationship with the pro- 
genitors of “‘topotypic”’ C. (C.) b. cavatus, 
although their similarities could just as well 
be a result of convergence. In Ohio and 
West Virginia, this putative subspecies ap- 
parently lacks the deeply excavate rostrum, 
and displays other characters that differ 
from those of the “‘typical’’ form (Jezerinac 
1985, Jezerinac et al. 1995). 

The combination of a narrow (but not 
obliterated), sparsely punctate areola, a la- 
dlelike rostrum, and a vaulted, unflattened 
carapace will serve to separate C. (C.) dav- 
idi from all other members of the subgenus 
except some C. (C.) b. cavatus, and some 
upland southern populations currently as- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


signed to C. (C.) b. bartonii (s.1.). Camba- 
rus (C.) davidi differs from C. (C.) b. ca- 
vatus and most populations of C. (C.) b. 
bartonii (s.1.) in its obtuse to nearly obso- 
lete suborbital angle, the shape of its anten- 
nal scale, and the presence of multiple tu- 
bercles on the dorsal surface of the merus 
of the cheliped. In some montane popula- 
tions of C. (C.) b. bartonii (s.1.) the cara- 
pace is relatively vaulted and the rostrum is 
deeply excavate and somewhat ladlelike. 
These populations, however, characteristi- 
cally have areolae that are broader and have 
more punctations across the narrowest part 
than does the areola of C. (C.) davidi. Also, 
in nearly all C. (C.) bartonii (s.1.) the la- 
mella of the antennal scale is much narrow- 
er than it is in C. (C.) davidi, and its distal 
margin is usually quite declivous from the 
base of the lateral spine to the mesial mar- 
gin. In addition, C. (C.) bartonii (s.1.) sel- 
dom displays prominent multiple tubercles 
on the dorsal surface of the merus. 

Roger E Thoma, whose knowledge of 
subgenus Cambarus is extensive, has sug- 
gested that a comparison of C. (C.) davidi 
with the burrower, Cambarus (Cambarus) 
ortmanni Williamson, 1907, would be use- 
ful. That species differs from C. (C.) davidi 
in many ways, including the following: 
Areola of C. (C.) ortmanni generally oblit- 
erated or nearly so, constituting 41.0 to 
44.0% of TCL; suborbital angle obsolete 
and lacking tubercle; cephalothorax mark- 
edly longer than abdomen; subdistal spine 
on mesial margin of carpus of cheliped 
thick and knoblike rather than long and 
acute; cervical region of carapace lacking 
strong, multiple tubercles; ventrolateral 
ridge of merus of cheliped usually lacking 
tubercles; and annulus ventralis and form I 
male gonopod quite different in configura- 
tion. 

Remarks.—Current evidence indicates 
that the range of C. (C.) davidi, limited to 
parts of the eastern Piedmont Plateau in two 
endemic North Carolina river basins, is dis- 
junct from that of other members of the 
subgenus. The nearest known North Caro- 


VOLUME 113, NUMBER 2 


lina populations of C. (C.) bartonii (s.1.) are 
in the mountains and eastern foothills. 
Whether or not C. (C.) davidi is indeed al- 
lopatric, however, will only be revealed by 
more extensive field work throughout the 
Piedmont Plateau. Specimens of subgenus 
Cambarus from seeps and burrows in the 
Dan River subdrainage of the Roanoke Riv- 
er basin resemble C. (C.) davidi in many 
respects, differ in others. The sample size 
of available adult specimens is far too small 
for accurate assignment of the Roanoke ma- 
terial at this time. 

Cambarus (C.) davidi has yet to be found 
in the Tar-Pamlico River basin, whose cray- 
fish fauna is as well known as that of the 
Neuse and is nearly identical (Cooper & 
Braswell 1995). Unfortunately, almost no 
sampling has been done in appropriate hab- 
itats within the Tar-Pamlico basin. 

Etymology.—I take great pleasure in 
naming this species for my son, David 
George Cooper, an avid naturalist who 
brought the species to my attention and who 
spent many hours tromping in mud and 
muck to collect quite a few of the existing 
specimens. 

Suggested vernacular name: Carolina la- 
dle crayfish. 


Acknowledgments 


I am indebted to David G. Cooper for his 
enthusiastic and persistent field work, as 
well as to those other biologists who col- 
lected specimens (their names are provided 
in the section on ‘‘Range and specimens ex- 
amined’’). I also express my sincerest grat- 
itude to Alvin L. Braswell, John E. Cooper, 
Jr., Martha Riser Cooper, and Don Howard, 
for their unstinting assistance. The manu- 
Script was greatly improved by the reviews 
of Roger E Thoma, Rafael Lemaitre, and 
an anonymous referee. Nancy Childs, 
NCSM, provided technical assistance in the 
final preparation of Figure 1. 


Literature Cited 


Bouchard, R. W. 1976. Geography and ecology of 
crayfishes of the Cumberland Plateau and Cum- 


441 


berland Mountains, Kentucky, Virginia, Tennes- 
see, Georgia, and Alabama, Part II: The genera 
Fallicambarus and Cambarus. Pp. 585—605 in 
J. W. Avault, Jr., ed., Freshwater crayfish. Lou- 
isiana State University Division of Continuing 
Education, Baton Rouge, Louisiana, 676 pp. 
Cooper, J. E. 2000. A new species of crayfish of the 
genus Cambarus, subgenus Cambarus (Deca- 
poda: Cambaridae), from the Broad River basin 
of North Carolina.—Journal of the Elisha 
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, & R. E. Ashton, Jr. 1985. The Necturus lewisi 
study: Introduction, selected literature review, 
and comments on the hydrologic units and their 
faunas.—Brimleyana 10:1—12. 
, & A. L. Braswell. 1995. Observations on 
North Carolina crayfishes (Decapoda: Cambar- 
idae).—Brimleyana 22:87—132. 
, & M. R. Cooper. 1995. A new species of cray- 
fish of the genus Orconectes, subgenus Procer- 


icambarus (Decapoda: Cambaridae), endemic 
to the Neuse and Tar-Pamlico river basins, 
North Carolina.—Brimleyana 23:65—87. 

Fabricius, J. C. 1798. Supplementum entomologiae 
systematicae. Hafniae: Proft et Storch, 572 pp. 

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. 

. 1914. Notes on the crayfishes in the United 
States National Museum and the Museum of 
Comparative Zodlogy with descriptions of new 
species and subspecies.—Memoirs of the Mu- 
seum of Comparative Zodlogy at Harvard Col- 
lege 40(8):351—427. 

Fitzpatrick, J. F, Jr. 1983. How to know the freshwater 
crustaceans. William C. Brown Company Pub- 
lishers, Dubuque, Iowa, 227 pp. 

Girard, C. 1852. A revision of the North American 
Astaci, with observations on their habits and 
geographical distribution.—Proceedings of the 
Academy of Natural Sciences of Philadelphia 6: 
87-91. 

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

. 1914. Cambarus bartonii carinirostris Hay. 
Pp. 384-385 in W. Faxon, Notes on the cray- 
fishes in the United States National Museum 
and the Museum of Comparative Zodlogy with 
descriptions of new species and subspecies.— 
Memoirs of the Museum of Comparative Zo- 
Glogy at Harvard College 40(8):351—427. 

Hobbs, H. H., Jr. 1956. A new crayfish of the genus 
Cambarus from North Carolina and South Car- 


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olina (Decapoda, Astacidae).—Journal of the 

Elisha Mitchell Scientific Society 72(1):61—67. 

. 1972. Crayfishes (Astacidae) of North and 

Middle America. Biota of freshwater ecosys- 

tems identification manual No. 9. U.S. Environ- 

mental Protection Agency, Washington, D.C., 

L7S.<pp: 

. 1989. An illustrated checklist of the American 
crayfishes (Decapoda: Astacidae, Cambaridae, 
and Parastacidae).—Smithsonian Contributions 
to Zoology 480: 1-236. 

Jezerinac, R. EF 1985. Morphological variations of 
Cambarus (Cambarus) bartonii cavatus (Deca- 
poda: Cambaridae) from Ohio, with a diagnosis 
of the Ohio form.—Ohio Journal of Science 
85(3):13 1-134. 

, & R. E Thoma. 1984. An illustrated key to 

the Ohio Cambarus and Fallicambarus (Deca- 

poda: Cambaridae) with comments and a new 
subspecies record.—Ohio Journal of Science 
84(3):120-125. 

, G. W. Stocker, & D.C. Tarter. 1995. The cray- 

fishes (Decapoda: Cambaridae) of West Virgin- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


ia.—Bulletin of the Ohio Biological Survey 
New Series 10:1—193. 

LeConte, J. 1856. Descriptions of new species of As- 
tacus from Georgia.—Proceedings of the Acad- 
emy of Natural Sciences of Philadelphia 7:400— 
402. 

Ortmann, A. E. 1931. Crawfishes of the southern Ap- 
palachians and the Cumberland Plateau.—An- 
nals of the Carnegie Museum 20(2):61—160. 

Thoma, R. FE, & R. FE Jezerinac. 1982. New distribu- 
tional records of crayfish (Cambarus and Fal- 
licambarus) from Ohio, including a new sub- 
species record.—Ohio Journal of Science 82(3): 
136-138. 

5 Oe . 1999. The taxonomic status and 
zoogeography of Cambarus bartonii cariniros- 
tris Hay, 1914 (Crustacea: Decapoda: Cambar- 
idae).—Proceedings of the Biological Society 
of Washington 112:97—105. 

Williamson, E. B. 1907. Notes on the crayfish of Wells 
County, Indiana, with a description of a new 
species. Pp. 749-763 in 31st Annual Report of 
the Department of Geology and Natural Re- 
sources of Indiana, 1906. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


113(2):443—457. 2000. 


Two new species of Hyalella (Crustacea: Amphipoda: Hyalellidae) 
from Death Valley National Park, California, U.S.A. 


Adam J. Baldinger, William D. Shepard, and Doug L. Threloff 


(AJB) Museum of Comparative Zoology, Harvard University, 26 Oxford Street, Cambridge, 
Massachusetts 02138-2902, U.S.A.; (WDS) Department of Biological Sciences, 
Sacramento State University, Sacramento, 

California 95819-6077, U.S.A.; (DLT) United States Department of the Interior, National Park 
Service, Death Valley National Park, Death Valley, California, 92328, U.S.A. 


Abstract.—Two new species Hyalella (Hyalella) muerta and Hyalella (Hy- 
alella) sandra are described from Death Valley National Park, California. Hy- 
alella (H.) muerta, the first North American hypogean hyalellid, is blind, lacks 
dorsal mucronations and antenna | is longer than antenna 2. Hyalella (H.) 
sandra collected from nearby epigean waters, also lacks dorsal mucronations, 
but has normal eye pigmentation and antenna 1 is shorter than antenna 2. 
Populations of both species rarely, if ever, coexist together in Death Valley 


National Park. 


Five species in the genus Hyalella occur 
in the continental United States. These are 
Hyalella (Hyalella) azteca (Saussure, 
1858); Hyalella (Hyalella) inermis Smith, 
1874; Hyalella (Hyalella) montezuma Cole 
& Watkins, 1977; Hyalella (Ayalella) tex- 
ana Stevenson & Peden, 1973; Hyalella 
(Hyalella) longicornis Bousfield, 1996. All 
are epigean species, have normal eye pig- 
mentation and have antenna 1 shorter than 
antenna 2. Only H. (H.) inermis and H. (H.) 
longicornis lack dorsal mucronations. 

Examination of recently collected mate- 
rial from Death Valley National Park, Cal- 
ifornia clearly shows the presence of two 
new species of Hyalella, both of which are 
newly described in this paper. 

In the figures, body parts are marked by 
the following abbreviations: A, antenna; 
Gn, gnathopod; LL, lower lip; UL, upper 
lip; Md, mandible; Mx, maxilla; Mxpd, 
maxilliped; P, pereopod; T, telson; U, uro- 
pod; Pl, pleopod; R, right; L, left. Type ma- 
terial is deposited in the Departments of In- 
vertebrate Zoology at the following muse- 
ums: National Museum of Natural History 


(USNM); Museum of Comparative Zoolo- 
gy, Harvard University (MCZ) and Califor- 
nia Academy of Sciences (CASIZ). 


Hyalella (AHyalella) muerta, new species 
Figs. 1-5 


Diagnosis.—Eyes absent. Antenna 1 lon- 
ger than antenna 2. Accessory flagellum ab- 
sent. Pereon lacking dorsal mucronations or 
spines. Sternal gills reduced in size and pre- 
sent on pereonites 3—7. Maxilla 1, inner 
plate narrow with 2 terminal plumose setae. 
Male gnathopod 2, carpus with strong pos- 
terior lobe with 4 submarginal setae; pro- 
podus robust and much larger than male 
gnathopod 1, palm with rectangular tooth 
near hinge of dactylus. Telson rounded, 
with 4 distal marginal setae. 

Description of male.—Body lacking dor- 
sal mucronations or spines. Eyes absent. 
Head cuboidal, equal to pereonites 1 and 2 
in length. Coxae 1—4 enlarged, subquadrate 
with distomarginal setae, coxae 5—6 with 
distinct anterior and posterior lobes. Sternal 
gills small, present on pereonites 3-7. 


444 


Antenna 1, 57% of total body length, pe- 
duncular ratio 1:0.7:0.6, flagellum 9 artic- 
ulate; accessory flagellum absent. Antenna 
2 shorter than antenna 1, 49% of body 
length, peduncle nearly equal in length to 
head, articles 4 and 5 equal in length, fla- 
gellum 8 articulate. 

Upper lip rounded, anterior margin with 
fine setae. Mandibles lacking palp; left in- 
cisor and lacina with 4 teeth, molar normal. 
Right incisor with 5 teeth, outer the largest 
and bifid, lacina with 3 teeth, spine row 
with 2 plumose setae, molar normal, with 
accessory plumose seta. Lower lip large, 
lacking inner lobes. Maxilla 1, inner plate 
narrow with 2 terminal plumose setae, inner 
margin finely setose; outer plate with 8 
strong apical serrate spines; palp vestigial, 
1 articulate. Maxilla 2, plates subequal in 
width; inner plate with 1 marginal and 12 
apical plumose setae, outer margin with fine 
setae distally; outer plate with 10 apical 
plumose setae, both margins with fine setae. 
Maxilliped, inner plate with 3 strong apical 
teeth, inner margin with 3 plumose setae 
distally; outer plate with 7 apical plumose 
setae, with marginal and submarginal plu- 
mose setae; palp 4 articulate, nearly 3X 
length of outer plate, inner margin of article 
2 with distal plumose setae, article 3 with 
distal setae, dactylus triangular with 3 distal 
setae. 

Gnathopod 1, basis elongate broadening 
distally, posterior lobe of carpus with 6 
marginal setae; propodus rectangular, wider 
and longer than carpus; palm and hind mar- 
gin equal in length, palm with marginal and 
submarginal setae. Gnathopod 2, basis elon- 
gate; posterior lobe of carpus strong with 4 
submarginal setae; propodus robust and 
much larger than that of gnathopod 1, distal 
anterior margin with triangular projection; 
palm with rectangular tooth near hinge of 
dactylus, with marginal and submarginal 
setae. Pereopods 3 and 4 similar in shape 
and size; carpus of both with distal anterior 
lobe and 2 setae. Pereopod 5, approximate- 
ly 70% length of pereopod 6. Pereopod 7 
slightly larger that pereopod 6. Pereopods 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


5—7 bases expanded posteriorly, stronger in 
pereopod 7; carpus with distal posterior 
lobe, with 3—4 spines. 

Pleopods 1-3 long and slender, pedun- 
cles with 2 coupling hooks, rami with plu- 
mose setae. Uropod 1, peduncle outer mar- 
gin with 4 bifurcate spines, inner margin 
with distal marginal seta; inner ramus with 
4 apical spines; outer ramus with 5 apical 
and 2 marginal spines. Uropod 2, 60% the 
length of uropod 1, peduncle with 2 mar- 
ginal spines and 1 marginal seta; inner ra- 
mus with 6 apical spines; outer ramus with 
5 apical spines. Uropod 3, peduncle with 3 
distal marginal spines; ramus with 2 apical 
spines and 1 apical seta. Telson rounded 
with 4 distal marginal setae. 

Female.—All features same as male ex- 
cept as noted below. Antenna 1, 43% of 
total body length; flagellum 9 articulate. 
Antenna 2, 34% of total body length; fla- 
gellum 9 articulate. Gnathopod 2 and pe- 
reopods 3—5 with oostegites. Gnathopod 1 
resembling male, posterior lobe of carpus 
with 8 marginal setae. Gnathopod 2, basis 
elongate; posterior lobe of carpus with 6 
submarginal setae; propodus longer than 
wide, palm and hind margin distinct. 

Etymology.—The specific epithet is de- 
rived from the Spanish muerta as a noun in 
apposition for the word death referring to 
Death Valley. 

Material examined.—Male holotype, 
3.28 mm, USNM 230433, Texas Spring, 
just uphill of the Texas Spring Camp- 
ground, Death Valley National Park, Inyo 
County, California, William D. Shepard, 20 
Jul 1994. Female (ovigerous) allotype, 3.28 
mm, USNM 230434, same data as holo- 
type. Male paratype, 3.28 mm, USNM 
230435, same data as holotype. Female 
(ovigerous) paratype, 3.24 mm, USNM 
230436, same data as holotype. 6 males, 8 
females (4 ovigerous) paratypes, USNM 
230437, same data as holotype. Male (4.00 
mm) and female (3.60 mm), paratypes, 
USNM 230438, Texas Spring tunnel, Death 
Valley National Park, Inyo County, Cali- 
fornia, 36°27'27.54"N,  116°50'14.44’W, 


VOLUME 113, NUMBER 2 


Perea ae 


Es an Ae 


Fig: 1. 


Doug L. Threloff, 19 Dec 1997. Paratypes, 
MCZ 25390, CASIZ 121603, Texas Spring 
tunnel, Death Valley National Park, Inyo 
County, California, 36°27'27.54’N, 
116°50'14.44"W, D. L. Threloff, 7 Dec 
if. Paratypes, MCZ 25391, CASIZ 
121604, Texas Spring tunnel, at discharge 
point, Death Valley National Park, 
Inyo County, California, 36°27'26.18’N, 
116°50'16.93"W, D. L, Threloff, 19 Dec 
Tu9T: 

Remarks.—Hyalella (H.) muerta is mor- 
phologically similar to Hyalella (Mesohy- 
alella) anophthalma Ruffo, 1957 and Hy- 
alella (Mesohyalella?) caeca Pereira, 1989; 
the former is known from a cave in Vene- 
zuela and the latter from Brazil. Both spe- 
cies have sternal gills on pereonites 2—7 and 
antenna 2 is longer than antenna 1. In con- 
trast, H. (H.) muerta has sternal gills on 
pereonites 3—7 and antenna | is longer than 
antenna 2. 


445 


Hyalella (Hyalella) muerta, male, 3.28 mm, USNM 230435. 


Hyalella (Hyalella) sandra, new species 
Figs. 6—10 


Diagnosis.—Eyes present. Antenna | 
Shorter than 2, flagellum 10-11 articulate; 
accessory flagellum absent. Antenna 2 
elongate, flagellum 20—24 articulate. Pere- 
on lacking dorsal mucronations or spines. 
Sternal gills on pereonites 3—7, approxi- 
mately % the size of coxal gills. Maxilla 1, 
inner plate narrow with 2 terminal plumose 
setae. Male gnathopod 2, carpus with strong 
posterior conical lobe with marginal setae. 
Uropod 3, ramus with 7 distal spines. Tel- 
son, rounded with 2 distal submarginal 
spines. 

Description of male.—Body lacking dor- 
sal mucronations or spines. Eyes pigment- 
ed. Head cuboidal, subequal to pereonites | 
and 2 in length. Coxae 1-3 enlarged, quad- 
rate with distomarginal setae, coxae 5—6 
with distinct anterior and posterior lobes. 


446 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


sal PP 


Rigs. 


Hyalella (Hyalella) muerta, male, 3.28 mm, USNM 230435. 


VOLUME 113, NUMBER 2 


Sternal gills on pereonites 3—7, approxi- 
mately % the size of coxal gills. 

Antenna 1, 25% of total body length, pe- 
duncular ratio 1:1:0.8, flagellum 11 articu- 
late each with small aesthetasc-like spines; 
accessory flagellum absent. Antenna 2 
much longer than antenna 1, 50% of body 
length, peduncle articles 3 and 4 equal in 
length to 5, flagellum 20 articulate. 

Upper lip rounded, anterior margin with 
fine setae. Mandibles lacking palp, molars 
normal with accessory plumose seta; left in- 
cisor and lacina with 5 teeth, spine row with 
3 plumose setae. Right incisor with 5 teeth, 
lacina with 3 teeth. Lower lip large, setose 
and lacking inner lobes. Maxilla 1, inner 
plate narrow, with 2 terminal plumose se- 
tae; outer plate with 9 strong apical serrate 
comb-like spines; palp vestigial, 1 articu- 
late. Maxilla 2, plates subequal in width; 
inner plate with 2 submarginal spines and 
apical plumose setae; outer plate with apical 
plumose setae. Maxilliped, left inner plate 
with 2 strong apical teeth, right plate with 
3; left outer plate with submarginal and api- 
cal plumose setae; palp 4 articulate, dacty- 
lus with two apical spines. 

Gnathopod 1, basis elongate; posterior 
lobe of carpus with 8 marginal setae; pro- 
podus equal in length to carpus; palm with 
submarginal setae, proximoposterior corner 
with distinct spine. Gnathopod 2, basis 
elongate; carpus with strong posterior con- 
ical lobe with marginal setae; propodus ro- 
bust and much larger than that of gnatho- 
pod 1; palm with marginal spines and setae, 
proximoposterior corner with tooth like 
spine. Pereopods 3 and 4 similar in shape 
and size; coxal plate of pereopod 4 su- 
bquadrate and much larger than that of pe- 
reopod 3; merus on both with distal anterior 
lobe with 3 and 2 spines respectively. Pe- 
reopod 5, approximately 75% the length of 
pereopod 6; basis expanded posteriorly with 
marginal and submarginal setae. Pereopod 
7 slightly smaller than pereopod 6. Pereo- 
pods 5-7 bases, anterior margins spinose, 
posterior margins expanded posteriorly, 


447 


much stronger in pereopod 7; merus and 
carpus of each distal posterior lobe spinose. 

Pleopods 1—3 long and slender, pedun- 
cles with 3 coupling hooks, except pleopod 
3 with 2, rami with plumose setae. Uropod 
1, peduncle with outer marginal with 4 bi- 
furcate spines, both margins with distal 
spine; inner ramus, with 2 marginal and 4 
apical spines; outer ramus with 3 marginal 
and 5 apical spines. Uropod 2, 75% the 
length of uropod 1, peduncle outer margin 
with 4 marginal spines, inner margin with 
a strong distal spine; inner and outer rami 
with 3 marginal and 4 apical spines. Uro- 
pod 3, peduncle with 3 distal marginal and 
1 submarginal spines; ramus with 7 apical 
spines. Telson rounded, with 2 distal sub- 
marginal spines and 5 small submarginal 
setae. 

Female (ovigerous).—All features same 
as male except as noted below. Antenna 1, 
27% of total body length; flagellum 7 artic- 
ulate. Antenna 2, 63% of total body length 
flagellum 14 articulate. Gnathopod 2 and 
pereopods 3—5 with oostegites. Gnathopod 
2, basis elongate; carpus longer than wide, 
lacking posterior conical lobe and with 9 
submarginal setae; palm and hind margin 
distinct. 

Etymology.—The specific epithet is a 
name in apposition in reference to the first 
author’s wife Sandra, for her support and 
encouragement towards his research en- 
deavors. 

Material examined.—Male holotype, 
4.80 mm, MCZ 25392, Travertine Spring, 
approximately 1.9 km southeast of Texas 
Spring, Death Valley National Park, 
Inyo County, California, 36°26'28.40’N, 
116°49'57.01"W, D. L. Threloff, 21 Dec 
1997. Female (ovigerous) allotype, 3.36 
mm, MCZ 25393, same data as holotype. 
Male paratype, MCZ 25394, same data as 
holotype, 4.88 mm. Female (ovigerous) 
paratype, 3.36 mm, MCZ 25395, same data 
as holotype. Male paratype, 4.80 mm, MCZ 
25435, same data as holotype. Male para- 
type, 4.72 mm, MCZ 25396, Texas Spring 
outflow, 34 m downstream of discharge 


448 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Ut 


Fig. 3. Hyalella (Hyalella) muerta, male, 3.28 mm, USNM 230435. 


point, Death Valley National Park, Inyo 25397, Texas Spring outflow, 13 m down- 
County, California, 36°27'25.44"N, | stream of discharge point, Death Valley Na- 
116°50'18.01"W, D. L. Threloff, 19 Dec tional Park, Inyo County, California, 
1997. Male paratype, 5.20 mm, MCZ 36°27'25.86"N, 116°50'17.20"W, D. L. 


VOLUME 113, NUMBER 2 


449 


Fig. 4. Hyalella (Hyalella) muerta, male, 3.28 mm, USNM 230435: Gnl, Gn2. Female, 3.24 mm, USNM 


230436: Gnl, Gn2. 


Threloff, 21 Dec 1997. Paratypes, USNM 
230439, CASIZ 121605, same data as ho- 
lotype. Paratypes, MCZ 25398, USNM 
230411, CASIZ 121606, Texas Spring out- 
flow, 34 m downstream of discharge point, 
Death Valley National Park, Inyo County, 
California, 36°27'25.44’N, 116°50'18.01’W, 
D. L. Threloff, 19 Dec 1997. Paratypes, 
MCZ 25399, USNM 230440, CASIZ 


121607, Texas Spring outflow, 13 m down- 
stream of discharge point, Death Valley Na- 
tional Park, Inyo County, California, 
IO LT 25.86 0 LIS 3017.20" WW,” D.*L. 
Threloff, 21 Dec 1997. 
Remarks.—Hyalella (H.) sandra is mor- 
phologically similar to H. (H.) longicornis, 
which is larger in size and is known from 
only the type locality in Utah. Examination 


450 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 5; 


of the type specimens of H. (H.) longicornis 
showed that the most significant differences 
between H. (H.) sandra and H. (H.) longi- 
cornis are: the ramus of uropod 3, H. (H.) 
sandra robust, with 7 apical spines and H. 
(H.) longicornis slender, with 4—5 apical se- 
tae; the telson, H. (H.) sandra with 2 sub- 
marginal spines and H. (H.) longicornis 
with 2 long slender apical setae; the flagel- 
lar articles of the male antenna 2, H. (H.) 
sandra with 20—24 and H. (H.) longicornis 
with 16; the palp of maxilla 1, H. (H.) san- 
dra vestigial, 1 articulate and H. (H.) lon- 
gicornis tall, length >2X the width; basis 
of gnathopod 2 posterior margin, H. (H.) 
sandra with 1—3 marginal setae and H. (H.) 
longicornis with 9 marginal setae. 
Discussion.—In 1874, Smith established 
the genus Ayalella, describing Hyalella 
dentata and HAyalella inermis, both from 
Colorado. Faxon (1876), working on South 
American hyalellids, regarded the genus AI- 
lorchestes as the senior synonym of Hy- 


Hyalella (Hyalella) muerta, male, 3.28 mm, USNM 230435. 


alella and determined H. inermis was a va- 
riety of H. dentata, calling it Allorchestes 
dentata var. inermis. Nearly 30 years later, 
Stebbing (1903) resurrected Hyalella and 
established H. inermis as a valid species, 
noting differences in the antennae, mouth- 
parts, gnathopods and pereopods. Weckel 
(1907), in re-examining the North Ameri- 
can hyalellid species, concluded that H. 
inermis, H. dentata and Hyalella faxoni 
Stebbing, 1903 were all junior synonyms of 
Hyalella knickerbockeri Bate, 1862 and 
mentioned that only H. azteca var. inermis 
lacked dorsal mucronations. Later, Barnard 
(1958) provided a list of the Hyalella spe- 
cies and listed H. faxoni and H. knicker- 
bockeri as junior synonyms of H. azteca 
and considered H. inermis a valid species. 
However, Bousfield (1958, 1973) conclud- 
ed that H. azteca is a single morphologi- 
cally variable species with the number of 
dorsal micronations varying from 1-3, that 
specimens totally lacking dorsal mycrona- 


VOLUME 113, NUMBER 2 


Fig. 6. Hyalella (Hyalella) sandra, male, 4.88 mm 


tions were H. azteca forma inermis, and 
that both taxa occurred throughout the Unit- 
ed States. Stevenson & Peden (1973) then 
described HAyalella texana from Texas, a 
species that coexisted with H. azteca. 
Shortly thereafter, Cole & Watkins (1977) 
described Hyalella montezuma from the 
Montezuma Well system in Arizona, but 
this species coexisted with H. azteca forma 
inermis. Holsinger (1981) provided a list of 
the 32 species of Hyalella and mentioned 
that most workers agreed that H. dentata, 
H. inermis and H. knickerbockeri are syn- 
onyms of H. azteca. Lastly, Bousfield 
(1996) described Ayalella (Ayalella) lon- 
gicornis that lacked dorsal mucronations 
and was known only from Utah [although 
table 1 of Bousfield (1996) gives distribu- 
tion as ““Texas’’]. 

Bousfield (1996) divided Hyalella into 
three subgeneric groups (Hyalella, Austroh- 
yalella, and Mesohyalella) based on geo- 
graphical distributions and morphological 
characters (i.e., body mucronations, the pro- 


45] 


, MCZ 25394. 


podus of gnathopods 1 and 2, rami of uro- 
pods 1 and 3, ornamentation of the telson). 
The plesiomorphic subgenera Austrohyalel- 
la and Mesohyalella are confined to conti- 
nental South America while the more apo- 
morphic subgenus Hyalella is endemic to 
the West Indies, Central and North America 
(Bousfield 1996). However, Bousfield 
(1996) mentions that northern distributed 
species of Mesohyalella show morphologi- 
cal similarities to species in the North 
American subgenus Hyalella. 

Both species described here exhibit some 
morphological characters similar to species 
in the subgenus Mesohyalella [H. (H.) 
muerta, smooth body, | plumose seta on the 
inner plate of maxilla 2; H. (H.) sandra, 
smooth body, 1 submarginal spine on the 
inner plate of maxilla 2, spines on the tel- 
son]. However, the presence of 5 sternal 
gills, the morphology of male gnathopod 1 
[H. (H.) muerta, propodus with 5 week and 
short facial setae, palm margin convex and 
short; H. (H.) sandra propodus with 5 week 


452 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


AUS vy Y 
ALAA AL 

y iy! A} f 
14% 
By) 


Wt, 
v4 Mp 
KY 


Fig. 7. Hyalella (Hyalella) sandra, male, 4.88 mm, MCZ 25394. 


and short facial setae, palm margin convex 
and short, palmer angle with 1 short spine] 
and the morphology of the female gnatho- 
pod 2 of both species (propodus long and 


slender) would place them in the subgenus 
Hyalella. In addition, as both new species 
lack copulatory spines on uropod 1, have a 
peduncle and ramus of uropod 3 subequal 


453 


VOLUME 113, NUMBER 2 


U1 


Fig. 8. HAyalella (Hyalella) sandra, male, 4.88 mm, MCZ 25394: Al, A2, U3, T, Pll. Male, 4.80 mm, MCZ 
ga4oa. U1, U2. 


454 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 9. Hyalella (Hyalella) sandra, male, 4.88 mm, MCZ 25394: Gnl, Gn2. Female (ovigerous), 3.36 mm, 
MEZ 25395: Gn, Gn2; 


VOLUME 113, NUMBER 2 


455 


Fig. 10. Ayalella (Hyalella) sandra, male, 4.88 mm, 


in length, and have setae on their telson 
support their subgeneric placement. Lastly, 
the North American distribution that defines 
the subgenus Hyalella as suggested by 
Bousfield (1996) are consistent with the 
known distribution of the two new species 
described here. 

HAyalella (H.) muerta and H. (H.) sandra 
occur in Death Valley National Park, Cali- 
fornia but rarely if ever coexist together. 
Hyalella (H.) muerta, the first North Amer- 
ican hypogean hyalellid, is blind, lacks dor- 
sal mucronations and antenna | is longer 
than antenna 2. Hyalella (H.) sandra, col- 
lected from epigean waters near Texas 
Spring, also lacks dorsal mucronations but 
has normal eye pigmentation and antenna | 
is shorter than antenna 2, as in the other 
North American Hyalella species. These 
two new species bring the total number of 
North American Hyalella taxa to seven. 

Specimens of H. (H.) muerta were col- 
lected in an artificial tunnel that was exca- 
vated in the 1930’s. The tunnel was most 
likely excavated in an effort to increase the 


MCZ 25394. 


volume of water that was being diverted as 
a potable water supply. Prior to the devel- 
opment of the tunnel, Texas Spring is be- 
lieved to have issued water directly from 
the local hillside. The interior of the tunnel 
is typically 1.5 m wide by 1.5—2.0 m high 
and is approximately 70 m in length. Flow 
into the upstream portion of the tunnel de- 
velops as water exits a fractured rock zone. 
The water runs along the floor of the tunnel 
in a stream that is 60—150 cm wide and 3— 
30 cm deep. Overburden above the tunnel 
consists of soft silt, 1-10 m thick. Collapse 
of the tunnel has been prevented through 
the installation of thick wooden cross mem- 
bers that support the ceiling and side wall 
surfaces of the tunnel. Specimens of H. (H.) 
muerta were found among the submerged 
roots originating from surface-inhabiting, 
riparian plants. In 1995, a one meter portion 
of the wooden tunnel structure collapsed. It 
is not known if the concurrent sediment in- 
put had any impact on the amphipod pop- 
ulation. 

At Texas Spring the water emerges from 


456 


local exposures of gravel and sand (Pistrang 
& Kunkel 1958). The rate of flow has al- 
ways been low, ranging from 0.2-0.5 cfs. 
Because Texas Spring provides water for 
human consumption, the water quality has 
been regularly tested. Miller (1977) provid- 
ed the following water chemistry analysis 
for Texas Spring water: temperature: 31°C; 
Silica: 25—40 mg/l; calcium 30 mg/l; sodi- 
um: 150 mg/l; bicarbonate 330 mg/l; dis- 
solved solids: 600—700 mg/l; pH: 7.5—8.5. 
Specimens of H. (A.) sandra were col- 
lected from Travertine Spring approximate- 
ly 1.9 km southeast of Texas Spring and 
13—34 m down stream from the Texas 
Spring discharge. In mid to late 1970’s, the 
potable water collection system at Texas 
Spring was replaced, and the entire spring 
flow was placed in a PVC pipe in an effort 
to eliminate the percolation of water into 
the ground. Between 1989 and 1994, Death 
Valley National Park maintenance person- 
nel diverted some of the piped water back 
onto the ground in an effort to re-establish 
a stream habitat. Maintenance personnel 
then transplanted benthic sediment and veg- 
etation from Travertine Spring to an area 
down stream of Texas Spring tunnel with 
the intentions of reinoculating the stream 
with aquatic invertebrates and plants. Pre- 
sumably, H. (H.) sandra was transported 
with the sediments that established an in- 
troduced population. Extensive sampling 
has revealed one live specimen of H. (H.) 
muerta occurring in the surface stream 
downstream of Texas Spring tunnel. Prelim- 
inary investigations therefore suggest that 
H. (H.) muerta and H. (H.) sandra rarely 
coexist in Death Valley National Park. 
Although Death Valley is one of the dri- 
est and hottest deserts in the New World, 
the climate there has not always been so 
harsh. During the Pleistocene the climate 
was cooler and wetter, similar to that found 
today around Lake Mono, 240 km (150 mi) 
to the north. Numerous large pluvial lakes 
occupied the many depressions in this area 
and at that time Texas Spring would have 
been at or only slightly above the shoreline 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


of pluvial Lake Manly. The aquatic com- 
munities currently found in the springs and 
streams of Death Valley are largely relicts 
of these Pleistocene and earlier communi- 
ties (Shepard 1992, 1993). Grayson (1993) 
provides an excellent account of both Pleis- 
tocene and Recent hydrology for the Death 
Valley area. 

In the desert southwest of the United 
States, it appears that H. (H.) azteca has 
been giving rise to new species via popu- 
lations that have been isolated in thermally 
constant waters. Thomas et al. (1994, 1997) 
have proposed ecological isolation for the 
species pair H. (H.) azteca and H. (H.) 
montezuma, based on DNA and behavior. 
Their behavioral studies led them to sepa- 
rate two lineages of Hyalella in north-cen- 
tral Arizona; swimmers that inhabit sub- 
mersed vegetation in lakes and clingers that 
inhabit springs dominated by emergent 
macrophytes. Jackson (1912) also noted 
two distinct locomotion behaviors. Hyalella 
(H.) muerta and H. (H.) sandra both fall 
into the clinger behavior category and like- 
ly have speciated from local epigean pop- 
ulations of H. (H.) azteca. 

Another undescribed Hyalella (Hyalella) 
species has also been found in Ash Mead- 
ows National Wildlife Refuge, 45 km to the 
east (in prep.). It also occurs in a warm 
spring. Because of the discovery of a num- 
ber of new species in such a small area, we 
suggest that aquatic biologists more care- 
fully collect and identify specimens in the 
future, particularly when dealing with ther- 
mally constant waters. 


Acknowledgments 


We thank Death Valley National Park for 
permission to collect. We also thank M. Zu- 
bowski (Royal Ontario Museum) for locat- 
ing the types of H. (A). longicornis and C. 
Serejo (Museu Nacional UF Rio De Janei- 
ro) for the loan of H. caeca and help in 
obtaining literature. The first author thanks 
M. E Gable (Eastern Connecticut State Uni- 
versity) for bringing this project to his at- 


VOLUME 113, NUMBER 2 


tention, E. A. Lazo-Wasem (Peabody Mu- 
seum of Natural History, Yale University) 
for helpful comments and discussion, and 
A. B. Johnston (MCZ) for the use of the 
departmental microscope. 


Literature Cited 


Barnard, J. L. 1958. Index to the families, genera and 
species of the Gammaridean Amphipoda (Crus- 
tacea).—Allan Hancock Foundation Publica- 
tions, Occasional Paper Number 19:1—145. 

Bate, C. S. 1862. Catalogue of the specimens of am- 
phipodous Crustacea in the collection of the 
British Museum, London, 399 pp. 

Bousfield, E. L. 1958. Fresh-water amphipod crusta- 
ceans of glaciated North America.—Canadian 
Field Naturalist 72:55—113. 

1973. Shallow-water Gammaridean amphi- 

pods of New England. Cornell University Press, 

Ithaca, New York, 312 pp. 

. 1996. A contribution to the reclassification of 
neotropical freshwater hyalellid amphipods 
(Crustacea: Gammaridea, Talitroidea).—Bollet- 
tino del Museo Civico di Storia Naturale de Ve- 
rona 20(1):175—224. 

Cole, G. A., & R. L. Watkins. 1977. Hyalella monte- 
zuma, a new species (Crustacea: Amphipoda) 
from Montezuma Well, Arizona.—Hydrobiolo- 
gia 52(2—3):175—-184. 

Faxon, W. 1876. Exploration of Lake Titicaca by Al- 
exander Agassiz and S. W. Garman. IV. Crus- 
tacea.—Bulletin of the Museum of Comparative 
Zoology 3:361—375. 

Grayson, D. K. 1993. The desert’s past, a natural pre- 
history of the Great Basin. Smithsonian Insti- 
tution Press, Washington, 356 pp. 

Holsinger, J. R. 1981. Amphipoda. Pp. 36—40 in S. H. 
Hurlbert, G. Rodriguez & N. D. Santos, eds., 
Aquatic Biota of Tropical South America, Part 
1: Arthropoda. San Diego State University, San 
Diego, California. 

Jackson, H. H. T. 1912. A contribution to the natural 
history of the amphipod, Hyalella nickerbockeri 
(Bate).—Bulletin of the Wisconsin Natural His- 
tory Society 10(1—2):49-—60. 

Miller, G. A. 1977. Appraisal of the water resources 
of Death Valley California-Nevada. U.S. Geo- 


457 


logical Survey, Open File Report No. 77-728, 
68 pp. 

Pereira, V. EF G. C. 1989. Uma nova espécie de anfi- 
pode cavernicola do Brasil-Hyalella caeca sp. 
n. (Amphipoda, Hyalellidae)—Revista Brasi- 
leira de Zoologia 6(1):49—55. 

Pistrang, M. A., & E Kunkel. 1958. A brief geological 
and hydrological reconnaissance of the Furnace 
Creek wash area, Death Valley National Mon- 
ument, California. United States Department of 
Interior Groplogical Survey, Ground Water 
Branch, 73 pp. 

Ruffo, S. 1957. Una nuova specie troglobia di Hyalella 
del Venezuela.—Annali del Museo Civico Di 
Storia Naturale Genova 69:363—369. 

Saussure, H. 1858. Mémoire sur divers crustacés nou- 
veaux des Antilles et du Mexique.—Memoires 
de la Societe Physique Histoire Naturelles Ge- 
néve 14:417—496. 

Shepard, W. D. 1992. Riffle beetles (Coleoptera: El- 
midae) of Death Valley National Monument, 
California.—Great Basin Naturalist 52(4):378— 
381. 

. 1993. Desert springs—both rare and endan- 
gered.—Aquatic Conservation: Marine and 
Freshwater Ecosystems 3:351—359. 

Smith, S. I. 1874. Report on the amphipod crusta- 
ceans.—Annual Report of the United States 
Geological Survey of the Territories Embracing 
Colorado (1873):608—61 1. 

Stebbing, T. R. R. 1903. Amphipoda from Costa 
Rica.—Proceedings of the United States Na- 
tional Museum 26(1341):925—931. 

Stevenson, M. M., & A. E. Peden. 1973. Description 
and ecology of Hyalella texana n. sp. (Crusta- 
cea: Amphipoda) from the Edwards Plateau of 
Texas.—The American Midland Naturalist 
89(2):426—436. 

Thomas, P. E., W. Blinn, & P. Keim. 1994. A test of 
an allopatric speciation model for congeneric 
amphipods in an isolated ecosystem.—Journal 
of the North American Benthological Society 
13(1): 100-109. 

1997. Genetic and behavioral divergence 
among desert spring amphipod populations.— 
Freshwater Biology 38:137—143. 

Weckel, A. L. 1907. The fresh-water Amphipoda of 
North America.—Proceedings of the United 
States National Museum 32(1507):25—58. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


113(2):458—464. 2000. 


Caecidotea cumberlandensis, a new species of troglobitic isopod from 
Virginia, with new records of other subterranean Caecidotea 
(Crustacea: Isopoda: Asellidae) 


Julian J. Lewis 


217 W. Carter Avenue, Clarksville, Indiana 47129, U.S.A. 


Abstract.—Caecidotea cumberlandensis is a subterranean asellid isopod de- 
scribed from two caves in Cumberland Gap National Historic Park in south- 
western Virginia. New records of several other subterranean Caecidotea are 
also discussed: C. incurva, C. jordani, C. barri, C. paurotrigonus and C. ter- 
esae. Comparison of the descriptions of C. paurotrigonus and C. dauphina 
suggest that these species are conspecific. 


In 1976 Dr. John Holsinger told me of a 
subterranean isopod collected from Cliff 
Cave in the Cumberland Gap area of south- 
western Virginia that had been identified by 
Fleming (1972a) as Asellus scrupulosus (a 
troglophilic species with eyes and pigmen- 
tation). Dr. Holsinger had related that this 
identification might be in error since the 
Cliff Cave specimen was eyeless and unpig- 
mented, and suggested that I look at the 
specimen if the opportunity presented. Flem- 
ing’s identification was subsequently reject- 
ed entirely by Holsinger & Culver (1988). 

In 1997 I visited the Smithsonian Insti- 
tution to assist in the sad task of curating 
collections that remained in the office of my 
mentor and friend, Dr. Thomas E. Bowman, 
at the time of his death. During this process 
I came upon the Cliff Cave specimen as 
well as another vial of isopods from Indian 
Cave, Lee Co., Virginia. Examination of 
these specimens proved Dr. Holsinger’s sus- 
picion correct, that the Cliff Cave speci- 
mens represented a distinct taxon new to 
science. 


Family Asellidae G. O. Sars, 1897 
Caecidotea Packard, 1871 
Caecidotea cumberlandensis, new species 
Figs. 1-3 
Asellus scrupulosus.—Fleming, 1972a: 

241. 


Caecidotea species A.—Holsinger & Cul- 
ver, 1988: 30-31, 37. 


Material examined.—Virginia: Lee Co., 
Indian Cave, David A. Hubbard, Jr., 16 Mar 
1993, 5.5 mm male holotype (USNM 
291204), 4.5 mm male paratype, 6 female 
paratypes (USNM 291205); Cliff Cave, 
Russell M. Norton, 24 Nov 1966, 5.8 mm 
male paratype (USNM 291206), including 
a glass slide labelled “‘8-K A. scrupulosus” 
and signed ““LEF”’ (L. E. Fleming) contain- 
ing the first and second pleopods. 

All specimens remain in the National 
Museum of Natural History, Smithsonian 
Institution. 

Description.—Eyeless, unpigmented, 
longest male 5.8 mm, female 4.7 mm; body 
slender, about 5.2 as long as wide. Head 
about 1.5 as wide as long, anterior margin 
concave, postmandibular lobes moderately 
produced. Pleotelson about 1.4 as long as 
wide, sides subparallel, caudomedial lobe 
moderately produced. 

Mandibles with 4-cuspate incisors and la- 
cinia mobilis, palp with rows of plumose 
setae on distal segments. Maxilla 1 with 5 
robust plumose setae on inner lobe, 13 
spines on outer lobe. Antenna | reaching to 
about mid-point of last segment of antenna 
2 peduncle, flagellum of 6-7 segments, es- 
thetes present on last 4 segments. Antenna 


VOLUME 113, NUMBER 2 


2, last segment of peduncle about 1.5 
length of preceding segment, flagellum of 
holotype with 40 segments. 

Male pereopod 1, propus 2.6 as long as 
wide, palmar margin with 2 large spines, 
processes absent; female pereopod 1.2% as 
long as wide, palmar margin similar to 
male. Pereopods 2-7 similar, with moderate 
setation, sexual dimorphism of pereopod 4 
for grasping negligible, carpus about 2.2 
as long as wide. 

Male pleopod 1, protopod about 0.5 
length of exopod, with 2-3 retinacula; ex- 
opod about 1.8 as long as wide; lateral 
margin slightly concave, distolateral margin 
setae not plumose. Pleopod 2 exopod prox- 
imal segment with 2 plumose setae, distal 
segment with 3—5 plumose setae along mar- 
gin. Endopod with rounded basal apophy- 
Sis; tip with 2 distinct processes extending 
subparallel to one another and approxi- 
mately perpendicular to the axis of the en- 
dopod: (1) lateral process subterminal, slen- 
der, tapering slightly, and (2) cannula beak- 
shaped. Pleopods 3—5, endopods present, 
but unremarkable. Pleopod 3 exopod distal 
margin with 3—4 short, non-plumose setae. 
Pleopod 4 exopod, proximal setae absent; 
sutures indistinct, suggestive of 2 barely 
discernible, unconnected false sutures. Ple- 
opod 5 exopod with faint transverse sutures. 
Uropods of male about 0.5 X length of pleo- 
telson, female similar. 

Etymology.—The name refers to the 
Cumberland Gap area in which the species 
occurs. The suggested vernacular name is 
the Cumberland Gap cave isopod. 

Range.—This species is known only 
from Cliff and Indian caves in the Cum- 
berland Gap National Historic Park, Lee 
Co., Virginia. Descriptions of the caves 
were presented by Holsinger (1975). The 
caves are about 500 meters apart, occur in 
the same rock formation (Greenbrier Lime- 
stone), and are probably disconnected parts 
of the large Cudjos-Cumberland Saltpeter 
Cave system (Holsinger, in litt.). The am- 
phipod Stygobromus cumberlandus occurs 
with C. cumberlandensis in Cliff Cave 


459 


(Holsinger 1978). The range of this amphi- 
pod also includes Scott and Wise counties 
in Virginia, suggesting the possibility of a 
wider range for C. cumberlandensis. 

Relationships.—Caecidotea cumberlan- 
densis most closely resembles two other sub- 
terranean species, C. bicrenata (northern Al- 
abama to southern Illinois; Lewis 1982a) 
and C. richardsonae (Tennessee and Virgin- 
ia; Steeves 1963). These three species have 
in common a male pleopod 2 endopod tip 
with a terminal beak-shaped cannula and 
subterminal lateral process, both extending 
approximately perpendicular to the axis of 
the endopod. These species are most easily 
separated from one another by the shape of 
the lateral process, which in C. cumberlan- 
densis is thin and tapered distally, in C. bi- 
crenata thicker and cylindrical, and in C. ri- 
chardsonae finger-shaped and overlapped by 
the cannula (the tip processes shown by 
Lewis & Bowman 1977 for C. richardsonae 
had been spread by pressure from a cover- 
slip; the appearance depicted by Steeves 
1963 is more typical). Differences and sim- 
ilarities of key structures in these three spe- 
cies are summarized in Table 1. 


Caecidotea jordani (Eberly, 1966) 


Material examined.—Indiana: Crawford 
Co., seep spring flowing from bank of Blue 
River at former site of Rothrock Mill, Wy- 
andotte, Julian J. Lewis, Victor M. Lewis, 
James J; Lewis, 30° Jul’ 1998, 5 males; 12 
females; same locality, Julian J. Lewis, Sal- 
isa T. Rafail, 1 Aug 1998, 4 males, 5 fe- 
males. 

Range.—The only previously known 
population (a spring under Jordan Hall on 
the campus of Indiana University, Bloo- 
mington) was extirpated by termiticides. 
The site reported above is a parafluvial hab- 
itat where water seeps from a gravel bed 
that extends under the floodplain of the 
Blue River. 


Caecidotea incurva (Steeves & Holsinger, 


1968) 


Material examined.—Tennessee: Blount 
Co., Whiteoak Blowhole Cave, W. Reeves, 


460 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 1. Caecidotea cumberlandensis, 4.5 mm paratype male, Indian Cave, Lee Co., Virginia (a—b, d—h, j); 
5.5 mm holotype male, same locality (c); 4.7 mm paratype female, same locality, (i): (a) habitus, (b) antenna 
1, (c) antenna 2, (d) maxilla 1, outer lobe, (e) maxilla 1, inner lobe, (f) mandibular palp, (g) left mandible, 
incisor and lacinia mobilis, (h) right mandible, incisor, (i) pleopod 2, (j) uropod. 


VOLUME 113, NUMBER 2 


461 


Fig. 2. 
mm male holotype, same locality, (b—c): (a) pereopod 1, (b) same, (c) pereopod 4, (d) same. 


12 Aug 1999, 2 males, 2 females; Rich 
Mountain Blowhole Cave, W. Reeves, 25 
Aug 1999, 2 males, 1 female; Virginia: 
Wythe Co., Early’s Cave, D. A. Hubbard, 
Jr., 7 Jul 1997, 6 males, 4 females; Camp- 
bell Cave, D. A. Hubbard, Jr., 14 Sep 1998, 
1 male, 1 female; Lone Ash Cave #2, D. A. 
Hubbard, Jr, 3 Nov 1997, 4 males, 13 fe- 
males; Mockley Cave, D. A. Hubbard, Jr., 
29 Mar 1999, 1 male, 1 female. 
Range.—This species was incompletely 
described, but the endopod of the male sec- 
ond pleopod is so distinct in appearance 
(Steeves & Holsinger 1968) that a fairly 
certain identification can be made. Holsin- 
ger & Culver (1988) reported this species 
in Virginia from McMullin Cave, Smyth 
Co., and Groseclose Cave Number l, 
Wythe Co. Other unpublished Virginia re- 
cords identified and provided by J. R. Hol- 
singer (in litt.) are Deep Spring and Bowles 


Caecidotea cumberlandensis, 4.7 mm female paratype, Indian Cave, Lee Co., Virginia, (a, d); 5.5 


Spring caves, Wythe Co., and Dolingers 
Cave, Washington Co. 


Caecidotea paurotrigonus (Fleming, 
1972b) 


Material examined.—Louisiana: St. 
Mary Parish, holes dug in moist area at for- 
est edge beside road, just outside entrance 
to parish park at Burn’s Point, at end of 
state highway 317, coll. D.W.D., 23 Aug 
1981, 1 male, 1 fragment, 9 females. 

Range.—This groundwater species was 
previously known from a single male de- 
scribed by Fleming (1972b) from a ditch in 
southwestern Mississippi. The new locality 
is the first report of a subterranean asellid 
from Louisiana. Identification of C. pau- 
rotrigonus is obscured by Fleming’s de- 
scription. Based on a single 16.7 mm male, 
the description provided drawings of key 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 3: 


Caecidotea cumberlandensis, 4.5 mm paratype male, Indian Cave, Lee Co., Virginia (a—c, e—g); 5.8 


mm male, Cliff Cave, Lee Co., Virginia (d): (a) pleopod 1, (b) pleopod 2, (c) same, endopod tip, (d) same, (e) 
pleopod 3 exopod, (f) pleopod 4 exopod, (g) pleopod 5 exopod. 


structures, but the quality of the drawings 
and the interpretation of the structures were 
both questionable. Modlin (1986) on the 
other hand provided a detailed description 
of Caecidotea dauphina based on a 7.5 mm 
male. Unfortunately, three of the four struc- 


tures figured by Fleming for C. paurotri- 
gonus bear a strong resemblance to that of 
C. dauphina: (1) pereopod 1 propods of 
very similar dimensions, margin in C. dau- 
phina with basal spine, medial and distal 
processes, identical in C. paurotrigonus ex- 


VOLUME 113, NUMBER 2 


463 


Table 1.—Comparison of selected structures of male C. cumberlandensis, C. bicrenata and C. richardsonae 


useful for separating the species. 


C. cumberlandensis 


C. bicrenata C. richardsonae 


Pereopod 1 propod 


palmar margin processes absent 
Pleopod | 

distolateral lobe absent 
Pleopod 2 exopod 3-5 setae 


Pleopod 2 endopod 


lateral process 
Proportion of uropod 


length to pleotelson 0.5 length 


cept the spine is replaced by a process (as 
typical of more mature specimens); (2) ple- 
opod 1 with essentially identical shapes, se- 
tation patterns and quantity of retinacula; 
(2) pleopod 2 endopod tips with short api- 
cal cannula directed mediad, and short, un- 
remarkable knob-like lateral process. I sus- 
pect that C. dauphina represents a well de- 
scribed juvenile of the poorly described, but 
conspecific C. paurotrigonus. Unravelling 
this will require redescription of C. pauro- 
trigonus and perhaps more male specimens 
for comparison from Mississippi and Ala- 
bama. 


Caecidotea barri (Steeves, 1965) 


Material examined.—Kentucky: Wood- 
ford Co., small spring 0.6 mile E. Clifton, 
in side valley above waterfall, elevation 
about 670 feet, Julian J. Lewis, Victor M. 
Lewis, James J. Lewis, 19 Feb 1995, 3 
males, 4 females. 

Range.—This species was previously 
known only from the type-locality, Clifton 
Cave, Woodford Co., Kentucky (Steeves 
1965), which was bulldozed shut by high- 
way workers. The new locality is across the 
valley from Clifton Cave. 


Caecidotea teresae Lewis, 1982b 


Material examined.—Indiana: Floyd Co., 
well on Grant Line Road, Julian J. Lewis, 
26 Apr 1995, 1 male, 1 female. 


thin, tapered distally 


present absent 
absent present 
12-15 setae 12—15 setae 


cylindrical, not tapered long digitiform 


1.5—2.0 length 1.5—2.0 length 


Range.This species was apparently extir- 
pated by termiticides at the two previously 
known localities on the campus of Indiana 
University Southeast, New Albany. The 
well from which the new specimens were 
taken is found in a pasture across Grant 
Line Road from the I.U.S. campus. Accord- 
ing to the owner, the well was hand dug at 
the end of the 19th century and was 21 feet 
deep, with 8 feet in soil and the bottom 13 
feet through New Albany Shale. 


Acknowledgments 


The description of Caecidotea cumber- 
landensis was prepared during a visit to the 
National Museum of Natural History fund- 
ed by a Smithsonian visiting scientist grant. 
Caecidotea jordani was rediscovered dur- 
ing the bioinventory of Blue River area 
caves funded by a Rodney Johnson/Kathar- 
ine Ordway Stewardship Endowment Grant 
from The Nature Conservancy. Caecidotea 
teresae was rediscovered during field work 
funded by the Indiana Non-game and En- 
dangered Wildlife Program. Caecidotea 
barri was found during field work funded 
by the U.S. Fish & Wildlife Service and 
Kentucky Nature Preserves Commission 
(Dr. Thomas C. Barr, Jr. principle investi- 
gator). I thank Dr. John R. Holsinger (Old 
Dominion University), Dr. Brian Kensley 
(Smithsonian Institution), Dr. Guy Magniez 
(University of Dijon), Mr. David Hubbard, 


464 


Jr. (Virginia Division of Mineral Resourc- 
es), and Mr. Allen Pursell (The Nature Con- 
servancy) for reading the manuscript and 
making suggestions on its improvement. 


Literature Cited 


Eberly, W. R. 1966. A new troglobitic isopod (Asel- 
lidae) from southern Indiana.—Proceedings of 
the Indiana Academy of Science 75:286—288. 

Fleming, L. E. 1972a. The evolution of the eastern 
North American isopods of the genus Asellus 
(Crustacea: Asellidae).—International Journal 
of Speleology 4:221—256. 

. 1972b. Four new species of troglobitic asel- 
lids (Crustacea: Isopoda) from the United 
States.—Proceedings of the Biological Society 
of Washington 84:489—499. 

Holsinger, J. R. 1975. Descriptions of Virginia 
Caves.—Virginia Division of Mineral Resourc- 
es, Bulletin 85, 450 pp. 

. 1978. Systematics of the subterranean amphi- 

pod genus Stygobromus (Crangonyctidae), Part 

II: Species of the eastern United States.— 

Smithsonian Contributions to Zoology 266:1— 

144. 

, & D. C. Culver. 1988. The invertebrate cave 

fauna of Virginia and a part of eastern Tennes- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


see: Zoogeography and ecology.—Brimleyana 
14:1-162. 

Lewis, J. J. 1982a. Systematics of the troglobitic Cae- 
cidotea (Crustacea: Isopoda: Asellidae) of the 
southern Interior Low Plateaus.—Brimleyana 8: 
65-74. 

. 1982b. A diagnosis of the Hobbsi Group, with 

descriptions of Caecidotea teresae, new spe- 

cies, and C. macropropoda Chase and Blair 

(Crustacea: Isopoda: Asellidae).—Proceedings 

of the Biological Society of Washington 95: 

338-346. 

, & T. E. Bowman. 1977. Caecidotea caroli- 
nensis, n. sp., the first subterranean water slater 
from North Carolina (Crustacea: Isopoda: Asel- 
lidae).—Proceedings of the Biological Society 
of Washington 90:968—974. 

Modlin, R. E 1986. Caecidotea dauphina, a new sub- 
terranean isopod from a barrier island in the 
northern Gulf of Mexico (Crustacea: Isopoda: 
Asellidae)—Proceedings of the Biological So- 
ciety of Washington 99:316—322. 

Steeves, H. R., III. 1963. The troglobitic asellids of the 
United States: The Stygius Group.—American 
Midland Naturalist 69:470—48 1. 

. 1965. Two new species of troglobitic asellids 

from the United States——American Midland 

Naturalist 73:81—84. 

, & J. R. Holsinger. 1968. Biology of three new 

species of troglobitic asellids from Tennes- 

see.—American Midland Naturalist 80:75-—83. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


113(2):465—479. 2000. 


Euphilomedes cooki, a new species of myodocopid ostracode from 
Moreton Bay, SE Queensland, Australia 


Elizabeth Harrison-Nelson and Louis S. Kornicker 


Department of Invertebrate Zoology, National Museum of Natural History, 
Smithsonian Institution, Washington, D.C. 20560-0163, U.S.A. 


Abstract.—A new species of myodocopid ostracode in the subfamily Phi- 
lomedinae from Moreton Bay, S.E. Queensland, Australia, is described and 
illustrated. A key is presented to the ten known species of Euphilomedes having 


primary furcal claws 1, 2, 4, and 6. 


This work describes and illustrates a new 
species of myodocopid ostracode from 
Moreton Bay, S.E. Queensland, Australia, 
which is north of Brisbane, near the south- 
ern end of the Great Barrier Reef. 

Disposition of specimens.—The holotype 
and paratypes have been deposited at the 
Queensland Museum (QM), South Bris- 
bane, Queensland. 


Superorder Myodocopa Sars, 1866 
Order Myodocopida Sars, 1866 
Suborder Myodocopina Sars, 1866 
Family Philomedidae Miiller, 1906 


This family contains two subfamilies, 
Philomedinae Miiller, 1906, and Pseudo- 
philomedinae Kornicker, 1967, both with 
representatives in the vicinity of S.E. 
Queensland, Australia (Kornicker 1994). 


Philomedinae Miiller, 1906 


This subfamily includes eight genera: 
Philomedes Liljeborg, 1853, Pleoschisma 
Brady, 1890, Scleroconcha Skogsberg, 
1920, Paraphilomedes Poulsen, 1962, Eu- 
philomedes Kornicker, 1967, Anarthron 
Kornicker, 1975, Igene Kornicker, 1975, 
and Zeugophilomedes Kornicker, 1983. A 
new species of Euphilomedes is described 
herein. 


Euphilomedes Kornicker, 1967 


Type species.—Euphilomedes nodosa 
Poulsen, 1962 (subsequent designation by 
Kornicker 1967). 

Composition.—Including the new spe- 
cies described herein this genus includes 19 
species plus one subspecies (Kornicker 
19912371 995e16). 

Distribution.—The genus is cosmopoli- 
tan except in Arctic and Antarctic waters. 
The known depth range is shallow water to 
2250 m, but members have been collected 
mostly from the continental shelf and upper 
slope (Kornicker & Harrison-Nelson 1997: 
14). Three species of the genus have been 
reported previously from the vicinity of 
Australia: E. corrugata (Brady 1897) from 
off Port Jackson and in Flinders Passage at 
depths of 3.7—18.3 m, E. walfordi Poulsen, 
1962, from the Coral Sea at a depth of 50 
m, and E. erynx Kornicker, 1995, from the 
continental slope off New South Wales at a 
depth of 220 m. The new species described 
herein is from Middle Banks, northern 
Moreton Bay, S.E. Queensland, Australia, 
at a depth of 15—25 m. 


Euphilomedes cooki, new species 
Figs. 1-9 


Etymology.—The species is named in 
honor of Stephen Cook formerly of the 
Queensland Museum, Australia, who col- 
lected some of the specimens. 


466 


Holotype.—Queensland Museum W2492, 
undissected adult female in alcohol. 

Type locality.—Miiddle Banks, northern 
Moreton Bay (27.02°S, 153.25°E), Queens- 
land, Australia, about 20 km offshore from 
the mainland, depth 15-25 m. All speci- 
mens part of Queensland Museum Registra- 
tion Number 11879. 

Paratypes.—12 adult females; 3 A-1 
males; 1 A-1 female. 

Distribution.—Middle Banks, northern 
Moreton Bay, S.E. Queensland, depth 15-— 
25 m, clean sand or sand and shell. Col- 
lected with Smith-MclIntyre grab in Novem- 
ber 1983 and November 1984. 

Description of adult female (Figs. 1-7a— 
7c).—Carapace elongate with shallow in- 
cisur (Fig. la). Left valve extends past right 
valve along free margin. Posterodorsal 
hinge area straight and without valve over- 
lap. 

Ornamentation: Carapace with pits of 
various sizes and shapes. Each pit with nar- 
row raised border. Some pits two or three 
times larger than others; pits larger in pos- 
terior half of valve. Valve edge with long 
single bristles. Long single bristles sparsely 
distributed on valve surface. 

Infold: Rostral and anteroventral infold 
(Fig. 1b,c,e, 7a), infold of caudal process 
and posteroventral infold (Fig. 1d,f, 7b), 
each with row of bristles. Anteroventral in- 
fold with several parallel ridges (Fig. 1c). 

Selvage: Broad lamellar prolongation of 
right valve in vicinity of rostrum with long 
marginal hairs; lamellar prolongation nar- 
rower and with shorter marginal hairs along 
ventral margin. Lamellar prolongation of 
left valve obscured. 

Central adductor muscle attachments 
(Fig. 1g): About 30 attachments on each 
side. These represented by large pits on 
each valve just anterior to midlength; most 
pits ventral to midheight. 

Carapace Size (mm): Average length 
1.89, range 1.81—1.95; average height 1.36, 
range 1.29-1.42; average height 72% of 
length, range 71-74; n = 11. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Height 
Length Height % length 
Holotype 1.94 39 TW 
Paratypes 1.84 i Wee) 74 
25 1.40 93 
LoS 1.42 73 
87 Sy, 71 
1.81 Vol 71 
Peo 136 A 
oS 1.41 2 
1.86 i335 73 
87 1has) 7 71 
1.81 29 7 


First antenna (Fig. 2a): 1st joint with dis- 
tal medial spines forming rows. 2nd joint 
with dorsal, ventral, and lateral spines, and 
3 bristles (1 dorsal, 1 ventral, 1 medial). 3rd 
joint short with 3 bristles (2 dorsal, 1 ven- 
tral). 4th joint with 6 bristles (2 dorsal, 4 
ventral). Sth joint about same length as 4th 
joint, with long sensory bristle with about 
7 short marginal filaments. Long medial 
bristle of 6th joint with base near dorsal 
margin and with long proximal and short 
distal spines. 7th joint: a-bristle similar to 
bristle of 6th joint; b-bristle with 2 fila- 
ments near midlength, 2 subterminal fila- 
ments, and bifurcate tip; c-bristle long with 
marginal filaments. 8th joint: d- and e-bris- 
tles long, bare, and with blunt tips; f- and 
g-bristles long with marginal filaments. Not 
all filaments of bristles of 7th and 8th joints 
shown in illustration. 

Second antenna: Protopodite bare. En- 
dopodite 2- or 3-jointed (Fig. 2b): 1st joint 
with row of 5 short bare proximal bristles 
and 1 very long spinous distal bristle; 2nd 
joint with very long spinous proximal bris- 
tle (bristle 2 % times length of long bristle 
of Ist joint) and short bare terminal bristle 
(the terminal bristle could be interpreted to 
be on a 3rd joint). Exopodite: Ist joint with 
minute straight medial terminal bristle; bris- 
tle of 2nd joint reaching well past 9th joint, 
with few dorsal spines and abundant short, 
fairly stout, ventral spines, no natatory 
hairs; bristles of 3rd and 4th joints with 
short slender distal spines, no natatory 


VOLUME 113, NUMBER 2 467 


a 


Fig. 1. Euphilomedes cooki specimen 12, adult female paratype: a, outline of complete specimen from left 
side, length 1.84 mm, note four epibionts attached to edge of valve. b—d, interior views of left valve: b, anterior 
end; c, anteroventral margin; d, posterior end. e, f, interior views of right valve: e, anteroventral margin; f, 
posteroventral margin. g, ends of central adductor muscles projecting from right side of body, anterior toward 
right. Abbreviation are: ant: antenna; Bo: Bellonci organ; cx: coxale; e: edge of valve; end: endopodite; ex: 
exopodite; epip: epipodite; fu: furca; gen: genitalia; gird: girdle; im: inner margin of infold; li:list; lv: lateral 
view; me: medial eye; mnd: mandible; mv: medial view; mx: maxilla; nabs: not all bristles shown; prot: pro- 
topodite; ul: upper lip; Y-scl: Y-sclerite. Roman numerals designate endites. Arrow on illustration indicates 
anterior. 


468 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


APACE 

ee = NAN ON 
oN 

7 
Vi 


\ 
/ 


\ prot 
\ 
7 
- Y 
Lh \ 
Z. 
\ Vi 
\ \ y 
SY ‘ 


Riga: 


Euphilomedes cooki specimen 12, adult female paratype: a, right Ist antenna, lv; b, left 2nd antenna, 
endopodite and distal part of protopodite, mv; c, right 2nd antenna, exopodial joints 8 and 9, only proximal 


parts of some bristles shown, lv; d, right furcal lamella, lv; e, left lateral eye, lv; f, posterior of body from right 
side; g, outline of unextruded egg, drawn at same scale as “‘f’’. (For abbreviations see Fig. 1.) 


VOLUME 113, NUMBER 2 


hairs; bristles of joints 5—8 longer than bris- 
tles of joints 2—4, with natatory hairs, no 
spines; 9th joint with 7 bristles (4 long and 
2 short with natatory hairs, 1 very short and 
with short marginal spines (Fig. 2c, only 
proximal parts of some bristles shown); 
joints 2—8 with row of terminal spines. 

Mandible (Fig. 3a, b): Coxale endite spi- 
nous, tip bifurcate, with small ringed bristle 
near base. Basale: medial surface and ven- 
tral and dorsal margins with rows of spines; 
medial surface with 5 bristles in proximal 
ventral corner (3 pectinate unringed, 2 
ringed and with long proximal and short 
distal spines), and 1 short ringed bristle 
closer to midlength with long proximal and 
short distal spines; dorsal margin with 3 
bristles (1 near midlength, 2 terminal); 7 
bristles with long proximal and short distal 
spines present on or near ventral margin (2 
longest bristles distal and with bases on 
ventral margin; 5 shorter bristles with bases 
slightly lateral). Exopodite slightly more 
than % length of dorsal margin of Ist en- 
dopodial joint, with distinct distal hirsute 
pad and few terminal spines, and 2 bristles 
(outer bristle with short marginal spines and 
about % length of inner bristle; inner bristle 
reaching midlength of 2nd endopodial joint, 
with long spines near midlength and short 
distal spines). 1st endopodial joint with me- 
dial spines and 4 ventral bristles. 2nd en- 
dopodial joint: dorsal margin with 2 long 
bristles in proximal group and 5 spinous 
bristles in distal group (2 long with basis 
on margin; 3 short with bases medial (prox- 
imal 2 with long spines, distal 1 with short 
spines)); ventral margin with bristles in 2 
groups (2 in proximal group, 3 in distal 
group); medial surface with spines forming 
rows. 3rd endopodial joint with 3 pectinate 
claws (dorsal claw short), and 3 ringed bris- 
tles. 

Maxilla (Figs. 3b, 4a—c): Precoxale and 
coxale with fringe of long hairs. Coxale 
with plumose dorsal bristle. Basale with 3 
distal bristles. Exopodite with 3 bristles 
(proximal short bristle bare, long middle 
bristle with long spines, other long bristle 


469 


with short spines). Ist endopodial joint with 
1 alpha-bristle and 4 beta-bristles. 2nd en- 
dopodial joint with 3 pectinate claw-like 
bristles and about 7 ringed bristles. Endites 
with stout spinous and pectinate bristles 
(endite I with 9 bristles, endite II with 6 
bristles, endite III with 8 bristles). 

Fifth limb (Figs. 4d—f, 5): Epipodite with 
46 bristles. Endite I with 5 bristles (only 3 
shown in illustration); endite II with 6 bris- 
tles; endite III with 8 bristles. Ist exopodial 
joint: anterior side with 2 bristles (with long 
spines) on distal edge; outer corner with 2 
small slender bristles with few marginal 
hairs; main tooth with proximal peg fol- 
lowed by 3 pointed teeth (teeth worn down 
on specimen 12) and 1 large squarish tooth; 
ringed spinous bristle proximal to teeth. 2nd 
exopodial joint: posterior side with 1 prox- 
imal bristle and group of 3 distal bristles 
(middle bristle long, others short (short 
bristle obscured on right limb of specimen 
12)). 3rd exopodial joint: inner lobe with 3 
bristles, outer lobe with 2 bristles with long 
spines. Fused 4th and 5th exopodial joints 
with 8 bristles. 

Sixth limb (Figs. 5a, 6a): Epipodite with 
3 spinous bristles. Endite I with 3 bristles; 
endite II with 4 bristles; endites III and IV 
each with 9 bristles. End joint with spines 
and hairs and 18 bristles (most with long 
proximal hairs and short distal spines). 

Seventh limb (Fig. 6b, c): Each limb with 
13-15 bristles: proximal group with 8—10 
bristles (specimen 12: 5 or 6 on peg side, 
4 on comb side; specimen 4 (right limb), 5 
on peg side, 3 on comb side), each bristle 
with 3—5 bells and marginal spines; termi- 
nal group with 5 bristles (3 on peg side, 2 
on comb side). Each bristle with 3—7 bells 
and marginal spines. Terminal comb with 
12 alate teeth; 2 small curved pegs (with 
proximal teeth) present opposite comb. 

Furca (Fig. 2d, f): Each lamella with 11 
claws: claws 1, 2, 4, and 6 primary; claws 
3, 5, 7-11 secondary. Primary claws with 
stout posterior teeth; teeth of claw 1 stouter 
than teeth of other primary claws and with 
stout medial teeth; secondary claws with 


470 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


a 


sw eSee 
I a 


—— OTE oTo—E—E=x=«: oo — 


Fig. 3. Euphilomedes cooki specimen 12, adult female paratype: a, right mandible, mv; b, left maxilla and 
mandible in place on body, nabs, mv; c, medial eye and Bellonci organ from left side. (For abbreviations see 
Fig. 1.) 


VOLUME 113, NUMBER 2 471 


Fig. 4. Euphilomedes cooki specimen 12, adult female paratype: a—c, right maxilla: a, complete limb, lv; b, 
bristle of distal end of Ist exopodial joint, mv; c, bristles and claws of 2nd endopodial joint, mv. d—f, left 5th 
limb, endites I, II, and III, respectively. (For abbreviations see Fig. 1.) 


472 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


An 


Fig. 5. Euphilomedes cooki specimen 12, adult female paratype: a, upper lip, left 6th limb, and right 5th 
limb in place on body, not all bristles shown. b, c, posterior views of distal end of right fifth limb, respectively. 
d, e, posterior and anterior views of distal end of left fifth limb, respectively. (For abbreviations see Fig. 1.) 


marginal spines. Right lamella with few Bellonci organ (Fig. 3c): Elongate with 
distal anterior spines and also spines medial short wide part (bearing striations) near 
and proximal to claw 1. Right lamella an- midlength and narrow tapered tip bearing 
terior to left by % width of claw 1. minute spines. 


VOLUME 113, NUMBER 2 473 


\ 
\ 
) 
kk 
Ly 
YE 
Gp 
\ 
} 
\ 


~ 


CUP IT/ 


p 
i 


Fig. 6. Euphilomedes cooki specimen 12, adult female paratype: a, right 6th limb, lv; b, right 7th limb; c, 
tip of left 7th limb, only proximal part of bristles shown. (For abbreviations see Fig. 1.) 


474 


Eyes: Medial eye with brown pigment 
(Fig. 3c). Lateral eye smaller than medial 
eye, unpigmented, with 4 divided amber- 
colored ommatidia (Fig. 2e). 

Upper lip (Figs. 5a, 7c):-Projecting 
slightly anteriorly, with anterior and lateral 
glandular processes. 

Genitalia (Fig. 2f): Small oval. 

Anterior of body: Convex. 

Posterior of body (Fig. 2f): Evenly 
rounded, bare. 

Y-sclerite (Fig. 2f): With ventral branch. 

Number and lengths of eggs: Specimen 
12 with about 16 unextruded eggs (Fig. 2g). 
Specimen 4 with 18 eggs in marsupium 
(lengths of 2 eggs 0.406 mm, 0.408 mm). 

Gut content: Specimen 12 with unrec- 
ognizable amber-colored particles in gut. 

Epizoa: Specimens 12 and 4 with vase- 
shaped protistans along posterior margin 
(Fig. la, f). 

Description of A-1 male (Fig. 7d—f).— 
Carapace similar in shape and ornamenta- 
tion to that of adult female. 

Carapace size (length, height in mm): 
1.59, 1.06, height 67% of length (specimen 
13); 1.58, 1.13, height 72% of length. 

Second antenna: Protopodite bare with 
narrow slightly curved proximal pivotal 
sclerite. Endopodite 3-jointed (Fig. 7d): Ist 
joint short with 4 small proximal ringed 
bristles and 1 distal long stout ringed bristle 
(broken off in illustrated endopodite (Fig. 
7d)); 2nd joint elongate with 3 small ringed 
bristles; 3rd joint elongate with 2 small 
ringed terminal bristles with short spines. 
Exopodite 9-jointed: Ist joint with small 
medial terminal spine; bristle of 2nd joint 
less than twice length of exopodite, with 
spines along ventral edge; bristles of joints 
3-8 fairly short, longer than bristle of 2nd 
joint, but less than twice length of exopod- 
ite, with ventral spines stouter than those of 
bristle of 2nd joint, without natatory hairs; 
9th joint with 6 bristles (2 short lateral at 
distal dorsal corner, 4 longer terminal (lon- 
gest at ventral end and less than twice 
length of exopodite)), all with marginal 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


spines, without natatory hairs; joints 2-8 
with row of spines along terminal edges. 

Fifth limb (Fig. 7e, f): Similar to that of 
adult female. 

Sixth limb: With 3 epipodial bristles. En- 
dite I with 3 bristles; endite II with 4 bris- 
tles; endite III with 8 bristles. 

Seventh limb: Limb with 12 tapered bris- 
tles with marginal spines: proximal group 
with 7 bristles (3 on comb side, 4 on peg 
side), each with 2 or 3 bells; terminal group 
with 5 bristle (2 on comb side, 3 on peg 
side), each with 2—5 bells. Terminus with 
comb of about 11 alate teeth opposite 2 
small pegs. 

Furca: Similar to that of adult female ex- 
cept with only 4 secondary claws following 
primary claw 6. 

Lateral eye: Well developed with many 
ommatidia and black pigment between 
them. 

Description of A-1 female (Figs. 8,9).— 
Carapace similar in shape and ornamenta- 
tion to that of adult female, but narrower. 

Carapace size (length, height in mm): 
1.58, 1.02; height 65% of length (specimen 
1): 

Remarks.—The absence of natatory hairs 
on bristles of the exopodite of the 2nd an- 
tennae of A-1 males and females indicates 
that the juveniles of this species are inca- 
pable of efficient swimming. 

Comparisons.—Each lamella of the furca 
of the new species E. cooki bears primary 
claws 1, 2, 4, and 6, secondary claws 3 and 
5, and 5 additional secondary claws follow- 
ing primary claw 6. Only one species of 
Euphilomedes having a similar distribution 
of primary and secondary claws has been 
described previously from the vicinity of 
Australia, E. walfordi, which is known from 
only the adult male, and so is not directly 
comparable in all morphological characters 
to the adult female and A-1 instar male of 
E. cooki described herein. However, the fol- 
lowing differences between the adult male 
of the E. walfordi and the adult female of 
E. cooki are considered significant: 1, The 
carapace length of the adult female cooki is 


VOLUME 113, NUMBER 2 475 


Fig. 7. Euphilomedes cooki specimen 2, adult female paratype: a, b, interior views of anterior and posterior 
ends of right valve, respectively; c, upper lip, anterior toward left. d, specimen 13, A-1 male paratype, left 2nd 
antenna, endopodite and distal part of protopodite, mv. e, f, right 5th limb, av: e, distal end endite I. (For 
abbreviations see Fig. 1.) 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


400M 20KY 200HM 20K¥ 


200HM OO = 190M ZOKY 


Fig. 8. Euphilomedes cooki, A-1 female paratype (specimen 1), left valve: a, lateral view of valve, length 
1.58 mm; b, dorsal view of valve, anterior to left; c, anterior view of valve, ventral to right; d, anteroventral 


margin of valve, lv, from ‘‘a’’; e, area in vicinity of central adductor muscle, from ‘‘a’’; f, detail of reticulations, 
from lower left part of ‘‘e’’. 


VOLUME 113, NUMBER 2 477 


——20FM 20K¥~—s«o 


7 * 
PE 
ek 


00 


40PM 2OKY¥ 


10rM 


Fig. 9. Euphilomedes cooki A-1 female paratype (specimen 1), left valve: a, detail of pits and bristles in 
upper right part of Fig. 8a; b, detail of bristle shown in upper right of ‘‘a’’; c, detail of bristle and reticulations 
in posterior part at midheight of Fig. 8a; d, detail of bristle and reticulations in anterior part at midheight of 
Fig. 8a; e, detail of bristles and reticulations near incisure in Fig. 8d. 


478 


1.73—1.95 mm compared to 3 mm for the 
adult male walfordi; 2, the outer surface of 
the carapace of the adult female cooki has 
few hairs, whereas that of the adult male 
walfordi, according to Poulsen (1962:371), 
‘‘is densely covered with short hairs’’. The 
seventh limb of the adult male walfordi has 
19 bristles without marginal spines (Poul- 
sen 1962:372), whereas that limb of the 
adult female cooki has 13-15 bristles with 
marginal spines (when differences occur in 
the number of bristles on the 7th limbs of 
male and female myodocopids, the male 
generally has fewer bristles than the fe- 
male). 

The key to species of Euphilomedes pre- 
sented below is restricted to those species 
having furcal lamellae with primary claws 
1, 2, 4, and 6, and secondary claws 3, 5, 
and 7+. 


Key to species of Euphilomedes having 
furcal claws 1, 2, 4, and 6 primary* 


1. Lower angle of posterior shell margin 
withosmialletoothr nce ca%\ tears sinc ie oe 

E. sinister (Kornicker, 1974) 

1. Lower angle of posterior shell margin 


a Xe: <0) 1e) .0), Xe) ware. 


VALCDOUE COOUME ona ex chins Ste Gk ae 2 
2. Seventh limb with less than 12 cleaning 
DEISCIES Se ean ee ner oe one ae cere 3 


2. Seventh limb with 13-15 cleaning bris- 
tles E, cooki, 1. sp. 
2. Seventh limb with more than 15 cleaning 
bristles E. walfordi (Poulsen, 1962) 
3. 2nd joint of endopodite of female 2nd 
antenna with only the long plumose bris- 
(1G 3 secret one ee E. africana (Klie, 1940) 
3. This joint with an additional distal bristle 


Late CSS OUND OMe iO! On Lh Oe OE et 


oe © ee 


4. The distal bristle is only a spine, shorter 
than: the width of- the jomt 5733, @. 2. 
E. japonica (Miller, 1890) 
4. This bristle slightly longer than width of 
(0.1 11 Cees emer eer 17 SSR Atop tum racer ae See oe) 
E. morini (Kornicker & Harrison-Nelson, 
1997) 
4. This bristle longer than twice the width 
ol the 4oinbos: ec eae es ote 5 
5. This bristle is placed dorsally on the 
joint E. nodosa (Poulsen, 1962) 


© #0) @ 8) @) 0 a 10 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


5. This bristle is placed on the narrow tip 
Of thé joint. o> 62 eee 6 
6. Furca with more than 12 pairs of claws 
E. smithi (Poulsen, 1962) 
6. Furca with 11 or less pairs of claws ... 7 
7. Shell with a dorso-posterior, chitinized 
PROCESS: ar E. producta (Poulsen, 1962) 
7. Shell without such process 


E. longiseta (Juday, 1907) 


©) 6% 'e, (0° <6) ‘0; 6) (0: 0) she) <0 Ke, 


* Key adapted in part from the key to spe- 
cies of the genus Euphilomedes in Poul- 
sen (1962:362). 


Acknowledgments 


Collections of specimens from Moreton 
Bay described herein were made by P. 
Saenger and S. Cook, Queensland Museum, 
Australia. We thank Matthew Kane, Smith- 
sonian Institution (SI) volunteer, for inking 
penciled camera lucida drawings of append- 
ages, Walter Brown (SI) for SEM micro- 
graphs of the carapace, and Andrew R. 
Parker, University of Oxford, for reviewing 
the manuscript. 


Literature Cited 


Brady, G. S. 1890. On Ostracoda collected by H. B. 
Brady, Esq., L.L.D.ER.S., in the South Sea Is- 
lands.—Transactions of the Royal Society of 
Edinburgh, 35(part 2):489—525. 

. 1897. A supplementary report on the crusta- 
ceans of the group Myodocopa obtained during 
the Challenger Expedition, with notes on other 
new or imperfectly known species.—Transac- 
tions of the Zoological Society of London 14: 
85-100. 

Juday, C. 1907. Ostracoda of the San Diego region, II. 
Littoral forms.—University of California Pub- 
lications in Zoology 3:135—156. 

Klie, W. 1940. Beitrage zur Fauna Eulitorials von 
Deutsch Stidwest-Afrika, II: Ostracoden von der 
Kiiste Deutsch-Stidwest-Afrika.—Kieler Meer- 
esforschungen 3:403—448. 

Kornicker, L. S. 1967. The myodocopid Ostracod fam- 
ilies Philomedidae and Pseudophilomedidae 
(New Family).—Proceedings of the United 
States National Museum 120(3580):1-—35. 

. 1974. Revision of the Cypridinacea of the 

Gulf of Naples (Ostracoda).—Smithsonian 

Contributions to Zoology 178:1—64. 

1975. Antarctic Ostracoda (Myodocopina) 


VOLUME 113, NUMBER 2 


Parts 1 and 2.—Smithsonian Contributions to 

Zoology 163:1—720. 

. 1983. Zeugophilomedes, a new genus of my- 

odocopine Ostracode.—Proceedings of the Bi- 

ological Society of Washington 96:478—480. 

. 1991. Myodocopid Ostracoda of hydrothermal 

vents in the Eastern Pacific Ocean.—Smithson- 

ian Contributions to Zoology 516:1—46. 

. 1994. Ostracoda (Myodocopina) of the SE 

Australian continental slope, Part 1.—Smith- 

sonian Contributions to Zoology 553:1—200. 

. 1995. Ostracoda (Myodocopina) of the SE 

Australian Continental Slope. Part 2.—Smith- 

sonian Contributions to Zoology 562:1—97. 

, & E. Harrison-Nelson. 1997. Myodocopid Os- 
tracoda of Pillar Point Harbor, Half Moon Bay, 
California.—Smithsonian Contributions to Zo- 
ology 593:1-53. 

Liljeborg, W. 1853. Ostracoda. Pp. 92—130 in De Crus- 


479 


taceis ex ordinibus tribus: Cladocera, Ostracoda 
et Copepoda in Scania Occurrentibus. Lund: 
Berlingska Boktryckeriet. 

Miiller, G. W. 1890. Neue Cypridiniden.—Zoologische 
Jahrbuecher 5:211-—252. 

1906. Die Ostracoden der Siboga-Expedi- 
tion.—Uitkomsten op Zod6logisch, Botanisch, 
Oceanographisch en Geologisch Gebied verza- 
meld in Nederlandsch Oost-Indié, 1899-1900 
30:1—40. Leiden: E. J. Brill. 

Poulsen, E. M. 1962. Ostracoda-Myodocopa, 1: Cy- 
pridiniformes-Cypridinidae.—Dana Report 57: 
1—414. Copenhagen: Carlsberg Foundation. 

Sars, G. O. 1866. Oversigt af Norges marine Ostra- 
coder.—Forhandlinger i Videnskabs-Selskabet I 
Christiania 8:1—130. [Preprint, 1865.] 

Skogsberg, T. 1920. Studies on marine ostracods, I: 
Cypridinids, Halocyprids, and Polycopids.— 
Zoologiska Bidrag fran Uppsala (supplement 
1): 1-784. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


113(2):480—485. 2000. 


A new record of Cornechiniscus madagascariensis Maucci, 1993 
(Tardigrada: Echiniscidae) from India 


Wataru Abe and Masatsune Takeda 


(WA, MT) Department of Biological Sciences, Graduate School of Science, 
The University of Tokyo; 
(MT) Department of Zoology, National Science Museum, 3-23-1 Hyakunincho, Shinjuku-ku, 
Tokyo, 169-0073 Japan 


Abstract.—Cornechiniscus madagascariensis Maucci, 1993 which has been 
known only from its type locality, Madagascar, is reported from Himachal 
Pradesh, northern India. Some taxonomic emendations to the original descrip- 


tion are made from the paratypes. 


Recently, through the courtesy of Dr. H. 
Dastych of the Zoologisches Institut und 
Zoologisches Museum der Universitat 
Hamburg, we had an opportunity to ex- 
amine an unidentified Cornechiniscus spe- 
cies collected from Himachal Pradesh, 
northern India. After the direct comparison 
with the paratypes of C. madagascariensis 
Maucci, 1993 deposited in the Museo Civ- 
ico di Storia Naturale di Verona, it was 
concluded that the specimen from India 
should be identified with C. madagascar- 
iensis Which has previously been known 
only from its type locality, Madagascar. In 
this paper, we describe this species in de- 
tail based on the specimen from India, and 
make some taxonomic emendations from 
the paratypes. 

The specimens mounted on the micro- 
slides were closely examined by a phase 
and Nomarski differential interference con- 
trast microscope (Zeiss Axiophot) at maxi- 
mum magnification (X2500), illustrated 
with the aid of camera lucida, and measured 
using an eyepiece micrometer. 

Terminology is mainly that used in Abe 
et al. (1998). Abbreviations used in the text 
are as follows, CT: Museo Civico di Storia 
Naturale di Verona (Italy), ZMH: Zoolo- 
gisches Institut und Zoologisches Museum 
der Universitat Hamburg (Germany). 


Genus Cornechiniscus Maucci & 
Ramazzotti, 1981 


Diagnosis.—Echiniscidae with horn- 
shaped cirrus A. Granulation on body sur- 
face consists of cuticular swellings. Pseu- 
dosegmental plate present. Secondary clava 
hemispherical. Venter with longitudinal cu- 
ticular grooves. 

Type species.—Echiniscus cornutus Ri- 
chters, 1907 


Cornechiniscus madagascariensis Maucci, 
1993 
Figs. 1, 2, 3A-E 


Cornechiniscus madagascariensis Maucci, 
1993:383, figs. 1—4.—Antananarivo, 
Madagascar. 


Material examined.—Paratypes: two 
adult females, Madagascar ‘‘Antsirabe, 8/v/ 
1989, Cornechiniscus lobatus madagascar- 
iensis Maucci, paratipo, 19/v’’ mounted in 
polyvinyl-lactophenol (CT 13915, 13918). 
One adult female, India ‘“‘Himal [sic Hi- 
machal] Pradesh, 1500 m asl., moss from 
rocks, Sep 1976, lg. J. Btoszyk, (T. 4), Cor- 
nechiniscus”’ (ZMH). 

Comparative material.—Cornechiniscus 
lobatus (Ramazzotti, 1943): syntypes: Italy 
**T-44, Pseudechiniscus cornutus f. lobata, 
Montirone (Abano), 12-5-42, G. Ramazzot- 
ti, 1 Muta con 2 uova, 6 [sic 7] Individui, 


VOLUME 113, NUMBER 2 


forma lobata, (Muschi su roccia), -solo 
Faure-, 24, TIPO, P. cornutus f. lobata’”’ 
(CP): 

Female from India.—Body length 295.0 
wm excluding leg IV, width 136.0 wm; body 
width : body length, 1:2.17. Eyespot well- 
marked, black, subelliptical, 9.7 4m wide, 
situated in posterior part of head plate. 
Body color evenly translucent in preserved 
material. 

Dorsal plates thick. Dorsal surface in- 
cluding intersegmental lateral plates wholly 
covered with fairly coarse granulation that 
consists of pillar-shaped cuticular granules; 
adjacent granules distantly spaced, inter- 
connected with conspicuous cuticular stri- 
ae; granules near edge of each plate much 
smaller than those at median part; granules 
on scapular, segmental paired, median, in- 
tersegmental lateral (1 and 2), pseudoseg- 
mental, and terminal plates ca. 2.5 wm in 
diameter; granules on head plate and ante- 
rior part of neck plate ca. 1.5 wm in di- 
ameter; granules on posterior part of neck 
plate ca. 0.2 um in diameter. Lateral region 
of neck plate and basis for cirrus A only 
with very fine, densely distributed puncta- 
tions that consist of pillar structures of epi- 
cuticle. 

Head plate well-developed, with two zig- 
zag-sutures, showing facetted appearance. 

Neck plate conspicuous, subdivided ver- 
tically into three parts; posterior part of 
neck plate developed as a lobe which is nar- 
rowly overlapping anteriormost part of 
scapular plate. 

Scapular plate well-developed, anterior 
with poorly defined W-shaped sculpture; 
posterior to W-shaped sculpture, scapular 
plate shallowly subdivided vertically at 
middorsal line of body; an oblique cuticular 
line at each lateral side. 

Median plate 1 clearly subdivided into 
anterior and posterior parts; anterior part 
fairly larger than posterior part; anterior and 
posterior parts trapezoidal and inverted sub- 
triangular in shape, respectively. 

Intersegmental lateral plates 1 and 2 sim- 
ilar to each other in size and shape. 


481 


Segmental paired plates II and III well 
developed, similar to each other in size and 
shape, both with lateral segmental plates; 
small, triangular spines developed posterior 
part of lateral segmental plates II and III 
each (spines C and D). 

Median plate 2 clearly subdivided into 
anterior and posterior parts; anterior part 
subpentagonal, fairly larger than posterior 
part; posterior edge of posterior part sinu- 
ate. 

Median plate 3 well-developed, rhom- 
boidal, undivided. 

Pseudosegmental plate clearly subdivid- 
ed vertically at middorsal line of body; lobe 
on its posterior margin well-developed, un- 
paired, but shallowly bilobate, 11.4 pm 
high, 33 wm wide at base; tips of lobe 
smooth, without spine. 

Terminal plate with obvious, long inci- 
sion which is slightly effaced near middor- 
sal line of body; minuscule, triangular spine 
developed posteriormost of incision (spine 
BE). 

Leg plates developed on outer surfaces 
of legs I-III and dorsal surface of leg IV; 
each leg plate with coarse granulation sim- 
ilar to those on dorsal plates, i.e. adjacent 
granules interconnected with cuticular stri- 
ae; granules larger but sparse on median 
part of leg plate, smaller but dense on pe- 
ripheral part. Plate of leg IV without true 
dentate collar, but with very blunt triangular 
process posterior to leg plate. Sensory or- 
gan on leg I conical, 6.3 um long, 1.7 wm 
wide at base; sensory organ on leg IV pa- 
pillate, 6.9 um long, 5.1 wm wide at base. 

Dense patches of cuticular granulation 
situated below mouth opening, between 
each pair of legs I-III, and around gono- 
pore; these patches each without perceptible 
edge. Venter longitudinally costate, with 
some shallow, linear grooves that devel- 
oped from cuticular patch between leg I to 
gonopore, along with midline of body. 

Mouth opening and cephalic sensory or- 
gans situated ventrally. Internal cirrus on- 
ion-shaped, tapering sharply toward tip, 8.6 
wm long, 5.1 4m wide at base; external cir- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Figs ©. 
view; C, cirrus A and primary clava on the left, dorsal view; D, sensory organ I; E, sensory organ IV; E left 
side of segmental paired plate III and pseudosegmental plate; G, external claw of leg I; H, internal claw of leg 
IV. (A-E, G, H: ZMH; adult female; Himachal Pradesh, N India. F: CT 13918; paratype adult female; Antsirabe, 
Madagascar). 


rus stout, tapering gradually, 21.4 wm long, 
4.0 wm wide at base; both internal and ex- 
ternal cirri without true cirrophores, but 
their proximal portions swollen in appear- 
ance; furthermore, internal and external cir- 
ri each with hemispherical cuticular swell- 
ing beneath own base; both sides of cutic- 


Cornechiniscus madagascariensis Maucci, 1993. A, habitus, dorsal view; B, cephalic region, ventral 


ular swellings of external cirri interconnect 
each other with arcuate cuticular line. Sec- 
ondary clava hemispherical, 7.4 wm in di- 
ameter, situated nearer to internal cirrus 
than to external cirrus. 

Cirrus A clearly longer than external cir- 
rus, 28.6 wm long, 9.7% of body length, 6.9 


VOLUME 113, NUMBER 2 


Aad 


ae +) 


. 
> 
* 


Fig. 2. Cornechiniscus madagascariensis Maucci, 
1993. Habitus, dorsal view. (ZMH; adult female; Hi- 
machal Pradesh, N India). Differential interference 
contrast. Scale = 50 wm. 


14m wide at base, directed laterad; cirrus A 
provided with normally developed cirro- 
phore; primary clava situated just behind 
cirrophore of cirrus A, clavate, curved, 
apex directed posteriad, 8.0 wm long, 4.8 
14m in diameter at base. 

Claws I-III distally curved, whereas claw 
IV less curved compared with claws I-III; 
all claws thickened basally, without spur; 
internal claw slightly longer than external 
claw on all legs; claws I-III 12.8-14.3 wm 
long; claw IV clearly longer than claws I- 
Ill, ca. 18 wm long. 

Female gonopore normal, consists of ro- 
sette-like structure, 16 wm in diameter. 


Anus large, situated posteriad, near base of 
legs IV. 


483 


Emendatory notes on the paratypes.— 
Body 282.1 and 322.4 wm long. Neck plate 
tripartite vertically. Scapular plate orna- 
mented with ill-defined W-shaped sculp- 
ture. Cuticular grooves on venter well- 
marked. Small spines developed at posi- 
tions C, D, and E. Other characters in con- 
cordance with Maucci (1993). 

Remarks.—We examined two paratypes 
(engths 295.0 and 322.4 wm) of C. mada- 
gascariensis. The specimen from India is 
almost identical with the paratypes includ- 
ing the following important characters: pat- 
tern of the dorsal granulation, shape and 
size of each claw, morphology of the ce- 
phalic and leg sensory organs. Although we 
were able to examine only one specimen 
from India, there seems to be no problem 
for us to identify the specimen with C. mad- 
agascariensis. 

Maucci (1993) considered that C. mad- 
agascariensis is most closely related to C. 
lobatus Ramazzotti, 1943, and we agree 
with him on this point. He pointed out that 
the consistent absence of the dorsal and lat- 
eral body spines is one of the most impor- 
tant characters to distinguish C. madagas- 
cariensis from C. lobatus. We confirmed, 
however, that, as in C. lobatus, the small 
spines are actually present at positions C, 
D, and E also in C. madagascariensis (both 
in the paratypes and specimen from India). 

There are some reports of C. lobatus pro- 
vided with cuticular striae among dorsal 
granules, viz. Binda & Pilato (1972) from 
Sicily (Italy), Dastych (1979) from Afghan- 
istan, and Moon & Kim (1991) from South 
Korea. Furthermore, we have verified re- 
cently that the cuticular striae are rather 
conspicuous, especially in the terminal 
plate, in the syntypes of C. lobatus (Fig. 
3F). The cuticular striae were not men- 
tioned in the original description (Ramaz- 
zotti, 1943) and subsequent redescription 
(Maucci, 1979). 

From the evidence discussed above, the 
presence or absence of body spines and cu- 
ticular striae among granules cannot be 
used as taxonomic characters in discrimi- 


484 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 3. A-E: Cornechiniscus madagascariensis Maucci, 1993. A, anterior body, dorsal view; B, pseudo- 
segmental and terminal plates, dorsal view; C, posterior body, dorsal view; D, venter between legs I to III, 
showing cuticular grooves; E, mouth and cephalic sensory organs. (A, B, E: ZMH; adult female; Himachal 
Pradesh, N India. C, D: CT 13918; paratype adult female; Antsirabe, Madagascar). F: C. lobatus (Ramazzotti, 
1943), pseudosegmental and terminal plates, dorsal view (CT; a syntype adult female). A, B, D—F: differential 
interference contrast; C: phase contrast. Scales = 20 wm (A-D, F), 8 wm (E). 


nating C. madagascariensis from C. loba- important criterion to distinguish C. mada- 


tus. gascariensis from C. lobatus (Fig. 3F). 
In C. madagascariensis, however, gran- 
ulation on body surface consists of fairly Discussion 
large, distantly spaced granules (Fig. 3A— In the original description of C. mada- 


C), and this can be considered as the most gascariensis, Maucci (1993) implied that 


VOLUME 113, NUMBER 2 


this species may be referred to a subspecies 
of a cosmopolitan species, C. lobatus Ra- 
mazzotti, 1943. This also can be inferred 
from his specimen labels on microslides 
(see Material examined). Considering the 
present record from India which is quite far 
from Madagascar, however, the subspecific 
status cannot be supported. We are in agree- 
ment with Maucci (1993) who recognized 
C. madagascariensis as a good species 
based on the differences mainly discussed 
above. 

We have confirmed that the ventral cutic- 
ular grooves, which have been omitted from 
the previous descriptions, are also common- 
ly developed in many other Cornechiniscus 
species (unpublished data). In Pseudechin- 
iscus, which is a closely related genus to 
Cornechiniscus, the venter is ornamented 
with net-like pattern in many species (Das- 
tych 1984, Kendall-Fite & Nelson 1996). It 
is thus mentioned at present that the lon- 
gitudinal groove character-condition is 
unique for the genus Cornechiniscus. 


Acknowledgments 


We thank Dr. Hieronymus Dastych of the 
Zoologisches Institut und Zoologisches 
Museum der Universitat Hamburg for loan 
of the valuable specimen and comments on 
the manuscript, to Prof. Roberto Bertolani 
of the Dipartimento di Biologia Animale, 
Universita di Modena for the arrangements 
of loan of the paratypes, and to Prof. Clark 
W. Beasely of the McMurry University and 


485 


Prof. Diane R. Nelson of the East Tennes- 
see State University for comments on the 
manuscript. 


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Abe W., K. Utsugi, & M. Takeda. 1998. Pseudechin- 
iscus asper, a new Tardigrada (Heterotardigra- 
da: Echiniscidae) from Hokkaido, northern Ja- 
pan.—Proceedings of the Biological Society of 
Washington 111:843—848. 

Binda, M. G., & G. Pilato. 1972. Tardigradi muscicoli 
di Sicilia. (IV Nota).—Bollettino delle Sedute 
della Accademia Gioenia di Scienze Naturali in 
Catania, Serie IV 11:47—60. 

Dastych, H. 1979. Tardigrada from Afghanistan with 
a description of Pseudechiniscus schrammi, sp. 
nov.—Bulletin de la Société des Amis des Sci- 
ences et des Lettres de Poznan, Série D: Sci- 
ences Biologiques 19:99—108. 

. 1984. The Tardigrada from Antarctic with de- 
scription of several new species.—Acta Zoolo- 
gica Cracoviensia 27:377—436. 

Kendall-Fite, K., & D. R. Nelson. 1996. Two new spe- 
cies of tardigrades from Short Mountain, Ten- 
nessee, USA.—Zoological Journal of the Lin- 
nean Society 116:205—214. 

Maucci, W. 1979. I Pseudechiniscus del gruppo cor- 
nutus, con descrizione di una nuova specie (Tar- 
digrada, Echiniscidae).—Zeszyty Naukowe Un- 
iwersytetu Jagiellonskiego, Prace Zoologiczne 
25:107-124. 

. 1993 [dated 1990]. Prime notizie su tardigradi 
«terrestri» del Madagascar con descrizione di 
tre specie nuove.—Bollettino del Museo Civico 
di Storia Naturale di Verona 17:381—391. 

Moon, S. Y., & W. Kim. 1991. Systematic study on 
the tardigrades from Korea: new records of five 
tardigrade species from Korea.—Korean Jour- 
nal of Systematic Zoology 7:225—232. 

Ramazzotti, G. 1943. Nuova varieta del tardigrado 
Pseudechiniscus cornutus.—Natura (Milano) 
34:89—90. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


113(2):486—492. 2000. 


Ophryotrocha lipscombae, a new species and a possible connection 
between ctenognath and labidognath-prionognath eunicean 
worms (Polychaeta) 


Hua Lu and Kristian Fauchald 


(HL)Department of Biological Sciences, the George Washington University, 
Washington, DC 20052, U.S.A.; 
(HL, KF) Department of Invertebrate Zoology, National Museum of Natural History, 
Smithsonian Institution, Washington, DC 20560-0163, U.S.A. 


Abstract.—One new species of dorvilleid, Ophryotrocha lipscombae, is de- 
scribed based on material from the U.S. Atlantic slope. This new species is 
unique in its presence of 5 pairs of accessory plates, which are never found in 
any other dorvilleids, but are present in all labidognath-prionognath euniceans. 
A brief discussion of the phylogenetic position of Ophryotrocha lipscombae in 


the Eunicida is provided. 


Ophryotrocha has been extensively stud- 
ied by many polychaetologists (e.g., Jumars 
1974, Akesson 1984, Hilbig & Blake 1991, 
Eibye-Jacobsen & Kristensen 1994, Pleijel 
& Eide 1996); this may due to a number of 
factors: Ophryotrocha is common in both 
shallow and deep water environments; fur- 
thermore, it is easily cultured for study of 
reproductive and developmental patterns; in 
addition, its phylogenetic position within the 
Dorvilleidae has turned out to be interesting. 
As a part of the study of Eunicida phylog- 
eny, a new Ophryotrocha species was found 
by chance. In a jar containing mixed small- 
sized eunicean specimens labeled only as 
‘‘Runicidae’’, not only one eunicid larva was 
found, but also two species of Ophryotrocha 
and one juvenile lumbrinerid could be iden- 
tified. All specimens were observed using 
stereo and compound light microscopes; il- 
lustrations were made using a camera lucida. 

Abbreviation.—Mx refers to maxillae. 
The species description is in a similar for- 
mat as that used by Hilbig & Blake (1991). 


Ophryotrocha lipscombae, new species 
Figs. 1-4. 


Material examined.—North Atlantic 
Ocean, United States, 110 miles south of 


Woods Hole, Massachusetts, 29 Jul 1977, 
39°47'N, 70°40’W, Alvin Dive 773, STA 
DOS-1 (N-34), 1830 m (Holotype, USNM 
186571; 7 paratypes, USNM 186572). 
North Atlantic Ocean, United States, 110 
miles south of Woods Hole, Massachusetts, 
15 Jun 1976, 39°47'N, 70°40’'W, Alvin Dive 
658, 1830 m (Paratype, USNM 186573). 

Description.—Holotype complete with 
31 chaetigers, 2.75 mm long, 0.41 mm wide 
(Fig. 1A). Other complete specimens 0.90— 
2.43 mm long, 0.17—0.44 mm wide (Figs. 
1B, C; 2). Largest specimen incomplete 
with width of 0.77 mm. Body slender, oval 
in cross section. Chaetigers two times wider 
than long throughout body. One ciliary gir- 
dle present on each peristomial ring and 
each chaetiger. Color whitish in specimen 
collected in 1976 and brownish in speci- 
mens collected in 1977. 

Prostomium distally bluntly triangular in 
smaller specimens (Fig. 1B), rounded in 
larger ones (Fig. 1A); two times longer than 
wide in holotype. Two short and stout, 
knob-like antennae, not reaching anterior 
end of prostomium. No evidence of palps 
and eyes observed in any specimen. Prosto- 
mium same length and width as peristom- 
ium. Peristomium consisting of two apo- 


VOLUME 113, NUMBER 2 


dous, achaetigerous rings, each ring slightly 
shorter than adjacent chaetigers. Distinct 
lateral incisions present between peristo- 
mial rings; first peristomial ring distinctly 
fused with prostomium. 

Parapodia uniramous, with one acicular 
lobe; inferiormost simple chaetae in a sep- 
arate chaetal lobe. Dorsal and ventral cirri 
absent. Chaetae of three types: type 1, 3—4 
serrated cultriform simple chaetae (Fig. 1E) 
tapering to slender distal teeth in supra- 
acicular fascicle (Fig. 1D), their subdistal 
end finely serrated; type 2, 4—7 (the number 
variable in different body regions and dif- 
ferent specimens) heterogomph bifid com- 
pound falcigers (Fig. 1F) arranged in two 
rows in subacicular fascicle, their blades 
short and serrated; type 3, single, slender, 
simple cultriform chaeta (Fig. 1G) emerg- 
ing from long chaetal lobe in inferiormost 
position, forming an angle with acicula at 
the base. Each parapodium with one sharply 
pointed acicula distinctly deeper into para- 
podium than other chaetae. Pygidium rela- 
tively long, as long as last two chaetigers 


in 24-chaetiger specimen. Anal cirri easily ' 


broken, one small knob-like pair observed 
on 24-chaetiger specimen. No distinct mid- 
dle pygidial stylus present. 

Mandible rod-like, black, two pieces 
forming X-shape. In 0.17 mm-wide speci- 
men, distal end of each mandible piece with 
about 20 small teeth (Fig. 3B); in 0.44 mm- 
wide specimen, distal end of mandible 
without teeth (Fig. 3C). Maxillae K-type, 
with 8 paired pieces in roughly four rows; 
on each side, MX-I, II, and V-VII each 
forming one main row, while Mx-III and 
Mx-IV each as a separate row sitting out- 
side (Fig. 4A). Mx-I (Figs. 3A, 4A) heavy, 
generally ice-tong shaped and facing each 
other, with 3 large teeth in addition to main 
fang. Mx-II (Figs. 3A, 4A) thin plates, 
forming an arc over distal part of Mx-l. 
Mx-II with about 12 large teeth and a few 
irregularly placed small teeth in the middle. 
Mx-V to Mx-VIII (Fig. 4A) smaller than 
Mx-lIl, but structurally similar, plate shaped, 
with 4—8 large teeth and several small teeth. 


487 


Mx-III and Mx-IV elongate and fang- 
shaped with additional 1 or 2 smaller teeth, 
sitting anterior to Mx-II and outside of Mx- 
V. One pair of short carriers (Figs. 3A, 4A) 
fused to each other, and to posterior end of 
Mx-I. Two rows of 5 sclerotinized black ac- 
cessory plates (Fig. 4A) in erect position 
inside main maxilla row, each plate corre- 
sponds to Mx-II and Mx-V to VIII. Jaw 
structure symmetrical. 

Remarks.—Ophryotrocha lipscombae 
can be easily distinguished from other 
Ophryotrocha species by the presence of 3 
teeth in addition to the main fang on Mx-l, 
the presence of five accessory plates and 
the anterior end formed by the prostomium 
and the first peristomial ring. The above 
features are novel in dorvilleids, especially 
the accessory plates which have never been 
reported in the Dorvilleidae but are present 
in all other major families of the Eunicida 
(personal observation). When it is com- 
pared to other dorvilleids, such as O. akes- 
soni and O. geryonicola, O. lipscombae ap- 
pears to have a larger jaw apparatus for a 
similarly sized specimen; its maxillae can 
reach through the anterior four to five chae- 
tigers, while those of the other species usu- 
ally reach through the anterior two chaetig- 
ers. The over-all jaw structure is similar in 
all four Ophryotrocha lipscombae_ speci- 
mens dissected whose size ranged from 
0.17 mm to 0.77 mm wide. 

This species reaches a relatively large 
size (as wide as 0.77 mm) among dorvil- 
leids. While no gametes have been ob- 
served, we do not believe these specimens 
to be juveniles of any other reported euni- 
cean worms. The type material is deposited 
in the National Museum of Natural History, 
Smithsonian Institution, Washington D.C. 

Geographic distribution.—110 miles 
south of Woods Hole, Massachusetts, At- 
lantic slope. 

Etymology.—The species is named after 
Prof. Diana Lipscomb of George Washing- 
ton University for her many contributions 
to systematic biology. 


488 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


fn 


EPR G 


Fig. 1. Ophryotrocha lipscombae: A, anterior end, 31-chaetiger specimen, dorsal view; B, anterior end, 18- 
chaetiger specimen, ventral view; C, posterior end, 24-chaetiger specimen, ventral view; D, parapodium 3, 
anterior view; E, supra-acicular chaetae; K compound falciger; G, most inferior simple chaeta. 


VOLUME 113, NUMBER 2 


3.0 


2.5 


= NS 
on ro) 


Body size (mm) 
= 


0.5 


0.0 


16 18 20 22 


Fig. 2. 


Discussion 


The eunicean worms share a set of com- 
plex jaws composed of a ventral mandible 
and dorsal maxillae, and they are classified 
into five types based on jaw morphology: 
labidognath (Ehlers 1868) includes Eunici- 
dae, Onuphidae, Lumbrineridae and Hart- 
maniellidae. This type has a pair of well 
separated short carriers; Oenonidae have 
prionognath jaws (Colbath 1989, Fauchald 
& Rouse 1997) characterized by the pres- 
ence of a median plate inside the muscular 
bulb; Dorvilleidae has 4 rows of maxillae 
and is called ctenognath; xenognath (Mier- 
zejewski & Mierzejewska 1975) and pla- 
cognath (Kielan-Jaworowska 1966) are rep- 
resented only by fossil taxa. The above 
classification of eunicean jaws might be ar- 
bitrary if cladistic tree-thinking is applied. 


489 


30 
Chaetiger number of complete specimen 


24 26 28 32 


Correlation between body size and chaetiger number in Ophryotrocha lipscombae. 


Usually ctenognath is considered as mono- 
phyletic, and it is a sister group of labidog- 
nath-prionognath taxa (Kielan-Jaworowska 
1966, Kozur 1970, Jumars 1974, Orensanz 
1990). The assessment of jaw homology 
can be relatively easily made between 
prionognath and labidognath (Orensanz 
1990), but both are generally considered 
difficult to compare to the ctenognath. The 
overall phylogenetic construction of Euni- 
cida by Tzetlin (1980) may be problematic 
when a cladistic analysis is performed 
based on additional evidence (unpublished 
data); however, his scheme connecting the 
K-type Ophryotrocha jaw with those of la- 
bidognath-prionognath jaws is reasonable 
based on the following evidence: first, the 
Mx-I of labidognath-prionognath is homol- 
ogous with the Mx-I of both P- and K-type 


490 


0.05mm 


0.05mm 


0.1mm 


rien 2: 


Se 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


a 


Ophryotrocha lipscombae: A. jaw structure in dorsal view, only Mx-I, Mx-II and the carrier are 


depicted; B. mandible of 18-chaetiger specimen, dorsal view; C. mandible of 31-chaetiger specimen, dorsal view. 


Ophryotrocha, based on extensive studies 
of larval and adult jaw morphology of Dor- 
villeidae, Onuphidae and Oenonidae (H. L., 
unpublished data); second, the accessory 
plates, or attachment lamellae (Paxton 
1986), are present in all members of the 
labidognath-prionognath taxon (personal 
observation), and the presence of such 
plates in Ophryotrocha lipscombae is the 
first record for any dorvilleid. Accessory 
plate numbered 5 pairs in O. lipscombae, 3 
pairs in Oenonidae (Fig. 4B) and Lumbri- 
neridae (Fig. 4C), and 2—3 in Onuphidae 
and Eunicidae. All accessory plates are lo- 
cated median to the corresponding maxillae 
in erect position; no plate corresponds to 
Mx-lI. 

The presence of accessory plates may 
provide a substantial primary homology for 
O. lipscombae and the labidognath-prion- 


ognath taxa; the relationship will be tested 
in a systematic study of Eunicida based on 
morphology, ontogeny and fossil data. At 
present, the new species is considered as a 
member of Ophryotrocha based on its over- 
all morphological characters. Though the 
phylogenies of Ophryotrocha (Pleyel & 
Eide 1996) and Dorvilleidae (Jumars 1974, 
Westheide 1982, Hilbig & Blake 1991, Ei- 
bye-Jacobsen & Kristensen 1994) have 
been studied using various methods, better 
overall understanding might be gained from 
a relatively broader study of the phylogeny 
of the order Eunicida, including detailed 
studies of members of all major groups. 


Acknowledgments 


The authors would like to thank Linda A. 
Ward, Cheryl Bright, William Moser, and 


VOLUME 113, NUMBER 2 


491 


Fig. 4. A. Detailed jaw structure of 31-chaetiger specimen, Ophryotrocha lipscombae, in ventral view; note 
that left and right sides are cut apart, the carrier is broken, and the whole structure is displayed in an unnatural 
way in order to show the detailed structure; B. jaw structure in ventral view, Arabella iricolor (USNM 10355); 
C. jaw structure in ventral view, Lumbrineris zonata (USNM 30611). 


other colleagues from the Department of In- 
vertebrate Zoology, NMNH, Smithsonian 
Institution, and Diana Lipscomb and the 
Systematics Group, Department of Biolog- 
ical Sciences, the George Washington Uni- 


versity. We thank Stephen Gardiner and 
Kirk Fitzhugh for their reviews. HL is sup- 
ported by the Research Enhancement Funds 
and Weintraub Research Fellowship from 
the George Washington University, and re- 


492 


search contract 9822uu04278 from the 
Smithsonian Institution. 


Literature Cited 


Akesson, B. 1984. Speciation in genus Ophryotrocha. 
Pp. 299-316 in A. Fischer and H. D. Pfan- 
nenestiel, eds., Polychaete reproduction, pro- 
gress in comparative reproductive biology, vol. 
29, Fortschritte der Zoologie (Stuttgart: Gustav 
Fischer Verlag). 

Colbath, G. K. 1989. Revision of the family Lysare- 
tidae, and recognition of the family Oenonidae 
Kinberg, 1865 (Eunicida; Polychaeta).—Pro- 
ceedings of the Biological Society of Washing- 
ton 102:116—123. 

Ehlers, E. 1868. Die Borstenwiirmer (Annelida: Chae- 
topoda) nach systematischen und anatomischen 
Untereuchungen dargestellt, W. Engelmann, 
Leipzig, 748 pp. 

Eibye-Jacobsen, D., & R. M. Kristensen. 1994. A new 
genus and species of Dorvilleidae (Annelida, 
Polychaeta) from Bermuda, with a phylogenetic 
analysis of Dorvilleidae, Iphitimidae and Di- 
nophilidae.—Zoologica Scripta 23:107—131. 

Fauchald, K., & G. Rouse 1997. Polychaeta system- 
atics: past and present.—Zoologica scripta 
26(2):71-138. 

Hilbig, B., & J. Blake 1991. Dorvilleid (Annelida: 
Polychaeta) from the U.S. Atlantic slope and 
rise. Description of two new genera and 14 new 
species, with a generic revision of Ophryotro- 
cha.—Zoologica Scripta 20 (2):147-183. 

Jumars, P. 1974. A generic revision of the Dorvilleidae 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


(Polychaeta), with six new species form the 
deep North Pacific.—Zoological Journal of the 
Linnean Society 54(2):101—135. 

Kielan-Jaworowska, S. 1966. Polychaete jaw appara- 
tuses from the Ordovician and Silurian of Po- 
land and a comparison with modern forms.— 
Palaeontologia Polonica 16:1—152. 

Kozur, H. 1970. Zur Klassifikation und phylogene- 
tischen Entwicklung der fossilen Phyllodocida 
und Eunicida (Polychaeta).—Freiberger For- 
schungshefte 260 C:35-81. 

Mierzejewski, P., & G. Mierzejewski 1975. Xenognath 
type of polychaete jaw apparatuses.—Acta Pa- 
laeontologica Polonica 20:437—443. 

Orensanz, J. M. 1990. The Eunicemorph polychaete 
annelids from Antarctic and Subantarctic seas. 
With addenda to the Eunicemorpha of Argen- 
tina, Chile, New Zealand, Australia, and the 
southern Indian Ocean.—Antarctic Research 
Series 21:1—183. 

Paxton, H. 1986. Generic revision and relationships of 
the family Onuphidae (Annelida: Polychae- 
ta).—Records of the Australian Museum 38:1— 
74. 

Pleijel, EF, & R. Eide 1996. The phylogeny of Ophry- 
otrocha (Dorvilleidae: Eunicida: Polychaeta).— 
Journal of Natural History 30:647—659. 

Tzetlin, A. B. 1980. Ophryotrocha schubravyi sp. n. 
and the problem of evolution of the mouth parts 
in the Eunicemorpha (Polychaeta).—Zoologi- 
cheskii Zhurnal 59:66—676. 

Westheide, E. 1982. /kosipodus carolinensis gen. et sp. 
new., an interstitial neotenic polychaete from 
North Carolina U.S.A., and its phylogenetic re- 
lationships within Dorvilleidae.—Zoological 
Scripta 11:117—126. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


113(2):493—499. 2000. 


Aphrodita bisetosa (Polychaeta: Aphroditidae), a new species of sea 
mouse from the southeastern Pacific Ocean off central Chile 


Nicolas Rozbaczylo and Elba Canahuire 


(NR) Departamento de Ecologia, Facultad de Ciencias Bioldégicas, P. Universidad Cat6lica de 
Chile, Casilla 114-D, Santiago, Chile, e-mail: nrozbac @ genes.bio.puc.cl; 
(EC) Facultad de Ciencias Biol6gicas, Universidad Nacional Mayor de San Marcos, Lima, Pert; 
presently: Programa de Magister en Ciencias menci6n Zoologia, Escuela de Graduados, 
Universidad de Concepcién 


Abstract.—A new species of Aphroditidae, Aphrodita bisetosa from the 
southeastern Pacific Ocean off central Chile, is described. Specimens were 
collected in soft bottoms, 27—37 km from the coast line in front of Los Vilos 
(31°56’S) and Papudo (32°31'S), at 250-400 m depth. The new species was 
compared with A. magellanica Malard, 1891 from the Magellan area, Chile 
and A. alta Kinberg, 1855 from Rio de Janeiro, Brazil, and the Antarctic region, 
with which it appears to be more closely related. 


Aphroditids are commonly named “‘sea 
mice,’’ because of the thick mat of very fine 
notopodial fibers which form a felt-like 
covering over the dorsum, giving them a 
furry appearance. Although they are rather 
large and conspicuous inhabitants of marine 
soft muddy bottoms, they are frequently ab- 
sent from most collections because of the 
great depths at which they usually live, and 
in general they have been poorly studied. 
The most recent and comprehensive study 
of aphroditids was by Hutchings & McRae 
(1993) on species found in Australian wa- 
ters and the Indonesian Archipelago. 

There is almost no information about 
aphroditids of the southeastern Pacific 
Ocean along the Chilean coast. Only one 
species, Aphrodita magellanica Malard, 
1891, has been previously recorded from 
the Magellan area (49°S) south to Cape 
Horn (56°S) in southern Chile (Rozbaczylo 
1985); the specimens, had been collected 
during the Challenger Expedition (1873- 
1876), and identified by McIntosh (1885) as 
Aphrodita echidna Quatrefages. 

Four specimens of aphroditids collected 
by commercial shrimp trawlers at two sites 
off central Chile were given to the first au- 


thor for study; after examination, they were 
considered as belonging to a new species. 


Materials and Methods 


Specimens were collected by the shrimp- 
ing boat Goden Wind, as part of a benthic 
survey obtained 27—37 km from the coast 
between Los Vilos and Papudo, in October 
1976. At 2 of 5 stations sampled, specimens 
of aphroditids were found in sandy-mud 
bottom (Fig. 1). Additional information on 
the accompanying macrofauna collected 
during the trawls can be found in Andrade 
(1986). 

Polychaetes were fixed in 4% formalin 
and preserved in 70% ethanol. Figures were 
prepared with a drawing tube on a Wild M- 
5 stereoscopic microscope and a Leitz com- 
pound microscope. 

Type specimens of the new species are 
deposited in the National Museum of Nat- 
ural History, Smithsonian Institution, 
Washington, D.C. (USNM), and the Sala de 
Sistematica, Departamento de Ecologfa, 
Pontificia Universidad Catdélica de Chile, 
Santiago (SSUC). 


494 


Aphrodita bisetosa, new species 
Figs. 1-3 


Material examined.—Central Chile: in 
front of Los Vilos,. Stieilecay 3156'S, 
71°49'W, 300—400 m, H. Andrade, coll., 14 
Oct 1976, holotype (USNM 186512) and 
female paratype (SSUC 6868); in front of 
Papudo, St. 3, ca. 32°31'S, 71°47'W, 250— 
280 m, H. Andrade, coll., 14 Oct 1976, par- 
atype (N° 2 USNM 186513) and female 
paratype (SSUC 6869). 

Description.—Holotype. Body ovate, 
arched dorsally, widest at setigers 14—16, 
with tapering caudal region (Fig. 2a); 27 
mm long, 16 mm wide, excluding setae, 
with 41 setigers. Dorsum with thick felt, ap- 
proximately 1.5 mm thick at middle region 
of body, with fine sediment entrapped giv- 
ing it greyish appearance. Ventral surface of 
body whitish, covered with minute spheri- 
cal papillae (Fig. 2d). 

Prostomium small, rounded with pair of 
ocular areas, pale to light brown in color, 
located on slightly raised prominences (Fig. 
2b). Median antenna with basal ceratophore 
and elongated style, slightly shorter than 
prostomium, attached dorsally, near anterior 
border of prostomium. Palps_ biarticulate, 
finely papillated, wide basally, tapering 
gradually, extending approximately to sixth 
setiger. Facial tubercle minutely papillated, 
approximately half of length of prostomi- 
um, partly hidden by palps dorsally and ex- 
tending ventrally as digitiform process over 
mouth (Fig. 2b, c). 

Elytra (Fig. 2e, f) 15 pairs, completely 
hidden by dorsal feltage, on setigers 2, 4, 
5, 1, 9, sex 25, 2831; imbricate, *complete- 
ly covering dorsum, semi-transparent; 
smooth except for few scattered microscop- 
ic digitiform papillae (Fig. 2g), mostly con- 
centrated on inner lateral area of upper sur- 
face and less on posterior area of elytra. 
First pair of elytra smallest, gradually in- 
creasing in size to approximately pair 7-9, 
then decreasing posteriorly. First and sec- 
ond pair of elytra ovate, longer than wide, 
with elytrophores attached centrally and lat- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


erally, respectively; following elytra wider 
than long; last three pairs noticeably longer 
than wide; last pair smaller. 

Dorsal tubercles from segment 6 to 30 
provided with branched fimbriated papillae 
on posterolateral margins (Fig. 3d). 

First setiger or tentacular segment with 
elongated, uniramous parapodia, flattened, 
projecting anteriorly and laterally to prosto- 
mium (Fig. 2b), with one tuft of fine silky 
fibers emerging dorsally forming the dorsal 
felt, and 3 fascicles of fine, faintly irides- 
cent, mud-covered capillary setae; one tuft 
supra-acicular and two others sub-acicular; 
with few scattered papillae covering para- 
podium (Fig. 2b). Parapodial dorsal and 
ventral cirri with cylindrical cirrophores 
and subulate styles. Dorsal cirri approxi- 
mately one-third length of palps. Ventral 
cirri slightly shorter than dorsal. 

Following setigers with biramous para- 
podia. Second setiger with first pair of el- 
ytra. Notopodia rectangular, with one tuft of 
fine silky fibers forming dorsal felt, 2 supra- 
acicular fascicles of stout acicular setae, 
with hooked tips, similar in shape and dis- 
tribution to that of parapodium 3, and a sub- 
acicular fascicle with capillary notosetae 
forming lateral fringe, and on posterior sur- 
face of notopodia at level of acicula, a small 
tuft of fine silky fibers forming dorsal felt. 
Neuropodia cylindrical, covered with spher- 
ical papillae; tip of acicula emerging at dis- 
tal end of neuropodial lobe. Neurosetae, 
brown, arranged in 3 tiers: upper consisting 
of 2—3 acicular setae, middle 1—2 acicular 
setae, and lower of numerous capillary se- 
tae (Fig. 31), slender bipinnate and spirally 
twisted, with two rows of thick teeth that 
continue in subdistal region as fine spines, 
and with distal end smooth. Ventral cirri su- 
bulate, located near base of neuropodium, 
about one and one-half length of neuropo- 
dia (Fig. 2d). 

Third setiger similar to second, but with 
dorsal cirri instead of elytra. Dorsal cirri 
long, subulate, approximately 3-4 times 
length of ventral cirri. 

Parapodia of middle region of body (Fig. 


VOLUME 113, NUMBER 2 


2 
< 
LL] 
O 
O 
Oo 
LL. 
O 
<x 
0. 


Fig, 1. 


3a, b) with notopodial lobe large, nearly tri- 
angular, with dorsal cirri with cirrophores 
large, basally bulbous, projecting on pos- 
terior faces of notopodia; styles subulate, 
long, slender, smooth, directed dorsally; 
dorsal cirri approximately 3—4 times longer 
than ventral cirri; notoacicula stout, light 
brown, emerging at vertex of distal end of 
notopodial lobe. 

Notopodia with tufts of fine silky fibers 
emerging dorsally forming dorsal felt, with 


495 


LOS VILOS 


PAPUDO 


VALPARAISO 


Map showing the stations (*) where specimens of Aphrodita bisetosa, new species, were found. 


hooked ends (Fig. 31); arranged in three 
main groups on cirrigerous segments, one 
above and one below upper fascicle of su- 
pra-acicular notosetae, and third one on 
posterior surface of notopodia at level of 
acicula; on elytrigerous segments appearing 
in two main groups, one between upper and 
lower fascicle of long supra-acicular noto- 
setae, and one on posterior surface of no- 
topodia at level of acicula. Three fascicles 
of notosetae present, one sub-acicular and 


496 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


i, @ees @\\\ 
\ AST 
NC |i 


| 
| 


Fig. 2. Aphrodita bisetosa, new species (Holotype USNM 186512). a, dorsal view, whole animal; b, dorsal 
view of prostomium and first segment, with first pair of elytra omitted (right palp and dorsal parapodial cirri 
missing); c, facial tubercle (Paratype SSUC 6869), antero-lateral view; d, anterior end (Paratype SSUC 6869), 
ventral view; e, anterior end (Paratype USNM 186513), dorsal view, with proboscis everted and feltage set 
aside, showing first three pairs of elytra, style of median antenna missing; f, ninth left elytron, inner surface 
(Paratype USNM 186513); g, papillae from ninth elytron; h, posterior segments around the anus (Paratype SSUC 
6869), dorsal view. Scales = 10 mm for a; 5 mm for d, e, f; | mm for b, h; 0.05 mm for g. 


VOLUME 113, NUMBER 2 497 


eg fle 


Fig. 3. Aphrodita bisetosa, new species, a, parapodium 14 (Paratype SSUC 6868), in anterior view (tufts of 
silky fibers are shown cut off); b, same parapodium in posterior view (f. p. = fimbriated papilla; s. p. = segmental 
papilla); c, conical papillae from notopodium of same parapodium; d, fimbriated papillae (Paratype SSUC 6869), 
from segments 6, 8, 10, and 12, respectively; e, capillary notoseta from parapodium 14; f, g, acicular notosetae 
of stout type, with hooked tips, from parapodium 14; h, acicular notoseta of stouter type, with straight tip, from 
parapodium 18; i, fine silky fiber, from parapodium 20; j, acicular neuroseta, with heavily bearded end, from 
lower tier, parapodium 22; k, acicular neuroseta, partially bearded, from lower tier, parapodium 14; 1, capillary 
bipinnate neuroseta, spirally twisted, from lower tier, parapodium 2; m, capillary, partially bipinnate neuroseta, 
from lower tier, parapodium 29. Scales = 5 mm for a, b, j; | mm for d, h; 0.5 mm for c; 0.01 mm for k, 1, m; 
0.005 mm for e, f, g, 1. 


498 


two supra-acicular. Sub-acicular fascicle 
made up of numerous iridescent capillary 
setae (Fig. 3e), extending laterally, with dis- 
tal end straight and pointed; most covered 
with fine silky fibers and fine mud resem- 
bling cotton in appearance. Two supraci- 
cular fascicles made up of 2 kinds of acic- 
ular setae emerging through dorsal feltage: 
many stout acicular setae with hooked tips 
(Fig. 3f, g) and 1—2 stouter acicular setae 
with straight tips (Fig. 3h), short, conical, 
dark brown to blackish, spine-like, extend- 
ing dorso-posteriorly (tips may be broken 
off), with fine silky fibers at base. In upper 
fascicle, stout protective notosetae present 
(Fig. 2a), long, brown-colored basally and 
shiny golden light brown color distally, ta- 
pering gradually, with flexible hooked tips 
(may be broken off), sometimes partially 
covered with fine silky fibers. In lower fas- 
cicle, stout protective notosetae short with, 
distal region having fine silky fibers and 
covered with fine mud presenting finger- 
like appearance, peppered with ferruginous 
color. 

Notopodia covered with two kinds of pa- 
pillae: globular, small, few, scattered on su- 
bacicular area near distal margin of noto- 
podial lobe, and conical (Fig. 3c), larger 
and more abundant, scattered on basal area 
of notopodial lobe. 

Neuropodia (Fig. 3b), cylindrical, ending 
distally in three step-like lobes, covered 
with minute spherical papillae. Ventral cirri 
small, subulate, located in middle region of 
neuropodia, approximately one-third the 
length of dorsal cirri. 

Neurosetae stiff, stout, dark brown, 
slightly curved distally, with heavily beard- 
ed ends (Fig. 3j), which can be broken; 
pointed subdistal tip visible through beard, 
exposed when the distal tip of beard ends 
lost (Fig. 3k). Neurosetae arranged in three 
tiers with 2 setae in upper, 2 in middle and 
4—6 in lower tier in anterior parapodium 
(setiger 4), 2 in upper, 3—4 in middle and 
5—9 in lower tier in middle parapodium (se- 
tiger 15), and 2 in upper, 2 in middle and 
7 in lower tier in parapodium of posterior 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


region of body (setiger 25). Upper group 
with stoutest and longest setae; lowest most 
slender and shortest. Neuroacicula, stout, 
light brown, emerging through upper lobe. 

Ventral neurosetae of posterior region of 
body, capillaries, with two rows of teeth 
subdistally (Fig. 3m), and distal region 
smooth with tips from slightly curved to 
straight. Neurosetae of more posterior setig- 
ers all capillaries. 

Anus (Fig. 2h) located dorsally, anterior 
to 2 small segments. Without anal cirri. 

Variation.—Additional material exam- 
ined ranges from 47—80 mm long, 20-35 
mm wide, excluding setae, with 42 setigers. 
Body varies from ovate to elongated in 
largest specimens. Fascial tubercle extends 
ventrally as digitiform process over mouth 
in all except one specimen (paratype SSUC 
6868). Only one specimen with everted 
proboscis (Fig. 2e) with opening surround- 
ed by numerous, leaflike, dichotomously 
branched papillae; chitinous jaws lacking. 
Elytra in largest specimens are light brown 
to cream in color; irregularly stained fer- 
rugineous to olive in colour on inner sur- 
face (Fig. 2f). Holotype without segmental 
papillae, but present in largest specimens as 
small knobs on posterior faces of parapodia 
between rami, on segment 14, and subse- 
quent setigers (Fig. 3b). 

In paratypes neurosetae varied as fol- 
lows: 2 setae in upper, 2—3 in middle and 
3—6 in lower tier in anterior parapodium 
(setiger 4), 2—3 in upper, 3—5 in middle and 
10—12 in lower tier in middle parapodium 
(setiger 15), and 2—3 in upper, 3—4 in mid- 
dle and 7—11 in lower tier in parapodium 
of posterior region of body (setiger 25). 

Etymology.—The species name bisetosa 
is derived from Latin bi-meaning double, 
and setosa meaning with bristles, because 
of the presence of two kinds of notosetae 
which emerge through the dorsal feltage. 

Distribution.—The species has been 
found at two sites off Central Chile: between 
27 and 37 km off the coast in front of Los 
Vilos, 300—400 m and in front of Papudo, 
250—280 m depth, in sandy-mud bottoms. 


VOLUME 113, NUMBER 2 


Remarks.—The new species Aphrodita 
bisetosa is characterized by two kinds of 
acicular setae emerging through dorsal fel- 
tage: numerous stout setae with hooked 
tips, and some stouter, short, dark brown, 
spine-like setae, with straight tips, forming 
two distinct rows of protective notosetae 
along each side of body. The acicular no- 
tosetae appear in two fan-shaped fascicles 
of different length: an upper fascicle of long 
setae, shiny golden brown-colored distally, 
extending dorsomedially, nearly touching 
medially, and a lower fascicle of short setae 
extending backwards, bearing a mud cover, 
peppered with ferruginous colour, present- 
ing finger-like appearance. 

Aphrodita bisetosa appears to be most 
closely related to A. magellanica Malard, 
1891, from the Magellan area and to A. alta 
Kinberg, 1855 from Rio de Janeiro, Brazil, 
and the Antarctic region. A. magellanica is 
the most similar to A. bisetosa n. sp. Both 
species have two rows of acicular notosetae 
along each lateral region of the body that 
are short, brown, and spine-like with 
straight tips emerging through the dorsal 
feltage. Both species have similar neurose- 
tae of the acicular type, brown, with beard- 
ed ends (which can be broken), and in both 
species the dorsal felt is formed of very fine 
silky fibers with hooked tips; the lateral re- 
gions are of ferruginous color. A. bisetosa 
differs from A. magellanica by having two 
kinds of protective notosetae; its spine like 
acicular notosetae are conical toward the 
tip, 1-2 of these notosetae are present in 
each supracicular tuft; ventral cirri of para- 
podia lack papillae; and the body has 41— 
42 setigers. In A. magellanica, in contrast, 
has only one kind of protective notosetae. 
The spine like notosetae are flattened to- 
ward the tip, 2—3 of these notosetae can be 
present in each supracicular tuft; ventral cir- 
ri of parapodia possess numerous papillae 
and the body has about 35 segments. 

Aphrodita bisetosa resembles A. alta in 
that the protective notosetae end in a hook, 
the acicular neurosetae of the middle region 
of the body are slightly curved and have 


499 


heavily bearded ends and in both species 
the proboscis bears leaflike papillae sur- 
rounding the opening. A. bisetosa differs 
from A. alta by the following characteristics 
present in the latter species: only one type 
of protective notosetae that do not project 
through the felting, only one type of orna- 
mentation in capillary neurosetae of second 
setiger; without segmental papillae and pro- 
boscis with jaws. 


Acknowledgments 


We are greatly indebted to Dr. Héctor An- 
drade who kindly made specimens available 
for study. We are specially grateful to Mrs. 
Clara Yanez for her skill and patience in pro- 
ducing the excellent illustrations of the new 
species. Two anonymous referees are thanked 
for their helpful comments and suggestions 
that greatly improved the manuscript. 


Literature Cited 


Andrade, H. 1986. Observaciones bioecoldgicas sobre 
invertebrados demersales de la zona Central de 
Chile. Pp. 41-56 en La Pesca en Chile. P. Ar- 
ana, ed., Escuela de Ciencias del Mar, Univer- 
sidad Cat6élica de Valparaiso. 

Hutchings, P, & J. McRae. 1993. The Aphroditidae 
(Polychaeta) from Australia, together with a re- 
description of the Aphroditidae collected during 
the Siboga Expedition.—Records of the Austra- 
lian Museum 45(3):279—363. 

Kinberg, J. G. H. 1855. Nya slagten och arter af An- 
nelider.—Ofversigt af Kongliga Vetenskaps- 
Akademiens Forhandlingar (Stockholm) 12: 
381-388. 

Malard, A. E. 1891. Sur une nouvelle aphrodite du Cap 
Horn, décrite a tort par M. McIntosh comme A. 
echidna (De Quatrefages).—Bulletin de la So- 
ciété Philomathique de Paris Sér. 8, 3:125—127. 

McIntosh, W. C. 1885. Report on the Annelida Poly- 
chaeta colleted by H.M.S. Challenger during 
the years 1873—1876. Jn Report on the Scientific 
Results of the Voyage of H.M.S. Challenger 
during the years 1873-1876 under the Com- 
mand of Captain George S. Nares, R.N., ER.S. 
and the Late Captain Frank Tourle Thomson, 
R.N., Zoology, 12(34):1-554, pls. 1-55, and 
la—39a. 

Rozbaczylo, N. 1985. Los Anélidos Poliquetos de 
Chile. Indice sinonimico y distribuci6n geo- 
grafica de especies.—Monografias Biolégicas 
3:1-284. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


113(2):500-—513. 2000. 


Exogone breviantennata Hartmann-Schrdéder, 1959 
(characters emended) (Annelida: Polychaeta: Syllidae), a new record 
for the Bahamas with a key to selected Exogone species 


Robert Zottoli and Charlene D. Long 


(RZ) Fitchburg State College, Fitchburg, Massachusetts 01420, U.S.A.; 
(CDL) The Buffum Group, 21 % Buffum Street, Salem, Massachusetts 01970, U.S.A. 


Abstract.—The syllid polychaete Exogone breviantennata Hartmann-Schréd- 
er, 1959 is reported from the Bahamas, where it was collected from epiphytic 
communities on submerged red mangrove roots in Osprey Lake and Reckley 
Hill Pond located on San Salvador Island. The external morphology of both 
paratype and Bahamian specimens is described and species characters emended. 
Observations on reproduction, biology, habitats and distribution are reported. 
The essential characteristics used to identify the genus Exogone are discussed 
and a key to Caribbean species with a dorsal cirrus on setiger 2 is provided. 


Several polychaete species were collect- 
ed from algal covered red mangrove roots 
in Osprey Lake and Reckley Hill Pond, 
landlocked bodies of water on San Salvador 
Island in the Bahamas. One of them, re- 
ported here for the first time from the Ba- 
hamas, E. breviantennata WHartmann- 
Schréder, 1959, has features unique to in- 
terstitial polychaetes normally associated 
with sand or mud. The species characters 
are emended based on examination of par- 
atype material and the specimens collected 
from the Bahamas. Included is a discussion 
of the important characters used to identify 
the genus Exogone and a key to species re- 
lated to E. breviantennata that are found in 
the Caribbean. 


Materials and Methods 


Two subtidal Rhiziophora mangle prop 
roots, extensively colonized by algae, were 
collected from Osprey Lake, San Salvador 
Island, Bahamas, 5 May 1996, and from 
Osprey Lake and Reckley Hill Pond (some- 
times given as Reckley Hill Settlement 
Pond), 19 Jun 1997. The polychaetes were 
removed, relaxed in 5% magnesium sulfate 
and fixed in a 5% formalin/seawater solu- 


tion. Specimens were transferred to 70% 
ethanol for long-term storage. Some of the 
specimens were mounted in Hoyer’s solu- 
tion in order to clear body tissue so that 
teeth and setae would be clearly visible 
(Zottol & Long 1998) or in Glycrogel 
Mountant (Gurr 1962), which allowed close 
examination of body surfaces. 
Measurements were made under com- 
pound or dissecting microscopes with a cal- 
ibrated grid. Body length was measured 
from the tip of the prostomium to the end 
of the pygidium; for twisted specimens, the 
grid was rotated along the body. Body 
width was measured from the tip of the 
right neuropodium to the left across the dor- 
sal surface of the fourth setiger. 
Comparative material was borrowed 
from the Zoologisches Museum, Hamburg 
(ZMH) and the National Museum of Nat- 
ural History, Smithsonian Institution 
(USNM), Washington, D.C., U.S.A. 


Identifying Specimens of the Genus 
Exogone 


Most syllids, because of their small size, 
are difficult to identify, requiring careful 
examination under a compound micro- 


VOLUME 113, NUMBER 2 


scope. For this and other reasons, Fauchald 
(1977:79) stated: “‘the identification of syl- 
lids [is] a time-consuming occupation.”’ 
Larger specimens should have the pharynx 
isolated and appropriate parapodia removed 
for examination under a compound micro- 
scope. Smaller specimens should be cleared 
in an appropriate medium such as Hoyer’s 
solution (see Zottoli & Long 1998) or 
mounted permanently in Glychrogel Moun- 
tant (Gurr 1962). During examination, it is 
important to keep in mind that what you see 
may be an artifact resulting from the degree 
of relaxation prior to fixation, the type of 
preservative used, or the length of time 
stored, especially with regard to the size 
and shape of soft parts and the appearance 
of setae (see Riser 1991:214). The quality 
of equipment, which determines the fine- 
ness of detail seen, and the experience of 
the investigator, may also influence what is 
observed. These factors may be responsible 
for the lack of detail in many original de- 
scriptions, which in some cases are now 
considered incomplete because they did not 
illustrate or discuss what are now consid- 
ered essential characters. 

The following is a discussion, on a char- 
acter by character basis, of features com- 
monly used to identify species of Exogone. 
Size should not be used to classify speci- 
mens because local and regional variations 
have been inadequately studied for most 
polychaetes. The degree of fusion of the 
palps must be viewed both dorsally and 
ventrally. The number of antennae (if not 
lost) is reliable, although antenna shape, 
length, and placement on the prostomium 
need to be measured on a large number of 
specimens; antennae are easily distorted 
due to relaxation and fixation techniques. 
Russell (1991) refuted the use of the num- 
ber of muscle cell rows of the proventric- 
ulus as a diagnostic character in the family 
Syllidae, and we have found sufficient var- 
iation in E. breviantennata to agree; the 
number of setigers in which the proventric- 
ulus is found varies according to how far 
the pharynx is retracted, thus making this a 


501 


questionable character for species distinc- 
tion. For comments regarding the useful- 
ness of nuchal organs in identifying some 
syllids, see Riser (1991). The shape of the 
neuropodia and the shape and position of 
the dorsal, ventral, and tentacular cirri are 
easy to determine even in small specimens. 
The presence or absence of dorsal cirri on 
setiger 2 in adult specimens is an important 
and stable species characteristic; however, 
the apparent absence of dorsal cirri on se- 
tiger 2 should be verified on several speci- 
mens as the structures can easily be de- 
tached. All parapodia should be scanned for 
modified setae of any kind and all setae 
should be carefully examined dorsally, lat- 
erally, and ventrally under oil immersion 
along the entire length of the animal. Setae 
of the same type can appear quite different 
from setiger to setiger depending on the de- 
gree of rotation. For example, blades may 
appear smooth if viewed from either a dor- 
sal or a ventral aspect but toothed from a 
lateral aspect. The tips of setal blades that 
appear unidentate may, on closer examina- 
tion, be seen to be bidentate. The type and 
extent of denticulation or spination on the 
setal shafts also become more apparent with 
increasing magnification. The importance 
of care in observing setae is illustrated in a 
comparison of figures 75—78 in the original 
description of E. breviantennata by Hart- 
mann-Schroéder (1959), which show no dec- 
oration on the setae, and, yet, painstaking 
reexamination of a type specimen revealed 
obvious decoration (blades and shafts). Fi- 
nally, every specimen in a collection should 
to be checked individually, to ensure that 
all belong to the same species (Perkins 
1981). 


Genus Exogone @Orsted, 1845 


Key characters.—Small worms (usually 
less than 3 mm in length but up to 8 mm); 
palps sometimes fused; three antennae and 
one pair of tentacular cirri; dorsal and ven- 
tral cirri well developed but shorter than se- 
tal lobes; all appendages papilliform or 


502 


ovoid; eversible pharynx with single ante- 
rior tooth (Fauchald 1977, Pascual et al. 
1996). San Martin (1991) reviewed this ge- 
nus based on his work with Cuban syllids 
and expanded the number of key characters 
to include presence or absence of eyes, ex- 
tent of palpal fusion, presence of two long 
anal cirri, smooth body surface, and repro- 
ductive habits. 

In summary and in decreasing order of 
reliability, we use the following characters 
to distinguish various species in the genus 
Exogone: presence or absence of a dorsal 
cirrus on setiger 2; position, shape, and 
length of the antennae; shape, size, devel- 
opment and placement of eyes; placement 
and relative length of the proboscis and pro- 
ventriculus; and shape and denticulation 
and spination of setae and acicula. 

Biology of the genus Exogone.—Mem- 
bers of this genus have been reported from 
soft and hard bottoms (Uebelacker 1984: 
30-37 through 30-43), associated with 
sponges (Pascual et al. 1996) and on un- 
shaded algal flats dominated by Caulerpa 
verticillata and Halimeda opuntia f. trilo- 
bata, reported by Russell (1991). In the Ca- 
nary and Madeira Islands, six species of 
this genus comprised 6.6% of all poly- 
chaetes collected from sponges. When they 
are associated with living in sponges, some 
members of this genus exhibit morpholog- 
ical characters such as decrease in the blade 
length of compound setae and/or blades 
fused with the shafts, as is the case in other 
syllid genera, e.g., Haplosyllis (Pascual et 
al. 1996). However, Exogone species that 
are not exclusively associated with sponges 
do not show these adaptations, e.g., E. brev- 
iantennata (see below). Members of the 
subfamily Exogoninae usually undergo re- 
productive metamorphosis that includes 
changes in various external structures (eyes, 
antennae, parapodial structures, and setae) 
and internal structures (nephridia and mus- 
culature) (Schroeder & Hermans 1975, Ku- 
per & Westheide 1998). Based on the lit- 
erature and our observations, E. brevian- 
nenata appears to be an exception in that 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


reproductive modifications have never been 
observed for either sex. 

Of interest is the fact that, based on the 
original descriptions of 34 species in this 
genus, there is about a one to one ratio of 
those that have a dorsal cirrus on setiger 2 
and those that lack this character. Of the 
latter group, many species were reported to 
have a minute median antenna; very few of 
those species were said to have a median 
antenna longer than the palps. Of the for- 
mer group, most species had a median an- 
tenna that reached at least to the tip of the 
prostomium, and many were much longer, 
reaching to or beyond the palps. An excep- 
tion to this is E. breviantennata, which has 
a shorter median antenna, not even reaching 
to the tip of the prostomium. 


Exogone breviantennata Hartmann- 
Schroder, 1959 
characters emended 
Figs. 1-5 


Exogone breviantenna Hartmann-Schréder, 
1959:125-127, figs. 75-78; San Martin, 
1991:728-729, 730-731, fig. 8; Nunez et 
al., 1992:47, fig. 3; Pascual et al., 1996: 
710; 77. Aablex2: 

Exogone occidentalis Westheide, 1974:113; 
Russell, 1991:59—61, fig. 4 (emended). 


Material examined.—El Salvador: para- 
type P-14590, ZMH; Belize: Twin Cays, 
West Bay, D. E. Russell, Nov 1993, Sta. M- 
4, 3 specimens labelled Exogone occiden- 
talis, USNM 102085; and Bahamas: San 
Salvador Island (from amongst filaments of 
the algae Batophora sp. and Vaucheria sp. 
taken from the prop roots of Rhiziophora 
mangle), Oyster Pond, R. Zottoli & C. D. 
Long, 6 specimens, 4 May 1996; Osprey 
Lake, R. Zottoli, 13 specimens, 5 May 
1996, USNM 186554; Reckley Hill Pond, 
R. Zottoh, 2 specimens, 5 May 1996 and 
17 specimens, 19 Jun 1997, USNM 
186555-186561; and southern edge of Bim- 
ini Lagoon, from plastic mesh sponges an- 
chored and floating in 1-3 feet of water, 
collected at intervals up to 11 months from 


VOLUME 113, NUMBER 2 


1970-1971, Amy Schoener, 12 specimens 
(labelled as Exogone verugera), USNM 
51542, and 5 specimens, USNM 51545. 
Key characters.—Syllid in the genus Ex- 
ogone with dorsal cirri on all setigers and 
median antenna shorter than prostomium. 
Description of the paratype.—Specimen 
1.95 mm long, 0.2 mm wide, 24 setigers 
(Fig. 1). New segment, without setae, form- 
ing between last setiger and pygidium. Pro- 
stomium about four times as wide as long; 
anterior and lateral margins rounded. Eyes 
not discernible. In the original description 
of type material, Hartmann-Schréder (1959) 
described and figured four eyes, one at each 
corner of a trapezoid, in the posterior half 
of the prostomium. Single median and two 
lateral digitiform antennae. Antennae simi- 
lar in shape to tentacular, dorsal, and ventral 
cirri. Palps completely fused dorsally, ex- 
tend anteriorly well beyond the prostomi- 
um. Tentacular segment short, about three 
times as wide as long; clearly separated 
from the prostomium, but not from the fol- 
lowing first setiger. One pair of lateral ten- 
tacular cirri, about 0.02 mm long and 0.01 
mm basal width. Uniramous parapodia 
(neuropodium only) as short lobes, similar 
on all setigers. Digitiform dorsal (about 
0.024 mm long and 0.02 mm basal width) 
and ventral (about 0.024 mm long and 
0.012 mm basal width) cirri. One dorsal 
simple seta (3 wm basal width) per setiger; 
terminates with short spines (Fig. 2A); tip 
slightly concave. One heterogomph spiniger 
(Fig. 2B) just below the dorsal simple seta; 
blade length 20 to 30 wm, increases from 
anterior to mid-body and then decreases 
posteriorly; blade with about 25 deeply cut 
teeth, rounded at the base. Terminal end of 
shaft where it meets the blade with cusps. 
Two to four heterogomph falcigers (Fig. 
2C) below the spiniger, blades 5-10 pm 
long, bidentate (first tooth much smaller 
than second), with about five deeply cut 
teeth; base rounded. Generally four falci- 
gers per setiger from anterior to mid-body 
setigers, then posteriorly diminishing to 
two; blade length increases in mid-body 


503 


and then decreases again (Table 1). Outline 
of single aciculum visible in some setigers; 
details not clear. Ventral simple seta (base 
width about 3 pm) in setigers 14—24; tip 
strongly hooked and with short spines (Fig. 
2D). Pygidium with two cirn, 0.13 mm 
long and 0.03 mm long. Pharynx short, re- 
tracted, extends forward from posterior por- 
tion of setiger 2; single dorsally attached 
tooth. Proventriculus 0.13 mm long and 0.1 
mm wide, extends from setiger 2 to 5, with 
about 13 muscle rows. 

Description of San Salvador Island spec- 
imens.—The following description is based 
on 32 complete specimens and 13 attached 
larvae, with 5—31 setigers, and ranging in 
length from 0.34 to 2.7 mm and 0.09 to 
0.22 mm in width; see Table 2 and Figs. 3— 
5. Prostomium (Fig. 3A) about twice as 
wide as long. Two pairs of dark red eyes 
placed lateral to the antennae; anterior- 
most, largest pair about half way along lat- 
eral antennal length; posterior-most pair at 
antennal bases. No obvious modification of 
eyes in sexually mature specimens. Median 
antenna 0.01 to 0.026 mm long and two 
lateral antennae 0.01 to 0.015 mm long, all 
short and originating slightly anterior to 
prostomial-tentacular segment junction. 
Tentacular segment about four times as 
wide as long. Tentacular cirri 0.01 to 0.024 
mm long (Figs. 3, 4). Digitiform dorsal 
(0.02 to 0.03 mm long) and ventral (0.01 to 
0.03 mm long) cirri on all setigers. 

Setal structure and pattern similar for all 
specimens examined. Dorsal simple seta 
(2.4 to 3 wm base width) terminates bluntly 
with about 15 short spines. Heterogomph 
spiniger blade length 17 to 28 wm; blades 
with approximately 30 teeth; spines on ter- 
minal end of shaft. One to four hetero- 
gomph falcigers (blade length 5 to 9 pm) 
and spines on terminal end of shaft. One 
ventral simple seta on posterior segments, 
found in all setigers in attached larvae (6— 
13 setigers) but are lost anteriorly in later 
stages. Single, smooth aciculum (6 um bas- 
al width) in each setiger, with rounded 
slightly concave tip. Pygidium with two lat- 


504 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


0.2 mM 


Fig. 1. Dorsal view of Exogone breviantennata paratype (P-14590, ZMH) from El Salvador, 1.95 mm long, 


0.2 mm wide, and with 24 setigers. 


eral cirri (0.05 to 0.15 mm long), some- 
times uneven in length, apparently due to 
regeneration. 

Pharynx (Figs. 3B, 4, 5) short, extends 
to setigers 1, 2, or 3 depending on degree 


of contraction; anterior rim with 10 soft pa- 
pillae in addition to the large, curved, point- 
ed, dorsally attached tooth, more easily 
seen when pharynx is everted. Proventric- 
ulus (0.07 to 0.2 mm long and 0.05 to 0.12 


VOLUME 113, NUMBER 2 505 


mS a eae 
ia 
a 
eee 


Fig. 2. Setae from Exogone breviantennata paratype (P-14590, ZMH) from El Salvador, 1.95 mm long, 0.2 
mm wide, and with 24 setigers. A. Distal portion of dorsal simple seta, 3 4m basal width; B. Heterogomph 
spiniger, 30 wm blade length; C. Heterogomph falciger, 10 zm blade length; D. Ventral simple seta, 3 4m basal 
width. 


506 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Table 1.—Length in ym of spinigers and blades of compound setae from right side of Exogone breviantennata 
paratype (P-14590, ZMH) from El Salvador, 1.95 mm long, 0.2 mm wide and with 24 setigers. utm = unable 


to measure; avg = average; 0 = not present. 


Setiger Heterogomph Heterogomph 


number spiniger falciger#1 
l 24 7. 
2, 24 dL 
3 30 Wf 
4 30 10 
») 30 10 
6 32 utm 
7-14 utm utm 
15 24 7 
16—20 utm utm 
2A 20 5 
22-24 utm utm 
Averages 26415 TASS 


mm wide) extends posteriorly from setigers 
1—4, 2—4, 2-5, 2—6, 3-5 or 3—6 (Figs. 1, 5) 
depending on degree of contraction during 
fixation; about 15 rows of muscle cells. 

Sexually mature females with internal 
eggs or ventrally attached eggs or larvae 
(Fig. 5). Six internal eggs, average diameter 
0.08 mm in setigers 12—17 of a 25-setiger 
female; five eggs 0.09 to 0.096 mm in di- 
ameter, attached ventrally, one to two per 
setiger on setigers 12, 14, and 15, of a 25- 
setiger female; and six eggs 0.08 to 0.1 mm 
in diameter, attached ventrally, two per se- 
tiger on setigers 15, 19, and 22 of a 31- 
setiger female. Five, 6-setiger larvae, (Fig. 
5) average length and width 0.43 mm and 
0.9 mm, one or two on setigers 12, 13, and 
15 of a 21-setiger female; eight, 5-setiger 
larvae, average length and width 0.26 mm 
and 0.11 mm, one each on setigers 11-14 
and 16—19 of a 27-setiger female. One 26- 
setiger male found with sperm packed in 
body cavity of setigers 18—25. No repro- 
ductive modifications observed for either 
sex. The only morphological differences 
between juveniles and sexually mature 
specimens are fewer setigers and more ven- 
tral simple setae in the younger stages (Ta- 
blew): 

Discussion.—Two species of Exogone 
that have a dorsal cirrus on the second se- 
tiger and a short median antenna have been 


Heterogomph Heterogomph Heterogomph 


falciger#2 flaciger#3 falciger#4 

Tl 6 5 

Ti 6 5 

a) 6 5 

6 6 6 

6 6 6 
utm utm utm 
utm utm 0 

7 5 0 
utm utm 0 

5 0) 0) 
utm utm 0) 

6:35 5.80 55 


reported from the Caribbean: E. brevianten- 
nata and E. exmouthensis. A third species, 
E. occidentalis, was not compared to E. 
breviantennata when it was described from 
the Galapagos Islands (Westheide 1974). 
When Russell (1991) emended Westheide’s 
(1974) description of E. occidentalis, based 
on a study of the type material and on spec- 
imens from a Belizean barrier reef, he de- 
scribed most setae as bearing spination as 
well as denticulation, thus bringing its de- 
scription closer to E. breviantennata. Our 
examination of the Belize specimens leads 
us to agree with San Martin (1991) who 
listed E. occidentalis as a junior synonym 
of E. breviantennata. 

We reexamined a paratype of E. brevian- 
tennata and found that the setae showed the 
spination and denticulation (also present in 
Bahamian specimens), which were referred 
to as indistinct in the type description. San 
Martin (1991) figured the denticulation of 
the blades of the compound setae and spi- 
nation on the shafts on most setae for E. 
breviantennata from Cuba. 

Exogone exmouthensis was described 
from Australia (Hartmann-Schroder 1980) 
without comparison to E. breviantennata 
and reported from Cuba by San Martin 
(1991) without description. He placed E. 
breviantennata and E. exmouthensis in sep- 
arate subgenera based on details of the se- 


VOLUME 113, NUMBER 2 507 


Table 2.—Morphological measurements of specimens of Exogone breviantennata from Osprey Lake and 
Reckley Hill Pond, San Salvador, Bahamas. Measurements in mm. Spec. = specimen; set. = setiger; provent. 
= proventriculus; vent.SS = setiger number on which ventral simple setae begin; coll. = collected; loc. = 
collection site; Loc. 1 = Osprey Lake; Loc. 2 = Reckley Hill Pond; utm = unable to measure. Figures for 
specimens numbered 33-37 are averages from 5 larvae attached to specimen #4 and those for specimens 38— 
45 are averages from 8 larvae attached to specimen #15. 


Spec. Set. Body Body Provent. Provent. Vent. Date 
# # length width length width SS coll. Loc. Comment 
38-45 5 0.30 1.00 0.07 0.05 05/05/96 2 on spec. 15 
33-37 6 0.43 0.09 0.07 0.05 ] 05/05/96 l on spec. 4 
16 6 0.34 0.10 utm utm 1 06/19/97 Z 
7 10 0.52 0.10 0.08 0.07 06/19/97 Z 
18 10 0.70 0.10 0.08 0.05 ] 06/19/97 Z 
19 12 0.63 0.12 utm utm 06/19/97 Z 
20 12 0.71 0.12 0.10 0.06 8 06/19/97 Z 
5 ie 0.85 0.17 0.12 0.09 4 05/05/96 l 
21 13 0.70 0.11 Onz 0.07 1 06/19/97 pe 
5 15 1.00 0.12 0.14 0.11 10 05/05/96 l 
PPD 16 0.88 0.14 O13 0.07 10 06/19/97 2 
1 16 1.20 On2 OS 0.1 9 05/05/96 ] 
23 17 1.03 0.14 0.11 0.07 i 06/19/97 2 
24 17 1.10 0.14 utm utm 10 06/19/97 2 
2 A] 1.35 0.16 0.14 0.1 9 05/05/96 1 
25 19 Lt 0.14 0.15 0.08 12 06/19/97 Ps with eggs 
7 19 1.40 0.18 0.15 0.09 IZ 05/05/96 
26 20 22 0.18 O17 0.10 utm 06/19/97 2 
24 20 1.30 0.16 0.14 0.08 13 06/19/97 2 
4 21 1.40 0.2 C5 Ont 22 05/05/96 l with larvae 
12 2A 2.10 utm OA7 0.1 14 05/05/96 2 
28 11 2.10 0.17 O11F 0.1 i) 06/19/97 2 
9 22 13/5 utm 0.2 0.1 16 05/05/96 l 
29 23 1.80 0.16 0.16 0.10 17 06/19/97 ou 
13 pee) Dine utm 0.17 0.1 14 05/05/96 2 
30 25 1.72 0.18 0.18 0.10 19 06/19/97 2 
6 jib 130 utm 0.18 Or 18 05/05/96 l with eggs 
51 25 2.00 0.16 utm utm 20 06/19/97 2 
32 26 1.90 0.19 0.20 0.10 20 06/19/97 2 with sperm 
8 pa | 1/95 0.20 O2 0.12 23 05/05/96 l 
14 2d pIp 2.) 22 0.19 O12 23 05/05/96 2 
15 pH Z.10 0.20 OF 0.1 22 05/05/96 2 with larvae 
11 a1 ZS 0.20 0.18 0.1 18 05/05/96 ] with eggs 


tae, e.g., head of the shaft of compound se- 
tae with or without spination, the determi- 
nation of which is dependent upon variables 
unassociated with morphology of the spec- 
imens themselves (see above). We have not 
seen specimens of E. exmouthensis and so 
are unable to comment on its relationship 
to E. breviantennata. 

Two other species, E. ovalis Hartmann- 
Schroder, 1960, and E. breviantennata 
ovalis Hartmann-Schroder, 1974, have been 


made synonyms of E. breviantennata (San 
Martin 1991). 

Most species of the genus Exogone that 
are described as having a dorsal cirrus on 
setiger two also have a median antenna that 
reaches at least to the tip of the prostomi- 
um, if not longer. In this regard, E. brev- 
iantennata is unique in that its median an- 
tenna is much shorter, not even reaching to 
the tip of the prostomium. 

Because new species are sometimes 


508 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


“VIZ 


Fig. 3. Anterior end of Exogone breviantennata from Reckley Hill Pond, San Salvador, Bahamas. 1.72 mm 
long, 0.18 mm wide, and with 25 setigers. A. Dorsal view of prostomium including first two setigers; B. Ventral 
view including first setiger, showing anterior end of pharynx with tooth. 


VOLUME 113, NUMBER 2 


509 


0.2 mm 


Fig. 4. Exogone breviantennata from Osprey Lake, San Salvador, Bahamas. 1.4 mm long, 0.18 mm wide, 
and with 19 setigers; lateral view of anterior end, showing tooth in extruded pharynx. 


erected for minor setal differences, we care- 
fully measured as many characters as pos- 
sible for the paratype (from El Salvador) 
and for specimens from Osprey Lake and 
Reckley Hill Pond on San Salvador. We 
found no significant differences from site to 
site. 

Habitat.—Type material collected in mud 
around mangroves in El Salvador (Hart- 
mann-Schroder 1959). In the Bahamas, E. 
breviantennata was found amongst filaments 
of the algae Batophora sp. and Vaucheria 
sp. and on the prop roots of Rhizophora 
mangle. Pascual et al. (1996) summarized its 
habitat as intertidal to shallow depths, sand 
and shell gravel, in sponges and amongst hy- 
droids on R. mangle roots, Halimeda sp. in 
beds of Thalassia testudinum, Lobophora 


variegata and calcareous crusts, Vermetus 
sp., photophilic algae and living inside the 
cavities of six species of infralittoral demo- 
sponges, where it constituted 0.6% of the to- 
tal number of polychaetes and 9.7% of Ex- 
ogone specimens extracted from the spong- 
es. These habitats define it as part of an in- 
terstitial and crevice fauna. 
Distribution.—Equatorial Pacific: Gala- 
pagos Islands (Westheide 1974), North East 
Pacific: La Herradura, El Salvador (Hart- 
mann-Schréder 1959); North Atlantic: Ba- 
hamas (reported here), Belize (Russell 
1991, as E. occidentalis), Canary and Ma- 
deira Islands (Pascual et al. 1996), and 
Cuba (San Martin 1991). San Martin (1991: 
731) says “‘Probably circumtropical.”’ 
Biology.—The lack of solid material in 


S10 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


0.2mm 


Fig. 5. Ventral view of Exogone breviantennata female with attached larvae from Osprey Lake, San Salvador, 
Bahamas, 1.4 mm long, 0.2 mm wide, and with 21 setigers. 


VOLUME 113, NUMBER 2 


the digestive tract of the Bahamian speci- 
mens suggests that they were fluid feeding, 
most likely using their tooth to penetrate 
algal filaments and then sucking out the 
contents with the aid of the muscular pro- 
ventriculus. See Figs. 3B and 4 for the tooth 
in position for feeding. In live material, the 
gut is often dark green along its entire 
length, thus supporting this interpretation. 
Pascual et al. (1996) reported that, in the 
Canary and Madeira Islands, spicules of the 
species of sponges from which specimens 
were collected were found in their gut, sug- 
gesting that the worms were actually eating 
sponge tissue. There did not seem to be any 
preterence for a particular species of 
sponge. They speculated as to whether 
polychaetes that are found with sponges 
function at times as parasites, mutualists, or 
commensals. Members of the subfamily 
Exogoninae had been previously thought to 
be selective deposit-feeders of mud or de- 
tritus (Uebelacker 1984:30—4). 

Reproduction.—San Salvadoran sexually 
mature females were found with up to six 
eggs or eight larvae (Fig. 5, Table 2) at- 
tached to their ventral surface. Extrusion of 
eggs from the body cavity and fertilization 
were not observed. One male was found 
with sperm. There was no evidence of re- 
productive modifications of any kind. Hart- 
mann-Schréder (1959) found gametes in, 
and eggs attached ventrally on, setigers 11— 
15 in the type material from El Salvador. In 
Belize, Russell (1991) found a sexually ma- 
ture female with oocytes in six setigers be- 
ginning about setiger 16. Since attached lar- 
vae from both San Salvador and Belize fe- 
males (Russell 1991) had no more than six 
setigers, they probably detach and become 
benthic at this stage. 

Previous Bahamian records of the genus 
Exogone.—Worldwide, nearly 50 species 
of the genus Exogone have been described 
(Pascual et al. 1996). As of 1992, 19 of 
these species that are considered valid have 
been reported from the Caribbean and as- 
sociated waters: E. arenosa Perkins, 1981; 
E. atlantica Perkins, 1981; E. brevianten- 


511 


nata Hartmann-Schroder, 1959; E. cariben- 
sis San Martin, 1991; E. dispar Webster, 
1879; E. exmouthensis Hartmann-Schroder, 
1980; E. gemmifera Pagenstecher, 1862; E. 
longispinulata San Martin, 1991; E. lourei 
Berkeley & Berkeley, 1938; E. naidina 
@rsted, 1845: E. naidinoides Westheide, 
1974; E. parhomoseta mediterranea San 
Martin, 1984; E. pseudolourei San Martin, 
1991; Exogone sp. A and sp. B, Uebelacker, 
1984; E. rolani San Martin, 1991; sp. A, 
San Martin, 1991; E. verugera Claparéde, 
1868; and E. wolfi San Martin, 1991. See 
Perkins & Savage (1975), Salazar-Vallejo 
(1992) and Camp et al. (1998) for referenc- 
es to Caribbean reports. Of these, two have 
been reported from the Bahamas: Exogone 
dispar (Webster) (based on Vittor & John- 
son 1977 and our examination of USNM 
51541, from Bimini Lagoon) and E. verug- 
era (Claparéde) (in the collections of the 
USNM, L. Ward, pers. comm., 29 Aug 
1995). San Martin (1991) notes that the 
original description of E. verugera is con- 
fused, and therefore not reliable, and syn- 
onymizes a series of Atlantic reports with 
E. breviantennata. Seven of the species re- 
ported from the Caribbean have a dorsal 
cirrus on the second setiger: E. arenosa, E. 
breviantennata, E. dispar, E. exmouthensis, 
E. lourei, E. pseudolourei, E. rolani, and E. 
wolfi. As any of the remaining six might be 
found in Bahamian waters, they are includ- 
ed in the key below. 


Key to Caribbean Exogone species with 
dorsal cirrus on second setiger 


la. Median antenna does not reach tip of 
prostomiumnisegas sok. cians i~.ce'u.. 
.. E. breviantennata Hartmann-Schroéder, 1959 
1b. Median antenna, much longer than lat- 
eral antennae, reaches to or exceeds tip 


C50 RUSS COREA ORIN Goss no a pitt ns acer sw oc ee 2 

2a. Tips of shafts of spinigers on setiger 2 

with greatly enlarged triangular process 
Pree hee eee oes eer ee eee i: 

2b. Tips of shafts on setiger 2 not greatly 
eilareed ax . seu 2). cs eee . +4 


3a. Superior simple setae with well-defined 


S12 


spine on tips; proventriculus with up to 
28 rows. of muscle cells. ssc, 4:6 
E. arenosa Perkins, 1981 
3b. Superior simple setae without spine; 
proventriculus with at most 20 rows of 
muscle cells: |. Were een pe eee ee 
.... E. lourei Berkeley & Berkeley, 1938 
4a. Dorsal simple setae of median and pos- 
terior setigers enlarged and modified, 
strongly bidentate; proventriculus short, 
with ca 15 muscle:cells rows ...... 
E. pseudolourei San Martin, 1991 
4b. Dorsal simple setae of median and pos- 
terior setigers not enlarged 
5a. Median antenna club shaped ........ 
ie ste Se: E. dispar (Webster, 1879) 


7 2e © © © © © © © ew ee 


bins eh er eye 


Sb. Median antenna elongate 2. ..5 s..o5< . 6 
6a. Dorsal simple setae similar throughout 
Ody chases. E. wolfi San Martin, 1991 


6b. Dorsal simple setae increasing in thick- 
ness posteriorly and changing in shape 
i. 4 Te E. rolani San Martin, 1991 


Acknowledgments 


We were alerted to the interesting poly- 
chaete fauna of the land-locked marine 
lakes of San Salvador, The Bahamas, by 
William Lindsay (Elmira College, New 
York), who, along with the staff of the Ba- 
hamian Field Station, have made this work 
possible. We thank them and Fitchburg 
State College for its continued support. We 
especially appreciate the constructive com- 
ments on the manuscript by Drs. Nathan 
Riser and David Russell. 


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. 1960. Polychaeten aus dem Roten Meer.— 

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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 
113(2):514—525. 2000. 


Podarke aberrans Webster & Benedict, 1887 - resolution, with 
descriptions of two new species in the genus Microphthalmus 
(Annelida: Polychaeta) 


Nathan W. Riser 


Marine Science Center, Northeastern University, Nahant, Massachusetts 01908, U.S.A. 


Abstract.—Reference is made to individuals with ‘“‘fan-shaped”’ caudal ap- 
pendages from two different regions in the Gulf of Maine and to two different 
life-history stages in the original description of Microphthalmus aberrans 
(=Podarke aberrans Webster & Benedict, 1887). Intertidal meiofaunal inves- 
tigations in the region over the past forty years have yielded three species that 
correspond to parts of the description. Two fragments of the ‘“‘adult form” 
labeled Type, from Eastport, Maine, upon which most of the type description 
is based, exist as one of the syntype slides deposited by Webster & Benedict 
in the National Museum of Natural History. The species has been encountered 
occasionally in clean coarse sand beaches along the coast of Maine, and in the 
subtidal of Nahant Bay, Massachusetts. A second species with a fimbriate anal 
lamella, M. aggregatus n.sp., occurs in the intertidal of Cape Cod Bay, Mas- 
sachusetts. The “‘half grown specimens”’, M. pettiboneae n.sp., to which the 
epithet aberrans has been incorrectly applied, is also present among the syntype 
slides, and has been collected in intertidal samples from New Brunswick, Can- 


ada to the south shore of Cape Cod, Massachusetts. 


Webster & Benedict (1887) assumed that 
badly damaged specimens from Province- 
town, Massachusetts collected in 1879, but 
not retained, were the same species as the 
one being described as Podarke aberrans 
from the Eastport area in Maine, because of 
the “peculiar fan-shaped’? anal lamella. 
One “‘adult’’ specimen was encountered by 
the authors at Eastport, and the description 
of Microphthalmus abberans is based upon 
that individual with additional remarks on 
the morphology of “‘half-grown speci- 
mens.” The figures accompanying the de- 
scription are of both the ‘adult’? and ju- 
venile form, and the type material of Mi- 
crophthalmus aberrans (Webster & Bene- 
dict, 1887) (=Podarke aberrans W. & B.) 
deposited in the National Museum of Nat- 
ural History (USNM), is a mixture of two 
species. Southern (1914) reported that he 
had examined the “original types”? depos- 
ited in the USNM and at Union College and 


that they did not agree with the type de- 
scription. He transferred the species from 
Podarke to Microphthalmus, accepting the 
specific epithet aberrans for the material 
available to him and furnished a figure of 
the notopodial setae to support his conten- 
tion; thus, we must assume that the slide of 
the specimen upon which Webster & Ben- 
edict based their description was not en- 
countered. Eliason (1920) accepted South- 
ern’s statements about the type material 
while describing material from the Ore- 
sund, which he ascribed to the Webster & 
Benedict species, thus establishing a pre- 
cedent of applying the epithet aberrans to 
a species that does not agree with the type 
description. The failure of subsequent au- 
thors to examine the type material has re- 
sulted in a perpetuation of this error. 

A few individuals fitting the description 
of Microphthalmus abberans have occurred 
in intertidal meiofaunal collections I have 


VOLUME 113, NUMBER 2 


made along the coast of Maine over the past 
forty years and in subtidal substrate from 
Nahant Bay, Massachusetts. Numerous 
specimens of a second species with a fim- 
briate anal lamella have routinely been en- 
countered in the intertidal dunes on the sand 
flat at Ellisville, Massachusetts, as well as 
on the beach at Manomet, Massachusetts 
across Cape Cod Bay from Provincetown. 
The initial description of Podarke aberrans 
refers to three different species belonging 
to the genus Microphthalmus, two of 
which, including the species that has born 
that name since Southern’s publication, 
must be described as new. 


Materials and Methods 


Specimens were obtained from substrate 
collected in 18 fl. oz. plastic bags for mei- 
ofaunal studies. Each sample was washed 
with fresh sea water in the laboratory and 
decanted onto 153 wm screens from which 
animals were removed for sorting. The sed- 
iment was then extracted with 7.5% MgCl, 
and decanted onto the screens from which 
the animals were washed into fresh sea wa- 
ter. Specimens were anaesthetized with 
7.5% MgCl, prior to fixation in Hollande’s 
cupri-picri-formal-acetic. Whole mounts 
were Stained with Ranvier’s picro-carmine 
or Mayer’s alcoholic HCl carmine; some 
were counterstained with alcoholic indigo- 
carmine. All measurements were obtained 
from living specimens. 

Material labeled Microphthalmus aber- 
rans in the collections of the Atlantic Ref- 
erence Centre (ARC), St. Andrews, New 
Brunswick, and the National Museum of 
Natural History were obtained on loan. The 
syntypes (USNM 447) of Podarke aberrans 
from Eastport, Maine deposited by Webster 
& Benedict consist of five slides and two 
specimens in alcohol. 


Microphthalmus aberrans (Webster & 
Benedict, 1887) 
Figs. 1-6 
Podarke aberrans Webster & Benedict, 
1887:713-—715, p.p., “‘adult form’’, figs. 14, 


515 


15, 18. Microphthalmus aberrans.—Petti- 
bone, 1963:104, p.p. 

Diagnosis.—Sexual individuals to 9 mm 
in length with 30—45 setigers. Ocelli ab- 
sent. Dorsal cirri more than four times as 
long as neuropodial lobe. Anal cirri slightly 
longer than dorsal cirri. Notopodia with up 
to 15 pointed simple and one pectinate seta 
and a sturdy aciculum. Presetal neuropodial 
ligule with thin aciculum associated with 
one or two bidentate simple setae (Fig. 3). 
Neuropodial setal bundle with thick acicu- 
lum, one apically bidentate simple seta and 
about 15 finely denticulated apically biden- 
tate falcigers. Falciger blades range from 6— 
32 wm in length. Testes in setigers 9 
through 16, ovaries in following setigers. 
Penis anterior to parapodia of setiger 3 on 
both sides, with tubular stylus 25 wm long, 
muscle bulb large with large posterior pa- 
pilla. Anal lamella with about 30 short, 
blunt, fimbriae. 

Lectotype.—USNM (447), slide 502, la- 
beled “‘Podarke aberrans adult fragments” 
(two fragments; one of three setigers from 
the female region of the body, and one of 
four setigers and anal lamella). 

Material examined.—Data and _ photo- 
graphs obtained from living specimens col- 
lected in the intertidal at Liberty Point, 
Robbinston, Maine; Crowe Neck, Cobs- 
cook Bay, Maine; Griffith’s Head, George- 
town, Maine, and 16 m depth off Egg Rock, 
Nahant Bay, Massachusetts, in addition to 
the lectotype slide, have made it possible to 
complete the description of the species. One 
adult (USNM 186552) collected on 27 Mar 
1980 at Liberty Point and one immature 
specimen (USNM 186533) encountered at 
Griffith’s Head on 20 Apr 1999, have been 
deposited, in alcohol, in the USNM. 

Morphological notes.—The acicula taper 
toward the slightly expanded apex; that of 
the presetal neuropodial lobe is hooded. 
The diameter of the presetal neuropodial 
aciculum is almost one-half that of the other 
two, <3 vs. 6 wm, however it runs at an 
angle, and its base varies in position from 
the same level to 4 wm distal to that of the 


516 


neuropodial aciculum. The acicula are all 
approximately the same length, 76 wm. The 
pectinate notopodial seta ranges from 91-— 
99 wm in length with the blade occupying 
almost one-half of the length. The blade is 
finely denticulated almost to the tip (Fig. 6). 
The difference in construction of the blade 
and shaft allows for great flexibility at the 
juncture so that at times the seta takes on 
the appearance of being jointed; the twisted 
appearance figured for the pectinate seta of 
M. bermudensis Westheide, 1973, (Abb. 4, 
fig. D) has also been encountered. The pec- 
tinate setae of species with several simple 
notopodial setae can rarely be recognized in 
the clump (Fig. 2) but can usually be dis- 
tinguished in some of the parapodia of liv- 
ing specimens under coverslip pressure. 
The simple notopodial setae vary in ap- 
pearance, depending upon the angle at 
which they are viewed and if they are re- 
tracted into the body or are projected. The 
majority of the simple notosetae of pre- 
served specimens are strongly hooked api- 
cally. The striated fibrillae forming the cor- 
tex spiral (180° ?) but can not be traced to 
the apex. This spiraling may account for the 
flexibility which results in the hooked con- 
figuration. 

There appear to be eight digitiform pa- 
pillae on the pharynx cap. The pharynx ex- 
tends posteriorly to setiger 5 which is oc- 
cupied by the ventriculus. The epithelium 
of the pharynx is thick and when the organ 
is inflated, develops longitudinal ridges 
(Fig. 1). There is a well-developed valve 
and sphincter separating the pharynx and 
ventriculus. The latter is globular and com- 
prised of radial muscle and glandular cells. 
Retractor muscles extend from the base of 
the pharyngeal cap to the body wall at the 
juncture of the third tentacular segment and 
first setiger at which point protractors orig- 
inate passing to insert at the juncture of the 
pharynx and ventriculus. Strongly devel- 
oped retractor muscles extend from the ven- 
triculus to the longitudinal muscles of se- 
tiger 8. 

One of the 18 sexually mature specimens 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


had an ovary on the left side, and testis on 
the right of setiger 16. 

Remarks.—tThe figure of the anal lamella 
in the description by Webster & Benedict 
has little relationship to the specimen on the 
lectotype slide, except that the midregion is 
different from the rest of the margin. The 
lamella of living animals (Fig. 5) is domed, 
and if not fully expanded projects over the 
fimbriae so that they may appear to be ab- 
sent medially for a variable extent of the 
margin (Fig. 4). A fold in the middle of the 
dorsal surface of the anal lamella on the 
lectotype may have been interpreted by 
Webster & Benedict as corresponding to the 
anal lamella of the “half grown’’ speci- 
mens. 

Pettibone (1963) described the general 
appearance of the notopodial simple setae 
on the adult slide of the type specimen of 
M. abberans but did not report observations 
on notopodial setation of other specimens 
which she identified as that species (some 
of which were mature). The simple noto- 
setae of preserved specimens of M. fragilis 
Bobretzky, 1870 are thin and sharply point- 
ed; some are slightly bent toward the apex; 
those of M. urofimbriatus Alikunhi, 1948 
were reported to be straight. [Hartmann- 
Schr6der (1960) stated that the setae of M. 
c.f. urofimbritta (sic) from the Red Sea 
were too delicate to allow for characteriza- 
tion. The clusters of notopodial simple setae 
present in M. aberrans (Fig. 1), M. fragilis, 
and M. urofimbriatus are so apparent that it 
is more likely that her worms belong to the 
M. similis group of species.] The apex of 
the pectinate seta of M. fragilis is long and 
slender beyond the denticulated portion of 
the blade. 

The species has not been routinely en- 
countered nor in significant numbers. The 
single specimen obtained from the subtidal 
of Nahant Bay was associated with Ophry- 
otrocha cf. gracilis Huth 1934, Parougia 
caeca (Webster & Benedict, 1884) and an 
eyeless species of Protodorvillea, all sub- 
tidal species in Nahant Bay but routinely 


VOLUME 113, NUMBER 2 517 


j 
Bee ae ie tr had 
-# yO te 3 
Pe Lea eee oes 


a 
Sa. tat 


ate ey 


Figs 1-10. 1—6; Microphthalmus aberrans, living specimens. |. Optical section of foregut region. 2. Noto- 
podial setal bundle in situ. 3. Paired simple neuropodial seta in situ. 4. Anal lamella, anal cirri detached, dorsal 
view. 5. Anal lamella second specimen, anal cirri detached, ventral view. 6. Blade of pectinate seta, dissociated 
preparation. 7-9; M. aggregatus, 7. Optical section of foregut region. Living specimen. 8. Anal lamella and anal 
cirri, living specimen. 9. Penis stylet and papilla, living specimen. 10; M. pettiboneae. 19 «wm falciger blades, 
slightly oblique. Fixed specimen. 


518 


present in the intertidal of the coast of 
Maine. 


Microphthalmus aggregatus, new species 
Figs. 7-9, 11-15 


?Podarke aberrans Webster & Benedict, 
1887:713 p.p., ““same form at Province- 
town, Massachusetts.”’ 

?Microphthalmus c.f. similis Westheide & 
Rieger, 1978; Westheide, 1979. 


Diagnosis.—Transparent except for 
opaque white gonads. Individuals with go- 
nads 3.2—7.1 mm long; with 31—43 setigers. 
Paired and median prostomial antennae ap- 
proximately same length; palps extend 
about same distance beyond prostomium as 
paired tentacles. Blackish brown ocellus 
about mid-way between bases of paired 
prostomial antennae and Ist tentacular cir- 
rus on each side. Dorsal cirri about four 
times length of neuropodium. Notopodium 
of first setiger with aciculum, following se- 
tigers with notopodial aciculum and pecti- 
nate seta. Neuropodium with bidentate fal- 
cigers and one or more simple setae. Phar- 
ynx bipartite; ventriculus oval to round, oc- 
cupying 3rd setiger. Testes bilobed, in 
setigers 9 through 16; ovaries in subsequent 
setigers to penultimate. Penis digitiform 
with eight stylets and dorsal sensory papilla 
(Fig. 9); anterior to parapodia of third se- 
tiger on each side. Anal lamella with digi- 
tiform fimbriae. 

Holotype.-—USNM_ 186534. Massachu- 
setts; Ellisville, intertidal coarse sand. 

Paratypes.—USNM_ 186535, 12 speci- 
mens in alcohol. Peabody Museum of Nat- 
ural History, Yale University, New Haven, 
Connecticut: YPM 24000, YPM 24001, 
stained whole mounts. 

Etymology.—Latin aggregatus, clus- 
tered; referring to the tendency to occur in 
large concentrations, and to aggregate when 
isolated in vitro. 

Material examined.—More than two 
hundred specimens from Ellisville, Massa- 
chusetts have been collected. Morphologi- 
cal data were obtained from 84. Additional 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


data were recorded from specimens collect- 
ed on Manomet Beach, Massachusetts 
(about 1 km North of Ellisville). 
Morphological notes.—The body tapers 
from the anterior end to the female region, 
but the interparapodial width is about con- 
stant. Bands of refringent granules in the 
dorsal epidermis are colorless and do not 
produce opacity. Pigment is restricted to 
black granules in the epithelial cells of the 
rectum. The setigers containing ovaries are 
inflated and have a larger diameter than the 
anterior setigers. The prostomial antennae 
and palps range from 0.13—0.15 mm in 
length; the median antenna is thinner and is 
occasionally forked. The palpophore is 
small, accounting for very little of the ex- 
tent of the palp. The dorsal tentacular cirri 
are longer than the ventral, viz., lst 0.20— 
0.22:0.13—0.16, 2nd 0.27:0.14-0.16, 3rd 
0.27—0.30:0.05—0.07 mm. Dorsal cirri of 
parapodia 0.24—0.27 mm long, except setig- 
er 1 which are shorter, rarely exceeding 
0.17 mm. Frequently, apparently as the re- 
sult of regeneration, individual tentacular 
and dorsal cirri are shorter than these mea- 
surements. The pleura of all setigers expand 
laterally and the cirrophores are elongate, 
capped distally with a lobe above the base 
of the cirrus (Fig. 11). This lobe contains 
numerous mucous cells. The pair of anal 
cirri to either side of the anus attain lengths 
up to 0.4 mm. (Fig. 8). The fully expanded 
anal lamella is about 0.25 mm broad with 
as many as 42 digitate fimbriae 0.03 mm 
long. Shorter fimbriae as well as those that 
are forked probably indicate injury. 
Ciliation on the prostomium is restricted 
to the nuchal organs (Fig. 11), each of 
which bears a small papilla on the anterior 
margin. A band of cilia extends across the 
dorsum between the bases of the second 
tentacular cirri but is interrupted medially 
on the subsequent four segments. Complete 
ciliary bands extend from the median edge 
of the lobe at the apex of the cirrophore 
(Fig. 11) across the dorsum of succeeding 
setigers. There is a lateral cluster of cilia 
anterior to each parapodium, and a lateral 


VOLUME 113, NUMBER 2 319 


* 


Figs. 11-14. Microphthalmus aggregatus, SEM. 11. Dorsal view, anterior end. 12. Ventral view, anterior 
end. 13. Anal lamella, ventral view. 14. Medium and short falciger blades. 


520 


patch posteriad. Ventrally, a band of cilia 
extends along the posterior edge of the 
mouth (Fig. 12), constituting the only con- 
sistent ciliation on the ventral surface. 

Cirrophores of the dorsal cirri are almost 
as long as the neuropodia. Notopodial lobes 
are not present. Notopodial setation consists 
of one or two pointed acicula 45-57 pm 
long and one or two pectinate setae 24—33 
wm long of which the comb occupies one- 
fourth to one-third of the length. The pre- 
setal lobes of the neuropodia are conical, 
about 22 ym long and are supported by a 
thin pointed aciculum 57 wm long accom- 
panied by a bidentate simple seta which 
moves independently, or sometimes in con- 
junction with the other neuropodial setae. 
A pointed simple or bidentate seta, or both, 
may be present in the neurosetal bundle, 
which consists of as many as nine apically 
bidentate falcigers (Fig. 14), most of which 
are 17—20 pm long and accompanied by 
one or more with short blades 7-8 wm long 
and usually one or more with blades in ex- 
cess of 36 wm long. Ventral cirri of the 
parapodia are thin and filiform ranging be- 
tween 35 and 42 um in length. 

The soft portion (cap) of the pharynx oc- 
cupies the first tentacular segment and bears 
eight simple digitiform papillae tightly 
packed together around a large and expan- 
sive surface. Retractor muscles pass directly 
from the base of the cap to the body wall 
of the third tentacular segment from 
whence, protractors extend to the base of 
the pharynx. The second region occupies 
the second tentacular segment and is mus- 
cular (Fig. 7) with an epithelium containing 
granular products. The third region extends 
from the third tentacular segment to the 
third setiger. The ventriculus is firmly at- 
tached to the adjacent wall of the body. 

Remarks.—In April and May, the ovaries 
are packed with yolky oocytes 35 wm in 
diameter, and the testes are primarily in ear- 
ly stages of spermiogenesis with few sper- 
matids undergoing flagellar development. 
By August, the testes are packed with fully 
developed sperm; ovarian segments contain 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


sperm in seminal receptacles; and the num- 
ber of oocytes is reduced. Ovaries contain 
two or three degenerating oocytes in No- 
vember and December, but the testes are 
packed with sperm. The animals have lived 
in the sorting bowls for up to eight months 
with biweekly water changes, but egg lay- 
ing has not occurred in the absence of a 
food source, and the gonads have been re- 
sorbed. Efforts to investigate posterior re- 
generation, as the result of the discovery of 
a specimen with a bifurcated posterior end 
(Fig. 15), in 1977, have been futile for the 
same reason. 

The major difference between M. aggre- 
gatus and M. similis Bobretzky, 1870, in 
addition to size and number of segments, is 
the distribution of the gonads. Ovaries of 
M. similis do not occur anterior to setiger 
20. About 10% of the M. aggregatus indi- 
viduals (7 of 92) had the ovary on the right 
side of setiger 16 and testis on the left, and 
the first testis on the right in setiger 8 in- 
stead of 9. One specimen was encountered 
with a mixed gonad in the right antimere of 
setiger 17 and ovary on the left. Some spec- 
imens of M. aggregatus have lacked pig- 
mented ocelli, while others have lacked 
them on one side, or have had two (one 
dorsal and one ventral) on one or both 
sides. Variation, as well as absence of ocelli 
in M. fragilis, was noted as an intraspecific 
variation by Bobretzky (1870). 

The divided pharynx of M. fragilis fig- 
ured by La Greca (1950:fig. 1) was not ap- 
parent in any of the three specimens from 
the Black Sea which I examined, but the 
pharynx of M. similis from there was all 
divided as in M. aggregatus. 


Microphthalmus pettiboneae, new species 
Fig. 10 


Podarke aberrans Webster & Benedict, 
1887: 713-715, p.p. ‘“‘juvenile form’’, 
fies: 16517, 19; 20: 

Microphthalmus aberrans Southern, 1914: 
46, fig. 7._Pettibone, 1963:104, p.p., figs. 
27a—b.—Clausen, 1986:184—186, figs. 


VOLUME 113, NUMBER 2 


pAC5 


16 


Figs. 15,16. 15. Microphthalmus aggregatus, with posterior bud, living specimen. Scale equals 0.09 mm. 16. 
Anal lamella of living Microphthalmus nahantensis Westheide & Rieger, 1987. Scale equals 0.045 mm. 


14, 19 (for additional synonymy see 
Hartmann-Schroder 1971). 


Diagnosis.—Variable intensity of brown 
pigmentation; mature individuals less than 
10 mm in length with 30-35 setigers; pair 
of ocelli present; dorsal cirri only slightly 
longer than neuropodial lobes; unpaired 
sucker-like penis; single pectinate and sim- 
ple seta with an aciculum in notopodium; 
testes in setigers 6 through 9; bilobed (me- 
dially indented) anal lamella. 

Syntypes.—USNM (447), slides 137, 
138, 501, and two in alcohol deposited by 
Webster & Benedict. 

Additional material deposited.—USNM 
(186536) from Beacon Point, Blacks Har- 
bour, New Brunswick, Canada: ARC 
9953141; 995314, from Pagan Point, St. 
Andrews, New Brunswick, Canada. 

Etymology.—The species is named for 
Dr. Marian Pettibone, who in 1953, was re- 
sponsible for my initial efforts to resolve 
the problem. 

Material examined.—Observations have 
been recorded from numerous living indi- 
viduals collected in intertidal coarse sand 


beaches between Beacon Point, Blacks Har- 
bour, New Brunswick, Canada and East 
Point, Nahant, Massachusetts, USA. Spec- 
imens identified by Pettibone as M. aber- 
rans (USNM 28413) from Southport Island, 
Maine; (USNM 32498) from Rye Beach, 
New Hampshire, and from Halifax, Nova 
Scotia, Canada (USNM 49180) were ex- 
amined and found to be M. pettiboneae, as 
were specimens from the subtidal (ARC 
9661230) of Lime Kiln Bay, Letang Estu- 
ary, New Brunswick, and (ARC 9953146) 
“off’? Navy Island, St. Andrews, New 
Brunswick in the Atlantic Reference Centre 
collection. 

Morphological notes.—The simple no- 
topodial seta tapers to a point, and is strong- 
ly developed, approximately 51 wm long 
and slightly more than 2 wm maximum di- 
ameter. The cutting edge of the pectinate 
seta is reinforced by 8—12 strongly devel- 
oped denticles, beyond which the blade is 
thin and serrations are not visible. Blades 
of the neuropodial falcigers are bidentate 
with serrated margins; one or two with fine 
denticulations are long and spiniger-like, 


522 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


measuring 75-85 wm in length; three to 
five, with strong denticulations, are about 
40 wm; two with strong denticulation are 
small, about 12 wm long; while some (Fig. 
10) approximate 20 wm in length. One or 
two simple neuropodial setae are present in 
the terminal 2—4 setigers. 

Remarks.—Westheide (1967) noted that 
the juveniles included in the type descrip- 
tion of M. abberans constituted a separate 
species characterized by the presence of 
eyes, short cirri, and a simple rounded anal 
lamella, features for which no ontogenetic 
evidence supported the possibility of 
change to the “‘adult’’” morphology. 

Denticulation of the blades of the pecti- 
nate setae of M. pettiboneae is difficult to 
show accurately in line drawings, and while 
the SEM micrograph of this seta in Clausen 
(1986) from a North Sea specimen, is 
slightly rotated, denticulation is similar to 
that of the New England forms. The strong- 
ly developed simple notopodial seta fre- 
quently projects from the body of preserved 
specimens and is the primary morpholog!- 
cal feature by which immature M. pettibo- 
neae can be distinguished from M. sczel- 
kowii (Table 1). Mature specimens of the 
two species are easily distinguished by o0o- 
cyte size and number (Table 1). 


M. scezelkowii 


<2 neuropodial cirri 


hemisphere 
paired 


>200 wm 
<2 


present 
9/10 


M. pettiboneae 
<2 neuropodial cirri 


hemisphere 
single 


<100 wm 
many 


present 
9/10 


M. nahantensis 


>3%X neuropodial cirri 


paired 
>200 wm 
<=13 


spatulate 
l 
9/10 


absent 


M. aggregatus 
>3 neuropodial cirri 


present 
fimbriate 
paired 
<100 pm 
many 
16/17 


Discussion 


The genus Microphthalmus is hermaph- 
roditic and its members are typical met- 
ofaunal animals attaching to hard particles 
with a caudal adhesive organ and coiling 
when disturbed. Intact specimens are infre- 
quently present in routine screenings of 
substrate, and pretreatment to release at- 
tached animals is essential in order to ob- 
tain complete specimens or population den- 
sities. 

An investigation similar to that of West- 
heide & Rieger (1987), which clarified the 
systematics of the M. listensis group of spe- 
cies, may demonstrate that the species listed 
in the synonymy of M. aberrans by Hart- 
mann-Schréder (1971) are M. pettiboneae 


M. abberans 
>3X neuropodial cirri 


absent 
fimbriate 
tor ls 
paired 
<100 wm 


Table 1.—Morphological features of New England species of Microphthalmus. 
many 
16/17 


Notopodial simple setae 


Dorsal cirri 


Penes 
Oocyte diameter 


Anal lamella 
Oocyte number 
M/f juncture 


Ocelli 


VOLUME 113, NUMBER 2 


or that speciation has occurred and they 
constitute different species. 

Very young individuals of M._ similis 
were described by Westheide (1967) from 
the intertidal at Sylt and Hartmann-Schréd- 
er & Stripp (1968) recorded sexually im- 
mature individuals up to 18 mm long and 
consisting of more than 60 segments, char- 
acteristics which agree with those of M. 
similis, from a depth of 21 m in the North 
Sea. Gonad distribution and penis charac- 
teristics have not been reported for the 
North Sea populations. The specimens list- 
ed as M. cf. similis by Westheide & Rieger 
(1978) from North Carolina, U.S.A. were 
small and immature. The distribution of go- 
nads in the two sexually mature specimens 
from the same region was not recorded by 
Westheide (1979). It is possible that they 
represent a southern distribution of M. ag- 
gregatus or constitute a separate species. 
Westheide (1977) noted that reproductive 
organs were significant but could not be in- 
cluded in a phylogenetic analysis of Mi- 
crophthalmus because of the absence of 
data for many species. However, the male- 
female (m/f) juncture of the five New Eng- 
land species (Table 1) supports the possible 
adelphotaxa indicated in his cladogram. 
The occurrence of ova in one antimere and 
sperm in the other of setigers at the m/f 
juncture of M. aggregatus has also been re- 
corded by Bobretzky (1880) for M. fragilis 
and M. similis and Clausen (1896) for M. 
cf. pettiboneae. 

The prostomium of Microphthalmus spe- 
cies is malleable, influencing its shape and 
the location of the palps and antennae. Re- 
traction of the prostomium into the peris- 
tomium may draw the median antenna back 
against or into the peristomial fold. The me- 
dian antenna is thin, rarely visible from 
ventral view unless bent to one side, and is 
often undetectable on preserved specimens. 

Ciliation has been described and figured 
for members of the Jistensis-group by 
Westheide & Rieger (1987), but other than 
for the present description of M. aggregatus 


523 


has not been recorded for other species in 
the genus. 

A pectinate notopodial seta is character- 
istic of members of the genus but is absent 
in M. hartmanae Westheide, 1977 and M. 
simplicichaetosus Westheide & Purschke, 
1992, species with fimbriate anal lamellae, 
dorsal cirri slightly more than twice as long 
as the neuropodium and in which strongly 
serrated simple setae are present in the neu- 
ropodium (Westheide 1977, Westheide & 
Purschke 1992). The pectinate notopodial 
setae may be involved in stimulating the 
glands on the cirrophore or in spreading se- 
cretions from those glands. The role of no- 
topodial setae in members of the genus with 
more than one or two simple setae in ad- 
dition to the comb seta is difficult to assess. 

The presence of the male copulatory or- 
gans in the third setiger is a generic char- 
acter; however, the morphology and num- 
ber of penes varies between species. Trans- 
mission electron microscopy (TEM) by 
Westheide (1979) demonstrated that the pe- 
nis stylets of M. cf. similis from North Car- 
olina are separate entities and not ridges in 
the wall of the penis. 

The presence of an aciculum in the pre- 
setal neuropodial lobe appears to be char- 
acteristic of the Microphthalminae sensu 
Hartmann-Schroder (1971) while absent 
from that lobe in the Hesionidae s.str. 

Westheide (1977), with ‘‘hesitation’’, 
postulated the fimbriate anal lamella as 
“‘ancestral’’. Dorsal cirri of the parapodia 
more than twice as long as the neuropodium 
appears to be characteristic of species in 
which the anal lamella is a large free fan as 
in the fimbriate species and listensis-group 
(Fig. 16). [The figure and description of the 
anal lamella of M. c.f. urofimbritta (sic) by 
Hartmann-Schréder (1960) is of a pre- 
served specimen with the lamella flexed so 
that the anal cirri and anus are terminal, and 
the fimbriate posterior margin has folded 
back beneath the terminal setigers. Artistic 
license is also apparent in the figure of the 
anal lamella of M. similis in La Greca 


524 


(1950) and M. abberans in Webster & Ben- 
edict (1887).] 

Gland openings on the anal lamella of 
fimbriate species are restricted to the ventral 
surface of the fimbrae (Fig. 13). The ar- 
rangement of these glands has not been re- 
corded for either the Jistensis-group nor for 
the species with a simple hemispherical 
(sometimes indented) lamella, and thus is 
not available for assessment at present. 

The ventriculus is attached by muscle fi- 
bers to the body wall, and is restrained pos- 
teriorly by the septum at the anterior end of 
the intestine. Telescoping of segments alters 
the topography as does eversion of the 
pharynx. Eversion in species with a bipar- 
tite pharynx (including the related genus 
Hesionides) does not significantly alter ven- 
tricular location; which is in setiger 2 in the 
listensis-group, 3 in a number of species, 5 
in aberrans, [the pharynx was reported by 
Westheide & Purschke (1992) to extend to 
setiger 4 in M. simplicichaetosus, which 
would place the ventriculus in 5], 12 in M. 
hamosus Westheide, 1982. The location of 
the ventriculus, or origin of the intestine, 
have not been routinely recorded in descrip- 
tions of species. 


Acknowledgments 


The constant advice and encouragement 
of Drs. Pettibone and Westheide throughout 
the years would have been in vain if I had 
not finally checked the type material of M. 
aberrans deposited in the USNM (courtesy 
of William Moser). The collection of Mi- 
crophthalmus species present at the Atlantic 
Reference Centre was obtained through the 
courtesy of Dr. G. Pohle. Translations of the 
two papers in Russian by Bobretsky were 
furnished by Dr. Pettibone. Alcoholic spec- 
imens of M. fragilis and M. similis from the 
Black Sea were generously furnished by Dr. 
T. Marinov in 1979 for comparative pur- 
poses. The manuscript was greatly en- 
hanced by the conscientious reviews of Drs. 
Brigitte Hilbig and Stephen Gardiner. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Literature Cited 


Alikunhi, K. H. 1948. On a new hermaphrodite species 
of Microphthalmus (Polychaeta-Hesionidae) 
from the sandy beach, Madras.—Journal of the 
Royal Asiatic Society of Bengal. Science 14: 
17-25. 

Bobretsky, N. 1870. On the fauna of the Black Sea. 
(In Russian).—Zapiski Kievskago Obshchestve 
Estestvoispytatelei 1:188—274. 

. 1880. Kutschenijn ob organach rasmnoschen- 
ija u annelid.—Zapiski Kievskago Obshchestve 
Estestvoispytatelei 6:61—86. 

Clausen, C. 1986. Microphthalmus ephippiophorus 
sp.n. (Polychaeta:Hesionidae) and two other 
Microphthalmus species from the Bergen Area, 
Western Norway.—Sarsia 71:177—191. 

Eliason, A. 1920. Biologisch-faunistische Untersu- 
chungen aus dem Oresund. V. Polychaeta.— 
Lunds Universitets Arsskrift, Avd. 2, 16(6):1— 
103. 

Hartmann-Schroéder, G. 1960. Polychaeten aus dem 
Roten Meer.—Kieler Meeresforschungen 16: 
69-125. 

1971. Annelida, Borstenwiirmer, Polychae- 

ta.—Tierwelt Deutschlands 58:1—594. 

, & K. Stripp 1968. Beitrage zur Polychaeten- 
fauna der Deutschen Bucht.—Veroffentlichun- 
gen des Instituts fiir Meeresforschung in Bre- 
merhaven 11:1—24. 

La Greca, M. 1950. Sulla presenza nel Mediterraneo 
di Lycastoides pontica (Bobr.), Microphthalmus 
fragilis (Bobr.) e Microphthalmus similis 
(Bobr.).—Annuario dell’Istituto e Museo di 
Zoologia della UnversitA di Napoli 2:1—16. 

Pettibone, M. H. 1963. Marine polychaete worms of 
the New England region. 1. Aphroditidae 
through Trochochaetidae.—Bulletin of the Unit- 
ed States National Museum 227:1—356. 

Southern, R. 1914. Archiannelida and Polychaeta, in 
Clare Island survey, pt. 47.—Proceedings of the 
Irish Academy Dublin 31:1—160. 

Webster, H. E., & J. E. Benedict. 1887. The Annelida 
Chaetopoda from Eastport, Maine.—Report of 
the United States Commissioner of Fisheries 
1885. Pp. 707-755. 

Westheide, W. 1967. Monographie der Gattungen He- 
sionides Friedrich und Microphthalmus Me- 
cznikow (Polychaeta, Hesionidae). Ein Beitrag 
zur Organisation und Biologie psammobionter 
Polychaeten.—Zeitschrift fiir Morphologie der 
Tiere 61:1—159. 

. 1973. Zwei neue interstitielle Microphthal- 

mus-Arten (Polychaeta) von den Bermudas.— 

Mikrofauna Meeresboden 14:1—16. 

. 1977. Phylogenetic systematics of the genus 

Microphthalmus (Hesionidae) together with a 

description of M. hartmanae nov. sp. Pp. 103— 


VOLUME 113, NUMBER 2 


113 in D. J. Reish & K. Fauchald, eds., Essays 
on polychaetous annelids in memory of Dr. 
Olga Hartmann.—Allan Hancock Foundation 
Special Publication. 

. 1979. Ultrastruktur der Genitalorgane inter- 
stitieller Polychaeten II. Mannliche Kopulation- 
sorgane mit intrazellularen Stilettstaben in einer 
Microphthalmus-Art.—Zoologica Scripta 8: 
111-118. 

, & G. Purschke 1992. Microphthalmus simpli- 
cichaetosus (Annelida:Polychaeta), a new he- 
sionid from the Northwestern American Pacific 


a20 


Coast with exclusively simple chaetae.—Pro- 
ceedings of the Biological Society of Washing- 
ton 105:132-135. 

, & R. M. Rieger 1978. Cuticle ultrastructure 
of hesionid polychaetes (Annelida).—Zoomor- 
phologie 91:1—18. 

, & . 1987. Systematics of the amphi- 
atlantic Microphthalmus-listensis-species-group 
(Polychaeta:Hesionidae): Facts and concepts for 
reconstruction of phylogeny and speciation.— 
Zeitschrift fiir zoologisches Systematik und Ev- 
olutionsforschung 25:12—39. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


113(2):526—531. 2000. 


Parapionosyllis cabezali, a new species of Exogoninae 
(Polychaeta: Syllidae) from Spain 


Julio Parapar, Guillermo San Martin, and Juan Moreira 


(JP) Departamento de Bioloxia Animal, Bioloxia Vexetal e Ecoloxia, Facultade de Ciencias, 
Universidade da Corufia, Campus da Zapateira, E-15071 A Coruna, Spain; 
(GSM) Laboratorio de Biologia Marina e Invertebrados. Departamento de Biologia, Unidad de 
Zoologia, Facultad de Ciencias, Universidad Aut6noma de Madrid, Canto Blanco, E-28049 


Madrid, Spain; (JM) Departamento de Ecoloxia e Bioloxia Animal, 
Facultade de Ciencias, Universidade de Vigo, E-36200, Vigo, Spain 


Abstract.—Parapionosyllis cabezali, a new species of Exogoninae (Poly- 
chaeta: Syllidae) from the Atlantic coast of Spain, is described. This new spe- 
cies is characterised by the shape of the simple and compound setae, small 
dorso-ventral gradation in the length of the blades, and by the presence along 
the body of two types of parapodial glands, which open dorsally separately 
from each other. 

Campoy (1982) identified three specimens collected in Punta Endata, Gui- 
puzcoa, NE Atlantic, Spain, as Parapionosyllis cf. gestans (Pierantoni, 1903). 
Subsequently, during a study of the taxonomy and ecology of the sublittoral 
soft bottom benthic polychaetous annelids in the Ria de Ferrol and Ria de 
Baiona, both located on the Atlantic coast of NW Spain, more specimens of 
this species were collected and reported by Parapar (1991), Parapar et al. (1994) 
and Moreira (1999). Examination by SEM of several specimens collected in 
the Ria de Baiona (NW Spain) has provided evidence that these organisms 
represent a new species. The specimens are characterized by the shape and 
dorso-ventral gradation in the length of the compound setal blades and by the 
presence along the body of two types of parapodial glands. In this paper, we 
describe and discuss this new species. The diagnosis of the genus was provided 


by Fauvel (1923) and San Martin (1984). 


Materials and Methods 


The type specimens (holotype and para- 
types) were collected by the junior author 
(JM) in littoral soft bottom areas of the Ria 
de Baiona (Galicia, NW Spain) at 9-12 m 
depth. Additional specimens were collected 
in Galicia, in the Ria de Ferrol. Samples 
were taken both by means of a naturalist 
rectangular dredge and a Van Veen dredge 
in Dec 1995 in the case of the type series 
and in 1987 and 1989 in the Ria de Ferrol. 
Samples with the letters VD denote sam- 
pling for the vertical distribution of animals 
in the sediment. In this study, samples were 
taken by scuba diving and by pushing a 50 


cm corer into the sediment. Animals taken 
from the sediment were fixed in 10% buff- 
ered formalin and preserved in 70% etha- 
nol. Body and setae measurements provided 
in the description refer to the holotype, the 
width was measured across the proventri- 
culous and excludes cirri, parapodia and se- 
tae. Observations, drawings, and measure- 
ments were obtained using a microscope 
with interference contrast optics (Nomar- 
sky). Drawings were made with the aid of 
a drawing tube. The SEM micrographs 
were taken at the Servicio Interdepartamen- 
tal de Investigacié6n of the Universidad Au- 
tonoma de Madrid, Spain. The types are de- 


VOLUME 113, NUMBER 2 


posited in the Museo Nacional de Ciencias 
Naturales de Madrid, Spain. 


Family Syllidae Grube, 1850 
Subfamily Exogoninae Rioja, 1925 
Genus Parapionosyllis Fauvel, 1923 

Parapionosyllis cabezali, new species 
Figss 1,72 


Parapionosyllis cf. gestans.—Campoy, 
1982: 267-269, fig. 18.—Parapar et al., 
1994: 96-97, fig. 2.—Moreira, 1999: 
319-323, fig. 26. [Not Parapionosyllis 
gestans Pierantoni, 1903]. 


Material examined.—Ria de Baiona (Ga- 
licia, NW Spain). Holotype and 23 para- 
types. Sta. 8, Dec 1995, coarse sand, 12 m 
(10 specimens); Sta. 9, Dec 1995, medium 
sand, 10 m (10); Sta. 12, Dec 1995, coarse 
sand, 9 m (4). 

Additional material.—Ria de Ferrol (Ga- 
licia, NW Spain), 39 specimens. Sta. 1, Jul 
1987, muddy sand, 15 m (1 specimen). Sta. 
3, Jun 1987, muddy sand, 11 m (1). Sta. 10, 
May 1987, muddy sand, 18 m (2). Sta. 11, 
Jul 1987, muddy sand, 20 m (1). Sta. 13, 
May 1987, coarse sand, 14 m (2). Sta. 
ISVD, Aug 1989, 20-25 cm in sediment 
gem, 17 m (1). Sta. 17, Jul: 1987, very 
eaarsersand, 25 mi (1). Stas, 22. Oct 1987, 
very coarse sand, 20 m (1). Sta. 22VD, Aug 
1989, O-—5 cm in sediment depth, 20 m (8). 
Sta. 22VD, Aug 1989, 5—10 cm in sediment 
depth (6). Sta. 22VD, Aug 1989, 10-15 cm 
in sediment depth (1). Sta. 22VD, Aug 
1989, 15—20 cm in sediment depth (2). Sta. 
26, Aug 1987, very coarse sand, 15 m (5). 
Sta. 37VD, Aug 1989, 0-5 cm in sediment 
depth, 15 m (7). Ensenada de Baiona (Ga- 
licia, NW Spain), 704 specimens. Sta. 8, 
Dec 1995, coarse sand, 12 m (327 speci- 
mens). Sta. 9, Dec 1995, medium sand, 10 
mm (120). Sta. 12, Dec 1995, coarse sand, 9 
m (236). Sta. 13, Dec 1995, medium sand, 
8 m (18). Sta. 14, Dec 1995, medium sand, 
9 m (3). 

Description.—Body relatively long and 
thin, with well marked segments. Holotype 


S27 


3.36 mm long, 0.24 mm wide with 39 seg- 
ments. Prostomium semicircular to pentag- 
onal (Figs. 1A, 2A); four eyes in an open 
trapezoidal arrangement and two very small 
anterior eyespots; eyes disappear soon after 
fixation in alcohol. Antennae bottle or nine- 
pin shaped, relatively long; median antenna 
somewhat longer than the lateral ones, ap- 
proximately of the same length as the pro- 
stomium and palps together; lateral anten- 
nae inserted ahead of anterior eyes; median 
antenna between posterior eyes, near the 
posterior margin of the prostomium. Palps 
robust, equal to or slightly longer than the 
prostomium, fused at bases but separated in 
the anterior one-half; a slight junction su- 
ture occasionally observed. Tentacular seg- 
ment distinct but shorter and narrower than 
the following segments; a pair of tentacular 
cirri similar in shape to antennae but ap- 
proximately one-half the length of the lat- 
eral ones. Two ciliated nuchal organs (Fig. 
2A, B) deeply imbedded between the pos- 
terior margin of the prostomium and the an- 
terior margin of the tentacular segment. 
Anterior dorsal cirri short and oval, with 
slightly enlarged bases (Figs. 1A, 2A); pro- 
gressively more elongated and slender; 
slightly longer than the parapodial lobes; 
posterior dorsal cirri thinner and longer 
than anterior ones (Fig. 1B). Two types of 
parapodial glands (Fig. 1A, B); the smaller 
ones beginning in the most anterior seg- 
ments, including the first setiger, with gran- 
ular content, and the larger ones with fi- 
brillar material, located near the central 
body line and starting from the proventric- 
ular segments. These parapodial glands 
open dorsally near the dorsal cirri, separate 
from each other; each pore is provided with 
a mobile, digitiform papilla which ejects the 
products of these glands (Fig. 2C, D). Both 
types of glands are dark red in colour. Para- 
podial lobes short and conical, ending in a 
small rounded papilla (Fig. 1C). Ventral cir- 
ri digitiform, long and slender, shorter than 
the parapodial lobes or similar in length. 
Most anterior parapodia have about nine 
compound setae; six or seven in the re- 


528 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 1. Parapionosyllis cabezali, new species. Holotype. A, anterior end and midbody, dorsal view; B, 
posterior end, dorsal view; C, parapodium of midbody, posterior view; D, anterior dorsal simple seta; E, anterior 
compound setae; EK aciculum; G, posterior dorsal simple seta; H, posterior compound setae; I, posterior ventral 
simple seta. Scale bar a: 0.25 mm for A, B and 60 pm for C; scale bar b: 25 wm for D to I. 


VOLUME 113, NUMBER 2 


WN 
nN 
\O 


t he 


. iia 
- 7 3 By 
F 1 rt Wiebe hej 
2 : r ~~ ? 
° F © 
| a | a, 
SU / 4 ij Sh 
Magn r — ———s ere A aqn 
Ox ,/ {/ 3000x 


Fig. 2. Parapionosyllis cabezali, new species. SEM micrographs. A, anterior end, dorsal view; B, detail of 
a nuchal organ opening; C, parapodial glands pores and papillae (arrows); D, detail of fibrilar material protruding 
from a parapodial gland; E. long bladed, dorsal compound seta from anterior parapodium; EK mid-length and 
short compound setae from midbody; G, posterior dorsal simple seta; H, posterior ventral simple seta. 


530 


maining anterior parapodia. Anterior dorsal 
compound setae with thick shafts and het- 
erogomph articulation (Fig. 1E); median 
and ventral shafts with hemigomph articu- 
lation (Fig. 2E, F). Shafts apparently 
smooth but having a few subdistal spines. 
Blades unidentate with spines on the mar- 
gin and a longer sub-distal spine more ev- 
ident on more dorsal and anterior setae. 
Dorso-ventral gradation in blade length; 
blades of the dorsal-most compound setae 
about 25 ym in most anterior setigers, 20 
im in the median ones and 15 pm in pos- 
terior-most ones; with moderately long 
spines on the margin; blades progressively 
shorter towards the ventral part of the para- 
podium (Fig. 1E, H), 10 wm in the ventral 
most seta with shorter spines on the margin. 
Solitary dorsal simple seta on each post- 
proventricular parapodium, slightly thicker 
than the shafts of compound setae, provided 
with various spines on the margin, one of 
which thicker and longer (Fig. 1D); most 
posterior dorsal simple setae have fewer 
spines on margin but are provided with a 
larger subdistal spine (Figs. 1G, 2G). Soli- 
tary ventral simple setae sigmoid, uniden- 
tate, apparently smooth (Fig. 11) but pro- 
vided with a few minute subdistal spines 
(Fig. 2H). Solitary aciculum per parapodi- 
um, relatively thick and with a rounded, en- 
larged and apparently hollow tip (Fig. 1F). 

Pharynx relatively long and thin, extend- 
ing through five or six segments (Fig. 1A); 
pharyngeal tooth located near the anterior 
margin; anterior end of pharynx surrounded 
by 10 papillae (Fig. 2A). Proventriculous 
shorter than the pharynx, extending through 
three or four segments, with about 22 rows 
of muscle cells. Pygidium small, semicir- 
cular, with two long and thin anal cirri, lon- 
ger than the antennae (Fig. 1B). Mature 
males provided with natatory capillary se- 
tae and sperm packets in each segment be- 
tween setigers 11 and 27; females carry 
eggs ventrally. 

Ecology.—Parapionosyllis cabezali, was 
found in Punta Endata in sublittoral muddy 
sand (70 m depth), in the Ria de Ferrol in 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


sublittoral muddy sand, coarse sand and 
very coarse sand (11—25 m depth) with or- 
ganic matter content between 0.12% and 
0.20% and reaching 20 cm depth in sedi- 
ment. In the Ria de Baiona this species was 
found in sublittoral medium sand and 
coarse sand (8—12 m depth). 
Remarks.—Campoy (1982) and, more re- 
cently, Parapar et al. (1994) reported this 
species as Parapionosyllis cf. gestans (Pier- 
antoni, 1903), due to its similarity to P. ges- 
tans [as described by Fauvel (1923) and 
Cognetti (1965)]. A more detailed study of 
the specimens revealed two main differenc- 
es between the species. P. cabezali shows 
a moderate dorso-ventral gradation in the 
length of the blades of the compound setae, 
while in P. gestans all blades are similar in 
length (Cognetti 1965). Unlike Campoy 
(1982), we consider the setae illustrated to 
be an accurate representation of the com- 
plete set of setae in each parapodium. In the 
same figure, the author provides detailed 
sketches of the setae of three other species 
(Parapionosyllis minuta, Parapionosyllis 
elegans and Parapionosyllis labronica), 
drawing the same number of setae for all of 
them and showing the variation in the size 
of the blades where it exists. Also, all spec- 
imens of P. cabezali had numerous and 
conspicuous parapodial glands, of two 
kinds and different in size, which are not 
described in the Mediterranean species. Per- 
kins (1981) also report the presence of 
paired parapodial glands in specimens of 
Parapionosyllis longocirrata (Webster & 
Benedict 1884) from the Atlantic coast of 
the United States, but this species differs 
from P. cabezali in that it has much larger 
dorsal cirri, in the shape and spinulation of 
the anterior dorsal simple setae and in the 
length of the dorsal-most compound setae 
blades (15 wm in P. longocirrata and 25 
wm in P. cabezali). Parapionosyllis brevi- 
cirra Day, 1954, a species also present in 
the Atlantic coast of the Iberian Peninsula 
(Parapar et al. 1994) and western Mediter- 
ranean Sea, occasionally shows granular 
and fibrillar inclusions in the dorsal part of 


VOLUME 113, NUMBER 2 


the parapodia (San Martin 1984), but this 
well known species has much longer dorsal 
compound setae blades (45 um) than P. ca- 
bezali. The type-series of P. gestans (Pier- 
antoni, 1903) was apparently lost, and it 
was not possible to examine any type spec- 
imens of this species. San Martin (1984) 
recorded 12 described species of Parapion- 
osyllis and provided a key for identification 
in which P. gestans (Pierantoni, 1903) was 
distinguished from P. cf. gestans of Cam- 
poy (1982). 

Distribution.—Parapionosyllis cabezali 
seems to be distributed along the Iberian 
Atlantic coast, from Basque Country (Cam- 
poy 1982, Aguirrezabalaga 1984), Galicia 
(NW Spain) in the Ria de Ferrol (Parapar 
et al. 1994) and Ria de Baiona (Moreira 
1999) to Punta Umbria (Huelva, SW Spain) 
(Rodriguez & Viéitez 1992), although it has 
not been reported in Portugal. 

Etymology.—The species is named in 
fond memory and honour of the late Luis 
Cabezal Gomez, lecturer in the Escuela Pol- 
itécnica Superior de Lugo (Universidade de 
Santiago, Spain), brother-in-law of the first 
author, in recognition of his love for nature 
and friendship. 


Acknowledgments 


This paper was partially supported by the 
projects “‘Aplicaciéns morfofuncionais da 
quetotaxia de anélidos acuaticos de Gali- 
cia” (XUGA10305B98) and ‘‘Fauna Ibéri- 
ca IV”’ (PB950235) supported by the Xunta 
de Galicia and Comision Interministerial de 
Ciencia y Tecnologia respectively. The 
drawings were made by J. Corbera, illustra- 
tor for the project “Fauna Ibérica’’, and the 
SEM micrographs were taken by E. Salva- 


bse 


dor of the Servicio Interdepartamental de 
Investigacion (U.A.M., Spain). 


Literature Cited 


Aguirrezabalaga, F 1984. Contribuci6n al conocimien- 
to de los Anélidos Poliquetos de la costa de 
Guiptizcoa.—Munibe 36:119—130. 

Campoy, A. 1982. Fauna de los Anélidos Poliquetos 
de la Peninsula [bérica.—Publicaciones de Biol- 
ogia de la Universidad de Navarra, Serie Zool- 
6gica 7(1—2):1—781. 

Cognetti, G. 1965. Sillidi mesopsammici delle secche 
della Meloria (Livorno).—Archivio Zoologico 
Italiano 50:65-—72. 

Fauvel, P. 1923. Polychétes errantes. Faune de France, 
5. Paul Lechevalier, Paris, 488 pp. 

Moreira, J. 1999. Anélidos Poliquetos de sustratos 
blandos infralitorales de la Ensenada de Baiona 
(Galicia). Unpublished M.S. thesis, University 
of Vigo, Vigo (Spain), 510 pp. 

Parapar, J. 1991. Anélidos poliquetos benténicos de la 
Ria de Ferrol (Galicia). Unpublished Ph.D. the- 
sis, University of Santiago, Santiago de Com- 
postela (Spain), 1104 pp. 

, G. San Martin, C. Besteiro & V. Urgormi, 
1994. Aspectos sistematicos y ecoldégicos de las 
subfamilias Eusylline y Exogoninae (Polychae- 
ta, Syllidae) en la Ria de Ferrol (Galicia, NO 
Espana).—Boletin de la Real Sociedad Espa- 
Nola de Historia Natural, Seccion Biologia 
91(1-4):91-101. 

Perkins, T. H. 1981. Syllidae (Polychaeta), principally 
from Florida, with descriptions of a new genus 
and twenty-one new species.—Proceedings of 
the Biological Society of Washington 93:1080— 
BLT: 

Pierantoni, P. 1903. La gestazioni esterna.—Archivio 
Zoologico 1(3—4):231—252. 

Rodriguez, C. V. & J. M. Viéitez. 1992. Macrofauna 
bentonica de los primeros metros del piso in- 
fralitoral de la costa de Punta Umbria (Huel- 
va).—Boletin del Instituto Espanol de Oceano- 
grafia 8(2):327-338. 

San Martin, G. 1984. Estudio biogeografico, faunistico 
y sistematico de los poliquetos de la familia Sil- 
idos (Syllidae: Polychaeta) en Baleares.—Uni- 
versidad Complutense, Madrid, Tesis Doctora- 
les, 187/84, 529 pp. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 
113(2):532—541. 2000. 


Paralarval gonatid squids (Cephalopoda: Oegopsida) from the 
Mid-North Atlantic Ocean 


Luisa I. Falcon, Michael Vecchione, and Clyde E E. Roper 


(LIF) Instituto de Ciencias del Mar y Limnologia, Universidad Nacional Autonoma de Mexico, 


Apdo. Postal 70-305, D.E 045510, D.E, Mexico and National Museum of Natural History, 
Research Training Program, Smithsonian Institution, Washington, D.C. 20560, U.S.A. 
(MV) National Marine Fisheries Service, National Systematics Laboratory, 
National Museum of Natural History, Washington, D.C. 20560-0153, U.S.A; 


(CFER) Department of Invertebrate Zoology-Mollusks, National Museum of Natural History, 


Smithsonian Institution, Washington, D.C. 20560-0118, U.S.A. 


Abstract.—Ninety six gonatid cephalopod specimens (Oegopsida: Gonati- 
dae) from the University of Amsterdam Mid-North Atlantic Plankton Expedi- 
tions were analyzed and two species were identified: Gonatus steenstrupi (Kris- 
tensen 1981) and Gonatus fabricii (Lichtenstein 1818). Gonatids were collected 
only in spring and summer, despite sampling in autumn and winter. This paper 
describes aspects of their development and reports their geographical distri- 
bution in the central North Atlantic Ocean. Chromatophore patterns were the 
most consistently useful characters for distinguishing between the species. 
Among 34 measurements, Tentacle Length (TtL) relative to Dorsal Mantle 
Length (ML) and number of suckers on Arms I-IV were useful for distinguish- 
ing specimens >13 mm ML. Both species develop hooks from suckers on the 
arms and tentacular clubs at ML >20 mm. Subtle differences were noted in 
the morphology of the funnel pads except in the smallest specimens. Specimens 
of G. steenstrupi >20 mm ML were collected at greater depths (250 to 995 
m) than the smaller specimens (found at depths <200 m). Our data suggest 
that 20 mm ML is the point of transition between paralarvae and juveniles of 
G. steenstrupi, because specimens larger than 20 mm ML have well defined 


hooks, and a juvenile vertical distribution is established. 


Knowledge of early-life-history stages is 
required for comprehensive understanding 
of the ecology of squids. These stages pre- 
sumably are most vulnerable to starvation 
and predation and they occupy a separate 
niche from older conspecifics (Vecchione 
1987). Because of controversy about use of 
the term “‘larva’’ for early life history stag- 
es in cephalopods, Young & Harman (1988: 
202) introduced the term “‘paralarva’’ for 
‘‘a cephalopod of the first post-hatching 
growth stage that is pelagic in near-surface 
waters during the day and that has a dis- 
tinctly different mode of life from that of 
older conspecific individuals.’’ The paralar- 


val concept includes both morphological 
and ecological features, in contrast to the 
definition of a “‘larva’’, which is based on 
morphological differences from the adults. 

The University of Amsterdam, Nether- 
lands, conducted four research expeditions 
in the North Atlantic Ocean between 55°N 
and 24°N approximately along 30°W lon- 
gitude. This research was designed to ‘“‘elu- 
cidate the patterns of latitudinal diversity, 
taxonomical variation below species level, 
vertical variation and interaction of climate, 
hydrographic features and ecology on mor- 
phological variation of marine plankton” 
(Van der Spoel 1981:1). The expeditions 


VOLUME 113, NUMBER 2 


were conducted during four consecutive 
years (1980-1983), each during a different 
season (Van der Spoel 1981, 1985; Van der 
Spoel & Meerding 1983). Discrete-depth 
samples were collected using opening/clos- 
ing nets. Cephalopod paralarvae sorted from 
the samples were donated to the National 
Museum of Natural History, Smithsonian In- 
stitution for systematic and ecological stud- 
ies. Shea (1995) sorted the material into 
families in preparation for subsequent stud- 
ies. The present paper reports on paralarval 
development and distribution of the squid 
family Gonatidae in these samples. 

The identification of paralarval cephalo- 
pod stages is difficult because of insuffi- 
cient collections which often come from in- 
adequate sampling devices and methods, 
and because of poorly understood taxono- 
my, even in adults (Vecchione 1987). This 
is true for the genus Gonatus in the North 
Atlantic Ocean in which only one arctic/ 
boreal species was recognized, Gonatus fa- 
bricii (Lichtenstein 1818), until Kristensen 
(1981) described Gonatus steenstrupi from 
boreal waters. At least five Gonatus species 
are found in the North Pacific Ocean and 
one in the Southern Ocean (Kristensen 
1981). The primary morphological charac- 
ters used to separate adults of the two North 
Atlantic species are (a) the presence or ab- 
sence of chromatophores on the ventral sur- 
face of the head, (b) the shape of the funnel 
organ, and (c) the patterns of hooks and 
suckers on the tentacular clubs. The onset 
of formation of hooks from suckers both on 
the tentacular clubs and arms I-III seems a 
good character to define the differences be- 
tween paralarvae and juveniles in gonatids 
(Young 1972, Kristensen 1977a). The pres- 
ence of hooks presumably indicates a 
change in feeding and therefore in the 
squid’s role in the oceanic trophic structure. 

The main goals of the present study were 
to identify the species of paralarval gona- 
tids in the Mid-North Atlantic collections, 
to analyze early-life-history features that 
could separate paralarvae from juveniles, 


533 


and to determine the distribution of the two 
species in these samples. 


Methods 


Gonatid specimens were arranged by size 
and their taxonomic identification began 
with the largest specimens, then proceeded 
sequentially to the smallest specimens. We 
looked in particular for previously unrec- 
ognized taxonomic characters for these par- 
alarvae, in addition to using various taxo- 
nomic guides (Kristensen 1981, Nesis 1987, 
Roper et al. 1984). Fifteen specimens were 
damaged and were excluded from some of 
the quantitative analysis. Characters were 
measured or counted on each of the 81 un- 
damaged gonatids following Roper & Voss 
(1983), including: 

ML (Dorsal Mantle Length); MW (Mantle 
Width); HL (Head Length); HW (Head 
Width); ED (Eye Diameter); FL (Fin 
Length); FW (Fin Width); TL (Total 
Length); TtL (Tentacle Length); CL (Club 
Length); AHI-IV (number of hooks on 
Arms I-IV); ASI-IV (Arm I-IV Sucker 
counts); ALI-IV (Arm I-IV Length); AWI- 
IV (Arm I-IV Width). 

We also measured the following characters: 


D (Dactylus Length); M (Manus Length); 
C (Carpus Length); CS (Club Sucker 
Length); ForgL (Funnel Organ Length); 
ForgW (Funnel Organ Width); ForgLl 
(Funnel Organ Dorsal Pad Length); 
ForgW1 (Funnel Organ Dorsal Pad Width). 
Characters on damaged specimens were 
measured when their condition permitted. 
Three specimens of different sizes from 
each of the two species were selected based 
on condition, and their third arms and ten- 
tacular clubs (left for G. fabricii and right 
for G. steenstrupi, because of specimen 
damage) were removed for scanning elec- 
tron microscopical (SEM) analysis. The 
specimens had been fixed in formalin and 
preserved in 45% isopropanol. Tissue for 
SEM was transferred through a dehydration 
series to 100% ETOH prior to critical-point 
drying, which was conducted using a Den- 


534 


ton Vacuum-1, Critical-Point Dryer. The 
arms and tentacular clubs were examined 
using a Leica 440 SEM to find morpholog- 
ical features that could separate the species. 


Results 


In total, 96 specimens of the family Gon- 
atidae were collected during the spring 
(1980) and summer (1983) cruises. No gon- 
atids were found in samples from the other 
two cruises. The presence of two chromato- 
phores on the ventral side of the head in G. 
fabricii and their absence in G. steenstrupi, 
the shape of the funnel organ and the de- 
velopment and pattern of hooks and suckers 
on the tentacular club were the basic fea- 
tures that we used to distinguish these two 
species. The 96 gonatids were separated 
into 43 G. fabricii and 38 G. steenstrupi, 
the remaining 15 specimens were too dam- 
aged to determine species with certainty. 
The 81 undamaged specimens ranged in 
size from 1.6 to 31.6 mm ML in G. steen- 
strupi, 3.3 to 24.1 mm in G. fabricii and 
3.0 to 10.8 mm for the damaged specimens. 

Gonatus fabricii paralarvae are charac- 
terized by the presence of a pair of round 
or oblong chromatophores on the ventral 
surface of the head slightly anterior to the 
ocular axis (see Kristensen 1981:67, fig. 3 
for specimens larger than those reported 
here). The dorsal pad of the funnel organ 
in this species has an inverted V-shape with 
very straight lateral sides (Fig. 1; cf. Kris- 
tensen 1981:69, fig. 5). After the largest 
specimen of G. fabricii (24.1 mm ML) was 
identified, published taxonomic characters 
could then be recognized in progressively 
smaller specimens. The presence of chro- 
matophores was the primary character used 
to identify the smallest specimens of G. fa- 
bricii. The funnel organ is so small in spec- 
imens <3.6 mm ML that its shape can not 
be determined confidently. Scanning elec- 
tron microscopy on specimens of 24.1, 
14.6, and 7.2 mm ML revealed that the pat- 
terns of hooks and suckers on the tentacular 
clubs of the largest G. fabricii were similar 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


to those described by Kristensen (1977a, 
1981). The largest specimen (24.1 mm ML) 
had one large hook, with three small hooks 
and a sucker proximal to the large hook. A 
concentration of small suckers occurs on 
both dorsal and ventral sides of the club, 
especially on the proximal end, where the 
suckers form a large cluster (Fig. 2d). 

The largest specimens of G. steenstrupi 
(e.g., 31.6 mm ML) were identified follow- 
ing the description of the holotype (Kris- 
tensen 1981). The shape of the funnel organ 
is characterized by a slight curve on the lat- 
eral edges of the dorsal pad (Kristensen 
1981: 83, fig. 20). This character can be 
seen in our largest specimens (Fig. 3). The 
absence of chromatophores on the ventral 
surface of the head is the primary distin- 
guishing character of the smallest G. steen- 
strupi paralarvae available, which could not 
be identified based on funnel-organ mor- 
phology. Scanning electron microscopy was 
used to examine specimens of 31.6, 19.2 
and 9.6 mm ML. The tentacular club of the 
largest specimen under SEM had the pat- 
tern of hooks and suckers described by 
Kristensen (1981) in which one large hook 
is preceded proximally with four small 
hooks and no suckers (Fig. 2c). At <20 mm 
ML the hooks are not yet well developed 
(Fig. 2a, b). Gonatus steenstrupi and G. fa- 
bricii smaller than 20 mm ML have similar 
sucker patterns in this central series on their 
tentacular clubs. 

The scatterplot of the relation between 
TtL and ML for all the undamaged speci- 
mens indicates a growth curve for G. steen- 
strupi that diverges from that of G. fabricii 
at ML >13 mm (Fig. 4). The relation be- 
tween TtL and ML is more linear for G. 
fabricii than for G. steenstrupi, perhaps be- 
cause of the lack of large G. fabricii spec- 
imens. However, all G. steenstrupi >13 mm 
ML had tentacles that were longer than 
those of similar-sized G. fabricii. Extension 
of a quadratic function line fitted to the G. 
fabricii data indicates that the difference in 
TtL between species likely continues at 
larger sizes, although a shift in growth pa- 


VOLUME 113, NUMBER 2 


Fig. 1. Growth series of G. fabricii. Scale bar = 1 mm. 


Nn 
eS) 
OY 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


-e. 5 
‘ ‘& 
“ie RAL AS og an Oa ; 
} bt 


ee 


rt 


Fig. 2. Tentacular clubs. a—c) G. steenstrupi, 9.6, 19.2, and 31.6 mm ML respectively, d) G. Sabet 24.1 
mm ML. Scale bars: a) 200 um, b) 300 wm, c) 1 mm, d) 1 mm. 


rameters at the transition between paralar-_ ular clubs are found in specimens of both 
vae and juveniles (Shea 1995) could either species larger than 20 mm ML. Kristensen 
reduce or increase interspecific differences. (1981) described major differences between 

Well-developed hooks on arms and ten- these species on specimens larger than 37 
tacular clubs and well-differentiated tentac- mm gladius length, which is equivalent to 


VOLUME 113, NUMBER 2 


Fig. 3. Growth series of G. steenstrupi. Scale bar = 


ML in gonatids. Our SEM analysis of arm 
III suckers for G. steenstrupi of 31.6 mm 
ML (Fig. 5a) and G. fabricii of 24.1 mm 
ML (Fig. 5c) shows that no obvious differ- 
ences occur, although the chitinous teeth of 


y 


1 mm. 


the internal ring of the suckers are sharper 
in G. steenstrupi. There is no difference be- 
tween species in this character on smaller 
specimens (Fig. 5b, d). 

The vertical distribution of the paralar- 


40 
Bi Gonmaius steensirupt 
@ Gonatus jabricti 
€ 30 
E 
< 
2 
@ 20 
& 
oO 
s 
eae 
2 40 
0 % 
0 10 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


20 
Mantle Length (mm) 


30 40 


Dorsal Mantle Length (ML) vs. Tentacle Length (TtL). Quadratic function lines are fitted to the 


scatterplots of the two species simply to identify trends in the data, rather than to test hypotheses of differences. 


vae was analyzed for each cruise (Fig. 6). 
During the spring cruise (1980), G. steen- 
strupi larger than 20 mm ML were cap- 
tured at the deepest sampling stations (250 
to 995 m). The specimens smaller than 20 
mm ML were collected in the upper 200 
m, although one specimen of 2.8 mm ML 
was caught between 390 and 510 m. Gon- 
atus fabricii differed in spring vertical dis- 
tribution, exhibiting greater variability in 
depths for all sizes. All G. fabricii speci- 
mens were captured shallower than 400 m 
although samples were collected from 


depths as great as 1750 m. The summer 
cruise (1983) showed a similar distribution 
for G. steenstrupi where specimens larger 
than 20 mm ML were caught between 490 
and 995 m and those smaller than 20 mm 
ML were found in the upper 50 m. Gon- 
atus fabricii were again found with greater 
variability in their vertical distribution, as 
specimens with 7.2 mm and 15.6 mm ML 
were collected at the deepest stations 
(1750 to 1000 m), while the rest of the 
sampled population was found in the upper 
100 m. 


VOLUME 113, NUMBER 2 


539 


Fig. 3. 
24.1 mm ML, d) G. fabricii, 14.6 mm ML. Scale bars: a) 20 wm, b) 20 wm, c) 30 wm, d) 10 pm. 


Discussion 


Van der Spoel (1981) reported that dur- 
ing the 1980 spring cruise subarctic polar 
water was present at depths greater than 
500 m north of 50°N. The 1983 summer 
cruise also may have sampled subarctic wa- 
ter north of 53°N, as well as an isothermal 
layer above the thermocline at depths great- 
er than 90 m near 55°N. Both the spring 
(1980) and summer (1983) cruises found a 
well marked northern branch of the North 
Atlantic Drift, although the southern branch 
was much more marked in 1983 (Van der 
Spoel 1985). The presence of Gonatus in 
these samples likely resulted from the pres- 
ence of cold subarctic water in the area. 


Suckers in Arm III. a) G. steenstrupi, 31.6 mm ML, b) G. steenstrupi, 19.2 mm ML, c) G. fabricii, 


Both Gonatus fabricii and Gonatus steen- 
strupi, occurred in a previously unreported 
distribution (54°53'54”"N 029°55'48"W to 
48°58'54”"N 030°01'18”W). Earlier reports 
(Kristensen 1977b, 1981: 62, fig. 1) listed 
their distribution as nearer to the coast and 
much more northerly, especially G. fabricii. 
This new distribution extends the known oc- 
currence of both species far offshore towards 
the Central North Atlantic Ocean and more 
southerly. 

The change in vertical distribution by G. 
steenstrupi larger than 20 mm ML and the 
coincident presence above that size of well- 
developed hooks on the arms and tentacular 
clubs of both species may define the tran- 


540 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 
Mantle Length (mm) 
1980 1983 
a 0 10 20 30 40 b 0 10 20 30 40 


Depth (m) 


Gonatus steenstrupi 


1500 


Fig. 6. Depth of capture vs Dorsal Mantle Length (ML). a) G. fabricii in spring 1980, b) G. fabricii in 
summer 1983, c) G. steenstrupi in spring 1980, b) G. steenstrupi in summer 1983. Diamonds indicate depth at 
opening of net, squares depth at closing of net, vertical lines, depth range sampled. 


sition between paralarvae and juveniles. 
Differences between the species on speci- 
mens larger than 20 mm ML include the 
pattern of hook development from suckers 
both on the tentacular clubs and the arms. 
Additional specimens are needed to confirm 
the apparent transition point between early- 
life-history stages for G. steenstrupi and to 
delineate such stages for G. fabricii. The 
results reported here, however, do indicate 
changes in both the morphological and eco- 
logical characteristics at sizes about 20 mm 
ML, particularly for G. steenstrupi. 


Acknowledgments 


The authors sincerely thank the Smith- 
sonian Institution Women’s Committee, 
which provided the funding for Luisa Fal- 
con’s internship in the National Museum of 
Natural History Research Training Pro- 
gram. We are grateful to Guido Pastorino 
for his help with the analysis of the data. 
We acknowledge the SEM laboratory staff 
at the NMNH, Walt Brown and Susann Bra- 


den, for their assistance on our research. 
Michael Sweeney, Division of Mollusks, 
NMNH was of invaluable help. The authors 
are grateful to the Research Training Pro- 
gram Coordinator, Mary Sangrey, for mak- 
ing this research possible. We appreciate the 
careful reviews of an earlier draft by Uwe 
Piatkowski and an annonymous reviewer. 


Literature Cited 


Kristensen, T. K. 1977a. Scanning electron microscopy 
of hook development in Gonatus fabricii (Lich- 
tenstein, 1818) (Mollusca: Cephalopoda). Vi- 
denskabelige meddelelser Fra Dansk naturhis- 
torisk Forening 140:111—116. 

. 1977b. Hatching, growth and distribution of 

juvenile Gonatus fabricii (Mollusca: Cephalop- 

oda) in Greenland waters.—Astarte 10:21—28. 

. 1981. The genus Gonatus Grey, 1840 (Mol- 
lusca: Cephalopoda) in the North Atlantic. A 
revision of the North Atlantic species and de- 
scription of Gonatus steenstrupi n. sp.—Steen- 
strupia 7(4):61—99. 

Nesis, K. N. 1987. Cephalopods of the World. [English 
translation from 1982 Russian edition]. TFH 
Publications, Neptune, New Jersey, 351 pp. 


VOLUME 113, NUMBER 2 


Roper, C. FE E., & G. L. Voss. 1983. Guidelines for 
taxonomic descriptions of cephalopod spe- 
cies—Memoirs of the National Museum, Vic- 
toria 44:49-—63. 

, M. J. Sweeney, & C. E. Nauen. 1984. FAO 
Species Catalogue. Cephalopods of the World. 
An annotated and illustrated catalogue of spe- 
cies of interest to fisheries. FAO Fishery Syn- 
opsis (125) vol. 3. 277 pp. 

Shea, E. K. 1995. The early life histories of three fam- 
ilies of cephalopods... and an examination of 
the concept of paralarva. Unpublished M.A. 
thesis, College of William and Mary, 134 pp. 

Vecchione M. 1987. Juvenile ecology. Pp. 61—84 in P. 
R. Boyle, ed., Cephalopod life cycles. vol. 2, 
Comparative Reviews. Academic Press, Lon- 
don. 

Van der Spoel, S. 1981. List of discrete depth samples 
and open net hauls of the Amsterdam Mid- 
North Atlantic Plankton Expedition 1980 (pro- 


541 


ject 101 A).—Bulletin Zoologisch Museum, 

Universiteit van Amsterdam 8 (1):1—10. 

. 1985. List of discrete depth samples and open 

net hauls of the Amsterdam Mid-North Atlantic 

Plankton expeditions 1982 and 1983 (Project 

101 A).—Bulletin Zoologisch Museum, Univ- 

ersiteit van Amsterdam 10 (17):129-—152. 

, & A. G. H. A. Meerding. 1983. List of dis- 
crete depth samples and open net hauls of the 
Amsterdam Mid-North Atlantic Plankton Ex- 
pedition 1981 (Project 101 A).—Bulletin Zool- 
ogisch Museum, Universiteit van Amsterdam 9 
(9):77-91. 

Young, R. E. 1972. The systematics and areal distri- 
bution of pelagic cephalopods from the seas off 
Southern California.—Smithsonian Contribu- 
tions to Zoology 97, 59 pp. 

, & R. E Harman. 1988. “‘Larvae’’, “‘paralar- 

vae”’ and “‘subadult”’ in cephalopod terminolo- 

gy.—Malacologia 29 (1):201-—207. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


113(2):542—560. 2000. 


Parabuccinum, a new genus of Magellanic buccinulid 
(Gastropoda: Neogastropoda), with a description of a new species 


M. G. Harasewych, Yuri I. Kantor, and Katrin Linse 


(MGH) Department of Invertebrate Zoology, National Museum of Natural History, 
Smithsonian Institution, Washington, D.C. 20560-0118, U.S.A.; 
(YIK) A. N. Severtzov Institute of Problems of Evolution, Russian Academy of Sciences, 
Leninski Prospect 33, Moscow 117071, Russia; 
(KL) Zoologisches Institut und Zoologisches Museum der Universitat Hamburg, 
Martin-Luther-King-Platz 3, D-20146 Hamburg, Germany 


Abstract.—A new genus, Parabuccinum, is established for four bathyal Ma- 
gellanic species, of which one, Parabuccinum rauscherti, is described as new. 
The remaining three species were originally described in the Antarctic genus 
Chlanidota, from which they differ in the morphology of their protoconch, 
operculum, radula, stomach, and male reproductive system. Parabuccinum ap- 
pears to be most closely related to the monotypic circumAntarctic genus Neo- 
buccinum, but may be distinguished on the basis of shell and penis morphology, 
as well as by the size and shape of the gland of Leiblein. Parabuccinum is 
endemic to the Magellanic Province, and is the first record of the subfamily 
Bucinulinae in the malacofauna of this region. 


The genus Chlanidota Martens, 1878 had 
been considered to be one of the more di- 
verse and wide-ranging members of the an- 
tiboreal buccinoidean radiation. Dell (1990) 
included 13 species within this genus, some 
tentatively, among them four species that he 
described as new. A more recent revision 
(Harasewych & Kantor, 1999) divided the 
genus Chlanidota into two subgenera, 
Chlanidota sensu stricto, with five species, 
and Chlanidota (Pfefferia) containing three 
species. Five species were excluded from 
Chlanidota on the basis of newly obtained 
anatomical data as well as radular and oper- 
cular morphology. 

Three of the species described by Dell 
(1990) (1.e., Chlanidota bisculpta, C. eltan- 
ini, and C. polyspeira), all from the Magel- 
lanic Province, differ substantially from all 
species of Chlanidota. In this publication, 
we propose a new genus to include these 
Species, as well as an additional species 
more recently collected by R/V Polarstern 
and R/V Vidal Gomaz and described herein. 


A review of each of the species included in 
this new genus is provided. 


Materials and Methods 


This report is based primarily on material 
collected by the United States Antarctic 
Program (USAP), housed at the National 
Museum of Natural History, Smithsonian 
Institution (USNM), and consisting mostly 
of the type material for the three species 
described by Dell (1990). Additional sam- 
ples, including alcohol preserved anatomi- 
cal material, were collected at two stations 
of the R/V Polarstern and one station of the 
R/V Vidal Gomaz. These samples are main- 
tained in the collections of the Zoologisches 
Institut und Zoologisches Museum der 
Universitat Hamburg (ZMH). 

In the material examined sections, ““spec- 
imen’’ denotes live collected material, 
while ‘shell’? refers to records containing 
only the dead collected shells. 

Shell and operculum measurements were 
obtained for representative specimens of 


VOLUME 113, NUMBER 2 


each species (n = 10, when available), as 
detailed in Harasewych & Kantor (1999: 
fig. 1). Anatomical descriptions are based 
on gross dissections of preserved material. 
Radulae were removed by dissection, 
cleaned using diluted bleach (NaCI1Q), coat- 
ed with carbon and gold, and examined us- 
ing a Hitachi $570 Scanning Electron Mi- 
croscope (SEM). 

Photographs were obtained using a Leaf 
Lumina Digital Scanning Camera. Optical 
and SEM images were processed using 
Photoshop Version 5.02 (Adobe). 

The following abbreviations are used in 
the text: SL—shell length, AL—aperture 
length, BWL-—body whorl length, SW— 
shell width. 


Systematics 


Order Neogastropoda Wenz, 1938 
Superfamily Buccinoidea Rafinesque, 
1815 


Harasewych & Kantor (1999:256) pro- 
vided a brief overview of the current state 
of buccinoidean systematics and summa- 
rized several of the more prevalent classi- 
fications. Until the phylogenetic relation- 
ships of the higher taxa of Buccinoidea are 
reassessed on a global scale, we provision- 
ally continue to use the family Buccinulidae 
and its subdivisions, as defined by Powell 
(1951), to include the many distinctive aus- 
tral buccinoidean taxa. 


Family Buccinulidae Finlay, 1928 


According to Powell’s (1951:151) con- 
cept of Buccinulidae, the chief characteris- 
tics of the group are tricuspid rachidian 
teeth and an operculum with a terminal or 
subterminal nucleus. In our ongoing studies 
on Buccinoidea, we were able to identify 
several anatomical characters, among them 
the prevalance of fused salivary glands, and 
a simple stomach that lacks a posterior mix- 
ing area, that will likely serve to distinguish 
Buccinulidae from Buccinidae and other 
buccinoidean families. 


543 


Subfamily Buccinulinae Finlay, 1928 


Powell (1951) subdivided the family into 
three subfamilies, the Cominellinae, Buc- 
cinulinae and Prosiphiinae based on the 
morphology of the lateral teeth of the rad- 
ula. Cominellinae were characterized as 
having bicuspid lateral teeth, Buccinulinae 
were diagnosed by their tricuspid lateral 
teeth, while Prosiphiinae were recognized 
by their lateral teeth with a long basal pro- 
jection. However, taxa included in Prosi- 
phiinae vary considerably in their radular 
morphology, suggesting that resolution on 
a fine scale may be possible within this 
group. 

The radular teeth of the four species in- 
cluded in the new genus Parabuccinum all 
have tricuspid lateral teeth, indicating that 
this genus is referable to the subfamily Buc- 
cinulinae. 

The geographic range of the subfamily 
Buccinulinae spans Australia, New Zea- 
land, Antarctica, and the eastern Pacific 
coast as far north as California (based on 
Powell’s (1951) inclusion of the genus Kel- 
letia in Buccinulidae). Parabuccinum ex- 
tends that range into the Magellanic Prov- 
ince. 


Genus Parabuccinum, new genus 


Type species.—Chlanidota bisculpta 
Dell, 1996. 

Description.—Shell small for family, 
reaching 10.2—16.4 mm, depending on spe- 
cies. Shell relatively thick, solid, elongate 
or elongate-ovate in outline, spire high 
(about 0.33 to 0.5 SL). Spiral sculpture of 
thin to prominent low cords. Axial sculp- 
ture ranges from fine growth lines to prom- 
inent, sigmoidal, weakly prosocline ribs, 
depending on species. Aperture narrowly 
oval. Siphonal notch broad, slightly re- 
curved dorsally, abaxial margin may form 
ridge along fasciole. Periostracum thin, 
smooth or finely hirsute, covering most of 
Shell. Operculum large (0.53—0.60 AL), 
Ovate, paucispiral, with nucleus rotated 
clockwise, nearly 180° along long axis of 


544 


operculum. Proboscis of moderate length. 
Radular ribbon long, triserial. Rachidian 
tooth with 3 large cusps (central cusp larg- 
est) emerging from shallowly arched, 
straight-sided basal plate. Lateral teeth with 
3 cusps, outer cusp longest, middle cusp 
shortest, closely adjacent to innermost cusp. 
Salivary glands small, fused. Valve of Lei- 
blein well defined. Gland of Leiblein very 
small, tubular. Stomach broadly U-shaped, 
without caecum. Penis long, with flattened 
distal surface, bordered by thickened edge. 
Papilla long, cylindrical, situated in middle 
of distal surface, surrounded by very nar- 
row circular fold at its base. 

Remarks.—Conchologically, species of 
Parabuccinum somewhat resemble juve- 
nile specimens of Chlanidota, but differ in 
having shells that are more slender and 
have a large, invariably well preserved 
protoconch, a relatively tall spire, and a 
very thin periostracum. In contrast, the 
protoconch of Chlanidota remains un- 
known, despite the large number of spec- 
imens (including juveniles <10 mm SL) 
that were studied (Harasewych & Kantor, 
1999). Anatomically, Parabuccinum dif- 
fers from Chlanidota in having a propor- 
tionally larger, paucispiral operculum, a 
very long and thin-walled siphon, a very 
small gland of Leiblein, a penis with a flat- 
tened distal surface with a thickened edge, 
and a stomach that overlies a portion of the 
esophagus. 

There are some conchological similari- 
ties between this new genus and Buccinella 
jucunda (Thiele, 1912), which differs in 
having a prosiphiine radula (Thiele 1912: 
pl. 16, fig. 13) and an operculum with a 
terminal nucleus (pl. 13, fig. 20). Parabuc- 
cinum, especially P. eltanini, also resembles 
Neobuccinum eatoni (Smith, 1875) in gen- 
eral shell outline, operculum and _ proto- 
conch morphology. 

Parabuccinum is endemic to the Magel- 
lanic Province, and has a bathymetric range 
of 247 to 2165 m. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Parabuccinum bisculptum (Dell, 1990) 
Figs. 1, 2A—E, 3, 4, 7A—B, Table 1. 


Chlanidota bisculpta Dell, 1990:185, 
figs 291,294, 295,75 he. 

Description.—Shell (Fig. 1) large for ge- 
nus (to 14.8 mm), solid, white, elongate, 
with angulated shoulder. Protoconch (Fig. 7 
A-B) large (diameter 1.33—1.67 mm), of 
about 2.25 smooth, glossy, convex, slightly 
flattened whorls. Protoconch diameter/pro- 
toconch height = 1.02—1.09. Transition 
from protoconch to teleoconch well marked 
by onset of axial and spiral sculpture. Te- 
leoconch of up to 3.25 convex whorls. Su- 
ture impressed. Spiral sculpture of raised, 
distinct, narrow cords (12-16 on body 
whorl, 5—7 on penultimate whorl), half the 
width of intervening spaces. Axial sculp- 
ture of raised, narrow, weakly sinuate, op- 
isthocline ribs (18—22 on first teleoconch 
whorl, 16—24 on subsequent whorls) that 
form nodules at intersections with spiral 
cords. Aperture large (0.51—0.65 SL), nar- 
row, ovate, deflected from shell axis by 15— 
19°. Outer lip, simple, rounded anteriorly, 
shouldered posteriorly. Columella ~0.5 to 
0.67 AL, weakly concave, with strong si- 
phonal fold. Callus of thick glaze overlying 
parietal region, siphonal fasciole. Siphonal 
notch narrow, slightly dorsally recurved, 
with nearly straight columellar and rounded 
apertural margins that define borders of fas- 
ciole. Ridge margin of fasciole formed by 
apertural margin of siphonal notch. Shell 
color chalky white, aperture weakly glazed. 
Periostracum thin, transluscent, light yel- 
lowish, thinly lallemose, with fine perios- 
tracal hairs produced at intersections of spi- 
ral threads with axial growth lines. Oper- 
culum (Fig. 1E) medium-sized (0.53 AL), 
elongate ovate, strongly coiled, with nucle- 
us rotated clockwise, nearly 180° to long 
axis of operculum. 

Anatomical data based on single imma- 
ture, male specimen (Fig. 1 I, R/V Polar- 
stern, Sta. 10-109), from which only a por- 
tion of the animal was recovered. 

External anatomy.—(Fig. 2 A—B). Man- 


VOLUME 113, NUMBER 2 


Fig. 1. Parabuccinum bisculptum (Dell, 1990). A—C. Holotype, USNM 860128, off Burdwood Bank, 
53°08'S, 59°23’'W, in 567-578 m (R/V Eltanin, Sta. 340). D—E. Paratype 1, USNM 860129, off Falkland (Mal- 
vinas) Islands, 51°56’'S, 56°39’ W, 855-866 m (R/V Eltanin, Sta. 557). D. Apertural view of shell. E. Operculum 
(coated with ammonium chloride). E Paratype 2, USNM 860129. Same locality as paratype 1. G—I. Off Cape 
Horn, 55°44.0’S, 66°14.5’W, 430-397 m (R/V Polarstern, Sta. 40-109; G, H. ZMH, I. USNM 892150). Scale 
bar = 5 mm for shells, 2 mm for operculum. 


546 


tle cavity spans just over 0.5 whorl, kidney 
(Fig. 2A, k) % whorl. Upper whorls of vis- 
ceral mass unknown. Columellar muscle 
comprises 1.5 whorl, thick and narrow, at- 
taching to shell at rear of mantle cavity. 
Foot large, oval, broader anteriorly. Body 
color very light yellow, nearly white, with- 
out pigmentation. Head small, tentacles 
(Fig. 2 A, B, ten) very long, narrow, widely 
separated, gradually tapering. Eyes absent. 
Siphon (Fig. 2 A, B, s) very long (0.75 
AL), thin-walled, open. 

Alimentary system.—Proboscis (Fig. 2 
C—D, pr) of moderate length when contract- 
ed (0.44 SL, 0.73 AL), smooth, unpig- 
mented. Proboscis sheath thin-walled, 
transparent. Mouth (Fig. 2 C, m) triangular 
slit. Proboscis retractors (Fig. 2 C, D, prr) 
broad, extremely thin, attached to middle 
part of rhynchodaeum on left side, to its 
base on right side. Buccal mass spans 
~=().67 proboscis length. Radular ribbon 
(Fig. 3) 1.55 mm long (0.39 AL), ~100— 
140 pm wide (~0.015 SL, 0.025 AL), tris- 
erial, consisting of about 60 rows of teeth. 
Rachidian teeth with 3 closely spaced cusps 
(central cusp slightly longer, wider than lat- 
eral cusps) on posterior portion of basal 
plate. Anterior margin of basal plate not 
thickened, overlaid by adjacent tooth. Lat- 
eral teeth usually with 3 cusps. Outer cusp 
roughly twice as long, half as wide as inner 
cusp. Intermediate cusp very thin, adjacent 
to, equal in length to inner cusp. In one 
specimen (Fig. 3 A—B) intermediate cusp 
split into two separate cusps of equal size 
along portion of radular ribbon. Salivary 
glands (Fig. 2 D, sg) small, fused, situated 
above nerve ring along left side of probos- 
cis. Valve of Leiblein (Fig. 2 C, vL) well 
defined, although not large, pyriform. 
Gland of Leiblein (Fig. 2 C, D, gL) very 
small, short, tubular, uncoiled, whitish, 
opening into oesophagus without constric- 
tion, just posterior to nerve ring (Fig. 2 C, 
nr). Oesophagus narrow, thin-walled, 
broader posteriorly. Stomach (Fig. 2 E, st) 
broad, simple, U-shaped, without caecum. 
Stomach simple, broad, overlies posterior- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


most oesophagus. Internal structures of 
stomach not well-preserved. 

Male reproductive system.—Specimen 
immature male, with very short, dorso-ven- 
trally flattened penis (Fig. 2B, p) lacking 
defined papilla. 

Type locality.—Off Burdwood Bank, 
53°08'S, 59°23’'W, in 567-578 m (R/V El- 
tanin, Sta. 340). 

Type material.—Holotype (Fig. 1 A—C), 
USNM 860128, from the type locality; Par- 
atypes 1-6, USNM 860129, Paratype 7, 
NMNZ ME .56615, off Falkland (Malvin- 
as) Islands, 51°56’S, 56°39'W, 855-866 m 
(R/V Eltanin, Sta. 557); Paratypes 8—9 
(Paratype 9 is a specimen of Parabuccinum 
rauscherti, new species, see below), USNM 
860130, Paratype 10, NMNZ MEF .56616, 
off Cape Horn, 56°06'S, 66°19’W, 384-349 
m (R/V Eltanin, Sta. 740). 

Material examined.—Type material in 
USNM. 1 specimen + 1 shell, USNM 
892150, 5 shells ZMH, off Cape Horn, 
55°44.0'S, 66°14.5'W, 430-397 m, 16 May, 
1996 (R/V Polarstern ANT XIII/4. Sta. 40— 
109). 

Distribution.—Off the Falkland (Malvin- 
as) Islands and Cape Horn at depths ranging 
from 349 to 866 m (Fig. 4). 

Remarks.—This species most closely re- 
sembles Parabuccinum polyspeirum and P. 
rauscherti, new species. Parabuccinum bis- 
culptum may be distinguished from P. poly- 
speirum by its slightly smaller protoconch 
(1.33—1.67 vs. 1.52—1.84 mm diameter), its 
strongly shouldered rather than rounded 
whorl profile, as well as by the presence of 
fewer, stronger spiral cords and more pro- 
nounced axial sculpture in the former (com- 
pare Tables 1, 4). Characters that may be 
used to differentiate P. bisculptum from P. 
rauscherti, new species, are discussed in the 
description of the new species. 


Parabuccinum polyspeirum (Dell, 1990) 
Figs. 4, 5, 7 E-E Table 2. 


Chlanidota polyspeira Dell, 1990:186, 
figs: 292.293.3113; 


VOLUME 113, NUMBER 2 547 


Fig. 2. Anatomy of Parabuccinum bisculptum (A-E, USNM 892150) and P. rauscherti (F—G, USNM 
880616). A, B. Body removed from the shell. A. Right view. B. Anterior view of foot-head. C. Right lateral D. 
Left lateral, views of anterior alimentary system (salivary glands removed to show valve of Leiblein in C). E. 
Lateral view of stomach. K Mantle complex of organs (partially preserved, missing part is indicated by dotted 
line). G. Anterior view of the body, mantle removed. Abbreviations: cme, cut mantle edge; ct, ctenidium; dg, 
digestive gland; gL, gland of Leiblein; h, heart; k, kidney; m, mouth; nr, circumoesophageal nerve ring; oe, 
oesophagus; op, operculum; os, osphradium; p, proboscis; poe, posterior oesophagus; prp, propodium; prr, pro- 
boscis retractors; s, siphon; sd, salivary duct; sg, salivary gland; st, stomach; ten, cephalic tentacles; vL, valve 
of Leiblein. 


Description.—Shell (Fig. 5) large for ge- 
nus (to 15.4 mm) solid, white, glossy, elon- 
gate, with rounded shoulder. Protoconch 
(Figs. 7 E-F) large (diameter 1.52—1.84 
mm), consists of ~2.5 smooth, glossy, con- 


vex, raised whorls. Protoconch diameter/ 
protoconch height = 1.12—1.28. Traces of 
weak spiral cords may be present in final 
0.25 whorl of protoconch. Transition from 
protoconch to teleoconch abrupt, marked by 


548 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Table 1— Parabuccinum bisculptum (Dell, 1990). Measurements of shell characters. Linear measurements 


in mm (nv = 9, including holotype). 


Character Mean 


Shell Length (SL) 9.9 

Body Whorl Length (BWL) 76 

Aperture Length (AL) 5.6 

Shell Width (SW) 5.4 

Protoconch diameter 1.53 

BWL/SL 0.76 

AL/SL 0.56 

SW/SL 0.54 

Number of axial ribs on Ist teleo- 19.5 
conch whorl 

Number of axial ribs on 2nd teleo- 19.7 
conch whorl 

Number of axial ribs on body 20.4 
whorl 

Number of spiral cords on body L357 
whorl 

Number of spiral cords on penul- 6 


timate whorl 


onset of pronounced spiral cords, followed 
within 0.5 whorl by first, weak axial ribs. 
Teleoconch of up to 3.25 convex whorls. 
Suture impressed. Spiral sculpture of nu- 
merous (19-26 on body whorl, 8—14 on 
penultimate whorl) closely-spaced, raised 
cords. Axial sculpture of raised, narrow, 
sinuate, weakly prosocline ribs (16—18 on 
first teleoconch whorl, increasing to 19-21 
on body whorl) that form faintly cancellate 
sculpture at intersection with spiral cords. 
Aperture large (0.53—0.63 SL), narrow, 
ovate, deflected from shell axis by 13-17°. 
Outer lip thin, evenly rounded. Columella 
=0.5 AL, weakly concave, with strong si- 
phonal fold. Callus of thin glaze (very thin 
in smaller specimens) overlying parietal re- 
gion, siphonal fasciole. Siphonal notch nar- 
row, slightly dorsally recurved, with nearly 
straight columellar and rounded apertural 
margins that form borders of fasciole. Ap- 
ertural margin of siphonal notch demarcates 
ridge margin of fasciole. Shell color chalky 
white, aperture weakly glazed. Periostrac- 
um very thin, smooth, translucent, tightly 
adherent to shell surface. Operculum un- 
known. 

Type locality.—Patagonian Shelf, NE of 


o Range Holotype 
1.56 6.6-14.8 14.8 
P33 5.0—11.6 11.6 
1.08 4.0-8.9 8.9 
0.83 3.9-8.2 8.2 
0.12 1733—1-67 153 
0.03 0.74—0.81 0.78 
0.05 0.51—0.65 0.60 
0.03 0.51—0.59 0.55 
ies? 18—22 — 
22535 16—24 19 
3:26 16—24 2S 
era 12-16 13 
0.53 5-7 6 

Islas de los Estados (Staten Island), 


54°04'S, 63°35’W, in 247-293 m (R/V EI- 
tanin, Sta. 369). 

Type material.—Holotype (Figs. 5, A-— 
C), USNM 860131, 3 paratypes, USNM 
860132, 1 paratype NMNZ ME56617, all 
from the type locality. 

Material examined.—Type material in 
USNM. 

Distribution.—Known only from the 
type locality (Fig. 4). 

Remarks.—The species is known from 
five shells. Dell (1990) regarded this taxon 
to be very similar to P. bisculptum, but dis- 
tinguished it on the basis of its narrower 
Shell, larger protoconch, more numerous 
spiral cords and axial sculpture that is 
weaker, and becomes obsolete on the anter- 
iormost part of the body whorl. 

Parabuccinum polyspeirum is known 
from a single station that represents the 
shallowest record for any species of Para- 
buccinum. 


Parabuccinum eltanini (Dell, 1990) 
Figs. 6,.8, 12, Table. 3. 


Chlanidota eltanini Dell, 1990:184-5, figs. 
290, 292, 297, 314. 


VOLUME 113, NUMBER 2 


i j a my 
» of ter’: ie ‘ = 
f “ wr ™ a 
Gt Oe Sect . i» i 5 > oe = 
BR Ss a See 4 , 
“a “ ; . ‘ \ 
Aish x we Sf”. Loe 
fey " ; Se oe og 
ns? A 
oe 


Radulae of Parabuccinum bisculptum (Dell, 1990). A. Dorsal, and B. right lateral (45°) views of the 
central portion of the radular ribbon taken from animal in Fig. 11 (USNM 892150). C, D. Dorsal views of the 
radulae of two specimens from R/V Polarstern, Sta. 40-109. Scale bars = 50 ym. 


Fig. 3. 


Description.—Shell (Fig. 6) largest of 
genus (to 16.4 mm), solid, white, ovate, 
with lower spire, rounded shoulder. Proto- 
conch (Fig. 8) very large (diameter 2.06— 
2.94 mm), dome-shaped, of about ~2.25 
smooth, low, whorls. Protoconch diameter/ 
protoconch height = 1.30—1.52. Border be- 
tween protoconch and teleoconch whorls 
well demarcated, protoconch well pre- 
served, teleoconch eroded. Teleoconch of 
up to 2.75 broadly convex whorls. Suture 
deeply impressed, with narrow, nearly flat- 


tened rim. Spiral sculpture of numerous 
(32-53 on body whorl, 17—24 on penulti- 
mate whorl), adjacent, very low, fine, nar- 
row, sometimes sinuous threads, covering 
entire shell surface. Axial sculpture limited 
to fine prosocline growth lines. Aperture 
large (0.65 SL), ovate to broadly ovate, de- 
flected from shell axis by 10—14°. Outer lip 
simple, evenly rounded. Columella <0.5 
AL, weakly concave, with broad siphonal 
fold. Callus consisting of thick glaze nar- 
rowly overlying parietal region, siphonal 


550 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


200 


Depth, m 


500 


Parabuccinum bisculptum (Dell, 1990) 
“9 = type locality © = material examined 


Parabuccinum polyspeirum (Dell, 1990) 


+ = type locality 


Fig. 4. 


700 


900 


[234356 3 
No. of specimens 


Geographical distribution and bathymetric ranges of Parabuccinum bisculptum (Dell, 1990) and 


Parabuccinum polyspeirum (Dell, 1990). Dashed line indicate 500 m isobath. The bathymetric range of P. 
bisculptum is shown in black, that of P. polyspeirum in gray. 


fasciole. Siphonal notch narrow, shallow, 
slightly dorsally recurved, its margins de- 
fining weak fasciole. Ridge margin of fas- 
ciole runs from apertural margin of siphon- 
al notch. Shell color chalky white, aperture 
glazed. Periostracum thin, yellowish, tightly 
adhering to the shell surface, bearing irreg- 
ularly spaced very short hairs. Operculum 
(Fig. 6H) large (0.60 AL), broadly ovate, 


strongly coiled, with nucleus rotated clock- 
wise, nearly 180° to long axis of operculum. 
Radula illustrated by Dell (1990:fig. 297). 

Type locality.—East of Falkland (Mal- 
vinas) Islands, 51°58’S, 56°38’W, in 845— 
646 m (R/V Eltanin, Sta. 558). 

Type material.—Holotype (Fig. 6, A—C), 
USNM 860124, paratypes 1-2, USNM 
860125, paratype 3, NMNZ ME56613, 


Table 2.—Parabuccinum polyspeirum (Dell, 1990). Measurements of shell characters. Linear measurements 


in mm. 


Character 


Shell Length (SL) 

Body Whorl Length (BWL) 

Aperture Length (AL) 

Shell Width (SW) 

Protoconch diameter 

BWL/SL 

AL/SE 

SW/SL 

Number of axial ribs on Ist teleoconch whorl 
Number of axial ribs on 2nd teleoconch whorl 
Number of axial ribs on body whorl 

Number of spiral cords on body whorl 
Number of spiral cords on penultimate whorl 


Holotype Paratype 1 Paratype 2 Paratype 3 
JI) 92 15.4 oN 8.1 
10.3 j bile fhe: 6.4 
72 8.2 =a | 5.0 
6s Fak 4.8 4.7 
1.84 1.84 1.75 152 
0.78 0.75 0.80 O79, 
0.55 0.53 0.63 0.62 
Ors2 0.50 0.53 0.58 

16 16 17 18 

22 18 24 2A 

21 20 19 19 

26 24 22 19 

14 11 8 8 


VOLUME 113, NUMBER 2 


SAI 
Nn 


Fie: 5. 
Patagonian Shelf, NE of Islas de los Estados (Staten Island), 54°04’S, 63°35'W, in 247-293 m D. Paratype 1. 
E. Paratype 2. Both from type locality. Scale bar = 5 mm. 


from the type locality. Paratypes 4—5 [par- 
atype 5 (SL 4.8 mm) is not a buccinoidean, 
but belongs in the family Cancellariidae 
(Admetinae)] USNM 860126, E off Falk- 
land (Malvinas) Islands, 51°56’S, 56°39’W, 
855-866 m (R/V Eltanin, Sta. 557). Para- 


Parabuccinum polyspeirum (Dell, 1990). A-—C. Holotype, USNM 860131, R/V Eltanin, Sta. 369, 


types 6-7, USNM 860127, paratype 8, 
NMNZ ME56614, E off Falkland (Malvin- 
as) Islands, 54°09’S, 52°08’W, 419-483 m 
(R/V Eltanin, Sta..1521): 

Material examined.—Type material in 
USNM. 


Table 3.—Parabuccinum eltanini (Dell, 1990). Measurements of shell characters of intact specimens. Linear 


measurements in mm. * = heavily eroded. 


Character 


Shell Length (SL) 

Body Whorl Length (BWL) 

Aperture Length (AL) 

Shell Width (SW) 

Protoconch diameter 

BWL/SL 

AL/SL 

SW/SL 

Number of spiral cords on body whorl 
Number of spiral cords on penultimate whorl 


Holotype Paratype Paratype 
USNM USNM USNM 
860124 860127 860125 
135 16.4 15.8 
113 13.6 13.0 

8.8 10.3 = 
8.8 10.0 9.6 
2.06* 2.94 2.24* 
0.84 0.83 0.80 
0.65 0.63 0.63 
0.65 0.61 0.61 

BY 33 38 
i 24 20 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 6. 
E of Falkland (Malvinas) Islands, 54°09’S, 52°08'W, 419-483 m (R/V Eltanin, Sta. 1521). D. Apertural view. 
E. Dorsal view of the protoconch. EF Lateral view of the protoconch showing transition between protoconch and 
teleoconch whorls. G-H. Paratype, USNM 860125, type locality, E off Falkland (Malvinas) Islands, 51°58’S, 
56°38'W, 845-646 m (R/V Eltanin, Sta. 558). G. Apertural view. H. Operculum (coated with ammonium chlo- 


Parabuccinum eltanini (Dell, 1990). A-C. Holotype, USNM 860124. D—-F Paratype, USNM 860127, 


ride). Scale bars = 1 cm for shells, 5 mm for protoconch, 2 mm for operculum. 


Distribution.—Off Falkland (Malvinas) 
Islands. Specimens collected at depths from 
419 to 866 m. (Fig. 12). 

Remarks.—This species differs from all 
congeners in having a smoothly ovate shell 
with a tall body whorl, a strongly prosoc- 
line outer lip, an extremely large, dome- 
shaped protoconch, and by the absence of 


axial sculpture. Dell’s (1990:fig. 297) line 
drawing of the radula of P. eltanini shows 
a lateral tooth with a narrow basal plate and 
inner cusp, which differs from those of oth- 
er Parabuccinum (Figs. 3, 10) and some- 
what resembles those of Chlanidota (see, 
eg., Harasewych & Kantor, 1999 figs. 10 A, 
C). Although we were unable to study the 


VOLUME 113, NUMBER 2 


anatomy of P. eltanini, we assign it to the 
genus Parabuccinum on the basis of the 
morphology of its shell, especially the char- 
acteristic protoconch, and operculum. 

Paratype 5, the smaller (SL = 4.8 mm) 
of the two paratypes from R/V Eltanin sta- 
tion 557 (USNM 860126), is not conspe- 
cific with the rest of the type series, but is 
a species referable to the cancellariid sub- 
family Admetinae. 


Parabuccinum rauscherti, new species 
Figs. 2F—G, 7C—D, 9-12, Table 4. 


Chlanidota bisculpta Dell, 1990 (par- 
tim):185. 

Description.—Shell (Fig. 9) small for ge- 
nus (to 10.2 mm), solid, white, broadly 
ovate to elongate, with flattened, shallow 
subsutural rim, pronounced, rounded to an- 
gulated shoulder. Protoconch (Fig. 7 C—D) 
small (diameter 1.07—1.2 mm), of about 
1.75 glossy, convex, inflated whorls. Pro- 
toconch diameter/protoconch height = 
1.10—1.31. Transition from protoconch to 
teleoconch well marked by onset of spiral 
followed immediately by axial sculpture. 
First teleoconch whorl thinner than proto- 
conch. Teleoconch of up to 3.25 convex 
whorls. Suture strongly impressed. Spiral 
sculpture of closely spaced cords varying in 
strength from strongly pronounced to near- 
ly smooth (16—40 on body whorl, 10—16 on 
penultimate whorl), generally broader than 
intervening spaces. Axial sculpture of 
strong, broad, slightly sinuous, orthocline 
ribs (13-17 on first teleoconch whorl, 15— 
20 on subsequent whorls). Nodules formed 
at intersections with spiral cords evident in 
early whorls, abbraded on body whorl. Ap- 
erture large (0.51—0.65 SL), narrow to mod- 
erately ovate, deflected from shell axis by 
12—15°. Outer lip evenly rounded, usually 
slightly thickened. Columella <0.5 AL, 
weakly concave, with strong, sharply de- 
flected siphonal fold. Callus of thin to thick 
glaze narrowly overlying parietal region 
and siphonal fasciole. Siphonal notch mod- 
erately broad, very slightly dorsally re- 


ae 


curved, with straight columellar, rounded 
apertural margins defining borders of fas- 
ciole. Ridge margin of fasciole formed by 
apertural margin of siphonal notch. Shell 
color chalky white, aperture weakly glazed. 
Periostracum very thin, tightly adherant, 
yellowish, weakly hirsute. Operculum (Fig. 
9D, G) medium-sized (0.49—0.53 AL), 
elongate ovate, strongly coiled, with nucle- 
us rotated clockwise, nearly 180° to long 
axis of operculum. 

Anatomical data based on single, poorly 
preserved male specimen (Fig. 9 E—E Par- 
atype 1) from which only a portion of the 
animal was recovered. 

External anatomy.—(Fig. 2F—G). Foot 
long (L/W ~2.5), oval, broad anteriorly, ta- 
pering posteriorly, with thickened propo- 
dium. Body color uniformly yellowish- 
white. Head small short, tentacles (Fig. 3G, 
ten) long, left longer than right, tapering 
distally. Eyes absent. Mantle cavity (Fig. 
3F) as wide as long. Mantle edge thickened, 
with scalloped edge. Siphon (Fig. 3F) very 
long (0.9 AL), thin walled, wide. Osphra- 
dium ~0.67 mantle length, wide, with 
curved axis. Ctenidium slighter longer, 0.67 
as wide as osphradium, spans about 0.75 of 
mantle length, formed of tall triangular la- 
mellae that become narrower toward mantle 
edge. 

Alimentary system.—Proboscis of mod- 
erate length in contracted position (0.35 SL, 
0.6 AL), smooth, unpigmented. Proboscis 
sheath very thin-walled, transparent. Mouth 
opening triangular. Proboscis retractors 
broad, extremely thin, asymmetrically at- 
tached to proboscis sheath. Buccal mass oc- 
cupies ~0.63 length of retracted proboscis. 
Radula (Fig. 10) 1.54 mm long (0.33 AL), 
~120-140 wm wide (~0.015-0.017 SL, 
0.026—0.030 AL), of 65 rows of teeth, pos- 
teriormost 3 nascent. Rachidian teeth tri- 
cuspid (central cusp slightly longer, as wide 
as lateral cusps). Lateral teeth usually with 
3 cusps. Outer cusp nearly twice as long, 
half as wide as inner cusp. Intermediate 
cusp thin, equal in length to inner cusp. In 
one specimen (Fig. 10 C) intermediate cusp 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


es 


S205 
Ss 


Fig. 7. Protoconchs of Parabuccinum. A-B. P. bisculptum (Dell, 1990), ZMH. C—D. P. rauscherti, new 
species. Paratype, ZMH 2813. E-E P. polyspeirum (Dell, 1990) Paratype in fig. 5 E. Arrows indicate transition 
from protoconch to teleoconch. Scale bar = 500 wm for all figures. 


VOLUME 113, NUMBER 2 Bi J 


Table 4.—Parabuccinum rauscherti, new species. Measurements of shell characters. Linear measurements in 
mm (n = 12, including types). 


Paratype Paratype 
Character Mean o Range Holotype ] 4 
Shell Length (SL) 8.4 0.67 7.6—9.6 ist 8.1 8.1 
Body Whorl Length (BWL) 6.5 0.63 5.4-7.4 6.1 6.3 6.2 
Aperture Length (AL) 4.7 0.49 4.2-5.5 4.2 4.7 4.6 
Shell Width (SW) 0 0.44 4.1-5.8 4.6 4.7 4.9 
Protoconch diameter Nis 0.05 1.07—1.20 1.07 1.18 1.13 
BWL/SL 0.76 0.03 0.71—0.80 0.79 0.78 0.77 
AL/SL 0.56 0.05 0.51—0.65 0.55 0.58 0.57 
SW/SL 0.60 0.04 0.54—0.67 0.59 0.58 0.60 
Number of axial ribs on Ist teleoconch whorl 15.0 15 13-17 13 15 14 
Number of axial ribs on 2nd teleoconch whorl 16.5 Ee 15-20 16 17 16 
Number of axial ribs on body whorl 77 1.95 15-20 16 17 17 
Number of spiral cords on body whorl 25.8 6.88 16—40 25 17 20 
Number of spiral cords on penultimate whorl jaa | 2.28 10-16 12 10 10 


fused with inner cusp along portion of rad- 
ular ribbon. Valve of Leiblein well defined, 
medium-sized, pyriform. Salivary ducts 
join oesophagus just anterior to the valve. 
Remainder of alimentary system poorly 
preserved. 

Male reproductive system.—Specimen 
mature male, with long, broad penis (Fig. 
11) extending length of mantle length. Dis- 
tal end of penis transversely flattened, bor- 
dered by thickened edge. Penial papilla 
(Fig. 11B, pap) long, cylindrical, surround- 
ed by narrow circular fold at the base. 


Type locality.—Off Cape Horn, 55°44.0’S, 
66°14.5'W, in 430-397 m. (R/V Polarstern, 
ANT XIII/4. Sta. 40-109). 

Type material.—Holotype (ZMH 2811), 
paratypes 1—3 (USNM 880616), paratype 4 
(ZMH 2812), paratypes 5—20 (ZMH 2813), 
paratypes 21-30 (ZMH 28134), from the 
type locality. 

Material examined.—Type material. R/V 
Eltanin. Sta. 339: Falkland (Malvinas) Is- 
lands, Beauchene Island, 53°05’S, 59°31’'W, 
512—586 m, 1 shell, USNM 870141; Sta. 
740, off Cape Horn, 56°06’S, 66°19'W, 


Fig. 8. 
USNM 860127, E of Falkland (Malvinas) Islands, 54°09'S, 52°08’W, 419—483 m (R/V Eltanin, Sta. 1521). See 
also Fig. 6 E-F Arrows indicate transition from protoconch to teleoconch. Scale bar = 1 mm. 


Protoconch of Parabuccinum eltanini (Dell, 1990) at half the magnification of figure 7. Paratype, 


556 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 9. Parabuccinum rauscherti, new species. A-D. Holotype, ZMH 2811, R/V Polarstern ANT XIII/4, 
Sta. 40-109, off Cape Horn, 55°44.0'S, 66°14.5’W, in 430-397 m. A-C. Shell. D. Operculum. E—G. Paratype 
1, type locality, USNM 880616. E—-F Shell. G. Operculum. H. Paratype 4, ZMH 2812, type locality. I-J. R/V 
Polarstern Sta. 40-114, off Cape Horn, 55°31.6'S, 65°56.8'W, 2165-2008 m, ZMH. Scale bar = 5 mm for 
shells, 2 mm for operculum. 


VOLUME 113, NUMBER 2 


ro | 


Fig. 10. Radula of Parabuccinum rauscherti, new species. A. Dorsal, and B. left lateral (45°) views of the 
central portion of the radular ribbon taken from paratype 1, USNM 880616 (Figs. 9, E—F). C. Dorsal and D. 
left lateral (45°) views of the central portion of the radular ribbon taken from paratype 4, ZMH 2812 (Figs. 9 
G). Scale bars = 50 pm. 


384-349 m, | shell (paratype of Chlanidota 
bisculpta), USNM 860130. R/V Polarstern 
ANT XIII/4: Sta. 40-114, off Cape Horn, 
55°31.6'S, 65°56.8’W, 2165-2008 m, 18 
May, 1996, ZMH, 29 shells. R/V Vidal 
Gomaz: Sta. 42, Canal Concepcion, 
50°35.7'S, 75°04.5'W, 250 m, 25 Oct 1996, 
ZMH, 5 specimens. 

Distribution.—Off the Falkland (Malvin- 
as) Islands, Cape Horn and northward along 
western coast of Chile to Canal Concep- 
cion. Shells were trawled at depths ranging 


from 349 to 2165 m. Live animals were col- 
lected from 349 to 532 m (Fig. 12). 
Etymology.—This new species is named 
after Martin Rauschert who developed the 
small dredge used to collect this species. Dr. 
Rauschert is an amphipod taxonomist who 
has worked for several years in the Antarc- 
tic and Magellanic regions. 
Remarks.—Parabuccinum rauscherti is 
most similar in shell sculpture structure to 
P. bisculptum from which it differs in hav- 
ing a smaller, slightly broader shell (SW/SL 


558 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 11. 


Parabuccinum rauscherti, new species. Critical point dried penis of paratype 1, USNM 880616 


(Figs. 9, E-F). A. Dorsal surface. Scale bar = 500 wm. B. Distal tip showing papilla. Scale bar = 200 pm. 


=().60 in P. rauscherti, 0.54 in P. bisculp- 
tum), a smaller protoconch (average diam- 
eter = 1.16 mminP. rauscherti, 1.53 in P. 
bisculptum) (compare Fig. 7A, B and 7C, 
D). Parabuccinum rauscherti also has more 
numerous and closely spaced spiral cords 


(average = 25.9 on body whorl, 12.1 on 
penultimate whorl) than P. bisculptum (av- 
erage = 13.8 on body whorl, 6.0 on pen- 
ultimate whorl) and less numerous (16.5 vs. 
19.8) but wider axial ribs. Parabuccinum 
rauscherti and P. bisculptum were taken to- 


300 
400 
500 
600 


700 


Parabuccinum eltanini (Dell, 1990) 
@) = type locality 


Parabuccinum rauscherti, new species 
© =type locality 


Fig. 12. 


@ = material examined 


@ = material examined 


800 


900 


12:34 
No. of specimens 


B 


2000 


Depth, m 


A 


510° 20730 
No. of specimens 


Geographical distribution and bathymetric ranges of Parabuccinum eltanini (Dell, 1990) and Par- 


abuccinum rauscherti, new species. Dashed line indicates 500 m isobath. A. Bathymetric range of P. rauscherti. 


B. Bathymetric range of P. eltanini. 


VOLUME 113, NUMBER 2 


gether at two stations off Cape Horn (R/V 
Polarstern Sta. 10-109 and R/V _ Eltanin 
Sta. 740). 

Parabuccinum rauscherti may be most 
readily distinguished from either P. poly- 
speirum or P. eltanini by its much smaller 
protoconch and pronounced axial sculpture. 
Parabuccinum rauscherti exhibits some 
variation in shell outline and the number of 
spiral cords. 

The smaller of the two paratypes of 
Chlanidota bisculpta, from R/V_ Eltanin. 
Sta. 740 (USNM 860130, SL = 6.9 mm) is 
actually a specimen of Parabuccinum raus- 
cherti. 


Discussion 


Despite the limited availability of the an- 
atomical data for Parabuccinum, it clearly 
shares anatomical features with other Buc- 
cinulinae, such as Chlanidota, Cavineptu- 
nea and several yet to be named genera. 
These similarities include proboscis mor- 
phology, tricuspid rachidian and lateral rad- 
ular teeth, a well defined valve of Leiblein, 
fused salivary glands, a characteristic crop- 
like structure of the posterior oesophagus, 
and a broadly U-shaped stomach, lacking a 
caecum (posterior mixing area). This stom- 
ach morphology was found in all antiboreal 
Buccinoidea studied thus far. Among north- 
ern hemisphere buccinoideans, a stomach 
lacking a caecum was recorded only in the 
northern Atlantic buccinid Colus gracilis 
(DaCosta, 1778) (see Smith 1967) and 
deep-water Arctic species Mohnia (Tacita) 
danielsseni (Friele, 1879) (see Lus 1981). 

Parabuccinum has a number of unusual 
features, among them a very long mantle 
siphon, a very small, nearly vestigial gland 
of Leiblein, and characteristic penis mor- 
phology previously unrecorded in Bucci- 
noidea. 

With the exception of Parabuccinum, the 
Magellanic buccinoidean fauna is com- 
prised of genera attributed to the buccinulid 
subfamily Cominellinae (e.g., Pareuthria, 
Falsitromina, Parficulina). As there are no 


559 


closely related South American or Magel- 
lanic Buccinoidea, Parabuccinum is likely 
derived from the Antarctic Buccinulinae. 
Powell (1951) attributed the following 
Antarctic genera to the subfamily Buccin- 
ulinae: Chlanidota, Pfefferia (a subgenus of 
Chlanidota, as demonstrated by Harase- 
wych & Kantor 1999), Neobuccinum, Prob- 
uccinum, Cavineptunea and Bathydomus. 
The differences between Parabuccinum 
and Chlanidota have been discussed above. 
Of the remaining Antarctic buccinuline 
taxa, only one—the monotypic genus Neo- 
buccinum, has a similar paucispiral oper- 
culum. The shell morphology of Neobuc- 
cinum eatoni (Smith, 1907) is similar to 
that of Parabuccinum, especially to that of 
P. eltanini. Both Neobuccinum and Para- 
buccinum have large, dome-shaped proto- 
conchs that appear to be more resistant to 
errosion that their teleoconchs. The radula 
of N. eatoni (illustrated by Numanami, 
1996:fig. 94C) resembles that of P. eltanini 
more that those of the other species of 
Parabuccinum in that the central cusps of 
the lateral teeth are larger and more distant 
from the inner cusps. Parabuccinum and 
Neobuccinum may be distinguished ana- 
tomically, particularly on the basis of penis 
morphology and the size and shape of gland 
of Leiblein. The penis of Neobuccinum has 
a broadly rounded distal end with a very 
small, blunt seminal papilla, while the 
gland of Leiblein in this species is large and 
well developed (unpublished observations). 
Neobuccinum eatoni has a circumantarc- 
tic distribution extending along the Scotia 
Arc, and to Kerguelen Island, as well as a 
broad bathymetric range (5 to 2350 m). De- 
spite the anatomical differences, we consid- 
er Neobuccinum to be the most promising 
candidate for sister group of Parabuccinum 
among the presently known Antarctic buc- 
cinoidean fauna. Parabuccinum thus rep- 
resents an Antarctic component in the Ma- 
gellanic malacofauna. 
The distinctive paucispiral opercula of 
Parabuccinum and Neobuccinum differ 
from those of other austral Buccinoidea, yet 


560 


resemble opercula of the deep-water boreal 
genus Mohnia (termed Mohnia type oper- 
culum by Bouchet and Warén 1985:171). 
Several species of Mohnia also have tricus- 
pid rachidian and lateral teeth (Bouchet & 
Warén 1985:178—-179) while at least one 
species [Mohnia (Tacita) danielsseni, see 
above] has a stomach lacking a caecum. 
Mohnia is readily distinguished from Par- 
abuccinum and Neobuccinum by numerous 
conchological (e.g., presence of siphonal 
canal) and radular (rachidian teeth with 
squarish basal plate and lateral cusps lost or 
closely juxtaposed to prominent central 
cusp) characters. It 1s, as yet, unclear 
whether these similarities are indicative of 
a close relationship between these antipodal 
taxa, or represent plesiomorphic characters 
within the Buccinoidea. 


Acknowledgments 


We thank Dr. James McLean for his help- 
ful comments on the manuscript. YIK 
gratefully acknowledges Mrs. Mabs Mango 
and Mr. and Mrs. Ed George for their kind 
hospitality during the preparation of this 
manuscript. 

This research was supported in part by a 
grant to YIK from the USAP Research Pro- 
gram Award (Biological Collections from 
Polar Regions at the National Museum of 
National History) [National Science Foun- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


dation (Office of Polar Programs) Cooper- 
ative Agreement OPP-9509761]. 


Literature Cited 


Bouchet, P., & A. Warén. 1985. Revision of the north- 
east Atlantic bathyal and abyssal Neogastropo- 
da excluding Turridae (Mollusca, Gastro- 
poda).—Bollettino Malacologico, Supplemento 
I: 123-296. 

Dell, R. K. 1964. Antarctic Mollusca.—Bulletin of the 
Royal Society of New Zealand 27:1—311. 
Harasewych, M. G., & Y. I. Kantor. 1999. A revision 
of the Antarctic genus Chlanidota (Gastropoda: 
Neogastropoda: Buccinulidae).—Proceedings 
of the Biological Society of Washington 112: 

253-302. 

Lus, V. Y. 1981. On the abyssal species Sipho (Si- 
Phonorbis) danielsseni (Friele) and Mohnia 
mohni (Friele) (Gastropoda: Buccinidae).—Tru- 
dy Instituta Okeanologii AN SSSR (Proceed- 
ings of the Instute of Oceanology of the USSR 
Ac. Sci.) 115:126—140 (in Russian). 

Numanami, H. 1996. Taxonomic study on Antarctic 
gastropods, collected by Japanese Antarctic re- 
search expeditions.—Memoirs of National In- 
stitute of Polar Research, Series E, Biology and 
Medical Science 39:1—244. 

Powell, A. W. B. 1951. Antarctic and SubAntarctic 
Mollusca: Pelecypoda and Gastropoda.—Dis- 
covery Reports 26:47—196, pls. 5—10. 

Smith, E. H. 1967. The Neogastropod stomach, with 
notes on the digestive diverticula and intes- 
tine.—Transactions of the Royal Society of Ed- 
inburgh, 67(2):23—42. 

Thiele, J. 1912. Die antarktischen Schecken und Mus- 
cheln.—Deutschen Sitidpolar-Expedition 1901— 
1903 13:183-—285. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 
113(2):561-571. 2000. 


Porites arnaudi, a new species of stony coral 
(Anthozoa: Scleractinia: Poritidae) from oceanic islands of the 
eastern Pacific Ocean 


Héctor Reyes-Bonilla and Juan P. Carricart-Ganivet 


(HRB) Universidad Aut6noma de Baja California Sur, Departamento de Biologia Marina, Apdo. 


Postal 19-B, La Paz, B.C.S. 23000, México; (JPCG) Departamento de Ecologia Acuatica, 
ECOSUR, Apdo. Postal 424, Chetumal, Q. Roo, 77000, México 


Abstract.—A new species of Porites (P. arnaudi) was found at depths of 7 
to 37 m at Clipperton Atoll, France, and the Revillagigedo Islands, México, 
oceanic islands located west of the American mainland (eastern Pacific). Col- 
onies are uncommon but very distinctive because of the presence of tiered 
plates, or laminae, expanding from a single base, smooth and concave surfaces, 
and a lack of living tissue on the inferior part of the plates, except on its 
growing edge. Calices are 0.8 to 1.4 mm in diameter, with wide walls. Cor- 
allites have a free triplet with all three septa about the same size. Lateral septa 
are fused in pairs and better developed than the rest. There are six to eight 
pali, and one or two denticles per septum, and the color of tissue is greenish- 
gray to dark brown. With this new species, the current total number of species 
in the genus Porites in the eastern Pacific reaches nine, with most species 


restricted to oceanic islands of the region. 


In the last 20 years, taxonomic studies on 
the scleractinian zooxanthellate fauna of the 
eastern Pacific have noticeably increased in 
number and quality (Wells 1983, Guzman 
& Cortés 1993, Hodgson 1995) and as a 
consequence, there is now fairly good 
agreement on coral identities since some 
key taxonomic problems have been re- 
solved (Squires 1959, Wells 1983, Veron 
1986). New research also has shown that 
the actual richness of coral species from the 
eastern Pacific is much higher than once 
thought [e.g., originally less than 20 species 
according to Veron (1993) and Paulay 
(1997)]. For example, México, Costa Rica 
and Panama have 20 to 25 species each and 
almost 40 are known to inhabit western 
America (Holst & Guzman 1993, Cortés & 
Guzman 1998, Reyes-Bonilla & Lé6pez- 
Pérez 1998). This perceived increase in 
richness comes from new records for par- 
ticular localities or areas (e.g., Reyes-Bon- 
illa 1992, Cortés & Guzman 1998) and by 


descriptions of new species (Budd & Guz- 
man 1994, Glynn 1999). 

Members of the genus Porites Link, 
1807 are widely distributed in the eastern 
Pacific, and they are one of the dominant 
corals in this region, both in abundance and 
species richness (Glynn 1997). To date, 
eight species have been reported: P. aus- 
traliensis Vaughan, 1918, P. baueri 
Squires, 1959, P. lichen Dana, 1846, P. lob- 
ata Dana, 1846, P. lutea Milne Edwards, 
1860, P. panamensis Verrill, 1866, P. rus 
Forskaal, 1775, and P. sverdrupi Durham, 
1947 (Guzman & Cortés 1993, Glynn 1997, 
Reyes-Bonilla 1999). Both P. panamensis 
and P. lobata are present in coral commu- 
nities from the Galapagos Islands, or the 
Ecuadorian mainland, to México (Reyes- 
Bonilla 1993, Glynn 1997), while P. sver- 
drupi and P. baueri are endemic to the Gulf 
of California and the Marias Islands, Méx- 
ico, respectively (Squires 1959, Reyes-Bon- 
illa & Lopez-Pérez 1998). Porites rus is an 


562 


Indo Pacific species that was reported from 
Costa Rica in the early 1980’s, but its pop- 
ulation may be extinct now (Glynn 1997, 
Cortés & Guzman 1998). The remaining 
three species are Indo Pacific also, but they 
have been observed in the Revillagigedo 
Archipelago, México, a group of four oce- 
anic islands located about 300 km south- 
west of the Baja California peninsula, or at 
Clipperton Island, the only true atoll of the 
eastern Pacific, 1300 km SW of the Mexi- 
can mainland (Glynn et al. 1996, Ketchum 
& Reyes-Bonilla 1997). 

In this paper, a new species of Porites is 
described. This coral is distinctive because 
its colonies have been found exclusively in 
the oceanic Clipperton and Revillagigedo 
Islands, eastern Pacific. This finding in- 
creases the known number of species of the 
genus Porites to nine, a number even great- 
er than that reported for the Atlantic-Carib- 
bean area (Weil 1992, Veron 1993). 

Abbreviations of repository institu- 
tions.—USNM: United States National Mu- 
seum of Natural History, Department of In- 
vertebrate Zoology, Smithsonian Institu- 
tion, Washington, D.C. 20560, U.S.A.; 
MHNUABCS: Museo de Historia Natural 
de la Universidad Aut6noma de Baja Cali- 
fornia Sur. Departamento de Biologia Ma- 
tina.«a; Paz .B:C:S3) 230803 Mexico: 
ECOCHBC: Colecci6n de Corales de El 
Colegio de la Frontera Sur, Grupo de Ecol- 
ogia Bentonica. Apartado postal 424, Che- 
tumal, Q. Roo, 77000, México. UMML: 
University of Miami Marine Laboratory. 
Rosenstiel School of Marine and Atmo- 
spheric Sciences. 4600 Rickenbacker 
Causeway, Miami, FL 33149, U.S.A. 


Order Scleractinia Bourne, 1900 
Suborder Fungiina Verrill, 1865 
Superfamily Poritoidea Gray, 1842 
Family Poritidae Gray, 1842 
Genus Porites Link, 1807 


Type species.—Porites polymorphus 
Link, 1807 = Madrepora porites Pallas, 
1766 (in part). 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Diagnosis.—Massive, ramose or encrust- 
ing cerioid coralla. Corallites with calices 
smaller than 2 mm in width, with two cy- 
cles of septa. Septa commonly bear spines 
and are formed by three or four trabeculae 
that in their innermost portions are differ- 
entiated as pali. A single trabecular colu- 
mella, absent in certain species. Colony for- 
mation by extratentacular budding. Coenos- 
teum poorly developed (Wells 1956). 

Distribution.—The genus is circumtrop- 
ical, recorded in the Pacific, Indian and At- 
lantic oceans, as well as in the Caribbean 
Sea (Veron 1993). Fossil records are known 
from the Cretaceous (?), Eocene to Recent, 
but are more abundant and dominant in reef 
communities since the Miocene (Wells 
1956, Veron 1995). In the eastern Pacific 
the genus is distributed from the Galapagos 
Islands (O°N), Ecuador, to the northern Gulf 
of California (31°N), México, including all 
adjacent oceanic islands (Wells 1983, 
Reyes-Bonilla 1993). 


Porites arnaudi, new species 
Figs. 1-6 


Etymology.—Named after Gustavo Ar- 
naud Vignon, the last Commander in Chief 
of the Mexican Navy garrison on Clipper- 
ton Island, who died tragically while on 
duty on 1916. 

Diagnosis.—(Terminology after Veron & 
Pichon 1982 and Veron 1986) Oval shaped 
colonies composed of tiered, thick plate- 
like laminae (at least 10 mm in width). Cor- 
allum originates from a single wide pedicel, 
with smooth, concave and undulated sur- 
faces, with no living tissue on the inferior 
parts, except on its growing edges. Coral- 
lites distinct, calices 0.8 to 1.4 mm wide, 
with walls less than 1 mm wide. Septa rare- 
ly bifurcated. Free triplet, although the ven- 
tral directive and one lateral can be joined. 
Lateral septa fused in pairs and better de- 
veloped than the rest. One or two conspic- 
uous denticles in the inner part of each sep- 
tum. Six to eight pali which do not reach 
the level of the wall; the palus of the triplet 


VOLUME 113, NUMBER 2 


563 


Fis. 71; 
b, side view. Both X0.49. 


and dorsal directive less developed than 
those of the lateral pairs. Two synapticular 
rings, the external more defined. Living tis- 
sue color greenish-gray to dark brown, with 
pale margins. 

Holotype.—USNM 100261 (Figs. la, b; 
4a, b; 5a, b): length 213 mm, width 148 
mm, height 111 mm, thickness of peripher- 


Porites arnaudi, holotype, USNM 100261. Recent, 30 m depth, Clipperton Atoll, France. a, top view; 


ical edge of folia 19 to 21 mm; collected 
25 Nov 1997 by Juan P. Carricart-Ganivet. 

Paratypes.—ECOCHBC 0107 (Fig. 2a, 
b), MHNUABCS 1044, 1076, 1092, 1100, 
1103 74581¢ (Pig. 3c; .d))- 1582: (Fig: Sa, b), 
1583, UMML 8.1475 (Fig. 6a, b). 

Type locality.—Northeast seaward slope 
(sensu Glynn et al. 1996), off Clipperton 


564 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 2: 
view; b, side view. Both X0.79. 


Atoll, France (10°18’N, 109°13’W), at 30 m 
depth. 

Description.—Colonies submassive, 
forming conspicuous tiered plates growing 
outwards from a central base (discoidal 
morphology; Figs. la, b; 2a, b); coralla can 
attain more than 40 cm in diameter and 
about 30 cm in height, being firmly at- 
tached to the substrate. Smooth, concave, 
and undulated upper surfaces, with thick fo- 


Porites arnaudi, paratype, ECOCHBC 0107. Recent, 30 m depth, Clipperton Atoll, France. a, top 


lia at least 10 mm in width (Fig. 3a—d). In- 
ferior surfaces devoid of living tissue ex- 
cept in their growing edges. Corallites 0.8 
to 1.4 mm diameter, conspicuous, with 
deep, polygonal calices (normally five to 
six sides) having six to eight deep-set pal, 
which do not reach the level of the wall. 
Intercalicular distances 0.8 to 1.7 mm (Figs. 
4a, b). Corallite walls less than 1 mm thick 
and composed of three rows of denticles. 


VOLUME 113, NUMBER 2 565 


Fig. 3. Porites arnaudi, paratypes, top and side views of laminae from the edge of larger colonies: a, b, 
MHNUABCS 1582; c, d, MHNUABCS 1581. Both recent, 21 m depth, Clipperton Atoll, France. All figures 
x0.30. 


566 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 4. Porites arnaudi, variation of skeletal structure in the holotype USNM 100261. Notice diagnostic 
characters such as polygonal calices and wide walls. a, X6; b, x9. 


Twelve septa rarely bifurcated (Figs. 5a, b; 
6a, b). The triplet normally has free mar- 
gins, although it can also have the directive 
and a lateral septum (generally that of the 
right side) fused in the inner section. Lateral 
septa arranged in four fused pairs, better de- 
veloped and longer than the rest, and in its 


innermost portion presents conspicuous pali 
that rarely reach the level of the corallite 
wall. Pali of the triplet and the dorsal di- 
rective septa less developed than those of 
lateral pairs, or may be absent (Fig. 6a, b). 
One or two denticles per septum, which can 
resemble small pali because of extensive 


VOLUME 113, NUMBER 2 


4 


ae 


Fig: 5. 


567 


Porites arnaudi, variation of skeletal structure in the holotype USNM 100261. Notice diagnostic 


characters such as prominent pali, inconspicuous columella, one denticle per septum and free triplet (arrow). a, 


X34; b, X66. 


development and their position near the cal- 
icular center. Columella quite variable, 
sometimes appearing as a small column or 
stylet, may be compressed dorsoventrally, 
or may be absent. In some, a second prom- 
inent protuberance about the same size as 
the columella may be present near the inner 


margin of the dorsal directive. Two synap- 
ticular rings, the external more defined than 
the inner palar. Tissue color from greenish- 
gray to dark brown, with pale margins. 
Distribution.—This species was first re- 
corded by Glynn et al. (1996) at Clipperton 
Atoll (southeast, northeast and southwest 


568 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 6. Porites arnaudi, SEM photos of the paratype UMML 8.1475. Recent, 18 m depth, Clipperton Atoll, 


France. a, group of calices X16. b, single calice X38. 


sites) and at San Benedicto (19°18'N, 
110°48’W) and Clarién islands (18°20’N, 
114°44’W), Revillagigedo Archipelago, un- 
der the name Porites sp. Later, Ketchum & 
Reyes-Bonilla (1997) confirmed its pres- 
ence in San Benedicto and Clarion, and also 
at Socorro Island (18°45’N; 111°00’W), 


calling it Porites sp. 1. However, no exact 
locations of collection or observation sites 
were presented in those papers. The holo- 
type (USNM 100261, collected by J. P. Car- 
ricart-Ganivet at 30 m depth, 25 Nov 1997) 
and six other specimens on which the cur- 
rent description is based (UMML 8.1475, 


VOLUME 113, NUMBER 2 


collected by P. W. Glynn at 18 m depth, 14— 
26 Apr 1994; ECOCHBC 0107 collected by 
A. Medina-Carcamo at 30 m depth, 25 Nov 
1997; MHNUABCS 1581, 1582, 1583 col- 
lected by H. Reyes-Bonilla at 21 m depth, 
26 Nov 1997) were found in the ‘‘Poritid 
zone” (sensu Glynn et al. 1996), at the 
northeast seaward slope, Clipperton Atoll. 
Other coralla of the same species were sam- 
pled at Playa Norte and Punta Tosca (So- 
corro Island; 18°47'25"N, 111°01'15’W, 
MHNUABCS 1044, collected by J. EK Cas- 
tellanos-Avila at 14 m depth, 30 Nov 
1991), Bahia Azufre (Clarion Island; 
18°20'47"N, 114°44'00"W, MHNUABCS 
1092, 1100, 1103, collected by J. Ketchum 
at 7, 7 and 10 m depth, 15 Dec 1994) and 
Roca Trinidad (San Benedicto Island; 
19°18'59"N, 110°49'00"W, MHNUABCS 
1076, collected by H. Reyes-Bonilla at 15 
m depth, 20 May 1995). As Porites arnaudi 
has not been observed elsewhere in the cen- 
tral Pacific or off American mainland, it is 
considered to be endemic of the oceanic 
Revillagigedos and Clipperton islands. 
Ecology.—This newly described coral is 
distributed at depths from 7 to 37 m, es- 
pecially in steep rocky slopes, and often 
overgrows adjacent coralla, mostly Porites 
lobata and Pavona varians Verrill, 1864 in 
Clipperton (Glynn et al. 1996) and P. lob- 
ata and Psammocora superficialis Gardiner, 
1898 in the Revillagigedos. It is not abun- 
dant; Glynn et al. (1996) mentioned that 
percent live cover was up to 7.4% in the 
southeast side of Clipperton, while in the 
Revillagigedos it is seldom found, although 
it is locally abundant at Playa Norte (So- 
corro Island), Bahia Azufre (Clarion Island) 
and Roca Trinidad (San Benedicto Island). 
Coralla commonly shows bite marks in the 
outer edges of the plates, most likely pro- 
duced by Arothron meleagris (Bloch & 
Schneider, 1801) or Melichthys niger 
(Bloch, 1786), both fishes that frequent lo- 
calities where Porites arnaudi has been 
found (Allen & Robertson 1994, Robertson 
& Allen 1996). Bioeroders and serpulid 
polychaetes were rare in colonies of this 


569 


species observed at Clipperton and practi- 
cally absent in those collected for this pa- 
per. In contrast, they were much more abun- 
dant in the Revillagigedos, in particular the 
bivalves Lithophaga spp., which commonly 
bore the bases of the plates. 
Comparison.—Because Porites arnaudi 
has been found in Clipperton and at the Re- 
villagigedo Archipelago, it must first be dif- 
ferentiated from all other conspecifics pre- 
sent in those localities: P. lobata, P. lutea, 
P. lichen, P. australiensis and P. panamen- 
sis (Ketchum & Reyes-Bonilla 1997, 
Reyes-Bonilla 1999). The coral here de- 
scribed is similar to P. lobata, as noted by 
Glynn et al. (1996), because both species 
have thick ledges with an inconspicuous 
columella, well developed dorsal directive 
septa and lateral pairs, palus of the dorsal 
directive septum smaller than those of the 
laterals, and septa of the triplet always free 
(Veron & Pichon 1982, Veron 1986; Figs. 
5a, b; 6a, b). Notwithstanding, the former 
can be identified by its plate-like tiered col- 
onies (Figs. la, b, 2a, b, 3a—d) which are 
not massive or hemispherical as in P. lob- 
ata. Also, it presents a well-defined syn- 
apticular ring and different tissue color 
(gray or brown in P. arnaudi and blue, pur- 
ple or green in P. lobata). A third differ- 
ence is that P. lobata has two denticles on 
each septum, whereas P. arnaudi normally 
has only one (Fig. 6b). Porites arnaudi and 
P. lutea differ noticeably because the latter 
is massive and has columniform lobes, shal- 
low corallites and thin walls. In addition, in 
P. lutea, pali reach the level of the walls, 
all septa of the triplet are fused by a trans- 
verse rod or in their margins (in a “‘tri- 
dent’), bifurcate septa often appear and 
columella is well developed. Porites arnau- 
di can be easily separated from P. austral- 
iensis because the latter has a large and 
well-developed columella and pali that 
reach the level of the wall denticles. Also, 
in the triplet, the former species has lateral 
septa about the same size as those of the 
ventral directive (Figs. 5a, b, 6a, b), while 
in P. australiensis the lateral septa are 


570 


smaller than those of the ventral directive, 
and occasionally have fused triplets instead 
of free ones. The morphology of the cor- 
allum is quite different also, as P. austral- 
iensis (like P. lobata) has massive, almost 
spherical colonies (Veron & Pichon 1982, 
Veron 1986). Porites panamensis can be 
distinguished from P. arnaudi principally 
because its colonies are very small (never 
larger than 40 cm height and diameter), en- 
crusting, massive or columnar, and its color 
is bright green instead of gray or brown. 
Other differences are that P. panamensis 
has fewer pali (normally five) and thin 
walls (Squires 1959). Porites lichen is an- 
other species quite similar to P. arnaudi, 
having six to eight pali and wide walls, one 
denticle per septum, thick ledges around 
colony bases, or appearing as laminar plates 
(Veron & Pichon 1982, Veron 1986). Nev- 
ertheless, they can be differentiated on the 
basis of the corallites of P. lichen which 
have septa of the triplet commonly fused in 
their margins, while directive dorsal septa 
of the same structure are shorter than lat- 
erals. Also, coralla of this species appear as 
columnar colonies, and when they occur as 
laminae, these are thin and develop only in 
the base of the coralla, a character never 
presented in P. arnaudi (Figs. la, b, 2a, b, 
3a—d). 


Acknowledgments 


Specimens for this work were collected 
in a series of visits to the Revillagigedos 
supported by UABCS, Comisi6n Nacional 
para el Conocimiento y Uso de la Biodiv- 
ersidad (contract FB342/H337/96), and 
Baja Expeditions, and during the cruise 
SURPACLIP-I to Clipperton and Socorro 
islands, under the direction of Vivianne So- 
lis Weiss (Instituto de Ciencias del Mar y 
Limnologia, Universidad Nacional Auton- 
oma de México, México City). James Ket- 
chum, Juan Francisco-Castellanos, Alejan- 
dro Granados, Miguel Angel-Garcia, Adri- 
an Medina-Carcamo, Leonardo Ortiz and 
Vivianne Solis collaborated in diving activ- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


ities. Peter W. Glynn, Stephen D. Cairns, 
James Ketchum and Andrés Lépez dis- 
cussed and commented on preliminary ver- 
sions of the manuscript. Special thanks to 
Scott Monks for correcting the English, and 
for his valuable comments on the first draft 
of the manuscript. Photographs taken by 
Humberto Bahena Basave at ECOSUR, and 
scanning electron microscope images taken 
at the USNM by Jarek Stolassky. 


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tropical eastern Pacific. University of Hawaii 
Press, Honolulu, Hawau, 332 pp. 

Budd, A. F, & H. M. Guzman. 1994. Siderastrea glyn- 
ni, a new species of scleractinian coral (Cnidar- 
ia: Anthozoa) from the eastern Pacific.—Pro- 
ceedings of the Biological Society of Washing- 
ton 107:591—599. 

Cortés, J.. & H. M. Guzman. 1998. Organismos de los 
arrecifes coralinos de Costa Rica: Descripcion, 
distribuci6n geogrdafica e historia natural de los 
corales zooxantelados (Anthozoa: Scleractinia) 
del Pacifico de Costa Rica.—Revista de Biolo- 
gia Tropical 46:55—92. 

Glynn, P. W. 1997. Eastern Pacific reef coral bioge- 
ography and faunal flux: Durham’s dilemma re- 
visited.—Proceedings of the 8th International 
Coral Reef Symposium, Panama 1:371—378. 

. 1999. Pocillopora inflata, a new species of 

Scleractinian coral (Cnidaria: Anthozoa) from 

the tropical eastern Pacific—Pacific Science 53: 

168-180. 

, J. E. N. Veron, & G. M. Wellington. 1996. 
Clipperton Atoll (eastern Pacific): oceanogra- 
phy, geomorphology, reef-building coral ecol- 
ogy and biogeography.—Coral Reefs 15:71-99. 

Guzman, H. M., & J. Cortés. 1993. Arrecifes coralinos 
del Pacifico oriental: revisi6n y perspectivas.— 
Revista de Biologia Tropical 41:535—557. 

Hodgson, G. 1995. Corales pétreos marinos (Tipo Cni- 
daria, Orden Scleractinia). Pp. 83-97 in W. Fi- 
scher, E Krupp, W. Schneider, C. Sommer, K. 
E. Carpenter, & V. H. Niem, eds., Guia FAO 
para la identificaci6n de especies para los fines 
de la pesca. Vol. 1. Algas e invertebrados, FAO, 
Rome, 646 pp. 

Holst, I, & H. M. Guzman. 1993. Lista de corales 
hermatipicos (Anthozoa: Scleractinia; Hydro- 
zoa: Milleporina) a ambos lados del istmo de 
Panama.—Revista de Biologia Tropical 41: 
535-540. 

Ketchum, J. T., & H. Reyes-Bonilla. 1997. Biogeog- 
raphy of hermatypic corals of the Revillagigedo 


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Archipelago, México.—Proceedings of the 8th 
International Coral Reef Symposium, Panama 
1:471—476. 

Paulay, G. 1997. Diversity and distribution of reef or- 
ganisms. Pp. 298-353 in C. E. Birkeland, ed., 
Life and death of coral reefs, Chapman & Hall, 
New York, 536 pp. 

Reyes-Bonilla, H. 1992. New records for hermatypic 
corals (Anthozoa: Scleractinia) in the Gulf of 
California, México, with an historical and bio- 
geographical discussion.—Journal of Natural 
History 26:1163—1175. 

. 1993. Biogeografia y ecologia de los corales 

hermatipicos (Anthozoa: Scleractinia) del Paci- 

fico de México. Pp. 207—222 in S. I. Salazar- 

Vallejo & N. E. Gonzalez, eds., Biodiversidad 

marina y costera de México, Comisi6n Nacional 

para el Conocimiento y Uso de la Biodiversidad 

& Centro de Investigaciones de Quintana Roo, 

Chetumal, 865 pp. 

. 1999. Corals and coral reefs of the Pacific 

coast of México. in J. Cortés, ed., Coral reefs 

of Latin America, Elsevier, Amsterdam. (in 
press.) 

, & A. Lépez-Pérez. 1998. Biogeography of the 
stony corals (Scleractinia) of the Mexican Pa- 
cific.—Ciencias Marinas 24:211-—224. 

Robertson, D. R., & G. R. Allen. 1996. The shorefish 
fauna of Clipperton Atoll (tropical eastern Pa- 
cific) and its zoogeographical affinities.—Coral 
Reels: 15:121—131. 


aM 


Squires, D. R. 1959. Corals and coral reefs in the Gulf 
of California.—Bulletin of the American Mu- 
seum of Natural History 118:370—431. 

Veron, J. E. N. 1986. Corals of Australia and the Indo 
Pacific. Angus & Robertson Publishers, Sydney, 
644 pp. 

. 1993. A biogeographic database of hermatyp- 

ic corals.—Australian Institute of Marine Sci- 

ence Mongraph Series 10:1—433. 

. 1995. Corals in space and time: the bioge- 

ography and evolution of the Scleractinia. Cor- 

nell University Press, Ithaca, 321 pp. 

, & M. Pichon. 1982. Scleractinia of Eastern 
Australia. Part IV. Family Poritidae.—Austra- 
lian Institute of Marine Science Monograph Se- 
ries 5:1-159. 

Weil, E. 1992. Genetic and morphological variation in 
Caribbean and eastern Pacific Porites (Antho- 
zoa: Scleractinia). Preliminary results.—Pro- 
ceedings of the 7th International Coral Reef 
Symposium, Guam 2:643—656. 

Wells, J. W. 1956. Scleractinia. Pp. F328—F444 in R. 
C. Moore, ed., Treatise of invertebrate paleon- 
tology, vol. EF Geological Society of America 
and University of Kansas Press, Lawrence, 
Kansas, 498 pp. 

. 1983. Annotated list of the Scleractinian cor- 

als of the Galapagos Islands. Pp. 212-295 in P. 

W. Glynn & G. M. Wellington eds., Corals and 

coral reefs of the Galapagos Islands. University 

of California Press, Berkeley, California, 330 


PP. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


113(2):572-578. 2000. 


A new species of Heterotella (Porifera: Hexactinellida: Euplectellidae) 
from the West Indies 


Henry M. Reiswig 


Redpath Museum & Biology Department, McGill University, 859 Sherbrooke St. West, Montreal, 
Quebec, Canada H3A 2K6, e-mail: cxhr@musica.mcgill.ca 


Abstract.—Heterotella pomponae, a new species, is reported from near S. 
Caicos Is., West Indies. This is only the second described species of the genus 
and the first from the Atlantic Basin. Heterotella was erected by Gray to ac- 
commodate Alcyoncellum corbicula Bowerbank from Isle de Bourbon (Re- 
union), southwest Indian Ocean. The genus is distinguished from other Cor- 
bitellinae by a microsclere which has controversial interpretation, as either a 
oxyhexactin with long spines or an oxyhexaster with irregularly originating 
terminal rays. The two species are distinguished by presence of sigmatocomes 
in H. pomponae and their absence in H. corbicula. 


‘The first taxonomically acceptable refer- 
ence to a sponge now included in the genus 
Heterotella was made by Bowerbank 
(1867:358), where he clearly referred the 
earlier published figure of a distinctive mi- 
crosclere (Bowerbank 1858, 1864) to a 
specimen labelled Alcyoncellum corbicula 
from Isle de Bourbon in the Paris Museum. 
The genus Alcyoncellum had a very con- 
voluted early history, originally attributed 
to a calcareous sponge and later considered 
to be an uncertain synonym of the hexac- 
tinellid genera Euplectella and Corbitella. 
The status of many of these previously con- 
founded generic names was clarified by 
Gray (1967), where he erected the genus 
Heterotella to contain only Bowerbank’s A. 
corbicula. The two closely related genera, 
Corbitella and Heterotella, were later re- 
viewed by Ijima (1902), who described the 
three then existing specimens of H. corbic- 
ula from Isle de Bourbon (Reunion) in the 
Museum National d’ Histoire Naturelle, Par- 
is. The earlier figured and still diagnostic 
microsclere of the genus was considered in 
some detail. Ijima interpreted the oxy- 
tipped spicule to be a hexactin with long, 
irregular spines, but noted that this could 
alternately be accepted as a oxyhexaster 


with irregularly arising terminal rays. Kirk- 
patrick (1910) reviewed the spicule in his 
attempt to systematize spicule nomencla- 
ture, and decided that it was an oxyhexaster 
since the axial canal did not extend to the 
ray tip. These distinctions remain arbitrary 
since there is no method of objectively dif- 
ferentiating spines from terminal rays. 

No other Heterotella specimens have 
been added to that type series collected 
from Reunion in 1819 and 1857. Here I de- 
scribe the first recent specimen of this ge- 
nus, collected in 1994 from S. Caicos Is- 
land, West Indies. The specimen represents 
a new species of Heterotella and the first 
member of the genus from a location other 
than Reunion, Indian Ocean. 


Materials and Methods 


The specimen of an unidentified lyssa- 
cine hexactinellid (the new species de- 
scribed herein) was obtained on loan from 
Harbor Branch Oceanographic Museum, 
Fort Pierce, Florida (HBOM). The type se- 
ries of H. corbicula was reviewed during a 
1997 visit to the Museum National 
d’ Histoire Naturelle, Paris (MNHN). Frag- 
ments of two of the MNHN specimens were 


VOLUME 113, NUMBER 2 


returned to Montreal where skeletal prepa- 
rations were made. Small pieces of dermal 
and gastral surfaces were either whole- 
mounted in balsam for light microscopy 
(LM) or digested in hot nitric acid. Large 
spicules in the resulting spicule suspensions 
were rinsed, spread on microscope slides 
and mounted in balsam. Smaller spicules 
were dispersed on 25 mm diameter, 0.2 ~m 
pore-size, nitrocellulose filters by filtration; 
the filters were rinsed, dried and mounted 
in balsam. Spicules were measured by com- 
puter via a microscope-coupled digitizer. 
Data are reported as: mean = St. deV. number 
(range). Spicule drawings were prepared 
from video-captured microscope images 
imported into a computer drawing program 
and traced on-screen. Samples for scanning 
electron microscopy (SEM) were nitric- 
acid-cleaned and mounted on stubs with ep- 
oxy. Following gold-palladium coating, 
specimens were viewed and photographed 
with a JEOL JSM-840 SEM. 


Systematics 


Subphylum Symplasma Reiswig & 
Mackie, 1983 
Class Hexactinellida Schmidt, 1870 
Subclass Hexasterophora Schulze, 1886 

Order Lyssacinosa Zittel, 1877 

Family Euplectellidae Gray, 1867 

Subfamily Corbitellinae Ijima, 1902 
Genus Heterotella Gray, 1867 


Type species.—Heterotella corbicula 
(Bowerbank, 1867). 

Diagnosis.—(from Tjima 1902:32 for H. 
corbicula, emended here). Corbitellinae of 
saccular shape, the lateral walls constricting 
distally to the margins of a terminal sieve- 
plate. With numerous, round, irregularly ar- 
ranged parietal oscula. Skeletal beams un- 
fused or totally fused by synapticula. Prin- 
cipalia parenchymalia as diactins; accesso- 
ria as diactins and hexactins. With 
distinctive microxyhexasters in which the 
slender terminal rays are irregularly undu- 
lating and do not all originate from a single 
point (often interpreted as microxyhexac- 


eH eo. 


tins with irregular spines). Floricome and 
graphiocome present, with or without sig- 
matocome; discohexaster lacking. 

Remarks.—A holotype has not been des- 
ignated for H. corbicula from among the 
original three specimens in the collections 
at MNHN. A lectotype is here designated 
as MNHN HX 23, Ijima’s specimen A, the 
specimen upon which he based his exten- 
sive spicule description. The specimen can 
be identified as consisting of only the top 
portion of a sponge from Isle de Bourbon 
donated by Mr. Leschinault, 1819. The two 
remaining specimens, MNHN HX 24 & 25, 
Ijima’s B and C, thus become paralecto- 
types. Lectotype designations are made 
here with the purpose of clarifying the ap- 
plication of the name to a taxon. 


Heterotella pomponae, new species 
Figs. 1-10, Table 1 


Material examined.—Holotype: HBOM 
0027-00019? km: -S) “ofes. Caicos’ IJs., 
21°28.84'N, 71°29.985'W, 467 m; col. A. 
Wright via RMS Johnson SeaLink I, dive 
3788, 31 Oct 1994; in alcohol. 

Shape.—Entire specimen with basal disc 
retrieved; broken during handling. Thin- 
wall, sac-shaped sponge (Fig. 1), 13 cm tall 
by 5.5 cm diam, attached to hard substrata 
by short basal disk supporting a short, rigid 
basal cup. Wall, 2.3 mm in maximum thick- 
ness, perforated by numerous primary pa- 
rietal oscula passing directly through the 
wall, diameter 1.7 + 0.3,; mm (1.1-2.1 
mm), spaced at 6.4 + 1.4,; mm (3.1—9.0 
mm); smaller openings passing obliquely or 
tortuously through the wall (secondary pa- 
rietal oscula) have diameter on the dermal 
side of 1.5 + 0.73, mm (0.4-—2.7 mm, Fig. 
2) and on the atrial side 1.0 + 0.3,, mm 
(0.5—2.0 mm, Fig. 3). Parietal oscula are ar- 
ranged without apparent pattern. The ter- 
minal osculum, 2.3 cm diam is covered by 
a sieve plate and bordered by a vertical 
marginal collar. 

Surface texture.—Dermal tissue is deli- 
cate and subject to detachment by gentle 


574 


Fig. 1. Heterotella pomponae n. sp. (holotype) 
cut-away diagram reconstructed from photographs, 
showing distribution of main parietal oscula, terminal 
sieve plate and location of sinuous diactins in the base 
of the atrium (wavy filled area). 


water currents; gastral tissue is firmer and 
more adherent to the skeletal framework. 
Both surfaces are smooth to eye but irreg- 
ularly lumpy under a dissecting micro- 
scope. Prostalia are lacking; the fine tips of 
sword hexactins projecting from the cush- 
ion-like white tissues can be perceived only 
with aid of a microscope; no regular grid 
arrangement is detectable on either surface. 
Living tissues with spicules are not present 
in the sieve plate or marginal collar; mar- 
ginalia absent, but may have existed in 
younger stage. 

Color.—White in life and preserved in 
ethanol. 

Skeleton.—The entire framework of the 
holotype (basal cup, lateral walls, sieve 
plate) has been solidified into a rigid struc- 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


ture by siliceous synapticulae so profuse as 
to obscure the nature of the primary frame- 
work-forming megascleres (Figs. 4-8). 
These are presumed to be diactins since 
hexactin nodes have not been found in this 
structure and diactins are known to be prin- 
cipalia of H. corbicula. Major siliceous el- 
ements of most of the framework appear 
randomly oriented (Fig. 4), but the outer 
layer contains major longitudinal bundles 
and the inner-most atrial bundles are trans- 
versely oriented. The outer-most strands of 
the framework are nodulated by secondary 
silicification (Fig. 5) while the middle and 
internal strands are smooth. Thickness of 
the framework is 1.6 + 0.2,, mm (1.2—1.9 
mm). In the basal region of the body (Fig. 
1), a dense matt of sinuous diactins joined 
by very short synapticulae or point cemen- 
tation, is deposited on the internal surface 
of the framework (Fig. 6); this matt nearly 
fills the mesh spaces (Fig. 7), occludes 
some parietal oscula and strengthens the 
basal framework. 

Elements of the sieve plate and marginal 
collar are rigidified and nearly filled to solid 
sheets by dense synapticulation (Figs. 8, 9). 
The marginal collar extends up from the 
body surface 1.9 + 0.4,, mm (1.4—2.4 mm), 
as a nearly solid sheet of fused, transverse- 
ly-oriented, siliceous spicules. Major 
strands of the body framework extend up 
the inner side of the collar and project out 
into the oscular plane as sieve beams (Fig. 
8). The sieve beams are unusual in their 
vertical sheet-like form, being taller than 
wide (septa-like), with beam height: 0.78 + 
0.4,, mm (0.2-1.9 mm), width: 0.25 + 
0.10,; mm (0.11—0.53 mm). Sieve pores are 
variable in shape and size (Fig. 9), but are 
mainly subtriangular with sides 1.5 + 0.4,, 
mm (0.6—2.3 mm). 

Loose spicules.—Diagrams of spicule 
form and dimensions are summarized in 
Figure 10 and Table 1. 

Megascleres: The major surface spicules 
of both dermal and gastral surfaces are 
smooth, sword-shaped hexactins with the 
short ray projecting from the tissue surface 


VOLUME 113, NUMBER 2 +75 


vine ;. 4 
> => 
- @ 


“a th ‘ 
i, A ner + 


Figs. 2-9. Heterotella pomponae n. sp. (holotype). 2-3. Photographs of dermal (2) and gastral (3) surfaces 
with tissues, showing differences in aperture distributions; scale equals 1 cm (LM). 4—5. External surface of 
acid-cleaned, fused framework of upper body wall; scale equals 1 mm (SEM). 6. External surface of basal region 
with meshes filled with sinuous diactins; scale equals 1 mm (SEM). 7. Internal surface of the cup base covered 
with dense pad of sinuous diactins; scale equals 1 mm (SEM). 8. Oblique vie of a fragment of the fused marginal 
collar and sieve plate showing continuity of sieve beams with wall bundles: scale equals 1 mm (LM). 9. Side 
view of a major sieve plate beam, top surface to left, showing complete fusion of constituent diactins; scale 0.1 
mm (LM). 


576 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 10. Loose spicules of Heterotella pomponae n. sp. (holotype). A. Superficial dermal and gastral sword 
hexactins grading to pentactin. B. Choanosomal regular hexactins. C. Centrotylote diactin. D. Ends of sinuous 
diactins. E. Oxyhexaster. F Floricome. G. Sigmatocome. H. Graphiocome centrum with long and short terminals 
(raphides). Scale bars: Left 100 1m bar applies to A, B; right 100 wm bar applies to C, D; 25 wm bar applies 


to E—-H. 


(Fig. 10A); a moderate number of pentac- 
tins (11%) may result by reduction of the 
distal ray. These occur in mixtures of thin 
and thick forms on both surfaces. True mar- 
ginalia were not found—the marginal collar 
was devoid of tissues or loose spiculation. 
Nearly equal-rayed, sparsely-spined hexac- 
tins (Fig. 1OB) occur throughout the choan- 
osome in small numbers; they vary greatly 
in size and may consist of two classes, but 
frequency analysis was not performed. 
Sparsely-spined, centrotylote diactins with 
acute tips (Fig. 10C) are abundant through- 
out the choanosome. Smooth, sinuous diac- 


tins with rounded tips (Fig. 10D) pack the 
lower framework. They occur as fragments 
up to 6+ mm long in loose spicule prepa- 
rations, presumably broken from‘their syn- 
apticular connections; loose complete spic- 
ules of this class have not been found. 
Microscleres: All of the following mi- 
croscleres, with exception of the short- 
rayed graphiocome noted below, are found 
in samples from both surfaces and the 
choanosome. The most common micro- 
sclere is a smooth, sharp-tipped form (Fig. 
10E) which can be regarded either as an 
oxyhexaster with very irregular secondary 


VOLUME 113, NUMBER 2 577 


Table 1.—Spicule dimensions of Heterotella pomponae holotype in zm; SD = standard deviation; n = number 
of measurements. 


Spicule type Length* + SD Range N Width + SD Range n 

Hexactin, thick sword 
distal ray 43° .24 8-101 SO 3 Se Ae «oa 
tangential ray Se. > 15 52-121 a0, SS) 2 Cl a5 3-o48- 25 
proximal ray 507, ='96 335-730 a0 Dy ae BD 

Hexactin, thin sword 
distal ray 90.25.17 35-118 50) (SOE OF VY 256 25 
tangential ray 106 + 18 67-141 50): SScteOGul 2788.) 25 
proximal ray 306.2 77 171-541 506 350 oP Oo 25 
Hexactin, regular ray Ist 281 64-318 50, 321. = aS. 2s 30 
Diactin, centrotylote S69) 257 320-1963 SUT yay 2 Eee. 3.8-12.3 50 
Diactin, sinuous — — —=/9 23.1 2'6.3'>' 9.1=40:2 - “50 
Oxyhexaster radius 50:5 + 6.8 B73-7hid 106. 20:04 Lt —eehy «25 
oxyhexaster secondary ray LSB. 45 5.8—28.7 100 120.3 OBS: 0 Zo 
Floricome radius Feel ae ce a 21644.6 357 54+1.0° 3474 25 
floricome centrum radius Ned = O8 5.49.0 100. ADE O83 lhieasS =o 
Sigmatocome radius |W ge Pike es 133—20'5) 6 100 0:6: 2502 O20 25 
sigmatocome centrum radius 6.8 + 0.8 5.1-9.5 100 — Ed 0.37 9106-16 (25 
Graphiocome centrum radius 82,209 6.3-11.1 IDO») (13:2 O3"°OS-19 25 
graphiocome long secondary ray 86 + 9 63-151 100° * 0.92.02 .01641:2"'" 25 
graphiocome short secondary ray 14.9 + 1.8 BIG —2-Or 100: 6) O6:2e10sike 04-09 125 


@ Radius for microscleres, ° Width of terminal flange. 


rays (preferred here) or an oxyhexactin with 
long spines. The secondary rays (or spines) 
number 1—4, most commonly 3, and ema- 
nate from an elongate swelling at their junc- 
tion. About 10% of these are without sec- 
ondary rays but bear the swelling seen in 
the astrose form. A typical floricome (Fig. 
10F) with usually 7 (6—9) terminal rays, 
bearing 3—5 claws on the terminal flange, is 
common. Although these occur in a wide 
size range, frequency analysis indicates 
these compose a single class. An uncom- 
monly occurring sigmatocome (Fig. 10G) 
bears 25—35 sigmoid terminals, uniform in 
length, from a single marginal whorl on 
each recurved primary capitulum. Graphio- 
come centra, with stumps of 30—50 terminal 
attachments scattered across the face of 
each discoid capitulum, (Fig. 10H) are un- 
common. The presumed raphide-like, long 
terminal rays are common on filters from 
all tissues, but the short versions are found 
only in gastral tissues. Intact graphiocomes 
have not been encountered. 
Etymology.—This species is named to 


honour Dr. Shirley Pomponi in recognition 
of her many years of invaluable contribu- 
tions to the knowledge of tropical marine 
biodiversity. 

Remarks.—The vary large number of 
shared characters between the new species 
and H. corbicula leaves little doubt that the 
new the West Indian form belongs to Het- 
erotella. Most of the differences between 
the two species can be attributed to their 
different stages of maturation. Only speci- 
men B of H. corbicula exhibits incipient 
spicule fusion in its lower portion, while the 
H. pomponae specimen has completely 
fused its lyssacine net into a rigid frame- 
work. The absence of free large choanoso- 
mal diactins and marginalia in H. pompon- 
ae are attributable to this terminal growth 
stage, where further body extension 
(growth) has ceased. Surface megascleres 
and microscleres are almost identical in 
form and size in both species. The dermalia 
are thicker (to 30 wm) and the accessory 
diactins longer (to 5 mm) in H. corbicula 
than in H. pomponae (to 6.7 wm and 2 mm 


578 


respectively). The distinctive oxyhexasters 
(oxyhexactins) are virtually identical in the 
two species. The most unambiguous differ- 
ence between the two is the presence of sig- 
matocomes in H. pomponae and their ab- 
sence in A. corbicula, confirmed by ex- 
amination of new spicule preparations of 
Ijima’s specimen A (the lectotype) and B 
(the nearly intact paralectotype) using filter 
techniques. Photographic records at HBOM 
suggest that H. pomponae is moderately 
common in the West Indian area. Younger 
specimens with incompletely fused skeletal 
networks are expected to be found in future 
collections. 


Acknowledgments 


I thank C. Adams for bringing the spec- 
imen to my attention, S. Pomponi and J. 
Reed for providing access to the specimen, 
photographs and data sheets, M. Kelly for 
providing transparencies and motivation, C. 
Lévi and K. Tabachnick for assisting in re- 
view of the Paris material, and the latter for 
his valuable comments on improving the 
manuscript. This study was supported by an 
operating grant from the Natural Sciences 
and Engineering Research Council of Can- 
ada. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Literature Cited 


Bowerbank, J. S. 1858. On the anatomy and physiol- 
ogy of the Spongiadae. Part 1: On the spic- 
ules.—Philosophical Transactions of the Royal 
Society of London 148:279-—332. 

. 1864. A Monograph of the British Spongia- 

dae, vol. 1. Ray Society, London, 290 pp. 

. 1867. On Alcyoncellum speciosum.—Proceed- 
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351-359. 

Gray, J. E. 1867. Notes on the arrangement of sponges, 
with the description of some new genera.—Pro- 
ceedings of the Zoological Society of London 
1867:492-558. 

Ijima, I. 1902. Studies on the Hexactinellida. Contri- 
bution II. (The genera Corbitella and Hetero- 
tella).—Journal of the College of Science, To- 
kyo Imperial University 17:1—34. 

Kirkpatrick, R. 1910. On Hexactinellida sponge spic- 
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Natural History (8)5:208—213. 

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hexactinellid sponges III. The taxonomic status 
of Hexactinellida within the Porifera.—Philo- 
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London B 301:419—428. 

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des Atlantischen Gebietes. Engelmann, Leipzig, 
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13:1—63. 


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CONTENTS 


Taxonomy and evolutionary relationships of Phillips’ small-eared shrew, Cryptotis phillipsii 
(Schaldach, 1966), from Oaxaca, Mexico (Mammalia: Insectivora: Soricidae) 

Neal Woodman and Robert M. Timm 

Two new species of flightless rails (Aves: Rallidae) from the Middle Pleistocene “crane fauna” of Bermuda 

Storrs L. Olson and David B. Wingate 

A new species of Anthias (Teleostei: Serranidae: Anthiinae) from the Galapagos Islands, with keys to 


Anthias and eastern Pacific Anthiinae William D. Anderson, Jr. and Carole C. Baldwin 
Description of two new blenniid fish species: Entomacrodus lemuria from the western Indian Ocean 
and E. williamsi from the western Pacific Ocean Victor G. Springer and Ronald A. Fricke 


Ammodytoides leptus, a new species of sand lance (Teleostei: Ammodytidae) from Pitcairn Island 
Bruce B. Collette and John E. Randall 
Neostrengeria binderi, a new species of pseudothelphusid crab from the eastern Andes of Colombia 


(Crustacea: Decapoda: Brachyura) Martha R. Campos 
Freshwater crabs (Brachyura: Potamoidea: Potamonautidae) from the rainforests of the Central African 
Republic, central Africa Neil Cumberlidge and Christopher B. Boyko 


The freshwater crabs of the Barbilla National Park, Costa Rica (Crustacea: Brachyura: Pseudothelphus- 
idae), with notes on the evolution of structures for spermatophore retention 
Gilberto Rodriguez and Ingemar Hedstr6m 
Acanthilia, a new genus of leucosioid crabs (Crustacea: Brachyura) from the Atlantic coast of the 
Americas Bella S. Galil 
Cambarus (Cambarus) davidi, a new species of crayfish (Decapoda: Cambaridae) from North Carolina 
John E. Cooper 
Two new species of Hyalella (Crustacea: Amphipoda: Hyalellidae) from Death Valley National Park, 
California, U.S.A. Adam J. Baldinger, William D. Shepard, and Doug L. Threloff 
Caecidotea cumberlandensis, a new species of troglobitic isopod from Virginia, with new records of 
other subterranean Caecidotea (Crustacea: Isopoda: Asellidae) Julian J. Lewis 
Euphilomedes cooki, a new species of myodocopid ostracode from Moreton Bay, SE Queensland, 
Australia Elizabeth Harrison-Nelson and Louis S. Kornicker 
A new record of Cornechiniscus madagascariensis Maucci, 1993 (Tardigrada: Echiniscidae) from 
India Wataru Abe and Masatsune Takeda 
Ophryotrocha lipscombae, a new species and a possible connection between ctenognath and labidognath- 
prionognath eunicean worms (Polychaeta) Hua Lu and Kristian Fauchald 
Aphrodita bisetosa (Polychaeta: Aphroditidae), a new species of sea mouse from the southeastern Pacific 
Ocean off central Chile Nicolas Rozbaczylo and Elba Canahuire 
Exogone breviantennata Hartmann-Schréder, 1959 (characters emended) (Annelida: Polychaeta: Syllidae), 
a new record for the Bahamas with a key to selected Exogone species 
Robert Zottoli and Charlene D. Long 
Podarke aberrans Webster & Benedict, 1887 - resolution, with descriptions of two new species in the 
genus Microphthalmus (Annelida: Polychaeta) Nathan W. Riser 
Parapionosyllis cabezali, a new species of Exogoninae (Polychaeta: Syllidae) from Spain 
Julio Parapar, Guillermo San Martin, and Juan Moreira 
Paralarval gonatid squids (Cephalopoda: Oegopsida) from the Mid-North Atlantic Ocean 
Luisa I. Falcon, Michael Vecchione, and Clyde F. E. Roper 
Parabuccinum, a new genus of Magellanic buccinulid (Gastropoda: Neogastropoda), with a description 


of a new species M. G. Harasewych, Yuri I. Kantor, and Katrin Linse 
Porites arnaudi, a new species of stony coral (Anthozoa: Scleractinia: Poritidae) from oceanic islands 
of the eastern Pacific Ocean Héctor Reyes-Bonilla and Juan P. Carricart-Ganivet 


A new species of Heterotella (Porifera: Hexactinellida: Euplectellidae) from the West Indies 
Henry M. Reiswig 


Front cover—from this issue, p. 375. 


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