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PROCEEDINGS
of the
Biological Society of
W ashington
VOLUME 8l
1968
0C1 20 1969
LIBRARIES
WASHINGTON
PRINTED FOR THE SOCIETY
EDITOR
Rocer F. CRESSEY
All correspciidence should be addressed to
the Biological Society of Washington, Smith-
sonian Institution, Washington, D.C. 20560
ALLEN Press INc.
LAWRENCE, Kansas 66044
rc rT
OFFICERS AND COUNCIL
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
FOR 1968-1969
OFFICERS
President
JOSEPH ROSEWATER
Ist Vice President
MEREDITH L. JONES
2nd Vice President
DANIEL M. COHEN
Recording Secretary
STANWYN G. SHETLER
Corresponding Secretary
RICHARD C. BANKS
Treasurer
HARALD A. REHDER
Council
Elected Members
THOMAS E. BOWMAN JAMES A. PETERS
K. C. EMERSON GEORGE A. LLANO
CLAYTON E. RAY
TABLE OF CONTENTS
Volume 81
Banarescu, Petru M., and Teodor T. Nalbant. Some new Chinese
Minnowsy.@kiscess) Gyprinitormes))) —/-ueslasle nee eeneeessmeuenie
Banks, Richard C., and Wayne H. Bohl. Pentland’s Tinamou in
ANgusraioe) « (Waniese Abner ys) Lia
Banse, Karl. Streptosyllis latipalpa, new species (Polychaeta,
Syllidae) from Puget Sound (Washington) —___________-__.
Banse, Karl, and Frederic H. Nichols. Two new species and
three new records of benthic polychaetes from Puget Sound
(Washimgtom)\ Sci) Ea gn ee
Barbour, Clyde D., and Salvador Contreras. Algansea monticola,
a new cyprinid fish from the Pacific Slope of Central Mexico
Berry, Frederick H., and Peter J. P. Whitehead. A new species
of sardine (Sardinella, Clupeidae) from the Marquesas
MST earn ES es aN EN Ns LE ae
Birdsong, Ray S. Microgobius crocatus, a new gobiid fish from
Pacific: Paniamia® 20.2 2:2 We ay: Se
Bohart, R. M. A synopsis of the American species of the genus
Oryitus; (Hymenoptera, Sphecidae)) jena
Bowman, Thomas E., Rudolph Prins, and Byron F. Morris. Notes
on the harpacticoid copepods Attheyella pilosa and A. carol-
inensis, associates of crayfishes in the Eastern United States
Branson, Branley A., and Donald L. Batch. An ecological study
on Valley-Forest spiders from Northern Kentucky —..__.
Brown, Walter C., and Michael J. Tyler. Frogs of the genus
Platymantis (Ranidae) from New Britain with descriptions of
TRE WW: STOC CT CS! 5 a cr NL SSRI ee
Buckett, John S., and Michael R. Gardner. Revision of the mil-
liped genus Wamokia Chamberlin from the Sierra Nevada of
Central California (Diplopoda: Polydesmida: Xystodesmidae )
Carter, Dilford C. A new species of Anoura (Mammalia: Chirop-
tera: Phyllostomidae) from South America
Chace, Fenner A. Jr. A new crab of the genus Cycloes
(Crustacea; Brachyura; Calappidae) from Saint Helena,
SoutiawA tla tice @ ce arn ye tee lita ae Come ne
Cole, Gerald A., and W. L. Minckley. A new species of ‘aquatic
isopod crustacean (genus Asellus) from the Puebla Plateau,
Centrale Miexice Vereen fe ete ar eink 10 eee in eee ee neater et
Collette, Bruce B. Daector schmitti, a new species of venomous
toadfish from the Pacific coast of Central America
335-346
485-490
151-154
223-230
101-108
209-222
191-196
431-438
197-208
511-538
427-430
605-612
755-760
155-160
Cressey, Roger F. A redescription of Hatschekia conifera,
Yamaguti 1939 (Copepoda, Caligoida), including the first
descriptioniofetne male. =teniawies oie | ee Rae Ee
Dawson, C. E. Two new wormfishes (Gobioidea: Microdes-
midae))) from’ the! Indian: Ocean) =
Deevey, Georgiana B. Bathyconchoecia, a new genus of pelagic
ostracods (Myodocopa Halocyprididae ) with six new species
from the deeper waters of the Gulf of Mexico
Dillon, William Arthur, and William J.- Hargis, Jr. Mono-
genetic trematodes from the Southern Pacific Ocean. Part IV.
Polyopisthocotyleids from New Zealand fishes: The families
Mazocraeidae, Diclidophoridae and Hexabothriidae —
Dixon, James R. A new species of Gecko (Sauria: Gekkonidae )
varoveoW idaves Ben? isin lence 2
Downey, Maureen E. A note on the Atlantic species of the
StarhishhyrO emus telat C ic ys seme teuesen! NLA NY nas 0 UUe NEN DE ems cs
Emerson, K. C., and Roger D. Price. A new species of Den-
nyus (Mallophaga: Menoponidae) from the Malaysian Spine-
(eT Sra URSA Gs dd ae Sept Uae ls NNN WSN SU ELI Ree ie
Emerson, K. C., and Roger D. Price. A new species of Parafeli-
cola (Mallophaga: Trichodectidae) from Mozambique —.
Fitzpatrick, J. F. Jr., and James F. Payne. A new genus and
species of crawfish from the Southeastem United States
@DecapodaywAstacicla ea Meiwalan etme ole luaa steed ents Me lpanetnia os
Gazin, C. Lewis. A new primate from the Torrejon middle
paleocene of the San Juan Basin, New Mexico
Genoways, Hugh H., and J. Knox Jones, Jr. A new mouse of
the genus Nelsonia from Southern Jalisco, Mexico...
Glass, Bryan P., and Robert J. Baker. The status of the name
WOCKS SuloMiaiiS Sexy) 2 ee
Glynn, Peter W. A new genus and two new species of sphaero-
matid isopods from the high intertidal zone at Naos Island,
IRAN ea VN aa en AS ace IN aN Wace I IR Ee LOM meth tee A A
Groves, Colin P. A new subspecies of white-handed gibbon
from Northern Thailand, Hylobates lar carpenteri new sub-
STDC CLES i meeemeiie ta lee ote are noe ities Se Ui aR ae ar oe Ati eee esta ier
Halpern, Jerald A. Biological investigations of the deep sea.
38. A new westem Atlantic Dipsacaster (Echinodermata
Asteroidea) with the distribution of known species
Hargis, William J. Jr., and William Arthur Dillon. Helminth
parasites of Antarctic vertebrates. Part IV. Monogenetic
trematodes from Antarctic fishes: The superfamily Capsaloidea
Veale cio] GBT GIN ie IN BNE oleae en, MIE ne See oe es Oe) eee ae
Hobbs, Horton H. Jr. Two new crayfishes of the genus Cam-
barus from Georgia, Kentucky, and Tennessee (Decapoda,
PS Gal CLG eC) a eee wee eA em RIE a LE a ee al delle MSO
173-178
53- 68
539-570
351-366
419-426
4\-— 44
87— 90
109-110
1l- 22
629-634
97-100
257-260
587-604
625-628
231-240
403-412
261-274
Holt, Perry C. New genera and species of branchiobdellid worms
(GNeoavelbyebays Olea ey cey))i
Humes, Arthur G. Lecanurius kossmannianus, a new cyclopoid
copepod parasitic in holothurians in Madagascar _______-
Humes, Arthur G., and Ju-Shey Ho. Cyclopoid copepods of the
genus Lichomolgus associated with octocorals of the family
Nieves tm IWehyenenir Loe
Humes, Arthur G., and Ju-Shey Ho. Cyclopoid copepods Es the
genus Lichomolgus associated with octocorals of the families
Xeniidae, Nidaliidae, and Telestidae in Madagascar _____-- =
Jones, Meredith L. Paraonis pygoenigmatica new species, a
new annelid from Massachusetts (Polychaeta: Paraonidae )
Kornicker, Louis S. Bathyal myodocopid Ostracoda from the
Northeastern Gulf of Mexico eae eae eee
Lawler, Adrian Russell, and William J. Hargis, Jr. Monogenetic
trematodes from the Southern Pacific Ocean. Part V. Mono-
pisthocotyleids from Australian fishes, the subfamily Tro-
Chopodinae: esas ee UE 2 a a sel
Lidicker, William Z. Jr., and W. I. Follett. Isoodon Desmarest,
291-318
179-190
635-692
693-750
323-334
439-472
367-402
1817, rather than Thylacis Mliger, 1811, as the valid generic ©
name of the short-nosed bandicoots (Marsupialia: Pera-
MVR L TAG!) ye soe ater ea SS
Loeblich, Alfred R. III. A new marine dinoflagellate genus
Cachonina, in axenic culture from the Salton Sea, California
with remarks on the genus Peridinium ____..--_-____»__
Loomis, H. F. New North American spiroboloid millipeds of the
families Atopetholidae and Messicobolidae —
Manning, Raymond B. Three new stomatopod crustaceans
lagoyoo} (dove) JbayeloonWielenversy gigsry Ne
Orr, Robert, T., and J. Dan Webster. New subspecies of birds
from Oaxaca (Aves: Phasianidae, Turdidae, Parulidae)
Pawson, David L. A new psolid sea cucumber from the Virgin
MST Ara CS see, LR Rs A ON AN
Peters, James A. A replacement name for Bothrops lansbergii
venezuelensis Roze, 1959 ( Viperidae, Serpentes) _.-_--___-_
Phillips, Carleton J., and Elmer C. Birmey. Taxonomic status
of the Vespertilionid genus Anamygdon (Mammalia; Chirop-
ETERS hai ea ei etek GL A, POM
Radwin, George E. New taxa of Western Atlantic Columbellidae
(GastropodaleProsobranchiay) pees eee ene Es
Ripley, S. Dillon, and D. S. Rabor. Two new subspecies of birds
from the Philippines and comments on the validity of two
ELE TS se EEA Oe SD SA PN IU Ae OP
251-256
91- 96
499-510
241-250
37- 40
347-350
319-322
491-498
143-150
Roper, Clyde F. E. Preliminary descriptions of two new species
of the bathypelagic squid Bathyteuthis (Cephalopoda:
(OLR 05 SSG EY) )) SANS ese a Sd ARETE SS ty IR Age AC CSP
Salmon, Michael, and Samuel P. Atsaides. Behavioral, morpholog-
ical and ecological evidence for two new species of fiddler
crabs (genus Uca) from the Gulf Coast of the United States
Schwartz, Albert. Geographic variation in the new world
geckkonid lizard Tarentola americana Gray
Schwartz, Albert, and Orlando H. Garrido. An undescribed sub-
species of Leiocephalus raviceps Cope (Sauria: Igaunidae)
PLOTINEWE Ste Tern Cul bye) ele tae AO DSR uN Ms Gola a al
Shear, William A. A new genus and species of milliped from
Illinois (Polydesmida, Euryuridae)
Short, Lester L., Jr. Variation of Ladder-backed Woodpeckers in
Southwestern) North s American se sste anne g cate wee inna,
Smith-Vaniz, William F. A new clingfish, Tomicoden rhabodotus
family Gobiesocidae, from the Lesser Antilles
Spangler, Paul J. A new species of Laccobius from the Greater
Antilles (Coleoptera: Hydrophilidae)
Springer, Victor G. The Indo-Pacific blenniid fish genus
Stanulus, with description of a new species from the Great
Barrier Reef (Blenniidae; Blenniinae; Salariini) __._...______
Springer, Stewart. Triakis fehlmanni, a new shark from the coast
oS coral aye RnR Ne IRAN So SLI
Thompson, Fred G. Ceochasma, a remarkable new land snail
from Colima, Mexico (Gastropoda, Prosobranchia, Helicini-
Cleves) pie dere a UREN sD RRR SNe ae Ove ei Octo aeRO BN ea
Walls, Jerry G. A new Faxonella from northeast Louisiana
Decapoda Astaciclae:)) 2a sem sunt tail 0 A a ds Ne
161-172
275-290
123-142
23- 30
479-484
1- 10
473-478
751-754
111-122
613-624
45— 52
413-418
BIOLOGICAL SOCIETY OF WASHINGTON
PROCEEDINGS
1040th Meeting—27 June 19638
EIGHTY-NINTH ANNUAL MEETING
President Ernest Lachner called the 1040th meeting and the
89th Annual Meeting of the Society to order at 3:08 p.m. in
room 43 MNH. Thirteen members were present.
The minutes of the previous annual meeting were approved
as published in the Proceedings, with correction of the date.
Treasurer Harald A. Rehder gave his report, which was ap-
proved.
The report of the Auditing Committee was read and accepted.
The Corresponding Secretary reported that 33 new members
had been elected during the year since the last annual meeting,
and read the names of three deceased members. The Cor-
responding Secretary reported the results of the election of of-
ficers.
A summary of the meeting scheduled for 11 October 1968
was presented by President Lachner and Daniel Cohen. A
motion that speakers at the meeting be provided lunch was
passed.
The meeting was adjourned at 3:36 p.m.
Richard C. Banks
Corresponding Secretary
Acting Recording Secretary
77. OF 77 | eS: ee eat
Vol. 81, pp. 1-10 SEC el April 30, 1968
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
VARIATION OF LADDER-BACKED WOODPECKERS
IN SOUTHWESTERN NORTH AMERICA
By Lester L. SHorr, Jr.
The American Museum of Natural History
New York, New York
As part of an investigation of interactions between the
Ladder-backed Woodpecker (Dendrocopos scalaris (Wagler ) )
and the Nuttall Woodpecker (D. nuttallii (Gambel)) it was
necessary to appraise the several races of the former species
occurring in the Southwest. Results of this analysis are re-
ported herein.
The races of the Ladder-backed Woodpecker were reviewed
by Oberholser (1911). Van Rossem (1942) described two
new subspecies, mojavensis and yumanensis, from the South-
west. These two forms have not been generally recognized,
although no one has criticized van Rossem’s data or provided
evidence against his tenets. I find that mojavensis and yuman-
ensis do in fact have some basis, although they do not merit
formal taxonomic recognition.
I measured and studied 550 specimens of D. scalaris from
the Southwest including the following: D. s. lucasanus (south-
ern Baja California )—60; D. s. eremicus (northern Baja Cali-
fornia )}—114; northern Baja California contact between D. s.
eremicus and D. s. cactophilus—11; and D. s. cactophilus—
365. Of the latter, 21 came from southwestern Utah and
northwestern Arizona, 107 from central and southeastern Ari-
zona, 25 from the upper Colorado River Valley (van Rossem’s
“yumanensis,’ in part), 84 from the lower Colorado River
Valley (“yumanensis”), 26 from the Providence Mountains
and vicinity, 26 from central southern California, and 76 from
central and from southwestern California (van Rossem’s “mo-
javensis’). Standard measurements were taken, including the
1—Proc. Bion. Soc. Wasu., Vou. 81, 1968 (1)
2 Proceedings of the Biological Society of Washington
TaBLE 1. A mensural comparison of southwestern male Ladder-backed
Woodpeckers taken from December through March.*
Form XK +2SE Range SD CV N
Wing length
eremicus (Southern ) 105.24+1.04 101.0—-108.0 1.96 1.86 14
“mojavensis”’ (CS California) 105.57+1.24 101.5-110.1 2.64 2.50 18
“yumanensis”’ (Lower Col. R.) 102.91+0.96 98.5-1071 2.38 2.31 24
cactophilus (C, SE Ariz. ) 103.14+0.68 97.8—-104.6 1.65 1.60 24
Tail length
eremicus (Southern ) 69.69=+0.88 67.5— 73.5 1.63 2.34 14
“mojavensis”’ (CS California ) 64.7441.16 62.0— 70.3 2.38 3.68 17
“yumanensis” (Lower Col. R.) 62.23+0.86 58.7— 67.8 2.09 3.36 23
cactophilus (C, SE Ariz. ) 61.77+1.02 55.8— 65.8 2.47 4.00 24
Bill length ( gonys)
lucasanus 16.31+0.32 15.0— 17.4 0.73 3.24 15
eremicus (Southern ) 17.73+0.50 16.1— 19.9 1.63 5.53 15
“mojavensis’”’ (CS California ) 16.68+0.36 15.6— 18.5 0.75 4.50 17
“yumanensis’”’ (Lower Col. R.) 15.75+0.36 14.4— 18.2 0.89 5.65 24
cactophilus (C, SE Ariz. ) 14.91+0.34 13.3- 16.6 0.87 5.84 25
Depth of back bar*
eremicus (Southern) 4.51+0.20 3.5- 6.0 0.59 13.08 38°
““mojavensis” (CS California ) 3.64+0.12 2.6- 4.3 0.36 9.89 42,
“‘yumanensis” (Lower Col. R.) 3.130.12 2.4- 4,1 0.42 13.42 AT
cactophilus (C, SE Ariz.) 3.6140.12 2.8- 4.7 0.44 12.19 56
* Measurements are in millimeters, except for CV. Figures are for specimens
collected from December through March, except for depth of back bar, where males
from all months were utilized. Symbols and abbreviations; K = Mean, SE =
Standard Error of the Mean, SD = Standard Deviation, CV = Coefficient of Vari-
ability, and N = number of specimens in the sample.
chord of the wing and the gonys of the bill. It is my expe-
rience that measurement of the gonys is more easily accom-
plished and involves less error in measuring than any other
bill length measurement in these woodpeckers; also, in mu-
seum specimens, the lower bill (which is involved in the
gonys measurement ) is less often broken than the upper bill.
A comparison of mensural characters of certain samples is
presented in Tables 1 and 2. The “mojavensis” and “yuman-
ensis’ samples are taken from the areas designated for these
races by van Rossem (op. cit., p. 24). Only specimens taken
from December through March were used in the tables
because these were most numerous in the samples secured.
Also, this period is represented by specimens in a state of
plumage wear intermediate between fresh-plumaged fall birds
(late August-November) and worn birds taken from April
Variation of Ladder-backed Woodpeckers 3
TABLE 2. A mensural comparison of southwestern female Ladder-backed
Woodpeckers taken from December through March.*
Form X + 2SE Range sD CV N
Wing length
eremicus (Southern) 102.43+0.86 100.0—106.9 1.42 1.39 11
“mojavensis”’ (CS California) 103.310.90 101.1—104.7 1.28 1.24 8
“yumanensis”’ (Lower Col. R.) 98.98+1.06 95.5-104.1 2.27 2.29 18
cactophilus (C, SE Ariz.) 100.59+0.92 96.2-106.2 2.25 2.24 24
Tail length
eremicus (Southern ) 68.5641.58 64.1-— 73.1 2.63 3.84 11
“mojavensis’” (CS California ) 64.21+1.54 59.7-— 66.1 AON | 3.44 8
“yumanensis’” (Lower Col. R.) 61.51+1.00 58.5— 64.7 2.12 3.45 18
cactophilus (C, SE Ariz. ) 61.62+0.86 58.0— 66.8 2.10 3.41 24
Bill length (gonys)
lucasanus 13.59+0.42 12.9— 16.0 0.77 5.67 14
eremicus (Southern) 15.82+0.60 14.7— 17.8 1.00 6.32 11
“mojavensis’” (CS California ) 14.44+0.50 13.38-— 15.3 0.71 4.92 8
“yumanensis” (Lower Col. R.) 13.34+0.36 12.7-— 14.5 0.72 5.40 18
cactophilus (C, SE Ariz. ) 12.92+0.26 11.2— 14.5 0.67 5.19 25
Depth of back bar*
eremicus (Southern) 4.50+0.18 3.4-— 6.0 0.54 12.00: 33
“mojavensis”’ (CS California ) 3.42+0.20 2.3- 4.7 0.55 16.08 31
“yumanensis”’ (Lower Col. R.) 2.99+0.08 1.9- 3.8 0.39 13.04 37
cactophilus (C, SE Ariz. ) 3.48+0.14 2.6— 4.8 0.49 14.08 51
* Measurements are in millimeters, except for CV. Figures are for specimens
collected from December through March, except for depth of back bar, where
females from all months were utilized. Symbols and abbreviations as in Table 1.
through July. Samples representing the latter periods were
also utilized, and data gathered from them. Such data yielded
results similar to those in Tables 1 and 2. Some of these data
(from birds taken April-July) were presented in Table 3.
D. s. eremicus (Oberholser). This northern Baja California
race exhibits longer wings and a longer tail than the southern
peninsular Baja California D. s. lucasanus (Xantus), and it
agrees with lucasanus in having a relatively longer tail than
that of other southwestern scalaris. The male tail: wing ratio
averages 0.65 to 0.67 in eremicus, as in lucasanus, versus 0.59
to 0.61 in all samples of cactophilus, “mojavensis” and “yuman-
ensis.” Compared with “mojavensis,” which occupies a range
contiguous with that of eremicus in the foothills of southern
California, eremicus has wings of about the same length, a
7 percent longer tail, a longer bill (not significantly longer in
4 Proceedings of the Biological Society of Washington
TABLE 3. A mensural comparison of certain samples of southwestern male
Ladder-backed Woodpeckers taken from April through early July.*
Form X + 2SE Range sD CV N
Wing length
eremicus (Southern ) 104.08+0.84 101.8—-106.3 1.32 1.27 10
eremicus (Northern) 103.55+——___ 100.0-107.0 — — 5
“mojavensis” 104.89+1.06 101.7—-110.1 2.14 2.04 16
Providence Mt. area 101.53+1.40 98.1-103.7 1.98 1.95 8
Mecca area 104.2641.78 101.0—108.4 2.67 2.56 9
“yumanensis’” (Lower Col. R.) 102.00+1.08 98.2-106.5 2.22 2.18 17
“yumanensis” (Upper Col. R.) 102.12+1.14 99.9-104.2 1.72 1.68 9
SW Utah area 104.18+——__ 102..8-106.4 — — 6
cactophilus (C, SE Ariz.) 101.74+0.88 98.4-105.3 1.88 1.85 18
Tail length
eremicus (Southern ) 68.138+1.58 64.8— 70.8 2.23 3.27 8
eremicus (Northern) 68.36+— 65.1— 71.3 — — 6
“mojavensis”’ 63.8641.12 59.7— 67.1 2.26 3.24 16
Providence Mt. area 60.50+1.68 58.0— 63.5 2.23 3.69 i
Mecca area 63.38+1.72 59.5— 66.0 2.43 3.83 8
“ygumanensis”’ (Lower Col. R.) 61.54+1.06 57.5— 65.1 2.24 3.64 18
“yumanensis” (Upper Col. R.) 62.09+1.00 60.4— 65.2 1.49 2.40 9
SW Utah area 63.97+—_ 60.6— 67.8 — — 6
cactophilus (C, SE Ariz. ) 60.12+1.10 56.3— 63.5 2.20 3.66 16
Bill length (gonys)
lucasanus 15.83+0.44 14.8— 17.0 0.73 4.61 11
eremicus (Souther ) 17.21+0.50 15.6— 18.7 0.84 4.88 11
eremicus (Northern ) 17.34+0.78 16.2— 19.0 1.09 6.23 i
“mojavensis”’ 16.81+0.44 15.9— 19.2 0.87 4.09 16
Providence Mt. area 16.50+0.60 15.0— 17.3 0.86 5.21 8
Mecca area 16.26+0.80 15.1— 19.1 1.19 7.32 9
“yumanensis’ (Lower Col. R.) 15.660.36 14.2-— 17.0 0.70 4.47 16
“yumanensis” (Upper Col. R.) 16.69+0.70 15.0— 18.3 1.06 6.35 9
SW Utah area 17.02+——_ 14.8— 18.2 — — 6
cactophilis (C, SE Ariz. ) 15.41+0.49 14.2— 17.2 0.86 5.58 18
Length of 4th rectrix
eremicus (Southern ) 58.8641.66 55.4— 61.8 2.35 3.99 8
eremicus (Northern ) 55.42—-—_ 51.8— 59.3 — — 6
“mojavensis” 55.934+1.14 52.1— 59.5 2.29 4.09 16
Providence Mt. area 57.38+1.18 54.0— 58.8 1.68 2.93 8
Mecca area 54.30+2.18 50.4— 58.1 3.07 5.65 8
“yumanensis’ (Lower Col. R.) 52.80+1.16 49.5— 57.6 2.40 4.55 17
“yumanensis’” (Upper Col. R.) 54.21+2.10 49.5— 58.1 3.16 5.18 9
SW Utah area 55.17+——_ 51.6— 57.2 — —- 6
cactophilus (C, SE Ariz.) 53.23+1.48 48.6— 59.5 2.88 5.41 15
Bill width at center of nostrils
eremicus (Southern ) 6.650.20 6.2— 7.2 0.64 4.81 11
eremicus (Northern ) 6.71+0.28 6.1-— 7.3 0.39 5.81 8
“mojavensis”’ 6.78+0.18 6.4— 7.4 0.36 5.31 16
Providence Mt. area 6.68+0.18 6:3= 40 0.25 3.74 8
Mecca area 6.660.24 6.3- 7.2 0.35 5.26 9
“yumanensis”’ (Lower Col. R. ) 6.5440.14 6.0— 7.1 0.32 4.89 18
“yumanensis”’ (Upper Col. R.) 6.59+0.26 6.1-— 7.3 0.40 6.07 9
SW Utah area 6.90+——_ 6.2— 7.5 — — 6
cactophilus (C, SE Ariz.) 6.22+0.24 5.4— 7.0 0.50 8.04 18
Variation of Ladder-backed Woodpeckers 5
TABLE 3 (Continued)
Form X + 2SE Range SD CV N
Depth of back bar*
eremicus (Southern ) 4.51+0.20 3.5— 6.0 0.59 13.08 38
eremicus (Northern) 4.40+0.34 3.3- 6.1 0.77 17.50 21
“mojavensis”” 3.64+0.12 2.6-— 4.3 0.36 9.89 42,
Providence Mt. area 3.490.26 2.5- 5.4 0.52 14.90 16
Mecca area 3.54+0.24 2.7— 4.3 0.47 13.28 14
“yumanensis”’ (Lower Col. R.) 3.1340.12 2.4- 4,1 0.42 13.42 47
“yumanensis’ (Upper Col. R.) 3.36+0.28 2.6-— 3.9 0.42 12.50 9
SW Utah area 3.330.28 2.7— 3.9 0.43 12.91 9
cactophilus (C, SE Ariz. ) 3.61+0.12 2.8— 4.7 0.44 12.19 56
* Measurements are in millimeters, except for CV. Figures are for specimens
collected from April through early July (excessively worn specimens considered
unmeasurable were disregarded), except for depth of back bar, where males from
all months were utilized. Symbols and abbreviations as in Table 1.
Tables 1 and 2, but all 3 male and 3 female eremicus samples
show a mean length of gonys greater than corresponding
samples of “mojavensis”), and significantly deeper black back
bars (hence a blacker back; Tables 1 and 2). In the latter
3 features eremicus differs by as much or more from other
samples of cactophilus (including “yumanensis”) than it does
from “mojavensis.” Additionally, eremicus tends to have a
slightly wider bill and less white in the primaries and wing
coverts than does “mojavensis.” However, there is more white
and less black barring in the tail of eremicus than in “mojav-
ensis’ (6 percent of eremicus, 38 percent of “mojavensis,’ 28
percent of “yumanensis,” and 53 percent of Arizona cactophilus
specimens have fully barred 5th rectrices). Analysis of the
patterns found in the outermost (6th) rectrices showed
that, while 80-90 percent of cactophilus and “yumanensis”
specimens have the normal barred condition of D. scalaris,
65 percent of “mojavensis” and only 50 percent of eremicus
specimens exhibited such a condition. The other patterns
found in eremicus were particularly varied, including some
tending toward the patterns of D. nuttallii (however, most of
the “abnormal” patterns found in “mojavensis” tend toward
those of D. nuttallii).
D. s. mojavensis (van Rossem). Specimens from Kern
County, California, south through the foothills and western
portion of the desert to San Diego County constituted this
6 Proceedings of the Biological Society of Washington
sample. This putative subspecies was compared above with
D. s. eremicus. Compared with D. s. “yumanensis” (van Ros-
sem), which occupies the lower Colorado River Valley and
immediately adjacent deserts, “mojavensis’ exhibits signif-
icantly longer wings, tail and bill. The tail: wing ratio of the
two samples is almost identical. The black back bars of
“mojavensis” are significantly deeper than those of “yuman-
ensis,’ and hence the back of this form is blacker than that
of “yumanensis.” However, “mojavensis” is virtually identical
to central and southeastern Arizona cactophilus in back
barring, and both of these have shallower black bars than
the blacker-backed eremicus (Tables 1 and 2). “Mojavensis”
tends to have a broader bill, less barred 6th rectrices, large
outer (5th) rectrices with about as much white, and blacker,
less spotted primaries than “yumanensis.”
D. s. yumanensis (van Rossem). This subspecies was based
(van Rossem, op. cit.) entirely on its paler (whiter), less black -
coloration than other D. s. cactophilus and D. s. “mojavensis.”
Comparison of “yumaiensis” with “mojavensis” and eremicus
has been accomplished above. Compared with cactophilus
specimens from central and southeastern Arizona, “yumanensis”
has: wings of about the same length; a slightly longer tail; about
the same tail : wing ratio; a bill which averages longer (males
only); a slightly wider bill; similar 6th rectrix patterns; sig-
nificantly whiter (less black-barred) 5th rectrices; and, a
tendency toward more white spotting and barring in the wings.
Additionally, the black back bars of “yumanensis” are signif-
icantly shallower than those of cactophilus, and hence “yuwman-
ensis’ has a very white back (Tables 1 and 2).
D. s. cactophilus (Oberholser). A sample of this race taken
in central and southeastern Arizona has been compared above
with D. s. eremicus, “mojavensis” and “yumanensis” (see also
Tables 1 and 2).
In Table 3 measurements of samples of the above four forms
taken from April through July are compared with certain other
critical samples. Only males are included in the table; avail-
able female samples gave similar results. The small sample
from the northern edge (roughly the region of Baja California
from Ensenada and the Sierra de Juarez south to a line passing
Variation of Ladder-backed Woodpeckers il
east from San Quintin) of the range of eremicus is very like
that of the southern sample. However, the northern eremicus
sample is more variable in outer rectrix markings, and both
sexes tend to have longer bills and more narrow bars than
do birds from the southern eremicus sample.
Samples from the Mecca area (Indio, Thermal and Mecca
in Riverside County, California), and from the Providence
Mountain region (San Bemardino County, California), and
several specimens from Calexico (Imperial County, Califor-
nia), Calipatria (Imperial County, California) and the west
side of Laguna Salada, Baja California, exhibit intermediacy
between the birds of the southern California foothills and
those of the lower Colorado River Valley. The Mecca area
specimens average very close to “mojavensis” in most features,
with extreme individuals appearing quite typical of “ywman-
ensis,’ although the black back bars are deeper, like those of
“mojavensis.”
Table 3 shows variation in populations of Ladder-backed
Woodpeckers inhabiting the Colorado River Valley and its
tributaries. The samples are from the Lower Colorado River
(from Blythe, California, south to the rivers mouth), the
Upper Colorado River (from Blythe north to the southern tip
of Nevada), and southwestern Utah (from the Virgin River
and its tributaries in western Washington County, Utah, and
northwestern Mohave County, Arizona). A cline of increas-
ing wing length from south to north is suggested by the data
in Table 3, as well as by data from the female samples not
included in the table. A corresponding cline is evident in tail
length, as indicated in the data (Table 3) for over-all tail
length and for length of the fourth rectrix. Male and female
samples likewise show such a cline for bill length. The
averages for length of gonys in the southwestern Utah sample
(mean of 6 males = 17.02 mm, mean of 7 females = 15.07
mm) were greater than those obtained for other samples of
southwestern D. scalaris, except D. s. eremicus and D. s.
lucasanus. A south to north cline of increasing bill depth is
also suggested by data from these three samples. The south-
western Utah and Upper Colorado Valley samples exhibited
intermediacy in back barring between Lower Colorado Valley
8 Proceedings of the Biological Society of Washington
“yumanensis,’ and both “mojavensis” and central Arizona
cactophilus.
Although various tendencies are evident in “mojavensis,”
“yumanensis,” and other populations of southwestern Ladder-
backed Woodpeckers currently assigned to the subspecies
cactophilus, they are not sufficiently marked to merit separate
subspecific recognition for any of them. The paler-backed
Colorado Valley population is otherwise very similar to that
of central Arizona, and there is great overlap in their various
characters. The populations assigned by van Rossem (op. cit.)
to “mojavensis” have many attributes of birds from south-
western Utah, as well as from central Arizona, and even of
“yumanensis.” Moreover, all of these populations resemble
one another, and differ from Baja California populations
(eremicus, lucasanus) in having a proportionally shorter tail.
They also have somewhat shorter bills, and much whiter backs
(i.e., significantly shallower black bars) than does eremicus. -
These differences from eremicus unite these variable popula-
tions, and separate them from that form. Therefore, it seems
prudent to continue to regard mojavensis and yumanensis as
synonyms of cactophilus, while recognizing eremicus as a
distinct subspecies.
D. s. cactophilus and D. s. eremicus meet along the border
of California and Baja California. A tendency toward cac-
tophilus has already been noted in the northern sample of
eremicus. All specimens examined from the California side of
the Mexican-United States border are definitely cactophilus.
Specimens from the Nachaguerro Valley (Univ. Calif. Mus.
Vert. Zool. no. 52146) and the west side of Laguna Salada
(M.V.Z. no. 52145), Baja California, are nearest cactophilus
in appearance, including tail: wing ratio. Two specimens
from Los Palmitos at the southeast end of the Sierra de Juarez
are nearer eremicus than cactophilus. A winter specimen
(U.S.N.M. no. 196279) from Ensenada, Baja California, is
intermediate in its features, and has a tail : wing ratio of 0.62
(cactophilus). It is possible that winter birds wander some-
what, so this bird may not represent the breeding population
in the vicinity of Ensenada. All small samples and individual
specimens from south of Ensenada, and from the east base of
Variation of Ladder-backed Woodpeckers 9
the Sierra de Juarez westward are referable to eremicus.
Three specimens from San Felipe, Baja California, are erem-
icus. The center of the area of intergradation between cac-
tophilus and eremicus appears to extend from the Pacific Coast
just south of the California-Baja California border eastward
across the north edge of the Sierra de Juarez, and southward
along the east edge of that range. There is probably no
contact between eremicus and cactophilus in the barren region
from the mouth of the Colorado River south to the vicinity of
San Felipe. Hence, there exists only a narrow area of contact
between the two subspecies in the foothills west, north, and
east of the Sierra de Juarez.
I am indebted to authorities of the following institutions for
the loan of specimens for this study: California Academy of
Sciences, University of California Museum of Vertebrate
Zoology (MVZ) at Berkeley, University of California at Los
Angeles (Dickey Collection), Carnegie Museum, Chicago
Natural History Museum, Los Angeles County Museum, San
Diego Museum of Natural History, San Diego State College
(Dept. of Biology), United States National Museum, Univer-
sity of Utah (Dept. of Zoology), and Zion National Park
Collection. Dean Amadon made helpful suggestions bene-
fitting the manuscript. I wish especially to thank authorities
of the U.S. Fish and Wildlife Service, which supported most
of the investigation, and of the U.S. National Museum, where
the studies were conducted.
LITERATURE CITED
OBERHOLSER, H.C. 1911. A revision of the forms of the ladder-backed
woodpecker (Dryobates scalaris [Wagler]). Proc. U.S. Nat.
Mus., 41: 139-159.
VAN RossEM, A. J. 1942. Four new woodpeckers from the western
United States and Mexico. Condor, 44: 22-26.
10 Proceedings of the Biological Society of Washington
Ud). OO7TS
h
Vol. 81, pp. 11-22 Slr : April 30, 1968
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
A NEW GENUS AND SPECIES OF CRAWFISH
FROM THE SOUTHEASTERN UNITED STATES
(DECAPODA, ASTACIDAE)!
By J. F. Frrzpatrick, JR.. AND JAMES F. PAYNE
Department of Zoology, Mississippi State
University, State College 39762
Taxonomically, the crawfishes assigned to the North Amer-
ican genus Cambarus have been recognized as the most dif-
ficult of the Cambarine, and remarkable convergence of
morphological form among them has caused students to make
tenuous assignments of relationships. The new species de-
scribed below has as its closest relatives the members of the
Cristatus Section (Hobbs, 1955:98) of the genus Cambarus:
C. cristatus Hobbs (1955:95), C. prominens Hobbs (1966:
110) and C. valleculus Fitzpatrick (1967:163). Further, this
new species provides information explaining the “distant rela-
tionships” (see below) of the members of the Cristatus Section
to other members of Cambarus.
This new crawfish could, on the basis of gonopodal charac-
ters and those of the annulus ventralis, be assigned to the
Limosus Section (Ortmann, 1905:108) of the genus Orcon-
ectes; however, it shares many more characters with the
members of the Cristatus Section (Hobbs, 1955:98) of the
genus Cambarus. Among these characters are the broad
antennal scale, the short-fingered chela with a cristiform row
of tubercles along the inner margin of the palm, and features
of the carapace which do not exist in any member of the
genus Orconectes. Only in the two members of the genus
Faxonella (Creaser, 1933:21) and in the members of the
Barbatus (Hobbs, 1942:35) and Planirostris (Penn, 1953:75)
1 Supported in part by NSF Grant GB-4719 to the senior author.
2—Proc. Biot. Soc. WAsH., Vou. 81, 1968 (11)
12 Proceedings of the Biological Society of Washington
Groups of the genus Procambarus is there a marked resem-
blance in the conformation of the carapace to that of this new
species and other Cristatus Section crawfishes. The annulus
ventralis, while unique, is within the limits of variability one
might expect in the Cristatus Section. The most distinctive
feature is the sternal modification of the first form male: the
conspicuous heavily setiferous lateral lobes of the sternum at
the bases of the third and fourth pereiopods have been
observed in no other members of the Cambarinae except in
the three species that compose the Cristatus Section.
In some species of Cambarus (C. striatus Hay, 1902:437;
C. reduncus Hobbs, 1956:61; and C. floridanus Hobbs, 1941:
114) the tips of the first pleopods are surrounded by a setifer-
ous hood. This hood, however, has a different morphological
source from the hood of Cristatus Section crawfishes. In all
of these the terminal elements of the pleopod of first form
males are relatively long and, in the absence of a sternal —
modification, would project beyond the ventral margin of the
coxae of the pereiopods. In C. striatus, C. reduncus and C.
floridanus the median portion of the sternum is depressed,
producing a cup-like depression in the third and fourth ster-
nites and “flaring” lateral margins. The setiferous hood of
these species arises principally from these lateral margins. On
the other hand, in members of the Cristatus Section the
sternites of the third and fourth pereiopod segments produce,
bilaterally, a longitudinal, submarginal ridge which forms a
depression to receive the pleopods. The setiferous hood arises
from these ridges and from the associated coxopodites. Thus,
these modifications seem to be two different solutions to the
same problem, with superficially similar end results.
The only true Orconectes-like characteristic of this new
species is the relatively straight central projection of the first
pleopod of the male. While the angle of the central projection
to the main shaft of the appendage in two species of Cambarus
[C. obeyensis Hobbs and Shoup (1947) and C. pristinus
Hobbs (1965)] is much less than in more typical members
of the genus, in neither is it less than 50 degrees. Further-
more, the central projection is slender and acuminate in C.
cristatus and its three relatives, whereas, in C. obeyensis, C.
New Genus and Species of Crawfish 13
pristinus and their relatives, it is more blade-like and has a
terminal notch. The latter two species, on the basis of several
characteristics, are clearly related to the “bartonii complex”
of the genus, rather than to the members of what has been
designated the Cristatus Section. We believe that the gono-
pods of neither of these assemblages could be conceived to
be of a type which was ancestral to the other and suggest that
in Hobbs’ (1967:11) diagram of relationships of related craw-
fishes, based on gonopodal characters, that the “Cristatus
Section” diverged from the pro-Orconectes line between those
branches leading to Procambarus pecki and pro-Cambarus.
The Cristatus Section, as previously recognized, exhibits
several apparently primitive characteristics. In all crawfishes,
a short, broad areola, such as is found in members of the
Cristatus Section, is considered primitive. The movable an-
nulus ventralis is found in the Cambarellinae, Procambarus,
Paracambarus, Troglocambarus, a few Cambarus (but none
so freely movable as cristatus and its relatives), and very
primitive Orconectes. It is simple and in some respects
resembles that of the primitive troglobitic Orconectes. The
straight central projection of the new species and the relative
slenderness of the appendage are also primitive characteristics.
Hobbs, in describing C. cristatus, was aware of the unique-
ness of this group. He stated (1955:98): “Cambarus cristatus
is not closely related to any described species of the genus.”
Neither Hobbs (1966:110) in his description of C. prominens
nor Fitzpatrick (1967:163) in describing C. valleculus mod-
ified this position. This conclusion suggested an independent
origin of the Section, but not until the discovery of the new
species described below with its primitive gonopod could this
be reasonably documented.
The angular reflection of the central projection in previously
described species of the Cristatus Section crawfishes, then,
represents a convergence (parallelism) in this characteristic
in the Cristatus Section and other members now assigned to
the genus Cambarus. Although this is the first recorded
incidence of parallelism in pleopod characteristics in Cam-
barinae crawfishes, other incidences of parallelism are known
to occur. Hobbs (1942:25) discussed the parallels of mor-
14 Proceedings of the Biological Society of Washington
phology in burrowing crawfishes in which the areola is long
and extremely narrow or obliterated and the rostral, post-
orbital and cervical spines are much reduced or absent. The
carapace similarities between members of the Barbatus and
Planirostris Groups of Procambarus, Faxonella, and the “Cris-
tatus Section” are mentioned above. In all of these there is
a reduction of marginal spines on the rostrum and a reduction
of the acumen. The margins of the rostrum tend to be converg-
ing or clypeate in outline. The cervical spines are reduced or
absent. The areola is moderately broad or broad. These
similarities are, however, indicative of ecologic habit rather
than evolutionary relationship. All generally occupy tem-
porary bodies of water and seem to dig simple burrows when
these waters dry. Interestingly, too, these species groups seem
to be, generally, complimentary in distribution. The Barbatus
and Planirostris Groups are distantly related members of the
genus Procambarus. The two species of Faxonella are not
closely related to any other crawfish. Fitzpatrick (1963:60)
noted that no other crawfishes possess the markedly unequal
terminal rami of the first pleopods. Further, only in Faxonella
are the terminal elements bent mesiad. Although the tips of
the central projections cross in Cambarus dissitus Penn (1955:
13), the primary bend of the terminal elements is caudad and
the mesial deflection is secondarily derived in what is clearly
a close relative of C. fodiens Cottle (1863:217), C. byersi
Hobbs (1941:118) and C. hedgpethi Hobbs (1948:224). As
we have indicated above, the “Cristatus Section” is unique.
Most contemporary students of crawfish evolution accept
the concept that a Procambarus-like population was ancestral
to the modern Cambarinae genera. The recent genera, then,
represent specializations from a basic stock. These ideas have
been discussed by Hobbs (1940:55); furthermore, Hobbs
(1967:2) has described a troglobitic species of the genus
Procambarus from Alabama and has further discussed the
origins of the generic stocks (pp. 8-15). The existence of two
primitive Cambarus-like species groups, which are at best
very distantly related, strongly suggests a dual emergence of
the Cambarus-like characters from a Procambarus-Orconectes
ancestral population. One of these groups is represented by
New Genus and Species of Crawfish 15
C. obeyensis, C. pristinus and their relatives, and the other by
C. cristatus and its relatives. In light of this apparently diphy-
letic origin of the genus as presently recognized, we propose
that a new genus be erected to receive the members of the
Cristatus Section and the new species described below:
Hobbseus new genus
Diagnosis: Rostrum broadly subovate and subspatulate. Areola 2.6-
7.5 times longer than wide and constituting 28.7-35.5 per cent of total
length of carapace. Cervical spines or tubercles lacking; suborbital angle
much reduced or absent. Antennal scale approximately one-half times
as broad as long with lamellar portion considerably broader than thick-
ened lateral portion. Sternum of first form male with conspicuous,
ventrally projecting, setiferous prominences at bases of third and fourth
pereiopods. Terminal elements of first pleopod of first form male almost
straight or bent; if bent, mesial process slender and projecting caudad
beyond central projection for considerable distance; central projection
acute to subsetiform, without subterminal notch. Cephalic margin of
annulus ventralis of female joined to sternum by flexible membrane.
Type-species: Here designated. Cambarus cristatus Hobbs, 1955:95
(USNM no. 96985).
Etymology: We take pleasure in naming this new genus in honor of
Dr. Horton H. Hobbs, Jr., in recognition of his efforts toward a better
understanding of crawfishes.
Range: This genus is known only from the watershed of the Tom-
bigbee River in Alabama and Mississippi and the upper portion of the
Pearl River watershed in Mississippi.
KEY TO THE SPECIES OF THE GENUS Hobbseus
1. Central projection of first pleopod of first form male almost
Strreat O'ln tiie ey led elk RUN Sa op ll Ne H. orconectoides, n. sp.
Central projection of first pleopod of first form males bent _______ 9)
2. Central projection bent at angle of 90° or less to axis of main
shatt/of appendages) Sa ea H. cristatus (Hobbs, 1955:95)
Central projection bent at angle greater than 90° to axis of main
CU aVEUA ECO) Bee) DY OVS) OVS Fe hope eae hen ND SU SUD Gy PE nat DOM Ne St ela MV SO 3
3. Central projection bent at approximately 110° angle to main
shaft of appendage; mesial process extending approximately 14
its length beyond tip of central projection; pleopods symmetrical
SDD AN TURE PROP DRED OO aa A H. prominens (Hobbs, 1966:110)
Central projection bent at approximately 120° angle to main
shaft of appendage; mesial process extending approximately 14
its length beyond tip of central projection; pleopods asymmetrical
H. valleculus (Fitzpatrick, 1967:163)
Fics. 1-12. Hobbseus orconectoides. (1) Mesial view of first pleopod
of holotype; (2) mesial view of first pleopod of morphotype; (3) dorsal
view of carapace of holotype; (4) lateral view of first pleopod of mor-
photype; (5) lateral view of first pleopod of holotype; (6) lateral view
of carapace of holotype; (7) ischiopodite and basipodite of third perei-
opod of holotype; (8) epistoma of holotype; (9) antennal scale of holo-
type; (10) annulus ventralis of allotype (sperm plug not illustrated);
(11) distal podomeres of cheliped of holotype; (12) sternal processes
and basal podomeres of third (III) and fourth (IV) pereiopods of para-
typic male, Form I. (Punctations and setation omitted from all figures. )
New Genus and Species of Crawfish 17
Hobbseus orconecioides new species
Diagnosis: Pigmented, eyes normal. Rostrum subplane or slightly
excavate, subspatulate without marginal spines or tubercles. Areola
9.64—-4.72, (avg. 3.54) times longer than wide, constituting 28.9-33.8
(avg. 31.9) per cent of entire carapace length. Suborbital angle obsolete.
Antennal scale more than one-half as broad as long, lamellar portion
widest distal to mid-length. Chela with row of cristiform tubercles along
inner margin of palm. First pleopod of male with central projection
almost straight, slender with acute tip; mesial process slender, distal half
directed at about 45 degree angle to main shaft of appendage; mesial
process extending distad beyond tip of central projection. First form
male with conspicuous setiferous lateral lobes on sternum at bases of
third and fourth pereiopods. Pleopods symmetrical. Annulus ventralis
(Fig. 10) freely movable; subovate; depression in cephalomedian portion
of cephalic third; sinus arising in caudal portion of depression, moving
sinuously caudad, disappearing near caudal margin.
Holotypic male, Form I: Body subovate, compressed laterally. Abdo-
men narrower than cephalothorax (8.1 and 9.0 mm, respectively). Width
of carapace less than depth in region of caudodorsal margin of cervical
groove (9.0 and 9.5 mm). Areola moderately broad (4.0 times longer
than wide), with irregular row of punctations mesial to each bran-
chiocardiac groove and irregular median longitudinal row, only three
punctations across narrowest part. Cephalic section of carapace 2.2 times
as long as areola; areola 31.2 per cent of total length of carapace.
Rostrum 1.3 times longer than wide, subspatulate, with tiny acute acumen,
but lacking marginal spines, tubercles or angles, margins not thickened
and only slightly elevated; tip of rostrum reaching tip of distal end of
peduncle of antennule; upper surface slightly excavate, with few setifer-
ous punctations; rostrum depressed cephalically (Fig. 6). Subrostral
ridges weak, barely evident in dorsal aspect.
Postorbital ridges well developed; cephalic ends lacking tubercles or
spines. Suborbital angle obsolete. Branchiostegal spine small, but acute.
Cervical tubercle nearly obsolete. Carapace moderately punctate dorsally
and cephalolaterally. Abdomen longer than carapace (23.1; 20.5 mm).
Cephalic section of telson with two acute spines in each caudolateral
corner.
Epistoma (Fig. 8) broader than long, subplane with elongate cephalo-
median tubercular projection. Antennules normal with small spine on
lower surface of basal segment. Antennae broken. Antennal scale (Fig.
9) broadest distal to midlength, 1.91 times longer than wide, terminating
distally in strong spine.
Chela (Fig. 11) with palm broad, only slightly inflated and somewhat
depressed, length of inner margin of palm equal to width; upper surface
of palm with prominent scattered squamous setiferous tubercles on inner
half, tubercles becoming two rows just above inner margin, remainder
of upper surface with less prominent tubercles; inner margin with single
18 Proceedings of the Biological Society of Washington
cristiform row of 18 closely placed tubercles; lower surface punctate,
fingers not gaping; upper and lower surfaces of both fingers with sub-
median ridge flanked by setiferous punctations; tubercle on lower surface
at base of dactyl lacking; opposable margin of movable finger with four
subequal tubercles along proximal two-thirds, and crowded minute
denticles along distal one-third; opposable margin of immovable finger
with one small, two large and one yet larger tubercle (proximal to distal )
along proximal two-thirds and crowded minute denticles along distal
one-third.
Carpus of cheliped longer than broad; grooved dorsally, with scattered
setiferous punctations, inner margin with numerous prominent cristiform
tubercles and one strong spine located on ventromedian margin at distal
two-thirds of its length; lower surface with strong acute spine on
cephalolateral margin. Upper surface of merus with row of five small
squamous tubercles along distal one-half and two acute spines on distal
margin; lower surface with row of 12 small subacute spines along mesial
margin and row of 12 small subacute spines along lateral margin, strong
acute spine in distal lateral corner and submedian row of three small
acute spines in distal one-tenth. Ischiopodite with row of five small
cristiform tubercles on inner margin.
Maxillipeds and coxae of third through fifth pereiopods heavily setose.
Ischiopodites of third sereiopods with strong simple hooks; hooks
extending proximally beyond bases of ischiopodites (Fig. 7). Prominent
setiferous projections at bases of third and fourth pereiopods (Fig. 12).
First pleopods reaching to coxopodites of third pereiopods when
abdomen flexed, lying deeply embedded between ventrally projecting
sternal projections; distal portion terminating in two parts; central projec-
tion corneous, tip acute, slender, nearly straight; mesial process non-
corneous, slender, tapering from base to tip, distal half directed caudad
approximately 45 degrees, extending beyond tip of central projection
(ates, al, 5)
Morphotypic male, Form II: Differs from holotype in following re-
spects: width of palm greater than length of inner margin; setiferous
tubercles of upper surface of palm less prominent; tubercles along oppos-
able margins of both fingers very weakly developed. Ischiopodite of
third pereiopod lacking hooks. First pleopod with both tips noncorneous,
stouter, shorter, and more blunt (Figs. 2, 4).
Allotypic female: Differs from holotype in following respects: width
of palm less than length of inner margin; inner margin of palm with
single cristiform row of 12 closely placed tubercles along proximal two-
thirds, row of three squamous tubercles along distal one-third; setiferous
squamous tubercles of upper surface of palm less prominent; tubercles
of opposable margins of both fingers less prominent.
Annulus ventralis freely movable, subovate; depression in cephalo-
median portion of cephalic third; sinus originating at caudal margin of
depression and moving sinuously caudad to disappear near caudal margin
(Fig. 10). Sperm plug in cephalic portion of sinus (not figured).
New Genus and Species of Crawfish 19
TABLE 1. MEASUREMENTS (in mm)
Holotype Allotype Morphotype
Carapace
Height 9.5 9.3 8.2
Width 9.0 8.6 7.6
Length 20.5 18.8 18.8
Areola
Length 6.4 6.4 6.1
Width 1.6 IBS ise
Rostrum
Length 5.5 5.0 5.7
Width 44 Al 4.3
Chela
Palm length (inner margin) G5) 4.2 5.4
Palm width ee 4.5 4.4
Palm length (outer margin) 15.7 9.2 11.9
Dactyl length 8.2 5.5 6.5
Type locality: Woodland pond near Rock Hill community, about 4%4
mi. N of Starkville, Oktibbeha County, Mississippi (R14E, T19N, Sec.
1). The pond is approximately 9 ft. deep, but H. orconectoides is found
only along the shore, usually associated with emergent vegetation and
shallow water.
Etymology: The trivial name of this species refers to the superficial
resemblance of the gonopod and annulus to members of the genus
Orconectes.
Disposition of types: The holotype, allotype and morphotype are
deposited in the United States National Museum (nos. 129191, 129193,
and 129194, respectively ). Most of the remaining specimens are topopara-
types which are located at the USNM (26 641, including the figured
paratype; 1411, 19; 14j; 19j) the Museum of Comparative Zoology
(26 61; 36 6Il; 599; 54 4j; 59 9j), Tulane University (2¢ 41;
36 SII; 59 9;5¢4 $j; 59 9j), the Ohio State Museum (26 61; 36 oI;
529; 56 6j; 529 9j), and Mississippi State University (136 ¢I1; 32
6 Sil; 399 2; 454 3j; 469 2j; and 38 unsexed juveniles). Other para-
types are located at Mississippi State University (1¢II; 29 9; 1éj).
The type series represents collections from September through May in
various years (1915-1967). None were collected in summer months.
Presumably they burrow during the dry summer months. A collection
from a burrow was made on 6 May 1967 at Rock Hill.
Variations: Very slight variation, almost exclusively in spinose and
tubercular ornamentation of the cheliped, was observed. All are within
the limits set by the types.
Relationships: These are discussed in preceding paragraphs.
20 Proceedings of the Biological Society of Washington
Associates: At the type locality, other crawfishes present are P. hayi
(Faxon, 1884:108) and O. mississippiensis (Faxon, 1884:123). At other
places P. acutissimus (Girard, 1852:91) and C. hedgpethi Hobbs inhabit
the same area.
Remarks on Range: H. orconectoides is never abundant and is mod-
erately plentiful only at or very near the type locality. Diligent search
has not located this species outside the watershed of Sand Creek, a small
creek system confined to Oktibbeha County, Mississippi, and an effluent
to Catalpa Creek which flows into Tibbie Creek. The latter forms the
boundary between Clay and Lowndes Counties, west of the Tombigbee
River. Such a restricted range is compatible with the primitive status
which we propose for this species.
Acknowledgments: We are grateful to Mesrs. James P. Thaxton and
Houston F. Lowery who helped with collections and Dr. Horton H.
Hobbs, Jr., for his critical evaluation of our ideas of the relationships of
this species.
LITERATURE CITED
Cotte, T. J. 1863. On the two species of Astacus found in upper
Canada. Canadian J. Industry, Sci., Arts. N.S. 8: 216-219.
Creaser, E. P. 1933. Descriptions of some new and poorly known
species of North American crayfishes. Occ. Pap. Mus. Zool.,
Univ. Michigan, no. 275: 21 p.
Faxon, W. 1884. Description of new species of Cambarus; to which is
added a synonymical list of the known species of Cambarus
and Astacus. Proc. Amer. Acad. Arts Sci. 20: 107-158.
Firzpatrick, J. F., Jr. 1963. Geographic variation in the crawfish
Faxonella clypeata (Hay) with the definition and defense of
the genus Faxonella Creaser. Tulane Stud. Zool. 10: 57-79.
1967. A new crawfish of the Cristatus Section of the genus
Cambarus from Mississippi (Decapoda, Astacidae). Proc.
Biol. Soc. Washington 80: 163-168.
Grrarp, C. 1852. A revision of the North American Astaci, with
observations on their habits and geographical distribution.
Proc. Acad. Nat. Sci. Philadelphia. 6: 87-91.
Hay, W. P. 1902. Observations on the crustacean fauna of Nickajack
Cave, Tennessee and Vicinity. Proc. U.S. Nat. Mus., 25
(1292): 417-439.
Hosss, H. H., Jr. 1940. On the first pleopod of the male Cambari.
Quart. J. Florida Acad. Sci., 5: 55-61.
1941. Three new Florida crayfishes of the subgenus Cam-
barus. Amer. Midl. Nat. 26: 110-121.
1942. The crayfishes of Florida. Univ. Florida Publ., Bio.
Sci. Ser. 3(2): 179 p.
1948. A new crayfish of the genus Cambarus from Texas,
with notes on the distribution of Cambarus fodiens (Cottle).
Proc. U.S. Nat. Mus. 98: 223-231.
New Genus and Species of Crawfish 21
. 1955. A new crayfish of the genus Cambarus from Missis-
sippi. Proc. Biol. Soc. Washington 68: 95-100.
. 1956. A new crayfish of the genus Cambarus from North
Carolina and South Carolina (Decapoda, Astacidae). J.
Elisha Mitchell Sci. Soc. 72: 61-67.
1965. A new crayfish of the genus Cambarus from Ten-
nessee, with an emended definition of the genus (Decapoda,
Astacidae). Proc. Biol. Soc. Washington 78: 265-274.
1966. A new crayfish from Alabama with observations on
the Cristatus section of the genus Cambarus (Decapoda,
Astacidae). Proc. Biol. Soc. Washington 79: 109-116.
1967. A new crayfish from Alabama caves with notes on
the origin of the genera Orconectes and Cambarus. Proc.
U.S. Nat. Mus. 123(3621): 17 p.
AND C. S. SHoup. 1947. Two new crayfishes (Decapoda,
Astacidae) from the Obey River drainage in Tennessee. J.
Tennessee Acad. Sci. 22: 138-145.
OrtTMANN, A. E. 1905. The mutual affinities of the species of the
genus Cambarus, and their dispersal across the United States.
Proc. Amer. Philos. Soc. 44: 91-136.
Penn, G. H. 1953. A new burrowing crawfish of the genus Procam-
barus from Louisiana and Mississippi. Tulane Stud. Zool.
1: 71-76.
1955. A new Cambarus of the diogenes Section from North
Louisiana. Tulane Stud. Zool. 3: 73-81.
99, Proceedings of the Biological Society of Washington
me 0673
Vol. 81, pp. 23-30 N April 30, 1968
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
AN UNDESCRIBED SUBSPECIES OF LEIOCEPHALUS
RAVICEPS COPE (SAURIA: IGUANIDAE)
FROM WESTERN CUBA
By ALBERT SCHWARTZ AND ORLANDO H. GARRIDO
Dept. of Biology, Miami-Dade Junior College,
Miami, Fla., U.S.A. 33167 and
Instituto de Biologia, Academia de Ciencias de Cuba,
Capitoli Nacional, La Habana, Cuba
Leiocephalus raviceps Cope is one of five Cuban members
of this exclusively West Indian genus. Long very poorly
represented in collections, L. raviceps has recently been shown
(Schwartz, 1960) to have a peculiarly disjunct distribution:
two subspecies, raviceps and uzzelli, occur along the southern
coast of Oriente Province between the Bahia de Santiago and
Cajobabo, whereas a third subspecies, klinikowskii, occurs on
the northern shore of the Peninsula de Hicacos in northern
Matanzas Province (Schwartz, 1960; Buide, 1966). Schwartz
(1960:81) noted that there seemed a good possibility that
other populations of L. raviceps remained to be discovered
throughout Cuba. This supposition has very recently been
shown to be correct, although the lizards were found far to
the west of the previously known most western population
(klinikowskii).
A colony of L. raviceps was discovered in 1965 by Sr. Miguel
L. Jaume, director of the Museo Felipe Poey, La Habana,
Cuba, and Dr. Zdenek Vogel, of Prague, Czechoslovakia.
Additional specimens from this population were collected by
the junior author and Sr. Jaume in 1967, so that there is now
an adequate series of these lizards available. The colony
occurs in the vicinity of Cortés, in western Pinar del Rio
Province, near the base of the Peninsula de Guanahacabibes,
3—Proc. Bio. Soc. WasH., Vou. 81, 1968 (23)
24 Proceedings of the Biological Society of Washington
removed some 320 kilometers to the southwest of L. r. klini-
kowskii in northern Matanzas Province. Specimens of this
distinctive new subspecies have been deposited in the collec-
tion of the Instituto de Biologia, Academia de Ciencias de
Cuba (IB), the Museo Felipe Poey (MFP) and in the collec-
tion of the senior author (ASFS). We are very grateful to
Sr. Jaume for assisting us in all ways in this project and to
Ing. Héctor Sagué D. for his cooperation and his making our
collaboration possible. We have examined, in addition to
specimens of the new subspecies, material of the subspecies
raviceps and uzzelli in the collection of Brigham Young Uni-
versity (BYU) and the senior author and wish to thank Wil-
mer W. Tanner for bringing to our attention the presence of
these specimens in the BYU collection.
The Pinar del Rio population of L. raviceps is quite different
from the three more eastern subspecies; we take very great
pleasure in honoring Sr. Jaume, one of the discoverers of these -
lizards, by naming them.
Leiocephalus raviceps jaumei new subspecies
Holotype: IB 1349, an adult male from San Waldo, 4 km N Cortés,
on the road between Cortés and Isabel Rubio, Pinar del Rio Province,
Cuba, one of a series collected 6 March 1967 by Orlando H. Garrido
and Miguel L. Jaume.
Paratypes: IB 1341-45, MFP (9 specimens), ASFS V11458-66, same
data as holotype; IB 179, IB 196—98, same locality as holotype, 3 May
1967, O. H. Garrido and M. L. Jaume.
Diagnosis: A subspecies of L. raviceps characterized by a combination
of small size (males to 61 mm, females to 49 mm snout-vent length),
lateral fields in both sexes overlaid with circular to subcircular inky
black spots from the neck to the groin and onto the upper surface of the
hindlimbs and basal lateral and ventral portions of the tail, and throats
of females and subadult and juvenile males dotted with black or brown
to dark gray discrete subcircular spots; low number of scales at one half
midbody, moderate number of fourth toe tricarinate subdigital scales,
modally 4 median head scales, modally 4 frontoparietal scales, the
frontoparietal row frequently incomplete, and high mean number of
loreal scales.
Distribution: Known only from the type locality.
Description of holotype: An adult male with the following measure-
ments (in millimeters) and counts: snout-vent length 61, tail 77, tip
regenerated; snout to anterior border of tympanum 13.4, head width
10.9; supraocular scales 7/7; loreals 6; temporals 12; enlarged auricular
New Subspecies of Cuban Lizard 25
scales 3/3; prefrontal row 3 scales, complete; 4 median head scales;
frontoparietal row 4 scales, incomplete; dorsal crest scales occiput to
vent 65, dorsal crest scales occiput to axilla 27; scales around half body
at midbody 33; fourth toe subdigital tricarinate scales 27/26; supraorbital
semicircles complete; parietal scales in contact.
Dorsum (as preserved) medium brown (whitish in life), with dorsal
crest scales cream, and dorsum densely punctate with creamy to pale
golden scales, especially from about midtrunk onto the base of the tail;
head scales paler yellowish tan (clear brownish in life), suffused with
whitish; lateral fields between fore- and hindlimbs slightly darker brown
than dorsum with three longitudinal rows of inky black circular or sub-
circular spots, those on the neck larger than those on the sides; of the
three longitudinal rows on the sides, the median row contains the largest
spots; sides of tail and upper surface of hindlimbs with black to very
dark brown (reddish to violet in life) spots; lateral fields additionally
with scattered cream scales and bordered below by a pale reddish tan
(grayish brown in life) longitudinal stripe between the fore- and hind-
limbs with two longitudinal series of small black dots below the pale
ventrolateral stripe; postocular spot solid, black, with a vertical white
(pure yellow in life) line separating its posterior third dorsally; throat
pale gray with some very small and vague gray flecks laterally, inter-
mixed with cream to white iridescent scales; chest and venter white,
with a few vague pale gray spots on the chest, venter immaculate except
for scattered iridescent cream to white scales laterally; underside of
fore- and hindlimbs concolor with venter, the former immaculate, the
latter with scattered brown flecks; tail cream below, rather heavily
punctate with dark gray spots basally, and concolor with dorsum above,
with about 17 faintly darker crossbands or widely opened chevrons to
the regenerated tip.
Variation: The series of 28 L. r. jaumei is composed of 19 males with
snout-vent lengths between 42 mm and 61 mm, and nine females with
snout-vent lengths varying from 32 mm to 49 mm; the sexes may be
distinguished externally by the presence of two pairs of enlarged postanal
scales in males. Scale counts (means and extremes) from this series
are: dorsal crest scales between occiput and vent 61-72 (mean 65.3);
dorsal crest scales between occiput and axilla 22-30 (26.4); one half
midbody scales 27-34 (30.5); fourth toe subdigital tricarinate scales
93-28 (25.4); loreals 3-8 (4.7); temporals 11-14 (12.4); supraoculars
7/7 (10 specimens), 5/6 (1), 6/6 (5), 6/7 (5), 7/8 (3); prefrontal row
3 scales, usually complete (24 of 27 specimens); median head scales 4—6
(mode 4); frontoparietal row 2—5 scales (mode 4), usually complete (15
of 26 specimens) but with a high frequency of incomplete rows; semi-
circles usually complete (22 of 25 specimens); and parietals usually in
contact (24 of 26 specimens).
Male L. r. jaumei (as preserved) vary in dorsal pattern. In younger
males, the dorsum is weakly zonate, with a central darker brown zone
(bisected by the cream median dorsal crest scales) and a pair of more
26 Proceedings of the Biological Society of Washington
lateral tan zones, often heavily flecked with cream scales. In older males,
the back becomes more unicolor tan to medium brown (grayish or
whitish in life) and the zonation is lost or at least greatly obscured by
the admixed dorsal cream colored scales. There is seldom any indication
of either dashes or herringbones in the dorsal pattern. The lateral fields
are always somewhat darker than the dorsum immediately above them
and are heavily marked with about two to four longitudinal series of
inky black, circular, subcircular or even ovoid spots, the spots extending
onto the base of the tail along its sides and onto the upper surface of
the hindlimbs, where the spots are more dot-like. The lateral fields also
contain scattered pale iridescent scales which were presumably creamy
in life. The postocular blotch is present and black in all males, but often
there is a pale to white (yellow in life) vertical bar which cuts off the
posterior third of the blotch from the anterior two thirds. The blotch
is delimited anteriorly by a vertical white (yellow in life) bar from the
posterior portion of the eye and the vertical white bar is bordered
anteriorly in turn by a vertical black to dark gray bar. The lateral
nuchal spots are usually somewhat larger than the more posterior spots,
and there are only one or two rows of blotches on the neck. Subadult
males (snout-vent lengths 42 mm to 50 mm) have the throat patterned .
with scattered dark gray ovoid spots, not aligned into longitudinal
series; with increasing age and size, these dots become increasingly less
obvious so that in large males they are absent or may be represented
only by some very vague grayish spots, intermixed with iridescent cream
scales, along the sides of the throat. The lateral margins of the venter
in subadult males likewise are distinctly punctate with dark brown dots;
in some adult males these persist along the sides of the venters, whereas
in others they disappear, leaving only some areas of iridescent paler
scales on the belly margins. The underside of the hindlimbs is speckled
with brown and iridescent pale scales as is also the underside of the
proximal portion of the tail.
Female L. r. jaumei are distinctly zonate dorsally, the central darker
brown region (which is longitudinally bisected by the pale median dorsal
crest scales) having a series of about twelve diagonally placed (apices
directed posteriorly) chevron remnants between the occiput and the
sacrum. The lateral fields resemble in detail those of the males, and
there is no sexual dichromatism nor ontogenetic change in the presence
of the circular black lateral spots. The lateral fields in females are clearly
bordered below by a longitudinal pale line between the limbs and there
are small black dots below this line which continue onto the abdomen as
scattered dark brown flecks. The throat in females is spotted with
circular to ovoid dark brown to dark gray spots which extend onto the
chest. The underside of the hindlimbs lacks dark brown spotting except
along the anterior margin of the thighs, and the underside of the tail
is brown spotted, at least proximally. A striking feature of female jaumei
(in contrast to the other subspecies) is the anterior continuation in most
specimens of the dorsal dark zone as a pair of fairly broad dark slightly
New Subspecies of Cuban Lizard on
divergent stripes onto the head shields as far as the supraocular scales;
thus the occipital region of female jaumei has a distinctly pale-and-dark
striped aspect.
Comparisons: L. r. jaumei, in having lateral fields with black spots,
is so very distinct from the remaining three subspecies that extended
comparison of pattern is unnecessary; raviceps, uzzelli, and klinikowskii
have been figured (Schwartz, 1960) and none of them has lateral black
spots or has the females and subadult males with spotted throats. L. r.
jaumei differs in many other details of pattern, and as far as known
lacks the herringbone or dashed patterns of the other subspecies (the
nominate subspecies also lacks dorsal dashes). In general intensity of
color, jaumei is apparently slightly darker than the pale sandy to almost
white klinikowskii, is paler than uzzelli, and is paler than raviceps, which
itself has a faded or washed-out appearance.
In dorsal crest scales between occiput and vent and between occiput
and axilla, and in fourth toe scales, jaumei does not differ significantly
from the other three subspecies. As far as one half midbody scales is
concerned, the much lower mean (30.5 + .64 = twice standard error
of mean) in jaumei is significantly different from those of klinikowskii
(35.3 + .65), uzzelli (32.4 + .70), and raviceps (32.3 + .70). The three
western subspecies are all larger than jaumei, with male raviceps reaching
a maximum size of 80 mm snout-vent length, uzzelli males to 71 mm and
klinikowskii males to 69 mm. Females of all races are comparably larger
than female jaumei. L. r. jaumei resembles raviceps and uzzelli in modally
having 7/7 supraoculars; klinikowskii modally has 6/6 supraoculars. In
modally having 4 median head scales, jaumei also differs from klinikowskii
(mode 6) but is like the two Oriente races. The mode of 4 frontoparietal
scales in jaumei is like that of uzzelli, but differs from raviceps and
klinikowskii (mode 5). L. r. jaumei has the highest incidence of incom-
plete frontoparietal row (42 per cent); frequencies in the other subspecies
are: uzzelli (35 per cent), raviceps (22 per cent), and klinikowskii (9
per cent). In usually having the supraorbital semicircles complete, jaumei
resembles raviceps and klinikowskii and differs from uzzelli which more
often has the semicircles incomplete. The higher loreal mean (4.7) in
jaumei is greater than those of klinikowskii (4.2), raviceps (3.6), and
uzzelli (3.3).
Remarks: The occurrence of L. raviceps far to the west of its pre-
viously known range is extremely interesting. Doubtless there are still
other localized populations of this species scattered in seldom visited or
remote localities throughout Cuba; the increasing knowledge of the
distribution of L. stictigaster (see Schwartz and Garrido, in press) in-
dicates that much is yet to be learned of the details of distribution of
Cuban Leiocephalus. The very distinctive lateral and ventral patterns
of jaumei suggest that this subspecies has been long separated from the
balance of the species to the east. The environs of the Peninsula de
Guanahacabibes continue to reveal an increasingly rich and varied
herpetofauna as exploration in that area goes forward. It is interesting
28 Proceedings of the Biological Society of Washington
that in this same general region (but not in the immediate region of
Cortés) there is an endemic subspecies of L. macropus (koopmani Zug;
see Schwartz and Garrido, 1967, for details), another Leiocephalus which
is predominantly eastern in distribution but which also has several non-
Oriente subspecies scattered over the balance of Cuba.
Through the courtesy of Ernest E. Williams at the Museum of Com-
parative Zoology at Harvard University, we have examined a short series
of L. raviceps from Baracoa, Oriente Province; the species has been
previously known from this immediate region by a single specimen in
the Museo y Biblioteca de Zoologia de la Habana, formerly under the
care of Sr. Jaume. The additional five Baracoa specimens (MCZ 13371-
75) are all males with snout-vent lengths between 61 and 66 mm; the
lizards have been long preserved and some details of pattern (and of
course color) are no longer clear. These lizards are geographically
closest to L. r. raviceps, but are separated from that subspecies by the
Sierra de Purial (unless L. r. raviceps is continuous along the south and
north Oriente coasts in this region via Cabo Maisi, an area where the
species remains uncollected). The Baracoa specimens are unusual in
that, of the five, four have supraocular counts of 6/6; modally L. r.
raviceps has 7/7 supraoculars, although 12 of 44 specimens of this sub- .
species have 6/6 supraoculars. All Baracoa lizards have 3 loreals. Both
of us have collected extciusively in the Baracoa area, yet neither of us
has secured L. raviceps in that region. Possibly these lizards are not
from the immediate area of the city itself; considering that L. raviceps
is regularly, in Oriente, an inhabitant of extreme xeric regions and that
the immediate environs of Baracoa are very mesic (with L. macropus
asbolomus Schwartz and Garrido the dominant Leiocephalus in coastal
areas), it seems probable that L. raviceps has a very limited distribution
in the Baracoa area. It seems equally likely that, when the species is
finally encountered in that region, the local population will be found to
differ from L. r. raviceps to the south.
As pointed out above and elsewhere (Schwartz, 1960:75-76; Schwartz
and Garrido, 1967:29) L. raviceps occupies extremely arid coastal areas
in Oriente; Schwartz (1960:79-81) and Buide (1966:7-9) commented
on the restriction of L. r. klinikowskii to white beach sands with sparse
vegetational cover. Buide (loc. cit:8) noted that the Matanzas lizards
were reluctant even to enter the more heavily wooded areas of the
peninsula but would do so if hard pressed or in danger. All eastern
localities of L. raviceps are coastal with the exception of a short series
in the southern foothills of the Sierra de Purial, north of Cajobabo.
The specimens of L. 7. jaumei were secured between 300 meters and
one kilometer to the north of the hamlet of San Waldo. The length of
the road between Cortés and Isabel Rubio is about 11 kilometers, but
for no more than 4 kilometers along this stretch of road the soils are
gray or ash-colored sands; on both sides of these gray sands are
found soils which are the typical yellow clays of this region. The gray
sands support a wooded growth, including a kind of palm as well as low
New Subspecies of Cuban Lizard 29
herbs. The area of gray sands, although narrow as far as the area
crossed by the road is concerned, may be much longer than the road
transect indicates. L. 1. jawmei was encountered by the junior author on
the bleakest portions of the sandy area. In the better wooded portion of
the gray sands were found L. stictigaster and Ameiva auberi.
L. r. jaumei began, on 6 March 1967, to emerge from their burrows
under fallen palm trunks at about 1000 hours, and were most active at
1100 hours. If unfrightened, the lizards remained near their burrows or
(preferably ) on the fallen palm trunks above the burrow mouths. When
disturbed, the lizards ran with great speed toward the burrow and many
remained 2 or 3 inches from the burrow mouth in an attitude of observa-
tion. Others quickly entered the burrows and remained there. At other
times, lizards which were disturbed away from their burrows ran beneath
the fallen palm trunks. One female contained an egg ready for deposi-
tion, but no juveniles were observed on 6 March. A pair was seen in
copulation.
We do not know if L. r. jaumei occurs elsewhere in the Cortés region
or in western Pinar del Rio Province. If this subspecies is really limited
to such precise an ecological situation as it seems to be, it may well
be that jaumei is indeed restricted to the area about the type locality.
It also seems logical to expect that there are other relict populations of
L. raviceps which remain yet unknown. If the species, outside of Oriente,
demands very specific soil and vegetational conditions for survival, the
discovery of other extra-Oriente populations of L. raviceps seems more
than probable, provided the collector stumbles by chance into the precise
locality where these ecological conditions are met and where the lizards
may occur. Persistence of relict extra-Oriente populations of two Leio-
cephalus, one of which (macropus) inhabits mesic forested situations
and the other (raviceps) bleak and xeric situations, in the general region
of the Peninsula de Guanahacabibes, is a most interesting phenomenon.
Considering also that western Pinar del Rio is occupied by two other
Leiocephalus (stictigaster, the clearly dominant Pinar del Rio species,
and carinatus), it would be most interesting to know the precise details
of distribution of all four species in this region, and the ecological inter-
relationships of the four species involved. Most remarkable is the fact
that the two species which are relict in this region (raviceps, macropus )
presumably at one time had an island-wide distribution (much as
carinatus does today); with changing geography and changing ecologies,
these two species are represented in western Pinar del Rio by limited
or extremely circumscribed populations. Further details of the distribu-
tion of L. raviceps in the area intervening between the known ranges of
klinikowskii and jaumei will be most interesting to determine.
Specimens examined (other than L. r. jaumei): L. r. raviceps, mouth
of Rio Yateras, 10 mi. E U.S. Naval Base, 43 (ASFS V11220-31, BYU
30258-63, BYU 30285-309); L. r. uzzelli, mouth of Rio Hatibonico,
14 (ASFS V11215-19, BYU 30241-42, BYU 30246-52).
30 Proceedings of the Biological Society of Washington
LITERATURE CITED
Bume, Mario S. 1966. Reptiles de la Peninsula Hicacos. Poeyana,
ser. A., no. 21: 1-12.
ScHWARTZ, ALBERT. 1960. Variation in the Cuban lizard Leiocephalus
raviceps Cope. Proc. Biol. Soc. Washington, 73: 67-82, 11
figs.
AND OrLaNpo H. Garripo. 1967. A review of the Cuban
iguanid lizard Leiocephalus macropus Cope. Reading Public
Mus. and Art Gallery, Sci. Publ. 14: 1-41, 4 figs.
(in press). Four new subspecies of Leiocephalus
stictigaster from Cuba. Nat. Mus. Canada.
We OG 75 ———————
Vol. 81, pp. 31-36 April 30, 1968
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
TWO NEW SUBSPECIES OF BIRDS FROM THE
PHILIPPINES AND COMMENTS ON THE
VALIDITY OF TWO OTHERS
By S. Ditton Rretey AND D. S. Rasor
Secretary, Smithsonian Institution, Washington, D.C. 20560
Research Professor in Biology, Institute of Research,
Mindanao State University, Marawi City, Philippines
In the course of visits to southern Mindanao from 1963 to
1966, Professor D. S. Rabor collected a number of specimens
now at the U.S. National Museum. Among these collections
which will eventually be reported on as part of a work on
Mindanao birds are new subspecies of an owl and a babbler
and series of two other species which demonstrate the in-
validity of previously described subspecies.
In the collection is one specimen of a Scops Owl from
Mindanao which differs from all described subspecies of this
widespread species and may be known as:
Otus scops mirus new subspecies
Holotype: USNM No. 519154, adult male collected by D. S. Rabor
19 April 1963, on Hilong-hilong Peak, Agusan Province, Mindanao
Island, Philippines. Original No. 35409.
Description: This is a typical small Otus scops in size and color tone,
similar to O. s. malayanus, except more coarsely barred and streaked with
blackish, the black streaks particularly heavy on the head, broader and
wider even than manadensis. The white feathers on the median wing
coverts are heavily tipped with black and brown, not clear black as
in sunia, malayanus and distans. The wing formula shows tenth (outer)
and first primaries equal in length, sixth and seventh equal and longest,
eighth longer than fifth, and fourth and ninth equal.
Below, this single specimen shows a good deal of white on the abdo-
men and undertail coverts, similar to malayanus although the chest is
coarsely streaked not finely vermiculated as in that subspecies. The
tarsus is feathered for more than half its length, the distal portion is bare.
4—Proc. Biot. Soc. WasH., Vou. 81, 1968 (31)
32 Proceedings of the Biological Society of Washington
Measurements: Wing 131, tail 58.5, culmen (from cere) 13, tarsus
22 mm.
Range: Known only from one specimen from the type locality some
twenty miles northeast of Butuan, northeastern Mindanao.
Etymology: The name is from the Latin mirus, meaning unusual or
striking.
Remarks: In 1941, Delacour (p. 41) questioned whether certain
Philippine forms of the Scops owl, listed by Peters (1949, pp. 92, 93)
properly belonged to the species Otus scops. The forms then known to
Peters consisted of: calayensis (Calayan); longicornis (Luzon); min-
dorensis (Mindoro); romblonis (Banton and Romblon); cuyensis (Cuyo)
and mantananensis (from Mantanani Island, off Sabah, north Borneo).
Peters (1940, p. 96) kept a separate species, Otus manadensis for the
forms sibutuensis (Sibutu); steerei (Tumindao); manadensis (Celebes
or Sulawesi); and related forms. About the same time Stresemann (1939,
pp. 320, 321) pointed out quite correctly that manadensis from Celebes
is indeed a subspecies of Otus scops and probably siaoensis should be
considered a synonym of it. Related forms included by Stresemann
(1940, pp. 429, 430) are Otus scops mendeni Neumann from Peling and
O. s. albiventris from the Lesser Sundas.
The only Moluccan form examined (Ripley, 1959, p. 4) was Otus
leucospilus, and we would keep this as a separate species contra Peters.
The Philippine owls of this assemblage have been considered by
Hachisuka [1934 (III), pp. 56-62] to belong to several species. His
arrangement is shown in Table 1.
Recently one of us (S.D.R.) has had the benefit of discussions with
Dr. A. L. Rand and has examined the material in the Field Museum
collection as well as Dr. Rand’s manuscript notes. From this discussion
it would appear that longicornis does not belong with the scops assem-
blage but is closer to bakkamoena, as is whiteheadi from Luzon. On the
other hand, mindorensis, still known from only one specimen from
Mindoro in the British Museum, is perhaps closer to Otus scops and
belongs with those species closely related to the scops assemblage.
Regarding the scops group of owls then, we would assume that aside
from Otus scops mirus, one should place the offshore island forms as
subspecies of Otus elegans of the Riukyu Islands with the following
subspecies, Otus elegans elegans; e. botelensis; e. calayensis (synonym
batanensis); e. romblonis; e. cuyensis; e. sibutuensis (synonym steerei);
and eé. mantananensis.
Otus elegans is a larger species (wing mostly over 155 and up to 175
mm) than Otus scops with more uniform dark upperparts and underparts,
very finely vermiculated throughout. In some specimens the abdomen
is paler, but always with vermiculations present.
Next to the Mindanao form then we would place Otus scops mana-
densis (synonym siaoensis); with related subspecies, sulaensis, mendeni,
kalidupae, morotensis, albiventris, and tempestatis. We would leave
Otus mindorensis as a separate species.
Two New Subspecies of Philippine Birds 33
Thus the derivation of scops-like owls in the eastern Sunda and Moluc-
can islands including Celebes, may well have come from the Philippines
via a series of invasions, the links of which seem to have been cut off.
A clue is the Ryukyu and Botel Tobago forms of elegans found on small
islands. Perhaps the presently defined species scops came from the north,
and remnant representatives may still be discovered in Luzon to connect
with this unique Mindanao form.
A Mindanao series of 3 males and 3 females of Micromacronus leytensis
recently described by Amadon (1962, pp. 3-5) from Mt. Lobi on Leyte
demonstrate that the Mindanao population represents a new subspecies.
Micromacronus leytensis sordidus new subspecies
Holotype: USNM No. 519070, adult male collected by D. S. Rabor,
2 Feb. 1964, on Mt. Matutum, Tupi, Cotabato Province, Mindanao
Island, Philippines, between 3,300 and 4,300 feet above sea level.
Original No. 39878.
Description: This form is larger and duskier, less bright in tone than
leytensis described from Mt. Lobi (1,500 ft. a.s.l.) on Leyte Island. The
specimens of sordidus have a slimmer bill, pale at the base, not all black.
They are pale greenish olive below, and lack the yellow forehead and
supercilium. The elongated white back and flank plumes are similar to
leytensis but shorter, not reaching the end of the tail. Females are
slightly paler below than are males.
Measurements (in millimeters )
wing tail culmen weight
M. l. leytensis (type) male 40.5 27 ND} —
female Al 28 10.5 —
M. l. sordidus (type) male 47 31 10 6.1 gms
2 males 43, 46 30, 31 10, 10 5.5, 6.5
2, females 43, 44 30, 31 10, 11 5.5, 5.9
Etymology: The name is from the Latin sordidus, referring to the dull
coloration.
A juvenile female with smaller measurements and soft, rather fluffy
plumage (June 27) would appear to be not long out of the nest. (Wing,
43; tail, 26.5; culmen, 9 mm; weight, 5.5 gms). Compared to the two
adult females it is more buffy above, with warm yellowish cinnamon
edges to the wing feathers. The specialized feathers of the back are
present although softer and shorter; not exceeding the wings in length.
Below, this specimen is dull whitish with traces of warm brown on the
breast and flanks. The specialized feathers of the flanks appear as soft
white filoplumes. As Amadon notes, this species would be difficult to
characterize as a babbler (Muscicapidae, Timaliinae). Perhaps only the
specialized feathers prevent it from being thought of as an uniquely
34 Proceedings of the Biological Society of Washington
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(sofoodsiodns sdoos jo y1ed) sisuatopuyw sniOC sisuaLopunu sdoos snjoC SISUBLOPUIUL SIULODIBUO] SNIC
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Two New Subspecies of Philippine Birds 35
plumaged leaf-bird (Aegithinidae). Basally these specialized feathers
appear somewhat downy in texture and in this area the soft barbs are
bordered with black, a point not mentioned by Amadon although it
appears in his photograph (1962, p. 4, fig. 1).
Rhabdornis inornatus Ogilvie-Grant
A series of this species taken on Leyte Island from 1,500-2,500 feet
altitude by Rabor between 31 May and 1 July measure:
wing tail culmen
3364 85-87 53-55 (tail moult May 31) 19-21
22° 85, 86.5 53, 54 17, 18
These specimens agree in size and color with R. inornatus inornatus
from Samar.
In addition Rabor has collected the species on Mindanao; in Surigao
del Sur in May, in Agusan, Mt. Hilong-hilong, in April, and in Cotabato
on Mt. Matutum in February. Altitudes ranged from 1,100 feet in
Surigao to 5,000 feet on Hilong-hilong peak. Two subspecies have been
described from Mindanao, alaris Rand from southeastern Mindanao in
Davao, and zamboanga Rand and Rabor from western Mindanao in the
Zamboanga Peninsula on Mt. Malindang. The latter subspecies has been
separated from alaris on the basis of duller brown coloration. There is
no difference in size (Rand and Rabor, 1960, pp. 436-437). Based on
these facts we do not feel that the presence of two subspecies on Min-
danao is maintained by this new series from northern and southeastern
Mindanao. Freshly moulted August birds from Davao Province appear
more ruddy-brown, but worn birds are inseparable throughout the island.
Ruddy or chocolatey-backed birds appear also from the west in the
Zamboanga series. Intermediate birds appear in our new series. The
differences thus appear too small to us to warrant separation.
Chloropsis flavipennis (Tweedale)
Among interesting species found by Rabor was Chloropsis flavipennis,
known from Cebu where it has not been recorded in recent years and
may be extinct (Rabor, 1959, p. 40). Currently recorded only from the
islands of Cebu and Mindanao (Delacour in Peters, 1960, p. 303) it
was a surprise to find also this species on the island of Leyte, from
whence a series was secured.
The Leyte birds were found from 800-2,500 feet altitude on Mt.
Lobi and the Lobi range, Burauen. An August female is noted as nesting.
In addition specimens were taken on Mindanao at the following localities:
Sibahay, Lanuza, Surigao del Sur, near sea level; Mount Mayo, Mati,
Davao Province; and Mt. Hilong-hilong, Agusan Province from 500-
4,200 feet altitude.
From these specimens we are inclined to feel that flavipennis is a
36 Proceedings of the Biological Society of Washington
monotypic species, and that it is unwise to separate the Mindanao birds
as mindanensis Salomonsen (1953, p. 270) named on the basis of larger
size alone. For comparison, we list below the measurements of the
specimens available (including specimens kindly loaned by the Camegie
Museum, and the American Museum of Natural History).
culmen (to base
wing tail : of s
Cebu 8 ¢ ¢ 88 -97(93) 68( worn )-75(72) 23.5-25( 24.5)
m 2) 22 85 -91 68-71.5 24 -25
Leyte 4 ¢ ¢ 91 -98(96) 69-73 23.5-24
n 32Q 89.5-90 66-68 22 -24
Mindanao 3 ¢ ¢ 98 -101 71-80 24 -25
" 3292 89 -95 T4717 24 -25
There is a slight continuous cline in size from Cebu to Leyte to
Mindanao with more than 25 per cent overlap when the sexes are
compared, males with males or females with females.
LITERATURE CITED
Amapon, D. 1962. A new genus and species of Philippine bird. Con-
dor, 64(1): pp. 1-5. 1 color plate.
DeELacour, J. 1941. On the species of Otus scops. Zoologica. Sci.
Contrib. New York Zool. Soc., no. 17, 26(2): pp. 133-142.
HacuisuKA, THE MARQuEss. 1934. The Birds of the Philippine Islands.
Part 3. H. F. and G. Witherby, London, pp. 1-256.
Peters, J. L. 1940. Check-list of birds of the World. Vol. 4. Harvard
Univ. Press, Cambridge, xii + 291 pp.
1960. Check-list of Birds of the World, vol. 9. Museum of
Comparative Zoology, Cambridge, xii + 506 pp.
Rasor, D. S. 1959. The Impact of Deforestation on Birds of Cebu,
Philippines, with New Records for that Island. Auk, vol. 76,
pp. 37-43.
Ranp, A. L. AND Rasor, D. S. 1960. Birds of the Philippine Islands:
Siquijor, Mount Malindang, Bohol, and Samar. Fieldiana:
Zoology, vol. 35, no. 7, pp. 223-441.
Rietey, S. D. 1959. Birds from Djailolo, Halmahera. Postilla, no. 41,
pp. 1-8.
SALOMONSEN, F. 1953. Miscellaneous Notes on Philippine Birds.
Vidensk. Medd. fra Dansk naturh. Foren., vol. 115, pp. 205—
281.
STRESEMANN, E. 1939. Die Vogel von Celebes. Journ. fiir Ornith.,
87(3): pp. 299-425.
1940. Die Vogel von Celebes. Journ. fiir Omith., 88(3):
pp. 299-487.
IG TF
aoe
Vol. 81, pp. 37-40 ____—= April 30, 1968
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
NEW SUBSPECIES OF BIRDS FROM OAXACA
(AVES: PHASIANIDAE, TURDIDAE, PARULIDAE)
By Rosert T. Orr AND J. DAN WEBSTER
California Academy of Sciences, San Francisco, California
and Hanover College, Hanover, Indiana
Southwestern Oaxaca is a region that shows considerable
endemism as regards its avifauna. This has been pointed out
by Phillips (1966), who recently described a number of new
subspecies of birds from there. The most important geograph-
ical feature influencing this evolutionary trend is the Sierra de
Miahuatlan, a southeastern extension of the Sierra Madre del
Sur separated from the backbone of that range by the Rio Verde.
The Valley of Oaxaca to the north, the Isthmus of Tehuantepec
to the east, and the Pacific Ocean to the south further isolate
this mountain system. In the course of preparing a report
based on a year of continuous field work in this region,
sponsored by the Western Foundation of Vertebrate Zoology,
the American Museum of Natural History, and the California
Academy of Sciences, several new subspecies of birds have
been noted. Three of these are described here. The holotypes
are all deposited in the collection of the California Academy
of Sciences but the paratypes, unless otherwise indicated, have
not yet been divided between the sponsoring institutions and
the Mexican government.
We wish to thank Dr. Ned K. Johnson and Dr. Peter L.
Ames for the use of certain pertinent specimens in the collec-
tion of the Museum of Vertebrate Zoology.
Family PHASIANIDAE
Colinus virginianus harrisoni new subspecies
Holotype: Adult male; no. 65512 California Academy of Sciences; 5.1
miles SW San Gabriel Mixtepec, 1,800 ft elevation, Oaxaca, Mexico;
5—Proc. Bion. Soc. Wasu., Vou. 81, 1968 (37)
38 Proceedings of the Biological Society of Washington
8 May 1965; collected by Warren Rook; original no. 4277; wing 101 mm;
weight 156.1 grams.
Geographic distribution: Known only from the vicinity of the type-
locality.
Diagnosis: Male, similar to Colinus v. atriceps (Ogilvie-Grant) in
having top of head, chin, and throat black and superciliary stripe
essentially missing, but differing in that breast is chestnut, streaked with
black, instead of being solid black. Female, similar to C. v. atriceps.
Measurements (in millimeters): Four adult males (including holo-
type) from type-locality: length of wing (chord), 101.5 (100-103);
length of exposed culmen, 13.8 (13.5-14.5); length of tarsus, 27.8
(27.3-28.5). One adult female from type-locality: length of wing
(chord), 101; length of exposed culmen, 14.0; length of tarsus, 29.8.
Weights (in grams): Three adult males from the type-locality, 151.7
(143-156.1); 1 adult female from the type-locality, 152.5.
Remarks: Colinus virginianus is one of the most geographically vari-
able species of birds in North America. At least 15 subspecies are
recognized as occurring in Mexico alone, and three of these are known
from the state of Oaxaca. These are C. v. atriceps, known only from
Putla, the type-locality; C. v. thayeri Bangs and Peters from. northeastern
Oaxaca; and C. v. coyolcos (P. L. S. Miiller) from the lower Pacific slope
of extreme eastern Oaxaca and western Chiapas. C. v. thayeri is one of
the white-throated races. C. v. coyolcos has the black essentially restricted
to the head and throat and a narrow white stripe on either side of the
head. The syntypes of C. v. atriceps were examined by one of us (Orr)
in the British Museum (Natural History) in the summer of 1966, and
the male differs from C. v. harrisoni in having the chest solid black
instead of streaked.
Specimens examined: The holotype and 4 paratypes (3 males and 1
female) from the type-locality.
Etymology: Named in honor of Ed N. Harrison of the Western Foun-
dation of Vertebrate Zoology, Los Angeles, California.
Family TURDIDAE
Turdus assimilis oaxacae new subspecies
Holotype: Adult female, no. 65513 California Academy of Sciences;
Jamaica Junction, 4 miles N San Gabriel Mixtepec, 2,400 ft elevation,
Oaxaca, Mexico; 20 November 1964; collected by J. Stuart Rowley,
original no. 3498; wing 119 mm, weight 79 grams.
Geographic distribution: Southern Oaxaca and the state of Guerrero.
Diagnosis: Differs from both Turdus a. assimilis Cabanis and T. a.
lygrus Oberholser in being more olivaceous, slightly darker, and less
brown dorsally, as well as more olivaceous on breast and flanks, also
darker than lygrus on breast and flanks. As contrasted with T. a. leu-
cauchen Sclater, oaxacae is paler and more olivaceous dorsally as well as
more olivaceous on breast and flanks. Eye ring orange in life as in
New Subspecies of Mexican Birds 39
assimilis and lygrus (fide notes and annotation on labels by J. S. Row-
ley), not lemon yellow as in lewcauchen.
Measurements (in millimeters); Three adult males from the Sierra de
Miahuatlan: length of wing (chord), 127 (124-130); length of tail, 95.8
(93.7-97.8); length of exposed culmen (2 averaged), 17.6 (16.7-18.5);
length of tarsus, 30.9 (30.7-31.3). Three adult females from the Sierra
de Miahuatlan: length of wing (chord), 125 (122-127); length of tail,
98.1 (94.5-101); length of exposed culmen, 18.0 (16.8-19.8); length
of tarsus, 31.3 (29.4-33.9).
Weights (in grams): Two adult males from the Sierra de Miahuatlan,
82.7 and 69.5; 3 adult females from the Sierra de Miahuatlan, 80.8
(79.0-86.7 ).
Remarks: Turdus assimilis was revised by Miller and Griscom (1925).
Since then Moore (1937) described T. a. renominatus from southern
Sinaloa. This was synonymized under T. a. lygrus in the Mexican Check-
list II (Miller et al, 1957). The range of lygrus was given as from
Sinaloa south to Oaxaca and east to Morelos.
The subspecies lygrus, as we now conceive it, ranges from Morelos to
Jalisco and north to central Sinaloa. A single specimen from Colima is
intermediate between lygrus and oaxacae. The ranges of assimilis and
leucauchen remain as stated in the Mexican Check-list II save that the
former is extended slightly south to central Oaxaca, on the basis of single
specimens from Cerro San Felipe and Progreso, and the latter is extended
slightly southwest to Cerro Baul in extreme eastern Oaxaca on the basis
of four specimens.
Foxing is severe in skins of this species but virtually all of the speci-
mens examined in the present study were collected since 1949.
Specimens examined: Thirty-one non-juvenals including the holotype
and 30 paratypes: Guerrero: Omilteme, 22 (4 in Calif. Acad. Sci. collec-
tion, 18 in Mus. Vert. Zool.); Oaxaca: Cerro Verde, 30 km E Santa Rosa
(Lachao Nuevo), 8,600 ft, 5; Jamaica Junction, 4 mi. N San Gabriel
Mixtepec, 2,400 ft, 1; 9 mi. W San Gabriel Mixtepec, 2,600 ft, 1; 5.1 mi.
SW San Gabriel Mixtepec, 1,800 ft, 1; Sinai, 10 km E Nopala, 7,200 ft, 1.
Etymology: Named after the Mexican State of Oaxaca.
Family PARULIDAE
Ergaticus ruber rowleyi new subspecies
Holotype: Immature male (skull not fully ossified); no. 65514 Cali-
fornia Academy of Sciences; 30 km E Lachao Nuevo, 8,600 ft elevation,
Oaxaca, Mexico; 27 October 1964; collected by Warren Rook; original
no. 3953; wing 62 mm; weight 8.6 grams.
Geographic distribution: High mountains of Guerrero and southern
Oaxaca.
Diagnosis: Brighter red throughout than either Ergaticus r. ruber
(Swainson) or E. r. melanauris Moore with auriculars silvery as in ruber.
Color of dorsum bright Ruby (Palmer and Reilly, 1956) rather than off
40 Proceedings of the Biological Society of Washington
Ruby in direction of Rose as in ruber, or off Ruby in direction of Scarlet
as in melanauris.
Measurements (in millimeters): Four males (including the holotype)
from the Sierra de Miahuatlan: length of wing (chord), 62.5 (61-64);
length of tail, 56.2 (55.8-60.6); length of exposed culmen, 7.4 (7.1-
7.7); length of tarsus, 18.3 (17.2-19). Four adult females from the
Sierra de Miahuatlan: length of wing (chord), 59.5 (57-62); length of
tail, 55.8 (53.2-58.3); length of exposed culmen (3 specimens), 7.6
(7.3-7.8); length of tarsus, 17.5 (17.0-18.7).
Weights (in grams): One adult male from the Sierra de Miahuatlan,
8.9; 4 adult females from the Sierra de Miahuatlan, 8.6 (8.49.0).
Remarks: We have examined two specimens of Ergaticus ruber from
Durango, an adult from 15 miles west of El Salto, and a juvenile from
Cerro Huehuento. Both confirm the race melanauris described by Moore
(1937) from Chihuahua and Sinaloa. The adult has the characters
ascribed to melanauris by Moore; the bird in juvenal plumage has the
auriculars metallic brown in contrast to the silvery color of comparable
specimens of ruber. Specimens examined from Morelos, Michoacan, and
Distrito Federal represent the typical race ruber.
It might be expected that the population of Ergaticus ruber in southern
Oaxaca would show some similarities to E. versicolor (Salvin) of Central
America. Comparison of specimens from Oaxaca with a series from the
vicinity of San Cristobal, Chiapas, fails to indicate this. The pink “frost-
ing” anteriorly and the black auriculars of E. versicolor are not exhibited
to any degree by any specimens of E. ruber.
Specimens examined: Eighteen, including the holotype and 17 para-
types, as follows: Guerrero: Omilteme, 6 (1 Calif. Acad. Sci., 5 Mus.
Vert. Zool.); Cuapongo, 1 (Mus. Vert. Zool.); Oaxaca: 30 km E Lachao
Nuevo, 8,600 ft, 1; Rio Molina, 7,300 ft, 10.
Etymology: Named after J. Stuart Rowley, Research Associate, Cali-
fornia Academy of Sciences.
LITERATURE CITED
Miter, A. H., FRIEDMANN, H., Griscom, L. AND Moore, R. T. 1957.
Distributional check-list of the birds of Mexico. Part II.
Pacific Coast Avif., no. 33, 436 p., 7 pls.
Miter, W. DEW. AND Griscom, L. 1925. Further notes on Central
American birds, with descriptions of new forms. Amer. Mus.
Novit. 184: 1-16.
Moors, R. T. 1937. Four new birds from northwestern Mexico. Proc.
Biol. Soc. Wash., 50: 95-102.
. 1937. New races of Myadestes, Spizella and Turdus from
northwestern Mexico. Proc. Biol. Soc. Wash. 50: 201-206.
PALMER, R. S. AND Rettiy, E. M., JR. 1956. A concise color standard.
Amer. Omith. Union, Albany, N. Y. 8 p., color chart.
Puitures, A. R. 1966. Further systematic notes on Mexican birds.
Bull. British Orith. Club 86: 86-94, 1 fig.
W667 3
Vol. 81, pp. 41-44 ™» April 30, 1968
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
A NOTE ON THE ATLANTIC SPECIES OF THE
STARFISH GENUS LINCKIA
By Maureen E.. DOWNEY
Smithsonian Institution, Washington, D.C.
The identities of those species of Linckia which have been
reported from the Atlantic have been in a somewhat confused
state for many years. The genus occurs in the Atlantic, Pacific,
and Indian Oceans, in tropical and subtropical waters. Five
species have been reported from the Atlantic in the area from
about 35° north to 35° south on the western side, and from
about 16° north to 16° south on the opposite side, from the
intertidal zone to depths in excess of 750 m.
A careful search through the literature and an examination
of specimens in the collections of the U.S. National Museum
and the Museum of Comparative Zoology at Harvard, as well
as of specimens kindly supplied by Jerald Halpern, Institute
of Marine Sciences, University of Miami; Lawrence McClos-
key, Duke University; and Maximo Cerame-Vivas, University
of Puerto Rico, has convinced me that not more than three
species of Linckia have been collected in the Atlantic to date.
A brief diagnosis of each follows.
Linckia guildingii Gray 1840
Ophidiaster ornithopus Miller and Troschel, 1842.
Scytaster stella Duchassaing, 1850.
Linckia ornithopus: Liitken, 1859.
Rays long, slender, well-rounded; usually five, sometimes four to seven;
often unequal, due to autotomous division. Dorsal plates thick, numer-
ous, somewhat convex, sometimes imbricate, covered with fine granules.
Papulae 3-9, papular areas small. Marginal plates larger than dorsals,
quadrangular, in two regular rows. Madreporite usually single, but may
be double. Comet forms are common. Illustrated in Verrill, 1915, pl.
XXVIII, Fig. 3.
6—Proc. Biot. Soc. WasH., VoL. 81, 1968 (41)
42 Proceedings of the Biological Society of Washington
Distribution: Florida to Brazil, and West Africa (Sao Thomé).
Shallow water.
Linckia bouwieri Perrier 1875
Linckia nodosa Verrill, 1915.
Rays long, slender, rounded, usually five. Many dorsal plates enlarged,
tumid, covered with granules. Pores per papular area 10-18. Marginals
in two somewhat irregular, sometimes discontinuous rows. Ocular plates
not very prominent, covered with granules. Madreporite single. Illus-
trated in Verrill, 1915, pl. XXIX, Figs. la-lb (as Linckia nodosa).
Distribution: North Carolina to Florida, and Cape Verde Islands.
35-130 m.
Linckia nodosa Perrier 1875
Lincki formosa Mortensen, 1933.
Linckia bullisi Moore, 1960.
non Linckia nodosa Verrill, 1915.
Rays very slender, rounded, long, five. Many dorsal plates enlarged,
tumid, covered with granules. Pores per papular area 2-8. Marginals
in two rather regular rows, many with a small spine or tubercle in center
of plate. Ocular plates prominent, raised, rounded, without granules.
Madreporite single. Illustrated in Moore, 1960, Fig. 1 (as Linckia
bullisi).
Distribution: Florida to Brazil, and St. Helena, 230-475 m.
Apparently Mortensen and Moore both overlooked Perrier’s Linckia
nodosa when describing their species. This, of course, was easy to do,
since Perrier gave the locality for L. nodosa as “unknown.” However,
L. formosa and L. bullisi agree with L. nodosa in having very slender
rays, a number of enlarged tumid plates, 2-8 pores per papular area, and
a small spine or tubercle on many of the marginal plates.
Linckia guildingii is widespread in the Caribbean and tropical Atlantic,
in shallow water. It is very common in reef areas. The other two species
are much rarer and occur mainly in deeper water. Linckia bouvieri has
been collected only from the Straits of Florida north to North Carolina
and on the other side of the Atlantic in the Cape Verde Islands. Linckia
nodosa, on the other hand, seems to occur only south of the Straits of
Florida and at St. Helena, an interesting reiteration of the affinity of the
St. Helena echinoderm fauna with the Antillean fauna.
Key to the Atlantic Species of Linckia
1. Large tumid plates occurring randomly on disc and arms _______ 2
INowtumid Slates! oe sa as cereal ee Le Linckia guildingii
2. Minute spines on some lateral plates; ocular plates without
Fes eh 01D (ats yyyteud ners bites RA WAL EEO Cyt eta ails MANTEL decile nd Linckia nodosa
No spines on lateral plates; ocular plates covered with granules
ete oP Al gee SARUM te ioe SAN = ORT LA QO ils Re RUN AEE Linckia bouvieri
Atlantic Species of Linckia 43
LITERATURE CITED
Criark, H. L. 1941. Reports on the Scientific Results of the Atlantic
Expeditions to the West Indies, under the joint auspices of
the University of Havana and Harvard University. Mem.
Soc. Cubana Hist. Nat. “Felipe Poey,” vol. XV, no. 1, p. 50.
Ducwassainc, P. 1850. Animales Radiates des Antilles. Paris.
Gray, J. E. 1840. A synopsis of the Genera and Species of the Class
Hypostoma (Asterias Linnaeus). Ann. Mag. Nat. Hist., vol.
vi, p. 284, London.
Lurxen, C. F. 1859. Bidrag til Kundskab. om de ved Kysterne af
Mellen-og Syd-Amerika levende Arter af Sostjerner. Vidensk.
Medd. dansk. naturh. Foren. Kbn., 1858, pp. 25-96.
Moore, D.R. 1960. Linckia bullisi, a new Asteroid from the Northeast
Coast of South America. Bull. Mar. Sci. Gulf and Carib.,
vol. 10, no. 4, pp. 414, 1 fig.
MorTENSEN, Tu. 1933. The Echinoderms of St. Helena (other than
Crinoids). Vidensk. Medd. dansk. naturh. Foren. Kbn., Bd.
93, p. 430, pl. XXII, Figs. 2-4.
MULLER, J. AND F. H. TroscueL. 1842. System der Asteriden, 3,
Braunschweig, 4°, 134 pp., 12 pls.
Perrier, E. 1875. Revision de la collection de Stellerides du Museum
d’Histoire Naturelle de Paris. Arch. Zool. Exper. et Génér.,
Tome IV, pp. 414-417.
VerRRILL, A. E. 1915. Report on the Starfishes of the West Indies,
Florida, and Brazil. Bull. Lab. Nat. Hist., S. U.I., vol. VII,
no. l, p. 93, pls. XIII, Figs. 2-2a; XXIX, Figs. la—lb.
44 Proceedings of the Biological Society of Washington
Go Bb Ze
Vol. 81, pp. 45-52 April 30, 1968
~SPROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
CEOCHASMA, A REMARKABLE NEW LAND SNAIL
FROM COLIMA, MEXICO (GASTROPODA,
PROSOBRANCHIA, HELICINIDAE )!
By Frep G. THOMPSON
Florida State Museum, University of Florida
While conducting field work in Colima during 1966 the
author collected an unusual helicinid snail that has several
shell peculiarities. Its radular structure substantiates both its
generic distinctness and its isolated phylogenetic position.
Features of its inner marginal teeth and its central teeth place
the genus in the subfamily Proserpininae, tribe Vianini, but
it is distinguished from all members of this subfamily by its
more primitive helicinoid capituliform complex. The genus
and species are described below:
Ceochasma new genus
Type-species: Ceochasma phrixina new species.
Generic Diagnosis: Readily distinguished from all other known mem-
bers of the family Helicinidae by a deep slitlike sinus that extends back
along the suture of the last 4% whorl. Schazicheila Shuttleworth and
Viana H. and A. Adams also have sinuses in the aperture, but the
structures differ in the three genera. Some species of Schazicheila have
a small notch in the upper corner of the peristome, accommodating a
small peglike extension of the operculum, and never much deeper than
the thickness of the operculum. Viana has a deep notch in the outer lip
about halfway between the upper corner of the peristome and the
periphery. The notch apparently serves as an interlocking device between
the two sexes during copulation. The notch develops late in the growth
of the shell, and has no similarity in location or structure to that of
Ceochasma.
Besides the characteristic siphon notch, the shell is readily distin-
1 This work was supported by the National Institutes of Health Research Grant
GM 12300.
7—Proc. Bion. Soc. WaAsuH., Vou. 81, 1968 (45)
46 Proceedings of the Biological Society of Washington
guished from other Mexican helicinids by its size, shape and sculpture.
In these characters it resembles some species of Helicina Lamarck, s. g.
Tristramia Crosse, sec. Oxyrhombus Crosse and Fischer, but is distin-
guished from all species of that section by its much coarser spiral chords.
Ceochasma has an operculum consisting of concentric rings with an
acentric nucleus, a well-developed calcareous plate with an oblique
lamellarlike projection along the columellar margin, and a horny plate
that extends beyond the margin of the calcareous plate. In these charac-
ters it resembles Tristramia, but the columellar projection is longer than
occurs in Tristramia, and the nuclear region of the calcareous plate is
thinner and more concave.
Etymology: The generic epithet is derived from the Greek xeéo,
meaning cleft, and xacua meaning gaping mouth, and alludes to the
clefted aperture. The gender of the name is feminine. The specific
name is taken from the Greek @¢p\e and évos, and refers to the rippled
appearance of the sculpture.
Ceochasma phrixina new species
Holotype: University of Florida Collections (UF) 20138, collected
2 August 1966 by Fred G. Thompson.
Paratypes: UF 20139 (16), UMMZ 228990 (5), Museo Nacional de
Mexico (3); collected with the holotype. UF 20140 (12); collected
29 May 1966 at the type-locality. UF 20141 (15); collected 30 May
1966 at the type-locality.
Type-locality: A collapsed limestone ridge 0.3 mi. E Tamala, Colima;
500 ft alt.
Tamala lies in the valley of a small stream that drains to the southeast
for a few miles into the Rio Salado. Immediately southeast of the town
is a low limestone ridge that consists of huge stone blocks and fragments
that resulted from a subterranean collapse of the ridge. The vegetation
in the area is a dense xeric scrub, with little ground cover except for a
moderate amount of debris and tuff around the base of the ledge and
among the boulders. At the time of my visit during the rainy season
snails were found at the base of the ridge aestivating close to the ground
on the boulders.
Description of shell (Figs. 1-6): Large. Solid, but not excessively
thick, depressed-conical, 0.66—-0.78 times as high as wide, acutely carinate
at periphery; sides of spire very weakly convex; base below periphery
strongly convex; aperture weakly auriculate, with upper lip separated
from preceding whorl by deep, narrow cleftlike sinus extending along
suture for about 4% of last whorl. Sinus present at all stages of post-
embryonic growth, cemented in from behind with progressive shell
growth. Aperture oblique in lateral profile. Lip with strong thick
reflection along peripheral and basal margin; in lateral profile lip mod-
erately recurved above and slightly more below periphery so that
periphery forms broad denticle. Columella moderately arched forward,
New Mexican Land Snail 47
Fics. 1-6. Ceochasma phrixina new species. 1-3, Holotype, UF
20138; 4-6, Paratype, UF 20139.
forming a weak denticle at junction with basal lip; columellar margin of
aperture weakly concave. Umbilical callus thin, glossy, smooth, less than
\% shell diameter, with weak, elongate dent at base of columella. Whorls
5.7-6.3 (usually more than 6.0). Embryonic whorls not clearly demar-
cated from later whorls. First whorl protruding, large, diameter 1.3-1.5
mm. First 2.5 whorls with moderately coarse arched incremental stria-
tions and wrinkles; remaining whorls with similar close striations and
wrinkles and strong spiral chords that parallel suture and continue to lip;
last whorl with 8-11 chords above, 12-19 chords below periphery, with
chords below periphery flattened and broader near umbilical region so
that spiral sculpture appears more as strongly impressed spiral grooves
than as raised chords.
The shell is dull in texture. Two color phases are about equally
represented in the type series. The holotype and most of the fresh para-
types are reddish pink dorsally and slightly lighter below. The suture is
nearly white, and the periphery and lip are white. The umbilical callus
is glossy white, but is thin along its periphery where it shows the under-
lying color of the shell. The remaining paratypes are similarly colored
except that the ground color is lemon-yellow.
Holotype Paratypes
height 14.3 mm 12.2-14.6 mm
width 20.4 18.4—20.4
aperture width 11.1 9.8-11.1
whorls 6.3 5.7— 6.3
48 Proceedings of the Biological Society of Washington
iG g
Fics. 7-9. Operculum of Ceochasma phrixina new species. 7, inner
surface; 8, outer surface; 9, parietal-columellar margin, scale equals
1 mm.
Operculum (Figs. 7-9): Concentric; nucleus acentric, near columellar
margin and slightly below middle. Horny portion reddish-yellow, extend-
ing beyond margins of calcareous plate. Latter well developed, white,
with low oblique lamellarlike projection along columellar margin, and
relatively deep impression over nuclear region where plate is thinnest.
Columellar margin with shallow but sharp furrow along base of lamellar
projection.
Reproductive System: (Terminologies follow Baker, 1926a). Male
(Fig. 10) secondary gonoduct greatly enlarged, convoluted, with first
prostate about %3 length secondary gonoduct. Right compartment with
numerous small globular lobes along left margin. Lobes becoming
increasingly complex near apex of first prostate. Second prostate about
one half length first prostate and slightly more stocky. Third prostate
relatively long, poorly demarcated, with conspicuously thicker walls than
second prostate. Accessory prostate almost as long as combined length
of first and second prostate; nearly uniform in width, except along some
terminal lobes. Basal segment elongate. Caecum about as long as second
prostate, very thin-walled.
Female (Fig. 11) with right limb of V-organ cylindrical and stocky,
left limb constricted near both apex and base. Accessory sperm sac
small, elongate, with short duct entering near base of left limb of V-
organ. Pedicel stout, well differentiated from left limb. Reception
chamber large, demarcated from uterus by deep furrow. Ventral bursa
large, with numerous small elongate lobes radiating from central duct.
Provaginal sac elongate, furrowed and lobed along outer margin; con-
New Mexican Land Snail 49
, = —V-organ
—<—ventral
pedicel—..
first reception chamber-— purge
prostate ne ‘> provaginal
i sac
~— secondary
gonoduct
f—accessory prostate
caecum—*
second
prostate
eS thind
© are prostate
—————d
Fics. 10, 11. Genitalia of Ceochasma phrixina new species. 10, male
system; 11, female system.
nected to reception chamber by short stout duct. Basal segment of
uterus very deeply pigmented. Adjacent rectum and cloacal region light
gray colored.
Radula (Figs. 12-21): (Terminologies follow Baker, 1922) R-central
(Fig. 12) a simple shieldlike structure with narrow, irregular cutting
edge along upper margin, small projection along basal margin. A- and
B-centrals have heavy cusp-bearing backs. A-central (Fig. 13) shoe-
shaped, lacks cusps along cutting edge. Occasionally one or two small
denticlelike knobs present, irregular in appearance and location. B-central
(Fig. 14) irregularly bootshaped, with 2-4 large pointed cusps along
cutting margin. Cusps vary in size, relatively small in some specimens.
C-central (Fig. 15) simple, spatulate, with single large horny cusp.
Capituliform complex (Figs. 16-17) consists of a large, irregular, trape-
zoidal comb-lateral (Fig. 16) and a very long accessory plate (Fig. 17).
The comb-lateral has about 6 well-defined cusps along its cutting margin,
and occasional small cusplike lumps may also be present. The accessory
plate attaches to the comb-lateral by a short extension that fits under
the dorsal crest of the comb-lateral. The main body also has a thin
reflection across the top of the main body. The accessory plate also
has a long lateral extension that is joined to the main body of the tooth
by a sharp sigmoid fold. The marginal teeth (Figs. 18-21) consist of
a series of 73-79 teeth per row on each side of the ribbon. The teeth
are elongate, sickle-shaped and simple (Fig. 18). The first 11-14
marginals are unicuspid (Fig. 19). The next 7-12 marginals are bicuspid
50 Proceedings of the Biological Society of Washington
Fics. 12-21. Radular teeth of Ceochasma phrixina new species. 12,
R-central; 13, A-central; 14, B-central; 15, C-central; 16, comb-lateral;
17, accessory plate; Figs. 16 and 17 are combined in their natural rela-
tionships to form the capituliform complex; 18, 3rd marginal; 19, 10th
marginal; 20, 11th marginal; 21, 21st marginal. Scale equals 100 microns
for Figs. 12-18, and 200 microns for Figs. 19-20.
(Fig. 20). The following 6-8 teeth are tricuspid (Fig. 21), and the -
remaining teeth have 4 or more cusps. The outermost marginals may
have as many as 6 cusps. The cusps on all of the marginals are long
and digitiform with rounded tips. The addition of cusps on the teeth
occurs along the outer margin of the tooth.
Relationship: Baker (1956: 28-30) divided the Helicinidae into two
subfamilies, the Helicininae and the Proserpininae, containing five and
three tribes, respectively. This classification is based upon earlier studies
of the radula (Baker, 1922: 35-64; 1926a: 35-54; 1926b: 449-451;
1928: 46; Pilsbry, 1927: 62-63; Thiele, 1927: 89). In the primitive taxa
all of the paired central, lateral and marginal teeth have well developed
acuminate cusps. The accessory plate of the capituliform complex is
relatively simple in its attachment to the lateral tooth, and has a long
lateral extension. The major evolutionary trend in the family is modifica-
tion and _ specialization of the capituliform complex and simplification
of the central teeth. The marginal teeth undergo relatively slight change
(Baker, 1923: 119). Other anatomical modifications have relatively
slight phylogenetic importance (Baker, 1926a: 35-54; 1928: 25-46).
The Helicininae include those groups in which all of the marginal teeth
have sharp, acuminate cusps, the capituliform complex as a comb-lateral
tooth in which the shank is located along the outer margin, and the
accessory plate is highly variable. The Proserpininae consists of those
groups in which the inner marginals are unicuspid, the outer marginals
have only a few large rounded cusps, the capituliform complex has a
T-lateral in which the shank is mesially located, and the accessory plate
is reduced in size and has a much reduced lateral wing.
The tribes of the Helicininae are not relevant to further discussion
because they have no bearing on the relationships of Ceochasma. The
New Mexican Land Snail Bill
extant tribes of the Proserpininae are the Stoastomini, Vianini and Proser-
pinini. The Stoastomini retain large cusps on all of the paired central
and lateral teeth. The T-lateral apparently is secondarily modified so
that it resembles a comb-lateral (Baker, 1928: 46). The shank of the
lateral tooth extends to the outer margin, but apparently originates from
near the center of the tooth. The Vianini is characterized by having a
typical T-lateral and generally a reduction or absence of cusps on the
paired centrals and lateral. The Proserpinini has a radula that is hardly
distinguishable from that of the Vianini, merely representing a more
advanced stage in the functional reduction of the central teeth. However,
the tribe is distinct in lacking an operculum and in having well developed
spiral lamella in the aperture.
Ceochasma clearly belongs to the subfamily Proserpininae on the basis
of its unicuspid inner marginal teeth and the lobate nature of the cusps
on the outer marginals. Its relationship within the subfamily is less clear
since it shares characteristics with the Stoastomini and the Vianini.
Ceochasma is similar to the Stoastomini in the appearance of its lateral
tooth. It is similar to the Vianini in that it has already lost cusps on
the A- and C-centrals. Ceochasma is more primitive than any known
genus in either tribe. It has an enlarged accessory plate that still bears
a pronounced lateral winglike extension, and its lateral tooth has all of
the characteristic features of a comb-lateral. I consider Ceochasma to
be a primitive member of the Vianini in which cusp reduction has already
occurred on the central teeth, but which retains a primitive capituliform
complex. It probably lies near the stem of the origin of the Vianini and
the Proserpininae collectively. The Stoastomini probably diverged prior
to this point, but underwent more advanced modifications in its capitu-
liform complex. If this is so then the stoastomid lateral tooth is modified
comb-lateral, and not a secondarily modified T-lateral as Baker has sug-
gested (1928: 46).
Other aspects of the soft anatomy and the operculum are of limited
use in establishing phylogenetic relationships because of the slight and
inconsistent variations that occur in these characters throughout most of
the family. However, in its reproductive structures Ceochasma strongly
resembles Tristramia, which Baker (1922: 39, 48) considered the most
primitive subgenus of Helicina. The female reproductive systems of both
groups possess a highly lobate ventral bursa, much more so than occurs
in most other groups of the family (Baker, 1926a: 42-44; 1928: 32-33).
Additional similarities to Tristramia are also evident in the structure of
the operculum. The calcareous plate is not as weakly developed in
Ceochasma as in Tristramia, but both are alike in having the horny plate
extend beyond the margins of the underlying calcareous plate. They are
also alike in the nature and location of the concentric growth rings on
the horny plate. Both genera occur together in Colima. In contrast, the
Vianini usually does not have the horny plate as extensively developed,
and the concentric growth rings are more centrally located or consist
of some direct variation thereof.
At
52 Proceedings of the Biological Society of Washington
Although the characteristics of the reproductive system and the
operculum cannot be heavily weighed, they do tend to strengthen the
allocation of Ceochasma as a primitive member of the Vianini.
LITERATURE CITED
Baker, H. B. 1922. Notes on the radula of the Helicinidae. Proc.
Acad. Nat. Sci. Phila., 74: 29-67, pl. 3-7.
1923. Notes on the radula of the Neritidae. ibid., 75: 117-—
178; pls. 9-14.
1926a. Anatomical notes on American Helicinidae. ibid.,
78: 29-56; pls. 5-8.
1926b. The radula of Prosperpina. ibid., 78: 449-451.
1928. Mexican mollusks collected for Dr. Bryant Walker in
1926, I. Occ. Pap. Mus. Zool. Univ. Mich., (1932): 1-65.
—. 1956 Family names for land operculates. Naut., 70: 28-31.
Pitssry, H. A. 1927. Notes on the genus Ceratodiscus. Naut., 41:
62-63.
THIELE, J. 1927. Handbuch der Systematischen Weichtierkunde, I.
Berlin: i-vi, 1-778.
GE OC HS:
Vol. 81, pp. 53-68 April 30, 1968
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
TWO NEW WORMFISHES (GOBIOIDEA:
MICRODESMIDAE) FROM THE INDIAN OCEAN
By C. E. Dawson
Gulf Coast Research Laboratory,
Ocean Springs, Mississippi
Among fishes I have examined in connection with current
studies for a revision of the gobioid family Microdesmidae, are
two undescribed species from the Seychelles and Maldive
Islands. Since each of these species contributes significantly
to our knowledge of the characteristics, species diversity and
distribution of the Indo-Pacific genus Gunnellichthys Bleeker,
I have decided to publish their descriptions at this time.
Gunnellichthys is widely distributed and locally abundant in
tidepool and reef habitats from Mozambique eastward to Fiji
and, questionably, the Line Islands. Generic characters were
most recently discussed by Schultz (1966) and although his
observations are, at best, incomplete, I propose to delay generic
diagnosis and description until I have had access to additional
study material.
Specimens reported here were obtained during the Inter-
national Indian Ocean Expedition and have been deposited in
the collections of the Academy of Natural Sciences of Phil-
adelphia (ANSP), the Field Museum of Natural History
(FMNH) and the Gulf Coast Research Laboratory (GCRL).
Head length is measured from tip of lower jaw to base of
uppermost pectoral ray; body depth measured at anal fin
origin; caudal fin length as distance between posterior margin
of hypural vertebra and tip of longest caudal ray. Measure-
ments are in millimeters (mm); proportions are shown as
percentages of standard length (SL) or head length (HL).
Appreciation is expressed to Dr. James E. Bohlke and Loren
8—Proc. Bion. Soc. WasuH., Vou. 81, 1968 (53)
54 Proceedings of the Biological Society of Washington
Fic. 1. Gunnellichthys curiosus ANSP 103611; holotype; 79.4 mm SL.
P. Woods for making this material available for study.
Acknowledgment is also made to Drs. Reeve M. Bailey and
C. Richard Robins for their critical comments on the manu-
script. Semidiagrammatic delineations are by Harry L. Moore,
Jr.
This study was supported in part by the American Philo-
sophical Society Penrose Fund Grant No. 4242.
Gunnellichthys curiosus new species
(Fig. 1)
Holotype: ANSP 103611; 79.4 mm SL; Curieuse Island, Seychelles,
just SE of point forming south end of Laraie Bay (Admiralty Chart no.
1072); from a depth of about 30 ft over sand and coral bottom; 23
February 1964; Sta. F-64, International Indian Ocean Expedition, Sey-
chelles Islands Program; James E. Bohlke et al. coll. This is the only
known specimen.
Diagnosis: A species of Gunnellichthys with short pelvic fins; lower
jaw with distinct, horizontally broad, lip; snout and lower jaw not
conspicuously narrowed at symphysis; close set to narrowly imbricate
New Indian Ocean Wormfishes 55
scales over most of body and head; snout and lower jaw clothed with
microscopic villi; three anteriormost dorsal spines more closely spaced
than successors; with several, prominent, long subvertical rows of papillae
on sides between anal fin origin and pectoral fin insertion; opercle with
dark spot; dark blotch on caudal fin base extending onto proximal caudal
fin; anterior dorsal fin with narrow brown to dusky marginal stripe, a
second stripe, or series of interradial blotches, below; body with a pale
midlateral longitudinal stripe; proximal pterygiophores of 1st two dorsal
spines inserted over interspace between 2nd and 3rd neural spines;
prezygapophyses low, not distinctly elevated.
Description: Dorsal spines 20, dorsal segmented rays 42, total dorsal
elements 62; anal rays 39; pectoral rays 15; pelvic fin I, 4; principal
caudal rays 15; vertebrae 26 + 32 = 58.
Measurements (mm) are followed by percent SL or HL in parentheses.
Caudal fin length 7.8 (9.8); least depth of caudal peduncle 2.7 (3.4);
body depth at anal fin origin 5.7 (7.2); predorsal length to tip of lower
jaw 13.2 (16.6); preanal length 41.6 (52.4); pectoral fin length 5.3
(6.7); pelvic fin length 2.6 (3.3); distance from pelvic insertion to anal
fin origin 29.1 (36.6); head length 12.1 (15.2). Diameter of fleshy
orbit 2.0 (16.5); distance from anterior margin of eye to tip of lower
jaw 4.0 (33.1); snout length 2.0 (16.5); postorbital length 6.1 (50.4);
tip of lower jaw to angle of gape 4.3 (35.5).
Body moderately elongate, tapering to slightly less than half body
depth at caudal peduncle; compressed, greatest breadth (7.6 percent
SL) at opercle, breadth at anal fin origin 3.4 percent SL; caudal fin
long, obtusely angular, middle rays longest; head depth subequal to
body depth; interorbital broad, about 7% eye diameter, laterally and
longitudinally convex; head curves evenly to snout tip without distinct
postorbital depression; eye prominent, lateral, diameter subequal to snout
length; lower jaw prominent, extends about eye diameter in advance of
snout tip; jaws and snout broadly rounded, not conspicuously narrowed
at symphysis; gape moderate, extends slightly past anterior margin of
eye, not strongly inclined; lips prominent, moderately thick and fleshy,
upper extends forward to below vertical from anterior naris, narrowed
and concealed in front by overhanging snout; lower jaw with horizontally
flattened lip crossing symphysis and expanded laterally into moderately
broad folds (this is a prominent and distinctive feature in both dorsal
and lateral aspect); anterior naris opens anterolaterally through short,
delicately flared, tubule located dorsoanterolaterally on snout margin;
posterior naris dorsolateral over preorbital and about % snout length
in advance of anterior margin of orbit; naris margin slightly elevated but
not distinctly tubiform; nares subequal in diameter, about 12 in eye;
jaws with outer series of large, separated, caniniform teeth in front, with
smaller close-set or contiguous, pointed teeth extending laterally to near
posterior angle of gape, apparently with two or more inner series of
close-set, pointed teeth in front; (dentition could not be clearly seen
without damage to specimen; inner margin of lower lip plicate, but
56 Proceedings of the Biological Society of Washington
os terse “ga Coie 5
Assigns (oe en eg
fer nice Gi) eG:
Fic. 2. Semidiagrammatic delineations of gill openings and pelvic
fins. Anterior dorsal fin elements omitted. Upper: Gunnellichthys
curiosus. Middle: Gunnellichthys viridescens. Lower: Gunnellichthys
curiosus.
New Indian Ocean Wormfishes 57
presence or absence of labial teeth is not yet confirmed); tongue thick,
round in front.
Gill opening (Fig. 2) moderate, its diagonal about half again as long
as pectoral fin base, slitlike, not tubiform, originating just below upper
angle of pectoral fin base and slightly in advance of insertion of upper-
most pectoral ray, margin continuing almost vertically downward across
pectoral base then anteriad as free flap of branchiostegal membrane to
unite ventrolaterally with isthmus in advance of pelvic fin insertion,
distance from lower pectoral angle to ventral terminus of gill opening
equivalent to 24 length of pectoral fin base; pectoral peduncle largely
concealed beneath opercle and gill membranes; median fins terminate
on anterior caudal peduncle, completely free from caudal fin, depressed
tips of posterior rays fail to reach posterior margin of hypural, extend
to 2nd or 3rd spinniform caudal fin element; dorsal fin origin above
anterior third of pectoral fin, little more than 4% eye diameter behind
vertical from upper angle of gill opening, first two spines about 30
percent shorter than successors; interspace between first 2 dorsal spines
narrow, interspace between 2nd and 3rd somewhat greater, but less
than that between 3rd and 4th, which is subequal to remaining inter-
spaces (ratios of first three interspaces approximate 1:1.5:2), 11 anterior
rays simple, subsequent rays branched; Ist anal fin element segmented
and finely branched at tip, remaining rays, except last two, well
branched; last two dorsal and anal fin rays simple, more closely spaced
than those preceding, bases separate; caudal fin formula 7 + 2+ 11 +
2+7 (counts of procurrent elements from radiograph); pectoral fins
rounded, left 2+ 12 + 1, right 2+ 11 + 2, 6th-7th rays longest; pelvic
fins (Fig. 2) inserted distinctly in advance of pectoral fins, separate,
innermost ray simple, shorter than next, which is both branched and
longest, 3rd and 4th rays progressively shorter, branched, outer spine
shorter and slender; anal fin origin beneath interspace between 25th and
26th dorsal fin elements.
Head covered with close-set to locally imbricate scales from nape
forward to just anteriad of posterior nares; preorbital, suborbital, entire
cheek and opercle similarly scaled to ventral margins; scales narrowly
impinge periphery of eye; narrow band of scales on mandible below
lateral expansion of lower lip, lower jaw apparently naked elsewhere;
non-imbricate scales close-set on posterior % of branchiostegal mem-
brane, anterior membrane and remainder of ventral head naked. Body
squamation similar, maximum scale diameter about 0.25 mm, everywhere
close-set to locally imbricate, not widely separated; squamation complete
over predorsal, chest, abdomen and lateral body except in immediate
vicinity of anal and genital openings; scales on pelvic fin base; sub-
triangular scale patch extends over proximal half of caudal fin; scales on
pectoral fin base extend well into gill opening. Snout, upper preorbital,
lower jaw and much of anterior ventral portion of head densely covered
with microscopic hyaline villi; under low magnification these regions
appear to be pubescent.
58 Proceedings of the Biological Society of Washington
Fic. 3. Semidiagrammatic delineation. Left—Anterior abdominal
vertebrae showing dorsal spines and proximal pterygiophores. Right—
Last abdominal and anterior caudal vertebrae together with the last
dorsal spine, anterior segmented rays, proximal and distal pterygiophores.
Upper: Gunnellichthys curiosus. Lower: Gunnellichthys viridescens.
A series of papillae crosses nape, another crosses interorbital to unite
with a series that follows dorsal orbital margin and runs forward to snout
tip, this latter series is paralleled by a short series near snout midline
and by another longitudinal series along outer margins of nares; a series
runs from anterior naris ventrad across preorbital and closely follows
anteroventral orbital margin; one or two vertical series on cheek; a
vertical series extends from near upper opercular angle to near predorsal
midline; two parallel series originate below angle of gape, pass antero-
laterally along lateral midline of lower jaw and across symphysis; a
ventrolateral series from near lower pectoral angle to just posteriad of
pelvic fin insertion; 8 or 9 long subvertical rows on lateral body between
pectoral axil and anal fin origin, originating on upper half of body and
extending to near ventral midline; sensory papillae indistinct or obsolete
posteriad of anal fin origin.
No predorsal interneurals. Proximal pterygiophores of first two dorsal
spines (Fig. 3) somewhat flattened, subhorizontal, without distinct
descending process, inserted over interspace between 2nd and 3rd neural
spines; proximal pterygiophore of 3rd dorsal spine with indistinct (in
radiograph) poorly ossified descending process inserted between 3rd
and 4th neural spines; subsequent proximal pterygiophores better ossified,
progressively more spinelike and each inserted between succeeding neural
spines. Distal pterygiophores begin immediately behind Ist segmented
dorsal ray and after 2nd anal ray; no supplemental proximal pterygio-
New Indian Ocean Wormfishes 59
Fic. 4. Radiograph of anterior body showing head, axial skeleton
and abdominal ribs. Upper: Gumnnellichthys curiosus. ANSP 103611;
holotype. Lower: Gunnellichthys viridescens. ANSP 103624; holotype.
phore between last dorsal spine and Ist ray. Abdominal neural spines
strong, slender, sharply pointed, of subequal length, each located pos-
teriad on centrum; prezygapophyses short, depressed in front, not dorsally
elevated; ribs apparently articulate with centra of first three vertebrae,
with parapophyses on remaining abdominal vertebrae; anterior ribs short,
those of 4th vertebrae about 4% longer than centrum, lengthening pos-
teriad until about thrice vertebral length at 19th or 20th vertebra; ribs
mostly at attitude of 45°; abdominal epipleurals distinct, subequal to
vertebral length or slightly shorter, caudal epipleurals not visible in
radiograph; skull (Fig. 4) elongate, not strongly elevated in front.
Frontals well ossified, reaching mesethmoid. Premaxillary pedicel strong,
blunt, not abruptly angled posteriad at tip. Posttemporals distinctly
elevated above anterior vertebrae. Pectoral radials 4. Branchiostegal
rays 5, innermost remote from next. Ceratohyal broadened posteriorly.
Dentary strong, with short, pointed terminal ventral process.
Ground color in alcohol, light tan; eye with black iris and gray-black
pupil; dark brown blotch, about %% eye diameter, on upper margin of
60 Proceedings of the Biological Society of Washington
opercle and lateral head (Fig. 1) its center about even with upper
pectoral angle; larger elongate lateral blotch on posterior caudal pe-
duncle and proximal half of caudal fin, ventrally expanded to near base
of lowermost segmented ray, terminating dorsally at 4th branched caudal
ray; dorsal fin with a narrow brown stripe extending posteriad to about
23rd ray, edging fin anteriorly, it is surmounted by narrow hyaline band
behind; first 7 dorsal spines and interradial membranes dusky brown,
subsequent dorsal elements lighter but at least partly delineated by
minute brown melanophores; each interradial membrane from 7th spine
to about 20th ray with conspicuous longitudinally elongate blotch, wider
than marginal stripe and separated above and below by immaculate
membrane; although interrupted by fin supports, blotches appear as
median dorsal fin stripe; anterior 25 or 26 anal rays delineated by fine
speckling of minute brown melanophores, remaining fin immaculate;
body and head shaded with greater or lesser concentrations of minute
brown melanophores; predorsal to snout and lateral head to about middle
of opercle dusky brown, melanophores less dense below, absent along
ventral third of opercle and cheek; lips and lower jaw finely speckled;
ventral head and branchiostegal membranes generally immaculate; few
scattered melanophores on chest; pelvic fins closely speckled proximally,
melanophores scattered toward fin tips; pectoral fin base with rather
dense patches of light brown melanophores above and below separated
by a narrow immaculate stripe; ventral patch continues into gill opening,
remainder of fin immaculate; lateral body from dorsal fin base to midline
dusky, somewhat lighter than head, finely speckled from dorsal fin origin
to caudal fin base; a generally immaculate stripe, bounded dorsally by
midline, extends from pectoral axil to caudal fin base, stripe broad under
pectoral fin, narrower along middle body, expanded over entire lower
half of body posterior to 26th anal ray, scattered melanophores intrude
stripe between pectoral tip and anal fin origin; remainder of lower sides
and the ventral abdomen dusky; scales on upper head, upper body and
medially on ventral abdomen are posteriorly margined with brown, scales
elsewhere usually without margination.
Etymology: The specific name refers to the curious body coloration
and the unusual pubescence of the anterior head.
Relationships: This species may be distinguished from Gunnellichthys
pleurotaenia Bleeker and G. copleyi J. L. B. Smith by its close-set to
imbricate squamation, its thick and flat, rather than thin and undif-
ferentiated lower lip, its obtusely angular, rather than rounded or trun-
cate, caudal fin, striking differences in coloration, including its lack of
a continuous dark lateral stripe and by differences in proportions, counts
and skeletal characters. G. curiosus agrees with these species in having
short pelvic fins, a broad anterior head configuration and a_ similar
spacing of the anterior dorsal spines. It is readily separated from G.
irideus J. L. B. Smith by its distinctive coloration, the rounded, rather
than narrow, snout and lower jaw, squamation and short pelvic fins.
Coloration of curiosus is superficially similar to that of G. monostigma
t=
New Indian Ocean Wormfishes 61
Fic. 5. Gunnellichthys viridescens. ANSP 103624; holotype; 53.6
mm SL.
J. L. B. Smith but the latter lacks the striped dorsal fin, the immaculate
lateral stripe and the large caudal blotch. Anterior dorsal spines are
regularly spaced in monostigma, the head is conspicuously narrowed in
front, scales are separated and there are differences in counts, propor-
tions and skeletal characters. Gunnellichthys curiosus appears to be
most closely related to G. pleurotaenia and G. copleyji.
Gunnellichthys viridescens new species
(Fig. 5)
Holotype: ANSP 103624; 53.6 mm SL; Seychelles, W. of NW tip of
Anonyme Island, between Anonyme and Mahé islands (Admiralty Chart
no. 1072); at depth of 35-50 ft; 11 February 1964; Sta. F-44, Inter-
national Indian Ocean Expedition, Seychelles Islands Program; James
E. Bohlke et al. coll.
Paratypes: ANSP 103625; (7) 35-53 mm SL; other data as for holo-
type. ANSP 109202; 50 mm SL; cleared and stained; other data as for
62 Proceedings of the Biological Society of Washington
TaBLe 1. Measurements of Gunnellichthys viridescens.+
Holotype Paratypes
mm % N Range (mm) X%
Standard length 53.6 14 29.8-64.4
Caudal fin length 4.9 9.1 12 3.2— 6.2 9.4
Least caudal peduncle depth 1.6 2.9 12 0.9— 2.0 2.9
Body depth at anal fin origin 3.0 5.6 12 1.8— 3.7 5)
Predorsal length (to tip of lower jaw ) 8.7 16.2 12 5.4-10.4 16.6
Preanal length 27.3 50.9 12 15.6—32.6 51.0
Pectoral fin length 3.6 6.7 119) 2.1-— 4.3 6.6
Pelvic fin length 1.4 2.6 12 1.0— 2.2 3.0
Pelvic insertion to anal fin origin 19.9 37.1 12 10.6—23.0 36.6
Head length 7.4 13.8 11 4.8— 9.7 14.8
Diameter of fleshy orbit 1.4 18.91 11 1.1— 1.6 19.3
Anterior margin of eye
to tip of lower jaw 2.0 27.0 10 1.5— 2.7 26.9
Snout length 1.4 18.9 8 0.9— 1.7 17.8
Postorbital length 4.0 54.0 10 3.2— 5.4 54.1
Fleshy interorbital width 0.8 10.8 10 0.5-— 1.1 9.6
Tip of lower jaw to angle of gape 2.6 35.1 10 1.7— 3.5 32.6
1From caudal fin length through head length percentages are of standard length;
remaining percentages are of head length.
holotype. GCRL V67:2186; 53 mm SL; other data as for holotype.
ANSP 103609; 51 mm SL; Mahé Island, Seychelles, SW of Souris
Island; coral sand and boulder bottom at depth of about 7 ft; 31 January
1964; Sta. F-13, International Indian Ocean Expedition, Seychelles
Islands Program. ANSP 103612; (3) 30-40 mm SL; Mahé Island, Beau
Vallon Bay, NNW of Hotel des Seychelles; from isolated patch of rock,
coral and sponge at a depth of 40-50 ft; 19 March 1964; Sta. F-119,
International Indian Ocean Expedition, Seychelles Islands Program.
FMNH 73909; 64.4 mm SL; Bushy Island, Addu Atoll, Maldive Islands;
from coral and sand bottom at a depth of 2-8 ft; 10 May 1964; Inter-
national Indian Ocean Expedition; L. P. Woods and D. L. Ray coll.
Diagnosis: A species of Gunnellichthys with short pelvic fins; lower
jaw with distinct horizontal expansion across symphysis; snout and lower
jaw not conspicuously narrowed in front; scales separate, non-imbricate;
snout, lower jaw and much of ventral head with blunt, microscopic,
papillae; four anteriormost dorsal spines more closely spaced than suc-
cessors; without prominent subvertical rows of lateral abdominal sensory
papillae; without conspicuous color markings but with faint, dark, median
stripe on anterior dorsal fin, pale lateral stripe, becoming slightly
darkened anteriorly with increase in size, margined dorsally by fine
dark line and an indistinct dusky median dorsal shading from snout
posteriad to caudal peduncle along each side of dorsal fin base; proximal
pterygiophores of first two dorsal spines inserted over interspace between
2nd and 3rd neural spines; anterior caudal prezygopophyses, sub-
triangular, elevated.
—————— se C~S
New Indian Ocean Wormfishes 63
TaBLeE. 2 Counts of Gunnellichthys viridescens.
Paratypes
Holotype N Range
Number of dorsal spines o1 15 20-21
Number of segmented dorsal rays 381 15 38-41
Total dorsal elements 59 15 59-61
Number of anal rays 37 SeelS 36-40
Anal fin origin beneath interspace
between dorsal elements 94/25 14 94/25-25/26
1 The last two rays of dorsal and anal fins are counted separately.
Description: Dorsal spines 20-21, dorsal segmented rays 38-41, total
dorsal elements 59-61; anal rays 36-40, pectoral rays 12-13; pelvic fin I,
4; principal caudal rays 15; vertebrae 24-25 + 33-34 = 57-59. See
Tables 1 and 2 for proportional measurements and counts.
Body moderately elongate, slender, depth at anal fin origin averages
5.4 percent SL, tapering to about 3 percent SL at caudal peduncle;
compressed, breadth at anal fin origin 2.6-3.7 percent SL, greatest
breadth (5.0 percent SL in holotype) at opercle; caudal fin rounded,
averages 9.4 percent SL; head 13.8-16.2 percent SL, its depth subequal
to body depth; interorbital moderately convex, width about 1% eye
diameter, with prominent, lateral, fleshy supraorbital ridges; head curves
evenly to snout tip without distinct postorbital depression; eye lateral,
diameter, averaging 19.3 percent head length, equal to or somewhat
greater than snout length; snout and lower jaw rounded, not conspicu-
ously narrowed at symphysis; gape moderate, extends (in holotype)
just posteriad of vertical from anterior margin of pupil, not strongly
inclined; lips not prominent, not conspicuously enlarged or fleshy; upper
lip generally narrow throughout, laterally exposed to near vertical from
anterior naris where lip is further narrowed and concealed across
symphysis by overhanging snout; lower lip somewhat horizontally
flattened across symphysis, undifferentiated anterolaterally and across
symphysis, with little posterolateral expansion; anterior naris minute,
opens anterolaterally through short simple tubule on dorsoanterolateral
snout margin; posterior naris dorsolateral over preorbital about 14 snout
length in advance of anterior orbital margin, with short simple tubule;
diameter of anterior naris 4% to 4% diameter of posterior naris which
is about 14 in eye diameter. Lower jaw with outer row of 3 or 4
enlarged, bluntly pointed, slightly recurved, separated caniniform teeth
on each side of symphysis (anteriormost smallest) followed by 12-15
smaller, more closely spaced, conical teeth, which decrease in size
posteriad; jaw anteriorly with 2 irregular inner series of smaller, slightly
recurved, pointed teeth, middle row becoming laterally obsolete; posterior
°4 of dentary with biserial, or terminally, uniserial dentition; upper jaw
teeth similar to lower jaw teeth, anterior caniniformes somewhat larger,
slightly protruding; inner margin of lower lip plicate, apparently with
64 Proceedings of the Biological Society of Washington
5 minute triangular teeth, one at symphysis between anteriormost canini-
form teeth, two on each side between bases of Ist three or four enlarged
teeth (labial teeth visible under 60 < magnification in holotype and
most paratypes, presence confirmed in cleared specimen); tongue thick
and rounded in front.
Gill opening (Fig. 2) moderate, its diagonal about 4% longer than
pectoral fin base, slitlike, not tubiform or greatly restricted; origin
slightly in advance of upper axil of pectoral peduncle, on level with
insertion of upper pectoral ray; margin extends almost vertically down-
ward to a point opposite 8th or 9th pectoral ray, then angles gradually
downward and forward, as free fold of branchiostegal membrane, unit-
ing ventrolaterally with isthmus in advance of pelvic fin insertion;
distance from lower pectoral axil to anteroventral terminus of gill open-
ing equivalent to 4% length of pectoral fin base; lower angle of gill
opening concealed by overhanging fold of branchiostegal membrane;
margin of membrane slightly produced at each branchiostegal ray;
peduncle of pectoral fin largely exposed, not concealed by opercle and
gill membranes. Dorsal and anal fins terminate on anterior caudal
peduncle, completely free from caudal fin; depressed tips of posterior
rays fail to reach posterior margin of hypural, extend to 3rd or 4th
spinniform caudal fin element; dorsal fin origin above anterior 1% of
pectoral fin or about 44 cye diameter posteriad of vertical from upper
angle of gill opening; first dorsal spine shortest, subsequent spines
gradually increase in length until 5th, which is about twice as long as
lst, remaining spines of subequal length; Ist dorsal ray about 14 longer
than last dorsal spine; remaining rays gradually increase in length to
about 35th, which is 4% longer than Ist; subsequent rays gradually
decrease in length; interspace between first 2 dorsal spines narrow, next
two interspaces increasingly longer, but shorter than 4th, which is sub-
equal to remaining interspaces (anterior interspace ratios vary somewhat,
those of two paratypes and holotype being 1:1.3: 1.7: 2.0, 1:1.2:
1.3: 1.6 and 1: 1.6: 2.0: 2.2); pelvic fins short, inserted in advance of
pectoral fin base near vertical from lower axil of pectoral peduncle,
separate, innermost pelvic ray shorter than next, which is the longest,
3rd and 4th rays progressively shorter, outer spine short and _ slender;
longest ray of right pelvic fin branched in holotype, 3 outermost rays
branched in FMNH 73909, all other pelvic fin rays in type series simple;
fin configuration slender but generally similar to that shown in Fig. 2;
anal fin origin beneath interspaces between 24th to 26th dorsal fin
elements.
Scales inconspicuous, somewhat embedded, well separated, seldom
touching and nowhere imbricate; maximum scale diameter about 0.2
mm; head with scattered scales on dorsum from nape forward to posterior
orbital margin, interorbital apparently naked except for occasional
isolated scales on orbital margin; snout and upper preorbital naked;
narrow series of scales on lower preorbital and suborbital from midline
of eye to posterior angle of gape, remaining suborbital, cheek and opercle
New Indian Ocean Wormfishes 65
generally scaled to ventral margins; scattered scales on gill membranes,
remainder of ventral head apparently naked. Body squamation generally
complete; naked area about anal and genital openings; chest scaled near
pelvic fin base but apparently naked elsewhere; caudal peduncle scaled
above and below, caudal fin apparently naked; scattered scales on
pectoral fin base continue into gill opening. Snout, upper preorbital,
lower jaw and much of ventral head with short, blunt, microscopic
papillae, without hyaline villi.
Series of larger dermal papillae inconspicuous, difficult to see; a
circumorbital series passes closely about eye with anterodorsal branch
extending forward to anterior naris; a series follows anterior preorbital
margin, terminating near angle of gape; a ventrolateral series along lower
jaw from tip to near posterior angle of cheek; a predorsal series continues
ventrad on each side to near upper pectoral axils; a few short, longi-
tudinal and subvertical series of 4 to 8 papillae on lateral head above
opercle; lateral body with short subvertical series of 3 to 6 widely
spaced papillae on myomeric impressions, other long or distinctive series
apparently lacking.
No predorsal interneurals. Proximal pterygiophores of first 4 dorsal
spines (Fig. 3) flattened, rectangular in lateral aspect, subhorizontal;
4th slightly decurved, 5th and subsequent abdominal proximal dorsal
pterygiophores progressively more spinelike with descending processes
interdigitating with succeeding neural spines; proximal pterygiophore
of Ist dorsal spine inserted slightly posteriad of 3rd neural spine, next
pterygiophore over interspace between 3rd and 4th neural spines, 3rd
and 4th proximal pterygiophores inserted over 4th and 5th neural spines;
distal pterygiophores begin immediately following lst segmented dorsal
ray and after Ist anal ray; no supplementary proximal pterygiophore
between last dorsal spine and Ist segmented dorsal ray. Abdominal
neural spines strong, slender, sharply pointed, of subequal length and
located posteriad on centrum; abdominal prezygopophyses short, little
elevated in front; anterior caudal prezygopophyses longer, elevated, sub-
triangular projections becoming reduced and subhorizontal posteriad of
26th abdominal vertebra; ribs apparently articulate with centra of Ist
three vertebrae, with parapophyses on remaining abdominals; anterior
ribs about 4% longer than centrum, attain maximum of about twice
vertebral length at 15th or 16th vertebra; remaining abdominal ribs of
subequal length except for last, which is about 4% longer than vertebra;
ribs mostly at an attitude of 45°; abdominal epipleurals distinct, slightly
longer than vertebra; epicentrals, subequal to vertebral length, on caudal
vertebrae; skull (Fig. 4) moderately elongate, not strongly elevated in
front. Frontals well ossified, reaching mesethmoid. Premaxillary pedicel
strong, blunt; tip spatulate in lateral aspect, not abruptly angled pos-
teriad. Posttemporals distinctly elevated above anterior vertebrae. Pec-
toral radials 4. Branchiostegal rays 5, innermost remote from the next.
Ceratohyal broadened posteriorly. Dentary strong, with short, pointed,
66 Proceedings of the Biological Society of Washington
well-ossified terminal, ventral process (foregoing observations from
cleared and stained paratype and radiographs ).
Ground color of holotype in alcohol, yellow-green; without conspicu-
ous markings; eye black with iridescent gray-black pupil. Other color
patterns are clearly distinguishable only under magnification. Pale
longitudinal stripe, slightly narrower than pupil diameter, originates
near posterior orbital margin, interrupted across opercle, continuing
posteriad to caudal fin base; stripe formed by local reduction of light
brownish micromelanophores, which, elsewhere, cover body and head
in varying concentrations; stripe margined dorsally by distinctive line
of slightly enlarged, darker brown, round melanophores predominately
arranged in single longitudinal series; micromelanophores usually sep-
arated, with closed centers anteriorly, becoming close-set or touching,
most frequently with open centers posteriad of anal fin origin. Predorsal
area dusky with concentration of brownish melanophores, which con-
tinues posteriad to caudal peduncle as a narrow dusky basal stripe on
both sides of dorsal fin; remainder of body sides and ventral abdomen
slightly speckled with pale-brownish melanophores; barlike concentration
of melanophores crosses suborbital and cheek broadens posteriad to give
entire opercle faintly dusky aspect; similar concentration begins on .
anterior 4% of upper lip, continues forward to symphysis of lower jaw;
scattered, darker brown micromelanophores present on peduncle of
pectoral fin; caudal, anal and paired fins immaculate; interradial mem-
branes of anterior dorsal fin with median, longitudinal, concentrations
of brownish melanophores, which, together, form a faint stripe; mem-
brane, above and below, as well as entire dorsal fin posteriad of vertical
from the 3rd or 4th anal fin ray, immaculate. Largest specimen (FMNH
73909) differs in having pale tan melanophores concentrated to form an
indistinct stripe, narrower than pupil diameter, from tip of lower jaw
to upper pectoral angle; similar melanophores frequently intrude pale
lateral stripe on remainder of body; dark line of melanophores on dorsal
margin of lateral stripe obsolete on anterior 7% of body. Color of
remaining paratypes similar to that of holotype.
Etymology: The specific name refers to the generally greenish appear-
ance of the type material.
Relationships: The absence of conspicuous color markings separates
Gunnellichthys viridescens from all known congeners except G. irideus
J. L. B. Smith from which it may be distinguished by having a rounded,
rather than conspicuously narrowed, snout and lower jaw; a thickened
lower lip and deeper lower jaw configuration; short, rather than long
pelvic fins; modally 59, rather than 55, vertebrae, and other features.
The configuration of the lower lip, irregular spacing of the anterior
dorsal spines and short pelvic fins suggest that viridescens is most closely
related to G. pleurotaenia Bleeker, G. copleyi J. L. B. Smith and G.
curtiosus.
Remarks: The first three or four segmented dorsal rays, as well as the
first anal element, are simple in the largest paratype. The remaining
ales
a soo Sy
ee ee
=r a ye
New Indian Ocean Wormfishes 67
specimens show a general positive correlation between increased SL and
the branching of dorsal and anal segmented fin rays. Incomplete branch-
ing of segmented dorsal and anal fin elements, together with a pre-
dominance of simple, rather than branched, pelvic fin rays, suggests
that none of the type series are fully developed. Larger specimens
probably have four branched pelvic fin rays and all segmented dorsal
and anal elements branched. Branching of fin rays is an ontogenetic
character in microdesmids and, in this species, anal fin elements begin
to branch prior to those of the dorsal fin; branching develops first in
the posterior fin, then proceeds anteriorly with growth. Adult color
patterns develop early in microdesmids (at 20-25 mm SL in some
species ) and apparently change little with increased size. It is improbable
that the color pattern of G. viridescens undergoes striking changes in
later life.
The presence of labial teeth has not been previously described in
microdesmid fishes; probably because of the paucity of study material
of many genera and species. Preliminary observations suggest that labial
teeth do not occur in the Eastern Atlantic and Western Hemisphere
species assemblage presently included in the nominal genus Microdesmus
Giinther.
The occurrence of Gunnellichthys viridescens in the Maldive Islands
constitutes the first record of the family Microdesmidae in the Eastern
Indian Ocean. Discovery of this species in both the Seychelles and
Maldives suggests that it may be a common component of the Indian
Ocean insular ichthyofauna.
LITERATURE CITED
ScHuLtTz, L. P. 1966. Fishes of the Marshall and Marianas Islands.
Bull. U.S. Nat. Mus. 202(3): 1-13.
68 Proceedings of the Biological Society of Washington
GIG 7s. Lae
Vol. 81, pp. 69-86 \ |} April 30, 1968
Pear rr
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
FROGS OF THE GENUS PLATYMANTIS (RANIDAE)
FROM NEW BRITAIN WITH DESCRIPTIONS
OF NEW SPECIES
By WALTER C. BROWN AND MICHAEL J. TYLER
Division of Systemic Biology, Stanford University and
Menlo College, Menlo Park, California; and
South Australian Museum, Adelaide, Australia
Zweifel (1960) commenting on the amphibian fauna of New
Britain as compared with that of New Guinea and the Solomon
Islands, conjectured that the Bismarck Archipelago had a more
recent origin in geologic time and consequently has a more
impoverished amphibian fauna, the result of overwater dis-
persal from both New Guinea and the Solomons. He also
noted (p. 1) that the frog fauna was probably poorly known.
Recent explorations, by one of us (Tyler), on New Britain
have added the family Microhylidae (Tyler, 1964) and
several new species of frogs to the island’s fauna. Included
are three new species and a new subspecies, described below,
belonging to the ranid genus Platymantis Giinther (Platyman-
tis replaces the name Cornufer Tschudi for the genus, Zweifel,
1967). These increase the number of species of Platymantis
known from New Britain to six, double the number known to
Zweifel (1960) and Gorham (1965) but still less than the nine
presently recognized from Bougainville Island in the Solomons
(Brown, 1965). With the exception of P. papuensis Meyer, as
treated in the present study, the species of New Britain are
distinct from those of the Solomons, though their closest
affinities, with the exception of P. boulengeri (Boettger) and
P. rhipiphalcus, new species, appear to be with Solomons’
species. This suggests a relatively long isolation of this seg-
ment of the fauna from related species in the Solomons and
New Guinea, but less long from the Solomons.
9—Proc. Biot. Soc. WasH., Vou. 81, 1968 (69)
70 Proceedings of the Biological Society of Washington
Measurements determined, to the nearest 0.1 mm, are based
on preserved specimens. Snout-vent length was measured
from tip of snout to vent, with the animal held flat, ventral side
down; head length from tip of snout to posterior edge of
tympanum; head breadth at angle of jaws; tibia length as
length of lower leg bone; eye diameter from anterior to pos-
terior edge of socket; third finger length from tip to base of
second subarticular tubercle; first finger length from tip to
base of subarticular tubercle. When ratios are stated in the
descriptions, the data are presented in the following manner:
head breadth/snout-vent length = average (number of speci-
mens: range). The most useful diagnostic characters are the
basis of the key that follows.
Key to Species of Platymantis from New Britain
1. Finger disks, except for Ist finger, considerably broader than
penultimate phalanx, about as broad or broader than toe disks;
4th finger, when adpressed, longer than either Ist or 2nd _______. 2
1’. Finger disks not or searcely broader than penultimate phalanx,
usually narrower than toe disks; 4th finger, when fingers are
adpressed, shorter than Ist or 2nd finger, or all three about equal
dn Verioth 22 = le ae a eee 3
2. Sides of head rather uniformly blackish; dorsum marked by
several very long, narrow folds; breadth of 3rd finger disk usually
more than 25 per cent of 3rd finger length; breadth of 3rd finger
disk greater than breadth of 3rd toe disk; head breadth usually
greater than 75 per cent of tibia length === rhipiphalcus
2’. Sides of head not uniformly blackish; dorsum relatively smooth
or marked by short folds; breadth of 3rd finger disk usually less
than 25 per cent of 8rd finger length; breadth of 3rd finger
disk usually slightly less than breadth of 3rd toe disk; head
breadth usually less than 75 per cent of tibia length ____ papuensis
3. Snout-vent length of mature females greater than 55 mm; head
breadth about equal to tibia length (except for juveniles); 3rd
finger length rarely as great as 30 per cent of head breadth;
diameter of eye 70 to 90 per cent of snout length, rarely more
than®s5*per fcent: 222 hal RR SAB EL Wished Se SAB boulengeri
3’. Snout-vent length of mature females rarely more than 50 mm;
head breadth usually less than 85 per cent of tibia length; 3rd
finger length 30 to 40 per cent of head breadth; diameter of eye
about equal to snout length, rarely less than 90 per cent _________ 4
4. Snout-vent length of mature males 18 to 25 mm; Ist finger about
as long as adpressed 2nd; Ist finger length 60 to 70 per cent of
Sraicfinger length matet ek aire Beta ee ee ae akarithymus
Frogs of the Genus Platymantis 71
4’. Snout-vent length of mature males more than 25 mm; Ist finger
usually longer than adpressed 2nd; Ist finger length 75 to 90
Percent Ohara mangers ere t hye yc a NA eae eT 5
5. Dorsum with numerous folds of short to moderate length; urm-
shaped, axillary pair of folds not present (Fig. 2a); eye diameter
usually greater than 35 per cent of head breadth; first finger
length usually less than 85 per cent of third finger length; pos-
terior surface of lower leg lacking conspicuous, blackish, longi-
urclira eal obyetra cli sereeeame sis onset iL a as AU mimicus
5’. Dorsum relatively smooth except for urnshaped, axillary pair of
folds; eye diameter usually less than 35 per cent of head breadth;
first finger length usually more than 85 per cent of third finger
length; posterior surface of lower leg with relatively conspicuous,
irregularly margined, blackish, longitudinal band gilliardi
Platymantis boulengeri (Boettger )
Cornufer boulengeri Boettger, 1892, Katalog der Batrachier-sammlung
im Museum der Senckenbergishen Naturforschenden Gesellschaft in
Frankfort am Main, p. 18: New Britain Island.
Definition: A large species for the genus, 6 mature females range
from about 66 to 79.7 mm snout-vent length; head much broader than
long; head breadth/tibia length = 1.00 (6: 0.845-1.11), at least for
females; snout round-pointed; canthus rostralis rather sharply rounded;
loreals strongly oblique, slightly concave; eye relatively small, its diam-
eter/snout length = 0.776 (6: 0.702-0.880) and diameter/head
breadth = 0.257 (6: 0.227-0.320); finger tips rounded, without dilated
disks; fingers relatively short (Fig. la), third finger length/head breadth
= 0.284 (6: 0.255—-0.303) and third finger length/snout length = 0.848
(6: 0.772-0.916); first and second fingers about equal in length; first
finger length/third finger length = 0.691 (6: 0.647-0.734); subarticular
tubercles, large, prominent, rounded; inner metacarpal tubercle broadly
oval; outer narrow, elongate; tibia relatively short, its length/snout-vent
length = 0.453 (6: 0.435-0.489 ); toes webbed at base; toe disks some-
what broader than penultimate phalanx; subarticular tubercles smaller
than those of fingers, prominent, somewhat pointed; inner metacarpal
tubercle about twice as broad as long; outer small, round; solar tubercles
small and scattered; dorsum marked by several very narrow, usually
broken ridges, more or less parallel posterior to the axillary region but
anteriorly diverging in a graceful fanlike pattern.
Color (in preservative): Dorsal ground color grayish tan to brown;
sides of head to posterior end of supratympanic fold, dark brown to
blackish; limbs marked by dark and light bands; venter heavily mottled
or flecked with grayish brown; in life, thoracic and throat regions and
posterior surface of thigh and lower leg reddish; light spots in thoracic
and throat regions pale yellow or grayish.
Remarks: This large species is not obviously, closely related to other
72 Proceedings of the Biological Society of Washington
a b
Fic. 1. (a) Platymantis boulengeri, inferior view of hand; (b) Platy-
mantis gilliardi, inferior view of hand.
known species. No size data are available for males, but the smallest
recorded mature female is 66 mm snout-vent length. Zweifel (personal
correspondence ) states inat the 54 mm specimen which he shows on the
graph (1960, p. 9) is an immature female.
Platymantis gilliardi Zweifel
Platymantis gilliardi Zweifel, 1960, Amer. Mus. Noviates, no. 2023, p. 10:
New Britain Island.
The 10 examples of this species, South Australian Museum nos. 6866,
6946, 7023, 7062-63, 7068, 7026 and Stanford University nos. 22877—78
collected at low altitudes in New Britain, makes possible a more adequate
definition in terms of variability.
Definition: A moderate-sized species for the genus, 7 mature females
range from 40.9 to 48.7 mm snout-vent length (a female measuring
39.9 mm is apparently not quite mature judging by the fact that the
oviducts are straight and undilated), 2 mature males measure 33.2 to
34.2 mm; head breadth/snout-vent length = 0.419 (10: 0.411-0.442);
head-breadth/tibia length = 0.815 (10: 0.765-0.866) snout rounded to
round-pointed; canthus rostralis rounded; loreals somewhat oblique,
slightly concave; eye moderate, its diameter/snout-length = 1.01 (10:
0.944-1.11), diameter/head breadth = 0.321 (10: 0.282-0.345); third
finger moderately long, its length/head-breadth = 0.303 (10: 0.283-
0.319); its length/snout-length = 0.944 (10: 0.855-1.02); finger tips
pointed or round-pointed, not or scarcely dilated (Fig. 1b); very shallow
circummarginal groove usually present, first finger, longer than second,
nearly as long as third when fingers are adpressed; first finger length/
third finger length = 0.873 (10: 0.833-0.919); subarticular tubercles,
Frogs of the Genus Platymantis 1
a b
Fic. 2. (a) Platymantis gilliardi, dorsal-fold pattern; (b) Platymantis
mimicus, dorsal-fold pattern.
large, prominent, rather pointed; inner metacarpal tubercle large, nearly
as broad as long; outer broadly oval; palmar tubercles relatively large
and prominent; tibia relatively long, its length/snout-vent length = 0.522
(10: 0.479-0.584); toe disks slightly broader than penultimate phalanx,
upper part separated from inferior pad by a marginal groove; subarticular
tubercles moderately large, strongly protruding, pointed; inner metatarsal
tubercle about twice as broad as long; outer strongly protruding, round;
solar tubercles prominent and numerous; dorsum relatively smooth except
for small granules and a pair of urn-shaped folds between post-orbital
and axillary regions (Fig. 2a).
Color (in preservative): Dorsal ground color variable, from light gray
to dark reddish-brown, relatively uniform or with a few black spots in
axillary and groin regions; sometimes with darker areas around prominent
dorsal folds or tubercles; limbs usually with moderately contrasting,
dark and light, transverse bands.
Remarks: The present series indicates that P. gilliardi is indeed a close
relative of P. acrochordus (Brown). As pointed out by Brown (1965,
p. 12), P. acrochordus differs in its denser dorsal rugosity, slightly smaller
size at maturity, slightly broader head relative to snout-vent length (for
acrochordus this ratio is 0.444 [11: 0.427-0.460]) or tibia length (for
acrochordus this ratio is 0.935 [13: 0.889-1.00]); somewhat shorter tibia
relative to snout-vent length (for acrochordus this ratio is 0.475 [13:
0.441-0.501]); and shorter snout relative to third finger length (for
acrochordus this ratio is 0.800 [13: 0.705-0.913]).
74 Proceedings of the Biological Society of Washington
a b
Fic. 3. (a) Platymantis mimicus, inferior view of hand; (b) Platy-
mantis akarithymus, inferior view of hand.
Platymantis mimicus new species
Holotype: South Australian Museum no. 6868, a mature male, col-
lected at Numundo Plantation, Willaumez Peninsula on the north coast
of New Britain Island, 19 January 1966, by M. J. Tyler.
Paratypes: Stanford University no. 22874, from same locality as
holotype; South Australian Museum nos. 7064 and 7069, Pomugu about
7 miles northwest of Kandrain, south coast of New Britain and no. 6864
from Gazelle Peninsula, northeast coast of New Britain.
Description: A moderate-sized Platymantis, 4 mature males measure
27 to 40 mm snout-vent length, single mature female 45.1 mm snout-
vent length; habitus relatively slender; head gradually tapering, snout
rounded, upper jaw round-pointed; head-breadth/snout-vent length =
0.404 (5: 0.386-0.420) and head-breadth/tibia length = 0.776 (5:
0.717—0.839 ); eye large, its diameter/snout length = 1.07 (5: 1.04—1.11)
and diameter/head breadth = 0.341 (5: 0.314—0.357), canthus rostralis
rounded, loreal region oblique, moderately concave; fold dorsal and
posterior to the tympanum; tympanum diameter/eye diameter = 0.542
(5: 0.500-0.578); fore limbs well developed; tips of fingers not or
scarcely dilated, rounded, with faint evidence of marginal groove separat-
ing dorsal and ventral portions; first finger usually longer than second
or fourth but shorter than third when fingers are adpressed (Fig. 3a);
first finger length/third finger = 0.801 (5: 0.737—0.838); third finger
relatively long, its length/snout length = 1.05 (5: 1.00-1.13) and
length/head breadth = 0.335 (5: 0.308-0.372); subarticular tubercles
strongly developed, rounded to pointed, supernumary palmer tubercles
moderately developed; metacarpal tubercles large, prominent, inner and
middle one broad, the outer more narrow; hind limbs relatively long;
Frogs of the Genus Platymantis 75
a
Fic. 4. (a) Platymantis papuensis schmidti, inferior view of hand;
(b) Platymantis rhipiphalcus, inferior view of hand.
tibia length/snout-vent length = 0.523 (5: 0.404—0.541); toes with
minute web at base; tips slightly dilated, rounded, ventral portion of
disk, separated from the dorsal portion by circummarginal groove; sub-
articular tubercles well developed, rather pointed; inner metatarsal
tubercle large, rather broad, outer small, conical; solar tubercles scattered,
prominent; skin of dorsum with numerous short to moderately long,
rather thick folds, (Fig. 2b) least conspicuous for the 44 mm male,
dorsum, between folds, and lateral surfaces with low, round granules,
venter with flat granules on thorax and abdomen.
Measurements of holotype (in mm): Snout-vent length 29.0; head
length to posterior edge of tympanum 11.1; head breadth 11.2; eye
diameter 4.0; tympanum diameter 2.4; snout length 3.6; third finger
length 3.8; first finger length 3.1; tibia length 14.9; hind limb length
48.5.
Color: Dorsal ground color grayish to dark reddish brown, relatively
uniform or with some lighter or darker blotches; light interorbital area
extending forward on snout; lips dark barred; lower lateral surfaces
marked by light, rounded spots on grayish background; venter rather
uniformly light on thorax and abdomen, on throat and head with dark
flecks; hind limbs marked by dark, transverse bands; under surface of
feet dark.
Etymology: The species name, mimicus, is from the Latin meaning
imitative.
76 Proceedings of the Biological Society of Washington
Remarks: This third, previously undescribed species of the acrochor-
dus-gilliardi group was first noted, during field work, to differ from
gilliardi in its distinct call and calling site. P. gilliardi calls from open
spaces on the ground, stems of low bushes such as ornamental crotons
or trunks of banana trees; the new species calls from concealed places
beneath piles of leaves and coconut husks. Though very similar in the
nature of the finger and toe disks and in general appearance, P. mimicus
may be distinguished from both gilliardi and acrochordus on the basis
of the slightly narrower head relative to the tibia length or snout-vent
length, the greater length of the third finger relative to the snout-length
or the head breadth (see ratios under definition and remarks for
gilliardi), and the absence of a conspicuous urnshaped pair of folds in
the axillary regions. From gilliardi, it differs further in the shorter first
finger relative to the third finger length, the more numerous dorsal
folds (Fig. 4), and the lack of a conspicuous, blackish, longitudinal band
on the posterior surface of the lower leg and from acrochordus in the
narrower head relative to tibia length (for acrochordus this ratio is
0.935 [13: 0.889-1.00] ).
Platymantis akarithymus new species
Holotype: South Australian Museum no. 7073, a mature male, col-
lected in the Pomugu area about seven miles northwest of Kandrian on
the south coast of New Britain Island by M. J. Tyler, 31 January 1966.
Paratypes: South Australian Museum no. 6982 and Stanford University
no. 22875 from same locality as holotype; South Australian Museum
no. 7066 and 7082 from near Malassait 40-++ miles west of Rabaul in
the Baining Ranges of Gazelle Peninsula on northeast coast of New
Britain Island.
Description: A small Platymantis, at least for males, 5 mature males
measure 18.5 to 24.2 mm snout-vent length; habitus moderately slender;
head about as broad as long, breadth/head length = 0.994 (5: 0.960—
1.04); head breadth/tibia length = 0.780 (5: 0.707-0.852); snout
pointed; upper jaw only slightly protruding; eye large, its diameter/snout-
length = 1.03 (5: 1.00-1.13) and diameter/head breadth = 0.391 (5:
0.360-0.421); tympanum distinct, its diameter/eye diameter = 0.535
(5: 0.500-0.556), and diameter/head breadth = 0.210 (5: 0.199—
0.232); canthus rostralis rounded, loreal region slightly oblique, moder-
ately concave; fold dorsal and posterior to the typmpanum; fore limbs
well developed; fingers rounded at tip, not or but slightly dilated; ventral
portion at tip delimited laterally and terminally by shallow groove, except
for inner finger; fingers without webs; subarticular tubercles strongly
developed, rather sharp pointed; transverse row of moderately developed
palmer tubercles; inner and middle metacarpal tubercles large, slightly
longer than broad, outer small, round to oval; first, second and fourth
finger about equal in length when fingers are adpressed (Fig. 3b); first
finger length/third finger length = 0.649 (5: 0.610-0.700); third finger
length/snout length = 0.993 (5: 0.947-1.03); hind limbs relatively long;
Frogs of the Genus Platymantis 77
tibia length/snout-vent length = 0.499 (5: 0.484-0.519); toes without
webs, tips not or but very slightly dilated, rounded, ventral portion of tip
separated from dorsal portion by groove laterally and terminally; subartic-
ular tubercles well developed, somewhat pointed; inner metatarsal tuber-
cle broadly oval, outer round and rather pointed; solar tubercles evident
but not well developed; skin of dorsum, upper lateral surfaces, and to
some degree limb surfaces marked by numerous prominent, round, oval
or somewhat elongate tubercles; venter, posterior to axilla, and ventral
and posterior thighs with small, rounded granules.
Measurements of holotype (in mm): Snout-vent length 24.0 head
length, measured to posterior edge of tympanum 9.8; head breadth 9.5;
eye diameter 4.0; tympanum diameter 2.2; snout length 4.0; third finger
length 4.1; tibia length 12.1; length of hind limb 40.7.
Color (in preservative): Dorsum generally darker on middorsal area
than on dorso-lateral or lateral surfaces; but with overall pattern of
irregular, light and dark blotches; most of short dorsal folds and tubercles
and immediately surrounding areas blackish; light patch present in
anterior orbital area; limbs and jaws barred with prominent light and
dark bands; bright red patch in life (fading in preservative) in groin
and on back of thigh for this small, all-male sample; venter is heavily
mottled with dark flecks, especially anterior to fore limbs.
Etymology: The species name akarithymus is from the Greek akares
meaning short and thymos meaning warty growth.
Remarks: Platymantis akarithymus apparently has its closest affinities
with that group of small to moderate species which have finger tips not
or but very slightly dilated. These include P. parkeri (Brown) and P.
aculeodactylus Brown from the Solomons, P. pelewensis Peters from the
Palau Islands, and P. dorsalis A. Duméril from the Philippines. It is most
similar to P. parkeri in the following combination of characters: shape,
small size and abundance of the dorsal tubercles; prominence and shape
of the tubercles on the undersurface of the hands and feet; the undilated
finger tips; and size at maturity (as measured by snout-vent length).
It differs from the latter in the greater snout-vent length at maturity,
the greater length of the tibia relative to the snout-vent length (for
parkeri, tibia length/snout-vent length = 0.458 [20: 0.428-0.484]) and
the larger tympanum relative to the eye diameter (for parkeri, tympanum
diameter/eye diameter = 0.640 [15: 0.560-0.700]) or the head breadth
(for parkeri, tympanum diameter/head breadth = 0.235 [15: 0.210-
0.255]). Also, the first and second fingers are about equal in length
in the sample of P. akarithymus available to us, whereas the first finger
is usually distinctly shorter than the second in P. parkeri.
Platymantis rhipiphaleus new species
Holotype: South Australian Museum no. 7071, a male, collected in
the Pomugu area about 7 miles northwest of Kandrian on the south
coast of New Britain Island by M. J. Tyler, 30 January 1966.
78 Proceedings of the Biological Society of Washington
Paratypes: South Australian Museum no. 7078 and Stanford Univer-
sity no. 22873, collected at San Remo plantation on Willaumez Peninsula
about 150 miles west of Rabaul on the north coast on New Britain by
M. J. Tyler, 20 January 1966.
Description: A moderately large Platymantis, at least for females, (no.
7078, measuring 41.4 mm snout-vent length, is immature, with small
ovaries and straight, undilated oviducts); the unique, mature male
measures 28.4 mm; habitus relatively slender; head depressed; snout
pointed; upper jaw protruding; head about as broad as or slightly broader
than long; head breadth/snout-vent length = 0.413 (3: 0.401-0.421),
head breadth/tibia length = 0.799 (3: 0.776—0.844); eye moderate, its
diameter/snout-length = 0.900 (3: 0.877—0.930); tympanum large, its
diameter/eye diameter = 0.663 (3: 0.580-0.750); tympanum diameter/
head breadth = 0.231 (3: 0.207-0.263); canthus rostralis rounded,
loreal moderately oblique; a fold dorsal and posterior to tympanum; fore
limbs well developed; tips of fingers depressed and moderately dilated
into rounded disks, ventral portion separated from dorsal by circum-
marginal groove; diameter of third finger disk/tympanum diameter =
0.439 (3: 0.400-0.483); diameter of third finger disk/third finger length
= 0.261 (3: 0.250-0.267); subarticular tubercles large and strongly -
developed as are inner, middle and outer metacarpal tubercles; low
fringe on outer edge of fourth finger and distal metacarpal region, first
finger slightly shorter than second which is shorter than fourth when
fingers are adpressed (Fig. 4b); third finger length/snout length = 0.974
(3: 0.938-1.00); first finger length/third finger length = 0.613 (3:
0.578-0.633); hind limbs relatively long, tibia length/snout-vent length
= 0.518 (3: 0.499-0.536); toes with minute web at base; tips of toes
depressed, dilated into slightly pointed disks somewhat smaller than
those of fingers, diameter of third finger disk/diameter of third toe
disk = 1.13 (3: 1.07-1.20); ventral portion of disk delimited from
dorsal by circummarginal groove, subarticular tubercles large, well
developed; inner metatarsal tubercle elongate, outer round; solar tubercles
small, low, not very numerous. Skin of dorsum with several narrow
ridges, relatively uninterrupted from head or axillary regions to groin,
diverging in gracefully curved, fanlike pattern anterior to axillary region;
skin granular on posterior venter and posterior surface of thighs.
Measurements of holotype (in mm): Snout-vent length 28.4; head
length measured to posterior edge of tympanum 11.8; head breadth 11.4;
eye diameter 4.0; tympanum diameter 3.0; snout length 4.5; third finger
length 4.5; diameter of third finger disk 1.2; diameter of third toe disk
1.0; tibia length 14.7; length of hind limb 47.5.
Color (in preservative): Dorsum variable, light grayish brown to dark
reddish, with darker interorbital bar; relatively uniform posterior to head,
or with few small darker spots or darker, broad, middorsal stripe; hind
limbs with rather indistinct dark cross bands; anal region grayish or
blackish; dark band of varying width from snout through ear; venter
light with faint specks of pigment for holotype, paratypes with expanded
Frogs of the Genus Platymantis 79
pigment cells, exhibiting brownish blotches, heaviest on chin, throat
and fore limb region.
Etymology: The species name, rhipiphalcus, is from the Greek rhipis
meaning fan and phalkes meaning rib; referring to the arrangement of
the narrow dorsal folds.
Remarks: The relatively long, very narrow dorsal folds which exhibit
a fanlike pattern anterior to the forelimbs also characterizes P. boulengeri
from New Britain and frequently P. corrugatus (Duméril) from the
Philippine Islands. In size of finger disks relative to finger length, P.
thipiphalcus resembles P. papuensis schmidti.
From the Philippine P. corrugatus, P. rhipiphalcus is readily dis-
tinguished by its more dilated finger tips and relatively shorter first
finger. The dilated finger tips also readily separate this species from
P. boulengeri. Other characteristics which readily distinguish it from
P. boulengeri are its narrower head relative to head or tibia length, its
larger eye relative to snout length or head breadth, and the longer tibia
relative to snout-vent length. In these proportions, only the 27 mm
juvenile of boulengeri is similar to the specimens of rhipiphalcus. The
juvenile boulengeri, for example, is the only specimen of this species
with a ratio of head breadth to snout-vent length below 0.430. The
largest female of P. rhipiphalcus (41.1 mm snout-vent length) with
small, straight oviducts, is clearly immature, though it might well have
become mature long before attaining a length of 66 mm, the size of the
smallest female of boulengeri in our series. The male (holotype) is
adult at 28.4 mm snout-vent length. Unfortunately, size at maturity for
males of boulengeri is not known.
P. macrops (Brown) and P. meyersi (Brown) both lack the distinc-
tive, long narrow dorsal folds. P. macrops is also distinguished by its
much smaller size and larger eye. P. meyersi is further distinguished by
the smaller disks relative to finger length (for meyersi this ratio is 0.127
[16: 0.11-0.156]), and relative to tympanum diameter (for meyersi this
ratio is 0.374 [18: 0.337-0.421]).
Platymantis papuensis Meyer
Platymantis corrugatus papuensis Meyer, 1874, Monatsber. Akad. Wiss.
Berlin, p. 1389: Biak Island, Dutch New Guinea.
A group of morphologically very similar species, which range from the
Pelau and Philippine Islands in the north to New Guinea and the Solomon
Islands in the south, have been assigned by numerous previous authors
to P. corrugatus or Rana rugata (e.g.) Werner 1900, van Kampen 1923,
and Hediger 1934. Loveridge (1948, p. 407) continued to follow this
broad interpretation of P. corrugatus, designating the New Guinean
population, as did Meyer, a subspecies under the name P. corrugatus
papuensis. Brown (1952, p. 51) placed corrugatus from the Philippines
and papuensis from New Guinea and the Solomons in different species.
Inger (1954, p. 351 ff.) came to the same conclusion, and pointed out
80 Proceedings of the Biological Society of Washington
that corrugatus and papuensis are actually less closely related than are
papuensis and meyeri (= dorsalis), another Philippine species. The
definition which follows is based upon samples from the populations of
New Guinea, the Bismarks, and the Solomons.
Definition: A moderately large species of the genus, 48 mature females
from these populations measure 39.8 to 63.6 mm snout-vent length, and
43 mature males measure 29.8 to 45.8 mm; habitus relatively slender;
head breadth slightly less than to slightly greater than head length; head
breadth/snout-vent length = 0.397 (48: 0.363-0.439), head breadth/
tibia length = 0.745 (48: 0.632-0.809); eye moderate, its diameter/
snout length = 0.543 (48: 0.446-0.653); tympanum distinct and moder-
ately large; its diameter/eye diameter = 0.543 (48: 0.446-0.571); snout
rounded to broadly round-pointed; loreals somewhat oblique and shal-
lowly concave; finger tips bluntly rounded or with depressed slightly
dilated disks; first finger varies from slightly shorter than to somewhat
longer than second, which is shorter than fourth when fingers are
adpressed; first finger length/third finger length = 0.732 (48: 0.578—
0.833); snout length/third finger length = 1.068 (48: 0.896-1.32);
subarticular tubercles moderate to large, strongly protruding, round;
inner metacarpal tubercle about half as broad to nearly as broad as long; .
outer broadly oval to nearly round; supernumary tubercles at base of
fingers prominent (Fig. 4a); hind limbs relatively long, tibia length/
snout-vent length = 0.535 (48: 0.473—0.593); toes with minute web at
base; tips of toes dilated into small, depressed disks; upper and lower
portions delimited by circummarginal groove; toe disks more broadly
dilated than, or about equal to, finger tips or disks, subarticular tubercles
well developed, somewhat pointed; inner metatarsal tubercle about half
as broad to nearly as broad as long; outer small to moderate, round;
solar tubercles low and scattered to relatively prominent and rather
numerous for the different races; skin of dorsum marked by sparse to
moderately numerous, short or moderately elongate folds; venter rela-
tively smooth.
Color (in preservative): Dorsal ground color variable, gray to grayish
red or brown, relatively uniform or with scattered dark blotches, lighter
middorsal area or dorsolateral stripes present in some instances; lateral
surfaces somewhat lighter, lips with or without distinct transverse bars;
hind limbs with relatively inconspicuous, dark, transverse bands; venter
uniformly light or with scattered dark flecks anteriorly.
Remarks: Moderately large size at maturity and pattern of the dorsal
folds, as well as the limited dilation of the digital tips, suggest possible
affinities of P. papuensis with P. myersi and P. solomonis Boulenger
among species sympatric with P. papuensis. As noted by Inger (1954,
p. 355), among nonsympatric species, P. papuensis probably has affinities
with P. dorsalis (= meyeri of Inger) from the Philippines and P. pele-
wensis Peters from the Palau Islands.
P. solomonis and P. myersi both occur in the Solomon Islands along
with P. papuensis weberi. Both myersi and solomonis are somewhat
Frogs of the Genus Platymantis 81
larger at maturity than papuensis (Table 1). P. myersi further differs
from P. papuensis in the lack of solar tubercles on the feet, its smaller
tympanum relative to the size of its eye (Table 1), and the slightly
greater webbing of its toes. From the sympatric (Solomons) population
of papuensis weberi, myersi also differs in the greater finger disk breadth
relative to toe disk breadth.
The finger tips of P. solomonis are bluntly rounded, not depressed,
with dorsal and ventral parts of the disk separated by a circummarginal
groove as is typically characteristic of P. papuensis weberi and P.
papuensis schmidti. In this characteristic, the structure of the finger
tips, P. papuensis from New Guinea appears more variable and many
specimens closely resemble P. solomonis. The ratio of shorter tibia
relative to snout-vent length (rarely as great as 50 per cent for P.
solomonis and rarely less than 50 per cent for P. papuensis) (Tables 1
and 2) also serves to distinguish these two species. The New Guinea
subspecies again exhibits the greatest overlap with solomonis for this
characteristic.
The subspecies of P. papuensis
Our comparison of samples of reasonable size of the three geograph-
ically isolated, sibling populations from New Guinea, the Bismarks and
the Solomons indicate close affinity in terms of similarities in a number
of anatomical characteristics. These include: (1) rather narrow, rela-
tively short or interrupted folds on the dorsum; (2) tibia length usually
more than 50 per cent of the snout-vent length; (3) head breadth about
40 per cent of the snout-vent length and about equal to the head length;
(4) length of third finger slightly less than to slightly greater than the
snout length; (5) diameter of eye about twice the diameter of the
tympanum, and 80 to 100 per cent of the snout length; (6) disks of
toes small, rounded, slightly broader than the subtending phalanx; (7)
finger tips bluntly rounded or slightly dilated and depressed into small
disks; (8) solar tubercles present, moderately or conspicuously protrud-
ing; (9) color pattern frequently exhibits broad dorsolateral light stripes.
It is interesting to note that the Solomons and New Guinea popula-
tions exhibit greater similarity in many characteristics than do either
with the intermediate New Britain population. However, our data shows
that the three populations can be distinguished on the basis of a combina-
tion of several characteristics. We therefore propose that all three be
given subspecific status in the species papuensis. The most useful dis-
tinguishing characteristics are given in the key to the subspecies.
The probable conspecific relationships of the populations of New
Guinea and the Solomons has been recognized previously, and the
population of the Solomon Islands designated as a subspecies under
the name Platymantis papuensis weberi Schmidt (Brown, 1952, p. 50).
This leaves the geographically intermediate sibling population in the
Bismark archipelago nameless, in the sense that it has not been referred
to either subspecies.
f the Biological Society of Washington
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84 Proceedings of the Biological Society of Washington
Inger (1954: p. 355) inadvertently referred this population to Cornufer
(= Platymantis) nova-britannea (a manuscript name of Schmidt)
creating a nomen nudem as noted by Zweifel (1960, p. 22). Zweifel
(op. cit.) further stated that he lacked adequate data to determine the
subspecific status of the New Britain population.
Key to subspecies of Platymantis papuensis
1. Finger tips dilated into disks broader than subtending phalanx,
breadth rarely less than 85 per cent of toe disk breadth, and
usually more than 35 per cent of tympanum diameter; first finger
length about equal to second, distinctly shorter than fourth,
aYonnan anova INGA Ioateniay papuensis schmidti
1’. Finger tips little dilated, not or slightly broader than subtending
phalanx, breadth usually less than 85 per cent of toe disk breadth,
usually Jess than 35 per cent of tympanum diameter; first finger
usually longer than 2nd, nearly equal to fourth 2)
2. Females immature at less than 50 mm snout-vent length; dorsal
folds narrow, numerous, elongate; known from Solomon Islands
RIE one DP aPIN UR oeneanD nia Wing Su pNtanel MON ST sce I Ns papuensis weberi
2’. Females mature between 40 and 50 mm snout-vent length; dorsal
folds not extremely narrow, scattered, mostly short; known from
INew, sGuineat ie alien Te ered Nat Mea eR papuensis papuensis
Platymantis p. papunesis Meyer
Platymantis corrugatus papuensis Meyer, 1874, Monatsber. Akad. Wiss.
Berlin, p. 139: Biak Id., Netherlands New Guinea.
Diagnosis: A moderately large Platymantis, 19 mature females measur-
ing from about 43 to 58 mm snout-vent length (a 39.3 mm specimen is
judged immature based on the undilated and relatively straight oviduct);
relatively short dorsal folds, most numerous anteriorly, solar tubercles
scattered, relatively inconspicuous, breadth of third finger tip about 70
to 90 (usually more than 80) per cent of breadth of third toe disk, less
than 20 per cent of 3rd finger length and about 30 per cent of tympanum
diameter (Table 2); known from New Guinea. Our sample is largely
from southeast New Guinea. )
Platymantis papuensis weberi Schmidt
Platymantis weberi Schmidt, 1932, Field Mus. Nat. Hist. Zool. Series, 18,
p. 178: Tulagi Id., Solomon Ids.
Diagnosis: A large Platymantis, 16 mature females measuring from
54 to 64 mm snout-vent length (a 48.8 mm specimen is judged immature,
based on the undilated and relatively straight oviducts); moderately
short dorsal folds over entire dorsum; solar tubercles numerous and
prominent; breadth of third finger tip 55 to 85 (usually less than 70)
per cent of breadth of third toe disk, less than 20 per cent of 3rd finger
Frogs of the Genus Platymantis 85
length and less than 30 per cent of tympanum diameter (Table 2);
known from the Solomon Islands.
Platymantis papuensis schmidti new subspecies
Holotype: South Australian Museum no. 7618, a gravid female, col-
lected at Talasea, Willaumez Peninsula on the north coast of New Britain
Island, by M. J. Tyler, January 1966. This locality is near sea level.
Paratypes: South Australian Museum nos. 6762-68, 6772-93, 6795,
6801, 6803-07, 6809-13, 6815-16, 6858-60, 6862, 6869, 6912-13, 6915,
6922-28, 7061, 7070, 7080, 7085, 7088-89, 7093, 7095, 7097, 7101-04,
7106, 7109, 7115 from the Willaumez Peninsula; 7615-23, 7625-74,
7677-78 from the Baining Ranges, north coast of New Britain Island;
nos. 7043, 7045, 7099, 7132, 7134-37, 7139, 7147-48, 7151 from the
Keravat area, Gazelle Peninsula, 20 miles west of Rabaul; American
Museum of Natural History nos. 64258—59, 64261 and 64264, Whiteman
Mountains, New Britain Islands; Stanford University nos. 22879, Wil-
laumez Peninsula and 22880—82, Baining Ranges, New Britain Island.
Diagnosis: A moderately large Platymantis, mature females measuring
from about 40 to 60 mm snout-vent length; males somewhat smaller;
tips of fingers dilated into moderate, depressed disks, much broader than
subtending phalanx; 3rd finger disk diameter 80 to 100 per cent of 3rd
toe disk, usually more than 20 per cent of third finger length, and 33
to 50 per cent of tympanum diameter; Ist finger length about 70 per
cent of 3rd finger length (Table 2); dorsal folds moderate to short,
usually scattered over entire dorsum; solar tubercles scattered, relatively
inconspicuous, known from New Britain Island.
Measurements (in mm) of holotype: Snout-vent length 56.9; head
length measured to posterior edge of tympanum 21.2; head breadth 21.8;
eye diameter 7.7; tympanum diameter 3.9; snout length 8.8; third finger
length 8.7; diameter of third finger disk 1.8; diameter of third toe disk
1.5; tibia length 32.7; length of hind limb 102.5.
Etymology: Named for Dr. Karl Schmidt who first recognized the
validity of this population as a distinct taxonomic entity.
ACKNOWLEDGMENTS
We wish to thank Dr. Emest Williams, Museum of Comparative
Zoology, Harvard University and Dr. Richard Zweifel, American Museum
of Natural History, New York, for their assistance in making available
pertinent material in the collections of their institutions. Drawings are
by Mr. Walter Zawojski, Stanford University.
The study of this genus of frogs has been a part of the senior author’s
program concerned with the herpetofauna of the Pacific Islands. This
program is sponsored by the National Science Foundation, grant No.
GB-4156. The junior author's visit to New Britain was made possible
by a grant generously provided by the Director and Trustees of the
South Australian Museum.
86 Proceedings of the Biological Society of Washington
LITERATURE CITED
BROWN, WALTER C. 1952. The amphibians of the Solomon Islands.
Bull. Mus. Comp. Zool., 107: 1-64, pls. 1-8.
1965. New frogs of the genus Cornufer (Ranidae) from
the Solomon Islands. Breviora, 218: 1-14, pls. 15-16.
GorHAM, STANLEY W. 1965. Fiji frogs (with synopses of the genera
Conufer and Platymantis). Zoologische Beitrage, 25: 381-
424, pls. 425-435.
Hepicer, Hemi. 1934. Beitrag zur Herpetologie und Zoogeographie
Neu Britanniens. Zool. Jahrb., Abt. Syst., 65: 441-582, 6
Figs.
IncER, Ropert F. 1954. Systematics and zoogeography of Philippine
amphibia. Fieldiana, Zool. 33: 183-531.
Kampen, P. N. van. 1923. The amphibia of the Indo-Australian
Archipelago. xii + 304 pp. Leiden, E. J. Brill Ltd.
LovermncE, ArTHUR. 1948. New Guinean reptiles and amphibians in
the Museum of Comparative Zoology and the United States
National Museum. Bull. Mus. Comp. Zool., 101: 305-430.
TyLer, Micuaet J. 1964. Transfer of the New Britain frog Hyla
brachypus (Werner) to the microhylid genus Oreophryne.
Mitt. Zool. Mus. Berl. 40 Heft 1: 3-8.
WERNER, Franz. J900. Die Reptilian und Batrachierfauna des Bis-
march-Archipels. Mitt. Zool. Mus. Berlin, 1, Heft 4: 1-132,
46 Figs.
ZWEIFEL, RicHarpD G. 1960. Results of the 1958-1959 Gilliard New
Britain Expedition 3. Notes on the frogs of New Britain.
Amer. Mus. Novitates, 2023: 1-27.
. Identity of the frog Cornufer unicolor and application of the
generic name Cornufer. Copeia, 1967, No. 1: 117-121.
74.0673
Vol. 81, pp. 87-90 eae April 30, 1968
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
A NEW SPECIES OF DENNYUS
(MALLOPHAGA: MENOPONIDAE) FROM
THE MALAYSIAN SPINE-TAILED SWIFT
By K.C. Emerson AND RoceEr D. PRICE
Arlington, Virginia and Department of Entomology,
Fisheries, and Wildlife, University of Minnesota,
St. Paul, Minnesota
Recently the authors obtained specimens of a new species
of Mallophaga, which is herewith described, illustrated, and
compared with a closely related species.
Dennyus major (Uchida, 1926)
Dennyus major (Uchida, 1926) is found on the Northern Spine-tailed
Swift, Chaetura caudacutus caudacutus (Latham). Uchida described
the species from specimens collected in Togakushimura and Chikumagun,
both in Nagano Prefecture, Japan. Nakagawa (1959) redescribed the
female from a specimen collected in Hokkaido, Japan. Both authors
noted that the species was the largest in the genus and had many
distinctive features. Uchida also noted the unusual genitalia, which we
have illustrated in Fig. 5. During this study, Dr. Theresa Clay lent us
a male and a female from the type series. Since Uchida did not designate
a holotype, the male with collection data “Takamatsuia major, Chaetura
c. caudacuta, JAPAN, Nagao, 1917, #162” is hereby designated lecto-
type and has been so labeled. The labels presently on these two slides
are not the original Uchida labels, which accounts for the incompleteness
of the locality data and for the fact that the generic name of the host
was changed from Hirundapus to Chaetura.
Dennyus giganteus new species
Holotype male: Total length, 3.13 mm. External morphology and
chaetotaxy as illustrated in Fig. 2. Sternal plate of prothorax as illustrated
in Fig. 4. Head, thorax (except for prosternal plate) and legs essentially
the same as in Dennyus major. All setae on the prosternal plate are
longer than in D. major. Abdominal chaetotaxy, as illustrated in Fig. 2,
with fewer short, fine marginal setae on segments I-VI than in D. major.
10—Proc. Brow. Soc. Wasu., Vou. 81, 1968 (87)
88 Proceedings of the Biological Society of Washington
See
i i
y
\Z = fay Ne if:
So Sy
Boe
Ln
: ;
2
5
Fics. 1-4. Dennyus giganteus, new species. 1, dorsal-ventral view of
female; 2, dorsal-ventral view of male; 3, male genitalia (less sac); 4,
sternal plate of prothorax. Fic. 5. Dennyus major (Uchida), male
genitalia (less sac).
New Mallophagan from Malaysian Swift 89
{ndentation of terminal abdominal segment deeper than in D. major.
Genitalia, less sac, as illustrated in Fig. 3.
Allotype female: Total length, 3.78 mm. External morphology and
chaetotaxy as illustrated in Fig. 1. Abdominal chaetotaxy, as illustrated
in Fig. 1, with fewer short, fine marginal setae on segments I-VI than
in D. major. Anal fringe with more heavy thick long peglike setae on
each side than D. major.
Type host: Chaetura gigantea gigantea (Temminck), Malaysian Spine-
tailed Swift.
Type material: Holotype male and two paratypes collected off the
type host on 14 October 1964 at Kabigaan Aborlan, Palawan, Philippines.
Allotype female and one paratype collected off Chaetura gigantea dubia
(McGregor) on 27 January 1966 at Dalton Pass, N. Vizcaya, Philippines.
Two paratypes collected off Chaetura gigantea dubia (McGregor) on
27 June 1965 at Dalton Pass, N. Vizcaya, Philippines. Two paratypes
collected off Chaetura gigantea indica (Hume) in October 1943 in
Punjab, India. The holotype and allotype will be placed in the collection
of the U.S. National Museum. The authors will each retain one pair
of paratypes from the Philippine material. The Indian material is in
the British Museum (Natural History).
Discussion: Peters (1940) recognized the genus Hirundapus for three
species of swifts: caudacutus, giganteus, and ernsti. He said of ernsti,
known only from the unique type, “Probably a race of giganteus, but
until more is known of its distribution, is better regarded as a distinct
species.” Several authors since Peters have included these swifts in the
genus Chaetura; but the large size, indented terminal abdominal segment,
and male genitalia are unique features of the species of Dennyus found
on Hirundapus as recognized by Peters.
LITERATURE CITED
Naxacawa, H. 1959. A redescription of Dennyus major (Uchida)
from the Needle-tailed Swift Hirundapus caudacutus cauda-
cutus (Mallophaga). J. Agri. Sci. Tokyo Nogyo Daigaku,
5: 23-27.
Peters, J. L. 1940. Check-list of birds of the World, Volume IV.
Harvard University Press. 291 pp.
Ucumpa, S. 1926. Studies on Amblycerous Mallophaga of Japan. J.
Col. Agri., Imperial Univ. of Tokyo, 9: 1-56.
90 Proceedings of the Biological Society of Washington
WA OG ZS
Vol. 81, pp. 91-96 ‘ April 30, 1968
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
A NEW MARINE DINOFLAGELLATE GENUS,
CACHONINA, IN AXENIC CULTURE FROM THE
SALTON SEA, CALIFORNIA WITH REMARKS
ON THE GENUS PERIDINIUM
By Aurrep R. Loesuicnu, III
Scripps Institution of Oceanography
University of California at San Diego
La Jolla, California 92037
An armored, photosynthetic dinoflagellate was collected
from a bloom in the Salton Sea, California (March 1966) by
W. F. Blankley. The species was isolated by means of a
Pasteur pipette into axenic culture and was generously placed
at my disposal for study. A clone was made from the axenic
culture and was the subject of all following observations.
Since isolation the culture has been maintained axenically in
an enriched sea water medium diluted to 75 per cent with
glass distilled water. The following enrichments were used
per liter of solution: KNO: 0.20 g, K2HPO,4:3H.0 0.046 g, soil
extract 15 mls (the solution of 1 part soil to 1 part distilled
water by weight is autoclaved and the supernatant used), P II
metals (Provasoli, 1964) 30 mls, vitamin By. 1 pg, thiamin
hydrochloride 1 mg, and biotin 2 »g. The clone is maintained
in a north window at room temperature (22—25°C).
Because the tabulation of this species differs from all pre-
viously described forms, a new genus and species is here
proposed. A culture of this alga (my strain no. 87; Blankley’s
original culture ) has been deposited at the Culture Collection
of Algae, Indiana University, Bloomington, Indiana, and has
been given the accession number IU No. 1564.
11—Proc. Brot. Soc. WaAsu., Vou. 81, 1968 (91)
92 Proceedings of the Biological Society of Washington
Fics. 1-3. Cachonina niei gen. et sp. nov. 1, Acetocarmine stained
cells showing the posterior position of the nucleus, the left one in end
view, the right in lateral view. Approx. X1200. 2 and 3, Chromosome
squash, drawing and photomicrograph at the same magnification as
Fig. 1.
Division PYRRHOPHYTA Pascher, 1914
OrvER PERIDINIALES Haeckel, 1894
FamiLy Peridiniaceae Diesing, 1850
Cachonina niei gen. et sp. nov.
Cellular morphology: The cell consists of a theca with a pellicle under-
neath. The pellicle at times can be seen detached from the theca in the
apical and cingular regions. This pellicle is present in addition to a
cytoplasmic membrane. All measurements have been made on iodine
(Lugol’s solution) fixed cells. The elongate, dorsal-ventrally flattened
cell has an average longitudinal dimension of 18 » (range 17-20 yz),
an average lateral dimension of 11.5 w (range 11-12 »), and an average
ratio of length to width of 1.61 (range 1.53-1.71). The spherical
nucleus, which is 6 « in diameter, remains posterior to the cingulum
(Fig. 1). From acetocarmine stained nuclei two chromosome counts
of 111, 112 were obtained (Figs. 2, 3); the chromosomes are short rods
of uniform size. Transversely and longitudinally directed flagella are
present.
The numerous yellow-brown plastids are parietally arranged. No
stigma is present, although stationary phase cells develop several parietal
orange colored bodies that presumably are degenerate plastids. The
cytoplasm of cells grown in high light intensity contains numerous
“starch” grains. No bioluminescence occurs.
Thecal morphology: The cell wall of Cachonina niei is very thin, and
has poorly marked sutures. In order to determine the tabulation, material
was fixed and stained with Lugol’s solution (Iz, KI); the wall was stained
brown and the sutures were revealed. Cells of this species may undergo
New Marine Dinoflagellate hs
ecdysis, thus leaving behind an empty theca. When stained with iodine,
these empty thecae were exceedingly useful for determination of the
plate pattern. The cell leaves the theca through the anterior intercalary
region and consequently, entire epithecae rarely were found. It is this
same anterior dorsal region in peridinoid cysts through which excystment
occurs (Evitt, 1967). Although the arrangement of the plates in the
epitheca was difficult to determine, a few free, intact epithecae were
observed that did reveal clearly the tabulation. In these free epithecae
the cingulum was never attached, indicating that the sutures between
the epitheca and cingulum are weaker than those between the cingulum
and the hypotheca. Figs. 4-7 show the arrangement of the thecal plates.
Plate 1’ is in contact only with 8” and 1” of the precingular series
giving an ortho arrangement (Fig. 4). Plate 2a contacts only plates 3”
and 4” of the precingular series, thus having a penta arrangement. In
contrast to members of Peridinium there is one extra plate in both the
apical series and the precingular series in Cachonina niei. The six plates
of the cingulum are all of approximately equal size. Of the four sulcal
plates, the posterior plate is largest. The left sulcal plate is narrow;
there is a small anterior plate above the left sulcal plate. The right
sulcal plate extends into the cingulum. Between the anterior and right
sulcal plates there is a ridged suture through which the flagella penetrate
the theca. The hypotheca has the normal arrangement for the Peridin-
iaceae. The plate tabulation is: pore plate, 5’, 3a, 8”, 6c, a.s., r.s., ls.,
DeSeossee oiae
Latin diagnosis: Cellula dorsali-ventraliter applanata, cingulum sinis-
traliter descendens, per dimidium latitudinis dispositum, sine collaribus.
Cormue et ornatus superficiales nulla. Cellula 18 w long., 11.5 w lat.;
ratio longitudinis et latitudinis 1.61. Plastides parietales multae. Nucleus
post cingulum; numerous chromosomatum 112. Theca tenuis. Suturae
non valde manifestae. Tabulatio thecalis: lamella pori apicalis, 5’, 3a,
8”, 6c, a.s., r.s., ls., p.s., 5/’, 2””. Habitatio: marina; cellulae in loco
Salton Sea, California dicto repertae.
Typus: Cachonina niei sp. nov.
Holotypus: Fig. 7.
Discussion: The two antapical and five postcingular plates of the
hypotheca place this organism in the family Peridiniaceae. Cachonina
niet has an ortho, penta arrangement on the epitheca similar to that of
members of the section Tabulata Jorgensen, 1912, of the genus Peridinium
Ehrenberg, 1830. The present form differs from species of Peridinium
in having five apical plates and eight precingular plates, as opposed to
four apical plates and seven precingular plates in Peridinium.
Much emphasis has been placed on the arrangement of the sulcal
plates and the number of cingular plates by Abé (1936) and Balech
(1959, 1963), but this information is lacking for the type species of
the genus Peridinium: P. cinctum (Miiller) Ehrenberg, 1830, a fresh-
water species. However, other freshwater species related to P. cinctum
94 Proceedings of the Biological Society of Washington
LSP ERS
Fics. 4-7. Cachonina niei gen. et sp. nov. 4, Apical view of the
tabulation. 5, Posterior view of the tabulation. 6, Dorsal view of the
tabulation. 7, Holotype, ventral view of the tabulation. Plate explana-
tion: p.p., pore plate; I’—5’, apical plates; Ja—3a, anterior intercalary
plates; 1”-8”, precingular plates; Ic—6c, cingular plates; a.s., anterior
sulcal plate; p.s., posterior sulcal plate; r.s., right sulcal plate; L.s., left
sulcal plate; 1”’-5”’, postcingular plates, 1””, 2””, antapical plates.
possess five cingular plates and lack an apical pore. All marine species
referred to the genus Peridinium that have been examined have only
three cingular plates plus a transitional plate, but do possess an apical
pore. Thus it appears that the genus Peridinium should be divided into
New Marine Dinoflagellate 95
two genera: (a) Peridinium, consisting of freshwater species resembling
the type species P. cinctum and (b) the marine species.
The marine species previously assigned to Peridinium whose tabulation
has been investigated logically belong to a genus whose type species
possesses an apical pore, three cingular plates and a transitional plate.
Among the genera considered to be synonymous with Peridinium, the
genus Archaeperidinium Jérgensen, 1912, is the earliest described whose
type species has the above three characters. Archaeperidinium minutum
(Kofoid) Jorgensen, 1912, is the type species of Archaeperidinium;
Balech (1964) has given a detailed analysis of the thecal plates of this
species. The type of Archaeperidinium has previously been placed in
the section Tabulata Jorgensen, 1912, of the genus Peridinium. The
genus Peridinium should be restricted to those non-pored species having
the same cingular tabulation as the type species of Peridinium. Archae-
peridinium is therefore the logical and nomenclaturally correct genus
for the marine, pored species with three cingular plates and a transitional
plate previously placed in Peridinium.
Scrippsiella Balech ex Loeblich, 1965 (Balech, 1959), has been estab-
lished for marine, apical pored species with six cingular plates but with
otherwise normal Peridinium epitheca and hypotheca. Cachonina has six
cingular plates in common with Scrippsiella but has a different number
of precingular and apical plates. Cachonina differs from the marine
Peridinium species (now referable to Archaeperidinium ) in the possession
of six cingular plates, but resembles it in the possession of an apical
pore. Archaeperidinium, Peridinium, and Scrippsiella all have seven
precingular and four apical plates, while Cachonina has eight precingular
and five apical plates.
Etymology: The generic and specific names are patronymics in honor
of Drs. Jean Cachon and Dashu Nie who have contributed greatly to
our knowledge of marine dinoflagellates. The gender is feminine.
Acknowledgments: This research was supported in part by Training
Grant GM 1065 from the National Institutes of Health, U.S. Public
Health Service. I thank Mr. W. F. Blankley, Scripps Institution of
Oceanography, La Jolla, California, for supplying the original subculture
of this organism. Drs. W. S. Drugg and A. R. Loeblich, Jr., Chevron
Research Corp., La Habra, California, kindly discussed the problem and
aided with the illustrations. Thanks are due to Laurel Loeblich and
Professor Helen Tappan Loeblich for critically reading the manuscript.
Special thanks are due to Professor B. E. Volcani, Scripps Institution of
Oceanography, La Jolla, California, for discussions of the subject and
for a critical reading of the manuscript.
LITERATURE CITED
Ase, T. H. 1936. Report of the biological survey of Mutsu Bay. 29.
Notes on the protozoan fauna of Mutsu Bay. II. Genus
Peridinium: Subgenus Archaeperidinium. Sci. Repts. Tohoku
Imperial Univ., ser. 4, 10(4): 639-686, text-figs. 1-102.
96 Proceedings of the Biological Society of Washington
BaLecH, Enrique. 1959. Two new genera of dinoflagellates from
California. Biol. Bull., Woods Hole, 116(2): 195-203, text-
figs. 1-3.
1963. Dos dinoflagelados de una laguna salobre de la
Argentina. Ministerio Educacion Nac., Univ. Nac. Plata,
Fac. Cienc. Nat. Mus., Notas Museo, Zool. no. 199, 20:
111-123, text-figs. 1-10.
1964. Tercera contribucion al conocimento del genero “Peri-
dinium”. Rev. Museo Argentino Cienc. Nat. “Bernardino
Rivadavia’, Inst. Nac. Inv. Cienc. Nat., Hidrobiol., 1(6):
179-195, pls. 1-3.
Evitt, W. R. 1967. Dinoflagellate studies. II. The archeopyle. Stan-
ford Univ. Publ. Geol. Sci., 10(3): 1-82, text-figs. 1-50, pls.
1-11.
JORGENSEN, EucEen. 1912. Bericht tuber die von der schwedischen
Hydrographisch-Biologischen Kommission in den schwedis-
chen Gewidssern in den Jahren 1909-1910 eingesammelten
Planktonproben. Svenska Hydro.-Biol. Komm. Skr., 4: 1-20.
LoepuicH, A. R., III. Dinoflagellate nomenclature. Taxon 14(1):
15-18.
Provasout, Luict. 1964. Growing marine seaweeds. Proc. Internat.
Seaweed Symp., 4: 9-17, appendix 1.
74. O67 3 on
Vol. 81, pp. 97-100 April 30, 1968
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
A NEW MOUSE OF THE GENUS NELSONIA
FROM SOUTHERN JALISCO, MEXICO
Hucu H. GENoways AND J. KNOx JONES, Jr.
In October 1966 and again in February 1967, Percy L.
Clifton of the Museum of Natural History, The University of
Kansas, collected terrestrial vertebrates on El Nevado de
Colima in the southern part of the Mexican state of Jalisco.
Among the mammals obtained were seven specimens of the
unique woodrat, Nelsonia neotomodon, which appear to rep-
resent a new subspecies that is named and described below.
Nelsonia neotomodon cliftoni new subspecies
Holotype: Adult female, skin and skull, no. 109,437 Museum of
Natural History, The University of Kansas; from 2.5 mi. ENE Jazmin,
6,800 ft, Jalisco; obtained on 20 October 1966 by Percy L. Clifton,
original no. 11,706.
Geographic distribution: Known only from the type locality and from
a place 4 mi. ENE Jazmin on the northwest slope of El Nevado de
Colima; probably also occurs on nearby Volcén de Fuego in Jalisco and
adjacent Colima.
Measurements: External and cranial measurements (in millimeters )
of the holotype, followed by those of another adult female, two young
adult females, and two young adult males, are, respectively: total length
251, 256, 234, 231, 238, 235; length of tail 126, 123, 110, 111, 111, 115;
length of hind foot 29, 29, 27, 27, 28.5, 27.5; length of ear 24, 24.5, 23,
23, 23, 23.5; greatest length of skull 32.8, 33.9, 32.0, 31.5, 31.8, 33.0;
zygomatic breadth 18.3, 18.6, 17.1, 17.1, 17.1, 17.9; mastoid breadth
13.5, 14.4, 13.4, 13.6, 13.3, 13.4; interorbital constriction 4.7, 4.7, 4.8,
4.8, 4.6, 4.9; rostral length 13.3, 13.8, 12.9, 12.6, 12.7, 13.2; rostral
breadth 5.5, 5.9, 4.8, 5.1, 5.0, 5.1; length of maxillary toothrow 6.5, 6.9,
6.3, 6.3, 6.6, 6.6; depth of skull 10.7, 11.2, 10.5, 10.6, 10.1, 11.0;
breadth of zygomatic plate 3.5, 3.5, 3.2, 3.4, 3.3, 3.3.
Comparisons: From WNelsonia neotomodon neotomodon (specimens
examined from Zacatecas and northern Jalisco), N. n. cliftoni differs in
many of the same ways—but not always to the same degree—as does
N. n. goldmani (see Hooper, 1954: 7-8, and Merriam, 1903: 80). The
most conspicuous ways in which cliftoni differs from neotomodon are in
12—Proc. Biot. Soc. WaAsH., VoL. 81, 1968 (97)
98 Proceedings of the Biological Society of Washington
50
KILOMETERS ~*
Fic. 1. Geographic distribution of Nelsonia neotomodon: (1) N. n.
neotomodon; (2) N. n. goldmani; (3) N. n. cliftoni. Localities of occur-
rence plotted for the subspecies neotomodon and goldmani are from
Hooper (1954: 6) and Baker and Greer (1962: 129).
being darker dorsally, in lacking a white tip on the tail, and in having
dusky (rather than whitish) hind feet, broad zygomatic plates with
correspondingly deep zygomatic notches, and larger auditory bullae.
Also, cliftoni averages larger than neotomodon both externally and
cranially.
From Nelsonia neotomodon goldmani of Cerro de Tancitaro and Cerro
Patamban, Michoacan, N. n. cliftoni differs principally in having a flatter,
less inflated braincase (and consequently a shallower skull reminis-
cent of the subspecies neotomodon), auditory bullae that are more
New Mexican Mouse 99
laterally directed posteriorly, and noticeably paler dorsal and _ lateral
coloration (October-taken cliftoni compared with the February-taken
holotype and paratypes of goldmani). The head and back of cliftoni is
nearly intermediate in coloration between that found in the darker
neotomodon and the paler goldmani. Considering the sides and cheeks,
cliftoni can be distinguished from goldmani in having a paler, more
ochraceous color to the pelage that is not sa buffy as in neotomodon.
In conventional measurements of the cranium, judging by the limited
material available for comparison, cliftoni averages larger than goldmani,
especially in greatest length of skull, zygomatic breadth, and _ rostral
length (12.2-12.6 in three goldmani, 12.6—13.8 in six cliftoni) and has
a shorter maxillary toothrow relative to the length of skull.
Remarks: We suspect that Nelsonia neotomodon cliftoni is restricted
geographically to the slopes of El Nevado de Colima and Volcan de
Fuego (Volc4n de Colima on some maps) in southern Jalisco and adja-
cent Colima. The range of cliftoni is isolated from that of neotomodon
by the interior basins of central Jalisco (see Fig. 1). From goldmani,
which occurs to the east in the high mountains of Michoacan, cliftoni
may be isolated by the basins of the Rio Tepalcatepec and Rio Tuxpan
and their tributaries and also in part by the broad valley containing the
playas of Zapotlan and Sayula. It is conceivable that intermittent gene
flow between goldmani and cliftoni has been maintained across the Sierra
del Tigre and associated pine-capped highlands south of Lago de Chapala
but this seems doubtful. Several field parties have collected in this area
in recent years and have not taken Nelsonia, although it is noteworthy
that the highest places in the Sierra del Tigre approximate the lowest
elevation (6,800 ft) at which cliftoni has been taken along the road
from Ciudad Guzman to Jazmin.
Our six specimens of cliftoni from the type locality were obtained on
19-21 October 1966 in cool, pine-oak forest. The area had been recently
logged and contained scattered cornfields. Nelsonia was taken in traps
baited with rolled oats that were set in various places along a brushy,
overhanging bank adjacent to a road. One was caught under a rock;
others were trapped in trails along the bank and under a woodpile.
Other species of rodents taken at the same place included Liomys irro-
ratus jaliscensis (J. A. Allen), Peromyscus boylii levipes Merriam, Pe-
romyscus hylocetes hylocetes Merriam, Peromyscus maniculatus labecula
Elliot, and Neotoma mexicana tenuicauda Merriam. A juvenile male
taken 4 mi. ENE Jazmin on 8 February 1967 was trapped in dense weeds
at the edge of a cornfield along with Reithrodontomys sumichrasti nert-
erus Merriam, Peromyscus hylocetes hylocetes, and Peromyscus mani-
culatus labecula. None of the four October-taken females (two adults,
two young adults) was reproductively active.
We take pleasure in naming the new subspecies for Percy L. Clifton,
field representative of the Museum of Natural History from 1961 to 1967,
whose collections of vertebrates from Jalisco and elsewhere in Mexico
have contributed materially to the research efforts of many scientists.
100 Proceedings of the Biological Society of Washington
Specimens examined: JALISCO: 2% mi. ENE Jazmin, 6,800 ft, 6
(KU 109434-38, 109440); 4 mi. ENE Jazmin, 7,700 ft, 1 (KU 111953).
Acknowledgments: Funds from the Kansas University Endowment
Association defrayed the cost of field work and a contract (DA—49-193-
MD-2215) from the U.S. Army Medical Research and Development
Command supported laboratory studies. Comparative material was made
available at the U.S. National Museum by Charles O. Handley, Jr.
LITERATURE CITED
Baker, R. H., AND J. K. GREER. 1962. Mammals of the Mexican state
of Durango. Publ. Mus. Michigan State Univ., Biol. Ser.,
9: 25-154, 4 pls., 6 Figs.
Hooper, E. T. 1954. A synopsis of the cricetine rodent genus Nelsonia.
Occas. Papers Mus. Zool., Univ. Michigan, 558: 1-12, 1 map,
1 Fig.
Merriam, C. H. 1903. Four new mammals, including a new genus
(Teonopus), from Mexico. Proc. Biol. Soc. Washington, 16:
79-82.
Ue OFS
Vol. 81, pp. 101-108 "April 30, 1968
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
ALGANSEA MONTICOLA, A NEW CYPRINID FISH
FROM THE PACIFIC SLOPE OF CENTRAL MEXICO
By Ciypr D. Barsour AND SALVADOR CONTRERAS
Dept. of Zoology and Entomology, University of Utah,
Salt Lake City, Utah 84112
Escuela de Ciencias Biologicas, Universidad de Nuevo Leon
Monterrey, Nuevo Leon
During the summer of 1963, eight series totaling 630 speci-
mens of an undescribed cyprinid were taken by us from the
headwaters of the Rio Colotlan and the Rio Juchipila, northern
tributaries to the Rio Grande de Santiago on the Pacific slope
of the states of Zacatecas and Jalisco. Subsequent study re-
vealed it to be a member of the genus Algansea, a small group
widely distributed in the Lerma-Santiago river system and
contiguous waters.
Algansea monticola new species
(Figs. 1-3)
Holotype: Tulane University no. 40869, a male 69.9 mm in standard
length, collected from the Rio Juchipila in Zacatecas near the town of
Jalpa at the Highway 70 bridge about one mile east of the intersection
with Highway 41, 30 June 1963, by Clyde D. Barbour and Salvador
Contreras.
Paratypes: Tulane University no. 30611, 62 specimens bearing the
same data as the holotype, now distributed as follows: British Museum
of Natural History, no. 1967. 10. 3. 1-2, 2 specimens; Facultad de
Ciencias Bioldgicas, Universidad de Nuevo Leén, Monterrey, N. L.,
México, no. FCB 1015, 2 specimens; Field Museum of Natural History,
no. 74073, 2 specimens; Instituto Nacional de Investigaciones Biolégico
Pesqueras, Laboratorio Biolégico, México, D. F., no. LBT 235, 2 speci-
mens; Tulane University, 49 specimens; University of Michigan, Museum
of Zoology, no. 184359, 3 specimens; United States National Museum,
no. 202161, 2 specimens.
The following collections from the State of Zacatecas are also desig-
nated as paratypes: Rio Juchipila at Mal Paso 16 mi. E Ciudad Garcia
Salinas, TU 30608 (81 specimens) including UMMZ 184356 (5), 28
13—Proc. Bion. Soc. WAsH., Vou. 81, 1968 (101)
‘Ysa, prepurys ur WU 6'Gg ‘ope ‘odAjoToYy “E980F NL “7joouow pasups]Y TOI
102 Proceedings of the Biological Society of Washington
New Cyprinid Fish from Mexico 103
June 1963. Tributary to Rio Juchipila 5 mi. S Villanueva on Hwy. 41,
TU 30609 (141 specimens) including UMMZ 184357 (10), 28 June
1963. Tributary to Rio Juchipila at Tabasco, TU 30610 (21) including
UMMZ 184358 (3), 29 June 1963.
Other specimens: Zacatecas. Reservoir behind dam across tributary
to Rio Colotlan 1.5-2 mi. S Tepechitlan, TU 30604 (1), 26-27 June
1963. Tributary to Rio Colotlan 2 mi. N Momax, TU 30605 (51), 27
June 1963. Tributary to Rio Colotlan 2 mi. S Tepetongo, in small pool
at foot of dam, TU 30607 (13), 28 June 1963. Jalisco. Rio Colotlan at
Santa Maria de los Angeles, TU 30606 (259) including UMMZ 184354
(15), 27 June 1963.
Diagnosis: A small species (maximum length about 90 mm S. L.)
which can be distinguished from all other known members of its genus
by characteristically having 7 rather than 8 or more rays in the dorsal
fin.
Description: Body robust, caudal peduncle relatively deep; snout
blunt, not overhanging upper lip; mouth oblique, included; maxillary
barbels rarely absent; eyes small; fins moderate, fleshy; scales small,
imbedded in small specimens; 6 to 11 gill rakers on first arch; pharyngeal
teeth 4-4.
Morphometric data for the holotype and four paratypes are given in
table 1. Combined counts for specimens from the Rio Juchipila and the
Rio Colotlan are as follows (the value including the holotype is italic-
ized): median lateral scales (206 specimens), 56 (in 2), 57 (3), 58 (5),
59M) 60 (4M) G15 (14), 62/119) 635(29)), eae 3), 65 (19), 66 (23),
Gy (ID), Gs} ls}, G2) CL), @@ (24), Wa (3), 7 (Bye Hee rays (220
specimens), 11 (1), 12 (0), 13 (1), 14 (2), 1 5 (42).1 6 (113), 17 (56),
18 (5); dorsal rays (234 specimens), 7 (213), 8 (21); anal rays (230
_ specimens ), 6 (8), 7 (217), 8 (5); total number of gill ses on the first
arch (234 specimens), 6 (9), 7 (70), 8 (77), 9 (64), 10 (12), 11 (2);
total number of vertebrae including the four fused tee and count-
ing the urostyle as one (77 specimens), 33 (3), 34 (1), 35 (6), 36
(40), 37 (25), 38 (2). The number of pores entering the cephalic
lateral line system were counted in 40 specimens as follows: supra-
temporal pores 4 (1), 5 (7), 6 (6), 7 (10), 8 (11), 9 (5); supraorbital
pores 7 (2), 8 (19), 9 (10), 10 (9) entering the left canal, 7 (4), 8
(19), 9 (13), 10 (4) entering the right canal; infraorbital pores 14 (2),
ls (7), (LD), We CL), WS (7), We) Cl) eke exael wl (aL), Ie (a), ils
(4), 16 (9), 17 (13), 18 (7), 19 (2), 20 (3) right; preoperculomandib-
wlenp jororress @) (@))s IO) (UL), 700 (1G), 1 (Sig a) (is), EOL) Wee eave
8 (2), 9 (3), 10 (8), 11 (19), 12 (4), 13 (4) right. Occasionally canals
were found to be disfigured; most commonly the supratemporal was
divided at the midline into separate left and right halves.
Color: Color variable but always dull. Dorsally, A. monticola is dusky
grey to grey-brown. A dark, narrow stripe extends along the middorsal
line from the nape to the caudal fin. Laterally, along the side of the
body but generally dorsal to the lateral line, a diffuse rust-orange band
104 Proceedings of the Biological Society of Washington
230
ie)
nN
(eo)
210
ie)
(o)
(e)
190
180
170
160
150
LENGTH OF PECTORAL FIN IN THOUSANDTHS OF THE STANDARD LENGTH
40 50 60 70 80
STANDARD LENGTH IN MM
Fic. 2. Sexual dimorphism in pectoral fin length of Algansea monti-
cola. The open circles represent males, the closed circles females.
New Cyprinid Fish from Mexico 105
Fic. 3. Known distribution of Algansea monticola. 1. Rio Grande de
Santiago; 2. Rio Colotlan; 3. Rio Juchipila; 4. Rio Verde. The closed
circles represent collection sites; the open circle the location of the town
of Moyahua de Estrada.
may be present between the dorsal margin of the operculum and the
midbase of the caudal fin. A small black spot is always present at the
latter point. The spot fades in alcohol and may be obscure in large,
preserved specimens. The ventral coloration is dirty white.
Etymology: The name monticola (Latin) means “a dweller in the
mountains” and alludes to the rugged nature of the area in which the
species occurs.
Sexual dimorphism: Although differences between the sexes have not
been reported for this genus, we found that breeding males had notice-
ably longer pectoral fins than did the females (fig. 2). Also, many of
the males had well developed breeding tubercles. These were minute,
white protuberances scattered over the top of the head and body, the
dorsal surface of the pectoral fins and on the dorsal and anal fins. No
differences in coloration were observed.
Ecology: Algansea monticola is ubiquitous in those parts of the head-
waters of the Rio Juchipila and Rio Colotlan sampled. The species was
taken over mud, sand, gravel and rocks, in open water and under banks,
in swift water and in pools and impoundments. The water temperature
106 Proceedings of the Biological Society of Washington
varied between 19°C and 27°C. The species was not associated with
any particular type of vegetation.
The fact that larger females carried large eggs and that testes and
tubercles were well developed in males suggests that spawning takes
place in late June and early July. The rust-orange coloration was
probably a reflection of spawning activity, although no collections have
been made at other times of the year. Turbid water precluded any
behavior observations.
Associated with Algansea monticola was a sucker, Moxostoma con-
gestum (Baird and Girard), a chub, Hybopsis alta (Jordan), a poeciliid,
Poeciliopsis infans (Woolman), a goodeid, Goodea sp., and two centrar-
chids, Chaenobryttus gulosus (Cuvier) and Micropterus salmoides (Lacé-
pede). The latter two species were taken in the reservoir south of
Tepechitlan and are introduced. No catfish were taken although they
were probably present.
Relationship: Algansea monticola is most similar to A. barbata ( Alvarez
and Cortés, 1964), a form found in the headwaters of the Rio Lerma,
and an undescribed species from Nayarit (R. R. Miller, ms.). All three
have barbels, few gill rakers and are small. It is probable, however,
that each evolved independently from a common stock and that the
similarities reflect adaptations to like environments. The eleven nominal
species of Algansea and A. monticola are currently under study by the
senior author and Robert R. Miller of the University of Michigan.
Distribution: Algansea monticola has been collected from the head-
waters of the Rio Juchipila and the Rio Colotlan (fig. 3). A collection
was made on 30 June 1963 in the Rio Juchipila at Moyahua de Estrada,
Zacatecas, at an elevation slightly below 1,200 meters, yielded only
Moxostoma congestum (Baird and Girard), Cyprinus carpio Linnaeus,
Hybopsis alta (Jordan), a catfish, Ictalurus pricei (Rutter), and a cichlid,
Cichlasoma sp., suggesting that the new species is absent from the lower
reaches of this river. Briggs and Miller (1960) made no mention of it
occurring in the Rio Grande de Santiago. Algansea Monticola has not
been taken in the Rio Verde to the east; its limits of distribution in the
tributaries to the west of the Rio Juchipila remain unknown.
Acknowledgments: Drs. R. R. Miller and R. D. Suttkus provided
much advice and critically read the manuscript. Dr. Miller also kindly
allowed us to examine specimens of his undescribed species of Algansea.
Dr. E. C. Raney provided helpful comments during the early stages of
this study. Dr. Kenneth Relyea and Mr. Glenn Clemmer, Tulane Univer-
sity, took the radiographs from which the vertebral counts were made.
(The former is now at the University of Jacksonville, Florida). The
cooperation of the Mexican government in providing a collecting permit
is also gratefully acknowledged.
Field work during the summer of 1963 and other travel was partially
financed by a grant from the National Institutes of Health (3—-T1-ES-
27-01S1) to Suttkus, a grant-in-aid from the Society of the Sigma
Xi-RESA (1963) and a grant from the University of Utah Research
New Cyprinid Fish from Mexico 107
TaBLE 1. Measurements! of Algansea monticola expressed
as thousandths of the standard length.
Paratypes
Holotype TU 30611
TU 40869
Sex of Q of fo) 2 Mean
Standard length (mm) 69.9 68.3 62.5 54.6 60.6 63.2
Head length Di2 265 250 269 262 264
Postorbital head length 140 142 130 150 140 140
Orbit length 53 47 47 53 53 50
Snout length 84 76 ria ah 79 79
Upper jaw length 2: 70 67 71 74 71
Fleshy interorbital width 100 92 91 103 96 96
Head width 179 179 150 158 167 167
Snout to dorsal origin 539 556 526 555 563 548
Snout to pelvic origin 559 548 531 548 559 548
Snout to anal origin 721 TAL 711 723 728 125
Greatest body lengih 273 268 251 267 282 268
Depth at occiput 183 179 171 183 178 179
Greatest body width 173 209 155 179 196 182
Caudal peduncle depth 136 124 136 132 134 132
Caudal peduncle length 216 214 206 216 218 214
Dorsal base 107 95 115 95 94 101
Anal base 86 75 93 88 83 85
Depressed dorsal length 225 187 227 211 198 210
Depressed anal length 176 143 192 172 162 169
Pectoral length 207 154 208 161 162 178
Pelvic length 143 122, 158 132 130 137
1 As described in Hubbs and Lagler (1947).
Fund (1967) to Barbour and a grant from the Banco de México (1963)
to Contreras.
LITERATURE CITED
ALVAREZ, J. AND M. T. Cortés. 1964. Una especie de Algansea cap-
turada en el Alto Lerma (Pisc., Cyprin.). Ciencia, Mex.
23(3): 104-106.
Briccs, JoHN C. AND Ropert RusH Mitier. 1960. Two new fresh-
water clingfishes of the genus Gobiesox from southern
Mexico. Occ. Pap. Mus. Zool. Univ. Mich. 616: 1-15.
Husss, Cart L. anp Karu F. Lacier. 1947. Fishes of the Great
Lakes Region. Bull. Cranbrook Inst. Sci. 26: 1-186.
108 Proceedings of the Biological Society of Washington
FH O73 ae
Vol. 81, pp. 109-110 April 30, 1968
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
A NEW SPECIES OF PARAFELICOLA
(MALLOPHAGA: TRICHODECTIDAE)
FROM MOZAMBIQUE
By K. C. EMERSON AND Rocer D. PRICE
Arlington, Virginia and Department of Entomology,
Fisheries, and Wildlife, University of Minnesota,
St. Paul, Minnesota
In 1966, the authors described Parafelicola africanus and
reviewed the characters of each species in the genus. Since
that time a series representing a sixth, and new, species has
been received. It is herewith described and illustrated.
Parafelicola neoafricanus new species
Holotype male: Total length, 1.22 mm. External morphology and
chaetotaxy as shown in Fig. 3. Genitalia, less sac, when contained within
the abdomen as shown in Fig. 2. Genital sac large and with numerous
small serrations.
Allotype female: Total length, 1.29 mm. External morphology and
chaetotaxy as shown in Fig. 1. Internal chamber of vulva prominent
and serrated.
Discussion: P. neoafricanus is without noticeable respiratory spiracles,
a feature shared only with P. acuticeps and P. africanus. The basal plate
of the male genitalia is shorter and wider in P. neoafricanus than in P.
africanus and P. acuticeps. The pseudopenis and the paramera of P.
neoafricanus are much thicker than those found in P. africanus and P.
acuticeps. The chaetotaxy of the terminal abdominal segment of the
male is more dense in P. africanus than in P. neoafricanus. The female
of P. neoafricanus has a tergal plate on all abdominal segments, the
female of P. africanus has a tergal plate only on segments I-V, and
P. acuticeps has a tergal plate on segments I—-VII. In our description of
P. africanus, it was incorrectly stated that the subgenital lobes were
without serrations. The subgenital lobes of: P. acuticeps are pointed,
without serrations; P. africanus are bluntly pointed, with inside and
outside serrations; and P. neéoafricanus are bluntly pointed, with inside
serrations. P. neoafricanus is larger, in both sexes, than P. africanus.
Type host: Genetta tigrina (Schreber, 1778).
14—Proc. Bion. Soc. WAsH., VoL. 81, 1968 (109)
110 Proceedings of the Biological Society of Washington
Fics. 1-3. Parafelicola neoafricanus, new species. 1, dorsal-ventral
view of female. 2, male genitalia. 3, dorsal-ventral view of male.
Type material: Holotype male, allotype female and nine paratypes
collected off the type host at Changara, Tete District, Mozambique on
4 November 1964 by H. J. Herbert (HJH-1019). The holotype and
allotype will be deposited in the U.S. National Museum.
LITERATURE CITED
EMERSON, K. C. aND R. D. Prick. 1966. A new species of Parafelicola
(Mallophaga: Trichodectidae) from the Small-spotted Genet.
Proc. Biol. Soc. Wash., 79: 231-233.
"ZL WOZKS:
Vol. 81, pp. 111-122 April 30, 1968
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
THE INDO-PACIFIC BLENNIID FISH GENUS
STANULUS, WITH DESCRIPTION OF A NEW
SPECIES FROM THE GREAT BARRIER REEF
(BLENNIIDAE; BLENNIINAE; SALARIINT)
By Vicror G. SPRINGER
Division of Fishes, U.S. National Museum,
Washington, D.C. 20560
This report is the result of an attempt to place taxonomically
an undescribed species of blenny that I collected at One Tree
Island, Capricorn Group, Great Barrier Reef, Australia. A
large series of the undescribed species was collected, as well
as a single specimen of a very similar species. The single
specimen was identifiable as either Fallacirripectes minutus
Schultz and Chapman (type-species of Fallacirripectes Schultz
and Chapman, 1960) or Stanulus seychellensis Smith (type-
species of Stanulus Smith, 1959). After examination of the
literature and additional material I decided that the new
species is congeneric with the single specimen, but there
remained the problem as to the exact identity of the single
specimen. My examination of the holotypes of F. minutus and
S. seychellensis has convinced me that the two species (gen-
era) are synonymous, as I had previously opined (Springer,
1967) based only on the literature. Fallacirripectes, then, is
a junior synonym of Stanulus. Schultz and Chapman (1960)
failed to compare Fallacirripectes with Stanulus in their orig-
inal description.
In addition, I find that Fallacirripectes wellsi Schultz and
Chapman (1960), is an ophioblennius larval stage of S. sey-
chellensis. The meristics and distribution of cirri found in
the holotype, and only known specimen, of F. wellsi are the
same as in S. seychellensis. The broad interorbital area relative
to other body parts, and the darker, pigmented pectoral fins
15—Proc. Bron. Soc. WasH., Vou. 81, 1968 (@ialsis)
112 Proceedings of the Biological Society of Washington
used by Schultz and Chapman to differentiate F. wellsi, are
larval characteristics I have noticed in the larval stages of
other blenniids.
The following abbreviations are used in this paper: CAS—
California Academy of Sciences, LACM—Los Angeles County
Museum, SI—standard length, USNM—wUnited States Na-
tional Museum.
Stanulus Smith
Stanulus Smith, 1959, Rhodes Univ. Ichthyol. Bull. 14, p. 246 (type-
species: S. seychellensis Smith, 1959, by original designation and
monotypy ).
Fallacirripectes Schultz and Chapman, 1960, U.S. Nat. Mus. Bull. 202,
pt. 2, p. 362 (type-species: F. minutus Schultz and Chapman, 1960,
by original designation ).
Description: Small species (to 39 mm SL) of tribe Salariini (Springer,
in press), with short simple cirri on each side of nape and on posterior
rim of anterior nostril; short simple supraorbital cirrus present (S.
talboti) or absent (S. seychellensis) above each eye; upper lip entire;
some pores in infraorbital series in pairs or groups; one pore in front
of each anterior nostril; preoperculomandibular series of pores simple;
predorsal commissural pores (see Springer, 1967) 5 or more in specimens
over 12 mm SL; lateral line continuous to point below or beyond posterior
end of spinous dorsal, continuing thence as short, separate, bipored
tubes to point below or anterior to mid-soft dorsal fin; imbricate scalelike
flaps variably present in anterior portion of lateral line. Dorsal fin spines
12 (14 in one specimen of S. talboti); last spine greatly reduced and
usually visible only in cleared and stained preparations or radiographs;
segmented dorsal fin rays 9 to 12; terminal dorsal ray bound by mem-
brane to caudal peduncle; dorsal fin deeply incised between spinous
and rayed portions. Anal fin spines 2; anal fin rays 10 to 13; last anal
ray split to base (here counted as one ray), supported by single proximal
pterygiophore (see Springer, 1967 for discussion of terminal anal ray
of blenniids); terminal anal ray free from caudal peduncle. Vertebrae
10 + 18 to 21 = 28 to 31; first neural spine on third vertebrae;
epipleurals on Ist to 11th through 14th vertebrae; pleurals on vertebrae
3 to 10 (rarely 11); parapophysial stays on vertebrae 8 to 10. Pectoral
rays 14 to 16 (15 in over 90 per cent of fins, both left and right sides);
pectoral radial formula (Springer, in press) 2-1-1 (based on 3 specimens
of S. seychellensis) or 2-0-2 (based on 2 specimens of S. talboti). Post-
cleithra 2 on each side. Pelvic fin rays I, 4. Caudal fin with 13 seg-
mented rays, middle 9 each branched once; upper procurrent rays 6 to
10, lower, 5 to 10; terminal vertebra with two epurals, hypural 5
(Nybelin, 1963) autogenous, hypurals 3 and 4 fused to urostyle, hypurals
1 and 2 fused together, but free from urostyle. Last haemal spine fused
to centrum of penultimate vertebrae. Gill-rakers 8 to 16. Pseudo-
Indo-Pacific Blenniids of the Genus Stanulus 113
branchial filaments 4 to 7. Lateral extrascapulars not fused to skull.
Circumorbitals 5. Basisphenoid present, complete (see Springer, in
press). Kinethmoid (Springer, in press) present in one of two cleared
and stained specimens of S. talboti. Canine on each dentary posteriorly.
Premaxillary and dentary comblike teeth typically salariine (Springer, in
press), small, difficult to count, numbering about 110 in upper jaw and
80 in lower jaw. Conical teeth present (S. talboti) or absent (S. sey-
chellensis) on vomer.
Relationships: Stanulus is most closely related to Entomacrodus Gill,
1859. There is considerable overall similarity between the two species
of Stanulus, especially S. talboti, and the species of Entomacrodus. The
main differences between the two genera are meristic: the presence of
typically 15 pectoral rays in Stanulus, 14 in Entomacrodus; 12 dorsal
spines in Stanulus, 13 in Entomacrodus; 9 to 12 dorsal rays in Stanulus,
13 to 18 in Entomacrodus; 10 to 13 anal rays in Stanulus, 14 to 19 in
Entomacrodus; 28 to 31 vertebrae in Stanulus, 33 to 36 in Entomacrodus.
In addition, the scalelike flaps of the anterior portion of the lateral line
in Stanulus are not present in Entomacrodus. All species of Entoma-
crodus have supraorbital cirri and usually all specimens of any species
have vomerine teeth, whereas S. seychellensis lacks both supraorbital
cirri and vomerine teeth in all specimens.
Schultz and Chapman (1960) considered Stanulus (as Fallacirripectes )
close to Cirripectes. The absence of a comb of nuchal cirri, fewer
epipleurals (17-22 in Cirripectes) and the general nature of the color
pattern and physiognomy of Stanulus lead me to place Stanulus closer
to Entomacrodus than to Cirripectes.
Differentiation of the species of Stanulus
Supraorbital cirrus absent; vomerine teeth absent; dorsal segmented
rays 9 to 11 (11 in 1 of 51 specimens); anal segmented rays 10 to 12
(12 in 3 of 51 specimens); vertebrae 28 to 29; gill-rakers 12 to 16;
continuous portion of lateral line ending at point below segmented
GCLOns all Mle Tea y SA st OO: eaten Sia a S. seychellensis
Supraorbital cirrus present; vomerine teeth present; dorsal segmented
rays 11 or 12 (11 in 2 of 45 specimens); anal segmented rays 12 or
13 (12 in 2 of 45 specimens); vertebrae 30 or 31; gill-rakers
12 to 16; continuous portion of lateral line ending at point below
postemorsdorsalefiny spines) ee een ene S. talboti new species
Stanulus seychellensis Smith
Fig. 1, Table 1
Stanulus seychellensis Smith, 1959, Rhodes Univ. Ichthyol. Bull., no. 14,
pp. 246-247 (La Digue, Seychelles Ids. ).
Fallacirripectes minutus Schultz and Chapman, 1960, U.S. Nat. Mus.,
Bull. 202, vol. 2, pp. 362-365 (Eman Id., Bikini Atoll, Marshall Ids. ).
114 Proceedings of the Biological Society of Washington
jeue punome vory ‘spurysy [[eYysieyY “Yoyarag] Woz “TS UU 9'9G
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Indo-Pacific Blenniids of the Genus Stanulus 115
TaBLE 1. Frequency distributions for certain meristic
characters of two species of Stanulus.
Dorsal Rays Anal Rays Vertebrae
© @ alt a 10 11 12 13 28 29 30 31
S. seychellensis 49> 1 = Lay B.S 13 jie =
S. talboti By OAS: VN OAR ayy, OPO
Pseudobranchial
Gill-rakers Filaments
8 9 10 11 12 13 14 #15 16 4 5 6 7
S sqycelass OO 9 2d 8 US 5 5 Sal w=
S. talboti Se Si te Sr toe Ibe Sh Oho 3 5) 6 oe)
Fallacirripectes wellsi Schultz and Chapman, 1960, U.S. Nat. Mus., Bull.
202, vol. 2, pp. 365-366 (half mile off Rongelap Id., Rongelap Atoll,
Marshall Ids. ).
Description: Color pattern. The following description is based on a
mature male, 24.0 mm SL, from Guam, which exhibited what appears
to be the most complete color pattern of all the specimens I examined.
This description is followed by comments on notable variations en-
countered in other specimens.
Head and body of a pale ground color. Upper lip with several
irregular dusky markings. Concentration of melanophores appearing
as dark spot on upper portion of opercle. Posterior to eye, head bears
two diffuse groups of melanophores in form of indistinct bands separated
by pale ground color. Underside of head exhibits pale dusky chevron
in form of Y. Arms of Y originate at corners of mouth, extend to
midventral side of head, meet and extend posteriorly for short distance
(shank of Y). Another pale mark extends from each opercle ventrally
onto ventral side of head. Ground color appears between these marks
and those forming arms of Y, thus forming pale chevron (in none of
the specimens were the dusky chevrons comparably as dark as they
may appear in S. talboti).
Side of body bears scattered diffuse dusky markings. Along midside
are five concentrations of melanophores that form spots. First, and
palest, spot beneath appressed pectoral fin; next, much darker, below
posterior spinous dorsal; next, not quite so dark as previous, below third
through fifth dorsal rays; next, about as dark as previous, below last
four dorsal rays. Previous two spots appear composed of two coalesced
spots. Last, and darkest, spot on caudal peduncle at caudal base.
Anteriorly and basally spinous dorsal fin bears dusky band that widens
and breaks up into irregular dusky marks posteriorly. Spinous dorsal
unmarked anteriorly and dorsally. Segmented-ray portion of dorsal bears
116 Proceedings of the Biological Society of Washington
Fic. 2. Stanulus talboti new species, USNM 201372 male, paratype,
37.0 mm SL, from One Tree Island, Great Barrier Reef, Queensland,
Australia. a, lateral view. b, enlarged view of scalelike flaps of anterior
lateral line; one flap folded anteriorly to reveal opening to lateral line.
c, anterior view of head.
Indo-Pacific Blenniids of the Genus Stanulus JULY
irregular, diffusely dusky band at about mid-height of fin, followed
distally by unmarked area and another smaller, dusky band. Tips of rays
unmarked. Anal fin spines enveloped in unpigmented fleshy rugosities;
urogenital papilla unpigmented. Segmented-ray portion of anal fin
generally dusky except anterior tips of rays, which are unmarked. Caudal
fin bears about three irregularly diagonal, dusky bands followed pos-
teriorly by broader, vertical, dusky band that covers end of caudal.
Pectoral fins unmarked; fleshy pectoral base dusky proximally, marking
extending ventrally onto prepelvic area. Unmarked, crescentic area
present just distal to dusky area on pectoral base, followed by narrow
dusky crescent just proximal to bases of pectoral rays. Pelvic fins
unmarked.
In other specimens the spots on the sides, except the most posterior,
are either absent or so diffuse that they are not recognizable. Many
specimens exhibit as their most prominent mark the spot at the caudal
base. Most specimens bear three dusky, vertical stripes on the middle
of the upper lip similar to those illustrated for S. talboti (Fig. 2C). The
illustrated specimen had a dark spot, not found in the other specimens,
over the posterior dorsal spines. In the specimen illustrated by Smith
(1959, plate 17) there are several dark markings on the head, body,
and fins, but these were not visible when I examined the specimen.
Predorsal commissural pores range in number from 3 to 21. They
tend to increase in number with increase in standard length. The cor-
relation coefficient for this character, based on 49 specimens, was 0.598.
The regression formula was Y = 0.519077X + 1.41733, where Y equals
the number of pores and X equals the standard length.
Size and sexual dimorphism. The largest specimen available, 27.7 mm
SL, is a mature male, as evidenced by the presence of fleshy rugosities
enveloping the anal spines. The smallest specimen, 11.0 mm SL, is an
ophioblennius larva with three canines on either side of the lower jaw.
The next smallest specimens, 11.7-12.1 mm SL, are juveniles, indicating
that metamorphosis occurs at a size between 11.0 and 11.7 mm. Males
probably attain a larger size than females. The largest female, of 15
available, was 23.4 mm SL. Five males, of 32 available, were larger
than the largest female.
Distribution (see Fig. 3 and material list). All specimens of S. sey-
chellensis were collected after 1947. All but a very few have come from
the surge zone. Because of the small size of the species and its occur-
rence in a habitat difficult to collect, I believe that the known distribu-
tion is an incomplete outline of the actual distribution. Another small
species that inhabits the surge zone, Entomacrodus thalassinus (Jordan
and Seale), is also known only from the Seychelles Islands in the Indian
Ocean (Springer, 1967). The only specimen of E. thalassinus known
from Australia was taken in the same collection as the only specimen
of S. seychellensis known from Australia, and both of these were taken
together with a relatively large number of S. talboti.
Material: La Digue, Seychelles Islands: Rhodes University (not cat-
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Indo-Pacific Blenniids of the Genus Stanulus 119
aloged, holotype of Stanulus seychellensis. One Tree Island, Great
Barrier Reef, Australia: USNM 201372 (1 specimen). Marshall Islands:
USNM 201554 (1); Eniwetok Atoll, LACM 6679-45 (3), 6679-54 (1);
Jieroru Island, USNM 142150 (2); Mui Island, USNM 142149 (5);
Bikini Atoll, USNM 142151 (1), 142153 (holotype of Fallacirripectes
minutus ); Eman Island, USNM 142152 (3, cleared and stained ); Ronge-
lap Island, USNM 142154 (holotype of Fallacirripectes wellsi). Guam,
Marianas Islands: CAS (all numbers are George Vanderbilt Foundation
register numbers) 1856 (4). Tapatuaitu, Kapingamarangi Atoll: CAS
356 (6). Raroia, Tuamotu Archipelago: CAS 113 (10). Ifaluk Atoll,
Carolina Islands: Falarik Islet, CAS 126 (3), 197 (2), 196 (5); Falalap
Islet, CAS 128 (2); Ulithi Atoll, CAS 482 (2).
Stanulus talboti new species
Fig. 2, Table 1
Description: Color pattern. The following description is based on the
illustrated specimen, a male, followed by mention of notable variations
encountered in the other specimens.
Body and head with pale ground color. Upper lip variably dusky
with three dusky stripes medianly and darker, dusky marking at each
corner of lip. Sides of head variably dusky with adumbrations of dark
spot just posterior to eye, followed by pale area and pale dusky stripe.
Underside of head exhibits two black spots on each side. Anterior two
spots just posterior to corners of lower lip, separated by narrow pale
area. Second pair of spots separated from anterior pair by crescentic
pale area. Posterior spots curved, extend medianly but fail to meet,
separated by a broad pale area. Preopercular and opercular areas vari-
ably dusky. Body bears number of irregular, diffusely dusky areas and
several dark spots.
Spinous dorsal fin dusky ventrally and dorsoposteriorly; dorsoanteriorly
unmarked. Segmented-ray portion of dorsal variably dusky with adum-
brations of darker and lighter, diagonal diffuse stripes coursing over it.
Anal spines enveloped in pale fleshy rugosities; segmented-ray portion
of anal fin dark dusky, darker distally except that anterior tips of rays
pale. Caudal fin evenly pale dusky. Pectoral fin bears irregular dusky
markings. Fleshy pectoral base dark dusky basally, followed distally
by narrow, crescentic pale area. Pelvics unmarked.
Other males vary primarily in having the various markings either
more or less intensified. The spots along the midside of the body may
be relatively larger and extend ventrally as paler, diffuse dusky mark-
ings, giving the appearance in some specimens of paired bands. The
markings on the head may be present only as diffuse dusky concentra-
tions of melanophores.
Females are usually much paler than males, and some bear almost
no distinct marks.
The three stripes on the upper lip (Fig. 2C) are present in all
specimens.
120 Proceedings of the Biological Society of Washington
Predorsal commissural pores range in number from 8 to 22. They
tend to increase in number with increase in standard length. The
correlation coefficient for this character, based on 42 specimens, was
0.510. The regression formula was Y = 0.449814X — 0.840596, where
Y equals the number of pores and X equals the standard length. When
compared with the regression line for the same character in S. seychel-
lensis, it can be seen that specimens of the latter species will usually
have more pores at any given standard length than will S. talboti (see
above). A covariance test of the two regressions indicates that the
slopes are not significantly different, but that the heights are signif-
icantly different at the 99.9 per cent level (F of slopes 0.19, F of heights
12.48, F 99.9 = 11.70).
Size and sexual dimorphism. The largest specimen available, 39.0 mm
SL, is a mature male, as evidenced by the presence of fleshy rugosities
enveloping the anal spines. The smallest specimen is a juvenile male,
19.4 mm SL. Males probably attain a larger size than females. The
largest female, of 15 available, was 33.6 mm SL. Thirteen males, of
27 available, were larger than the largest female.
Distribution (see Fig. 3 and distribution under S. seychellensis). All
specimens are known only from One Tree Island, Great Barrier Reef.
The species occurs in the surge zone at depths of about 5 to 10 meters.
Because of the difficulty of collecting such a habitat, it is probable that
the species is distributed more widely than is presently recorded.
Holotype: USNM 202421, adult male, 29.4 mm SL, from south reef
face (ocean side), about one mile from One Tree Island, Great Barrier
Reef, Queensland, Australia, coll. 22 November 1966, by V. G. Springer
and party. Depth: approximately 5 meters; bottom living coral and
coral rubble. Dorsal fin XII, 12; anal II 13; pectorals 15-15, pelvic I,
4-I, 4; caudal 13, predorsal commissural pores 11; vomerine teeth 6;
gill-rakers 12, pseudobranchial filaments 7; vertebrae 10 + 21.
Paratypes: USNM 201371, 19 specimens collected with holotype.
USNM 201372, 25 specimens (including 2 cleared and stained), off
reef on west side of One Tree Island, just shoreward of drop off (all
Stanulus were taken from a barren surge trench about 2.5 meters below
coral surface, which was 10 meters below surface), 7 December 1966,
by V. G. Springer and party, depth 8 to 12 meters, bottom living coral,
Some of the above paratypes will be deposited at a later date in the
Australian Museum, Sydney.
Named for my friend, Dr. Frank H. Talbot, who participated in the
collection of the specimens, and who organized the expedition that
made the collections possible.
Acknowledgments: Appreciation is here extended to Dr. J. L. B.
Smith, Rhodes University, who generously lent me the holotype of
Stanulus seychellensis. Dr. D. K. Caldwell and Mr. R. J. Lavenberg,
LACM, and Dr. William Eschmeyer and Miss Pearl Sonoda, CAS, lent
specimens in their care. Fig. 1 was drawn by Ann Hoskins and Fig. 2
by Martha Hatch Lester. Dr. B. B. Collette offered critical comments
—
Indo-Pacific Blenniids of the Genus Stanulus WAIL
that improved the manuscript. This study was supported by a grant
from the TFH Fund of the Smithsonian Institution.
ADDENDUM
While this paper was in press Dr. J. E. Randall provided me with
nine specimens of S. seychellensis (Bishop Museum no. 5896) he col-
lected from the reef front on the ocean side of Igurin Island, Eniwetok
Atoll. Eight were males, 22.5-30.7 mm SL, and one was a female, 21.1
mm SL. Randall also made available to me a color photograph he took
of one of the males, 30.7 mm SL, while the specimen was in fresh
condition. The ground color of this specimen was bright white and was
overlain on the body by a coarse brownish-orange reticular pattern. The
fleshy pectoral base bore a bright orange spot distally separated by a
narrow, bright white line from the duskier proximal portion of the base.
The head was also brownish orange except for a large, deep orange
hexagonal marking surrounding a dark gray area on the opercle. Almost
all of the brownish-orange, orange and gray areas vanished after pres-
ervation, which may account for the differences I noted above between
the specimen illustrated by Smith (1959) and his illustration.
LITERATURE CITED
NyBELIN, O. 1963. Zur Morphologie und Terminologie des Schwanz-
skelettes der Actnopterygier. Ark. Zool., vol. 15, pp. 485-516.
ScHuLtz, L. P. anp W. M. CHapMan. 1960. Subfamily Salariinae.
In Fishes of the Marshall and Marianas Islands. U.S. Nat.
Mus. Bull. 202, pt. 2, pp. 302-372.
SmirH, J. L. B. 1959. Fishes of the families Blenniidae and Salariidae
of the western Indian Ocean. Dept. Ichthy. Rhodes Univ.,
Ichthy. Bull. 14, pp. 229-252.
SPRINGER, V. G. 1967. Revision of the circumtropical shorefish genus
Entomacrodus (Blenniidae: Salariinae). Proc. U.S. Nat.
Mus., vol. 122, no. 3582, pp. 1-150.
In press. Osteology and classification of the fishes of the
family Blenniidae. U.S. Nat. Mus. Bull. 284.
122 Proceedings of the Biological Society of Washington
AHF, O6 ZZ
Vol. 81, pp. 123-142 April 30, 1968
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
GEOGRAPHIC VARIATION IN THE NEW WORLD
GECKKONID LIZARD TARENTOLA
AMERICANA GRAY
By ALBERT SCHWARTZ
Dept. of Biology, Miami-Dade Junior College,
Miami, Florida 33167
Tarentola americana Gray is the sole New World member
of the otherwise Old World genus Tarentola. Although the
first name proposed for the New World species is T. (as
Platydactylus) americana Gray (1831), this name was based
upon a specimen supposedly taken in New York, and ameri-
cana was long considered a distinct species which had not
been subsequently collected. Later, Duméril and Bibron
(1836) redescribed (using the same specimen as the holotype)
T. americana, naming the species Platydactylus milbertii. Fin-
ally, Gundlach and Peters (1835) described Platydactylus
americanus var. cubanus from Cuba. The Antillean Tarentola
was consistently called T. cubana (Barbour, 1914; Barbour,
1916; Barbour and Ramsden, 1919; Alayo, 1951), but Love-
ridge (1944) relegated this name to the synonymy of T.
americana, since he considered that all three names (ameri-
cana, milberti, cubana) referred to the same species. Later
authors (Alayo, 1955; Schwartz and Ogren, 1956; Ruibal,
1957; Buide, 1966, 1967) have consistently used T. americana
for these geckos.
T. americana was long considered a rare (or local) inhab-
itant of Cuba. Barbour (1914: 259) commented that the
species was seldom found; at that time the Museum of Com-
parative Zoology at Harvard University had material only
from Santiago de Cuba, and Barbour had seen a single speci-
men from Cienfuegos. Barbour also reported that Gundlach
had found only two specimens in his entire life in Cuba, one
16—Proc. Biot. Soc. WasH., Vou. 81, 1968 (123)
124 Proceedings of the Biological Society of Washington
from San Diego de los Bafios in the Sierra del Rosario in
Pinar del Rio Province and the other from Cabo Cruz in
Oriente Province. Barbour and Ramsden (1919: 116-117)
stated that Tarentola was “generally a very rare lizard,”
although they stated that there were particular localities where
the species was easily obtained (such as at Puerto Escondido
near the Bahia de Guantanamo in Oriente Province). They
stated that the species was extremely rare outside of a few
stations in Oriente, but that it had been collected in the
westernmost Cuban province, Pinar del Rio; thus the lizard
was shown to have an islandwide distribution. Ruibal (1957:
257) secured T. americana abundantly on the cays of the
Laberinto de los Doce Leguas off the south coast of Camagiiey
Province; later, Buide (1966: 4) noted that he had collected,
in only three visits to the locality, about 60 specimens from
Cueva Ambrosio on the Peninsula de Hicacos on the north
coast of Matanzas Province. Far from being rare outside of
Oriente, T. americana can be secured in some numbers, pro-
vided its habitat requirements are known.
Barbour (1916: 219-220) was the first to report Tarentola
(as T. cubana) from the Bahama Islands; he had a single
specimen, collected by C. J. Maynard, from U Cay, of the
Allan’s Harbour Cays near Highborn Cay, in the northern
portion of the chain of Exuma Cays. Later, Barbour (1930:
82) reported the lizard from Andros Island in the Bahamas.
Although Rabb and Hayden (1957: 23) collected fourteen
specimens of T. americana, all were from Leaf Cay of the
Allan’s Harbour Cays, so that the known distribution of the
species in the Bahamas was not considerably extended by
their fine series. Although T. americana is presently fairly
well represented in American collections from most Cuban
provinces (with the exceptions of Habana and Matanzas
provinces), there has been an extreme paucity of specimens
of this lizard from its Bahaman range, with the material
reported by Rabb and Hayden being the only Bahaman series
available. Although T. americana had only been reported
from two regions (Andros; Exuma Cays) it was expected that
its range would ultimately be shown to include many (if not
all) of the islands on the Great Bahama Bank, at least. Pre-
Variation in New World Lizard 125
vious lack of sufficient Bahaman material has prevented
comparison of Cuban and Bahaman specimens; fortunately
there are now available a total of 44 Bahaman specimens so
that variation in T. americana can now be clarified.
Almost half of the Cuban T. americana examined are the
result of collections made in Cuba by myself and parties
between the years 1956 to 1960; these specimens are in the
collection of the American Museum of Natural History
(AMNH). I have also borrowed material in the collections
of the Camegie Museum (CM), Museum of Comparative
Zoology (MCZ), University of Florida, Florida State Museum
(UF/FSM), and the Museum of Zoology, University of Mich-
igan (UMMZ). For the loans of these lizards I wish to thank
Charles M. Bogert and George W. Foley, Neil D. Richmond
and Clarence J. McCoy, Jr., Walter Auffenberg, and Charles
F. Walker and Arnold G. Kluge. The large number of Baha-
man individuals now in collections are in part due to the
activities of Wayne King, Neil D. Richmond, Richard Thomas,
and C. Rhea Warren; material collected by Messrs. Thomas
and Warren are in the Albert Schwartz Field Series (ASFS),
whereas that collected by Dr. King is in the UF/FSM collec-
tion; the series collected by Dr. Rabb and Mr. Hayden has
already been mentioned. My Cuban collections were made
under National Science Foundation Grants G-—3865 and G-
6252; during my Cuban field work I had the assistance of
John R. Feick, William H. Gehrmann, Jr., Ronald F. Klinikow-
ski, David C. Leber, James D. Smallwood, Barton L. Smith,
Richard Thomas, and George R. Zug, to all of whom I am
grateful for their cooperation.
I have taken the following counts and measurements on all
adult and subadult T. americana. A total of 130 specimens is
available from Cuba, the Bahamas, and the Isla de Pinos; the
species has not been previously reported from the latter island,
although it was expected there.
1) Sex. The external sexual characteristics of T. americana are rather
subtle, but once they are understood, all specimens, regardless of age,
can be sexed without difficulty. Just posterolaterally to the vent, there
is in males a pair of semilunar series of enlarged (in respect to the scales
both anteriorly and posteriorly to them) scales, usually three in number,
126 Proceedings of the Biological Society of Washington
which project boldly from the lateral and ventral margins of the tail
and are on a rounded prominence. In females, these scales are not so
enlarged nor so conspicuous nor on an enlarged prominence. In males,
the scales are usually dead white and contrast sharply with the buffy
or brownish color of the tail, whereas in females the scales are often
concolor with the remainder of the tail surface in that area. Although
these color differences are not absolute, the bold and tooth-like appear-
ance of the semilunar scales in males is a striking feature of that sex.
Verification of these sexual differences was made by extrusion of hemi-
penes in males and visible shelled eggs in females. Sex of even very
small T. americana can be determined with a high degree of accuracy,
if the student is aware of the changes in size of these semilunar scales
in respect to the size of the animal.
The hemipenis of T. americana has not been previously described. As
extruded (ASFS V11232), the organ is relatively short, extending for
about the length of seven or eight ventral caudal scales, and is rather
strongly bifurcate for its distal third. The basal half is smooth; the distal
half is minutely calyculate on both the sulcate and non-sulcate surface.
The sulcus spermaticus divides at that point where the hemipenis
becomes bifurcate, each branch extending thence onto the terminal .
surface of the weakly crenulate apices; each of these branches forks
once more, on the apex, or, perhaps more properly, on each apex are
two weak grooves which radiate from the termination of the fork of the
sulcus spermaticus. The hemipenis is quite simple in total aspect,
although as extruded it is rather robust.
2) Snout-vent length, in millimeters.
3) Longitudinal rows of enlarged dorsal scales at midbody. Although
this count has been employed by other students of the genus Tarentola
(Loveridge, 1944; Grandison, 1961), the situation in T. americana is
such that a perfectly accurate count of these scales is not possible. The
dorsal scales, in adults, are keeled and prominently set off from the much
smaller interstitial paving scales; as these enlarged scales progress down
the sides, and especially at the extreme ventrolateral area, they become
smaller and stud-like, less obvious, and more like the ventrals in size.
Although the ventrolateral “dorsals” are quite different from the smooth
and imbricate ventrals, ventrolaterally the former become more like both
the interstitial scales and the ventrals. In one population, there are
additionally about four to eight rows of transitional scales scattered
between the ventrolateral “dorsals” and the ventrals, so that the arrange-
ment of scales in this whole ventrolateral area, with its conglomeration
of differently sized scales, all of which intergrade with one another in
size, makes an accurate count of enlarged dorsal rows difficult indeed.
Loveridge (1944: 19) gave the range of dorsal tubercles as 13 to 17
in 15 Cuban and two Bahaman specimens, whereas in a much larger
series, my counts vary between 12 (MCZ 11877, a specimen examined
by Loveridge) and 24 (ASFS V11232, a Bahaman specimen not avail-
able to Loveridge). Although there are mean and modal differences in
Variation in New World Lizard 197
dorsal counts as I made them, I strongly doubt that any two investigators
would count the same number of longitudinal rows of dorsal scales on
many specimens.
4) Number of ventral scales, counted transversely at midbody. The
ventral scales of T. americana are smooth, rounded, and imbricate. The
transition from the ventrolateral enlarged studs and interstitial scales
to the smooth ventrals is fairly abrupt, but, if a count is made from what
appears to be the area of transition on one side to that of the other,
a great source of error is encountered; one is simply not sure which of
the transitional scales should be considered ventrals and which slightly
enlarged and smoother interstitials. Fortunately, a more finite boundary
for ventrals occurs in T. americana. Laterally, the belly is set off from
the sides by a pair of longitudinal folds, one on each side, similar in
aspect to the belly disc in some frogs. When preserved, the lateral
limits of the venter are strongly shown by these lateral folds; even in
specimens which were injected upon preservation, the folds are dis-
cernible and make excellent landmarks for beginning and ending the
ventral counts. Since these counts are easily made and the limits on
each side quickly and accurately determined, they are far more reliable,
and I have placed much more emphasis upon them than on the dorsal
counts.
5) Interorbital scales. This count includes the superciliary scales on
each side.
6) Gulars. Grandison (1961: 3) counted the number of transverse
gulars between the ears. I have counted the longitudinal number of
gulars contained within the distance between the posterior border of the
orbit and the anterior border of the auricular opening, this distance set
off by vernier calipers on the center of the throat at the level of the
ears. The gulars in T. americana are small, smooth, rounded, and closely
juxtaposed. This count was not taken on many very small lizards. As
in any situation where a particular distance (eye-ear, in this case) is
laid out on another body part, there is a possibility that heterogonic
growth of the head in reference to the gulars might create artificial
differences between adult and juvenile lizards. This is not the case in
T. americana; likewise, there are no sexual differences in this count.
The differences in number of gulars are so striking that it would be
difficult indeed to obscure them.
7) Number of lamellae on first and fourth toes. These data were
taken on most specimens, but the variation within any series is fairly
broad and the count appears meaningless as far as populations of T.
americana are concerned.
8) Scales between the first sublabials, in contact with the mental.
In effect, this is a postmental count, since only those scales (gulars) in
contact with the mental and the first pair of sublabials were counted.
The count varies from 0 (sublabials in contact behind mental) to 4 (four
scales in contact with the mental and enclosed between the first sub-
labials ).
128 Proceedings of the Biological Society of Washington
These counts have been variously useful in defining the subspecies of
T. americana. The differences in scutellation between the Cuban and
Bahaman populations of this lizard are so striking that, for instance,
comparison of lizards of comparable snout-vent lengths easily demon-
strates the difference in size of ventrals, a fact verified by the counts
themselves. The color and pattern differences between the two basic
populations are, for the most part, less easy to explain but are nonetheless
real. ;
Systematic Accounts
Tarentola americana Gray, 1831
Platydactylus americanus Gray, in Griffith, Animal Kingdom,
9:48 (type locality—“New York”).
Platydactylus Milbertii Duméril and Bibron, 1836, Erpét. Gén.,
3:325 (type locality—“New York’).
Platydactylus (Tarentola) americanus var. cubanus Gundlach
and Peters, 1865, Monatsb. Akad. Wiss. Berlin: 384 (type
locality—Cuba; here restricted to Cabo Cruz, Oriente Prov-
ince, Cuba, one of two localities whence Gundlach and Peters
had material at the time of the proposal of the name
cubanus).
Type-locality: “New York”; considering the period (1831) when the
holotype of P. americanus was collected, that the holotype was in the
Paris Museum at that time and was still there at the time that Duméril
and Bibron (1836) renamed the same individual P. Milbertii, and that
it was sent to the Paris Museum by a M. Milbert, it seems likely that
the specimen originated in one of the Cuban ports, and possibly even
that port with which there was considerable French trade at that time;
note that Duméril and Bibron stated that “Cette espéce nous a été
envoyée de New-Yorck par M. Milbert,” not that the specimen was
collected in that city (although it may have been). Although both
La Habana and Santiago de Cuba were prominent ports of that period,
it seems more likely that the lizard originated in the vicinity of Santiago
de Cuba. Tarentola is common in that region (Alayo, 1951), and trade
between Santiago de Cuba and the previously French city of Port-au-
Prince in Haiti was flourishing, due in part to presence of French coffee
plantations, begun by exiles from Saint-Domingue during the revolution
on Hispaniola (1791), in the Sierra de la Gran Piedra and other uplands
in the vicinity of Santiago de Cuba. Marrero (1951: 608) stated that
the slave rebellion in Saint-Domingue “was a notable factor in the
growth and progress of Santiago. In 1808 the city had more than 20,000
inhabitants and new zones of cultivation, principally of coffee and cacao,
had been developed in the mountainous areas of the Sierra Maestra,
with Santiago as the commercial center. The French influence on the
culture and customs of Santiago was notable.” It seems very likely that
the original specimen of T. americana was collected in the Santiago de
Cuba region and was sent to Paris via New York. I therefore hereby
Variation in New World Lizard 129
restrict the type locality of Tarentola americana Gray to the vicinity of
Santiago de Cuba, Oriente Province, Cuba.
Tarentola americana americana Gray
Type-locality (by restriction): Vicinity of Santiago de Cuba, Oriente
Province, Cuba.
Distribution: Throughout the island of Cuba (known in the literature
from all provinces) and the Isla de Pinos (see, however, further discus-
sion of the status of the Isla de Pinos specimens), and the Laberinto de
las Doce Leguas (and probably many other off-shore islands and cays).
See Fig. 1.
Definition: A subspecies of T. americana characterized by large size
(males to 111 mm, females to 100 mm snout-vent length on Cuba, larger
on the Isla de Pinos), high number of transverse ventral scales (usually
30 to 47), high number of interorbital scales, high number of gular
scales, scales bordering the mental posteriorly usually 1; ventrolateral
area between enlarged lateral scales and ventrals covered with from
four to eight vertical rows of small interstitial scales which differ sharply
from the enlarged laterals and grade into the ventrals; dorsal colors
primarily shades of tan to brown, not gray to black, and pale bands on
unregenerated tails vaguely streaked with darker (brown).
Remarks: The series of 80 Cuban (in contradistinction to Isla de
Pinos) T. a. americana has the following measurements and counts:
largest male (AMNH 76545—Pinar del Rio) 111 mm snout-vent length,
largest female (MCZ 59319—Las Villas) 100 mm snout-vent length;
enlarged dorsal rows 12-22 (mean 17.4, mode 17), ventrals 27-47
(mean 33.5), interorbitals 10-17 (14.3), gulars 17-31 (24.6), first toe
lamellae 10-17 (13.8), fourth toe lamellae 9-17 (14.5); scales between
sublabials modally 1 (40 specimens), but other counts include 0 (11),
9} (QO). BY (Wa eral Zo (Cib)),
Very few Cuban T. a. americana have snout-vent lengths greater than
100 mm; only 11 males have snout-vent lengths in excess of that dimen-
sion, and only one female reaches that size (although three other females
have snout-vent lengths between 90 and 100 mm). There is thus some
sexual dimorphism in overall size, but it is not striking. Although the
total range of ventral scales is 27 to 47, only 6 of 72 Cuban specimens
have counts below 30, and the mean (33.5 + 1.01) is high.
Although there is rather limited material from throughout Cuba, there
does not seem to be any west-east trend in number of ventral scales (or
in numbers of other scales on which I have taken data; I emphasize
ventrals because they are numerous and the count is easily and accurately
taken). The Oriente sample includes both extremes (27 and 47), the
low extreme from a specimen on the southern Oriente coast east of the
Bahia de Guantanamo (AMNH 17727) and the high extreme from a
specimen on the same coast between the Bahia de Santiago and the
Bahia de Guantanamo (AMNH 95981). The modal number of ventrals
in Oriente is 37, whereas that in Las Villas is 30 or 33 (both with equal
130 Proceedings of the Biological Society of Washington
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Variation in New World Lizard 131
incidences); this might indicate that there is a trend toward higher
number of ventrals in the east. The Pinar del Rio material, limited as
it is (six specimens), includes specimens with high counts of 39, 43,
and 47 (AMNH 81207), so the east-west cline seems to be reversed
in the extreme western portion of the island.
The number of scales between the sublabials and bordering the mental
posteriorly is modally 1 in all samples; both the Las Villas and Oriente
samples have equal incidences (57.1 per cent) of this character. How-
ever, the material from the Laberinto de las Doce Leguas is bimodal,
with 0 scales in this position having an equal incidence as 1 scale (6
specimens in each category of 16 total specimens). The Doce Leguas
specimens likewise include the only lizard with 4 scales in this position
of all Tarentola examined.
Description of the pattern of T. a. americana is made difficult by
the extreme changes and obfuscation of pattern with ontogeny. In
essence, the pattern consists of the following elements. The dorsal
ground color is tan, with a series of four or five dark brown transverse
bands across the back between the limbs, at times with some pale tan
stippling along the margins of the bands. There is an additional dark
brown band across the neck and there may be a second band across
the neck behind the occiput. The head is tan and in younger individuals
has a dark brown figure, made up of fine lines, which resembles an
arrow-head, with its apex between the eyes and its open ends on the
occiput. Anterior to the apex of the arrow-head may be a single median
line which proceeds toward the snout and, if fully developed, forks
toward the lores on each side. There is a dark brown postocular stripe,
margined above by tan to buffy, which proceeds posteriorly over the
shoulder and forms the lateral margins of the one (or two) nuchal bands
as well as (usually) the first body band above the forelimbs. The
unregenerated tail is banded with about eight or nine dark brown bands,
which alternate with pale tan bands; the latter are regularly smeared
with longitudinal dark brown markings, which at times are almost striae.
The limbs are irregularly marbled or flecked with tan and brown; the
underside is ochraceous tan in adults.
The above color and pattern notes summarize the most brightly
patterned condition in T. a. americana. Many individuals have the dorsal
bands much obscured or virtually absent, with a resultant only vaguely
crossbanded lizard. The head markings may include the figure described
above as well as a more posterior dark line from between the bases of
the cephalic arrow-head onto the neck; in other lizards the head pattern
is grossly fragmented or obscured or absent. Lizards lacking the head
pattern are most often large adults, but other large adults still have the
head pattern present and prominent. The postocular dark line is a
regular feature at all ages; some specimens have an additional para-
median pair of dark nuchal stripes, derived from portions of the nuchal
blotch, of equal width and intensity as the postocular stripes. Variation
in pattern details is legion, but the basic colors are always tans and
132 Proceedings of the Biological Society of Washington
browns (rather than grays and black), and at least some of the pattern
elements described above are present on most individuals.
Special comment remains to be made on the scutellation of the lower
sides. As noted previously, the enlarged dorsals decrease in size as
one progresses ventrolaterally, so that on the lower sides they are smaller
than on the back but still somewhat enlarged (compared to the inter-
stitial scales) and stud-like. In T. a. americana, between these lowermost
enlarged “dorsals” and the fold which laterally delimits the venter and
its smaller, smooth, and rounded scales, there is an area of small flattened
scales. These scales are arranged in from four to eight vertical rows
(depending upon the ventral extent of the enlarged scales above them),
and these small scales grade gradually into the ventral scales, so that
the scales on the ventrolateral folds are typical of those of the venter,
not of the transitional scales. As will be seen later, the Bahaman popula-
tions lack this feature of scutellation.
The six lizards from the Isla de Pinos are three adult and one juvenile
males and two adult females. These lizards differ in several ways from
their Cuban relatives. First, the largest specimens of all T. americana
are in this Isla de Pinos series, despite much greater numbers of speci-
mens from both Cuba and the Bahama Islands. The largest Isla de Pinos
male has a snout-vent length of 120 mm, and the two other adult males
have snout-vent lengths of 119 and 100 mm. The largest female T.
americana is an Isla de Pinos individual with a snout-vent length of
111 mm, and the other female from that island has a snout-vent length
of 105 mm. Although the differences in snout-vent lengths between
this short Isla de Pinos series and the Cuban material are not exception-
ally great, it may be more than coincidence that the largest Tarentola
are from the Isla de Pinos. Secondly, the adults of both sexes from the
Isla de Pinos are virtually patternless, having only the vaguest indication
of crossbars or none at all. Two adult individuals with complete or
nearly complete tails show no tail banding, the tails being concolor with
the dorsum. Even the regularly consistent postocular dark stripe in
Cuban specimens is suppressed and absent in these Isla de Pinos adults.
Finally, the juvenile individual (snout-vent length 62 mm) had the
“pale” tail crossbands pale orange in life, a color not recorded in Cuban
specimens. The juvenile specimen is, however, patterned like Cuban
T. a. americana, or, perhaps more properly, the Isla de Pinos juvenile
falls within the variation shown by Cuban lizards.
Scale counts on the six Isla de Pinos T. americana are: enlarged dorsal
rows 17-29 (mean 19.0, mode 20), ventrals 35-37 (35.5), interorbitals
13-16 (15.0), gulars 19-31 (26.2), first toe lamellae 14-16 (14.5),
fourth toe lamellae 14-16 (14.8); scales between sublabials modally 2
(4 specimens), but two specimens have 1 scale. Although the means
of all counts are higher for the Isla de Pinos specimens, and the modal
condition of scales between the postlabials is 2 (rather than 1 as in
Cuba), the small series from the Isla de Pinos precludes any statistical
analysis of these data. I think it highly likely that additional material
Variation in New World Lizard 133
from the Isla de Pinos will show that the populations there differ rather
markedly in size, pattern, and scutellogical detail from their Cuban
relatives.
Several authors (Schwartz and Ogren, 1956; Hardy, 1957; Buide,
1966) have pointed out that in Cuba T. americana is regularly associated
with caves (where the animals rest in the twilight zone near the entrances
during the day) and cliffs (where the animals forage, in addition to
the cave mouths, at night). Ruibal (1957) reported the occurrence of
T. americana inside the old erect dead trunks of mangroves (Lagun-
cularia) on the Laberinto de las Doce Leguas off the Camagiiey coast.
These two habitats—cliffs and caves, and dead tree trunks—are the
customary places where T. a. americana has been found. Other sites
where lizards have been collected include inside a newly constructed
storage shed and in a hollow royal palm (Roystonea). Both Alayo
(1951: 107) and Buide (1967: 25) cited the capture of T. americana
in the roofs of native bohios. All these latter situations are generally
vertical faces as are the previously reported situations. Barbour (1914:
259) reported taking a specimen at Cabo Cruz under a stone, and
Barbour and Ramsden (1919: 117) noted that numbers of specimens
had been secured at Puerto Escondido in and under dead dry cactus.
On occasion I have encountered T. a. americana on the ground, where
it is often associated with rough limestone rocks which are imbedded
in rocky soils. Near Guane in Pinar del Rio, a lizard was collected in
such a situation, although there were nearby a cave and low rocky cliffs.
Perhaps the most unusual situation wherein I have taken T. a. americana
is that of the specimen from west of Cayuco on the Peninsula de Guana-
hacabibes; this individual was taken on a live but hollow tree in dense
hardwood forest. There were no caves or cliffs in the immediate area,
but the ground surface at this locality was rocky. Apparently T. a.
americana exists in some regions where caves and their associated cliffs
are absent, provided there is cover for diurnal retreats in the form of
either large and chunky limestone rocks on the ground or hollow trees.
T. a. americana deposits its eggs in small hollows in the roofs and
lower walls of caves or in small but fairly extensive rocky cavities (large
enough to admit a person) in cliff faces. The eggs, of which two is
the clutch, are white, hard-shelled, and adherent to the tops or sides of
the cavities wherein they are deposited; most often there are large
numbers of eggs deposited in the same immediate area, and the cave
or cavity floor beneath the eggs is littered with many broken egg shells,
which fall to the ground after the young have hatched. Im this charac-
ter, T. a. americana resembles Anolis lucius (see Allen and Neill, 1957,
and Hardy, 1957, whose observations I can readily confirm) and Anolis
bartschi, both of which have communal egg deposition areas and both
of which occur syntopically with Tarentola on the same cliffs and cave
entrances. However, A. lucius and A. bartschi are allopatric, so that
Tarentola occurs only with one species or the other at any single locality.
134 Proceedings of the Biological Society of Washington
A hatchling T. a. americana (MCZ 68937) has a snout-vent length of
24 mm.
No specimens of T. a. americana indicate its occurrence in the Cuban
or Isla de Pinos uplands; most records are coastal, but this may only
indicate the greater accessibility of coastal habitats, greater facility for
collecting this species there, or more common occurrence of suitable
habitats (caves and cliffs) at lower elevations or along the coast. The
species occurs at San Vicente in the Sierra de los Organos at an elevation
of about 1,000 feet (305 meters); Barbour and Ramsden (1919: 117)
reported its occurrence at San Diego de los Bafios in the adjacent Sierra
del Rosario and at Los Negros in the northern foothills of the Sierra
Maestra in Oriente. Koopman and Ruibal (1955: 6) reported a single
fossil Tarentola dentary from pre- or post-Columbian cave deposits in
the Sierra de Cubitas in Camagiiey Province. Apparently the species
occurs at least at intermediate elevations in some (and perhaps all) of
the mountain massifs in Cuba.
Specimens examined: Cuba, Pinar del Rio Province, 18 km W Cayuco,
1 (AMNH 81204); 3.5 km NE Guane, 1 (AMNH 81207); cliffs near
San Vicente, 2 (AMNH 76545-46); 0.5 mi. (0.8 km) S San Vicente,
1 (AMNH 78204); 4 km S Rancho Mundito, 2 (AMNH 81205-06); ©
Las Villas Province, Soledad, 1 (MCZ 43801); Guajimico, 16 mi. (25.6
km) SE Soledad, 2 (AMNH 78205-06 ); Trinidad, 7 (AMNH 78207-13);
Finca Morales, 8 mi. (12.8 km) NW Trinidad, 8 (AMNH 78214-21);
8 mi. (12.8 km) NW Trinidad, 1 (AMNH 95980); 6 km W Trinidad,
1 (MCZ 59320); 12-13 km from Topes de Collantes, 1 (MCZ 59319);
Cueva de Caguanes, Punta Caguanes, 1 (AMNH 81208); Camagiiey
Province, Cayo Cabeza del Este, 7 (MCZ 56358, 56365-70); Cayo
Levisa, 8 (MCZ 57297, 56373-74, 56376-77, 56380, 56382, 56384);
Oriente Province, Cabo Cruz, 1 (MCZ 8506); Santiago de Cuba, 2
(MCZ 6919, 19770); Morro Castle, Santiago de Cuba, 3 (UMMZ 90721);
Playa Juragua, 3.7 mi. (5.9 km) E Siboney, 6 (AMNH 83823—3 speci-
mens, 95981—3 specimens); United States Naval Base, Guantanamo
Bay, 5 (MCZ 93702, 68936-37, 67917, UMMZ 115733); Puerto Escon-
dido, 1 (MCZ 9435); between Janeo and Imias, 2 (AMNH 17727-28);
Caleta, costa sur, Baracoa, 1 (AMNH 17720); Cabo Maisi, 1 (AMNH
12855); Cabo Maisi, San Lucas (not mapped), 2 (AMNH 17725-26);
Cabo Maisi, La Patana (not mapped), 10 (MCZ 11871, 11873~-75,
11877-79, 96530, UF/FSM 21936, UMMZ 50073); Yaitresito, Baracoa
(not mapped), 2 (MCZ 47047, 96331); Isla de Pinos (Habana Prov-
ince), just W Nueva Gerona, east base, Sierra de las Casas, 2 (AMNH
81198, 81203); 1 mi. (1.6 km) SSW Nueva Gerona, east base, Sierra
de las Casas, 3 (AMNH 81199-201); Bibijagua, 1 (AMNH 81202).
Tarentola americana warreni new subspecies
Holotype: United States National Museum (USNM) 160725, an
adult male, from Gray’s Settlement, Long Island, Bahama Islands, one
Variation in New World Lizard 135
Fic. 2. Map of the Bahama Islands, showing (solid circles) localities
whence specimens of T. a. warreni have been examined.
of a series collected between 27 and 29 May 1967 by C. Rhea Warren.
Original number V11233.
Paratypes: ASFS V11232, V11234, same data as holotype; ASFS
V8587-91, CM 45821, USNM 160724, same locality as holotype, 23-30
January 1965, C. R. Warren; ASFS V10839, east of Salt Pond, Long
Island, Bahama Islands, 13 October 1966, R. Thomas.
Associated specimens: Bahama Islands, Eleuthera, lighthouse, South-
east Point, 1 (MCZ 96078); Andros, 1 (MCZ 19566); Exuma Cays,
U Cay, Allan’s Harbour Cays, 1 (MCZ 12330); Leaf Cay (= S. W.
Allan’s Cay), Allan’s Harbour Cays, 18 (ASFS V10704—-05, AMNH
80131-37, CM 41135-36, UMMZ 117391—7 specimens); Warderick
Wells Cay, 10 (UF/FSM 23096-105); Ragged Islands, Great Ragged
Island, Duncantown, 2 (UMMZ 11802425).
Distribution: Islands of the Great Bahama Bank, Bahama Islands;
known from the islands of Eleuthera, Andros, Exuma Cays (U Cay or
S. W. Allan’s Cay, Leaf Cay, Warderick Wells Cay), Long, and Great
Ragged. The species has not heretofore been reported in the Bahamas
from Long Island or the Ragged Islands. See Fig. 2.
Definition: A subspecies of T. americana characterized by small size
(males to 92 mm, females to 88 mm snout-vent length), low number of
transverse ventral scales (usually 23 to 29), low number of interorbital
136 Proceedings of the Biological Society of Washington
scales, low number of gular scales, scales bordering mental posteriorly
variable by population but modally 3 on Long Island (see detailed
discussion); area between enlarged lateral scales and ventrals (ventro-
laterally) lacking transitional scales, the enlarged laterals abutting
directly on the ventrals at the ventrolateral fold; dorsal colors most often
combinations of gray and black (although tans and browns occur in
some samples), and pale bands on unregenerated tails with black dots,
occasionally fused into short black dashes or longitudinal streaks, very
bold and clear against the white band color.
Description of holotype: An adult male, left hemipenis extruded,
snout-vent length 81 mm, tail about 92 mm, complete and unregenerated;
enlarged dorsal scale rows 21, ventral scales 28, interorbital scales 13,
gular scales 20, first toe lamellae 12, fourth toe lamellae 13, 3 scales
between sublabials and bordering the mental. Dorsum in life very pale
gray, almost white, with very scattered black dots on the dorsal scales;
head concolor with back, but with a broad white preocular band across
the snout (including the lores anterior to the eyes); postocular stripe
absent, but a white postocular line present from the posterior upper
margin of the eye across the temples; dorsal crossbanding barely evident,
the grayish band remnants separated by narrowed bands of very pale ©
gray to white; limbs gray, vaguely marbled with darker and with some
still darker gray dots on the hindlimbs; tail with eight “dark” bands,
the more proximal ones pale gray, grading to darker gray distally; pale
tail bands white, with scattered jet black dots enclosed therein, the
black dots occurring also on the “dark” bands as well; venter in life
pale ochraceous.
Variation: Combined data for all (44) specimens of T. a. warreni
are: maximum snout-vent length in males 92 mm (ASFS V8588, Long
Island) in females 88 mm (UF/FSM 23103, Warderick Wells Cay);
enlarged dorsal rows 16-24 (mean 19.6, mode 19), ventrals 23-31
(26.8), interorbitals 11-16 (13.4), gulars 14-23 (18.6), first toe lamellae
12-16 (13.1), fourth toe lamellae 11-17 (13.4).
The scales between the sublabials and bordering the mental show the
following pattern. On Long Island, the mode is 3 (eight of 11 specimens;
range 0-3). On Warderick Wells Cay, the mode is 1 (seven of 10
specimens; range 0-2). On the Allan’s Harbour Cays, there is a weak
mode of 2 (eight specimens) with 0 having an almost equal incidence
(seven specimens; range 0-3). The single Andros specimen has 2 scales
in this position, the Eleuthera specimen 1, and the two Great Ragged
specimens have counts of 1 and 2. Combining data for all Bahaman
specimens, nine specimens have counts of 0, 12 have counts of 1, 13
have counts of 2, and 10 have counts of 3. It is obvious that there is
a great degree of modal variation in this count, depending on the island
involved. Of great interest is the fact that, within the Exuma Cays, the
Allan’s Harbour Cays are almost bimodal (0 and 2), whereas Warderick
Wells Cay some 50 miles (80 km) to the southeast has a strong modality
of 1. Admittedly in all cases the samples are rather small, but the
Variation in New World Lizard ISi7
evidence for interisland variation seems conclusive. It is of course con-
ceivable that, when much additional material is collected, it will be
possible to show that there is more than one subspecies of T. americana
on the Bahama Islands; the material presently available does not suggest
this.
The basic colors of T. a. warreni are grays and blacks, although some
specimens, even from Long Island, were recorded as having tans and
browns in life. Except in small lizards (ASFS V10705, for instance),
the dorsum does not show the dark crossbands which customarily are
demonstrated by T. a. americana. If these crossbands are present, they
are usually faint (ASFS V8590) and the interband pale bars are fairly
broad but not too well defined. The dark postocular stripe is brown to
black and is often fragmented and occasionally absent, as in the holotype;
if the latter is the case, the white line dorsal to the postocular stripe
remains as a fairly conspicuous cephalic feature. The most common
dorsal condition is the presence of more or less scattered black speckling,
often arranged into rather vague short lines or dashes, over the entire
dorsum; these scattered dorsal markings tend additionally to obscure
the dark dorsal crossbands. The head figure described for T. a. ameri-
cana is almost always absent in warreni; some specimens (ASFS V8588 )
show remnants of it. If the head figure is present, it is always very
broken and barely definable. The ventral color varies from pale ochra-
ceous to yellow or pinkish and is not tan as in T. a. americana. The
dark dotting on unregenerated tails of warreni is a conspicuous feature
of all adults. The general aspect is of a gray-and-white-banded tail with
scattered black dots, occasionally fused into short longitudinal but
equally dark dashes. Although these dots and/or dashes are about
equally distributed over the entire upper surface of the tail, they stand
out more boldly against the white rather than the gray bands. The
black dotting is most prominent after about the third basal dark band,
so that specimens lacking the major portion of the tail barely show this
feature and do not show it at all on the regenerated portion of the tail,
which is uniformly striate longitudinally above and below.
Comparisons: The simplest method for differentiating americana from
warreni is the examination of the lower sides above the ventrolateral
folds. In warreni, the enlarged laterals abut directly against the smaller
and smooth ventrals, whereas in americana, these two scale types are
separated by several rows of transitional scales. Some warreni may have
a few intermediate scales scattered throughout this transitional area,
but none has the four to eight rows which occur in americana. The two
subspecies differ in number of transverse ventrals: only two of 44 warreni
have counts above 30, whereas only six of 80 americana have counts
below 30. Mean numbers of ventrals are statistically different (ameri-
cana—33.5 + 1.01; warreni—26.8 + .60). Mere gross examination of
specimens of americana and warreni of comparable snout-vent lengths
will easily demonstrate this difference in almost all individuals; the
much larger ventrals of warreni are obvious. Likewise, the smaller
138 Proceedings of the Biological Society of Washington
number of gulars (mean 18.6) and the fewer interorbitals in warreni
(mean 13.4) differ significantly from comparable means in americana
(gulars 24.6, interorbitals 14.3). In general, warreni has larger scales
and thus has lower counts than americana.
The pattern and color differences are more subtle. The generally gray
to black colors of warreni differ from the tans to browns of americana,
although some warrent were recorded as being brown and tan. In
general, the dorsa of warreni are not heavily crossbarred as are those of
americana but are rather much paler and unicolor, with suppression of
the dark postocular line; some americana, however, lack obvious dorsal
pattern, notably those from the Isla de Pinos. The dorsum of warreni
is often longitudinally streaked or striate, whereas that of americana is
seldom so. The most easily determined color and pattern differences are
the black and white (warreni) versus brown and tan to buff (americana)
unregenerated tails; the presence of discrete black dots or dashes on the
tails of warreni distinguish it from the only vaguely streaked tails of
americana. Except for the last chromatic character, the two subspecies
overlap as far as pattern is concerned. This is all the more remarkable
since so many Antillean reptile subspecies are defined primarily on color
and secondarily on scutellogical details. The reverse is true in T. ameri-
cana, where the scales offer far better means of differentiation than do
color and pattern. Nevertheless, although difficult to quantify, there
are color and pattern differences between the two subspecies.
Remarks: The series of T. a. warreni from the type locality was col-
lected there by Warren in two visits. The earlier series was taken in
abandoned stone houses, the lizards resting during the day in crannies
and crevices between the stones; at night the lizards were actively
foraging on the house walls and rafters. On the second visit to these
houses, over two years later, Warren found them stacked to the ceiling
with cut wood, so that he was unable to secure lizards inside the houses
themselves. The specimen from east of Salt Pond on Long Island was
taken by Thomas in Cocos trash on the ground at the base of a coconut
tree. The two ASFS Leaf Cay lizards were secured around the edges
of limestone slabs in palm woods. Rabb and Hayden (1957: 23) col-
lected their series on Leaf Cay under large limestone slabs. Since Leaf
Cay is exceptionally well strewn with rocks and slabs, T. a. warreni is
doubtless abundant there.
The scattered nature of Tarentola records in the Bahamas surely does
not reveal the complete range of the species in that archipelago. Con-
sidering that the first Bahaman Tarentola was collected by G. Sabille
in 1900 and that no further material was collected in quantity until the
Rabb and Hayden specimens (1953), it is obvious that Bahaman Taren-
tola are not easily secured except under the most favorable circum-
stances. The facts that no recent collector has taken the species on
Andros and that only very recently was the Eleuthera specimen secured,
suggest that in some areas the species is uncommon. Certainly the
Bahamas are becoming better known herpetologically, but many details
Variation in New World Lizard 139
of distribution remain to be clarified. An interesting sidelight is that
Richard Thomas, aware that T. americana had been collected on three
of the Exuma Cays, made special effort to secure specimens on Great
and Little Exuma where he searched cave entrances and cliffs for these
animals—typical Cuban situations. He found no Tarentola on these two
islands in these situations; in fact, insofar as I know, no Bahaman
Tarentola have been taken in cave mouths or on adjacent cliffs, niches
which Tarentola frequents in Cuba. Possibly there are ethological or
ecological differences between the subspecies americana and warreni.
I know of no eggs of T. a. warreni having been collected and none of
the series is a hatchling; the smallest specimen has a snout-vent length
of 31 mm (UMMZ 118025).
Discussion
Tarentola americana has been shown to be composed of two popula-
tions, whose divergence has been primarily in scutellogical details rather
than color and pattern as is often the case with Antillean reptile sub-
species. There is a possibility that the Isla de Pinos Tarentola will
ultimately be shown to differ from their Cuban relatives, but present
collections are inadequate for assessment of the apparent chromatic and
pattern differences between these populations. Additionally, it is also
possible that more Bahaman specimens will demonstrate differences
between Tarentola throughout the Great Bank islands, although this
seems less probable to me than nomenclatural recognition of the Isla de
Pinos lizards.
Since T. americana is the only New World representative of an other-
wise Old World genus, it seems likely that the West Indies species has
long been in residence on Cuba. The apparent lack of differentiation
into recognizable subspecies on Cuba suggests also that T. americana
is extremely stable genetically. On the other hand, as far as the Bahamas
are concerned, T. americana is presumably one of the relatively recent
immigrants (from Cuba) into that archipelago; I (MS) have suggested
that the present Bahaman herpetofauna is composed of two elements,
one an old and relict fauna, occurring on islands south of the Crooked
Island Passage and some other islands on their own banks, and a rela-
tively new and recent fauna, derived principally from Cuba, whose
members are not, or only subspecifically, different from their Cuban
relatives. This newer herpetofauna arrived in the Bahamas during the
Pleistocene or post-Pleistocene, since it is generally agreed that the
Great Bank was completely submerged during the Pliocene and may
have been submerged, at least partially, during the Pleistocene. If so,
then the strong scale differences between americana and warreni have
had to evolve relatively recently and relatively quickly. Those Bahaman
Tarentola which are geographically closest to Cuba (specimens from
Great Ragged Island, removed about 75 miles—120 km—from the north
shore of Cuba) show no approach to T. a. americana in scalation nor
140 Proceedings of the Biological Society of Washington
pattern. Such differences are all the more striking when the uniformity
of T. a. americana is considered. Perhaps Tarentola is an old Bahaman
relict, which persisted on one of the outer Great Bank islands (such as
Eleuthera, where it still occurs today) which are part of the Bahaman
volcanic arc. Such a longer period (Pliocene ?) of separation from its
Cuban congeners might account for the differences in scutellation which
segregate T. a. warreni from the nominate subspecies.
LITERATURE CITED
AuLAyo DALMAu, Pastor. 1951. Especies herpetolégicas halladas en
Santiago de Cuba. Bol. Hist. Nat. Soc. Felipe Poey, 2(7):
106-110.
1955. Lista de los reptiles de Cuba. Museo Charles T.
Ramsden, Universidad de Oriente, mimeographed, pp. 1-29,
7 pls.
ALLEN, Ross, AND WILFRED T. Nemu. 1957. The gecko-like habits of
Anolis lucius, a Cuban anole. Herpetologica, 13(3): 246-
247.
Barsour, THomMas. 1914. A contribution to the zodgeography of the
West Indies, with especial reference to amphibians and —
reptiles. Mem. Mus. Comp. Zool., 44(2): 209-359, 1 pl.
1916. Additional notes on West Indian reptiles and amphib-
ians. Proc. Biol. Soc. Washington, 29: 215-220.
1930. A list of Antillean reptiles and amphibians. Zoologica,
11(4): 71-116.
AND CHARLES T. RAMSDEN. 1919. The herpetology of Cuba.
Mem. Mus. Comp. Zool., 47(2): 71-213, 15 pls.
Bume, Mario S. 1966. Reptiles de la Peninsula Hicacos. Poeyana,
ser. A, 21: 1-12.
1967. Lista de los anfibios y reptiles de Cuba. Torreia,
new ser., 1: 1-60.
CocuraNn, Doris M. 1934. Herpetological collections from the West
Indies made by Dr. Paul Bartsch under the Walter Rathbone
Bacon Scholarship, 1928-1939. Smithsonian Misc. Coll., 92
(7): 1-48.
Granpison, ALICE G. C. 1961. Preliminary notes on the taxonomy of
Tarentola annularis and T. ephippiata (Sauria: Gekkonidae).
Zool. Meded. Leiden, 38(1): 1-14, 1 Fig., 3 pls.
GuNDLACH, JUAN, AND WILHELM PETERS. 1865. Uber einige neue
Saugethiere . . . Amphibien . . . und Fische. . . . Monatsb.
Akad. Wiss. Berlin, “1864,” pp. 381-399.
Harpy, Jerry D., Jr. 1957. Observations on the life history of the
Cuban lizard Anolis lucius. Herpetologica, 13(3): 241-245.
KoopMAN, Kart F., AND RopoLtro RurpAt. 1955. Cave-fossil verte-
brates from Camaguey, Cuba. Breviora, Mus. Comp. Zool.,
46: 1-8.
Variation in New World Lizard 141
LovERIDGE, ARTHUR. 1944. Certain American geckos of the genus
Tarentola. Copeia, 1: 18-20.
Marrero, Levi. 1951. Geografia de Cuba. Alfa, La Habana, pp.
i-xvi, 1-736, 478 Figs.
Rass, GreorcE B., AND Exuis B. HAypEN, Jr. 1957. The Van Voast-
American Museum of Natural History Bahama Islands expedi-
tion. Record of the expedition and general features of the
islands. Amer. Mus. Novitates, 1836: 1-53, 15 Figs.
Ruma, Ropotro. 1957. A preliminary investigation of the ecology
and taxonomy of Cuban lizards. Year Book Amer. Philo.
Soc., pp. 256-258.
ScHWARTZ, ALBERT. MS. The geckos (Sphaerodactylus) of the south-
em Bahama Islands. Ann. Carnegie Mus.
AND Larry H. Ocren. 1956. A collection of reptiles and
amphibians from Cuba, with the description of two new
forms. Herpetologica, 12(2): 91-110.
STEJNEGER, LEONHARD. 1917. Cuban amphibians and reptiles collected
for the United States National Museum from 1899 to 1902.
Proc. U.S. Natl. Mus., 53: 259-291, 128 Figs.
142, Proceedings of the Biological Society of Washington
‘Ei Mte ZS
Vol. 81, pp. 143-150 core April 30, 1968
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
NEW TAXA OF WESTERN ATLANTIC
COLUMBELLIDAE (GASTROPODA,
PROSOBRANCHIA )
By Gerorce E. RApwIn
Smithsonian Predoctoral Intern, Division of Mollusks,
United States National Museum, Washington, D.C.
The Columbellidae is one of a number of prosobranch
families whose classification is much in need of careful
revision. The last monographic treatment of the family
(Kobelt, 1897) was extensive, but was essentially a catalogue
of species with no attempt to establish their possible inter-
relationships. In order to establish such relationships, a
taxonomic revision of the Columbellidae of the western At-
lantic has been undertaken as the present author's doctoral
dissertation (Radwin, unpublished ).
During the course of the above investigation, the colum-
bellid components of the mollusk collections of the Academy
of Natural Sciences of Philadelphia, the American Museum
of Natural History, the Museum of Comparative Zoology,
Harvard, and the United States National Museum have been
examined. Considering the lack of attention that has been
given in the past to the family in question, it is, perhaps, not
too surprising that five new taxa have been uncovered. These
include two new species, two new subgenera and one new
genus.
Rhombinella new genus
Type-species: Buccinum laevigatum Linné, 1758 (Figs. 2, 9)
Shell moderately large (12-20 mm) and buccinoid, spire acute and
moderately high, whorls strongly convex, and sutures impressed. Body
whorl large, aperture broad, apertural lip usually unthickened and,
rarely, weakly denticulate on its inner surface. Columella straight and
smooth, weak anal groove present, and siphonal canal very short. Shell
17—Proc. Bron. Soc. WasH., Vou. 81, 1968 (143)
144 Proceedings of the Biological Society of Washington
Fics. 1-5: 1. Anachis (Costoanachis) fenneli, shell. 2. Rhombinella
laevigata, shell. 3. Anachis (Suturoglypta) pretrii, shell. 4. Anachis
(Parvanachis) rhodae, shell. 5. Anachis (Parvanachis) obesa, shell.
surface smooth with a dull buff-colored periostracum. Shell straw-
colored with axial red, brown, and white flammulations whose precise
pattern varies greatly from one specimen to another.
The radula is typically stenoglossate with a 1-1-1 formula. The
median teeth are flat, sub-rectangular plates, each of which is flanked
by a pair of lateral teeth. The lateral teeth are heavy, sub-rectangular
plates with three large cusps and five to seven smaller denticles appended
to the most proximal of these.
Remarks: The type species of this monotypic genus, in many cases,
has been assigned to Nitidella Swainson, 1840. There are numerous
distinctions regarding both shell and radula which may be used to
separate these two genera.
The aperture in Nitidella nitida (Fig. 7), the type species of Nitidella,
is much narrower than that of Rhombinella. The apertural lip of Nitidella
is thickened and denticulate, in contrast to the typically thin, non-
denticulate apertural lip of Rhombinella. The sutures in Nitidella are
New Columbellid Snails 145
very shallow imparting a flat-sided appearance to its rather low spire.
Those of Rhombinella, on the other hand, are impressed, imparting a
buccinoid appearance to its moderately high spire.
The radular distinctions here are even more striking than those of the
shell features (see Figs. 8, 10). The lateral radular tooth of Nitidella
comprises a sharp distal cusp; a broad, blunt medial cusp; and a long
flat proximal plate. The lateral tooth of Rhombinella, in contrast, com-
prises a sharp distal cusp, a broad second cusp ending in a sharp point,
a broad third cusp bearing five to seven sharp denticles, and a flat
proximal plate which is shorter than that of Nitidella. The lateral teeth
of both genera have a basal extension, bent into a plane at right angles
to that of the cusps. In addition to the above, the body of the lateral
tooth of Rhombinella is notably broader than that of Nitidella.
The only western Atlantic species assigned to Rhombinella is the type-
species, R. laevigata (Linné). There is also a subspecies, R. laevigata
hendersonii Dall, 1908, found on the southwestern coast of Cuba. It
may be distinguished from the nominate subspecies by its higher, more
acute spire; duller coloration; and the fact that the third cusp of its
lateral radular tooth always bears five denticles, whereas, that of the
nominate subspecies consistently bears seven denticles.
Anachis (Suturoglypta) new subgenus
Type-species: Columbella pretrii Duclos, 1846 (Fig. 3)
Shell small (4-8 mm), and fusiform. Spire high and acute, whorls
generally almost flat-sided, sutures squarely incised. Body whorl mod-
erately small (less than % total shell length), aperture narrow, apertural
lip thickened and denticulate on its inner surface. Columella slightly
to moderately bent and nondenticulate. Sculpture predominantly of
axial ribs, distinctly raised and almost square in cross-section. Color
generally white with brown flammules in many specimens. The radula
is typically stenoglossate with a flat, sub-rectangular median plate;
flanked by a pair of sigmoid, distally-bicuspid lateral teeth.
Remarks: This group shows remarkable variability at the species level
with regard to the number and spacing of the axial ribs. The three
species here assigned to Anachis (Suturoglypta): A. (S.) albella (C. B.
Adams, 1850), A. (S.) iontha (Ravenel, 1861), and A. (S.) pretrii
(Duclos, 1846) have been considered by many authors to be a single
species. They are, nevertheless, distinct species distinguished by slight
but consistent shell and radular differences.
The characters which are most diagnostic in distinguishing Anachis
(Suturoglypta) from other Anachis groups are the distinctly fusiform
shell; the strong, square-cut axial ribs; and the squarely incised sutures.
Anachis (Parvanachis) new subgenus
Type-species: Buccinum obesum C. B. Adams, 1845 (Fig. 5)
Shell small (5-10 mm) and obese. Spire moderately high and acute,
whorls generally flat-sided with incised sutures. Length of body-whorl
146 Proceedings of the Biological Society of Washington
10
o.imm
Fics. 6-10: 6. Anachis (Costoanachis) fenneli, dissociated radula
teeth. 7. Nitidella nitida, shell. 8. Nitidella nitida, radular dentition. 9.
Rhombinella laevigata, shell. 10. Rhombinella laevigata, radular denti-
tion. Note: Scale below figure 10 applies to all figures of radulae.
New Columbellid Snails 147
approximately equal in length to that of spire. Apertural lip markedly
thickened and denticulate on its inner surface. Siphonal canal short to
moderate in length and constricted, columella straight or slightly bent
and denticulate. Shell sculpture generally consisting of prominent axial
ribs, in most species crossed by spiral cords.
The radula has a 1-1-1 formula with a flat sub-rectangular median
plate, flanked by a pair of sigmoid, tricuspid lateral teeth. The two
most distal cusps of the lateral tooth are similar to those of most other
columbellid genera. The proximal cusp, however, is strongly down-
hooked; a feature which is characteristic of the subgenus.
Remarks: The new subgenus proposed here is intended to include
those small, stout, Anachis-like columbellids with inflated body whorls
and heavily thickened apertural lips. The shell proportions in this
subgenus are considerably different from those in Anachis s. s. or
Costoanachis as they impart a foreshortened or “dumpy” appearance to
the shell. In addition, the strongly down-hooked proximal cusp of the
lateral radular tooth serves to distinguish Parvanachis from Costoanachis
Sacco, another Anachis-like group, to which A. obesa has been assigned
by Gardner (1948, p. 229) and others. Other western Atlantic species
of Parvanachis include Anachis (Parvanachis ) isabellei (Orbigny, 1839),
A. (Parvanachis) ostreicola (Sowerby, 1882), and A. (Parvanachis )
rhodae n. sp.
Anachis (Parvanachis) rhodae new species (Fig. 4)
Holotype: Mus. Comp. Zool., Harvard, 261479, L: 7.7 mm, W: 3.2 mm
Shell moderately small (6-8 mm) and obese. Spire high (1% total
shell length) and acute; whorls slightly convex, each “set-in” from the
one below it, imparting a terraced appearance; sutures impressed. Body
whorl large and inflated, apertural lip ventricose and strongly denticulate
on its interior surface. Columella strongly denticulate and sharply bent
with a small but heavy callus, siphonal canal short and sharply bent,
slight anal groove present. Sculpture of low axial ribs and several
obsolete spiral grooves on the anterior part of the body whorl. Color
yellow-white with varying amounts of chestnut-brown blotches. Proto-
conch of one full, bulbous, translucent white whorl. The radula of this
species is unknown. Despite the apparent freshness of most of the
thirty-five shell specimens in the type lot, none of them contained any
animal material.
Remarks: A hitherto unsuspected new species inhabiting the northern
coast of Hispaniola, this species, named for the author's wife Rhoda, is
known from only a single locality, Puerto Plata, Dominican Republic.
Two lots from this locality were found in the collection of the Mollusk
Department of the Museum of Comparative Zoology, Harvard. Fourteen
paratypes (USNM 677645) have been deposited in the collection of
the Division of Mollusks, U.S. National Museum.
The present species strongly suggests a link between Strombina
148 Proceedings of the Biological Society of Washington
(Sincola) (Olsson and Harbison, 1953, pp. 230-31) and Anachis (Par-
vanachis). It lacks the typical hump and the heavily callused and
emarginate aperture of Strombina, but otherwise might be considered
quite close to Sincola. On the other hand, the overall shell form and
the appearance of the apertural lip strongly resemble those of A. obesa,
the type species of Parvanachis. The present placement of A. rhodae
must remain provisional until the radula and the soft parts have been
examined.
Anachis (Costoanachis) fenneli new species (Fig. 1)
Holotype: USNM 539122. L:6.3 mm, W: 2.8 mm
Shell small (5-7 mm). Spire acute and slightly more than one-half
total shell length, spire whorls slightly convex with weakly impressed
sutures. Body whorl expanded, angular, and asymmetrical, the left side
slightly more expanded than the right. Aperture broad, apertural lip
denticulate on its inner surface, a distinct anal sinus present. Columella
straight and weakly denticulate. Sculpture consisting of broad, strong
axial ribs crossed by fine, shallow, spiral grooves. Yellowish-white
ground color with numerous anastomosing brown blotches. Protoconch .
of almost three full, glossy-brown whorls. The radula (Fig. 6) is
typically stenoglossate with a 1-1-1 formula. The median tooth is a
flat, sub-rectangular plate; flanked by a pair of sigmoid, bicuspid lateral
teeth. In details the laterals are very similar to most other Coastonachis
species. The only notable difference is the smaller size of the entire
lateral tooth.
Remarks: This new species is named in honor of Dr. William E.
Fennel and is known from a single lot of about thirty-five specimens, all
collected alive by Dr. D. M. Cochran near Nictheroy, State of Rio de
Janeiro, Brazil in April, 1935. Its uniqueness is evident in its unusually
large, angular body whorl, its prominent axial, and distinctive spiral
sculpture, its color pattern, and its multiwhorl protoconch. In addition,
some minor radular distinctions have been noted.
This is probably the “Anachis sparsa Reeve” treated by Marcus and
Marcus (1962, p. 339, pl. 1, Fig. 2). Unfortunately, no specimens of
this species were obtained from Dr. Marcus because of difficulties in
the mailing of parcels from Brazil. Nevertheless, the figures of its shell
and radula in the above-mentioned paper, plus the fact that the range
of A. sparsa is not known to extend to South America, leaves little doubt
that the Marcuses could not have had that species.
LITERATURE CITED
Apams, C. B. 1845. Spec. Nov. Conch. Jamaica Report. Synop. Proc.
Bost. Soc. Nat. Hist. 2, pp. 1-17.
1850. Notes on the synonymy of certain marine shells.
Contrib. to Conch. 1(4), pp. 53-56.
‘>
New Columbellid Snails 149
Daut, W. H. 1908. A new West Indian Nitidella. Nautilus 22, pp.
31-32.
Ductos, M. 1846. (in Chenu) Illustrations Conchyliologiques. 4, pls.
1-27, Paris.
Garpner, J. 1948. Mollusca from the Miocene and Lower Pliocene
of Virginia and North Carolina. Part 2. Scaphopoda and
Gastropoda. U.S. Geol. Sury. Prof. Paper # 199B, pp. 179-
310, pls. 24-38.
Kospetr, W. 1897. (in) Martini and Chemnitz Conchylien-Cabinet.
3 (Id), 343 p., pls. 1-44, von Bauer and Raspe, Nuremberg.
LinnE, C. von. Systema Naturae. ... 10° edition 1, 824 p., Stockholm.
Marcus, E. anp E. Marcus. 1962. Studies on Columbellidae. Bol.
Fac. Fil. Cién. Letr. Univ. S40 Paulo # 261, Zoologia # 24,
pp. 335-402, 8 pls.
Ousson, A. AND H. Harsison. 1953. Pliocene Mollusca of S. Florida.
Acad. of Nat. Sci. Phila. Monogr. # 8, 457 pp., 65 pls.
Orsicny, A. 1839. Voyage dans l’Amerique Meridionale Atlas, livr.
44, pls. 61, Figs. 18-21, Paris.
Rapwin, G. 1968. A systematic revision of the family Columbellidae
in the western Atlantic. Unpublished doctoral dissertation
George Washington Univ. 255 p., 54 pls.
RAVENEL, E. 1861. Descriptions of new recent shells from the coast
of South Carolina. Proc. Acad. Nat. Sci. Phila. pp. 41-42.
SowerBy, G. B. 1882. Descriptions of new species of shells in the
collection of Mr. J. Cosmo Melvill. Proc. Zool. Soc. Lond.
pp. 116-121.
SwaInson, W. 1840. A treatise of Malacology 419 p., Longman et al.
London.
150 Proceedings of the Biological Society of Washington
£L Die ose
Vol. 81, pp. 151-154 \ April 30, 1968
PROCEEDINGS.
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
STREPTOSYLLIS LATIPALPA, NEW SPECIES
(POLYCHAETA, SYLLIDAE) FROM PUGET
SOUND (WASHINGTON)!
By Karu BANSE
University of Washington, Seattle
Among, the polychaetes collected from intertidal sand near
Seattle, Washington, in 1956 to 1957 by Wieser (1959), there
is a previously overlooked, well-preserved specimen of an
undescribed species of Streptosyllis Webster and Benedict, a
genus not yet reported from the Pacific Ocean. The exact
locality of collection can no longer be determined. The holo-
type is deposited in the United States National Museum
(USNM 36509).
Genus Streptosyllis Webster and Benedict
Streptosyllis latipalpa new species
Description: The holotype has 26 setigers and is 1.8 mm long (with-
out cirri). The greatest width is 0.27 mm with parapodia, and 0.12 mm
without. The palps, which are large for a member of this genus, are
fused at the base and rounded anteriorly (Fig. 1A). The median
antenna inserts in the middle, the lateral antennae insert near the anterior
margin of the prostomium. Eyes cannot be distinguished. The peri-
stomium is well separated from the prostomium and carries two pairs of
tentacular cirri. These cirri, as well as the dorsal cirri of the following
segments, tend to be slightly thicker distally than proximally. Usually
they are irregularly pseudoannulated, although occasionally, they are
truly annulated. The ventral cirri of the first four setigers are short,
whereas those of the remaining setigers are slightly longer than the
parapodia (Fig. 1B). The pharynx is short and straight (not everted )
and apparently without a tooth, but the specimen is fairly opaque. The
proventricle extends from the anterior border of the fourth setiger into
1 Contribution No. 460 from the Department of Oceanography, University of
Washington, Seattle, Washington 98105. The preparation of this paper was sup-
ported by National Science Foundation Grant No. GB-4902 to K. Banse and M.
M. Pamatmat.
18—Proc. Brox. Soc. WasuH., VoL. 81, 1968 (151)
152 Proceedings of the Biological Society of Washington
c ca.0.01mm(C-G)
0.05mm
_
Fic. 1. Streptosyllis latipalpa, new species: A, Anterior end, dorsal
view; setae schematic; B, twelfth parapodium; setae schematic; C,
acicula from third parapodium; D, compound seta from third para-
podium; E, compound seta from twelfth parapodium; F, dorsal simple
seta from third parapodium; G, dorsal simple seta from twelfth para-
podium.
the eighth setiger. There is only a median cirrus, about four times as
long as wide, on the pygidium; long paired cirri, common to other
members of the genus, may have fallen off. Heavy aciculae (Fig. 1C)
occur in setigers 2 to 5. In the first four setigers there are about a dozen
compound setae per parapodium. The shafts seem to end in three knobs
opposite to the insertion of the very short blades (Fig. 1D). From the
fifth setiger onward, the blades of most of these setae become longer
(two to five times as long as shown in Fig. 1D), until in the ninth or
tenth setiger they are about as long as shown in Fig. 1E. In addition,
one or two setae in the last setigers have very large blades of about 35 u
length. In the 12th setiger the shafts of all setae apparently end only in
one tip each, opposite the insertion of the blades. The blades are bifid,
and at least some also have a spur below the tips (Fig. 1E). A dorsal
simple seta is present from the first setiger, which in anterior segments
is slightly bent at the tip (Fig. 1F). It is straight in median and posterior
New Species of Polychaete 153
segments (Fig. 1G). In all, there are about half a dozen setae per
parapodium in the posterior region.
The name refers to the broad palps.
Diagnosis: A species of Streptosyllis Webster and Benedict with
broad, anteriorly rounded palps, and heavy aciculae in setigers 2 to 5.
Differential Diagnosis: Six species of Streptosyllis are listed by Hart-
man (1959, 1965): S. arenae Webster and Benedict; S. bidentata South-
em; S. cryptopalpa Hartmann-Schréder; S. reducta Hartmann-Schréder;
S. varians Webster and Benedict; and S. websteri Southern. After
inspecting the type of S. varians, Southern (1914) and Pettibone (1963)
have pointed out that the animals described by Saint-Joseph (1895)
under this name must be a different form; Southern observed the
similarity with S. websteri.
The new species is similar to S. arenae, S. cryptopalpa, and S. websteri,
including the form described by Saint-Joseph (1895) as S. varians, in
that it has heavy aciculae in setigers 2 to 5. It is well distinguished
from these species, apart from details of setation, by its broad, rounded
palps visible from above. S. varians likewise has large palps which,
however, are pointed anteriorly; also, it has heavy aciculae in about 20
setigers. There are no palps and heavy aciculae in S. reducta. The latter
species, as well as S. cryptopalpa, has a tooth in the pharynx which is
not reported for the species described earlier.
LITERATURE CITED
Hartman, O. 1959. Catalogue of the Polychaetous Annelids of the
World. I. Occas. Papers, Allan Hancock Found. Publ., 23:
1-353.
. 1965. Catalogue of the Polychaetous Annelids of the World.
Supplement 1960-1965 and appendix. Occas. Papers, Allan
Hancock Found. Publ., 23 (Suppl.): 1-197.
PETTIBONE, M. H. 1963. Marine polychaete worms of the New Eng-
land region. 1. Aphroditidae through Trochochaetidae. Bull.
U.S. Nat. Mus., 227: 1-356.
SAINT-JosEPH, A. DE. 1895. Les annélides polychétes des cétes de
Dinard. IV. Ann. Sci. Nat. Zool., (7) 20: 185-272.
SouTHERN, R. 1914. Clare Island Survey. 47. Archiannelida and
Polychaeta. Proc. R. Irish Acad., 31: 1-160.
Wieser, W. 1959. Free-living nematodes and other small invertebrates
of Puget Sound beaches. 179 pp., University of Washington
Press, Seattle.
154 Proceedings of the Biological Society of Washington
iG OCT SF
Vol. 81, pp. 155-160“ x_“/Bpy April 30, 1968
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
DAECTOR SCHMITTI, A NEW SPECIES OF
VENOMOUS TOADFISH FROM THE PACIFIC
COAST OF CENTRAL AMERICA
By Bruce B. CoLLETTE
Bureau of Commercial Fisheries Ichthyological Laboratery,
U.S. National Museum, Washington, D.C. 20560
In my recent review (Collette, 1966) of the venomous
toadfishes (Batrachoididae, Thalassophryninae ), I tentatively
referred to a specimen from Costa Rica as a variant of Daector
reticulata (Gunther) although I believed that it might rep-
resent an undescribed species. I did not describe it because
I had only the single specimen, which closely resembled D.
reticulata in color pattern. A second, larger, specimen, from
Panama has come to my attention and I am now convinced
that these two fish represent an undescribed species. I take
great pleasure in naming the new species in honor of the
energetic collector of the holotype—Dr. Waldo L. Schmitt.
Daector schmitti new species
(Fig. 1)
Diagnosis: A species of Daector most similar to D. reticulata but
differing in having fewer second dorsal rays (22 vs. 25-27); anal rays
(21 vs. 24-26); and caudal vertebrae (23 vs. 26-28). The filaments on
the head of D. schmitti extend farther laterally than in D. reticulata—
beyond the infraorbital lateral line canal, instead of to the canal. D.
schmitti has a color pattern similar to that of the smaller sizes (80 mm)
of D. reticulata but loses the reticulations and develops a spotted pattern
reminiscent of the Atlantic species Thalassophryne maculosa Giinther at
larger sizes (130 mm). Precaudal vertebrae 8. Pectoral fin glands 6.
Comparisons: D. schmitti has a much more extensive distribution of
filaments on the head and anterior part of the body than does D.
reticulata although the filaments are larger in D. reticulata (Fig. 2).
Filaments cover a greater area in larger specimens of both species. They
extend farther laterally in D. schmitti—below the infraorbital canal on
19—Proc. Bion. Soc. Wasu., Vou. 81, 1968 (155)
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*(qysue] piepuvys Jo osejzue01ed ul) DyDjNOIAa4 ‘Gq YUMA IW4pWYIS 40J0aDq Jo UOsSTIedUIOD oLjoULOYdIOJYN—'[ ATA],
Fic. 1. Daector schmitti. Left, SU 14949, paratype, 80.1 mm SL,
Golfo de Nicoya, Costa Rica. Right, USNM 144869, holotype, 130 mm
SL, Secas Islands, Panama.
the head, and below the lateral line on the body in the region of the
second dorsal fin origin. In D. schmitti, the filaments cover much of the
membrane over the dorsal spines and extend posteriorly as far as the
eighth ray in the second dorsal fin. In eight large (185-252 mm SL)
158 Proceedings of the Biological Society of Washington
Fic. 2. Dorsal view sf heads of Daector reticulata (left, USNM
81698) and D. schmitti (right, USNM 144869) showing distribution of
filaments.
specimens of D. reticulata, the filaments extend posteriorly no farther
than the second or third ray in the second dorsal fin.
D. schmitti agrees with both D. reticulata and D. gerringi (Rendahl)
in having one fewer ray in the second dorsal fin than the number of
caudal vertebrae. D. dowi (Jordan and Gilbert) has the same number
of second dorsal fin rays as caudal vertebrae.
Morphometrically, D. schmitti has a longer, wider head, greater
interorbital distance, longer pelvic fins, and greater snout—pectoral and
snout—-pelvic distance than does D. reticulata (Table 1).
Types: Holotype—U. S. Nat. Mus. 144869, 130 mm standard length;
Panama, Secas Islands, from channels in coral tidal flat; 6 February
1935; VELERO 434-35. Paratype—Stanford Univ. 14949, 80.4 mm; Costa
Rica, Golfo de Nicoya, Negritos Island, tidepools; 13 February 1947;
M. B. Schaefer.
Discussion: The low number of caudal vertebrae and anal and second
dorsal rays tend to bridge the gap between Daector which has high
counts and Thalassophryne which has low counts. D. schmitti, however,
agrees with D. reticulata, D. gerringi, and D. dowi in the three diagnostic
characters of the genus: possession of discrete pectoral fin glands (see
Fig. 1 in Collette, 1966), 8 precaudal vertebrae instead of 7, and a
submarginal stripe in either the second dorsal or anal fin or both.
q
-
CS ed
New Central American Toadfish 159
Dr. William A. Bussing, University of Costa Rica, has informed me
that he has not found any toadfishes in the many tidal pool and inshore
rotenone collections made between Bahia Culebra and Punta Mala. He
suggests that D. schmitti may be restricted to island shores.
Acknowledgments: I am grateful to Mr. Robert E. Trist, Division of
Fishes, U.S. National Museum, for calling the holotype to my attention
and to Dr. Warren C. Freihofer, Stanford University, for lending me the
paratype of D. schmitti. The figures were drawn by Mrs. Mildred H.
Carrington. Drs. Daniel M. Cohen and Victor G. Springer have com-
mented on the manuscript.
Addendum: After this manuscript was sent to the printer, Mr. Robert
J. Lavenberg of the Los Angeles County Museum discovered a third
specimen of D. schmitti: LACM 22360, 20.0 mm SL; Costa Rica, Port
Culebra, off South Viradore Islands, 10°35’N, 85°43’W; 10 fms. over
sand and shells; 25 February 1934; VeLERo 257-34. It agrees exactly
with the other two types in counts: D II, 22; A 21; vertebrae 8 + 23 =
31. Like the holotype, this paratype was also collected by Dr. Schmitt
from an island while on one of the Velero III expeditions (Fraser, 1943a,
1943b).
LITERATURE CITED
CoLLEeTTE, Bruce B. 1966. A review of the venomous toadfishes, sub-
family Thalassophryninae. Copeia 1966 (4): 846-864.
FRASER, C. McLEAN. 1943a. General account of the scientific work of
the Velero III in the eastern Pacific, 1931-41. Part II.
Geographical and biological associations. Allan Hancock
Pacific Exped. 1(2): 49-258.
1943b. Ibid. Part III. A ten-year list of the Velero III
collecting stations. Ibid. 1(3): 259-445.
160 Proceedings of the Biological Society of Washington
Ny
Wo OO7S
Vol. 81, pp. 161-172 April 30, 1968
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
PRELIMINARY DESCRIPTIONS OF TWO NEW
SPECIES OF THE BATHYPELAGIC SQUID
BATHYTEUTHIS (CEPHALOPODA: OEGOPSIDA )
By CiypE F. E. Roper
Division of Mollusks, Smithsonian Institution,
Washington, D.C. 20560
Two new species were discovered during the course of a
study on the systematics and distribution of the world-wide
deep-sea squid genus Bathyteuthis. Both species occur in the
eastern Pacific Ocean, one from the tropical waters of the Bay
of Panama and the other from the cooler waters off southern
California. The species are briefly described here so that the
descriptions and names will be available for use by workers
currently engaged in regional surveys, e.g., that in preparation
by R. E. Young, Institute of Marine Sciences, University of
Miami. Detailed descriptions will be presented in the more
comprehensive study (Roper, 1968).
I am grateful to the following persons for supplying speci-
mens: E.. Bertelsen (Carlsberg Foundation, Dana collections ),
G. L. Voss (Institute of Marine Sciences, University of Miami,
ELTANIN and Pittspury collections), R. E. Young (1.MS.,
VELERO collections from the University of Southern Califor-
nia). The ELTANIN material was collected under the sponsor-
ship of the Office of Antarctic Programs, National Science
Foundation. I wish to thank Constance Stolen for her accu-
rate preparation of the illustrations. R. E. Young, J. Rose-
water, and R. B. Manning kindly read the manuscript and
made valuable comments.
The standard measurement of size is mantle length, abbre-
viated ML. Abbreviations for ships are Exr. for ELTanin,
Pit. for Pittspury, D for DANa and V for VELERO.
20—Proc. Biot. Soc. WasH., Vou. 81, 1968 (161)
Piatt 1. Bathyteuthis bacidifera. Holotype, female, 37 mm ML,
Ext. 34. A. Dorsal view. B. Ventral view.
Two New Squid 163
Bathyteuthis bacidifera n. sp.
Plates 1-4, 7 G, H
?Benthoteuthis megalops, Chun, 1910, pp. 185-199, pls. 24-27 (pars;
station 221, 18 mm specimen only).—Pfeffer, 1912, pp. 325-331
(pars; using Chun’s description ).
MATERIAL STUDIED.
Sex ML, mm Ship Sta. Location Date Depth, m Gear
Holotype:
Q 37 ET. 34 07°47’S 81°23’W 7VI62 £683 10’ IKMT
Paratypes:
ce) 37 ExT. 34 07°47’S 81°23’W 7VI62 683 10’ IKMT
ce) 34 Ext. 54 18°23’S 72°39’W 16 VI62 1373 10’MW Beam
oS 28 Pin. 510 06°54’N 79°57’W 3 V 67 3182 40’ Otter
© 26 D1208 XIV 06°48’N 80°33’W 12 1 22 1550 $1501
1 Stramin tow-net of 150 cm diameter.
Diagnosis: Protective membranes on arms lacking; long, free, finger-
like trabeculae present; tentacles and clubs relatively long; suckers on
arms numerous; sucker rings with 18—34 protuberances; gills long, broad.
Description: Mantle short, broad, bullet-shaped; width about 50%
of length. Fins short, small, paddle-like, circular in outline, subterminal;
anterior and posterior fin lobes project well beyond bases of fins (pl. 1).
Funnel broad basally, tapers anteriorly (pls. 1B, 2A), extends to a level
between posterior margins of eye openings; bridles weak. Posterior end
of funnel groove with small median orifice (pl. 2F). (This unusual
structure is currently being investigated.) Funnel component of locking
apparatus simple, long, narrow; sulcus anteriorly deeper and narrower
than posteriorly (pl. 2A, B). Mantle component a simple, low ridge
(pl. 2A, C). Funnel organ an inverted Y with narrow, rounded pos-
terolateral limbs and broad, narrow anterior limb; a flat apical papilla
present (pl. 2E). Head short, broad with large, anterolaterally-directed
eyes. Eye openings circular; optic sinus lacking. Olfactory papilla
small. Nuchal folds lacking.
A small, flat, ovoid photophore embedded on base of each of dorsal
six arms (pl. 1A). Arms short, conical, not attenuate; ventral arms
shortest, others subequal. A fleshy web connects basal third of arms
(pl. 3A, B, F) except between ventral arms. Swimming keels present
on all arms. Protective membranes lacking proximally, but trabeculae
are modified as long, fleshy, blunt, finger-like cirri (pl. 3A, B), becom-
ing shorter, broader, more lobate distally; protective membranes normal
toward arm tips. Arm suckers small to minute, numerous, arranged
biserially basally, becoming 3- to 4-rowed distally (pl. 3A, B). Inner
rings of largest arm suckers 0.12-0.16 mm in diameter with 20-34
minute closely packed teeth grading from long and truncate distally to
stubby and knobby proximally; smaller suckers have fewer teeth (pl.
4B, C, D, F, G). Tentacles long, thin, muscular, 125%-150% of ML;
164 Proceedings of the Biological Society of Washington
PLATE 2. Bathyteuthis bacidifera. Holotype, B—F, and a paratype,
A, G, females, 37 mm ML, Etr. 34. A. Open mantle cavity. B, C.
Funnel and mantle components of locking apparatus. D. Nuchal car-
tilage. E. Funnel organ and funnel valve. F. Pore of funnel groove.
G. Gladius.
tentacular clubs long, narrow, undifferentiated into carpus, manus, or
dactylus (pl. 4A). Club suckers numerous, minute, closely-packed, in
8-10 rows. Largest sucker rings 0.08-0.10 mm in diameter with 8-10
minute, widely-spaced truncate to knob-like teeth (pl. 41). Protective
membranes absent; weak swimming keel present.
Buccal connectives attach to dorsal, dorsal, ventral, dorsal edges of
arms I-IV respectively (pl. 3F). Sucker rings from 7 buccal lappets
Two New Squid 165
Piate 3. Bathyteuthis bacidifera. Holotype, A, B, F, and a paratype,
C, D, E, females, 37 mm ML, Eur. 34. A. Brachial crown with buccal
membrane expanded. B. Right arm I. C. Upper mandible. D. Lower
mandible. E. Radula. F. Brachial crown showing connectives of the
buccal membrane.
0.08-0.10 mm in diameter with 8-12 minute, widely-spaced, truncate
teeth (pl. 4H). Buccal mass and beaks relatively small, beaks with
strong rostra, weak lamellae (pl. 3C, D). Radula with 7 transverse rows
of pointed teeth, 2 rows of marginal plates (pl. 3E). Rhachis of gladius
long, slender, with rolled, rod-like lateral edges; vane broad, thin, weak;
conus absent (pl. 2G).
166 Proceedings of the Biological Society of Washington
Puate 4. Bathyteuthis bacidifera. Holotype, female, 37 mm ML,
Ext. 34. A. Tentacular club. B, C. Inner sucker rings from Arm I. D,
E. Inner and outer sucker rings from Arm IJ. F. Inner sucker ring from
Arm III. G. Inner sucker ring from Arm IV. H. Buccal sucker ring.
I, J. Inner and outer sucker rings from left tentacular club.
Color maroon.
Cement body of spermatophore cigar-shaped with flaring lip anteriorly;
base of ejaculatory apparatus barrel-shaped (pl. 7G, H). Sperm mass
63%-66% of total length of spermatophore, cement body 17%-20%,
ejaculatory apparatus 15%—-20%. Hectocotylus absent. Gills long, broad.
Holotype: United States National Museum, No. 576148.
Yaoi
“a
¢
f
.
i
Two New Squid 167
Puate 5. Bathyteuthis berryi. Holotype, male, 49 mm ML, VELERO
8714. Ventral view.
168 Proceedings of the Biological Society of Washington
PLATE 6. Bathyteuthis berryi. Holotype, A, C-F, H, male, 49 mm
ML, VELERO 8714, and a paratype, B, G, juvenile, 19 mm ML, VELERO
10976. A. Arm I. B. Tentacular club. C-—F. Inner sucker rings from
Arms I-IV. G. Inner sucker ring from tentacular club. H. Buccal sucker
ring.
Two New Squid 169
Type locality: Off northern Peru at 07°47’S 81°23’W. USNS ELTanin
Sta. 34, 7 June 1962.
Distribution: Bathypelagic in the productive waters of Eastern Pacific
Equatorial Water Mass; possibly in the Indian Ocean Equatorial Mass
(based on Chun’s (1910) single specimen).
Etymology: The specific name bacidifera is a neo-Latin word meaning
“bearing little rods”; this is derived from the old Latin baculum, a staff,
stick or rod, the diminutive -idiwm, and -fer, a suffix meaning bear,
carry. The name alludes to the outstanding characteristic of the species,
the rod-like trabeculae.
Bathyteuthis berryi n. sp.
Plates 5—7A-F
MATERIAL STUDIED
Sex ML, mm Ship Sta. Location Date Depth, m1
Holotype:
of 49 V. 8714 33°14’45’N 118°37’20”W 7 VI 63 1200
Paratypes:
of 23 V. 10540 29°05’04’N 118°12’00”W 6 IV 65 1300
of 20 V. 10377 33°25’/00”N 118°50’45”W 24 II 65 1100
ie) 19 V. 10976 32°35’00”N 120°35’06”W 17 II 66 1300
1 Estimated depths of capture. All specimens were captured by a 10’ Isaacs-Kidd
midwater trawl.
Diagnosis: Protective membranes on arms present, well developed and
fleshy proximally, no free trabeculae; suckers on arms extremely numer-
ous; sucker rings with 10-14 protuberances; gills long and broad.
Description: Mantle very plump, robust, bullet-shaped; width 50% of
length. Fins rounded, short, separated, with free anterior and posterior
lobes (pl. 5). Funnel very large, prominent; extends to level between
anterior margins of eye openings. Small pore at base of funnel groove.
Funnel component of locking apparatus simple, with shallow sulcus;
mantle component with low ridge. Funnel organ Y-shaped with short,
broad limbs; apical papilla spatulate. Head long, narrow. Eyes large,
directed anterolaterally; eye openings circular, lacking optic sinus.
Olfactory papilla minute. Nuchal folds absent.
A single, small, simple photophore embedded at base of each of dorsal
six arms, most readily seen in juvenile and larval specimens. Arms long,
slender, attenuate, joined basally by a deep web except between ventral
arms; adult arm formula 4=3=2=1. Swimming keels low on arms
I-III, well-developed as tentacular sheath on IV. Thick, fleshy, ruffle-
like protective membranes well-developed basally on arms, diminishing
distally; no free trabeculae. Arm suckers small to minute, extremely
numerous (275 on dorsal six arms, 150 on arms IV of holotype); sucker
rows increase from 1 proximally to 3-4 along mid-portion of arms (pl.
6A). Inner sucker rings with 10-14 small, low, rounded or subtriangular,
knob-like teeth (pl. 6C-F). Tentacles missing from holotype; tentacles
170 Proceedings of the Biological Society of Washington
Puate 7. Bathyteuthis berryi, A-F. Holotype, male, 49 mm ML,
VeELERO 8714. B. bacidifera, G-H, Paratype, male, 28 mm ML, PiLts-
BuRY 510. A. Radula. B. Gladius. C. Upper mandible. D. Lower
mandible. E. Spermatophore. F. Enlarged section of spermatophore.
G. Spermatophore. H. Enlarged section of spermatophore.
available only from small specimen (19 mm ML, VELERO 10976); long,
robust, with short, unexpanded, undifferentiated club with 7-8 transverse
rows of numerous minute suckers (pl. 6B); inner rings smooth, scalloped,
or with small, low, subtriangular teeth (pl. 6G).
Two New Squid wal
TaBLE 1. Distinguishing features of the three species of Bathyteuthis.
Character abyssicola bacidifera berryi
Free trabeculae absent present absent
Protective membranes present absent present
Arm Suckers? few (100) numerous (150) extremely
numerous (275)
Sucker Ring
Dentition (Arms ) 8-18, truncate 18—34, truncate 10-14,
subtriangular
Arms short, blunt short, blunt long, attenuate
Gills short, narrow long, broad long, broad
Spermatophore 68-72; 6-8; 63-66; 17-20; 72; 8; 20
proportions? 20-25 15-20
Tentacles and short long missing from
clubs material
1 The numbers in parentheses represent the approximate number of suckers on
each of the six dorsal-most arms from specimens of about the same size (49 mm
ML).
2 The size of the sperm mass, cement body and ejaculatory apparatus respectively,
expressed as a percentage of the total length of the spermatophores.
Buccal lappets possess 4-6 small suckers with about 10 small, low,
papilla-like teeth (pl. 6H). Buccal connectives attach dorsally to arms
I, II, IV, ventrally to III. Beaks small, with strong, curved rostra, weak
lamellae (pl. 7C, D). Radula with 7 transverse rows of teeth, 1-2 rows
of platelets (pl. 7A). Rhachis of gladius long, slender; vane very broad,
thin, weak; conus absent (pl. 7B).
Color maroon.
Cement body elongate, vase-shaped with collar at junction with short,
bell-shaped end of spiral filament (pl. 7E, F). Sperm mass = 72% of
spermatophore length, cement body = 8%, ejaculatory apparatus =
20%. Hectocotylus absent. Gills long, broad.
Holotype: University of Southern California. U.S.C. Hancock col-
lections, AHF cephalopod type No. 10.
Type locality: Catalina Basin, 10.9 miles SSW of West End Light,
Catalina Island at 33°14’45”N 118°37’/20”W. VeELERO Sta. 8714, about
1200 m.
Distribution: Bathypelagic in the waters off Southern California.
Etymology: The specific name, berryi, is given in honor of Dr. S.
Stillman Berry who has contributed a lifetime of study to malacology
and teuthology.
Discussion: The addition of two new species brings to three the num-
ber of species that belong to Bathyteuthis. The originally described
species, B. abyssicola Hoyle, 1885, has a broad geographic distribution
in the bathypelagic zone of the major oceans of the world; distinct
172 Proceedings of the Biological Society of Washington
geographic populations occur in the Atlantic, tropical eastern Pacific,
and Antarctic Oceans. The species is particularly abundant in the
nutrient-rich waters of the Antarctic Ocean. B. bacidifera is sympatric
with B. abyssicola in the Panama Bay region of the tropical eastern
Pacific, while B. berryi occurs alone in the colder water mass to the
north off California. Systematic problems at the familial and generic
levels, geographical variation, and environmental aspects of geographic
and bathymetric distribution are discussed in Roper (1968).
The three species of Bathyteuthis are distinguishable by several
features, the most prominent of which are listed in Table 1.
The most striking and easily recognized character of bacidifera is the
presence of long, finger-like trabeculae on the arms that have no inter-
connecting protective membrane. This feature is apparent even on the
smallest larva available (6 mm ML) and readily separates the species.
Both abyssicola and berryi possess thick, fleshy protective membranes.
Considerable variation exists in the membranes, but they are always
present and connect unmodified trabeculae.
B. berryi is most readily distinguished from abyssicola by the extreme
abundance of suckers on the arms and by long, wide gills. (The signif-
icance of gill size is discussed in Roper, 1968). The arms of the holotype
of berryi are 5-7 mm longer than the arms of abyssicola of the same
mantle length (49 mm), and they are more attenuate. This trend holds
in all specimens available. When material with tentacles in tact becomes
available, differences in the clubs may be found. Although it is difficult
to demonstrate quantitatively with the limited number of specimens on
hand, the mantle of berryi appears to be slightly more plump than that
of abyssicola.
LITERATURE CITED
Cuun, C. 1910. Die Cephalopoden. I. Teil Oegopsida. Wiss. Ergebn.
Deutschen Tiefsee-Exped. “Valdivia, 18(1): 1-410. Atlas of
61 plates.
Roper, C. F. E. 1968. Systematics and Zoogeography of the world-
wide bathypelagic squid Bathyteuthis (Cephalopoda: Oegop-
sida). Bull. U.S. Nat. Mus. In press.
:
(Gf, OO67D
Vol. 81, pp. 173-178 = < __ April 30, 1968
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
A REDESCRIPTION OF HATSCHEKIA CONIFERA,
YAMAGUTI 1939, (COPEPODA, CALIGOIDA),
INCLUDING THE FIRST DESCRIPTION
OF THE MALE
By Rocer F. CREssEy
Smithsonian Institution, Washington, D.C.
In 1939 Yamaguti described a new species of Hatschekia
based on 8 females collected from Stromateoides argenteus
(= Pampus argenteus) in Japan. Until now this has been the
only report of Hatschekia conifera and the male has remained
unknown. During Cruise 14 of the R. V. Anton Bruun off
Chile I collected sufficient material of this species to elaborate
the description of the female and describe the male for the
first time.
I wish to acknowledge the National Science Foundation—
US Program in Biology for supporting the field portion of this
work.
Hatschekia conifera Yamaguti, 1939
Material studied: 93 females and 12 males collected from the gill
filaments of a single specimen of the fish Cubiceps caerulus collected
during Cruise 14 of the R. V. Anton Bruun at 33°02’S 74°37'W off Chile.
The host was tentatively identified by Dr. Richard Haedrich. The
material has been deposited in the United States National Museum.
Description: Female: body form as in Figure la. Total length 2.78
mm (2.59-2.88 mm); greatest width .90 mm (.88—.91 mm) (based on
an average of 5 specimens).
Cephalon about twice as wide as long, comprising about one-eighth
total length, and separated from rest of body. First and second thoracic
segments fused and weakly separated from posterior portion of body.
Remainder of thoracic segments fused with genital segment comprising
three-fourths of total length. Dorsal body surface without surface
ornamentation. Posterior corners of genital segment produced (see Fig.
lb). Abdomen (Fig. 1b) small, one-segmented. Caudal rami bearing
5 setae, 4 terminal and 1 on median outer edge; rami held at a wide
21—Proc. Biot. Soc. WasuH., Vou. 81, 1968 (173)
174 Proceedings of the Biological Society of Washington
Fic. 1. Hatschekia conifera, female: a, dorsal view; b, abdomen and
caudal rami; c, first antenna; d, second antenna; e, mandible, first maxilla
and second maxilla; f, maxilliped.
ee ee ee ee a ee
Male of Hatschekia conifera 175
Fic. 2. Hatschekia conifera, female: a, leg 1; b, leg 2, male; c, ventral
view; d, caudal ramus; e, distal portion of second antenna.
——
—
Fic. 3. Hatschekia conifera, male: a, posterior corner of genital
segment; b, leg 1; c, leg 2; d, leg 3.
Male of Hatschekia conifera 177
angle to body. First antenna (Fig. 1c) 3-segmented; each of last 2
segments incompletely subdivided into 2 segments, all segments bearing
naked spines as in the figure. Second antenna (Fig. ld) in form of
a stout claw; basal 3 segments mostly covered with short spinules, no
setae on inner margins of segments. Mandible, first maxilla, and second
maxilla (Fig. le) small and weakly developed. Maxilliped (Fig. 1f)
4-segmented, each of last 3 segments with seta on inner margin, claw
bifid.
Legs 1 and 2 biramose. Leg 3 reduced to knob with 2 setae. Leg 4
absent. Leg 1 (Fig. 2a) with each ramus weakly divided into 2 seg-
ments; basipod with stout spine near base of endopod, exopod with 1
spine on outer distal corner of basal segment, terminal segment with
6 spines; endopod with 6 spines on terminal segment only. Leg 2 (Fig.
2b) with each ramus 2 segmented; exopod with strong spine on outer
distal corner of basal segment and 5 spines on terminal segment, endopod
with an inner spine on basal segment and 5 spines on terminal segment.
All spines naked. Both legs with an interpodal plate.
Eggs strings of the usual caligoid type; eggs uniseriate containing
30-45 eggs. Length of strings variable but usually at least as long as
body.
Male: body form as in Fig. 2c. Total length 1.12 mm; greatest width
.26 mm (based on a single specimen). Cephalon as long as wide,
comprising about one-fourth total length. Thoracic segments 1 and 2
separated. Genital segment fused with remaining thoracic segments.
Ventral surface of genital segment covered with delicate scales. Abdomen
l-segmented and as in female. Caudal rami (Fig. 2d) longer than wide
(70 u X 30); each ramus with 6 setae, one outer lateral, 2 subterminal,
and 3 terminal (terminal-most setae stout, with short plumosities ).
Oral area generally as in female. First antenna armed as in female
but each segment relatively longer than corresponding one in female.
Second antenna (Fig. 2e) with a prominent process near midpoint of
inner margin of second segment; each of last two segments with a single
seta, tip in form of a claw. Maxillae as in female except that posteriorly
directed setae are longer. Maxilliped as in female.
Legs 1 and 2 biramose, each ramus 2-segmented. Leg 1 (Fig. 3b)
with striated scale-like processes on basipod and rami; exopod first
segment with seta on outer distal comer, second segment with 6 plumose
setae; endopod first segment unarmed, second segment with 6 plumose
setae. Leg 2 (Fig. 3c) with scales as in leg 1; exopod first segment
with stout seta on outer distal corner, second segment with 5 plumose
setae; endopod first segment with plumose seta on inner distal corner,
second segment with 5 plumose setae. Leg 3 (Fig. 3d) reduced to a
lateral process on genital segment bearing 3 setae, longest plumose.
Single setae on posterior corner of genital segment (Fig. 3a) probably
represents leg 4.
Remarks: The diagnosis of the genus Hatschekia as given by Yama-
guti, 1963, p. 135 describes both sexes of this genus as having the head
178 Proceedings of the Biological Society of Washington
and first thoracic segment separated with the remainder of the thoracic
segments fused with the genital segment. Also Yamaguti states that the
mouth parts and swimming legs of the male are as in the female. In light
of this present study certain changes in the generic diagnosis of both sexes
are in order. The first 2 thoracic segments of both sexes are fused,
separated from the head anteriorly and the remaining thoracic segments
(fused with genital segment) posteriorly. The male differs from the
female by the presence of setae on the second antenna of the male, the
longer setae on the maxillae of the male, and the long plumose setae
on legs 1 and 2 of the male.
The genus Hatschekia has had nearly 75 species assigned to it. Most
of these are known by the female only. As far as I have been able to
determine the only other species from which males are known are H.
iridescens Wilson 1913 and H. prionoti described erroneously by Pearse
in 1947 as a female.
The collection used in this study from a stromateoid fish together with
Yamaguti’s original material from a stromateoid suggests a possible
affinity of this parasite for that host group.
LITERATURE CITED
Pearse, A. S. 1947. Parasitic copepods from Beaufort, North Carolina.
J. Elisha Mitchel Sc. Soc. 63(1): 1-16.
Wison, C. B. 1913. Crustacean parasites of West Indian fishes and
land crabs with description of new genera and species. Proc.
U.S. Nat. Mus. 44: 189-277.
Yamacutl, S. 1939. Parasitic copepods from fishes of Japan. Pt. 5.
Caligoida, III. Vol. Jub. Pro. Prof. S. Yokhida, 2: 443-487,
33 pls.
1963. Parasitic Copepoda and Branchiura of Fishes. John
Wiley and Sons, N. Y., 1104 pp.
oo
eS. a
:
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Vol. 81, pp. 179-190 30 August 1968
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
LECANURIUS KOSSMANNIANUS, A. NEW CYCLOPOID
COPEPOD PARASITIC IN HOLOTHURIANS
IN MADAGASCAR |
By ArtTHurR G. HuMEs
Department of Biology, Boston University, Bossa We
The lichomolgid copepod Lecanurius intestinalis Kossmann,
1877, was found by Semper at Bohol in the Philippine Archi-
pelago in the intestine of the holothurian Actinopyga lecanora
(Jaeger). The name of this host was given by Kossmann as
Miilleria lecanura Jager. His generic name Lecanurius was
evidently based upon an incorrect spelling of the specific
name. Another species of Lecanurius, found in Actinopyga
lecanora and A. miliaris (Quoy and Gaimard ) in northwestern
Madagascar, is described below.
The specimens were obtained by slitting the ventral body
wall of freshly collected holothurians while holding each over
a pail of weakly alcoholized sea water and thus saving the
fluid escaping from the body cavity. The entire holothurian
was then rinsed in this water. The contents of the pail were
subsequently strained through a fine net (after first removing
coarse fragments of viscera) and the copepods picked from
the sediment. Unfortunately, since the intestines of the holo-
thurians often broke, the exact location of the copepods,
whether in the body cavity or in the intestine, is not known.
All collections were made by the author, that in 1960 during
an expedition sponsored by the Academy of Natural Sciences
of Philadelphia and those in 1967 during field work aided by
a grant (GB-5838) from the National Science Foundation.
This grant has also supported the study of the specimens.
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. The abbreviations used are:
22—Proc. Biou. Soc. WAsH., Vou. 81, 1968 (179)
180 Proceedings of the Biological Society of Washington
FicurEs 1-7. Lecanurius kossmannianus new species, female. 1, body,
dorsal (A); 2, urosome, ventral (A); 3, area of attachment of egg sac,
dorsal (B); 4, caudal ramus, dorsal (C); 5, right egg sac, incomplete,
ventral (A); 6, left egg sac, incomplete, ventral (A); 7, rostral area,
ventral (C).
New Cyclopoid Copepod 181
A, = first antenna, Az = second antenna, MXPD = maxilliped,
and P,-P, = leg 1-leg 4. The measurements of the length of
the body have been made from specimens in lactic acid and
do not include the setae on the caudal rami.
I thank Dr. Elisabeth Deichmann of the Museum of Com-
parative Zoology, Harvard University,.for the identifications
of the holothurians.
FAMILY LICHOMOLGIDAE KOSSMANN, 1877
Genus LECANURIUS KossMANN, 1877
Leeanurius kossmannianus new species
(Figs. 1-32)
Type material: 4 22 and3 ¢ @ from 2 Actinopyga lecanora (Jaeger )
in 1 m under coral, east of Ambariotelo, a small island almost between
Nosy Komba and Nosy Bé, Madagascar. Collected July 20, 1967. Holo-
type @, allotype, and 5 paratypes (3 992, 2 66) deposited in the
United States National Museum.
Other specimens: From Actinopyga lecanora: 1 @ and 1 @ from 2
hosts, in 1 m, Pte. Ambarionaomby, Nosy Komba, near Nosy Bé, June 8,
1967; 2 $3 from 3 hosts, in 1 m, same locality, July 6, 1967; 2 9 9
from 1 host, in 3 m, Andjiabe, on the southern shore of Nosy Komba,
July 8, 1967; 1 @ and 1 ¢ from 3 hosts, in 1 m, Nosy N’Tangam, near
Nosy Bé, July 21, 1967; and 1 @ from 3 hosts, in 1 m, Ankify, on the
mainland of Madagascar opposite Nosy Komba, July 22, 1967. From
Actinopyga miliaris (Quoy & Gaimard): 1 @ from 25 hosts, in 1 m,
among the “sea grass” Cymodocea, Ambatoloaka, Nosy Bé, May 28,
1967; 1 @ and 2 $2 from 70 hosts, same locality, June 1, 1967; 1 9
and 2 ¢ 4 from 60 hosts, same locality, June 15, 1967; and 1 ¢ from
50 hosts, in 1 m, Antsamantsara, north of Madirokely, Nosy Bé, Oct. 31,
1960.
Female: Body (Fig. 1) elongated and flattened dorsoventrally, with
a broad and somewhat triangular cephalosome. Length 2.64 mm (2.40—
2.77 mm) and greatest width (at the level of the segment of leg 1)
0.97 mm (0.88-1.01 mm), based on 8 specimens. Margin of the head
indented on both sides. Segment of leg 1 very weakly separated from
the head, the separation indicated only by an indistinct and incomplete
transverse dorsal line. Tergal areas of segments of legs 1-4 becoming
progressively narrower posteriorly, their epimera rounded as indicated
in the figure. Ratio of length to width of the prosome 1.30: 1.
Segment of leg 5 (Fig. 2) 165 » x 352 uw. Between this segment and
the genital segment no ventral intersegmental sclerite. Genital segment
(Figs. 1 and 2) elongated, in dorsal view indented laterally near the
junction of its anterior two-thirds, expanded posteriorly to form two
lateral lobes, and then abruptly constricted in its posterior third (where
there is ventrally a transverse line). Greatest length of the genital
Ficures 8-15. Lecanurius kossmannianus new species, female. 8, first
antenna, ventral (D); 9, second antenna, anterior (B); 10, labrum,
ventral (E); 11, mandible, posterior (E); 12, paragnath, ventral (F);
13, first maxilla, posterior (E); 14, second maxilla, posterior (E); 15,
maxilliped, antero-inner (E).
New Cyclopoid Copepod 183
segment 540 yu, greatest width 396 yu, width in its anterior third 300 uz,
and width at the level of the lateral indentations 280 u. Areas of attach-
ment of the egg sacs situated dorsally and posteriorly on the lateral lobes,
each area (Fig. 3) bearing two naked setae 18 and 21 uw in length. Three
postgenital segments 220 u xX 198 uw, 198 uw X 176 uw, and 188 uw x 180 u
from anterior to posterior. First and second of these segments with a
transverse line ventrally. Anal segment with a row of minute spinules
along its posteroventral margin on each side, and bearing a pair of
relatively large dorsolateral setules.
Caudal ramus (Fig. 4) elongated, about 4 times longer than wide,
tapering slightly distally, its greatest length 265 yu, its greatest width
near its base 83 w, and its width near middle 65 yw. Mid-dorsal seta
(corresponding to the outer lateral seta in other lichomolgids) 63 u long,
distal dorsal setae 58 u, outermost terminal seta 68 u, innermost terminal
seta 64 uw, and the two long median terminal setae 211 » (outer) and
330 u (inner). All setae naked. A minute lateral spinule 6 u long on
the outer proximal area of the ramus, and the distal outer corner of the
ramus with a dorsal group of small spiniform processes and a marginal
row of minute spinules.
Dorsal surface of prosome and urosome with a few small hairs as in
Fig. 1. Urosome longer than the prosome, the ratio being 1 : 1.20.
Egg sacs in the single ovigerous female collected very long and slender,
but incomplete. Eggs in the right egg sac (Fig. 5) in a linear series, the
distalmost egg 78 u in diameter. This incomplete egg sac 1.5 mm long.
Eggs in the left sac (Fig. 6) partly linear and partly in two rows.
Rostral area (Fig. 7) not well defined posteroventrally.
First antenna (Fig. 8) 7-segmented, 470 u in length, with a sclerite
on the ventral side of the third segment suggesting an intercalary seg-
ment. A slight flexure in the first antenna at the level of the third
segment. Lengths of the segments (measured along their posterior non-
setiferous margins): 42 4 (99u along the anterior edge), 138 u, 36 u, 78
h, 52 pw, 38 w, and 29 uw respectively. Formula for the armature: 4, 13, 6,
3, 4 + I aesthete, 2 + 1 aesthete, and 7 + 1 aesthete, as in many
other lichomolgids. All setae naked.
Second antenna (Fig. 9) 4-segmented, 237 u long, including the claw.
Armature: 1, 1, 3 + claw, and 7. Second segment 109 wu along its outer
edge. Claw 109 u along its axis. Last segment with five of the seven
setae arising from a small distal lobe.
Labrum (Fig. 10) with two posteroventral lobes.
Mandible (Fig. 11) having at the base of the pectinate blade a blunt
process on the convex side and a pointed process on the concave side.
Paragnath (Fig. 12) a small lobe with a few small spinules, located
close to the inner side of the base of the first maxilla. First maxilla (Fig.
13) with three terminal setae and a subterminal hyaline seta. Second
maxilla (Fig. 14) 2-segmented. First segment unarmed. Second segment
having on its outer (ventral) margin a small proximal hyaline seta, on
its posterior surface a naked seta jointed basally, on its inner (dorsal)
184 Proceedings of the Biological Society of Washington
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BoA RO Rt AY
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17
Ficures 16-19. Lecanurius kossmannianus new species, female. 16,
area between maxillipeds and first pair of legs, ventral (D); 17, mid-
sternal areas of legs 1-4, with intercoxal plates pushed somewhat poste-
riorly, ventral (C); 18, leg 1, anterior (D); 19, leg 2, anterior (D).
margin a seta with hyaline serrations along its distal edge, and terminat-
ing in a short lash bearing two long teeth arising from a hyaline expan-
sion followed by a small hyaline tooth. Maxilliped (Fig. 15) 3-seg-
mented. First segment unarmed. Second segment rather tumid in outline
New Cyclopoid Copepod 185
and bearing two naked setae. Third segment with two naked setae and
terminating in a spiniform process with a slightly recurved tip.
Area between the maxillipeds and the first pair of legs (Fig. 16) not
protuberant; a sclerotized line between the bases of the maxillipeds.
Ventral sclerite of the segments of legs 1 and 2 projected posteriorly in
a median pointed process, which is surrounded by a small semilunar
sclerite (Fig. 17). Ventral sclerite of leg 3 without this pointed process
and only a trace of the semilunar sclerite present. Ventral sclerite of
leg 4 not evident and the semilunar sclerite absent.
Legs 1-4 (Figs. 18, 19, 20, and 22) with 3-segmented rami. Armature
as follows (Roman numerals indicating spines, Arabic numerals setae) :
P; protopod 0-1 1-0 exp I-00 [1 § IILI,4
end @O-1 O-l I,5
P, protopod O-l1 1-0 exp I-00 I-11 _ III,I,5
end OQ-l 0-2 LJIL,3
P; protopod 0-1 1-0 exp J-0 [1 £JILI5
end 0-1 0-2 ~ J[,III,2
P, protopod O-1 1-0 exp J0 J[-1 £JILI5
end) (0=1)) 0=1. 1,1,1,1,1
Inner seta on the coxa of legs 1-3 long and plumose but in leg 4 short
(24 «) and naked. Inner margin of the basis in leg 4 naked, but in
legs 1-3 bearing a row of hairs. Two proximal outer spines on the
exopods of legs 1-3 with prominent spinules; remaining spines fringed
with fine spinules. Distalmost spine of the exopod of leg 3 (Fig. 21)
and leg 4 with the inner margin bearing short spinules and with a short
distal fringe. Endopod of leg 4 nearly equal in length to the exopod.
First segment 68 u X 44 wu, with a slightly haired inner distal seta 88 u
long. Second segment 68 » X 39 u, with a shorter inner distal very
minutely barbed seta 39 u. Third segment 101 » x 42 u, bearing five
elements from outer to inner: a fringed spine 26 yu, a terminal fringed
spine 56 uw, a terminal barbed seta 112 uw, a subterminal barbed seta 91 uy,
and a minutely barbed seta 13 uw. Inner margin of the second segment
naked, but this margin of the third segment with a few hairs.
Leg 5 (Fig. 23) with a relatively small subrectangular free segment
78 « X 42 uw in greatest dimensions (about 1.86 times longer than wide),
bearing two terminal setae, the outer 55 uw and naked, the inner 106 yu
and finely barbed. Segment without ornamentation except for a terminal
row of minute spinules. Seta on the body near the free segment 44 yu
and naked, with a row of small spinules between it and the free segment.
Leg 6 probably represented by the two setae near the attachment of
each egg sac (see Fig. 3).
Color in life in transmitted light opaque and slightly brownish, eye
red, intestine brown, eggs light tan.
Male: Body (Fig. 24) resembling in general form that of the female.
Length 1.97 mm (1.92-2.08 mm) and greatest width 0.79 mm (0.75-
186 Proceedings of the Biological Society of Washington
{QE
>
or OIE a
20
SN
e Ss
Ficures 20-23. Lecanurius kossmannianus new species, female. 20,
leg 3, anterior (D); 21, distalmost spine on exopod of leg 3, anterior
(G); 22, leg 4, anterior (D); 23, leg 5, dorsal (B).
FicurE 24. Lecanurius kossmannianus new species, male. 24, body,
dorsal (A).
New Cyclopoid Copepod 187
0.81 mm), based on 7 specimens. Ratio of length to width of the pro-
some 1.13: 1.
Segment of leg 5 (Fig. 25) 86 u xX 208 uw. No ventral intersegmental
sclerite. Genital segment (Fig. 25) only a little longer than wide, 341 u
>< 335 u. (In males where the genital segment does not contain formed
spermatophores the segment is somewhat longer, as in Fig. 24, where
the dimensions are 363 » X 308 uw.) Four postgenital segments 133 » x
156 w, 151 w X 153 uw, 127 w x 140 w, and 135 w x 151 yw from anterior
to posterior.
Caudal ramus (Fig. 25) similar in form to that of the female but
relatively a little shorter, 198 » x 58 uw, or 3.4 times longer than wide.
Dorsal surface of the body ornamented with hairs as in the female.
Urosome longer than the prosome, the ratio being 1 : 1.52.
Rostral area, first antenna, second antenna, labrum, mandible, parag-
nath, first maxilla, and second maxilla like those of the female. Maxil-
liped (Figs. 26 and 27) 4-segmented, assuming that the proximal part
of the claw represents a fourth segment. First segment unarmed. Second
segment with two short naked setae and two groups of two or three
short stout spines. Third segment small and unarmed. Claw 242 wu along
its axis (including the terminal lamella), with a spiniform prominence
(keeled on its posterior surface) on its concave margin, and bearing
two very unequal naked setae proximally.
Exopods of legs 1-3 and both rami of leg 4 segmented and armed as
in the female. Endopods of legs 1-3 with the same arrangement of
spines and setae as in the female, but showing sexual dimorphism. (Spine
and setal formula of the male is thus similar to that of the female,
taking into account the fusion of the second and third segments of the
endopods in legs 1 and 2.) Endopod of leg 1 (Fig. 29) 2-segmented,
with the last two segments fused, but the original articulation still in-
dicated by an outer marginal spiniform process and a transverse row of
spinules. Endopod of leg 2 (Fig. 30) 2-segmented, with the original
articulation between the last two segments indicated as in leg 1. Endopod
of leg 3 with the four spines on the last segment somewhat different in
length from those in the female; these spines from proximal to distal in
the male 10 uw, 11 uw, 17 w, and 23 u, in the female 27 u, 33 u, 44 uw, and
55
Leg 5 (Fig. 31) resembling that of the female, but the free segment
smaller, 42 u X 21 uw (about twice as long as wide), its two terminal
setae 39 uw (outer) and 83 uw (inner).
Leg 6 (Fig. 32) a posteroventral flap on the genital segment bearing
two slender naked setae 36 uw and 44 wu in length.
Spermatophore not observed.
Color in life similar to that of the female.
(This species is named for Dr. Robby Kossmann, who first described
the genus Lecanurius. )
Comparison with Lecanurius intestinalis Kossmann: There are several
features whereby L. kossmannianus appears to be distinct from L. intes-
188 Proceedings of the Biological Society of Washington
EES
, male.
urosome, dorsal (H); 26, maxilliped, antero-inner (B); 27, maxilliped,
postero-outer (B); 28, claw of maxilliped, anterior (EK); 29, endopod of
leg 1, anterior (B); 30, endopod of leg 2, anterior (B); 31, leg 5, dorsal
(E); 32, leg 6, ventral (D).
25,
Ficures 25-32. Lecanurius kossmannianus new species
New Cyclopoid Copepod 189
tinalis. However, since Kossmann’s specimen was evidently a male (judg.
ing from his figure of the maxilliped), only males can be compared. In
L. kossmannianus the length of the body of the male is greater (1.97
mm) than in L. intestinalis (1.46 mm), the cephalosome is laterally
indented (but approximately semicircular in L. intestinalis), the genital
segment is a little longer than wide or its length is nearly equal to its
width (but twice as wide as long in Kossmann’s figure of L. intestinalis ),
the claw of the maxilliped has one pointed prominence (but two blunt
protuberances in L. intestinalis), and the free segment of leg 5 is 42 uw
long (but 130 » in L. intestinalis).
The armature of legs 1 and 4 of L. kossmannianus differs in certain
respects from that of L. intestinalis. Kossmann mentioned two setae on
the inner side of the second segment of the endopod of leg 1, but in
the Madagascan species there is only one such seta. (Perhaps this is a
lapsus by Kossmann, since the presence of two setae on the inner side
of this segment would be indeed remarkable in a lichomolgid copepod. )
In the exopod of leg 4 of Kossmann’s species the outer spines on the
first two segments are absent; on the endopod the inner element on the
first segment is absent, that on the second segment is a spine rather than
a seta, and the elements on the last segment are 3 spines and 2 setae
(in his text) or 3 spines and 3 setae (in his figure).
The males of both L. intestinalis and L. kossmannianus show striking
similarities in their general body form, and in the nature of the second
antenna, mandible, and second maxilla.
Relationship of Lecanurius to Scambicornus: The genus Lecanurius
shows certain characters which are similar to those of Scambicornus
Heegaard, 1944 (= Preherrmannella Sewell, 1949), a genus of which
many species live externally on holothurians. Stock (1964) has already
suggested that the two genera are related, noting the similarity in the
second antennae (the third segment with a large claw) and in the
armature (5 elements) of the third segment of the endopod of leg 4. In
addition, both genera have a bladelike mandible, without a long lash,
and both have 2-segmented endopods in legs 1 and 2 of the male.
There are, however, important differences between Lecanurius and
Scambicornus. In Lecanurius the body is elongated, with a much
broadened cephalosome, while in Scambicornus the form of the body is
more typically lichomolgid. In Lecanurius the egg sacs (as far as known)
are very long and the eggs are in a more or less linear series, rather than
sacciform with massed eggs as in Scambicornus. In Lecanurius the
mandible bears processes at both sides of the base of the blade, rather
than a single process (on the concave side) or none as in Scambicornus.
In Lecanurius the lash of the second maxilla is short and bears only a
few large teeth, whereas in Scambicornus it is longer and has a row of
several teeth. In Lecanurius the five elements on the last segment of
the endopod of leg 4 consist of both spines and setae, but in Scam-
bicornus these elements are spines. Lecanurius lives inside the holo-
thurians, while Scambicornus lives on the outside of these hosts.
190 Proceedings of the Biological Society of Washington
In view of these differences it seems best to retain Kossmann’s genus
Lecanurius for L. intestinalis and L. kossmannianus. If in the future
species intermediate between the two genera (in respect to the several
differences listed above) are discovered, Lecanurius might be considered
as a senior synonym of Scambicornus. This position is untenable now,
however, in the present state of knowledge of these genera.
Hosts of Lecanurius: Both L. intestinalis and L. kossmannianus live
in members of the genus Actinopyga. In fact, both of these copepods are
known from the same species of host, A. lecanora, though in widely
separated regions,—L. intestinalis in the Philippines, and L. kossman-
nianus in Madagascar. The new species occurs also in Actinopyga miliaris
in Madagascar, although apparently less commonly than in A. lecanora.
Sixteen specimens were recovered from 14 A. lecanora, but only 8 from
205 A. miliaris.
A single copepodid of what is possibly a Lecanurius was found in
another holothurian, Synapta maculata (Chamisso and Eysenhardt), at
Ambariobe, near Nosy Bé, October 6, 1963 (during field work of the
U.S. Program in Biology, a part of the International Indian Ocean
Expedition). The immaturity of this specimen made it impossible to
determine the genus with certainty, but some of its features suggest
Kossmann’s Lecanurius.
Associated with both hosts of L. kossmannianus there is a species of
Scambicornus, S. campanulipes (Humes and Cressey, 1961), which lives
on the exterior of A. lecanora and A. miliaris in Madagascar (Humes,
1967).
LITERATURE CITED
HeEEGAARD, P. 1944. A new copepod (Scambicornus hamatus) para-
sitic on a Japanese holothurian. Vidensk. Medd. fra Dansk
naturh. Foren., 107: 359-366.
Humes, A. G. 1967. A new species of Scambicornus (Copepoda, Cy-
clopoida, Lichomolgidae) associated with a holothurian in
Madagascar, with notes on several previously described spe-
cies. Beaufortia, 14 (173): 135-155.
Humes, A. G., AND R. F. Cressey. 1961. Copépodes cyclopoides du
genre Preherrmannella parasites d’holothuries et d’un oursin
a Madagascar. Mém. Inst. Sci. Madagascar, 1959, sér. F,
3: 25-65.
KossMANN, R. 1877. Entomostraca (1. Theil: Lichomolgidae). In:
Zool. Ergeb. Reise Kiistengeb. Rothen Meeres, erste Halfte,
IV, pp. 1-24. Leipzig.
SEWELL, R. B. S. 1949. The littoral and semi-parasitic Cyclopoida, the
Monstrilloida and Notodelphyoida. John Murray Exped.
1933-34, Sci. Repts., 9 (2): 17-199.
Stock, J. H. 1964. On Scambicornus Heegaard, 1944, a senior syn-
onym of Preherrmannella Seymour Sewell, 1949 (Copepoda,
Cyclopoida). Beaufortia, 10 (123): 183-192.
O0TE
Vol. 81, pp. 191-196 30 August 1968
Mr
PROCEEDINGS j AUG 9
OF THE
BIOLOGICAL SOCIETY OF i Deca
ee ie
— ae
MICROGOBIUS CROCATUS, A NEW GOBIID FISH
FROM PACIFIC PANAMA?
By Ray S. Brrpsonc
Institute of Marine Sciences, University of Miami,
Miami Florida
The genus Microgobius is distributed from Baja California
to Ecuador in the Pacific and from Chesapeake Bay to Natal,
Brazil in the Atlantic. No species is common to both oceans.
A new species, herein described, is the eighth known Pacific
representative of this genus. There are six species in the
Atlantic.
Many species of Microgobius are sexually dimorphic in fin
and body pigmentation, larger mouth and elongation of the
dorsal spine filaments in the male, and greater development
of the fleshy nuchal crest in the female. As M. crocatus is
known from a single male, the nature of these characters in
the females may vary from the following account.
The single specimen of M. crocatus has been compared with
material of both sexes throughout the known range of each of
the nominal Pacific species of Microgobius. Those Pacific
species which I consider to be valid are listed in the compara-
tive materials section. A review of the entire genus is nearing
completion.
The manuscript benefited from the suggestions of C. R.
Robins, Institute of Marine Sciences, University of Miami;
F. H. Berry, Bureau of Commercial Fisheries, Tropical Atlantic
Biological Laboratory; and E. A. Lachner, U.S. National Mu-
seum. Field work in Panama was supported by The National
Science Foundation (NSF-GB-4389, C. R. Robins, principal
investigator) and by The National Geographic Society.
1 Contribution No. 938 from the Institute of Marine Sciences, University of Miami.
23—Proc. Biot. Soc. WaAsH., Vou. 81, 1968 (191)
192 Proceedings of the Biological Society of Washington
Comparative Material: Microgobius tabogensis Meek and Hildebrand:
USNM 81844 (holotype), PANAMA, Taboga Island, S. E. Meek and S. F.
Hildebrand, 1 (36.2); USNM uncat., San Francisco Beach near Panama
City, S. F. Hildebrand, 13 February 1937, 2 (16.0-22.3); USNM uncat.,
C. Z., Venado Beach, S. F. Hildebrand, 26 February 1937, 3 (30.3-
31.5); USNM uncat., C. Z., Miraflores Locks (lower chamber), S. F.
Hildebrand, 26-29 March 1937, 26 (17.0-33.1); UCLA W54-45, costa
r1cA, Golfo de Nicoya, Isla Caballo, Erdman Cove, C. Peterson and party,
22 November 1952, 58 (15-35); UCLA W54—254, mexico, Baja, Bahia
Magdalena, Turtle Inlet, M. A. Newman and party, 81 (29.3-45.0).
Microgobius miraflorensis Gilbert and Starks: FMNH 8489, panaMa,
Rio Culebra, S. E. Meek and S. F. Hildebrand, 19 March 1911, 2 (29.7-
33.2); USNM 81845, Rio Culebra, S. E. Meek and S. F. Hildebrand,
19 March 1911, 2 (32.8-33.8); UMMZ 173398, Mexico, Guerrero,
Laguna Coyuca, about 10 mi. NW of Acapulco, R. R. and M. Miller, 21
March 1957, 9 (24.5-31.0); UCLA W52-44, Sonora, Gulf of California,
Estero, about 2 mi. W of Yavarros, A. O. Flechsig and party, 9 February
1952, 17 (20.9-36.3).
Microgobius curtus Ginsburg: USNM 107292 (holotype), ECUADOR,
Salada, Guayaquil, W. L. Schmitt, 1-2 October 1926, 1 (29.9); USNM-
88788 (paratypes), same data as USNM 107292, 5 (18.7-33.9); UMML
23812, PANAMA, Pacific Ocean, about 1.3 mi. SSE of Miraflores Locks
on west side of Panama Canal near town of Cocoli, R. Birdsong, T.
Fraser, T. Murray, 14 May 1967, 1 (36.1); UMML 23811, C. Z., Mira-
flores Locks at base of spillway, R. Birdsong, T. Fraser, 11 May 1967,
1 (39.3); UMML 23813, C. Z., bay SSE of Miraflores Locks on W side
of canal E of Cocoli, R. Birdsong and D. Dean, 17 May 1967, 1 (39.4).
Microgobius brevispinis Ginsburg: USNM 81842 (holotype), PANAMA,
Balboa, in tide pools, S. E. Meek and S. F. Hildebrand, 7 February 1912,
1 (42); USNM 81841 (paratypes), Panama City, in tide pools, S. E.
Meek and S. F. Hildebrand, 19 February 1912, 4 (42.2-48.7); USNM
81843, Panama City, in tide pools, S$. E. Meek and S. F. Hildebrand,
21 March 1912, 2 (45.3-50.1); UMML 23810, C. Z., Kobbe Army Base,
Venado Beach, R. Birdsong and party, 12 May 1967, 4 (14.8—47.6);
SIO 62-106, MExico, Baja, Magdalena Bay, Isla Margarita, 44 (34.6—
63.0).
Microgobius cyclolepis Gilbert: USNM 44370 (holotype), Gulf of
California, 30°37'30”N, 113°07’W, 7 fms., ALBATROss sta. 3020, 24 March
1889, 1 (44.8); SIO 64-875, Mexico, Baja, Bahia Almejas, field no.
B6411-12, TR 122, 109 (24.9-43.5).
Microgobius erectus Ginsburg: UCLA W50-43, Mexico, Sonora, Gulf
of California, 80 mi. S Guaymas, near Boca del Rio Mayo, A. O. Flechsig,
27-29 January 1950, 2 (39.8-47.4); UMML 23809, panama, Bay of
Panama, 8°38.6’N, 78°51.9’W, 17 fms., R/v PILLSBURY sta. 535, 6 May
1967, 23 (28.4-37.3); UMML 23808, Bay of Panama, 7°50.7'N, 80°
09.8’W, 9-10 fms., R/v PILLSBURY sta. 492, 2 May 1967, 11 (28.5-44.6).
Microgobius emblematicus (Jordan and Gilbert): FMNH 8488,
TN nn H ice OT aes
New Gobiid Fish 193
PANAMA, Balboa, S. E. Meek and S. F. Hildebrand, 5-6 May 1911, 1
(39.7); FMNH 8487, Balboa, S. E. Meek and S. F. Hildebrand, 31
January 1912, 1 (30.0); UMML 23806, C. Z., Kobbe Army Base, Venado
Beach, R. Birdsong and party, 12 May 1967, 59 (24.1-48.7); UCLA
W51-36, Mexico, Sinaloa, Gulf of California, Astillero at Mazatlan, B.
W. Walker and party, 29 January 1951, 2 (29.7-33.3).
Microgobius crocatus new species
(Fig. 1)
Diagnosis: A species of moderate size (30.6 mm SL) having no dark
spot on body below origin of spinous dorsal; lips, pelvic distal margin
and anal margin yellow; interorbital distance broad (7.8 mm in head);
eye large (3.1 mm in head); scales cycloid except for a small patch of
weakly ctenoid scales under P:; lateral scale rows about 49.
Description: D. VII-I,17; A. 1,17; P: 23-23; Pz 1,5, the inner rays
completely joined; P: with well-developed frenum; caudal fin with 17
segmented rays, 15 branched rays; gill-rakers of first arch 4+ 14;
vertebrae 11 + 16. Head, nape and area below anterior spinous dorsal
naked. Tongue deeply notched. Anterior teeth in two rows in both
jaws, one row posteriorly; outer tooth row enlarged, caninoid; no teeth
on vomer or palatines. Mouth large (upper jaw 1.6 in head), inclined
about 45° from horizontal axis. Branchiostegals 5 (1 on shaft of cerat-
ohyal, 3 on enlarged portion of ceratohyal, 1 on epihyal). Third to fifth
dorsal spines with elongate filaments, longest reaching end of soft dorsal.
Cephalic lateral-line canals (Fig. 2): supraoccipital canals separate
except at juncture with single, enlarged medial pore; each supraoccipital
canal terminating in a single nasal pore; infraorbital canal obsolete,
containing a single pore at juncture of supraorbital canal with lateral
canal; lateral canal with 2 pores; preopercular canal short, with 2 pores.
Anterior nostril tubular. Fleshy nuchal crest absent. Pectoral fin 3.6 mm
in SL; pelvic fin 4.3 mm in SL.
Color Description: Body dusky, greenish yellow dorsally, becoming
white ventrally; belly bluish. Two small yellow spots on body under
dorsal edge of appressed pectoral fin. Yellow spot on mid-pectoral base,
another on mid-line of nape. Two iridescent blue, longitudinal stripes
on cheek; two iridescent blue spots on anterior edge of opercle, and two
orange-yellow spots on posterior edge; lips dusky yellow; chin dusky.
Spinous dorsal fin dusky with yellowish cast to basal portion; clear
slash originating at base of each spine and running at slight diagonal to
each spine; distal portion of fin dark with orange-red cast to filaments.
Soft dorsal dusky, with narrow, dark distal margin and orange cast.
Pelvic fins dusky with distal yellow margin. Anal fin dusky with yellow
distal margin and a submarginal dark stripe. Caudal fin with yellow
ventral stripe and several orange spots dorsally; median caudal rays
dusky orange.
194 Proceedings of the Biological Society of Washington
-yueurg ‘olporg ‘opeut “TS wet g'OE “LEGZOS INNSN 4 M[04
‘snqoIOLI SMGOBOLI
Al
mo) iG |
New Gobiid Fish 195
Fic. 2. Cephalic lateral-line canals of Microgobius crocatus. A.—
lateral view: supraoccipital canal (soc); lateral canal (lat); infraorbital
canal (inf); preopercular canal (pop). B.—dorsal view: medial supra-
occipital pore (a); nasal pore (b); infraorbital pore (c); lateral pores
(d, e).
196 Proceedings of the Biological Society of Washington
TaBLeE 1.—Comparison of Microgobius crocatus
with Microgobius tabogensis.+
Species
Character Microgobius crocatus Microgobius tabogensis
Spot below D1 origin? absent present
Body dorsum? no dark dashes 5-6 dark dashes on body
along dorsal fin bases
Spinous dorsal fin? 7 clear slashes basal row of dark spots
Soft dorsal fin? narrow dark border, no dark border,
no basal stripes 2-3 basal stripes
Anal fin? dark submarginal stripe no dark submarginal stripe
Pelvic fins? dusky with pale margin completely dusky
Interorbital width 7.8 in head 13.4 in head (10.8-17.3 )
Ctenoid scale patch under Pi — small. moderately large
Second dorsal fin i M7 I, 16 (rarely 17)
Anal fin I ey I, 16 (rarely 17)
1 Males only.
2 Pigmentation in preserved specimens.
Etymology: The name crocatus (Latin) alludes to the yellow markings
on the fins, body and lips.
Holotype: USNM 202587 (30.6 mm SL, male); PANAMA, Pacific
Ocean; approximately 1.3 mi. SSE of Miraflores Locks on west side of
Panama Canal near town of Cocoli; muddy tidal slough; depth of capture
4 ft.; 14 May 1967; original field no. RSB-PAN-11; collected by R.
Birdsong, T. Fraser, T. Murray.
Habitat: A tidal slough that receives some freshwater through a drain- .
age ditch emptying the incompleted auxiliary lock of the Miraflores
system near Cocoli. The bottom consists of soft mud and detritus to a
depth of several feet. The shores are thickly overgrown with mangrove.
There is a moderate surge during peak tidal flow. One specimen of
Microgobius curtus was among the other species collected with the
holotype.
Relationships: Microgobius crocatus appears to be most closely related
to M. tabogensis with which it is sympatric. They are similar in all
meristic features and differ primarily in interorbital width and pigmenta-
tion pattern (Table 1). Features of color and pattern are diagnostic
and highly stable in the genus Microgobius. This appears to be true of
all brightly colored species of gobiid fishes which display a large degree
of sexual dimorphism in pigmentation.
.0O676
Nol. 81, pp-197+208 30 August 1968
Vite i 10LO
AY §- x &y te Oa!
SO BIBLOGIEAL SOCIETY OF WASHINGTON
AN ECOLOGICAL STUDY ON VALLEY-FOREST
SPIDERS FROM NORTHERN KENTUCKY
By Brantey A. BRANSON AND Donacp L. BatcH!
Department of Biological Sciences, Eastern Kentucky
University, Richmond, Kentucky
As far as can be discerned no one has compared the spider
faunas of relict vs disturbed forest regions in the Cumberland
Mountains. A number of studies have been carried out on the
ecology of soil arachnids in forests (Gasdorf and Goodnight,
1963; Elliot, 1930; and others) and several taxonomic lists are
available for certain forested regions, such as that of Jones
(1940), but such lists are not available for any Kentucky
locality.
The Study Area: The Cumberland Mountains of Kentucky offer widely
variable habitats, certain parts of them representing some of the oldest
undisturbed habitat in North America. Parts of the Red River Drainage
in the northern part of the state are of this nature. Particularly instruc-
tive are some tributary canyon systems of the middle fork of the stream,
lying in Wolfe and Powell counties. In this region, the authors have
commenced a long-term ecological analysis, special emphasis being
accorded the Arachnida, Chilopoda, Diplopoda and Mollusca.
In the canyon systems alluded to, the valley floors are mostly underlaid
by undifferentiated Mississippian rocks, whereas the valleys themselves
are incisions through the much interdigitated Lee Formation, directly
underlaid by the Breathitt Formation; the latter is exposed on the floors.
The Breathitt Formation consists primarily of gray, micareous siltstones,
gray, subgraywacke sandstone, dark and light claystones containing some
ironstone concretion, some limestone (sparse), chert and coal. The
rugged, narrow valleys with steep cliffs are supported by this formation.
The overlying Lee Formation forms even more massive cliffs, from 200
to 300 feet high (vertical), and consist mostly of resistant sandstone and
siltstone, relatively little clay, ironstone and limestone being present.
The canyon systems here considered head approximately 4 mile
1 Supported by Eastern Kentucky University faculty grant 5-3-372-6.
24—-Proc. Biot. Soc. WasH., Vou. 81, 1968 (197)
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northwest of Pine Ridge, Wolfe County, Kentucky (Map 1). These are
associated with Tight Hollow Creek, Mill Creek and the Middle Fork
of Red River, the latter lying in Powell County. The Tight Hollow
system drains 0.558 square mile, and it is three-pronged, roughly Y-
shaped. The south arm, which also receives a short, deep bifurcation,
heads at 1,040 feet mean sea level (MSL), and extends 2.4 miles to its
confluence with the northern arm. The cliffs at the head are of sand-
stone, averaging slightly more than 200 feet in height and are vertical,
concave risers. Below this point, the cliffs are of variable height, but
always steep, the enclosed valley narrow and rocky. The northern arm
heads at 1,240 feet MSL and makes contact with the southern arm at
1,030 feet two miles below the origin. It is similar in topography to the
southern arm. Following this, the lower arm (Fig. 1) extends 2,225 feet
to Mill Creek Canyon, at 900 feet MSL. From this point, the canyon
meanders between steep walls for 12,000 feet to open into the much
wider canyon of the Red River, receiving the small canyons of Doe
Branch, Black John Creek, and Doublecave Branch, all of these being
very similar to Tight Hollow.
Generally speaking, these canyon systems lie in mixed mesophytic
forest land. The dominant trees are Betula lenta Linnaeus, Tsuga cana-
densis Carr, Liriodendron tulipifera Linnaeus, Acer rubrum Linnaeus,
Cornus florida Linnaeus and Magnolia fraseri Walt. On the south ridge
Ilex opaca Ait is abundant from station 6 upgrade. From stations 1
through 5, Rhododendron maximum Linnaeus is exceedingly dense, up
to 2,117 main stalks per acre being counted in some places. Below
station 5, this species is progressively replaced by mountain laurel (Fig.
2). A similar relationship was observed between the hemlock and tulip
tree (Fig. 3), and also hemlock versus pines (White, Virginia). This
is a reflection of past logging. Stations 1 through 5 lie in a relatively
undisturbed, relictual area, whereas the stations below 5 have been
variously disturbed by farming, deforestation and some rather insignif-
icant silver mining.
Of the ground vegetation various species of Polypodiaceae, Sedium,
Lygodium, Lycopodium and numerous mosses are the most abundant
in the upper stations, with some dense stands of Equisetum arvense
Linnaeus along the creek and thick growths of Mitchella repens Linnaeus
on the drier slopes. Downstream, several grasses have invaded clearings.
In the undisturbed areas there is a thick litter of leaves and needles, in
addition to logs in various stages of decomposition.
Collecting Stations: Collections were secured from fourteen localities,
as indicated in Figure 1. The limits of each station are demarked by
arrows. A swath-transect, set at right angles to the long axis of the
valley, was followed from crest to crest. Temperature readings, air at
standard level, soil at six inches, are in centigrade.
Station 1. 26 February 1966. The valley is narrow with vertical cliffs
and several old talus slides; numerous fallen logs; vegetation dense, moss,
ferns, Liverworts; heavily shaded by Tsuga, Magnolia, and a large pop-
200 Proceedings of the Biological Society of Washington
2500
2400
2200
2000
1800
MOUNTAIN
1600 RHODODENDRON
1400
NUMBER PER ACRE
Ce
°
°
600
400
200
TRANSECT
Ficure 2. Importance values of Rhododendron versus Mountain Laurel.
ulation of Rhododendron and Ilex; considerable ground seepage. Tem-
perature: soil, 3.0; air, 2.0.
Station 2. 5 March 1966. Valley walls steep, V-shaped in section;
many flat sandstones; considerable topsoil (washed in from above);
vegetation as in station 1. Temperature: soil, 5.0; air, 5.5.
Station 3. 12 March 1966. Valley walls less declivitous; U-shaped;
much organic debris; vegetation as in station 1. Temperature: soil, 6.0;
air, 15.0.
Valley-forest Spiders 201
Station 4. 19 March 1966. Slope of valley walls continues to amelio-
rate; a modest floodplain (100 feet in width) produced, composed of
gravel, mud, large and small sand-rocks, and deep organic litter. Tem-
perature: soil, 7.5; air, 10.0.
Station 5. 26 March 1966. Nearly identical to station 4. Temperature:
soil, 6.0; air, 10.0.
Station 6. 2 April 1966. Valley much wider, slope of walls more
gentle, supporting numerous flowing springs; Tsuga partially replaced
by white pine, and Rhododenron by Kalmia. Temperature: soil, 7.0;
air, 10.7.
Station 7. 9 April 1966. Similar to station 6, but valley reverts to a
narrow V-configuration, with steep walls; considerable ironwood and
maple; practically no Rhododendron; large boulders, rubble, dead logs.
Temperature: soil, 6.0; air, 7.8.
Station 8. 23 April 1966. Mill Creek Valley; wide, the slope of walls
only moderate; floodplain two to three hundred yards wide, gently
inclined toward the creek and covered with grasses, sedges, sumac,
various weedy species, Lobelia, willow and dogwood; some boulders,
small stones, rubble and organic siltation. Temperature: soil, 16.0; air,
19.7.
Station 9. 14 May 1966. Similar to station 8, but walls even less
steep; floodplain with more grass and much sand; a few abandoned
silver mines. Temperature: not taken.
Station 10. 22 May 1966. Similar to station 9, but valley wider.
Temperature: soil, 11.5; air, 21.5.
Station 11. 25 June 1966. Here the valley has been impounded by
a small dam. Around the margins of the lake are many dead trees,
standing and fallen. Otherwise conditions are quite similar to those at
station 9, though the valley is somewhat deeper and narrower, since the
average slope is increased. Just below this point, Mill Creek empties into
the Middle Fork of Red River. Temperature: soil, not taken; air, 34.0.
Station 12. 11 June 1966. Valley of Middle Fork of Red River. Valley
much wider, floodplain nearly a mile wide in places, under agriculture
in places (corn and tobacco), and numerous old-field seres evident;
willow, sycamore, tulip trees, hawthorn, few pines and young Tsuga.
Temperature: soil, 17.0; air, 23.7.
Station 13. 2 July 1966. Valley wide, floodplain about 80 feet wide
on either side of river; weeds, dead leaves, flat sandstones; vegetation
as in station 12. Temperature: soil, 22.5; air, 32.0.
Station 14. 9 July 1966. Similar to station 13, but stream bank heavily
overgrown by kudzu. Temperature: soil, not taken; air, 30.2.
Acknowledgments: We are indebted to Dr. H. W. Levi and his stu-
dent, Mrs. J. E. Coyle (Museum of Comparative Zoology, Harvard)
for identifying most of the specimens, and for suggestions. In the field,
we were accompanied by Messrs. P. W. Hake, C. J. Moore ( University
of Delaware), R. T. Schaaf (University of Illinois), J. S. Stacy (Ken-
tucky Park Service), and John Egan.
202 Proceedings of the Biological Society of Washington
200
180
160
140}
120
100
80
60
NUMBER PER ACRE
40
TRANSECT
FicurE 3. Importance values of hemlock versus tulip tree.
Methods: At each collecting station shown in Figure 1, a 12-foot wide
swath, located at the center of each area, was hand collected from one
rim of the valley to the other. Most of our efforts were directed toward
securing specimens from the ground stratum, although vegetation was
searched up to a height of six feet.
No attempt has been made to analyze in detail seasonal effects,
although the authors are quite aware of its profundity as regards popula-
tion fluctuations (Gasdorf and Goodnight, 1963). However, Figure 4
demonstrates an obvious increase in both total numbers and the percent-
age-composition of mature versus immature forms. It is perhaps interest-
ing to note the relatively large figures shown on the graph for March 13.
This probably reflects an unusual warming tendency during the week
immediately preceding, in which the air temperature climbed to approxi-
Valley-forest Spiders 203
9 e Oo
MAY JUNE JULY
APRIL
MARCH
FEB
SHOR SRC OMum oS Yo ne OC ae)
NUMBER SPECIMENS
Ficure 4. Spiders collected at study areas in Wolfe and Powell coun-
ties, Kentucky as a reflection of advancing season. Solid circles, total
numbers collected; open circles, immature forms as percentages of total.
mately 15° C. Following this, temperatures fell back to 10° C and
remained there until the middle of April.
Consequently, in the following discussion two primary objectives are
met. (1) The fauna of the undisturbed area (Stations 1 through 5) is
compared with that of the progressively more disturbed (Stations 6
through 14). A third objective should probably be included, i.e., report-
ing the species included; very little work has been attempted on the
Kentucky Arachnida. (2) The general ecology of the species collected
is discussed.
Results: Seventy-seven species of spiders were collected during a six-
month period, from February through July, from the study area delin-
eated above. These species were distributed through 61 genera and 19
families. Of the total number collected, 70 percent were females, 17
204 Proceedings of the Biological Society of Washington
TasBLe 1. Comparison of spiders taken by random swath collecting
in a northern Kentucky valley system, 1966. Numbers refer
to collecting stations.
Species
Antrodiaetus
Ariadna bicolor
Achaearnea tepidariorum
Latrodectus mactans
Theridion rupicola
Theridion albidum
Frontinella communis
Pityohyphantes phrygianus
Helophora insignis
Lepthphantes sp.
Linyphia marginata
Linyphia maculata
Microneta varia
Ceraticelus sp.
Erigone
Micrathena gracilis
Micrathena mitrata
Meta menardi
Mangora placida
Mangora maculata
Neoscona sp.
Epeira raji
Aranea sp.
Theridiosoma sp.
Theridiosoma gemmosum
Leucauge venusta
Tetragnatha versicolor
Tetragnatha elongata
Tetragnatha sp.
Cybaeus
Cicurina sp.
Cicurina robusta
Cicurina brevis
Agelenopsis pennsylvanicus
Wadotes sp.
Wadotes carolinus
Wadotes calearatus
Coras sp.
Coras lamellosus
Coras montanus
Coras tangynus
Calymmaria sp.
Undisturbed
2,5
1,3
Disturbed
9
12, 14
7
12
13
i}, Mal
9, 10, 12
14
1565 Uk,
9, 10, 11
6
7,10, 14
6
11, 12, 13, 14
7, 8, 10
6
6
9
vi
Shared
|
Species
Tegenaira sp.
Hahnia. cinerea
Dolomedes sp.
Dolomedes scriptus
Dolomedes tenebrosus
Dolomedes urinator
Dolomedes vittatus
Pirata sp.
Pirata montanus
Pirata maculatus
Pardosa sp.
Trocosa sp.
Schizocosa sp.
Schizocosa crassipes
Lycosa helluo
Zelotes duplex
Drassyllus virginianus
Anahita sp.
Anahita punctulatus
Micaria sp.
Trachelas sp.
Clubiona sp.
Clubionoides texana
Castianeira longipalpus
Agroeca minuta
Phrurotimpus alarius
Anyphaena peéctorosa
Oxyptila sp.
Xysticus elegans
Xysticus ferox
Philodromus rufus
Habrocestum pulex
Habrocestum acerbum
Thiodina sp.
Pellenes
Habronattus sp.
Paraphidippus sp.
Neon nelli
Maevia vittata
Valley-forest Spiders 205
TABLE 1 (Continued )
Undisturbed Disturbed Shared
4 6 X
13
13, 14
3
4 6 X
12
12
3 12
9,10
13
4,5 GOP 2013 xX
fe)
9
NO, ey, 1 TY 14
8, 10, 11
10
LOM13
oh 4b Gh Tal TB Lay x
10
4
1
5
9
11
13
9, 11,14
13
11
IS
ea
2
5
12,14
10
10
9
3,4
8
Ge, 1
percent were males, and 13 percent were immature forms. Since most
of our efforts were confined to species occupying the ground stratum
those data for web-building or foliage hunters, such as Frontinella com-
munis and Micrathena, are not significant.
206 Proceedings of the Biological Society of Washington
As noted by Whitcomb, Exline, and Hite (1963), and as illustrated by
Table 1, lycosids have a decided preference for more or less open areas.
The arrangement of our collecting stations, beginning in the narrow,
steeply declivitous portion of the valley system and extending down-
grade into the wider, less declivitous portion, is strongly reflected by
spider distribution, as illustrated in the Table. This is probably because
of an increase in habitat diversity in the lower stretches, and because
considerably more moisture is available around the year. The narrow
parts of the valleys become very dry, except precisely on the floor, during
the summer months.
This same picture is obvious in the whole spider fauna (Table 1),
both from the standpoint of numbers of species present and numbers of
individuals. When the effect of advancing season is considered, it does
not appear that the relict, undisturbed area has retained a significantly
different spider fauna than the areas in the disturbed.
EcoLocicAL ANNOTATIONS
Antrodiaetus species. Lives in burrows, mostly beneath fallen logs and
deep litter. Males observed on surface during late winter.
Ariadna bicolor (Hentz). Primarily found beneath the loosened bark of
dead trees, fallen or standing.
Achaearanea tepidariorum (Koch). This species is seldom found far
from human habitations. Stations 12 and 14 are both in close prox-
imity to various buildings. Our specimens were secured beneath large
rocks.
Latrodectus mactans (Fabricius). The single specimen was found be-
neath a rock near an abandoned farm site.
Theridion rupicola Emerton. Beneath a large piece of bark on the
ground.
Theridion albidum Banks. In a clump of partridge pea, near valley floor.
Frontinella communis (Hentz). In web on a small white pine.
Lepthyphantes sp. In leaf litter.
Linyphia marginata Koch. On base of white pine.
Linyphia maculata Emerton. On face of upright rock wall.
Pityohyphantes phrygianus (Koch). On ground, open floor of valley;
not under trees.
Helophora insignis (Blackwell). On tall grass at the margin of Mill
Creek.
Microneta varia (Blackwell). Under dead leaves on ground; deep shade.
Ceraticelus sp. On ground in leaf litter.
Erigone sp. On ground in leaf litter.
Micrathena gracilis (Walckenaer ).
Micrathena mitrata (Hentz).
Neoscona sp.
Aranea sp.
The last four forms were all secured from orb webs on low vegetation.
Valley-forest Spiders 207
Eveira raji (Scopoli). This species builds a large orb web, but hides in
a rolled leaf about five feet above the ground.
Mangora placida (Hentz). In a small orb web located approximately
eight inches above the ground.
Mangora maculata (Keyserling). In a small orb web at about shoulder
height above the ground.
Meta menardi (Latreille). Under a deeply placed monolith of large size.
Theridiosoma sp.
Theridiosoma gemmosum (Koch).
The last two forms were removed from some liverworts on the face of
a cliff, near the valley floor.
Leucauge venusta (Walckenaer ).
Tetragnatha versicolor Walckenaer.
Tetragnatha elongata Walckenaer.
The last three forms were all found on limbs and twigs extending out
above the creeks. When approached, they flatten themselves against
twigs, their legs held parallel to the long axis of the body.
Agelenidae. All the funnelweb spiders in this area were secured from
the ground level, either running or under stones, logs, and debris.
Dolomedes. The four species of this genus are more or less restricted to
stream and pond margins. Several specimens were removed from the
water. Since first order streams in this region periodically cease run-
ning in the summer, the greater abundance of these forms in the lower
parts of the canyon system is probably correlated with the constant
' water supply found there.
Lycosidae. With the exception of Pirata montanus Emerton, which pre-
fers shaded ravines, all the wolf spiders here discussed are species of
the lower altitudes, at the edge of the forest, and in cleared areas.
Zelotes duplex Chamberlin. Under a stone in tall grass on the floodplain.
Drassyllus virginianus Chamberlin. On leaf litter.
Anahita punctulatus. Under rocks, large pieces of bark on the ground,
and dead logs.
Micaria sp.
Clubiona sp.
Clubionoides texana.
Castiameira longipalpus (Hentz).
Agroeca minuta Banks.
Phrurotimpus alarius (Hentz).
The last six forms were all secured from leaf litter, chiefly on the slopes.
Trachelas sp. Beneath a clump of grass near the creek.
Anyphaena pectorosa Koch. In Scirpus, about a foot above the ground,
near the creek.
Oxyptila sp. In leaf litter.
Xysticus ferox (Hentz). Under rocks and on ragweeds.
Xysticus elegans Keyserling. Wandering on the ground.
208 Proceedings of the Biological Society of Washington
Philodromus rufus Walckenaer. On a fallen log.
Habrocestum pulex (Hentz).
Habrocestum acerbum.
Lhe last two species were removed from bare rocks.
Thiodina sp.
Pellenes sp.
The last two forms were found in leaf litter. -
Habronattus sp. In tall grass near the creek.
Neon nelli Peckham. On fallen pine needles.
Maevia vittata (Hentz). Under logs and fallen bark.
LITERATURE CITED
Extiot, F. R. 1930. An Ecological Study of the Spiders of the Beech-
Maple Forest. Ohio J. Sci., 30: 1-22.
Fircu, H.S. 1963. Spiders of the University of Kansas Natural History
Reservation and Rockefeller Experimental Tract. Misc. pub.
U. Kansas, 33: 1-202.
GasporF, E. G., AND C. J. Goopnicut. 1963. Studies on the Ecology
of Soil Arachnids. Ecology, 44: 261-268.
Jones, S. E. 1940. An annotated list of the spiders of an east-central
Illinois forest (Wm. Trelease Woods University of Illinois).
Trans. Illinois State Acad. Sci., 33: 216-220.
Wuitcoms, W. H., H. Exuine, anp M. Hite. 1963. Comparison of
spider populations of ground stratum in Arkansas pasture and
adjacent cultivated field. Proc. Arkansas Acad. Sci., 17: 34—
39.
74.6673
Vol. 81, pp. 209-222 30 August 1968
Ne yee OF SARDINE (SARDINELLA,
; AE) FROM THE MARQUESAS ISLANDS!
By Freperick H. Berry AND PETER J. P. WHITEHEAD
Bureau of Commercial Fisheries, Tropical Atlantic Biological
Laboratory, and British Museum (Natural History)
A sardine from the Marquesas Islands in recent years has
been introduced to the Hawaiian Island area as a bait fish
and has successfully reproduced there. A study has been
completed on many aspects of its biology (Nakamura and
Wilson, MS). In several publications this species has been
referred to or discussed under the name Harengula vittata
(Valenciennes) and other binomials (see synonymy ).
Of the three closely related genera Sardinella, Herklotsich-
thys, and Harengula, the Marquesan sardine is best included
in Sardinella, as defined by Chan, 1965a, Berry, 1964, and
Whitehead, 1963 and 1964.
Recent studies of the Indo-Pacific species of Sardinella have
better defined many of the valid species and have clarified
the status of many nominal species (Dutt, 1959 and 1961;
Chan, 1965a and 1965b; Whitehead, 1965 and 1966; White-
head, Boeseman, and Wheeler, 1966; Whitehead, 1967). We
find that the Marquesan sardine lacks a valid scientific name,
and we describe it here as a new species.
Sardinella marquesensis new species
Figs. 1-6
Harengula sundaica (non Bleeker), Kendall and Goldsborough, 1911:243
(specimens from Nukahiva, MCZ 29507 and USNM 50794).
Sardinella jussieu (non Lacépéde ), Fowler, 1928:30 (in part; based on
Nukahiva specimens, USNM 65861; excluding other references and
distributions ).
1 Contribution No. 65 Tropical Atlantic Biological Laboratory, Bureau of Com-
mercial Fisheries, Miami, Florida 33149.
25—Proc. Bion. Soc. WasH., Vou. 81, 1968 (209)
‘OFGIOS WNSA ‘Spuryst sesonbieyy ‘vaArpT NY~NN “ovyorey, “TS wut ,OT “edAzofoy ‘srsuasanbipue Djjauypung "TAMAS
ton
ing
| Society of Wash
gica
210 Proceedings of the Biolo
A New Sardine 211
FicurE 2. Outline of second (posterior) supramaxilla of S. marque-
sensis. The anterior (shaft) end is to the left.
Sardinella melanura (non Cuvier), Fowler, 1928:30 (in part; based on
Nukahiva specimens, MCZ 29507; excluding other specimens and ref-
erences ).
Harengula vittata (non Valenciennes ), Fowler, 1941:596 (in part; based
on Nukahiva specimens, USNM 65861; excluding other references ).
Wilson and Rinkel, 1957:110 (the Marquesan sardine, caught as a
bait species). Wilson, Nakamura, and Yoshida, 1958:71 (Marquesan
bait species). Murphy, 1960:185 (introduction into Hawaiian waters ).
Yoshida, 1960:37 (Marquesan bait species). Hida and Morris, 1963:
431 (Marquesan sardine in Hawaii).
Diagnosis: A Sardinella with overlapping scales on median predorsal
ridge (not a single median row); 8 pelvic finrays; body scales with 3 to
6 anterior interrupted transverse grooves, and with exposed portion not
perforated, greatly produced, or fimbriated; 58 to 85 gillrakers on lower
limb of first arch (at 80 to 107 mm SL); no dark tips on caudal fin
lobes. Supplemental diagnostic characters: Head length 25.5 to 28.3%
SL (3.5 to 3.9 times into SL); body depth 22.9 to 28.5% SL (3.5 to 4.4
times into SL); 16 or 17 prepelvic and 12 or 13 postpelvic ventral scutes;
ca. 30 to 60 epibranchial and ca. 23 to 35 ceratobranchial gillrakers on
first arch (at 80 to 107 mm SL).
Holotype: USNM 201946, 107 mm SL, caught at Taiohae Bay, Nuku
Hiva, Marquesas Islands, 08°55’ S, 140°06’ W, 14 September 1956, on
Charles H. Gilbert cruise 30, station 45, by bait seine, preserved by
Donald W. Strasburg.
Description of the Holotype: Dorsal rays 17. Anal rays 21. Pectoral
rays 14 on each side. Pelvic rays 8 on each side. Lateral scale rows ca.
42. Transverse scale rows ca. 12. Ventral scutes: prepelvic 16, post-
pelvic 12. Frontoparietal striae ca. 7 on each side. Gillrakers on lateral
side of first arch, epibranchial 39, ceratobranchial 30, hypobranchial ca.
41. Branchiostegals 7 on each side. Pseudobranchial filaments 15. Gill
filaments on lateral side of first arch, upper limb ca. 25, lower limb ca.
45. [Measurements in millimeters, followed by percent of standard length
in parentheses:] Standard length 107. Fork length 115 (107.5). Total
212 Proceedings of the Biological Society of Washington
FicurE 3. Outline of the striae on the frontoparietal and occiput
region of S. marquesensis.
length 130 (121.5). Head length 28.0 (26.2). Eye diameter 7.9 (7.4).
Pupil diameter 3.6 (3.4)— Snout length 8.0 (7.5). Upper jaw length
11.7 (10.9). Lower jaw length 13.7 (12.8). Opercle depth 14.5 (13.6).
Pseudobranch length 6.6 (6.2). Gillraker length, at angle on lower limb
4.9 (4.6). Longest dorsal ray length (5th) 15.3 (14.3). Dorsal-fin base
length 15.2 (14.2). Longest anal ray length (4th) 4.7 (4.4). Anal-fin
base length 16.6 (15.5). Pectoral fin length 16.9 (15.8). Pelvic fin
length 11.6 (10.8). Pelvic axillary scale length 8.5 (7.9). Snout to
dorsal-fin origin 47.5 (44.4). Snout to pelvic-fin origin 53.5 (50.0).
Snout to anal-fin origin 77 (72.0). Dorsal-fin origin to caudal base 62
(57.9). Body depth, at occiput 22.0 (20.6), at dorsal-fin origin 28.2
(26.4), at anal-fin origin 20.3 (19.0), least depth of peduncle 9.8 (9.2).
Body width at dorsal-fin origin 12.4 (11.6).
Paratypes: MARQUESAS ISLANDS. Nuku Hiva, Taiohae Bay, 08°55’ S,
140°06' W, Gilbert cruise 35, station 11, 13 October 1957, ANSP 109213
(1 specimen, 95 mm SL), BMNH 1967.11.4.1-2 (2, 95-103), BPBM
5592 (1, 98), CAS 24046 (1, 101), FMNH 74076 (1, 85.5), RMNH
25824 (1, 95), MCZ 45899 (1, 98), MNHN 1967-591 (1, 105), SU
66228 (1, 97), TABL 101827 (2, 93-101), UMMZ 186866 (1, 95),
USNM 201947 (2, 88-106). Hiva Oa, Atuona, 09°49’S, 139°03' W,
Gilbert cruise 32, station 26, 27 January 1957, ANSP 109212 (1, 62),
BMNH 1967.11.4.3 (4, 53-59.5-62-67 ), BPBM 5593 (2, 60.5-63), CAS
24045 (2, 62-63.5), FMNH 74075 (1, 62), RMNH 25825 (2, 58.5-
64.5), MCZ 45898 (1, 62), MNHN 1967-590 (2, 60-63.5), SU 66229
(1, 59.5), TABL 101828 (3, 48.5-55.5-67), UMMZ 186867 (1, 62),
USNM 201948 (3, 53.5-56.5-64). Nuku Hiva, Taiohae Bay, collected
with the holotype, USNM 179874 (19, 56.5-96.5). Nuku Hiva, Taiohae
A New Sardine 913
\ |
& WA QS
ss
\ YV W *
FicurE 4. Outline of posterior 5 anal-fin rays of S. marquesensis.
Bay, 08°55’ S, 140°06’ W, Gilbert cruise 43, station 90, 14 March 1959,
TABL 101829 (7, 77.5-91). Tahu Ata, Vai Tahu, 09°07’ S, 139°06’ W,
Gilbert cruise 50, station 33, 24 November 1960, TABL 101830, (39,
45.5-97.5). Nuku Hiva, Taiohae Bay, 08°55’ S, 140°06’ W, Gilbert cruise
50, station 32, 22 November 1960, TABL 101831 (26, 74-88). Nuku
Hiva, Taiohae, 26 September 1929, ANSP 52615 (1, 59). Nuka Hiva,
17 September 1899, USNM 65861 (25, 33.5-61.5), MCZ 29507 (7,
43-52 ).
Description of the Species: Dorsal-fin rays 16 (7 specimens) or 17
(22); the first 3 or usually 4 rays unbranched; the last ray divided to
its base. Anal-fin rays 18 (1), 19 (16), 20 (11), or 21 (1); the first 3
or rarely 2 rays unbranched; the last ray divided to its base. Pectoral-fin
rays 13 (4), 14 (20), or 15 (17) on the left side; of 41 specimens, 6
with one more ray on right side than on left, 3 with one more ray on left;
the first ray unbranched. Pelvic-fin rays 7 on both sides (39), 8 on left
side and 7 on right (1), and 7 left and 8 right (1); the first ray un-
branched. Caudal-fin rays 10 + 9 principal, 9 or 10 + 7 or 8 secondary.
Branchiostegal rays 6 on both sides (41), 6 left and 7 right (3). Scales
in lateral series ca. 35-42 (no specimens with all scales remaining).
Ventral scutes, 16 prepelvic + 12 postpelvic (1), 17 + 12 (22), 17 +
13 (15). Gill filaments on lateral side of first arch ca. 21 to 25 upper
and 40 to 45 lower at 88 to 107 mm SL. Pseudobranchial filaments ca.
14 to 19, tending to increase with increase in body length.
Body proportions in percent of standard length from 11 selected speci-
mens 67-107 mm SL, except as noted. Head length 25.5 to 28.3 (as in
diagnosis). Eye diameter 7.3 to 8.2. Snout length 7.2 to 8.1. Upper
jaw length 10.9 to 11.6. Lower jaw length 12.1 to 13.6. Pectoral-fin
length 15.8 to 17.6. Pelvic-fin length 10.7 to 12.2. Pelvic axillary scale
length 6.9 to 8.2. Dorsal-fin base length 11.5 to 15.1. Anal-fin base
length 14.3 to 16.5. Snout to dorsal-fin origin 43.4 to 47.3. Snout to
pelvic-fin origin 50.0 to 54.2. Snout to anal-fin origin 72.0 to 76.8.
Caudal base to dorsal-fin origin 55.7 to 59.0. Caudal base to pelvic-fin
origin 52.4 to 55.2. Caudal base to anal-fin origin 27.0 to 30.4. Depth
at occiput 20.4 to 23.1. Depth at dorsal-fin origin 24.8 to 27.1, tending
214 Proceedings of the Biological Society of Washington
to increase slightly with increase in body length. Depth at anal-fin origin
17.5 to 20.0. Least depth of peduncle 9.1 to 10.2. Body width at dorsal-
fin origin 10.1 to 11.6. Length of gillraker at angle on lower limb 4.0
tOnorlle
Body compressed, the belly strongly keeled. Snout usually slightly
shorter than eye diameter. Maxilla smooth along lower edge; the second
(posterior) supramaxilla somewhat rounded posteriorly, but with the
point of junction at the shaft of the ventral part of the expanded portion
anterior to that of the dorsal part (Fig. 2); no hypomaxilla. No teeth in
upper jaw or on palate or tongue; a few recurved canines near symphysis
of lower jaw. Upper jaw ending at about a vertical through the anterior
part of the pupil.
Pseudobranch with crescentic base and a slightly elevated basal ridge;
its length about equal to eye diameter. Gillrakers relatively slender,
tapering markedly at anterior ends of upper and lower limbs. Fronto-
parietal region (Fig. 3) as described under generic relationships. Bran-
chiostegal membrane on left side overlapping externally at isthmus with
that on right side.
Dorsal and anal fins with low scaled basal sheaths; dorsal-fin origin
nearer to tip of snout than to caudal base; anal-fin origin about midway -
between pelvic-fin origin and caudal base, or slightly nearer to pelvic;
last 2 rays of anal fin not markedly enlarged but more branched than
preceding rays (Fig. 4). Pectoral-fin tips failing to reach pelvic-fin
origin by a distance greater than snout length and less than post-orbital
head length; a recess present beneath dorsal part of pectoral fin. Pelvic-
fin origin beneath about anterior third or fourth of dorsal-fin base; pelvic
fin inserted about midway between pectoral-fin origin and anal-fin origin
or slightly closer to anal fin; pelvic axillary scale present, its length about
three-fourths that of pelvic fin. Scales covering bases of pectoral, pelvic,
and caudal fins, extending onto dorsal and anal lobes of caudal fin to
near fork of fin.
Body scales. Exposed portion only slightly eroded or not eroded poste-
riorly and not fimbriated, and lacking perforations (Fig. 5). Anterior
(imbedded) portion of scale with a continuous basal transverse groove
and with the more anterior transverse grooves all interrupted and increas-
ing in number both from anterior to posterior part of the body and with
increase in length of fish. Scales from the midlateral side of the body,
on a 103-mm specimen with 2 to 4 anterior interrupted transverse grooves
anterior to the dorsal-fin origin, 4 or 5 between the origins of the dorsal
and anal fins, and 5 or 6 on the peduncle. Scales taken from peduncle
with 2 anterior interrupted grooves on a 3l-mm specimen, 4 at 70 mm,
and 5 or 6 at 103 mm.
Predorsal scales. Scales between the occiput and the dorsal-fin origin
along the middorsal ridge are not arranged in a single median row typical
of the subgenus Amblygaster (Chan, 1965a:106; 1965b, Fig. 1). Neither
are they exactly as described as “a double row” for the subgenus Clu-
peonia (Whitehead et al., 1966:47; Chan, 1965a, 1965b). They are
A New Sardine DALES
FicurE 5. Scale from the midlateral part of the body above the anal-
fin base of S. marquesensis. The imbedded portion is at the top; the
dotted line marks the termination of the circuli.
basically irregular and not strictly median. In one 95-mm paratype, 19
scale-pockets (scales missing) lie in part over the median predorsal ridge;
8 of them are mainly on the left side of the fish, 6 are on the right, and
5 are essentially median; the 5 median scales are in consecutive series
starting about one-half way back in the area. Other type specimens have
similar but varying arrangements of predorsal scales.
Pigmentation. Dorsal third of body darkly pigmented, usually marked
in its ventral portion by a slightly wide and a narrow longitudinal stripe,
and grading into small, less dense melanophores on the midlateral part
of the body, with the ventral third to half of the body unpigmented. Top
of head and snout dark, with melanophores grouped along anterior part
of preorbital, premaxilla, and maxilla, and along mandible; melanophores
scattered below eye and on opercular bones; tip of lower jaw and frenum
dark. Dorsal and caudal fin with scattered melanophores; pectoral,
pelvic, and anal fins unpigmented. Peritoneum very dark. Scattered
melanophores on tongue, inside mandibles, on palate, on inside of gill
covers (especially grouped along pseudobranch base), and in humeral
area beneath gill cover. No marked spots on body; the two stripes men-
tioned above not very pronounced, and not obvious or formed on smaller
specimens; the caudal fin without dark tips on the lobes.
Gillrakers. The number of gillrakers on the lateral side of the first gill
arch increases with increase in body length. This increase is depicted
for a series of specimens between 31 and 107 mm SL in Fig. 6. The
gillrakers on the upper limb (over the epibranchial bone) are shown on
the top portion of the graph; the stippled area indicates our estimation,
from this relatively small sample, of the number of gillrakers that might
be expected at any standard length within this size range. The lower
216 Proceedings of the Biological Society of Washington
UPPER LIMB) ————>
s
T T T
——Se
=
—-9
———-~-f_@
—_____
—_—e
——
—2
~4
es
GILLRAKERS
ie!
T =
+ —
LOWER LIMB
8
if
—o—
=|
[a aes SS: a ae
70 }—
L A-Hypobranchial
80 —
| Ir
90K
L
60h @-Epibranchial | | |
l %-Ceratobranchial
OME
100 as | 1 | 1 | i ! fe ! 1 | 1 ! 1 | n
te) 30 40 50 80 90 100
60 70
STANDARD LENGTH (mm)
Ficure 6. Depiction of gillraker ontogeny in S. marquesensis, show-
ing the number of gillrakers located over each element of the first gill
arch (epibranchial above; ceratobranchial and hypobranchial cumula-
tively below) for 52 specimens of various standard lengths.
portion of the graph shows the number of gillrakers (increasing down-
ward on the graph) on the lower limb (i.e., over the ceratobranchial
and hypobranchial bones); the number of gillrakers over the hypo-
branchial has been added to that over the ceratobranchial for each speci-
men (this number is associated above with the epibranchial count for
each specimen); the number of gillrakers plotted for hypobranchial thus
represents the total number of gillrakers on the lower limb. The stippled
areas again represent the estimated range in number of rakers that might
be expected at any standard length. The increase in number of gillrakers
over the epibranchial and hypobranchial bones is expected, because the
small, graduated rakers over the anterior ends of each of these bones
indicates that these are the areas of continued formation of additional
gillrakers. The stippled area representing the complement of gillrakers
over the ceratobranchial bone also indicates an increase in gillraker
number with increase in standard length; but because of the relatively
similar lengths of gillrakers over the ceratobranchial, it is apparent that
A New Sardine D7
no new gillrakers can be formed there. We theorize that the increase
over the ceratobranchial is produced by the growth anteriorly of the
bone so that its anterior end comes to lie under gillrakers that previously
were over the posterior end of the hypobranchial. A similar condition
was observed in the thread herring genus Opisthonema (Berry and
Barrett, 1963).
On the basis of our estimates of gillraker complements, the following
number of gillrakers could be expected at the body sizes indicated:
Standard length 30 40 50 60 70 SO 90 100
Epibranchial 9-13 13-20 17-25 21-29 24-34 26-40 29-48 33-58
Ceratobranchial 11-15 13-18 16-21 18-25 20-28 22-31 24-35 27-38
Lower limb 21-26 27-35 34-45 40-53 44-61 49-70 54-81 59-89
Occurrence or absence of gillrakers on the five gill arches in S. marque-
sensis is typical of that in other species of Sardinella. An 88-mm SL
specimen of S. marquesensis had the following numbers of upper +
~
lower limb gillrakers on the lateral and medial sides of the 5 arches:
arch lateral medial
Ist 37 + 62 0+ 0
2nd 39 + 80 0+ 0
3rd 37 + 65 24 + 0
4th 26 + 48 56 + 30
5th 54 + 30
Vertebrae. Forty-two vertebrae (including the terminal ural) in 79
specimens. Two other specimens had 43 vertebrae, but both had abnor-
mal structures associated with the centra. Six other specimens had 44
vertebrae; 5 of these were abnormal, but the sixth specimen apparently
had normal structure. X-rays show usually 9, rarely 8, supraneurals
(predorsals ) anterior to the interneurals.
Distribution: S. marquesensis was apparently endemic in the Mar-
quesas, from where it was first recorded in 1899. In 1957 it was
introduced into Hawaiian waters. It is the only known species of
Clupeidae occurring in the Marquesas Islands, where it apparently
replaces the closely related S. melanura, widely distributed from the
Society Islands westward into the Indian Ocean.
Relationships with Other Species: Sardinella marquesensis is most
closely related to Sardinella melanura (Cuvier, 1829), a species with
prominent black tips on the lobes of the caudal fin. Chan (1963b:7)
described S. nigricaudata as a new species from New Hebrides, making
three Indo-Pacific species of the Sardinella-Herklotsichthys group with
black-tipped caudal-fin lobes; the other two, long acknowledged species
were S. melanura (Cuvier) and Herklotsichthys vittata (Valenciennes,
1847). Whitehead (1967) concluded that vittata of Valenciennes was
a junior synonym of melanura of Cuvier, questioned the validity of
nigricaudata of Chan, and suggested that there was only a single, wide-
ranging species with black caudal tips (i.e., melanura) from the Indian
Ocean and the Philippines to Tahiti. Currently we subscribe to White-
218 Proceedings of the Biological Society of Washington
head’s suggestion. We further add that all specimens we have seen of
S. melanura, from throughout its range, have no appreciable variation in
the black-tipped lobes.
S. marquesensis, with one exception, is similar in morphological charac-
ters to S. melanura. Compared to specimens of S. melanura (ANSP
86245) from Tahiti (about 750 miles from the Marquesas Islands), S.
marquesensis has slightly greater gillraker complements and tends to
have a less deep body, especially under the termination of the dorsal fin.
The black-tipped caudal-fin lobes that characterize S. melanura are
absent in S. marquesensis. The scarcely and uniformly pigmented lobes
of S. marquesensis (and the lack of black tips) are constant not only in
the type specimens we have examined, but also in all live specimens
observed by Bureau of Commercial Fisheries personnel in the Marquesas
and in progeny of that stock that has reproduced in Hawaiian waters
(E. L. Nakamura, personal communication). We conclude that this
striking difference in caudal pigmentation between the two species
represents a trenchant genetic difference, and requires specific distinction
of the two forms.
Generic Allocation: Most of the morphological characters proposed to
separate the genera Sardinella, Herklotsichthys, and Harengula are sub-
jective, and most of these characters are not precise (as they have been
defined) in certain species within these genera. This statement holds
particularly in Sardinella marquesensis, which is more intermediate in
characters between Sardinella and Herklotsichthys than other species
previously studied as critically. [These two genera lack the hypomaxillae,
present in the American Harengula; hence the latter need not be further
considered here.] The four characters used in recent publications to
distinguish between Sardinella and Herklotsichthys are discussed below
in relation to S. marquesensis.
1. Relative size of terminal two anal-fin rays: in Sardinella more
extensively branched and elongated than preceding rays; in Haren-
gula relatively similar in size and branching to preceding rays.
This character was used by Regan (1917) and was qualified by
Chan (1966a:107-109). In some species of Sardinella the excessive
branching, thickness, and length of the last two anal-fin rays are
extremely pronounced, whereas in Herklotsichthys the last several
rays are almost identical to those preceding, and the antepenulti-
mate ray may be longer than the penultimate ray (except that the
terminal ray is branched to its base). In other species of Sardinella
the last two rays are almost intermediate between these two
extremes. In S. marquesensis, the antepenultimate ray is the same
length as or slightly longer than the penultimate ray (but not as
long as the ultimate ray), but the two terminal rays are slightly
thicker and more branched than those immediately preceding (Fig.
4). In this character, S. marquesensis is almost intermediate be-
tween the two genera.
2. Transverse grooves on body scales: the basal transverse groove is
A New Sardine 219
continuous in both genera; in Herklotsichthys the other (anterior)
transverse grooves in the anterior field are reported to be continu-
ous from dorsal to ventral margin of the scale on all scales from
the midlateral side of the body; in Sardinella the other transverse
grooves are interrupted on body scales of all species in the area
anterior to a vertical of the dorsal-fin origin (in some species
transverse grooves are interrupted on all body scales, but in other
species the grooves may be partially or incompletely interrupted in
the area posterior to a vertical of the dorsal-fin origin, Chan, 1965a:
107-109); S. marquesensis has all grooves (other than the basal
transverse groove) interrupted on all midlateral body scales (Fig.
5). This character would imply, on the basis of previous accounts,
a relation only to Sardinella, and would exclude Herklotsichthys.
We know, however, that previous accounts are partly erroneous,
because certain midlateral body scales posterior to the cleithrum
and anterior to a vertical of the dorsal-fin origin in Herklotsichthys
(two species examined, as well as four species of Harengula) have
interrupted transverse grooves. This observation tends to discount
further the morphological and taxonomic separation of Herklot-
sichthys from Sardinella.
Shape of the second supramaxilla: in Sardinella the junction of
the shaft of the second supramaxilla with its expanded posterior
portion is aligned almost dorsally and ventrally and the dorsal and
ventral parts of the expanded portion are almost equal in area
(described variously as rectangular, almond-shaped, paddle-shaped,
nearly circular); in Herklotsichthys the point of junction of the ven-
tral part of the expanded portion is well anterior to that of the
dorsal part, and the ventral part is larger than the dorsal (described
as somewhat crescentic) (Chan, 1966a:108-109). This character is
also subjective to a degree, as the shape of the expanded portion of
some species of Sardinella departs from the “nearly circular” shape
toward that of Herklotsichthys. Here again, S. marquesensis tends
to be intermediate (Fig. 2); most of the expanded portion has a
nearly circular outline as in Sardinella, but the ventral part of the
expanded portion joins the shaft distinctly anterior to the junction
of the dorsal part—characteristic of Herklotsichthys.
Number of fronto-parietal striae: cited as 3-7 in Herklotsichthys;
7-14 in Sardinella (Whitehead, 1964:41). The individual striae
are easier to distinguish and to count in some species of the two
genera than in others. In S. marquesensis they are especially
confused; there is much branching and anastomosing, and more
irregularity in thickness in the striae (Fig. 3) than in most other
species examined; and they are even less well-defined in larger
than in smaller specimens. Our approximate counts of the striae
in S. marquesensis range from six to eight on one side, which is
intermediate between the ranges given for the two genera, but
the high count of eight favors Sardinella.
220 Proceedings of the Biological Society of Washington
Acknowledgments. Eugene L. Nakamura, U.S. Bureau of Commercial
Fisheries Biological Laboratory, Honolulu, first called this problem to
our attention. The following persons made specimens available for
study: Ernest A. Lachner, U.S. National Museum (USNM); James E.
Bohlke and James C. Tyler, Academy of Natural Sciences of Philadelphia
(ANSP); Giles W. Mead and Myvanwy M. Dick, Museum of Compara-
tive Zoology, Harvard University (MCZ); George S. Myers and Warren
C. Freihofer, Natural History Museum, Stanford University (SU). In
addition to these four museums, paratypes have also been deposited at
the British Museum of Natural History (BMNH); Bernice P. Bishop
Museum (BPBM); California Academy of Sciences (CAS); Field Mu-
seum of Natural History (FMNH_); Rijksmuseum van Natuurlijke Historie
(RMNH); Muséum National d’Histoire Naturelle (MNHN); Tropical
Atlantic Biological Laboratory (TABL); and University of Michigan
Museum of Zoology (UMMZ). Grady W. Reinert, Tropical Atlantic
Biological Laboratory, prepared the illustrations. The manuscript bene-
fited from reviews by Robert V. Miller and Daniel M. Cohen.
LITERATURE CITED
Berry, F. H. 1964. A hypomaxillary bone in Harengula (Pisces: Clu- -
peidae). Pac. Sci., 18 (4): 373-377.
Berry, F. H., anv I. BArvetrr. 1963. Gillraker analysis and speciation
in the thread herring genus Opisthonema. Inter-Amer. Trop.
Tuna Comm., 7 (2): 113-150.
Cuan, W. L. 1965a. <A systematic revision of the Indo-Pacific clupeid
fishes of the genus Sardinella (family Clupeidae). Jap. J.
Ichthyol., 12 (3/6): 104-118.
1965b. Ibid. Jap. J. Ichthyol., 13 (1/3): 1-39.
Dutt, S. 1959. Biometric studies on Sardinella spp. off Waltair coast.
I. Sardinella fimbriata Val. Proc. First All-India Congr.
Zool., 2: 286-298.
1961. Biometric studies on Sardinella spp. off Waltair
coast. 2. Sardinella gibbosa Blkr. Jour. Zool. Soc. India,
13 (1): 78-89.
Fowier, H. W. 1932. The fishes obtained by the Pinchot South Seas
Expedition of 1929, with descriptions of one new genus and
three new species. Proc. U.S. Nat. Mus., 80 (art. 6, no.
2906): 1-16.
1941. Contributions to the biology of the Philippine Archi-
pelago and adjacent regions. The fishes of the groups
Elasmobranchii, Holocephali, Isospondyli, and Ostarophysi
obtained by the United States Bureau of Fisheries steamer
“Albatross” in 1907 to 1910, chiefly in the Philippine Islands
and adjacent seas. Bull. U.S. Nat. Mus., no. 100, 13:i-x +
1-879.
1928. The fishes of Oceania. Mem. B. P. Bishop Mus.,
10:i-iii + 3-540.
A New Sardine I
Hipa, T. S., anp R. A. Morris. 1963. Preliminary report on the
Marquesan sardine, Harengula vittata, in Hawaii. Pac. Sci.,
17 (4); 431-437.
KENDALL, W. C., AND E. L. GotpsBporoucH. 1911. Reports on the
scientific results of the expedition to the tropical Pacific, in
charge of Alexander Agassiz, by the U.S. Fish Commission
steamer “Albatross,” from August, 1899, to March, 1900,
Commander Jefferson F. Moser, U.S. N., commanding. XIII.
The shore fishes. Mem. Mus. Comp. Zool., 26 (7): 241-243.
Murpnuy, G. I. 1960. Introduction of the Marquesan sardine, Haren-
gula vittata (Cuvier and Valenciennes), to Hawaiian waters.
Pac. Sci., 14 (2): 185-187.
Nakamura, E. L., AnD R. C. Witson. A contribution to the biology of
the Marquesan sardine (Sardinella marquesensis). Unpub-
lished manuscript.
WHITEHEAD, P. J. P. 1963. Herklotsichthys Whitley 1951 to replace
Harengula Valenciennes 1847 for Indo-Pacific species ( Pisces:
Clupeidae). Ann. Mag. nat. Hist., Ser. 13, 6: 273-284.
1964. A redescription of the holotype of Clupalosa bulan
Bleeker, and notes on the genera Herklotsichthys, Sardinella,
and Escualosa (Pisces: Clupeidae). Ann. Mag. nat. Hist.,
Sem digi; 30-41.
1965. A review of the elopoid and clupeoid fishes of the Red
Sea and adjacent regions. Bull. Br. Mus. nat. Hist., (Zool. ),
12 (7): 225-281.
1966. The elopoid and clupeoid fishes in Richardson’s
‘Ichthyology of the Seas of China and Japan.’ Bull. Br. Mus.
nat. Hist., (Zool.), 14 (2): 15-44.
1967. The clupeoid fishes described by Lacépéde, Cuvier,
and Valenciennes. Bull. Br. Mus. nat. Hist. (Zool.), Suppl.
2: 1-180.
WHITEHEAD, P. J. P., M. BoEsSEMAN, AND A. WHEELER. 1966. The
types of Bleeker’s Indo-Pacific elopoid and clupeoid species.
Zool. Verhandl. Leiden, no. 84, 1-152.
Witson, R. C., E. L. NAkAMuURA, AND H. O. YosHipa. 1958. Mar-
quesas area fishery and environmental data, October 1957—
June 1958. Fish & Wildl. Serv., Spec. Sci. Rep.—Fish., no.
283, 1-105.
Witson, R. C., anp M. O. RinxeL. 1957. Marquesas area oceanog-
raphic and fishery data, January—March 1957. Fish & Wildl.
Serv., Spec. Sci. Rep.—Fish., no. 238, 1-136.
Yosuipa, H. O. 1960. Marquesas area fishery and environmental data,
January—March 1959. Fish & Wildl. Sery., Spec. Sci. Rep.—
Fish., no. 348, 1-37.
999, Proceedings of the Biological Society of Washington
MEE OG] 3
Vol. 81, pp. 223-230 30 August 1968
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
TWO NEW SPECIES AND THREE NEW RECORDS
OF BENTHIC POLYCHAETES FROM PUGET
SOUND (WASHINGTON)! <//
By Kart BANSE AND FREDERIC H. NICHOLS
Department of Oceanography, Univ. of Washington,
Seattle, Washington
We describe here two new polychaete species, give three
new records for Puget Sound, Washington, and comment on
one species known already from this region. The animals
were found among the polychaetes of twenty-one 0.1 m? grab
samples taken primarily in spring 1963 (see Lie, in press).
Locations and full descriptions of the sampling stations, includ-
ing full species lists, can be found in that paper.
FAMILY PHYLLODOCIDAE
Eulalia (Pterocirrus) macroceros (Grube)
Figure la—c
E. (Sige) macroceros.—Berkeley and Berkeley, 1948, p. 48.
E. macroceros.—Banse, 1959, p. 423.
Three specimens were found in April on station 2 near Seattle (mean
depth, 172-216 m; silt). There is also material of this species in the
U.S. National Museum, collected by E. and C. Berkeley near Nanaimo,
and by M. H. Pettibone in the San Juan Archipelago and Puget Sound,
from the intertidal zone to 60 m depth, from rocky and sandy bottoms,
as well as from “mud and algae” (USNM Nos. 26846—49; 26851; 26853;
32460; 32673).
The prostomium is cordiform. The occipital notch is not always
distinct except by its color. The second and third tentacular segments
1 Contribution No. 466 from the Department of Oceanography, University of
Washington, Seattle, Washington 98105. The material was collected with support
from U.S. Public Health Service Grant GM 10817. The preparation of the paper
was partially supported by Contract AT(45-1)-1725 of the U.S. Atomic Energy
Commission (ref. RLO-1725-101). The support is gratefully acknowledged. The
U.S. National Museum kindly lent us specimens for study. Drs. M. L. Jones and
M. H. Pettibone offered friendly criticism of the manuscript.
26—Proc. Brow. Soc. WasH., Vou. 81, 1968 (223)
224 Proceedings of the Biological Society of Washington
0.05mm
t_____
0.025 mm
(f-i)
To)
0.015 mm
Ficure 1. Eulalia macroceros: a, papilla from proboscis; b, outline
of median parapodium; c, detail of seta. Protodorvillea recuperata new
species: d, dorsal view of anterior end slightly from the right side; e,
posterior view of 40th parapodium of paratype; f—i, setae from 40th
parapodium: f, capillary supraacicular seta; g, Y-shaped seta; h, upper-
most and i, lowermost compound subacicular setae.
Two New Polychaetes 225
are separated dorsally and are clearly visible. There is a small parapodial
lobe, without setae, on the third tentacular segment. The proboscis is
fully covered, feltlike, with papillae about 120 » long (Fig. la). Dorsal
cirri are broadly lanceolate (Fig. 1b) and reach almost to the mid-
dorsum. The shafts of the setae are rounded with numerous small spines
which are slightly bent inward (Fig. lc). The blades are long (approxi-
mately 150 «) and strongly serrated throughout their length. Complete
animals are a few centimeters long. All specimens belong to the Mediter-
ranean species, which is not identical with Sige fusigera Bergstrom from
Northern Europe (Banse, 1959).
Numerous eggs of 65 to 70 uw diameter occur in June and August; all
material from the U.S. National Museum was collected in the summer.
The color in alcohol is yellow-brown with a uniform violet-brown
dorsum.
FAMILY DORVILLEIDAE
Protodorvillea recuperata new species
Figure 1d-i
Types: Holotype, USNM 36282. Paratypes (2), USNM 36283. All
types from station 5, 47°10’48” N, 122°50’'00” W; 15-37 m depth, in
medium sand (May, 1963).
‘Description: The holotype has 67 setigers and is 5.3 mm long (with-
out anal cirri). Its greatest width, at about the 30th setiger, is 0.60 mm
with and 0.34 mm without parapodia. A paratype, also complete, has
55 setigers and is 4.8 mm long. There are some additional specimens.
The blunt prostomium (Fig. 1d) carries two club-shaped antennae
and two pseudo-annulated palps with small, oval terminal joints. Only
one of the preserved specimens has eyes. No noteworthy features beyond
those common for Protodorvillea and Stauronereis could be seen in the
undissected jaws. The outer row of the upper jaws contains 20 to 25
loose plates. The prostomium is followed by two apodous segments. All
subsequent segments have parapodia with dorsal and ventral cirri inserted
near their tips, and a distinct presetal lip (Fig. le) which occasionally
may be withdrawn. There are no aciculae in the dorsal cirri. A capillary
seta with fine teeth (Fig. 1f) and a dorsal Y-shaped seta (Fig. lg) are
present from the first setiger. Ventrally there are one, rarely two,
compound setae with fairly long blades (Fig. lh), two to four similar
setae with blades about half as long, and a lowermost compound seta
with a short blade (Fig. li). The shafts of these setae have serrated
tips; the tips of the blades are bifid with subterminal spines resembling
those of some Hesionidae. The pygidium is flattened and carries one
pair of annulated dorsal cirri about 0.5 mm in length, and another pair
of about 0.1 mm length.
The specimens are without color. The intestines are almost empty,
but boluses in the posterior part of some animals contain some small
sand grains.
226 Proceedings of the Biological Society of Washington
The name refers to an accident during the preparation of the descrip-
tion.
Diagnosis: A small Protodorvillea species with short, smooth, club-
shaped antennae. Slender parapodia with a presetal lip. A dorsal cirrus
and a Y-shaped seta from the first parapodium onward.
Habitat: Southern Puget Sound, on stations 5, 6, and 8; 10 to 40 m
depth, on medium sand with 10 to 25 percent mud, and some gravel.
Differential Diagnosis: The genus Protodorvillea has been established
by Pettibone (1961). Since then, P. biarticulata has been described by
Day (1963), and P. minuta by Hartman (1965); synonyms have been
discussed by Banse and Hartmann-Schréder (1964). P. recuperata can
be distinguished from the other species with antennae, well-developed
palps, and dorsal cirri on the first setiger, i.e. P. biarticulata Day, P.
gracilis (Hartman), and P. kefersteini (McIntosh), as follows: The
antennae are fairly short and not articulated as in P. biarticulata; the
articulation is indistinct also in P. kefersteini. There is a presetal lip
which is replaced by a postsetal lip in P. kefersteini; lips are entirely
absent in P. gracilis.
Stauronereis japonica ( Annenkova)
Staurocephalus japonica.—Ushakov, 1955, p. 246.
One specimen from station 8 in southern Puget Sound (10-21 m
depth, medium sand). New for the eastern Pacific. Previously known
from the Seas of Japan and Okhotsk.
FAMILY TEREBELLIDAE
Lanassa venusta (Malm)
L. venusta.—Hessle, 1917, p. 205.
There are six poorly to moderately well-preserved specimens. Eye-
spots are not visible. The lateral lappets are very small. Capillary setae
start on the fourth segment, and are found on 11 setigers. They have
narrow wings and finely serrated tips. The uncini are arranged in double
rows in the four first abdominal setigers as described by Hessle (1917).
A specimen collected in May carried moderately numerous eggs.
Found on stations 3 and 4 near Seattle, and 6 in southem Puget
Sound. A doubtful identification comes from station 8 nearby. Exclud-
ing the shallow, sandy last-named station, the depth range of the records
is 17-40 m, very fine to fine sand. New for Puget Sound. Previously
known in the North Pacific from the Bering Sea.
Lysilla pacifica Hessle
L. pacifica—Ushakov, 1955, p. 403.
Found on station 4 near Seattle. 10-18 m depth, very fine sand. New
for the eastern North Pacific. Previously known from the Sea of Japan
and the Bonin Islands.
Two New Polychaetes 2271
FAMILY SABELLIDAE
Chone bimaculata new species
Figure 2a-j
Types: Holotype, USNM 36284. Paratypes, USNM 36281 (1) and
36280 (4). All from station 5, 47°10’48” N, 122°50'00” W; 15-37 m
depth, on medium sand (May, 1963).
Description: There are several complete, mature specimens in addition
to the types. Mature females with eight thoracic and 40 to 45 abdominal
setigers are up to 17.5 mm long, of which 7.5 mm are contributed by
the tentacular crown. The greatest width of the body is 1.0 to 1.1 mm.
The tentacular crown consists of about eight pairs of pinnate radioles
connected by a very thin membrane for at least half of their length.
The radioles have a flange beyond the membrane but the free end
beyond the pinnate section is narrow (Fig. 2a). The free end, 2 mm
long in the holotype, contributes one-fourth to one-third of the total
length of the radioles. Ventrally and dorsally there are one to two pairs
of free filaments, about half as long as the tentacular crown.
The collar (Fig. 2b) is higher ventrally than laterally. Dorsally it is
folded; in specimens other than the holotype, the folding is more clearly
seen than shown in the figure. Setigers are weakly biannulate. On the
first setiger, there are otocysts, but no eyespots; on the second setiger
is a girdle of glandular cells, posterior to the setae. The fecal groove
is not very distinct. The pygidium is pointed (Fig. 2c).
In the thoracic setigers, except for the first one, there are five to six
limbate setae (Fig. 2d), six to seven spatulate setae with long tips (Fig.
Ye), and the same number of bayonet setae (Fig. 2f) per ramus. The
latter setae accompany the spatulate bristles, just reaching to the base
of their blades. There are six to eight long-handled hooks in setigers 2
through 8, with delicate wings at the backs of their crowns (Fig. 2g).
In the abdomen there are two bundles of three or four capillary setae
on each side; these have broad wings but otherwise are more slender
than those of the thorax (Fig. 2h). About 12 uncini (Fig. 2i) occur
in the midabdomen per ramus.
The color of the preserved animal is pale yellow. Approximately 10
eggs per segment are found in the thorax and the anterior two-thirds of
the abdomen; they are polygonal, up to 200 u X 150 u.
To distinguish the new species more easily from C. suspecta Kroyer,
specimens were stained in a dark-green solution of methyl green in 70
to 80 percent ethyl alcohol for about 15 min, following the method used
by Hofsommer (1913). The color was differentiated for some time in
70 to 80 percent alcohol. The most intensive coloration in the three
animals checked is found laterally on the collar. The margins of the
collar, an area at the distal, ventral side of the collar, the dorsal folded
part of the collar, and the ring on the second setiger, are free of glands
that are stained. Less clearly distinguished from the diffuse color of
the stained epidermis is an unstained ring around the middle of each
setiger. On the ventral side of one well-preserved specimen occur
228 Proceedings of the Biological Society of Washington
0.02 mm
Ficurre 2. Chone bimaculata new species: a, tip of radiole; b, dorsal
view of anterior end of the body with base of tentacular cirri; c, dorsal
view of posterior end of body; d, thoracic capillary seta; e, thoracic
spatulate seta; f, thoracic bayonet-shaped seta; g, thoracic uncinus; h,
abdominal capillary seta; i, abdominal uncinus; j, color pattern of stained
animals (see text); body outline schematic.
Two New Polychaetes 229
unstained areas parallel to this ring, from the third thoracic setiger
through the first few abdominal setigers (Fig. 2j).
The name refers to the two glandular areas on the collar, which are
very distinct in stained specimens.
Diagnosis: A Chone species with a girdle of glands on the second
setiger. Tentacular crown relatively long; free ends of the radioles long
and thin. Collar protruding ventrally farther than laterally; folded
dorsally. Spatulate setae with a long mucron and accompanied by
bayonet setae.
Differential Diagnosis: The following Chone species are known to
have a glandular ring on the second setiger: C. cincta Zachs, recently
considered to be a synonym of C. teres Bush (Chlebovich, 1916); C.
filicaudata Southern; C. paracincta Hartmann-Schréder; C. rosea Hart-
mann-Schréder; and C. suspecta Kroyer. All have spatulate setae with
distinct points, but bayonet-shaped bristles have been reported only for
C. filicaudata, C. striata, C. suspecta, and C. teres. Chone filicaudata is
distinguished by the abdominal appendage and its abdominal uncini,
which have narrow gaps between the rostra and the bases. Chone teres,
when adult, is very much larger (56 mm) than all other species with
glandular rings, and among the group with bayonet bristles it is the
only species where the tentacular crown is relatively short (about one-
seventh of the total length). Chone striata has a relatively short tentacular
crown (4.8 mm out of 16 mm) and an almost level collar; also the free
ends of the radioles are broad. The glandular ring of C. suspecta is
uniquely situated on the third setiger; also, the bayonet bristles are not
smooth as in C. bimaculata. Finally, the mucus glands on the collar of C.
suspecta and of C. bimaculata are arranged differently. In the former
species the collar is uniformly stained excepting the margin which
remains whitish. There is a dark spot near each bundle of setae on
the first setiger. Paired, small, unstained areas, parallel to the full
unstained ring in the middle of the segments, appear on the second
setiger.
LITERATURE CITED
BansE, K. 1959. Uber die Polychaeten-Besiedlung einiger submariner
Hohlen. Ergebnisse der Gsterreichischen Tyrrhenia-Expedi-
tion, 1952. Teil XII. Pubbl. Staz. Zool. Napoli, vol. 30
(Suppl.), 417-469.
BaNnsE, K., aND G. HARTMANN-SCHRODER. 1964. Synonyms of Proto-
dorvillea egena (Ehlers) (Eunicidae, Polychaeta). Proc.
Biol. Soc. Wash., vol. 77, 241-242.
BERKELEY, E., AND C. BERKELEY. 1948. Annelida, Polychaeta Errantia.
Canad. Pac. Fauna, no. 9b(1). Fish. Res. Bd. Canada,
1-100.
CuLeEBovicu, V. V. 1961. [Littoral polychaetes from the Kurile Is-
lands.] Akad. Nauk SSSR, Issled. Dal’nevost. morei SSSR,
vol. 7, 151-260. [In Russian.]
230 Proceedings of the Biological Society of Washington
Day, J. H. 1963. The polychaete fauna of South Africa. Part 8. New
species and records from grab samples and dredgings. Bull.
Brit. Mus. (Nat. Hist.) Zool., vol. 10, 383-445.
HartTMan, O. 1965. Deep-water benthic polychaetous annelids off
New England to Bermuda and other North Atlantic areas.
Allan Hancock Found. Publ., Occas. Pap. 28, 1-378.
HesstE, C. 1917. Zur Kenntnis der terebellomorphen Polychaeten.
Zool. Bidr. Uppsala, vol. 5, 39-258. —
HorsomMeEr, A. 1913. Die Sabelliden-Ausbeute der Poseidon-Fahrten
und die Sabelliden der Kieler Bucht. Wiss. Meeresunters,
Abt. Kiel, N. F., vol. 15, 305-364.
Liz, U. Impress. A quantitative study of the benthic infauna in Puget
Sound (Washington), 1963-1964. FiskDir. Skr. HavUnders,
vol. 14.
PETTIBONE, M. H. 1961. New species of polychaete worms from the
Atlantic Ocean, with a revision of the Dorvilleidae. Proc.
Biol. Soc. Wash., vol. 74, 167-186.
MEE OCTS
Vol. 81, pp. 231-240 30 August 1968
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
Z Zh i |
BIOLOGICAL INVESTIGATIONS OF THE DEEP SEA.
38. A NEW WESTERN ATLANTIC DIPSACASTER
(ECHINODERMATA, ASTEROIDEA), WITH THE
DISTRIBUTION OF KNOWN SPECIES® |
By JeRALD A. HALPERN
Institute of Marine Sciences, University of Miami
Four sea stars collected north of the Little Bahama Bank by
the R/V Gerda of the Institute of Marine Sciences, University
of Miami, represent a new species of Dipsacaster. The dis-
covery of this species extends the range of the genus Dipsa-
caster to the western Atlantic (Table 1).
The astropectinid genus Dipsacaster was erected by Alcock
(1893a:87) for a new species, D. sladeni, from the Andaman
Sea. Dipsacaster may be distinguished from related genera by
the inferomarginal plates, which project beyond the supero-
marginals, forming the lateral margins. Other distinguishing
characters are the large actinal intermediate areas, the large
papular area, the presence of an anal aperture, gonads that
extend far along the arms, and a large madreporite covered
by paxillae.
This research was supported by the National Geographic
Society through a grant for investigations of the deep-sea
fauna, and by the National Science Foundation through grant
GB-4936. The biological operations of R/V Gerda have been
supported by National Science Foundation grant GB-1204.
The author is grateful to these agencies. Thanks are also
extended to Lowell P. Thomas, Gilbert L. Voss, and Frederick
M. Bayer for their assistance.
The terminology employed has been adopted mainly from
the works of Walter K. Fisher, Ailsa M. Clark, and F. Jen-
1 Contribution No. 947 from the Institute of Marine Sciences, University of Miami.
27—Proc. Brow. Soc. WAsH., Vou. 81, 1968 (231)
232. Proceedings of the Biological Society of Washington
OW
FicurE 1. a, Holotype, abactinal view, 0.9 .—b, Paratype, abactinal
view, 3.5&.
senius Madsen. The number of marginal plates represents
those from one arm tip to the tip of an adjacent arm.
The abbreviations USNM and UMML refer to the United
States National Museum and the museum of the Institute of
A New Atlantic Starfish 233
Marine Sciences (University of Miami Marine Laboratory ),
respectively.
Dipsacaster antillensis new species
Material Studied: Holotype: R = 44 mm, r = 16 mm, R/r = 2.7;
27°56’ N, 78°40’ W, 860-897 m, R/V Gerda sta. G-182, 22 July 1963,
USNM E10854. Paratypes: 27°49’N, 78°45’W, 824 m, R/V Gerda sta.
G-403, 20 September 1964, UMML 40.210, 3 specimens.
Diagnosis: Inferomarginal plates up to four times as wide as long,
covered by hemispherical granules; plates with tuft of one to four short,
blunt, poorly developed spines on distal portion of free edge. Median
areas of arms distinctive, due to their elevation and smaller paxillae.
Arms tapering gradually.
Description: Five arms. R = 44 mm, r = 16 mm, R/r = Q2.7r.
The general form is stellate; the arms taper gradually to a bluntly
pointed tip. The body is flat, with a slightly elevated carinal ridge along
the arms. The interbrachial arcs are wide, round.
The abactinal surface is paxillose, extending to the terminal plates
on the arms. The paxillae are regularly arranged, and conspicuously
smaller along the median areas of the arms. Each paxilla consists of a
convex base, from which a short, broad pedicel arises. Each pedicel
bears about twenty long, thin, laterally directed spinelets around its
periphery; and about six short, stout, blunt, dorsally directed spinelets
in the middle. Papulae are found throughout the abactinal surface;
there are usually six surrounding each plate.
There are forty-five marginal plates in each series. The marginal
fascioles in the interbrachial arcs are wide and deep.
The massive superomarginal plates are three to four times as broad
as long in the interbrachial arc, diminishing in width distally, so that
they are twice as broad as long in the middle of the arms and almost
square near the terminal plate. The superomarginals are completely
covered by large, hemispherical, closely crowded granules. The terminal
plate is small; its distal end is notched, proximal end truncate.
The inferomarginal plates correspond with and are broader than the
superomarginals. They are covered by large, round, crowded granules,
becoming spiniform in the fasciolar grooves and toward the free edge;
these spinelets are delicate and thin in the fasciolar grooves, larger and
lanceolate along the free edge. There is a tuft of one to four short,
stout, conical or truncate spines in the distal corner of the free edge.
The actinal intermediate area is large, the plates being arranged in
a regular series parallel to the inferomarginals. The actinal plates are
strongly carinate; each is covered by a tuft of spinelets. These spinelets
are slender and pointed outward along the periphery of the plate; they
are shorter, stouter, often clavate, and facing the marginals in the center
of the plate. A wide channel separates the actinal and adambulacral
plates.
234 Proceedings of the Biological Society of Washington
|
Ficure 2. a, Paratype, abactinal surface, 8; note madreporite at
lower left.—b, Paratype, actinal view, 3x.
The adambulacral plates are moderately large and have a curved
furrow margin. There are six to seven long, blunt, compressed furrow
spines; the longest spines are in the center of each series. There are
two or three irregular subambulacral rows of three to five tapering,
A New Atlantic Starfish 235
FicurE 3. a, Paratype, actinal view, 3.5><.—b, Paratype, actinal view,
IOLOS<.
pointed spines. The subambulacral spines are about half as long as
and more slender than the furrow spines.
The mouth plates are large and prominent. Each mouth plate bears
seven to ten closely crowded furrow spines; these spines are strongly
236 Proceedings of the Biological Society of Washington
TABLE 1.
Species
D. eximius
Fisher, 1905
D. anoplus
Fisher, 1910
D. borealis
Fisher, 1910
D. laetmophilus
Fisher, 1910
D. nesiotes
Fisher, 1906
D. pretiosus
(Déderlein, 1902)
D. grandissimus
Goto, 1914
D. magnificus
(H. L. Clark, 1916)
D. diaphoris
Fisher, 1913
D. imperialis
Fisher, 1917
D. pentagonalis
Alcock, 1893a
D. sladeni sladeni
Alcock, 1893
D. farquharsoni
Macan, 1938
D. sladeni capensis
A. M. Clark, 1952
Distribution
off California, Monterey Bay to
San Diego, 377—961 m
Washington to San Diego, 549—-1,464 m
Bering Sea, 2,200 m
Pribilof Islands, 220-640 m
Sea of Okhotsk, Kurile Islands, 2,000 m
Bering Sea and south of Aleutian
Islands, 221-642 m
Pribilof Islands, 220-235 m
southeast of Alaska Peninsula, 1,272 m
Hawaiian Islands, 518-564 m
Molucca Islands, 498-527 m
Tokyo ard Sagami Bays, Japan, 20-200 m
Uraga Channel, Misaki, Toyama Bay,
off Miho, Japan, 20-480 m
off Misaki, Japan, 640 m
Great Australian Bight, 146-220 m
Cook Strait, New Zealand, 101-115 m
Philippine Islands, Sulu Sea, 701—1,473 m
Philippine Islands, 622 m
Andaman Sea, 205 m
Andaman Sea, 458 m
Maldives, 229 m
Cape of Good Hope, 110-329 m
Cape of Good Hope, 240-631 m
Cape of Good Hope, 190-403 m
off East London, S. Africa, and Cape
of Good Hope, 170-326 m
off Cape Town, S. Africa, 329-357 m
Distribution of known species of Dipsacaster.
Source
Fisher
(1911, p. 90)
Fisher
(1911, p. 100)
Baranova
(1957, p. 156)
Djakonov
(1950, p. 28)
Djakonov
(1958, p. 286)
Fisher
(1911, p. 94)
Baranova
(1957, p. 157)
Fisher
(1911, p. 97)
Fisher
(1906, p. 1028)
Fisher
(1919, p. 145)
Déderlein
(1921, p. 22
Okada et al.
(1966, p. 154)
Goto
(1914, p. 257)
H. L. Clark
(1916, p 32)
Fell
(1958, p. 6)
Fisher
(1919, p. 153)
Fisher
(1919, p. 148)
Alcock
(1893a, p. 172)
Alcock
(1893a, p. 172)
Macan
(1938, p. 343)
Bell
(1905, p. 243)
H. L. Clark
(1923, p. 247)
H. L. Clark
(1925, p. 3)
Mortensen
(1933, p. 237)
A. M. Clark
(1952, p. 194)
A New Atlantic Starfish 237
compressed and stouter than the adambulacral furrow spines. The two
spines behind the median spines are very stout; the rest of the mouth
plate pair is covered by short, blunt spines which increase in size toward
the suture.
The madreporite is very large, and is covered by paxillae. It is located
in the middle of an interradius, close to the superomarginals. The anal
pore is in an elevated area in the center of the disk; the paxillae on
this anal elevation are short and crowded.
There are no pedicellariae. Superambulacral ossicles are present as
slender rods. The tube feet are large and pointed; the ampullae are
double. The gonads are in a longitudinal dichotomous series on either
side of the median radial area, extending slightly beyond the middle of
the arm.
Measurements, ratios, and number of marginal plates in each of the
paratypes as follows:
R (mm) r (mm) R/r Marginals
30 elt 2.7 36
36 14 2.6 38
45 19 2.4 36
Distribution: It is known only from a very small area just north of
the Little Bahama Bank, with a bathymetric range of 824-897 m.
Remarks: The only previous record of Dipsacaster in the Atlantic
Ocean is that of D. sladeni (Table 1). However, this can not be con-
sidered a true Atlantic species. It ranges from the Andaman Sea to the
Cape of Good Hope, the latter being only a western extension of its
Indian Ocean range. Thus, Dipsacaster antillensis represents the first
species of the genus known from the Atlantic Ocean proper.
Dipsacaster antillensis does not appear to be closely related to any of
the known species of Dipsacaster. The extremely broad marginal plates
serve to distinguish it from all the others except D. pretiosus (Déderlein ),
D. grandissimus Goto, and D. farquharsoni Macan. The inferomarginals
of the first two are spinose; in the last, the arms are more strongly
tapered and the inferomarginal spines are at least four times larger than
in D. antillensis.
LITERATURE CITED
Aucock, A. 1893a. Natural history notes from H. M. Indian Marine
Survey Steamer “Investigator,” Commander C. F. Oldham,
R. N., commanding. Series 2, No. 7. An account of the col-
lection of deep-sea Asteroidea. Ann. Mag. nat. Hist., (6)
11: 73-121, pls. 4-6.
1893b. Natural history notes from H. M. Indian Marine
Survey Steamer “Investigator,” Commander C. F. Oldham,
R. N., commanding. Series 2, No. 9. An account of the deep-
sea collection made during the season of 1892-93. J. Asiat.
Soc. Beng., 62: 169-184, pls. 8, 9.
238 Proceedings of the Biological Society of Washington
Bett, F. Jerrrey. 1905. The Echinoderma found off the coast of
South Africa. Part 2. Asteroidea. Mar. Invest. S. Afr., 3:
2A 253"
BarAnova, Z. I. 1957. Echinoderms of the Bering Sea. Invest. Far-
east Seas USSR, 4: 149-266, 19 figs. (In Russian.)
Cuark, AitsA McGown. 1952. Some echinoderms from South Africa.
Mrans. roy. Soc. S. Ati) Go.) 193=22) thigsy al —oeaplaelivn
Cuark, Husperr LyMAn. 1916. Report on the sea lilies, starfishes,
brittle stars, and sea urchins obtained by the F.I.S. “En-
deavour” on the coasts of Queensland, New South Wales,
Tasmania, Victoria, South Australia, and Western Australia.
Fisheries, Sydney, 4 (1): 1-123, pls. 1-44.
1923. The echinoderm fauna of South Africa. Ann. S. Afr.
Mus., 13: 235-310, pls. 8-18.
1925. Echinoderms from the South African fisheries and
marine biological survey. Part 2. Sea-stars (Asteroidea).
Invest. Rep. Fish. Mar. biol. Surv. S. Afr., 4: 1-33, pls. 1-7.
Dyaxonoy, A. M. 1950. Starfish of the Soviet Union. Tableaux ana-
lytiques faune USSR, 34: 1-203, figs. 1-212. (In Russian.)
1958. Echinodermata, excluding Holothuroidea; collected |
by the Kurile-Sakhalin expedition in 1947-49. Invest. Far-
east Seas USSR, 5: 271-357, 40 figs. (In Russian. )
DO6DERLEIN, Lupwic. 1902. Japanische Seesterne. Zool. Anz., 25:
326-335.
1921. Die Asteriden der Siboga-Expedition. I. Porcellanas-
teridae, Astropectinidae, Benthopectinidae. Siboga Exped.,
46 (91): 1-47, pls. 1-13, 7 text figures.
Fett, H. BArRAcLoucH. 1958. Deep-sea echinoderms of New Zea-
land. Zool. Publ. Vict. Univ. N. Z., 24: 1-40, pls. 1-5.
FisHER, WALTER KeENrRIcK. 1905. New starfishes from deep water off
California and Alaska. Bull. Bur. Fish., Wash., 24: 291-320.
1906. The starfishes of the Hawaiian Islands. Bull. U.S.
Fish Comm., 1903, 23 (3): 987-1130, pls. 1-49.
1910. New starfishes from the North Pacific. I. Phanero-
zonia. Zool. Anz., 35: 545-553.
1911. Asteroidea of the North Pacific and adjacent waters.
Part 1. Phanerozonia and Spinulosa. Bull. U.S. Nat. Mus.,
76 (1): 1-420, pls. 1-192.
1913. Four new genera and fifty-eight new species of star-
fishes from the Philippine Islands, Celebes, and the Moluccas.
Proc. U.S. Nat. Mus., 43: 599-648.
1917. New starfishes from the Philippine Islands, Celebes,
and the Moluccas. Proc. biol. Soc. Wash., 30: 89-93.
1919. Starfishes of the Philippine Seas and adjacent waters.
Bull. U.S. Nat. Mus., 100 (3): 1-547, pls. 1-155.
Goro, SEiraAro. 1914. Japanese Asteroidea. J. Coll. Sci. Tokyo, 29
(1): 1-808, pls. 1-19.
eee a
A New Atlantic Starfish 239
Macan, T. T. 1938. Asteroidea. Sci. Rep. John Murray Exped., 4:
323-435, pls. 1-6, 12 text figures.
MortTENSEN, THEODOR. 1933. Papers from Dr. Th. Mortensen’s Pacific
Expedition 1914-16. 65. Echinoderms of South Africa. (As-
teroidea and Ophiuroidea). Vidensk. Medd. dansk naturh.
Foren. Kbh., 93: 215-400, pls. 8-19, 91 text figures.
OKADA, YAICHERO, ICHITARO SAKAMOTO, RYOHEI AMANO, AND YOSHIAKI
Tominaca. 1966. Preliminary report of the benthic bio-
logical survey in Suruga Bay. J. Fac. Oceanogr. Tokai Univ.
(1966), 1: 135-155, pls. 1-4.
940 Proceedings of the Biological Society of Washington
Y.0€ 74
Vol. 81, pp. 241-250 30 August 1968
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
THREE NEW STOMATOPOD CRUSTACEANS
FROM THE INDO-MALAYAN AREA
By Raymonp B. MANNING
Smithsonian Institution, Washington, D.C.
In his monograph on the Indo-West Pacific stomatopods,
Kemp (1913) noted that Lanchester’s (1901) record of Chlo-
ridella chlorida from the Malay Peninsula was probably in-
correct. In April 1967, I had the opportunity to visit the
Museum of Zoology, Cambridge, England, and examine that
specimen as well as the small series of specimens which
Lanchester reported as Lysiosquilla spinosa. Both lots rep-
resent undescribed species which are described and illustrated
herein.
Among the unidentified stomatopods in the collection of
the Division of Crustacea, U.S. National Museum are two
specimens representing two new species of Clorida. One of
these specimens is conspecific with the specimen Lanchester
recorded from the Malay Peninsula while the other belongs
to another undescribed species, characterized below.
I am indebted to Dr. C. B. Goodhart and Dr. K. A. Joysey
for their assistance during my visit to Cambridge and for the
loan of the specimens. The illustrations were made by my
wife, Lilly. Support of this study by the Office of Oceanog-
raphy and Limnology, Smithsonian Institution, is gratefully
acknowledged.
In the descriptive accounts, TL refers to Total Length and
CL to Carapace Length. Other terms have been defined in
earlier papers (Manning, 1966, 1968).
Austrosquilla malayensis new species
Figure 1
Lysiosquilla spinosa.—Lanchester, 1901, p. 554 [not L. spinosa (Wood-
Mason, 1875)].
28—Proc. Biou. Soc. WaAsH., Vou. 81, 1968 (241)
242, Proceedings of the Biological Society of Washington
FicurEe 1. Austrosquilla malayensis new species, male holotype: a,
anterior portion of body; b, ischiomeral articulation of left raptorial
claw; c, last abdominal somite, telson, and uropod; d, outline of telson
in ventral view; e, basal prolongation of uropod, ventral view. (Setae
omitted. )
Holotype: 1 6, TL 17.0 mm; Pulau Bidan, Penang, Malay Peninsula
[Malaysia]; Skeat Expedition, 1899; Museum of Zoology, Cambridge.
Paratypes: 1 6, CL 3.0 mm; 3 9, TL 15.8-17.3 mm; data as in
holotype.
Description: Eye small, cornea subglobular, set obliquely on stalk,
faintly overhanging stalk ventrolaterally; eyes extending almost to end
of antennular peduncle; ocular scales erect, fused.
Antennular peduncle short, less than half as long as carapace; dorsal
processes of antennular somite visible lateral to rostral plate as sharp,
anteriorly-directed spines.
Antennal scale small, ovate, about 4 carapace length; antennal
Three New Stomatopods 243
papillae not clearly visible in type-series, but at least 1 ventral papilla
present.
Rostral plate triangular, slightly broader than long, without apical
spine.
Carapace smooth, narrowed anteriorly, without carinae or spines.
Dactylus of claw with 16-21 teeth, antepenultimate not longer than
penultimate; outer margin of dactylus flattened or slightly concave
proximally, convex distally, with rounded, obtuse basal prominence;
propodus fully pectinate, with 4 proximal movable spines, second small-
est, remainder subequal; dorsal ridge of carpus terminating in blunt
lobe; ischium almost as long as merus, with sharp subdistal spine on
outer ventral surface.
Mandibular palp absent; 4 epipods present.
Lateral processes of sixth and seventh thoracic somites subtruncate,
rounded anterolaterally and posterolaterally; basal segments of first 2
walking legs with inner and outer spines, last leg in male with outer
spine only; eighth somite with, at most, a low obtuse tubercle on ventral
midline.
Abdomen smooth, depressed, unarmed dorsally; sixth somite with
small, fixed ventrolateral spine overhanging articulation of each uropod.
Telson broader than long, dorsal surface with median and 4 lateral
obscure, obtuse marginal prominences; posterior armature on each side
of midline, consisting of 7-10 submedian denticles, 1 movable submedian
tooth, 4 fixed intermediate denticles, first and third shorter than second
and fourth, 1 fixed intermediate tooth, 1 fixed lateral denticle, and the
fixed lateral tooth; most teeth and denticles submarginal.
Basal segment of uropod with inner and outer carina, inner terminat-
ing in dorsal tooth; proximal segment of exopod shorter than distal, with
4 curved, movable spines on outer margin, distal spine extending past
midlength of distal segment; inner distal lobe of proximal segment of
exopod with 3 stiff setae; endopod slender, triangular, with strong fold
on outer proximal margin; inner spine of basal prolongation longer than
outer.
Color: Preserved specimens completely faded.
Measurements: Holotype, TL 17.0 mm; other measurements, in mm:
carapace length, 3.1; rostral plate length, width, 2.2, 2.4; fifth abdominal
somite width, 3.3; telson length, width, 2.0, 2.9.
Discussion: Although the five specimens reported here are obviously
young and in very poor condition, I have no hesitation in assigning them
to a new species. All specimens clearly show the distinguishing features
of the genus, the ischial spine on the claw and the four fixed intermediate
denticles on the telson.
Austrosquilla now contains three species. A. osculans (Hale) and
A. vercoi (Hale), both redescribed by Manning (1966), occur only off
Australia. A. malayensis differs from both other species in lacking
posterolateral spines on the sixth abdominal somite and in having more
than one dorsal prominence on the telson. The new species resembles
244 Proceedings of the Biological Society of Washington
Ficure 2. Clorida malaccensis new species, female holotype: a, ante-
rior portion of body; b, carpus, propodus, and dactylus of raptorial claw;
c, outline of exposed thoracic somites; d, last abdominal somite, telson,
and uropod; e, basal prolongation of uropod, ventral view. (Setae
omitted. )
A. vercoi in the large number of teeth on the claw and the triangular
rostral plate, but it further differs from that species in having more
submedian denticles on the telson (7-10 instead of 6) and fewer stiff
setae on the inner, distal lobe of the proximal segment of the uropodal
exopod (3 instead of 11).
Etymology: The name is derived from the type-locality.
Clorida malaccensis new species
Figure 2
Holotype: 1 2, TL 55.8 mm; Strait of Malacca; November, December
1961; U.S. Navy Hydrographic Office; USNM 124769.
Description: Eye small but elongate, extending to end of first segment
of antennular peduncle; stalk inflated, greatest width about half eye
Three New Stomatopods 245
length; cornea small, bilobed, broader than distal portion of stalk, comea
width slightly more than half eye length; ocular scales rounded, fused
medially.
Antennular peduncle elongate, more than half as long as carapace;
dorsal processes of antennular somite produced into short, sharp, ante-
riorly-directed spines.
Antennal peduncle elongate, first segment extending about to cornea;
scale slender, curved, less than half as long as carapace.
Rostral plate longer than broad, triangular, lateral margins sinuous,
converging on rounded apex.
Carapace narrowed anteriorly, lateral margin strongly concave; cara-
pace completely lacking carinae except for marginals on posterolateral
borders and anteriorly-reflected portions of marginals; posterior margin
slightly concave; anterior margins of lateral plates slope posterolaterally
to strong anterolateral spines which do not extend to base of rostral
plate.
Raptorial claw stout; dactylus with 5 teeth, proximal very small, outer
margin of dactylus with 2 lobes, proximal small, distal much larger,
rectangular; dorsal ridge of carpus terminating in low tubercle.
Mandibular palp and 3-4 epipods present.
Exposed thoracic somites without submedian carinae, last 3 somites
with low, inconspicuous, intermediate carinae; lateral process of fifth
somite a slender spine, directed anterolaterally; fifth somite also with
slender ventrolateral spine on each side; lateral processes of sixth and
seventh somites rounded laterally, unarmed posteriorly; ventral keel of
eighth somite low, rounded.
Abdomen broad, depressed, lacking submedian carinae on first 5
somites; abdominal carinae spined as follows: submedian, 6; inter-
mediate, 5-6; lateral, 5-6; marginal, 6; dorsal surface of sixth somite
rough, irregular, between submedian and intermediate carinae; sixth
somite with sharp ventrolateral spine in front of articulation of each
uropod.
Telson broader than long, with 3 pairs of sharp marginal teeth, sub-
medians with movable apices; carinae of marginal teeth short, tuberculate
dorsally; dorsal surface of telson with about 7 irregular, curved rows of
tubercles and several tubercles scattered on anterior surface; denticles
sharp, 3-4, 8, 1; ventral surface of telson with long, well-developed
postanal keel.
Outer margin of proximal segment of uropodal exopod with 6 slender,
movable spines, last not extending beyond midlength of distal segment;
endopod slender, elongate; basal prolongation of uropod with 8-9 slender,
fixed spines on inner margin and broad rounded lobe on outer margin
of longer inner spine.
Color: Body completely covered with small, dark chromatophores
arranged in no discernible pattern; uropodal endopod dark; distal fifth
of proximal segment and inner half of distal segment of uropodal exopod
dark.
246 Proceedings of the Biological Society of Washington
Ficure 3. Clorida latispina new species, a-e female holotype: a,
anterior portion of body; b, eye, enlarged; c, lateral processes of exposed
thoracic somites; d, last abdominal somite, telson, and uropod; e, basal
prolongation of uropod, ventral view; f, telson of male paratype. (Setae
omitted. )
Measurements: Female holotype, TL 55.8 mm. Other measurements,
in mm: carapace length, 11.7; cornea width, 1.5; eye length, 2.9; stalk
width, 1.4; rostral plate length, width, 2.3, 1.6; fifth abdominal somite
width, 14.1; telson length, width, 9.0, 12.1.
Discussion: Clorida malaccensis resembles both C. microphthalma (H.
Milne-Edwards) and C. chlorida (Brooks), and it would key out to the
couplet containing those two species in the key provided by Manning
(1968). It differs from both in (a) color pattern, for neither of the
other two species has a pattern in which the body is completely
covered with small chromatophores; (b) in having a large prominent
lobe on the outer margin of the dactylus of the claw; (c) in eye shape
(see below); and (d) in having tubercles on the sixth abdominal somite
Three New Stomatopods 247
between the submedian and intermediate carinae. In addition, the
postanal keel in C. malaccensis is more prominent than in either of the
two other species. The shape of the claw will distinguish C. malaccensis
from all other species in the genus.
The eye of C. microphthalma differs in that the mesial margins are
not as convex as in C. malaccensis. The cornea is much smaller in C.
microphthalma (% greatest stalk width) than in C. malaccensis (sub-
equal to stalk width).
It seems likely that one of the epipods on ne holotype has been lost
and that normally four are found on each side.
The unusually-shaped claw of C. malaccensis resembles that found
in Acanthosquilla multifasciata (Wood-Mason), and a similar lobe on
the dactylus is found on males of some species of Harpiosquilla. The
hairs on the propodus and the peculiarly-shaped dactylus may both be
features which differ in males and females.
Etymology: The name is derived from the type-locality.
Clorida latispina new species
Figure 3
Chloridella chlorida.—Lanchester, 1901, p. 554 [not C. chlorida (Brooks,
1886) ].
PSquilla latreillei—Seréne, 1952, text fig. 15, pl. 1 (Fig. 4) [part].
Holotype: 1 9, TL 61.7 mm; China Sea, off Hongkong; 21°44’ N,
114°48’ E; Albatross Sta. 5303; 34 fms; black mud; 9 August 1908;
USNM 77929.
Paratype: 1 6, TL 77.1 mm; Khota Bharu, Kelantan, Malay Peninsula,
[Malaysia]; Skeat Expedition, 1899; Museum of Zoology, Cambridge.
Description: Eye small, not extending to middle of first segment of
antennular peduncle; stalk inflated, width almost *4 eye length; mesial
margin of stalk straight, outer margin convex; cornea very small, bilobed,
set obliquely on stalk, width % stalk width; ocular scales small, fused
into a rounded plate in male, a bilobed plate in female.
Antennular peduncle elongate, longer than carapace and rostral plate
combined; dorsal processes of antennular somite visible lateral to rostral
plate as small anterolaterally-directed spines.
Antennal peduncle elongate, first segment extending well beyond eye;
antennal scale slender, curved, length less than 1% carapace length.
Rostral plate short, rounded anteriorly, length % greatest width or
less.
Carapace strongly narrowed anteriorly, lateral margins concave, median
and intermediate carinae completely absent; posterior fourth of carapace
with sharp reflected portions of marginal carinae; posterior portion of
lateral carinae present in female, represented by longitudinal swelling
in male; posterior margin of carapace concave; anterior margins of
lateral plates sloping posterolaterally to well-developed anterolateral
spines which do not extend to base of rostral plate.
248 Proceedings of the Biological Society of Washington
Raptorial claw stout; dactylus with 5 teeth, outer margin of dactylus
convex or flattened, with basal notch; propodus of females with long
hairs proximally; dorsal ridge of carpus terminating in low spine.
Mandibular palp and 4 epipods present.
Exposed thoracic somites with faint submedian carinae on eighth
somite only; last 3 somites with well-developed intermediate carinae,
unarmed posteriorly; lateral process of fifth thoracic somite a short,
slender spine directed anterolaterally; fifth somite also with small ventro-
lateral spine on each side; lateral processes of sixth and seventh somites
not bilobed, obliquely truncate, each with posterolateral spine; antero-
lateral margin of process of seventh somite more angular than that of
sixth; median ventral keel of eighth somite an inconspicuous tubercle.
Abdomen broad, depressed, first 5 somites each with submedian
carinae, carinae divergent on each somite; abdominal carinae spined as
follows: submedian, 6; intermediate, 5-6; lateral, 5-6; marginal, 3-5;
dorsal surface of sixth somite between submedian and intermediate
carinae rough, irregular, with several short, oblique swollen ridges; sixth
somite with ventrolateral spine in front of articulation of each uropod.
Telson broader than long, with 3 pairs of sharp marginal teeth, sub-
medians with movable apices; prelateral lobes present; dorsal surface
irregular, with minute pits and tubercles; denticles sharp, 2-3, 7-8, 1; —
ventral surface ornamented with sharp postanal keel only; dorsal sculp-
ture of telson differing ii: male and female as noted below.
Telson of male: Length about 44 greatest width; median carina in-
flated; each marginal tooth with tuberculate basal swelling, those of
intermediate teeth largest; each side of dorsal surface with longitudinal
row of tubercles converging under apex of median carina and 3 curved,
inflated rows of tubercles between the submedian and intermediate
teeth; fourth row, at level of intermediate teeth, indicated on 1 side;
median third of telson, anterior to notch in median carina, with swollen
patch of tubercles.
Telson of female: Length about % greatest width; median carina
sharp; bases of marginal teeth tuberculate but not inflated; dorsal carina
of prelateral lobe sharp; each side of dorsal surface with longitudinal
row of sharp tubercles converging under apex of median carina and
3 curved rows of sharp tubercles between submedian and intermediate
teeth; median third of telson, anterior to notch in median carina, with
U-shaped carinae, open posteriorly.
Outer margin of proximal segment of uropodal exopod with 7 slender
movable spines, last not extending to midlength of distal segment; proxi-
mal segment of uropodal exopod shorter than distal, with short longi-
tudinal dorsal carina; endopod slender, curved; basal prolongation with
7-8 slender fixed spines on inner margin and broad rounded lobe on
outer margin of longer inner spine; basal segment with spine (female)
or angular projection (male), on proximal inner margin at articulation
of endopod.
Three New Stomatopods 249
Color: Completely faded in both preserved specimens.
Measurements: Male paratype, TL 77.1 mm; female holotype, TL
61.7 mm. Other measurements of male, in mm: carapace length, 13.3;
cornea width, 0.9; eye length, 2.4; stalk width, 1.8; rostral plate length,
width, 1.4, 2.5; fifth abdominal somite width, 17.3; telson length, width,
12.3, 15.4.
Discussion: C. latispina is most closely related to C. bombayensis
(Chhapgar and Sane) described in 1967 and based on a series of four
females taken in shallow water off Bombay, India. Both C. latispina
and C. bombayensis can be distinguished from all other species in the
genus by the presence of the posterolateral spines on the lateral processes
of the sixth and seventh thoracic somites. C. decorata Wood-Mason also
has spines on these lateral processes, but in that species the anterolateral
angles of the lateral processes are armed rather than the posterolateral
angles.
C. latispina differs from C. bombayensis as follows: (a) the spines
of the dorsal processes of the antennular somite are smaller; (b) the
rostral plate is rounded anteriorly, not triangular; (c) the lateral process
of the fifth thoracic somite is slenderer, smaller, and not strongly curved
anteriorly; (d) submedian carinae are present on the eighth thoracic
somite but not on the seventh; and (e) the area between the submedian
and intermediate carinae on the sixth abdominal somite is sculptured
rather than smooth.
In his redescription of C. depressa (Miers), Seréne (1952, text fig. 15,
pl. 1, fig. 4) figured a specimen from the Australian Museum collection
which he identified as C. latreillei. The specimen may be referable to
either C. bombayensis or C. latispina, for the posterolateral spines on
the lateral processes of the sixth and seventh thoracic somites are clearly
shown in the figures. However, it may also represent a distinct species,
for in text fig. 15 submedian carinae are shown on the sixth, seventh,
and eighth thoracic somites rather than on the eighth only as in C.
latispina, or the seventh and eighth, as in C. bombayensis.
One of the specimens identified as C. microphthalma and figured by
Seréne (1952, text fig. 17) in the same paper may represent still another
distinct species, for it also has posterolateral spines on the lateral
processes of the sixth and seventh thoracic somites and apparently lacks
submedian carinae on the thorax as well.
As in some other species of Clorida, the telsons of adult males and
females have markedly different appearances. To illustrate these dif-
ferences, telson of both sexes, are shown in Figure 3. In addition the
female differs from the male in having numerous straight hairs on the
proximal portion of the propodus of the claw.
In the key to the species of Clorida published by me in 1968, C.
latispina would key out to C. bombayensis.
Etymology: The name refers to the spinose lateral processes of the
sixth and seventh thoracic somites.
250 Proceedings of the Biological Society of Washington
LITERATURE CITED
Cuuapcar, B. F., AND S. R. SANE. 1967. Two new species of Squilla
(Stomatopoda) from Bombay. Crustaceana, 12 (1): 1-8,
figs. 1-2.
Kemp, S. 1913. An account of the Crustacea Stomatopoda of the
Indo-Pacific region, based on the collection in the Indian
Museum. Mem. Indian Mus., 4: 1-217, 10 text figs., pls.
1-10. f
LANCHESTER, W. F. 1901. On the Crustacea collected during the
“Skeat Expedition” to the Malay peninsula, together with a
note on the genus Actaeopsis. Part I. Proc. Zool. Soc. Lon-
don, 1901: 534-574, pls. 33-34.
MANNING, RayMonpD B. 1966. Notes on some Australian and New
Zealand stomatopod Crustacea, with an account of the species
collected by the Fisheries Investigation Ship Endeavour.
Rec. Australian Mus., 27 (4): 79-137, figs. 1-10.
1968. Stomatopod Crustacea from Madagascar. Proc. U.S.
Nat. Mus., 124 (3641): 1-61, figs. 1-16.
SERENE, R. 1952. Etude d’une collection de stomatopodes de l’Aus-
tralian Museum de Sydney. Rec. Australian Mus., 23 (1):.
1-24, figs. 1-33, pls. 1-3.
OG TS
Vol. 81, pp. 251-256 30 August 1968
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
ISOODON DESMAREST, 1817, RATHER THAN
THYLACIS ILLIGER, 1811, AS THE VALID GENERIC
NAME OF THE SHORT-NOSED BANDICOOTS
(MARSUPIALIA: PERAMELIDAE )
By Wituiiam Z. LIDIcKER, Jr., AND W. I. FOLLETT
Museum of Vertebrate Zoology, University of
California, Berkeley, and California Academy
of Sciences, San Francisco
In recent years, two generic names have been widely used
for the three or more species of short-nosed bandicoots, which
inhabit large parts of Australia and southeastern New Guinea.
The older name, Thylacis Illiger, 1811, has been used, for
example, by Simpson (1945), Laurie and Hill (1954), Marlow
(1958), and Ziegler and Lidicker (1968); the later name,
Isoodon Desmarest, 1817, by Troughton (1947), Tate (1948),
Mackerras and Mackerras (1960), and Marlow (1962).
Since differing interpretations of the historical record con-
tinue to prolong nomenclatural instability in this group of
marsupials, we have taken a fresh look at the entire problem,
and have examined essentially everything written on these
animals up to 1888. It was in this year that Oldfield Thomas
formally designated Perameles nasuta Geoffroy, 1804, as the
type-species of Perameles, and Didelphis obesula Shaw and
Nodder, 1797, as the type-species of Thylacis, thus pre-
cipitating the current confusion. As a result of our research,
we have become convinced that Isoodon Desmarest, 1817,
rather than Thylacis Illiger, 1811, is the valid generic name
of the short-nosed bandicoots. To record the evidence sup-
porting this conclusion, we must review briefly the major
events in the nomenclatural history of the three nominal
genera concerned.
29—Proc. Bio. Soc. WasH., VoL. 81, 1968 (251)
252 Proceedings of the Biological Society of Washington
Discussion: The first bandicoot known to science was
described as Didelphis obesula by Shaw and Nodder (1797) .1
The nominal genus Perameles was erected by Geoffroy
(1803), who included therein two recognizable species, but
did not mention either by a specific name. In these circum-
stances, Perameles Geoffroy, 1803, is to be regarded as a
nominal genus without originally included nominal species
(see Art. 69 a i).?
Geoffroy (1804) redescribed the genus Perameles, including
therein Didelphis obesula Shaw and a new species described
by Geoffroy, in the same paper, as Perameles nasuta (These
were the same two species included, but not by name, in
Perameles Geoffroy, 1803). These two nominal species are to
be regarded as those first subsequently referred to the nominal
genus Perameles Geoffroy, 1803 (see Art. 69 a ii). Geoffroy
(1804) did not designate or indicate the type-species of the
nominal genus Perameles. Therefore, any zoologist could sub-—
sequently designate as the type-species either one of the two
nominal species first subsequently referred to this genus (see
Art. 69 a). The earliest subsequent designation that we have
found was that of Lesson (1828:197), who stated unequiv-
ocally “. . . Perameles, avait pour type le Perameles nasuta,
Geoff., . . .” This type-designation fixed the generic name
Perameles upon the long-nosed bandicoots, a conclusion af-
firmed by subsequent authors, both contemporary with Lesson
(e.g. Desmerest, 1849:577) and modern (e.g. Iredale and
Troughton, 1934:18).
The nominal genus Thylacis was erected by Illiger, 1811,
who cited as generic synonyms “Perameles Geoff. Tiedem.”
[= Perameles Geoffroy, 1803, and Perameles Tiedemann,
1808], and who included the same two species, Didelphys
[sic] obesula Shaw and Perameles nasuta Geoffroy.
Demonstrably, Illiger proposed the generic name Thylacis
as a replacement name for Perameles Geoffroy, 1803, and
Perameles Tiedemann, 1808, which he regarded as violating
1 Date from Sherbom (1895:376), and also independently derived from a study of
the several dated plates in the work.
2The abbreviation “Art.” is used in this paper to designate an Article of The
International Code of Zoological Nomenclature (1964).
On a Bandicoot Genus 953
the rules of the Linnaean botanical philosophy. Illiger (1811:
xvi) stated, “Nomina generica, quae rejeci, secundum regulas
codicis nostri, philosophiae nempe botanicae Linnaeanae,
quibus contradicunt, digesta jam enumerabo.” We translate
this, “I shall now enumerate separately those generic names
that I have rejected, [which] according to the rules of our
code, obviously violate those of the Linnaean botanical
philosophy.” This statement is followed by an enumeration
of 15 categories of rejected names. One such category (page
xvii) reads as follows: “N[omen] g[enericum] cui syllaba
una vel altera praeponitur (aut aufertur) ut aliud genus, quam
antea, significat, excludendum est.” We translate this, “A
generic name to which one syllable or another is prefixed (or
[from which it is] removed) so that it signifies another genus
than [it did] formerly, is to be excluded.” In this category
Illiger listed Perameles (which he apparently considered
objectionable because it represented the generic name Meles
with syllables prefixed )—expressly demonstrating his purpose
of substituting Thylacis as a replacement name. Further evi-
dence for his intent is provided by the fact that Illiger’s
description of Thylacis matches in all significant details Geof-
froy’s original description of Perameles.
If a zoologist proposes a new generic name expressly as a
replacement for a prior name, both nominal genera must have
the same type-species, and (subject to the requirement that
the type-species must be a species eligible for fixation as the
type of the earlier nominal genus), type-fixation for either
applies also to the other, despite any statement to the contrary
(see Art. 67 i). Thus when Lesson (1828:197) designated
Perameles nasuta Geoffroy as the type-species of the nominal
genus Perameles Geoffroy, 1803, that type-fixation applied
also to the nominal genus Thylacis Illiger, 1811, which had
been proposed expressly as a replacement for Perameles
Geoffroy, 1803.3
The type-species of Thylacis Illiger, 1811, having once been
8 This applies also to Oken’s (1816) invalid name Thylax, which was proposed as
still another alternative for Perameles. Oken (1816) was rejected for nomenclatural
purposes by Opinion 417 of the International Commission on Zoological Nomen-
clature.
254 Proceedings of the Biological Society of Washington
fixed by Lesson (1828:197) in conformity with the provisions
of the Code, was not subject to change (see Art. 61). The
action of Oldfield Thomas (1888:227) in designating “P.
obesula” as type of Thylacis Mliger, 1811, was therefore void.
Thylacis Iliger, 1811, remains a junior objective synonym of
Perameles Geoffroy, 1803 (see Art. 61 b). This was the con-
clusion reached by Mackerras and Mackerras (1960), who,
however, did not state their supporting evidence or reasoning,
and therefore left the matter open to question.
Didelphis obesula Shaw and Nodder, 1797, is currently
regarded as generically distinct both from Didelphis Linnaeus,
1758, and from Perameles Geoffroy, 1803. The oldest generic
name with which it may properly be combined is Isoodon
Desmarest, 1817.
Thus the three nominal genera concerned are the following:
Perameles Geoffroy, 1803
Without originally included nominal species. (Art. 69 a i)
First subsequently referred species (through Geoffroy,
1804): Didelphis obesula Shaw and Nodder, 1797, and
Perameles nasuta Geoffroy, 1804. (Art. 69 a ii)
Type-species, by subsequent designation of Lesson (1828:
197): Perameles nasuta Geoffroy, 1804. (Art. 69 a)
Thylacis Mliger, 1811
Proposed expressly as a replacement name for Perameles
Geoffroy, 1803.
Originally included species: Didelphys [sic] obesula Shaw
and Nodder, 1797, and Perameles nasuta Geoffroy, 1804.
Type-species, through designation by Lesson (1828:197) of
the type-species of Perameles Geoffroy, 1803: Perameles
nasuta Geoffroy, 1804. (Art. 67 i)
Isoodon Desmarest, 1817
Only originally included species, and therefore type-species
by monotypy: Didelphis obesula Shaw and Nodder, 1797.
(Art. 68 c)
Acknowledgments: We are indebted to Lillian J. Dempster,
of the California Academy of Sciences, and to Basil Marlow,
of the Australian Museum, for assistance with the literature,
On a Bandicoot Genus 255
and to Alan C. Ziegler, now with the Bishop Museum, Hono-
lulu, for technical assistance with this project.
LITERATURE CITED
Desmanrsst, A.G. 1817. In Nouveau Dictionnaire d’Histoire Naturelle
. (nouv. ed.). Paris, 595 pp., vol. 16; 542 pp., vol. 18;
610 pp., vol. 25.
. 1849. In Dictionnaire Universel d’Histoire Naturelle, vol. 9.
Paris, 776 pp.
Greorrroy, E. 1803. Note sur les genres Phascolomys et Perameles,
nouveaux genres d’animaux a bourse. Bull. des Sciences, par
la Soc. Philomathique 3 (80): 249-250.
1804. Sur un nouveau genre de mammiféres a bourse,
nommé Péraméles. Ann. du Mus. National d’Hist. Nat. 4
(12): 56-65.
InuicEr, J. K. W. [llligeri D., Caroli] 1811. Prodromus Systematis
Mammalium et Avium. ... Berlin, 301 pp.
INTERNATIONAL CODE OF ZOOLOGICAL NOMENCLATURE ADOPTED BY THE
XV INTERNATIONAL ConcREss oF ZooLocy. 1964. London,
176 pp.
IREDALE, T., AND E. LEG. Troucuton. 1934. A check-list of the
mammals recorded from Australia. Aust. Mus. Mem., 6: 1-
122.
Lauriz, E. M. O., ano J. E. Hm. 1954. List of Land Mammals of
New Guinea, Celebes and Adjacent Islands 1758-1952.
London, 175 pp.
Lesson, R. P. 1828. In Dictionnaire Classique d’Histoire Naturelle,
vol. 13. Paris, 648 pp.
Mackerras, I. M., AND M. J. MAcKERRAS. 1960. Taxonomy of the
common short-nosed marsupial bandicoot of eastern Queens-
land. Aust. Jour. Sci., 23 (2): 51-53.
Martow, B. J. 1958. A survey of the marsupials of New South Wales.
C.S.1.R. O. Wildlife Res., 3 (2): 71-114.
1962. Marsupials of Australia. Brisbane, 140 pp.
Oxen, L. 1816. Lehrbuch der Naturgeschichte, 3, Zoologie (2). Jena,
1272 pp.
Suaw, G., AND F. P. Nopper. 1797. The Naturalist’s Miscellany: or
coloured figures of natural objects; drawn and described
immediately from Nature, vol. 8. London, 180 pp. + 46
pls., 1796-97.
SHERBORN, C. D. 1895. On the dates of Shaw and Nodder’s ‘Natural-
ist’s Miscellany.’ Ann. Mag. Nat. Hist. (ser. 6), 15: 375-376.
Simpson, G. G. 1945. The principles of classification and a classifica-
tion of mammals. Bull. Amer. Mus. Nat. Hist., 85: 1-350.
Tate, G. H. H. 1948. Results of the Archbold expeditions. No. 60.
Studies in the Peramelidae (Marsupialia). Bull. Amer. Mus.
Nat. Hist., 92 (6): 315-346.
256 Proceedings of the Biological Society of Washington
Tuomas, O. 1888. Catalogue of the Marsupialia and Monotremata in
the Collection of the British Museum (Natural History).
London, 401 pp.
TIEDEMANN, D. F. 1808. Zoologie. Zu seinen Vorlesungen entworfen.
Erster Band. Allgemeine Zoologie, Mensch und Saugthiere.
Landshut, 612 pp.
Troucuton, E. 1947. Furred Animals of Australia. New York, 374
pp. 5
ZIEGLER, A. C., AND W. Z. LivickER, Jr. 1968. Keys to the genera of
New Guinea Recent land mammals. Proc. Calif. Acad.
Sciences, in press.
x
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Vol. 81, pp. 257-260 30 August 1968
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
THE STATUS OF THE NAME MYOTIS
. SUBULATUS SAY?
By Bryan P. Giass AND RoBert J. BAKER
Zoology Department, Oklahoma State University
The name subulatus was first applied to a North American
Myotis by footnotes in James (1823) account of Long's
expedition from Pittsburgh to the Rocky Mountains. Ap-
parently James took the description verbatim from Say’s field
notes, and the description has quite properly been attributed
to Say. The type locality is about 40 miles above the Pur-
gatoire, on the banks of the Arkansas River, slightly east of
the 104th meridian. Say’s notes suggest that he collected at
a place where a flowing creek enters the river. The place
most nearly fitting this description lies somewhere between
the towns of Fowler and Manzanola in Otero County and
Olney Springs in Crowley County, Colorado. This is a more
precise definition than the one used by Miller and Kellogg
(1955), where the locality is given as “Arkansas River, near
La Junta, Otero County, Colorado.” Actually La Junta is some
25 miles east of this locality. James recorded that the party
had passed the Wharf (= Huerfano) during the morning of
July 20, and had ridden 26 miles along the river during the
day. The next morning they traveled only approximately
10 miles, where they camped at a ford on the north bank,
remaining there until the morning of the 24th. This places
the type locality of Myotis subulatus in the vicinity of the
mouth of Apishapa Creek, which enters the Arkansas from
the south.
The type specimen is not mentioned in Say’s notes as having
1 Contribution number 450, Department of Zoology, Oklahoma State University,
Stillwater, Oklahoma 74074.
30—Proc. Biot. Soc. WaAsuH., Vou. 81, 1968 (257)
258 Proceedings of the Biological Society of Washington
been preserved, but in another footnote (James, 1823:14) it
was indicated that the type of the band-tailed pigeon and
other natural history specimens acquired on this expedition
were placed in the Philadelphia Museum. This collection,
popularly known as Peale’s Museum, was later destroyed by
fire and the type of M. suwbulatus with it, if such was ever
actually preserved.
LeConte (1855) applied the name Vespertilo subulatus Say
to bats in the tidewater country near Riceboro, Liberty
County, Georgia. Miller and G. M. Allen (1928) have in-
dicated that LeConte presumed that he had two species, to
one of which he applied the name V. subulatus Say, but they
suspected that all LeConte’s specimens were actually Myotis
lucifugus. Whatever the species actually was, it certainly was
not the saxicolous species currently bearing the name M.
subulatus, which is absent from the southeastern United
States.
Harrison Allen (1864) applied Say’s name to the eastern
form of the long-eared Myotis, which usage was accepted
until the revision of the genus by Miller and G. M. Allen
(loc. cit.) wherein they correctly rejected M. subulatus for
the eastern long-eared Myotis in favor of the name M. keeni
Merriam 1895, which is currently accepted, and allocated M.
subulatus instead to the form currently bearing the name.
They based this change (op. cit. p. 25) in part, on their
imperfect knowledge of the bats known to occur in south-
eastern Colorado.
To date only two species of Myotis are known from the
plains where Say made his collection. These are the species
currently known as M. subulatus, and M. yumanensis. Both
are known from Cimarron County, Oklahoma and the former
also by one specimen from Baca County, Colorado about 12
miles north of the Oklahoma border.
Upon reading Say’s description, one who is familiar with
both species is impressed by the fact that the characters fit
M. yumanensis as well as, or better than, they do M. subulatus
(sensu Miller and Allen, 1928 et. auct.), particularly in its
being cinereous rather than chestnut, having a normal-sized
rather than dwarfed foot, with toes beset with hair, and in
Status of Myotis subulatus 259
its manner of flight. Say described this bat as flying—‘“rapidly
in various directions, over the surface of the creek”—. In the
writers’ experience M. yumanensis flies close to the surface of
streams, but follows a rather steady path directly above the
water. M. subulatus seems not to be a compulsive stream-
flier, and is often high enough to be seen against the sky.
Because of the questionable identity of Say’s specimen one
of us (Glass and party) attempted to collect bats near the
mouth of Apishapa Creek May 15, 1967. Several bats were
seen coursing the water in typical “yuwmanensis” fashion, and
two taken in mist nets proved to be M. yumanensis. However,
one or two bats were also seen flying some 10-12 feet above
the ground, and these could well have been M. subulatus.
Unfortunately none were collected. It seems to the writers
that the species now called M. yumanensis is as likely to have
been the species Say collected as is the bat presently called
M. subulatus.
In view of the involved history of use of the name Myotis
subulatus, its present dubious assignment, and the threat that
it poses to the well-established name M. yumanensis, the
writers feel that the name should be considered nonassign-
able. The species currently bearing the name should therefore
be called Myotis leibii, published as Vespertilio leibii Audubon
and Bachman (1842). Relegation of the name subulatus to
synonymy requires that the subspecies of this taxon be as
follows:
Myotis leibii leibii Audubon & Bachman 1842. Type locality
Erie County, Ohio.
Myotis leibii ciliolabrum Merriam 1886. Type locality near
Banner, Trego County, Kansas.
Myotis leibii melanorhinus Merriam 1890. Type locality
Little Spring, North base of San Francisco Mountain,
Coconino County, Arizona, altitude 8,250 feet.
The writers earlier proposed to the International Commis-
sion (Glass and Baker, 1965) that this name be suppressed
and that M. yumanensis be placed on the official list. How-
ever, the proposal was withdrawn after numerous expressions
of disapproval, primarily because the action amounts to resolv-
260 Proceedings of the Biological Society of Washington
ing a taxonomic problem by legalistic nomenclatural proce-
dure. Secondly, the proposal as made would protect M.
yumanensis from justifiable nomenclatural challenge in the
future.
LITERATURE CITED
ALLEN, H. 1864. Monograph of the bats of North America. Smith.
Misc. Coll., (165): 1-85.
AupbuBON, J. J., AND J. BACHMAN. 1842. Descriptions of new species
of quadrupeds inhabiting North America. Jour. Acad. Nat.
Sci. Phila., Sr. 1, 8: 284.
Guass, B. P., AND R. J. BAKER. 1965. Vespertilio subulatus Say, 1823:
Proposed Suppression. Bull. Zool. Nomen., 22 (3): 204-205.
James, E. 1823. Account of an expedition from Pittsburgh to the
Rocky Mountains, Volume 2. H. C. Carey and I. Lea,
Philadelphia.
LeConte, J. 1855. Observations on the North American species of
bats. Proc. Acad. Nat. Sci. Phila., 431-438.
MeErRIAM, C. H. 1886. Description of a new species of bat. from the
western United States (Vespertilio ciliolabrum sp. nov.).
Proc. Biol. Soc.. Washington, 4: 1-4, December 17.
1890. Results of a biological survey of the San Francisco
Mountain region and desert of the Little Colorado, Arizona.
N. Amer. Fauna, 3: 46.
MILLER, G. S., AND G. M. ALLEN. 1928. North American bats of the
general Myotis and Pizonyx. Bull. U.S. Nat. Mus., 144:
218 pp.
, AND R. Kettocc. 1955. List of North American recent
mammals. Bull. U.S. Nat. Mus., 205: 954 pp.
OC TS
Vol. 81, pp. 261-274 30 August 1968
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
TWO NEW CRAYFISHES OF THE GENUS, CAMBARUS
FROM GEORGIA, KENTUCKY, AND TENNESSEE
(DECAPODA, ASTACIDAE )
By Horton H. Hosss, Jr.
Smithsonian Institution, Washington, D.C.
It seems likely that no more than two-thirds of the existing
species of the genus Cambarus have been described. Because
the ranges and limits of variation of so few of the 58 recorded
species and subspecies are understood, most systematists have
hesitated to add further confusion to the already chaotic state
of our knowledge of the genus. Unlike the situation which
obtains in the genus Procambarus, the secondary sexual fea-
tures of Cambarus, particularly the first pleopod of the male,
exhibit, by comparison, few conspicuous modifications from
basic patterns. Additional difficulties derive from the prob-
able fact that adaptations to different types of habitats ex-
ploited by Cambarus have been markedly similar in several
evolutionary lines. As a result, the recognition of species
must be based more on different combinations of these
adaptive features than on characters that are unique. Thus,
in the absence of adequate series of those species which have
already been described, little progress could have been made
toward gaining an understanding of the composition of the
genus.
Compounding the innate taxonomic difficulties was the
unfortunate practice among several workers of designating
new “forms, as well as some old ones, as subspecies of
previously named species—just because they resembled one
another. This has resulted in bringing together totally separate
lineages into a single species complex. While such procedures
have probably been utilized in many different groups of
31—Proc. Bion. Soc. WasH., Vou. 81, 1968 (261 )
262 Proceedings of the Biological Society of Washington
organisms, here they have resulted not only in clouding species
concepts but also in retarding our understanding of relation-
ships of the members of the genus.
With the acquisition of several large collections of Cambarus
by the Smithsonian, representatives of all the known members
of the genus (many of them in large series) have been made
available so that those specimens which have borne the label
“Cambarus sp?” now can be analyzed with some hope of
determining whether or not they can be assigned to previously
described taxa.
In view of the fact that so many of the so-called subspecies
that have been recognized in the genus are based on super-
ficial considerations, it seems reasonable to propose that,
except in those taxa for which there is conclusive evidence
of integradation between populations, subspecific designa-
tions be elevated to the specific level. Surely no additional
subspecific categories should be proposed for this genus, -
which are based solely on superficial resemblances.
Specimens of one of the two species described here was
first collected in 1938, and, in their report on the crayfishes
of the Big South Fork of the Cumberland River, Hobbs and
Shoup (1942) referred to it as “Cambarus bartonii bartonii,
Regionally Restricted Variant 5.” At that time, we knew that
it was related to Ortmann’s “Cambarus bartonii_ striatus”
(1931), but we were unable to determine how close those
relationships actually are. Consequently, we adopted what
we considered to be a conservative view in reporting it as a
variant of the typical subspecies. In retrospect, however, it
is clear that we lent countenance to and compounded an
error of judgment that had already been made. At best, the
affinities of bartonii with striatus and our variant are remote,
and most assuredly they are not members of the same species!
Furthermore, there is every reason to believe that our variant,
although allied to C. striatus Hay (1902b:437), is not a race
of that species.
Cambarus sphenoides new species
Cambarus bartonii bartonii “Regionally Restricted Variant 5,” Hobbs and
Shoup, 1942: 640-641, Plate 1, figs 5 and 8, and Fig. 1.
Cambarus “unidentified species,’ Hobbs, 1965: 271 (in part).
Two New Crayfish 263
Diagnosis: Body pigmented, eyes small but well developed. Rostrum
with convergent margins and adults devoid of marginal spines or tuber-
cles at base of acumen. Areola 4.9 to 11.6 times longer than broad,
comprising 32.8 to 37.0 per cent of entire length of carapace and bear-
ing two to four punctations across narrowest part. Lateral surface of
carapace with or without small cervical spine with adjacent tubercles
or with spine reduced to tubercle. Suborbital angle acute or rounded.
Postorbital ridges with or without small cephalic tubercles or spines.
Antennal scale between 2.3 and 2.5 times longer than broad. Chela with
two rows of tubercles along mesiodorsal portion of palm; lateral margin
of hand costate, and both fingers with well defined longitudinal ridges
on upper surfaces. First pleopod (Figs. 2, 6) of first form male with
central projection recurved at angle much greater than 90 degrees,
tapering only slightly and with distinct subterminal notch; mesial process
inflated and terminating in two to four small subangular prominences.
Annulus ventralis (Fig. 10) asymmetrical with caudosinistral wall
strongly convex caudolaterally; caudal portion slightly movable and
bearing tilted, reverse S-shaped sinus.
Holotypic Male, Form I: Body subcylindrical, not strongly depressed.
Abdomen narrower than thorax (17.3 and 19.6 mm). Greatest width
of carapace greater than depth at caudodorsal margin of cervical groove
(19.6 and 16.5 mm). Areola 7.2 times longer than wide with two
longitudinal rows of punctations. Cephalic section of carapace 1.7 times
as long as areola (length 36.4 per cent of entire length of carapace).
Rostrum excavate dorsally with convergent, slightly thickened margins
devoid of marginal spines or tubercles. Acumen not distinctly set off
from basal portion of rostrum and terminating in small tubercle; upper
surface with usual submarginal punctations and others scattered between.
Subrostral ridges moderately prominent and visible in dorsal aspect to
base of acumen. Postorbital ridges moderately prominent, grooved
dorsolaterally, and rounded cephalically, devoid of tubercles or spines.
Sinistral suborbital angle broadly rounded, dextral one obtuse. Bran-
chiostegal spine small and acute. Carapace punctate dorsally and
tuberculate laterally with row of tubercles along ventral margin of
cephalic portion of cervical groove; cervical spines reduced to small
tubercles. Abdomen slightly shorter than carapace (36.2 and 37.6 mm).
Cephalic section of telson with two spines in each caudolateral corner.
Dorsal surface of telson and uropods not conspicuously setose. Epistome
(Fig. 7) distinctly broader than long with small subacute cephalomedian
projection, lateral portions elevated, central portion slightly convex ven-
trally and bearing setae. Antennules of usual form with small spine on
ventral surface of basal segment at base of distal third. Antenna extend-
ing caudad to fourth abdominal tergum. Antennal scale (Fig. 8) 2.3
times longer than broad, broadest distal to midlength with widest
lamellar area about 2.5 times width of thickened lateral portion, latter
terminating in small acute spine (dextral one broken).
Left chela (Fig. 9) (right regenerated) depressed but with palm
264 Proceedings of the Biological Society of Washington
Fics. 1-11. Cambarus sphenoides new species (pubescence removed
from all structures illustrated). 1, Lateral view of carapace of paratypic
male, form II. 2, Mesial view of first pleopod of holotype. 3, Mesial
view of first pleopod of morphotype. 4, Basis and ischium of third
pereiopod of holotype. 5, Lateral view of first pleopod of morphotype.
6, Lateral view of first pleopod of holotype. 7, Epistome of holotype.
8, Antennal scale of paratypic male, form II. 9, Dorsal view of distal
podomeres of cheliped of holotype. 10, Annulus ventralis of allotype.
11, Dorsal view of carapace of holotype.
Two New Crayfish 265
somewhat inflated; distal three-fourths of lateral margin of hand costate.
Lateral portions of dorsal and ventral surfaces of palm punctate, mesio-
dorsal portion tuberculate with mesial row of eight tubercles and more
lateral one of six; third and fifth members of mesial row slightly ventral
to level of remaining six. Mesial portion of ventral surface tuberculate,
more distal tubercles larger. Fingers not conspicuously gaping and both
with well defined longitudinal ridges dorsally and ventrally. Opposable
margin of fixed finger with five tubercles, third from base largest; both
fingers with narrow band of crowded minute denticles along distal two-
fifths, interrupted on fixed finger by distal tubercle and on dactyl by
distal two; corresponding margin of dactyl with eight tubercles, fourth
from base largest; mesial surface of dactyl with row of six tubercles
decreasing in size distally, row flanked proximally by other tubercles
and distally by linear series of punctations.
Carpus of left cheliped longer than broad with arc-like oblique
longitudinal furrow dorsally and conspicuously punctate dorsally and
laterally; mesial surface with large spine having a smaller one at its
base, and another proximal to latter; dorsomesial surface with several
tubercles. Lower distal margin with large tubercle on lateral articular
knob and another mesial to it; three smaller tubercles present between
latter and large spine on mesial surface.
Merus of left cheliped with mesial and lateral surfaces punctate.
Upper surface tuberculate with four somewhat prominent ones grouped
subdistally. Ventrolateral margin with row of five tubercles and ventro-
mesial margin with 11. Row of three tubercles on ischium correspond-
ing to mesial row on merus.
Hooks on ischia of third pereiopods only (Fig. 4); hooks simple and
not opposed by tubercle on basis but extending proximad of distal margin
of latter. Coxa of fourth pereiopod with caudomesial protuberance; fifth
without prominence.
Sternum moderately deep between third, fourth, and fifth pereiopods,
and with tuft of plumose setae between bases of third and fourth.
First pleopods (Figs. 2, 6) symmetrical and reaching coxa of third
pereiopods when abdomen is flexed. (See diagnosis for description. )
Morphotypic Male, Form II: Differs from holotype in following re-
spects: rostrum more acuminate and subrostral ridges not evident so
far cephalically; areola 8.4 times longer than broad with few scattered
punctations between two lateral rows; inner margin of palm of chela
with six tubercles, and fifth tubercle from base largest on opposable
surface of dactyl. As usual, hooks on ischiopodites of third pereiopod
and protuberance on coxa of fourth reduced. First pleopod (Figs. 3, 5)
with neither element corneous, and central projection heavier and lack-
ing subterminal notch. (See measurements, Table 1.)
Allotype Female: Differs from holotype in following respects: sub-
rostral ridges less conspicuous but slightly evident dorsally to base of
acumen; cervical tubercles smaller; areola constituting only 34.6 per
cent of entire length of carapace and 5.5 times longer than wide, bear-
266 Proceedings of the Biological Society of Washington
ing three rows of punctations; mesial area of palm of left chela with
inner row of six tubercles and more lateral one of five; merus with six
tubercles in ventrolateral row. Hooks on third pereiopods and _pro-
tuberances on coxae of fourth lacking. (See measurements, Table 1.)
Sternum between fourth pereiopods V-shaped and moderately deep.
Annulus ventralis (Fig. 10) about 1.5 times longer than broad, firmly
fused to stemum cephalically but caudal half movable; caudal wall
asymmetrical with sinistral one inflated and convex caudoventrally,
dextral one concave; cephalomesial area with distinct longitudinal median
furrow, sinus originating sinistrally at end of furrow, curving gently
caudomesially and making U-tum cephalomesially before crossing median
line; second broad U-turn crossing median line and ending on caudal
wall of annulus almost on median line.
Type-Locality: Tributary to Clear Creek (Emory River drainage), 11.2
miles north of Crossville, Cumberland County, Tennessee, on U.S. Rte.
127. There the stream is approximately 10 feet wide, and the slightly
turbid water flows with a moderate current over a rock and gravel
bottom. The stream is partially shaded by a mixed stand of Quercus
and Pinus.
Disposition of Types: The holotypic male, form I, the allotypic female, _
and the morphotypic male, form II, are deposited in the United States
National Museum (nos. 129325, 129326, 129327, respectively) as are
the paratypes which consist of 2 ¢, form I, 10 ¢, form II, 27 9, 27
juvenile ¢, 32 juvenile 9, and 1 @ with eggs.
Size: The largest male has a carapace length of 38.4 mm, the largest
female, 42.0 mm, and the smallest first form male, 31.3 mm.
Range and Specimens Examined: Although the range of this species
is poorly known, it is widespread in the Cumberland River drainage on
the Plateau and Eastern Highland Rim and is present, but apparently
less abundantly so, in the Sequatchie River system.
KENTUCKY—Jackson County, 0.1 mi. S of Owsley Co. line on
St. Rte. 30 (4 ¢ II, 3 9, 8 juv. ¢). Lauret County, 1.3 mi. SW of
Greenmount on St. Rte. 30 (1 ¢ II, 3 9, 4 juv. ¢). TENNESSEE—
BLEDSOE County, 11.3 mi. SE of Spencer on St. Rte. 30 (1 4 II, 2 9,
7 juv. 6,4 juv. 9). CumMBERLAND County, type-locality (1 46 I, 2 @?);
6.5 mi. E of White Co. line on U.S. Rte. 70 (2 ¢ II, 1 9, 1 juv. ¢);
Caney Fork River at U.S. Rte. 70 (2 6 II, 2 2,1 juv. ¢ ); Scotts Creek,
7.0 mi. N of Crossville on U.S. Rte. 127 (2 @ ); 3.9 mi. E of White Co.
line on U.S. Rte. 70 (5 2, 1 juv. 6); Clear Creek at Jones Ford (1 ¢
II); No Business Creek S of Clark Range (2 ¢ I, 4 6 II, 7 9, 2 juv.
6, 2 juv. @ ); six localities on Clear Creek (7 6 II, 12 9, 1 juv. 2);
four localities on No Business Creek (to Clear Creek) (1 9, 5 juv. 6,
4 juv. 2); Grassy Cove (1 6 II, 1 9, 2 juv. 2); White Creek, SE of
Grassy Cove (1 ¢ Il, 1 juv. 6); Daddys Creek near Crossville (1 ¢ II,
1 92, 1 juv. ¢); Creek on Draw Road between U.S. Rtes. 127 and 70
(1 2); Fentress Co. line on U.S. Rte. 127 (1 6 I, 1 6 II); 14.1 mi.
N of Cumberland Courthouse on U.S. Rte. 127 (1 ¢ II, 1 juv. 2);
ee ee ee ee
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Two New Crayfish 267
TABLE 1. Measurements (mm) of Cambarus sphenoides.
Holotype Allotype Morphotype
Carapace:
Height 16.5 16.1 16.0
Width 19.6 19.3 13.0
Length 37.6 - 36.7 30.3
Areola:
Width 1.9 8} 1.3
Length 13.7 G7 10.9
Rostrum:
Width 6.0 5.9 6.6
Length 8.7 Cos 5.0
Left Chela:
Length of inner margin of palm 9.5 8.5 6.4
Width of palm 14.4 13.0 10.0
Length of outer margin of hand 29.6 26.5 20.2
Length of dactyl 19.2 Ip 13.4
4.1 mi. W of Crossville on U.S. Rte. 70 (3 ¢ II, 1 @). DEKats
County, Sink Creek, Blue Springs Community (2 juv. @). FENTREsS
County, Big Hurricane Creek west of Clark Range (2 ¢ II, 5 9, 10 juv.
6, 13 juv. 2, 1 @ with eggs); 9.9 mi. S of Jamestown on U.S. Rte.
127 (1 6 IL,1 9, 3 juv. 6, 1 juv. 2); Mill Creek east of Clark Range
(1 9, 2 juv. 6, 6 juv. 2); Long Branch, 1.5 mi. E of Grimsley (2 juv.
6, 5 juv. 2); Little Crab Creek near Jamestown (1 9, 2 juv. 9).
Marion County, 2 mi. E of Sewanee on U.S. Rte. 41 (2 ¢ II, 3 9,
1 juv. ¢, 1 juv. 2). Morcan County, Clear Fork River at Gatewood
Ford (1 2). Overton County, East Fork of Obey River at Cliff Springs
(5 juv. 6, 5 juv. 2); 3.3 mi. SE of Hilham on St. Rte. 85 (2 juv. 6,
4 juv. 2). Purnam County, Hurricane Creek W of Clark Range (1 ¢
Il, 4 9, 12 juv. 6, 17 juv. 2); Meadow Creek E of Monterey (1 9,
2 juv. 2); East Fork of Obey River and Meadow Creek (2 9, 1 juv.
@ ). Rua County, Morgan Creek, 4 mi. E of Bledsoe Co. line on St.
Rte. 30 (1 9). VAN BurEN County, Long Branch of Rocky River W of
Spencer (1 92, 1 juv. @ ); 8.4 mi. E of Spencer on St. Rte. 30. WARREN
County, 11.9 mi. SW of McMinnville on St. Rte. 55 (1 juv. 2); 6.1 mi.
SW of McMinnville on St. Rte. 55 (2 juv. ¢); Charles Creek below
County Rd. 4399 (1 ¢ II, 1 9, 1 juv. 6, 1 juv. 2). Wurre County,
1.1 mi. S of Putnam Co. line on State Rte. 42 (2 ¢ Il, 3 @); Clifty
Creek, near Clifty (1 ¢ II).
Variations: The shape of the rostrum ranges from sharply acuminate
to one with subangular contractions at the base of the acumen; in the
young of some populations, small, acute marginal tubercles are present.
The areola varies from 32.8 to 37.0 per cent of the entire length of the
268 Proceedings of the Biological Society of Washington
carapace and from 4.9 to 11.6 times longer than broad; few individuals,
however, have an areola more than 8.0 times longer than broad. Cervical
tubercles are always comparatively small, but occasional individuals
exhibit acute ones. The suborbital angle is almost always obtuse if
present, but in some juveniles it is produced into a small spine. Differ-
ences also occur in the numbers of tubercles on the chelipeds, but,
except in regenerated appendages, the numbers are scarcely more variable
than differences noted in the primary types.
Color Notes: The thorax and abdomen of the holotype were almost
concolorous reddish brown with paler rostral margins, postorbital ridges,
and indistinct bands along the caudolateral margins of the cervical
groove. As in most crayfishes, the branchiostegites are paler ventrally.
The chelae were only slightly lighter than the carapace, and the tubercles
and spines were about the same color as the rostral margins. The remain-
ing pereiopods were pale tan proximally and ventrally but otherwise
also dark brown.
Life History Notes: Three of the four first form males listed above
were collected in April, the other in July. A single female carrying eggs
was found on April 27, 1945.
Relationships: Cambarus sphenoides has its closest affinities with Cam- —
barus striatus Hay, from which it differs in possessing a shorter, broader
areola, and the central pivjection of the first pleopod of the first form
male possesses a subterminal notch, in this respect resembling the new
species described below. It differs from the latter, however, in lacking
marginal spines on the rostrum and in possessing a longer, narrower,
and less punctate areola. (Compare Figs. 11 and 20.)
Etymology: The name sphenoides is derived from Greek: spheno—
wedge; oides—like; so named because of the shape of the chela.
Crayfish Associates: Collected with Cambarus sphenoides in one or
more localities were Cambarus friaufi Hobbs (1953:24), Cambarus parv-
oculus Hobbs and Shoup (1947:142), Cambarus tenebrosus Hay
(1902a:232), two undescribed members of the genus Cambarus and one
of the genus Orconectes.
Remarks: In discussing this species, Hobbs and Shoup (1942:640)
indicated that in the Big South Fork it occurs “. . . in the drainage of
the Clear Fork River, the principal western tributary. . . . The stream
is characterized by a low total alkalinity averaging only 10.8 parts per
million from 9 stations where methyl orange tests were made. The water
flows over bed rock and rubble of siliceous nature; there is good shade
and excellent cover under rock ledges and in crevices. Aquatic vegeta-
tion is extremely limited, and there is a paucity of bottom fauna. The
Clear Fork River is considered by us a typical upper Cumberland
Plateau stream which flows over weathered sandstone through an ex-
tremely unproductive agricultural area.”
Although comparable data are not available for localities other than
those in the Clear Fork, C. sphenoides is a stream inhabitant occurring
among rock litter and burrowing into the submerged banks.
Two New Crayfish 269
Cambarus halli new species
Diagnosis: Body pigmented, eyes well developed. Rostrum with
slightly convergent margiis bearing marginal spines or tubercles at base
of acumen. Areola 3.1 to 3.9 times longer than broad, comprising 28.9
to 33 per cent of entire length of carapace, and bearing five to seven
punctations across narrowest part. Lateral surface of carapace with
moderately strong cervical spine and two or three small cervical tubercles.
Suborbital angle obtuse and rounded. Postorbital ridges with cephalic
spines or tubercles. Antennal scale about 2.5 times longer than broad.
Chela with two rows of tubercles along mesiodorsal portion of palm;
lateral margin of hand costate, and both fingers with well defined
longitudinal ridges on upper surfaces. First pleopod (Figs. 13, 17) of
first form male with central projection recurved at angle much greater
than 90 degrees, not markedly tapering distally and with a distinct sub-
terminal notch; mesial process inflated but less bulbous than in most
related species; tip rounded and entire. Annulus ventralis (Fig. 21)
almost symmetrical and with caudal portion slightly movable and _ bear-
ing tilted, reverse S-shaped sinus.
Holotypic male, form I: Body subovate, depressed. Abdomen nar-
rower than thorax (14.0 and 16.3 mm). Greatest width of carapace
greater than depth at caudodorsal margin of cervical groove (16.3 and
13.2 mm). Areola broad (3.1 times longer than wide) with many
punctations, six or seven across narrowest portion. Cephalic section of
carapace twice as long as areola (length of areola 33 per cent of entire
length of carapace). Rostrum excavate dorsally with convergent, slightly
thickened margins bearing pair of tubercles at base of moderately long
acumen, latter terminating in corneous, slightly upturned tip; upper
surface with usual submarginal punctations and with others scattered
over entire area. Subrostral ridges weak and visible in dorsal aspect to
level of posterior third of base of eyestalk. Postorbital ridges moderately
prominent, grooved dorsolaterally, and terminating cephalically in cor-
neous tubercles. Suborbital angle broadly rounded. Branchiostegal
spines moderately strong and acute. Carapace punctate dorsally and
tuberculate laterally with row of tubercles along ventral margin of
cephalic portion of cervical groove; cervical spine strong with several
tubercles above and below it. Abdomen longer than carapace (31.4 and
30.3 mm). Cephalic section of telson with two spines in dextral and
three in sinistral caudolateral corners. Dorsal surfaces of telson and
uropods conspicuously setose.
Epistome (Fig. 18) subequal in length and breadth, with subacute
apex, lateral areas elevated, and central portion slightly convex ventrally.
Antennules of usual form with small spine on ventral surface of basal
segment slightly distal to midlength. Antenna broken in holotype but
extending caudad to end of telson in morphotype. Antennal scale (Fig.
19) 2.6 times longer than broad, broadest at midlength with widest
lamellar area approximately twice width of thickened lateral area, latter
terminating in acute spine.
270 Proceedings of the Biological Society of Washington
Fics. 12-22. Cambarus halli new species (pubescence removed from
all structures illustrated). 12, Lateral view of carapace of paratypic
male, form II. 13, Mesial view of first pleopod of holotype. 14, Mesial
view of first pleopod of morphotype. 15, Basal portions of third, fourth,
and fifth pereiopods of holotype. 16, Lateral view of first pleopod of
morphotype. 17, Lateral view of first pleopod of holotype. 18, Epistome
of holotype. 19, Antennal scale of paratypic male, form II. 20, Dorsal
view of carapace of holotype. 21, Annulus ventralis of allotype. 22,
Dorsal view of distal podomeres of cheliped of holotype.
Two New Crayfish rail
Right chela (Fig. 22) depressed but with palm inflated; distal three-
fourths of lateral margin of hand costate. Ventral surface and lateral
portion of dorsal surface of palm mostly punctate, mesiodorsal portion
tuberculate with two distinct rows of six tubercles each. Ventral surface
with three tubercles—one proximomesial to ventral condyle of dactyl,
another at level of condyle but proximal to base of gap between fingers,
and third on ridge at base of dactyl. Fingers slightly gaping and both
with well-defined longitudinal ridges dorsally and ventrally. Opposable
margin of fixed finger with five tubercles, third from base largest; cor-
responding margin of dactyl with six, fourth from base largest; both
fingers with single row of minute denticles along most of distal half
interrupted on each by distalmost tubercle. Mesial surface of dactyl
with row of tubercles, except for fourth, decreasing in size distally; row
flanked proximomesially by other tubercles, and distally by linear series
of punctations.
Carpus of right cheliped longer than broad, with oblique longitudinal
furrow dorsally and conspicuously punctate dorsally and laterally; mesial
surface with one large spine and smaller acute tubercle proximal to it;
dorsomesial surface with row of three small subsquamous tubercles.
Lower distal margin with large spine on lateral articular knob and large
tubercle mesial to it, smaller tubercle present proximal to latter.
Merus of right cheliped with mesial and lateral surfaces punctate.
Upper surface with three subdistal tubercles, two large and spiniform.
Ventrolateral margin with two tubercles and ventromesial margin with
row of eight (left with three and nine, respectively). Row of four
tubercles on ischium corresponding to mesial row on merus.
Hooks on ischia of third pereiopods only (Fig. 15); hooks simple
and not opposed by tubercle on basis but extending proximad of distal
end of latter. Coxa of fourth pereiopod with prominent caudomesial
protuberance; fifth without prominences.
Sternum moderately deep between third, fourth, and fifth pereiopods
and with moderate tuft of plumose setae between bases of third and
fourth pereiopods.
First pleopods (Figs. 13, 17) symmetrical and reaching coxa of third
pereiopods when abdomen is flexed. (See diagnosis for description. )
Morphotypic Male, Form II: Differs from holotype in following re-
spects: marginal tubercles on rostrum more acute; areola only 31.7 per
cent of entire length of carapace but 3.7 times longer than broad;
cephalic section of telson with two spines in each caudolateral corner.
Hooks on ischiopodites of third pereiopods and protuberance on coxa
of fourth reduced. First pleopod (Figs. 14, 16) with neither element
corneous, both proportionately much heavier and central projection
lacking subapical notch. (See measurements, Table 2. )
Allotypic Female: Differs from holotype in following respects: ros-
trum and postorbital ridges provided with spines instead of tubercles;
areola 30.7 per cent of entire length of carapace and 3.4 times longer
than wide; cephalic section of telson with two spines in each caudo-
272, Proceedings of the Biological Society of Washington
lateral corner; chela with only five tubercles in each row along mesial
and dorsomesial surfaces of palm; major tubercles on opposable margins
of finger of chela much reduced; mesial surface of carpus of cheliped
with moderately conspicuous tubercle at base of major spine; ventral
surface of merus with only six tubercles in mesial row. Hooks on third
pereiopod and protuberance on coxa of fourth lacking. (See measure-
ments, Table 2.)
Sternum between fourth pereiopods broad and shallow. Annulus
ventralis (Fig. 21) twice as broad as long, rather firmly fused with
sternum cephalically, but caudal half slightly movable; cephalomesial
area with distinct slightly oblique furrow extending caudosinistrally to
cephalic end of sinus, latter making broad U-turn dextrally on transverse
ridge across median line then turing gently caudodextrally, and finally
caudally and slightly caudosinistrally to caudal margin of annulus.
Type-locality: Small tributary of the Tallapoosa River, 1.3 miles south
of the River on U.S. Rte. 27, Haralson County, Georgia. There the
creek is some six feet wide and six inches deep, and the slightly cloudy
water flows over a sand bed littered with rocks. The stream is shaded
by several species of trees and shrubs among which are Liquidambar
styraciflua, Acer sp., and Alnus rugosa. No other crayfishes were col-—
lected at this locality.
Disposition of Types: The holotypic male, form I, the allotypic female,
and the morphotypic male, form II, are deposited in the United States
National Museum (nos. 129288, 129289, 129290, respectively) as are
the paratypes, which consist of 3 6, form I; 4 ¢, form II; 4 9; 11
juvenile ¢; and 10 juvenile 2. No additional specimens of this species
are available.
Size: The largest male has a carapace length of 30.6 mm, the largest
female, 33.3 mm; and the smallest first form male, 30.3 mm.
Color Notes: Cephalic portion of carapace dark brown with cream
tubercles laterally. Margins of rostrum and postorbital ridges red.
Branchiostegites tan, areola dark olive brown. Abdomen dark olive with
narrow transverse pinkish yellow band on caudal margin of each tergum;
pleura with cream margins. Chela olive with basal articular tubercle
and those at base of dactyl red; tubercles on mesial surface of palm
orange; fingers dark green basally fading rapidly to cream; tubercles on
mesial surface of dactyl orange. Entire ventral surface of animal greenish
cream.
Range and Crayfish Associates: This species is known from only three
localities in the headwaters of the Tallapoosa River in Georgia—the type
locality; the Tallapoosa River at U.S. Rte. 27 in Haralson County; and
the same river on Georgia Rte. 101 in Paulding County. In the latter
two localities it was sharing the stream with Procambarus spiculifer
(LeConte) and Cambarus latimanus (LeConte), respectively.
Relationships: Cambarus halli seems to combine characters of C.
spicatus Hobbs (1956b:116) and its allies (C. acuminatus Faxon 1884:
113 and C. robustus Girard, 1852:90) with those of Cambarus latimanus
Two New Crayfish 273
TABLE 2. Measurements (mm) of Cambarus halli.
Holotype Allotype Morphotype
Carapace:
Height NS2 11.5 10.5
Width 16.3 13.0 12.6
Length SOS 28.0 25.9
Areola:
Width So D155 2).
Length 10.0 8.6 8.2
Rostrum:
Width el 4,2 4.4
Length 8.3 7.8 Us
Chela:
Length of inner margin of palm 6.7 4.2 4.3
Width of palm 10.7 6.4 6.6
Length of outer margin of hand 21.0 15.1 14.6
Length of dactyl 3.2 9.9 9.1
(LeConte, 1856:402) and its relatives. The cervical spine, marginal
rostral tubercles, and the short, broad, punctate areola of C. halli bestow
upon it an aura of primitiveness that is not so obvious in C. latimanus
and its kin (C. striatus Hay, C. reduncus Hobbs, 1956a:61, and C.
sphenoides). Cambarus halli shares with C. spicatus the primitive fea-
tures indicated but differs from it chiefly in the shorter chela and the
greater curvature of the central projection of the first pleopod.
While the adults of C. latimanus have small cervical spines, the
marginal rostral spines are absent and the areola is much narrower and
less punctate than that of C. halli, C. spicatus, and its relatives. It is
perhaps significant that the young of C. latimanus possess rostral spines
which are lost before the crayfish reach maturity. Also, while the areola
is narrower and has fewer punctations than do C. halli and C. spicatus,
it is much broader than those of C. striatus and C. reduncus. In the
latter two and C. latimanus, the central projection of the first pleopods
tapers from the base and lacks a subterminal notch which is present
in all of the other species mentioned.
Etymology: This species is named in honor of my good friend and
former student, Edward Taylor Hall, Jr., who has assisted me on several
collecting trips and who was with me when this new species was dis-
covered.
Acknowledgments: I wish to express my appreciation to the following
persons who have donated or assisted in the collection of specimens of
the species described above: J. F. Fitzpatrick, Jr., E. E. Ford, E. T.
Hall, Jr., W. Hildreth, P. C. Holt, Virgie F. Holt, J. W. Parsons, Jean
E. Pugh, C. S. Shoup, J. M. Stubbs, S. R. Telford, and W. R. West.
274 Proceedings of the Biological Society of Washington
LITERATURE CITED
Faxon, WALTER. 1884. Descriptions of new species of Cambarus; to
which is added a synonymical list of the known species of
Cambarus and Astacus. Proc. Amer. Acad. Arts and Sci., 20:
107-158.
GrrarD, CHARLES. 1852. A revision of the North American Astaci,
with observations on their habits and geographical distribu-
tion. Proc. Acad. Nat. Sci., Philadelphia, 6: 87-91.
Hay, WiL.L1AM Perry. 1902a. Observations on the crustacean fauna
of the region about Mammoth Cave Kentucky. Proc. U.S.
Nat. Mus., 25 (1285): 223-236, 1 fig.
1902b. Observations on the crustacean fauna of Nickajack
Cave, Tennessee, and vicinity. Proc. U.S. Nat. Mus., 25
(1292): 417-439, 8 figs.
Hosss, Horton H., Jr. 1953. Two new crayfishes from the Highland
Rim in Tennessee (Decapoda, Astacidae). Journ. Tenn.
Acad. Sci., 28 (1): 20-27, 18 figs.
1956a. A new crayfish to the genus Cambarus from North
Carolina and South Carolina (Decapoda, Astacidae). Journ.
Elisha Mitchell Sci. Soc., 72 (1): 61-67, 11 figs.
1956b. A new crayfish of the Extraneus Section of the
genus Cambezus with a key to the species of the section
(Decapoda, Astacidae). Proc. Biol. Soc. Wash., 69: 115-
122, 11 figs.
1965. A new crayfish of the genus Cambarus from Ten-
nessee with an emended definition of the genus (Decapoda,
Astacidae). Proc. Biol. Soc. Wash., 78: 265-273, 12 figs.
Hosgs, Horton H., Jr., AND CHARLES S. SHoup. 1942. On the cray-
fishes collected from the Big South Fork of the Cumberland
River in Tennessee during the summer of 1938. Amer. Mid.
Nat., 28 (3): 634-655, 8 figs.
1947. Two new crayfishes (Decapoda, Astacidae) from the
Obey River drainage in Tennessee. Journ. Tenn. Acad. Sci.,
22 (2): 138-145, 22 figs.
LECONTE, JOHN. 1856. Descriptions of new species of Astacus from
Georgia. Proc. Acad. Nat. Sci., Philadelphia, 7: 400-402.
OrRTMANN, A. E. 1931. Crawfishes of the southem Appalachians and
the Cumberland Plateau. Ann. Carnegie Mus., 20 (2): 61-
160.
aie
Y.O06 7S
Vol. 81, pp. 275-290 30 August 1968
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
BEHAVIORAL, MORPHOLOGICAL AND ECOLOGICAL
EVIDENCE FOR TWO NEW SPECIES OF FIDDLER
CRABS (GENUS UCA) FROM THE GULF COAST
OF THE UNITED STATES!
By MiIcHAEL SALMON AND SAMUEL P. ATSAIDES
Department of Zoology, University of Illinois,
Champaign, Illinois, and University of Maryland,
College Park, Maryland
The fiddler crab, Uca pugnax (Smith) is a common form
found along the Atlantic coast of the United States (Crane,
1943). The southern limit of its range on the Atlantic seaboard
varies from year to year, depending upon local climatic con-
ditions, but has been designated as Crescent Beach (Tashian
and Vernberg, 1958) and Daytona Beach (Salmon, 1967),
Florida. At points farther south on the east and west coast of
Florida, U. pugnax is replaced by U. rapax, a closely related
tropical species.
The reported range of U. pugnax was extended to the Gulf
of Mexico by Rathbun (1918) on the basis of three specimens
(USNM 2259) collected from Grand Isle, Louisiana. She
remarked, however, that these individuals were morpholog-
ically atypical of forms found on the eastern seaboard. Shortly
after the publication of her monograph in 1918, Rathbun
examined a few individuals from Mississippi (USNM 74902)
and Texas (USNM 72132) which she identified as U. pugnax.
Salmon (1967) found morphologically similar forms on the
1 Supported by Grant GB-6771 from the National Science Foundation.
2 This paper is a contribution of the Cape Haze Marine Laboratory. Miss Jocelyn
Crane and L. P. Holthuis kindly examined some of our specimens for identification.
Figures 1 and 5 were drawn by Miss Alice Boatwright. We gratefully acknowledge
the hospitality we received from the staff of the Institute of Marine Science (Texas)
and the Gulf Coast Research Laboratory (Mississippi) during this study. Drs. R.
Cressey and F. Chace kindly criticized the manuscript.
32—Proc. Bion. Soc. WasuH., Vou. 81, 1968 (275)
276 Proceedings of the Biological Society of Washington
D
Fic. 1. Uca longisignalis new species, holotype male. A, dorsal view.
B, front view. C, major cheliped, inner view. D, same, outer view. E,
minor cheliped, inner view. F, same, outer view.
west coast of Florida at Yankeetown and farther north, but he
did not extend his observations beyond the Florida coastline.
Salmon (1967), while completing a study of the display of
Florida fiddler crabs, noted differences between the waving
gestures of populations of fiddler crabs on the east and Gulf
coast of Florida. Further, he was able to record nocturnal
Two New Fiddler Crabs O17
10 x
@ PUGNAX
xX LONGISIGNALIS iy
e VIRENS x x
9 ry
x x
x
x
= 8 xX ‘
~~ 7 ‘ ° @%o, e x e o e
é x ox XxX bts . @ ee ; x
X KX xy X ® Cy Bao e ry e e
6 zs x rae : ° ° 0 :
x x X eo reac hd
oy Y @. eo
FRONT WIDTH
J2 13 14 IS 16 17 18 19 20 2l 22
CARAPACE WIDTH (MM)
Fic. 2. Scatter plot of carapace width and corresponding front width
in Uca longisignalis new species, U. virens new species, and U. pugnax.
sounds from males on the Gulf coast but had been unsuccess-
ful three years previously in attempts to record these signals
from North Carolina populations (Salmon, 1965). He sug-
gested that populations of U. pugnax on either side of the
Florida peninsula, which were geographically isolated, might
be diverging into two species.
The study reported here was carried out in order to examine
both Gulf and Atlantic coast populations of U. pugnax in more
detail so that their similarities and differences in behavior and
morphology could be characterized. Our results indicate that
the Atlantic coast populations are distinct from those found
along the Gulf coast of the United States. Further, we present
evidence which suggests that the Gulf coast populations are
themselves divisible into two distinct species. Uca virens new
species, is found from Texas to Mississippi, and U. longisig-
nalis new species, occurs from Louisiana to the West coast of
Florida.
278 Proceedings of the Biological Society of Washington
26 x
x
25 Xk
24 XPINS
@ PUGNAX 2, ¥
23 X LONGISIGNALIS x® s
e VIRENS argh
DACTYLOPOD LENGTH (MM)
Is 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38
PROPOPOD LENGTH (MM)
Fic. 3. Scatter plot of dactylopod and corresponding propopod lengths
in Uca longisignalis new species, U. virens new species, and U. pugnax.
MATERIALS AND METHODS
Field observations were carried out during early June, 1967 and in
late August, 1966 and 1967. Gulf coast populations were studied and
collected at the following localities: Port Aransas, Texas; Cameron,
Louisiana; Ocean Springs, Mississippi; and Yankeetown, Florida. Atlantic
coast populations were studied at Daytona Beach and St. Augustine,
Florida.
The waving displays of males were filmed with a 16-mm camera and
a 120-mm telephoto lens for later frame-by-frame analyses. The behavior
of species showing relatively slow waving gestures was also quantified by
announcing the beginning and end of the display into a tape recorder,
then transcribing the data to a level recorder for measurement. The
following parameters of waves by lone males were measured: general
form, duration, interwave interval, and number of waves completed in a
series. Air temperatures during these observations ranged from 23°—
Sis? (Cy
Recordings of nocturnal sounds produced by males were made with
a Uher 4000-L tape recorder at speeds of 1% inches per second. Contact
microphones were used for transducers, as described previously (Salmon,
1965). The temporal properties of the sounds were measured from
————————— et
Two New Fiddler Crabs 979
oscillographs made with a Fairchild oscilloscope (701) and Grass Kymo-
graph camera (C-4) at film speeds of 25 mm/sec. The following pa-
rameters of the sounds from males were measured: sound duration,
number of pulses per sound, and intervals between consecutive sounds
produced in a series.
A number of morphological characters were measured in order to
characterize the populations from different localities. These included
the width of the front, width of the carapace as measured at its greatest
dorso-lateral extension, the length of the propodus (from the tip of the
finger to the proximal edge), and the length of the dactyl (from the
tip of the movable finger to the upper point of articulation to the
propodus). Additionally, the number, form, and distribution of the
spoon-tipped hairs on the merus of the second maxilliped were examined
in specimens from each of the several conspecific populations.
Distribution and Ecology of Atlantic and Gulf Coast Populations: We
studied several populations of U. pugnax at Daytona Beach and St.
Augustine, Florida. Their coloration and habitat preferences were similar
to those described earlier by Tashian and Vernberg (1958) and Salmon
(1967) for Florida residents.
Gulf coast populations could be separated into two forms on the basis
of their distribution. Uca virens new species was found at Port Aransas,
Texas, Cameron, La., and Ocean Springs, Miss., while U. longisignalis
new species was found at Cameron, La., Ocean Springs, Miss., and at
Yankeetown, Florida.
Differences in preferred habitat between the two Gulf forms were
slight but could be recognized in the areas where the two forms over-
lapped. Uca virens new species was found in muddy sand while U.
longisignalis new species was more common in mud and, occasionally,
muddy-clay substrates. Both forms were intertidal, and both seemed to
prefer the more open areas than those of dense vegetation. Habitat
preferences with respect to substrate were correlated with differences
in the number and form of the spoon-tipped hairs contained on the
second maxillipeds (see below).
Uca longisignalis new species
Material Examined: A total of 125 specimens collected from Yankee-
town, Florida and Ocean Springs, Mississippi. Holotype male, USNM
121599; paratype male USNM 122204. Type locality: Ocean Springs,
Mississippi.
Etymology: Longisignalis—alluding to the extended duration of the
waving display and sounds produced by the males.
Morphological Description: (Fig. 1) Carapace moderately arched
and widest at antero-lateral margins. Frontal margin evenly convex.
Surface of carapace smooth; H-form depression moderately outlined.
Antero-lateral margins distinct, tuning at obtuse angles and continuing
in convergent lines to point opposite middle of cardiac region. Another
280 Proceedings of the Biological Society of Washington
Fic. 4. Uca longisignalis new species. A, inner view of second maxil-
liped showing merus and more distal segments. B, front view of spoon-
tipped process. C, inner view of tip of left abdominal appendage. Uca
virens new species. D, inner view of second maxilliped showing merus
and more distal segments. E, front view of spoon-tipped process. F,
inner view of tip of left abdominal appendage. Many additional groups
of bristles on maxillipeds and abdominal appendages deleted for purposes
of clarity.
2 Gio meget PST OE aes
Two New Fiddler Crabs 281
short ridge present on oblique margin of carapace, just above last two
pairs of walking legs. Oblique margins strongly convergent, forming
less than right angle when extended. Carapace about 2.3 times wider
than front, but these proportions varying with size (see Fig. 2).
Minor cheliped serrated along distal two-thirds of inner margins of
both fingers, except near flattened spooned tips. Fingers with distinct
gap when closed. Scattered row of hairs extending along serrated margins
of both fingers.
Outer surface of major cheliped smooth, but covered with many small,
flat tubercles. Propodus about 1.46 times as long as dactyl, but propor-
tions varying with size (Fig. 3). Submarginal ridge on palm weakly
developed, lacking distinct tubercles and extending from proximal edge
of propodus to lower margin at level below articulation with dactyl.
Teeth of both fingers small; single enlarged tooth prominent near center
of immovable finger. Tip of propodus blunt and obtuse, containing 4-5
prominent teeth. Inner surface of palm with oblique ridge of small
tubercles, often arranged in irregular double row for short portions near
pollex and extending close to Jateral margin of carpal cavity.
Carpus with oblique ridge on inner surface continuing proximally
along upper margin of carpus almost to articulation with merus. Ridge
containing single continuous row of small tubercles.
Supplementary Specific Characters: (Fig. 4) Spoon-tipped hairs on
second maxilliped totalling 25-35, arranged in 4 to 5 rows and confined
to narrow zone on median inner edge of merus. Spoon-tip processes
bearing 6—8 enlarged lateral lobes.
Abdominal appendages of male long, slender. Arm of appendage
moderately covered with hairs. Pore opening smooth and relatively
narrow.
Color: Anterior portion of carapace between eyestalks bright turquoise,
blending into blue-green band posteriorly. Posterior portion of carapace
dark brown to black. Major cheliped of displaying males white at tips
of fingers. Inside of palm speckled with tan below and dark olive green
above. Outer surface of propodus speckled olive green. Ambulatories
dark brown to black. Females with less intense turquoise coloring and
white chelae.
Uea virens new species
Material Examined: A total of 70 specimens collected from Port
Aransas, Texas and Ocean Springs, Mississippi. Holotype male, USNM
121598; paratype male USNM 122205. Type locality: Port Aransas,
Texas.
Etymology: Virens—alluding to the green band on the anterior border
of the carapace.
Morphological Description: (Fig. 5) Carapace moderately arched and
widest just behind antero-lateral margins. Frontal margin transverse and
angulate, forming obtuse angle at either side of front. Surface of cara-
pace smooth; H-form depression shallow. Antero-lateral margins prog-
282 Proceedings of the Biological Society of Washington
D
Fic. 5. Uca virens new species, holotype male. A, dorsal view. B,
front view. C, major cheliped, inner view. D, same, outer view. E,
minor cheliped, inner view. F, same, outer view.
ressively less distinct posteriorly and blending into smooth surface of
carapace lateral to cardiac region. Another short ridge, just above last
two pairs of walking legs, weakly developed if present. Oblique margins
weakly convergent, forming right angle when extended. Carapace about
2.8 times wider than front, but proportions varying with size (Fig. 2).
Minor cheliped serrated along distal one-half of inner margin of both
fingers, except near flattened spooned tips. Distinct gap present when
Two New Fiddler Crabs 283
5.0 SEC.
LONGISIGNALIS
Fic. 6. Above: Topography of waving gestures displayed by males
of the three species. Graph below each diagram indicates vertical posi-
tion of tip of major cheliped as a function of time during the wave, with
deflections indicating a “jerk.”
Below: Temporal pattern of consecutive waves produced in a series
by single males of each species. The deflections indicate the beginning
and end of single wave.
fingers are closed. Scattered row of hairs extending along serrated
margins of both fingers.
Outer surface of major cheliped smooth, but covered with many small,
flat tubercles. Propodus about 1.88 times as long as dactyl, but propor-
tions varying with size (Fig. 3). Distinct tuberculate submarginal ridge
on palm, extending from proximal edge of propodus to lower margin
at level below articulation with dactyl. Teeth of both fingers well
developed; usually single enlarged tooth present on propodus, %3 of way
to tip. Tip of propodus blunt and containing 3-4 large teeth. Inner
surface of palm with oblique ridge of large tubercles, often arranged in
irregular double row for short portions near pollex, and extending close
to lateral margin of carpal cavity.
Carpus with oblique ridge on inner surface extending from lower
margin, where it contains 3-8 tubercles, to upper surface; there, it con-
284 Proceedings of the Biological Society of Washington
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Two New Fiddler Crabs 285
tinues proximally along upper margin of carpus with a group of 3-8
tubercles almost to point of articulation with merus.
Supplementary Specific Characters: (Fig. 4) Spoon-tipped hairs on
merus of second maxilliped totalling 50-75 and arranged in 5-6 rows.
These distributed along distal half of inner edge of merus, but most
concentrated medially. Spoon-tipped processes bearing 5-7 enlarged
lateral lobes.
Abdominal appendages of males similar to U. longisignalis. However,
pore opening wider and surrounded by sharp spines.
Color: Anterior border of carapace white. Anterior surface of carapace
with green band, extending posteriorly over 4% of dorsal surface. Posterior
surface of carapace deep brown. Major cheliped of displaying males
white at fingers. Palm purple, outside of propodus red-brown above
and blue-green at base. Ambulatories speckled dark brown. Females
with brown carapace and body; chelae colored grey-white.
Rathbun’s specimens (labeled as U. pugnax) from Texas (USNM
72132) are U. virens. Her collections from Mississippi (USNM 74902)
and Louisiana (USNM 2259) are U. longisignalis.
DIFFERENCES IN COURTSHIP BEHAVIOR
Waving Display: The waving display of the three forms was distinct
in all populations we observed, and individuals of each form could be
easily identified on the basis of these movements (Fig. 6). In U. virens,
the major cheliped was raised vertically in 3-6 distinct jerks, then
immediately returned to the front in one, or rarely, two jerks. The waves
were short in duration (about three seconds) and consecutive waves
were displayed with scarcely any pause. In U. longisignalis the waves
were executed in 8-15 less pronounced jerks as the claw was raised and
lowered. Each wave was rather long in duration (4-8 seconds), and
there was a pause of several seconds between consecutive waves. The
displays of Atlantic U. pugnax were executed without any jerking and,
in contrast to U. virens, the claw was returned to the front at about the
same rate as it was initially extended. Successive waves in Atlantic
populations were each separated by a pause of about two seconds. The
temporal properties of the waves of the three forms are summarized in
Table 1.
There appeared to be little variation in the temporal patterns of wav-
ing displays within conspecific populations found in different locales.
However, U. longisignalis observed at Yankeetown, Florida, showed
consistently shorter wave durations, but no other significant deviations,
than did conspecifics observed at Ocean Springs, Mississippi. The
former were observed during the late summer of 1966, while the latter
were studied in June during the height of the breeding season. Yankee-
town forms showed displays that were often incomplete and executed
at low intensity. It is quite probable that these factors account for the
differences between the two populations. The values for the Atlantic
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286 Proceedings of the Biological Society of Washington
Two New Fiddler Crabs 287
coast forms were comparable to estimates of two seconds (Crane, 1943)
and 2.5 seconds (Salmon, 1965) given for wave durations of U. pugnax
from New York and North Carolina, respectively.
Sound Signals: Males of all three forms produced sounds during
nocturnal low tides for periods of up to several hours. Males could be
observed for shorter periods while producing sounds under the illumina-
tion of a flashlight. In all the crabs, the sounds were produced by
movements of the ambulatory legs rather than by rapping of the major
cheliped against the substrate, as described for U. pugilator, U. speciosa,
and several other tropical species (Salmon, 1967; Crane, 1943). Sound
production by movements of the ambulatories was also characteristic of
U. burgersi Holthuis? and U. rapax (see Salmon, 1967), both closely
related to U. pugnax (according to Crane, 1943).
Each sound, as described here, was composed of several pulses pro-
duced in a series (Fig. 7). In U. pugnax, the sounds were usually com-
posed of 2-3 pulses and intervals between successive sounds averaged
about 7 seconds. In U. virens, most sounds contained two pulses, but
the intervals between successive sounds were about three seconds. In
U. longisignalis, the sounds containing 5-14 pulses, and successive sounds
in a series were each separated by long intervals (8-10 seconds). The
temporal properties of the sonic signals of the three forms are sum-
marized in Table 2.
The signals described above were all produced by lone males. Tactile
stimulation by other crabs, or by touching the ambulatories of sonic
males with a fine twig, elicited responses from the males typical of
other species (Salmon, 1965, 1967), i.e., the stimulated males began
producing sounds more rapidly and moved several millimeters into their
burrows.
DIscussION AND CONCLUSIONS
The behavioral, ecological, and morphological characteristics of pop-
ulations of fiddler crabs, heretofore designated as U. pugnax, indicate
that these forms should be separated into three species: the nominate
form found on the Atlantic coast, and two new species on the Gulf
coast of the United States. The differences between the three forms do
not suggest any sign of intergradation, and the forms on the Gulf coast
are as distinct from one another as they are from other closely related
and well established species, such as U. rapax and U. burgersi. Although
we have compiled a number of lines of evidence to support our con-
clusions, it should be pointed out that it was the distinctiveness of the
waving display of the three forms that first suggested the problem. The
results of our study support Crane’s (1941) original contention that
these displays are species-typical movements within the genus.
In most cases, there was strikingly little variation in the morphology
3 Specimens referred to as U. mordax by Salmon, 1967 are synonymous with U.
burgersi Holthuis.
288 Proceedings of the Biological Society of Washington
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Two New Fiddler Crabs 289
and behavior of conspecific forms making up widely separated popula-
tions along the Gulf coast. These results are consistent with those
obtained by Crane in similar studies of other species in the tropics
(1941). Crane (in Tashian and Vernberg, 1958) found little evidence
of variation in displays within single species throughout their range,
and a noticeable absence of any tendency toward subspeciation in the
genus.
All three forms are members of Crane’s (1941) “Group 2” species,
i.e., they have waves of rather long duration, usually executed with
various degrees of “jerking.” In addition, they produce sounds by move-
ments of the ambulatories as do U. rapax and U. burgersi (see Salmon,
1967). The morphological similarities of the Group 2 species are dis-
cussed in detail by Crane (1941).
It appears that the evolution of sound production in Uca has developed
in at least two independent lines, one including the Group 2 species,
and another including U. pugilator, U. speciosa, and U. spinicarpa as
well as several tropical species. The latter produce sounds by striking
the base of the major cheliped against the substrate (rapping). It is
likely that a detailed study of the many species of fiddlers found in the
tropics may reveal other mechanisms of sonic emission. These mech-
anisms may be characteristics of other species groups, and should enable
the formulation of a more detailed explanation of relationships and the
evolution of courtship behavior in the genus, especially when combined
with additional behavioral, morphological, and ecological data.
LITERATURE CITED
CranE, J. 1941. Eastern Pacific Expeditions of the New York Zoolog-
ical Society. XXVI. Crabs of the Genus Uca from the west
coast of Central America. Zoologica, 26: 145-207.
1943. Display, breeding and relationships of fiddler crabs
(Brachyura, Genus Uca) in the northeastem United States.
Zoologica, 28: 217-223.
Ho.tuuis, L. B. 1967. Ona new species of Uca from the West Indian
region (Crustacea, Brachyura, Ocypodidae). Zoologische
Mededelingen, 42: 6, 51-54.
Ratusun, M. J. 1918. The Grapsoid crabs of America. U.S. Nat.
Mus. Bull. No. 97: 1-447.
Satmon, M. 1965. Waving display and sound production in the court-
ship behavior of Uca pugilator, with comparisons to U. minax
and U. pugnax. Zoologica, 50: 123-149.
1967. Coastal distribution, display and sound production
by Florida fiddler crabs (Genus Uca). Anim. Beh., 15: 449-
459.
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.
290 Proceedings of the Biological Society of Washington
HOG Js
Vol. 81, pp. 291-318 30 August 1968
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
NEW GENERA AND SPECIES OF
BRANCHIOBDELLID WORMS
(ANNELIDA: CLITELLATA )
By Perry C. Hott!
Visiting Research Associate, Smithsonian Institution,
Washington, D.C.
In the taxonomic study of any group of animals, the dis-
covery of possibly primitive survivors of ancestral stocks
commands attention. The new genera and species of bran-
chiobdellid worms described herein are such animals. This
paper treats of them as a necessary prelude to further con-
siderations of evolution within the order Branchiobdellida.
In addition to the financial support of the National Science
Foundation (grant GB-372) and release from academic duties
afforded by a Visiting Research Associateship at the Museum
of Natural History, Smithsonian Institution, I should like to
acknowledge the help given me by Drs. Alejandro Villalobos
Fiqueroa and James J. Friauf, Jr., who collected specimens of
three of the species described in this paper. Mrs. Virgie F.
Holt helped collect specimens of the other species and with
the preparation of the manuscript. Dr. Villalobos and Dr.
Horton H. Hobbs, Jr., very kindly identified the host crayfish.
Dr. Hobbs, as for nearly all of my writings, has given gener-
ously of his time in a critical reading of the manuscript.
The methods and terminology used in the taxonomic study
of the branchiobdellids have been described before (inter alia,
Holt, 1953, 1960a, pp. 57, 62-64; 1968; Hoffman, 1963, pp.
277-292). Since, however, within what Stephenson, 1930,
calls a homogeneous group of annelids, thirteen genera are
now known and three new ones are erected in this paper,
1 Department of Biology, Virginia Polytechnic Institute, Blacksburg, Virginia.
33—Proc. Biou. Soc. WAsH., VoL. 81, 1968 (291)
292 Proceedings of the Biological Society of Washington
some attention to what constitutes a genus is necessary and
appropriate.
In most species groups there is a distinctive generic facies,
illustrated by such genera as Ceratodrilus Hall, 1914, and
Xironogiton Ellis, 1919, produced by features of body form
and ornamentation. Yet, some characters, presumably the
result of convergent evolution, appear among some species
of two or more genera otherwise considered distinct. An
example is the presence of dorsal projections among species
of Ceratodrilus and Pterodrilus Moore, 1895a. Such characters
as this and others (raised dorsal ridges on the prosomites of
trunk segments, peristomial lobes and tentacles, a flattened
trunk, etc.) are too inconsistently correlated with features of
the reproductive systems to be used reliably in the recognition
of supraspecific taxa. They are, however, along with the
general shape and appearance of the jaws, responsible for the
superficial resemblances among species of most genera. Some
genera, separated by what has been thought to be fundamental
differences in the reproductive systems, are indistinguishable
in external appearance, for instance, species of Ankyrodrilus
Holt, 1965, and Xironodrilus Ellis, 1918 ( Holt, 1965, pp. 9-10).
With exceptions dictated by previous usage of generic names
and the distinctiveness of groups of species characterized by
jaw shape, external ornamentation and coherence of geo-
graphical distribution, e.g. Ceratodrilus (Holt, 1960a, pp. 57-
58) and Pterodrilus (Holt, 1968b), the genera of branchiob-
dellids are based upon what are considered basic differences
in the reproductive systems. This approach allows the devel-
opment of theories of phylogenetic relationships that cannot
be derived from a classification based upon non-genital char-
acters. The genera now known, defined by these characters,
seem to represent separate lineages.
The important features of the female system are associated
with the spermatheca. This organ is subject to considerable
intraspecific variability accounted for, presumably, by differ-
ences in its degree of distension with spermatozoa. Further-
more, constant differences that occur, the presence or absence
of ental processes and various types of glandular or muscular
thickenings of the wall of the spermathecal bulb, are not
New Branchiobdellid Worms 293
correlated invariably with differences in the male genitalia
regarded as generic characters. Yet it seems to be true that
a spermatheca other than one composed of a simple muscular
ectal duct and thin-walled bulb is associated with primitive
features of the male reproductive system.
The most significant generic characters, then, are thought
to be those of the male genitalia. The importance of the dif-
ferences between the eversible as opposed to a non-eversible
penis was recognized early (Moore, 1895b, p. 498). The haz-
ards associated with this distinction arise from the difficulty
of being sure in preserved animals of the exact functioning
of the bursal complex housing the penis. It is rare that speci-
mens in collections are found with everted or protruded penes,
and observations of copulation in living animals, at least for
North American species, have not been reported. Further,
there exist species in which the penis is intermediate in struc-
ture between those which are indubitably eversible and those
which are clearly protrusible. Since, however, these differences
of the bursa and penis can be associated with other features
of the male system, they are of generic value.
Among other features of the male genitalia believed to be
important is the place of entry of the vasa deferentia into the
spermiducal gland. In one group of genera the deferent ducts
enter the spermiducal gland directly at its ental end. In
other genera the ducts enter the gland somewhere along its
length; that is, there is a blindly ending ental projection of
the gland beyond the region at which the vasa deferentia
enter it.
Finally, the structure referred to as the prostate, a divertic-
ulum of the spermiducal gland may or may not be present.
If present, it may vary from a slight prostatic protuberance
produced by a few apparently non-glandular cells enclosing
a small cavity on the anterodorsal side of the spermiducal
gland to a large, tubular, blindly-ending gland that opens in
common with the spermiducal gland into the ejaculatory duct.
The presence or absence of the prostate, the major differences
in its degree of separation from the spermiducal gland, and
its histological differentiation, or lack thereof, seem to be
valid generic characters.
294 Proceedings of the Biological Society of Washington
The permutations of a relatively few characters within a
group of species that are collectively distinguished by a com-
mon pattern of the male genitalia may be numerous. The
genus Cambarincola Ellis, 1912, for instance, with a distinc-
tive male system that remains basically similar throughout
the genus, contains more than thirty easily recognizable spe-
cies (23 nominal species and several undescribed ones in my
collections). On the other hand, the state of knowledge of
the branchiobdellids and perhaps the pattern of evolution
within the order is such that several genera are, as now known,
monotypic or composed of only a few species. The bran-
chiobdellids present relatively few characters suitable for
taxonomic analysis and the decision as to what constitutes a
genus among them admittedly involves elements of arbitrar-
iness and subjectivity. The system that I am developing, of
which this paper constitutes a part, is based upon theories of
phylogeny that have been presented in preliminary form —
elsewhere (Holt, 1968a).
Sathodrilus new genus
Type-species: Sathodrilus carolinensis new species, here designated.
Diagnosis: Medium-sized branchiobdellid worms (known species 1.6
to 4.6 mm in length) with two pairs of testes; unpaired nephridiopore
on dorsum of segment III; body terete, without peristomial tentacles or
dorsal projections on trunk segments; spermiducal gland with vasa
deferentia entering entally; prostate, if present, consisting of bulb-like
prostatic protuberance on anterior or dorsal border of spermiducal gland;
ejaculatory duct present; penis eversible, but attached by cytoplasmic
strands to inner wall of penial sheath and without cuticular hooks;
spermatheca with or without ental process.
Etymology: From Greek, sathon, one with a large penis, and drilus,
a penis, by extension worm. Masculine.
Affinities: Among the branchiobdellids with eversible penes, rudi-
mentary (or vestigial) prostates or none, vasa deferentia entering the
ental end of the spermiducal gland and a common or unpaired anterior
nephridiopore are Sathodrilus, Ceratodrilus, Oedipodrilus Holt, 1967a,
Magmatodrilus Holt, 1967b, and a second new genus erected herein,
Tettodrilus. Of these, Sathodrilus is most closely related to Ceratodrilus
and is the unnamed relative of the latter mentioned previously (Holt,
1960a, p. 58). The two genera differ most strikingly in the presence
of dorsal projections on segments II to VIII and the peristomial tentacles
of Ceratodrilus. The jaws of the two genera are also quite different:
those of Sathodrilus are subtriangular in shape, as is common among
New Branchiobdellid Worms 295
other related genera (Figs. 2b, c, 3b, 4c, 6c, d), while those of Ceratod-
rilus are subrectangular with the tooth-bearing border slightly concave
(Holt, 1960, Figs. 15-16).
Oedipodrilus has a distinct prostate, though it is incompletely divided
from the spermiducal gland, and a completely eversible, tubelike penis
that is unattached to the inner wall of the penial sheath portion of the
bursa. In general, species of Oedipodrilus -are smaller, more gracile
animals than those of Sathodrilus and often have dorsal ridges on the
prosomites of some or all body segments.
In Magmatodrilus, the vasa deferentia enter the large and very long
spermiducal gland entally, but there is no prostate or prostatic pro-
tuberance. The most striking difference between these not too closely
related genera is in the bursae. That of Magmatodrilus is exceptionally
long, but the penial sheath is relatively very short. The penis, however,
contrary to my former statement (Holt, 1967b, pp. 3, 4), is probably
eversible, since it and the penial sheath have the structure of those of
Sathodrilus. Most of the bursa of Magmatodrilus is composed of a thick
muscular layer enclosing the atrial canal leading into the atrium proper.
The jaws are entirely different in appearance from those of Sathodrilus.
They are subrectangular with a 6/5 dental formula.
Sathodrilus is more distantly related to Tettodrilus (Figs. 9a, b, c, d).
The prostate of Tettodrilus is incompletely divided from the spermiducal
gland; that is, it does not extend to a common junction with the latter
and the ejaculatory duct, but it is distinct with a well-defined prostatic
bulb. In addition, the bursae and penes of the two genera differ
markedly: the bursa of Tettodrilus is, in external appearance and shape,
like that of the genus Cambarincola, but the penial sheath encloses an
eversible penis that is different in structure from that of Sathodrilus
(see below, p. 313).
The genera Ellisodrilus Holt, 1960b, Cambarincola and Pterodrilus
have prostates, but mostly short, ovoid bursae, and all have protrusible
penes. The other genera of the branchiobdellids lack prostatic pro-
tuberances or prostates, and, if the vasa deferentia enter the ental end
of the spermiducal gland, the penis is protrusible.
Remarks: Species of Sathodrilus are described herein from localities
in South Carolina, Georgia, and Mexico. The distribution may be more
extensive than these records indicate, but species of the genus as defined
are localized and rare; for although the branchiobdellid fauna of North
America is comparatively poorly known, these are the only occurrences
of Sathodrilus from over 1,600 locality records represented in my collec-
tions. This is the sort of distribution expected for a primitive group and
Sathodrilus appears to stand close to the ancestral stocks of the genera
Cambarincola and Pterodrilus and, in addition, to be closely related to
the likewise localized species of Ceratodrilus and the possibly more
advanced and certainly widespread, but still in some ways primitive,
Oedipodrilus.
296 Proceedings of the Biological Society of Washington
FicureE 1. Sathodrilus carolinensis: a, lateral view of reproductive
systems; b, longitudinal optical section of bursa and penis. Abbrevia-
tions: b, bursa; ba, bursal atrium; dl, deferent lobe; ejd, ejaculatory duct;
f, male funnel; af, inner atrial fold; p, penis; prt, prostatic protuberance;
ps, penial sheath; sb, spermathecal bulb; sd, spermathecal duct; sg, sper-
miducal gland; vd, vas deferens.
Sathodrilus carolinensis new species
Figures 1, 2
Type-specimens: Holotype, USNM 37107, one paratype, USNM
37108, and one paratype, PCH 1333, from Cambarus latimanus
(LeConte) and Cambarus sp. taken in a small, sandy stream about 11.5
miles southwest of Anderson, Anderson County, South Carolina, on U.S.
Highway 29, by Perry C. and Virgie F. Holt, 21 March, 1961.
Diagnosis: Slender, terete worms; upper lip entire, lower with shallow
median indentation; no oral papillae; jaws small, delicate, dental formula
5/4; bursa large, about 34 body diameter in length, bursal atrium about
¥% total bursal length, penial sheath expanded, greater in diameter than
bursal atrium; eversible penis attached by numerous thin strands to inner
wall of penial sheath (Fig. 1b); ejaculatory duct short, less than %4
length of bursa, thin-walled; spermiducal gland about twice its greatest
diameter in length, tapering ectally from junction of ental-most and
median thirds, with short, but distinct deferent lobes forming most of
ental third of organ; prostatic protuberance at region of greatest diameter
of spermiducal gland; spermatheca long, subequal to body diameter in
length, clavate or spatulate, ectal duct widening gradually, spermathecal
New Branchiobdellid Worms 297
.
om
FicureE 2. Sathodrilus carolinensis: a, holotype; b, upper jaw; c,
lower jaw.
bulb not externally set off from ectal duct, thin-walled without ental
process.
Etymology: The adjectival form of Carolina.
Description: Sathodrilus carolinensis is a medium-sized, slender worm.
The three individuals of the type series have the following dimensions:
total length, 3.1-4.1 mm; greatest body diameter (segment VIII), 0.39-
0.57 mm; head length, 0.43-0.48 mm; head diameter, 0.27-0.30 mm;
diameter, segment 1, 0.28-0.30 mm; diameter, sucker, 0.31-0.34 mm.
The holotype is the longest of these specimens, but not consistently the
greatest in other dimensions.
In external appearance (Fig. 2a), specimens of S. carolinensis are not
remarkable. The head appears slender, although actually it is of rather
usual proportions among the branchiobdellids. The peristomium, as
usual, is divided by lateral indentations into dorsal and ventral lips. The
upper lip is without lobes or projections; the lower is marked by a
shallow, median indentation. Oral papillae are absent. Externally, the
peristomium, again as usual, is set off by a shallow furrow; otherwise,
there is one annular groove surrounding the head at about its midlength
that is marked internally by what Ellis (1919, p. 243) called a “pharyn-
geal diverticulum” (actually an inner ring-like indentation of the wall
of the pharynx) that I shall hereafter refer to as a pharyngeal sulcus.
In some branchiobdellids there are more than one.
The jaws are small (Fig. 2b, c). The upper bears five sharply pointed
298 Proceedings of the Biological Society of Washington
teeth, the lower four. Though not shown in the outline drawing, these
teeth are separated by deep valleys in the body of the jaws that extend
almost to their base (cf. Fig. 7d of the jaws of Cronodrilus ogygius).
The anterior nephridiopore is located as usual on the dorsum of
segment III and is clearly visible in the holotype, but not in the other
specimens.
A fairly well defined clitellum is present on segment VII. There are
no supernumerary muscles in the prosomites of body segments, con-
sequently the body outline is rather smooth.
The spermiducal gland is provided with well-marked, but relatively
small, deferent lobes (Hoffman, 1963, pp. 286-287). These unite only
a slight distance ental to the position of the prostatic protuberance.
From the prostatic protuberance which marks the region of greatest
diameter of the spermiducal gland, the latter narrows gradually so that
its outer or ectal end as it joins the ejaculatory duct is no greater in
diameter than the latter.
The prostatic protuberance is composed of a small group of cells that
appear to have been evaginated by some force from the surrounding
glandular epithelial cells. These cells of the prostatic protuberance are,
however, much more finely granular than the cells of the epithelium of -
the spermiducal gland. There is a very small, but distinct space enclosed
by the cells of the prost2tic protuberance that is continuous by a very
narrow canal with the lumen of the spermiducal gland. The prostatic
protuberance is located on the anterior border of the spermiducal gland
and is about % the length of the latter from its ental end (including
the deferent lobes in the total length of the spermiducal gland), but
median to and only slightly ectal to their junction.
The ejaculatory duct is short and thin-walled.
The bursa is marked (Fig. 1b) externally by a constriction between
the penial sheath and atrial regions. The penial sheath, composing about
2% of the length of the organ, is somewhat greater in diameter than the
atrial region of the bursa, but is quite thin-walled. The penis is clearly
eversible: that of the holotype, indeed, is partially everted into the
lumen of the bursal atrium. The penis is composed of a thin tubular
membrane (Fig. 1b) apparently continuous with the inner cuticular
lining of the bursal atrium and the cuticle of the outer body surface.
This is attached by several thin strands to the inner wall of the penial
sheath. The penial sheath is, as usual, covered by the peritoneum and
its wall is composed of a layer of muscle the strands of which run
lengthwise of the organ and continue as the muscular layer of the
ejaculatory duct. These muscles are continuous with, and apparently
are derived from, the outer circular muscles of the body wall. Internal
to them, the penial sheath is lined with a layer of irregular cells with
granular cytoplasm. Processes of these cells produce the strands that
attach the inner wall of the everted penis to that of the penial sheath.
In the uneverted position of the penis, most of the interior of the penial
sheath portion of the bursa is composed of intercellular spaces.
New Branchiobdellid Worms 299
The atrium of the bursa is enclosed by a relatively thick layer of
muscle. A narrow canal leads from the outside into an expanded cavity
which is the atrium proper. It is not clear that this portion of the atrium
is eversible, but by analogy with specimens of Oedipodrilus with everted
penes, it is not: the penis as it everts is simply exserted through the
outer canal of the atrium. There is an inwardly projecting fold of the
atrium, but it is not prominent (see below, p- 301).
The spermatheca is elongate club-shaped, extends to the dorsal wall
of the coelom in segment V, and is in actuality, since it lies obliquely in
the coelom and bends around and slightly over the gut, at least as long
as the diameter of the segment. It gradually increases in diameter from
the outlet pore almost to its ental end, and there is no external indication
of the boundary between the spermathecal duct and the bulb. The
spermathecal duct is relatively long and histologically similar to that
described for other branchiobdellids (Holt, 1960a, p. 70) in which it is
composed of muscle layers and lined with tall glandular cells, the inner
ends of which project separately into the lumen. The bulb is thin-walled
and has no features of note.
Variation: The three known specimens of Sathodrilus carolinensis
differ in size and there are minor differences between them in propor-
tions. From such a limited series, nothing else can be said about the
variability of the species.
Affinities: The other new species of Sathodrilus described below are
the closest known relatives of S. carolinensis. For a discussion of the
affinities of all these species, see S. veracruzicus (p. 307, below).
Hosts: Cambarus latimanus (LeConte) and C. species.
Distribution: S. carolinensis is known only from the type-locality.
Material Examined: The type-series.
Sathodrilus villalobosi new species
Figure 3
Type-specimens: Holotype, USNM 37101, four paratypes, USNM
37102, and four paratypes, PCH 208, from Paracambarus paradoxus
(Ortmann) taken at Tetela de Ocampo, Puebla, Mexico, by Alejandro
Villalobos F., May, 1941.
Diagnosis: Medium-sized, terete branchiobdellids; upper lip entire,
lower with shallow median indentation; no oral papillae; jaws heavy
and dark, dental formula 1/4; bursa large, subequal to body diameter
in length, bursal atrium less than 1% total bursal length, diameter of
penial sheath less than that of bursal atrium; eversible penis composed
of tube attached by few strands to inner wall of penial sheath; ejacula-
tory duct short, about % length of spermiducal gland; length of sper-
miducal gland about twice its diameter, about 144 that of the bursa,
tapering toward each end from prostatic protuberance, and without
deferent lobes; spermatheca with long, narrow spermathecal duct and
globose bulb.
300 Proceedings of the Biological Society of Washington
FicurE 3. Sathodrilus villalobosi: a, holotype; b, lateral view of jaws,
upper jaw above; c, lateral view of reproductive systems; d, longitudinal
optical view of bursal atrium and ectal end of penis.
Etymology: I take pleasure in naming this Mexican species in honor
of its discoverer, Dr. Alejandro Villalobos F. of the Universidad Nacional
Autonoma de México, in appreciation of his hospitality and help in
collecting branchiobdellids in Mexico.
Description: Sathodrilus villalobosi is a medium-sized worm. Five
individuals of the type-series have the following dimensions: total length,
2.2-2.5 mm; diameter, segment VI, 0.42-0.49 mm; head length, 0.33-
0.51 mm; head diameter, 0.34-0.45 mm; diameter, segment I, 0.32-0.39
mm; diameter, sucker, 0.32-0.39 mm.
The head is, in all specimens, short and thick, and though this
appearance is exaggerated in the type series, it reflects the actual pro-
New Branchiobdellid Worms 301
portions of the animals. Other than a similar proportionally greater
thickness of the body segments, there is little worthy of comment about
the external appearance of the animals. The upper lip is entire; the
lower has a shallow median indentation. There are no oral papillae
detectable in the type series, but each of a series of fifteen specimens
collected later at Agua Fria show these papillae. The peristomium is
set off by a deep encircling furrow. There is one other shallow groove
surrounding the head and a single pharyngeal sulcus internally.
The jaws (Fig. 3b) are distinctive. They are larger and darker than
those of S. carolinensis. The upper is triangular in both lateral and
en face view and bears on its apex a large tooth. The sides of the upper
jaw, which in most species with such jaws bear other teeth, are straight
or slightly wavy and lateral teeth are represented by at most very low
elevations. The lower jaw is triangular in lateral view and subrectangular
in en face view. It bears two low rounded teeth on each side of the
median line and two very low knobs lateral to these.
The anterior nephridiopore cannot be seen in the holotype, but a
careful examination of other topotypical specimens shows it to be un-
paired on the dorsum of segment III.
The body outline is smooth. The clitellum is not unusually prominent.
Deferent lobes are absent, or at least undetectable, and the vasa
deferentia enter the narrow ental end of the spermiducal gland close
together. The latter is small, about % of the length of the bursa and
% its own length in diameter, narrowed at each end, and widest at the
level of the prostatic protuberance.
The prostatic protuberance is located on the spermiducal gland about
1% the length from the ental end of the latter. It is composed of a small
group of cells with a lumen between them that communicates with that
of the spermiducal gland. The cells of the prostatic protuberance do not
differ in appearance from those of the spermiducal gland.
The ejaculatory duct is short, about % the length of the spermiducal
gland, and somewhat expanded. Its structure does not depart from that
of other species.
The bursa (Figs. 3b, c) is large and subequal to the body diameter
in length. The atrial region is somewhat shorter than 14 of the total
length of the bursa. The diameter of the penial sheath is less than that
of the atrial region and the penial sheath is not as expanded as that of
S. carolinensis (cf. Fig. la).
The penis, an eversible tube, is longer than the penial sheath, looped
and coiled entally within the cavity of the latter. A few strands attach
it to the inner wall of the sheath, particularly at the ectal end. It is
lined by a continuation of the cuticular lining of the bursal atrium
and has an apparently muscular wall of some thickness (Fig. 3d).
Internally, the bursal atrium is characterized by a thick muscular fold
that extends into the atrium and divides it into two compartments which
communicate by the central space defined by the inner edge of the fold
and narrow clefts extending from this space radially into the fold. This
302 Proceedings of the Biological Society of Washington
feature of the bursa is not prominent in S. carolinensis (cf. Figs. 1b
and 3d), and has been described for other species (Holt, 1949, pp. 544,
554, figs. 9, 16; Hoffman, 1963, p. 290, fig. 3). This atrial fold is
itself eversible in these other species (ibid.); whether it is in S. caro-
linensis and S. villalobosi is unknown: it is not obvious that it is, but
there is no doubt that the penis is eversible.
The spermatheca consists of a long spermathecal duct of the usual
composition and a short, expanded, subglobose bulb set off by a con-
striction from the duct. There is no ental process.
Variation: Other than differences in size, differences in proportions
that appear to be the effects of the method of killing and preservation
and variations in the shape of the spermathecal bulb, and the prom-
inence of the atrial fold depending upon the degree of expansion of the
bursal atrium, there are no detectable variations in the material I have
seen.
Affinities: See p. 307 below.
Hosts: Sathodrilus villalobosi is known from Paracambarus paradoxus
(Ortmann) and Procambarus contrérasi (Creaser).
Distribution: In addition to the type-locality, Sathodrilus villalobosi
has been taken from Procambarus contrerasi in Arroyo de San Diego,
3 km southeast of Agua Fria, Puebla, Mexico, by Alejandro Villalobos
F., 23 October 1948, and by Alejandro Villalobos F., Patricio Gonzales
K., and Perry C. and Virgie F. Holt, 14 July 1962.
Material Examined: Other than the types listed above, 18 specimens
from the type locality, PCH 208, two, PCH 202, and 15, PCH 1593,
from Agua Fria, Puebla, in all, 35 specimens, have been studied.
Sathodrilus megadenus new species
Figures 4, 5
Type-specimens: Holotype, USNM 37109, two paratypes, USNM
37110, and two paratypes, PCH 1346, from Cambarus latimanus
(LeConte) taken in a small stream, 3.1 miles north of Buchanan, Haral-
son County, Georgia, on U.S. Highway 27, by Perry C. and Virgie F.
Holt, 25 March 1961.
Diagnosis: Stout, medium-large worms; lips entire or slightly lobed;
no oral papillae; jaws of medium size, dental formula 5/4; bursa large,
about %4 body diameter in length; length of bursal atrium about % total
bursal length; penial sheath subglobose to pyriform, greater than bursal
atrium in diameter; eversible penis attached by many strands to inner
wall of penial sheath; ejaculatory duct long, subequal in length to penial
sheath; spermiducal gland very long, length greater than body diameter,
diameter about 14 its length, without deferent lobes; prostatic bulb
present; spermatheca with spermathecal bursa and ental process, bulb
cylindrical, of moderate size, total length about %4—4% body diameter.
Etymology: From Greek, mega, large, and adenos, gland, in reference
to the large size of the spermiducal gland.
ai
New Branchiobdellid Worms 303
PE
a d
Ficure 4. Sathodrilus megadenus: a, lateral view of reproductive
systems; b, longitudinal optical section of prostatic protuberance and
portion of spermiducal gland; c, lateral view of upper jaw; d, oblique
view of lower jaw.
Description: Sathodrilus megadenus is a medium-large, stout worm.
The five specimens of the type series have the following dimensions:
total length, 3.9-4.6 mm; greatest body diameter, segment VI, 0.7-1.0
mm; head length, 0.8-1.0 mm; head diameter, 0.5-0.7 mm; diameter,
segment I, 0.5-0.6 mm; diameter, sucker, 0.5-0.6 mm. The head is
about 14 of the total length and almost as great in diameter. Segment
VI, instead of the egg-bearing segment VII, has the greatest diameter.
Segment I and the sucker are subequal to the head in diameter. Those
dimensions confer a distinctive appearance within the genus upon
individuals of S. megadenus (p. 297 and 300 above; Figs. 2a, 3a).
In other respects, the external appearance is not remarkable (Fig. 5).
It is not possible in lateral view (and all the specimens are so mounted )
to be sure that the lips are entire, but they appear to be so or at most
only slightly lobed. The head shows little sign externally of segmenta-
tion: the peristomium is set off by a shallow encircling furrow and, at
the region of the single pharyngeal sulcus, there is externally a still more
obscure shallow furrow. The prosomites of at least the three anterior
body segments are slightly raised. The clitellum is most prominent on
segment VI instead of VII.
The anterior nephridiopore is unusually prominent in whole mounts
for such large worms, but otherwise is unremarkable.
The jaws (Figs. 6c, d) are not unusual. The upper one is distinctly
triangular in lateral and en face view with five sharply pointed teeth of
which the median is the largest; the lower one is triangular in lateral
view, subtriangular in en face view and bears four teeth, of which the
paramedian ones are the largest.
304 Proceedings of the Biological Society of Washington
Ficure 5. Sathodrilus megadenus: holotype.
The vasa deferentia enter the spermiducal gland directly; deferent
lobes are absent. The large spermiducal gland, however, is the most
striking character of the animal. It extends from the lower anterior
border of the bursa posterodorsally over the bursa, ending at its junction
with the ejaculatory duct. opposite its beginning (Fig. 4a). The total
length exceeds the greatest diameter of the body. It is also greater in
diameter than those of related species. The diameter increases somewhat
ectally, but not greatly, and it ends abruptly at its junction with the
ejaculatory duct.
The prostatic protuberance is not easily seen in whole mounts, but it
is present as a group of flattened epithelial cells enclosing a small lumen
that communicates with that of the spermiducal gland (Fig. 4b) ap-
proximately at the midlength of the latter (Fig. 4a).
The ejaculatory duct is subequal in length to that of the penial sheath
portion of the bursa and runs ventrad from the upper end of the latter
along its posterolateral border (Fig. 4a). The epithelial lining and
enclosing layer of muscle of the ejaculatory duct are prominent.
The bursa is quite large, though the greater diameter of segment VI
makes it appear proportionally smaller than those of the other species
of the genus. The penial sheath is subglobose, composes about % of
the bursa, is thin-walled and much of its interior is occupied by spaces
between the penis and the apparently muscular strands that attach the
penis to its inner wall (cf. S. carolinensis, p. 298 and Fig. 1b).
The penis is difficult to interpret in my material. It appears to have
a longitudinally folded cuticular lining and a thin investment of cyto-
plasm and to be somewhat longer, hence, slightly looped, than the
enclosing penial sheath. There are numerous strands that converge from
the inner wall of the penial sheath to attach to the penis. There is,
however, no doubt as to its eversibility and essential similarity to those
of other members of the genus.
sia all
New Branchiobdellid Worms 305
The atrial region of the bursa widens somewhat abruptly from the
outlet canal and its ental-most portion, the atrium proper, is enclosed
by a thick layer of muscle. I cannot determine whether the atrial fold
described above (p. 301) for S. villalobosi is present or not.
The spermatheca of S. megadenus is distinctive (Fig. 4a). There is
a prominent sphincter formed of circular muscles around the outer
portion of the spermathecal duct just ental to the spermathecal pore.
The spermathecal duct is relatively narrow, but heavily muscular and
its inner wall projects as a circular fold into a short, somewhat expanded,
ental portion of the duct that is lined with a glandular epithelium typical
of the lining of the spermathecal duct in other species. An everted
“spermathecal bursa” (Holt, 1960a, p. 64) has never been seen, but the
structure of the ectal part of the spermathecal duct of S. megadenus
strongly suggests eversibility. The middle third of the spermatheca
makes up the spermathecal bulb which is thin-walled and filled, as usual,
with spermatozoa. The ental third is a narrow cylindrical process with
a thick wall that almost obliterates its lumen. Apparently, the wall of
this ental process is muscular, but this cannot be asserted with certainty
in the absence of histological studies.
Variations: Other than minor differences in size, there are no detect-
able variations in my material.
Affinities: See p. 307 below.
Host: Cambarus latimanus (LeConte ).
Distribution: Sathodrilus megadenus is known only from the type-
locality.
Material Examined: The five specimens of the type-series.
Sathodrilus veracruzicus new species
Figure 6
Type-specimens: Holotype, USNM 37105, three paratypes, USNM
37106, and three paratypes, PCH 1623, from Procambarus hoffmanni
(Villalobos), taken from Coyutla, Veracruz, by Alejandro Villalobos F.,
16 April 1949.
Diagnosis: Small, slender, terete worms; lips entire; dental formula
5/4; bursa long, exceeding body diameter in length, penial sheath about
34 total length, less in diameter than bursal atrium; eversible penis with
thickened wall attached by few strands to penial sheath; ejaculatory
duct short, about % length of spermiducal gland; spermiducal gland
small, about % as long as bursa; no prostatic protuberance; spermatheca
long, spermathecal duct subequal to body diameter in length, bulb
cylindrical, long, no ental process.
Etymology: Of, or pertaining to, Veracruz.
Description: Sathodrilus veracruzicus is a small, slender, terete worm.
Four specimens of the type series have the following dimensions: total
length, 1.6-1.8 mm; greatest diameter, segment VI, 0.28-0.34 mm; head
length, 0.26—0.29 mm; head diameter, 0.18-0.21 mm; diameter, segment
1, 0.19-22 mm; diameter, sucker, 0.20-0.22 mm.
306 Proceedings of the Biological Society of Washington
Ficure 6. Sathodrilus veracruzicus: a, lateral view of reproductive
systems; b, holotype; c, lateral view of jaws; d, en face view of jaws,
upper jaw above.
The head of S. veracruzicus is not of unusual size or proportions. The
lips are entire. The holotype appears to have at least some indistinct
oral papillae, but these cannot be seen in other specimens. The external
encircling furrow of the head is shallow and there is only one correspond-
ing pharyngeal sulcus.
The jaws are of usual size for a small worm. The dental formula is
5/4; both jaws are triangular in lateral view, subrectangular in en face
view (Fig. 6c, d).
The single nephridiopore occurs dorsally on segment III.
There are no dorsal ridges of the prosomites, but the intersegmental
furrows are prominent. Clitellar glands are located upon the dorsa of
segments VI and VII, but they are not prominent.
The vasa deferentia enter the spermiducal gland together at its ental
end. There are no deferent lobes. The spermiducal gland is very small,
about 1% the length of the bursa. The cells composing it are almost
cuboidal (instead of the usual columnar type) and the lumen is dilated,
perhaps a stage in a secretory cycle. The gland is only slightly tapered
New Branchiobdellid Worms 307
at its ends and there is no sign at all of a prostate or prostatic pro-
tuberance.
The ejaculatory duct is short, but not as short as it appears to be in
fore-shortened view (Fig. 6a). It is about 4 the length of the sper-
miducal gland.
Relatively, the bursa is very long, its length exceeding the body
diameter. The penial sheath constitutes about °4 of this length. The
penis consists of a looped tube, with a wall of noticeable thickness, that
is attached by a few strands to the inner wall of the penial sheath. The
bursal atrium is expanded; its length is subequal to the diameter of the
coelom in which it lies. Its wall is heavily muscular and folded, but it
is not possible to determine clearly the inner structure of the atrium in
the available specimens.
There is no spermathecal bursa. The spermathecal duct is long,
subequal to the body diameter in length, with the usual glandular lining
and muscle layers. The bulb is thin-walled, cylindrical and bent cephalad
over the gut (Fig. 6a).
Variation: Except for differences in size and the possible presence of
oral papillae in some specimens, no variations have been observed.
Affinities: Among the four species of Sathodrilus described herein,
S. carolinensis and S. villalobosi are most closely related. S. megadenus
and S. veracruzicus stand somewhat apart in different ways from the
others.
S. veracruzicus is the smallest in body size, S. megadenus the largest,
and the others lie in between, with S. carolinensis as a somewhat longer
and more slender animal. Any phylogenetic significance that these dif-
ferences in size and proportions may have is certainly obscure, but the
smaller size and more slender proportions of S. veracruzicus are consonant
with its other primitive features.
The jaws of all four species are similar in general shape and all, except
S. villalobosi, have a 5/4 dental formula. Those of S. veracruzicus and
S. carolinensis may more nearly approach the primitive conditions, but
those of S. megadenus are not markedly different and the heavier jaws
with a 44 dental formula of S. villalobosi, apparently an advanced
character, can easily have been derived from a common ancestor with
jaws like those of S. carolinensis or S. veracruzicus.
Oral papillae are apparently present in S. villalobosi and S. veracruz-
icus and absent in the other species.
The spermiducal gland of S. megadenus is very different from those
of the other species in its much greater absolute and relative size. That
of S. carolinensis is the only one provided with deferent lobes, pre-
sumably a primitive feature, but that of S. veracruzicus lacks a prostatic
protuberance, a still more primitive characteristic. S. villalobosi has a
spermiducal gland with a prostatic protuberance and without deferent
lobes. There are some minute but significant differences in the prostatic
protuberances of the three species that have them. The cells composing
the prostatic protuberance of S. villalobosi differ from those of the
308 Proceedings of the Biological Society of Washington
spermiducal gland (as far as can be determined in whole mounts) only
in being displaced to the outer surface of the gland; those of S. caro-
linensis appear more finely granular than the obviously secretory cells
of the spermiducal gland; and, finally, those of S. megadenus form a
distinctly flattened epithelium (Fig. 4b).
The ejaculatory ducts of all species are much alike. That of S. me-
gadenus is unusual in its length, which is consonant with the unusual
length of the spermiducal gland. ;
The bursa and its components unify the genus. In all the species the
penial sheath region is large and encloses an eversible penis. The penial
sheath of S. villalobosi is less in diameter than the atrial region; in the
other species the reverse is true. The wall of the penis is thin, apparently
composed only of cuticle with minute attachments of cytoplasmic
strands, in S. carolinensis; in the other species it is thicker. The most
striking bursal modification is found in S. villalobosi with its prominent
inner atrial fold which is present, but not so greatly developed in S.
carolinensis, and possibly is absent in the other species.
The spermatheca of S. megadenus has a spermathecal bursa and an
ental process; the other species lack these structures and their sper-
matheca differ only in the shapes and relative sizes of the spermathecal
ducts and bulbs. There does seem to be a relationship between such
features of the spermatheca as those possessed by S. megadenus and
large bursae with eversible penes and, on the basis of current views of
phylogeny, they are primitive. In all the species of Sathodrilus, the
spermathecal ducts are long and roughly correspond in length to the
estimated length of the everted penes.
S. veracruzicus is the most primitive of these species, if it is admitted
that the presence of a prostatic protuberance indicates an advance over
its absence. S. carolinensis and S. villalobosi stand at about the same
level of evolutionary advance and S. megadenus is the most specialized
of the four. But such conclusions, although supported by the evidence,
are not as important as the more general one that we have in these
species a group of primitive forms that are the survivors of the stock
that gave rise to the seven genera that collectively constitute the major
portion of the North American branchiobdellid fauna.
Host: Procambarus hoffmanni (Villalobos).
Distribution: Known only from the type locality.
Material Examined: The type series.
Remarks: At one time, I considered S. veracruzicus a member of the
genus Oedipodrilus and so stated in referring to the distribution of the
latter (Holt, 1967a, p. 60). Oedipodrilus is composed of a number of
species with collectively a wide geographical range that does not, how-
ever, to my knowledge, include Veracruz.
Cronodrilus new genus
Type-species: Cronodrilus ogygius new species, here designated.
Diagnosis: Branchiobdellid worms with two pairs of testes; unpaired
—————————
New Branchiobdellid Worms 309
nephridiopore on dorsum of segment III; body terete, without peristomial
tentacles or projections on trunk segments; spermiducal gland with vasa
deferentia entering ectad to ental end; no prostate or prostatic pro-
tuberance; ejaculatory duct present; penis eversible, lying free in long,
slender penial sheath; spermatheca with ental process.
Etymology: From Greek, Kronos, the father of Zeus, hence, an ancient
ancestor, and drilus, a worm. Masculine.
Affinities: The North American genera of branchiobdellids with
spermiducal glands that extend entally beyond the junction of the vasa
deferentia with them are Bdellodrilus Moore, 1895b, Ankyrodrilus Holt,
1965, and Xironogiton Ellis, 1919. In addition, Cirrodrilus Pierantoni,
1905, from eastern Asia, and Branchiobdella Odier, 1823, of eastern
Asia and Europe, have this arrangement of the vasa deferentia and the
spermiducal gland. Cronodrilus differs from all these genera in the
smaller size of its spermiducal gland and the shortness of the blindly-
ending portion ental to the entry of the vasa deferentia. Cronodrilus and
Bdellodrilus further differ in the structure of the jaws, those of Bdel-
lodrilus are unusually modified (Moore, 1895b, fig. 9); in the numerous
aggregations of glandular cells, including lateral and bursal ones, present
in Bdellodrilus (Moore, 1895b, pp. 505-506, 522-523) and absent in
Cronodrilus; in the branched spermatheca of Bdellodrilus; and in the
generally thinner and more glandular body wall of Bdellodrilus which
confers a distinctly parasitic appearance upon the genus. Cronodrilus
and Ankyrodrilus differ in jaw structure. The jaws of Ankyrodrilus are
more nearly rectangular and the teeth are differently arranged. The
bursa and penis of Ankyrodrilus are unlike those of any other branchiob-
dellid and though externally the bursa has the form found in genera with
eversible penes, it is not clear that the penis of Ankyrodrilus is eversible
(Holt, 1965, p. 16, fig. 9), while that of Cronodrilus is. In addition,
species of Ankyrodrilus are flattened, not terete as in Cronodrilus, and
the anterior nephridia open separately instead of by a common pore.
The posterior segments of species of Xironogiton are flattened; the
anterior nephridia open separately; the bursa, but not the penis, is
eversible; the spermatheca is greatly reduced in size: in all of these
respects Xironogiton differs from Cronodrilus. Cirrodrilus is an east
Asian genus and includes all the species assigned by Yamaguchi (1934)
to Stephanodrilus. The latter name is a junior synonym of Cirrodrilus
(Holt, 1967b, p. 3). Both Branchiobdella and Cirrodrilus differ from
Cronodrilus in having two anterior nephridiopores. The penis of Cir-
rodrilus is attached to the inner wall of the penial sheath (Yamaguchi,
1934, pp. 194-195), but that of Cronodrilus lies free in it. Branchiob-
della has only one testicular segment (V) and both the penial sheath
and spermiducal gland are excessively elongated, narrow tubes. Cronod-
rilus has two testicular segments. The penial sheath and spermiducal
gland, while elongated and narrow, are not as excessively so as those
of Branchiobdella. All these genera are related in various ways, par-
ticularly in the nature of the spermiducal gland and the absence of any
310 Proceedings of the Biological Society of Washington
Ficure 7. Cronodrilus ogygius: a, holotype; b, lateral view of re-
productive systems; c, lateral view of male system viewed from side
opposite that shown in 7b to show ejaculatory duct; d, oblique view of
jaws, upper to the left.
type of prostate. Except for the common outlet of the anterior nephridia
and the unbranched spermatheca, presumably advanced characters,
Cronodrilus has the characteristics one would expect to find in the com-
mon ancestral stock of the genera with which it is compared.
Cronodrilus ogygius new species
Figures 7, 8
Type-specimens: Holotype and three paratypes, USNM 37103, three
paratypes, PCH 1346, from Cambarus latimanus (LeConte), taken in
a small stream, 3.1 miles north of Buchanan, Haralson County, Georgia,
on U.S. Highway 27, by Perry C. and Virgie F. Holt, 25 March 1961.
Diagnosis: As for genus.
Etymology: From Greek, ogygios, of or pertaining to Ogyges, leg-
endary king of Thebes, hence, primeval or ancient.
Description: Cronodrilus ogygius is a medium-sized worm of relatively
slender proportions. Four animals of the type series (USNM 37104)
New Branchiobdellid Worms 311
Ficure 8. Cronodrilus ogygius: a, longitudinal optical section of
bursal atrium and ectal end of penis; b, longitudinal optical section of
junction of ejaculatory duct and penial sheath, and ental end of penis.
have the following dimensions: total length, 2.7-3.3 mm; greatest diam-
eter (either segment VI or VII), 0.39-0.56 mm; head length, 0.45-0.49
mm; head diameter, 0.25-0.31 mm; diameter, segment 1, 0.26-0.31 mm;
diameter, sucker, 0.29-0.30 mm.
The head is of usual proportions. The lips are entire, but the margin
of the lower is slightly concave. There are no oral papillae. The external
furrow encircling the head is shallow. There is only one pharyngeal
sulcus. There are no supernumerary muscles in the prosomites of body
segments and, consequently, the body outline is smooth (Fig. 7a). The
clitellar glands of segment VI and VII are not prominent. The neph-
ridiopore is on the dorsum of segment III.
The jaws are of usual size and the dental formula is 5/4, but the
teeth are unusually large in proportion to the size of the jaws, sharply
pointed, and subequal in length; the median ones slightly the longer.
They are separated by deep grooves in the body of the jaws that extend
almost to its base (Fig. 7d).
The vasa deferentia enter the spermiducal gland about %4 of the total
length of the latter from its ental end (Fig. 7c). The spermiducal gland
is long and slender, about ¥ of its length in diameter. There is no
prostate or prostatic protuberance.
The ejaculatory duct is prominent (Fig. 7c), subequal to the sper-
miducal gland in diameter, about %4 its length, and set off by a slight
constriction at its junction with the penial sheath of the bursa.
312 Proceedings of the Biological Society of Washington
The bursa is distinctive. The penial sheath region is long, about %4
the diameter of the body in length, and very slender (Fig. 7b). The
wall of the penial sheath is composed of an outer longitudinal and an
inner circular layer, both presumably muscular. The cuticular lining
forms the eversible penis which lies free of the wall of the penial sheath,
at least at its ental and ectal ends, except for the attachments to the
atrium and ejaculatory duct (Fig. 8a, b), but along its midlength the
wall of the penis and that of the penial sheath are so close to each
other that their separateness cannot be determined. It is assumed, on
the basis of the structure of the penis and penial sheath, that the penis
is free, except at the ental and ectal ends, of its sheath and everts
freely to form a double-walled cuticular tube. The atrium and its outlet
canal make up about % of the total length of the bursa. It is not
certain that the usual inner atrial fold is present and the atrium is almost
surely noneversible.
The spermatheca (Fig. 7b) has a long spermathecal duct. The bulb
is of hardly greater diameter than the duct, but of the usual construction.
An ental process is only slightly set off by a shallow constriction from
the bulb and has a thick wall and narrow lumen. The exterior surface
of the ental process appears wrinkled in the holotype.
Variation: The only observable variation is in the spermatheca. It is
possible that the ental process is not a constant feature, but, rather is
only present when the spermathecal bulb is incompletely filled with
spermatozoa. Such processes do, however, seem to be constant features
in some species, but an ental process of the spermatheca is not detectable
in some of the specimens of the type series. Nonetheless, the congruence
in other characters among these specimens is such that there can be no
reasonable doubt as to their conspecificity.
Affinities: Since, at this time, Cronodrilus is a monotypic genus, the
affinities of C. ogygius are those of the genus as discussed above.
Host: Cambarus latimanus (LeConte).
Distribution: Known only from the type locality.
Material Examined: The type specimens listed above.
Remarks: It is noteworthy that one collection from the Piedmont of
Georgia should yield two such unusual branchiobdellids as Sathodrilus
megadenus and C. ogygius. Though it is to be expected that both species
will be found in other localities, it is likely that both are localized and
rare.
Tettodrilus new genus
Type-species: Tettodrilus friaufi new species, here designated.
Diagnosis: Branchiobdellid worms with two pairs of testes; unpaired
nephridiopore on dorsum of segment III; body terete, without peristomial
tentacles, or dorsal projections on body segments; spermiducal gland
slender, vasa deferentia enter entally; prostate non-differentiated, with
bulb, incompletely divided from spermiducal gland; bursa ovoid to
pyriform, penial sheath not externally demarkated from bursal atrium;
New Branchiobdellid Worms 313
short eversible penis enclosed by muscular tube projecting into atrial
lumen; ejaculatory duct present; spermatheca short, without ental process.
Etymology: From Greek, tetta, little father or daddy, and drilus,
worm, in reference to the phylogenetic significance of the genus. Mas-
culine.
Affinities: Tettodrilus is a primitive relative of the genus Cambarincola
and its specialized descendants, the species of the genus Pterodrilus.
Pterodrilus has been accorded monographic treatment (Holt, 1968b)
and its relationships with Cambarincola discussed. Herein it is only
necessary to compare Tettodrilus and Cambarincola and refer to the
still more primitive Sathodrilus, to the aberrant Ellisodrilus, and Oedi-
podrilus.
The distinctive features of Sathodrilus in this context are the small
and primitive spermiducal gland, the ental entry of the vasa deferentia
into the gland, the rudimentary prostatic protuberance in some species,
and the eversible penis. The spermiducal gland of Tettodrilus is slender
with prominent deferent lobes at its ental end, obviously a tube formed
by the union of the vasa deferentia, the epithelial wall of which has
become glandular and rather like that, except for the relatively lesser
diameter, of S. carolinensis. The prostatic protuberance of such species
as S. carolinensis has become a true prostate, but one that is still incom-
pletely divided from the spermiducal gland; that is, it does not empty
into the ejaculatory duct at the place where the latter arises from the
gland, but into the lumen of the spermiducal gland some distance entally
to the ectal end of the gland. The prostate is slender, undifferentiated,
but there is a prostatic bulb at its ental end that seems to be comparable
to the prostatic protuberance in species of Sathodrilus. The penes of
both Sathodrilus and Tettodrilus are eversible, but the two differ. The
penis of Tettodrilus, unlike that of Sathodrilus, is enclosed in a very
short penial sheath which is not separated from the bursal atrium by
an external constriction. The eversible cuticular lining, that is the penis
itself, is enclosed in a cylindrical, projecting (into the bursal atrium)
tube that may be homologous with the inner atrial fold of other genera
(see above, p. 302), but, more likely, is the evolutionary precursor of
the protrusible penis (Fig. 9b). The jaws are similar, though those of
Tettodrilus are more acutely triangular.
Tettodrilus resembles Ellisodrilus in the undivided and undifferentiated
prostate, but otherwise, in the absence of a spermatheca and presence
of a muscular, protrusible penis in Ellisodrilus, these genera are not
alike. Oedipodrilus, likewise, has the same type of prostate, but the
long, eversible penis of the latter is quite different from that of Tet-
todrilus.
The external (to the organs themselves) shape and arrangement of
the components of the reproductive systems of Tettodrilus and Cam-
barincola are remarkably similar. They differ in the following ways:
the spermiducal gland of Tettodrilus is a less compact, slenderer tube
than in any known species of Cambarincola; the prostate is incompletely
314 Proceedings of the Biological Society of Washington
divided from the spermiducal gland in Tettodrilus, but in species of
Cambarincola it empties in common with the spermiducal gland into
the ejaculatory duct; the bursa of Tettodrilus has the shape of that of
Cambarincola and the ejaculatory ducts of the two are alike, but the
penis of Tettodrilus retains its eversible character in contrast to the
protrusible, cone-shaped, muscular penis of Cambarincola (Holt, 1949,
p. 554). The jaws of the only known species of Tettodrilus and those
of many species of Cambarincola are essentially identical; and the 5/4
dental formula of T. friaufi is most likely the primitive dental formula
in the genus Cambarincola. Tettodrilus, in short, has the features ex-
pected in a primitive stock ancestral to the dominant genus Cambarincola
and its specialized derivatives, but deserves, however, generic status on
the basis of its primitive spermiducal gland and prostate and unusual
eversible penis.
Tettodrilus friaufi new species
Figure 9
Type-specimens: Holotype, USNM 37099, and one paratype, USNM
37100, from Orconectes rusticus mirus (Ortmann), O. rhoadesi Hobbs,
Cambarus striatus Hay, and C. tenebrosus Hay,? taken about 8.5 miles —
south of Lewisburg, Marshall County, Tennessee, in a small stream, on
U.S. Highway 431, by Perry C. and Virgie F. Holt, 18 April 1960;
one paratype, PCH 1007, from Cambarus striatus Hay and C. tenebrosus
Hay, taken in a stream tributary to the Harpeth River, 2.4 miles south
of Franklin, Williamson County, Tennessee on U.S. Highway 431, by
Perry C. and Virgie F. Holt, 18 April 1960; one paratype, PCH 1008,
from Cambarus striatus Hay, taken in a small stream, 5.3 miles south
of Franklin, Williamson County, Tennessee, on U.S. 431, by Perry C.
and Virgie F. Holt, 18 April 1960.
Diagnosis: As for genus.
Etymology: I am pleased to name this species for my friend, Dr.
James J. Friauf, Jr., who collected the first recognized specimens of it.
Description: Tettodrilus friaufi is a medium-sized worm of graceful
proportions. The specimens of the type series have the following dimen-
sions: total length, 2.2-2.7 mm; greatest diameter (segment VI), 0.40-
0.48 mm; head length, 0.41-0.49 mm; head diameter, 0.26-0.29 mm;
diameter, segment 1, 0.27-0.33 mm; diameter, sucker, 0.26—0.29 mm.
The upper lip bears four lobes. Oral papillae are present, 10-12 in
the holotype (they are difficult to count with certainty). There is only
one encircling furrow of the head other than that which bounds the
peristomium and it is very shallow. There is one pharyngeal sulcus.
Segments II and III have weak supernumerary muscles, but the dorsal
ridges of these segments are not prominent and the body outline is
smooth. The clitellum is detectable, but not prominent, on the dorsa
2 These specimens came from the sediments taken from the jar in which specimens
of all four species of crayfish were collected.
New Branchiobdellid Worms 315
Ficurt 9. Tettodrilus friaufi: a, lateral view of reproductive systems;
b, longitudinal optical section of bursa and penis; c, holotype; d, jaws,
upper jaw above.
of segments VI and VII. The nephridiopore is usually easy to see on
the dorsum of segment III.
The dental formula is 5/4. The median tooth of the upper jaw and
the paramedian ones of the lower jaw are large and sharp. The upper
jaw is noticeably larger than the lower. In color, both are a medium
shade of brown.
The vasa deferentia are expanded and become glandular to produce
deferent lobes of the spermiducal gland. The latter is about 1% the body
diameter in length, slender, and frequently slightly bent so that it
presents an irregular outline. It is subequal in diameter to the ejaculatory
duct at the junction of the two.
The prostate appears in whole mounts to be histologically identical
to the spermiducal gland, arises from the gland at about the border of
the ectal and median thirds of the latter and extends entally to the
junction of the deferent lobes. Its ental end contains a cavity, presum-
ably equivalent to the prostatic bulb of many species of Cambarincola
(Holt, 1949, p. 553; 1960, p. 63). In diameter, it is about 94 that of
the spermiducal gland.
The ejaculatory duct is subequal in length and diameter to the sper-
miducal gland and an inner layer of circular muscle is prominent. It is
set off from the penial sheath region of the bursa by a slight constriction.
The bursa is ovoid-pyriform. The ejaculatory duct passes almost
directly into the atrium as a muscular tube about 1% as long as the
bursa which encloses the eversible cuticular penis (Fig. 9b). This tube
may be eversible, though it need not be for the penis to be everted.
The outer end of the penis lies free within its enclosing tube and is a
double-walled cylinder of cuticle that is transversely folded several
times at the level of the junction of the ejaculatory duct and the bursa
(Fig. 9b) so that it is capable of further eversion. But in the everted
condition, it cannot be of any great length. The inner atrial fold is
poorly developed and not at all prominent.
There is little to note concerning the spermatheca. The spermathecal
duct is not unusually long, extending hardly farther than the ventral
316 Proceedings of the Biological Society of Washington
border of the gut, and is of the usual structure, that is, there is no
spermathecal bursa or other modifications of it. The bulb of the
spermatheca is obovate. The muscle layer of the bulb is unusually thick
and readily apparent at higher magnifications (400 ) in whole mounts.
There is no ental process.
Variation: There is no detectable variability, other than size, in the
available material.
Affinities: The affinities of the genus Tettodrilus have been discussed
above. Nothing further can be added: T. friaufi is not closely related
to any known species of either Sathodrilus or Cambarincola.
Hosts: Cambarus striatus Hay, C. tenebrosus Hay, Orconectes rhoadesi
Hobbs, and O. rusticus mirus (Ortmann). It is known from the field
data that T. friaufi certainly occurs on both C. striatus and C. tenebrosus,
but it is not certain that it occurs on the two species of Orconectes
named.
Distribution: Streams of the Nashville Basin in Middle Tennessee.
Other than localities mentioned above, T. friaufi has been taken from
Cambarus tenebrosus in Percy Warner Park, near Nashville, Davidson
County, Tennessee, by James J. Friauf, Jr., 22 March 1947.
Material Examined: In addition to the holotype and three paratypes, -
several poorly preserved specimens from Percy Warner Park, Davidson
County, Tennessee, PCH 161, have been examined.
In anticipation of future discussions of the phylogeny of the
branchiobdellids, the primitive nature and interesting distribu-
tion of the new genera and species described above deserve
brief comment here. The species of Sathodrilus, particularly
S. carolinensis, appear to be survivors of the stock that gave
rise on one hand to Ceratodrilus and on the other to Oedipod-
rilus. By way of S. veracruzicus, Sathodrilus appears to be
related to the relictual monotypic genus Magmatodrilus of
northern California. Cronodrilus may well be a remnant of
the stock that gave rise to all the branchiobdellids with the
spermiducal glands that are produced entally beyond their
junction with the vasa deferentia, including those of Asia
and Europe except Caridinophilia Liang, 1963. Tettodrilus
seems to be ideally suited to be an ancestor of Cambarincola
and the latter’s descendant genus Péterodrilus. As to their
distribution: these putatively primitive forms occur in an arc
around the southern end of the Appalachian Mountains and
disjunctively in Mexico. It is tempting to theorize that those
in the foothills of the Appalachians, remaining near their
original homes, are relicts of ancient groups that gave rise to
New Branchiobdellid Worms 317
the modern branchiobdellid fauna and that those in Mexico
are remnants of likewise ancient stocks that somehow at an
early time reached and have survived in the streams of the
eastern flanks of the Sierra Madre Oriental.
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 (2267):
241-265.
Hartt, Maurice C. 1914. Descriptions of a new genus and new
species of the discodrillid worms. Proc. U.S. Nat. Mus.,
48 (2071): 187-193.
HorFMAN, RicHArD L. 1963. A _ revision of the North American
annelid worms of the genus Cambarincola (Oligochaeta:
Branchiobdellidae). Proc. U.S. Nat. Mus., 114 (3470):
271-371.
Hott, Perry C. 1949. A comparative study of the reproductive
systems of Xironogiton instabilius instabilius (Moore) and
Cambarincola philadelphica (Leidy) (Annelida, Oligochaeta,
Branchiobdellidae). Jour. Morphol., 84: 535-571.
. 1953. Characters of systematic importance in the family
Branchiobdellidae (Oligochaeta). Va. Jour. Sci, N.S., 4
(2): 57-61.
1960a. The genus Ceratodrilus Hall (Branchiobdellidae,
Oligochaeta), with the description of a new species. Va.
Outre, St, We Ss Ill (Ae BSEYae
. 1960b. On a new genus of the family Branchiobdellidae
(Oligochaeta). Amer. Midl. Nat., 64 (1): 169-176.
. 1965. On Ankyrodrilus, a new genus of branchiobdellid
worms (Annelida). Va. Jour. Sci., N.S., 16 (1): 9-21.
. 1967a. Ocdipodrilus oedipus, new genus, new species (An-
nelida, Clitellata: Branchiobdellida). Trans. Amer. Microsc.
Soc., 86 (1): 58-60.
1967b. Status of genera Branchiobdellida and Stephanod-
rilus in North America with description of a new genus
(Clitellata: Branchiobdellida). Proc. U.S. Nat. Mus., 124
(3631): 1-10.
. 1968a. The Branchiobdellida: epizo6tic annelids. The Biol-
ogist, 50 (3-4): 79-94.
. 1968b. The genus Pterodrilus (Annelida: Branchiobdel-
lida). Proc. U.S. Nat. Mus. (in press).
318 Proceedings of the Biological Society of Washington
Lianc, YAN-Lin. 1963. Studies on the aquatic Oligochaeta of China.
1. Descriptions of new naids and branchiobdellids. Acta
Zool. Sinica, 15: 560-570.
Moores, J. Perry. 1895a. Pterodrilus, a remarkable discodrilid. Proc.
Acad. Nat. Sci. Philadelphia for 1894: 449-454.
1895b. The anatomy of Bdellodrilus illuminatus, an Amer-
ican discodrilid. Jour. Morphol., 10 (2): 497-541.
Opier, A. 1823. Mémoire sur le Branchiobdelle, nouveau genre
d’Annelides de la famille des Hirudinées. Mem. Soc. d’Hist.
Nat. de Paris, 1: 69-78.
PiERANTONI, UMBERTO. 1905. Cirrodrilus cirratus, n.g., n. sp. parassita
dell Astacus japonicus. Ann. del Mus. Zool. R. Univ. Napoli,
CNGS|) 2 CS) Ea 3:
STEPHENSON, J. 1930. The Oligochaeta. Oxford Univ. Press, Oxford.
YamMacucut, Hmeyr. 1934. Studies on Japanese Branchiobdellidae with
some revisions on the classification. Jour. Fac. Sci., Hokkaido
Imp. Univ., Ser. 6, 3 (3): 177-219, pls. 12-13.
HOGS
Vol. 81, pp. 319-322 30 August 1968
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
A REPLACEMENT NAME FOR BOTHROPS
LANSBERGII VENEZUELENSIS ROZE,
1959 (VIPERIDAE, SERPENTES)
By James A. PETERS
Smithsonian Institution, Washington, D.C.
Hoge (1966) published a list of “all recognized species” of
Bothrops, in which two distinct taxa carried the trivial name
“venezuelensis.” This resulted from the difficulty in determin-
ing which of the two was the earlier name. This problem has
now been resolved.
In 1952 Sandner described Bothrops venezuelensis from the
Serrania de El Avila, Federal District, Venezuela. The descrip-
tion (published in a Colombian journal which is extremely
difficult to obtain) currently is available in the United States
only in xerox copies. Apparently copies of this description
were not distributed widely enough to bring it to the attention
of other herpetologists until 1961. Certainly it was not seen
by Roze in 1959, when he described Bothrops lansbergii vene-
zuelensis from Caripito, Monagas, Venezuela, or he would not
have used the primary homonym. In 1961, Sandner and Romer
described Bothrops pifanoi, also from “Serrania de El Avila El
Papelon,” in a paper dated December 18, 1961, and Sandner
described Bothrops venezuelae, again from “Serrania de El
Avila,” in a paper dated December 20, 1961. These two papers
are mentioned in the review of Venezuelan snakes by Roze
(1966) in a footnote on p. 272, but as I have pointed out
(Peters, 1967), Roze did not have an opportunity to change
his text materially after about 1959. Sandner (1966) reviewed
the three nominal species venezuelensis Sandner, 1952; pifanoi
Sandner and Romer, 1961; and venezuelae Sandner, 1961. He
concluded that only one valid species is involved, and that it
34—Proc. Biou. Soc. WasH., Vou. 81, 1968 (319)
320 Proceedings of the Biological Society of Washington
should be known as Bothrops venezuelae Sandner, 1961. In
this review, he also attempted to justify using venezuelae as
a replacement name for venezuelensis, and to prove that vene-
zuelae and pifanoi were published simultaneously. He used
Article 33 of the International Code of Zoological Nomen-
clature as the basis for the change from venezuelensis to
venezuelae, arguing that these are identical names, having the
same stem, “venezuel—,” with the change in the suffix con-
sidered a “justified emendation” in the sense of Article 33.
The article in question, however, limits justified emendations
to the “correction of an incorrect original spelling.” Although
Sandner felt that his earlier name “venezuelensis” was unsatis-
factory, and wished to change it to “venezuelae,” it is clear
that the original name was correctly formed, and correctly
spelled, in accord with Recommendation IV of Appendix D
of the Code. Sandner’s proposed change must be rejected, and
the name of the taxon must stand as Bothrops venezuelensis .
Sandner, 1952, which seems to fulfill the criteria of the Code
for publication and availability, even though no library known
to me received copies before 1961. Following Sandner, 1966,
the names Bothrops pifanoi Sandner and Romer, 1961, and
Bothrops venezuelae Sandner, 1961, are synonyms of this
taxon.
Because the name Bothrops lansbergii venezuelensis Roze,
is a primary homonym of Bothrops venezuelensis Sandner,
I asked Dr. Roze to publish a new name for his subspecies.
He responded by asking me to review the whole problem.
I am pleased, therefore, to have an opportunity to continue
the association of his name with a subspecies from the country
whose reptile fauna he has studied so carefully, and I shall
call it:
Bothrops lansbergii rozei nomen novum
Bothrops lansbergii venezuelensis Roze, 1959, p. 11. Type-locality: Ca-
ripito, Monagas, Venezuela, 50 m. (Holotype: Amer. Mus. Nat. Hist.
59489.) Preoccupied by Bothrops venezuelensis Sandner Montilla,
1952.
Bothrops lansbergii venezuelensis—Hoge, 1965 (1966): p. 123.
Bothrops lansbergii venezuelensis—Roze, 1966: p. 278 (q.v. for earlier
synonymy ).
On Bothrops lansbergii venezuelensis 321
Distribution: Drier coastal areas of Venezuela from Monagas State
to Zulia State, according to Roze, 1966.
Still another nominal subspecies should be mentioned here.
In 1934, Bricefio described Bothrops Neuvoiedii Venezuelenzi,
based upon a series of specimens from the Rio de Oro region,
Venezuela. This taxon has been synonymized with Bothrops
atrox atrox by Roze, 1966. Both the specific and the sub-
specific name were obviously misspelled. The species was
meant to be “neuwiedi,” and for the subspecies the author
intended to use the adjective formed from the geographical
name plus the ending -ensis. Since, however, no corrected
spelling appeared in the original paper, and no errata were
published later, it seems legitimate to accept the spelling as
published. The name does not assume priority over the two
more recent uses of the epithet venezuelensis by Sandner or
Roze, because it differs in two letters, and the Code (1964,
Art. 57d) states that “. . . the difference of a single letter is
sufficient to prevent homonymy.” It can remain in the syn-
onymy of Bothrops atrox atrox, or be revived by a later author,
without effecting the other similarly spelled names.
LITERATURE CITED
BricENo Rossi, A. L. 1934. El problema del ofidismo en Venezuela.
Bol. Ministerio Salubr. Agric. Cria, Venezuela, 1 (14):
1079-1177, and 2 (15): 15-103.
Hocr, A. R. 1966. Preliminary account on Neotropical Crotalinae
[Serpentes Viperidae]. Mem. Inst. Butantan, 32, 1965
[1966]: 109-184.
Peters, J. A. 1967. On Venezuelan Snakes. Copeia, 1967 (2): 496—
498.
Roze, J. A. 1959. Taxonomic notes on a collection of Venezuelan
reptiles in the American Museum of Natural History. Amer.
Mus. Novitates No. 1934: 1-14.
. 1966. La taxonomia y zoogeografia de los ofidios de Vene-
zuela. Univ. Centr. Caracas.
SANDNER Montitia, F. 1952. Serpientes Bothrops de Venezuela.
Monog. Cien. Inst. Terapeut. Exp. Lab. “Veros,” Bogota,
Col., 9 (21): 5 unnumbered pages.
. 1961. Ofidios ponzofiosos de Venezuela—Bothrops vene-
zuelae. Sp. nov. Noved. Cient. Cont. Ocas. Mus. Hist. Nat.
La Salle, Caracas No. 30: 1-36.
. 1966. El caso de anomalia en Bothrops venezuelae. Publ.
Inst. Invest. Ofidiol. Venez., Caracas, No. 1: 1-16.
322 Proceedings of the Biological Society of Washington
AND M. Romer. 1961. Ofidios ponzofosos de Venezuela—
Bothrops pifanoi sp. nov. Noved. Cient. Cont. Ocas. Mus.
Hist. Nat. La Salle, Caracas No. 29: 1-15.
0678
Vol. 81, pp. 323-334 30 August 1968
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
PARAONIS PYGOENIGMATICA NEW SPECIES,
A NEW ANNELID FROM MASSACHUSETTS
(POLYCHAETA: PARAONIDAE)
By MerepitTH L. JONES =
Smithsonian Institution, Washington, D.C.
In his comprehensive study of the Paraonidae, in general,
and of Aricidea jeffreysii (McIntosh), in particular, Cerruti
(1909) synonymized Levensenia Mésnil, 1897, with Paraonis
Grube, 1873, and recognized the latter and Aricidea Webster,
1879, as the two valid genera of the family. In addition, he
reduced Cirrophorus Ehlers, 1908, to subgeneric status in the
genus Aricidea and proposed a new subgenus, Paraonides, for
the genus Paraonis.
This classification of the Paraonidae was followed for more
than fifty years, and was recognized by Fauvel (1927), E. and
C. Berkeley (1956), Hartman (1959), Day (1961), and Petti-
bone (1963). Hartman (1957), in addition to proposing a
third subgenus, Aedicira, for the genus Aricidea, redefined
Paraonis (Paraonides) Cerruti.
It remained for Day (1963) to raise the subgenera to generic
rank with the statement (p. 420) that “... as the whole family
includes some fifty species it is suggested that the subgenera
be given full generic rank.” Hartman (1965a), with some few
exceptions, concurred with Day as to the generic status of the
former subgenera and proposed two new genera, Aparaonis
and Paradoneis, the latter including Paraonides as earlier
redefined (Hartman, 1957); all seven genera were listed in
Hartman’s catalogue supplement (Hartman, 1965b). Laubier
(1965, p. 469), in his report of the presence of Cirrophorus
in the Mediterranean, followed Day in the recognition of
genera, with the comment that “Ce point de vue parait
35—Proc. Brow. Soc. WAsH., VoL. 81, 1968 (323)
324 Proceedings of the Biological Society of Washington
justifié . . . ,” and Glémarec (1966), in describing a new
species of Paradoneis from Brittany also accepted Day’s judg-
ment. Finally, Day (1967) continued the recognition of the
former subgenera as genera, with the exception of Paradoneis
Hartman which he returned to Paraonides Cerruti. Thus,
according to some authors there are presently seven, six, or
two genera in the family Paraonidae, and generic criteria are
the presence or absence of modified setae in neuropodia or
notopodia and/or the presence or absence of a median pro-
stomial antenna.
It was on the basis of the above that at the outset of the
present study, when a paraonid species was found which
possessed modified setae in both neuropodia and notopodia,
it was felt that this represented a new genus. However, a
survey of the literature has presented a number of questions
which have made this decision less straight-forward than
originally thought.
Paradoneis Hartman has been taken to include Paradoneis
lyra (Hartman, 1965) with modified setae comprised only of
lyrate notosetae while Paradoneis armata (Glémarec, 1966)
has modified setae represented by notopodial lyrate and
acicular aristate forms and Paraonis (Paraonides) harpagonea
(Stérch, 1967) has lyrate setae of two types, as well as biden-
tate hooks, as modified setal types. In addition, Paraonis
(Paraonides) neapolitana (Cerruti, 1909) is said to have
broadly limbate setae as modified setae; this would hardly
be considered “modified” in the same sense as lyrate or acic-
ular aristate setae.
Further, Paraonis platybranchia (Hartman, 1961) is said
to lack modified setae in either neuropodia or notopodia, and
would appear to be a species of Aparaonis (Hartman, 1965)
which, according to both a key to genera and the generic
diagnosis, has only “pointed” setae in neuropodia and noto-
podia. However, a figure of a parapodium of Aparaonis
reveals neuropodial acicular spines, also referred to in the
description, which may represent modified setae and, thus,
suggest that Aparaonis should be synonymized with Paraonis.
In considering paraonids with a medial prostomial antenna,
Aricidea (Aedicira) (Hartman, 1957) was said to have no
;
‘
.
4
i,
i
New Massachusetts Annelid 325
0.2 mm
FicureEs 1-3. Paraonis pygoenigmatica new species. 1, Dorsal view of
anterior region. 2, Lateral view of right side of anterior region. 3, Ven-
tral view of anterior region.
modified setae, although a figure of the type species ( Aricidea
pacifica Hartman, 1944, Fig. 9) suggests that a second type
of seta, perhaps “modified,” is present. Hartmann-Schroder
(1962) has described a species of Aricidea (Aedicira) which
possesses acicular (modified ?) neurosetae. Finally, there
would appear to be at least three types of “modified” setae
326 Proceedings of the Biological Society of Washington
0.2 mm
8
7
YZ ru
GY
Ze
= O
9 10
Ficures 4-11. Paraonis pygoenigmatica new species. 4, Right first
setiger, anterior view. 5, Right second setiger, posterior view. 6, Right
sixth setiger (first branchial), anterior view. 7, Left sixteenth setiger,
anterior view. 8, Left twenty-second setiger (last branchial), posterior
view. 9, Left twenty-third setiger (first postbranchial), anterior view.
10, Right forty-first setiger, anterier view. 11, Right fifty-ninth setiger
(fourth to last), anterior view. Lumen of gut is indicated in all figures;
setae with swellings near proximal ends are specialized; setae drawn with
T-shaped tips represent broken setae.
New Massachusetts Annelid 327
described for various species of Aricidea Webster (abruptly
attenuated, pseudoarticulate, acicular, and acicular with mu-
cronate tips) and for Cirrophorus Ehlers (acicular, with a
frail sheath, or acicular aristate).
In view of the discussion above it was considered prudent
to be taxonomically conservative in the present case and to
beg the question of the validity of genera and subgenera. For,
until a critical comparison and an evaluation of the taxonomic
value of the so-called modified setae can be carried out, I
prefer to consider that there are two genera, Paraonis and
Aricidea, which comprise the Paraonidae and that there is too
much confusion in the literature to allow the definition of
subgenera.
The present study was prompted by my attention being
drawn to a collection of paraonids taken in the course of the
Cape Cod Bay Census by personnel of the Systematics-
Ecology Program (= SEP), Marine Biological Laboratory,
Woods Hole, Massachusetts. I am indebted to these individ-
uals, Dr. David C. Grant, Resident Biclogist, and Dr. Mel-
bourne R. Carricker, Director, for allowing me the opportunity
of observing this material and for the gift of a part of their
material to the collections of the United States National
Museum. I am further grateful to Dr. Marian H. Pettibone
and Nancy M. Cramer, both of the Division of Worms, for
their criticisms and comments on this work.
Paraonis pygoenigmatica new species
Differential Diagnosis: Paraonis with limbate and non-limbate capil-
lary setae in the anterior region; with laneolate branchiae beginning on
the sixth setiger and extending to the 20th to 25th setiger; with setae
provided with a sub-basal tooth present in neuropodia and notopodia of
the posterior branchial region continuing to the posterior end of the body;
with postsetal notopodial lobes beginning on the sixth (= first branchial )
setiger; with no postsetal neuropodial lobes; pygidium with from three
to eight anal cirri.
Type Locality: The collection upon which the present research is
based is comprised of 61 specimens. The holotype (USNM 37457) and
paratypes (USNM 37458) were collected at SEP station 1910-E3 in
Cape Cod Bay (lat. 41° 48.7'N; long. 70° 04.6’ W), 17 May 1967, in
hard-packed sand, at a depth of approximately seven meters (50 speci-
mens). Other paratypes were collected at SEP stations 1930-E2 (lat.
328 Proceedings of the Biological Society of Washington
41° 50.0’ N; 70° 30.4’ W, 21 November 1967, eleven meters, nine speci-
mens), 1030-E (lat. 41° 49.0’ N; long. 70° 30.4’ W, 21 November 1967,
thirteen meters, one specimen), and 1608-E3 (lat. 41° 51.9’ N; long. 70°
01.8’ W, 6 September 1967, six meters, one specimen). Paratypes not
deposited in the collections of the U.S. National Museum have been
returned to the reference collection of the Systematics-Ecology Program,
Marine Biological Laboratory, Woods Hole.
Description: Complete representative specimens vary from 0.25 to
0.33 mm in diameter, at the widest part of the anterior region, and are
18 mm (for 77 setigerous segments, 16 mm (81 setigers), 13 mm (79
setigers), 12 mm (71 setigers), and 9 mm in length (for 62 setigers).
One ovigerous specimen, incomplete posteriorly, is 7 mm long (for 62
setigers) and 0.25 mm at its widest.
The generally brownish appearance of specimens preserved in alcohol
is due to the presence of small pigment granules, most dense on the
dorsal surface, and least dense on the ventral surface. There is a basic
segmental distribution of the pigment granules, in that there appear to
be few, if any, present in the intersegmental regions.
The prostomium is conical and poorly delimited from the peristomium
(Figs. 1-3). The anterior area of the prostomium appears to be set off
from the basal part by a deep transverse furrow. This buttonlike pro-
stomial tip appears to be retractile into a subapical pocket formed by the
adjoining surface of the prostomium; the margin of this pocket is smooth,
except for a pair of very small lateral notches (Fig. 2). A pair of darkly
pigmented eyespots are embedded ventrolaterally in the posterior region
of the prostomium. The peristomium is achaetous and is provided with
a pair of dorsolateral nuchal slits. As mentioned above, the peristomium
is not sharply set off from the prostomium; indeed, there is only a rather
faint ventrolateral segmental furrow separating the two. The mouth is
“T”-shaped and opens in the anterior third of the peristomium (Fig. 3);
a medial furrow runs posteriorly to the first setiger.
The prebranchial region is comprised of five setigerous segments and
is circular in cross-section (Figs. 4 and 5). There are no cirri or any
other parapodial appendages present in this region. The branchial region
extends from the 6th (Fig. 6) to the 20th to 24th setiger (Fig. 8); it is
flattened dorsoventrally anteriorly and grades to a more circular cross-
section posteriorly (Figs. 6-8). In addition to paired branchiae, each
setiger bears a pair of cirriform postsetal notopodial lobes. The post-
branchial region is comprised of from 40 to 50 setigers which are circular
to subcircular in cross-section (Figs. 9-11) and segmentally constricted
to give the posterior body a somewhat moniliform aspect; each setiger
bears a pair of postsetal notopodial lobes. The length to width ratio of
the prebranchial segments is approximately 1 : 2, that of the branchial
segments is approximately 1 : 4, and the ratio for the postbranchial seg-
ments varies from | : 3 in the region immediately following the branchial
area to approximately 1: 1 in the posterior portion.
The pygidium is rounded and bears a heavily ciliated, dorsally placed
New Massachusetts Annelid 329
me
SSS
—I S—,
SS
SS
0.05 mm ee SS
Ficures 12-15. Paraonis pygoenigmatica new species. 12, Dorso-
lateral view of pygidium and last four setigers of a specimen with usual
complement and disposition of three anal cirri. 13, Dorsal view of pygid-
ium and last four setigers of a specimen with supernumerary anal cirri.
14, Ventral view of pygidium of same. 15, View of two anal cirri show-
ing pattern of ciliation. Figs. 12-14, scale A; Fig. 15, scale B.
subterminal anal opening. In addition, there are usually three anal cirri,
a pair dorsolaterally and a single one ventromedially (Fig. 12). How-
ever, there may be as many as four pairs of anal cirri disposed over the
surface of the extreme posterior face of the pygidium (Figs. 13 and 14);
the most medial pair are ventrally placed and the most lateral are at
about the level of the anal opening. The tips of the anal cirri are pro-
vided with extensible, possibly articulated, apical processes (Fig. 15).
The distal portions of the anal cirri also bear tufts of cilia (Figs. 12-15)
which suggest the presence of sense organs.
The branchiae are lanceolate and of approximately the same size
(Figs. 1, 2, 6, and 7), except for the much smaller posterior-most pair
(Fig. 8). The branchiae are oriented in a parasagittal plane and each
bears a central medial ciliated tract. Both the medial and the lateral
surfaces of the branchiae have a barred appearance due either to parallel
wrinklings of the surface or to a well-ordered distribution of the cellular
elements comprising the tissue of the branchiae.
330 Proceedings of the Biological Society of Washington
Ficures 16-23. Paraonis pygoenigmatica new species. 16, Three
types of neurosetae of the first setiger. 17, Notoseta (type 1) of the
first setiger showing extent of granular surface of sheath-like structure.
18, Notoseta from second setiger. 19, Neuroseta from sixth setiger with
presumed separation indicated by heavy-line. 20, Neuroseta from sixth
setiger showing (a) basal separation between sheath or limbation and
axis, (b) hypothetical cross-section of separation area if there is an actual
separation, (c) hypothetical cross-section if the sheath or limbation is
rolled. 21, Specialized notoseta from twenty-second setiger. 22, Special-
ized neuroseta from forty-first setiger. 23, Specialized notoseta from
fifty-ninth setiger.
There is a rather extensive pattern of ciliation present in the anterior
region. The prostomial tip, as well as the main portion of the anterior
region of the prostomium, is strewn with a number of ciliated tufts
(Figs. 1 and 3) which probably represent sensory structures. In addition
New Massachusetts Annelid 331
there are dorsal, transverse ciliated bands present on the prostomium
and at the posterior margin of each of the prebranchial setigers (Figs. 1
and 3) and between the branchiae of the following region (Figs. 6 and
7). The peristomium bears a pair of oblique, lateral, ciliated bands
which are associated with the nuchal slits at their dorsal ends. The main
function of these bands, in conjunction with branchial cilia, is probably
concerned with the maintenance of respiratory water currents.
The setae are arranged in fascicles which are: not obviously palisaded
and which are located from the middle to the posterior third of a given
segment. The setae in both the neuropodia and notopodia of the first
setiger are of three kinds: (1) short capillary setae, broadly uni- or
bilimbate, which are disposed in an anterior series (Fig. 16, a), (2)
long, narrowly unilimbate capillary setae (Fig. 16, b) which, along with
(3) short non-limbate capillary setae (Fig. 16, c), are arranged in a
posterior series. It will be noted that types (1) and (2) have been
referred to as “limbate”; observations under oil-immersion suggest that
the structures which give the impression of being flattened, typical
limbations when examined with lower magnifications, may actually be
sheath-like structures, not unlike the hoods of some hooded hooks of
other families, e.g., the Spionidae and Magelonidae. Such has also been
observed to be the case when structures appearing to be limbate capillary
setae of certain spionid polychaetes were carefully observed with oil-
immersion optics. In addition, the limbation or sheath may present a
granular appearance (Fig. 17) suggestive of a finely spinous surface
(cf. Jones, 1962, Figures 73-76, Fauvel, 1927, Figures 12, m and n;
Figure 14, i, and Mésnil, 1896, Plate 7, Figure 12, Plate 9, Figures 7
and 13, et seq.).
Capillary setae of all three types mentioned above are present on all
of the remaining prebranchial setigers, although the setae of type (2)
may become less typical (Fig. 18) with a less readily distinguishable axis
and a less distinct axial termination, giving the impression of an extension
of the “limbation” beyond the setal axis (Figs. 17, et seq.).
All three types of capillary setae are present in the anterior portion
of the branchial region. However, there is a progressive decrease in the
number of the setae of type (1) along the length, until these are con-
fined to the neuropodial fascicle by the 16th setiger and are lacking
altogether by the last branchial setiger. In the branchial region there
may be a further modification of the setae of type (2). At times there
may be a discrete separation between the “limbation” and the axis (Fig.
19) and this may be so pronounced as to give the appearance of a slit
(Figs. 20a and b); it is difficult to ascertain the nature of this structure
with certainty and it may be that this is not in fact a slit but a space
between the axis of the setae and a rolled portion of the sheath or
limbation (Fig. 20c).
In the posterior branchial region there are no setae of type (1) and
setae of a fourth type are present. These appear to be derived from the
modified setae of type (2) (Figs. 19 and 20) and bear a “limbation” as
332 Proceedings of the Biological Society of Washington
well as a closely applied basal tooth (Figs. 21-23). More posteriorly
these toothed setae come to be the dominant setal type, until in the
pygidial region they are nearly the only type present in both neuropodia
and notopodia (Figs. 8-14). In addition to the increased numbers of
these setae toward the posterior region, there is also an increase in the
relative length of the basal tooth; in the last branchial setiger the ratio
of the tooth length to the total length is approximately 1 : 13, further
posterior, 1:8, and in the extreme posterior region, 1:4 or 1:5.
Shortly after the appearance of the first of these toothed setae in the
branchial region, they are to be found in both notopodia and neuropodia,
where they continue to be present to the end of the body (Figs. 9-14).
These are taken to represent the so-called “modified” or “specialized”
posterior setae of other paraonids.
Etymology: The specific epithet refers to the occurrence of pygidia
with supernumerary anal cirri; this condition is truly enigmatic.
LITERATURE CITED
BERKELEY, EDITH, AND Cyri. BERKELEY. 1956. Notes on Polychaeta
from the east coast of Vancouver Island and from adjacent
waters, with a description of a new species of Aricidea. Jour.
Fish. Res. Bd. Canada, 13: 541-546.
Cerruti, Atrizio. 1909. Contributo all’anatomia, biologia e sistem-
atica delle Paraonidae (Levinsenidae) con particolare ri-
guardo alle specie del golfo di Napoli. Mitt. Zool. Stat.
Neapel, 19: 459-512.
Day, Joun H. 1961. The polychaete fauna of South Africa. Part 6.
Sedentary species dredged off Cape coasts with a few new
records from the shore. Jour. Linn. Soc. London, 44: 463—
560.
1963. The polychaete fauna of South Africa. Part 8. New
species and records from grab samples and dredgings. Bull.
British Mus. (Nat. Hist.) Zool., 10: 381-445.
1967. A Monograph on the Polychaeta of Southern Africa.
Publication No. 656. Trustees of the British Museum (Nat.
Hist.), London, 878 pp.
FAUVEL, PreRRE. 1927. Polychétes sédentaires. Addenda aux errantes,
archiannélides, myzostomaires. In Faune de France, Paris,
Le Chevalier, 16: 1-494.
GiemMarec, MicHEL. 1966. Paraonidae de Bretagne. Description de
Paradoneis armata nov. sp. Vie et Milieu, Sér. A: Biol. mar.,
17: 1045-1052.
HarTMANn, Otca. 1957. Orbiniidae, Apistobranchidae, Paraonidae and
Longosomidae. Allan Hancock Pacif. Expeds., 15: 205-393.
1959. Catalogue of the polychaetous annelids of the world.
Allan Hancock Found. Publs., Occas. Paper, no. 23, pp. 1-
628.
1965a. Deep-water benthic polychaetous annelids off New
New Massachusetts Annelid 333
England to Bermuda and other North Atlantic areas. Allan
Hancock Found. Publs., Occas. Paper, no. 28, pp. 1-378.
1965b. Catalogue of the polychaetous annelids of the world.
Supplement 1960-1965 and index [= Supplement and index
to the catalogue of the polychaetous annelids of the world,
including additions and emendations since 1959]. Allan
Hancock Found. Publs., Occas. Paper, no. 23, pp. 1-197.
HARTMANN-SCHRODER, G. 1962. Zweiter Beitrag zur Polychaetenfauna
von Peru. Kieler Meeresforsch., 18: 109-147, 20 figs.
Jones, MerepirH L. 1962. On some polychaetous annelids from Ja-
maica, the West Indies. Bull. Amer. Mus. Nat. Hist., 124:
169-212.
LavusiER, Lucien. 1965. Sur la présence du genre Cirrophorus (Poly-
chétes, Paraonidae) en Méditerranée. Bull. Soc. Zool. France,
90: 469-477.
Mesnit, FeLrx. 1896. Etudes de morphologie externe chez les an-
nélides. I. Les spionidiens des cotes de la Manche. Bull.
Sci. France Belgique, 29: 110-287.
PETTIBONE, MARIAN H. 1963. Marine polychaete worms of the New
England region. I. Families Aphroditidae through Trocho-
chaetidae. U.S. Natl. Mus. Bull., 227: 1-356.
334 Proceedings of the Biological Society of Washington
74.0678
Vol. 81, pp. 335-346 30 August 1968
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
SOME NEW CHINESE MINNOWS
(PISCES, CYPRINIFORMES )
By Perru M. BaNnarescu AND Tropor ‘Fo NALBANT
Academy of the People’s Rumanian Republic, Bucharest
Through the kindness of Dr. Ernest A. Lachner, Division of
Fishes, U.S. National Museum, we received for study large
series of east Asian Cyprinidae in the collections of the
National Museum. Many of these specimens were mentioned
in recent revisional papers on some genera belonging to the
subfamilies Gobioinae and Cultrinae. Among the U.S. Na-
tional Museum collections were large series collected in 1961
by Kuntz and Wells on Taiwan Island. In these series, we
found representatives of a new species and a new subspecies
of Gobioinae. Another new species, belonging to the Cultrinae,
was discovered among USNM specimens collected by D. C.
Graham in Szechwan, upper Yangtze drainage. We also iden-
tified specimens in the collections of the Zoological Museum
in Hamburg as a Gobiobotia from Fukien, Southeast China.
This fish is closely related to the new species of Gobiobotia
from Taiwan.
We thank Dr. E. A. Lachner for loan of specimens and for
furnishing us information on specimens in the U.S. National
Museum and for copies of papers; Prof. Dr. C. Kosswig for
obtaining a financial grant for the senior author which allowed
him to travel and study in Germany, and Dr. Kosswig and
Dr. W. Ladiges for facilitating study of the cyprinids in the
collections of the Zoological Institute and Museum in Ham-
burg; and Dr. Bruce B. Collette, Bureau of Commercial
Fisheries Ichthyological Laboratory for providing photographs
and radiographs of specimens, and for other valuable assist-
ance.
36—Proc. Bion. Soc. WasH., Vou. 81, 1968 (335 )
336 Proceedings of the Biological Society of Washington
SUBFAMILY GOBIOINAE
Genus Gobiobotia Pappenheim
Until recently only 11 species of the highly specialized genus Gobio-
botia were known—eight from China and three from Korea. Most of
these have restricted ranges. A single one, G. pappenheimi, is widely
distributed from the Amur drainage to the Yangtze. Three new species
were described by Banarescu and Nalbant (1966a) from China (main-
land and adjacent islands). The species here described as new is the
14th within the genus.
Gobiobotia intermedia new species
Diagnosis: A species of Gobiobotia with 7 branched rays in the dorsal
fin; 37-39 scales in lateral line, 5-514 between lateral line and dorsal
fin origin, 244-3 between lateral line and pelvic origin; body rather
elongate; slender and low caudal peduncle; dorsal fin slightly in advance
of pelvic fins, rarely exactly above them; distance from pectoral to pelvic
origin greater than that from pelvic to anal origin; second ray of pectoral
elongate; first pair of mental barbels inserted either slightly in advance
or at the same level or even slightly behind the insertion of the maxillary
barbel; scales on back in front of dorsal fin slightly keeled; eyes large;
snout equal to or shorter than postorbital distance.
Affinities: This new species resembles Gobiobotia kolleri Banarescu
and Nalbant in body shape and color pattern, differing from it in having
the second pectoral ray elongate and the pelvic fins closer to the anal
fin than to the pectoral origin; on the other hand, it resembles G. kia-
tingensis Fang and G. ichangensis Fang in general body form and
elongate second pectoral ray, differing from them in number of scales
and from G. kiatingensis also in its larger eyes.
It is remarkable that among the specimens of G. intermedia intermedia
from Ping Tung Hsien, Taiwan, there are some in which the first pair
of mental barbels is inserted slightly behind the maxillary barbels, other
ones in which it is inserted at the same level; some on which the pelvics
are inserted exactly below, others in which they are inserted slightly
behind dorsal origin; the same variation in the position of pelvic fins
occur in Fukien specimens of G. intermedia fukiensis. Both characters
—position of the insertion of first mental barbel and position of pelvic
fins—were previously considered as constant species characters in the
genus of Gobiobotia.
Etymology: We chose the name intermedia because this new species
is intermediate between G. kolleri, G. naktongensis, G. kiatingensis, and
G. ichangensis.
Range: Taiwan Island and Minkiang drainage in Fukien Province,
southeast China.
Gobiobotia intermedia intermedia new subspecies
Figs. 1 and 3
Holotype: USNM 200245, one specimen, apparently immature, 48.0
New Chinese Minnows 337
mm SL, collected by Kuntz and Wells in an irrigation ditch near Ping
Tung, Ping Tung Hsien, Taiwan, 25 January 1962.
Paratypes: USNM 202593, five specimens, 41.0-51.0 mm SL.
Diagnosis: A subspecies of Gobiobotia intermedia with 37-38 scales
in the lateral line, body depth 14-18 percent SL, first mental pair of
barbels inserted at the same level or slightly behind insertion of maxillary
barbels.
Description: D 3/7; A 2/6; L. lat. 37 38.
2%-3
Maximum body depth 14.4—17.8 percent SL (x: 15.8); length of caudal
peduncle 20.7-24.2 percent (22.5); least depth of caudal peduncle 7.7—
9.7 percent (8.9). Dorsal profile only slightly convex, ventral profile
horizontal. Head wider than deep, its length 23.4-26.8 percent SL
(24.78); snout shorter than postorbital distance, its length 8.1-9.6 per-
cent SL (8.65) and 34.5-35.6 percent of head length (35.7); eye diam-
eter 5.5-6.9 percent SL (6.1), 22.4-25.8 percent of head length (24.6)
and 87.0-100.0 percent of interorbital width (94.0). Maxillary barbels
reaching under anterior fourth of eye; their length is 6.1-7.7 percent SL
and 25.6-29.1 percent of head length. First pair of mental barbels
inserted in some specimens slightly behind insertion of maxillary barbels,
in other specimens at the same level; they reach slightly beyond insertion
of second pair and behind vertical from anterior margin of eye; their
length is 3.1-3.4 percent SL, 14.3-16.3 percent of head length. Second
pair of mental barbels inserted behind vertical from anterior mar-
gin of eye, reaching far beyond insertion of third pair and almost
to vertical from posterior margin of eye; their length is 4.3-5.3 percent
SL and 17.9-20.0 percent of head length. Third pair inserted below
middle of eye, reaching to or almost to margin of preopercle; their length
is 5.5-8.0 percent SL and 22.4-28.4 percent of head length. Space
between mental barbels papillose.
Origin of dorsal about equidistant between tip of snout and middle of
caudal peduncle and between posterior margin of eye and last anal ray.
Origin of pelvic fins below or slightly behind that of dorsal, nearer anal
than pectoral origin; anal origin much nearer pelvic origin than caudal
base. Pectoral fins in some specimens not reaching, in others reaching
or even passing beyond pelvic origin; pelvic fins usually reaching or
almost reaching anal fin origin. Lobes of caudal equal or almost equal.
Predorsal distance 43.1—47.7 percent SL (x: 45.6); preanal distance 66.5—
71.0 percent (68.6); preventral distance 47.1-51.5 percent (48.5);
distance from pectoral to pelvic origin 24.4—27.8 percent (26.25); dis-
tance from pelvic to anal origin 20.8-25.2 percent (22.32); length of
pectoral fin 22.1-29.1; length of pelvic fin 19.7-22.4 percent; base of
dorsal fin 13.9-16.6 percent; height of dorsal 19.3-23.6 percent; base of
anal fin 9.8-10.8 percent; height of anal 15.4-19.5 percent. Second
pectoral ray elongate. Vent much closer to pelvic axil than to anal fin
origin. Dorsal scales in front of dorsal fin slightly keeled.
338 Proceedings of the Biological Society of Washington
Ficures 1-4. 1, Gobiobotia intermedia intermedia new subspecies
Ping Tung, Taiwan. Holotype (USNM 200245). 2, Gobiobotia inter-
media fukiensis new subspecies Fukien. Holotype (HZS 4044). 3,
Gobiobotia intermedia intermedia. Ventral view of mouth. 4, Gobiobotia
intermedia fukiensis. Ventral view of mouth.
Color Pattern: (Specimens in alcohol): light gray yellowish above,
yellow whitish below; scales above lateral line bordered with blackish.
A row of 7-8 round spots on sides along lateral line, in some specimens
more of less confluent in a longitudinal stripe. Upper side of head dark
gray. Minute, hardly distinct spots on dorsal and caudal fin; lower
fins unspotted.
New Chinese Minnows 339
Gobiobotia intermedia fukiensis new subspecies
Figs. 2 and 4
Holotype: Zoolo. Staat. Zool. Mus., Hamburg, 4044, one specimen,
70.0 mm SL, Minkiang drainage, Fukien Province, 19 October 1907,
obtained from Consul Siemssen.
Paratypes: ZSZM 4045 (5, 61.0-89.2) same locality and data.
Diagnosis: A subspecies of Gobiobotia intermedia with 37-39 scales
in the lateral line, body depth 20-23 percent SL, first pair of mental
barbels inserted slightly in advance of maxillary barbels.
5-54
Description: D 3/7; A 2/6; L. lat. 37 —
244-3
Body depth 20.4—-22.8 percent SL (x: 22.2); length of caudal peduncle
21.0-21.8 percent (21.4); least depth of caudal peduncle 8.2-9.1 per-
cent (8.65); head length 25.3-26.2 percent (25.9); snout slightly shorter
than or equal to postorbital distance, its length 9.6—-10.7 percent SL
(10.1) and 37.0-41.0 percent of head length (39.0). Maxillary barbels
reaching in most specimens under middle of eye. First pair of mental
barbels inserted slightly in advance of maxillary barbels, reaching beyond
insertion of second pair and almost to, or even slightly beyond vertical
from middle of eye; second pair reaching under or almost under posterior
margin of eye; third pair reaching under or even beyond margin of
preopercle. Length of maxillary barbels 4.5-6.5 percent SL and 23.4—
30.0 percent of head length; first pair of mental barbels 4.5-6.5 percent
SL and 17.3-25.2 percent of head length; second pair 7.4—9.4 percent
SL and 28.2-36.6 percent of head length; third pair 9.8-12.6 percent
SL and 37.6—45.0 percent of head length.
Origin of dorsal about equidistant between tip of snout and middle
of caudal peduncle and between anterior margin of eye and _ vertical
from posterior margin of anal fin; origin of pelvic fins exactly under
dorsal fin origin or slightly behind it. Second pectoral ray elongate.
Predorsal distance 43.4-47.0 percent SL (x: 45.2); preanal 69.5-72.5
percent (70.9); preventral distance 44.6-49.2 percent (46.2); distance
from pectoral to pelvic origin 23.0-27.6 percent (25.2); distance from
pelvic to anal origin 23.5-26.5 percent (25.2); length of pectoral fins
21.4-24.6 percent; length of pelvic fins 17.4—20.0 percent; base of dorsal
15.6-17.5 percent; height of dorsal 18.2-23.0 percent; base of anal 8.4—
8.9 percent; height of anal 14.9-17.7 percent. Vent much nearer pelvic
axil than anal origin. Dorsal scales in front of dorsal fin slightly keeled
(the specimens apparently not mature).
Color Pattern: As in the nominal subspecies, but lateral spots hardly
39.
distinct and not confluent.
Microphysogobio Mori
A revision of the species of Microphysogobio was recently published
by Banarescu and Nalbant (1966b), who recognized nine species, includ-
340 Proceedings of the Biological Society of Washington
ing the Taiwan Pseudogobio brevirostris Giinther; they considered Pseu-
dogobio fukiensis Nichols and three other Chinese forms hitherto ascribed
to Pseudogobio, as well as the Korean Microphysogobio koreensis Mori
as subspecies of M. brevirostris, while they ascribed a series of specimens
from Taichung, Taiwan, to the Upper Yangtze Pseudogobio obtusirostris
Wu and Wang, which they considered a species of Microphysogobio,
closely related to but specifically distinct from Microphysogobio brevi-
rostris. :
The examination of recent series from Taiwan leads us to conclusions
somewhat different from those expressed in the recent revision.
Two specimens, USNM 191286, 78.0 and 82.0 mm SL (Figs. 5, 7):
from a small stream in coastal plain at Hsin-chu Hsien, Chu-Tung agrec
with the lectotype and paratype of Pseudogobio brevirostris recorded by
Banarescu and Nalbant: they have 39-40 scales in the lateral line; 21%
between the lateral line and pelvic fin origin, depth 19.3-23.0 percent
SL; head 21.8-23.8 percent, eye 5.5-6.2 percent, 25.0-26.0 percent of
head length and 78.0—87.5 percent of interorbital width. In color pattern
these specimens differ rather sharply from fukiensis and the other Chinese
gudgeons included by Banarescu and Nalbant within M. brevirostris as
subspecies, but approach the specimens from Taichung, Taiwan, ascribed
by us to M. obtusirostris in having no lateral spots but a broad continuous >
dark stripe from eye almost to caudal base and an even more intensive
dark round spot on caudal base. The same stripe can be recognized in
the badly preserved paratype of brevirostris figured by Banarescu and
Nalbant (1966b, Pl. I, Fig. 1) who did not pay enough attention to
the color pattern. These two specimens belong thus to the nominal sub-
species Microphysogobio brevirostris brevirostris (Giinther ).
A series of specimens from western coastal plain of Taiwan, described
below as a new subspecies, have exactly the same color pattern but a
much deeper body (depth 21.7—28.0 percent) and 35-37 scales, while
those from Taichung, North Taiwan are intergrades, having depth 21.6—
24.1 percent SL, scales 37-39, rarely 40. It is our present opinion that
the Upper Yangtze Pseudobogio obtusirostris Wu and Wang is also a
subspecies of M. brevirostris, approaching the west and north Taiwan
populations of M. brevirostris in number of scales in the lateral line,
number of scales between lateral line and pelvic axil (3, as against 2—
2% in M. brevirostris brevirostris) and deep body. It seems, from the
figure by Wu and Wang, that P. obtusirostris also has a longitudinal
stripe, (yet slighter than in brevirostris ), the small, sparse blackish minute
spots mentioned by Wu and Wang in obtusirostris are present also in
brevirostris and even the black spot above the pectoral fin mentioned by
Wu and Wang as a characteristic of obtusirostris can be recognized in
a few Taiwan specimens of brevirostris.
An active process of subspeciation within the species Microphysogobio
brevirostris took place on Taiwan. We have no exact information on the
locality from which Gunther obtained the type-specimens (these are
labelled only “Formosa” ); the specimen from Tamsui River recorded by
New Chinese Minnows 341
Oshima (1919) belongs also to the nominal subspecies and those from
Hsin-chu Hsien; we could not locate these two localities on the map,
but presume they are in the eastern coastal plain. The new subspecies
M. brevirostris alticorpus occurs in the western coastal plain and the
intergrades in Taichung, north Taiwan. Because west Taiwan is opposite
the mainland and the upper Yangtze, M. brevirostris obtusirostris is
morphologically closer to the west Taiwan M. brevirostris alticorpus than
to the probably east Taiwan M. brevirostris brevirostris, one can assume
that the species arrived first in west Taiwan, when this island was still
connected to the mainland, spread northwards along the western plain,
then southwards along the eastern plain. During dispersal, the body
became less deep and the scales smaller. Further intensive collections,
from Taiwan rivers are necessary in order to clarify this problem.
Microphysogobio brevirostris alticorpus new subspecies
Figs. 6 and 8
Holotype: USNM 192926, one specimen, 63.0 mm SL, collected by
Kuntz and Wells in a small stream and roadside ditch a few miles away
from Chia-I-Hsien, western coastal plain of Taiwan Agriculture area;
March 1961.
Paratypes: USNM 202592 (66 specimens, 36.0-60.7) same locality
and collector.
Diagnosis: A subspecies of Microphysogobio, differing from M. brevi-
rostris brevirostris in its deeper body (depth 21.7-28.0 percent SL),
scales in lateral line 35-38, between lateral line and pelvic fin origin 3,
eye 4.9-6.3 percent SL and 52-73 percent of interorbital width.
5
Description: D 3/7; A 2/6; L. lat. 35 — 37 (38).
3
Body deeper than in other forms of Microphysogobio, maximum depth
21.7-28.0 percent SL (x: 25.32) in specimens 52-63 mm long; caudal
peduncle length 15.7-20.9 percent (17.94); least depth of caudal
peduncle 11.1-14.3 percent (12.98); head 20.1-23.2 percent (21.84);
snout blunt, its length 7.1-8.2 percent SL (7.59) and 32.5-39.4 percent
of head length (34.8); a notch in front of nostrils, eye high, supero-
lateral, smaller than in the nominal subspecies, its diameter 4.9-6.2 per-
cent SL (5.51), 24.2-27.0 percent of head length (25.23) and 52.5-73.0
percent of interorbital width (65.3). Barbel minute, about three times
eye diameter. Interorbital flat. Mouth small, inferior, horse-shoe
shaped; jaws covered by a horny sheath. Papillae on upper lip in one
row, distinct, the median pair only slightly larger than the other. Lower
lip with a pair of irregular mental pads (ovoid or more or less rectan-
gular), often confluent on most of their length, and hardly distinct
papillae in many rows on the sides on mental pads.
Origin of dorsal fins in most specimens, slightly nearer tip of snout
than vertical from last anal ray; origin of pelvic fins behind that of
342 Proceedings of the Biological Society of Washington
Ficures 5-9. 5, Microphysogobio brevirostris brevirostris (Gunther).
Hsin-chu Hsien, Chu-Tung, Taiwan (USNM 191286). 6, Microphyso-
gobio brevirostris alticorpus new subspecies. Chia-I, West Taiwan. Holo-
type (USNM 192926). 7, Microphysogobio brevirostris brevirostris.
Ventral view of mouth. 8, Microphysogobio brevirostris alticorpus. Ven-
tral view of mouth. 9, Ancherythroculter wangi (TCHANG). Suifu,
Szechwan (USNM 87457).
New Chinese Minnows 343
dorsal fin. Caudal forked; dorsal fin slightly concave; pectoral fins not
reaching pelvic axil, pelvic fins not reaching anal origin.
Predorsal distance 43.0-74.5 percent (73.0); preventral distance 49.0—
55.5 percent (52.9); distance from pectoral to pelvic origin 27.1-33.9
percent (30.66); distance from pelvic to anal origin 20.7—23.2 percent
(21.72); length of pectoral fins 24.1-26.4 percent (25.36); length of
pelvic fins 18.7-21.4 percent (20.09). Breast in front of pectoral origin
naked. Pharyngeal teeth in one row, usually 5 on each side.
Color Pattern: (Specimens in alcohol) light grayish above, yellowish
below; scales above lateral line bordered with blackish. An intensive,
1-1% scales broad dark stripe extends from opercle almost to caudal
base; a more intensive almost blackish spot on base of caudal. Upper
part of head dark. In some specimens a blackish spot present above
base of pectoral, in a few another spot present on opercle immediately
in front of pectoral fin. Several hardly distinct rows of spots on dorsal,
caudal and pectoral and, quite slightly, on pelvics and anal.
SUBFAMILY CULTRINAE
Ancherythroculter Wu
This genus was described by Wu in his recent (1964) volume on the
Cyprinidae of China, for three species: a new species described by him
as A. nigrocauda, Erythroculter wangi Tchang, and Chanodichthys kue-
matsui Kimura. This genus differs from Erythroculter in its bipartite
swim-bladder (as against tri-partite in Erythroculter).
A single specimen of the species included by Wu in Ancherythroculter
was available to us: A. wangi USNM 87457, Suifu, Szechwan, 112.0
mm SL (Fig. 9); this species approaches Erythroculter mongolicus in
general body and head shape, and differs from it in its bipartite swim-
bladder.
The delimitation of genera within the Cultrinae is rather arbitrary.
The species included within Ancherythroculter differed from Erythro-
culter mainly in the bipartite swim-bladder, also in having usually fewer
branched anal rays (most species of Ancherythroculter have no more
than 22 branched anal rays, but nigrocauda has 23-27, kurematsui 21-26;
Erythroculter usually has 23 or more, but E. ilishaeformis has 21-25, E.
mongolicus 18-22) and fewer scales (usually 50-65, but A. nigrocauda
has 66-70; most Erythroculter have more than 73 scales, but E. hypse-
lonotus has 60-66). The new species A. brevianalis also approaches
Hemiculter; in A. brevianalis, as well as in all other species of Anchery-
throculter and of Erythroculter, the lateral line is almost straight, while
in most species of Hemiculter it is bent abruptly downwards; but in
H. liui and, to a lesser extent, in H. bleekeri, the lateral line is almost
straight. Most species of Hemiculter have 11-14 branched anal rays
and 43-58 scales, but H. krempfi has 16-18 branched anal rays, H. dispar
up to 17 and H. liui 75 scales; the species of Ancherythroculter have
usually 21-27 branched anal rays and 57-70 scales, but A. nigrocauda
344 Proceedings of the Biological Society of Washington
Holotype (USNM 87462).
throculter brevianalis.
Suifu, Szechwan.
11, Scale from the caudal peduncle of Anchery
10, Ancherythroculter brevianalis Banarescu new species.
Ficures 10-11.
(Drawn by Mrs. M. Partatescu ).
New Chinese Minnows 345
has 23-27 branched anal rays, A. brevianalis 15-20 branched anal rays
and 50-60 scales. i
The scales on the caudal peduncle of both Ancherythroculter wangi
and A. brevianalis (Fig. 10), are somewhat intermediate, as to the
position of the focus, between those of Erythroculter (with an almost
central focus) and those of Hemiculter (with a more basal focus); the
basilateral angles of the scales of both Ancherythroculter are marked
much better than those of Hemiculter and Erythroculter.
It is remarkable that the range of all four species of Ancherythroculter
is restricted to the Upper Yangtze drainage.
Ancherythroculter brevianalis Banarescu new species
Figs. 10 and 11
Holotype: USNM 87462, one specimen, 92.0 mm SL, collected by
D. C. Graham at Suifu, Szechwan, October-November 1924.
Paratypes: USNM 202690 (2, 72.0-89.0), same data. USNM 86891
(1, 88.3), same locality and collector, 15.1II-15.1V. 1929. USNM 91620
(6, 51.0-77.5), same locality and data. USNM 87458 (1, 116.2 mm),
same locality, October-November 1924. USNM 91625 (6, 46.2-68.0),
same locality, 15.11I-15.IV. 1929.
Diagnosis: A species of Ancherythroculter with 16-20, rarely 15 or
21, branched anal rays, 50-60 lateral line scales and 12-14 gill rakers.
10-11
— 60;
3-4
Sp. br. 12-14; D. phar. 4.4.2-2.4.5; 4.4.1-1.4.4; 5.4.2-2.4.4 or 4.4.2-
2.4.4,
Body rather oblong and convex; body depth 22.0-29.2 percent SL
(x: 25.95; in holotype 26.1 percent); length of caudal peduncle 16.3-
20.0 percent (x: 17.7; in holotype 20.0 percent); least depth of caudal
peduncle 9.8-11.2 percent (x: 10.46; in holotype 10.9 percent); width
of body 37.0-51.0 percent of depth. Both profiles more or less similar,
dorsal one slightly more convex.
Head compressed, much deeper than wide; its length 25.0—27.4 per-
cent SL (x: 25.6; holotype 26.6); snout rather short and bluntly pointed,
its length 6.5-7.5 percent SL (x: 7.0; holotype 6.75) and 25.0-29.4
percent of head length (x: 26.9; holotype 25.2). Eye lateral, its diameter
6.5-8.8 percent SL (x: 7.8; holotype 7.6), 24.8-33.3 percent of head
length (x: 29.5; holotype 28.6 percent and 80.0-108.0 percent of inter-
orbital width (x: 95.2; holotype 87.5 percent). Mouth terminal, oblique,
its cleft reaching under nostrils, the insertion of mandible behind vertical
from anterior margin of eye but before the vertical from middle of eye.
A small symphyseal knob present and a corresponding excavation on
upper lip.
Origin of dorsal fin about equidistant between tip of snout and base
of caudal (in some specimens slightly nearer to tip of snout, in others
to base of caudal). Origin of pelvic fins before dorsal fin origin; origin
Description: D III/7; A 2 (15) 16-20 (21); L. lat. 50
346 Proceedings of the Biological Society of Washington
of anal fin slightly behind vertical from last dorsal ray but before the
vertical from tip of depressed dorsal. Tip of pectoral fins not reaching
pelvic fin origin in most large specimens, reaching it in some smaller
specimens. Pelvic fins reaching or almost reaching anal fin origin. Edge
of dorsal fin straight. The variation of number of branched anal rays
is irregular; among the 17 specimens available (holotype and 16 para-
types) there were: 15 branched rays (one specimen), one (the holo-
type) with 16, 17 (4), 18 (2), 19 (2), 20 46), and one with 21
branched rays (x: 18.5 + 0.41; = 1.68). Predorsal distance 48.0—52.0
percent SL (x: 49.5; holotype 51.0 percent); preanal distance 58.5-64.0
percent; preventral distance 42.5-46.0 percent; distance from pectoral
to pelvic origin 19.2-23.2 percent; distance from pelvic to anal origin
15.2-20.3 percent; length of pectorals 18.3-22.0 percent; length of
pelvics 15.2-18.4 percent; base of dorsal to 10.6-14.3 percent; length
of last dorsal spine 13.3-20.4 percent; length of first branched dorsal
ray 17.2-23.8 percent; base of anal 17.5-26.6 percent (holotype 17.6
percent); height of anal 11.8—15.7 percent.
A scaleless keel present between pelvic and anal fins. Lateral line
slightly arched from opercle to above posterior part of abdominal keel,
then slightly ascending; on caudal peduncle straight, equidistant from
ventral and dorsal side. Swim-bladder bipartite.
Color: (Specimens preserved in alcohol) yellowish, somewhat darker
above; a hardly distinct but broad longitudinal stripe from opercle to
caudal base. Fins unspotted. In life the animal is probably silvery.
LITERATURE CITED
Banarescu, P., anD T. NALBANT. 1966a. Notes on the genus Gobio-
botia (Pisces, Cyprinidae) with description of three new
species. Annot. Zool. Botan., Bratislava, 1-16.
1966b. Revision of the genus Microphysogobio (Pisces,
Cyprinidae). Vestnic Cs. Spol. Zool., Praha, (3): 194-209.
Osumia, M. 1919. Contributions to the study of freshwater fishes of
the Island of Formosa. Ann. Camegie Mus., (2-4): 169-328.
Wu, Hs. W. 1964. The cyprinid fishes of China. Vol. 1. Technical
Printing House, Shanghai, 228 pp., 78 pls. (in Chinese).
Wu, Hs. W., ann F. F. Wanc. 1931. On a collection of fishes from
the upper Yangtze Valley. Contr. Biol. Lab. Sci. Soc. China,
Nanking, zool. ser., (6): 221-237.
men Oo) 5
Vol. 81, pp. 347-350 30 August 1968
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
A NEW PSOLID SEA CUCUMBER
FROM THE VIRGIN ISLANDS
By Davin L. Pawson
Smithsonian Institution, Washington, D.C.
Holothurians of the family Psolidae have the dorsal surface
of the body invested in a continuous covering of imbricating
calcareous scales. The mouth and anus are dorsal, and the
ventral surface of the body (the sole) is soft, surrounded by
tube feet, by means of which the animal attaches itself to a
hard substrate. The family comprises about 100 species, and
in most the area between the oral and anal apertures is covered
by four or more scales. Within the genus Psolus, only P.
macrolepis Fisher from bathyal depths in the Hawaiian Islands
was known to possess two scales between the oral and anal
apertures. The new species described below shares this feature
with P. macrolepis, but is quite distinct in other respects, and
appears to have no known close relatives.
The type series of the new species was found in the collec-
tion of the Universitetets Zoologiske Museum, Copenhagen,
Denmark during the summer of 1965. I am grateful to Drs.
F. Jensenius Madsen and Bent Hansen of that institution for
their help during my stay there.
ORDER DENDROCHIROTIDA GRUBE, 1840
Famity PsoLtmaAr PERRIER, 1902
Psolus megaloplax new species
Material Examined: Three specimens from off Christiansted, St. Croix,
Virgin Islands; 90-360 meters; bottom type unknown, but specimens
attached to small rocks; collected by Dr. Th. Mortensen, 17 February
1906. Holotype: total length 17 mm, greatest width 13.5 mm, height
9mm. Paratypes: larger specimen length 21 mm, width 18.5 mm, height
at anterior end 6 mm; smaller specimen (juvenile) length 4.5 mm, width
4 mm. All specimens in the Universitetets Zoologiske Museum, Copen-
hagen.
37—Proc. Biot. Soc. WAsH., Vou. 81, 1968 (347)
348 Proceedings of the Biological Society of Washington
__0-Imm j
,O:2mm_,
Ficures 1-6. 1, Holotype of Psolus megaloplax, dorsal aspect (partly
diagrammatic; tentacles not shown); 2, oral area of larger paratype; 3,
anal area of larger paratype; 4, deposits from sole; 5, perforated plates
from tentacles; 6, curved rods from tentacles.
New Sea Cucumber 349
Diagnosis: Two or three scales between oral and anal apertures. Mar-
ginal fringe of small dorsal scales absent. Oral aperture with five radial
and five interradial valves. Deposits of sole four-holed buttons of average
length 0.10 mm, with low rounded marginal knobs.
Description: Holotype and juvenile with tentacles contracted; larger
paratype with expanded tentacles. Body approximately circular, dorsal
surface slightly arched; sole contorted to closely fit hard substrate. Mouth
and anus dorsal. Mouth covered by five large triangular oral interradial
valves, below and between which are five radial valves, which project
into middle of oral field. Anal aperture covered by several valves, vary-
ing in number and size. Elsewhere, dorsal surface covered by large
scales up to 5.0 mm in diameter, which extend to edge of dorsal surface,
so that marginal fringe of smaller scales is absent; 2-3 scales between
oral and anal apertures (Fig. 1). Scales scarcely overlapping; edges of
adjacent scales held together by leathery skin. All scales covered by
fine pattern of ridges, scarcely visible to naked eye; all with covering of
evenly scattered, firmly fixed granules. Granules do not occur at extreme
edges of plates. Tentacles richly branched, all apparently subequal in
size.
Sole more or less circular, thin, translucent, surrounded by 2-3 rows
of tube feet. Midventral radius naked, except at extreme anterior (five
tube feet) and posterior (four tube feet) ends. Color in alcohol light
brown, distinctly lighter at sutures between plates; tentacles light yellow.
Juvenile specimen similar to adults, but dorsal scales smooth, lacking
granules.
Calcareous deposits of sole closely scattered four-holed buttons (Fig.
4) of average length 0.10 mm, with low rounded marginal knobs. Some
plates with more or less than four holes, and in other margin may be
tumid rather than actually knobbed. Tentacles with perforated plates
(Fig. 5) of varying shape, averaging 0.13 mm in length, and large curved
rods (Fig. 6) up to 0.50 mm long, many of which have a single perfora-
tion at each end.
Remarks: It is unlikely that this species could be confused with any
other Caribbean psolids. Of those species in the Caribbean area which
have five distinct interradial oral valves, only P. megaloplax has less than
four scales between the oral and anal apertures. P. megaloplax differs
from the Hawaiian species P. macrolepis in lacking a marginal fringe of
smaller scales from the dorsal surface, in having fewer perforations in
the deposits of the sole, and in having rods of a different type in the
tentacles.
LITERATURE CITED
Fisuer, W. K. 1907. The holothurians of the Hawaiian Islands. Proc.
U.S. Nat. Mus., 32: 637-744, pls. 66-82.
350 Proceedings of the Biological Society of Washington
ioe /3
Vol. 81, pp. 351-366 30 August 1968
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
MONOGENETIC TREMATODES FROM THE
SOUTHERN PACIFIC OCEAN, PART IV.
POLYOPISTHOCOTYLEIDS FROM-NEW ZEALAND
FISHES: THE FAMILIES MAZOCRAEIDAE,
DICLIDOPHORIDAE AND HEXABOTHRIIDAE?
By Wit1am ArtHuR DILLON? AND WILLIAM J. Harcis, Jr.
Virginia Institute of Marine Science,
Gloucester Point, Virginia
This is the fourth paper of a series on monogenetic trema-
todes of fishes of the southern Pacific Ocean. This installment
deals specifically with several species belonging to the families
Mazocraeidae Price, 1936, Diclidophoridae Cerfontaine, 1895,
and Hexabothriidae Price, 1942 of New Zealand fishes. The
scope, organization, and purpose are the same as for the first
installment.
Materials and Methods: Methods used in the preservation and the
preparation of the monogenetic flukes for identification and study are
essentially the same as those given by Dillon and Hargis (1965).
All measurements were made with the use of a filar micrometer and
are given in microns unless otherwise noted. In indicating these measure-
ments the mean is given, followed by the range (minimum and maxi-
mum.) in parentheses. The number of measurements used in the calcula-
tions appears in parentheses before these data. Measurements of curved
structures are across the lines subtending the greatest arcs described by
those structures.
In the measurements to follow, length—of the body, its appendages
and most internal organs—refers to the distance along the anteroposterior
axis except where otherwise noted. Width refers to a measurement made
1 This entire research is supported by grants GA-13853, with amendments, and
GA-235 under the United States Antarctic Research Program of the National Science
Foundation.
Contribution No. 276 from the Virginia Institute of Marine Science, Gloucester
Point, Virginia 23062.
2 Present address: University of Southern Mississippi, Southern Station, Box 1631,
Hattiesburg, Mississippi 39401.
38—Proc. Biou. Soc. WaAsH., Vou. 81, 1968 (351)
352 Proceedings of the Biological Society of Washington
at right angles to the length, i.e., along the dextrosinistral axis. The
lengths of cirri, accessory pieces, genital ducts, anchors, hooks, and spines
are along the longest axes of those structures regardless of orientation.
Clamps, because of their differing shapes and variable orientation in
relation to the anteroposterior axis of the body, present special problems.
In this installment, the length of clamps is regarded as the greatest
dimension of the sclerotized framework; the width is taken as the greatest
dimension at right angles to the length.
Camera lucida drawings were used to facilitate identification and in
preparation of the plates.
SUBORDER POLYOPISTHOCOTYLEA ODHNER, 1912
SUPERFAMILY DICLIDOPHOROIDEA Price, 1936
FAMILY MAZOCRAEIDAE PRICE, 1936
Neogrubeinae new subfamily
Diagnosis: Mazocraeidae. Body elongate, merging inconspicuously
with posthaptor. Prohaptor a pair of buccal suckers placed laterally in
walls of buccal funnel. Posthaptor armed with four pairs of clamps and
two pairs of anchors; clamps similar in framework, unequal in size;
clamps on one side of body (usually right side) open, large, with center
piece of clamp parallel to dextrosinistral axis; clamps on other side of
body smaller, closed, with center piece of clamp parallel to antero-
posterior axis and at right angles to the large, open clamps. Clamps of
a modified mazocraeid-type; anchors Kuhnia-like, dissimilar in size.
Testes numerous, pre-, para-, and postovarian, extending into posthaptor.
Ovary tubular, folded, with mature end directed anteriorly. Vagina
apparently absent. Genital atrium armed as in Kuhnia. Eggs with fila-
ments at both poles.
Type Genus: Neogrubea new genus.
Discussion: Of the six subfamilies (Clupeocotylinae, Grubeinae, Kuhni-
inae, Mazocraeinae, Mazocraeoidinae, and Neomazocraeinae) described
by Price (1961), Neogrubeinae most closely resembles, as the name
suggests, Grubeinae—especially in the following characters: (1) large,
open modified mazocraeid-type clamps on one side, (2) much smaller,
closed clamps on the opposite side, (3) clamp framework, (4) two pairs
of dissimilar anchors, (5) Kuhnia-type genital atrium, and (6) eggs with
filaments at both ends.
Neogrubeinae differs from Grubeinae as follows: (1) posthaptor with
four pairs of clamps rather than four on one side and one on the other,
(2) testes numerous, pre-, para-, and postovarian, extending into post-
haptor, (3) vagina apparently absent, and (4) mature end of ovary
directed anteriorly.
Although trematodes were recovered from Seriolella porosa Guichenot
and S. brama (Gunther), the description herein is based only on that
population from S. porosa since the specimens from S. brama were rela-
tively immature and in poor condition. Comparison between the two
Southern Pacific Trematodes Pt. IV 303
populations indicated that they were similar morphologically but differed
in the size of most of the significant characters. A close study of good,
mature specimens from S. brama should be undertaken in order to deter-
mine if the two populations under discussion are definitely conspecific.
Neogrubea new genus
Diagnosis: Neogrubeinae. With characters of the subfamily.
Type Species: Neogrubea seriolellae new species.
Neogrubea seriolellae new genus, new species
Figs. 1-6
Hosts: Seriolella porosa Guichenot (type host), Silver Warehou, and
S. brama (Gunther), Warehou; family Stromateidae.
Location: Gills.
Localities: (1) Cape Campbell, Marlborough Province, South Island;
10 statute miles NE of Cape Campbell (60 fathoms; hard mud), and
(2) Akaroa, Canterbury Province, South Island; Akaroa Harbour (4-10
fathoms; mud-—sand—mussels ).
Number Studied: 50.
Holotype: USNM Helm. Coll. No. 71191.
Paratypes: USNM Helm. Coll. No. 71192 (5 specimens).
Description: Body elongate, (8) 8,520 (6,800-11,000) long by (8)
1,690 (1,310-1,970) wide; sides tapered gently anteriorly; widened
posteriorly to merge inconspicuously with posthaptor. Prohaptor a pair
of buccal suckers (6) 97 (90-105) long by (6) 85 (80-94) wide, placed
laterally in walls of buccal funnel. Posthaptor bearing four pairs of
clamps and two pairs of anchors on a small terminal lappet. Clamps
similar in framework, unequal in size; clamps on one side of body
(usually the right side) open, large, (19) 816 (493-1,187) long by (19)
717 (501-976) wide, with center piece of clamp parallel to dextro-
sinistral axis; clamps on other side of body smaller, closed, (17) 531
(429-615) long by (17) 392 (324-350) wide, with center piece of
clamp parallel to anteroposterior axis and at right angles to the large,
open clamps. (The side with the larger clamps, right or left, varies
individually, but the internal organs maintain a constant orientation
regardless of this variance. When the parasite is attached to a gill
filament of the host, the larger, open clamps appear to overlap the ex-
posed margin of the filament while the smaller, closed clamps are tightly
closed on the lamellae of the filament.) Clamp structure of modified
mazocraeid-type (Figs. 4-5). Anchors (Fig. 6) dissimilar in size; outer
anchors (8) 65 (53-73) long; inner anchors (7) 53 (48-62) long.
Mouth subterminal, ventral. Pharynx (5) 77 (73-84) long by (5)
70 (62-79) wide; esophagus long, with lateral branches. Gut bifurcated;
crura ramified medially and laterally, extending into posthaptor.
Testes numerous, follicular, pre-, para-, and postovarian; testes extend-
ing into posthaptor. Vas deferens tightly coiled posteriorly, loosely coiled
354 Proceedings of the Biological Society of Washington
MEY,S a 6
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Ficures 1-6. Neogrubea seriolellae n. gen., n. sp. 1, Whole mount,
ventral view. 2, Genital corona. 3, Egg. 4, Large, open clamp; ventral
view. 5, Small, closed clamp; ventral view. 6, Anchors.
anteriorly, running anteriorly in midline to genital atrium. Genital corona
consisting of a central, ring-shaped muscular piece (5) 97 (88-107)
long by (5) 87 (82-91) wide, armed centrally with 10-14 spines (6)
18 (16-19) long, and a pair of curved muscular pieces anterolateral to
muscular ring, each with one large antlerlike spine (5) 35 (32-38) long.
Southern Pacific Trematodes Pt. IV 305
Ovary tubular, folded, with mature end directed anteriorly; oviduct
proceeding anteriorly, joining with genito-intestinal canal from right crus
and duct from vitelline reservoir before entering ootype. Ootype dorsal
to vitelline reservoir, surrounded by Mehlis’ glands; uterus ventral, run-
ning anteriorly in midline to genital atrium. Vagina apparently absent.
Vitellaria follicular, extending from level near gut bifurcation to end of
posthaptor; transverse vitelloducts fusing medially to form vitelline
reservoir. Eggs somewhat fusiform, with filaments at both ends; eggs
(2) 349 (340-357) long (measurement exclusive of filaments). Ex-
cretory tubules observed on right and left side near level of genital
atrium. Brain situated posterodorsally to pharynx.
FAMILY DICLIDOPHORIDAE CERFONTAINE, 1895
SUBFAMILY EURYSORCHIINAE YAMAGUTI, 1963
GENUS EURYSORCHIS MANTER AND WALLING, 1958
Eurysorchis australis Manter and Walling, 1958
Host: Seriolella brama (Gunther), Warehou and S. porosa Guichenot,
Silver Warehou; family Stromateidae.
Location: Gills.
Localities: (1) Cape Campbell, Marlborough Province, South Island;
10 statute miles NE of Cape Campbell (60 fathoms; hard mud), and
(2) Akaroa, Canterbury Province, South Island; Akaroa Harbour (4-10
fathoms; mud—sand—mussels ).
Previously Reported Host and Locality: Seriolella brama (Gunther)
from Wellington, New Zealand (Manter and Walling, 1958).
Number Studied: 20.
Homoeotypes: USNM Helm. Coll. No. 71193 (4 specimens).
Discussion: Manter and Walling (1958) described Eurysorchis aus-
tralis from the gills of Seriolella brama collected at Wellington, New
Zealand. In the present study this species was recovered from the gills
of S. brama and S. porosa, the latter being a new host record for this
species.
SUBFAMILY DICLIDOPHORINAE CERFONTAINE, 1895
Genus DICLIDOPHORA DieEsinc, 1850
Diclidophora coelorhynchi Robinson, 1961
Host: Coelorhynchus australis (Richardson), Rat Fish; family Macrou-
ridae.
Location: Gills.
Locality: Cape Campbell, Marlborough Province, South Island; 10
statute miles NE of Cape Campbell (60 fathoms; hard mud).
Previously Reported Host and Locality: Coelorhynchus australis
(Richardson) from Cook Strait, New Zealand (Robinson, 1961).
Number Studied: 43.
Discussion: A study of the original description established the con-
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Southern Pacific Trematodes Pt. IV 307
specificity of Robinson’s (1961) worms with the population in the present
collection.
SUPERFAMILY POLYSTOMATOIDEA PRICE, 1936
FamMity HexABOTHRUDAE Price, 1942
Genus Hexabothrium Nordmann, 1840
Hexabothrium akaroensis new species
Figs. 7-13
Host: Galeorhinus australis Macleay, Southern Tope or School Shark;
family Carcharhinidae.
Location: Gills.
Locality: Akaroa, Canterbury Province, South Island; Akaroa Harbour
(4-10 fathoms; mud-sand—mussels ).
Number Studied: 5.
Holotype: USNM Helm. Coll. No. 71194.
Paratype: USNM Helm. Coll. No. 71195.
Description: Body elongate, (4) 4,600 (4,330-5,100) long (measure-
ment not including appendage) by (2) 915 (840-990) wide. Cuticle
relatively thick; sometimes undulant. Prohaptor an oral sucker, (4) 272
(246-299) long by (4) 370 (295-467) wide; sclerotized papillae on
rim and inner surface of oral sucker; preoral hood or membrane mod-
erately developed. Posthaptor (4) 1,190 (1,120-1,260) long by (4)
810 (730-980) wide, armed with three pairs of sclerotized suckers, with
each sucker possessing a large hooklike sclerite; rims of suckers with
minute, conical, sclerotized papillae; striae in lumen of sucker. Hooklike
sclerites typically hexabothriid in shape nearly equal in size; anteriormost
sclerite (opposite to appendage) (4) 316 (293-339) long; middle sclerite
(4) 358 (352-364) long; posteriormost sclerite (adjacent to appendage)
(3) 313 (293-327) long. Hooklike sclerites with spines along their
lateral margins. Appendage (4) 809 (573-1,013) long, armed with two
anchors and two barrel-shaped suckers; anchors (4) 67 (66-71) long;
suckers (4) 261 (236-286) long.
Mouth slightly subterminal, ventral. Pharynx (2) 107 (99-115) long
by (2) 82 wide; esophagus short. Gut bifurcated, with medial and
lateral branches; crura confluent posteriorly prior to entering posthaptor
and appendix; not branched in posthaptor or appendix.
Testes numerous, postovarian; number of testes not discernible. Vas
deferens loosely coiled anteriorly, tightly coiled posteriorly, extending
anteriorly in midline to proximal end of cirrus. Muscular cirrus (4) 171
(163-180) long; distal end of cirrus armed with a circle of minute,
curved spines (number of spines not clearly discernible); prostate glands
emptying into base of cirrus. Genital atrium (4) 63 (55-72) long by
(4) 64 (57-66) wide, situated (4) 462 (406-506) from anterior end.
Uterus usually distended, containing numerous eggs; some oncho-
miracidia free in uterus, suggesting ovoviviparity. Seminal receptacle
present. Vagina double, opening just lateral to crura, slightly posterior
358 Proceedings of the Biological Society of Washington
2i
0.2
23
0.2
24
0.08
Ficures 14-25. Erpocotyle squali (MacCallum, 1931) Price, 1942.
14, Whole mount, ventral view. 15, Anteriormost sclerite. 16, Middle
sclerite. 17, Posteriormost sclerite. 18, Egg. 19, Anchor. Erpocotyle
callorhynchi (Manter, 1955) Yamaguti, 1963. 20, Whole mount, ventral
view. 21, Anteriormost sclerite. 22, Middle sclerite. 23, Posteriormost
sclerite. 24, Anchors. 25, Eggs.
Southern Pacific Trematodes Pt. IV 309
to level of genital atrium. Vitellaria follicular, co-extensive with intestine,
extending from level near base of cirrus to confluent crura; vitellaria
apparently not extending into posthaptor or appendage. Eggs (4) 124
(113-134) long by (4) 54 (52-57) wide, with filament at one pole;
filament almost twice as long as egg.
Discussion: Hexabothrium akaroensis n. sp. differs from H. appendic-
ulatum (Kuhn, 1829) Nordmann, 1840, the type species, as follows:
(1) smaller body size, (2) larger anchors, (3) larger hooklike sclerites,
and (4) host. (Anchor and sclerite size of H. appendiculatum was
extrapolated from the figures of Cerfontaine, 1899.) H. akaroensis n. sp.
differs from the other species in the genus; H. canicula (Cerfontaine,
1899) Price, 1942 and H. musteli (MacCallum, 1931) Price, 1942, as
follows: (1) eggs smaller, (2) anchors larger, (3) hooklike sclerites
larger, and (4) host.
Genus Erpocotyle van Ben. and Hesse, 1863
Erpocotyle antarctica (Hughes, 1928) Price, 1942
Figs. 26-31
Synonym: Squalonchocotyle antarctica Hughes, 1928.
Host: Mustelus antarcticus Guenther, Gummy Shark; family Carcha-
rhinidae.
Location: Gills.
Localities: (1) Akaroa, Canterbury Province, South Island; Akaroa
Harbour (4-10 fathoms; mud-sand—mussels), (2) Timaru, Canterbury
Province, South Island; 10 statute miles ENE of Timaru (9 fathoms;
sand) and (3) Timaru, Canterbury Province, South Island; 23 statute
miles ENE of Timaru (16 fathoms; sand—mud).
Previously Reported Host and Localities: Mustelus antarcticus from
Port Phillip Bay, Victoria, Australia (Hughes, 1928) and from New
Zealand (Manter, 1955).
Number Studied: 50.
Homoeotypes: USNM Helm. Coll. No. 71196 (5 specimens).
Description: Body elongate, (20) 10,600 (8,500-13,200) long (mea-
surement not including appendage) by (20) 1,150 (910-1,460) wide.
Cuticle thick, crenulated; surface of body, especially anterior portion,
covered with fleshlike papillae. Prohaptor an oral sucker, (20) 444
(390-573) long by (20) 591 (500-819) wide; oral sucker strongly
muscular, with rims bearing sclerotized papillae; inner surface of sucker
with tubercles. Preoral hood or membrane well developed, armed with
sclerotized papillae. Posthaptor armed with three pairs of sclerotized
suckers, with each sucker possessing a large hooklike sclerite; conical
papillae not observed on rims of suckers; striae in lumen. Hooklike
sclerites typically hexabothriid in shape, dissimilar in size; anteriormost
sclerite (opposite the appendage), (20) 544 (442-789) long; middle
sclerite (20) 654 (535-910) long; posteriormost sclerite (adjacent to the
appendage), (20) 576 (485-812) long. Hooklike sclerites bearing spines
360 Proceedings of the Biological Society of Washington
along their lateral margins. Disklike sclerites located in posthaptor.
Appendage (20) 1,110 (900-1,630) long, armed with two anchors and
two barrel-shaped suckers; anchors (20) 54 (51-60) long; suckers (10)
286 (213-380) long by (10) 200 (150-273) wide.
Mouth subterminal, ventral. Pharynx (10) 162 (147-171) long by
(10) 125 (99-147) wide; esophagus short. Gut bifurcated, with medial
and lateral branches; crura confluent posteriorly, prior to entering post-
haptor and appendix; unbranched in either.
Testes numerous, approximately 91-100, postovarian, between intes-
tinal crura. Vas deferens sinuous, broadened posteriorly, narrowed
anteriorly, extending anteriorly in midline to proximal end of cirrus;
distal end of vas deferens expanded. Cirrus muscular, (10) 500 (364-—
642) long; base of cirrus surrounded by prostate glands. Genital atrium
(10) 217 (182-257) long by (10) 205 (164-267) wide, receiving uterus
(ventrally) and cirrus (dorsally); genital atrium situated (10) 866
(740-1,044) from anterior end.
Ovary irregularly multilobed or branched; oviduct extending anteriorly
from mature end of ovary, fusing with ducts from vitelline reservoir, and
genito-intestinal canal before entering ootype. Uterus extending anteriorly
in midline to genital atrium. Seminal receptacle present; duct from -
seminal receptacle not observed (judging from other members of the
group it probably joins the oviduct prior to entering ootype). Vagina
double, opening ventrally near lateral margins of body just posterior to
level of genital atrium. Vitellaria co-existensive with intestine, with some
vitellaria extending into posthaptor and appendage (not observed in all
specimens); transverse vitelloducts fusing in midline to form vitelline
reservoir. Eggs fusiform, with filaments at both ends; eggs in utero (12)
214 (182-243) long by (12) 73 (61-83) wide (measurement exclusive
of filaments); filaments approximately half the length of the main body
of the eggs, not united in chains.
Discussion: This species was inadequately described by Hughes
(1928) from Mustelus antarcticus collected at Port Phillip Bay, Victoria,
Australia. Manter (1955) added a few details to Hughes’ description
and a new locality record.
The above redescription is given because the original figures and
description were incomplete.
Erpocotyle callorhynchi (Manter, 1955) Yamaguti, 1963
Figs. 20-25
Synonym: Squalonchocotyle callorhynchi Manter, 1955.
Host: Callorhynchus milii Bory, Elephant Fish; family Chimaeridae.
Location: Gills.
Localities: (1) Timaru, Canterbury Province, South Island; 10 statute
miles ENE of Timaru (9 fathoms; sand), (2) Timaru, Canterbury
Province, South Island; 23 statute miles ENE of Timaru (16 fathoms;
sand—mud), and (3) Timaru, Canterbury Province, South Island; 52
statute miles ENE of Timaru (28 fathoms; sand).
Southern Pacific Trematodes Pt. IV 361
Previously Reported Hosts and Localities: (1) Callorhynchus capensis
Dumeril from Capetown, South Africa, and (2) C. milii Bory from
Wellington, New Zealand (Manter, 1955).
Number Studied: 16.
Homoeotypes: USNM Helm. Coll. No. 71197 (5 specimens).
Description: Body elongate, (7) 8,038 (6,370-9,500) long (measure-
ment exclusive of appendage) by (7) 1,068 (924—1,370) wide. Cuticle
fairly thick, occasionally undulated; surface of body, especially anterior
surface, covered with papillae. Prohaptor an oral sucker (7) 219 (176—
248) long by (7) 282 (171-349) wide; sclerotized papillae on rim and
inner surface of oral sucker; preoral hood or membrane weakly de-
veloped. Posthaptor armed with three pairs of sclerotized suckers with
each sucker possessing a large hooklike sclerite; rims and inner surface
of suckers armed with sclerotized papillae; striae in lumen; suckers with
membranous flange. Hooklike sclerites typically hexabothriid in shape
with posteriormost sclerite (adjacent appendage) always smaller than
other two pairs; anteriormost sclerite (opposite appendage) (12) 476
(390-540) long; middle sclerite (11) 470 (382-554) long; posteriormost
sclerite (adjacent appendage) (11) 356 (293-462) long. Hooklike
sclerites with spines along lateral margins. Appendage armed with two
anchors and two barrel-shaped suckers; anchors (7) 52 (50-55) long.
Disklike sclerites in posthaptor of some specimens.
Mouth ventral, slightly subterminal. Pharynx (5) 91 (78-100) long
by (5) 81 (71-86) wide; esophagus short to medium in length,
branched. Gut bifurcated; crura with medial and lateral branches,
confluent posteriorly prior to entering posthaptor and appendix.
Testes follicular, 46-60 in number, postovarian, between intestinal
crura. Vas deferens sinuous, extending anteriorly in midline to proximal
end of cirrus; cirrus weakly muscular. Genital atrium (5) 97 (82-110)
long by (5) 90 (79-101) wide, situated (6) 636 (589-687) from
anterior end.
Ovary irregularly lobed with mature end directed anteriorly; oviduct
extending anteriorly, joining genito-intestinal canal and vitelline reservoir
before entering ootype. Uterus extending anteriorly in midline to genital
atrium. Seminal receptacle not observed. Vagina double, each opening
ventrally between midline and lateral margin near level of genital atrium.
Vitellaria co-extensive with intestine; not extending into posthaptor or
appendage; transverse vitelloducts fusing in midline to form vitelline
reservoir. Eggs fusiform, (9) 180 (171-187) long by (9) 70 (59-81)
wide; eggs connected by their filaments. Brain posterodorsal to pharynx.
Discussion: A comparison of USNM Helm. Coll. No. 37447 (holo-
type) and No. 37447 (paratype), from South Africa, and No. 37448
(paratype), from New Zealand, with our population from Callorhynchus
milii indicated that the present specimens are conspecific with Erpocotyle
callorhynchi (Manter, 1955) Yamaguti, 1963. Our population agreed
in all particulars with the paratype from Callorhynchus milii from New
Zealand. In the comparison of the holotype and paratype from C.
Ficunres 26-31. Erpocotyle antarctica (Hughes, 1928) Price, 1942.
26, Whole mount, ventral view. 27, Egg. 28, Anchor. 29, Anteriormost
sclerite. 30, Middle sclerite. 31, Posteriormost sclerite.
Southern Pacific Trematodes Pt. IV 363
capensis (from South Africa) with our population fom C. milii, one
difference was noted. In those specimens from C. capensis the poimis
of the hooklike sclerites were considerably longer.
According tp Manter (1955) the eggs of Erpocotyle callorhynchi are
not comnected by their filaments. It appears that this is erroneous because
in the two paratypes from his collections, which contamed numerous
egss in uigro, and m our population of worms the eggs are commecied
by their Glamenis.
Prior to the report of Manter (1955), members of the genus Erpocotyle
(= Squalonchocotyle) had been found only on selachians. They are now
known to be present on both selachians and chimaerids. This distribution
likely reflects the historic genetic relationship between these two groups
of soft-bodied fishes.
Erpocoiyle squali (MacCallum, 1931) Price, 1942
Figs. 14-19
Synonyms: Squalonchocoityle squali MacCallum, 1931; S. acanthi
MacCallum, 1931.
Host: Squalus lebruni (Valliant), Dogfish Shark; family Squalidze.
Location: Gills.
Localities: (1) Timaru, Canterbury Province, South Islend; 23 statute
miles ENE of Timaru (16 fathoms: sand—mnud), (2) Akaroa, Canterbury
Province, South Island: Akaroa Harbour (4-10 fathoms: mud—sand—
mussels), and (3) Taiaroa Heads (mouth of harbor ti» Dumedm), Oigo
Province, South Island; 3 statute miles E of Taiaroa Heads (17 fathoms;
fine sand).
Previously Reporied Host and Locality: From the gills of Squalus
acanthias Linnaeus collected at Woods Hole, Mass. (MacCallum, 1931:
Price, 1942).
Number Studied: 56.
Homoeoiypes: USNM Helm. Coll. No. 71198 (5 specimens).
Description: Body elongate, (10) 5,489 (4,310-6.680) long (measure-
ment not including appendage) by (10) 778 (620—1,060) wide. Cuticle
relatively thin, sometimes undulant; surface of body, especially anterior
portion, covered with fleshlike papillae. Prohaptor an oral sucker (13)
215 (183-269) long by (13) 308 (247-392) wide; oral sucker muscular,
with rims and inner surfaces bearing minute, sclerotized papillae. Preoral
hood or membrane slightly developed. Posthaptor armed with three
pairs of sclerotized suckers, with each sucker possessing a large hooklike
sclerite; sclerotized papillae not observed on rims of suckers; striae in
lumen. Hooklike sclerites typically hexabothriid im shape, nearly equal
in size; anteriormost sclerite (opposite appendage) (14) 428 (337-494)
long: middle sclerite (14) 427 (366—480) long; posteriormost sclerite
(adjacent appendage) (15) 395 (325-480) long. Hooklike sclerites
with a few spimes along lateral margins. Disklike sclerites located in
posthaptor. Appendage (9) 802 (641-952) long, armed with two
anchors and two barrel-shaped suckers; anchors (10) 67 (61-72) long.
364 Proceedings of the Biological Society of Washington
Mouth ventral, slightly subterminal. Pharynx (12) 102 (84-117) long
by (12) 75 (61-89) wide; esophagus short, with slight branching. Gut
bifurcated, with medial and lateral branches; crura confluent posteriorly,
prior to entering posthaptor and appendix; unbranched in either.
Testes 40-60 in number, postovarian, between intestinal crura. Vas
deferens sinuous, broadened and tightly coiled posteriorly narrowed and
loosely coiled anteriorly, extending anteriorly in midline to proximal end
of cirrus. Cirrus weakly muscular, (6) 117 (74-167) long; base of cirrus
surrounded by prostate glands. Genital atrium small, (8) 31 (26-36)
long by (8) 31 (25-37) wide, receiving uterus (ventrally) and cirrus
(dorsally); genital atrium situated (11) 503 (420-655) from anterior
end.
Ovary irregularly looped; distal end of ovary directed anteriorly.
Oviduct convoluted, extending anteriorly from mature end of ovary,
fusing with ducts from genito-intestinal canal and vitelline reservoir
prior to entering ootype; ootype dorsal to vitelline reservoir, surrounded
by Mehlis’ glands. Seminal receptacle present. Vagina double, ventral,
opening just lateral to crura near level of genital atrium. Vitellaria
follicular, co-extensive with crura and not extending into posthaptor or
appendage. Eggs fusiform, (8) 231 (210-247) long by (8) 104 (87--
135) wide (measurement not including filaments), with filament (usually
less than length of egg) at both poles; filaments not forming chains.
Discussion: A comparison of USNM Helm. Coll. Nos. 8133, 8134, and
8135 (a total of eight specimens) with our population from Squalis
lebruni indicated that the present specimens are conspecific with Erpoc-
otyle squali (MacCallum, 1931) Price, 1942.
The above redescription is given because: (1) previous descriptions
of this species (MacCallum, 1931; Price, 1942) were incomplete; and
(2) certain discrepancies exist between those descriptions and the type
material on which they were based.
This fourth of a series on monogenetic trematodes from the
southern Pacific Ocean discusses seven species of monogenetic
trematodes from New Zealand waters. Two new species are
described: Neogrubea seriolellae and Hexabothrium aka-
roensis. Erpocotyle antarctica (Hughes, 1928) Price, 1942,
E. callorhynchi (Manter, 1955) Yamaguti, 1963, and E. squali
(MacCallum, 1931) Price, 1942 are redescribed. Eurysorchis
australis Manter and Walling, 1958 and Diclidophora coelo-
rhynchi Robinson, 1961 are reported.
In Mazocraeidae Price, 1936, the subfamily Neogrubeinae is
established to accommodate the new genus Neogrubea.
A new locality record is reported for Erpocotyle squali and
new host records are reported for E. squali and Eurysorchis
australis.
Southern Pacific Trematodes Pt. IV 365
LITERATURE CITED
CERFONTAINE, P. 1899. Les Onchocotylinae. (Contribution a l’étude
des octocotylidés. 5.) Arch. Biol., 16 (3): 345-478.
Ditton, W. A., AND W. J. Harcis, Jk. 1965. Monogenetic trematodes
from the southern Pacific Ocean. I. Monopisthocotyleids
from New Zealand fishes, p. 229-249. In Biology of the
Antarctic Seas II, Ant. Res. Ser. 5.
Hucues, W. K. 1928. Some trematode parasites on the gills of Vic-
torian fishes. Proc. Roy. Soc. Victoria, N.S., 41 (1): 45-54.
MacCa.tium, G. A. 1931. Four new species of trematode worms of
the subfamily Onchocotylinae. Proc. U.S. Nat. Mus., 79:
1-8.
Manter, H. W. 1955. Two new monogenetic trematodes from ele-
phant fishes (Callorhynchus) from South Africa and New
Zealand, p. 211-220. In Essays in the natural sciences in
honor of Captain Allan Hancock. Univ. Southern California
Press.
Manter, H. W., AnD G. WaALLING. 1958. A new genus of monogenetic
trematode (family Diclidophoridae) from a New Zealand
fish. Proc. Helminthol. Soc. Wash., 25 (1): 45-47.
Price, E. W. 1942. North American monogenetic trematodes. V. The
family Hexabothriidae, n. n. (Polystomatoidea). Proc. Hel-
minthol. Soc. Wash., 9 (2): 39-56.
. 1961. North American monogenetic trematodes. IX. The
families Mazocraeidae and Plectanocotylidae. Proc. Biol. Soc.
Wash., 74: 127-155.
Rosrnson, E. S. 1961. Some monogenetic trematodes from marine
fishes of the Pacific. Trans. Amer. Microscop. Soc., 80 (3):
235-266.
366 Proceedings of the Biological Society of Washington
B7Lf.06735
Vol. 81, pp. 367-402 30 August 1968
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
MONOGENETIC TREMATODES FROM THE
SOUTHERN PACIFIC OCEAN. PART V.
MONOPISTHOCOTYLEIDS FROM-AUSTRALIAN
FISHES, THE SUBFAMILY TROCHOPODINAE??
By ApriAN RussELL LAWLER AND WILLIAM J. Harcis, Jr.
Parasitology Section, Virginia Institute of Marine
Science, Gloucester Point, Virginia
In 1958 the Parasitology Section of the Virginia Institute of
Marine Science undertook a study of the host specificity,
zoogeography and systematics of the monogenetic trematodes
and other parasites of Antarctic vertebrates. Subsequently
two field expeditions were conducted, one at NAF McMurdo
(1959) and the other at Wilkes Station (1961), Antarctica.
As research progressed it was decided to include comparisons
of parasites from Antarctic hosts with those from hosts taken
from nearby ocean bottoms and shoal waters of adjacent land
masses. To this end comparison collections were made of
fishes from New Zealand, Australia, southern Chile, Drake
Passage, Gulf of Guinea, and the Indian Ocean.
This paper on the Monogenea of certain Australian marine
fishes is the first, of several planned, resulting from the Aus-
tralian expedition which was undertaken as part of the United
States Antarctic Research Program of the National Science
Foundation. Subsequent publications in this series will include
parasites from both sides of the Antarctic convergence.
This study is based on microscopic examination and system-
atic considerations of monogenetic trematodes collected from
the gills of 1,909 host individuals representing 137 species.
Parasites representing approximately 130 species and 17 fam-
1 Contribution No. 277 from the Virginia Institute of Marine Science.
2 Supported by grants GA-13853, with amendments, and GA-235 under the United
States Antarctic Research Program of the National Science Foundation.
39—Proc. Brow. Soc. WasH., VoL. 81, 1968 (367 )
368 Proceedings of the Biological Society of Washington
ilies were found on 83 of these host species. The remaining
54 were not parasitized by monogentic trematodes. Only six
parasite species from five host species are discussed here. The
others and a special discussion of ecological aspects will be
the subjects of later works.
Examination of the literature reveals that in comparison to
the total number of hosts and monogeneids reported from
elsewhere in the world little is known about the monogeneid
fauna of the Australian area. MacCallum (1917), Johnston
' and Tiegs (1922), Hughes (1928), Johnston (1929, 1930a,
1930b, 1931, 1934a, 1934b, 1937), Murray (1931), Woolcock
(1936), Sandars (1944, 1945, 1947), Hargis and Dillon (1965),
and Young (1967a, 1967b, 1967c, 1968) have described or
reported monogenetic trematodes from fishes of Australian
waters. Robinson (1961) noted that the majority of Mono-
genea known from the Southern Hemisphere were reported
from Australia. Other works concerning Monogenea from
the Southern Hemisphere are as follows: Brinkmann (1952)
and Cordero (1944) from Chile; Manter and Prince (1953)
and Laird (1958) from Fiji and the New Hebrides; and
Blanchard (1847), Johnston (1931), Manter (1955), Manter
and Walling (1958), Robinson (1961), Dillon and Hargis
(1965a, 1965b), and Hargis and Dillon (1965) from New
Zealand. Up to the present time a total of 79 species of
monogenetic trematodes has been reported from Australian
waters, 12 of these being reported by Johnston and Tiegs
(1922) from freshwater fishes.
The research reported herein is a continuation of a long-
range study of certain aspects of host specificity, zoogeog-
raphy, and phylogeny of monogenetic trematodes being con-
ducted by members of this Institute.
Materials and Methods: Host collections were made near Tweed Heads
and Ulladulla, New South Wales; Lakes Entrance, Victoria; Gladstone,
Queensland; Dunalley and Hobart, Tasmania; and Port Kenney, South
Australia. The fish were procured from commercial fishery operations
in which Danish seines, hand lines, and gill nets were used. Messrs.
William Stanley Wilson and William Saunders, the field collectors, ac-
companied the vessels and took the host specimens as they came on
board. Collections were made from March through July of 1962.
Mr. Wilson identified the fish with the aid of experienced fishing
Southern Pacific Trematodes Pt. V 369
vessel captains and using the keys and descriptions of Waite (1923),
Graham (1956), Parrott (1957, 1958, 1959), and Roughley (1953).
Since the vessel captains could help with or verify identifications, and
since all species captured are relatively common, it is believed that host
identifications are reliable. Scientific names of hosts are those given by
Munro (1958), Roughley (1953), and Parrott (1959).
In the locality descriptions given below, the nearest town or prominent
geographical feature and its province are given first, followed by the
approximate site of capture of the host. The place of capture is followed
by the depth and bottom type in parentheses. Distance is in statute
miles.
The monogenetic trematodes were collected using a procedure outlined
by Hargis (1953). This technique works best when the gill arches are
separated from each other before immersion in the relaxant. However,
according to Mr. Wilson, such separation was generally not possible
because of trying shipboard conditions. As a result, relaxation of worms
on internal gill surfaces not readily bathed by the Chloretone-seawater
solution was somewhat variable. Such unevenness of relaxation probably
also resulted from the varying physiological conditions of the worms
themselves at the time of killing, since all hosts could not be processed
at the same time. The trematodes were killed, fixed, and preserved by
adding AFA (aceto-formalin-alcohol). This technique has proven advan-
tageous when collecting large numbers of hosts as it facilitates rapid
handling.
The parasites were removed from the gill material and sediment with
the aid of a stereomicroscope and stored in vials containing a solution
of 5% glycerol in 70% ethanol.
For preparation of whole mounts the worms were removed from the
preservative, hydrated, overstained, destained, dehydrated, cleared, and
mounted. The parasites were stained with one of the following: (1)
Reynolds’ double stain (Delafield’s hematoxylin plus alum cochineal);
(2) alum cochineal; (3) Harris’ hematoxylin; and (4) Harris’ hema-
toxylin with either sodium bicarbonate for “bluing” or eosin as a counter-
stain. As many stains as possible were used on each species to study the
various internal and external structures. The following gave good results:
(1) Reynolds’ double stain for internal structures and glands, (2) alum
cochineal as a general stain, (3) Harris’ hematoxylin, alone or with
sodium bicarbonate for reproductive structures, (4) the latter stain, with
eosin as a counterstain, for reproductive organs and to reveal posthaptoral
septation. If worms were few, only alum cochineal and/or Reynolds’
double stain were used.
The worms were overstained and immediately destained using a solu-
tion of two to four drops of concentrated HC1 in 100 ml of 30% ethanol.
This procedure afforded better control of the amount of stain retained
by the specimen than progressive staining. After dehydration, the worms
were cleared in deacidified beechwood creosote and mounted permanently
in Piccolyte.
370 Proceedings of the Biological Society of Washington
Only those specimens which were well-relaxed and possessed clear
morphological characters were used for identification and study. Diag-
noses and descriptions were based on adult individuals, sexual maturity
being the criterion for adulthood. Sexual maturity was determined by
either (1) the presence of an egg in utero, (2) attainment of the same
approximate size and morphological condition as individuals with eggs,
or (3) by the apparently mature condition of the gonads (especially
the ovary) where no egg was observed.
All measurements were made with the use of a filar micrometer and
are given in microns unless otherwise noted. In indicating these measure-
ments the mean is given, followed by the range (minimum and maxi-
mum) in parentheses. In cases where more than five specimens were
measured, the standard deviation (S), standard error (Sz), and the
interval estimate at the 95% level (t.sSz) follow the range. For con-
venience the alphabetical symbols SE and CL are established for standard
error (Sz) and confidence limits or interval estimate at the 95% level
(t.osSz), respectively. The number of measurements used in the calcula-
tions appears in parentheses before these data. Measurements of curved
structures were across the lines subtending the greatest arcs described
by these structures. All egg measurements were taken of the main portion .
of the egg capsule, exclusive of the filament. The value of filament
length as a taxonomic character is doubtful since great variation occurs
within a species. Measurements of soft parts, which are subject to
contraction and expansion in life and shrinkage in death, can be con-
sidered of value for comparison only when specimens have been similarly
treated or where differences between individuals or groups of individuals
being compared are great. Measurements of hard parts are thought to
be free of such vagaries and are considered more reliable as taxonomic
characters. Camera lucida and microprojector drawings were used to
facilitate identification and in the preparation of figures.
The taxonomic scheme employed is essentially that of Sproston (1946)
and Yamaguti (1963). The work of Bychowsky (1957) has also been
considered and his conclusions discussed where possible.
The morphological terminology used in the descriptions is that of
Hargis (1958), who compiled a list of useful terms from his own studies
and from the literature.
ORDER MONOGENEA CARUS, 1863
SUBORDER MONOPISTHOCOTYLEA ODHNER, 1912
SUPERFAMILY CAPSALOIDEA PRICE, 1936
FAMILY CaApsALIDAE BArrp, 1853
Subfamily Trochopodinae (Price, 1936) Sproston, 1946, emend.
Synonyms: Trochopinae Price, 1936; Megalocotylinae Bychowsky,
1957.
Diagnosis: Capsalidae. Diagnosis that of Yamaguti (1963) except as
Southern Pacific Trematodes Pt. V 371
follows: (1) pharynx with or without constrictions; and (2) common
genital opening marginal or proximal to margin behind left prohaptor.
Type Genus: Trochopus Diesing, 1850.
Discussion: The above emendation is made to accommodate Allo-
sprostonia tauvinae new genus, new species. In addition, the figure of
Sprostonia squatinae (MacCallum, 1921) Bychowsky, 1957 as illustrated
by MacCallum (1921) shows definite constrictions of the pharynx, which
Yamaguti (1963) overlooked in his diagnosis of the subfamily Trocho-
podinae.
Price (1936) proposed a new subfamily in the Capsalidae, the group
Trochopinae which included the genera Trochopus Diesing, 1850, and
Macrophyllida Johnston, 1929. He suppressed Megalocotyle Folda, 1928
as a synonym of Trochopus Diesing, 1850. Upon re-examination of the
large number of species of Trochopus, Price (1939) decided that they
fall into two groups and therefore reinstated the genus Megalocotyle
Folda, 1928.
Dr. Baylis, in personal communication with Sproston (Sproston, 1946),
pointed out that the correct spelling of the subfamily name was Trocho-
podinae. Sproston (1946) then emended the subfamily and corrected
the name according to Dr. Baylis’ suggestion.
Bychowsky (1957) split the subfamily Trochopodinae (Price, 1936)
Sproston, 1946 into two subfamilies: (1) Megalocotylinae Bychowsky,
1957, with Megalocotyle Folda, 1928 as the type genus, and (2) Trocho-
podinae (Price, 1936) Sproston, 1946, with Trochopus Diesing, 1850 as
the type genus. He did this on the basis of whether there were an
unequal or equal number, in the former and latter respectively, of
peripheral depressions separated by septa on the posthaptor.
Yamaguti (1963) considered Megalocotylinae Bychowsky, 1957 as a
synonym of Trochopodinae (Price, 1936) Sproston, 1946, combining the
two. He also proposed two new genera, Allomegalocotyla and Pseudo-
megalocotyla.
After careful study of the generic characters (Table 1) which may
be considered taxonomically significant (type of prohaptor, number of
septa, position of prostate reservoir, testicular arrangement, position of
vaginal opening) within the subfamily Trochopodinae (Price, 1936)
Sproston, 1946, the authors accept, with the emendations previously
made and to follow, the rearrangement as proposed by Yamaguti (1963).
The genus Trochopella, as proposed by Euzet and Trilles (1962), is
considered to be a synonym of Trochopus Diesing, 1850. As can be
ascertained from Table 1, all the major taxonomic characters of the
genus Trochopella are the same as those of Trochopus. Although Euzet
and Trilles (1962) did not mention the position of the prostate reservoir
in the text, their drawing indicates its presence inside of the cirrus pouch.
Genus Trochopus Diesing, 1850
Synonym: Placunella Beneden and Hesse, 1863.
Diagnosis: Trochopodinae. Diagnosis that of Yamaguti (1963).
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Southern Pacific Trematodes Pt. V 373
Type Species: Trochopus tubiporus (Diesing, 1836) Beneden and
Hesse, 1863.
Trochopus hobo Yamaguti, 1942
Figs. 1-10
Host: Chelidonichthys kumu (Lesson and Garnot), Red Gurnard or
Kumukumu or Latchet; family Triglidae.
Locality: Lakes Entrance, Victoria; 45 statute miles ESE of Lakes
Entrance (65-75 fathoms).
Previously Reported Host and Locality: Chelidonichthys kumu (Lesson
and Garmot); Hamazima, Japan.
Gear Used: Danish seine.
Location: Gills.
Number Studied: 19.
Number Measured: 12.
Homoeotypes: USNM Helm. Coll. No. 71199 (four specimens ).
Redescription: Body elliptical, flattened dorsoventrally, (6) 3,640
(2,860-4,670), S = 707, SE = 289, CL = 742 long by (8) 2,100 (1,490-
3,000), S = 553, SE = 195, CL = 462 wide. Cuticle fairly thin and
smooth. Prohaptor a pair of ventrolateral suckers; left sucker (8) 503
(405-602), S = 68.6, SE = 24.3, CL = 57.4 long by (11) 501 (416-
636), S = 71.5, SE = 21.6, CL = 48.0 wide; right sucker (9) 503 (399-
600), S = 74.1, SE = 24.7, CL = 57.0 long by (12) 501 (423-638), S
= 70.3, SE = 20.3, CL = 44.7 wide. Posthaptor a subsessile, concavo-
convex sucker, opening ventrally, (10) 1,430 (1,120-1,950), S = 298,
SE = 94.2, CL = 213 long by (10) 1,530 (1,220-2,070), S = 285,
SE = 90.1, CL = 204 wide, divided by septa into one small, crescent-
shaped central and ten peripheral depressions (loculi); margin of post-
haptor a strong muscular rim, surrounded by scalloped marginal mem-
brane (11) 135 (110-173), S = 19.4, SE = 5.84; CL = 13.0 wide;
armed with 3 pairs of dissimilar anchors and 14 marginal hooks. First
anchors large, robust, left (6) 103 (85.5-116), S = 11.4, SE 4.65,
Gi = 19/0 long and right (5) 104 (914-117), S = 10.5, SE = 479,
CL = 13.1 long, with slightly knobbed and tapered external tips and
bifid (a slightly pointed ventral and a blunt dorsal) internal tips; second
pair of anchors elongate, left (4) 96.2 (81.0-111) long and right (7)
(96.8) (78.7-116), S = 10.8, SE = 4.07, CL = 9.97 long, more robust
than third pair with slightly recurved blunt tips; third pair of anchors
elongate, left (1) 110 long and right (2) 97.6 (84.0-111) long, with
recurved pointed tips. Posthaptoral hooks (5) 9.91 (7.96-12.4), S =
2 Ola 04> Chi —)2-88rlongs
Pharynx muscular, (10) 385 (273-543), S = 88.9, SE = 28.1, CL =
63l6llone by, (10/420) (3032662), S| = 10eSE —"341) Chl = 77.2
wide, papillated internally. Mouth subterminal, anteroventral to pharynx.
Esophagus short; gut bifurcated, crura with medial and lateral dendritic
branching, not confluent posteriorly.
Two testes, juxtaposed, entire, fenestrated, oval in outline, left (8) 534
374 Proceedings of the Biological Society of Washington
Ficures 1-10. Trochopus hobo Yamaguti, 1942. (Scale is in mm.)
1, Whole mount, ventral view (posthaptor twisted bringing posterior
loculus about 135° out of alignment in the clockwise direction). 2, Re-
productive organs and terminal genitalia, ventral view. 3, Left first
anchor. 4, Right first anchor. 5, Left second anchor. 6, Right second
anchor. 7, Left third anchor. 8, Right third anchor. 9, Egg in utero,
ventral view. 10, Left prohaptoral sucker, ventral view.
Southern Pacific Trematodes Pt. V 375
(383-696), S = 117, SE = 41.2, CL = 97.5 long by (8) 391 (286-
592), S = 110, SE = 38.9, CL = 92.0 wide and right (8) 529 (385-
696), S = 118, SE = 41.7, CL = 98.7 long by (8) 379 (267-572), S
= 119, SE = 42.2, CL = 99.7 wide, equatorial in position. Vasa
efferentia anastomosing in midline to form convoluted vas deferens; vas
deferens proceeding anteriorly, dorsal to left arm of vitelline reservoir
and vagina, whence it turns right and passes dorsal to ootype and cirrus
pouch prior to entering cirrus pouch. Cirrus obliquely situated just
posterior to pharynx. Cirrus complex consisting of cirrus and prostate
reservoir in cirrus pouch. Prostate reservoir ventral to cirrus proper, with
small duct leading to prostatic cells surrounding proximal end of cirrus
pouch. Uterus connecting to cirrus pouch at level of posterior border of
pharynx forming genital atrium, latter emptying via marginal genital
pore on left just above level of pharynx. Three glands of Goto in midline
just posterior to testes.
Ovary pretesticular, oval, entire, median, (9) 323 (222-419), S =
66.8, SE = 22.3, CL = 51.4 long by (9) 225 (159-291), S = 46.2,
SE = 15.4, CL = 35.5 wide, with intemal chamber (= seminal recep-
tacle of Meserve, 1938) containing mature ova; oviduct passing from
internal chamber, dorsal to right arm of vitelline reservoir and sinuously
to ootype. Ootype obliquely situated between cirrus and vagina, sur-
rounded by Mehlis’ gland cells; uterus short, opening into genital atrium.
Vagina long, with bulbous swelling at proximal end, opening marginally
on left just posterior to common genital pore.
Vitellaria follicular, extending from level of prohaptoral suckers to
very near posterior end of body proper. Transverse vitelloducts fusing
medially to form vitelline reservoir immediately anterosinistral to ovary;
vitelloducts confluent just posterior to testes. Egg in utero polyhedral,
(1) 147 long by (1) 136 wide, with convoluted basal filament.
Brain anterodorsal to pharynx; three pairs of nerves passing into
prohaptoral region. Four granular eyespots located dorsal to brain, the
first pair smaller and closer together than the posterior pair. Excretory
pores at level of distal end of ootype, opening dorsolaterally.
Discussion: The worms in this collection, though larger than those in
the original description, are probably conspecific with Trochopus hobo
Yamaguti, 1942. However, our population differs from Yamaguti’s (1942)
description in the following: (1) presence of 14 marginal hooks instead
of 16; (2) presence of crescent-shaped central loculus; (3) lengths of
anchors; (4) prostate reservoir inside cirrus pouch instead of outside;
and (5) presence of membranous flap around each prohaptoral sucker.
A small crescent-shaped central loculus was illustrated by Palombi (1949)
for Trochopus tubiporus (Diesing, 1836), the type species of the genus.
Although Yamaguti (1942) did not mention such a central loculus, he
did illustrate a small slit in his drawing. The apparent discrepancy in
anchor sizes as shown in Table 2 (our population, although composed
of larger specimens, having smaller second and third anchors than
Yamaguti’s population) is hard to explain. The worms of our sample
376 Proceedings of the Biological Society of Washington
TaBLeE 2. Comparison of measurements of Trochopus hobo
Yamaguti, 1942.
Yamaguti (1942) Present sample
Entire body (L) 1,800-2,900 2,,860-4,670
(WwW) 500-1,000 1,490-3,000
Posthaptor 430-780 (dia.) (L) 1,120-1,950
(W) 1,220-2,070
Marginal membrane (W) 30-60 110-173
Anchor 1 (L) 60-85 L-(L) 85.5-116
(W) 11-15 R-(L) 91.4-117
Anchor , 2 (L) 115-170 L-(L) 81-111
(W) 13-21 R-(L) 78.7-116
Anchor 3 (L) 130-210 L-(L) 110
(W) 27-50 R-(L) 84-111
Hooks (L) 10 7.96-12.4
Prohaptoral suckers (L) 180-350 L-(L) 405-602
(W) 130-270 (W) 416-636
R-(L) 399-600
(W) 423-638
Pharynx (L) 120-260 273-543
(W) 160-310 303-662
Testes (L) 230-360 L-(L) 383-696
(W ) 120-200 (W) 286-592
R-(L) 385-696
(W) 267-572
Ovary (L) 114-210 229-419
(W) 80-175 159-291
Egg 110-150 (dia.) (L) 147
(Woe s1386
(L) = Length. (W) = Width. L= Left. R = Right. (dia.) = diameter.
are otherwise identical to those of Yamaguti (1942), except for size
differences.
The three “oval giant cells of unknown nature” (Yamaguti, 1942)
were observed in our population. They are probably glands of Goto.
Nineteen parasites were recovered from 29 host specimens (Table 6).
This study represents a new locality record for this species.
Genus Sprostonia Bychowsky, 1957, emend.
Diagnosis: Capsalidae, Trochopodinae.
Diagnosis that of Yamaguti
(1963) except as follows: Posthaptor divided by seven primary (con-
Southern Pacific Trematodes Pt. V 377
nected to central loculus) septa, of which the posterolaterals are either
trifid or bifid, with either four or six secondary (incomplete) septa.
Pharynx notched marginally into eight lobes. Vagina opening close to
genital pore. Parasitic on elasmobranchs and teleosts.
Type Species: Sprostonia squatinae (MacCallum, 1921) Bychowsky,
1957.
Discussion: The above emendation is made to accommodate Sprostonia
longiphallus n. sp. In addition, though not mentioned by Bychowsky
(1957), the pharynx of S. squatinae as figured by MacCallum (1921)
is notched marginally into eight lobes, as is the pharynx of S. longiphal-
lus. Yamaguti (1963) wrote that the vagina opened “close to, or together
with, genital pore.” The phrase, “or together with,’ should not be
included in the diagnosis, as Price (1937) stated that the vagina had
its opening “immediately posterior to genital aperture,” and all specimens
of S. longiphallus have vaginal openings proximal and posterior to the
genital pore.
The genus Sprostonia Bychowsky, 1957 includes one previously de-
scribed species, S. squatinae, described by MacCallum (1921) as Acantho-
cotyle squatinae from the host Squatina squatina (Linn.) from Singa-
pore. Price (1937) redescribed it from specimens that MacCallum
(1921) used for his original description, renaming it Trochopus squa-
tinae. Later Price (1939) reassigned it to the genus Megalocotyle Folda,
1928. Brinkmann (1940), apparently in ignorance of Price’s work (1936,
1937, 1939), also placed it in the latter genus. Sproston (1946) con-
sidered its inclusion in this genus as “very doubtful.” Bychowsky (1957)
isolated this species into the new genus Sprostonia Bychowsky, 1957, on
the basis of (a) the more complex nature of the posthaptor with its
incomplete septa and (b) the paired anterolateral suckers united antero-
dorsally by a common hood.
Sprostonia longiphallus new species
Figs. 11-16
Host: Epinephelus tauvina (Forskal), Greasy Cod or Estuary Rock-
Cod; family Epinephelidae.
Locality: Gladstone, Queensland; 36 statute miles ENE of Gladstone
(9 fathoms, coral).
Gear Used: Hand line.
Location: Gills.
Number Studied: 83.
Number Measured: 19.
Holotype: USNM Helm. Coll. No. 71200.
Paratypes: USNM Helm. Coll. No. 71201 (five specimens ).
Description: Body elliptical, flattened dorsoventrally, (19) 3,650
(2,910-4,930), S = 494, SE = 113, CL = 238 long by (19) 1,310
(1,060-1,600), S = 135, SE = 31.0, CL = 65.1 wide. Cuticle fairly
thin and smooth. Prohaptor a pair of ventrolateral suckers, united antero-
dorsally by a common hood; left sucker (15) 143 (124-163), S = 12.4,
378 Proceedings of the Biological Society of Washington
Os
BOSE sheet
ie
14
t
—————
CS
Ficures 11-16. Sprostonia longiphallus n. sp. (Scale isin mm.) 11,
Whole mount, ventral view. 12, Reproductive organs and terminal geni-
talia, ventral view. 13, Left first anchor. 14, Right first anchor. 15,
Right second (dorsal to third anchor) and right third anchor. 16, Egg
in utero, ventral view.
Southern Pacific Trematodes Pt. V 379
SE = 3.20, CL = 6.87 long by (15) 192 (156-231), S = 20.5, SE =
5.30, CL = 11.4 wide, right sucker (13) 141 (116-160), S = 9.58,
SE = 2.66, CL = 5.79 long by (13) 184 (161-222), S = 16.3, SE =
4.52, CL = 9.86 wide. Region anterior to brain glandular. Posthaptor
a subsessile, concavo-convex, oval sucker, opening ventrally, (19) 624
(514-829), S = 84.9, SE = 19.5, CL = 40.9 long by (19) 790 (626-
994), S = 115, SE = 26.4, CL = 55.4 wide, divided by seven primary
(connected to central loculus) septa, the posterolateral bifid, with two
secondary (incomplete) septa extending inward from muscular rim sub-
dividing posterior marginal loculus, with a secondary septum dividing
each of the anterior and anterolateral marginal loculi; margin of post-
haptor a strong muscular rim, surrounded by a delicate, scalloped mar-
ginal membrane (19) 52.5 (38.8-66.9), S = 8.50, SE = 1.95, CL =
4.10 wide; armed with 3 pairs of dissimilar anchors and 14 marginal
hooks. First anchors large, robust, left (15) 136 (108-165), S = 17.4,
SE = 4.49, CL = 9.64 long; right (16) 142 (117-174), S = 145,
SE = 3.62, CL = 7.71 long, external tips pointed, internal blunt, and
enlarged in middle; second pair of anchors elongate, left (14) 50.9
(43.9-61.0), S = 5.39, SE = 1.44, CL = 3.11 long, more robust than
third pair, with tips slightly recurved, blunt; third pair of anchors
elongate, left (14) 55.4 (46.9-70.2), S = 7.10, SE = 1.90, CL = 4.10
long, tips recurved and pointed. (Second and third pairs of anchors
proximated, arising subequally on posthaptor and are sometimes hard to
distinguish from each other.) Posthaptoral hooks (19) 6.01 (4.49-8.16),
S = 0.991, SE = 0.227, CL = 0.478 long. Posthaptor sometimes with
small, round, disclike sclerites near first pair of anchors.
Pharynx muscular, (19) 270 (227-367), S = 35.8, SE = 8.21, CL =
17.2 long by (19) 236 (184-281), S = 29.9, SE = 6.85, CL = 14.4
wide, constricted into eight distinct lobes. Mouth subterminal, ventral,
immediately anterior to pharynx. Esophagus very short; gut bifurcated,
crura with medial and lateral dendritic branching, not confluent poste-
riorly.
Two testes, juxtaposed, entire, fenestrated, oval in outline, left (19)
328 (263-451), S = 53.6, SE = 12.3, CL = 25.9 long by (19) 296
(217-370), S = 44.6, SE = 10.2, CL = 21.5 wide, and right (19) 327
(247-441), S = 50.6, SE = 11.6, CL = 24.4 long by (19) 289 (216-
365), S = 41.2, SE = 9.45, CL = 19.9 wide, postequatorial in position.
Vasa efferentia anastomosing in midline to form vas deferens, which
runs anteriorly between vagina and base of cirrus pouch to level of distal
end of ootype, right and posteriorly to parallel cirrus pouch, ventral to
proximal end of cirrus and across posterior of prostate reservoir (which
it is assumed to enter). Cirrus pouch elongate, its proximal end lying
left of median line and directed anteriorly, extending diagonally across
body to right of median line as far as ovary, then tuming anteriorly.
Cirrus very long and slender, eversible, coiled in cirrus pouch. Prostate
reservoir lying in median field, right of proximal end of cirrus, and
connected to cirrus by a duct which continues throughout its length as
380 Proceedings of the Biological Society of Washington
TABLE 3. Comparison of measurements of Sprostonia species.
S. squatinae S. squatinae S. longiphallus
MacCallum (1921) Price (1937) n. sp.
Entire body (L) 4,500 2,900-3,500 2,910—4,930
(W) 1,600 1,300-1,500 1,060—1,600
Posthaptor 1,000 (dia.) 544-599 (dia.) (L) 514-829
~ (W) 626-994
Marginal membrane (W) —— 47-57 38.8-66.9
Anchor 1 (L) 140 143-148 L 108-165
R 117-174
Anchor 2 (L) —— 24-38 L 43.9-61.0
Anchor 3 (L) —- 42 L 46.9-70.2
Hooks (L) — — 4.49-8.16
Prohaptoral suckers —_— (L) 95 L-(L) 124-163
(W) 172 (W) 156-231
R-(L) 116-160
(W) 161-222
Pharynx 800 266 (dia. ) (L) 227-367
(W) 184-281
Testes 1,120 L-(L) 400 L-(L) 263-451
(W) 340 (W) 217-370
R(dia.) 340 R-(L) 247-441
(W) 216-365
Ovary 480 190 (dia.) (L) 124-208
(W) 187-278
Egg 140 (W) 120 (L) 105
(W) 68.5—-79.4
(L) = Length. (W) = Width. L = Left. R = Right. (dia.) = diameter.
ejaculatory duct. Prostatic cells surround prostate reservoir except on
left. Cirrus pouch joining uterus near margin to form genital atrium
opening to outside via the common marginal genital pore on left at
anterior level of brain. Glands of Goto irregular in outline, on each side
of midline posterior to testes.
Ovary immediately pretesticular, oval, entire, median, (19) 170 (124—
208), S = 22.2, SE = 5.09, CL = 10.7 long by (19) 233 (187-278),
S = 29.6, SE = 6.79, CL = 14.3 wide, having internal chamber (=
seminal receptacle of Meserve, 1938) containing mature ova; oviduct
passing from chamber dorsal to right arm of vitelline reservoir, ventral
to vas deferens on left to proximal end of cirrus, then dorsal to vas
deferens and ootype. Ootype oblique, between cirrus and vagina, sur-
rounded by Mehlis’ gland cells; uterus very long, opening into genital
atrium. Vagina muscular, elongate, wavy near distal end, with base at
level of posterior end of prostate reservoir, opening marginally just
posterior to common genital pore; connected by small duct to right side
of vitelline reservoir.
ee
Southern Pacific Trematodes Pt. V 381
Vitellaria follicular, extending from level of anterior part of pharynx
to near posterior end of body. Transverse vitelloducts fusing to left of
median line to form vitelline reservoir immediately anterolateral to
ovary. Vitelloducts confluent postpharyngeally and_posttesticularly.
Egg in utero polyhedral, (2) 105 long by (2) 74.0 (68.5-79.4) wide,
with convoluted basal filament.
Brain anterior to pharynx, three pairs of nerves passing into prohaptoral
region. Four granular eyespots dorsal to brain, first pair smaller and
closer together than posterior pair. Excretory pores at level of anterior
of ootype, opening dorsolaterally.
Discussion: The present species is very similar to Sprostonia squatinae
(MacCallum, 1921) Bychowsky, 1957, from the gills of Squatina squatina
(Linn.), but differs in the following respects: (1) glands of Goto
present; (2) vitelloducts confluent postpharyngeally and _posttestic-
ularly; (3) testes fenestrated; (4) posterolateral septa bifid instead of
trifid; (5) second and third pairs of anchors larger (Table 3); (6)
prohaptoral suckers larger (Table 3); (7) egg smaller (Table 3); (8)
vas deferens passing dorsal to left arm of vitelline reservoir and ventral
to prostate reservoir, which it apparently enters; (9) proximal end of
vagina more anterior and also connected by a small duct to the right
side of the vitelline reservoir; and, (10) host.
The septation of the posthaptor is very close to that described by
Price (1937) for Sprostonia squatinae (MacCallum, 1921) Bychowsky,
1957. The major difference is that the posterolateral septa are bifid
instead of trifid. This observation was facilitated by the use of eosin as
a counterstain on several specimens. It is possible that Price (1937)
observed something similar to the “obsolescent radii devoid of special
musculature,” described by Johnston (1930a) for Macrophyllida antarc-
tica, passing from the anterolateral secondary septa to the posterolateral
bifid septa instead of an actual elevated septum. Such radii were
observed on several of the specimens in our sample.
There is some confusion over anchor classification in the paper by
Price (1937), where those designated as the second and third pairs are
really the third and second, respectively. In other Trochopodinae the
coarse, slightly-curved and blunt-tipped anchor is designated as the
second anchor while the finer anchor with the recurved and sharp tip
is the third. In many capsalids these anchors do occupy the second
and third positions behind the first anchor along the anteroposterior axis
of the posthaptor. In Sprostonia, however, they lie side-by-side rather
than in tandem. The terms “second anchor” and “third anchor” denote
specific morphological entities with definite embryological histories as
do “femur” and “tibia” in vertebrate anatomy. They cannot be mixed.
The arrangement of the posterior two pairs of anchors in Sprostonia
Bychowsky, 1957 can thus be best explained by a migration backward
of the second pair or a migration forward of the third pair, resulting
in their lying side-by-side.
This description increases the number of species in the genus Sprostonia
382 Proceedings of the Biological Society of Washington
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Ficures 17-20. Allosprostonia tauvinae n. gen., n. sp. (Scale is in
mm.) 17, Whole mount, ventral view. 18, Reproductive organs and
terminal genitalia, ventral view. 19, Left first anchor. 20, Left second
(dorsal to third anchor) and left third anchor.
Bychowsky, 1957 to two: the type species from Squatina squatina
(Linn.) (Singapore, Malaysia); and S. longiphallus n. sp. from Epineph-
elus tauvina (Forskal) (Gladstone, Queensland, Australia). This non-
rigid supraspecificity (Hargis, 1957) of Sprostonia Bychowsky, 1957
could possibly represent paleoecological or neoecological relationships
between the hosts.
Southern Pacific Trematodes Pt. V 383
A total of 83 parasites was recovered from 7 host specimens (Table
6).
Allosprostonia tauvinae new genus, new species
Figs. 17-20
Host: Epinephelus tauvina (Forskal), Greasy Cod or Estuary Rock-
Cod; family Epinephelidae.
Locality: Gladstone, Queensland; 36 statute miles ENE of Gladstone
(9 fathoms, coral).
Gear Used: Hand line.
Location: Gills.
Number Studied and Measured: 2.
Holotype: USNM Helm. Coll. No. 71202.
Paratype: USNM Helm. Coll. No. 71203.
Description: Body elliptical, flattened dorsoventrally, 2,860 (2,620—
3,100) long by 1,250 (1,140-1,360) wide. Cuticle fairly thin and
smooth. Prohaptor a pair of ventrolateral suckers, united anterodorsally
by a common hood; left sucker 142 (125-159) long by 161 (151-172)
wide and right sucker 148 (136-161) long by 164 (154-175) wide.
Head glands in three major areas, in body between prohaptoral suckers
and posterolateral to each sucker. Posthaptor a subsessile, concavo-
convex, oval sucker, opening ventrally, 607 (518-696) long by 782
(712-851) wide, divided by five septa into one central and five pe-
ripheral depressions (loculi); margin of posthaptor a strong muscular
rim, surrounded by a delicate, scalloped marginal membrane 49.2 (46.5—
51.8) wide; armed with 3 pairs of dissimilar anchors and 14 marginal
hooks. First anchors large, robust, left of larger specimen 128 long and
right of larger specimen 122 long, with pointed external tips and blunt
internal tips and enlarged in the middle; second pair of anchors elongate,
left of larger worm 75.1 long and right of smaller worm 65.3 long, more
robust than third pair with slightly recurved blunt tips; third pair of
anchors elongate, left of larger worm 81.8 long and right of smaller
worm 70.0 long, tips recurved and pointed. (Second and third pairs
of anchors lie side by side, arising at about the same level on the post-
haptor, being sometimes hard to distinguish from each other.) Post-
haptoral hooks 6.12 long. Small, round, disclike sclerites on posthaptor
near first pair of anchors.
Pharynx muscular, 168 (163-172) long by 215 (204-226) wide,
constricted (or notched) into five distinct lobes, papillated internally.
Mouth subterminal, ventral, immediately anterior to pharynx. Esophagus
very short; gut bifurcated, crura with medial and lateral dendritic
branching, not confluent posteriorly.
Two testes, juxtaposed, entire, fenestrated, oval in outline, left slightly
larger, 483 (471-494) long by 324 (319-330) wide, than right, 468
(454-482) long by 301 (293-309) wide, equatorial in position. Vasa
efferentia anastomising in midline to form vas deferens; vas deferens
proceeding anteriorly dorsal to left margin of ovary and left arm of
384 Proceedings of the Biological Society of Washington
vitelline reservoir and vagina whence it turns right and passes dorsal to
cirrus pouch, entering it near the proximal end. Vas deferens convoluted
from level of ovary to proximal end of vagina, where it straightens out.
Seminal vesicle a continuation of the vas deferens in the ventral side of
the cirrus. Cirrus obliquely situated just posterior to pharynx, 293 (281—
304) long by 70.7 (70.0-71.4) wide. Cirrus complex consisting of cirrus,
ejaculatory duct, and seminal vesicle in cirrus pouch. Prostate reservoir
separate from cirrus complex, on right side of body extending longi-
tudinally between proximal end of cirrus pouch and ovary, containing
strongly developed prostatic cells; duct passing from anterior end of
prostate reservoir around proximal end of cirrus and continuing ventrally
in cirrus. Prostatic cells around prostate reservoir except for left side.
Uterus connecting to cirrus pouch immediately at distal end of cirrus
forming a genital atrium opening outside via the common submarginal
genital pore on left above level of pharynx. Glands of Goto on each
side of the midline immediately posterior to testes.
Ovary pretesticular, oval, entire, median, 173 (153-192) long by 196
(163-229) wide, having internal chambers (= seminal receptacle of
Meserve, 1938) containing mature ova; oviduct passing from internal
chambers dorsal to right arm of vitelline reservoir and connecting to -
duct coming from right side of reservoir, proceeding anteriorly to ootype.
Ootype obliquely situated between cirrus and vagina, surrounded by
Mehlis’ gland cells; uterus short, opening into genital atrium. Vagina
muscular, passing anteriorly to distal end of ootype, turning left and
continuing to submarginal pore directly posterior to common genital
pore, probably connected via a small duct to anterior end of vitelline
reservoir.
Vitellaria follicular, extending from level of anterior pair of eyespots
to near posterior end of body. Transverse vitelloducts fusing medially
to form vitelline reservoir immediately anterolateral to ovary. Transverse
vitelloduct separating ovary from testes. Egg in utero polyhedral, 136
long by 125 wide, with convoluted basal filament.
Brain anterior to pharynx; three pairs of nerves passing into pro-
haptoral region. Four granular eyespots located dorsal to brain, first
pair smaller and closer together than posterior pair. Excretory pores
posterior to level of proximal end of vagina, opening dorsolaterally.
Allosprostonia new genus
Diagnosis: Trochopodinae. Body elliptical, flattened dorsoventrally.
Prohaptor a pair of anterolateral suckers united anterodorsally by a
common hood. Prohaptor subsessile, with scalloped marginal membrane;
ventral surface divided by five septa into central loculus and five pe-
ripheral loculi. Three pairs of dissimilar anchors (posterior two pairs, the
second and third anchors, lying side by side) and 14 marginal hooks.
Two pairs of eyes. Pharynx constricted (or notched) into five distinct
lobes. Intestinal crura with medial and lateral dendritic branches, not
Southern Pacific Trematodes Pt. V 385
confluent posteriorly. Two testes, fenestrated, juxtaposed. Vas deferens
joining seminal vesicle in cirrus pouch. Prostatic reservoir strongly
developed, outside of and connected to cirrus pouch. Cirrus short. Com-
mon genital atrium opening near left margin of body below left pro-
haptor. Ovary oval, median, pretesticular, separated from testes by a
transverse vitelloduct. Ootype surrounded proximally by Mehlis’ gland
cells; uterine egg with a single, convoluted polar filament. Vagina fairly
long, tubular, slightly swollen proximally, and probably united with
vitelline reservoir by a narrow duct; vaginal opening close to genital pore
submarginally. Vitellaria co-extensive with intestinal branches; vitelline
reservoir large, immediately anterolateral to ovary. Parasitic on marine
teleosts.
Type Species: Allosprostonia tauvinae n. sp.
Discussion: Allosprostonia n. gen. varies from every other group of the
subfamily Trochopodinae (Price, 1936) Sproston, 1946 in characters
that are presently regarded as generic in rank: (1) posthaptor divided
by five septa into one central and five peripheral loculi; (2) prohaptor
consists of a pair of anterolateral suckers united anterodorsally by a
common hood; (3) genital openings slightly submarginal; and, (4)
pharynx constricted into five lobes.
Allosprostonia tauvinae n. sp. is apparently related to Sprostonia
Bychowsky, 1957 in that it also possesses: (1) a pair of anterolateral
suckers united anterodorsally by a common hood; (2) strongly developed
prostate reservoir lying longitudinally between the cirrus and the ovary;
and, (3) three pairs of anchors similar in shape and location (the
posterior two pairs, the second and third anchors, lying side by side).
It is significant that Allosprostonia tauvinae n. sp. was found on the
same host as Sprostonia longiphallus n. sp. Although Macrophyllida
antarctica (Hughes, 1928) Johnston, 1929 also has five septa, these are
more weakly developed (see redescription below) and the anchors,
rather than being in the posterolateral septa, are in the posterior loculus
on obsolescent radii devoid of special musculature (Johnston, 1930a).
The five septa of Allosprostonia tauvinae n. sp. are considered homolo-
gous to the five septations of Sprostonia Bychowsky, 1957 which originate
around the central loculus. Because of the apparent close relationship
between Sprostonia Bychowsky, 1957 and the present species, the name
Allosprostonia tauvinae n. gen., n. sp. is proposed.
Two parasites were collected from seven host specimens (Table 6).
Genus Macrophyllida Johnston, 1929, emend.
Synonym: Macrophylla Hughes, 1928.
Diagnosis: Trochopodinae. Diagnosis that of Yamaguti (1963) except
as follows: (1) posthaptor bearing three pairs of anchors; anterior pair
at junction of posterior septa with that surrounding central loculus, two
posterior pairs in large posterior loculus; (2) vasa efferentia coming
from anterior margin of right testis and posterior margin of left; (3)
glands of Goto present to left and posterior to right testis; (4) cirrus,
386 Proceedings of the Biological Society of Washington
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25
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26
©
2
°o
22
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Ficures 21-26. Macrophyllida antarctica (Hughes, 1928) Johnston,
1929. (Scale is in mm.) 21, Whole mount, ventral view (posthaptor
twisted bringing posterior anchor-bearing loculus slightly out of align-
ment in the clockwise direction). 22, Reproductive organs and terminal
genitalia, ventral view. 23, Left first anchor. 24, Right second anchor.
25, Right third anchor. 26, Right third anchor.
Southern Pacific Trematodes Pt. V 387
uterus, and vagina opening close together in marginal depression on
left at posterior level of pharynx; and, (5) vaginal opening separate
from, but close to, common genital pore.
Type Species: Macrophyllida antarctica (Hughes, 1928) Johnston,
1929.
Discussion: The above emendation was made in order to correct the
generic diagnosis of Macrophyllida Johnston, 1929, as given by Yamaguti
(1963).
Hughes (1928) described a new genus and species, Macrophylla
antarctica, from the gills of the Gummy Shark, Mustelus antarcticus,
from Port Phillip Bay, Victoria, Australia. She found this species on only
two host specimens of about 100 examined. As the generic name was
found to be preoccupied, Macrophyllida was proposed in its place by
Johnston (1929), who later (1930a) redescribed and figured the species
from specimens sent to him by Dr. O. W. Tiegs.
Macrophyllida antarctica (Hughes, 1928) Johnston, 1929
Figs. 21-26
Synonym: Macrophylla antarctica Hughes, 1928.
Host: Mustelus antarcticus Gunther, Gummy Shark; family Galeo-
rhinidae.
Locality: Dunalley, Tasmania; 30 statute miles ENE of Hobart (15
fathoms, weed/mud).
Previously Reported Host and Locality: Mustelus antarcticus Gunther;
Port Phillip Bay, Victoria, Australia.
Gear Used: Danish seine.
Location: Gills.
Number Studied: 2.
Number Measured: 1.
Homoeotype: USNM Helm. Coll. No. 71204.
Redescription: Body elongate, flattened dorsoventrally, 5,510 long by
1,250 wide. Cuticle fairly thin and smooth. Anterior end broadly
rounded, separated from rest of body by constrictions on each side at
level of midpoint of pharynx, with pair of extensive glandular areas
almost meeting in front, extending backwards around pharynx to near
its posterior margin, and containing numerous ducts; greatest width of
prohaptoral region 1,220. Posthaptor a subsessile, concavo-convex, nearly
circular sucker, opening ventrally, diameter 962, divided by five weakly
developed septa into one central and five peripheral loculi; margin of
posthaptor a weak muscular rim surrounded by marginal membrane up
to 111 wide; armed with 3 pairs of dissimilar anchors, first pair at
junction of posterolateral septa with that surrounding central loculus,
and the second and third pairs of anchors located in the large posterior
loculus; and 14 marginal hooks. First anchors minute, left 9.59 long,
external tips blunt, internal tips bifid; second pair of anchors elongate,
slender, left 82.0 long and right 79.4, with blunt slightly recurved tips;
388 Proceedings of the Biological Society of Washington
third pair of anchors elongate, wider than second, right 61.8 long, with
recurved sharp tips. (Posthaptor with large cell between second and
third anchors.) Posthaptoral hooks 10.8 long, each with a domus.
Pharynx muscular, 329 long by 403 wide, margin partially constricted,
containing long, slender cells internally. Mouth subterminal, ventral,
anteroproximal to pharynx. Esophagus short; gut bifurcated, crura with
only lateral dendritic branching, not confluent posteriorly.
Testes two, oval, entire, obliquely tandem, left smaller, 210 long by
214 wide, diagonally anterior to right, latter 210 long by 223 wide,
preequatorial in position. Vasa efferentia emerging from anteromedial
margin of right testis, posterior margin of left, and fusing to right of
left at its posteromedial margin to form vas deferens. Latter running
to left of ovary, dorsal to vagina and distal end of ootype, then right to
pass dorsally to cirrus (entrance of vas deferens in cirrus not seen).
Ejaculatory duct (origin not evident) running inside cirrus to its tip.
Cirrus subobliquely situated between right crus of intestine and ootype.
Cirrus complex consisting of a sclerotized cirrus and prostate reservoir
at proximal end of cirrus in cirrus pouch. Several large prostatic cells
outside cirrus pouch, connected by small ducts to prostate reservoir.
Uterus connecting to cirrus pouch very near left margin, forming genital .
atrium opening outside via common marginal genital pore on left at level
of posterior margin of pharynx. Glands of Goto oval, large, situated
next to and slightly dorsal to right testis.
Ovary pretesticular, entire, nearly circular except for anterior extension,
309 long by 287 wide, having internal chamber (= seminal receptacle
of Meserve, 1938) containing mature ova; convoluted oviduct passing
from internal chamber to proximal end of ootype. Ootype obliquely
situated between cirrus and vagina, surrounded by Mehlis’ gland cells;
uterus thick-walled, tapering distally, opening into genital atrium. Vagina
long, slender, looped immediately anterolateral to ovary, containing duct
with constriction at level of distal end of ootype (constriction apparently
surrounded by circular muscles and conceivably functioning as a sphinc-
ter); opening proximal to genital pore into a common ventral marginal
depression having a muscular vaginal lip.
Vitellaria follicular, extending from near posterior level of pharynx
almost to end of body, smaller in intercecal field than in extracecal.
Vitelline reservoir(s) not observed. Egg not observed.
Brain anterior to pharynx. Four granular eyespots dorsal to brain,
first pair smaller and closer together than posterior pair. Excretory pores
opening dorsolaterally near margin at posterior level of pharynx.
Discussion: The worms in this collection appear conspecific with
Macrophyllida antarctica (Hughes, 1928) Johnston, 1929.
Though smaller than those in the original description (Table 4), the
worm measured is considered mature due to the presence of mature ova.
The worms in the present collection differ from Hughes’ (1928) descrip-
tion in the following: (1) presence of large glands of Goto; (2) posses-
sion of a small anterior pair of anchors on the posthaptor; (3) vas
Southern Pacific Trematodes Pt. V 389
TaBLE 4. Comparison of measurements of Macrophyllida antarctica
(Hughes, 1928) Johnston, 1929.
Hughes Johnston Present
(1928) (1930a) sample
Entire body (L) 13,150 14,800 5,510
(Ww) 1,300-2,500 2,800 1,250
Posthaptor — 2,000 962,
Marginal membrane (W) —— — Tee
Anchor 1 (L) ——= ——— L 9.59
Anchor 2 (L) —. 95 Le oo
R 79.4
Anchor 3 (L) ——. — R 61.8
Hooks (L) ——. —- 10.8
Prohaptoral region (W) — —— 1,220
Pharynx (L) —_— 700 329
(W) ——. 850 403
Testes L-(L) —— 600 210
(W) eG 360 214
R-(L) —. 600 210
(Ww) ease 450 293
Ovary (L) —— 600 (dia. ) 309
(W) — —. 287
Egg (L) 220 ——. —_—
(W) 107 —
(L) = Length. (W) = Width. L= Left. R = Right. (dia.) = diameter.
efferens from left testis coming from the posterior margin instead of the
anterior; and, (4) the presence the 14 marginal hooks, each having a
domus (Llewellyn, 1963). Hughes (1928) also illustrated Laurer’s
canals near the posterior end of the body proper in her figure. These,
in reality, are lateral branches of the gut and not Laurer’s canals, which
are found in digenetic trematodes. Johnston (1930a) mentioned the
other obvious mistake of Hughes (1928), who mistook the uterus for
the vagina, but failed to mention items 1, 2, and 4 above in his redescrip-
tion. Both Hughes (1928) and Johnston (1930a) indicated that the vas
efferens emerges from the anterior margin of the left testis. In our
specimens it comes from the posterior margin. The connection of the
vas efferens to the left testis in this fashion can best be explained by a
rotation of this testis 180 degrees to the right, resulting in the tandem
arrangement of the testes. This is further evidenced by the left gland
of Goto moving forward to lie next to the right testis and posterior to
the left.
The nature of the genital openings as described Johnston (1930a)
and Yamaguti (1963) is not clear. According to Johnston (1930a) there
are three sex apertures. Yamaguti (1963) wrote “cirrus, uterus and
390 Proceedings of the Biological Society of Washington
Mediavagina forsteri n. gen., n. sp. (Scale is in mm.)
FicureEs 27-35.
27, Whole mount
ventral view (posthaptor twisted bringing posterior
2
2
28
loculus about 125° out of alignment in the clockwise direction).
Reproductive organs and terminal genitalia, ventral view. 29, Left first
anchor. 30, Right first anchor. 31, Left second anchor. 32, Right second
anchor. 33, Left third anchor. 34, Right third anchor. 35, Egg in utero,
ventral view.
Southern Pacific Trematodes Pt. V 391
vagina opening close together into muscular common genital atrium
which in turn opens on the left margin.” In the two worms of the
present collection the genital atrium is formed by the junction of the
cirrus pouch and the uterus close to the left margin. The vagina opens
near but posterior to and separate from the genital pore in the depression.
The common genital pore is directed somewhat posteriorly and the
vaginal pore anteriorly. A single vaginal lip (Johnston, 1930a stated
that two such lips occurred in his sample) is situated on the anterolateral
edge of the depression.
The presence of the large cells between the posterior two pairs of
anchors as mentioned by Johnston (1930a) is verified in this study.
Johnston stated that this cell was probably a multipolar nerve cell. Its
function is unknown. Johnston also noted a resemblance of the long
gland cells of the pharynx with those described by Heath (1902) from
Entobdella squamula (Heath, 1902) Johnston, 1929. They have a large
nucleus and a granular protoplasm which stains well.
Sproston (1946) mentioned “the two posterior radii of typical Cap-
salids being virtually suppressed” in referring to those radii on which the
posterior two pairs of anchors are located. Johnston (1930a) wrote that
the disc of M. antarctica (Hughes, 1928) Johnston, 1929, has five
peripheral depressions and the “posterior loculus is large and is crossed
by two obsolescent radii devoid of special musculature.” Bychowsky
(1927) considered these rays to represent the disappearing posterior
septa of the rest of Megalocotyle as supported by the presence of anchors
in them. We accept Bychowsky’s theory.
Two parasites were recovered from 17 host specimens (Table 6).
This monotypic genus occurs on Mustelus antarcticus of the family
Galeorhinidae.
Mediavagina forsteri new genus, new species
Figs. 27-35
Host: Latridopsis forsteri (Castelnau), Silver Trumpeter or Bastard
Trumpeter; family Latridae.
Locality: Hobart, Tasmania; 25 statute miles E of Hobart (2-3
fathoms, mud).
Gear Used: Gill net.
Location: Gills.
Number Studied: 14.
Number Measured: 11.
Holotype: USNM Helm. Coll. No. 71205.
Paratypes: USNM Helm. Coll. No. 71206 (two specimens).
Description: Body elliptical, flattened dorsoventrally, (9) 1,880
(1,490-2,340), S = 307, SE = 102, CL = 236 long by (9) 573 (356-
712), S = 135, SE = 44.9, CL = 104 wide. Cuticle fairly thin and
smooth. Prohaptor a pair of slightly pedunculated ventrolateral suckers,
united in the median prepharyngeal region; left sucker (8) 169 (139-
206), S = 23.5, SE = 8.32, CL = 19.7 long by (8) 175 (130-210),
392 Proceedings of the Biological Society of Washington
TABLE 5. Comparison of measurements of Mediavagina species.
M. foresteri n. sp. M. macropteri n. sp.
Entire body (L) 1,490-2,340 2,560-3,140
(W) 356-712 518-699
Posthaptor (L) 358-720 568-756
(W) 360-746 605-766
Marginal membrane (W) 2.4,5-55.5 59.3-63.3
Anchor 1 L-(L) 26.7-40.8 34.2—38.0
R-(L) 26.1-37.3 31.0-39.0:
Anchor 2 L-(L) 46.9-74.9 58.9-72.8
R-(L) 50.0-76.5 54.9-66.0
Anchor 3 L-(L) 38.6-51.4 37.3-44.3
R-(L) 37.3-51.8 —-
Prohaptoral suckers L-(L) 139-206 258-260
(Ww) 130-210 213-239
R-(L) 151-197 256-282
(Ww) 120-216 218-265
Pharynx (L) 121-265 190-226.
(Ww) 114-265 175-183
Testes: Anterior (L) 112-240 97.9-129
(W) 117-255 76.2-114
Posterior (L) 111-211 90.1-142
(W) 100-224 ZALMAN OIL
Ovary (L) 73.6-143 155-196
(W) 92.6-168 140-172
Egg (L) 108-126 112-149
(W) 86.5-105 88.4-115
(L) = Length. (W) = Width. L= Left. R = Right.
S = 23.7, SE = 8.37, CL = 19.8 wide, right sucker (8) 169 (151-197),
Se AGAR SE 75 OS Cl 13:5 longs byn(9)) pal on CL 0—O1G) sas —
99.8, SE = 9.94, CL = 22.9 wide. Region before posterior border of
pharynx glandular. Posthaptor a subsessile, concavo-convex, oval sucker,
opening ventrally, (10) 565 (358-720), S = 111, SE = 35.2, CL =
79.6 long by (10) 624 (360-746), S = 138, SE = 43.7, CL = 98.8
wide, divided by five weakly-developed septa into one central and five
peripheral depressions (loculi); margin of posthaptor a muscular rim
surrounded by a delicate marginal membrane (10) 42.1 (24.5-55.5),
S = 11.9, SE = 3.77, CL = 8.53 wide; armed with 3 pairs of dissimilar
anchors and 14 marginal hooks. First anchors fairly small, robust, left
Southern Pacific Trematodes Pt. V 393
(7) 31.1 (26.7-40.8), S = 5.28, SE = 2.00, CL = 4.89 long and right
(4) 32.8 (26.1-37.3) long, with external tips knobbed and pointed,
internal bifid; second pair elongate, with slightly recurved blunt tips,
left (6) 62.1 (46.9-74.9), S = 9.08, SE = 3.71, CL = 9.53 long and
right (6) 63.8 (50.0-76.5), S = 9.53, SE = 3.89, CL = 10.0 long, more
robust than third pair; third anchors elongate, left (4) 45.6 (38.6—-51.4)
long and right (4) 45.0 (37.3-51.8) long, tips recurved and pointed.
Pharynx muscular, (11) 213 (121-265), S = 50.0, SE = 15.1, CL =
33.6 long by (11) 195 (114-265), S = 49.7, SE = 15.0, CL = 33.4
wide. Mouth subterminal, immediately anteroventral to pharynx. Esoph-
agus short; gut bifurcated, crura with medial and lateral dendritic
branching, not confluent posteriorly.
Testes two, tandem, entire, slightly fenestrated, oval in outline, anterior
Gielp) G2 Cl2=940) Si 3285 SE 1979s Clie 2 leSalongy bye (@luly)
181 (117-255), S = 35.3, SE = 10.6, CL = 23.7 wide, posterior (11)
5570 CSOT) Se OOM Sh Sai) Cle 195) long bya Cll) 168
(100-224), S = 32.6, SE = 9.82, CL = 21.9 wide, equatorial in posi-
tion. Vasa efferentia not observed. Vas deferens traced from dorsal to
left side of vitelline reservoir forward, proceeding right and turning left
just posterior to ootype, becoming convoluted, extending to distal end
of ootype where it straightens, then executes a wide loop, passing dorsal
to ootype and entering cirrus dorsally. Cirrus obliquely situated just
posterior to pharynx. Cirrus complex consisting of cirrus, seminal vesicle,
ejaculatory duct, and prostate reservoir in cirrus pouch. Prostatic cells
around prostate reservoir. Uterus connecting to cirrus pouch near left
margin, opening to genital atrium via common marginal genital pore on
left at posterior level of pharynx. Glands of Goto in midline, variable
in position, either posterior to, or to right of, posteriormost testis.
Ovary pretesticular, oval, entire, median, (11) 117 (73.6-143), S =
24.1, SE = 7.26, CL = 16.2 long by (11) 138 (92.6-168), S = 23.9,
SE =7.19, CL = 16.0 wide, having internal chamber (= seminal re-
ceptacle of Meserve, 1938) containing mature ova; oviduct passing from
internal chamber dorsal to right arm of vitelline reservoir and sinuously
to ootype. Connection of vitelline reservoir to oviduct not observed.
Ootype obliquely situated immediately posterior to cirrus pouch, sur-
rounded by Mehlis’ gland cells; uterus short, opening into genital atrium.
Vaginal pore between vitelline reservoir and ovary, median in position.
Vaginal tube not observed.
Vitellaria large and follicular, extending from posterior level of pharynx
to near posterior end of body. Transverse vitelloducts fusing medially
to form vitelline reservoir immediately anterolateral to ovary. Egg in
utero polyhedral, (7) 121 (108-126), S = 6.24, SE = 2.36, CL = 5.77
long by (7) 98.9 (86.5-105), S = 6.43, SE = 2.43, CL = 5.94 wide,
with convoluted basal filament.
Brain dorsal to anterior part of pharynx. Four granular eyespots
located dorsal to brain, first pair smaller and closer together than poste-
rior pair.
394 Proceedings of the Biological Society of Washington
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Ficures 36-43. Mediavagina macropteri n. sp. (Scale is in mm.)
36, Whole mount, ventral view. 37, Reproductive organs and terminal
genitalia, ventral view. 38, Left first anchor. 39, Right first anchor.
40, Left second anchor. 41, Right second anchor. 42, Left third anchor.
43, Egg in utero, ventral view.
Southern Pacific Trematodes Pt. V 395
Mediavagina macropteri new species
Figs. 36-43
Host: Nemadactylus macropterus (Bloch and Schneider), Jackass Fish
or Tarakihi; family Cheilodactylidae.
Locality: Lakes Entrance, Victoria; 45 statute miles ESE of Lakes
Entrance (65-75 fathoms).
Gear Used: Danish seine.
Location: Gills.
Number Studied: 5.
Number Measured: 4.
Holotype: USNM Helm. Coll. No. 71207.
Paratypes: USNM Helm. Coll. No. 71208 (two specimens).
Description: Body elliptical, flattened dorsoventrally, (3) 2,800
(2,560-3,140) long by (3) 587 (518-699) wide. Cuticle fairly thin
and smooth. Prohaptor a pair of slightly pedunculated ventrolateral
suckers, united in median prepharyngeal region; left sucker (2) 259
(258-260) long by (2) 226 (213-239) wide, right sucker (2) 269
(256-282) long by (2) 241 (218-265) wide. Region anterior to poste-
rior border of pharynx glandular. Posthaptor a subsessile, concavo-
convex, oval sucker, opening ventrally, (4) 674 (568-756) long by (4)
689 (605-766) wide, divided by five weakly-developed septa into one
central and five peripheral depressions (loculi); margin of posthaptor
a muscular rim, surrounded by a delicate marginal membrane, (2) 61.3
(59.3-63.3) wide; armed with 3 pairs of dissimilar anchors and 14
marginal hooks. First anchors fairly small, robust, left (4) 36.3 (34.2-
38.0) long and right (4) 35.9 (31.0-39.0) long, external tips knobbed,
internal tips bifid; second pair elongate, left (3) 65.0 (58.9-72.8) long,
right (3) 61.1 (54.9-66.0) long, more robust than third pair, tips
slightly recurved and blunt; third pair of anchors elongate, left (2) 40.8
(37.3-44.3) long, tips recurved and pointed.
Pharynx muscular, (3) 205 (190-226) long by (3) 178 (175-183)
wide. Mouth subterminal, immediately anteroventral to pharynx. Esoph-
agus short; gut bifurcated, crura with medial and lateral dendritic
branching, not confluent posteriorly.
Two tandem testes, entire, oval in outline, anterior (4) 114 (97.9-129)
long by (4) 87.8 (76.2-114) wide, and posterior (4) 117 (90.1-142)
long by (4) 83.8 (71.1-101) wide, equatorial in position. Vasa ef-
ferentia not observed. Vas deferens traced from dorsal to left side of
vitelline reservoir forward, looping just posterior to ootype, passing
sinuously to left of ootype before running dorsal to ootype and entering
cirrus dorsally. Seminal vesicle a continuation of the vas deferens in the
ventral side of the cirrus. Cirrus obliquely situated just posterior to
pharynx. Cirrus complex consisting of cirrus, seminal vesicle, and prostate
reservoir in cirrus pouch. Prostatic cells around prostate reservoir. Small
duct from prostate reservoir leading to large duct in cirrus passing dorsal
to seminal vesicle and leading to end of cirrus. Uterus connecting to
396 Proceedings of the Biological Society of Washington
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cirrus pouch near left margin forming a genital atrium opening via the
common marginal genital pore on left at level of posterior border of
pharynx. Glands of Goto in midline, slightly to the right and posterior
to posteriormost testis.
Ovary pretesticular, oval, entire, median, (4) 169 (155-196) long by
(4) 151 (140-172) wide, having internal chamber (= seminal receptacle
of Meserve, 1938) containing mature ova; oviduct passing from internal
chamber dorsal to right arm of vitelline reservoir and making a loop on
left of ootype before entering it. Connection of vitelline reservoir to
oviduct not observed. Ootype obliquely situated immediately posterior
to cirrus pouch, surrounded by Mehlis’ gland cells; uterus short, opening
into genital atrium. Vaginal pore just anterior to vitelline reservoir,
median in position. Vaginal tube not observed.
Vitellaria large and follicular, extending from posterior level of pharynx
to near posterior end of body. Transverse vitelloducts fusing medially
to form vitelline reservoir immediately anterosinistral to ovary. Main
vitelloducts very large, confluent anteriorly behind pharynx. Egg in utero
polyhedral, (6) 129 (112-142) long by (6) 104 (88.4115) wide,
with convoluted basal filament. Eggs occurring either singly or doubly
in ootype-uterus complex.
Brain anterodorsal to pharynx. Four granular eyespots located dorsal
to brain, first pair smaller and closer together than posterior pair.
Mediavagina new genus
Diagnosis: Trochopodinae. Body elliptical, flattened dorsoventrally.
Prohaptor a pair of slightly pedunculated anterolateral suckers. Post-
haptor subsessile, with delicate marginal membrane; ventral surface
divided by five weakly-developed septa into central loculus and five
peripheral loculi. Three pairs of dissimilar anchors on posterior two
septa, 14 marginal hooks. Two pairs of eyes. Pharynx globular. Intes-
tinal crura with medial and lateral dendritic branches, not confluent
posteriorly. Testes two, tandem, equatorial in position. Vas deferens
convoluted, making large loop on right just posterior to ootype, joining
seminal vesicle in cirrus pouch. Prostatic reservoir inside cirrus pouch.
Cirrus fairly short. Common genital atrium opening on left margin at
posterior level of pharynx. Ovary pretesticular, oval, entire, median.
Ootype surrounded proximally by Mehlis’ gland cells; uterine egg with
a single, convoluted polar filament. Vaginal pore opening ventrally in
midline in region of vitelline reservoir. Vitellaria follicular, co-extensive
with intestinal branches; vitelline reservoir immediately anterosinistral to
ovary. Parasitic on marine teleosts.
Type Species: Mediavagina forsteri new species.
Discussion: Mediavagina n. gen. varies from every other genus of the
subfamily Trochopodinae (Price, 1936) Sproston, 1946 in characters that
are presently regarded as generic in rank. These characters are: (1)
posthaptor divided by five weakly-developed septa into one central and
398 Proceedings of the Biological Society of Washington
five peripheral loculi; (2) prohaptor consists of a pair of slightly pedun-
culated anterolateral suckers; (3) vaginal pore located medially in the
region of the vitelline reservoir; and, (4) testes tandem. Due to the
above differences the name Mediavagina n. gen., concerning the location
of the vaginal pore, is proposed.
Mediavagina n. gen. is apparently most closely related to Macrophyllida
Johnston, 1929 in that it possesses: (1) five weakly-developed septa,
(2) tandem testes and, (3) prostate reservoir inside cirrus pouch.
The two new species of this new genus, M. forsteri n. sp. and M.
macropteri n. sp., possess differences besides different hosts that are
considered specific in rank. M. macropteri differs from the type species,
M. forsteri, in the following (Table 5): (1) body larger; (2) testes
smaller; (3) egg larger; (4) uterus sometimes with two eggs; and, (5)
vaginal pore anterior to vitelline reservoir instead of posterior.
Fourteen specimens of Mediavagina forsteri were recovered from 21
specimens of Latridopsis forsteri (Castelnau) (Table 6). A total of 5
specimens of Mediavagina macropteri were collected from 31 specimens
of Nemadactylus macropterus (Bloch and Schneider) (Table 6).
This fifth of a series on Monogenea from fishes of the
southern Pacific Ocean discusses six species from five host
species collected in Australian waters. Four monogeneids,
Sprostonia longiphallus, Allosprostonia tauvinae, Mediavagina
forsteri, and M. macropteri, are described for the first time.
In addition, Trochopus hobo Yamaguti, 1942 and Macrophyl-
lida antarctica (Hughes, 1928) Johnston, 1929 are redescribed.
Two new genera, Allosprostonia and Mediavagina are de-
scribed. Allosprostonia has been proposed to include those
Trochopodinae with five posthaptoral septa and paired pro-
haptoral suckers united by a common hood. Mediavagina is
necessary to accommodate those species having five posthap-
toral septa, tandem testes, and a median vagina. The sub-
family Trochopodinae (Price, 1936) Sproston, 1946 is emended
to accommodate members of the genus Allosprostonia since
they resemble other Trochopodinae more closely than other
groupings in the family Capsalidae.
New locality records are herewith established for Trochopus
hobo Yamaguti, 1942 and Macrophyllida antarctica (Hughes,
1928) Johnston, 1929.
LITERATURE CITED
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Southern Pacific Trematodes Pt. V 399
BRINKMANN, A., Jk. 1940. Contribution to our knowledge of the mono-
genetic trematodes. Bergens Mus. Aarb. Naturvitenskaplig-
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1952. Some Chilean monogenetic trematodes. Repts. Lund
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BycHuowsky, B. E. 1957 (1961). Monogenetic trematodes, their sys-
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Corpero, E. H. 1944. Dos nuevos especies de trematodos mono-
genéticos de los plagidstomos de la costa Uruguaya. Cali-
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GraHAM, D. H. 1956. A treasury of the New Zealand fishes. A. H.
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1958. A revised, annotated list of terms useful for mor-
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Harcis, W. J., Jk., AND W. A. Ditton. 1965. Monogenetic trematodes
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400 Proceedings of the Biological Society of Washington
Heatu, H. 1902. The anatomy of Epibdella squamula sp. nov. Proc.
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Hucues, W. K. 1928. Some trematode parasites on the gills of Vic-
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. 1930a. The anatomy of the trematode, Macrophyllida ant-
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1930b. A new species of trematode of the genus Anoplo-
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1934a. New trematodes from South Australian elasmo-
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1937. Report on the Trematoda. Australasian Antarctic
Expedition 1911-1914. Scientific Reports, Series C, 10 (1): .
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Jounston, T. H., ano O. W. Tiecs. 1922. New gyrodactyloid trem-
atodes from Australian fishes, together with a reclassification
of the superfamily Gyrodactyloidea. Proc. Linn. Soc. N.S.
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melanesiensis n. sp. a monogenetic trematode from Fiji and
the New Hebrides. Can. J. Zool., 36 (2): 167-173.
LLEWELLYN, J. 1963. Larvae and larval development of monogeneans,
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Dawes, Academic Press, London and New York.
MaAcCatium, G. A. 1917. Some new forms of parasitic worms. Zoo-
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pathologica, 1: 137-204.
Manter, H. W. 1955. Two new monogenetic trematodes from ele-
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20 (2): 105-112.
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Vol. 81, pp. 403-412 30 August 1968
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
HELMINTH PARASITES OF ANTARCTIC
VERTEBRATES. PART IV. MONOGENETIC
TREMATODES FROM ANTARCTIC FISHES:
THE SUPERFAMILY CAPSALOIDEA PRICE, 19361
By Wiu1AM J. Harcis, JR. AND WiLLIAM ARTHUR DILLON?
Virginia Institute of Marine Science
Gloucester Point, Virginia
This paper on the Monogenea (Carus, 1863) is the fourth
in a series dealing with certain trematodes from fishes of
Antarctic waters. It treats three species belonging to the
superfamily Capsaloidea Price, 1936. The scope, organization,
and purpose are the same as for Part II of this series ( Hargis
and Dillon, in press).
Materials and Methods: Methods involved in the preservation and the
preparation of the Monogenea for identification and study are essentially
the same as those given by Dillon and Hargis (1965).
All measurements were made with the use of a calibrated filar mi-
crometer and are given in microns unless otherwise noted. In indicating
these measurements the mean is given, followed by the range (minimum
and maximum) in parentheses. The standard deviation (S), standard
error (Sz), and the interval estimate at the 95% level (t.osSz) follow the
range. For convenience the alphabetical symbols SE and CL are utilized
for the formal mathematical designations for standard error (Sz) and
1 This entire research is supported by grants GA-13853, with amendments, and
GA-235 under the United States Antarctic Research Program of the National Science
Foundation, to whom our thanks are due. The writers also wish to thank the follow-
ing individuals for their valuable contributions to this study: Dr. Willis L. Tressler
for the initial collection from Wilkes; Messrs. W. Stanley Wilson and William J.
Saunders, formerly of the Parasitology Section of the Virginia Institute of Marine
Science, for the collection of host materials from McMurdo and Wilkes; Dr. Harry
L. Holloway of Roanoke College for the collection of additional specimens from
McMurdo; Dr. Hugh DeWitt and Dr. Donald E. Wohlschlag for the identification of
host fishes.
Contribution No. 275 from the Virginia Institute of Marine Science, Gloucester
Point, Virginia.
2 Present address: University of Southem Mississippi, Southern Station, Box 1631,
Hattiesburg, Mississippi 39401.
40—Proc. Bio. Soc. WAsH., VoL. 81, 1968 (403)
404 Proceedings of the Biological Society of Washington
confidence limits or interval estimate at the 95% level (t.oSz), respec-
tively. The number of measurements used in the calculations appears
in parentheses before these data. Measurements of curved structures were
made across lines subtending the greatest arcs described by those struc-
tures. In the measurements to follow, length—of the body, its appen-
dages and most internal organs—refers to the distance along the
anteroposterior axis except where otherwise noted. Width refers to a
measurement made at right angles to the length, ie., along the dextro-
sinistral axis. The measurements of the lengths of cirri, genital ducts,
anchors and hooks were made along the longest axes of those structures
regardless of orientation.
Camera lucida drawings were used to facilitate identification and in
the preparation of the figures.
Station Locations: In general, collections at McMurdo Station were
made in nearby McMurdo Sound from shore or ice while those at Wilkes
Station were made in the waters around the Windmill Island group
through ice or from the Institute’s specially built research vessel, R/V
Octans. Actual locations are given immediately below for greater pre-
cision since the area is not widely known and the habitats from which
hosts were collected ranged from shelf to oceanic. The first letter in
each station number gives the base of operations (M = NAF McMurdo;
W = Wilkes Station), and the second letter (and sometimes a third
letter) gives the collecting locality. The station designation is followed
by: (1) general locality, and (2) precise location of each sampling
station. For trapping stations, latitude and longitude are presented.
Beginning as well as ending positions are provided for trawl runs. Also
presented are: (1) date of sampling, (2) type of sampling gear used,
(3) depth in fathoms at sampling location (if the sampling was done
over a straight course, the depth range is given) and, (4) a general
description of the nature of the bottom where known.
Station M-SA: McMurdo Sound. 77° 42'S, 166° 15’E. October
1959 to February 1960. Trap. Depth 150 fathoms.
Station M-SC: McMurdo Sound. 77° 48’S, 166° 30’ E. October
1959 to February 1960. Trap. Depth 300 fathoms.
Station M-M: McMurdo Sound. 77°51’S, 166° 38’E. October
1959 to February 1960. Trap. Depth 5 fathoms.
Station M-H: McMurdo Sound. 77° 55’ S, 166° 39’ E. During 1964—
1965.
STATION W-BA: Ramp Cove. 66° 17'S, 110° 32’ E. 12 January 1958.
Hook and line. Depth 2.5-3.3 fathoms.
Station W-C: Windmill Islands. 66° 16’00” S, 110° 31'00” E to
66° 16°00” S, 110° 31’ 34” E. 25 January 1961. Trawl. Depth 30-50
fathoms. Rock.
Station W-G: Windmill Islands. 66° 14'25”S, 110° 27'40” E to
66° 14’ 40” S, 110° 28’ 25” E. 13 February 1961. Trawl. Depth 25-35
fathoms. Rock.
STATION W-H: Windmill Islands. 66° 14’05”S, 110° 27'30” E to
Trematodes from Antarctic Fishes 405
66° 14’ 20” S, 110° 28’15” E. 13 February 1961. Trawl. Depth 25-35
fathoms.
StaTION W-I: Windmill Islands. 66° 13'12”S, 110° 27'45” E to
66° 13’ 30” S, 110° 28’25” E. 22 February 1961. Trawl. Depth 20-30
fathoms. Rock.
Station W-J: Windmill Islands. 66° 15’59”S, 110° 32'17” E. 10
March 1961. Trap. Depth 20 fathoms.
STATION W-K: Windmill Islands. 66° 15’52”S, 110° 34’ 26” E. 12
March 1961. Trap. Depth 10 fathoms.
STATION W-L: Windmill Islands. 66° 15’55” S, 110° 34/50” BE. 16
March 1961. Trap. Depth 12 fathoms.
STATION W-M: Windmill Islands. 66° 15’ 56” S, 110° 35’05” E. 2-3
June 1961. Trap. Depth 9 fathoms.
Station W-O: Windmill Islands. 66° 15’50”S, 110° 35’01” E. 1-3
September 1961. Trap. Depth 7 fathoms.
STaTION W-P: Windmill Islands. 66° 15’54”S, 110° 35’01” E. 5
September 1961. Trap. Depth 6 fathoms. Sand—rock.
Station W-Q: Windmill Islands. 66° 15’58”S, 110° 33’ 43” E. 6
September 1961. Trap. Depth 33 fathoms. Rock.
STaTION W-R: Windmill Islands. 66° 20’ 41” S, 110° 27'52” E. 11
October 1961. Trap. Depth 98 fathoms.
Station W-S: Windmill Islands. 66° 20'59”S, 110° 26’59” FE. 2
December 1961. Trap. Depth 200 fathoms. Mud.
StaTIon W-AO: Windmill Islands. 66° 14’ 20”S, 110° 34'06” E to
66° 14’ 16” S, 110° 34/14” E. 15 December 1961. Trawl. Depth 1-6
fathoms. Mud-rock.
ORDER MONOGENEA CARUS, 1863
SUBORDER MONOPISTHOCOTYLEA ODHNER, 1912
SUPERFAMILY CAPSALOIDEA PRICE, 1936
Faminty CapsALIpAE Barrp, 1853
Subfamily Trochopodinae (Price, 1936) Sproston, 1946
Discussion: The subfamily Trochopodinae was reviewed in a previous
paper (Dillon and Hargis, 1965, p. 241-243). Since that review we
have concluded that Bychowsky (1957) and Dollfus and Euzet (1964)
were correct in transferring Pseudobenedenia Johnston, 1931 from Bene-
deniinae Johnston, 1931 to this subfamily. Also, recent studies indicate
that the genus Trochopella Euzet and Trilles, 1962 is similar in every
taxonomically important feature to Trochopus Diesing, 1850 as charac-
terized by Yamaguti (1963). Therefore, Trochopella is a synonym of
Trochopus. With these changes the subfamily Trochopodinae, as pres-
ently understood, contains the following genera: Allomegalocotyla
Yamaguti, 1963, Macrophyllida Johnston, 1923, Megalocotyle Folda,
1928, Pseudobenedenia Johnston, 1931, Pseudomegalocotyla Yamaguti,
1963, Sprostonia Bychowsky, 1957, and Trochopus Diesing, 1850.
406 Proceedings of the Biological Society of Washington
Genus Pseudobenedenia Johnston, 1931
Diagnosis: Trochopodinae. Prohaptor consisting of head organs (or
glandular areas) and a pair of ventrolateral suckers. Posthaptor a con-
cavo-convex, circular disc, divided by septa into one central depression
and 6 (P. nototheniae Johnston, 1931) or 7 peripheral depressions;
armed with 3 pairs of anchors and 14 marginal hooks. Crura weakly
or strongly branched. Testes two, usually juxtaposed (may range from
juxtaposed to tandem position, as in P. shorti n. sp.). Cirrus complex
consisting of muscular cirrus, prostate reservoir, and seminal vesicle in
cirrus pouch. Vaginal aperture situated slightly to left of midline, near
anterior border of vitelline reservoir. Eyespots present or absent.
Discussion: The modified diagnosis of the genus Pseudobenedenia
given by Dollfus and Euzet (1964) is generally correct. Our specimens
of P. nototheniae Johnston, 1931 apparently lack the seventh (anterior-
most) septum, but due to the obscure septation in some capsalids, this
septum may exist. Extensive collections now being made in Antarctica
may settle this question.
Pseudobenedenia shorti new species
Figs. 1-10
Hosts: Trematomus bzrnacchii Boulenger, T. hasoni Boulenger, and
T. centronotus Regan; family Nototheniidae. Rhigophila dearborni
DeWitt (see below.); family Zoarcidae.
Location: Gills.
Localities: Trematomus bernacchii from stations M-SA, M-M, M-H,
W-C, W-G, W-H, W-I, W-J, W-L, W-M, W-O, W-P, W-R, W-S, and
W-AO. T. hansoni from stations M-SC, M-SA, M-M, M-H, W-K, W-L,
W-Q, and W-S. T. centronotus from stations M-H and W-S, and
Trematomus sp. from station W-BA. Rhigophila dearborni from station
M-H.
Number Examined and Studied: 1,183.
Holotype: USNM Helm. Coll. No. 70956.
Paratypes: USNM Helm. Coll. No. 70957 (4 specimens).
Description of Adult: Body elongate, cylindrical, (20) 2,150 (1,550-
2,520), S = 240, SE = 54, CL = 113 long by (20) 590 (450-710),
= 84, SE = 19, CL = 40 wide; cuticle fairly thin and smooth. Pro-
haptor consisting of head organs connected by ducts to cephalic glands
and a pair of ventrolateral, muscular suckers, (20) 253 (148-347), S =
66.3, SE = 14.8, CL = 31.0 long by (20) 246 (159-322), S = 60.6,
SE = 13.5, CL = 28.3 wide. Posthaptor a concavo-convex, circular disk,
opening ventrally, (17) 620 (400-780), S = 116, SE = 28, CL = 59
long by (17) 700 (490-920), S = 127, SE = 31, CL = 66 wide, divided
by septa into one central and 7 peripheral depressions; margin of post-
haptor a strong muscular rim, surrounded by a delicate membrane;
posthaptor armed with 3 pairs of anchors and 14 marginal hooks; post-
haptor slightly pedunculated. Anteriormost anchors (19) 127 (90-
Trematodes from Antarctic Fishes 407
181), S= 23.1, SE = 5.3, CL = 11.1 long, with bifid base, strong shaft
and rounded to pointed tips. (Tips of the anteriormost anchor pair are
spatulate in the onchomiracidium, becoming less spatulate during early
and late juvenile stages and more pointed in large adults, Figs. 4-8.)
Middle anchors (18) 260 (199-308), S = 28.8, SE = 6.8, CL = 143
long; posteriormost anchors (16) 203 (148-247), S = 25.3, SE = 6.3,
CL = 13.4 long. Marginal hooks (10) 14 (12-15), S = 0.8, SE = 0.25,
CL = 0.6 long. Tissues of posthaptor with disclike sclerites.
Mouth ventral, subterminal, opening into muscular buccal funnel; rim
of aperture serrated; wall of buccal funnel armed with conical papillae.
Pharynx (18) 159 (129-198), S = 19.8, SE = 4.7, CL = 9.9 long by
(18) 185 (156-216), S = 15.8, SE = 3.7, CL = 7.8 wide; esophagus
short or non-existent. Gut bifurcated; crura weakly branched and ap-
parently not confluent posteriorly.
Testes two, ranging from juxtaposed to tandem position, (12) 103
(87-122), S = 11.3, SE = 3.3, CL = 7.3 long by (12) 92 (83-118),
S = 4.7, SE = 1.4, CL = 3.1 wide. Vasa efferentia anastomosing in
midline to form vas deferens which proceeds anteriorly for a short
distance prior to dilating immediately behind ovary; then passing around
left side of ovary, crossing to right side of body and back to left side,
and finally forming numerous loops before entering cirrus pouch. Cirrus
complex consisting of muscular cirrus, prostate reservoir, and seminal
vesicle in cirrus pouch; cirrus pouch (10) 272 (216-315), S = 39.4,
SE = 12.5, CL = 28.3 long. Glands of Goto not observed. Cirrus and
uterus opening to outside via common genital pore, located along left
side of pharynx.
Ovary (13) 139 (92-231), S = 38.6, SE = 10.7, CL = 23.3 long by
(13) 119 (76-185), S = 35.3, SE = 9.8, CL = 21.3 wide; oviduct
convoluted, proceeding anteriorly to ootype. Ootype relatively large,
apparently surrounded by Mehlis’ glands; uterus extending diagonally
from ootype to genital atrium. Vaginal opening small, located slightly
to left of midline, near anterior border of vitelline reservoir; short vaginal
duct proceeding into vitelline reservoir. Vitellaria follicular, somewhat
large; transverse vitelloducts fusing medially to form vitelline reservoir.
Eggs fusiform, (10) 303 (277-310), S = 9.7, SE = 3.1, CL = 7.0 long
by (10) 114 (106-122), S = 5.7, SE = 1.8, CL = 4.1 wide (measure-
ment exclusive of filament), with a long filament at one pole. (The
filaments of the eggs are wrapped around the peduncle, which is behind
the posthaptor in a standard ventral view, Fig. 10. This unusual habit
has also been noted in Ancyrocotyle vallei by Parona and Perugia in
1895 and Parona and Monticelli in 1903.) Brain dorsal to oral aperture;
eyespots absent.
Description of Onchomiracidium (Fig. 9): Body elongate, cylindrical,
(2) 270 (252-288) long. Prohaptor with a pair of ventrolateral suckers
and head organs. Posthaptor well defined, circular, armed with three
pairs of anchors and 14 marginal hooks; anteriormost anchors (10) 23
(21-27) long, with spatulate tips wider than shaft or bifid roots; marginal
408 Proceedings of the Biological Society of Washington
Ficures 1-10. Pseudobenedenia shorti n. sp. 1, Whole mount, ventral
view. 2, Middle anchor. 3, Posteriormost anchors. 4-6, Anteriormost
anchors. 7, Anteriormost anchor from late juvenile (showing two dif-
ferent views). 8, Anteriormost anchor from early juvenile. 9, Oncho-
miracidium emerging from egg. 10, Showing filaments of eggs wrapped
around peduncle. (Scales are in millimeters. )
Trematodes from Antarctic Fishes 409
hooks (10) 14 (13-16) long, without accessory sclerotized processes.
Pharynx located in anterior %4 of body proper; gut not observed. Eye-
spots and cilia absent.
Etymology: This species is named in honor of Dr. Robert B. Short.
Discussion: Pseudobenedenia shorti n. sp. can be distinguished from
P. nototheniae, the type species, by: (1) general body shape; (2) shape
of the posteriormost anchors; and (3) smaller prohaptoral suckers,
posthaptor, pharynx, and testes.
Though we have indicated Rhigophila dearborni as a host of this
species it is likely that the record on which it is based is erroneous.
The single set of gills from which the worms were taken were separated
from the main body of the specimen after collection, but before exam-
ination, and given a number. Much later a carcass bearing the matching
number was identified as R. dearborni by Dr. Hugh DeWitt, then of
the University of Southern California at Los Angeles.
We believe that this is an erroneous host record because: (1) P.
shorti occurred on only one Rhigophila dearborni specimen though 101
of these fish were collected and examined in all, (2) a copepod, Clavel-
lodes intermedius (Quidor, 1906) Wilson, 1915, which Zwerner (1966)
believes to be specific for nototheniid fishes, was also collected from this
particular host, and (3) none of the other monogeneids or copepods
regularly occurring on R. dearborni were present on this specimen. If
this single record is eliminated, P. shorti has been reported only from
the several species of the genus Trematomus listed above.
Because the host record under discussion cannot be more positively
challenged and we do not wish to reinforce our own notions of host-
specificity by eliminating the record, it remains. It is hoped that later
collections will support our conclusions that Pseudobenedenia shorti
occurs only on species of the genus Trematomus.
Pseudobenedenia nototheniae Johnston, 1931
Host: Trematomus bernacchii Boulenger; family Nototheniidae.
Location: Skin.
Localities: Stations M-M and M-SA.
Previously Reported Hosts and Localities: Notothenia colbecki from
Antipodes Island (Johnston, 1931) and from the Island of Auckland
(see Dollfus and Euzet, 1964, p. 1, footnote 1). N. macrocephala (= N.
angustata) from Antipodes Island and Macquarie Island (Johnston,
1931). N. rossi collected in the Kerguelens (Dollfus and Euzet, 1964).
Number Examined and Studied: One adult; four juveniles.
Discussion: Examination of the one adult specimen and four immature
specimens from the skin of Trematomus bernacchii revealed that they
are probably conspecific with Pseudobenedenia nototheniae Johnston,
1931.
Careful studies of the posthaptors of all worms failed to reveal the
anteriormost septum which was described by Dollfus and Euzet (1964)
but not by Johnston (1931). Additional specimens are needed for study
410 Proceedings of the Biological Society of Washington
before we can challenge the description of Dollfus and Euzet in this
regard. More extensive collections are now being made in Antarctica
and a final decision on this matter may be available for inclusion in a
later work.
Pseudobenedenia sp.
Host: Trematomus borchgrevinki Boulenger; family Nototheniidae.
Location: Gills. i
Locality: Station M-H.
Number Studied: 253.
Discussion: The 253 juvenile specimens of an unidentified capsalid
recovered from T. borchgrevinki are tentatively assigned to the genus
Pseudobenedenia because of the glandular regions anterior to the pro-
haptoral suckers and the similarities of the anchors to those of P. shorti.
Pseudobenedenia sp. differs from the juveniles of P. shorti in the follow-
ing respects: (1) pharynx of juvenile specimens larger than that of the
juveniles (or adults) of P. shorti, (2) presence of two eyespots (eyespots
were absent in all stages of development in P. shorti), and (3) a slight
difference in the relative lengths of the roots of the anteriormost anchors.
Because of these differences we are convinced that this population
represents a species unknown to science. However, since all of the
worms are early juveniles whose anatomical features are not developed
sufficiently to permit more positive identification, confirmation of this
thesis must await acquisition of adult specimens.
LITERATURE CITED
BycHowsky, B. E. 1957. Monogenetic trematodes, their systematics
and phylogeny. [in Russian] Izdanie Akad. Nauk USSR,
Moskva-Leningrad, 509 p. (English Translation from the
Russian by P. C. Oustinoff, edited by W. J. Hargis, Jr., 627
p. American Institute of Biological Sciences, Washington,
D. C., 1961.)
DitLon, W. A., AND W. J. Harcis, Jr. 1965. Monogenetic trematodes
from the southern Pacific Ocean, 1. Monopisthocotyleids
from New Zealand fishes, p. 229-249. In Biology of the
Antarctic Seas II, Antarctic Res. Ser., 5.
Dou.rus, R. Pu., aNnp L. Euzet. 1964. Complement a la description
de Pseudobenedenia nototheniae T. H. Johnston, 1931
(Trematoda Monogenea) parasite d’un teleosteen du genre
Notothenia Richardson des Kerguelen. Bull. Mus. Nat. Hist.
Natur. 2° Ser., 36 (6): 849-857.
Harcis, W. J., JR.. AND W. A. Ditton. Helminth parasites of Antarctic
vertebrates, II, Monogenetic trematodes from Antarctic
fishes: The superfamily Gyrodactyloidea Johnston and Tiegs,
1922. (In press.)
Jounston, T. H. 1931. New trematodes from the subantarctic and
Antarctic. Aust. J. Exp. Biol. Med. Sci., 8: 91-98.
Trematodes from Antarctic Fishes 411
Parona, C., aND F. S. Monticetui. 1903. Sul genere Ancyrocotyle
(n.g.). Arch. Parasitol., 7 (1): 117-121.
Parona, C., AND A. PeruciA. 1895. Supra due nuove specie di Trem-
atodi ectoparassiti di pesci marini. Boll. Mus. Zool. Anat.
Comp. R. Univ. Genova, 31: 1-4.
YamacutTt, S. 1963. Systema Helminthum. IV. Monogenea and
Aspidocotylea. John Wiley & Sons, New York—London,
699 p.
ZWERNER, D. E. 1966. Parasitic copepods of some Antarctic and New
Zealand fishes. (Unpublished Master’s Thesis, School of
Marine Science, College of William and Mary, Virginia In-
stitute of Marine Science, Gloucester Point, Virginia, U.S. A.)
412 Proceedings of the Biological Society of Washington
Vol. 81, pp. 413-418 30 December 1968
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
A NEW FAXONELLA FROM NORTHEAST LOUISIANA
(DECAPODA, ASTACIDAE)
By Jerry G. WALLS
Department of Biological Sciences, McNeese State College,
Lake Charles, Louisiana
The genus Faxonella (Creaser, 1933) was diagnosed by
Fitzpatrick (1963), who recognized two species, F. clypeata
(Hay, 1899) and F. beyeri (Penn, 1950). The new species
here described brings to three the number of known species.
Two of these, F. beyeri and F. creaseri (new species), are
endemic to Louisiana.
I would like to thank Dr. Joe B. Black, McNeese State
College, for the loan of specimens and for his patience and
assistance in preparing this description. Thanks are also due
to Messrs. Norman Arnold and Frank Amsden for providing
transportation and assistance in collecting.
Faxonella creaseri new species
Diagnosis: Lateral rostral spines and cervical spines absent; rostrum
short and broad, one and one-half or fewer times as long as broad.
Shape of chela (Fig. 4) and carapace as in Faxonella clypeata and F.
- beyeri; areola variable, 25.9-33.1 percent of carapace length (average
30.0), and 2.3-5.3 times as long as broad (average 3.8). Male with
copulatory hooks on third pereiopods only. First pleopod of form I
male (Figs. 1-3) terminating in two rami: central projection long,
flattened, bent mesially; mesial process much shorter, only about one-
half as long as central projection, removed from and paralleling central
projection. In usual position, terminal elements of left pleopod overlap
those of other (Fig. 3). Annulus ventralis (Fig. 5) immovable, subovate,
with U-shaped depression between cephalic tubercles; caudal region
depressed, with contours as figured.
Holotypic male, form I: Body subovate, not depressed. Abdomen
narrower than thorax. Width of carapace at level of caudal margin of
cervical groove slightly less than height. Areola of moderate width,
41—Proc. Brot. Soc. Wasu., Vou. 81, 1968 (413)
414 Proceedings of the Biological Society of Washington
Tae a
nap SEIT te
Ficures 1-9. Faxonella creaseri new species. 1. Mesial view of first
pleopod of male, form I. 2. Caudal view of first pleopod of male, form
I. 3. Caudal view of first pleopods of male, form I, in situ. 4. Chela
of male, form II. 5. Annulus ventralis. 6. Mesial view of first pleopod
of male, form II. 7. Caudal view of first pleopod of male, form II.
8. Mesial view of first pleopod of male, form II, with shrunken mesial
process. 9. Caudal view of first pleopod of male, form II, with shrunken
mesial process.
New Louisiana Crawfish 415
TABLE 1. Measurements of Faxonella creaseri (in mm.).
Holotype Allotype Morphotype
Carapace
Height 6.7 8.0 6.0
Width 6.4 7.4 5.5
Length 14.5 18.1 14.0
Rostrum ;
Length 3.0 41 ay
Width Dd) 3.5 Bd)
Areola
Length 4.8 5.8 4.2
Width 0.9 15 1.0
Right chela (Left )
Length (outer margin) 8.5 9.0 7.0
Length (inner palm) 3.7 4.3 3.1
Width of palm 3.0 4.2 D5)
Length of dactyl 4.6 5.0 3.8
Abdomen, to tip of telson 16.2 18.8 Set
5.3 times longer than wide, with two or three punctations in nar-
rowest part; cephalic portion of carapace about 2.2 times as long as
areola; length of areola 33.1 percent of total length of carapace. Rostrum
without lateral spines; widest at base, very gradually converging toward
tip; margins raised but not thickened; no median carina present, but
distinct impressed median line visible toward tip; upper surface
distinctly concave. Acumen slightly upturned; short but distinct.
Postorbital ridge distinct, ending bluntly without spine. Branchiostegal
spine small, indistinct. Cervical area without spines, although low
tubercles present.
Antennule with small spine on ventromesial margin about midlength
of basal segment. Antenna of usual form but broken. Antennal scale
extending beyond tip of rostrum and closely resembling that of F.
clypeata and F. beyeri.
Chela with palm inflated; closely resembling that of F. clypeata and
F. beyeri in all respects; inconspicuous tubercles present on proximal
cutting edges of both fingers; fingers meeting evenly for most of length
when closed, gaping slightly at base.
Hooks on ischiopodites of third pereiopods only; hooks acute and as
long as greatest diameter of ischiopodites.
Abdomen 110 percent of carapace length; cephalic region of telson
with three spines in left and four spines in right caudolateral angles.
First pleopod (figs. 1-3) reaching almost to coxa of first pereiopod
when abdomen is flexed. Tip terminating in two rami as follows:
central projection corneous, long, slender, strongly compressed and
416 Proceedings of the Biological Society of Washington
flexible toward tip; tip recurved ventrally; mesial process noncorneous,
about one-half length of central projection, slender, with acute apex
and situated parallel to central projection but removed from mesial
edge of latter by at least width of mesial process. In usual position
(fig. 3) both rami are crossed, the mesial processes extending laterally
beyond lateral edges of central projections.
Allotypic female: Similar in most respects to holotype, but chela
proportionately shorter, broader, and more depressed; fingers stouter,
with distinct tubercles on cutting edges, gaping narrowly throughout
their length. Areola wider, 3.9 times as long as wide, with four to six
punctations in narrowest part.
Annulus ventralis (fig. 5) subovate with greatest length in transverse
axis; immovable except for slightly free lateral edges. Surface contours
as illustrated. Anterior margin produced into two conspicuous tubercles,
the larger strongly excavated on its caudal surface; tubercles not meeting
in midline, producing a U-shaped gap in cephalic view.
Morphotypic male, form II: Similar to holotype except for areola and
secondary sexual characters. Areola 30 percent of length of carapace
and 4.2 times as long as broad. Hook on third pereiopod blunt, length
only one-half maximum diameter of ischiopodite. First pleopod (Figs.
6-7) stouter, noncorneous; length and disposition of elements similar —
to holotype, except mesial process only one-quarter length of central
projection.
Measurements: See Table 1.
Type-locality: Louisiana, Caldwell Parish, roadside ditch 3.0 mi. N.
of Grayson on U.S. Hwy. 165.
Specimens examined: Thirty-two, all from Caldwell Parish, as follows:
3.9 mi. N. Caldwell-LaSalle Par. line on U.S. Hwy. 165, 22 July 1967,
J. G. Walls and N. Amold (16 I, 2é II, 19); 6.6 mi. N. Caldwell-
LaSalle Par. line on U.S. Hwy. 165, 22 July 1967, J. G. Walls and N.
Arnold (16 I, 4é Il, 42); 0.9 mi. N. Caldwell-LaSalle Par. line on
U.S. Hwy. 165 (La. Hwy. Dept. Rest Area), 29 July 1967, J. G. Walls
and F. Amsden (7¢ I, 4¢4 II, 59, including morphotype); type-
locality, 29 July 1967, J. G. Walls and F. Amsden (1¢ I, 29, including
holotype and allotype). All the above are designated paratypes.
Disposition of types: The holotype male, form I, allotype female, and
morphotype male, form II are deposited in the United States National
Museum (nos. 129215, 129216 and 129214, respectively). Paratypic
series consisting of one male, form I, one female and one male, form
II are deposited in the following collections: private collection of Dr.
J. F. Fitzpatrick, Jr., Mississippi State University; Tulane University;
Museum of Comparative Zoology; Instituto de Biologia (Mexico);
Australian Museum (Sidney); and the private collection of Dr. Joe
B. Black, McNeese State College. The remaining specimens are in the
collection of the author.
Color and habitat notes: Body translucent, pale brown with darker
New Louisiana Crawfish 417
specks; pair of grayish-brown stripes extending from behind eyes to
tip of telson. Stripes bordered laterally on abdomen by stripes of vivid
reddish-brown. Lateral surfaces of carapace grayish, almost white.
Chela tinged with light orange. Some specimens with patch of reddish-
brown on cephalolateral region of carapace.
All specimens were taken from temporary pools in ditches or culverts.
Vegetation, if present, was sparse and consisted of roadside grasses.
Associated species were Procambarus acutus (Girard, 1852), Procambarus
tulanei Penn (1953), and an undescribed subspecies of Cambarus
hedgpethi Hobbs (1948).
Variations: The specimens agree in most significant respects. Greatest
variation is shown in the width of the areola, which varies from 2.3
to 5.3 times as long as wide, averaging 3.8 times. When more speci-
mens have been taken it may be shown that this variation is correlated
with age and/or sex. Size range of carapace length (in mm.): ¢ 4,
13.5-17.6 (av. 15.8); 99, 12.8-18.6 (av. 15.0). Some form II males
show shrinkage of the mesial process (figs. 8-9).
Relationships: Faxonella creasert is in many respects intermediate
between F. beyeri and F. clypeata. Like F. clypeata it possesses a long
central projection, but it also resembles F. beyeri in the long mesial
process which extends parallel to the central projection. F. creaseri
is distinct from both F. beyeri and F. clypeata in that the mesial
processes overlap not only each other, but also the central projections.
It is interesting to note that F. creaseri is found at the same latitude as
F. beyeri, and both species are found near the edges of the trans-
Mississippian range of F. clypeata (Fitzpatrick, 1963), F. beyeri to the
west and F. creaseri to the east. Based on collections by Dr. Joe B.
Black and me, both F. beyeri and F. creaseri are believed to be the
only Faxonella found within their ranges. Neither species shows signs
of intergradation with F. clypeata where their ranges are in close
proximity, although the possibility of such intergradation does exist.
Both F. beyeri and F. creaseri are common within their ranges and no
obvious reasons exist for their seemingly isolated distributions.
Etymology: I take pleasure in naming this new species in honor of
Prof. Edwin P. Creaser, who rediscovered Faxonella clypeata and first
recognized Faxonella as distinct from his Faxonius (= Orconectes Cope,
1872).
LITERATURE CITED
Corr, E. D. 1872. On the Wyandotte Cave and its fauna. Amer.
Nat., 6: 406—422.
Creaser, E. P. 1933. Descriptions of some new and poorly known
species of North American crayfishes. Occ. Pap. Mus. Zool.,
Univ. Michigan, No. 275: 1-21.
Fitzpatrick, J. F., Jr. 1963. Geographic variation in the crawfish
Faxonella clypeata (Hay) with the definition and defense of
418 Proceedings of the Biological Society of Washington
the genus Faxonella Creaser (Decapoda, Astacidae). Tulane
Stud. Zool., 10(1): 57-79.
Gmarp, CHARLES. 1852. A revision of the North American Astaci,
with observations on their habits and geographical distribu-
tion. Proc. Acad. Nat. Sci. Philadelphia, 6: 87-91.
Hay, W. P. 1899. Description of two new species of crayfish. Proc.
U.S. Nat. Mus., 22: 121-123.
Hosss, H. H., Jr. 1948. A new crayfish of the genus Cambarus from
Texas with notes on the distribution of Cambarus fodiens
(Cottle). Proc. U. S. Nat. Mus., 98: 223-231.
Penn, G. H. 1950. A new crawfish of the genus Orconectes from
Louisiana. Jour. Washington Acad. Sci., 40(5): 166-169.
. 1953. A new crawfish of the genus Procambarus from
Louisiana and Arkansas. Jour. Washington Acad. Sci., 43(5):
163-166.
Vol. 81, pp. 419-426 30 December 1968
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
A NEW SPECIES OF GECKO (SAURIA: GEKKONIDAE)
FROM THE BAY ISLANDS, HONDURAS
By JAmes R. Dixon
Department of Wildlife Science, Texas AUM University,
College Station, Texas
I began a revisionary study of the genus Phyllodactylus in
1956 and subsequent to this revision a new species of gecko,
P. insularis, was described from Half Moon Cay, British Hon-
duras (Dixon, 1960). Later Dixon (1964) compared P. insularis
with three specimens of Phyllodactylus from Roatan Island,
the largest of the four islands that comprise the Bay Islands
chain. At that time differences were noted in color, color
pattern, body proportions, lamellae of the fourth toe and snout-
vent length.
An analysis of 19 individuals (including two hatchlings )
from Islas Roatan and Guanaja collected by Meyer and
Wilson in July, 1967, reveals several additional differences
between insularis and the Bay Islands population. The popula-
tion from the Bay Islands appears to represent an undescribed
species and since it seems to prefer palm trees to other kinds
of habitat, I propose that it be known as
Phyllodactylus palmeus new species
Holotype: Louisiana State University 16986, adult ¢ from 0.5 km.
N. Roatan, Isla de Roatan, ca. 25 mts., collected by John R. Meyer
and Larry D. Wilson on July 14, 1967, original number JRM 1807
(Figure 1).
Paratypes: LSU 16987-92, all topotypes; TCWC 24016, 0.5 km. N.
Roatan, Isla de Roatan, ca. 10 mts.; LSU 16993-94, 3 km. N. Roatan,
Isla de Roatan, ca. 25 mts.; LACM 38512-13, 1 km. N. Roatan, Isla de
Roatan, ca. 10 mts.; LACM 38514—15, southeast shore, opposite Guanaja;
Isla de Guanaja; LACM 38516-20, La Playa Hotel, Isla de Guanaja;
MVZ 52402, Bay Islands; BM(NH) 1889.11.13.42-43, Roatan Islands;
all from Honduras.
42-——Proc. Brow. Soc. WaAsH., Vou. 81, 1968 (419)
420 Proceedings of the Biological Society of Washington
FicureE 1. Dorsal view of the holotype of Phyllodactylus palmeus
(LSU 16986).
New Honduras Gecko 421
Diagnosis: A rather large Phyllodactylus, maximum snout-vent length,
76 mm., that differs from its closest relative, P. insularis, by having
48-60 (M = 52.9) transverse ventral scales from throat to anus rather
than 60-66 (M = 63.9); tubercles in a paravertebral row from rear of
head to base of tail number 39-47 (M = 44.1) rather than 46-52 (M =
49.5); light ash-gray ground color rather than dark chocolate brown;
white spot below eye absent; enlarged tubercles present on thigh; head
width almost twice as wide as deep rather than head width and depth
about equal.
Description of holotype: Rostral two-thirds as high as wide, its dorsal
edge with a slight posterior curve and with a vertical groove about
half the depth of rostral; internasals somewhat rounded, their median
edges in broad contact, bordered posteriorly by five granules and _ post-
nasal of each side; nostril surrounded by rostral, labial, internasal and
two postnasals; first labial in broad contact with lower edge of nostril;
shallow depression between internasals and moderate depression in
frontal region; 14 scales between nostril and eye; scales in posterior
loreal region about three to four times larger than interorbital scales;
29 scales across snout between third labials; 19 scales between anterior
edge of orbits and 24 midorbital scales; eye large, its diameter con-
tained in snout length slightly less than two times; eyelid with two
rows of granules and one larger outer row of scales, last four or five
of which are pointed; diameter of ear contained in eye diameter two
and one-half times; ear opening not denticulate, scales on the anterior
and posterior borders rounded, subequal in size; rear of head with
many larger tubercles intermixed with granular scales; six supralabials
and 4+ 5 infralabials to a point below center of eye; mental bell-
shaped, slightly longer than wide, bordered posteriorly by two _post-
mentals; postmentals much wider than long, their median edges in
narrow contact with one another; postmentals followed by tranverse
row of seven smaller scales, followed by second row of 11 smaller
scales; postmentals contact only the first labial on each side.
Dorsum with 16 longitudinal rows of enlarged, keeled tubercles
that are somewhat flat, paravertebral row with 45 tubercles from rear
of head to base of tail, 28 between axilla and groin; paravertebral rows
separated from each other by three to four rows of granules; five rows
of tubercles reach to rear of head on right side, three on left, eight
rows to base of tail; each tubercle of enlarged dorsal series separated
from preceding tubercle by one to two granules; postanal enlarged scales
number three on each side; venter with 32 longitudinal and 56 trans-
verse rows of scales.
Dorsal surface of upper arm with flattened scales, forearm with larger
tubercles dispersed among smaller flattened scales; dorsal surface of
thigh with seven to nine tubercles among the granular scales; lower
leg with 10 to 12 tubercles dispersed among granular scales; lamellae
formula for hand 7-9-11-11-9, foot 7-10-12-13-10; claw hidden when
422 Proceedings of the Biological Society of Washington
—, SSS =: == 7 oes oes Coal TooeesT
48 49 50 51 52 53 5455 56 57 58 59 60 6I 6263 64 65 66 G7 C8 Sr 89
<< = Se
SCALES BORDERING
B a B = = POSTMENTALS
TRANSVERSE VENTRAL.
SCALES — a c (/--5
26 27 28 29 30 3] 32 33 3435
A =
SCALES BORDERING
5678S
fe
SS
ik
A
B = = LONGITUDINAL B INTERNASALS
VENTRAL SCALES
—— c
19 20 21 22 23 24 2526 27 28 293031 32 12 13 14 15 16
A a ao os
SCALES ACROSS LAMELLAE
B
wo 6 Cl c ATED
THIRD LABIALS
31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 II 12 13 14 15 16
le ee CSE
= SCALES BETWEEN
PARAVERTEBRAL B — | = = B = NOSTRIL and EYE
TUBERCLES from
Eo: c Am | co (ler
15 16 17 18 19 20 2l 22 23 24 25 26 27 28 2930 16 17 18 19 20 2! 22 23 24 25 26 27 28
= SCALES of MID-ORBITAL
(SS SS] —<— = S ot mi
earaverteora, = 8 ESR 5 +
TUBERCLES between
AXILLA ond GROIN c Cc
FicureE 2. A statistical analyses of 10 characters of three populations
of Phyllodactylus: A = tuberculosus (33 specimens), B = palmeus (22
specimens ), C = insularis (7 specimens). The horizontal line represents
the known range of variation, vertical line the mean; hollow rectangle
represents two standard deviations on either side of the mean, solid
rectangle two standard errors.
viewed from below; terminal pad large, slightly longer than wide,
truncate.
Measurements in mm.; snout-vent length 72, axilla-groin length 33.5;
leg length 26.5, arm length 22, tail length 83, head length 20.5, head
depth 9, head width 16, snout length 8, eye diameter 4.6, ear diameter
1.8.
Color in alcohol: Ground color ash-gray; dorsum with 10 narrow
grayish brown crossbands, interrupted mid-dorsally, with two paraverte-
bral and two lateral broken lines of similar color that tend to form a
loose rectangular pattern of lines from rear of head to hind limbs; upper
New Honduras Gecko 423
arm with one to two obscure grayish brown bands; lower arm with two
distinct grayish brown bands; thigh with obscure spotting of ground
color and grayish brown, head light ash, with inverted V-shaped, grayish
black mark from rostral to posterior edge of eye, two arms of “V”
passing along inner edge of eye; frontal and parietal regions with fine
grayish black reticulations intermixed with ground color; grayish black
line from lateral edge of rostral through middle of eye to behind ear;
labials cream with minute black dots concentrated near the center of
each labial; ventral surfaces cream white with ventral surfaces of
digits grayish; tail with six narrow black crossbands separated by
whitish interspaces that are two times width of dark bands; one-third of
tail regenerated; lower surfaces of tail grayish white, regenerated portion
mottled cream and black.
Variation: The snout-vent length of adult @ 2 range from 55 to 69
mm., adult ¢ ¢ from 60 to 76 mm.; postmentals number two in all
specimens except one, which has four; postmentals contact only the
first labial on each side in all specimens except one, which has the
postmental contacting two labials on one side; midorbital scales vary
from 21 to 28 (M = 23.7); scales across snout between third labials
vary from 25 to 31 (M = 28.5); longitudinal and transverse scales of
the venter vary from 28 to 33 (M = 30.4) and 48 to 60 (M =52.9)
respectively; transverse row of scales following postmentals vary from
6 to 9 (M=7.3); scales bordering posterior edge of internasals vary
from 6 to 9 (M =7.1); scales between nostril and eye vary from 12 to
15 (M = 13.5); lamellae of fourth toe vary from 13 to 15 (M = 13.7);
number of tubercles in paravertebral row from rear of head to base of
tail and between axilla and groin vary from 39 to 50 (M = 44.1) and
22-9299 (M = 26.4), respectively; number of tuberculate rows across
base of tail vary from 6 to 8, with 8 rows present in about 60 per cent
of the sample; number of supralabials to a point below center of eye
varies from 5 to 7, with 6 occurring in 90 per cent of sample; rows of
tubercles on the tail are usually 6 for the first four to six whorls, 4 for
about five whorls, 2 for about four whorls, and tubercles generally absent
for the last four whorls.
The dorsal color pattern is variable, ranging from broken crossbands
to a tessellated pattern. The dorsal head pattern is constant, with the
inverted V-shaped mark of the snout present in all specimens. One
specimen has an almost uniform dorsal coloration, with four thin
paravertebral dark lines from nape to level of groin. The color of the
belly is cream white in all specimens.
Hatchlings of P. palmeus are generally darker in dorsal color than
the adults. The transverse bands are more distinct and contrasting
strongly with the ground color. The distal half of the tail is almost
completely black for one-half its length; the proximal half cream white
with two to four pale gray crossbands.
Comparison: Phyllodactylus palmeus is more closely related to P.
424 Proceedings of the Biological Society of Washington
insularis of Half Moon Cay, British Honduras, than to mainland Hon-
durian P. tuberculosus (Figure 2). Most characters of squamation of
P. palmeus and P. insularis show some degree of overlap. The major
scale differences are found in the number of transverse ventral scales
and tubercles in a paravertebral row from rear of head to base of tail
(see diagnosis). In addition, the ground coloration of the two species
is different, P. insularis is chocolate brown and P. palmeus, ash gray.
Phyllodactylus insularis has a small white spot~below the eye, con-
trasting with brownish labials, while P. palmeus has whitish labials and
no spot below the eye. The head proportions of P. insularis and P.
palmeus are different, insularis having a head almost as deep as wide
while that of palmeus is almost twice as wide as deep.
The new species is significantly distinct from mainland Hondurian
P. tuberculosus in the following characters of squamation: Lower num-
ber of transverse ventral scales; higher number of midorbital scales;
scales across snout between third labials, tubercles in paravertebral row
from rear of head to base of tail and from axiJla to groin. However,
P. palmeus resembles P. tuberculosus in coloration, head proportions,
and body size.
Habitat and Natural History: On the morning of July 14, 1967, Meyer
and Wilson found seven specimens of P. palmeus in a grove of standing, —
dead palm trees. The palms were hollow, approximately 10 to 15 feet
in height, and the lizards were taken from the hollow trunks. Two
eggs were found in the same area in a rotting palm stump. Two addi-
tional specimens were seen on the side of a living tree on the evening
of the 14th, but only one was captured. Several additional specimens
were taken from rotting palm trees on successive days in the same
general area where the first specimens were found.
P. palmeus was also found during the day under the bark of trees,
under a piece of tin, and beneath piles of palm fronds. One was found
at night on a log overlying a stream and one on the stem of a banana
plant.
On July 18, Meyer found six eggs in a rotten palm stump. Four
eggs had hatched and two appeared to be in the process of hatching.
The latter two eggs hatched shortly after they were taken. It has been
my experience that Phyllodactylus deposit only two eggs at a time. The
oviducts of the female usually contain several developing eggs at the
time the two fully developed eggs are laid. Meyer apparently found
three clutches of eggs laid by three different females at about the same
time. Three clutches of eggs in the same palm stump may represent
a communal nest, and optimum conditions (in this case, the rotting
palm stump) are sought out by several females for egg deposition.
The eggs taken on July 14th measured 9.5 x 12.1 mm. The eggs
contained well developed embryos, without pigment, and half the size
of the hatchlings.
New Honduras Gecko 495
LITERATURE CITED
Dixon, J. R. 1960. The discovery of Phyllodactylus tuberculosus
(Reptilia: Sauria) in Central America, the resurrection of
P. xanti, and description of a new gecko from British Hon-
duras. Herpetologica, 16(1): 1-11.
1964. The systematics and distribution of lizards of the
genus Phyllodactylus in North and Central America. New
Mexico State University Sci. Bull., 64(1): 1-139.
426 Proceedings of the Biological Society of Washington
G73
Vol. 81, pp. 427-430 30 December 1968
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
A NEW SPECIES OF ANOURA (MAMMALIA:
CHIROPTERA: PHYLLOSTOMIDAE) FROM
SOUTH AMERICA!
DitForp C. CARTER
Department of Wildlife Science, Texas Atv-M University,
College Station, Texas
Three specimens of Anoura taken in Peri in 1964 by a
collecting party from Texas A&M University are not referable
to any of the recognized species of Anoura. Subsequently,
two additional specimens were found, one in the Field Museum
of Natural History; the other in the Museum of Zoology,
Louisiana State University. For this undescribed species, I
propose the name
Anoura brevirostrum new species
Holotype: adult female, skin and skull, no. 11882, Texas Cooperative
Wildlife Collection, from 31 km S Tingo Maria, 850 m, Huanuco, Peri;
collected 25 August 1964 and prepared by Dilford C. Carter, original
number 5470. Selected measurements in millimeters: head and body,
64; foot, 9; ear from notch 14 (preceding measurements were taken in
the field); forearm, 40.3; third metacarpal, 41.5; first phalanx of third
digit, 13.9; second phalanx of third digit, 22.6; fourth metacarpal, 40.3;
first phalanx of fourth digit, 10.1; fifth metacarpal, 35.8; first phalanx
of fifth digit, 8.2; tibia, 14.2; calcar, 3.6; greatest length of skull
including incisors, 23.1; zygomatic breadth, 9.5; braincase breadth, 9.3;
postorbital constriction, 4.6; length of maxillary toothrow, 8.2; greatest
width across upper molars (M*-M*), 5.6; greatest width across canines,
4.7. Weight, 12.5 grams.
Description: A medium-sized Anoura with a relatively short rostrum.
Dorsal coloration gray, individual hairs with basal two thirds white,
distal third gray with slightly paler tips. Ventral coloration paler than
1 This contribution results from an investigation supported in whole by Public
Health Service Research Grant No. AI-03743, from the National Institute of
Allergy and Infectious Diseases. Contribution No. 7227 of the Texas Agricultural
Experiment Station.
43—Proc. Biot. Soc. Wasu., Vou. 81, 1968 (427)
428 Proceedings of the Biological Society of Washington
dorsal; individual hairs on thoracic region with basal half very pale
gray, distal half pale gray with very pale gray tips; on abdominal region
with basal two thirds dark gray and distal third very pale gray, almost
white. Membranes blackish, ears and noseleaf gray. Interfemoral
membrane narrow, with dense fringe of hairs extending two to three
millimeters beyond edge. Tail absent. Skull with zygomatic arch
complete and moderately well developed; basisphenoid pits rather well
developed and separated by relatively narrow ridge; first upper premolar
greatly reduced in size; upper molars with well developed protocone;
hypocone weak on M7’, all but absent on M7?, absent on M®°; first lower
premolar largest of the lower premolars, with main cusp posterior to
middle of tooth.
Comparisons: Anoura brevirostrum is readily separated from A.
geoffroyi and A. caudifer in which the first lower premolar is no larger
than the other two lower premolars and the main cusp of the first
lower premolar is anterior to the middle of the tooth. A. brevirostrum
is most similar to A. cultrata, but differs from it in the following ways:
tail absent; rostrum relatively shorter (shortest of the four species of
Anoura) with a more crowded toothrow; first upper premolar very
small and its dorso-ventral dimension obviously greater than its antero-
posterior dimension (the opposite is the case in A. cultrata); hypocone
more weakly developed? on M’; and hypocone all but absent on M?
whereas in A. cultrata the hypocone is rather well developed for species
of Anoura and essentially no less developed on M? than on Mt’.
Measurements: See Table 1.
Distribution: Known from the type-locality; Cordillera Azul, Hudnuco,
Peri; and Gramales (SW of Bucaramanga), Santander, Colombia. The
type-locality is situated in the foot hills of the eastern slope of the
Andean highlands and the three specimens from there were taken
on a rather steep hillside where second growth brush was two to three
meters high. The vegetation in the general area was second growth
and cut-over rain forest. Logging operations were in progress but
diminishing. The other Peruvian specimen came from the pass in the
Cordillera Azul (about 45 km NE of Tingo Maria) through which the
road from Tingo Maria to Pucallpa passes. The elevation there is about
600 m and the vegetation in 1964 was similar to that at the type-locality
but less disturbed. The Colombian specimen is from the west slope
of the Cordillera Oriental which is east of the Magdalena River. The
elevation is not recorded on the tag and I have been unable to find the
exact locality on maps of that area.
Etymology: Latin, brevis, short + rostrum, snout.
Specimens examined: A. brevirostrum: PERU: 19 mi. S Tingo Maria,
850 m, Huanuco, 16, 2° 2, TCWC; Divisoria en Cordillera Azul, about
600 m, Huanuco, 19, LSU. COLOMBIA: Gramales, Santander, 19,
FMNH. A. cultrata: PANAMA: Cerro Mali, 4100 and 4700 ft, Darién,
1g, 3292, USNM; Cerro Tacarcuna, 4100 ft, Darién, 16, USNM;
New South American Bat 499
TABLE 1. Means and extremes in millimeters for selected variates of
the holotype and four paratypes of A. brevirostrum compared with the
means and extremes of the same variates of A. cultrata.
A. brevirostrum A. cultrata
Variate No. Min. Mean Max. Min. Mean Max. No.
Weight Oy TLAS) TIS TDS
Head & body 4 6l 62.5 64 76 84.6 94 3
Forearm 5 38.7 39.9 40.9 ANG Wer AS y(n Ae2 alley
Metacarpal III 5 392 414 42.8 43.0 45.0 46.9 15
Metacarpal IV Bie yO Zale
Metacarpal V CR CEE | BIO) BRO. BHO ZB Ibs
Tibia Silo ome So Ome el Ane, SIO Sea Go lip:
Calcar BS Be 3.6 3.8 4.0 1
Greatest skull
length 5 23.1 23.4 24.0 25.8 263 269 14
Zygomatic breadth 5 9.5 9.7 10.0 NOG: Oss UL =
Cranial breadth 5 9.2 9.3 9.6 98 10.0 10:2 £14
Postorbital
constriction 5 46 4.7 4.8 4.9 5.2 buf) 14
Width of rostrum
above orbits 5 416 4.8 aD) 5.0 Dal ba hs
Maxillary toothrow 5 7.8 8.0 8.2 8.8 9.1 94 14
Greatest width
across molars ys} 5.5 5.7 Bott 5.9 G3 14
Greatest width
across canines 5 4.7 47 4.8 4.7 ED) fey 18)
Length of PM: BH ey LAY Ta 1.75 185 1.90 13
Tacarcuna Village camp, 3200 ft, Darién, 26 6, 4292 (including
holotype), USNM; Cerro Punta, 5300 ft, Chiriqui, 12, USNM; Cuesta
de Piedra, Chiriqui, 1 6, USNM; about 20 mi. SSW Changuinola, Bocas
del Toro, 12, USNM; Rio Changena camp, Bocas del Toro, 1¢ (juv),
USNM. COSTA RICA: 18 mi. NE Naranjo, 3100 ft, Alajuela, 14,
TCWC.
Remarks: In addition to the above specimens, I have compared the
two paratypes from the type locality with the following type specimens:
Anoura geoffroyi Gray: sex unknown, skin and skull, BMNH no. lla,
Brazil. Glossonycteris lasiopyga Peters (= Anoura geoffroyi lasiopyga):
syntypes: one ¢, mounted skin with skull, ZMB no. 3565 and one sex
unknown, skin and skull (skull lostP), ZMB no 3564, both from
Cuernavaca, 12 meilen von Mexico, Mexico, August 1866. Glossophaga
caudifer Geoffroy (= Anoura caudifer): holotype: 9@skin only, MNHN
no. 937, Brésil. Lonchoglossa wiedi aequatoris Lonnberg (= Anoura
430 Proceedings of the Biological Society of Washington
caudifer): holotype: ¢, skin and skull, NR no. 6, Tlambo, 5000 ft,
Gualea, Ecuador, 20 April 1913.
Rode (1941) did not include the specimen MNHN no. 937 as one
of the types in the Paris Museum (Muséum National d’Histoire Naturelle)
and there may be some question as to whether this specimen is the
holotype of Glossophaga caudifer since the Paris Museum’s A-series
catalog of about 1868 lists the specimen as having come to the Museum
from A. Saint-Hilaire, 1822, and the date of publication for the name
is 1818. However, the date 1822 is probably the date of registration
in a catalog no longer extant and probably has nothing to do with the
date of capture or receipt of the specimen at the Museum. The holotype
of G. caudifer was obtained by Auguste de Saint-Hilaire and the
Delalande brothers while they were in Rio de Janeiro, Brazil and I think
it reasonable to assume that the specimen in question is indeed the
holotype of this species.
The Colombian specimen of Anoura brevirostrum, taken in December
1945, is a female, now a skin and skull, but was preserved originally in
fluid and the pelage is pale reddish-brown in color. The tag is without
notation concerning its reproductive status. The following reproductive
data were recorded for the other specimens of A. brevirostrum. The ~
two females from the type-locality, taken 25 and 26 August 1964, were
neither pregnant nor lactating. The Peruvian female taken from the
Cordillera Azul on 21 August 1967 was lactating. The one male caught
at the type-locality on 26 August 1964 had testes 4 mm in length.
I wish to express my thanks to Drs. Joseph Curtis Moore, Field Mu-
seum of Natural History (FMNH) and George H. Lowery, Museum of
Zoology, Louisiana State University (LSU) for the loan of specimens
in their charge. I am especially grateful to Dr. Charles O. Handley Jr.,
United States National Museum (USNM); Mr. John E. Hill, British
Museum (Natural History) (BMNH); Dr. Francois de Beaufort, Mu-
séum National d’Histoire Naturelle (MNHN); Dr. Georg Stein, Institut
fiir Spezielle Zoologie und Zoologisches Museum der Humboldt-Univer-
sitét zu Berlin (ZMB); and Dr. Alf G. Johnels, Naturhistoriska Riksmu-
seum (NR) for the opportunity to examine specimens in their charge
and. the hospitality extended to me. Also, I wish to thank Dr. Handley
for kindly loaning me a copy of his manuscript description of Anoura
brevirostrum.
LITERATURE CITED
Rope, P. 1941. Catalogue des types de mammiféres du Muséum Na-
tional d’Histoire Naturelle, II Ordre des Chiroptéres. Mu-
séum National d’Histoire Naturelle, pp. 72-105.
Vol. 81, pp. 431-438 30 December 1968
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
A SYNOPSIS OF THE AMERICAN SPECIES OF THE
GENUS ORYTTUS (HYMENOPTERA, SPHECIDAE)
By R. M. Bowarr
Department of Entomology, University of California, Davis
The American species of Oryttus were assigned by Pate
(1938) to Harpactostigma Ashmead of which the type species,
velutinus Spinola, is Chilean. However, the close relationship
to Oryttus from Europe is obvious, and Harpactostigma seems
hardly worth distinguishing, even as a Chilean subgenus.
Oryttus is composed of about a dozen species of slender,
medium-sized wasps, with long antennae and mostly with
some rough thoracic sculpture. Seven species occur in the New
World and five of these have the forefemur of the male greatly
expanded. Females of Oryttus have the forepulvillus much
larger than the pulvilli of the other legs. Contrastingly, the
males have the hindpulvilli the largest. Another morphological
curiosity is the presence in males of at least 5 of the known
species of one midtibial spur instead of two spurs as usual in
Gorytini. Other features are: inner eye margins essentially
parallel or converging a little below; forewing spotted or
clouded; hindwing cu-a strongly curved near cubitus; tergite
I at least a little constricted toward base of II, curved gently
toward base in profile; male sternite VIII bispinose apically
and usually protruding.
Nests are provisioned with cicadellids or fulgorids.
Holotypes of the new species are deposited in the California
Academy of Sciences, San Francisco (O. lapazae) or the En-
tomology Museum of the University of California at Davis
(O. yumae).
Kry to NEw WoRrRLD sPEcIES OF Oryttus
1. Hindfemur with non-black area more than one-half yellow, or ab-
44_Proc. Biot. Soc. WaAsH., Vou. 81, 1968 (431)
432 Proceedings of the Biological Society of Washington
dominal markings whitish; male with two midtibial spurs; male
forefemur with ventral depression, if any, occupying no more
lovin lokell Grvsdomlbe (Gates, &) io W)) 2
Hindfemur with non-black area more than one-half red, abdomen
without white markings; male with only one mid-tibial spur; male
forefemur with ventral depression occupying basal two-thirds or
BOTS | (CHI SS y NeoA|) Ate ea AN I OR Ae 4
2. Tergite I in dorsal view about as broad basally as apically; abdomi-
nal bands whitish; femora red; scutum all black and coarsely stri-
atopunctated (Chile: jee a eeesaene een eee velutinus (Spinola)
Tergite I in dorsal view much broader toward apex; scutum yellow
laterally and finely to moderately punctate (North America) __ 3
3. Propodeum with “cheeks” only slightly roughened; male forefemur
not depressed beneath basally (fig. 6) (s. California )
I SAB A RRL NCO RU 2 STs Oa Re umbonatus (Baker )
Propodeum with “cheeks” strongly and closely roughened; male
forefemur with basal one-half beneath hollowed (fig. 5) and
conspicuously hairy (Shasta Co. to Nevada Co., California )
eR NL WN A Ae IE mirandus (W. Fox)
4. Propodeal “cheeks” polished; male forefemur nearly one-half as
road sas Tom (igs cB) ee Daa INN 5
Propodeal ss cheeksjigaug nemie clyyeeee eee eae 6
5. Abdomen predominantly black, marked with yellow (e. of 100th
Tug (cig ColPs 0) nea casera ae cP SNUG ACen | Aa See gracilis Patton s. s.
Abdomen predominantly yellow, marked with red and black (w.
Oe UOCdn sneChem)) gracilis arapaho Pate
6. Scutum coarsely punctate, punctures considerably larger than those
of interocellar area; male forefemur more than one-half as broad
as long, greatest breadth near middle of femur (fig. 1) (Baja
(OPM Rorgautals- hy (Gpatolo))) Lan ee lapazae R. Bohart
Scutum with fine to moderate punctation, punctures not larger
than those of interocellar area; male forefemur with greatest
lovqserchdny, Joveywornel oonrakelley (Gate, Bho) 0
7. Propodeum red except for enclosure; wings strongly bicolored;
male forefemur about three times as long as broad (fig. 4);
flagellar article VIII shorter along midventral line than preceding
two articles together (fig. 9) (sw. U.S.) yumae R. Bohart
Propodeum black with two large yellow spots; male forefemur
about twice as long as broad (fig. 2); flagellar article VIII along
midventral line about as long as preceding two articles together
(Eig OD KGwe MUSA)? £ se Rees Corer see laminiferus (W. Fox)
Oryttus gracilis gracilis (Patton)
Hoplisus gracilis Patton, 1879. Canad. Ent. 11:210.
Female holotype, Southington, Connecticut (type lost).
The mostly red legs, polished propodeal “cheeks”, small subantennal
triangle (about one-fourth as broad as frons at that point (fig. 12), and
American Species of Oryttus 433
WS xn [ TAG (i, fer
FicurEs. 1-7, male forefemur from beneath. Fic. 1, O. lapazae.
Fic. 2, O. laminiferus. Fic. 3, O. gracilis. Fic. 4, O. yumae. Fic. 5, O.
mirandus. Fic. 6, O. umbonatus. Fic. 7, O. velutinus. Fics. 8 to 10,
last five articles of male flagellum from beneath. Fic. 8, O. velutinus.
Fic. 9, O. yumae. Fic. 10, O. laminiferus. Fics. 11 and 12, center of
male face. Fic. 11, O. lapazae. Fic. 12, O. gracilis. Fics. 13 and 14,
female pygidium. Fic. 13, O. yumae. Fic. 14, O. laminiferus. Fics. 15
—17, male metapleuron. Fic. 15, O. gracilis. Fic. 16, O. lapazae. Fic.
17, O. yumae.
434 Proceedings of the Biological Society of Washington
coarsely reticulate upper metapleural suture (fig. 15) characterize the
species. The typical form has very dark wings and tergites II and fol-
lowing black with narrow yellow bands. This species, as a whole, shows
much the same color range as its largely sympatric, but more common
relative, Pseudoplisus phaleratus (Say). The distribution of the typical
subspecies covers much of eastern United States to the 100th meridian.
Oryttus gracilis arapaho (Pate)
Harpactostigma arapaho Pate, 1938. Trans. Amer. Ent. Soc. 64:
67. Female holotype, Roggen, Colorado (Mus. Comp. Zool.
Harvard). New status.
Harpactostigma rutilum Pate, 1938. Trans. Amer. Ent. Soc. 64:69.
Male holotype, El Paso, Texas (Acad. Nat. Sci. Phil.). New
synonymy.
An examination of Pate’s types as well as new material have shown
the relationship to O. gracilis. The extensive red markings and mostly
yellow abdomen characterize the subspecies which occurs from western
Texas and Kansas to Colorado. New records are: one pair, Roggen,
Colorado, 8 July 1933 (M. James, L. Ireland); one male, Brookville, ©
Kansas, 11 June 1950 (C. D. Michener).
Oryttus laminiferus (W. Fox)
Gorytes ruficornis Provancher, 1888 (nec G. ruficornis Latreille,
1805). Addit. Corr. Faune Ent. Canada Hym., p. 273. Holo-
type female, “Californie” (Laval Univ., Quebec).
Gorytes laminiferus W. Fox, 1895. Proc. Acad. Nat. Sci. Phil. 47:
532. Holotype male, Washington Territory (Acad. Nat. Sci.
Phil. ).
Gorytes flavicornis Baker, 1907. Invertebrata Pacifica 1:162.
Holotype male, Claremont, California (Pomona College).
Gorytes rufulicornis Maid] and Klima, 1939. Hym. Cat., Pars 8,
Sphecidae 1:102. New name for ruficornis.
This species, while not at all common, is the most widespread of the
western forms. I have seen specimens from California, Oregon, Nevada,
Idaho, and Utah.
The mainly black and yellow body, reddish-brown wings, sculptured
propodeal “cheeks”, broadly expanded male forefemur (fig. 2), relatively
long male flagellar article VIII (fig. 10), and striatopunctate female
pygidium (fig. 14) distinguish the species.
Oryttus lapazae R. Bohart new species
Male holotype: Length 11 mm. Mostly orange-red and yellow, a
little black. Yellow are: scape in front, face nearly to midocellus, pro-
notum all across, scutellum and metanotum mostly, forelegs and midlegs
American Species of Oryttus 435
extensively, hindfemora distally, hindtibia outwardly, hindtarsus mostly,
tergites I to III broadly at apex, IV to V mostly; black are: broad area
around ocelli, scutellum in front, stain along metapleural suture, pro-
podeal enclosure, coxae partly; wings mostly brown, stigma and costal
cell more reddish. Pubescence fine, silvery to fulvous, inconspicuous.
Punctation fine to moderate, coarse on scutum where punctures are 0.5
to 2.0 diameters apart, close and smaller between ocelli, smaller and
scattered on scutellum, mesopleuron and toward apices of tergites. Fla-
gellar article VIII modified as usual but only about 1.5 times as long as
either VII or IV in midline; frons narrowed slightly and gradually below,
least interocular distance a little more than eye breadth in front view,
subantennal triangle one third of frons breadth (fig. 11); propodeum
closely and coarsely sculptured, enclosure obliquely striate; metapleural
suture finely punctate throughout (fig. 17); forefemur expanded, flat-
tened, concave beneath, free edge thin and curving forward, greatest
breadth near middle and about three-fifths length of femur (fig. 1); mid-
tibia with only one apical spur; tergite I in dorsal view nearly one-half
as broad as long and about one-half as broad as tergite II; points of
sternite VIII separated by about a midocellus diameter.
Holotype male (CAS) La Paz, Baja California, Mexico, 7 October
1955 (F. X. Williams).
Known only from the type, O. lapazae is distinguished by the unusual
forefemoral breadth (fig. 1) which is greatest near the middle. Also
characteristic is the coarsely punctate scutum. The dark and rather
evenly colored wings are found also in gracilis. The almost wholly red
and yellow body pattern is shared with yumae.
Oryttus mirandus (W. Fox)
Gorytes mirandus W. Fox, 1892. Canad. Ent. 24:152. Holotype
male, Nevada (Acad. Nat. Sci. Phil.).
From 1892 until recently this species has been known only from the
type. New records are from California: male, Cayton, Shasta Co., 21
July 1918 (E. P. Van Duzee); female, Berkeley, 4 August 1938 (N.
Hardman); female, near Quincy, 16 July 1949 (P. D. Hurd); male,
Sierraville, 26 July 1956 (R. M. Bohart); male, Sattley, 22 July 1949
(E. I. Schlinger); female, Carnelian Bay, Placer Co., 22 July 1959 (R.
M. Bohart).
The black and yellow aspect, restricted, but unusually hairy depression
beneath the male forefemur, two midtibial spurs in the male, roughened
propodeal “cheeks”, and small subantennal triangle are distinguishing.
Superficially, there is a close resemblance to O. umbonatus which has
similar markings and mostly clear wings. Differences in male forefemora
(figs. 5, 6) and propodeal roughness are obvious, however, as indicated
in the key.
436 Proceedings of the Biological Society of Washington
Oryttus umbonatus (Baker )
Hoplisoides umbonatus Baker, 1907. Invertebrata Pacifica 1:163.
Holotype male, Claremont, California (Pomona College).
Gorytes femoratus Bradley, 1920. Trans. Amer. Ent. Soc. 46:119.
Holotype male, Claremont, California (Cornell University).
New synonymy.
The species is known from perhaps a dozen specimens from drier areas
of San Bernardino, Los Angeles, and Riverside counties of southern Cali-
fornia. New localities with dates of May and June are: Lebec (E. I.
Schlinger); Claremont; The Gavilan, Riverside Co. (P. H. Timberlake,
R. C. Bechtel); San Jacinto Mts. (E. V. Stahl, C. D. Michener); Victor-
ville (P. H. Timberlake).
The male has no depression beneath the slightly broadened forefemur
(fig. 6) and the propodeal “cheeks” are smooth with a few faint wrinkles.
As in mirandus, the male has two midtibial spurs, and the extensively
clear wings are similar. However, the propodeal sculpture is differenti-
ating in both sexes as indicated in the key.
Oryttus velutinus (Spinola)
Hoplisus velutinus Spinola, 1851. In Gay, Hist. fis. Pol. Chile,
Zool. 6:337. Holotype male, Central Provinces, Chile (type
supposedly at University of Turin, Italy).
In this species the basal tergite is unusually constricted and there are
subbasal hair patches on sternites IV to VI of the male. Also, the male
forefemur is simple (fig. 7), the male flagellum is only a little modified
(fig. 8), the midtibiae are two-spurred in both sexes, and the pronotal
ridge is more rounded than in other species and less closely appressed
to the scutum. These features may warrant recognition of Harpacto-
stigma as a subgenus. Im sculpture O. velutinus is much like such Old
World species as O. infernalis (Handlirsch).
Authenticated records are all from Chile, especially from the provinces
of Santiago, Valparaiso, and Coquimbo.
Pate (1938) with some reservations, placed Gorytes unicinctus Bréthes
from Argentina in the genus Harpactostigma along with velutinus. Re-
cently (Bohart, 1967), I have placed wnicinctus in Liogorytes R. Bohart.
Oryttus yumae R. Bohart new species
Male holotype: Length 10 mm. Head and thorax mostly orange, ab-
domen mostly yellow. Clypeus and lower frons, tibiae in front, prothorax
mostly, scutellum and postnotum yellow; pattern on vertex, irregular
spots on scutum, metapleuron partly, propodeal enclosure, spot above
insertion of gaster, tergites I and II basally, black; wings partly clear,
stain darkest along veins, in marginal cell, second and third submarginal
cells, and out to wing tip; stigma and costal cell light reddish. Pubes-
American Species of Oryttus 437
cence fine, silvery, inconspicuous. Punctation mostly fine, punctures
much smaller and farther apart on scutum than between ocelli, scattered
and fine on scutellum, metanotum and tergites, moderate and sparse on
mesopleuron, coarsely irregular and close on propodeum posteriorly. Fla-
gellar article VIII modified as usual, but only about 1.5 times as long
as either VII or IX in midline; frons narrowed below, least interocular
distance more than eye breadth in front view, subantennal triangle about
as in figure 11; propodeal enclosure obliquely striate; metapleural suture
stitch-like (fig. 17); front femur expanded (fig. 4), somewhat flattened,
concave beneath and pilose on basal two-thirds, greatest breadth at distal
two-thirds where it is a little over one-third of femoral length; midtibia
with only one apical spur; tergite I in dorsal view about one-half as
broad as long and about one-half as broad as tergite II; points of sternite
VIII separated by about 1.5 midocellus diameters.
Female: About as in male. Length 11 mm. Clypeus, legs mostly and
tergite VI orange. Least interocular distance about 1.5 times eye breadth
in front view, frons narrowed only a little below. Pygidium broad, well
rounded apically, shiny with punctures well spaced (fig. 13).
Holotype male (UCD), 5 mi. se. of Bouse, Yuma Co., Arizona, on
Asclepias erosa, 14 May 1966 (M. A. Cazier, J. H. and J. M. Davidson).
Paratypes, one female (UCD), same data as type, one female (Arizona
State Univ., Tempe), 18 mi. s. Gila Bend, Arizona, on Viguiera deltoidea,
8 May 1965 (M. Cazier, M. A. Mortenson, J. M. Davidson).
The moderately expanded forefemur (fig. 4) is distinctive for the
male. In addition the roughened red propodeum with black enclosure is
shared only by lapazae which has a much more coarsely punctate scutum.
Reddish examples of arapaho approach yumae in appearance but the
former have a much narrower subantennal triangle and a smooth, yellow-
marked propodeum.
LITERATURE CITED
Bouart, R. M. 1967. New genera of Gorytini. Pan-Pacific Ent. 43:
155-161.
Pate, V. S. L. 1938. Studies in the Nyssonine wasps, III. A revision
of the genus Harpactostigma. Trans. Amer. Ent. Soc. 64:
57-77.
438 Proceedings of the Biological Society of Washington
4. G7 3
Vol. 81, pp. 439-472 30 December 1968
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
BATHYAL MYODOCOPID OSTRACODA FROM THE
NORTHEASTERN GULF OF MEXICO
By Louis S. KorRNICKER
Smithsonian Institution, Washington, D.C.
Myodocopid ostracods of the deeper waters of the Gulf of
Mexico are virtually unknown ... only | species, Cypridina fla-
tus Tressler, 1949, having been previously reported from 1200
meters near Tortugas (Tressler, 1949, p. 336, p. 431). There-
fore, I was quite pleased to receive from Dr. Willis E. Pequeg-
nat and Mr. Thomas J. Bright a small collection containing
myodocopid ostracods collected in a mid-water trawl that acci-
dentally dragged along the bottom at a depth of 1000-1200
meters for 1.5 hours during the Texas A&M University cruise
66-A-9 of the R/V Alaminos on July 11, 1966. The Myodo-
copida are described in the systematic part of this paper. Os-
tracods in the sample are listed below:
Order Myodocopida
Suborder Myodocopina
Superfamily Cypridinacea
Tetragonodon rhamphodes new
species 19
Paramekodon poulseni new
species 19
Bathyvargula optilus new
species YN MN uh
Suborder Halocypridina
Superfamily Halocypridacea
Conchoecia atlantica (Lubbock) 22 9
Conchoecia valdiviae Miller 22 9
Conchoecia macrocheira Miller 1 ¢
45—Proc. Bion. Soc. WaAsuH., Vout. 81, 1968 (439)
440 Proceedings of the Biological Society of Washington
Order Podocopida (ident. by Drs. R. H. Benson
and R. F. Maddocks )
Suborder Podocopina
Bairdoppilata ?hirsuta (Brady) 1°, 4 MT shells
Bairdia new species 222, 9 MT shells,
_ 2 single valves
Echinocythereis echinata (Sars) 1 single valve
Four specimens of bottom fish collected in the trawl con-
tained ostracods in their stomachs or intestines: Nezumia
hildebrandi (2 specimens ), Dicrolene intronigra (1 specimen),
and Dicromita agassizii (1 specimen). Nezumia hildebrandi
contained 4 specimens of the podocopid Krithe sp. (identified
by Dr. J. E. Hazel), 1 specimen of a halocyprid (new genus)?
and 3 specimens of the myodocopid Philomedes sp. in the
intestine, and 1 specimen of Bathyvargula optilus and 1
specimen of a halocyprid (new genus) in the stomach.
Dicrolene intronigra contained in its intestine 1 specimen of
the cladocopid ostracod Polycopsis sp. The intestine of Dicro-
mita agassizii contained 5 specimens of a halocyprid (new
genus) and 1 specimen belonging to the family Cypridinidae
(not Bathyvargula; furca missing from specimen).
Because the trawl was open in the water as well as on the
bottom, it is not possible to specify with certainty the depths
at which those ostracods not inside fish were collected. The
absence of Conchoecia in the stomachs and intestines of bottom
fish suggests that specimens of that genus collected free in
the trawl were living above the bottom. The presence of a
specimen of Bathyvargula optilus in the stomach of a
bottom fish indicates that members of the species collected in the
trawl were obtained at the bottom. The absence of natatory
setae on the exopodites of the 2nd antennae suggests that
Tetragonodon rhamphodes is a bottom dweller. It cannot
be determined with certainty at what depth specimens of
Paramekodon poulseni were collected, but it was probably on
the bottom.
I have taken the opportunity to redescribe herein the type-
species of Paramekodon Brady and Norman 1896, which is in
a This new genus of halocyprid with several new species is being described
separately by Dr. Georgiana Deevey.
Gulf of Mexico Ostracods 44]
the collection of the Hancock Museum, Newcastle-on-Tyne.
I wish to thank Dr. Georgiana Deevey for assistance in
identification of the halocyprids, Mr. John R. A. Gray of the
Hancock Museum for permission to study the holotype of
Paramekodon inflatus, Dr. J. P. Harding of the British Museum
(Natural History ) for permission to study the shell of a speci-
men of T. ctenorhynchus. I wish also to thank Mrs. Raymond
B. Manning for preparing text-figures for publication from my
penciled camera lucida drawings. Criticism of the manuscript
by Drs. Thomas E. Bowman and Raymond B. Manning are
greatly appreciated. I am grateful to Dr. Willis E. Pequegnat
and Mr. Thomas J. Bright, Department of Oceanography,
Texas A&M University, for the ostracods and station data
from the Gulf of Mexico. These were collected on the R/V
Alaminos under contract Nonr 2119 (04) of the Office of Naval
Research.
Listed below are known depths from which specimens have
been reported belonging to the three genera of Myodocopina
represented in the sample from the Gulf of Mexico.
Species Locality Depth (meters) Reference
Tetragonodon ctnoryhnchus Atlantic Ocean, 636-1485 Brady, 1884-1887
Mauritania to
Gibralta
Tetragonodon ctnoryhnchus Atlantic Ocean, off 92-152 Baker, (MS, 1965)
Puerto Rico
Tetragonodon ctnoryhnchus Colon Harbor, 7 Bold (1967)
Panama
Tetragonodon rhamphodes Gulf of Mexico 1000-1200 herein
Paramekodon inflatus Atlantic Ocean, off 1435 Brady and Nor-
Spanish Sahara man, 1896
Paramekodon poulseni Gulf of Mexico 1000-1200 herein
Bathyvargula parvispinosa West Indies, St. 900 Poulsen, 1962
Croix
Bathyvargula walfordi Tasman Sea 610 Poulsen, 1962
Bathyvargula optilus Gulf of Mexico 1000-1200 herein
Hydrographic data from Station XV slightly downslope of where
the ostracods described herein were collected are presented below:
Dissolved O2
Depth (m) Temp, °C Sal. %o ml/1
0 28.66 36.210 4.56
25 30.00 36.272 4.78
50 21.71 36.359 4.77
100 18.74 36.384 4.35
201 14.14 35.821 2.87
301 11.54 35.422 2.68
442, Proceedings of the Biological Society of Washington
401 9.40 35.118 2.76
601 6.84 34.888 2.94
801 5.48 34.895 3.00
1,000 4.84 34.935 4.18
1,251 4.39 34.977 4.45
Sediment: Foraminiferal lutite.
ORDER MYODOCOPIDA POKORNY, 1953
SUBORDER MYODOCOPINA SARS, 1866
SUPERFAMILY CYPRIDINACEA Barrp, 1850
Family Cypridinidae Baird, 1850
Bathyvargula Poulsen, 1962
Type-species: Bathyvargula parvispinosa Poulsen, 1962 Dana Report
No. 57, p. 216, figs. 103-104, designated herein. Gender: feminine.
Discussion of classification: Bathyvargula optilus n. sp., described
herein, has some morphological characters of the genus Paravargula
Poulsen, 1962, and others of the genus Bathyvargula. Characteristic of
Paravargula is the presence of spines along the ventral margin of the
bristle on the 2nd joint of the exopodite of the 2nd antenna. Character- -
istic of Bathyvargula are the reduced lateral eyes and the strongly de-
veloped medial spine in the terminal pair on the ventral margin of the
mandible. Both Paravargula and Bathyvargula have a pair of posterior
tusk-like processes on the upper lip and an unjointed or weakly 2-
jointed endopodite on each 2nd antenna. These two genera are very
closely related. I have assigned the new species described herein to
Bathyvargula rather than Paravargula primarily because of the strongly
developed medial spine of the mandible. The medial eye of B. optilus
is much larger than those on previously described species of both
Bathyvargula and Paravargula. As males of B. optilus are not present
in the collection it is not possible to ascertain whether they have blade-
like filaments of the sensory bristle of the lst antenna, which are
peculiar to the genus Bathyvargula.
Diagnosis (amended): This genus was defined by Poulsen, (1962,
p. 215). In order to include B. optilus, it is necessary to expand
the genus to include species having spines along the ventral margin
of the bristle on the 2nd joint of the exopodite of the 2nd antenna.
Bathyvargula optilus new species
Plate la—-f, 2a, b, Figures 1, 2
Etymology: The specific name is from the Greek optilos: eye, in
reference to the large middle eye on this species.
Holotype: USNM 122076; valves and some appendages in alcohol,
remaining appendages on slides; 2 with eggs in brood chamber.
Paratypes: USNM 122077; complete 2 with eggs in brood chamber in
alcohol. USNM 122078; juvenile, in alcohol.
ene
Gulf of Mexico Ostracods 443
e
PLATE 1. a-f, Bathyvargula optilus, holotype, L = 3.70mm: a, com-
plete specimen; b, complete carapace without soft parts, lateral
view; c, anterior left valve, medial view; d, caudal process left valve,
medial view; e, fifth limb; f, maxilla (same scale in microns: e, f.)
Type-locality: Gulf of Mexico; Lat. 28°13’N, Long. 87°16’W; on
bottom at depth of 1000-1200 M.
Description of female:
Shape (fig. la): Carapace oval in lateral view with greatest height
near middle; anterior with broadly rounded rostrum and fairly deep
444 Proceedings of the Biological Society of Washington
LEG. \
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Fic. 1. Bathyvargula optilus, complete specimen: a, outline show-
ing eggs, lateral eye and position of muscle scars; b, anterior, medial
view; c, shell pattern; d, caudal process, medial view; e, muscle scars
right valve, lateral view. Right lst antenna: f. medial view (bristles of
8th joint not shown); g, lateral view 8th joint. 2nd antenna: h, bristle
on 2nd joint of expodite; i, joints 6-9 left exopodite, lateral view; j, right
endopodite, medial view; k, joints 8-9 right exopodite, lateral view. Man-
dible: 1, tip of coxale endite; m, right medial view; n, claws of left end
joint, medial view; 0, ventral spine of 2nd joint of right endopodite; p,
ventral bristles of Ist joint of right endopodite; q, joints 2-3 of right
Gulf of Mexico Ostracods 445
incisur (fig. 1b); posterior with elongate caudal process below valve
middle (fig. 1d).
Ornamentation: Surface smooth, but faintly reticulate under high
magnification (fig. lc); hairs not present; shells thin, not strongly
calcified (no microconcretions observed).
Inner lamella: Infold broad in area of rostrum and caudal process,
narrower elsewhere; infold behind rostrum with about 21 bristles in-
cluding 1 long bristle on list (fig. 1b); infold of anteroventral and
ventral margins with about 16 short stout bristles followed by about
32 somewhat longer bristles being fairly equally spaced from incisure
to caudal process; list on infold in front of caudal process with about
18 blunt backward pointing processes followed by 4-5 short bristles
(fig. 1d).
Selvage: Wide lamellar prolongation with smooth outer margin present
along anterior and ventral margins, being quite broad in area of
incisur.
Muscle scars: Central muscle scars in anteroventral part of shell
indistinct, but consisting of about 12 individual scars (fig. le).
Size: @ holotype—length 3.70 mm, height 2.19 mm; @ paratype
with eggs in brood chamber—length 3.65 mm, height 2.01 mm; juvenile
paratype (probably @ )—length 2.97 mm, height 1.63 mm.
First antenna (fig. 1f, g): 3rd and 4th joints each with a dorsal
and ventral bristle; sensory bristle of 5th joint with 10 long proximal
filaments, 2 shorter distal filaments, and bifurcate tip; medial bristle of
6th joint longer than joint and with few marginal spines; 7th joints
with 3 bristles: a-bristle with few marginal spines and about same
length as bristle of 6th joint; b-bristle with 5 filaments, the proximal
2 having marginal teeth; c-bristle with 10 filaments; 8th joint with 4
bristles: d- and e-bristles bare and about half length of c-bristle and
about one and one-half length of b-bristle, f- and g-bristles each with
8-11 filaments and about same length as c-bristle; some filaments of c-,
f- and g-bristles with marginal teeth; surfaces of 2nd and 3rd joints with
short spines.
Second antenna (fig. Ih-k): Protopodite with short bare medial
bristle; endopodite weakly 2-jointed: Ist joint with 3 bare proximal
bristles, 1 long and 2 short, and 1 long distal bristle with spines; 2nd
joint short and with extremely long terminal bristle. Exopodite with
usual 9 joints: bristle of 2nd joint with 12-13 spines along ventral
margin, 1-2 minute spines, proximally on dorsal margin, and bifurcate
tip (fig. lh) reaching 7th joint; bristles of joints 3-9 with natatory
hairs and without spines; 9th joint with 3 long bristles and 1 shorter
<
endopodite, medial view; r, left lateral eye; s, genitalia; t, upper lip,
oblique view; u, medial eye above and rod-shaped organ below. (All
figures from holotype except e which is from paratype. Same scale in
IMUCKONS aes eA eb es jy Tat us nay my qos; k. os ps)
446 Proceedings of the Biological Society of Washington
dorsal bristle equal in length to joints 4-9; joints 2-8 with short teeth in
row along distal margins; joints 2-9 with basal spines; basal spines
smaller on proximal joints.
Right Mandible (fig. Il, m, o-—q): Coxale endite spinose with 2
terminal spines with marginal teeth; short process present at base of
endite; dorsal margin of basale with 1 bristle distal to middle and 2
terminally, all with short spines distally; ventral margin of basale with
2, spinous a-bristles, 1 long and 1 short, both on medial surface, 1 short
spinous b-bristle on lateral surface, 2 c-bristles, 1 short and bare, other
long and spinous, 2 d-bristles, 1 short with short spines, 1 long with long
spines; exopodite slightly longer than Ist joint of endopodite and with
2 spinous bristles; tip of exopodite hirsute. Endopodite: ventral margin
of Ist joint with 4 bristles, 2 long and 2 short, long bristles spinous,
short bristles bare, shorter of short bristles peglike and with filament at
tip; ventral margin of 2nd joint spinous and with 3 groups of bristles
having 1, 1, and 2 bristles, all short; medial bristle of terminal group
broad and with teeth along inner margin; dorsal margin of 2nd _ joint
with 7 long bristles and about 14 short bristles; end joint with 3
claws and 3 bristles: broad proximal part of ventral bristle with spines
along ventral margin; proximal part of ventral margin of lateral claw
with rounded tip, probably deformed; outer medial claw broken on
specimen examined.
Left Mandible: Similar to right mandible except for lateral claw of
end joint of endopodite not being deformed and outer medial claw
not being broken (fig. In).
Maxilla: (fig. 2a-e): Coxale with stout plumose bristle. Exopodite
large with 3 bristles: proximal bristle almost reaching tip of terminal
bristles and plumose proximally; 1 terminal bristle plumose proximally,
other with few short spines. Endopodite 2-jointed: 1st joint with cutting
edge having 3 large teeth (considerable variation between left and right
limbs of specimen examined), 2 alpha bristles consisting of long and
plumose outer bristles and shorter inner bristle with few short spines
along outer margin, and 3 beta-bristles, consisting of long outer bristle
and 2 short inner bristles, all with stout spines along inner margins;
2nd joint with 4 a-bristles consisting of 2 stout outer bristles having strong
spines along middle of inner margin and 2 slender and bare inner bristles,
and 3 b-, c-, and d-bristles, all with spines along inner margins.
Fifth limb (fig. 2f): Epipodial appendage with 49 hirsute bristles;
disto-anterior process on protopodite short and with lobate surface:
anterior group of bristles on Ist joint consisting of short bare bristle
and long spinous bristle; 1 hirsute bristle present between anterior
group of bristles and thumb-like process on protopodite; main tooth
with peg and 6 pectinate teeth; bristle with few marginal spines and
hairs present near peg; 2nd joint with 3 rows of pectinate bristles having
4 bristles in each row; 1 short spinous bristle present posterior to middle
Gulf of Mexico Ostracods 447
Fic. 2. Bathyvargula optilus: Right maxilla: a, medial view; b,
lateral view; c, lateral view showing exopodite; d, tooth on Ist joint of
endopodite. Left maxilla; e, tooth on Ist joint of endopodite, medial
view. Fifth limb: f, anterior view of left limb; g, h, posterior view of
right limb. Sixth limb: i, epipodial bristles on right limb; j, medial view
of left limb. Seventh limb: k, distal part; 1, enlargement of comb. m,
furca, right lamella; n, brush-shaped organ; 0, eggs. (All figures from
holotype. Same scale in microns: a-c, f, i-k, m, 0; d, e, g, h, n.)
448 Proceedings of the Biological Society of Washington
row; 2 very long and 1 short spinous bristles present anterior to the
pectinate bristles; 1 spinous bristle present near middle of disto-anterior
margin of 2nd joint; outer lobe of 3rd joint with 3 bristles: proximal
bristle with long spines proximally and short spines distally, terminal
bristles longer and bare; 4th joint united to 5th joint; 4th joint with
5 bristles, 5th joint with 2 bristles and spinous terminal process near
margin of 4th joint.
Sixth limb (fig. 2i, 7): 2-3 small bare bristles present in place of
epipodial appendage; Ist endite with 3 bristles, 2 short medial and 1
long terminal; 2nd endite with 3 short medial bristles and 2 long terminal
bristles; 3rd and 4th endites each with 3 terminal bristles; end joint
with 17 marginal bristles, spines in clusters along disto-lateral margin,
and hairs on medial surface.
Seventh limb (fig. 2k, 1): Distal group with 9 bristles, 5 ventral, 4
dorsal, each with 3-6 bells; proximal group with 4 bristles, 2 on each
side, each with 3-4 bells; terminal comb with 8 elongate teeth with
digitate tips; elongate incurved peg present opposite comb; teeth and
peg annulate and with marginal spines.
Furca (fig. 2m): Each lamella with 9 claws separated from lamella
by suture; 2nd claw almost same length as Ist, each remaining claw
smaller than previous claw; minute spines present along anterior margin
of each lamella and in clusters on lateral surface.
Genitalia and brush-like bristles: Genitalia with attached ovoid struc-
tures (spermatophores?) (fig. 1s). Brush-like cluster of 10 minute
bristles present in vicinity of genitalia (fig. 2n).
Upper lip (fig. It): Anterior part large, unpaired and with crenulate
margin; posterior tusks paired, each with broad base and glandular
openings on lateral surface.
Lateral eye (fig. Ir): Eyes small, pigmented, each with about 5
ommatophores.
Medial eye and rod-shaped organ (fig. lu): Medial eye large, pig-
mented, tapering anteriorly. Rod-shaped organ ovoid with narrow ridges
distally.
Eggs: Holotype with 31 oval eggs of approximately same size in
brood chamber (fig. la); in addition, about 14 minute eggs with
distinct nuclei present inside body (fig. 20). Paratype with about 12
eggs in brood chamber.
Comparisons: Only 2 species have previously been described in the
genus Bathyvargula—B. parvispinosa Poulsen, 1962, and B. walfordi
Poulsen, 1962. B. optilus differs from these species in having spines
along the ventral margin of the bristle on the 2nd joint of the 2nd
antenna, a larger medial eye, more claws on the furca, and a considerably
larger carapace.
Family Philomedidae Miiller, 1912
Discussion of classification: In 1967 (Kornicker, 1967) I proposed
the family Pseudophilomedidae containing the genera Pseudophilomedes
Gulf of Mexico Ostracods 449
and Paramekodon. One of the morphological criteria upon which the
new family was established was the presence of only 2 endites on the
maxilla compared to 3 in genera of Philomedidae. Further studies of
the species Pseudophilomedes foveolatus has convinced me that what I
described as a short lobe with 3 bristles located distally of the coxale
endite (Kornicker 1967, p. 15) is probably a vestigal 3rd endite. I have
learned also from the present study that each maxilla of Paramekodon
inflatus and Paramekodon poulseni has a 3rd endite at least half the
length of the 2nd endite. Therefore, the Pseudophilomedidae is reduced
to subfamily status herein, and along with the Philomedininae comprise
the family Philomedidae.
The elongate tooth of the 2nd joint of the female 5th limb, the
relatively few bristles on the basale and endopodite of the mandible,
the reduced 3rd endite of the maxilla and the relatively few bristles
of the endites, the elongate process on the end joint of the maxilla
of some species, the relatively few bristles on the infold of the shell
behind the rostrum and the hirsute bristles on the infold in front of the
caudal process serve to distinguish the Pseudophilomedinae from the
Philomedinae.
Subfamily Philomedinae Miiller, 1912
Tetragonodon Brady and Norman, 1896
Type-species: Bradycinetus ctenorhynchus Brady, 1887 Fonds Mer,
vol. 4, p. 199, pl. 12, figs. 3-5 (pl. 12 is missing from volumes I ex-
amined and may not have been published), designated by Sylvester-
Bradley 1961, p. Q400. Gender: Masculine.
Type-locality: Atlantic Ocean, off Spanish Sahara.
Bradycinetus Sars, 1866 (part): Brady, 1887, p. 199.
Philomedes G. W. Miiller, 1894 (part): Muller, 1912, p. 25.
Tetragonodon Brady and Norman, 1896, p. 667: Sylvester-Bradley 1961,
p. Q400.
Discussion of classification: Muller (1912, p. 25) referred Tetra-
gonodon to Philomedes. Skogsberg (1920, p. 348) did not entirely
agree with Miiller, and stated, “With regard to Tetragonodon it does
not seem impossible to me that it must be regarded as a special unit,
perhaps as a subgenus of the genus Philomedes. These questions can,
however, only be decided after a renewed investigation of these forms.”
Poulsen (1962, p. 339) stated, “Obviously it is quite correct to include
Tetragonodon in Philomedinae, but the descriptions of the species are
so insufficient that the position within the sub-family cannot be deter-
mined. The reduced endopodite of the 2nd female antenna may indicate
a closer relation to the genus Paraphilomedes.”
The endopodite of the 2nd antenna of Tetragonodon rhamphodes,
described herein, is 2-jointed, not reduced as in T. ctenorhychus. It
differs from other species of Philomedinae having a 2-jointed endopodite
in bearing only 1 bristle on the Ist joint. It also differs from most
species in the subfamily in having only 1 bristle on the 2nd joint of the
450 Proceedings of the Biological Society of Washington
lst antenna, a squarish tooth on the 2nd joint of the 5th limb, and
fewer than 10 bristles on the end joint of the 6th limb. The carapace
has a characteristic beak-like rostrum, and the infold behind the rostrum
bears only 4 long bristles. These differences are of sufficient magnitude
to warrant retention of the genus.
Diagnosis of Tetragonodon (emended):
Carapace with beak-like rostrum and a caudal process. Infold behind
rostrum with only 4 long bristles; infold in front of caudal process with
leaf-like bristles. First antenna with only 1 bristle on 2nd joint. Second
antenna: endopodite 1-2 jointed and with total of 1 to 3 bristles;
bristles of exopodite without marginal spines or natatory hairs, and
relatively short; 9th joint of exopodite with only 3 bristles. Second joint
of 5th limb with large quadrate tooth. End joint of 6th limb not pro-
longed posteriorly and bearing fewer than 10 bristles. Furca with 9-10
claws, all separated from lamella by suture.
Discussion of type-species: Brady (1884-1887, p. 164) presented a
list of ostracods collected by the Travailleur and Talisman expeditions
of 1881 to 1883. The list contains the taxon “Bradycinetus Stenorynchus
Id. n. sp.,’ which was collected at 8 stations. Brady (1887, p. 199)
described the species “Bradycinetus ctenorynchus.” He did not indicate
that it was a new species, se presumably it is the same species previously
listed, but with a different spelling. The description refers to Plate
XII, but this plate is missing from three volumes I have examined and
may not have been published.
Brady and Norman (1896, p. 667-669) proposed a new genus
Tetragonodon containing two species, T. ctenorhyncus and T. erinaceus.
Both were noted as being new species, but presumably the first was
previously described by Brady in 1884-1887 (Although, as discussed
below, the date of collection given by Brady and Norman for T.
ctenorynchus does not agree exactly with that given by Brady for B.
ctenorynchus). Miller (1912, p. 33) considered them to be synonyms,
and Sylvester-Bradley (1961) designated B. ctenorynchus as the type-
species of Tetragonodon.
Brady (1884-1887, p. 166) listed the following 8 stations at which
specimens of Bradycinetus Stenorynchus [sic] (= T. ctenorynchus ) were
collected by the Travailleur and Talisman:
Date Depth (m)
1882 11 July 932
1882 25 July 770
1882 7 July 1976
1882 I July 1485
1882 18 July 932
1882 30 July 636
1882 29 Aug. 2995
1883 14 July 1485
Gulf of Mexico Ostracods 451
Collections were made by the Travailleur in 1880-1882 and by the
Talisman in 1883. Accordingly, the first 7 dates listed above refer to
collections of the Travailleur and the last date to a collection of the
Talisman.
Brady and Norman (1896, p. 669), following the description of
T. ctenorynchus state: “The exact locality is unknown to us. The
specimens were dredged by the “Talisman, July 11, 1883, in 932
metres, and received by us from the Marquis de Folin; the data would
lead us to suppose that T. ctenorhynchus was taken off the coast of
Morocco.”
Data in the preceding paragraph is in agreement with that on the
label of a specimen (1900-3-6-454) in the British Museum which
carries the following information: “Tetragonodon ctenorhynchus Brady
and Norman, Type @, Talisman, 11-7/83, 932 meters, Marquis de
Folin.”
The day, month, and depth in Brady and Norman’s 1896 statement
and on the label of the specimen in the British Museum agree with the
first item in Brady’s list (11 July, 932 m), but do not agree in the
year. The list has 1882 instead of 1883.
Smith (1888, p. 111) listed the stations dredged by the Talisman
in 1883. Five dredgings were made on July 11 but none at 932 m.
Six were made on July 12; the first of these was from 932 m. If this
is the station from which the specimen was collected, it is necessary
to conclude that Brady and Norman (1896) and the label at the
British Museum is off 1 day, and that Brady’s list for this date is off a
year and a day. As the Travailleur did not collect on July 11 (or 10)
and did not collect at a depth of 932 m on July 12 (Anonymous, 1883,
p. 33), it is probably that the holotype was collected by the Talisman
on July 12, 1883.
The position of the Talisman during the collection made on July 12,
1883 at 932 m was Lat. 23°00’N, Long. 17°30’W (Smith 1888, p. 111).
This station was off the coast of Spanish Sahara, not Morocco. The
bottom consisted of greenish muddy sand; bottom temperature was 7.0°C.
In Table 1, I have attempted to correct Brady’s list of stations at which
specimens of T. ctenorhynchus were collected.
Baker (1965, MS) reported T. ctenorynchus from the outer shelf
(92-152m) off Porto Rico. About 56 specimens were collected, but all
without “soft parts.” Until specimens with appendages are collected
from that locality, I think it necessary to consider the specific identifica-
tion doubtful. Dr. N. C. Hulings kindly allowed me to measure the
right valve figured by Baker, (pl. X, fig. 15) and I found it to have a
length of 0.88 mm, and height of 0.50 mm (height as percent of length
56.8). This is much smaller than the right valve of the British Museum’s
specimen of T. ctenorynchus (length 1.83, height 1.18; height as percent
of length 64.5), but the valves could be from specimens at different
stages of development. The left valve of the British Museum specimen
452. Proceedings of the Biological Society of Washington
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Gulf of Mexico Ostracods 453
PuateE 2. a, b, Bathyvargula optilus, holotype: a, 6th limb; b,
frontal organ (below), medial eye (above). c-e, Tetragonodon rham-
phodes, holotype, L = 2.11 mm: ec, right valve, medial view (calcareous
microconcretions formed while specimen was in glycerine); d, anterior
left valve, lateral view; e, caudal process left valve, lateral view. f, Con-
choecia atlantica, complete specimen, L = 3.86 mm. (Scale in microns. )
has a length of 1.90 mm and height of 1.26 mm (height as percent of
length 66.3). Measurements of both the Porto Rico and British Museum
specimens are based on dry valves. Brady (1887, p. 199) gave the length
454 Proceedings of the Biological Society of Washington
Fic. 3. Tetragonodon rhamphodes, complete specimen: a, outline
showing representative surface pits. Left valve: b, medial view; c, caudal
process, medial view; d, anterior, lateral view; e, anterior, medial view;
f, surface structure near posterior, lateral view. Left Ist antenna: g,
medial view (b-g bristles not shown); h, tip of b-bristles; i, bristle of end
joints, medial view; j, enlargement of bristles on joints 7-8, medial view.
Second antenna: k, exopodite of right limb, medial view; 1, medial view
showing endopodite; m, enlargement of spine on Ist joint of exopodite;
n, joints 8—9 of right limb, lateral view; 0, joint 8-9 of left limb, medial
view. (All figures from holotype. Same scale in microns: a, b; c-f, i,
I, Ie im, sg In, Fy st)
Gulf of Mexico Ostracods 455
Fic. 4. Tetragonodon rhamphodes: a, endopodite of right 2nd an-
tenna, medial view. Mandible: b, left limb, medial view; c, enlargement
of tip of exopodite, medial view; d, enlargement of 2 terminal claw of
right limb showing spines, lateral view. Maxilla: e, right limb, lateral
view; f, terminal end right limb, medial view; g, lateral view. Fifth
limb: h, anterior view right limb; i, endopodites I, II of left limb; j,
posterior view left limb; k, main tooth of Ist joint on right limb; |, an-
terior view left limb. (All figures from holotype. Same scale in microns:
a, e-j; 1; Cc, d, k.)
as 2.2 mm; Brady and Norman (1896, p. 668) gave length as 2 mm
and height as 1.3 mm (height as percent of length 65).
Bold (1967, p. 44) reported a specimen of T. ctenorhynchus from
Colon Harbor, Panama. Through the courtesy of Drs. Lewis G. Nichols
456 Proceedings of the Biological Society of Washington
and Joseph E. Hazel, I had the opportunity to examine and measure
this specimen—a left valve with tip of caudal process missing (No. 8286,
Louisiana State University). The length is 0.75 mm; height 0.48 mm
(height as percent of length—64). The specimen is much smaller than
that described by Brady (1887). The condition of the valve and
absence of soft parts does not permit certain specific or generic identifica-
tion. Its reticulate ornamentation is very similar to that of the specimen
described by Baker (1965, MS) and T. rhamphodes described herein.
Tetragonodon rhamphodes new species
Plate 2c-e, Figures 3, 4, 5a-e
Etymology: The specific name is from the Greek rhamphodes: beak-
like, in reference to the slender rostrum of the carapace.
Holotype: USNM 122073, 2 without eggs; valves and some ap-
pendages in alcohol, remaining appendages on slides.
Type-locality: Gulf of Mexico; Lat. 28°13’N, Long. 87°16’W; on
bottom at depth of 1000-1200 M.
Description of female:
Shape (fig. 3a): Carapace oval in lateral view with greatest height
in front of middle; anterior with long beak-like rostrum; posterior with
caudal process below middle of valve; shell forming lateral fold over
rostrum (fig. 3d, e).
Ornamentation: Surface with rather large punctae, otherwise smooth
(fig. 3f); hairs fairly abundant over valve surface, especially near edge,
those on posterior surface of valve being longer than elsewhere.
Inner lamella (fig. 3b): Infold broad in area of rostrum and caudal
process, narrower elsewhere; infold behind rostrum with 4 long bristles
in row and 2 smaller bristles at lower margin, infold immediately
ventral to rostrum with 1 small bristle; infold in front of caudal process
with 5 leaf-like bristles near top and 3 small bristles near lower inner
margin (fig. 3c); striations present on infold along anteroventral margin.
Selvage: Wide lamellar prolongation with slender spines present
along anterior, ventral, and posterior margins of each valve; prolongation
divided below rostrum.
Size: @ holotype—length 2.11 mm, height 1.32 mm, height as per-
cent of length 62.6.
First antenna (fig. 3g-j): Ist joint with short spines in clusters on
lateral surface; 2nd joint with 1 spinous dorsal bristle; 3rd joint with
1 ventral and 2 dorsal bristles, all with spines; 4th joint with 2 spinous
bristles, 1 ventral and 1 dorsal; joints 2—4 with spines in clusters on
medial and lateral surfaces; 5th joint with ventral sensory bristles having
3 proximal and 4 terminal filaments; 6th joint with short spinous
bristle medially; 6th joint not separated from 5th by visible suture;
7th joint with spinous a-bristle about twice length of bristle on 6th
joint, b-bristle with 1 proximal and 3 terminal filaments, c-bristle with
3 proximal and 3 terminal filaments; 8th joint width d- and e-bristles
Gulf of Mexico Ostracods 457
of equal length, and f- and g-bristles, each with 2 proximal and 3
terminal bristles.
Second antenna (figs. 3k-0; 4a): Endopodite 2-jointed: Ist joint
with 1 short spinous bristle; 2nd joint with long spinous anterior bristle
and short terminal spine-like bristle. Exopodite: distal margin of elongate
Ist joint with short medial spine with blunt tip; bristle of 2nd joint
reaching past 9th joint, joints 2-8 with comb of short spines along
distal margin and 2 or more short spines on ventral corner; bristles of
joints 2-9 without spines or natatory hairs and relatively short; 9th joint
with 1 short and 2 long bristles (9th joint and short bristle of joint on
right appendage smaller than on left).
Mandible (fig. 4b-d): Coxale endite with bifurcate tip and spinous.
Basale with 12 bristles: dorsal margin with 1 near middle and 2 terminal;
ventral margin with 4 having long and short marginal spines; medial
surface with 4 in cluster proximally and 1 nearer middle. Exopodite
with hirsute tip reaching past middle of Ist joint of endopodite and
with 2 spinous bristles, outer bristle longer than inner bristle.
Endopodite: 1st joint with 3 spinous bristles on ventral margin, 1
short, 2 long; ventral margin of 2nd joint with 2 subterminal and 2
terminal bristles; dorsal margin of 2nd joint with 2 spinous bristles
proximally, and 5 bristles near middle; end joint with 2 large subequal
claws with few minute teeth along concave margin (fig. 4d), 1 short
spine-like claw at dorsal corner, and 3 bristles. Basale and 2nd _ joint
of endopodite with spines in clusters on medial and lateral surfaces.
Maxilla (fig. 4e-2): Precoxale and coxale with anterior fringe of
hairs; coxale with long hirsute anterior bristle; 1st endite with 6 spinous
bristles; 2nd endite with 7 bristles; 3rd endite with 7 terminal and 1
proximal bristle; basale with 3 bristles along distal margin; exopodite
with 1 short and 2 long bristles. Endopodite: Ist joint with 1 alpha-
bristle and 2 beta-bristles; 2nd joint with 3 a-, 2 b-, 2 c-, and 3 d-
bristles.
Fifth limb (fig. 4h-l): Main tooth of Ist joint consisting of distal
tooth with small triangular tooth proximally on anterior surface, and 2
teeth with small anterior spine-like teeth; small tusk-like tooth present
proximal to main tooth; distal margin of Ist joint with 2 spinous bristles;
short spinous bristle present on anterior surface of Ist joint near main
tooth; 2 long stout bristles present near tusk-like tooth of 1st joint; 2nd
joint with large quadrate tooth; 3rd joint with 2 spinous outer bristles
and 3 inner bristles; 4th joint with 8 spinous bristles; Ist endite with
2 or 3 bristles; 2nd endite with about 9 bristles, 3rd endite with about
6 bristles; epipodial appendage with about 49 hirsute bristles.
Sixth limb (fig. 5a, b): Ist endite with 3 short bristles; 2nd endite
with 3-4 spinous bristles; 3rd and 4th endites each with 8—9 spinous
bristles. End joint with total of 8-9 bristles: 5-6 bristles followed by
space, then, a single bristle followed by another space and 2 long hirsute
bristles. Surfaces of appendage hirsute.
458 Proceedings of the Biological Society of Washington
Fic. 5. Tetragonodon rhamphodes: 6th limb: a, right limb, medial
view; b, left limb, medial view. Seventh limb: c, distal part; d, en-
largement of comb. Furca, e, lateral view of left lamella and medial view
of claw 1 of right lamella. Paramekodon inflatus right valve: f, outline;
g, incisure left valve, medial view; h, surface pits; i, rostrum right valve,
lateral view; j, caudal process right valve, lateral view. First antenna: k,
end joints left limb, medial view; 1, medial view. Second antenna: m,
joints 2-3, left exopodite, lateral view; n, left limb, lateral view; 0,
endopodite of right limb, medial view. (All figures from holotype. Same
scale in microns: a-c, e, h, 1; g, i, j, m, 0; d, k.)
Gulf of Mexico Ostracods 459
Fic. 6. Paramekodon inflatus mandible: a, right limb, lateral view;
b, enlargement of coxale endite; c, distal end left limb, lateral view.
Maxilla: d, left limb, medial view; e, right limb, medial view; f, enlarge-
ment of distal end left limb, medial view. Fifth limb: g, posterior view
right limb; h, posterior view left limb. i, 6th limb; j, 7th limb, distal
part; k, enlargement of terminus; 1, furca. (All figures from holotype.
Same scale in microns: a, i, j, 1; b-e, g, h; f, k.)
Seventh limb (fig. 5c, d): 4 bristles in distal group, 2 on each side;
6 bristles in proximal group, 3 on each side; all bristles with 1-4
bells and marginal spines; terminus consisting of comb with 12 teeth
opposed by 3 pegs.
Furca (fig. 5e): Each lamella with 10 claws, each separated from
460 Proceedings of the Biological Society of Washington
lamella by suture; claw 1 with lateral and medial spines; distal medial
spine of claw 1 being extremely large; claws 2-9 with sharp spines
along posterior margins; claws 4-7 with few short spines along anterior
margins; each lamella with hairs along anterior and ventral margins.
Eyes: Lateral eyes absent; medial eye and rod-shaped organ not
observed, probably lost during dissection.
Ecology: The relatively short stout bristles without natatory hairs on
the exopodite of the 2nd antenna indicate that the females of the species
are incapable of swimming, and therefore must dwell on the bottom.
Comparisons: Only 2 species have been referred to Tetragonodon—
T. ctenorhynchus (Brady, 1886?) and T. erinaceus Brady and Norman,
1896. The carapace of T. rhamphodes differs from the former in having
a rostrum that is much less acuminate, and from the latter in having
a punctate surface, rather than one covered with the acute spinous
processes described by Brady and Norman (1896, p. 669). T. erinaceus
is known only from its shell and was correctly considered species dubia
by Miller (1912, p. 51). The endopodite of the 2nd antenna of T.
ctenorhynchus is single jointed and bears only 1 bristle (Brady and
Norman, 1896, p. 668) whereas, the endopodite of T. rhamphodes is two
jointed and bears 3 bristles. Specimens of T. ctenorhynchus described
by Baker (1965, MS) and Bold (1967) are known only from their
shells which apparently are considerably smaller than T. rhamphodes.
Subfamily Pseudophilomedinae Kornicker, 1967
Paramekodon Brady and Norman, 1896
Type-species: Paramekodon inflatus Brady and Norman, 1896, p. 67,
pl. 59, figs. 1-10, by monotypy. Gender: masculine.
Paramekodon Brady and Norman, 1896, p. 670.
Pseudophilomedes G. W. Miiller, 1894 (part): Miller, 1912, p. 33.
Pseudophilomedes G. W. Miiller, 1894 (part): Skogsberg, 1920, p. 348
Pseudophilomedes G. W. Miiller, 1894 (part): Sylvester-Bradley, p.
Q399.
Paramecodon Brady and Norman, 1896: Poulsen, 1962, p. 339, p. 345.
Pseudophilomedes G. W. Miiller, 1894 (part): Kornicker, 1967, p. 3.
Discussion of classification: With the exception of Poulsen (1962,
p. 399) most investigators have considered Paramekodon to be a synonym
of Pseudophilomedes G. W. Miiller, 1894 (see discussion in Kornicker,
1967, p. 3). I recognize the validity of the genus herein not on the
basis of the diagnosis given by Brady and Norman (1896, p. 670), which
is essentially the same as that given for Pseuwdophilomedes by Miller
(1894, p. 211), but because of the absence of a b-bristle on the 7th
joint of the Ist antenna of known species of Paramekodon. It is ap-
parent from Brady and Norman’s description and illustration of the
Ist antenna of P. inflatus that it lacks a b-bristle; however, they did
not mention this detail in their diagnosis. A b-bristle is present on the
Gulf of Mexico Ostracods 461
Ist antenna of species of Pseudophilomedes, and I am unaware of any
other genus in the Cypridinacea lacking this bristle.
Diagnosis (emended): 7th joint of Ist antenna without b-bristle;
natatory bristles of 2nd antenna with ventral spines proximally and long
hairs distally: endite III of maxilla at least half length of endite II.
Sth joint: inner margin of distal tooth of Ist joint with 2 pectinate
teeth (always?); inner margin of 2nd joint with at least 1 quadrifid
tooth.
Paramekodon inflatus Brady and Norman,1896
Figures 5f-o, 6
Holotype: @, appendages on 1 slide, valves on another at Hancock
Museum, Newcastle-on-Tyne; both slides contain labels with following
information: “Paramekodon inflatus, type, 1435M, 9 July 1883, Travail-
leur.” Slides are without numbers. Holotype by unique specimen.
Type-locality: Atlantic Ocean, off Spanish Sahara, at depth of 1435m.
Paramekodon inflatus Brady and Norman, 1896, p. 670, pl. 59, figs. 1-10.
Pseudophilomedes inflata (Brady and Norman, 1896): G. W. Miller
1912, p. 34 [diagnosis, key].
Material: Two slides from the Hancock Museum, Newcastle-on-Tyne,
England, containing appendages and valves of holotype. An anastomising
pattern of air chambers surrounds most appendages, but details of
morphology are discernible. The valves are somewhat compressed on
the slide and details difficult to perceive; the left valve is fragmented.
Discussion of type-species: In their description of the species, Brady
and Norman (1896, p. 672) state, “A single specimen was dredged
by the “Talisman’ off the coast of Morocco, July 6, 1883, in 1434 métres
(Marquis de Folin).” According to the list of dredging stations as-
sembled by Smith (1888, p. 983), the Talisman did not dredge in the
area of Morocco on July 6, in fact, the ship may not have dredged
anywhere on that day.
As noted above, labels on the microscope slides containing the holotype
bear the following information: “1435m, 9 July 1883, Travailleur.”
This information differs from that given by Brady and Norman (1896)
in the depth (1435 m compared to 1434m), the date (July 9 compared
to July 6) and the ship (Travailleur instead of Talisman). As the
Travailleur did not collect in 1883, it is apparent that either the ship
name or the year is incorrect on the labels.
The Talisman made 7 dredgings on July 9, 1883; one of these was
at 1435 m, which coincides with depth information given on the label
of the holotype, and is only 1 meter off the depth given by Brady
and Norman (1896) for the depth at which their single specimen was
collected. The Travailleur did not collect deeper than 614 m on either
July 6 or July 9, 1882. Therefore, I conclude tentatively that the date
of collection stated in their paper as July 6 should have been July 9
462 Proceedings of the Biological Society of Washington
(possible the number 9 was inverted in printing); and that the collection
was made aboard the Talisman, not Travailleur.
The position of the station dredged on July 9 at 1435 m was Lat.
25°39'N Long. 16°06’W, which is off Spanish Sahara, and the bottom
was composed of muddy sand, corals, and shells (Smith, 1888, p. 983).
Redescription of holotype:
Shape (fig. 5£): Carapace oval in lateral view with greatest height
near middle, quite broad in dorsal view; lower margin of rostrum
forming obtuse angle with anterior margin of valve below rostrum;
posterior margin with caudal process below middle.
Ornamentation: Surface with punctae, otherwise smooth (fig. 5h);
hairs scattered over valve surface and along margins.
Inner lamella: Infold behind rostrum with 4 long spinous bristles
in row and 2 small bristles at lower margin (fig. 5i); infold in front
of caudal process with about 8 pores in row (Some of these probably
contain hirsute setae (fig. 5j), but preservation of valve is too poor
to see them).
Selvage: Wide lamellar prolongation present with slender spines
along outer margin (fig. 5g).
Size: @ holotype—measured length of compressed valve under slide
1.69 mm. Length of dried specimen—1.73 mm, height 1.07 mm (from
Brady and Norman, 1896, pl. 49, fig. 1, using scale of 30 ).
First antenna (fig. 5k, 1): 2nd joint with 1 spinous dorsal bristle; 3rd
joint with 1 long ventral and 1 short dorsal bristle, both with marginal
spines; 4th joint with 3 bristles, 2 ventral and 1 dorsal, all with spines;
joints 2-4 with clusters of spines on surfaces; 5th joint with sensory
bristle having about 4 filaments; 6th joint minute, not sharply demarked
from 5th joint, and with 1 short spinous bristle medially; 7th joint
reduced and fused to 8th, with spinous a-bristle about twice length of
bristle on 6th joint, no b-bristle, and c-bristle with about 5 filaments;
8th joint with long bare d- and e-bristles of almost equal length, and f-
and g-bristles with about 4 filaments (filaments on bristles of end joints
could not always be seen clearly on specimen).
Second antenna (fig. 5m-o): Endopodite single jointed with 3
short spinous bristles proximally and 1 long hirsute bristle terminally.
Exopodite: distal margins of joints 2-8 with combs of slender spines;
bristles on joints 2-8 with a few hairs proximally on dorsal margin
followed by fairly long spines along the ventral margins and then long
natatory hairs along both margins; 9th joint with 2 bristles, 1 long with
spines proximally along the ventral margin and natatory hairs distally
along both margins and 1 short with marginal spines.
Mandible (fig. 6a—-c): Coxale endite bifurcate with each prong
widely separated from the other and spinous; outer prong acuminate
and with spine at tip, inner prong rounded at tip and more spinous
than outer prong; basale with 7 spinous bristles, 4 ventral, 3 dorsal, and
with spines in clusters on medial and lateral surfaces; exopodite with 2
Gulf of Mexico Ostracods 463
bristles, outer bristle about half length of inner bristle. Endopodite:
Ist joint with 3 spinous bristles on ventral margin, 1 long, 2 short; 2nd
joint with 3 bristles near middle of dorsal margin, 1 short bristle sub-
terminal on ventral margin, and 2 short terminal bristles on lateral
surface near ventral margin, all bristles with spines; end joint (fig. 6c)
with 1 long claw with minute spines distally along concave margin, and
total of 4 bristles; clusters of spines present on surfaces of Ist and 2nd
joints of endopodite.
Maxilla (fig. 6d-f): Coxale with long hirsute anterior bristle and
fringe of hairs along outer margin; endite I with 3 short stout pectinate
spines, 1 long stout spinous bristle, and 2 short widely separated spinous
bristles; endite II with 2 stout pectinate spines and 2 slender bristles;
endite III with 1 long proximal bristle and 3 distal bristles, 1 short
and 2 long; basale with 3 long bristles along distal margin; exopodite
consisting of 3 bristles, 1 short and 2 long. Endopodite: Ist joint with
1 alpha and 1 beta-bristle, both with spines; end joint with 2 a-, 1-2 b-
bristles, and 1 long stout c-bristle.
Fifth limb (fig. 6g, h): (Both appendages are mounted on slide with
posterior up so that teeth of Ist joint are not clearly visible; the ap-
pendage is apparently quite similar to the same appendage of
Paramekodon poulseni described in this paper): Ist joint with large
pectinate proximal tooth and distal tooth having large tooth followed
by 2 smaller teeth. 2nd joint: inner margin with large fang-like tooth
followed by small tooth with 2-3 prongs, 1 trifid or quadrifid tooth,
and then 1 quadrifid tooth; posterior surface with 3 spinous bristles, 2
near middle and 1 proximal to them. 3rd joint with a pair of spinous
bristles; 4th joint with 4 spinous bristles, 3 long and 1 short; epipodial
appendages with 40-44 hirsute bristles.
Sixth limb (fig. 61): 1st endite represented by 3 short bristles, 2nd
endite represented by short lobe with 1 spinous bristle; 3rd endite with
5 spinous bristles; 4th endite with 4 spinous bristles (1 missing in fig.
6i); end joint with 6 spinous anterior bristles followed by a space and
then 2 long hirsute bristles; 1 short bristle present in place of epipodial
appendage; surface of appendage hirsute.
Seventh limb (fig. 6j, k): 6 bristles in distal group, 3 on each side;
2 bristles in proximal group, 1 on each side; all bristles with 1-5 bells
and marginal spines distally; terminus consisting of comb with single
tooth having a spinous tooth at each side facing a similar but slightly
smaller comb.
Furca (fig. 61): Each lamella with 10 claws separated from lamella
by suture; 3rd claw smaller than 4th; claws 1, 2, 4 with backward point-
ing spines in row on lateral and medial sides and slender hairs along
convex margins; hairs present on margins of lamellae at base of claws
and following claws.
464 Proceedings of the Biological Society of Washington
Paramekodon poulseni new species
Figures 7-9
Etymology: The species is named in honor of Prof. Erik M. Poulsen
who has contributed greatly to the study of myodocopid Ostracoda.
Holotype: Adult 2 USNM 121760; valves and some appendages in
alcohol, remaining appendages on slides.
Type-locality: Gulf of Mexico; lat. 28°13’N, long. 87°16’W; on or
near bottom at depth of 1000-1200 M.
Diagnosis: Paramekodon with 2 proximal bristles on endopodite of
2nd antenna, quadrifid tooth in middle of inner margin of 2nd joint of
5th limb, 3 bristles on endite I of 6 limb, and 9 claws on each lamella
of furca.
Description of adult female:
Shape (fig. 7a, b): Carapace oval in lateral view with greatest height
and length near middle, quite broad in dorsal view, width about 4
length; lower margin of rostrum forming obtuse angle and continuous
with anterior margin of valve below rostrum; posterior margin of valve
with caudal process below valve middle.
Ornamentation (fig. Te): Surface with small punctae, otherwise
smooth; hairs scattered over valve surface and along margins.
Inner lamella: Infold broad in area of rostrum, ventral margin and
caudal process, narrower along posterior margin; infold behind rostrum
with 4 long spinous bristles in row and 2 small bristles at lower margin
(fig. 7£); infold immediately ventral to rostrum with 1 small bristle;
infold in front of caudal process with 6 hirsute setae in row near middle
and cluster of 2 small bristles at inner edge (fig. 7c); a cluster of 2
small bristles present at inner margin of infold above and below caudal
process; striations present on infold along ventral margin.
Central muscle scars (exact number and position could not be deter-
mined with certainty): consisting of about 20 individual scars below
and anterior to center of each valve (fig. 7d, g).
Selvage: Wide lamellar prolongation with slender spines along outer
margin along anterior, ventral, and posterior margins of each valve;
prolongation divided below rostrum.
Size: @ holotype—length 1.81 mm, height 1.21 mm.
First antenna (fig. 7h, i): 2nd joint with 1 spinous dorsal bristle;
3rd joint with 1 long ventral bristle and 1 short dorsal bristle, both
with marginal spines; 4th joint with 1 ventral and 1 dorsal bristle, both
with spines; joints 2-4 with clusters of spines on surfaces; 5th joint with 1
long sensory bristle with filaments, spines near base of sensory bristle;
6th joint minute and not separated from 5th by suture, with 1 short
spinous bristle medially; 7th joint reduced and fused to 8th, with
spinous a-bristle about twice length of bristle on 6th joint, no b-bristle,
and c-bristle with about 3 filaments and forked tip; 8th joint with long
bare d- and e-bristles of equal length, f-bristle with 2 proximal and 3
Gulf of Mexico Ostracods 465
SET ERTS
iN) we
\! K
VG
NIAC
Fic. 7. Paramekodon poulseni complete specimen: a, outline show-
ing position of muscle scars; b, dorsal view showing representative sur-
face pits (valves slightly open) c, caudal process left valve, medial view;
d, muscle scars right valve, lateral view; e, surface pits near middle of
right valve; f, anterior left valve, medial view; g, muscle scars left
valve, lateral view. First antenna: h, lateral view right limb; i, medial
view distal end left limb. Second antenna: j, medial view right limb;
k, enlargement of joints 1—3; 1, medial view endopodite right limb. (All
figures from holotype. Same scale in microns: a, b; d-h; c, k, 1.)
466 Proceedings of the Biological Society of Washington
Fic. 8. Paramekodon poulseni mandible: a, medial view right limb;
b, enlargement of distal end; c, enlargement of coxale endite; d, lateral
view of basale of right limb. Maxilla: e, lateral view right limb; f, en-
largement of distal end; g, medial view endite I of right limb; h, medial
view endite I left limb. Fifth limb: i, posterior view left limb; j, anterior
view right limb; k, enlargement of tip of tooth on 2nd joint; 1, posterior
view right limb. (AIl figures from holotype. Same scale in microns: a,
d; b, C, €, h-j, Ie fe g, k.)
Gulf of Mexico Ostracods 467
Fic. 9. Paramekodon poulseni: a, fifth limb, posterior tooth on inner
margin of Ist joint of right limb. Sixth limb: b, medial view left limb;
c, lateral view right limb (endite I displaced; hairs on bristles not
shown). Seventh limb: d, distal part; e, f, lateral and medial view of
terminus. g, furca; h, medial eye, rod-shaped organ, anterior end of
body; i, genitalia; j, brush-shaped organ. (All figures from holotype.
Same scale in microns: a, e, f; b-d, g, h; i, j.)
distal filaments and forked tip, g-bristle with 1 short proximal filament
and 1 long and 2 short distal filaments, tip forked.
Second antenna (fig. 7j-1): Endopodite 2 jointed with 2 short spinous
bristles on joint 1, and 1 long hirsute terminal bristle on joint 2.
Exopodite: Distal margin of Ist joint with short medial spine; distal
margins of joints 2-8 with combs of slender spines; bristles on joints
2-8 with a few hairs proximally on dorsal margin followed by fairly
long spines along the ventral margin and then natatory hairs along both
margins; 9th joint with 2 bristles, 1 long with spines proximally along
the ventral margin and natatory hairs distally along both margins, and 1
short with marginal spines.
Mandible (fig. 8a—d): Coxale endite with bifurcate tip with each
prong widely separated from the other and spinose; outer prong
acuminate and with spine at tip, inner prong rounded at tip and more
spinous than outer prong; basale with 6—7 spinous bristles, 3-4 ventral,
3 dorsal, and with spines in clusters on medial and lateral surfaces;
exopodite with 2 spinous bristles, outer bristle about half length of inner
bristle. Endopodite: Ist joint with 3 spinous bristles on ventral margin,
1 long, 2 short; 2nd joint with 3 long spinous bristles near middle of
dorsal margin, 1 short spinous bristle subterminal on ventral margin,
468 Proceedings of the Biological Society of Washington
Fic. 10. Conchoecia atlantica, complete specimen: a, outline; b,
dorsal view (valves open as in ventral view); c, ventral view; d, endop-
odite left 2nd antenna, medial view (all bristles of end joint broken);
e, rod-shaped organ. Conchoecia valdiviae, complete specimen: f, out-
line; g, endopodite left 2nd antenna, medial view; h, rod-shaped organ,
ventral margin to right. Conchoecia macrocheira, complete specimen: i,
outline; j, endopodite (all bristles broken on end joint); k, right Ist an-
tenna, lateral view (4 bristles of end joint broken); 1, rod-shaped organ.
(Same scale in microns: a-c, f, i; d, g, h, j, k; e, 1.)
and 2 short spinous terminal bristles on lateral surface near ventral
margin; end joint (fig. 8b) with 1 long claw with minute spines distally
along concave margin, and total of 4 additional bristles and bristle-like
claws; clusters of spines present on surface of Ist and 2nd joints of
endopodite.
Maxilla (fig. 8e-h): Precoxale and coxale with anterior fringe of
hairs; coxale with long hirsute anterior bristle; 1st endite with 3 short
stout pectinate spines, 1 long stout spinous bristle, and 2 short spinous
Gulf of Mexico Ostracods 469
proximal bristles; 2nd endite with 2 stout pectinate spines and 2 slender
spinous bristles; 3rd endite with 1 proximal and 3 distal spinous bristles;
basale with 3 spinous bristles along distal margin; exopodite consisting
of 3 spinous bristles, 1 short and 2 long. Endopodite: Ist joint with 1
alpha-bristle and 1 beta-bristle, both with spines; end-joint with 2 spin-
ous a-bristles, 2 spinous b-bristles, and 1 stout c-bristle with short spines
distally along the convex margin.
Fifth limb (figs 8i-1; 9a): 1st joint with proximal tooth consisting
of 1 large and 3 small pectinate teeth and bearing 4 short bristles, and
distal tooth having large tooth followed by 2 smaller pectinate teeth
and 1 spine along the inner margin. 2nd joint: inner margin with large
fang like tooth followed by 1 small bifid tooth and 2 quadrifid teeth;
posterior surface with 3 spinous bristles, 2 near middle, 1 long and 1
short, and 1 long bristle proximal to them. 3rd joint with 2 spinous
bristles; 4th joint with 4 spinous bristles, 3 long and 1 short; endites
represented by 1 short spinous bristle; epipodial appendage with about
40 hirsute bristles.
Sixth limb (fig. 9b,c): Ist endite with 3 short spinous bristles; 2nd
endite with 1-2 spinous bristles; 3rd endite with 5-6 spinous bristles;
Ath endite with 4 spinous bristles; end joint with 9-10 spinous bristles;
surfaces of appendage hirsute; 1 spinous bristle present in place of epi-
podial appendage (observed only on 1 appendage, but may have been
lost from other during dissection ).
Seventh limb (fig. 9d-f): 6 bristles in distal group, 3 on each side;
2 bristles in proximal group, 1 on each side; all bristles with 2-5 bells
and marginal spines; terminus consisting of comb with large tooth with
shorter spinous tooth on each side opposing comb with large spinous
tooth with shorter tooth on each side, latter teeth each with frond of
spines at base.
Furca (fig. 9g): Each lamella with 9 claws, each separated from la-
mella by suture; 3rd claw smaller than 4th; claws 1 and 2 with spines
in row on lateral and medial sides of concave margins and slender hairs
along convex margins; each lamella with spines along dorsal margin and
hairs along ventral margin following claw 7.
Rod-shaped organ (fig. 9h): Segmented proximally and with short
spines in vicinity of tip.
Genitalia (fig. 91): Paired ovals, each in sclerotized sheath.
Eyes (fig. 9h): Medial eye pigmented, lateral eyes absent.
Brush-shaped organ (fig. 9j): 8 minute bristles, some with marginal
spines. (These small bristles were observed on integument dissected
from the vicinity of the genitalia).
Gut content: The gut contained numerous minute spines blunt at
one end acute at other. These are probable polychaete spines (Meredith
L. Jones, personal communication ).
Egg: Single egg containing well developed embryo present in brood
chamber.
470 Proceedings of the Biological Society of Washington
Comparisons: P. poulseni differs from P. inflatus in having 2 instead
of 3 small bristles on the endopodite of the 2nd antenna. The endopodite
of the 2nd antenna of P. poulseni has a small 2nd joint which seems to
be absent on P. inflatus. The 1st endite of the right maxilla of P. poul-
seni has 2 small closely spaced bristles; on P. inflatus these bristles are
separated by a wide space. The 4th joint of the Ist antenna of P. poul-
seni bears only 1 ventral bristle; P. inflatus has 2 in this position. The
furca of P. poulseni bears 9 claws compared to 10 on P. inflatus. Dif-
ferences also occur in the number of bristles on the 6th limbs.
SUBORDER HALOCYPRIDINA SKOGSBERG, 1920
SUPERFAMILY HALOCYPRIDACEA Dana, 1852
Family Halocyprididae Dana, 1852
Conchoecia atlantica (Lubbock, 1856)
Plate 2f, Figures 10a-e
Material: 29° 9 USNM 122079, complete specimen in alcohol, length
3.86 mm, height 1.88 mm; USNM 122080, specimen on slides and in
alcohol, length 3.52 mm, height 1.79 mm.
Conchoecia valdiviae Miiller, 1906
Figures 10f-h
Material: 29 9 USNM 122082, length 5.7 mm, height 3.1 mm;
USNM 122083, length about same (shell distorted); both specimens
preserved in alcohol.
Conchoecia macrocheira Miller, 1906
Figures 10i-]
Material: 19 USNM 122081, length 3.37 mm, preserved in alcohol.
LITERATURE CITED
ANONyMous. 1883. Sondages de l’aviso le Travailleur dans le golfe
de Gascogne. Annales Hydrographiques, ser. 2, vol. 5, pp.
4-35.
Baker, J. R. 1965. [MS]. Recent marine ostracod assemblages of
Puerto Rico. Thesis submitted to the Graduate School of
Texas Christian University. vi pp. 76, pl. I-XI, figs. 1-6.
Boip, W. A. vAN DEN. 1967. Ostracoda from Colon Harbor, Panama.
Caribbean Journal of Science, vol. 6, no. 2, March-June
1966, pp. 43-64, 4 pls. 1 fig.
Brapy, G. S. 1884-87. Les Crustacés-Ostracodes des expeditions du
Travailleur et du Talisman de 1881 a 1883. Les Fonds de
la Mer, vol. 4, pt. 3, pp. 164-175.
1887. Les ostracodes nouveaux des explorations du Tra-
vailleur et du Talisman. Les Fonds de la Mer, vol. 4, pt. 3,
pp. 194-200, pls. 12, 14, 15, [pl. 12 may not have been
published].
Gulf of Mexico Ostracods 471
Brapy, G. S., AND NorMAN, A. M. 1896. A monograph of the marine
and freshwater Ostracoda of the North Atlantic and of north-
western Europe, 2-4: Myodocopa, Cladocopa, and Platy-
copa. Trans. Roy. Dublin Soc., ser. 2, vol. 5, pp. 621-746,
pls. 50-68.
Kornicker, L. S. 1967. The myodocopid ostracod families Philo-
medidae and Pseudophilomedidae (New Family). Proc. U. S.
Nat. Mus. vol. 121, no. 3580, 35 pp. 12 figs., 1 pl.
Luspock, J. 1856. On some Entomostraca collected by Dr. Sutherland
in the Atlantic Ocean. Trans. Entomological Soc. Lond., New
Ser., vol. 14, pp. 8-42, pls. II-XII.
MUutier, G. W. 1894. Die Ostracoden des Golfes von Neapel und
der angrenzended Meeres-Abschnitte. Monogr. 21 in Fauna
und Flora des Golfes von Neapel, viii + 404 pp., 40 pls.
1906. Ostracoda in: Wissenschaftliche Ergebnisse der deut-
schen Tiefseeexpedition, vol. 8, pt. 2, pp. 29-154, pls. 5-35.
1912. Ostracoda in Das Tierreich. Preuss. Akad. Wiss. zu
Berlin, vol. 31, xxxiii + 434 pp., 92 figs.
Pokorny, Viapimir. 1953. A contribution to the taxonomy of the Pale-
ozoic Ostracods. Sbornik Ustredniho Ustavu Geologického,
Svazek XX, oddil Paleontologicky, pp. 213-232.
PouusEen, E. M. 1962. Ostracoda-Myodocopa, 1: Cypridiniformes-Cyp-
ridinidae. Copenhagen, Carlsberg Foundation, Dana Report
no. 57, 414 pp. 181 figs.
Sars, G. O. 1866. Oversigt AF Norges Marine Ostracoder. Norske
Vidensk.-Akad. forhandlinger (1865) 1866, pp. 1-130.
SxocsBERG, T. 1920. Studies on marine ostracods, 1: Cypridinids,
halocyprids and polycopids, Zool. Bidr. Uppsala, suppl., vol.
1, 784 pp., 153 figs.
SmirH, SANDERSON. 1888. Lists of dredging stations in North Amer-
ican waters from 1867 to 1887, (Extracted from the Annual
Report of the Commissioner of Fish and Fisheries for 1886),
pp. 871-1017, Washington: Government Printing Office.
SYLVESTER-BRADLEY, P. S. 1961. Ostracoda (part). Pt. 3 in Arthro-
poda of Part Q in Moore, Treatise on invertebrate paleontol-
ogy, xxiii + 442 pp., 334 figs.
472 Proceedings of the Biological Society of Washington
Vol. 81, pp. 473-478 30 December 1968
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
A NEW CLINGFISH, TOMICODON RHABDOTUS
FAMILY GOBIESOCIDAE, FROM THE
LESSER ANTILLES*
By WituiAM F. SmIrH-VANIZ
Tropical Atlantic Biological Laboratory, Bureau of
Commercial Fisheries, Miami, Florida
Collections of shore fishes by Victor G. Springer, obtained
under the auspices of the Breden-Archbold-Smithsonian Bio-
logical Survey of Dominica, contained several undescribed
species, among which is a distinctive new Tomicodon. The
new species brings to 11 the total known in the genus and
represents a second species from the Atlantic Ocean.
The following description is based on the uniform plan and
sequence previously adopted for this order of fishes (Briggs,
1955). Methods and terminology also follow those described
in that work. Measurements included in the description were
taken by measuring individual body parts and arithmetically
dividing the results into the standard length.
For the privilege of examing specimens in their care I thank
Drs. Warren C. Freihofer, Stanford University, C. Richard
Robins, Institute of Marine Sciences, University of Miami, and
Ernest A. Lachner and Victor G. Springer, United States Na-
tional Museum (USNM).
Tomicodon rhabdotus new species
Fig. 1
Holotype: USNM 201804, 38.4 mm standard length (SL), collected
just north of Bibay River, Bout Sable Bay, Dominica, bottom black sand
and few rocks, 0-2% ft., 13 November 1964, by V. G. Springer, R. H.
Reckeweg, and P. Spangler (VGS 64-26).
1 Contribution No. 74, Tropical Atlantic Biological Laboratory, Bureau of Com-
mercial Fisheries, Miami, Florida 33149 and Contribution No. 971, Institute of
Marine Sciences, University of Miami, Miami Florida, 33149.
46—Proc. Biot. Soc. WasH., Vou. 81, 1968 (473)
474 Proceedings of the Biological Society of Washington
w
Fic. 1. Tomicodon rhabdotus. USNM 199561; paratype; 18.7 mm
SL. (see text for data). A. Dorsal view. B. Lateral view.
475
A New Clingfish
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476 Proceedings of the Biological Society of Washington
TABLE 2. Measurements of the holotype and two paratypes of Tomicodon
rhabdotus (in thousandths of standard length).
Holotype Paratype Paratype
USNM 201804 USNM 199561 USNM 199560
Standard length (mm.) 38.4 18.7 15.9
Greatest body depth 126 126 107
Caudal peduncle depth 073 072 072
Caudal peduncle length 095 091 084
Head length 355 396 404
Head width 302 333 333
Snout length 094 086 075
Eye diameter 057 083 087
Bony interorbital 079 076 073
Dorsal-caudal length 284 259 248
Pectoral fin length 133 144 155
Postdorsal-caudal length 090 083 082
Disc length 293 297 309
Paratypes: USNM 199560, one specimen 15.9 mm SL, just north of
Rav Anse Cola River and a little south of Colihaut, Dominica, bottom
algal-covered rocks and barren boulders, 0-20 ft., 31 October 1964, V.
G. Springer and R. H. Reckeweg (VGS 64-14). USNM 199561, one
specimen 18.7 mm SL, Grand Bay, just south of Stowe, Dominica, bottom
rocks and boulders, 0-15 ft., 2 November 1964, V. G. Springer and R.
H. Reckeweg (VGS 64-16).
Diagnosis: A medium-sized Tomicodon with 5 pairs of trifid incisors
in upper jaw and 4 pairs in lower jaw. Anus slightly closer to posterior
margin of disc than to anal fin origin. Dermal flap extending from
margin of anterior nostril comparatively large, simple, and broad at distal
end; its height equal to or greater than distance between anterior and
posterior nostrils. Distinctive pattern of narrow, pale, vertical stripes on
posterior half of sides of body. Dorsal rays 9, anal rays 6, pectoral rays
18 (17 or 18), and caudal rays 8. Vertebrae 14 + 14 or 15 = 28 or 29,
including hypural.
Description: Body moderately compressed, depth 7.9-9.3 in SL.
Caudal peduncle short, least depth 1.2-1.5 in length. Head depressed,
length 2.5-2.8 and width 3.0-3.3 in SL. Snout moderately shallow with
a rounded outline, 3.8-5.4 in head length. Teeth in front of upper jaw
consist of 5 pairs of trifid incisors in all specimens, followed on each
side by 2 (1 or 2) well-developed canines. Lower jaw with 4 pairs of
trifid incisors in all specimens, followed on each side by 3 (2 or 3) well-
developed canines.
Greatest diameter of eye 0.8-1.4 in bony interorbital width and 4.7—
6.2 in head. Subopercular spine moderately developed, but hidden by
A New Clingfish ATT
TABLE 3. Frequency distribution of vertebrae in three species of
Tomicodon.
Species 28 29 36 31 32 33 34 35 36 37 38 No. mean
T. rhabdotus 1 2 3. = =28.66
T. humeralis 11 5412 1 78 30.08
T. f. fasciatus 3 31 9 By Bk A CGP
1 Although several specimens of T. fasciatus exhibited unusually high vertebral
counts, they appeared to be otherwise typical.
heavy skin of opercle region. Upper attachment of gill membrane op-
posite 5th (5-6) pectoral ray.
Dorsal-caudal distance may extend forward as far as the posterior
margin of pectoral fin. Postdorsal-caudal distance 1.8-2.0 in dorsal
length. Disc length 3.2-3.4 in SL. Disc region A with 6-7 rows of flat-
tened papillae across its width; 9-10 rows across width of disc region B.
Coloration: No color observations were made at the time of collection.
Specimens examined shortly after being placed in isopropanol, exhibited
a rich olive-green ground color. The light diagnostic markings (fig. 1)
were pale yellow, perhaps as a result of fading. After several months in
alcohol, the ground color faded to brown. Configuration of the pale
body markings is nearly identical in all specimens; the markings were
more conspicuous, however, in the two small paratypes which were
noticeably darker than the holotype.
Etymology: From the Greek, rhabdotus, meaning striped.
Discussion: As indicated in Table 1, the new species is not closely
related to T. fasciatus (Peters), the only species of the genus known
previously from the Atlantic Ocean. Of the Eastern Pacific species of
Tomicodon, rhabdotus is probably most closely related to T. humeralis
(Gilbert), endemic to the Gulf of California. Although the two species
are easily differentiated, they have in common low vertebral and
pectoral-ray counts, and a high dorsal-ray count. Configuration of lateral
markings is similar in the two species. The major difference is that in
rhabdotus the lateral stripes are nearly vertical, whereas in humeralis
they are diagonal.
The apparent restricted distribution of Tomicodon rhabdotus and the
similarities it shares with T. humeralis suggest that it has only recently
invaded the Atlantic Ocean. In contrast, the wide distribution and the
occurrence of two forms, subspecifically distinct, indicate that T. fasci-
atus has long been established in the Atlantic.
LITERATURE CITED
Briccs, Joun C. 1955. A monograph of the clingfishes (order Xenop-
terygii). Stanford Ichthy. Bull. 6: 1-224, 114 figs., 15 maps.
478 Proceedings of the Biological Society of Washington
Vol. 81, pp. 479-484 30 December 1968
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
A NEW GENUS AND SPECIES OF MILLIPED
FROM ILLINOIS (POLYDESMIDA, EURYURIDAE)
By WiLiIAM A. SHEAR
Dept. of Biology, Concord College, Athens, West Virginia
Although the members of the Subfamily Euryurinae, Family
Euryuridae, are remarkably similar in non-sexual characters,
the structure of the male gonopods provides reliable and easily
defined criteria for the recognition of genera. In Singuliurus
Causey, 1955, the gonopods are simple, falcate, and the
acropodite is subequal to the prefemur in length (Causey,
1955). Euryurus C. L. Koch, 1847, also has the prefemur and
acropodite subequal, and differs from Singuliurus only in that
the gonopod retains a tibiotarsal remnant and bears a process
near the end of the prefemur. Auturus Chamberlin, 1952,
has the acropodite one-fourth or less the length of the pre-
femur. Illiniurus, the new genus proposed here, presents a
mixture of these characters, perhaps best summarized by the
following key, which can be used to separate the known North
American genera of Euryurinae:
la. Gonopod blunt in lateral view, acropodite broadly expanded
and flaring, usually widest part of gonopod
AMA NIELS Moca Mia thatynce esuiul UMMA nOs Dy Auturus Chamberlin, 1942.
1b. Gonopod with tip acute in lateral view, widest at joint between
ACFOPOGItE wand POTEET poeta MeN NE a aia ol 2
2a. Tibiotarsal remnant exceeding solenomerite
PN ET SN PRO ME CU RUN PANO Illiniurus n. gen.
2b. Tibiotarsal remnant not exceeding solenomerite, or absent .. 3.
3a. Tibiotarsal remnant present, exceeded by solenomerite __.
DI ANG Be TN VG NCU RNC URE Sh I Euryurus C. L. Koch, 1847.
3b. Tibiotarsal remnant absent —-- Singuliurus Causey, 1955.
Despite the similar appearance of the many species of
euryurine millipeds, a search of the literature reveals no
47—Proc. Biot. Soc. WasuH., Vou. 81, 1968 (479)
480 Proceedings of the Biological Society of Washington
adequate description of their structure. The fairly detailed
description of Illiniurus beattyi (new species) that follows will
serve to characterize most species of the subfamily.
I am indebted to Dr. Joseph Beatty, Carbondale, Illinois,
for sending the large collection of millipeds which contained
the specimens of the new genus. The type species is named in
his honor. I also thank Dr. Richard Hoffman, Radford, Vir-
ginia, who first recognized the genus as new.
FAMILY EURYURIDAE POCOCK
SUBFAMILY EURYURINAE HOFFMAN
Illiniurus new genus
Diagnosis: Non-sexual characters as described for type species, distinc-
tive in details of gonopod structure from other genera of subfamily.
Coxae of gonopods globose, articulated with each other by lightly
sclerotized membrane. Prefemur only slightly longer than acropodite,
line of contact between acropodite and prefemur distinct on ventral side,
seminal canal not curved in passing line. Prefemur without processes.
Acropodite apically acute, with mesal margin incurved, concave
mesodorsally, ending in three subequal, short processes, of which
the solenomerite is the iongest. Distinguished from Auturus by having
the prefemur and acropodite subequal in length, and the acropodite
acute; from Euryurus by the lack of a prefemoral process and having
the tibiotarsal remnant exceed the solenomerite; from Singuliurus by the
presence of a tibiotarsal remnant. Type species, Illiniurus beattyi new
species, by monotypy.
Illiniurus beattyi new species
Figures 1-4
Type Specimens: Male holotype and female paratype (Mus. Comp.
Zool.), from Clear Springs, Union Co., Illinois; J. M. Nelson, collector,
28 October 1966.
Description of Male Holotype:
Structure: Body of head and 20 segments, 24.5 mm long, 3.9-4.0 mm
wide. Head (Fig. 1) large, smooth, without setae on cranial portion,
Epicranial suture distinct on frons, becoming obsolete between antennal
sockets. Face smooth, vertical, setation as follows: interantennal 1-1,
subantennal 1-1, frontal 1-1, genal 2-2, clypeal 7-7, labral 6-6.
Antennal sockets rimmed, separated by slightly more than their diameter.
Antennae short, stout, reaching posterior border of third segment when
fully extended posteriad. First and seventh antennal segments globose,
others clavate. Two dorsal macrosetae on antennal segment one, three
dorsal, three ventral on segment two, first two segments otherwise
New Illinois Milliped 481
glabrous, or nearly so. Distal segments progressively more setiferous.
Sensory cones four, set loosely in socket.
Collum slightly larger and broader than second body segment, smooth,
margined anteriorly, distinctly depressed transversely about one-fourth
its length from anterior edge, anterior edge straight, posterior edge
procurved, vaguely emarginate near the sides, gradually curving posteriad
laterally. Sides flared and raised posteriorly, depressed and _ nearly
vertical anteriorly. Lateral edges distinctly margined.
Body segments glabrous, faintly tuberculate, evenly convex dorsally.
Metazonites greater in diameter than prozonites, stricture strongly im-
pressed dorsally. Paranota about one-fourth body width, strongly de-
pressed anteriorly on segments two to five, gradually becoming nearly
horizontal; paranota overlapping on segments 16-19, otherwise well-
separated. Scapulorae marginal anteriorly, becoming submarginal and
blending into submarginal swellings laterally. Prolateral tooth set off
by obtuse notch. Pore formula normal, pores opening laterally on
edges of paranota, peritremata obsolete, surrounded by swellings.
Epiproct large, as long as segment 19 dorsally, broadly spatulate,
somewhat margined dorsally, surface convex anteriorly, nearly flat pos-
teriorly. Periprocts planoconvex, with distinct, but not thickened, lips.
Dorsal setae only slightly closer to margin than ventral setae. Hypoproct
flat, semicircular, paramedian setose tubercles small, indistinct.
482, Proceedings of the Biological Society of Washington
Prozonites minutely hispidulose ventrally, interzonal furrow nearly
straight. Sterna becoming glabrous, subcoxal elevations pronounced.
Coxal sockets rimmed, equidistant in both leg pairs except on segment
18, where posterior coxae are closer together.
Sides of prozonites becoming smooth, stricture distinct laterally. Sides
of metazonites smooth, except for raised line extending from posterior
coxal socket to posterior edge of paranota. Tuberculate ridge above
coxal sockets on all segments. Stigmata immediately anterio-dorsal to
coxal sockets, subequal in size, anterior stigma oval, posterior subcircular.
Stigmatic margins raised, contiguous with coxal condyles (Fig. 2).
Coxae globose, subequal in length with prefemora, unspined. Prefemora
unspined, but both coxae and prefemora bearing ventrodistal macrosetae.
Femora clavate, twice length of prefemora, setose beneath. Postfemora
globose. Tibiae and tarsi subcylindrical, tarsi twice as long as tibiae,
tapering, flattened laterally, heavily setose beneath, slightly so above.
Pretarsi long, one-third length of tarsi, sclerotic.
Sternite of segment six depressed, concave, anterior sternites otherwise
unmodified. Socket of gonopodal segment large, oval, prozonite reduced
to narrow strip. Lateral edges of socket forming distinctly raised lobes.
Gonopods (Fig. 3) large, extending in situ to anterior leg pair of sixth
segment. Coxae of gonopods large, globose, joined by lightly sclerotized
membrane. Prefemur slightly longer than acropodite, setose, constricted
midway in its length, ~vithout processes. Line of contact between
prefemur and acropodite distinct only on ventral side. Acropodite
more heavily sclerotic than prefemur, glabrous, curved transversely with
concave side mesodorsad, ending in three processes. Longest, most
lateral process probably represents tibiotarsal remnant; seminal groove
ending on middle (solenomerite) division, which bears third process, a
short pointed lamina, mesally.
Coloration: Coloration in life unknown, but probably agreeing
closely with other genera of subfamily: olive black with red or orange
paranota and red or orange spot in midline of each segment (based on
Virginia specimens of Euryurus leachii). After preservation in alcohol,
dorsum light brown, paranota and median spot white to clear testaceous,
dark brown line running diagonally from light median spot to posterior
origins of paranota. Epiproct unmarked, light brown. Collum light
brown, dark brown submarginal spots on sides.
Description of Female Paratype:
26.5 mm long, 4.0 mm wide. Structure very close to that of male, but
segments more arched dorsally, body of generally more robust appear-
ance. Anterior surface of prozonal sternite of segment three rugose,
cyphopods as in Fig. 4, valves flattened, oval, with single, median
receptacle.
Distribution: Known only from the type locality, Clear Springs, Union
Co., Illinois. The vial which contained the types also contained a single
New Illinois Milliped 483
male of Auturus evides (Bollman). The type specimens have been
deposited in the American Museum of Natural History, New York.
LITERATURE CITED
Causey, N. B. 1955. New records and descriptions of polydesmoid
millipeds (Order Polydesmida) from the eastern United
States. Proc. Biol. Soc. Washington 68: 21-30.
484 Proceedings of the Biological Society of Washington ] |
M
q
P|
Vol. 81, pp. 485—490 30 December 1968
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
PENTLAND’S TINAMOU IN ARGENTINA
(AVES: TINAMIDAE)
By RicHarp C. BANKs AND WAyNE H. Bou.
Bureau of Sport Fisheries and Wildlife; U.S. National Museum,
Washington, D.C., and Foreign Game Investigation Program,
Utah State University, Logan, Utah
Field work in Argentina in 1965 and 1966 by personnel of the
Foreign Game Investigation Program, of the Bureau of Sport
Fisheries and Wildlife, had as one of its results the accumula-
tion of a collection of nearly 300 specimens of tinamous,
which have been deposited in the U.S. National Museum.
Among these was a series of 15 Pentland’s Tinamou, Notho-
procta pentlandii (Gray), from the provinces of Mendoza,
San Luis, and Cordoba, at or near the southern end of the
range of the species (Hellmayr and Conover, 1942; Olrog,
1963). Comparison of these southern specimens with others
from the more northerly provinces of Catamarca, Tucuman,
and Salta and from Bolivia revealed that two distinct southern
populations were represented—a previously undescribed one
in Mendoza and one in the Sierras de San Luis and Cordoba.
There are thus three subspecies in Argentina, as follows.
Nothoprocta pentlandii pentlandii (Gray)
Range: From the Bolivian departments of Chuquisaca, Santa Cruz,
Cochabamba, and La Paz south along the Andean foothills of western
Argentina to southern Catamarca.
Comments: Comparison of this subspecies with others is made below.
This is the only subspecies currently listed as occurring in Argentina by
Olrog (1963), who did not recognize the next form. The type locality is
in the Bolivian Andes.
Nothoprocta pentlandii doeringi Cabanis
Range: Sierras de San Luis and Cordoba, central Argentina.
Diagnosis: Similar to the northerly nominate race in size (Table 1)
48—Proc. Biou. Soc. Wasu., Vou. 81, 1968 (485)
486 Proceedings of the Biological Society of Washington
but grayer in the general tone of the dorsum, resulting from gray rather
than brown feather edgings; more heavily spotted on the upper breast,
and with a less extensive pale area centrally on the abdomen.
Comments: Nothoprocta doeringi was named by Cabanis (1878) on
the basis of a specimen sent to him from Sierra Chica, near Cordoba.
Sclater (1879) noted that this population was closely allied to N.
pentlandii, and Sclater and Hudson (1889) considered the name
doeringi synonymous with pentlandii. The name applied by Cabanis
to the population of the Sierra Cordoba has seidom been used since
that time. Examination of the series of four fully grown birds available
to us from that mountain range reveals that that population is distinct
from the nominate form, as indicated above. Further, five specimens
from the Sierra de San Luis, in San Luis Province, are not separable
from the Cordoba birds. The name bestowed by Cabanis is available
and must apply to these populations, which are apparently isolated from
other populations by lowlands containing unsuitable habitat.
One specimen of N. p. pentlandii from near La Merced, Catamarca,
which of those examined is the nearest geographically to the Cordoba
population, shows an approach to N. p. doeringi in the grayness of the
dorsal coloration.
As far as we are aware, the occurrence of this species of tinamou in.
the Province of San Luis has not previously been reported.
Nothoprocta pentlandii mendozae new subspecies
Holotype: USNM 530855; female, Quebrada de Canota, Mendoza
Province, Argentina; collected 7 May, 1966, by Wayne H. Bohl, original
number 100.
Diagnosis: Differs from N. p. pentlandii and N. p. doeringi by having
a much longer bill (Table 1) and additionally from pentlandii by a
slightly longer wing, on the average. The general tone of the dorsal
coloration is gray, rather than brown, this appearance resulting from
the color of the lateral edges of the back feathers. In this respect,
mendozae is similar to doeringi but quite distinct from pentlandii. The
breast is more heavily spotted than in pentlandii and the abdomen is
darker than in either pentlandii or doeringi, being suffused with pale
gray or grayish-brown rather than buff.
Range: Known definitely only from the northwestern corner of the
Province of Mendoza, Argentina. Other reports of N. pentlandii from
Mendoza (Sanzin, 1917; Reed, 1919) and from northern Neuquen
(Olrog, 1963) probably also apply to birds of this subspecies.
Comments: Bill size is the most distinctive character of the subspecies
N. p. mendozae. The bill averages approximately 15 per cent longer
in members of this population than in either pentlandii or doeringi
(Table 1), and there is no overlap in the samples available to us.
Wing length is slightly greater in mendozae than in pentlandii, but
Pentlands Tinamou 487
TABLE 1. Measurements (mm.) and weight (gm.) of subspecies of
Nothoprocta pentlandii in Argentina.
pentlandii doeringi mendozae
N 10 9 6
Wing Range 120.2-142.8 123.0-143.2 134.5-145.7
Mean 133.6 135.4 139.3
N 9 9 5
Bill Range 21.0—24.0 21.0-25.1 26.3-27.6
Mean 22.6 23.3 26.9
Tarsus plus N 10 9 6
Middle Toe Range 60.2-67.8 60.2-68.4 65.2-68.6
Mean 65.0 65.1 66.9
N 2 10 14
Weight Range 955-305 211-330 301-384
Mean 280.0 265.7 339.9
doeringi is intermediate in this regard. Similarly, the length of the tarsus
and middle toe combined is greater, on the average, in mendozae than
in the other subspecies. The general impression of greater size in
mendozae is borne out by a consideration of weight. Fourteen specimens
of mendozae averaged 340 gm., whereas ten doeringi averaged only 266
gm. In both instances, equal numbers of each sex were considered; not
all birds weighed were saved as specimens. Two examples of pentlandii,
both males, weighed 255 and 305 gm.
DIscussION
Pentland’s Tinamou is a bird of moderate to high elevations; all speci-
mens taken in the course of field work by FGIP personnel were at
elevations of 3000 feet or above. Thus the low country in the extreme
northwest portion of Cordoba and the extreme southeast part of
Catamarca, containing the Salinas Grandes, would be an effective barrier
providing for the isolation of N. p. doeringi from N. p. pentlandii.
Similarly low-lying country to the west of the Sierra de San Luis, either
to the east or west of the Alto Pencosco, separates doeringi from
mendozae. The Rio Bermejo and its tributaries in northern San Juan
Province represent a possible barrier between mendozae and pentlandii;
the country is considerably more arid to the south and west of the rios
Troya, Bermejo, and Desaguadero than to the north and east.
Olrog (1963) has divided Argentina into a number of ornithological
zones. The distribution of the subspecies of Pentland’s Tinamou out-
lined in the previous paragraph fits well the zonation mapped by Olrog.
488 Proceedings of the Biological Society of Washington
Nothoprocta p. pentlandii is distributed roughly in the “Subzona Sub-
tropical Occidental” (p. 14), N. p. doeringi in the “Subzona Chaquena
Occidental” and N. p. mendozae in the “Subzona Chacoandina” (p. 26).
The three subspecies discussed above were also compared with a pop-
ulation from well north of the range of N. p. pentlandii, called N. p.
oustaleti Berlepsch and Stolzmann by Hellmayr and Conover (1942).
The sample of this form examined averages considerably larger than
any of the southern races of N. pentlandii in length of wing and of
tarsus and middle toe. Differences in color and in feather pattern are
so great that it is difficult to consider oustaleti conspecific with the forms
of pentlandii seen. Indeed, oustaleti was considered to be a distinct
species by Peters (1931).
Special comment should be made on the methods of measurement
used in this study. The scutellation of the tarsus and middle toe is
extremely variable in the vicinity of the juncture of these two elements.
This variability made it difficult, if not impossible, to obtain measure-
ments of either segment that would be comparable from one individual
to another. However, a single scute could be used as the boundary
point between tarsus and toe on a single specimen, and adding the
measurements for these two segments yields a figure more nearly com-
parable between populations than either measurement alone. The bill -
chord was measured from the tip to the point of the “V” formed by
the cere. Sexual dimorphism in size is slight (a maximum of 3 per cent)
with a population, and measurements for both sexes are combined be-
cause of the small number of specimens available.
We wish to acknowledge gratefully the assistance in the field of Srs.
Luis Arturo, Ernesto Roco, and Peter Miles. Dr. Gardiner Bump
provided assistance in many ways.
Specimens Examined
N. p. pentlandii. (9)—Bolivia: Cochabamba: Vacas, 2752 and 3800
m. (3); Arani, 2752 m. (1); Pocona, 2700 m. (1). Argentina: Salta:
25 mi. W Salta, 5500 ft., (1); Tucuman: San Pedro de Colalao (2);
Catamarca: near La Merced, 4000 ft. (1).
N. p. doeringi. (10)—Argentina: Cordoba: La Falda, 3000-3200 ft.,
(3); La Cumbre (1 plus 6 chicks); San Luis: vic. El Volcan, 3000-
3400 ft. (4); 7 mi. SW San Francisco, between 2400 and 2800 ft. (1).
N. p. mendozae. (6)—Argentina: Mendoza: Quebrada de Canota,
4800-6200 ft. (5); 2 mi. SW Puesto Hoyada, 4500 ft. (1).
N. p. ? oustaleti. (6)—Peru: Lima: San Mateo (2); Huinco (1);
Apurimac: Andahuaylas Hda., La Laguna, 3040 m. (1); Ecuador: Loja:
Malacatos, 5000 ft. (2).
Bolivian specimens of pentlandii and all oustaleti were borrowed
from the Chicago Museum of Natural History through the courtesy of
Emmet R. Blake. All other specimens are in the U. S. National Museum.
Pentland’s Tinamou 489
LITERATURE CITED
Capanis, J. 1878. Ueber eine Sammlung von Végeln der Argentinis-
chen Republik. Jour. f. Ornith., 26: 194-199.
HeLuimayr, C. E., AnD B. Conover. 1942. Catalogue of Birds of the
Americas and the Adjacent Islands. Part I, No. 1, Field
Mus. Nat. Hist., Zool. Ser., Vol. 13, pt. I, no. 1.
Oxtroc, C. C. 1963. Lista y Distribucion de las Aves Argentinas.
Opera Lilloana IX, Instituto Miguel Lillo, Univ. Nac. de
Tucuman. 377 p.
Peters, J. L. 1931. Check-list of Birds of the World. Vol. 1. Harvard
Univ. Press, Cambridge. 345 pp.
REED, C. S. 1919. Breves notas acera de nidos y huevos de algunas
aves de la cordillera de Mendoza. Hornero, 1: 267-273.
SANZIN, R. 1917. Lista de aves Mendocinas. Hornero, 1: 147-152.
SciaTer, P. L. 1879. [On a collection of birds.] Proc. Zool. Soc.
London for 1879: 460-461.
ScLATER, P. L., anp W. H. Hupson. 1889. Argentine Ornithology.
Vol. II. London. 251 pp.
490 Proceedings of the Biological Society of Washington
SM
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Vol. 81, pp. 491-498 30 December 1968
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
TAXONOMIC STATUS OF THE VESPERTILIONID
GENUS ANAMYGDON (MAMMALIA; CHIROPTERA)
By Carteton J. PHitiips AND ELMER C. BIRNEY
Museum of Natural History,
The University of Kansas, Lawrence
Miller (1907) named the vespertilionid subfamily Kerivou-
linae and included in it the genera Kerivoula Gray and
Phoniscus Miller. Chrysopteron Jentink was added to the
subfamily (Jentink, 1910), but later was considered a subgenus
of Myotis Kaup and therefore a vespertilionine by Tate (1941la,
1941b). Anamygdon, a monotypic genus known from a single
specimen, was named as a kerivouline by Troughton (1929).
Hill (1965) recently reviewed Kerivoula and Phoniscus but
did not judge the taxonomic status of Anamygdon. Ryan
(1965), who also studied Kerivoula and Phoniscus, considered
Anamygdon a distinct genus on the basis of descriptions by
others.
The subfamily Kerivoulinae is distinguished from the
Vespertilioninae primarily by means of sternal characteristics.
In the Kerivoulinae the length of the sternum is much less
than twice the breadth of the presternum. Furthermore, “only
four or five ribs articulate with the sternum” in kerivoulines
(Miller, 1907: 232), whereas in vespertilionines the sternum is
slender, “considerably more than twice [the] greatest width of
[the] presternum,” and “six ribs [are] connected with [the]
sternum’ (Miller, op. cit.: 197).
In addition to the sternal features listed above, Kerivoula
and Phoniscus are characterized by having three well-de-
veloped upper premolars in line with the axis of the toothrow.
Chrysopteron differs in having a reduced middle upper pre-
molar that is located slightly internal to the toothrow, and lacks
the sternal characteristics of kerivoulines.
49—Proc. Biot. Soc. WasH., Vou. 81, 1968 (491)
492. Proceedings of the Biological Society of Washington
Anamygdon was named on the basis of one specimen from
Roviana Island in the Solomons. Although this specimen had
external features and dentition generally like those of Myotis,
only five ribs were attached to the sternum (Troughton, 1929:
85). After careful study of his specimen, Troughton (op.
cit.: 86) decided that its “premolars definitely ally it with
Chrysopteron.” On the basis of sternal and dental character-
istics he concluded (op. cit.: 98) that Anamygdon represented
a kerivouline branch “nearest to the Vespertilioninae.” Further-
more, Troughton (op. cit.: 85) pointed out that because
Anamygdon agreed well with published descriptions of Myotis
moluccarum |= M. adversus moluccarum], it was possible that
some specimens assigned to M. moluccarum would prove to be
representatives of Anamygdon. This seemed especially likely
to Troughton because the sternal characteristics of Anamygdon,
which apparently allied it with the Kerivoulinae, could easily
have been overlooked by other workers.
Thomas (1923: 253) failed “to see any reason for distinguish-
ing [Chrysopteron] from Myotis.” Subsequently, Tate (1941a:
547; 1941b: 584) allocated Chrysopteron subgeneric rank
under Myotis, and thereby placed it in the Vespertilioninae.
Tate (1941b: 586) thought that Anamygdon, with its “Keri-
voula-like” sternum and “Myotis-like” premolars, represented
a lineage independently derived from near the common origin
of Myotis and Kerivoula.
In 1964, Philip Temple, who collected zoological specimens
in the Southwest Pacific for the Entomology Department of
the Bernice P. Bishop Museum, Honolulu, Hawaii, obtained
five specimens referable to A. solomonis in sea caves near
Toumoa on Fauro Island (6° 55’ S, 157° 07’ E) in the British
Solomon Islands Protectorate. Study of these specimens has
enabled us to re-evaluate the taxonomic status of this nominal
kerivouline genus.
Specimens listed herein are deposited in the Bernice P.
Bishop Museum (BPB-BSIP), the American Museum of Nat-
ural History (AMNH), the Australian Museum (AM), the
United States National Museum (USNM), and the British
Museum (BM). The Bishop Museum specimens originally
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494 Proceedings of the Biological Society of Washington
Fic. 1. Diagramatic representation of ventro-lateral aspect of the >
sterna of Anamygdon solomonis (BPB-BSIP 23757, upper) and Kerivoula
hardwickii depressa Milier (USNM 17909, lower) showing attachment
of ribs and clavicle.
were stored in alcohol; two later were prepared as museum
skins with skulls removed, and the cranium of a third was
extracted and cleaned so that morphological features could
be studied. All measurements given herein are in millimeters
and were taken by Phillips.
TAXONOMIC STATUS OF ANAMYGDON
We compared our five specimens of Anamygdon with the
careful, detailed description of the holotype of A. solomonis
given by Troughton (1929). Furthermore, Basil Marlow
compared one of our specimens directly with the holotype
in The Australian Museum. In all ways (dimensions, ex-
ternal features including the tragus, and cranial and dental
characteristics ) our specimens agreed with the holotype.
The sterna of Anamygdon and Kerivoula are compared in
Fig. 1. In Anamygdon the length of the sternum is not quite
twice as great as the breadth of the presternum, but in
Taxonomic Status of Anamygdon 495
Kerivoula the sternum is much less than twice the breadth of
the presternum. Number of ribs connected to the sternum can
be counted easily, but a standard system must be used. For
example, the first rib, which nearly is obscured by the clavicle,
might not be counted, and the last rib could be counted as
two because two ribs join together a few millimeters before
articulation. Direct comparison revealed that the sternum of
Anamygdon differs little, if at all, from sterna of numerous
species of Myotis. In Myotis, five ribs attach directly to the
body of the sternum. Clearly, Miller's (1907: 197, 232) com-
ments regarding sternal characteristics of the Kerivoulinae
and the Vespertilioninae would have been enhanced by an
illustration.
Comparative external and cranial measurements of speci-
mens of Anamygdon solomonis and Myotis adversus moluc-
carum are given in Table 1. Specimens of Anamygdon agree
well with a specimen (AMNH 99901) of Myotis adversus
moluccarum from Nissan in the Solomons and with character-
istics given by Thomas (1915: 170-171) for the holotype of
moluccarum. It is our conclusion, therefore, that Anamygdon
solomonis is synonymous with Myotis moluccarum, which is
currently regarded as a subspecies of M. adversus. Asian
species of Myotis, including adversus, are in need of taxonomic
review. The assignment of specimens herein discussed is as
nearly correct as is possible until such a review is undertaken.
A revised synonymy (based on literature judged most perti-
nent) of Myotis adversus moluccarum is as follows:
Myotis adversus moluccarum
Leuconoe moluccarum Thomas, Ann. Mag. Nat. Hist., ser. 8, 15: 170,
January 1915 (holotype from Ara, Kei Islands); Sanborn, Publ. Field
Mus. Nat. Hist., Zool. Ser., 18: 25, 12 February 1931.
[Myotis] moluccarum, Troughton, Rec. Australian Mus., 17: 85, 26
June 1929.
Anamygdon solomonis Troughton, Rec. Australian Mus., 17: 89, 26 June
1929 (holotype from Roviana, Solomon Islands); Laurie and Hill,
List of land mammals of New Guinea, Celebes and adjacent islands,
p. 77, 30 June 1954.
M[yotis]. moluccarum, Tate, Bull. Amer. Mus. Nat. Hist., 78: 551,
29 December 1941.
496 Proceedings of the Biological Society of Washington
Myotis moluccarum, Tate, Bull. Amer. Mus. Nat. Hist., 78: 590, 31
December 1941.
A[namygdon]. solomonis, Tate, Bull. Amer. Mus. Nat. Hist., 78: 590,
31 December 1941.
Myotis adversus moluccarum, Laurie and Hill, List of land mammals of
New Guinea, Celebes and adjacent islands, p. 69, 30 June 1954.
Anamygdon [solomonis], Ryan, J. Mamm., 46: 517, 25 August 1965.
ACKNOWLEDGMENTS
We are grateful to Hobart M. Van Deusen and Karl F. Koopman
of the American Museum of Natural History for arranging the loan of a
specimen of Myotis adversus and for allowing us to dissect the rib cage,
and to Basil Marlow and Ellis LeG. Troughton, The Australian Museum,
for comparing one of our specimens with the holotype of Anamygdon
solomonis, and for their helpful suggestions in discussions and cor-
respondence. J. Linsley Gressitt, the Bernice P. Bishop Museum, kindly
allowed us to study specimens collected by his expeditions, which were
supported by a contract (DA-MD-49-193-62-G65) with the United
States Army Medical Research and Development Command. A National
Science Foundation travel grant, to Phillips through the Committee on.
Systematics and Evolutionary Biology at The University of Kansas,
enabled comparative studies of specimens of Kerivoula in the U. S. Na-
tional Museum. J. Knox Jones, Jr., The University of Kansas Museum
of Natural History, reviewed the manuscript, and Charles O. Handley,
Jr. allowed us to study comparative material in the U. S. National
Museum.
LITERATURE CITED
Hint, J. E. 1965. Asiatic bats of the genera Kerivoula and Phoniscus
(Vespertilionidae ), with a note on Kerivoula aerosa Tomes.
Mammalia, 29: 524-556, December.
JenTinK, F. A. 1910. Chrysopteron bartelsii, novem genus et nova
species, from Java. Notes Leyden Mus., 32: 73-77.
Miter, G. S., Jr. 1907. The families and genera of bats. Bull. U. S.
Nat. Mus., 57: xvii + 1-282, 29 June.
Ryan, R. M. 1965. Taxonomic status of the vespertilionid genera
Kerivoula and Phoniscus. J. Mamm., 46: 517-518, 26
August.
Tate, G. H. H. 1941a. A review of the genus Myotis (Chiroptera )
of Eurasia, with special references to species occurring in the
East Indies. Bull. Amer. Mus. Nat. Hist., 78: 537-565, 29
December.
1941b. Notes on vespertilionid bats of the subfamilies
Miniopterinae, Murininae, Kerivoulinae, and Nyctophilinae.
Bull. Amer. Mus. Nat. Hist., 78: 567-597, 31 December.
Taxonomic Status of Anamygdon 497
Tuomas, O. 1915. Two species of Leuconoe. Ann. Mag. Nat. Hist.,
ser. 8, 15: 170-172, January.
1923. On some small mammals, chiefly bats, from the East
Indian Archipelago. Ann. Mag. Nat. Hist., ser. 9, 11: 250—
255, February.
TroucutTon, E. LeG. 1929. A new genus and species of bat (Keri-
voulinae) from the Solomons, with a review of the genera
of the sub-family. Rec. Australian Mus., 17: 85-99, 26
June.
498 Proceedings of the Biological Society of Washington
Vol. 81, pp. 499-510 30 December 1968
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
NEW NORTH AMERICAN SPIROBOLOID MILLIPEDS
OF THE FAMILIES ATOPETHOLIDAE AND
MESSICOBOLIDAE
By H. F. Loomis
U. S. Dept. of Agriculture, Miami, Florida
The publication of R. L. Hoffman and B. S. Orcutt’s
synopsis of the family Atopetholidae (1960), putting the dis-
connected genera in formal keys; giving comparative diagnoses
and notes on them, as well as redescribing several of the earlier
species in the light of their current knowledge, has greatly
facilitated work on this large group whose principal range
was then our southwestern tier of States and Mexico. Since
1960 only two species of established genera have been added
to the family and these do not materially affect the range or
generic concepts.
The species of the order Spirobolida described herein have
been accumulating for many years without a treatment having
been prepared in which to include them appropriately. It is
desirable that they be recognized now since they represent
three new genera and a total of seven new species. Several
more or less important characters affecting known genera and
species have been seen and are mentioned.
All holotype and paratype specimens have been deposited
in the myriapod collection of the U. S. National Museum.
FAMILY ATOPETHOLIDAE
SUBFAMILY ARINOLINAE
Uvalida new genus
Type-species: Uvaldia intersecta new species.
Diagnosis: Running to Arinolus Chamberlin in Hoffman and Orcutt’s
key (1960), but males differing especially from Arinolus in lacking
constricted segments behind segment 1 and having no enlargement at
segment 6 or 7. Males differ also in having telopodite of anterior
50—Proc. Biot. Soc. WasuH., Vou. 81, 1968 (499)
500 Proceedings of the Biological Society of Washington
gonopods hidden from in front by coxite; there is no distal projection for
seminal channel of posterior gonopods; the produced coxal lobes of legs
3-7 are broad, thin, and simple.
Description: Body large, its anterior end neither noticeably constricted
behind segment 1 nor enlarged at segments 6 and 7 in males.
Head with ocelli in rounded groups separated by less than three times
the diameter of one group. Antennae quite slender and barely exceeding
posterior margin of segment 1. ‘
Segment | strongly emarginate below eye; 1-3 deep striae in posterior
surface above lateral angle which much exceeds ventral surface of
segment 2. Except at extremities, segments have a transverse constric-
tion containing a slight furrow marking suture between mid- and
hindbelts; constriction becoming more obvious in its descent; hindbelt
convex with posterior margin thick, the middorsal suture lightly im-
pressed on most segments, lateral suture also impressed. Lateral in-
terstrial ridges not continued behind posterior margin. Pores small,
located in or behind the transverse suture at or very little below its
intersection with the usually straight, continuous lateral suture except
on segment 6 where pore is well below the discontinuous lateral one.
Two segments preceding the last one telescoped as far as pores. Last
segment of male broadly rounded at apex, in female definitely more
angular, surpassed by the inflated anal valves which meet in a very
broad, deep groove; prcanal scale broadly rounded at apex Claws of
legs 1-3 of both sexes decidedly larger than others.
Coxite of each anterior gonopod unusually short, the relative area
much less than in any other genus of the family; telopodite hidden be-
hind coxite. Sternum very broad above the basal yoke, its area greater
than in other genera. Outer joint of each posterior gonopod thin, simple,
and widening from base to apex; seminal tube apparently opening from
a short apical trough instead of a separately produced process. Coxae
of legs 3-7 produced into broad and thin lobes of which those of fifth
legs are broadest; those of seventh legs thickest.
Etymology: Generic name in reference to county where collected.
Uvaldia intersecta new species
Holotype: Male, USNM myriapod collection no. 3296.
Type-locality: Holotype, 2 other males, 1 lacking a moult of maturity,
female and several young, Uvalde, Uvalde Co., Texas, 20 November
1911, O. F. Cook.
Description: Holotype 47 mm long, 4.5 mm wide, with 45 segments;
other male with 43 segments; female 53 mm long, 6 mm wide, 45
segments.
Head with ocelli in rounded groups of 7-8 rows—3-5—6—7—6—4—2 in
holotype; 2—4-5-5-5-5-5-3 in female. Clypeal fovea 5-5. Antennal
joint 2 longest, subequal joints 4 and 5 next, joint 6 shortest, joint 1
decidedly thickest.
—_
Ul eS Shalit
ol es
New Millipedes 501
Segments 1 and 2 shown in Fig. 1. Body segments without spines at
end of ventral striae. Surface of hindbelt finely and sparsely punctate;
midbelt less punctate. Segment 7 of male not greatly modified behind
gonopods, merely being raised into a straight, low, transverse ridge or
swelling.
Gonopods shown in Figs. 2-4. Sternum rather heavily chitinized for a
considerable distance of median area above basal yoke, its apex also
obviously chitinized. Coxae of legs 3-5 shown in Fig. 5.
Etymology: Latin intersecta is in reference to the location of each
repugnatorial pore in or near the intersection of the interzonal suture
with the lateral sulcus.
Some doubt arose over including the female in this species as pores
of the anterior segments are slightly in front of the transverse suture
but farther back they are as found in males. Also last segment is quite
abruptly rounded behind, differing materially from males. These
characters may indicate another species or merely individual or sexual
variation within this one.
SUBFAMILY EURELINAE
Cenirelus fluvialis new species
Holotype: Male, USNM myriapod collection no. 3297.
Type-locality: Holotype and 2 females, Comstock, Val Verde Co.,
Texas, March 1911, W. P. Carr.
Diagnosis: Distinguished from C. kerrensis (Chamberlin) by the
more quadrate anterior gonopods, differences of the posterior ones, and
of the coxae of male legs 3-7.
Description: Number of segments 50-51, diameter 6.5-7 mm, the
specimens broken.
Frontal median sulcus of head much more impressed than that of
vertex; clypeal fovea 4-4 and 5-5 in females, 5-6 in male. Eyes sub-
triangular rather than rounded, ocelli of male 6-7—7—7-6—4-2, beginning
at edge of segment 1, the lower ocellus in first five rows twice the diam-
eter of the upper one. Antennal joints quite evenly decreasing in length
from second to sixth, the latter narrowest; only joints 1 and 2 glabrous.
Segment 1 much as in kerrensis but the lateral limit produced farther;
side above it smooth or with a few striations. On following segments
the suture between mid- and hindbelt quite strongly impressed on sides,
more lightly so across dorsum; hindbelt gradually elevated behind it to
the thickened posterior margin; dorsal median suture impressed across
hindbelt but less so across midbelt; lateral suture strongly impressed
across hindbelt. The oblique spines terminating the lateral ridges begin
on segment 17-20 and do not project on the two preanal telescoped
segments where the pores are barely visible in females but not in male;
hindbelts of both telescoped segments much shortened in both sexes.
Last segment evenly rounded at apex, distinctly exceeded by the
valves which are broadly convex and meet in a broad deep groove.
502 Proceedings of the Biological Society of Washington
Fics. 1-5. Uvaldia intersecta new species. 1, lower side of male
segments 1 and 2; 2, right anterior gonopod, anterior view; 3, same,
posterior view; 4, right inner gonopod with basal element considerably
foreshortened, anterior view; 5, basal joints of male legs 3-5, anterior
view. Fics. 6-8. Centrelus fluvialis new species. 6, left anterior gonopod,
anterior view; 7, same, posterior view; 8, right inner gonopod, anterior
view. Fics. 9-11. Comanchelus camporum new species. 9, anterior
gonopods, anterior view; 10, same, posterior view; 11, right inner
gonopod, anterior view. Fics. 12-13. Comanchelus lobatus new species.
12, right anterior gonopod, anterior view; 13, same, posterior view.
New Millipedes 503
Preanal scale with an indefinite submedian transverse depression in the
essentially flat surface; anterior half smooth, posterior portion variously
striate; apex very broadly rounded to almost emarginate.
Gonopods as shown in Figs. 6—8; coxites of anterior gonopods more
quadrangular than shown for kerrensis, their telopodites (posterior
plates) also apparently more quadrate. Ventral ridge of segment 7
transverse, its posterior median portion directed strongly ventrocaudad
and hiding the deeply emarginate segmental border.
Legs 1 and 2 of male with claws large and heavy; those of third legs
half as long, followed by very short conical ones, a fourth as long, on
legs 4-7. Coxal lobes of third legs fully chitinized, broadly spatulate,
directed somewhat as described by Hoffman and Orcutt (1960) for
kerrensis but apparently less caudally bent. Coxae of legs 4-6 about
half as high as on third legs, very thin, broad, increasing in width
caudad, and rounded-truncate at apex; coxae of seventh legs several
times as thick as preceding ones and at least twice as high, evenly
rounded at apex.
Etymology: Latin fluvialis associates the species with the nearby Rio
Grande.
Comanchelus camporum new species
Holotype: Male, USNM myriapod collection no. 3298.
Type-locality: Holotype, a slightly younger male, 3 females found
24 mi. E of Llano, Llano Co., Texas, 4 April 1964, J. C. Loomis.
Diagnosis: Several characters of this species associate it with Eurelus
Cook but others, particularly relating to the gonopods, more closely
approach Comanchelus, as exemplified by Hoffman and Orcutt’s geno-
type, hubrichti, and accordingly it is placed in this genus. The out-
standing characters which associate it with Eurelus, however,—the lobed
coxae of the anterior male legs and reduced claws of legs 4—7, are not
found in hubrichti. This mixture of generic characters may require with-
drawal of Comanchelus into the elder genus, especially since when
Comanchelus was erected its authors observed that its “two species which
although generally similar to Eurelus soleatus appear to diverge enough
to justify their recognition as a separate group.”
Description: Size varying from 40-43 mm long and about 4.5 mm
wide, with 45-46 segments; color in life probably slate gray to slightly
olivaceous.
Head with surface shining, very sparsely punctate; posterior half of
vertex finely sulcate medianly, anterior half smooth; front much more
coarsely and deeply sulcate; ocelli 40 or more, in a round patch;
clypeal fovea 44, 4-5, 4-6, hence possibly 6-6; emargination of clypeus
slightly broader than a right angle. Antennae not slender, capable of
reaching posterior margin of segment 1; length of joints as in hubrichti.
Segment 1 strongly emarginate below lower corner of eye and bounded
by a heavy rim; lateral angle acutely rounded, surface above it not
504 Proceedings of the Biological Society of Washington
striate but one or two very short rudimentary striae present in posterior
margin adjacent to lateral angle of some specimens; surface shining and
finely punctate. Outer pleural corner of segment 2 not produced beyond
limit of segment 1 and much less acute than in original illustration of
hubrichti.
Succeeding segments with forebelt finely striate; midbelt evenly and
finely short-aciculated lengthwise and with a few tiny punctations inter-
mixed on posterior half; hindbelt more shining and with more and larger
punctations but no aciculations; mid- and hindbelts separated by im-
pressed suture in a slight but evident constriction; all sutures between
segmental divisions obvious as light lines in dark bodywall; lateral striae
adjacent to legs ending in an angulation not produced beyond margin;
posterior margin of segments thick and rising abruptly from supplementary
margin. Last segment very broadly rounded at apex. Anal valves
strongly inflated their anterior halves rugulose-punctate, posterior por-
tions generally smooth and strongly shining. Preanal scale broadly
truncated at apex, sides somewhat rounded. Ventral setae on leg joints
]—]—2—2-2-8.
Gonopods as shown in Figs. 9-11.
Male legs 1 and 2 with long heavy claws, three fourths as long
on next pair and decreasing thereafter to half as long on seventh pair
where they are half as long as on remaining legs. Coxae of third legs
caudally produced as long, thin, narrow, subspatulate, distally deflexed
lobes slightly notched on outer side near middle, and separating the
lobes of both fourth and fifth coxae which are much as described by
Hoffman and Orcutt for E. soleatus. Seventh coxal lobes quite broad,
thick, and high, especially at mesal limit; lobes of sixth coxae broader,
thinner, and only about half as high. Third joint of anterior legs not
lobed at base; last joint of legs 4—7 imperceptibly flattened dorsally.
Etymology: Latin camporum is in reference to the type locality being
in the West Texas plains- Llanos (Spanish).
Comanchelus lobatus new species
Holotype: Male, USNM myriapod collection no. 3299.
Type-locality: Holotype, another male, 1910, W. P. Carr; 2 females,
April 1911, O. F. Cook; both collections, San Antonio, Texas.
Diagnosis: The posterior gonopods show relationship with C. hubrichti
from which it may be readily distinguished by the large coxal lobes
of the anterior male legs.
Description: Males 53 mm long, 6 mm wide, 45-46 segments; fe-
males 53-56 mm long, 7 mm wide, 44 segments.
Eyes with 34-36 ocelli in seven rows, counting from back of head;
ocellus nearest lower posterior corner larger than others. Antennae
with joint 2 scarcely longer than joint 3; joints 4 and 5 subequal and a
little shorter; joint 6 shorter and narrower than inner ones. Frontal
area deeply sulcate at middle and with two or three to six or eight
coarse transverse striations; vertigial sulcus faint; clypeal fovea 5-5.
New Millipedes 505
Segment 1 deeply and evenly emarginate in front from behind eye
to lateral angle; lateral surface with one or two to five or six short
striae. Anterior flange of segment 2 not reaching as low as angle of
segment 1. Following segments with transverse constriction from which
the faintly convex hindbelt is gradually elevated; midbelt finely punctate;
hindbelt finely but less punctate above, lower sides with 13 or 14~20
strong striations with intervals ending in rounded but not protruding
shoulders. Dorsal suture lightly impressed across hindbelt; second
transverse suture lightly impressed across dorsum, more strongly so on
sides. Three preterminal segments moderately telescoped in females
but more strongly so, and with pores hidden, in males.
Last segment broadly rounded behind and surpassed by anal valves.
Anal scale transverse to emarginate at apex, each side emarginate; surface
with submedian transverse depression behind which it is slightly
swollen and longitudinally striate.
Gonopods as shown in Figs. 12-14; back surface of outer joint of
posterior gonopod smooth and shining.
Ventral ridge of male segment 7 nearly flat, its posterior limit high
above the deeply emarginate posterior border of the segment; surface
strongly striate, the adjacent surface each side with coarse, more or less
lunate depressions.
Male legs 1 and 2 with claws long and quite slender; those of third
legs about half as long. Coxae of third legs with lobes broad and thick
at base, distal half thinned from in front and narrowed inwardly to an
erect acute apex rising close to that of opposite lobe. Lobes of coxae 4
and 5 thin, but broader, shorter, subquadrate, highest mesally. Lobes of
sixth coxae wider but a little shorter and thicker; those of seventh coxae
narrower and shorter than the sixth but considerably thicker, and
broadly rounded at apex.
Etymology: Latin lobatus is in reference to the large coxal lobes of
the anterior male legs.
Mannobolus new genus
Type-species: Mannobolus peninsularis new species.
Diagnosis: Most closely related to Eurelus but the apical portion of
the inner gonopods is thin, simple, decidedly boat-like, and somewhat
curved; unusually large basal portion also appears capable of some
movement; coxal apodemes absent and possibly broken from both
specimens. Claws of anterior male legs reduced in length after the
first two pairs but none rudimentary.
Description: Small, rather slender species. Sides of segment 1
sharply narrowed, curving mesad, and somewhat clasping lower portion
of mandibulary cardo; anterior marginal rim narrow. Segment 2 not
reaching lower angle of segment 1. Principal body segments strongly
constricted, the constriction lacking an impressed sulcus; pores in
midbelt; ridges formed by lateral striae of midbody segments continued
506 Proceedings of the Biological Society of Washington
Fic. 14. Comanchelus lobatus new species, left inner gonopod, an-
terior view. Fics. 15-18. Mannobolus peninsularis new species. 15,
first and part of second segment, lateral view; 16, anterior gonopods,
anterior view; 17, same, posterior view; 18, right inner gonopod, anterior
view. Fics. 19-20. Petenobolus antiquorum new species. 19, preanal
scale; 20, right gonopods, anterior view. Fics. 21-24. Petenobolus
mayanus new species. 21, right anterior gonopod, anterior view; 22, left
telopodite of anterior gonopod and inner gonopod, posterior view; 23,
right inner gonopod, anterior view; 24, coxa and next two joints of
third male leg, posterior view.
New Millipedes 507
as upraised spines behind margin. Two segments before the last strongly
telescoped.
Anterior gonopods with tips almost meeting above chitinized apex
of sternum which is separated from the basal yoke by a semi-membranous
area. Posterior gonopods as in diagnosis.
Coxal lobe of third male legs somewhat as in Eurelus, high, thin,
the tip bent caudad; lobes of next three legs about half as high, their
tips decreasingly bent back; lobes of seventh legs higher than on
preceding three pairs, vertical, and at least twice as thick.
Etymology: The generic name combines that of the collector of the
species with the suffix of Spirobolus, used widely in naming genera
within its order.
Mannobolus peninsularis new species
Holotype: Male, USNM myriapod collection no. 3300.
Type-locality: Holotype and another male, both broken, from
Comondu, Baja Califomia, February 1923, Wm. R. Mann.
Description: Largest and most badly broken male (holotype) ap-
parently with 46 segments, 3 mm wide; other male 2.8 mm wide, ap-
parently with 41 segments but one or two credited to holotype may
belong to this male.
Head with strong median sulcus on vertex widely separated from
similarly strong one on front. Clypeal fovea 4-4. Eyes separated by
about three times diameter of one eye; ocelli in rows 2-5-6—7—7-6-4,
beginning nearest vertigial sulcus, forming round group. Antennal joint
2 distinctly longest, joint 6 narrower and shorter than the subequal
intervening ones.
Segment 1 narrowly rimmed along anterior emargination and slenderly
angled below (Fig. 15); lower sides curving inward, somewhat clasping
lower part of mandibulary cardo. From segment 2 or 3 to those tele-
scoped at back end of body a pronounced constriction is evident from
base of legs across dorsum but is deepest just above pores which are
in its bottom on anterior segments but on its front slope farther back.
Lateral suture impressed behind pore across hindbelt; interbelt sutures
nowhere impressed. Beginning at about segment 14 or 15, and con-
tinuing to caudal segment 5 or 6, four or five ridges formed by the
lateral striae are continued beyond margin as obliquely raised spines.
Last segment very broadly and evenly rounded behind; considerably
exceeded by the broad anal valves which are somewhat flattened be-
hind and meet in a narrow, shallow goove. Anal scale broadly rounded
behind except for its nearly straight median portion.
Gonopods shown in Figs. 16-18. Legs 1 and 2 with long, quite slender
claws, claws gradually shortening on succeeding legs, only about half
as long on sixth legs, broken from seventh legs. Coxae of third legs
elevated into thin, basally constricted lobes narrowing distally, the
acute tips bent caudad; lobes of fourth and fifth coxae about half a
508 Proceedings of the Biological Society of Washington
high with less acute tips decreasingly bent caudad; lobes of sixth legs
a little higher, the rounded apex not bent; lobes of seventh legs higher
and at least twice as thick as those of legs 4-6, rounded and highest
mesally.
Etymology: Latin peninsularis alludes to the peninsula of Baja
California from which the species came.
FAMILY MESSICOBOLIDAE
Petenobolus new genus
Type-species: Petenobolus antiquorum new species.
Diagnosis: Related to Oxobolus Chamberlin but easily separated by
the acutely angled last segment and preanal scale, and by differences
of the gonopods, particularly the broad granular tips of the anterior
ones, and the setose tips of their telopodites.
Description: Body large and stout. Head with rounded eyes; outer
row of ocelli with one or two at least twice the diameter of ocelli of
inner row. Antennae short, crassate, the outer joints noticeably flattened;
sense cones 4, in a zigzag line. Clypeal setae 44 or 5-5, with an
additional one far in front on each side in the angle adjacent to labrum.
Labral setae 12 or 13-22. Gnathochilarium widest near apex; mentum .
swollen at apex, broadly concave below it.
Segment 1 narrowing Jaterally to an abruptly rounded angle; a heavy
rim from angle to lower corner of eye. Segment 2 extending far below
1, as in Oxobolus, and deeply excavated in anterior angle. Succeeding
segments with sulcus between mid- and hindbelt faintly impressed
across dorsum. Pore of segment 6 at level of those following. Surface
immediately below each ventral stria extended behind margin as an
acute spine, the spines reaching to level of pores on caudal segments.
Apex of last segment and of preanal scale subequally acute, the former
much exceeded by the anal valves which have prominent thickened
margins.
Anterior gonopods large; inner portion of each broad, moderately
extended, surface finely and evenly granulate; telopodite with a dense
cluster of setae on outer side of apical process. Inner gonopods with
outer joint somewhat spoon-like; seminal duct in a longitudinal median
ridge along the concave face, terminating in a projecting spur. Coxae
of legs 3-6 distally lobed; coxae of seventh legs not lobed or as broad
as those preceding. Second joint of some pregenital legs with a rounded
tubercle on ventral face adjacent to coxa.
Etymology: Generic name in reference to the Dept. of Peten, in
which the species were found, plus suffix of Spirobolus.
Petenobolus antiquorum new species
Holotype: Male, USNM myriapod collection no. 3301.
Type-locality: Holotype, Uaxactun, Dept. of Peten, Guatemala, 29
March 1922, O. F. Cook and H. F. Loomis.
wasieleial
can
New Millipedes 509
Description: Length 84 mm, width 11 mm, with 41 segments; surface
of body almost smooth and strongly shining.
Head with eyes more nearly round than triangular or oval; com-
posed of 25 ocelli in rows, 2-4-5-5—5—4, counting outwardly, the two
posterior ocelli in outer row largest. Antennae short and stout, a little
flattened; joints decreasing in length as follows- 2—3(1—4—5 subequal)-6-7;
sense cones in a zigzag row rather than a quadrangle. Frontal sulcus
moderately impressed. Clypeal setae five one side, none opposite, a
supplementary one in each angle adjacent to labrum; labral setae 7-6.
Mandibulary stipes and cardo strongly depressed under antennae.
Gnathochilarium approximately quadrate; mentum moderately concave,
its apex slightly swollen.
Sides of segment 1 narrowed, the lower angle sharply rounded, edge
above it emarginate to lower corner of eye. Segment 2 descending well
below segment 1 anteriorly, its front and lower margin much thickened,
the latter rounded behind without a definite posterior angle; inner sur-
face deeply depressed below and behind angle of segment 1. Last
segment ending in an acute angle exceeded by thickened margins of anal
valves. Anal scale shown in Fig. 19, its apex resembling that of last
segment.
Gonopods as shown in Fig. 20; inner one with apex truncated in
lateral view and with a distinct shoulder on outer side near base of
joint; seminal spur prominent. Second joint of legs 3 and 4 with a
round tubercle ventrally adjacent to coxa; pregenital legs otherwise
as in next species.
Etymology: Latin antiquorum refers to the fact that the species was
found among the ancient ruins of Uaxactun.
Petenobolus mayanus new species
Holotype: Male, USNM myriapod collection no. 3302.
Type-locality: Holotype, consisting of head and next 23 segments,
Uaxactun, Dept. of Peten, Guatemala, 29 March 1922, O. F. Cook
and H. F. Loomis.
Description: Comparisons throughout this description are with
antiquorum. Body 12 mm wide. Head with ocelli in rows—2-3-4-5-5+4,
the last row with two enlarged ocelli. Frontal sulcus more impressed
and extending farther upward, to between antennae which are more
flattened, as easily seen in end view of joints; joint 2 noticeably longer,
joints 3—5 with basal constriction longer and more obvious. Clypeal
setae 4-5; labral setae 9-10. Mandibulary stipes and cardo only
moderately depressed. Gnathochilarium obviously broader than long;
mentum more concave, apex more strongly swollen. Anterior rim of
segment 1 more sharply limited behind by a deeper furrow; front
margin straighter; lateral angle not as narrow. Segment 2 with depression
under segment 1 deeper; anterior angle a little more abruptly rounded;
ventral margin straighter and ending in a slight but noticeable angula-
510 Proceedings of the Biological Society of Washington
tion with the posterior margin, the thickened ventral margin flatter
and less sharply defined.
Gonopods shown in Figs. 21-23; inner gonopod with outer joint
more continuous with basal one, lacking a lateral shoulder, apex more
rounded, seminal spur smaller and less protruding. Coxae of third legs
produced into conic lobes; legs 4 and 5 with broader and less produced
lobes; legs 3-6 with a rounded tubercle on ventral face of joint 2
adjacent to coxa (Fig. 24).
Etymology: Mayanus is in allusion to the Mayan inhabitants of the
country where the species was collected.
LITERATURE CITED
HorrMan, R. L. ann B. S. Orcutr. 1960. A synopsis of the
Atopetholidae, a family of spiroboloid millipeds. Proc. U.S.
Nat. Mus., no. 3426, vol. 111, pp. 95-166, illus.
Vol. 81, pp. 511-538 30 December 1968
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
REVISION OF THE MILLIPED GENUS WAMOKIA
CHAMBERLIN FROM THE SIERRA NEVADA OF
CENTRAL CALIFORNIA (DIPLOPODA:
POLYDESMIDA: XYSTODESMIDAE)
By Joun S. Buckertr AND MicHAEL R. GARDNER
California Department of Agriculture Sacramento, California,
and University of California Davis, California
The genus Wamokia Chamberlin consists of a moderately
sized group of milliped species occupying a small region of
the central Sierran Foothills of California. They may be
recognized by the dull yellow ground color with a dark mid-
dorsal stripe and orange paranota.
Unlike most groups of western diplopods, Wamokia has been
almost totally neglected by students of the Diplopoda. De-
scribed by R. V. Chamberlin in 1941, the original diagnosis
distinguished the genus adequately, but left some doubt about
the identity of placera Chamberlin, the type species. An
illustration given of the gonopod from the ventral aspect is
of assistance for identification, but does not show enough of
the important femoral endite. However, a sketch of the type
prepared by Richard L. Hoffman and loaned to us has been
of great value in fixing the identity of placera. Chamberlin
and Hoffman (1958) misspelled the genus as Waimokia in their
checklist and cited the type locality as occurring in Tulare
County, though it is actually in E] Dorado County. Buckett
(1964) corrected the latter error although he, too, misspelled
the generic name.
This work was initiated through an agreement with Dr.
Hoffman, who had discovered two undescribed species of
Wamokia in some xystodesmid material we loaned him for
his revisionary work. Subsequently, further field work was
51—Proc. Bion. Soc. Wasu., Vou. 81, 1968 (Sila)
512 Proceedings of the Biological Society of Washington
carried out which revealed the existence of four other un-
described species thus making a total of seven entities in the
genus.
Under the variation section of each species, measurements
are given to help characterize populations. Whenever enough
material existed, five males and five females chosen at random
were used to compute the values. For the “average population
length” the total of the 10 specimens was used. For length/
width ratios, full width, including paranota, was used.
ECOLOGY
The genus Wamokia is composed of a group of species
which range from elevations of 500’ on the Cosumnes River
to 3200’ at Foresthill, Placer County. Yet the entire known
range of the genus does not exceed 50 miles in the greatest
dimension.
A number of aspects of the habitat are shared by all the.
species in the genus. Wamokia are almost always associated
with the bright ochre Aiken Clay (see Storie, 1927) soil which
dominates the Sierran Foothills of El Dorado, Placer and
Nevada Counties, specimens being found actually within the
soil or in the zone between soil and litter. It is quite possible
that the geographic boundaries of the genus are influenced
by the distribution of this soil type. A common denominator of
all collections is the occurrence of Quercus spp., usually
wislizenii A.DC. and kelloggii Newb. in the immediate vicinity
of the collecting spot. The immediate habitat of Wamokia
populations appears to be influenced strongly by vegetation
in connection with the soil type.
In general, Wamokia species occur in large populations and
are easily collected. Often these populations occur in relatively
restricted areas on slopes and are probably exploiting an
optimum habitat not immediately evident to the observer.
Ecological relationships with other species: Most Californian
xystodesmid species occupy mutually exclusive geographic
regions, probably a result of competition for the same niche;
such is the case with species of Wamokia. Immediately south
of the Cosumnes River, the distribution of Wamokia ceases
|
( 4
Milliped Genus Wamokia 513
and Amplocheir Chamberlin becomes abundant and continues
south through Madera County. Like Wamokia, Amplocheir
species inhabit areas of clayey soil and shallow leaf litter, yet
the soil and vegetation are qualitatively different than in areas
inhabited by Wamokia. The closest northern neighbor of
Wamokia appears to be Hybaphe Cook, which occurs in
Tehama and Shasta Counties in California.
Only two other genera of xystodesmids are known to occur
sympatrically with Wamokia. Sigmocheir Chamberlin which
occurs throughout the central and southern Sierras, was col-
lected on the same hillside with W. hoffmani new species, at
the Cosumnes River southeast of Latrobe. Also, an undescribed
species of Hybaphe, to be described elsewhere, occurs in the
American River Canyon near Auburn. Near the bottom of the
south wall of the canyon, where vegetation is luxuriant and the
soil rich, the Hybaphe species is found. Higher on the wall,
where the slope is steeper, soil leached, and litter thin, one
finds many W. discordis new species, and very few Hybaphe.
Presumably, competition with Hybaphe and Sigmocheir is
avoided in these cases by different preferential habitats.
TAXONOMIC CHARACTERS
Gonopods: As in most groups of millipeds, the gonopods ex-
hibit obvious interspecific differences. As with any structure
which is so genetically pliable, however, considerable variation
occurs intraspecifically.
The telopodite branches about one-third its length from
the hinged connection to the coxa into two structures, a long,
narrow tibiotarsus carrying the seminal canal, and a femoral
endite of varying shapes and sizes, usually with a mesally
curving distal process and a smaller lateral process.
The tibiotarsus is simplest, hence offering fewer diagnostic
characters. On the lateral margin distad of the femoral process
is a spine in some species. In specimens of discordis or sierrae
new species, the spine may be present or absent. In species
which constantly possess it, variation is expressed in relative
size of the spine, whether it is simple or bifid, and even its
position on the tibiotarsus.
514 Proceedings of the Biological Society of Washington
The distal portion of the tibiotarsus curves strongly mesad
in all species except remota new species, the nature of this
curvature being characteristic of the species.
The femoral endite possesses a serrate ventrobasal margin
in remota and dentata new species, the other species all having
smooth margins in the specimens examined. The body of the
prefemoral endite basically possesses two processes, a distal
process curving mesad, and a small lateral process. One
species, hoffmani, possesses a basal process as well, which
projects mesad. The terminology used for these structures
was constructed only for convenience, so homologies are not
necessarily presumed between related groups for all structures
bearing the same terms.
Variation in the femoral endite is trenchant, the size, shape
and curvature of the distal process being generally constant
within species, the only exception being discordis, in which the
apex may be acute or rounded. The lateral process is useful
only with regard to general size. Although well developed in
most specimens of remota, in one specimen the lateral process
is represented only by a rounded bump. In discordis it may
be simple or bifid, and the length may vary up to 50 percent.
Cyphopods: The mesoproximal aspect of the cyphopods
displays amazing differences between some species, although
the usefulness of these structures is partly canceled by the
strong variability present. Nonetheless, most female specimens
can be identified by the cyphopods.
Sexual dimorphism: During our field work with Wamokia,
we were soon able to easily distinguish male from female
specimens from a considerable distance, the following char-
acteristics making this possible. Female length/width ratios
are lower than in males. Accentuating this disparity, each
paranotum of females projects out less than 25 percent the
width of the body cylinder, whereas in males the paranota
approach 40 percent of the cylinder width. Also, females
possess very thin legs, whereas in males, the legs are quite
robust. Females are also slightly longer than males on the
average.
Milliped Genus Wamokia 515
Genus Wamokia Chamberlin
Wamokia Chamberlin, 1941. Bull. Univ. Utah, Biol. ser. 6(5):14.
Waimokia Chamberlin and Hoffman, 1958, Lapsus calami. Bull. U. S.
Nat. Mus. No. 212:53; Buckett, 1964, Lapsus calami, Ann. list
Diplopoda California, p. 10.
Type species: Wamokia placera Chamberlin, by original designation.
Diagnosis: Wamokia may be recognized by the small size (19-32 mm),
thin exoskeleton with orange-brown coloration and with orange, acute
paranota; second and third legs of male with coxal processes present and
sternal processes lacking; male gonopods with coxae joined firmly to-
gether, telopodite with long, simple tibiotarsus and simple or complex
prefemoral process.
In order to facilitate future comparison of xystodesmid genera, a
thorough description of the genus is presented below.
Description: Color in life usually beige on prozonites, medium grey
on metazonites and bright orange on both surfaces of lateral and posterior
margins of paranota; sometimes orange color extends across metazonite;
telson orange dorsally; a dark dorsomedial longitudinal line present.
Head oval, with coronal suture smooth, prominent, dividing between
antennae into two frontal sutures which are evident only for distance
between antennae; one pair of long supra-antennal setae present on
each side of coronal sutures; frons with a pair of setae on each frontal
suture, upper setae between antennae and lower seta just ventromesad
of antennae; frons smooth, but with shallow transverse sculptured lines;
lower part of frons with transverse row of setae; another row of setae
lining lateral margin of facial shield; clypeus with transverse row of
about twenty setae; labrum also with about twenty setae; genae smooth,
with impressed groove parallel to supra-antennal groove; antennae
widely separated, distance between them equal to length of second
antennal segment; first antennal segment subcircular and broader than
other segments, with two setae longer than segment; second segment
long, proximally curving slightly dorsad, then curving slightly in opposite
direction, with long distal setae; segments 3-5 subequal in size and
shape, slightly shorter than segment 2, narrow proximad and wide
distad; segment 6 subequal in length to segment 2, but narrower, more
densely setose; segment 7 small, densely setose, truncate; four terminal
sense cones set in concavity of apex.
Body subcylindrical, paranota moderately developed, projecting out
from body on each side 20 percent to 40 percent width of cylinder,
projecting downward at about thirty degrees from horizontal at just
above mid-body height, posterior margin higher than anterior margin;
dorsa of tergites smooth; prozonites separated from metazonites by
shallow groove dorsally, this groove disappearing below paranotal
margins; repugnatorial pores round, opening laterally on paranota near
posterior margin; pleural areas smooth, coriaceous; anterior tracheal
516 Proceedings of the Biological Society of Washington
remota
discordis
placera dentata
falcata
Fic. 1. Dendrogram showing relationships within species of Wamokia.
aperture oval, vertically elongate; posterior aperture oval, less elongate.
Sterna smooth, raised out slightly from level of body mesially, but much
laterally, the coxae projecting forty-five degrees down from horizontal;
sterna not produced up on legs, a transverse groove present on sternum
between legpairs on each segment.
Collum wider than head but narrower than following segments, with
anterior margin curving in even semicircular arc, the posterior corners
even with posterior margin of segment; collum not much arched, not
reaching ventrad to paranotal margin of second tergite; segments 3-8
with antero-lateral corners increasingly rounded, posterolateral corners
increasingly produced; near posterior end of body much of lateral
margins of paranota nearly straight; segments 17 and 18 progressively
smaller and narrower, segment 19 much reduced, paranota often not
.
%
qt)
Milliped Genus Wamokia 517
sierrae
|
NEVADA
2
PLACER
remota
|
B |
discordis
EL DORADO
placera
2 a falcata
= een oO
\ @ A
\ ® dentata
\ hoffmani ®
\ @
AMADOR
Fic. 2. Distribution map of species of Wamokia.
projecting caudad beyond posterior corners of segment 18; telson sub-
triangular, broader than long, with two pairs of dorsal setae, one pair
of setae on lateral margin cephalad, one pair laterad of posterodorsal
setae, and two pairs of apical setae; anal valves smooth, distinctly
surpassed by mucro, with two pairs of setae lining lips; preanal scale
518 Proceedings of the Biological Society of Washington
large, sub-lenticular, about half as long as broad, with a pair of setae
on posterior margin; tergum of twentieth segment visibly ringing entire
segment anteriorly.
Legs long, femur exceeding lateral margin of paranota; male with leg
segments much broader and more robust than female; legs with coxa
cylindrical, slightly longer than broad; prefemur subequal in length
to coxa, lateral margin bulging; femur the longest segment, very narrow
basally and widening apically; postfemur the shortest segment and
narrower than previous segments; tibia equal in length to postfemur,
but narrower; tarsus narrowest segment, and about as long as prefemur;
tarsal claw large and curved mesially, legs generally sparsely setose,
the tarsus more densely setose, and prefemur possessing a long seta
based just proximad of distal spine; distal subtriangular prefemoral
spine beginning on segment 9 and progressively increasing in size
posteriorly, the posterior leg of each segment with slightly greater spine
development; second coxae of male each with a short, subcylindrical
ventral process; third coxae with larger pair of cylindrical processes
produced anteroventrad; legs 1 and 2 with coxae adjacent, projecting
ventrad; legs 3-7 with coxae increasingly separated; legs 8-28 with
coxae separated by about two coxal widths, the posterior legpair of
each segment separated slightly more than anterior legpair.
Gonopod socket large, oval, extended cephalad three-fourths width
of prozonite and caudad to within two-thirds coxal width of posterior
margin of segment.
Gonopods large, wholly filling socket, with telopodite and much of
coxa extruding from body cavity, the telopodite joined to coxa at ninety
degrees and produced cephalad, projecting between seventh legs; coxae
very large, broad, with a short coxal apodeme produced proximad from
mesal margin of coxa; telopodite rigid, but fixed to coxa at a flexible
joint.
Coxae of gonopods joined together by transverse muscle tissue as
well as the sclerotized sternal connective where coxae meet distally.
Telopodite with femoral endite of varying shapes emerging from the
femoral region and situated cephalad of tibiotarsus; tibiotarsus long and
thin for most of its length, often apically curving mesad and sometimes
again distad, and sometimes with a lateral spine located about mid-
length; femoral endite composed of main body from which project two
or three processes, a distal process which projects mesodistad, a basal
process which projects mesad and a small lateral process which projects
laterad; solenite emerging from coxa distally on anterior face mesad of
telopodite base, immediately arching up to telopodite away from body
of gonopod, proceeding distad in seminal canal mesad of femoral endite,
thence along center of tibiotarsus to apex.
GENERIC RELATIONSHIPS
Characters of Wamokia appearing most significant at the generic level
are the following: gonopods with coxae large and rigidly joined together
ML Oe aa
a
Milliped Genus Wamokia 519
Fics. 3-6. 3. W. placera, male. Dorsal aspect of tenth tergite.
4. W. placera, bases of second and third legs. 5. W. dentata, paratype
male. Gonopods in vertical position. 6. W. dentata, paratype male.
Gonopods, posterior aspect.
Milliped Genus Wamokia pal
with a sternal structure; telopodite movably hinged to coxa and bent at
right angle; second and third legs of male with rounded coxal projections;
second legs of female with cylindrical processes on coxae; dorsum smooth,
with cuticle thin and paranota with acute posterolateral corners; color
yellow-brown with orangetipped paranota.
On the basis of the characters given above, Wamokia appears to be
closely related to Amplocheir, an inhabitant of the more southerly Sierra
Nevada of California. Differing only in the configuration of the gonopod
telopodite and presence of dorsal tubercles, Amplocheir can be easily
mistaken for Wamokia in the field. That the genera should be maintained
with separate status is a belief based primarily on the discontinuity in
the gonopods, which is considerable. A third genus in this group is
Paimokia Chamberlin (sensu stricto). Sharing most characters with
Wamokia and Amplocheir, Paimokia differs in having the dorsum well-
tubercled, and the paranota rounded rather than angular. Motyxia
Chamberlin and the Xystocheir group (Cheirauxus Chamberlin, Delocheir
Chamberlin) appear to be more remotely related to Wamokia, with
Motyxia pior Chamberlin possessing sternal processes on the fourth
legs, no coxal processes on the third legs, rounded paranota and a
differently shaped gonopod aperture, with gonopod coxae joined near
their bases. Xystocheir acuta Cook from Palo Alto, California, possesses
sternal processes on third and fourth legs, a larger, much more robust
body with much thicker cuticle, tuberculate dorsum and a differently
shaped gonopod aperture.
SPECIES RELATIONSHIPS
The species of Wamokia may be broken into two general groups,
which differ by the presence or absence of a torsion in the femoral
endite of the male gonopods. The four southern species, dentata, falcata,
hoffmani, and placera lack the torsion and form a rather close group,
although hoffmani possesses the unusual basal process. Further to the
north, the species discordis, sierrae and remota form a group in which
the femoral endite is twisted at the base so that the distal process curves
oppositely from that of the other species group. Setting the small, simple
femoral process as the primitive condition, falcata would be the least
modified species and discordis or remota the most modified. A dendro-
gram expressing these ideas is presented (see fig. 1).
<
Fics. 7-10. (Right male gonopods). 7. W. falcata, paratype; anterior
aspect. 8. W. falcata, paratype; mesal aspect. 9. W. placera; anterior
aspect. Collected 3 miles southwest of Placerville, E] Dorado County.
10. W. placera, holotype male; mesal aspect. Collected from Placerville,
El Dorado County.
522 Proceedings of the Biological Society of Washington
Kry To THE SPECIES OF Wamokia BASED ON MALE GONOPODS
1. Distal process of femoral endite rounded, or arched strongly
caudad, often exceeded in length by lateral process
Pe eld Winch ce Rt eh ER ML et discordis Buckett & Gardner, n. sp.
Distal process curving mesodistad, much longer than lateral
PDEOCESS 12220 aU el ied cal ida CR RE Fyne op Ren ce ee Se A 2
2. Femoral endite with two processes projecting mesad, a falcate
distal process, and a truncate basal process ~
Ea NN ee Atha ea aah CRA RO GH hoffmani Buckett & Gardner, n. sp.
Femoral endite with only one mesially projecting process __._- 3
3. Distal process of femoral endite arching mesad, with lateral margin
(6{0) Shi, =) Gane RE MR ROR Re ee Same ne a Pteetae OS Un Ra WMS AE 4
Distal process of femoral endite arching distad, with lateral margin
COTGAVE sre ee PRIN UES LUN Pera ESN ela Bd Uh RI ea 6
4. Tibiotarsus lacking lateral spine; lateral process of femoral endite
subequal in width to distal process placera Chamberlin
Tibiotarsus possessing distinct lateral spine; lateral process of
femoral) endite-sminu ter; Wes. ee NE ee 5
5. Mesal margin on base of femoral endite serrate; lateral spine basad
of apex of distal process ______ dentata Buckett & Gardner, n. sp.
Mesal margin on base of femoral endite entire; lateral spine distad
of apex of distal process falcata Buckett & Gardner, n. sp.
6. Tibiotarsus not much arched mesad; lateral spine prominent; mesal
margin of base of femoral endite serrate
UD CAA Rea a eli tee ar POW egae remota Buckett & Gardner, n. sp.
Tibiotarsus strongly curved mesad; lateral spine minute or obsolete;
mesal margin entire _____.___ sierrae Buckett & Gardner, n. sp.
Wamokia discordis Buckett and Gardner new species
(Figures 11, 12, 18, and 28)
Holotype: Male, 1 mile east of Auburn, Placer County, California,
placed in the Type Collection of the Department of Entomology, Uni-
versity of California, Davis (UCD).
Diagnosis: Similar to sierrae and remota in that the femoral endite
curves so that the mesal margin is convex, but distinguished from them
by the broad cephalic face of the femoral endite, with a short, sharply
curved or apically rounded distal process, and a lateral process which
almost equals distal process in length.
Description: Head with vertex and frons equally trans-microstriate;
coronal suture well developed, frontal sutures not apparent; lower part
>
Fics. 11-14. (Right male gonopods). 11. W. discordis, paratype;
anterior aspect. Collected 1 mile north of Cool, El Dorado County.
12. W. discordis, paratype; anterior aspect. Collected 1 mile east of
Auburn, Placer County. 13. W. dentata, paratype; anterior aspect.
14. W. dentata, paratype; mesal aspect.
523
Milliped Genus Wamokia
Pir 7 SF ey
524 Proceedings of the Biological Society of Washington
of frons with irregular transverse row of 8 setae; clypeus with irregular
row of 22 setae; labrum with row of 30 setae; gular groove very shallow;
a series of 6 setae present along lateral margin of facial shield between
labrum and ventrodistal corner of antennal groove. Tergites smooth,
with minute impressed longitudinal striae; collum with front margin
evenly curving, caudolateral corners not surpassing posterior margin
of segment; segment 19 narrower than anterior width of segment 18,
with paranota of 19 projecting caudad only slightly beyond posterior
margin of segment; telson subtriangular, with 14 setae normally placed.
Pleural area smooth; sterna low medially, pronounced laterally, smooth
except for minute impressed longitudinal striae, a distinct transverse
groove present between legpairs of each segment; legs as described for
genus, with numerous setae on ventral surfaces of first three segments;
legs separated by distance of less than three coxal lengths; second
legs with a pair of small, mesal, cylindrical coxal processes on posterior
surface, a minute pair of swellings on anterior surface; third legs with
coxal lobes larger and flatter than those of second legs, these bearing
small setae and lacking lateral spine.
Gonopods with tibiotarsus narrow, curving evenly mesad beyond
femoral endite, recurving slightly apically, tip acuminate; femoral branch
narrow basally, widening to broad surface, rounded dorsally and
ventrally, with an elongate, straight lateral process and a broader distal
process curving sharply distad and narrowing abruptly to acute apex;
mesal margin of femoral endite smooth.
Female: As in male, but second legs with coxa thin, subtriangular,
with small ventral process apically, mesad of joint with prefemur;
cyphopod aperture as wide as sternal area of sixth segment, closed
along posterior margin by thin sclerotized bar; socket narrowed mesally,
thus longest in region of each cyphopod; anterior margin of cyphopod
aperture curving evenly; cyphopods as in figure 28.
Specimens examined: Holotype male: CALIFORNIA: Placer County:
1 mile east of Aubum, 28 January 1968 (M.R., R.C., J.L., B.W. & K.B.
Gardner). Paratypes: 28 males, 32 females, same data as holotype; El
Dorado County: 8 males, 6 females, 1 mile north of Cool, 4 February
1968 (M.R.G.); 1 male, 3 females, 1 mile west of Greenwood, 4
February 1968 (M.R.G.); 6 males, 6 females, 2 miles west of George-
town, 4 February 1968 (M.R.G.).
Variation: Statistics: Cool population: males, length 20-23.5mm;
length/width 5.4-6.0; females, length 21.5-26mm; length/width 5.0—5.3;
>
Fics. 15-18. (Right male gonopods). 15. W. remota, paratype;
mesal aspect. Collected 3 miles southwest of Foresthill, Placer County.
16. W. remota, paratype; anterior aspect. Collected from Foresthill,
Placer County. 17. W. remota, paratype; anterior aspect. Collected 3
miles southwest of Foresthill, Placer County. 18. W. discordis, paratype;
mesal aspect. Collected 1 mile east of Auburn, Placer County.
525
Milliped Genus Wamokia
526 Proceedings of the Biological Society of Washington
average population length 23.0mm. Auburn population: males, length
21.5-28mm, length/width 6.1-6.8; females, length 21.5-30mm, length/
width 5.1-5.6; average population length 23.4mm. Populations east of
Cool: males, length 25-28mm, length/width 6.0-6.1; females, length
28-32mm, length/width 5.2-5.7; average population length 28.2mm.
Perhaps the greatest infraspecific variability of the genus occurs in
this species. Specimens from Auburn vary in degree of development
of the lateral process of the femoral endite, most specimens with this
process being a well developed simple or bidentate spine, although
one specimen, probably a mutant, lacked the lateral process on both
gonopods. The distal process of the femoral endite is always sharply
curved and acutely pointed. The lateral spine of the tibiotarsus is
minute on some specimens, lacking on others.
Specimens from Cool often possess a rather distinct lateral spine on
the tibiotarsus, with the femoral endite exhibiting a rounded distal
margin rather than notched as in the Auburn population. The distal
process is sharply recurved so that it appears rounded from the
anterior aspect (see fig. 11). Populations east of Cool have the
distal margin of the femoral endite notched or acutely produced; the
lateral process is short and bidentate to long and simple; the lateral
spine of the tibiotarsus is small but distinct.
Ecology: Collected at 1,000’ on the American River at Auburn and
2,200’ west of Georgetown, discordis occupies perhaps the most diverse
range of conditions found in the genus. Near the top of the northeast-
facing canyon wall of the American River, the soil is red, duff is shallow,
Quercus wislizenii and kelloggii low and scrubby, and Arctostaphylos
common. At the highest elevation, near Georgetown, where the hillside
remains partially covered with snow, Pinus spp. provide the dominant
ground litter, although tall oaks and smaller herbaceous plants abound
on the hill also. The ground litter is generally thick (2-8cm). Speci-
mens were found in areas of the slope where low brush was minimal, and
oak leaf litter the most prevalent. The large size of individuals at the
higher elevations is possibly a reflection of the richer food source
available.
Distribution: Four collections have been made in a line 12 air-miles
long between Georgetown and Auburn, with one collection on the west
bank of the American River eliminating the river as an effective barrier.
Closely bordered by remota to the north and placera to the south,
discordis is probably a localized form.
>
Fics. 19-22. (Right male gonopods). 19. W. sierrae, paratype;
anterior aspect. 20. W. sierrae, paratype; mesal aspect. 21. W. hoffmani,
paratype; anterior aspect. Collected 1 mile north of Shingle Springs, El
Dorado County. 22. W. hoffmani, paratype; mesal aspect.
527
Milliped Genus Wamokia
Zim
ee SSS
iii
ae
Pip =
Pye ee
528 Proceedings of the Biological Society of Washington
Wamokia hoffmani Buckett and Gardner new species
(Figures 21, 22, and 27)
Holotype: Male, 1 mile north of Shingle Springs, El Dorado County,
California; deposited in Type Collection, (UCD).
Diagnosis: Resembling falcata and dentata in the small distal process
and very small lateral process of the femoral endite, but distinguished
from these and other known members of the genus by the possession of
two processes projecting mesad from the femoral endite. The lateral
process may be present or absent.
Description: Holotype male: Color in life orange on lateral tips of
collum and other segments, this color extending across posterior part of
metatergite, a longitudinal medial black stripe present from collum to
antepenultimate segment. Head with smooth, deep coronal suture,
dividing between antennae to inconspicuous frontal sutures; vertigial
area dorsad of antennae with a number of small wrinkles; 4 supra-
antennal, 9 frontal, 20 clypeal and 22 labral setae present; 7 setae along
lateral margin of facial shield between labrum and ventral margin of
antennal groove; gular area with broad, shallow groove parallel to
antennal groove. Collum typical of genus, much exceeded ventrally by
second segment; tergites generally smooth with fine longitudinal striae
on anterior half of metatergite, posterior half with about seven low
longitudinal wrinkles on each side of segments of caudal half of body;
segments 17-19 reduced in size, telson much surpassing anal valves and
possessing regular 14 setae; anal valves well developed, with 2 pair of
setae. Metazonites ventrally sunken at point of contact with prozonites,
sterna raised out from body level mesally about one-fourth their lateral
heighth, a shallow longitudinal groove present on sterna between anterior
and posterior legs of each segment; posterior and anterior pairs of legs
separated by the same distance, equal to two coxal widths.
Gonopods with lateral spine located on tibiotarsus about 1.2 x distance
from origin of femoral endite to its apex; tibiotarsus strongly curving
mesally distad of lateral spine, the apex acuminate and slightly recurved;
femoral endite short, broad, with acute distal process longest, projecting
mesocaudad; basal process more directly mesad, truncate with uneven
mesal margin; lateral process distinct, narrow and acute.
Female: As in male, except for secondary sexual characters; second
legs with coxa flat, a distinct cylindrical process present; cyphopod
aperture with posterior margin evenly rounded, a low mesal swelling
present; cyphopods as in figure 27.
Specimens examined: Holotype male: CALIFORNIA: El Dorado
County: 1 mile north of Shingle Springs, 20 January 1968 (J. S. Buckett
& M. R. Gardner). Paratypes: 14 males, 34 females, same data as
holotype; 2 males, 2 females, 2 miles north of Shingle Springs, 16 April
1965 (M.R.G.); 37 males, 19 females, 4 miles southwest of Shingle
Springs, 4 February 1968 (M.R.G.); 2 males, 2 females, 6 miles south
of El Dorado, 28 November 1964 (J.S.B. & M.R.G.); Amador County:
i
a
a
:
4
Milliped Genus Wamokia 529
Fics. 23-26. (Female cyphopods, mesal aspect). 23. W. dentata,
paratype. 24. W. placera. Collected 2 miles north of Placerville, El
Dorado County. 25. W. remota, paratype. Collected 3 miles southwest
of Foresthill, Placer County. 26. W. placera, paratype female; sternum
and basal segments of second legs.
2 males, 2 females, Cosumnes River, southeast of Latrobe, 26 November
1965 (M.R.G.).
Variation: Population from 4 miles southwest Shingle Springs: males
length 21-24mm, length/width 5.4-5.9; females length 23-25mm,
length/width 4.8-5.6; average population length 23.3mm. Population
530 Proceedings of the Biological Society of Washington
from 1 mile north of Shingle Springs: males, length 23-26mm, length/
width 5.6—-6.2; females length 23-28mm, length/width 4.8-5.5; average
population length 24.5mm.
Specimens from southwest of Shingle Springs differ from the holotype
by possessing a longer distal than basal process on the femoral endite,
and either lacking the lateral process entirely or exhibiting a small,
rounded protrusion. Also in this population the lateral spine of the
tibiotarsus is minute, and variable in its position. Specimens from
Cosumnes River possess a distinct lateral process on the prefemoral
endite and a large lateral spine, which is positioned lower on the
tibiotarsus than in other populations. Individuals from south of El
Dorado have the lateral process either lacking or small and spiculate;
the lateral spine of the tibiotarsus is prominent and low as in individuals
of the Cosumnes population.
Ecology: W. hoffmani has been collected at elevations of 600’ to
1,400’ in areas of red soil with plant cover of mixed stands of Quercus
wislizenii and kelloggii, and with occasional specimens of Pinus sabiniana.
Populations of millipeds were dense at least at the two localities near
Shingle Springs, numbers averaging about 20 individuals per square
yard of ground.
Distribution: This species possesses the greatest known distribution, ©
extending longitudinally from the south bank of the Cosumnes River
to 2 miles north of Shiugle Springs, a distance of about 15 air-miles.
Latitudinally, the distribution is 8 miles wide.
Wamokia placera Chamberlin
(Figures 3, 4, 9, 10, 24, and 26)
Wamokia placera Chamberlin, 1941. Bull. Univ. Utah, biol. ser., 6(5):
14, fig. 27.
Type: Holotype: 9 miles north of Placerville, El] Dorado County,
California, probably in the vicinity of Coloma; presently in the collection
of the late R. V. Chamberlin.
Diagnosis: Characterized by the development of the femoral endite
into an almost straight, acute distal process and a broad, prominent
lateral process which projects lateroproximad, and is subequal in width
to distal process.
Description: Male: Head with usual coronal suture, frontal sutures
weak; head smooth, but with very fine, impressed vertical lines on
vertex; short horizontal lines on frons and gula; frons with irregular row
of 9 setae; clypeus with row of 23 setae and labrum with 22 setae;
gular region with shallow, although distinct, groove parallel to lateral
>
Fics. 27-30. (Female cyphopods, mesal aspect). 27. W. hoffmani,
paratype. Collected 1 mile north of Shingle Springs, El] Dorado County.
28. W. discordis, paratype. Collected 1 mile east of Auburn, Placer
County. 29. W. sierrae, paratype. 30. W. falcata, paratype.
532 Proceedings of the Biological Society of Washington
margin of facial shield; a row of 6 right and 4 left setae present along
lateral margin of gula from clypeus to half the distance toward antennal
groove; antennae of normal proportions, spaced by distance equal to
length of third segment. Collum wider than head, anterior margin
evenly curving with posterior corners back as far as posterior margin
of segment. Tergites generally smooth, with short, longitudinal etchings;
last four segments becoming reduced, telson with usual 14 setae, much
surpassing well developed anal lips; preanal scale longer than half its
width, obtusely angular caudad. Sterna raised very little mesially, half
a coxal width laterally, with fine longitudinal striations and a shallow
transverse groove present between legpairs of each segment. Legs
long, of usual segmental proportions, on postgenital segments separated
by about 2.3 coxal lengths; second legs with small pair of ventral coxal
processes, third legs with broader, longer processes located relatively
further laterad than those of leg 2.
Gonopods with telopodite distally narrowing to half width of femoral
region; tibiotarsus curving distad and slightly mesad for two-thirds its
length, then strongly curved inward and slightly proximad, the apex
acuminate and curved again mesodistad, lateral spine lacking; femoral
endite very narrow near base, produced strongly cephalad away from
tibiotarsus, with distal process long, straight and acute, and lateral
process curved proximad, short, but stout, its basal area as wide as
distal process; mesal margin of endite smooth, not produced mesad, and
separated from femur for much of its length.
Female: As in male, except for the following sexual characters:
posterior margin of cyphopod socket curved, meeting mesad between
cyphopods; second legs with coxa thin, subtriangular, with a distinct
cylindrical distal process; cyphopods as in figure 26.
Specimens examined: CALIFORNIA: El] Dorado County: 9 males,
8 females, 2.1 miles north of Placerville, 4 February 1968 (M. R.
Gardner); 7 males, 12 females, 3 miles southwest of Placerville, 13
January 1968 (J. S. Buckett & M. R. G.).
Variation: Population southwest of Placerville: males, length 22.5—
28mm, length/width 5.6—-5.9; females, length 26-30mm, length/width
5.0-5.4; average population length 26.8mm. Population north of
Placerville: males, length 27-29mm, length/width 5.3-5.9; females,
length 26—3lmm, length/width 5.2-5.5; average population length
28.5mm.
Of the two collections we have made, variability in the gonopods is
insignificant, a slight difference in the curvature of the tibiotarsus being
the only evident difference between the two populations.
Ecology: Collections have been made by us at elevations around 1,600’.
The type collection, if from near Coloma, was made at about 800’. The
localities at which we collected placera were on northeast facing 30°
slopes, with the oaks Q. wislizenii and kelloggii dominating the com-
position of the ground litter, the soil being red Aiken Clay.
Milliped Genus Wamokia 533
Distribution: We have made two collections 4 miles apart in the
vicinity of Placerville. Including the type locality, 9 miles north of
Placerville, the known range exceeds a little over 10 air-miles in a north-
south direction, and will probably not be found to extend much further.
Wamokia dentata Buckett and Gardner new species
(Figures 13, 14, and 23)
Holotype: Male, 3 miles northwest of Somerset, El Dorado County,
California; deposited in Type Collection, (UCD).
Diagnosis: Similar to falcata in configuration of the distal and lateral
processes of the femoral endite, but distinguished by the dentate mesal
margin of the femoral endite and more basal location of lateral spine
or telopodite.
Description: Holotype male: Color typical of genus, orangish-grey
dorsally with orange on both surfaces of lateral and posterior margins
of paranota. Head with coronal suture smooth, prominent, dividing
between antennae to two frontal sutures which are evident only for
distance between antennae and lower seta just ventromesad of antennae;
frons smooth, but with shallow transverse sculptured lines; lower part
of frons with transverse row of 8 setae; 5 setae in row along lateral
margin of facial shield from clypeus half the distance to antennae;
clypeus with transverse row of 22 setae; labrum with transverse row of
33 setae; genae smooth, with impressed groove parallel to supra-
antennal groove; antennae widely separated, distance between them
equal to length of second antennal segment; antennal segments as in
generic description. Collum wider than head but narrower than follow-
ing segments, anterior margin curving in even semicircular arc, the
posterior corners even with posterior margin of segment; segments 17
and 18 progressively smaller and narrower, segment 19 much reduced,
paranota projecting caudad only slightly beyond posterior corners of
segment 18; telson subtriangular, broader than long, with 14 setae; anal
valves smooth, much surpassed by mucro, with 2 pairs of setae lining
lips; preanal scale large, sublenticular, about half as long as broad, with
a pair of setae on posterior margin. Sterna smooth, raised out from
level of body and curved with lateral margins much more produced
than mesal area, the coxae projecting at 45° from horizontal, a transverse
groove present; legs with prefemoral spine beginning on segment 9 and
progressively increasing in size posteriorly, the posterior leg of each
segment with greater spine development; second coxae each with large
cylindrical process produced anteroventrad.
Gonopods similar to placera, with tibiotarsus long and narrowing
evenly though rapidly from connection with femur, and bent mesad near
distal margin, with apex curved again distad; lateral spine of tibiotarsus
minute; femoral endite with basal region small, but distal process large,
curving evenly mesodistad and narrowing evenly to acuminate apex;
534 Proceedings of the Biological Society of Washington
mesal margin of femoral endite serrate; lateral process elongate and
truncate, though small.
Female: Female with prominent setose processes on flat second coxae;
cyphopod socket with posterior rim swollen mesally. Cyphopods as in
figure 23.
Specimens examined: CALIFORNIA: El Dorado County: Holotype
male, 3 miles northwest of Somerset, 20 January 1968 (J. S. Buckett &
M. R. Gardner). Paratypes: 3 males, 15 females; same data as holotype.
Variation: Male, length 22-24mm, length/width 5.3-6.0; female,
length 25-28mm, length/width 5.1—5.6. Gonopods not noticeably vari-
able.
Ecology: The one collection of this species was made on a gently
(ca. 12°) sloping hillside covered with Q. wislizenii, with thin surface
litter. Notably, this is the only collection of Wamokia made in an area
of black soil.
Wamokia falcata Buckett and Gardner new species
(Figures 7, 8, and 30)
Holotype: Male, Camino, E] Dorado County, California; deposited
in Type Collection, (UCD).
Diagnosis: Distinguished from dentata, its closest relative, by the
lack of a serrate mesal margin of the femoral endite, and by the presence
of prominent lateral spine located well distad of the apex of femoral
endite.
Description: Holotype male: Color faded in alcohol. Head with
frontal sutures barely evident, with frontal and supra-antennal setae
present; lower part of frons with transverse row of 8 setae; clypeus
with 20 setae in transverse row; labrum with 23 setae in transverse row;
5 setae in row along lateral margin of facial shield from clypeus to
half the distance to antennae; gena with groove very shallow, barely
evident; antennae of normal configuration, separated by distance equal
to length of third antennal segment, first, second and third segments
with long distal setae. Collum with posterior margin convex laterally,
concave mesially, with rather sharp posterior corners; tergites smooth;
nineteenth segment small, exceeded by paranota of eighteenth segment;
telson with 2 pairs of lateral setae, posterior pair just cephalad of
posterior dorsal setae, with 2 pairs of terminal setal sockets, though
setae missing; telson surpassing anal valves, which are smooth, with 2
pairs of setae lining anal lips. Sterna almost even with prozonites
mesially, raised out laterally, longitudinal wrinkles present mesad of leg
sockets; prozonites separated from metazonites by narrow rim ringing
segment, followed by shallow groove. Legs long, extending well beyond
lateral margins of segment, the relative length of segments from longest
to shortest being 3, 2, 6, 5, 1, 4; both anterior and posterior coxae
with row of 10-12 stout setae and 1 long seta.
Milliped Genus Wamokia 535
Gonopods large, well exserted from socket, with tibiotarsus broad
proximad of large lateral spine, the narrow distal continuation arching
mesocephalad, the apex acuminate, recurved; femoral endite much
shorter than tibiotarsus, arising mesocephalad from half the overall
length of telopodite, projecting strongly laterad to lateral margin of
tibiotarsus, then curving evenly mesodistad and narrowing gradually
to acuminate apex, a small rounded lateral process present; the mesal
margin of base of femoral endite much produced, entire, concealing
part of tibiotarsus from cephalic aspect.
Female: Second leg with coxa flat, subtriangular, and with a promi-
nent elongate distal process; cyphopod aperture with anterior margin
essentially straight for much of its length, with small, abruptly pro-
duced mesal process. Cyphopods as in figure 30.
Specimens examined: CALIFORNIA: El Dorado County: Holotype
male, Camino, 24 December 1964 (S. Seminoff). Paratypes: 1 male,
2 females, same data as holotype.
Distribution: Known only from one collection, this entity is bounded
on the west by placera, and on the south by dentata. The situation
to the north is not known.
Wamokia remota Buckett and Gardner new species
(Figures 15, 16, 17, and 25)
Holotype: Male, 8 miles northeast of Auburn, Placer County, Cali-
fornia; deposited in Type Collection, (UCD).
Diagnosis: Distinguished from most species in the genus by the
presence of a long distal process, subequal to tibiotarsus in length and
exceeding it in width; tibiotarsus not curving strongly mesad as in other
species of the genus. Distinguished from sierrae by the presence of a
prominent lateral spine and serrate mesobasal margin of femoral endite.
Description: Holotype male: Color in life bright orange on tips of
collum and paranota; beige across prozonites dorsally, light brown across
metatergites, telson orange.
Head with 2 pairs of supra-antennal setae, uneven transverse row of
8 frontal setae, about 20 prominent clypeal setae and 23 prominent
labral setae; head smooth and shining above, sparsely transversely
sculptured medially, this fine sculpturing dense on ventral third; antennae
well spaced, slightly less than length of second segment apart. Dorsum
of tergites smooth except for short, minute longitudinal striae; collum
with anterior margin forming smooth arc, with least curvature mesad
over head, posterior margin sinuous, convex laterad and concave mesad;
segments 17-19 increasingly reduced, paranota of 19 small, rounded lobes
barely projecting caudad beyond posterior margin of segment; telson
with usual 4 dorsal, 6 lateral, 4 apical setae; anal lips moderately
prominent. Sterna finely striate, on postgenital segments medially low,
barely raised out from level of body, with a longitudinal and transverse
groove, laterally more pronounced, lacking longitudinal wrinkles; legs
536 Proceedings of the Biological Society of Washington
separated by distance equal to two coxal lengths, posterior legs of median
body segments slightly more separated than anterior legs; pregenital
legs with sterna more produced; prefemur of legs with apical spine,
femur becoming very long at posterior end of body.
Gonopods with tibiotarsus broad, shorter than in other species,
curving slightly mesad though much exceeded by femoral endite;
tibiotarsus with apex notched, truncate, and with prominent lateral
spine near base; femoral endite curving laterad, then curving strongly
mesad; endite exceedingly broad, equal to twice or more breadth of
tibiotarsus distad of lateral spine, near apex curving distad toward apex
of tibiotarsus and evenly narrowing to sharply acute apex; lateral process
of endite as large as lateral spine of tibiotarsus, located proximad and
directed laterad; mesal margin of base produced, strongly dentate.
Female: Coxa of second legs with prominent cylindrical ventral
processes; posterior margin of cyphopod socket of two convex arcs
meeting mesad between cyphopods. Cyphopods as in figure 25.
Specimens examined: CALIFORNIA: Placer County: Holotype
male, 8 miles northeast of Auburn, 28 January 1968 (M. R., R. C., J. L.,
B. W. & K. B. Gardner). Paratypes: 29 males, 33 females, same data
as holotype; 11 males, 16 females, 3 miles southwest of Foresthill, 28
January 1968 (same collectors as preceding); 1 male, Foresthill, 24
April 1965 (J. S. Buckett).
Variation: Auburn population: males, length 22-24.5mm, length/
width 5.3-6.8; females, length 22.5-28mm, length/width 5.0-5.4; aver-
age population length 24.7. Population 3 miles southeast Foresthill:
males, length 21.5-24mm, length/width 5.7-6.9; females, length 24—
27mm, length/width 4.9-5.8; average population length 24.0mm.
In the gonopods, variability is found in the lateral process of the
femoral endite. In the single specimen from Foresthill, a slight bump
lies in the place of the distinct spine-like lateral process found in other
specimens. Specimens from northeast of Auburn are very similar to
those southeast of Foresthill. One difference lies in the length of the
lateral spine of some of the Auburn specimens greatly exceeding those
of the Foresthill population and being the most strongly developed in
the genus. The spine can be either simple or bidentate.
Ecology: Collections of remota have been made at elevations of
approximately 2,000, 3,000 and 3,200’, the latter two localities being
above snow line much of the winter. At the localities 8 miles north-
east of Auburn and 3 miles southwest of Foresthill, specimens were
collected in areas of red clay soil, with Q. wislizenii, Q. kelloggii the
dominant vegetation. Specimens occurred in areas of the hillside where
oak ground litter was apparently deepest, with as many as 25 individuals
occurring in a single square yard.
Distribution: Known from localities spanning 9 miles in latitude in
Placer County just north of the Middle Fork of the American River,
Milliped Genus Wamokia 537
which probably forms the southern boundary. The northern boundary
remains unknown.
Wamokia sierrae Buckett and Gardner new species
(Figures 19, 20 and 29)
Holotype: Male, 1 mile southwest of Grass Valley, Nevada County,
California; deposited in Type Collection, (UCD).
Diagnosis: Distinguished from remota by the long tibiotarsus which
curves strongly mesad, the apices overlapping in situ, with lateral spine
lacking or minute, and by the untoothed mesal margin of the base of the
femoral endite; distinguished from other species by the very long
blade-like distal process, with prominent lateral process arising directly
from base of distal process.
Description: Holotype male: Head with typical distinct coronal
suture, 1 pair of supra-antennal setae on each side; vertex and frons
marked by short, impressed striations, otherwise smooth; a pair of setae
present on frontal suture below antennae; setae between antennae
lacking; frons with uneven transverse row of 10 setae above clypeus;
clypeus with 26 transverse setae, labrum with 28 setae; gena with
shallow groove parallel to margin of facial shield; a row of about 10
setae on lateral margin of gena; antennae separated by distance equal
to length of second antennal segment; sixth antennal segment longer
than second antennal segment. Collum anteriorly curving in even posterior
arc, lateral corners round, even with posterior margin of segment;
paranota of mid-body segments evenly rounded antero-laterally, the
posterior corners acutely produced; on posterior half of body, pore
bearing segments with lateral margins of paranota swelled; segments
smooth dorsally, marked only by fine longitudinal striations; segments
17-20 progressively reduced, paranota of penultimate segment pro-
jecting caudad only slightly; anal tergite subtriangular, with usual 14
setae; anal lips weakly produced, with two pairs of setae lining lips.
Preanal scale suboval, with 2 apical setae. Sterna essentially flat,
coriaceous, low medially but raised out laterad to half coxal width, a
deep transverse groove between legpairs; legs as described for genus,
distal 3.5 segments extending beyond lateral margins of paranota; legs
2 and 3 with rounded coxal processes, those of second legs much the
smaller and projected caudad, those of third legs projected ventrad.
Prefemoral spines prominent in postgenital segments.
Gonopods with tibiotarsus narrowing beyond femoral endite, lateral
spine lacking on right gonopod, minutely represented on the left;
tibiotarsus curving laterad, then evenly mesad, the apex acute, curved
distad; femoral endite narrowing beyond base, then abruptly dividing
into broad blade-like distal process and straight lateral process; distal
process subequal to tibiotarsus mesally, but much exceeded by it distally,
the lateral margin concave; lateral process straight, acute, projecting
directly laterad.
538 Proceedings of the Biological Society of Washington
Female: Cyphopod aperture with posterior margin nearly straight,
but with low mesal swelling present; coxa of second legs more rounded
than in other species, possessing short ventral cylindrical processes.
Cyphopods as in figure 29.
Specimens examined: CALIFORNIA: Nevada County: Holotype
male, 1 mile southwest of Grass Valley, 10 February 1968 (J. S. Buckett
& M. R. Gardner). Paratypes: 5 males, 7 females, same data as
holotype. :
Variation: Males: length 21—-23mm, length/width, 5.6—-5.8; females:
length 23-24mm; length/width 4.8-5.6. Average population length
22.9mm. The lateral spine of the tibiotarsus is the only noticeable
variable gonopod character. In most specimens, the spine is absent,
but on the holotype, a minute remnant is present on the left gonopod,
though absent on the right gonopod.
Ecology: W. sierrae inhabits the most unusual econiche of the known
species in the genus. At 2,500’ elevation, the area is typical pine forest
composed of Pinus ponderosa and P. jeffreyi, with low shrubs and
occasional scrubby Quercus wislizenii and kelloggii. The collection was
made beneath the largest Q. wislizenii, where the ground was covered
with a thick mat 3-4” thick of oak leaves and pine needles fused with
fungus. Most specimens were in the zone between the red soil and the
litter mat, although a few occurred in the mat itself.
Distribution: With coly one collection known, sierrae is set off by
over 15 air-miles from the nearest collection of Wamokia. This northern
area of the generic range is where future collecting should be con-
centrated for the possible discovery of new species.
LITERATURE CITED
Buckett, J. S. 1964. Annotated list of the Diplopoda of California.
Simmons Pub. Co., Davis, California, 34 pp.
CHAMBERLIN, R. V. 1941. New western millipeds. Bull. Univ. Utah
biol. ser. 6(5): 1-23. figs. 1-30.
, AND R. L. Horrman. 1958. Checklist of the millipeds of
North America, Bull. United States Natl. Mus. No. 212:
1-236.
Storiz, R. E. anp D. F. Trusset. 1927. Soil Survey of the Placerville
Area, California. United States Dept. Agric. ser. 1927 No.
34, 35 pp. + map.
Vol. 81, pp. 539-570 30 December 1968
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
BATHYCONCHOECIA, A NEW GENUS OF PELAGIC
OSTRACODS (MYODOCOPA HALOCYPRIDIDAE )
WITH SIX NEW SPECIES FROM THE DEEPER
WATERS OF THE GULF OF MEXICO
By GrorcIANA B. DEEVEY*
Guest Investigator, Smithsonian Institution, Washington, D.C.
Specimens of ostracods taken from the stomach and intestines
of bottom fish, collected at depths of 1000 and 3165 m in the
Gulf of Mexico, were sent by Dr. Thomas Bright, then of
Texas A&M University, to Dr. Louis Kornicker at the U. S. Na-
tional Museum. Since nine of these specimens belonged to
the family Halocyprididae, Dr. Kornicker submitted them to
me for identification. The nine specimens, all mature in-
dividuals with the possible exception of one female, proved
to represent six new species of a new genus. Eight of the
specimens were found in the intestines of three species of
bottom fish, Nezumia hildebrandi, Bassozetus normalis, and
Dicromita agassizii. The shells of most of the specimens were
soft and flexible and although all the soft tissues had been
dissolved away, the exoskeleton, including that of the ap-
pendages, was more or less intact. One specimen was found in
the stomach of Nezumia hildebrandi; this specimen still re-
tained a good deal of its musculature. With one exception, the
smaller ostracods, 0.93-1.66 mm long, were found in the
intestine of Dicromita agassizii, collected at a depth of 1000 m.
The two largest specimens, 3.1 and 5.2 mm long, were taken
from the intestine of Bassozetus normalis, obtained in a dredge
sample from 3165 m depth. The smallest specimen, 0.85 mm
long, was found in the intestine of Nezumia hildebrandi, and
* Present Address: Institute of Oceanography, Dalhousie University, Halifax,
Nova Scotia, Canada.
52—Proc. Brow. Soc. WasH., Vou. 81, 1968 (539)
540 Proceedings of the Biological Society of Washington
a 1.9 mm specimen was taken from the stomach of this fish;
the last two specimens were females, the others males. A
specimen of Bathyvargula sp., a benthic form, was also found
in the stomach of Nezumia, and another Cypridinid ostracod
was taken from the intestines of Dicromita (Kornicker, in
press ).
I am very grateful to Dr. Kornicker and Dr. Bright for
permission to study and describe these specimens, which were
collected by Texas A&M University under Contract NONR
2119(04) of the Office of Naval Research. I am deeply in-
debted to Dr. Raymond Manning, Chairman of the Depart-
ment of Invertebrate Zoology, for granting me office space and
the privileges of the laboratory in the Division of Crustacea,
and to all the members of the staff of the Division of Crustacea
for innumerable kindnesses to me during my months at the
U. S. National Museum. I also thank Dr. Kornicker and Dr.
T. E. Bowman for criticizing the manuscript and Dr. Fenner
Chace for advice on etymology. This work was supported by
grant GB-6879 from the National Science Foundation.
THE CLASSIFICATION OF THE HALOCYPRIDIDAE
Poulsen (1962) agreed with Skogsberg (1920) in dividing Sars’
Myodocopa into two suborders, the Cypridiniformes and the Halocypri-
formes, the latter of which includes the truly pelagic marine forms. The
suborder Halocypriformes contains but one family, the Halocyprididae,
and this includes two subfamilies, the Thaumatocyprinae and the Con-
choeciinae. Hitherto, the subfamily Conchoeciinae has included four
genera: Conchoecia Dana, Halocypris Dana, Archiconchoecia Miller,
and Euconchoecia Miiller. Since the new species here described are
apparently living at considerable depths, probably not far above the
bottom, and at least within the depth ranges of the bottom fish that ate
them, the name Bathyconchoecia is given to the new genus to which
they belong. Two previously described species assigned to the genus
Euconchoecia, E. lacunosa Miiller and E. darcythompsoni Scott, also
belong in this new genus.
Miller's (1912) key to the genera of Conchoeciinae is amended as
follows to include the new genus:
Key TO THE GENERA OF THE CONCHOECUNAE
1. The two terminal, often fused, segments of the first antenna with
5 setae: or sensory: filaments, 2002.0 ie eae 2
New Genus of Ostracod 541
1. These segments with more than 5 setae or filaments 3
2. Basal segment of endopodite of second antenna with a tubercle
(processus mammillaris) on its anterior margin Conchoecia
2. This segment without a processus mammillaris Halocypris
3. Terminal segments of first antenna with 6 similar sensory fila-
TTTETA ES eae =A eet rash Sede a nemMMa NEON CER AULA ER AYA anaes Ma Archiconchoecia
3. These segments with 20 or more setae and sensory filaments ____ 4
4. Terminal segments of first antenna with 20-40 setae and filaments;
LUT CAVE Walt ij aiiee lays) emmeete: seu baennererual IE MeL YIN er Ne Euconchoecia
4. These segments with more than 100 setae and filaments; furca
TAL aticto ts CO RED ATG) se See NEAL RO CN Bathyconchoecia
Bathyconchoecia new genus
Shell distinctively sculptured, especially in smaller species, pattern
of tiny pits characteristic; dorsal concavity approximately at center of
hinge line, rostrum and rostral incisure well developed. Shell glands
symmetrical, at or near postero-dorsal corners. Frontal organ lacking or
reduced to a peg. First antenna of 4-5 segments (segmentation may
not be clear), 4th segment ventrally with large oval cluster of over
200 sensory filaments and distally with an unusually large plumose seta;
male 5th segment with principal seta and 3 filaments, female’s with
principal seta and 2 filaments. Basal segment of endopodite of second
antenna with 2 bristles distally, 1 exceptionally long; last endopodite
segment with 2 setae and 3 filaments, and in male also a hook-like
clasping organ on right endopodite, clasping organ of left endopodite
smaller or lacking. Mandible characteristically lacks protruding distal
articular process of coxa; 3 plumose setae at distal end of basis. Sixth
limbs not markedly sexually dimorphic. Furca with 8 long weak claws
and an exceptionally long unpaired bristle behind the claws; Ist claw
jointed or with indications of jointing.
Type-species: Bathyconchoecia paulula new species.
Etymology: From Greek “bathys” = deep, plus Conchoecia, meaning
deep-living Conchoecia. Feminine gender.
Shell: The members of this genus are characterized by the sculpturing
of the shell, which is especially distinctive in the 3 smaller species. All
have a pattern of tiny pits over the shell, although traces of this were
hard to find on 2 of the specimens. Aside from this, the smaller species
are strikingly sculptured with reticulations and cross-striations, such as
were described by Miller (1908) for B. lacunosa. Scott (1909) said
that the shell was ornamented with faint delicate reticulations in both
sexes of B. darcythompsoni. Although the shape of the shell varies, all
the species have well developed rostrums, the rostrum usually protrud-
ing well beyond the anterior margin of the shell, and a deep rostral
incisure. Also, all the species have a dorsal concavity at approximately
the middle of the hinge line. This was made especially evident by the
transparency of the shells, but may not be so noticeable in intact freshly
542 Proceedings of the Biological Society of Washington
caught individuals. The asymmetrical glands are situated symmetrically,
as in Euconchoecia, at or near the postero-dorsal corners.
Frontal organ: No well developed frontal organ has been found, but
in some of the specimens a small peg, situated near the base of the
first antenna, may represent the remnant of the frontal organ. Con-
cerning the frontal organ of B. lacunosa, Miiller wrote: “Frontalorgan
des 2 auf einen kurzen Zapfen reduziert (?).” Scott made no mention
of a frontal organ in B. darcythompsoni. All other Conchoeciinae have
frontal organs, although these are most highly developed in the genus
Conchoecia. In Euconchoecia and Archiconchoecia the frontal organ is
an undifferentiated rod which extends just beyond the tip of the first
antenna, whereas in Halocypris it is 2-segmented. The single species,
Thaumatocypris echinata, of the subfamily Thaumatocyprinae lacks a
frontal organ.
First antenna: This consists of 5 segments in the males and probably
also in the females. The 4th segment bears distally a large plumose
seta and ventrally the large cluster of sensory filaments. The species
here described had at least 250 such filaments arranged like the facets
of a compound eye, in 10 or more rows of 25 or more filaments per row.
Viewed dorsally or ventrally this cluster is oval in outline, and from
each “facet” a sensory filament is produced. These filaments project
out stiffly for a short distance, then become very weak and flabby and
so transparent that it is virtually impossible to determine their length,
unless they are clumped together. In the male the 5th segment produces
the principal seta and 3 filaments; in the single obviously mature female
it was possible to distinguish only the principal seta and 2 filaments
on the last segment. In the genus Euconchoecia males and females have
a cluster of 20-24 filaments on the 4th segment, but these are each
borne individually and apparently arranged in 3 rows of 7 or so each.
Second antenna: As in Halocypris, Euconchoecia, and Archiconchoecia,
there is no processus mammillaris on the basal segment of the endopodite,
which bears on its dorsal distal margin 2 spines or setae, one of which is
usually quite long. In the female the 2nd endopodite segment bears 2
setae and 3 filaments. Mature females of Euconchoecia have only 2
setae and 1 filament on the 2nd endopodite segment. In males of
Bathyconchoecia the 2nd segment of the endopodite of the right
second antenna has 2 setae and a hook-like clasping organ, and from
the proximal portion of the clasping organ 3 filaments are produced.
The 2nd segment usually has 2 short fat spines beneath the curve of
the clasping organ. The left endopodite may lack the hook of a clasping
organ, but from its basal section 3 filaments are similarly produced.
Mandible: This is very similar in the 4 smaller species, but in the
largest species, B. foveolata n. sp., the last segment of the endopodite is
exceptionally small and the incisor edges of the basis and coxa are
different from those of the other 5 species. In B. sagittarius n. sp. the
incisor edge of the coxa is different in structure from that of the other
New Genus of Ostracod 543
species, but the basis and endopodite segments are similar to those of
the smaller species. In general, the incisor surface of the basis and
the endopodite of the mandible are fairly similar in structure to those
of the other genera of Concnoeciinae. These species of Bathyconchoecia,
however, have 3 plumose setae on the dorsal distal part of the basis in
the location where, according to Iles (1961, Fig. 5), in Conchoecia a
single seta on a papilla represents the remnant of the exopodite.
Halocypris globosa, Euconchoecia chierchiae, and Archiconchoecia striata
also have a single seta in this location. The distal articular process of
the coxa, which occurs in other genera of Conchoeciinae, is lacking in
all but one (B. sagittarius n. sp.) of these species of Bathyconchoecia.
Maxilla: on the 2nd endopodite segment there are 4 setae between
the 2 thicker claw-setae. Euconchoecia chierchiae and E. aculeata also
have 4 setae, but in Conchoecia, Halocypris, and Archiconchoecia striata
there are 3 setae in this location.
Fifth limb: This is fairly similar throughout the Conchoeciinae. These
species of Bathyconchoecia, except B. foveolata n. sp., have an ex-
ceptionally long seta arising from a papilla at the dorsal distal end of
the Ist exopodite segment.
Sixth limb: This is clearly sexually dimorphic in Conchoecia and
Euconchoecia, whereas in Halocypris globosa and Archiconchoecia striata
the sixth limbs of males and females are fairly similar. For Bathycon-
choecia the data are insufficient, but there is no marked sexual dimor-
phism. In males of Conchoecia and Euconchoecia the 3 setae on the last
exopodite segment are exceptionally long, as long or longer than the
limb itself, and plumose. These setae were lost from the sixth limb of 2
of the Bathyconchoeécia males, but in the B. paulula n. sp. and B.
foveolata n. sp. males, and also in male B. darcythompsoni (Scott), 1
seta is exceptionally long and the other 2 considerably shorter. In the
B. kornickeri n. sp. male all 3 setae are long, 1 somewhat shorter than
the other 2. Males and females may have an exceptionally long seta
arising from a papilla at the dorsal distal end of the Ist exopodite
segment.
Seventh limb: This is not distinctive in the Conchoeciinae. Except
in one instance (the B. kornickeri n. sp. male, which had both setae of
the same length) 1 of the 2 long setae of the last segment is longer
than the other.
Furca: This is distinctive and differs from that of other Conchoeciinae
in that the 8 pairs of claws are exceptionally long, slim and weak. In
other genera the last pairs of claws are quite small, but in Bathycon-
choecia these claws may be more like setae and not much shorter than
the first several pairs. In the larger species, the 1st claw is 5 or 6-jointed.
Scott’s (1909, Pl. IV, Fig. 11) figure of the furca of B. darcythompsoni
shows the Ist claw 5-jointed. Indications of jointing of the 1st claw are
found in the smaller species. Of the other genera of Conchoeciinae,
indications of jointing are seen only in Euconchoecia. Behind the claws
544 Proceedings of the Biological Society of Washington
Fic. 1. Male Bathyconchoecia paulula n. sp. a-—e, Lateral, anterior,
posterior, ventral and dorsal views of shell. f, First antenna. g, Second
segment of endopodite of left second antenna (setae cut off). h, Penis.
i, Endopodite of right second antenna (setae cut off). Scale on i for g
and i; at left margin for a—e; at right margin for f and h. Scales in mm.
New Genus of Ostracod 545
there is a single exceptionally long unpaired bristle. Such a bristle may
occur in the other genera, but it is usually very short. In Euconchoecia
there are only 7 claws on the furca, but there is a rounded bump be-
tween the Ist and 2nd claws similar to the process with which a claw
articulates.
All members of the suborder Halocypriformes lack eyes.
The genus Bathyconchoecia at present includes 8 species, ranging
in length of mature specimens from 0.93-5.2 mm. Of these the 3 smaller
species here described, B. paulula n. sp., B. laqueata n. sp., and B.
kornickeri n. sp., 0.93-1.66 mm long, form a natural group, being fairly
similar in shape and in the sculpturing of the shell and character of
the appendages. In all probability B. lacunosa (Miller) belongs in
this group. Miiller (1908) described this species from a 1.6 mm female,
possibly a juvenile, collected at the Gaussstation at 66°2’9”S, 89°38’E.
Both B. galerita n. sp. and B. sagittarius n. sp., here described from a
1.9 mm female and a 3.1 mm male, respectively, differ from all the
others in some respects. The two largest species, B. foveolata n. sp., here
described from a 5.2 mm male, and B. darcythompsoni (Scott), 4.7 mm
long, may possibly be grouped together. Scott (1909) described the
latter species from 2 males, a female, and 2 immature males collected
from a depth of 1140 m at 59°36’N, 7’W. The distribution of the
genus therefore extends from almost 60°N in the Atlantic to the Gulf of
Mexico and to 69°S in the Antarctic.
Of the group of smaller species, the most complete material is avail-
able for the smallest, since 2 mature males and a female, probably a
stage V juvenile, were taken. This species, B. paulula n. sp. is there-
fore designated as the type species.
A key to the species of Bathyconchoecia follows.
KEY TO THE SPECIES OF THE GENus Bathyconchoecia
1. Length of mature specimens greater than 2.5 mm _—__--.-____- 2
1. Length of mature specimens less than 2.5 mm ___. 4
2. Length of mature specimens around 3 mm _.....-
2aakength of matureyspecimens-over 4.5 mm) 2 eee 3
3. Male rostrum bifid, postero-dorsal corners of shell with a point
OTESPITIE |e eae eae eee ALE a I B. darcythompsoni (Scott)
3. Male rostrum not bifid, postero-dorsal corners rounded _.._-
a bananas avian pea ION eens ne SY CAN AN TANION Ge) et TORRE 1 Ree B. foveolata n. sp.
4. Length of mature specimens less than 1.55 mm ___. 5
4. Length of mature specimens greater than 1.5 mm —_... 6
5. Mature specimens 1 mm long or less, height of shell around 70%
of length, postero-dorsal corners bluntly rounded _.. B. paulula n. sp.
5. Mature specimens over 1.2 mm long, height 55-60% of length,
postero-dorsal corners of shell sharply right-angled —_____.
ePaper UBL PI nL te RINNE cleke aN ae ee B. laqueata n. sp.
546 Proceedings of the Biological Society of Washington
66M, GG
COGi ny,
aa g
pines ca
Fic. 2. Male Bathyconchoecia paulula n. sp. a, Coxa of mandible.
b, Basis and endopodite of mandible. c, Furca. d, Basis and endopodite
of maxilla. e, Sixth limb. f, Fifth limb. g, Pattern of sculpturing above
ventral margin; arrow points anteriorly. Scale at upper right for c and d;
at bottom for a, b, e, f, g. Scales in mm.
6. Shell strikingly sculptured with reticulations forming polygonal
cells): filled wath) taxyy, pits ewes i ea Sa i
6. Shell not strikingly sculptured though punctate pattern may be
OTE SE Mn typ 2 a NN IC A ea B. galerita n. sp.
New Genus of Ostracod 547
7. Flange of sculpturing on posterior margin beneath shell glands
EOSIN IE Nid ees SSH RAMA SAA ON Oa SOIREE B. kornickeri n. sp.
7. No flange of sculpturing on posterior margin _._ B. lacunosa (Miiller )
Bathyeconchoecia paulula new species
Figures 1-3
Holotype: Mature male, 0.95 mm long by 0.7 mm high (Figs. la-e,
g-i, 3c). Two slides. USNM 123209.
Paratypes: Mature male, 0.93 mm long by 0.7 mm high (Figs. lf,
2a-g). Two slides. USNM 123210. Female, possibly a stage V juvenile,
0.85 mm long by 0.6 mm high (Fig. 3a, b, d-l1). Two slides. USNM
123211.
Type-locality: Both males were found in the intestine of Dicromita
agassizii, a bottom fish collected at 1000 m depth in a midwater trawl
(hit bottom) at 28°15’N, 87°02’W on 11 July 1966. The female was
taken from the intestine of Nezumia hildebrandi, a bottom fish also
collected at 1000 m depth in a midwater trawl at the same location
and on the same date.
Etymology: The specific name “paulula” is the diminutive form of
the Latin “paulus” = little, and refers to the small size of this species.
Description of male (Figs. 1, 2, 3c):
Shell (Fig. la-e): Height around 70% of length, shoulder vaults
smoothly rounded, antero-ventral and postero-ventral corners rounded,
postero-dorsal corners bluntly rounded, rostrum bent sharply downwards
to about half the shell height and pointed at the tip, dorsal margin
with a concavity at the hinge line which starts at approximately half
the shell length and extends for nearly a quarter of the length. Shell
with striking sculpture, reticulations and cross-striations forming a
pattern of polygonal cells filled with tiny pits (Fig. 2g). These striations
occur over the whole surface but are especially visible along the anterior,
ventral and posterior margins and on and underneath the rostrum. The
shell glands are at the postero-dorsal corners, the openings obscured by
the sculpturing.
First antenna (Fig. 1f): This consists of 5 segments. The 4th segment
bears distally the large feathered seta and ventrally the oval cluster, 0.1
mm long, of sensory filaments which in this species appear to number
250-300, arranged in 10-12 rows of approximately 25 per row. The
tiny 5th segment bears the principal seta and the 3 shorter setae, which
become weak and flabby filaments like those of the cluster.
Second antenna (Figs. lg, i, 3c): This species has an exceptionally
long basal segment to the exopodite, which is 71-72% of the length of
the shaft (Fig. 3c). The length of the basal segment of the endopodite
is approximately 29% that of the shaft. The basal segment bears
distally 2 bristles, one of which, as is the case in this genus, is excep-
tionally long (Fig. li). Of the 2 terminal setae on the 2nd endopodite
segment, the shorter is more than half as long as the longer (Fig. 3c). In
548 Proceedings of the Biological Society of Washington
Fic. 3. Bathyconchoecia paulula n. sp. a, Lateral view of female
shell. b, Female first antenna. c, Male right second antenna. d, Basis
of female mandible. e, Endopodite of female left second antenna. f, Fe-
male fifth limb. g, Endopodite of female maxilla. h, Coxal endites of
female maxilla. i and j, Coxa of female mandible. k, Portion of female
coxal endite of maxilla. 1, Last segment of female seventh limb. Scale
at bottom left for a; on c for c; on e for e and k; at upper left for b, d, f,
h, i, j, 1. Scales in mm.
New Genus of Ostracod 549
the genus Euconchoecia the longer of these 2 setae is exceptionally long,
longer than the length of the shell. The right clasping organ (Fig. 1i)
is slim and curves evenly beyond the basal section which bears the
filaments. Only 2 filaments were found on both endopodites in this
species, but the 3rd may possibly have broken off or been obscured by
the 2 long setae. Two tiny spinules are present on the 2nd segment,
beneath the curve’ of the right clasping organ. The left clasping organ
has a small hook (Fig. 1g); aside from the 2 tiny spinules on the left
2nd endopodite segment, this species has an extra protrusion like a
short fat curving spine, and this structure has not been found in the
other species. In my experience only Euconchoecia males have a similar
but more rounded protrusion in this location.
Mandible (Fig. 2a, b): The structure of the mandible is very similar
in the 4 smaller species of Bathyconchoecia. All lack the protruding
distal articular process of the coxa which is found in other Conchoeciinae.
Also the longest setae on the 3rd endopodite segment are exceptionally
long in these species. Of the 2 longest setae on this segment, the longer
is around 0.35 mm long and the other approximately 0.25 mm long in
the B. paulula male.
Maxilla (Fig. 2d): The longer claw on the 2nd endopodite segment
is unusually long and slim. On the Ist endopodite segment there is a
group of 4 setae distally, as in all the other species here described
except B. sagittarius. This species differs from the other 3 smaller
species in having a group of 4 setae near the basis, instead of 3 as in
those species. Also, 3 of the large bristles of the coxal endite are bent
characteristically sideways, as is illustrated for the female in Figure 3k.
Fifth limb (Fig. 2f): The last exopodite segment has 2 claw setae
of unequal length and a shorter weaker seta. The lst exopodite segment
has one exceptionally long seta, which arises from a tubercle and extends
beyond the tips of the claw-setae. There are 2 claws on the endopodite.
The epipodial appendage has 3 groups of 4 plumose setae each.
Sixth limb (Fig. 2e): The last exopodite segment has one very long
claw-seta and 2 shorter setae of equal length. The Ist segment has
dorsally a very long seta arising from a tubercle. The epipodial ap-
pendage has 3 groups of setae arranged in a pattern of 6, 5, 5 setae.
The seventh limb was missing.
Penis (Fig. lh): This is relatively short and bluntly rounded at the
tip.
Furca (Fig. 2c): This has 8 pairs of long slim weak claws and an
exceptionally long unpaired bristle behind the claws. The furcal
lamella is covered with fine hairs from above the 3rd claw downwards.
The function of the oval area behind the 2nd claw is not known. The
last several pairs of claws are exceptionally long and slim. There are
indications of jointing on the Ist claw, but the joints are not separated.
Description of female (Fig. 3a, b, d-l):
Shell (Fig. 3a): Height 70 percent of length, similar in shape to that
550 Proceedings of the Biological Society of Washington
* 7
LETT
Fic. 4. Male Bathyconchoecia laqueata n. sp. a—d, Lateral, dorsal,
anterior and ventral views of shell. e, First antenna. f, Fifth limb.
g, Penis. h, Sixth limb lacking the 4th exopodite segment. Scale at top
center for a—d; at left center for e—h. Scales in mm.
of the male, but with the maximum height in the anterior half. The
shell of this specimen was stiff and brittle, not flexible like those of the
other specimens, the entire surface covered with tiny pits. If striations
had been present, they were lost. This specimen, 0.85 mm long by 0.6
mm high, is probably a stage V juvenile.
New Genus of Ostracod dol
First antenna (Fig. 3b): This consists probably of 5 segments, the
4th bearing distally the large plumose seta and ventrally the cluster
of sensory filaments. These filaments are arranged in 10-12 rows of
around 25 per row, so 250-300 filaments are present, as in the male.
The tiny last segment bears the principal seta; if 2 weaker and smaller
setae were present, they were obscured by the large cluster of filaments.
Second antenna: This has the same relative proportions as that of the
male illustrated in Figure 3c, the basal segment of the exopodite being
three-quarters as long as the shaft. The basal segment of the endopodite
is 30% the length of the shaft, and bears distally 2 bristles, both of
which were broken (Fig. 3e) on this specimen. The 2nd endopodite
segment has only the 2 distal setae, both of which were broken, and 1
filament. This specimen resembles the females of Euconchoecia in
having only 1 filament, but in this case it may indicate that this female
is a stage V juvenile, since female B. lacunosa (Miller) and B. galerita
n. sp. have 2 setae and 3 filaments on the 2nd endopodite segment.
Mandible (Fig. 3d, i, j): This is similar to that of the male.
Maxilla (Fig. 3g, h, k): This is also similar to that of the male; the
larger claw of the 2nd endopodite segment is also exceptionally long,
and 3 of the bristles of the coxal endite are bent characteristically side-
ways.
Fifth limb (Fig. 3f): As in the male, the last segment of the exopo-
dite has 2 claw-setae and a smaller seta, and the Ist segment has an
exceptionally long seta arising from a tubercle, which extends well beyond
the tips of the claw-setae. The endopodite has only 1 strong claw,
possibly a juvenile character.
Sixth limb: This appears to be similar to that of the male, except that
the long seta arising from a papilla on the 1st exopodite segment was
much longer, at least twice as long as the limb itself.
Seventh limb (Fig. 31): One of the 2 long setae of the last segment
is a little shorter than the other.
The furca is similar to that of the male (Fig. 2c).
Remarks: B. paulula n. sp. differs from all the other species in its
small size, in that the height of the shell is at least 70% of the length,
and in that the basal segment of the exopodite of the second antenna
is exceptionally long, approximately three-quarters the length of the
shaft.
Bathyconchoecia laqueata new species
Figures 4-5
Holotype: Male, 1.3 mm long by 0.75 mm high (Figs. 4, 5a—c, e, h).
Two slides. USNM 123212.
Paratype: Male, 1.28 mm long (Fig. 5d, f, g). Two slides. USNM
123213.
Type-locality: Both males were found in the intestine of Dicromita
agassizii, a bottom fish taken in a midwater trawl at 1000 m depth (hit
bottom) at 28°15’N, 87°02’W on 11 July 1966.
552 Proceedings of the Biological Society of Washington
Fic. 5. Male Bathyconchoecia laqueata n. sp. a, Endopodite of right
second antenna. b, Basis and endopodite of maxilla. c, Coxa of mandible.
d, Precoxal and coxal endites of maxilla. e, Second segment of endopodite
of left second antenna (setae and filaments cut off). f, Right clasping
organ (filaments cut off). g, Basis and endopodite of mandible. h, Molar
and incisor surfaces of coxa of mandible. Scale at bottom left for a,
c, g, h; at upper center for b, d; at right center for e, f. Scales in mm.
New Genus of Ostracod 503
Etymology: The specific name is derived from the Latin “laqueatus” =
fretted, paneled, pitted, and refers to the sculpturing of the shell of
this species.
Description of male: Shell (Fig. 4a—d): Height 57-58 percent of
length, shoulder vaults smoothly rounded, antero-ventral and _ postero-
ventral corners rounded, dorsal margin with a concavity at the hinge
line which starts just behind the midline, postero-dorsal corners sharply
right-angled with shell glands opening on the dorsal side. The shell
is strikingly sculptured, with striations forming irregular polygonal cells
filled with tiny pits, as is indicated in Figure 4a-d. These striations
are particularly noticeable on the rostrum and the anterior, ventral and
posterior margins. The underside of the rostrum is also similarly sculp-
tured. The sculpturing is similar to that of B. paulula n. sp. and B.
kornickeri n. sp., and also to that described for B. lacunosa (Miiller).
First antenna (Fig. 4e): This consists of 5 segments, the first 2
relatively short. It was not possible to distinguish any remnant of a
frontal organ from the pointed thickenings of the exoskeleton. There are
some fine hairs on the anterior margin of the 3rd segment or the posterior
margin of the 4th. The cluster of sensory filaments on the 4th segment
is slightly over 0.1 mm long and consists of 10-12 rows of 26-28
filaments each, so approximately 250-350 filaments are present. The
principal seta is bare of spinules and approximately 0.7 mm long. Figure
4e shows the relative lengths of the other setae and filaments.
Second antenna (Fig. 5a, e, £): The basal segment of the exopodite
is rather long, around 65 percent of the length of the shaft. The long
exopodite setae are approximately 0.6 mm long. The basal segment
of the endopodite, which is 38—40 percent of the length of the shaft, has
a more lumpy shape than that of the other species. It bears distally 2
setae, the longer approximately twice the length of the shorter one.
The right clasping organ is slim and strongly curved and bears, from
its proximal section, 3 filaments whose relative lengths are shown in
Figure 5a. The second segment of the endopodite has 2 short fat spines
beneath the curve of the clasping organ, and bears 2 long setae, the
longest of which was 0.8 mm long. The left clasping organ (Fig. 5e)
has a small hook.
Mandible (Fig. 5c, g, h): The longest seta of the 3rd segment of
the endopodite is exceptionally long (0.32 mm) in this species, as it is
in the other smaller species. The incisor and molar surfaces of the
coxa are also similar to those of the other smaller species.
Maxilla (Fig. 5b, d): The basal segment of the endopodite has a
cluster of 3 setae near the basis, as in B. galerita n. sp. and B. kornickeri
n. sp. The maxilla most closely resembles that of the latter species,
except that 2 bristles of the precoxal endite that are quite small in
B. laqueata are long and slim in B. kornickeri, as well as in B. galerita.
Fifth limb (Fig. 4f): The endopodite has 2 claws. One of the 2
claw-setae on the 3rd exopodite segment is longer, the shorter one
504 Proceedings of the Biological Society of Washington
Fic. 6. Male Bathyconchoecia kornickeri n. sp. a—b, Lateral and
ventral views of shell. c, Rostrum and portion of anterior margin of
shell. d, First antenna. e, Endopodite of left second antenna. f, Endopo-
dite of right second antenna (setae cut off). g, Endopodite of left
second antenna (setae cut off). h, Sixth limb. Scale at upper left for
a and b; at top center for c, d, e, h; at lower center for f, g. Scales in mm.
New Genus of Ostracod 500
similar in length to the seta of this segment. The Ist exopodite segment
has an exceptionally long seta arising from a papilla.
Sixth limb (Fig. 4h): The last segment was broken off. This limb
differs from that of the other smaller species in that the seta arising
from a tubercle on the Ist segment is much shorter.
Penis (Fig. 4g): This is fairly similar in shape to that of the B.
paulula male.
The seventh limb and furca were lost. ©
The female is not known.
Remarks: This species differs from the others in that the postero-
dorsal corners of the shell are sharply right-angled, with the shell glands
opening on the dorsal side.
Bathyconchoecia kornickeri new species
Figures 6—7
Holotype: Male, 1.66 mm long by 0.92 mm high. One slide. USNM
123214,
Type-locality: Intestine of Dicromita agassizii, a bottom fish taken
in a midwater trawl at 1000 m depth (hit bottom), at 28°15’N, 87°02’W
on July 11, 1966.
This species is named for Dr. Louis S. Kornicker.
Description of male: Shell (Fig. 6a—c). Height 55-56 percent of
length, antero-ventral and postero-ventral corners rounded, postero-
dorsal corner definite, dorsal margin straight in lateral view but with a
concavity at the hinge line at approximately half the shell length,
rostrum pointed and curving sharply downward. The sculpturing of the
shell is striking, and is similar to that of B. paulula n. sp. and B. laqueata
n. sp. The entire surface of the shell is punctate, and especially on and
beneath the rostrum and on the anterior margins striations form an
irregular pattern of polygonal cells, each with the small pits arranged
inside (Fig. 6c). Just below the postero-dorsal corner and below the
symmetrical shell glands the sculpturing is prolonged in a flange for a
short distance on the posterior margin.
First antenna (Fig. 6d): This consists of 5 segments, although the
basal segment is quite short and not clearly delimited from the 2nd
segment. No remnant of a frontal organ was found. The 4th segment
bears distally the plumose seta and ventrally the large oval cluster of
sensory filaments, of which there are around 250-300 in all, arranged
in 10-12 rows of 24—25 filaments per row, as in B. paulula n. sp. The
exact length of the fine filaments could not be determined, but they
were probably considerably longer than is shown in Figure 6d. The
5th segment bears the principal seta and the 3 shorter setae, the relative
lengths of which are illustrated in Figure 6d.
Second antenna (Fig. 6e, f, g): The length of the basal segment of
the exopodite is around 63 percent that of the shaft; the long exopodite
bristles are 0.75-0.8 mm long. The basal segment of the endopodite is
556 Proceedings of the Biological Society of Washington
———
Fic. 7. Male Bathyconchoecia kornickeri n. sp. a, Coxa of mandible.
b, Basis and endopodite of mandible. c, Basis and endopodite of maxilla.
d, Coxal and precoxal endites of the maxilla. e, Last segment of seventh
limb. Scale at left center for a—-e. Scale in mm.
33-34 percent the length of the shaft. The relative lengths of the setae
and filaments of the left endopodite are shown in Figure 6e. The
longer seta of the 2nd segment is more than twice as long as the shorter
seta; though broken near the tip it measured 1.45 mm in length. The
2nd endopodite segment also has 2 short fat spinules as in the other
species. The right clasping organ is strong and sharply curved (Fig.
6f) and bears 3 filaments. On the left endopodite (Fig. 6g) the 3
filaments were produced from the basal portion of the clasping organ;
no hook was present, but it may have been broken off.
New Genus of Ostracod 507
Mandible (Fig. 7a, b): This is similar to that of the other smaller
species.
Maxilla (Fig. 7c, d): This resembles most closely that of B. laqueata
n. sp., and, as in B. laqueata and B. galerita n. sp., there is a cluster
of 3 setae on the basal segment of the endopodite near the basis. As
in the other species, except B. sagittarius n. sp., there are 4 setae on the
distal margin of the lst endopodite segment. The coxal and precoxal
endites are similar to those of the other smaller species, but have
shorter and fewer bristles than the 2 larger species, B. sagittarius n. sp.
and B. foveolata n. sp.
Sixth limb (Fig. 6h): This differs from that of the other known males
of Bathyconchoecia in that the 3 setae on the last exopodite segment
are all long, one somewhat shorter than the other 2. As in most of the
other species, the seta borne on a tubercle on the Ist segment is quite
long. The epipodial appendages of the sixth limb consist of 3 groups
of plumose setae in the pattern of 5, 5, 7 setae each. Those of the
fifth limb have groups of 4, 4, 5 setae each.
Seventh limb (Fig. 7e): This differs from that of the other species
in that the last segment has 2 long setae of the same length, approxi-
mately 0.3 mm long.
The fifth limb, penis and furca were lost.
The female is not known.
Remarks: This species is similar in size and in the sculpturing of the
shell to B. lacunosa, which Miller (1908) described from a female speci-
men, possibly a stage V juvenile, 1.6 mm long, in his report on the
ostracods of the Deutsche Siidpolar Expedition. Since this sculpturing
occurs in several species and Miiller did not mention the flange of
sculpturing on the posterior margin nor describe the appendages so
that they could be compared with those of this male, it is impossible to
determine whether these two specimens are conspecific.
Bathyconchoecia galerita new species
Figures 8-9
Holotype: Female, 1.9 mm long by 1.2 mm high. Two slides. USNM
123215.
Type-locality: Stomach of Nezumia hildebrandi, a bottom fish taken
in a midwater trawl at 1000 m depth on 11 July 1966 at 28°15/N,
87°02/W.
Etymology: The specific name is derived from the Latin “galeritus” =
wearing a hood, and refers to the hood-like shape of the rostrum of this
species.
Description of female: In the other specimens, taken from fish
intestines, all traces of soft tissues were gone, leaving only the exoskele-
tons of the appendages. This is the only specimen from a fish stomach
and much of the musculature remained, or was in process of being
dissolved.
508 Proceedings of the Biological Society of Washington
Pe =
AS
S
Clay eae
ay aw :
Fic. 8. Female Bathyconchoecia galerita n. sp. a—b, Lateral and
ventral views of shell. c, Right second antenna. d, Coxa of mandible.
e, Basis and endopodite of mandible. f, Sixth limb. Scale at upper right
for c-f; at lower right for a, b. Scales in mm.
Shell (Fig. 8a, b): Height three-fifths the length, antero-ventral and
postero-ventral corners rounded, dorsal margin nearly straight but with
a concavity at the hinge line at approximately half the shell length,
rostrum relatively large and rounded at the tip. Very faint traces of
sculpturing in a pitted scale-like pattern, as in the other species, were
found around the rostral area, so in life the shell may have been covered
with a punctate pattern. No indications of striations or other sculpturing
New Genus of Ostracod 559
Fic. 9. Female Bathyconchoecia galerita n. sp. a, First antenna.
b, Coxal and precoxal endites of maxilla. c, Basis and endopodite of
maxilla. d, Fifth limb. e, Furca. Scale at right for a; at lower left for
b-e. Scales in mm.
were found. The shell glands are at the postero-dorsal corner of each
shell. The body of the animal compactly filled the shell, and the
posterior part of the body was tightly crammed with 4 large eggs
0.30-0.33 mm in diameter.
560 Proceedings of the Biological Society of Washington
Fic. 10. Male Bathyconchoecia sagittarius n. sp. a—b, Lateral views
of right and of left shells. c, Penis. d, Furca. e, Mandible. f, Fifth
limb. Scale at upper left for a and b; at bottom left for c-f. Both
scales in mm.
First antenna (Fig. 9a): This consists of 4 to 5 segments. The first
2 (?) segments are short, and the tiny last segment which bears the
principal seta and the 2 smaller setae is not clearly separated from the
Ath segment. The anterior edge of the 3rd segment has several rows
of long hairs laterally. The 4th segment bears distally the large plumose
seta and ventrally the cluster of sensory filaments, which consisted of
New Genus of Ostracod 561
12-15 rows of 35-38 filaments each, so there were at least 400-500
filaments in the cluster.
Second antenna (Fig. 8c): The shaft is stout, and the length of the
basal segment of the exopodite is 54-55 percent that of the shaft. The
long setae of the exopodite segments are approximately 0.8 mm long.
The basal segment of the endopodite, which is around 26 percent the
length of the shaft, has distally 2 setae, the larger one well over twice
as long as the other. The last segment bears 2 long setae and 3 filaments.
Figure 8c shows the relative lengths of the setae and filaments, except
for the longest seta which was broken on both endopodites.
Mandible (Fig. 8d, e): This is very similar to that of the 3 other
smaller species.
Maxilla (Fig. 9b, c): This is distinctive, in that the claws of the
2nd endopodite segment are short and stumpy, unlike those of any other
of these species. On the Ist endopodite segment there is a cluster of
only 3 setae near the basis, as in B. laqueata n. sp. and B. kornickeri
Nn. sp.
Fifth limb (Fig. 9d): The last exopodite segment has 2 strong
curved claw-setae and a weaker seta, all of about the same length. The
Ist segment differs from that of the other smaller species in having 9
setae as well as a long seta which arises distally from a tubercle. The
endopodite has 2 claws, and the endopodite and protopodite, which
are not clearly separated, together have 9 setae.
Sixth limb (Fig. 8f): The last exopodite segment has 2 relatively
short claw-setae and a weaker seta, all of about the same length. The
lst segment has distally a very long seta borne on a tubercle.
The seventh limb was lost.
Furca (Fig. 9e): This has 8 pairs of long weak claws, which decrease
gradually in length posteriorly, and a long unpaired bristle. There is a
small circular area of unknown function near the base of the Ist claw.
The Ist claw shows indications of jointing, but is not actually separated
into joints.
The male is not known.
Remarks: This species is similar in shape to B. lacunosa (Miller),
but lacks the sculpturing described for this species. Also the relative
lengths of the filaments and setae of the endopodite of the second
antenna are different from those figured by Miller (1908, Pl. X. Fig. 6)
for B. lacunosa. B. galerita n. sp. differs from all the other species in
the lack of sculpturing on the shell and in the character of the maxilla
and the number of setae on the Ist exopodite segment of the fifth limb.
Bathyconchoecia sagittarius new species
Figures 10-11
Holotype: Male, 3.1 mm long by 1.6 mm high. Three slides. USNM
123216.
Type-locality: Intestine of Bassozetus normalis, an abyssal bottom fish
562 Proceedings of the Biological Society of Washington
Fic. 11. Male Bathyconchoecia sagittarius n. sp. a, First antenna.
b, Cross-section of fragment of cluster of filaments of the first antenna.
c, Second segment of endopodite of left second antenna (setae and
filaments cut off). d, Endopodite of right second antenna (setae and 1
filament cut off). e, Coxal and precoxal endites of the maxilla.
f, Endopodite segments of the maxilla. g, Seventh limb. Scale at top
left for a and g; at left center for b, at top right for c-f. Scales in mm.
New Genus of Ostracod 563
taken in a dredge at a depth of 3165 m on 27 July 1967 at 25°26.5’N,
86°06’W.
Etymology: The specific name is derived from the Latin noun
“sagittarius” = archer, bowman. This species is named for Dr. Thomas
E. Bowman.
Description of male: This specimen was crumpled into a ball, and
part of the posterior margin of the shell was damaged, but most of the
appendages were essentially intact.
Shell (Fig. lla, b): Height approximately half the length, dorsal
margin slightly curved, antero-ventral and _ postero-ventral corners
rounded, postero-dorsal corner of the right shell produced in a blunt
point, rostrum large, rounded, protruding well forward and curving down
to approximately half the shell height. In the condition in which it was
found, the shell is pliable, not brittle. Sculpturing covered the entire
surface, consisting primarily of fine striations running dorso-ventrally,
with some cross-striations producing polygonal or diamond-shaped cells,
particularly on and beneath the rostrum and on and near the anterior
margin. Indications of the pattern of tiny pits were also present, but it
is impossible to determine whether the entire surface of the shell had
been punctate as in most of the other species. The posterior margins of
both shells were damaged, but enough of the margins remained to show
that the entire posterior margin was edged with a single row of dentate
structures which started at the postero-dorsal corner and became smaller
ventrally, ending at the postero-ventral corner. It was not possible to
determine the location of the shell glands; presumably they are just below
the postero-dorsal corners.
First antenna (Fig. lla, b): There is a tuft of long hairs ventrally
at the distal end of the 3rd segment. The 4th segment bears distally
the large plumose seta and ventrally the cluster of sensory filaments
which are arranged in approximately 12 rows of 25-28 filaments per
row, so around 300 filaments are present. Figure 11b shows a cross-
section of a fragment of the cluster; from each facet of the cluster a
filament is produced which extends stiffly for a short distance, then
becomes very weak and flabby. The tiny 5th segment bears the
principal seta and 3 shorter setae which become flabby sensory fila-
ments. The principal seta of each first antenna was broken, so its length
is not known, but it is at least as long as is illustrated in Figure lla.
No remnant of a frontal organ was found on this specimen.
Second antenna (Fig. llc, d): The length of the basal segment of
the exopodite is 60 percent that of the shaft. The basal segment of the
endopodite of the right second antenna, which is nearly 40 percent the
length of the shaft, has distally 2 setae, both of which were broken in
this specimen, but evidently they are exceptionally long. The 2nd
endopodite segment has distally 2 long setae, the longer of which was
broken at a length of 1.9 mm. The right clasping organ (Fig. 11d)
is a large and strongly curved hook which bears at its bend 3 filaments
564 Proceedings of the Biological Society of Washington
Fic. 12. Male Bathyconchoecia foveolata n. sp. a, Lateral view of
right shell. b, Sculpturing of shell below shell gland. c, Sixth limb.
d, Mandible. e, Furca. f, Penis. g, Seventh limb. Scale below a for a;
at lower left for e and f; on b for b; at upper right for c, d, g. Scales in
mm.
of varying lengths. The 2nd segment also has 2 tiny spinules beneath
the curve of the clasping organ, and another spinule at the base of the 2
long setae. The 2nd segment of the left endopodite (Fig. llc) also
has these 3 spinules, but the base of the left clasping organ bore only
the 3 filaments, and no hook was present.
New Genus of Ostracod 565
Mandible (Fig. 10e): The basis and endopodite are similar to the
other species described here, except B. foveolata n. sp. This species
differs from all the others in having a relatively sharp, knob-like distal
articular process on the coxa, and also in the structure of the incisor and
molar surfaces of the coxa.
Maxilla (Fig. lle, f£): This species differs from all the others in
having a row of 7 setae at the distal end of the Ist endopodite segment
plus a group of 3 more setae, making 10 in all, in the location where
the other species have no more than 4 setae (compare, for example,
Fig. 2d with Fig. 11f). There are more bristles on the coxal and
precoxal endites (Fig. lle) than in any of the smaller species; these
bristles are shorter and fatter than in B. foveolata n. sp.
Fifth limb (Fig. 10f): This differs from that of all the others here
described, with the possible exception of B. foveolata n. sp., in that there
appear to be 4, instead of 3, exopodite segments. At the joint between
the Ist and 2nd segments there is an exceptionally long seta arising from
a tubercle. The last segment has 2 claw-setae, one slightly longer than
the other, and a slightly shorter seta. The 3rd segment has 13 setae,
instead of 4—5, as in the other species. The endopodite has 2 claws
and 4 curved spines. The separation between the endopodite and
protopodite is not clear, but together there are 9 setae, 5 of these plumose.
Sixth limb: This also has more setae than in the other species. The
setae were broken from the last segment, but the 3rd segment has a
total of 8 setae (other species have 2), the 2nd segment has 12 setae
(other species have 4-5), and the Ist segment has 5 setae plus an
exceptionally long seta arising from a tubercle at the distal end.
The epipodial appendages of the 5th limb have 3 groups of plumose
setae in the pattern of 5, 4, 4 setae each; those of the sixth limb have
3 groups of 7, 5, 5 setae each.
Seventh limb (Fig. 11g): This bears distally 2 long setae, one 0.6
mm long, the other nearly 0.8 mm long.
Penis (Fig. 10c): The shape is distinctive. The distal end is straight
and truncates the long axis at 45°.
Furca (Fig. 10d): This has 8 pairs of exceptionally thin weak claws,
the last 2 pairs more like setae than claws, and an unusually long un-
paired bristle behind the claws. The distance between the 6-jointed Ist
claw and the 2nd claw is exceptionally great, and an extra bump is
present in front of the 2nd claw.
The female is not known.
Remarks: This species differs from all the others in the type of
sculpturing, in the coxa of the mandible, and in the greater number of
setae or bristles on the maxilla and fifth and sixth limbs.
Bathyconchoecia foveolata new species
Figures 12-13
Holotype: Male, 5.2 mm long by 2.2 mm high. Two slides. USNM
IPRPIT
566 Proceedings of the Biological Society of Washington
Fic. 13. Male Bathyconchoecia foveolata n. sp. a, Fifth limb.
b, First antenna. c, Basis and endopodite segments of maxilla. d, Pre-
coxal and coxal endites of maxilla. e, Endopodite of right second antenna
(setae cut off). Scale at bottom right for a—e. Scale in mm.
Type-locality: Intestine of Bassozetus normalis, a bottom fish taken in
a dredge at 3165 m depth on 27 July 1967 at 25°26.5'N, 86°06’W.
Etymology: The specific name is derived from the Latin “foveola” =
little pits, and refers to the sculpturing of the shell of this species.
Description of male: Shell (Fig. 12a, b): This is a large species, the
New Genus of Ostracod 567
height of the shell two-fifths the length, antero-ventral and postero-
ventral corners rounded, ventral margin straight, dorsal margin nearly
straight with a slight concavity at the hinge line at approximately half
the shell length. The rostrum is large and rounded and projects well
beyond the anterior margin for approximately one-sixth the shell length.
The entire surface of the shell is covered with a sculpturing of tiny pits
in a small scale-like pattern, as is indicated on the rostrum in Figure
12a and shown more highly magnified in Figure 12b, which also shows
the single row of dentate structures which extends down the posterior
margin from the shell gland to just anterior to the postero-ventral corner.
The shell glands are situated symmetrically just below the postero-
dorsal corner of each shell.
First antenna (Fig. 13b): This consists of 5 segments and is similar
in structure to that of the other males here described. The basal seg-
ment has a small peg-like structure antero-distally, which may represent
a remnant of the frontal organ or be only a thickening of the integument.
At the dorsal distal margin of the 3rd segment there is a row of sharply
curved spines, and a row of hairs extends down the lateral distal margin
of this segment and continues posteriorly on the ventral surface. Some
long hairs are also present on the 4th segment above the cluster of
sensory filaments. This cluster is 0.35 mm long, and consists of 16—20
rows of approximately 35 filaments per row, making a total of 550-700
filaments in all. These filaments were tangled in debris and are un-
doubtedly considerably longer than is shown in Figure 13b. Aside from
the large plumose seta borne antero-distally on the 4th segment, the
small 5th segment has the large principal seta and 3 smaller setae. The
2 shorter setae were approximately 0.9 and 1.0 mm long; the longer
setae were broken.
Second antenna (Fig. 13e): The length of the basal segment of the
exopodite is 60 percent that of the shaft, the basal segment of the
endopodite 33.5 percent of shaft length. The endopodite of the left
second antenna was broken off, but that of the right is illustrated in
Figure 13e. Both the long setae of the 2nd segment were broken. The
right clasping organ is large and strong and curves around the 2nd
segment, 3 filaments being produced from its basal section. Two short
fat spinules are present beneath the curve of the clasping organ and 2
more spinules are situated at the base of the 2 long setae. The larger
of the 2 setae of the 2nd segment has some tiny spinules down its
length. This seta, broken near the tip, was 3.5 mm long; the shorter
seta, also broken near the tip, was 1.3 mm long. Both of the bristles
at the distal end of the basal segment were also broken.
Mandible (Fig. 12d): This differs from all the other species. The
structure of the molar and incisor surfaces of the coxa and the incisor
surface of the basis is different. Also, the last endopodite segment is
proportionately much smaller and has fewer setae than the other species.
568 Proceedings of the Biological Society of Washington
It resembles the others, however, in having a group of 3 plumose setae
at the distal end of the basis.
Maxilla (Fig. 13c, d): The endopodite is similar to that of the other
species, except B. sagittarius n. sp. The bristles of the coxal and pre-
coxal endites (Fig. 13d) are much longer and slimmer than those of
B. sagittarius n. sp. and are also more numerous than in the smaller
species.
Fifth limb (Fig. 13a): The endopodite has 2 strong claws, and is not
clearly separated from the protopodite. The basal segment of the exopo-
dite is not as definitely separated into 2 segments as in the B. sagittarius
male, so it is difficult to decide whether the exopodite has 3 or 4 seg-
ments. As in B. sagittarius, at the possible joint between the Ist and 2nd
exopodite segments a long seta is produced. The last exopodite segment
has 2 relatively short sub-equal claw-setae and a shorter seta. The
epipodial appendages of the fifth limb have 3 groups of 5, 4, 4 plumose
setae each.
Sixth limb (Fig. 12c): The exopodite has 4 segments, the last seg-
ment bearing 1 long seta and 2 shorter ones about half as long as the
longest. The Ist segment has a long seta, but this is not exceptionally
long as in some of the species. The epipodial appendage has 3 groups
of setae arranged as 7, 5, 5 setae each.
Seventh limb (Fig. 12g): This bears distally 2 long setae, the longer
1.7 mm long, the shorter 1.15 mm long.
Penis (Fig. 12 £): This is long and slim, bluntly rounded at the tip.
Furca (Fig. 12e): The first several pairs of claws are exceptionally
long, weak and slim, the last several pairs quite short, and there is an
unpaired bristle behind the claws. The Ist claw is 5-jointed, with
indications of a 6th joint. Near the base of the 2nd claw there is a
tiny circular area. The furcal lamella is covered with hairs.
The female is unknown.
Remarks: This species is in the same size range as B. darcythompsoni
(Scott). In the latter species, however, the male rostrum is bifid and
there is a small point at the postero-dorsal corner of each shell. B.
foveolata n. sp. is distinctive in the size, shape, and sculpturing of the
shell and differs from the other species primarily in the structure of
the mandible.
The two previously described species, B. lacunosa (Miller )
and B. darcythompsoni (Scott), were originally placed in the
genus Euconchoecia because the sensory filaments on the last
2 segments of the first antenna were numerous. Miiller (1908,
p. 10, Pl. X, Figs. 1-8) noted that in the shape and sculpturing
of the shell and the structure of the appendages lacunosa
differed widely from Euconchoecia chierchiae, the type species.
His description, particularly of the sculpturing of the shell
New Genus of Ostracod 569
and of the first and second antenna and mandible show that
this species belongs in the group of smaller species of Bathy-
conchoecia, together with B. paulula n. sp., B. laqueata n. sp.,
and B. kornickeri n. sp. From Scott’s (1909, p. 128, Pl. III,
Fig. 19, Pl. IV, Figs. 1-12) description and figures of darcy-
thompsoni it is evident that this species also belong to the
genus Bathyconchoecia.
The removal of these 2 species to the genus Bathyconchoecia
leaves in the genus Euconchoecia only 2 very closely related
species, E. chierchiae Miller and E. aculeata (Scott). These
species live in the upper waters of warmer seas and are
tropical-subtropical in distribution. E. chierchiae has been
recorded in the Atlantic from 42°N to 20°S, and, according to
Miiller (1906), E. aculeata from 10°N-11°S in the Indian
Ocean. E. chierchiae was described from the coast of Brazil
(Miiller, 1890), and E. aculeata from the Gulf of Guinea
(Scott, 1894). In the Sargasso Sea (Deevey, 1968) E.
chierchiae occurred seasonally over an extreme possible tem-
perature range of 16.6-27.3°C. These are small species, around
1 mm long, with delicate shells, lacking sculpturing. In the
females the body of the animal does not fill the shell, and,
as in no other known Halocyprid, large eggs are carried be-
tween the back of the body and the shell.
The species of Bathyconchoecia are apparently living at
considerable depths, either on or just above the bottom in
regions not easily sampled with plankton nets. That three
bottom fish should eat 9 specimens of 6 new species may
indicate that many more species still are to be found. The
species here described were taken at depths of 1000 and 3165
m. B. lacunosa was collected on 2 December 1902 at the
Gaussstation, 66°2’9’S, 89°38’E in the Indian Ocean sector of
the Antarctic in a qualitative sample, probably a bottom
sample, since Miller (1908) noted that the bottom net samples
obviously caught pelagic species while being pulled up. B.
darcythompsoni was collected in August 1907 from a depth of
1140 m in the Atlantic north of Scotland. In the shape of the
shell, the presence of a well-developed rostrum and in the
character of the furca these forms are immediately recogniz-
570 Proceedings of the Biological Society of Washington
able as belonging to the family Halocyprididae; however,
these are pelagic species that, particularly in the extraordinary
development of the large cluster of sensory filaments on the
4th segment of the first antenna, have become specialized
for living in the depths of the sea.
LITERATURE CITED ~
Deevey, G. B. 1968. Pelagic ostracods of the Sargasso Sea off
Bermuda. Peabody Mus. Nat. Hist. Yale Univ., Bull. 26., pp.
1-125, 65 Figs.
Ines, E. J. 1961. The appendages of the Halocyprididae. Discovery
Rep. Vol. XXXI, pp. 229-326.
Kornicker, L. S. In Press. Bathyal myodocopid Ostracoda from the
northeastern Gulf of Mexico. Proc. Biol. Soc. Wash. Vol. 81,
no. 45, pp. 439-472.
Muuxer, G. W. 1890. Ueber Halocypriden. Zool. Jahrb. Abt. Syst.,
Geog. Biol. Vol. V (2), pp. 253-280, Pls. 28-29.
. 1906. Ostracoda. Wissensch. Ergeb. d. Deutschen Tiefsee-
Expedition auf dem Dampfer “Valdivia” 1898-1899. Vol.-8,
pp. 1-154, Pls. V-XXXV.
1908. Die Ostracoden der Deutschen Siidpolar-Expedition
1901-1903. Deutsche Siidpolar-Expedition 1901-1903, Vol.
X, Zoologie II, pp. 53-181, Pls. IV—XIX.
1912. Ostracoda. In Das Tierreich, Lief. 31, pp. 1-434.
Berlin.
Poutsen, E. M. 1962. Ostracoda-Myodocopa Part I Cypridiniformes-
Cypridinidae. Dana-Report 57, 414 pp., 181 Figs.
Scotr, T. 1894. Report on Entomostraca from the Gulf of Guinea,
collected by John Rattray, B.Sc. Linn. Soc. London, Trans.,
Ser. 2, Vol. VI, Zool., pp. 1-161, Pls. 1-15.
1909. On some new and rare Entomostraca from the Scottish
Seas. Ann. Mag. Hist., Vol. III, Eighth Series, pp. 122-130,
Pls. I-IV.
SxocsBERG, T. 1920. Studies on marine ostracods Part I (Cyprininids,
Halocyprids and Polycopids). Zool. Bidr. Uppsala, Suppl.
I, 784 pp., 153 Figs.
Of —
Vol. 81, pp. 571-586 30 December 1968
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
NOTES ON THE HARPACTICOID COPEPODS
ATTHEYELLA PILOSA AND A. CAROLINENSIS,
ASSOCIATES OF CRAYFISHES IN THE
EASTERN UNITED STATES
By THomas E. BowMAN, RUDOLPH PRINS, AND Byron F’. Morris
Division of Crustacea, Smithsonian Institution; Department of
Biology, Western Kentucky University, Bowling Green,
Kentucky; Biology Department, Dalhousie University,
Halifax, Nova Scotia
Among the North American freshwater copepods described
by Chappuis are 2 species of the harpacticoid genus Attheyella.
A. pilosa was described from Mammoth Cave, Kentucky, and
Donnaldson Cave, Indiana (Chappuis, 1929a); A. carolinensis
was described from Chapel Hill, North Carolina (Chappuis,
1932). An amplified description of A. carolinensis from the
type locality was given by Coker (1934), and the species was
later reported from Mountain Lake, Virginia (Carter, 1944)
and, erroneously, from Doe Run, Kentucky (Prins, 1964).
The only published report of A. pilosa since its discovery is a
record of a single female from Luchil Cave, Yucatan (C. B.
Wilson, 1936). The Yucatan specimen is not in the collections
of the Smithsonian Institution, and a record so distant from the
type-locality cannot be accepted without confirmation.
Reexamination of the Doe Run specimens, which Prins
found on crayfishes, showed them to be A. pilosa rather than
A. carolinensis. In an attempt to shed some light on the distri-
bution of the 2 species and on the life history of A. pilosa we
have examined harpacticoids found in association with cray-
fishes from a number of localities, mostly in the southeastern
United States, including 22 samples from Doe Run. Since
A. pilosa and A. carolinensis are similar in many respects and
53—Proc. Biot. Soc. WasH., Vou. 81, 1968 (571)
572 Proceedings of the Biological Society of Washington
since the original descriptions do not point out adequately the
differences between the 2 species, we take this opportunity to
redescribe and contrast their diagnostic characters.
Attheyella pilosa Chappuis
Figs. 1, 2, 3a4j
Attheyella (Brehmiella) pilosa Chappuis, 1929a, pp. 53-55, figs. 5-11;
1929b, p. 488 [in list]; p. 507 [in key]; 1931, pp. 348-351, figs. 5-11
[reprint of original description and figs.]; 1933, p. 19 [in list]—Lang,
1948, p. 984, fig. 3 [copied from Chappuis].
Attheyella (Ryloviella) pilosa Chappuis——Borutsky, 1952, pp. 282-283
[p. 260 in 1964 translation], fig. 79: 4-6 [copied from Chappuis].
Attheyella pilosa Chappuis.—Pennak, 1953, p. 403 [listed in footnote].
—M. S. Wilson, 1956, p. [in list]—Barr, 1968, p. 159.
Attheyella (Mrazekiella) pilosa Chappuis—M. S. Wilson and Yeatman,
1959, p. 847, fig. 29.201 [copied from Chappuis].
[?] Attheyella pilosa Chappuis.—C. B. Wilson, p. 79.
Attheyella carolinensis Chappuis.—Prins, 1964, pp. 370-371.
Female body length without caudal setae usually 0.58-0.70 mm
(extremes measured: 0.53 and 0.76 mm). Male body length usually
0.52-0.64 mm (extremes measured 0.46 and 0.68 mm). Posterior
margins of body segments serrate. Body clothed with minute spinules.
Urosomites 2-5 with rows of slender spines anterior to serrate posterior
margin, spine row sometimes incomplete middorsally on urosomite 5,
spines and serrations interrupted ventrally by leg 6 on male urosomite 2.
Anal operculum with numerous minute serrations. Caudal ramus conical,
length about 3 times width of distal margin, dorsal ridge well de-
veloped, dorsal spine inserted at about proximal third of ramus, medial
margin with row of fine setae, lateral part of dorsal surface with
delicate spines, more numerous in male. Middle caudal seta about 14
length of body. Nuchal organ oval. Genital field as in fig. 1f; leg 6 a
single seta.
Female Ist antenna 7-merous, rarely 8-merous when long distal seg-
ment divided by suture; Ist segment with curved row of fine spinules
on dorsal surface and curved row of larger spinules on ventral surface;
esthete of 4th segment reaching just beyond apex of appendage; all
segments except 4th and 5th with a single sparsely plumose seta on
anterior margin. Male Ist antennae with moderately enlarged 4th
segment.
2nd antenna with l-merous exopod bearing 4 setae and 2 surface
spines. Mandible with l-merous palp 1 bearing 4 setae; gnathal lobe
armed from ventral to dorsal with a robust bicuspid tooth, a slightly
less robust tricuspid tooth, several smaller teeth with sharp cusps, and
a dorsal seta. Proximal lobe (precoxa) of 1st maxilla bearing 9 spines;
middle lobe (coxa) with a single spine; distal lobe with 7 spines.
Harpacticoid Copepods 573
ON ary reeponnt
yt Veil
UMA I
le! AG
Fyn i yn we! rie ie
D rhe
'" rag natth eR ET
i a Ue Wont an A
(
i
(aaa Dar re ‘|
petty,
tn ¢ tn Hh 1 ry
Fic. 1. Attheyella pilosa Chappuis: a, male, lateral; b, anterior end
of female, dorsal; c, posterior urosome of male, dorsal; d, posterior
urosome of female, dorsal; e, female caudal ramus, ventral; f, genital
field.
574 Proceedings of the Biological Society of Washington
Proximal lobe of 2nd maxilla with 2 setae, one with recurved tip bearing
long setules; 2nd lobe with 3 setae, 2 of them similar to those of proximal
lobe; endopod with strong terminal spine flanked by 2 setae, endite
bearing 3 setae. Ist segment with proximal seta, anterior margin serrate.
Sternal plate inconspicuous, with smooth margin. Leg 1, exopod
shorter than Ist segment of endopod; 3rd segment of endopod shorter
than that of A. carolinensis. Legs 2-4 like those of A. carolinensis;
endopod of male leg 2 without subterminal lateral spine or 2nd _ seg-
ment; leg 3 endopod 2-merous in female, 3-merous in male, medial
process of 2nd segment doubly curved with minute terminal barb; 2nd
segment of leg 4 endopod usually with 4 setae in addition to subterminal
spine, occasionally with 5 setae. Female leg 5 with low basal expansion
of proximal segment bearing 5 setae of which most lateral is by far the
shortest; distal segment less than twice as long as wide, with rows of
surface spines and 5 marginal setae, medial seta only slightly longer
than longest distal seta. Male leg 5 with 2 setae on basal expansion,
lateral half as long as medial, sometimes absent. Leg 6 a low plate
with 2 strong pinnate setae and a slender plumose outer seta.
Relationships: Of the 9 species of Attheyella known from North
America (according to Wilson and Yeatman, 1959), only A. americana
(Herrick), A. carolinensis Chappuis, A. illinoisensis (S. A. Forbes),
A. obatogamensis (Willey), and A. pilosa have been found in the
United States east of the Mississippi River. Three of these species are
readily distinguishable from A. pilosa. In A. americana and A. illinoisensis
the caudal ramus is short, about as wide as long, and in A. obatogamensis
the female caudal ramus has a prominent spiniform process on the
medial margin. A. carolinensis, however, closely resembles A. pilosa,
but can be distinguished from it as follows:
A. pilosa
1. Caudal ramus narrower, with
comb of fine setae along medial
A. carolinensis
1. Caudal ramus broader, with-
out setal comb on inner mar-
to
margin and from few to many
minute triangular spines on
dorsal surface.
Spine row anterior to serrate
margin of urosomites complete
dorsally (may be incomplete
gin, with spiniform setae on
lateral margin and in row ex-
tending posteriad from dorsal
seta.
Spine row limited to ventral
and lateral parts of urosomites.
in @ urosomite 5). 3. 83rd segment of endopod of
3. 3rd segment of endopod of leg leg 1 longer.
1 shorter. 4. Basal expansion of @ leg 5
4. Basal expansion of @ leg 5 with 4 setae.
with 5 setae. 5. Distal segment of @ leg 5
5. Distal segment of 2 leg 5 short longer and narrower, about 2.6
and wide, about 1.7 times as
long as wide; longest seta only
slightly longer than medial seta.
times as long as wide; longest
seta more than twice as long
as medial seta.
Harpacticoid Copepods 575
Fic. 2. Attheyella pilosa Chappuis: a, antenna 1, female; b, antenna
1, male; c, antenna 2, male; d, gnathal lobe of mandible, female; e,
maxilla 1, female; f, maxilla 2, female; g, maxilliped, male.
576 Proceedings of the Biological Society of Washington
A more detailed comparison than we have made would undoubtedly
reveal other differences, but those listed above were most obvious to
us. In practice we have found contrasting items 2 and 5 most useful,
since the ornamentation of the caudal ramus (item 1) is often obscured
by adhering particles of detritus. The spine rows of the urosomites are
easily seen in specimens cleared in glycerine or lactic acid, and the
relative lengths of the setae of the distal segment of leg 5 are usually
visible in undissected specimens viewed laterally.
Localities of specimens examined (major drainage systems listed in
parentheses). VIRGINIA.—Giles Co.: Sinking Creek, Newport (New
River), H. H. Hobbs, Jr. Smyth Co.: Middle Fork, Holston River
(Tennessee R.), 2.5 mi. E of Marion, R. D. Ross. White Oak Branch,
just above confluence with North Fork of Holston River, Chatham Hill
(Tennessee R.), W. Harman & H. H. Hobbs, Jr. SOUTH CAROLINA.—
Greenville Co.: In pools of cascading stream about 7 mi. W of Cleve-
land on US 276, (Saluda R.), H. H. Hobbs, Jr. GEORGIA.—Catoosa
Co.: Tributary to Chicamauga Creek crossing Ga. 2 4.1 mi. W of
junction of Ga. 71 and Ga. 2 (Tennessee R.), H. H. Hobbs, Jr. Catoosa
Springs, 2 mi. E of Ringgold (Tennessee R.), S. Peck, A. Fiske. Chatooga
Co.: Blowing Springs Cave, 2.5 mi. E of Cloudland (Tennessee R.), J. R.
Holsinger. Floyd Co.: Stream about 5 mi. W of Rome on Ga. 20
(Coosa R.), H. H. Hobbs, Jr. Stream 2 mi. W of Junction of Ga. 100
N and Ga. 20 on Ga. 20 (Coosa R.), H. H. Hobbs, Jr. Polk Co:
Tributary to Cedar Creek at northern city limits of Cedartown on Ga.
100 (Coosa R.), H. H. Hobbs, Jr. TENNESSEE.—Unicoi Co.: Erwin
Highway, Buffalo Creek; Erwin Fish Hatchery (both Tennessee R.);
P. C. Holt and class. Jackson Co.: Roaring River, 8-9 mi. above mouth
(Cumberland R.). KENTUCKY.—Hardin Co.: Nelson Cave (Ohio R.),
T. C. Barr. Edmonson Co.: Mammoth Cave (Ohio R. via Green R.),
Chappuis (1929a). Hart Co.: Cub Run Cave (Ohio R. via Green R.),
Barr (1968). Meade Co.: Doe Run (Ohio R.), R. Prins. INDIANA.—
Lawrence Co.: Donnaldson Cave (Wabash R.), Chappuis (1929a).
ILLINOIS.—Ogle Co.: Kilbuck Creek, 16.4 mi. W of Kingston (Mis-
sissippi R.), P. C. Holt.
Attheyella carolinensis Chappuis
Fig. 3k-—l
Attheyella (Brehmiella) carolinensis Chappuis, 1932, pp. 226-229, figs.
1-10.—Lang, 1948, pp. 985-986, fig. 392.1 [copied from Chappuis
and from Coker].
Attheyella carolinensis Chappuis.—Coker, 1934, pp. 116-118, pls. 10-
11.—Carter, 1944, p. 158.—Pennak, 1953, p. 407, figs. 256C [from
Coker].
Attheyella (Ryloviella) carolinensis Chappuis.—Borutsky, 1952, p. 283
[p. 261 in 1964 translation], fig. 79: 7-9 [copied from Coker].
Harpacticoid Copepods 577
Fic. 3. Attheyella pilosa Chappuis: a, leg 1, female; b, leg 2, female;
c, leg 3, female, distal segment of endopod; d, leg 3, male, endopod; e,
leg 4, female; f, leg 4, endopod, from another female; g, leg 4, male,
endopod; h, leg 5, female; i, leg 5, male; j, leg 6, male. Attheyella
carolinensis Chappuis: k, leg 1, female, endopod; 1, leg 5, female.
578 Proceedings of the Biological Society of Washington
Attheyella (Mrazekiella) carolinensis Chappuis—M. S. Wilson and Yeat-
man, 1959, p. 847, fig. 29.200 [copied from Coker].
Coker (1934) has provided a well illustrated account of this species,
but a few points should be clarified. In Coker’s account of the head
appendages the Ist antenna is called “antenna” and the 2nd antenna is
called “antennnule.” His drawing of leg 1 (pl. 11, fig. 1) omits the row
of setae along the medial margin of the Ist segment of the endopod.
A nuchal organ identical in appearance to that of A. pilosa is present
altho not illustrated by Coker.
Localities of specimens examined (major drainage systems listed in
parentheses). WEST VIRGINIA.—Greenbriar Co.: Fuller's Cave,
37°56/00"N, 80°25'38”W, Paul J. Starr. VIRGINIA.—Mountain Lake
(county and drainage system uncertain), Marjorie E. Carter (1944).
Carrol Co.: Stream flowing S from plateau 10 mi. S of Hillsville on
US 221, elevation 3000 ft. (Ohio R. via New River), H. H. Hobbs,
Jr. Smyth Co.: McHenry Creek off Va. 91, Saltville (Tennessee R.
via Holston R.), J. R. Cunningham. NORTH CAROLINA.—Orange
Co.: Chapel Hill (Cape Fear R.), R. E. Coker (1934). Transylvania
Co.: Thompson River headwaters, 1.1 mi. N of Bohaynee Church
(Savannah R. via Keowee R.), R. Prins. Jackson Co.: Whitewater
River, off NC 107 (Keowee R.), R. Prins. Tributary from Fairfield
Lake off US 64 (Keowee R.), R. Prins et al. East Fork Chatooga River
at N.C.-S.C. state line off N.C. 107 (“old” Tugaloo R.), R. Prins et al.
SOUTH CAROLINA.—Pickens Co.: Tributary to Oolenoy River, 0.3
mi. E of SC 11 off US 69 (Saluda R.), R. Prins, G. E. Dillard. Green-
ville Co.: About 7 mi. W of Cleveland on US 76 in pools of cascading
stream (Saluda R.), H. H. Hobbs, Jr. South Saluda River at boundary
of Pickens and Greenville Cos. on SC 11 (Saluda R.), H. H. Hobbs,
Jr. Oconee Co.: Chauga River at Cassidy Bridge on SC 290 (“old”
Tugaloo R.), R. Prins, J. R. Cunningham, V. H. McCaskill. Mud Creek
at SC 172 (Little R.), R. M. Shealy. Ramsey Creek at Chauga River
on US 76, about 5 mi. W of Westminster (Savannah R.), H. H. Hobbs,
Jr. Small stream between Long Creek and Chatooga R. on US 76
Chatooga R.), H. H. Hobbs, Jr. GEORGIA.—Rabun Co.: Timpson
Creek, about 6 mi. W of Clayton, on US 76 (Savannah R.), H. H.
Hobbs, Jr. Small tributary to Chatooga River immediately W of crossing
on US 76 (Chatooga R.), H. H. Hobbs, Jr. Towns Co.: Stream at
junction of Ga. 17 and Ga. 66, 12 mi. S of Hiawassee (Hiwassee R.),
H. H. Hobbs, Jr. Stream at northern city limits of Young Harris
(Hiwassee R.), H. H. Hobbs, Jr. Union Co.: 1.7 mi. NE of Blairs-
ville on US 76 (Hiwassee R.), H. H. Hobbs, Jr. Fannin Co.: Small
stream 3 mi. N of Blue Ridge on Ga. 5 (Hiwassee R.), H. H. Hobbs,
Jr. 7 mi. S of Morganton on Ga. 60, seepage area and mountain spring
(Hiwassee R.), H. H. Hobbs, Jr. Gilmer Co.: Hells Creek, running
into Carter's Reservoir (Cooawattee R.), H. H. Hobbs, Jr. Pickens
Co.: Tributary to Talking Rock Creek, 1.7 mi. E of boundary of
Harpacticoid Copepods 579
)
if
wes
Fic. 4. Distribution records of Attheyella pilosa (stars) and A.
carolinensis (triangles). Some records in Georgia are omitted because
of the scale; these are shown in fig. 5.
Pickens and Fannin Cos. on Ga. 156 (Coosawattee R.), H. H. Hobbs,
Jr. KENTUCKY.—Bell Co.: Mill Creek, near Pineville (Cumberland
R.), Donald L. Batch.
DISTRIBUTION OF ATTHEYELLA PILOSA AND A. CAROLINENSIS
Locality records for the two harpacticoid species are shown in figure
4, All records except the type-localities are from collections that we
have examined. In only one instance did both species occur in a single
580 Proceedings of the Biological Society of Washington
;
Fic. 5. Distribution records of Attheyella pilosa (squares) and A.
carolinensis (circles) in northern Georgia and parts of adjacent states.
collection; this was from a stream belonging to the Saluda River drainage
system near Cleveland, Greenville Co., S. C. The associated crayfishes
were Cambarus asperimanus and an undescribed species of Cambarus
belonging to the C. bartonii group. Our records are far too inadequate
to support any detailed generalizations concerning the distribution
patterns, but the preponderance of A. pilosa west of the Appalachians
and of A. carolinensis in the drainage systems of streams flowing into
the Atlantic is obvious. The area for which we have the most detailed
information on the occurrence of the two species is in northern Georgia
and northwestern South Carolina, where collections of crayfishes have
been obtained by H. H. Hobbs, Jr. and by R. Prins. Figure 5 shows
the distribution of the two species in this region and some of the
principal tributaries of the major drainage systems. Besides occurring
in the upper drainage systems of the Savannah and Saluda Rivers,
A. carolinensis is present in streams leading to the Tennessee and Coosa
Rivers. Progressing down-stream in the drainage of the Coosa, and
perhaps in that of the Tennessee, A. carolinensis is replaced by A. pilosa.
The record of A. carolinensis is southeastern Kentucky (Mill Creek,
Cumberland River drainage, near Pineville, from Cambarus distans)
upsets the otherwise rather neat east-west separation of the two species.
The limits of drainage systems, however, do not necessarily impose
barriers to their movements. Some species of crayfishes are known to
Harpacticoid Copepods 581
20
INFESTATION
%
JAN |FEB | MAR! APR! MAY IJUNE!JULY | AUG § SEPT! OCT I NOV !DEC IJAN |! FEB! MAR | APR ! MAY
AUG SEPT.
aE oe oO ENCE OY tne ear
Fic. 6. Incidence of infestation of Orconectes r. rusticus with
Attheyella pilosa at Doe Run, Kentucky.
migrate over land during damp weather, presumably carrying their
associates with them. Moreover, migration between stream systems
thru underground connections is a likely possibility for the two
harpacticoids. The type-locality of A. pilosa is subterranean, and both
species are inhabitants of spring-fed streams. Finally, the role of
stream piracy on distribution cannot be overlooked. It really is not
surprising that the distribution patterns of the two species do not
coincide exactly with those of drainage systems.
Tue BioLtocy or ATTHEYELLA PILOSA AT DoE RuN
As reported by Prins (1964), A. pilosa (called A. carolinensis by
Prins) was collected at Doe Run in “dense reddish assemblages from
the pleopods, the bases of the coxae of the pereiopods, the hairs around
the sterna, and various other places on the under-surfaces of Cambarus
tenebrosus Hay .. . and Orconectes rusticus rusticus (Girard).” Between
November 1962 and September 1964 Prins collected 22 samples from
3 collecting sites on Doe Run: Station I, at the source of the stream;
Station IA, 4 mile downstream, and Station II, 2 miles downstream
(Minckley, 1963). These collections, which were made during every
month except October and December, have been studied in an attempt
to obtain information on the life history of A. pilosa. From each sample
a number of specimens (between 50 and 160) were selected randomly.
For each specimen selected the sex and developmental stage were
determined, and the total length (tip of rostrum to end of caudal rami)
was measured.
Seasonal occurrence: Of the crayfish hosts, only Cambarus occurred
at stations I and IA; both Cambarus and Orconeéctes occurred at station
II. The incidence of infestation was usually much higher in Cambarus,
perhaps because of their habit of spending more time in burrows in the
banks and on the channel bottom where they may be more accessible
582 Proceedings of the Biological Society of Washington
Ee fora S| oe eee IZANOVIG2
ee 16 Feb 63
___ es 23 Mar
EEze ey 26 June
Se ee 22 Nov
ee | es 6 June
20 June eal pee [Saleem
ee 27 June
ee | pe 8 July
Fic. 7. Variation in length of Attheyella pilosa populations at Doe
Run, Kentucky. For each diagram the horizontal line represents the
sample range, the central vertical line the sample mean, the inner white
rectangles the value of 2 standard errors; the distance from the mean
to the edge of an outer rectangle equals the value of 1 standard devia-
tion. Stations indicated by outer rectangle: station I—white, station
IA—stippled, station II—black.
to the copepods. Seasonal variations in the incidence of infestation of
Cambarus, which molts thruout the year in Doe Run, were very erratic.
On the other hand, infestation of Orconectes is greatest in the winter
when it is torpid and rarely molts, and least in midsummer when molting
is most frequent (fig. 6). This suggests that Attheyella as has been
suggested for branchiobdellid worms (Hobbs, Holt, and Walton, 1967,
p. 13) does not return to the host when the old exoskeleton is shed in
molting. In contrast, Hobbs, Holt, and Walton (1967, p. 12) report
that after a molt the entocytherid ostracods leave the host’s recently
molted skeleton and make their way to the new one.
Only adult males and females of Attheyella were found, in spite of a
careful search for copepodids. Immature specimens must be free-living
on the stream bottom. In the harpacticoid Nitocrella divaricata (Chap-
Harpacticoid Copepods 583
puis), which lives in the gill chamber and on the body surface of the
European crayfish Astacus astacus (L.), both copepodids and adults
are found on the host; only the nauplii are free-living (Chappuis, 1927).
At present we do not know to what extent association with a crayfish
host is mandatory for A. pilosa, since a systematic search has not been
made for free-living specimens in Doe Run.
Females with egg sacs were found in samples collected during
February, March, April, June, and September. Breeding probably
occurs thruout the year.
Sex ratio: In most samples the numbers of males and females were
nearly equal. The percentage of males varied from 25 (5 May 1964)
to 65 (22 Nov. 1964, sta. IA). We feel that these fluctuations are
fortuitous rather than indicative of significant variations in the sex
ratio. The percentage of males in all samples combined was 49.8.
Population structure: Size-frequency histograms were constructed
for all samples in an attempt to identify the generations comprising the
populations and to trace their emergence and decline from month to
month. The composition of the samples proved to be so complex that
the desired information could not be extracted with any confidence.
The histograms had from 2 to several peaks, not all clearly defined, and
it usually was not possible to correlate the peaks of one sample with
those of another.
Such a picture is to be expected from a multivoltine species that
breeds during most or all of the year and has a short period between
generations (probably not more than 1 or 2 months). This results in
several generations, perhaps 5-10 during a year. The size composition
of one generation may differ from that of another generation in the
same local population, and the size composition also may be different
in comparable generations from different local populations. As it moves
about, a crayfish may encounter and be infested by several different
local populations of Attheyella. Thus the harpacticoid assemblages on a
crayfish may be so heterogeneous that it is extremely difficult to identify
the component populations.
Fig. 7 shows the seasonal variation in length of A. pilosa at the 3
stations. We can offer no explanation for the fluctuations other than
the heterogeneity of the samples. The statistical variability of the samples
does not seem excessive; the coefficient of variability ranged from 2.4
to 8.4, but usually lay between 3 and 6. We could find no relationship
between size and water temperature; this is not surprising since the
annual range in temperature in the cool, spring-fed Doe Run was
slight: Station I, 2.6°C, Station IA, 4°C, and Station II, about 8°C.
ACKNOWLEDGMENTS
Many of the harpacticoids were sorted for us from crayfishes by
Margaret Walton at the request of Horton H. Hobbs, Jr., who collected
them. We are grateful to Dr. Hobbs for his cooperation and to Miss
584 Proceedings of the Biological Society of Washington
Walton for patiently undertaking this tedious task. Field work at Doe
Run by Prins was supported in part by U. S. Atomic Energy Commission
under contract No. AT-(40-1)-2595 with the University of Louisville,
Louisville, Kentucky. For critical reviews of the manuscript we are
grateful to Drs. Gerald C. Cole, John R. Holsinger, and Perry C. Holt.
LITERATURE CITED
Barr, THoMas C., Jr. 1968. Ecological studies in the Mammoth Cave
system of Kentucky. Intern. J. Speleology 3(1+2): 147-
204, pls. 37-64.
Borutsky, E. V. 1952. Crustacea, vol. 3, no. 4, Freshwater Harpacti-
coida. Fauna SSSR, Zool. Inst. Akad. Nauk SSSR, N.S., no.
50, 424 pp. [English translation by Israel Program for
Scientific Translations, 396 pp., published in 1964].
Carter, Marjorie E. 1944. Harpacticoid copepods of the region of
Mountain Lake, Virginia (with description of Moraria
virginiana n. sp.). J. Elisha Mitchell Sci. Soc., vol. 60, no. 2,
pp. 158-166, pls. 65-67.
Cuappuis, P. A. 1927. Harpacticiden aus der Kiemenhohle des Flus-
skrebses. Archiv fur Hydrobiologie, vol. 17, no. 3, pp. 515-
520.
1929a. Copépodes cavernicoles de Amérique du Nord.
(note préliminaire). Bull. Soc. Stiinte Cluj (Romania), vol.
4, pt. 2, pp. 51-57.
1929b. Die Unterfamilie der Canthocamptinae. Archiv
fur Hydrobiologie, vol. 20, pp. 471-516.
. 1931. Biospeologica no. 16, Campagne spéologique de C.
Bolivar et R. Jeannel dans Amérique du Nord (1928). 4,
Crustacés Copépodes. Arch. Zool. Expér. Gén., vol. 71, no.
3, pp. 345-360.
1932. Eine neue Attheyella aus Nordamerika. Attheyella
(Brehmiella) carolinensis n. sp. Zool. Anz., vol. 98, nos. 7/8,
pp. 226-229.
1933. Biospeologica no. 59, Copépodes (premiére série),
avec l’énumération de tous les Copépodes cavernicoles connus
en 1930. Arch. Zool. Expér. Gén., vol. 76, no. 1, pp. 1-57.
Coxer, Rosert FE. 1934. Contribution to knowledge of North Ameri-
can freshwater harpacticoid copepod Crustacea. J. Elisha
Mitchell Sci. Soc., vol. 50, no. 1-2, pp. 75-141, pls. 1-15.
Hosss, Horton H., Jr., Hott, PERry C., AnD WALTON, MARGARET.
1967. The crayfishes and their epizootic ostracod and
branchiobdellid associates of the Mountain Lake, Virginia,
region. Proc. U. S. Nat. Mus., vol. 123, no. 3602, pp. 1-84.
Lanc, Karu. 1948. Monographie der Harpacticiden. Stockholm, vol.
1, pp. 1-896, figs. 1-361; vol. 2, pp. 897-1682, figs. 362—
607, maps 1-378.
Harpacticoid Copepods 585
MinckteEy, W. L. 1963. Studies of the biology of a spring stream.
Doe Run, Meade County, Kentucky. Wildlife Mongr., No.
11, pp. 1-124.
PENNAK, RoperT W. 1953. Fresh-water invertebrates of the United
States. 769 pp., Ronald Press, New York.
Prins, RupotpH. 1964. Attheyella carolinensis Chappuis (Copepoda;
Harpacticoida) on freshwater crayfishes from Kentucky.
Trans. American Microscop. Soc., vol. 83, no. 3, pp. 370-371.
In press. Comparative ecology of the crayfishes Orconectes
rusticus rusticus and Cambarus tenebrosus in Doe Run, Meade
County, Kentucky. Intern. Rev. Ges. Hydrobiol.
Witson, Cartes B. 1936. Copepods from the cenotes and caves
of the Yucatan Peninsula, with notes on cladocerans. Carnegie
Inst. Washington Publ. no. 457, pp. 77-88.
Witson, Mitprep S. 1956. North American harpacticoid copepods. 1.
Comments on the known freshwater species of the Cantho-
camptidae. 2. Cantocamptus oregonensis n. sp. from Oregon
and California. Trans. American Microscop. Soc., vol. 75,
no. 3, pp. 290-307.
Witson, Miuprep S., AND Harry C. YEATMAN. 1959. Free-living
Copepoda, pp. 735-861. In W. T. Edmondson (ed.), Ward
and Whipple’s freshwater biology, 1248 pp., John Wiley &
Sons, New York.
586 Proceedings of the Biological Society of Washington
@ /—>
Vol. 81, pp. 587-604 30 December 1968
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
A NEW GENUS AND TWO NEW SPECIES OF
SPHAEROMATID ISOPODS FROM THE
HIGH INTERTIDAL ZONE AT
NAOS ISLAND, PANAMA
By PETER W. GLYNN
Smithsonian Tropical Research Institute, P.O. Box 2072,
Balboa, Canal Zone
The three new species of sphaeromatid isopods described in
this paper were found living together, in association with a
large barnacle in the high intertidal zone, at Naos Island on
the Pacific shore of Panama (79°32’08”’W; 8°54’57”N ). Two
of the species can be placed readily in the genus Dynamenella
of the Eubranchiatae. The relationship of the third form is
not so clear-cut; it has been described here as a new genus and
species in the platybranchiate group. A search for members
of the Sphaeromatidae was undertaken in this area in connec-
tion with previous studies of shore forms, especially symbiotic
species, in tropical Atlantic waters (Menzies and Glynn in press;
Glynn, in press). Appreciation is expressed for the opportunity
provided by I. Rubinoff and I. E. Wallen, Smithsonian Institu-
tion, to initiate these studies on the Panamanian isopod fauna.
Critical comments offered by R. J. Menzies (Florida State Uni-
versity ) and T. E. Bowman (Smithsonian Institution ) helped
to improve the manuscript; their remarks are also gratefully
acknowledged.
Formalin washings of cobbles and boulders, and other likely
retreats (small and sparse patches of encrusting algae; Chiton
stokesii Broderip, 1832, the only large intertidal chiton; oyster
shells, both living and dead of the two species Ostrea iridescens
Hanley, 1854 and O. conchaphila Carpenter, 1856; the large
pulmonate Siphonaria gigas Sowerby, 1825) were obtained
systematically at all horizons in the intertidal belt over the
54—Proc. Biot. Soc. Wasu., Vou. 81, 1968 (587)
588 Proceedings of the Biological Society of Washington
Fic. 1. Type-locality at Naos Island, Panama (2 April 1968). The
vertical rock face is dominated by clusters of both living and dead
Tetraclita squamosa panamensis. The substratum is fine grained, igneous
rock (dacite).
: - seta
period January 18-23, 1967. At that time numerous isopods
were found only with the abundant and large sessile cirriped
Tetraclita squamosa panamensis Pilsbry, 1916 on the upper
shore near and above mean sea level (Fig. 1). (More recently
one species of Dynamenella was collected from dense growths
of the barnacle Chthamalus). The sites occupied on Tetraclita,
at low water when the collections were made, were the spaces
between the basal plate and substratum, the parietal canals
within the parapet and inside dead, empty tests, peripherally
where the parapet joins the basis. These various niches also
provide refuge for an overlapping micro-fauna composed of
elements from the upper and lower reaches of the tide. In-
cluded in this assemblage are juvenile gastropods, Onchidella,
Lasaea, sipunculids, anemones, mites, the springtail Anurida
maritima, pseudoscorpions, etc. Although hundreds of individ-
uals of the three species of isopods were found inhabiting less
than 20 barnacles growing in juxtaposition, attempts to collect
New Panama Isopods 589
the isopods from several other barnacle clusters in the same
vicinity were without success.
The factors responsible for the patchy occurrence of the
isopods, their interspecific interactions, and the symbiotic rela-
tions with the barnacle host, are questions which immediately
come to mind. These animals may provide good material for
the investigation of such problems.
FAMILY SPHAEROMATIDAE DAHL, 1916
Cephalon with well developed epistome, not divided into frontal
plate and clypeus, and rarely fused with upper surface of cephalon.
Peduncle of Ant’ (first antennae) three-jointed, of Ant’ five-jointed.
Mouth parts biting or gnawing, never really suctorial; second joint of
maxillipeds at least in males and immature specimens without external
expansion; mouth parts in females with brood rather frequently strongly
metamorphosed and useless for nutrition. Peraeon with seven somites,
all free; marsupial lamellae only on second, third and fourth “epimera’,
rarely wanting. The brood develops in pouches and not out free in
the marsupium. All pleopods (Plp) lamellar; all endopods, and at least
exopods of Plp* and Plp? unjointed; at least both rami of Plp' and Plp?
fringed with long plumose setae; both rami of Plp‘ and Plp® (or endopod
of Plp* and Plp’) without such setae, but adapted for a respiratory
function. Pleon and pleotelson combine functionally to form a vault in
which the pleopods are enclosed. Uropods with rami unjointed, these,
at least in females, generally depressed, sometimes one or even both
wanting. Body can be rolled more or less completely into a_ ball
(slightly modified from Hansen, 1905).
Group EUBRANCHIATAE
Plp* and Plp*® have both rami subsimilar, with deep, essentially
transverse folds, often of fleshy aspect, without PMS (plumose marginal
setae); exopod of Plp® generally distinctly two-jointed, with the sub-
apical squamiferous protuberance on lower surface very high. Plp* have
both rami closely set with long PMS at least on distal margin. Endopod
of Plp’ at least rather broad, scarcely ever half as long again as broad.
End of pleotelson at least emarginate, generally with notch or with a
slit terminating in a foramen (Hansen, 1905).
Genus Dynamenella Hansen, 1905
Type-species—Dynamenella perforata (Moore, 1901).
Diagnosis: Both sexes rather similar in aspect, without real processes;
abdomen with notch which is semicircular or oblong in female, in male
narrow in distal part, while proximal part constitutes a transverse
foramen; uropods subsimilar in both sexes, with rami lamellar. Mouth-
590 Proceedings of the Biological Society of Washington
Md
Fic. 2. Dynamenella josephi new species; A. male holotype, length
3.2 mm; B. enlarged view of pleonal suture; C. pleotelson of female
allotype, length 2.5-mm; P’, seventh peraeopod and enlarged view of
apical segments; P', first peraeopod and enlarged view of apical seg-
ments; Ant’, first antenna and enlarged view of apical segments; Ant,
second antenna; Md, mandible and mandibular palp; Mx’, second
maxilla; Mxp, maxilliped; Mx’, first maxilla.
parts similar in both sexes; male with appendix masculinum on endopod
of Plp*; marsupial lamellae overlap. each other somewhat. Hansen also
noted, but incorrectly, that exopod: of Plp* is unjointed and basal joint
of Ant’ is of usual shape, not expanded in a free. plate.
New Panama Isopods 591
Fic. 3. Dynamenella josephi new species; Plp’, first pleopod; Plp’,
second pleopod and stylet with enlarged view of tip; P, penes; Plp’,
third pleopod; Plp*, fourth pleopod; Plp’, fifth pleopod.
Remarks: The subsequent description of several species of Dy-
namenella has somewhat modified Hansen’s original concept of the
genus. For example, the pleotelson in males of D. tropica (Loyola e
Silva, 1960), D. acutitelson and D. barnardi (Menzies and Glynn, in
press) is provided with just a simple indentation, lacking a transverse
foramen. Further, the female of D. acutitelson var. glabrothorax is with-
out any notch on the pleotelson, but rather has the terminal border
drawn out as a blunt spine. Typically the exopod of Plp* is jointed.
592 Proceedings of the Biological Society of Washington
Dynamenella josephi new species
Figures 2 and 3
Diagnosis: Sexes dimorphic in pleotelson; mature male with slit and
transverse foramen on posterior border of pleotelson; female with smooth,
ovate border or at most with faintly perceptible indentation. First
pleonal somite in both sexes with a pair of excrescences near midline;
anterior border of pleotelson with two small pairs of tuberculate ridges;
rear half of pleotelson with single pair of elongate, tuberculate ridges
lateral to midline. These ridges are smooth and rounded in female,
but more irregular and ragged in male. Tuberculate carina on midline
of pleotelson, extending forward about half its length. Pleonal suture
with a single, forwardly directed incision. This incision flexes medially
near anterior termination. Male with a pair of circular, pigment-free
spots near midline on seventh peraeonal somite. Both rami of Plp’-Plp®
with PMS. Exopod of Plp* jointed. Plp* and Plp°® with both rami
membranous and provided with conspicuous transverse folds. Exopod
of Plp* with cleft on upper, lateral margin; endopod with apical spine.
Exopod of Plp’ jointed; three prominent, squamiferous protuberances
with one apical member bluntly spinose. Stylet nearly twice length of
endopod of Plp*. Penes long, one and one-half times the length of Plp’,
and tenuiform.
Measurements: Male holotype, length 3.2 mm, width 1.4 mm. Fe-
male allotype (illustrated), length 2.5 mm, width 1.2 mm.
Type-locality: Naos Island, Panama, 22 January 1967; holotype
(USNM 122850), plus 27 males, 35 females, 307 young paratypes
(USNM 122851).
Distribution: Type-locality and 12 specimens from Tortola Island,
Panama (halfway between Taboga Island and the Palo Seco leper colony
on the mainland), 25 September 1967, upper intertidal zone from
under Tetraclita squamosa panamensis; 35 specimens from Périco Island,
Panama, collected from under cluster of Chthamalus sp. in the upper
intertidal zone, 30 March 1968.
Supplementary descriptive notes: Surface of body covered with
minute granulations which are especially evident on the pleon of large
males. Body sparsely setose. Second through seventh peraeonal somites
subequal in size; first peraeonal somite nearly twice the length of
second and tapering in width toward cephalon which is slightly over
half the width of mid-peraeonal somites. Eyes large, occupying postero-
lateral angles of cephalon. Posterior border of seventh peraeonal somite
with flange on either side of midline. Central portion of pleotelson
inflated in both sexes, forming a conspicuous, hemispherical elevation.
Posterior border of the pleotelson in small males (ca. 1.8 mm and less)
bears an incipient cleft only. Larger males, of around 2.3 mm in length
and greater, have both a cleft and foramen. The development of the
mature pleotelson is very similar to that in Dynamenella perforata
(Moore, 1901; Glynn, in press). Pleonal suture similar in both sexes.
New Panama Isopods 593
Female without pair of circular, pigment-free spots on seventh
peraeonal somite. Uropods broad in mature male, with relatively
smooth border; exopods reach posterior border of pleotelson and
endopods extend noticeably beyond this. Female uropods smaller, not
quite reaching to posterior edge of pleotelson. Second, third and fourth
articles of Mxp (maxilliped) palp slightly expanded; endite with a
single coupling hook. Mx’ (first maxilla) exite with four teeth. Mx?
(second maxilla) and Md (mandible) also typical for genus, as illus-
trated. Female with equally well developed mouth parts. Ant’ slightly
shorter than Ant®. Flagellum of Ant' with eight articles and five
aesthetascs apically; flagellum of Ant? with 13 articles. P' (first
peraeopod) shortest, about one-half the length of P’. Blood sinuses
very evident on rami of all pleopods. Exopod of Plpt with 22 PMS and
a single, simple, blunt spine near base; endopod with 16 large and two
small PMS, medial border of endopod with numerous short setae.
Exopod of Plp? with 22 PMS; endopod with 15 PMS, four located
medially along groove in which stylet rests. Stylet smooth and tapering
except for scattered, minute spines terminally. Exopod of Plp* jointed
near terminal border and with 21 PMS; endopod with 11 PMS and
medial border with minute setae along entire length. Exopod of Plp‘
with seven prominent respiratory folds and lateral margin provided
with few, minute setae; endopod with four well developed respiratory
folds. Respiratory folds six in number on two-jointed exopod of Plp’,
lateral border of exopod sparsely setose; individual folds not readily
discernible but apparently eight in number and best developed on
endopod.
Affinities: Dynamenella josephi shows a very close resemblance to
D. perforata (Moore, 1901; Menzies and Glynn, in press) and D. antonii
(Loyola e Silva, 1960). The mature males of the three species are very
similar in appearance. However, whereas D. perforata and D. antonii
have pleonal sutures with two incisions, D. josephi can be readily
distinguished by a simpler suture of only one incision. The dimorphic
character of the pleotelson in the two sexes, and the development of
the slit and foramen in males of D. perforata and D. josephi, follow
essentially the same pattern. The pleopods are likewise strikingly
similar in these species. The stylet is long and tapering, the exopod of
Plp* is jointed marginally, the endopod of Plp* has an apical spine, and
the exopod of Plp® bears three prominent, squamiferous protuberances
and is jointed marginally. Finally, the penis is long, filiform and
bifurcated to the base.
The deficient description of Clianella elegans Boone, apparently a
closely related form, does not permit an entirely critical comparison with
D. josephi. The following relevant characters at least indicate that the
two species are distinct: basal articles of Ant’ swollen; frontal margin
of cephalon carinate and sculptured; foramen in pleotelson surrounds
an overhanging, blunt, round, tooth-like projection (Boone, 1923).
594 Proceedings of the Biological Society of Washington
Etymology: josephi, from the middle name of my son, who at an early
age preferred to “experiment” with Dynamenella perforata rather than
play with toys.
Dynamenella setosa new species
Figures 4, 5 and 6
Diagnosis: No obvious difference in the appearance of the two sexes.
Body covered with dense growth of long setae. Posterior border of
pleotelson denticulate and with a narrow slit communicating with
circular foramen. Central portion of pleotelson inflated. Pleonal suture
with a long incision leading forward for about half its length then
flexing medially; a second, shorter incision joins main cleft at point of
flexure. Uropods large, serrate and extend well beyond posterior border
of pleotelson. Both rami of Plp'-Plp* with PMS. Exopod of Plp* jointed.
Both rami of Plp* and Plp’? membranous and provided with deep-set,
transverse folds. Exopod of Plp’ jointed; one squamiferous protuberance
located laterally and two apically. Stylet long and tapering, nearly twice
the length of endopod of Plp*? (paratype specimen). Penis bifurcate
to base, long (exceeding length of Plp*) and tenuiform.
Measurements: Male holotype, length 3.1 mm, width 1.5 mm. Fe-
male allotype (illustrated), length 2.9 mm, width 1.5 mm.
Type-locality: _Naos Island, Panama, 22 January 1967; holotype
(USNM 122854) plus 1 male and 14 female paratypes (USNM 122855).
Distribution: Type-locality and 35 specimens from Tortola Island,
Panama, 25 September 1967. Hypotypes also associated with Tetraclita
squamosa panamensis in the upper intertidal zone.
Supplementary descriptive notes: Cephalon, peraeonal somites five
through seven, and pleon amply provided with minute tubercles and
their associated setae. Peraeonal somites two through seven subequal in
size; first somite twice the length of second. Cephalon and _ first
peraeonal somite equal in length. Eyes large, located at postero-lateral
angles of cephalon. Central, anterior margin of pleotelson depressed in
both sexes. The extent of development of the penes and stylet in the
holotype indicates that this specimen was not fully mature. Larger male
paratype (broken in half), with major features of pleotelson developed
to same extent as in holotype, does however show minor differences in
posterior half of body which should be noted. These differences include
a pair of circular, pigment-free spots near midline on posterior border of
seventh peraeonal somite (large females lack these); upward extension
of posterior margin of seventh peraeonal somite as flange; a pair of
ragged, spinous projections near midline of first pleonal somite; two
pairs of similar projections in line with above on pleotelson. A hyaline
rim is formed around foramen in pleotelson in large individuals of both
sexes. Mxp palp with second, third and fourth articles slightly ex-
panded; endite with a single coupling hook and two rows of teeth
(three in each). Mx* exite with five teeth ard three setae, endite with
New Panama Isopods 595
Ant!
Fic. 4. Dynamenella setosa new species; A. male holotype, length
3.1 mm; B. enlarged view of pleonal suture; C. pleotelson of female
allotype, length 2.9 mm; Mxp, maxilliped; Ant’, first antenna; Ant’,
second antenna; Mx’, first maxilla (near moult).
three setae. Mx” normal for genus. Md normal, with incisor heavily
armed with teeth and well developed molar. Females also with well
developed mouth parts. Ant' about two-thirds the length of Ant’.
Flagellum of Ant’ with nine articles and eight aesthetascs terminally.
Ant® with 14 articles in the flagellum. P’ comparatively short and stout,
596 Proceedings of the Biological Society of Washington
P| p2
Fic. 5. Dynamenella setosa new species; Plp'’, first pleopod; Plp’,
second pleopod with incompletely formed stylet; S, fully formed stylet
from paratype specimen; P, penes; p, penes from paratype specimen
shown at lower magnification; Plp*, third pleopod; Md, mandible and
mandibular palp (near moult).
rear peraeopods longer and more slender. Blood sinuses conspicuous
over surface of all pleopods. Plp' exopod with 20 PMS plus one simple,
short spine; endopod with 13 PMS. Plp? exopod with 23 PMS; endopod
with 10 PMS and incompletely formed stylet (holotype). Fully mature
stylet appears to possess one beveled edge. Exopod of Plp* jointed and
New Panama Isopods 597
Pl p4
Fic. 6. Dynamenella setosa new species; Plp’, fifth pleopod; Plp‘,
fourth pleopod; Mx*, second maxilla; P', first peraeopod; P‘, seventh
peraeopod.
with 27 PMS; endopod with 11 PMS. Both rami of Plp* with five pairs
of respiratory folds. Both rami of Plp® with six pairs of respiratory folds.
Lateral border of exopod of Plp° sparsely setose; minute, squarish
scales with marginal spines shield surfaces of squamiferous areas.
Affinities: Dynamenella eatoni (Miers), like D. setosa, also has the
pleotelson of both sexes nearly identical. Even though this condition
is unlike that in the type-species of the genus (D. perforata), Menzies
(1962) and Hansen (1905) decided to retain this species in Dynamenella.
598 Proceedings of the Biological Society of Washington
The pleonal suture in D. setosa, with two distinct incisions, is more
complex than in D. josephi. In this respect D. setosa is structurally
similar to D. dianae (Menzies) where the main, forwardly directed
incision appears to branch.
Etymology: setosa, from saetosus (L.), named for the conspicuous,
long setae covering the dorsum.
Group PLATYBRANCHIATAE
Plp* and Plp® have both rami completely without transverse folds,
and their exopods are unjointed; endopod of Plp* at most with a few
short terminal plumose setae, exopod of same pair rarely with numerous
long marginal plumose setae ( Tecticeps), in most genera both rami with-
out plumose setae; both rami of Plp® without plumose marginal setae,
and the exopod has the squamiferous protuberances slightly in relief
and in rare cases without spines or even wanting. Plp* have sometimes
plumose marginal setae on both rami as Plp*, sometimes with endopod
nearly naked or with both rami naked. Endopod of Plp’ rarely broad,
most frequently narrow. (End of abdomen sometimes with a rounded
notch, often truncate, rounded, or acute.) (Hansen, 1905).
SECTION CASSIDININI
Body much or exceedingly depressed; thorax considerably or strongly
expanded; margin of thorax, anterior part of abdomen, uropods and
sometimes the two proximal joints of Ant’ constituting a nearly con-
tinuous border ciliated with a more or less conspicuous rim of short
protruding setae. Eyes well developed. Two proximal joints of Ant’
with anterior part protruding, visible from above in at least almost
whole length, frequently much expanded in front, depressed. Mandibles
with masticatory process well developed. Anterior pairs of legs without
prehensile band. Endopod of Plp* at least somewhat longer than broad,
sometimes very narrow. Both rami of Plp* with several plumose setae
on terminal margin; exopod unjointed or two-jointed. Both rami of Plp*
and Plp’® without setae, subsimilar in aspect, respiratory. Posterior
margin of abdomen short; a real notch always wanting. Marsupial
lamellae wanting; brood in a chamber formed by two external pouches.
(Hansen, 1905).
Striella new genus
Type-species: Striella balani, new genus, new species.
Diagnosis: Intermediate species with characters about equally divided
between the hemibranchiate and platybranchiate groups. Body oval,
strongly depressed and fringed with dense, setose growth. Cephalon
confluent with and immersed in first and largest of peraeonal somites.
No obvious sexual differences. Pleotelson terminates posteriorly in
blunt point; minute incision faintly visible terminally. Outer rami of
uropods about two-thirds length of endopods. Ant’ roughly two-thirds
New Panama Isopods 599
the length of Ant®. Proximal joints of Ant’ not inflated or readily visible
in dorsal view. Neither does epistome protrude beyond cephalon. Lobes
of Mxp palp not produced on inner margin. Md with strongly developed
molar process and elongate, toothed incisor. P’'—P* with natatory setae.
Plp' endopod extremely elongate, about three times longer than wide.
Appendix masculinum on Plp? inflated and projecting slightly beyond
upper margin of endopod. All pleopods unjointed, and Plp* and Plp’
without transverse, respiratory folds. Penis long, tenuiform and bifurcate
to base.
Etymology: Striella, gender feminine, derived from combining the
initials of the Smithsonian Tropical Research Institute, behind whose
marine facility these animals were first collected.
Striella balani new species
Figures 7, 8 and 9
Diagnosis: Cephalon and first peraeonal somite equal in length.
Pleotelson essentially smooth with slightly raised, hemispherical area at
midline near forward border. Pleonal suture with two short incisions
which converge posteriorly. Marginal setose fringe around body best
developed in larger individuals. Flagellum of Ant’ with eight articles
and five aesthetascs; flagellum of Ant® with 14 articles (illustrated left
Ant? had seven terminal articles broken off; flagellum of right Ant? is
composed of 14 articles). Mouth parts very similar to those in
Cassidinidea Hansen and Dies Barnard (possibly a synonym of Cassi-
dinidea according to Menzies and Frankenberg, 1966), i.e. second, third
and fourth articles of Mxp palp not lobed, and Md with strongly formed
incisor and molar process. Plp'—Plp* with PMS; only two coupling
setae on medial margin of basis. Plp* exopod with 21 PMS and one
spine; narrow endopod with 13 PMS. Plp? exopod with 22 PMS;
endopod with 19 PMS, five aligned medially adjacent to stylet. Lateral
border of stylet hyaloid above and thickened below; minute spines in
couplets along medial border. Plp* exopod with 23 PMS; endopod with
10 PMS. Plp* and Plp® unjointed, without setae or respiratory folds, but
with conspicuous blood sinuses. A pair of creases extend across base of
exopod of Plp*; endopod with an apical spine. Plp°’ exopod with three
squamiferous protuberances, each with a delicate pattern of striae;
upper pair of protuberances capped with minute, horny spines.
Measurements: Male holotype, length 2.6 mm, width 1.8 mm. Fe-
male allotype (illustrated), length 2.1 mm, width 1.1 mm.
Type-locality: Naos Island, Panama, 22 January 1967; holotype
(USNM 122857) plus 37 males, 34 females, 42 young paratypes, USNM
122858.
Distribution: Known only from type-locality.
Affinities: Certain difficulties are met in trying to assign this species
to the present system of classification. In many respects it is similar
to Exosphaeroma and Pseudosphaeroma of the hemibranchiate group.
600 Proceedings of the Biological Society of Washington
Fic. 7. Striella balani new genus, new species; A. male holotype,
length 2.6 mm; B. enlarged view of pleonal suture; C. pleotelson of
female allotype, length 2.1 mm; P’, first peraeopod; P’, seventh
peraeopod; Ant’, first antenna; Ant®, second antenna (terminal section
of seven articles missing ).
Several of the characters already described, however, including absence
of branchiae on Plp* and Plp*, and the unjointed exopods of these
pleopods, necessitates placement of Striella with the Platybranchiatae.
Although the respiratory folds in Exosphaeroma alba are especially
weakly developed in this genus, examination of the pleopods in topotype
New Panama Isopods 601
Fic. 8. Striella balani new genus, new species; Plp’, first pleopod;
Plp*, second pleopod and enlarged view of stylet; P, penes; Plp*, third
pleopod; Plp*, fourth pleopod; Plp*, fifth pleopod with enlarged view
of apical, squamiferous protuberances.
specimens reveals their unquestioned presence. This is not the case in
Striella. It should be pointed out, though, that some species related to
Exosphaeroma are without folds (Menzies, 1954) and that Monod
(1931) has warned against placing too much emphasis on the presence
or absence of articulations.
602 Proceedings of the Biological Society of Washington
Mx! Mx? MXp Md
Fic. 9. Striella balani new genus, new species; Mx’, first maxilla;
Mx’, second maxilla; Mxp, maxilliped; Md, mandible and mandibular
palp.
Within the platybranchiates Striella satisfies all the characters of the
Cassidinini except one, namely the proximal joints of Ant’ are not
expanded or easily viewed from above. Further, assignment to the
genus Cassidinidea is not possible because the epistome does not
protrude as a broad plate separating Ant‘, and the uropodal exopod is
not greatly reduced in size.
Similarities with Exosphaeroma include at least the tenuiform, bifur-
cated penis; the inflated stylet; and the apical spine on the endopod
of Plp*. Except for the presence of respiratory folds and the dimorphic
character of the pleotelson, Chilton’s (1909) description of Pseudo-
sphaeroma also shares much in common with Striella.
With the present and incomplete understanding of the relationships
within the Sphaeromatidae, the proper assignment of transitional taxa
like Striella is vexing. While it is here suggested that this species
belongs to the platybranchiate line, one cannot dismiss the possibility
of descent from the hemibranchiates with emphasis on development
of an ovoid and depressed body structure.
Etymology: balani, from balanos (Gr.), because the first specimens
were collected from large clusters of barnacles.
LITERATURE CITED
Boone, Peart Lee. 1923. New Marine tanaid and isopod Crustacea
from California. Proc. Biol. Soc. Wash., vol. 36, pp. 147-156.
CuHILtTon, CHartes. 1909. The Crustacea of the Subantarctic Islands
of New Zealand. Subantarct. Is. New Zealand, vol..2, -axt.
26, pp. 601-671, 19 figs. vis
GLYNN, PETER W. In press. Ecological studies on chiton associations
in Puerto Rico with special reference to sphaeromid isopods.
Bull. Mar. Sci., 58 ms. pp., 12 figs., 13 tabs.
New Panama Isopods 603
Hansen, H. J. 1905. On the propagation, structure, and classification
of the family Sphaeromidae. Quart. J. Micr. Sci., New Ser.,
vol. 49, no. 1, pp. 69-135, 1 pl.
LoyoLta E Sinva, J. pe. 1960. Sphaeromatidae do litoral Brasileiro
(Isopoda-Crustacea). Bol. Univ. Parana, no. 4, pp. 1-182,
28 figs., 1 tab.
Menzies, Ropert J. 1954. A review of the systematics and ecology
of the genus “Exosphaeroma’, with the description of a new
genus, a new species, and a new subspecies (Crustacea,
Isopoda, Sphaeromidae). Am. Mus. Novitates, no. 1683,
pp. 1-24, 12 figs., 4 tabs.
1962. The zoogeography, ecology, and systematics of the
Chilean marine isopods. Repts. Lund Univ. Chile Exped.
1948-49, 42. Lunds Univ. Arsskrift. N.F., Avd. 2, vol. 57,
no. 11, pp. 1-162, 51 figs., 6 tabs.
Menzies, RoBert J., AND DirK FRANKENBERG. 1966. Handbook on
the common marine isopod Crustacea of Georgia. Univ.
Georgia Press, pp. viii + 1-93, 27 figs., 4 pls.
Menzies, Ropert J., AND PETER W. GiyNnn. In press. The common
marine isopod Crustacea of Puerto Rico, a handbook for
marine biologists. Stud. Fauna Curacao other Carib. Is.,
vol. 27, no. 104, pp. 1-133, 43 figs.
Monop, THeoporE. 1931. Tanaidacés et isopodes aquatiques de
l Afrique occidentale et septentrionale. (3° pt., 1) Sphaero-
matidae. Mem. Soc. Sci. Nat. Maroc, no. 29, pp. 1-91, 74
figs.
Moors, H. F. 1901. Report on Porto Rican Isopoda. U. S. Fish
Comm. Bull. 1900, vol. 2, pp. 161-176, 5 pls.
RICHARDSON, Harrier. 1905. A monograph on the isopods of North
America. Bull. U. S. Nat. Mus., no. 54, pp. liii + 1-727,
740 figs.
604 Proceedings of the Biological Society of Washington
Yi
Vol. 81, pp. 605-612 30 December 1968
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
A NEW CRAB OF THE GENUS CYCLOES (CRUSTACEA;
BRACHYURA; CALAPPIDAE) FROM SAINT HELENA,
SOUTH ATLANTIC OCEAN
By FENNER A. CHACE, Jr.
Smithsonian Institution, Washington, D. C.
Since publication of the report on Saint Helena collections
received between 1958 and 1964 (Chace, 1966), Arthur
Loveridge has continued to send material to the Smithsonian
Institution in the hope of enhancing our all too limited knowl-
edge of the marine fauna of that remote island. A recent ship-
ment contained the crab described below, by far the largest
and perhaps the most beautiful species of Cycloes De Haan
yet known; it rivals in size and coloration many of the more
familiar box crabs of the genus Calappa Weber.
It is a pleasure to name this species for the collector, Rev.
R. de Wet. I must reiterate my gratitude to Mr. Loveridge
for his continuing important contributions and also thank my
carcinological colleagues at the Smithsonian Institution, es-
pecially Henry B. Roberts, as well as Daniele Guinot of the
Muséum National d Histoire Naturelle in Paris, for helping to
solve systematic and bibliographic problems associated with
this description.
Cycloes deweti new species
Material: Male holotype (Smithsonian Institution, Division of
Crustacea cat. no. 122769) from off north coast of Saint Helena; 10-15
fm (18-27 m), in spiny lobster trap; 17 January 1968; Rev. R. de Wet.
Description: Carapace (Fig. 1) 1.12 times as broad as long in midline
(not including frontal projections), broadest anterior to level of lateral
teeth. Surface of carapace irregularly granulate everywhere except in
furrows; frontal and epigastric regions more finely and densely granulate
than remainder of surface. Mesogastric, protogastric, epigastric, hepatic,
and cardiac regions delimited by longitudinal or oblique furrows, those
bordering cardiac region most pronounced; gastric and cardiac regions
55—Proc. Biot. Soc. Wasu., Vou. 81, 1968 (605 )
606 Proceedings of the Biological Society of Washington
Fic. 1. Cycloes deweti, male holotype, x 0.56.
not separated; branchiat region irregularly ridged by five longitudinally
curved furrows subparallel to anterolateral margin; surface otherwise
slightly uneven, with several obscure elevations surmounted by clusters
of granules. Frontal projections (Fig. 2a) blunt and elongate, lateral
margins oblique, mesial margins subparallel, thereby forming distinctly
U-shaped median sinus. Dorsal margin of orbit sharply separated by
subrectangular bend from frontal margin, with distinct short fissure
slightly lateral to mid-width. Anterolateral margin bearing three or four
enlarged compound teeth in anterior half of length; posterior portion
rather uniformly dentate, but three or four regularly spaced teeth slightly
more upstanding than others. Lateral tooth prominent, sharp, and
directed dorsolaterally.
Each chela (Figs. 2c, d) bearing nine dentiform lobes dorsally,
proximal lobe widely bilobate, second obscurely so; extensor surface
with eight or nine low but distinct elevations in dorsal half, without
longitudinal ridges but with row of close-set tubercles extending hori-
zontally from proximoventral tooth about halfway to base of fixed
finger; most of extensor surface covered with scattered granules or
low tubercles becoming larger and more crowded ventrally but disap-
pearing near double row of tubercles along ventral margin. Stridulating
band on inner portion of extensor margin of dactyl of right chela (Fig.
2e) composed of 32 transverse ridges and tubercles. Merus of cheliped
bearing two large sharp teeth in line with proximoventral tooth on
chela; distoventral margin of merus with fringe of long hairs. Walking
legs smooth, each with fringe of long hairs on extensor margin of
New St. Helena Crab 607
merus; fringe continuing onto proximal half of carpus of two posterior
legs only; left posterior leg smaller than right, apparently regenerating.
Abdomen of unique male specimen (Fig. 2b) slightly distorted;
first two somites free, first with uninterrupted sinuous granular ridge
and fringe of hairs; second somite trilobate in profile, each lobe bearing
row of granules and fringe of hairs; third, fourth, and fifth somites
fused; fourth somite markedly narrower proximally than distal width
of third; median depression bounded by lateral elevations between third
and fourth somites; fifth somite more than twice as broad proximally
as long in midline; sixth somite free, probably about as long as broad
proximally if not distorted; telson sinuously triangular, less than one and
one-half times as long as broad.
First pleopod (Figs. 2h, i) elongate, sinuously tapered tube with
irregularly triangular opening at tip; fields of minute horny denticles
extending proximally from tip on mesial and lateral surfaces for less
than one-fifth of length of terminal segment of appendage. Second
pleopod (Figs. 2j, k) rather strongly convex toward midline in shorter
proximal portion; distal slender portion variably recurved, bent com-
pletely back on left side to form buttonhook with tip directed toward
base of appendage, less sharply bent on right side with tip directed
toward midline of sternum; endpiece armed with seven spinules on
anterior surface near tip; small bifid spine followed by two microscopic
spinules on mesial margin of appendage at juncture of proximal and
distal portions.
Color: After preservation in formalin for three months, carapace with
dark red vermiculate pattern of partially anastomosing lines and blotches
on pinkish gray or cream background, granules often white, especially
anteriorly. Chelipeds similarly colored dorsally, changing to irregular
isolated red bands and spots on white background on ventral half of
chelae and to plain white ventral to upper longitudinal row of tubercles;
flexor surface of chelae with irregular blotches of dark red on white
background, largest spots on palm proximodorsal to base of movable
finger and on proximal portion of movable finger. Walking legs white
with broad, irregular, longitudinal stripe of dark red on anterior and
posterior surfaces of carpus, propodus, and proximal portion of dactyl
of all legs; dark color progressively reduced on posterior surface of
merus from posterior to anterior walking legs; dark color present on
anterior surfaces of meri of two anterior pairs of walking legs but not
on meri of two posterior pairs. Pterygostomian region with diagonal
row of three red spots below orbit, diminishing in size laterally, and
interrupted row of dots and stripes on subhepatic region. Outer
maxilliped (Figs. 2f, g) with roughly trapezoidal red stripe visible
through pubescence on merus and additional colored areas visible when
appendage is denuded, especially on palp and exopod. Sternum with
red coloration in extreme anterior portion and paired spots on margins
of sternites bearing third and fourth pereiopods. Abdomen with paired
608 Proceedings of the Biological Society of Washington
Fic. 2. Cycloes deweti, male holotype. a, front. b, abdomen.
c, right chela. d, left chela. e, inner surface of dactyl of right chela.
f, right outer maxilliped. g, same, denuded. h, right first pleopod,
posterior view. i, tip of same. j, right second pleopod, anterior view.
k, tip of same. (b-d, x 0.6; e-g, x 1.2; a, h, 7, X 2.4; i, k, x 10.)
New St. Helena Crab 609
spots proximal to elevations between third and fourth somites, near
distal margins of fourth and fifth somites, near center of sixth somite,
and single median spots at distal end of sixth somite and near center
of telson.
Measurements: Overall length of carapace 72.7 mm; length in mid-
line 70.3 mm; maximum width 78.8 mm; width between lateral teeth
73.5 mm.
Range: Known only from the unique type-specimen from off Saint
Helena.
Remarks: As indicated in the key that follows, Cycloes deweti is
very similar to C. bairdii from American waters. It apparently differs
from that species in its much larger size, vermiculate rather than
spotted color pattern, sparser granulation, more strongly produced frontal
lobes, and broader abdomen, especially at the sixth somite. C. bairdii
is variable in all of these characters, as well as in the prominence of the
elevations on the carapace and chelae. Eastern Pacific specimens
commonly have more pronounced elevations than do those from the
western Atlantic, especially on the extensor surfaces of the chelae, where
they often form a carina-like row paralleling the ventral margin and
turning dorsally at the distal end near the base of the movable finger.
I have not seen Atlantic specimens with such a continuous ridge, but
the smoother form found in the Atlantic also occurs in the Pacific,
sometimes at the same locality as the more strongly sculptured variety,
as noted by Rathbun (1937, p. 229). The variety atlantica proposed
by Verrill (1908, p. 423) may prove to be a valid subspecies when
adequate series of both forms become available for systematic analysis,
but I doubt that it is specifically distinct.
If the Atlantic form represents a separate taxon, apparently the oldest
name available for it is Mursia balguerii Desbonne in Schramm, 1867.
Mme. Daniéle Guinot rediscovered the unique type-specimen of this
species in Desbonne’s collection of dried crabs in the Muséum National
d’Histoire Naturelle in Paris and graciously sent it to me for examina-
tion. It is an unusually smooth specimen of the species; the row of
close-set tubercles extending horizontally from the proximal tooth on
the extensor surface of the chela is obscure and barely distinguishable
from the scattered tubercles dorsal and ventral to it.
The largest specimen of C. bairdii mentioned by Rathbun (1937)
is a female with a carapace width of 49 mm from the Gulf of Panama.
The carapace fragment from Isla Socorro, Mexico, however, represents
a specimen that must have had a carapace width of about 53.5 mm.
The largest C. granulosa that I have seen is a male from Hawaii with a
carapace width of 49 mm. The only specimen of C. cristata examined,
a male syntype from the Canary Islands, has a carapace width of
32.3 mm.
It is possible that the characters here used to distinguish C. deweti
are directly related to size and that the Saint Helena specimen will
610 Proceedings of the Biological Society of Washington
eventually be found to represent only an unusually large C. bairdii.
For the present, however, it seems best to treat it as a distinct species
rather than to extend the size range of C. bairdii, perhaps erroneously,
by nearly 50 per cent and the geographic range by some 4,000 miles.
There has long been confusion about the valid name of the genus
of box crabs in which the carapace is not strongly produced laterally,
either as a large spine (Mursia Leach) or as an expansion over the
welking legs (Calappa Weber and Paracyclois Miers), or in which the
chelipeds are not armed with an unusually long meral spine (as in
Acanthocarpus Stimpson). Apparently both Cycloes De Haan and
Cryptosoma Brullé were proposed for this genus in the year 1837, the
former for an Indo-Pacific species, the latter for a crab from the
eastern Atlantic (see Monod, 1933, p. 40, and 1956, p. 114, footnote).
Which name has priority may never be determined satisfactorily, but
Cryptosoma was used ten years earlier for a genus of beetles. Although
Neave (1939) suggests that the beetle name may have been an incorrect
subsequent spelling of Cryptostoma and hence without nomenclatural
status, both Cryptostoma and Cryptosoma are apparently in current
use for coleopteran genera. It therefore seems best to reject the latter
name for the crab as a junior homonym and to recognize Cycloes as
the valid name of the genus.
KEY TO THE SPECIES OF Cycloes
1. Lateral tooth of carapace sharper but little if at all larger than
marginal teeth immediately anterior to it; stridulating band on
inner extensor margin of movable finger of right chela composed
of less than 20 ridges and tubercles; first pleopod of male straight
in distal portion, not sinuous, lateral margin of distal opening
Straig nity 2200 c0A0 in Nos TE ky pele C. granulosa De Haan, 1837.
India to Japan and Hawaii (see Sakai, 1965, p. 50, pl. 20, fig. 3
[as Cryptosoma granulosum] ).
Lateral tooth of carapace at least twice as large as marginal teeth
immediately anterior to it; stridulating band on inner extensor
margin of movable finger of right chela composed of more than
25 ridges and tubercles; first pleopod of male sinuous, lateral
Maren Of distal Opera SCO VC Kage eee 2
2. Carapace (including frontal projections) longer than broad, deeply
sculptured anterior to level of lateral teeth, elevations surmounted
by subconical granular bosses especially prominent on concentric
ridges on branchial regions; dorsal two-fifths of extensor surface
of chelae with similar conical bosses inclined dorsally _________
Fi REE NA COROT EOIN SN ae ee, Sd em ears C. cristata (Brullé, 1837).
Eastern Atlantic islands from the Madeiras to the Cape Verdes
(see Guinot-Dumortier & Dumortier, 1961, pp. 558-561, figs.
1-4).
Carapace broader than long, not deeply sculptured; extensor sur-
face of chelae without strong conical bosses. 3
New St. Helena Crab 611
3. Carapace densely granulate, more of surface covered with granules
than not; frontal projections not strongly produced, subtriangular,
median sinus broader than long; sixth abdominal somite longer in
midline than proximal width; color pattern spotted —..__________
1 NE AS GI ECTS aie) AL eA vee) C. bairdii Stimpson, 1860.
Eastern Pacific from Baja California to Ecuador, including the
Galapagos Island; western Atlantic from off North Carolina
and the Bermudas to Barbados and Isla de Providencia (see
Rathbun, 1937, p. 225, pl. 69, figs. 3, 4, and Garth, 1946, p.
362, pl. 62, figs. 7, 8).
Carapace sparsely granulate, more of surface without granules than
with, except on elevations; frontal projections strongly produced,
mesial margins subparallel distally, median sinus longer than
broad; sixth abdominal somite about as long in midline as wide
proximally; color pattern vermiculate, especially posteriorly __
Ab aa SNR WANS 1 (Sr ei REIRSON EN hE RL a C. deweti.
Saint Helena (see above).
LITERATURE CITED
BRULLE. 1837-1839. Crustacés. In Barker-Webb, P., et S. Berthelot,
Histoire naturelle des Iles Canaries, 2 (2) Entomologie: 13-
18, 1 pl.
Cuace, F. A., Jk. 1966. Decapod crustaceans from St. Helena Island,
South Atlantic. Proc. U. S. Nat. Mus., 118 (3536): 623—
661, 15 Figs.
Gartu, J. S. 1946. Littoral brachyuran fauna of the Galapagos
Archipelago. Allan Hancock Pacific Expeditions, 5 (10):
i-iv, 341-601, 1 Fig. in text, 39 pls.
Gurnot-DumortTiER, D., Er B. DuMortier. 1961. Description d’un
appareil stridulatoire dans le genre Cycloés De Haan
(Crustacea, Brachyura, Oxystomata, Calappidae). Bull. Mus.
Nat. Hist. Nat., Paris, ser. 2, 32 (6): 558-561, 4 Figs.
Haan, W. ve. 1833-1850. Crustacea. In Siebold, P. F. de, Fauna
Japonica. i—xvii, i-xxxi, ix—xvi, 1-243, 71 pls.
Monon, T. 1933. Sur quelques crustacés de l'Afrique Occidentale.
Bull. Com. Et. Hist. Sci. Afr. Occ. Fr., 15 (2-3): 1-93,
26 Figs.
1956. Hippidea et Brachyura ouest-africaines. Mém. Inst.
Fr. Afr. Noire, 45: 1-647, 884 Figs.
Neave, S. A. 1939. Nomenclator zoologicus, 1: i-xiv, 1—957.
Ratusun, M. J. 1937. The oxystomatous and allied crabs of America.
U. S. Nat. Mus. Bull. 166: i-vi, 1-278, 47 Figs. in text,
86 pls.
Saxar, T. 1965. The crabs of Sagami Bay. i-xviii, 1-206, 1-92,
1-32, 26 Figs. in text, 100 pls.
612 Proceedings of the Biological Society of Washington
ScHramMM, A. 1867. Crustaces de la Guadeloupe d’aprés un manuscript
du Docteur Isis Desbonne comparé avec les échantillons de
crustacés de sa collections et les derniéres publications de
Mm. Henri de Saussure et William Stimpson. i-ii, 1-60,
8 pls.
Stmmpson, W. 1860. Notes on North American Crustacea, in the
Museum of the Smithsonian Institution. No. II. Ann. Lyc.
Nat. Hist. New York, 7: 177-246, 2 pls.
VeRRILL, A. E. 1908. Decapod Crustacea of Bermuda; I. Brachyura
and Anomura. Trans. Connecticut Acad. Arts Sci. 13:
299-474, 67 Figs. in text, 20 pls.
Vol. 81, pp. 613-624 30 December 1968
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
TRIAKIS FEHLMANNI, A NEW. SHARK FROM THE
COAST OF SOMALIA
By STEWART SPRINGER
Bureau of Commercial Fisheries Ichthyological Laboratory
U. S. National Museum, Washington, D. C.
A 46-cm adult female specimen of a remarkable new shark
(Triakidae) collected off the coast of Somalia during the
International Indian Ocean Expedition (I.I.O.E.) is described
here and assigned to the genus Triakis.
Smith (1957) referred to species grouped by Bigelow and
Schroeder (1948) in Triakis as an uneasy assemblage. This
is an understatement in view of the remarkable diversity in
the group noted by Smith (1957), Kato (1968), and L. J. V.
Compagno (in correspondence). The addition of the new
species places more strain on currently used definitions of
genera and families of galeoid sharks and emphasizes the
need for review of characters that are traditional for their
recognition. For the purposes of this description, however, I
follow Bigelow and Schroeder (1948) with some exceptions
and make only those amendments required by the addition
of the new species.
The family Triakidae and the genus Triakis as provisionally
redefined here includes species with functional gill-rakers
as well as species without them, and also includes species with
as many as 7 cusps on lower jaw teeth toward the angles of
the jaws. The gill-rakers of the new species are not homologous
with denticle-derived structures in Cetorhinus and Rhincodon
that Bigelow and Schroeder evidently had in mind as typical
shark gill-rakers, but externally resemble those found in some
squaloid species (see Daniel, 1934, p. 154, fig. 147). In the
new species the structures are skin-covered, springy projections
56—Proc. Brot. Soc. WaAsH., Vou. 81, 1968 (613)
614 Proceedings of the Biological Society of Washington
extending and interdigitating across the inner branchial aper-
tures from both sides of the branchial bars. They are short
but presumably serve to make the branchial basket more
effective as a strainer of small food particles. Structures of
this kind are present but developed to varying degrees in
several small triakid and scyliorhinid species; in Triakis
barbouri ( Triakidae ) and in Halaelurus canescens ( Scyliorhini-
dae), for example, they are nearly as strong as in the new
species, whereas, in Cephalurus (Scyliorhinidae) they appear
as mere mounds of tissue or as short villi. These structures are
absent from the various species of Mustelus and from T.
scyllia, T. semifasciata, and T. maculata. Teeth toward the
angles of the lower jaw of the new species have seven or
more cusps, a condition that it shares with T. attenuata.
The new species is named in recognition of the important
contributions of Dr. H. Adair Fehlmann in setting high stan-
dards for field treatment of shark specimens collected for
study.
Triakis fehlmanni new species
(Figs. 1, 2, 3, 4, and 5C)
Holotype: A 46-cm sexually mature female, U.S.N.M. 202969, col-
lected on I.I.O.E. cruise 9 of the R/V Anton Bruun at station 463, 17
December 1964, in a trawl dragged at a depth from 70 to 170 m at
11° 24’ North Latitude, 51° 35’ East Longitude, southwest of Cape
Guardafui, Somalia.
Material examined: the holotype is the only specimen of the species
known. It was compared with examples of Triakis scyllia Muller and
Henle, 1841; T. semifasciata Girard, 1854; T. maculata Kner and Stein-
dachner, 1867; T. barbouri Bigelow and Schroeder, 1943; T. (Neotriakis)
sinuans J. L. B. Smith, 1957; T. (Eridacnis) radcliffei H. M. Smith, 1913;
T. venusta (= Calliscyllium venustum Tanaka 1912, probably also =
Proscyllium habereri Hilgendorf, 1904); T. henlei Gill, 1862; and several
species of Mustelus. I did not see specimens of T. attenuata Garrick,
1954 or T. acutipinna Kato, 1968, but both were given thorough and
modern descriptions and present no difficulty in diagnosis. Specimens
of Hemitriakis leucoperiptera Herre, 1923, were not available for
comparison. I agree, however, with L. J. V. Compagno (in correspon-
dence ) that it cannot be placed in Triakis.
Diagnosis: The distinctive color pattern of this species sets it apart
from all other sharks. It is also unique among galeoid sharks in having
615
New Shark from Somalia
“TOUBe AA “EY Arey Aq Burmviq,
‘adAjoroy a[vUloF wo
oF
“
sotdeds Mau
«
luubULyYyaf SLYDLLT.
‘T ‘ong
616 Proceedings of the Biological Society of Washington
"saat
Fic. 2. Camera lucida outlines of teeth of Triakis fehlmanni new
species. Upper row, teeth from upper left jaw (numbers below tooth
groups indicate number of tooth row counting from symphysis). Lower
row, teeth from lower left jaw.
cs by
large inner and outer nasal flaps that are irregularly scalloped along
their posterior margins.
T. fehlmanni resembles T. scyllia, T. semifasciata, T. maculata, and
T. acutipinnis in general body form, fin position, and fin shape as
viewed laterally, although the lower caudal lobe is not produced to
the extent that it is in adults of some of these species. It differs from
them in that it has a wider and more flattened head, a wider mouth
with smaller and more numerous teeth, very short labial furrows, short
but functional interdigitating gill-rakers, and a comparatively shallow
subocular gutter fully lined with denticles instead of a deep and in-
completely denticle-lined subocular pouch.
The similarity of the new species to the various species of Mustelus
in body form, and in fin shape and position is also striking. Further-
more, the large number of tooth rows in T. fehlmanni and their pattern
of tooth arrangement in quincunx with more functional rows near the
symphysis of the lower jaw are similar to that in species of Mustelus.
T. fehlmanni differs from species of Mustelus in that it has multicuspid
and sharp-pointed teeth, a wider head and mouth, shorter labial fur-
rows, and functional interdigitating gill-rakers.
The gill-rakers of T. fehlmanni have slender, pointed tips but arise
from wide bases, those of the anterior rim of one gill arch alternating
in position with those of the posterior rim of the adjacent gill arch.
Furthermore, they are as large and numerous around the anterior gill
apertures as around the posterior ones and are also present around the
spiracular pocket. In contrast, the gill-rakers of squaloid sharks, although
proportionally higher, arise from slender bases chiefly on the posterior
rims of gill arches and, if present, are usually larger on the posterior
arches or are confined to posterior arches as in some_ species of
Centrophorus. No observations were made on the comparative histology
of the gill-rakers of galeoid and squaloid sharks.
Triakis barbouri, T. sinuans, T. radcliffei and T. venusta have the
New Shark from Somalia 617
db VE
A B Cc
1mm
Fic. 3. Camera lucida outlines of 10th upper jaw tooth counting
from symphysis. A—anterior face. B—lateral aspect. C—posterior
aspect.
anal fin slightly in advance or nearly under the origin of the second
dorsal fin, whereas in T. fehlmanni the anal origin is appreciably pos-
terior to the origin of the second dorsal.
Triakis fehlmanni may be distinguished easily by its color and com-
paratively short body, from the plain-colored and elongate T. attenuata.
T. fehlmanni and T. attenuata are similar in that both have broad,
flattened heads, wide mouths with short labial furrows, and numerous
small teeth of somewhat similar shape arranged in similar patterns.
In both species the origin of the anal fin is posterior to the origin of
the second dorsal.
Description: Proportional dimensions of holotype are in percentages
of total length except as indicated. Measurements were made as out-
lined in Bigelow and Schroeder (1948).
Tip of snout to: front of mouth, 6.5; eye, 5.9; spiracle, 11.5; first
gill opening, 18.3; fifth gill opening, 22.8; origin pectoral, 22.0; origin
first dorsal, 33.7; origin pelvics, 43.0; origin second dorsal, 58.9; origin
anal, 61.7; origin upper caudal lobe 76.5; anterior end cloacal opening,
45.9.
Greatest width of: head, 13.7; trunk at pectorals, 10.0; trunk at
pelvics, 6.3; trunk at origin caudal 2.0.
Greatest height of: head at spiracles, 7.2; trunk at pectorals, 10.0;
trunk at pelvics, 8.3; trunk at origin caudal 3.0.
Eyes: horizontal diameter, 4.3; vertical diameter 1.7; distance be-
tween supraorbital rims, 3.9.
Spiracles: greatest diameter, 0.9; least distance from eye, 1.2; distance
between, 9.1.
618 Proceedings of the Biological Society of Washington
OBS.
OSs
ee oe
——EZSS
1mm
Fic. 4. Camera lucida outlines of dermal denticles from shoulder
area of Triakis fehlmanni new species.
Mouth: width, 9.3; length, 3.9; length upper labial furrow, 0.2;
length lower labial furrew, 1.1.
Nasal apertures: level of anterior ends to tip of snout, 3.7; level of
posterior ends to front of mouth, 0.7; minimum distance between, 3.5;
greatest length (diagonal), 3.3.
Gill slits: height of first 2.2; height of fifth, 2.0.
First dorsal fin: \ength base, 8.3; length posterior tip, 3.5; height,
7.2; length anterior margin, 9.8.
Second dorsal fin: length base, 9.3; length posterior tip, 2.8; height,
6.3; length anterior margin, 11.7.
Anal fin: length base, 7.0; length posterior tip, 2.6; height, 3.0;
length anterior margin, 7.0.
Pectoral fins: width base, 5.0; length anterior margin, 13.9; greatest
width, 10.4.
Pelvic fins: length (origin to rear tip), 9.8.
Caudal fin: upper margin, 23.0; anterior margin lower lobe, 8.9;
tip to notch, 5.9.
Distance between fin bases: first to second dorsal, 17.8; pectoral to
pelvic, 19.0; pelvic to anal, 11.7; anal to lower caudal, 6.5; second
dorsal to upper caudal, 8.3.
Posterior tips of fins to fin origins: pectorals (appressed) to pelvics,
7.0; pelvics to anal, 8.3; anal to lower caudal, 4.1; second dorsal to
upper caudal 5.4.
Teeth: upper, 86 rows; lower, 88 rows; width base, widest upper, 0.9
mm.; height, highest upper, 0.5 mm.; width, widest lower, 0.8 mm.;
height, highest lower, 0.5 mm.
New Shark from Somalia 619
1mm Imm 2mm i 2mm Smm
Fic. 5. Camera lucida outlines made from radiographs through the
longitudinal axis of a single trunk vertebra to show calcification pattern.
A—from Eridacnis radcliffei, a 25-cm female. B—from Triakis barbouri,
a 29-cm female. C—from T. fehlmanni new species, type, a 46-cm
female. D—from T. semifasciata, a 59-cm female. E—from Mustelus
canis, an adult female about 150 cm (stippled and dark areas indicate
calcification ).
Vertebrae: total number, 137; number “monospondylous,’ 39; num-
ber precaudal, 90; number caudal, 47; A-value (see Springer and
Garrick, 1964, p. 79), 128; B-value, 111; greatest diameter, 4.2 mm.
Gill-rakers: height of largest, 1.2 mm.; 6 or 7 on each side of ventral
arms and 2 or 3 on each side of dorsal arms of gill bars 2, 3, and 4.
A small species; 46-cm holotype is sexually mature; head long, very
wide, depressed; five gill slits, 4th over pectoral origin; abdominal
body segment short, approximately circular in cross-section at pectoral
level, trunk tapering toward tail, somewhat compressed posteriorly; no
precaudal pits or keels; caudal axis slightly but definitely elevated; no
nasal barbels or thickened areas on nasal flaps; nostrils not connected
with mouth by groove; length caudal more than 2 times length first
dorsal base; first dorsal base entirely in advance of origin pelvics.
Snout broadly rounded, slightly narrowed at nostrils, its length from
front mouth about 0.7 times width mouth; large nasal apertures in
posterior half snout, separated by about 0.5 times snout length; outer
and inner nasal flaps large, their posterior borders in apposition and
irregularly crenulate; pores of underside of snout moderately prominent,
forming Y-shaped pattern in single row with base near front of mouth.
Eyes large, about 2 times as long as high, their lengths about half
length of snout, situated well above rim of head, angle of view as much
dorsal as lateral, with a strong but shallow and fully denticle-lined
gutter below eye extending from anterior end of eye nearly to spiracle;
supraocular crest present, not extended over orbit as a shelf; interorbital
distance only about 55 percent of interocular distance; spiracle
moderately large, its height about half height of eye, located immediately
posterior to eye and separated from eye by a distance a little more than
greatest diameter of spiracle.
Mouth very wide, its width more than 2 times its length, about equal
620 Proceedings of the Biological Society of Washington
to length of second dorsal base; lower jaw sharply arched at symphysis;
labial furrows very short, length of lower labial furrow slightly less than
greatest diameter of spiracle and about 5 times length of upper; inside
of mouth light colored with one anterior row and 2 medial rows of low
papillae; no oral denticles; branchial basket large, a series of springy,
papilla-like gill rakers on both sides of each branchial bar interdigitating
across internal branchial apertures; 6 or 7 gill rakers, some scarcely
more than mounds of tissue, along each side of ventral section of
complete gill bars, 3 along each side of each dorsal section, smaller
numbers around spiracular pocket; branchial bars and internal surface
of pharynx without denticles.
Teeth small, numerous (83/86 rows); those of central part of both
jaws tricuspid, generally symmetrical, middle cusp much the longest,
slender, erect, pointed; lateral cusps short, most teeth wider than high;
series continuous across symphyses without marked change in tooth
shape or size; anterior faces of central teeth of both jaws with a series
of striations running from roots into enamel surfaces; the striations of
upper laterals reaching nearly to tips, those of lower teeth short; central
teeth of both jaws with posterior flange-like extension of roots (Fig. 3)
surrounding nutrient canal. Upper teeth in 4 to 5 functional series;
teeth tricuspid and nearly symmetrical in middle of jaw, generally lack-
ing an accessory cusp om side toward symphysis and main cusp oblique
from about 20th row (counting from symphysis) to jaw corners. Lower
jaw teeth in 5 to 7 functional series; functional area broader at
symphysis forming pavement-like occlusion surface of tricuspid symetri-
cal teeth; teeth transitional at about 20th row to 5 to 7 or more cusped
teeth, comb-like, cusps subequal, main cusp only slightly larger and
higher than others and located asymmetrically as the 5th, 6th, or 7th
cusp toward the jaw-corner side of each tooth.
Dermal denticles (Fig. 4) of shoulder area small, less than 0.5 mm,
only a little longer than broad; closely imbricate with 3 posteriorly-
directed short points, middle point longest; points little elevated, a
moderately strong central ridge with a weaker ridge on each side;
external surfaces uniformly covered, variation in denticle size and
shape similar to that in lower galeoids—that is, ventral denticles
generally without accessory points, ridges obsolete; denticle points some-
what longer toward tail; denticles of leading edges of fins and around
apertures smaller, smoother, and rounder than on flanks. No caudal
crest; skin relatively smooth to touch except when rubbed toward the
head; color pattern produced by skin chromatophores; denticles uniformly
semitransparent.
Pectorals broad, tips and inner corner broadly rounded, distal margins
straight, distance from tips of appressed pectorals to origin pelvics
about 1.5 times horizontal diameter of eye. Pelvics relatively small,
their outer contours broadly rounded, posterior tips rounded and not
attenuate; no apron connecting fins; origins slightly anterior to tip of
New Shark from Somalia 621
first dorsal but posterior to end of first dorsal base. First dorsal origin
about over free inner angle of pectorals, apex rounded, posterior point
sharp but short; slightly higher but about equal to second dorsal in area.
Distance between dorsal bases slightly less than distance from pectoral
axilla to pelvic origin. A depression or trench in the skin extending
between dorsal bases and continuing from second dorsal base to caudal
origin. Shape of second dorsal similar to that of first but slightly lower,
its base slightly longer; distance from its tip to origin upper caudal
about 1.5 times horizontal diameter of eye; end of its base about over
end of anal base; tip extending only slightly beyond anal tip. Anal
about 0.5 times as high as dorsals, its base shorter, its origin posterior
to second dorsal origin by distance about equal to vertical diameter of
eye. Caudal slightly less than 0.25 times total length, strongly notched
below near caudal tip; its lower anterior section not projecting as a
definite lobe.
Vertebral column comparatively slender; greatest diameter of mono-
spondylous vertebrae about 4.2 mm (about 6.7 mm in T. semifasciata
of equal total length); transition from monospondyly to diplospondyly
in pelvic region clearly marked, alternation of length of vertebrae not
apparent. Pattern of vertebral calcification (see Fig. 5) intermediate
in complexity between that of Triakis barbouri and T. semifasciata.
Visceral cavity short, its lining not pigmented; liver size moderate, 2
liver lobes well separated posteriorly, extending about 0.75 of distance
to cloaca; intestinal valve a spiral with about 10 turns; one ovary and
2, oviducts functional; nidamental glands small as compared to those of
most scyliorhinids such as Galeus of about equal total length. Holotype
has, in each oviduct, one very thin-walled, transparent, 45-mm by 17
mm egg-case enclosing an amorphous mass of egg-yolk material (no
embryos detected); single ovary appears to have much amorphous yolk
material except for several spherical yolk masses about 7 mm in diameter.
It is not clear from the condition of the specimen whether reproduction
is Oviparous or Ovoviviparous.
The holotype observed a few months after formalin preservation was
marked by a series of rich brown dorsal saddles interspersed by smaller
brown markings of bars and round or ovoid spots of varying size on a
ground color of light tan. Ventral surfaces were yellowish white with
some indistinct spots on the lower sides of the pectorals and caudal
trunk. Color changes after transfer to alcohol were not great.
Comments on natural history and relationships.
The stomach of the holotype contained only the partially digested
cephalothorax of one unidentified crustacean. The upper portion of the
valvular intestine contained some granular material (about 1 mm
diameter), possibly incompletely digested invertebrate shells.
The combination of large mouth, small teeth, and a large branchial
chamber with interdigitating gill-rakers suggests that this shark feeds
622 Proceedings of the Biological Society of Washington
on very small invertebrates. The presence or absence of gill-rakers
has rarely been noted in descriptions of sharks except for the kinds of
denticle-derived gill-rakers in the huge plankton-feeding whale sharks
and basking sharks. To see gill-rakers in most sharks it is necessary
to open the branchial chamber. This dissection is best accomplished
by cutting preserved and well-hardened specimens through the angle
of the jaws and gill arches on the right side back to the pelvic girdle.
I recommend this procedure as standard, at Ieast in first laboratory
examination of a shark species. Even with unique specimens such as
the type of T. fehlmanni no great harm is done and access is provided
for the search for additional identification characters sorely needed by
the shark taxonomist. A cut on the right side of the shark leaves intact
the left side preferred by illustrators.
T. fehlmanni resembles various species of scyliorhinids in its basic
color pattern, tooth shape, heterodonty, slender vertebral column, clear
and abrupt transition from monospondyly to diplospondyly, prominent
pore system on the ventral side of the snout, and degree of development
of a subocular gutter, as well as in its size at maturity and in its
scarcity from tropical continental shelf collections. All of these char-
acters as represented in T. fehlmanni point up the difficulty in the
separation of the families Scyliorhinidae and Triakidae. My own
separation of the two families is arbitrary. In the Triakidae the first
dorsal fin base is entirely in advance of the pelvic fins, whereas, in the
Scyliorhinidae the first or in one species the only dorsal fin, has some
part of its base over or posterior to the pelvic origin.
Tanaka (1915) reported that Triakis venusta (= Calliscyllium
venustum; probably also = Proscyllium habereri) is oviparous and con-
sequently should be placed in the Scyliorhinidae instead of in the
Triakidae where it has been left by most authors. He evidently re-
garded the character of egg laying as of overriding importance. Cadenat
(1959) reported, however, that of the two morphologically very similar
species, Galeus melastomus and G. polli, one lays eggs and the other
(G. polli) produces living young. By the arbitrary definition I have
used, both T. venusta and T. fehlmanni definitely fall in the Triakidae
as can easily be determined by external examination.
Several graded series of structural differences may be identified from
the small, usually slope-dwelling and demersal scyliorhinids though
the usually medium-size triakids that usually live in waters of the
continental shelf to the generally large shallow-water or surface-dwelling
carcharhinids. Among the differences are gradations from small to
large size; increase in the amount of calcification and strength of the
vertebral column; a shift from small multi-cusped teeth toward uni-
cuspid teeth, blade-like in the upper jaw; a development of the
nictitating lower eyelid through a series of stages accompanying loss
of the spiracle; a change from egg-laying to the development of large
young nourished in later stages of embryonic life by a pseudoplacenta;
New Shark from Somalia 623
a loss in the relative volume and perhaps also in complexity of the
Ampullae of Lorenzini; and an increase in the length of the clasper-
siphons accompanying a moderate lengthening of the claspers, a loss
of clasper hooks or reduction of clasper spines; and a change in fin
positions and shapes. Also, accompanying an adaptive radiation from
the kind of demersal life that characterizes the scyliorhinids to the kind
of life of the surface-dwelling carcharhinids, are improvements in
locomotion through changes in fin shape and position and through the
increased buoyancy provided by a relatively larger amount of liver oil.
Existing galeoid sharks provide a fine example of an evolutionary
series from which general trends can be demonstrated. Unforunately,
but as might be expected, the course of evolution of galeoids seems
to have proceeded with shifts in direction and rate for different organ
systems. The existing galeoids do not lend themselves to separation
neatly into families either by groups of characters or by marked dis-
continuities in the degree of specialization of one particular character.
Attempts to produce a phylogenetic classification through emphasis of
one character over another breaks down as more species and specimens
from the outer continental shelf and upper continental slopes become
available. The vertebral calcification pattern as used by White (1937),
for example, may be a better indicator of size at maturity or depth of
habitat than of phylogeny.
T. fehlmanni has many characteristics in common with scyliorhinid
species, perhaps more than with triakid species. My use of the position
of the first dorsal fin as the primary character for the separation of
members of the families Triakidae and the Scyliorhinidae has the
practical advantage that fin position is easily determined on any speci-
men and that, among presently known species, variational overlaps are
not known. So long as it is understood that galeoid families are grouped
for convenience in discussion and are not separated by conspicuous
gaps in character continuity no harm is done.
LITERATURE CITED
BicELow, Henry B. AND WILLIAM C. SCHROEDER. 1948. Sharks.
Fishes of the western North Atlantic. Sears Found. Mar.
Res., Mem. 1(1): 59-576.
CapENAT, J. 1959. Notes dichthyologie ouest-africaine—Galeus polli
espéce nouvelle ovovivipare de Scylliorhinidae. Bull. de
VI.F.A.N. 21(ser. A, no. 1): 395-409.
DANIEL, J. FRANK. 1934. The elasmobranch fishes. Univ. Cal. Press,
Berkeley, 1-332.
Garrick, J. A. F. 1954. A new species of Triakis (Selachii) from
New Zealand. Trans. Roy. Soc. New Zealand, 82(3):
695-702.
Herre, ALBERT W. C. T. 1923. Notes on Philippine sharks, I.
Philippine Jour. Sci., 23: 67-73.
624 Proceedings of the Biological Society of Washington
Kato, Susumu. 1968. Triakis acutipinna (Galeoidea, Triakidae), a
SmitTH, J.
new species of shark from Ecuador. Copeia, 1968, No. 2:
319-325.
L. B. 1957. A new shark from South Africa. South
African Jour. Sci., 53(10): 261-264.
SPRINGER, Vicror G. ANnp J. A. F. Garrick. 1964. A survey of
vertebral numbers in sharks. Proc. U. S. Nat. Mus., 116
(3496): 73-95.
TANAKA, SuHicGEHO. 1915. Calliscyllium venustum is oviparous. Zoologi-
cal Magazine, Tokyo. 27(323): 506-507. [In Japanese,
Trans. by Peter Miyake]
Waite, EF. Grace. 1937. Interrelationships of the elasmobranchs with
a key to the order Galea. Bull. Amer. Mus. Nat. Hist. 74(2):
25-138.
ys
Vol. 81, pp. 625-628 30 December 1968
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
A NEW SUBSPECIES OF WHITE-HANDED GIBBON
FROM NORTHERN THAILAND, HYLOBATES LAR
CARPENTERI NEW SUBSPECIES
By Coin P. GROVES
Primate Biology Program, Smithsonian Institution
16 Grafton Road, Enfield, Middx., England
A study of the gibbon specimens in the U.S. and British
collections has shown that an undescribed subspecies of the
white-handed gibbon, Hylobates lar Linnaeus, 1771, exists in
the Chiengmai and Loei districts of northern Thailand. This
form is by now well-known as to behaviour (Carpenter, 1940),
skeleton (Schultz, 1944) and dentition (Frisch, 1960); and
since gibbons from Thailand are finding their way in in-
creasing numbers into zoological gardens and Primate research
facilities in the United States, it seems important to describe
and define the new subspecies without delay.
Hylobates lar carpenteri new subspecies
Holotype: WHarvard, Museum of Comparative Zoology (MCZ) no.
41430, adult male skin, skull and skeleton; skin in dark phase. Mt.
Angka, N. Thailand, 3,400 ft.; collected by Asiatic Primate Expedition
of 1937.
Diagnosis: A subspecies of Hylobates lar with sharply distinct dark
and light colour phases, all individuals having a white ring of hair
round the face and the hands and feet white, sometimes as far as the
wrists and ankles; dark phase a very dark chocolate brown, the tips of
the hairs being blackish and their bases (up to one-half the total length
of the hair) silvery-brown; pale phase creamy-white, with the basal
one-quarter to one-third light grey. Hair on body much longer than
in other subspecies.
Distribution: Northern and part of northeastern Thailand.
Specimens seen: From Chiengmai district (Chieng Dao, Champee,
Kun Wang, Mt. Angka, Doi Nangkeo) 144 skins, skulls and skeletons
mostly in the museum of Comparative Zoology, Harvard, but a few
57—Proc. Biot. Soc. WasH., Vou. 81, 1968 (625)
626 Proceedings of the Biological Society of Washington
of the osteological specimens in the Anthropology Department, Uni-
versity of California at Berkeley; from Loei district (Ban Muang Khai,
Mt. Namlang, Mt. Phak Kinak, Mt. Lomlo, Ban Bo, Ban Na Muang)
16 skins and skulls in the U.S. National Museum; from Nakawu, one
skin and skull in American Museum of Natural History; from Khun Tan
mountains, one skin and skull in U.S. National Museum; from Siken,
near Korat, one skin and skull in U.S. National Museum (a somewhat
divergent specimen). In all, 163 skins and skulls, 144 skeletons.
Remarks: In the northwest, the distribution of this subspecies ap-
proaches the Burmese border; in the northeast, the Mekong river may
form the boundary. South of about 17°N, the river Mae Nam Ping
appears to form a boundary between this subspecies and the neighbour-
ing H. lar entelloides I. Geoffroy, 1842, but in the latitude of Chiengmai
it crosses the river at Mt. Angka and Doi Nangkeo, and _ possibly
intergrades with entelloides between these two localities and Myawadu
(16°40’N), the northernmost locality for the latter, although inter-
mediate specimens are lacking. In the southwest its distribution abuts
on that of Hylobates pileatus Gray, 1861, which according to field
observations kindly communicated by Dr. Gershon Berkson is a distinct
biological species. !
The new subspecies requires detailed comparison only with Hylobates
lar lar Linnaeus, 1771. and H. I. entelloides, occurring respectively in
Malaya and in the southwest Thailand and Tenasserim region. With
both of these it shares the sharply distinct colour phases, the white
hands and feet, and the complete white face-ring. In the dark phase,
however, H. I. lar is at once distinguished by its much lighter, medium
brown hue, with the basal half to two-thirds of the hairs light grey-
brown and the terminal portion yellow-brown; while in the dark phase
of H. I. entelloides the general colour is dark brown (not as dark as
carpenteri) with the basal portion very slightly lighter and greyer. In
the pale phase, H. |. lar is nearly white, less creamy than carpenteri
and lacking the grey basal portion; H. /. entelloides in light phase is
more honey-coloured with uniform tone on the individual hairs. Thus,
the new subspecies is nearer to its Thai neighbour in the dark phase,
but to the Malay form in the pale phase. In all three, the dark and
pale phases are sharply distinct, without intermediates, bearing no
relation to sex and not undergoing change during the life cycle (in
contradistinction to H. pileatus); in this sense the term “phase” is
unfortunate for H. lar, though not inappropriate for certain other species,
such as pileatus. The significance of dichromatism in gibbon popula-
tions is quite unknown.
The new subspecies may equally be distinguished by its much longer
hair. In both H. I. lar and entelloides the hair length between the
shoulders varies from 29-56 mm., but in H. |. carpenteri 79-103 mm.
The only specimen of carpenteri without this character is the specimen
from Siken (USNM no. 241423), in which the hair is approximately
New Subspecies of Gibbon 627
55 mm. long; this locality is in the lowlands of east-central Thailand;
other specimens were collected in montane areas, up to 8,000 ft. on Mt.
Lomlo and from 3,400 to 5,700 ft. on Mt. Angka. The other subspecies
of H. lar may equally inhabit mountainous areas; but the only sub-
species with hair-length approaching the new form is the Javan H. l.
moloch Audebert, 1800, which the hair length between the shoulders
measures 50-70 mm.
The newly described subspecies is named in honour of C. R. Car-
penter, whose behaviour study of gibbons in the Chiengmai area re-
mains a classic in the field of Primate behaviour after nearly thirty
years. The 144 Harvard specimens of H. l. carpenteri were collected
on the Asiatic Primate Expedition, in the course of which Dr. Carpenter’s
behaviour study was carried out.
LITERATURE CITED
CaRPENTER, C. R. 1940. A field study in Siam of the behaviour and
social relations of the gibbon. Comp. Psychol. Monogr.
16, no. 5.
Frisco, J. E. 1960. Dental variability in a population of gibbons
(Hylobates lar). In Brothwell, ed., Dental Anthropology,
pp. 15-28.
Scuuttz, A. H. 1944. Age changes and variability in gibbons. Am.
J. phys. Anthrop. 26: 389-408.
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Vol. 81, pp. 629-634 30 December 1968
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
A NEW PRIMATE FROM THE TORREJON MIDDLE
PALEOCENE OF THE SAN JUAN BASIN, NEW MEXICO
By C. Lewis GAzIN
Smithsonian Institution, Washington, D. C.
Recent review of Paleocene materials from the San Juan
Basin of New Mexico, that had been set aside for further
study, disclosed the presence of certain forms new to the
Torrejon fauna. One of these is a moderately large primate,
which seems, among other Paleocene torms, to be most closely
related to Paromomys, first described by Gidley (1923) from
the Fort Union beds in Montana, and further investigated by
Simpson (1937 and 1955).
Primates had not been known from the Torrejon beds of
the Nacimiento formation in New Mexico until reported from
the Angel Peak basin by R. W. Wilson (1951 and 1956).
These were cited (1956) only as “Palaechthon?, n. sp.” and
“phenacolemurid, n. gen. and sp.” without description, but
mentioned as found in the Deltatherium or lower zone of the
Torrejon. Primate remains found by Franklin Pearce and
myself in 1949 were obtained from the upper or Pantolambda
zone in the Arroyo Torrejon, essentially the type section or
area for the middle Paleocene. I have not examined the
Angel Peak materials but those from the Arroyo Torrejon
would not be referred to Palaechthon. Only the single speci-
men cited as “phenacolemurid” remains in doubt as to the
possibility of its representing the form described below.
FamMiLy PAROMOMYIDAE Simpson, 1940
Torrejonia new genus
Type: Torrejonia wilsoni! new species.
Generic characters: Close to Paromomys but lower premolars more
1 Named for Robert W. Wilson who was first to recognize primates in the
Torrejon beds.
58—Proc. Biot. Soc. WasuH., Vou. 81, 1968 (629)
630 Proceedings of the Biological Society of Washington
elongate anteroposteriorly. P. with relatively narrower and more elongate
talonid. Anterior lower molars with moderately elevated trigonids with
essentially three low transverse crests defining two very small basins.
Posterior wall of trigonids relatively flat and erect. Talonid basins of
anterior lower molars large, relatively deep and enclosed. Weak
hypoconulid formed by flexure of posterior crest of M:, and _ slight
medial elevation of posterior crest on Me. External cingular crest de-
veloped about protoconid, extending lingually well below anterior crest
of trigonid, very subdued posteriorly.
Discussion: The dental formula for Torrejonia is not known, but it
is evident that there were at least three premolars, as well as three molars,
and in all probability the formula corresponded to that for Paromomys,
not as in Phenacolemur. Paromomys had an enlarged lower incisor,
not nearly so procumbent as in Phenacolemur, followed by a moderately
large single rooted tooth, presumably the canine, and a two rooted
second premolar. In Torrejonia segments of two alveoli are seen anterior
to P;. Possibly these are both of Ps. There may have been a large
incisor as in Paromomys but there is no incisor alveolus beneath the
posterior premolars as in Phenacolemur.
P; is relatively much larger and more elongate anteroposteriorly than
in Paromomys, and its greatest width is slightly more forward. This
tooth in both forms shews a sharp anterior crest above, which part way
down turns lingually and disappears. There is no definable parastylid on
either. Both show a narrow, somewhat flattened posterior wall with a
well defined posterolateral crest and a slight talonid. The talonid
is relatively a little narrower in Torrejonia.
P: in Torrejonia is also relatively larger and more elongate than in
Paromomys, and the greatest width is well forward, at the primary cusp
rather than essentially across the talonid. In both forms the anterior
crest, as in Ps, is sharp above and at its anterior extremity turns
abruptly inward and disappears, but without a definable parastylid.
The talonid of Ps is a little larger and more deeply basined than in
Paromomys, but shows a similar development of the posteroexternal
(hypoconid ) and posterointernal (entoconid) cusps. The posteroexternal
cusp joins the well defined posteroexternal crest on the primary cusp.
This crest is not so medially placed as in Paromomys. The posterointernal
crest of the primary cusp in Torrejonia is double in its upper part but,
as in Paromomys, there is no evidence of a metaconid, such as seen in
Palaechthon, or still better developed in Plesiolestes.
The anterior molars M: and Mz strongly resemble these teeth in
Paromomys but M: is a little longer with respect to Mz than in
Paromomys. The paraconid of M:; is conical, placed high and well
forward. The trigonid of Ms is more compressed anteroposteriorly
with the paraconid small and close to the metaconid. The posterior or
third small crest of the trigonid in both molars is better defined in
Torrejonia and the posterior wall of the trigonid is somewhat flatter, or
New Primate from New Mexico 631
less inflected, and appears more erect. An external cingular crest, not
seen on the premolars, is developed around the protoconid and carried
across the anterior surface of the molars well below the anterior crest
of the trigonid, relatively a little lower with respect to this crest than
in Paromomys. The lingual walls of the teeth are without a cingular
crest.
The large talonid basins of the molars are deeper and more elongate
than in Paromomys, in which respect there is a somewhat stronger
resemblance to Omomys. A certain rugosity or folding of the enamel
in the walls of the basin essentially matches in detail that in Paromomys,
but is less emphasized. The hypoconid is prominent with strong oblique
crests, the anterior showing a very small cuspule near its extremity with
a slight notch separating it from the posterior wall of the trigonid.
The entoconid is a little lower than the hypoconid and more smoothly
crescentic. There is no conical hypoconulid but its position on Miz is
indicated by a flexure of the posterior rim of the talonid basin where
it is joined by the posterolingual extremity of the decidedly weak
posterior portion of the external cingular crest. On Mz this position is
denoted by a slightly raised portion of the posterior rim, which as
in M: is a little more elevated than in Paromomys.
Torrejonia makes an approach toward Palaechthon in the relatively
slender, elongate P:, but lacks the parastylid and metaconid exhibited
by the latter. Moreover, the narrow talonid of Ps in Palaechthon is less
basined but developed more as a high transverse crest posteriorly. Ps is
relatively very much smaller in Palaechthon. Resemblance in the lower
molars is seen in the flat, relatively erect posterior wall of the trigonids
and in the large, deeply basined talonids, but the trigonids in Palaechthon
are relatively much more elevated with higher cusps and the talonid
basins are less well closed lingually.
An isolated upper molar (fig. 2) believed to represent Torrejonia
is of a size comparable to the molars in the lower jaw. Its form
suggests the second rather than the first of the molar series, but this
is uncertain. The tooth appears relatively short anteroposteriorly in
comparison with its striking width. The outer cusps are conical and
moderately high, much as in Paromomys, but perhaps a little better
separated. The talon, however, shows a larger protocone, which would
correspond to the larger talonid basins of the lower molars in Torrejonia.
As in Palaechthon, the protocone is more medially placed, fore and aft,
than in Paromomys. There are three crests extending laterally from
the protocone, the median as a well defined rib toward the broad
central basin, and the others toward the accessory cuspules. The small
accessory cuspules are sharply distinct from the crests of the protocone,
as well as from the outer cusps. The metaconule, though no larger
than the protoconule, is, as in Palaechthon, much better defined than
in Paromomys. The cingulum externally is essentially as in Paromomys,
but is much weaker on the anterior wall of the tooth. Posteriorly the
632 Proceedings of the Biological Society of Washington
cingulum is well developed but not so widely separated from the crest
between the protocone and metaconule. On the posterior slope of the
protocone the lingual extremity of the cingulum turns toward the apex
of the protocone but does not actually reach it. In Paromomys the
protocone is characterized by a prominent, elongate crest from the apex
to the lingual extremity of the posterior cingulum. This crest is relatively
much shorter in Palaechthon. The lingual wall of the Torrejon molar
is only slightly bilobed and, as in Palaechthon, is not nearly so askew.
Torrejonia wilsoni new species
(Figs. 1 and 2)
Type: Left ramus of mandible, USNM No. 25255, including Ps-M2.
Horizon and locality: Upper fossiliferous zone of the middle Paleocene
Torrejon beds, Nacimiento formation. East branch of the Arroyo
Torrejon, San Juan Basin, New Mexico.
Specific characters: Size appreciably larger than Paromomys maturus
Gidley (1932), much larger than the paromomyids Palaechthon alti-
cuspis Gidley (1923) and Plesiolestes problematicus Jepsen (1930). A
close approach is made in size of molars, particularly M2, with the
plesiadapid Pronothodectes simpsoni Gazin (1956) from the early Tif-
fanian of the Bison Basin, Wyoming. Other characters of specific im-
portance not recognized.
Discussion: In addition to the isolated upper molar, USNM No.
25256 (fig. 2) discussed above, there is also an isolated Mz, USNM
No. 25257, referred to this species. Both are from the upper level of
the Torrejon in the west branch of the Arroyo Torrejon. The isolated
Mz is relatively unworn and slightly larger than this tooth in the type
lower jaw. The grooves or rugosity on the lingual slope of the hypoconid
are better developed than in the type and are also evident on the labial
slope of the entoconid. There is, moreover, better evidence of a
cuspule on the crista obliqua, but less evidence of an hypoconulid. The
outer cingular crest is perhaps a little better defined posteriorly.
Measurements (in mm.) of lower teeth in T. wilsoni (USNM No.
25255, type) in comparison with those for a specimen of P. maturus
(U.S.N.M. No. 9292).
trigonid talonid
length width width
P;, T. wilsoni : P. maturus 2.6:1.8 ids) 9 18} —:—
P,, T. wilsoni : P. maturus 3.3 : 2.8 1.9 ;:— 1.8: 1.8
M,, T. wilsoni : P. maturus 3.5 : 3.0 2.5 > 2.0 Ds) 8 PAD)
M2, T. wilsoni : P. maturus 3.3: 3.1 2.6 : 2.2 SY 8 P58
The isolated Mz of T. wilsoni (USNM No. 25257) is 3.5 mm. long
by 2.7 mm. wide at the trigonid and 2.9 mm. across the talonid. The
isolated upper molar (USNM No. 25256) believed to be T. wilsoni
New Primate from New Mexico 633
Fics. 1-2. Torrejonia wilsoni, new genus and species. 1, left ramus
of mandible (U.S.N.M. No. 25255), with Ps-Me, type specimen, occlusal
and lateral views (x4). 2, right upper molar (U.S.N.M. No. 25256),
occlusal view (4).
Fic. 3. Paromomys maturus Gidley. Left ramus of mandible
(U.S.N.M. No. 9292), occlusal and lateral views (x4).
Drawings by Lawrence B. Isham, scientific illustrator, Department of
Paleobiology, U. S. National Museum.
634 Proceedings of the Biological Society of Washington
is 2.8 mm. long anteroposteriorly across the external portion and 2.5
mm. long across the midsection. Its greatest transverse diameter per-
pendicular to the outer wall is 4.7 mm.
LITERATURE CITED
Gazin, C. L. 1956. Paleocene mammalian faunas of the Bison Basin
in south-central Wyoming. Smithsonian Misc. Coll., vol. 131,
no. 6, pp. 1-57, 2 figs., pls. 1-16.
Gwotey, J. W. 1923. Paleocene primates of the Fort Union, with
discussion of relationships of Eocene primates. Proc. U. S.
Nat. Mus., vol. 63, art. 1, pp. 1-38, pls. 1-5.
Jepsen, G. L. 1930. Stratigraphy and paleontology of the Paleocene
of northeastern Park County, Wyoming. Proc. Amer. Philos.
Soc., vol. 69, no. 7, pp. 463-528, pls. 1-10.
Smepson, G. G. 1937. The Fort Union of the Crazy Mountain field,
Montana, and its mammalian faunas. U. S. Nat. Mus. Bull.
169, pp. i-x, 1-287, figs. 1-80, pls. 1-10.
1940. Studies on the earliest primates. Bull. Amer. Mus.
Nat. Hist., vol. 77, art. 4, pp. 185-212, figs. 1-8.
1955. The Phenacolemuridae, new family of early primates.
Bull. Amer. Mus. Nat. Hist., vol. 105, art. 5, pp. 411-441,
pls. 30-35, tables 1-6.
Witson, R. W. 1951. Preliminary survey of a Paleocene faunule
from the Angels Peak area, New Mexico. Univ. Kans. Publ.,
Mus. Nat. Hist., vol. 5, no. 1, pp. 1-11, 1 fig.
1956. A new multituberculate from the Paleocene Torrejon
fauna of New Mexico. Trans. Kans. Acad. Sci., vol. 59, no.
1, pp. 76-84, figs. 1, la. 1 table.
Vol. 81, pp. 635-692 30 December 1968
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
CYCLOPOID COPEPODS OF THE GENUS
LICHOMOLGUS ASSOCIATED WITH OCTOCORALS OF
THE FAMILY ALCYONITDAE IN MADAGASCAR
By Artuur G. HuMEs AnD Ju-SHEY Ho
Dept. of Biology, Boston University, Boston, Mass.
Three species of Lichomolgus are already known to be as-
sociated with octocorals of the family Alcyoniidae in Mada-
gascar. These are L. decorus Humes and Frost, 1964, from
Cladiella laciniosa (Tixier-Durivault), L. squamiger Humes
and Frost, 1964, from Sinularia polydactyla (Ehrenberg), and
L. protentus Humes and Frost, 1964, from Sarcophyton
globosum Tixier-Durivault. The last named host was listed by
Humes and Frost (1964) as Sarcophyton sp., but has since
been described as new by Tixier-Durivault (1966). This paper
concerns seven new species of Lichomolgus, two species which
are redescribed, and a new host record for L. squamiger, all
washed from various species of Alcyoniidae in the region of
Nosy Bé in northwestern Madagascar.
All collections were made by A. G. Humes, those in 1960
during an expedition sponsored by the Academy of Natural
Sciences of Philadelphia, and those in 1963-64 as part of the
U.S. Program in Biology of the International Indian Ocean
Expedition.
The study of the specimens has been aided by a grant
(GB-5838) from the National Science Foundation of the
United States.
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. The abbreviations used are:
A, = first antenna, As = second antenna, MXPD = maxilliped,
and P; = leg 1.
59—Proc. Biou. Soc. WAsH., Vou. 81, 1968 (635)
636 Proceedings of the Biological Society of Washington
All descriptions are based on type material. The measure-
ments of the length of the body have been made in all cases
from specimens in lactic acid and do not include the setae
on the caudal rami. The lengths of the segments of the first
antenna have been measured along their posterior non-
setiferous margins.
We are indebted to Mme. A. Tixier-Durivault of the Muséum
National d’Histoire Naturelle, Paris, for the identifications of
the octocorals collected in 1960, and to Dr. J. Verseveldt,
Zwolle, The Netherlands, for the determinations of those col-
lected in 1963-64. We thank Dr. John O. Corliss of the Uni-
versity of Illinois for the identification of the suctorian
attached to L. singularipes.
The new copepods described in this paper comprise the
following:
1) Lichomolgus cristatus new species
from Sinularia leptoclados (Ehrenberg).
2) Lichomolgus adelphus new species
from Sinularia whiteleggei Liittschwager, S. pedun-
culata Tixier-Durivault, and S. polydactyla
(Ehrenberg ).
3) Lichomolgus hetaericus new species
from Cladiella pachyclados (Klunzinger) and C.
krempfi Hickson.
4) Lichomolgus insolens new species
from Lobophytum crassum Marenzeller.
5) Lichomolgus spathophorus new species
from Sarcophyton glaucum (Quoy and Gaimard).
6) Lichomolgus incisus new species
from Sarcophyton ehrenbergi Marenzeller.
7) Lichomolgus singularipes new species
from Parerythropodium rubiginosum Verseveldt.
Of the following two species, females are redescribed and
males described for the first time:
1) Lichomolgus dentipes Thompson and A. Scott, 1903
from Sinularia humesi Verseveldt.
2) Lichomolgus foxi Gurney, 1927
from Cladiella krempfi Hickson, C. laciniosa
Copepods Associated with Octocorals 637
( Tixier-Durivault ), and C. pachyclados (Klun-
zinger ).
The following is a new host record:
1) Lichomolgus squamiger Humes and Frost, 1964
from Sinularia whiteleggei Liittschwager.
FAMILY LICHOMOLGIDAE KOSSMANN, 1877
Genus LICHOMOLGUS THORELL, 18591
Lichomolgus dentipes Thompson and A. Scott, 1903
Figures 1-28
This species was established on the basis of one female found in
washings of dredged invertebrates in Ceylon. It has not been reported
again, and the original specimen is presumably lost along with the rest
of the Andrew Scott collection (see Humes and Ho, 1967, p. 209).
The male has been unknown until now.
Both sexes of L. dentipes (872 2 and 1456 6) were washed from
one colony of Sinularia humesi Verseveldt, collected in 2 m, off
Ampombilava, Nosy Bé, Madagascar, 26 September 1964. One hundred
and fifty-six specimens (549 2, 102¢ ¢) have been deposited in the
United States National Museum; 35 specimens (152 2, 204 6) in the
Zodlogisch Museum, Amsterdam; and the remaining specimens in the
collection of A. G. Humes.
Female: Body (fig. 1) with rather broad prosome. Length 0.94
mm (0.91-0.95 mm) and greatest width 0.57 mm (0.56-0.58 mm),
based on 10 specimens. Ratio of length to width of prosome 1.14: 1.
Segments of legs 1-3 with irregularly serrate hyaline lateral margins.
Segment of leg 4 narrow and mostly concealed beneath dorsum of
preceding segment; terminating posteriorly on each side in a hyaline
spiniform process.
Segment of leg 5 (fig. 2) 78 u xX 187 w. Genital segment wider than
long, 100 « X 148 u, in dorsal view rounded laterally and abruptly
constricted in its posterior fifth. Areas of attachment of egg sacs situated
dorsolaterally on posterior part of expanded region. Each area (fig. 3)
bearing two naked spiniform setae 7 uw and 11 » long, with a recurved
sclerotized pointed process between them. Three postgenital segments
39 uw X 78 pw, 31 uw x 73 uw, and 65 uw xX 70 uw, from anterior to posterior.
Caudal ramus (fig. 4) slightly longer than wide, its greatest dimen-
1The year of publication of Thorell’s genus has been cited by some authors
as 1859, by others as 1860. The source of this confusion lies in the fact that
Thorell published Lichomolgus as a new generic name in two different papers
(1859, p. 340, and 1860, p. 64). Although the title page of Arg. 16 of the
Ofversigt af Kongl. Vetenskaps-Akademiens Foérhandlingar for 1859 bears the
year 1860, the actual year of publication for Lichomolgus appears to be 1859,
since the last page of no. 8 (in which Thorell’s paper appears) bears that year.
638 Proceedings of the Biological Society of Washington
A 0.2MM °
Cc 0.02 MM
Fics. 1-5. Lichomolgus dentipes Thompson and A. Scott, 1903, fe-
male: 1, body, dorsal (A); 2, urosome, dorsal (B); 3, area of attachment
of egg sac, dorsal (C); 4, caudal ramus, dorsal (C); 5, rostral area,
ventral (B).
sions being 33 u X 26 uw. Outer lateral seta 50 u, pedicellate dorsal seta
40 uw, outermost distal seta 55 wu, innermost distal seta 109 mw, and the
two long median terminal setae 220 » (outer) and 360 uw (inner) and
both inserted between dorsal (unornamented ) and ventral ( with marginal
row of spinules) flaps. All these setae with lateral hairs except outer
lateral seta which is naked.
Copepods Associated with Octocorals 639
0.1 MM
D
F 0.05 MM
Hee 6-12. (rea dentipes Thompson and A. Scott, 1903,
female: 6, first antenna, ventral (D); 7, second antenna, anterior
(inner) (D); 8, labrum, ventral (E); 9, mandible, posterior (C); 10,
paragnath, ventral (C); 11, first maxilla, posterior (C); 12, second
maxilla, posterior (E).
Ratio of length of prosome to that of urosome 2.24 : 1.
Egg sacs incomplete on females examined. Each egg about 55 wu in
diameter.
Rostral area (fig. 5) without well defined posterior margin.
First antenna (fig. 6) 7-segmented, 326 uw long. Lengths of segments:
640 Proceedings of the Biological Society of Washington
i}
HU
dl SS
“Ses
SSS SS is
cS
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a
Ze
Fics. 13-17. Lichomolgus dentipes Thompson and A. Scott, 1903,
female: 13, maxilliped, posterior (EF); 14, area between maxillipeds and
first pair of legs, ventral (D); 15, leg 1 and intercoxal plate, anterior
(F); 16, leg 2 and intercoxal plate, anterior (F); 17, leg 3 and inter-
coxal plate, anterior (F).
30 » (58 uw along anterior margin), 138 uw, 21 », 50 uw, 23 uw, 19 pw, and
17 uw respectively. Formula for armature: 4, 13, 6, 3, 4+ 1 aesthete,
2-+ 1 aesthete, and 7 + 1 aesthete.
Second antenna (fig. 7) 4-segmented, last segment moderately
elongated, 71 uw along its outer edge, 40 uw along its inner edge, bearing
Copepods Associated with Octocorals 641
UK Ss Ze e,
SS. i
Fics. 18-22. Lichomolgus dentipes Thompson and A. Scott, 1903,
female: 18, leg 4 and intercoxal plate, anterior (F); 19, endopod of
leg 4, anterior (C); 20, leg 5, dorsal (E). Male: 21, body, dorsal
(G); 22, urosome, dorsal (D).
distally five small hyaline elements and a single claw 44 w along its
greatest axis.
Labrum (fig. 8) with two broad posteroventral lobes. Mandible
(fig. 9) with flagellum very reduced, and represented only by a small
pointed process. Paragnath (fig. 10) a small hairy lobe. First maxilla
Fics. 23-28. Lichomolgus dentipes Thompson and A. Scott, 1903,
male: 23, caudal ramus, dorsal (H); 24, first antenna, dorsal (F); 25,
second antenna, posterior (outer) (EF); 26, maxilliped, outer (F); 27,
endopod of leg 1, anterior (C); 28, leg 5, dorsal (H).
(fig. 11) with two long unequal naked terminal elements and a small
subterminal one. Second maxilla (fig. 12) 2-segmented. Maxilliped
(fig. 13) 3-segmented, the second segment with two very unequal
setae, and the third with two terminal spiniform elements (one lacking
a distinct articulation) and a small seta. Area between maxillipeds and
first pair of legs (fig. 14) not protuberant.
Copepods Associated with Octocorals 643
Legs 1-4 (figs. 15-18) with trimerous rami except for 2-segmented
endopod of leg 4. Armature of legs as follows (Roman numerals =
spines, Arabic numerals = setae):
P: protopod 0-1 1-0 exp I[I-O I-1 IILI,4
end OQ-l1 O-1 I,5
P2 protopod 0-1 1-0 exp IO I[-1 IILI5
end 0-1 0-2 L,]I,3
P; protopod 0-1 1-0 exp I-O [1 MIILJ,5
end 0-1 0-2 LII,2
P: protopod O0-1 1-0 exp I-0 I[-1 MIII,I5
end Q-1 II
Inner seta on coxa of leg 4 very short (6 uw), blunt, and naked.
Outer seta on basis of legs 3 and 4 unusually long (up to 90 wz).
Terminal spine on last segment of endopod of leg 1 a little longer than
the segment, naked, and recurved. Endopod of leg 4 (fig. 19) slender,
first segment 22 u X 9 uw, with inner distal seta 33 wu, second segment
47 uw long (including processes) and 7 u wide at middle, with the two
terminal spines 18 u (outer) and 36 uw (inner). Hairs along outer margin
of second segment arranged in two rows.
Leg 5 (fig. 20) with free segment 74 wu long, its proximal area ex-
tended obliquely and inwardly to form a very large toothlike process.
Oblique length from outer base of segment to tip of process 67 u. Two
terminal naked setae 34 w and 37 uw. Row of spinules along outer edge
of segment.
Leg 6 probably represented by the two spiniform setae near areas of
attachment of each egg sac (see fig. 3).
Color in life in transmitted light translucid, eye red, egg sacs light
gray.
Male: Body (fig. 21) with prosome less broadened than in the
female. Length 0.58 mm (0.56-0.60 mm) and greatest width 0.28
mm (0.26—0.30 mm), based on 10 specimens. Ratio of length to width
of prosome 1.27: 1.
Segment of leg 5 (fig. 22) 21 uw xX 81 uw. Genital segment rounded in
dorsal view, 122 uw X 133 uw, with a small constricted area posteriorly.
First postgenital segment 15 uw x 46 uw, second 14 uw xX 41 uw. Third and
fourth postgenital segments fused, third probably represented by the
anterior constricted part, 11 «xX 35 w, and fourth being the much
broadened posterior region, 39 u X 55 wu.
Caudal ramus (fig. 23) about as long as wide, 17 u x 19 u.
Ratio of length of prosome to that of urosome 1.6: 1.
Rostral area like that of female. First antenna (fig. 24) similar to that
of female, but two aesthetes added on second segment, and one on
third, so that the formula is 4, 13+ 2 aesthetes, 6, 3+ 1 aesthete,
4+-1 aesthete, 2+ 1 aesthete, and 7+ 1 aesthete. Second antenna
(fig. 25) resembling that of female, but inner surface of second
segment with a row of spinules and a raised membranous lamella.
644 Proceedings of the Biological Society of Washington
Labrum, mandible, paragnath, first maxilla, and second maxilla like
those of female. Maxilliped (fig. 26) slender and 4-segmented (assuming
that the proximal part of claw represents a fourth segment). Claw
124 uw long (measured along its axis). Area between maxillipeds and
first pair of legs like that in female.
Legs 1-4 segmented as in female and with same spine and setal
formula except for endopod of leg 1 (fig. 27), which has arrangement
of 0-1; 0-1; 11,4. Last segment of endopod of leg 1 bearing two
terminal barbed spines, with a slender process between them. Endopod
of leg 4 as in female.
Leg 5 (fig. 28) with a slender free segment 27 uw X 6 wu, lacking the
large toothlike process seen in the female. Two terminal setae 24 uw and
22 uw.
Leg 6 (see fig. 22) a posterolateral flap on ventral surface of genital
segment bearing two naked setae 17 uw and 22 wu long.
Spermatophore not observed.
Color in life like that of female.
Remarks: The brief original description of the female of L. dentipes
by Thompson and A. Scott (1903, p. 281, pl. XVI, figs. 27-30), based
on one female, fits almost exactly the specimens from Sinularia. Although
it is impossible to compare them with the single Ceylonese female (which
no longer exists), the specimens from Madagascar appear to be identical
with L. dentipes. Probably Thompson and A. Scott’s specimen, ob-
tained in washings of dredged invertebrates, actually came from an
octocoral, perhaps even Sinularia.
Lichomolgus cristatus new species
Figures 29-50
Type material: 5729 and 10 copepodids from one colony of
Sinularia leptoclados (Ehrenberg), in 1 m, west of Pte. Mahatsinjo,
Nosy Bé, Madagascar, collected 2 November 1960. Holotype and 45
paratypes deposited in the United States National Museum and the
remaining paratypes in the collection of A. G. Humes.
Female: Body (fig. 29) with moderately broadened prosome, the
sides of the cephalosome more or less parallel instead of rounded.
Length 1.00 mm (0.95-1.10 mm) and greatest width 0.48 mm (0.45—
0.50 mm), based on 10 specimens. Ratio of Jength to width of prosome
1.37: 1. Segment of leg 1 separated dorsally from head by a transverse
furrow. Epimeral areas of segments of legs 1-4 rounded posteriorly
as in figure.
Segment of leg 5 (fig. 30) 96 w xX 166 uw. Ventrally between this
segment and genital segment a weak intersegmental sclerite. Genital
segment shorter than wide, 94 u xX 146 uw, in dorsal view rounded on
either side and constricted posteriorly. Areas of attachment of egg
sacs situated dorsally in midregion of segment. Each area (fig. 31)
bearing two small naked setae 6 uw long. Three postgenital segments
Copepods Associated with Octocorals 645
Fics. 29-33. Lichomolgus cristatus, new species, female: 29, body,
dorsal (A); 30, urosome, dorsal (B); 31, area of attachment of egg sac,
dorsal (C); 32, caudal ramus, dorsal (E); 33, rostral area, ventral (B).
57 uw X 70 uw, 36 uw X 60 pw, and 60 uw X 50 wu, from anterior to posterior.
Caudal ramus (fig. 32) about 4.5 times longer than wide, its greatest
dimensions being 89 u X 20 w. Outer lateral seta 80 » long and naked,
pedicellate dorsal seta 38 «4 and naked, outermost distal seta 112 « with
lateral hairs proximally, innermost distal seta 156 mu with hairs along
inner margin, and the two long median terminal setae 325 yu (outer)
646 Proceedings of the Biological Society of Washington
Fics. 34-40. Lichomolgus cristatus, new species, female: 34, first
antenna, anteroventral (D); 35, second antenna, posterior (outer)
(F); 36, labrum, ventral (E); 37, mandible, posterior (C); 38,
paragnath, ventral (C); 39, first maxilla, posterior (C); 40, second
maxilla, posterior (EF).
with a few spinules on outer margin and 400 uw (inner) without spinules.
A few surficial spinules on ramus.
Dorsal surface of prosome and urosome with very few hairs. Ratio
of length of prosome to that of urosome 1.7: 1.
Copepods Associated with Octocorals 647
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Fics. 41-45. Lichomolgus cristatus, new species, female: 41, maxil-
liped, posterior (E.); 42, area between maxillipeds and first pair of legs,
ventral (F); 43, leg 1 and intercoxal plate, anterior (F); 44, spine on
last segment of endopod of leg 1, anterior (H); 45, leg 2 and intercoxal
plate, anterior (F).
Egg sac (fig. 29) elongated, 380 » x 100 uw, reaching beyond caudal
rami, with each egg about 50 u in diameter.
Rostral area (fig. 33) with an extremely delicate broadly rounded
posteroventral margin.
First antenna (fig. 34) 7-segmented, 342 » in length. Lengths of
segments: 55 uw (68 uw along anterior margin), 130 pw, 33 pw, 47 uw, 25 yu,
648 Proceedings of the Biological Society of Washington
ett
N
N
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iS
Fics. 46-50. Lichomolgus cristatus, new species, female: 46, leg 3
and intercoxal plate, anterior (F); 47, leg 4 and intercoxal plate, anterior
(F); 48, endopod of leg 4, anterior (C); 49, leg 5, dorsal (E); 50,
free segment of leg 5, dorsal (E).
21 w, and 18 » respectively. Formula for armature as in L. dentipes.
Many setae haired as in figure.
Second antenna (fig. 35) 4-segmented, with last segment moderately
elongated, 70 uw along its outer edge, 42 uw along its inner edge, and
19 « wide at middle. First segment with a small inner seta, second
with a similar seta and outer surficial spinules, third with three setae,
Copepods Associated with Octocorals 649
and fourth with six elements: five small hyaline elements and a
terminal recurved claw 46 uw along its greatest axis. All setae naked.
Labrum (fig. 36) with two broad and rather truncated posteroventral
lobes.
Mandible (fig. 37) with very reduced flagellum and _ resembling
closely that of L. dentipes. Paragnath (fig. 38) a small hairy lobe. First
maxilla (fig. 39) 1-segmented, with two unequal terminal elements.
Second maxilla (fig. 40) 2-segmented, large first segment unarmed,
second segment with a very small setule near its proximal outer margin,
a naked seta on its posterior surface, an outer distal spine as long as
the lash and prominently barbed along one edge, and the segment
produced distally to form a lash bearing along its inner edge a
proximal spine, followed by a row of unusually long rather hyaline
setae, and then a graduated row of smaller spinules. Maxilliped (fig.
41) 3-segmented, first segment with a few small spinules, second with
a few spinules and two very unequal setae (the short seta naked, the
long one with barbules distally), and third with two terminal spiniform
barbed elements (the inner one without a distinct articulation) and a
minute setiform process.
Area between maxillipeds and first pair of legs (fig. 42) not pro-
tuberant; a sclerotized line between bases of maxillipeds.
Legs 1-4 (figs. 43, 45-47) segmented as in L. dentipes, and having
same spine and setal formula. Inner seta on coxa of legs 1-3 long and
plumose, but in leg 4 very short (8 «) and finely barbed. Inner margin
of basis in legs 1-3 with row of hairs, but this margin naked in leg 4.
Outer seta on basis unusually long in legs 3 and 4. Spine on last seg-
ment of endopod of leg 1 (fig. 44) somewhat recurved with prominent
spinules on outer margin and a proximal fringe of much smaller spinules
on inner margin. Endopod of leg 4 (fig. 48) slender, first segment
25 « X 9 w with its inner distal feathered seta 45 wu, second segment
48 u long (including processes ) and 8.5 « wide at middle, its two terminal
unequal barbed spines being 15 « (outer) and 34 uw (inner), the latter
more strongly barbed along inner side than outer side. Row of long
hairs along outer margins of both segments.
Leg 5 (fig. 49) with elongated free segment, 68 uw long, 17 » wide
at the slight proximal inner expansion (fig. 50), 12 « wide at midregion.
Two terminal naked setae 40 uw and 42 uw. Naked seta on body near
insertion of free segment 60 uz.
Leg 6 probably represented by the two setae near areas of attachment
of each egg sac (see fig. 31).
Color in life in transmitted light translucid, eye red, egg sacs gray.
Male: Unknown.
Etymology: The specific name cristatus, from Latin = crested, refers
to the crest of unusually long setae on the proximal part of the terminal
lash of the second maxilla.
Comparison with related species: The crest of long setae on the
650 Proceedings of the Biological Society of Washington
terminal lash of the second maxilla sets this species apart from all
other known species in the genus. In some respects L. cristatus
resembles L. dentipes, for example, the second antenna, labrum,
mandible, maxilliped, and endopod of leg 4.
Since the male of L. cristatus is unknown, comparisons with L.
aegyptius Gurney, 1927, and L. vagans Gurney, 1927, of which only
males are known, cannot be made directly. In L. aegyptius, however,
the male has three claws and two setae on the end of the second
antenna, the caudal ramus is very short (nearly as broad as long), and
the slender mandible has a long flagellum. In L. vagans the male has
two claws on the second antenna, the caudal ramus is only a little longer
than wide, and the formula for the third segment of the exopod of leg
4 is II,1,5. It would appear very unlikely, therefore, that the new species
could be the same as either of Gurney’s species.
Lichomolgus adelphus new species
Figures 51-66
Type material: 519 2, 344 4, and 11 copepodids from one colony
of Sinularia whiteleggei Liittschwager, in 2 m, Antsamantsara, northwest
of Madirokely, Nosy Bé, Madagascar, collected 31 October 1960. Holo-.
type @, allotype, and 62 paratypes (372 9, 254 6) deposited in the
United States National Museum, and the remaining paratypes in the
collection of A. G. Humes.
Other material: 52 9, 114 6, and 4 copepodids from one colony of
Sinularia pedunculata Tixier-Durivault, in 3 m, Pte. Ambarionaomby,
Nosy Komba, near Nosy Bé, 3 October 1960; and 109 2, 94 6, and 4
copepodids from one colony of Sinularia polydactyla (Ehrenberg), in
15 m, Tany Kely, a small island south of Nosy Bé, 30 August 1964.
Female: Body (fig. 51) resembling that of L. squamiger Humes and
Frost, 1964. Length 1.29 mm (1.26-1.37 mm) and greatest width 0.55
mm (0.51-0.59 mm), based on 10 specimens. Ratio of length to width
of prosome 1.58: 1. Epimeral areas of segments of legs 1-4 some-
what more angulate posteriorly than in L. squamiger.
Segment of leg 5 (fig. 52) 101 w xX 229 uw. Ventrally between this
segment and genital segment no intersegmental sclerite. Genital segment
153 w long, in dorsal view expanded in anterior part (width 187 uw)
and constricted with parallel sides posteriorly (width 109 uw). Areas
of attachment of egg sacs situated dorsolaterally near middle of segment.
Each area with two very unequal naked elements, as in L. squamiger.
Three postgenital segments 73 u X 99 uw, 57 uw X 91 pw, and 78 uw xX 88 y,
from anterior to posterior.
Caudal ramus (fig. 53) about as long as wide, 35 u X 23 uw in greatest
dimensions. Outer lateral seta 77 wu, pedicellate dorsal seta 50 4p,
outer most distal seta 121 mw, innermost distal seta 300 », and the two
long median terminal setae 470 « (outer) and 685 uw (inner), both
inserted between slight unornamented dorsal and ventral flaps. All
Copepods Associated with Octocorals 651
Fics. 51-55. Lichomolgus adelphus, new species, female: 51, body,
dorsal (1); 52, urosome, dorsal (G); 53, caudal ramus, dorsal (C); 54,
tip of second antenna, anterior (inner) (C); 55, maxilliped, postero-
inner (E).
setae naked. A few surficial small setules and refractile points on ramus.
Dorsal surface of prosome and urosome with a few short hairs. Ratio
of length of prosome to that of urosome 1.85: 1.
Form of egg sac unknown, since all ovigerous females collected had
broken sacs. Each egg about 52 uw in diameter.
Rostral area as in L. squamiger.
652 Proceedings of the Biological Society of Washington
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Fics. 56-60. Lichomolgus adelphus, new species, female: 56, area
between maxillipeds and first pair of legs, ventral (D); 57, endopod
of leg 3, anterior (F); 58, endopod of leg 4, anterior (F); 59, leg 5,
dorsal (F). Male: 60, body, dorsal (A).
First antenna similar to that of L. squamiger. Lengths of segments:
55 uw (88 uw along anterior edge), 143 uw, 44 uw, 104 u, 47 w, 36 w, and 31 w
respectively. Formula for armature as in L. squamiger (and also like
L. dentipes, given above).
Second antenna resembling that of L. squamiger, but inner distal
edge of first segment a little more swollen. Fourth segment 104 uw along
Copepods Associated with Octocorals 653
Fics. 61-66. Lichomolgus adelphus, new species, male: 61, urosome,
dorsal (B); 62, second segment of second antenna, anterior (inner)
(E); 63, maxilliped, outer (B); 64, endopod of leg 1, anterior (E); 65,
last segment of endopod of leg 2, anterior (E); 66, spermatophore,
empty and attached to female, dorsal (B).
outer side, 66 uw along inner side, and 21 » wide; bearing distally six
small hyaline elements (as in L. squamiger, though only three such
elements indicated in Humes and Frost’s figure 174) and a terminal
claw 52 u long (fig. 54).
Labrum, mandible, paragnath, first maxilla, and second maxilla as
654 Proceedings of the Biological Society of Washington
in L. squamiger. Maxilliped (fig. 55) closely resembling that of L.
squamiger with only minor differences in ornamentation and with third
segment bearing in addition to the two large terminal elements a small
seta (present also in L. squamiger, but not shown in Humes and Frost’s
figure 180).
Area between maxillipeds and first pair of legs (fig. 56) slightly
protuberant; a sclerotized line between bases of maxillipeds.
Legs 1-4 segmented as in L. squamiger and with the same spine and
setal formula (also same as in L. dentipes, given above). Legs 1 and
2 like those of L. squamiger. Leg 3 with exopod like that of L.
squamiger, but last segment of endopod (fig. 57) relatively shorter
and of a slightly different form than in that species. Leg 4 also with
exopod similar to that of L. squamiger; endopod (fig. 58) with first
segment 47 uw long (not including processes) and 30 mu wide, with
inner distal seta 107 u, second segment 125 u long (including processes),
19 » wide at midregion, with only two terminal spiniform processes
instead of three as in L. squamiger. Two terminal spines of endopod
40 uw (outer) and 69 w (inner).
Leg 5 (fig. 59) with free segment elongated, slender, and bowed,
195 uw X 26 u, distinctly longer and more slender than in L. squamiger.
Convex surface with two rows of scalelike spines along proximal half
merging to a single row in distal half. Two terminal naked setae 36 u
(outer) and 73 u (inner). Seta on body near insertion of free segment
39 uw and naked.
Leg 6 as in L. squamiger.
Color in life in transmitted light translucid, with orange globules in
prosome, eye red.
Male: Body (fig. 60) resembling that of L. squamiger. Length 0.98
mm (0.95-0.99 mm) and greatest width 0.34 mm (0.33-0.35 mm),
based on 10 specimens. Ratio of length to width of prosome 1.77: 1.
Segment of leg 5 (fig. 61) 52 « X 112 uw. No ventral intersegmental
sclerite. Genital segment a little longer than wide, 200 » x 185 u,
similar in form to that of L. squamiger. Four postgenital segments
36 uw X 71 w, 39 uw xX 70 wu, 27 uw X 62 w, and 36 u X 60 yu, from anterior
to posterior.
Caudal ramus similar to that of female, but a little wider than long,
22 uw X 26 uw.
Dorsal surface of prosome and urosome with a few small hairs. Ratio
of length of prosome to that of urosome 1.58: 1.
Rostral area as in L. squamiger, with two lateral anterior processes
more prominent than in female, as in that species.
First antenna like that of L. squamiger, with same formula for arma-
ture (also same as for L. dentipes, given above). Second antenna as
in L. squamiger, but lacking the obtuse spines seen in that species and
having two rows of slender spines along inner surface of second segment
(fig. 62).
Copepods Associated with Octocorals 655
Labrum, mandible, paragnath, first maxilla, and second maxilla as in
female. Maxilliped (fig. 63) very long and slender (about 600 wu in-
cluding claw when extended). Second segment with a single inner
row of spines; two inner setae as in L. squamiger, the proximal one
with a fringe of spinules along its proximal edge as in that species.
Claw 297 wu along its axis, longer than in L. squamiger. Area between
maxillipeds and first pair of legs as in L. squamiger.
Legs 1-4 segmented as in female, and the spine and setal formula
as in that sex except for last segment of endopod of leg 1 (fig. 64)
which is I,],4 (as in L. squamiger). Sexual dimorphism also in last
segment of endopod of leg 2 (fig. 65), where spines and spiniform
processes are very different from those of L. squamiger. Legs 3 and
4 resembling those of L. squamiger.
Leg 5 (see fig. 61) similar to that of L. squamiger, free segment
being 50 u X 9 uw. Leg 6 as in L. squamiger.
Spermatophore (fig. 66), attached to female and empty, 151 » x
86 uw, not including neck.
Color in life as in female.
Etymology: The specific name adelphus, from Greek aée\¢és = sisterly
or brotherly, alludes to the close relationship of this species to L.
squamiger.
Comparison with related species: Like L. squamiger, the new species
is close to L. spinipes (Sewell, 1949), known only from a single female
(now lost) found in weed-washings in the Nicobar Islands. However,
like L. squamiger, L. adelphus is distinct from L. spinipes. The distinc-
tions are largely the same as those mentioned by Humes and Frost
(1964, pp. 147-148) in comparing L. squamiger with L. spinipes.
Several characters may be used to distinguish L. adelphus from L.
squamiger: the form of the last segment of the endopod of leg 3 in
the female, the presence of only two terminal spiniform processes on
the endopod of leg 4 in both sexes, the more elongated slender bowed
free segment of leg 5 in the female, the absence of obtuse spines on
the second antenna of the male, and the sexual dimorphism in the
last segment of the endopod of leg 2 in the male.
Lichomolgus foxi Gurney, 1927
Figures 67-88
Gurney described this species on the basis of one female taken at Port
Taufiq in the Suez Canal. This specimen was supposed to have been
deposited in the British Museum (Natural History), but upon examina-
tion of the vial in the museum’s collection labeled “Lichomolgus foxi
n. sp.” we have found only a fragment of an unknown lichomolgid and
a specimen of Corycaeus. Thus the type specimen appears to have
been misplaced or lost. The male has been unknown until now.
In Madagascar the first author collected specimens referable to L. foxi
as follows: 1969 9, 2124 6, and 74 copepodids from several colonies
656 Proceedings of the Biological Society of Washington
e:
Ch
COorariioes
(7 ry x
KI COL
5] NS
LY <@' 4
a
(>
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S=S50L]
H)
ee
Fics. 67-71. Lichomolgus foxi Gurney, 1927, female: 67, body,
dorsal (1); 68, urosome, dorsal (B); 69, area of attachment of egg sac,
dorsal (E); 70, caudal ramus, dorsal (E); 71, first antenna, dorsal (B).
of Cladiella krempfi Hickson, in 1 m, west of Pte. de Tafondro, Nosy
Bé, 5 December 1963 (specimens deposited in the United States
National Museum, the Zodlogisch Museum, Amsterdam, and the British
Museum (Natural History) ); 25992, 194 4, and 7 copepodids from
one colony of Cladiella laciniosa (Tixier-Durivault), in 1 m, Nosy
Kisimany, a small island 27 km southwest of Nosy Bé near the mainland
of Madagascar, 4 October 1960 (specimens in the U.S.N.M.); and 2092 @
Copepods Associated with Octocorals 657
Fics. 72-77. Lichomolgus foxi, Gurney, 1927, female: 72, second
antenna, anterior (inner) (D); 73, labrum, with paragnaths in dashed
lines, ventral (F); 74, mandible, posterior (E.); 75, first maxilla, anterior
(E); 76, second maxilla, posterior (FE); 77, maxilliped, posterior (E).
and 96 6 from one colony of Cladiella pachyclados (Klunzinger), in 1
m, Ambariotelo, a small island almost between Nosy Bé and Nosy
Komba, 15 May 1964 (specimens in the U.S.N.M.).
Female: Body (fig. 67) with slender prosome. Length 1.16 mm
(1.04-1.27 mm) and greatest width 0.51 mm (0.49-0.54 mm), based
on 10 specimens. Ratio of length to width of prosome 1.57 : 1. Segment
658 Proceedings of the Biological Society of Washington
Fics. 78-82. Lichomolgus foxi Gurney, 1927, female: 78, leg 1 and
intercoxal plate, anterior (D); 79, leg 2 and intercoxal plate, anterior
(D); 80, last segment of endopod of leg 3, anterior (D); 81, leg 4,
anterior (D); 82, leg 5, dorsal (F).
of leg 1 separated dorsally and laterally from head by a transverse
furrow. Epimeral areas of segments of legs 1-4 as in figure.
Segment of leg 5 (fig. 68) 80 u x 143 uw. Between this segment and
genital segment a short ventral intersegmental sclerite. Genital segment
156 w X 135 uw, only a little longer than wide, and not greatly expanded
laterally in dorsal view. Areas of attachment of egg sacs located
Copepods Associated with Octocorals 659
=D)
SSS
88
Fics. 83-88. Lichomolgus foxi Gurney, 1927, male: 83, body,
dorsal (A); 84, urosome, dorsal (B); 85, maxilliped, outer (D); 86, last
segment of endopod of leg 1, anterior (E); 87, endopod of leg 4,
anterior (E); 88, spermatophore, empty and attached to female, dorsal
(G).
dorsolaterally in middle of segment. Each area (fig. 69) bearing two
small naked setae 10 » and 12 wu long, partially covered in dorsal view
by sclerotized processes. Region between two areas of attachment
with sclerotized lines and setules as indicated in figure. Three postgenital
segments 52 uw X 86 un, 36 uw xX 78 wu, and 52 uw xX 78 uw, from anterior to
posterior.
660 Proceedings of the Biological Society of Washington
Caudal ramus (fig. 70) about as long as wide, 43 ux 37 w in
greatest dimensions. Outer lateral seta 180 mw and naked, pedicellate
dorsal seta 46 uw and lightly feathered, outermost distal seta 195 wu with
hairs on proximal inner side, innermost distal seta 350 mu with hairs
on both sides proximally, and the two long median terminal setae 600 u
(outer) and 790 uw (inner), both with coarse spinules on both sides
except near bases and both inserted between unornamented dorsal and
ventral flaps. :
Dorsal surface of prosome and urosome with a few small hairs. Ratio
of length of prosome to that of urosome 2.1 : 1.
Egg sac (fig. 67) elongated, 420 u x 140 uw, reaching well beyond
caudal rami. Each egg about 46 uw in diameter.
Rostral area resembling that of L. cristatus.
First antenna (fig. 71) 7-segmented, 563 w long. Lengths of seg-
ments: 55 uw (91 uw along anterior edge), 179 u, 36 uw, 91 uw, 73 w, 55 p,
and 38 uw respectively. Formula for armature as in three previous species.
All setae naked.
Second antenna (fig. 72) 4-segmented. Longest seta on third segment
characteristically bent. Last segment moderately elongated, 88 » along
its outer edge, 59 w along its inner edge, and 23 wu wide, bearing distally
two short and three rather long setae and two unequal terminal claws,
the stouter claw 46 wu along its axis, the more slender claw 33 uz.
Labrum (fig. 73) with two somewhat obtusely pointed posteroventral
lobes. Mandible (fig. 74) with basal region distal to constriction bearing
on its convex margin a distally directed pointed scalelike process
ornamented with a row of spinules, followed by a serrated fringe, and
on its concave margin a row of slender spinules joining a group of
somewhat stouter spinules near base of flagellum; flagellum elongated
with lateral spinules. Paragnath (fig. 73) a small hairy lobe. First
maxilla (fig. 75) with three long terminal naked setae and a minute
subterminal seta. Second maxilla (fig. 76) 2-segmented, first segment
unarmed, second with a small setule on proximal inner margin, a
surficial posterior seta finely barbed along one edge, an outer distal
spine with prominent spinules mostly along one edge, and the segment
produced distally to form a lash with dentiform spines proximally and
fine bilateral spinulation distally. Maxilliped (fig. 77) 3-segmented,
armed as in L. dentipes, though details of form and ornamentation
somewhat different.
Area between maxillipeds and first pair of legs similar to that in L.
cristatus, but slightly protuberant; a sclerotized line between bases of
maxillipeds.
Legs 1-4 (figs. 78-81) segmented as in three previous species and
with same spine and setal formula. Inner seta on coxa of legs 1-3 long
and plumose, but in leg 4 very short (8 «) and naked. Inner margin
of basis with row of hairs in legs 1-3, but naked in leg 4. Outer spines
of exopod of leg 1 with strong spinules along proximal edges. Endopod
Copepods Associated with Octocorals 661
of leg 4 shorter than exopod. First segment 37 wu X 32 mu (not including
processes) with inner distal naked seta very short (11 m) and turned
anteriorly (so that in casual examination it might appear to be absent).
Second segment 65 uw xX 26 w (greatest width) bearing two terminal
very unequal spines, outer 23 mu and naked, inner 61 uw with an outer
finely barbed fringe and a narrow inner lamella. Both segments with
outer margins haired, and second segment with a row of fine spinules
near insertions of terminal spines. ;
Leg 5 (fig. 82) with elongated free segment, 101 uw in length, its
proximal area expanded inwardly (width here 33 uw) but distal two-
thirds of segment slender (15 mw at widest point). Segment bearing
two naked terminal setae 94 uw and 99 uw and short spinules along its
outer margin. Seta on body near insertion of free segment 45 w and
feathered. Expansion of free segment partially covered in dorsal view
by a posterolateral extension of dorsum of body segment.
Leg 6 probably represented by the two setae near areas of attachment
of each egg sac (see fig. 69).
Color in life in transmitted light translucid to slightly opaque, eye
red, egg sacs gray.
Male: Body (fig. 83) similar in general shape to female. Length
0.93 mm (0.88-0.96 mm) and greatest width 0.32 mm (0.31—0.33 mm),
based on 10 specimens. Ratio of length to width of prosome 1.66: 1.
Segment of leg 5 (fig. 84) 42 u x 91 uw. No ventral intersegmental
sclerite. Genital segment longer than wide, 211 u x 172 u, its lateral
borders in dorsal view slightly irregular. Four postgenital segments
32 uw X 57 w, 28 uw X 56 wu, 20 uw X 56 uw, and 29 uw x 56 yu, from anterior
to posterior.
Caudal ramus similar to that of female, but smaller, 26 u x 24 wu.
Dorsal surface of prosome and urosome with a few small hairs. Ratio
of length of prosome to that of urosome 1.46: 1.
Rostral area as in female.
First antenna similar to that of female, but with two aesthetes
added on second segment and one on fourth segment, so that formula
is same as for L. dentipes and L. adelphus. Second antenna resembling
that of female, but with small spinules added on inner surface of first,
second, and fourth segments.
Labrum, mandible, paragnath, first maxilla, and second maxilla
like those of female. Maxilliped (fig. 85) resembling in general form
and armature that of L. dentipes. Claw 185 wu along its axis (including
terminal lamella).
Area between maxillipeds and first pair of legs as in female.
Legs 1-4 segmented as in female, and spine and setal formula as in
that sex except for last segment of endopod of leg 1 (fig. 86) which
is 11,4. No sexual dimorphism in legs 2 or 3. Endopod of leg 4
(fig. 87) closely resembling that of female.
Leg 5 (see fig. 84) with slender free segment 41 » x 8 uw, without
662 Proceedings of the Biological Society of Washington
a proximal expansion. Outer margin with fewer spinules than in female.
Two terminal naked setae 31 uw (inner) and 63 wu (outer).
Leg 6 (see fig. 84) a posterolateral flap on ventral surface of genital
segment bearing two naked setae 30 u and 39 u long.
Spermatophore (fig. 88), attached to female and empty, elongated,
220 w X 90 uw (not including neck).
Color in life as in female.
Remarks: As nearly as can be determined, the specimens from
Cladiella in Madagascar represent the species described by Gurney
(1927) as L. foxi. In Gurney’s description, based upon a single female,
several significant points of similarity with the Madagascan specimens
may be noted: the body shape and size, the proportional lengths of
the segments of the first antenna, the bent seta on the third segment
and the two unequal claws on the last segment of the second antenna,
the strong spinules on the proximal edges of the outer spines of the
exopod of leg 1, and the form of the caudal ramus.
There are two rather perplexing apparent differences. Gurney
stated that the first segment of the endopod of leg 4 had no seta, but
added that “it may have been broken off.” In his figure 113E he showed
two small spiniform processes at the region where the seta would
normally be. In the Madagascan females the very small seta at this
point is often directed anteriorly and could easily be overlooked when
the leg is examined on a slide in flat view. We think it probable that
one of Gurney’s spiniform processes may represent this seta, since it
would be very unusual in Lichomolgus for two spiniform processes to
be present here. The length of the second segment of the endopod in
leg 4 is relatively shorter and the two terminal spines are less unequal
in Gurmey’s figure than in the Madagascan females. However, such
small differences may be attributable to the technique of drawing.
Leg 5 as shown in Gurney’s figure 113G in general resembles that in
the Madagascan specimens, but shows an inner proximal expansion of
somewhat different form, the terminal setae are relatively shorter, and
the outer margin is said to be hairy. The form of the expansion in his
figure does not appear to be significantly different from that in the
Madagascan females, since the variation could be introduced by the
angle at which the leg was drawn. The terminal setae in the
Madagascan females are very delicate distally and those shown in
Gurney’s figure may not represent their entire length. The “hairy”
nature of the outer margin may simply be an interpretation of the rather
small spinules seen here in the Madagascan material.
Admittedly we are interpreting these features in relation to what
we know of the females from Madagascar, but in the absence of types
or other specimens we have come to the conclusion that the apparent
differences in the endopod of leg 4 and in leg 5 are probably not
significant, and that our specimens from Cladiella actually represent
L. foxi.
Copepods Associated with Octocorals 663
Lichomolgus hetaericus new species
Figures 89-106
Type material: 749 9, 186 6, and 1 copepodid from one colony of
Cladiella pachyclados (Klunzinger), in 1 m, Ambariotelo, a small island
almost between Nosy Komba and Nosy Bé, Madagascar, collected 15
May 1964. Holotype @, allotype, and 62 paratypes (509 9, 12¢ ¢)
deposited in the United States National Museum, and the remaining
paratypes in the collection of A. G. Humes.
Other material: 999 from several colonies of Cladiella krempfi
Hickson, in 1 m, west of Pte. de Tafondro, Nosy Bé, 5 December 1963.
Female: Body (fig. 89) with moderately broadened prosome. Length
0.84 mm (0.78—-0.89 mm) and greatest width 0.42 mm (0.40—-0.45 mm),
based on 10 specimens. Ratio of length to width of prosome 1.27: 1.
Segment of leg 1 separated dorsally and laterally from head by a
transverse furrow. Epimeral areas of segments of legs 1-4 as in figure.
Segment of leg 5 (fig. 90) 62 u x 109 uw. No ventral intersegmental
sclerite. Genital segment (fig. 90) about as long as wide, 107 u x 112 n,
in dorsal view moderately expanded laterally. Areas of attachment of
egg sacs situated laterally near middle of segment. Each area (fig. 91)
bearing two small naked spiniform setae 6 w and 4.5 uw long, and partly
covered dorsally by a flap of dorsum of genital segment. Three
postgenital segments 34 uw X 62 uw, 26 w X 60 w, and 29 uw x 57 pw, from
anterior to posterior.
Caudal ramus (fig. 92) nearly quadrate, 22 u xX 23 mw in greatest
dimensions. Outer lateral seta 133 » and naked, dorsal pedicellate seta
80 « and apparently naked, outermost distal seta 79 mw and naked,
innermost distal seta 143 « and haired proximally, and the two long
median terminal setae 395 » (outer) and 495 w (inner), both with
lateral spinules except near bases and inserted between dorsal and
ventral unornamented flaps.
Dorsal surface of prosome and urosome with a few small hairs. Ratio
of length of prosome to that of urosome 2.15: 1.
Egg sac (fig. 89) elongated, 660 » x 187 u, nearly three times length
of urosome. Each egg about 49 uw in diameter.
Rostral area as in L. cristatus.
First antenna similar to that of L. foxi. Lengths of segments: 34 pu
(68 » along anterior margin), 151 pu, 26 uw, 65 uw, 47 w, 32 w, and 29 pw
respectively. All setae naked. Second antenna also resembling that of
L. foxi, with seta on third segment bent as in that species. Last segment
75 uw along outer side, 47 wu along inner side, and 17 wide at middle.
Stouter claw 36 u, more slender claw 29 uw.
Labrum (fig. 93) with two rather pointed posteroventral lobes.
Mandible (fig. 94) resembling that of L. foxi, but distal end of
scalelike process on convex margin of basal region ornamented with
spinules. Paragnath and first maxilla (fig. 94) similar to those in
L. foxi. Second maxilla (fig. 95) resembling that of L. foxi, but proximal
664 Proceedings of the Biological Society of Washington
Fics. 89-95. Lichomolgus hetaericus, new species, female: 89, body,
dorsal (A); 90, urosome, dorsal (B); 91, area of attachment of egg
sac, dorsal (C); 92, caudal ramus, dorsal (C); 93, labrum, ventral (E);
94, mandible and first maxilla, posterior (C); 95, second maxilla,
posterior (E).
spines on lash more slender. Maxilliped (fig. 96) similar to that of
L. foxi, but first segment without ornamentation, long seta on second
segment relatively shorter, and a spiniform process between two large
terminal elements.
Area between maxillipeds and first pair of legs not protuberant and
similar to that in L. cristatus.
Copepods Associated with Octocorals 665
Fics. 96-101. Lichomolgus hetaericus, new species, female: 96,
maxilliped, anterior (C); 97, leg 1 and intercoxal plate, anterior (F);
98, leg 2, anterior (F); 99, last segment of endopod of leg 3, anterior
(E); 100, leg 4, anterior (F); 101, leg 5, dorsal (C).
Legs 1-4 (figs. 97-100) segmented as in four previous species, with
same spine and setal formula except for last segment of exopod of leg
4 which is IILI,5. Inner seta on coxa long and plumose in legs 1-3,
but extremely minute (3 ~) and naked in leg 4. Outer seta on basis
long (70 uw) in legs 1, 3, and 4. Inner margin of basis in leg 4 naked.
Proximal outer spine on third segment of exopod in legs 1-3 distinctly
666 Proceedings of the Biological Society of Washington
Fics. 102-108. Lichomolgus hetaericus, new species, male: 102,
body, dorsal (A); 103, urosome, dorsal (D); 104, second maxilla,
posterior (C); 105, maxilliped, outer (F); 106, last segment of endopod
of leg 1, anterior (C). Lichomolgus insolens, new species, female: 107,
body, dorsal (A); 108, urosome, dorsal (B).
shorter than distal spines. Endopod of leg 4 (fig. 100) rather slender,
shorter than exopod. First segment 22 u X 12 uw, with its inner distal
seta 20 uw and very lightly feathered. Second segment 44 uw X 11 uw (in-
cluding processes and width taken at widest point), two terminal fringed
spines 11 w (outer) and 23 w (inner). Hairs along outer margins of
both segments and minute spinules near insertions of terminal spines.
Copepods Associated with Octocorals 667
Leg 5 (fig. 101) with small unornamented free segment 27 » < 12 n,
its two terminal naked setae 46 » (outer) and 55 mw (inner). Seta
on body near free segment 50 uw and lightly feathered.
Leg 6 probably represented by the two small setae near areas of
attachment of each egg sac (see fig. 91).
Color in life as in L. foxi.
Male: Body (fig. 102) with prosome more slender than in female.
Length 0.62 mm (0.59-0.66 mm) and greatest width 0.25 mm (0.24—
0.27 mm), based on 10 specimens. Ratio of length to width of prosome
1.53): 1.
Segment of leg 5 (fig. 103) 26 u x 65 uw. No ventral intersegmental
sclerite. Genital segment about as long as wide, 139 » x 130 4u, its
lateral borders in dorsal view only slightly rounded. Four postgenital
segments 18 uw X 41 uw, 17 uw X 40 uw, 13 uw X 40 uw, and 19 uw xX 43 uw, from
anterior to posterior.
Caudal ramus resembling that of female, but smaller, 19 u x 18 x.
Dorsal surface of prosome and urosome with a few small hairs. Ratio
of length of prosome to that of urosome 1.62: 1.
Rostral area as in female. First antenna like that of female, but two
aesthetes added on second segment and one on fourth segment, so that
formula is same as in L. dentipes, L. adelphus, and L. foxi. Second
antenna similar to that of female, but, as in L. foxi, a few small
spinules added on inner surface of segments 1, 2, and 4.
Labrum, mandible, paragnath, and first maxilla like those of female.
Second maxilla similar to that of female but proximal spines on lash
coarser and longer (fig. 104). Maxilliped (fig. 105) resembling that
of L. foxi, but claw (122 «) shorter.
Area between maxillipeds and first pair of legs like that of female.
Legs 1-4 segmented as in female, with same spine and setal formula,
except for last segment of endopod of leg 1 (fig. 106) which has arrange-
ment of I,J,4. Proximal outer spine on third segment of exopod in legs
1-3 short as in female. Legs 2 and 3 as in female, without sexual
dimorphism.
Leg 5 (see fig. 103) with small unornamented free segment 17 » X
7 w, its two terminal setae 28 uw (outer) and 20 mw (inner). Seta on
body near free segment 33 ». All setae naked.
Leg 6 (see fig. 103) a posterolateral flap on ventral surface of
genital segment bearing two naked setae 24 uw and 26 wu long.
Spermatophore not observed.
Color in life like that of female.
Etymology: The specific name hetaericus, from Greek éracpixds =
comradely or sociable, alludes to the occurrence of this species along
with L. foxi on the same colonies of octocorals.
Comparison with related species: L. hetaericus may be distinguished
from all other known species of Lichomolgus by the combination of the
following characters: the nearly quadrate caudal ramus, the second
668 Proceedings of the Biological Society of Washington
antenna with two short terminal claws, the mandible with a relatively
long flagellum and without a strong tooth on its basal part, leg 1
showing sexual dimorphism (I,I,4) in the male, legs 1-3 with the
proximal outer spine on the third segment of the exopod reduced, the
formula for the third segment of the exopod of leg 4 as II,I,5, and the
free segment of leg 5 in the female with a length to width ratio of
2.3: 1 and lacking a proximal expansion or surficial ornamentation.
Two species of Lichomolgus described from -southeastern India by
Ummerkutty (1962) might at first glance be confused with L.
hetaericus, but differ from the new species in significant details.
In L. brevifurcatus Ummerkutty, 1962, the female is 1.6 mm in length
and the male 1.5 mm; the genital segment of the female is “barrel-like
and does not become narrowed in the posterior half’; and the proximal
outer spine on the third segment of the exopod in the swimming legs
is not reduced. In L. indicus Ummerkutty, 1962, the female is 1.05 mm
and the male 1.00 mm; there are only two postgenital segments in the
female and three in the male; there is no sexual dimorphism in leg 1;
and the endopod of leg 4 differs from that of L. hetaericus in details
of shape and armature. L. indicus shows a reduction (as in the new
species) of the proximal outer spine on the third segment of the
exopod of leg 1 (Ummerkutty’s Pl. XII, fig. 9) and leg 2 (his
Pl. XIII, fig. 2).
Lichomolgus insolens new species
Figures 107-127
Type material: 189 9, 164 6, and 17 copepodids from one colony
of Lobophytum crassum Marenzeller, in 1 m, opposite Ambariotsimara-
mara, a small island on the southern coast of Nosy Bé, Madagascar,
collected 12 June 1964. Holotype 9, allotype, and 23 paratypes (129 9,
1144) deposited in the United States National Museum, and the
remaining paratypes in the collection of A. G. Humes.
Other material (from Lobophytum crassum): 1699, 156 6, and 4
copepodids, from one colony, in 10 cm, Ambafaho, Nosy Bé, 25 Septem-
ber 1964; and 129 9, 16¢ 6, and 6 copepodids, from one colony, in
1 m, Nosy N’Tangam, near Nosy Bé, 5 September 1963.
Female: Body (fig. 107) rather weakly sclerotized and with
broadened prosome. Anterior border of head with a minute median
notch. Length 1.11 mm (1.06—-1.19 mm) and greatest width 0.52 mm
(0.48-0.58 mm), based on 10 specimens. Ratio of length to width of
prosome 1.40: 1. Segment of leg 1 separated from head by a weak
dorsal furrow. Epimeral areas of segments of swimming legs formed
as in figure.
Segment of leg 5 (fig. 108) 78 » xX 138 yu. Between this segment
and genital segment a weak intersegmental sclerite ventrally. Genital
segment (fig. 108) longer than wide, 153 » x 121 wu, broadest in its
anterior half and tapering gradually posteriorly. Areas of attachment
ay
Copepods Associated with Octocorals 669
Fics. 109-115. Lichomolgus insolens, new species, female: 109,
area of attachment of egg sac, dorsal (F); 110, caudal ramus, dorsal (C);
111, egg sac, ventral (A); 112, first antenna, dorsal (B); 113, mandible,
posterior (C); 114, second maxilla, posterior (EF); 115, maxilliped,
anterior (E).
of egg sacs situated dorsolaterally at level of junction of anterior two
thirds of segment. Each area (fig. 109) with two small spiniform naked
setae 5 uw and 8 wu in length. Three postgenital segments 68 » x70 un,
47 w X 62 w, and 74 uw < 55 uw, from anterior to posterior.
Caudal ramus (fig. 110) nearly twice as long as wide, 43 uw x 23 yu
Dy? Yee
‘ Ox =
a Si Se
oF j
_— pa 118
Sk Vi
Fics. 116-120. Lichomolgus insolens, new species, female: 116,
leg 1 and intercoxal plate, anterior (F); 117, last segment of endopod
of leg 2, anterior (E.); 118, leg 3, anterior (F); 119, leg 4 and intercoxal
plate, anterior (F); 120, leg 5, dorsal (E).
in greatest dimensions. Outer lateral seta 70 mw and naked, dorsal
pedicellate seta 38 « and lightly feathered, outermost distal seta 104 u
and naked, innermost distal seta 203 w and haired proximally, and the
two long median terminal setae 290 u (outer) with a few outer spinules
and 430 uw (inner) with a few inner spinules, both inserted between
the usual dorsal (unornamented) and ventral (with a short row of
spinules) flaps.
Copepods Associated with Octocorals 671
Fics. 121-127. Lichomolgus insolens, new species, male: 121, body,
dorsal (A); 122, urosome, dorsal (B); 123, second antenna, posterior
(outer) (F); 124, maxilliped, outer (F); 125, last segment of endopod of
leg 1, anterior (E); 126, last segment of endopod of leg 2, anterior (E);
127, last segment of endopod of leg 3, anterior (E).
Dorsal surface of prosome and urosome with a few hairs. Ratio of
length of prosome to that of urosome 1.64: 1.
Egg sac (fig. 111) elongated, 462 » x 176 uw, reaching well beyond
ends of caudal rami. Each egg about 47 « in diameter.
672 Proceedings of the Biological Society of Washington
Rostral area as in L. protentus.
First antenna (fig. 112) 350 uw long and resembling that of previous
species, with the same formula for armature. Lengths of segments:
50 uw (57 uw along anterior edge), 135 uw, 32 uw, 50 uw, 30 uw, 28 uw, and 19 pu
respectively. All setae naked.
Second antenna similar to that of L. protentus. Last segment 48 uw
along its inner side, 66 w along its outer side, and 20 u wide. Claw
39 uw along its axis. -
Labrum as in L. protentus. Mandible (fig. 113) with basal region
distal to constriction bearing on its convex edge a small sclerotized
process followed by a scalelike process with spinules and then by a
serrated fringe, and on its concave margin a row of long slender setules
and at base of lash a toothlike prominence (on anterior surface); lash
elongated with spiny concave margin proximally and barbed distally.
Paragnath and first maxilla as in L. protentus. Second maxilla (fig.
114) resembling in major respects that of L. protentus, but first tooth-
like spine at base of lash larger and not in line with others. Maxilliped
(fig. 115) much like that of L. protentus, but lesser of two terminal
elements smaller than in that species, so that with casual observation
there may appear to be only one large terminal element. _
Area between maxillipeds and first pair of legs generally like that of
L. protentus, but not protuberant; a line between bases of maxillipeds.
Legs 1-4 (figs. 116-119) segmented as in previous species, but
spine and setal formula differing as follows:
P: protopod 0-1 1-0 exp I-0 I-1 IILI4
end 0-1 0-1 I5
P2 protopod 0-1 1-0 exp I-O I-1 JIILJI,5
end 0-1 0-2 LIL3
P, protopod O0-1 1-0 exp JI-0O I-11 ILI5
end 0-1 0-2 L112
P. protopod 0-1 1-0 exp I-O I-1 II,1,5
end 0-1 LJ
Outer margin of coxa of leg 1 with a prominent protrusion. Inner
seta on coxa in legs 1-3 long and plumose, but in leg 4 only 6 uw long
and naked. Inner margin of basis of leg 4 naked except for a minute
sensillum. Outer spines on exopod of leg 1 with coarse spinulation
as in L. protentus. Third segment of exopod of leg 3 with only two
outer spines (fig. 118) as in leg 4. Endopod of leg 4 (fig. 119) slender
and a little longer than exopod. First segment 32 wu x 13 uw, with inner
distal plumose seta 44 uw. Second segment 68 «X13 uw (including
terminal processes), bearing distally an outer naked seta 25 mw and an
inner fringed spine 35 », and resembling the endopod of L. protentus.
Leg 5 (fig. 120) with unornamented free segment about 3.6 times
longer than wide, 43 uw X 12 uw, with width at very slight inner proximal
expansion 14 yw; bearing two terminal naked setae 40 u (outer) and 53 u
(inner). Seta on body near free segment 50 uw and naked.
Copepods Associated with Octocorals 673
Leg 6 probably represented by the two setae near areas of attachment
of each egg sac (see fig. 109).
Color in life in transmitted light translucid, prosome sometimes finely
speckled with reddish orange, intestine brown, eye red, ovary opaque
gray, egg sacs gray or speckled with reddish orange.
Male: Body (fig. 121) with prosome less broadened than in female.
Anterior border of head smooth, without median notch. Length 0.89
mm (0.84-0.91 mm) and greatest width 0.33 mm (0.29-0.34 mm),
based on 10 specimens. Ratio of length to width of prosome 1.49: 1.
Segment of leg 1 incompletely separated from head. Lateral borders
of cephalosome showing a few slight indentations.
Segment of leg 5 (fig. 122) 38 u x 74 uw. No ventral intersegmental
sclerite. Genital segment longer than wide, 192 ux 164 yw. Four
postgenital segments 31 uw X 48 uw, 31 uw X 47 w, 21 wu X 44 w, and 36 ux
45 uw, from anterior to posterior.
Caudal ramus similar to that of female, but smaller, 30 u x 19 u.
Surfaces of prosome and urosome with a few small hairs as in female.
Ratio of length of prosome to that of urosome 1.46: 1.
Rostral area as in female.
First antenna as in female, but two aesthetes added on second segment
and one on third segment, so that formula is same as for males of
previous species and of L. protentus. Second antenna (fig. 123) like
that of female, but second segment with a long striated inner membra-
nous lamella.
Labrum, mandible, paragnath, first maxilla, and second maxilla re-
sembling those of female. Maxilliped (fig. 124) slender, as in L.
protentus. Claw 126 uw along its axis, with a series of obtuse hyaline
knobs on its concave surface.
Area between maxillipeds and first pair of legs as in female.
Legs 1-4 segmented as in female, with same spine and setal formula
as in that sex except for last segment of endopod of leg 1 which has
arrangement of LI,4 (fig. 125). Sexual dimorphism seen also in
endopods of legs 2 and 3. Last segment of endopod of leg 2 (fig. 126)
with three spines shorter and broader than in female, with first two
having coarser spinulation. (In female lengths of these spines from
proximal to distal are 18 », 17 uw, and 17 uw; in male 13 yp, 14 uw, and
12 «). Last segment of endopod of leg 3 (fig. 127) with middle spine
modified (compare with fig. 118). Endopod of leg 4 as in female.
Leg 5 (see fig. 122) with small unornamented free segment, 20 u x
7 w, without a proximal expansion. Two terminal naked setae 28 uw and
31 uw, and naked seta on body near free segment 35 yu.
Leg 6 (see fig. 122) a posterolateral flap on ventral surface of
genital segment bearing two naked setae about 23 » long.
Spermatophore (see fig. 107), attached to female, elongated and
somewhat irregular, 177 « x 86 uw (not including neck).
674 Proceedings of the Biological Society of Washington
Color in life in transmitted light more translucid than in female, eye
red.
Etymology: The specific name insolens, from Latin = contrary to
custom, refers to the unusual formula of II,I,5 for the last segment of
the exopod of leg 3 in this species.
Comparison with related species: L. insolens appears to be closely
related to L. protentus, a species associated with Sarcophyton in
Madagascar. In both there are striking similarities in the mandible,
the maxilliped of the female, the nature of the outer spines on the
exopod of leg 1, and the armature of the endopod of leg 4. The new
species differs from L. protentus, however, in several important features:
its smaller size, the proportions of the genital segment in the female,
details of the mouthparts, the presence of only two outer spines on
the last segment of the exopod of leg 3, the unornamented leg 5 in
the female, the membranous lamella on the second segment of the second
antenna in the male, and the series of hyaline knobs on the claw of the
maxilliped of the male.
Like L. protentus, L. insolens seems to approach L. robustus Thompson
and A. Scott, 1903, described from one female found in washings of
dredged invertebrates in Ceylon. However, in L. robustus the formula
for the last segment of the exopod of leg 4 is III,I,5, and the two terminal
elements on the endoped of this leg appear to be different (T. and S.,
PI. XVI, fig. 20).
Although interspecific variation in the number of outer spines on the
last segment of the exopod of leg 4 occurs in Lichomolgus (the formula
for the segment being either III,I,5 or II,1,5), it is unusual that the
corresponding segment of leg 3 should show a reduction from III,1,5
to II,I,5, as is the case in the new species. Another species, L. curtiramus
Bocquet and Stock, 1962, shows a similar armature on the third segment
of leg 3.
Lichomolgus spathophorus new species
Figures 128-147
Type material: 139 9 and 172 ¢ from two colonies of Sarcophyton
glaucum (Quoy and Gaimard), in 0.5 m, Tany Kely, a small island
south of Nosy Bé, Madagascar, collected 23 June 1963. Holotype 9,
allotype, and 18 paratypes (79 2, 11é¢ 4) deposited in the United
States National Museum, and the remaining paratypes in the collection
of A. G. Humes.
Female: Body (fig. 128) with expanded prosome. Length 0.98 mm
(0.95-1.04 mm) and greatest width 0.59 mm (0.57-0.61 mm), based
on 10 specimens. Ratio of length to width of prosome 1.17: 1. Seg-
ment of leg 1 distinctly separated from head. Epimeral areas of seg-
ments of legs 1-4 as in figure.
Segment of leg 5 (fig. 129) 60 uw xX 146 uw. Between this segment
and genital segment a short ventral intersegmental sclerite. Genital
(Bay
Copepods Associated with Octocorals 675
Ay
i I;
|
Uh
se AT
AON JN
UK |
ee, > )
Fics. 128-134. Lichomolgus spathophorus, new species, female: 128,
body, dorsal (A); 129, urosome, dorsal (B); 130, area of attachment
of egg sac, dorsal (C); 131, caudal ramus, dorsal (C); 132, rostral area,
ventral (B); 133, second antenna, posterior (outer) (D); 134, mandible,
posterior (E).
segment (fig. 129) 117 uw x 140 wu, wider than long, a little broadened
in its anterior four-fifths in dorsal view, but abruptly narrowed in its
posterior fifth. Areas of attachment of egg sacs located almost laterally
in posterior half of segment. Each area (fig. 130) with two naked
spiniform setae 10 uw and 8 wu long and a large pointed bladelike process.
676 Proceedings of the Biological Society of Washington
i
139 iN 136
at a
\y) 7 Fal!
\ “Me A
Fics. 135-141. Lichomolgus spathophorus, new species, female: 135,
second maxilla, posterior (EK); 136, maxilliped, posterior (E); 137, area
between maxillipeds and first pair of legs, ventral (D); 138, leg 1 and
intercoxal plate, anterior (D); 139, leg 2, anterior (D); 140, last
segment of endopod of leg 3, anterior (D); 141, leg 4, anterior (D).
Three postgenital segments 26 » X 89 yu, 21 uw X 83 uw, and 31 uw Xx 78 4,
from anterior to posterior.
Caudal ramus (fig. 131) quadrate, 30 u« x 30 u in greatest dimensions.
Outer lateral seta 44 » and naked, pedicellate dorsal seta 45 m and
feathered, outermost distal seta 104 « with proximal inner spinules,
Copepods Associated with Octocorals 677
Fics. 142-148. Lichomolgus spathophorus, new species, female: 142,
leg 5, dorsal (F); 143, leg 5, dorsal (F). Male: 144, body, dorsal (A);
145, urosome, dorsal (B); 146, maxilliped, inner (E); 147, last segment of
endopod of leg 1, anterior (E). Lichomolgus incisus, new species, fe-
male: 148, body, dorsal (A).
innermost distal seta 180 u with spinules along both sides proximally,
and the two long median terminal setae 375 » (outer) with spinules
on inner midregion only or along both sides and 500 uw (inner) with
spinules along both sides proximally. Both terminal setae inserted
between dorsal (unornamented) and ventral (with a marginal row of
spinules ) flaps.
678 Proceedings of the Biological Society of Washington
Dorsal surface of prosome and urosome with a few small hairs.
Ratio of length of prosome to that of urosome 2.58 : 1.
Egg sacs broken in all ovigerous females seen, but apparently elongated
and containing many small eggs.
Rostral area (fig. 132) with broadly rounded posteroventral margin.
First antenna similar to that of L. foxi. Lengths of segments: 35 yu
(71 uw along anterior margin), 143 uw, 23 uw, 64 uw, 55 uw, 39 w, and 27 uw
respectively. All setae naked. Armature as in EL. foxi.
Second antenna (fig. 133) 4-segmented and slender. Last segment
elongated, 122 w along its outer edge, 94 uw along its inner edge, and
18 » wide, bearing distally five small hyaline elements and a claw 58 u
along its axis. All setae naked.
Labrum resembling that of L. foxi. Mandible (fig. 134) much like
that of L. foxi, but with scalelike process on convex basal region more
acutely pointed. Paragnath and first maxilla as in L. foxi. Second
maxilla (fig. 135) 2-segmented, and armed as in figure. Maxilliped
(fig. 136) 3-segmented, with armature resembling in general that of
L. foxi.
Area between mazxillipeds and first pair of legs (fig. 137) not
protuberant; a sclerotized line between bases of maxillipeds.
Legs 1-4 (figs. 138-141) segmented as in previous species, with
spine and setal formula as in L. hetaericus. Inner seta on coxa of legs
1-3 long and plumose, but in leg 4 short (18 «), somewhat spiniform,
and finely barbed. Inner distal corner of coxa of leg 1 slightly
protuberant. Inner margin of basis of legs 1-4 with hairs. Outer
spines on exopod of leg 1 with coarse spinulation along proximal margins.
Endopod of leg 4 (fig. 141) a little shorter than exopod. First segment
33 uw X 25 w (without processes), its inner distal seta relatively short
(37 mw) and somewhat spiniform, with hairs proximally but naked
distally. Second segment 85 uw long (including processes), 23 m in
greatest width, and 17 uw in least width. Two unequal terminal spines
26 » (outer) with a delicate fringe and 54 w (inner) with a coarsely
serrated hyaline fringe. Hairs along outer margins of both segments
and short hairs on inner margin of second segment. A row of minute
spinules near insertions of terminal spines.
Leg 5 (fig. 142) with elongated free segment, 122 uw x 20 wu, having
a slight proximal inner expansion, where width is 22 yu. Outer convex
surface with numerous small spines. Two naked terminal setae 57 mu
(outer) and 81 uw (inner). Plumose seta on body near free segment
38 uw. Apparent variation in size of proximal expansion, some females
showing free segment of leg 5 as in fig. 143, where dimensions of
segment are 112 uw X 16 uw, and width at expansion 27 uw. Other females
showing expansion intermediate in size between those in figs. 142
and 143.
Leg 6 probably represented by the two setae near areas of attachment
of each egg sac (see fig. 130).
hae
Wee
Copepods Associated with Octocorals 679
Color in life in transmitted light slightly amber, eye red, egg sacs
opaque gray.
Male: Body (fig. 144) with prosome expanded nearly as in female.
Length 0.79 mm (0.72—0.86 mm) and greatest width 0.39 mm (0.33-
0.44 mm), based on 10 specimens. Ratio of length to width of prosome
1.26: 1.
Segment of leg 5 (fig. 145) 33 «x 105 uw. A very short ventral
intersegmental sclerite. Genital segment 174 uw x 187 uw. Four post-
genital segments 22 uw X 60 w, 22 uw xX 64 uw, 18 w xX 64 pw, and 29 ux
66 uw, from anterior to posterior.
Caudal ramus as in female, 25 uw x 24 u.
Surface of prosome and urosome with a few small hairs. Ratio of
length of prosome to that of urosome 2.0: 1.
Rostral area as in female. First antenna similar to that of L. foxi,
with three aesthetes added as in that species. Second antenna as in
female, but with small spinules on inner surfaces of first, second, and
fourth segments.
Labrum, mandible, paragnath, first maxilla, and second maxilla as in
female. Maxilliped (fig. 146) moderately stout and armed as in
previous species, with a long claw 151 w along its axis.
Area between maxillipeds and first pair of legs as in female.
Legs 1-4 segmented as in female and with same spine and setal
formula, except for last segment of endopod of leg 1 (fig. 147) which
has the arrangement of I,],4. Legs 2-4 as in female.
Leg 5 (see fig. 145) with free segment 62 «x8 w, without a
proximal expansion, and with extremely small spinules on outer surface.
Two terminal naked setae 58 » and 33 uw. Seta on body near free seg-
ment 33 p.
Leg 6 (see fig. 145) a posterolateral flap on ventral surface of
genital segment bearing two naked setae 50 yu long.
Spermatophore not observed.
Color in life similar to that of female.
Etymology: The specific name spathophorus, from Greek oré6n =a
broad blade and gopéw = to carry, refers to the bladelike process on the
area of attachment of each egg sac.
Comparison with related species: L. spathophorus may be distin-
guished from all but two species of Lichomolgus on the basis of a
combination of three features: the single short claw on the second
antenna, the formula of II,I,5 for the third segment of the exopod of
leg 4, and the short quadrate caudal ramus. The remaining two species
have other characters that show them to be distinct from L. spathophorus.
L. anomalus A. Scott, 1909, differs in having the endopod of leg 1
in the male geniculate between the second and third segments, the
third segment being elongated (about 3:1 in A. Scott’s Pl. LXVII,
fig. 14). L. elegans Thompson and A. Scott, 1903, of which the male
is unknown, is larger (female 1.5 mm), has a less expanded prosome,
680 Proceedings of the Biological Society of Washington
and the genital segment has different proportions and shape than in
the new species.
Lichomolgus incisus new species
Figures 148-167
Type material: 2499, 104 6, and 3 copepodids from one colony
of Sarcophyton ehrenbergi Marenzeller, in 0.5 m, Andilana, Nosy Bé,
collected 9 August 1963. Holotype ¢@, allotype, and 24 paratypes
(199 2, 54 4) deposited in the United States National Museum, and
the remaining paratypes in the collection of A. G. Humes.
Female: Body (fig. 148) with expanded and relatively short
cephalosome, its anterior border broadly rounded in dorsal view.
Length 1.03 mm (0.90-1.08 mm) and greatest width 0.42 mm (0.40-
0.43 mm), based on 10 specimens. Ratio of length to width of prosome
1.41: 1. Segment of leg 1 separated from head by an incomplete dorsal
transverse furrow. Epimeral areas of legs 1-4 rounded.
Segment of leg 5 (fig. 149) 52 «x 117 uw. Between this segment
and genital segment a short ventral intersegmental sclerite. Genital
segment (fig. 149) only a little wider than long, 114 u x 121 uw. Areas
of attachment of egg sacs situated dorsolaterally in anterior half of
segment. Each area (fig. 150) bearing two minute naked setae about
4 w long. Three postgenital segments 52 u X 68 uw, 44 uw xX 63 yw, and
56 uw X 61 u, from anterior to posterior.
Caudal ramus (fig. 151) elongated, 77 u long, 24 uw wide near base,
and 19 uw wide distally. Setae relatively short. Outer lateral seta 35 u,
dorsal pedicellate seta 28 u, outermost distal seta 33 u, innermost distal
seta 82 uw, and the two long median terminal setae 105 uw (outer) and
150 » (inner), both inserted between unornamented dorsal and ventral
flaps. All setae naked.
Dorsal surface of prosome and urosome with a few small hairs. Ratio
of length of prosome to that of urosome 1.42: 1.
Egg sac in one ovigerous female elongated, 450 u x 175 yu, reaching
just beyond ends of caudal rami and containing many eggs about 52 w
in diameter.
Rostral area as in L. cristatus.
First antenna (fig. 152) 7-segmented, but relatively short (232 uw).
Armature as in all previous species. Lengths of segments: 25 wu (41 pu
along anterior margin), 61 uw, 21 uw, 31 uw, 30 uw, 21 pw, and 17 uw respec-
tively. All setae naked.
Second antenna (fig. 153) 4-segmented. Last segment relatively
short, 46 » along its outer edge, 31 uw along its inner edge, and 19 uw
wide, bearing distally four small hyaline elements and a terminal claw
33 » along its axis. All setae naked.
Labrum as in L. foxi. Mandible (fig. 154) resembling that of L.
spathophorus, but scalelike process on convex side of basal region not
pointed distally. Paragnath and first maxilla as in L. protentus. Second
Copepods Associated with Octocorals 681
Fics. 149-157. Lichomolgus incisus, new species, female: 149,
urosome, dorsal (B); 150, area of attachment of egg sac, dorsal (C);
151, caudal ramus, dorsal (E); 152, first antenna, ventral (F); 153,
second antenna, posterior (outer) (F); 154, mandible, posterior (E);
155, second maxilla, posterior (E); 156, maxilliped, posterior (E); 157,
area between maxillipeds and first pair of legs, ventral (D).
maxilla (fig. 155) resembling that of L. spathophorus. Maxilliped (fig.
156) 3-segmented. Second segment with two unequal naked setae, the
longer seta only about four times the length of the other. Third
segment with the usual two terminal spiniform elements (one without
an articulation) and a small subterminal seta; outer side of segment
swollen and membranous.
682 Proceedings of the Biological Society of Washington
wa
Fi
|
]
Jo \\
\
Fics. 158-164. Lichomolgus incisus, new species, female: 158, leg 1
and intercoxal plate, anterior (F); 159, leg 2, anterior (F); 160, last
segment of endopod of leg 3, anterior (E); 161, leg 4 and intercoxal
plate, anterior (F); 162, leg 5, dorsal (C). Male: 163, body, dorsal
(A); 164, urosome, dorsal (B).
Area between maxillipeds and first pair of legs (fig. 157) not pro-
tuberant; a sclerotized line between bases of maxillipeds.
Legs 1-4 (figs. 158-161) segmented as in previous species, with
spine and setal formula as follows:
Pi: protopod O0-1 1-0 exp I-O I-1 IILI,4
end OQ-l1 O-1 I5
Copepods Associated with Octocorals 683
Pz protopod 0-1 1-0 exp I-0O J[-1 III,I,5
end 0-1 0-2 LJII,3
Ps; protopod 0-1 1-0 exp I-O I-1 ILLS (or I1,1,5)
end 0-1 0-2 L,II,2
Ps protopod OQ-1 1-0 exp I-O I-1 IL,15
end 0-1 1,I
Inner seta on coxa of legs 1-3 long and plumose, but in leg 4 short
(5 w) and naked. Inner margin of basis of legs 1-4 without row of
hairs, though a minute setule (sensillum ?) may be present. Third
segment of exopod of leg 3 with II,I,5 or IJI,I,5 (on either or both
legs of same individual), though most frequently II,I,5 (in 7 of 8 legs
in 4 females). Endopod of leg 4 (fig. 161) shorter than exopod. First
segment 25 u X 17 uw, with inner distal seta short (19 w) and naked.
Second segment 39 u X 17 yu, its two terminal elements being an outer
naked seta 29 w and an inner barbed spine 18 uw. Both segments with
hairs along outer margins and second segment with a row of spinules
near insertions of terminal elements.
Leg 5 (fig. 162) with small free segment 34 ux 15 uw, tapering
slightly distally, without proximal expansion and unornamented. Two
terminal setae 38 uw (outer) and 42 w (inner). Seta on body near
insertion of free segment 36 uw. All setae naked.
Leg 6 probably represented by the two setae near areas of attachment
of each egg sac (see fig. 150).
Color in life in transmitted light translucid except for slightly opaque
cephalosome, eye red, egg sacs opaque gray.
Male: Body (fig. 163) resembling in general form that of female.
Length 0.90 mm (0.86—0.92 mm) and greatest width 0.29 mm (0.29-
0.30 mm), based on 10 specimens. Ratio of length to width of prosome
1.69 : 1.
Segment of leg 5 (fig. 164) 44 u x 88 uw. No ventral intersegmental
sclerite. Genital segment 156 uw xX 146 uw, subquadrate in dorsal view.
Four postgenital segments 44 u X55 uw, 44 wu xX 49 w, 34 w X 47 pw, and
49 w xX 49 uw, from anterior to posterior.
Caudal ramus as in female, but slightly shorter, 68 u x 19 u.
Surface of prosome and urosome with a few small hairs. Ratio of
length of prosome to that of urosome 1.12: 1.
Rostral area as in female. First antenna similar to that of female but
three aesthetes added, so that formula is same as for males of previous
species. Second antenna resembling that of female but second segment
(fig. 165) bearing a notched lamella along inner surface.
Labrum, mandible, paragnath, first maxilla, and second maxilla like
those of female. Maxilliped (fig. 166) with only one row of spines
along inner surface of second segment; claw relatively short, 115 u
along its axis (including terminal lamella), and not strongly recurved.
Area between maxillipeds and first pair of legs as in female.
AS
Fics. 165-171. Lichomolgus incisus, new species, male: 165, second
segment of second antenna, posterior (outer) (C); 166, maxilliped,
inner (F); 167, last segment of endopod of leg 1, anterior (E).
Lichomolgus singularipes, new species, female: 168, body, dorsal (A);
169, urosome, dorsal (D); 170, area of attachment of egg sac, dorsal
(C); 171, caudal ramus, dorsal (C).
Legs 1-4 segmented as in female and with same spine and setal
formula. Sexual dimorphism showing in last segment of endopod of leg
1 (fig. 167), where terminal spiniform process is much longer than in
female. Last segment of endopods of legs 2 and 3 also with longer
terminal processes than in female.
Copepods Associated with Octocorals 685
Leg 5 (see fig. 164) with free segment 24 uw x 8 wu, unornamented,
its two terminal setae 25 w and 35 uw. Seta on body near free segment
25 uw. All setae naked.
Leg 6 (see fig. 164) a posterolateral flap on ventral surface of genital
segment bearing two naked setae 22 uw and 24 wu.
Spermatophore not observed.
Color in life similar to that of female, but cephalosome more
translucid.
Etymology: The specific name incisus, from Latin = cut into or
notched, alludes to the notched lamella on the second segment of the
second antenna of the male.
Comparison with related species: Two features of L. incisus may be
used as recognition characters,—ihe swollen and membranous outer
side of the last segment of the maxilliped in the female and the notched
lamella along the inner surface of the second segment of the second
antenna of the male. These occur in no other known species of
Lichomolgus.
The new species has several features which suggest a relationship to
L. insolens. Both species have a reduction in the number of outer
spines on the third segment of the exopod of leg 3, the terminal armature
of the endopod of leg 4 consisting of a spine and a seta, and a lamella
or membrane along the inner surface of the second segment of the
second antenna of the male.
Lichomolgus singularipes new species
Figures 168-187
Type material: 409 9 from several encrusting colonies of Parery-
thropodium rubiginosum Verseveldt, in 2 m, Ambariobe, a small island
almost between Nosy Komba and Nosy Bé, Madagascar, collected 4
October 1964. Holotype and 30 paratypes deposited in the United States
National Museum, and the remaining paratypes in the collection of
A. G. Humes.
Female: Body (fig. 168) with rather slender prosome. Length 0.95
mm (0.88-1.01 mm) and greatest width 0.40 mm (0.39-0.42 mm),
based on 10 specimens. Ratio of length to width of prosome 1.86: 1.
Segment of leg 1 separated from head by a dorsal transverse furrow.
Epimeral areas of segments of legs 1-4 rounded.
Segment of leg 5 (fig. 169) 57 1» x 120 uw. Between this segment
and genital segment a very short ventral intersegmental sclerite. Genital
segment a little longer than wide, 120 u x 107 yu, in dorsal view not
greatly expanded, with its posterior eighth set off by transverse dorsal
ridges and at first glance suggesting a postgenital segment. Areas of
attachment of egg sacs situated dorsolaterally just posterior to middle
of segment. Each area (fig. 170) with two spiniform naked setae 11 yu
and 17 uw long. Three postgenital segments 26 » X 68 py, 21 wu xX 63 u,
and 29 u x 61 wu, from anterior to posterior.
686 Proceedings of the Biological Society of Washington
Fics. 172-179. Lichomolgus singularipes, new species, female: 172,
first antenna, ventral (D); 173, second antenna, posterior (outer) (D);
174, labrum, ventral (F); 175, mandible, posterior (E); 176, paragnath,
posterior (C); 177, first maxilla, posterior (EK); 178, second maxilla,
posterior (E.); 179, maxilliped, anterior (E).
Caudal ramus (fig. 171) quadrate, 24 w x 25 uw in greatest dimensions.
Outer lateral seta 83 w and naked, dorsal pedicellate seta 40 mw and
feathered, outermost distal seta 95 u and naked, innermost distal seta
161 « with proximal spinules, and the two long median terminal setae
275 w (outer) and 360 uw (inner) with strong spinules except near tips,
Copepods Associated with Octocorals 687
Fics. 180-187. Lichomolgus singularipes, new species, female: 180,
area between maxillipeds and first pair of legs, ventral (D); 181, leg 1
and intercoxal plate, anterior (D); 182, leg 2, anterior (D); 183, last
segment of endopod of leg 3, anterior (D); 184, leg 4 and intercoxal
plate, anterior (D); 185, leg 5, dorsal (E); 186, spermatophore, empty
and attached to female, dorsal (D); 187, Ophryodendron sp., attached
to caudal ramus, dorsal (B).
both inserted between dorsal (unornamented) and ventral (with a mar-
ginal row of very small spinules) flaps.
Dorsal surface of prosome and urosome with a few small hairs. Ratio
of length of prosome to that of urosome 2.62 : 1.
688 Proceedings of the Biological Society of Washington
Egg sac (fig. 168) elongated, 500 u <x 200 uw, reaching almost to
tips of ramal setae, containing numerous (fewer than in previous species )
moderately large eggs about 60 u in diameter.
Rostral area as in L. spathophorus.
First antenna (fig. 172) 7-segmented and armed as in all previous
species. Setae naked and rather long. Lengths of segments: 33 yu
(61 w along anterior margin), 115 uw, 22 uw, 61 w, 51 uw, 40 w, and 23 up
respectively. ;
Second antenna (fig. 173) 4-segmented. Last segment 80 wu along its
outer edge, 53 uw along its inner edge, and 20 u wide, bearing distally
five hyaline setae (three of them long) and two unequal terminal claws,
the stouter claw 36 uw along its axis, the more slender claw 43 u. All
setae naked.
Labrum (fig. 174) with two moderately elongated lobes. Mandible
(fig. 175) similar to that of L. foxi. Paragnath (fig. 176) a small hairy
lobe. First maxilla (fig. 177) with three terminal elements and one
subterminal. Second maxilla (fig. 178) with lash bearing several large
proximal teeth, followed by a row of small spines; tip of lash naked.
Maxilliped (fig. 179) resembling in general form that of L. commodus
Humes, 1964, with two setae on second segment extremely unequal,
the shorter seta naked, the longer one with a row of 6-9 long erect
spinules. Last segment with two rather slender spiniform terminal
elements and a smail seta. Ornamentation on first two segments of
maxilliped as in figure.
Area between maxillipeds and first pair of legs (fig. 180) not
protuberant; a sclerotized line between bases of maxillipeds.
Legs 14 (figs. 181-184) segmented as in previous species, with
spine and setal formula as in L. hetaericus and L. spathophorus (the
formula for last segment of exopod of leg 4 being II,I,5). Inner seta
on coxa of legs 1-3 long and plumose, but in leg 4 shorter (23 ) and
naked. Inner margin of basis of legs 1-3 with row of hairs, but in leg
4 naked. Outer seta on basis of leg 2 shorter than in other legs and
naked. Endopod of leg 4 (fig. 184) shorter than exopod. First segment
33 « X 26 uw (not including processes), its inner distal feathered seta
50 uw. Second segment 70 uw X 21 uw (length including processes and
width taken at middle), its two unequal terminal barbed spines 32 uw
(outer) and 67 u (inner). Both segments with hairs along outer margins
and second segment with a row of spinules near insertions of terminal
spines.
Leg 5 (fig. 185) with rather broad free segment, 81 u xX 34 uw in
greatest dimensions, its outer surface with scalelike spines and its inner
margin abruptly expanded a short distance from distal end of segment,
the contour of margin from this point to base of segment being irregular
as in figure. Two terminal naked setae 68 » (outer) and 82 wu (inner).
Seta on body near insertion of free segment 44 w and naked. A row of
small spines near this seta.
oe al
Copepods Associated with Octocorals 689
Leg 6 probably represented by the two setae near areas of attachment
of each egg sac (see fig. 170).
Spermatophore (fig. 186), attached to female and empty, elongated,
125 uw X 60 uw (not including neck).
Color in life in transmitted light slightly opaque, eye red, ovary gray,
egg sacs gray.
Male: Unknown.
Etymology: The specific name singularipes; from Latin singularis =
remarkable, extraordinary, and pes = foot, refers to the unusual form of
leg 5.
Comparison with related species: L. singularipes may be distinguished
by the form of leg 5 from all other species in the genus having similarly
a quadrate caudal ramus, two short claws on the second antenna, and
the formula II,I,5 on the last segment of the exopod of leg 4. The
maxilliped of the new species resembles that of L. commodus Humes,
1964, but leg 5 in the latter is very different (ratio about 4:1 with a
slight proximal inner expansion).
L. singularipes cannot be directly compared with L. vagans Gurney,
1927, and L. aegyptius Gurney, 1927, of which only males are known.
In L. vagans the length of the male is 1.07 mm, the second antenna is
unusually long and slender, and the caudal ramus is a little longer than
wide: In the male of L. aegyptius the second antenna has terminally
three claws and two setae and the form of the mandible (Gumey’s
fig. 112D) is distinctly different from that in L. singularipes. It is
very unlikely, therefore, that the new species could be the same as
either of Gurney’s species.
Epibiotic protozoan on L. singularipes: Thirteen of the 40 females
collected (33 per cent) had large suctorians, identified by Dr. John
O. Corliss as Ophryodendron sp., attached to the caudal ramus. These
epibionts were not specially stained, but observed only incidentally in
dissections in lactic acid. The greatest dimensions of the specimen
drawn (fig. 187), which shows three buds, were 275 uw X 78 yp, the
length being almost one-third that of the copepod.
The suctorians were attached always to the dorsal surface of the caudal
rami, and there was never more than one on each ramus. They were
distributed on the 13 females as follows: 8 copepods had one on the
right caudal ramus, 4 copepods had one on the left ramus, and one
copepod had two suctorians, one on each ramus.
Since only female copepods were collected, it is not known whether
male copepods also bear these suctorians.
Lichomolgus squamiger Humes and Frost, 1964
This species has been previously recorded from Sinularia polydactyla
(Ehrenberg) at Nosy Bé, Madagascar (Humes and Frost, 1964).
New host record: From Sinularia whiteleggei Liittschwager: 369 9,
2446 6, and 19 copepodids from one colony, in 2 m, Pte. Lokobe, Nosy
Bé, 16 August 1960.
690 Proceedings of the Biological Society of Washington
Two minor additions should be made to the original description by
Humes and Frost. In their figure 174 of the second antenna of the
female, there are six small hyaline elements near the terminal claw, and
in their figure 180 of the maxilliped of the female there is a small seta
on the last segment in addition to the two terminal elements. These
features have been verified by studying a paratype of L. squamiger.
LITERATURE CITED -
BocguEt, C. ann J. H. Srocx. 1962. Copépodes parasites d’in-
vertébrés des cdtes de France. XIV. Description d’un
nouveau copépode cyclopoide, Lichomolgus curtiramus n. sp.
Koninkl. Nederl. Akad. Wetensch.—Amsterdam, Proc., ser.
C, vol. 65, no. 3, pp. 244-249.
Gurney, R. 1927. Zoological results of the Cambridge expedition to
the Suez Canal, 1924, XXXIII: Report on the Crustacea:
Copepoda (Littoral and semi-parasitic). Trans. Zool. Soc.
London, vol. 22, pt. 4, pp. 451-577.
Humes, A. G. 1964. New species of Lichomolgus (Copepoda, Cyclo-
poida) from sea anemones and nudibranchs in Madagascar.
Cahiers O.R.S.T.O.M. Océanogr., 1963 (série Nosy Bé II),
no. 6, pp. 59-130.
Humes, A. G. anp R. W. Frost. 1964. New lichomolgid copepods
(Cyclopoida) associated with alcyonarians and madrepo-
rarians in Madagascar. Cahiers O.R.S.T.O.M. Océanogr.,
1963 (série Nosy Bé II), no. 6, pp. 131-212.
Humes, A. G. ann J.-S. Ho. 1967. New species of Stellicola (Copepoda,
Cyclopoida) associated with starfishes in Madagascar, with
a redescription of S. caeruleus (Stebbing, 1900). Bull. Brit.
Mus. (Nat. Hist.), Zool., vol. 15, no. 5, pp. 201—295.
KossMANN, R. 1877. Entomostraca (I. Theil:Lichomolgidae). In:
Zool. Ergeb. Reise Kiistengeb. Rothen Meeres, Erste Halfte,
IV, pp. 1-24.
Scott, A. 1909. The Copepoda of the Siboga expedition. Part I.
Free-swimming, littoral and semi-parasitic Copepoda. Siboga
Exped., 29a, pp. 1-323.
SEWELL, R. B. S. 1949. The littoral and semi-parasitic Cyclopoida,
the Monstrilloida and Notodelphyoida. No. 2 of vol. 9 in
John Murray Expedition 1933-34, Scientific Reports, pp.
17-199.
Tuompson, I. C. anp A. Scorr. 1903. Report on the Copepoda col-
lected by Professor Herdman, at Ceylon, in 1902. Rept.
Gov. Ceylon Pearl Oyster Fish. Gulf of Manaar, suppl. rept.
no. 7, pp. 227-307.
THORELL, T. 1859. Till kannedomen om vissa parasitiskt lefvande
Entomostraceer. Ofversigt K. Vetensk.-Akad. Forhandl.,
Arg. 16, no. 8, pp. 335-362.
Copepods Associated with Octocorals 691
1860. Bidrag till kinnedomen om krustaceer, som lefva i
arter af slagtet Ascidia L. K. Svenska Vetensk. Akad. Handl.,
n. f., vol. 3, no. 2, pp. 1-84.
TrxreR-DurRIvVAULT, A. 1966. Octocoralliaires. Faune de Madagascar,
fasc. 21, pp. 1-456.
Ummerkutty, A. N. P. 1962. Studies on Indian copepods 5. On
eleven new species of marine cyclopoid copepods from the
south-east coast of India. J. Mar.: Biol. Ass. India, 1961,
vol. 3, nos. 1 & 2, pp. 19-69.
692 Proceedings of the Biological Society of Washington
Vol. 81, pp. 693-750 30 December 1968
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
CYCLOPOID COPEPODS OF THE GENUS
LICHOMOLGUS ASSOCIATED WITH OCTOCORALS
OF THE FAMILIES XENIIDAE, NIDALITDAE, AND
TELESTIDAE IN MADAGASCAR
By ArtHur G. Humes anp Ju-SHey Ho
Dept. of Biology, Boston University, Boston, Mass.
In two previous papers (Humes and Ho, 1968b and 1968c)
several new species of Lichomolgus were described, all
associated with octocorals of the families Nephtheidae and
Alcyoniidae in the region of Nosy Bé, in northwestern Mada-
gascar. This paper deals with seven new species associated
with the aleyonacean families Xeniidae ( Anthelia, Heteroxenia,
and Xenia) and Nidaliidae (Siphonogorgia) and with the
telestacean family Telestidae (Telesto and Coelogorgia) from
the same geographical area.
All collections were made by A. G. Humes, those in 1960
during an expedition sponsored by the Academy of Natural
Sciences of Philadelphia, and those in 1963-64 as part of the
U. S. Program in Biology of the International Indian Ocean
Expedition.
The study of the specimens has been aided by a grant
(GB-5838) from the National Science Foundation of the
United States.
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. The abbreviations used are:
A, = first antenna, Az = second antenna, MXPD = maxilliped,
and P; = leg 1.
All descriptions are based on type material. The measure-
ments of the length of the body have been made in all cases
from specimens in lactic acid and do not include the setae on
60—Proc. Biou. Soc. Wasu., Vou. 81, 1968 (693)
694 Proceedings of the Biological Society of Washington
the caudal rami. The lengths of the segments of the first
antennae have been measured along their posterior non-
setiferous margins.
We are indebted to Mme. A. Tixier-Durivault of the
Muséum National d’Histoire Naturelle, Paris, for the identifica-
tions of the octocorals collected in 1960, and to Dr. J. Verse-
veldt, Zwolle, The Netherlands, for the determinations of those
collected in 1963-64.
The new copepods described below are as follows:
1. Lichomolgus verseveldti new species
from Heteroxenia elisabethae Kolliker.
2. Lichomolgus triquetrus new species
from Anthelia gracilis (May).
3. Lichomolgus glabripes new species
from Xenia umbellata Lamarck.
4, Lichomolgus longispinifer new species
from Siphonogorgia pendula Studer.
5. Lichomolgus hians new species
from Siphonogorgia pendula Studer.
6. Lichomolgus telestophilus new species
from Telesto arborea Wright and Studer.
7. Lichomolgus clavatus new species
from Coelogorgia palmosa Milne Edwards and
Haime.
FAMILY LICHOMOLGIDAE KOSSMANN, 1877
Genus Lichomolgus Thorell, 1859
Lichomolgus verseveldti new species
Figures 1—26
Type material: 929 2, 254 4, and 7 copepodids from colonies of
Heteroxenia elisabethae Kolliker, in 1 m, Pte. Lokobe, Nosy Bé, Mada-
gascar, collected 13 August 1960. Holotype 2, allotype, and 81 paratypes
(652 2, 164 6) deposited in the United States National Museum, and
the remaining paratypes in the collection of A. G. Humes.
Other specimens (all from Heteroxenia elisabethae): 1699, 46 4,
and 3 copepodids, in 1 m, Pte. Lokobe, 12 August 1960; 149 9, 1
copepodid, in 1 m, Pte. Lokobe, 1 November 1960; and 402 9, 254 ¢,
and 3 copepodids, in 1 m, Pte. Ambarionaomby, Nosy Komba, near Nosy
Bé, 27 September 1964.
Copepods Associated with Octocorals 695
6
a
ae
a\ a
ke
:
0
al
\
AVA\\
\
Le =
0.05 MM
Cc
dorsal (A); 2, urosome, dorsal (B); 3, area of attachment of egg sac,
dorsal (C); 4, caudal ramus, dorsal (C); 5, rostral area, ventral (D);
6, first antenna, ventral (D); 7, second antenna, posterior (outer) (FE).
Female: Body (fig. 1) with prosome not unusually broadened.
Length 1.20 mm (1.12-1.25 mm) and greatest width 0.54 mm (0.51-—
0.58 mm), based on 10 specimens. Ratio of length to width of prosome
1.47: 1. Segment of leg 1 distinctly separated from head dorsally by a
696 Proceedings of the Biological Society of Washington
Fics. 8-16. Lichomolgus verseveldti, new species, female: 8, labrum, —
ventral (F); 9, mandible, posterior (C); 10, paragnath, ventral (C);
11, first maxilla, posterior (C); 12, second maxilla, posterior (F); 13,
maxilliped, posterior (F ); 14, area between maxillipeds and first pair of
legs, ventral (D); 15, leg 1 and intercoxal plate, anterior (D); 16, leg —
2, anterior (D).
Copepods Associated with Octocorals 697
Fics. 17-22. Lichomolgus verseveldti, new species, female: 17, third
segment of endopod of leg 3, anterior (E); 18, leg 4 and intercoxal
plate, anterior (D); 19, leg 5, dorsal (E). Male: 20, body, dorsal (G);
21, urosome, dorsal (D); 22, first antenna, ventral (E).
transverse furrow. Epimeral areas of segments of legs 1-4 as shown
in figure.
Segment of leg 5 (fig. 2) 68 u x 187 uw. Between this segment and
genital segment a slight ventral intersegmental sclerite. Genital segment
698 Proceedings of the Biological Society of Washington
0.03 MM
H 0.02 MM
Fics. 23-30. Lichomolgus verseveldti, new species, male: 23, maxil-
liped, inner (F); 24, outer element on second segment of maxilliped,
posterior (H); 25, endopod of leg 1, anterior (C); 26, spermatophores,
attached to genital segment of female, lateroventral (B). Lichomolgus
triquetrus, new species, female: 27, body, dorsal (A); 28, urosome,
dorsal (D); 29, area of attachment of egg sac, dorsal (I); 30, caudal
ramus, dorsal (I).
Copepods Associated with Octocorals 699
only slightly longer than wide, 200 » x 190 u, widest in its anterior half
and narrowed posteriorly, where the dorsal surface has two transverse
lines simulating a segment but the ventral surface is smooth. Areas of
attachment of egg sacs lateral in position just behind widest portion of
segment. Each area (fig. 3) bearing two naked setae about 12 yu long.
Three postgenital segments 55 uw & 101 uw, 34 uw x 91 uw, and 40 uw x 90 yu,
from anterior to posterior.
Caudal ramus (fig. 4) slightly wider than long, 31 « x 34 u in greatest
dimensions. Outer lateral seta 86 » long and naked, pedicellate dorsal
seta 33 » and naked, outermost distal seta 130 « and naked, innermost
distal seta 260 u with lateral spinules (mostly along inner edge), and
the two long median terminal setae 495 » (outer) and 620 uw (inner),
both with strong lateral spinules and both inserted between unornamented
dorsal and ventral flaps. A minute setule on proximal outer margin of
ramus.
Dorsal surface of prosome without hairs; surface of urosome with only
a few hairs and refractile points. Ratio of length of prosome to that of
urosome 2.25; 1.
Egg sac (fig. 1) elongated, 570 u x 230 uw, with many eggs, each
about 50 uw in diameter.
Rostral area (fig. 5) with a well defined broadly rounded postero-
ventral margin.
First antenna (fig. 6) 7-segmented, 495 u long. Lengths of segments:
50 » (77 uw along anterior margin), 138 uw, 35 uw, 78 uw, 74 uw, 56 w, and 37 uw
respectively. Formula for armature: 4, 13, 6, 3, 4+ 1 aesthete, 2+ 1
aesthete, and 7+ 1 aesthete. All setae naked.
Second antenna (fig. 7) 4-segmented, last segment moderately
elongated, 90 u along its outer edge, 57 u along its inner edge, 23 » wide,
bearing distally five small hyaline elements and two slightly unequal
claws 55 « and 46 uw in length. Distalmost of three setae on third segment
with a blunt tip. All setae naked.
Labrum (fig. 8) with two posteroventral lobes. Mandible (fig. 9)
with basal region distal to constriction showing on its convex margin a
scalelike protrusion ornamented with a row of spinules, followed by a
serrated fringe, and on its concave margin a row of slender spinules.
Flagellum rather short and coarsely barbed. Paragnath (fig. 10) a
small hairy lobe. First maxilla (fig. 11) with three long terminal and
one short subterminal setae, all naked. Second maxilla (fig. 12) 2-seg-
mented. First segment unarmed, second with a minute setule on its
proximal outer (ventral) surface, a surficial posterior seta finely barbed
along one edge, an inner (dorsal) distal spine with several large spinules
along its distal edge, and the segment produced distally to form a lash
with strong teeth along one edge proximally, grading into fine bilateral
spinulation distally. Maxilliped (fig. 13) 3-segmented. First segment
with a small posterior patch of spinules, second with an inner patch of
spinules and two barbed setae, and third with a small naked seta and
700 Proceedings of the Biological Society of Washington
two terminal barbed spiniform elements, one without a distinct articula-
tion.
Area between maxillipeds and first pair of legs (fig. 14) not pro-
tuberant; a sclerotized line between bases of maxillipeds.
Legs 1-4 (figs. 15-18) with trimerous rami except for endopod of
leg 4 which is 2-segmented. Armature as follows (Roman numerals =
spines, Arabic numerals = setae):
P, protopod 0-1 1-0 exp J-0O i-1 IILI,4
end O-1 0-1 I5
P. protopod 0-1 1-0 exp JI-0O I-1 III,15
end 0-1 0-2 LJII,3
P; protopod 0-1 1-0 exp I-O JI-1 MIII,15
end 0-1 0-2 L,II,2
P, protopod Q0-1 1-0 exp I-0O I[-1 ILI5
end O-I II
Inner seta on coxa of legs 1-3 long and plumose, but in leg 4 short
(17 w) and naked. Outer margin of coxa of leg 1 showing a slight
protrusion. Outer seta on basis short and naked in leg 2, longer and
with lateral hairs in other legs. Inner margin of basis with row of hairs
in legs 1-3, but naked in leg 4. Three middle spines on exopod of leg
1 with spinules along their proximal edges much stronger than in legs
2-4. Endopod of leg 4 (fig. 18) a little shorter than exopod. First
segment 50 « X 40 uw (including spiniform processes), with its inner
distal element a slender naked spine 36 uw long. Second segment 97 uw
(including processes) X 38 u (greatest width), its two unequal terminal
spines 34 w (outer), with a slight distal fringe, and 72 mw (inner),
with a prominent fringe. Both segments with outer margins haired,
and second segment with a row of minute spinules on proximal inner
margin and another row near insertions of terminal spines.
Leg 5 (fig. 19) with rather broad free segment, 122 u long, with its
proximal area expanded inwardly, width here being 55 uw. Segment
bearing two terminal naked setae 91 uw (outer) and 114 w (inner) and
ornamented with small spinules on its outer surface. Seta on body near
base of free segment 66 » and naked; a few small spinules near its
insertion.
Leg 6 probably represented by the two setae near areas of attachment
of each egg sac (see fig. 3).
Color in life in transmitted light opaque to translucid, eye red, ovary
gray, eZg sacs gray.
Male: Body (fig. 20) with prosome a little more slender than in
female. Length 0.90 mm (0.86—0.94 mm) and greatest width 0.35 mm
(0.33-0.36 mm), based on 10 specimens. Ratio of length to width of
prosome 1.79: 1.
Segment of leg 5 (fig. 21) 33 u x 106 uw. No ventral intersegmental
sclerite. Genital segment about as long as wide, 203 wu x 195 uw. Four
a
Copepods Associated with Octocorals 701
postgenital segments 25 wu X64 uw, 22 uw X 64 uw, 17 w X 62 w, and 24 ux
63 u», from anterior to posterior.
Caudal ramus much like that of female, 21 u x 25 uw, with a few
inner proximal hairs on outermost distal seta.
Dorsal surface of prosome smooth; surface of urosome with a few
small hairs as in figure. Ratio of length of prosome to that of urosome
1.62: 1.
Rostral area as in female.
First antenna (fig. 22) resembling that of female, but with two
aesthetes added on second segment and one on fourth segment, so
that formula is 4, 13 +2 aesthetes, 6, 3+ 1 aesthete, 4+ 1 aesthete,
2-+1 aesthete, and 7+ 1 aesthete. Second antenna like that of female,
but with small spinules added on inner surface of first, second, and
fourth segments.
Labrum, mandible, paragnath, first maxilla, and second maxilla like
those of female. Maxilliped (fig. 23) 4-segmented, assuming that
proximal part of claw represents a fourth segment. First segment
unarmed. Second segment with two setae, one slender and naked,
the other (fig. 24) stout, spiniform, obtusely tipped, and spinulose; and
with two rows of moderately long spines. Third segment short and
unarmed. Claw slender, 168 w along its axis including terminal lamella,
with a suggestion of division midway along its length, and bearing two
proximal setae, that on inner surface long with a finely barbed tip, the
other on concave margin short and naked.
Area between maxillipeds and first pair of legs as in female.
Legs 1-4 segmented as in female, with spine and setal formula as in
that sex except for last segment of endopod of leg 1 (fig. 25) which is
1,1,4. This endopod slightly geniculate. No sexual dimorphism in legs
2 or 3. Endopod of leg 4 as in female.
Leg 5 (fig. 21) with slender free segment 42 wu x 11 yp, without a
proximal expansion. Outer surface with a few minute spinules. Two
terminal naked setae 30 w (inner) and 77 uw (outer). Seta on body
near free segment 30 w and naked.
Leg 6 (fig. 21) a posterolateral flap on ventral surface of genital
segment bearing two naked setae 66 wu and 50 wu long.
Spermatophore (fig. 26), attached to female in pairs, elongated,
210 « X 117 uw, not including neck.
Color in life as in female.
Etymology: This species is named for Dr. J. Verseveldt, who has
identified many octocorals which were the hosts for copepods.
Comparison with related species: There are 40 species of Lichomolgus
which, like L. verseveldti, have two claws on the second antenna and the
formula II,1,5 for the third segment of the exopod of leg 4. Thirty-five
of these have a seta (usually feathered) on the first segment of the
endopod of leg 4, thus differing from the new species. In none of these
is the proximal inner expansion (when present) of leg 5 like that in
702 Proceedings of the Biological Society of Washington
L. verseveldti. L. securiger Humes, 1964, an associate of the nudibranch
Doris mabilla Abraham in Madagascar, has a fifth leg somewhat
resembling that of the new species, but differs in having a long
flagellum on the mandible, spinules rather than strong teeth on the
lash of the second maxilla, two extremely unequal setae on the second
segment of the maxilliped of the female, a feathered seta on the first
segment of the endopod of leg 4, relatively short aesthetes on the first
antenna of the male, and the free segment of leg 5 in the male being
about 2: 1 instead of about 4:1 as in the new species.
The remaining five species in the genus (which similarly have two
claws on the second antenna and the formula II,I,5 on the last segment
of the exopod of leg 4) have a spine (rather than a seta) on the first
segment of the endopod of leg 4, as in the new species. These species,
the first five described by Humes and Ho (1968b), associated with
nephtheid octocorals in Madagascar, may be distinguished from L.
verseveldti by the combination of two characters,—the longer claws
on the second antenna in both sexes and the form of leg 5 in the female.
Lichomolgus triquetrus new species
Figures 27-48
Type material: 74299, 2766, and 10 copepodids from Anthelia
gracilis (May), on stems of Cymodocea in 0.5 m, Antsakoabe, Nosy Bé,
Madagascar, collected 12 July 1964. Holotype @, allotype, and 75
paratypes (559 9, 208 4) deposited in the United States National
Museum, and the remaining paratypes in the collection of A. G. Humes.
Female: Body (fig. 27) resembling that of L. verseveldti. Length
1.12 mm (1.01-1.23 mm) and greatest width 0.45 mm (0.42-0.47 mm),
based on 10 specimens. Ratio of length to width of prosome 1.58: 1.
Segment of leg 1 separated from head by a dorsal furrow. Epimeral
areas of segments of legs 1-4 as in figure.
Segment of leg 5 (fig. 28) 64 u x 148 uw. Between this segment and
genital segment a very slight ventral intersegmental sclerite. Genital
segment a little longer than wide, 166 u x 143 yu, in dorsal view broadest
in its anterior third and tapering gradually posteriorly. Areas of attach-
ment of egg sacs situated laterally near middle of segment. Each area
(fig. 29) with two naked unequal setae, the longer one 21 w and
attenuated, the shorter one 10 » with a rather blunt tip. Three post-
genital segments 50 uw < 80 uw, 33 uw X 69 uw, and 39 u x 66 uw, from an-
terior to posterior.
Caudal ramus (fig. 30) quadrate, 30 u x 28 uw in greatest dimensions.
Outer lateral seta 52 « and naked, pedicellate dorsal seta 25 uw and naked,
outermost distal seta 68 « with only a few proximal outer spinules,
innermost distal seta 115 uw with inner spinules, and the two long median
terminal setae 240 uw (outer) and 385 uw (inner), both naked and both
inserted between unornamented dorsal and ventral flaps.
ee
Copepods Associated with Octocorals 703
Fics. 31-39. Lichomolgus triquetrus, new species, female: 31,
second antenna, posterior (outer) (F); 32, mandible, posterior (C);
33, paragnath, ventral (C); 34, first maxilla, anterior (C); 35, second
maxilla, posterior (C); 36, maxilliped, posterior (F); 37, leg 1 and
intercoxal plate, anterior (E); 38, leg 2, anterior (E); 39, third segment
of endopod of leg 3, anterior (E).
Fics. 40-48. Lichomolgus triquetrus, new species, female: 40, leg 4
and intercoxal plate, anterior (E); 41, leg 5, dorso-inner (C). Male:
42, body, dorsal (G); 43, urosome, dorsal (D); 44, maxilliped, outer
(F); 45, third segment of endopod of leg 1, anterior (C); 46, third
segment of endopod of leg 2, anterior (C); 47, leg 5, dorsal (1); 48,
spermatophore, as seen inside genital segment of male, dorsal (D).
Copepods Associated with Octocorals 705
Dorsal surface of prosome and urosome with a few hairs. Ratio of
length of prosome to that of urosome 1.94: 1.
Egg sac (fig. 27) elongated, 473 uw x 204 uw, reaching well beyond
caudal rami and containing about 22-26 moderately large eggs, each
approximately 68 u in diameter.
Rostral area as in L. verseveldti.
First antenna also like that of L. verseveldti, with same formula for
armature. Lengths of segments: 23 « (57° along anterior margin),
112 pw, 22 uw, 49 uw, 46 pw, 35 uw, and 22 uw respectively. All setae naked.
Second antenna (fig. 31) 4-segmented, last segment not greatly elon-
gated, 58 uw along its outer edge, 40 uw along its inner edge, 23 w wide,
bearing distally six small hyaline elements and a single relatively short
claw 31 wu along its axis. All setae naked.
Labrum as in L. verseveldti. Mandible (fig. 32) with fewer spinules
on protrusion of convex margin of basal part distal to constriction and
with flagellum longer than in L. verseveldti. Paragnath (fig. 33) a
small hairy lobe. First maxilla (fig. 34) with four elements as in L.
verseveldti. Second maxilla (fig. 35) in general like that of L.
verseveldti, but with inner distal spine of second segment with three
rows of slender spinules. Maxilliped (fig. 36) lacking patches of spinules
on first and second segments and the two setae on second segment
naked and smaller than in L. verseveldti.
Area between maxillipeds and first pair of legs as in L. verseveldti,
but slightly protuberant; a sclerotized line between bases of maxillipeds.
Legs 1-4 (figs. 37-40) segmented as in L. verseveldti, and with same
spine and setal formula except for leg 4 where exopod is I-O; I-1; II,1,5
and endopod is 0-1; II. Inner seta on coxa of leg 4 short (15 ») and
naked, instead of long and plumose as in legs 1-3. Outer seta on basis
short in leg 2, as in L. verseveldti, and inner margin of basis naked in
leg 4 as in that species. Endopod of leg 4 (fig. 40) shorter than exopod.
First segment 40 u X 39 wu (including spiniform processes), with inner
distal element a seta 38 u long which is lightly feathered in some speci-
mens and apparently naked in others. Second segment 70 uw X 33 wu in
greatest dimensions including processes, its two unequal terminal fringed
spines 35 uw (outer) and 54 uw (inner). Both segments with outer margins
haired and second segment with a row of minute spinules near insertions
of terminal spines.
Leg 5 (fig. 41) with rather triangular free segment, 63 » X 39 wu in
greatest dimensions, bearing two relatively short terminal naked setae
29 » (outer) and 31 uw (inner). Small spinules on outer surface of
segment. Seta on body near base of segment 44 w and naked, a few
small spinules near its insertion.
Leg 6 probably represented by the two setae near areas of attachment
of each egg sac (see fig. 29).
Color in life in transmitted light translucid except for fine reddish
brown speckling, eye red, ovary gray, egg sacs greenish gray.
706 Proceedings of the Biological Society of Washington
Male: Body (fig. 42) similar in general form to that of female, but
prosome a little more slender. Length 0.89 mm (0.85-0.96 mm) and
greatest width 0.29 mm (0.28-0.30 mm), based on 10 specimens.
Ratio of length to width of prosome 1.85: 1.
Segment of leg 5 (fig. 43) 41 u x 98 uw. No ventral intersegmental
sclerite. Genital segment about as long as wide, 205 » x 195 uw. Four
postgenital segments 31 uw X 58 uw, 25 w X 55 pw, 23 uw X 52 wu, and 25 ux
50 uw, from anterior to posterior.
Caudal ramus similar to that of female, but aes 23 w X 22 mw,
Dorsal surface of prosome and urosome with a few hairs. Ratio of
length of prosome to that of urosome 1.58: 1.
Rostral area as in female.
First antenna as in L. verseveldti, with three aesthetes added as in that
species. Second antenna as in female, without sexual dimorphism.
Labrum, mandible, paragnath, first maxilla, and second maxilla as in
female. Maxilliped (fig. 44) with two setae on second segment at-
tenuated, one of them with small spinules along one edge, the other
naked. Claw 168 uw along its axis including terminal lamella, with its
outer proximal surface faintly striated. Otherwise maxilliped similar to
that of L. verseveldti.
Area between maxillipeds and first pair of legs as in female.
Legs 1-4 segmented as in female, with spine and seta] formula as in
that sex except for last segment of endopod of leg 1 (fig. 45) which
is I,1,4. (One male showed the formula J,5 on both first legs, but the
usual condition is I,I,4.) Last segment of endopod of leg 2 (fig. 46)
with same formula as in female, but two terminal spines modified.
Legs 3 and 4 as in female.
Leg 5 (fig. 47) with an elongated and somewhat rectangular free
segment, 33 » X 10 uw with nearly parallel sides. Two terminal elements
very unequal, the outer one a naked seta 31 uw long, the inner one a
delicately barbed spine 12 ». (In the same male mentioned above which
showed the abnormal armature of the last segment of the endopod of
leg 1, the fifth leg had the unusual condition of two nearly equal naked
setae, as in the female.) Outer surface of segment with small spinules.
Seta on body near free segment 30 mw and naked.
Leg 6 (see fig. 43) a posterolateral flap on ventral surface of genital
segment bearing two naked setae about 33 wu long.
Spermatophore (fig. 48), seen only inside genital segment of male,
elongated with an unusually thick wall.
Color in life as in female.
Etymology: The specific name triquetrus, from Latin = three-cornered,
triangular, refers to the rather triangular form of the free segment of
leg 5 in the female.
Comparison with related species: In six species of Lichomolgus the
second antenna bears terminally a single claw and a few small hyaline
elements and the formula for the last segment of the exopod of leg 4
‘tl
Copepods Associated with Octocorals 707
is III,I,5, as in the new species. All of these differ from L. triquetrus,
however, in significant respects. In L. dentipes Thompson and A. Scott,
1903, there is a large toothlike process on leg 5 of the female. (This
species has been redescribed by Humes and Ho, 1968c.) In L.
protentus Humes and Frost, 1964, the caudal ramus of the female is
twice as long as wide, and the formula for the endopod of leg 4 is
0-1; 1,1. In L. spinipes (Sewell, 1949) leg 5 of the female is long and
curved, and the formula for the endopod:of leg 4 is 0-1; 1,I. In
L. squamiger Humes and Frost, 1964, the areas of attachment of the
egg sacs are dorsolateral, the endopod of leg 4 is a little longer than the
exopod, and leg 5 in the female is elongated with broad scalelike
spines. In the two remaining species, L. cristatus and L. adelphus,
both described by Humes and Ho (1968c), leg 5 in the female is
elongated, at least five times longer than wide.
In L. elegans Thompson and A. Scott, 1903, the second antenna of the
female has a single claw as in the new species. The armature of the
exopod of leg 4 is undescribed. (The male is unknown.) This Ceylonese
species differs from L. triquetrus in the genital segment having “a wedge-
shaped notch near the centre on each side,” in the second segment of
the first antenna being relatively longer, and in the inner distal seta
on the first segment of the endopod of leg 4 being longer and more
conspicuously feathered.
Lichomoilgus glabripes new species
Figures 49-68
Type material: 5399, 3766, and 33 copepodids from Xenia
umbellata Lamarck, in 1 m, Andilana, Nosy Bé, Madagascar, collected
9 August 1963. Holotype 9, allotype and 69 paratypes (402 9, 294 6)
deposited in the United States National Museum, and the remaining
paratypes in the collection of A. G. Humes.
Female: Body (fig. 49) with a moderately slender prosome. Length
1.33 mm (1.26—-1.42 mm) and greatest width 0.53 mm (0.51-0.55 mm),
based on 10 specimens. Ratio of length to width of prosome 1.46: 1.
Segment of leg 1 separated from head by a dorsal furrow. Epimeral
areas of segments of legs 14 as in figure.
Segment of leg 5 (fig. 50) 99 u x 200 uw. Between this segment and
genital segment a short ventral intersegmental sclerite. Genital segment
longer than wide, 200 » xX 151 yu in greatest dimensions, its posterior
part in dorsal view somewhat constricted (99 « wide). Areas of attach-
ment of egg sacs located dorsolaterally just anterior to middle of seg-
ment. Each area (fig. 51) with two short naked spiniform setae 7 p»
and 4.5 » long. Three postgenital segments 75 mu X 87 yw, 62 uw xX 78 uy,
and 70 u X 78 yu, from anterior to posterior.
Caudal ramus (fig. 52) elongated, 110 uw X 32 yu, or 3.4 times longer
than wide. Outer lateral seta 112 » and naked, pedicellate dorsal seta
33 w and lightly feathered, outermost distal seta 120 u and naked, inner-
708 Proceedings of the Biological Society of Washington
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Fics. 49-57. Lichomolgus glabripes, new species, female: 49, body,
dorsal (A); 50, urosome, dorsal (B); 51, area of attachment of egg sac,
dorsal (1); 52, caudal ramus, dorsal (F); 53, rostral area, ventral (E);
54, second antenna, posterior (outer) (E); 55, labrum, ventral (F);
56, mandible, posterior (F); 57, second maxilla, posterior (F).
Copepods Associated with Octocorals 709
Fics. 58-64. Lichomolgus glabripes, new species, female: 58,
maxilliped, posterior (F); 59, leg 1 and intercoxal plate, anterior (E);
60, leg 2, anterior (D); 61, third segment of endopod of leg 3, anterior
(E); 62, leg 4 and intercoxal plate, anterior (D); 63, leg 5, dorsal
(E). Male: 64, body, dorsal (G).
710 Proceedings of the Biological Society of Washington
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Fics. 65-70. Lichomolgus glabripes, new species, male: 65, urosome,
dorsal (D); 66, second antenna, posterior (outer) (F); 67, maxilliped,
inner (E); 68, third segment of endopod of leg 1, anterior (C).
Lichomolgus longispinifer, new species, female: 69, body, dorsal (G);
70, urosome, dorsal (E).
Copepods Associated with Octocorals 711
most distal seta 130 « with prominent inner proximal hairs, and the
two long median terminal setae 200 « (outer) and 263 » (inner), both
naked and inserted between an unornamented dorsal flap and a ventral
flap with a marginal row of minute spinules.
Dorsal surface of prosome and urosome with very few hairs. Ratio
of length of prosome to that of urosome 1.42 : 1.
Egg sac (fig. 49) elongated oval, 396 u x 220 uw, reaching just beyond
anal segment and containing about 15-17 eggs, each approximately 105 yu
in diameter.
Rostral area (fig. 53) with broadly rounded posteroventral margin.
First antenna 340 w long, with segmentation and armature as in
L. verseveldti and L. triquetrus. Lengths of segments: 25 u (55 u along
anterior edge), 105 wu, 28 uw, 46 uw, 47 w, 35 uw, and 24 yu respectively. All
setae naked. Second antenna (fig. 54) 4-segmented and slender, last
segment 61 u along its outer edge, 44 uw along its inner edge, 18 » wide,
bearing distally six small hyaline elements and a single short claw 40 yu
along its axis. All setae naked.
Labrum (fig. 55) with two broad and outwardly angular lobes.
Mandible (fig. 56), paragnath, first maxilla, second maxilla (fig. 57),
and maxilliped (fig. 58) resembling in major respects those of L.
triquetrus.
‘Area between maxillipeds and first pair of legs as in L. verseveldti;
a sclerotized line between bases of maxillipeds.
Legs 1-4 (figs. 59-62) segmented as in the two previous species,
with spine and setal formula as in L. triquetrus (exopod of leg 4 being
I-0; I-1; IILI,5). Coxa of leg 1 with a sclerotized boss on outer
posterior surface. Inner seta on coxa of leg 4 short (20 uw) and naked,
instead of long and plumose as in legs 1-3. Inner margin of basis of leg
4 naked, but haired in legs 1-3. Endopod of leg 4 shorter than exopod.
First segment 51 u xX 43 w (including spiniform processes), with inner
distal feathered seta 66 u long. Second segment 98 u x 38 u in greatest
dimensions (including processes), its two unequal terminal fringed
spines 37 u (outer) and 64 uw (inner). Both segments with outer margins
haired and second segment with a terminal row of minute spinules.
Leg 5 (fig. 63) with a long bowed free segment 135 » X 31 yw in
greatest dimensions, without fine ornamentation. Two terminal naked
setae 44 uw (outer) and 60 uw (inner). Seta on body near free segment
58 uw and naked. (In ovigerous females leg 5 is held erect so that its
curvature fits around the ends of the egg sacs.)
Leg 6 probably represented by the two setae near areas of attachment
of each egg sac (see fig. 51).
Color in life in transmitted light opaque to transparent, eye red, egg
sacs gray.
Male: Body (fig. 64) resembling that of female. Length 1.04 mm
(1.00-1.07 mm) and greatest width 0.35 mm (0.34-0.35 mm), based on
10 specimens. Ratio of length to width of prosome 1.67: 1.
712 Proceedings of the Biological Society of Washington
Segment of leg 5 (fig. 65) 47 « X 107 uw. No ventral intersegmental
sclerite. Genital segment longer than wide, 205 » x 169 uw. Four post-
genital segments 36 uw X 60 uw, 39 u X 55 uw, 31 uw x 52 uw, and 43 u X 53 4,
from anterior to posterior.
Caudal ramus similar to that of female, but relatively shorter, 73 u X
23 w, or 3.17: 1.
Dorsal surface of prosome and urosome with only a few hairs. Ratio
of length of prosome to that of urosome 1.33 : 1.
Rostral area as in female.
First antenna as in female, but with three aesthetes added as in the
two previous species. Second antenna (fig. 66) like that of female, but
with scalelike spines added on first two segments, those on first segment
arranged in an arc.
Labrum, mandible, paragnath, first maxilla, and second maxilla as in
female. Maxilliped (fig. 67) slender. Second segment bearing a row
of spines and two moderately long naked setae. Claw not greatly re-
curved, 189 uw along its axis including terminal lamella.
Area between maxillipeds and first pair of legs as in female.
Legs 1-4 segmented as in female, with spine and setal formula as in
that sex except for last segment of endopod of leg 1 (fig. 68) which
is 11,4. Legs 2-4 as in female, without sexual dimorphism. .
Leg 5 (fig. 65) with an elongated straight unornamented free seg-
ment, 44 u X 11 yu, its two naked terminal setae 30 » and 33 uw. Naked
seta on body near free segment 40 u.
Leg 6 (fig. 65) a posterolateral flap on ventral surface of genital seg-
ment bearing two naked setae 40 wu and 36 uw long.
Spermatophore not observed.
Color in life as in female.
Etymology: The specific name glabripes, from Latin glaber = without
hairs, bald, and pes = foot, alludes to the absence of fine ornamentation
on leg 5.
Comparison with related species: In seven species of Lichomolgus
the second antenna bears terminally a single claw and a few small
hyaline elements and the formula for the last segment of the exopod of
leg 4 is III,I,5, as in L. glabripes. Each of these species may, however,
be readily differentiated from L. glabripes. In L. dentipes Thompson
and A. Scott, 1903, the free segment of leg 5 in the female has a large
toothlike process. In L. protentus Humes and Frost, 1964, the armature
of the endopod of leg 4 is 0-1; 1,1. In L. spinipes (Sewell, 1949) the
caudal ramus is wider than long, and the fourth endopod is 0-1; 1,1. In
L. squamiger Humes and Frost, 1964, the caudal ramus is 1 : 1, and leg
5 of the female bears scales. In L. cristatus Humes and Ho, 1968c, the
genital segment of the female is wider than long, the mandible has a
very short flagellum, and the lash of the second maxilla has a crest of
long spinules. In L. adelphus Humes and Ho, 1968c, the caudal
ramus is about 1:1, the genital segment of the female is wider than
Copepods Associated with Octocorals 718
long, and leg 5 of the female has scalelike spines. In L. triquetrus,
described above, the caudal ramus is about 1:1, and the free segment
of leg 5 is triangular and ornamented with small spines.
L. elegans Thompson and A. Scott, 1903, of which only the female
is known, has a single claw on the second antenna. The armature of
the exopod of leg 4 is undescribed. However, this Ceylonese species
differs from L. glabripes in having the caudal ramus about as broad as
long, and in the presence of “a wedge-shaped notch near the centre on
each side” of the genital segment.
Lichomolgus longispinifer new species
Figures 69-89
Type material: 8399, 2844, and several copepodids from one
colony of Siphonogorgia pendula Studer, in 10-12 m, south of Andjiabe,
Nosy Komba, near Nosy Bé, Madagascar, collected 30 July 1964.
Holotype 9, allotype, and 80 paratypes (6092 9, 204 6) deposited in
the United States National Museum, and the remaining paratypes in the
collection of A. G. Humes.
Other specimens: 399 9, 474 4, and several copepodids from one
colony of Siphonogorgia pendula, in 20 m, Banc de Dzamandzar, near
Nosy Bé, 16 September 1964.
Female: Body (fig. 69) with prosome not much broadened; urosome
relatively short. Length 0.88 mm (0.84—0.90 mm) and greatest width
0.43 mm (0.40-0.44 mm), based on 10 specimens. Ratio of length to
width of prosome 1.60: 1. Segment of leg 1 well separated from head
dorsally. Epimeral areas of segments of legs 1 and 2 rather angular
posteriorly, those of segments of legs 3 and 4 rounded.
Segment of leg 5 (fig. 70) 47 u X 101 yu. Between this segment and
genital segment no ventral intersegmental sclerite. Genital segment
about as long as wide, 104 u x 101 yu, in dorsal view gradually broadened
from anterior to posterior, then rather abruptly constricted in its posterior
fourth. Areas of attachment of egg sacs dorsal in position near midregion
of segment. Each area (fig. 71) bearing two naked setae 11 » and 9 uw
long and two pointed processes. Three postgenital segments 26 u x 56 uy,
21 w X 52 w, and 28 » x 51 u, from anterior to posterior.
Caudal ramus (fig. 72) a little wider than long, 20 uw Xx 22 w in
greatest dimensions. Outer lateral seta 77 mu and naked, pedicellate
dorsal seta 66 uw and lightly feathered, outermost distal seta 105 » with
lateral hairs proximally, innermost distal seta 143 » and similarly haired,
and the two long median terminal setae 440 » (outer) and 495 uw (inner),
both strongly haired along their midregions and both inserted between
an unornamented dorsal flap and a ventral flap with a marginal row of
minute spinules.
Dorsal surface of prosome and urosome with very few hairs. Ratio
of length of prosome to that of urosome 3.33: 1.
Egg sac (fig. 69) slender and elongated, 363 u x 101 u, reaching far
714 Proceedings of the Biological Society of Washington
Dy
: aN
Fics. 71-79. Lichomolgus longispinifer, new species, female: 71,
area of attachment of egg sac, dorsal (I); 72, caudal ramus, dorsal (1);
73, first antenna, dorsal (D); 74, second antenna, anterior (inner)
(F); 75, mandible, posterior (C); 76, first maxilla, posterior (C); 77,
second maxilla, posterior (C); 78, maxilliped, posterior (C); 79, area
between maxillipeds and first pair of legs, ventral (E).
Copepods Associated with Octocorals 715
RR Ni
CR
SN
Fics. 80-84. Lichomolgus longispinifer, new species, female: 80, leg
1 and intercoxal plate, anterior (F); 81, leg 2, anterior (F); 82, leg 3,
anterior (F); 83, leg 4, anterior (F); 84, leg 5, dorsal (F).
beyond caudal rami and containing many eggs, each about 47 w in
diameter.
Rostral area similar to that in L. verseveldti and L. triquetrus.
First antenna (fig. 73) long and slender, 437 uw in length, with seg-
mentation and armature as in the three previous species. Lengths of
716 Proceedings of the Biological Society of Washington
Fics. 85-90. Lichomolgus longispinifer, new species, male: 85,
body, dorsal (G); 86, urosome, dorsal (E); 87, maxilliped, postero-
inner (C); 88, endopod of leg 1, anterior (C); 89, third segment of
endopod of leg 2, anterior (C). Lichomolgus hians, new species, female:
90, body, dorsal (G).
Copepods Associated with Octocorals 717
segments: 42 uw (65 uw along anterior edge), 164 u, 27 uw, 68 w, 55 mw, 37 p,
and 21 w respectively. All setae naked. Second antenna (fig. 74) 4-
segmented, last segment 55 uw along its outer edge, 28 u along its inner
edge, 23 « wide, bearing distally five short hyaline elements and two
short terminal claws, one stout and 39 wu along its axis, the other more
slender and 44 uw. All setae naked.
Labrum resembling that of L. verseveldti. Mandible (fig. 75) with
basal region strongly constricted, the scalelike protrusion on convex side
distal to constriction pointed and ornamented with a row of minute
spinules. Paragnath as in L. verseveldti. First maxilla (fig. 76) with
four setae. Second maxilla (fig. 77) similar to that in L. triquetrus
and L. glabripes, but with slender spines instead of strong teeth along
terminal lash. Maxilliped (fig. 78) resembling that of L. verseveldti
but differing in details of ornamentation.
Area between maxillipeds and first pair of legs (fig. 79) slightly
protuberant; a sclerotized line between bases of maxillipeds.
Legs 1-4 (figs. 80-83) segmented as in all previous species and with
spine and setal formula as in L. verseveldti. Inner seta on coxa of leg
4 short (7 w) and naked. Inner margin of basis of leg 4 naked. First
segment of exopod of leg 1 with outer spine unusually long, 44 u, a
little more than twice length of spine on second segment (21 sz).
Spinulose fringe on this long spine narrower and more delicate than on
more distal spines. Spine on first segment of exopod of leg 2 only
slightly longer than others; proximal spine on third segment of this
exopod slightly shorter than adjacent spines; lengths of the five outer
spines on exopod being 25 uw, 20 uw, 18 uw, 20 uw, and 18 yp, from proximal
to distal. Endopod of leg 4 shorter than exopod. First segment 37 u x
17 uw (including spiniform processes), with a short naked inner distal
spinelike element 18 » long. Second segment 66 u xX 17 wu in greatest
dimensions (including processes), its two unequal terminal spines 22 u
(outer) and weakly barbed distally, and 50 w (inner) with an outer
spinulose fringe and an inner narrow smooth lamella. Both segments with
outer margins haired and second segment with a terminal row of minute
spinules.
Leg 5 (fig. 84) resembling in general form that of L. cuneipes Humes
and Ho, 1968b. Free segment elongated, 106 uw in greatest length and
21 mw wide near middle. Proximal inner expansion (smaller than in
L. cuneipes) in the form of a bent thumb. Two terminal naked setae
39 w (outer) and 56 uw (inner). Outer surface of segment with short
spines. Seta on body near free segment 44 uw and lightly feathered.
Leg 6 probably represented by the two setae near areas of attachment
of each egg sac (see fig. 71).
Color in life in transmitted light faintly rose or lavender, sometimes
translucid, eye red, egg sacs light gray.
Male: Body (fig. 85) with prosome more slender than in female.
Length 0.67 mm (0.64-0.70 mm) and greatest width 0.24 mm (0.23—
718 Proceedings of the Biological Society of Washington
0.28 mm), based on 10 specimens. Ratio of length to width of prosome
176i
Segment of leg 5 (fig. 86) 34 u x59 uw. No ventral intersegmental
sclerite. Genital segment 148 » x 130 yu, only a little longer than wide.
Four postgenital segments 15 uw x 44 w, 15 w x 44 uw, 11 w xX 42 pw, and
17 uw X 44 uw, from anterior to posterior.
Caudal ramus similar to that of female, 15.5 u x 19 u.
Dorsal surface of prosome and urosome with very few hairs. Ratio
of length of prosome to that of urosome 1.89 : 1.
Rostral area as in female. First antenna like that of female, but with
three aesthetes added as in the three previous species. Second antenna
similar to that of female, but bearing a few short spinules along inner
surface of second and fourth segments.
Labrum, mandible, paragnath, first maxilla, and second maxilla as in
female. Maxilliped (fig. 87) resembling in general form that of L.
cuneipes and L. gentilis Humes and Ho, 1968b. Claw 142 w along its
axis including terminal lamella.
Area between maxillipeds and first pair of legs as in female.
Legs 1-4 segmented as in female, with same spine and setal formula
except for last segment of endopod of leg 1 (fig. 88) which is I,I,4.
Leg 2 showing slight sexual dimorphism, with outer terminal spiniform —
process on third segment of endopod larger than in female and turned
outwardly (fig. 89). Leys 3 and 4 as in female, without sexual dimor-
phism.
Leg 5 (fig. 86) with free segment straight and elongated, 32 u x 6 un,
without ornamentation. Two terminal naked setae 29 uw and 14 uw, and
seta on body near free segment 20 uz.
Leg 6 (fig. 86) a posterolateral flap on ventral surface of genital
segment bearing two naked setae 39 uw and 30 uz.
Spermatophore not observed.
Color in life in transmitted light translucid, eye red.
Etymology: The specific name longispinifer, from Latin = bearing a
long spine, refers to the unusually long spine on the first segment of the
exopod of leg 1.
Comparison with related species: The new species appears to be
unique in the genus in having an unusually long outer spine on the first
segment of the exopod of leg 1. It may be further separated from all
but two species of Lichomolgus on the basis of a combination of the
following four characters: the second antenna with two short claws, the
formula for the last segment of the exopod of leg 4 being II,I,5, the
endopod of leg 4 being 0-I; II, and the caudal ramus being nearly
quadrate, a little wider than long. The two species which share these
features with L. longispinifer are L. cuneipes Humes and Ho, 1968b, and
L. verseveldti, described above. In L. verseveldti, however, the form
of the fifth leg and genital segment in the female is very different and
the lash of the second maxilla bears stout teeth rather than slender
Copepods Associated with Octocorals 719
spines. The new species seems closely related to L. cuneipes (associated
with the octocoral Stereonephthya acaulis Verseveldt in Madagascar).
In L. cuneipes, however, all ramal setae are naked, the teeth on the lash
of the second maxilla are partly slender and partly stout, the genital
segment of the female has a different form, and the fifth leg of the
male bears fine ornamentation.
Lichomolgus hians new species
Figures 90-108
Type material: 2389 9, 3764, and several copepodids from one
colony of Siphonogorgia pendula Studer, in 10-12 m, south of Andjiabe,
on the southern shore of Nosy Komba, near Nosy Bé, Madagascar,
collected 30 July 1964. Holotype ¢@, allotype, and 225 paratypes
(2002 2, 254 4) deposited in the United States National Museum,
and the remaining paratypes in the collection of A. G. Humes.
Other specimens (also from Siphonogorgia pendula): 789 2, 1126 é,
and several copepodids from one colony, in 20 m, Banc de Dzamandzar,
near Nosy Bé, 16 September 1964.
Female: Body (fig. 90) with moderately broadened prosome. Length
0.93 mm (0.86-1.03 mm) and greatest width 0.49 mm (0.46-0.51
mm), based on 10 specimens. Ratio of length to width of prosome
1.37: 1. Segment of leg 1 separated incompletely from head. Epimeral
areas of segments of legs 1-4 as in figure.
Segment of leg 5 (fig. 91) 65 w x 131 uw. Between this segment and
genital segment a slight ventral intersegmental sclerite. Genital segment
112 » X 109 uw, about as long as wide, and resembling that of L.
varirostratus Humes and Ho, 1968b. Areas of attachment of egg sacs
situated dorsally near middle of segment. Each area bearing two naked
setae 10 uw and 11 uw long and a prominent unguiform process. Three
postgenital segments 29 u x 67 nw, 24 uw X 60 uw, and 26 u x 57 wu, from
anterior to posterior.
Caudal ramus (fig. 91) quadrate, 23 « x 24 w in greatest dimensions.
Outer lateral seta 66 « long and naked, pedicellate dorsal seta 50 « and
lightly feathered, outermost distal seta 110 » and naked, innermost
distal seta 200 » with hairs along both sides of its proximal half, and
the two long median terminal setae 308 uw (outer) and 440 w (inner),
both with strong lateral spinules (stronger than in L. longispinifer)
except in distal third, and both inserted between an unornamented dorsal
flap and a ventral flap with a marginal row of minute spinules. A
minute spinule on proximal outer margin of ramus.
Dorsal surface of prosome and urosome with very few hairs. A small
crescentic sclerotization in tergum of segment of leg 4. Ratio of length
of prosome to that of urosome 3.23: 1.
Egg sac seen complete in only one female, where it measured 495 » x
165 uw, reaching almost to ends of ramal setae, and containing many
720 Proceedings of the Biological Society of Washington
SS
ERRSY
LD
Pop
Va
IX.
(WU
y)
Ne
Fics. 91-98. Lichomolgus hians, new species, female: 91, urosome,
dorsal (EF); 92, rostral area, ventral (D); 93, first antenna, dorsal (D);
94, second antenna, anterior (inner) (E.); 95, mandible, posterior (C);
96, first maxilla, posterior (C); 97, maxilliped, postero-inner (C); 98,
third segment of endopod of leg 2, anterior (F).
Copepods Associated with Octocorals Al
Fics. 99-105. Lichomolgus hians, new species, female: 99, third
segment of endopod of leg 3, anterior (F); 100, leg 4, anterior (E);
Male: 101, body, dorsal (G); 102, urosome, dorsal (E); 103, second
antenna, anterior (inner) (F); 104, maxilliped, outer (F); 105, endopod
of leg 1, anterior (C).
722 Proceedings of the Biological Society of Washington
Fics. 106-113. Lichomolgus hians, new species, male: 106, third
segment of endopod of leg 2, anterior (C); 107, endopod of leg 4,
anterior (C); 108, leg 5, dorsal (1). Lichomolgus telestophilus, new
species, female: 109, body, dorsal (G); 110, urosome, dorsal (D); 111,
area of attachment of egg sac, dorsal (1); 112, rostral area, ventral
(D); 113, second antenna, anterior (inner) (E).
Copepods Associated with Octocorals 723
eggs about 52 » in diameter. Egg sacs in other ovigerous females in-
complete, as in figure 90.
Rostral area (fig. 92) raised ventrally, with a well defined postero-
ventral border.
First antenna (fig. 93) 7-segmented, 441 u long, and slender. Lengths
of segments: 40 » (69 u along anterior margin), 133 uw, 31 », 68 wu, 62 pn,
48 uw, and 30 w respectively. Formula for armature as in the four
previous species. All setae naked.
Second antenna (fig. 94) 4-segmented, last segment 86 wu along its
outer edge, 50 uw along its inner edge, 24 uw wide, bearing distally five
small hyaline elements and two moderately long unequal claws, the
stouter one 65 uw along its axis, the more slender one 78 uw. Third seg-
ment with a spiniform seta and two longer slender setae. Groups of
small spinules on first and second segments. All setae naked.
Labrum and paragnath as in L. verseveldti. Mandible (fig. 95) re-
sembling that of L. longispinifer. First maxilla (fig. 96) slender with
four elements as in the four previous species. Second maxilla as in L.
varirostratus. Maxilliped (fig. 97) resembling in general form that of L.
longispinifer, but the two terminal elements equal and widely divergent,
suggesting open blades of scissors.
Area between maxillipeds and first pair of legs as in L. verseveldti.
Legs 1-4 segmented as in the four previous species, and spine and
setal formula as in L. verseveldti and L. longispinifer. Inner seta on
coxa of leg 4 short (7 «) and naked. Inner margin of basis of leg 4
naked. Rami of leg 1 as in L. varirostratus. Rami of legs 2 and 3 also
similar to those in L. varirostratus, but last segment of endopod slightly
different (figs. 98, 99). Endopod of leg 4 (fig. 100) shorter than
exopod. First segment 43 u X 32 w (including processes), with its
distal inner finely barbed spine 44 uw long. Second segment 78 » long
(including spiniform processes), its greatest width 32 w and least width
17.5 uw; its two unequal terminal spines 31 » (outer) and 57 uw (inner),
the fringe on the inner spine more coarsely spinulose on its outer than
on its inner margin. Both segments with outer margins haired, second
segment with a few weak hairs on proximal half of inner margin and
with a terminal row of minute spinules.
Leg 5 (fig. 91) suggesting that of L. varirostratus, but differing in
details of form and ornamentation. Free segment elongated, 117 u x 26 uw
in greatest dimensions. Outer surface of segment with many scalelike
spines. Two terminal naked setae 41 uw (outer) and 69 yu (inner). Seta
on body near free segment about 50 uw and naked.
Leg 6 probably represented by the two setae near areas of attachment
of each egg sac (see fig. 91).
Color in life in transmitted light translucid or slightly opaque, eye
red, egg sacs light gray.
Male: Body (fig. 101) with prosome more slender than in female.
Length 0.73 mm (0.70-0.78 mm) and greatest width 0.28 mm (0.27—
724 Proceedings of the Biological Society of Washington
0.29 mm), based on 10 specimens. Ratio of length to width of prosome
1.70: 1.
Segment of leg 1 (fig. 102) 39 « x 75 uw. No ventral intersegmental
sclerite. Genital segment 151 » x 133 yu, only a little longer than wide,
with its lateral margins sometimes slightly irregular. Four postgenital
segments 25 uw X 48 uw, 18 uw X 46 uw, 11 w xX 45 w, and 19 uw x 46 pn, from
anterior to posterior.
Caudal ramus similar to that of female, 20 « x 20 u in greatest dimen-
sions.
Dorsal surface of prosome and urosome with very few hairs. Ratio
of length of prosome to that of urosome 1.90: 1.
Rostral area as in female. First antenna similar to that of female, but
with three aesthetes added as in the four previous species. Second
antenna (fig. 103) resembling that of female, but last segment relatively
more elongated, its outer margin 82 uw, its inner margin 56 uw, and its
width 17 yw, and both margins with minute spinules. Second segment
with small spines along inner surface.
Labrum, mandible, paragnath, first maxilla, and second maxilla as in
female. Maxilliped (fig. 104) slender. Second segment with two naked
setae and two rows of spinules. Claw 159 w along its axis (including
terminal lamella), only slightly recurved, its proximal inner surface ~
faintly striated.
Area between maxillipeds and first pair of legs as in female.
Legs 1-4 segmented as in female, with same spine and setal formula
except for last segment of endopod of leg 1 (fig. 105) which is I,I,4.
Last segment of endopod of leg 2 (fig. 106) showing slight sexual
dimorphism, with outer terminal process larger than in female, and
bearing minute spinules. Leg 3 entirely as in female. Endopod of leg
4 (fig. 107) with second segment relatively more slender than in female.
First segment 31 » X 21 uw including processes, its spine 32 u; second
segment 60 u long, 15.5 u in greatest width, and 10 wu in least width, its
two terminal spines 17 » (outer) and 47 wu (inner).
Leg 5 (fig. 108) with free segment 35 u xX 8 uw, bearing two terminal
setae, outer 34 w and naked, inner 16 w and finely barbed. Surface of
segment with a few small spines. Seta on body near free segment 29 u
and lightly feathered.
Leg 6 (fig. 102) a posterolateral flap on ventral surface of genital
segment bearing two slender setae 31 « and 40 u long.
Spermatophore not observed.
Color in life as in female.
Etymology: The specific name hians, from the Latin word hio = to
stand open or gape, alludes to the two divergent terminal elements on
the maxilliped of the female.
Comparison with related species: L. hians may be separated from all
but seven species of Lichomolgus on the basis of a combination of four
characters: the caudal ramus being about as long as wide, the second
Copepods Associated with Octocorals Hs,
antenna having two terminal claws, the formula for the last segment
of the exopod of leg 4 being II,I,5, and the formula for the endopod of
leg 4 being O-I; II. Of the remaining species L. varirostratus Humes
and Ho, 1968b, L. exilipes Humes and Ho, 1968b, and L. gentilis Humes
and Ho, 1968b, have the two claws on the second antenna about as
long as or longer than the greatest length of the fourth segment (instead
of distinctly shorter than this segment as in the new species), and the
outer spine on the last segment of the endopod of leg 1 in the males
of these species is differently formed. In L. fissisetiger Humes and Ho,
1968b, L. cuneipes Humes and Ho, 1968b, L. verseveldti, and L.
longispinifer (both described above) the fifth leg of the female has a
distinct proximal inner expansion.
Although in several respects L. hians resembles rather closely L.
varirostratus, particularly in the form of the genital segment in the fe-
male, the mandible, the second maxilla, the maxilliped in the female,
and legs 1-4, there are many differences between the two species. The
distinctions include not only the two features mentioned above, but also
(in L. varirostratus) the sexual dimorphism in the rostral area, the exact
nature of leg 5 in both sexes, and the two unornamented long terminal
setae on the caudal ramus.
Lichomolgus telestophilus new species
Figures 109-127
Type material: 139 9, 224 6, and 4 copepodids from one colony of
Telesto arborea Wright and Studer, in 4 m, northeastern coast of Antany
Mora, Isles Radama, Madagascar, 14°06’10”S, 47°45’10”E, collected 30
September 1964. Holotype °, allotype and 26 paratypes (99 9,174 4)
deposited in the United States National Museum, and the remaining
paratypes in the collection of A. G. Humes.
Female: Body (fig. 109) with moderately slender prosome. Length
0.91 mm (0.85-0.95 mm) and greatest width 0.46 mm (0.42-0.50
mm), based on 10 specimens. Ratio of length to width of prosome
1.49:1. Segment of leg 1 distinctly separated from head by a
transverse furrow. Epimeral areas of segments of legs 1-4 more or
less rounded posteriorly.
Segment of leg 5 (fig. 110) 68 » X 133 yw. Between this segment
and genital segment a weak ventral intersegmental sclerite. Genital
segment 138 u X 133 u, about as long as wide and somewhat expanded
laterally in its midregion. Areas of attachment of egg sacs located
dorsolaterally near middle of segment. Each area (fig. 111) bearing
two unequal naked setae 4 u and 12 wu in length and a small spiniform
process. Three postgenital segments 30 uw X 80 yw, 23 uw xX 75 pw, and
33 « X 72 uw, from anterior to posterior.
Caudal ramus (fig. 110) quadrate, 32 uw X 31 yu in greatest dimensions.
Outer lateral seta 104 uw, pedicellate dorsal seta 36 uw, outermost distal
seta 151 uw, innermost distal seta 221 u, and the two long median terminal
726 Proceedings of the Biological Society of Washington
Fics. 114-120. Lichomolgus telestophilus, new species, female: 114,
mandible, posterior (C); 115, first maxilla, posterior (C); 116, second
maxilla, posterior (C); 117, maxilliped, posterior (C); 118, leg 1,
anterior (E); 119, leg 2, anterior (E); 120, third segment of endopod
of leg 3, anterior (E).
Copepods Associated with Octocorals 127
Fics. 121-127. Lichomolgus telestophilus, new species, female: 121,
leg 4, anterior (E); 122, leg 5, dorsal (F). Male: 123, body, dorsal
(G); 124, urosome, dorsal (D); 125, maxilliped, inner (F); 126, endopod
of leg 1, anterior (C); 127, leg 5, dorsal (I).
setae 429 uw (outer) and 529 w (inner), both inserted between an un-
ornamented dorsal flap and a ventral flap with a marginal row of
minute spinules. Ornamentation of these setae similar to that in L.
longispinifer.
728 Proceedings of the Biological Society of Washington
Dorsal surface of prosome and urosome with very few hairs. Ratio
of length of prosome to that of urosome 2.62 : 1.
Egg sacs incomplete on all females collected. Each egg about 43 u in
diameter.
Rostral area (fig. 112) slightly raised ventrally and broadly tongue-
shaped.
First antenna 377 uw long, and resembling that of L. verseveldti.
Lengths of segments: 23 » (55 uw along anterior margin), 88 u, 26 u,
65 pw, 62 uw, 45 pw, and 36 w respectively. All setae naked. Second
antenna (fig. 113) 4-segmented, last segment elongated, 94 uw along its
outer edge, 64 uw along its inner edge, and 21 mw wide, bearing distally
five hyaline elements and two slightly unequal claws 57 uw and 53 wu in
length. One of three setae on third segment angularly bent. All setae
naked. Inner surfaces of all four segments with minute spinules.
Labrum as in L. verseveldti and L. hians. Mandible (fig. 114) rather
like that of L. hians, but with shorter barbs on flagellum than in that
species. Paragnath a small hairy lobe as in L. verseveldti. First maxilla
(fig. 115) with four naked elements. Second maxilla (fig. 116) some-
what resembling that of L. verseveldti, but inner distal spine near base
of lash rather blunt and finely barbed, instead of being attenuated and
ornamented with large spinules as in that species. Maxilliped (fig. 117) ©
with two setae on second segment very unequal, the larger seta with
two rows of erect spinuies in its midregion and more finely barbed
distally, the smaller seta naked. Third segment with two unequal ter-
minal barbed elements, one without an articulation.
Area between maxillipeds and first pair of legs as in L. verseveldti.
Legs 1-4 (figs. 118-121) segmented and armed with spines and
setae as in L. verseveldti, L. longispinifer, and L. hians. Inner seta on
coxa of leg 4 short (9 ») and naked. Inner margin of basis of leg 4
naked. Rami of all four legs resembling in general aspects those of
L. fissisetiger Humes and Ho, 1968b. Endopod of leg 4 (fig. 121) much
shorter than exopod. First segment 42 wu X 34 w (including processes),
with its distal inner finely barbed spine 33 » long and setiform. Second
segment 77 » long (including spiniform processes), its greatest width
33 uw and least width 19 uw; its two unequal terminal fringed spines 45 pu
(outer) and 83 uw (inner). Both segments with outer margins haired
and second segment with a terminal row of minute spinules.
Leg 5 (fig. 122) suggesting that of L. fissisetiger. Free segment
elongated, 133 » in length, width at expansion 42 yu, width distally 21 wu.
Two terminal setae 50 uw (outer) and naked and 81 uw (inner) with a
slight lamella along its midregion. Outer surface of free segment with
many small spines. Seta on body near free segment 46 uw and lightly
feathered.
Leg 6 probably represented by the two setae near areas of attachment
of each egg sac (see fig. 111).
Color in life in transmitted light slightly opaque, eye red.
Copepods Associated with Octocorals 729
Male: Body (fig. 123) resembling in general form that of female.
Length 0.75 mm (0.72—0.79 mm) and greatest width 0.31 mm (0.29-
0.31 mm), based on 10 specimens. Ratio of length to width of prosome
1.60: 1.
Segment of leg 5 (fig. 124) 36 u x 86 uw. No ventral intersegmental
sclerite. Genital segment 174 u x 161 uw, only a little longer than wide.
Four postgenital segments 15.5 uw x 50 yw, 14 w x 52 uw, 12 w X 50 w, and
20 uw X 53 wu, from anterior to posterior.
Caudal ramus similar to that of female, 24 uw x 23 u in greatest dimen-
sions.
Dorsal surface of prosome and urosome with very few hairs. Ratio
of length of prosome to that of urosome 1.91: 1.
Rostral area as in female. First antenna resembling that of female,
but with three aesthetes added as in the five previous species. Second
antenna similar to that of female but inner surficial spinules coarser
that in that sex.
Labrum, mandible, paragnath, first maxilla, and second maxilla as in
female. Maxilliped (fig. 125) slender and resembling in most respects
that of L. hians. Claw 130 wu along its axis including terminal lamella.
Area between maxillipeds and first pair of legs as in female.
Legs 1-4 segmented as in female, with same spine and setal formula
except for last segment of endopod of leg 1 (fig. 126) which is 1,I,4.
Legs 2-4 as in female, without sexual dimorphism.
Leg 5 (fig. 127) with free segment 38 uw xX 8.5 uw, bearing two terminal
elements, the outer setiform and naked, 48 uw long, the inner spiniform
with an outer barbed fringe and a few inner barbules, 25 uw. Surface
of segment with a few small spines. Seta on body near free segment
20 uw and lightly feathered.
Leg 6 (fig. 124) a posterolateral flap on ventral surface of genital
segment bearing two slender naked setae 24 uw and 55 uw long.
Spermatophore not observed.
Color in life as in female.
Etymology: The specific name telestophilus is a combination of the
generic name of the host and the Greek word ¢i\os = loving.
Comparison with related species: Only four species of Lichomolgus
have, as in the new species, the following combination of characters:
the second antenna with two short claws (shorter than the fourth
segment), the third segment of the exopod of leg 4 with the formula
II,1,5, the inner distal element on the first segment of the endopod of
leg 4 a spine and not feathered, and the dimensions of the caudal ramus
in the ratio of about 1: 1. These are L. cuneipes Humes and Ho, 1968b,
L. verseveldti, L. longispinifer, and L. hians. All four may be readily
distinguished from L. telestophilus, however. The free segment of the
fifth leg of L. cuneipes has a large wedge-shaped expansion. In L.
verseveldti this segment is relatively short and broad. In L. longispinifer
the outer spine on the first segment of the exopod of leg 1 is unusually
730 Proceedings of the Biological Society of Washington
long. In L. hians the two terminal elements on the maxilliped of the
female are widely divergent and the two setae on the second segment
of this appendage have a different size and ornamentation.
L. telestophilus might at first glance be confused with L. botulosus
Stock and Kleeton, 1963, from octocorals in the Mediterranean, but the
two species are easily separated by the nature of the lash on the second
maxilla and the ornamentation of the inner distal element on the first
segment of the endopod of leg 4.
Lichomolgus clavatus new species
Figures 128-148
Type material: 229 2, 198 6, and 1 copepodid from one colony of
Coelogorgia palmosa Milne Edwards and Haime, in 2 m, near village of
Ampasindava, northern end of Nosy Sakatia, near Nosy Bé, Madagascar,
collected 8 October 1960. Holotype @ allotype, and 35 paratypes
(192 2, 164 6) deposited in the United States National Museum, and
the remaining paratypes (dissected) in the collection of A. G. Humes.
Other specimens (all from Coelogorgia palmosa): 3929, 26 6, and
2, copepodids from one colony, in 1 m, Ambariobe, near Nosy Bé, 3
November 1960; and 1¢ from one colony, in 1 m, Ambariobe, 17
December 1963.
Female: Body (fig. 128) with broad prosome and slender urosome.
Length 2.27 mm (2.19-2.37 mm) and greatest width 1.09 mm (1.01-—
1.16 mm), based on 10 specimens. Ratio of length to width of prosome
1.08: 1. Segment of leg 1 almost completely fused with head. Epimeral
areas of legs 1-4 as in figure; tergum of segment of leg 3 overlapping
segment of leg 4.
Segment of leg 5 (fig. 129) 110 uw xX 264 w. Between this segment
and genital segment no ventral intersegmental sclerite. Genital segment
242, w long, in dorsal view not expanded laterally, its greatest width
238 w and its least width in its posterior third 204 uw. Areas of attach-
ment of egg sacs situated laterally in front of middle of segment. Each
area (fig. 130) bearing two naked setae about 13 uw jong. Three post-
genital segments 112 u X 177 uw, 78 w X 164 uw, and 221 uw x 174 u, from
anterior to posterior.
Caudal ramus (fig. 131) elongated, 286 u xX 77 w in greatest dimen-
sions, or 3.7 times longer than wide. All setae relatively short. Outer
lateral seta 42 uw and naked, pedicellate dorsal seta 39 w and lightly
feathered, outermost distal seta 47 « with lateral hairs proximally, inner-
most distal seta 61 mw and haired, and the two median terminal setae
85 » (outer) and 127 w (inner), both naked and inserted between an
unornamented dorsal flap and a ventral flap with a marginal row of
minute spinules.
Dorsal surface of prosome and urosome with small refractile points.
Ratio of length of prosome to that of urosome 1.18: 1.
Copepods Associated with Octocorals 731
Fics. 128-133. Lichomolgus clavatus, new species, female: 128,
body, dorsal (J); 129, urosome, dorsal (G); 130, area of attachment
of egg sac, dorsal (C); 131, caudal ramus, dorsal (D); 132, rostral
area, ventral (G); 133, first antenna, dorsal (D).
Egg sac (seen on only one female) elongated, 1090 u x 340 uw, reach-
ing to ends of ramal setae, and containing many eggs, each about 57 u
in diameter.
Rostral area (fig. 132) weakly developed.
732 Proceedings of the Biological Society of Washington
Fics. 134-141. Lichomolgus clavatus, new species, female: 134,
second antenna, anterior (inner) (D); 135, labrum, with paragnaths
indicated by dashed lines, ventral (E); 136, mandible, posterior (E);
137, first maxilla, anterior (C); 138, second maxilla, posterior (E);
139, maxilliped, posterior (E); 140, area between maxillipeds and first
pair of legs, ventral (B); 141, leg 1 and intercoxal plate, anterior (D).
Copepods Associated with Octocorals 733
Fics. 142-148. Lichomolgus clavatus, new species, female: 142,
leg 2, anterior (D); 143, third segment of endopod of leg 3, anterior
(E); 144, leg 4 and intercoxal plate, anterior (D); 145, leg 5, dorsal
(E). Male: 146, body, dorsal (A); 147, urosome, dorsal (G); 148,
maxilliped, inner (E).
734 Proceedings of the Biological Society of Washington
First antenna (fig. 133) 7-segmented, 554 uw long. Lengths of seg-
ments: 55 uw (97 uw along anterior margin), 180 pu, 39 uw, 85 wu, 64 uw, 57 p,
and 32 mw respectively. All setae naked. Second antenna (fig. 134)
4-segmented, last segment moderately elongated, 114 uw along its outer
edge, 75 uw along its inner edge, and 33 uw wide, bearing distally six
small hyaline elements and a claw 72 wu along its axis. All setae naked.
Labrum (fig. 135) rather deeply indented. Mandible (fig. 136) with
basal region distal to constriction showing on its-convex margin a scale-
like sclerotization with a row of long blunt spinules, and on its concave
margin a row of spinules interrupted by a sclerotization. Near base of
flagellum a dentate fringe (not rather delicately serrated as in many
other species). Flagellum elongated and finely barbed. Paragnath
(fig. 135) a small hairy lobe. First maxilla (fig. 137) with three long
terminal setae and a small subterminal element, the innermost terminal
seta finely barbed along one edge. Second maxilla (fig. 138) with first
segment unarmed. Second segment with a minute setule on its proximal
outer (ventral) surface, a surficial naked seta, an unusually long inner
(dorsal) seta bearing a row of erect spinules, and the segment ter-
minating in a relatively short lash bearing two large spines near its
base and two spikelike spinules along its midregion. Maxilliped (fig.
139) with two unequal naked setae on second segment; one of the
terminal elements long and clawlike.
Area between maxillipeds and first pair of legs as in figure 140.
Legs 1-4 (figs. 141-144) segmented and armed as in L. verseveldti,
L. longispinifer, L. hians, and L. telestophilus, except that formula for
endopod of leg 4 is 0-1; II. Coxa of leg 1 with an outer protuberance.
Inner seta on coxa of leg 4 long (112 uw) and feathered, instead of
much reduced as in many other species. Inner margin of basis of all
four legs with a row of hairs. Outer spines on exopod of leg 1 with
unusually strong lateral spinules. Endopod of leg 4 (fig. 144) a little
shorter than exopod. First segment 52 «X55 pw (including spinous
processes), with its distal inner seta long (155 mw) and feathered.
Second segment 122 u x 45 uw (including processes), its tip obliquely
truncated; its two terminal fringed spines 60 u (outer) and 70 wu (inner)
in one female, 66 w and 68 uw in another. Both segments with outer
margins haired and second segment with a terminal row of spinules.
Leg 5 (fig. 145) with an elongated unornamented free segment with-
out a proximal inner expansion, 170 uw X 32 yw in greatest dimensions
(ratio 5.3: 1), its two terminal setae 40 » (outer) and 55 wm (inner).
Seta on body near free segment 40 uw. All three setae naked.
Leg 6 probably represented by the two setae near areas of attachment
of each egg sac (see fig. 130).
Color in life in transmitted light slightly opaque to transparent, eye
red, egg sacs opaque.
Male: Body (fig. 146) with prosome less expanded than in female.
Length 1.74 mm (1.60-1.80 mm) and greatest width 0.64 mm (0.58-
Copepods Associated with Octocorals 735
0.70 mm), based on 10 specimens. Ratio of length to width of prosome
1.28: 1.
Segment of leg 5 (fig. 147) 60 u x 211 uw. No ventral intersegmental
sclerite. Genital segment subquadrate, 244 uw x 252 uw. Four post-
genital segments 86 uw X 133 uw, 83 uw x 125 », 55 w xX 114 w, and 169 ux
138 uw, from anterior to posterior.
Caudal ramus similar to that of female, 239 u x 60 x.
Dorsal surface of prosome and urosome with small refractile points
as in female. Ratio of length of prosome to that of urosome 1: 1.
Rostral area as in female. First antenna like that of female, but three
long asethetes added (fig. 146) as in six previous species, proximalmost
aesthete as long as entire first antenna. Second antenna, labrum,
mandible, paragnath, first maxilla, and second maxilla as in female.
Maxilliped (fig. 148) slender. Claw 140 wu along its axis.
Area between maxillipeds and first pair of legs as in female.
Legs 1-4 segmented and armed as in female. Leg 1 not showing
sexual dimorphism.
Leg 5 (fig. 147) with free segment 62 u xX 14 uw, without ornamenta-
tion, its two terminal setae 38 w and 43 uw. Seta on body near free
segment 36 uw. All three setae naked.
Leg 6 (fig. 147) a posterolateral flap on ventral surface of genital
segment bearing two naked setae 39 w and 47 u.
Spermatophore not observed.
Color in life as in female.
Etymology: The specific name clavatus, a Latin word meaning
furnished with nails or spikes, alludes to the spikelike elements on the
short lash of the second maxilla.
Comparison with related species: L. clavatus may be distinguished
from all other species in the genus by the combination of the following
features: the broad prosome and slender urosome, the moderately
elongated caudal ramus (3.7: 1 in the female) with relatively short
setae, the second antenna with a single claw, the formula of II,I,5 on
the last segment of the exopod of leg 4, a long feathered inner seta on
the coxa and on the first segment of the endopod of leg 4, the elongated
free segment of leg 5 (5.3: 1 in the female), and the absence of sexual
dimorphism in legs 1—4.
The new species is larger than most Lichomolgus. The females of
only three other species reach or exceed a body length of 2 mm: L.
gigas Thompson and A. Scott, 1903 (2 mm), L. magnificus Humes,
1964 (3.06 mm), and L. pterophilus Stock, 1962 (2.06 mm).
Three features of L. clavatus appear to be unique among the species
in the genus, as far as they are known. These are: the long blunt
spinules on the convex side of the mandible, the short lash with strong
spikelike spines on the second maxilla, and the long feathered inner
seta on the coxa of leg 4.
736 Proceedings of the Biological Society of Washington
REMARKS ON LICHOMOLGUS ASSOCIATED WiTH OCTOCORALS
Almost one-third of the known species of Lichomolgus (34 out of
105) are associated with octocorals,—25 with Alcyonacea, 4 with
Pennatulacea, 2 with Stolonifera, 2 with Telestacea, and 1 with
Gorgonacea. It seems very probable that the small number of species
reported from Gorgonacea reflects not the lack of preference of the
copepods for these hosts but rather the very few observations as yet made
on lichomolgids associated with them.
Among the species of Lichomolgus from octocorals there are several
external anatomical features which are worthy of note. Only in this
series of species does the endopod of leg 4 have the formula 0-1; 1,I
(as in L. protentus Humes and Frost, 1964, L. incisus Humes and Ho,
1968c, and L. insolens Humes and Ho, 1968c). Nine of the species
(L. cuneipes Humes and Ho, 1968b, L. exilipes Humes and Ho, 1968b,
L. fissisetiger Humes and Ho, 1968b, L. gentilis Humes and Ho, 1968b,
L. hians, L. longispinifer, L. telestophilus, L. varirostratus Humes and
Ho, 1968b, and L. verseveldti) have this formula as 0-I; II, a condition
not found in other Lichomolgus, except perhaps in L. anomalus A. Scott,
1909. (If Scott’s fig. 15, pl. LXVII, is correct, the inner element on
the first segment of the endopod of leg 4 is a spine. Such an armature
suggests to us that L. anomalus, obtained from washings of dredged
invertebrates from a depth of 1595 m, was actually associated with an
octocoral.) In all 34 species the second antenna is 4-segmented. In five
species (L. aculeatus, L. exilipes, L. fissisetiger, L. gentilis, and L.
varirostratus, all described by Humes and Ho, 1968b), the two claws
on the second antenna are very long (as long as or longer than the
last segment). In two species (L. cristatus Humes and Ho, 1968c, and
L. dentipes Thompson and A. Scott, 1903) the flagellum of the mandible
is extremely short and reduced to a small pointed process. In three
species (L. conjunctus Humes and Ho, 1967c, L. decorus Humes and
Frost, 1964, and L. organicus Humes and Ho, 1967c, the convex margin
of the mandible has a prominent toothlike process, replacing the scale-
like spinulose area seen in other species.
Two features of these species, the number of claws on the second
antenna and the formula for the last segment of the exopod of leg 4,
vary without correlation. Hence they are useful in an artificial key, but
probably do not indicate natural groups. Eleven species have one
claw with the formula II,I,5 in four of them and III,I,5 in seven.
Twenty-three species have two claws with II,I,5 in nineteen and III,1,5
in four.
In three species among those from octocorals, L. trispinosus Stock,
1959, L. cristatus Humes and Ho, 1968c, and L. singularipes Humes
and Ho, 1968c, males are unknown. In the remaining 31 species sexual
dimorphism in the formula for the endopod of leg 1 (1,5 in the female,
1,1,4 in the male) occurs in all but four (L. pteroidis Della Valle, 1880,
Copepods Associated with Octocorals 137
L. pterophilus Stock, 1962, L. serratipes Ummerkutty, 1962, and L.
clavatus).
Characters especially useful in distinguishing the species of Lichomol-
gus associated with octocorals are: the number of claws on the second
antenna and their length in relation to the last segment, the length of
the flagellum and the nature of the convex margin of the mandible
(whether a toothlike process or a spinulose scalelike crest), sexual
dimorphism in the formula for the last segment of the endopod of leg 1,
the formula for the last segment of the exopod and for the endopod of
leg 4, the length to width ratio and the shape of the free segment of leg
5 in the female (presence or absence of a proximal inner expansion), and
the length to width ratio and the nature of the armature of the caudal
ramus.
Keys To THE 52 SpEcIES oF LICHOMOLGUS KNOWN FROM THE REGION OF
Nosy Bt, MADAGASCAR
The following keys include those species not only from octocorals but
from all other invertebrate hosts as well. The number of species now
known from Madagascar comprises about half of the known species
in the genus.
These artificial keys are intended only as aids in identification. In
many instances it has been necessary to rely upon qualitative rather
than quantitative characters for distinctions. Final determination of a
species should be made by reference to the original description and
figures, or to actual specimens, if available.
The copepod referred to by Humes and Cressey (1961) as Lichomol-
gus oreastriphilus (Kossmann, 1877), from asteroids at Nosy Bé, is in
reality a Stellicola and therefore not included in the keys.
The host for L. protentus was cited in the original description by
Humes and Frost (1964) as Sarcophyton sp., but has since been
described as S. globosum n. sp. by Tixier-Durivault (1966).
Descriptions of several new species of octocorals listed as hosts may
be found in the work of Verseveldt (1968).
FEMALES
1. Last segment of exopod of leg 4 with formula II,I5 — 9)
Last segment of exopod of leg 4 with formula III,I,5 Al
Za SeCOndyantenmeas witli Claw) eee 3
Seconqmantennamwithy 2) Clays) see eee sean ee eae ee 20
3. A large sometimes toothlike process on convex margin of
rsa ve Weta I 0) (=\peLil sD eT Tek SON DF ae dee Re, ea RR 4
Convex margin of mandible without such a process or at most
Withee —Ancmallidigitit orm) processes!) aa es lle wene ues ee 6
4. A setiferous sphere on second segment of second maxilla (from
corals, Pavona angulata, P. danai, P. cactus, and P. Pvenusta) _.
elinedia els Yds BW CP L. actinophorus Humes and Frost, 1964
738 Proceedings of the Biological Society of Washington
Without: such iaxsphere sive hoist 00 ee 5
5. Free segment of leg 5 small, 34 « x 13 uw, without proximal inner
expansion (from coral, Alveopora sp.) —......--__---__-
UWE Viste ile Oe Nee Nee Pe L. campulus Humes and Ho, 1968a
Free segment of leg 5 larger, 90 uw X 22 uw, with a rounded
proximal inner expansion (from coral, Psammocora contigua) _..
i 0 A = NR SU ae SEU L. rhadinus Humes and Ho, 1967a
6. Convex margin of mandible with 2-4 small digitiform processes _. 7
Convex margin of mandible without such processes _....._- 10
7. A prominent posteroventral lobe on first postgenital segment
(from corals, Acropora scherzeriana, A. cytherea, and Acropora
SPs 8) eee ee ee EW L. lobophorus Humes and Ho, 1968a
Without such a.sloboe 2 8
8. Segments 3 and 4 of second antenna fused; free segment of leg
5 strongly arched (from coral, Acropora palifera) __.._..___.
eae aka Eee al ha rae L. arcuatipes Humes and Ho, 1968a
Segments 3 and 4 of second antenna not fused; free segment of
leg +5. mot arched |: eb oo el PA 0 ae 9
9. Free segment of leg 5 small, 36 u x 17 uw, ratio about 2: 1 (from
corals, Goniopora sp. and Porites sp.) _.._-- sha el ;
dv RUG EA MMOLE GAN) 4.5 Laie a seein L. digitatus Humes and Ho, 1968a
Free segment of leg 5 elongated, 203 u x 30 u, ratio about 6.8 : 1
(from corals, Porites sp. cf. P. andrewsi, P. sp. cf. P. nigrescens,
Biel Jonesy (Ue; SOTA). Gjo)) ee
Be ie 1 Sit Ac AOL eee 3 L. prolixipes Humes and Ho, 1968a
10. Formula for endopod of leg 4= 0-1; 1,J 11
Formula for endopod of leg 4= 0-1; Il _ 12
11. Seta on first segment of endopod of leg 4 naked; third segment
of maxilliped with swollen membranous outer edge (from
octocoral, Sarcophyton ehrenbergi) _......-..-------
SP Ade cg SU 0 ae ay aoe oe L. incisus Humes and Ho, 1968c
Seta on first segment of endopod of leg 4 feathered; third seg-
ment of maxilliped without swollen membranous outer edge
(from octocoral, Lobophytum crassum) _....-.--------
oN i RAN les Was il Bird aah Rn L. insolens Humes and Ho, 1968c
12. Length exceeding 2 mm (2.19-2.37 mm); inner coxal seta long
and feathered (from octocoral, Coelogorgia palmosa)
SU A a Sas sc A a L. clavatus Humes and Ho, above
Length distinctly less than 2 mm; inner coxal seta much reduced
and either naked or at most with minute barbules _......_.-___- 13
13. Free segment of leg 5 minute, less than 25 w long, and not
reaching anterior border of genital segment 14
Free segment of leg 5 large, at least 80 u in length, reaching
beyond anterior border of genital segment —......-_. 15
14. Rostral area broadly rounded and slightly triangular; terminal
15.
16.
Wi,
18.
19.
bo
i)
Copepods Associated with Octocorals 739
setae on caudal ramus haired (from pelecypods, Asaphis rugosa
and Standella solanderi) _..... L. asaphidis Humes, 1959
Rostral area forming a long triangular beak; terminal setae on
caudal ramus naked (from pelecypod, Chama iostoma) _....
PORARUARS EOL liye Runa M NN. ONS at L. chamarum Humes, 1968
Second antenna 3-segmented (third and fourth segments
PUTS lh) pes ese ie te aR Se 2 A dE SR 16
Second antenna 4-segmented _....__ 18
Prosome broad, about as long as wide, with urosome flexed
beneath it; outer spines on exopods of legs 2—4 with smooth
lamellae (from corals, Stylophora pistillata, S. mordax, and
ING HOGNOUES S)OW)), atc L. crassus Humes and Ho, 1968a
Prosome not unusually broad, ratio about 1.3: 1; urosome held
posteriorly; outer spines on exopods of legs 2-4 with finely
Spintlosembamell ae pace mend a Eee IS Ne ana 17
Caudal ramus with ratio 9.1:1 (from corals, Stylophora
pistillata, S. mordax, and Acropora sp.) --...--..----—--—------
1 i Us L. geminus Humes and Ho, 1968a
Caudal ramus with ratio 5 : 1 (from corals, Seriatopora subseriata,
Sn CGuojouGe, eiatel Wanaojieme Won) ee
ule reneaee Nea aca tO LE L. compositus Humes and Frost, 1964
Caudal ramus as long as wide (from octocoral, Sarcophyton
UEUCULIT) eee aa L. spathophorus Humes and Ho, 1968c
@audalaramusilonger.than! wide sss sel ie easa OS ene 19
Free segment of leg 5 with a proximal inner expansion (from
sea anemone, Rhodactis rhodostoma) _---------------------
MMA aa Zi ts LAM nt Ey L. simulans Humes and Ho, 1967b
Free segment of leg 5 without a proximal inner expansion (from
sea anemone, Rhodactis rhodostoma) __...----------------------
1 AAR RO SAY ate a NO L. politus Humes and Ho, 1967b
Convex margin of mandible with a strong toothlike process __ 21
Convex margin of mandible without such a process _.- ob)
Caudal ramus 36 u X 24 w (1.5: 1); genital segment broadest
near middle (from octocoral, Tubipora musica)
MH Des Blea Bae aE ten Vi h L. organicus Humes and Ho, 1967c
Caudal ramus 28 uw X 23 uw (1.22: 1); genital segment broadest
more posteriorly (from octocoral, Tubipora musica)
BUN SII SUSU oA OR L. conjunctus Humes and Ho, 1967c
Ratio of caudal ramus distinctly more than 1.5:1] 23
Ratio of caudal ramus distinctly less than 1.5: 1, often 1:1 or
eventawicen thanglomeiy se. SUN eee, BS isis Me ee ee 26
Longer claw on second antenna longer than greatest length of
fourth segment (from octocorals, Nephthya aberrans, N. sphaero-
phora, N. tixierae, N. crassa, and Litophyton arboreum) __.
PU CMAN Ported iLL ase cA Ah L. aculeatus Humes and Ho, 1968b
740 Proceedings of the Biological Society of Washington
26.
2.
29.
30.
31.
Longer claw on second antenna shorter than greatest length of
fourth: segment ills iia 7 he CT Na EDs 94
Caudal ramus 3.3: 1 (from octocorals, Lemnalia sp., L. flava,
L. elegans, L. amabilis, L. africana, and Paralemnalia thyrsoides )
a ot Tea ORRIN i RP L. spinulifer Humes and Frost, 1964
Caudaliramusyalso witli 1eS, = 9 ssa ee 25
Free segment of leg 5 83 uw X 35 uw, without a proximal inner
expansion (from sea anemones, Stoichactis giganteum and
JRO AGUS Taare) eo L. cuspis Humes, 1964
Free segment of leg 5 75 uw X 19 uw, with a proximal inner expan-
sion where width is 31 uu (from zoantharians, Palythoa tuber-
Culosaandweaeliscia) jeamaanen L. inaequalis Humes and Ho, 1966
Longer claw on second antenna about as long as or longer than
greatest lengthy of second se game nt an aera 27
Longer claw on second antenna shorter than greatest length of
second segments) 8 ik REC Tees ate reaieg ge 30
Two long terminal setae on caudal ramus naked; a prominent
unguiform process (twice the length of the setae) on area
of attachment of egg sac (from octocorals, Dendronephthya
mucronata, D. regia, D. stocki, and D. kollikeri)
EN Senne SO) Cea OD, RES NY L. varirostratus Humes and Ho, 1968b
Two long terminal setae on caudal ramus haired; without such a
prominent unguiform process on area of attachment of egg sac _. 28
. Free segment of leg 5 with a weak proximal inner expansion;
longer claw on second antenna distinctly longer than greatest
length of fourth segment (143 u to 130 ») (from octocorals,
Dendronephythya mucronata, D. kollikeri, D. stocki, Stereone-
phthya (acaulis. and Se papyiaced) ee ae cee
HAN AN eS goes ed AMS beat nad tt L. gentilis Humes and Ho, 1968b
Free segment of leg 5 with a well developed proximal inner
expansion; longer claw on second antenna about as long as
greatest) lengthy ofa fourths isecmaenb ewmnilesic sil ae Oneness nen Onna 29
Free segment of leg 5 148 uw long; lateral contour of genital
segment in dorsal view rounded (from octocorals, Stereonephthya
acaulis, S. papyracea, and Lemnalia elegans)
ELS MRA SH ah sa CU L. fissisetiger Humes and Ho, 1968b
Free segment of leg 5 172 w long; lateral contour of genital
segment in dorsal view angular rather than rounded (from
octocorals, Dendronephthya mucronata, D. regia, D. stocki, and
D skollikerni\ Ak) Ua Leoni L. exilipes Humes and Ho, 1968b
Formulator; endopodof leg, 4.101. yi manasa nna 31
Formula’ for endoped (of les 4 101k Illi jesse ae a a 35
Free segment of leg 4 117 u X 26 uw, without a proximal inner
expansion (from octocoral, Siphonogorgia pendula) __.......
sl ee Ma Al UN Dal lO L. hians Humes and Ho, above
Free segment of leg 5 with a distinct proximal inner expansion _ 32
32.
33.
34.
35.
36.
37.
38.
39.
40.
Copepods Associated with Octocorals 7A
Free segment of leg 5 with a broad rounded expansion 33
Free segment of leg 5 with a pointed expansion, directed distally _ 34
Genital segment broadest near middle; free segment of leg 5
rather slender (from octocoral, Telesto arborea) __........-
hE Tia AY eNO ge L. telestophilus Humes and Ho, above
Genital segment broadest anterior to middle; free segment of
leg 5 moderately broad (from octocoral, Heteroxenia elisabethae)
A EY OY Se ee OLS Pe L. verseveldti Humes and Ho, above
Setae on caudal ramus naked; genital segment broadest near
middle; spine on first segment of exopod of leg 1 of usual length
(from octocoral, Stereonephthya acaulis) _
ILS is EU SCO. ‘cuneipes Humes and Ho, 1968b
Setae on caudal ramus mostly haired; genital segment broadest
posterior to middle; spine on first segment of exopod of leg 1
unusually long (from octocoral, Siphonogorgia pendula) __.
ANE UP, ae nea ew L. longispinifer Humes and Ho, above
Free segment of leg 5 small, less than 50 wu long 36
Free segment of leg 5 large, more than 80 uw long _.....-. 37
Proximalmost outer spine on third segment of exopod of legs
1-3 shorter than others; seta on first segment of endopod of leg
4 very lightly feathered; free segment of leg 5 27 nx 12 u
(from octocorals, Cladiella pachyclados and C. krempfi) __...
SUS DALY MAT tN NL L. hetaericus Humes and Ho, 1968c
Proximalmost outer spine on third segment of exopod of legs
1-3 not shorter than others; seta on first segment of endopod
of leg 4 conspicuously feathered; free segment of leg 5 49 up x
20 uw (from nudibranch, Phyllidia trilineata)
SDE NANPA alee) Pea eT oe MAL a CORN NOE i A NON L. venustus Humes, 1959
Caudal ramus a little longer than wide, ratio 1.3: 1 (from
nudibranch, Trevelyana rubromaculata) _
puoreae PUTA NAN Ue HAN MeO EN EN G1 Salute NS Ue L. sensilis Humes, 1964
Caudal ramus about as long as wide or a little wider than long _ 38
Free segment of leg 5 without a basal expansion 39
Free segment of leg 5 with a proximal inner expansion ________ 40
Free segment of leg 5 broad, 81 uw X 34 4, with relatively large
scalelike spines on outer surface (from octocoral, Parerythropodium
rubiginosum) ____....---------- L. singularipes Humes and Ho, 1968c
Free segment of leg 5 slender, 98 uw <X 22 u, with small spines on
outer surface (from nudibranch, Phyllidia trilineata) —
UTES ON ASIEN REO) Gl OMT L. patulus Humes, 1959
Expansion slight; outer surface of free segment of leg 5 with
small spinules (from nudibranchs, Hexabranchus orbicularis and
Dondopsis\ruber) mse sn L. commodus Humes, 1964
Expansion large and rounded; outer surface of free segment
of leg 5 smooth (from nudibranch, Doris mabilla) _-
URN IRMA SMUb A ONIN ged MLE YO NT AS L. securiger Humes, 1964
742 Proceedings of the Biological Society of Washington
Ale (Second! antennas wathielync le vyjeame meee alas ashen 42
ecornel einer Satie) 2 GEN 48
42. Formula of endopod of leg 4=0-1; 1,I (from octocoral,
Sarcophyton globosum) __.- L. protentus Humes and Frost, 1964
Hormulavofsendopodioflegy4 = 0=1- si) Sain eee nnn 43
43. Caudal ramus distinctly longer than wide (at least 3.5: 1) 44
Caudaljramus fabout asi longias nwaic cheese eee 45
44, Setae on first antenna haired; mandible with an extremely short
flagellum; lash of second maxilla with a crest of long spinules
(from octocoral, Sinularia leptoclados)
SRY s Nel Me eee Ee ered L. cristatus Humes and Ho, 1968c
Setae on first antenna smooth; mandible with moderately long
flagellum; without crest of long spinules on lash of second
maxilla (from octocoral, Xenia umbellata)
AL AN eR L. glabripes Humes and Ho, above
45. Free segment of leg 5 with an inner expansion __...._...___- 46
Free segment of leg 5 lacking an inner expansion —_____________. 47
46. Inner expansion in the form of a large tooth (from octocoral,
Sinularia humesi) —- L. dentipes Thompson and A. Scott, 1903
Inner expansion large, not well delimited, and giving a triangular
appearance to the segment (from octocoral, Anthelia gracilis )
RIE RAR AS Nl ioe cea Ra ea ally L. triquetrus Humes and Ho, above
47. Free segment of leg 5 195 u X 26 uw, with spines on outer surface
arranged in two rows proximally and in one row distally (from
octocorals, Sinularia polydactyla, S. pedunculata, and S. white-
leg een) ine se NE ON es Rae L. adelphus Humes and Ho, 1968c
Free segment of leg 5 82 u X 34 uw, with spines on outer surface
not arranged in rows (from octocorals, Sinularia polydactyla and
Sawhitelegege?)) mwas aes L. squamiger Humes and Frost, 1964
48. Body length 3 mm; caudal ramus 7.8: 1 (from sea anemone,
Stoichactis giganteum) __.....--- L. magnificus Humes, 1964
Body length less than 1.5 mm; caudal ramus 2 : 1 or less, in some
even, widerthan long ie eee a oe eee ea 49
49. Seta on first segment of endopod of leg 4 naked; mandible with
a spiniform or toothlike process on convex margin 50
Seta on first segment of endopod of leg 4 feathered; mandible
without a spiniform or toothlike process 51
50. Free segment of leg 5 short, 33 « X 15 w, with an inner expan-
sion but unornamented (from octocoral, Cladiella laciniosa)
RURK Lene p alters! Ore te eV ire yWLL COM eas L. decorus Humes and Frost, 1964
Free segment of leg 5 elongated, 101 u X 15 uw, with an inner
expansion and outer surface with spinules (from octocorals,
Cladiella krempfi, C. laciniosa, and C. pachyclados)
CL Si NEAR kG SIAR ee eC ee L. foxi Gurney, 1927
51. Inner surface of segment 2 of second antenna with small spines;
Copepods Associated with Octocorals 743
caudal ramus wider than long, 24 u xX 30 uw (from nudibranch,
AOS SCAnG)) L. audens Humes, 1959
Inner surface of segment 2 of second antenna smooth; caudal
ramus longer than wide, 55 ux 31 uw (from sea anemone,
Stoichactis giganteum) L. gemmatus Humes, 1964
MALES
(The males of L. cristatus Humes and Ho, 1968c, and L. singularipes
Humes and Ho, 1968c, are unknown, and hence are not included
in the key. )
1) Kast segment of exopod with formula ID5 2s 2
Last segment of exopod with formula IIILI,5 — 40
2 Saconel sinvsorny wat I CEN? ee 3
Seconcl ain vain O dh ww eee 20
3. A large sometimes toothlike process on convex margin of
ANAT GL fo yeni eee Mew ales ALU ae SURE os Me Mao te 4
Convex margin of mandible without such a process or at most
with~2—4) small digitiform) processes) ee 6
4. A setiferous sphere on segment 2 of second maxilla
10.
11.
PAE i Ei NESE ONY L. actinophorus Humes and Frost, 1964
Wathoutisuchpausphere ye... 2) leah ee 5
Last segment of endopod of leg 1 =1,I,4; free segment of leg
1S 6 5< 6 me Gamal sens Jeehs 9 il
ADS AL ASR ae OS SE aa ae L. rhadinus Humes and Ho, 1967a
Last segment of endopod of leg 1=1,5, as in female; free
segment of leg 5 15 1» x 9 uw; caudal ramus 3.2:1
Lele iE NUCAie Rieaeed ote on ae Oma L. campulus Humes and Ho, 1968a
Convex margin of mandible with 2—4 small digitiform processes 7
Convex margin of mandible without such processes __.._._.____- 10
A prominent posteroventral lobe on second postgenital segment _
PEMOMLASN ES i ph ait she sNabet, RE, L. lobophorus Humes and Ho, 1968a
Nithoutssuchinat lobe gauss tS Se eee ene dues Dee) S 8
Segments 3 and 4 of second antenna fused; segment 2 of maxil-
liped with two rows of prominent spinules —
MMe ais a ee BRAS ood tet ia, L. arcuatipes Humes and Ho, 1968a
Segments 3 and 4 of second antenna not fused; segment 2 of
maxilliped with only one row of prominent spinules 9
Caudalaramuse2 i: lone L. prolixipes Humes and Ho, 1968a
Caudaleramus) 4:3 31 L. digitatus Humes and Ho, 1968a
Hormulastonsencopod) of les 40) EE 11
Fornnulastormendopodsof ler 40-1) 1h ee 1D
Seta on first segment of endopod of leg 4 naked; inner margin
of segment 2 of second antenna with a notched lamella; concave
eaverot claw, ofmmaxillipedysmooth = ene
Ee Sib aE, Eee GS eS L. incisus Humes and Ho, 1968c
744 Proceedings of the Biological Society of Washington
12.
13.
14.
15.
16.
Np
18.
119}
Seta of first segment of endopod of leg 4 feathered; inner
margin of segment 2 of second antenna with a striated membrane;
concave edge of claw of maxilliped with hyaline knobs ______
SSR Rah ASS EIR AA RU L. insolens Humes and Ho, 1968c
Length 1.74 mm (1.60-1.80 mm); inner coxal seta long and
FEASTS WN A MRI PLU en area L. clavatus Humes and Ho, above
Length not exceeding 1.55 mm; inner coxal seta much reduced
and either naked or at most with minute barbules ___________. 13
Last segment of endopod of leg 1 with formula I,5, as in
Perri ll ey i ES ee oH | URS a SUT ENG ete 14
Last segment of endopod of leg 1 with formula I,I,4 16
Caudal ramus 4.4 : 1, its setae relatively long and naked _________
Cul? anita edie ada OL a Ean EU ASU tee L. chamarum Humes, 1968
Caudal ramus at least 5.13: 1, its setae otherwise 15
Two terminal setae on caudal ramus not broadened, ornamented
with hairs; concave margin of claw of maxilliped smooth; free
segment of leg 5 minute, 18 uw xX 12 uw _.. L. asaphidis Humes, 1959
Two terminal setae on caudal ramus naked and slightly broad-
ened; concave margin of claw of mazxilliped with hyaline
dentation; free segment of leg 5 larger, 35 uw X 10 w
Precis Mee OMEN Jebel CaM 92 SAL Lt L. crassus Humes and Ho, 1968a
Second antenna with third and fourth segments fused 17
Second antenna 4-segmented _. 18
Outer of two terminal spines on last segment of endopod of
second leg not modified _._. L. compositus Humes and Frost, 1964
Outer of two terminal spines on last segment of endopod of
second leg modified (truncated and broadened at tip)
Te SAN BU AEN ENON Nea UO L. geminus Humes and Ho, 1968a
Caudal ramus about vas) lonewas |wicdes seme
sii OOS REP a IEA eR ennrce Ae L. spathophorus Humes and Ho, 1968c
Caudal! ‘ramus’ distinctly longer thaniwide) 2 a ae 19
Length 1.33 mm (1.26-1.40 mm); caudal ramus 138 pu x 36 yu,
TALIO SEO ested get RAL NO L. politus Humes and Ho, 1967b
Length 0.96 mm (0.85-1.08 mm); caudal ramus 52 u x 34 u,
TALIO | Mc) sili Coc wee net che aees L. simulans Humes and Ho, 1967b
Convex margin of mandible with a strong toothlike process _.. 21
Convex margin of mandible without such a process ___.._____- SY)
Caudal ramus 31 yu X 21 u, ratio 1.48: 1; free segment of leg
5 36 u xX 9 yw, ratio 4:1 L. organicus Humes and Ho, 1967c
Caudal ramus 25 uw X 21 wu, ratio 1.19: 1; free segment of leg
5) ONS} hs SKS My Tee) Sis) BIL L. conjunctus Humes and Ho, 1967c
Longer claw on second antenna about as long as or longer than
greatest lengthy of fourth) segmiemt) suamueenialee enn eee enna 23
Longer claw on second antenna shorter than greatest length of
fourth) Seeiment: i SUC Le ei ais Nae Nhe Ge len 7
. Caudal ramus 2: 1; terminal spine on third segment of endopod
a
25.
26.
27.
OX),
30.
3l.
32.
33.
Copepods Associated with Octocorals 745
of leg 1 concave with two rows of strong spinules
Dees TIL 6 Ces IRR A CAL LS L. aculeatus Humes and Ho, 1968b
Caudal ramus about 1:1; terminal spine on third segment of
endopod of leg 1 otherwise 24
Rostral area pointed, not broadly rounded as in female; two
terminal setae on caudal ramus naked; free segment of leg 5
33 uw X 9 pw, ratio 3.66: 1 __ L. varirostratus Humes and Ho, 1968b
Rostral area rounded; two terminal setae on. caudal ramus haired;
free segment of leg 5 with ratio of 5-5.77: 1 — 25
One of two setae on second segment of maxilliped terminating
in several pointed spiniform elements; free segment of leg 5
WithmrationOhoiincele jee L. fissisetiger Humes and Ho, 1968b
Both setae on second segment of maxilliped normally attenuated;
free segment of leg 5 with ratio close to 5: 1 (not more than
530/31) dy AES Seo a ONO RM UU NOOR La ales 26
Outer spine on last segment of endopod of leg 1 angularly bent;
genital segment with lateral margins rounded posteriorly ___
pe calisal Lien dimen Na IN L. exilipes Humes and Ho, 1968b
Outer spine on last segment of endopod of leg 1 not strongly
bent; genital segment with lateral margins angular posteriorly ___.
nA) a ES Oe L. gentilis Humes and Ho, 1968b
Spine on first segment of exopod of leg 1 unusually long (about
twice length of succeeding spines)
CODUESU OU NALA UES ea ot La L. longispinifer Humes and Ho, above
Spine on first segment of exopod of leg 1 of usual length 28
Setae on caudal ramus naked _.._....- 29
Setae on caudal ramus haired __.....---- 30
First segment of endopod of leg 4 with slightly barbed spine __
tei NL a eR IESE L. cuneipes Humes and Ho, 1968b
First segment of endopod of leg 4 with feathered seta ______.
OPS SMe YESS eRe eR TUN |e VLR IR L. venustus Humes, 1959
Endopod of leg 4 with formula 0-1; II, first segment with a
FeEatherednsebate = We seen ek ee ult ual sa ALB ea Agr nel ae ee 31
Endopod of leg 4 with formula 0-I; II, first segment with a very
finelybarbed jon makeduspine, seer We ae eae eweu eee 38
One of two setae on second segment of maxilliped strongly
modified, with base swollen and spinose and distal part slender
AN CWOT AE Ci wena ies al evel eA aeRO L. cuspis Humes, 1964
Neither of two setae on second segment of maxilliped so
TORLOVGADALSTOL ease Ae UE WS UN CU NUL OMS Ace Seer NOt Rr ea 32
Second antenna like that of female, without ornamentation
Cll ec geet sua NN eh a NE L. inaequalis Humes and Ho, 1966
Second antenna with ornamentation on inner surface of second
SE OG TMA iat te ean RAD ere AR a NIC ll LAE a a eS alae ath ein 33
This ornamentation consisting of hairs _. L. commodus Humes, 1964
746 Proceedings of the Biological Society of Washington
This ornamentation consisting of spinules (in some cases knob-
MN Gi) iy Leek es ek AEN IN TOS AAR
345 .CaudalGrammus long ergnthvam| evil cl eee sme alee
Caudal ramus with ratio close to 1: 1, or wider than long _____
35. Caudal ramus 1.3: 1; inner surface of segment 2 of second
antenna, with small knobsin(spinul estan) am mnaneenansne canner
PATS AN AE oil A ee al acs L. sensilis Humes, 1964
Caudal ramus 2.44: 1; inner surface of segment 2 of second
antenna with strong spinules __ L. spinulifer Humes and Frost, 1964
36. Inner distal spine on segment 2 of second maxilla with spinules
on one side; seta on first segment of endopod of leg 4 very
lightly feathered _.- L. hetaericus Humes and Ho, 1968c
Inner distal spine on segment 2 of second maxilla with spinules
on both sides; seta on first segment of endopod of leg 4 well
feathered: eth leuncusend pte A see se ole atse Okt IN an da ele
37. Free segment of leg 5 small, 20 » x 9 uw, without fine ornamenta-
rat es te i ee ae Ra oe L. securiger Humes, 1964
Free segment of leg 5 larger, 87 u x 14 uw, with small spinules
ONMOULET SUTLa Ce —aamakenme man neenE ee AESS L. patulus Humes, 1959
38. Longer claw on second antenna about 80 per cent of greatest
length of fourth segment; slight sexual dimorphism in third
segment of endopod of leg 2 _.. L. hians Humes and Ho, above
Longer claw on second antenna about 61 per cent of greatest
length of fourth segment; without sexual dimorphism in third seg-
ment) of endopodofi leg. 2 pe ssi oe
39. One of two setae on second segment of maxilliped with a blunt
finely spinulose tip —__. L. verseveldti Humes and Ho, above
Both setae on second segment of maxilliped attenuated and
Naked, keds uss peu tears L. telestophilus Humes and Ho, above
20); Secemel ainsi wrdn J) Geyy
Seconel mime wadn O Cewyg 2
Al Hormulay forsendopodsof leew 4— 0-1-1 51-] at ee ee
PE Sec AUG tal EOS Oars UA AOS TIEN) 0 Dh L. protentus Humes and Frost, 1964
Hormulay for endopodgok leg e4 == 10—1 ili ae ee
42. Mandible with extremely short flagellum; last postgenital seg-
ment much wider than preceding three __.._---________
AE) ARE RY EN Os L. dentipes Thompson and A. Scott, 1903
Mandible with long flagellum; last postgenital segment not notice-
ably wider than preceding segments
43. Segment 2 of second antenna with inner surface unornamented,
as in female; seta on first segment of endopod of leg 4 naked _.
SA OM TEA. aa L. triquetrus Humes and Ho, above
Segment 2 of second antenna with inner surface ornamented
with spinules or spines; seta on first segment of endopod of
Leow. feathered ss cs ste jee BCA aoe
44. Segment 2 of second antenna with few scalelike spines on inner
45.
46.
47.
48.
49,
Copepods Associated with Octocorals TAT
Surbaceyy 22 eens Le eee) L. glabripes Humes and Ho, above
Segment 2 of second antenna with many spinules on inner
SULT CO Bats ase Mievmmne me Me sd MUERTE RS 45
Last segment of endopod of leg 2 with spiniform process be-
tween two terminal spines broadly triangular
SEMIN atts 6.50 BAMA ata | L. adelphus Humes and Ho, 1968c
Last segment of endopod of leg 2 with spiniform process between
two terminal spines in the form of a bent thumb
BU AVN SRI LE L. squamiger Humes and Frost, 1964
Body length 2.74 mm; caudal ramus 6.7: 1 _.........-
SS hg SE Me a EU a ne NS L. magnificus Humes, 1964
Body length not exceeding 1 mm; caudal ramus less than 2: 1 _. 47
Last segment of endopod of leg 1 with formula I,II,3; segment
2 of second antenna with refractile knobs on inner surface __
PA UHUD LCE AUN cg RASS TOWN IE USAC A L. gemmatus Humes, 1964
Last segment of endopod of leg 1 with formula I,I,4; segment
2 of second antenna without refractile knobs on inner surface __. 48
Seta on first segment of endopod of leg 4 naked; genital segment
longer thamwade tills sol) ia L. foxi Gurney, 1927
Seta on first segment of endopod of leg 4 feathered; genital
Segment n Ot long err ehary wi G ey eevee al wnteaby tse cava en ieee 49
Segment 2 of second antenna with smooth inner surface, as
in female; seta on first segment of endopod of leg 4 weakly
feathered in lise umn iavageal L. decorus Humes and Frost, 1964
Segment 2 of second antenna with spinules on inner surface;
seta on first segment of endopod of leg 4 well feathered __-
L. audens Humes, 1959
LITERATURE CITED
DELLA VALLE, A. 1880. Sui Coriceidi parassiti, e sull’anatomia del
gen. Lichomolgus. Atti R. Accad. Lincei, ser. 3, Mem. Cl.
Sci. Fis., Math. Nat., 5: 107-124.
Gurney, R. 1927. Zoological results of the Cambridge expedition to
the Suez Canal, 1924. XXXIII. Report on the Crustacea:
Copepoda (Littoral and semi-parasitic). Trans. Zool. Soc.
London, 22: 451-577.
Humes, A. G. 1959. Copépodes parasites des mollusques a Madagas-
car. Mém. Inst. Sci. Madagascar, 1958, sér. F, 2: 285-342.
1964. New species of Lichomolgus (Copepoda, Cyclopoida)
from sea anemones and nudibranchs in Madagascar. Cahiers
O.R.S.T.O.M. Océanogr., 1963 (série Nosy Bé II), no. 6,
pp. 59-130.
1968. Two new copepods (Cyclopoida, Lichomolgidae)
from marine pelecypods in Madagascar. Crustaceana, suppl.
1, Studies on Copepoda, pp. 65-81.
748 Proceedings of the Biological Society of Washington
Humes, A. G. AnD R. F. Cressry. 1961. Lichomolgus oreastriphilus
(Kossmann), un copépode cyclopoide parasite des étoiles de
mer a Madagascar. Mém. Inst. Sci. Madagascar, 1959, sér.
F, 3: 83-92.
Humes, A. G. anp B. W. Frost. 1964. New lichomolgid copepods
(Cyclopoida) associated with alcyonarians and madrepo-
rarians in Madagascar. Cahiers O.R.S.T.O.M. Océanogr.,
1963 (série Nosy Bé II), no. 6, pp. 131-212.
Humes, A. G. anv J.-S. Ho. 1966. New lichomolgid copepods
(Cyclopoida) from zoanthid coelenterates in Madagascar.
Cahiers O.R.S.T.O.M. Océanogr., 4(2): 3-47.
1967a. New cyclopoid copepods associated with the coral
Psammocora contigua (Esper) in Madagascar. Proc. U. S.
Nat. Mus., 122(3586): 1-32.
1967b. Two new species of Lichomolgus (Copepoda,
Cyclopoida) from an actiniarian in Madagascar. Cahiers
O.R.S.T.O.M. Océanogr., 5(1): 3-21.
1967c. New cyclopoid copepods associated with the al-
cyonarian coral Tubipora musica (L.) in Madagascar. Proc.
U. S. Nat. Mus., 121(3573); 1-24.
1968a. Lichomolgid copepods (Cyclopoida) associated with
corals in Madagascar. Bull. Mus. Comp. Zool., Harvard
Univ., 136(10): 353-413.
1968b. Cyclopoid copepods of the genus Lichomolgus
associated with octocorals of the family Nephtheidae in
Madagascar. Proc. U. S. Nat. Mus., 125(3661): 1-41.
1968c. Cyclopoid copepods of the genus Lichomolgus as-
sociated with octocorals of the family Alcyoniidae in Mada-
gascar. Proc. Biol. Soc. Wash., vol. 81: 635-692.
KossMANN, R. 1877. Entomostraca (1. Theil:Lichomolgidae). In
Zool. Ergeb. Reise Kiistengeb. Rothen Meeres, erste Halfte,
IV, pp. 1-24.
Scotr, A. 1909. The Copepoda of the Siboga Expedition. Part 1.
Free-swimming, littoral and semi-parasitic Copepoda. Siboga
Exped., 29a: 1-323.
SrEweELt, R. B. S. 1949. The littoral and semi-parasitic Cyclopoida,
the Monstrilloida and Notodelphyoida. No. 2 of vol. 9 in
John Murray Expedition 1933-34, Scientific Reports, pp.
17-199.
Stock, J. H. 1959. Copepoda associated with Neapolitan inverte-
brates. Pubbl. Staz. Zool. Napoli, 31/1: 59-75.
1962. Lichomolgus pterophilus n. sp., a cyclopoid copepod
associated with the East Indian sea-pen Pteroeides.
Beaufortia, 9(105): 155-163.
Stock, J. H. anp G. Kuereron. 1963. Copépodes associés aux in-
Copepods Associated with Octocorals 749
vertébrés des cdtes du Rousillon. 2.-Lichomolgidae ecto-
associés d’octocoralliaires. Vie et Milieu, 14(2): 245-261.
Tuompson, I. C. anp A. Scotr. 1903. Report on the Copepoda col-
lected by Professor Herdman, at Ceylon, in 1902. Rept.
Gov. Ceylon Pearl Oyster Fish. Gulf of Manaar, suppl. rept.,
no. 7, pp. 227-307.
THoRELL, T. 1859. Till kamnedomen om vissa parasitiskt lefvande
Entomostraceer. Ofversigt K. Vetensk.-Akad. Foérhandl.,
Arg. 16, no. 8, pp. 335-362.
TrxmeR-DurivauLt, A. 1966. Octocoralliaires. Faune de Madagascar,
21: 1-456.
Ummerkutty, A. N. P. 1962. Studies on Indian copepods 5. On
eleven new species of marine cyclopoid copepods from the
south-east coast of India. J. Mar. Biol. Ass. India, 1961,
3(1 and 2): 19-69.
VERSEVELDT, J. 1968. Preliminary note on some new Octocorallia from
Madagascar. Koninkl. Nederl. Akad. Wetensch.-Amsterdam,
Proc., ser. C, 71(1): 52-59.
750 Proceedings of the Biological Society of Washington
>
Vol. 81, pp. 751-754 30 December 1968
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
A NEW SPECIES OF LACCOBIUS FROM THE GREATER
ANTILLES (COLEOPTERA: HYDROPHILIDAE )
By Paut J. SPANGLER!
Smithsonian Institution, Washington, D.C.
The genus Laccobius, in the Western Hemisphere, occurs
from southeastern Alaska and Canada through the United
States and into Mexico. Most of the described species are
found in the United States but an apparent hiatus occurs
along the Gulf Coast states from Louisiana to Florida. In his
revision of the American species of the genus Laccobius,
D’Orchymont (1942) stated that although the means of disper-
sion are great enough the genus does not appear to be repre-
sented from the vast region of the southeastern United States
or from the Antilles. He emphasized the fact that the genus
was absent from Florida, a state whose water beetle fauna
had been well explored and reported upon by Leng and
Mutchler (1918) and Blatchley (1919). However, examina-
tion of older collections not seen by D’Orchymont and recent
surveys of aquatic beetles made in the Greater Antilles
revealed the presence of a single new species of Laccobius
on Puerto Rico, Jamaica, Haiti, the Dominican Republic and
Cuba.
Present evidence suggests that this new species reached the
Greater Antilles from Mexico or Central America where a
number of species of Laccobius are known to occur. This
assumption is supported by the fact that Laccobius is still
rare from Gulf Coast states and has not been found in Florida
even after the recent and very thorough study of the Floridian
water beetles by Young (1954). The interesting new species
from the Greater Antilles is described below.
1 This study was made possible in part by grant GB-1697 from the National
Science Foundation and by grant 465 from the American Philosophical Society.
61—Proc. Brot. Soc. WasH., Vou. 81, 1968 (751)
752 Proceedings of the Biological Society of Washington
Fics. 1-2. Laccobius antillensis new species, holotype, male genitalia:
(1) ventral view, (2) dorsal view.
Laecobius antillensis new species
Figures 1 and 2
Length of holotype male 2.75 mm.; greatest width 1.4 mm. Color of
head piceous except iateral testaceous spot anterior to each eye;
pronotum with piceous disc but broadly testaceous laterally; scutellum
piceous; elytron testaceous except humeral macula, two maculae at
midlength near suture, one macula near lateral margin and almost all
punctures of serial rows of punctures dark brown. Venter dark brown
except hypopleura and epipleura testaceous. Labial palpus light
testaceous; maxillary palpus light testaceous with apices brownish;
antenna light testaceous except last three segments of club darker brown.
Leg testaceous except coxa, trochanter and basal two-thirds of femur
brown.
Head distinctly microreticulate, moderately coarsely and moderately
densely punctate, more densely so adjacent to eyes, punctures on frons
separated by two or three times their width; labrum very feebly
microreticulate, densely punctate with punctures smaller than those of
head, feebly emarginate medially. Pronotum slightly more than twice
as broad as long; distinctly microreticulate; moderately coarsely and
moderately densely punctate; punctures slightly larger than those of head
and those on disc separated by one to three times their width; finely
margined laterally; submarginal row of punctures across base. Elytron
without surface sculpture but with 10 serial rows of coarse punctures and
a few coarser punctures scattered between rows; serial punctures slightly
larger than those on head; finely margined laterally. Scutellum sparsely
punctate, punctures smaller than those of pronotum or elytra. Front
leg with coxa, trochanter and basal third of femur covered with
New Species of Laccobius 753
hydrofuge pubescence. Fore tarsus consisting of four segments; first
segment expanded apically; second segment darker in color, expanded,
with pubescent pad beneath. Middle leg with only coxa and trochanter
pubescent; femur with few, small, seta-bearing punctures. Hind leg
with coxa pubescent; femur impunctate. Venter with hydrofuge pubes-
cence except posterior half of first and entire second abdominal sternum
smooth. Prosternum longitudinally carinate. Mesosternum strongly pro-
tuberant medially, protuberance with strong anterior tooth and carinate
posteriorly. Posterior half of first and entire second abdominal sternum
shallowly concave; anterior and posterior edges of concavity lined with
a sparse row of long, fine hairs; hyaline substance in concavity similar
to that found in Chaetarthria.
Aedeagus (fig. 1 & 2) as illustrated.
Female: Similar to male but tarsus of fore leg normal, five segmented,
not broadened, without pubescent pad beneath.
Variations: The two maculae on each elytron, near the suture, and
the lateral macula vary in size and degree of distinctness. In a few
specimens, the anteriormost and lateral maculae are practically absent
and the posteriormost macula is greatly reduced.
Holotype: Male, Puerto Rico, Juncos, Rio Gurabo, XII-30-1962,
Paul J. Spangler, USNM type No. 69502, deposited in the U. S. National
Museum.
Allotype: Same locality and data as holotype.
Paratypes: Same locality and data as types, 21¢ 6, 272 9; PUERTO
RICO: nr. Buenos Aguas, Rt. 56, Km. 15.5, Rio de Bayamon, VIII-19-
1961, 16, 12, Flint and Spangler; Comerio, Rt. 156, Rio de la Plata,
VIII-15-1961, 2¢ 6, 19, Flint and Spangler; nr. San Lorenzo, Rt. 181,
Km. 16, VIII-21-1961, 106 ¢, 799, Flint and Spangler; nr. San
Lorenzo, Rt. 181, Km. 9.1, VIII-21-1961, 7466, 499, Flint and
Spangler; Toro Negro, Rt. 149, Km. 41.8, VIII-24-1961, 16¢ 6, 169 9,
Flint and Spangler; Toro Negro, Rt. 149, Km. 40.7, VIII-24-1961,
1¢, Flint and Spangler; Bayamon, Rt. 174, Km. 13.5, VIII-19-1961,
964, 7929, Flint and Spangler; Bayamon, Rt. 174, Km. 8.7, VIII-19-
1961, 3546, 2699, Flint and Spangler; nr. San German, Rio
Guanajibo, VIII-14-1961, 244 6, 249 9, Flint and Spangler; nr. Paso
Seco Jct., Hwy. 31, Km. 15.4, 1-8-1963, 1¢, 19, roadside stream, 400’
elev., Paul J. Spangler; Magueyes, I-1-1963, 19, Paul J. Spangler.
JAMAICA: Gordontown, II-4-1937, 19, Sta. 384, Chapin and Black-
welder; St. Catharine Parish, Hyde nr. Bog Walk, VII-23-1962, 8é ¢,
829, T. Farr, O. and R. Flint; Whitfield Hall, Blue Mts., nr. 4500 ft.,
VIII-13-20-1934, 399, Darlington (MCZ). HAITI: P. au Prince,
1934, 16, Darlington (MCZ); Ennery, nr. 1000 ft., IX-6-11-1934, 19,
Darlington (MCZ); La Hotte, N.E. foothills, 3-4000 ft., X-12-1934, 19,
Darlington (MCZ). DOMINICAN REPUBLIC: 25 Km. S. Pto. Plata,
VI-1938, 299, Darlington (MCZ). CUBA: Soledad, Cienfuegos,
X-17-1926, 14, Darlington (MCZ); Soledad, Cienfuegos, XII-2-1926,
754 Proceedings of the Biological Society of Washington
Arimao R., 19, Darlington (MCZ); Soledad, Cienfuegos, X-19-1926,
Arimao R., 192, Darlington (MCZ); Soledad, Cienfuegos, VIII-12-1934,
Arimao R., 1¢, Darlington (MCZ).
Comparative notes: This species runs to couplet 5 in D’Orchymont’s
(1942) key, but it possesses some of the characters given in both
rubrics of the couplet. It resembles agilis (Randall) by the “punctation
of the elytra less irregular, more or less chromatically aligned in fine
brown lines,” but differs from it by its smaller size and more elliptical
shape. From agilis and the three species that key through rubric 5’,
ellipticus Le Conte, mexicanus Orch. and californicus Orch., this new
species may be distinguished by the median lobe of the aedeagus which
is very broad (fig. 1 & 2) from base to apex instead of being slender.
I am grateful to Dr. P. J. Darlington, Jr., for the opportunity to
study the Laccobius he collected in the West Indies. These paratypic
specimens are indicated by the abbreviation (MCZ) and are in the
Museum of Comparative Zoology, Cambridge, Massachusetts. All other
paratypic specimens are from the collections of the U. S. National
Museum.
Paratypes will be deposited in the California Academy of Sciences,
British Museum (Natural History), Canadian National Collection,
Museum Nationale de Histoire Naturelle de France, Institut Royal des -
Sciences Naturelle de Belgique, Agricultural Experiment Station, Uni-
versity of Puerto Rico, and in the collections of Drs. R. Mouchamps
and F. N. Young.
LITERATURE CITED
BLATCHLEY, Wituis S. 1919. Insects of Florida Va. Supplementary
Notes on the Water Beetles. Bull. American Mus. Nat.
Hist. 41(4): 305-322.
LENG, CHARLES W., AND ANDREW J. MutTcHLER. 1918. Insects of
Florida V. The Water Beetles. Bull. American Mus. Nat.
Hist. 38: 73-116.
D’OrcHYMonT, ARMAND. 1942. Revision des Laccobius Américains
(Coleoptera Hydrophilinae Hydrobiini). Bull. Mus. Roy.
Hist. nat. Belgique 18(30): 1-18.
Younc, Frank N. 1954. The Water Beetles of Florida. Univ. Florida
Stud., Biol. Sci. Ser. 5(1): ix + 238.
Vol. 81, pp. 755-760 30 December 1968
PROCEEDINGS
OF THE
BIOLOGICAL SOCIETY OF WASHINGTON
A NEW SPECIES OF AQUATIC ISOPOD CRUSTACEAN
(GENUS ASELLUS) FROM THE PUEBLA PLATEAU,
CENTRAL MEXICO
By GerALD A. CoLe anp W. L. MINCKLEY
Department of Zoology, Arizona State University, Tempe
On 18 December 1966, Robert R. Miller and Minckley
briefly visited the Rio Cosala at the edge of the village of San
Martin Texmelucan, Puebla, México. A small collection of
invertebrates included the new asellid isopod described be-
low, and extends the known world distribution of the genus
Asellus, formerly given as no farther south than 30° latitude
(Birstein, 1964), to just below 20° latitude.
The Rio Cosala drains from the eastern slopes of the volcano
Iztaccihuatl and enters the Rio Atoyac just downstream from
San Martin Texmelucan. The larger stream then passes
through the portion of the Puebla Plateau lying between
Iztaccithuatl and Popocatépetl on the west and Volcan La
Malinche on the east, then bends south and west to flow
into the Rio Balsas, and thence to the Pacific Ocean. At 2,300
meters above mean sea level, where the collection was made,
the Rio Cosala was clear and cool and was flowing about 0.33
cubic meters per second. The banks were gently sloping and
grassy, and the stream was lined by a narrow rim of emergent
aquatic vegetation (Nasturtium, Ludwigia, and sedges). Bot-
toms were of sand and mud in pools and gravel on riffles,
with substantial deposits of detritus (twigs, leaves, etc.)
present in all areas. The isopods were sieved most abundantly
from the aquatic plants, but also were present in debris on,
and lateral to, swifter water. Other abundant animals were
insects (Hemiptera, Ephemeroptera, and Trichoptera) and a
talitrid amphipod, genus Hyalella. The area was locally
¢
62—Proc. Biou. Soc. WaAsH., Vou. 81, 1968 (755)
756 Proceedings of the Biological Society of Washington
Ficures 1-21. Asellus puebla, new species. 1. Holotype, 6.9 mm.
2. Maxilliped, allotype, 6.5 mm. 3. First pereopod, male, 8.6 mm.
4. First pereopod, propodus and dactylus, male, 8.6 mm. 5. First
pereopod, allotype. 6. First pereopod, propodus and dactylus, allotype.
7. Second pereopod, male, 8.6 mm. 8. Fourth pereopod, lateral aspect,
male, 8.6 mm. 9. Fourth pereopod, allotype. 10. Fourth pereopod,
merus, median aspect, male, 8.6 mm. 11. Seventh pereopod, male, 8.6
mm. 12. Second pereopod, dactylus, male, 10.3 mm. 13. First pleopod,
New Mexican Isopod 757
disturbed by persons doing laundry and children wading, but
was otherwise unmodified.
Travel to México was supported, in part, by NSF Grants
GB-2461 (to Minckley) and GB-4854X (to R. R. Miller), and
preparation of the figures for this manuscript was funded by
NSF Grant GB-6477X (to Minckley).
Asellus puebla new species
Figs. 1-21
Material: All specimens were obtained 18 December 1966 from the
Rio Cosala, west edge of San Martin Texmelucan, Puebla, México, by
R. R. Miller and W. L. Minckley. The holotype, a 6.9-mm male, an
allotype, an ovigerous female 6.5 mm long, and one lot of paratypes
are deposited at the U. S. National Museum. Other lots of paratypes
are at the Nacional Coleccién de México, Mexico, D. F., and at the
National Museum of Canada, Ottawa.
Description: A small species of Asellus, largest male 10.3 mm long,
largest ovigerous female 6.5 mm long (allotype). Body (Fig. 1) from
2.6 to 2.8 x width. Telson length subequal to width. Eye small, longer
than broad.
Antenna 1: 10 to 11 flagellar segments, reaching to distal border
of peduncle of antenna 2 when reflected.
Antenna 2: flagellum with about 72 segments, reaching to 7th
pereonite when reflected.
Mandible: right incisor with 4 teeth, spine row with 5 to 6 dentate
spines distally and 11 to 12 plumose spines proximally; left mandible
with 4-toothed incisor and lacinia of 4 teeth, spine row of 12 to 14
stout, plumose setae. Palp segment 1, narrowest basally, with 6 inner,
marginal setae, and one distal, facial seta; palp segment 2, setae on
distal half of inner margin in 2 ranks, about 10 marginal and 10 sub-
marginal setae, proximal 3 setae smooth, others with short plumosities;
palp segment 3, inner margin with about 18 pectinate spines.
Maxilla 1: apex of outer plate with 11 spines, all denticulate except
2 outer and 1 innermost, and 2 smooth, subapical, facial setae; inner
plate, apex with 2 robust, circumplumose setae, with denticulate apices,
and 3 slender, plumose setae.
Maxilla 2 has no distinctive features.
male, 8.8 mm. 14. Second pleopod, anterior aspect, male, 10.3 mm.
15. Second pleopod, endopod apex, anterior aspect, male, 8.8 mm.
16. Second pleopod, endopod apex, posterior aspect, male, 8.8 mm.
17. Second pleopod, allotype. 18. Third pleopod, male, 8.6 mm.
19. Fourth pleopod, male, 8.6 mm. 20. Fifth pleopod, male, 8.6 mm.
21. Uropod, male, 8.6 mm.
758 Proceedings of the Biological Society of Washington
Maxilliped: inner plate apex densely setose, with 4 to 6 coupling
spines in males; oostegite in ovigerous females with 18 apical setae,
each distally plumose (Fig. 2). Palp, inner margins of segments 2 to 5
extremely setose; segment 1, with 2 to 4 outer setae and 1 inner seta;
segment 2, with 3 outer setae and one at distal corner; segment 3,
with 2 outer setae and one at distal corner.
Pereopod 1 (Figs. 3-6): palmar margins of male propodus with 2 to
3 proximal, robust spines, directed obliquely posteriad, one large, acute
process near middle, and 1 short, blunt process just distad. Propodus
of female with 2 sharp spines at proximal corner of palmar margin and
only the short, blunt process at mid-palmar surface. Posterior margin
of dactylus with 10 to 15 teeth.
Pereopods 2-7 (Figs. 7-12): dactyls bearing 3, rarely 4, spines.
Anterior margin of propodus 5-7 armed with large spine inserted about
midway. Pereopod 4, shortest, much stouter in male than in female,
merus with row of posterior spines on inner surface in males.
Pleopod 1 (Fig. 13): basal segment with 6 to 8 coupling spines;
distal segment slightly larger than basal, with concave outer surface
and broadly rounded, truncate, distal margin; inner surface smooth;
distal portion of outer surface and apex with 16 to 20 smooth, relatively
short, setae.
Pleopod 2 (Figs. 14-17): protopod of male 1.3 x longer than wide,
longer than exopod; endopod reaches slightly past midlength of exopod;
protopod with 2 medio-distal, smooth setae. Endopod narrow, almost
straight but curving gently laterad; lateral apophysis prominent, no
medial apophysis except a postero-basal, cuplike expansion. Tip of
endopod, median process curving laterad, overriding lateral process;
cannula protruding distally and pointing slightly mesad, no caudal
process. Exopod, proximal segment with 3 lateral setae; distal segment
with 16 to 19 plumose setae. Pleopod 2 of female with lateral sinuosity
in exopod, appearing somewhat bilobed with a straight inner margin;
outer and apical margins with 12 plumose setae; inner margin with 2
short setae on distal third.
Pleopod 3 (Fig. 18): exopod with about 16 terminal, plumose setae,
outer margin with about 16 smooth setae and 2 or 3 distal, plumose
setae.
Pleopods 4-5 (Figs. 19-20): with a few smooth setae on basal,
outer margins; endopods unarmed in pleopods 3-5.
Uropod (Fig. 21): protopod and endopod subequal; exopod 0.75 x
length of endopod in larger specimens, rami more nearly subequal in
small specimens. Protopod narrowest basally; rami tapered.
Color in life gray, mottled, with lighter underparts. Eye black,
reflecting violet in sunlight.
Etymology: The name “puebla’ is for the intermontane plateau from
which the new species is known.
New Mexican Isopod 759
Discussion
The relationships of Asellus puebla to other described species of
epigean Asellus are not apparent. Subgenera of this group have recently
been criticized (Chappuis, 1953, 1955; Bowman, 1967), and may be,
for the most part, invalid. However, if they are utilized, the new species
is related to the Eastern American group of species (subgenus Conasellus
Stammer). Only one other epigean form is described from the Pacific
drainage of North America, A. tomalensis Harford, and it may belong
to the subgenus Mesoasellus Birstein of Western America and Eastern
Asia. The presence of A. puebla in the westward-draining Rio Balsas
obviously reflects dispersal from the east.
Four of the described epigean Asellus from Eastern North America
resemble A. puebla in one or more respects. Asellus brevicauda
(Forbes), A. dentadactylus Mackin and Hubricht, A. kenki Bowman, and
A. oculata (Mackin and Hubricht) have at least slightly concave lateral
margins of the distal segment of the first pleopod of the male. The
first three species differ from A. puebla in having the distal margin of
that segment armed with long, plumose setae rather than with relatively
short, smooth setae. Asellus oculata differs markedly from A. puebla
(and the others) in details of the male second pleopod and _ the
dactylus of the first pereopod.
Asellus puebla appears elongate and relatively narrow in life, reminis-
cent of some troglobitic species of the genus. Bowman (1967) reviewed
relative proportions and ratios of body length : width, and telson length :
width, in many epigean and hypogean species of Asellus. Asellus kenki,
a form living in the sources of springs, was intermediate between the
surface and subterranean forms. Asellus puebla is obviously an epigean
form based on these criteria. The eye of A. puebla is, however, only
slightly larger than that of A. kenki (see Bowman, 1967, Fig. 1).
LITERATURE CITED
Bmstet, J. A. 1964. Freshwater isopods (Asellota). Fauna of U. S.
S. R., Crustacea 7: 1-148 (Original published 1951, Zool.
Inst. Acad. Sci. SSSR 47: 1-140; Israel Progr. Sci. Transl.,
1964).
Bowman, T. E. 1967. Asellus kenki, a new isopod crustacean from
springs in the eastern United States. Proc. Biol. Soc. Wash.
80: 131-140.
Cuappuis, P. A. 1953. Sur la systematique du genre Asellus. Notes
Biospéol., 8: 67-79.
1955. Remarques générales sur le genre Asellus et descrip-
tion de quatre espéces nouvelles. ibid. 10: 163-182.
760 Proceedings of the Biological Society of Washington
*
INDEX
VOLUME 81
Se ee
(New names in boldface; see p. v for list of authors)
A (Parvanachis) rhodae _ 144, 147
abberans, Nephthya ___..-_.___-_______- 739 SParsay eo slus wakes Beate A 148
abyssicola, Bathyteuthis _ _ 171 (Suturoglypta) 145
acanthi, Squalonchocotyle _ 363 (Suturoglypta) albella ___ 145
acanthias, Squalus _________ 363 ae iontha ___ 145
Acanthocarpus _ (Suturoglypta) pretrii ___ 145
Acanthosquilla multifasciata 7 Anahita punctulatus —___________ 207
acaulis, Stereonephthya ______ 719,740 Amamygdon —______ 491
AGG SeilyaeT — 199 SOlOMONIS ee 492
acerbum, Habrocestum _______..___- 208 Ancherythroculter _______________________. 343
Achaearanea tepidariorum __._. 206 brevianalis ________________ 345
acrochordus, Platymantis kurematsui 343
Acropora) eo eee nigrocauda, === 343
cytheria _____ wangi ___-------- Be AS
palifera _.. andrewsi, Porites _____________------------__. 738
scherzeriana angulataseeayonay oan 737
actinophorus, Lichomolgus __ 737,743 Amkyrodrilus —_______-_________ 309
INS Tay ap fee a ce ele ee ea oe 179 ~«=©Anolis —-___. eS ee ed 133
SCAT OLA pee 179 bartschi -_---- 133
ATS |, oe RS 179 (lich 133
aculeata, Euconchoecia -—-......- 543 anomalus, Lichomolgus __------------- 679
aculeatus, one _.. 736, 739, one Anoural Sie ae ee 427
aculeodactylus, Platymantis _.....- 77 brevirostrum _________---_-_-__-__- 427
acuminatus, Cambarus __--.........- 272 Caudifert sees eee 428
acuta, Xystocheir CSE ean Par 521 cultrata __ _ 428
acuticeps, Parafelicola __..........- 109 geoffroyi _-.-_._--- _ 428
acutipinna, Triakis -.______________- _ 614 geoffroyi lasiopyga 429
acutissimus, Procambarus -_.._.._.--_- 20 antarctica, Erpocotyle _- 359
acutitelson, Dynamenella —__.-----.---- 591 Macrophylla _____-
acutus, Procambarus ____.._________-____- A417 Macrophyllida __. 2
adelphus, Lichomolgus ___ 650, 707, 747 Mustelusiy = ee
adversus, Myotis __......---_--- 492, Squalonchocotyle
(Aedicira ), Aricidea ________ antarcticus, Mustelus ________
aegyptius, Lichomolgus __- Antheliay=
africana, Lemnalia ____._._.. os GRECTNG
africanus, Parafelicola antillensis, Dipsacaster
agassizii, Dicromita _............ 440, 539 antillensis, Laccobius ------.
agilis, Laccobius —-_-.------__------____- 754 antiquorum, Petenobolus _-
Agroeca minuta 207 =antonii, Dynamenella -__-_-----
akarithymus, Platymantis __ 70, 74, 76 Antrodiaetus
akaroensis, Hexabothrium 357. Anurida maritima —______.
alarius, Phrurotimpus Gees eee iF 207. Anyphaena pectorosa -_.
alba, Exosphaeroma -__..._--_---- GO0hObata00 se
albella, Anachis ____-_____---__----_--___- 145 appendiculatum, Hexabothrium
albidum, Theridion __.......___. GYiYs AARNE
PAU erm Cet ees ees TS, 101 arapaho, Harpactostigma
monticola ______ arborea, Telesto —__----------------
parbatae cn cie sai hee 0 arboreum, Litophyton —___------------------ 739
Allomegalocotyla __._....__. Archaeperidinium minutum _----.--- 95
Allosprostonia __.__ Archiconchoecia ______------------------------ 540
tauvinae __ striata _____.. 543
alla, Iskyoeogy Arctostaphylos == Ee 526
slticorpus Microphysogobio brevi- arcuatipes, Lichomolgus _____- 738, 743
OSEVIS DV aah ea ere EE 341 arenae, Streptosyllis _____----------------- 153
aluouspis' Palaechthon __.._________ 632), sargenteus) Pampus 173
AVEO pOlapi san tin oe PS 738 ‘ Stromateoides -_--------------—- 173
amabilis, Lemnalia ___________ 749 ~~ +Ariadna bicolor —-.----------------------- 206
(Amblygaster), Sardinella 214 Aricidea ——____-___-_---------------------- 323
americana, Atheyella = 574 (Nedicira) ee 323
Platydactylus 123 (Cirrophorus) __-----------------—- 323
Tarentola - _ 123 jeffreysii —________________- 323
Amplocheir Daciticay {sie eee ee 325
Anachis DATTA OL US pests oe ae ee 499
3 armata, Paradoneis ______----------------- 324
(Parvanachis) __-------------- 145 arvense, Equisetum __-----------------. 199
(Parvanachis) isabellei ____- 147 asaphidis, Lichomolgus __..... 739, 744
(Parvanachis) obesa _-------- 147 = Asclepius erosa -.-.---------------------------- 437
(Parvanachis) ostreicola _. 147 (ASM iiss es soe en 755
61
<M 1 1HSCNV > S 7
S ly
\ JUN 1 4970 }
~
“LLB RARIES
a __— EEE
762 Proceedings of the Biological Society of Washington
brevicauda)) Sue 759
(Conasellus) ___- _. 759
dentadactylus __ 159
kenkiy sei ee 759
(Mesoasellus) __ 59)
Oculata= aaa S59
puebla _____ _ 757
tomalensis _________ 759
asperimanus, Cambarus _. - 580
assimilis, Turdus _ 38
IXSENGTG AGES a 583
atlantica, Conchoecia 468, 470
atroxse Bothrops) eee een 321
attenuata, Triakis __ . 614
Attheyella _._______. 571
americana, (else Datewise 574
carolinensis ____ 571, 576
Tin O1sensis) yee 574
obatogamensis __.----- 574
pilosay ea ee ee ee 571, 572
audens, Lichomolgus __.-
australis, Coelorhynchus
Eurysorchis ____ 355
Galeorhinus _ Biz
Austrosquilla _________. . 241
malayensis ___ _ 241
osculans _________ 243
vercoi ____ _ 243
Auturus ____-_____ A479
evides|i iio a ae 483
B
bacidifera, Bathyteuthis _.._ 163
bairdii, Cycloes
bakkamoena, Otus ___
balani, Striella 598, 599
balguerii, Mursia _.......-.- 609
barbata, Algansea 106
barbouriiriakis) = aa 614
bartonii, Cambarus _____ 13, 262, 580
IbaxtschisAm olis eee eee 133
Bassozetus normalis __ 539, 561
Bathyconchoecia _____________ _.. 540, 541
darcythompsoni ___________________ 541
foveolata _.. 542, 565
galerita ___ BY Dz/
kornickeri __ 543, 555
NACUMOS Ae eset AL Len eer ee 541
laqueata = en aes 545, 551
paulula (2.2 ei 541, 547
Sagittarius _______________ 542, 561
Bathyteutiii sien eee eens 161
abyssicola __ 2) 1e/ik
bacidifera __ 163
berryi _______ _ 169
MELal Op shee Le eel eee 163
Bathyvargula _______ _.. 442, 540
optilus 439, 442
PaLviSpinos aye 448
walfordi _______ 448
Bdellodrilus ________ 309
beattyi, Illiniurus _____ 480
bernacchii, Trematomus _. 406
berryi, Bathyteuthis __________________ 169
Betulavlentay iwi hale 199
beyeriyHaxonellay 413
biarticulata, Protodorvillea ___._______ 226
loyioolkars, ANmtevoboyey 206
bidentata, Streptosyllis __--____________ 153
bimaculata, Chone 227
bleekeri, Hemiculter 343
bombayensis, Clorida __.._.-_-.-_____- 249
borchgrevinki, Trematomus — 410
Bothrops) 3232 ae 319
atroOxe sa trOXxG ee eee 321
lansbergii rozei 320
lansbergii venezuelensis ____. 319
neuvoiedii venezuelenzi __ 321
pifanol | Seana ees
venezuelae ________
venezuelensis
boulengeri, Cormufer
Platymantis _____ 69, 70, 71, 3
bouvieri, Linckia ____.__-___----_-_-
boylii, Peromyscus ees La Ue
Bradycinetus ctenorhynchus _
brama, Seriolella ___.____________.
Branchiobdella ____-__--___-_____
brevianalis, Ancherythroculter ______ 345
brevicauda, Asellus ______________...__. 759
brevifurcatus, Lichomolgus __________ 668
brevirostris, Microphysogobio __-___- 340
Pseudogobio _______-___----__-__- 340
brevirostrum, Anoura ___. 427
brevispinis, Microgobius _____ OD,
iskovecorbaysron _. VA43
laevigatum __ _. 143
obesum ______ _.. 145
bullisi, Linckia _ <A)
burgersi- cay eae ee Prs)7/
Cc
Cachonina niei __..--- 92
cactus, Pavona _________. me /Si7/
caerulus, Cubiceps - BS lye
Calappai vate Ae Aue __. 605
californicus, Laccobius __— a, 4!
Calliscylium venustum _____- _.. 614
callorhynchi, Erpocotyle ___. _.. 360
Squalonchocotyle _.. 360
Callorhynchus _._--_--- _._. 360
capensis ______ _.. 361.
SOU bu Popeater GI _. 360
Cambarincola __ _.. 294
Cambarus = 11
acuminatus __ D2.
asperimanus ________________-_-______ 580
bartonii ___________ __ 13, 580
bartonii bartonii ---________ 262
bartonii striatus __. _.. 262
cristatus _. 11
dissitus ___ 14
distans ________ _.. 580
floridanus __ 12
fodiens _________ 14
friaufi __ ___ 268
hall Oe aa 269
hedgpethi _. 14, 20, 417
latimanus _________---__-___ 72, 296
Obey.ensis. 2. eee ea 12
parvoculus __ _. 268
pristinus _______ 12
prominens) 2 1
reduncus) a) es 12, 273
Tobustus) 22 eae any 27
sphenoides __ 262
Splcatus.) 22 eee ae O72:
striatus _____ _ 12, 262, 314
tenebrosus - 268, 314, aaa
valleculus) 2 ees
campanulipes, Scambicomus 190
camporum, Comanchelus ___________-- 503
campulus, Lichomologus ____- 738, 743
canadensis, Tsuga _____________-___-_--_- _ 199
canescens, Halaelurus 614
canicula, *Hexabothrium E359)
capensis, Callorhynchus _ 361
carinatus, Leiocephalus 29
carolinensis, Atheyella _______. 571, 576
carolinensis, Sathodrilus ______ 294, 296
carpenteri, Hylobates lar __-___ 625
Carpio Cy prin ste eennD 106
G@assidinidea,) 22-2. ae ee 599
Castiameira longipalpus _____________- 207
caudacutus, Chaetura _____.._.__.____- 87
Enrundapus) 89
Index
caudifer, Anoura ______ _.. 428
Glossophaga _ __. 429
Gentrelus ae ee ee ne 501
flavialis| 2222 sc ie ee 501
kerrensisy)) 2-4 o ee ee 501
centronotus, Trematomus __________ 406
G@eochasma y=) 8 eee 45
phrixinay) 23 eee 46
@ephalirusqe =) oe eee eee _ 614
@eraticelus rs 22 206
@eratodmlus) e223) eee 292
@etorbiniis pes es es Ss 613
Chaenobryttus gulosus _.._...______ 106
Chaetarthria) = SS eee 753
Chacturayseen hs oe ee ae 87
caudacutus caudacutus ___ 87
gigantea dubia —_..._________- 89
gigantea gigantea ____________ 89
gigantea indica —__..______ 89
Chamayiostomay eee 739
chamarum, Lichomolgus _____ 739, 744
Chanodichthys kuematsui _______ 343
Cheirauxus oe ie 521
Chelidonichthys kumu ___.....__- 373
chierchiae, Euconchoecia —__-_______ 543
Chiton stokesii ________________________ S aey/
chlondas,Clorida) 2 a= Ess 246
Ghloridella) 22 aes 241
Chloridella chlorida —___.__-________ 241
Chloropsis flavipennis _._..._.---_-- = 835
mindanensis _...._._----t—(—ti«'G
Ghone wei nls hoe es PPI/
bimaculata 227
CinCtasie a eae oe en ene e: 229)
filicaudata
paracinctay == se
TOSCA Weer ee ies ines ae
striata
SuSpectay= ar
teres
Chrysopteron
Chthalamusi-= ee aaa
Cichlasomat es is a ae
cincta, C@hone —______
cinctum, Peridinium
Cirripectes yess eee eee
Gizrophorus}2 2 eee _ 323
(Cirrophorus), Aricidea 323
Cladiella ____ eee ee
laciniosa _______
pachyclados
clavatus, Lichomolgus __ 730
Clavellodes intermedius
Clianella elegans __....-______
cliftoni, Nelsonia neotomodon
Cloridag eames Ee OAL
bombayensis __.-_.________-
chlorida _____________.
decorata -_______
depressay es i ee O4G
latispina) oo 247
latreillet (eee re 249
malaccensis 244
microphthalma __.--._-__ 246
Glubionaye aw es Seeaee 207
Clubionoides texana —_____________________
(Clupeonia), Sardinella
clypeata, Faxonella ____.____.________
Coclogorgial =a Sea 693
palmosa 22) St ee ie eae 730
coelorhynchi, Diclidophora ______ 355
Coelorhynchus australis _..___-__--_---—s« 855
colbecki, Notothenia ____________ 409
Colinuispimee sn Secale a eh eee 37
virginianus atriceps _._.__ 88
virginianus coyolcos _______ 38
virginianus harrisoni _______ 37
virginianus thayeri ________ 38
Columbella pretrii _..- Cid 455
Comanchelus —__________ _.. 503
camporum ___ _.. 503
hubrichti _____ e503
lobatus) jae seke eS moa 504
commodus, Lichomolgus _ 688, 741, eae
communis, Frontinella
compositus, Lichomolgus ____- 739, Fad
(Conasellus), Asellus _.._-_._-_-_--- ss 759
conchaphilassOsirea) 587
Conchoccia ae 440, 540
atlanticaye = eee 468, 470
mactocheiray =e 470
Vall divi a eis se OR eT 470
congestum, Moxostoma —__.--_- 106
conifera, Hatschekia = 173
conjunctus, Lichomolgus _ 736, 739, 744
contigua, Psammacora __..- 738
contrerasi, Procambarus 302
copleyi, Gunnellichthys as) _.. 60, 66
Gornuters 22 ee ee 69
boulengeri ________ a7 |
nova-britannea __ TNO A
@omus) florida 2 ae 199
corrugatus, Platymantis — 79
@ory.Caeus pea ea Mae eee Si 655
(Costoanachis), Anachis 144
crassa, Nephthya —________________.. 739
crassum, Lobophytum
crassus, Lichomolgus __
creaseri, Faxonella ____
cristata, Cycloes ________
cristatus, Cambarus __-
Mobbseus) se ee
cristatus, Lichomolgus — 644, 707, 742
crocatus, Microgobius __________ 191, 193
Cronodrilus)=3ese a ee 308
ogygius __- =, Silt)
cryptopalpa, cient ee 5S
@ryptosomag = ee __. 610
@ryptostomal 222. eee eee 610
ctenorhynchus, Bradycinetus __-___ 449
Tetragonadon __________ 441, 449
cubana, Tarentola 123
Cubiceps caerulus
cultrata, Anoura _-_______
cuneipes, Lichomolgus __ 717, 741, is
curiesus, Gunnellichthys — _________-
curtiramus, Lichomolgus __---_-___- 674
curtus, Microgobius ia oie Retell 192
cuspis, Lichomolgus ___________. 740, ae
bairdii __ __. 609
cristata _ _.. 609
deweti __ . 605
granulosa _______ _ 609
cyclolepis, Microgobius ——_---_-------_- 192
C€ymodocea oS 181, 702
@ypridinayehl ats eee ea 439
C@yprinusicarpio EEE 106
cytheria, Acropora —__----------------- 738
D
TB) Re CEOT Nastia Rea 155
COW es EON 158
POM Giese ote ee 158
ireqe(onleyeey a alas
schnittic 2) Ceca 155
danalPav Onaga anes 737
darcythompsoni, Bathyconchoecia _ 541
Euconchoecia -__---__----------_--- 540
dearborni, Rhigophila ________ 406
decoratase @lonid age ee 249
decorus, Lichomolgus 635, 736, 742, 747
Delochern ee ne ee 521
Deltatherium( = 629
deltoidea, Virguiera -_________________ 437
764 Proceedings of the Biological Society of Washington
Dendrocopos EE 1 Equisetum arvense ___________---... _ 199
TRUTEE LNT eee ee a 1 erectus, Microgobius —__..-...---___. 192
SCalarisy, toe Seon ea Nas il Ergaticus) (22 e 39
sealaris cactophilus _____ 1 ruber melanauris ___________ 39
scalaris eremicus __------------- 1 ruber rowleyi __-.-_-_----__- 39
scalaris lucasanus ____---_-____-- 1 ruber, tuber. 39
scalaris mojavensis ____________ 1 versicolor _________ ol) 40)
scalaris yumanensis -_--_--..--- 1 Enigone |) ee 206
Dendronephthya _.----_-_-_---_____- 740 erinaceus, Tetragonodon __. __.. 450
kollikeri oe ee a ee 740 ermsti, Hirundapus ____.____. 89
MUCTOMAtAy see 740 ~—erosa, Asclepius ___.__-____----__ 437
TERA few theo Ree ee #40) . ‘Erpocotyle aa a ee 359
stocki __ .... 740 antarcticay =a 359
STB) mara yur sy eee eee eee ES 87 callorbynchi ________-___________. 360
giganteus —_______----_____-_____-- 87 squalt =e eae 363
TA) OF pe ee Sf Exythroculter: 2 343
dented acts Asellus): 222-7 = 759 hypselonotus __________________- 343
dentata, Wamokia _______ 514, 522, 533 ilishaeformis _______.-___-_---_____. 343
dentipes, Lichomolgus 637, 707, 742, 746 mongolicus __._____.---_-_--------__ 343
depressa, Clorida _____-___-__-_--------_-- 249 wantl) 0 eee 343
deweti, Cycloes _________--------------------- 605 Eulalia (Pterocirrus) macroceros -. 223
dianae, Dynamenella _________---____-- 598 Eurelus soleatus
Diclidophora coelorhynchi ____---------- 355 Eurysorchis australis ___
Dicrolene intronigra ______.-_-_-_---- 440 unyarus eee
Dicromita agassizii __ _. 440, 539 leachii _____-
Didelphis obesula ________-----_--------_--- 251 evides, Auturus 483
ND YG GE die eR HC Se ee Reem La 599 _exilipes, Lichomolgus __ 725, 740, Tis
digitatus, Lichomolgus ____ _ 738, 743 Fixosphaeromay = 599
Dipsacasters ee ee meee 231 albag222. 8a.) a eae B00
antillensis ______---_--__--_-___-_- 233
farquharsoni ____._____-__-____- 237 F
grandissimus 237 faleata, Wamokia _
pretiosus —___----------------------- 237 — Fallacirripectes
sladeni( 0 =e ee 231 minutus
discordis, Wamokia ____ 513, 522, 522 SOP ie Pe EEO
dispar, Hemiculter 343 farquharsoni, Dipsacaster
dissitus, Cambarus -__. fasciatus, Tomicod
distans, Cambarus F
divaricata, Nitrocrella 582 Be Eee cera
Dolomedes) clypeata _
Doridopsis ruber ereascril
Dorishmabila eS Moat lo
dorsalis, Platymantis _---._-- 77, 80 fehlmanni, Triakis
dowi, Daector —-_------------------------- 158 — femoratus, Gorytes
Drassyllus virginianus ___________- 207% fenneli” Anachis)
duplex} Zelotes)=— ee 207 ferox, Xysticus got
Dynamenella ___ 587, 589 filicaudata, Chone 229
acutitelson —__.----------------------- 591 fissisetiger, Lichomolgus _ 725, 740, ee
antonii __ 593 flatus, Cypridina woe Jenene So ee
dianae __- - 598 ibe, beam, | 730
eatoni | ______--------------------------- 597 — flavicornis, Gorytes ______ 434
JOSeD hie es 592 flavipennis, Chloropsis =a 35
perforata ___------------------ 589)2 florday\ Cornus) <a een 199
setosa ______________________________- 594 floridanus, Cambarus _____--__- 12
tropica a 591 fluvialis, Centrelus 501
fodiens, Cambarus ___.-_-_--_--_____ 14
, LOLIMOS A ee Ck 42
eatoni, Dynamenella __________ 597 forsteri, Latridopsis __...___________ 391
ehrenbergi, Sarcophyton -------- 680, 738 forsteri, Mediavagina __-_ 391
elegans, Clianella —________________ 593 foveolata, Bathyconchoecia __ 542, 565
Lemnalia _---—------------.-~ 740 foxi, Lichomolgus 655, 742, 747
Lichomolgus aa 679, 707 fraseri, Magnolia eee! Prd ie Liens) 199
eee 32 friaufi. Cambarus __ 268
6 RYSHCUS nnn anna 207 friaufi, Tettodrilus _. 312, 314
elisabethae, Heteroxenia _____ 694, 741 Tvs Gone
ellipticus, Laccobius —______-__ 754 fukiensis, Gobiobotia intermedia _ 339
Ellisodrilus _____-_____-_---------- 295,312 — fukiensis, Pseudogobio ______ 340
elongata, Tetragnatha ___________________- 207 fusigera, Sige 995
emblematicus, Microgobius -__.-------- 192 aR iOet: who we
Entobdella squamula _______----------- 391 G
entomacrod use 113 Galeorhinus australis ________._-____ 357
thalassint sie een 117 _~—s gailerita, Bathyconchoecia ___ 545, 557
Eoconchoecia _________-- __. 540 Galevs) 22 are ee 621
aculéata) ss eee 543 melastomus eee 622
Chierchiae: 2202. 543 poli eee 622
darcythompsoni a 540 geminus, Lichomolgus ________- 739, 744
lacunosal = ee 540 gemmatus, Lichomolgus ____. 743, T47
peira:rayiee 2 ee eal 207 gemmosum, Theridiosoma ___---- 207
Epinephelus tauvina __________---_______- Siian eGenetta ti Grin aye een 109
Index
gentilis, Lichomolgus ________ 718, 740, ioe
geoffroyi, Anoura _......-----
gerringi, Daector ___ = 158
gigantea, Chaetura -______- as toh?)
giganteum, Stoichactus _. _._ 740
giganteus, Dennyus ________ Sli,
giganteus, Hirundapus ___. mos; eK8)
gigas, Lichomolgus _________. 3.5)
Siphonarial ee 587
gilliardi, Platymantis _____________ Gils 7A
glabripes, Lichomolgus _ 707, 742, 747
glaucum, Sarcophyton 674, 7
globosa, Halocypris _____-
globosum, Sarcophyton
Glossonycteris lasiopyga -________-_____- 429
Glossophaga caudifer ____. _.. 429
Gobiobotia ___-_____________. _.. 335
ichangensis ___ _.. 336
intermedia = 336
intermedia fukiensis ______ 339
intermedia intermedia ______ 336
kiatingensis —__.-______---__--_-_____ 336
kolleri __________. __ 336
naktongensis ____ _.. 336
pappenheimi ____. _._ 336
Goniopora __.___..______. ==> 18%}
Goodea ____- _ 106
Gorytes _______.------____ ... 434
femoratus ___. _ 436
flavicomis _____ _.. 434
laminiferus ___- . 434
mirandus ______.- 435
ruficornis _______ _... 434
rufulicornis _________----_.--_____- 434
gracilis, Anthelia _____ OD 42,
IH oplisus\s 2 eee 432
Micrathena ___ . 206
Oryttus .. 432
Protodorvillea _______. == 226
grandissimus, Dipsacaster -___- =) DBY/
granulosa, Cycloes __.._...-.. . 609
guildingii, Linckia ________. WAT
gulosus, Chaenobryttus ___ 106
Gunnellichthys —_________-_---___-__--______-_- 53
copleyi -____. 60, 66
Curiosus() 22 aoe
irideus ______. 60, 66
monostigma oa
pleurotaenia _ 60, 66
viridescens _______-______________ 56, 61
H
habereri, Proscyllium _______....__...-- 614
Habrocestum —____________-____- _. 208
acerbum ___ . 208
pulex? 2 es .. 208
Habronattus _ 208
Halaelurus canescens __.- _ 614
halli, Cambarus _____.___. _ 269
Halocypris __________. ._ 540
globosa -_----_-----___.. . 543
hardwickii, Kerivoula ____......_.__. 494
Harengula vittata __..__ 209, 211
Harpactostigma ___________-_-_------ 431
arapaho __ 434
rutilum _______ . 434
unicinctus ___ - 436
velutinus __________. .. 431
harpagonea, Paraonis _ _ 323
Harpiosquilla _________--___--__-_----_---___-_- 247
harrisoni, Colinus virginianus ______- 37
hasoni, Trematomus __.......___- 406
Hatschekia _______-_--__-_-_. = alesse)
conifera _______. = 73
iridescens ____ _ 178
prionoti __-.-.. ._ 178
hayi, Procambarus __._---........-..-.--_- 20
hedgpethi, Cambarus _________ 14, 20, 417
Helicinay 2 2 a eS 46, 51
Helophora insignis —___.._.....-.---_---_- 206
Hemiculter; (ose eee 343
Dbleekerij 2S -e2 See hie ee 343
Gispanh 22 Sioa eee inka s 343
kremp fi. ee 343
| Wie ee eee ns cule aes Oe ED 343
Hemitriakis leucoperiptera __ _.. 614
henlei, Triakis _.. 614
Herklotsichthys __. .... 209
Witte tee eee ee ie 038 217
hetaericus, Lichomolgus _ 663, 741, 746
Heteroxenia 693
elisabethae —_____--------.--_-_. 694, 741
Hexabothrium -~____--------_---_----___-------_- 57
akaroensis __________ 6
appendiculatum _.. 859
canicula ____________ ee OOD
mMUsteli pee Se .... 359
Hexabranchus orbicularis ____________ 741
hians, Lichomolgus ______ 719, 740, 746
hildebrandi, Nezumia ________. 440, 539
Hirundapus) se eee 87, 89
Caudacutiusiy ee 89
ermmstivy, Soon = hehe)
giganteus _ =4 GY)
Hobbseus ___________ cot el5
cristatusy) ea as 15
orconectoides ________________ 15, 17
prominens ____---____-------------_-_- 1
valleculus __ a all:
hobo, Trochopus _______-----------------___- 373
hoffmani, Wamokia ____ 513, 522, 528
hoffmanni, Procambarus ______.....-__- 305
Hoplisoides umbonatus -___.--- 436
Hoplisusiyos Se eee 432
gracilis ___ 432
velutinus -__-__----_-.. 436
hubrichti, Comanchelus __ 503
humeralis, Tomicodon -_____._.._.. 477
humesi, Sinularia __________. » 742
Hy alellay oe eee ae ee eee 755
Hybaphe -__-.-_- 513
Hybopsis alta __. 106
Hvlobatesi ae 625
lar carpenteri ___ 625
lar entelloides _____. _. 626
larg lary eens leet 626
lar moloch __.- 627
pileatus ____---.-___ 626
hylocetes, Peromyscus -- 99
hypselonotus, Erythroculter ___. 343
I
Ictalurusspricei= = 106
TWEXKOp ACA Sika OR wea neers ._ 199
ilishaeformis, Erythroculter __._. 343
Tiniurns) ees ee een ee 480
beattyi\ 20 ei _ 480
illinoisensis, Atheyella -___.___. 574
inaequalis, Lichomolgus ______- 740, 745
incisus, Lichomolgus ___ 680, 738, 743
indicus, Lichomolgus -_---__----_----_--_. 668
infans, Poeciliopsis -------- .. 106
infemalis, Oryttus _________. . 436
inflata, Pseudophilomedes -__.___--____. 461
inflatus, Paramekodon _... 441, 458, 461
inornatus, Rhabdornis -___--_---__---.--_- 35
insignis, Helophora -----------.----------.- 206
insolens, Lichomolgus -... 668, 738, 744
insularis, Phyllodactylus ____----_-__. 419
intermedia, Gobiobotia _____------------.- 336
intermedia, Gobiobotia intermedia _ 336
intermedius, Clavellodes
intersecta, Uvalida _______.
intestinalis, Lecanurius
intronigra, Dicrolene _____-
iontha, Anachis
766 Proceedings of the Biological Society of Washington
lostoma,, Chama EE 739
iridescens, Hatschekia ___ NZS}
QOstre ai SS ae ae aed 587
irideus, Gunnellichthys ________- 60, 66
irroratus, Liomys 99
isabellei, Anachis
SOO Oni yh ee eS ee
J
japonica, Stauronereis — 226
jaumei, Leiocephalus raviceps _-_ 24
jefireyis Pinus) eee See aE 538
jeffreysii, Aricidea ____ 828}
josephi, Dynamenella __ — bEY
jussievalsarainel ages 209
K
DE Fe Wg ae eR Rea ER
keeni, Myotis __
kefersteini, Protodorvillea_
kelloggii, ‘Quercus
kenki, Asellus ___—
Kerivoula: este oo. ee ee ee
hardwickii depressa _____-_- 494
kerrensis, Centrelus _--_-___________ 501
kiatingensis, Gobiobotia —_- ___. 336
kollens (Gobiobotay 336
kollikeri, Dendronephthya ___
koreensis, Microphysogobio ________ 340
kornickeri, Bathyconchoecia _ 543, 555
kossmannianus, Lecanurius _______ 181
krempfi, Cladiella —_-_________
Hemiculter __ ah
Ker thie feces = RI rea ase e
ee Chanodichthys
Kuhn a ae eee E
ean Chelidonichthys —______________ 373
kurematsui, Ancherythroculter -___. 343
L
Maccobiusyc2 2 aS Aa eee 751
agilise =a ee eee 754
antillensis _______________________- 752
californicus 254.
ellipticus - — They!
MVEXCAT Se 754
laciniosa, Cladiella _____ 635, 742
lacunosa, Bathyconchoecia _______ 541
Huconchoecia, as 540
laevigata, Rhombinella = 145
laevigatum, Buccinum —_.---__ 143
laminiferus, Gorytes —___.________ 434
Oryttus)\( See 434
anvaSSayhv CUS tape 226
lansbergii, Bothrops —_--_-____________ 319
lapazaes Oryttus| ee 434
laqueata, Bathyconchoecia __ 545, 551
larwHylobatess ae es
[easaedy saan ee
lasiopyga, Glossonycteris
latimanus, Cambarus —______-______
latipalpa, Streptosyllis ___
latispina, Clorida ________
latreillei, Clorida ___
Latridopsis forsteri ____-
Latrodectus mactans _
leachii, Euryurus ____
lebruinise Squalus =a
lecanora, Actinopyga —________ eee
lecanura, Miilleria —_._.____ 179
Tecanurius oe ei ae 179
intestinalis _________ 179, 187
kossmannianus ___________________
leibits My.Otis eee
Vespertilo _
Leiocephalus ________
carinatus __
Macropus w= ee
TaViCeps .2= 24s ee ee 23
raviceps jaumei 24
raviceps klinikowskii __.___ 23
raviceps raviceps ___________._ 23
raviceps uzzelli ________ = 2B)
SHICHigaster, 5 ane E47 /
Bemnaliqa yi ee Sas
africana yee
amabilis
elegans
flavanii aii aA
lenta, Betula —
Eepthyphantes\ =a _ 206
leptoclados, Sinularia ______ 644, 742
Leucauge venusta _.._-_ _ 207
Leuconoe moluccarum -_-_________. 495
leucoperiptera, Hemitriakis _______. 614
leucospilus, Otus __..._-- BET 1)
Levensenia __--_--__------__--__-_- _.. 323
leytensis, Micromacronus ___________ 33
Pichomoleusy ee ae eee 635
actinophorus ______-___-_-- 737, 743
aculeatus _________ 736, 739, 745
adelphus ___.____- 650, 707, 747
aefyptius) ee Gd 0
anomalus
arcuatipes ee 738, 743
asaphidis _______________ __ 739, 744
audens\—=-2— = aa ea 743, TAT
brevifurcatus —--.____--_ 668
campulus) 738, 743
chamaruny = eee 739, 744
clavatus _____-____ 730, 738, 744
commodus ___- es 688, 7AT, 745
compositus ____--________ 73 9, 744
conjunctus ____ _ 736, 739, 744
Crassus) eee. eee 739, 744
cristatus ________ 644, 707, 742
cuneipes ____ Bs 717, 7Al, 745
curtiramus) eae 674
CUSpIS) = eee 740, 745
decorus ______ 635, 736, 742, 747
dentipes ____ 637, 707, 742, 746
dicsitatus es 738, 743
elévansi(=) ee 679, 707
exilipes ______________ 725, 740, 745
fissisetiger __________ 725, 740, 745
foxa\t- tees ke 655, 742, 747
FEMINUS ye ee 739, 744
gemmatus __....._._ 743, 747
gentilis __.- 718, 740, ae
707, 742, a4
hetaericus _____ 663, 741, 746
hians 719, 740, 746
inaequalis _——___ 0, 745
incisus 680, 738. 743
INGICUS ea 668
insolens ___________ 668, 738, 744
lobophorus —__--------_____- 738, 743
longispinifer ___ 713, 741, 745
magnificus __________- 735, 742, TAT
oreastriphilus __________________-- 737
organicus ________- 736, 739, 744
patulus. = eee 741,
politus) =e 739, 744
prIolixipes 738, 743
protentus 635, 672, 707, 742, ae
pteroidis) = ee 736
pterophilus __
rhadinus ____________
TODUSEUS ee i eee _ 674
securiger __________ 702, 741, 746
sensilis; (== 741, 746
serratipes,..3.— ae ee 737
simulans ga 739, 744
singularipes _ 636, 685, 736, 741
spathophorus ____- 674, 739, 744
Index
spinipes _____________________- 655, 707
spinulifer ___ z
Squamicers =e
635, 689, 707, 742, 747
telestophilus _____ 725, 741, 746
triquetrus _______ 702, 742, 746
trispinosus
vagans ____________
varirostratus
venustus = ee
verseveldti
Lincki formosa ______
Binckia yes
bouvieri ________
ernithopus -___-
Kinyphia ase
maculata _____
marginata eae
Liogorytes unicinctus ____ —
Liriodendron tulipifera ___
liscia, Palythoa _________.
iui, Hemiculter _________ _. 343
lobatus, Comanchelus ___. 504
Teobieliaigpiaion echt. nol BUS 201
lobophorus, Lichomolgus _____.
Lobophytum crassum _______________ 668, 738
Lonchoglossa wiedi aequatoris ___ 429
longicornis, Otus __.......--_ 32
longipalpus, Castiameira _ Be 2.070
longiphallus, Sprostonia __ _ 377
longiphallus, Squatina _______________. 382
longisignalis, Uca ___________ QT, 279
longispinifer, Lichomolgus 713, 742, 745
lucifugus, Myotis ac a RRND ANB Ss Re 5
lucius, Anolis __________
Ludwigia ___-___-__-
Lycopodium
Lygodium —_...-___.
lyra, Paradoneis ___
Lysilla pacifica ___
Lysiosquilla spinosa
mabillaseD Oris eee 702, 741
macrocephala, Notothenia _____.________ A
macroceros, Eulalia _________
macrocheira, Conchoecia _
macrolepis, Psolus ________ _._ 347
Macrophylla ___---_--__- - 385
antarctica) 2 ee ees 387
Macrophyllida _ 371, 381, 385, 398, 405
antarctica’ “2 ee 38 itp 387
macrops, Platymantis ___________________
macropteri, Mediavagina _____ 395
macropterus, Nemadactylus ____- 395
macropus, Leiocephalus _____. 28
mactans, Latrodectus ___. _. 206
maculata, Linyphia __ 2206
Mangora ________ __ 207
Synapta ____ _. 190
faa kis) jee tie ie ee _. 614
maculosa, Thallassophryne __ _ 155
Maevia vittata -__-_.______________ __ 208
Magmatodnlus) ee 294
magnificus, Lichomolgus _ 735, 742, 747
Magnolia fraseri --__.______________________ 199
HALA OLS HL) CLITA VATS ee 87
malaccensis, Clorida —_.......-._.___- 244
malayensis, Austrosquilla ____________- 241
Manporayye => 2055 Dini eset ees 207
maculata __ _.. 207
placida (22a -. 207
maniculatus, Peromyscus -__.--_--------- 99
Mannobolus ____....-- 505
peninsularis — = 567
marginata, Linyphia _ 206
maritima, Anurida ____. _ 588
Marquesensis, Sardinella _________ 209
maturus, Paromomys —__--------- 632
maximum, Rhododendron _____--_--____ 199
mayanus, Petenobolus _--_ 509
Mediavagina ______--_-------_--.-
forsteri ____
macropteri
megadenus, Sathodrilus
Megalocotyle ____---_----__-_..
megaloplax, Psolus ______--__--__--__-__
megalops, Bathyteuthis
melanura, Sardinella ___._________
melastomus, Galeus
MeN ATGing Net ay pee eee eee 207
mendozae, Nothoprocta pentlandii _ 486
(Mesoasellus), Asellus
Meta menardi ee ee
mexicana, Neotoma ----------------------
mexicanus, Laccobius
meyersi, Platymantis
Micaria __-___-
Micrathena ___---
gracilis __
mitrata ___
Microgobius
brevispinis -___-
crocatus _____.
curtus
cyclolepis
emblematicus
tabogensis
Micromacronus leytensis sordidus _ 33
Microneta varia -__---------------------------
microphthalma, Clorida
Microphysogobio _______---------------------
brevirostris alticorpus ________
brevirostris brevirostris
brevirostris obtusirostris
KOreensis)) eae ae
obtusirostris __
Micropterus salmoides -_----------------
milbertii, Platydactylus ________------_----
miliaris, Actinopyga -_---------------------
milii, Callorhynchus -_--_---------.-----
mimicus, Platymantis ______-_- 71, 73, 74
mindanensis, Chloropsis -_----------------- 36
mind orensiss Otusy se 32
minuta, Agroeca ___.
Protodorvillea
minutum, Archaeperidinium ----.__- 95
minutus, Fallacirripectes ___---- 111, 113
miraflorensis, Microgobius ----------- 192
mirandus, Gorytes -_-------------------------- 435
Oryttus 435
mirus, Otus scops —--------------------------- 31
mississippiensis, Orconectes -------.---- 20
Mitchella repens ___------------------------—- 199
mitrata, Micrathena —-.-_.______ 206
moluccarum, meuconoen ae 495
My OtiSy se ee 492,
mongolicus, Erythroculter _--______--- 343
monostigma, Gunnellichthys __.-- 60
fomoyayesbayntcy, [Pb 207
monticola, Algansea ____------------------- 101
mordax, Stylophora ___-_-___-------------- 739
Moty xia pion) 220 ee 521
Moxostoma congestum __---_------------- 106
mucronata, Dendronephthya _ 740
Miilleria lecanura ___-___-__-------------- 179
multifasciata, Acanthosquilla ___-- 247
Mursiayibal ewe rite 609
MUSICaywhuUbIpOray ee ee 739
768 Proceedings of the Biological Society of Washington
musteli, Hexabothrium — 359 optilus, Bathyvargula __________ 439, 442
Mustelus 022 = orbicularis, Hexabranchus _________ aay
antarcticus __ Orconectes _____-___--------------
My otis) 22s eee mississippiensis ____---_--------_--
adversus mollucarum ____ 492, 495 Thoadeési" 22 =a ee ee
keen) 2 ae eee 258 rusticus mirus ____________________
leibii ciliolabrum ___________ 259 rusticus rusticus
leibii leibil) 2s ee 259 orconectoides, Hobbseus
leibii melanorhinus __---_-____- 259 _ oreastriphilus, Lichomolgus___________ 737
ere fcr 1s ee 258 organicus, Lichomolgus ___ 736, 739, 744
moluccannn = 492, omithopus, Lickia ______.____________. 41
VALI ANETISI S| 258 Ophidiaster _____________-________- 41
Onyttus ee ee ee 431
N gracilis arapaho __ _. 434
INS Cured tira ae no gracilis gracilis _. 432
nasuta, Perameles ______. infernalis noano-- -- 436
neapolitana, Paraonis laminiferus ---.- - 434
nelli, Neon ___.._______- lapazaes =. 2 ee eee 434
Nelsonian eer nee OB is, demi mirandus) — 435
neotomodon cliftoni _—____ 97 umbonatus was en ence n nese ne nn enn 436
neotomodon goldmani _______ 97 Welutin is jee 436
neotomcdon neotomodon _.. 97 yumae ___--_--_---_-___-_---_--_-------- 436
Nemadactylus macropterus ________- 395 osculans, Austrosquilla ____________ 243
neoafricanus, Parafelicola ________ 109 Or nnn nnn nena nnn nnn 587
Neogrubea's ese a 353 conchaphila ros 587
seriolellae _....-_.__-_-___-._ 353 s iridescens | wane nn nanan nanan nnn =n an 587
Neogrubeinae ss 352 ostreicola, Anachis ____-_________________ 147
Neonine] li sae elena eee eae 0 SOLS 31, 32
NGOSCOR A in nn eS De: | a 206 bakkamoena --_-__--------------- 32
Neotoma mexicana tenuicauda __ 99 elegans batanensis —________- 32
neotomodon, Nelsonia _____......___- 97 elegans botelensis 32
Nephthy ate naiemeit cc. err ea 739 elegans calayensis ______------- 32
ADETLAT Sen ee 739 elegans cuyensis 32
CLASS At ee ee 739 elegans elegans —_____.___.--.. 32
Sphacrophoral eens 739 elegans mantananensis ________ 32
PiXIGTA Gulp nad MEO TD 739 elegans romblonis _-_-.-------- 32
neuvoiedii, Bothrops —_.____________ 321 elegans sibutuensis _________. 32
Nezumia hildebrandi __ 440, 539 elegans steerei —__.....-_._-. 32
niei, Cachonina 92 leucospilus —_____----_---_---- 32
nigrescens, Porites == 738 longicornis -__------_-_--_--------- 32
nigricaudata, Sardinella 217 mindorensis —_____--_____--__- 32
nigrocauda, Ancherythroculter _____ 343 scops albiventris 32
rtetorA, INiatotk oo 144 scops)distans) 31
Nitidellaynifiday= 2&2. neaNoennONn Ne 144 scops kalidupae -—---_- 32
Nitocrella divaricata ___---._-..--_-._____. 582 sScops malayanus nena nn nnn nanan ane 31
nodosasildinckiae =... senna 42 scops manadensis ________- 31, 32
normalis, Bassozetus ___-__--_-_--___-_-____- 539 SCOpS mendeni wononnnnnnn nnn =-- === — 32
Nothoproctay 2 ae 485 ScOpS MIFUS --------------_-----__--_- 31
pentlandii doeringi _ _ 485 scops morotensis -_______________ 32
pentlandii mendozae 486 SCOPS S1aOeMSIS ___------------------- 32
pentlandii oustaleti 488 scops sulaensis —----__-_- 32
pentlandii pentlandii _. 485 SCODSBSUD1a p= 31
INOtothiers 1a Winans aa esti 20 MERSIN 409 scops tempestatis —____--_-- 32
Colbcclauaanns a409 whiteheadi 32
macrocephalay === = aaa 409 Oxobolus anna nnn nn nnn nnn nn nn nnn nnn 508
nototheniae, Pseudobenedenia _ 406, 409 >> 701 EEE 207
nova-britannea, Cormufer ________. 84 Oxyrhombus -----.-------------------------—-- 46
nuttallii, Dendrocopos __________ 1 P
O pachyclados, Cladiella 657, 741
oaxacae, Turdus assimilis ______________- 38 pacifica, Aricidea —_______________. 325
obatogamensis, Atheyellay === 574 E ys 336
obesawAnachis 2 es ee 144 Paimokia ——________ 521
obesula, Didelphis —______________ 251 LPalaechthon —__ = 629
obesum;, Buccinum: = 145 -,___alticuspis ——-_--___-____-__- 632
obeyensis, Cambarus ---------------- 12 + palifera, Acropora —__________ 738
obtusirostris, Microphysogobio __-_- 340 Palmeus, Phyllodactylus __ 419
Pseudogobio __..-._-_ 340 palmosa, Coelogorgia —-----.--------
octoptera, Seriatopora -__---.-------_-- 739 Palythoa ——-
oculatalyASell 1s) ee 759 Sc
Oedipodrilus «294 fubercnloca
ogygius, Cronodrilus ___ __ 308, 310 Pampus argenteus __-
Omomys A es eho! = dee oe Ae 630 Pantolambda ee
QOnchidellayeio2 Gee ees 588 pappenheimi, Gobiobotia —__-----.-.-. _ 836
Opaca,, Wlexy = se eae a 199 papuensis, Platymantis ______. 69, 70, 79
Ophidiaster ornithopus __. bt Al papyracea, Stereonephthya
Ophryodendron _________--_-____-____ 689 Paracambarus ____..._.__-
Opisthonemay = 217 Paradoxus) ee
Index
paracincta, Chone __________-___----__..-- 229
Paracycloisi ces 2s e ines A 610
Paradoneis yg eG 323
armatay = eee 324
ya ete eee sd en ee ee 324
paradoxus, Paracambarus __.____.______- 299
Paratelicolale oe Mee Nee ae 109
acuticeps |) 220s eee ean 109
ALTE CATLUS ee 109
neoafricanus __-__ 109
Paralemnalia thyrsoides ____--__. 740
Paramekodon _________--_-___------___. 440, 460
inflatus —___.________ 441, 458, 461
poulseni ____________ 439, 440, 464
iParaonides) jo. eee a Ba 324
(Paraonides), Paraonis ___________________ 323
Para Gnis pers ok Be ee Sine ae 323
harpagonea ___..._--_-______-.- 324
neapolitana _______------_--__---__- 324
(Baraonides)) 2 323
platybranchia ______--_-------_----- 324
Ppygoenigmatica _--__--- 327
RAT AV ATE Ul aye et ame meade ne 442,
Parerythropodium rubiginosum 685, 741
parkeri, Platymantis ____________--____- 77
Paromomys ______ _ 629
maturus _ 632
(Parvanachis), Anachis _______-___.. 145
parvispinosa, Bathyvargula ________ 448
parvoculus, Cambarus ____..__--___- 268
patulus, Lichomolgus __________ 741, 746
paulula, Bathyconchoecia ___ 541, 547
TERETE tA RE Ot NE Ne NY 737
fanculata: 2 ee 737
CACTUS je eee er ue era: 737
Gana see ae ae 737
VETUUS ta eee are AE aw 737
pecki, Procambarus _____--_.__--_- 13
pectorosa, Anyphaena ___..__.____ 207
pedunculata, Sinularia ____. 650, 742
pelewensis, Platymantis ______.______- 77
Rellenesie see ee Nee 208
pendula, Siphonogorgia ________ 713, 740
peninsularis, Mannobolus _____ 505, 507
pentlandii, Nothoprocta __....-_.--. 485
Perameles nasuta -__-_---------------------- 251
perforata, Dynamenella ______________. 589
Peridinium cinctum _______-_----------------- 93
SPOT OMS CUS) eee ae een 99
boylii levipes _________-__--_-_-___ 99
hylocetes hylocetes ___________ 99
maniculatus labecula __-____ 99
Petenobolus 508
antiquorum —__.-.-__ 508
mayanus ____..--- 509
phaleratus, Pseudoplisus __...--_.-_- 434
Phenacolemur _______-_-__---- _. 630
Philodromus rufus ______-------_----------- 208
Philomedes ______-_--- 440, 449
IPhOniSCUS re ee ee 4
phrixina, Ceochasma _____._____-__- 46
Phrurotimpus alarius _____________-_-__-_- 207
phrygianus, Pityohyphantes __________. 206
Phyllidia trilineata _______________ _._ 741
Phyllodactylus)” 22 419
TFISULATIS Up eee eee ES 419
Pal meus so eee ae eee 419
tuberculosus -___--------------------- 422,
pifanoi, Bothrops ------.
pileatus, Hylobates -__
pilosa, Atheyella
SADT US (pees IS 0 a a eS
jeffreyi _____-_-----------
ponderosa
sabiniana
pior, Motyxia —_---_.-__-..
Pirata montanus
pistillata, Stylophora __----.--..---------_-- 739
769
Pityohyphantes phrygianus ____________ 206
placera, Wamokia ______ “B11, 522, 530
placida, Mangora __________________________ 207
Placunella, ee Ee ae 371
platybranchia, Paraonis _______.__. 324
Platy.clacty Lusi meen cs aT 123
americana. Seer eet 123
Mil bertiiy yess sae ee 123
Platydoris scaber __...--..----_.__- 743
IB Vai yam 2 Gi see 69
acrochordus —___----__--__-_------- 73
aculeodactylus _________________
akarithymus _______- 70, 74
boulengeri _._ 69, 70, 71, 79
COTRU GA TUS he eee Ene
dorsalise 22 eee 77, 80
gilliardi (2 ils CA
IMAClOpS |e 79
meyersi ______ RENN ADE RI eee
mimicus ____-__________ 71, 73, 74
papuensis ______________ 69, 70, 79
papuensis papuensis ___________ 84
papuensis schmidti 75, 79, 84, 85
papuensis weberi _______-___- 81 , 84
parkeri _____ g evr e
pelewensis ___- us Sate 77
rhipiphaleus _________ 69, 70, 77
SOlOmOniS\ye eee 80
Weberti ace eis A LN 81
Plesiolestes)) 22.20 yk ee 630
problematicus ___-----_-_-__-__ 632
pleurotaenia, Gunnellichthys __. 60, 66
Poeciliopsis infans —_________---_-_____- 106
politus, Lichomolgus __._________- 739, 744
pollaesGaleus ses ee Ss Se 622
Roly copsis eset a eee. 440
polydactyla, Sinularia __________ 635, 742
ponderosa, Pinus 5
TROT ESS Ae eats EEE cee e ette
andrewsi
nigrescens
porosa, Seriolella 352
poulseni, Paramekodon _ 439, 440, 464
Preherrmannella _______..-----_--_-----___- 189
pretiosus, Dipsacaster _________-------------
pretmis Anachis
Columbella __-_--_--_--
pricei, Ictalurus -_____-_-__----------------
prionoti, Hatschekia _________-------------
pristinus, Cambarus -___-_------------------
problematicus, Plesiolestes
Procambarus) 22 ae
acutissimus —2_____-:—---
A CUICUS He ea Re Le ee
contrerasi
prolixipes, Lichomolgus
prominens, Cambarus
Hobbseus
Pronothodectes simpsoni -_----------------
Proscyllium habereri ------------------------
protentus, Lichomolgus
635, 679, 707, 742, uae
Protodorvilleay EEE 225
ibiarticulatay eee 226
gracilis
kefersteini) 232 = ees 226
minuta
recuperata
Psammocora contigua -_-------------------- 738
Pseudobenedenia _______---_---------------- 405
nototheniae
shorti
Pseudogobio
770 Proceedings of the Biological Society of Washington
brevirostris
fukiensis _____________
obtusirostris
Pseudomegalocotyla
Pseudophilomedes __________.
ata), obsess
Pseudoplisus phaleratus
Pseudosphaeroma
FES OL us pee ee ee
macrolepis
megaloplax __.___________________
Pterodrilus)- 32 ae
pteroidis, Lichomolgus
pterophilus, Lichomolgus ___________- 735
puebla, Asellus __.._.._..._-- 757
pouesent enioye, WWI 287
puSMaxy Wica pani we ee eee 275
pulex, (Habrocestum)) ae 208
punctulatus, Anahita 207
Pygoenigmatica, Paraonis ___________ 327
Q
@uercus eke ee Se ee 512
kelloggii ie eee 512
Wis liz erie) eee eee 512
R
radcliffei, Triakis (Eridacnis) ___ 614
Radianthus xitteri) eee 740
TajinE pelras evs Oe Ae ee ee 207
Ranavinucatayse ess tee tu Ce 79
TDP) AX ANU) Cea yee ae rae eae a ee 275
raviceps, Leiocephalus 23
recuperata, Protodorvillea __________ 225
reducta, Streptosyllis ____.____________. 153
reduncus, Cambarus ____________ 12, 273
regia, Dendronephthya ____________ 740
Reithrodontomys sumichrasti ~ ner-
LOTUIS nals eae ae ereree 99
remota, Wamokia _________ 514, 522, 535
repens, Mitchella —________.__________ 199
reticulata, Daector _____.____________
IRbabd omnis se eke a ces
The(Onae AO)
inornatus alaris _________________
inornatus zamboanga __.._
rhabdotus, Tomicodon ___..._._
rhadinus, Lichomolgus _______
rhamphodes, Tetragonodon ________
Ei eee sDreataLn 439, 440, 449, 456
Rhigophila dearborni 406
Rhincod on} 2220 1) seas as See 613
rhipiphaleus, Platymantis __ 69, 70, 77
rhoadesi, Orconectes ae ROCA
Rhodactis rhodostoma _____.
rhodae, Anachis ___---_-_______
Rhododendron maximum
thodostoma, Rhodactis _.____________-
Rhombinella) 2 ee
ME va sata ies seein Ea
laevigata hendersonii ________ 145
ritteri,) Radianthus) 2) eee 740
robustus, Cambarus _____..___ Ls, BAF(Bd
Lichomolgus _________ a GS
rosea, Chone 2) 0a oe 229
rowleyi, Ergaticus ruber __ a BY)
rozei, Bothrops lansbergii _- _.. 320
ruber, Doridopsis _________-- AN:
Tabgep ovens i ae ae 39
rubiginosum, Parerythropodium 685, 741
rubromaculata, Trevelyana _____- TAL
ineloyabhaay, Jofeye
ruficornis, Gorytes __
rufulicornis, Gorytes _
rufus, Philodromus __
Lugataw Rana seas
rupicola, Theridion __-
rusticus, Orconectes _____.______-
rutilum, Harpactostigma ________ 434
sabiniana, Pinus ______________-.._-__ 530
sagittarius, Bathyconchoecia _ 542, 561
salmoides, Micropterus 106
Sarcophyton __.-
ehrenbergi ________________
glaucum) =e
globosum __
Sardinellay =a
( Ambly gaster )
(Clupeonia )
jussieu _________
marquesensis
melanura _______
nigricaudata ___
sundaicay es
Sathodrilus _
carolinensis __
Megadenus ____
veracruzicus ___
villalobosi _____
scaber, Platydoris _______
scalaris, Dendrocopos ____.
Scambicornus _______________
campanulipes _____
Schazicheila Ss Bae
scherzeriana, Acropora —_________ 738
schmidti, Platymantis papuensis __
ERC 75, 79, 84, 85
schmitti, Daector __......-- 155
scops, Otus ____---_ _. 31, 32
Scrippsiellay) =e eee 95
scyllia, Triakis __ _.. 614
Scytaster(stellay Al
securiger, Lichomolgus _ 702, 741, 746
Sediumm(tis 2 Sees Ae eee eee 199
semifasciata, Triakis 614
sensilis, Lichomolgus __________- 741, 746
SeriatoporaY) 2 Se eee ae 739
octoptera __. 39)
subseriata (U2 a ee 739
Seriolella 352
brama ___ __. 352
OLOSa ee _.. 352
seriolellae, Neogrubea _____ —— 5p
serratipes, Lichomolgus _ 737
setosa, Dynamenella _._- 594
seychellensis, Stanulus __________ 111, 113
shorti, Pseudobenedenia ________________ 406
sierrae, Wamokia _______ 513, 522, 537
Sive -fusigeray) sana a ee 225
Sigmocheini) =e ae — 513
simpsoni, Pronothodectes _______ 632
simulans, Lichomolgus _______- 739, 744
(Sincola), Strombina —.-..__-_-__-___ 148
singularipes, Lichomolgus _________
GOS ITT e US 636, 685, 736, 741
Singlitiqneg | Lo _ 479
sinuans, Triakis (Neotriakis) _____-
Sinularia
humesi __
leptoclados
polydactyla
whiteleggei
650, 689, 742
Siphonariay gig asi see eee
Siphonog orga ee
pendul ay 2-2 oe eae
sladeni, Dipsacaster _______-_-_---____
solanderi, Standella ______-_-__-__--
soleatus, Eurelus __-_..--____-______
solomonis, Anamygdon
Platyam antic eeee
sordidus, Micromacronus leytensis 33
sparsay Anachis ys. ee 148
Index tal
spathophorus, Tichomoleus pies Ue tek
2) 3 Le eee a 674, 739, 744
speciosa Wea, 22 Ue 287
sphaerophora, Nephthya —__--------_- 739
sphenoides, Cambarus
spicatus, Cambarus ________________--______
spiculifer, Procambarus
spinicarpa, Uca ______--______-
spinipes, Lichomolgus eS
spinosa, Lysiosquilla ____._______________ 24
spinulifer, Lichomolgus —____- 740, 746
Spirobolus 22) 26 2 eRe esi 507
Sprostonia -_----.-_-----___
longiphallus
squatinae ___..--_
squali, Erpocotyle
Squalonchocotyle
Squalonchocotyle —______---__.--
acanthi ____ 363
antarctica _____ 359
callorhynchi 360
SCAM a oe De ee 363
SUT ea as bgp ea ea ee ae ed 363
acanthias _- _-. 363
Lebrun ee eee 363
squamiger, Lichomolgus ______-_--___-
me eee 635, 689, 707, 742, 747
squamosa, Tetraclita ____________.____.. 588
squamula, Entobdella __ _.. 391
Squatinag ee _._ 382
longiphallus _ _.. 382
Sua tin a ee RU el 382
squatinae, Sprostonia ____ 371, 381
Stantella solanderi ___.___._---------_-_- 739
Stanulusy ae een 111, 112
seychellensis ________--___-_- 111,113
talboti _..-
Stauronereis __-_---
japonica ___
stella, Scytaster ___
Stellicola
Stephanodrilus __.
Stereonephthya ___
acaulis __.__..
papyracea
stictigaster, Leiocephalus _- Me
stocki, Dendronephthya _______ el
Stoichactus giganteum __. aunt
stokesii, Chiton __---_-__------ as
Streptosyllis ______ eee
arenae ___ uses
bidentata ______- sete
cryptopalpa __- ee
latipalpa _______ jes
reducta __ wee
varians __ ny
websteri _________--_- ae)
striata, Archiconchoecia _- we
@honey ieee Sloe Sa
striatus, Cambarus ________-_- 12, 262,
SS tree gg ES
balani __-----_-_____ deity
Stromateoides argenteus _ aot
Strombina (Sincola) ---_. se
Stylophora ____-_-----_---_- as
mordax __. zine!
pistillata ____ £29
subulatus, Myotis _____________________
Mespertilo) 2223 eee
subseriata, Seriatopora —_.____________
sumichrasti, Reithrodontomys ____-
sundaica, Harengula -__----___-___
suspecta, Chone _-_-___-____-___
(Suturoglypta), Anachis __
Synapta maculata ____._-__-___
Sylar ec eee Lee lee AEN esac ei
T
tabogensis, Microgobius ________________- 192
talboti, Stanulus
Tarentola
americana
americana americana _
americana warreni _____ 134
cubanay (see Aenea 123
tauvina, Epinephelus __________________ 377
tauvinae, Ailosprostonia _-.____ 383
Mecticepsy tee ess eee ea 598
PR CLEStO Sees ER ie aa ee INE 693
arborea ee 725, 741
telestophilus, Lichomolgus 725, 741, 746
tenebrosus, Cambarus ___ 268, 314, 581
tepidariorum, Achaearanea ______ 206
teressiChone ye ae ATS 229
Tetraclita squamosa panamensis —_ 588
Metragnathaw ae 207
elongata 2 Sans eS 207
VeETSICOlON je Ee 207
Tetragonodon _______--____________ 440, 449
ctenorhynchus _________ 441, 449
ETINACCUS ee ek a 450
rhamphodes _ 439, 440, 449, 456
Tettodrilus === 294
friauhige Nw a
texana, Clubionoides _...-_--.-_
thalassinus, Entomacrodus __
Thalassophryne maculosa ______.
Wh eridion oN ee ly
albid umn 2) aa es oe
UPICOLa yy ee eros
Mheuidigsoms gemmosum
hod inaye asker Ow ee Seabee oe
Thylacis LES I oe ERNE Oe cae sea
thyrsoides, Paralemnalia —___________.
tigrina, Genetta ______________--_-___.
tixierae, Nephthya ___. me
tomalensis, Asellus ___-_._.__-_-_____
AROmMICOd Oxi ee
LAS CLA CS) ese aes
humeralis) ee
rhabdotus __.---------
Torrejoniay | 200s) elie eas a
WALSOMD Se ey
Arar Che ais ee 2 ea Bee 8 207
(Erematomus; (500s oe ee
bermacchiie sea Se
borchgrevinki _____.
centronotus -___----
has oni et es
Trevelyana rubromaculata __
oLalcis yea Ss UN MEN ae
acutipinna ________-________
attenuata -__-.
barbourls 2s eee
(Eridacnis) radcliffei
fehlmanni _—
henlei ________-
maculata essere
(Neotriakis ) sinuans _
Scylhia nessa
semifasciata
venusta —____-
trilineata, Phyllidia ____.
triquetrus, Lichomolgus
trispinosus, Lichomolgus
Tristramia ___-
Trochopella __.
‘txrochopus) 22 eee
hoboy == sea ee
tubiporus ___.
Troglocambarus _._
tropica, Dynamenella __
Tsuga canadensis _____.
tuberculosa, Palythoa __________.
tuberculosus, Phyllodactylus
772 Proceedings of the Biological Society of Washington
Tubipora musica
tubiporus, Trochopus
tulipifera,
urdus. =e
assimilis
lygrus
oaxacae
renominatus
assimilis
assimilis
assimilis
1 DIC rs ge en a OO ko
burgersi ___________
longisignalis __.
pugilator
pugnax ___
rapax eee
speciosa
spinicarpa
WITGNS (02 2S oe
umbellata, Xenia
umbonatus, Hopliseides od pele 436
Oly thus ee eee 436
unicinctus, Hamactostiena hse Ne 436
[iogorytes!| 2222 ee 436
Uvalidat 220. na eal ell 499
intersecta 22.22 sa ee 500
V
vagans, Lichomolgus
valleculus, Cambarus
Hobbseus _______-
valviviae, Conchoecia
varia, Microneta _______-_--.
varians, Streptosyllis 53
varirostratus, Lichomolgus 719, 740, 745
velutinus, Harpactostigma _ 431
foplisus oo es Sa eee 436
Oryttus 2 2 eee. eet 436
venezuelae, Bothrops ___-__-__---.__- 319
venezuelensis, Bothrops —__-_-----..-._- 319
venusta, Lanassa —___-____-_-_______-__.__- 226
Leucauge _.. Pe 20%
Ravona yyw aes 737
Bra tel a tfc a ee 614
venustum, Calliscyllium —_--__.___ 614
venustus, Lichomolgus __________. 741, 745
veracruzicus, Sathodrilus _________ 305
vercoi, Austrosquilla ___.._________. 243
verseveldti, Lichomolgus _ 694, 741, 746
versicolor, Ergaticus ______...______.--__--_-
Tetragnatha
Vespertilos 2223 ee 258
eibii _____.. ae _... 259
subulatus ___ _-.. 258
Wianay 22 os ae ee 45
Viguiera deltoidea —____...-_-_-___- 437
villalobosi, Sathodrilus ___________ 299
Wwirens, Uca. 2-2) beset _._ 277, 281
virginianus, Colinus _______..._-_____- 37
virginiatus, Drassyllus ________________- 207
viridescens, Gunnellichthys 59, 61
vittata, Harengula -__--.._.--..... 09.
Herklotsichthys _
Maeviai. <a eee
W
Waimokiaw= ==. ae
walfordi, Bathyvargula _.
Wamokia
hoffmani
placera -...--.---.--.-
remota _____- 5
sierrae
wangi, Ancherythroculter ______________
Erythroculter —______--_--_--___-__-
warreni, Tarentola americana _____-
weberi, Platymantis
websteri, Streptosyllis ___.
whiteheadi, Otus
whiteleggei, Sinularia _._. 650, 689, 742
wiedi, Lonchoglossa 429
wilsoni, Torrejonia ___
wislizenii, Quercus _____----_--------------_-
x
Kemi ay ites eae ee 693
umbellata ___ 707, 742
MGronodrilus; eee 2
Xironogiton —__. 292, 309
Mey sticus 2 dee 207
elegans) 20 aa 207
ferox® i222 .. 207
Xystocheir acuta —______.-____---___-__-__ 521
Y
yumae, Oryttus ______---_------------ 436
yumanensis, Myotis ___.-..-------------.- 258
Z
Zelotes duplex _____.____--.-.-----_-------_- 207
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3.9088 01205 1991
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