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PROCEEDINGS
of the
Biological Society of
Washington
VOLUME 87
1974
WASHINGTON
PRINTED FOR THE SOCIETY
EDITOR
AvustTIN B. WILLIAMS
ASSOCIATE EDITORS
Classical Languages Invertebrates
GrorcE C. STEYSKAL THomas FE. BOWMAN
Insects Vertebrates
RosertT D. GORDON Victor G. SPRINGER
and
WILLIAM R. TAayYLor
All correspondence should be addressed to the
Biological Society of Washington, Smithsonian Institution
Washington, D. C. 20560
ALLEN Press INC.
LAWRENCE, Kansas 66044
OFFICERS AND COUNCIL
of the
BIOLOGICAL SOCIETY OF WASHINGTON
FOR 1974-1975
OFFICERS
President
THOMAS E. BOWMAN
Vice President
CLYDE J. JONES
Secretary
LOUIS S. KORNICKER
Treasurer
OLIVER S. FLINT
Custodian of Publications
JOHN H. MILES
COUNCIL
Elected Members
RICHARD C. BANKS FENNER A. CHACE, JR.
ISABEL C. CANET CLAYTON E. RAY
PAUL J. SPANGLER
iu
TABLE OF CONTENTS
Volume 87
Baumann, Richard W. What is Alloperla imbecilla (Say)?
Designation of a neotype, and a new Alloperla from eastern
North America (Plecoptera: Chloroperlidae) —.____
Bogan, Michael A. Identification of Myotis californicus and M.
leibi imy southwestern North Ament cays esas ee
Bowman, Thomas E. The “sea flea” Dolobrotus mardeni n.
gen., n. sp., a deep-water American lobster bait scavenger
(Amphipoda: Eusinridae)) 250 2
Bowman, Thomas E., and George A. Schultz. The isopod
crustacean genus Munnogonium George and Stromberg, 1968
(Munnidae, “Asellota’) (2 ee Se
Child, C. Allan. Hedgpethius tridentatus, a new genus and
new species, and other Pycnogonida from Key West, Florida,
UGA co AE cL ia eR ee
Collette, Bruce B. Potamorrhaphis petersi, a new species of
freshwater needlefish (Belonidae) from the upper Orinoco
and. Rio, Negro. 22): we
Collette, Bruce B. A review of the coral toadfishes of the genus
Sanopus with descriptions of two new species from Cozumel
Island, México. 0000 on eh
Cowan, Richard S. A revision of the genus Bocoa (Caesal-
pinioideae-Swartzieae)) 222 8 Ee
Cressey, Roger F. A redescription of Hermilius pyriventris
Heller (Copepoda: Caligoida) with the first description
of the’ male eee ee
Dawson, C. E. Pseudophallus brasiliensis (Pisces: Syngnath-
idae), a new freshwater pipefish from Brazil
Deevey, Georgiana B. Pelagic ostracods collected on Hudson
70 between the equator and 55°S in the Atlantic __.....___-_-__
Emerson, K. C., and Roger D. Price. A new species of Tricho-
dectes (Mallophaga: Trichodectidae) from the yellow-
throated marten (Martes flavigula) ~--- = ees
Flint, Oliver S., Jr. Studies of neotropical caddisflies, XIX:
the genus Cailloma (Trichoptera: Rhyacophilidae) _____
Fraser, Thomas H. Redescription of the cardinal fish Apogon
endekataenia Bleeker (Apogonidae), with comments on pre-
vious usage of the name.) Ue
Gomon, Martin F. A new eastern Pacific labrid (Pisces),
Decadon melasma, a geminate species of the western Atlantic
Dy pellets: ise SEE)
257-264
49-56
129-138
265-272
493-500
31-40
185-204
235-244
405-410
351-380
77-80
473-484
3-10
205-216
Gopalakrishnan, T. C. A review of the copepod Scottocalanus
securifrons (T. Scott) and a note on its synonym Scolecithrix
cuneifrons Willey (Calanoida: Scolecithricidae) -----......
Gordon, Robert D. Additional notes on the genus Glaresis
m@oleoptera;,. Scarabaeidae) Leet su ee Neal
Herald, E. S., and C. E. Dawson. Micrognathus erugatus, a
new marine pipefish from Brazil (Pisces: Syngnathidae) __
Heyer, W. Ronald. Vanzolinius, a new genus proposed for
Leptodactylus discodactylus (Amphibia, Leptodactylidae) __.
Heyer, W. Ronald, and M. Judith Diment. The karyotype of
Vanzolinius discodactylus and comments on usefulness of
karyotypes in determining relationships in the Leptodactylus-
complex (Amphibia, Leptodactylidae) —___________
Hoffman, Richard L. A new polydesmid milliped from the
southern Appalachians, with remarks on the status of Dixi-
desmus and a proposed terminology for polydesmid gonopods
Holt, Perry C. An emendation of the genus Triannulata Good-
night, 1940, with the assignment of Triannulata montana to
Cambarincola Ellis 1912 (Clitellata: Branchiobdellida) __
Humes, Arthur G. Cyclopoid copepods (Lichomolgidae) from
Sauconaceans in Madagascar 0 ae ee
imtonmations tor contributors 2 2
Jones, Meredith L. Brandtika asiatica new genus, new species,
from southeastern Asia and a redescription of Monroika afri-
cane (Monro) (Polychaeta: Sabellidae)
Kingsolver, John M., and Donald R. Whitehead. Biosystem-
atics of Central American species of Ctenocolum, a new
genus of seed beetles (Coleoptera: Bruchidae) _...
Light, William J. Occurrence of the Atlantic maldanid Asychis
elongata (Annelida, Polychaeta) in San Francisco Bay, with
BOUMnentSsONMItSeSyNODYIMNY) 22.2) Oe ee ee
Light, William J. A new species of Pseudopolydora (Poly-
chaeta, Spionidae) from the Solomon Islands __.......______.
Lynch, John D. A new species of Eleutherodaciylus (Am-
phibia: Leptodactylidae) from the Pacific lowlands of
Ecuador
McLaughlin, Patsy A., and Julie H. Brock. A new species of
hermit crab of the genus Nematopagurus (Crustacea:
Decapoda: Paguridae)) from Hawai
Miller, Robert Rush. Cichlasoma regani, a new species of cich-
lid fish from the Rio Coatzacoalcos basin, México _....._._
Novak, Allen, and Michael Salmon. Uca panacea, a new spe-
cies of fiddler crab from the Gulf coast of the United States
(9)
273-282
91-94
27-30
81-90
327-336
345-350
57-72
411-438
|-2
217-230
283-312
175-184
389-394
381-388
245-256
465-472,
313-326
Olson, Storrs L. A new species of Nesotrochis from Hispaniola,
with notes on other fossil rails from the West Indies (Aves:
Beallichae,): <2 2 5 ocala lie el pce eet el oes ee
Orejas-Miranda, Braulio R., and George R. Zug. A new tricolor
Leptotyphlops (Reptilia: Serpentes) from Peru ____--
Robbins, C. Brian. Comments on the taxonomy of the West
African Taterillus (Rodentia: Cricetidae) with the descrip-
tion, Of anew sSDECIES 22k eae Oke ee
Ross, Charles A., and Franklin D. Ross. Caudal scalation of
Central American Crocodylus ____. eee me BE mererree Mere
Schwartz, Albert, and Orlando H. Garrido. A new Cuban
species of Sphaerodactylus (Gekkonidae) of the nigropunc-
tatus:; COMpIeRX, feces Ee ee
Serafy, D. Keith. A new heart-urchin, Araeolampas atlantica,
new genus, new species, from the North Atlantic and notes
on the closely related Homolampas
Steyskal, George C. Euxesta mazorca, new species, associated
with ears of maize in South America (Diptera, Otitidae) __.
Thomas, Richard. A new species of Typhlops (Serpentes:
Typhlopidae)) (from Hispaniolay 2
Vagvolgyi, Joseph. Eight new Ashmunellas from the south-
western United States (Pulmonata: Polygyridae) —
Williams, Austin B. Allactaea lithostrota, a new genus and
species of crab (Decapoda: Xanthidae) from North Carolina,
USAGI ie
Williams, Austin B. Two new axiids (Crustacea: Decapoda:
Thalassinidea: Calocaris) from North Carolina and the
Straits of Florida 2.2. 2)
Williams, Austin B. A new species of Hypsophrys (Decapoda:
Homolidae) from the Straits of Florida, with notes on related
eras 30 DSN te
439-450
167-173
395-404
231-234
337-343
BIOLOGICAL SOCIETY OF WASHINGTON
PROCEEDINGS
1046th Meeting—13 March 1974
NINETY-FIFTH ANNUAL MEETING
The meeting was called to order by Vice President Thomas E. Bowman
in the absence of President Victor Springer at 2:10 p.m. in Room 43
of the National Museum of Natural History.
Reading of the minutes of the previous annual meeting was dis-
pensed with, since they were printed in the Biological Society of Wash-
ington Proceedings.
Treasurer Oliver Flint read the Financial Report. He thanked Mau-
reen Downey for serving as Acting Treasurer for the past 6 months and
for developing figures for the report. In answer to a question as to
why an endowment fund repayment had not been made during the
past year, Treasurer Flint said that it was an oversight and that $1700
is to be repaid in 1974.
Clayton Ray asked the Treasurer what percent of page-charges for
papers published in the Proceedings came from the Smithsonian In-
stitution. Treasurer Flint said that about one-half, but that some pay-
ments of the previous year were not yet paid. In answer to a question
from Stanwyn Shetler, Treasurer Flint said that page-charges were
requested from all authors, but that payment was not a condition for
publication.
Daniel Cohen, Chairman of the Financial Committee, stated that
the Financial Report had been audited and found to be correct. A
motion made to accept the report was seconded and passed. He then
presented the budget for 1974.
Austin Williams, Chairman of the Editorial Committee, presented a
report stating, Volume 86 of the Proceedings for 1973 consisted of 45
papers totalling 524 pages. The first issue of Volume 87, consisting
of 14 papers, is now in galley proof. Ten of these are paid papers.
Twelve papers for the second issue have been received and are under
review. He also said that instructions entitled, “Information for Con-
tributors,” will be included in Volume 87. George Watson suggested
that the instructions not be republished every year, but that enough
reprints should be printed to last about 5 years.
Secretary Louis Kornicker gave results of the election for Officers
and Council members for 1974: President, Thomas E. Bowman; Vice
President, Clyde J. Jones; Secretary, Louis S. Kornicker; Treasurer,
Oliver S. Flint; Members of the Council, Richard C. Banks, Isabel C.
Canet, Fenner A. Chace, Clayton E. Ray, and Paul J. Spangler.
vik
Secretary Louis Kornicker gave a report on membership (see Ap-
pendix in Treasurer’s Report). Daniel Cohen asked how a new life
member could have been elected since that category had been elimi-
nated by the council some years back. He suggested that the Secretary
check minutes of old meetings to ascertain whether the category of
Life Member was still extant.
Vice President Bowman reported that the Council had 3 meetings
during the past year and that a proposed Bicentennial Symposium had
been cancelled because of being unable to obtain someone willing to
organize it, and because of general lack of interest by possible partic-
ipants approached. The subject of the symposium was to have been
concerned with biological, geological and ecological changes that have
taken place in the Washington area since 1776. He also announced that
“The Panamic Biota” volume is being reprinted because of demand
and should be available shortly. A policy statement concerning a pro-
posed Sea-level Canal in Panama was distributed to other societies and
was published in Science. A letter concerning this statement was re-
ceived from the President of the Panama Canal Company and is avail-
able to interested members.
Marian Pettibone made a motion thanking Officers and Council mem-
bers for their efforts on behalf of the Society during the past year, and
also Paul Spangler, who had served as Editor of the Proceedings until
it was taken over by Austin Williams in the fall. The motion was sec-
onded and passed.
Vice President Bowman then turned the meeting over to incoming
President Bowman.
A motion to adjourn was seconded and passed.
Louis S. Kornicker
Secretary
iii
~ 0673
Vol. 87, No. 0, pp. 1-2 25 April 1974
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
INFORMATION FOR CONTRIBUTORS
1. Content. The Proceedings of the Biological Society of Wash-
ington contains papers bearing on systematics in the biological
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The establishment of new taxa must conform with the require-
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Authors are expected to be familiar with these codes and to comply
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0—Proc. Biot. Soc. WasuH., Vou. 87, 1974 (aL)
2 Proceedings of the Biological Society of Washington
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mr C6 7/3
Vol. 87, No. 1, pp. 3-10 25 April 1974
PROCEEDINGS
ORMInE
BIOLOGICAL SOCIETY OF WASHINGTON
REDESCRIPTION OF THE CARDINAL FISH APOGON
ENDEKATAENIA BLEEKER (APOGONIDAE), WITH
COMMENTS ON PREVIOUS USAGE OF THE NAME
By Tuomas H. FRASER!
J. L. B. Smith Institute of Ichthyology,
Rhodes University, Grahamstown, South Africa
In 1852 Pieter Bleeker described Apogon endekataenia from
a single specimen (57 mm total length) taken at Banka in
Indonesia. Subsequently, fifteen other specimens were added
to the jar containing the type. According to the Auction
Catalogue (Hubrecht, 1879) in which Bleeker’s fish collection
was offered for sale, ten specimens of A. endekataenia were in
the A series, two specimens each in the B, C and D series and
none in the E series. Whitehead et al. (1966: 7-14) indicate
some basic procedures involved in attempting to determine
which specimens are type material in the Bleeker collections.
In the case of A. endekataenia the criteria utilized were total
length and the remains of faded color pattern. Fortunately,
only one specimen (55 mm total length) was close to the 57
mm length given by Bleeker. The others were of various sizes,
the nearest three 45, 63 and 64 mm total length. (I cannot
account for the presence of two additional specimens in the
jar unless a counting or typographic error was made by
Hubrecht.) The 55 mm specimen is considered to represent
Bleeker’s original specimen from Banka. This agrees with
the opinion of M. Boeseman cited in Smith (1961).
The identification of the putative holotype fundamentally
changes the concept of Apogon endekataenia; most past records
are considered to be erroneous or doubtful identifications.
The specimens subsequently added to the type jar were of
1 Present address: Division of Fishes, National Museum of Natural History, Wash-
ington, D. C. 20560 U.S.A.
1—Proc. Biot. Soc. WaAsu., Vou. 87, 1974 (3)
4 Proceedings of the Biological Society of Washington
two species and may have been the source of some errors.
Descriptions in all major faunal reviews examined indicate
other species, or are composites of two or more species
(Bleeker, 1874, 1873-76, Weber and de Beaufort, 1929; Fowler
and Bean, 1930; Smith, 1961). Lachner (1953) did not reach
a decision concerning the identity of Apogon endekataenia
but he noted that none of the specimens available to him con-
formed to the figure in Bleeker’s Atlas (1873-76). A re-
description of the species is necessary and will contribute to
the eventual resolution of the dark-striped complex of species
in Apogon.
I thank M. Boeseman, Rijksmuseum van Natuurlijke His-
torie (RMNH) for permission to examine Bleeker’s material
and for his co-operation and valuable help during my stay
at Leiden. A. C. Wheeler made possible the loan of speci-
mens at the British Museum (Natural History) (BMNH).
E. A. Lachner and M. M. Smith critically read a draft of the
manuscript. Funds for the taxonomic study of fishes and attend-
ing aspects were provided by the South African Council for In-
dustrial and Scientific Research and by Rhodes University.
Counts and measurements were made according to Hubbs
and Lagler (1958) except that the body depth was taken from
the dorsal-fin origin to the pelvic-fin base. Measurements
were made with dial calipers. The last dorsal ray and last
anal ray, each split to its base, was counted as one element.
The gillraker at the angle was included in the count for the
lower arch.
Apogon endekataenia Bleeker, 1852
Figure 1
Description: For general body shape see Figure 1. Range of pro-
portions (as percentages of standard lengths, putative holotype values in
parentheses): body depth (33) 33-36; head length (36) 35-38; eye
length (12) 11-12; snout length (8) 7-9; bony interorbital width (7)
6-7; upper jaw length (18) 17-18; caudal peduncle depth (14) 14-16;
caudal peduncle length (25) 22-26; first dorsal spine length (4) 1-4;
second dorsal spine length (12) 9-12; third dorsal spine length 19-23;
fourth dorsal spine length 16-20; last dorsal spine length (17) 15-17;
first anal spine length (2) 2; second anal spine length (13) 12-13;
pectoral fin length (24) 23-25; pelvic fin length (21) 20-22.
Meristic counts and ranges for nine specimens examined: dorsal fin
Redescription of Cardinal Fish 5
Fic. 1. A. Apogon endekataenia, illustration from Bleeker 1873-—
1876. B. Apogon endekataenia, putative holotype, RMNH 5593 (43.9
mm SL). C. Apogon endekataenia, BMNH 1892.9.2.18—-20, one of three.
VII-I, 9; anal fin II, 8; pectoral fin 14; pelvic fin I, 5; well-developed
gillrakers 13-15 (2-3 + 11-12), usually 13-14, including rudiments
18-21 (2-4 + 2-3; 11-12 + 2-4), putative holotype 3 + 3-12 +4 3;
pored lateral-line scales (25) 24-25; longitudinal scale rows above
lateral line (25) 24-25; transverse scale rows above lateral line 2; trans-
verse scale rows below lateral line (6) 6-7; median predorsal scales (3)
38—4; cireumpeduncular scale rows 12 (5 + 2+ 5).
6 Proceedings of the Biological Society of Washington
Teeth in villiform bands on premaxilla and dentary; one or two rows
on vomer and palatine; none on ectopterygoid, endopterygoid or basihyal.
Vertebrae 10 + 14. Five free hypurals, one pair of slender uroneurals,
three epurals, a free parhypural. Three predorsals, two spines on first
dorsal pterygiophore. Seven branchiostegal rays. Basisphenoid present.
Supramaxilla absent.
Life colors unknown. In 70% ethanol six dark, longitudinal stripes
present from dorsal-fin base to abdomen; large round basicaudal spot
slightly off center (dorsally) of lateral line (Figure 1): 1. median stripe
on head splitting at dorsal fin and extending along dorsum at base of
fins, fading on caudal peduncle; 2. stripe extending from snout along
occiput and nape paralleling above the lateral line and reaching nearly
to basicaudal spot; 3. post-ocular stripe from upper part of eye and just
below lateral line extending onto caudal peduncle; 4. median stripe ex-
tending from eye above pectoral fin along side nearly to basicaudal spot;
5. stripe extending from lower part of eye and head through pectoral fin
along side nearly to basicaudal spot; 6. stripe extending from lower jaw
below pectoral fin along abdomen and onto last several anal-fin rays.
Complete dark stripe present in basal portion of soft dorsal fin but no
counterpart in soft anal fin. Peritoneum silvery, intestine black.
Distribution: Discounting the doubtful and erroneous reports, Apogon
endekataenia is known from Borneo and Banka Island, near Sumatra, and
recently I have seen material from Thailand at the California Academy of
Sciences. Fowler (1937: 232-234, Fig. 23) identified two specimens
from Bangkok as Apogon robustus (Smith and Radcliffe, 1911); how-
ever, the illustration appears similar to A. endekataenia but with extra
stripes. It has not been reliably reported in the western Indian or cen-
tral Pacific Oceans.
Material examined: Putative holotype. RMNH 5593, 43.9 mm SL,
55 mm total length, Banka Island. Other material. BMNH 1892.9.2.18—20.
(3 specimens, 52.1-62.1 mm SL), Borneo, X-ray. RMNH 23966 (5,
51.5-67.6 mm SL), East Indies.
Remarks: Apogon endekataenia differs from other dark-striped Apogon
(Nectamia) in possessing the following combination of characters: a
large, round, basicaudal spot, no dark stripe in the anterior part of the
anal fin, and a long post-ocular stripe. Smith (1961: 400) believed that
Bleeker’s illustration (1873-76: Pl. 32, fig. 2) was inaccurate. The
illustration and putative holotype (Figs. 1A and B) agree better than
figures by other authors of specimens purported to be A. endekataenia.
The putative holotype also agrees with Bleeker’s original description.
Of the 18 specimens examined in the Bleeker collections, five speci-
mens in RMNH 23966 are conspecific with the putative holotype and
one other specimen in RMNH 23966 was unidentifiable; two specimens
in RMNH 23968 may be conspecific with the type but are too poorly
preserved to be certain; and all nine in RMNH 23967 are a different
species, probably Apogon cookii Macleay, 1881.
Smith (1961) experienced difficulty in separating closely related spe-
Redescription of Cardinal Fish
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§ Proceedings of the Biological Society of Washington
cies in the dark-striped complex of Apogon and believed that he was
dealing with a polychromic species. He did not see the putative holo-
type but did see two specimens (out of five) which are here con-
sidered conspecific with the holotype. Nevertheless, he believed these
specimens were identical with his relatively fresh material. None of the
material so identified at the J. L. B. Smith Institute is Bleeker’s species
but rather Apogon cookii and Apogon taeniophorus Regan, 1908, common
species in the western Indian Ocean (see Table 1 for comparisons). The
illustrations in Smith (1961) are of A. cookii. Both of these species also
appear to be widespread in the eastern Indian and western Pacific
Oceans, but have been misidentified or given junior synonyms, a prob-
lem that will be discussed elsewhere.
Weber and de Beaufort (1929) probably based their description on
the collection of mixed species at RMNH although it is not so stated.
Their description is insufficient to determine whether more than one
species was included.
Fowler and Bean (1930) examined only the type of A. wilsoni
(Fowler, 1918), a different species.
Some of the dark-striped, seven-spined species of Apogon have re-
ceived varied treatment: consideration as polychromic forms of the
same species (Jordan and Seale, 1906; Smith, 1961), as subspecies
(McCulloch, 1915), and as different species (Radcliffe, 1911; Lachner,
1953). McCulloch’s treatment would seem to be least acceptable on
practical and theoretical grounds because several of these supposed
subspecies are sympatric over a wide geographic range. The ideas of
either a single variable species or several different species are more
useful hypotheses to test at this stage. Radcliffe (1911) pointed out the
changes in number of stripes in Apogon multilineatus (Bleeker, 1865)
with growth, and it appears that stripes and the basicaudal spot may be
altered with growth in other species. These changes are not described
for other striped species and must be investigated.
LITERATURE CITED
BLEEKER, P. 1852. Bijdrage tot de kennis der ichthyologische fauna
van het eiland Banka. Nat. Tijdschr. Ned. Ind. 3:443—460.
1873-76. Percoides I. Priacanthiformes, Serraniformes, Gram-
misteiformes, Percaeformes, Datniaeformes. Atlas Ichthyol.
Indes. Orient. Neerland. 7:1—126, pls. 249-320.
—. 1874. Revision des especes Indo-Archipelagiques du groupe
des Apogonini. Verh. Holl. Maatsche. (3)2(1):1-82.
Fowter, H. W. 1937. Zoological Results of the Third De Schauensee
Siamese Expedition. Part VIII—Fishes obtained in 1936.
Proc. Acad. Nat. Sci. Phila. 89:125-264, 300 figs.
, AND B. A. BEAN. 1930. The fishes of the families Amiidae,
Chandidae, Duleidae, and Serranidae, obtained by the United
States Bureau of Fishes Steamer Albatross in 1907 to 1910,
Redescription of Cardinal Fish 9
chiefly in the Philippine Islands and adjacent seas. Bull. U. S.
Nat. Mus. 100( 10) :i—ix, 1-334, 27 figs.
Husss, C. L., anp K. F. Lacter. 1958. Fishes of the Great Lakes
Region. Cranbrook Inst. Sci. (revised ed.) Bull. 26:i—lxi, 1-213,
44 pls., 251 figs.
Huprecur, A. A. W. 1879. Catalogue des collections formees et
laissees par M. P. Bleeker. Leiden iv + 71 pp.
JorDAN, D. S., AND A. SEALE. 1906. The fishes of Samoa. Description
of the species found in the Archipelago with a provisional
checklist of the fishes of Oceania. Bull. Bur. Fish. 25: 175-488,
pls. 23-53, 111 figs.
LacHNER, E. A. 1953. Family Apogonidae: cardinal fishes. (In:
Schultz, L. P., Fishes of the Marshall and Marianas Islands).
Bull. U. S. Nat. Mus. 202(1):412—498, figs. 68-84, tabs. 38-44.
McCuttocu, A. R. 1915. Report on some fishes obtained by the
F.1I.S. Endeavour on the coast of Queensland, New South Wales,
Victoria, Tasmania, South and South Western Australia. Biol.
Res. Endeavour 3(3):97—170, pls. 13-37.
RapcuiFFE, L. 1911. Notes on some fishes of the genus Amia, family
of Cheilodipteridae, with descriptions of four new species from
the Philippine Islands. Proc. U. S. Nat. Mus. 41(1853): 245-
261, pls. 20-25, 3 figs.
SmitH, J. L. B. 1961. Fishes of the family Apogonidae of the western
Indian Ocean and the Red Sea. Ichthyol. Bull. Rhodes Univ.
29:373-418, pls. 46-52, 11 figs.
Weser, M. C. W., anp L. F. pE BeAurorr. 1929. The fishes of the
Indo-Australian Archipelago. E. J. Brill, Leiden 5:i-xiv, 1-458,
98 figs.
WHITEHEAD, P. J. P., M. BOESEMAN, AND A. C. WHEELER. 1966. The
types of Bleeker’ Indo-Pacific elopoid and clupeoid fishes.
Zool. Verhand. 84:1—152, 19 pls., 3 tabs.
10 Proceedings of the Biological Society of Washington
ws O63
Vol. 87, No. 2, pp. 11-18 25 April 1974
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
A NEW SPECIES OF TYPHLOPS (SERPENTES:
TYPHLOPIDAE) FROM HISPANIOLA
By RicHARD THOMAS
Museum of Zoology, Louisiana State University,
Baton Rouge, Louisiana 70808
Since the description (Thomas, 1965) of Typhlops syntherus,
it has become evident to me that the Hispaniolan Typhlops,
then regarded as T. lumbricalis Linnaeus, actually comprises
two species. One of these, known from only a few localities in
eastern Hispaniola but better represented from the Cul de
Sac Plain of Haiti, differs in no major morphological feature,
other than size, from the Cuban and Bahamian populations
known as Typhlops lumbricalis. The other species, un-
described, appears to be restricted to the southwestern
quadrant of Hispaniola.
Aside from three old specimens, the hypodigm of this un-
described species was obtained through the sponsorship and
efforts of Albert Schwartz (ASFS designates the Albert
Schwartz Field Series) and of Ernest E. Williams of the
Museum of Comparative Zoology (MCZ). To Albert Schwartz
I am greatly indebted for his support of my own efforts in
the field and laboratory leading to the discovery of this new
form. I also thank Lewis D. Ober, who generously provided
some crucial specimens of T. lumbricalis (LDO designates
the Lewis D. Ober private collection) and Douglas A. Ross-
man for his careful critical reading of the manuscript. For the
loan of specimens used in this study I wish to thank Richard
G. Zweifel and George R. Foley of the American Museum of
Natural History (AMNH), Edmond G. Malnate of the Acad-
emy of Natural Sciences of Philadelphia (ANSP), Alice G. C.
Grandison of the British Museum (Natural History) (BMNH),
Neil D. Richmond and Clarence J. McCoy of the Carnegie
2—-Proc. Biot. Soc. WasuH., Vou. 87, 1974 (11)
12 Proceedings of the Biological Society of Washington
Museum (CM), Ernest E. Williams of the Museum of Com-
parative Zoology (MCZ), and the late James A. Peters and
George R. Zug of the National Museum of Natural History
(USNM). Without the capable field work of Robert K.
Bobilin, Patricia A. Adams, Ronald F. Klinikowski, Mark D.
Lavrich, and David C. Leber so extensive a series of the new
species would not have been collected. Material obtained
from 1968 through 1971 was collected under National Science
Foundation Grants GB 7977 and B-023603 to Albert Schwartz.
The undescribed species is the sixth known species of
Typhlops from Hispaniola; in reference to this fact I propose
the name:
Typhlops hectus new species
Holotype: MCZ 81149, taken at Martineau, ca. 9 km (airline) W
Jérémie, Dépt. du Sud, Haiti, 12 March 1966 by Richard Thomas.
Original number ASFS V9145.
Paratypes: HAITI. Derr. pu Sup: ASFS V9276-77, ca. 10 km WSW
Moron, 475 m, 13 March 1966, R. Thomas; MCZ 74907-14, Marfranc
near Jérémie, 27 December 1966, D. Hill and F. Vuilleumier; MCZ
64779, Carrefour Sanon near Jérémie, 14 December 1960, Luc and George
Whiteman; ASFS V9147, Jérémie, 12 March 1966, native; ASFS V9495,
Jérémie, 19 March 1966, native; ASFS V9601-12, Jérémie, 21 March
1966, natives; ASFS V25284, beach area within 1 km E Jérémie, 1 July
1971, Elie Joseph; ASFS V9389, 2 km SE Jérémie, 15 March 1966, native;
MCZ 64773-78, *Place Négre near Jérémie, 14-15 December 1960, Luc
and George Whiteman; MCZ 70044, *La Source near Jérémie, Decem-
ber 1962, George Whiteman; MCZ 64780, *Laye near Jérémie, 14-15 De-
cember 1960, Luc and George Whiteman; MCZ 70045, Tosia near (about
33 km SW) Jérémie, December 1962; ASFS V9348, ca. 5 km (airline)
SE Marché Léon, 670 m, 15 March 1966, R. Thomas; ASFS V9514—16,
ca. 7.5 km (airline) SSE Roseaux, est. 2 km W La Bastille, 20 March
1966, R. Thomas; ASFS V9519, ca. 3 km (airline) SW Corail, 20 March
1966, R. Thomas; MCZ 25552, Ile Grande Cayemite; ASFS V26459, Ile
Grande Cayemite, vicinity of Anse 4 Macon, 6 August 1971, native; ASFS
X3070, Camp Perrin, 26 July 1962, native; USNM 157928—29, ASFS
X3141-52, Camp Perrin, 29 July 1962, native; MCZ R-123873-74,
Marceline; ASFS V26251-53, Fond des Négres, 1 August 1971, R.
Thomas; ASFS V26223, Paillant, 548 m, 31 July 1971, R. Thomas; MCZ
66323-24, CM 37953, *Butéte near Miragodne, 16 August 1961, Luc
Whiteman; MCZ 66325, *Mingrette near Miragodne, August 1961, Luc
Whiteman; MCZ 66326-28, CM 37921—22, *Pemel near Miragoane, 14
August 1961, Luc Whiteman. DEPT. DE L’Oursr: ASFS V982]1, ca. 2.4
km S Trouin, 243 m, 29 March 1966, R. Thomas; ASFS V8370-7]1, ca.
A New Typhlops from Hispaniola 13
Fic. 1. A, dorsal and lateral views of the head of ASFS V9604 (T.
hectus); B, dorsal and lateral views of the head of ASFS V8395 (T.
lumbricalis). Bar equals one mm.
7 km (airline) W Pétionville, N versant Morne lH6pital, ca. 609 m, 1
March 1966, R. Thomas; ASFS V8375, 9.6 km (road) W Pétionville,
N versant More I|’H6pital, 822 m, 1 March 1966, native.
Non-paratypic specimens: HAITI. Dtpr. pE L’Ourst: ASFS V8295-
96, ca. 7 km SE Mirebalais; ASFS V26542—48, V26577, 3.55 km SW
Lascahobas, 274 m, 15 August 1971, native. REPUBLICA DOMINI-
CANA. Prov. SAn Juan: ASFS V512-15, Rio Arriba del Norte; ASFS
V21567, 4 km N Sabaneta; ASFS V396—98, 7 km W Vallejuelo, 792 m.
Prov. SAN RAFAEL: ASFS V21570-71, 1 km E Hondo Valle. Prov.
INDEPENDENCIA: ASFS V20924, 15 km W Puerto Escondido; AMNH
41265-66, Duverge. Prov. BaraHona: ASFS V30493, El Iguito, 2.6
km NE Fondo Negro; ASFS V35641, El Iguito, 3.1 km NE Fondo Negro;
AMNH 51496, above Delmonte’s Finca (near Barahona).
Localities indicated with an asterisk are not mapped; it is not certain
whether the unmapped localities reckoned from Miragoane are in the
Dépt. du Sud or the Dépt. de ]’Ouest.
Definition: A species of Typhlops having the preocular in contact
with upper labial three (as opposed to contacting upper labials two and
14 Proceedings of the Biological Society of Washington
three), two postoculars, a primary scale row number of 20 with reduction
to 18 rows absent or occurring far posteriorly, and high mid-dorsal scale
counts (284-328). Further distinguished by a clavate rostral shape
(narrow, posteriorly expanded and blunt); the posterior nasals being
roughly parallel sided, not divergent in dorsal aspect; and the preocular
being sharply pointed anteriorly (Fig. 1).
Distribution (Fig. 2): Southwestern Hispaniola south of the Cul
de Sac-Valle de Neiba Plain (except for the Barahona Peninsula) and
the southwestern part of the north island (that portion of Hispaniola
north of the Cul de Sac-Valle de Neiba Plain). One certain record for
the Cul de Sac-Valle de Neiba Plain (extreme eastern part).
Description of holotype: Total length 192 mm, tail length 4.6 mm;
307 mid-dorsal scales between rostral and caudal spine, 295 midventral
scales posterior to mental and including last scale of anal flap; 20 scale
rows anteriorly, reducing to 19 rows at ventral scale 277 and to 18 at
ventral scale 279 (with some redividing and re-fusing of scale rows
before final fusion); 2 postoculars and one parietal on each side. Colora-
tion pale gray dorsally, fading out on lower sides to become unpigmented
ventrally.
Variation: The paratypes vary in total length from 95 to 230 mm and
in mid-dorsal scale counts from 284 to 324. The modal primary scale
row number is 20 reducing to 18 at the level of midventral scale 194 to
288; 46 specimens (60 percent) have no reduction. A number of speci-
mens show reduction abnormalities shortly anterior to the vent: the first
reduction step (20 to 19 rows) may be made without the final (19 to
18 rows), or irregular fusion and dividing of the median and paramedian
ventral rows may occur. Three specimens have 22 scale rows initially
with reduction to 20 rows occurring from 35 to 131 ventrals posterior to
the mental. A few have short regions of 22 scale rows anteriorly due to
irregular division and fusion, and a few others have 21 scale rows for
a short distance in the neck region. There are from 5 to 15 (mode 8-10)
fewer ventrals than middorsals. Postoculars are 2/2 (66 specimens), 2/3
or 3/2 (2), or 3/3 (1); parietals are 1/1 (16), 1/2 or 2/1 (20), or 2/2
(25). Coloration is pale gray to tan but varies somewhat in intensity of
pigmentation. The coloration may be bicolor with the transition from the
pigmented dorsum to the unpigmented venter occurring in a narrow
mid-lateral zone of one or two scale rows; or the pigmentation may extend
well onto the venter, occasional individuals having almost entirely pig-
mented venters with irregular non-pigmented areas occurring along the
midventral region. In the more extensively pigmented individuals the
intensity of pigmentation gradually decreases ventrally. Some bicolor
specimens have an indistinct collar of dark pigment across the throat.
Twenty-four specimens from several scattered localities north of the
Cul de Sac-Valle de Neiba Plain plus six from the eastern portion of the
south island and the Valle de Neiba agree with Typhlops hectus in the
configuration of the rostral-nasal complex, in having pointed preoculars,
high longitudinal scale counts, and in having the reduction from 20 to 18
15
A New Typhlops from Hispaniola
‘UIR[ VQION Op 2I[BA-9VS 9p [ND oy} Jo spurl] oyeurrxoidde oy} soyeorpul SurTyoyeyY [v}I0A OY, “E ‘Sly UL poydeis snzoay
ordAyeivd-uou Jo sojdures “e—-v sioqjey, ‘sngzoay ordAyered-uou 10} soiyeooy, ‘sojsueLy MOT[OY ‘snzoay oidAyered 10J sotppeooy, oye
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16 Proceedings of the Biological Society of Washington
240 260 280 300 320
Fic. 3. Upper graph: Variation in reduction level (ventral scales
posterior to mental). Range only is shown for samples of non-paratypic
hectus (a-e); line histograms (smallest vertical unit represents one
individual) are given for lumbricalis (L.) and paratypic hectus. The
uppermost horizontal line of a pair indicates range in level of final reduc-
tion step (19 to 18 scale rows); the lowermost line of a pair indicates
the range in initial reduction step (18 to 19 rows). Crosses on lower
range line for paratypic hectus indicate specimens that reduce to 19
but not to 18 scale rows; + 46 and + 1 indicate specimens that do not
reduce from 20 scale rows. Lower graph: Line histograms showing
variation in mid-dorsal scale counts of T. lumbricalis (L.) and T. hectus
(H); the smallest vertical unit represents one individual. Histograms
a—e show counts for geographic samples of non-paratypic hectus indi-
cated in Fig. 2. The dashed line in both upper and lower parts separates
graphs for lumbricalis and hectus.
A New Typhlops from Hispaniola 17
TasLE 1. Meristic and color variation among the Hispaniolan species of
Typhlops.
Pre- Post-
ocular ocular Scale Middorsal
number number rows scales Coloration
lumbricalis 1 1-2 20-18 231-294 bicolor
20-20 ss bicolor to
hectus 1 2 90-18 284-328 Pee a
pusillus 2 2; 20-18 952-315 bicolor to
unicolor
syntherus 1 2 WON) 304-353 bicolor
sulcatus 1 1 20-20 400-452 bicolor
capitulatus 1 2; 20-20 358-418 unicolor
scale rows situated relatively far posteriorly. Despite an average far
posterior reduction level, these specimens are somewhat heterogeneous in
this respect (Fig. 3). The four from the eastern south island are most
like paratypic hectus in reducing far posteriorly (one does not reduce).
Of those from the north island, the specimens from the northern Valle
de San Juan reduce to 18 rows farther anteriorly than those in any other
sample and are therefore least like paratypic hectus in this respect. The
sample from the vicinity of Lascahobas broadly overlaps the other samples
of both paratypic and non-paratypic hectus in reduction level, and the
three specimens from the Hondo Valle and Vallejuelo region are inter-
mediate. One of the two specimens from E] Iguito in the eastern Valle
de Neiba reduces far anteriorly and the other far posteriorly. Some of
the samples of non-paratypic hectus are more heavily pigmented than
paratypic hectus, and all have higher average mid-dorsal counts (Fig. 3).
Two specimens of non-paratypic T. hectus have divided preoculars uni-
laterally (AMNH 41266 from Duvergé and ASFS V398 from 7 km NW
Vallejuelo); AMNH 41265 from Duvergé has an incompletely divided
preocular on one side; ASFS V396 and V398 (Vallejuelo) and V513
(Rio Arriba) have indications of a groove on one preocular. Divided
preoculars characterize Typhlops pusillus Barbour, a widespread
Hispaniolan species. The significance of divided preoculars in specimens
of T. hectus will be discussed in a comprehensive study of Antillean
Typhlops.
Because of the isolation, heterogeneity in reduction level, and divergence
of these eastern and northern samples from the balance of the specimens
of T. hectus, I have not designated them as paratypes. It is hoped that
the variation in these populations will be elucidated by more compre-
hensive sampling.
Comparisons: Typhlops hectus is readily distinguishable from four of
the other Hispaniolan species on rather gross morphological features
18 Proceedings of the Biological Society of Washington
(Table 1). From lumbricalis, however, hectus cannot be absolutely dis-
tinguished on the basis of meristic features. There is a small amount
of overlap between the two species in mid-dorsal counts (Fig. 3). In
reduction level lumbricalis and paratypic hectus are almost completely
separable; the non-paratypic samples are intermediate. The differences
in head scale shape are subtle; some specimens are difficult to distinguish
on head scale shape alone. In hectus the rostral is narrow in dorsal view,
almost parallel sided but with a slight terminal expansion; in lumbricalis
it may be narrow but is usually broader than in hectus, and the widest
point is near the middle (Fig. 1). The preocular in hectus is more
sharply pointed (the anterior edges form a smaller angle) than that of
lumbricalis. A correlate of the low preocular angle in hectus is that the
lateral edges of the postnasals are more nearly parallel with one another
and with the lateral edges of the rostral, whereas they are divergent in
lumbricalis (Fig. 1). In addition, the Cul de Sac lumbricalis typically
have four parietals (32 of 37); only 25 of 61 paratypic hectus have
four parietals. Ten of 11 of the eastern Hispaniolan lumbricalis have the
first parietal (only two specimens have paired parietals and these only
unilaterally ) greatly elongated and fused with the upper postocular, a
condition not found either in hectus or other lumbricalis. T. lumbricalis
also is a larger, stouter species than T. hectus.
Although lumbricalis and hectus have partially overlapping geographic
ranges, their microsympatry is not established. At no time have the two
been taken together, despite the collection of fair numbers of lumbricalis
(and three other species) from the Cul de Sac. Microsympatry between
lumbricalis and hectus will probably be most easily demonstrated in the
area between the Cul de Sac and Pétionville.
Comparative material: Typhlops lumbricalis—HAITI, D&pr. DE
L OvEst: MCZ 62637, 81150, CM 38886, ASFS V8185-86, Manneville;
USNM 117270-72, 117275, ASFS V24346, Trou Caiman; BM 1948.
1.6.63-64 (2), Pont Beudet; USNM 75893, 123792, MCZ 51426, 62631-
33, Port au Prince; MCZ 65812, near Port au Prince; ASFS V8394—97,
Chateau Blond, 6.4 km NE Pétionville, 160 m; ASFS V22440-44, LDO
7-6464-69, 4.8 km N Pétionville; ASFS V24346, 1.6 km NW Pétionville;
MCZ 68571, Colombier near (22 km E) Saltrou. REPUBLICA DO-
MINICANA, Prov. PEDERNALEs: ASFS V2604, 11 km SW Los Arroyos,
443 m; ASFS V2708, 21 km N Pedernales, 243 m. Prov. Azua: ASFS
X8047, 2.9 km W, thence 16.4 km N, Azua. Prov. SAN PEDRO DE
Macoris: AMNH 13630, San Pedro de Macoris. Prov. La Veca: ASFS
V35897-900, V35904—05, 1 km W Jayaco, 274 m; ASFS V14143, 4 km
S La Vega. Prov. SaAaMANA: USNM 55298, Sanchez. Prov. PuErro
PLata: USNM 10276, Puerto Plata.
LITERATURE CITED
Tuomas, RicHarD. 1965. A new species of Typhlops from the Bara-
hona Peninsula of Hispaniola. Copeia 1965(4):4386—-439.
mez) G 7 3
Vol. 87, No. 3, pp. 19-26 25 April 1974
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
ALLACTAEA LITHOSTROTA, A NEW GENUS AND
SPECIES OF CRAB (DECAPODA: XANTHIDAE)
FROM NORTH CAROLINA, U.S.A.
By Austin B. WILLIAMS
National Marine Fisheries Service Systematics
Laboratory, Washington, D.C. 20560
In 1963 an unusual female crab of the family Xanthidae was
brought to my attention in a collection of invertebrates made
during a survey of benthic fauna off North Carolina, U.S.A.,
by M. J. Cerame-Vivas, then at Duke University Marine
Laboratory. Description was delayed until males could be
found and studied to allow more complete analysis of the
species.
Genus Allactaea, new genus
Areoles of carapace prominent, indicated by large and small lobules
often arranged in triads, tending to proliferate with increasing age.
Mouth frame roughly quadrate, widest anteriorly. Third maxillipeds
separated; merus wider than long, with anteroexternal corner produced,
anterior margin sinuous, internal corner notched for insertion of palp.
Lacinia of first maxillipeds moderate in size. Fused basal antennal seg-
ment extending into inner orbital hiatus. First gonopods of male
terminating in a more or less flat and moderately developed lobe pre-
ceded by a tuft of long hairs in a nearly single file.
Type-species: Allactaea lithostrota.
Etymology: From the Greek “allos” other, and “actaea.” The gender
is feminine.
Allactaea lithostrota, new species
Figures 1-3
Holotypic male: Carapace xanthoid (Fig. 1B), wider than long, arched
anteroposteriorly, regions prominently indicated by smooth raised lobules
arranged in a radiating pattern eminating laterally and anteriorly from
urogastric region. Smooth tracts between lobules almost completely
3—Proc. Bion. Soc. WAsuH., Vou. 87, 1974 (19)
20 Proceedings of the Biological Society of Washington
Het
Fic. 1. A, right and left chelae of holotype ¢, frontal view; B, holo-
type ¢, dorsal view; C, allotype ¢@, dorsal view; mm scales (upper,
A-B; lower, C).
obscured by feltlike covering of clavate hairs anterior to urogastric
region; hairy covering much thinner posteriorly.
Front fairly prominent and somewhat produced at each side of
median, narrowly U-shaped notch; a dorsal, submarginal row of about 8
A New Xanthid Crab from North Carolina 21
coalesced lobules on each side extending from antennal notch to near
midline giving front a transversely doubled appearance (Fig. 2).
Lobules, exclusive of margin, arranged as follows (Fig. 1B): 2 F (see
Rathbun, 1930, Fig. 3), 2 well separated laterally; 1 M, 3 completely
coalesced posteriorly, less so anteriorly; 2 M, evidence of 8 or 9 clumped
in two groups, a longitudinally oriented and fused group of 3 near
midline having a deep lateral cleft, and a more loosely clumped group of
3 and rudimentary fourth along lateral margin of region with 2 associated
lobules in intermediate space; 3 M, a row of 6 in midline, 4 in transverse
basal row, each series more or less fused; 4 M, 3 separated in crescentic
transverse row; 1 L, solitary and small near anterolateral border; 2 L, 2
closely approximated and among largest individual lobules; 3 L, 2
approximately size of 1 L and not coalesced on right side, but coalesced
into single large lobule on left side; 4 L, solitary and small near fourth
and fifth anterolateral teeth; 5 L, a complex of 2 groups of 3 nearly
coalesced, among largest on carapace; finally, a complex composed of
1 P, 2 P, 1 R and 3 R arranged in irregular, transverse, lunate rows with
fairly symmetrical fusion of lobules within rows. Orbital margin thickened,
raised and smooth dorsally with fissures closed but evident as depressons;
outer orbital hiatus a broadly open notch; obtuse suborbital tooth followed
laterally by irregularly increasing suborbital rim. Eyestalks with a tuft
of clavate hairs in corneal emargination preceded by 2 spiniform
tubercles and a number of much smaller tubercles at base of cornea. Four
prominent anterolateral teeth exclusive of outer orbital widely separated
and reduced to anteriorly directed mammillary lobules; on margin a bit
below these a row of smaller slender intercalary teeth; hepatic and sub-
branchial region lightly tuberculate. Posterolateral border bearing a
row of lobules.
Epistome with 2 tubercles on lower border of each antennular socket,
mesial one largest. Mouth frame roughly quadrate (Fig. 2), widest
anteriorly, anterior margin smoothly and sharply cristate and shaped
into a low angular arch on each side. Third maxillipeds separated, di-
vergent anteriorly; ischium with a shallow, smooth, longitudinal, ventral
depression; merus broader than long with slightly sinuous anterior
margin, notched for reception of strong palp and produced anterolaterally,
tubercles clustered on ventral surface along mesial border and emargin-
ation for palp but scattered elsewhere centrally and near anterior margin.
Lacinia of first maxillipeds moderate in size covering roughly 40 percent
of endostomial surface. Inner orbital hiatus filled by a club-shaped
extension of fused basal antennal segment and first movable antennal
segment.
Sternal plastron relatively smooth, bearing scattered clavate hairs. Ab-
domen with telson wider than long, tip rounded, segments 3-5 fused.
Chelipeds (Fig. 1A) dissimilar, right larger than left. Palm and carpus
of each ornamented on outer surfaces with well separated distally di-
rected lobules varying from sharpest along dorsal crest of carpus and palm
to flared at top with narrower bases on external surface of carpus and
22 Proceedings of the Biological Society of Washington
Fic. 2. Portion of frontal, suborbital and mouth field regions of
allotype @ showing edge of front, antennules, left antenna and eye,
epistome, left third maxilliped, lacinia of right first maxilliped in situ with
overlying mouthparts turned aside, and outline of mandibles in situ.
dorsal portion of palm, becoming smaller, sharper and tending to align-
ment in longitudinal rows on outer surface of palm. Carpus with a
spiniform lobule at inner angle and another below it; anterior border
with a few small teeth hidden in hairs. Dorsal and outer surfaces of
chela, carpus and merus with spaces between lobules and tubercles
covered with dense coating of clavate hairs, inner surfaces smooth.
Fingers dark colored, a crest of tubercles at base of each dactyl; teeth
distinct but forming a continuous row on occlusal surface of each finger,
proximal teeth of major chela somewhat molariform. Merus of each
cheliped with a dorsal crest of sharp forward-curved spines flanked
laterally by a few tubercles.
Second and fifth pereopods with external surfaces covered by a dense
ragged coat of clavate hairs (third and fourth pairs of pereopods
missing). Dorsal crest of merus, carpus and propodus of each leg armed
with erect, well separated, distally trending spines; a secondary lateral
row of spiniform tubercles nearly hidden in hairs on carpus and propodus.
A New Xanthid Crab from North Carolina U8}
SS SS SSS
S) &
Fic. 3. First right gonopod of paratype ¢ (USNM 143774), terminal
portion; A, mesial view, B, lateral view; scale = 0.1 mm.
Dactyl of second legs densely hairy on all surfaces, of fifth legs nearly
naked on inner surface, both with darkened corneous tip.
First gonopod of male (Fig. 3) rather stout, approximated along middle
part but tips diverging ventrolaterally, terminating in a membranous
rounded lobe. An irregular row of articulated retrogressive spines along
sternal edge near tip grading from scattered and smallest proximally to
24 Proceedings of the Biological Society of Washington
denser, longer and more slender distally, and merging finally into an
irregular subterminal file of elongate hairs; a broader tract of similar
scattered spines on mesioventral surface and a lateral cluster at base of
terminal lobe. In addition, a row of slender short spinules on mesial
surface following course of sperm duct.
Measurements in mm: Holotypic male: greatest width of carapace at
level of fourth anterolateral tooth, 16.2, length in midline, 11.6; telson
width, 1.8, length, 1.6.
Allotypic female: Lobular pattern (Fig. 1C) basically same as in
holotype but specimen larger and more omate suggesting greater age.
Coalescence of lobules varying in minor details from that in holotype
with introduction of additional small solitary lobules at edge of and
between areoles, especially behind orbits, on branchial regions and antero-
lateral borders. Lunate arrangement or rows of lobules behind urogastric
region in 1 P, 2 P, 1 R, 2 R and 3 R not so evident as in holotype.
Epistome with tendency to formation of a row of tubercles on lower
border (Fig. 2) of antennular socket, 3 on right side, 2 on left. Lobulation
at corners of buccal frame and subhepatic region much bolder and merus
of third maxillipeds with ventral tubercles stronger than in holotype.
Sternal plastron lightly punctate and hairy on exposed parts becoming
ornamented with obsolescent granules along anterior and anterolateral
borders at base of third maxillipeds and chelipeds.
Abdomen ovate, densely edged with long hairs; segments lightly
covered externally with clavate hairs except on worn raised portions.
Telson broadly rounded, broader than long.
Chelipeds with disparity in size and shape more pronounced than in
holotype. Lobules not pointed but many with tips rounded, flattened
and flared forming knobs. Anterior edge of carpus with a row of spines,
a strong spine at inner angle followed by a second almost equal below
it. Proximal teeth of major chela more molariform than in holotype.
Remaining pereopods essentially as in holotype, each with crest of
distally trending spiniform lobules, many of them club shaped, along
upper border of merus, carpus and propodus; crest most pronounced in
coalesced ornate lobules on superodistal angle of merus. Each leg with
an external flanking row of lobules on carpus and propodus, and a
scattering of lobules on merus of fifth leg. Lower edges of each merus
with ornamentation grading from proximal granules to attenuated distal
lobules. Legs clothed in dense growth of clavate hairs, longest and most
ragged along crests.
Measurements in mm: Allotypic female: greatest width of carapace
at level of fourth anterolateral tooth, 28.1, length in midline, 18.1; telson
width, 4.9, length, 4.0. Ovigerous female paratype: greatest width of
carapace at fourth and fifth anterolateral teeth, 17.2, length in midline,
le
Type locality: Approximately SE Cape Lookout, N. C., 33°43’ N,
76°40’ W, 90 m-33°42.7’ N, 76°40.2’ W, 110 m, Eastward Sta. 1087.
A New Xanthid Crab from North Carolina 25
Material studied: Specimens studied are confined to the type series
deposited in collections of the National Museum of Natural History
(USNM), Washington, D. C., and University of North Carolina Institute
of Marine Sciences (UNC-IMS), Morehead City, N. C.
USNM 143770 6 holotype; USNM 143772 ¢ juv. paratype, Eastward
Sta. 1087, 27 Apr. 1965, L. R. McCloskey, col. USNM 143771 off Cape
Lookout, N. C., 91 m, Oct. 1963, 2 allotype, M. Cerame-Vivas, col.
USNM 143773 32°39’ N, 78°32’ W, 91-95 m, 28 July 1960, 2 (ov)
paratype, R/V Silver Bay Sta. 2266, clam dredge. USNM 143774
33°25.7’ N, 77°01’ W, 60-70 m, 17 Mar. 1969, ¢ paratype (frags.),
Eastward Sta. 11552, dredge, N. Chamberlain, col. UNC-IMS 2299,
34°11’ N, 76°10’ W, 60 m, 4 May 1969, @ paratype, Eastward Sta.
11943, F. J. Vernberg, col. UNC-IMS 2366, about 34°08’ N, 76°10’ W,
50 m, 27-30 Apr. 1966, 2 juv. (dry) paratype, Eastward, C. E. Jenner
and class, col.
Name: The name is from the Greek “lithostrotos” meaning inlaid with
stones, a term descriptive of the symmetrical lobate ornamentation.
Remarks: Allactaea lithostrota is unusual among numerous species of
the actaeid group in that lobules of the carapace are large, smooth, and
either clustered or solitary in each area. Tending to be rather sharply
pointed in juveniles, the lobules become progressively blunter as they
increase in size, many of them finally becoming flattened knoblike ex-
pansions on somewhat constricted bases. In addition, new small lobules
proliferate with increasing size. Clusters on frontal and medial areas
tend to be tripartite with an opening in the cluster directed forward and
outward. All lobulations of the carapace tend to be peripherally directed
away from the urogastric area.
Surface ornamentation indicates close relationship to none of the
American actaeas, but characters of the mouth field and orbito-antennal
region are similar to Actaea de Haan 1833 (emend.) and those of the
first male gonopod resemble Paractaea Guinot, 1969 (Guinot, 1967;
1969; 1971). On the basis of these comparisons, the mouth field of
Allactaea lithostrota has a remote resemblance to that of Indo-Pacific
Actaea calculosa (H. Milne Edwards, 1834) and A. glandifera Rathbun,
1914 (Guinot, 1969), and the pleopods to those of western Atlantic
Paractaea rufopunctata nodsa (Stimpson, 1860) (Guinot, 1969).
Allactaea lithostrota may be confined to the Lithothamnion reeflike
structure described by Menzies, Pilkey, Blackwelder, Dexter, Huling and
McCloskey (1966), and Menzies, George and Rowe (1973) in warm
water near the edge of the continental shelf off North Carolina.
I am indebted to M. J. Cerame-Vivas, N. A. Chamberlain, C. E. Jenner,
L. R. McCloskey and F. J. Vernberg for gifts of specimens, H. B. Roberts
for counsel, F. A. Chace, Jr., B. B. Collette and I. Pérez Farfante for
criticism of the manuscript, and M. Diéguez for drawings and assembly
of figures. The type, allotype and two paratypes have been transferred
to USNM from UNC-IMS lots 2365, 1899 and 2374.
26 . Proceedings of the Biological Society of Washington
LITERATURE CITED
Gurnor, D. 1967. Recherches préliminaires sur les groupements naturels
chez les crustacés décapodes brachyoures. III. A propos des
affinités des genres Dairoides Stebbing et Daira de Haan. Bull.
Mus. Natl. Hist. Nat. Paris Ser. 2, 39(3):540-563.
—. 1969. Sur divers Xanthidae notament sur Actaea de Haan et
Paractaea gen. noy. (Crustacea Decapoda Brachyura). Cah.
Pac. No. 3:223—267.
1971. Recherches préliminaires sur les groupements naturels
chez les crustacés décapodes brachyoures. VIII. Synthése et
bibliographie. Bull. Mus. Natl. Hist. Nat. Paris Ser. 2, 42(5):
(for 1970) : 1063-1090.
Menzies, R. J., R. Y. GEorcE AND G. T. Rowe. 1973. Abyssal environ-
ment and ecology of the world oceans. John Wiley and Sons,
New York. xxiii + 488 pp.
, O. H. Pmxey, B. W. BLACKWELDER, D. DEXTER, P. HULING
AND L. McCtoskey. 1966. A submerged reef off North Caro-
lina. Int. Rev. Gesamten Hydrobiol. 51(3):393—431.
Ratusun, M. J. 1930. The cancroid crabs of America of the families
Euryalidae, Portunidae, Atelecyclidae, Cancridae and Xanthidae.
U.S. Nat. Mus. Bull. 152: 1-609.
MX G&S (ENC er
Vol. 87, No. 4, pp. 27-30 25 April 1974
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
MICROGNATHUS ERUGATUS, A NEW MARINE
PIPEFISH FROM BRAZIL (PISCES: SYNGNATHIDAE)
By E. S. Heratp' anp C. E. Dawson
Steinhart Aquarium, California Academy of Sciences,
San Francisco, California 94118, and
Gulf Coast Research Laboratory Museum
Ocean Springs, Mississippi 39564
In the course of recent fieldwork by the junior author in
Central and South America, we have been fortunate to obtain
a single specimen of an unusual marine pipefish. Characters of
this new species are so distinctive that we do not believe it
necessary to delay its description until more specimens are
available.
The holotype has been deposited in the Museum de Zoologia
da Universidade de Sao Paulo (MZUSP), Brazil. Measure-
ments are in millimeters (mm).
Appreciation is expressed to Dr. Naercio A. Menezes
(MZUSP) for making this specimen available for study. The
entire J. L. Almeida family of Salvador, Brazil is thanked for
their hospitality and active participation in the field. This
work was supported in part by National Science Foundation
Grant GB 31053X.
Micrognathus erugatus, new species
Figure 1
Holotype: MZUSP 9408 (64 mm SL, female); Brazil, Bahia, rock
tidepool about 1 km S of Arembepe, approx. 12°45’ S, 38°10’ W, 0-1.3
m, chemical ichthyocide; 26 August 1972, C. E. Dawson and party.
Diagnosis: Dorsal-fin rays 19, located on 0.5 trunk rings and 4.5 tail
rings; trunk rings 20; tail rings 36; pectoral-fin rays 13 (2); anal-fin
rays 2; caudal-fin rays 10; head length 5.9; snout length 1.9; length of
1 Deceased.
4—Proc. Biot. Soc. WasH., Vou. 87, 1974 (27)
28 Proceedings of the Biological Society of Washington
Fic. 1. Head and anterior body of Micrognathus erugatus MZUSP
9408; holotype; 64 mm SL.
dorsal-fin base 4.9; caudal-fin length 2.0; body ridges smooth, poorly
developed.
Description: Head ridges indistinct, only snout, supraorbital and
opercular ridges visible. Opercle with 20 or more rows of minute pock-
like depressions radiating from the indistinct median ridge. Excurrent
gill openings extend to top of head. Body ridge pattern typical of
Micrognathus, i.e., lateral trunk ridge deflected ventrad at anal ring
thence continuous with inferior tail ridge; superior tail ridge deflected
at dorsal fin to become lateral tail ridge, ending free near middle of last
trunk ring. Body ridges rounded, indistinct; intermedial scutellar plates
on trunk and anterior portion of tail equal about half width of individual
rings. Pectoral cover plate without ridges but with a few pocklike de-
pressions; pectoral-fin length equals about 1.75 trunk rings. Head-in-SL
10.8; snout-in-head 3.1; dorsal-fin base-in-head 1.2.
Color in alcohol mainly dark brown with indications of 5 darker bands
on upper sides and dorsum of trunk and 10 on tail; tip of upper jaw
white; narrow white cross bar below angle of gape; white surrounds
nares and continues as narrow, irregular bar across snout; streaks of
white radiate from eye and one streak extends posteriad as narrow blotch
on opercle; pectoral-fin base and top of head mottled with white; pale
blotches bisect dark trunk and tail rings. Dorsal-fin rays streaked with
brown near bases, fin otherwise pale; caudal fin mainly brown, but
uppermost ray and distal margin white.
Etymology: erugatus, from the Latin e, not, -- rugatus, wrinkled, in
allusion to the almost smooth head and body.
Discussion: The genus Micrognathus now includes the subgenera
Micrognathus, Anarchopterus and Minyichthys (Herald and Randall,
1972). Counts of 13-17 trunk rings occur in all but the two known spe-
cies of the Pacific Minyichthys, which is characterized by 19-21 trunk
rings and well-developed head and body ridges. The 20 trunk rings of
M. erugatus suggest a close relationship with Minyichthys, but its smooth
head and poorly developed ridges are atypical of Minyichthys. In the
latter characters, Micrognathus erugatus is similar to the smooth-bodied
Atlantic American Anarchopterus, but the anal fin is absent in this
subgenus.
A New Pipefish from Brazil 29
The unique holotype of Micrognathus erugatus was collected with eight
specimens of the wide-ranging (Florida—Brazil) M. vittatus (Kaup).
This is the only Atlantic representative of the subgenus Micrognathus,
although seven of its nine Pacific congeners fall in this characteristically
sharp-ridged subgenus. The western Atlantic species Micrognathus
dawsoni (Herald, 1969) was originally placed in the subgenus Micro-
gnathus, but its body ridges are not sharp edged and it is the only long-
snouted species in a genus otherwise noted for short-snouted forms.
The Atlantic Micrognathus group presents somewhat of an enigma in
that there are now five species, representing four intrageneric trends, and
two species (M. dawsoni and M. erugatus) do not fall within described
subgeneric limits. These two species are known only from unique
female holotypes and additional study material will hopefully help
clarify relationships within the genus.
LITERATURE CITED
Heratp, E. S. 1969. A new pipefish from the Virgin Islands, Micro-
gnathus dawsoni. Occ. Pap. Calif. Acad. Sci. 73:1-3.
, AND J. E. RANDALL. 1972. Five new Indo-Pacific pipefishes.
Proc. Calif. Acad. Sci., 4th Ser. 39(11):121—140.
30 Proceedings of the Biological Society of Washington
Mn fa. CJD F 7
Vol. 87, No. 5, pp. 31-40 25 April 1974
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
POTAMORRHAPHIS PETERSI, A NEW SPECIES OF
FRESHWATER NEEDLEFISH (BELONIDAE)
FROM THE UPPER ORINOCO AND
RIO NEGRO
By Bruce B. COLLETTE
National Marine Fisheries Service Systematics Laboratory
National Museum of Natural History,
Washington, D.C. 20560
While reviewing the South American freshwater needlefish
genus Potamorrhaphis, an undescribed species was found from
the upper tributaries of the Orinoco in Venezuela and Co-
lombia and the upper Rio Negro in Brazil. Because completion
of the generic review has been delayed, I have decided to
describe the new species separately. It is herein named in
honor of the late Dr. James A. Peters, who was best known as
an authority on Neotropical amphibians and reptiles. He
directly aided my studies of the South American freshwater
needlefishes Potamorrhaphis and Pseudotylosurus, and I take
great pleasure in naming this species after him.
Study materials are in the collections of the Academy of
Natural Sciences, Philadelphia (ANSP); the American Mu-
seum of Natural History (AMNH); California Academy of
Sciences, San Francisco (CAS and SU); Muséum National
d'Histoire Naturelle, Paris (MNHN); INDERENA, Bogota;
Museo de Biologia, Universidad Central de Venezuela, Caracas
(MBUCV); and the National Museum of Natural History
(USNM). This paper is one in a series on the systematics
of the Synentognathi; see Collette (1966) on Belonion and
Collette (in press) on Pseudotylosurus for studies on the
other two genera of South American freshwater needlefishes.
Discovery of a second species of Potamorrhaphis and sepa-
5—Proc. Brot. Soc. WAsH., Vou. 87, 1974 (31)
32 Proceedings of the Biological Society of Washington
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34 Proceedings of the Biological Society of Washington
TaBLE 2. Measurements (as percent of body length) of Potamorrhaphis
guianensis from the lower Orinoco and P. petersi.
P. guianensis P. petersi
Character Range x N Range x N
Body length (mm) 87.8 119 1OL9 23. (92770 162) MeO
Head length 55.2 59.2 56.91 11. 47.8 (Sb. O lesa
Snout length 37.0. 42.10 "40152 1 S39) Valor oma
Postorbit length NOS pA Be 28} 94 11.1 AGO 9
Preopercle length Sto GO Olin 3 5.2) | 15.9 aaa)
Orbit length AVON Olly Age) PAB 3.9 | AiG 424
Interorbit width AW 6) EN IIA BAS} AD AQY Ai529
Head depth Bee, Ga) OM WB A9 60 554 9
Head width Sele > Oo aS AT (6.30 or A0men
Pectoral-fin length 10337 13S .27 11 s6r as 9.7 118 1061 9
Pelvic-fin length dh TR BAD), 8} 6.2)" 73°) Gione9
Pectoral fin to
pelvin fin 47.4 52.0 4944 23 435 468 44.94 9
Pelvic fin to
caudal-fin base 46.0 51.4 48.97 23 516 542 53.18 9
ration (Collette, in press) of Pseudotylosurus microps
(Ginther) from Ps. angusticeps (Gtinther) raises the total
number of species of freshwater synentognaths known from
South America east of the Andes to seven. As previously noted
(Collette, 1966), the other three include two neotenic needle-
fishes Belonion dibranchodon Collette (described from the
same Rio Atabapo locality as three paratypes of P. petersi)
and B. apodion Collette plus the halfbeak Hyporhamphus
brederi (Fernandez-Yépez).
Potamorrhaphis petersi, new species
Figures 1-3
Diagnosis: A species of the genus Potamorrhaphis that differs from
the other nominal species of the genus, P. guianensis (Schomburgk) and
P. eigenmanni Miranda Ribeiro, primarily in having the posterior portion
of the body greatly elongated. This is best reflected in the higher counts
(Table 1): greater total number of vertebrae (77-85 vs. 64-77);
caudal vertebrae (38-44 vs. 28-37); dorsal-fin rays (37-43 vs. 27-36);
and anal-fin rays (32-39 vs. 24-31). Predorsal scales are more nu-
merous than in the Orinoco population of P. guianensis (110-141, x 128
vs. 97-112, x 102).
Morphometrically (Table 2), this difference is shown by the distance
30
‘[izeig “uozeury
TOMOT “TQNI— ONY “WU BOT “LEG6LT WNSN ‘sisuaupins -g ‘0 “eIquIC[OD “ee OLY “WU ETT “ESSOTT dSNV ‘Stsuaunins gq -q
‘ejonzoueA ‘Oo0UTIO Joddn ‘eping ‘ij Jeou “YyySue] Apog wu FET ‘6196 HNIWY ‘edAyered ‘isuajad srydoysowpjog ‘e ‘[ ‘ol
eo)
alee aA
Bal le pte a .—
Sn SAO
A New Needlefish from South America
WY NN
CAN WMS
WW KK SX S MSV
36 Proceedings of the Biological Society of Washington
Fic. 2. Diagrammatic representation of pigment patterns of five
specimens of Potamorrhaphis petersi. From top to bottom: USNM
210546, holotype, 162 mm BL, junction Rio Guaviare and R. Inirida;
USNM 210547, paratype, 137 mm, same data; AMNH 9619, paratype,
133 mm, upper Orinoco; MBUCV-V-6132, paratype, 124 mm, Rio
Casiquiare; and INDERENA G.A-P-0066, 111 mm BL, Rio Tomo.
from the pelvic-fin origin to the caudal-fin base (P:-C) being much
greater than the distance from the pectoral-fin origin to the pelvic-fin
origin (P:—P.) in P. petersi (PC 51.6-54.2% vs. Pi-P2 43.5-46.8%
body length). These distances are about equal in P. guianensis from the
lower Orinoco (P:—-C 46.0-51.4% vs. P:—P2 47.4--52.0% body length). In-
creasing the P:-C distance in P. petersi also increases the body length
37
A New Needlefish from South America
‘(S}Op) sisuaupina
‘d
jo suonejndod ureyj10u pue (sieys) ssvajzad srydpy.lownjog JO UOTNGIISIq ‘¢ ‘oI
sisusunin6 @
Issojed 4
siydpyssowpjog
aN
ce
S3OWNOS Va
Ay
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4 i
SS SA
XQ Rivinobvs Vf AEE SS
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. VOIUINI SS
auyiAvN
2p) .
38 Proceedings of the Biological Society of Washington
thereby decreasing other body proportions. The type-series of P. petersi
is completely separated from the Orinoco population of P. guianensis by
having a smaller Pi-P. distance and is nearly completely separated in
having a narrower interorbital distance and a shorter head (Table 2).
Other proportions are also smaller but overlap with the Orinoco popu-
lation of P. guianensis.
Potamorrhaphis petersi is also distinct in its pigment pattern. The
deeper lying brown pigment that forms bars (Fig. lc) or a continuous
lateral band (Fig. 1b) in P. guianensis is absent and replaced by
scattered large black melanophores that appear to be more superficial.
These melanophores vary in number, size and position from specimen
to specimen (Fig. 2) and from side to side in a single specimen.
Types: Holotype—USNM 210546, 162 mm body length, Colombia,
Laguna Coco northeast of Puerto Inirida, pool near junction of Rio
Guaviare and Rio Inirida, 17 January 1972, P. Cala. Paratypes: USNM
210547 (1: 137), same data as holotype. AMNH 9619 (1: 133), Vene-
zuela, Cano Pescado, about 8 km north of Esmeralda near Mt. Duida, 9
March 1929, G. H. Tate. MNHN 87-655-6 (2: 97.5-123), and USNM
210861 (1: 108), Venezuela, Amazonas, Rio Atabapo at San Fernando
de Atabapo, October 1886; J. Chaffanjon. MBUCV-V-6132 (1: 124),
Venezuela, Amazonas, Cafo Beripamoni, tributary of Rio Casiquiare, 29
January 1969, F. Mago L., J. Moscé, A. Machado. CAS 27587 (1: 92.7),
Brazil, Rio Negro, Sao Gabriel rapids, above Camanaos, 1 February
1925, C. Ternetz.
Other specimen examined: INDERENA G.A-P-0066 (1: 111), Co-
lombia, Rio Tomo near entrance into Orinoco.
Comparative material: Only the data on Orinoco Potamorrhaphis
guianensis is presented here; extensive material from the other parts of
the range will be reported on in a subsequent generic review. Seven
series containing a total of 24 specimens (62.7-113 mm body length)
were taken in the lower Orinoco in Cafto Quiribana at Caicara by C.
Ternetz in April and May, 1925: SU 52684, SU 52683, CAS 28322, SU
52686, CAS 28323, SU 58818, USNM 209303. One specimen was
recently collected in a western tributary of the Orinoco: ANSP 116533;
119 mm; Rio Meta, Cafo Emma at Finca El Viente south of Matuzal;
4°08’ N, 72°39’ W, 18 March 1973, J. E. Bohlke, W. Saul, and W.
Smith-Vaniz.
Discussion: Discovery of P. petersi adds another species to the list
of fishes common to the upper Orinoco, upper Rio Negro, and the
connecting Rio Casiquiare (Mago Leccia, 1971). Potamorrhaphis is
probably derived from an inshore marine needlefish similar to Strongylura
and has secondarily developed a larger number of caudal vertebrae and
dorsal- and anal-fin rays (Collette, 1966: 21). In its even higher counts,
P. petersi seems to be a specialized derivative of the wide-spread P.
guianensis. Additional collecting is needed in the upper Orinoco, Rio
Tomo to the Rio Meta and in the upper Rio Negro to ascertain the
A New Needlefish from South America 39
distributional limits of P. petersi and to determine if it is sympatric with
P. guianensis in the Orinoco or Amazon.
The INDERENA specimen from the Rio Tomo has the lowest counts
of vertebrae and dorsal- and anal-fin rays of the small series of P. petersi.
It also has traces of the usual P. guianensis pigment pattern underlying
the large scattered melanophores characteristic of P. petersi. This speci-
men is, therefore, not designated a paratype because of these hints of
possible intergradation with downstream populations of P. guianensis.
However, the recently collected ANSP specimen of P. guianensis from the
Rio Meta agrees well with the 24 specimens from Caicara on the lower
Orinoco.
Additional information about four collections of P. petersi is available
from the following references: Chaffanjon (1889), for the Rio Atabapo
specimens; La Monte (1929), for the AMNH specimen from near Mt.
Duida; Bohlke (1953), for Ternetz material from the upper Orinoco and
Rio Negro; and Mago Leccia (1971), for the Casiquiare specimen.
Acknowledgments: Museum material and information on_ localities
was kindly provided by: Drs. Donn E. Rosen and Gareth J. Nelson and
Miss Avis Rumney (AMNH); Dr. James E. Bohlke and Messrs. William
Saul and William Smith-Vaniz (ANSP); Drs. William N. Eschmeyer,
Warren C. Freihofer, and Tomio Iwamoto and Miss Pearl Sonoda (CAS);
and Sr. Francisco Mago Leccia (MBUCV). Dr. Neal R. Foster (ANSP)
put me in contact with Dr. Plutarco Cala (Universidad Nacional de
Colombia, Bogota) who lent me the INDERENA specimen and graciously
donated two important specimens of P. petersi to the USNM collections.
Mrs. Keiko Hiratsuka Moore drew the figures except for Fig. lc which
was drawn by Mrs. Mildred Carrington; Mr. George Clipper made most
of the radiographs and counted vertebrae; and Drs. Daniel M. Cohen,
Victor G. Springer, and Stanley H. Weitzman commented on the
manuscript.
LITERATURE CITED
BOHLKE, JAMES E. 1953. Studies on fishes of the family Characidae.
No. 5. A review of the genus Microschemobrycon with descrip-
tions of two new species. Ann. Mag. Nat. Hist., ser. 12, 6:
841-849.
CHAFFANJON, J. 1889. L’Orénoque et la Caura. Relation de voyages
exécutés en 1886 et 1887. Librairie Hachette, Paris, 351 pp.
CoLLETTE, BrucE B. 1966. Belonion, a new genus of fresh-water
needlefishes from South America. Amer. Mus. Nat. Hist.
Novitates No. 2274, 22 pp.
in press. South American needlefishes (Belonidae) of the
genus Pseudotylosurus. Zool. Meded., Leiden.
La Monte, FraNcEsca. 1929. Two new fishes from Mt. Duida, Vene-
zuela. Amer. Mus. Nat. Hist. Novitates No. 373, 4 pp.
Maco Leccia, Francisco. 1971. La ictiofauna del Casiquiare. Rev.
Defensa Naturaleza (Caracas) 1(4):5—10.
40 Proceedings of the Biological Society of Washington
ae
me Oe
Vol. 87, No. 6, pp. 41-48 25 ‘April 1974
PROCEEDINGS
OP Une
BIOLOGICAL SOCIETY OF WASHINGTON
A NEW HEART-URCHIN, ARAEOLAMPAS ATLANTICA,
NEW GENUS, NEW SPECIES, FROM THE NORTH
ATLANTIC AND NOTES ON THE CLOSELY
RELATED HOMOLAMPAS
By D. Kerry Serary!
New York Ocean Science Laboratory
Drawer EE, Montauk, New York 11954
INTRODUCTION
In the summer of 1971 two lots of an undescribed species of
Homolampas were discovered in the collections of the National
Museum of Natural History, Smithsonian Institution (USNM).
Unfortunately, both specimens were badly broken. An attempt
was made to reconstruct the specimens, but in neither case
was a complete specimen obtained. A similarly incomplete
lot was found in the Museum of Comparative Zoology at
Harvard University (MCZ). During 1972 three more lots of
fragments were discovered in the unidentified echinoid collec-
tions of the University of Miami, School of Marine and
Atmospheric Sciences (UMML). A description of this new
species has been withheld for three years in the unfulfilled
hope of finding a complete specimen. A complete specimen
may be very difficult to obtain since the species occurs in
deep water and has a very fragile test.
A review of Homolampas revealed two distinct species
groups. One group, including H. fragilis and H. lovenioides,
has a small test (less than 35 mm TL), no peripetalous
fasciole, deeply sunken areoles that form internal pouches,
globiferous pedicellariae, non-crenulate primary tubercles
1 The first part of this work was done in the Department of Zoology, University of
Maine at Orono.
6—Proc. Brot. Soc. WaAsH., Vou. 87, 1974 (41)
42 Proceedings of the Biological Society of Washington
which do not occur on interambulacrum 5 and three (rarely
four) genital pores. The other group, including H. fulva, H.
glauca, H. hastata, H. rostrata and the new species, has a
large test (up to 110 mm TL, except for H. hastata which is
less than 35 mm TL), peripetalous fasciole, slightly sunken
areoles that form slight internal bulges, no globiferous pedi-
cellariae, crenulate primary tubercles extending on to inter-
ambulacrum 5 and four genital pores. On the basis of the
peripetalous fasciole alone, Mortensen (1950: 265) believed
that Homolampas may include two distinct genera, but as
the species were poorly known, he retained the single genus.
The presence of the peripetalous fasciole and the other char-
acters mentioned above are believed to merit generic dis-
tinction. Since the type-species is Homolampas fragilis (A.
Agassiz, 1869) the group including H. fragilis and H.
lovenioides is included in Homolampas, strict sense. The
other group including the five remaining species comprise the
new genus described herein. A key to the species of Homo-
lampas s.s. and Araeolampas n. gen. is given below.
Kry TO THE SPECIES OF Homolampas s.s. AND Araeolampas N. GEN.
1. No peripetalous fasciole; no primary tubercles on interambulacrum
5; deeply sunken areoles forming internal pouches; 3 (rarely 4)
genital pores; globiferous pedicellariae present; primary tubercles
non-crenulate; small forms generally less than 35 mm TL
2s ABTA va Pie eo ip ee lee eee ee Homolampas s.s. 2
1’ Peripetalous fasciole present; primary tubercles on interambulacrum
5; areoles not forming internal pouches; 4 genital pores; no glo-
biferous pedicellariae; primary tubercles crenulate; generally
large forms up to 110 mm TL (except A. hastata, which is less
than: 35mm (1s) 3 ae eee Araeolampas n. gen. 3
2. Posterior prolongation of labrum not reaching beyond second pair
of adjoining ambulacral plates (tropical western Atlantic)
MEN rs Paes ser Ek volte! Homolampas fragilis (A. Agassiz, 1869)
2’ Posterior prolongation of labrum reaching middle of third pair
of adjoining ambulacral plates (Malay Region)
CIMT AIM of Bile Meee RR OA a Homolampas lovenioides Mortensen, 1948
3. Posterior end of test forming a rostrum surrounded by subanal
fasciole; periproct visible from above (Indonesia) —.------__--
Sine esl sal Wedica hd hp OR RC Araeolampas rostrata (de Meijere, 1902)
3’ Posterior end of test not forming a rostrum; periproct not visible
from above .2.2.2 3s ee ee ee 4
A New Heart-Urchin from North Atlantic 43
4, Only a single primary tubercle in each series of interambulacra 1,
4 and 5; small forms less than 35 mm TL (tropical eastern
ACTIC)) pees ee Araeolampas hastata (A. Agassiz, 1898 )
4’ Several primary tubercles in each series of interambulacra 1, 4
aval Be lava iwormane wre wo, JMC) more Ib; 5
5. Frontal notch shallow (about 2 percent TL) (Indian Ocean)
_ erro Araeolampas glauca (Wood-Mason & Alcocks, 1891)
HaErontalsnoteh deep (about 5 percent PI) 220. 6
6. Tridentate pedicellariae with very narrow blade (Figure 1d and
e); periproct small (width plus height about 15 percent TL);
pores of ambulacra double adapically (central and eastern
IRACHIC) preter ti Loe Araeolampas fulva (A. Agassiz, 1879)
6’ Tridentate pedicellariae with wide blade (Figure lc); periproct
large (width plus height 19 to 24 percent TL); pores of ambulacra
single adapically (North Atlantic) _.. Araeolampas atlantica n. sp.
Homolampas, s.s.
Diagnosis: Small forms up to 35 mm TL; test fragile with a distinct
frontal notch; ambulacral plates flush with interambulacral plates, not
sunken to form petals; ambulacral pores double, placed obliquely near
inner edge of plates; primary tubercles perforate, non-crenulate and
present on interambulacra 1—4; primary tubercles with deeply sunken
areoles that form internal pouches, as in Lovenia; primary spines long,
curved, uniformly dentate; subanal fasciole present; apical system with
3 (rarely 4) genital pores; peristome anterior; labrum with long
posterior prolongation reaching to second or third pair of adjoining
ambulacral plates; anterior portion of sternum naked of primary spines;
globiferous, tridentate, ophicephalous and_ triphyllous pedicellariae
present; sphaeridia not placed in pits.
Type-species: Lissonotus fragilis A. Agassiz, 1869.
Araeolampas, new genus
Diagnosis: Small to large forms up to 110 mm TL; test fragile with
distinct frontal notch; ambulacral plates flush with interambulacral
plates, not sunken to form petals; ambulacral pores single or double,
placed obliquely near inner edge of plates; primary tubercles perforate,
crenulate, present on interambulacra 1-5; primary tubercles with slightly
sunken areoles forming slight internal bulges; primary spines long, curved,
unilaterally dentate; peripetalous and subanal fasciole present; apical
system with 4 well-developed genital pores; peristome anterior; labrum
with long posterior prolongation; anterior portion of sternum naked of
primary spines; tridentate, rostrate and triphyllous pedicellariae present;
sphaeridia not placed in pits.
Type-species: Homolampas fulva A. Agassiz, 1879.
Etymology: The generic name is derived from the Greek araeo, thin,
and the Greek lampas, f., lamp. Gender, feminine.
44 Proceedings of the Biological Society of Washington
oosD
0.5mm
Q 5
6,
)
ot
04
0
6
0
b
0
0
Wh
Fic. 1. Pedicellariae. a—c, Araeolampas atlantica n. sp., 92 mm TL,
USNM 24598, holotype; d-f, A. fulvua, 108 mm TL, USNM E889.
a, rostrate pedicellaria; b, small tridentate pedicellaria; c, large tridentate
pedicellaria; d, large tridentate pedicellaria; e, large tridentate pedicellaria
(side view of 1d); f, rostrate pedicellaria.
Araeolampas atlantica, new species
Figures la—c, 2a and b
Material Examined: 1 specimen (Holotype), 92 mm TL; off Virginia,
Albatross station 2105, 37°50’ N, 73°03.8’ W; 2575 m; 6 November 1883;
USNM 24598. — fragments of 1 specimen; off the Azores, Atlantis sta-
tion 20, 37°50.5’ N, 26°00’ W; 2585 m; 18 August 1948; USNM E7948.
— fragments of 1 specimen; west of Dry Tortugas, Gulf of Mexico; 3545
m; MCZ 2910. — 1 specimen, 82 mm TL and fragments of 2 specimens;
west of Georgia, Pillsbury station P-120, 31°48’ N, 76°38’ W; 1920 m;
USNM E13095. — fragments of 1 specimen; west of Gabon, Gulf of
Guinea, Pillsbury station P-292, 0°12’ N, 5°11’ E; 3595 m; 23 May 1965;
USNM E13096. — fragments of 1 specimen; north of Haiti, Pillsbury
station P-1429, 21°19.2’ N, 73°45.5’ W; 2532 m; USNM E13097.
Description: Test large (up to 92 mm TL), fragile; aboral aspect heart-
shaped (test width about 86 percent TL and test height about 37 percent
A New Heart-Urchin from North Atlantic 45
Frc. 2. a, aboral aspect of Araeolampas atlantica n. sp., 92 mm TL,
USNM 24598, holotype; b, lateral aspect of A. atlantica n. sp. (holo-
type); c, aboral aspect of A. fulva, 108 mm TL, USNM E889; d, lateral
aspect of A. fulva.
TL); periproct large (width plus height 19 to 24 percent TL), not
visible from above; slight indentation above periproct at ambitus; pos-
terior part of test truncate vertically; frontal notch deep (5 percent TL):
apical system anterior (32 percent TL from anterior ambitus) with 4
46 Proceedings of the Biological Society of Washington
TasLE 1. Morphometric characters of Araeolampas atlantica
and A. fulva.*
Distance
Test apex to Periproct Depth
—_———__——- anterior ———_———__ frontal
Species length width height ambitus width height notch
A. atlantica 91.9 86.0 37.0 31.9 Wore MILE 4.9
Sik5 33.0 FB MILL 4A
A. fulva 107.9 80.2 39.9 30.7 8.4 6.9 4.8
* Test length in mm; all other measurements expressed as percent TL.
well developed genital pores; peripetalous fasciole present; primary
spines restricted to interambulacral areas and to within the peripetalous
fasciole except for interambulacrum 5 (Loven’s system) where they
occur outside the peripetalous fasciole; areoles of primary tubercles about
2.8 percent TL: posterior part of areoles of primary tubercles about
2.8 percent TL; posterior part of areole deeply sunken, anterior part
slightly sunken; areoles produce slight internal bulges, but not deep
internal pouches as in Homolampas; primary tubercles perforate and
crenulate; subanal fasciole present; tridentate pedicellariae with wide
valves (Figure Ib and c); rostrate and triphyllous pedicellariae the same
as in A. fulva; pores of ambulacra near apical system single; oral surface
unknown.
Etymology: The specific name refers to occurrence in the Atlantic
Ocean.
Distribution: In the western North Atlantic A. atlantica occurs from
off Virginia to north of Haiti, and in the eastern North Atlantic from the
Azores to off Gabon. It is known from 1920 to 3595 m.
Taxonomic Affinities: The closest relative of A. atlantica (Figure 2a
and b) is A. fulva (Figure 2c and d) from the Pacific. A. fulva differs
from the new species by having tridentate pedicellariae with narrow
blades (Figure 1d and e), a smaller periproct and double ambulacral
pores adapically. The morphometric characters of these two species are
shown in Table 1. The Indian Ocean species A. glauca can be distin-
guished from A. atlantica by its shallow frontal notch.
DISCUSSION
Agassiz (1881) considered many of the generic differences discussed
above for Homolampas s.s. and Araeolampas n. gen. to be due to age. He
assumed that large specimens of H. fragilis would acquire a peripetalous
fasciole and four genital pores. Although some specimens of H. fragilis
do have four genital pores, these specimens are not necessarily the largest
specimens. No remnants of a peripetalous fasciole have ever been ob-
served on H. fragilis while small specimens of A. fulva and A. atlantica
n. sp. have both a distinct peripetalous fasciole and four well-developed
A New Heart-Urchin from North Atlantic AT
genital pores. The peripetalous fasciole is one of the more conservative
fascioles. No instance has ever been cited where this fasciole was lost
during ontogeny, as has been the case for some other fascioles (e.g.
subanal and lateroanal). All other generic characters mentioned above
are also present on small A. fulva and A. atlantica n. sp.
The large specimen identified by H. L. Clark (1941) as H. fragilis
(MCZ 2910) is really A. atlantica.
ACKNOWLEDGMENTS
I thank R. H. Chesher of the Marine Research Foundation for dis-
covering the first specimen of the new species and encouraging me to
describe it. H. B. Fell (MCZ), D. L. Pawson (USNM) and G. L. Voss
(UMML) provided me with specimens. F. J. Fell and D. L. Pawson
critically read the manuscript. Partial support was provided by NSF
grant GB-3532 to H. B. Fell and a Sigma Xi Grant-in-Aid of Research
to D. K. Serafy.
LITERATURE CITED
Acassiz, A. 1881. Report on the scientific results of the voyage of the
H.M.S. Challenger during the years 1873-76; Report on the
Echinoidea. Challenger Rep. 3(1):1-321, 45 pls.
Ciark, H. L. 1941. Reports on the scientific results of the Atlantis
Expeditions to the West Indies, under the joint auspices of the
University of Havana and Harvard University: The echino-
derms (other than holothurians). Mem. Soc. Cubana Hist. Nat.
15(1):1-154, 10 pls.
MortTENSEN, T. 1950. A Monograph of the Echinoidea. 5(1). Spat-
angoida I. C. A. Reitzel, Copenhagen, 432 pp., 25 pls.
48 Proceedings of the Biological Society of Washington
Vol. 87, No. 7, pp. 49-56 25 April 1974
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
IDENTIFICATION OF MYOTIS CALIFORNICUS AND
M. LEIBII IN SOUTHWESTERN NORTH AMERICA
By MicHaEL A. BOGAN
Bird and Mammal Laboratories
National Museum of Natural History
Washington, D.C. 20560
Within temperate North America there are, perhaps, no two
bats more similar than Myotis californicus (Audubon and
Bachman) and Myotis leibii (Audubon and Bachman). Over
much of their respective ranges they are allopatric, and speci-
mens may be identified on strictly geographic grounds. How-
ever in the southwestern United States and Mexico where their
ranges overlap broadly and where californicus is known to be
highly variable, problems of identification arise. Most students
of southwestern bats agree that the two kinds are closely
related but nonetheless distinct. For example, Findley (1972)
includes the two species in the “Leibii group” of the subgenus
Selysius within Myotis.
Most problems with the allocation of specimens appear to
center in the southwestern portion of New Mexico and ad-
jacent Arizona, Texas and Mexico (Barbour and Davis, 1969;
Anderson, 1972; Findley et al., in manuscript), probably due
to the increased variability of californicus in this area (Bogan,
1973). The collection of good series of both kinds of bats from
Hidalgo County, New Mexico, by J. D. Druecker and others
from the University of New Mexico offered the chance to
ascertain the status of the two bats in this area and, hopefully,
to establish reliable quantitative procedures for identifying
these bats, both in the field and in the museum.
METHODS AND MATERIALS
Thirty-four M. leibii and 50 M. californicus, all adults, from various
localities in Hidalgo County, New Mexico, were examined. Sexes were
7—Proc. Biot. Soc. WasH., Vou. 87, 1974 (49)
50 Proceedings of the Biological Society of Washington
TaBLE 1. Traditional characters used to separate Myotis californicus
and M. leibii.
Character californicus leibii
Facial coloration Brown Black
Ear color Brown Black
Dorsal pelage Dull-tipped, not Shiny-tipped,
burnished or glossy glossy or
burnished
Third metacarpal As long as forearm Shorter than
(over 30.5 mm) forearm (less
than 30.3 mm)
Braincase Rounded Flattened
Forehead profile Rises abruptly Rises gradually
Skull size Smaller Larger
combined for this analysis, as I have found that sexual dimorphism is not
marked in these two species. The fourteen characters utilized, measured
as described in Bogan (1973), are: total length, tail length, hindfoot
length, ear length, forearm length, tibia length, condyle-premaxillary
length, condylocanine length, maxillary toothrow length, cranial breadth,
cranial depth, least interorbital breadth, rostral breadth, rostral length.
The specimens were initially identified using only published characters
(Table 1). The crania were examined closely to determine additional
characters for identification. Bats were grouped into samples for sta-
tistical analysis in two ways: first, using just two samples, Hidalgo
County californicus and Hidalgo County leibii; and second, by sub-
dividing the two species into samples from eastern, central and western
Hidalgo County. These samples correspond to the three major mountain
ranges in Hidalgo County and likewise to most of the productive collecting
localities in the county (Findley and Traut, 1970).
Frequency distributions for each variable were prepared to examine
normality of distributions and degree of overlap between californicus
and leibii. Means and variances were then calculated for the two species.
Additionally, bivariate scattergrams were prepared for most pairs of
variables.
The data matrix was subjected to a two-group discriminant analysis
(BMD04M ) which gives a discriminant function as well as the dis-
criminant coefficients for each variable enabling the identification of
new specimens. Next, a stepwise discriminant analysis (BMD07M)
yielding five canonical variates and a classification matrix based on the
posterior probabilities and generalized distance values was performed,
allowing an estimate of the degree of phenetic overlap. Additionally,
the data were subjected to a principal component analysis (BMD0O1M)
and to a numerical taxonomic analysis (NTSYS, developed by F. James
Identification of Two Myotis Species 51
TABLE 2. Basic statistics for 14 variables of Myotis californicus and
M. leibii from Hidalgo County, New Mexico. The mean plus or minus
one standard deviation and the range (in parentheses) are shown for
each sample.
M. californicus M. leibii
Character (N = 50) (N = 84)
Total length 84.52 + 4.69 89.06 + 4.16
(73.0-94.0) (80.0-99.0)
Tail length 39.00 + 3.03 41.50 + 2.71
(32.0-45.0) (37.0-49.0)
Hindfoot length 6.82 + 0.92 7.71 = 0.62
(5.0-8.5) (6.0-9.0)
Ear length 137/22 1-03 14.70 + 0.97
(11.0-15.0) (12.0-16.0)
Forearm length COAG) a= 10 33.49 + 0.93
(30.45-35.22) (31.28-35.77)
Tibia length 13.98 = 0.83 13.88 = 0.60
(12.30-15.75) (13.11-15.34)
Condyle-premaxillary length 12.69 + 0.27 IBS se OM
(12.10-13.33) (12.73-13.88)
Condylocanine length 11.98 + 0.26 12.60 + 0.26
(11.50-12.60) (11.99-13.07)
Maxillary toothrow length 4.99 = 0.13 5.26 + 0.13
(4.64-5.25) (5.02-5.57)
Cranial breadth GP az Oz 6.49 + 0.13
(5.95-6.65) (6.24-6.79)
Cranial depth 4.58 + 0.17 4,42 + 0.13
(4.19-4.95) (4.13-4.80)
Interorbital breadth 3.09 + 0.09 3.25 == 0.14
(2.90-3.28) (2.91-3.52)
Rostral breadth 4.81 + 0.13 Sroot== 038
(4.51-5.10) (5.00-5.63)
Rostral length Foo = 028 5.56 + 0.26
(4.32-6.00) (5.02-6.30)
Rohlf and associates). These last two analyses supported the discriminant
analyses and therefore are not reported herein.
RESULTS AND DISCUSSION
An impressive array of investigators (Hall and Kelson, 1959; Cockrum,
1960; Barbour and Davis, 1969; Anderson, 1972; Armstrong, 1972;
Findley et al., in manuscript) have used the same basic set of characters
[first presented by Miller and Allen (1928)] to separate Myotis cali-
fornicus and M. leibii (Table 1). In the Southwest, however, no single
CRANIAL DEPTH
TABLE 3. Discriminant coefficients for original
in computing discriminant scores for unknowns.
Proceedings of the Biological Society of Washington
9.5
5.0
4.5
4.0
Fic. l.
M. californicus
45 4f¢ 90
9.9
ROSTRAL BREADTH
60
Relationships of two cranial measurements in Myotis cali-
fornicus and M. leibii. Measurements are in millimeters.
Variable
Total length
Tail length
Hindfoot length
Ear length
Forearm length
Tibia length
Condyle-premaxillary length
Condylocanine length
Maxillary toothrow length
Cranial breadth
Cranial depth
Interorbital breadth
Rostral breadth
Rostral length
variables to be used
See text for details.
Coefficient
0.00508
0.00019
—0.02895
—0.01790
0.01046
0.02176
—0.18182
0.02349
—0.06011
—0.00339
0.30601
0.14769
—0.43839
0.07154
Identification of Two Myotis Species 53
californicus leibii
FREQUENCY
33 BO) TBE ae | qn fee
DISCRIMINANT FUNCTION
Fic. 2. Frequency distribution of Hidalgo County samples of Myotis
californicus and M. leibii on the discriminant function computed by the
two-group discriminant analysis.
qualitative character provides certain identification. Rather, all such
characters must be considered simultaneously in a typically time-
consuming operation. The quantitative character most often cited, that
of third metacarpal length, was so variable within californicus that it
was not further considered as an effective means to distinguish cali-
fornicus from leibii. The basic statistics and amount of character over-
lap between the two species in Hidalgo County are shown in Table 2.
The character showing the least amount of overlap in my analysis is
rostral breadth as measured at the junction of M:i—M2; 96% of cali-
fornicus not exceeding 5.0 mm, and 92% of leibii equalling or exceeding
5.2 mm.
Most of the bivariate scattergrams were useless for separating the two
species. The exceptions were rostral breadth plotted against other skull
parameters; the most discriminating being rostral breadth against cranial
depth (Figure 1). This useful combination is a reflection of Myotis
leibii having a flattened skull, and M. californicus possessing a more
globose skull. I have subsequently plotted rostral breadth against cranial
depth for numerous specimens of both species from elsewhere in the
southwestern U. S. and Mexico with equally good separation. Only rarely
do specimens overlap the 52° line separating the clusters of leibii and
californicus.
The results of the bivariate scattergram of Figure 1 were extended by
performing a discriminant analysis to maximize the separation of the two
species of bats. The results of this analysis are presented in Figure 2,
and show a clear separation between the two groups. This analysis
yielded the discriminant coefficients listed in Table 3 which can be used
in determining identification of unknowns. The procedure, described in
54 Proceedings of the Biological Society of Washington
: / L
East
-3.300 i LWest
| / Least Central
ib
LWest Central
leibii
californicus
6) -|.800 1.200 4.200 7.200
-3.300 -.300 2.700 5.700
JL
Fic. 3. Distribution of Myotis californicus and M. leibii on the first
two canonical variates computed by the stepwise discriminant analysis.
Each species is represented by three samples from western, central and
eastern Hidalgo County. Lines enclose all bats within a sample.
- 4.80
-6.300
Choate (1973) and elsewhere, is to multiply the value for each character
by the discriminant coefficient, sum these values, and then plot the
value on the discriminant function as shown in Figure 2. As Choate
(1973) points out, the importance of extreme care in taking the measure-
ments cannot be overstressed.
The multiple stepwise discriminant analysis using six groups (three
californicus and three leibii samples) is of interest for at least two
reasons. First, since the variables are entered in a stepwise fashion, the
analysis enables the investigator to determine which variables possess the
greatest discriminating power; and second, since the samples are sub-
divided it is possible to define a five-dimensional space within which the
bats are dispersed. As seen in Table 4 the variable possessing the greatest
discriminatory power among the 14 original variables is rostral breadth.
Figure 3 is a plot of the six samples on the first two canonical variates
and clearly shows that the two species do not overlap. The first two
canonical variates account for 84% and 8% of the variance, respectively,
and the first four variates account for 98.7% of the total variance.
These analyses demonstrate the distinct nature of the two taxa, and
thus support the opinions of other investigators, and they enable the
Identification of Two Myotis Species 5D
TaBLE 4, Rank order of variables as determined by the stepwise dis-
criminant analysis. Those variables at the top of the column possess the
greatest discriminatory power.
Rank Variable
i Rostral breadth
D Hindfoot length
3 Cranial depth
4 Ear length
3S Maxillary toothrow length
6 Total length
7 Cranial breadth
8 Tibia length
9 Rostral length
10 Condyle-premaxillary length
ILL Condylocanine length
12 Interorbital breadth
13 Forearm length
14 Tail length
rapid allocation of specimens of unknown identity. Proper identification,
however, requires the presence of a clean, intact skull. It is disappointing
that this study did not reveal any completely reliable character facilitating
positive identification of these species in the field. Field identification
still requires the utilization of traditional characters coupled with con-
siderable experience. The most important feature is that leibii is
usually darker in color than californicus. The exceptions are found in
populations of dark-colored californicus occurring at higher elevations in
the southwestern U. S. and Mexico. Furthermore, californicus usually
appears to be distinctly smaller and more aggressive than leibii.
Having demonstrated that the two taxa are distinctive the intriguing
question is, how are they partitioning the environment in areas of
sympatry? Black (1972), based on his study of bat food habits, sug-
gests that one member of the pair might be a “beetle strategist” while
the other may prey more heavily on moths. Husar (1973) has recently
described such a situation in Myotis evotis and M. auriculus, two very
similar bats with rather narrow regions of overlap in the southwest.
Geluso (1972) has shown that californicus and leibii are physiologically
different in urine concentrating abilities and in kidney morphology. These
investigations suggest that californicus and leibii are at least as different
ecologically and physiologically as they are morphologically and they
may be minimizing competition through differences in diet, roosting or
foraging sites, or foraging times. Such partitioning should be demon-
strable through observable differences in dietary, behavioral, or physio-
logical parameters.
56 Proceedings of the Biological Society of Washington
ACKNOWLEDGMENTS
Several people and institutions were instrumental in bringing this
report to completion. The University of New Mexico and the Smithsonian
Institution provided computer time. James S. Findley generously assisted
in initial stages of identification of bats. Alfred L. Gardner, Charles O.
Handley, Jr., Clyde Jones, Richard W. Thorington, Jr., and Don E.
Wilson all read and commented on the manuscript. To all the above
I am grateful.
LITERATURE CITED
ANDERSON, S. 1972. Mammals of Chihuahua. Bull. Amer. Mus. Nat.
Hist. 148:149—410.
ARMSTRONG, D. M. 1972. Distribution of mammals of Colorado. Mono-
graph, Mus. Nat. Hist., Univ. Kansas 3:1-415.
Barsour, R. W. ann W. H. Davis. 1969. Bats of North America.
Univ. Press Kentucky, Lexington. pp. 285.
Buack, H. L. 1972. A north temperate bat community: structure,
food habits and prey community. Unpubl. Ph.D. Dissertation,
Univ. New Mexico, Albuquerque.
Bocan, M. A. 1973. Geographic variation in Myotis californicus in
the southwestern United States and Mexico. Unpubl. Ph.D.
Dissertation, Univ. New Mexico, Albuquerque.
CuHoateE, J. R. 1973. Identification and recent distribution of white-
footed mice (Peromyscus) in New England. J. Mammal. 54:
41-49.
Cocxrum, E. L. 1960. The recent mammals of Arizona: their tax-
onomy and distribution. Univ. Arizona Press, Tucson. viii +
276 pp.
FINDLEY, J. S. 1972. Phenetic relationships among bats of the genus
Myotis. Systematic Zool. 21:31-52.
FINDLEY, J. S., A. H. Harris, C. J. Jones anD D. E. Witson. Mammals
of New Mexico. Manuscript.
FINDLEY, J. S. and G. L. Traut. 1970. Geographic variation in
Pipistrellus hesperus. J. Mammal. 51:741-765.
Getuso, K. N. 1972. Urine concentrating abilities, kidney structure
and drinking water dependence of insectivorous bats. Unpubl.
Ph.D. Dissertation, Univ. New Mexico, Albuquerque.
Hau, E. R. anp K. R. KEtson. 1959. The mammals of North America.
Vol. I. Ronald Press, New York. xxx + 546 + 79 pp.
Husar, S. L. 1973. Behavioral character displacement: evidence of
food limitation in bats. Unpubl. M.S. Thesis. Univ. New
Mexico, Albuquerque.
Miter, G. S., Jr. anD G. M. ALLEN. 1928. The American bats of the
genera Myotis and Pizonyx. U. S. Nat. Mus. Bull. 144:viii ++
218 pp.
Vol. 87, No. 8, pp. 57-72 25 April 1974
PROCEEDINGS
OF TEE
BIOLOGICAL SOCIETY OF WASHINGTON
AN EMENDATION OF THE GENUS TRIANNULATA
GOODNIGHT, 1940, WITH THE ASSIGNMENT OF
TRIANNULATA MONTANA TO CAMBARINCOLA
ELLIS 1912 (CLITELLATA:
BRANCHIOBDELLIDA )
By Perry C. Hout
Department of Biology and
Center for Systematic Collections
Virginia Polytechnic Institute and State University
Blacksburg, Virginia 24061
The nominal genus Triannulata Goodnight, 1940, is com-
posed of two species, T. magna Goodnight, 1940, the type-
species, and T. montana Goodnight, 1940. Superficially, these
large worms resemble members of the genus Cambarincola
Ellis, 1912, and I thought they were species of the latter when
I began this study. But magna is a representative of a dis-
tinctive, and as of now, monotypic, genus, while montana is
indeed a member of the genus Cambarincola. The following
redefinition of the genus Triannulata, a redescription of T.
magna and the reassignment and emended description of
montana are offered as a part of my ongoing effort to describe
and classify the North American branchiobdellid fauna.
Other than the original treatment of these species (Good-
night, 1940:56-58), their possible inclusion in such compendia
as Pennak (1953:300) and brief statements in Hoffman
(1963:281, 295), Liang (1963:570) and Holt (1969:195),
nothing else, to my knowledge, has been written about them.
In former works (Holt, 1965; 1968a; 1969), I ignored
Triannulata, believing its species to belong to Cambarincola.
The illustrations herein are so oriented that the anterior of
all animals, or parts thereof, is to the reader’s right. Measure-
ments given are approximations, roughly correct to the nearest
8—Proc. Biot. Soc. WaAsH., VoL. 87, 1974 (57)
58 Proceedings of the Biological Society of Washington
0.1 mm. The animals are large and the details shown in the
illustrations are essentially free-hand sketches, based upon pro-
portions established with the camera lucida. Further, the
drawings are conventionalized (as in all my works on the
branchiobdellids): stippling indicates glandular cells or struc-
tures; line hatching, muscular structures or investments. The
initials “PCH” indicate collection numbers of branchiobdellids
in the collections of the VPI&SU Center for Systematic
Collections.
I am grateful to Mrs. Virgie F. Holt, my wife and constant
field companion, for helping in the collecting of topotypical
material and other specimens treated herein; to Dr. Marian H.
Pettibone, Curator, Division of Worms, Department of In-
vertebrate Zoology, National Museum of Natural History,
Smithsonian Institution, for allowing me to examine the holo-
types of Triannulata magna and Cambarincola montanus; to
Dr. Horton H. Hobbs, Jr., for supplying the identifications of
the host animals and for a critical reading of the manuscript; to
the National Science Foundation (grant GB-372) for financial
support of my field work.
Goodnight (1940:56) presented the following diagnosis of
the genus Triannulata: “With the characteristics of the sub-
family; spermatheca not bifid; no accessory sperm tube; body
cylindrical, not flattened; head roughly triangular in shape
with protruding lips; major annuli of most segments redivided
to give the appearance of three annuli per segment; this is
especially evident in the median segments and moderately
contracted specimens; anterior nephridia opening to the out-
side through separate pores in the dorsal half of segment III.”
When this diagnosis is examined in the light of our knowl-
edge of the branchiobdellids as it now exists, it is found to be
inadequate.
“With the characteristics of the subfamily” means that the
worms produce spermatozoa in both segments V and VI
(Goodnight, 1940:27), instead of possessing testes and male
funnels only in segment V. The latter is true of species of the
Eurasian genus Branchiobdella Odier, 1823, but is not known
for any North American genus (Holt, 1967:8). In including
the statement, “no accessory sperm tube” in his diagnosis,
An Emendation of Triannulata 59
Goodnight was referring to what is now known as the prostate
(Holt, 1960:63). Triannulata magna clearly does not possess
a prostate; montana is characterized by a very large and dis-
tinctive one. The only known genus with a bifid spermatheca
is the monotypic, North American Bdellodrilus Moore, 1895.
Members of the American genera Xironodrilus Ellis, 1918, and
Xironogiton Ellis, 1919, are flattened, rather than terete, and
are not at all closely related; the remaining 15 genera of the
order, including Triannulata, are composed of animals with
cylindrical bodies. The “triangular” shape of the head (“cone-
shaped” would be more descriptive) with protruding lips are
minor features, worthy of mention only as specific characters,
and cannot be taken seriously as distinctive of a genus, since
many intrageneric variations of this type are known among
the branchiobdellids (cf. Hoffman’s 1963 treatment of
Cambarincola). The same objection applies to the use of
“redivided” major annuli as a generic character. Not all of
the segments of members of the two species that Goodnight
placed in Triannulata present a tripartite appearance; in
those that do, the redivision takes the form, mainly ventrally,
of a rather minute diminution of the diameter of the anterior
annulus immediately anterior to the usual distinct sulcus that
sets off the minor (posterior and shorter) annulus of a seg-
ment. This tripartite appearance of some segments may simply
be a consequence of the large size of the worms and is also
characteristic of the anterior segments of the likewise large
Cambarincola ingens Hoffman, 1963, and Stephanodrilus
(= Cirrodrilus) truncatus Liang, 1963. [See Holt (1967:2-3)
for a discussion of the synonymy of Cirrodrilus Pierantoni,
1905, and Stephanodrilus Pierantoni, 1906].
Goodnight’s belief that Triannulata is characterized by
separate nephridiopores on the dorsum of segment III must be
considered carefully; the manner of opening of the anterior
nephridia is a recognized generic character. T. magna is ge-
nerically distinct on the basis of features of the reproductive
systems, but Goodnight appears to be mistaken in his statement
that the dorsum of segment III bears two nephridiopores. The
holotype is mounted with the dorsal side uppermost on the
slide and I cannot find two nephridiopores; rather, middorsally,
60 Proceedings of the Biological Society of Washington
there appears to be a single pore, but the nephridia are very
difficult to see in this specimen and impossible to trace un-
mistakably to their opening. Immature specimens from the
type locality, that I am confident on the basis of body shape
and the structure of the jaws are conspecific with the type,
have only one nephridiopore. In all branchiobdellids, the
nephridiopores are frequently difficult to locate unambiguously
in specimens mounted entire. Often, not always, their position
can only be determined in mature specimens by means of
serial sections. Though Goodnight (1940:8-9) mentioned the
use of serial sections, nowhere does he refer to their use in the
discussion of any species in his monograph or present any
drawings or photographs of sections. This is noticeably true
of his treatment of T. magna. Unfortunately, I was able to take
only a few specimens of T. magna in the limited time I could
devote to collecting in Washington and Oregon. Of these,
I chose to dissect for a study of the reproductive systems all
but one which is mounted entire. Yet for the reasons cited, I
am convinced that Triannulata is partly characterized by a
common opening of the anterior nephridia. The point is im-
portant for any consideration of the phylogenetic relations of
the genus. |
Part of these conclusions were anticipated by Liang (1963:
570) on the basis of an anlysis of Goodnight’s descriptions.
That is, Liang was unable to distinguish between Goodnight’s
description of Triannulata and Stephanodrilus (= Cirrodrilus)
and placed both Triannulata and Stephanodrilus (= Magmato-
drilus Holt, 1967) obscurus (Goodnight, 1940) together in the
invalid genus Stephanodrilus. Liang adequately described and
illustrated the reproductive systems of his Chinese worms,
accepting my earlier contentions that only detailed study of
the reproductive systems furnish an adequate basis for the
classification of the branchiobdellids (Holt, 1949, et seq.;
Hoffman, 1963.
Triannulata Goodnight, 1940
Triannulata Goodnight, 1940:56-58—Pennak, 1953:300.—Hoffman,
1963:281, 295.—Holt, 1969:195.
Stephanodrilus.—Liang, 1963:570 [in part].
An Emendation of Triannulata 61
Type-species: Triannulata magna Goodnight, 1940:56-57, by original
designation.
Diagnosis: Moderately large branchiobdellids (about 4.5 mm in aver-
age length); 2 pairs of testes; unpaired nephridiopore on dorsum of
segment III; body terete, without peristomal tentacles or dorsal pro-
jections, head large, lips (peristomium) prominent; some, mostly III-V,
segments superficially triannulate; spermiducal gland large, subspherical
to subcubical, vasa deferentia entering entally, with small deferent lobes,
without prostate or prostatic protuberance; no ejaculatory duct; bursa
with large ectal spherical atrium, long muscular eversible penial sheath;
spermatheca with thick outer muscular wall, internally essentially filled
with tall columnar epithelial cells, lumen reduced.
Distribution and affinities: With the removal of montana from the
genus, Triannulata becomes monotypic. Goodnight (1940:57) records
T. magna from two localities in Washington and three in Oregon. I
have material from the type-locality and one other locality in Washington.
The species is probably widespread in the Cascade and Coastal Ranges of
the Pacific Northwest.
The affinities of the genus must be sought among a group of pre-
sumably primitive branchiobdellids among the Sathodrilus—lineage whose
relationships have been discussed (Holt, 1969:195-198; 1973:35):
Sathodrilus Holt, 1968b; Ceratodrilus Hall, 1914; Magmatodrilus. In
addition, the Asiatic genus Caridinophila Liang, 1963, must be included
as a possible relative of these genera.
On the assumption that an eversible penis and the ental entry of the
vasa deferentia into the spermiducal gland are primitive features, Satho-
drilus has been placed near the beginning of a lineage that culminates
in such genera as Pterodrilus and Cambarincola with single anterior
nephridiopores, well defined ejaculatory ducts, prostates always present,
ental points of entry of the vasa deferentia into the spermiducal gland
and protrusible, cone-shaped muscular penes.
The species of Sathodrilus have single anterior nephidiopores, ejacu-
latory ducts that usually are short and relatively thick, prostates or
prostatic protuberances may or may not be present, the vasa deferentia
always enter the ental end of the spermiducal gland and the penes are
eversible, though their finer structure differs among the species that are
included in the genus. Sathodrilus ranges from northwestern South
Carolina to southern Mexico (Holt, 1973a; 1973b).
Ceratodrilus is composed of two allopatric species from the Great
Salt Lake and Snake River drainages. The anterior nephridiopore is
single. A well defined, but relatively short though prominent ejaculatory
duct, a prostatic protuberance, ental entry into the spermiducal gland of
the vasa deferentia and an eversible penis are shared with other members
of the Sathodrilus lineage. The genus is distinguished by long peristomal
and dorsal tentacles and appendages (Holt, 1960).
Magmatodrilus, a monotypic genus from northern California, is
similar to Ceratodrilus in the above respects, except that the places of
62 Proceedings of the Biological Society of Washington
a aa eecr
a ee
Fic. 1. Triannulata magna. a, holotype, ventral view; b, same, outline
of ventral jaw; c, same, ventral view of reproductive systems.
entry of the vasa deferentia into the spermiducal gland are widely
separated, the latter is long and slender without any indications of a
prostatic protuberance, the penial sheath enclosing the eversible [not
protrusible, contra Holt, 1967] penis is much shorter, tentacles and
dorsal projections are absent (Fig. 4).
The Chinese genus Cardinophila has no spermatheca, the spermiducal
gland is small and there are no vasa deferentia (the vasa efferentia
enter the spermiducal gland at four separate places) and an ejaculatory
duct is present (Liang, 1963: 569). Nothing is known of the penis of
C. unidens.
The condensed account just given of the major features of these genera
constitute convincing evidence of their close phylogenetic relationships.
Their generic distinctiveness is attested by the differences mentioned in
addition to those in the overall facies of the jaws, the presence (in
Ceratodrilus) or absence of peristomial and dorsal body appendages and
striking, but difficult to describe succinctly (see Liang, 1963, and Holt,
1960; 1967; 1968a; 1969) variations in the minor features of the
reproductive systems.
An Emendation of Triannulata 63
is Ee v = ue
Zz f ; \ \S
Uf | i Ne
Fic. 2. Triannulata magna. Latero-dorsal view of reproductive sys-
tems from a dissection: ba, bursal atrium; ps, penial sheath; sb, sperma-
thecal bulb; sd, spermathecal duct; sq, spermiducal gland; vd, vas
deferens.
It is obvious that Triannulata is related to this group of genera. The
absence of an ejaculatory duct in Triannulata immediately separates it
from the others. Or, if one wishes to consider what I have called the
penial sheath of Triannulata an ejaculatory duct, the eversibility of the
ejaculatory duct does so. But until more study is devoted to these struc-
tures of the male reproductive system of the branchiobdellids, I cannot
more precisely place the genus in the Sathodrilus lineage. I can only
say that these four genera are closely related, that they appear to be
phylogenetic relics and that guesses as to which is nearer in structure
to the postulated ancestor of branchiobdellids with a single anterior
nephridiopore are futile.
Triannulata magna Goodnight, 1940
Figures 1-3
Type-specimens: Holotype, USNM 20567 from Naches, Washington,
on Pacifastacus sp.
Diagnosis: As for the genus.
Description: The worms are large, the holotype, the only mature speci-
men I have seen from the type-locality, has the following dimensions:
total length, 3.8 mm; greatest diameter, segment VII, 1.2 mm; head
length, 0.9 mm; head diameter, 0.9 mm; diameter, segment I, 0.6 mm;
64 Proceedings of the Biological Society of Washington
Fics. 3-4. Longitudinal section of male copulatory apparatus. Fig.
3. Triannulata magna. Fig. 4. Magmatodrilus obscurus: ba, bursal
atrium; ejd, ejaculatory duct; p, penis; pl, lumen of penis; ps, penial
sheath.
diameter, sucker, 0.8 mm. A specimen from Cowlitz County, Wash-
ington, is somewhat larger with the following dimensions: total length,
4.9 mm; greatest diameter, 1.2 mm; head length, 1.1 mm; head diameter,
1.0 mm; diameter, segment I, 0.7 mm; diameter, sucker, 0.9 mm. The
other two mature specimens from Cowlitz County that I have examined
were dissected for a study of the jaws and reproductive systems, but
were of comparable size.
The peristomium is divided by lateral indentations into upper and
lower lips that lack emarginations or lobes, but that noticeably protrude
and are less in diameter than the head. There are no detectable oral
papillae. The head, execpt for the region of lessening diameter of the
peristomium, has no external sulci and internally there is oa one,
though prominent, pharyngeal sulcus.
The anterior annuli of the trunk segments are only very slightly
An Emendation of Triannulata 65
greater in diameter than the posterior ones, hence there are no noticeable
dorsal ridges. The anterior annuli of segments I-V are subdivided, a
subdivision that appears rather superficial and does not involve the longi-
tudinal muscles. The anterior nephridiopore has been mentioned above.
The jaws are subequal in size, triangular in en face view, without
lateral teeth, therefore, a 1/1 dental formula. They are dark brown.
The following description of the reproductive system is based upon
dissected material and it is difficult or impossible to present the relative
size of its components in comparison to the diameter of the animals.
The spermiducal gland is large, subspherical, almost subcubical in
shape. Its ventral border is at the level of the dorsal border of the
bursa and the vasa deferentia enter it at opposite angles of its ventral
(ental) side. There are small, indistinct deferent lobes that are not at
all prominent. A prostrate is totally absent.
The penis of the branchiobdellids is the ectal end of the ejaculatory
duct. For these traditionally separately described organs in Triannulata
magna, two interpretations are immediately suggested by the bursal-
penial complex: (1) the ejaculatory duct is absent and the penial sheath
is long and in its totality eversible as the penis; or (2) the ejaculatory
duct is itself a heavily muscular, eversible organ with only its ectal
portion representing the penis. In actuality, neither interpretation is
satisfying. I have chosen to interpret the eversible, presumably intro-
mittent portion of the male copulatory apparatus of T. magna as the
penis and deny the animal an ejaculatory duct. At a deeper level of
interpretation the problem is probably semantic. It is most likely
that in the ancestors of the branchiobdellids a muscular tube, undiffer-
entiated into penial sheath, penial and ejaculatory duct portions, was
everted as the intromittent organ through a somewhat heavier muscular,
but small, bursal atrium. T. magna, with an increase in the muscularity
of this intromittent organ, has, then retained this postulated ancestral
arrangement.
In any case, adopting the first of the interpretations above for the
sake of simplicity of description, the eversible penis, comprised of what
in other members of the order would be known as the ejaculatory duct
plus the penis, is composed of, other than the investing peritoneum, a
prominent outer layer of encircling muscles and a much thicker layer
of longitudinal (in reference to the organ itself) ones. The lumen is
distinct throughout and convoluted, particularly ectally.
The bursa is spherical, with a rather short outlet duct, and is composed
almost entirely of the atrial portion: the penis projects outward only a
short distance into what in other branchiobdellids is the penial sheath.
The organ is, nonetheless, relatively large.
The spermatheca has a comparatively short ectal duct; most of the
organ consists of a clavate bulb with a thick muscular investment and
very tall columnar glandular cells which almost obliterate the lumen,
leaving only a minute space near the median portion for the storage of
spermatozoa.
66 Proceedings of the Biological Society of Washington
OV
Fic. 5. Cambarincola montanus. a, lateral view of animal from
Douglas County, Oregon; b, same, lateral view of jaws; c, same, lateral
view of reproductive systems.
Variation: In the limited material at my disposal, no significant varia-
tions are detectable. Segment VI may be sometimes triannulate, but on
the other hand it may always be so and the extra “annulus” is obscured
in the holotype and other specimens I have seen.
Affinities: Discussed above as those of the genus.
Distribution: As discussed above for the genus.
Hosts: Pacifastacus |. leniusculus (Dana, 1852) and P. I. klamathensis
(Stimpson, 1857).
Material examined: The holotype, several immature topotypical speci-
mens (PCH 1811) taken on Pacifastacus leniusculus klamathensis from
the Naches River just above its confluence with the Tieton River,
Yakima County, Washington, 13 August 1964, by Perry C. and Virgie F.
Holt.—4 specimens (PCH 1814) taken from the Kalami River on P. I.
leniusculus about 8 miles south of Kelso, Cowlitz County, Washington,
15 August 1964, by Perry C. and Virgie F. Holt.
Cambarincola Ellis, 1912
Astacobdella Leidy, 1851:206.
Branchiobdella——Moore, 1894:427 [in part].
Bdellodrilus.—Pierantoni, 1912:24 [in part].
Cambarincola.—Ellis, 1912:481; 1919:263.—Hall, 1914:190.—Stephen-
son, 1930:801.—Yamaguchi, 1932:454; 1934:189.—Goodnight, 1940:
30.—Holt and Hoffman, 1959:97.—Hoffman, 1963:271.—Hobbs, Holt
and Walton, 1967:52.—Holt, 1969:197; 1973a:84; 1973b:9.
Diagnosis (modified from Hobbs, Holt and Walton, 1967:52): Body
terete without specialized projections other than peristomial tentacles in
some species; anterior nephridia opening through common pore on
An Emendation of Triannulata 67
dorsum of segment III; deferent ducts entering ental end of spermiducal
gland; prostate and ejaculatory duct both present; penis non-eversible;
bursa subpyriform to obcordate; spermatheca present, never bifid.
Type-species: Cambarincola macrodonta Ellis, 1912, by original
designation.
Cambarincola montanus (Goodnight, 1940), new comb.
Figure 5
Triannulata montana Goodnight, 1940:57.—Pennak, 1953:300.—Hoff-
man, 1963:281, 295.—Liang, 1963:570.—Holt, 1969:195.
Type-specimens: Holotype, USNM 2056, from the Kalami River, Wash-
ington, on Pacifastacus sp.
Description: Goodnight’s description of Triannulata magna is con-
fined almost entirely to features of the body and jaws and omits any
diagnostic reference to the reproductive systems. In the following
emended description of Cambaricola montanus, I shall quote all of
Goodnight’s relevant statements while adhering to my previously de-
veloped format for species descriptions.
Specimens of Cambarincola montanus are large worms. Averages of 5
mature individuals from Polk County, Oregon, selected at random, have
the following dimensions (ranges in parentheses): total length, 5.8 mm
(4.8-6.3 mm); greatest diameter, 0.8 mm (0.6—-1.0 mm); head length,
1.0 mm (0.9-1.4 mm); head diameter, 0.7 mm (0.5-0.8 mm); diameter
segment I, 0.6 mm (0.4—0.7 mm); diameter, sucker, 0.6 mm (0.5-0.8
mm). These measurements are concordant with Goodnight’s description
(1940:57) of a worm 5.0 mm long.
The peristomium (lips) are provided with 4 dorsal tentacles, 2
lateral lobes on each side and 4 ventral lobes, which accords with Good-
night’s (1940:57) statement “peristomium divided into twelve lobes . . .
which may be extended into tentacular appendages, dorsal longer than
ventral or lateral.” Experience with a variety of species, e.g., those
of Ceratodrilus (Holt, 1960) and Cambarincola fallax Hoffman, 1963,
enables one to distinguish between lobes and tentacles of the lips. No
oral papillae are detectable. The peristomium, lateral indentations of
which form the lips of the branchiobdellids, is set off from the remainder
of the head by a marked narrowing in diameter, often with about 3
annular indentations. There are no other external sulci of the head and
only one prominent internal (pharyngeal) sulcus. The marked narrow-
ing of the peristomium and the relatively large diameter of the head in
contrast to the lesser diameter of trunk segment I confer a distinct
cone-shaped appearance to the head.
The anterior (major) annuli of the trunk segments are not noticeably
greater in diameter than the posterior ones, there are, therefore, no
dorsal ridges, but in segments III to V the major annuli are subdivided
ventrally to give a triannulate appearance to these segments. The anterior
nephridiopore is not prominent, but contra Goodnight (1940:57) it is
68 Proceedings of the Biological Society of Washington
a single pore located in the usual position on the dorsum of the major
annulus of segment III.
The jaws are massive in appearance, but not disproportionately large,
and dark brown. They are triangular in shape in en face view and sub-
retangular to rounded triangular in lateral view. The usual dental
formula is 1/1 with prominent blunt teeth. Younger, though large, speci-
mens from the Kalami River, the type-locality, have a dental formula of
5/5, but the lateral teeth are obscure and probably wear away with age,
so I cannot dispute Goodnight’s statement that the dental formula is 7/5.
There may well be this much variation with age in the number of
lateral teeth.
Goodnight (1940:57) has nothing further to say of any diagnostic
value, remarking merely that the reproductive organs are in their normal
position in segments V and VI. The male reproductive system of
Cambarincola montanus furnishes, however, one of the most distinctive
features of the species. In its totality it is of normal proportional size
for the genus, but the worms are large and its components are often
compressed underneath the gut.
The spermiducal gland, though, is relatively small, in length, about
1/3 the diameter of segment VI, usually flexed and partially obscured in
whole mounts by the prostate. The vasa deferentia enter the gland at
widely separated regions, that is, there is a prominent anterior deferent
lobe. Otherwise, it is not unusual.
The prostate is the truly remarkable feature of the male system.
Proportionately huge, it exceeds somewhat the spermiducal gland in
length and is at least 144 times the diameter of the latter. Yet it is
composed of only the usual single layer of columnar glandular cells
which are highly vacuolated, that is, differentiated. There is no ental
bulb of the prostate.
The ejaculatory duct is relatively short and thick. The bursa is
somewhat more than 14 the diameter of its segment, a little more than
Y% its length in breadth and subpyriform in shape. Internally, its
structure is typical of that of other species of the genus, a short penial
sheath enclosing a cone-shaped protrusible penis, a rather short atrial
portion and relatively long bursal “outlet canal.”
The spermatheca is composed of a long ectal duct and a globose bulb.
Because of the bending of the ectal duct, the total length of the
spermatheca is difficult to estimate, but it is about 14 the diameter of
segment V.
Variation: There is considerable variation in the size of mature
animals, but all are larger than those of most members of the genus. The
reproductive systems may appear to differ from one worm to another,
because of the differing positions in which they lie with reference to the
gut. The peristomial tentacles may vary in degree of extension, which is
of no consequence; they are always distinctly tentaculate and borne on the
dorsal lobes of the upper lip. More significantly, the jaws of most speci-
mens appear to bear only one tooth each, with undulations along the
An Emendation of Triannulata 69
normally tooth-bearing margins, but as remarked, this is probably a
function of wear and the dental formula in young animals is either 7/5
or 5/5, possibly varying between these two.
Affinities; At this stage of our knowledge of the genus Cambarincola,
it is futile to speculate as to which of its species is closest to C. montanus.
The differentiated prostate, the peristomal tentacles, and almost surely
the large total size mark it as an advanced member of the genus. The
lack of a prostatic bulb removes it from the philadelphicus section
(Hoffman, 1963), yet in overall facies C. montanus is closest to mem-
bers of this no longer valid section (Holt, 1973a, b) of the genus. Of
the much better known eastern species of the genus, C. ingens Hoffman,
1963, rivals or exceeds C. montanus in size, but the prostate is much
longer than the spermiducal gland, less in diameter, and possesses an
ental bulb in the former. C. fallax Hoffman, 1963, has four dorsal
peristomial tentacles and a 5/5 dental formula, but otherwise is a much
smaller worm without the discordance in size of the prostate (which also
has an ental bulb) and the spermiducal gland. Further comments must
await a detailed study of the genus in western North America. For the
present, the size of the total animal, the prominence of the tentacles of
the upper lip and the size and nature of the prostate readily separate
C. montanus from all of its known congeners.
Hosts: Pacifastacus leniusculus leniusculus (Dana, 1852), P. l.
klamathensis (Stimpson, 1857), P. I. trowbridgii (Stimpson, 1857).
Distribution: Streams of the Coastal and Cascade Ranges of the
Pacific drainage in western North America from Santa Barbara County,
California, to northern Washington.
Material examined (all collected by Perry C. and Virgie F. Holt): 5
specimens (PCH 1110) taken on Pacifastacus leniusculus klamathensis
from Myrtle Creek, 8.3 miles south of Tiller, on state highway 42, Douglas
County, Oregon, 11 July 1960.—7 specimens (PCH 1113) taken on P. l.
klamathensis from stream tributary to the Umpqua River, 12.6 miles south
of junction of state highway 23 and U. S. highway 99 on U. S. 99,
Douglas County, Oregon, 11 July 1960.—2 specimens (PCH 1116)
taken on P. |. leniusculus from Mary’s River at Philomath, Benton County,
Oregon, 12 July 1960.—2 specimens (PCH 1119) taken on P. l.
klamathensis from a small tributary to the Yaquina River, 14.4 miles east
of Toledo, Lincoln County, Oregon, 12 July 1960.—5 specimens (PCH
1124) taken on P. 1. klamathensis from South Yamhill River, 1.6 miles
west of Valley Junction, Polk County, Oregon, 13 July 1960.—5 speci-
mens (PCH 1127) taken on P. I. klamathensis from Butte Creek, on
state highway 213 at Marion-Clackamas County line, Oregon, 13 July
1960.—5 specimens (PCH 1130) taken on P. 1. klamathensis from Gray’s
River on U. S. Highway 830, Wahkiakum County, Washington, 14
July 1960.—One specimen (PCH 1133) taken on P. I. klamathensis from
Humptulips River at Humptulips, Gray’s Harbor County, Washington,
16 July 1960.—2 specimens (PCH 1137) taken on P. I. trowbridgii from
Mill Creek, 0.9 miles south of Forks on U. S. Highway 101, Clallam
70 Proceedings of the Biological Society of Washington
County, Washington, 16 July 1960.—3 specimens (PCH 1813) taken on
P. |. trowbridgii from the Chehalis River at Adna, Lewis County, Wash-
ington, 15 August 1964.—4 specimens (PCH 1814) taken on P. l.
leniusculus from the Kalami River about 8 miles south of Kelso, Cowlitz
County, Washington, 15 August 1964.
LITERATURE CITED
Evuis, Max M. 1912. A new discodrilid from Colorado. Proc. U. S.
Nat. Mus. 42(1912):481—486.
1918. Branchiobdellid worms from Michigan crayfish. Trans.
Amer. Microsc. Soc. 37:49-51.
1919. The branchiobdellid worms in the collections of the
United States National Museum, with descriptions of new
genera and new species. Proc. U.S. Nat. Mus. 55:241—265.
GooDNIGHT, CLARENCE J. 1940. The Branchiobdellidae of North
American crayfishes. Illinois Biol. Monogr. 17(3):1-75.
Hatt, Maurice C. 1914. Descriptions of a new genus and a new
species of the discodrilid worms. Proc. U. S. Nat. Mus. 48(2071):
187-193.
Hosss, Horton H., Jr., Penny C. HOLT AND MARGARET WALTON. 1967.
The crayfishes and their epizoGtic associates of the Mountain
Lake, Virginia, region. Proc. U. S. Nat. Mus. 123(3602):1-84.
HoFFMAN, RicHarp L. 1963. A revision of the North American annelid
worms of the genus Cambarincola (Oligochaeta: Branchiobdel-
lidae). Proc. U. S. Nat. Mus. 114(3470) :271-371.
Hott, Perry C. 1949. A comparative study of the reproductive sys-
tems of Xironogiton instabilius instabilius (Moore) and Cam-
barincola philadelphica (Leidy). (Annelida, Oligochaeta,
Branchiobdellidae). J. Morph. 84(3) :535-572.
——. 1960. The genus Ceratodrilus Hall ( Branchiobdellidae, Oligo-
chaeta ), with the description of a new species. Virginia J. Sci.,
ON:S: )) 11(2)2538=77,
—. 1965. The systematic position of the Branchiobdellidae (An-
nelida: Clitellata). Syst. Zool. 14(1):25-32.
———. 1967. The status of the genera Branchiobdella and Steph-
anodrilus in North America with description of a new genus.
Proc. U.S. Nat. Mus. 124( 3631 ):1-—10.
—. 1968a. The Branchiobdellida: epizodtic annelids. The Biol-
ogist 50( 2-3 ):79-94.
———. 1968b. New genera and species of branchiobdellid worms
(Annelida: Clitellata). Proc. Biol. Soc. Wash. 81:291-318.
—. 1969. The relationships of the branchiobdellid fauna of the
southern Appalachians. in Holt, Perry C. (ed.). The distri-
butional history of the biota of the southern Appalachians.
Part I: Invertebrates. Res. Div. Monogr. 1, Virginia Poly-
technic Institute: 191-219.
An Emendation of Triannulata 71
—. 1973a. Epigean branchiobdellids (Annelida: Clitellata) from
Florida. Proc. Biol. Soc. Wash. 86(7):79-104.
—. 1973b. A summary of the branchiobdellid (Annelida: Clitel-
lata) fauna of Mesoamerica. Smithsonian Contr. Zool., 142:
i-iii, 1-40.
—, anpD Ricwarp L. Horrman. 1959. An emended description
of Cambarincola macrodonta Ellis, with remarks on the diag-
nostic characters of the genus (Oligochaeta: Branchiobdellidae ).
J. Tennessee Acad. Sci. 34(2):97—104.
Leiwy, JosEpH. 1851. Contributions to helminthology. Proc. Acad.
Nat. Sci. Phila. 5:205—209.
Lianc YAN-LIN. 1963. Studies on the aquatic Oligochaeta of China.
I. Descriptions of new naids and branchiobdellids. Acta
Zoologica Sinica 15( 4) :560-570.
Moore, J. Percy. 1894. On some leech-like parasites of American
crayfishes. Proc. Acad. Nat. Sci. Phila., (for 1893) :419-428.
1895. The anatomy of Bdellodrilus illuminatus, an American
discodrilid. J. Morph. 10(2):497-540.
Opirer, AucusTE. 1823. Memoire sur le branchiobdelle, nouveau
genre d’Annelides de la famille de Hirudinees. Mém. Soc. Hist.
Nat. Paris 1:71-78.
PENNAK, RoBERT W. 1953. Fresh-water Invertebrates of the United
States. The Ronald Press, New York, 769 pp.
PIERANTONI, UMBERTO. 1905. Cirrodrilus cirratus, n.g. n.s.p. Parasita
dell’ Astacus japonicus. Ann. Mus. Zool. R. Univ. Napoli,
(N.S.) 1(31):1-3.
1906. 2 nuovi discodrilidi del Giappone e della California.
Ann. Mus. R. Zool. Univ. Napoli. (N.S.) 2:11.
1912. Monografia dei Discodrillidae. Ann. Mus. Zool. R.
Univ. Napoli, (N.S.) 3(24):1-28.
STEPHENSON, J. 1930. The Oligochaeta. Oxford Univ. Press, Oxford,
978 pp.
YamacucHi, Hipeyi. 1932. A new species of Cambarincola, with re-
marks on spermatic vesicles of some branchiobdellid worms.
Proc. Imp. Acad. [Japan] 8(9):454—456.
—. 1934. Studies on Japanese Branchiobdellidae with some re-
visions on the classification. Journ. Fac. Sci., Hokkaido Imp.
Univ. ser. VI. Zool. 3(3):177-219, pls. 12, 13.
72 Proceedings of the Biological Society of Washington
Vol. 87, No. 9, pp. 73-76 25 April 1974
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
EUXESTA MAZORCA, NEW SPECIES, ASSOCIATED
WITH EARS OF MAIZE IN SOUTH AMERICA
(DIPTERA, OTITIDAE)
By Georce C. STEYSKAL
Systematic Entomology Laboratory, Agr. Res. Serv.,
U.S. Department of Agriculture, Washington, D. C1
Specimens of a species of Euxesta have been received on a
few occasions from Colombia, Ecuador, and Peru accompanied
with the notation that they were in some manner associated
with ears of corn (maize, Zea mays Linnaeus), although exact
data concerning their relations with the plant are not avail-
able. A few specimens of the species have been in the
United States National Museum collection for many years
determined as Euxesta obliquestriata Hendel, which was de-
scribed from Brazil, Paraguay, and Bolivia. The species herein
described, however, differs in several ways from Hendel’s
description and figure.
Euxesta mazorea, new species
Figures 1-2
Male and female. Length of wing 4-O—4.83 mm. Agrees with E.
obliquestriata Hendel in basic pattern of wing, interantennal space of
same color as lower parts of face (not dull black), wholly dark metallic
blue-green abdomen, and largely black legs.
Head as in Fig. 2; largely yellowish, metallic greenish on frontal
triangle, parafrontal plates, vertex, and occiput; middle of face and sides
of clypeus infuscated to variable extent; antenna usually wholly orange,
but 3rd segment sometimes a little infuscated above; front largely
almost polished, narrow orbits silvery tomentose.
Thorax wholly metallic greenish, mesoscutum lightly pale grayish
tomentose with indistinct more blackish arcuate postsutural transverse
band, especially evident above wing bases; scutellum blackish. Angle
1 Mail address: c/o U. S. National Museum, Washington, D. C. 20560.
9—Proc. Biot. Soc. WaAsH., VoL. 87, 1974 (73)
74 Proceedings of the Biological Society of Washington
Fics. 1-2, Euxesta mazorca, new species: 1. wing; 2. lateral view,
head of holotype, with anterodorsal view of front.
between line connecting posterior dorsocentrals bristles and each acro-
stichal bristle 45°.
Legs largely metallic greenish; forecoxa (except basilateral oval dark
spot), most of inner face of forefemur, fore- and midbasitarsi, hind-
basitarsus (except more or less of tip), and distal end of forefemur tawny.
Wing as in Fig. 1; medial hyaline band attaining costa in nearly full
width, impinging on only small apical part of Ist vein (R:); preapical
hyaline band somewhat narrower than apical brown band; section of
Ath vein crossing preapical hyaline band about equal in length to apical
section in brown area; length of apical brown area in discal cell less
A New Dipteran on Maize in South America 75
than half as great on 5th vein as on 4th vein; brown area in base of 2nd
costal cell squarish, not extended along costa; extension of tip of anal
cell about as long as greatest width of cell.
Abdomen wholly metallic bluish green, very lightly pale tomentose.
Holotype @, allotype, and 1 @ and 3 ¢@ paratypes, ECUADOR:
Quito, Estacién Experimental Santa Cataline, August, 1966, feeding on
corn ear (V. Vazquez), No. 72830 in United States National Museum.
Also examined: ECUADOR: 1 9, 1 sex P (head and abdomen missing ),
Azogues; 3 2, 1 6, Paute [Canar Province, near Azogues], 18 June
1959, in corn buds (G. Merino); COLOMBIA: 12, 3 4, Obonuco, 16
January 1973, mazorca choclo [= green ear of corn] (B. Yanguatin);
1 ¢, Sonson, Antioquia, 16 November 1955, 2500 meters, Solanum
andigenum (L. Posada); PERU: 1 sex ? (abdomen missing), Ajipampa,
Cajamarca, EF] Chota mountain, 1800 meters, 29 January 1968, trap with
hydrolyzed protein (B. G. Casanova C.); 1 9, Huadquifia, Cuzco [at
base of Cerro Macchupicchu], 5000 meters, 30 July 1911. All specimens
are in the United States National Museum collection.
The species-name is an American Spanish word for “ear of maize,”
treated as Latin and used as a noun in apposition.
76 Proceedings of the Biological Society of Washington
Vol. 87, No. 10, pp. 77-80 25 April 1974
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
A NEW SPECIES OF TRICHODECTES (MALLOPHAGA:
TRICHODECTIDAE) FROM THE YELLOW-THROATED
MARTEN (MARTES FLAVIGULA)
By K. C. Emerson AND RocER D. PRICE
2704 North Kensington Street, Arlington, Virginia 22207,
and Department of Entomology, Fisheries, and Wildlife,
University of Minnesota, St. Paul, Minnesota 55101
Dr. Howard B. Emery and his colleagues of the Arun Valley
Wildlife Expedition recently collected a series of Mallophaga
in Nepal on the Yellow-throated Marten. We believe that these
lice represent an undescribed species and herewith describe
and illustrate it.
Trichodectes emeryi, new species
Figures 1-5
Holotype male: Total length, 1.28 mm. External morphology and
chaetotaxy as shown in Figure 1. Anal region as shown in Figure 4.
Genitalia as in Figure 3; genital sac large, with prominent spines;
parameres unmatched in size and form, each curved outwardly.
Allotype: Total length, 1.64 mm. Female paratypes varying in length
from 1.41 to 1.64 mm, reflecting differences in mounting. External
morphology and chaetotaxy as shown in Figure 2, with ventral terminalia
as in Figure 5.
Discussion: Trichodectes emeryi does not cloesly resemble any known
species, although it does have a number of characters in common with
certain other species. The head, in both sexes, resembles in shape that
of T. octomaculatus Paine, 1912, found on the Raccoon, and T. pinguis
Burmeister, 1838, found on the European Brown Bear. The chaetotaxy
of T. emeryi consists of smaller setae on the dorsum and lateral margins
of the head than T. pinguis, and the forehead is somewhat more rounded
than that of T. octomaculatus. Setae on the posterior margin of the
pterothorax are more prominent and numerous than T. pinguis and T.
octomaculatus.
Dorsal abdominal chaetotaxy is of the type found on T. octomaculatus
and T. fallax Werneck, 1948, taken from the Crab-eating Raccoon; how-
10—Proc. Brot. Soc. WasH., Vou. 87, 1974 (CT)
78 Proceedings of the Biological Society of Washington
Fics. 1-5. Trichodectes emeryi, new species. 1, dorsal-ventral view of
male. 2, dorsal-ventral view of female. 3, male genitalia. 4, dorsal
terminalia of male. 5, ventral terminalia of female.
ever, both sexes of T. emeryi usually have three prominent setae (less
often two or four) on the posterolateral margins of segments II-VI,
instead of two, and the female lacks any medial tergal setae on III-
VIII, instead of having the single seta on each side as the other two
species or the male of T. emeryi. Ventral abdominal chaetotaxy of
New Louse from Yellow-throated Marten fig
segments II-VI is heavier and more numerous than that of T. octo-
maculatus and T. fallax.
The female genital region is distinctive, as shown in Figure 5. Male
genitalia have uniquely structured parameres and sac. T. zorillae Stobbe,
1913, found on the Libyan Striped Weasel, is the only previously known
species with dissimilar parameres and an armed genital sac. However,
the parameres of T. emeryi are short, stout, and curved outwardly, whereas
the parameres of T. zorillae are long, slender, and curved inwardly at
the distal tip.
Type-host: Martes flavigula (Boddaert, 1785).
Type-material: Holotype male, allotype, and 22 paratypes collected
off the type-host at Sankhuwa Sabha, Nepal, on 8 February 1973. The
holotype and allotype will be deposited in the National Museum of
Natural History, Smithsonian Institution.
80 Proceedings of the Biological Society of Washington
Vol. 87, No. 11, pp. 81-90 25 April 1974
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
VANZOLINIUS, A NEW GENUS PROPOSED FOR
LEPTODACTYLUS DISCODACTYLUS
(AMPHIBIA, LEPTODACTYLIDAE )
By W. RonaLp HEYER
Division of Amphibians and Reptiles, National Museum
of Natural History, Smithsonian Institution,
Washington, D.C. 20560
The relationships of Leptodactylus discodactylus Boulenger
have been obscure. In the first analysis of the relationships of
discodactylus since Boulenger’s description, I (1970) placed
this species in the Melanonotus group of the genus Lepto-
dactylus, based primarily on external morphology. This re-
lationship was questioned when certain life history data were
gathered (Heyer and Bellin, 1973). In a recent analysis of
the relationships of the genus Adenomera within the subfamily
Leptodactylinae (Heyer, 1974), I concluded that disco-
dactylus was most closely related to Lithodytes lineatus, and
with a certain amount of hesitation redefined Lithodytes to
include discodactylus. Since then, the karyotype has been
determined for discodactylus (Heyer and Diment, in prep.)
and I have been able to examine more material of Lithodytes
lineatus, which has resulted in a change of one character state.
With this additional information, I believe that there is now
sufficient evidence to firmly establish the generic identity of
discodactylus.
Previously (Heyer, 1974), primitive and derived states
were defined for 50 taxonomic characters, and polarities of
character states were inferred for members of the subfamily
Leptodactylinae. Alternate phylogenies were constructed based
on shared derived character states. One phylogeny was chosen
that was thought to best represent the actual relationships
11—Proc. Brot. Soc. WasH., Vou. 87, 1974 (81)
82 Proceedings of the Biological Society of Washington
within the subfamily. The analysis showed that Adenomera,
Leptodactylus, Lithodytes lineatus, and discodactylus con-
stitute a tight taxonomic cluster. It is the relationships among
these taxa that need to be reevaluated; this study, therefore,
is limited to examining the relationships among these four taxa.
MertTHOops AND MATERIALS
The same methods and data are used as in Heyer (1974)
with the following modifications: 1) If all four taxa share
the same character state, that character is not used; 2) If a
derived state is unique to one taxon, it is not used as it gives
no information on common ancestry; 3) The units of com-
parison are different, necessitating a redescription of the
character states (see below). In the previous study, the unit
used in the analysis was the species. In this study, as generic
relationships are the focus, the unit of comparison is the genus;
4) Four characters (pterygoid-parasphenoid overlap; iliacus
externus muscle; adductor longus muscle; gluteus muscle),
were shown to be suspect in the original determination of
direction of change of states (Heyer, 1974), and as no addi-
tional evidence has been accumulated to resolve these char-
acters, they are omitted in the present study.
Character state descriptions: External vocal sacs—Primitive
state: No external vocal sac. State 1: No external vocal sac
or indications of lateral vocal folds. State 2: No external vocal
sac, indications of vocal folds, or well-developed lateral vocal
sacs. The direction of change of character states is: P>1—2.
Male thumb—Primitive state: Nuptial adspersities present
in form of pad. State 3: No nuptial adspersities. State 4:
Either spines on thumb, or in case of Fuscus group, no
adspersities (see Heyer, 1974). The directions of change of
character states are: 3<-P—4.
Body glands—Primitive state: No well-defined glands. State
5: No glands and/or dorsolateral folds.
Toe disks—The dorsal toe disks of Lithodytes lineatus
have dorsal scutes, similar to those of Hylodes and distinct
from the longitudinal grooves in the disks of discodactylus.
Primitive state: No disks. State 6: No disks and/or disks
lacking dorsal grooves or scutes. State 7: Toe disks with
A New Genus of Leptodactylid Frog 83
scutes on dorsal surface. State 8: Toe disks with longi-
tudinal grooves on dorsal surface. The directions of change
of character states are: P—6-—>7.
\
8
Toe web—Primitive state: Toes fringed. State 9: Toes
fringed or free. State 10: Toes free. The direction of change
of character states is: P—9—10.
Egg pigment—Primitive state: Melanocytes present or
absent. State 11: No melanocytes.
Geniohyoideus lateralis muscle—Primitive state: No lateral
flare or slip. State 12: Lateral flare or slip present.
Sternohyoideus muscle origin—Primitive state: Single or
double slip from sternum. State 13: Double slip from sternum.
Sternohyoideus muscle insertion—Primitive state: Lateral
edge of hyoid plate. State 14: Some fibers near midline of
hyoid plate.
Gracilis minor—Primitive state: Broad. State 15: Rudi-
mentary.
Frontoparietal fontanelle—Primitive state: None. State 16:
Small fontanelle present or absent.
Anterior articulation of vomer—Primitive state: Vomer
articulating with maxilla or premaxilla or neither. State 17:
Vomer articulating with maxilla or premaxilla.
Sphenethmoid-optic foramen relationship—Primitive state:
Posterior extent of sphenethmoid far from optic foramen. State
18: Posterior extent of sphenethmoid far from to bordering
optic foramen.
Anterior extent of sphenethmoid—Primitive state: To mid-
vomer. State 19: To mid-vomer or more anteriad.
Terminal phalanges—Primitive state: Knobbed. State 20:
T-shaped.
Karyotypes—Heyer and Diment (in prep.) argued that two
aspects of karyotypes yield phylogenetic information on the
genera Adenomera and Leptodactylus: diploid number and
presence or absence of acrocentric chromosomes. The prim-
itive karyotype was argued to have a diploid number of 26
with acrocentric chromosomes.
Diploid number—Primitive state: 2N = 26 or 24. State 21:
84 Proceedings of the Biological Society of Washington
2N = 22. State 22: 2N = 18. The direction of change of states
is: P2122.
Acrocentric chromosomes—Primitive state: Present. State
93: Present or absent. State 24: Absent. The direction of
change of character states is: P—~23—24.
RELATIONSHIPS
The four taxa have the following advanced states: Adeno-
mera—l, 3, 6, 10, 11, 12, 14, 15, 16, 18, 19, 20; Leptodactylus—
2, 4, 5, 9, 12, 16, 18, 19, 21, 23; lineatus—3, 5, 7, 10, 11, 13, 14,
17, 20, 22, 24; discodactylus—3, 8, 11, 13, 15, 17, 20, 21.
Two taxa pairs share the most derived character states (7):
Adenomera-lineatus and lineatus—discodactylus. As Lepto-
dactylus and discodactylus only share one derived state, the
best phylogeny using the Adenomera-lineatus pair cluster is
that shown in Fig. 1, A. Two alternate phylogenies are possible
using the lineatus—discodactylus cluster (Fig. 1, B, C). Adeno-
mera shares 6 states with Leptodactylus (Fig. 1, C) and 5
states with discodactylus (Fig. 1, C, D). The phylogenies
figured (A-E) are all of the reasonable possibilities.
In evaluating which phylogenies are likely to give a truer
reflection of the actual relationships among these taxa, three
criteria may be applied to the phylogenies. The first criterion
was used in constructing the phylogenies—maximizing the
number of shared character states, or in Hennig’s terms, seek-
ing sister-groups. Using this criterion, the phylogenies of
Fig. 1, A, B, C are preferred over those of Fig. 1, C and D.
A second criterion is the number of convergences of character
states required by each phylogeny. The fewer the number,
the more likely that the phylogeny is correct. The number of
convergences in each of the phylogenies pictured are: Fig.
1, A and B, 13 convergences; Fig. 1, C, 11 convergences; Fig.
1, D and E, 15 convergences. A third criterion is to evaluate
the ancestral clusters in terms of non-convergent character
states. That is, if a character state is already convergent in
>
Fic. 1. Five possible phylogenies for Adenomera, Leptodactylus,
Lithodytes lineatus, and discodactylus.
A New Genus of Leptodactylid Frog
A
Adenomera Lineatus
LIZ AS MGSO Seo seiliaceacoe4,
9,10,14 discodactylus
8,13,15,17;21
3,6,11,20 Leptodactylus
132,4,539,12,16,18,19,21 ,23
B
lineatus discodactylus
5,7,.9,10,14,. 8,15
2252324
Adenomera
WSU 62 1,9,10,12,14,
; 15,16,18,19
3,6,11,20 Leptodactylus
ee Se errr nae
C
lineatus discodactylus Adenomera Leptodactylus
5,7,9,10,14 8,15 3,6,10,11 25455321,23
22523524 14,15,20
1,9,12,16,18,19
SLASUU SSS l/pAU all
D
Adenomera discodactylus
1,9,10,12,14,16,18,19 8,13,17,21
Ss
15 Lineatus
5,7;,9,10,13,14,17521 522,23,24
BAG5 U1 520) Leptodactylus
1,2,4,5,9,12,16,18,195,21 ,23
E
Adenomera discodactylus Leptodactylus lineatus
1,9,10,12,14, 8,13,17,321 152,4,12,16,18,19 SRON OSs sili.
16,18,19 17,20,22,24
3,6,11,155,20 559521523
86 Proceedings of the Biological Society of Washington
a given phylogeny, its phylogenetic information content is not
as great as non-convergent character states. A set of four
shared derived character states that are not convergent in
the phylogeny is more robust than a set of four derived states
that are convergent within the total phylogeny. The phylog-
enies of Fig. 1, A and B are strong, with six non-convergent
states ancestral to either Adenomera and lineatus (Fig. 1, A)
or lineatus and discodactylus (Fig. 1, B) and four non-
convergent states ancestral to Adenomera, lineatus, and
discodactylus. The phylogeny of Fig. 1, C is almost as strong
as Adenomera and Leptodactylus share five non-convergent
states; however, lineatus and discodactylus share but two non-
convergent states. The phylogeny of Fig. 1, D has the same
non-convergent states ancestral to Adenomera, discodactylus,
and lineatus that are found in the phylogenies of Fig. 1, A
and B. Within the cluster of these three taxa, the phylogeny
of Fig. 1, D is weakest, as Adenomera and discodactylus only
share one non-convergent state while each of these taxa shares
two non-convergent states with lineatus (Fig. 1, A, B). The
phylogeny of Fig. 1, E is weak, Adenomera and discodactylus
sharing only one non-convergent state and lineatus and Lepto-
dactylus sharing two non-convergent states.
The phylogenies of Fig. 1, A, B, and C are the most robust.
All three have the maximum number of shared derived states
for a species pair. The phylogenies of Fig. 1, A and B have the
most shared non-convergent ancestral states, while that of
Fig. 1, C has the least number of convergent states. The
phylogenies of Fig. 1, D and E are much weaker in comparison
and are not further considered.
It is necessary at this point to reiterate my concept of a
genus: it should 1) be monophyletic, 2) be reasonable in size
(number of species), and 3) represent a distinct adaptive com-
plex if possible. Criterion 2 does not apply here, as Adeno-
mera, Leptodactylus and Lithodytes combined have about 40
species. As developed elsewhere (Heyer, 1973, 1974), Adeno-
mera is a wet forest genus, Leptodactylus is a savanna genus
and each has evolved in response to the very different selective
pressures associated with the two environments. The resolu-
tion of how many genera are represented among Adenomera,
A New Genus of Leptodactylid Frog 87
Leptodactylus, discodactylus, and lineatus thus hinges on
adaptive complexes, which information is not complete for
discodactylus and lineatus. The available evidence is sufficient
to determine the broad adaptive relationships of lineatus and
discodactylus, however. No egg clutches have been found for
either form, so it is not known whether either has a foam nest.
In both, the eggs are non-pigmented (determined from ad-
vanced ovarian eggs), which strongly suggests that the eggs
are hidden in some fashion. In the case of lineatus, the eggs
are large and of the same size found in Eleutherodactylus,
which has direct development. The eggs are small in disco-
dactylus and are probably laid at the male calling sites, which
are at the edge of seepage areas in naturally occurring en-
closures formed by tree roots and leaves (Heyer and Bellin,
1973). The calling sites of lineatus are not known, but collec-
tions indicate that they are terrestrial and/or standing water
breeders. With what is known, discodactylus can be cate-
gorized as having adaptive complexes distinct from both
Adenomera and Leptodactylus. Briefly, discodactylus is
adapted to the slow moving stream way of life, Adenomera
is adapted to the terrestrial, wet-forest way of life, and Lepto-
dactylus is adapted to xeric environments.
Combining this information with the phylogenies of Fig.
1, A, B, C, the following nomenclatural decisions are possible.
If Adenomera, Leptodactylus, and discodactylus are ge-
nerically distinct, then lineatus would have to be a distinct
genus according to the phylogeny of Fig. 1, A, or it could be
combined with discodactylus in a common genus in the
phylogenies of Fig. 1, B and C. Lithodytes lineatus has a
number of derived states in addition to those shared with
discodactylus, however, including dorsolateral folds, free toes,
and a derived karyotype. These unshared derived states,
together with the probability that lineatus and discodactylus
differ in mode and habitat of egg and larval development
convince me that lineatus and discodactylus are generically
distinct.
A comment might be helpful in explaining why I think this
decision is the proper one to make. In the previous study
(Heyer, 1974), the evidence then available dictated that
88 Proceedings of the Biological Society of Washington
discodactylus was certainly not a Leptodactylus. To remove
discodactylus from the genus Leptodactylus, a decision had to
be made to include the species in the genus Lithodytes or to
name a new genus for it. I chose the former course because
it was nomenclaturally conservative and I hoped that more in-
formation on life history and karyotypes would resolve the
matter. Combining lineatus and discodactylus in a common
genus was believed to be an unsatisfactory solution, however,
as the two species appear very dissimilar. The karyotypic
evidence further supports the generic differentiation of these
two species. The relationships of discodactylus have been of
concern because externally discodactylus looks like members
of the Melanonotus group of the genus Leptodactylus, or if
the toe fringe were removed, like members of the genus
Adenomera. It is apparent that the similarities of discodactylus
to certain species of Leptodactylus and Adenomera are based
on shared primitive character states, however. The analysis of
derived shared character states clearly demonstrates the
divergent evolutionary pathways of these three taxonomic
units.
TAXONOMIC CONCLUSIONS
Four genera are recognized in the Leptodactylus-complex:
Adenomera, which contains five species, Leptodactylus, which
contains about 35 species, Lithodytes, which contains one
species, and a genus for discodactylus. Definitions of Adeno-
mera and Leptodactylus may be found in Heyer (1974);
a definition of Lithodytes may be found in Lynch (1971). As
no generic name has been proposed for discodactylus, a new
genus is described as follows:
Vanzolinius, new genus
Type species: Leptodactylus discodactylus Boulenger, 1883.
Diagnosis: Vanzolinius is unique among leptodactylid frogs in possess-
ing a bony mesosternum and expanded toe disks with longitudinal grooves
on the dorsal surfaces. All other genera with a bony mesosternum either
do not have toe disks or, if disks are present, either do not have any
dorsal modifications or have dermal scutes.
Definition: Pupil horizontal; tympanum distinct; vocal sac internal;
male thumb without nuptial adspersities; body without well defined
glands; toes disked with dorsal surfaces with 3-5 longitudinal grooves;
A New Genus of Leptodactylid Frog 89
tarsal fold present; metatarsal tubercles neither pronounced nor cornified;
toes with lateral fringes; eggs lacking melanophores; large clutch size
(> 1000 eggs); depressor mandibulae condition DFsq (large slip
originating from dorsal fascia, small slip originating from squamosal area);
geniohyoideus medialis continuous medially; geniohyoideus lateralis
without lateral flare or slip; anterior petrohyoideus insertion on edge
of hyoid; sternohyoideus origin with distinct slips from anterior meso-
sternum and another from posterior meso, and/or xiphisternum; sterno-
hyoideus insertion near lateral edge of hyoid; omohyoideus insertion on
hyoid plate and fascia between posterolateral and posteromedial processes
of hyoid; tendon of semitendinosus confluent with posterior portion
of sartorius insertion on knee and tendons of gracilis minor and major
passing dorsad to tendon of semitendinosus; iliacus externus extending
from % to full length of iliac bone; tensor fasciae latae insertion pos-
terior to iliacus externus on iliac bone; gracilis minor narrow; interior
and exterior portions of the semitendinosus uniting in common tendon
distally, exterior portion larger or equal to interior (smaller) portion;
sartorius moderately developed; accessory head of adductor magnus
without distinct tendon; adductor longus well developed; gluteus
insertion on cruralis and knee; quadratojugal well developed, con-
tacting maxilla; maxillary teeth present; nasals widely separated; no
frontoparietal fontanelle; zygomatic ramus of squamosal just longer
than, just shorter than, or equal to otic ramus; vomerine teeth present;
vomers not in medial contact; vomer articulation with premaxilla and/or
maxilla; posterior extent of sphenethmoid widely separated from optic
foramen; sphenethmoid extending anteriorly to middle of vomers;
occipital condyles widely separated; no anterior processes of hyale; alary
process of hyoid somewhat narrow and stalked; posterolateral process
of hyoid present; ilium with well developed dorsal crest; terminal
phalanges T-shaped, expanded; mesosternum a bony style; xiphisternum
entire, single; diploid chromosome number 22, one pair of acrocentric
chromosomes.
Etymology: The genus, masculine in gender, is named for Dr. Paulo
E. Vanzolini, in recognition of his work on the South American herpeto-
fauna.
Content: Monotypic. For further details of morphology and dis-
tribution, see Heyer (1970).
ACKNOWLEDGMENT
George R. Zug, National Museum of Natural History, Smithsonian
Institution, critically read the manuscript.
LITERATURE CITED
Boutencer, G. A. 1883. On a collection of frogs from Yurimaguas,
Huallaga River, northern Peru. Proc. Zool. Soc. London 1883:
635-638.
90 Proceedings of the Biological Society of Washington
Heyer, W. R. 1970. Studies on frogs of the genus Leptodactylus
LYNCH,
(Amphibia, Leptodactylidae). VI. Biosystematics of the
Melanonotus group. Contrib. Sci., Los Angeles County Mus.
Natur. Hist. 191:1—48.
1973. Systematics of the Marmoratus group of the frog
genus Leptodactylus (Amphibia, Leptodactylidae). Contrib.
Sci., Los Angeles County Mus. Natur. Hist. 251:1-50.
1974. Relationships of the Marmoratus species group (Am-
phibia, Leptodactylidae) within the Subfamily Leptodactylinae.
Contrib. Sci., Los Angeles County Mus. Natur. Hist. 253:1—46.
AND M. S. Betuin. 1973. Ecological notes on five sympatric
Leptodactylus (Amphibia, Leptodactylidae) from Ecuador.
Herpetologica 29:66-72.
J. D. 1971. Evolutionary relationships, osteology, and zoo-
geography of leptodactyloid frogs. Univ. Kansas Mus. Natur.
Hist. Misc. Publ. 53:1-238.
wna
Vol. 87, No. 12, pp. 91-94 25 April 1974
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
ADDITIONAL NOTES ON THE GENUS GLARESIS
(COLEOPTERA: SCARABAEIDAE)
By Rosert D. GorDON
Systematic Entomology Laboratory,
Agricultural Research Service, USDA‘
Since the genus Glaresis was revised for the U. S. and
Canada (Gordon, 1970), specimens have been examined that
extend the known ranges of some species. The most interesting
record is of a specimen of G. ecostata Fall collected by H.
Howden at Angostura Dam, Hot Springs, South Dakota in
July 1968, along with specimens of G. canadensis Brown and
G. inducta Horn. The latter two species would be expected at
that locality, but ecostata had previously been recorded only
from California, Arizona and New Mexico. I collected another
specimen of ecostata at Angostura Dam in August 1973. Speci-
mens of G. phoenicis Fall have been examined from Mesquite,
Nevada, and White Sands, New Mexico. G. confusa Brown
was previously known only from the unique type (Gordon,
1970), but specimens from two additional localities have since
been observed. The data on these specimens is as follows:
Bull Creek, 5000’, 15 mi. S Hanksville, Utah, VII-6-1968, A. T.
Howden; Cochise Co., Arizona, 5 mi. N Benson, VII-26-1969,
A. R. Hardy, F. G. Andrews, J. W. Smith. Of 15 Glaresis
specimens from Benton, Mono Co., California, sent to me by
Alan Hardy, 13 proved to be G. medialis Gordon, a species
previously known only from Utah (Logan and Richfield)
and Washington (Toppenish). The other two specimens are
G. clypeata Van Dyke and G. mendica Horn. Hardy also sent
four Glaresis specimens from another locality, these being the
type material of the new species described below.
1 Mail address: c/o U. S. National Museum, Washington, D. C. 20560.
12—Proc. Brot. Soc. WaAsH., Vou. 87, 1974 (91)
92 Proceedings of the Biological Society of Washington
Notes on Glaresis 93
(f\
6
Fic. 6. Glaresis arenata, new species (holotype), ventral view of
male genitalia.
The antenna of species of Glaresis has been recorded in the
literature as having nine segments. In my 1970 paper I
accepted this without examining the antenna myself. I am
indebted to Paul Ritcher, Oregon State University, for calling
my attention to the fact that Glaresis has ten-segmented
antennae. I have personally confirmed this by examining the
antenna of five species of Glaresis.
In addition to Dr. Ritcher, I wish to thank Alan Hardy,
California Department of Agriculture, for the loan of speci-
mens; Henry Howden, Carleton University, Ottawa, and E. C.
Becker, CNC, Ottawa, for data obtained from specimens in
their care.
The Scanning Electron Microscope time for this paper was
supported in part by the University of Maryland Center of
Material Research, Department of Mechanical Engineering
and Electron Microscope Central Facility, College Park,
Maryland.
Glaresis arenata, new species
Male: Length 3.80 mm, greatest width 2.05 mm. Form elongate,
widest posterior to middle of elytra. Color pale brownish yellow, head
and pronotum slightly darker than elytron, reddish brown. Head smooth,
extremely feebly tuberculate; anterior clypeal margin straight, distinctly
<
Fics. 1-5, Glaresis arenata, new species (paratype): 1, head; 2, apex
of mesotibia; 3, teeth on posterosuperior margins of hind trochanter and
femora; 4, enlarged view of teeth on posterosuperior margin of hind
femora; 5, hind tibia.
94 Proceedings of the Biological Society of Washington
margined with traces of tubercles, anterior angle angulate, not toothed
(Fig. 1); mandible slender, sinuate. Pronotum slightly more than half as
long as wide; short, longitudinal carinae present except in impressions;
surface nearly smooth, impressions feeble; lateral and posterior margins
with fringe of coarse setae; a distinct fovea medially near lateral margin
and a feeble, longitudinal, median impression present. Elytron smooth;
interval nearly without costa, flat with tiny, widely interrupted remains
of a costa present; sparsely pubescent. Mesotibia sinuate on outer
margin, five spines present on outer margin from middle nearly to apex,
projecting at apex (Fig. 2). Posterosuperior margin of hind trochanter
with a single, large tooth near femur (Fig. 3); posterosuperior margin of
hind femur with two small teeth (Figs. 3, 4). Hind tibia broadly tri-
angular, a strong projection on outer margin medially, elongate tubercles
irregularly scattered over surface, inner margin strongly pubescent (Fig.
5). Genitalia with apex of aedeagus rounded in ventral view (Fig. 6),
curved upward in lateral view.
Female: Length 3.56 mm, greatest width 2.00 mm. Similar to male
in external appearance.
Type-material: Holotype, male, sand dunes, 7 mi. SW Kelso, San
Bernardino Co., California, IV-20-21-1969, fluorescent black light, M. S.
Wasbauer and R. R. Pinger (USNM). Paratypes, three, same data as for
holotype except one collected IV-16-17-1969 and one collected beating
Larrea divaricata. Paratypes in collections of USNM, California Acad-
emy of Sciences and the collection of Alan Hardy, Sacramento, Calif.
Variation: Length 3.25 to 3.80 mm, greatest width 1.78 to 2.05 mm.
The number of spines on the outer margin of the mesotibia varies from
four to five, and one specimen of the type series had a feeble second tooth
present on the posterosuperior margin of the trochanter.
Remarks: This species goes to couplet ten in the key to species
(Gordon, 1970) and would key to cartwrighti Gordon. G. arenata
differs from cartwrighti in having the apex of the aedeagus broadly
rounded, the elytral intervals not distinctly costate, the teeth on postero-
superior margin of hind femur widely separated, the outer margin of
hind tibia strongly projecting medially and the outer margin of middle
tibia with spines present, distinctly projecting apically. The lack of dis-
tinct elytral costae is unusual, and in this respect arenata resembles
medialis Gordon and, to a lesser degree, ecostata Fall, but these two
species do not resemble arenata in any other characteristics.
LITERATURE CITED
Gorpon, R. D. 1970. <A review of the genus Glaresis in the United
States and Canada (Coleoptera: Scarabaeidae). Trans. Amer.
Entomol. Soc. 96:499-517.
Vol. 87, No. 13, pp. 95-128 25 April 1974
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
A REVISION OF THE GENUS BOCOA
(CAESALPINIOIDEAE-SWARTZIEAE )
By RicHarp S. CowANn
Smithsonian Institution, Washington, D.C. 20560
INTRODUCTION
Aublet (1775) described the genus Bocoa, with a single
species, B. proudcensis, in a supplemental section of his classic
work on the plants of French Guiana. When Schreber (1789-
1791) and Willdenow (1800) considered the species of what
now is Swartzia, no mention was made of Bocoa, and, except
for the confusion with Inocarpus and Etaballia discussed
below, the genus was ignored thereafter. In fact, taxa con-
generic with B. prouacensis were described subsequently as
species of Swartzia by Bentham (1840) and others. Tulasne
(1843) even redescribed the genus under a new name,
Trischidium, based on a species published by Bentham three
years earlier. It remained for Amshoff (1939) to recognize
the affinities of Bocoa, although she considered it a synonym
of Swartzia. My earlier study of this subtribe convinced me
of the distinctness of the two genera and in my monograph of
the species of Swartzia (Cowan, 1968), I excluded several as
taxa belonging to Bocoa; the new combinations were not made,
pending further study of specific characteristics.
If one consults Index Kewensis, Bocoa will be found listed
as a synonym of Inocarpus Forster, an Old World genus of
which one species is widely cultivated on islands of the Pacific
Basin. Another synonym that appears under this genus is
Etaballia Bentham. Although the three genera were main-
tained by Bentham as separate taxa in the beginning, he was
less than convinced in his 1862 paper on the subject and later
in the Genera Plantarum he synonymized his genus under
13—Proc. Bron. Soc. WAsH., Vou. 87, 1974 (95)
96 Proceedings of the Biological Society of Washington
Inocarpus. Comparison of herbarium material of the species of
Etaballia and Inocarpus does raise considerable doubt con-
cerning their separateness and, in fact, they have been con-
sidered to be congeneric by some later workers. Most re-
cently, Rudd (1970) concluded they should be treated as
separate genera, pending further detailed studies.
Whether or not two genera are involved, the species of
Etaballia and Inocarpus are clearly papilionate legumes and
have little similarity to the caesalpinioid genus Bocoa, except
for the unifoliolate leaves of many of the taxa in question. The
flowers of B. prouacensis, the only species Bentham accepted
in the genus, are quite distinctive in being apetalous, other
species of the genus having a single obovate petal; in con-
trast, flowers of Inocarpus and Etaballia have five, strap-like
petals and a very different androecium. The latter genera
have indehiscent fruits while those of Bocoa are dehiscent
in two valves along both sutures.
The characteristics separating Bocoa and Swartzia can be
set out clearly in a key:
Anthers oval or oblate to oblong, dorsifixed; stamens usually
many and dimorphic, the filaments several times the length of
the anthers; seeds arillate; herbage not foetid; leaflets rarely
alternate 200 oy Swartzia
Anthers linear to narrowly oblong, basifixed; stamens 7-30, the
filaments and anthers of similar length; seeds usually exarillate;
herbage often foetid in fresh state; leaflets often alternate to
subopposite =... 20. DS eee Bocoa
There is greater similarity morphologically between these
genera than there is between Swartzia and other genera of
this complx. One is tempted to think of Bocoa as a phylo-
genetic derivative of Swartzia, derived from such species as
S. arborescens (Aublet) Pittier and S. auriculata Poeppig,
both of which have monomorphic stamens. However, I doubt
that the situation is anything so simple and I have no “hard”
evidence on which even to speculate profitably with respect to
the true relationship of these genera. Similarly, the order of
the species in the following treatment is largely arbitrary, but
generally, evolutionary loss of flower parts is viewed as derived
and connation of parts more advanced than totally free mem-
bers of a series.
A Revision of the Genus Bocoa O77
POLLEN MORPHOLOGY
In my monograph of Swartzia, I described pollen shapes
(Cowan, 1968:6) as “globose .. . to oval, elliptic, deltoid-
globose, or deltoid.” These forms were from observations of
pollen in lacto-phenol aniline-blue on slides prepared by an in-
experienced technician (the author). I am now convinced
that deltoid-globose or deltoid grains do not occur in the
genus or its near-relatives. This conclusion is based on study
of the pollen of fourteen species of Swartzia, representing all
the major subgeneric groupings, four species of Bocoa, two of
Aldina, and one each of Zollernia, Lecointea, Candolleo-
dendron, and Holocalyx. I am greatly indebted to Joan
Nowicke, Julianne Piraino, and Dieter Wasshausen for the
preparation of the pollen samples for study by both scanning
electron and light microscopy, as well as for their consultative
assistance in my analysis of the data.
I undertook this preliminary review of the pollen of the
Swartzieae because of my long held suspicion that this collec-
tion of genera is an unnatural assemblage which has two
characters in common but whose members are not otherwise
closely related—an entire calyx and an indefinite number of
stamens. It seemed possible that palynology might be help-
ful in sorting out the several taxa. However, this has not
proven to be the case and this fact, in addition to greater
familiarity with the constituent genera, leads me to believe
now that there may indeed be considerable evolutionary
relationship among them. Certainly pollen morphology does
not refute this viewpoint, for the similarities palynologically
are great.
To document this conclusion and to provide a brief summary
of the diversity observed among the pollens of the limited
sampling of species enumerated below, the following account
and illustrations are offered. It is not intended to be a com-
prehensive, analytical description of pollen morphlogy in
Bocoa or of any of its relatives. Rather, it is hoped that this
presentation, however incomplete, will stimulate further, more
detailed palynological investigations of the caesalpinioid
legumes.
98 Proceedings of the Biological Society of Washington
All species examined had pollen grains which were: monads
with the longest axis ca. 18-30 wm, spheroidal to subprolate
or prolate, 3-colporate, the colpi short, elongate, or syncolpate,
the os mostly circular and frequently prominent, sometimes
bridged by the ektexine, ektexine more or less psilate, punctate,
finely or coarsely rugose, or striato-reticulate.
Based on a combination of colpus length and ektexine
pattern, five pollen types can be recognized. Unfortunately,
these groupings, appear to correlate with nothing else and
certainly provide no new insights into the phylogeny of the
Swartzieae. Also, it should be emphasized that these pollen
types are not well separated and that palynological investi-
gation of additional species of either Bocoa or Swartzia may
reduce the already limited distinctions among these groupings:
Group I: Colpi short or elongate but not syncolpate.
Type I-A: Ektexine rugose.
Type I-B: Ektexine striato-reticulate, sometimes very
finely so.
Type I-C: Ektexine punctate.
Group II: Colpi syncolpate.
Type III-A: Ektexine rugose.
Type II-B: Ektexine punctate.
Following is a listing of the species examined in each pollen-
type, the number of the illustration in parentheses just after
each name, any special notes about the morphology, citation
of the voucher collection, and the source herbarium for the
sheet sampled:
Comments Voucher
_Species of Type I-A:
Bocoa alterna (Fig. 1) Ducke 24203 (US)
B. mollis (Fig. 2) Blanchet 2774 (G)
B. prouacensis Stahel 69 (G)
B. racemulosa (Fig. 3) Snethlage 10058 (G)
Swartzia apetala Thick-walled grains Santos et al. 24166 (US)
S. aptera Many grains abnormal Maguire et al. 47056 (US)
S. brachyrachis Some grains syncolpate, Ducke 17038 (US)
others not
S. flaemingii (Fig. 4) Finely rugose Froes 11753 (US)
S. guianensis (Fig. 5) Very coarsely rugose Boyan 67 (US)
and thick-walled
A Revision of the Genus Bocoa 99
Comments Voucher
Species of Type I-B:
S. amplifolia Very finely striato- Klug 3122 (US)
reticulate
S. panacoco (Fig. 6) Finely reticulate Cardona 1142 (US)
S. cuspidata (Fig. 7) Os prominent Maguire et al. 60409 (US)
S. jorori (Fig. 8) Cardenas 5211 (US)
S. arborescens (Fig. 9) Very finely reticulate, Lizots.n. (US)
appearing psilate by
light microscopy
S. leptopetala Finely striato- Wurdack et al. 41367
reticulate (US)
Aldina latifolia (Fig. 10) Maguire/Wurdack 35595
(US)
A. macrophylla Wurdack/ Adderley 43431
(US)
Zollernia vogelii (Fig. 11) Handro 45344 (US)
Cyathostegia matthewsti (Fig. 12) Ferreyra 89lla (US)
Species of Type II-A:
Swartzia benthamiana (Fig. 13) Thick-walled with Irwin 57641 (US)
os prominent
Species of Type II-B:
Swartzia panamensis (Fig. 14) Punctate Standley 55317 (US)
S. grandifolia Coarsely punctate Holt/Blake 673 (US)
It is interesting to note that all the species of Bocoa have the
same pollen-type, in common with several Swartzia species
representing disparate phylogenetic groupings into which the
genus is divided. Also the species of Aldina, Cyathostegia,
and Zollernia have similar pollen morphology which appar-
ently is slightly different from that of all other species ex-
amined. (Cyathostegia is now considered a separate genus but
was earlier viewed as a part of Swartzia.)
The pollens of the three other genera in this complex do
not fit into the “classification” above. The pollens of Lecointea
(Ducke 1702-US) and Holocalyx (Pedersen 1422a-US) are
more similar to each other than to those of any of the other
taxa studied: under light microscopy the grains appear thin
walled, more or less psilate, 3-colporate, the colpi not con-
tinuous at the poles. On the other hand, the pollen of Can-
dolleodendron (Froes 32364-US) is intermediate in several
respects, just as is the gross morphology of other parts of the
plant: the grains are prolate and rugose but some are syn-
colpate while others (fewer) are not.
100 Proceedings of the Biological Society of Washington
The following figures are arranged by pollen-type to
facilitate review of the observations made. The arrangement
represents, then, an attempt to organize the data for report-
ing purposes, rather than to suggest phylogenetic relationships
of the taxa in each grouping or between groupings.
SYSTEMATIC TREATMENT
Bocoa Aublet
Bocoa Aublet, Pl. Guiane Fr. Suppl. 38, t. 391. Jun. 1775.
Trischidium Tulasne, Ann. Sci. Nat. (Ser. 2) 20:141, t. 4. Sep. 1843.
Shrub or small to large tree up to 30 m tall and 25 cm in diameter,
the wood and leaves foetid, the branchlets glabrous or strigulose to
tomentulose and glabrescent, the stipules minute and more or less tri-
angular or lanceolate and up to 5 mm long; leaves 1- to 9-foliolate,
imparipinnate, the petioles and rachis terete or occasionally flat or
canaliculate on the upper surface, leaflets alternate to opposite, the
blades glabrous to tomentulose, oval, elliptic, rounded, sometimes ovate-
elliptic or lanceolate-elliptic, the base rounded to acute, the apex acute to
acuminate or obtuse and retuse to emarginate; inflorescences racemose,
infrequently compound racemes, ramuligerous just below the leaves
of the current season, or occasionally axillary, sometimes very short but
up to 10 cm long, the axis glabrous, puberulous, strigulose, or tomentulose,
the pedicel scars strongly raised, the bracts caducous to persistent, tri-
angular to semicircular, about 1 mm long and wide; bracteoles absent;
pedicels 0.5-8 mm long at anthesis, becoming nearly twice as long with
mature fruits, pubescent like the inflorescence axis; buds ellipsoid to
oblong-ellipsoid, more or less pubescent externally, glabrous within,
opening into 3 or 4 segments, free or remaining united in basal half,
reflexed to strongly revolute after anthesis; petal present or lacking,
white and usually glabrous, caducous to persistent, the claw 2-6 mm
long, the blade round to oblate, 4-8 mm long and 3-7 mm wide;
stamens 7-30, uniform, the filaments glabrous, sometimes shortly united
basally, 2-6 mm long, the anthers oblong, basifixed, 1.5-4 mm long,
0.5-0.9 mm wide, glabrous; gynoecium glabrous to densely strigulose, the
stigma truncate to capitellate, sometimes obliquely so, the style about
as long as the elliptic ovary, the ovules 10-14, in two rows but only
the one nearest the style maturing, the gynophore basicentric, to 2 mm
long; fruit oval, inflated, one-seeded, the surface usually reticulate-veined,
glabrous to densely strigulose, 1-3 cm long, the seed yellow brown,
tan, or black, oval, arillate in two species but usually exarillate, the
funicle elongate (to 3 m) in the type species.
Type-Species: Bocoa prouacensis Aublet, Pl. Guiane Fr. (Suppl.)
Soh ts Goll, U1.
A Revision of the Genus Bocoa
Be
Fic. 1. Bocoa alterna—Type I-A. a. Equatorial view. x 3500. b.
Portion of surface. >< 10,000.
102 Proceedings of the Biological Society of Washington
Fic. 2. Bocoa mollis—Type I-A. a. Equatorial view, showing prom-
inent os. x 3900. b. Polar view. x 3900.
A Revision of the Genus Bocoa 103
Fic. 3. Bocoa racemulosa—Type I-A. a. Polar view, showing short
colpi. xX 3700. b. Equatorial view. x 3700.
104 Proceedings of the Biological Society of Washington
Fic. 4. Swartzia flaemingii var. psilonema—Type J-A. a. Polar
view. X< 3000. b. Equatorial view. x 3000.
Fic. 5. Swartzia guianensis—-Type I-A. Polar view. x 3000.
b. Slightly oblique equatorial view, showing very coarsely rugose
ektexine. < 3000.
106 Proceedings of the Biological Society of Washington
Fic. 6. Swartzia panacoco var. cardonae—Type I-B. a/b. Equa
torial views. x 3000.
A Revision of the Genus Bocoa
Fic. 7. Swartzia cuspidata—Type I-B. a. Oblique polar view, show-
ing striato-reticulate ektexine. > 3000. b. Equatorial view. x 3000.
108 Proceedings of the Biological Society of Washington
Fic. 8. Swartzia jorori—Type I-B. a. Oblique equatorial view.
< 3000. b. Portion of surface of ektexine, showing relatively narrow,
interwoven muri. xX 10,000.
A Revision of the Genus Bocoa 109
Fic. 9. Swartzia arborescens—Type I-B. a. Equatorial view, showing
prominent os and discrete orbicules. > 3600. b. Enlargement of mid-
section of colpus. x ca. 10,000.
110 Proceedings of the Biological Society of Washington
Fic. 10. Aldina latifolia—Type I-C. a. Equatorial view. x 3000.
b. Oblique polar view. x 3000.
A Revision of the Genus Bocoa feet:
Fic. 11. Zollernia vogelii—Type I-C. a. Polar view. Xx 3000. b.
Slightly oblique equatorial view. x 3000.
112 Proceedings of the Biological Society of Washington
Fic. 12. Cyathostegia matthewsii—Type I-C. a. Slightly oblique
equatorial view. >< 3000. b. Equatorial view. x 3000.
A Revision of the Genus Boco
Fic. 13. Swartzia benthamiana—Type I-A. a. Polar view, showing
syncolpate condition. x 3000. b. Equatorial view. > 3000.
114 Proceedings of the Biological Society of Washington
Fic. 14. Swartzia panamensis—Type II-B. a. Polar view, showing
syncolpate condition. x 3000. b. Oblique equatorial view. >< 3000.
A Revision of the Genus Bocoa 115
KEY TO THE SPECIES
im leeaveseunitoliolate or titoholate) =. 2 2}
eavesiwithi more than 13 leaflets 22208 ee 4
2. Flowers lacking a petal; leaf rachis and petioles about equal in
length or leaves unifoliolate; seeds obviously arillate; large trees
WARIO RINOe Call emcee ee ee ee 5
Flowers with a petal; leaf rachis several times as long as the petioles;
seeds exarillate; shrub or small tree 3. B. racemulosa
3. Leaves trifoliolate, the stipules deciduous, lanceolate, 3-4.5 mm
long; inflorescence tomentulose; stamens 25-30; funicle not
GOING AUC Clem Mem eerie Sek se eet ln 4. B. viridiflora
Leaves unifoliolate, the stipules persistent, broadly triangular, 1.5
mm or less in length; inflorescence glabrous; stamens 7-10;
HUMIC oWIb Oo TN MONG fee Ne 5. B. prouacensis
4. Leaflets alternate, the apices acute to acuminate, about 7-13 cm
JMS? ei Ni a ec ere oe 5
Leaflets opposite to subopposite, the apices obtuse to rounded,
usually at least some emarginate or retuse, 0.5-5 cm long 6
5. Petal deciduous or caducous; stamens 13-15; stigma obliquely
capitellate; inflorescence 30-50 mm long; calyx segments free
throughoutcdiem length 28s 1. B. alterna
Petal persistent even with old flowers; stamens 6-11; stigma trun-
cate; inflorescences 3-10 mm long; calyx segments not separating
irom oaSaleemotsleme tines. 200 Neel ee Bee 2. B. limae
6. Branchlets, lower surface of leaflets, and inflorescence axes puber-
ulous to tomentulose; leaflets 35-55 mm long, the lower ones oval
to elliptic; inflorescence axes 9-20 mm long; calyx segments be-
coming strongly revolute; ovary and fruit glabrous _ 6. B. mollis
Branchlets, lower surface of leaflets, and inflorescence axes minutely
strigulose, glabrous or glabrescent; leaflets 3-22 mm long, the
lower ones round; inflorescence axes 2—2.5 mm long; calyx seg-
ments reflexed; ovary and fruit strigulose __________ 7. B. decipiens
1. Bocoa alterna (Bentham) Cowan, comb. nov.
Swartzia alterna Bentham, Hook. Jour. Bot. 2:89. 1840.
Description: Shrub or small tree 3-8 m tall, the trunk slender, to 10
cm in diameter, the bark when bruised with a foetid odor, the branchlets
puberulous to strigulose, sometimes glabrescent, the stipules usually per-
sistent, more or less triangular 0.3-0.4 mm long, strigulose externally;
petioles terete to subterete, (5—)12-18(-25) mm long, more or less
puberulous, sometimes glabrescent, the rachis (3—)6—8(—11.5) cm long,
terete to subterete, more or less puberulous; leaflets (3—)5—7, subopposite
to alternate, always with one terminal on the rachis, the petiolules (1—)2-
4(-6) mm long, usually sparingly puberulous, the blades mostly elliptic
to ovate-elliptic, sometimes narrowly elliptic, lanceolate-elliptic, or ovate
(4.5—)7—12(-20.5) cm long, (2.8—)3.5-6(-7.5) cm wide, the base
116 Proceedings of the Biological Society of Washington
cuneate and acute or rounded but finally acute, the apex acute to
acuminate with the tip acute to obtuse, usually glabrous except pu-
berulous on the costa above, occasionally soft-puberulous on the
undersurfaces generally, often somewhat lucid on one or both surfaces,
the venation prominulous on both surfaces or subobscure above, the costa
impressed on the upper surface, salient beneath; inflorescences (1—)3-5
(—8) em long, racemose or sparsely branched panicles of racemes, usually
several arising together at the same node below the leaves of the current
season but sometimes also axillary, the axis strigulose or puberulous, the
flower scars prominently raised, the bracts about triangular and 1 mm
long and wide, persistent or tardily deciduous, the pedicels 4-7 mm long
in flower, 6-13 mm long in fruit, sparingly strigulose or puberulous, the
buds 5-6 mm long, 3 mm in diameter, ellipsoid, sparsely strigulose except
for the tufted apex; calyx segments 3 or rarely 4, free to the base,
revolute tightly at anthesis, glabrous on the inner surfaces; petal one,
white, glabrous, obovate to oblanceolate, the claw (1—)3-6 mm long,
the blade 5-8 mm long, 3-5 mm wide; stamens (12—)13-15(-—18), the
filaments filiform, 3.5-5 mm long, usually free but sometimes shortly
joined basally, the anthers narrowly oblong, 3-4 mm long, 0.7—0.8 mm
wide; gynoecium glabrous or occasionally with a few hairs on the ovary
and rarely puberulous, the stigma obliquely capitellate, the style 3-5.5
mm long, the ovary oblong, 1.5-4.5 mm long, 1-2.5 mm wide, the
gynophore 0.8-2 mm long, ovules 14-16 in each ovary but only one
(usually nearest the style) maturing; fruits dehiscent, usually glabrous,
rarely puberulous, more or less reticulate-veined on the surface, oval to
round in outline, 1-2 cm long, 1-1.5 cm wide, the seed black, oval to
oblong-oval, 8-10 mm long, 5—5.5 mm in diameter, exarillate.
Type-Collection: 1. Riedel 1 (holotype K), “Barra do Rio Negro in
Brasilia.” The specimens cited below as “Riedel s.n.” are from the
type locality and may be isotypes.
Distribution: Guyana, Amazonian Brazil, and western Peru in dense
primary or open, dry secondary forest on sandy or clay soils above
annual flooding by the rivers in the lowlands and up to 800 m elevation.
Discussion: Since there are small local morphological differences
among the collections representing the species sensu lato, the following
are cited under each of several unnamed “phases” I have distinguished.
They are not worthy of formal recognition but should not be passed
over without notice.
Key to PHASES OF B. ALTERNA
1. Leaflets soft-pubescent on the lower surfaces
eR Gn sence | nc A Ne “Pubescent Leaflet Phase”
Leaflets glabrous or minutely puberulous on costa 2
2 Eiriitss pubescent mamas. aie nna enanne nna “Pubescent Fruit Phase”
Eruits glabrous. 2... ee eee 3
3. Fruits about 10 mm long on pedicels 4-5 mm long —_------------------
. “Small Fruit Phase”
A Revision of the Genus Bocoa M7
MNES OS ..F NY EE ee ut “Acuminate Leaflet Phase”
Leaflets acute to acuminate, elliptic, ovate-elliptic, or lanceolate-
GIONS = 2 a Se SE eek eee a et “Glabrous Phase”
(1) The “Glabrous Phase” is glabrous in most parts, particularly the
leaflets and the gynoecium. Collections representing this phase are:
GUYANA: basin of Essequibo R., near mouth of Blackwater Creek,
December 1937, A. C. Smith 2828 (A, K, NY); Wawitau, Kanuku Mtns.,
September 1948, Wilson-Browne 176 (Record No. 5669 (K, NY). PERU:
Depto. Junin, Puerto Bermudez, July 1929, Killip & Smith 26502 (US).
BRAZIL-AMAZONAS-vicinity of Manaus: Igarapé do Binda, April 1956,
Coelho (INPA No.) 3740 (US); Barcelos, September 1962, Duarte 6978
(US); Flores, January 1941, Ducke 371 (MADw); Cachoeira Grande,
April 1937, Ducke 454 (A,F,LAN,MG,MO,NY,US); Cachoeira below
Taruma, 21 September 1956, Francisco & Luiz 4188 (IAN); Estrada do
Mundu, February 1945, Froes 20465 (IAN,K,NY,US); Pai Raimundo,
Rio Demeni, Froes & Addison 29016 (IAN); Rio Taruma, Aug. 1949,
Froes 24946 (IAN); Flores, March 1924, Kuhlmann 96 (P, U) and 1607
(RB,US); Cachoeira baixa de Taruma, September 1956, Mello & Coelho
(INPA No.) 4188 (US); Kms 65-70 da Rodovia, Manaus-Itacoatiara,
22 Oct. 1963, Oliviera 2742 (IAN); Estrada do Aleixo, March 1947,
ires 213 (US); Road Igarapé Leaé 25 km north of Manaus, November
1966, Prance et al. 3137 (U,US); estrada Manaus-Itacoatiara km 70,
October 1960, Rodrigues & Coelho 1877 (US); Estrada do Taruma,
January 1962, Rodrigues & Chagas 4112 (US); Igarapé do Buiao, No-
vember 1962, Rodrigues & Chagas 4723 (US); Km 22 da estrada
Manaus-Caracaroi, November 1962, Rodrigues et al. 4811 (US); Manaus,
Schwacke 3942 (RB); between Santarem and Barra do Rio Negro
along the Amazon River, October 1850, Spruce s.n. (BM,G,K,NY,RB);
Paruna-miri dos Ramos, Oct. 1850, Spruce 1106 (K,P). PERNAMBUCO:
Recife, Dois Irmaos, Feb. 1948, Ducke 2116 (IAN).
(2) The “Pubescent Leaflet Phase” has leaflets that are softly pubes-
cent on the undersurfaces, but like the typical phase in other respects.
All the collections are from BRAZIL-AmMazonas-Manaus: B.A.M., BR
17, Km 9, December 1955, Dionisio 3076 (MG); Flores, October 1936,
Ducke 296 (A,F,IAN,K,MG,MO,NY,US); between the city and Flores,
July 1929, Ducke (HJBR No.) 23342 (G,K,RB,US); near the city, August
1931, Ducke (HJBR No.) 24203 (G,K,P,RB,U,US); road to Aleixo,
August 1936, Krukoff 7944 (A,BM,G,K,MO,P,U,US); Margem do
Igarapé do Buiao, July 1956, Luis 13979 (IAN); Sept. 1945, Pires &
Black 930 (IAN); Barra do Rio Negro, Riedel s.n. (A,G,K,P); Igarapé
do Santa Maria, June 1962, Rodriguez and Chagas 4473 (US), cultivated
at Rio de Janeiro, Dec. 1878, Glaziou 9767 (K,P,U).
(3) The “Acuminate Leaflet Phase” differs in having relatively
narrower leaflets that have long-acuminate apices and it occurs in the
Kanuku Mountains of Southwestern Guyana: March—April 1938, A. C.
Smith 3147 (F,IAN,K,NY,P,U,US) and 3575 (G,F,IAN,K,MADw,MO,
118 Proceedings of the Biological Society of Washington
NY,U,US). One additional Guyana collection without locality should be
assigned here: Robt. Schomburgk 293 (Rich. Schomburgk 585) (K,P).
(4) The “Pubescent Fruit Phase” is separable only by the puberulous
fruits and is represented by a single collection from Guyana: basin of
Essequibo River near mouth of Onoro Creek, December 1937, A. C.
Smith 2772 (A,F,G,K,MADw,MO,NY,U,US).
(5) The “Small Fruit Phase” has noticeably smaller fruits, somewhat
shorter pedicels, and longer inflorescences: Serra do Navio, Terr. do
Amapa, Rio Amapari, Brazil, November 1954, Cowan 38157 (F,NY).
(6) The “Unifoliolate Phase” is identical with the “typical phase”
except that the leaves are uniformly unifoliolate. The only two collections
are from the Brazil-Guyana boundary in the Akarai Mountains, at 600—
800 m elevation between the drainages of the Rio Mapuera and Shodikar
Creek: A. C. Smith 2991 (A,F,C.K,MADw,MO,NY,U,US) and 2996
(A,F,G,K,MO,NY,P,U,US).
The closest relationship of B. alterna is with B. limae, which shows
once again a familiar distributional pattern (in the legumes at least )—
a wide-ranging species with a disjunct, very similar, related species in the
coastal forest of southeastern Brazil. The two taxa differ in the length
of the inflorescence, persistence of the petal, and the post-anthesis union
of the calyx-segments.
2. Bocoa limae Cowan, sp. nov.
Description: Frutex 1.5-2 m altus, trunco ca. 3 cm diametro, ramulis
strigulosis, stipulis caducis, non visis; petioli 3-4(-6) mm longi, sparse
strigulosi, subteretes, rhachibus 4.5-8 cm longis, sparse strigulosis, sub-
teretibus; foliola 4-7, alterna, imparipinnata, petiolulis 1-2 mm _ longis,
laminis superioribus amplioribus, 6—20 cm longis, 2.5—7.5 cm latis, laminis
inferioribus minoribus, 3-5.5 cm longis, 1.5-3 cm latis, laminis omnibus
ellipticis, ad basim angustatis acutis vel tandem obtusis, ad apicem argute
vel obtuse acutis, sparse strigulosis infra sed mox glabrescentibus, lucidis,
manifeste venulosis in ambobus paginis; inflorescentiae ramuligerae vel
axillares, axe 3-5(-10) mm longo, striguloso, bracteis deciduis, tri-
angularibus, strigulosis extus; pedicelli 4—5.5 mm longi, strigulosi, ala-
bastris oblongis, sparse strigulosis; calycis segmenta 3 vel 4, revoluta, ad
basim incomplete fissa; petalum unum, album, persistens, glabrum,
unguiculo 3.5-4 mm longo, lamina rotundata sed attenuata ad basim,
5-7 mm diametro; stamina 6-11, filamentis 3-5 mm longis, antheris
anguste oblongis, ca. 4 mm longis et 1 mm latis; gynoecium glabrum,
stigmate truncato, stylo ca. 5 mm longo, ovario elliptico, ca. 3.5 mm
longis et 1.8 mm latis, gynophoro ca. 1 mm longo; fructus ovalis, glaber,
reticulatus, ca. 18 mm longus et 12 mm latus, rostellatus, semine uno,
nigro, exarillato, ovali ca. 10 mm longo et 7 mm lato.
Type-Collection: A. Lima 65-4270 (holotype US, sheet no. 2639742,
isotype IPA), “Bahia. Salvador. Areias em torno da Lagéa do Abaeté,”
Brazil, 26 January 1965.
Distribution: Coastal brush-forest from Pernambuco to Bahia, Brazil,
A Revision of the Genus Bocoa IL,
on sandy soils. BRAZIL-PERNAMBUCO: Recife, woods near Dois Irmaos,
29 October 1949, Lima 49-369 (IPA); along road to Aldeia, 17 March
1952, Lima 52-1007 (IPA, US); mata de Dois Irmaos, 2 June 1966,
Soares 2185 (US).
Vernacular Name: “Feijao brabo” (Soares 2185).
Discussion and Etymology: There is a great resemblance of B. limae
(named for the collector of most of the known material of the new taxon)
to B. alterna but close examination reveals a number of differences in
addition to the obvious geographic disjunction. Unlike its nearest relative,
the new species has a very short inflorescence, at times the flowers almost
seeming to arise from the branchlets directly; the calyx segments do not
open completely, remaining partially united basally; the petal is persistent,
even with quite old flowers; the stamens are fewer, and the stigma is
truncate; vegetatively, the leaflets are obviously venulose and the costa is
salient on both surfaces.
3. Bocoa racemulosa (Huber) Cowan, comb. nov.
Swartzia racemulosa Huber, Bol. Mus. Goeldi 5:395. 1909.
Description: Shrub or small tree, the branchlets sparingly strigulose
minutely, glabrescent, the stipules not seen, the petioles 2-10 mm long,
sparingly strigulose but glabrescent soon, rachis 2.5-6 cm long, glabrous,
subterete; leaflets 1-3, the lateral ones smaller (6-11 cm long, 3-7 cm
wide) ovate, with petiolules 1-3 mm long, the terminal leaflet 12—23.5
cm long, 5-12 cm wide, the petiolules 3-4 mm long, base of leaflets
rounded-obtuse (some lateral ones) to rounded and acute, or simply
acute, the apex sharply to bluntly acute, sometimes emarginate, glabrous,
venation prominulous; inflorescences 2.5-3 cm long, ramuligerous,
racemose, minutely strigulose, the bracts persistent, ovate-triangular,
about 1 mm long, minutely strigulose externally; pedicels about 2.5 mm
long in flower and 4 mm long with fruit, minutely strigulose sparingly,
the calyx minutely strigulose externally, opening only about half of length
in 2-4 segments, ca. 3.5-4 mm long, segments erect or somewhat reflexed,
the calyx usually falling in one piece; petal white, glabrous, obovate, 7
mm long (fide Huber), cuneate-unguiculate; stamens 12—14, the filaments
2-3 mm long, the anthers oblong, 2.5-2.8 mm long, 0.5 mm wide;
gynoecium glabrous, the stigma capitellate, the style 3.5 mm long, the
ovary elliptic, ca. 2 mm long and 0.8 mm wide, the gynophore ca. 1 mm
long; fruit ca. oval in outline and 1.5 cm long, 1 cm wide, reticulate-
veined on surface, glabrous, the seed exarillate, black, oval-oblong.
Type-Collection: A. Ducke 7870 (holotype presumably in Brazil but
not at Museu Goeldi in Belém (fide the curator), isotypes F,G), “in
silvis flumenis Trombetas ad vicum Oriximina, Para,” Brazil, 8 December
1906.
Distribution: Lowland forest of Para in east-central Brazil, near the
junction of the Amazon and the Rio Trombetas: Sao Jorge, Municipio
de Faro, November 1950, Black & Ledoux 50-10669 (IAN,US); Orixi-
120 Proceedings of the Biological Society of Washington
mina, 23 November 1907, Ducke 8865 (BM,G,MG); Rio Tapajoz,
Mangabal, February 1917, Ducke 11184 (HAMP No. 16755) (BM,G,
MG,P,RB,U,US); S. Luiz do Rio Tapajoz, December 1915, Ducke 11185
(HAMP No. 15856) (BM,MG,RB); Obidos, November 1919 (fl.), June
1926 (frt.), Ducke 11186 (G,K,P,RB,U,US); Rio Tapajoz, Villa Braga,
November 1908, Snethlage 10058 (BM,G,MG).
Discussion: While clearly related to B. alterna, this species is easily
distinguished by its dimorphic leaflets. The pair of lateral ones is ovate
and about half the length of the elliptic terminal one. In addition, the
calyx segments in B. racemulosa only separate about halfway and they
are more or less reflexed, in contrast to the totally free, retrorsely revolute
segments in B. alterna.
4. Bocoa viridiflora (Ducke) Cowan, comb. nov.
Swartzia viridiflora Ducke, Arch. Inst. Biol. Veg. 2:44. 1935.
Description: Tree to 30 m tall and 43 cm diameter, with glabrescent,
lucid branchlets, strigulose at very early stages, the stipules deciduous,
lanceolate, 3-4.5 mm long, 1-1.5 mm wide, strigulose externally at first
but soon glabrescent; leaves glabrous, trifoliolate (unijugate and im-
paripinnate ), the petioles 1-4 cm long, terete, the rachis 2—4.3 cm long,
terete; lateral leaflets opposite, occasionally subopposite, petiolules 3-6
mm long, transversely rugose, the blades subcoriaceous, lucid, 7-15 cm
long, 3-6.5 cm wide (the terminal leaflet larger), elliptic, the base acute,
the apex acuminate to bluntly acute, costa and 3-5 pairs of primary
veins more or less salient on both surfaces; inflorescences 5-9 cm long, to
20 cm long in fruit, ramuligerous, axes albo-tomentulose, the peduncle
3-4 mm long, the bracts persistent, darker colored, semicircular, cucullate,
about 1 mm long and broad, puberulous externally; flowers greenish,
fragrant, the pedicels tomentulose, about 1 mm long; calyx segments
deciduous, sparingly tomentulose externally, glabrous on inner surfaces;
petal absent; stamens equal, 25-30, the filaments 6 mm long, the anthers
basifixed, oblong ca. 1.5 mm long and 0.5 mm wide, gynoecium glabrous,
the stigma capitellate, the style 3-4 mm long, the ovary oblong, 3 mm
long 1.5 mm wide, the gynophore 1.5-2 mm long; fruits oval in outline,
ca. 3 cm long and 1.5 cm in diameter, the surface tessellate-scaly, the
“scales” darker, the seed oval, only slightly smaller than the fruits,
the aril small.
Lectotype-Collection: A. Ducke (H.J.B.R. No.) 24219 (flowering por-
tion). (Lectotype RB, isolectotypes F,G,K,MADw,NY,P,U,US), “circa
Manaos (civ. Amazonas) in silvis loco alto ultra Flores,” Brazil, 29 No-
vember 1932.
Distribution: Vicinity of Manaus, Brazil, southern Guyana and Suri-
name, in riverine lowland forest. BRAZIL-Amazonas: Vicinity of
Manaus above Flores, 18 April 1933, Ducke (H.J.B.R. No.) 24219
(fruiting portion) (F,G,MADw,NY,P,US) and 1 March 1946, Ducke
1923 (A,F,IAN,K,MG,NY,US). BRAZIL-ParaA: Tinguelim km 21, 30
A Revision of the Genus Bocoa 121
March 1970, Silva 3024 (IAN). GUYANA: “Plot 3, hill top 1000 feet
high, 7 miles E. of Onoro mouth, Upper Essequibo,” 30 September 1952,
Guppy 304 (Record No. 7280) (NY). SURINAME: Fallawatra, 8 Nov.
1971, Jimenéz-Saa 1568 (K).
Choice of a lectotype is required in this instance because Ducke in-
cluded two collections under one number, probably from the same tree
but this is not at all certain; both parts of the type collection are cited
here.
The collection from Guyana is sterile but it almost certainly represents
this species.
Vernacular Names: “Gombeira amarela” (Silva 3024).
Discussion: This is one of the two species of the genus which are char-
acterized by apetalous flowers and arillate seeds. It is distinct from its
near relative, B. prouacensis, by the number of leaflets, stipule size, and
number of stamens. Like B. racemulosa, although less pronounced, B.
viridiflora has trifoliolate leaves with the terminal one the largest. This
species has the largest stipules, the longest inflorescences, and the
largest number of stamens.
5. Bocoa prouacensis Aublet, Pl. Guiane Fr. Suppl. 38, t. 391. 1775.
Swartzia prouacensis (Aublet) Amshoff, Meded. Bot. Mus. & Herb. Rijks.
Univ. Utrecht 52:40. 1939.
Swartzia minutiflora Kleinhoonte, Rec. Trav. Bot. Neer]. 22:408. 1925.
Description: Tree 14-30 m tall, the trunk 19-25 cm diameter, straight
and cylindrical, the bark brownish-grey with small rectanglar scales, the
branchlets glabrous, the stipules persistent, rigid, 0.7-1.3 mm long and
wide, broadly triangular-ovate, acute, glabrous; leaves glabrous, uni-
foliolate, the petiole subterete, non-alate, 6-16 mm long or occasionally
totally suppressed, petiolules 4-7 mm long, transversely corrugate, the
blades coriaceous, 9—-18(-—21) cm long, 4.5-8(-10) cm wide, mostly
elliptic but sometimes elliptic-ovate or broadly ovate, sometimes asym-
metric, the base rounded and obtuse to subobtuse, infrequently acute,
the apex bluntly acute, infrequently acuminate, the costa and its primary
branches more or less salient on both sides of the leaflet-blade; in-
florescences 2-5 cm long, ramuligerous, axes glabrous, the bracts per-
sistent, cucullate, broadly ovate, about 0.6-0.8 mm long and 1 mm wide,
glabrous except for ciliolate margin, bracteoles lacking; pedicels 0.3-0.6
mm long, rarely to 2 mm long, glabrous, the calyx opening nearly to the
base in three, more or less equal segments which often fall in one piece;
petal absent; stamens 7-10, uniform, the filaments 4.5 mm long, the
anthers oblong, 1.3-1.5 mm long, ca. 0.5 mm wide; gynoecium glabrous,
the stigma capitellate, the style filiform, 1.7-2 mm long, the ovary 1.8—2
mm long, 0.8—1 mm broad, oblong, the gynophore 1—-1.5 mm long; fruit
oval in outline, 2.5-3 cm long, 1.5-2.5 cm wide, dehiscent, the seed
yellow or light brown (fide Amshoff) 1-2 cm long, the aril white,
laciniate, the funicle filiform, 1.5-3 m long.
122 Proceedings of the Biological Society of Washington
Type-Collection: J. B. C. F. Aublet s.n. (BM), “Habitat in sylvis
Caux,” French Guiana.
Distribution: Suriname and French Guiana in lowland forest. SURI-
NAME: Marechalskreek, 9 July 1921, BW 5441 (U) and 21 Jan.
1922, BW 5769 (U); Brokopondo District, 8 km ESE of Brownsweg
Village, 26 Mar. 1965, van Donselaar 2273 (U); Tapanahoni River, 11
Nov. 1918, Gonggrypp 4175 (IAN,U); Nassau Mts., 9 Mar. 1949,
Lanjouw & Lindeman 2545 (U); Boven Coesewijne, ca. 20 km SW
of Poika, Schulz 7926 (U); Sectie “O,” 23 Apr. 1915, Suriname
Forestry Department 342 (K,MO,U ),—23 Oct. 1915, 1191 (U) (type
of Swartzia minutiflora Kleinhoonte),—27 Oct. 1915, 1198 (K,MO,
U),—Nov. 1915, 1434 (U),—22 Nov. 1915, 1560 (U),—26 Feb. 1916,
1660 (U),—22 Dee. 19116, 2559 (U) 5 Mar. 1917, ou qunurs) ae
29 Aug. 1918, 3974 (U),—2 Sept. 1918 3984 (NY,U,US)._4 Sep
1918, 3986 (IAN,U),—9 Dec. 1918, 4185 (U),—30 May 1919, 4326
(U),—28 Nov. 1919, 4455 (MO,U),—28 Nov. 1919, 4477 (U),—
21 Jan. 1920, 4521 (U),—6 Mar. 1920, 4569 (IAN,K,U,US),—3
Nov. 1920, 4781 (U),—12 Nov. 1921, 5556 (U),—17 May 1922, 5839
(K,NY,U),—16 Mar. 1923, 6084 (IAN,U); Brownsberg Forest Re-
serve, Suriname Forestry Department 6130 (NY,U); Sectie “O,”’ Dec.
1942, Suriname Woodherbarium (Stahel) 69 (A,IAN,K,MADw,NY,U );
Brownsberg Forest Reserve, 10 Oct. 1969, Tawjoeran 12581 (U).
FRENCH GUIANA: St. Laurent, Feb. 1956, BAFOG 339M (U),
7236 (U), and 7249 (P,U), 7 Feb. 1956, Bena 1117 (U); Placeau
no. 2—Carreau no. 56—Route de Mana,” 3 Jan. 1956, French Guiana
Forestry Department 7126 (NY,P,U), 26 Mar. 1956, 7415 (U); Sinna-
mary River, Crique Grégoire, 28 Apr. 1968, Oldeman B-1614 (P);
Camopi River, 1 km downstream from Saut Ouasseye, 11 Dec. 1967,
Oldeman 2625 (P); Karouany, 1859, Sagot 1210 (BM,K,P); with-
out locality or date, Wachenheim s.n. (BM,K,P,US).
Vernacular Names: “Bois bobo,” “caux bois boco” (Aublet s.n.);
“boko” (Oldeman B-1614); “aie oudou” (F. G. For. Dept. 7126);
“itikiboroballi hohorodikoro,” “ijzerhart” (Sur. Woodherb. 69); “yzerhart”
(Sur. For. Dept. 4326).
Discussion: Bocoa prouacensis is one of the two most frequently
collected species in the genus and certainly one of the most distinctive.
One characteristic alone separates it from all its relatives; the single
seed in each fruit at the time it dehisces is suspended on a thread up
to three meters long. (An interesting description of this phenomenon
is given by Stahel in the Journal of the New York Botanical Garden,
vol. 45:265-268. 1944). This attribute plus an obvious, fleshy aril
is surely of some adaptive significance, probably for dispersal by
animals. The aril in this species, and in the preceding, links the
genus to Swartzia, most of whose species have arillate seeds. The
unifoliolate leaves of B. prouacensis serve to separate it from all its
near-relatives, the closest of which is probably B. alterna.
A Revision of the Genus Bocoa 123
6. Bocoa mollis (Bentham), Cowan, comb. nov.
Swartzia mollis Bentham, Hook. Joum. Bot. 2:89. 1840.
Trischidium vestitum Tulasne, Ann. Sci. Nat. (Ser. 2) 20:14], t. 4.
Sept. 1843.
Swartzia cearensis Ducke, Anais da Academia Brasileira de Ciencias 31:
295. 30 June 1959.
Description: Shrub or small tree 2—4 m tall, foetid in fresh state, the
young branchlets tomentulose densely; stipules early caducous, minute,
ca. 0.5 m long, densely strigulose; petioles 6-11 mm long, tomentulose,
terete, the rachis 25-50 mm long, tomentulose, terete; leaves impari-
pinnate, the leaflets 5-9, 3.5-5.5 cm long, 1.5-3 cm wide, subopposite
or rarely opposite, the petiolules 0.7-1.5 mm long, tomentulose, the
blades oval to elliptic or ovate to lanceolate, the base rounded, obtuse
or slightly cordate, the apex obtuse, retuse, to sub-emarginate, puberu-
lous to tomentulose beneath, rarely only on costa, puberulous to
strigulose above, the venation subobscure to barely prominulous, plane
except the costa salient on lower surface; inflorescences ramuligerous,
racemose, the axes 9-20 mm long, densely tomentulose-pilosulose, the
bracts early deciduous, broadly triangular-ovate or semicircular in
outline, about 1 mm long and wide, densely strigulose externally,
glabrous within, the pedicels 4-12 mm long at anthesis, 7-15 mm long
with mature fruits, pilosulose; buds elliptic in outline, subappressed-
tomentulose, ca. 5 mm long and 3.5 mm diameter, the calyx opening
only about half-way into 3 or 4 segments; petal one, deciduous to
caducous, white, the claw 2-6 mm long, sparingly villosulose externally
or glabrous, the blade glabrous, oblate, 4-7 mm long and 5-7 mm
wide; stamens (12—)20-22, the filaments 4—7 mm long, slightly joined
basally, the anthers narrowly oblong, 2-3 mm long, 0.7-0.9 mm wide;
gynoecium glabrous except sometimes a few long hairs on ovary, the
stigma capitellate, sometimes obliquely so, the style 2-3.5 mm long,
the ovary elliptic, ca. 3.5 mm long and 2 mm wide, the gynophore
ca. 1 mm long; fruit obliquely ellipsoid, beaked with the 1-2 mm
long remnant of style, 10-15 mm long, 8-10 mm diameter, reticulate,
the carpophore about 1-4 mm long; the seed tan to black, exarillate,
5.0.-7 mm long, 4.5-5 mm diameter, shiny, ellipsoid to subrotund.
Type-Collection: J. S. Blanchet 2774 (holotype K, isotypes F
(frag.), G,K,P) (also the type collection of Trischidiwm vestitum
Tulasne). “In Brasilia prope Utinga, Fazenda in Certao de San
Francisco,’ Bahia, 1839.
Distribution: Near-interior of Ceara and Rio Grande do Norte to
Bahia, Brazil, on sandy soils in scrub forest. BRAZIL-Crara:
Baturité, 11 Apr. 1909, Ducke 1980 (BM,G,US); Itaitinga, Pé do
Serrote da Pedreira, 11 August and 23 December, 1955, Ducke 2467
(IAN,INPA,K,MG,NY,US) (type of Swartzia cearensis Ducke) and
Ducke 2582 (IAN,K,NY,RB,US); along the road from Morada Nova
to Bixoba, 14 February 1960, Lima 60-3443 (IPA,US); no specific
124 Proceedings of the Biological Society of Washington
locality, Dias da Rocha (HJBR No.) 11183 (RB); without locality or
date, Allemao & Cysneiros 497 (P). Rio GRANDE po Norte: Acu, along
the margin of the left bank of the Rio Acu, Lima 60-3505 (IPA).
PERNAMBUCO: Margem da estrada Ibimirim-Joazeiro dos Candidos,
Lima 50-700 (IPA); Manicobal, Fazenda Contenda, 20 June 1952,
Lima & Magalhaes 52-1050 (IPA). Banta: Antonio B. de Oliviera
(HJBR No.) 68404 (RB); Raxo da Catarina, Labouriou 912 (F,
RB,US). Mrnas Gerais; 4 km N of Vasante de San Francisco,
Municip. Januaria, 29 Sept. 1953, Magalhaes 6082 (IAN,RB).
Vernacular Names: “Brinquinho” (Oliveira 68404).
Discussion: This species and B. decipiens form a small subgeneric
grouping that may represent a separate phylogenetic offshoot of the
genus. Both have several pairs of emarginate leaflets and small fruits,
and both apparently are shrubs to small scrubby trees in the arid parts
of southeastern Brazil. The smaller, glabrous or glabrescent leaflets,
longer inflorescence, strongly revolute calyx segments, and _ larger
number of stamens amply separate B. mollis from its closest relative.
7. Bocoa decipiens Cowan, sp. nov.
Swartzia decipiens Holmes, Pharm. Journ., ser. 4, 3:2. 4 July 1896.
(Provisional Name)
Diagnosis: Frutex foliorum foliolis basalibus rotundatis 3-9 mm
longis, 5-5.5 mm latis; gynoecium sicut fructus strigulosum.
Description: Shrub with the branchlets minutely strigulose, the
stipules minute; petioles 1-2 mm long, lightly canaliculate on upper
surface, minutely strigulose, the rachis 9-28 mm long, minutely
strigulose, obviously canaliculate-marginate on the upper surface; leaf-
lets 5-7, imparipinnate, the laterals subopposite, the petiolules 0.2—0.5
mm, glabrous, the blades glabrous, smaller and rounded near the base of
the leaves but progressively larger and more elliptic toward the leaf
apex, the basal ones 3-9 mm long, 5-5.5 mm wide, the upper ones
and the terminal one 14-22 mm long, 6-10 mm wide, the base acute
except the basal ones rounded-obtuse, the apex rounded, emarginate,
glabrous or sparingly puberulous minutely on the costa, the venation
subobscure, the costa salient on both surfaces; inflorescences axillary,
racemose, few-flowered, the axis 2-5 mm long, strigulose, the bracts
persistent, ca. 0.5 mm long, the fruiting pedicels 8-10 mm _ long,
glabrous; calyx segments 3 or 4, glabrous, separating to the base, about
4.5 mm long, reflexed; stamens 8-10, glabrous, the filaments 2.5-3
mm long; gynoecium densely strigulose; fruit more or less strigulose,
the basal 1-2 mm of the style persistent as a beak, the body of the
fruit 8-9 mm long and wide, broadly oval in outline, the surface
more or less reticulate, gynophore and carpophore densely strigulose,
0.5-1 mm long; seed black, exarillate, oval in outline, 5 mm _ long,
3.5-4 mm wide.
Type-Collection: A. Lima & M. Magalhaes 52-1075 (holotype US,
A Revision of the Genus Bocoa 125
sheet no. 2639739, isotype IPA), “Nos campos da Serra Araripe, prox.
Est. Exp. Araripina,” Pernambuco, Brazil, 24 June 1952.
Distribution: Known certainly only from the type collection but
circumstantial evidence in the following discussion indicates a broader
range.
Discussion: In the latter part of the 19th Century, Europe was
importing for medicinal purposes large quantities of one species of
Pilocarpus (Rutaceae). As demand exceeded supply, another species
of the same genus gradually came into commerce, as well as other
unrelated adulterant plants. In his discussion of one of these, E. M.
Holmes, who for 50 years was Curator of the Materia Medica Mu-
seum of the Pharmaceutical Society in London, noted in the paper
referred to above, “Some of the more’ recent importations of these
leaves (Pilocarpus microphyllus) have contained a few bales of leaves
almost indistinguishable from them to the eye of the casual observer,
. attention has probably been directed to them by their not yielding
pilocarpine.” He was encouraged by W. B. Hemsley at Kew to
look among the legumes for the indentity of the plant parts in the
“few bales” and it is very much to Holmes’s credit that he correctly
related his scraps to Swartzia, three species in particular—S. mollis,
S. pilulifera, and S. matthewsii. He concluded that his material “prob-
ably” represented an undescribed species which he characterized quite
adequately, considering the quality of his material. However, the
last sentence leaves considerable doubt with respect to the validity of
the publication of the new species: “Until further specimens are pro-
curable from which it might be determined if only one petal and five
stamens are present, the plant might be provisionally named Swartzia
decipiens.” Article 34 of the Code states (in part), “A name is not
validly published . . . (2) when it is merely proposed in anticipation
of the future acceptance of the group concerned .. .” Having con-
cluded that Holmes’s description is invalid, I have supplied a Latin
diagnosis and as full an English description as possible to make the
same specific epithet available in Bocoa where the species properly
belongs. Its very small leaflets and pubescent fruits serve to distinguish
it readily from all the other species in the genus.
The choice of type collection indicated above requires some explana-
tion, for ordinarily I would probably have used the older material
on which Holmes had based his description. However, this was not
practical in this instance. I have seen no material certainly studied
by Holmes and used by him in drawing up his description, although
there are two sheets at Kew which have been viewed as “type
material.” One is a small packet with a label bearing the following
notation: “Swartzia decipiens Holmes / fruits picked out of the leaves
in commerce. March 1897 / Comm E. M. Holmes / See Ph. Journ.
July 4.96. p. 2.” If one were to accept his description as validly
published, this fragment would have to be viewed as the type. In
addition, there is at Kew a “type” sheet bearing several small packets
126 Proceedings of the Biological Society of Washington
and a note “with 2 samples (A & B) of ‘genuine Paraguay Jaborandi
leaves’ / comm. by J. W. Drysdale & Co. / recd. 21.3.01.” On the
note there are sketches in pencil of calyx segments, a gynoecium, a
flower with pedicel, calyx segments, and ten antherless filaments (the
anthers, petal, and gynoecium are omitted from the flower sketch);
there is also a representation of a partly opened ovary with ten ovules.
The Berlin Herbarium likewise had one or more sheets of fragments
obtained from the same source, the drug importing firms of Western
Europe; copies of the photograph made by Macbride of the Berlin
sheet are in several world herbaria but only the one at the Field Museum
has a packet of the fragments from the original source. All these ma-
terials I have seen certainly represent the new species.
The small leaflets, fewer stamens, and smaller strigulose fruits
serve to separate this poorly known taxon from B. mollis, with which it
shares numerous characteristics. In fact, one can speculate, appar-
ently rather safely, that B. decipiens is an evolutionary derivative
of B. mollis.
LITERATURE CITED
AmsHoFF, G. J. H. 1939. On South American Papilionaceae. Meded.
Bot. Mus. & Herb. Rijks. Univ. Utrecht 52:1-78.
AuBLET, F. 1775. Histoire des plantes de la Guiane Francaise (Sup-
plement). 38, pl. 391.
BENTHAM, G. 1840. Contributions towards a flora of South Amer-
ica—Enumeration of plants collected by Mr. Schomburgk in
British Guiana. Hooker’s J. Bot. 2:38—102.
1862. On Inocarpus. J. Linn. Soc. London 6:146—150.
Cowan, R. S. 1968. Swartzia (Leguminosae, Caesalpinioideae,
Swartzieae). Flora Neotropica 1:1-228.
Rupp, V. E. 1970. Etaballia dubia, a new combination. Phytologia
20:426—428.
ScHREBER, J. C. D. 1789-1791. Genera plantarum 1:364-365; 2:
518.
TuLasneE, L. R. 1843. Nova quaedam genera Leguminosarum. Ann.
Sci. Nat. (Ser. 2) 20:141. Sept. 1843.
WILLDENow, C. L. 1800. Species plantarum 2:1219-1221.
List oF EXSICCATAE
The figures in parentheses refer to the number assigned to each taxon.
Allemao & Cysneiros Bena, P.
A497 (6) Ney (5)
Aublet, K. B. C. F. Black, G. A. & Ledoux, P.
sn. (5) 50-10669 (3)
BAFOG Blanchet, J. S.
339M (5); 7236 (5); 7249 (5) 2774 (6)
A Revision of the Genus Bocoa
Coelho, D.
3740 (1)
Cowan, R. S.
38157 (1)
van Donselaar, J.
DONS ((S)))
Dias da Rocha
11183 (6)
Dionisio
3076 (1)
Duarte, A. P.
6978 (1)
Ducke, A.
QI (ibe Sl Gye
1923 (4); 1980 (6); 2116 (1);
2467 (6); 2582 (6); 7870 (3);
8865 (3); 11184 (3); 11185 (3);
11186 (3); 15856 (3); 16755
(oyemoo42e ee): 24203) (1b):
24219 (4)
Francisco & Luis
4188 (1)
French Guiana Forestry Depart-
ment
MAGN (S) 27415 (5)
Froes, R. L.
20465 (1); 24946 (1)
Froes, R. L. & Addison
29016 (1)
Glaziou, A. F. M.
9767 (1)
Gonggrijp, J. W.
4175 (5)
Cuppy, N. G. L.
304 (4)
Jiménez-Saa, H.
1568 (4)
Killip, E. P. & Smith, A. C.
26502 (1)
Krukoff, B. A.
7944 (1)
Kuhlmann, J. G.
96 (1); 1607 (1)
Labouriau, L.
O27 (G6)
Lanjouw, J. & Lindeman, J. C.
2545 (5)
454 (1);
127
Lima, A.
49-369 (2); 50-700 (6); 52-
1007 (2); 60-3443 (6); 60-3505
(6); 65-4270 (2)
Lima, A. & Magalhaes, M.
52-1050 (6); 52-1075 (7)
Luis
13979 (1)
Magalhaes, M.
6082 (6)
Melo, F. & Coelho, L.
4188 (1)
Oliveira, A. B.
68404 (6)
Oliviera, E.
DA (11)
Oldeman, R. A. A.
B-1614 (5); 2625 (5)
Pires, J. M.
213 (1)
Pires, J. M. & Black, G. A.
930 (1)
Prance,-G. T. et al.
Cs (1)
Riedel, L.
I (i) ssn, (il)
Rodrigues, W. et al.
MSTA (ils ZU) (ibs 448s (il ))e
AV 23) 481i)
Sagot, P.A.
1210 (5)
Schwacke, C. A.
3942 (1)
Schomburgk, Rich.
585 (1)
Schomburgk, Robt.
293 (1)
SchulZ7 pee
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Silva, N.T.
3024 (4)
Smith, A. C.
DTA (IL )e Seeks} (Ibe Ql (IL is
INS (i)e Silay Cis Baris (3)
Snethlage
10058 (3)
128 Proceedings of the Biological Society of Washington
Soares, J. 4455 (5); 4477 (5)s 4521) (5k
ZNSon E23) 4569 (5); 4781 (5); 5441 (1);
Spruce, R. 5996 (5); 5769 (5); 5839 (5);
1106 (1); s.n. (1) 6084 (5); 6130 (5)
Stahel, GC. Suriname Woodherbarium
69 (5) aa
; Tawjoeran, J.
Suriname Forestry Department 12581 (5)
342/15) 1S 5) OSI (S) Sane esas
1434 (5); 1560 (5) 1660 (5); am, (3)
2552 (5); 2743 (5); 3984 (5); Wilson-Browne, G.
3986 (5); 4185 (5); 4326 (5); 176 (1)
x
Vol. 87, No. 14, pp. 129-138 25 April 1974
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
THE “SEA FLEA” DOLOBROTUS MARDENI N. GEN.,
N. SP., A DEEP-WATER AMERICAN LOBSTER
BAIT SCAVENGER (AMPHIPODA:
EUSIRIDAE)
By THomas E.. BOWMAN
Smithsonian Institution, Washington, D.C. 20560
Amphipod crustaceans called “sea fleas” or “sand flies” are —
well known to American and Canadian lobstermen. These
amphipods frequently scavenge the bait placed in lobster
pots; they will also attack and devour trapped lobsters that
have been wounded, but apparently do not molest uninjured
lobsters.
Fishermen have been catching American lobsters in baited
traps for well over a century, and we may suppose that
amphipod scavengers have been feeding on the bait and have
been noticed by fishermen for about the same length of time.
Yet the voluminous literature on the delectable Homarus
americanus contains no references to “sea fleas” until the
appearance in recent decades of a few notes on these bait
scavengers (Templeman, 1954, 1958; Scarratt, 1965; Shave,
1966). Thus far 2 species of “sea fleas” have been identified:
Anonyx sarsi Steele & Brunel and Orchomenella pinguis
(Boeck). Both are members of the family Lysianassidae,
notorious for its scavengers in the cooler waters of the world
oceans (references in Vader, 1972; see also Hessler, Isaacs and
Mills, 1972; Bowman and Manning, 1972; Paul, 1973). Two
additional amphipod species were collected recently from
baited lobster pots by Luis Marden, Chief, Foreign Editoral
Staff, National Geographic Magazine, while collecting infor-
mation for an article about the American lobster (Marden,
1973). Mr. Marden kindly sent his specimens to me, and later
forwarded other samples taken from lobster pots by William
14—Proc. Brot. Soc. WAsH., VoL. 87, 1974 (129)
130 Proceedings of the Biological Society of Washington
Dion, Chief Engineer of the lobster trawler Mars. One of Mr.
Marden’s samples, collected near Petit Manan light, Maine
(about 44°22’N), at a depth of about 20 m, contained nu-
merous specimens of Pontogeneia inermis (Krgyer) and a
_ single specimen of Calliopius laeviusculus (Krdyer). The other
samples, collected from lobster pots set at greater depths at
the edge of the continental shelf in the vicinity of Atlantis
Canyon, south of Nantucket Island, Massachusetts, and
almost due east of Philadelphia, Pennsylvania, contained speci-
mens of the new genus and species described below.
EusIRIDAE (SENSU BARNARD, 1972)
Dolebrotus, new genus
Antenna 1: slightly shorter than antenna 2, with small, apparently
unarticulated accessory flagellum. Labium: without inner lobes.
Mandible: 3rd segment of palp not widening distally, about *4 length
of 2nd segment. Maxilla 1: inner lobe with fully setose inner margin.
Maxilla 2: inner lobe with oblique row of setae. Pereopods 1 and 2
(gnathopods): subchelate; propus sublinear, elongate, especially in
pereopod 1; carpus not lobate, sublinear, elongate but shorter than
propus; coxa of pereopod 1 slightly narrowing distally. Pereopods 5-7:
basis and propus subequal in length. Uropod 3: rami subequal. Telson:
cleft to midlength.
Type-species: Dolobrotus mardeni, new species.
Etymology: From the Greek “dolos” (= “bait”) and “brotos” (=
“eating ). Gender, masculine.
Dolobrotus mardeni, new species
Figures 1-27
Material examined (all from lobster pots set in the vicinity of Atlantis
Canyon ):—Holotype, 7.0 mm ovigerous 9, (USNM 149127) and
several hundred paratypes (149216), 2-4 miles W of Atlantis Canyon,
39°52’N, 70°12’W, depth 170-256 m, received 16 February 1973. 10
paratypes from this collection have been deposited in the National Mu-
seum of Natural Sciences, Ottawa, Canada.—25 paratypes (USNM
142192), vicinity of Atlantis Canyon, exact locality unknown, depth
90 m, received 22 August 1972.—Several hundred paratypes (USNM
=>
Fics. 1-11. Dolobrotus mardeni. 1, head, lateral; 2, head, dorsal;
3, antenna 1, medial; 4, antenna 2; 5-7, epimera of pleonites 1-3; 8,
labrum; 9, left mandible; 10, incisor and lacinia of left mandible; 11,
incisor of right mandible.
A New Deep-water Lobster Bait Scavenger 131
132 Proceedings of the Biological Society of Washington
149217), northern edges of Atlantis Canyon, depth 75-165 m, received
4 May 1978.
Etymology: Named for Luis Marden, who first collected these amphi-
pods and brought them to my attention.
Diagnosis: With the characters of the genus.
Additional description: Length of largest adult 8.5 mm. Eyes: diffi-
cult to discern in preserved specimens, without dark pigment; roughly
oval in shape, with regular scattered ommatidia. Head: rostrum short,
triangular in dorsal view, rounded in lateral view; lateral lobe with
sloping dorsal margin and truncate vertical margin; anteroventral corner
smoothly rounded. Pereon: smooth dorsal profile. Pereonites 1 and 2
with anterior part of dorsum elevated above posterior part. Pleon:
epimera with posteroventral corners pointed; posterior margin above
point with shallow concavity in pleonite 2, with deeper concavity in
pleonite 3; pleonites 2 and 3 each with 3 spines along ventral margin,
pleonite 2 with 2 additional spines anterior and slightly dorsal to mar-
ginal spines.
Antenna 1: slightly shorter than antenna 2; 2nd peduncular segment
about half as long as Ist, distal margin produced medially into rounded
lobe; 3rd peduncular segment subequal in length to 2nd, but narrower;
accessory flagellum somewhat less than twice as long as wide, without
basal suture separating it from 3rd peduncular segment, bearing 4 apical
setae; flagellum with 19-26 segments. Antenna 2: about as long as
head and pereon combined; 4th peduncular segment produced into
rounded medial lobe; 5th peduncular segment subequal in length to
Ath; flagellum with about 22 segments.
Epistome: pointed above, rounded anteriorly, separated from labrum
by lateral notches. Labrum: rounded anteriorly; posterior margin
evenly rounded, armed with close-set spinules.
Left mandible: incisor with 7-8 teeth; lacinia with 5 teeth; spine row
with 6-7 spines. Right mandible: incisor with 8 teeth; spine row with
about 8 spines, 2 spines adjacent to incisor broad, truncate, resembling
noncuspidate lacinia. Labium: outer lobes densely setose; mandibular
processes rather narrow, sublinear. Maxilla 1: inner lobe with about 17
setae along medial margin and a much longer terminal seta; outer lobe
with 11 strongly pectinate terminal spines; palp with 11 terminal spines.
Maxilla 2: inner lobe with 3 robust apical spines; outer lobe with about
9 stout spines along medial margin; palp with moderately broad 2nd
segment and acutely produced distolateral corner of 3rd segment.
Pereopod 1: slender, subchelate, resembling that of Leptamphopus;
coxa slightly narrowing distally, with rounded distal margin; carpus
about %4 as long as propus, both with densely setose posterior margins,
their combined length subequal to that of basis. Pereopod 2: slender,
subchelate; basis about %4 longer than propus, widening distally; carpus
about “4 as long as propus and slightly broader, both with densely
setose posterior margins. Pereopods 3-4: basis slightly longer than
propus, nearly linear; dactyl with inner and outer facial setae distally.
A New Deep-water Lobster Bait Scavenger 133
Fics. 12-20. Dolobrotus mardeni. 12, labium; 13-15, maxilla 1:
13, inner lobe; 14, apex of outer lobe; 15, apex of palp; 16, maxilla 2,
inner lobe (setae omitted except those in oblique row); 17, maxilliped;
18, pereopod 1; 19, pereopod 2; 20, palm and dactyl of pereopod 2
(most setae omitted).
134 Proceedings of the Biological Society of Washington
Fics. 21-27. Dolobrotus mardeni. 21-24, pereopods 4—7; 25, uropod
1; 26, uropod 2; 27, uropod 3 and telson.
135
A New Deep-water Lobster Bait Scavenger
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Pereopods 5-7: P7 > P6 > P5; basis broadly expanded, slightly shorter
than propus; dactyls with inner and outer facial setae distally.
Uropod I: reaching distal end of uropod 3 and slightly beyond distal
end of uropod 2; protopod slightly longer than exopod, both margins
armed with spines, more numerous on lateral margin; endopod sublinear,
slightly longer than exopod, both margins armed with spines, apex bear-
ing 2 spines, lateral half as long as medial. Uropod 2: exopod similar
to uropod 1, but protopod subequal in length to exopod, only %4 length of
endopod; exopod with 2 spines on medial margin. Uropod 3: rami
lanceolate, with spines on both margins but not at apex. Telson: about
¥ longer than wide; incised to midlength; each half of apex with minute
indentation bearing spinule.
Relationships: Dolobrotus belongs to the family Eusiridae sensu
Barnard (1972). Barnard (1964) had previously expanded the Eusi-
ridae to incorporate the Pontogeneiidae. The addition of the Calliopiidae
in 1972 was anticipated by his statement (1964:57), “The Calliopiidae
are simply a mixture of eusirid and pontogeneiid types having the telsonic
lobes completely fused.” Bousfield (1973), however, did not accept
Barnard’s combining the Pontogeneiidae with the Eusiridae [Barnard’s
(1972) addition of the Calliopiidae appeared after Bousfield’s book was
in press]. My limited experience in gammaridean taxonomy deters me
from taking part in the argument; I will only point out that those
amphipod genera that appear to be closest to Dolobrotus include both
Calliopiidae (Bouvierella, Leptamphopus, Oradarea) and Pontogeneiidae
(Djerboa, Schraderia). A comparison of the principal features of all
these genera is given in Table 1.
Distribution of lobster bait scavengers: The 2 species of Lysianassidae
are panarctic species that occur also in Atlantic boreal waters. Orcho-
menella pinguis ranges south to North Carolina, but Anonyx sarsi only
reaches Rhode Island (Steele and Brunel, 1968; Bousfield, 1973). Both
are found on sandy bottoms, O. pinguis to depths of more than 100 m
and A. sarsi down to about 50 m. Two of the reported attacks on
trapped lobsters took place in Seal Cove, Grand Manan, New Brunswick,
and the depths were shallow. Templeman’s (1954) reported attacks by
an amphipod later identified as Orchomenella pinguis (Templeman,
1958) took place at 3 fathoms (5.5 m), and the predation by Anonyx
sarsi reported by Scarratt (1965) was on live lobsters that had been
loaded into crates and left floating overnight. Shave (1966) did not
give depths for his “sand fleas,’ identified by Roland Wigley as
Orchomenella and Anonyx.
Pontogeneia inermis is an Arctic boreal species that ranges south in
the western Atlantic to Long Island Sound. Bousfield (1973) char-
acterized it as “An essentially pelagic cold-water species; clings to sub-
merged plants and algae, from lower intertidal levels to more than 10
m.” I am not aware of any previous reports of its having been attached
to bait.
Dolobrotus mardeni appears to inhabit greater depths than the other
A New Deep-water Lobster Bait Scavenger 137
known amphipod bait scavengers and probably has not been a pest to
lobstermen until the recent development of setting pots at depths of
about 100 to 350 m. Thus far it is known only from the vicinity of
Atlantis Canyon (about 40°N, 70°10’W). A rather similar amphipod,
Leptamphopus sarsii Vanhoffen, was recently taken in baited traps near
Bergen, Norway, at 450 m, together with several species of Lysianassidae
(Vader, 1972).
Predation on lobsters by amphipod bait scavengers: “Sea fleas” are
a problem to lobster fishermen not only because they devour the bait
in lobster pots, but also because in some circumstances they will attack
trapped lobsters. Templeman (1954) noted that the shells of trapped
lobsters that had been eaten by amphipods always had a new appearance
and were not hard. He believed that these recently molted lobsters had
been wounded by hard-shelled lobsters in the same pot, and that
amphipods had been attracted to the wounds. Scarrett (1965) observed
a lobster placed in an aquarium with a number of Anonyx. After 6
hours the lobster had not been attacked, but had eaten about a dozen
amphipods. Scarrett concluded that healthy lobsters can resist attack
except when their movements are restricted. Lobster fishermen also
report that only wounded lobsters are attacked.
ACKNOWLEDGMENTS
I am grateful to Luis Marden and William Dion for collecting the
specimens used in this study, to J. L. Barnard for discussions on the
relationships of Dolobrotus mardeni, and to E. L. Bousfield for informa-
tion on published accounts of amphipod bait scavengers.
LITERATURE CITED
BARNARD, J. Laurens. 1964. Revision of some families, genera and
species of gammaridean Amphipoda. Crustaceana 7(1):49-74.
1972. Gammaridean Amphipoda of Australia, Part I. Smith-
sonian Contrib. Zool. 103:1-333.
BousFIELD, Epwarp L. 1973. Shallow-water gammaridean Amphipoda
of New England. Cornell University Press, Ithaca. xiv -++
312 pp.
BowMan, THoMAS E.. AND RAYMOND B. Manninc. 1972. Two arctic
bathyal crustaceans: the shrimp Bythocaris cryonesus new spe-
cies, and the amphipod Eurythenes gryllus, with in situ photo-
graphs from Ice Island T-3. Crustaceana 23(2):187-201, pl. 1.
HeEssLER, Ropert R., JoHN D. Isaacs, AND Eric L. Muus. 1972.
Giant amphipod from the abyssal Pacific Ocean. Science 175:
636-637.
Marpen, Luis. 1973. The American lobster, delectable cannibal.
Natl. Geogr. Mag. 143(4) :462-487.
PauL, ALLEN Z. 1973. Trapping and recovery of living deep-sea
amphipods from the Arctic Ocean floor. Deep-Sea Res. 20(3):
289-290.
138 Proceedings of the Biological Society of Washington
ScaRRATT, Davip J. 1965. Predation on lobsters (Homarus ameri-
canus) by Anonyx sp. (Crustacea, Amphipoda). J. Fish. Res.
Board Can. 22(4):1103-1104.
SHAVE, Paut A. 1966. Sand fleas are pests to our lobstermen. Fal-
mouth Enterprise (Massachusetts), 25 Nov. 1966, p. 6.
STEELE, DonaLp H. AND PIERRE BRUNEL. 1968. Amphipoda of the
Atlantic and Arctic coasts of North America: Anonyx (Lysi-
anassidae). J. Fish. Res. Board Can. 25(5):943—1060.
TEMPLEMAN, WILFRED. 1954. Lobsters eaten by sea-fleas. Prog. Rep.
Atlantic Coast Sta., Fish. Res. Board Can. No. 60:14—15.
1958. Lobsters eaten by sea-fleas. A further note. Prog.
Rep. Atlantic Coast Sta., Fish. Res. Board Can. No. 69:16.
VapvEerR, Wim. 1972. Notes on Norwegian marine Amphipoda. 5.
New records of Leptamphopus sarsii (Calliopiidae). Sarsia
50:25-28.
cs
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i
MypyeS ds
Vol. 87, No. 15, pp. 139-166 22 October 1974
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
EIGHT NEW ASHMUNELLAS FROM THE
SOUTHWESTERN UNITED STATES
(PULMONATA : POLYGYRIDAE)
By JosEpH VAGVOLGYI
Staten Island Community College,
Staten Island, New York 10301
INTRODUCTION
During the summers of 1967 and 1968 a search was made
for Ashmunellas in the low mountains of Arizona, New Mexico
and Texas. These mountains had been studied previously by
very few malacologists, presumably because their barren ap-
pearance discouraged potential collectors. The mountains are
deceptive, however, for many sustain land snail communities
in protected canyons, rock slides or near the peak areas.
Among the many species of land snails found, 8 forms of the
genus Ashmunella proved to be new to science. Description
of these forms and a critical reexamination of the taxonomy
of several related Ashmunella species follows.
ACKNOWLEDGMENTS
I am very grateful to Miss Cathy Corrigan, Messrs. Joe
Jarvis and Laszlo Simon who accompanied me on the field
trips; Dr. Joseph Rosewater, who made the collection of the
United States National Museum of Natural History (USNM)
available for comparative studies; Dr. Robert Robertson, who
allowed me to use the collection of the Academy of Natural
Sciences of Philadelphia (ANSP); Mrs. Morgan C. Rulon,
who helped me to obtain Pilsbry’s field notes; Dr. Kenneth J.
Boss, who sent specimens of Ashmunella bequaerti for com-
parison. I am also very grateful to Messrs. J. M. Cox, Jack
Hayter, Lotto Turner, Deryl York and many other ranchers
15—Proc. Bion. Soc. WasuH., Vou. 87, 1974 (139)
140 Proceedings of the Biological Society of Washington
who allowed collecting on their property, and the Headquar-
ters of the Army, the Fort Bliss Military Reservation and the
White Sand Missile Range who gave access to restricted
areas. Special thanks are due to the rangers who escorted us
in these areas and whose fabulous knowledge of the terrain
helped our work considerably. The work was supported by
Grant-in-Aid #89-2-B from the research foundation of the
State University of New York; Grant #4838, Penrose Fund,
from the American Philosophical Society.
‘TERMINOLOGY
The measurements and symbols used in this paper are
explained in Fig. 1.
DESCRIPTIONS
The descriptions start with that of the nominal subspecies
of Ashmunella kochi. Its description is necessary to provide
a comparative basis for the descriptions of three new sub-
species of A. kochi which follow, and to allow a more mean-
ingful discussion of their relationships.
Ashmunella kochi Clapp
Ashmunella kochi kochi Clapp
Plate I, Figures 1, la; Figure 4a
Ashmunella kochii Clapp, 1908: 77, pl. 8, figs. 1-3. Black Mountains,
San Andres Range, Doma Ana County, New Mexico. W. E. Koch.
This locality appears erroneous, see discussion of distribution below.
>
Fic. 1. Terms and standards of description. a, Shell in top view.
1, width; 2, reflected lip; 3, constriction or furrow behind lip; 4, ascend-
ing portion of last whorl; 5, whorl number; measured under dissecting
microscope, using circular scale divided into 10 parts, apex of the shell
pointing upward and being held in the center of scale; shell shown has
5.4 whorls; 6, embryonic shell; shell shown has 1.5 embryonic whorls;
7, granules, arranged in regressing radial rows; 8, striae, regressing
radial in direction. b, Frontal view, shell tilted. 9, lip swelling; 10, outer
lip; 11, outer lip tooth; 12, outer lamella of lower lip; 13, spout between
outer and inner lower lip lamellae; 14, lower lip; 15, inner lamella of lower
lip; 16, fulcrum or inner lamella, inside of shell; 17, umbilicus; 18, lower
parietal lamella; 19, upper parietal lamella; shell shown has medium
New Ashmunellas from southwestern U.S. 141
=
b 3
| G
J
thick lip swelling, wide and medium high outer lip tooth, medium large
lower lip lamellae, long fulcrum, medium large lower parietal lamella
and small upper parietal lamella; 20, upper lip. c, Frontal view, shell
held horizontally. 21, spire; 22, height of shell; 23, left shoulder of last
whorl; angularity of last whorl measured here, shell shown is rounded;
24, elevation of spire; (shell shown has 10° angle = flat) 7.5°-22.4°
Slat, 22.5°-37 4° = moderately elevated: d, Aperture shape. 25,
triangular; 26, auricular; 27, standing oval; 28, round; 29, rectangular;
30, angular; all but 29 round below. e, Angularity of last whorl, measured
at left shoulder. 31, rounded; 32, slightly angular; 33, angular; 34, keeled.
142 Proceedings of the Biological Society of Washington
George H. Clapp Collection, Pittsburgh, Pa. Lot labelled “Topotype,
original lot [probably paratype],” ANSP 97640, 2 specimens, examined.
Ashmunella kochi Clapp.—Pilsbry, 1940: 976, figs. 566a—c.
Ashmunella kochi amblya Pilsbry, 1940: 977, fig. 566d. Pine Spring
Canyon, Guadalupe Mountains, Culberson County, Texas. Type
ANSP 164659, 1 specimen, examined.
Definition: Ashmunella k. kochi as defined here includes the current
A. k. kochi and those populations of the current A. k. amblya which live
in the Guadalupe Mountains, but excludes those living in the San
Andres Range. The latter are considered a separate subspecies, A. k.
sanandresensis; see below.
Material examined: 7 samples, 1-30 specimens each, 77 specimens
altogether, from the ANSP and my own collection.
Description of shell: Size and Shape: Width 15.9-21.5 mm; spire
flat and only very slightly convex or plane in contour; apex sometimes
so depressed that it is level with right shoulder of last whorl when shell
is held horizontally; height of shell 5.6-7.9 mm, height-width ratio,
0.30—0.40; last whorl slightly angular or angular on left shoulder, slightly
or not at all ascending and then briefly but abruptly descending before
aperture, constricted by a deep furrow behind outer and lower lips.
Growth pattern: The 4.9-5.7 whorls first increase gradually, but last
whorl expands suddenly, particularly in larger specimens (Fig. 2b);
coiling usually looser than average, whorl—width ratio, .25-.34; embry-
onic shell has 1.4—1.6 whorls.
Umbilicus: Narrow at beginning but becoming suddenly wider in
last half of last whorl, diameter here reaching 4.0-7.1 mm; umbilicus—
width ratio varies from .24 to .34.
Aperture: Angular to triangular—auricular above, round below; its
plane slanted down and backward, somewhat concave when viewed from
side; outer and lower lips reflected; lip swelling medium thick to thick,
receded in upper lip, marginal in outer and lower, thus upper lip ap-
pears sharp, outer and lower, round.
Armature: Consists of 6 elements as follows: outer lip tooth, outer
and inner lower lip lamella, upper and lower parietal lamella, and
fulcrum (lamella deep inside of last whorl near umbilicus). Outer lip
tooth medium high and variable in width, moderately or deeply receded
and slanted down and inward; 2 lower lip lamellae medium large or
large, outer one usually larger than inner one, lip swelling usually pro-
truding between them to form a little spout; upper parietal lamella
present in traces or completely absent, lower large and long; fulcrum
moderately to well developed, % to % of a whorl in length.
Sculpture: First whorl of embryonic shell smooth except near suture
where it bears short radial striae; rest of embryonic shell covered with
fine striae, some of which break up into rows of fine granules; this
pattern persists for 1 or 2 more whorls, after that granules gradually dis-
appear and striae become more widely and irregularly spaced.
Reproductive anatomy: As characteristic of Ashmunella, the sperma-
New Ashmunellas from southwestern U.S. 143
theca is long, cylindrical, without a terminal widening; the epiphallus
is long, and the flagellum, vestigial (Fig. 4a). Three internal ridges
run through the entire length of the epiphallus (see cross sections).
All is in essential agreement with Pilsbry’s drawing of the genitalia of
A. k. amblya (1940: 916, from Pine Spring Canyon, Guadalupe Moun-
tains, Texas). However, in my specimen the penial retractor is only at-
tached to the penis and that region of the epiphallus next to it, whereas
in Pilsbry’s it is also attached to a point farther above on the epiphallus.
The difference may be due to individual variation.
Taxonomy: Ashmunella k. amblya of Pilsbry is combined with A. k.
kochi because their types are inseparable in shell features such as
width, height, umbilicus, whorl number, apertural characteristics, etc.
(Bigs. 2a, 2b).
Distribution and ecology: Ashmunella k. kochi is restricted to the
Guadalupe Mountains of Texas (Fig. 3). We found it in a very narrow
and deep branch of Pine Spring Canyon, on the northern slope of
Guadalupe Peak (8,751 ft.), at about 6,800-7,000 ft. of elevation, about
3 mi. west of Pine Spring. There was a nice stand of oak in the canyon.
In spite of the heavy rains that fell prior to our visit, we only could find
20 live adult specimens in 4—5 hours of search. The snails were hiding
under rocks and logs. Pilsbry described some of the localities he collected
at in the area as “dry, stony hills,” “terraced butte on dry canyon,” etc.
The type-locality of Ashmunella k. kochi reportedly lies in the Black
Mountains, San Andres Range, New Mexico. This locality appears
erroneous for the following reasons. First, the type-specimens of A. k.
kochi most closely match the types of A. k amblya, reportedly collected
in Pine Spring Canyon, Guadalupe Mountains, which appear authentic
because they match other populations collected in that same mountain
range. Second, populations of A. k. kochi other than the types do not
occur in the San Andres Range; they seem only to occur in the Guadalupe
Mountains. Third, in the San Andres Range, of which Black Mountains
are a part, another subspecies of A. k. kochi occurs, A. k. sanandresensis;
this subspecies differs significantly from the types of A. k. kochi. Fourth,
another subspecies, A. k. cornudasensis occurs in the Cornudas Mountains,
which lie halfway between the areas of the two subspecies in question
(Fig. 3), which fact makes it unlikely that A. k. kochi could occur in
identical forms at both ends of the range. All these considerations seem
to indicate to me that the types of A. k. kochi do, in fact, come from
the Guadalupe Mountains, not from the San Andres Range as accepted
today. But until further studies I leave the status quo unchanged.
Ashmunella kochi cornudasensis, new subspecies
Plate I, Figures 2—2a; Figure 4b
Holotype USNM 701251, deposited in the United States National
Museum of Natural History, Washington, D.C. Paratypes in the collec-
tion of the author. Type-locality: Wind Mountain, Cornudas Mountains,
Otero County, New Mexico.
144 Proceedings of the Biological Society of Washington
Material examined: 1 sample, 3 specimens altogether, from my own
collection.
Description of shell: Size and shape: Width of shell 17.9-18.2 mm;
spire flat or moderately elevated, very slightly convex or plane in out-
line; height 7.4-8.1 mm, height—width ratio 0.41-0.45; last whorl
angular at left shoulder, descending briefly before aperture, constricted
by a deep furrow behind outer and lower lips.
Growth pattern: The 5.3-5.4 whorls widen gradually; coiling is loose,
whorl—width ratio varies from 0.29 to 0.30; embryonic shell has
1.4—1.5 whorls.
Umbilicus: Very narrow at beginning, almost point-like, widening
suddenly in last part of last whorl; umbilical diameter here is 3.8-4.2
mm, umbilicus—width ratio, 0.21-0.23.
Aperture: Auricular or angular above, round below, its plane concave
and strongly slanting down and backward in side view; outer and lower
lips reflected; lip swelling medium thick, receded from edge of lip above
outer lip tooth, marginal below it, therefore former region appears
sharp, the latter, round.
Armature: Consists of same elements as in nominal subspecies; outer
lip tooth medium large, wide, receded and slanted down and inward;
2 lamellae on lower lip medium large, outer usually stronger than inner;
lip swelling forms a spout between them; lower parietal lamella medium
high and long, upper, very weak and short; they converge inward but
do not fuse at their inner end; fulcrum moderately large.
Sculpture: Embryonic whorls smooth except below suture where very
fine, short striae are visible; postembryonic whorls bear a fine striation.
Reproductive anatomy: Reproductive organs essentially as in Ash-
munella k. kochi (Fig. 4b) except that spermatheca is somewhat shorter
and penial retractor is attached to epiphallus both near penis and at a
point farther above; when the retractor contracts, it throws the lowest
section of the epiphallus into a loop. Also, a minor ridge is noticeable
in cross sections of the middle epiphallus beside the 3 major ridges, and
the penis has much thicker walls and smaller ridges than in nominal
subspecies.
Similarities and differential diagnosis: Ashmunella k. cornudasensis
closely resembles A. k. kochi in apertural features and armature. Its
higher spire, narrower umbilicus and angular last whorl clearly differ-
entiate it, however, from that subspecies (Table 1). In the angularity
of the last whorl it resembles A. b. mudgei; this appears to be a super-
ficial parallelism, however, because the growth pattern and the apertural
features are very different in the two forms.
Name: The name cornudasensis refers to the Cornudas Mountains
where the subspecies was found.
Distribution and ecology: Ashmunella k. cornudasensis is a geographic
isolate of A. kochi (Fig. 3). It is only known from Wind Mountain,
Cornudas Mountains, New Mexico, about 18 mi. north of Cornudas,
Hudspeth County, Texas. We collected on the northwestern slope, in a
New Ashmunellas from southwestern U.S. 145
small, very steep canyon, some hundred feet below the peak (the eleva-
tion of which is 7,280 ft.). The vegetation was rather sparse in the
habitat, and so were the snails. In several hours of search we found
only 3 live specimens; one was crawling on the underside of a boulder
we turned over, 2 others were copulating under another big rock. The
ground was still moist from storms a few days before, and we expected
to find more snails. It is possible that additional specimens would have
been found at slightly higher elevations.
Ashmunella kochi sanandresensis, new subspecies
Plate I, Figures 3-3a; Figure 4c
Holotype USNM 701252, deposited in the collection of the United
States National Museum of Natural History, Washington, D.C. Paratypes
in the collection of the author. Type-locality: Western slope of San
Andres Peak, San Andres Range, Dona Ana County, New Mexico.
Definition: Ashmunella k. sanandresensis includes those populations
of the former A. k. amblya which live in the San Andres Range, but
excludes those which live in the Guadalupe Mountains. The latter is
synonymized here with A. k. kochi. The reason for this procedure is
that the San Andres populations differ so much from the Guadalupe
populations (Fig. 2a; Table 1) that they have to be treated as a separate
taxonomic unit.
Material examined: 2 samples, 8-23 specimens each, 31 specimens
altogether, from the collection of the Academy of Natural Sciences of
Philadelphia and my own collection.
Description of shell: Size and shape: Width of shell 14.5-18.4 mm;
spire flat or sometimes moderately elevated, very slightly convex or
plane in contour; height 5.2-6.7 mm, height-width ratio, 0.34—-0.42;
last whorl slightly angular at left shoulder, gently ascending and then
briefly but abruptly descending before aperture, constricted behind outer
and lower lips.
Growth pattern: The 5.1-5.7 whorls increase gradually; coiling aver-
age in tightness, whorl-width ratio varies from 0.31—0.36; embryonic
shell consists of 1.5-1.6 whorls.
Umbilicus: Narrows at beginning but suddenly expanding in last part
of last whorl, reaching a diameter of 5.7-6.5 mm and an umbilicus-width
ratio of .30—.38; however sudden this expansion may appear, it is less
sudden than in the related species A. auriculata. This is shown by the
fact that the first part of the umbilical diameter (U1, Fig. 2c) is nearly
equal to the second part (U2), values for Ul being 2.2-3.4 mm and
U2, 2.4-3.4, whereas in A. auriculata U2 values are definitely greater
than U1 values.
Aperture: Rounded auricular or rounded angular above, round below,
its plane concave and slanting down and backward when viewed from
side; outer and lower lips reflected; lip swelling medium thick, receded
in region above outer lip tooth, marginal below it, hence former region
is sharp, latter, round.
146 Proceedings of the Biological Society of Washington
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New Ashmunellas from southwestern U.S.
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148 Proceedings of the Biological Society of Washington
Armature: Consists of same elements as in Ashmunella k. kochi; outer
lip tooth medium large, rather narrow, receded from margin of aperture,
slanted down and inward; lower lip lamellae medium large although
outer lamella usually slightly larger than inner; both protrude slightly
from aperture, and there is often a spout between them formed by a pro-
trusion of lower lip swelling; upper parietal lamella very weak and very
short, sometimes only in traces, while lower one is moderate or strong
and long; both converge inward but remain separate; fulcrum medium
large.
Sculpture: Embryonic whorls covered from suture to suture with
closely packed, fine striae many of which are broken up into long, trans-
verse granules; exception is apex of shell which is smooth; on postem-
bryonic whorls both striae and granules become more widely spaced,
and on last whorls the granules gradually disappear.
Reproductive anatomy: Ashmunella k. sanandresensis is like A. k.
cornudasensis in all features except that the lower section of the epiphallus
is wider and has a much wider lumen with 4 ridges (Fig. 4c). This is
remarkable since in shell features, especially in the height and the
angularity of the last whorl the subspecies is closer to the nominal sub-
species than to A. k. cornudasensis. It may be that the penis is contracted
to a different degree in the dissected specimens, and this makes the
difference.
Differential diagnosis: The distinguishing characteristics of the sub-
species are summarized in Table 1.
Name: The name sanandresensis refers to San Andres Range where
the subspecies occurs.
Distribution and ecology: Ashmunella k. sanandresensis is a geo-
graphic isolate of A. kochi (Fig. 3). It is known from 2 localities in
the San Andres Range, both on the western slope of San Andres Peak
(8,239 ft.), near Ropes Spring, 28 mi. northeast of Las Cruces, Dona
Ana County, New Mexico. Pilsbry and Ferriss first found the snails in
1922; they collected in a ravine southeast of the spring, rather deep under
stones; elevation was not given. I found the snails for the second time
in 1967 at a locality far above the spring, at the foot of a low, north
facing cliff, under small rocks, deep in the dirt, under brush oak; eleva-
tion was about 7,000 ft. The terrain generally was rocky, with sparse
vegetation. Despite the heavy storm of a few days before, I found very
few live specimens.
The specimens that Pilsbry and Ferriss collected are noticeably larger
than the ones I found (16.5 versus 15.2 mm mean sample width). The
reason for the size difference is not known.
Ashmunella kochi caballoensis, new subspecies
Plate I, Figures 4—4a; Figure 4d
Holotype and one paratype deposited in the collection on the United
States National Museum of Natural History, Washington, D.C., USNM
New Ashmunellas from southwestern U.S. 149
701253 and 706822, respectively. Other paratypes in the collection of
the author. Type-locality: Brushy Mountain, Caballo Mountains, Sierra
County, New Mexico.
Material examined: 1 sample, 30 specimens, from my collection.
Description of shell: Size and shape: Shell width 15.3-20.5 mm;
spire moderately elevated or flat, sometimes so flat that right shoulder
of last whorl is level with apex of shell when held horizontally; shell
height 6.0-8.2 mm, height-width ratio, 0.34—0.46; last whorl round or
slightly angular at left shoulder, at the most (as in the flattest specimens )
slightly ascending and then briefly but abruptly descending before
aperture, constricted by deep furrow behind outer and lower lips.
Growth pattern: The 5.2-5.7 whorls are evenly although fairly rapidly
expanding; whorl-width ratio 0.28—-0.34, coiling looser than average;
embryonic whorls number 1.5—1.7.
Umbilicus: Narrow at beginning, but expanding suddenly in last part
of last whorl reaching diameter of 4.0-7.0 mm; umbilicus-width ratio
varies from 0.26 to 0.34.
Aperture: Round, rounded-angular or rounded-auricular above, round
below; plane of aperture concave to varying degrees and slanting down
and backward when viewed from side; outer and lower lips reflected;
lip swelling medium thick or thick, receded in region above outer lip
tooth while marginal below it, thus lips appear sharp in former region,
round in latter.
Armature: Consists of 6 elements as usual in species; outer lip tooth
medium large, moderately wide or wide, receded and often slanted down
and inward; lower lip lamellae both large or very large with outer being
usually larger than inner lamella; lip swelling often forms a spout be-
tween them; lower parietal lamella medium large or large and long, upper,
small or very small and short, sometimes only present in traces; the two
parietal lamellae converge inward but fuse only exceptionally; fulcrum
may be large, small or totally absent.
Sculpture: Embryonic whorls smooth except below the suture where
they bear very fine, regularly arranged striae; rest of shell with a some-
what coarser and more irregular striation.
Reproductive anatomy: Agrees in all essential features with Ashmu-
nella k. cornudasensis (Fig. 4d); the differences are that one of the 3
internal ridges is slightly subdivided in the lower region of the epiphallus,
and that the penis has very thick walls and 6-7 small ridges.
Differential diagnosis: The distinguishing features of the subspecies
are summarized in Table 1.
Name: The name caballoensis refers to Caballo Mountains, the
type-locality of the taxon.
Distribution and ecology: Ashmunella k. caballoensis is a geographic
isolate of its species (Fig. 3). It occurs only on Brushy Mountain, Caballo
Mountains, New Mexico. We collected it on the north face just below
the peak, at an elevation of about 7,200-7,300 ft. The snails live under
the small, loose rocks and dirt that accumulated at the foot of the huge
150 Proceedings of the Biological Society of Washington
cliffs forming the peak. The vegetation was sparse with juniper, brush
oak, creosote, yucca, etc. being the more conspicuous elements. The soil
was still wet from the rains that fell a few days before our visit, yet we
only could find 18 live, adult specimens, most of them deep underground.
Empty shells were abundant.
Taxonomy of Ashmunella kochi: The species as defined here consists
of 4 subspecies, 3 of which are newly described, while the remaining
one, the nominal subspecies, is the result of the synonymization of 2
previously recognized taxa. Considering that the 4 taxa recognized as
valid in this paper are but geographic isolates of the same entity, each
with some morphological distinction, some investigators may wish to
rank them as separate full species. My reasons for treating them as con-
specific subspecies are 2. First, the 4 taxa in shell features differ from
one another less than any one of them does from other, related forms,
e.g. A. auriculata. Second, the differences in reproductive anatomy be-
tween them are minute.
Ashmunella auriculata, new species
Plate I, Figures 5—5a
Holotype USNM 701254, deposited in the collection of the United
States National Museum of Natural History, Washington, D.C. Paratypes
in the collection of the author. Type-locality: Boulder Canyon, Organ
Mountains, Dona Ana County, New Mexico.
Material examined: 1 sample, 13 specimens, from my own collection.
Description of shell: Size and shape: Width of shell 11.5-12.9
mm; spire flat, sometimes so much so that ascending portion of last whorl
may be higher than apex when shell is held horizontally; spire very
slightly convex or plane in outline; height of shell 4.3-5.0 mm, height-
width ratio 0.35-0.40; last whorl slightly angular at left shoulder,
ascending decidedly and then descending abruptly before aperture, con-
stricted from suture to suture behind lips.
Growth pattern: The 4.8-5.3 whorls increase slowly, gradually; coiling
moderately tight; whorl-width ratio ranges from 0.40 to 0.42; embryonic
shell consists of 1.5—1.6 whorls.
Umbilicus: Narrow at beginning but suddenly very greatly expanding
in last part of last whorl which renders it oval shaped, and makes diameter
of last part of umbilicus greater than that of first part (Fig. 2c, U2 and
U1 values); umbilical diameter 3.5-4.4 mm, U1 values 1.3-1.9 mm,
U2, 2.1-2.6 mm; umbilicus-width ratio 0.30-0.35
Aperture: Rounded-auricular or standing oval-auricular above, round
below, its plane concave in side view and slanted down and backward;
lips reflected all around; lip swelling medium thick, marginal or slightly
receded above outer lip tooth, marginal or protruding below it, hence lips
are sharp in former region, round or bulging in latter.
Armature: Composed of 5 elements: outer lip tooth, moderately
wide and high, sometimes bicuspid, slightly or deeply receded and
New Ashmunellas from southwestern U.S. JUS
slanted down and inward; outer and inner lamellae on lower lip, medium
large, or large, sharp blade protruding from aperture; often a spout
between them formed by protrusion of lower lip swelling; upper parietal
lamella short and weak, lower parietal lamella very strong and long;
they converge inward but do not fuse at their inner end. The fulcrum
which forms the sixth element in the armature of the related species
A. kochi is missing.
Reproductive anatomy: Unknown. There are no live specimens
available for study.
Similarities and differential diagnosis: Ashmunella auriculata closely
resembles A. kochi in apertural features. However, the high whorl-width
ratio, the pronounced ascension of the last whorl upon the penultimate
one, the suture-to-suture constriction behind the completely reflected
lips, the oval umbilicus, the standing oval-auricular aperture and the
absence of the fulcrum are more than enough to distinguish this species
from any of the 4 subspecies of A. kochi (Table 1).
Taxonomy: Considering that in apertural features Ashmunella auricu-
lata is similar to A. kochi, and that it is isolated but occurs near the
area of one of the subspecies of A. kochi, A. k. sanandresensis, some
malacologists may wish to consider it just another subspecies of that
species. As the great number of distinguishing features listed above
indicates, however, A. auriculata is more different from any subspecies
of A. kochi than the latter are from each other. Therefore, it seems best
to rank it as a full species. While the facts that the ascension of the
last whorl, the shape of the umbilicus and that of the aperture may be
correlated characters (and a slight ascension also is noticeable in some
specimens of A. kochi) weaken this conclusion somewhat, they do not
seem to invalidate it.
Name: The name auriculata refers to the shape of the aperture,
which is more auricular than in related species.
Distribution and ecology: Ashmunella auriculata is a geographic
isolate of the A. kochi group. To date it has been found only in Boulder
Canyon, in the southern part of Organ Mountains, about 14 mi. east of
Las Cruces, Dona Ana County, New Mexico (Fig. 3). The snails were
found in the narrow upper part of the canyon, at about 6,500-6,800 ft.
of elevation, under dirt and small rocks that accumulated at the foot
of the precipitous eastern wall of the canyon. The cliffs provided some
shelter, but otherwise the canyon was exposed to the scorching sun
and was very dry and barren.
The discovery of Ashmunella auriculata in Organ Mountains was sur-
prising since another species of Ashmunella, namely A. organensis of
Pilsbry, was already reported from there, and only a small area is involved.
I first assumed that some error was made in labelling the localities, and
checked both my collection and Pilsbry’s field book. I have found
some discrepancies and some data are missing from the latter; never-
theless, the localities appear correct. Thus it seems that 2 species of
Ashmunella occur in Organ Mountains.
152 Proceedings of the Biological Society of Washington
Ashmunella macromphala, new species
Plate I, Figure 7; Figure 5a
Holotype and 1 paratype deposited in the collection of the United
States National Museum of Natural History, Washington, D.C., USNM
701256 and 706823, respectively. Other paratypes in the collection of
the author. Type-locality: Cooks Peak, Luna County, New Mexico.
Material examined: 1 sample, 30 specimens, from my collection.
Description of shell: Size and shape: Width of shell 12.5-14.9 mm;
spire flat or sometimes moderately elevated, plane or slightly convex
in outlines; height of shell 5.4-6.5 mm, height-width ratio 0.40-0.51,
last whorl slightly angular or angular on left shoulder, briefly descending
before aperture, constricted by deep furrow behind lips in region of outer
lip tooth and below it.
Growth pattern: The 5.0-5.4 whorls are evenly expanding and moder-
ately tightly coiled; remarkably, the whorl number varies independently
of the width of the shell (Fig. 2a), whereas usually the two characters
are positively correlated: It is possible, however, that correlation would
become evident if more samples were studied; whorl-width ratio 0.35-
0.43; embryonic shell has 1.5—1.6 whorls.
Umbilicus: Wide from very beginning so that one can see underside
of first whorls; in last part of last whorl umbilicus widens even more,
reaching a diameter of 2.7—-4.3 mm, and an umbilicus-width ratio of
0.21-0.29.
Aperture: Round, rounded-angular or rounded-auricular above, round
below; its plane slanted down and backward, concave in side view,
sometimes pronouncedly so; lips reflected in region of outer lip tooth
and below it, thus paralleling extent of constriction behind lips; lip
swelling medium thick at most, receded above outer lip tooth, marginal
below it, therefore lip appears sharp in former region while round in latter.
Armature: Consists of 5 elements: outer lip tooth, medium large and
medium wide or wide, receded and slanted down and inward; outer
and inner lower lip lamellae both medium large although outer usually
somewhat larger than inner; lower lip often bears a spout-like pro-
tuberance between these two lamellae; fourth and fifth elements are
2 parietal lamellae which converge inward although remaining separate
along their entire length; upper parietal lamella very short and small,
sometimes only in traces; lower parietal lamella medium large and long.
Sculpture: Embryonic whorls completely covered with fine, closely
packed striae which sometimes break up into long transverse granules;
post embryonic whorls bear a similar sculpture except that striae are
somewhat coarser and more widely spaced.
Reproductive anatomy: The long epiphallus, the vestigial flagellum
and the long, cylindrical spermatheca are typical ashmunelloid features
(Fig. 5a). The inner structure of the epiphallus is characterized by 3
longitudinal ridges which run through its entire length. The distin-
guishing features are: the hermaphroditic gland is not subdivided; the
New Ashmunellas from southwestern U.S. 153
penial retractor is attached to a long section of the epiphallus above
the penis, thus contraction of the retractor muscle forces the epiphallus
into at least 2 loops; the penis sac is quite large in comparison to the
vagina, its wall is relatively thin and its lumen wide, with 2 major
and 7 or 8 minor internal ridges; the atrium is small.
Similarities and differential diagnosis: In regard to the aperture and
the armature, Ashmunella macromphala is very similar to A. kochi. Its
main distinguishing feature is the umbilicus which is wide throughout
whereas in the A. kochi group it is narrow at the beginning and wide
only in the last part of the last whorl. In the structure of the epiphallus,
too, an agreement exists between the 2 species. The large penis clearly
distinguishes A. macromphala from the A. kochi group. In fact, the shape
and structure of the penis somewhat resembles that of A. mogollonensis
(Pilsbry 1940, Fig. 524:3).
Taxonomy: Ashmunella macromphala is considered a full species be-
cause of the distinguishing features in its shell and in its reproductive
anatomy, and because of its isolated distribution. Its nearest relatives
appear to be the members of the A. kochi group, with which it shares
apertural features and epiphallic structure, and possibly A. mogollonen-
sis, with which it agrees in some features of the penis.
Name: The name macromphala refers to the most distinguishing
feature of the species, the large umbilicus.
Distribution and ecology: Ashmunella macromphala is only known
from Cooks Peak, New Mexico (Fig. 3). We collected the snails on the
very steep northern slope of the peak, at about 6,900-7,000 ft. of eleva-
tion, at the edge of 2 huge rock slides, from under the rocks and the
debris accumulated between them. Groups of oak bordered the rock
slides, providing food and shelter for the snails. The debris was dry, it
did not rain for weeks prior to our visit; yet we found 80-90 live adult
snails at one of the rock slides. At the other, which was located merely
a quarter of a mile to the east, we only found 20-30 empty shells.
Ashmunella animasensis, new species
Plate I, Figure 8
Holotype USNM 701282, deposited in the collection of the United
States National Museum of Natural History, Washington, D.C. Para-
type in the collection of the author. Type-locality: Animas Peak, Hidalgo
County, New Mexico.
Material examined: 1 sample, 2 specimens, from my own collection.
Description of shell: Size and shape: Width of shell 11.9-12.0 mm;
spire moderately elevated and slightly convex in contours; height of
Shell 5.4-6.0 mm, height-width ratio 0.45-0.50; last whorl slightly
angular on left shoulder, very slightly descending before aperture, con-
stricted by a deep furrow behind lips below right shoulder.
Growth pattern: The 6.1-6.2 whorls increase very slowly, gradually,
coiling very tight, whorl-width ratio in both shells 0.51; embryonic shell
consists of 1.4 whorls.
154 Proceedings of the Biological Society of Washington
Umbilicus: Narrow at beginning but suddenly expanding in last part
of last whorl, its diameter here reaches 2.3-2.4 mm; umbilicus-width
ratio 0.19-0.20.
Aperture: Rectangular or auricular-rectangular, its plane slanted down
and backward and hardly concave in side view; outer and lower lips
reflected; lip swelling medium thick, receded in upper lip, nearly marginal
in outer, and slightly protruding in lower; therefore upper lip is sharp,
the outer and lower, round.
Armature: Consists of 1 tooth in outer lip, 2 in lower, and 1 lamella
on parietal wall; outer lip tooth low but very wide, sometimes bifid,
slightly receded and slanted down and inward; outer lip tooth of lower
lip medium sized, inner lip tooth a small structure, both situated near
middle of lower lip; lip swelling protrudes between them like a spout;
parietal lamella moderately or strongly developed, straight; it corresponds
in position to lower parietal lamella of the Ashmunella kochi group. —
Sculpture: The 2 shells available for description are somewhat worn
and therefore fine details of the sculpture are difficult to discern; but
apparently the sculpture solely consists of fine striae both on the
embryonic and postembryonic whorls.
Reproductive anatomy: Not known.
Relationships and differential diagnosis: Ashmunella animasensis is
most similar to A. proxima of the Chiricahuas in size and coiling pattern,
but to A. pilsbryana of the Mogollons, in apertural features. The true
relations are probably to the Chiricahua species, because that range
lies only 25 mi. from Animas Peak, whereas the Mogollons are about
120 mi. away.
Ashmunella animasensis is not identical with A. tetrodon animorum
of Pilsbry and Ferriss, 1917.
Name: The name animasensis refers to Animas Peak, where the
species was discovered.
Distribution and ecology: Ashmunella animasensis is restricted to
Animas Peak, New Mexico (Fig. 3). It is a geographic isolate, pre-
sumably of A. proxima which lives in the nearby Chiricahua Mountains.
The collecting site was on the western side of Animas Peak (8,519 ft.),
somewhat below the summit, at an estimated elevation of 7,500-8,000 ft.
Pine, oak and juniper grew in abundance on the slope, indicating a fair
amount of precipitation. At the time of our visit, however, everything
was dry. We had to dig about 2 ft. in loose dirt and rock before we found
a couple of live adult specimens and a few empty shells.
Ashmunella salinasensis, new species
Plate I, Figure 9; Figure 5b
Holotype USNM 701257, deposited in the collection of the United
States National Museum of Natural History, Washington, D.C. Para-
types in the collection of the author. Type-locality: Salinas Peak,
Socorro County, New Mexico.
New Ashmunellas from southwestern U.S. 155
Material examined: 1 sample, 20 specimens, from my collection.
Description of shell: Size and shape: Width of shell 12.9-14.6
‘mm; spire flat or moderately elevated, its contours plane or very slightly
convex; height of shell 6.0-7.3 mm, height-width ratio 0.44-0.51; last
whorl slightly angular on left shoulder, very slightly descending before
aperture, constricted behind lips; constriction very shallow near origin
of upper lip (where latter fuses with parietal wall), considerably deeper
elsewhere.
Growth pattern: The shell is formed by 4.8-5.2 gradually to rather
rapidly expanding whorls; coiling moderately tight, whorl-width ratio
0.35-0.38; embryonic shell has 1.3-1.6 whorls.
Umbilicus: Narrow at beginning but suddenly expands in last part
of last whorl where its diameter is 2.8-3.8 mm; umbilicus—width ratio
varies between 0.22 and 0.28.
Aperture: Round, rounded-rectangular or rounded-auricular above,
round below, its plane slanted down and backward, exceptionally with
slight concavity in region of outer lip tooth when viewed from side;
upper lip slightly, the outer and lower, very much reflected; lip swelling
medium thick or thick, running near edge of lips except near origin of
upper lip where receded, hence lips swollen except in latter region
where sharp.
Armature: Consists of 3 elements: outer and lower lip teeth and
parietal lamella; outer lip tooth has degenerated into a barely recog-
nizable thickening of lip swelling or completely disappeared; lower lip
tooth is a minuscle protuberance or again may be completely absent;
parietal lamella, corresponding to lower parietal lamella of A. kochi,
may be very short and low, may be present in traces or may be com-
pletely absent; about 74 of specimens have no armature at all.
Sculpture: Embryonic whorls ornamented with fine striation except
on smooth apex; some striae may break up into rows of long, transverse
granules; this pattern persists for first 2 or 3 postembryonic whorls; there-
after moderately strong ribs replace striae, numbering 33-39 on last whorl.
Reproductive anatomy: Spermatheca long, cylindrical, without a
terminal sac, epiphallus long, and flagellum vestigial, as usual in
Ashmunella (Fig. 5b). Upper epiphallus characterized internally by
3 longitudinal ridges and several minute wrinkles, which features occur
in many congeners. However, spermatheca is unusually swollen (had the
specimen just mated?) and middle region of epiphallus has a section
about 3.7 mm long inflated to about 1% times normal diameter with
walls and 3 internal ridges being stretched out thin; particularly, this latter
feature is distinctive. Penis has 3 major, subdivided ridges and 4-6
minor ones.
Similarities and differential diagnosis: Ashmunella salinasensis is
very similar in shell features to the A. rhyssa group. It is distinguishable
from the latter by its smaller and flatter shell and much wider umbilicus.
Anatomically it is difficult to relate to any group; the inflation of the
middle epiphallus is a unique feature. It is fair to say that nothing in
156 Proceedings of the Biological Society of Washington
the anatomy contradicts the conclusion concerning its relationships
reached on conchological grounds.
Taxonomy: On the basis of its isolated distribution and moderate
degree of distinction in shell morphology, A. salinasensis could be ranked
either as a subspecies of A. rhyssa or as a separate species. Its uniqueness
in anatomical features lends support to the latter alternative.
Name: The name “salinasensis” refers to Salinas Peak where the
species was found.
Distribution and ecology: Ashmunella salinasensis is a geographic
isolate of the A. rhyssa group (Fig. 3). It occurs solely on Salinas Peak
(8,958 ft.), New Mexico, while the A. rhyssa group is widespread in
the Sierra Blanca Mountains, about 35 mi. to the east. The collecting
site was on the northern slope of the peak, near the summit, at about
8,500-8,800 ft. elevation. The road leading to the summit was met
here by the upper end of a huge rock slide. Several groups of oak trees
occurred at the edge and inside the rock slide, forming small green islands
on the otherwise barren mountainside. We found many live snails in
these patches, under rocks in the dirt and litter. We also collected at
another rock slide further down along the road, at about 7,200-7,800 ft.
elevation on the northwestern slope, but there found only Sonorella.
Ashmunella pasonis polygyroidea, new subspecies
Plate I, Figure 10
Holotype USNM 701225, deposited in the collection of the United
States National Museum of Natural History, Washington, D.C. Para-
types in the collection of the author. Type-locality: Franklin Mountain,
El Paso County, Texas.
Material examined: 1 sample, 3 specimens, from my collection.
Description of the shell: Size and shape: Width of shell 14.1-14.8
mm; spire nearly or completely flat, plane or very slightly convex in
outline; height of shell varies from 4.9 to 5.1 mm, height-width ratio
0.34-0.35; last whorl angular at left shoulder, ascending and then
briefly but very abruptly descending before aperture, with deep con-
striction behind outer and lower lips, with a shallow constriction or none
behind upper lip.
Growth pattern: The 5.0-5.1 whorls expand slowly, gradually; coiling
moderately tight, whorl-width ratio 0.34-0.36; embryonic shell consists
of 1.4-1.5 whorls.
Umbilicus: Very narrow at beginning, but expands rapidly in last part
of last whorl, reaching diameter of 4.3-4.6 mm; umbilicus-width ratio
varies from 0.30-0.32.
Aperture: Round or rounded-auricular above, round below, its plane
slanted down and backward, deeply concave in region of outer lip
tooth; upper lip moderately reflected, outer and lower ones more so;
lip swelling moderately thick or thick, receded above outer lip tooth,
marginal below.
New Ashmunellas from southwestern U.S. 157
Armature: Unique in that dentition resembles that of Ashmunella
kochi, while parietal features resemble those of A. pasonis. Thus, upper
lip toothless, outer lip bears a tooth, and lower lip bears 2 lamellae;
outer lip tooth medium large, medium wide, situated near right upper
corner of aperture, slightly receded and slanted down and inward; outer
lamella of lower lip medium large or large, inner lamella small or medium
large; both rather short, in 1 specimen so much so that they resemble
teeth (i.e. tubercles) more than lamellae (i.e. folds perpendicular to the
peristome); both situated far away from origin of lower lip (where lip
joins parietal wall), thus outer lamella is quite close to outer lip tooth;
parietal wall features a callus, roughly triangular in shape, slightly
raised at its frontal edge; lower boundary of callus formed by a very
large lamella which runs in a straight line from near origin of lower lip
across parietal wall towards outer lip tooth; upper boundary formed by
a slight thickening which fuses with lower lamella at its inner end; a
short fulcrum situated about 0.2 whorls behind aperture inside last
whorl. It must be emphasized here that a parietal callus is characteristic
of the related genus Polygyra, but it is extraordinary in Ashmunella.
Sculpture: The embryonic shells are worn in all specimens hence their
sculpture cannot be described with certainty; they appear to be smooth
with short striae near the sutures; the rest of the shell bears fine striation;
no granules or scales are discernible.
Reproductive anatomy: Not known.
Similarities and differential diagnosis: Ashmunella pasonis polygyroidea
resembles A. k. sanandresensis in size, shape, growth pattern, sculpture
and dentition, and A. p. pasonis (Plate I, Fig. 11) in parietal features.
This combination is unique and it distinguishes the subspecies from any
other form of Ashmunella.
Evolution and taxonomy: The newly recognized form presumably
originated by hybridization between A. p. pasonis and A. k. sanandresen-
sis, or some other subspecies of A. kochi. The former of the putative
parents lives in Franklin Mountain, the latter, in the San Andres Range,
near enough to allow hybridization. The assumption that hybridization
occurred is based on the fact that the new subspecies combines features
from each of its putative parents. This contention has not been sub-
stantiated by statistical analysis or breeding experiments. The small
number of shells available for study prevents a statistical treatment; the
breeding experiments so far undertaken have not yielded any results
since neither inter- nor intraspecific matings were observed (4 experi-
ments, each involving 19-32 specimens). Alternative explanations also
are possible.
Whether or not the newly described form originated through hy-
bridization, it is best to classify it as a subspecies of A. pasonis. The
latter is the only other form of Ashmunella, beside the taxon under dis-
cussion, which has a polygyroid parietal lamella, and the conspecific
status would bring out this fact. Furthermore, both of the
Ashmunella pasonis subspecies occur in the same mountain, separated by
158 Proceedings of the Biological Society of Washington
PLATE I. 1-la, Ashmunella k. kochi, Guadalupe Mountains, Texas,
« .82; 2-2a, Ashmunella k. cornudasensis, Cornudas Mountains, New
Mexico, x .82; 3-3a, Ashmunella k. sanandresensis, San Andres Range,
New Mexico, 3, x .82, 3a, x .88; 44a, Ashmunella k. caballoensis,
Caballo Mountains, New Mexico, 4, x .88, 4a, x .82; 5-5a, Ashmunella
auriculata, Organ Mountains, New Mexico, 5, < .93, 5a, X.88; 6, Un-
described Ashmunella from Chisos Mountains, Texas, < .82; 7, Ashmu-
nella macromphala, Cooks Peak, New Mexico, x .88; 8, Ashmunella
animasensis, holotype and paratype, Animas Peak, New Mexico, xX .88;
New Ashmunellas from southwestern U.S. 159
only a few miles from one another, and it appears most unlikely that 2
forms of Ashmunella could acquire a polygyroid parietal callus and occur
near one another without being related to one another. It seems less
logical to consider the new subspecies conspecific with the other putative
parent, A. k. sanandresensis, because it is more isolated geographically
from the latter form. In light of the above arguments, it also appears
illogical to rank it as a full species. Still another possibility is to leave
the subspecies, a presumed hybrid, without any taxonomic recognition.
However, the taxon does deserve some recognition because it has some
geographic isolation and thus an evolutionary potential with already
acquired morphological distinction.
The occurrence of a parietal callus in both Ashmunella pasonis
polygyroidea and the analogous A. p. pasonis poses another evolutionary
problem. As it has been pointed out earlier, a parietal callus is not known
in any other form of Ashmunella while it is characteristic of the related
genus Polygyra. It may be that intergeneric hybridization is involved,
or that independent evolution brought about this phenomenon; this
problem is not solved at the present time. It may be added that A. p.
pasonis resembles Polygyra in the location of the lip teeth and the de-
velopment of the whole peristome region as well, which facts may lend
some more credence to the assumption of intergeneric hybridization.
Name: The name polygyroidea has been chosen to express the
similarity of the subspecies to the genus Polygyra in apertural features,
especially in regard to the parietal callus.
Distribution and ecology: Ashmunella pasonis polygyroidea is only
known from Franklin Mountain, just north of El Paso, Texas (Fig. 3).
We collected in a large canyon on the east side of the mountain, above
Cottonwood Spring, at an elevation of about 6,000 ft., under the rocks
of a rock slide on the north-facing slopes; the vegetation was very
sparse. We only found a few empty shells.
Reproductive anatomy of Ashmunella bequaerti, mudgei and A. p.
pasonis: These taxa are not new to science. Cheatum described the
former in 1971 as a full species, A. mudgei; Drake described the latter
in 1951 also as a full species, Polygyra pasonis. Neither author described
the anatomy of the reproductive organs, however.
Reproductive anatomy of Ashmunella b. mudgei: The spermatheca is
cylindrical, the flagellum very small, attached, and the epiphallus, long
as usual in Ashmunella (Fig. 5c). The upper portion of the epiphallus
has 1 major and 7-9 minor ridges, which pattern is rather unusual. The
<
9, Ashmunella salinasensis, Salinas Peak, New Mexico, x .88; 10, Ash-
munella pasonis polygyroidea, Franklin Mountain, Texas, x .82; 11, Ash-
munella p. pasonis, Franklin Mountain, Texas, < .82; 12, Ashmunella b.
bequaerti, Davis Mountains, Texas, < .93; 13, Ashmunella b. mudgei,
Davis Mountains, Texas, x 1.1.
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middle epiphallus has 3 thin, large ridges and 6-8 small ones; the penis
has 1 very large ridge, 1 intermediate in size and about 10 very small ones.
Reproductive anatomy of Ashmunella p. pasonis: This taxon was
originally described as a Polygyra, based on shell features, which indeed
are very much more those of a Polygyra than an Ashmunella (Drake,
1951). Soon after its description, the taxon was transferred to the
genus Ashmunella by Drake (1952) who cited Wendell O. Gregg’s
observation that anatomically the taxon belongs to Ashmunella. Gregg’s
observations have not been published, however, thus the position of the
taxon has remained somewhat uncertain. As the description below will
show, the taxon indeed belongs to Ashmunella.
The spermatheca, the flagellum and the epiphallus are as charac-
teristic in Ashmunella (Fig. 5d). The upper epiphallus shows 3 major
ridges and several wrinkles in cross section, which is also a common
feature in the genus. The middle epiphallus has, however, 2 of the
major ridges subdivided and has in addition a small ridge. The penis
is very large, blunt, and horseshoe-shaped.
The conspecificity of Ashmunella bequaerti and A. mudgei: The two
forms in question were originally described as separate species (Clench
and Miller, 1966; Cheatum and Fullington, 1971). Considering the
great degree of morphological difference between them, their at least
<
Fic. 2. Variation of width, whorl number, umbilicus and width of
last whorl in 4 species of Ashmunella. a, Scatter diagram of variation in
width (W) vs. whorl number (Wh) in Ashmunella kochi, A. auriculata,
A. macromphala and A. bequaerti. Symbols as explained in diagram;
horizontal lines indicate range of variation; one dot may represent more
than one specimen. Note how close the type of A. k. amblya is to those
of A. k. kochi; this is the reason for synonymizing it with the latter.
Also note the lack of correlation between width and whorl number in
A. macromphala. b, Scatter diagram of variation in width of last whorl
(L) vs. width of shell (W) in A. k. kochi, A. k. sanandresensis and A. k.
caballoensis. Symbols as above, except that a common sign is used for
specimens of A. k. sanandresensis. Note that while there is an overlap
between the smaller specimens of the nominal subspecies and specimens
of the other 2 taxa, the larger specimens of A. k. kochi have a relatively
wider last whorl (i.e. the regression curve of A. k. kochi is steeper than
that of the 2 other taxa). The great size of these specimens is due to
the disproportionately wide last whorl not to more whorls (cf. Fig. 2a).
c, Scatter diagram of variation in diameter of first and last part of
umbilicus (U1 vs. U2) in A. k. sanandresensis and A. auriculata.
Measurements as explained on the diagram; symbols as explained above.
The more the umbilicus expands in the last part of the last whorl, the
greater the U2 value becomes relative to Ul, as in A. auriculata.
162 Proceedings of the Biological Society of Washington
Sierra Blanca Mts.
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Fic. 3. Distribution of Ashmunellas described or discussed in text.
Symbols as shown in figure. Abbreviations: A., Alamogordo; C., Carls-
bad; CO., Cornudas; D., Deming; E.P., El Paso; L.C., Las Cruces;
R., Roswell; S.C., Silver City; T.C., Truth or Consequences; V.H., Van
Horn; other abbreviations are of states. Note: not all mountain ranges
are shown in area.
partial geographic isolation, and the lack of evidence for interbreeding,
this procedure was logical. The discovery of a shell in the USNM collec-
tion (USNM 251921) has, however, changed this picture. This shell
(Plate I, Fig. 6) contains a mixture of the characteristics of the two
forms under discussion thus bridging the gap between them. Because
of practical reasons, formal taxonomic recognition is not extended to
this shell (or to the taxon of which it is a sample); but a somewhat
shortened description of the properties of this shell is presented here.
>
Fic. 4. Anatomy of reproductive organs of Ashmunellas described
or discussed in text. a, Ashmunella k. kochi, Guadalupe Mountains,
Texas; b, Ashmunella k. cornudasensis, Cornudas Mountains, New Mex-
ico; c, Ashmunella k. sanandresensis, San Andres Range, New Mexico;
d, Ashmunella k. caballoensis, Caballo Mountains, New Mexico. Insets
New Ashmunellas from southwestern U.S. 163
show sections of the epiphallus and penis sac made where indicated by
heavy lines; additional inset to Fig. 4b shows enlargement of the talon,
to Fig. 4c, location of the hermaphroditic gland in situ. 1, hermaphroditic
gland; 2, hermaphroditic duct; 3, albumen gland; 4, talon; 5, uterus;
6, oviduct; 7, spermatheca; 8, vagina; 9, prostate; 10, vas deferens; 11,
flagellum; 12, epiphallus; 13, penis sac; 14, atrium; 15, penis; 16, penis
retractor; 17, genital orifice; 18, mantle; 19, foot; 20, eye. Note: the
insets were drawn under 20 X magnification; their dimensions are pro-
portionate in relation to one another, but not to the rest of the diagrams.
164 Proceedings of the Biological Society of Washington
Fic. 5. Anatomy of reproductive organs of Ashmunellas described
or discussed in text. a, Ashmunella macromphala, Cooks Peak, New
Mexico; b, Ashmunella salinasensis, Salinas Peak, New Mexico; ec,
Ashmunella b. mudgei, Sawtooth Mountain, Davis Mountains, Texas;
d, Ashmunella p. pasonis, western side of Franklin Mountain, Texas.
Insets show sections of the epiphallus, penis sac and penis made where
indicated by heavy lines. Additional inset to Fig. 5a shows the soft body,
New Ashmunellas from southwestern U.S. 165
Width of shell 13.9 mm; spire flat and convex in outline; height 5.7
mm; last whorl keeled on left shoulder; whorl number 6.0, coiling tight.
Umbilicus wide at beginning so that underside of first whorls can be
seen from below; umbilical diameter in last whorl 3.4 mm.
Aperture angular above, round below; upper lip slightly, outer and
lower, widely reflected; lip swelling moderately thick; outer lip tooth
high and very wide, outer lamella of lower lip large, inner lamella,
medium large; upper parietal lamella very thin and long, lower lamella
strong and long.
The sculpture can only be tentatively described because the shell of
the only available specimen is somewhat worn; the embryonic whorls seem
to have only granules, packed tightly, the first postembryonic whorls
bear both granules and striae, and the last whorl, only striae.
As can be seen from the foregoing description, the shell in question
is similar to A. mudgei (Plate I, Fig. 13) in shape, sculpture and growth
pattern (Fig. 2a). It does not have, however, the exceedingly heavy lip
swelling and lower parietal lamella characteristic of that form, its um-
bilicus is wide at the beginning whereas that of A. mudgei is narrow, and
it is somewhat smaller than the latter. At the same time, this shell in
question agrees with A. bequaerti (Plate I, Fig. 12) in the formation of
the umbilicus, but lacks the more compressed, carinated shape, more
angular aperture, very widely separated parietal lamellae and_pre-
dominantly granular sculpture of that form.
As to the locality of the shell in question, the museum label reads:
“Chisos Mts., Texas, V. Bailey.” This certainly is possible but it has to
be pointed out that Chisos Mountains lie outside the known area of
distribution of the genus and that several collectors who worked in the
Chisos Range have failed to find any Ashmunella there. Thus this locality
report may be in error. It is possible though that future collecting will
disclose the species in the Chisos Range. This range is sizeable and it
provides a variety of habitats some of which appear suitable for Ash-
munella. (The range lies in the Big Bend National Park, Brewster County,
Texas; highest point is Emory Peak, 7,777 ft.)
Whether this shell in question comes from Chisos Range or from
Davis Mountains, where both A. bequaerti and A. mudgei occur, its
significance is in the fact that it provides a mixture of the morphological
characters of these two species and thereby it indicates their conspecificity.
<
with the shell broken away and the apical 24% whorls removed to expose
the hermaphroditic gland in situ; to Fig. 5b, the intact soft body with the
shell broken away, and the position of the albumen gland and the tip
of the epiphallus with the flagellum in situ; to Fig. 5c, the atrium,
partially and completely opened; and to Fig. 5d, the tip of the penis as
seen from the genital orifice.
166 Proceedings of the Biological Society of Washington
Therefore, I propose that their taxonomic rank be changed from species
to subspecies, and their names be A. b. bequaerti and A. b. mudgei.
LITERATURE CITED
Cueatum, E. P. 1971. A new species of Ashmunella from the Davis
Mountains in West Texas. Nautilus 84(3):107—109.
CuHEATUM, E. P. ann R. W. Fuuuincron. 1971. The aquatic and land
Mollusca of Texas, Part I. The recent and Pleistocene members
of the gastropod family Polygyridae in Texas. Bull. Dallas Mus.
Nat. Hist. 1: vi + 74 pp.
Ciapp, Georce H. 1908. New land shells from Arizona and New
Mexico. Nautilus 22:76-78.
CLENCH, W. J. AND W. B. Minter. 1966. A new species of Ashmunella
from West Texas (Mollusca : Pulmonata). Breviora 244:1-6.
Drake, R. J. 1951. A new species of Polygyra from West Texas.
Revista de la Sociedad Malacologica “Carlos de la Torre,’
Habana, 8:44—46.
. 1952. Change in genus for Polygyra pasonis. Ibid. 9:30.
Pinspry, H. A. 1936. Land shells from Texas and New Mexico.
Nautilus 49:100—102.
—. 1940. Land Mollusca of North America (North of Mexico).
Vol. I, Part 2. Acad. Nat. Sci. Philadelphia, Monograph
No. 3; vi + 575-994 + ix pp.
Pitspry, H. A. anp E. P. CHeatum. 1951. Land snails from the
Guadalupe Range, Texas. Nautilus 64:87-90.
Pinspry, H. A. AND J. H. Ferriss. 1917. Mollusca of the southwestern
states, VIII: The Black Range, New Mexico. Proc. Acad. Nat.
Sci., Philadelphia, 69:83-107.
Vol. 87, No. 16, pp. 167-174 22 October 1974
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
A NEW TRICOLOR LEPTOTYPHLOPS
(REPTILIA: SERPENTES) FROM PERU
By Brautio R. OreyAS-MrrRANDA AND GEORGE R. ZuG
Division of Reptiles and Amphibians, National Museum
of Natural History, Washington, D.C. 20560
Two Peruvian species of Leptotyphlops possess color pat-
terns of red, yellowish white, and black. L. rubrolineatus
(Werner, 1901:6) has a body pattern of alternating red and
black longitudinal stripes; yellow pigment is confined to the
rostrum, chin and throat, and tail. L. teaguei ( Orejas-Miranda,
1964:4) has a body pattern of red, black, and yellow longi-
tudinal stripes. Black stripes predominate with a thin mid-
dorsal black stripe flanked on each side by an equally thin
black stripe. These three stripes lie on a red field bordered
by dorsolateral black stripes and yellow lateral stripes; ventro-
laterally and ventrally the body is black. Recently one of us
(B.O-M.) discovered in the Museo de Historia Natural “Javier
Prado” (MHNJP) three Leptotyphlops specimens with a tri-
color pattern similar to that of L. teaguei. Scutellation differ-
ences indicate that these specimens represent a distinct species.
Leptotyphlops tricolor, new species
Holotype: MHNJP 0669, Peru: Ancash Department, Huaylas Province,
Huaylas District: Yunca Pampa, (2700 m), collected by Acacio Ramos
on 24 February 1966.
Paratypes: MHNJP 0670 and USNM 195853, same data as holotype.
Diagnosis: A Leptotyphlops of the tesselatus species group (Orejas-
Miranda, 1964:4) characterized by broad contact between supraoculars
and first pair of labials, differing from other members of the group
except L. teaguei by its tricolor pattern of brick red, black, and cream
longitudinal stripes, and differing from L. teaguei by absence of three
narrow middorsal black stripes, presence of a cream colored mental
region bisected by black stripe, and a higher number of dorsal scales
(more than 300).
16—Proc. Briou. Soc. WasH., VoL. 87, 1974 (167)
168 Proceedings of the Biological Society of Washington
Fic. 1. Leptotyphlops tricolor (MHNJP 0669). A, B, C, dorsal,
lateral, and ventral views of head, respectively; D, lateral view of tail;
E, ventral view of cloacal region; F, dorsal view at midbody. Heavy
stippling, black; light stippling, brick red; no stippling, cream white.
Description of the holotype: An adult male with total length of 295
mm and tail length of 14 mm; total body length/midbody diameter 60;
head slightly depressed with snout projecting a third of entire head
length beyond lower jaw; body subcylindrical, slightly tapered cranially
and caudally; tail tip with spine mucronate. Body tricolor, a broad, dorsal,
red stripe, a pair of narrow dorsolateral black stripes, a pair of lateral
cream stripes, and a black venter (Figure 1).
Rostral triangular, dorsal apex not extending to transverse line between
anterior border of eyes; nasal completely divided with naris in center of
suture, supranasal longer than infranasal (approximately two times);
A new Leptotyphlops from Peru 169
supralabial border formed by rostral, infranasal, anterior labial, ocular
and posterior labial; rostral, ocular and posterior labial of subequal width
at supralabial border; infranasal and anterior labial subequal, their width
at least half that of former three scales; anterior labial single, two and
a half times as high as wide, in contact dorsally at level of eye with
supraocular; ocular one and a half times as high as wide; eye at level
of maximum width of ocular and in anterodorsal half of ocular without
reaching nasal border; posterior labial trapezoidal, widest at supralabial
border; prefrontal, frontal, interparietal and interoccipital subequal and
slightly smaller than more posterior middorsal scales; large supranasal
and supraocular subequal and three quarters the size of the subequal
parietal and occipital; eye in contact with supraocular-ocular suture;
mental followed by four sublabials on each side; first pair of sublabials
separated by a postmental.
Middorsal scales 303 plus rostal and caudal spine; body covered with
14 scale rows, reducing to 12 rows at anal plate and to 10 rows at the
level of third subcaudals; anal plate enlarged and triangular with apex
free posteriorly, anteriorly bordered by five rows of scales; 19 subcaudal
scales.
Color pattern in alcohol: The body pattern consists of six longitudinal
stripes. The middorsal brick red stripe occupies four scale rows, the
middle three rows and half of each the two adjacent scale rows. The
dorsolateral black stripes cover the halves of each scale on two adjacent
rows. The lateral cream stripes lie on half of one scale row and entirely
on the next adjacent ventral row. The ventral black stripe covers the
five ventralmost scale rows.
The stripes lose their regularity on the head and tail. The wide
ventral stripe ends abruptly on the neck; the throat has a few black
spots, and a narrow black stripe extends from these spots to the mental.
The mental is cream, although the dorsal halves of the first two pairs of
sublabials are black. The dorsolateral stripes gradually angle dorsally
as they approach the head, so that by the tenth scale row they lie over
a single scale row rather than two halves. Also at this level, the dorso-
lateral stripe begins to widen medially, first as a few scattered black
spots and cranially as solid black scales. On the head, this dorsolateral
black stripe covers most of the occipital, parietal, and supraocular scales,
dips ventrally onto the ocular to enclose the eye, and extends anteriorly
across the dorsal tip of anterior supralabial onto the ventral half of the
supranasal. The cream-colored rostral is immaculate except for a faint
black mustache at the labial border. The prefrontal and frontal bear
medial smudgelike black spots. The dorsal stripe begins to change from
cream to red at the fifth middorsal scale, and the change is complete by
the tenth middorsal scale.
Caudally, the ventral black stripe ends abruptly on the enlarged pre-
anal scale; thus the ventral half of the tail including the caudal spine
is cream colored. The dorsolateral black stripes converge and fuse
dorsally above the spine. Black pigment has invaded the dorsal red
170 Proceedings of the Biological Society of Washington
Fic. 2. Distribution of Leptotyphlops tenellus.
stripe to produce three faint black stripes that fade out as they approach
the base of the tail.
Variation: Both paratypes are smaller than the holotype, a total length
of 269 mm and 222 mm, tail length of 12 mm and 11 mm for the
MHNJP and USNM specimens, respectively. The midbody diameter
into total body length is 89 and 71, respectively. They possess 302 and
310 middorsal scales plus rostral and spine, and 18 and 21 subcaudals,
respectively. The number of scale rows around body and tail is the same
for all three specimens. Similarly the shape, position, and number of
head scales is constant for all specimens. The only deviation occurs in
the USNM specimen in which the postmental is enlarged to the right
so that the second sublabial touches it.
The color patterns of the paratypes are similar to that of the holotype;
differences are discernible only by close examination. The MHNJP
specimen was preparing to shed when it was preserved, so color bright-
ness is dulled; however the three narrow black stripes in the middorsal
red field appear more prominently on the tail than on the body
and also appear for a short distance in the neck region. On both of
the paratypes the black pigment of the venter extends across the vent
as a midventral stripe on the first one or two subcaudals. The black chin
and throat stripe are continuous with the ventral body stripe and tend to
A new Leptotyphlops from Peru 171
wn: 4
10°
| | | | i {
80° 78° 76°
Fic. 3. Distribution of Peruvian members of tesselatus species group.
L. rufidorsus is intermediate between this group and albifrons group.
Open circle, rubrolineatus; solid circle, tricolor; open square, tesselatus;
solid square, rufidorsus; triangle, teaguei; shaded area, altitude greater
than 2500 m.
be wider. The black rostral mustache of the USNM paratype is pro-
portionately larger than that of the holotype and continuous across the
infranasal with the dorsolateral stripe.
Discussion: Members of the tesselatus species group share: a supra-
172 Proceedings of the Biological Society of Washington
labial border of rostral, infranasal, anterior supralabial, ocular and pos-
terior supralabial, an enlarged supraocular in broad contact with the
anterior supralabial thereby preventing ocular-nasal contact, and ten
scale rows around the middle of the tail. Currently five species comprise
this group: L. rubrolineatus, L. teaguei, L. tenellus, L. tesselatus, and
L. tricolor. L. tenellus is widespread in northern South America (Figure
2), whereas the other four are confined to western Peru (Figure 3)
and generally to the immediate vicinity of their type localities.
L. rufidorsus (Taylor, 1940:532) is not included in the tesselatus
group because the holotype has an asymmetry of left and right supra-
oculars, and the second known specimen (Schmidt and Walker, 1943)
possesses ocular-nasal contact, albeit narrow (Orejas-Miranda, 1964).
However, the enlargement of the supraocular reduces the amount of
ocular-nasal contact in rufidorsus and suggests that rufidorsus is inter-
mediate between the albifrons and tesselatus groups. These two groups
are similar in most characteristics but differ primarily by the broad
ocular-nasal contact of the albifrons group.
L. rufidorsus and three of the tesselatus group species (rubrolineatus,
teaguei and tricolor) share a striped pattern with a red dorsal stripe.
The reddish stripe occurs only in these Peruvian species. Because the
four species are related, their sharing of a similar color pattern is not
unexpected. But why is the dorsum red, when red does not appear in
any other leptotyphlopid groups, and why is this color pattern restricted
to western Peru? The three colors, red, cream and black, occur either
together or in combinations of two in many fossorial snakes; however,
these colors are usually in transverse bands or blotches and are considered
to be aposematic or mimetic patterns. The pattern of these leptotyphlo-
pids is possibly mimetic. R. Thomas (in litt.) suggested that some
Leptotyphlops bear a strong resemblance to small millipeds; therefore
mimicry is a possibility.
If the tesselatus members prove to be polymorphic, teaguei may be
the red-striped morph of tenellus. In spite of the strong similarity in
color pattern to teaguei, tricolor has approximately 60 to 70 more mid-
dorsal scales, a number which does not support a close relationship.
Likewise tricolor has 40 to 50 more middorsals than either rufidorsus or
tesselatus and is probably not closely related to them. L. tricolor is a
distinctive new leptotyphlopid in the Peruvian fauna.
Key TO SPECIES OF THE TESSELATUS GROUP
1. A bicolor pattern of cream white and dark brown; dorsal and
ventral coloration pattern identical ~~ 9 Se Eee 2
A tricolor pattern of cream white, brick red and black; dorsal and
ventral coloration dissimilar 2.) 28
2. Middorsal scales more than 250; generally small, body diameter
55 times or more into total body length = tesselatus
Middorsal scales less than 250; generally large, body diameter
less than 49 times into total body length _.. tenellus
A new Leptotyphlops from Peru 173
3. Body bicolor, a black venter bordered above by a red dorsum;
cream white areas confined to head and tail __.____. rubrolineatus
Body tricolor, a black venter separated from a red dorsum by a
cream white lateral, and a black dorsolateral stripe on each side _ 4
4, Middorsal scales less than 250; five black dorsal stripes, the middle
three thin and enclosed within dorsal red stripe; throat com-
Pleve lypilol icky eed La Pik oe Us ee WI a Ss teaguei
Middorsal scales more than 300; dorsal red stripe bordered by
dorsolateral black stripe, three middle stripes absent; throat cream
white with a midventral black stripe _...___/_»_____ tricolor
ACKNOWLEDGMENTS
We wish to thank Dr. Ramon Ferreyra and Ms. Nelly Carrillo de
Espinoza for allowing us to examine these specimens from the Museo de
Historia Natural “Javier Prado” (MHNJP), Lima Peru, and for per-
mitting us to retain one of the types in the National Museum of Natural
History (USNM). Dr. Albert Schwartz and Mr. Richard Thomas pro-
vided critical reviews of the manuscript. Mr. George Steyskal informed
us of the zoological nomenclature committee’s decision to consider all
generic names ending in -ops as masculine (1972, Bull. Zool. Nomencl.
29 (A177).
LITERATURE CITED
OrEyAS-Miranpa, R. 1964. Dos nuevos Leptotyphlopidae de Sur
America. Com. Zool. Mus. Hist. Nat. Montevideo 8(103):1-7.
Scumipt, K. P. anp W. F. Warxer. 1943. Snakes of the Peruvian
coastal region. Zool. Ser. Field Mus. Nat. Hist. 24(27):297-
324.
Taytor, FE. H. 1940. Herpetological miscellany. Univ. Kansas Sci.
Bull. 26( 15 ):489-549.
WERNER, F. 1901. Reptilien und Batrachier aus Peru und Bolivien.
Abh. Ber. konig. zool. anthr.-ethn. Mus. Dresden (1900/01)
92) a1
174 Proceedings of the Biological Society of Washington
Vol. 87, No. 17, pp. 175-184 22 October 1974
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
OCCURRENCE OF THE ATLANTIC MALDANID
ASYCHIS ELONGATA (ANNELIDA, POLYCHAETA )
IN SAN FRANCISCO BAY, WITH
COMMENTS ON ITS SYNONYMY
By Wiu1aM J. Licut
California State University, San Francisco, San Francisco,
California 94132, and Department of Invertebrate Zoology,
California Academy of Sciences, San Francisco,
California 94118
A maldanid polychaete of the genus Asychis Kinberg,
commonly found in San Francisco Bay, California, has pre-
viously been misidentified in the literature (Berkeley and
Berkeley, 1960 and Hartman, 1969). Based on numerous,
recently collected specimens, and by comparison with extensive
material on loan from the National Museum of Natural History,
the specimens can now be correctly identified as Asychis
elongata (Verrill), previously known only from the Atlantic
coast of the United States. The synonymy is discussed below.
Asychis elongata may have been introduced into San Francisco
Bay through shipments of the Virginia oyster, Crassostrea
virginica (Gmelin). It is now a conspicuous element of the
benthos throughout the bay.
I wish to thank Dr. Curtis Newcombe, San Francisco Bay
Marine Research Center (MRC), and Dr. David Liu, De-
partment of Toxicology, Stanford Research Institute (SRI),
for making specimens available to me. Dr. Liu also provided
extensive field data. Drs. Robert Beeman and Claude Alex-
ander, California State University, San Francisco, gave much
helpful advice. I am especially grateful to Dr. Marian H.
Pettibone for the generous loan of material from the National
Museum of Natural History (USNM), for a xeroxed copy of
part of an unpublished manuscript by J. Percy Moore, and for
17—Proc. Biot. Soc. WasH., VoL. 87, 1974 (175)
176 Proceedings of the Biological Society of Washington
the immense encouragement, help and advice she has offered
me. The manuscript has benefited from the critical reading
of James T. Carlton, California Academy of Sciences. The
material from San Francisco Bay is deposited in the California
Academy of Sciences, the National Museum of Natural His-
tory, the Allan Hancock Foundation and the Zoological Insti-
tute, Academy of Sciences of the USSR, Leningrad.
MALDANIDAE
Asychis elongata ( Verrill )
Figures 1, 2
Maldane elongata Verrill, 1873:343, 365, 371, 377, 609.— Webster, 1879:
259; 1880:124.—Verrill, 1881:302, 318, 322, pl. 9: fig. 1—Andrews,
1892:294.—Hartman, 1944:355, pl. 58: figs. 6, 6a.
Maldanopsis elongata.—Verrill, 1900:659.—Pettibone, 1963:183.
Asychis elongata——Arwidsson, 1907:262.
Branchioasychis colmani Monro, 1939:352-354, fig. 283.
Branchioasychis americana Hartman, 1945:40-—42, pl. 9: figs. 1-4; 1951:
105.—Day, 1973:104.
Asychis amphiglypta—Berkeley and Berkeley, 1960:360.—Hartman,
1969:421, figs. 1-5 (part). [Not Maldane amphiglypta Ehlers, 1897.]
Material examined: Catirornia: San Francisco Bay, 11-17 m, in
fine sands with silt-clay component through equal parts silt-clay to al-
most pure silt, salinities varying from 7—23%., sediment temperature
11.3-13.8°C., March 1973, SRI Sta. 5a, 5c (Redwood City Harbor),
5b, 5d (South Bay, south of San Mateo Bridge), 4 (Hunter’s Point), 3
(Oakland Inner Harbor), 2b (Carquinez Strait), 20 September 1973,
HP-6 (Hunter's Point); MRC Sta. 18.3, 18 November 1971 (Hunter’s
Point) and 18.1, 22 August 1973 (Hunter’s Point)—many specimens.
South San Francisco Bay, 3-6 m, W. A. Newman, M. Leberman, col-
lectors, 11 specimens (USNM 40864-5; as A. amphiglypta by the Berke-
leys). Martner: Morgan’s Bay, L. N. Taxiarchis, collector, 2 specimens
(USNM 49275). Ruope Istanp: Off Newport, 20 m, dark, soft, fetid
mud, bottom temperature 21.7°C. (see Smith and Rathbun, 1886:919),
Fish Hawk Sta. 834, 27 August 1880, 2 specimens (USNM 11854; iden-
tified by A. E. Verrill). Massacnusetts: Woods Hole, wharf dredgings,
August 1883, 10 specimens (USNM 10122; identified by A. E. Verrill).
Hadley Harbor, M. H. Pettibone, collector, 22 August 1953, 3 specimens
(USNM 44138). Detaware: E of Millsboro, Sussex County, channel
>
Fic. 1. Asychis elongata. a, anterior end, left lateral view; b, anterior
end, dorsal view; c, distal portion of spirally fringed capillary.
Atlantic polychaete in San Francisco Bay Me
178 Proceedings of the Biological Society of Washington
marker no. 34, R. D. Jones, collector, 16 February 1972, 1 specimen
(USNM 49200). Vircinta: H. E. Webster, collector, 8 specimens
(USNM 539). Norro Caronina: Beaufort, E. A. Andrews, collector,
1885, 4 specimens (USNM 4883). FLorima: Alligator Harbor, A. J.
Kohn, collector, 21 February 1959, 1 specimen (USNM 34240). Tampa
Bay, J. L. Taylor, collector, 1963, 5 specimens (USNM 45665). Key
West, near bridge of U.S. Naval Hospital, M. L. Jones, collector, 3 May
1964, 2 specimens (USNM 34260). Muississrpp1: Between Biloxi and
Ship Island, 4.6 and 6.8 m (15 and 22 ft), R. L. Caylor, collector, 27
July 1949, 2 specimens (USNM 22094; as B. americana by O. Hartman).
Description (San Francisco Bay specimens): 22 somites, including
fused prostomium and peristomium, first achaetous somite, 19 setigers
and pygidium, no achaetous pre-anal somites. Overall length to 215
mm by 3 mm wide; most specimens considerably smaller, averaging
62-90 mm long by 1.5 mm wide.. Lateral cephalic lobes set off from pos-
terior cephalic lobe by deep lateral notches, former confluent with and
indistinguishable from anterior margin of prostomium; margin of cephalic
plate otherwise entire, smooth, with thin, sharply defined lip (Fig. la, b).
Prostomium forming upturned prow in lateral view (Fig. la) and, to-
gether with lateral cephalic lobes, defining unbroken semicircle in dorsal
aspect (Fig. 1b). Paired nuchal organs completely separated from
margins of cephalic plate (unlike most Asychis), forming small crescentic
or V-shaped glandular depressions. First setiger rolled into thick, com-
plete collar projecting anteriorly around posterior portion of first, achae-
tous somite. Achaetous somite and first 3 setigers completely biannulate,
next 3-4 somites incompletely biannulate, heavily glandularized laterally
and ventrally, dorsum smooth and _ blister-like, occasionally somewhat
transparent; parapodial tori greatly expanded and glandular over this
region. Somites to this point from 1-3 times as long as broad, thereafter
gradually lengthening posteriorly in most individuals up to and including
setiger 17, reaching length-width ratios of 13:1, with some variation de-
pending upon state of contraction of specimen. Usually 3 longitudinal
rows, 1 dorsal and 2 lateral, of rounded, biscuit-like tubercles and papillae
(Fig. 2b), commencing as early as setiger 6 in some specimens; these
rows of papillae occasionally obscured, exhibiting more diffuse pattern.
From setigers 6-14 dorsal transverse folds of tissue forming epaulettes
between notopodial postsetal lamellae at segmental nodes; epaulettes
sometimes faint or absent. Pygidium well developed, trumpet-shaped,
dorsal lobe flared, petaloid, separated from ventral lobe by deep, V-
shaped lateral notches; margins of both lobes completely smooth or very
faintly crenulate (Fig. 2a), margins forming thin lips. Anus situated
on posterior tip of long, rectangular dorsal mound (Fig. 2b).
=>
Fic. 2. Asychis elongata. a, posterior end, left lateral view; b, pos-
terior end, dorsal view; c, spatulate companion seta, showing lateral keel;
d, limbate companion seta.
Atlantic polychaete in San Francisco Bay 179
028 mm
180 Proceedings of the Biological Society of Washington
Notosetae of 3 types: (1) simple, eccentrically sheathed capillaries,
appearing unilimbate upon first inspection, occurring on setigers 1 to 7
or 8; (2) spirally fringed capillaries becoming gradually elongated on
more posterior somites, with sheaths splitting into fimbriae and forming
well expanded, double spiral at distal ends (Fig. 1c), these spirally
fringed capillaries occurring in setigers 8-19, becoming extremely long
posteriorly; (3) long, whiplike, capillary companion setae with greatly
expanded, flattened, spatulate bases, usually with winglike keel lateral
to spatulate portion (Fig. 2c), occasionally limbate (Fig. 2d), or without
keel, being evenly expanded proximally; found between above two types
throughout length of worm. Uncini of typical configuration for Asychis,
in neuropodia of setigers 2-19, up to 30-40 per ramus.
Color in alcohol: pale white, generally with iridescent blue-black or
greyish-purple mottlings and blotches on cephalic plate and dorsum of
anterior segments, pigmentation sometimes quite extensive; individuals
occasionally unmarked; specimens collected from Hunter’s Point (20
September 1973) dark grey-brown, completely opaque. Tubes long,
thick, of black mud and clay, often collected without animals.
Remarks: The animals from San Francisco Bay conform in every
respect to Asychis elongata from the eastern coast of North America,
although the latter tend toward a much larger size. A complete indi-
vidual from Hadley Harbor, Massachusetts (USNM 44138) measures
365 mm in length, and Moore (MS: 870) commented that specimens
may range up to more than 450 mm, although 150-300 mm is most
common. Atlantic specimens are generally much darker, at least in the
northern part of their range (North Carolina to Maine) and vary from
light flesh-brown, red- and yellow-brown, dark red-brown and dark
chocolate-brown to light grey. Four specimens from San Francisco Bay
(SRI Sta. HP-6) are indistinguishable both in color and form from
Atlantic examples. The anterior mottled pigmentation is generally red-
dish-brown to purple-grey or blue-black and the anterior segments are
nearly always strongly iridescent. Specimens previously identified as
Branchioasychis americana from North Carolina south through Florida
and into the Gulf of Mexico are lighter in color, ranging from yellow-
brown, yellow-ochre and flesh to cream-white; the anterior mottled pig-
mentation tends to be much reduced, and is usually blue- to sepia-black.
A cream-white specimen from Alligator Harbor, Florida (USNM 34240)
was apparently dark red in life, as indicated by the collector, A. J. Kohn.
Branchioasychis americana Hartman was described from specimens
from North Carolina and southern Florida and included the record of
Maldane elongata by Andrews from Beaufort (see below). Specimens
from these localities, including some identified as B. americana by Hart-
man ( Mississippi, USNM 22094, and the specimens recorded by Andrews,
USNM 4883) were carefully examined. They conform closely in every
respect to Asychis elongata with the exception of the presence of “gills.”
The dense gill-like diverticula, found on the dorsal and lateral surafces
Atlantic polychaete in San Francisco Bay 181
of setigers 6 to 10 or 11, are arranged in a more or less distinct pattern
(although this is somewhat obscured by their density) of parallel lines
running postero-anteriorly from the middorsal line at an angle of 45°
from the longitudinal axis of the worm. The pattern exhibited on one
side is a mirror image of that seen on the other. It is not clear whether
these dorsal outgrowths represent branchiae or some other type of
structure; they appear in some individuals and not in others. Of 8
specimens collected by H. E. Webster from Virginia (USNM 539), one
specimen exhibits the dense feltlike dorsal outgrowths characteristic of
Branchioasychis. Andrews (1892:294) remarked that a fragment in his
collection of Maldane elongata exhibited these “slender papillae” over
several somites. Of the 4 specimens collected by Andrews (USNM
4883), 2 of these bear dense dorsolateral outgrowths. Branchioasychis
Monro, 1939, with type-species B. colmani Monro, 1939, from British
Honduras, was separated from Asychis Kinberg, 1867, on the basis of
these so-called branchiae. Monro (1939:354) mentions Andrews’ speci-
mens as possibly synonymous with B. colmani. Branchioasychis colmani
is herein synonymized with Asychis elongata, since there are no sig-
nificant differences between Monro’s excellent description (1939:352—
354, Fig. 283) and the characters seen in A. elongata. It seems clear
that Branchioasychis Monro is a junior synonym of Asychis Kinberg.
The presence of these dorsal outgrowths, found in some specimens
and not in others throughout its range and within single samples from
the same locality, may be due to an expression of developmental pheno-
typic flexibility. It is curious that the species tends towards greater
plasticity and a greater development of the dorsal diverticula in the
southern part of its range.
Moore (MS: 869) included Maldane elongata Verrill in the synonymy
of Maldane urceolata (Leidy) [= Clymene urceolatus Leidy, 1855].
Hartman (1959:450, 455) refers Leidy’s species to Asychis; however,
C. urceolatus possessed 26 segments according to Leidy (1855:145),
whereas the genus Asychis is not known to have more than 19 setigers.
I have been advised by Dr. Robert Robertson, The Academy of Natural
Sciences of Philadelphia, that a thorough search failed to turn up the
holotype of Clymene urceolatus Leidy and that it must be presumed lost.
Because of the inadequacy of the original description and the lack of
figures, this taxon is in substantial doubt, and, in concurrence with a
suggestion by Dr. Marian Pettibone (personal communication), Clymene
urceolatus Leidy is herein declared a nomen dubium.
Distribution: Maine to Florida, Gulf of Mexico, British Honduras.
San Francisco Bay, California (new record). In clay, silt and sandy
mud, intertidal to 37 meters.
LITERATURE CITED
ANDREWS, E. A. 1892. A report upon the Annelida Polychaeta of
Beaufort, North Carolina. Proc. U.S. Nat. Mus. 14(852):
227-302.
182 Proceedings of the Biological Society of Washington
Arwipsson, I. 1907. Studien iiber die skandinavischen Maldaniden.
Zool. Jahrb., Suppl. 9:1-308.
BERKELEY, EF. AND C. BERKELEY. 1960. Notes on some Polychaeta
from the west coast of Mexico, Panama, and California. Canad.
Jour. Zool. 38( 2) :357—362.
Day, J. H. 1973. New Polychaeta from Beaufort, with a key to all
species recorded from North Carolina. NOAA Tech. Rept.
NMFS CIRC-375, 140 pp.
Enuers, E. 1897. Polychaeten. In: Ergebnisse der Hamburger
Magalhaenischen Sammelreise. Friedrischson & Co., Hamburg,
148 pp.
Hartman, O. 1944. New England Annelida. Part 2. Including the
unpublished plates by Verrill with reconstructed captions. Bull.
Amer. Mus. Nat. Hist. 82(7):327-344.
——. 1945. The marine annelids of North Carolina. Bull. Duke
Univ. Mar. Sta., No. 2:1-54.
—. 1951. The littoral marine annelids of the Gulf of Mexico.
Publ. Inst. Mar. Sci. 2( 1) :7—124.
—-. 1959. Catalogue of the polychaetous annelids of the World.
Part II. Allan Hancock Found. Publ., Occ. Pap. 23:255-628.
———. 1969. Atlas of the sedentariate polychaetous annelids from
California. Allan Hancock Foundation, Univ. S. California, Los
Angeles, 812 pp.
Kinperc, J. 1867. Annulata nova. Oecfvers. K. Vetenskaps Akad.
Stockholm, Foérhandl]. 23:337-357.
Lewy, J. 1855. Contributions towards a knowledge of the marine in-
vertebrates of the coast of Rhode Island and New Jersey. Jour.
Acad. Nat. Sci. Philadelphia 3(2):135-158.
Monro, C. C. A. 1939. Polychaeta of the “Rosaura” expedition. Novit.
Zool., Tring 41(4):345-354.
Moors, J. P. MS. The polychaetous annelids of the Woods Hole re-
gion. Unpublished, deposited in the National Museum of
Natural History, 1,032 pp.
PeTTIBONE, M. H. 1963. Marine polychaete worms of the New
England region. 1. Aphroditidae through Trochochaetidae.
Bull. U.S. Nat. Mus., Part 1, No. 227:356 pp.
SmiTH, S. AND R. Ratusun. 1886. Lists of the dredging stations of
the U.S. Fish Commission from 1871 to 1879, inclusive, with
temperature and other observations. U.S. Fish Comm. Rept.,
Pt. XIV, 1886:877—-1017.
VERRILL, A. E. 1873. Report upon the invertebrate animals of Vinyard
Sound and the adjacent waters, with an account of the physical
characters of the region. U.S. Fish Comm. Rept. for 1871-72:
295-778.
———. 1881. New England Annelida. Pt. 1. Historical sketch, with
annotated lists of species hitherto recorded. Trans. Connecticut
Acad. Arts Sci. 4:285-324.
Atlantic polychaete in San Francisco Bay 183
. 1900. Additions to the Turbellaria, Nemertina and Annelida
of the Bermudas, with revisions of some New England genera
and species. Trans. Connecticut Acad. Arts Sci. 10:595-671.
WesstEerR, H. E. 1879. Annelida Chaetopoda of the Virginian coast.
Trans. Albany Inst. 9:202—272.
1880. Annelida Chaetopoda of New Jersey. New York State
Mus., Ann. Rept. 32:101-128.
184 Proceedings of the Biological Society of Washington
SS
Vol. 87, No. 18, pp. 185-204 22 October 1974
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
A REVIEW OF THE CORAL TOADFISHES OF THE
GENUS SANOPUS WITH DESCRIPTIONS OF TWO NEW
SPECIES FROM COZUMEL ISLAND, MEXICO
By Bruce B. COLLETTE
National Marine Fisheries Service Systematics Laboratory,
National Museum of Natural History, Washington, D.C. 20560
Two species of western Atlantic toadfishes have been known
only from their original descriptions: Opsanus barbatus Meek
and Hildebrand, 1928, based on the holotype from Colon,
Panama, and Opsanus astrifer Robins and Starck, 1965, based
on five specimens from the Turneffe Islands, Belize (British
Honduras). Smith (1952) named the genus Sanopus for O.
barbatus because of its higher number of dorsal fin rays (24-27
in Opsanus vs. more than 30 in Sanopus). O. astrifer was
transferred to Sanopus based on its agreement with O. barbatus
in high number of fin rays and absence of distinct pectoral fin
glands (Collette, 1966). New material of S. barbatus and S.
astrifer collected by George C. Miller and David W. Green-
field, respectively, stimulated Collette to write a short paper
diagnosing Sanopus and its two species. At about the same
time, Walter A. Starck II was planning to describe two new spe-
cies of toadfishes that he had collected around Cozumel Island,
Mexico. Independently, Craig Phillips obtained specimens of
one of these species and planned to describe it. In the interests
of having a complete review of the genus, we have combined
forces with Collette writing the review and Starck and Phillips
participating in the description of the two new species. A total
of 26 specimens of Sanopus is now available: 12 S. barbatus,
7 S. astrifer, 6 S. splendidus n.sp. Collette, Starck, and Phillips
and 1 S. johnsoni n.sp. Collette and Starck.
18—Proc. Bron. Soc. Wasu., Vou. 87, 1974 (185)
186 Proceedings of the Biological Society of Washington
A review of coral toadfishes 187
MetrHops AND MATERIALS
At the subfamily level, important characters in the Batra-
choididae include presence or absence of photophores and of
hollow spines connected to venom glands (Smith, 1952; Col-
lette, 1966). At the generic level, characters such as the pres-
ence or absence of scales and the nature of the glands around
the pectoral fin region (Collette, 1966) are important. Other
useful characters include number of vertebrae (precaudal and
caudal), number of fin rays (dorsal, anal, and pectoral),
number and shape of teeth, shape of anterior nostril tube
(Roux, 1971), number of lateral lines, number of opercular
and subopercular spines, and number and shape of barbels,
flaps, and cirri on the head and body. Within Sanopus, the
most important species characters are the relative branching
of barbels on the chin and head and the color pattern. Num-
bers of fin rays and vertebrae are more valuable than morpho-
metric characters.
Counts of vertebrae and dorsal and anal fin rays were made
with the aid of radiographs. All dorsal and anal elements were
counted. Vertebral counts include the two hypural vertebrae.
Two factors cause difficulties in counting teeth. Teeth are
frequently broken or missing, leading to low counts when
estimating the number of empty sockets. High counts result
from counting extra replacement teeth. To arrive at the best
estimate for the number of teeth on the premaxillary, palatine,
and dentary bones, counts for left and right sides were aver-
aged and rounded off to the nearer higher whole number.
These averaged numbers are used in the species descriptions
(except for the vomer, for which total counts are given).
Papillae were counted along the dorsal lateral line from above
the opercular spine to the caudal base and along the ventral
lateral line from between the pectoral and pelvic fin origins
to the caudal base. Standard length was measured from the
tip of the upper jaw to the caudal base. Head length extends
<
Puate I. Sanopus splendidus Collette, Starck and Phillips, n.sp.,
Cozumel I., México. Photograph by Hans Flaskamp.
188 Proceedings of the Biological Society of Washington
to the most posterior part of the opercular flap. Head width
Was measured at its maximum point. Orbit length is the maxi-
mum length of the clear part of the orbit. Interorbital width
is the least distance between the clear parts of the two eyes.
Snout to second dorsal and snout to anal are straight line
distances to the origins of these fins. Pectoral fin length is
measured from the upper base of the fin to the tip of the
longest ray; pelvic fin length from the outer base to the tip
of the longest ray.
Material examined is in the following collections: ANSP
(Academy of Natural Sciences, Philadelphia); CAS (Cali-
fornia Academy of Sciences, San Francisco); FMNH (Field
Museum of Natural History, Chicago); MCZ (Museum of
Comparative Zoology, Harvard University); SIO (Scripps In-
stitution of Oceanography, La Jolla, California); UCR (Uni-
versity of Costa Rica, San Jose); UMML (University of
Miami, Rosenstiel School of Marine and Atmospheric Science,
Miami); UP (Departamento de Biologia Marina, Universidad
de Panama); and USNM (U.S. National Museum of Natural
History, Washington).
Sanopus Smith
Sanopus Smith, 1952:314 (type-species, by original designation, Opsanus
barbatus Meek and Hildebrand).
Diagnosis: A genus of the subfamily Batrachoidinae (lacks hollow
dorsal and opercular spines connected to venom glands and _ lacks
photophores ). Closest to Opsanus, which it resembles in six characters:
body scaleless; three solid spines in first dorsal fin; an axillary pore
(leading to a group of glands) present behind pectoral fin; anterior
nostrils at the ends of narrow tubes arising from flaps projecting beyond
upper jaw; two opercular and one subopercular spines; second dorsal
rays 24 or more. Differs from Opsanus in several important characters
as summarized in Table 1. No discrete glands on the posterior surface
of the pectoral fin between bases of the upper fin rays. Only one fin
ray supported by the last dorsal and anal pterygiophore (usually two in
Opsanus). The species of Sanopus are longer-bodied than the species
of Opsanus as reflected in the higher numbers of vertebrae; dorsal and
anal fin rays; papillae along the dorsal and ventral lateral lines. Corre-
lated with the larger size of Sanopus, adults develop more teeth: pre-
maxillary 29-44 vs. 20-28; dentary 21-32 vs. 13-21; palatine 12-22
vs. 9-15; and total vomerine teeth 7—15 vs. 5-10.
A review of coral toadfishes 189
TaBLE 1. Diagnostic characters of Opsanus and Sanopus.
Character Opsanus Sanopus
Pectoral fin glands 6-16 well-developed absent
glands between upper
rays on posterior
surface of fin
Number of rays on last usually 2 always 1
dorsal and anal
pterygiophores
Precaudal vertebrae 9 or 10 11, occasionally 12
Caudal vertebrae Q3=25 26-29
Total vertebrae 31-36 ov—41
Dorsal fin rays =o 29-34
Anal fin rays 19=23 24-28
Upper lateral line papillae 24-30 30-41
Lower lateral line papillae 21-26 MS
Premaxillary teeth (one side) 20-28 29-44
Dentary teeth (one side) 13=2)1 Pile oe,
Sanopus barbatus (Meek and Hildebrand)
Figs. 2 and 3
Opsanus barbatus Meek and Hildebrand, 1928:917-918 (original descrip-
tion; Colon, Panama; USNM 81009; pl. 91).—Walters and Robins,
1961:13 (comparison with other species of Opsanus).—Robins and
Starck, 1965:249-250 (comparison with O. astrifer).
Sanopus barbatus.—Smith, 1952:314 (type-species of Sanopus Smith).
Collette, 1966:848 (genus recognized as valid).—Randall, 1968:285
(listed ).
Diagnosis: S. barbatus is similar to S. johnsoni and different from
S. astrifer and S. splendidus in having branched chin barbels, brown
dorsum with mottled belly, and generally higher counts. It differs from
S. johnsoni primarily in the nature of the branching of the barbels. The
main barbels are shorter and thicker in S. barbatus and the branches are
also short and thick (compare Figs. lc and Id). The papillae along the
lateral line are much shorter and wider in S. barbatus than in S. johnsoni
(compare Fig. 5c with 5d). No cirri are present on the interorbital
region of S. barbatus as they are on S. johnsoni. Pectoral fin shorter than
in S. johnsoni (157-166 vs. 175 thousandths of SL, Table 2).
Frequency distributions of counts (except teeth) presented in Table
3. Dorsal fin rays III, 31-34, usually III, 33; anal fin rays 25-28; pectoral
fin rays 20-22, usually 20-21; vertebrae (11-12) + (28-29) = 39-41,
190 Proceedings of the Biological Society of Washington
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Fic. 2. Anterodorsal view of head of Sanopus barbatus (UCR 442-2,
370 mm SL, Uvita I., Costa Rica).
usually 40; upper lateral line papillae 32-38; lower lateral line papillae
27-34; dentary teeth 21-32 (x 25.4); palatine teeth 12-22 (x 16.4);
premaxillary teeth 29-39 (x 34.3); and total vomerine teeth 8-15 (x 9.0).
Changes with growth: Comparisons can be made between seven large
specimens (213-370 mm SL) and five juveniles (two, 92.1-97.4 mm,
and three 28.7-36.0 mm). Juveniles have larger eyes, longer pectoral
and pelvic fins, narrower heads and interorbital distances and shorter
snout to second dorsal and snout to anal origin distances (Table 2).
Meristically, counts of fin rays, vertebrae, and lateral line papillae do
not change with size. The number of teeth on all four toothed bones
<
Fic. 1. Diagrammatic anterodorsal views of heads of four species of
Sanopus. A. S. astrifer (USNM 209720, 245 mm SL, Glovers Reef,
Belize). B. S. splendidus n.sp. (holotype, USNM 205944, 245 mm SL,
Cozumel I., México). C. S. barbatus (USNM 211322, 250 mm SL,
Bonacca, Honduras). D. S. johnsoni n.sp. (holotype, USNM 205945, 253
mm SL, Cozumel I., México).
192 Proceedings of the Biological Society of Washington
A B C
Fic. 3. Diagrammatic views of left and right sides of caudal fin of
three large Sanopus barbatus to show caudal ocelli. A. USNM 81009,
329 mm SL, Panama, holotype B. UP 736, 355 mm SL, Panama.
C. UCR 442-2, 370 mm SL, Costa Rica.
increases with size: dentary from 15-18 to 21-32; palatine from 10-13
to 12-22; vomer from 5—9 to 8-15; and premaxillary from 15—21 in the
three smallest specimens (and also the 97.4 mm specimen) to 29-39 in
the seven adults (and also the 92.1 mm specimen).
The papillae along the lateral line are all simple in the four smallest
specimens (28.7-92.1 mm), the anteriormost few are branched in the
97.4 mm specimen, most are branched in the 213 and 215 mm speci-
mens, and they are branched the length of the lateral line in the five
largest specimens (306-370 mm). The branching of the barbels on
the head becomes increasingly complex with age (compare Fig. lc cf
the head of a 250 mm SL specimen with Fig. 2 of a 370 mm specimen).
Specimens smaller than 250 mm have reduced branching. Caudal ocelli
(Fig. 3) are present on the tails of the three largest specimens (329,
355, and 370 mm) but not on the nine smaller specimens (28.7—306 mm).
Range: Central American coast of Caribbean Sea. Originally described
from a specimen from Colon Reef, Panama, and now also known from
Costa Rica and Honduras, as well as several additional Panamanian
localities.
Material Examined: PanamA: USNM 81009 (329); Colon Reef;
S. E. Meek and S. F. Hildebrand; 6 May 1912; holotype. UP 736 (355);
Maria Chiquita, about 17 km NE of Colon; 5 May 1973. MCZ 44549
(28.7); Canal Zone, Ft. Sherman; I. Rubinoff; 16 June 1964. MCZ
44550 (36.0); Canal Zone, Ft. Sherman; I. Rubinoff; 26 July 1964.
SIO 67-45 (2, 29.5-92.1); Toro Point; R. Rosenblatt, I. and R. Rubinoff;
23 March 1967. UMML 22522 (97.4); Canal Zone; Pulpit Point, W
Shore Limén Bay; R/V Pillsbury 319; 4 July 1966.
193
A review of coral toadfishes
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194 Proceedings of the Biological Society of Washington
Cosra Rica: UCR 442-2 (370); Limon Prov.; Isla Uvita off Puerto
Limon; W. and M. Bussing and R. Nishimoto; 4 October 1970.
Honpuras: UMML 31257 (306); The Town, Guanaja (Bonacca);
G. C. Miller; 5 May 1967. USNM 211322 (3, 213-306); The Town,
Guanaja (Bonacca); G. C. Miller, 6 May 1967.
Sanopus johnsoni Collette and Starck, new species
Fig. 4
Diagnosis: S. johnsoni is most similar to S. barbatus in general color
pattern (uniform brown with reticulated belly), counts of fin rays and
vertebrae, measurements, and in having branched chin barbels. The
chin barbels are longer and thinner in S. johnsoni than in S. barbatus
and the branches are longer, thinner, and fewer in number (compare
Fig. ld and 1c). The papillae along the lateral line are longer and
thinner in S. johnsoni than in any of the other species of the genus
(compare Fig. 5a-c with 5d). There are a number of long, thin cirri
on the interorbital region in S. johnsoni that are absent in S. barbatus.
They are also absent in S. splendidus and only a few are present in S.
astrifer. The pectoral fin is longer than in S. barbatus (175 vs. 157-166
thousandths of SL, Table 2).
Dorsal fin rays III, 32; anal fin rays 25; pectoral fin rays 22; vertebrae
1] + 28 = 39; upper lateral line papillae 37; lower lateral line papillae
32; dentary teeth 20-21; palatine teeth 14-15; and total vomerine teeth
10. Premaxillary teeth more numerous (R-38, L-48) than the range
in S. barbatus (29-37).
Etymology: Named in honor of Mr. J. Seward Johnson in recognition
of his generous and extensive patronage of marine science.
Habitat: The holotype was speared at night as it rested part way out
of a small cave under a coral head. Another individual was seen among
the rock and coral on top of the reef near the same location on another
night dive. It retreated into a crevice after a few seconds exposure in
the beam of the diving light. The stomach of the holotype was empty
but the intestine was greatly distended, chiefly with parts of the spines
and test of a small specimen of sea urchin, Diadema antillarum.
Holotype: USNM 205945 (253); México; Quintana Roo; Cozumel
I., Palancar Reef near S end of island, just inside drop-off at 8 m;
Walter A. Starck, II; 30 September 1968.
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Fic. 5. Anterior portion of lateral line in four species of Sanopus
showing differences in shape of papillae that protect the neuromast
dorsally and ventrally. A. S. astrifer (USNM 209720). B. S. splendidus
n.sp. (USNM 205944). C. S. barbatus (USNM 211322). D. S. johnsoni
nsp. (USNM 205945).
A review of coral toadfishes 195
196 Proceedings of the Biological Society of Washington
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Sanopus astrifer (Robins and Starck)
Enger, (6)
Opsanus astrifer Robins and Starck, 1965:247—250 (original description;
Turneffe Is., British Honduras; USNM 259421-F1; fig. 1, table 1).
Greenfield and Greenfield, 1973:564 (reported from Glovers Reef,
Belize ).
Sanopus astrifer—Collette, 1966:848 (transferred to Sanopus ).—Randall,
1968:285 (listed).
Diagnosis: Differs from the other three known species of Sanopus in
having the dorsum dark with prominent white spots. Most similar to
S. splendidus in having unbranched chin barbels and generally lower
counts. The barbel at the corner of the mouth is about the same size
as those on the chin, not wider and longer as in S. splendidus (compare
Figs. la and 1b). S. astrifer has the broadest head of the four species
(323-333 vs. 290-316 thousandths of SL) but is similar to the other
species in most other measurements (Table 2).
Frequency distributions of counts (except teeth) are presented in
Table 3. Dorsal fin rays III, 30-31; anal fin rays 24—25; pectoral fin
rays 21-22, usually 22; vertebrae 11 (rarely 12) + (26-28) = 37-39,
usually 37-38; upper lateral line papillae 37-41; lower lateral line
papillae 33-36; dentary teeth 23-24 (x 23.7); palatine teeth 13-16
(x 14.8); premaxillary teeth 33-44 (x 39.0); and total vomerine teeth
10-11 (x 10.7).
Changes with growth: The three largest specimens (233-245 mm SL)
were compared with the four smallest (one 72.9 mm, three 24.7—37.6
mm). Morphometrically (Table 2), the four smallest specimens had
longer heads, orbits, pectoral fins, and pelvic fins and smaller head
width and snout to anal distances. Meristically, counts of fin rays, verte-
brae, and lateral line papillae do not change within this size range. The
number of teeth increases on the vomer (8-9 vs. 10-11), dentary (18-19
vs. 23-24), and premaxillary (32-34 teeth in the 72.9 mm specimen vs.
36-44), but not on the palatine (14-16 vs. 13-16).
Habitat: The types and three specimens recently collected by Dr.
Greenfield all came from caves or hollows underneath Montastrea coral
heads, M. annularis in the case of the Greenfield specimens. Most speci-
mens were taken over sand but one was resting on coral (Robins and
Starck, 1965; D. W. Greenfield, pers. comm.). The individual taken
on the coral “produced loud grunts and thumps” when speared by Dr.
Greenfield.
Range: Caribbean, Belize (British Honduras). Described from
Turneffe Is., and now also known from Glovers Reef (Greenfield and
Greenfield, 1973).
Material Examined: Belize (British Honduras). USNM 259421-F1
(holotype, 236); UMML 9415 (paratype, 73); and ANSP 102736 (3
paratypes, 25-38); Turneffe Is., 200 yards ESE of Cay Bokel in a coral
cave at 5 m; W. A. Starck II and E. N. Belcher III; 30 June 1961.
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A review of coral toadfishes
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FMNH 71318 (1, 233); Glovers Reef, Southwest Cay outside of lagoon;
D. W. and T. Greenfield 71-66; 18 August 1971. USNM 209720 (1,
245); Glovers Reef, patch reef inside of lagoon off Long Cay at 3 m;
D. W. Greenfield 70-128; 22 December 1970.
Sanopus splendidus Collette, Starck and Phillips, new species
Plate 1 and Fig. 7
Opsanus superbus Korthaus, 1973:240 (Cozumel; color photograph,
nomen nudum).
Diagnosis: The most brightly colored species of Sanopus. Differs
from the other three known species in having all the fins, except the
first dorsal, broadly bordered with bright orange-yellow in life (white
in preservative). Body ground color dark bluish-gray with numerous
alternating narrow and wider dark-bordered crossbars on head and body
posterior to the second dorsal fin origin (Pl. 1 and Fig. 7). The barbel
at the corner of the mouth is much wider and longer than in the other
species of Sanopus (Fig. 1). Similar to S. astrifer in having conical,
unbranched chin barbels and in lower average counts of second dorsal
rays and caudal and total vertebrae (Table 3). Interorbital width
narrower (69-77 thousandths of SL compared to 84—95 in the other
three species, Table 2).
Dorsal fin rays III, 29-30; anal fin rays 25; pectoral fin rays 21-23,
modally 22; vertebrae 11 + 26 = 37; upper lateral line papillae 30-34;
lower lateral line papillae 28-30; dentary teeth 23-32 (x 27.7); palatine
teeth 13-19 (x 16.5); premaxillary teeth 35-42 (x 39.2); and total
vomerine teeth 7-13 (x 10.8).
Habitat: The three specimens collected by the Starcks, plus another
individual seen in the field, were all at depths of 10-15 m on the narrow
shelf of shoal water along the western side of Cozumel Island. None
were seen by the Starcks at greater depths during numerous dives along
the outer edge of the shelf. Each was in a small cave under a coral
head, facing outwards with its head just inside the entrance. An indi-
vidual observed at night rested partly outside of its cave. The stomachs
of these three specimens were empty but the intestines of two contained
small fish bones, small gastropods, and bristles that may be setae of
polychaete worms. Hans Flaskamp (pers. comm.) reported that the
species was common in sand-floored caves under rocks or corals at
depths of 10-25 m.
Range: Known only from Cozumel Island, Quintana Roo, México.
None were seen by the Starcks in several days diving along the main-
land coast of Yucatan.
Holotype: USNM 205944 (245); Cozumel Is., about 200 m offshore
of middle of W coast of island at 10 m; Jo D. Starck; 28 September 1968.
Paratypes: ANSP 117316 (225); Palancar Reef near S end of island,
just inside drop-off at 15 m; Jo D. Starck; 26 July 1970. UMML 29141
(217); same locality at 8 m; Walter A. Starck II; 1 October 1968. USNM
201
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A review of coral toadfishes
202 Proceedings of the Biological Society of Washington
205606 (@, 252); Cozumel Is., 12 m from shore off seaward coast at
3.5—4.0 m; N. Neilson; 6 January 1970. USNM 205607 ( 2, 246); same
locality; N. Neilson; August 1970. CAS 29110 (250); Cozumel Is.;
N. Neilson; 1971.
DISCUSSION
The lack of pectoral fin glands is the primary reason that Sanopus is
recognized as distinct from Opsanus. As previously shown (Collette,
1966:848), the presence or absence of pectoral fin glands and of the
axillary gland is a useful character at the generic level in the Batrachoidi-
dae. A suite of other characters (Table 1) shows the four species of
Sanopus to be much more closely related to each other than to any of
the four known species of Opsanus: beta Gimther in Goode and Bean,
pardus Goode and Bean, phobetron Walters and Robins, and tau (Lin-
naeus ). Furthermore, the two genera occupy different habitats and are
almost completely separated geographically: Sanopus lives on coral reefs
along the Caribbean coast of Central America; Opsanus is found in the
Gulf of Mexico to Yucatan (plus one record of O. beta from Belize,
Greenfield and Greenfield, 1973), the Bahamas, and along the east
coast of the United States over a variety of bottom types other than
coral reefs.
Sanopus can be divided into two species groups. 1. S. barbatus and
S. johnsoni are sombre-colored brown toadfishes, frequently with mottled
bellies. Both species have branched chin barbels and higher average
numbers of second dorsal rays, caudal vertebrae and total vertebrae.
2. S. astrifer and S. splendidus are strikingly patterned and have conical,
unbranched barbels and lower average numbers of dorsal rays, caudal
vertebrae and total vertebrae. The only case of sympatry in Sanopus is
of S. splendidus and S. johnsoni at Cozumel; however, the ranges of S.
astrifer and S. barbatus almost overlap. S. barbatus extends from Panama
to Honduras and S. astrifer occurs in Belize.
COMPARATIVE MATERIAL EXAMINED
Opsanus beta. FLoripaA: USNM 158548 (1, 186); Clearwater. USNM
21477 (lectotype of O. beta, 200); Pensacola. USNM 23541 (paralecto-
type of O. beta, 185); Punta Rassa. ANSP 68629 (1, 175); Sand Key;
holotype of O. vandeuseni Fowler. Texas: USNM 746 (2 paralectotypes
of O. beta, 104-145); Indianola. México: USNM 192217 (1, 101);
Yucatan, Ascension Bay. USNM 199218 (1, 73); Yucatan, Ascension
Bay. USNM 192354 (2, 44-58); Quintana Roo, Mujeres Is.
Opsanus pardus. FLoripa: USNM 22317 (2 syntypes of O. pardus,
314-324); Pensacola. USNM 73173 (1, 116); Anclote. USNM 142837
(1, 76); Pepperfish Keys. UMML 189 (1, 182); Dry Tortugas. UMML
4220 (1, 200); Ft. Myers.
Opsanus phobetron. BAHAMA Is.; Bimini: 5 paratypes of O. phobetron.
USNM 170961 (1, 127). USNM 170962 (1, 87). ANSP 79480 (1,
A review of coral toadfishes 203
119). ANSP 79481 (1, 114). UMML 498 (1, 114). Also: UMML
927128 (1, 129); Bimini. MCZ 34708 (1, 25); Eleuthera. Cusa: MCZ
12751-4 (4, 56-154).
Opsanus tau.. NEw York: USNM 48976 (1, 131); Peconic Bay.
MarytanpD: USNM 45460 (1, 143); Cornfield Harbor. Vzircinia:
USNM 91202 (2, 74-83); Lewisetta. Grorcia: USNM 258056-F10
(1, 118); Sapelo Is. FLorma: USNM uncat. (1, 125); Cape Canaveral.
USNM 18034 (1, 161); Matanzas R. Inlet.
ACKNOWLEDGMENTS
The initial stimulus to write this paper was provided by George C.
Miller (NMFS Southeast Fisheries Center, Miami) and David W. Green-
field (Northern Illinois University) by their collecting fresh material of
S. barbatus and S. astrifer, respectively. Information and material was
provided by John Bleck (University of California, Los Angeles), James
E. Bohlke (ANSP), William Bussing (UCR), C. E. Dawson (Gulf
Coast Research Laboratory, Ocean Springs, Miss.), William N. Esch-
meyer (CAS), Richard H. Goodyear (UP), Robert K. Johnson (FMNH),
John E. McCosker (CAS), C. Richard Robins (UMML), Richard
Rosenblatt (SIO), and Robert Schoknecht (MCZ). W. A. Starck, II
acknowledges the National Geographic Society for its support of his
work in the western Caribbean Sea. The original color photograph of
Sanopus splendidus was taken by Hans Flaskamp. It appears here
through the courtesy of Herr Flaskamp and Edith Korthaus, editor of
Das Aquarium, where the photograph was originally published. The
figures were drawn by Keiko Hiratsuka Moore and the radiographs were
taken by George Clipper, Susan Stephens, Eugenia Bohlke, and Frank
McKinney. Drafts of the manuscript were reviewed by Daniel M. Cohen,
Thomas H. Fraser, Carter R. Gilbert, David W. Greenfield, George C.
Miller, C. Richard Robins, and Victor G. Springer.
LITERATURE CITED
CoLLeTTE, Bruce B. 1966. A review of the venomous toadfishes, sub-
family Thalassophryninae. Copeia 1966 (4) :846—-864.
GREENFIELD, Davin W. AND TERRY GREENFIELD. 1973. Triathalas-
sothia gloverensis, a new species of toadfish from Belize
(= British Honduras), with remarks on the genus. Copeia
1973 (3) :560-565.
Kortuaus, Epirw. 1973. Anglerfisch Opsanus superbus Phillips. Das
Aquarium No. 48:240.
MEEK, SETH EF). AND SAMUEL F. HILDEBRAND. 1928. The marine fishes
of Panama. Part III. Field Mus. Nat. Hist. Publ. no. 249,
Zool. Ser. 15(3):709—1045.
RANDALL, JOHN E. 1968. Caribbean reef fishes. T.F.H. Publ., Inc.,
Jersey City, N.J. 318 pp.
- Rosins, C. RicHARD AND Wa ttTeR A. Starck, II. 1965. Opsanus
204 Proceedings of the Biological Society of Washington
astrifer, a new toadfish from British Honduras. Proc. Biol. Soc.
Wash. 78:247—250.
Roux, Cu. 1971. Révision des poissons marins de la famille des
Batrachoididae de la cédte occidentale Africaine. Bull. Mus.
Nat. Hist. Nat., ser. 2, 42(4):626—-643.
SmitH, J. L. B. 1952. The fishes of the family Batrachoididae from
South and East Africa. Ann. Mag. Nat. Hist., ser. 12, 5:313-
339.
WALTERS, VLADIMIR AND C. RicHarRD Rosins. 1961. A new toadfish
(Batrachoididae ) considered to be a glacial relict in the West
Indies. Amer. Mus. Novitates No. 2047, 24 pp.
Vol. 87, No. 19, pp. 205-216 22, October 1974
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
A NEW EASTERN PACIFIC LABRID (PISCES),
DECODON MELASMA, A GEMINATE SPECIES OF
THE WESTERN ATLANTIC D. PUELLARIS!
By Martin F. GoMON
Smithsonian Institution, Washington, D.C. 20560, and
Rosenstiel School of Marine and Atmospheric Sciences,
University of Miami, Miami, Florida 33149
The genus Decodon was erected by Giinther (1861) for
Cossyphus puellaris Poey 1860, a species differing from other
members of the genus Cossyphus (= Bodianus) primarily in
lacking scales on the membrane of the soft dorsal fin. For
over a hundred years Decodon was thought to be monotypic
with a distribution restricted to the deeper shelf waters of the
Western Atlantic between North Carolina and Brazil. Recent
collections from along the Pacific Coast of Central and northern
South America in depths similar to those in which D. puellaris
is often taken in the Atlantic, however, have yielded a second,
geminate species. As in many other closely related labrids,
differences between the two species are largely in color pattern
while counts and measurements are nearly identical. Further-
more, two species from the Indo-West Pacific presently placed
in allied genera are referable to Decodon, though not treated
here.
METHODS
Terminology follows that of Hubbs and Lagler (1958) ex-
cept: caudal-ray count includes dorsal unsegmented rays +
dorsal segmented, unbranched rays + segmented, branched
rays + ventral segmented, unbranched rays + ventral unseg-
mented rays; pectoral-fin rays are indicated with unbranched
1 Contribution No. 1701 from the Rosenstiel School of Marine and Atmospheric
Sciences, University of Miami.
19—Proc. Brio. Soc. WasuH., Vou. 87, 1974 (205 )
206 Proceedings of the Biological Society of Washington
rays in lower case Roman numerals and branched, segmented
rays in Arabic (the dorsalmost pectoral ray in labrids is typi-
cally short, unsegmented and unbranched; the second ray is
typically long, segmented and unbranched); first lateral-line
scale is the anteriormost free, pored scale not attached to upper
end of posterior opercular membrane; scales below lateral line
are counted diagonally forward and upward from the first
moderately sized body scale laterally adjacent the first anal-fin
spine to but not including the lateral line (small scales in vi-
cinity of anus and first anal-fin spine are ignored for standard-
ization of counts); gill-raker counts include rudiments; orbital
length is a horizontal measurement. Figures enclosed by
parentheses following meristic counts indicate number of spec-
imens or structures (e.g., pectoral fins) exhibiting count.
Measurements were made with needlepoint dial calipers and
recorded in millimeters; morphometric dimensions are given
as ranges in percent of standard length; all body length
measurements are expressed as the standard length (SL) in
mm. Color descriptions of the new species are based on photo-
graphs and color notes supplied by Lloyd T. Findley and
transparencies loaned by Richard H. Rosenblatt; comparative
color transparencies of D. puellaris were made available by
Patrick L. Colin.
Type specimens and study material are deposited in the
following institutions: Academy of Natural Sciences of Phila-
delphia (ANSP); California Academy of Sciences (CAS); Los
Angeles County Museum (LACM); Museum of Comparative
Zoology, Harvard University (MCZ); Scripps Institution of
Oceanography, University of California (SIO); University of
Arizona (UA); University of Costa Rica (UCR), also tem-
porarily housing Searcher specimens; School of Marine and
Atmospheric Sciences, University of Miami (UMML); USS.
National Museum of Natural History, Smithsonian Institution
(USNM); University of California at Los Angeles (W).
ACKNOWLEDGMENTS
My sincere gratitude is extended to those individuals who
helped in obtaining specimens, photographs and field data:
Patrick L. Colin, C. Richard Robins and Susan J. Stevens,
Geminate labrids from Western Hemisphere 207
School of Marine and Atmospheric Sciences, University of
Miami; Lloyd T. Findley, University of Arizona; Richard H.
Rosenblatt, Scripps Institution of Oceanography, University
of California; William Bussing, University of Costa Rica;
Phillip C. Heemstra, Academy of Natural Sciences of Phila-
delphia; Leslie W. Knapp, Smithsonian Oceanographic Sorting
Center; Robert J. Lavenberg and Camm C. Swift, Los Angeles
County Museum. I thank Ernest A. Lachner, Smithsonian In-
stitution, Washington, D.C., and C. Richard Robins for criti-
cally reviewing the manuscript.
Decodon Gunther
Giinther 1861, p. 384. Type-species by monotypy, Cossyphus puellaris
Poey 1860.
Decodon puellaris, the new species and two species from the Indo-West
Pacific form a natural group within the bodianine labrids (e.g., Bodianus,
Choerodon, Pimelometopon), here recognized at the generic level. A
treatment of the species occurring in the Indo-West Pacific, together with
an examination of the general systematic status of Decodon, is deferred
until a study of the bodianine labrids by the author is complete. A
combination of the following characters separates Decodon from other
closely related genera of bodianines: absence of scales on basal portion
of dorsal and anal-fin membranes; dorsal-fin rays XI, 9-10; anal-fin rays
III, 10; pored lateral-line scales 26-30; trunk scales deciduous; predorsal
area scaled from dorsal-fin origin to anterior nostrils; preoperculum
completely scaled.
Decodon melasma, new species
(Figure 1)
Holotype: USNM 210859 (139.6), Mexico, Gulf of California, Baja
California Sur, 3 miles (4.8 km) off Buena Vista, small reef surrounded
by sand, depth 55 fathoms (100 m), collected by Ray Cannon and
Boyd W. Walker, 15-16 March 1961, field station no. W61-33, hook
and line.
Paratypes: USNM 210860 (4, 56.2-106.9), Mexico, Gulf of Cali-
fornia, Baja California Sur, Bahia Santa Inex, 26° 56.9’ to 58.3’N, 111°
DI MmrOnos ow, and) 26°) 5912) to 5iNe Le ail. to) S102. water
depth 40-45 fathoms (82 m), 40’ otter trawl, collected by W. Baldwin
and party, field station no. RR 65-51, 14 July 1965, 1410-1510 and 1535-
1615 hours; CAS 28809 (4, 52.2-150.1), LACM 33722-1 (4, 59.7-149.3)
and SIO 65-305 (50, 37.8-200.0), same data as USNM 210860.
Other material: Gulf of California, SIO 65-250 (6, 78.6-128.9),
65-257-50 (1, 87.7), 65-294 (1, 118.8), 65-304 (4, 69.6-147.8), 65-313
208 Proceedings of the Biological Society of Washington
(GE Asks), Giseon (il, Ql), Gsarih (ily Bro), Gass (4 440 199.8)), UA
67-59-4 (2, 68.7-86.2), 70-21-5 (4, 1983-1361), 70-419) (4s eato™
126.8), W61-79 (1, 132.8), W63-148 (2, 129.2-129.5): Costa Rica,
UCI sie) (IE zn), OSI (CL, sd), BH (@, 1O44—167.3)),
510-5 (1, 116.7), Searcher 484 (1, 61.6), 485 (4, 36.8-88.7); Isla del
Coco, Searcher 521 (4, 100-139), 522 (1, 169), 539 (1, 55.2); Gulf
of Panama, UMML 26461 (1, 104.3), 26462 (1, 94.5), USNM 210711
(1, 48.2), 210712 (1, 48.7), 210713 (1, 126.4); Ecuador, USNM 210710
(2, 122.4-198.4).
Diagnosis: Pectoral-fin rays ii, 15, rarely ii, 16; lateral-line scales 28;
adults with vertically elongate dark spot below seventh dorsal-fin spine
extending from just above lateral line (eighth pored lateral-line scale)
to midside; juveniles with up to 6 vertical dusky bars on side of body,
usually confined to dorsal half.
Description: Dorsal- fin rays XI, 10 (72), XII, 9 (1); anal-fin rays
IIT, 10° (72), 1M, 9 (1); caudal-fin rays 942). 10, 0) Soll
1 (1) 12 (61), 13 (1) 42 4773 is) 2 sioner co eatoneeyy
pectoral-fin rays ii, 15 (133), ii, 16 (9), ii, 12 (1), ii, 14 (1); pelvic-fin
rays I, 5; vertebrae I-17 = 28) (61), 1-" IS 295 Ge elateralelime
scales 28; scales above lateral line 2%; scales below lateral line 8; caudal
peduncular scales 16; predorsal scales approximately 20; gill rakers 5 (3),
BB), 7 (6) 28 (3), 20 Gis), 1) 0G), 1 (2D).
Body compressed, elongate, greatest depth in large specimens at about
level of dorsal-fin origin, body depth 20.3-31.6, increasing with increase
in SL; caudal peduncle 11.6—14.8, increasing with increase in SL. Scales
on trunk large, cycloid, deciduous; scales not extending onto bases of
dorsal and anal fins; lateral line smoothly curved, uninterrupted; lateral-
line scales posteriorly notched, possessing unbranched medial canal with
single posterior pore.
Head blunt, length 32.2-38.0; forehead gently curved, curve more
pronounced in large specimens; snout 6.1—11.8, increasing with increase
in size; eye 6.9-10.6, decreasing with increase in SL. Head almost
totally scaled; scales on nape and forehead small, reaching anteriorly to
just in front of anterior nostrils; underside of head and chest with small
scales reaching anteriorly to symphysis of jaw; small scales covering
cheeks and infraorbital area forward to angle of mouth; preopercle scaled
to edge; preopercular edge finely serrate; operculum mostly covered by
large deciduous scales; opercular membrane naked posteriorly. Opercular
membranes united and free from isthmus; ventral end of gill opening
reaching anteriorly to below posterior part of eye. Pseudobranch present;
diagonal series of 5 or 6 fleshy, gill raker-like appendages present on
inner surface of operculum opposing ventral gill rakers of first arch; gill
rakers on first gill arch moderately long, narrow, usually simple; dorsal-
most gill raker on lower limb of arch occasionally bifurcate. Lower lip
narrow, not differentiated from chin at tip of jaw; upper lip broad,
smooth, not fleshy, covering lower lip completely when mouth closed.
Teeth in jaws caniniform, more or less uniserial (Figure 2). Upper jaw
209
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210 Proceedings of the Biological Society of Washington
with two widely spaced, large, curved canines on either side of symphysis
at anterior outer margin of premaxillary, followed posteriorly by row of
approximately 5 to 8 equally short canines arising from narrow bony
ridge on ventral edge of premaxillary; ridge prominent anteriorly, forming
relatively tall cutting edge along mesial side of bases of anterior canines,
becoming less distinct posteriorly; several small teeth often present along
mesial side of bony ridge; usually one or two (0-3) anteriorly curved
canines present at posterior end of upper jaw. Lower jaw with two large
curved canines on each side of symphysis at anterior outer margin of
dentary; first canine directed anteriorly next to symphysis; second canine
widely spaced and larger than first, angled slightly laterally; large
canines followed by row of 3 to 5 short posteriorly longer canines
arising from narrow bony ridge similar to that in upper jaw, and in
turn, followed by second row of shorter canines of equal length; second
series of short canines based on separate bony ridge immediately posterior
to first ridge, also forming cutting edge with short blunt teeth in
large specimens.
Dorsal fin low, continuous, origin slightly anterior to posterior extent of
opercular margin; first dorsal-fin spine shortest, subsequent spines slightly
longer and nearly equal in length; 11th dorsal-fin spine 10.5—12.6; each
spine with filamentous membrane at tip; dorsal fin pointed posteriorly,
not reaching base of caudal fin. Anal fin low; spines progressively longer
posteriorly, 3rd anal-fin spine 8.6-11.7, fin pointed posteriorly, not
reaching base of caudal fin. Caudal fin rounded to truncate in small
specimens, truncate with dorsal and ventral rays produced in large
specimens. Depressed pectoral-fin tip reaching perpendicularly below
7th or 8th dorsal-fin spine; length of pectoral fin 17.2-21.3. Pelvic-fin
tip reaching slightly more than half way from pelvic-fin origin to anus;
pelvic-fin length 12.0-18.0, increasing with increase in SL.
A comparatively small species of bodianine; largest specimen ex-
amined 200.0 mm SL.
Reproduction: Adults appear to be synchronous hermaphrodites in
practically all but very large individuals. Six of seven small adult para-
types (98.3-106.9) examined have a large, well developed ovary with
advanced oocytes on each side of the body and a lobe of testicular tissue
about one-fourth or one-fifth the size of each ovary present next to the
ovary of the left side, no testis on the right. The seventh specimen has
a single testis approximately the same size as the testicular lobe in each
of the other specimens and two strands of connective tissue which appear
to represent spent ovaries. Smaller adult specimens consistently have
two distinct ovarian lobes and a single testicular lobe. The ovary does
not appear to degenerate at any particular adult size as further ex-
emplified by two paratypes, 149.3 and 150.1 mm SL, with well-
developed ovaries, and the holotype, 138.3 mm SL, with only a single
narrow testis on the right side and no vestige of ovarian development.
The largest specimen examined, a paratype (200.0), has testicular tissue
proportionate in size to that of other paratypes and no sign of functional
Geminate labrids from Western Hemisphere all
Fic. 2. Dentition in upper and lower jaws of D. melasma. Premaxil-
lary: (a) ventral view, (b) lateral view. Dentary: (c) dorsal view,
(d) lateral view. USNM 210713, 126.4 mm SL. Line indicates 5 mm.
ovaries. The highly convoluted appearance of the testis in this specimen,
however, suggests that development of testicular tissue may also have
taken place in the walls of the spent ovaries.
Color in alcohol: (Specimens larger than about 130 mm) Head and
212 Proceedings of the Biological Society of Washington
body pale; underside of head, throat and chest with underlying white
pigmented layer visible; moderately large, often faint, diffuse, dusky
spot present on dorsal midline of nape just anterior to dorsal fin origin;
vertically elongate dark spot dorsolaterally on body below seventh dorsal-
fin spine, extending from just above lateral line almost to midside. Fins
transparent.
(Specimens smaller than about 130 mm) Markings better developed
in smaller specimens, fainter in larger ones. Head and body pale; large
dusky spot present on nape anterior to dorsal-fin origin; as many as
six, usually faint, broad, dusky, vertical bands present on dorsal half of
body; bands reaching or nearly reaching dorsal body contour dorsally
and usually extending to or just below lateral midline ventrally; very
small specimens with bands reaching ventral body contour; anteriormost
band positioned below second dorsal-fin spine, second band below
seventh dorsal-fin spine, third band below ninth or tenth dorsal-fin
spine, fourth band below fifth segmented dorsal-fin ray, fifth band on
caudal peduncle just posterior to dorsal fin, fourth and fifth bands often
very faint or absent. Small, dusky spot present middorsally on caudal
peduncle just anterior to caudal fin, remnant of sixth band on basal edge
of caudal fin in very small specimens. Anal, caudal, pectoral and pelvic
fins transparent; dorsal fin transparent in specimens larger than about
45 mm, smaller specimens with large dark spot in center of dorsal fin
usually extending from first to third segmented rays.
Color in life: Head and body pinkish red above, white below. Head
pinkish red on snout and top of head, becoming pinkish white below
level of eye and white below level of mouth. Three curved, moderately
broad, chrome yellow stripes on head; dorsalmost stripe originating on
dorsal margin of orbit, encircling orbital margin dorsoposteriorly, then
curving posteriorly across preopercle and opercle to opercular margin
at upper level of pectoral base where it turns ventrally and forms short,
broad, yellow posterior margin on gill flap; second stripe anteriorly con-
fluent with corresponding stripe of opposite side across snout from
anterior edge of orbit, encircles orbital margin ventrally and is directed
ventroposteriorly from ventroposterior edge of orbit across cheek to
ventroposterior angle of gill flap; third stripe forms narrow margin on
free edge of upper lip (confluent with that of opposite side anteriorly )
and is directed posteroventrally from angle of mouth, becoming broader
ventrally and terminating just below ventral edge of preopercle. Chrome
yellow blotch on nape at anterior end of lateral line. Faint purplish
blue area on opercle above upper level of pectoral-fin base. Trunk
pinkish red above level of posterior lateral line, becoming pinkish white
ventrolaterally and white ventrally. Large, vertically elongate, black
spot on body below seventh dorsal-fin spine extending from just above
Fic. 3. Distribution of D. melasma and D. puellaris.
Geminate labrids from Western Hemisphere Dalles
* = D. melasma
@ OD. puellaris
214 Proceedings of the Biological Society of Washington
lateral line (eighth pored lateral-line scale) almost to midside (approxi-
mately 2 to 2% scale rows high).
Pectoral fin pinkish red. Pelvic fin dusky white; bases of membranes
between inner rays chrome yellow. Dorsal fin pinkish red on spinous
portion, pinkish white on anterior segmented ray portion with chrome
yellow highlights to rays and whitish posteriorly. Anal fin white with
narrow, horizontal, chrome yellow stripe parallel with ventral body con-
tour near base of fin. Caudal fin pinkish red with some chrome yellow
on bases of outer rays.
Distribution: Present only in the Eastern Pacific from the northern
Gulf of California to off Ecuador (Fig. 3). The absence of specimens
from the central portion of this range (Fig. 3) is probably related to
poor collecting efforts. This species is rather common in shrimp-trawl
hauls made in depths of 40 to 160 m. Areas of capture usually have
flat sandy to slightly soft bottoms often with some rocky rubble or
patch reefs, very similar to those areas in the Western Atlantic inhabited
by D. puellaris.
Etymology: melasma, a Greek noun meaning black spot, in reference
to the characteristic single black mark on each side of the body in large
specimens. Melasma is here used as a noun in apposition.
Relationships: D. melasma and D. puellaris differ in counts only in
the number of pectoral-fin rays, ii, 14 (ii, 15 in 2 of 50 counts) in
D. puellaris versus ii, 15 (ii, 16 in 9 of 142 counts) in D. melasma, and
number of lateral-line scales, 28 in D. melasma versus 27 in D. puellaris.
Proportional measurements in specimens of the two species show no
significant differences.
Preserved specimens of D. melasma and D. puellaris are virtually
identical in appearance with the exception of the presence of a single,
dark, lateral body mark in large specimens and several vertical, diffuse,
dark bands on the sides of smaller individuals of D. melasma, in contrast
to the virtual absence of dark pigment in all specimens of D. puellaris
(both D. melasma and D. puellaris possess a dark spot on the anterior
branched, segmented-ray portion of the dorsal fin in specimens smaller
than about 45 mm). Life colors of both are basically pinkish red dorsally
grading to white ventrally with distinct chrome yellow markings on the
head, body and fins. Specimens of D. puellaris possess a number of
chrome yellow spots on the operculum and apparently on most of the
lateral body scales. These spots are not present in specimens of D.
melasma. Specimens of D. puellaris also have a basal row of yellow
spots on the anal fin rather than the basal stripe of D. melasma, and
possess a narrow yellow line along the distal margin of the fin as well
as a number of yellow spots scattered on the midportion of the fin.
The two species also appear to differ in dorsal-fin coloration, although
available photos are not sufficiently clear to evaluate this aspect ade-
quately. Both species, however, possess an identical pattern of yellow
stripes radiating from the eye and have a distinctive chrome yellow
upper lip.
Geminate labrids from Western Hemisphere 215
Both appear to inhabit the same ecological niche in the same depth
range (a relatively unique ecological distribution for bodianines, as most
species are shallow water reef forms or are associated with sheer rocky
conditions ) and are distributed almost identically along their respective
coasts of North, Central and South America in continental shelf conditions.
Adults of both species have an identical blunt head profile in contrast
to the more typically pointed, Bodianus-like head of their two congeners
occurring in the Indo-West Pacific. This fact suggests a closer and more
recent ancestry of the new world species than of either with the Indo-
West Pacific species. The greatest similarities in structure, appearance
and ecology of the two probably point to their recent evolution, thus
affording a good example of a geminate species pair of the tropical
Western Atlantic and Eastern Pacific faunas as envisioned by Jordan
(1905) and discussed by Robins (1972). An interesting aspect con-
cerning the two species lies in their rather deep vertical distribution,
making them the first documented consistently deep water geminate
pair; all other Central American geminates occur in shallow marine or
estuarine habitats (Jordan, 1905).
COMPARATIVE MATERIAL
Comparative material of Decodon puellaris includes: South Carolina,
USNM 151868 (1, 97); Georgia, USNM 156600 (1, 63); northeast
Florida, UMML 29567 (1, 164); eastern Florida, UMML 1236 (1, 73);
Florida Keys, UMML 10206 (2, 41-43); Tortugas, UMML 20527 (3,
33-89), USNM 116960 (5, 45-135); northwest Florida, UMML 4925
(5, 74-115), USNM 156599 (1, 62), 158826 (2, 112-143), 210636 (1,
145), 210638 (2, 114-117); Louisiana, USNM 185304 (1, 138); Texas,
USNM 156616 (1, 58), 185441 (1, 179), 210634 (4, 132-171), 210637
(2, 116-143); Mexico, USNM 210644 (2, 108-116); Yucatan, UMML
28613 (4, 36-73); Honduras, UMML 2712 (1, 93), USNM 185045
(1, 71), 185260 (15, 52-103); Panama, USNM 210645 (2, 64-101),
210646 (1, 91); Colombia, UMML 16295 (2, 96-112), 22290 (4, 39-
62), USNM 210640 (3, 109-116), 210641 (1, 92), 210643 (1, 92);
Venezuela, USNM 185297 (1, 122); Guyana, USNM 210635 (1, 133),
210639 (1, 105), 210642 (1, 122); Surinam, UMML 4974 (1, 142):
Brazil, USNM 185031 (2, 80-86), 185052 (1, 83), 185065 (1, 91),
185192 (2, 67-70); Cay Sal Bank, UMML 9968 (3, 51-79); Cuba,
ANSP 90775 (1, 148), MCZ 14339 (2, 140.0-151.5, types), USNM
Wee ass), 12564 (1, 199), 94941 (1) 183). Jamaica, UMMIE
29888 (1, 157); Dominican Republic, UMML 29564 (2, 40-89);
Grenada, USNM 178125 (1, 122).
LITERATURE CITED
GtntTHer, A. 1861. A preliminary synopsis of the labroid genera. Ann.
Mag. Nat. Hist., 1861 (3) ser. 8:382-389.
Husss, C. L. anp K. F. Lacter. 1958. Fishes of the Great Lakes Re-
216 Proceedings of the Biological Society of Washington
gion. The University of Michigan Press, Ann Arbor. i-xv,
1-135 pp., 44 color pls., 251 figs.
Jorpan, D.S. 1905. A Guide to the Study of Fishes. Henry Holt Co.,
New York. Vol. I:i—xxvi, 1-624 pp., 393 figs.
1908. The law of geminate species. Amer. Nat., 42:73—80.
Pory, F. 1860. Poissons de Cuba, especes nouvelles. In Memorias
sobre la historia natural de la Isla de Cuba, acompanadas de
sumarios latinos y extractos en frances, etc. Habana. Vol. 2,
Chpt. XLIX: 115-356.
Ropins, C. R. 1972. The state of knowledge of the coastal fish fauna
of the Panamic Region prior to the construction of an inter-
oceanic sea-level canal. In M. L. Jones, ed. The Panamic biota:
Some observations prior to a sea-level canal. Bull. Biol. Soc.
Wash., 2:159-166.
Vol. 87, No. 20, pp. 217-230 22, October 1974
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
BRANDTIKA ASIATICA NEW GENUS, NEW SPECIES,
FROM SOUTHEASTERN ASIA AND A REDESCRIPTION
OF MONROIKA AFRICANA (MONRO)
(POLYCHAETA: SABELLIDAE)
By MerepiTH L. JONEs
Smithsonian Institution, Washington, D.C. 20560
In 1939, Monro reported on a collection of polychaetes from
near the mouth of the Congo River in West Africa. Among
these were specimens of a small sabellid from Matadi, approxi-
mately 130 km upstream from the Congo River mouth, in what
is now the Democratic Republic of the Congo. Monro con-
sidered these to be a new species of Manayunkia, M. africana,
in spite of the presence of true spatulate thoracic setae (on the
basis of there being a trend from mere limbation toward a
spatulate condition among various described species of
Manayunkia) and in spite of the presence of two (rather than
three) abdominal setigers. Further, he noted that specimens
of M. africana constructed incrusting, “. . . intertwining sandy
tubules” (Monro, 1939:220) on the shells of the freshwater
gastropod, Hydrobia plena, that the worms were found in areas
of swift current, that they were not confined to snail shells, but
also found on rocks, and that additional dried tubes of the
worms were collected in the M’Pozo River, further upriver
from Matadi.
No additional collections of M. africana have been reported
in the literature. Hartman (1951:389), in connection with a
study of the sabellid polychaetes of the Fabriciinae, considered
the presence of a palmate membrane on the branchial crown
to be sufficient grounds to separate M. africana from
Manayunkia and established the new genus Monroika, based
on Monro’s description.
20—Proc. Bion. Soc. WasH., VoL. 87, 1974 (a7)
218 Proceedings of the Biological Society of Washington
During an extensive study of southeast Asian specimens of
the polychaete genus Caobangia and the molluscs associated
with it (Jones, 1969, 1974), curious incrusting sandy tubes
were observed on the surface of the shells of the viviparid
snail, Mekongia jullieni (Deshayes), and on the valves of the
unionid bivalve, Hyriopsis delaportei (Crosse and Fischer),
both collected by Dr. R. A. M. Brandt from the Mekong River
in southern Laos. Perusal of the southeast Asian freshwater
snails in the collections of the Division of Molluscs, National
Museum of Natural History, Smithsonian Institution, yielded
other snails with similar incrusting tubes, i.e., Brotia cf. baccata
(Gould) and the paratype of Melania pagodula Gould
[= Brotia pagodula (Gould)] both from the Salween River
drainage.
The sandy tubes found associated with the four molluscan
species mentioned above contained the dried remains of a
sabellid similar to Monroika and were in from fair to very poor
condition. Drying notwithstanding, it was possible to observe
a number of similarities and differences relative to Monro’s
description of Manayunkia africana and it was felt necessary
to re-examine Monro’s material deposited in the British Mu-
seum (Natural History) and, subsequently, to redescribe the
type-species of Monroika, as well as to establish a new genus,
Brandtika, for the Asian specimens.
It is with great pleasure that I thank Dr. R. A. M. Brandt,
School of Tropical Medicine, Bangkok, Thailand, for the Lao-
tian molluscs which bore the tubes of Brandtika, Dr. J. David
George, British Museum (Natural History) (BMNH) for
arranging the loan of Monro’s types of Manayunkia africana,
and Dr. Marian H. Pettibone, National Museum of Natural
History (USNM), my colleague and ever ready and ever
willing manuscript reader.
FAMILY SABELLIDAE
SUBFAMILY FABRICIINAE
Monroika Hartman, 1951; emended
Type-species: Manayunkia africana Monro, 1939, by original designa-
tion. Gender: feminine.
Diagnosis: Fabriciine sabellids with 8 thoracic and 2 abdominal
setigers; with long limbate capillary setae on all 10 setigers, with short
limbate capillary setae on setigers 1, 2, 6-8, with spatulate setae on
Polychaete worms from southeast Asia 219
setigers 2-5, with neuropodial long-handled hooks on setigers 2-8, and
with notopodial long-necked uncini on setigers 9-10; branchial crown
of simple radioles united by high palmar membrane (?).
Remarks: Examination of Monro’s syntypes and other material from
the original collection of M. africana have allowed the recognition of
two kinds of limbate capillary setae, and the emendation herein reported
results from this. The presence of “. . . fairly numerous, possibly about
a dozen [simple filamentous radioles] ... and... a high palmar mem-
brane” (Monro, 1939:220) could not be confirmed, for, as Monro stated,
“.. the state of preservation is poor and the more delicate parts such
as the gills are partly disintegrated’ (Monro, 1939:220).
In contrast to Hartman’s judgment, I feel that the crucial generic
differences between Monroika and Manayunkia lie in the type and distri-
bution of setae and the number of thoracic and abdominal setigers, rather
than in having “. . . a weblike membrane in the crown” (Hartman, 1951:
389), which has not been unequivocally demonstrated.
Monroika africana ( Monro)
Figures 1, 2
Manayunkia africana Monro, 1939:220-—223.—Wesenberg-Lund, 1958:29
[list of fresh- and brackish-water polychaetes].
Monroika africana.—Hartman, 1951:389-390.
Material examined: Democratic Republic of the Congo, the Congo
River at Matadi (05°49’S, 13°27’E) E. Dartevelle, collector; syntype
mounted on microscope slide (BMNH 1939.7.17.1); three syntypes
mounted on microscope slide (BMNH 1989.7.17.2); additional speci-
mens, (USNM 50913) dissected from sandy tubes on snails labelled as
follows: “Manayunkia africana—[BMNH] 1939.7.17.12 Holotype [sic]
—RMatadi, Congo River.”
Type-material: Syntypes are from 2 to 3 mm in length and from 0.2
to 0.3 mm in width at their widest part. Little detail of the branchial
crown can be discerned. One of the syntypes (BMNH 1939.7.17.2) is
in two pieces. None of the syntypes appears to be the specimen figured
by Monro (1936: Fig. 1).
Description: The body is small, having 10 setigerous segments, 8
thoracic, with notopodial setae and neuropodial hooks, and 2 abdominal,
with notopodial hooks and neuropodial setae; the anterior end is pro-
vided with a branchial crown (Fig. 1). There is a collar which bears
what appear to be shallow dorsolateral incisions (Fig. lc, d), as well as
a mid-dorsal notch and groove (Fig. 1d).
The first setiger is provided only with unilimbate capillary notosetae
of 2 types, 3-5 longer ones (Fig. 2a) and 2-4 shorter ones (Fig. 2b;
Tables 1, 2). The following 7 thoracic setigers all bear longer limbate
capillary notosetae (Fig. 2f, i, m, r); the 2 abdominal setigers bear long
capillary notosetae (Fig. la) which may be smooth or provided with
extremely fine limbations. There are from 1-4 short limbate capillary
220 Proceedings of the Biological Society of Washington
Polychaete worms from southeast Asia 22h
notosetae on setiger 2 and from 1-3 or 4 on setigers 6-8 (Fig. 1n, p, s).
Spatulate notosetae occur on setigers 2-5 (Fig. 2d, g, k). Occasional
short limbate capillary notosetae were found on setigers 3 and 5 (Fig.
2j). Limbations tend to become progressively less well-developed toward
the posterior end.
There are 2—5 long-handled avicular notopodial hooks on setigers 2-8.
These have a stout, sinuous manubrium surmounted by a large central
fang topped by series of small denticles, becoming smaller, more distal
to the central fang (Fig. 2c, e, h, 1, 0, q, t). When viewed en face it is
apparent that there is some asymmetry in the placement of the denticles
above the central fang (Fig. 21). The more posterior thoracic hooks
apparently have fewer small denticles. Notopodial hooks of the abdominal
region disposed in what might be called “uncinigerous” rows, from 14—41
per row, dorsal to the long capillary neurosetae (Fig. la). The hooks
are provided with rather broad long necks (Fig. 2u). Each abdominal
hook bears a single proximal, relatively large, central tooth surmounted
distally by many small teeth, in about 10-11 rows, with about 5 in
each row.
Remarks: The buff-colored sand tubes of M. africana are found at
the apical ends of the associated snails (Hydrobia plena). There is a
suggestion of a dorsal transverse groove between setigers 3 and 4 (Fig.
1d), and, in the case of 2 of the syntypes, there appears to be an internal
septum between setigers 5 and 6. Eggs were noted in one specimen and
these were restricted to the anterior end, extending between setigers 3-6
(Fig. 1b). Striated muscles were found associated with the rows of
abdominal hooks. There appears to be a single medial ventral blood
vessel in the region of the first setiger; this is formed by the union of
the lower ends of a vascular loop which seems to be closed dorsally
(Fig. lb, d). There are 2 pairs of brown granular structures, also in
the region of the first setiger, which may prove to be “ventral sacs”
functioning in tube formation. Spatulate setae may appear to be “spatu-
late” due to their usual orientation and, when differently disposed, may
appear to be limbate (cf. Fig. 2d, g, k, and Fig. 3p).
Brandtika new genus
Type-species: Brandtika asiatica new species. Gender: feminine.
Diagnosis: Fabriciine sabellids with 8 thoracic and 2 abdominal
eS
Fic. 1. Monroika africana (USNM 50913): a, Left lateral view of
entire specimen, branchial crown lacking (figure split between setigers
5 and 6 to fit page); b, left lateral view of second specimen showing
some internal structures (slightly twisted); c, ventral view of same
(setae omitted); d, dorsal view of same (VL= vascular loop, VV =
ventral vessel, VS = ventral sac (?), LM = longitudinal muscle bundle,
E = eggs, G = gut).
222, Proceedings of the Biological Society of Washington
eyequi] suo]
ATS “VE d A+ 0G “VE ATS “VE ALT ‘VG A+ PIL “VG ad ‘VE ¢ G PIV
aly Ve d AGS “VE AGE “AT “VS AST “VG AQT “VE 406 “VP d T Pav
d d ag ‘ag ‘vp =. dg “ag vy ag “ag “VG ag ‘aZ ‘ve ap “aE “VG d 8 4L
d d ap ‘ag‘ve dp “8g “VE aZ ‘ag “VE ag ‘aq ‘VE dz “AZ “VG d L UL
d az ‘aq ‘ve ag‘ae ‘vp ag ‘ap ‘vs az ‘ATI ‘VS az ‘az ‘vg ag ‘aT ‘ve dj 9 UL
d ag ‘og “aT ‘vp dp Op ‘ve = ap “09 “vg ag ‘Vy ag ‘og ‘ve dg ‘OF ‘ve ag ‘9g “VE S 4L
ag ‘OP ‘VP ap ‘og ‘vG ag‘oPp‘vp dg ‘og vg ag ‘9g “vg ay ‘Op ‘vp dg ‘0g ‘vp ag “og ‘vg y UL
ag ‘og ‘Vg ag ‘og ‘vg ag‘og ‘vp dg ‘0g ‘vg ag ‘Og ‘v7 ap ‘Op ‘vp ap “op “vg ag ‘op “aT ‘Vy € UL
ap ‘op ‘vg ac‘op‘al ‘vp ap‘os‘vp dp ‘Op ‘vg ag ‘og ‘aT ‘vq ag ‘0g ‘ag ‘vp ap “ap “VG dp ‘VP @ UL
ag ‘VP ap “VP ap ‘VE ap “V7 gg “VP dG “VE qZ “VG ag VP T UL
v € G I € G T (ad4quds T) zoquina
(sueuttoeds Gc) (sodAyuds ¢) Be ge aes
sodAjuAs-u0N T'LI'L'6861
HNWN@
‘uoumloods Y IF & UIOIZ POATIop
‘~ UUINjOO “QT pue G SIeBIES 10F soljUuy “(e[qeyUNoOouN = g ‘pammosqo = xX ‘syooy [eIpodojou [eurwiopqe = a ‘syooy
[erpodoimeu repnorAe ofoei0y, = a ‘ovjosojou oyejnyeds = 9 ‘ovj}osojou Areyideo oyequily y0Ys = a ‘ovzoSoO}OU Areyjideo
Vv) vupoufo py10lu0yy JO SuauToeds ur peAresqo sejes Fo sedAy Jo UOTNQIAsIp pue sIoqUINN “[ ATdv |]
Polychaete worms from southeast Asia 228
TABLE 2. Comparison of summarized numbers and distribution of types
of setae observed in specimens of (a) Monroika africana, (b) Brandtika
asiatica, and (c) Brandtika sp. (A = long limbate capillary notosetae;
B = short limbate capillary notosetae; C = spatulate notosetae; D =
thoracic avicular neuropodial avicular hooks; EK = abdominal notopodial
hooks; F = pilose neurosetae ).
Setiger -- Monroika africana -—~—~éBrandltika asiatica ——-Brandtika sp.
Th 1 3-5A, 2-4B 4A, 3B ?
Th 2 4-5A, 1-4B, 2-4C, 3-5D 4-5A, 2C, 45D e
Th3 4-5A, (1B), 3-5C, 3-5D 4-5A, 2C, 5D ?
Th4 45A, 2-5C, 3-5D 4-5A, 2-3C, 4D 6A, 2C, 7D
Th5 3-5A, (1B), 2-6C, 24D 5-6A, 2-3C, 3-4D 5A, 2C, 6D
Th6 2-4A, 14B, 2-3D 3-4A, 2-3C, 2F 3A, IC, 2F
Th 7 2-3A, 1-3B, 24D 2-4A, 2C, 1-2F ?
Th 8 24A, 2-3B, 3-5D 2-4A, 2-4C, 1-2F 5A, 4C, 4F
Abd 1 2-4A, (1B), 18-41E 3A, 17-20E 3A, 34E
Abd 2 2-3A, 144-21E 2A, 14-15E 3A, 24E
setigers; with long limbate capillary setae on all 10 setigers, with short
limbate capillary setae only on setiger 1, with spatulate setae on setigers
2-8, with neuropodial long-handled hooks on setigers 2-5 and short,
abruptly geniculate, pilose neurosetae on setigers 6—8, and with notopodial
long-necked uncini on setigers 9-10.
Etymology: It is with pleasure that I name this genus in honor of
Dr. R. A. M. Brandt, School of Tropical Medicine, Bangkok, Thailand,
in appreciation for the collecting of the numerous snails from Thailand
and Laos which provided me with these sabellids, as well as specimens
of Caobangia.
Remarks: Brandtika appears to be closely related to Monroika, as
regards the presence of similar thoracic notopodial and abdominal neuro-
podial hooks and the presence of thoracic spatulate setae. Brandtika
differs, however, in that spatulate notosetae are not replaced by short
limbate capillaries in the posterior thoracic region, whereas long-handled
avicular neuropodial hooks are replaced by short pilose neurosetae in
the posterior 3 thoracic setigers.
Brandtika asiatica, new species
Figure 3
Monroika sp. Jones, 1974: Fig. 5a.
Material examined: Laos: Mekong River at Ban Don Det, south of
Khong Island, Sithandone (14°07’N, 105°48’E), associated with the
224 Proceedings of the Biological Society of Washington
Fic. 2. Monroika africana (USNM 50913): a, Long limbate noto-
seta from first setiger; b, short limbate notoseta from same; c, three
neuropodial avicular hooks from setiger 2, viewed from above; d, spatulate
notoseta from setiger 3 (limbation not in profile); e, two neuropodial
hooks from same; f, long limbate notoseta from setiger 4; g, spatulate
notoseta from same; h, neuropodial hook from same; i, long limbate
notoseta from setiger 5 (limbation not in profile); j, short (?) limbate
notoseta from same; k, three spatulate notosetae from same; l, three
hooks from same (diagrams below right and left hooks show disposi-
tion of denticles); m, long limbate notoseta from setiger 6; n, two short
limbate notosetae from same; 0, neuropodial hook from same; p, short
limbate notoseta from setiger 7; q, neuropodial hook from same;
r, long limbate notoseta from setiger 8; s, short limbate notoseta from
Polychaete worms from southeast Asia 225
viviparid gastropod Mekongia jullieni (Deshayes) (USNM [Molluscs]
704235), R. A. M. Brandt, collector, 13 March 1968—holotype (USNM
50914), paratypes (USNM 50915), dried specimens; Mekong River at
Ban Seo, south of Khong Island, Sithandone (14°06’N, 105°51’E), asso-
ciated with the unionid bivalve Hyriopsis delaportei (Crosse and Fischer )
(USNM [Molluscs] 704236), R. A. M. Brandt, collector, 6 March 1968
—paratypes (USNM 50916), dried specimens.
Type-material: The holotype is 1.75 mm long, excluding the branchial
crown, and 0.13 mm wide at its widest part; the branchial crown is totally
dried. Paratypes are from 1.50-3.10 mm long, excluding the branchial
crown, and 0.14—0.27 mm wide; branchial crowns are 0.15-0.18 mm.
Description: The body is small, with 10 setigerous segments, 8 tho-
racic and 2 abdominal (Fig. 3a). The anterior end has a branchial
crown whose details can not be determined, due to the dried condition
of the specimens. A collar is present, and its margin is entire except for
a deep mid-dorsal cleft which is continuous with a dorsal groove extending
to about the level of setiger 2 (Fig. 3a).
The first setiger is provided only with unilimbate capillary notosetae
of 2 types, about 4 long ones (Fig. 3b) and 3 short ones (Fig. 3c;
Table 2). The following 7 thoracic setigers all bear 2-6 long limbate
capillary notosetae (Fig. 3f, i, 1, 0); the 2 abdominal setigers bear long
geniculate limbate capillary neurosetae (Fig. 3r, t). Only the first setiger
bears short limbate capillary setae; on setigers 2-8, these are replaced
by 2-4 spatulate notosetae (Fig. 3g, j, m, p).
There are 3-5 long-handled avicular neuropodial hooks on setigers
9-5. Each of these has a stout, sinuous manubrium with a large, single,
central fang surmounted by a series of smaller denticles (Fig. 3d, e,
h, u). An en face view reveals that there is a pronounced asymmetry in
the placement of the denticles above the central fang (Fig. 3d, e, wu).
Notopodial hooks of the abdominal region are in “uncinigerous’ rows,
from 14—20 per row, dorsal to the long geniculate capillary neurosetae.
The hooks are provided with long necks (Fig. 3s). Each abdominal
hook is provided with a single, proximal, relatively large, central tooth
surmounted distally by many small teeth in about 10 rows; the number
of denticles per row decreases distally from about 3 proximally (just
above the single tooth) to about 7-8 distally.
Unique pilose neurosetae appear on setigers 6-8, where they replace
the neuropodial avicular hooks of the more anterior thoracic segments.
They are short and sharply bent to nearly right angles, relative to the
slightly curved shaft; their distal ends present a pilose appearance
(Fig. 3k).
Remarks: Tubes of B. asiatica were present over several of the apical
&
same; t, three neuropodial hooks from same; u, abdominal notopodial
hook.
226 Proceedings of the Biological Society of Washington
Polychaete worms from southeast Asia 2251
whorls of Mekongia and formed extensive incrustations on the valves of
Hyriopsis; in both cases, they were associated with the aberrant sabel-
liform polychaete Caobangia brandti (Jones, in press). Two types of
granular dermal inclusions are present in the anterior thoracic region of
some specimens of B. asiatica.
Brandtika sp.
Figure 4
Material examined: Thailand: Salween River at Ban Chao Noi, Mae
Hong Son Province (approximately 18°26’N, 97°35’E), associated with
Brotia cf. baccata (Gould) (USNM [Molluscs] 420480b), H. M. Smith,
collector, 14 January 1933 (USNM 50917). Burma: Thoungyin River
[a tributary of the Salween], associated with paratype of Melania pago-
dula Gould [= Brotia pagodula (Gould)] (USNM [Molluscs] 611238)
(USNM 50918).
Fragments of from 1 to 3 setigers each were obtained from tubes
similar to those of B. asiatica. In only a few instances was it possible to
determine which setigers were being examined and then, only by making
the assumption that the types of setae and their linear distribution along
the body of these worms was the same as for B. asiatica. Thus, because
a posterior end consisted of 3 setigers, the terminal 2 of which had rows
of hooks and the other had short pilose setae, it was possible to decide that
these were setigers 8-10. Similarly, it was possible to determine that
another series of 3 setigers consisted of the setigers 4-6, because of the
shift from neuropodial avicular hooks to pilose setae between setigers
5 and 6. Unfortunately, not all of the setae encountered were oriented
so that they could be illustrated. However, no setae were found which
were radically different from those found in specimens of B. asiatica.
ێ
Fic. 3. Brandtika asiatica new genus, new species (a-t, USNM
50915; u, USNM 50916): a, Right dorsolateral view of entire specimen;
b, long limbate notoseta from first setiger; c, short limbate notoseta
from same; d, three neuropodial avicular hooks from setiger 2 (diagrams
show disposition of denticles); e, same, from setiger 3; f, long limbate
notoseta from setiger 5; g, two adjacent spatulate notosetae from same;
h, neuropodial hook from same; i, long limbate notoseta from setiger
6; j, two spatulate notosetae from same; k, three pilose neurosetae from
same; |, long limbate notoseta from setiger 7; m, two adjacent spatu-
late notosetae from same; n, two pilose neurosetae from same; o, long
limbate notoseta from setiger 8; p, three spatulate notosetae from same;
q, pilose neuroseta from same; r, slightly geniculate, long limbate seta
from setiger 9; s, two notopodial abdominal hooks from same; t,
geniculate long limbate neuroseta from setiger 10; u, thoracic avicular
hook. Scale A, a; scale B, b—u.
228 Proceedings of the Biological Society of Washington
[| ff
Fic. 4. Brandtika sp. (USNM 50917): a, Spatulate notoseta from
setiger 4; b, pilose neuroseta from setiger 6; c, long limbate notoseta
from setiger 8; d, two spatulate notosetae from same; e, two pilose
neurosetae from same; f, long limbate neuroseta from abdominal
setigers 9 or 10; g, two notopodial abdominal hooks from same.
0 0.05 mm
[Reed DS ESS [ea
Due to the lack of even one whole specimen, I do not feel that these
specimens should be considered to be conspecific with B. asiatica or
necessarily a different species from B. asiatica, even though they occur
in a quite different drainage system than B. asiatica. It should be noted
that the two different species of Caobangia occur in these same two
drainages, i.e., C. brandti in the Mekong, as well as the Chao Phraya
of Thailand, and C. smithi in the Salween (Jones, 1974). Thus, I
offer the following merely to add to our present knowledge of the genus
Brandtika.
Preliminary description: A fourth setiger included 6 long limbate
capillary notosetae, 2 spatulate notosetae (Fig. 4a), and 6 neuropodial
avicular hooks, all similar to their counterparts in B. asiatica. The ad-
jacent setiger 5 bore 5 long capillary notosetae, 2 spatulate notosetae,
and 6 neuropodial hooks; setiger 6 had 3 long capillary notosetae, 1
spatulate notoseta, and 2 pilose neurosetae (Fig. 4b). In the posterior
section mentioned above, setiger 8 had 5 long capillary notosetae (Fig.
4c), 4 spatulate notosetae (Fig. 4d), and 4 pilose neurosetae (Fig. 4e);
setiger 9 carried 3 long, slightly sinuous, limbate capillary neurosetae
(Fig. 4£) and about 34 long-necked notopodial hooks (Fig. 4g); setiger
10 was provided with 3 nearly straight capillary neurosetae and about
20 notopodial hooks.
Potential specific differences between these specimens and B. asiatica
Polychaete worms from southeast Asia 229
(Table 2) may lie in the number of neuropodial avicular hooks in the
anterior thorax, i.e., 6-7 hooks in setigers 4-5 in Brandtika sp. compared
with 3—4 hooks in the same setigers in B. asiatica, and in the number
of abdominal hooks on setigers 9 and 10, i.e., 24-24 in Brandtika sp. vs.
15-20 in B. asiatica, as well as in the shape of the abdominal capillary
setae (cf. Fig. 4f vs. Fig. 3r, t) and of the abdominal notopodial hooks
(cf. Fig. 4g vs. Fig. 3s). Confirmation or refutation of this possibility
must await the collection of better preserved material.
LITERATURE CITED
HarTMan, Ouca. 1951. Fabricinae (Feather-duster polychaetous an-
nelids ) in the Pacific. Pacific Sci., 5:379-391.
Jones, MEREDITH L. 1969. Boring of shell by Caobangia in freshwater
snails of southeast Asia. Amer. Zool., 9:829-835.
1974. On the Caobangiidae, a new family of the Polychaeta
with a redescription of Caobangia billeti Girard. Smithson.
Contrib. Zool., No. 175.
Monro, C. C. A. 1939. On a collection of Polychaeta from near the
mouth of the River Congo. Rev. Zool. Bot. Africaines, 32:
213-225.
WESENBERG-LUND, ELIsE. 1958. Lesser Antillean polychaetes, chiefly
from brackish water with a survey and bibliography of fresh
and brackish-water polychaetes. Stud. Fauna Curacao Other
Caribbean Isl. No. 30, pp. 1-41.
230 Proceedings of the Biological Society of Washington
x
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Vol. 87, No. 21, pp. 231-234 22, October 1974
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
CAUDAL SCALATION OF CENTRAL AMERICAN
CROCODYLUS
Cartes A. Ross AND FRANKLIN D. Ross
Bird and Mammal Laboratories, Bureau of Sport Fisheries
and Wildlife, National Museum of Natural History,
Washington, D.C. 20560, and Goldfarb Library, Brandeis
University, 415 South Street, Waltham, Massachusetts 02154
Two species of Central American crocodiles, Crocodylus
acutus Cuvier, 1807, and C. moreletii Duméril, 1851, have been
confused owing to variation in their presumed diagnostic char-
acters. The examination of a large number of live and pre-
served individuals shows that these species can be differenti-
ated on the basis of their caudal scalation. Specimens of C.
acutus were examined from Belice, Guatemala, Honduras,
Mexico, and Nicaragua; C. moreletii was examined from
throughout its range—Belice, Guatemala, and Mexico.
The presence of caudal irregularities in C. moreletii was
used by King and Brazaitis (1971) to aid in the identification
of commercial crocodilian skins. However, they failed spe-
cifically to note that caudal irregularities occur on some C.
acutus. In both species, one or more scales have been noted
between the caudal whorls in the proximal half of the tail
(Fig. 1). No more than three proximal irregularities occur
in C. acutus, as determined from the examination of 63 speci-
mens; each irregularity may consist of from one to three scales.
Furthermore, these irregularities are restricted to the lateral
surface of the tail (Table 1). Crocodylus moreletii also ex-
hibits proximal caudal irregularities; however, these are always
on, but not limited to, the ventral surface. These ventral sur-
face irregularities are variable. They may be either a single
scale, or two or more scales arranged laterally. Extensive,
21—Proc. Brou. Soc. WasuH., Vou. 87, 1974 (231)
232 Proceedings of the Biological Society of Washington
—
Oo
Fic. 1. Ventro-latero views of anterior caudal scalation of C. acutus
(A), and C. moreletti (B), showing caudal irregularities. Anterior
is to the right of the figure.
complex ventral irregularities have been noted in which the
normal number of proximal ventral whorls was nearly doubled.
Some specimens of C. moreletii exhibit lateral caudal irregu-
larities similar to those of C. acutus, but these are always ac-
companied by ventral irregularities.
No mention of proximal caudal irregularities was made in
the type descriptions of C. moreletii (Duméril and Duméril,
1851; A. Duméril, 1852). The second description (1852) was
accompanied by a plate which shows ventral proximal caudal
irregularities extending laterally. King and Brazaitis (1971)
state that C. moreletii exhibits a caudal irregularity 66 per
cent of the time. All the specimens we examined possessed
ventral proximal caudal irregularities and 87 per cent exhibited
irregularities extending laterally (Table 1). This condition is
visible in animals of all ages.
Caudal scalation of crocodiles 233
TaBLE 1. Numbers of Central American crocodiles exhibiting caudal
scalation irregularities.
Ventral
Sample extending
size None Lateral Ventral laterally
Crocodylus acutus 63 56 i 0 0
Crocodylus moreletii 111 0 many* 111 97
_ * Number not noted, however, always occurring in conjunction with ventral
irregularities.
As determined by our sample, C. acutus exhibits regular
caudal scalation 89 per cent of the time. Eleven per cent ex-
hibit proximal lateral caudal irregularities (Table 1), but no
ventral caudal irregularities have ever been noted.
ACKNOWLEDGMENTS
We wish to acknowledge the financial support of Robert L.
Whiston, Queensland, Australia; Dr. G. F. De Witte for the
loan of the co-types of Alligator lacordairei Preudhomme de
Borre (=C. acutus) housed in the Académie Royal de
Belgique, Professor Miguel Alvarez del Toro for permitting
the examination of live specimens in his charge at the Instituto
de Historia Natural, Tuxtla Gutierrez, Chiapas, Mexico. Spe-
cial gratitude is due the Division of Amphibians and Reptiles
of the National Museum of Natural History for providing re-
search space and encouragement. We would like to thank Dr.
George R. Zug for his critical reading of the manuscript and
advice.
LITERATURE CITED
DuMeErIL, C. anp A. Dumeri. 1851. Catalogue Méthodique de la
Collection des Reptiles. Paris. :1—224.
DumeriL, A. 1852. Descriptions des Reptiles Nouveux or Imparfait
Connus de la Collection du Muséum D’Historie Naturelle.
Archives de Muséum 6:209-588.
Kinc, F. W. anv P. Brazairtis. 1971. Species Identification of Com-
mercial Crocodilian Skins. Zoologica 56( 2 ):15-70.
934 Proceedings of the Biological Society of Washington
Vol. 87, No. 22, pp. 235-244 22 October 1974
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
A REDESCRIPTION OF HERMILIUS PYRIVENTRIS
HELLER (COPEPODA: CALIGOIDA) WITH THE
FIRST DESCRIPTION OF THE MALE
By Rocer F. CrEssEy
Smithsonian Institution, Washington, D.C. 20560
The genus Hermilius was described by Heller in 1868 to
accommodate a new species of caligid copepod collected from
the catfish Arius acutus in Java. Since then 3 additional
species have been described; H. longicornis Bassett-Smith 1898,
H. armatus Capart 1959, and H. youngi Kabata 1964. Kabata
suggested that H. armatus might be synonymous with H.
pyriventris. Examination of the material described below
and its comparison with Capart’s description of H. armatus
has led me to this conclusion also.
Several specimens of Hermilius pyriventris Heller were
sent to me by Dr. I. Paperna of the Hebrew University of
Jerusalem. Additional specimens were collected by the author
from the gills of catfish from India and the Philippines
contained in the fish collections of the Smithsonian Institution.
A review of the literature concerned with the genus
Hermilius revealed that no adequate description of the type-
species exists. Pillai (1961) described a collection containing
material which he identified as H. pyriformis but which
Kirtisinghe (1964) showed to be H. longicornis Bassett-Smith.
All drawings were made with the aid of a Wild Drawing
Tube.
All material has been deposited in the collections of the
Smithsonian Institution.
The illustrations were made by Hillary Boyle.
22—Proc. Briou. Soc. WasH., VoL. 87, 1974 (235 )
236 Proceedings of the Biological Society of Washington
Hermilius pyriventris Heller
Figures 1-21
Hermilius pyriventris Heller, 1868, p. 186.—Bassett-Smith, 1899, p. 445.
—Brian, 1924, p. 393.—Capart, 1959, p. 91.
Hermilius armatus Capart, 1959, p. 90 non H. pyriventris—Pillai,
1961 (= H. longicornis Bassett-Smith).
Material studied: 6 females from the gills of Netuma macrocephalus
and 11 females from the gills of N. thalassinus, both hosts collected
off the coast of Kenya. 16 females and 2 males from the gills of Arius
sp. collected at Vizakhapatnam, India. 11 females from the gills of
Tachysurus sp. (USNM 160551) from the Philippines.
Description: Female—Body form as in Figures 1 and 2. Measure-
ments (in mm) for total body length and width as below.
No. of Avg. Avg. Range of
specimens length width ength
N. thalassinus Vipingno, Kenya 5 2.05 0.56 1.84-2.38
N. thalassinus Diani, Kenya 3 5.40 1.33 5.03-5.77
N. macrocephalus Vipingno, Kenya 3 3.94 1.11 3.57-4.48
Trachysurus sp. Philippines 9 3.78 1.05 3.33-4.16
Arius sp. India 11 3.86 1.14 3.44-4.24
The specimens in the two collections from Kenya varied significantly
in size but no morphological differences could be found between them
—hence they are considered to be of the same species here. Measure-
ments below are from a specimen in the Vipingno collection.
Cephalon about one-half of total body length, folding laterally to
envelope that part of gill to which it is attached. (This modification
is not uncommon to parasitic copepods which attach to gill filaments,
Lernanthropus, Metataeniacanthus, and Alicaligus for example). Ceph-
alon supported by several heavily sclerotized rods making it rather
rigidly folded laterally and difficult to flatten for study. Genital
segment (Fig. 3) longer than wide (968 x 679), widest anteriorly.
Abdomen (Fig. 4) 2-segmented, segments 1364 and 124 in length
=
Fics. 1-5. Hermilius pyriventris Heller, female: 1, dorsal; 2, lateral;
3, genital segment and abdomen, ventral; 4, abdomen, ventral; 5,
caudal ramus, ventral.
Fics. 6-12. Hermilius pyriventris Heller, female, cont: 6, first
antenna; 7, second antenna; 8, mandible; 9, first maxilla; 10, second
maxilla; 11, maxilliped; 12, sternal furca.
Fics. 13-16. Hermilius pyriventris Heller, female, cont: 13, leg 1;
14, leg 2; 15, leg 3; 16, leg 4.
Redescription of a caligoid copepod Zon
238 Proceedings of the Biological Society of Washington
Redescription of a caligoid copepod 239
ee 7 77
240 Proceedings of the Biological Society of Washington
Ya:
Redescription of a caligoid copepod 241
respectively, each considerably wider than long. Caudal rami (Fig. 5)
longer than wide (59u x 47); each with 6 setae; terminal 3 short
and. pyriform; lateral and subterminal setae plumose, of about equal
length.
First antenna (Fig. 6) 2-segmented; basal segment 94 in length,
bearing many stout finely plumose setae; terminal segment bearing
12 slender naked setae and one aesthete. Second antenna (Fig. 7) 4-seg-
mented; segments measure 89u, 295u, 260u and 183 long respectively.
Second segment with fine striations along inner edge. Terminal claw
of 2 segments; first with short seta near base and accessory process
near inner distal corner, second recurved to complete claw. Mandible
(Fig. 8) of typical caligid type with 13 teeth. First maxilla (Fig. 9)
a long sclerotized, posteriorly directed, process with 2 short setae near
base; process situated lateral to and nearly as long as mouth tube.
Second maxilla (Fig. 10) small, 2-segmented; terminal segment
with short spinelike process near inner distal. corner and_ terminal,
finely serrate, spine. Maxilliped (Fig. 11) slender, 3-segmented, not
heavily sclerotized; second segment with short barblike process on inner
margin and short stout spine on inner distal comer, terminal segment
slightly recurved bearing basal seta and short accessory barb near tip.
Sternal furca (Fig. 12) weakly developed, tines spatulate.
Leg 1 (Fig. 13) basipod with patch of spinules and 2 plumose setae,
one inner and one on outer distal corner; exopod 2-segmented, first
segment with short spine on outer distal corner, second segment with
3 inner plumose setae and 3 terminal setae, each with inner hyaline
membrane; endopod reduced to short spinulose process. Leg 2 (Fig.
14) exopod 3-segmented, first segment with inner seta and short outer
distal spine, second segment with inner seta and longer outer distal
spine, third segment with 5 terminal to inner setae and 3 terminal spines,
outermost directed inwardly across other 2; endopod 3-segmented,
first segment with inner seta, second segment with 2 inner setae,
third segment with 6 setae, outer edges of last 2 segments with short
blunt spinules. Leg 3 (Fig. 15) exopod 3-segmented, first segment
with heavily sclerotized inwardly directed spine, second segment with
short inner seta and outer distal spine, third segment with 7 short
setae, second and third outer shorter and naked; endopod 3-segmented,
first segment lobate, covering part of second and third segments,
second segment short with inner seta, third segment with 6 setae.
Leg 4 (Fig. 16) basipod with outer short seta; exopod 2-segmented,
first segment with outer distal spine, second segment with 3 terminal
<
Fics. 17-22. Hermilius pyriventris Heller, male: 17, dorsal; 18,
genital segment and abdomen; 19, caudal ramus, ventral; 20, second
antenna; 21, leg 1; 22, leg 4.
242 Proceedings of the Biological Society of Washington
spines, innermost naked. Leg 5 absent. Leg 6 represented by 2 short
plumose setae lateral to area of egg string attachment.
Egg strings of usual caligoid form.
Male.—Body form as in Figure 17. Total length and greatest
width 1.53 x 0.68 mm. Cephalon about one-half of total body length.
Cephalon folded laterally as in female to envelope gill filaments.
Genital segment (Fig. 18) longer than wide (40lu x 212), spermato-
phores easily visible within. Abdomen 1-segmented, longer than wide
(1244 x 100”). Caudal ramus (Fig. 19) bearing 6 plumose setae,
2 lateral, 1 subterminal, and 3 terminal (longest seta 472u long);
each ramus longer than wide (59u xX 47).
Cephalic appendages as in female except second antenna. Second
antenna (Fig. 20) 3-segmented (4 in female), without accessory
process on claw.
Leg 1 (Fig. 21) similar to female except endopod relatively larger
and 3 terminal exopod spines relatively shorter. Legs 2 and 3 as in
female. Leg 4 (Fig. 22) similar to female except terminal spines of
male relatively longer and without fringes. Leg 5 represented by 2
setae near midlateral margin of genital segment. Leg 6 represented
by 2 setae at posterior corners of genital segment.
Remarks: H. pyriventris females can be easily separated from
H. longicornis and H. youngi by the nature of the second antenna
(4-segmented in pyriventris and 3-segmented in the other 2). The
pyriform terminal setae of the caudal rami of pyriventris further
separates them from females of longicornis and youngi. So far the
genus has only been reported from species of marine catfish (Arridae)
and has not been collected from waters adjacent to North and South
America. I have examined many catfish from the Gulf of Mexico
without finding Hermilius.
LITERATURE CITED
BAssETT-SmitTH, P. W. 1898. Further new parasitic copepods from
fish in the Indo-tropical region. Ann. Mag. Nat. Hist. 2(7):
77-98.
1899. A systematic description of parasitic copepoda found
on fishes, with an enumeration of the known species. Proc.
Zool. Soc. London, pt. 2. 438-507.
Brian, A. 1924. Parasitologia mauritanica, 1: Copepoda. Bull.
Com. Etud. Hist. Sci. Afrique Occid. Francaise, no. d, Sept.,
365-445.
Capart, A. 1959. Copépodes parasites. Fasc. 5 of vol. 3 in Expédi-
tion océanographique belge dans les eaux cétiéres africaines
de Atlantique Sud (1948-1949). Inst. Roy. des Sci. Nat. de
Belg. 57-126.
HELLER, C. 1868. Crusteceen. No. 8 in vol. 2 of Zoologischer Theil
in Reise der Oesterreichischen Fregatte Novara. ... , 1-280.
Redescription of a caligoid copepod 243
Kapata, Z. 1964. Copepoda parasitic on Australian fishes. I. Hermi-
lius youngi sp. nov. (Caligidae). Ann. Mag. Nat. Hist.
7(13) :609-618.
KirTIsINGHE, P. 1964. A review of the parasitic copepods of fish
recorded from Ceylon with descriptions of additional forms.
Bull. Fish. Res. Sta., Ceylon. 17(1):45-132.
Pruuar, N. K. 1961. Copepods parasitic on South Indian fishes.
Pt. 1. Caligidae. Bull. Cent. Res. Inst., Univ. of Kerala,
8:87-130.
244 Proceedings of the Biological Society of Washington
i, Ae
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Vol. 87, No. 23, pp. 245-256 22 October 1974
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
A NEW SPECIES OF HERMIT CRAB OF THE
GENUS NEMATOPAGURUS (CRUSTACEA:
DECAPODA: PAGURIDAE) FROM
HAWAII
By Patsy A. McLAUGHLIN AND JULIE H. Brock
Rosenstiel School of Marine and Atmospheric Science,
University of Miami, Miami, Florida 33149, and Department
of Zoology, University of Hawaii, Honolulu, Hawaii 96822
The genus Nematopagurus was established by A. Milne-
Edwards and Bouvier (1892) for Nematopagurus longicornis
A. Milne-Edwards and Bouvier from the western Mediter-
ranean and eastern Atlantic from Spain to Cape Verde
(A. Milne-Edwards and Bouvier, 1892; 1899, 1900). Subse-
quently, Alcock (1905) assigned Catapagurus muricatus
Henderson, 1896 from Ceylon to Nematopagurus and de-
scribed four additional species from several localities in
the northern Indian Ocean. De Saint Laurent (1968) pro-
posed, without discussion, that Melin’s (1939) Eupagurus
(Catapagurus) vallatus from Japan also should be assigned
to Nematopagurus. Recently, Lewinsohn (1969) reported
N. muricatus and N. squamichelis Alcock from the Gulf of
Aqaba, Red Sea, and described another new species, Nemato-
pagurus diadema Lewinsohn from the same area. The dis-
covery of a new species of this genus from Hawaii is the first
record of Nematopagurus in this easternmost part of the
Indo-west-Pacific Ocean.
This species, Nematopagurus spinulosensoris n. sp., de-
scribed herein, is particularly noteworthy, not only as indica-
tive of the considerable extension of the range of Nemato-
23—Proc. Bron. Soc. WasuH., Vou. 87, 1974 (245 )
246 Proceedings of the Biological Society of Washington
pagurus, but because, as its name implies, it possesses certain
unique sensory structures, unknown in other pagurids.
The holotype and certain paratypes are deposited in the
National Museum of Natural History, Smithsonian Institution
(USNM). Additional paratypes are deposited as follows:
Bernice P. Bishop Museum (BPBM), Honolulu, Hawaii;
Allan Hancock Foundation (AHF), University of Southern
California; and Rijksmuseum van Natuurlijke Historie, Leiden,
The Netherlands (RMNH). A single measurement, the shield
length (SL) has been made for each specimen.
The authors are indebted to Dr. Paul J. Struhsaker, National
Marine Fisheries Center, Honolulu, Hawaii, for providing
specimens from the collections of the National Marine
Fisheries Service; to Dr. Chanan lLewinsohn, Tel-Aviv
University, Tel-Aviv, Israel, for providing comparative
specimens of Nematopagurus muricatus (Henderson); and
particularly to Dr. Robin M. Overstreet, Gulf Coast Research
Laboratory, Ocean Springs, Mississippi, for his serial sections
and preliminary interpretation of the sensory structures of
this species. This research has been supported, in part, by
Research Grant GB 40062 from the National Science Founda-
tion and by a Hawaii Biomedical Support Grant 1970/1971.
Nematopagurus spinulosensoris new species
Figures 1-3
Holotype: 6 (SL = 8.6 mm), NOAA station TC-33-15, 21°02’N,
156°46’W, 31 October 1967, 241-254 m, USNM 149299.
Paratypes: 16 (SL =5.2 mm), off Oahu, 23 September 1971, 382 m,
RMNH; 64, (SW = 5.1-7.4 mm), 29, 1 ovigerous (SL = 5.2, 6.5
mm), NOAA station TC-33-15, 21°02’N, 156°46’W, 31 October 1967,
241-254 m, USNM, BPBM, AHF, RMNH.
Type-Locality: Off Molokai, Hawaii, 21°02’N, 156°46’W.
Diagnosis: Chelae and carpi of both chelipeds, each with sensory-
modified spines on dorsal surface. Right chela with several irregular
rows of spines on dorsal surface. Carpi of ambulatory legs each with
row of strong spines dorsally.
Description: Shield (Fig. la) longer than broad; anterolateral
margins sloping or slightly terraced; anterior margin between rostrum
and lateral projections concave; posterior margin truncate or roundly
truncate; dorsal surface with numerous tufts of stiff setae; anterolateral
angle produced, subacute or acute. Rostrum usually obtusely rounded,
A new hermit crab from Hawaii 247
(ie oe ee
AHIR emma
Fic. 1. Nematopagurus spinulosensoris n. sp., 6 paratype: a, shield;
b, left chela and carpus (dorsal view)
view); d, 2nd left dactyl and propodus (mesial view); e, spine of
chela with sensory process. Scales equal 3 mm (a), 5 mm (b-—-d) and
0.5 mm (e).
; c, 2nd left pereiopod (lateral
248 Proceedings of the Biological Society of Washington
occasionally obtusely triangular; often not exceeding lateral projections,
occasionally slightly exceeding lateral projections; usually with few
moderately long setae. Lateral projections prominent, obtusely
triangular, each with prominent submarginal spine.
Ocular peduncles moderately short, two-thirds to three-fourths
length of shield; dorsal and dorsomesial surfaces frequently with
few tufts of setae; corneae usually strongly dilated. Ocular acicles
acutely triangular, moderately slender, with prominent longitudinal
furrow; terminating acutely or subacutely, and with very strong sub-
marginal spine; separated basally by three-fourths to entire basal
width of one acicle.
Antennular peduncles moderately long, exceeding ocular peduncles
by one-half to two-thirds length of ultimate segment; ultimate segment
with longitudinal row of setae on dorsal surface in distal half, occa-
sionally on distal two-thirds to four-fifths; penultimate segment with
few scattered setae; basal segment with small spine or spinule at
ventrodistal margin, partially obscured by tuft of long stiff setae,
lateral face proximally, with very strong, slender, acute spine.
Antennal peduncles moderately short, exceeding ocular peduncles
by one-fifth to one-half length of ultimate segment; with supernumerary
segmentation (c.f. McLaughlin, 1974). Fifth and fourth segments
with numerous tufts of moderately long setae. Third segment with small
to moderately strong spine at ventrodistal angle, partially obscured
by tuft of very long stiff setae. Second segment with dorsolateral
distal angle produced, terminating in strong simple or bifid spine,
lateral margin occasionally with small spine distally, mesial face
with tufts of long stiff setae and occasionally also with 1 or 2 small
spines; dorsomesial distal angle with 1, occasionally 2, small to
moderately strong spines, mesial face with long stiff setae. First
segment with small to moderately strong spine on lateral face distally;
ventrodistal margin with 1-5 small to moderately strong spines.
Antennal acicle moderately long, usually reaching beyond proximal
half of ultimate peduncular segment; terminating in acute or blunt
spine; mesial margin with tufts of long stiff setae and occasionally
with 1 or 2 small spines. Antennal flagella long, overreaching tip of
right cheliped; occasionally few articles each with 1 or 2 very short
setae or bristles.
Mandible without distinguishing characters. Mazxillule (Fig. 3a)
with proximal endite subquadrate; endopodite with 1 moderately long
=>
Fic. 2. Nematopagurus spinulosensoris n. sp., a-e, & paratype:
a, right chela and carpus (dorsal view); b, 3rd left pereiopod (lateral
view); c, 3rd left dactyl and propodus (mesial view); d, 4th right
pereiopod (lateral view); e, coxae of 5th pereiopods with sexual
A new hermit crab from Hawaii 249
po 1 tq
|}__1—_—_e,f
tubes (ventral view). f, @ paratype, coxae of 5th pereiopods and
paired gonopods of Ist abdominal segment (ventral view). Scales
equal 5 mm (a—d) and 1 mm (e, f).
250 Proceedings of the Biological Society of Washington
bristle on well developed internal lobe, external lobe vestigial or
absent. Maxilla (Fig. 3b) with moderately broad endopodite usually
equalling scaphognathite in distal extension. First maxilla (Fig. 3c)
with basal segment of exopodite very slender. Second mazxilliped
(Fig. 3d) with basis-ischium fusion complete. Third maxilliped
(Fig. 3e) with basis-ischium incomplete; basis usually unarmed;
ischium with crista dentata well developed, with accessory tooth;
merus with strong spine on dorsodistal margin, ventrodistal angle
with cluster of denticulate bristles. Sternite of third maxillipeds
unarmed, sparsely setose.
Chelipeds subequal, right usually somewhat larger than left; chelae
and carpi of both chelipeds with numerous sensory-modified spines
(Figs. 1b, e, 2a) on dorsal surfaces. Right cheliped (Fig. 2a) elongate,
moderately slender. Dactyl moderately short, slightly shorter than or
equalling length of palm; cutting edge with row of calcareous teeth
and usually with few comeous teeth distally, terminating in small
corneous claw; slightly overlapped by fixed finger; dorsal surface
generally flattened, with few tufts of setae, dorsomesial margin, or
dorsal surface mesially, usually slightly elevated and with irregular
longitudinal row of unmodified small spines or tubercles, frequently
becoming obsolete distally; mesial face with scattered tubercles or
spines on proximal third and scattered tufts of setae; ventral surface
usually with 3 or 4 rows of tufts of stiff bristles. Palm slender,
moderately long, two-thirds to four-fifths length of carpus; dorsomesial
margin with irregular single or double row of usually unmodified
moderately strong spines, and tufts of long stiff setae; dorsal surface
with several irregular rows of usually modified spines, extending onto
fixed finger proximally, and scattered tufts of setae; dorsolateral margin
with single or double row of moderately strong, usually modified
spines, extending onto fixed finger as single row of blunt unmodified
spines or tubercles, decreasing in size distally; lateral face frequently
weakly tuberculate and with scattered tufts of long stiff setae; ventral
surface usually tuberculate, particularly mesially, with scattered
tufts of long stiff setae; mesial face with 1 or 2 irregular rows of
small tubercles or unmodified spines dorsally and tufts of stiff setae.
Carpus moderately long, usually equalling or slightly exceeding length
of merus; dorsomesial margin with row of very strong, slender, often
curved or hooked, unmodified spines and tufts of long setae; dorsal
surface with very irregular rows of moderately strong, usually
modified spines, more numerous distally, and scattered tufts of long
stiff setae; dorsolateral margin not noticeably delimited, lateral face
often with low protuberances and tufts of long stiff setae dorsally
and distally, occasionally with moderately strong spines, laterodistal
margin with acute spine; ventral surface usually somewhat tuberculate
and with numerous tufts of long stiff setae; mesial face with numerous
low tubercles or ridges and tufts of stiff setae. Merus_ sub-
A new hermit crab from Hawaii 251
triangular; dorsal margin with irregular row of transverse ridges,
often becoming somewhat spinulose distally, and with tufts of long
stiff setae, distal margin usually with 2 or 3 strong acute spines;
lateral face with scattered tufts of long stiff setae, ventrolateral margin
with row of strong, acute, often curved or hooked spines; mesial face
usually setose, distal margin and ventromesial face distally with
few, small, usually acute spines; ventral surface usually with low
tubercles or ridges and tufts of long stiff setae, occasionally with
few moderately strong spines. Ischium with strong acute spine at
ventrolateral distal angle; ventral, lateral and mesial surfaces setose,
ventromesial margin often with few small spines. Coxa with clump
of long stiff setae at ventromesial distal angle, 1-3 small tubercles
or spinules on ventroproximal angle, ventrolateral margin occasionally
with few small spines or spinules, ventrolateral distal angle usually
with 1 or 2 acute spines.
Left cheliped (Fig. 1b) moderately long, usually reaching beyond
proximal half of dactyl of right; moderately slender, fingers arched
or somewhat depressed distally. Dactyl moderately long, one and
one-half to twice length of palm; cutting edge with row of small
corneous teeth, terminating in small cormneous claw; slightly over-
lapped by fixed finger; dorsal surface somewhat convex, with short row
of small unmodified spines or spinulose tubercles usually in midline
or occasionally on dorsomesial margin proximally and with 2 or 3
rows of tufts of long stiff setae; mesial face slightly spinulose or
tuberculate proximally and with tufts of long stiff setae ventrally
and distally; ventral surface with 2 or 3 rows of stiff settae. Palm
moderately long, one-third to one-half length of carpus; dorsomesial
margin with single or double row of moderately small, frequently
modified, acute spines; dorsal surface somewhat convex, midline with
2 or 3 irregular rows of moderately strong, usually modified spines
extending onto fixed finger as single row proximally, usually with
1 or 2 irregular rows of modified spines laterally and mesially, occa-
sionally only with scattered spines; dorsolateral margin with double or
triple row of small modified spines proximally becoming single row
of small unmodified spines or spinulose or blunt tubercles on fixed
finger distally, and with long stiff setae; lateral, mesial and ventral
surfaces usually somewhat tuberculate and with tufts of long stiff
setae. Carpus long, equalling or exceeding length of merus; dorso-
mesial margin with row of strong, slender, often curved, usually
unmodified spines; dorsal surface with 2 or 3 irregular rows of moderately
strong modified spines proximally, tending to cluster distally, distal
margin occasionally with 1 or 2 spines; dorsolateral margin with
single or double row usually modified spines and tufts of long stiff
setae; lateral face with scattered modified spines in dorsal half, tufts
of long stiff setae ventrally, distal margin ventrally with moderately
strong acute spine; mesial and ventral surfaces frequently with low
252 Proceedings of the Biological Society of Washington
TINT
Z
e Y
C YZ
Fic. 3. Nematopagurus spinulosensoris n. sp., ¢ paratype, a-e,
mouthparts (left, internal view): a, maxillule; b, maxilla; c, Ist
maxilliped; d, 2nd maxilliped; e, 3rd maxilliped. f, sternite of 3rd
pereiopods; g, telson. Scale equals 1 mm.
A new hermit crab from Hawaii 253
protuberances or ridges and tufts of long stiff setae. Merus moderately
long, subtriangular; dorsal surface with low transverse ridges, often
becoming spinulose distally, and tufts of stiff setae, distal margin
with 1-3 acute spines; lateral and mesial faces with low transverse
ridges and tufts of long stiff setae; ventromesial and _ ventrolateral
margins each with row of strong, acute, sometimes curved spines.
Ischium with row of small spines and tufts of long setae on ventro-
mesial margin, ventrolateral distal angle with acute spine and tuft
of long setae. Coxa with clump of long stiff setae at ventromesial
distal angle and often 1 or 2 acute spines at ventrolateral distal angle.
Second (Fig. lc, d) and third (Fig. 2b, c) pereiopods usually
overreaching right cheliped, right slightly longer than left, generally
similar in structure and armature. Dactyls long, slender, one and
one-fourth to one and one-half length of propodi; in lateral view,
slightly curved ventrally; in dorsal view, somewhat twisted; each
terminating in strong corneous claw; dorsal surfaces each with row
of strong corneous spines, increasing in size distally, and tufts of
long stiff setae; mesial faces each with faint longitudinal sulcus
proximally flanked by 1 or 2 rows of corneous spinules above and few
corneous spinules below; lateral faces each sometimes with faint longi-
tudinal sulcus proximally and usually with 1 or 2 rows of tufts of setae;
ventral surfaces each with row of strong corneous spines increasing in
size distally. Propodi moderately long, one-third to twice length of carpi;
dorsal surfaces each with irregular row of transverse ridges and tufts of
long stiff setae; mesial and lateral faces usually with 1 or 2 rows of tufts
of setae; ventral surfaces each with 1 or 2 rows of tufts of long stiff
setae (P:) or row of small corneous spinules and tufts of long setae
(Ps). Carpi two-thirds to three-fourths length of meri; dorsal surfaces
each with row of strong spines; mesial and ventral surfaces each with
few scattered tufts of long setae; lateral faces each with 1 or 2 rows
of stiff setae. Meri laterally compressed; dorsal surfaces each with
row of transverse protuberances or ridges and long stiff setae; lateral
and mesial faces usually with few tufts of setae, lateral distal margin
(P2) usually with strong acute spine ventrally; ventral surface usually
with 1 to several small to moderately strong spines and tufts of setae
(P2) or row of transverse ridges and tufts of long setae (Ps). Ischia
each usually with row of long setae on ventral margin and row of
small spines (Pe left). Coxae each with row of tufts of long setae
on ventromesial and ventrolateral margins, ventral surface distally also
with long setae. Sternite of third pereiopods (Fig. 3f) semisubcir-
cular, anterior margin with long stiff setae.
Fourth pereiopods (Fig. 2d) subchelate; without apparent pre-
ungual process; propodal rasp of single row of comeous scales or
teeth.
Fifth pereiopods weakly chelate.
Males with well developed, elongate, filiform sexual tube on right
coxa of fifth pereiopods (Fig. 2e) left with vas deferens usually
254 Proceedings of the Biological Society of Washington
slightly protruded. Pleopods unpaired, pl:—pl; with exopodites well
developed; endopodites moderately well developed, smallest on pls.
Females with paired gonopores; first pleopods (Fig. 2f) paired,
pl.-pl., unpaired, with exopodite moderately well developed, endopodite
reduced.
Telson (Fig. 3g) with posterior lobes usually somewhat asymmetrical,
left usually slightly larger than right, subtriangular or subquadrate;
separated by very shallow median cleft; terminal and usually also
lateral margins weakly calcified, terminal margins somewhat oblique,
each with numerous small calcareous spines marginally and _ several
stronger calcareous acute or blunt spines submarginally; lateral margins
unarmed or occasionally each with row of small calcareous spines or
spinules. Anterior lobes unarmed.
Color: In life: Chelipeds and ambulatory legs generally vivid
salmon-pink, bordering on iridescent; antennal flagella bright yellow.
In preservative: Shield pale orange or straw-colored; ocular peduncles
light orange with dark orange ring proximally. Chelipeds very
pale orange with white spines; carpi with darker red-orange proximally
and ventrally. Ambulatory legs pale orange with lighter longitudinal
stripes on dactyls and propodi; carpi pale orange with darker red-orange
proximally; meri pale orange and white. Color fading in time to
straw-color.
Distribution: Known only from the Hawaiian Islands, 180-250
meters.
Etymology: The specific name, spinulosensoris, is derived from
the diminutive of the Latin, spina, meaning spine, and sensor, meaning
pertaining to sense, and depicts the sensory-modified spines of the
chelipeds.
Remarks: As previously indicated, N. spinulosensoris is particularly
distinctive in the development of sensory structures on the majority
of the spines on the dorsal surfaces of the chelae and carpi of the
chelipeds. Although presumed sensory structures are known from
several pagurid genera (cf. De Saint Laurent, 1970; McLaughlin,
1974) none of these structures have been asssociated with modifications
of the spines. In this species these structures appear as chitinous
“tear-drop” like processes, generally on the proximal faces of the
spines. Preliminary serial sections have confirmed the sensory character
of these processes; however, their fine structure is still under study
and will be described in a subsequent report.
N. spinulosensoris can be distinguished from all other species of
the genus particularly by the presence of sensory-modified spines
on the dorsal surfaces of the chelae and carpi of the chelipeds. In
addition, the presence of several irregular rows of spines on the dorsal
surface of the right chela distinguishes this species from all others
except N. muricatus; however, the presence of a row of spines on
the dorsal surface of the carpus of each ambulatory leg immediately
distinguishes N. spinulosensoris from the latter species.
A new hermit crab from Hawaii 255
LITERATURE CITED
Aucocx, A. 1905. Catalogue of the Indian decapod Crustacea in
the collection of the Indian Museum. Part II. Anomura.
Fasc. 1. Pagurides. i-xi:197 pp. Calcutta.
HENDERSON, J. R. 1896. Report on the Paguridae from the H. M.
Indian Survey steamer “Investigator,” Commander C. F.
Oldham, R.N. commanding. Series II, no. 24. Jour. Asiatic
Soc. Bengal, 65(2):516-536.
LEWINSOHN, Cu. 1969. Die Anomuren des Roten Meeres (Crustacea
Decapoda: Paguridea, Galathedeidea, Hippidea). Zool.
Verhandl., No. 104:1-214.
McLaucuuin, P. A. 1974. The Hermit Crabs (Crustacea: Decapoda:
Paguridea) of Northwestem North America. Zool. Verhandl.
No. 130:1-396.
Metin, G. 1939. Paguriden und Galatheiden von Prof. Sixten
Bocks Expedition nach den Bonin-Inseln 1914. K. Svenska
Vetenskapsakad. Handl., 18(2):1-119.
MiILNE-EDwarps, A. AND E. L. Bouvier. 1892. Observations
préliminaires sur les Paguriens recueillis par les Expéditions
francaises du “Travailleur” et du “Talisman.” Ann. Sci.
Nat. Zool. (7)13:185-226.
—. 1899. Crustacés Décapodes provenant des Campagnes de
l’“Hirondelle” (Supplément) et de la “Princesse Alice”
(1891-1897). Rés. Camp. Monaco, 13:1-106.
——. 1900. Crustacés Décapodes. 1: Brachyures et Anomoures.
Expéditions scientif. “Travailleur’ et “Talisman,” pp. 1-396.
SAINT LAURENT, M. bE. 1968. Revision des genres Catapaguroides
et Cestopagurus et description de quatres genres nouveaux.
I. Catapaguroides A. Milne-Edwards et Bouvier et Deca-
phyllus nov. gen. (Crustacés Décapodes Paguridae). Bull.
Mus. Natl. Hist. Nat. Paris, (2) 39 (5,6):923-954, 1100-1119.
——. 1970. Révision des genres Catapaguroides et Cestopagurus
et description de quatre genres nouveaux. IV. Solenopagurus
De Saint Laurent (Crustacés Décapodes Paguridae). Bull.
Mus. Natl. Hist. Nat. Paris (2) 41 (6):1448-1458.
956 Proceedings of the Biological Society of Washington
Vol. 87, No. 24, pp. 257-264 22, October 1974
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
WHAT IS ALLOPERLA IMBECILLA (SAY)?
DESIGNATION OF A NEOTYPE, AND A
NEW ALLOPERLA FROM EASTERN
NORTH AMERICA (PLECOPTERA:
CHLOROPERLIDAE )
By RicnHarp W. BAUMANN
Smithsonian Institution, Washington, D.C. 20560
Alloperla imbecilla (Say) is the type-species of the genus
Alloperla Banks (1906). It was described by Thomas Say
from the Ohio River at Cincinnati but the description is
very brief and does not include anything about the genitalia.
Needham and Claassen (1925) redescribed what they thought
was this species using specimens from New York because
no type-specimen was available. Their description and figures
of male and female genitalia served to fix the concept of
A. imbecilla until now.
Recently while studying Alloperla specimens at the National
Museum of Natural History it became apparent that two
species were included under the name Alloperla imbecilla
(Say). Since no type-specimen was known, as complete
a study as possible was undertaken using most existing
specimens. The results obtained indicate that a type-specimen
does not exist and that the true A. imbecilla is restricted to
the Ohio River drainage and nearby localities along the
Allegheny Front. A second, previously undescribed species,
is distributed all along the eastern portion of North America
from Canada to the southeastern United States.
I am indebted to the following people for helping me with
this study: Dr. Edwin F. Cook, University of Minnesota
(UM); Dr. Paul H. Freytag, University of Kentucky (UK);
24——Proc. Biou. Soc. WasH., VoL. 87, 1974 (25170)
258 Proceedings of the Biological Society of Washington
Dr. Arden R. Gaufin, University of Utah (UU); Dr. Peter P.
Harper, University of Montreal (UMC); Dr. Robert L. Jeanne,
Boston University (BU); Mr. J. E. H. Martin, Canadian
National Collection (CNC); Dr. Charles H. Nelson, Univer-
sity of Tennessee, Chattanooga (CHN); Dr. L. L. Pechuman,
Cornell University (CU); Dr. Selwyn S. Roback, Academy
of Natural Sciences, Philadelphia (ANSP); Mr. Martin A.
Tkac, Kent State University; Dr. F. Christian Thompson,
American Museum of Natural History (AMNH); Dr. Charles
A. Triplehorn, Ohio State University (OSU); Dr. George E.
Wallace, Carnegie Museum (CM); Mr. Donald W. Webb,
Illinois Natural History Survey (INHS); Ms. Janice C. White,
Museum of Comparative Zoology (MCZ); Dr. Glenn B.
Wiggins, Royal Ontario Museum (ROM). Special thanks
are given to Dr. Freytag and Dr. Triplehorn for allowing the
type-specimens to be deposited at the National Museum of
Natural History (USNM).
Alloperla imbecilla (Say)
Figures 1-6
Sialis imbecilla Say, 1823:165.
Perla imbecilla, Hagen, 1861:31.
Alloperla imbecilla, Banks, 1906:175; 1907:13 (in part).—Claassen,
1940:186 (in part) —Walker, 1947:1385.—Gaufin, 1956:323; 1964:
40 (in part).—lIllies, 1966:432 (in part).
Male: Macropterous. Length of forewings 7.5-8.5 mm; length of
body 7-8 mm. General color light pale green (dirty white in alcohol);
legs yellow; antennae brown; cerci yellow; head and pronotum yellow
brown, eyes and ocelli prominent chestnut brown. Head as wide as
prothorax; ocellar triangle equilateral; posterior ocelli slightly closer
to eyes than to each other. Pronotum wider than long, with very
faint rugose markings. Abdomen without dark dorsal markings;
abdominal segments covered by dense hairs. Epiproct long slender
and lightly sclerotized, basal %3 covered with hairs; dorsal aspect
broadest at base and apex, narrowest at midline, lateral margins of
apex serrated and heavily sclerotized, tip bearing numerous stout
hairs which appear as a single spine under low magnification (Figs.
2 & 3); lateral aspect broad at base, tapering gradually toward apex,
apical portion enlarged, bearing sclerotized serrations on dorsal margin,
stout hairs plainly visible at tip (Figs. 1 & 4).
Female: Macropterous. Length of forewings 8.5-9.5 mm; length
of body 8-9 mm. Color and general morphology similar to male.
Alloperla imbecilla and relative 259
Fics. 1-6. Alloperla imbecilla (Say): 1, male terminalia, lateral;
2, male terminalia, dorsal; 3, epiproct, dorsal; 4, epiproct, lateral;
5, female terminalia, ventral; 6, female terminalia, lateral. Fics. 7-10.
Alloperla atlantica sp. nov.: 7, male terminalia, lateral; 8, male
terminalia, dorsal; 9, epiproct, dorsal; 10, epiproct, lateral.
Subgenital plate formed from posterior median portion of seventh
sternum, base broad, 1% width of sternum 8, tapering abruptly to
narrowly rounded apex, extending over stemum 8 beyond midline
(Fig. 5); lateral aspect slightly broader at base, width nearly constant
to rounded tip, appearing as stout fingerlike projection (Fig. 6).
Material; Neotype, male, Ohio, Adams Co., Lower Twin Creek,
Vastine Run, 10-V-1963, F. J. Moore, USNM #73053). Additional
260 Proceedings of the Biological Society of Washington
confirmed records: New York: Tompkins Co., Newfield, 5-VI-1923,
P, 16 (CU). Onto: Adams Co., same data as neotype, 2¢ ¢ (OSU)
(UK); Geauga Co., Stebbins Gulch, 20 to 27-V-1972, M. A. Tkac,
12346, 11992 (USNM). Vimermta: Bath Co., Blowing Spring
Cmpg., 8 miles W. Warm Springs, 18 to 20-V-1963, Flint & Field,
13, 19 (USNM). West VircintA: Summers Co., Bluestone State
Park, near Hinton, 21 to 22-V-1963, Flint & Field, 1¢, 42 2 (USNM).
Alloperla atlantica, new species
Figures 7-10
Alloperla imbecilla, Banks, 1907:13 (in part); 1908:150.—Needham
and Claassen, 1925:125.—Procter, 1938:53.—Claassen, 1940:186.
—Harden, 1942:319.—Procter, 1946:54—Sprules, 1948:74.—Ricker,
1948:411.—Harden and Mickel, 1952:59.—Brinck, 1958:56.—Gaufin,
1964:40 (in part).—Illies, 1966:432 (in part ).—Hitchcock, 1968:43.
—Ricker, et al. 1968:1116.
Male: Macropterous. Length of forewings 7.5-8.5 mm; length of
body 6-8 mm. General color and morphology similar to A. imbecilla.
Epiproct long slender and lightly sclerotized, basal 24 covered with
hairs; dorsal aspect broadest at base, tapering gradually to pointed
apex, tip rounded and naked (Figs. 8 & 9); lateral aspect tapering
from broad base to pointed tip, apical % naked and often slightly bent
downward (Figs. 7 & 10).
Female: Macropterous. Length of forewings 8-9 mm; length of
body 8-9.5 mm. Color and general morphology similar to male.
Subgenital plate formed from posterior median portion of seventh
sternum, base broad, 4% width of sternum 8, tapering abruptly to narrowly
rounded apex, extending over sternum 8 nearly to posterior margin;
lateral aspect slightly broader at base, width nearly constant to nar-
rowly rounded tip, appearing as stout fingerlike projection with slightly
pointed tip.
Material: Holotype, male and allotype, Maryland, Frederick Co.,
Fishing Creek, Mountaindale, 4-VI-1958, P. H. Freytag (USNM
# 73054). Paratypes: CANADA: New Brunswick: York Co.,
Fredericton, 12-VI-1933, J. L. McDunnough, 4 ¢ 6, 622 (CNC).
Nova Scotia: Cape Breton Co., Baddeck, 23-VI-1936, J. L. McDun-
nough, 1¢ (CNC); Baddeck Forks, Cape Breton Island, 30-VI-1936,
J. L. McDunnough, 12 (CNC). Queens Co., White Point Beach,
23-VIT-1934, J. L. McDunnough, 12 (CNC). Onrario: Nipissing
Dist., Costello Lake, Algonquin Provincial Park, 7 to 10-VI-1939,
W. M. Sprules, 26 6, 699 (INHS). Parry Sound Dist., Kearney,
22-VI-1926, F. P. Ide, 12 (CNC). Quesec: Brome Co., Knowlton,
20-VI-1927, G. S. Walley, 1¢ (CNC); 4-VI-1936, 1¢ (CNC);
12-VI-1928, G. H. Fisk, 19 (CNC). Gatineau Co., Masham Hills,
5-VI to 1-VIII-1935, F. P. Ide, 36 6, 592 (ROM); 27-VI to 8-VIII-
Alloperla imbecilla and relative 261
1935, G. E. Bucher, 11¢ ¢, 62 2 (ROM). Huntingdon Co., Covey
Hill, 15 to 25-VI-1924, G. S. Walley, 3¢ ¢, 29 9 (CNC). Montmorency
Co., Montmorency River, 14-VII-1958, R. Malouin, 19 (CNC);
Barriere Brook, Laval, 28-VII-1958, 19 (CNC); Goudreault River,
Laval, 14-VII-1958, R. Malouin, 1¢, 19 (CNC); Dauphine River,
Orleans Island, 13-VI-1957, R. Malouin, 19 (CNC); La Fleur River,
Orleans Island, 10-VII-1958, R. Malouin, 19 (CNC); Moulin River,
Orleans Island, 6-VII-1958, R. Malouin, 2¢ ¢ (CNC); St. Patrick
Brook, Orleans Island, 26-VI-1957, R. Malouin, 16, 222 (CNC);
24-VI to 4 VIII-1958, 156 6, 1022 (CNC); 4-VI to 10-VII-1959,
33646, 15992 (CNC); 28-VI-1960, 229 (CNC). Saguenay Co.,
Nabisipi River, 4-VII-1962, G. Shooner, 346 6, 599 (UMC). Terre-
bonne Co., du Diable River, Mont Tremblant Provincial Park, 28-V
to 4-VIII-1959, A. Robert, 94 6, 3699 (UMC); 25-VI-1960, 19
(CNC); Mont Tremblant Provincial Park, 3 to 18-VI-1958, A. Robert,
156 4, 2399 (UMC); 19607, 11¢¢, 1199 (UMC). Ungava,
Great Whale River, 7 to 20-VII-1949, V. R. Vockeroth, 2¢ ¢, 529
(CNC). UNITED STATES: Gerorcra: Rabun Co., Burton, 21-V-1911,
J. C. Bradley, 16, 12 (CU). ? Co, S. Georgia, ?-V-1931, P. W.
Fattig, 1¢ (CU). Mare: Hancock Co., Bar Harbor, Mt. Desert
Island, 13 & 14-VI-?, C. W. Johnson, 36 6, 19 (BU). MaAnybanp:
Baltimore Co., Baltimore, 16-V-1938, E. G. Fisher, 2¢ 6, 19 (ANSP).
Frederick Co., Fishing Creek, Mountaindale, 4-VI to 1-VII-1958,
P. H. Freytag, 62 9 (UK); Little Hunting Creek, 25-V-1958, P. H.
Freytag, 16 (UK). Massacuusetrs: Franklin Co., Johnsons Brook,
Colrain, 16-VI-1968, C. H. Nelson, 14 (CHN). Hampshire Co.,
Westfield River, Rt. 9, Cummington, 17-VI-1965, F. R. Holbrook,
1g, 19 (CHN). Micnican: Marguette Co., Marguette, 25-VII-?,
Hubbard & Schwarz, 1¢ (USNM). Minnesota: Cook Co., ?, 5-VIII-
1939, H. T. Peters, 14, 39 2 (UM); Kimball Creek, 22-VII-1948,
P. H. Harden, 24 6, 2929 (UM); Temperance River, 6 miles from
Lake Superior, 22-VII-1948, P. H. Harden, 344, 3292 (UM);
Kadunce Creek, Grand Marais (dead in spider web), 8-VIII-1929,
C. T. Schmidt, 34 6, 499 (UM) (INHS); Devil’s Track River,
Grand Marais (dead in spider web), 7-VIII-1935, C. T. Schmidt, 64 ¢,
329 (UM) (INHS); Devil’s Track River, Gunflint Trail, 11-VI-1969,
H. Bell, 1¢, 499 (USNM); Little Devil’s Track Creek, Grand
Marais, C. T. Schmidt, 1599 (UM) (INHS). Lake Co., Stewart
River, Boy Scout Camp, N. Two Harbors, 9-VI-1966, H. Bell, 1¢,
19 (USNM); 20-VI-1966, 1¢, 429 (USNM). St. Louis Co.,
French River, 22-VII-1948, P. H. Hardin, 22 9 (UM). New Hamp-
SHIRE: Coos Co., Imp Creek, Pinkham Notch, near Mt. Washington,
23-VI-1941, Frison & Ross, 19 (INHS); Gorham, 23-VI-1941, Frison
& Ross, 1¢, 19 (INHS). Grafton Co., Franconia, ?, A. T. Slosson,
19 (AMNH); Bog Brook, North Woodstock, 21-VI-1941, Frison &
Ross, 12 (INHS); Whitcherville Brook, near Benton, 21-VI-1941,
262 Proceedings of the Biological Society of Washington
Frison & Ross, 2292 (INHS). Merrimack Co., Ragged Mountain
Brook, East Andover, 16-VI-1973, D. N. Alstad, 34 6, 39 9 (USNM).
New York: Albany Co., Catskill Creek, Rt. 145, near Cooksburg,
16-VI-1964, Nelson & Root, 1¢ (CHN). Essex Co., Alcohol Brook,
Adirondacks Lodge, 1-VII-1923, ?, 1¢ (CU); trib. Au Sable River,
Keene, 20-VI-1941, Frison & Ross, 1¢ (INHS); Tahawus, 20-VI-1941,
Frison & Ross, 1¢ (INHS). Fulton Co., Trammel Brook, near Strat-
ford, 18-VII-1935, ?, 1¢ (CU). Greene Co., Ashland, Rt. 23, 19-VI-
1964, Nelson & Root, 246 6, 129 (CHN). Hamilton Co., Sacandaga
River, Sport Island, 14-VI-1910, Alexander, 16, 19 (MCZ); Bear
Brook, near Blue Mt. Lake, Adirondack State Park, 19-VI-1941, Frison
& Ross, 36 ¢, 12 (INHS). Tompkins Co., Ithaca, 1 to 7-VII-?,
N. Banks, 12 (MCZ); 9-VII-1924, ?, 14, 599 (CU). Ulster Co.,
Chichester, Rt. 214, 16-VI-1964, Nelson & Root, 246 ¢, 19 (CHN).
NortH Caronina: Swain Co., Smokemont Cmpg., 11-V-1944, Frison
& Ross, 8646, 19 (INHS). Transylvania Co., Whitewater River,
Rt. 171, 18-V-1970, O. S. Flint, Jr..2¢ ¢,19 (USNM). PENNSYLVANIA:
Sullivan Co., Little Loyalsock Creek, Dushore, ?, 1¢ (UU). SourH
Carouina: Pickens Co., Six Mile Creek, Hwy. S-160, 1-V-1968,
P. H. Carlson, 12 (USNM). Oconee Co., Chattooga River, Rt. 28,
8-V-1969, J. C. Morse, 16 (USNM). TENNEsSEE: Monroe Co.,
Tellico River, 21 miles SE, Tellico Plains, 18-VI-1969, W. L. Peters,
464,12 (UU) (USNM). Polk Co., 12.8 miles N. jct. Rt. 30/64
on Rt. 30, Cherokee National Forest, 9-V-1970, C. H. Nelson, 16
(CHN); .6 mile N. Parksville Lake Cmpg., Rt. 30, Cherokee National
Forest, 11-VI-1970, C. H. Nelson, 26 ¢ (CHN). Sevier Co., Elkmont
Cmpg., 12-VI-1935, H. H. Ross, 1¢ (INHS); LeConte Creek, Gatlin-
burg, 14-V-1939, Frison & Ross, 46 ¢ (INHS); Fighting Creek
Gap, Gatlinburg, 27-V-1934, T. H. Frison, 19 (INHS); 15-V-1939,
Frison & Ross, 1¢ (INHS); Fork of Little Pigeon River, Gatlinburg,
27-V-1934, T. H. Frison, 66 6, 922 (INHS); Little Pigeon River,
Gatlinburg, 29-V-1934, T. H. Frison, 26 6, 12 (UU); 12-VI-1935,
H. H. Ross, 599 (INHS); Gatlinburg, 14-V-1939, Frison & Ross,
14, 19 (INHS); 13-VI-1940, T. H. Frison, 16, 19 (INHS); 11-V-
1944, Frison & Ross, 1¢ (INHS). Vermont: Orange Co., Waits
River, West Topsham, 21-VI-1941, Frison & Ross, 16 (INHS).
Windham Co., Townshend, 16-VII-1934, H. Kahl, 192 (CM).
There are also literature records (as A. imbecilla) from Newfoundland
(Labrador) Brinck (1958) and Connecticut Hitchcock (1968), which
likely represent A. atlantica but were not confirmed in this study.
Diagnosis: Alloperla atlantica may be separated from A. imbecilla
in the male adult stage by the detailed structure of the epiproct.
The epiproct of A. atlantica has a naked apex (Figs. 9 & 10), while
the epiproct of A. imbecilla has darkly sclerotized serrations on the
dorsolateral margins of the apex (Figs. 3 & 4). The female adults
are very similar and cannot always be separated with certainty. How-
Alloperla imbecilla and relative 263
ever, the subgenital plate of A. atlantica tends to be slightly longer
and thinner with a more pointed apex as shown in Needham and
Claassen (1925).
The female of Alloperla usa Ricker has a subgenital plate of similar
shape but can be distinguished because of its larger size and the
more flattened appearance of the projecting subgenital plate in
lateral view. Ricker (1952) gives the length of A. usa females as
8.5-9.5 mm but most specimens are larger attaining up to 12 mm.
LITERATURE CITED
Banks, N. 1906. On the perlid genus Chloroperla. Entomol. News
17:174—175.
—. 1907. Catalogue of the neuropteroid insects (except
Odonata) of the United States. Amer. Entomol. Soc., Phila-
delphia. 50 p.
——. 1908. A list of neuropteroid insects from North Carolina.
Proc. Entomol. Soc. Wash. 9:149-156 (1907).
Brincx, P. 1958. On a collection of stoneflies (Plecoptera) from
Newfoundland and Labrador. Opuscula Entomol. 23:47-58.
CuaassEN, P. W. 1940. A catalogue of the Plecoptera of the world.
Memoir Cornell Univ. Agr. Exp. Sta. 232:1-235.
Gaurin, A. R. 1956. An annotated list of the stoneflies of Ohio
(Plecoptera). Ohio Jour. Sci. 56:321-324.
1964. The Chloroperlidae of North Amerika. Gewédsser
Abwisser 34/35:37—49.
Hacen, H. 1861. Synopsis of the Neuroptera of North America,
with a list of South American Species. Smithsonian Misc.
Collect., Philadelphia, 347 p.
Harpen, P. H. 1942. The immature stages of some Minnesota
Plecoptera. Ann. Entomol. Soc. Amer. 35:318-331.
and C. E. Micxen. 1952. The stoneflies of Minnesota
(Plecoptera). Univ. Minnesota Agr. Exp. Sta. Tech. Bull.
201: 1-84.
Hircucocx, S. W. 1968. Alloperla (Chloroperlidae: Plecoptera) of
the Northeast with a key to species. Jour. New York Entomol.
Soc., 76:39-46.
Inuims, J. 1966. Katalog der rezenten Plecoptera. Das Tierreich.
Lieferung 82. Walter de Gruyter & Co., Berlin, 632 pp.
NEEDHAM, J. G. anp P. W. Craassen. 1925. A monograph of the
Plecoptera or stoneflies of America north of Mexico. Thomas
Say Found., Entomol. Soc. Amer. 2:1-397.
Procter, W. 1938. Biological survey of the Mount Desert Region.
VI. The insect fauna. Wistar Inst. Anat. Biol., Philadelphia.
A496 p.
1946. Biological survey of the Mount Desert Region. VII.
264 Proceedings of the Biological Society of Washington
RICKER,
RICKER,
Say, T.
The insect fauna. Wistar Inst. Anat. Biol., Philadelphia,
566 p.
W. E. 1948. Stoneflies of the Maritime Provinces and
Newfoundland. Trans. Roy. Canad. Inst., 26:401-414 (1947).
1952. Systematic studies in Plecoptera. Indiana Univ. Publ.
Sci. Ser. 18:1—200.
W. E., R. Matouin, P. Harper and H. H. Ross. 1968.
Distribution of Quebec stoneflies (Plecoptera). Natur.
Canad. 95:1085-1123.
1823. Description of insects belonging to the order Neuroptera
Lin., Latr. collected by the expedition authorized by J. C.
Calhoun, Secretary of War, under the command of Major
S. H. Long, p. 160-165. In Western Quarterly Reporter,
Vol. 2 (reprint 1883, LeConte ed. p. 170-175).
SpRULES, W. M. 1948. An ecological investigation of stream insects
in Algonguin Park, Ontario. Univ. Toronto Stud. Biol.
Ser. 56:1-81 (1947).
WaLKER, J. D. 1947. A list of the stoneflies, Plecoptera, known to
occur in southeastern Ohio. Ohio Jour. Sci. 47:134—136.
Vol. 87, No. 25, pp. 265-272 22, October 1974
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
THE ISOPOD CRUSTACEAN GENUS
MUNNOGONIUM GEORGE AND
STROMBERG, 1968 (MUNNIDAE,
ASELLOTA)
By Tuomas E. BOWMAN AND GEORGE A. SCHULTZ
Smithsonian Institution, Washington, D.C. 20560,
and 15 Smith St., Hampton, N. J. 08827
The genus Munnogonium was established by George and
Stromberg (1968) for their new species of Munnidae, Munno-
gonium waldronense, from the San Juan Islands, Washington.
The generic name was based on the belief that the new genus
was intermediate between Munna and Pleurogonium, or at
least . . . “combines a number of characters of both . . .”.
The genus actually appears to be closest to Austrosignum
Hodgson (1910). If the diagnosis of Austrosignum by Norden-
stam (1933; repeated by Menzies and Barnard, 1959) is
compared to that of Munnogonium by George and Stromberg
(1968), only 3 differences of possible generic value can be
found. 1. Coxae visible dorsally on pereonites 2-7 in Munno-
gonium, but only on pereonites 5-7 in Austrosignum. 2.
Ocular processes slender in Austrosignum, short and partly
hidden in dorsal view by base of antenna 1 in Munnogonium.
3. Mandible with 3-segmented palp in Austrosignum, without
palp in Munnogonium. A brief discussion of these differences
follows.
As shown in Figures 1-2, coxae in Munnogonium are visible
dorsally on pereonites 2-7 in the male and on pereonites 5-7
in the female. In the latter, the anterior pereonites expand
laterally in conjunction with development of the marsupium,
with the result that the coxae come to lie beneath the pleura
where they are not visible in dorsal view. Validity of the
first difference seems questionable.
25—Proc. Brow. Soc. WasH., Vou. 87, 1974 (265)
266 Proceedings of the Biological Society of Washington
Length and slenderness of the ocular processes vary among
the known species of Austrosignum, and in A. falklandicum
Nordenstam (1933) these processes are short and partly
covered by the base of antenna 1 as in Munnogonium. We
doubt the significance of this character.
We are left with the presence or absence of a mandibular
palp as the only difference between species of Austrosignum
and Munnogonium, and this seems to us to be a character of
generic value. Species of Austrosignum which lack a mandib-
ular palp and should therefore be transferred to Munnogonium
are the following: A. grande Hodgson, 1910; A. tillerae
Menzies and Barnard, 1959; A. globifrons Menzies, 1962;
A. erratum Schultz, 1964; and A. maltiniit Schiecke and Fresi,
1972. Hodgson (1910, p. 72) considered Austrimunna incisa
Richardson (1908) “most closely allied to” if not identical
with his species A. grande. In his list of species of Austro-
signum, Menzies (1962, p. 50) included A. incisa (Richard-
son) and Paramunna dubia Hale (1937), but gave no reasons
for transferring them to Austrosignum. It was not definitely
stated in the original descriptions whether either has or does
not have a mandibular palp.
George and Stromberg (1968) pointed out the possibility
that Austrosignum tillerae Menzies and Barnard might be
a Munnogonium, but because of the brevity of the description
and the fact that the type-series was temporarily unavailable
to them, they did not make the transfer. We have examined
the 5 paratypes of Austrosignum tillerae (the holotype
could not be found) and find that not only is this species a
Munnogonium, but it is conspecific with M. waldronse. In
support of this finding we give below some descriptive notes,
based on specimens from Puget Sound (Figs. 1-20) and
on paratypes from southern California (Figs. 21-31).
Munnogonium tillerae (Menzies and Barnard)
Figures 1-31
Austrosignum tillerae Menzies and Barmard, 1959, pp. 8-9. fig. 1 A—C.
Munnogonium waldronense George and Strémberg, 1968, pp. 226—230,
figs. 1-2.
Material examined: Paratypes of Austrosignum tillerae (499 1¢),
from southern California (Figs. 21-31). More than 50 specimens of
The isopod genus Munnogonium 267
Fics. 1-10. Munnogonium tillerae. 1, Male, dorsal (1.45 mm).
2, Female, dorsal (1.42 mm). 3, Eye and bases of antennae 1 and 2,
dorsal. 4, Male telson, dorsal. 5, Female (1.2 mm) operculum and
telson, ventral. 6, Left male antenna 1, dorsal. 7, Left male antenna
2, dorsal. 8, Left mandible, male. 9, Labium, mandibular surface.
10, Maxilla 1, male.
M. tillerae collected by William Shapeero in Puget Sound, about
1350 m NW of West Point, Seattle, Washington, November 1965
(Figs. 1-20). Depth about 6 m; bottom of fine to muddy sand with
many seapens (Leioptilus quadrangularis) present. ‘Ten specimens
(566, 5292) from a sandy mud bottom at about 6 m depth in
Shilshole Bay, off Seattle, Washington, collected in the summer of 1970
268 Proceedings of the Biological Society of Washington
Fics. 11-20. Munnogonium tillerae. 11, Maxilla 2, male. 12,
Maxilliped, male. 13, Pereopod 1, male. 14, Pereopod 2, male. 15,
Pereopod 2, female, basis. 16-19, Pleopods 1-4, male. 20, Right
uropod, dorsal.
by William Shapeero. The longest male was 1.3 mm and _ longest
female 1.6 mm long.
Supplemental description of specimens from Puget Sound: Length
of A. tillerae paratypes, 9 1.0-15 mm, ¢ 1.2 mm; Puget Sound
specimens 0.8-1.8 mm. Ocular processes longer and more _ slender
The isopod genus Munnogonium 269
Fics. 21-32. Munnogonium tillerae. 21, Female paratype (1.5 mm).
22, Pleopod 1, male. 23, Paratype male (1.2 mm). 24, Female
operculate pleopod. 25, Maxilliped. 26, Antenna 2. 27, Antenna l.
28, Labium. 29, Dactylus pereopod 7, male. 30, Dactylus pereopod 2,
male. 31, Pereopod 1, male. 32, Munnogonium erratum. Pereopod 1,
holotype, male.
then shown by Menzies and Barnard, partly to almost completely
concealed in dorsal view by proximal segments of antenna 1. Pereonites
5-7, lateral expansions separated by distinct gaps except in holotype 2
of A. tillerae. Coxae visible in dorsal view on pereonites 2-7, except
270 Proceedings of the Biological Society of Washington
in some ovigerous 2 @ in which they are not visible in pereonites 2-4.
Dorsal surface of pereon with scattered setae (not shown by previous
authors). Antenna 1 with 4-segmented flagellum (3-segmented in
holotype of A. tillerae). Antenna 2 with 6-segmented flagellum
(5-segmented in holotype of A. tillerae). Incisor of mandible with
4 teeth (3 teeth in allotype of M. waldronense). Lower lip with
distal setose lobe on either side, separated by notch from small blunt
lobe covered with fine setules. Other mouthparts agree with those
described by George and Strémberg, with slight differences in numbers
of setae. Pereopod 2 sexually dimorphic; posterior margin of basis
produced into flange ending distally in rounded lobe in male; flange
absent in female. Female operculum with scattered setae on ventral
surface and row of setae on convex proximal part of each lateral
margin. Male pleopods 2—4, not previously illustrated, as in Figures
17-19. Endopod of pleopod 3, 1.5 times as wide as exopod; apex
with seta inserted in notch; lateral margin with 2 setae on distal
half. Exopod of pleopod 3 narrowly pyriform, with terminal seta.
Endopod of pleopod 4 much wrinkled; exopod with convex lateral
margin, concave medial margin, and slender pointed apex.
Munnogonium erratum (Schultz)
Figure 32
Austrosignum erratum Schultz, 1964, pp. 307-310, figs. 1-11.
Munnogonium erratum (Schultz ).—George and Stromberg, 1968, p. 230.
The holotype, thus far the only specimen discovered, was reexamined,
and found to be a Munnogonium, distinct from the other Pacific coast
species, M. tillerae. The most obvious difference is the much greater
separation of the lateral parts of the pereonites in M. erratum, especially
those of pereonites 4-7. The absence of a mandibular palp is confirmed.
Antenna 1 is broken at the fourth flagellar segment, giving this segment
the flattened appearance shown by Schultz (1964, fig. 8). Pereopod
1 resembles that of M. tillerae (compare figs. 31 and 32).
ACKNOWLEDGEMENTS
We are grateful to William Shapeero, Department of Fisheries and
Wildlife, Oregon State University, for the fine collection of Munno-
gonium tillerae from Puget Sound.
LITERATURE CITED
GrorcE, RosperT Y., and JARL-OvE STROMBERG. 1968. Some new
species and new records of marine isopods from San Juan
Archipelago, Washington, U.S.A. Crustaceana 14(3) :225-254.
Hate, Hersert M. 1937. Isopoda and Tanaidacea. Australasian
Antarctic Exped. 1911-14, Sci. Repts. ser. C.—Zoology
and Botany 2(2):1-45.
The isopod genus Munnogonium DAT)
Hopcson, T. V. 1910. Crustacea, IX, Isopoda. National Antarctic
Exped. 1901-1904, 5(9):1-77, pls. 1-10, London.
Menzies, Roserr J. 1962 Reports of the Lund University Chile
Expedition 1948-49, 42. The zoogeography, ecology, and
systematics of the Chilean marine isopods. Lunds Universitets
Arsskrift, N. F. (2)57(11):1-162.
, and J. Laurens BARNARD. 1959. Marine Isopoda on coastal
shelf bottoms of southern California; systematics and
ecology. Pacific Nat. 1(11):1-35.
NorpENsTAM, AKE. 1933. Marine Isopoda of the families Serolidae,
Idotheidae, Pseudidotheidae, Arcturidae, Parasellidae and
Stenetriidae mainly from the South Atlantic. Further Zool.
Res. Swedish Antarctic Exped. 1901-1903, 3(1):1-284,
pls. 1-2.
RicHARDsON, Harriet. 1908. Isopodes (2° Mémoire). Expéd.
Antarctique Francaise (J. Charcot):1—8, Paris.
SCHIECKE, ULRICH, AND EucGENIO Fresi. 1972. Record of the asellote
isopod Austrosignum Hodgson from the Bay of Naples:
Austrosignum maltinii n. sp. (Paraselloidea, Munnidae).
Crustaceana, supplement 3, Studies on Peracarida:31-38.
ScHuLTz, GrorcE A. 1964. Some marine isopod crustaceans from
off the Southern California coast. Pacific Sci. 18(3):307-314.
972, Proceedings of the Biological Society of Washington
Vol. 87, No. 26, pp. 273-282 22, October 1974
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
A REVIEW OF THE COPEPOD SCOTTOCALANUS
SECURIFRONS (T. SCOTT) AND A NOTE ON
ITS SYNONYM SCOLECITHRIX CUNEIFRONS WILLEY
(CALANOIDA: SCOLECITHRICIDAE)
By T. C. GopALAKRISHNAN
Regional Centre of National Institute of Oceanography
(C.S.I.R.) Cochin, India
ABSTRACT
Scottocalanus securifrons from the International Indian
Ocean Expedition Collections is described and compared with
that of T. Scott 1893, A. Scott 1909, Tanaka 1961, Lophothrix
securifrons Wolfenden 1904 and Scolecithrix cuneifrons Willey
1918. A thorough examination of the descriptions of securi-
frons and cuneifrons and the examination of 2 syntypes of
S. securifrons from the British Museum revealed a doubtful
existence of Scolecithrix cuneifrons as a distinct species. Hence
it is considered as a synonym of S. securifrons in agreement
with Vervoort (1965).
INTRODUCTION
While studying the scolecithricid copepods collected during
the International Indian Ocean Expedition 1960-65 (IIOE) I
encountered a number of specimens of both sexes belonging
to the genus Scottocalanus Sars. Upon comparing the ILOE
specimens with published descriptions of species of Scotto-
calanus, I found good agreement with S. securifrons (T. Scott,
1893) and also with S. cuneifrons (Willey, 1918). The IIOE
specimens are assigned to the older S. securifrons and de-
scribed below.
26—Proc. Brot. Soc. WasuH., Vou. 87, 1974 (273)
274 Proceedings of the Biological Society of Washington
Scottocalanus securifrons (T. Scott)
(Figure 1 a—g, Figure 2 a—c, Figure 3 a—h)
Scolecithrix securifrons T. Scott, 1893, p. 47, pl. 4, figs. 40-56, pl. 5,
fig. 1 [2 only, ¢ = Scottocalanus helenae (Lubbock )|—Giesbrecht
and Schmeil, 1898, p. 49.—van Breemen, 1908, p. 76, fig. 88 [2
only|.—Canu, 1896, p. 425.—Thompson, 1903, p. 20.—Norman 1903,
p. 137.—Cleve, 1904, p. 197.—Cons. Explor. Mer, 1909, p. 99.—
Jespersen, 1940, p. 36.
Lophothrix securifrons Wolfenden, 1904, p. 120, pl. 9, figs. 12-15.
Lophothrix securifrons (T. Scott).—Wolfenden, 1911, p. 268.
Scottocalanus acutus Sars, 1905, p. 7.
Scolecithrix cuneifrons Willey, 1918, p. 194, figs. 17-24.
Scottocalanus securifrons (T. Scott).—Sars, 1905, p. 7 [by implication];
1912, p. 654; 1924-1925, p. 160-162, pl. 45 figs. 1-8.—Pearson, 1906,
p. 19.—Farran, 1908, p. 57; 1920, pp. 18, 21; 1926, p. 267; 1929, p.
251.—Paulsen, 1909, p. 137.—A. Scott, 1909, p. 104, pl. 25 figs. 1-9,
pl. 28 figs. 1-9.—Stebbing, 1910, p. 529.—With, 1915, p. 220, pl. 8
fig. 13, text figs. 71-73.—Cons. Explor. Mer, 1916, p. 57.—Lysholm
and Nordgaard, 1921, p. 21.—Rose, 1929, p. 26; 1933, p. 144, fig. 144;
1942, p. 148.—Wilson, 1936, p. 91; 1950, p. 340—Tanaka, 1937, p.
259, figs. Qa—c; 1953, p. 132; 1961, p. 141-148, fig. 106; 1969, p. 275.
—Leavitt, 1938, p. 384.—Lysholm, Nordgaard and Wiborg, 1945, p.
26.—Sewell, 1947, p. 143.—Fraser and Saville, 1949a, pp. 61, 63.—
Brodsky, 1950, p. 242, fig. 152.—Wiborg, 1955, p. 51.—Hida and
King, 1955, p. 11.—Marques, 1956, p. 15; 1958, p. 225; 1959, p. 211.
—Heinrich, 1958b, p. 1029.—Vinogradov, 1960, p. 502.—Grice, 1962,
p. 213, pl. 19 figs. 12-15.—Grice and Hart, 1962, appendix, tab.
—Owre, 1962, p. 492.—Vervoort, 1965, p. 36.—Fleminger, 1967, p.
194.—Owre and Foyo, 1967, p. 63, figs. 98, 400-403, 409, 410.
—Park, 1970, p. 476.—Bowman, 1971, p. 34.
Description of the specimens: Female. Length 4.2 mm. Head and
first thoracic segment, 4th and 5th thoracic segments fused. Head with
high median crest (Fig. 1c). Last thoracic segment produced posteriorly
into triangular expansion terminating in sharp pointed spine on either
side. Rostrum bifid at tip (Fig. 2 a). Abdomen 4-segmented. Genital
segment swollen ventrally at mid-length; its ventral posterior margin
overlapping the following segment (Fig. 1 d). Posterolateral margins of
genital segment furnished with spines which are absent on dorsal and
ventral side (Fig. 1 e). A. Scott (1909) shows spines on the posterior
margin of the genital segment present on the dorsal as well as lateral
surfaces. Posterior margins of 2nd and 3rd abdominal segments with a
hyaline fringe (Fig. 1 e). Anal segment very short. Caudal rami
almost as wide as long, each with 4 setae.
First antenna with 23 separate segments, when the partly separated
8th and 9th segments are counted as one (Fig. 1 a). First maxilla with
Review of Scottocalanus securifrons 275
OES ab.cdf.&
| o°3mm le
Fic. 1. Scottocalanus securifrons. a, Ist antenna 2; b, Ist antenna
é; c, frontal profile 2; d, last thoracic segment and genital segment 9;
e, genital segment 9, enlarged; f, frontal profile ¢; g, last thoracic
segment ¢.
numbers of setae on different lobes as follows (Fig. 2 b): Inner lobe
1 with 12 setae of which 3 are on posterior surface; inner lobe 2 with 2
setae; inner lobe 3 with 3 setae; basipod 2 with 5 setae; endopod seg-
ment 1 with 3 setae; endopod segments 2 and 3 together with 4 setae;
exopod with 8 setae; outer lobe with 9 setae. Exopod segment with fine
surface hairs at distal end. Second maxilla endopod with 4 bud-like and
276 Proceedings of the Biological Society of Washington
Fic. 2. Scottocalanus securifrons. a, rostrum @; b, Ist maxilla 9;
c, distal portion of 2nd maxilla 9.
3 vermiform filaments (Fig. 2 c). Swimming legs 1-4 as shown in
figures (Figs. 3 a—d). .
Fifth pair of legs asymmetrical; subapical spine of left leg thicker
than that of right leg. Subapical spines with two rows of teeth (Fig. 3 e).
Male. Length 4.57 mm. General appearance similar to that of female.
Head with high median crest (Fig. 1 f). Last thoracic segment terminat-
ing in small spine on either side (Fig. 1 g). Abdomen 5—segmented.
Posterior margin of 2nd to 4th abdominal segments with hyaline fringe
(Fig. 1 g). First antenna with 20 segments when fused 8th to 12th
segments, partly divided by 2 incomplete sutures between segments 8
and 9 and 11 and 12, are counted as one (Fig. 1 b). 5th leg as illustrated
(Fig. 3 fh). The small teeth on the inner margin of the proximal joint
of the endopod of the left leg figured by A. Scott are not present in
Tanaka’s specimens, Willey’s specimens or in the IIOF, specimens.
DISCUSSION
The IIOE specimens agree well with descriptions of S. securifrons by
T. Scott (1893, @ only), A. Scott (1909), Wolfenden (1904), and
Tanaka (1961). They also appear to conform to Willey’s (1918) de-
scription of S. cuneifrons. Willey was aware of the similarity of his
S. cuneifrons to S. securifrons, and stated that he was at first inclined
to identify his specimens as S. securifrons, but decided to establish a
Review of Scottocalanus securifrons 201
o'15mm In
Fic. 3. Scottocalanus securifrons. a, Ist leg 9; b, 2nd leg @; c, 3rd
leg 2; d, proximal portion of 4th leg 9; e, 5th leg 9; f, left 5th leg
6; g, right 5th leg ¢; h, left 5th leg ¢, 2nd joint of Re, enlarged.
new species for them because the male fifth legs of his specimens differed
from those of S. securifrons. Apparently he was referring to T. Scott’s
male, which is now believed to belong to a different species, Scotto-
calanus helenae (Lubbock), and not to the male described by A. Scott
which had a fifth leg very similar to that of S. cuneifrons. Curiously,
278 Proceedings of the Biological Society of Washington
Willey did not cite either T. Scott (1893) or A. Scott (1909) in his
bibliography.
In order to be certain about the identity of T. Scott’s specimens, I
requested a loan of them from The British Museum (Natural History).
Although Scott (1893) reported S. securifrons from 5 “Buccaneer”
stations, Dr. Roger J. Lincoln reported that the British Museum had
only 2 females, the male being missing. Upon examining these 2 females,
I found that one of them is not S. securifrons but another species of
Scottocalanus, possibly S. australis Farran (1936). The remaining female
is herewith designated as the lectotype of Scolecithrix securifrons T. Scott
in order to obviate further confusion.
Distribution: The species has a fairly wide distribution, and has
been recorded from the Atlantic, Pacific and Indian Oceans. It has been
recorded from the central and southern part of the Arabian Sea (Sewell,
1947), from the Indian Ocean off Port Shepstone, South Africa (Cleve,
1904a), from many localities in the eastern part of the Malay Archipelago
(A. Scott, 1909, and Snellius Expedition), from the Philippine Islands
region (Wilson, 1950), from the central equatorial Pacific, 00°03’N,
157°00’E (Grice, 1962), from the California Current region (Fleminger,
1967), from Sagami and Suruga Bays, Izu region, Japan (Tanaka, 1937,
1961) from surface waters off Three Kings Islands, New Zealand
(Farran, 1929), from many localities in the West-Pacific ranging from
off Peru to the Galapagos Islands region (Wilson, 1950) and from the
Far Eastern and Polar Seas of the U.S.S.R. (Brodsky, 1950). The
specimens described in this paper are from around the central part of
the equatorial Indian Ocean (03°29’N, 77°54’E).
In the Atlantic S. securifrons is widely distributed over large areas,
penetrating at least as far north at the Atlantic slope off Cabot Strait,
42°31’N, 63°31.5’W (Willey, 1918) and 43°18’N, 60°11’W (Rose, 1929).
Also known from 47°47’-63°08’N, 8°00’-26°20’W (With, 1915; Lysholm
and Nordgaard, 1921); the South coast of Iceland, 63°08’N, 21°30’W;
62°47’N, 15°03’W (Paulsen, 1909; Jespersen, 1940); the Faroe Channel,
+ 60°N, 7°W (Norman, 1903; Wolfenden, 1904); 28°-58°N, 7°-50°5’W
(Lysholm, Nordgaard and Wiborg, 1945); NNW of Achill Head, Ireland
(Norman, 1903); 52°06’-54°33’N, 10°30’—-15°53.9’W (Thompson, 1903;
Farran, 1908); off the south-west coast of Ireland (Farran, 1920);
27°43’-47°43’N, 8°06’-42°40.5°W (Sars, 1925); Bermuda (Wilson,
1936); 30°08’N, 31°19’W (Sars, 1912); Florida current (Owre, 1962);
between Bermuda and New York (Grice and Hart, 1962); Bay of Biscay
+ 47°N, 8°W (Farran, 1926); 44°17’N, 4°38’W (Canu, 1896); 20°41’N,
31°53’W; 17°28’N, 29°42’W; 16°24’N, 28°53’W (Wolfenden, 1911);
Gulf of Guinea (T. Scott, 1894); off Angola (Marques, 1956, 1958,
1959); 26°59’S, 17°06’W and 35°10’S, 2°33’W (Wolfenden, 1911). It
has been captured in the northern part of the North Sea during the
periodical plankton investigations, 1902-1908 (Cons. Explor. Mer 1909,
Scottish area). It has been recorded from the southeastern United
States between Cape Hatteras and Southern Florida (Bowman, 1971).
Review of Scottocalanus securifrons 279
ACKNOWLEDGMENTS
I am grateful to Dr. N. K. Panikkar, former Director, National Institute
of Oceanography, India for allowing me to take the material to the U. S.
National Museum and for his interest in the progress of this study. My
sincere thanks are due to Dr. Janet M. Bradford for her encouragement
and to Dr. T. E. Bowman for his valuable suggestions and helpful
criticism during the preparation of this paper. Thanks are also due to
the U. S. National Muesum for providing all facilities to pursue this work.
LITERATURE CITED
Bowman, T. E. 1971. The distribution of calanoid copepods off the
southeastern United States between Cape Hatteras and southern
Florida. Smithsonian Contrib. Zool. No. 96:1-58.
BREEMEN, P. J. van. 1908. VIII. Copepoden. Nordisches Plankton
4(8):1-263.
Bropsky, K. A. 1950. Calanoida of the Far Eastern Seas and Polar
Basin of the U.S.S.R. Keys to the Fauna of the USSR No. 35:
1-442. (Translation: Israel Program for Scientific Transla-
tions, 1967.)
Canu, E. 1896. Copépodes. In R. Koehler, ed. Résultats scientifiques
de la Campagne du “Caudan” dans le Golfe de Gascogne, Fasc.
2. Ann. Univ. Lyon 26:421—437, pl. 18.
Creve, P. T. 1904. The plankton of the South African seas. I. Copep-
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sal
Vol. 87, No. 27, pp. 283-312 22 October 1974
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
BIOSYSTEMATICS OF CENTRAL AMERICAN SPECIES
OF CTENOCOLUM, A NEW GENUS OF SEED
BEETLES (COLEOPTERA: BRUCHIDAE)
By JoHN M. KincsoLver! AND DoNnaLp R. WHITEHEAD?
National Museum of Natural History, Washington, D. C. 20560
INTRODUCTION
This paper is the third in a series concerned with a group
of related seed beetle genera predatory on leguminous plants
in the New World tropics; see Kingsolver and Whitehead (in
press ) for a review of Caryedes, and Whitehead and Kingsolver
(in press) for a review of Gibbobruchus. As in those papers,
the major purpose is to make known the fauna of Central
America in general and of Costa Rica in particular, in con-
junction with ecological studies of the beetles and their host
plants being conducted by D. H. Janzen.
In this paper, we treat only the species of Ctenocolum known
from Mexico and Central America, an area including the
northernmost part of the range of the genus. Two species
groups are distinguished, each represented in this area by four
species. Our respective contributions are as in our Caryedes
paper, but with some nongenital drawings contributed by K.
Conway and L. Heath.
Materials and methods: We studied about 250 specimens,
most of which are deposited in the United States National
Museum of Natural History, Washington (USNM). All Amer-
ican specimens of the host plant subtribe Lonchocarpinae
(tribe Dalbergiae, family Fabaceae) in the United States
National Herbarium were examined for additional data. Meth-
ods are as described previously (Kingsolver and Whitehead,
1 Systematic Entomology Laboratory, IIBII, Agr. Res. Serv., USDA.
2 Organization for Tropical Studies.
27—Proc. Biot. Soc. WaAsuH., Vou. 87, 1974 (283 )
284 Proceedings of the Biological Society of Washington
in press; Whitehead and Kingsolver, in press). Distribution
records are mapped for each species. Holotypes of the five
species described as new in this paper are deposited in the
USNM.
Acknowledgments: We again thank D. H. Janzen for pro-
viding reared Costa Rican material, for criticizing the manu-
script, and for providing financial assistance from NSF grants
GB-7819 and GB-35032X. Additional fresh material was pro-
vided by C. D. Johnson and M. Sousa. We are indebted also
to P. M. Hammond for important notes on the type specimens
of C. salvini.
Type material of previously described species belonging to
Ctenocolum is housed in the British Museum (Natural His-
tory), London; Naturhistoriska Riksmuseum, Stockholm; and
Zoological Museum, Moscow State University, Moscow. We
thank the curators of these collections for allowing us to study
their material.
Additional material was loaned by the following individuals
and institutions: D. E. Bright, Canadian National Collection,
Ottawa (CNC); H. R. Burke, Texas A&M University, College
Station (TAMC); H. F. Howden, Carleton University, Ottawa
(HFHo); C. D. Johnson, Northern Arizona University, Flag-
staff (NAUF); H. Reichardt, Museu de Zoologia da Univer-
sidade, Sao Paulo (MZSP); and G. E. Wallace, Carnegie Mu-
seum, Pittsburgh (CarP).
TAXONOMY
Ctenocolum, new genus
Type-species: Pachymerus tuberculatus Motschoulsky 1874.
Derivation of name: The name Ctenocolum is derived from Greek:
Ktenos = comb, + Kolon = leg, in reference to the comb-shaped pecten
characteristic particularly of the type-species and its relatives.
Diagnostic combination and relationships: Ctenocolum is one of the
American genera of Bruchinae with pronotum campaniform and gibbous,
hind tibia arcuate, and external carina of hind femur obsolete or repre-
sented by row of teeth. We previously placed this genus (as the
“Caryedes” tuberculatus group) in the “Caryedes’” assemblage (King-
solver and Whitehead, in press) in which the abdomen and pygidium
lack glabrous polished areas and the antennal scrobe of the gena is as
long as or longer than the width of the antennal fossa. Of the three
genera placed in the “Caryedes’ assemblage, Meibomeus and Caryedes
Some Central American seed beetles 985
differ from Ctenocolum by having the synapomorphous condition of
lateral lobes of male genitalia not deeply divided. Conversely, an
increased number of teeth of the hind femoral pecten is synapomorphous
for members of the genus Ctenocolum; either 6-10 teeth with second
and following forming a crescentic profile or 8-16 teeth with second
and following forming a linear profile. The hind tibial mucro is short,
but is neither as short as in Meibomeus nor longer than apical width of
tibia as in most Caryedes.
Description: Body robust; length from pronotum to pygidium 2.0-4.6
mm, about 1.5 times greater than maximum width; width across elytra
1.4-3.6 mm, nearly equal to maximum length of elytron. Head moder-
ately elongate, strongly constricted behind eye, postocular lobe short.
Eye prominent, larger in male than in female or not, interocular ratio
about 0.12-0.24 in male, 0.15-0.24 in female; ocular sinus deep, 4—10
rows of facets behind sinus. Frons with median carina prominent,
alutaceous, extended forward onto base of clypeus, triangularly ex-
panded basally. Frontoclypeal region broad, pentagonal; sides parallel
behind, convergent in front; distance from apex of clypeus to apex of
superior lobe of eye not or slightly greater than length of eye from apex
of superior lobe to base. Gena between base of mandible and antennal
fossa about as long as width of antennal fossa, glabrous. Antenna of
male extended to or beyond elytral humerus, outer articles transverse to
elongate, eccentric to serrate or subflabellate; antenna of female extended
to about elytral humerus, outer articles transverse to slightly elongate,
moderately eccentric to serrate. Pronotum campaniform, sides shallowly
and irregularly concave; median and lateral basal gibbosities low to
moderately elevated, median sulcus shallow to moderately deep; median
anterior gibbosities low, not or shallowly sulcate between; median basal
lobe shallowly emarginate; lateral carina obsolete to distinct. Scutellum
about square, bidentate or truncate. Intercoxal process of prosternum
narrow, acute. Metasternum shallow, broadly rounded in profile. Meso-
sternal lobe broad, flat, apex rounded or truncate. Elytra together
subquadrate; striae deep, punctate; striae three and four strongly de-
flected laterad before base, abbreviated basally by high bidentate gib-
bosity; striae four and five abbreviated apically, coalescent or not; striae
seven, eight, and nine limited basally by humeral gibbosity; interval nine
flat to moderately convex apically, not carinate; disc convex. Front and
middle legs slender, not sex-dimorphic, front coxae contiguous apically,
middle coxae widely separated. Hind coxa densely punctate, middle
third glabrous: Hind femur strongly swollen, external ventral margin
finely dentate to coarsely dentate-carinate to base; ventral sulcus de-
veloped to or nearly to base, deep in apical half; internal ventral margin
with two or more small teeth before pecten; pecten raised, either with
7-9 apical teeth separated from basal tooth by distinct gap, or with
12-16 evenly spaced teeth. Hind tibia strongly carinate, arcuate, flanged
ventrally from near middle to mucro; mucro shorter than apical width of
286 Proceedings of the Biological Society of Washington
tibia; external apical margin oblique or not, lateral coronal tooth sharply
developed or not; hind basitarsus longer than four outer tarsal articles
together. Abdominal sterna telescoped in male, last sternum emarginate,
abdomen not otherwise sex-dimorphic; sterna without polished lateral
areas. Pygidium various, without speculum, dimorphic or not in pattern.
Male genitalia with median lobe broad, not fractured before apex, not
strongly arcuate; internal sac various, with or without distinctive sclerites;
lateral lobes broadly divided; median lobe with strap- or blade-like hinge
sclerites; basal strut of tegmen narrow, with median keel; median lobe
with characteristic patch of setae above apical orifice; basal hood of
median lobe broad.
Distribution: Ctenocolum is restricted to tropical America including
the West Indies, and is most diverse in species in Central America and
northern South America. Eight species are known from Mexico and
Central America; there probably are several additional species from
this area. We recognize two species groups, each of which includes
four of the Central American species.
Remarks: So far as known, larval Ctenocolum attack seeds of mem-
bers only of the subtribe Lonchocarpinae (tribe Dalbergiae, family
Fabaceae). No other bruchid genera are known to attack this group
of plants. The only known host genera for species from Mexico and
Central America are Lonchocarpus and Piscidia, but South American
species occur in Bergeronia and Muellera also. Pods of all of these plants
are indehiscent, and beetles emerge through circular exits cut through
the seed and pod walls.
Key To Species OF CTENOCOLUM FROM MEXICO AND CENTRAL AMERICA
1. Pecten with 8-16 teeth, second and following linear in
profile; hind tibial apex oblique or nearly so, lateral coronal
tooth not distinguished from mucro by deep notch; internal
sac of male genitalia with large distinctive sclerites (Tuber-
culatum ‘Group )) -:40.0¢). we 2.
AY Pecten with 6-10 teeth, second and following crescentic in
profile; hind tibial apex with strong lateral coronal tooth
distinguished from mucro by deep notch; internal sac of male
genitalia without large sclerite (Crotonae Group) _...- D.
2 (1). Eye with about 10 rows of facets behind sinus; external
ventral denticles of hind femur greatly reduced; vestiture
of pronotum and hind femur largely bright orange ..........-..-
SSI ULNA 2. A UN an ec eR ee 1. Ctenocolum acapulcensis n. sp.
De Eye with about 3-6 rows of facets behind sinus; external
ventral denticles of hind femur strongly developed; vestiture
of pronotum and hind femur yellowish or whitish ................ 3.
3 (2’). Eye with about 3 rows of facets behind sinus; vesiture of
elytral intervals two and three uniform from behind scutellum
to midlength, in most specimens dark; female pygidium with
Some Central American seed beetles 287
contrasting longitudinalspattenny cece ee ee eee
Sh 0 Ie 4, Ctenocolum tuberculatum (Motschoulsky ).
By Eye with about 6 rows of facets behind sinus; vestiture of
elytral intervals two and three variegated behind scutellum;
female pygidium without distinctive pattern ......................--.... A,
4 (3’). Dorsoapical angles of antennal articles 8-10 dark; sclerite of
male endophallus not produced, with lateral hooks ................
saci besSe ea ea Ra eae 3. Ctenocolum salvini (Sharp).
4’ Antennal articles 8-10 uniformly colored externally, dark in
most specimens; sclerite of male endophallus produced,
without lateral hooks .................... 2. Ctenocolum janzeni n. sp.
5 (1’). Elytron (Fig. 5) with extensive whitish sutural maculation
Pree Oo ers Oh re EA) a 5. Ctenocolum colburni n. sp.
By Elytron (Fig. 6-8) without extensive whitish sutural macu-
eA Tel Ora epee werent RE a Ls a le AAG pe tpl a gc mi LE Fu lags, 6.
6 (5’). Apex of outer tooth of basal elytral gibbosity nearer to
elytral base than to apex of inner tooth ................0......200000--00+-
acionde ses LEE ee Me Cal adeRO Ie Aaa 6. Ctenocolum biolleyi n. sp.
6’ Apex of outer tooth of basal elytral gibbosity nearer to apex
Of inner tooth than’ to) elytral base 2... =e. Te
7 (6’). Base of notch between teeth of basal elytral gibbosity equi-
distant between elytral base and apex of inner tooth ............
ascii SSN at se MN PIO Ree EDs a 7. Ctenocolum martiale n. sp.
ee Base of notch between teeth of basal elytral gibbosity nearer
apex of inner tooth than elytral base ..........................---000--00-+
PRE ee etn 8 St 8. Ctenocolum crotonae (Fahraeus).
THE TUBERCULATUM GROUP
Diagnostic combination: The Tuberculatum Group is sharply dis-
tinguished from the Crotonae Group by the following characters of the
hind leg and the male genitalia. Hind femur: pecten with 8-16 evenly
spaced teeth, all teeth except first regular in profile. Hind tibia: lateral
coronal tooth not sharply developed, not set off from mucro by deep
emargination, apex oblique. Male median lobe: internal sac with large
complex sclerite; ventral valve acute, sides not or barely sinuate.
Additional characteristics of the Tuberculatum Group are the follow-
ing: eye of male enlarged in some species; ocular sinus shallow to deep,
3-10 rows of facets behind sinus; antenna of male flabellate in some
species; lateral subbasal tubercles of pronotum moderate to strong;
placement of outer tooth of basal elytral gibbosity not or slightly basad
of inner tooth; and hind femur with external ventral series of denticles
reduced in some species.
Distribution: Northern Mexico to northern South America. Four
species are known from Mexico and Central America and are treated
in this paper. We have examined specimens of three additional species
from Colombia and Venezuela; all three are undescribed.
288 Proceedings of the Biological Society of Washington
Taxonomic notes: This group is quite poorly known, as none of the
species are represented by adequate numbers and samples. However,
there are no evident taxonomic problems, as all the species are readily
separable by characteristics of the male genitalia, especially by the form
of the sclerite of the internal sac.
Remarks: Larvae of members of the Tuberculatum Group develop
in seeds of Lonchocarpus and Piscidia. We suspect that the range of
hosts for each species is limited. Only one species is known from more
than one host species, and it is known from three closely related host
species.
Little is known of flight activity. Specimens of two South American
species were taken at black lights in Feburary, and another specimen
of one of them was collected in July.
1. Ctenocolum acapulcensis, new species
Description of holotype male: Ctenocolum, Tuberculatum Group.
Length 4.66 mm. Width 3.2 mm. Integument largely dark rufous or
rufopiceous, elytra largely piceous with rufopiceous variegation; an-
tennal articles 3-10 infuscated, article 11 pale; front and middle legs
and hind tarsus rufotestaceous. Vestiture variegated; pattern of dorsum
(Fig. 1); bright orange on base of head, pronotum, mesepisternum,
metepisternum, dorsal surfaces of all femora, and elytral intervals one
and two; sparse and dark on most of elytra and hind tibia; pale or nearly
white on most of pygidium, abdomen, and lower surface of hind femur.
Head (Fig. 9); antenna flabellate (Fig. 16); eye large, interocular ratio
about 0.15; ocular sinus shallow, 10 rows of facets behind sinus. Pro-
notum and elytra (Fig. 1); lateral subbasal gibbosities strongly de-
veloped. Hind leg (Fig. 24); pecten with about 12 teeth, first tooth
not much larger than second; external ventral series of denticles much
reduced. Pygidium (Fig. 31). Male genitalia (Fig. 43-44).
Type material: Holotype male, “Bot. N°226.”, “Acapulco. Mex.”,
“Chittenden Collection’, “MEX. Guerrero. Acapulco vic. X.1894-
I1I.1895 E. Palmer #226”, “Lonchocarpus eriocarinalis Micheli USNM
Herb.”; in United States National Museum of Natural History, Wash-
ington, type no. 72797.
Paratype, Costa Rica (Fig. 58), 1; in USNM. COSTA RICA. Guana-
caste: 6 km. n. Bagaces, 3.XI.1970, D. H. Janzen #268, reared ex
Lonchocarpus eriocarinalis Micheli, live adult extracted from seed
9.111.1971.
Remarks: Ctenocolum acapulcensis is named for the type locality.
The only other bruchid species known from this host species is C.
martiale, one specimen of which was found in the same herbarium
specimen with the holotype of C. acapulcensis.
Ctenocolum acapulcensis is the only Central American member of the
genus with male antenna flabellate and with external ventral denticles
of hind femur nearly obsolete. An undescribed South American species
Some Central American seed beetles 289
is closely related but differs conspicuously by having only about 8 teeth
in the femoral pecten and by having whitish rather than bright orange
dorsal vestiture.
2. Ctenocolum janzeni, new species
Description: Ctenocolum, Tuberculatum Group. Length 3.3-3.8 mm.
Width 2.1-2.4 mm. Integument largely rufopiceous, elytra largely
piceous with rufopiceous variegation; antenna rufotestaceous, outer
surfaces of articles 8-10 infuscated; base of head with bilobate piceous
mark; front and middle femora mottled, front and middle legs otherwise
rufotestaceous; hind tarsus rufous. Vestiture variegated; pattern of
dorsum (Fig. 2); nearly white on most of pygidium and abdomen; light
tan on mesepisternum, metepisternum, dorsal surface of hind femur;
dark tan or orange brown on much of pronotum and in elytral variega-
tion. Head (Fig. 11); antenna not flabellate (Fig. 17); eyes large, sex-
dimorphic, interocular ratio about 0.17 in male and 0.23 in female;
ocular sinus deep, 6 rows of facets behind sinus. Pronotum and elytra
(Fig. 2); lateral subbasal gibbosities moderately developed. Hind leg
(Fig. 26); pecten with 13-16 teeth, first tooth much larger than second
and following; external ventral denticles of hind femur strongly de-
veloped. Pygidium (Fig. 34). Male genitalia (Fig. 45-47).
Type material: Holotype male, “C.R. Puntarenas Prov., 4 mi. n. jet.
to Puntarenas 6.III.1972 DHJanzen’, “reared ex Piscidia fruits, em. by
20.V1.1972 CODE: VI-20-1972-X”; in United States National Museum of
Natural History, Washington, type no. 72798.
Paratypes, Mexico to Costa Rica (Fig. 57); 8; in CarP, CNC, USNM.
MEXICO. Jalisco: Guadalajara, McConnell. Oaxaca: Carrizal, 25.XII.
1940, F. Miranda #1043 (MEXU), ex herbarium specimen of Piscidia
grandifolia (Donn. Sm.) I. Johnst. San Luis Potosi: El Salto de Agua,
28-30.VII.1960, H. Howden. Sinaloa: no locality specified, M. Narvaez
M. & A. E. Salazar #3129 (US), ex herbarium specimen of Piscidia
mollis Rose. Sonora: Ciudad Obregon, 29.1X.1933, H. S. Gentry #305M
(US), ex herbarium specimen of Piscidia mollis Rose; Torres, 10.11.1903,
F. V. Coville #1659 (US), ex herbarium specimen of Piscidia mollis
Rose. Tamaulipas: 20 mi. n. Nuevo Morelos, 3.VII.1954, D. G. Kissinger.
COSTA RICA. Puntarenas: 4 mi. n. jct. to Puntarenas, 6.III.1972, D. H.
Janzen #VI-20-1972-X, reared ex Piscidia carthagenesis Jacq., emerged
by 20.VI.1972.
Remarks: Ctenocolum janzeni is named for D. H. Janzen, collector
of the holotype.
We report C. janzeni from three species of Piscidia and suspect that
it also attacks seeds of the other Mexican and Central American Piscidia
species. Various other Ctenocolum attack Piscidia in South America
and the West Indies, and Janzen reared C. crotonae from the same
plant that produced the holotype and a paratype of C. janzeni in
“osta Rica.
290 Proceedings of the Biological Society of Washington
3. Ctenocolum salvini (Sharp), new combination
Bruchus salvini Sharp 1885:446. Type-locality: Capetillo, Sacapete-
quez, Guatemala. Type-depository: British Museum (Natural His-
tory), London.
Pseudopachymerus salvini: Pic 1913:12.
Caryedes salvini: Blackwelder 1946:758.
Remarks: We have no host records for this species. Only the type
specimens from Capetillo (Fig. 58) are known and we are unable to
give a full redescription. The bright coppery vestiture (see Sharp 1885,
pl. 26, Fig. 5) is distinctive. Head: bilobate dark mark at base as in
C. janzeni; antenna (Fig. 18) not flabellate, articles 8-10 infuscated
only at apical angles; eye large and ocular sinus deep, not or slightly
sex-dimorphic, ocular ratio about 0.18—-0.20; ocular sinus deep, 6 rows
of facets behind sinus. Hind leg: pecten with about 12 teeth as in
C. tuberculatum, first tooth much larger than second and following;
external ventral denticles of femur strong. Pygidium: pattern not sex-
dimorphic. Male genitalia (Fig. 39-42). The pattern of elytral vestiture
is strongly variegated as in C. janzeni, but differs from the pattern of
both C. janzeni and female C. tuberculatum in that the post-scutellar
dark macula is widely broken by the pale vestiture of intervals one and
two.
4. Ctenocolum tuberculatum (Motschoulsky), new combination
Pachymerus tuberculatus Motschoulsky 1874:244; Sharp 1885:440 (as
synonym of B. longicollis Fahraeus but “. . . will prove to be one of the
varieties of this species”). Type-locality: Panama. Type-depository:
Zoological Museum, Moscow State University, Moscow.
Pseudopachymerus longicollis var. tuberculatus: Pic 1913:11.
Caryedes longicollis var. tuberculata: Blackwelder 1946:758.
Bruchus serratissimus Sharp 1885:443. Type-locality: Paso Antonio,
Escuintla, Guatemala. Type-depository: British Museum (Natural
History ), London. New synonymy.
Pseudopachymerus serratissimus: Pic 1913:12.
Caryedes serratissima: Blackwelder 1946:758.
Description: Ctenocolum, Tuberculatum Group. Length 2.5-3.9 mm.
Width 1.7-2.3 mm. Integument largely dark rufous or rufopiceous,
dorsum with some piceous variegation; antennal articles 8-10 strongly
infuscated externally, articles 4—7 slightly infuscated; front and middle
femora faintly banded, front and middle legs and hind tarsus rufo-
testaceous. Vestiture of dorsum (Fig. 3-4) and pygidium (Fig. 32-33)
sex-dimorphic, pattern more strongly developed in female; black brown
dorsally in median maculations of pronotum, elytra, and pygidium of
female, paler in male; tan or orange brown most of rest of dorsum;
whitish ventrally. Head (Fig. 10); antenna (Fig. 19-20) not flabellate;
eyes large, not sex-dimorphic, interocular ratio 0.21-0.24; ocular sinus
Some Central American seed beetles 291
deep, 3 rows of facets behind sinus. Pronotum and elytra (Fig. 3-4);
lateral subbasal gibbosities moderately developed. Hind leg (Fig. 25);
pecten with about 12 teeth, first tooth much longer than second and
following; external ventral denticles of hind femur strongly developed.
Pygidium (Fig. 32-33). Male genitalia (Fig. 48-50).
Material examined: Mexico to Costa Rica (Fig. 59) and Venezuela,
110. MEXICO. Jalisco: 34 mi. nw. Barra de Navidad, 9.III.1973, C. D.
Johnson #438-73, reared ex PLonchocarpus, emerged by 22.III-13.VI.
1973. Yucatan: Mérida, 24.VII.1944, N. Souza, through G. N. Wolcott
44-19729, ex Lonchocarpus longistylus Pittier. COSTA RICA. Guana-
caste: Camnas area, 2.1J.1969, D. H. Janzen, reared ex Lonchocarpus:
7 km. n. Canfas, 6.11.1970, D. H. Janzen reared ex Lonchocarpus; La
Pacifica, 7 km. n. Canfas, 2.IIJ.1972, D. H. Janzen #VI-20-1972-XXV,
reared ex Lonchocarpus, emerged by 20.VI.1972; 1.3 mi. e. Playa Coco,
3.XII.1970, D. H. Janzen #307, reared ex Lonchocarpus costaricensis
Donn. Sm.; Santa Rosa, 15.1JI.1972, D. H. Janzen #VI-19-1972-XXVIII,
reared ex Lonchocarpus, emerged by 19.V1.1972. VENEZUELA. Aragua:
45 km. e. Sombrero, 30.1.1964, W. H. Cross, “cultivated cotton.”
Remarks: Specimens of Ctenocolum tuberculatum are readily dis-
tinguished from specimens of C. acapulcensis, C. janzeni, and C. salvini
by having only 3 facet rows behind the ocular sinus rather than 6 or
more. We have examined specimens of two closely related species from
Colombia and Venezuela which agree in eye structure but differ in
structure of the sclerite of the internal sac of the male genitalia and, in
at least one of these two species, lack of dark median stripe on female
pygidium.
THE CROTONAE GROUP
Diagnostic combination: The Crotonae Group is well distinguished
from the Tuberculatum Group by each of the following characteristics.
Hind femur: pecten with 6-10 teeth, teeth from second of series to last
gradually increased in size to middle then decreased to apex. Hind
tibia: lateral coronal tooth sharply developed, set off from mucro by
deep emargination. Male median lobe: internal sac without large com-
plex sclerite; ventral valve truncate, sides strongly sinuate.
Further characteristics of the Crotonae Group include the following:
eyes not sex-dimorphic; ocular sinus deep, 4—5 rows of facets behind
sinus; antenna not sex-dimorphic, or male antenna elongate; lateral sub-
basal gibbosities of pronotum weak to moderate; placement of outer
tooth of basal elytral gibbosity well basad of inner tooth; and hind
femur with external ventral series of denticles strong in all species.
Distribution: Northern Mexico to southern South America, and West
Indies. Four species are known from Mexico and Central America and
are treated in this paper. At least two Central American species extend
to South America. Several additional species occur in South America
and the West Indies.
Taxonomic notes: Though this group is less poorly known than the
292 Proceedings of the Biological Society of Washington
Tuberculatum Group, various taxonomic problems are evident. The four
known Central American species are separable from one another without
difficulty, but southern limits for one of them, C. crotonae are uncertain.
The name Bruchus podagricus Fabricius (1801) pertains, we think, to
a closely related West Indian form; type material of this form (in
Copenhagen) needs to be studied further. Several other South American
forms are also closely related, and indeed may not be distinct from C.
crotonae.
Remarks: Larvae of species of the Crotonae Group devour seeds of
Bergeronia, Lonchocarpus, Muellera, and Piscidia. The host ranges for at
least some of the species are rather broad. Three of the known Central
American species are known from two or more species of Lonchocarpus,
and one of them is known also from Piscidia.
We know little about adult activity. One specimen of a Brazilian
form was taken at a light trap in January. Other specimens of South
American and West Indian forms were collected at various times through-
out the year.
5. Ctenocolum colburni, new species
Description: Ctenocolum, Crotonae Group. Length 2.5-2.8 mm.
Width 1.6-1.9 mm. Integument largely rufous; dorsum of head, anterior
median portion of pronotal disc, and outer two-thirds of elytron piceous;
hind tibia rufopiceous; antennal articles 1-5 partly and articles 6-11
wholly infuscated externally; front and middle legs testaceous. Vestiture
mixed black and white, variegated in density; black only on dorsal
surface as shown, characteristic (Fig. 5). Head (Fig. 13); antennal
form as in C. crotonae (Fig. 21-22); eyes large, interocular ratio about
0.18-0.19; ocular sinus deep, 5-6 rows of facets behind sinus. Pronotum
and elytra (Fig. 5); lateral subbasal gibbosities of pronotum nearly
obsolete; basal elytral gibbosities displaced far behind base, paired teeth
of each relatively small and proximate. Hind leg (Fig. 27); pecten
with about 6 teeth, wide gap between first and second. Pygidium (Fig.
36). Male genitalia (Fig. 51-52).
Type material: Holotype male, “HOND. Cuyamel 17.VIII.1924
MACarleton,” “Lonchocarpus pentaphyllus (Poir.) D. C. USNM herb.”;
in United States National Museum of Natural History, Washington,
type no. 72800.
Paratypes, Mexico to Costa Rica (Fig. 62), 8; in USNM. MEXICO.
Veracruz: Coyame, Lake Catemaco, 14.VII.1971, Clark et al. GUATE-
MALA. Chimaltenango: road between Chimaltenango and San Martin
Jilotepeque 1500-1900’, along stream, 25.XI.1938, P. C. Standley #57907
(NY), ex herbarium specimen of Lonchocarpus purpureus Pittier. HON-
DURAS. Colén: Cuyamel, 17.VIII.1924, M. A. Carleton #634 (US), ex
herbarium specimen of Lonchocarpus pentaphyllus (Poir.) DC. COSTA
RICA. Heredia: San Miguel, 20.IIJ.1971, D. H. Janzen #656, reared ex
Lonchocarpus velutina Benth.
Some Central American seed beetles 293
. janzeni.
2C
>
us
Fic. 1-2, Ctenocolum spp., habitus. 1, C. acapulcens
PLATE l.
294 Proceedings of the Biological Society of Washington
; 4, male.
3, female
°
°
Fic. 3-4, Ctenocolum tuberculatum, habitus
PLATE 2.
Some Central American seed beetles 995
as NS
5, C. colburni; 6, C. martiale.
Fic. 5-6, Ctenocolum spp., habitus
- J gt) Ke
ySeucpe S
PLATE 3.
996 Proceedings of the Biological Society of Washington
Seni
Se,
¢
PSP a>: By
3
8, C. crotonae.
1;
7, C. biolley
Fic. 7-8, Ctenocolum spp., habitus
PLATE 4.
Some Central American seed beetles 297
Puate 5. Fic. 9-15, Ctenocolum spp., head: 9, C. acapulcensis;
10, C. tuberculatum; 11, C. janzeni; 12, C. martiale; 13, C. colburni;
14, C. biolleyi; 15, C. crotonae.
298 Proceedings of the Biological Society of Washington
16
20
21
22
23
PLATE 6. Fic. 16-23, Ctenocolum spp., antenna: 16, C. acapulcensis,
male; 17, C. janzeni, male; 18, C. salvini, male; 19, C. tuberculatum,
male; 20, same, female; 21, C. crotonae, male; 22, same, female; 23,
C. martiale, male.
Some Central American seed beetles 299
PLATE 7. Fic. 24-26, Ctenocolum spp., hind leg: 24, C. acapulcensis;
25, C. tuberculatum; 26, C. janzeni.
300 Proceedings of the Biological Society of Washington
;
27, C. colburn
°
°
Ctenocolum spp., hind leg
?
. 27-30
iolley
FIG
PLATE 8.
30, C. crotonae.
?
i
b
5 Osh, (G
. martiale
28, C
301
Some Central American seed beetles
acapul-
C
C
mvagye. eal
Ctenocolum spp., pygid
-38,
31
tuberculatum
Fic.
PLATE Q.
.
>
janzent
34, C.
33, same, female;
male;
>
colburn
a Aan G
35, C. crotonae
Censts
i
38, C. biolley
>
37, C. martiale
1;
C
?
36
>
302 Proceedings of the Biological Society of Washington
Tan
AES
ON
Some Central American seed beetles 303
Remarks: Ctenocolum colburni is named to honor John Colburn
Bridwell’s outstanding contributions to knowledge of the Bruchidae.
This particularly handsome species is immediately distinguishable
from all other members of the genus by the color pattern. No other
Central American species is closely related. An undescribed South
American species has similar elytral and pronotal gibbosities, femoral
pecten, and male genitalia but lacks a dark spot on the pronotum and
has a more diffuse elytral pattern. This undescribed species is repre-
sented in USNM material by a long series from Guyana labelled as from
Muellera seeds; this host record needs confirmation.
6. Ctenocolum biolleyi, new species
Description of female: Ctenocolum, Crotonae Group. Length 4.4
mm. Width 2.8 mm. Integument largely dark rufous, pronotum and
elytra with piceous maculation; (antennal color not known); front and
middle legs rufotestaceous. Vestiture variegated; mostly pale tan on
venter, pleura, and pygidium; elytra and central part of pronotal disc
variegated mostly with brown orange and black (Fig. 7). Head (Fig.
14); (antennal form not known); eyes large, interocular ratio about 0.23;
ocular sinus deep, 5 rows of facets behind sinus. Pronotum and elytra
(Fig. 7); lateral subbasal gibbosities of pronotum moderate; basal elytral
gibbosities not displaced far behind base, paired teeth of each large,
outer tooth much nearer to base than to inner tooth. Hind leg (Fig. 29);
pecten with about 10 teeth, no gap between first and second but second
minute. Pygidium (Fig. 38).
The male of this species is unknown.
Type material: Holotype female, “San Jose Costa Rica,” “P. Biolley
Collector,” “Buchus? Teosite (Rheana luxurians”’; [sic], “Chittenden
Collection”; in United States National Museum of Natural History, Wash-
ington, type no. 72801.
Paratype, Costa Rica (Fig. 58), 1; in USNM. COSTA RICA. San
Jose: San Jose, P. Biolley.
The holotype lacks the head, and the paratype lacks the elytra.
Measures of length and width are based on the holotype and of
interocular ratio on the paratype.
Remarks: Ctenocolum biolleyi is named for Paul Biolley, whose
turn-of-the-century collections contributed greatly to knowledge of Costa
Rican invertebrates. This species, the largest of the Crotonae Group, is
related to C. crotonae and C. martiale but differs markedly from both by
the peculiar positioning of the teeth of the basal elytral gibbosities.
<
PuatE 10. Fic. 39-44, Ctenocolum spp., male genitalia: 39, C.
salvini, median lobe; 40, same, lateral lobes; 41, same, median lobe,
lateral aspect; 42, same, lateral lobes, lateral aspect; 43, C. acapulcensis,
median lobe; 44, same, lateral lobes.
304 Proceedings of the Biological Society of Washington
—
LEIALAL-
<S =
Some Central American seed beetles 305
7. Ctenocolum martiale, new species
Description: Ctenocolum, Crotonae Group. Length 2.6—4.4 mm.
Width 2.0-2.6 mm. Integument largely piceous above, dark rufous or
rufopiceous below; antenna externally with articles 4-10 piceous and
other articles infuscated in male, and with articles 4-10 but slightly
infuscated in female; front and middle legs and hind tarsus rufous in
male and with front and middle femora and tibiae extensively infuscated,
much paler in female. Vestiture sex-dimorphic, variegated; dorsum (Fig.
6) with orange brown and black pattern on pronotum, and with elytra
almost wholly black except for orange brown in scutellar region and
small white areas (male) or with extensive orange brown and white
variegation (female); pygidium light orange brown basally, light tan
apically; mesepisternum, metepisternum, and dorsum of hind femur
light orange brown; venter white, sparse. Head (Fig. 12); antenna
(Fig. 23) serrate, extended to about elytral humerus in female and to
about elytral basal third in male; eyes large, interocular ratio 0.21-0.23;
ocular sinus deep, 4—5 rows of facets behind sinus. Pronotum and
elytra (Fig. 6); lateral subbasal gibbosities of pronotum moderate; basal
elytral gibbosities not displaced far behind base, paired teeth of each
large, outer tooth about as close to base as to inner tooth. Hind leg
(Fig. 28); pecten with about 8 teeth, no gap between first and second
but second much smaller. Pygidium (Fig. 37). Male genitalia (Fig.
53-54).
Type material: Holotype male, “ex Lonchocarpus cruentus Lundell
M. Sousa #3353,” “MEX. Ver. Laguna del Majahual, Los Tuxtlas, 100
m. 30 Nov. 1967”; in United States National Museum of Natural History,
Washington, type no. 72799.
Paratypes, Mexico (Fig. 61) and Trinidad, 13; in CNC, USNM.
MEXICO. Guerrero: Acapulco, X.1894-III.1895, E. Palmer #589
(US), ex herbarium specimen of Lonchocarpus constrictus Pittier;
Acapulco, X.1894-ITI.1895, E. Palmer #226 (US), ex herbarium speci-
men of Lonchocarpus eriocarinalis Micheli. Oaxaca: Loma del Chivo,
50 m., Chiltepec, Tuxtepec, 9.V.1968, M. Sousa #3664, ex herbarium
specimen of Lonchocarpus cruentus Lundell. Veracruz: Laguna del
Majahual, 100 m., Los Tuxtlas, 30.XI.1967, M. Sousa #3353, ex her-
barium specimen of Lonchocarpus cruentus Lundell. TRINIDAD. 3 mi.
w. Mayaro, 14.VIII.1969, H. & A. Howden.
Remarks: We name this species, which destroys seeds of Loncho-
carpus, after Mars, the Roman god of war: from Latin, “Martialis’ =
<
PLATE 11. Fic. 45-50, Ctenocolum spp., male genitalia: 45, C.
janzeni, median lobe; 46, same, sclerite of endophallus, lateral aspect;
47, same, lateral lobes; 48, C. tuberculatum, median lobe; 49, same,
lateral lobes; 50, same, endophallus, lateral aspect.
306 Proceedings of the Biological Society of Washington
WSEAS
Anas gio esp Same
See
99
S
~
RSs
SYN ON
SSX 2
VES AS
= a5
—— >
dg
SSR
SS
See ey ‘
Te Sar
Cet tS
Peg
SOS Ale
Se ENS
Some Central American seed beetles 307
“of Mars.” We dedicate Ctenocolum martiale to Mario Sousa, who in
the course of studies of Lonchocarpus provided us with important ma-
terial of this and other species of Ctenocolum.
8. Ctenocolum crotonae (Fahraeus), new combination
Bruchus crotonae Fahraeus 1839:123. Type-locality: Brazil. Type-
depository: Naturhistoriska Riksmuseum, Stockholm.
Pseudopachymerus crotonae: Pic 1913:10.
Caryedes crotonae: Blackwelder 1946:758.
Bruchus pictifemur Sharp 1885:446. Type-locality: Jalapa, Veracruz,
Mexico. Type-depository: British Museum (Natural History), London.
New synonymy.
Pseudopachymerus pictifemur: Pic 1913:11.
Caryedes pictifemur: Blackwelder 1946:758.
Description: Ctenocolum, Crotonae Group. Length 2.0-3.4 mm.
Width 1.4-2.0 mm. Integument and vestiture color and variegation much
as in C. martiale except that sex-dimorphism is much less developed
(most strongly in South America); integument of males and females
varied from largely rufous to largely piceous; antennae in most specimens
not strongly infuscated; pygidium in some specimens with vestiture
orange brown nearly to apex. Head (Fig. 15); antenna (Fig. 21-22)
serrate, extended to about elytral humerus in female and to about basal
third of elytron in male; eyes large, interocular ratio 0.17—0.20; ocular
sinus deep, 4—5 rows of facets behind sinus. Pronotum and elytra (Fig.
8); lateral subbasal gibbosities of pronotum moderate; basal elytral
gibbosities not displaced far behind base, paired teeth of each large, outer
tooth closer to inner tooth than to base. Hind leg (Fig. 30); pecten
with 7-9 teeth, no gap between first and second but second much
smaller. Pygidium (Fig. 35). Male genitalia (Fig. 55-56).
Ctenocolum crotonae is distinguished from the related C. martiale by
the male genitalia, with conspicuous differences in form of ventral valve,
form of lateral lobe apices, and armature of internal sac; by larger eye
size and smaller interocular ratio; and by positioning of the teeth of the
basal elytral gibbosities. Several South American forms are much less
distinct and perhaps are conspecific but are here distinguished from
C. crotonae by having more reduced pronotal markings rather than a
nearly continuous central macula.
Material examined: Mexico to Costa Rica (Fig. 60) and northern
South America, 100. MEXICO. Campeche: head of Rio Candelaria,
<
PuaTE 12. Fic. 51-56, Ctenocolum spp., male genitalia: 51, C.
colburni, median lobe; 52, same, lateral lobes; 53, C. martiale, median
lobe; 54, same, lateral lobes; 55, C. crotonae, median lobe; 56, same,
lateral lobes.
308 Proceedings of the Biological Society of Washington
PuaTE 13. Fic. 57-59, Ctenocolum spp., distribution records in
Central America: 57, C. janzeni (open symbol = state record); 58, C.
acapulcensis (1), C. salvini (2), C. biolleyi (3); 59, C. tuberculatum
(s = type locality of serratissimus, t = type locality of tuberculatum).
Some Central American seed beetles 309
Puate i4. Fic. 60-62, Ctenocolum spp., distribution records in
Central America: 60, C. crotonea (open symbol = type locality of picti-
femur); 61, C. martiale; 62, C. colburni.
310 Proceedings of the Biological Society of Washington
Rancho San José, Rio San Pedro, 27.VJ.1957, O. G. Enriquez #B48
(US), ex herbarium specimen of Lonchocarpus hondurensis Benth.
Chiapas: Ejido Morelos, La Libertad 60 m., 20.IIJ.1968, M. Sousa
#3657 (US), ex herbarium specimen of Lonchocarpus rugosus Benth.
Jalisco: 34 mi. nw. Barra de Navidad, 9.III.1973, C. D. Johnson
# 438-73, reared ex Lonchocarpus nitidus (Vog.) Benth., emerged by
12.IV-23.V1.1973. Sinaloa: Mazatlan, 14.VIII.1965, H. Burke & J. Meyer.
EL SALVADOR. La Unidén: Volcan Conchagua, 27-29.V.1958, L. J.
Bottimer #101lu, reared ex Lonchocarpus. San Salvador: San Salvador,
650-850 m., 20.XXI.1921-4.1.1922, P. C. Standley #19254 (US), ex
herbarium specimen of Lonchocarpus rugosus Benth. COSTA RICA.
Guanacaste: La Pacifica, 7 km. n. Cafias, 2.III.1972, D. H. Janzen
# VI-20-1972-XXV, reared ex Lonchocarpus, emerged by 20.VI.1972;
Santa Rosa, 15.]II.1972, D. H. Janzen #VI-19-1972-XXVIII, reared ex
Lonchocarpus, emerged by 19.VI.1972; Santa Rosa, 1% km. from highway,
15.11.1972, D. H. Janzen #VI-19-1972-XXIV, reared ex Lonchocarpus,
emerged by 19.VI.1972. Puntarenas: 4 km. n. Puntarenas junction,
6.111.1972, D. H. Janzen #VI-20-1972-X, reared ex Piscidia cartha-
genesis Jacq., emerged by 20.VI.1972. VENEZUELA. Distrito Federal:
Caracas, 22.XII.1944, interception #EQ#087748 ex Lonchocarpus mar-
garitensis Pittier. TOBAGO. Interception #PQ#022305 (Washington )
ex “seeds of black manoe,” 17.VIII.1933. GUYANA. Interception
#EQ#A33506 ex “seeds of Peltophorum dosyrachis,’ 20.XI1I.1935; in-
terception (from botanic garden) #EQ#A33506 ex Lonchocarpus
pentaphyllus (Poir.) DC., 20.X.1935; interception (from botanic garden)
#BPQ#030932. BRAZIL. Mato Grosso: Borda da Mato Sertaozinho,
12.1.1960, Medeiros & Pereira. ECUADOR. Intercepted at Honolulu ex
“tree seeds.”
The specimen from “Peltophorum dosyrachis” is doubtless mislabelled,
as it bears the same quarantine interception numbers as a series from
Lonchocarpus pentaphyllus.
Remarks: There appear to be several closely related forms in the
West Indies and South America. The male genitalia of these probably
different species differ from the genitalia of C. crotonae particularly
in structures of the internal sac. Also, dark markings of the pronotum
are reduced to a pair of broken longitudinal vittae, rather than con-
fluent or nearly so as in C. crotonae.
Ctenocolum crotonae is known from more species of host plants than
is any other member of the genus, and it is the only species of Ctenocolum
so far known from more than one host genus. The records of Loncho-
carpus margaritensis and L. pentaphyllus are based on quarantine inspec-
tions and need confirmation. Several collections of reared material from
undetermined species of Lonchocarpus include both C. crotonae and C.
tuberculatum, and one rearing from Piscidia carthagenensis included
C. janzeni as well as C. crotonae.
Some Central American seed beetles 311
DISCUSSION
Since several of the Central American species of Ctenocolum are known
from only one or two localities each, several additional species are likely
to be found. Much remains to be learned about the natural history of
the genus. We can confidently state only that the genus is restricted to
various members of the subtribe Lonchocarpinae (tribe Dalbergiae,
family Fabaceae), and that no other New World bruchids are known to
use these plants as hosts. We can draw no useful conclusions about
geographic relationships, save that the greatest evolutionary activity for
both species groups seems to be in Central America and northwestern
South America. Similarly, distributional data are of no help in a
phylogenetic reconstruction of the relationships of the species.
We recognize two apparently natural species groups, but relationships
of their included species are unclear and perhaps cannot be adequately
worked out until South American species are studied. The Tuberculatum
Group may be defined as monophyletic by having as an apomorphous
characteristic a large, complex, symmetric sclerite in the male endo-
phallus. The Crotonae Group may similarly be defined by having the
ventral valve of the male median lobe truncate and by having the pecten
of the hind femur crescentic in outline.
Within the Tuberculatum Group, C. tuberculatum is distinguished by
having the eye not sex-dimorphic, the male interocular ratio over 0.20,
and the male lateral lobes divided to near the base (probably all
plesiomorphous states); and by the ocular sinus deep, with 3 facet rows
behind the sinus, and by the male lateral lobes not rolled over at the
tips (probably apomorphous states). Perhaps the form of the ocular
sinus (deep in C. tuberculatum, moderate in C. janzeni and C. salvini,
and shallow in C. acapulcensis) is a true morphocline, with the deep
sinus a plesiomorphous condition. Among the other three species in
Central America, C. acapulcensis stands out by having the male antenna
flabellate, the ocular sinus shallow and with about 10 facet rows behind
the sinus, and the external ventral denticles of the hind femur obsolete.
All of these conditions, however, are apomorphous, and the possession
of plesiomorphous states of these characters does not indicate an un-
equivocal relationship between C. janzeni and C. salvini. The sclerite
of the male endophallus is relatively simplified in C. acapulcensis and
C. salvini, perhaps as a synapomorphous condition. Similarly, the eyes
are strongly sex-dimorphic in C. acapulcensis and C. janzeni, an equally
plausible synapomorphy.
Among Central American members of the Crotonae Group, C. colburni
is certainly the most distinctive species and perhaps therefore is the least
related, but we have discovered no convincing synapomorphous condi-
tions to relate the other three species. A clue to relationships among
C. crotonae, C. martiale, and C. biolleyi lies in the probable morpho-
cline of increasing basal displacement of the outer tooth of the basal
elytral gibbosity. Thus, C. martiale and C. biolleyi, in which this dis-
312 Proceedings of the Biological Society of Washington
placement is greatest, are probably the most closely related species of
the Crotonae Group.
LITERATURE CITED
BLACKWELDER, R. E. 1946. Checklist of the coleopterous insects of
Mexico, Central America, the West Indies, and South America.
Bull. U. S. Nat. Mus. 185:551-763.
FAuRAEuS, O. I. von. 1839. In C. J. Schoenherr, Genera et species
curculionidum cum synonymia hujus familiae. Paris 5:1—456.
KINGSOLVER, J. M. AND D. R. WHITEHEAD. (In press). Classification
and comparative biology of the seed beetle genus Caryedes
Hummel (Coleoptera: Bruchidae). Trans. Amer. Ent. Soc.
MortscHoutsky, V. 1873 (1874). Enumération des nouvelles espéces
de Coléoptéres rapportés de ses voyages. Bull. Soc. Imp. Nat.
Moscou 46:203—252.
Pic, M. 1913. Coleopterorum catalogus, Pars 55, Bruchidae. Junk,
Berlin, 74 pp.
SuHarRP, D. 1885. Biologia Centrali-Americana, Insecta, Coleoptera,
Bruchidae 5:437-504.
WHITEHEAD, D. R. AND J. M. KiNcGsoLvER. (In Press). Biosystematics
of the North and Central American species of Gibbobruchus
(Coleoptera: Bruchidae: Bruchinae). Trans. Amer. Ent. Soc.
al
ore
Wolk, GI, ING, Yeh joes Sls = sw 22 October 1974
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
UCA PANACEA, A NEW SPECIES OF FIDDLER CRAB
FROM THE GULF COAST OF THE UNITED STATES
By ALLEN Novak AND MICHAEL SALMON
Department of Zoology, University of Illinois,
Urbana, Illinois 61801
This paper documents the presence of a new species of
fiddler crab, Uca panacea, on the Gulf Coast of the United
States. A morphological description of the larval and adult
specimens is given here, with special reference to those char-
acters which distinguish the new form from U. pugilator, with
which it has been confused.
Most of the North American fiddler crabs (Genus Uca)
are found along the Atlantic and Gulf Coasts of the United
States. Rathbun (1918) first attempted to list all of these
species and describe them. She listed 9 species, including Uca
pugilator (Bosc), the most common form, which is found
from Cape Cod, Mass., to Florida and along the Gulf Coast
komme xas:
Since her initial study, some new species have been de-
scribed. Tashian and Vernberg (1958) designated U. pugnax
and U. rapax (considered by Rathbun to be subspecies) as
distinct species. Salmon and Atsaides (1968) described two
new species, U. longisignalis and U. virens, as distinct from
the Atlantic populations of U. pugnax with which they had
previously been confused. Subsequently, these findings were
confirmed by Selander, Johnson and Avise (1971) employing
as their measure electrophoretic analyses of variation in
muscle proteins.
Rao and Fingerman (1968) described two variants in colora-
tion pattern among “U. pugilator” from the Panacea area.
Each variant exhibited different chromatophoric response to
28—Proc. Brot. Soc. WAsH., Vou. 87, 1974 (313)
314 Proceedings of the Biological Society of Washington
black and white backgrounds. The study by Selander, et al.
(1971) led to the discovery that Gulf Coast specimens,
thought to be U. pugilator, could be placed in two distinct
groups. That finding led to this morphological study. In
another paper (in preparation), behavioral differences be-
tween the two forms are considered. The results indicate
that U. panacea new species exists sympatrically but distinct
from U. pugilator on the Gulf Coast of the United States.
METHODS
Uca panacea new species was found and studied within a
ten-mile radius of the Florida State University Marine Labora-
tory, Carrabelle, Florida. Several large populations were
found on the beach areas of Saint Mark’s wildlife preserve,
just east of Panacea, Florida. Animals from this location were
used in this study.
About 150 individuals were collected and brought back
alive to the University of Illinois, Urbana. The sample con-
sisted of males and females from 1.1 to 1.8 cm in carapace
width. The crabs were maintained in a large, 400 liter aquar-
ium, 1.9 m long by 58 cm wide), provided with a bank of
sand. These crabs, which were observed for behavioral studies,
were also measured.
Characters measured with vernier calipers (accurate to 0.01
mm) were the following: width of front; width of carapace
at its greatest dorsolateral extension; on large chela—length
of propodus (from tip of finger to proximal edge), dactyl
length (from tip to upper point of articulation with propodus ),
and dactyl width from ventral tooth ridge to dorsal margin
midway from tip to point of articulation with propodus. Re-
gressions for these measures were established for each species.
The figures were then subjected to statistical analysis by
comparison of regression slopes. In other cases, means of
certain measurements were compared, using a t-test. The 0.05
level was chosen to reject the null hypothesis of no significant
difference between the samples.
The number, distribution, and form of the spoon-tipped
hairs on the merus of the second maxilliped were examined.
Samples of 14 maxillipeds from 10 males and 4 females from
A new fiddler crab from Gulf of Mexico BS
Fic. 1. Uca panacea, new species, allotype female. Top, dorsal view;
bottom, frontal view.
U. panacea new species were photographed and compared to
4 maxillipeds from U. pugilator, as well as previous descrip-
tions of the latter ( Miller, 1961). The hair tips for both species
were photographed under a scanning electron microscope
(magnification 450-900 x). Hairs used for electron micros-
copy were taken from fresh maxillipeds that had been placed
in 95% ethyl alcohol for at least 72 hours. The hairs were
mounted on pedestals and gold-palladium coated in a vacuum
chamber before being photographed.
Five egg-bearing females were sent to the Duke University
Marine Laboratory, Beaufort, North Carolina. The larvae
were reared to the second crab stage at a temperature of 25
Ge cheess OG msalimity, OL oo) ppt. and sohnotopernod: 12) 1h: 2D)
using established techniques (Costlow and Bookhout, 1962).
A few (10-20) larvae at each stage were preserved in a solu-
316 Proceedings of the Biological Society of Washington
Fic. 2. Uca panacea, new species, holotype male. Top, dorsal view;
bottom, frontal view.
tion of 4% formalin buffered to pH 7 with hexamethylenamine.
Larval description is based primarily upon preserved speci-
mens, although details were often checked against the mor-
phology of living larvae.
Ueca panacea, new species
Etymology: Panacea, alluding to the location where this animal was
first discovered.
Distribution: Specimens of U. panacea described here were collected
from Saint Mark’s Wildlife Preserve just east of Panacea, Florida. They
are known to occur west to Texas (Selander, personal communication ).
Types: Deposited in the Smithsonian Institution. Holotype, male,
USNM 150096; allotype, female, USNM 150097; paratypes, 50 males,
50 females, USNM 150098.
Sample size: 250 preserved and 150 live animals of various sex and
size combinations were used for this study.
Morphological description: (Figs. 1 and 2) Carapace moderately
A new fiddler crab from Gulf of Mexico OMly/
Fic. 3. Uca panacea, new species. Left, major cheliped: top, in-
terior view; bottom, exterior view. Right, minor cheliped: top, exterior
view; bottom, interior view.
arched, widest at anterolateral margins. Lateral margins parallel. Frontal
margin transverse, angulate, forming obtuse angle on either side of
protruding front. Surface of carapace smooth. H-form cardiac depres-
sion moderately outlined. Anterolateral margins distinct but blending
into smooth surface of carapace posterior to middle of cardiac region.
A ridge above last 2 paris of walking legs on oblique lateral wall of
carapace. Oblique and anterolateral margins converging anteriorly to
form less than right angle. Carapace about 2.8 times wider than front
but these proportions vary with size.
Minor cheliped (Fig. 3) slightly serrated along inner margin of lower
finger except near flattened spoon tip. Distinct gap present when fingers
are closed. Scattered row of hairs extending along margins of both
fingers.
Outer surface of major cheliped (Fig. 3) smooth but covered with
many small flat tubercles. Propodus about 1.30 times as long as dactylus
but proportions vary with size. Dactyl 8.25 times as long as wide but pro-
portions vary with size and maturity. Submarginal ridge on palm not
developed and lacking distinct tubercles. Teeth of both fingers well
developed. Inner row of teeth continuing along palm usually extending
close to lateral margin of carpal cavity.
Carpus with oblique ridge on inner surface continuing proximally
along upper margin of carpus with slightly developed tubercles to point
of articulation with merus. Lower margin having distinct ridge con-
taining tubercles extending proximally to point of articulation with merus.
Supplementary specific characteristics: (Figs. 4 and 5) Spoon-tipped
hairs of second maxilliped totaling about 160, projecting beyond inner
margin of merus with many more non-projecting shorter hairs, Hairs
confined to wide zone on median and distal inner edge of merus. Spoon-
318 Proceedings of the Biological Society of Washington
Fic. 4. Electron micrographs of spoon-tipped hairs from the second
maxilliped. Left (A and C), Uca pugilator; right (B and D), U.
panacea, new species. Note that tips are cup-shaped in the former, but
more spatulate in the latter.
tipped process bearing 8-9 enlarged lateral lobes which continue as
smaller lobes and extend down the shaft of the hair (Fig. 4).
Abdominal appendage with well developed arm. Curvature of tip
continuing to follow that of shaft of appendage (Fig. 5).
Color: Carapace light grey-brown to uniformly creamy olive brown,
being slightly darker posteriorly and laterally; the cardiac H-form de-
pression is uniformly rust-red. Major cheliped of males white at fingers.
Outside propodus orange to purple, palm usually being slightly lighter.
Inner surface of carpus deep orange to red. Ambulatories white and
speckled or uniform light red. Females with same color pattern. Color
pattern of all animals altered with environmental conditions. Often
carapace is considerably darker. When carapace is light colored, a pair
of small grey spots can occasionally be seen slightly anterior to H-form
depression.
A new fiddler crab from Gulf of Mexico Bile)
Fic. 5. Right abdominal appendage, seen from an anterior view.
Right, Uca panacea, new species; left, U. pugilator.
Morphology of larvae: First zoea (Fig. 6-1); carapace dorsoventrally
elongated, bearing dorsal and rostral spines. Rostral spine longer than
dorsal. No lateral spines present. Carapace smooth with no sharp
depressions or protuberances. Abdomen composed of 5 segments, 5th
segment coalesced with telson. First abdominal segment predominantly
covered by carapace. First and second segments show blunt lateral
spines. Posterolateral margins of abdominal segments 2-5 overlap fol-
lowing segment with short blunt points, posterodorsal margin of each
segment bears short hairs. Exopodite of maxilliped bears 4 natatory
setae. Eyes sessile.
Second zoea (Fig. 6-II); carapace primarily unchanged with excep-
tion of slight protuberance just posterior to dorsal spine. Growth and
expansion have occurred in carapace. Dorsal surface of first abdominal
segment bears single setae centrally. Posterolateral edges of abdominal
segments longer than in Ist stage and more heavily constructed. Ex-
opodite of maxilliped bears 6 plumose setae. Eyes are now stalked.
Third zoea (Fig. 6-III); carapace enlarged with no apparent lengthen-
ing of dorsal and rostral spines over 2nd stage. Expansion of carapace
gives dorsal spine stout appearance. Rostral spine still longer than
dorsal. Carapace maintains smooth surface. Three setae appear in
posteroventral margin of carapace; 3 setae are borne by, and are equally
spaced along dorsal margin of first abdominal segment. First abdominal
segment partially covered by carapace. Abdomen now composed of 6
segments, 6th bearing telson. Segments 3-5 have posterolateral projec-
tions which appear as short spines. Exopodite of maxilliped typically
bears 8 plumose setae.
Fourth zoea (Fig. 6-IV); carapace remains smooth. Rostral and
dorsal spines maintain previous size as carapace and abdomen continue
to grow. First abdominal segment (hidden under carapace) bears 6
320 Proceedings of the Biological Society of Washington
mm
Frontal and lateral views of the
Uca panacea, new species.
Fic. 6.
five zoeal stages.
A new fiddler crab from Gulf of Mexico oui
Fic. 7. Uca panacea, new species. Lateral (A) and dorsal (B)
view of megalopa.
setae equally spaced along entire dorsal margin. Pleopod buds appear
on abdominal segments 2-5. Spiny posterolateral projections on seg-
ments 3-5 are maintained. Maxilliped typically bears 10 plumose setae.
Fifth zoea (Fig. 6-V); carapace remains smooth. Eyes large, stalked.
First abdominal segment (under carapace) bears 11 setae equally spaced
along dorsal margin. Pleopods are more developed and occur on seg-
ments 2-5. Posterolateral margins of abdominal segments 3-5 have
short spines which overlap following segment. Exopodite of maxilliped
typically has 12 plumose setae.
Gross morphology of megalopa (Fig. 7); carapace oval when viewed
from above, rounded on dorsal surface by 3 protuberances running from
anterior to posterior surface. Chelipeds formed but small. Abdomen
consists of 6 segments (Ist segment hidden by carapace) plus telson.
Segments 2-6 have bisegmented pleopods. Pleopods typically bear 15
setae each segment.
Gross morphology of first crab: (Fig. 8-A) Carapace round when
viewed from above, extension of carapace between stalked eyes has
formed. Depressions and protuberances slight and not well defined.
Chelipeds are developed but small and equal in size. Abdomen (not
visible in figure) carried under body where it fits into midventral
depression.
Gross morphology of second crab: (Fig. 8-B) Carapace round when
viewed from above, with grooves for horizontal reception of eyestalks.
Extension of carapace between eyestalks now fully developed. “U’-
322 Proceedings of the Biological Society of Washington
a
1 mm
Fic. 8. Uca panacea, new species. A, dorsal view of first crab stage;
B, anterodorsal view of second crab stage.
shaped slight depression sometimes apparent on dorsal surface of cara-
pace, beginning posterior to eyestalks and arching posteriorly where
arcs meet middorsally. Abdomen remains folded under ventral portion
of body where it is retained in a depression.
A new fiddler crab from Gulf of Mexico 323
COMPARISON WiTH UCA PUGILATOR
While U. pugilator in the Carrabelle-Panacea area of Florida com-
monly reach a size greater than that of the largest U. panacea, the typical
size of an adult U. panacea is in keeping with that of most mature U.
pugilator. A typical carapace width for U. panacea in Carrabelle,
Florida, is 1.6 cm, while U. pugilator commonly attains a width of 2.0
em and occasionally larger. From a random sample of 500 animals
collected, the greatest carapace width of U. pugilator was 1.91 cm while
that of U. panacea was 1.63 cm.
Regressions were established using carapace width as a function of
width of front. A random sample (20 of each sex for each species) was
drawn and the regressions compared using F-test to establish whether
the two regressions could be drawn from the same population. In both
sexes the statistical analysis indicated that the slopes of the regressions
of U. panacea (b = 3.1088, 9 2, and 1.9254, ¢ 4) and those of U.
pugilator (b = 2.3674, 2 9, and 2.822, ¢ 6) were different (compari-
son of females: Fu%3 = 5.7771, p = .025; comparison of males: F,361
== (2547, 1 = Ol)
Unfortunately, carapace pigmentation is variable within the two
species. However, distinctive characteristics do exist in the form of the
cardiac H-form depression, carpal pigmentation, the spots on the carapace
and overall carapace color, especially when the animals are in a lightened
state. A most distinctive characteristic is the rust-red cardiac depression
on the back of U. panacea. This differs from U. pugilator, which typically
has a dark grey-brown cardiac depression, and is maintained even when
the animals assume the dark coloration.
Occasionally U. panacea is seen with a pair of small grey spots
slightly anterior to the H-form depression. This marking is common in
U. pugilator also, but two additional zones of purple pigmentation are
located just anterior and lateral to the grey spots.
When animals assume light carapace pigmentation, differentiation
becomes simple. Uca panacea appears creamy olive-brown compared to
U. pugilator (creamy powder blue-grey carapace with deep purple
spots). The difference in cardiac H-form is at a maximum then, with
U. panacea bright rust-red compared to the greyish-brown of U. pugilator.
The broad flat inner margin of the carpus is orange to deep red in
U. panacea and distinct from the rest of the appendage segments. The
carpus of Uca pugilator, while having some red pigmentation, does not
present as bright a color as in U. panacea.
The major cheliped of male U. panacea appears more slender than
that of U. pugilator. Measurements confirmed this. Mean dactyl widths
were compared between 15 crabs of each of the two forms, using a
sample in which dactyl length was held between 1.55 and 1.59 cm. The
mean + s was 0.189 + 0.007 cm for U. panacea, and 0.221 + 0.006 cm
for U. pugilator (t = 191.3, p < .005). Mean dactyl lengths were also
compared, again using 15 crabs/sample, and holding carapace width
324 Proceedings of the Biological Society of Washington
between 1.43 and 1.49 cm. For U. panacea, the mean was 1.51 + 0.06
cm; for U. pugilator, 1.32 + 0.14 cm. These values were also signif-
icantly different (¢ = 131.4, p < .005). Thus, in the major cheliped of
U. panacea, the dactyl is more slender and longer than in U. pugilator.
Of the 30 animals measured, the range and mean of the dactyl length:
width proportion was 7.75-8.78, 8.25 for U. panacea; and 6.80-7.48,
7.09 for U. pugilator.
The overall proportion of dactyl length to propodus length is only
slightly different between the two species. Regressions were established
using dactyl length as a function of propodus length. With a random
sample of 30 for each species, the comparison of slopes (U. panacea,
b = 0.864, U. pugilator, b = 0.800) was not significant (Fujs61 = 2.3132,
p < .10). The tooth midway along the finger of the propodus is in
most cases smaller in U. panacea than it is for U. pugilator.
Spoon-tipped hairs of both species are equal in abundance and
distribution on the second maxilliped. In five U. panacea examined,
148-162 spoon-tipped hairs were counted extending beyond the inner
margin of the merus. Crane (1943) describes U. pugilator as having
150-200 spoon-tipped hairs extending beyond the inner margin of the
merus. Photographs under the scanning electron microscope however,
indicate different morphology for the two species (Fig. 4). The spoon-
tipped hairs of U. panacea are broader and shallower than those of U.
pugilator. Lateral lobes of the former extend down the shaft of the hair,
while in the latter, none are seen along the shaft.
Abdominal appendages of adult males are similar, but a difference
is apparent in the angles and ridges at the tip of the appendage (Fig. 5).
Six males of both species were compared. The appendage of U. panacea
continues to follow the curvature of the shaft to the tip of the appendage,
while in U. pugilator, it diverts away from the arm. This causes the tip
of U. pugilator to form a concave arch on the side opposite the arm.
DISCUSSION
The larval as well as adult descriptions given here form a base of
reference for species identification. The only previous description of
larval development in Uca was done by Hyman (1922) for U. pugilator.
Crane (1940) primarily refers to Hyman’s description when comparing
the Genus Ocypode to Uca. But Hyman did not rear his animals in the
laboratory. Hence, his data are suspect, both in terms of uncontrolled
environmental (salinity, temperature, nutrition) influences as well as
species identification.
The distinct differences in the form of the spoon-tipped hair is of
prime importance. According to Crane (1941), and Miller (1961), this
character is correlated with feeding adaptations to different substrates.
Thus the differences found here are indicative of niche specialization
between the two forms.
Another important character, again indicative of species distinctive-
ness, is the difference in shape of the abdominal appendages. This slight
A new fiddler crab from Gulf of Mexico 325
difference in form may, in conjunction with other adaptations, prevent
interbreeding. No intermediate forms, which could indicate hybridiza-
tion, were found in the field.
Color pattern is a distinctive characteristic differentiating the two
species, and it is now known that fiddler crabs have color vision (Hyatt,
1973). Characteristics such as the bright color of the carpus on the
major chela may coordinate vitally with the mating display of the male.
The description presented above indicates that U. panacea and U.
pugilator are closely related and have probably diverged only recently.
There is little doubt, based upon the combined evidence from _ this
description and the biochemical study of Selander et al. (1971), that
isolation is complete. The courtship displays of both species are also
distinct and will be described in a separate paper. From an evolutionary
viewpoint, this behavioral evidence must be considered the most con-
clusive for speciation (Crane, 1943; Cain, 1954; Mayr, 1966).
Acknowledgments: We thank Dr. R. Selander, Department of
Zoology, University of Texas at Austin, for his help in this study. The
Director, Staff, and Dr. W. F. Herrnkind of Florida State University
Marine Laboratory provided valuable advice, and the use of facilities
and materials. Dr. John Costlow of the Duke University Marine Labora-
tory gave us use of larval rearing facilities. Scanning electron microscope
photographs were taken by Richard Forester, Department of Paleontology,
University of Illinois. Figures 1-3 and 6-8 were drawn by Carol Coope,
Departmental Artist. This work was supported by grant GB-20766X
from the National Science Foundation.
LITERATURE CITED
Cain, A. J. 1954. Animal Species and Their Evolution. Hutchinson’s
University Library, London.
CostLow, J. D. Jr., anp C. G. Booxuour. 1962. Larval development
of Sesarma reticulatum Say reared in the laboratory. Crusta-
ceana, 4:281—-295.
Crane, J. 1940. Eastern Pacific Expeditions of the New York Zoologi-
cal Society. XVIII. On the post-embryonic development of
Brachyuran crabs of the Genus Ocypode. Zoologica, 25:65—82.
—. 1941. Eastern Pacific Expeditions of the New York Zoologi-
cal Society. XXVI. Crabs of the Genus Uca from the west
coast of Central America. Zoologica, 26:145—207.
—. 1943. Display, breeding and relationships of the fiddler crabs
(Brachyura, Genus Uca) in the northeastern United States.
Zoologica, 28:217-223.
Hyatt, G. W. 1973. Physiological and behavioral studies of color
vision in fiddler crabs (Brachyura, Ocypodidae, Genus Uca).
Ph.D. thesis, University of Illinois, Urbana.
Hyman, O. W. 1922. Adventures in the life of a fiddler crab. Smith-
sonian Report for 1920, pp. 443-460.
326 Proceedings of the Biological Society of Washington
Mayr, E. 1966. Animal Species and Evolution. Harvard University
Press, Cambridge.
Minter, D. C. 1961. The feeding mechanism of fiddler crabs, with
ecological considerations of feeding adaptations. Zoologica, 46:
89-100.
Rao, K. R., anp M. FINGERMAN. 1968. Dimorphic variants of the
fiddler crab Uca pugilator and their chromatophore responses.
Proc. Louisiana Acad. Sci. 31:27-39.
Ratusun, M. J. 1918. The Grapsoid crabs of America. Smithsonian
Institute, Bull. U. S. Nat. Mus. 97: 1-447.
SALMON, M., AND S. P. ArsaripEs. 1968. Behavioral, morphological
and ecological evidence for two new species of fiddler crabs
(Genus Uca) from the Gulf Coast of the United States. Proc.
Biol. Soc. Wash. 81:275—290.
SELANDER, R., W. JOHNSON, AND J. AvisE. 1971. Biochemical popula-
tion genetics of fiddler crabs (Uca). Biol. Bull. 141:402.
TasHian, R. E., AnD F. J. VERNBERG. 1958. The specific distinctness
of the fiddler crabs (Uca pugnax (Smith) and Uca rapax
(Smith) ) at their zone of overlap in northeastern Florida.
Zoologica, 43:89-92.
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Vol. 87, No. 29, pp. 327-336 22 October 1974
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
THE KARYOTYPE OF VANZOLINIUS DISCODACTYLUS
AND COMMENTS ON USEFULNESS OF KARYOTYPES
IN DETERMINING RELATIONSHIPS IN
THE LEPTODACTYLUS-COMPLEX
(AMPHIBIA, LEPTODACTYLIDAE)
By W. Ronatp HEYER AND M. JupirH DIMENT
Division of Amphibians and Reptiles, National Museum of
Natural History, Smithsonian Institution,
Washington, D. C. 20560, and
Biology Department, Pacific Lutheran University,
Tacoma, Washington 98447
Information on karyotypes of several species of the frog
genera Adenomera and Leptodactylus recently has become
available. Bogart (In press) described and figured the karyo-
types of 16 species of Leptodactylus and 3 species of Aden-
omera (reported as the marmoratus group members of the
genus Leptodactylus). We have obtained information on the
karyotypes of 9 species of Leptodactylus and Vanzolinius dis-
codactylus. The purpose of this paper is to describe the karyo-
type for the previously unreported V. discodactylus and offer
alternatives to the relationships among the Leptodactylus-
complex to those proposed by Bogart (In press). It should be
pointed out that the genera Adenomera, Leptodactylus, and
Vanzolinius have been considered to all belong to the genus
Leptodactylus until recently.
MetTHOops AND MATERIALS
The technique and terminology used in preparation and
description of the karyotypes follows Patton (1967). Ap-
proximately 50 cells were examined from marrow, spleen, or
testis tissue of the specimens. The material was examined
29—Proc. Bion. Soc. WAsH., Vou. 87, 1974 (SRA)
328 Proceedings of the Biological Society of Washington
Fic. 1. Karyotype of Vanzolinius discodactylus, specimen WRH 464.
Chromosome pair number 3 with overlap.
using a phase contrast microscope and the better metaphase
figures were photographed. The chromosomal spreads used in
the analysis were those with few or no overlapping chromo-
somes in the condensed state. Each arm of every chromosome
was measured and the chromosomes were grouped in pairs
according to their size and arm ratio. Our preparations did
not allow the location of secondary constrictions.
The specimens were taken from the following localities:
Leptodactylus bolivianus, melanonotus—Costa Rica. Lepto-
dactylus bufonius, chaquensis, fuscus, latinasus—Argentina:
Salta; Embarcacion. Leptodactylus mystaceus, pentadactylus,
wagneri, Vanzolinius discodactylus—Ecuador: Napo; Limon-
cocha. Specimens and slides will be deposited at the Natural
History Museum, Los Angeles County.
THE Karyotype oF Vanzolinius discodactylus
The karyotype of Vanzolinius discodactylus is characterized
as follows: diploid number = 22; three pair of metacentrics
(Fig. 1, chromosome pair numbers 5, 8, 9); four pair of sub-
metacentrics (Fig. 1, chromosome pair numbers 1, 2, 6, 10);
three pair of subtelocentrics (Fig. 1, chromosome pair numbers
3, 4, 7); one pair of acrocentrics (Fig. 1, chromosome pair
number 11); fundamental number = 42.
ré
Fic. 2. Karyotype of Leptodactylus chaquensis, specimen WRH 1402.
Karyotypic Comparisons oF Leptodactylus SpEctEs
The karyotypes of the species of Leptodactylus we exarnined
agree with previously published accounts in that all species
have a diploid number of 22.
Although Barbieri (1950) reported the diploid number of
L. chaquensis as 2n = 22, he did not include a metaphase
figure of the karyotype. We include a metaphase figure (Fig.
2) of this species for comparative purposes. There are four
pair of metacentrics (Fig. 2, chromosome pair numbers 1, 6,
8, 11); five pair of submetacentrics (Fig. 2, chromosome pair
numbers 2, 5, 7, 9, 10); and two pair of subtelocentrics (Fig.
2, chromosome pair numbers 3, 4).
Our analysis of chromosome morphology based on cen-
trometric position differs considerably from previously pub-
lished accounts (Table 1). These differences could be ac-
counted for in three different ways. First, the different
karyotypes could be the result of different techniques. Bogart
(In press) used the corneal squash technique while we have
used the marrow and spleen cell suspension technique. Sec-
ond, the differences could be due to geographic variation
within species. Third, the differences might be due to different
interpretations of morphology and of which pairs are ho-
330 Proceedings of the Biological Society of Washington
TasLE 1. Comparison of karyotypes for species of Leptodactylus.
Meta- Submeta- Subtelo- Acro-
Source Species centrics centrics centrics centrics
Bogart" bolivianus’ 8 2) 1 0
Present Study " 4 5 2; 0
Bogart bufonius 7 y 2 0
Present Study " 6 2 3 0
Bogart fuscus il 3 1 0
Present Study " 3) 3 3 0
Bogart melanonotus 6 3 D 0
Present Study " 4 4 3 0
Bogart mystaceus 5 3 3 0)
Present Study " 4 4 3 0
Bogart pentadactylus 7 3 1 0
Denaro’ " 4 3 4 0
Present Study t" 2) 6 3 0
Bogart wagneri 2 4 if 4
Present Study " 2 2 3 4
1 Bogart (In press).
2 Reported as insularum.
3 Denaro (1972).
mologous. We think that the third explanation could very
well account for many of the differences noted in the karyo-
types of Table 1. Until homologues can be determined ob-
jectively, we think that the karyotype data must be interpreted
extremely conservatively.
RELATIONSHIPS
There are two aspects of the karyotypes that appear to be
conservative in the sense that they are not influenced by
interpretations by different workers and also appear to pro-
vide information of possible use in predicting relationships.
These aspects are the diploid number and the presence or
absence of acrocentric chromosomes. Character states of these
two karyotypic characters are assigned as follows:
Diploid number of chromosomes: Character state a =
diploid number of 26. This is the primitive state of the family
(Lynch, 1971, p. 37). State A = diploid number of 24 and is
Karyotypes in Leptodactylus-complex Soll
L. albilabris, bolivianus, bufonius
chaquensis, fuscus, geminus, gracilis,
knudseni, labialis, melanonotus,
mystaceus, mystacinus, ocellatus,
pentadactylus, rhodonotus
AX*B
L. latinasus, "natalensis,"
podicipinus, wagneri;
V. discodactylus
oe
A. marmorata
|
A. andreae, hylaedactyla
ab
Fic. 3. Proposed relationships based on karyotypes. Lower case
letters indicate primitive states, capital letters indicate derived states,
asterisks indicate secondarily derived states. Character a is the diploid
number, character b is presence or absence of acrocentric chromosomes.
Also see text. Based on data from Barrio (1973), Bogart (In press),
Denaro (1972), Heyer (1972), and present study.
derived. State A* = diploid number of 22 and is secondarily
derived.
Acrocentric chromosomes: Character state b = acrocentric
chromosomes present. This is the primitive state, generally
recognized by chromosome workers, although there is no in-
formation to support this from the primitive members of the
family Leptodactylidae. Character state B = acrocentric chro-
mosomes absent and is derived.
A phylogeny based on these two characters is presented
(Fig. 3). Certain comparisons of this phylogeny with previ-
ously proposed species groupings (Heyer, 1968) are of interest.
As only two characters are used, the clusterings of species at
certain steps of the phylogeny are large, particularly for the
last. The most advanced cluster contains all of the members
of the ocellatus (L. bolivianus, chaquensis, ocellatus) and
pentadactylus (knudseni, pentadactylus, rhodonotus) species
332 Proceedings of the Biological Society of Washington
groups examined. All species examined of the fuscus group (L.
albilabris, bufonius, fuscus, geminus, gracilis, labialis, mysta-
ceus, mystacinus) except L. latinasus are in this group, and
one member of the melanonotus group (L. melanonotus) is
also represented in the most advanced group. The rest of the
melanonotus group members ( “natalensis,” podicipinus, wag-
neri) are in an ancestral grouping. This generally corresponds
to the scheme previously proposed (Heyer, 1969), in which
the melanonotus species group is among the most primitive
species groups in the genus. The two exceptions to this agree-
ment of phylogenetic schemes are L. melanonotus and L. latin-
asus. In the case of L. melanonotus, the advanced state of the
karyotype may indicate that L. melanonotus has diverged from
podicipinus and wagneri to a greater degree than expected on
the basis of standard morphological analysis. This explanation
makes biogeographic sense as L. melanonotus has a Middle
American distribution pattern, while podicipinus and wagneri
are South American species.
We think that the karyotype evidence gives greater insight
to the relationships among the species of the melanonotus
group, but that the karyotypes should not be emphasized more
than standard morphological evidence. For example, L. me-
lanonotus should not be removed from its group members
podicipinus and wagneri because it has a more derived karyo-
type. We are also hesitant in recognizing species within the
melanonotus group based only on karyotypic evidence. Bogart
(In press) and Denaro (1972) found different numbers of
acrocentric chromosome pairs in the two geographic samples
of L. podicipinus they examined. We prefer to give a con-
servative explanation to this karyotypic variation and consider
the two karyotypes to represent the same species. Similarly,
until more evidence is gathered, we prefer to consider the
karyotype Bogart (In press) reported as “natalensis” to be a
geographic variant of wagneri, differing in acrocentric number.
Because L. podicipinus and wagneri show geographic differ-
ences in the number of acrocentrics, both species would appear
to be ideal systems for detailed population study by the use
of karyotypes to determine evolutionary patterns within the
Karyotypes in Leptodactylus-complex 333
species, as has been successfully used in some species of small
mammals (e.g. Patton, 1972) and Sceloporus (e.g. Cole, 1972).
Members of the genus Adenomera apparently have the
most primitive karyotypes of the Leptodactylus-complex. The
karyotype of Vanzolinius is comparable to the more primitive
karyotypes of Leptodactylus species. Karyotypic information
is limited in delineating relationships at the generic level
within the Leptodactylus-complex.
Bogart (In press), using chromosome morphology based
upon centromere position, postulated relationships that crossed
many of the species groups lines that were based on other
morphological evidence (Heyer, 1968). We think that his
analysis is overextended as he is basing homologies on mea-
surements. As Pathak, Hsu, Shirley, and Helm (1973) demon-
strated, homologies based on karyotype measurements can be
incorrect (in this case banding indicated the errors based on
measurements). Atchley (1972) has also demonstrated the
pitfalls of predicting relationships from karyotypes when the
homologues are not known with certainty. The conservative
use of karyotype information as used herein is more consistent
with relationships of the taxa based on other data sets.
Bogart (In press) postulated that because Adenomera had
a primitive karyotype, terrestriality (no free swimming, feeding
larva) is primitive within the Leptodactylus-complex. We
disagree with this hypothesis for two reasons. First, the
homologous chromosomes are not known. It may be that the
karyotype of members of the genus Adenomera is actually
derived. Second, there is no reason why a group of frogs can-
not retain a primitive karyotype while having a derived life
history pattern. The morphological evidence (Heyer, 1969,
1972, 1974) is quite conclusive that terrestriality is a derived
condition in the Leptodactylus-complex; we see no reason to
overthrow this evidence in light of the supposedly primitive
karyotypes of Adenomera species.
We conclude that until homologous pairs of chromosomes
can be determined accurately (banding probably will give
this information), 1) the information content of karyotypes in
elucidating relationships among Leptodactylus species and
genera of the Leptodactylus-complex is limited, and 2) the
334 Proceedings of the Biological Society of Washington
karyotype evidence must be interpreted within the framework
of other morphological and ecological evidence and can not
profitably be used as an entirely independent set of informa-
tion in elucidating relationships.
ACKNOWLEDGMENTS
We thank Keith A. Berven, University of Maryland, for
assistance in the field in Ecuador and Argentina and for doing
much of the initial scanning of the microscope slides. The
Summer Institute of Linguistics allowed the use of their base
camp at Limoncocha for a field site. Dr. and Mrs. Raymond
F. Laurent and Miriam Heyer aided the field work in Ar-
gentina. Roy W. McDiarmid, University of South Florida,
kindly provided the material examined from the Costa Rican
specimens. James P. Bogart, Louisiana Tech University,
greatly aided our efforts by supplying us his data prior to
publication. Robert L. Bezy, Natural History Museum, Los
Angeles, made several suggestions on an earlier draft and
took the photographs for Figs. 1 and 2. Support from National
Science Foundation Grant GB 27280 administered by Pacific
Lutheran University is gratefully acknowledged.
LITERATURE CITED
ATCHLEY, W. R. 1972. The chromosome karyotype in estimation of
lineage relationships. Syst. Zool. 21:199-209.
Barsieri, F. D. 1950. Observaciones sobre los cromosomas y esper-
matozoides en algunos batracios del genero Leptodactylus.
Acta Zool. Lilloana 9:455—462.
Barrio, A. 1973. Leptodactylus geminus, una nueva especie del grupo
Fuscus (Anura, Leptodactylidae). Physis 32(84):199-206.
Bocart, J. P. In press. A karyosystematic study of frogs in the genus
Leptodactylus (Anura: Leptodactylidae). Copeia.
Cote, C. J. 1972. Chromosome variation in North American fence
lizards (Genus Sceloporous; undulatus species group). Syst.
LOO, Plessis,
Denaro, L. 1972. Karotypes of Leptodactylidae anurans Jour.
Herpetol. 6:71-74.
Heyer, W. R. 1968. Biosystematic studies on the frog genus Lepto-
dactylus. Ph.D. Dissertation, Univ. Southern California, 234 p.
——. 1969. The adaptive ecology of the species groups of the
Karyotypes in Leptodactylus-complex 330
genus Leptodactylus (Amphibia, Leptodactylidae). Evolution
23:421—428.
——. 1972. The status of Leptodactylus pumilio Boulenger (Am-
phibia, Leptodactylidae) and the description of a new species
of Leptodactylus from Ecuador. Contrib. Sci., Los Angeles
County Mus. 231:1-8.
———. 1974. Relationships of the marmoratus species group (Am-
phibia, Leptodactylidae ) within the Subfamily Leptodactylinae.
Ibid. 253: 1—46.
Lyncu, J. D. 1971. Evolutionary relationships, osteology, and zoo-
geography of leptodactyloid frogs. Univ. Kansas Mus. Nat.
Hist. Misc. Publ. 53:1-238.
ICRECAKom Onl Ga Hsu, le iSHiREEY AND) J, DS Herne, Wik 1973:
Chromosome homology in the climbing rats, genus Tylomys
(Rodentia, Cricetidae). Chromosoma 42:215—228.
Patron, J. L. 1967. Chromosome studies of certain pocket mice, genus
Perognathus (Rodentia: Heteromyidae). Jour. Mammal. 48:
27-37.
——. 1972. A complex system of chromosomal variation in the
pocket mouse Perognathus baileyi Merriam. Chromosoma 36:
QAV—255.
336 Proceedings of the Biological Society of Washington
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Vol. 87, No. 30, pp. 337-344 22, October 1974
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
A NEW CUBAN SPECIES OF SPHAERODACTYLUS
(GEKKONIDAE) OF THE NIGROPUNCTATUS
COMPLEX
By ALBERT SCHWARTZ AND ORLANDO H. GARRIDO
Miami-Dade Community College, Miami, Florida 33167,
and Academia de Ciencias de Cuba, Instituto de Zoologia,
La Habana, Cuba
The nigropunctatus complex of the gekkonid genus Sphaero-
dactylus occurs in the Bahama Islands (S. nigropunctatus
Gray), Hispaniola (S. stejnegeri Cochran), and Cuba (S.
nigropunctatus, S. torrei Barbour, S. alayoi Grant, S. inter-
medius Barbour and Ramsden, and S. ruibali Grant). The last
revision of the group is that of Thomas and Schwartz (1966);
they, however, used the name “decoratus complex” for this
group of species. Later Thomas and Schwartz (in press), after
examination of the long-neglected holotype of S. nigropuncta-
tus, showed that this name, which has priority over S. de-
coratus Garman, should be applied to the species which
formerly was called S. decoratus and which occurs in both the
Bahamas and Cuba. Thomas and Schwartz (1966:25) also
suggested that perhaps S. cinereus Wagler should be included
within this complex, but neither they nor others have studied
that species in detail.
In Cuba, the center of Sphaerodactylus diversity is in the
eastern portion of the island, in Camagiiey and Oriente Prov-
inces. This is true not only for the genus but also for the
nigropunctatus complex. Of the five Cuban species of this
complex, all occur in Oriente, and ruibali, torrei, and alayoi
are restricted to that province. S. intermedius has a peculiarly
disjunct distribution, occurring in extreme southwestern Ori-
ente Province (vicinity of Cabo Cruz) and in northern Habana
30—Proc. Bion. Soc. Wasu., Vol. 87, 1974 (SB)
338 Proceedings of the Biological Society of Washington
and Matanzas Provinces far to the northwest. The systematic
status of S. alayoi remains in doubt; no further specimens of
this taxon have been collected since the Thomas and Schwartz
review. Those authors commented that S. alayoi was a distinct
form, but whether it should be regarded as a separate species
or as a subspecies of S. nigropunctatus would depend in part
upon learning details of distribution of both forms in Oriente.
Gradually, additional material from the southern Oriente
coast has reached collections in the Instituto de Zoologia,
Academia de Ciencias de Cuba. Previously, S. ruibali has been
known only from the United States Naval Base, both east and
west of Bahia de Guantanamo. More recent specimens taken
by the junior author and Jorge de la Cruz have extended the
range of this species to the east as far as Baitiquiri (two speci-
mens) and Loma de Macambo between Imias and San Antonio
del Sur (one specimen). These newly collected specimens
agree in most details with long series of topotypes from the
east side of Bahia de Guantanamo on the United States Naval
Base; but specimens, taken primarily by Peter J. Tolson, from
the west side of Bahia de Guantanamo differ in several re-
spects from topotypical material and likely represent a new
subspecies of S. ruibali.
A series of six specimens of Sphaerodactylus was taken by
Luis de Armas in 1973 at Cabo Maisi, Cuba’s extreme eastern
point. This locality is about 60 kilometers northeast of the
nearest locality for any other member of the nigropunctatus
complex (ruibali, at Loma de Macambo). At first, we con-
sidered that this series of geckos was a well marked subspecies
of S. ruibali. Further comparisons, however, suggest that they
are more closely related to S. intermedius but are separated
from the nearest locality for that species (Rio Puerco) not
only by about 330 kilometers but also by the intervening dis-
tributions of S. torrei, S. alayoi, S. nigropunctatus, and S. rui-
bali. Although we are reluctant to add still another named
population to this complex of geckos on the southern Oriente
coast, the Cabo Maisi specimens are so distinctive that we
cannot associate them nomenclaturally with any other named
species. To complicate matters even further, the Oriente speci-
mens of S. intermedius number only two and are old and faded;
A New Cuban Gecko 339
additionally, they may not be identical with the Habana-
Matanzas segment of the species (the type-locality of S. inter-
medius). Since much of this xeric coastal area of Oriente is
still difficult of access, remote from most populous areas, and
collecting there is difficult, we feel justified in naming the
Cabo Maisi series as a distinct species, despite the presently
insoluble uncertainties involved with Sphaerodactylus from
this entire area. Accordingly, we name the Cabo Maisi lizards
in honor of Sr. de Armas who collected the type-series.
Sphaerodactylus armasi, new species
Holotype: Instituto de Zoologia (IZ), Academia de Ciencias de Cuba,
4089, an adult female, from Cabo Maisi, Baracoa, Oriente Province, Cuba,
taken 6 October 1973 by Luis de Armas.
Paratypes: YZ 4093-94, ASFS (Albert Schwartz Field Series)
V28442-43, and NMC (National Museum of Canada) 15849, same data
as holotype.
Definition: A moderate size species of Sphaerodactylus (males to
26 mm, females to 30 mm snout-vent length), characterized by the
combination of: a short blunt snout; dorsal scales small and granular,
45 to 60 between axilla and groin; ventral scales smooth and imbricate,
31 to 41 between axilla and groin; 74 to 89 scales around body at mid-
body; usually 4 enlarged supralabials to eye center; 1 internasal scale.
Head pattern in females diffuse grays and white without a distinct
pattern (even in female-patterned juveniles) except for a dark gray to
black postocular line and a dark loreal line which in fully adult females
may expand to give a solid dark loreal region; dorsum strongly cross-
banded pale and dark, with five pale (whitish in preserved specimens )
crossbands, at times composed of transverse series of isolated pale elon-
gate dots, bordered by black bands on either side, the most anterior
black bands and their included pale band forming a collar immediately
posterior to the ear opening; a pair of large pale ocelli on the base of
the tail, followed by two additional tail bands of two fused ocelli (no
tail is complete). Males dull grayish brown dorsally, more or less
vaguely spotted with dark gray, the upper surface of the head at times
with scattered black irregular spots, extending onto the lores and snout,
and. onto the sides of the throat.
Variation: The series of S. armasi consists of three adult females,
two adult males, and one juvenile. The largest male (ASFS V28443)
has a snout-vent length of 26 mm, the largest female (ASFS V28442)
30 mm, and the juvenile 20 mm. Dorsals between axilla and groin are
45 to 60, ventrals between axilla and groin 31 to 41, scales around body
at midbody 74 to 89, fourth toe lamellae 7 to 11, almost always 4 en-
larged supralabials to eye center (one specimen with 3 enlarged supra-
340 Proceedings of the Biological Society of Washington
labials bilaterally), internasal scale usually 1 (one specimen with 2),
escutcheon in two males 3 to 10 < 6 to 15. All ventral scales smooth,
including gular and pectoral scales; dorsal scales small and granular,
showing very slight keeling dorsolaterally.
The two males are generally undistinguished. The dorsal color (as
preserved) is dark tan to grayish, with vague to moderately prominent
dark blotches densely appressed and almost lepoard-like. One male
(IZ 4093) has the upper surface of the head heavily marked with black
spots, these extending laterally and ventrally onto the sides of the throat,
whereas the other male has the head and throat unmarked. The heavily
spotted male also has a tiny pair of pale ocelli, outlined with dark gray,
on the neck, and both males have a pair of almost-fused postsacral ocelli,
the tails showing vague further indications of more distal transverse
bands.
The three females and the female-patterned juvenile are very dis-
tinctive. Basically, the pattern consists of a series of five pale transverse
bands, at times showing their origin from a series of elongate (anterior
to posterior) pale spots which fuse with each other laterally to give a
continuous pale band. Each pale band is bordered both anteriorly and
posteriorly by dark gray to black edging, which may be irregular or
scalloped due to the nature of the pale crossbands (i.e., that they repre-
sent fused rows of pale spots). Of the five pale bands, one is nuchal,
one is more or less scapular, and three are on the body. The anterior
dark border of the first pale band lies just behind the auricular opening.
There is a prominent pair of large pale, but dark-bordered, ocelli post-
sacrally, and the unregenerated portions of the tails likewise have at
least two prominent irregular pale crossbands which represent pairs of
fuse docelli. The juvenile is like the females in dorsal pattern, but the
pale transverse body bands are less prominent than in adults. The head
in females is mottled with shades of gray and white dorsally and has
no clear-cut pattern except for the presence of a dark postocular line,
and in the juvenile a dark loreal line. The latter line, in adult females,
is so expanded that the entire loreal region is black. The holotype has
some large pale areas on the temporal region, and the dark postocular
line is much obscured. The venter (including the chin and throat) are
immaculate, but there may be some uniform dark stippling (ASFS
V28442) on the throat. .
Comparison: S. armasi requires comparsion primarily with S. ruibali,
that species of the complex which is closest geographically. In S. ruibali,
dorsals between axilla and groin are 42 to 60, ventrals between axilla
and groin 30 to 43, scales around midbody 69 to 80, fourth toe lamellae
6 to 12, supralabials to eye center usually 4 (occasionally 3 or 5), 1
internasal, and escutcheon 5 to 10 * 19 to 28. None of these counts
is strikingly different between the two species; however, S. armasi has
a higher number of scales around the body at midbody (74 to 89) than
does S. ruibali (69 to 80); there is, however, considerable overlap.
A New Cuban Gecko 341
It is in pattern that S. armasi differs most strongly from topotypical
and eastern S. ruibali. In the S. ruibali populations east of Bahia de
Guantanamo, males are finely stippled dorsally and lack any sort of
head pattern; there is no tendency for these males to have spotting on
the body. In females from the same region, there is a nuchal pattern of
three fine dark transverse bands not including a pale ocellar band, and
occasionally there is a fourth dark band across the back at the axilla.
The female head pattern in S. ruibali consists of a clearly delineated
dark canthal line which continues as a distinct postocular line; the loreal
region is never generally black. The top of the head is vaguely patterned
with pale dots or merely variegated smudges of grays. Both sexes of
S. ruibali are shown in Thomas and Schwartz, 1966, fig. 19. In addi-
tion, S. ruibali lack ocellate tails, although there may be (rarely) a pair
of faint ocelli in the sacral region in males.
Referring to Thomas and Schwartz, 1966, fig. 18, which shows two
female S. intermedius from northern Habana and Matanzas Provinces,
one can see the similarities between that species and S. armasi. The
dorsal pattern in the Habana specimen shows a diagrammatically trans-
versely banded lizard, with five pale crossbands (which do not show
indications of a fused ocellar origin) and with distinct and sharp-edged
black borders; there is a postsacral pair of ocelli. The Matanzas specimen
illustrated shows the same sort of pattern but only the axillary pale band
is clearly shown. However, the head pattern of S. intermedius is sharply
delineated, and consists mainly of a complete or incomplete black
U-shaped figure, along with additional dark cephalic lines including a
loreal and a postocular line. As far as scale counts are concerned, S.
intermedius has 49 to 62 dorsals between axilla and groin, 31 to 44
ventrals between axilla and groin, 65 to 77 scales around midbody,
internasals usually 1 (0 to 2), and enlarged supralabials 4 (mode) or
5. S. armasi is comparable in scale counts to S. intermedius, but the
midbody counts are higher in S. armasi (74 to 89) than in S. intermedius
(65 to 77) with less overlap than between S. armasi and S. ruibali.
In addition, S. intermedius reaches a greater snout-vent length (35 mm)
than does S. armasi. It should be recalled that the major portion of
the known distribution of S. intermedius lies far to the northwest along
the northern coast of Habana and Matanzas Provinces, but that there
are two old and faded specimens from the Cabo Cruz region which
Thomas and Schwartz assigned to S. intermedius.
The three other species of the nigropunctatus complex which occur
in southern Oriente Province are S. nigropunctatus, S. alayoi, and S.
torrei. Females of all these species are vividly crossbanded and also
have distinct dark head patterns, a feature which is absent in S. armasi.
All species are quite comparable in scale counts; for instance, all (in-
cluding S. armasi) regularly have 1 internasal scale and 4 enlarged
supralabials to eye center as the modal conditions. Dorsal scales are
less in S. ¢. torrei (45 to 49), the subspecies closest geographically to
342 Proceedings of the Biological Society of Washington
S. armasi, and these two species are distinguishable by this feature as
well as in details of pattern. S. alayoi (whose status as a distinct species
remains questionable) differs from S. armasi in its vividly marked cross-
banded dorsum (three dark but uniformly hollowed crossbands), head
pattern, and slightly larger size (32 mm in S. alayoi). From S. n. strate-
gus (the subspecies of S. nigropunctatus closest geographically to S.
armasi), S. armasi differes in smaller size (S. n. strategus to 40 mm
snout-vent length), quite different dorsal and head patterns in females,
and lesser number of dorsal scales between axilla and groin (45 to 60
versus 50 to 67). In addition, none of the species with which S. armasi
might be confused locally is distinctly short-snouted; this is especially
true of S. ruibali, which is a long-headed and attenuate-snouted species.
Remarks: Very much yet remains to be learned concerning the
distributions and variation of the southern Oriente Sphaerodactylus.
In those species about which we have some ecological information,
S. torrei and S. nigropunctatus are mesophilic or at least inhabitants
of shaded situations. S. ruibali on the other hand inhabits open and
more xeric areas. S. intermedius (along its northern Habana-Matanzas
distribution) likewise is mesophilic and most closely associated with
rock rubble; it appears to some extent to be saxicolous. The series of
S. armasi was secured in xeric situations associated with a species of
Agave. Also secured syntopically with S. armasi was S. notatus atactus
Schwartz; this species is tolerant of a wide variety of ecological situa-
tions, from semi-mesic coastal woods to open and exposed beaches.
S. notatus is widespread in Cuba (where it is less common in the western
and central portions of the island), Isla de Pinos, throughout many of
the Bahama Islands, and the Swan Islands; it also occurs on the con-
tinental mainland in extreme southeastern Florida and the Florida Keys.
Its broad geographical distribution is due at least in part to its ecologcial
tolerances in contrast to other species of Sphaerodactylus which are
often more rigidly restricted ecologically.
One further comment is pertinent. Specimens of S. ruibali from west
of Bahia de Guantanamo (an area where the species has not previously
been recorded) show strong differences in dorsal and cephalic pattern
from specimens taken east of the bay. It is likely that these S. ruibali
are subspecifically different from their eastern relatives. However, they
do not approach S. armasi in complexity of dorsal pattern and are typ-
ically long-snouted S. ruibali. They show no trends toward the very few
Cabo Cruz S. intermedius, and indeed the hiatus between the two species
in this area is about 220 kilometers. Thomas and Schwartz (1966:24)
suggested that S. ruibali and S. intermedius are related (S. armasi seems
to be close to this species-pair), but there is as yet no unequivocal evi-
dence that S. ruibali should be regarded as a subspecies of S. intermedius.
Certainly, the southern Oriente coast of Cuba and its adjacent lower
montane slopes well merit intensive collecting for the nigropunctatus
complex of Sphaerodactylus.
A New Cuban Gecko 343
LITERATURE CITED
Tuomas, R., AND A. ScHwartTz. 1966. The Sphaerodactylus decoratus
complex in the West Indies. Brigham Young Univ. Sci. Bull.
7(4):1-26, 20 figs.
—. (Im press). The status of Sphaerodactylus gilvitorques Cope
and Sphaerodactylus nigropunctatus Gray (Sauria:Gekkonidae ).
Jour. Herpetol.
344 Proceedings of the Biological Society of Washington
Vol. 87, No. 31, pp. 345-350 22, October 1974
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
A NEW POLYDESMID MILLIPED FROM THE
SOUTHERN APPALACHIANS, WITH REMARKS ON
THE STATUS OF DIXIDESMUS AND A PROPOSED
TERMINOLOGY FOR POLYDESMID GONOPODS
By Ricuarp L. HorrmMan
Radford College, Radford, Virginia 24141
The diplopod generic name Dixidesmus, presently compre-
hending some eleven named taxa endemic to eastern United
States, was established by R. V. Chamberlin in 1943. The
genus was defined as being very similar to Pseudopolydesmus
in gonopod structure, “. . . but differing in having a spine or
spine-like process in the angle above the papillose cushion and
in having on outer side, at or distad of the level of the cushion,
a retrorse tooth or barb with a spur on or near its base, the
latter rarely abortive or absent.” Dixidesmus has subsequently
come into general use by American diplopodologists despite
its foundation upon a single character (the second cited by
Chamberlin admitted to be variable).
Since 1951 I have had material of an undescribed species
which completely negates the supposed difference between
the two “genera” mentioned and, in my view, requires the
abolition of Dixidesmus. Publication of this species has been
repeatedly deferred in the hope it might be included in a re-
vision of the American forms, but as the completion of that
revision still seems remote, I wish to place the new taxon on
record in order to justify combination of Dixidesmus with
Pseudopolydesmus in a general classification of Diplopoda
now in preparation.
Genus Pseudopolydesmus Attems
Polydesmus (nec Latreille 1802) American authors prior to 1943.
Pseudopolydesmus Attens, 1898, Denk. Akad., Wien, 67: 270, 479.—
31—Proc. Biou. Soc. WAsH., VoL. 87, 1974 (345 )
346 Proceedings of the Biological Society of Washington
Brolemann, 1916, Ann. soc. ent. France, 84: 569.—Verhoeff, 1931,
Zool. Anz., 94: 305.—Attems, 1940, Tierreich, 70: 139.—Carl, 1941,
Zool. Anz. 133: 291.—Chamberlin, 1943, Bull. Univ. Utah, biol. ser.,
8(2): 17. Type species, Polydesmus canadensis Newport, 1844 (? =
serratus Say, 1821).
Dixidesmus Chamberlin, 1943, Bull. Univ. Utah, biol, ser., 8(2): 18.—
Chamberlin & Hoffman, 1958, U. S. Nat. Mus. Bull. 212:65. Type
species, Dixidesmus tallulanus Chamberlin, by original designation.
[New synonymy. ]
Diagnosis: Gonopod large, falcate in general shape, with prominent
endomerite subtended by small dentiform process (m 1) on median side;
prostatic groove running distad from median to lateral side, debouching
not into a small fossa within endomerite but into a deeply invaginated
groove (“Samenslauch,” Carl) which extends proximad to base of fem-
oral region, and is then reflexed distad to open on distad side of endo-
merite. Apex of telopodite with a linear series of flattened setae, and,
usually, a small cluster of spiniform setae from base of distalmost process
on the median side (m 4).
Species: Approximately ten, all endemic to North America east of
the Great Plains, concentrations of species in the southern Appalachians
and in the Ozark region.
Remarks: The combination of the names Dixidesmus and Pseudo-
polydesmus is suggested here because of the discovery of a species which
in some ways resembles “D.” branneri (e.g. fusion of lateral telopodite
processes e 2 and e 3) but which has the proximal lateral process (e 1)
reduced to a small point or in many specimens, absent entirely. As this
process was the criterion upon which Dixidesmus rested, and as the
examination of numerous species of American polydesmids has revealed
no other basis for making generic distinctions, there seems no justifica-
tion for continued recognition of Dixidesmus.
This merger will bring the following names into combination with
Pseudopolydesmus for the first time: branneri Bollman, 1887; catskillus
Chamberlin, 1947; christianus Chamberlin, 1946; echinogon Chamber-
lin, 1942; erasus Loomis, 1943; gausodicrorhachus Johnson, 1954;
humilidens Chamberlin, 1943; nitidus Bollman, 1887; penicillus Cham-
berlin, 1943; phanus Chamberlin, 1951; sylvicolens Chamberlin, 1943;
and tallulanus Chamberlin, 1943. The great majority of these names are
either subspecific in rank or are strict synonyms, and will be accounted
in a forthcoming paper. Bibliographic reference to all of these names
is to be found in the “Checklist of the Millipeds of North America”
(Chamberlin & Hoffman, 1958).
Pseudopolydesmus collinus, new species
Figures 1, 2
Diagnosis: A medium-sized species of the genus (length to 23 mm),
with the following diagnostic gonopod characters: process m 1, m 2, and
Remarks on Diplopods and a new species 347
m 4 present; e 1 vestigial, sometimes wanting; e 2 and e 3 basally fused
and stipate; e 4 present.
Type-material: Male holotype, ¢ and 9 topoparatypes from “Pin-
nacles of Dan,” + 4 miles southwest of Vesta, Patrick County, Virginia;
22 April 1972, R. L. Hoffman & L. S. Knight, leg. (Hoffman collection ).
Holotype: Adult male, 22.0 mm in length, 3.8 mm in maximum
width, W/L ratio 17.3%. Segmental widths across paranota as follows:
Segment 1-2.3 mm Segment 10-3.7 mm
2-2.6 12-3.5
4—3,2, 14-3.4
6-3.8 16-3.2
8-3.8 18-2.0
Color in life dorsally uniform light brown, legs and sterna lighter, no
suffusion of reddish pigmentation on paranota.
Segmental sculpture typical for the family. Most metaterga with three
transverse series of convex areas middorsally: a large area occupying the
anterior half of the tergum between stricture and transverse sulcus, four
smaller pentagonal areas immediately behind the sulcus, and six smaller
areas along the posterior edge of the segment. On the posteriormost
segments these latter areas project slightly caudad, producing a subserrate
edge between the paranota. Latter moderately broad, the anterior and
lateral edges forming a continuous curvature with four minute lateral
denticles; peritreme broad, convex, the pore small and located in a
dorsolateral depression behind middle of peritrematic length. Dorsum
of paranota with an almost circular prominent convexity.
Sterna bicruciately impressed, each quadrant moderately setose; sterna
between legs 3, 4, 5, 6, 9, 11, and 13 produced into two prominent
conical paramedian lobes, the lobes anterior to gonopods profusely
setose; sterna between legs 7 and 10 produced into subhemispherical
subcoxal swellings a little further apart than the other lobes. Prefemora
and femora of most legs somewhat enlarged, ventral sides of tibiae and
tarsi with a row of moderate-sized sphaerotrichomes, preceded on the
anterior side by a series of much smaller but similar setae.
Coxae of gonopods large, mostly exserted, directed caudal parallel to
each other and to median axis of body, with the usual concavity on the
ventral side for retraction of telopodites, lateral ventral edge with two
macrosetae. Telopodite of normal polydesmid form, arcuate caudally,
the prefemur densely setose but without processes, acropodite slender,
with a terminal series of macrosetae and endomerite at midlength, medial
side with three marginal processes (m 3 apparently absent) and lateral
with four (e 1 vestigial or absent, e 2 and e 3 fused and carried on a
common stalk, e 4 of normal form and position, see Figs 1 and 2).
Remarks: An account of female genitalia is deferred for the planned
revision of the genus. Externally the female is similar to the male in
size, color, and peripheral characteristics.
348 Proceedings of the Biological Society of Washington
Fics. 1-2. Pseudopolydesmus collinus, n. sp.: 1, left gonopod of male
topoparatype, mesal aspect; 2, the same gonopod, lateral aspect.
Fic. 3. Pseudopolydesmus branneri (Bollman): left gonopod of speci-
men from Roan Mountain, Tennessee, slightly oblique caudo-mesal aspect,
to show complete series of telopodite processes.
Remarks on Diplopods and a new species 349
This species has been taken in association with both Pseudopolydesmus
serratus and P. branneri in western Virginia, as well as in biotopes in
which those two species were absent. P. collinus is easy to recognize
in the field, being about 3-5 mm shorter than serratus which it otherwise
resembles, and 2-4 mm smaller than branneri which is characterized by
having a black middorsum and reddish paranota. No observations sug-
gest any difference in the phenology of the three, nor, overtly, in habitat
preferences.
The specific name reflects the fact that although the species occurs
in the southern Appalachians it clearly prefers moderate elevations and
has not been found so far above 3600 ft.
Distribution: The species is known to me from the following counties
(detailed citations deferred for a later account): Kentucky: Carter Co.;
West Virginia: McDowell Co.; Virginia: Alleghany, Floyd, Franklin,
Henry, Montgomery, Patrick, and Pittsylvania counties. It seems to be
most abundant in central southwestern Virginia on and just east of the
Blue Ridge, but occurs on river bluffs well out into the Piedmont, and
extends westward to easternmost Kentucky.
Gonopop TERMINOLOGY
The characterization of polydesmid gonopods has long been impeded
by the lack of standardized names for the various spines, branches, or
processes which adorn the posterior face of the telopodite. Although
some names have been used for European species, they are usually
vernacular in origin and often not easily translatable, in any case often
long and cumbersome.
It has long seemed possible to me to devise a somewhat different
approach which I venture to present here in a brief outline. First of
all, the invention of meaningful names of classical origin is not only
difficult but perhaps at the present time a little premature, so that one
alternative is to develop a numerical system of designation reflecting
the actual location of the processes as well as providing an indication of
serial homologies between species.
The telopodite of the gonopod in polydesmids, as seen in a caudal or
obliquely caudal aspect (Fig. 3) is fairly broad at its base and gradually
tapers to a much thinner distal end, although even there it retains the
dimension of breadth, and may be correctly stated to have two distinct
sides: ectal and mesal. Except for the large setiferous endomerite, which
occupies a median position on the gonopod, the various smaller processes
all originate on one side or the other and this fact provides an initial
dichotomy in their classification: a given process is either mesal or
ectal in its place on the telopodite and may be indicated by the prefix
m or é.
Arbitrarily taking as a standard that species which is most complex
in terms of number of processes, such as P. branneri, we can apply a
numerical designation to those on each side starting with the most
300 Proceedings of the Biological Society of Washington
proximal. On each side there appear to be no more than four processes,
although in some forms one or more may be suppressed, or two may be
coalesced and carried by a common pedicle. Thus, in P. branneri, the
processes on the mesal side are m 1, m 2, m 3, and m 4, on the ectal
side e 1, e 2+-3, and e 4, by which it is indicated that 2 and 3 are joined
at the base. In the American species, process m I is generally present
near the base of the endomerite, and e 4 is usually present beside the
terminal row of setae. M 4 usually carries several setae at its base and
is easy to recognize as a result. The greatest variation thus appears to
occur with processes 2 and 3 on each side.
Using the above terminology, the original generic diagnosis for
Dixidesmus quoted above could be rewritten as follows: “Gonopod with
process e I present, spiniform or acicular; process e 2 present, usually
with a smaller e 3 at its base.”
It is generally necessary to examine a gonopod from both sides, as
frequently it is impossible to see all of the processes due to concealment
of a small process by juxtaposition of a larger one; this is shown very
clearly in Figs. 1 and 2.
LITERATURE CITED
CHAMBERLIN, R. V. 1943. On some genera and species of American
millipeds. Bull. Univ. Utah, biol. ser., 8(2):1—20, figs. 1-42.
, AND R. L. Horrman. 1958. Checklist of the millipeds of
North America. U.S. Nat. Mus. Bull. 212:i-iii, 1-236.
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Vol. 87, No. 32, pp. 351-380 22. October 1974
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
PELAGIC OSTRACODS COLLECTED ON HUDSON 70
BETWEEN THE EQUATOR AND 55°S
IN THE ATLANTIC
By GEorGIANA B. DEEVEY
Florida State Museum, University of Florida,
Gainesville, Florida 32611
Zooplankton samples were collected in the South Atlantic
on the first two legs of the CSS HUDSON 70 Expedition.
Between the equator and 55°S, samples were obtained rou-
tinely along the 30°W Longitude parallel (Fig. 1). The first
leg of the expedition left Halifax in mid-November 1969 and
arrived at Rio de Janeiro in mid-December. The second leg
left Rio on December 20th, proceeded down the 30°W Longi-
tude parallel to 55°S, then passed northeast of South Georgia
and up the 40°W Longitude parallel to 47°S, crossed the
Argentine Basin to the Rio Plata and arrived in Buenos Aires
in mid-January, 1970. Between the equator and 25°S, zoo-
plankton samples were collected for the micropaleontology
program by vertical tows, 200-0 m, with a % meter net of No.
8 mesh. South of 27°30’S, hauls were made with % m nets of
No. 6 mesh to varying depths with up to 4,000 m of cable out,
and the 200-0 m vertical tows were also obtained.
Some 70 species of pelagic ostracods were recorded, of
which two belong to the family Cypridinidae and the rest to
the family Halocyprididae. Since many species of ostracods
are bathypelagic, only 33 species were found in the 200-0 m
tows collected between the equator and 25°S. Eight species
(Conchoecia acuminata, C. allotherium, C. atlantica, C. con-
centrica, C. echinata, C. inermis, C. incisa and C. stigmatica)
were not noted south of 16°S. Six more species (C. ctenophora,
C. echinulata, C. microprocera, C. macrocheira, C. parthenoda
32——Proc. Brou. Soc. WasH., Vou. 87, 1974 (351)
352 Proceedings of the Biological Society of Washington
:
is
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Tee South
60° 50° seen 30° 20°W
Fic. 1. Map of zooplankton station locations for Legs 1 and 2 of
HUDSON 70.
and C. pseudoparthenoda) did not occur south of 30°S. Be-
tween 30°S and 40°S, 42 species were recorded, half of which
were not noted south of the subtropical convergence. Between
the subtropical and subantartic convergences, around 42°S-
49°S, 27 species were present in the samples, although 9 of
these were not found south of 45°S. South of the subantarctic
convergence 18 species occurred between 50°S and 55°S. Two
Pelagic Ostracods from South Atlantic
TABLE 1.
vertical tows.
303
Station data, Leg 1, Halifax to Rio de Janeiro. 200-0 m
Station Date Latitude Longitude Time of day No. species
1 XI1/30/69 0°02’N 29°58.3’W 10.40 AM 12
2 XI/30 0°51’S 30°W 9.00 PM 21
3 XII/1 2°29’S 30°W 10.20 AM 18
4 XII/1 3°32’S 30°07’W 10.00 PM 16
5 XII/2 4°55’S 30°W 10.20 AM 16
6 XII/2 6°04’S 30°W 10.15 PM 18
7 XIT/3 COAUS 30°03’W 10.20 AM 14
8 XIT/3 8°30.8’S 30°W 10.20 PM 14
9 XII/4 9°59’S 30°W 10.30 AM 11
10 XII/4 10°59’S 29°59’ W 10.00 PM 16
11 XII/5 12°26’S 30°07’W 10.00 AM 10
12 XII/5 13°29’S 30°W 10.30 PM 16
13 XII/6 14°55’S 30°07’W 10.30 AM o
14 XII/6 15°47’S 29°59’ W 10.15 PM 13
15 XII/7 17°32’S 30°W 1.30 PM 8
16 XII/7 18°19.5’S 30°04’W 10.50 PM 11
17 XII/8 20°S 30°W 8.15 AM 10
18 XIT/8 20°02.6’S 30°31.3’W Midnight 17
19 XII/9 22,°30.2’S 29°58’ W 2.15 PM 9
20 XII/9 23°21’S 29°55’W 9.45 PM 19
21 XII/10 25°S 30°W 8.30 AM 14
22, XII/10 24°52.5’S 30°08’ W 8.15 PM 15
23 XII/11 24°37’S 32°30.6’W 8.15 AM 12
24 XIT/11 24°17.3’S 32°44’W 9.00 PM 16
25 XII/12 24°13.4’S 35°W 7.15 AM 15
species, C. isocheira and C. belgicae, were found only south
of the subantarctic convergence. The total number of species
recorded therefore decreased markedly between the sub-
tropical and subantarctic convergences. HUDSON 70 speci-
mens extend the southward ranges in the Atlantic for 27
species, some by only a few degrees. The northward range
was extended for two species, C. belgicae and C. major (= C.
plactolycos major). C. plactolycos (= C. plactolycos placto-
lycos), described from the Indian Ocean, is now known from
the Atlantic. A few species, such as C. elegans and C. skogs-
bergi, increased in size farther south. Larger forms began
354 Proceedings of the Biological Society of Washington
TABLE 2. Station data, Leg 2, Rio de Janeiro to Buenos Aires.
Total M No.
Station Date Latitude Longitude of cable species
12 XII/23/69 27°31.6’S 30°07’'W 450 18
13 XII/24 30°00’S 29°59’W 1400 32
14 XII/25 30°25.05’S 30°10.5’W 1500 30
15 XII/26 35°00’S 30°04.7’'W 1800 36
16 XII/27 37°28’S 29°59.2’W 2500 30
17 XII/28 39°55’S 29°53’W 2500 30
18 XITI/29 42°30’S 29°57.5’W 2500 24
19 XII/30 45°07.3’S 29°59’W 2500 15
20 XII/31 47°35’S 29°53.4’W 2500 9
21 1/1/70 49°03.4’S 29°48.7’W 2500 14
22 1/2 50°33.3’S 29°35.4’W 2500 12
23 1/3 52°04’S 30°04’W 2500 t
24 1/4 53°21 29°59.5’W 2500 6
25 1/5 54°55’S 29°23’W 2500 11
26 1/8 51°28’S 39°57’W 3000 14
TT 1/8 49°59’S 39°58’W 1000 9
28 1/9 48°59’S 39°52.4’W 500 6
29 1/10 47°59’S 39°59.6’W 400 7
30 1/10 47°03’S 40°04.6’W 3000 13
30A Y/1l 45°10.6’S 41°10.8’W 4000 16
30B 1/12 44°12,.4’S 42°46’W 4000 25
31 1/13 41°46.5’S 46°26.4’W 2500 31
to appear in the region of the subtropical convergence, but
the increase in size was gradual; two distinct size ranges for
a species were not noted.
The station data for Legs 1 and 2 are given in Tables 1 and
2. Table 2 shows the decrease in numbers of species/station
south of the subtropical convergence, and the increase in
numbers of species as the ship proceeded north over the
Argentine Basin. Table 3 lists the species of ostracods, to-
gether with the stations from which they were recorded and
their overall latitudinal distribution. All specimens have been
deposited in the Florida State Museum.
I am indebted to Gordon Riley, Director of the Institute of
Oceanography at Dalhousie University, for arranging for my
travel between Halifax and South America, to C. R. Mann
300
Pelagic Ostracods from South Atlantic
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358 Proceedings of the Biological Society of Washington
and R. J. Conover of the Bedford Institute of Oceanography
for inviting me to participate on Leg 2 of HUDSON 70, and
to G. Vilks for permission to examine the ostracods in the
200-0 m vertical tows. This work was partially supported by
grants GB-15575 and GA-36512 from the National Science
Foundation.
THE SPECIES OF OSTRACODS
Suborder CYPRIDINIFORMES Skogsberg
Family CyprminmaAE Dana
Gigantocypris muelleri Skogsberg. One gravid female, 19.9 mm long,
and 7 juvenile specimens were taken in hauls with 2,500 m of cable out,
between 37°30’S and 53°30’S.
Distribution: 63°38’N-62°S in the Atlantic.
Macrocypridina castanea (Brady) s. str. One female 5.9 mm long
and a female 6.2 mm long were found at 30°S and 35°S. Poulsen (1962)
designated the Indo-Pacific form as M. castanea rotunda.
Distribution: 56°N—35°S in the Atlantic.
Suborder HALOCYPRIFORMES Skogsberg
Family HALocypripIpDAE Dana
Subfamily ARCHICONCHOECINAE Poulsen
Archiconchoecia cucullata (Brady). A number of specimens were
taken in tows with 750-4000 m of cable out, between 30°S and 50°30’S.
Distribution; 52°N-50°30’S in the Atlantic; Indian Ocean. These
specimens extend the southward range.
A. striata Miller. This tiny species was common in the 200-0 m hauls
taken between the equator and 25°S and was present to 37°30’S, 30°W
and at about 42°S in the Argentine Basin; it did not occur south of
the subtropical convergence.
Distribution: 36°N-—42°S in Atlantic; Indian and Pacific Oceans and
Mediterranean. These specimens extend the southward range.
Subfamily HaLocyprinAE Poulsen
Halocypria globosa Claus. H. globosa occurred from 20°S—42°S,
30°W, and in the Argentine Basin between 42°S and 45°S. Females
were caught at shallow depths, but males (Fig. 2h) were taken only
in samples collected with 750-2500 m of cable out.
Distribution: 64°N-37°S in the Atlantic; Pacific and Indian Oceans.
These specimens extend the range south to 45°S.
Halocypris brevirostris (Dana). This species was present in all 200-0
m tows obtained between the equator and 25°S, and was found down to
49°S, extending the southward range in the Atlantic. Between the
Pelagic Ostracods from South Atlantic 309
equator and 32°30’S, adult specimens were 1.15-1.25 mm long. Females
up to 1.45 mm long appeared at 35°S; between 37°30’S and 42°30’S
females were 1.5-1.6 mm long.
Distribution: 60°N—49°S in the Atlantic; Pacific and Indian Oceans.
Fellia cornuta dispar (Miiller). Muller (1906a) described two sub-
species of F. cornuta, the typical form, F. cornuta typica (1906a) = F.
cornuta cornuta (1912), which has a sharp horn on each side of the
shoulder vaults (Fig. 2a), and F. cornuta dispar for some southern
hemisphere males which lacked these projections and had rounded
shoulder vaults. Both Miiller and Poulsen (1969a) recorded only males
as lacking the sharp horns. Miiller’s specimens were found south of 29°S
in the Atlantic and Indian Oceans, and Poulsen’s south of 30°S in the
Pacific off New Zealand. The typical horned form has been recorded
from around 20°N-38°S in the Atlantic, around the equator in the Indian
Ocean and Indo-Pacific regions, and from around 30°N-—45°S in the
Pacific. Both males and females, lacking the sharp lateral processes,
were found between 30°S and 45°S, in hauls with 1100-2500 m of cable
out; no typical forms were taken. Juvenile specimens 1.30-1.45 mm long
(Fig. 2f) had small lateral points on the shoulder vaults, not as large and
curved as in the typical form (Fig. 2a). Specimens 2.0-2.1 mm long
had either no points or very tiny points (Fig. 2g). Mature males 2.9-3.1
mm long (Fig. 2d, e) and females 3.00-3.35 mm long (Fig. 2b, c) had
completely rounded shoulder vaults. This subspecies can therefore be
distinguished from the typical form in the juvenile stages.
Distribution: 29°S—45°S in the Atlantic; south of 29°S in Pacific
and Indian Oceans.
Subfamily CONCHOECINAE Poulsen
Most Halocyprids belong to the genus Conchoecia, for which over 90
species have been described. Miiller’s (1906a) system of grouping
species together into more or less natural groups of related forms will
be followed here.
Spinifera Group Miller
C. spinifera Miiller. This species was recorded from the equator to
40°S, 30°W and at about 42°S in the Argentine Basin, extending the
recorded southward range. It did not occur south of the subtropical
convergence.
Distribution: 52°N—42°S in the Atlantic; Indian and Pacific Oceans.
C. oblonga (Claus). Miiller (1906a) distinguished two forms of this
species, Form A in which the right asymmetrical gland opens at the
postero-ventral corner, and Form B in which the gland opens just an-
terior to this location. In the Canary Island region Angel (1969) found
Form A at shallower depths than Form B. The two forms were not dis-
tinguished in the HUDSON 70 material. C. oblonga was found from
the equator to 37°S and at about 42°S in the Argentine Basin, extending
its southward range.
360 Proceedings of the Biological Society of Washington
Fic. 2. a, Ventral view of female Fellia cornuta cornuta from the
Caribbean Sea; b and c, Anterior and dorsal views of female F. cornuta
dispar; d and e, Lateral and ventral views of male F. cornuta dispar;
f, Dorsal view of 1.3 mm juvenile; g, Ventral view of 2.0 mm juvenile;
h, Lateral view of Halocypria globosa male; i, Lateral view of Con-
choecia allotherium male; j, Endopodite of left second antenna of C.
allotherium male; k, Right clasper; 1, Capitulum of frontal organ;
m, Furca; n, Distal part of proximal secondary seta of male first antenna.
Scale on b for a—e and g, on h for f and h, on i for i, on right margin for
l and m, at bottom for j, k, and n. Scales in mm.
Pelagic Ostracods from South Atlantic 361
Distribution; 48°N—42°S in Atlantic; Indian and Pacific Oceans.
C. allotherium Miller. Males of this species were taken in two of the
200-0 m tows from 0°51’S and 15°42’S. This species is similar in size
and shape to C. oblonga Form A (Fig. 2i); also the 4th claw on the
furca is curved as in C. oblonga (Fig. 2m), and Miiller was unable to
differentiate the females of the two species. The male is distinguished
by a double row of spinules near the tip of the proximal secondary seta
(Fig. 2n) of the first antenna, and the frontal organ is bare of spinules
(Fig. 21).
Distribution: 18°N-35°S in Atlantic; 10°N-—10°S in Indian Ocean.
C. aequiseta Miiller and C. hirsuta Muller. Females of these two
species are indistinguishable, and males are differentiated only on the
basis of long hairs on the distal bristle of the first endopodite joint of
the second antenna in C. hirsuta. This bristle is bare in C. aequiseta
males and in females and juveniles of both species. Females and
juveniles identified as C. aequiseta were found between 30°S and 37°30'S,
30°W; a C. dequiseta male was taken at 42°S in the Argentine Basin.
C. hirsuta males were present in samples from 35°S, 30°W and from
42°S—44°S in the Argentine Basin, so both forms occur south to the
region of the subtropical convergence.
Distribution: For C. aequiseta 32°N-31°S and for C. hirsuta 29°N-
35°S in the Atlantic; Indian Ocean. The HUDSON 70 males extend
the southward range to 42°S and 44°S, respectively.
C. echinata Miller. This species occurred from the equator to 20°S.
Distribution: 37°N-29°S in Atlantic; Pacific and Indian Oceans.
C. inermis Miiller. A single specimen was taken at 13°30’S in a 200-0
m night haul, extending slightly the known southward range.
Distribution: 43°N-13°30’S in Atlantic; Indian Ocean.
C. mamillata Miller. This species was taken in hauls with 1100-4000
m of cable out from 30°S—37°30’S, 30°W, and from 42°S—44°S in the
Argentine Basin.
Distribution; 48°N-56°S in the Atlantic; Indian and Pacific Oceans.
Elegans Group Miller
C. elegans Sars. This was one of the commonest species, and occurred
from the equator to 55°S. Between the equator and 25°S, males and
females were 1.2-1.3 mm long; between 30°S and 40°S larger specimens
up to 1.60-1.65 mm long were found; south of 45°S females were 1.5—1.8
mm long and males 1.75—1.85 mm in length.
Distribution: 80°N-55°S in Atlantic; Indian and Pacific Oceans.
Procera Group Miiller
C. microprocera Angel. C. microprocera was taken at 14 stations be-
tween the equator and 25°S in the 200-0 m vertical tows.
Distribution: 32°N-25°S in Atlantic.
362 Proceedings of the Biological Society of Washington
C. procera Miiller. This species occurred from the equator to 40°S
and was not found south of the subtropical convergence.
Distribution: 32°N-—40°S in the Atlantic; Mediterranean and Indian
Ocean.
C. brachyaskos Miiller. C. brachyaskos was recorded from 37°30’S to
55°S, 30°W and from 42°S—45°S in the Argentine Basin, from hauls
with 1000-4000 m of cable out. Females (Fig. 3a) were 1.5-1.6 mm
and males (Fig. 3b) 1.4-1.5 mm long. Although larger than the speci-
mens recorded from the Sargasso Sea (Deevey, 1968), the frontal organs
(Fig. 3c, d) and male claspers (Fig. 3f, g) were similar. Also the
armature (Fig. 3e) of the male first antenna had no more teeth or
spinules than that of the smaller males.
Distribution: 32°N-56°30’S in the Atlantic; Indian and Pacific
Oceans.
Acuminata Group Miller
C. acuminata (Claus). This species was taken in only 4 of the 200-0
m tows collected between 2°26’S and 6°S.
Distribution: 46°N-37°S in the Atlantic; Pacific and Indian Oceans.
Obtusata Group Miiller
C. obtusata Sars var. antarctica Miller. C. obtusata is a bipolar
species. C. obtusata antarctica was found from 40°S to 50°30’S, 30°W
and from 42°S to 51°30’S in the Argentine Basin, in hauls with 100-200
m of cable out, as well as the deeper tows. The length range for males
(Fig. 3j) was 1.25-1.35 mm and for females (Fig. 3i) 1.65-1.85 mm.
The antarctic form differs from the arctic form primarily in the shape
of the frontal organs (Fig. 3n, 0) and the right male clasper (Fig. 3m)
and usually in the shape of the posterior end of the shell. The capitulum
of the frontal organs of both males and females is sharply pointed, and
the right male clasper is bent at a more acute angle than in the arctic
forms. The armature of the principal seta of the male first antenna
(Fig. 3p) is apparently similar.
Distribution: 23°S-68°S in Atlantic; 26°S-68°S in the Pacific. C.
obtusata obtusata occurs chiefly north of 60°N in the Atlantic and Arctic
Ocean and at 58°N in the Skager Rak and Cattegat.
Rotundata Group Miller
This group is greatly in need of taxonomic revision. Miller originally
included 7 species in this group, which differs from all others in the
position of the asymmetrical glands: C. macromma, C. pusilla major,
C. pusilla minor, C. glandulosa, C. kyrtophora, C. nasotuberculata, C.
rotundata and C. isocheira. C. rotundata has since been separated into
3 species (Iles, 1953): C. rotundata, C. skogsbergi Iles and C. tereti-
valvata Iles, and Rudyakov (1962) has described C. abyssalis from the
Pacific. Eleven species were found in the HUDSON 70 samples, in-
Pelagic Ostracods from South Atlantic 363
Fic. 3. a and b, Lateral views of female and male Conchoecia
brachyaskos; c and d, Capitulum of frontal organs of male and female;
e, Armature of male first antenna; f and g, Right and left male claspers;
h, Female furca; i and j, Lateral views of female and male C. obtusata
antarctica; k, Endopodite of male left second antenna; | and m, Left
and right male claspers; n, Capitulum of frontal organs of two males;
o, Capitulum of frontal organs of two females; p, Armature of principal
seta of male first antenna; gq, Endopodite of female second antenna;
r, Lateral view of female C. gaussae. Scale on a for a, b, i and j, at upper
right for h, beside e for e, below p for p, at bottom center for c, d, f, g,
k-o and q, on r for r. Scales in mm.
364 Proceedings of the Biological Society of Washington
cluding two new forms, which, due to insufficient material, will not be
described in this report. Of the known species, only C. abyssalis and
C. pusilla were not recorded.
C. teretivalvata Iles (Figure 4). This species was recorded from 5°S
in the 200-0 m tows and from 30°S—49°S, 30°W and from 42°S—48°S
in the Argentine Basin. As in the case of C. skogsbergi, smaller and
larger forms were present, and the larger forms may eventually prove
to be a different species, but males of both sizes had the large penis
(Fig. 41) shaped as figured by Miller (1894, Pl. 6, Fig. 20). Females
and males, 0.85-0.90 mm long (Fig. 4b, g, h), occurred down to
42.°30’S and at 42°S in the Argentine Basin. Larger specimens 1.0-1.1
mm long (Fig. 4a, j) were found from 40°S—49°S, between the sub-
tropical and subantarctic convergences. Smaller males had 9-10 pairs
of teeth on the principal seta of the first antenna, and larger males 10-11
pairs of teeth (Fig. 4e). Larger and smaller males had claspers of
similar shape (Fig. 4c, d, i).
Distribution: Mediterranean; warmer Atlantic to 29°S in Benguela
Current. These specimens extend the range southward.
C. rotundata Miiller. This species, as described from the Sargasso Sea
(Deevey, 1968), occurred from the equator to 35°S. The length range
for females was 0.87-0.95 mm and for males 0.80-0.85 mm (Fig. 5h).
Distribution: 32°N-35°S in Atlantic; tropical Pacific.
C. skogsbergi Iles. C. skogsbergi was recorded from 30°S—55°S,
30°W and from 42°S-47°S in the Argentine Basin, from all stations
where samples were collected with 1000 or more meters of cable out.
Smaller specimens, females 1.10—1.25 mm long and males 1.05-1.20 mm
long, such as those described from the Sargasso Sea (Deevey, 1968)
were found from 30°S-—35°S where larger specimens 1.35-1.50 mm long
were also present. South of 35°S females were 1.4-1.7 mm long and
males 1.35-1.75 mm long (Fig. 5b). Males 1.7 mm long had 14 pairs
of teeth on the principal seta of the first antenna, whereas males 1.25
mm long had 13 pairs of teeth.
Distribution: 32°N-—65°S in the Atlantic.
C. nasotuberculata Miller. This species (Fig. 5a) was found from
the equator to 42°30’S, just south of the subtropical convergence.
Distribution: 18°N-—40°S; Indian Ocean and Mediterranean. These
specimens extend the range slightly south.
C. kyrtophora Miller. This is not the species described (Deevey,
1968) as C. kyrtophora from the Sargasso Sea. One female 0.93 mm
long (Fig. 5g) and one male 0.95 mm long were collected at 40°S in a
haul with 2500 m of cable out. This species is closely related to C.
nasotuberculata, but the female is distinguished by lacking the lateral
bumps on the shell that are characteristic of female C. nasotuberculata.
Distribution: 14°N-35°S in Atlantic; Indian Ocean. These speci-
mens extend the range 5° south.
C. macromma Miller. Two females 1.40 (Fig. 5c, d) and 1.27 mm
long and one male 1.50 mm long were collected at 44°12.4’S, 42°46’W
Pelagic Ostracods from South Atlantic 365
Fic. 4. Conchoecia teretivalvuata. a, A 1.10 mm male; b, 0.92 mm
female; c, Endopodite of right second antenna of 1.07 mm male;
d, Right clasper of 1.05 mm male; e, Frontal organ and first antenna
of 1.05 mm male; f, Frontal organ and first antenna of 0.92 mm
female; g and h, Lateral and dorsal views of 0.85 mm male; i, Left and
right claspers of 0.9 mm male; j, 1.1 mm female; k, Capitulum of
frontal organ of 0.9 mm male; 1, Penis of 1.0 mm male. Scale on a for a,
b, g, h and j, at upper right for cf, i, k and 1. Scales in mm.
366 Proceedings of the Biological Society of Washington
Fic. 5. a, Lateral view of 0.85 mm female Conchoecia nasotuber-
culata; b, 1.6 mm male C. skogsbergi; c and d, Lateral and dorsal views
of 1.4 mm female C. macromma; e and f, Capitulum of frontal organs of
male and female C. macromma; g, Lateral view of 0.93 mm female
C. kyrtophora; h, 0.85 mm male C. rotundata; i, 0.93 mm female C.
isocheira; j and k, Lateral and posterior views of 1.95 mm female
C. glandulosa. Scale on b for a, b, g—i, on c for c and d, in center for
e and f, on j for j andk. Scales in mm.
Pelagic Ostracods from South Atlantic 367
in a haul with 4000 m of cable out. Although these specimens are larger
than those described by Miiller from 24°N-3°S in the Atlantic, the
position of the asymmetrical glands, the frontal organs (Fig. 5e, f), and
the structure of the first and second antennae agree with his figures.
Iles (1953) listed one female from a 750-1000 m haul at 25°S in the
Benguela Current, but did not give its size or any corroboratory data.
Distribution: 24°N-25°S in the Atlantic; Indian Ocean. These speci-
mens extend the range to 44°S.
C. glandulosa Miiller. One female 1.95 mm long (Fig. 5j, k) was
caught at 44°S in the Argentine Basin in a haul with 4000 m of cable out.
Distribution: 32°N-64°S in the Atlantic; Pacific and Indian Oceans.
C. isocheira Miller. C. isocheira (Fig. 5i) occurred from 49°S-55°S,
in hauls with a minimum of 500 m of cable out. The length range for
females was 0.95-1.05 mm and for males 0.84—1.00 mm.
Distribution: 53°S-71°S in the Atlantic, Indian and Pacific Antarctic.
The HUDSON 70 specimens extend the range a few degrees to the north.
Curta Group Miller
C. curta Lubbock. This species was found from the equator to 35°S,
and did not occur south of the subtropical convergence.
Distribution: 48°N-37°S in the Atlantic; Indian and Pacific Oceans
and Mediteranean.
C. echinulata (Claus). C. echinulata was present from 11°S-30°S.
Distribution: 34°N-37°S in Atlantic; Indian Ocean.
C. stigmatica Miiller. Juvenile specimens of this species were noted
from the equator to 10°S in the 200-0 m tows.
Distribution: 0°39’N-31°S in Atlantic; Indian and Pacific Oceans.
C. acuticosta Miiller. This species occurred from 32°30’S—45°S, 30°W
and from 42°S-44°S in the Argentine Basin, primarily in hauls with
1000-4000 m of cable out.
Distribution: 27°N-36°S in the Atlantic; Indian and Pacific Oceans.
These specimens extend the range to 45°S.
Bispinosa Group Miiller
C. haddoni Brady and Norman. C. haddoni was present from 35°S to
51°30’S, 30°W, and from 42°S—47°S in the Argentine Basin, in hauls
with 1000-4000 m of cable out. The length range noted for females
was 2.8-3.1 mm and for males 2.30-2.55 mm. Miller noted that the
southern forms were larger than those from the North Atlantic, where
Angel (1970) recorded length ranges of 2.15-2.53 mm for females and
1.75-2.00 mm for males.
Distribution: 60°N-51°30’S in Atlantic; Indian Ocean.
C. bispinosa Claus. C. bispinosa was found from the equator to 40°S,
30°W and at 42°S in the Argentine Basin. Ramirez and Moguilevsky
(1971) identified all their specimens, collected between 36° and 38°S,
as C. secernenda, but their smaller males around 2.1 mm long were
368 Proceedings of the Biological Society of Washington
evidently C. bispinosa, since they noted 29 pairs of teeth on the principal
seta of the male first antenna. Apparently, as in the case of C. haddoni,
specimens of C. bispinosa are larger in the South Atlantic than in the
North. HUDSON 70 females were 1.85-2.20 mm long and males
1.90-2.00 mm long. Angel (1970) recorded length ranges of 1.8-2.1
mm for females and 1.65—1.88 mm for males from around 29°N-33°N
in the eastern North Atlantic. Females from the Sargasso Sea were
1.80-1.95 mm long and males 1.65-1.80 mm long (Deevey, 1968).
Distribution: 43°N-—42°S in the Atlantic.
C. secernenda Vavra. This species was caught only in several 200-0
m tows between 8°30’S and 27°30’S, 30°W, and apparently is not as
abundant in the South Atlantic as in the North. Judging by their large
size (females 2.6 mm and males 2.3 mm mean length) Poulsen’s (1969b )
specimens identified as C. bispinosa from the Gulf of Guinea are pre-
sumably C. secernenda; he did not state the number of teeth on the
principal seta of the male first antenna, but believes C. secernenda and
C. bispinosa to be synonymous.
Distribution: 37°N-38°S in the Atlantic.
C. atlantica (Lubbock). C. atlantica was found only between 0°51’S
and 7°27’S, 30°W.
Distribution: 40°N-37°S in the Atlantic; Indian and Pacific Oceans.
Gaussae Group Skogsberg
Skogsberg (1920) included C. gaussae and C. incisa in this group,
both of which Miiller (1906a) had put in the Bispinosa Group. In this
report C. edentata, which Miiller assigned to the Acuminata Group, is
also included. These three species differ from all others in having large
compound glands on the mid-ventral margin of both shells, as well as
the usual asymmetrical glands. Poulsen (1969b) has provisionally in-
cluded C. congolensis Poulsen in this group, although it lacks the glands
on the ventral margin.
C. edentata Miiller. Three females, 1.65, 1.73, 1.73 mm long, and two
juvenile specimens 1.40 and 1.45 mm long were collected between 23°S
and 55°S. Miiller described this species from a 1.7 mm male and an
immature female. Barney (1921) recorded 4 specimens from a 0—1000
m haul at about 72°S in the Pacific Antarctic, and Iles (1953) a single
female from a 250-500 m tow at 23°S in the Benguela Current.
Rudyakov (1962) gave a supplementary description of a 1.83 mm female
and 2 males, 1.43-1.45 mm long.
Distribution: 0°39’N-56°S in the Atlantic; 55°S—62°S in the Indian
Ocean; 72°S and 44°N-50°N in the Pacific.
C. incisa Miiller. A single female, 2.37 mm long, was caught in a
200-0 m tow at 3°40’S, 30°W. Miiller (1906a) described this species
from a 2.15 mm male and a 2.5 mm female from the Indian Ocean.
Poulsen (1969b) has since recorded 3 females, 2.5-2.6 mm long, from
a station near the equator in the Gulf of Guinea. Angel (1969) noted
this species from the Canary Island region.
Pelagic Ostracods from South Atlantic 369
Distribution: 29°N-3°40’S in the Atlantic; 2°S—26°S in the Indian
Ocean.
C. gaussae Miiller. One female, 3.45 mm long, was caught at 32°30’S
in a haul with 750 m of cable out, and another female, 3.85 mm long
(Fig. 3r), was taken at 51°30’S in a tow with 3000 m of cable out.
Miiller (1908) described the species from a single male, 3.1 mm long,
from 35°11’S, 2°43’E. Skogsberg (1920) described the female from a
3.6 mm specimen from 50°S in the Atlantic. A subspecies, C. gaussae
curilensis, has been described from the Kuril Kamchatka region from two
females 4.08—4.18 mm long (Rudyakov, 1962). The only other recorded
specimens are two 4.0 mm females and a 3.3 mm male from 2°S-6°35’S
in the Gulf of Guinea (Poulsen, 1969b).
Distribution: 2°S—51°30’S in the Atlantic; 46°31’N, 154°22’E in the
Pacific
Loricata Group Miller
C. loricata loricata (Claus). A single male, 1.8 mm long, was found
at 35°S in a haul with 1800 m of cable out.
Distribution: 46°N-37°S in the Atlantic; Indian Ocean and Medi-
terranean.
C. ctenophora Miiller. One female, 2.7 mm long, was taken at 30°S
in a haul with 1400 m of cable out.
Distribution: 18°N-35°S in the Atlantic; near equator in Indian
and Pacific Oceans.
Serrulata Group Skogsberg
C. concentrica Miller. A single female, 1.55 mm long, was found at
0°51’S.
Distribution: 38°N-3°S in the Atlantic; Malay Archipelago and
Indian Ocean.
C. serrulata Claus. This species occurred from 40°S-51°30’S, in hauls
with 100 or more meters of cable out. It was most numerous between
47°S and 49°S, over the convergence in subantarctic waters in the
Argentine Basin. The length range for females was 1.40-1.65 mm and
for males 1.25-1.35 mm.
Distribution: 10°S-58°S in the Atlantic; to 59°S in the Indian Ocean
and to 68°S in the Pacific.
Magna Group Miiller
C. magna Claus. C. magna occurred from the equator to 42°30’S, and
was present in almost all 200-0 m tows. Females had a length range of
1.7-2.0 mm and males of 1.6-1.9 mm.
Distribution: 52°N-55°S in the Atlantic; Indian and Pacific Oceans
and Mediterranean.
C. lophura Miiller. This species, easily recognized by the large group
of gland cells on the ventral margin at the postero-ventral corner of the
370 Proceedings of the Biological Society of Washington
shell, was found from 30°S—45°S, in hauls with 750-2500 m of cable out.
Distribution: 46°N-—48°S in the Atlantic; Pacific and Indian Oceans
and Mediterranean.
C. parvidentata Miller. C. parvidentata occurred from 35°S—48°S, to
the region of the subantarctic convergence. Females had a length range
of 2.55—2.85 mm and males of 2.15—2.50 mm.
Distribution: 31°N—48°S in Atlantic; Indian Ocean.
C. hyalophyllum Claus. This species was recorded from 6°S in a
200-0 m tow and from 27°30’S—42°30’S. Females were 1.65-1.80 mm
long and males 1.55-1.70 mm long.
Distribution: 52°N-—48°S in the Atlantic; Indian Ocean.
C. macrocheira Miiller. C. macrocheira was noted on only two oc-
casions, at 0°51’S and at 27°30’S, in hauls with 200-450 m of cable out.
Distribution: 60°N-34°S in the Atlantic; Indian and Pacific Oceans.
C. subarcuata Claus. This species was recorded from 8°31’S—37°30'S,
and did not occur south of the subtropical convergence.
Distribution: 59°N-56°S in the Atlantic; Indian and Pacific Oceans.
C. parthenoda Miiller. C. parthenoda was found from the equator to
30°S. The two specimens described as C. parthenoda by Ramirez and
Moguilevsky (1971, p. 657, Pls. 12-13) may represent C. magna; at least
in the case of the male the size is too great and the left asymmetrical
gland is not placed far enough forward on the dorsal margin.
Distribution: 37°N-30°S in the Atlantic; Indian and Pacific Oceans.
C. pseudoparthenoda Angel. This species occurred from the equator
to 30°S in 200-0 m tows. The male specimen figured as C. parthenoda
from Barbados (Deevey, 1970) is now recognized as C. pseudoparth-
enoda. Also, in all probability Poulsen’s (1969b) description and figures
of his male C. parthenoda represent C. pseudoparthenoda.
Distribution: 14°N-30°S in the Atlantic.
C. spinirostris Claus. C. spinirostris was recorded from the equator to
40°S, 30°W and to 42°S in the Argentine Basin, and was not found south
of the subtropical convergence.
Distribution: 45°N-24°S in the Atlantic; Pacific and Mediterranean.
These specimens extend the range to 42°S.
C. porrecta Claus. This species was found from the equator to 37°30’S,
and also did not occur south of the subtropical convergence. Because of
their size, the specimens identified by Poulsen (1969b) as C. spin-
irostris must be C. porrecta; Poulsen does not separate these two species.
Distribution: 41°N-2°N in the Atlantic; Mediterranean. These
specimens extend the range to 37°30’S.
Mollis Group Miiller
C. amblypostha Miller (Figure 6). This species occurred from 30°S—
40°S, 30°W and from 42°S—47°S in the Argentine Basin, in hauls with
1100-3000 m of cable out. Poulsen (1969b) recorded this species from
1°S—17°S in the Gulf of Guinea, from 670-800 m depths. Miller (1906a)
Pelagic Ostracods from South Atlantic 371
Fic. 6. Conchoecia amblypostha. a, Lateral view of female; b,
Endopodite of male right second antenna; c, Left clasper; d, Female
first antenna; e, Endopodite of female second antenna; f, Capitulum
of female frontal organ; g, Posterior view of female; h, Male penis;
i, Armature of distal secondary seta of male first antenna; j, Capitulum
of male frontal organ; k, Lateral view of male. Scale on a for a, g, k,
below i for b-f, i, j, at upper right for h. Scales in mm.
372 Proceedings of the Biological Society of Washington
was unable to differentiate the females (Fig. 6a, g) from those of C.
mollis, but males are distinguished by the armature of the first antenna,
particularly the distal secondary seta (Fig. 6i) and the shape of the penis
(Fig. 6h). No C. mollis males were noted. Females had a length range of
3.30-3.75 mm and males of 3.2-3.5 mm. The female frontal organ (Fig.
6f£), first and second antennae (Fig. 6d, e) are indeed similar in structure
and spination to those of female C. mollis. The male claspers (Fig. 6b,
c) are also similar in the two species, but the male C. mollis lacks the
long hairs on the “b” bristle of the endopodite of the second antenna
that are present in male C. amblypostha.
Distribution: 1°S—37°31’S in the Atlantic; Indian Ocean. These
specimens extend the range to 47°S.
C. dichotoma Miiller. A single immature female, 2.04 mm long, was
taken at 42°30’S in a haul with 2500 m of cable out. Poulsen (1969b)
recorded 3 specimens from 0°50’S.
Distribution: 0°50’S—35°S in the Atlantic. This specimen extends the
range to 42°30’S.
C. plactolycos Miiller (Figure 7). Muller described two subspecies of
C. plactolycos: C. plactolycos typica (1906a) = C. plactolycos plactoly-
cos (1912) and C. plactolycos major (1906a, 1908). Specimens of both
of these forms were collected in the HUDSON 70 samples and are
easily distinguishable, so that each form should be raised to the rank
of species. According to rule, C. plactolycos plactolycos thus becomes
C. plactolycos Miiller and C. plactolycos major is named C. major Miller.
Miller described C. plactolycos from a single male 2.3 mm long from
29°S in the Indian Ocean. One male, 2.5 mm long, was taken at
44°12.4’S, 42°46’W in a haul with 4000 m of cable out. This specimen
(Fig. 7c, d, g-k) agrees with Miiller’s description in the shape of the
shell, location of the glands, shape of the frontal organ and clasping
organs and the armature of the first antenna (Miiller, 1906a, p. 114, PI.
25, Figs. 14-20). C. plactolycos is closely similar to C. major, but is
smaller in size and the sculpturing of the shell is difficult to discern.
The principal seta of the male first antenna has 33 pairs of short fat
teeth (Fig. 7i) as against 43 pairs for the C. major male (Fig. 8i) and
the distal secondary seta is not armed (Fig. 7d) as in the larger form.
The medial glands on the posterior margin are much more visible in the
C. plactolycos male (Fig. 7k). A single female, 2.7 mm long (Fig. 7a, b),
taken at 41°44.6’S, 46°33’W, resembles the male in shape and sculptur-
ing. The female differs from that of C. major in having fewer spinules
on the basal segments of the first antenna (Figs. 7f and 8d) and in
lacking spinules on the anterior side of the principal seta. Unfortunately,
the distal part of the capitulum of the frontal organ was broken off
(Fig. Te).
Distribution: 42°S—44°S in the Atlantic; 29°S in the Indian Ocean.
C. major (= C. plactolycos major) Miller (Figure 8). C. major was
recorded from 49°S—55°S, 30°W and from 44°S-47°S in the Argentine
Pelagic Ostracods from South Atlantic 373
Fic. 7. Conchoecia plactolycos (= C. plactolycos plactolycos). a and
b, Lateral and ventral view of female; c, Capitulum of male frontal
organ; d, Male first antenna; e, Proximal part of capitulum of female
frontal organ; f, Female first antenna; g, Male penis; h, Male right
clasper; i, Armature of principal seta of male first antenna; j, Endopo-
dite of male left second antenna; k, lateral view of male. Scale on a
for a and b, on k for k, in center for j and h, at bottom right for i, on
right margin for c-g. Scales in mm.
Basin in hauls with 2500-4000 m of cable out. The length range for
females was 3.1-3.6 mm and for males 2.97-3.30 mm. Apparently this
is the first record of this species since Miller (1906a, 1908) described it
from 56°S in the Atlantic and 62°S-65°S in the Indian Ocean. The
374 Proceedings of the Biological Society of Washington
Fic. 8. Conchoecia major (= C. plactolycos major). a, Lateral view
of female; b, Endopodite of male right second antenna; c, Male left
clasper; d, Frontal organ and first antenna of female; e, Capitulum of
male frontal organ; f, g and h, Posterior, ventral and lateral views of
male; i, Armature of male first antenna. Scale above h for a, fh, beside
i for i, at left center for b—e. Scales in mm.
sculpturing on the shell (indicated in Fig. 8a, h) is much more definite
than in C. plactolycos; other differences have been noted above.
Distribution: 56°S in Atlantic; Indian Ocean. These specimens ex-
tend the range north to 44°S.
Pelagic Ostracods from South Atlantic 370
C. borealis Sars var. antipoda Miller. C. borealis antipoda occurred
from 35°S—55°S, 30°W and from 42°S—51°28’S in the Argentine Basin,
in hauls with 1000—4000 m of cable out. The length range for males was
2.80-3.05 mm and for females 3.05-3.35 mm. The antarctic form is
larger than the arctic form for which Skogsberg (1920) gave measure-
ments of 2.1-2.3 mm for males and 2.4—2.9 mm for females. The female
has more sharp-edged shoulder vaults (Fig. 9a, b) than the male (Fig.
9c, h). The male appears to differ from male C. borealis Sars, as figured
by Skogsberg (1920, Fig. 135-4), in having fewer teeth on the principal
seta (Fig. 9e); 40-45 pairs of teeth were noted, whereas Skogsberg
described 50-55 pairs of teeth on the principal seta of males from
Lofoten. The clasping organs and frontal organ (Fig. 9d, f, g) are
similar to those of the arctic form.
Distribution: 1°S-—65°S in the Atlantic and Antarctic Oceans.
Imbricata Group Miiller
C. imbricata (Brady). This species was recorded at 18°S, 24°S and
from 27°30’S—40°S, and was not found south of the subtropical con-
vergence.
Distribution: 63°N-—55°S in the Atlantic; Pacific and Indian Oceans.
C. ametra Miiller. C. ametra was taken from 30°S—40°S, 30°W and
rom 42°S—44°S in the Argentine Basin, in hauls with 1100-4000 m of
cable out. A length range of 3.75-4.10 mm was noted for males and of
4.5—-4.8 mm for females.
Distribution: 60°N-37°S in the Atlantic; Indian and Pacific Oceans.
These specimens extend the range south to 44°S.
C. plinthina Miiller. A single juvenile specimen, 4.1 mm long, was
found at 42°30’S in a haul with 2500 m of cable out. This is the largest
species of this group.
Distribution: 48°N-31°S in the Atlantic; Indian and Pacific Oceans.
This specimen extends the range to 42°30’S.
C. symmetrica Miller. C. symmetrica occurred from 35°S—55°S, 30°W
and from 42°S—47°S in the Argentine Basin, in hauls with 1000-4000 m
of cable out. The length range for females was 4.1-4.5 mm and for
males 3.7—4.0 mm.
Distribution: 0-55°S in the Atlantic; Indian Ocean and Pacific
Antarctic.
Alata Group Miiller
C. hettacra Miiller. C. hettacra (Fig. 9i-k) occurred from 49°S-—55°S,
30°W and from 42°S-51°30’S in the Argentine Basin, at shallow depths,
since it was taken in the 200 and 100 m tows. The length range for
females was 2.25-2.45 mm and for males 1.80-1.95 mm.
Distribution: 47°S-57°S in the Atlantic, so these specimens extend
the range north to 42°S; 43°S—70°S in the Indian Ocean.
C. valdiviae Miiller. This species occurred from 32°30’S—42°30’S,
376 Proceedings of the Biological Society of Washington
Fic. 9. a and b, Lateral and posterior views of female Conchoecia
borealis antipoda; c, Lateral view of male; d, Capitulum of male frontal
organ; e, Armature of male first antenna; f and g, Right and left male
claspers; h, Posterior view of male; i, Lateral view of female C. hettacra;
j, Armature of C. hettacra male first antenna; k, Lateral view of male;
] and m, Lateral views of female and male C. chuni. Scale on a for a-—c
and h, beside f for f and g, on lower right margin for d, e, and j, on i
for i, k-m. Scales in mm.
Pelagic Ostracods. from South Atlantic O77
Fic. 10. Conchoecia belgicae. a, b and c, Lateral, ventral and pos-
terior views of female; d, Female frontal organ and first antenna; e,
Capitulum of male frontal organ; f, Armature of principal seta of male
first antenna; g and h, Right and left male claspers; i, Endopodite of
female second antenna; j, Endopodite of male right second antenna;
k, Lateral view of male. Scale above k for a—c and k, at bottom for d, e,
i, j, on right margin for f-h. Scales in mm.
378 Proceedings of the Biological Society of Washington
30°W and from 42°S—45°S in the Argentine Basin. C. valdiviae is ex-
ceptionally large and bulky, deep red in color when alive. Most speci-
mens were immature, but one female was 5.35 mm long and one male
5.0 mm in length.
Distribution: 28°N in the Gulf of Mexico (Kornicker, 1968) to 47°S
in the Atlantic; to 58°S in the Indian Ocean.
C. belgicae Miiller (Figure 10). One 2.8 mm male was obtained at
51°30’S, 40°W, in a haul with 600 m of cable out, and one female 2.9
mm long was taken at 55°S in a haul with 1100 m of cable out. These
specimens agree with Miller's (1906b) and Skogsberg’s (1920) de-
scriptions and figures. Miiller’s “Belgica” females were 2.4—2.7 mm long
and his males 2.4-2.6 mm long, but speciemns from the Gausstation
(66°2.9’S, 89°38’E) were larger: females 2.6—-3.0 mm long and males
2.6-2.8 mm in length. Figure 10 illustrates the shape of the male and
female shells and the structure of the first and second antennae. Ac-
cording to Skogsberg, the male first antenna has 27-30 pairs of teeth,
similar in shape to those of the C. hettacra male (Fig. 9j). The HUDSON
70 specimen had 26 teeth in lateral view, the two rows unevenly paired
(Fig. 10f).
Distribution: Muller described this species from 70°S-71°S in the
Pacific Antarctic, and obtained it later from 65°S-66°S in the Indian
Antarctic. Skogsberg’s specimens were from about 66°S in the Atlantic.
Barney (1921) reported it abundant at a number of stations in the Ross
Sea and McMurdo Sound and also at two stations north of New Zealand.
The HUDSON 70 specimens extend the range in the Atlantic north to
51°30'S.
Daphnoides Group Miller
C. daphnoides (Claus). C. daphnoides was recorded from about 5°S
and 23°S to 40°S, 30°W, and at 42°S in the Argentine Basin, and there-
fore was not found south of the subtropical convergence. Males ranged
in length from 2.8-3.1 mm and females from 4.8-5.6 mm.
Distribution: 60°N-37°S in the Atlantic; Indian and Pacific Oceans.
These specimens extend the range to 42°S.
C. chuni Miiller. C. chuni (Fig. 91, m) was taken from 32°30’S-55°S,
30°W, and from 42°S-50°S in the Argentine Basin, in shallow as well
as deeper hauls. The length range for males was 1.40-1.55 mm and
for females 2.0-2.4 mm.
Distribution: 26°S-53°S in the Atlantic; 2°S—44°S in the Indian
Ocean, and to 64°S in the Pacific. These specimens extend the range
slightly to 55°S.
LITERATURE CITED
AncEL, M. V. 1969. Planktonic ostracods from the Canary Island
region; their depth distributions, diurnal migrations, and com-
munity organization. J. Mar. Biol. Assoc. U. K., 49:515-553.
Pelagic Ostracods from South Atlantic 379
1970. The redescription of Conchoecia bispinosa Claus, C.
haddoni Brady & Norman and C. secernenda Vavra from the
North Atlantic. Crustaceana, 18:147-166.
Barney, R. W. 1921. Crustacea. Part V. Ostracoda. Brit. Antarctic
(Terra Nova) Exped., 1910, Zool. Nat. Hist. Rep., 3(7):175-
189.
Derevey, G. B. 1968. Pelagic ostracods of the Sargasso Sea off Ber-
muda. Peabody Mus. Nat. Hist., Yale, Bull. 26, 125 p., 65 figs.
——. 1970. Pelagic ostracods (Myodocopa Halocyprididae) from
the North Atlantic off Barbados. Proc. Biol. Soc. Washington,
82:799-824.
Ines, E. J. 1953. A preliminary report on the Ostracoda of the
Benguela Current. Discovery Rep. 26:259-280.
Kornicker, L. S. 1968. Bathyal Myodocopid Ostracoda from the
northeastern Gulf of Mexico. Proc. Biol. Soc. Washington, 81:
439-472.
MULLER, G. W. 1894. Die Ostracoden des Golfes von Neapel. Fauna
und Flora des Golfes von Neapel, Monogr. 21:1—404, 40 Pls.
1906a. Ostracoda. Wissensch. Ergeb. d. Deutschen Tiefsee-
Expedition auf dem Dampfer “Valdivia” 1898-1899, 8:1-154,
Pls. 5-35.
1906b. Ostracoden. Expedition Antartique Belge. Resultats
du Voyage du S. Y. Belgica en 1897—1898-1899. Rapp. Scient.,
Zool., 7 p., 1 Pl.
1908. Die Ostracoden der Deutschen Siidpolar-Expedition
1901-1903. Deutsche Siidpolar-Expedition 1901-1903, X, Zool.
II: 53-181, Pls. 4-19.
1912. Ostracoda. Das Tierreich, Lief. 31, 434 p., Berlin.
PouLseN, E. M. 1962. Ostracoda-Myodocopa Part I Cypridiniformes-
Cypridinidae. Dana-Report No. 57, 414 p., 181 figs.
1969a. Ostracoda-Myodocopa Part IIIA Halocypriformes-
Thaumatocypridae and Halocypridae. Dana-Report No. 75, 100
p., 40 figs.
1969b. Ostracoda-Myodocopa from the eastern tropical At-
lantic. Vidensk. Meddr. Dansk Naturh. Foren. 132:129-197.
RAMiReEz, F. C., anp A. Mocurtevsxy. 1971. Ostracodos planctionicos
hallados en aguas oceanicas frente a la Provincia de Buenos
Aires (Resultados de la XLI Comissae Oceanografica Costa
Sul). Physis, 30( 81) :637-666.
Rupyakov, Yu. A. 1962. Ostracoda Myodocopa (family Halocypridae )
from the N. W. Pacific Ocean. Trudy Inst. Okeanol., 58:172-
201. 14 figs.
SxocsBERG, T. 1920. Studies on marine ostracods Part I (Cypridinids,
Halocyprids and Polycopids). Zool. Bidr. Uppsala, Suppl. I,
784 p., 153 figs.
380 Proceedings of the Biological Society of Washington
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Vol. 87, No. 33, pp. 381-388 31 December 1974
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
A NEW SPECIES OF ELEUTHERODACTYLUS
(AMPHIBIA: LEPTODACTYLIDAE) FROM THE
PACIFIC LOWLANDS OF ECUADOR
By Joun D. LyNcu
School of Life Sciences, The University of Nebraska, Lincoln
Soon after my description of Eleutherodactylus croceoin-
guinis (Lynch, 1968) from Provincia Napo, Ecuador, Dr.
Charles F. Walker called my attention to a series of a dimin-
utive eleutherodactyline frog that he had collected in 1962
in the vicinity of Santo Domingo de los Colorados, Prov.
Pichincha, Ecuador. His frogs were superficially similar to
E. croceoinguinis in size (less than 25 mm SVL) and color
pattern (presence of bright color patches on the concealed
surfaces of the limbs). At that time I had not done field
work in western Ecuador but had seen two examples of this
frog collected along the Rio Baba. Field work in 1968 and
1970 provided me the opportunity to collect additional ma-
terial at Santo Domingo de los Colorados as well as at several
other localities in western Ecuador and to make ecological
observations of this frog which is here named
Eleutherodactylus walkeri, new species
Holotype: The University of Kansas Museum of Natural History (KU)
131652, an adult male collected at Las Palmas (junction of highways
28 and 30), Prov. Pichincha, Ecuador, 920 m, 8 August 1970 by J. D.
Lynch.
Paratypes: KU 131653-63, California Academy of Sciences 134064-
66, collected syntopically with holotype; University of Michigan Museum
of Zoology 131675 (10 specimens), Hacienda Delta, ca. 8 km SE Santo
Domingo de los Colorados, Prov. Pichincha, Ecuador.
Diagnosis: (1) skin of dorsum finely shagreened, no dorsolateral
folds, that of venter coarsely areolate, discoidal folds prominent; (2)
33—Proc. Bion. Soc. WAsuH., VoL. 87, 1974 (381)
382 Proceedings of the Biological Society of Washington
tympanum visible externally, its length one-third eye length; (3) snout
subacuminate in dorsal view, rounded in lateral profile; (4) upper
eyelid slightly narrower than interorbital distance (IOD); no cranial
crests; (5) prevomerine odontophores oval, bearing teeth; (6) males
with vocal slits and median subgular vocal sac; (7) first finger shorter
than second, digits bearing discs on large, non-emarginate pads, discs
wider than long; subarticular tubercles simple; (8) fingers bearing
weak, keel-like lateral fringes; (9) ulnar tubercles obscure or lacking;
(10) inner tarsal tubercle minute, outer edge of tarsus with faint
tubercles; no heel tubercles; (11) inner metatarsal tubercle oval,
length 2-3 times width, 5-6 times size of conical outer metatarsal
tubercle; plantar surfaces bearing 0-3 supernumerary tubercles; (12)
toes bearing weakly defined lateral fringes, not basally webbed; digits
bearing discs and pads, pads slightly smaller than those of fingers;
(13) color pattern polymorphic; most common pattern is: dorsum gray
to brown with dark brown markings; limbs barred; venter cream with
brown flecking; posterior surface of thighs pale to medium brown,
sometimes with black patches enclosing cream spots; groin black with
large cream (yellow in life) spots; (14) adults small, males 15.3-17.9
mm, females 20.8-25.3 mm SVL.
E. walkeri is the only small South American eleutherodactyline frog
now known having visible tympana and yellow spotting on a black or
dark brown field in the groin. E. carvalhoi Lutz and E. croceoinguinis
are superficially similar frogs but have concealed tympana.
Description: Statements listed in the diagnosis are not repeated be-
low unless additional information is provided. Head about as wide as
body or slightly wider than body; head longer than wide; head width
33.7-87.6 percent SVL (x = 35.6); tip of snout extending beyond lower
jaw; snout short, eye-nostril distance slightly less than eye length; can-
thus rostralis moderately sharp, slightly concave or straight; loreal
region concave, sloping abruptly to lip; lips not flared; nostrils directed
laterally, protuberant; interorbital space flat; frontoparietals complete,
not produced laterally into crests; upper eyelid width 74.5-100.0 per-
cent IOD (x = 87.2); tympanum visible, its upper third usually con-
cealed by supratympanic fold; tympanum round, its length 24.1-34.7
percent eye length (x = 31.3), not sexually dimorphic in size or
shape, separated from eye by distance equal to tympanic diameter;
tongue large, longer than wide, weakly notched or entire along pos-
terior border, posterior two-fifths to one-third not adherent to floor of
mouth; choanae large, round, not concealed by palatal shelf of maxillae;
prevomerine teeth present on small (one-third size of a choana), low,
odontophores lying medial and posterior to choanae; 2-3 teeth per
odontophore, arranged in a row (or clump) across posterior edge of
odontophore.
Skin of dorsum, upper flanks, and limbs smooth to finely shagreened,
that of lower flanks and venter coarsely areolate; shank of males 52.1-
A new frog from Ecuador 383
54.4 percent SVL (x = 53.1), of females 44.9-57.0 percent (x = 49.7);
forearm lacking prominent ulnar fringe or tubercles, no antebrachial
tubercle; palmar tubercle bifid, larger than oval thenar tubercle; palm
bearing a few supernumerary tubercles; subarticular tubercles subconical,
round; all fingers bearing discs on simple pads; pad of thumb only
slightly wider than digit, pads on fingers 2-4 nearly twice width of
digit below pad.
Heel lacking tubercles in most individuals; when heel tubercles pres-
ent, small and non-conical; most individuals have a minute inner tarsal
tubercle and a series of small outer tarsal tubercles but a few indi-
viduals lack tarsal tubercles (possibly due to preservative); toe pads
wider than long but less than twice width of digit below pad; pads of
toes 3-5 largest; all toe pads smaller than finger pads (2-4) but larger
than thumb pad.
Preserved color pattern: Gray to brown dorsally, pale cream (with
indefinite infuscation on throat) to dark gray ventrally; labial bars,
canthal and supratympanic stripes, interorbital bar, scapular and sacral
chevrons, and lumbar bar dark brown (Fig. 1); limbs with oblique dark
brown bars separated by broader, paler interspaces; anal triangle dark
brown to black, contiguous with dark brown posterior surfaces of thighs;
posterior surfaces of thighs with or without black-ringed cream spots
(spots occur in 5 percent of specimens from Las Palmas and vicinity
of Santo Domingo de los Colorados, Prov. Pichincha, and in 65 percent
of specimens from Balzapamba, Prov. Bolivar); groin and anterior sur-
faces of thighs dark brown to black with colorless spots; concealed shank
brown with cream spots.
E. walkeri exhibits pattern polymorphism (Fig. 2). Three pattern
morphs are clearly evident (A—C), whereas a fourth (D) may not be
distinct from the most common morph (A). The morphs are as follows:
Morph A. Pattern of interorbital bar, scapular and sacral chevrons,
and lumbar bar; flanks and side of head not darker than
dorsum and top of head. (Fig. 1).
Morph B. Flanks and side of head much darker than dorsum and
top of head; dark vertebral stripe from tip of snout to
vent bordered by paravertebral dark stripes. No chevrons
or bars present.
Morph C. As in A except for fine, cream, dark-brown-edged verte-
bral stripe passing through chevrons and bars.
Morph D. As in A except for broad, pale cream, dorsolateral stripes;
chevrons and bars occur between dorsolateral stripes.
At three localities, samples are adequate to compute frequencies of the
morphs. Morph A is the most common at each locality (Santo Domingo
de los Colorados and vicinity—86.4%; Las Palmas—93.3%; and Balza-
pamba—76.9% ). The frequencies at the three localities are as follows
(data given as numbers of A:B:C:D): Santo Domingo de los Colorados
384 Proceedings of the Biological Society of Washington
Fic. 1. Eleutherodactylus walkeri new species (KU 120230).
and vicinity—19:0:1:2; Las Palmas—14:1:0:0; and Balzapamba—
SOFororle
Color in life: Gray, brown, rusty-brown, or black with black or dark
brown markings; dorsal surfaces irregularly flecked with cream; venter
dark gray to dull black with pale gray flecking; groin and anterior
and posterior surfaces of thighs (latter variable with respect to spotting )
dark brown to black with lemon yellow spots; iris chocolate-brown
reticulated with black or gray-bronze above, brown below; dark brown
horizontal streak through eye.
Natural history: All specimens collected were found in forested or
forest-analogue habitats. Perhaps due to its small size, E. walkeri has
not been found on the forest floor but has been found on herbs 10-30
cm above the ground. One individual was taken at night about 2.5 m
above the forest floor on an elephant-ear plant. The specimens from Las
Palmas were found on sticks and ferns in the spray zone of a small
waterfall (ca. 6-7 m in height) where the vegetation was constantly
buffeted by wind and spray. At Balzapamba, all but two individuals
were collected by day in the cut-off stumps of banana plants. Walker
found males calling both by day and night and described the call as
“a weak, single-syllabled ‘tink’, quite Acris-like in quality.”
A new frog from Ecuador 385
Fic. 2. Color pattern morphs of Eleutherodactylus walkeri new
species. Top row, left to right, KU 120221 (morph C), 120215 (morph
A), and 131628 (morph B). Lower row, left to right, KU 120221,
120215, 131628.
The only evidences of breeding were calling males (11 August 1962; 4
August 1968) at Hacienda Delta and Santo Domingo de los Colorados
and an amplectic pair (KU 131652-53) at the type-locality (8 August
1970). The female of the pair (KU 131653) is the largest specimen
examined (25.3 mm SVL) and has large, yellow ovarian eggs.
The known altitudinal range for E. walkeri is 220 m (Estacion Bio-
logica Rio Palenque) to 1270 m (ESE of Balzapamba). Collections
made on the Pacific slopes of the Andes at Tandapi, Prov. Pichincha
386 Proceedings of the Biological Society of Washington
(1450 m), Apuela, Prov. Imbabura (1650 m), Guaranda, Prov. Bolivar
(2500 m), and La Esperanza, Prov. Cotopaxi (1500 m), include several
eleutherodactyline species but not E. walkeri; these localities are pos-
sibly too high for E. walkeri. The leptodactylid sympatrants of E.
walkeri include two largely terrestrial species, Barycholos pulcher (di-
urnal) and E. achatinus (nocturnal), and several arboreal species (all
nocturnal) including E. areolatus, E. latidiscus, and E. ornatissimus
among the named forms. Of these, only E. achatinus occurs at eleva-
tions above 1270 m (found at Tandapi).
Measurements of the holotype in mm: SVL 17.9; tibia 9.4; head width
6.4; head length 6.6; upper eyelid width 1.6; IOD 2.1; tympanum length
0.8; eye length 2.5.
Etymology: Named for Charles F. Walker.
Other specimens examined: In addition to the type-material, the
following have been examined. ECUADOR, Prov. Bolivar: Balzapamba,
800 m, KU 131613-51; 6 km ESE Balzapamba, 1270 m, KU 142039-40.
Prov. Cotopaxi: 20.3 km W Pilalo, 830 m, KU 142034-38. Prov. El Oro:
32.6 km SSE Portovelo, 990 m, KU 142041. Prov. Los Rios: Estacion
Biologica Rio Palenque, 56 km N Quevedo, 220 m, KU 147567-68. Prov.
Pichincha: 3 km E Dos Rios, 1050 m, KU 142033; 3.7 km E Dos Rios,
1190 m, KU 142031-32; Rio Baba, 4 km E, 10 km S Santo Domingo de
los Colorados, Univ. Illinois Mus. Nat. History 93560-61; Santo Domingo
de los Colorados (vicinity of Hotel Zaracay), 660 m, KU 120215-21,
120227-31.
Remarks: Few species of Cochran and Goin’s (1970) Group II (skin
of venter areolate, first finger shorter than second) are as small as E.
walkeri. Of the Group II species, only E. carvalhoi (northwestern Brazil
and adjacent Colombia) and E. croceoinguinis (eastern Ecuador and
adjacent Peru) are likely to be confused with E. walkeri. These three
species have bright color patches in the groin (yellow in carvalhoi and
walkeri, orange in croceoinguinis). The bright spots are irregular in
shape and number in walkeri (Fig. 2), two are found on each side in
croceoinguinis (one on the flank and one on the anterior thigh—both
round), and one on the flank (round) in carvalhoi. The tympana are
concealed in carvalhoi and croceoinguinis and neither species exhibits
pattern polymorphism as in walkeri.
E. trachyblepharis (Boulenger) resembles E. walkeri in having visible
tympana but lacks bright color patches on the concealed surfaces and
does not exhibit pattern polymorphism. E. trachyblepharis has nar-
rower digital pads and is slightly smaller than E. walkeri; E. trachy-
blepharis is currently known only from the Pastaza valley on the Ama-
zonian versant of the Ecuadorian Andes. E. martiae Lynch, from the
Amazon basin in Ecuador, resembles E. walkeri in size, structure of the
hands and feet, and in exhibiting a similar pattern polymorphism. Un-
like E. walkeri, E. martiae has a short, rounded snout, lacks color
patches in the groin, does not have visible prevomerine odontophores,
A new frog from Ecuador 387
and has a concealed tympanum. The several other dwarf Eleuthero-
dactylus known from northwestern South America will not be confused
with E. walkeri. E. acuminatus Shreve has a protruding snout and
concealed tympana; E. diastema (Cope) and E. moro Savage have short,
fringed digits; E. gularis (Boulenger) has short, fringed digits and
papillae at the tips of the digits (as in E. areolatus, see Lynch, 1971);
E. paululus Lynch has a protruding snout and is much smaller; and
E. pseudoacuminatus Shreve has a truncate snout and concealed pre-
vomerine odontophores. None of these species has yellow spotting in
the groin or on the concealed limbs and none is polymorphic in color
pattern.
Acknowledgments: Field work in Ecuador was supported by grants
from the Penrose Fund (Grant no. 5549) of the American Philosophical
Society, a Watkins Museum of Natural History Grant, The University of
Kansas, and by the University of Nebraska Research Council. For loan
of specimens, I am grateful to William E. Duellman, Dorothy Smith,
Hobart M. Smith, and Charles F. Walker. Thomas H. and Patricia R.
Fritts aided me in field work. I am especially grateful to Charles F.
Walker who relinquished his specimens and field notes permitting me
to more completely describe this frog.
LITERATURE CITED
CocHran, D. M. anv C. J. Gorn. 1970. Frogs of Colombia. Bull.
U.S. Nat. Mus., 288: 1-655.
Lyncu, J. D. 1968. Two new frogs of the genus Eleutherodactylus
from eastern Ecuador (Amphibia: Leptodactylidae). J. Her-
petol., 2:129-35.
—. 1971. Redescriptions of three little-known Eleutherodactylus
from northwestern Ecuador (Amphibia: Leptodactylidae).
Trans. Kansas Acad. Sci., 73: 169-80.
388 Proceedings of the Biological Society of Washington
me OG 7_>
Vol. 87, No. 34, pp. 389-394 31 December 1974
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
A NEW SPECIES OF PSEUDOPOLYDORA
(POLYCHAETA, SPIONIDAE) FROM
THE SOLOMON ISLANDS
By WituiaM J. LicuT
Department of Invertebrate Zoology
California Academy of Sciences
San Francisco, California 94118
A new spionid polychaete of the genus Pseudopolydora
Czerniavsky was collected in large numbers from the north
coast of Guadalcanal Island in a lagoon at the mouth of the
Lunga River by Peter E. Gibbs during the Royal Scientific
Expedition to the Solomon Islands in 1965. The animals were
taken from the interstices of a sponge growing around sub-
merged tree roots, in association with the cosmopolitan ser-
pulid, Mercierella enigmatica Fauvel. The new species was
found in water of very low salinity (5.6 %.). A more complete
account of the ecology and faunal associations of this worm
may be found in a monograph on the polychaetes of the
Solomon Islands (Gibbs, 1971). I wish to thank Dr. Gibbs
who kindly provided the specimens upon which this paper
is based and in whose honor the species is named.
Pseudopolydora gibbsi, new species
Figures 1-8
Pseudopolydora sp., Gibbs, 1971: 173-174.
Diagnosis: Prostomium anteriorly entire, no occipital or nuchal cirrus;
all notosetae, except posterior hooks, smooth simple capillaries, first
present on setiger 1; stout, curved, notopodial hooks in posterior 15-17
segments; neuropodial crotchets from setiger 8, 6-8 per ramus; setiger
5 with 2 types of modified spines: (1) anterior row of stout spathe-
like setae, with hollow ends bearing acuminate tips, and (2) posterior
row of heavy, curved falcigers; modified spines of setiger 5 not ar-
34—Proc. Brow. Soc. WAsH., VoL. 87, 1974 (389 )
390 Proceedings of the Biological Society of Washington
0.58mm
0.164m™m
Fics. 1-8. Pseudopolydora gibbsi, new species. 1, anterior end, dorsal
view, palpi removed; 2, posterior end, dorsal view; 3, major spine from
posterior row of setiger 5; 4, major spine from anterior row of setiger 5;
5, major spine from anterior row of setiger 5, broken example; 6, anterior
end, dorso-lateral view, showing arrangement of modified spines of
setiger 5; 7, posterior notopodial hook; 8, neuropodial hooded crotchet.
ranged in double U-shaped row; branchiae from setiger 6, 10-14 pairs;
pygidium much reduced, consisting of pair of small, ventral lappets.
Description: Largest specimen (holotype) 9 mm long by 0.5 mm
wide at broadest point (setiger 5), but most examples only half this
size.
Prostomium wide, anteriorly entire, with 2 pairs of eyes arranged
A new polychaete from Solomon Islands 391
trapezoidally, anterior pair twice as far apart as posterior pair; caruncle
extending to middle of setiger 2, occasionally reaching anterior border
of setiger 3 (Fig. 1). No trace of occipital or nuchal cirri. First setiger
completely suppressed dorsally and ventrally by prostomial caruncle
and peristomium, respectively. Well developed, extrusible proboscis
present.
Body attenuated over anterior and posterior regions, attaining greatest
width over median somites, beginning approximately with setiger 5.
Notopodial postsetal lamellae of typical configuration, well formed over
first 2-4 setigers, or may be much reduced. Neuropodial postsetal
lamellae prominent on setigers 1 and 2, often reduced on 3 and absent
on 4. Minute notosetal fascicle present on setiger 1. All setae, except
modified spines of setiger 5, neuropodial crotchets and _ specialized
posterior notopodial hooks, consisting of simple, alimbate, smooth, long
capillaries numbering about 6 per fascicle.
Setiger 5 with 2 rows of modified major spines forming gentle arc
(Fig. 6); anterior (ventral) row consisting of 5-6 stout, spathe-like,
distally acuminate setae (Fig. 4), these spines often broken or worn
(Fig. 5); posterior (dorsal) row consisting of 3-4 heavy falcigers with
tip at right angles to main axis of shaft (Fig. 3). Both dorsal and
ventral fascicles of capillary setae also present.
Hooded neuropodial crotchets commencing on setiger 8, 6-8 per
ramus, with well developed manubrium (Fig. 8). Posterior 15-17
notopodia bearing 1 or 2 stout, sickle-shaped hooks on each side, to-
gether with 2-3 long capillaries (Figs. 2, 7).
Branchiae beginning on setiger 6, generally fully developed from
that segment, typically 10-14 pairs, this arrangement not always sym-
metrical, there often being more on one side than other. Pygidium
quite reduced, consisting merely of pair of ventral lappets (Fig. 2).
Color in alcohol: Fleshy white, occasionally with few very faint
flecks of dark pigment on dorsum of anterior segments.
Type material: Holotype (ZB. 1970.880) and 26 paratypes are de-
posited in Porifera-Polychaeta Section of British Museum (Natural
History ).
Type-locality: Mouth of Lunga River, Guadalcanal Island (9° 24’
30” S, 160° 01’ 30” E), from interstices of sponge, in association with
Mercierella enigmatica.
Remarks: Pseudopolydora gibbsi resembles P. antennata (Claparéde,
1870) in the configuration of the anterior modified setae on setiger 5.
It also resembles P. reishi Woodwick, 1964, in that the modified major
spine series does not form a U- or J-shaped double row, as generally
seen in species of Pseudopolydora. P. gibbsi may be differentiated from
P. antennata by the absence of an occipital cirrus and in exhibiting an
entire prostomium. It is distinguished from P. reishi and P. antennata
in the absence of a pygidial funnel. P. pigmentata Woodwick, 1964,
also lacks an occipital cirrus, but there is only one pair of eyes, no noto-
392 Proceedings of the Biological Society of Washington
setae on setiger 1 and the anterior row of modified setae on setiger 5
consists of limbate capillaries.
The hooded neuropodial crotchets exhibit an overall configuration
intermediate between that of Pseudopolydora and the genus Polydora
Bosc; the apical tooth is less closely imbricated over the main fang than
in most species of Pseudopolydora but more so than in most species of
Polydora, with both forming an angle of 90° with the main shaft.
Their commencement on setiger 8 is consistent with Pseudopolydora.
The presence of only a few simple capillaries differs from the dense,
3-tiered fascicles of heteromorphic setae generally seen in species of
Pseudopolydora, such as P. kempi (Southern, 1921) and P. pauci-
branchiata (Okuda, 1937; 232, 234, Fig. 12b; see also Imajima and
Hartman, 1964: 287; Light, 1969: 544).
The posterior notopodial hooks are unique for species of this genus.
Their close resemblance to similar hooks in Boccardia hamata ( Webster),
Blake, 1966, as well as to those of the Polydora hoplura-colonia com-
plex (see Light, 1969: 540), indicates that such specialization of the
posterior notosetae must be incipient in the entire polydorid line,
having developed independently several times. Similarly, the tendency
towards reduction of the pygidium into small lappets is also seen in all
three genera.
Pseudopolydora gibbsi appears to be quite specialized by virtue of
the development of the major spines in the anterior row of setiger 5,
the reduced number of crotchets in each ramus, the absence of an oc-
cipital cirrus and the presence of highly modified posterior notopodial
hooks. The only other species of Pseudopolydora known to possess mod-
ified posterior notosetae is P. corallicola Woodwick, 1964, from Eniwetok
Island. In this form the posterior notosetae are awl-shaped and grouped
together in conical bundles, a condition very similar to that seen in the
Polydora armata-caulleryi complex (see Light, 1969: 542).
LITERATURE CITED
BLAKE, J. A. 1966. On Boccardia hamata (Webster), new combina-
tion (Polychaeta, Spionidae). Bull. S. Calif. Acad. Sci. 65(3):
176218410 fie
CLAPAREDE, EF. 1870. Les Annélides Chétopodes du Golfe de Naples.
Mem. Soc. Phys. Hist. Nat. Genéve 20 (pt. 1):1-225, 31 pls.
Gipss, P. EF. 1971. The polychaete fauna of the Solomon Islands. Bull.
British Mus. (Nat. Hist.), Zool. 21(5):99-211, 17 figs.
ImajiMA, M., ano O. Hartman. 1964. The polychaetous annelids of
Japan. Part II. Allan Hancock Foundation Publ., Occ. Pap. 26:
239-452, pls. 36-38.
Licut, W. J. 1969. Polydora narica, new species, and Pseudopolydora
kempi californica, new subspecies, two new spionids ( Annelida:
Polychaeta) from central California. Proc. Calif. Acad. Sci.
36(18) :531-550, 13 figs., 2 tables.
A new polychaete from Solomon Islands 393
OxupA, S. 1937. Spionoform polychaetes from Japan. Jour. Fac. Sci.
Hokkaido Imp. Univ., ser. VI, Zool. 5:217—254, 27 figs.
SOUTHERN, R. 1921. Fauna of the Chilka Lake. Polychaeta of the
Chilka Lake and also of fresh and brackish waters in other
parts of India. Mem. Indian Mus. 5:563-659, 13 pls., 18
textfigs.
Woopwickx, K. H. 1964. Polydora and related genera (Annelida,
Polychaeta) from Eniwetok, Majuro, and Bikini Atolls, Marshall
Islands. Pacific Sci. 18:146-159, 28 figs., 3 tables.
394 Proceedings of the Biological Society of Washington
=
my, OG 75
Vol. 87, No. 35, pp. 395-404 31 December 1974
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
COMMENTS ON THE TAXONOMY OF THE WEST
AFRICAN TATERILLUS (RODENTIA: CRICETIDAE)
WITH THE DESCRIPTION OF A NEW SPECIES
By C. Brian Rossins
Division of Mammals, National Museum of Natural
History, Washington, D.C. 20560
The taxonomy of West African rodents of the genus Tater-
illus is confused. Allen (1939) and Ellerman (1941) listed
four taxa: T. gracilis gracilis (Thomas, 1892), T. g. angelus
Thomas and Hinton, 1920, T. nigeriae Thomas, 1911, and T.
lacustris (Thomas and Wroughton, 1907). Rosevear (1969)
recognized two species, T. gracilis and T. nigeriae, allocating
T. lacustris as a subspecies of T. gracilis. Taterillus pygargus
(F. Cuvier, 1838), long considered a species of Gerbillus oc-
curing in Egypt and the Sudan, has been shown by Petter
(1952) to be a species of Taterillus. The holotype of T.
pygargus is from Senegal. This taxon was overlooked by
Rosevear (1969).
Even though Ellerman (1941) and Rosevear (1969) recog-
nized more than one species of Taterillus, they suggested that
the West African forms could be a single polymorphic species.
Cytogenetic data and additional study specimens demonstrate
the presence of at least four distinct species. Near topotypes
of T. gracilis from Senegal have a karyotype of 2N = 36/37,
FN = 44 (Matthey and Jotterand, 1972; Petter et al., 1972).
Chromosomal information is also known on Taterillus from
Upper Volta (Matthey and Petter, 1970; Matthey and _ Jot-
terand, 1972), Ivory Coast (Petter, pers. comm.), and Ghana
(Robbins unpublished data). These specimens have a karyo-
type of 2N = 36/37, FN = 42, and have been identified as
35—Proc. Bio. Soc. WaAsH., VOL. 87, 1974 (395 )
396 Proceedings of the Biological Society of Washington
T. gracilis. Chromosomal information has been reported from
near topotypes of T. pygargus (2N = 22/23, FN = 37-40)
from Senegal (Matthey and Jotterand, 1972; Petter e¢ al.,
1972). Specimens from the Cameroon with a karyotype of
2N = 28, FN = 44 have been referred to T. lacustris (Tranier
et al., 1974). A Taterillus with a karyotype of 2N = 30, FN =
36, reported as T. nigeriae has been found in Mauritania
(Matthey, 1969).
Petter (1972) revised all of the species and subspecies of
Taterillus utilizing chromosomal information. Based on a
greater number of specimens and additional karyotypic in-
formation, I have come to different conclusions regarding the
taxonomy of West African Taterillus. The large number of
West African specimens of Taterillus now available are a
result of the Smithsonian Institutions’ African Mammal Pro-
ject under the direction of Dr. H. W. Setzer. Among these
are 115 specimens from Mauritania where Taterillus was pre-
viously known only by the single specimen reported by Mat-
they (1969) and Petter (1970). The purpose of this report
is to clarify the identities of the West African Taterillus.
MATERIALS AND METHODS
The specimens used in this study, except for a single indi-
vidual from the Museum National d'Histoire Naturelle, Paris
(MNHP), which was reported as a 2N=30 karotype by
Matthey (1969), are housed in the National Museum of Nat-
ural History, Division of Mammals (USNM), and were all
prepared as conventional museum study skins. Twenty-four
measurements as applied to Taterillus by Robbins (1973),
were made of various dimensions of the skull. Measurements
are in millimeters, weights in grams, and capitalized color
terms are those of Ridgway (1912).
Computer analyses were performed through the Information
Systems Division, Smithsonian Institution. The Smithsonian-
developed DSTAT univariate analysis program yielded stan-
dard statistics (mean, standard error, standard deviation, vari-
ance, coefficient of variation, and range), as well as Student's
“t’ test, F-test for homogeneity of variance, and one-way
analysis of variance. The BMDO7M (Dixon, 1973) multivar-
Comments on West African Taterillus 397
iate program performed the step-wise discriminant function
analysis. Results from multivariate analysis are shown in Table
1. A scattergram of the first two canonical variates and char-
acter vectors are presented in Figure 2. Vectors were deter-
mined by multiplying the pooled within-groups standard
deviation by the coefficients for canonical variates one and
two for the most useful characters (Power and Tamsitt, 1973).
An analysis of the West African Taterillus, which includes a
reexamination of the Mauritanian specimen discussed by
Matthey (1969) and Petter (1970), reveals the presence of a
heretofore undescribed species that may be known as:
Taterillus arenarius, new species
Holotype: Adult male, skin and skull, USNM 401919, from Tiguent,
Trarza Region, Mauritania; obtained 9 April, 1967, by C. B. Robbins,
original number 799.
Etymology: The name arenarius refers to the sandy substrate char-
acterizing the areas where the species occurs.
Distribution: Mauritania, Niger, and presumably Mali (Fig. 1).
Definition; Interauricular, interorbital, and rostral areas same color
as dorsum and varying from Sayal Brown to Snuff Brown; circumorbital
region, postauricular patches, mystacial and pectoral areas, fore and
hind limbs, and entire underparts white; cheeks and sides Cinnamon-
Buff; dorsal hairs plumbeous basally, the brown pigmented portion
only 2 to 3 mm in length, and some hairs finely tipped with dark brown;
fore and hind limbs have five digits with claws, plantar surfaces naked
except for a narrow band of white hairs; pinna of ear long and almost
naked, color almost the same as the dorsum, and anterior margin with
short buff colored hairs; vibrissae long and composed of both white
and dark brown hairs; tail long and uniformly Cinnamon-Buff basally
with the dorsal hairs interspersed with darker brown hairs grading to a
terminal Mummy Brown pencil; ventrally, the Cinnamon-Buff color
grades to white in the region of the pencil.
Skull relatively large for the genus and moderately robust; zygomata
heavy; lachrymals large; molariform teeth medium-sized; auditory bul-
lae large and inflated; parapterygoid fossae large and deep but not
markedly flared; rostrum relatively long and slightly expanded anterior
to the infraorbital shield; nasals relatively broad and long; braincase
flattened.
Measurements: Measurements of the holotype (age class 5; see Rob-
bins, 1973) followed by averages and extremes of eight adults (age class
4) from the type-locality are, respectively: total length 289, 279 (269-
300); head and body length 128, 117 (111-124); length of tail 161,
163 (158-176); length of hind foot 33, 33 (32-34); length of ear from
398 Proceedings of the Biological Society of Washington
Hed ‘a i i
MALI
Sst SSG
an ae
Sessten |
Sei ea
Fa,
GUINEA
Fic. 1. Distribution of T. arenarius with known localities shown by
closed circles. Type-localities for the other species and subspecies of
West African Taterillus are: T. pygargus, closed triangle; T. g. gracilis,
open triangle; T. g. angelus, x; T. g. nigeriae, open circle with dot en-
closed; T. lacustris, open circle.
notch 22, 21 (19-22); weight 59, 46 (42-52); greatest length of skull
(GLS) 36.9, 35.6 (34.9-36.3); occipitonasal length 35.7, 34.3 (33.4—
35.3); basilar length 26.9, 25.7 (25.2-26.5); zygomatic breadth 18.2, 17.6
(17.4-17.8); condylobasilar length 28.8, 27.5 (27.1-28.2); cranial
breadth 14.6, 14.4 (13.6-14.7); least interorbital constriction 6.0, 6.1
(5.8-6.6); breadth of rostrum 4.5, 4.6 (4.4—4.8); greatest breadth across
bullae (GBAB) 13.9, 13.4 (12.8-13.8); greatest width across upper
molars 7.3, 7.1 (6.7-7.3); length of diastema 9.0, 8.9 (8.6-9.6); palatilar
length 15.3, 15.0 (14.6-15.3); postpalatal length (PPAL) 11.3, 10.6
(10.0-11.0); length of anterior palatine foramina 6.2, 5.9 (5.5-6.5);
length of palatal bridge 7.4, 7.3 (7.0-7.7); length of auditory bulla
(LAB) 10.1, 9.4 (9.0-9.7); breadth of auditory bulla (BAB) 6.2, 5.8
(5.6-6.1); length of nasals 15.0, 14.8 (14.1—15.3); length of rostrum 13.4,
12.9 (12.7-13.6); frontonasal length 28.2, 27.2 (26.4-28.3); depth of
cranium 13.7, 13.6 (13.0-13.8); length of posterior palatine foramina
(LPPF) 3.8, 3.7 (3.2-4.2); alveolar length of first molar (ALM1) 2.8,
2.7 (2.4-3.1); alveolar length of maxillary toothrow 5.1, 5.1 (5.0-5.4).
Comparisons: Taterillus arenarius can be distinguished from T.
pygargus, where the two occur together, by its markedly longer and
more robust skull; flatter, less rounded braincase; broader rostrum and
slightly larger incisors; more bulbous bullae; slightly smaller cheek
Comments on West African Taterillus 399
teeth; parapterygoid fossae larger (deeper) but not so widely flaring;
and paler dorsal coloration. At Garak, Mauritania, where T. arenarius
is sympatric with T. pygargus, the latter are darker dorsally and lat-
erally—Snuff Brown and Tawny-Olive, respectively.
Taterillus arenarius differs from T. gracilis by its broader, more ro-
bust skull although the skulls are similar in length. The bullae of T.
arenarius are more inflated anteriorly, antero-laterally, and ventrally.
The pelage color of T. arenarius is conspicuously paler.
Taterillus arenarius closely resembles T. lacustris, from which it can
be distinguished by its somewhat narrower nasals, flatter braincase, and
more inflated bullae. The pelage color is similar.
Remarks: A specimen of Taterillus from Bou Rjeimat, Mauritania,
has a reported karyotype of 2N = 30, FN = 36 (Matthey, 1969) and
was identified by Matthey (1969), Petter (1970), and Petter (1972)
as T. nigeriae. Inasmuch as the karyotype of T. nigeriae was not known,
and since its geographic range is closer to Mauritania than any other
known species of Taterillus, this allocation was made. However, an
examination of available specimens of T. nigeriae, including the holo-
type, demonstrates that T. arenarius is distinct. Taterillus nigeriae is
darker in dorsal coloration and occupies a different habitat.
Taterillus arenarius is restricted in distribution to the vegetated sandy
habitats of the northern Sahel Savanna and Sub-Desert of West Africa.
It is replaced southward by either T. pygargus or T. gracilis. Where
sympatric with T. pygargus, this species occupies the sandier habitats
while T. pygargus is found in areas with a harder substrate vegetated
with tall grasses. Eastward, T. arenarius is replaced by T. lacustris.
Other rodents occurring with T. arenarius are: Gerbillus (Gerbillus)
gerbillus, G. (G.) pyramidum, G. (G.) nigeriae, G. (Hendecapleura)
mauritaniae, G. (H.) amoenus, Tatera guineae, T. kempi, T. gambiana,
T. wellmani, Desmodilliscus braueri, Jaculus jaculus, J. deserti, Arvi-
canthis niloticus, and Euxerus erythropus.
Specimens examined: Mauritania (97). Inchiri Region: Bou Rjeimat,
1 (MNHP 247); Trarza Region: 11 km N Nouakchott, 5 (USNM
411120-411124); 6 km E Nouakchott, 3 (USNM 401117-401119);
Tiguent, 48 (USNM 401893-401939); Garak, 10 (USNM 401940;
401941; 401950-401956; 401958); Brakna Region: 3 km S Aleg, 14
(USNM 401960-401973); Gorgol Region: Kaedi, 10 (USNM 401974-—
401983); Guidimaka Region: Passe de Soufa, 6 (USNM 401989;
401994401996; 401998; 401999). Niger (15). Agadez Region: 5 km
NE Agadez, 7 (USNM 482626-482632); 30 km S In-Gall, 3 (USNM
482633482635 ); Tahoua Region: 120 km S In-Gall, 5 (USNM 482636-
482640).
Gazetteer: (Localities plotted in Figure 1)
1. Aleg,3kmS 17°02’N., 13°55’W.
2. Bou Rjeimat 19°04’N., 15°08’W.
3. Garak 16°33’N., 15°46’W.
400 Proceedings of the Biological Society of Washington
TABLE 1. Discriminant coefficients for characters most useful in separat-
ing three West African species of Taterillus. For explanation of abbrevia-
tions see text and Fig. 2
Character A vs. P A vs. G P vs. G
BAB 9.287 11.552 2.265
GBAB — 6.197 — 0.334 5.863
PPAL —11.921 —20.116 — 8.195
GLS 4.758 — 7.975 —12.715
ALMI1 14.857 22.794 8.237
LAB — 6.624 — 2.408 4.216
LPPF —14.407 —13.813 0.594
4. Kaedi 16°09’N., 13°30’W.
5. Nouakchott, 6 km E 18°09’N., 15°58’W.
6. Nouakchott, 11 km N 18°13’N., 16°01’W.
7. Passe de Soufa 15°56’N., 12°00’W.
8. Tiguent 17°16’N., 16°01’W.
9. Agadez, 5 km NE 17°02’N., 08°02’E.
10. In-Gall, 30 km S 16°33’N., 06°52’E.
11. In-Gall, 120 km S 15°45’N., 06°36’E.
Analysis of Variation and Diagnostic characteristics of the Species:
Table 1 and Figure 2 show the results of the discriminant analysis on
three species of Taterillus. The specimens of T. gracilis and T. pygargus
were age class 4 animals (see Robbins, 1973) of known karyotype rep-
resenting several localities in Senegal. The specimens of T. arenarius
were also age class 4 from several Mauritania localities, but most were
from the type-locality.
The mean and standard error for seven cranial characters in their
order of inclusion in the discriminant analysis are shown in Table 2.
These seven cranial characters were selected as the minimum necessary
for discriminating the three species because of sample size. In addition,
by step seven, the F-matrix in the discriminant analysis showed sig-
nificant differences between the three groups at the 95 per cent level.
It should be noted that three of the cranial characters in the dis-
criminant analysis reflect auditory bulla dimensions. Taterillus arenarius
and T. pygargus have bullar shapes which are more similar to each
other than either are to T. gracilis. In areas where T. arenarius and
T. gracilis are sympatric (as well as areas where T. pygargus and
T. gracilis occur together), comparisons of bullar shape and size are
the most reliable method of distinguishing between the species. The
length and direction of the cranial character vectors (Fig. 2) show the
relative importance of each variable in discriminating between the three
species. When a clean, intact skull is available, specimens of unknown
identity can be correctly allocated.
Comments on West African Taterillus 401
TABLE 2. Means and standard error for seven cranial characters in their
order of inclusion in the discriminant analysis. For explanation of ab-
breviations see text.
T. arenarius (n= 15) T. pygargus (n=9) T. gracilis (n= 7)
Standard Standard Standard
Character Mean Error Mean Error Mean Error
1. BAB 5.70 0.084 5.00 0.087 5.30 0.117
2. GBAB 13.40 0.109 13.80 0.137 13.30 0.157
3. PPAL 10.50 0.149 10.50 0.070 10.90 0.146
4, GLS 35.40 0.190 35.00 0.102 35.30 0.261
5. ALM1 2.66 0.081 2.83 0.041 2.77 0.047
6. LAB 9.40 0.082 9.30 0.087 9.40 0.084
7. LPPF 3.60 0.121 3.60 0.095 3.50 0.094
The chromosomes of T. arenarius are only known from the female
reported under the name T. nigeriae by Matthey (1969). Its karyotype
(as reported by Matthey, 1969) consists of one pair of large sub-
telocentrics, three pairs of medium-sized to small metacentrics, and ten
pairs of medium-sized acrccentric autosomes. The X-chromosomes are
large submetacentrics. The Y-chromosome is unknown.
w
£ T. GRACILIS
nN
CANONICAL VARIATE 2 (19.3%)
Oo —
!
=
-2
| 3 2) -| 0 2 3 4
CANONICAL VARIATE 1 (80.7%)
Fic. 2. Projection of the first two canonical variates for three species
of West African Taterillus. Vectors show relative contributions of the
characters in the discriminant analysis. For explanation of abbreviations
see text.
402 Proceedings of the Biological Society of Washington
On gross morphology, the four West African species of Taterillus
appear to be closely related. However, their different karyotypes do
not affirm a common derivation. The 2N = 28, FN = 44 karyotype of
T. lacustris could have been easily derived from the 2N = 36, FN = 44
karyotype of T. gracilis through four fusions. However, serum protein
analysis (Tranier et al., 1974) shows that T. lacustris is more closely
related to T. pygargus than to T. gracilis. The 2N = 30, FN = 36
karyotype of T. arenarius and the 2N = 22, FN=40 karyotype of
T. pygargus could also have been derived from the T. gracilis karyotype,
but would have required a complex series of translocations and in-
versions. The 2N = 22, FN = 40 karyotype of T. pygargus could have
been derived from the 2N = 30, FN =36 karyotype of T. arenarius
through two inversions and four fusions. Speculation on the relationships
of these four species based on their karyotypes is premature at this time,
and should await additional information from banding pattern analysis.
ACKNOWLEDGMENTS
Special thanks must go to Henry W. Setzer for allowing me to study
the Taterillus collection at the Smithsonian, for the opportunity to collect
in West Africa, for computer time, and for commenting on the manu-
script. Francis Petter (MNHP) kindly provided specimens and karyo-
type information. Dan Piecesi and Charles Roberts of the Smithsonian
Information Systems Division helped with the computer programs. Don
E. Wilson and Michael A. Bogan tutored me on multivariate statistics
and read the manuscript. Alfred L. Gardner critically reviewed the
manuscript and offered numerous helpful suggestions. My wife Norrie
provided the illustrations. This paper is based on specimens collected
under the U.S. Army Medical Research and Development Command
Contract No. DA-49-193-MD-2738, and the research was partially funded
by National Science Foundation Grant GB 35143 to the author.
LITERATURE CITED
ALLEN, G. M. 1939. A checklist of African mammals. Bull. Mus.
Comp. Zool. 83:1-763.
Dixon, W. J. (Ed.) 1973. BMD biomedical computer programs.
Univ. Calif. Press, Berkeley. vii + 7783p.
ELLERMAN, J. R. 1941. The families and genera of living rodents.
British Mus. (Nat. Hist.), London. 2:xii + 690p.
MatrHey, R. 1969. Chromosomes de Gerbillinae. Genres Tatera et
Taterillus. Mammalia 33:522-528.
, AND F. Petter. 1970. Etude cytogenetique et taxonomique
de 40 Tatera et Taterillus provenant de Haute-Volta et de
Republique Centrafricaine (Rongeurs, Gerbillidae). Mammalia
34:585—597.
, AND M. JoTrrerRAND. 1972. L’analyse du caryotype permet de
reconnaitre deux especes cryptiques confondues sous le nom
Comments on West African Taterillus 403
de Taterillus gracilis Th. (Rongeurs—Gerbillidae). Mammalia
36: 193-209.
Petrer, F. 1952. Note sur un type de F. Cuvier:Gerbillus pygargus,
“la Gerbille du Senegal”. Mammalia 16:37-39.
——. 1970. Capture d’un Taterillus en Mauritanie. Mammalia 34:
325-326.
——. 1972. Preliminary identification manual for African mammals
17. Rodentia:Gerbillidae (excluding the genera Tatera and
Gerbillurus). J. Meester Ed., Smithsonian Institution, Wash.
D. C. 23p.
—., A. Pou.et, B. Husert, anp F. Apam. 1972. Contribution a
Vetude des Taterillus du Senegal, T. pygargus (F. Cuvier,
1832) et T. gracilis (Thomas, 1892) (Rongeurs, Gerbillides ).
Mammalia 36:210-213.
Power, D. M., AND J. R. Tamsirr. 1973. Variation in Phyllosomus
discolor (Chiroptera:Phyllostomatidae). Canadian Jour. Zool.
51:461-468.
Rmeway, R. 1912. Color standards and color nomenclature. Published
by the author. iii + 43p., 53 pl.
Rozssins, C. B. 1973. Nongeographic variation in Taterillus gracilis
(Thomas) (Rodentia:Cricetidae). Jour. Mamm. 54:222-238.
RosEvEsaR, D. R. 1969. The rodents of West Africa. British Mus.
(Nat. Hist.), London. xii + 604p.
TRANIER, M., B. HuBErT, AND F. Perrer. 1974. Taterillus de Youest
de Tchad et du nord du Cameroun (Rongeurs, Gerbillides).
Mammalia 37:637-641.
404 Proceedings of the Biological Society of Washington
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Vol. 87, No. 36, pp. 405—410 31 December 1974
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
PSEUDOPHALLUS BRASILIENSIS (PISCES:
SYNGNATHIDAE), A NEW FRESHWATER
PIPEFISH FROM BRAZIL
By C. E. Dawson
Gulf Coast Research Laboratory Museum
Ocean Springs, Mississippi 39564
I recently received eight specimens of an undescribed syng-
nathine tailpouch pipefish of the genus Pseudophallus Herald
(1940) from the Rio Tocantins, a major tributary of the lower
Rio Amazonas, Brazil. Although the genus is treated in a
manuscript being prepared for the Sears Foundation series
“Fishes of the Western North Atlantic,” there will be con-
siderable delay in completion of that work. Description of
the new species is, therefore, given at this time in order to
make the name available.
Type-material has been deposited in collections of the
Museu de Zoologia da Universidade de Sao Paulo (MZUSP),
National Museum of Natural History, Smithsonian Institution
(USNM) and Gulf Coast Research Laboratory Museum
(GCRL). Measurements (mm) or proportions referred to
standard length (SL) or head length (HL) are given for the
holotype followed, in parentheses, by the range for the re-
mainder of the type-series; counts from the holotype are
marked with an asterisk (*). Coloration is described from
alcohol preserved specimens. Data used in species compari-
sons are from my observations on almost all specimens of
Pseudophallus now available in Central and North American
collections. Lists of these examined materials are omitted
here, but will be included in the forthcoming general treat-
ment of the genus mentioned above.
36—Proc. Bion. Soc. WAsuH., Vou. 87, 1974 (405 )
406 Proceedings of the Biological Society of Washington
I thank the curators of the various institutions that have
loaned comparative material. Special acknowledgment is due
Dr. Naércio A. Menezes (MZUSP) for providing the speci-
mens of the new species which were obtained by the MZUSP
Expedicao Permanente da Amazonia. Drawings are by Anne
Langenfeld (GCRL). This work was in part supported by
National Science Foundation Grant GB-31053X.
Pseudophallus brasiliensis, new species
Figure 1
Holotype: MZUSP 10278 (85 mm SL, male); Brazil, Para, Rio
Tocantins, Igarapé Ind, Faro de Panaquera; 01°52’S, 49°10’W; 1 Sept.
1970.
Allotype: USNM 212058 (70, female); taken with holotype.
Paratypes: MZUSP 10627 (1 specimen, 43 mm SL); taken with holo-
type. MZUSP 10279 (2, 60-63), GCRL 12755 (2, 56-66); Brazil,
Para, Rio Tocantins, Igarapé Mapara, Parana, Samuuma; 02°05’S,
49°20’W; 5 Sept. 1970. MZUSP 10280 (1, 54); Brazil, Para, Rio
Tocantins, Igarapé Acicuré at Cameta; 02°15’S, 49°30’W; 7 Sept. 1970.
Diagnosis: Dorsal fin rays 28 to 31, located on 6.75 to 7.25 tail
rings; trunk rings 13; tail rings 31 to 32; median head ridges, opercular
and pectoral cover plate ridges, trunk and tail ridges somewhat ‘elevated,
distinct; length of dorsal fin in head 0.9 to 1.0; snout without a longi-
tudinal brownish stripe.
Counts and measurements: Dorsal fin rays 28(in 1), 29*(5), 30(1),
31(1); pectoral fin rays 13(4), 14*(8), 15(4); caudal fin rays 10(8),
trunk rings 13(8); tail rings 31(3), 32*(5); trunk rings covered by
dorsal fin 0(8); tail rings covered by dorsal fin 6.75*(3), 7.0(3),
7.25 (2). Head length 9.5 (5.8-8.1); snout length 3.4 (2.3-3.1); least
breadth of snout 0.9 (0.6—0.9); pectoral fin length 1.8 (1.0-1.3); caudal
fin length 3.5 (2.3-2.9); length of dorsal fin base 10.1 (5.8-9.1). Head
length in SL 8.9 (7.4-8.6); snout length in HL 2.8 (2.5-2.7); dorsal
fin base in HL 0.9 (0.9-1.0).
Description: Median snout ridge smooth, somewhat elevated, extends
from rear of articulation of upper jaw to about anterior third of inter-
orbital (Fig. 2); supraorbital, frontal, nuchal and prenuchal ridges low
but distinctly elevated and prominent in all specimens; opercular ridge
prominent, complete in 6 specimens (extends across 34 of opercle in
remainder) margined above and below by radiating striae; pectoral
cover plate with a distinct anteroventrally directed ridge margined by
radiating striae; trunk and tail ridges well defined, their edges distinctly
granular or scalloped under 30 magnification; width of intermedial
scutellar plates on trunk and anterior portion of tail equals about a
third the distance between proximal margins of adjacent scutellae. Body
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407
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A new freshwater pipefish from Bra
408 Proceedings of the Biological Society of Washington
Fic. 3. Delineation of posterior trunk and anterior tail rings of Pseudo-
phallus brasiliensis, illustrating ridge pattern, dorsal fin location and anal
papilla of USNM 212058; allotype.
ridge pattern typical of Pseudophallus, i.e., lateral trunk ridge continuous
across anal ring, uniting with superior tail ridge near right angle from
posterior end of dorsal fin, usually with some irregular degree of de-
flection at or near anal ring (Fig. 3). Lateral line neuromasts indis-
tinct. Dorsal fin originates at middle of first tail ring; anal fin absent;
abdomen somewhat V-shaped in both sexes. Anal papilla of mature
females much enlarged (Fig. 3), apparently smooth over its distal
third, the middle portion distinctly papillose under 30 magnification;
anal papilla measured 1.9 mm (from posterior basal angle to tip) in
allotype, 2.4 mm in 66 mm fish (GCRL 12755). Brood pouch of holo-
type covers 13 tail rings and contains eggs through 12 rings; maximum
diameter of 5 measured eggs with eyed larvae was 1.1 mm.
Ground color tan to brown, markings brown to near black, generally
darkest on venter. Markings highly variable, generally distinctly blotched
and mottled with brown (Fig. 1), most frequently with up to 13 ir-
regularly spaced, narrow, pale bands on trunk and tail; head irregularly
blotched, snout without a lateral stripe; iris crossed by up to 10 brown
bars somewhat narrower than interspaces; dorsal and pectoral fin rays
irregularly edged with brown melanophores; caudal fin dark brown with
narrow pale margin. Proximal half of anal papilla of mature females
heavily pigmented with brown melanophores, distal portion pale to
nearly white with few flecks of brown.
Etymology: Named brasiliensis, in reference to the Brazilian type-
locality.
Comparisons: Principal features differentiating Pseudophallus brasili-
ensis are given in the key below.
Discussion: The amphi-American genus Pseudophallus now includes
four closely related forms that appear to be largely restricted to fluvial
409
A new freshwater pipefish from Brazil
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410 Proceedings of the Biological Society of Washington
habitats. All exhibit considerable overlap in meristic or morphometric
characters but they appear to represent distinct taxa. Meristic features,
such as frequencies of dorsal fin rays or total body rings (Table 1 and
2), alone do not permit identification of single specimens, but little dif-
ficulty is encountered when multicharacter comparisons are employed.
The late Earl S. Herald (unfinished Ms.) treated the Atlantic P. mindii
as a subspecies of the Pacific P. starksi and retained species rank for
the other Pacific form, P. elcapitanensis. He was, however, unaware of
the new Brazilian form and had not seen much of the material examined
by me. My data indicate a modal shift from 40 to 39 dorsal fin rays
between Mexican and Costa Rican-Panamanian populations of P. starksi,
but there is no evidence of clinal variation in the other species. The
Pacific forms are sympatric in Costa Rica and Panama and sympatric
populations of P. mindii and P. brasiliensis may occur in Brazil. Pending
collection of more extensive series from Atlantic drainages, I retain
species rank for each of the four nominal forms of Pseudophallus.
Provisional key to the genus Pseudophallus
I Drunk rings modally 145(99%)- Raciticespecies »)
Trunk rings modally 13 (78%); Atlantic species _....-___. 3
2. Dorsal fin rays 37 to 44 (x = 40); dorsal fin covers 9.0 to 11.0
(x = 9.7) rings; trunk rings 13 to 15 (x = 14); usually with a
distinct, broad, brown stripe on snout and short ventrally directed
brown bar or blotch behind eye; Baja California Sur, México
tOSE Cua or tesa uremic eel ne P. starksi (Jordan and Culver )
Dorsal fin rays 30 to 35 (x = 33); dorsal fin covers 6.75 to 7.50 (x
= 7.1) rings; trunk rings 14; without postorbital bar or stripe on
snouts ‘Costa Rica and) Ramana) ee
Reig aie nnots SPE IN a P. elcapitanensis (Meek and Hildebrand)
3. Dorsal fin rays 33 to 41 (x = 36); dorsal fin covers 7.50 to 10.25
(x = 8.7) rings; length of dorsal fin base in head 0.6 to 1.0
(x = 0.7); rings total 46 to 51 (x= 48); head ridges usually
indistinct or obsolete in adults; usually with postorbital blotch
and stripe on snout; Cuba, Jamaica, Guatemala to Panama, Vene-
mole, smael Jeimavaill P. mindii (Meek and Hildebrand )
Dorsal fin rays 28 to 31 (x = 29); dorsal fin covers 6.75 to 7.25
(x= 7.0) rings; length of dorsal fin base in head 0.9 to 1.0
(x = 1.0); rings total 44 to 45 (x = 45); head ridges prominent
in adults; without postorbital bar or stripe on snout; Rio To-
Cantins: (Brazile ae eee P. brasiliensis new species
LITERATURE CITED
HeErAtp, E. S. 1940. A key to the pipefishes of the Pacific American
coasts with descriptions of new genera and species. Univ. S.
Calif., Allan Hancock Pac. Exped. 9(3):51-64.
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Vol. 87, No. 37, pp. 411-438 31 December 1974
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
CYCLOPOID COPEPODS (LICHOMOLGIDAE) FROM
GORGONACEANS IN MADAGASCAR
By ArrHur G. HuMEs
Boston University Marine Program,
Marine Biological Laboratory,
Woods Hole, Massachusetts 02543
Lichomolgid copepods associated with Gorgonacea have
been reported from the West Indies (Humes and Stock, 1973),
Bermuda (Humes, 1973a), and the Mediterranean coast of
France and Spain (Stock and Kleeton, 1963). In addition three
lichomolgid copepods are known to be associated with Gor-
gonacea in Madagascar. These are Acanthomolgus astrictus
Humes and Stock, 1973, from Acanthogorgia aspera Pourtalés,
Acanthomolgus hales Humes and Stock, 1973, from Soleno-
caulon tortuosum Gray, and Metaxymolgus cinctus Humes
and Stock, 1973, from Psammogorgia ramosa Kikenthal.
This paper includes the descriptions of three new species of
Acanthomolgus and one new species of Paramolgus, along
with new records for certain species of Acanthomolgus, Meta-
xymolgus, and Telestacicola. All these were collected by the
author from shallow-water gorgonaceans in the vicinity of
Nosy Bé in northwestern Madagascar. The specimens were
recovered from the sediment obtained after washing the hosts
in approximately 5 per cent ethyl alcohol in sea water.
The field work in 1963-64 was carried out while the author
was chief scientist in Madagascar for the U. S. Program in
Biology of the International Indian Ocean Expedition. The
work in 1967 was supported by a grant (GB-5838) from the
National Science Foundation. The study of the material has
been aided by another grant (GB-8381X) also from the Na-
37—Proc. Broun. Soc. WAsH., Vou. 87, 1974 (411)
412 Proceedings of the Biological Society of Washington
tional Science Foundation. I wish to acknowledge with many
thanks the generous assistance given by the staff of the Centre
O.R.S.T.O.M. de Nosy Bé, where the field work was done.
I am greatly indebted to Dr. J. Vervoort, Rijksmuseum van
Natuurlijke Historie, Leiden, for the identification of the gor-
gonacean hosts.
The observations and measurements were made on speci-
mens cleared in lactic acid. Measurements are in microns
unless otherwise specified. The dimensions of the body are
based on ten specimens and the body length does not include
the setae on the caudal rami. The lengths of the segments of
the first antennae are taken along their posterior nonsetiferous
margins. All figures have been drawn with the aid of a camera
lucida. The letter after the explanation of each figure refers
to the scale at which it was drawn.
Acanthomolgus Humes and Stock, 1972
Acanthomolgus mopsellae, new species
Figures 1-17
Type-material: 285 292, 470 464, and 311 copepodids from one
colony of the gorgonacean Mopsella rubeola Wright and Studer, in 3 m,
Pte. Lokobe, Nosy Bé, Madagascar, 3 June 1967. Holotype 9, allotype,
and 567 paratypes (201 22, 366 66) deposited in the National
Museum of Natural History (USNM), Washington; 90 paratypes (40
292,50 ¢2) in the Zodlogisch Museum, Amsterdam; the remaining
paratypes in the collection of the author.
In the following description those features not mentioned may be
assumed to be essentially like those of Acanthomolgus hians (Humes
and Ho, 1968a).
Female: Body (Fig. 1) with a moderately slender prosome. Length
0.98 mm (0.89-1.02 mm) and the greatest width 0.49 mm (0.46—0.56
mm). Ratio of the length to the width of the prosome 1.45:1. Ratio of
the length of the prosome to that of the urosome 2.91:1.
Segment of leg 5 (Fig. 2) 73 x 156. Between this segment and the
genital segment no ventral intersegmental sclerite. Genital segment 143
>
Fics. 1-6. Acanthomolgus mopsellae, new species, female: 1, dorsal
(A); 2, urosome, dorsal (B); 3, genital area, dorsal (C); 4, caudal
ramus, dorsal (C); 5, first antenna, with arrows indicating positions of
three aesthetes added in the male, dorsal (B); 6, second antenna, an-
terior (inner) (D).
413
Copepods from gorgonaceans in Madagascar
414 Proceedings of the Biological Society of Washington
>< 143. Genital areas located dorsolaterally (more laterally than in A.
hians). Each area (Fig. 3) bearing two naked setae about 9 long;
spiniform process small instead of prominent and unguiform as in A.
hians. Three postgenital segments from anterior to posterior 42 x88,
31 x 83, and 34 x 75. Posteroventral border of the anal segment smooth.
Caudal ramus (Fig. 4) 27 x 31, wider than long. Outer lateral seta
143 and naked, the dorsal seta 30 and delicately haired, the outermost
terminal seta 198 and naked, the innermost terminal seta 275 with lateral
spinules, and the two long median terminal setae 495 (outer) and 583
(inner), both with strong spinules along their midregions.
Egg sacs (Fig. 1) variable in size. Those in the figure approximately
320 x 175, but in another female about 495 x 250. Each of the many
eggs about 47 in diameter.
Rostrum resembling that in A. hians. First antenna (Fig. 5) 425,
with the armature as in A. hians. Lengths of the seven segments: 34
(77 along the anterior margin), 100, 29, 67, 69, 48, and 35 respectively.
Second segment 100 x 28.5, shorter and broader than in A. hians, where
it is 130 x 22.
Second antenna (Fig. 6) with the formula 1, 1, 3, and II + 5, lacking
minute spinules on the first and second segments. Fourth segment 99
along its outer side, 66 along its inner side, and 31 wide. Stouter claw
73 along its axis, the more slender claw 83.
Labrum, mandible, paragnath, first maxilla, and second maxilla like
those of A. hians. Two setae on the second segment of the maxilliped
(Fig. 7) 22 and 43 (in A. hians 28 and 66), the longer seta sharply
bent distally. Two terminal elements on the third segment less divergent
than in A. hians.
Legs 1 and 2 like those of A. hians. Leg 3 also resembling that species
except for the relative lengths of the three spines on the third segment
of the endopod (Fig. 8), which are 38, 35, and 45 from outer to inner
(31, 24, and 36 in A. hians). Leg 4, with the formula for the exopod
I-0; I-1; II, I, 5, like that in A. hians except for details of the endopod
(Fig. 9). First endopod segment 48 x 36 (including the spiniform
processes) and the inner distal spine 30. Second endopod segment 93
long (including the spiniform processes), 32 in greatest width and 23
in least width, with the two terminal spines 31 (outer) and 72 (inner).
>
Fics. 7-10. Acanthomolgus mopsellae, new species, female: 7, maxil-
liped, postero-inner (E); 8, third segment of endopod of leg 3, an-
terior (D); 9, endopod of leg 4, anterior (D); 10, leg 5, dorsal (D).
Fics. 11-17. Acanthomolgus mopsellae, new species, male: 11, dorsal
(F); 12, maxilliped, outer (D); 13, third segment of endopod of leg 1,
anterior (E); 14, third segment of endopod of leg 2, anterior (E); 15,
endopod of leg 4, anterior (D); 16, leg 5, dorsal (EF); 17, spermato-
phores, attached to female, lateral (F).
415
Copepods from gorgonaceans in Madagascar
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416 Proceedings of the Biological Society of Washington
Leg 5 (Fig. 10) with a slender free segment 140 long, 28 wide proxi-
mally, and 22 wide distally. Two terminal setae 130 and 122. Seta on
the body adjacent to the free segment 47. Spines on the free segment
a little more slender and less scalelike than in A. hians.
Living specimens in transmitted light opaque, the intestine brown,
the eye red, the egg sacs gray.
Male: Body (Fig. 11) with a moderately slender prosome as in the
female. Length 0.86 mm (0.83-0.89 mm) and the greatest width 0.36
mm (0.32-0.39 mm). Ratio of the length to the width of the prosome
1.52:1. Ratio of the length of the prosome to that of the prosome 1.58:1.
Genital segment 230 x 210. Caudal ramus 20 x 25.
Rostrum like that of the female. First antenna similar to that of the
female, but three aesthetes added as in A. hians. Second antenna re-
sembling that of the female, but the fourth segment a little more slender,
105 along its outer side, 74 along its inner side, and 25 wide.
Labrum, mandible, paragnath, first maxilla and second maxilla like
those of the female. Maxilliped (Fig. 12) resembling closely that of
A. hians. Claw 180 along its axis including the terminal lamella.
Leg 1 like that of the female except for the third segment of the endo-
pod (Fig. 13) which has the formula I, I, 4 instead of I, 5. Inner spine
with a row of spinules along the inner side but the outer side smooth,
thus differing from A. hians where both margins are barbed. Leg 2
like that of the female, with the third segment of the endopod (Fig.
14) not showing an enlarged outer terminal process as in the male of
A. hians. Leg 3 like that of the female. Leg 4 resembling that of the
female except for dimensions of the endopod (Fig. 15). First segment
37 xX 24 (including the spiniform processes) and its inner distal spine
26. Second segment 70 long (including the processes), 21 in greatest
width, and 15 in least width, with the two terminal spines 25 (outer)
and 64 (inner).
Leg 5 (Fig. 16) with a long slender free segment 57 x 10.5 bearing
a few minute spines. Two terminal elements 81 x 23 and the seta on
the body adjacent to the free segment approximately 30.
Leg 6 like that of A. hians.
Spermatophore (Fig. 17) oval, 200 <x 100, not including the neck.
Living specimens colored as in the female.
Etymology: The specific name mopsellae is the genitive form of the
generic name of the host.
Comparison with other species in the genus: In the keys published by
Humes (1973a) to both sexes of the 17 known species of the genus
Acanthomolgus, the new species from Mopsella falls in the same position
as A. hians, a species from the alcyonacean Siphonogorgia pichoni Verse-
veldt at Nosy Bé, Madagascar (for the correct name of the host see
Humes and Stock, 1973). Although the two species have certain sim-
ilarities, careful study shows that there are many consistent significant
differences. The more important and easily recognized of these are
summarized in Table 1.
Copepods from gorgonaceans in Madagascar A417
TaBLE 1. Comparison of distinguishing characters of Acanthomolgus
mopsellae and Acanthomolgus hians.
FEMALE
genital areas
length of second
segment of first
antenna
free segment of
leg 5
MALE
length of claw on
maxilliped
inner of 2 spines
on third segment
A. mopseliae
situated far apart,
each with a small
pointed process
relatively short,
ratio 3.5:1
slender, with 2
terminal setae
nearly equal
in length
180 microns
with barbs along
inner side only
A. hians
closer together, each
with a prominent
unguiform process
more elongate, ratio
5.9:1
stouter, with 2
terminal setae
very unequal
159 microns
with barbs along
both sides
of endopod of
leg 1
outer terminal process small enlarged
on third segment of
endopod of leg 2
ratio of length to 5.4:1 44:1
width of free segment
of leg 5
Acanthomolgus combinatus, new species
Figures 18-41
Type-material: 114 99, 278 ¢ 4, and many copepodids from one
colony of Echinogorgia sasappo (Esper), in 10 m, Pte. Lokobe, Nosy
Bé, Madagascar, 18 July 1967. Holotype @, allotype, and 384 para-
types (110 2 9, 274 4 4), deposited in the National Museum of Nat-
ural History (USNM), Washington; the remaining paratypes (dissected )
in the collection of the author.
Other specimens (all from Echinogorgia sasappo): 3 2@, 10 64,
and a few copepodids from one colony, in 10 m, Nosy N’Tangam, near
Nosy Bé, 1 January 1964; 4 99, 18 ¢ 4, and a few copepodids from
418 Proceedings of the Biological Society of Washington
one colony, in 25 m, Tany Kely, a small island to the south of Nosy Bé,
14 August 1964; 55 9 9,57 ¢ 6, and a few copepodids from one colony,
in 25 m, Tany Kely, 14 August 1964; 1 9,2 6 46, and a few copepodids
from one colony, in 13 m, opposite Antsiabe, Nosy Komba, near Nosy
Bé, 2 September 1967.
Female: Body (Fig. 18) resembling in general form that of A. mop-
sellae. Length 1.10 mm (1.02-1.19 mm) and the greatest width 0.60
mm (0.58-0.62 mm). Ratio of the length to the width of the prosome
1.47:1. Ratio of the length of the prosome to that of the urosome 3.06:1.
Segment of leg 5 (Fig. 19) 86 x 174. Between this segment and the
genital segment no ventral intersegmental sclerite. Genital segment 143
< 169, in dorsal view broadest in its midregion, posterior to which the
lateral margins are obliquely truncated, then abruptly constricted near
the end of the segment. Genital areas located dorsolaterally along the
truncated margins. Each area (Fig. 20) bearing two short naked setae
11 and 20 and a small spiniform process. Adjacent to the area two
spiniform papillate processes, one larger than the other. Three post-
gential segments from anterior to posterior 22 x 91, 23 x 86, and 34
< 84. Posteroventral border of the anal segment bearing a row of mi-
nute spinules on each side.
Caudal ramus (Fig. 19) quadrate, 34 x 34. Outer lateral seta 112
and the dorsal seta 39, both naked. Outermost terminal seta 230 and
naked, the innermost terminal seta 350 with spinules along the inner
side. Two long median terminal setae 570 and 690, both with strong
lateral spinules along their midregions. A minute setule on the proximal
outer side of the ramus. |
Body surface bearing only a few small hairs (sensilla) as in Fig. 18.
Egg sac (Fig. 21) 550-600 x 240, reaching nearly to the ends of
the longest setae on the caudal ramus, and containing many eggs about
50 in diameter.
Rostrum (Fig. 22) in ventral view broadly rounded posteroventrally.
First antenna (Fig. 23) 492 long. Lengths of the seven segments:
36 (75 along the posterior margin), 135, 31, 78, 76, 62, and 35 re-
spectively. Formula for the armature the same as in other species of
the genus: 4, 13, 6, 3, 4 + 1 aesthete, 2 + 1 aesthete, and 7 + 1 aes-
thete. All the setae naked.
Second antenna (Fig. 24) 4-segmented with the formula 1, 1, 3, and
Ii + 5. All the setae naked. Fourth segment 86 along its outer edge,
=>
Fics. 18-28. Acanthomolgus combinatus, new species, female: 18,
dorsal (A); 19, urosome, dorsal (G); 20, genital area, dorsal (E); 21,
egg sac, dorsal (F); 22, rostrum, ventral (B); 23, first antenna, ventral
(G); 24, second antenna, posterior (outer) (D); 25, first maxilla, pos-
terior (C); 26, second maxilla, posterior (E); 27, maxilliped, posterior
(E); 28, leg 1 and intercoxal plate, anterior (B).
419
Copepods from gorgonaceans in Madagascar
420 Proceedings of the Biological Society of Washington
50 along its inner edge, and 27 wide. More slender terminal claw 55
and the stouter claw 57. First and second segments lacking fine orna-
mentation.
Labrum like that in A. hians and A. verseveldti (Humes and Ho,
1968a). Mandible similar to that in A. telestophilus (Humes and Ho,
1968a). Paragnath resembling that of A. verseveldti. First maxilla
(Fig. 25) with three elements. Second maxilla (Fig. 26) 2-segmented.
First segment unarmed. Second segment with a small setule on its
proximal outer (ventral) surface, a surficial posterior seta finely barbed
along one edge, a long inner (dorsal) distal setiform spine with spinules
along one edge; the terminal lash with a broad hyaline lamella fringed
with small teeth. Maxilliped (Fig. 27) lacking ornamentation on the
first segment. Two naked setae on the second segment 6.5 and 28.5.
Small third segment with a small naked seta and two nearly equal ter-
minal spinelike elements.
Ventral area between the maxillipeds and the first pair of legs like
that in A. verseveldti.
Legs 1-4 (Figs. 28, 30, 31, and 32) segmented and armed as in other
species in the genus. Outer spines on the exopod of leg 1 having con-
spicuous lateral barbs (Fig. 29). Inner coxal element of leg 4 a short
naked seta 10 long; inner margin of the basis of this leg smooth. Exopod
196. First segment of the endopod 53 x 35 (including the spiniform
processes) and the inner distal spine 31 and very finely barbed. Second
segment 107 x 32 (greatest width and including the spiniform pro-
cesses). Outer terminal spine 27.5 and finely barbed; inner terminal
spine 68 and more coarsely barbed, the barbs along the outer margin
being larger than those on the inner margin. (In another female these
spines 33 and 72.)
Leg 5 (Fig. 33) with a long slender free segment having a rounded
proximal inner expansion. Segment 143 long, 47 in greatest width at
the expansion and 18 in least width distally. Two terminal setae 169
(with a narrow outer lamella) and 135. Seta on the body near the in-
sertion of the segment about 44 and apparently naked. Near this seta
a patch of small spines. Free segment ornamented on its outer dorsal
surface with short spines.
Leg 6 represented by the two setae and the spiniform process on the
genital area.
>
Fics. 29-33. Acanthomolgus combinatus, new species, female: 29,
spines on exopod of leg 1, anterior (E); 30, leg 2, anterior (B); 31,
third segment of endopod of leg 3, anterior (B); 32, leg 4 and intercoxal
plate, anterior (B); 33, leg 5, dorsal (D). Fics. 34-36. Acanthomolgus
combinatus, new species, male: 34, dorsal (F); 35, urosome, dorsal (G),
36, second antenna, posterior (outer) (D).
AQT
Copepods from gorgonaceans in Madagascar
422 Proceedings of the Biological Society of Washington
Living specimens in transmitted light opaque and slightly reddish,
the eye red, the egg sacs dark gray.
Male: Body (Fig. 34) resembling that of A. mopsellae. Length 0.93
mm (0.83-0.99 mm) and the greatest width 0.37 mm (0.34-0.39 mm).
Ratio of the length to the width of the prosome 1.50:1. Ratio of the
length of the prosome to that of the urosome (in the specimen drawn)
1.94:1. In another male the postgenital segments less contracted (as
in Fig. 35) and the ratio 1.70:1.
Segment of leg 5 (Fig. 35) 47 x 109. No ventral intersegmental
sclerite. Genital segment 224 x 213. Four postgenital segments from
anterior to posterior 29 x 57, 21 x 62, 15.5 x 60, and 23.5 x 60.
Caudal ramus 22 x 24, and armed as in the female.
Fine ornamentation of the body resembling that of the female.
Rostrum like that of the female. First antenna resembling that of
the female, but three aesthetes added as in the male of A. mopsellae.
Second antenna (Fig. 36) similar to that of the female, but slight sexual
dimorphism indicated by the minute spinules along the inner surface
of the second segment and by the more slender proportions of the fourth
segment (79 along the outer side, 53 along the inner side, and 47 wide).
Terminal claws 55 and 53.
Labrum, mandible, paragnath, first maxilla, and second maxilla like
those in the female. Maxilliped (Fig. 37) slender and 4-segmented
(assuming that that proximal half of the claw represents a fourth seg-
ment). First segment unarmed. Second segment with two rows of spines
and two naked setae, one with a minutely trifurcated tip (Fig. 38).
Small third segment unarmed. Claw 168 including the small terminal
lamella, and bearing proximally two unequal setae.
Ventral area between the maxillipeds and the first pair of legs like
that of the female.
Legs 1-4 like those of the female, except for the third segment of
the endopod of leg 1 (Fig. 39) which has the formula I, I, 4, the two
spines being coarsely barbed. On the second segment of the endopod
of leg 4 (Fig. 40) the two terminal spines 27.5 and 55. (In another
male these spines 25 and 52.)
Leg 5 (Fig. 41) with a long slender free segment 65 x 8, ornamented
with a few small spines. Inner terminal seta 28 and barbed along its
outer side, the outer seta 70 and naked.
=>
Fics. 37-41. Acanthomolgus combinatus, new species, male: 37,
maxilliped, inner (D); 38, seta on second segment of maxilliped, outer
(H); 39, endopod of leg 1, anterior (D); 40, endopod of leg 4, an-
terior (D); 41, leg 5, dorsal (E). Fics. 42-46. Acanthomolgus arc-
tatipes, new species, female: 42, dorsal (F); 43, urosome, dorsal (D);
44, genital area, dorsal (H); 45, caudal ramus, dorsal (1); 46, first
antenna, dorsal (D).
Copepods from gorgonaceans in Madagascar 493
424 Proceedings of the Biological Society of Washington
Leg 6 resembling that in A. telestophilus. Two setae 65 and 39.
Spermatophore not observed.
Living specimens having a color similar to that of females.
Etymology: The specific name combinatus, Latin = combined or
united, refers to the incorporation in this species of several characters
found in related forms.
Comparison with other members of the genus: The new species bears
many similarities with A. telestophilus, a species from the telestacean
Telesto arborea Wright and Studer at Nosy Bé, Madagascar. In fact, in
the keys of Humes (1973a) it falls with A. telestophilus. There are,
however, several ways in which A. combinatus differs significantly from
that species. These are summarized in Table 2.
Acanthomolgus arctatipes, new species
Figures 42-63
Type-material: 99 99, 57 66, and many copepodids from one
colony of Echinogorgia sasappo (Esper), in 10 m, Pte. Lokobe, Nosy
Bé, Madagascar, 18 July 1967. Holotype 9, allotype, and 149 para-
types (95 99, 54 $4) deposited in the National Museum of Natural
History (USNM), Washington; the remaining paratypes (dissected) in
the collection of the author.
Other specimens (all from Echinogorgia sasappo): 16 92 and a
few copepodids from one colony, in 10 m, Nosy N’Tangam, near Nosy
Bé, 1 January 1964; 49 99,13 66, and a few copepodids from one
colony, in 25 m, Tany Kely, a small island to the south of Nosy Bé, 14
August 1964; 33 99,1 ¢ from one colony in 25 m, Tany Kely, 14
August 1964; 6 9° 2,1 6 from one colony, in 13 m, opposite Antsiabe,
Nosy Komba, near Nosy Bé, 2 September 1967.
Female: Body (Fig. 42) resembling in general form that of A. astrictus
Humes and Stock, 1973. Length 0.74 mm (0.66-0.80 mm) and the
greatest width 0.34 mm (0.29-0.37 mm). Ratio of the length to the
width of the prosome 1.52:1. Ratio of the length of the prosome to
that of the urosome 2.05:1.
Segment of leg 5 (Fig. 43) 47 x 93. Between this segment and the
genital segment no ventral intersegmental sclerite. Genital segment
99 x 90, in dorsal view with gently rounded sides. Genital areas located
dorsolaterally at the middle of the segment. Each area (Fig. 44) bear-
ing two naked setae 7 and 11 and a small spiniform process. Medial
to the genital area an elongated lobe with a mucronate tip. Three post-
genital segments from anterior to posterior 29 x 54, 23 x 52, and 24
< 52. Posteroventral border of the anal segment bearing a row of small
spinules on each side.
Caudal ramus (Fig. 45) quadrate, 21 x 21. Outer lateral seta 52,
the dorsal seta 17, the outermost terminal seta 70, the innermost ter-
minal seta 130, and the two long median terminal setae 247 (outer)
and 380 (inner), both inserted between dorsal (with a smooth margin)
Copepods from gorgonaceans in Madagascar 495
TABLE 2. Comparison of distinguishing characters of Acanthomolgus
combinatus and Acanthomolgus telestophilus.
FEMALE
length of body
genital segment
outermost terminal
seta on caudal ramus
second antenna
first maxilla
second maxilla
longer of 2 setae on
second segment of
maxilliped
MALE
length of body
second antenna
setae on second
segment of
maxilliped
dimensions of free
segment of leg 5
A. combinatus
1.10 mm
(1.02—1.19 mm)
sides obliquely trun-
cated posterior to
middle
naked
inner surfaces of all
4 segments smooth;
all three setae on
third segment
straight
with 3 elements
lash having hyaline
lamella with small
marginal teeth
smooth
0.93 mm
(0.83-0.99 mm)
with inner spinules
on second segment
only
one with trifurcated
tip, other attenuated
65 x 8 microns
A. telestophilus
0.91 mm
(0.85-0.95 mm )
sides rounded rather
than truncated
with lateral spinules
minute inner spin-
ules on all 4 seg-
ments; one of setae
on third segment
angularly bent
with 4 elements
lash with graduated
teeth; no lamella
with spinules
0.75 mm
(0.72—0.79 mm )
with inner spinules
on all 4 segments
both attenuated
38 x 8.5 microns
and ventral (with a marginal row of spinules) flanges. All the setae
naked.
Body surface bearing only a very few small hairs (sensilla) as in
Fig. 42.
426 Proceedings of the Biological Society of Washington
Egg sac (Fig. 42) 350 x 145, reaching to more than half the length
of the longest ramal setae, and containing numerous eggs 45-47 in
diameter.
Rostrum like that in A. astrictus.
First antenna (Fig. 46) 285 long. Lengths of the seven segments:
21 (44 along the anterior edge), 81, 21, 36, 44, 34, and 25 respectively.
Formula for the armature like that in A. combinatus. All the setae
naked.
Second antenna (Fig. 47) segmented and armed as in the previous
two species. One of the three setae on the third segment characteris-
tically bent. Fourth segment 62 along the outer side, 43 along the inner
side, and 14 wide. Stouter claw 42 along its axis, the more slender claw
38. All the setae naked.
Labrum (Fig. 48) with two broad posteroventral lobes.
Mandible (Fig. 49) basically similar to that of A. astrictus, but the
convex side of the base having an expanded hyaline area without spinules
followed by a minutely serrated portion. Paragnath (Fig. 48) a small
hairy lobe. First maxilla (Fig. 50) with four elements, one of them with
small spinules. Second maxilla (Fig. 51) resembling in general structure
that of A. astrictus, but the lash having along its convex side an expanded
hyaline lamella with a finely dentate margin. Maxilliped (Fig. 52) also
resembling that of A. astrictus but the first segment lacking spinules and
the two setae on the second segment more unequal, 7.5 and 22, with a
ratio of approximately 1:3.
Ventral area between the maxillipeds and the first pair of legs like
that in A. astrictus.
Legs 1—4 segmented and armed as in A. astrictus with only minor
differences in details. In the endopod of leg 1 (Fig. 53) the terminal
process on the third segment having a slightly different form than in
that species. In leg 4 (Fig. 54) the inner coxal seta only 2 long. Exo-
pod 112. First segment of the endopod 28.5 x 22 (including the ter-
minal spiniform processes) and its inner distal spine (extremely finely
barbed) 13. Second segment 55 x 17.5 (greatest width and the length
including the terminal spiniform processes). Two terminal spines un-
equal, the outer 21 and slender with very small barbs, the inner 42
and stouter with more prominent barbs along the outer side than the
inner side.
Leg 5 (Fig. 55) with an elongated free segment 86 long. At the
=>
Fics. 47-54. Acanthomolgus arctatipes, new species, female: 47,
second antenna, posterior (E); 48, labrum, with paragnaths indicated
by broken lines, ventral (C); 49, mandible, anterior (1); 50, first maxilla,
anterior (C); 51, second maxilla, posterior (1); 52, maxilliped, pos-
terior (I); 53, endopod of leg 1, anterior (E); 54, leg 4, anterior (D).
Copepods from gorgonaceans in Madagascar A427
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428 Proceedings of the Biological Society of Washington
proximal beaklike inner expansion the width 15.5. At the slender neck
the width only 6.5. Width increasing distally to the widest point of 15.5.
Two terminal setae 44 and smooth and 78 with a narrow lamella. Seg-
ment ornamented outwardly with small spines. Adjacent seta on the
body, held erect and difficult to measure, about 35 and naked.
Leg 6 represented by the two setae and the spiniform process on the
genital area.
Living specimens in transmitted light opaque, the eye red, the egg
sacs gray.
Male: Body (Fig. 56) slender. Length 0.61 mm (0.57—-0.65 mm) and
the greatest width 0.21 mm (0.19-0.23 mm). Ratio of the length to the
width of the prosome 1.77:1. Ratio of the length of the urosome to that
of the prosome 1.60:1.
Genital segment (Fig. 57) 24 x 56. No ventral intersegmental sclerite.
Genital segment 110 x 107. Four postgenital segments from anterior
to posterior 19 x 38.5, 19 x 36, 11 x 35.5, and 14 x 36.
Caudal ramus resembling that of the female but smaller, 17 x 17.
Fine ornamentation on the body like that of the female.
Rostrum like that of the female. First antenna similar to that of the
female, but three aesthetes added as in the males of A. mopsellae and
A. combinatus. Second antenna (Fig. 58) resembling that of the female,
but sexually dimorphic in having minute spinules on the inner margins
of the first, second, and fourth segments.
Labrum, mandible, paragnath, first maxilla, and second maxilla like
those of the female. Maxilliped (Fig. 59) resembling that of A. astrictus,
but both setae on the second segment naked. Claw 117 along its axis.
Ventral area between the maxillipeds and the first pair of legs like
that in the female.
Legs 1-4 segmented and armed as in the female, except for the last
segment of the endopod of leg 1 (Fig. 60) where the formula is I, I,
A instead of I, 5 as in the opposite sex. Two terminal spines on the endo-
pod of leg 4 (Fig. 61) having proportions different from those in the
female, the outer being 13, the inner 34 long.
Leg 5 (Fig. 62) with a rectangular free segment 25 x 6.5 ornamented
with a few small spinules. Two terminal elements 32 and smooth, and
15.5 with barbs along the outer edge.
=>
Fic. 55. Acanthomolgus arctatipes, new species, female: 55, leg 5,
dorsal (C). Fics. 56-63. Acanthomolgus arctatipes, new species, male:
56, dorsal (F); 57, urosome, dorsal (D); 58, second antenna, posterior
(E); 59, maxilliped, outer (E); 60, endopod of leg 1, anterior (C);
61, endopod of leg 4, anterior (E); 62, leg 5, dorsal (1); 63, leg 6,
ventral (E). Fic. 64. Paramolgus ellisellae, new species, female: 64,
dorsal (F).
429
Copepods from gorgonaceans in Madagascar
430 Proceedings of the Biological Society of Washington
Leg 6 (Fig. 63) consisting of the usual posteroventral flap on the
genital segment bearing two naked setae 17 and 41.
Spermatophore not observed.
Living specimens colored as in the female.
Etymology: The specific name arctatipes, Latin arctatus = constricted
and pes = foot, alludes to the constricted leg 5 in the female.
Comparison with other members of the genus: Only three species of
Acanthomolgus have on the free segment of leg 5 in the female an inner
proximal distally directed process separated sharply from the more distal
margin of the segment. These are A. astrictus from the gorgonacean
Acanthogorgia aspera Pourtalés, A. cuneipes (Humes and Ho, 1968b)
from the alcyonaceans Stereonephthya acaulis Verseveldt and Dendro-
nephthya mucronata (Pitter), and A. longispinifer (Humes and Ho,
1968a) from the alcyonacean Siphonogorgia pichoni Verseveldt, all in
Madagascar. The new species differs from these, however, in many
features. It seems to be closest to A. astrictus, but is easily separated
from that species which has lateral spinules on the two long median setae
on the caudal rami.
The expanded hyaline area without spinules on the convex side of
the mandible and the hyaline finely dentate lamella on the lash of the
second maxilla distinguish A. arctatipes from the other three species just
mentioned. These species lack a similar hyaline expansion on the man-
dible and have large teeth but no lamella on the lash of the second
maxilla. In addition, A. longispinifer has lateral spinules on the two
long median setae of the caudal rami and the outer spine on the first
segment of the exopod of leg 1 is unusually long. A. cuneipes has a
relatively shorter fourth segment in the second antenna and the free
segment of leg 5 in the female has a different form, without a constricted
neck as in A. arctatipes.
Acanthomolgus astrictus Humes and Stock, 1973
Specimens collected: From the gorgonacean Acanthogorgia aspera
Pourtalés: 15 99,16 ¢ 6, and 7 copepodids from one colony, in 4 m,
Pte. Lokobe, Nosy Bé, 24 December 1963; 28 99, 17 66, and 7
copepodids from one colony, in 40 m, Banc de Cinq Métres, near Nosy
Bé, 19 August 1964; 56 2 9, 22 ¢ 8, and 4 copepodids from one colony,
in 8 m, Nosy Ovy, Isles Radama, south of Nosy Bé, 13°59’S, 47°46.5’E,
>
Fics. 65-73. Paramolgus ellisellae, new species, female: 65, urosome,
dorsal (B); 66, genital area, dorsal (1); 67, caudal ramus, dorsal (E);
68, rostrum, ventral (B); 69, first antenna, with arrows indicating posi-
tions of aesthetes added in the male, dorsal (D); 70, second antenna,
posterior (E); 71, labrum, with paragnaths indicated by broken lines,
ventral (E.); 72, mandible, posterior (1); 73, first maxilla, anterior (1).
Copepods from gorgonaceans in Madagascar 431
432 Proceedings of the Biological Society of Washington
30 September 1964. From the gorgonacean Muricella rubra robusta
Thomson and Simpson: 77 2 9, 44 44, and 4 copepodids from one
colony, in 15 m, Tany Kely, near Nosy Bé, 30 August 1964; 3 92,1 ¢
from one colony, in 10 m, Nosy N’Tangam, near Nosy Bé, 1 June 1964.
Muricella rubra robusta is a new host for this species.
Acanthomolgus hales Humes and Stock, 1973
Specimens collected: 157 992, 190 66, and 103 copepodids from
one colony of the gorgonacean Solenocaulon tortuosum Gray, in 18 m,
on sand in the pass at Pte. Lokobe, Nosy Bé, 14 July 1967.
Metaxymolgus cinctus Humes and Stock, 1973
Specimens collected (all from the gorgonacean Psammogorgia ramosa
Kikenthal): 17 99, 43 6 6, and 99 copepodids from one colony, in
2m, northern end of Nosy Sakatia, near Nosy Bé, 19 August 1963; 22
992, 73 86, and 206 copepodids from one colony, in 2 m, east of
Ambariotelo, near Nosy Bé, 20 July 1967; 4 99, 60 $6, and 132
copepodids from two colonies, in 15 m, Nosy Iranja, southwest of Nosy
Bé, 9 August 1967.
Paramolgus Humes and Stock, 1972
Paramolgus ellisellae, new species
Figures 64-90
Type-material: 30 22, 38 6 4, and 63 copepodids from one colony
of the gorgonacean Ellisella ramosa (Simpson), in 24 m, Banc des Fréres,
Isles Mitsio, northeast of Nosy Bé, Madagascar, 12°58’S, 48°28’E, 17
August 1967. Holotype @, allotype, and 60 paratypes (26 2? 2, 34 64)
deposited in the National Museum of Natural History (USNM), Wash-
ington; the remaining paratypes (dissected) in the collection of the
author.
Other specimens: 30 2 2, 23 6 6, and 22 copepodids from one colony
of Ellisella ramosa, in 25 m, north of Ankazoberavina, near Nosy Bé,
13°27.6’S, 47°58.2’E, 24 August 1967.
Female: Body (Fig. 64) with a broad flattened prosome. Length 0.81
mm (0.75-0.86 mm) and the greatest width 0.43 mm (0.37-0.46 mm).
Epimeral areas of the segments of legs 1-4 rounded. Ratio of the length
to the width of the prosome 1.09:1. Ratio of the length of the prosome
to that of the urosome 1.65:1.
=>
Fics. 74-79. Paramolgus ellisellae, new species, female: 74, second
maxilla, anterior (C); 75, maxilliped, anterior (C); 76, area between
maxillipeds and first pair of legs, ventral (D); 77, leg 1 and intercoxal
plate, anterior (E); 78, leg 2, anterior (E.); 79, third segment of endopod
of leg 3, anterior (E).
433
Copepods from gorgonaceans in Madagascar
434 Proceedings of the Biological Society of Washington
Segment of leg 5 (Fig. 65) 73 x 112. Between this segment and
the genital segment no ventral intersegmental sclerite. Genital segment
96 long, in dorsal view moderately widened in its anterior three-fourths
(greatest width 122) and abruptly narrowed in its posterior fourth
(width 74). Genital areas situated dorsolaterally. Each area (Fig. 66)
bearing two naked setae about 8 long and a small blunt spiniform pro-
cess. Nearby a large pointed bladelike process. Three postgenital seg-
ments from anterior to posterior 42 x 68, 31 x 65, and 26 x 57. Pos-
teroventral border of the anal segment smooth.
Caudal ramus (Fig. 67) elongated, 60 <x 26, with the ratio being
about 2.3:1. Outer lateral seta 47 and naked. Dorsal seta 38 and
sparsely feathered. Outermost terminal seta 88, the innermost terminal
seta 115, and the two long median terminal setae 210 (outer) and 400
(inner), both inserted between smooth dorsal and ventral flanges. All
four terminal setae having lateral spinules.
Body surface with only a very few small hairs (sensilla) as in Fig. 64.
Egg sac not observed.
Rostrum (Fig. 68) broad and weakly delimited posteroventrally.
First antenna (Fig. 69) 7-segmented, with the formula 4, 13, 6, 3,
4 + ] aesthete, 2 + 1 aesthete, and 7 + 1 aesthete. Lengths of the
segments: 24 (49 along the anterior margin), 104, 24, 38, 34, 27, and
24 respectively. All the setae naked.
Second antenna (Fig. 70) 180 long and 4-segmented with the formula
1, 1, 3, and I + 5. Fourth segment 60 along the outer side, 45 along
the inner side, and 17 wide. Claw 23 along its axis. All the setae naked.
Labrum (Fig. 71) with two divergent posteroventral lobes, each with
a small rounded terminal hyaline process.
Mandible (Fig. 72) having on the convex side of the base a scalelike
area with a row of spinules followed by a deeply insected dentate fringe,
and. on the concave side a row of long spinules. Lash long with only a
few spinules proximally. Paragnath (Fig. 71) a small hairy lobe. First
maxilla (Fig. 73) with four elements. Second maxilla (Fig. 74) 2-
segmented. First segment unarmed. Second segment bearing a small
proximal inner setule, a naked surficial posterior seta, and an outer distal
spine with long spinules along the distal edge and a few on the proximal
edge. Segment produced distally to form a lash with graduated teeth
>
Fics. 80-81. Paramolgus ellisellae, new species, female: 80, leg 4 and
intercoxal plate, anterior (E); 81, leg 5, dorsal, but free segment dorso-
ventral (EF). Fics. 82-90. Paramolgus ellisellae, new species, male: 82,
dorsal (F); 83, urosome, dorsal (B); 84, second antenna, posterior (E);
85, maxilliped, outer (D); 86, third segment of endopod of leg 1, an-
terior (E); 87, third segment of endopod of leg 2, anterior (E); 88,
third segment of endopod of leg 3, anterior (E); 89, leg 5, dorsal, but
free segment dorsoventral (E); 90, leg 6, ventral (E).
435
Copepods from gorgonaceans in Madagascar
436 Proceedings of the Biological Society of Washington
along the distal edge and with bilateral spinules distally. First tooth
on the lash much larger than the others. Maxilliped (Fig. 75) 3-seg-
mented. First segment unarmed. Second segment bearing two naked
setae 5 and 50. Third segment bearing a seta 14 and terminating in two
unequal spiniform processes.
Ventral area between the maxillipeds and the first pair of legs (Fig.
76) not protuberant.
Legs 1-4 (Figs. 77, 78, 79, and 80) segmented and armed as in other
species of the genus, for example, Paramolgus politus (Humes and Ho,
1967). In leg 1 (Fig. 77) the coxa having a small outer posterior pro-
trusion and the outer spines on the exopod being prominently barbed.
In leg 2 (Fig. 78) the three spines on the third segment of the endopod
10, 9, and 11 from proximal to distal; in leg 3 these spines 10, 9, and
12 (Fig. 79). In leg 4 (Fig. 80) the exopod 95 long. Inner element on
the coxa reduced to a minute setule 5 long. Inner margin of the basis
smooth instead of haired as in the preceding legs. First segment of the
endopod 30 long (including the spiniform processes) and 16 wide, with
its inner distal feathered seta 33. Second segment 54 long (including
the spiniform processes) and 13 wide. Two terminal barbed spines 15.5
(outer) and 27.5 (inner).
Leg 5 (Fig. 81) having a free segment 80 long, 39 wide at the large
rounded proximal inner expansion, and 18 wide distally. Two terminal
setae 57 and 81. Adjacent seta on the body 57. All the setae naked.
Outer distal surface of the free segment bearing a few small spines.
Leg 6 represented by the two setae and spiniform process on the
genital area.
Living specimens in transmitted light opaque, the eye red.
Male: Body (Fig. 82) with the prosome less expanded than in the
female. Length 0.71 mm (0.67-0.73 mm) and the greatest width 0.30
mm (0.28-0.32 mm). Ratio of the length to the width of the prosome
1.30:1. Ratio of the length of the prosome to that of the urosome 1.49:1.
Segment of leg 5 (Fig. 83) 29 x 64. No ventral intersegmental
sclerite. Genital segment 143 x 122. Four postgenital segments from
anterior to posterior 26 x 47, 29 x 47, 21 x 43, and 18 x 42.
Caudal ramus resembling that of the female, but shorter, 42 x 21,
with the ratio 2:1.
Fine ornamentation of the body like that of the female.
Rostrum similar to that of the female. First antenna resembling that
of the female but two aesthetes added so that the formula is 4, 13 + 1
aesthete, 6, 3 + 1 aesthete, 4 + 1 aesthete, 2 + 1 aesthete, and 7 + 1
aesthete.
Second antenna (Fig. 84) showing sexual dimorphism in having a
few spinules along the inner surfaces of the second and fourth segments
and in the three setae on the third segment being finely barbed along
one edge.
Labrum, mandible, paragnath, first maxilla, and second maxilla like
Copepods from gorgonaceans in Madagascar 437
those of the female. Maxilliped (Fig. 85) slender and 4-segmented (as-
suming that the proximal part of the claw represents the fourth segment).
First and third segments unarmed. Second segment bearing a naked
seta, a spiniform seta finely barbed along one side, and two rows of
spines. Claw 142 along its axis (including the small terminal lamella),
showing a weak subdivision about midway, and bearing proximally two
very unequal setae, the longer seta with short barbules distally.
Ventral area between the maxillipeds and the first pair of legs like
that of the female.
Legs 1-4 segmented and armed as in the female, except for the third
segment of the endopod of leg 1 (Fig. 86) where the formula is I, I, 4
instead of I, 5 as in the opposite sex. These two spines 13 and 22 re-
spectively. Slight sexual dimorphism in leg 2 where the three spines on
the third endopod segment (Fig. 87) are 14, 14, and 23 and in leg 3
where these spines (Fig. 88) are 15.5, 15.5, and 28.5 respectively.
Leg 5 (Fig. 89) with a small free segment 33 x 11, armed terminally
with a short spiniform finely barbed seta 15 with a narrow inner lamella,
and a longer naked seta 34.
Leg 6 (Fig. 90) consisting of the usual posteroventral flap on the
genital segment bearing two naked setae 24 and 28.
Spermatophore not observed.
Living specimens colored as in the female.
Etymology: The specific name ellisellae is the genitive form of the
generic name of the host.
Comparison with other species in the genus: Humes and Stock (1973)
listed seven described species in the genus Paramolgus, but one of these,
P. anomalus (A. Scott, 1909) was included with reservations. Two ad-
ditional species have recently been described (Humes, 1973b).
The large rounded proximal expansion on the free segment of leg 5
in the female distinguishes Paramolgus ellisellae from all other species
in the genus. Both sexes may be separated from all other species by the
proximalmost tooth on the lash of the second maxilla being greatly en-
larged. In other Paramolgus this tooth is not conspicuously enlarged
and forms part of a graduated series with the succeeding teeth.
Although species of Paramolgus are known from various cnidarians
(actiniarians, aleyonaceans, and antipatharians), P. ellisellae is the first
record from gorgonaceans.
Telestacicola angoti Humes and Stock, 1973
Specimens collected: From Suberogorgia reticulata (Ellis and So-
lander): 14 99,6 66 from one colony, in 8 m, Pte. Lokobe, Nosy
Bé, 25 July 1967. From Suberogorgia suberosa (Pallas): 1 9 from
one colony, in 17 m, near the black buoy north of Pte. Ambarionaomby,
Nosy Komba, near Nosy Bé, 5 August 1967.
These two gorgonaceans are new hosts for T. angoti.
438 Proceedings of the Biological Society of Washington
LITERATURE CITED
Humes, A.G. 1973a. Cyclopoid copepods of the genus Acanthomolgus
(Lichomolgidae) associated with gorgonians in Bermuda. Jour.
Nat. Hist. 7:85-115.
1973b. Cyclopoid copepods (Lichomolgidae) from octocorals
at Eniwetok Atoll. Beaufortia 21(282):135-151.
———., AND J.-S. Ho. 1967. Two new species of Lichomolgus (Co-
pepoda, Cyclopoida) from an actiniarian in Madagascar.
Cahiers O.R.S.T.O.M. sér. Océanogr. 5(1):1-21.
—. anp J.-S. Ho. 1968a. Cyclopoid copepods of the genus Lich-
omolgus associated with octocorals of the families Xeniidae,
Nidaliidae, and Telestidae in Madagascar. Proc. Biol. Soc.
Wash. 81:693-750.
———, AND J.-S. Ho. 1968b. Cyclopoid copepods of the genus Lich-
omolgus associated with octocorals of the family Nephtheidae
in Madagascar. Proc. U. S. Nat. Mus. 125(3661):1-41.
———, anD J. H. Srocx. 1972. Preliminary notes on a revision of
the Lichomolgidae, cyclopoid copepods mainly associated with
marine invertebrates. Bull. Zool. Mus., Univ. Amsterdam 2:
121-133.
————, AND J. H. Strocx. 1973. A revision of the family Lichomol-
gidae Kossmann, 1877, cyclopoid copepods mainly associated
with marine invertebrates. Smithsonian Contr. Zool. no. 127:
i-v, 1-368.
Scotr, A. 1909. The Copepoda of the Siboga Expedition. Part 1.
Free-swimming, littoral and semi-parasitic Copepoda. Siboga
Exped. Monogr. 29a:1-323.
Stock, J. H., anp G. KLEErron. 1963. Copépodes associés aux in-
vertébrés des cétes du Rousillon. 2. Lichomolgidae ecto-
associés doctocoralliaires. Vie et Milieu 14(2):245-261.
v
v
)
Wir OCT S5
Vol. 87, No. 38, pp. 439-450 31 December 1974
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
A NEW SPECIES OF NESOTROCHIS FROM
HISPANIOLA, WITH NOTES ON OTHER FOSSIL RAILS
FROM THE WEST INDIES (AVES: RALLIDAE)
By Storrs L. OLson
Smithsonian Institution, Washington, D. C. 20560
In the National Museum of Natural History (USNM) are
extensive collections of vertebrate remains from cave and
midden deposits in the Greater Antilles. Most of these were
obtained in the 1920's under the auspices of Gerrit S. Miller.
A few papers concerning some of these remains were published
by Miller and Alexander Wetmore, after which the collections
were set aside by Miller. Subsequently they have been largely
neglected. I have recently attempted some rough sorting of
the avian material in these collections which has resulted in
several significant discoveries of fossil birds. Because much
yet remains to be done, the present report is preliminary, the
main purpose being to put on record a new species of the
distinctive ralline genus Nesotrochis. There appear to be other
undescribed rails and new birds in the collections, but anal-
ysis of these will have to await completion of the sorting of
specimens and associating them with their often scanty and
scattered data.
A REVIEW OF THE GENUS NESOTROCHIS WETMORE
Nesotrochis debooyi WETMORE
This large flightless rail was first described by Wetmore
(1918) from aboriginal kitchen middens in the Virgin Islands.
It was later found in middens and cave deposits in Puerto Rico.
The following list gives the locality, references and location of
specimens for each of the known occurrences of the species
(key to abbreviations in acknowledgments ).
38—Proc. Biou. Soc. WAsH., Vou. 87, 1974 (439)
440 Proceedings of the Biological Society of Washington
Puerto Rico
Cueva Clara near Morovis Wetmore, 1922a AMNH
Cueva San Miguel near Morovis Wetmore, 1922a AMNH
Cueva Torano near Utuado Weimore, 1922a AMNH
Hacienda Jobo near Utuado Wetmore, 1922a AMNH
Midden at Barrio Canas, ca. 2.4 Wetmore, 1938 YPM,
km E of Ponce USNM
Rio Piedras Specimens examined by Olson MCZ
Cave at Ciales Specimens examined by Olson MCZ
Shell heap at Aguirre Specimens examined by Olson MCZ
Virgin Islands
St. Thomas; midden at Magen’s Bay Wetmore, 1918; USNM
( type-locality ) Brodkorb, 1974 and UF
St. Croix; midden at mouth of Salt River Wetmore, 1918 USNM
St. Croix; midden ca. 2.4 km E of Wetmore, 1937 USNM
SW Cape
St. Croix; midden near Christiansted Wetmore, 1927 USNM
St. John; midden at Francis Bay Brodkorb, 1974 UF
PVirgin Gorda (based on hearsay ) Nichols, 1943
The skeletal elements known for this species include the
femur, tibiotarsus, tarsometatarsus, humerus and fragments of
pelvis. It has been suggested that the Puerto Rican birds may
have been larger than those of the Virgin Islands (Wetmore,
1938; Brodkorb, 1974), but there is not sufficient material to
confirm this, especially since there appears to be marked sexual
dimorphism in size. The not unlikely possibility that West
Indian aborigines may have transported this species from one
island to another must also be entertained.
N. debooyi apparently was an important food item of the
aborigines of Puerto Rico and the Virgin Islands, and it prob-
ably persisted there at least until late in the last century. Wet-
more (1927) attributed to this species legends of a bird called
the “carrao” which hunters in Puerto Rico prior to 1912 used to
run down with dogs. The name “carrao” is now applied to
the Limpkin (Aramus guaruana), a wary bird of strong flight
that is unlikely to have been captured by dogs. To Wetmore’s
account may be added the report of Nichols (1943), who was
told by the natives of Virgin Gorda that in the past “flightless
waterhens” had been common and many had been killed
with sticks.
Fossil rails from West Indies 44]
TaBLE 1. Measurements (mm) of limb bones of the three species of
Nesotrochis. Those of N. picapicensis from Fischer and Stephan (1971).
N. debooyi N. picapicensis N. steganinos
n range mean n range n range mean
Length
tarsometatarsus 6 71.4— 75.5 73.2 2 62.0-63.2 1 ————— 59.5
Length
tibiotarsus 13 104.6-124.0 117.5 2 88.0-96.2
Distal width
tibiotarsus 1 AS 1322) 1273 6 9:3=10;0 3) 8:4— 97-92.
Length femur 8 72.3- 80.3 76.7 2 58.5-68.3 _—W—W————
Length humerus 3 58.3— 63.7 61.2 4 49.0-50.0 4 43.2-49.2 46.9
Length ulna $$ So] — 33,8
Nesotrochis picapicensis (Fischer and Stephan)
Fischer and Stephan (1971) described three new species of
birds, including two rails, in Pleistocene deposits from Pio
Domingo cave, near Sumidero, Pinar del Rio, Cuba. Their
paper fails to compare or even discuss many of the Antillean
taxa with possible affinity to their fossil species. Consequently,
their generic assignments are incorrect.
The flightless rail that Fischer and Stephan named Fulica
picapicensis was based on femora, tibiotarsi, tarsometatarsi,
humeri, pelves, a few vertebrae, a proximal end of an ulna!
and a partial cranium. A humerus was designated as the
holotype. I have had only their illustrations, description and
tables for comparison, but these indicate clearly that picapi-
censis shows no more than a familial resemblance to the coots
(Fulica). Furthermore, the broad distal end of the tarsometa-
tarsus, with widely flared trochleae, particularly the inner
one, is a feature peculiar among rails to Nesotrochis and in-
dicates, along with the other skeletal elements illustrated, that
picapicensis certainly belongs in Nesotrochis rather than in
Fulica. Nesotrochis picapicensis differs from N. debooyi in
its much smaller size (Table 1), more pointed intercotylar
knob and in the heavier and straighter shaft of the humerus.
No doubt more differences could be discerned upon com-
parison of actual specimens of N. picapicensis.
1 This is not from a rail and is possibly columbid,
442 Proceedings of the Biological Society of Washington
Nesotrochis steganinos, new species
Figures 1, 2
Holotype: Complete right tarsometatarsus USNM 205609 (Fig. 1).
Collected in Pleistocene cave deposits (Cave I in Miller's notes) near
St. Michel de L’Atalaye, Département de L’Artibonite, Haiti, Hispaniola,
in 1925 by G. S. Miller.
Measurements of holotype: Overall length 59.5 mm, proximal width
9.9, proximal depth 10.1, least width of shaft 4.3, least depth of shaft
2.7, width through trochleae 10.3, depth through trochleae 6.7.
Paratypes: Incomplete left tarsometatarsus (USNM 205610), three
left humeri (205676, 205687, 205691), two proximal ends of left humeri
(205689, 205690), two distal ends of right humeri (205684, 205677),
left ulna (205688), two distal ends of left tibiotarsi (205678, 205685 ),
two distal ends of right tibiotarsi (205679, 205680), right femur lack-
ing distal end (205681), incomplete left femur (205682), shaft of right
femur of juvenile (205683). These came from the same group of caves
as the holotype, details of which are discussed by Miller (1922, 1929).
A left humerus (205686) of this species was found in a tray of speci-
mens with no data. It is almost certainly from one of the Haitian col-
lections. Matrix adhering to it is similar to that on specimens labelled
“San Raphael”—a locality about ten miles north of St. Michel.
Etymology: Gr. steganos, through a covered passage or tube + inos,
sinew—in reference to the closed hypotarsal canals.
Diagnosis and description: Clearly referable to the genus Nesotrochis
on the basis of the expanded distal end of the tarsometatarsus with
widely flaring trochleae and particularly the more anterior and distal
placement of the inner trochlea. Much smaller than N. debooyi and
averaging somewhat smaller than N. picapicensis (Table 1).
Tarsometatarsus with ossified ring below internal cotyla as typical
of most rails. Distinctive in having a sheet of bone completely enclosing
both of the tendinal canals on the internal side of the hypotarsus (Fig.
lc, e), whereas in all other known rails, except Paraortygometra porzan-
oides from the Miocene of France, the posterior canal is open. In
medial view (Fig. lc) the area anterior to the hypotarsus is more deeply
excavated, with the result that the ridge descending along the shaft from
the external cotyla is much narrower and more distinct than in N.
debooyi. In proximal view (Fig. le) the internal cotyla is deeper and
much more distinctly set off from the hypotarsus than in N. debooyi,
and the intercotylar knob and the depression posterior to it are both
better developed. In anterior view the tarsometatarsus differs from
N. debooyi and N. picapicensis in that the internal cotyla and the proxi-
momedial part of the shaft below it do not project as abruptly medially
beyond the line of the shaft, and the distal foramen is relatively smaller.
The tendinal bridge of the tibiotarsus is not higher on the internal
side in N. steganinos as it is in N. debooyi, and the opening distal to
Fossil rails from West Indies 443
Fic. 1. Nesotrochis steganinos, new species, holotype tarsometatarsus,
USNM 205609: a, anterior view; b, posterior view; c, medial view;
d, lateral view; e, proximal view; f, distal view; a—d natural size; e and
f, twice natural size.
the bridge is not slanted (Fig. 2d). Both of these species differ from
N. picapicensis in having this opening smaller and more constricted.
The few tibiotarsi of N. steganinos differ considerably in size.
Conformation of the humerus in N. steganinos shows clearly that
the species was flightless. The humerus is shorter and heavier than in
N. debooyi and more similar to that of N. picapicensis, but differs from
both of these species in having the entepicondyle produced much farther
distally and internally. There appear to be two distinct types of humeri
in N. steganinos—larger ones with a shallow elongated brachial de-
pression (Fig. 2a) and smaller ones with a very deep, rounded brachial
depression (Fig. 2b). These at first seemed so disparate that I took
them to be from entirely different species. However, they are alike in
all but these two respects and one of the specimens (205691) is some-
what intermediate. It seems best, therefore, to refer all of these humeri
to N. steganinos, since there is no other indication of the presence of two
species of Nesotrochis in the deposits. The size differences are in accord
with observed size differences in the hindlimb, while differences in the
brachial depression are possibly indicative of a sexual dimorphism that
involved more than size.
444 Proceedings of the Biological Society of Washington
Fic. 2. Nesotrochis steganinos, new species, paratypes: a-c, wing
elements (upper row, anconal view; lower row, palmar view); a, hu-
merus, USNM 205676; b, humerus, USNM 205687; c, ulna, USNM
205688; d, distal end of tibiotarsus, USNM 205678. All figures natural
size.
The ulna referred to N. steganinos (Fig. 2c) appears to be the only
specimen of that element of Nesotrochis yet recognized. It is short and
markedly curved, with a much reduced olecranon and a distinctly flat-
tened internal surface of the shaft.
The remains of N. steganinos were found in deposits containing great
quantities of bones of large extinct rodents and other mammals. These
Fossil rails from West Indies 445
accumulations of bones are attributed to predations of the gigantic ex-
tinct barn owl, Tyto ostologa Wetmore (1922b), bones of which occur
in the same deposits. The owl must also have taken the flightless rails
from time to time.
On the basis of the elements so far known, particularly the tarsometa-
tarsus, the genus Nesotrochis appears to be quite distinct from other
genera of rails. Its affinities are uncertain. Wetmore (1922a) at first
considered Nesotrochis to be an offshoot of the primitive South American
wood-rails of the genus Aramides, but later he felt it to be closer to the
gallinules (1937). I fully concur that Nesotrochis is not related to
Aramides, while a gallinule-like ancestor is a possibility. The discovery
of a complete skull of any of the species of Nesotrochis would probably
shed much light on the affinities and adaptations of the genus.
OTHER Fosst RAILS FROM THE WeEsrT INDIES
Cyanolimnas cerverai Barbour and Peters
The rail Cyanolimnas cerverai was first described from skins
obtained in the Zapata Swamp of Cuba (Barbour and Peters,
1927) and all subsequent non-fossil specimens have come from
this same vicinity. I have elsewhere discussed the relation-
ships of the genus (Olson, 1973). The species has generally
been regarded as flightless (e.g. Ridgway and Friedmann,
1941), although Bond (1942) reported that he saw one flutter
about ten feet across a canal, and that another observer had
also seen a bird fly for short distances. On morphological
grounds Cyanolimnas cerverai would be classed as a flightless
species, since the pectoral girdle and wing are as reduced as
in other species of rails that are considered to be flightless
(iige3))):
In addition to describing “Fulica” (= Nesotrochis) picapi-
censis, Fischer and Stephan (1971) named another, smaller
rail represented in the fossil material from Pio Domingo cave
in Pinar del Rio as Rallus sumiderensis. They had a very large
series of this species and the morphology of the wing and
pectoral girdle indicated to them that it was flightless. If
they were aware of the existence of Cyanolimnas cerverai,
they gave no indication of it in their paper.
In the USNM is a collection of bones from a cave deposit
in the Caballos Mountains on the Isle of Pines that contains
a large series (at least 40 individuals) of a rather small rail
446 Proceedings of the Biological Society of Washington
Fic. 3. Body skeleton of Cyanolimnas cerverai, AMNH 3930. Note
the reduced pectoral girdle and carina, and the obtuse coraco-scapular
angle typical of flightless rails.
with similarly reduced wings and pectoral girdle. The various
elements agree well with illustrations of “Rallus sumiderensis”
in Fischer and Stephan (1971). In Table 2 these specimens
are compared with “Rallus sumiderensis” and with the scanty
comparable material of Cyanolimnas from the Zapata Swamp
(tarsal measurements from skins and a single body skeleton).
There is close agreement between the three groups except in
the tarsal measurements. Published measurements of tarsus
length in “Rallus sumiderensis’ do seem to be somewhat
smaller than in the Zapata Swamp specimens but there is over-
lap with the Isle of Pines material.
The single available femur of Cyanolimnas cerverai from
the Zapata Swamp is generally similar to specimens from the
Isle of Pines but is somewhat more slender and curved, with
a heavier, more rounded head that is not as markedly under-
cut on the anteroventral side. This is not a large enough sam-
ple to establish that there were differences between the pop-
Fossil rails from West Indies 447
TABLE 2. Lengths of limb bones (mm) of three samples of
Cyanolimnas cerverdai.
Pinar del Rio* Isle of Pines Zapata Swamp
n range n range mean n range mean
Tarsometatarsus 39 32.0-38.3 17 35.4-42.6 39.1 8 39.5-46.7 42.2’
Femur 2 41.5-44.2 15 42.1-50.6 466 1 ————— 48.1’
Coracoid 1 — 19.3 2 17.7-19.6 18.7. 1 ————— 19.6’
Humerus 26 32.4-36.7 18 32.7-38.1 35.2
Ulna 1 — 264 15 22.8-29.6 26.1
Carpometacarpus 1 — 17.5 £2 18.0-18.2 18.1
* “Rallus sumiderensis’ from Fischer and Stephan (1971).
1 Measured from skins.
2 From body skeleton AMNH 3930.
ulations of the Zapata Swamp and the Isle of Pines. While
recognizing that further study may disclose temporal or geo-
graphic variation within the species, I feel that it is best to
consider the Isle of Pines specimens and “Rallus sumiderensis”
as belonging to Cyanolimnas cerverai, until better material
becomes available.
Barbour (1928) was of the opinion that Cyanolimnas cer-
verdi originated in the Zapata Swamp. It is now evident that
this was not the case. Two other distinctive birds, the finch
Torreornis inexpectata and the wren Ferminia cerverai, were
also originally described from the Zapata Swamp. Later, a
well-marked new race of Torreornis was discovered on the
arid coast of extreme southeastern Cuba (Spence and Smith,
1961). It thus seems fairly certain that all three of these spe-
cies were formerly more widely distributed in Cuba and that
the present populations are relicts.
Fulica podagrica Brodkorb
The species described by Brodkorb (1965) as Fulica podag-
rica from the Pleistocene of Barbados requires further study;
after a perfunctory examination of the type material, I be-
lieve that it is probably a composite. The holotype humerus
is clearly that of a coot and seems to differ from available
specimens of Fulica americana and F. caribaea only in its
slightly heavier shaft and the slightly more proximal position
448 Proceedings of the Biological Society of Washington
of the ectepicondyle. Historical specimens of coots taken on
Barbados were mentioned by Brodkorb (1965) as possibly
belonging to F. podagrica, but he was unable to locate them.
They were later discovered by Phillips (1967) who identified
them as Fulica caribaea, thus establishing that this species
once occurred on the island.
Most, if not all, of the leg elements assigned to F. podagrica,
upon which most of the diagnosis of that species was based,
appear to be from a wholly different rail that was larger and
heavier than indicated by the type humerus of F. podagrica.
The distal part of a tarsometatarsus (UF 7468), for example,
has a wider shaft, narrower trochleae, smaller and more proxi-
mally located distal foramen, and a longer and deeper scar for
the hallux than does Fulica, and it evidently does not pertain
to that genus.
Through an unfortunate mixup, the specimen Brodkorb
(1965) gives as “Fulica leucoptera’ is actually Gallinula chlor-
opus garmani. Since much of the original characterization of
F. podagrica rested on comparisons with this specimen, there
is further cause to reappraise F. podagrica.
ACKNOWLEDGMENTS
This paper is dedicated to Alexander Wetmore, the father
of Antillean paleornithology, who called to my attention much
of the type material of Nesotrochis steganinos. For loan of
specimens and for information concerning them I am indebted
to Pierce Brodkorb and S. David Webb, University of Florida
(UF); Malcolm C. McKenna and Dean Amadon, American
Museum of Natural History (AMNH); Raymond A. Paynter,
Jr.. Museum of Comparative Zoology (MCZ); and Fred C.
Sibley, Yale Peabody Museum (YPM). I thank William B.
Robertson for directing me to the Nichols reference. The
manuscript was criticized by Pierce Brodkorb, Robert J. Emry,
John Farrand, Jr., Clayton E. Ray, and Alexander Wetmore.
The photographs are by Victor E. Krantz.
RESUMEN
Se hacen breves consideraciones acerca de la especie extinta
de ralido no volador Nesotrochis debooyi, de Puerto Rico y
Fossil rails from West Indies 449
las Islas Virgenes, y se afaden nuevas localidades a su dis-
tribucién. Fulica picapicensis Fischer y Stephan, hallada en
cuevas pleistocénicas del oeste de Cuba, se asigna al género
Nesotrochis. Se describe una nueva especie, Nesotrochis steg-
aninos, encontrada en depdsitos cavernicolas de la Espanola.
La especie fosil Rallus sumiderensis Fischer y Stephan, des-
crita en base de restos procedentes de cuevas del oeste de
Cuba, es la especie viviente Cyanolimnas cerverai, y otros
fosiles hallados en la Isla de Pinos, Cuba, se asignan tambien
a esta misma especie que hasta ahora se conocia sdlo de la
Ciénaga de Zapata, Cuba. E] material tipo de Fulica poda-
grica Brodkorb, del Pleistoceno de Barbados, parece que
incluye dos especies y, por tanto, debe estudiarse con mas
detalle.
LITERATURE CITED
BaArsour, T. 1928. Notes on three Cuban birds. Auk 45:28-32.
, AND J. L. Perers. 1927. Two more remarkable birds from
Cuba. Proc. New England Zool. Club 9:95-97.
Bonp, J. 1942. Additional notes on West Indian birds. Proc. Acad.
Nat. Sci. Philadelphia 94:89-106.
Bropxors, P. 1965. Fossil birds from Barbados, West Indies. Jour.
Barbados Mus. Hist. Soc. 31:3-10.
—. 1974. Bird remains from pre-Columbian middens in the Vir-
gin Islands. Quart. Jour. Florida Acad. Sci. 35[1972]:239-240.
FiscHER, K., AND B. STEPHAN. 1971. Weitere Vogelreste aus dem
Pleistozan der Pio-Domingo-Hohle in Kuba. Wiss. Zeitsch.
Humboldt-Univ. Berlin, Math.-Nat. R. 20:593-607.
Miutiter, G. S. 1922. Remains of mammals from caves in the Republic
of Haiti. Smiths. Misc. Coll. 74(3):1-8.
——. 1929. A second collection of mammals from caves near St.
Michel, Haiti. Smiths. Misc. Coll. 81(9):1-30.
Nicuots, R. A. 1943. The breeding birds of St. Thomas and St. John,
Virgin Islands. Mem. Soc. Cubana Hist. Nat. 17:23-37.
Ouson, S. L. 1973. A classification of the Rallidae. Wilson Bull. 85:
381-416.
PuHituips, A. R. 1967. Some Antillean coots (Fulica) in the Cambridge
University and British Museums. Bull. Brit. Ornithol. Club 87:
35-36.
Ripcway, R., AND H. FrrepMAN. 1941. The birds of North and Middle
Amenica, WOl, @ Bull, UW. So Nai, Ming, BOs wa
SPENCE, M. J., AND B. L. Smiru. 1961. A subspecies of Torreornis
inexpectata from Cuba. Auk 78:95-97.
450 Proceedings of the Biological Society of Washington
Wermore, A. 1918. Bones of birds collected by Theodoor de Booy
from kitchen midden deposits in the islands of St. Thomas and
St. Croix. Proc. U. S. Nat. Mus. 54:513=522)
1922a. Bird remains from the caves of Porto Rico. Bull.
Amer. Mus. Nat. Hist. 46:297—333.
1922b. Remains of birds from caves in the Republic of Haiti.
Smiths. Misc. Coll. 74(4):1-4.
1927. The birds of Porto Rico and the Virgin Islands. Co-
lymbiformes to Columbiformes. Sci. Surv. Porto Rico and
Virgin Islands, Vol. 9, Part 3:243—406.
1937. Ancient records of birds from the island of St. Croix
with observations on extinct and living birds of Puerto Rico.
Jour. Agric., Univ. Puerto Rico 21:5-16.
1938. Bird remains from the West Indies. Auk 55:51—55.
mar. Sos
Vol. 87, No. 39, pp. 451-464 31 December 1974
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
TWO NEW AXIIDS (CRUSTACEA: DECAPODA:
THALASSINIDEA: CALOCARIS) FROM NORTH
CAROLINA AND THE STRAITS OF FLORIDA
By Austin B. WILLIAMS
National Marine Fisheries Service Systematics Laboratory,
National Museum of Natural History, Washington, D.C. 20560
Benthic biological sampling near the edge of the continental
shelf off Cape Lookout, North Carolina, during cruises of the
Duke University, NSF-sponsored, R/V Eastward has produced
a few specimens of a new species of axiid “lobster” belonging
to the genus Calocaris. A second new species, represented by
a single specimen from the western edge of the Bahama Bank,
was taken during deep-water shrimp exploration with a 40-foot
flat trawl in the Straits of Florida by the then U.S. Fish and
Wildlife Service, Bureau of Commercial Fisheries vessel Com-
bat. These species added to the four North Atlantic Axiidae
discussed by Boesch and Smalley (1972) reinforce their con-
clusion that the genus Calocaris as now constituted represents
more than one group.
Calocaris (Calastacus) jenneri, new species
Figures 1-10
Holotype male: Integument microscopically granulate. Carapace com-
pressed; nearly smooth, scattered rugosities anteroventrally on branchio-
stegite and cephalic part; almost straight along middorsal line in lateral
view except gradually deflexed from gastric region to tip of rostrum;
midline narrowly rounded and elevated posteriorly but not keellike;
cephalic groove obsolete ventrolaterally; branchial groove inconspicuous;
branchiostegite emarginate ventrally and posteriorly; scattered setae
overall but tufts anterior to branchial region, densest on gastric region.
Rostrum slightly exceeding eyes, tip upturned but no larger than any
of 3 asymmetrically spaced marginal spines on each side; with low
39—Proc. Biot. Soc. WAsH., VOL. 87, 1974 (451)
452 Proceedings of the Biological Society of Washington
"G “Tomoy ‘pF ‘g ‘T “teddn :wur % = sojeog ‘ueyz rey ‘fF ‘B9] Puoddes ‘EG SMOIA IOUUT
‘PeAOUI9I svJOS YPM vfoYyO JOUTUE FO sIoBuIZ ‘g ‘AorA [ereyey “T eyeut edAyojoy “wauual (snopyspjpD) siuv00j]09 “~—-[ ‘SOI
Two new axiids—Calocaris 453
median dorsal, interrupted keel reaching from level of midlength of
eyestalks to cephalic groove; ventrally with fairly deep median keel;
notched laterally at base where rostrum merges with gastric region
(orbit) for reception of eyestalks in erect position.
Gastric region with 2 dorsal carinae on each side of middorsal keel
more or less interrupted by spines and gaps; lateral pair interrupted at
about midlength by blunt spine, but continued to near base of rostrum,
ending in low ridge on left but a broken spine on right at base of rostrum;
row of scattered tubercles between middorsal keel and submesial carinae.
Abdomen and telson together about 1.5 times length of carapace (parts
of segments 4-6 broken). Segments smooth and almost uniformly arched
transversely (but with low lateral carina most prominent and oblique on
first segment, becoming obsolescent posteriorly ), bearing scattered clumps
of long, fine setae; first segment short, fitted for articulation with cara-
pace, with pleuron drawn posteroventrally to acute tip; pleura of second
to fifth segments with anteroventral margins broadly rounded, ventral
margin of second and third nearly straight, posterior corners of 2-5
nearly rectangular; sixth with broadly rounded pleuron.
Eyestalks almost cylindrical, slightly flattened mesiodorsally, cornea
well developed.
Antennular peduncle with second article almost reaching tip of rostrum.
Antennal peduncle with penultimate article extending beyond anten-
nular peducle; first (coxal) article with a short ventral spine; strong
dorsolateral spine of second article reaching proximal border of cornea;
antennal thorn (acicle) slender, curved, exceeding cornea and reaching
distal border of penultimate article of antennular peduncle; third article
with slender mesioventral spine reaching about ™% length of fourth
article; terminal article about °%4 length of fourth article; flagellum
reaching beyond chelipeds, but broken.
Endopods of third maxillipeds pediform, densely setiferous ventrally
and mesially; distal article unarmed; merus with 3 ventral spines pro-
gressively larger and more acute distad; ischium with 2 small spines on
ventral margin proximally, and along mesial margin a prominent crest
of about 17 strong spines terminating in distally advanced spine curved
mesad; coxa with ventral spine.
First legs strong, chelate, asymmetrical; with numerous tufts of con-
spicuous, silky, plumose setae obscuring fingers on both inner and outer
surfaces, becoming progressively shorter to crest of merus on outer sur-
face but leaving inner proximal part of hand bare; submarginal row of
setae on inner face of carpus and ventral border of merus. Right chela
strongest, fingers stout; fixed finger nearly straight, crushing edge with
small proximal tooth and larger bilobed tooth distally; dactyl curved, tip
closing inside fixed finger, strong tooth opposing bilobed tooth of fixed
finger and a stronger tooth proximal to it; crest of propodus with few
spines and tubercles, inner face with patch of strong tubercles near base
of fixed finger; carpus with sharp dorsal spine at articulation with hand;
454 Proceedings of the Biological Society of Washington
Fics. 5-6. Calocaris (Calastacus) jenneri, major (right) chela of
holotype male: 5, external view; 6, internal view. Scale = 5 mm.
merus with upper margin keeled, low tooth near crest and row of 5
spines on proximoventral margin. Left chela similar except more slender
and shaped as cutter; fingers agape proximally, distal half with opposed
edges straight and finely serrate; prominent triangular tooth at middle
of fixed finger diverging outward from cutting edge, opposite it an in-
ternal shoulder on dactyl; low rounded teeth on both fingers near artic-
ulation of dactyl; externally, and nearly concealed in pubescence, a
prominent row of coalesced tubercles and spines parallel to proximal
margin of fixed finger near articulation of dactyl; tubercles at bases of
setal tufts on inner palm proximal to fixed finger.
Second legs chelate (left missing), flattened, merus with 3 unequal
ventral spines; fingers corneous-tipped, cutting edges with fine spinules
most dense distally, arranged full length of fixed finger, only on distal
half of dactyl; dense tufts of setae on fingers and longer tufts of setae
on margins elsewhere.
Two new aie @ aloes 455
Fics. 7-10. Calocaris (Calastacus) jenneri: 7, anterior carapace in
dorsal view, holotype male; 8, sternite of first abdominal segment and
adjacent parts, holotype; 9, sternite of first abdominal segment, allotype
female; 10, appendices interna and masculina with part of second pleo-
pod, paratype male (UNC-IMS 2484). Scales: horizontal, 7, 8, 9 =
2 mm; vertical, 10 = 0.2 mm.
Third and fourth legs similar, flattened; dactyls slightly curved, a
comb of corneous spinules laterally, 3rd with 5 on left, 6 on right, 4th
with 6, tips corneous; propodus with combs of corneous spinules ventro-
laterally, 3rd with 7, 4th with 8 on left, 9 on right, and 3 or 4 smaller
accessory combs laterally; scattered setae on all articles.
Fifth legs smaller in diameter as well as length, cylindrical; dacty]
curved, corneous-tipped, with spinules laterally and on propodus distally;
scattered setae longest and densest distally.
Biramous, setose pleopods on abdominal segments 2—5, lacking on first
segment; endopod of each with appendix interna equipped with hooked
setules at tip; second pleopod with an appendix masculina bearing long
non-plumose setae on distal third (appendix missing from left side).
456 Proceedings of the Biological Society of Washington
Tail fan with dense, long, plumose setae on caudal margin and tufts
elsewhere. Telson longer than broad, with submesial pair of thickened
longitudinal ribs; caudal margin evenly rounded, lateral margins with
proximal lobate process articulating with mesial branch of uropod and
interrupted distally by 3 progressively more closely spaced spines; sub-
marginal spine each on marginal process, at base of first marginal spine
(movable), mesial to it on submesial rib (fixed), and an unequal pair
near distolateral corners (movable). Uropods as long as telson. Mesial
branch with thickened longitudinal ribs bearing 4-5 spines, lateral mar-
gins with 2—4 spines progressively longer distally. Lateral branch with
transverse distal suture bordered by row of strong, fixed spines; sub-
marginal movable spine at distolateral corner; 2 thickened longitudinal
ribs, lateral one broadest and bearing scattered small spines; lateral mar-
gins with 2-4 spines and slight unevenness distally.
Allotype female: Much as holotype in general appearance but differ-
ing as follows: Rostrum with asymmetrical marginal spines, 2 on left,
1 plus rudiment on right. Ridging and tuberculation of gastric area less
pronounced. Both chelae somewhat less setose and less tuberculate.
Major chela with dactyl closing completely inside fixed finger, tips cross-
ing; fixed finger with a prominent tooth near midlength and 3 low ones
proximally, shearing edge distally with 1 low broad tooth; dactyl with
broad, low proximal tooth, a notch distal to it followed by smooth shear-
ing edge. Minor chela with fingers separated by narrower proximal
diastema, distal shearing edges of fingers faintly serrate; single sharp
spine on external surface of palm proximal to articulation of dactyl.
Small uniramous pair of pleopods on sternite of first abdominal segment;
pleura of 4th and 5th segments more acutely tipped.
Measurements in mm: Holotype ¢. Carapace: length including ros-
trum, 12.4; width, 6.3; height, 6.3. Length articles of legs, L = left,
R = right:
IR 2R 3L 4L 5L
dactyl 6.9 1.9 1S 1.4 1.1
propodus 14.4 4.3 3.8 4.1 3.8
carpus 4.4 4.3 3.6 3.3 2.6
merus 6.5 7.0 6.9 5.0 3.5
total 32.2 ies 15.8 13.8 11.0
Allotype @. Carapace: length including rostrum, 10.5; width, 4.4;
height, 4.6. Right chela: length dactyl, 4.0; propodus, 7.9.
Type-locality: E Cape Lookout, N. C., 34°18’N, 76°01.2’W-34°17.1’N,
76°01.3’W, 100-85 m, Eastward Stn. 12885, Cape Town dredge, 3
October, 1969, C. E. Jenner and class.
Material studied: Specimens studied are confined to the type series
deposited in collections of the National Museum of Natural History
(USNM), Washington, D.C., and University of North Carolina Institute
of Marine Sciences (UNC-IMS), Morehead City, N. C.
Two new axiids—Calocaris 457
USNM 150472 holotype ¢; USNM 150474 allotype 92°, paratype ¢
(cl 7.5 mm [est., rostrum broken], cl 8.4 mm), SW Cape Lookout, N. C.,
33°43.N, 76°40.7’W-33°43.4’N, 76°40.3’W, 100 m, Eastward Stn.
1089, small biological trawl, 27 April, 1965, Menzies. UNC-IMS 2484
paratype ¢ and @ (cl each 11.3 mm), E Cape Lookout, N. C., 34°43’N,
76°40’W-33°42.7’N, 76°40.2’W, 90-110 m, Eastward Stn. 1087, small
biological trawl, 27 April, 1965, Menzies. USNM 150473 paratype ¢
(cl 9.6 mm), SSW Cape Lookout, N. C., 33°43.3’N, 76°42’W-33°44'N,
76°41’W, 91-98 m, Eastward Stn. 10789, Cape Town dredge, 4 No-
vember, 1968, J. and W. Vernberg.
Variation: Ommamentation (setae, tubercles, spines) increases in size
and complexity with increasing body length. In one male (USNM
150473) the proximal tooth of the major propodal finger is fairly well
developed, but other teeth are rudimentary. The diastema on fingers
of the minor chela is also well developed in this specimen, but it is
not pronounced in smaller specimens examined. The hands of this male
are reversed, the left stronger, but there is no evidence of alteration in
the minor chela to indicate regeneration. The rostrum may be symmetri-
cally or asymmetrically toothed.
Color in life: Ground color “flesh” to off-white (pinkish white) with
longitudinal trending, cross-banded spots of orange-red. Most dense
orange-red color on posterior part of carapace where red is also deepest
in intensity. Abdomen with first segment pale, others progressively
darker caudad; lightest areas at articulations dorsally, deepest colors at
bases of pleura. Basal articles of antennules and antennae with bands
of color equal in intensity to darker parts of abdomen. Color on legs
more washed out. Pubescence light greenish tan. (From holotype. )
Name: This species is named in honor of Professor Charles E. Jenner,
Department of Zoology, University of North Carolina, Chapel Hill, who
has contributed much to knowledge of marine invertebrates of the Car-
olinas and who with one of his classes collected the holotype.
Calocaris (Calastacus) oxypleura, new species
Figures 11-18
Holotype male: Integument microscopically granulate. Carapace com-
pressed; bearing a sparse, short pubescence in tufts originating from
rather evenly scattered, nearly obsolescent tubercles, longer setae on
gastric region and in grooves; almost straight along middorsal line in
lateral view except rostrum very slightly deflexed; most broadly rounded
dorsally in gastric region, narrowing posteriorly; midline slightly raised,
almost keellike in short section near posterior margin but interrupted
at cervical groove, posterior part of gastric region and middle of pos-
terior carapace; cephalic groove deepest dorsally but evident throughout
length; branchial groove obsolescent but forming broad, shallow de-
pression.
458 Proceedings of the Biological Society of Washington
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‘WW T = ZI “IomMoy ‘wu Z = ET ‘TT
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Two new axiids—Calocaris 459
Rostrum slightly exceeding eyestalks, tapered and narrow; edges up-
turned; tip slightly damaged but apparently ending in an uptumed spine
approximately equal to pair of adjacent marginal spines, behind these
a pair of slightly larger spines at midlength, and at broadening base a
still larger pair; marginal notch (orbit) at base for reception of eye-
stalks in erect position where rostrum merges with gastric region.
Gastric region with 2 interrupted carinae on each side of raised mid-
line; midline with moderate anterior spine followed by obsolescent spine;
each lateral carina originating in diverging rostral base and armed with
3 buttressed spines, 1 above ocular notch, 1 behind notch, and 1 remote
posterior spine; each submesial carina consisting of 3 remote buttressed
spines in nearly straight line; area between carinae smooth except for
scattered tufts of setae.
Abdomen and telson together about 1.5 times length of carapace
(parts of abdominal segments 2-6 badly broken); segments smooth and
uniformly arched dorsally, first short, fitted for articulation with cara-
pace; low discontinuous longitudinal carina laterally between segmental
articulations; pleura of segments 1-5 drawn posteroventrally to acuminate
tips and with uniform margins except as follows: anterior margin of
first segment slightly sinuous, of long second segment quite sinuous,
of segments 3—5 with single spine in middle; sixth segment with pleuron
subrectangularly produced, a tiny blunt spine anterior to tip of pleural
lobe.
Eyestalks slightly flattened mesiodorsally, left slightly longer than
right; cornea dilated.
Antennular peduncle with second article reaching anteriorly to same
level as tip of rostrum (estimated rostral length).
Antennal peduncle with penultimate article extending beyond anten-
nular peduncle; first (coxal) article with short, slender, terminal ventral
spine; strong dorsolateral spine of second article reaching beyond distal
margin of cornea; antennal thorn (acicle) slender (bent unnaturally)
exceeding antennular peduncle; third article with small, distal, mesio-
ventral spine; terminal article % length of penultimate fourth article;
flagellum missing.
Endopods of third maxillipeds pediform, densely setiferous ventrally
and mesially, a few long setae on carpus and merus laterally; distal
articles unarmed; carpus with small distoventral spine laterally; merus
with 3 ventral spines progressively larger and more acute distad; ischium
with 2 small spines on ventral margin, and along mesial margin a prom-
inent crest of about 16 spines terminating in an enlarged distal spine
curved mesad; coxa with ventral spine.
First legs strong and asymmetrical, chelae elongate. Each rather
thinly clothed with tracts of conspicuous, tufted, long setae, thickest
on dactyls (but nearly as thick on fixed finger of right chela), dorsal
aspect of palms, external surface of carpus and distal exposed part of
merus; sparser but equally long tufts on inner surface of palms (except
central inflated part), in row along groove on ventrolateral margin of
460 Proceedings of the Biological Society of Washington
ae
Fics. 14-16. Calocaris (Calastacus) oxypleura, chelae of holotype
male in external view: 14, left; 15, right; 16, right merus. Scale =
So mm.
hands, and on ventral edge of merus; tufts more evenly distributed on
right than on left chela. Each hand with: row of 4 nearly equidistant
spines on dorsal margin of palm and laterally displaced one near articu-
lation of dactyl; tracts of irregularly sized tubercles on both inner and
outer surfaces of palms, distally near base of fixed finger (one spiniform
tubercle on each surface of right chela much larger and sharper than
remainder) and trailing posteroventrally, especially on inner surface;
low but sharp ridge along ventrolateral margin, sharpest proximally but
diminishing to obsolescence near tip of fixed finger. Right chela slender,
fingers agape nearly throughout length but tips broken; dactyl stouter
than fixed finger, its prehensile edge with fine but uneven serrations
most dense distally; fixed finger with prominent triangular tooth at %
Two new axiids—Calocaris 461
Fics. 17-18. Calocaris (Calastacus) oxypleura, holotype male: 17,
appendices interna and masculina with part of second pleopod; 18, an-
terior carapace in dorsal view. Scales: upper, 18 = 2 mm; lower, 17 =
0.25 mm.
length diverging outward from prehensile edge, proximal to it a row of
widely spaced, fairly evenly distributed tuberculiform teeth, and distal
to it a row of fine, irregular but much more closely spaced teeth. Left
chela stouter, palm moderately inflated; dactyl stouter than fixed finger;
462 Proceedings of the Biological Society of Washington
opposed edges of fingers with central diastema, proximal %4 of fingers
armed with irregularly elongate sectorial teeth and weak molar complex
on fixed finger, distal half of both with shearlike edges, tips broken.
Carpus of each cheliped with 4 strong dorsal and 4 ventrolateral spines.
Merus of right cheliped with 5 spines on inner crest and 1 laterally near
articulation with carpus, left merus with 4 spines on inner crest; both
meri with 2 spines on external crest (left proximal one broken). Each
ischium with single curved spine on inner crest.
Remaining legs missing except for disarticulated, chelate left second;
chela nearly bare on inner surface but clothed with long, dense setae
elsewhere, most prominent on dactyl and along ventrolateral aspect;
fingers bent ventrally at slight angle to palm, opposed edges spooned;
dactyl about % length of palm; both carpus and merus slightly bent to
fit curve of carapace, prismatic in cross section; distal third of carpus
with long setae ventrally; external surface of merus covered with ciliated,
transverse rugosities.
Biramous, setose pleopods on abdominal segments 2-5, lacking on
first segment; endopod of each with appendix interna equipped with
hooked setules at tip; second pleopod with an appendix masculina bearing
long nonplumose setae along entire length, most dense distally.
Tail fan with dense, long, plumose setae on caudal margin (many
broken) and scattered tufts elsewhere. Telson about ™%4 longer than
wide, evenly rounded caudally; lateral margins with proximal lobate pro-
cess articulating with uropodal endopods and armed with 4 spines, 1
surmounting lobate process, 2 well deparated on middle reaches, and
1 at posterolateral corner; mesial to latter an unequal pair of movable
spines; median longitudinal sulcus fairly broad; suggestion of a pair of
submesial thickened longitudinal ribs in proximal half and on these
a small submesial pair of proximal spines, larger spine near center on
each side. Uropods as long as telson. Mesial branch with thickened
longitudinal ribs, central rib bearing 4 nearly equidistant spines, caudal-
most one nearly on terminal margin; lateral marginal rib with 4 spines
progressively longer distally and strong spine at caudolateral angle (spine
twinned and broken on right side). Lateral branch with transverse distal
suture bordered by row of small spines; 2 thickened longitudinal ribs
centrally; lateral margin with 4 unevenly spaced spines, distalmost
flanked mesially by submarginal movable spine.
Measurements in mm: Carapace: length including rostrum, 18.8;
width, 6.1; height, 7.0. Length abdomen and telson (estimate), 23.
Length articles of first legs: right propodus, 21; right dactyl, 14.4; left
propodus, 19.8; left dactyl, 10.0—of second leg: dactyl, 0.94; propodus,
2.94; carpus, 6.25; merus, 7.31.
Type-locality: [Straits of Florida W of Riding Rocks] 25°15’N,
79°13’W, 365 m, Combat Stn. 445, 23 July, 1957 (USNM 101651 holo-
type 3).
Name: The name is from the Greek “oxys” acute, and “pleura” side,
for the abdominal pleura with prominent acuminate tips.
Two new axtids—Calocaris 463
Remarks: I have described this species with some hesitation because
the imperfect holotype is the only specimen, but the characters are so
distinctive that there seems little likelihood that it could be confused
with another species. Specimens of most axiids are rare in collections,
and knowledge of the group seems benefitted by any evidence available.
The specimen is well preserved but damaged mechanically. The rostral
tip and tip of each antennal thorn have seemingly been jammed and
pushed to the right. A number of spines on the carapace and appendages
are broken, as are many setae. The legs present are disarticulated from
the body and tips of the chelipeds broken off. The abdomen has been
crushed and twisted, most damage inflicted on segments 4—6, splitting
the terga and cracking some pleura. The lateral branch of the left
uropod is missing.
The large chelae, though apparently worn, are distinctive in shape.
The left chela is the major, and the right the slender, forcepslike minor.
No other minor chela resembles this one among published illustrations
of Axiidae or specimens available for study. The hand may be re-
generated, but there is no evidence for or against this.
Discussion: In the western Atlantic, 3 species of Calocaris belonging
to the subgenus Calastacus are now known from waters off the south-
eastern United States, the 2 described here and C. hirsutimana Boesch
and Smalley, 1972, known also from Guyana. All have conspicuously
bearded chelae. Boesch and Smalley (1972) found the gills of C.
hirsutimana as listed for Calocaris by Gurney (1942: 149) except with
“podobranch and arthrobranch on second maxilliped well developed and
epipod of second pereiopod lacking a podobranch.” The large male
specimen of C. hirsutimana listed below agrees with Gurney’s gill formula
except that legs 3 and 4 have epipods and that of 3 has a small podo-
branch. Calocaris jenneri has the same gill formula as C. hirsutimana;
danger of breakage prevented examination of gills in C. oxypleura.
Calocaris jenneri is much less spinose on the lower palm of the chelae
and along the edges of the pleura than C. hirsutimana, and it lacks the
elongate antennal thorn and median spine on the telson; but C. jenneri
does have much more prominently armed tooth rows and _ accessory
spines on the chelae in both males and females (females of C. hirsut-
imana are unknown). While the long antennal thorn and somewhat
setose carapace link C. oxypleura to C. hirsutimana, the acuminate tips
on the pleura, open forcepslike chelae, and lack of a median spine on
the telson clearly separate it from the latter. Boesch and Smalley (1972)
drew parallels between Axiopsis and Calastacus, pointing out that ir-
regularities in the middorsal keel make it a poor generic character, and
related C. hirsutimana to 2 Indo-Pacific species of Axiopsis, implying
thereby some importance in the long antennal thorn. Calocaris jenneri
has a short antennal thorn yet shares other features with C. hirsutimana.
It is clear that present generic definitions are inadequate.
New locality record: USNM 150475 Calocaris (Calastacus) hirsut-
464 Proceedings of the Biological Society of Washington
imana, 1 @ (mature, broken), Tortugas shrimp grounds [Florida], fisher-
man, April, 1964, to D. Tabb, to A. J. Provenzano.
Acknowledgments: I am indebted to D. M. Dexter, C. E. Jenner,
R. J. Menzies, and J. and W. Vernberg for gifts of specimens, to Fenner
A. Chace, Jr., and Isabel Pérez Farfante for critical reading of the
manuscript, and to Maria M. Diéguez for preparing the llustrations.
The holotype, allotype and a series of paratypes of Calocaris jenneri
were transferred to USNM from UNC-IMS lots 2376, 2564 and 2579.
Addendum: After this paper was in press a second male specimen
was found in material from the then U. S. Fish and Wildlife Service,
Bureau of Commercial Fisheries R/V Oregon, Stn. 6460, 14°12’N,
81°57.5°W [Quita Sueno Bank off Nicaragua], 146 m, 2 December
1967, 66-foot trawl, by Michelle de Saint Laurent. The specimen agrees
well with the holotype, is considered a paratype (USNM 151211), and
differs in minor details as follows: Chelipeds and left fifth leg at-
tached, remainder of legs missing. Carapace length 18.5 mm, extreme
tip of rostrum broken, spines of rostrum asymmetrically arranged. Ab-
dominal pleura with a few small submarginal tubercles laterally. Chelae
with tracts of spines and/or spiniform tubercles on both inner and outer
surfaces of palms in same position as tracts of tubercles in holotype;
left (major) chela with tips of fingers each tapered to a point and curved
toward each other, crossing when closed; right (minor) chela with sim-
ilar tip on dactyl, tip of fixed finger broken; left carpus with 3, right
with 4 ventrolateral spines; meri each with 5 spines on inner crest. Fifth
leg slender; brush of long setae on dactyl; propodus with a few long,
mesial setae on distal end, a dense patch of short setae terminally on
lower margin, and an elongate patch of longer ones distolaterally; length
of dactyl 1.3, propodus 4.1, carpus 3.2, merus 4.4, ischium 2.5 mm.
Tail fan with dense fringe of long, plumose setae largely intact, minor
differences from holotype in spination.
LITERATURE CITED
BorescH, D. F., anp A. E. SMALLEY. 1972. A new axiid (Decapoda,
Thalassinidea) from the northern Gulf of Mexico and tropical
Atlantic. Bull. Mar. Sci. 22(1):45-52.
Gurney, R. 1942. Larvae of decapod Crustacea. Ray Society, Lon-
don, No. 129:306 pp.
NS
) GE ANSO. E
Vol. 87, No. 40, pp. 465-472 31 December 1974
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
CICHLASOMA REGANI, A NEW SPECIES
OF CICHLID FISH FROM THE
RIO COATZACOALCOS BASIN, MEXICO
By Rosert RusH MILLER
Museum of Zoology, The University of Michigan,
Ann Arbor, Michigan 48104
In 1957 and 1959, collections were made in the upper part
of the Rio Coatzacoalcos basin near the crest of the Isthmus
of Tehuantepec, Oaxaca, México. One paper on cichlid fishes
collected there (Miller and Nelson, 1961) gave the specific
traits of a seemingly localized endemic of the Thorichthys
group, Cichlasoma callolepis (Regan). The present paper
describes another localized endemic, this time of the Theraps
group, which I am pleased to name for the late British ichthy-
ologist, C. Tate Regan, in recognition of his classic studies
on American cichlids.
Study materials are in the collections of the Museum of
Zoology of The University of Michigan (UMMZ), the National
Museum of Natural History (USNM), and the Museum of
Natural History of The University of Kansas (KU). The lat-
eral-series scale count begins with the first scale in the upper
lateral line, goes to the last one in that line, then continues
with the scale on the lower lateral line that is next behind
the transverse row that extends downward and forward from
the last pored scale of the upper lateral line. The last scale
recorded is at the end of the lower lateral line, where it meets
the base of the caudal fin (this is not necessarily the last pored
scale, since pored scales often extend onto the caudal fin).
This method is equivalent to Regan’s “longitudinal series,”
as redefined by Trewavas (1935: 72), and is employed to
40—Proc. Bion. Soc. WaAsH., VOL. 87, 1974 (465 )
466 Proceedings of the Biological Society of Washington
make use of his extensive scale-count data on American cich-
lids (Regan, 1905a—c, 1906a—b). The transverse-series scales
are counted, as described by Regan, from the base of the first
dorsal spine downward and backward to, but not including,
the scale row on the midline of the abdomen. Fin-ray counts
and measurements are made according to the methods pre-
scribed by Hubbs and Lagler (1958).
Cichlasoma regani, new species
Figure 1
Cichlasoma sp. Miller (1966: 793), upper part of Rio Coatzacoalcos
basin, Atlantic slope of Isthmus of Tehuantepec, México.
Diagnosis: A moderately small, rather slender-bodied, large-scaled
species of the Theraps group of Regan (1906-08: 17), distinguished
from other members of this group by lacking both clearcut vertical bars
and a horizontal stripe on the side of the adult, with 32 (31-33) scales
in lateral series, typically XVII,13 elements in dorsal and VI,9 in anal
fins, 3 + 8 or 9 gill rakers, rather somber life colors, and the adult
with 2 or 3 blotches along upper side below dorsal fin and a prominent
dark spot at caudal base.
Holotype: UMMZ 184756 (adult female, 80.5 mm S. L.), from Rio
Almoloya where crossed by the Trans-Isthmian Highway about 33 km
north of southern terminus of that road, at 95° 01’ W Long., 16° 45’ N
Lat., Oaxaca, México, taken 25 February 1959 by R. R. Miller and R. J.
Schultz; elevation approximately 250 meters.
Paratypes: UMMZ 184757 (11, 36-67 mm), same data as holotype.
UMMZ 178529 (male, 37 mm), same locality as holotype, 27 March
1957, R. R. and M. Miller. USNM 102255 (male, 95 mm), from Rio
Malotengo, upper tributary Rio Coatzacoalcos, 6.4 km _ southeast of
Matias Romero, Oaxaca, 21 December 1935, T. MacDougall. KU 1573-
1574 (2 females, 116 and 145 mm), from 60 km southwest of Jesus
Carranza (Rio Jaltepec drainage), Oaxaca, 26-30 March 1948, W. W.
Dalquest; elevation, 150 meters.
Description: Form and color pattern are shown in Figure 1 and me-
ristic data are given below. Proportional measurements appear in
Table 1.
Dorsal spines, XVI (1), XVII (15); dorsal soft rays, 12 (1), 13 (12),
14 (3); anal spines, V (1), VI (15); anal soft rays, 9 (16); pectoral
rays in both fins (splint along uppermost ray excluded), 15 (11), 16
(21). Lateral scales, 31 (2), 32 (13), 33 (1); scales in upper lateral
line, 20 (1), 21 (5), 22 (6), 23 (4), and in lower lateral line, 9
(1), 10 (3), 11 (5), 12 (5), 13 (2); scale-row overlap of the upper
lateral line on the lower, 2 (2), 3 (3), 4 (5), 5 (4), 6 (2); transverse
scales, 18 (1), 19 (3), 20 (7), 21 (5); lateral line to soft dorsal origin
A new cichlid fish from México 467
Fic. 1. Cichlasoma regani, holotype, UMMZ 184756, female, 80.5 mm
standard length, from Rio Almoloya, Oaxaca, México.
(not including scales on scaly sheath along fin base), 2% (6), 3 (7),
3% (3); lateral line to origin of anal fin, 8 (4), 9 (10), 10 (2); scale
rows between bases of pectoral and pelvic fins, 5 (7), 6 (8), 7 (1);
scales around caudal peduncle, 8+9=19 (8), 9+9= 20 (6); scale
rows on cheek, 4 (7), 5 (9). Gill rakers on first arch, upper limb,
Sm les) a4 @2))-nlower limb 6) (6). 9) (0) totaly” 15). 12 lors
(1). Vertebrae, 13 (1), 14 (14), 15 (1) precaudal; 16 (3), 17 (12),
18 (1) caudal; and 30 (2), 31 (14) total.
Body slender, elongate, becoming deeper with age, predorsal contour
strongly arched in marked contrast to the gently sloping (concave)
preventral contour. Highest point along back at or near origin of dorsal
fin. Base of dorsal fin increasing in length with age. Mouth horizontal,
low (well below level of ventral rim of orbit), lower lip thicker than
upper (from about half to slightly greater than half diameter of pupil).
Upper jaw extends to vertical from a point between nostril and anterior
rim of orbit; jaws equal anteriorly or lower slightly included. Fold of
lower lip not continuous, but frenum narrow. Dorsal origin above in-
sertion of pectoral fin, spines increasing in length rapidly to about fifth
to eighth, then more gradually to last (longest) which varies from
slightly less than half (in juveniles) to half (in adults) head length;
soft part of fin when laid back extending from anterior fourth to half
of caudal fin. Pectoral shorter than head, extending from middle of to
well beyond midlateral spot or to between anus and anal-fin origin;
pelvic extending from anus to slightly beyond base of second anal
spine. Caudal fin rounded to slightly emarginate. Caudal peduncle
deeper than long. Depth of preorbital varying from about one-half
mgton
l Society of Wash
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Proceedings of the Biolo
468
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A new cichlid fish from México 469
diameter of orbit in juveniles to greater than orbit in large adults. Pre-
maxillary process extending backward between a point on posterior part
of snout to above middle of pupil. Along base of caudal rays one sup-
plementary lateral line, lying either above or below principal lateral
line, occurs in 6 juvenile-to-adults, and two supplementary lateral lines
occur in 2 half-grown, holotype, and 2 largest specimens; the other
types lack this feature.
Teeth: Jaw teeth conic, without a posterior cusp (present in Her-
ichthys), bluntly pointed (sometimes peg-like). Teeth of outer series
in upper jaw number 7 to 9 on each side, increasing rather regularly
in size anteriorly; in lower jaw anterior 4 outer teeth enlarged, subequal,
and sharply differentiated from smaller lateral ones. Occlusal surface
of lower pharyngeal bone of a 67-mm specimen with dentigerous por-
tion about 1.3 times broader than long, the teeth arranged in about 24
rows; those teeth in the two median rows, especially posteriorly, heavy
and rounded (molariform) but most of them bear remnants of a central
cusp, and those on rows on each side of these specialized (crushing)
teeth also enlarged.
Color pattern: Up to seven vertical bars (mostly rather obscure) in
young, four in juveniles, the broadest only about half diameter of eye,
the third broadest and darkest dorsally extending from below dorsal
fin nearly to venter or to anal fin, decreasing in number with age (4
in juveniles, 2-3 in small adults, 1-2 in large adults) and represented
in larger specimens chiefly by prominent dark spots or blotches, 2 or
3 in number, along upper lateral line (Fig. 1), the largest blotch below
10th to 14th dorsal spines and above tip of extended pectoral fin; the
upper lateral line runs through this blotch. Interradial membranes of
soft dorsal and caudal fins with rows of prominent dark spots, weakly
developed on soft anal fin. A prominent ovate to nearly round black
spot occurs on base of caudal fin and adjacent part of caudal peduncle,
extending over most of side of latter. Top of head with two prominent,
dark brown, curved saddles, anterior one across snout to anterodorsal
part of orbit and posterior one largely between eyes; anterior saddle
widest at its midpoint, about two-thirds diameter of pupil, whereas
posterior one narrowest there, about one-fourth to one-third diameter
of pupil.
Color in life: At time of capture, no bright colors were noted on
body or fins. Dorsal, caudal, and anal fins had rusty orange spots.
Horizontal rows of spots along side following scale rows were black,
each scale with a spot at or near its base, the rows becoming less prom-
inent toward abdomen. Body greyish olive to greenish to golden yellow
(on sides). Sexes not distinguished in field.
Habitat and Associates: The Rio Almoloya is a clear winding stream
of moderate gradient, about 6 meters wide on the average, with pools
attaining a maximum depth of 3 meters. At the time of our visits late in
the dry season it consisted of swift, rocky riffles alternating with long
pools and some sluice-like sections. Nearly a fourth of the stream was
470 Proceedings of the Biological Society of Washington
shaded with marginal vegetation along the steep banks. The bottom in-
cluded much sand and silt in the pools, alternating with rocks and
boulders on the riffles. There was also some bedrock. A green alga
and localized growths of Nasturtium were the only aquatic plants noted.
A fairly rich fish fauna was secured at this locality, including 18 species
in 8 families: Characidae (Astyanax fasciatus), Pimelodontidae (Rham-
dia guatemalensis), Belonidae (Strongylura hubbsi Collette, 1974),
Poeciliidae (Poecilia mexicana and P. sphenops, Heterandria bimacu-
lata, Poeciliopsis fasciata and P. gracilis), Atherinidae (Archomenidia
bolivari and Xenatherina schultzi), Mugilidae (Agonostomus monticola),
Cichlidae (Cichlasoma aureum auct., C. callolepis, C. bulleri, C. zona-
tum, C. salvini), and Eleotridae (Gobiomorus dormitor).
Distribution: Cichlasoma regani is known from the Rios Almoloya,
Malotengo, and Jaltepec, upper tributaries of the Rio Coatzacoalcos on
the Atlantic slope of the Isthmus of Tehuantepec.
Acknowledgments: I am grateful to the staff of the National Museum
of Natural History for facilities and working space, the John Simon
Guggenheim Memorial Foundation for support as a Guggenheim Fel-
low while preparing the manuscript, the U. S. National Science Founda-
tion (Grant GB-4854) for field support, and Louis P. Martonyi for
the photograph. Mexican officials kindly gave permission to collect
fishes in their country.
LITERATURE CITED
CoLLeTTE, B. B. 1974. Strongylura hubbsi, a new species of fresh-
water needlefish from the Usumacinta Province of Guatemala
and México. Copeia 1974(3):611-619.
Husps, C. L., anp K. F. Laciter. 1958. Fishes of the Great Lakes
region. Bull. Cranbrook Inst. Sci. 26:i-xi, 1-213.
Miniter, R. R. 1966. Geographical distribution of Central American
freshwater fishes. Copeia 1966(4):773-802.
, AND B. C. NEtson. 1961. Variation, life colors, and ecology
of Cichlasoma callolepis, a cichlid fish from southern Mexico,
with a discussion of the Thorichthys species group. Occ. Pap.
Mus. Zool., Univ. Mich. No. 622:1-9.
REGAN, C. T. 1905a. <A revision of the fishes of the South-American
cichlid genera Crenacara, Batrachops, and Crenicichla. Proc.
Zool. Soc. London, 1905, pt. 1:152-168.
—. 1905b. A revision of the fishes of the South-American cichlid
genera Acara, Nannacara, Acaropsis, and Astronotus. Ann.
Mag. Nat. Hist. ser. 7, 15:329-347.
—. 1905c. A revision of the fishes of the American cichlid genus
Cichlosoma and of the allied genera. Ann. Mag. Nat. Hist.
ser. 7, 16:60—-77, 225-243, 316-340, 433-445.
—. 1906a. A revision of the South-American cichlid genera
Retroculus, Geophagus, Heterogramma, and Biotoecus. Ann.
Mag. Nat. Hist. ser. 7, 17:49-66.
A new cichlid fish from México A471
——. 1906b. A revision of the fishes of the South-American cichlid
genera Cichla, Chaetobranchus, and Chaetobranchopsis, with
notes on the genera of the American Cichlidae. Ann. Mag.
Nat. Hist. ser. 7, 17:230-239.
1906-08. Pisces. In: Biol. Cent.-Amer. 8:i—xxxii, 1—203.
Trewavas, E. 1935. A synopsis of the cichlid fishes of Lake Nyasa.
Ann. Mag. Nat. Hist. ser. 10, 16:65-118.
472 Proceedings of the Biological Society of Washington
Be 7: C67 OD
Vol. 87, No. 41, pp. 473-484 31 December 1974
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
STUDIES OF NEOTROPICAL CADDISFLIES, XIX:
THE GENUS CAILLOMA (TRICHOPTERA:
RHYACOPHILIDAE)
By OLiver S. FLINT, Jr.
Smithsonian Institution, Washington, D.C., 20560
All of the 17 recognized Neotropical genera of Rhyacophil-
idae belong to the subfamily Hydrobiosinae, a subfamily which
also occurs in southern and eastern Asia, Fiji, Australia, Solo-
mon Islands, New Caledonia, and New Zealand. The larvae
of all species are free-living, and, as far as is known, predaceous
and rheophilic.
One genus, Atopsyche, is known from the southwestern
United States, south throughout Central and South America,
including the Greater Antilles, and as far south as the Province
of Catamarca in Argentina. Yet this genus has not been col-
lected, and probably does not occur, in Chile or southern
Argentina. Conversely, 14 of the 16 remaining genera are
strictly limited to these southernmost regions of Chile and
Argentina. Of the remaining 2 genera, Iguazu is known from
outside Chile by only 1 specimen labelled Misiones, Argentina,
that may be mislabelled. This genus, then, is most likely
limited also to the Chilean Subregion. The other genus, Cail-
loma, actually has a distribution that broadly overlaps the
two Neotropical subregions. Examples of this genus have been
collected near the eastern tip of Isla de los Estados (Staten
Island) off the tip of Tierra del Fuego, and north along the
Andes at least into Ecuador. Why Atopsyche, whose larvae
may occur on the same rock as those of Cailloma in northern
Argentina, has not been able to accomplish the same bridging
of the two Subregions, remains unanswered.
41—Proc. Bio. Soc. Wasu., Vou. 87, 1974 (473)
474 Proceedings of the Biological Society of Washington
I have frequently collected adults and immatures of this
genus in my travels in Chile, especially during my most re-
cent visit in Argentina in 1973. In fact, a special trip was
taken to Mendoza for the primary purpose of collecting ad-
ditional material of the unrecognizable Cailloma lucidula
(Ulmer), the results of which are summarized in the discus-
sion under this species. As a result I now possess larvae and
pupae, males and females, of all 3 known species of this genus.
This paper, then, brings together illustrations, keys, distribu-
tions, and new synonymies in the genus Cailloma.
I wish to thank Prof. Dr. H. Weidner for the loan of the
type of Cailloma lucidula (Ulm.), and Dr. J. Unzicker for
the loan of the type of C. brunosa Ross & King. Mr. L. M.
Druckenbrod prepared the excellent illustrations of the larval
heads and pronota.
Genus Cailloma Ross & King
Cailloma Ross & King, 1951, p. 507.—Ross, 1951, p. 112; 1953, p. 154;
1956, p. 125.—Flint, 1971, p. 12.—Fischer, 1971, p. 149.
Genus A Flint, 1963, p. 463; 1971, p. 12.
Type species: Cailloma brunosa Ross & King (= lucidula Ulmer).
Adult: Spurs 2,4,4. Venation nearly identical in both sexes, except
that the discal cell in the forewing of the female is typically longer than
in the male. Forewing with Re,s branched well beyond s, Ras branched
at s; Mie and Ms, both branched about midway to wing margin. Hind-
wing with Ress apparently arising from R: with cell Cus» considerably
inflated. Abdomen without lateral processes; seventh sternum of both
sexes with a small wart bearing enlarged setae. Male genitalia with
cercus small, filicercus elongate, paracercus short; clasper one-segmented;
aedeagus short, simple, with a single internal spine. Female genitalia
not produced into an elongate ovipositor.
Larva: Mandibles with a single tooth on dorsal carina, ventral carina
lacking on right mandible, a simple ridge on left. Head %4 as wide as
long. Prosternum with sclerite small, % to ¥% as wide as distance be-
tween legs. Foreleg with apicoventral process of femur short and
slender; tibia and tarsus fused. Mid and hindlegs similar in size and
structure; claw with basal seta short, broad, and curved. Lateral sclerite
of anal proleg 11% times as long as broad; claw without ventral teeth,
but with basoventral seta broad and curved.
Pupa: Mandibles curved; inner margin serrate with larger teeth at
midlength. Hook plates anteriorly on segment 2 to 6 or 7, not raised
Neotropical caddisflies—Cailloma AT5
above body surface, each with many small teeth; posterior plates on
segments 4 and 5, each broader than long, low, and with many small
teeth.
Recognition: The adults of the genus are easily recognized by their
venation, especially the short apical forks of Ress, Mii2, and Maia in the
forewing and the apparent origin of Res from Rs in the hindwing. The
other genera that may be confused with Cailloma on the basis of vena-
tion are Parachorema, Stenochorema, and Nolganema. The males of
the first 2 genera possess a narrow reflexed costal cell and lack all
vestiges of crossveins in the forewing. In addition, the claspers in the
males of Parachorema are clearly 2-segmented and in Stenochorema
possess a basal complex. The genus Nolganema Navas is not fully
recognizable from the original description, but clearly falls into this
complex of genera. Navas does state however, that the type possesses
sternal processes, which are lacking in Cailloma but present in some de-
gree in the other genera.
The larvae of few Neotropical Hydrobiosines have been described,
but based on undescribed material in the collection here, the following
characteristics will probably serve for recognition of the larvae of
Cailloma: presence of a single dorsal tooth and reduction of ventral
cutting edges of mandibles; shape and structure of forelegs, especially
fusion of tibia and tarsus; very small size of prosternal sclerite; modifica-
tion of basal setae of mid and hind legs; and short lateral plate of anal
prolegs, whose claw lacks ventral teeth and possesses a modified basal
seta.
The larvae described as genus A from Ecuador in 1963 are clearly
those of a species of Cailloma. They are in an early instar, at which
time the pronotal halves are fused and the head spots discrete. Early
instar larvae in any collection that I examined have the appearance of
these from Ecuador, thus they cannot be identified to species at this
time.
In general, characteristics of the mandibles, hook plates, and apical
lobe offer the best characteristics for recognition of the pupae whenever
recognition has been possible.
Biology: Immatures of the species in this genus are commonly found
on and under the rocks in cold, rather small streams in the Andes. C.
lucidula (Ulm.) is a dominant, and at times exclusive, inhabitant of
small, tumbling, brooks of the puna grassland areas of Tucuman and
Mendoza, Argentina (and probably similar areas in the other north-
western provinces). Dr. Spangler informs me that the stream near
Cochabamba, Bolivia, was 1 to 2 meters wide, with riffles but at a
rather low gradient and also in the puna. According to Dr. Hodges,
the two Ecuadorian localities where small larvae were taken were also
of similar nature and in the paramo, a similar low-type vegetation zone
of high elevations.
However, many of the collection sites in southern Chile and Argentina
are in the Nothofagus forest, which characteristic is especially true for
476 Proceedings of the Biological Society of Washington
C. rotunda Flint. Two sites in southern Chile near Punta Arenas are
most instructive in this connection. One site, collected on Jan. 10, 1965
was at an elevation where the forest was well developed with the small
stream, slightly over a half meter wide, flowing in and alongside the
forest. The site collected on Feb. 26, 1966 was at a slightly lower ele-
vation, in open grassland. At the first site only C. rotunda Flint was
taken, at the second only C. pumida Ross. Yet, one night in central
Chile at the fish hatchery at Rio Blanco males of all three species came
to the light. There are, however, many springs and streams up to sev-
eral meters in width in the vicinity, some of which are shaded by trees
planted at the fish hatchery, others in the more open, natural scrub-
type vegetation. It is thus still possible for some degree of ecological
difference in the breeding sites at this one locality.
KEY TO SPECIES, ADULTS
1. With claspers and aedeagus (males) __.__---- ee 2
Without these structures (females) — 4
2. Clasper broadest subapically, apex nearly round ____. rotunda Flint
Clasper narrowing apicad, apex bluntly pointed _ 3
3. Apicoventral lip of aedeagus produced into a long narrow point
NUN a UE RUR STAR Daal MVS Ua RNG Ss SE lucidula (Ulmer)
Apicoventral lip of aedeagus broad and bluntly bifid __. pumida Ross
4, Midventral membranous process of ninth sternum greatly surpass-
ing ‘apex of ninth jteroum) 20 ee ee rotunda Flint
Midventral process only attaining apex of ninth tergum _________
ar aN eel ee Ds SN lucidula (Ulmer) & pumida Ross
KEY TO SPECIES, LARVAE
1. Head with frontoclypeus unmarked anteriorly __._..... rotunda Flint
Head with frontoclypeus bearing a dark transverse bar or with
dark lateral spots anteriorly ____.. lucidula (Ulmer) & pumida Ross
KEY TO SPECIES, PUPAE
1. Hook plate 6A about twice as wide as long; without 3 large apico-
lateral setae on apical lobe —_.. rotunda Flint
Hook plate 6A about as wide as long; apical lobe with 3 large setae
apicolaterally. ee lucidula (Ulmer) & pumida Ross
Cailloma lucidula (Ulmer)
Figures 1-4, 12-17, 19, 21-96
Atopsyche lucidula Ulmer, 1909, p. 73; 1913, p. 404.—Jorgensen, 1919,
p. 392.—Fischer, 1960, p. 161.
Cailloma lucidula (Ulm.).—Ross & King, 1951, p. 507.—Ross, 1956,
p. 125.—Fischer, 1971, p. 149.
Neotropical caddisflies—Cailloma A477
Fics. 1-9. Cailloma lucidula (Ulmer): 1, male genitalia, lateral; 2,
aedeagus, lateral; 3, clasper, ventral; 4, wings. C. pumida Ross: 5, male
genitalia, lateral; 6, aedeagus, lateral; 7, clasper, ventral. C. rotunda
Flint: 8, male genitalia, lateral; 9, clasper, ventral.
Cailloma brunosa Ross & King, 1951, p. 507.—Ross, 1956, p. 125.—
Fischer, 1971, p. 149. [NEW SYNONYMY/].
Cailloma angustipennis Schmid, 1955, p. 122.—Fischer, 1971, p. 149.
—Knutson & Flint, 1971, p. 315. [NEW SYNONYMY].
478 Proceedings of the Biological Society of Washington
Through the kindness of Prof. Dr. Weidner, I have been able to clear
and study the type female of A. lucidula Ulmer. It is absolutely typical
of the complex containing this species and C. pumida Ross, the females
of which I am unable to tell apart. I visited the type locality of Po-
trerillos, Mendoza, Argentina within a few days of the day of the year
of the collection of the type in the hope of collecting additional ma-
terial which would permit the resolution of this problem. I found larvae
and pupae commonly in the small streams above Potrerillos, and ob-
tained adults at the light on several nights. All the male metamorpho-
types and adult males, save one, pertain to the species previously known
as either C. brunosa Ross & King or C. angustipennis Schmid. There is
one adult male of C. pumida Ross in the collection. Therefore, on the
overwhelming preponderance of this species at the type locality near
the date of collection of the type, I identify the type of A. lucidula with
the species here treated.
I have studied the type of C. brunosa Ross & King, loaned by the
Illinois Natural History Survey and returned to the Field Museum of
Natural History (FMNH), and C. angustipennis Schmid, located at the
National Museum of Natural History (USNM). They are clearly con-
specific with C. lucidula, which shows little variation over its extensive
range.
This species and the following, C. pumida Ross, are separable with
certainty only in the male sex; the females, larvae, and pupae seem
to be identical in the two species. The aedeagus, which offers the surest
means of recognition, has its apicoventral lip drawn out into a long
slender point in C. lucidula, but in C. pumida the lip is broader and
bluntly bifid. In addition the clasper is broader apically and in dorsal
aspect bears a smooth, slightly cupped dorsomesal area in C. lucidula.
Material; PERU: San Ignacio, Cailloma, 1 Sept. 1939, K. P. Schmidt,
6 holotype of C. brunosa Ross & King (FMNH).
BOLIVIA: CocHasamsa; 48 km north of Cochabamba, 8490’ (2570
m), 10 May 1969, P. & P. Spangler, 45 larvae, 4 prepupae, 7 pupae,
1g 22 metamorphotypes.
ARGENTINA: TucumMan; Rt. 307, La Angostura, 1850 m, 11 Oct.
1973, O. S. Flint, Jr., 1¢ 29, 1 larva. Rt. 307, km 76, west of Tafi
del Valle, 2600 m, 11 Oct. 1973, O. S. Flint, Jr., 1 larva, 5 prepupae,
6 pupae, 1¢ metamorphotype. Rt. 307, Abra de Infiernillo, 2970 m,
11 Oct. 1973, O. S. Flint, Jr., 17 larvae. Camino a Amaicha (Rt. 307)
km 92, 31 Oct. 1964, Teran, 19 (Inst. Lillo). Rt. 307, km 97.5, Los
Cardones, 2700 m, 11 Oct. 1973, O. S. Flint, Jr., 1 prepupa, 5 pupae,
26 12 metamorhpotypes.
MeEnpoza; Potrerillos, 26 Dec. 1907, P. Jorgensen, 2 holotype of A.
lucidula Ulmer (Hamburg Mus.). 4 km south of Potrerillos, 1450 m,
18 Dec. 1973, C. M. & O. S. Flint, Jr., 20 larvae, 2 prepupae, 7 pupae,
26 12 metamorphotypes. Rio Blanco, 10 km south of Potrerillos, about
1500 m, 18 Dec. 1973, C. M. & O. S. Flint, Jr., 9 larvae, 6 prepupae,
Neotropical caddisflies—Cailloma 479
16 7
Fics. 10-18. Cailloma rotunda Flint: 10, female genitalia, lateral;
11, female genitalia, ventral. C. lucidula (Ulmer): 12, female genitalia,
lateral; 13, female genitalia, ventral; 14, apical lobe of male pupa, dorsal;
15, apical lobe of female pupa, dorsal; 16, pupal mandible, dorsal; 17,
pupal hook plates, dorsal. C. rotunda Flint: 18, pupal hook plate 6A,
dorsal. (A = anterior, P = posterior, 2-7 = segment number).
480 Proceedings of the Biological Society of Washington
16 pupae, 6¢ 592 metamorphotypes. Above El Salto, southwest of Po-
trerillos, about 1600, m, 20 Dec. 1973, C. M. & O. S. Flint, Jr., 21¢
49, 10 larvae, 16 prepupae, 25 pupae, 3¢ 42 metamorphotypes.
CHILE: Cogumpo; Las Hedionditas, 10 Jan. 1966, L. E. Pena G.,
1é 29. Aconcacua; Rio Blanco, 10 Mar. 1968, Flint & Pena, 3¢é.
SANTIAGO; E] Alfalfal, 29 Feb. 1968, Flint & Pena, 1¢. El Manzano,
26 Oct. 1951, L. E. Pena G., ¢ holotype C. angustipennis Schmid
(USNM).
Cailloma pumida Ross
Figures 5-7, 27
Cailloma pumida Ross, 1956, p. 125.—Schmid, 1958, p. 190.—Flint,
1967, p. 45.—Knutson & Flint, 1971, p. 315.—Fischer, 1971, p. 149.
Cailloma erinaceus Schmid, 1957, p. 382; 1958, p. 190.—Fischer, 1971,
p. 149.
Cailloma pumida Ross is very similar to C. lucidula (Ulm.); indeed,
I am able to tell only the males apart. However, I have collected males
of both these as well as C. rotunda Flint together, indicating an ab-
solute sympatry and validity of the species. The characters differentiat-
ing males of the former two species are discussed under C. lucidula.
Material. ARGENTINA: Menpoza; Rio Blanco, 8 km south of
Potrerillos, 18 Dec. 1973, C. M. & O. S. Flint, Jr., 1¢. NEuQuEN; Rio
Quilquihué, at Quilquihué, 26 Jan. 1974, O. S. Flint, Jr., 5¢ 29. Rio
Quilquihué, at Lago Lolog, 22-23 Jan. 1974, O. S. Flint, Jr., 2 pre-
pupae. Arroyo Cordoba Grande, at Caleufu, 3 Feb. 1974, O. S. Flint,
Jr., 1 larva, 1 prepupa. Rio NrEcro; Arroyo Nireco, near San Carlos de
Bariloche, 5 Mar. 1957, J. Illies, 446; same, but 31 Dec. 1957, 34; same,
but 3 May 1958, L. E. Pena G., 546 29; same, but 26 May 1958, 1¢;
same, but 17 June 1958, 146; same, but 15 Oct. 1958, 1¢. Lago Nahuel
Huapi, near Arroyo Nireco, San Carlos de Bariloche, 25 Sept. 1957, J.
Illies, 1¢, 14 metamorphotype. Rio Nirihuau, at Estacion Nirihuau,
11 Feb. 1974, O. S. Flint, Jr., 1 ¢ 19. Brook 5 km south of Rio Villegas,
7 Feb. 1974, O. S. Flint, Jr., 5 larvae, 1 ¢ 19 metamorphotypes. Cascada
Mallin Ahogado, near El] Bolsdn, 9 Feb. 1974, O. S. Flint, Jr., 26,
392, 2 larvae. CHusput; Rio Epuyen, near Hoyo de Epuyen, 10 Feb.
1974, O. S. Flint, Jr., 19, 4 larvae, 2 prepupae.
CHILE: Cogurmso; Los Molles, 1 Oct. 1967, L. E. Pena G., 146
19. Canela Baja, 23-24 Oct. 1961, L. E. Pena G., 14. Rivadavia, 16
May 1952, L. E. Pena G., 1¢. Aconcacua; Los Andes, 12 Nov. 1957,
J. Illies, 1¢. Rio Blanco, 10 Mar. 1968, Flint & Pena, 1¢. SANtTrAGO;
El Alfalfal, 29 Feb. 1968, Flint & Pena, 26. O’Hicci1ns; La Leonera,
26-28 Dec. 1954, L. E. Pena G., 26 59 including ¢ holotype 9 allo-
type of C. erinaceus Schmid (USNM). Nuss; Las Trancas, 10-11 Feb.
1956, L. E. Pena G., 16. CHiLoxE; Ancud, 6 Jan. 1952, L. E. Pena G.,
16. MAGALLANEs; Tributary to Rio de las Minas, at water supply in-
Neotropical caddisflies—Cailloma 481
Fics. 19-25. Cailloma lucidula (Ulmer): 19, larval head and pro-
notum, dorsal. C. rotunda Flint: 20, larval head and pronotum, dorsal.
C. lucidula (Ulmer): 21, larval mandibles, dorsal; 22, larval foreleg,
posterior; 23, larval hindleg, posterior; 24, larval anal proleg, lateral;
25, larval prosternite, ventral.
482 Proceedings of the Biological Society of Washington
Fics. 26-28. Recorded distribution of: 26, Cailloma lucidula (Ulmer);
27, C. pumida Ross; 28, C. rotunda Flint.
take above Punta Arenas, 26 Feb. 1966, O. S. Flint, Jr., 1 larva, 9 pre-
pupae, 66 pupae, 3¢ metamorphotypes.
Cailloma rotunda Flint
Figures 8-11, 18, 20, 28
Cailloma rotunda Flint, 1967, p. 46—Knutson & Flint, 1971, p. 315.
This recently described species is only distantly related to the other
two species of Cailloma. The male is easily recognized by the very
broad and rounded claspers and the differently shaped paracercus and
aedeagus. The female genitalia also are diagnostic, especially the much
longer mesal process from the ninth sternum and the general shape and
structure of the vagina and associated supports. The larva is less cer-
tainly determined, but lacks all dark marks anteriorly on the fronto-
clypeus, which marks appear to be present in some degree on the other
species. The pupa of rotunda is easily recognized by the reduction or
loss of the hook-plate on 7A and the general increase in size of the
other plates, as well as the loss of the 3 pairs of long setae on the apical
lobe.
Material; ARGENTINA: Tierra DEL FuEco; Bahia Buen Suceso,
23-26 April 1971, Flint & Hevel, 1¢ 29. Isla de los Estados, Bahia
Crossley, 26-30 April 1971, Flint & Hevel, 2 prepupae, 17 pupae, 1¢
12 metamorphotypes. Isla de los Estados, Bahia Capitan Canepa, 1-3
May 1971, Flint & Hevel, 5 larvae, 1 prepupae, 1¢ 19 metamorpho-
types. Isla de los Estados, Bahia Blossom, 10 May 1971, Flint & Hevel,
Neotropical caddisflies—Cailloma 483
1 prepupa. Isla de los Estados, Puerto Ano Nuevo, 19 May 1971, Flint
& Hevel, 1 larva, 5 pupae, 1¢ metamorphotype.
CHILE: Aconcacua; Rio Blanco, 10 Mar. 1968, Flint & Pena, 1¢.
SANTIAGO; FE] Alfalfal, 29 Feb. 1968, Flint & Pena, 14; same, but 12—
13 Oct. 1969, Flint & Barria, 19. Near Los Valdes, 4000 m, 25 Mar.
1958, J. Illies, ¢ holotype (USNM). Near Banos Morales, 2000 m, 25
Mar. 1958, J. Illies, 1¢. Los Valdes, tributary to Rio Volcan, 7 Mar.
1958, J. Illies, 4 larvae. Nusie; 2.7 km north of Las Trancas, 27 Jan.
1967, Irwin & Stange, 26. MAGALLANEs; Tributary to Rio de las Minas,
14 km north of Punta Arenas, 10 Jan. 1966, Flint & Cekalovic, 13 larvae,
8 prepupae, 13 pupae, 3¢ 32 metamorphotypes.
LITERATURE CITED
Fiscuer, F. C. J. 1960. Necrotaulidae, Prosepididontidae, Rhyacophil-
idae. Trichopt. Catalog. I:1-168.
1971. Supplement to Vol. I and IJ. Trichopt. Catalog. XII:
1-311.
Furnt, O. S., Jr. 1963. Studies of Neotropical Caddis Flies, I: Rhy-
acophilidae and Glossosomatidae (Trichoptera). Proc. U.S.
Nat. Mus. 114(3473):453—478.
1967. Studies of Neotropical Caddis Flies, II: Trichoptera
Collected by Prof. Dr. J. Illies in the Chilean Subregion. Beitr.
Neotr. Fauna 5:45-68.
1971. Studies of Neotropical Caddisflies, XII: Rhyacophil-
idae, Glossosomatidae, Philopotamidae, and Psychomyiidae
from the Amazon Basin (Trichoptera). Amazoniana 3:1-67.
JORGENSEN, P. 1919. Los Trichopteros Argentinos. Reun. Nac. Soc.
Argent. Cienc. Nat. 1916:389-399.
Knutson, L., anp O. S. Fuint, Jr. 1971. Pupae of Empididae in pupal
cocoons of Rhyacophilidae and Glossosomatidae ( Diptera-
Trichoptera). Proc. Ent. Soc. Wash. 73:314—320.
Ross, H. H. 1951. The Origin and Dispersal of a Group of Primitive
Caddisflies. Evolution 5:102—-115.
—. 1953. On the Origin and Composition of the Nearctic Insect
Fauna. Evolution 7:145-158.
———. 1956. Evolution and Classification of the Mountain Caddis-
flies. University of Illinois Press, Urbana. 195 pages.
Ross, H. H., anp E. W. Kinc. 1951. A Key to the World Genera of
of the Caddisfly Tribe “Hydrobiosini’. Acta Zool. Lill. 12:
501-508.
ScHMip, F. 1955. Contribution a Ja connaissance des Trichoptéres
Neotropicaux. Mem. Soc. Vaud. Sci. Nat. 11:117—160.
— —. 1957. Contribution a étude des Trichoptéres Neotropicaux
II. Beitr. Entomol. 7:379-398.
—. 1958. Contribution a l’étude des Trichoptéres Neotropicaux
III. Mitt. Zool. Mus. Berlin 34:183-217.
484 Proceedings of the Biological Society of Washington
Utmer, G. 1909. Argentinisch Trichopteren. Z. wiss. Insekt Biol. 5:
(3-16) 120=194)
—. 1913. Verzeichnis der sudamerikanischen Trichopteren mit
bemerken ueber einzelne Arten. Deut. Entomol. Z. 1913:383-
414,
M4206 7S
Vol. 87, No. 42, pp. 485-492 31 December 1974
PROCEEDINGS
OF HIE
BIOLOGICAL SOCIETY OF WASHINGTON
A NEW SPECIES OF HYPSOPHRYS (DECAPODA:
HOMOLIDAE) FROM THE STRAITS OF FLORIDA,
WITH NOTES ON RELATED CRABS
By Austin B. WILLIAMS
National Marine Fisheries Service, Systematics Laboratory,
National Museum of Natural History, Washington, D.C. 20560
A crab taken in a 40-foot trawl during exploratory drags
for royal red shrimp, Hymenopenaeus robustus Smith, by the
then U.S. Fish and Wildlife Service, Bureau of Commercial
Fisheries, R/V Silver Bay SW of Dry Tortugas, Fla., on 8
June, 1959, belongs to an undescribed species and is the first
specimen of the genus Hypsophrys Wood-Mason, 1891, to be
reported from the Atlantic. The two other known species in
the genus, H. superciliosa Wood-Mason, 1891, and H. longipes
Alcock and Anderson, 1899, are confined to the Indo-Pacific
region (Alcock, 1900; Serene and Lohavanijaya, 1973). The
well-preserved animal shows an external feature on the che-
lipeds that is also evident in the previously known species,
represented by faded specimens preserved for over 4 of a
century. The terminology of Ihle (1913), revised by Serene
and Lohavanijaya (1973), is followed in describing features
on the carapace.
Genus Hypsophrys Wood-Mason, 1891
To Alcock’s (1900) definition of Hypsophrys the following should
be added: palm of each cheliped with smooth, oval spot on inner and
outer surface at base of fixed finger.
Hypsophrys noar, new species
Figures 1-12
Holotype male: Carapace ovate-oblong, tending toward quadrangular
in dorsal aspect, flattened and deep; distinctly subdivided into regions;
42--Proc. Bion. Soc. WASsH., VoL. 87, 1974 (485 )
486 Proceedings of the Biological Society of Washington
Fic. 1. Hypsophrys noar, dorsal view. x 0.5.
median raised regions glabrous; remainder of body clothed with rather
stiff pubescence of variable length and density.
Rostrum narrow, decurved, exceeding eyes, distinctly bifurcated,
each short tip continuous with raised margin in turn continuing on
carapace as raised orbital margin; reinforced ventrally by median keel
fused with interantennular septum, but distinguished from it by suture
line.
New homolid crab from Florida Straits A87
Orbital margins each interrupted by prominent intermediate spine
and stronger supraorbital spine, and ending ventrally near still larger
antennal spine.
Gastric region humped and prominent; obscurely subdivided into
central subregion flanked by larger lateral subregions; pair of epigastric
spines at each side diagonally in line with supraorbital spine, a low
tubercle anterolateral to each; an eminence behind and to each side of
rostral base surmounted by prominent tubercle with scattered smaller
tubercles on slopes.
Lineae homolica originating near edge of orbit above antennal spine,
passing around gastric region and continuing sinuously to disappear near
posterior margin of carapace. Deeply inscribed cephalic groove sep-
arating gastric region from posterior carapace and, ventral to linea
homolica on each side, separating hepatic region bearing 3 strong spines
from more posterior somewhat triangular region bearing strong dorso-
lateral spine and 2 tubercles in line below it. Post-cervical groove sep-
arating triangular from branchial region bearing strong posterolateral
spine (broken on right) and smaller spines and tubercles behind it
(more on right than left).
Eyes well developed.
Antennular peduncles with proximal article greatly inflated, much
more slender distal articles nearly cylindrical, terminal article shortest.
Antennae longer than carapace; antennal peduncle with proximal
article greatest in diameter, with spine at anterolateral corner; middle
article longer but more slender and terminal article less than half length
of middle one.
Pediform third maxillipeds lacking spines, setose, especially on pre-
hensile surfaces.
Chelipeds subequal, right slightly larger, elongate and spiny. Carpus
and chela about equal in length to merus-ischium. Palm irregularly
spiny, spines tending to be arranged in rows, lower border of palm with
fixed finger offset obliquely at angle of about 5-8°; an ocellated, hair-
less, smooth, oval spot at base of fixed finger on both inner and outer
surfaces (right chela, outer 3.50 x 4.25—inner 2.63 x 4.13 mm) having
central partially darkened portion surrounded by lighter, irregular hyaline
ring, and that in turn by wider white border. Fingers dark colored along
opposed edges and portion of other surfaces; curving inward from origins
to tips; opposed edges thin, sharp, and closely fitted with toothless
shearlike edges distally but with alternately meshing teeth proximally.
Fixed (propodal) finger with prominent, dark tooth near base and
smaller light colored tooth proximal to it on distal edge of palm; dactyl
longer than fixed finger and closing inside it, with subconical dark tooth
closing distal to large propodal tooth, and proximally an inconspicuous
tooth meshing between propodal teeth. Carpus irregularly spiny on
upper surface; merus-ischium subcylindrical, with numerous short and
sparser long spines tending to alternate in longitudinal rows.
488 Proceedings of the Biological Society of Washington
Fics. 2-4. Hypsophrys noar: 2-3, right chela, 2, external view, 3,
ventral view of fingers showing paired spots at base; 4, telson and part
of sixth abdominal segment with attached barnacles. Scales = 5 mm:
upper, 2, 3; lower, 4.
Second, third and fourth legs similar to each other, longer than che-
lipeds, proximal articles oval in cross section, distal ones flattened;
fourth leg having shorter coxal and basal articles than preceding legs;
each merus bearing dorsal row of well separated, strong, acute, curved
spines; propodi with pair of movable spines mesiodistally at articulation
with dactyls; each dactyl slightly curved, with row of slender corneous
spines along prehensile margin, ending in an acute corneous tip.
Fifth legs dorsal in position, much more slender and shorter than
preceding legs; carpus and propodus about equal in length and together
about as long as merus; propodus branching into shallow, asymmetrical
Y-shape terminally, one branch bearing dactylar condyle, other ter-
minating in densely spined, setose, spooned pad forming stationary mem-
ber of subchela opposing extremely short, curved, clasping dactyl; dactyl
closing at about 90° to axis of leg, length twice width of propodus (left
detached), bearing row of closely arranged, strong, curved, corneous
spines on ventral edge and tip.
New homolid crab from Florida Straits A89
Fics. 5-9. Subchela of fifth leg: 5, Hypsophrys noar, distal articles
of right leg in dorsal view; 6, same, subchela enlarged; 7, left, ventral
view; 8, H. longipes, left, ventral view, syntype 2, BMNH 1899.1.20.14;
9, H. superciliosa, same, 9, USNM 42696. Scales: 5 = 2 mm; 6, 7 =
il sanromg @, ©) == Oss saatene
Abdomen large, ovate in outline, composed of 6 free segments; first
segment dorsal and quite narrow; fifth segment broadest and longest.
Telson large, essentially triangular, but almost trilobate with slightly
mucronate naked terminal lobe and rounded proximolateral lobes having
setose margins.
First and second pairs of gonopods large and conspicuous. First gono-
pods diverging tangentially distolaterad from articulation as stout, flat-
490 Proceedings of the Biological Society of Washington
Fics. 10-12. Hypsophrys noar, male gonopods: 10, first, caudal view;
11, same, terminal aspect of tip; 12, second, cephalic view. Scale =
2 Tama,
tened plate, but becoming aligned with central axis of body and rolled
into tube progressively flattened distally and drawn into low, shoulder-
like, somewhat corneous distomesial ridge angling distolaterally into
prominent and more membranous distolateral lobe; secondary lobular
shoulder on distocaudal aspect, and (concealed from view caudally but
visible terminally ) a small central triangular lobe folded in from cephalic
surface and nestled among other terminal foliations; suture line on caudal
surface marking edge of rolled tube; short, dense setae on lip of broadly
open proximal aperture, and longer setae distally.
Second gonopods shorter and more compact than first pair, bent tan-
gentially mesad from flattened basal portion and formed into thick
nearly cylindrical plunger terminating in asymmetrical, flared, truncate
New homolid crab from Florida Straits AQ]
head tilted mesially; central concavity of head membranous and furred
with short, soft setae; longitudinal mesial tuft of longer setae on cylin-
drical distal portion.
Measurements in mm: Carapace: length from rostral notch to pos-
terior margin, 39.7; width excluding spines, 37.7; height, 20.4; maxi-
mum height of body, 24.3. Length of right hand, 42.2; left, 40.9. Aver-
age length of articles in legs 2-4: merus, 50; carpus, 19; propodus, 40;
dactyl, 25.
Type-locality: SW of Dry Tortugas, Fla., 24°11’N, 83°21.5’W, 400
fm (732 m), Silver Bay Stn. 1196.
Material studied: The holotype deposited in the U.S. National Mu-
seum of Natural History (150816) is the only known specimen of this
species. It was compared with specimens of H. superciliosa and H.
longipes.
Name: The name is a noun in apposition from the Greek “noar”
meaning phantom or specter, referring to the fancied appearance of
the crab (Liddell and Scott, 1940, p. 1177).
Remarks: Hypsophrys noar is larger than reported specimens of super-
ciliosa (ov @, cl 19 mm) but about the same size as H. longipes (ov @,
cl 38 mm) (Alcock, 1900). In both H. longipes and H. noar the rostrum
is bifid, and the carapace conspicuously spined; H. superciliosa has a
simple rostrum, and the body is somewhat less spiny. Hypsophrys
longipes has spines on the third maxillipeds but the other species lack
these. The body in H. superciliosa is smooth and nearly devoid of setae
except for a few long ones; H. longipes has a sparse covering of very
short setae in addition to a larger number of longer ones, and H. noar
has a dense coat of short setae and still more numerous longer ones. The
oval spots on the chelae are concave in H. longipes and H. superciliosa
but flat in H. noar. Superficial structure suggests that these spots may
be photophores.
The specialized subchela on each fifth leg is distinct in the 3 species
(Figs. 6-9). Unlike that in H. noar, the propodus is not Y-shaped
distally in either H. superciliosa or H. longipes, but its mesial edge is
angled so that the dactyl must swing through an arc of nearly 135° to
close. In both, the propodus has strong spines bordering the surface
against which the dactyl closes. In H. superciliosa the dactyl bears a
few curved corneous spines ventrally on the occlusal surface, as well
as some opposed fringing setae on the propodus, but in H. longipes
the dactyl has only a strong corneous tip and row of weak spines on
the edge of the occlusal surface. In specimens of H. longipes and H.
superciliosa examined, the linea homolica extends nearly to the posterior
edge of the carapace.
The species of Hypsophrys can be distinguished by the following key:
IL, ROSaMETIN Gihanyo) hy yoxommcetol ee superciliosa.
JRYORSCTeTTARY [Spc Ee NE SR TO ee ee SNe OR SS aOR, MPT Re el 2
492 Proceedings of the Biological Society of Washington
2. Propodus of last legs as long as carpus; third maxillipeds without
SITUS SSS el ee ea nd noar.
Propodus of last legs much shorter than carpus; third maxillipeds
Wlda, Gomes Cm Wsclomonn einel meng longipes.
Fastened to setae on the abdomen and left cheliped of the holotype
of H. noar are a number of small pedunculate barnacles, Poecilasma
inaequilaterale Pilsbry.
ACKNOWLEDGMENTS
I am indebted to Henry B. Roberts who first recognized H. noar as
a probably undescribed species, provided the photograph for Fig. 1,
and helped with other details of study. R. W. Ingle, British Museum
(Natural History) (BMNH), and V. K. Prem Kumar and K. K. Tiwari,
Zoological Survey of India, loaned specimens from their institutional
collections for comparative study, and William A. Newman identified
the commensal barnacle. Fenner A. Chace, Jr., and Isabel Pérez Far-
fante critically read the manuscript, and Maria M. Diéguez prepared
the drawings.
LITERATURE CITED
Aucock, A. 1900. Materials for a carcinological fauna of India. No.
5. The Brachyura primigenia or Dromiacea. Jour. Asiat. Soc.
Bengal 68, pt. 2 (3) [1899]:123-456.
———, AND A. R. S. AnprErRson. 1899. Natural history notes from
H. M. Royal Indian Marine Survey Ship ‘Investigator, Com-
mander T. H. Heming, R. N., Commanding. Series III, No. 2.
An account of the deep-sea Crustacea dredged during the
surveying-season of 1897-98. Ann. Mag. Nat. Hist. ser. 7,
Bo ILA
Inve, J. KE. W. 1913. Die Decapoda Brachyura der Siboga-Expedition.
I, Dromiacea. Siboga-Exped. Monogr. 39b, 96 pp., 4 pls.
LippELL, H. G., Anp R. Scotr. 1940. A Greek-English lexicon, a new
(ninth ) edition. Oxford Univ. Press, London. xlviii -+- 2111 pp.
SERENE, R., AND P. LOHAVANIJAYA. 1973. The Brachyura (Crustacea:
Decapoda) collected by the Naga Expedition, including a
review of the Homolidae. Naga Rep. 4(4):187 pp., 21 pls.
Woop-Mason, J., AND A. Atcockx. 1891. Natural history notes from
H. M. Indian Marine Survey Steamer “Investigator,” Comman-
der R. F. Hoskyn, R. N., commanding.—No. 21. Note on the
results of the last season’s deep-sea dredging. Ann. Mag. Nat.
Hist. ser. 6, 7( 27) :258-272.
BG. G6 72>
Vol. 87, No. 43, pp. 493-500 31 December 1974
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
HEDGPETHIUS TRIDENTATUS, A NEW GENUS AND
NEW SPECIES, AND OTHER PYCNOGONIDA FROM
KEY WEST, FLORIDA, U. S. A.
By C. ALLAN CHILD
Smithsonian Institution, Washington, D. C. 20560
The following specimens were collected during repeated
surveys for marine annelids by Dr. Meredith L. Jones, of the
Smithsonian Institution, on the islands of Key West, Florida.
They include five species, two of which are new. This is rather
surprising, because the area 60 miles west of Key West was
intensively collected for marine organisms for many years
during the time the Carnegie Institute Marine Laboratory was
located in the Dry Tortugas Islands. These collections were
reported on by Hedgpeth (1948), and hardly any new species
have come to light from southern Florida since that time.
That the present collection contains 40 percent new species
attests to the careful hand collecting techniques used by Dr.
Jones.
I wish to thank Dr. Jones for making the specimens avail-
able for study. They are deposited in the collections of the
U. S. National Museum of Natural History (USNM).
AMMOTHEIDAE Dourn, 1881
Hedgpethius, new genus
Diagnosis: Ammotheidae: Ascorhynchus-like with minutely papil-
lose body surface, without trunk or lateral process tubercles. Anterior
trunk segment longer than combined length of posterior 3 segments.
Ocular tubercle near extreme anterior of first segment; first lateral
processes at extreme posterior of first segment, imparting “long necked”
appearance. First 2 trunk segments articulated, last segment only faintly
articulated. Proboscis with large ventrodistal mouth opening and 3
43—Proc. Brot. Soc. WAsH., VoL. 87, 1974 (493 )
494 Proceedings of the Biological Society of Washington
anterior pointing tubercles arranged laterally and ventrally around its
largest circumference. Scape 1-segmented, very short; chela vestigial,
ovoid, without fingers. Palp 6-segmented. Oviger (female) rudimentary,
3-segmented. Propodus with large auxiliary claws more than twice
length of very reduced main claw.
Type-species: H. tridentatus, n. sp.
Etymology: Named for Joel W. Hedgpeth in recognition of his en-
during contributions to the phylogeny and systematics of the Pycno-
gonida. Gender, masculine.
Relationships: This genus is closely related to Ascorhynchus, but has
the following peculiarities which immediately remove it from the many
described species of that genus: It has 6 rather than 9 palp segments,
and peculiarly degenerate ovigers which are alike on both specimens
examined. The proboscis tubercles are not found on any Ascorhynchus
species and may be similar in nature to the female sexual (?) alar pro-
cesses of Anoplodactylus portus and A. digitatus. Perhaps they exist
only on females of the new genus. The presence of dominant auxiliary
claws flanking a reduced main claw is not found in any species of
Ascorhynchus, a genus without auxiliary claws. The genus Ammothella
has 1 species, A. biunguiculata, in which this claw configuration exists,
but here the main claw is reduced to a tiny tubercle which does not
appear to have articulation. There is no further resemblance between
Hedgpethius and Ammothella. The extremely long first trunk segment
is similar to some species of Ascorhynchus, although with Hedgpethius
it is longer in relation to the other trunk segments than with any As-
corhynchus species.
The genus Hedgpethius is placed tentatively in the Ammotheidae
because, although the presence or absence of complete male ovigers is
unknown, it conforms with all other definitions of this heterogeneous
group of genera.
Hedgpethius tridentatus, new species
Figure 1
Material examined: Holotype, USNM 149797, ovigerous female;
Paratype, USNM 149798, female: Key West, just west of bridge to
Fleming Key, south shore among intertidal and subtidal rocks, 24 March
1968, Sta. KW-5.
Description: With the characters of the genus. First body segment
0.3 longer than combined length of posterior 3 segments. Lateral pro-
cesses short, less than half trunk diameter, separated by their own diam-
eter, all but posterior pair armed with single dorsodistal seta. Ocular
tubercle a low truncate cone near anterior of ocular segment. Eyes
large, unpigmented in alcohol. Abdomen short, slightly bulbous distally,
not extending beyond fourth lateral processes, armed with 2-3 short
dorsodistal setae.
Pycnogonids from Key West, Florida 495
Fic. 1. Hedgpethius tridentatus, new genus, new species: a, dorsal
view of trunk; b, dorsal proboscis; c, chelifore; d, oviger; e, oblique
view of trunk; f, third leg; g, terminal segments of third leg; h, palp.
Proboscis large, ovoid, without constrictions, carried pointing pos-
teriorly as in Ascorhynchus. Two dorsolateral and 1 ventral anterior-
pointing tubercles surmount widest diameter of proboscis at %4 its length.
Anterior to dorsolateral tubercles are 2 bulges in same longitudinal axes.
Mouth at anterior tip, extending ventrally for % length of proboscis;
lips flat, distinct.
496 Proceedings of the Biological Society of Washington
Scape l-segmented, short, heavily papillose, armed with single dorso-
distal seta. Chela vestigial, carried anaxially, ovoid, without trace of
fingers.
Palp 6-segmented. First segment no longer than wide; second longest,
inflated toward proximal end, with hint of suture line towards distal
end; third twice as long as wide, armed with 2 ectal setae as long as
segment diameter. Fourth segment thin, 54 length of second, armed with
2 stout ectal setae or spines distally. Fifth as wide as long, unarmed;
sixth 6 times its diameter, armed distally and ectally with 7-8 setae.
Distal setae longer than 3 times diameter of segment.
Oviger (female) rudimentary, 3-segmented, arising on ventral surface
of ocular segment just anterior to first lateral processes. First and third
segments subequal; second 3 times as long, L-shaped. All segments
extremely small.
Third leg of moderate length, femur swollen with eggs. Femur sub-
equal in length to tibiae, tibiae equal, armed with several short dorsal
setae and 1-2 setae longer than segment diameter, several lateral and
ventral setae shorter than segment diameter. Propodus thin, without
heel or large sole spines, armed with long dorsal setae arising from
tubercles and several lateral and distal setae. Main claw tiny, curved
ventrally under propodus. Auxiliaries large, over twice main claw length,
strongly curved. Propodal sole armed with single row of 7-8 setae equal
to segment diameter.
Measurements (holotype) in mm:
Ocular segment length 0.50
Total trunk length (anterior tip to tip of 4th lateral processes ) 84
Trunk width (across 2nd lateral processes ) Bl
Proboscis length 40
Abdomen length .10
Third leg: Tibia 1 A2
Coxa 1 0.07 Tibia 2 42,
Coxa 2 19 Tarsus 05
Coxa 3 13 Propodus 19
Femur 44 Auxiliary claw .08
Distribution: Known only from the type-locality, Key West, Florida.
Etymology: The specific name refers to the three tooth-like tubercles
on the proboscis.
Remarks: This species first appeared to be an aberrant Ascorhynchus
undergoing regeneration, but the second specimen with the same set
of characters did not support this view. The peculiar tuberculate pro-
boscis is unlike that of any other pycnogonid, to my knowledge, and
may or may not be confined to the female. The ovigers appear as they
would on most juvenile pycnogonids, but present an enigma on a mature
female with eggs. Perhaps both specimens have damaged ovigers that
Pycnogonids from Key West, Florida 497
are undergoing regeneration. Only further collecting of male and female
adults can solve these perplexing problems.
Achelia sawayai Marcus
Achelia (Pigrolavatus) sawayai.—Fry and Hedgpeth, 1969: 104 [litera-
ture].
Material examined: One female, just west of bridge to Fleming Key,
south shore among intertidal and subtidal rocks, 24 March 1968, Sta.
KW-3. Eight males, 6 females, 2 juveniles, scrapings of culvert wall
carrying water from Cow Key Channel to ponds west of Airport, 0.3-
0.6 meter, 14 May 1967, Sta. KW-5. Two males, 2 females, on rocks
in above culvert, Sta. KW-6. One male, 1 female, 1 juvenile, Aero
Palms Club at base of Trumbo Point, subtidal under rocks, 30 March
1968, Sta. KW-6. Two females, east side of Fleming Key near boat
dock, with Thalassia and Halimeda, subtidal, 20 May 1967, Sta. KW-9.
Remarks: These specimens are typical and inseparable from their
neighbors in the Dry Tortugas.
Ammothella appendiculata (Dohm)
Ammothella appendiculata.—Stock, 1955: 250 (literature).
Material examined: One subadult, just west of bridge to Fleming
Key, south shore among intertidal and subtidal rocks, 24 March 1968,
Sta. KW-5. One male, 2 females, 3 juveniles, scrapings of culvert wall
carrying water from Cow Key Channel to ponds west of Airport, 0.3-
0.6 meter, 14 May 1967, Sta. KW-5.
Remarks: These specimens appear to be the more slender stage II
described by Stock (1955: 251-2).
Ammothella sp.
Material examined: One juvenile, near bridge to U. S. Naval Hos-
pital, associated with black ascidians on subtidal rock, 3 May 1964.
Remarks: This is unidentifiable, but probably is A. appendiculata.
PHOXICHILIDIIDAE Sars, 1891
Anoplodactylus jonesi, new species
Figure 2
Material examined: Holotype, USNM 150619, ovigerous hermaphro-
dite (?): Key West, southwest tip, off Fort Taylor, intertidal and sub-
tidal, 24°32’55”N, 81°48’37”W, 21 June 1963, Sta. 2.
Description: Body unsegmented, but with marked indentations sug-
gesting segmentation. Lateral processes glabrous, separated by their
own diameter. Ocular tubercle low truncated cone pointing anteriorly.
498 Proceedings of the Biological Society of Washington
Fic. 2. Anoplodactylus jonesi, new species: a, dorsal view of trunk;
b, terminal segments of third leg; c, third leg; d, chela; e, lateral view
of trunk, with enlargement of dorsal pore; f, oviger with enlargement
of terminal segment.
Pycnogonids from Key West, Florida 499
Eyes large, unpigmented. Two (sensory?) papillae on dorsolateral
crown of ocular tubercle. Abdomen slightly longer than fourth lateral
processes, erect, without setae.
Proboscis cylindrical, slightly constricted anteriorly.
Chelifore as long as proboscis, thin. Scape armed with 2 lateral setae
and 1 dorsodistal seta. Chela small, fingers curved, tips overlap when
closed, armed with 2-3 setae on hand and movable finger and 4—5 mi-
nute teeth on each finger.
Oviger segments 1 and 2 subequal; segment 3 longest, with proximal
constriction and 2 distal setae. Terminal segments increasingly shorter,
armed with 2-4 setae. Ultimate segment slightly shorter than segment
2, armed with 2 recurved setae.
Legs robust, tibiae subequal, femur slightly longer, each armed with
few dorsal and ventral setae and single long dorsodistal feathered seta
on tubercle. Propodus straight with small heel bearing 3 spines. Sole
with lamina over almost entire length, armed with few setae. Claw
slightly curved. Auxiliary claws robust, short.
Hermaphrodite characters: Ovigers of male, but lacking femoral
cement glands. Female genital pores on coxa 2 of all four legs and
mass of eggs in swollen femora. Two swollen pores immediately be-
hind ocular tubercle may or may not have some sexual function. No
evidence of glands seen beneath cuticle under pores.
Measurements in mm:
Trunk length (chelifore insertion to tip 4th lateral processes ) 0.90
Trunk width (across Ist lateral processes ) 52
Proboscis length (ventral ) 00
Third leg: Tibia 1 A8
Coxa 1 0.13 Tibia 2 AQ
Coxa 2 22 Tarsus O07
Coxa 3 .20 Propodus oi}
Femur 53 Claw 24
Distribution: Known only from its type-locality, Key West, Florida.
Etymology: Named for its collector, Meredith L. Jones.
Remarks: There is nothing adherent on the ovigers which could prove
them functional, and without the usual femoral cement glands of males
in this genus I hesitate to call this specimen a true hermaphrodite. This
single pycnogonid has all of the normal female characters, including
eggs, and it is possible the ovigers are just the result of early embryonic
misdetermination. There is nothing I could find in the leg that takes
the form of a testis, such as that figured by Marcus (1952, p. 30, fig. 9)
for Ascorhynchus corderoi. If subsequent collecting proves this to be
a functional hermaphrodite, then Anoplodactylus jonesi will only mark
the second record of this phenomenon in an otherwise dioecious group.
Besides the evident bisexual characters of this specimen, there is no
other outstanding character that separates it from many others of this
500 Proceedings of the Biological Society of Washington
large, cumbersome genus. Its two closest geographical associates on
the American coast are Anoplodactylus parvus and A. petiolatus, and
although it compares favorably in size to these two, chelifore and oviger
length ratio differences separate it from A. parvus.
Propodus shape, ocular tubercle size, segment length ratios, and the
presence of lateral process tubercles on A. petiolatus serve to separate
A. jonesi from that species.
Anoplodactylus pectinus Hedgpeth
Anoplodactylus pectinus Hedgpeth, 1948; 234-236, fig. 34.—Stock, 1955:
935, fig. 11.—Arnaud, 1973: 955-957.
Material examined: Two females, Fleming Key, on east side near boat
dock, with Thalassia and Halimeda, subtidal, 20 May 1967, Sta. KW-9.
Remarks: This is an easily recognized species and these specimens
are typical of Hedgpeth’s Tortugas males and Stock’s Virgin Island
females. Why these should turn up in the Indian Ocean at Madagascar
(Amaud, 1973) only serves to emphasize our extreme lack of knowledge
concerning transport and zoogeography of the majority of pycnogonid
species. So many species have appeared to be endemic to some areas,
but later turn up in widely disparate places, sometimes as if to con-
found our various efforts to compartmentalize, that the value of dis-
tribution maps becomes questionable.
LITERATURE CITED
ArNAvuD, F. 1973. Pycnogonides des Récifs Coralliens de Madagascar.
4.—Colossendeidae, Phoxichilidiidae et Endeidae. Tethys 4(4):
953-960, figs. 1-8.
Fry, W. G., anp J. W. HeEpcPETH. 1969. The fauna of the Ross Sea,
7. Pycnogonida, 1. Colossendeidae, Pycnogonidae, Endeidae,
Ammotheidae. New Zealand Oceanogr. Inst. Mem. 49:1-139,
figs. 1-209, tables 1-16.
HeEpcretTH, J. W. 1948. The Pycnogonida of the western North At-
lantic and the Caribbean. Proc. U. S. Nat. Mus. 97:157-342,
figs. 4-342, charts 1-3.
Marcus, E. pu Bois-REyMonp. 1952. A hermaphrodite Pantopod.
Anais Acad. Bras. Ciéne. 24(1):23-30, figs. 1-9.
Stock, J. H. 1955. Pycnogonida from the West Indies, Central Amer-
ica, and the Pacific Coast of North America. Vidensk. Medd.
Dansk naturh. Foren. 117:209-266, figs. 1-26, tables 1-2.
OTF 0673
Ashmunella animasensis
“INDEX TO NEW TAXA
Youn 86 PeUe oe
(New taxa indicated in boldface; n. c. = new combination )
PHANEROGAMAE
LEGUMINOSAE
BOCOAvAl Lena ti Curie ad MeReuieonaideeeh. wh MN wae eae AD TT ER
CLC HPT TDS ee eae sl rca ete i eA ee
limae
SOUYOUU FIGs caso ee nue LN A en eee eT RU See Neal bana
ROW ACEMSIS Tlie Cid an kia e nc) SOE a SS Sh Saree
AC CIAAUI OSTA sr acti ee eet She A RAE AR
NUTACLUEL OHA Tair Reatalelawmeds as ed My SN tral nn lal vhs itis 9 le oe
MOLLUSCA
GASTROPODA
BRULEE CULL Cape erst Whee hme ction ce eed bien ate i hee oo a AR
kochit-caballoensts tec ro ici cy kn an pect
kochi cornudasensis
kochi sanandresensis
ATS CEO 111 Jo Lied cas ee wa ace eo my PN ee ee
pasonisepolygyroideay ssn vate. 2 we eee
SALIMASEISIS re eset Aleines ia 4 wl a IAN nena roan ROMER,
ANNELIDA
POLYCHAETA
[ Breer I Peer idea ete Utena ER ke id NS a eee See ON (eer re eRe Te meee
POST IES AA Gi Tel dit eset toes AMG cz oe VON EAE ot Re oo Tse UR etn ey SRT
BScUGOpOly Gora @ibWest cme a sie eon ce ee cad ae ia
BRANCHIOBDELLIDA
(Chavanll of weriaveco) ka), TaaVOvaNeNaNUIS Gay, Chee eee
ARTHROPODA
PYCNOGONIDA
Anioplodacty lis si @mesi i 5.50 ie oe
EW eh ope ER UN S ype es 9 ae ow Ld tN
LAR TH Cay ch et iT hohe sDeaiae emia Dk Ane RECUR Gle Meany MR Mee Oy Br Win, sks Ieee
67
CRUSTACEA
Acanthomolgus aretatipes 2 20 ns e loh ee ae 424
CO TTR TD an RU UNS ta PRN lata LS Ue Al7
mrrn en pose ca ee RE ii 412
CALL E:T 6:1 ct Tei nme ee JON ed es one ULM SM MRCS ee Net WN 19
Ta R ros Gar 2a ee Tce Nr a a OR 19
Calocaris!((‘Calastacus))) yemme ra) ieee ere rs ae arene Se 451
CORY po ua an 25 a a La Rte A457
Dodo Dy re tus TT RS 130
TU OEE) 0 U2) 1) Meee aN Mae NUN ADDN OM a bras: CU ed 130
Hey pop lirrys) rveo gare p00 8 dS Sn ee Se 485
Munnogoniumsgloloitr ors rat (cp aea 266
OTATIGE Ms Ce dat cr ee ISL At a 266
THVAQVEARTUEL The Crea nse Oe 266
cll evgcVou cet Cum eeneUD a MCU Aas OMNI CR ee R ANCL eh i 266
Nematopagurus spinulosensoris _____________-__--_________-_-___------------------- 246
Paramol gus’ ellise lee NT Ce IM at el oncom 432,
Uca pamace a 8s eM 2 I SEES A io EW 316
DIPLOPODA
Pseudopolydesmus collinus) 2.200 iC ee en ee 346
INSECTA
Alloperla atl emitieee 000) i ee 260
Gtemiccobunrm 2522 NTT 284
aAcapwlCensis | 2.07 A ae 288
Voto yi UN RS he A 2 ck 303
eo Ry cure wk ig DY 292
Crotonae MNCs .2 cose ae a le 307
janizend 2)oe eu) ee aa 289
wmar nal e020 a 0 aE 305
Sall vray or CA, ila ALE NN Se 290
tuberculatum ny. Cyst ee eh ae ee 290
Buxesta: riazorca hie sich 2) IR Uo) RO a ere 73
Glaresis’ arena satan) jist 2 0 20k ame UN URE a ec 93
Trichodeéctes.emiery ix ..0.2 aie al Ne de ae ee ii
ECHINODERMATA
ECHINOIDEA
AsvecoTaarnn pass i226 0 hh tI a 43
euGDca rat eee 02 Rg aca 2 ga 44
Pislva Hes io) RI NR a ho ere cea 43
real hi fer: Wi a0 Reale auDMO AMMO UUNLEE A Cieire taninMmontc ny TOE Es 43
Wnastata sn) oe oN eae DR a A ee cee 43
OSU Es ie Ih kA 42,
CHORDATA
PISCES
CGrenlasomacre sama!) Awe he ea ch US EE, Ca ed eee ae 466
Wecoc omeprnne le sunny oe cee eee ee ee ee ee 207
Microvnathus \erugabus 200 J oT
ROtAMOEDMAp MIS Pelers! 2.62.28. Ns a 34
Bseudophiallus: brasiliensis, 2200.02 406
SSAA OPTS 4) CORNET S CORR pes re see a oe ag HI Te oe ee 194
SLE KA CUCU S essen ee a ial ree a ee 200
AMPHIBIA
Mleutherodactylus. walkeri: 2.0500 eee 381
Sphiacrodackylustarmast oes ei ea ie he wee a ee 339
NEW AT) DAI TAS) SE SOS TDs fe Di Pe wae ah 2, AUP Set Fer 88
Giseodacty lus ei nape eee Neate ee sede ee ne 88
REPTILIA
ieptotyphlops: tricolor, 2.2. re 167
“WSs ky NS SCAT eae ES Ss ion, ese SP AL Mie Get ake 12
MAMMALIA
atecrilltiswarenarstes eles ogee ee a ee 397
INEsotrocnisistemaninos) 22 0 2 ee 442
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